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A Generalized Discriminant for Sexing Fulmarine Petrels from External Measurements

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A Generalized Discriminant for Sexing Fulmarine Petrels from External Measurements The Auk 110(3):492-502, 1993 A GENERALIZED DISCRIMINANT FOR SEXING FULMARINE PETRELS FROM EXTERNAL MEASUREMENTS J. A. VAN FRANEKER•,2 AND C. J. F. TER BRAAK•,3 •Institutefor Forestryand Nature Research(IBN-DLO), P.O. Box167, NL-1790 AD Den Burg(Texel), The Netherlands; 2Institutefor TaxonomicZoology, University of Amsterdam,The Netherlands; and •AgriculturalMathematics Group (GLW-DLO), Wageningen, The Netherlands AnSTg•cr.--Discriminant analysiscan use morphometric differencesbetween known male and female birds to predict the sex of unknown individuals in field studies.Geographic variation in size and shapeoften limits the predictive value of a discriminant function to the populationfrom which it was derived. Specificdiscriminant functions for populationsof five speciesof fulmarine petrels(Northern Fulmar,Fulmarus glacialis; Southern Fulmar, F. glacial- oides;Antarctic Petrel, Thalassoicaantarctica; Cape Petrel, Daption capense;and Snow Petrel, Pagodromanivea) assigned 81 to 98%of birdsin the samplesto the correctsex, but the validity of eachdiscriminant applied to alternativepopulations remained questionable. Our approach to overcomethis limitation is to combinedata from the different speciesinto a single dis- criminant. Adequate performanceof this generalized discriminant in samplesof different speciesshows its validity for use in other populationsof any of thesespecies. The generalized function calculatesthe discriminant scoresfor individual fulmarine petrels as: Y = HL + 2.38BD+ 0.41TL- 0.21CL,where HL is head length, BD is bill depth, TL is tarsuslength and CL is bill length (measurementsin millimeters). The cut point to split sexesis different in each sample and may be calculateddirectly from discriminant scores,without reference to sexedbirds, by using a maximum-likelihoodmethod. Depending on species,the gener- alized method resultsin 84 to 97%correct classifications and can be applied to other popu- lations of fulmarine petrels without requiring samplesof birds of known sex. Received19 November1991, accepted20 November1992. FULMARINE PETRELS,like most seabirds, lack terrestrial birds (e.g. Anderson 1975, Green plumage charactersby which sexesmay be rec- 1982).Various approaches are possible.The most ognized. Small differences in mensural char- usual for discrimination between sexes is to acters, however, may reveal sufficient dimor- constructa single formula that calculatesa dis- phism to distinguish sexes. From known criminant score for each individual on the basis correlations between sex and measurements in of its measurements.The cut point to partition a sampleof sexedbirds, a discriminantanalysis scoresinto male and female groups is usually (Sokal and Rohlf 1981) can weight characters taken as the midpoint of the interval between for their power to distinguishgroups (sexes) of the group means of sexedmales and females. unknown individuals. Unfortunately, there are somedrawbacks that Many statistical packages supply computer prevent general usageof publishedsex discrim- programs for discriminant analysis. Comput- inants. Many bird speciesshow considerable erized data processinghas promotedthe useof variation in size over their geographicalrange. discriminant analysis. The method has been As a consequence,the cut point calculatedfor successfullyapplied to a wide variety of seabird one population may not be appropriatefor an- speciesfrom different groups, including pen- other. Also, geographicvariation may involve guins (Scolaroet al. 1983, Gales 1988, Williams shape,which couldaffect the weighting of char- 1990, Williams and Croxall 1991), divers (Okill actersin the discriminantformula. For example, et al. 1989), tube-nosed birds (Dunnet and An- Northern Fulmars (Fulmarus glacialis) from derson 1961, Brooke 1978, Copestakeand Crox- Spitsbergenare not only considerablysmaller all 1985,Sagar 1986, Johnstone and Niven 1989), in overall size, but also have relatively shorter gulls (Shugart 1977,Ryder 1978,Fox et al. 1981, bills than Northern Fulmars from Britain: rel- Nugent 1982, Coulson et al. 1983, Hanners and ative to head length, bill length is about 4% Patton 1985, Schnell et al. 1985), skuas (Hamer shorter in the Spitsbergenbirds (van Franeker and Furness 1991) and also to freshwater and and Wattel 1982, unpubl. data). Wynne-Ed- 492 July1993] SexingFulmarine Petrels 493 T^BLE1. Samplesof birdsused for discriminantfunction analysis. Total numbers,with numbersof sexed males and females in parentheses. Sex determined by Species Locality Dissection Observation Northern Fulmar Netherlands 247 (117, 130) -- Northern Fulmar Jan Mayen 32 (12, 20) -- Southern Fulmar Ardery Island 27 (13, 14) 103 (51, 52) AntarcticPetrel Ardery Island 11 (6, 5) 66 (30, 36) CapePetrel Ardery Island 30 (19, 11) 32 (16, 16) Snow Petrel CaseyStation -- 32 (15, 17) wards (1952) described further variation in bill METHODS shapesand in levels of sexual dimorphism in populations of the Northern Fulmar. Age-de- Samples of sexed fulmarine petrels are character- pendent variation in size or shape can be a fur- ized in Table 1. Northern Fulmars, beached in the Netherlands between 1980 and 1988, were dissected ther complicatingfactor (e.g. Coulson et al. 1983, for morphological and pollution-related studies (van Scolaro et al. 1983). The uncertainties induced Franeker 1983, 1985). Several other North Atlantic by these types of variation urged authors to fulmar populations were studied, but an adequate caution against the unchecked application of sample of sexed birds was available only from Jan their sexdiscriminant to other populations(e.g. Mayen (van Franeker et al. 1986). Southern Fulmars, Fox et al. 1981, Nugent 1982, Witt et al. 1984, Antarctic Petrels,Cape Petrelsand Snow Petrelswere Gales 1988, Hamer and Furness 1991). studied in 1984-1985 and 1986-1987 at the Australian Since 1980 the first author has worked on baseCasey (66øS, 110øE) and the nearby Ardery Island, several projects involving five species of ful- Wilkes Land, Antarctica (van Franeker et al. 1990). marine petrels: Northern Fulmar, Southern Ful- For Antarctic species,two groups of sexedbirds were mar (F. glacialoides),Antarctic Petrel (Thalassoica available: birds sexed by dissection;and birds sexed by observing characteristic behaviors or external antarctica),Cape Petrel (Daption capense),and anatomy. Data on the latter group involved cloacal Snow Petrel (Pagodromanivea). All studies re- evidence (Serventy 1956, Boersmaand Davies 1987), quired knowledge of the sex of birds handled copulation position, egg laying, incubation shifts in the field. In some field projects,birds sexed (Pinder 1966), known sex of partner and, for Snow by dissectionor by field observationsallowed Petrel, voice level (Bretagnolle1990). Although gen- the construction of a discriminant function that erally reliable, a small risk for incidental misinter- could be applied to other birds within the pop- pretation of such observationsmay occur. Therefore, ulation. However, in other study populations, our calculationsin this paper have often been made it was not possible to sex an adequate sample separatelyfor dissectedbirds, and for all sexed birds of birds. In spite of a good general impression (dissection + observation). Skins of dissectedspeci- mens from known breeding localities have been de- of sexual dimorphism in fulmarine petrels, we posited in the care of the Zoological Museum of the were unable to give a reliable prediction of sex- Institute ot Taxonomic Zoology, University of Am- es of birds in those populations where sexed sterdam. individuals were missing. The same problem Measurements.--Measurementsof the following will alsobe encounteredin future projects.Thus, characterswere taken from fresh corpsesor live spec- we decided to reconsider all our data in an at- imens(Fig. 1;Cramp and Simmons1977): bill length tempt to construct a reproducible method to (CL), from edge of implantation of feathers to most discriminate sexes in populations of the ful- distant part of curve of hook (accuracy_+ 0.1 mm); bill marine petrelswithout requiring birds of known depth (BD), from angle of gonysto dorsalsurface of sex. The method calculates(1) a single gener- hook(ñ 0.1 ram);head length (HL), from supraoccip- alized discriminant from data on sexed birds of ital to front edge of bill (_+1ram); tarsus length (TL), from middle of midtarsal joint to distal end of tar- a number of different populations, and (2) pop- sometatarsus(+0.5 mm); wing length (WL), maxi- ulation-specificcut points without referenceto mum flattened chord, from carpal joint to tip of lon- sexedbirds. The predictive power of the meth- gest primary (ñ 1 ram); tail length (TA), central tail od is demonstratedfor populationsof five spe- feathersfrom point where emerging from skin to tip cies of fulmarine petrels. (_+1 ram). Lengths of wing and tail were not always 494 vANFRAIqEKER AND TER BP,•A•< [Auk,Vol. 110 HL fulmarine petrels (Appendix), the sign of the coeffi- cients was chosen in such a way that a score of an unsexed bird above the cut point indicates a male, and values below indicate a female. Discriminant analyseswere carried out with both the original measurementsand their logarithms.Log- arithmic transformation did not improve results and is not discussedfurther. Multiple-regressionanalysis in Genstat supplies information on outliers, such as birds with malelike measurements but sexed as fe- males.Outliers were checkedfor errorsin data entry, but were not omitted from analyses. Reliability of sexassignment by discriminantfunc- tions
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