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Taxonomy and Identification of Steamer-Ducks (Anatidae : Tachyeres)

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Taxonomy and Identification of Steamer-Ducks (Anatidae : Tachyeres) Taxonomy and Identification of STEAMER-DUCKS (Anatidae: Tachyeres) Bradley C. Livezey and Philip S. Humphrey u^C ^^, ^ Ernst Mayr lArary Zoology - P ^ ^ %weiim of Compara»ve Harvard University THE UNIVERSITY OF KANSAS. MUSEUM OF NATURAL HISTORY PUBLICATIONS The University of Kansas Publications. Museum of Natural History, beginning with volume I in 1946. was discontinued with volume 20 in 1971. .Shorter research papers formerly published in the above series are now published as The Universitv of Kansas Museum of Natural History Occasional Papers. The University of Kansas Museum of Natural History Miscellaneous Publications began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural llisioiy were initiated in 1970. Authors shouldcontact the managing editor regarding style and submission procedures before manuscript submission. All tiianuscripts are subjected to critical review by intra- and extramural specialists; final acceptance is at the discretion of the Director. This publication is printed on acid-free paper. Occasional Papers and Miscellaneous Publications are lypeset using Microsoff " Word and Aldus PageMaker" on a Macintosh computer. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publications by addressing the E.xchange Librarian, The University of Kansas Library, Lawrence. Kansas 6604.'S-2K0() LISA. Individuals may purchase separate numbers from the Office of Publications, Muscimh oI Nalmal History, The LIniversily of Kansas, Lawrence, Kansas 6604.^-24.S4 I SA. f-'ninl Cover: ihc jour species oj Sieaiueriliicks asjollows: I pper lefi \\ liiieheiiiled I'll^^litlcss Sieanier- (liicks (I'hoto;^rapli h\ R Siraiiei ki: Cpper rii^hl—Moiielluiue I- li;^luless Steamer -iliieks iPlioloi^rapli h\ — ('. D. /.immenmin): Lower left iatklaiul l-'lii;li!less Slcamer-dueks I Plioloi;riiph In- B l.i\e:e\): unci Lower rii>lu—Llyliii; Stcamer-diiek (Photoiiraph by (!. L. Niieclilerlein). oc so in TtiK UNivi-;Rsin oi Kansas Museum oi- Naturai. History MoNOCK \I'H No. S 10 Docciiiher 1^)42 Taxonomy and Identification of Steamer-Ducks (Anatidae: Tachxercs) Bradley C. Livezey and Philip S. Humphrey Museum (if Natural llisioiy The ihiivcrsiix of Kansas LawiciHC. Kansas hM)45-2454 USA MUSEI'M OF Natlral Hisiorv Dyche Mali Thh UNi\r:RsiTV ni- K\nsas Lawri \i I . K \nsas MONOGRAPHS EJilor fur litis issue: RiLluird [•. Jolinsloii Mdiuiiiiiii; lulilar: Joseph "I'. Collins MCZ JUL 9 3 UNlVcI-SlTY Monogia|ih No. S Pp. i-l2.'i; 2y figures (2 in eolor); 13 tables: 1 appemiix Published 10 December 1992 * ISBN: 0-89338-042-3 © 1992 H\ Ml sci'M or NAruRAr HisroKi DVCHE Hali. Till-: University or Kansa.s Lawrenci;. Kansa.s 66045-2454, USA Primi i> m Usrvi Rsiiv oi Kansas Pkintini; Shrvick l.AWR[-N( I . Kansas 1 contp:nts INTRODUCTION 1 ACKNOWLEDGMENTS 1 METHODOLOCilCAL NOTES 2 Specimens - Molls and Plumages 2 Measiiremenls 2 Slalistical Analyses 3 ORNITHOLOC.ICAL HISTORY OF THE STEAMER-DLICKS 3 Early Deseiiplions 3 Flighted and Flightless Species 4 A Second Flightless Species 1 A Fourth Species 12 Overview 14 SYSTEMATIC REVIEW 15 Relationships ol l\it. Iiwres 15 Synonomies lor lac hycres 16 Genus Tcuiiycirs 18 Flying Steamer- Duck tTachyercs piitachoniciis) 20 Falkland Flightless Steamer-Duck iT(i< hvcirs Ivuchyplcins) 27 Magellanic Flightless Steamer- Duck iTdclncifs piciicrcs) 30 White-headed Flightless Steamer-Duck iTik Incnw Icucm cpluiliis) 35 MOLTS AND PLUMAGES 36 History of Study 36 Comments on Descriptions 46 Ten llVCIVS pilhll hdiiicus 48 Tachycrc.s hnuiiyplcnis 54 Tackycrvs picncrcs 56 Tachycics Icmoccphalus 59 DIAGNOSTIC CHARACTERS 63 Generic Characters 63 Detennination of Age 65 Qualitative Characters of Species 67 Quantitative Characters of Species 75 KEYS TO IDENTIFICATION OF SPECIMENS 90 Skins 90 Skeletons 91 FIELD IDENTIFICATION 92 Introductory Comments 92 Ecological Segregation 92 General Appearance 93 Behavior 93 Vocalizations 94 STEAMER-DLiCKS IN CAPTIVITY 95 PROSPECTS FOR THE FUTURE 97 SUMMARY 98 RESUMEN 99 SPECIMENS EXAMINED 101 LITERATURE CITED 105 APPENDIX 121 ) INTRODl'CTION Durmy our leii-scar stLkl\ ol the sieaiiicr-diiLks prone toinersmiplilicalion ("lumping ol s|iecies'"). view {Tcichycrc.s ). a moderately speeiali/ed Soulli Aiiici i- ap|iarentl\ because of the traditional that can genus of shelducks (Tadorninae: IJve/es s|iecies of waterfowl should be reatlily distin- 1986a). our piiniar\ inierests have lieen llie mor- guishable using plumage characters, even under phological bases and adapli\e signiticanee ol held conditions. tlighllessness. geographic variation, sexual diiiioi- Ihe present pa|iei has six major objectives: ( I |ihisni, and phylogenetic relationships ol the loui to present a history of the study of the genus. species. ,-\s these investigations progressed, hovv- v\liich in turn will provide an insight into the its e\ei-. :t became apparent that basic taxoiiomic complicated taxonomy that has been applied to issues, including alpha-ta\ononi\ and the ilesig- members; (2) to provide generic aiul specific syn- nation of types, and the data and techniques asso- onviifies and species accounts; (.^i tii present the ciated with identihcation of specimens, required a\ailable information on the molts and plumages attention. Taxonomic difficulties ha\e |ilagued the of 7(/(7;\'('/(',s: (4) to cimipare the diagnostic char- study of steamer-ducks since their discovery in the acters of the four species and to construct keys to late sixteenth centur\. and certain of the present- the identification of specimens; (."i) to summari/e tla\ problems in s\steniatics of the genus ha\e information on field identification aiul asiculture their origins in this history of controversy. This of steamer-ducks; and lb) to present a subject- state of affairs is in part due to the three most indexed bibliography of the considerable but scat- is that salient characteristics of the genus Tacliycrcs: { 1 ) tered literatLire on steamer-ducks. Our hope large si/e. which exacerbated the difficulties o\ the work will enable ornithologists, some of whom their concerns, to identify collection and study of specimens; ( 2 ) widespread have shared with us flightlessness. an often conspicuous characteristic steamer-ducks in the haiul anil (uinler giiod condi- thai principally created taxonomic conlusitm; and tions) in the field. Should this endeavor prove [?i) drab gray-and-white plumage, which offered successful, we anticipate that these unique birds fev\ clues to the recognition and diagnosis ol will receive the continuing siiidv that they de- species. The taxonomy ofTiichyens also has been serve. ACKNOWLEDGMENTS Our research on steamer-ducks was supported M. Gallardo. R. A. Bockel. and .1. Navas (Museo during 1980-85 by National Science Foundation Argentino de Ciencias Naturales. Argentina). G. grants DEB-8()-1240.\DEB-8 1-1 7942. and BSR- PincheiraandT. G. Iriarte (Universidad de Chile), 83-199{)(). the Kansas University Endowment and M. M. L'Huillier. D. Bersalobec. S. Canessa. Association. Southwestern College, the British C. Montenar, and D. Nunez D. (instituto Profes- Broadcasting Corporation. W. Saul, M. C. Thomp sional de Osorno. Chile). Essential arrangements son. L. A. Osborne. R. T. Peterson. T. Mastm. K. were made by G. A. Giaroli. F. Liibhe and F. Villar. Hamilton, and the Humphrey family. Livezey also Housing and laboratory facilities were arranged was supported in part by NSF grant BSR-85 I6b2.v by O. Kuhnemann, D. Nunez D.. P. Medina, and J. Collecting permits were issued by E. O. Gonzalez Sesti; B. Mayer and F. V. T. .1. Fauring provided a Ruiz (Direccion Nacional de Fauna Sylvestre. home and field support at Puerto Melo. Logistic Argentina). L. O. Saigg de Chiaha (Proteccion help and friendship were provided in the Falkland Ambiental. Chubut. .Argentina). D. H. Soria Islands by A. F. G. Douse (Stanley ) and A. and Y. (Ganaderia. Santa Cruz. Argentina). E. Cruz Davis (Lively Island). We also are grateful for the (Servicio Agricola y Ganadero. Chile), and Sir R. assistance and hospitalitv of B. Rosso. L. Orquera. Hunt (Civil Commissioner. Falkland islands ). Field E. Plana. G. Piacentino, J. E. Bonczak. R. E. work in Argentine national parks was made pos- Caferata. A. Fernandez. R. Landivar. G. C. Sarccda. sible by .\. Tarak (Direccion Nacional de Parques F. Erize. R. Straneck. W. Conway. Y. Lucero. P. NaciiMiales). Logistic support was provided by J. Canevari. M. A. E. Rumboll. R. Hall. H. Niemever. — UNIV. KANSAS MUS. NAT. HIST. MONOGRAPH NO. S M. SallalxTiv A.. A. Veloso. J. Ballastcros C. and Naturales. Buenos Aires) prin ided tield record- the (iihson I'amily. Spcciiiicns were made avail- ings of vocalizations of steamer-ducks, and J. W. able by tlio l-icld Museum of Natural History Hartiy (1-lorida State Museum. Gainesville) pre- (Chieago). British Museum of Natural liistor\ pared sonograms of selected recordings. I'npub- (Tring). American Musemn of Natural llistor\ lishetl observations of steamer-ducks inciirporated (New York). San Diego Museum of Natural His- into the liistribution maps were pro\ ideti by P. tory, Peabody Museum of Natural Flistory (Yale C'anevari. J. C. Chebe/. X. Ferrer. O. Kiilinemann. University). Museum of Comparative Zoology T. Narosky. M. Nores. and R. Straneek. We also (Harvard I'niversity ). Museum of Zoology (Llni- thank R. Mengei. M. Jenkinson. and their col- versity of Michigan), and U. S. National Museum leagues ft)r preparation of speciinens; P. Rasmussen of Natural History. R. Straneek (Laboratorio de for executing Figure '->: and K. McManness, K. Sonidos Naturales. Museo .Argentino de Ciencias C'orbin. and M. Schmal/ for t\ ping. METHODOLOCilCAL N()TP:S Specimens Measurements Much of our work on steamer-ducks has been Masses were measured w ithin 1 2 hours of col- based on observations made and specimens col- lection of the birds using spring scales. Most skin lected during our Held v\(nk. We made five trips measurements used are standard (Baldwin ct al. during 1979-84 (the first with Max C.Thompson), I9.il). although a few comments are needed.
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