April 1982] ShortCommunications 365

chology and education, 2nd ed. New York, PAYNE,R. B. 1977. The ecologyof brood parasitism McGraw-Hill Book Co. in birds. Ann. Rev. Ecol. Syst. 8: 1-28. HERGENRADER,G. L. 1962. The incidence of nest- PIeMAN, J. 1977. Destruction of eggs by the Long- parasitism by the Brown-headed Cowbird on billed Marsh Wren (Telmatodytespalustris palus- roadside nesting birds in Nebraska. Auk 79: tris). Can J. Zool. 55: 1914-1920. 85-88. ROBERTSON,R. J., & R. F. NORMAN. 1977. The func- HILL, R. A. 1976. Host-parasiterelationships of the tion and evolution of aggressivehost behavior Brown-headed Cowbird in a prairie habitat of towards the Brown-headed Cowbird (Molothrus west-central Kansas. Wilson Bull. 88: 555-565. ater). Can. J. Zool. 55: 508-518. MAYFIELD,H. F. 1961. Vestiges of a proprietary in- ROT•STEIN,S. I. 1975. An experimentaland taleon- terest in nests by the Brown-headedCowbird omic investigation of avian brood parasitism. parasitizing the Kirtland's Warbler. Auk 78: Condor 77: 250-271. 162-166. Received13 July 1981, accepted19 October1981.

Evidenceof a BorealAvifauna in Middle Tennesseeduring the Late

PAUL W. PARMALEE AND WALTER E. KLIPPEL Departmentof Anthropology,University of Tennessee,Knoxville, Tennessee 37916 USA A faunistically significant, stratified deposit environment for an extended period of time in this was discoveredduring an archaeologicalsite survey area during the Late Wisconsinan (Parmalee and of the proposedTennessee Valley Authority (TVA) Klippel 1981). Columbia Dam that will impound approx- The avifauna from Cheek Bend Cave is extremely imately 30 km of the Duck River in middle Tennes- diversified and reflects both open grasslandor sa- see. This small cave, known as Cheek Bend Cave vanna and forest habitat. Remains of aquatic birds (40MU261), is in a limestone bluff 20 m above the (e.g. Pied-billed Grebe, Podilymbuspodiceps, and at river bed, 13 km east-southeastof Columbia, Maury least three speciesof ducks) are rare in the deposit, County. Testing for the presenceof aboriginal oc- although severalspecies associated with riverine or cupation began in September 1978 and, as a result marsh habitats are represented.Most notable among of encounteringhuge quantitiesof animal bone and theseare the Yellow Rail (Coturnicopsnoveboracensis) shell at considerabledepths that appearednot to be and the BlackRail (Laterallusjamaicensis), both prob- attributable to man, was continued until May 1979. ably rare transients in Tennessee,although their Three 1 x 2-m excavationunits placedalong the east modern distribution in the Midsouth is poorly wall revealed stratified deposits of animal remains known. We identified a minimum of approximately to a depth of about 4.5 m. 60 species,representing 28 families, from the avian On the bases of the speciescomposition within sample. Elementsof numerous passerinespecies of and among strata, differing fill zones, and certain the families Parulidae and, particularly, Fringillidae otherstratigraphic features, the depositreflecth two comprised 75-95% of the avian samples from the distinct and -defined major episodesof fill. The levels. Except for the PassengerPigeon top 2 m represent the Holocene epoch and contain (Ectopistesmigratorius) and the Ruffed Grouse (Bo- remains of only extant modem species.In contrast, nasa umbellus),all of the identified species repre- the faunal assemblagein the bottom 2 m, although sented in the Holocene levels are still a part of the it includes some speciesthat still inhabit the cave avifauna of middle Tennessee.A comprehensivere- area (presently a deciduous Western Mesophytic port dealingwith the stratigraphyand chronologyof Forest),contains many that are now extirpatedand CheekBend Cave, its archaeologicalcomponent, and reflect a prairie habitat (Prairie Chicken, Tympanu- the total faunal assemblageis being preparedfor the chuscupido; 13-lined ground squirrel, Spermophilus TennesseeValley Authority. tridecemlineatus,pocket gopher, Geomys sp.) or a bo- Several speciesof birds identified from elements real environment (e.g. arctic shrew, Sorexarcticus; recovered in the bottom 2 m of the deposit are es- water shrew, Sorexpalustris; red-backed vole, Cleth- pecially significant,because they not only represent rionomysgapperi ; yellow-cheekedvole, Microtusxan- new stateor regionalrecords but alsobecause they thognathus; red squirrel, Tamiasciurushudsonicus ; suggesta boreal and habitat in this region northern flying squirrel, Glaucomyssabrinus; and during the Wisconsinan.Five speciesworthy of spe- heather vole, Phenacomysintermedius). Recovery of cial comment are the Hawk Owl (Surnia ulula), Boreal a partial carapaceof a wood turtle (Clemmysinsculpta) Owl (Aegoliusfunereus), Saw-whet owl (Aegolius also suggests,based on the presentcenter of distri- acadicus),Gray Jay (Perisoreuscanadensis), and bution of the wood turtle, a well-established boreal Grosbeak (Pinicola enucleator). In addition to these 366 Short Communications [Auk, Vol. 99

TABLE1. Avian speciesidentified from the late Pleis- Siskin (Carduelispinus) also suggestextensive conif- tocenestrata of Cheek Bend Cave, Maury County, erous forests in the vicinity of Cheek Bend Cave. Tennessee. Althoughmost species comprising the avian assem- blage reflecta forest habitat, the presenceof mead- Falconidae owlark (Sturnellasp.), Horned Lark (Eremophilaal- AmericanKestrel, Falco sparverius pestris), Prairie Chicken, Sharp-tailed Grouse Tetra onida e (Pedioecetesphasianellus), and somerodents such as Ruffed Grouse, Bonasa umbellus the pocketgopher (Geomyssp.) is indicative of these Sharp-tailedGrouse, Pedioecetes phasianellus a forestedareas having been interspersedwith short- Greater Prairie Chicken, Tympanuchuscupido grassprairie or savanna. The Hawk Owl lives in Canada and Alaska in co- Rallidae niferous forests mixed with scrub, tamarack Yellow Rail, Coturnicopsnoveboracensis , and muskegs(Bent 1938). This owl is a rare Scolopacidae and sporadicwinter vagrant in northern Plains and CommonSnipe, Capellagallinago • Midwest states; there are no historic records of it American Woodcock, Philohela minor from Tennessee and no Pleistocene records from Columbidae anywhere. Identification of Surniaulula from Cheek PassengerPigeon, Ectopistesmigratorius Bend Cave is basedon a completephalanx (phalanx I, digit I) and premaxilla(Fig. 1). Distribution of the Strigidae Boreal Owl is similar to that of the Hawk Owl, as is Barred Owl, Strix varia its choiceof habitat. The only fossil(Holocene?) rec- Boreal Owl, Aegoliusfunereus a ord of A. funereusis from ShelterCave, New Mexico Saw-whetOwl, Aegoliusacadicus • Hawk Owl, Surnia ulula a (Brodkorb 1971). Evidence of its occurrence in his- toric times in Tennesseeis lacking. Diagnostic ele- Apodidae ments of at least two individuals of the Boreal Owl Chimney Swift, Chaeturapelagica • from Cheek Bend Cave consistof the proximal two- Picidae thirds of a right and shaft sectionof a left tarsometa- CommonFlicker, Colaptesauratus a tarsus, distal one-third of a right and distal end of Red-belliedWoodpecker, Melanerpes carolinus a left tibiotarsus,distal end of a right femur, head Hairy Woodpecker,Picoides villosus of a left humerus, distal end of a left ulna, proximal Tyrannidae end of a left radius, sectionsof a lower jaw, articular EasternPhoebe, Sayornis phoebe end of a left scapula,coricoidal articular facet and manubrium sectionof a sternum,and 18 phalanges. Alaudidae Although the Saw-whet Owl is not restrictedto Horned Lark, Eremophilaalpestris • the northern coniferous forests, its center of distri- Corvidae bution and greatestabundance lies in the woodland Blue Jay, Cyanocittacristata areas acrosssouthern Canada, the Great Lakes states, Gray Jay, Perisoreuscanadensis • and in the northeastern United States. It is a rare Sittidae winter residentin the Midsouth;there are only four reported sight recordsof this owl in the Nashville Red-breasted Nuthatch, Sitta canadensisa area (ca. 60-80 km north of Cheek Bend Cave) (Rig- Turdidae gins 1977).All remainsof A. acadicuswere recovered Robin, Turdusmigratorius from the same late Wisconsinan levels (300-400 cm Icteridae below the surface) in which the bones of the Hawk Owl and Boreal Owl occurred. Remains of the Meadowlark, Sturnellasp. Screech Owl (Otus asio), a common inhabitant of Fringillidae primarily deciduousforests of easternNorth Amer- PurpleFinch, Carpodacuspurpureus ica, were found in the Holocene levels of Cheek Bend Pine Grosbeak, Pinicola enucleator• Cave but not in the Pleistocene levels. Elements of Pine Siskin, Carduelispinus a the Saw-whet Owl consist of the distal section of a cf. Towhee, Pipilo erythrophthalmus cf. White-throatedSparrow, Zonotrichia albicollis right and left coracoid,articular end of a right scap- ula, proximalone-half of a right and distal one-third Remainspresent only in late Pleistocenestrata. of a left radius, distal end of a left ulna, a complete right tibiotarsus, distal one-third of a right tarso- metatarsus,and sevenphalanges. In easternNorth America both the Gray Jay and five, remainsof 23 other specieswere identifiedfrom Pine Grosbeakoccur throughout the coniferousfor- the strata (Table 1). In addition to ests of southern Canada, and in the United States the three owls, Gray Jay, and Pine Grosbeak,the they seldom appear farther south than the upper Red-breasted Nuthatch (Sitta canadensis)and Pine Great Lakes and northern New England states. Like April1982] Shor•Communications 367

a c j

a

b b d e

A B

a

e g b c d f

c

Fig. 1. Avianfossils from Cheek Bend Cave, Maury County, Tennessee. A. Surniaulula: a, premaxilla; b, phalange.B. Aegoliusacadicus: a, tibiotarsus; b, radii; c, coracoids; d, ulna; e, tarsometatarsus.C. Aegolius funereus:a, sternum;b, humerus;c, radius;d, ulna;e, femur;f, tarsometatarsi;g, tibiotarsi.

the Hawk Owl and Boreal Owl, these two passerines teau, basedon the analysesof pollenrecovered from are associated with a boreal habitat and seldom wan- sedimentcores from Anderson Pond (White County) der or migrategreat distances. Identification of Pin- and Mingo Pond(Franklin County), Delcourt (1979) icola enucleator was based on sections of the lower was able to show that several major vegetational jaw andleft carpometacarpus (distinct from Pheuticus changeshave taken placeduring the past 25,000 and Hesperiphona)and that of Perisoreuscanadensis . During the Wisconsinanglacial maximum, on limb elementsincluding the distal ends of two about 19,000-16,300yr BP, borealtaxa of jack pine, left humeri and two complete left carpometacarpi. spruce,and fir weredominant, and "Between16500 Fossilrecords of thesepassetines are lacking(Brod- and12500 yr BP,mixed coniferous deciduous forests korb 1978). coveredthe landscapebetween 34øN and 37øNlati- In a recentstudy of the late Pleistocenevegetation tude" (Delcourt1979: 276). In his discussionof the of the easternHighland Rim and CumberlandPla- vegetationof easternNorth America18,000 yr ago, 368 Short Communications [Auk, Vol. 99

Wright (1981:122) statesthat "Borealforest formed CheekBend Cave (40MU261)and the analysisof the a belt perhaps1000 km broad, starting at or near the faunal materialswere made possible. [Laurentide] ice front in the north and ending south near a line extendingfrom central Georgia around LITERATURE CITED the southernend of the AppalachianHighlands to BEtat, A. C. 1938. Life histories of North American Tennessee.... "Recovery of remains of predomi- birds of prey (part 2). U.S. Natl. Mus. Bull. No. nately boreal small mammals from the lower levels 170. of Cheek Bend Cave and those of four avian species BROVICORB,P. 1971. Catalogueof fossil birds: part whose present primary northern ranges coincide, 4 (Columbiformes through Piciformes). Bull. coupledwith pollen analysesdata, suggestthat the Florida State Mus. 15: 163-266. Nashville Basin was characterizedby climatic con- ditions that included somewhat lower annual mean ß 1978. Catalogueof fossilbirds: part 5 (Pas- seriformes).Bull. Florida State Mus. 23: 139-228. temperaturesthan exist today and a climax vegeta- DErCOURt,H. R. 1979. Late vegetation tion dominated by coniferousforest. Although the history of the eastern Highland Rim and adja- possibility exists that the former presenceof the cent Cumberland Plateau of Tennessee. Ecol. Hawk Owl, BorealOwl, Saw-whetOwl, Gray Jay, and Pine Grosbeak at this cave site was the result of Monogr. 49: 255-280. PARMALEE, P. W., & W. E. KLtPPEL. 1981. Remains southward-wanderingvagrants, the small mammal of the wood turtle Clemmysinsculpta (Le Conte) assemblageand the remains of at least two individ- from a Pleistocenedeposit in Middle Tennessee. uals of A. funereusand P. canadensissuggest a locally Amer. Midi. Natur. 105: 413-416. establishedpopulation of these boreal birds during RIGGINS,J. N. 1977. Fourth Nashville area record of the Wisconsinan. Saw-whet Owl. Migrant 48: 12-13. We wish to thank W. M. Wright for the photo- WRtGHT, H. E., JR. 1981. Vegetation east of the graphsand B. W. Creechfor typing the manuscript. Rocky Mountains 18,000 years ago. Quaternary We gratefullyacknowledge the TennesseeValley Au- Research 15: 113-125. thority under whose auspices(Contracts No. TVA TV-49244A and TVA TV-53013A) the excavation of Received27 May 1981, accepted20 October1981.

Flightlessnessin Flying Steamer-Ducks

PHILIP S. HUMPHREY AND BRADLEy C. LIVEZEY Museumof NaturalHistory and Department of Systematicsand Ecology, University of Kansas, Lawrence, Kansas 66045 USA Steamer-duckscomprise a single, widely distrib- that Flying Steamer-Ducks"are capable of getting uted, flying species,Tachyeres patachonicus (Falk- well into the air within a spaceof 2 meters, even in lands and Fuego-Patagonia),and three flightless a dead calm," and "they rarelydived at the approach species, T. brachypterus(Falklands), T. pteneres of... [Beck's]craft, but preferred to take wing .... (Tierra del Fuego and the southerncoast of Chile), some of them flew heavily, not rising far above the and the recentlydiscovered (Humphrey and Thomp- water, while others mounted high into the air and son 1981) T. leucocephalus(coast of Chubut, Argen- passedquite out of sight." Reynolds(in Lowe 1934: tina). This paper presentsdata on wing loading and 474) wrote that "Tachyerespatachonicus is capableof useof flight for escapeby FlyingSteamer-Ducks and sustainedand rapid if somewhatheavy flight.... evidencethat someindividuals are permanentlyin- At times T. patachonicusmay well be taken for a capableof flight. flightlessbird, and some of the confusiondistin- Flying Steamer-Ducksoccur in freshwater and guishingbetween the two species[patachonicus and marine coastalenvironments in Patagonia,Tierra del pteneres]is probably due to the unaccountablere- Fuego, and the Falkland Islands (Weller 1976, Hum- luctance to rise so often demonstrated by this phrey and Thompson1981). They feed on sessileor species." slow-moving molluscs,crustaceans, and insect lar- During field seasonsin 1979 (Humphrey and Max vae, which they often obtain by diving, using hind C. Thompson)and 1980-1981(Humphrey and Liv- feet and partly spreadwings for propulsion(Murphy ezey), 57 specimensof Flying Steamer-Duckswere 1936:960; Humphrey and Livezey unpubl. data). collectednear Ushuaia, Tierra del Fuego (5-17 No- Weller (1976: 47) noted the "hesitancy of ... vember 1979, 24-27 December 1980, 4-6 January Flying SteamerDucks to fly .... "Murphy (1936: 1981), Puerto Deseado, Province of Santa Cruz (10-16 970), writing of Beck's collecting activities, stated October 1979, 20-26 January1981), and Puerto Melo,