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The Use of the Wings and Feet by Diving Birds

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The Use of the Wings and Feet by Diving Birds 234 TOWNSE.WD,Useof Wingsand Feet by DivingBirds. [.July['Auk THE USE OF THE WINGS AND FEET BY DIVING BIRI)S2 BY CHARLES W. TOWNSEND• M.D. BIRDSthat diveand swimunder water may be dividedinto two main classes,--thosethat habituallyuse the wingsalone for sub- aqueouspropulsion, and thosethat use the feet alone. 'The followingpaper includesmy own observationswhich have been madeonly on wildbirds and have ' been recorded at thetime, and of recordsgiven •ne by other observers,as well as of thosecollected from literature. Thcse two last named include observations on captivebirds in tanks,which I believeare of value,for, as we shall see,the birdsthat habituallyuse their wingsalonc under water whenin the wild state,and those that habituallyuse the feetalone, follow this rule even when confined. In watchingwild birds divingI have concludedthat thosethat spreadtheir wingsjust as they enterthe water,use them under water,while those that keepthe wingstightly pressed to the sides, and often executegraceful curves in diving, sometimesleaping clear of the water, dependon the feet alone. The tr•tth of this conclusionI have confirmed in someinstances by obscrvingthe bird under water; in other instancesthc conclusionhas been borne out by the obscrvationof others,so that the rule is, I think,a goodone. Sinceformulating this rule I havefound that EdmundSelous (15) eq•ressesthis same idea when he says:"This openingof thewings in the momentof divingis, I believe,a suresign that theyare used as fins or flippersunder water." And again (16): "On the other hand cormorants,shags, and mergansers,birds which do not use their wingsin this way, dive in a quitedifferent manner. Instead of the sudden,little, splashyduck, as described,they make a smoothgliding leap forwards and upwards,rising a little from the water,with the neckstretched out, and wingspressed closc to the sidesto enter it again back foremost,like a curvedarrow, thus describingthe segmentof a circle." In thc Loonsand Grebes,the wingsare small,but the legsare large and powerful. The femur is short and stout,thoroughly Read before the Nuttail Ornithological Club, February 15, 1909. Vol.1909XXVI] ] TO•VNS•ND, Use o[ Wingsand Feet by Diving Birds. 235 imbeddedin musculartissue, while the tibia is longand provided with a crestor keel on the anteriorsurface of the proximalend, to which the powerfulmuscles that movethe tarsusand toesare attached. In the Grebesthere is a very largepatella. In these respectsthis groupresembles the fossilHesperornis, a toothedbird with wingsrepresented by mere vestiges,but onethat wasstrongly specializedfor propulsionthrough the water by meansof the feet alone. The Loon and the Red-throatedLoon in divingkeep the wings closelypressed to the sides,and disappearbelow the surfacewithout a splash,propelled apparently by the leg actionalone. I have not beenable to seethese shy birds under water, but the inference that they do not usetheir wingsthere is confirmedby Mr. C. H. Townsend(25) who saysin regardto a tame Loon observedin the NexvYork Aquarium: "In exploringthe bottomof the pool, or in pursuitof Killifishesit swamunder water with its wi•gs closely folded- neverin use." The italicsare his. The pool in which the Loon was confinedwas of ample size to displaythe agility of both bird and fish; it was twenty-eightfeet long and three feet deep. Evans (3) statespositively both of Loonsand Grebesthat "whensubmerged they do not usethe pinions." C. LloydMorgan (27) says: "It mustbe rememberedthat Grebesand diversdo not use the wingsfor progressionunder water." Lea (8) saysof the Loon: "It usesits largewebbed feet only,when swimmiug[under water]." Althoughprogression by the fee[ aloneseems to be the rule in this group,yet it is a fact that at timesthe wingsalso are used. Thus Mr. C. Wm. Beebe,whose opportunities at the New York Zo61ogicalPark areunusually great, writes me under date of Nov.6, 1908, that "Grebes and Loons do [use their wings] at times of emergencyto turn quickly,or getup a burstof speed." Mr. W•n. Brewster tells me, and the observation has bceu con- firreedby others,that Loonsin divingunder boatsto avoidbeing hemmedin, or in hurryingfrom their nests,use their wingsas well as their feet, and this I think has givenrise to the idea that 'they alwaysuse their wings. I havealso been told by thosewho have observedyoung Loons, that theyuse both wings and feet in (living. A noteof th{ssort was given me by Mr. F. H. Allen, who wasim- pressedby the "quadropedal action" of theyoung bird under water. 236 TOWNSEND,Use o/ Wingsand Feet by DivingBirds. 1July['Auk I have watchedHorned, Holboell'sand Least Grebesdiving, and as in the caseof the Loonsthe wingsare alwayskept closely pressedto theside. Whilethe Grebesare able to sinkmysteriously in the water, or to dive with very little motionof the body,they oftenleap clearof the water and executegraceful curves or arcsas theydescend into the water. At suchtimes the feetare stre•rched out behind. In the shallowwater of a pondat closerange I have seenthe powerfulleg strokesof the Least Grebeas it progressed underwater, while the wingswere seen to be 1notionlessat the sides. On severaloccasions at IpswichI havebeen treated to an excellent view of Horned Grebesas they dove,borne up in the clear trans- parentwater of an advaucingwave before it brokeon the beach. In this casethe wingswere not in use,they werepressed close to the sktes,while the Grebesadvanced rapidly by leg strokesalone. JosephKittredge, Jr. (7) in describingthe actionsof a Horned Grebe in a small tidal pool says: "He doveimmediately .... In this casecertainly, he did not usehis wings in swimming." Evans, as alreadyquoted, states that Grebesdo not usetheir wingsunder water, and in anotherplace (4) he says: "in divingthe feet alone act as oars." Mr. A. C. Bent in replyto a (p•estionon this subject kindly wrote me as follows: "While collectingin the big Grebe rookeriesin SaskatchewanI had ample opportunitiesto observe the WesternGrebes swimming under water at shortrange, often almostat my feet; I am quitesure that they generallyswam with the wingsclosed, and swamvery rapidly. I am quitesure that I have seenthe Loonsdo the samething, thoughI have no doubt that they bothuse their wingsoccasionally." Very different is the case of the Aleidm, the Auks, Puffins and Guillemots, birds with stout rounded bodies and short necks. In all the membersof this family that I haveseen, the wingsof the divingbirds are not pressedto the sidesbut spreadout for action as they go belo,•vthe surface,and I have actuallyseen the wings used under water in some instances. The Puffin on the Labrador coastallows of closeapproach, and onecan plainlysee the wings usedvigorously as the bird descendsunder water. That the wings aloneare usedis statedby Selous(17) whosays: "I havebeen able to follow the Puffin downwardsin its dive, and at once noticed that the legs,instead of beingused, were trailed behind, as in flight, Vol.1909XXVI] J TOWnSEa-U,Use o[ Wingsand Feet by Diving Birds. 237 so that the bird'smotion was a gemfineflight throughwater, un- assistedby the webbedfeet." The Black Guillemotor Sea Pigeonplainly usesits wingsunder water, for it goesflown with a very ol)viousflop, spreadingboth wings.as if to pushthe water away. Selous(22) says: "In swim- ming under water the Black Guillemotuses its wingsonly--the rosered legstrail behindit, a fadingfire as it goesdown." Mr. James L. Peters writes me that he "watched a Black Guillemot off Nahant. We werealmost directly above him and, while we could seehis wingsnicely, his feet were quite hidden. When he dove he spreadhis wingsand usedthem under water." The Razor-billedAuk and the Murres spreadtheir wingsin diving, but I have not seenthese birds under water. Selous(24) saysof thesebirds: "Whilst watchi•g the Guillelnots[Common 5/furres]on the ledges,one of them flew down into the sea, just below,which was like a great, clearbasin, and tht•sgave me the first opportunityI have yet had of seeinga Guillemotunder water. It progressed,like the Razorbilland Puffin,by repeatedstrokes of its wings,which were not, however,ot•tspread as in flight,but held as they are when closed,para]leI, that is to say, roughlyspeaking, with the sides,from,which they were movedontwards, and then back,witch a flap-likemotion, as thoughattached to themall along. Thus the flight throughthe water is managedin a very different way from the flightthrough the air." In anotherplace (20) he says: "Razorbills also dive briskly,opening the wings.... One remarksthen that the wingsare movedboth together--flapped or beaten--so that the bird really flies throughthe water. In flight,however, they are spread straight out without a bendin them, whereashere they are all the while flexedat the joint, wing raised from a•rdbrought downwards again towards the sidesin the same positionin whichthey repose against them when closed." Of the Dovekieor Little Auk I have had the opportunityto make some interestingobservations. In diving the wings are spreadout as in othersof the group,and, from a near-byrock, I haveplainly seen it usingits wingsas it swamunder water. As to the feetI cannotsay, for I haveno notenor distinctremembrance as regardsthese members. One is apt to assumethat the feetas wellas the wing's are used in thesebirds under water, but the careful 238 TOWNSraND,Use o/ Wingsand Feetby DivingBirds. [July[Auk observationsof Selouson other membersof the groupshow that the wings only are used. The GreatAuk, with wingsreduced to flipper-likeproportions, doubtlessadvanced rapidly throughthe water
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