CSIRO PUBLISHING

www.publish.csiro.au/journals/emu Emu, 2006, 106, 141146

Sex determination and sexual size-dimorphism in Southern

Giant-Petrels (Macronectes giganteus) from Patagonia, Argentina

S. CopelloA,D, F. QuintanaA,B and G. SomozaC

A
Centro Nacional Patagnico (CONICET), (U9120ACF) Puerto Madryn, Argentina.
B
Wildlife Conservation Society, New York, NY 10460, USA.
C
Instituto de Investigaciones Biotecnolgicas-Instituto Tecnolgico de Chascoms (IIB-INTECH),
CONICET-Universidad Nacional de General San Martn, (B7130IWA) Chascoms, Argentina.
D
Corresponding author. Email: scopello@cenpat.edu.ar

Abstract. We used discriminant analysis of a series of external morphometric measurements, together with
confirmation by genetic methods, to develop a simple method to determine the sex of fledgling and adult Southern
Giant-Petrels (Macronectes giganteus) in colonies in northern Patagonia. We also described morphometric
parameters (bill-length, bill-depth, tarsal length, wing-length, and weight) for different classes of age and sex and
analysed differences in size between male and female chicks over the growth period. Southern Giant-Petrels showed
conspicuous sexual size-dimorphism, with males being significantly larger than females, this difference being
apparent within 8 weeks of hatching. In adults, males were 40% heavier than females and consistently larger in all
measurements (% sexual size-dimorphism 515%). For fledglings, bill-length was the most dimorphic variable
(15.2%). The discriminant analysis of four external measurements correctly identified the sex of more than 90% of
adults and fledglings. Bill-length alone was sufficient to predict sex of adults and fledglings and can thus serve as
a simple and effective tool for sexing Southern Giant-Petrels.

Introduction biology techniques constituting a reliable tool to assign sex

The determination of sex is important for ecological studies
where gender-specific variation is expected in the phenom-
ena studied. Given the usual limited opportunity to sex live
birds using morphological or behavioural methods in the
field, determination of sex by genetics or discriminant func-
tion can be particularly valuable. Body measurements have
been widely used to discriminate gender using predictive
functions (Coulson et al. 1983; Evans et al. 1993; Evans and
Cavanagh 1995; Phillips and Furness 1997; Renner et al.
1998; Lo Valvo 2001) and, more recently, DNA-based tech-
niques have proved successful in determining sex in a variety
of seabird species, some studies combining DNA-based
work with behavioural or morphometric measurements (Van
Franeker and Ter Braak 1993; Torlaschi et al. 2000; Velando
et al. 2000; Lo Valvo 2001; Bertellotti et al. 2002; Quintana
et al. 2003).
Most seabirds are sexually monomorphic although some
procellariiforms, notably the albatrosses (Diomedeidae), ful-
marine petrels (Fulmarus spp., Thalassoica antarctica,
Daption capense and Pagodroma nivea) and giant-petrels
(Macronectes spp.) are markedly dimorphic in size (Van
Franeker and Ter Braak 1993; Gonzlez-Sols et al. 2000a;
Shaffer et al. 2001). However, even in such cases, gender
determination by discriminant function analysis could prove
useful in sexing borderline individuals, with molecular
for discriminant function development. Few of the existing
studies on albatrosses and large petrels, however, have
employed either or both techniques to determine the sex of
chicks, fledglings or adults (Hedd et al. 1998; Berrow et al.
1999; Shaffer et al. 2001; Double et al. 2003).
The Southern Giant-Petrel (Macronectes giganteus) is
widely distributed throughout the southern oceans, breed-
ing on several subantarctic islands, the Antarctic Peninsula,
Malvinas (Falkland) Islands, and southern Chile and
Argentina (Harrison 1983). Although there is information
on morphometric parameters and sexual dimorphism of
breeding adult Southern Giant-Petrels in some colonies
(Voisin and Bester 1981; Hunter 1984; Gonzlez-Sols et al.
2000a), there is no published information for the colonies
of Patagonia, and no discriminant functions have been
applied to determine the sex for other age-classes. In this
study, we used genetic sexing techniques (assumed correct)
in the development of predictive morphometric discrimi-
nant function models for fledgling and adult Southern
Giant-Petrels from two neighbouring colonies (47 km apart)
located at the northern extreme of this species distribution.
In this work we also describe morphometric parameters for
adults and fledglings of each sex and we analyse differences
in size between male and female chicks over the growth
period.

Royal Australasian Ornithologists Union 2006 10.1071/MU05033 0158-4197/06/020141

142 Emu S. Copello et al.

Materials and methods
As part of a wider study on the breeding biology and foraging ecology
of the Southern Giant-Petrel, adult birds and fledglings (birds older than
90 days) were captured during four breeding seasons (from October to
April, 19992002) at two colonies in coastal Patagonia, Argentina (Isla
Arce, 4500S, 6529W, and Isla Gran Robredo, 4508S, 6603W).
A total of 104 birds (40 adults and 64 fledglings) was used to take exter-
nal measurements and collect blood samples for sex determination. For
a detailed description of blood extraction, sample storage and DNA
analysis see Quintana et al. (2003). Fledglings were sampled every
season between the first and the second week of April to guarantee they
were >90 days old according to the hatching date reported by Quintana
et al. (2005) for the studied colonies. During the 2002 breeding season,
a sample of 21 chicks of known age was measured after the first 10 days
of life up to the sixteenth week of age, with measurements being taken
every 10 days, to analyse differences in size between males and females
during the growth period.
We took five body measurements: wing-length (WL), measured
from the carpal joint to the tip of the longest primary; bill-length (BL),
measuring exposed culmen; bill-depth (BD), measured at the nostril;
tarsal length (TL), measured from the middle of the mid-tarsal joint to
the distal end of the tarsometatarsus; and body-weight (BW).
Measurements of both wing and tarsus were taken on the right side of
the body. Wing-length was measured using a stopped ruler ( 1 mm),
while bill-depth, and length of bill and tarsus were measured using
digital Vernier calipers ( 0.01 mm). All birds were weighed to the
nearest 25100 g using Pesola spring balances.
One-way ANOVAs or MannWhitney U-tests were used to deter-
mine whether the overall external morphology varied with sex, and
between adults and fledglings of the same sex (Sokal and Rohlf 1995).
The percentage of dimorphism between sexes in each measure was cal-
culated as: [(xm x f)/xf] 100, where x m and x f are mean values for
males and females respectively. We used the statistical package
STATISTICA 6.0 (StatSoft. Inc. 2001) to apply the discriminant analy-
sis to the biometric data of birds of known sex. Body-weight was not
included in the discriminant analysis because it may vary throughout
the year and during the breeding season (Croxall 1995). The perfor-
mance of each variable was evaluated using Wilks Lambda, which
decreases as discriminatory power increases and with the allocation rate
of the model (the percentage of individuals correctly identified for each
sex). The measurements that best discriminated between sexes were
selected to derive a discriminant function model. For each discriminant
function the associated cutting score (cut-off value) was calculated (see
Phillips and Furness 1997). Birds with a score higher than the cut-off
value were classified as males, and those with a lower score classified
as females. Only the classification of new cases allows assessment of
the predictive value of the classification functions, so we used a cross-
validation method (jackknife) leaving out one case, and then classified
that case using the function derived from the remaining birds
(Tabachnick and Fidell 1996).
Results
Sexual dimorphism
In fledglings, males were significantly larger than females in
all body measurements (P < 0.0001) except for WL and BW
(P > 0.05) (Table 1). Of the five body measurements, only
BL did not overlap between sexes. The degree of sexual
dimorphism for fledglings ranged from 3% to 15%, with BL
(15.2%) being the most dimorphic variable (Table 1).
Adult males were significantly larger and heavier than
females for all body measurements (P < 0.01) (Table 1). The
range of bill-dimensions (length and depth) did not overlap
between sexes. The degree of sexual dimorphism for adults
ranged from 5% to 40%, depending on the body measure
(Table 1). BW showed the greatest sexual size-dimorphism
(40%), followed by BL (15.2%) and BD (14.1%). TL and
WL were the least dimorphic variables (8.7% and 5.1%
respectively) (Table 1).
The sexual dimorphism of chicks (males larger than
females) was apparent from the eighth week of life for all
body measures (P < 0.01), except for WL and BW, which
were similar between sexes in all age-classes (P > 0.05). The
percentage of dimorphism in bill-size ranged from 2% to
13% and from 6% to 14% for BL and BD respectively. These
two measurements were clearly the most dimorphic variables
from the eighth week after hatching. Sexual dimorphism in
bill-size was already apparent 4 weeks after hatching and
showed a marked increase with time (Fig. 1).
Discriminant analysis
All measured characteristics taken alone significantly pre-
dicted the sex of fledglings (F > 134.1, P < 0.0001), except
for WL (F = 3.2, P > 0.05). Bill-length was the most accurate

Table 1. Body measurements (mean s.d., and n) and percentage dimorphism of adults and fledglings of the Southern Giant-Petrel
from Northern Patagonia, Argentina
n.s., not significant (one-way ANOVA or MannWhitney U test)

Fledglings Adults
Variable Males Females Sex comparison % Males Females Sex comparison %
Test P dimorphism Test P dimorphism
Wing-length 46.3 2.5 44.8 3.2 175 n.s. 3.4 51.7 1.2 49.1 0.9 10.2 < 0.01 5.1
(cm) (n = 26) (n = 19) (n = 3) (n = 8)
Bill-length 93.2 2.4 80.9 2.6 381.3 <0.0001 15.2 95.5 2.6 82.9 2.1 305.1 < 0.001 15.2
(mm) (n = 37) (n = 27) (n = 18) (n = 22)
Bill-depth 26.8 1.1 23.8 0.9 133.9 <0.0001 12.6 29.2 1.0 25.6 0.8 173.3 < 0.0001 14.1
(mm) (n = 37) (n = 27) (n = 18) (n = 22)
Tarsal length 91.7 2.6 84.8 1.8 136.4 <0.0001 8.1 92.2 2.1 84.8 3.6 16.0 < 0.0001 8.7
(mm) (n = 36) (n = 27) (n = 15) (n = 22)
Weight (kg) 4.2 0.9 3.7 0.6 261 n.s. 13.5 3.5 0.3 2.5 0.2 113.3 < 0.0001 40.0
(n = 26) (n = 27) (n = 15) (n = 21)
Sex determination and dimorphism in Southern Giant-Petrels Emu 143

indicator of sex, correctly classifying 100% of cases (Wilks
Lambda = 0.14). Bill-depth and TL had less discriminatory
power, correctly classifying 95% and 96% of fledglings
respectively (for both cases Wilks Lambda = 0.31).
Discriminant analysis provided three highly significant func-
tions including only a single variable (BL, BD or TL). The
resulting functions (with associated cut-off values) were:
D1 = 35.395 + BL 0.402 (0.150)
D2 = 25.083 + BD 0.981 (0.425)
D3 = 47.755 + TL 3.089 (0.027)
The jackknife validation process provided the same
classifications as those produced by discriminant analyses.
Given that the ranges of BL showed no overlap between
sexes, adults are easily sexed by simple measurement of the
bill. At the studied colonies, adult birds with bills longer than
92 mm can be considered as males. However, we performed
a discriminant analysis as a complementary method of
assigning sex. All characteristics measured alone signifi-
100
(a)
90
80
Bill length (mm)
cantly predicted the sex of Southern Giant-Petrels (F > 10.2,
P < 0.0001). BL and BD were the most accurate indicators of
sex, correctly classifying 100% of adult birds (Wilks
Lambda = 0.11 and 0.18 respectively). TL and WL had less
discriminatory power, correctly classifying 92% and 94% of
adults respectively (Wilks Lambda = 0.13 and 0.47 respec-
tively). The resulting functions (with associated cut-off
values) including only one single measure were:
D1 = 39.050 + BL 0.441 (0.44)
D2 = 31.507 + BD 1.156 (0.78)
D3 = 25.530 + TL 0.325 (0.87)
The jackknife validation process provided the same clas-
sifications as those produced by discriminant analyses.
Discussion
Sexual dimorphism
Our results show a conspicuous sexual size-dimorphism in
Southern Giant-Petrels, which is apparent even during the
first weeks of life, and was evident for three (BL, BD and
TL) of the five body measures considered in this study. For
adults, males were 40% heavier than females and consis-
tently larger than females in all other measurements taken
(515%) (Table 1). With the Northern Giant-Petrel
(Macronectes halli), the Southern Giant-Petrel shows one of

the largest sexual size-dimorphisms of seabirds, and consti-

70
tutes the second largest degree of dimorphism of its family
60 (Procellariidae) behind its sibling species (see Hunter 1984,
1987; Gonzlez-Sols et al. 2000a, 2000b; Gonzlez-Sols
50 2004). The degree of sexual dimorphism of adult Southern
Giant-Petrels found at the Patagonian colonies was similar to
40
that reported at other breeding locations (Table 2) (Voisin
30
1988; Gonzlez-Sols et al. 2000a).
2 4 6 8 10 12 14 16 Sexual dimorphism in birds may reduce intersexual food
30 competition through the evolution of different phenotypes
(b) (e.g. bill-shape or bill-size, or both, body-size) that facilitate
28
the separation of feeding niches between sexes (Selander
26 1972). In fact, intersexual differences in foraging and
Bill depth (mm)

feeding ecology have been observed in both species of giant-

24
petrel. Several studies have highlighted sex-related dif-
22 ferences in the diet of the two species, with males feeding
almost exclusively on penguins and seal carrion, and females
20
taking primarily fish and cephalopod prey (Hunter 1983;
18 Hunter and Brooke 1992; Forero et al. 2005). Both species
also show remarkable sexual segregation in foraging areas
16
during the breeding period, as shown by satellite telemetry
14 studies, with males foraging mainly along the coast and in
2 4 6 8 10 12 14 16
coastal waters and females foraging offshore (Gonzlez-
Age (weeks) Sols et al. 2000a, 2000b; Quintana and Dell' Arciprete 2002;
Fig. 1. Growth-curves for (a) bill-length and (b) bill-depth of
Patterson and Fraser 2003; Quintana et al. 2005).
Southern Giant-Petrels from northern Patagonia, Argentina.
, males; Of the linear measurements analysed in our study, bill-
, females. Values are means 95% confidence intervals. dimensions showed the greatest sexual dimorphism, with
 144
Emu

Table 2. Measurements (mean standard deviation, range and n) and percentage dimorphism of males and females of Southern Giant-Petrels from different localities

Locality Bill-length (mm) Bill-depth (mm) Tarsal length (mm) Wing-length (cm) Reference
Males Females % Males Females % Males Females % Males Females %

Gough Island 95.2 2.4, 83.3 2.9, 14.3 30.0 1.1, 27.3 0.7, 9.9 95.3 2.0, 88.3 3.1, 7.9 50.7, 48.4, 4.7 Voisin and Bester 1981
(43S 10W) 91.598.5 79.588 28.532 2728.5 9299) 8595 8.553.5 47.849.1
(n = 17) (n = 12) (n = 17) (n = 12) (n = 17) (n = 12) (n = 4) (n = 2)
Arce and Gran 95.5 2.6, 82.9 2.1, 15.2 29.2 1.0, 25.6 0.8, 14.1 92.2 2.1, 84.8 3.6, 8.7 51.7 1.2, 49.1 0.9, 5.1 This study
Robredo Islands 91.7101.2 80.1287.3 27.530.9 24.427.1 87.796.9 80.599.1 50.753.0 47.850.8
(45S 65W) (n = 18) (n = 22) (n = 18) (n = 22) (n = 15) (n = 22) (n = 3) (n = 8)
Iles Crozet 104.7 4.0, 89.1 3.3, 17.5 33.0 2.0, 28.6 1.0, 15.4 102.4 2.6, 95.1 6.8, 7.7 53.3 1.4, 49.5 2.4, 7.7 Voisin and Bester 1981
(46S 51W) 97111 8494 3036 27.530 97107 88111 49.255.5 46.052.2
(n = 16) (n = 8) (n = 12) (n = 7) (n = 14) (n = 8) (n = 15) (n = 7)
Islas Malvinas 97.9 3.2, 84.4 2.5, 16.1 30.2 1.6, 25.9 0.9, 16.6 93.0 1.5, 84.0 2.2, 10.7 51.1 1.0, 48.5 0.9, 5.4 Voisin 1982
(Falkland Islands) 94.5102 8089 28.532.5 24.527.5 91.595 79.586 50.052.5 47.050.0
(52S 59W) (n = 5) (n = 10) (n = 5) (n = 10) (n = 5) (n = 10) (n = 5) (n = 10)
South Georgia 102.3 2.3, 88.0 2.7, 16.2 100.1 2.3, 91.1 1.9, 9.9 53.9 0.8, 50.8 1.2, 6.1 Gonzlez-Sols 2000a
(54S 36W) 98.4108.6 84.494.9 95.4105.1 87.494.1 52.755.5 48.552.7
(n = 19) (n = 20) (n = 19) (n = 20) (n = 19) (n = 20)
Macquarie Island 101.9 2.1, 89.7 2.4, 13.6 102.7 1.7, 94.2 1.9, 9.0 55.2 1.1, 52.7 0.8, 4.7 Voisin and Bester 1981
(54S 158E) 95.8105 84.693.5 99.5105.3 90.198 53.457.7 51.354.0
(n = 20) (n = 21) (n = 20) (n = 21) (n = 20) (n = 21)
Signy Island 101.4 2.4, 87.4 3.0, 16.0 96.3 1.7, 87.1 5.4, 10.6 55.3 1.1, 51.9 1.5, 6.5 Conroy 1972
(60S 45W) 97.4108.2 8297 9499 8295 53.457.1 49.854.1
(n = 66) (n = 73) (n = 13) (n = 13) (n = 13) (n = 13)
Antarctica 98.5 2.0, 89.1 4.1, 10.5 34.3 3.3, 31.7 4.0, 8.2 94.7 5.1, 89.0 3.3, 6.4 51.0 2.5, 50.1 1.3, 1.8 Voisin and Bester 1981
Peninsula 96102 85.595 96102 26.535 85101 8493 46.555.2 47.751.5
(64S 61W) (n = 9) (n = 8) (n = 8) (n = 8) (n = 9) (n = 6) (n = 8) (n = 8)
Terre Adlie 98.2 4.4, 85.2, 15.3 30.1 1.8, 27.5, 9.4 96.0 3.3, 92.0, 4.3 53.4 1.5, 50.6, 5.5 Voisin and Bester 1981
(66S 140E) 92104 8389 2732 2728 9199 8994 51.754.9 9.351.4
(n = 6) (n = 3) (n = 5) (n = 3) (n = 6) (n = 3) (n = 5) (n = 3)
Frazier Island 98.9 2.3, 88.0 2.1, 12.4 99.3 2.3, 90.4 2.2, 9.8 54.2 0.8, 51.2 0.7, 5.9 Voisin and Bester 1981
(66S 110E) 96103.1 85.890.4 95.8102 87.893.8 53.055.0 50.051.8
(n = 7) (n = 5) (n = 7) (n = 5) (n = 7) (n = 5)
S. Copello et al.
Sex determination and dimorphism in Southern Giant-Petrels
wing-length being least conspicuously different for adults,
fledglings and chicks more than 8 weeks old. The same trend
was also reported for the two species of giant-petrels in other
locations, suggesting past directional selection pressures for
bill-dimensions and wing-length in males and females
respectively (see Gonzlez-Sols and Croxall 2005). Larger
bills may make males more efficient in competing for access
to carcasses, and larger wings in females (with respect to the
body-size) may make them more efficient at foraging longer
distances than males (see Gonzlez-Sols and Croxall 2005).
Discriminant analysis
The discriminant analysis we conducted provided high allo-
cation rates in adults and fledglings (> 90%). Similar allo-
cation rates have also been found for adults in other
procellariforms, such as albatrosses, shearwaters and fulma-
rine petrels (Van Franeker and Ter Braak 1993; Berrow et al.
1999; Genovart et al. 2003) and also in fledglings (Hedd
et al. 1998). For adults and fledglings, BL was sufficient to
predict the sex and can serve as a simple and effective tool
for quickly sexing Southern Giant-Petrels in the field. This
finding accords with previous studies on giant-petrels that
also show that bill-length is a key characteristic for differen-
tiating the sexes (Hunter 1984, 1987). The ability to sex
fledglings by simple measurement of the bill-length is also a
convenient and cheap way to monitor fledgling sex-ratio in
giant-petrels. Given that measures of two different parts of
the body (bill and tarsus) were appropriate to distinguish sex,
the functions presented here could prove extremely useful for
determination of gender in museum specimens, carcasses
and birds killed incidentally in fisheries, particularly in cases
where only few body parts are available or in a condition to
be measured.
The discrimination functions to sex Southern Giant-Petrel
adults and fledglings reported in this paper constitute an
easy, rapid, inexpensive and minimally stressful technique.
However, studies of several seabirds species have noted geo-
graphical variation in morphometrics (Bost and Jouventin
1990; Murie et al. 1991; Gandini et al. 1992; Kerry et al.
1992; Van Franeker and Ter Braak 1993; Waugh et al. 1999)
and suggest that sex determination by discrimination func-
tions must be interpreted carefully given that geographical
variation may invalidate the functions applicability. A
review of previously published morphometric data of the
Southern Giant-Petrel from ten locations (Table 2) indicates
that birds from the Patagonian colonies have smaller dimen-
sions than conspecifics at Antarctic and subantarctic
colonies although they are similar to those reported at
colonies located in the same latitudinal range, such as Gough
Island (4320S 954W) and Islas Malvinas (Falkland
Islands) (5200S 5930W) Islands. Although there are
morphological differences between locations, the relative
size-dimorphism between sexes is remarkably consistent
within each measurement. Furthermore, bill-dimensions (the
Emu 145
most accurate indicator of sex) were the most dimorphic
measurements in all locations (10.517.5% and 8.216.6%
for BL and BD respectively; Table 2). Thus, identification of
sex using bill-dimensions appears to be a generally applica-
ble technique for populations of Southern Giant-Petrel.
Acknowledgments
Research was part of wider research on the Southern Giant-
Petrel on the Patagonian coast of Argentina funded by the
Wildlife Conservation Society, Agencia Nacional de
Promocin Cientfica y Tecnolgica (ANPCyT), Consejo
Nacional de Investigaciones Cientficas y Tecnolgicas
(CONICET) and Ecocentro Puerto Madryn. We thank the
Direccin de Fauna y Flora Silvestre of Chubut Province for
issuing the work permits to capture and handle birds. We
would like to thank A. Sapoznikow, N. Suarz, M. Uhart,
A. Sabatini, J. Braun, J. E. Sala, G. Morales, F. Marquez,
A. Steinfurth, I. Zimmer and I. Escapa for their help in the
field and G.C. Lpez for her technical assistance in the lab-
oratory. Special thanks to J. Owen and R. Vera for logistical
help. T. Svagelj, P. Yorio and R. P. Wilson are also acknowl-
edged for their valuable comments on early versions of this
manuscript.
References
Berrow, S. D., Huin, N., Humpidge, R., Murray, A. W. A., and
Prince, P. A. (1999). Wing and primary growth of the wandering
albatross. Condor 101, 360368.
Bertellotti, M., Tella, J. L., Godoy, J. A., Blanco, G., Forero, M. G.,
Donazr, J. A., and Ceballos, O. (2002). Determining sex of magel-
lanic penguins using molecular procedures and discriminant func-
tions. Waterbirds 25, 479484.
Bost, C., and Jouventin, P. (1990). Evolutionary ecology of Gentoo
Penguins (Pygoscelis papua). In Penguin Biology. (Eds L. S. Davis
and J. T. Darby.) pp. 85109. (Academic Press: San Diego, CA.)
Conroy, J. W. H. (1972). Ecological aspects of the biology of the giant
petrel Macronectes giganteus (Gmelin) in the maritime Antarctic.
British Antarctic Survey. Scientific Reports 75, 174.
Coulson, J. C., Thomas, C. S., Butterfield, J. E. L., Duncan, N.,
Monaghan, P., and Shedden, C. (1983). The use of head and bill
lenght to sex live gulls Laridae. Ibis 125, 549557.
Croxall, J. P. (1995). Sexual size dimorphism in seabirds. Oikos 73,
399403.
Double, M. C., Gales, R., Reid, T., Brothers, N., and Abbott, C. L.
(2003). Morphometric comparison of Australian Shy and New
Zealand White-capped Albatrosses. Emu 103, 287294.
doi:10.1071/MU03012
Evans, D. R., and Cavanagh, P. M. (1995). Identifying the sex of
Massachusetts Herring Gull by linear measurements. Journal of
Field Ornithology 66, 128132.
Evans, D. R., Hoopes, E. M., and Griffin, C. R. (1993). Discriminating
the sex of Laughing Gulls by linear measurements. Journal of Field
Ornithology 64, 472476.
Forero, M. G., Gonzlez-Sols, J., Hobson, K. A., Donzar, J. A.,
Bertellotti, M., Blanco, G., and Bortolotti, G. R. (2005). Stable iso-
topes revel trophic segregation by sex and age in the southern giant
petrel in two different food webs. Marine Ecology Progress Series
296, 107113.
146 Emu
Gandini, P., Frere, E., and Holik, T. M. (1992). Implicancias de las
diferencias en el tamao corporal entre colonias para el uso de las
medidas morfomtricas como mtodo de sexado en Spheniscus
magellanicus. El Hornero 13, 211213.
Genovart, M., McM inn, M., and Bowler, D. (2003). A discriminant
function for predicting sex in the Balearic Shearwater. Waterbirds
26, 7276.
Gonzlez-Sols, J. (2004). Regulation on incubation shifts near hatch-
ing by giant petrels: a timed mechanism, embryonic signalling or
food availability? Animal Behaviour 67, 663671. doi:10.1016/
j.anbehav.2003.08.008
Gonzlez-Sols, J., and Croxall, J. P. (2005). Differences in foraging
behaviour and feeding ecology in giant petrels. In Sexual
Segregation in Vertebrates: Ecology of the Two Sexes. (Eds K. E.
Ruckstuhl and P. Neuhaus.) pp. 92111. (Cambridge University
Press: Cambridge, UK.)
Gonzlez-Sols, J., Croxall, J. P., and Wood, A. G. (2000a). Foraging
partitioning between giant petrels Macronectes spp and its relation-
ship with breeding population changes at Bird Island, South
Georgia. Marine Ecology Progress Series 204, 279288.
Gonzlez-Sols, J., Croxall, J. P., and Wood, A. G. (2000b). Sexual
dimorphism and sexual segregation in foraging strategies of north-
ern giant petrels, Macronectes halli, during incubation. Oikos 90,
390398. doi:10.1034/j.1600-0706.2000.900220.x
Harrison, P. (1983). Seabirds: An Identification Guide. (Croom
Helm:London.)
Hedd, A., Gales, R., and Brothers, N. (1998). Reliability of morphome-
tric measure for determining the sex of adult and fledgling shy alba-
trosses, Diomedea cauta cauta, in Australia. Wildlife Research 25,
6979. doi:10.1071/WR96121
Hunter, S. (1983). The food and feeding ecology of the giant petrels
Macronectes halli and M. giganteus at South Georgia. Journal of
Zoology 200, 521538.
Hunter, S. (1984). Breeding biology and population dynamics of giant
petrel Macronectes at South Georgia (Aves: Procellariiformes).
Journal of Zoology 203, 441460.
Hunter, S. (1987). Species and sexual isolating mechanisms in sibling
species of giant petrel Macronectes. Polar Biology 7, 295301.
doi:10.1007/BF00443947
Hunter, S., and Brooke, M. D. L. (1992). The diet of giant petrels
Macronectes spp. at Marion Island, Southern Indian Ocean.
Colonial Waterbirds 15, 5665.
Kerry, K. R., Agnew, D. J., Clarke, J. R., and Else, G. D. (1992). Use of
morphometric parameters for the determiantion of sex of Adelie
penguins. Wildlife Research 19, 657664. doi:10.1071/WR9920657
Lo Valvo, M. (2001). Sexing adult Corys Shearwater by discriminant
analysis of body measurements on Linosa Island (Sicilian Channel),
Italy. Waterbirds 24, 169174.
Murie, J. O., Davis, L. S., and McLean, I. G. (1991). Identifying the sex
of Flordland Crested Penguins by morphometric characters.
Notornis 38, 233238.
Patterson, D. L., and Fraser, W. (2003). Satellite tracking Southern
Giant Petrels at Palmer Station, Antarctica. Microwave Telemetry,
Inc. 4, 34.
http://www.publish.csiro.au/journals/emu
S. Copello et al.
Phillips, R. A., and Furness, R. W. (1997). Predicting the sex of Parasitic
Jaegers by discriminant analysis. Colonial Waterbirds 20, 1423.
Quintana, F., and Dell Arciprete, P. (2002). Foraging grounds of south-
ern giant petrels (Macronectes giganteus) on the Patagonian shelf.
Polar Biology 25, 159161.
Quintana, F., Uhart, M., Somoza, G., Cassar, C., Gandini, P., and Frere,
E. (2003). Sex determination of adult rock shags by molecular
sexing and morphometric parameters. Journal of Field Ornithology
74, 370375.
Quintana, F., Schiavini, A., and Copello, S. (2005). Estado poblacional,
ecologa y conservacin del Petrel Gigante del Sur (Macronectes
giganteus) en Argentina. El Hornero 20, 2534.
Renner, M. J., Valencia, L. S., Davis, L. S., Saez, D., and Cifuentes, O.
(1998). Sexing of adults Gentoo Penguins in Antarctica using
morphometric. Colonial Waterbirds 21, 444449.
Selander, R. K. (1972). Sexual selection and dimorphism in birds. In
Sexual Selection and the Descent of Man, 18711971. (Ed.
B. Campbell.) pp. 180229. (Heinemann: Chicago, IL.)
Shaffer, S. A., Weimerskirch, H., and Costa, D. P. (2001). Functional
significance of sexual dimorphism in Wandering Albatrosses,
Diomedea exulans. Functional Ecology 15, 203210. doi:10.1046/
j.1365-2435.2001.00514.x
Sokal, R. R., and Rohlf, F. J. (1995). Biometry. (W. H. Freeman: New
York.)
Tabachnick, B. G., and Fidell, L. S. (1996). Using Multivariate
Statistics. (Harper Collins Publishers: New York.)
Torlaschi, C., Gandini, P., Frere, E., and Martinez Peck, R. (2000).
Predicting the sex of Kelp Gulls by external measurements.
Waterbirds 23, 518520.
Van Franeker, J. A., and Ter Braak, C. J. F. (1993). A generalized dis-
criminant for sexing fulmarine petrels from external measurements.
Auk 110, 492502.
Velando, A., Graves, J., and Freire, J. (2000). Sex-specific growth in the
European Shag Stictocarbo aristotelis, a sexuall dimorphic seabird.
Ardea 88, 127136.
Voisin, J. F. (1982). Observations on the Falkland Islands Giant Petrels,
Macronectes giganteus solanderi. Le Gerfaut 72, 367380.
Voisin, J. F. (1988). Breeding biology of the Northern Giant Petrel
Macronectes halli and the Southern Giant Petrel M. giganteus at Ile
de la Possession, Iles Crozet 19661980. Cormorant 16, 6597.
Voisin, J. F., and Bester, M. N. (1981). The specific status of giant
petrels Macronectes at Gough Island. In Proceedings of the
Symposium on Birds of the Sea and Shore. University of Cape
Town, 1979. (Ed. J. Cooper.) pp. 215222. (African Seabird Group:
Cape Town, South Africa.)
Waugh, S. M., Prince, P. A., and Weimerskirch, H. (1999).
Geographical variation in morphometry of black-browed and grey-
headed albatrosses from four sites. Polar Biology 22, 189194.
doi:10.1007/s003000050409
Manuscript received 28 June 2005, accepted 28 March 2006