Sexual Dimorphism in the Adult South African (Cape) Fur Se^\ Arctocephaluspusilluspusillus (Pinnipedia: Otariidae): Standard Body Length and Skull Morphology C. L. Stewardson', T. Prvan^, M. A. Meyer^ and R. J. Ritchie''* 'Botany and Zoology, Australian National University, Canberra, ACT 2601, Australia. (PresentAddress, Fisheries and Marine Sciences Program Bureau ofRural Sciences, The Department of Agriculture, Fisheries and Forestry, Canberra, ACT 2601, Australia). NSW ^Department ofStatistics, Macquarie University, 2109, Australia. ^Marine and Coastal Management (MCM), Rogge Bay, Cape Town, South Africa. NSW 'School ofBiological Sciences, The University ofSydney, 2006, Australia. *CorrespondingAuthor: Raymond J. Ritchie, School ofBiological Sciences, The University ofSydney, NSW 2006, Australia, email [email protected]. Stewardson, C.L Prvan, T., Meyer, M.A. and Ritchie, R.J. (2010). Sexual dimorphism inthe adult SouthAfrican (Cape) farsealArctocephaluspusilluspusillus (Pinnipedia: Otariidae): standard body length and skull morphology. Proceedings oftheLinnean Society ofNew South Wales 131, 119-140. We examine differences in standardbody length and skull morphology ofmale (n= 65) andfemale (n= 18) SouthAfrican (Cape) fur seals, Arctocephahispiisilhispiisilliis, from the coast of southern Africa with the aim to develop an objective method for determining the sex offur seal skulls. Males were found to be significantly larger than females in standard body length, with K-means cluster analysis successfully identifying 2 relatively homogeneous groups. Principal component analysis (covariance matrix) showed that the underlying data structure for male and female skull variables was different, and that most ofthis variation was expressed in overall skull size rather than shape. Males were signiticantly larger than females in 30 ofthe 31 skull variables. Breadth ofbrain case was significantly different for the genders. Relativetocondylobasal length,males were significantly largerthan females in 13 ofthe 31 skull variables used in the present study. These were gnathion to posterior end ofnasals, breadth at preorbital processes, least interorbital constriction, breadth at supraorbital processes, greatest bicanine breadth, breadth of palate at postcanine 1 and 3, calvarial breadth, mastoid breadth, gnathion to anterior of foramen infraorbital, gnathion to posterior border of preorbital process, height of skull at base of mastoid and height ofmandible atmeatus. In males, these variableswere associatedwith the acquisition anddefenseofterritory(e.g., largeheadsize andmass; increasedstructuralstrengthoftheskull; increased bite capacity). Two skull ratio parameters, breadth ofbraincase/condylobasal length and length ofupper postcanine row/condylobasal length were significantly higher in females compared to males. Based solely on the skull data, mature males can be reliably distinguished from immature males and females using both (a) Classification and Regression Tree (CART) and (b) Hierarchical ClusterAnalysis. Both approaches had difficulty in reliably distinguishing immature males from females. The Classification and Regression Tree methodwas the more successhil in correctly distinguishing immature males from females. Manuscript received October 2009, accepted for publication 21 April 2010. 1 KEYWORDS: Arctocephahis pusilhis pusilhis, identification of sex, muldvariate analysis, Otariidae, polygyny,Pinnipeds,principlecomponentandcladisticanalysis,sexualdimorphism,skullmorphometries, SouthAfrica fur seal, standard body length. SEXUAL DIMORPHISM IN ARCTOCEPHALUS PUSILLUS PUSILLUS INTRODUCTION Adult male fur seals are about 3 to 5 times heavier and about 1/4 longer than adult females Sexual dimorphism isa form oFnon-gcographic (Stirling, 1983; David, 1989; Boncss, 1991; Guinet et variation tliat can be generated in a species by al., 1998; Arnould and Warneke, 2002; Stewardson et the process of sexual selection (Bartholomew, al., 2009). Large body size is in itselfan intimidating 1970; Alexander et al., 1979; Stirling, 1983). form of display to discourage rival males from Highly polygynous species such as fur seals, sea attempting an actual physical challenge and in the lions and elephant seals, generally exhibit a high event of a physical challenge is advantageous in degree of sexual dimorphism (Laws, 1953; competitive interactions and enables breeding bulls Ralls, 1977; Alexander et al., 1979; Stirling, to remain resident on territories for longer periods 1983; Sirianni and Swindler, 1985; McLaren, of time without feeding (Rand, 1967; Miller, 1975; 1993; Arnould and Warneke, 2002). Differences Payne, 1978, 1979; Stirling, 1970, 1983). Strong fore- in reproductive success among males ofthese species quarters, enlarged jaw and neck muscles, robust are large, and competition for access to females canines, increased structural strength of the skull, is intense. Selection pressure appears to favour and long, thick neck hair (protective mane or wig), the development of traits that enhance male also appear to be potentially advantageous in the fighting ability, including intimidating body acquisition and maintenance of territory; quan- size, weaponry and skin thickness (Laws, 1953; titative information on these features, however, are Bartholomew, 1970; Le Boeuf, 1974;Alexander et al., lacking (Miller, 1991). 1979;McCann, 1981; Stirling, 1983). Here we examine morphological differences Breeding Southern fur seals (Arctocephaliis between skulls (n = 31 variables) of male (n = spp.) are among the most territorial of animals, 65) and female (n = 18) South African (Cape) fur are strongly sexually dimorphic in body size, sealsArctocephaluspusilluspusillus, from the coast polygynous and gregarious (Peterson, 1968; Harrison of southern Africa. Body length information was etal., 1968; Stirling, 1970; Bryden, 1972; Alexander also included in analyses where available. Where et al, 1979; Bonner, 1981; McKenzie et al., 2007). possible, comparisons aremadetothe closelyrelated In the southern hemisphere, breeding status male Australian fur seal Arctocephaluspusillus doriferus fur seals (beachmasters) generally arrive at the (King, 1969; Brunner, 1998ab, 2000; Brunner et rookeries around November to establish ter- al., 2002; Arnould and Warneke, 2002; Brunner et ritories. Pregnant females arrive soon after. Once al., 2004; Stewardson et al, 2008, 2009) and other females are present in the male's territory, males otarid species for which morphological data are guard females until they come into oestrus post- available such as the Steller sea lion {Eumetopias partum. Females give birth within one week of jubatus) (Winship et al., 2001). coming ashore and then mate with the nearest For many life history, conservation and male during the short breeding (pupping/ mating) ecological studies it is important to be able to season (Guinet et al., 1998). Males seldom leave determine the sex of skull material in museum the territory until the breeding season is over (Rand, collections, skulls of animals found dead or 1967; Stirling, 1970; Miller 1974; Peterson, 1968; accidentally killed in fishing operations or killed in Harrison et al, 1968; Bonner, 1981). After mating, other ways. Often only the skull is available. We the territorial system gradually breaks down and show that two types of multivariate analysis [(a) males return to sea to replenish their physiological Classification and Regression Tree (CART) reserves. Males do not care for their young. and (b) Hierarchical Cluster Analysis] can be used When establishing territories, male fur seals to objectively distinguish mature male, immature threaten each other with vocal and visual displays, male and female skulls ofthe South African fur seal emphasising their size, to intimidate competitors {A. pusillus pusillus). By extension the approach (Bonner, 1968; Stirling, 1970; Stirling and Warneke, could be applied to other fur seals, particularly the 1971; Miller, 1974; Shaughnessy and Ross, 1980). Australian fur seal {A. pusillus doriferus) and the Much time is spent in making visual and vocal threats New Zealand fiir seal {A. australisforsteri). to rival males and chasing them away, but fights may develop, occasionally resulting in severe injury or death (Rand, 1967; Stirling, 1970; Shaughnessy MATERIALSAND METHODS and Ross, 1980; Trillmich, 1984; Campagna and Le Boeuf, 1988). Collection ofspecimens South African (Cape) fur seals {Arctocephalus 120 Proc. Linn. Soc. N.S.W., 131, 2010 C.L. STEWARDSON, T. PRVAN, M.A. MEYERAND R.J. RITCHIE pusilhis pusilhis) were collected along the Eastern Museum records Cape coast of South Africa between Plettenberg The data setonthe males used inthepresent study Bay (34° 03'S, 23° 24'E) and East London (33° has alreadybeenpublished in (Stewardson et al., 2008) 03'S 27 54'E), fromAugust 1978 to December 1995 and fiarther details can be found in Stewardson (2001). (Stewardson et al., 2008, 2009), and accessioned The list ofmale specimens used in the present study is at the Port Elizabeth Museum (PEM). Specimens shown in Appendix 1. There were 39 adult males, 24 were collected dead or dying from the coastline and immature sub adult males and twojuvenile males only some from accidental drowning in fishnets; none 2 years old. No standard body length measurements were deliberately killed (cf. Guinet et al., 1998). were available on four (4) of the adult males (PEM Routine necropsies were performed and biological 2004, PEM 2007, PEM 2013, PEM 2036) but it is parameters recorded, based on recommendations unlikely that any adult male skulls wouldbe assigned of the Committee on Marine Mammals (1967). tothewrongsexbecausematuremale skulls aremuch Animals were aged from incremental lines observed largerthan females and more heavily built. However, in the dentine of upper canines (Stewardson et al., there were no SBL measurements available on four 2008, 2009). The sample was supplemented with (4) of the immature males (PEM 2006, PEM 2009, measurements from 1 1 known-aged adult males PEM 2010 and PEM 2014). This raises some doubts (animals tagged as pups) from Marine and Coastal aboutthecertaintythatthesespecimenswerecorrectly Management (MCM), Cape Town. The specimens identified as males. Generally if the SBL had been MCM from the collection have accession numbers determined, the genitalia would have been available MCM MCM MCM beginningwith (e.g. 1809). The for examination. The raw data set for the females collection also housed 5 tag-aged adult females (18 adults, 4 juveniles and sub adults) is shown in and 3 tag-aged sub adult/juvenile females. Appendix 2 and the means and standard deviations All animals considered adults had reached full inAppendix 3.All the female carcasses were complete reproductive capacity, i.e., males > 8 y (Stewardson enough forreliable determination oftheir sex. et al., 1998; Stewardson et al., 2008, 2009) and females > 3 y (J.H.M. David, pers. comm.). When Skull variables age was not known, males > 170 cm (Stewardson et A total of 32 skull measurements were recorded al., 2008, 2009) and females > 135 cm (Guinet et al., (Table 1). However, one ofthese variables, height of 1998; J.H.M. David, pers. comm.) were considered sagittal crest, was not examined statistically because fully adult males and females and included in the there were few measurements for females and also analysis as adults even iftheir dentition age was less becausewehavefoundthatsagittalcrestmeasurements than 8 y for males. South African fur seals > 12 y seemtoprovidelittleusefialinformationinmaleskulls cannot be aged from counts of growth layer groups (Stewardson et al., 2008). Thus, statistical analysis (GLG) in the dentine of upper canines because of was conducted on 31 of the 32 variables. Skull closure of the pulp cavity. Estimated longevity for preparation and measurement procedures follow male SouthAfrican Furseals is c. 20y(Wickens, 1993; Stewardson et al. (2008). Stewardson et al., 2008, 2009). There is much less information on the longevity offemale South African Statistical analyses fiar seals (despite the large numbers ofanimals that are Six methods ofanalyses were employed. Firstly, shot in culling and hunting operations) but Wickens two sample t-tests (assuming equal variance) were (1993) based on zoo records concluded that females used to test the hypothesis that the mean value of a could liveto c. 30 y. skull variable was significantly different for males Australian male fur seals {A. pusilhis dorifems) and females against an appropriate alternative alsohaveasimilarlifespanofabout20yearsbutfemale y^ ^ 0' HTiales f'^females' "I'r^males f'^femafes' 1" ^females Australian fiar seals based on age tags are currently > 111^3,^3). Since more than 1 skull variable was being knownto live towell over20y (Amould andWameke, considered, the Bonferroni correction was used - the 2002). Seal life spans in arange ofseal species average experiment-wise error rate was divided by the total about 15 to 20 y for males and in excess of20 y for number oftests performed (Cochran, 1977). females (New Zealand fur seal {A. australlsforsteri), Secondly, K-means clustering, a non- McKenzie et al., 2007; Antarctic fur seal {A. gazelld), hierarchical cluster analysis was used to classify Payne, 1978, 1979); Steller sea hon {Eumetopias observations into 1 of2 groups based on some ofthe jubatus), Winship et al., 2001). skull variables. Observations on some of the skull variables fromboth sexeswerepooled sothat initially there is a single cluster with its centre as the Proc. Linn. Soc. N.S.W., 131, 2010 121 SEXUAL DIMORPHISM IN ARCTOCEPHALUS PUSILLUS PUSILLUS moan vector of the variables considered. These PLUS (MathSoft, Inc., Seattle, 1999, 5.1). observations were then assigned at random to two Fifthly, the data mining approach. Classification sets. Step entails calculating the mean vector of and Regression Trees (CART), a technique that 1 the variables considered (ccntroid) for each set. generates a binary decision tree, was used to classify Step 2 entails allocating each observation to the the observations. In this approach, the set of data is clusterwhosecentroid is closest to that observation. progressivelysub-dividedbasedonvaluesofpredictor These two steps are repeated until a stopping variables into groups that contain higher proportions criterion is met (there is no further change in the of "successes" and higher proportions of "failures". assignment of the data points). Before doing this The relative importance of the predictor variables all variables were standardised. Closest neighbour is assessed in terms of how much they contribute to (similarity) was measured using Euclidean distance successful splits into more homogeneous sub-groups. (Johnson and Wichem, 1992). The groupings ofskull The classification is most commonly earned out using variables we considered were dorsal, palatal, lateral the Gini criterion, which always selects the split that and mandibular. We also used k-means clustering to maximises the proportion of "successes" in one of classify observations into 1 of2 groups using standard the groups (Petocz, 2003). Data mining techniques body length. are attractive because no distributional assumptions Thirdly, plots oflog,ofeachskullvariableagainst are needed, data sets can have missing data and log^ofstandard body length (SBL) for the genders analyses are less time consuming. The training data were examined. 'Robust' regression (Huber M- usedto createthe binary decision setwas the setofall Regression) was usedto fit straight lines (log y = log a animals that have already been determined to be adult + b log x) to the transfoiTned data (Weisberg, 1985; males, immature males and mature females. SPSS Myers, 1990). Clementine 12.0 was used for the analysis. Fourthly, principal component analysis (PCA) Finally, Minitab was also used to perform was used. Oneuseful application ofPCAisidentifying hierarchical clustering and produce dendrograms the most important sources ofvariation in anatomical showing the degree ofsimilarity ofthe skull data for measurements for various species (Jackson, 1991; males, females and immature males. In general, the Jolliffe, 2002). When the covariance matrix is used and conclusions reached were similar to those from the the data has not been standardized the first principle CART analysis: it was possible to distinguish mature component (PC)usuallyhas all positive coefficients males from immature males and mature females but and according to Jolliffe (2002) this reflects the it was not possible to clearly distinguish immature overall 'size' of the individuals. The other PCs males from females. usually contrast some measurements with others Unless otherwise statedvalues are means quoted and according to Jolliffe (2002) this can often be ± standard errors with the number of data points in interpreted as reflecting certain aspects of 'shape', brackets. which are important to the species. Skull measurements were recorded in the same units; therefore a covariance matrix was used to RESULTS calculate PCs (however this gives greater weight to larger,andhencepossiblymorevariablemeasurements Standard body length (SBL) because the variables are not all treated on an equal SBL ranged from 157-201 cm in males (n = footing). Genders were examined separately 33, SBL was not recorded for 4 of the adult males) because the grouped PCA was quite different, and 135-179 cm in females (n = 18). Mean lengths in most cases, to either the separate male PCA or were 182.9 ± 2.3 (n = 33) and 149.1 ± 2.5 (n = 18), female PCA. respectively. The two sample t-tests on our data PCA and two sample t-tests were calculated in indicated that adult males were significantly larger Minitab (Minitab Inc., Slate College, 1999, 12.23). than adultfemales (Table 1). Theratioofmean female K-means cluster analyses for skull variables and SBL to mean male SBL was 1:1.23. SBL were calculated in Minitab (Minitab Inc., K-means clusteranalysis successfully identified2 Slate College, 1999, 12.23) and in SPSS (SPSS Inc., relatively homogeneous groups from the pooled Chicago, Illinois, 1989-1999, 9.0.1), respectively. data, i.e., cluster 1,predominantlymales andcluster2, This was necessary because Minitab could only predominantly females (Table 2). Ofthe 18 females, perform K-means cluster analysis for 2 or more 17 (94%) were correctlyclassified. Ofthe 33 males, 28 variables, therefore SBL (a single variable) was (85%) were correctly classified. analysed in SPSS. The regressions were fitted in S- 122 Proc. Linn. Soc. 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All variables except standard body length (SBL) were standardised (dorsal, palatal and mandibular). Skullvariables Sex Cluster1 Cluster2 n Male 22 (96%) (4%) 23 1 Dorsal Female 11 (100%) 11 Male 24 (92%) 2 (8%) 26 Palatal Female 17(100%) 17 Male 28 (80%) 7 (20%) 35 Lateral Female 10(100%) 17 Male 25 (93%) 2 (7%) 27 Mandibular Female (6%) 16(94%) 17 1 Male 28 (85%) 5(15%) 33 Standardbodylength Female (6%) 17(94%) 18 1 Skull variables of mandibular tooth row (M29) (Table 1). Height Absolute skull size: two sample t-tests ofsagittal crest (L27) was not examined statistically The two sample t-tests indicated that 30 of because there were too many skulls with missing or the 31 mean skull variables were significantly damaged sagittal crests. larger in males than in females, i.e., we reject H^ in favour ofH^: \i^^^^^^>\i^^^^^^^(Table 1, Fig. 1). Mean value Relative skull size: two sample t-tests of breadth of brain case (D9) was not significantly When skull variables were analysed relative to different for the genders (Table 1). The coefficient condylobasal length (CBL, Dl), males were found of variation (C.V.) was larger in males, with the to be significantly larger than females for 13 (43%) following exceptions: least interorbital constriction variables: (1) gnathion to posterior end of nasals (D7), breadth of brain case (D9), gnathion to (D3), (2) breadth at preorbital processes (D8), (3) anterior of foramen infraorbital (L24) and length least interorbital constriction (D7), (4) breadth at supraorbital processes (D8), (5) 250 greatestbicanine breadth (PI2), (6) .' breadth of palate at postcanine 1 200 Fig. 1: Mean values of31 skullvar- iables for male and female South African fur seals. Numbers cor- 150 respond to skull variables listed in Table 1 (numbers 1-9 correspond to parameters Dl to D9, 10-23 to OJ PIO to P23 and 24-32 to L24 to 100 L32). Numbers above the dashed line, males > females; numbers on the line, males = females; numbers 50 below the line, females > males. Minitab could only perform K- means cluster analysis if there was > 2 variables, therefore SBL (a single variable) was analysed in o 50 100 150 200 SPSS. SBL was not recorded for 4 Females (cm) ofthe 39 males (i.e., n = 35). 126 Proc. Linn. Soc. N.S.W., 131, 2010 1 ex. STEWARDSON, T. PRVAN, M.A. MEYERAND R.J. RITCHIE The remaining 15 (50%) variables were not significantly different for the genders (Table 1). Since males were larger than females in 'absolute size', this suggested that the 15 variables were proportionate to CBL regardless of sex, i.e., the ratio relative to CBL (Dl) was E significantly different for the genders. CO The coefficient of variation o for values 'relative to CBL' OS was larger in males for about 1/3 rd of all variables (Table 1). Exceptions were breadth at pre- orbital processes (D6), least interorbital constriction (D7), palatal notch to incisors (PIO), breadth of zygomatic root of maxilla (P14), breadth of palate at postcanine 5 (P17), gnathion to foramen infraorbital (L24), 0.4 0.6 0.8 gnathion to hind border of Females (cm) preorbital process (L25), length of mandible (M28) and length of mandibular tooth row Fig. 2: Mean values of 30 skull variables, relative to condylobasal (M29). The coefficients of 2 of length, for male and female South African fur seals. Numbers corre- these variables (least interorbital spond to skull variables listed in Table 1 (numbers 1-9 correspond to constriction (D7) and length of parameters Dl to D9, DlO-23 to PIO to P23 and P24-32 to L24 to L32). mandibular tooth row (M29)) Numbers above the line, males > females; numbers on the line, males were considerably larger in = females, numbers below the line, females > males. females in both 'absolute size' and size 'relative to CBL' (M29/ (P15), (7) breadth ofpalate at postcanine 3 (P16), Dl and D7/D1). (8) calvarial breadth (P21), (9) mastoid breadth (P22), (10) gnathion to foramen infraorbital K-means cluster analysis (L24), (11) gnathion to hind border of preorbital K-meansclusteranalysissuccessfullyidentified process (L25), (12) height of skull at bottom of 2 relatively homogeneous groups from the pooled mastoid (L26) and (13) height of mandible at data, i.e., cluster 1, predominantly males and cluster meatus (M31) (Table 1, Fig. 2). Differences between 2, predominantly females (Table 2). Classification the genders were highly significant (P < 0.001); apart based on dorsal, palatal and mandibular obser- from gnathion to foramen infraorbital (L24) and vations was highly successful in recapturing the 2 height of skull at bottom of mastoid (L26), which groups. Classification based on lateral observations were significant at the 5% level (Table 1). was less successful. Breadth of brain case (D9) was significantly Apart from 1 mandibular variable, all females different in 'absolute size' for males and females, were correctly classified. The majority ofmales were but 'relative to CBL' parameterD9/D1 forfemales correctly classified with the following exceptions - was larger than males (Table 1). Length of upper dorsal, 2 palatal, 2 mandibular and 7 lateral variables postcanine row (Pll) was larger in 'absolute size' in were incorrectly classified as females (Table 2). males, but 'relative to CBL' Pll/Dl in females was Misclassification occurred in small males only. larger than in males (Table 1). Proc. Linn. Soc. N.S.W., 131, 2010 127 SEXUAL DIMORPHISM IN ARCTOCEPHALUS PUSILLUS PUSILLUS Fig. 3 Fig. 4 4.2 #• 4.1 E E E 5.5 4.0- •.'*. ^ >^ C 5.4 D jvi. 3.9 OJ • ftT* • if> 5.3 DQ 3.8- X"' Q o <cU J/ >O, 5.2 'c 3.7- oJ (o0 /•• # .v>- 5.1 CO 3.6- ••• X" • ><^ > 3.5- 4.4 4.5 4.6 4.7 4.8 4.9 5.0 5.1 5.2 5.3 03 /•y y Ln Seal Body Length (cm) <1u— / • ^• CD 3.4- X c y _l • ^ 3.3- Fig. 5 4.4 4.5 4.6 4.7 4.8 4.9 5.0 5.1 5.2 5.3 5.1 Ln Seal Body Length (cm) 5.0 Figs. 3, 4 & 5: Bivariate plot of: (3) log [CBL (Dl) 4.9 (mm)] on log (SBL(cm)); (4) log [greatest bicanine breadth (P12) (mm)] on log (SBL (cm)); (5) log 4.8-1 [mastoid breadth (P22) (mm)] on log (SBL (cm). Circles, males. Squares, females. c 4.7 0) o 4.6 The first 3 PCs accounted for most of the "o variation. The first PC (PCI) can be interpreted w 05 4.5 as a measure of overall skull size while PC2 and PC3 define certain aspects of shape (Table 3). 4.4 Interpretations for the first 3 PCs for the 2 genders are given in Table 4, together with the percentage of 4.3-1 total variation given by each PC. The variances of 4.2 corresponding PCs forthe two genders do vary and I ' "'I 1 ' I ' 1 I I interpretations are dissimilar for mostpairs ofPCs. 4.4 4.5 4.6 4.7 4.8 4.9 5.0 5.1 5.2 5.3 Ln Seal Body Length (cm) Determining the gender ofan isolated skull It is claimed that it is often possible to make a visual determination of the gender of an isolated Linear regression South African fur seal skull, provided the skull is All transformed variables were regressed on from an adult animal (Brunner, 1998ab). However, log, (SBL in cm). Three variables that best depicted visual identification based on morphology of the maximum discrimination between the sexes, using skull alone can be misleading, e.g., young adult regression, are given in Figs. 3, 4 and 5. These males can be mistaken for larger, older females were CBL (Dl), greatest bicanine breadth (P12) and sex determination of a pup from examining and mastoid breadth (P22). These plots (males the skull alone would be very difficult. A more closed black circles, females grey squares) clearly objective procedure in determining sexes of skulls show pronounced sexual dimorphism in adult South would be desirable. In most practical situations if African fur seals, supporting findings of the two- the carcass was available for examination, the sex sample t-test and K-means cluster analysis. would usually be detenninable, however for many museum specimens only the skull is available. The Principal component (PC) analysis 128 Proc. Linn. See. N.S.W., 131, 2010