234 TOWNSE.WD,Useof Wingsand Feet by DivingBirds. [.July['

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 otherobservers, 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 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 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 ,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 ,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 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 ; 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 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 feet are 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 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 , 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 by the action of thesemembers only. The ,akhough entirely distinct froin the Auks, fly throughthe water with their attenuatedflipper-like wings, and, accordingto Beebe(1) whohas watched them in tanks,"make but little use of their feet in swimming,only occasionallyaiding the tail in steering." Evans (5) says: "When submerged,the wings act as paddleswith alternatingrotary action, and the feet as rudders,but on the returnto the surfacethe latter naturallybecome propellers." Lea (9) saysof Penguins:"Their flight maybe watchedand studiedin the largeglass tanks at the 'Zoo'. .... With short,rapid strokes of its paddle-wingsit dartsthrough the water leavinga trail of glisteningbubbles behind, and shootsforward with the speedof a fish, turning more rapidly than almostany bird of the air by the strokesof the wingsalone, the legsfloating apparentlyinert in a line with the gleamingbody, or givingan occasionalupward kick to forceit to greaterdepths." Someof the Terns,in plungingfor fish,disappear entirely under water,and as theirfeet are comparativelyfeeble, and astheir wings are partiallyopen in the plunge,it is possiblethat theselatter are usedto someextent in aidingtheir progress, although it is probable that the impulseof the plungealone is all that is necessary. Amongthe Tubinares,some of the Petrelsand the Shearwaters plungeunder water. Darwin (2) saysof a diving Petrel, Pele- canoidesberardi: "When disturbed it divesto a distance,and on comingto the surface,with the samemovement takes flight. After flyingby the rapid movementsof its shortwings for a spacein a straightline, it drops,as if struckdead, and divesagain .... It would undoubtedlybe mistaken for an auk, ,when seen from a distance,either on the wing,or whendiving and quietlyswimming aboutthe retiredchannels of Tierra del Fuego." Lea (10) states of the Capped Petrel, (E.•trelatahcesitata, now nearly extinct: "It poisesitself in the air for a momentat a heightof twentyor twenty-fivefeet, and then, folding its wings,takes a headerinto the water. The actualplunge is madewith the wingsopen, and Vol.1909XXVI'] J Tow•sr•D, Useo[ Wingsand Feet by Diving Birds. 239 they are used under water much in the samemanner as during flight." The plungeof the Gannet,with wingspartly open, is so swift and oftenfrom sogreat a height,that it seemsprobable that there is no needof eitherwings or feet underwater, but that the initial impulseof the plunge,which must be greaterthan that of gravity alone,is sufficientto enablethe bird to catchits prey. Cormorantsexecute graceful curves in divingwith wingsclose pressedto the sides,often throwing themselves clear of the water with their powerfulfeet. I have never seen them under water but haveconclusive evidence from literatureon this point. F.W. Headley(6) says: "The Cormorantuses his feet aloneto propel him [in diving]striking with bothsimultaneously, and holdinghis wings motionless,though slightly lifted from the body. The positionof thewings must have given rise to the idea,common amongfishermen, that the Cormorantflies under water.... But whenyou see him in a tank youcan have no doubtthat the legsare the propellers."Mr. Beebe,in a letter,confirms this observation from his experiencewith Cormorantsin tanks. Selous(23), speakingof Shags,both adultsand full grownyoung, observed in cavernsin the ShetlandIslands says: "Others, whose young were still with themon the nest,although full-fledged and ahnostas big asthemselves, plunged, attended by theseinto the water ..... It was easyto followthese birds as theyswam midway between the sur- faceof thewater and the white pebbled floor of the cavern,and I wasthus able to confirmmy previousconviction that the feet alone are usedby them in swimming,without any helpfrom the wings, whichare keptall the whileclosed. I havemany times observed this before,but neverso clearlyor for sucha lengthof time." Lea (11) saysof Shags:"This speciesalso may be watchedat the 'Zoo'. It alwaysbegins its diveby jumpingup in the waterand taking a header,and then strikeshard upwardswith both feet. You will seethat it doesnot useits wingsat all for swimming,but holdsthem quitestill, lifted just a little away frmn its body. It strikesout with both feet simultaneously,and in this differsfrom the Darters(Plot•.•), which adopt an alternatestroke, as you may see for yourselfby visitingthe Diving-birds'House at feeding- time." In thisconnection it is interestingto notethat Cormorants 240 Tow•se_•u,Use o/ Wingsand Feet by DivingBirds. L['AukJuly in risinginto the air in flightkick awaythe beachor the waterwith bothfeet together. This I haveproved by examiningthe imprints of their feet in the sand of a beach. The Dartersor Anhingas,as just referredto by Lea, usethe feet alone under water. Among the Ducks, both classesof divers are found. The American and the Red-breasted]5•erganser both dive like the . They oftenleap clear of the water, in gracefulcurves, with their wingscleaving closely to the sides. At othertimes the leap is much curtailed,or they sink beneaththe surfacewithout apparenteffort. I shouldinfer, therefore, that the wingswere not usedunder water, and this inferenceis borneout by the following from Selous(16): "The merganserdives like the shagor cormor- ant -- thoughthe curvedleap is a little less.vigorous -- and swims, like them, withoutusing the wings. His food beingfish,....he usuallyswims horizontally, sometimes only just beneaththe surface, and, as he comesright into the shallow inlets, when the water ahnostlaps the shore,he canoften be watchedthus gliding in rapid pursuit." The other membersof this order that do not, I believe,use the wings under water are the Redhead, Greater and Lesser and Ring-neckedSeaups, Whistler and Bufflehead,while those that do usethe wingsare the Old Squaw,Harlequin Duck, the threeEiders the threeScoters and possiblythe Ruddy Duck. This list I have madeout frownmy own observationsof the way the wingsare held in thesebirds as they dive, and in a few casesthe birdshave been observedunder water. The only memberof our easternsea or divingducks omitted frotn the list is the Canvasback,which I have neverseen dive. It is probablethat it acts as doesthe nearly relatedRedhead. It is interestingto noticethat the two classesare groupedseparately in the A. O. U. Check-List,the first classbeing placedtogether at the beginningof the list, the other classat the end of the list. Someof theseDucks feed largely on the bottomon shellfishand crustaceans,or on vegetablematter, while a few of them feed largelyon fish. The latter birdswould naturallydevelop the swiftestform of propulsion. Mergansers,Whistlers and Burfie- heads,largely fish-eating Ducks, progress by the feet alone,while Vol.1909xxvIq TownsEnD,Use o/ Wingsand Feet by Diving Birds. 241

Eidersand Scoters,almost entirely bottom feeders, use the wings. Many of the others,however, cannot be dividedin thisway. Redheadsin divingkeep their wingsclose to their sidesand Mr. Beebewrites that thesebirds, as observedby him in tanks,do not usethe wingsunder water. The Seaups,both Greaterand Lesser, in the sameway keep the wingsclose to the sidesin diving,and sometilnesleap clear of the water or disappearwith scarcelyan effort. Mr. (7. M. Allen tellsme that he watcheda captiveRing- neck Scaupin a small pool, and was able to observenot only the cleancut leap and (live with wingsclose to the side,but the rapid progressunder water by the useof the feet alone. The Whistler or AmericanGolden-eye, is alsoa gracefuldiver, and, as far as I have observed,always keeps its wingsclose to its sidesin (living. As it disappearsfrom sightit oftensends up a little spurt of water by the powerfulaction of its feet. I shouldtherefore conclude that the Whistlerhabitually swims under water by the use of its feet alone. Mr. Win. Brewster, however, tells me that he has seen it make use of its wingsin (living, but this was probablyat times whenthe bird washard pressed,and it actedas doesthe Loon in similarcircumstances. The Buffieheadalso keeps its wingsclose to its sidesin diving,and sometimesleaps clear of the waterbefore it disappearsfrom sight. Oht Squaws,on the otherhand, opentheir wingsbefore diving as plainlyas do the Guillemotsand the Puffins. On oneoccasion, while I was watchingsome Old Squawssporting in the water off Nahant, chasingeach other on and just belowthe surface,I dis- tinctlysaw the wingof oneof themcut the waterfrom belowlike the fin of a great fish. A Harlequin Duck, that I saw on the Labrador Coast,opened its wingsas it clove. I have watchedthe Northern Eider, the AmericanEider and the King Eider dive, and all opentheir wingsfor subaqueousflight as they go down. Of the Eider Selous(15) has madesome satisfactory observations. He says: "Their dive is a suddendip down,and in the act of it theyopen the wings,which they use under water, as canbe plainly seenfor a little way belowthe surface." Our threeScore(s, --American, White-wingedand Surf Scoters --all open their wingsas they dive. I onceshot and slightly woundeda Surf Scorerthat wasstanding on the edgeof the beach 242 Tow•'sE•D,Use o[ Winejsand Feetby Divin(jBirds. [.July[Auk

at Ipswich. He tookto the shallowwater and dovewhere I could plainlysee him flyingalong under water using his wings. My observationsof the Ruddy Duck lead me to think that althoughits wingsare often closeto the sidesat the beginningof the dive,they are openedjust as the bird goesunder the surface. The shortbut strongwing in this bird wouldsuggest adaptation for subaqueousflight. The curiousSteamer Duck, Tachyerescinereus, of the Straitsof Magellan,that in the adult stateat leastis unableto fly, but flops along the water by the use of its wings,and divesawkwardly, probablyuses its wingsunder water. Darwin (2) saysthat he is "nearlysure" that thisbird usesthe wingsalternately. • The group of River Ducks obtain their food by dippingtheir headsand necksbelow the surfaceof the water, but occasionally these birds dive. I have observed close at hand smni-domesticated decoyBlack Ducks sportingtogether, and diving awkwardly. In this caseboth wingsand feet wereused. As regardsthe Mal- lard, the followingnote kindly communicatedto me by Mr. Win. L. Finleyin a letterdated January 4, 1909,is of considerableinterest. "While in the lake regionof southernOregon, on two different occasionsI saw a youngMallard duck swimmingunder water. He lookedto me exactlylike a frog. He wasnot manydays old. He usedhis little wingsas if theywere two front feet, and he we•t throughthe water like a streak." It is evidentfrom the context that the feet were used as well as the wings. Teal occasionally dive, especiallywhen wounded,but I have no observationsto record as to the method used." The Rail family showan interestingdiversity in the mannerof diving. The AmericanCoot, Fulica, with its largelobed feet has evidentlyperfected the feet method,for with its feeblewings close to its sides,it oftenleaps out of the waterand describesan arc, the bill enteringthe water as the feet leaveit. At other timesit dis- appearswithout any leap, and all degreesbetween these two extremes are to be found. Mr. Beebe writes me that Coots in

• The alternate action of the wings has already been referred to in the case of Penguins. I 11aYefor 8Ollle years been aliilost convinced that Chimney Swifts use the wings alternately in flight. 2 I recently watched a wounded Brant dive and swim under water. In this case the wings were flapped slowly and the feet used rapidly. Vol.1909 XXVI'] j TOWNSr•D,Use o/ Wingsand Feet by Diving Birds. 243

tanksuse the feet and not the wingsunder water, and Selous(18) believesthat the Cootbelongs "to the cormorant-schoolof diving." This he infers from the manner of its enteringthe water. He apparentlyhas not seenit underwater. I haveno observationsof my ownto recordon Railsproper and Gallinulesbut Selous(21) saysof the EnglishMoorhen, Gallinula chloropus,that he "may followno fixedplan in his diving,for I have certainlyseen him using his feet only under water, and I believeI have also seenhim usinghis wings." Lea (12) saysof the Moorhen: "After diving, it flies throughthe water at a great pace." Amongthe Shore-birdsthe ymmgof the SpottedSandpiper are saidto useboth wings and feet in diving. The impetusof the Kingfisheris probablyall that is neededby titisbird in itsplunge, but it ispossible that it occasionallyscra•nbles a bit with its wingsunder water,-- its feet canhardly be of use. AmongPasserinc birds the waterOusel is the onlydiver, and it is well known that this bird usesits wingsunder water, and many observersstate that it usesalso its feet (13). From the result of these studies it seems reasonable to conclude that diving birds tend to specializein two directions,--either towardsthe useof the feet alone,or of the wingsalone. The ques- tion naturallyarises as to which line is superior,which has pro- ducedthe swiftestdiving bird,-- the linethat haslead to theuse of the feet aloneor that whichhas lead to the useof the wingsalone ? It is evidentthat a methodof divingwhich leaves the wingsunim- pairedin sizeor formfor theuse in theair isa desirableone, a•d this is possiblewhere the feetalone are used. In mostfishes pro- pulsionis fromthe rear by meansof the tail, for the pectoralfins, which correspondto the birds'wings, are usedchiefly for balanc- ing. When the fish swimsfast thesefins are kept closeto the sides. Amongmammals the cetaceanshave developedgreatest speedin divingand swimming under water, and here also the tail is the propulsivepower, while the anterior extremitiesare used chieflyfor balancing. The modernscrew propeller is superiorto the old side-wheeler. In Hesperornisthe wing is a merevestige, reduced to a slender humerusonly, withouteven articulating facets on the distalend. TOWNSEND,Use o/ Wingsand Feet by Diving Birds. [.July[Auk

The tibia, however,is of greatstrength and size,provided with a keelor crestfor theattachment of powerfulmuscles, and thepatella is enormouslydeveloped. It is evidentthat Hesperornispursued its prey underwater by meansof the feetalone, and that through manygenerations it had graduallylost the useof the wings,which must have been,therefore, a hindrancerather than a help in its subaqueousflight. It had longsince given up aerialflight. Loons and Grebes,however, although apparently allied to Hesperornis, do at times,as we have seen,use their wingsin additionto their feet under water, yet it seemsto me probablefrom the evidence adducedthat as a rulethey progress by the feet alone. The young appearto usethe wings as well as the feet habitually. Thesefacts wouldseem to indicatethat the methodof posteriorpropulsion in Loonsand Grebeshas not beenlong developed nor permanently fixed,and that the youngshow the ancestralor primitiveform of locomotion.The closerese•nblanee in the legsof the Loonsand Grebeson the one hand, and Hesperornison the other would suggesteither a caseof parallelismfrom similarfunctions, or that they were all descendedfrom the samestock. In the 'Birds of Essex County' (26) I spoke of the Loon as "approachingthe winglessconditions." The presentstudies would, however, lead me to believethat the Loon in perfectingthe methodof posterior propulsionunder water, has no needto reducethe sizeof its wings for usethere. It can,however, with advantageincrease their size, providedit doesnot usethem under water, for the wingsare now sosmall that on calmdays it is unableto riseinto the air. Cormorantson the otherhand have for so longa time perfected the posteriorpropulsion method that they do not use the wings underwater evenapparently when young. In consequencethey havebeen able to retainlarge wings for aerialflight. That they can developgreat speedunder water and are very expert fish- catchers is well known. The other line of evolution,the subaqueousflight by anterior propulsion,or by theuse of thewings alone, reaches its height in the Penguins,and probablyin the extinctGreat Auk, two birdswidely separatedgenetically, but convergingto the same result in this particular. Both birds in developingspeed under water by the useof the wings,reduced them in sizeto the proportionsof seal's VoL1909 XXVI'] J TowNsr•.•-•),Useo/ Wingsand Feet by Diving Birds. •945 flippers,--most markedly so in the caseof the Penguins,--thereby showingthat large wings are not only unnecessary,but even a hindrancein subaqueousflight. In attailfingthis end they were obligedto sacrificeaerial flight. This the Penguinswere able to do owingto the absenceof landmammals in theirantarctic breeding grounds. The same conditionsexisted for the Great A•k at its chief breedingplace in this countryon Funk Island, until the arrival of that most destructive land mammal, the white man. The DivingPetrel of theStraits of Magellanis a birdthat appears to be in dangerof sacrificingaerial for sub-aqueousflight, and illustratesthe inconveniencesof this line of evolution. Nichol (14) saysof this bird, after describingits shortflights in the air and its diving: "In appearanceit remindsone forcibly of the little auk.... The wingsare very small and weak, the bird, doubtless,is losing the powerof flight." In the caseof the existingAlcidm and of the other birds that habituallyuse the wingsalone in diving,it would be interesting to determinewhether they are ableto progressunder water as fast as thosebirds that usethe feet alone,for the Alcidmare trying to makethe sametool work for two purposes,to propelthem in the air as well as in the water. One is i•npressedwith the imperfection of their wingsfor bothpurposes, when one watchesa Puffinen- deavoringto get out of the way of a steamer. First the bird dives and flies under water. Then in alarm it rises to the surface and attemptsto ascendinto the air on its wings,but unlessthere is a strongwind to act on its small aSroplanes,it soongives up the attemptand flopsdown into the water again. Althoughit would be difficult to prove, it would seemto me reasonableto suppose that the compressedpointed body of the Loon, with the air ex- pelled from beneath the flattened feathers, would make faster progressby feet actionalone, than by the wingsor by the wings and feet combined,unless the wingswere reducedto the propor- tionsof flippers. It is possiblethat the occasionaluse of the wings observedin thesebirds may be explainedby fright, whichcauses them to "lose their heads," and return to the ancestral form of progression,to a reptilianscramble so to speak,without increasing the speedof theirprogress. It couldalso be arguedthat the wings of Loonsare now so reducedin sizethat their usein emergencies 246 Tow•-sE•D,Use o[ Wingsand Feet by Diving Birds. [.July[Auk

under water is a help and not a hindrance. Experimentson captivebirds in tanksmight determine these facts. That Loonsare able to progressfaster under water than on the surfaceI have concludedfrom suchobservations as the following (26): "Thus on one occasionI waswatching a Loon swimming about, dippinghis head under water from time to time on the lookoutfor food. The cry of anotherLoon was heardat a dis- tanceand my friend immediatelydove in the directionof the other, and, appearingon the surfacefor a moment,dove again and again until he reachedhis companion.At anothertime on the Maine Coastwhile watchinga flock of youngRed-breasted Mergansers swimmingoff the shore,I noticeda movementas of a large fish on the water outside. The Mergansersat onceflapped in alarm alongthe surface of the watertowards the shorewhere I washidden, and I soonsaw that a Loon was chasingthem, followingthem underwater." Theoreticallya Loon shouldbe able to go faster under water than on the surface, for on the surface the bird is retardedby the wavesin front and the eddiesbehind, and the fasterit goesthe moreit is retardedby thesefactors. The subject of the resistanceof submergedbodies has been exhaustively studiedby navalarchitects, and it has beenshown that a prop- erly shaped body completelysubmerged under ideal circum- stances with the wave eliminated •neets with little resistance besidesfriction. The fact that a Loon when swimmingrapidly on the surfaceis apt to depressits body in the water so that its back is awashseems to favor this contention. It •nay be argued that the bird doesthis to avoidobservation or to escapebeing shot, but it certainlyswims faster when thus submerged. Under water the divingbird has a great advantagein beingable to assumea shapebest adapted to cleavingthe liquidmedium. Incidentallyit maybe remarkedthat the Loon,in perfectingits legsfor useunder water, has disabled itself for walkingon the land, but as it usuallybuilds its neston or closeto the water, it can well afford to sacrifice terrestrial locomotion. The combineduse of wingsand feet,a reptilianform of progr.es- sion, would naturallybe found amongbirds that had not fully specializedin eitherdirection. Amongliving birds the Cormorant and the Penguinrepresent the extremesof specializationfor the rot.1909xxvI'l J TOWNSSND,Use o[ Wingsand Feet by Diving Birds. 247 posterior and anterior extremity respectively. Where either habit is not firmly establishedwe shouldexpect at timesa return to the primitivemethod, and we shouldexpect to find it in young birds. This is well shown in the case of the Loon. We should ex-pectto find it at all timesin beginnersin the art of diving,i.e., amongbirds whoseancestry in the diving line is not a long one. The Mallard, the Black Duck, the Gallinule,the SpottedSand- piperand the Water Ouzelmay perhaps illustrate this contention. In conclusionthe following tentative inferencesfrom these preliminarystudies may be setdown. 1st. That progressionby both the wings and feet under water in divingbirds is theprimitive method, and is therefore to belooked for amongbeginners and youngbirds. 2d. That specializationtowards the useof the wingsalone leads to a diminutionin the size of the wings, and finally to a form of bird that is flightlessin the air; for wingsof flipperproportions, too smaflfor aerial flight, are more efficientthan large wingsfor subaqueousflight, as witnessthe Great Auk and Penguins. 3d. That specializationtowards the use of the feet alone is probablybest adapted for the mostrapid progression under water, and this methodmay leavethe wingsundiminished in sizefor use in the air. The apparentexception, Hesperornis, with powerful feet but with wingsdegenerated to vestigesthrough disuse, serves but to confirmthe inferenceof the superiorityunder water of feet action alone.

REFERENCES.

1. Beebe, (1. W. The Bird: Its Form and Function. New York, 1906, p. 4. 2. Ds,rwin, (lbs,ties. Voyage of the Beagle. New York, 1871, p. 200. 3. Eva,ns, I. I-I. Cambridge Nat. Hist. Vol. IX, Birds, London, 1900, p. 52. 4. Loc cit., p. 54. 5. Loc. cit., p. 56. 6. I-Ie•aley, F.W. Life and Evolution, 1907, p. 125. 7. Kit•rerlge,Joseph. The Wren. B?dl.Norf,•lk Bird Cl?•b,I, 1909,p. 5. 8. Lea,,John. The Romanceof Bird Life. London, 1909, p. 210. 9. Loc cit., pp. 202, 203 10. Loc cit., p. 211. 11. Loc. cit., p. 203. 248 R•c•_•ioN•),Ornithological Writings o/ Rafinesque. [July[Auk

12. Loc. cit., p. 205. 13. Newton, Alfred. Diet. of Birds. Article: Ousel. 14. Nichol,M.J. Three Voyagesof a Naturalist. London, 1908, p. 160. 15. llelous,Edmund. Bird Watching. London, 1901, p. 148. 16. Loc. cit., p. 153. 17. Loc. cit., p. 152. 18. Loc. cit., p. 159. 19. Loc. cit., p. 148. 20. Loc. cit., pp. 150-151. 21. Loc. cit., p. 156. 22. Selous, Edmund. The Bird Watcher in the Shetlands. London, 1905, p. 72. 23. Loc. cit., p. 50. 24. Loc. cit., p. 168. 25. Townsen•l,C.H. Bird Lore, X, 1908, p. 171. 26. Townsend,C.W. Birds of EssexCounty. Cambridge,1905, p. 80. 27. Morgan, C. Lloyd. Habit and Instinct. London, 1896, p. 69.

A REPRINT OF THE ORNITHOLOGICAL WRITINGS OF C. S. RAFINESQUE.

PaR• II. •

BY CHARLES W. RICHMOND.

R^•q•F•S;•UE,with his brother, left Leghorn in March, 1802, and landedin Philadelphiaon the 18thof April. He at oncebegan to botanize, collect specimens,and make drawings. He says ('Life of Travel,' p. 17): "My brotherhad becomea sportsman, and procuredme manybirds. I wantedto undertakethe Orni- thologyof theUnited States, finding many of themnew or unknown, or badly described. I continuedalso to study the Snakesand Reptiles,communicating some of them to Daudinfor his work on Reptiles." So, it appears,Rafinesque narrowly escaped being the Father of AmericanOrnithology. However,this plan, like many othersprojected by him, fell through,and on his first visitto the

For Part I see antea, pp. 37-55.