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Morphological and Vocal Variation Among Subspecies of the Black-Faced Sheathbill ’

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Morphological and Vocal Variation Among Subspecies of the Black-Faced Sheathbill ’ 818 SHORT COMMUNICATIONS ashi Matsudo, David Nobel, and Charles Francis for R. GOLDIZEN.1994. Parentage analysis of multi- providing blood samples. S. C. Lougheed, C. J. Oliar- male social groups of Tasmanian Native Hens nyk and three anonymousreviewers provided helpful (Tribonyx mortierii): genetic evidence for monog- comments on the manuscript. amy and polyandry. Behav. Ecol. Sociobiol. 35: 363-371. LITERATURE CITED GYLLENSTEN,U. B., AND H. A. ERLICH. 1988. Gener- ation of single-strandedDNA by the polymerase BROOKE, M. DE L., AND N. B. DAVIES. 1987. Recent chain reaction and its application to direct se- changesin host usage by cuckoos Cuculus cano- quencing of the HLA-DQ-A locus. Proc. Natl. rus in Britain. J. Anim. Ecol. 56:873-883. Acad. Sci. 85:7652-7656. BROOKE,M. DE L., AND N. B. DAVIES. 1988. Egg mim- MORITZ,C., T E. DOWLING,AND W. M. BROWN.1987. icry by cuckoos Cuculus canorus in rel%on to Evolution of animal mitochondrial DNA: rele- discriminationbv hosts. Nature 335:630-632. vance for population biology and systematics. CABOT,E. 1990. Thi eyeball sequenceeditor, version Annu. Rev. Ecol. Syst. 18:269-292. 1.09d.Dept. Biology,Univ. Rochester,Rochester, NY MOUM, T, AND S. JOHANSEN.1992. The mitochondrial DESJARDINS,R., AND P MORAIS. 1990. Sequenceand NADH dehydrogenasesubunit 6 (ND6) gene in gene organization in the chicken mitochondrial murres: relevance to phylogenetic and population genome: a novel gene order in higher vertebrates. studiesamong birds. Genome 35:903-906. J. Mol. Biol. 212:599-634. MOUM, T., S. JOHANSEN,K. E. ERIKSTAD,AND J. E EDWARDS,S. V. 1993. Mitochondrial gene genealogy PIA~T. 1994. Phylogeny and evolution of the auks and gene flow among island and mainland popu- (subfamily Alcinae) based on mitochondrialDNA lations of a sedentarysongbird, the Grey-crowned sequences.Proc. Natl. Acad. 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BULLOUGH,AND A. SarusCrane (Grus antigone). Auk 113:655-663. The Condor 99:818-825 0 The CooperOmlthological Society 1997 MORPHOLOGICAL AND VOCAL VARIATION AMONG SUBSPECIES OF THE BLACK-FACED SHEATHBILL ’ JOELBRIED AND PIERRE JOUVENTIN Centre National de la Recherche Scientique,’ Centre dEtudes’ Biologiques de Chiz& 79360 Beauvoir sur Niort, France, e-mail: [email protected] Abstract. The Black-faced Sheathbill (Chionis mi- and Kerguelen-Heard) on geographic and morpholog- nor) is a sedentary and polytypic species. Four allo- ical criteria. In the eastern group (Kerguelen-Heard), patric subspeciesare known, each breeding on one ar- corresponding to higher latitudes, sheathbills were chipelago in the Southern Indian Ocean. To evaluate larger and heavier, following Bergmann’s Rule. The the degree of isolation of these four subspecies,mor- sheathbills from Iles Kerguelen also had a lower- phometricsand vocalizationsof adult birds of Iles Ker- pitched voice than those from Iles Crozet, consistent guelen and Crozet were compared with those of the with their larger body size. Moreover, the birds from other localities (Prince Edward and Heard Islands). the southernmostlocality (Heard Island) had a shorter Two groupswere distinguished(Prince Edward-Crozet culmen, consistentwith Allen’s Rule, but longer tarsi and deeper sheaths.Within the western group (Prince Edward-Crozet), and at Iles Kerguelen, there also was ’ ’ Received 8 July 1996. Accepted 13 March 1997. variability on a microgeographicalscale. Differences SHORT COMMUNICATIONS 819 Kerguoelen Crozet ;:a Prince Edward He&d \ ANTARCTICA FIGURE 1. Breeding range of the Black-faced Sheathbill in the SouthernIndian Ocean. Island namesindicated in bold. between subspeciesof Black-faced Sheathbill there- may be achieved by combining morphological and fore could be due not only to environmentalcorrelates ethological studies. of latitude, but also to possible genetic drift. The four subspecies are allopatric and do not differ in their METHODS breeding scheduleor in their general behavior and diet, Field work was conductedat Iles Kerguelen (48”27’- suggestingthat differentiation may be recent and main- 5o”S, 68”37’-70”35’E) in late December 1992, and at ly due to geographicalisolation. Ile de la Possessionin the eastern part of the Crozet archipelago(45”50 ’-46”3O’S, 50”-53”21’E) in Novem- Key words: speciation, reproductive isolation, ber 1994. Adult Black-faced Sheathbills were caught morphometrics,vocalizations, Chionis minor, Black- using a shepherd’s hook. We identified them as adults faced Sheathbill. becausethey were ringed, and known to have bred in the past or to be at least 3 years old, which is the The biological speciesconcept (Mayr 1970) considers minimum age of first breeding (Burger 1980a, Jouven- speciesas groupsof naturalpopulations capable of in- tin et al. 1996). Measurements (wing length, tarsus terbreedingand separatedfrom other similar groupsby length, culmen length, culmen width, and sheath mechanisms of reproductive isolation, such as geo- depth) were taken for each individual, following Bur- graphical, morphometrical,coloration, behavioral, and ger (1980b). Black-faced Sheathbills are sexually di- phenological. However, variability within the range of morphic with males larger than females (Burger a given species also can occur. Thus, geographical 1980b). Therefore, birds were sexed from size and bill variations in body size and coloration are well docu- shape index (i), the latter defined as follows (Warham mented in seabirds,particularly in Procellariiforms or 1972, Burger 198Ob): in the White-tailed Tropicbird Phaethon lepturus (de1 i = culmen length X culmen width Hoyo et al. 1992). Geographicalreproductive isolation X sheath depth X 0.1 also seems to play a major role in the Black-faced Sheathbill Chionis minor. Four subspeciesusually are Morphometrics of birds from Iles Kerguelen were recognized from body size, leg coloration and carun- comparedwith those from Prince Edward Islands (data cles, each subspeciesbreeding on a remote archipelago from Marion Island, A. E. Burger, pers. comm.) and in the Southern Indian Ocean (Marchant and Higgins Be de la Possession,using univariate statistics(Krus- 1993): C. m. marionensisat Prince Edward Islands, C. kal-Wallis H-tests and multiple comparisonpairwise z- m. crozettensisat Iles Crozet, C. m. minor at Iles Ker- tests; Scherrer 1984). We obtained the length of the guelen, and C. m. nasicornisat Heard Island (Fig. 1). appendagesrelative to individual body size by dividing Black-faced Sheathbills are sedentary and have never tarsusand culmen length by wing length (see Aldrich been recorded on the nearestcontinent or more than a and James 1991). However, wing length varies in the few kilometers offshore, becauseof their poor flying course of the year due to molt or abrasionof the outer abilities (Verheyden and Jouventin 1991). primaries. We measuredwing length during the austral This paper provides new and more accurate infor- spring and assumedthat the abrasionstage of the outer mation about biometry of Chionis minor than previ- primaries was similar in both Iles Crozet and Kergue- ously available. New insight into the speciation of len, which allowed us to compare these two localities Chionis minor and the role of reproductive isolation only. Multivariate analyseswere conducted using the 820 SHORT COMMUNICATIONS SAS statisticalpackage (SAS Institute 1988); variables tarsi were significantly longer on Iles Crozet (Mann- were not transformed.A principal componentanalysis Whitney U-test, z = 3.69, P < 0.001, see Table 1). on correlation matrix between the variablesused (Dig- For Iles Kerguelen birds, measurementsfrom skins by and Kempton 1987) was conducted, using the were significantly smaller than those obtained from PRINCOMP procedure. Discriminant analyses also live birds (Mann-Whitney U-test, all P < O.OOl), ex- were performed by running a stepwise analysis in or- cept for culmen length and sheathdepth (sheath depth der to determine the two most discriminantparameters was significantly higher for skins in females: Mann- (STEPDISC procedure), before subsequentuse in the Whitney u-test, z = 2.39, P < 0.02). Kerguelen spec- discriminant function analysis (nonparametricmethod imens had a longer culmen than those from Heard Is- of the DISCRIM procedure using uniform kernel and land (at least for males, Mann-Whitney U-test, U = a posteriori cross-validationtechniques for estimating 14.5, rz, = 9, n, = 10, P < 0.01). Conversely, the latter error rates in classification). tended to have longer tarsi and deeper sheathsthan the Data from skins of birds from Heard Island and Iles former, althoughdifferences were not significant,prob- Kerguelen
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