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Vol.•8] GlvFom), and Habits of Pigeons 239

TAXONOMY AND HABITS OF PIGEONS

BY E. Wo GIFFORD TI-IE taxonomicgroupings of the speciesof pigeonsand dovesare basedprimarily on structuralcharacters. Classifications differ in the rank assignedto these characters. In the last fifty years the ranking has fluctuated between extremes. Thus, Salvadori (1895) divides the Columbae into the suborders Columbae and Didi. W. L. Sclater (1950) reducesthese suborders to rank, Colum- bidae and Raphidae. Salvadorirecognizes five familieswithin his suborderColumbae: Treronidae (fruit pigeons), ('true' pigeons),Peristeridae ('ground doves'), Gouridae (crownedpigeons), and Didunculidae(tooth-billed pigeon). Sclater,dealing with Afri- can only, embracesthe first three in his family Columbidae, which carries the implication that he must include therein also Salvadori's Gouridae and Didunculidae. He does not even accord Salvadori'sfamilies the rank of subfamilies. Peters's(1937) classi- ficationfollows Sclater's scheme of two families:Raphidae and Colum- bidae, but divides the latter into four subfamilies: , Columbinae, Gourinae, and Didunculinae. Within the Columbinae, Peters'sorder of listing is virtually that of Salvadori'sfamilies Treronidae, Columbidae, Peristeridae, Gouridae, Didunculidae, and I might add, of Salvadori's subfamilies as well. The taxonomic pendulumalternately swinging from lumping to splittingof genera will perhapsin time prove to be a cultural phenomenonof the sameorder as the alternationsin dressfashions (cf. Richardsonand Kroeber, 1940). Bestsuited for thisdiscussion is the hierarchicalgrouping set forth by Salvadori. Naturally, I am concernedonly with his suborder Columbae, which he divides into five families: Treronidae, Columbi- dae, Peristeridae,Gouridae, and Didunculidae.! havenever kept members of Gouridae nor of Didunculidae. I have had a few Tre- ronidae,but neverbred them. The ensuingremarks concern some of the characteristicactions of the living birds and the extent to which theseactions coincide with the taxonomicgroupings. On the whole there is somedegree of correlationbetween Salvadori's groupings and the actionsof the living birds. The followingbrief noteson characteristicactions of certainpigeons and dovesare basedon ob- servationsof individualsof a numberof ,which the writerhas kept as aviary inmatesat varioustimes over a considerablenumber of years. 240 G•FFORD,Taxonomy and Habits o[ Pigeons I'Auk!. April

SUBORDER COLUMBAE Certain activitiesof pigeonsare well known and serve to dis- tinguishthem as one of the best-markedmajor groups of birds. Such are: (1) feedingthe youngby regurgitation;(2) drinking by immers- ing the bill and suckingin the water, often at a singledraught; (3) division of incubationduties, males sitting during the day, femalesduring the night; (4) monogamyand mating for life. A high degreeof 'paternalinstinct' is manifestedin the feeding and care of the young. In some individual pairs, at least, I have observedonly the malesfeeding the youngafter they have left the nest. Indeed, I have observedfemale indica and Oena capensisfly away from pleadingyoung and leave them to the male parentsto feed. Contrariwise,I haveseen a hen Macropygiaunchall, that had laid only infertile ,repeatedly feed a young Tympanis- tria tyrnpanistria. Consequently,what I refer to as 'paternal in- stinct' is a tendencyrather than a hard and fast rule. The duration of family life in which parentsand offspringform a socialunit is brief and is usuallyterminated when the younghave moltedpartially from juvenal into adult . After that, adults and youngignore one another,or the youngare driven away when coming too close to the adults. Sometimesthe young may drive the parentsaway. The reciprocalattitude of parentsand youngbe- comesthat of unrelatedbirds. While the young are still in juvenal plumage,it is a commonthing for them to perch with the parents, often between the parents. Frequently the parents show marked solicitude, the 'head-, just as they frequently do one another's. Sometimesdoves of other speciesshow similar interestin strangesquabs. In an aviary with permanentboxes for nesting,recognition and defenseof property rights prevail during nesting only. Once the young leave the nest, the parentscease to drive intrudersfrom its vicinity. Most dovesand pigeonsthat ! have kept are enthusiasticbathers, especiallywhen first rains comeafter a long dry spell. They lie on the sidewith one wing raised,so as to allow the dropsto strike under the wing. They will do the samething whenspray is suppliedwith a hose. They are equally ardent sun-bathers,lying belly down with tail and one wing spreadagainst the ground,or lying on the sidewith the upper wing partly spread. Duration of incubation is somewhatvariable. For the family Gouridae it is four weeks, for the subfamily Caloenadinaeit is vol.1•1 •8'[ Gieeoe.v, Taxonomy and Habits o! Pigeons tln'ee weeks, for the ½olumba about sixteen days, for the Peristeridae two weeks.

F.s,Mx•.¾ T•U•RONXD.•E TO live indefinitelyin captivity,fruit pigeonsrequire soft food. They can be inducedto partakeof seedby mixing it with the soft foodand graduallyincreasing the quantityuntil they are entirelyon a seeddiet. On sucha seeddiet, however,they do not thrive and usuallydie after a year or two. One of their mostconspicuous characteristics is graspingwith the toes;some species are almostparrot-like in their ability to hold them- selvesupside down or in a verticalposition, whence the Germanchar- acterizationof Papagei-tauben.Grasping with the toes is also a characteristicof squabsof all speciesof the familiesColumbidae and Peristeridaeand is particularlymanifested when one liœtsa œroma nest. It would seemalmost that this is an infantilepigeon characteristicthat is held overin adult life in the family Treronidae.

FAMILY COLUMBIDAE SubfamilyC'olumbinae.-Next to the Treronidae,these pigeons grasp with their toes to greatestextent and are most arboreal or perching in their habits. There seemsinsufficient differentiation in the habits of the variousspedes of ½olumbato warrant elevating the severalsubgenera to genericrank aswas done by Ridgway(1916). All Columbinaewhich I have kept partookof seedreadily. Sub[amilyMacropygiinae.--Experience with thissubfamily is limited to three species: unchall, M. tenuirostris,and M. ru[a. They are verylong-tailed, short-legged birds, which, in their prefer- encefor perching,substantiate Salvadori's placing them next to the Columbinae,rather than next to the genusGeopelia of the family Peristeridae,where Baker (1913,3: 237,252) puts them. On the groundthey are clumsy,for their shortlegs and long tails militateagainst an easygait. Often the longtail is carriedslightly elevated,apparently to avoiddragging on the ground. In cooing to the hen, the cockexpands his cropregion extensively so that it touchesthe ground. He bowsslightly, raises and lowersthe front of his bodyslowly. His tail is kept horizontaland slightlyspread. Their actionssuggest that they belongwith the Columbidaerather that with the Peristeridae. 242 Gxrro•.•,Taxonomy and Habits oi Pigeons L[Auk April

FAMILY PERISTERIDAE The wide variation in behavior in this family seemsto warrant groupinginto subfamilies,although within certainof thesethe be- havior is not so uniform as might be expected. All that I am ac- quaintedwith feedone anotherjust prior to coition. Sub[arnilyZenaidinae.-This American group is quite homogene- ous. The malesof the speciesthat I have kept make conspicuous displayof the iridescentneck and cropregions when cooing. The expansionof theseparts gives prominence to the brilliant coloring. Sub[arnilyTurturinae.-This Old World group is anotherhomo- geneousone in habits. In payinghis addressesto the hen, the cock pointshis bill somewhatdownward, expands his throat,and bows rapidlyand repeatedly,sometimes taking a few stepstoward her if shemoves away. The tail is only slightlyexpanded if at all, and is not elevated above the normal level. Sub[arnilyGeopeliinae.-The genus is Asiatic,Malaysian, and Australian in distribution, while Scardafella is American. Gyrnnopeliaerythrothorax, the single species of a monotypicgenus, put by Salvadoriin thissubœamily, has been placed now in the genus Metriopeliaunder the nameof Metriopeliaceciliae (Peters, 3: 102). That it does not belong in the subfamily Geopeliinaeis apparent from its short tail and different behavior. That it belongswith the other Metriopeliaefrom the standpointof habits,I am unprepared to say,as I have neverkept other Metriopeliae. In appearanceand actionsGyrnnopelia erythrothorax is very differentfrom Geopelia and Scarda[ella. It runs asrapidly as a quail on the ground,suggest- ing Lophophapsin this respect. In cooingit doesnot spreadand elevate the tail to a nearly vertical position as do the Geopeliae. (Incidentally,Stictopelia cuneata certainly agrees with the genus Geopeliain thisrespect.) In perchingand cooingGyrnnopelia raises and lowers the tail slightly without spreadingit. Indeed, in the mating seasonit keepsits tail in motion like a pipit. In doing this the tail is not moved far from a horizontal position. This behavior is different from anything I have seen in Geopelia. I think its re- movalfrom Geopeliinaeis warrantedon the basisof actions. Except for humeraliswhich has a -likecall, the speciesof Geopelia havea croaking,more or lessfrog-like coo. Subfamily Peristerinae.--ThisAmerican group comprisesseveral genera,of which I have experienceonly with Charnaepelia(Colurn. bigallina),Claravis, and the afore-mentionedMetriopelia ceciliae. Charnaepeliaehave one very characteristicaction, the rapid twitch- ing of onewing by the male,when he is payinghis address to the hen. Vol.•8] G•Fro•o,Taxonomy and Habits ol Pigeons 243 t94• -•

This takesplace either when sittingon a perchbeside her or when cooingto her. This is leastnoticeable in C. cruziana,but very char- acteristicof C. passerina,C. minuta, and C. talpacoti. C. cruziana has at timesbeen placedin a monotypicgenus, Eupelia (Ridgway, 1916:989). Unlike other Chamaepeliae,it has a call that is so like thecroaking of a frogas to be mistakenfo• suchif thebird is not visible. Claravispretiosa has two noticeablydistinctive acts. When alight- ing, the head is dartedforward, then quickly pulled back to normal position. In cooingto the hen, the cocklifts his feet very high and puts them down slowly. Sub[amilyPhabinae.--This Old World group is African, Asiatic, MalaysJan,and Australian in distribution. It is divided taxonomi- cally into two groups,distinguished by presenceor absenceof a pair of dark transversebars on the rump. In the first are the African genera Oena, Tympanistria,and Chalcopelia (now ), and the AsiatioAustraliangenus ½halcophaps. From the standpointof actionsthese should be distinctgenera without a doubt. Oena capensisis a very long-taileddove, quite butterfly-likein its a•rial activities,one of whichis a hovering,descending flight with tail spreadon the part of the malewhen disporting before the female. Whenalighting, both sexes raise and lower the tail. In cooingto the hen, the cocklowers his head,elevates his tail withoutspreading it, and movesboth wings slightly and rhythmicallyat the rate of about one hundred times a minute. Tympanistriatympanistria, the , is rightlynamed for its call,which is producedwhile the malestands perfectly still with thesides of thethroat much expanded and palpitating rhythmi- callyas he pipeshis call. Sometimes,in displaying to the hen,the cockelevates his unspread tail slightly.There is no bowingmotion as with the Turturinae. Chalcophapsis as distinctiveas Oenain its matingactivities. The displayof the cockto the hen can bestbe describedas a ridicu- louslyslow jumping-jack movement in which the cockraises and lowershimself vertically several times beside the hen beforemount- ing her. Of the seconddivision of the Phabinae,I havekept only , Lophophaps,Ocyphaps, , and . Unfortunately the last wasrepresented by femalesonly. The two speciesof Phapshave a verybrilliant display, the male spreadingwings and tail in a partiallyvertical position, so that the iridescentcoloring on the wingsshows to advantage.At the same 2z•z• G•rror•,Taxonomy and Habits oi Pigeons Li'Auk Ap151

tinhehe makesa sortof waddlingmotion from sideto sideor he may take a few jumping stepsforward toward the hen. Sinhilarityof activitiesjustifies placing these two speciesin one genus,rather than in two as Mathews (1913: 18) has done. Mon0typicOcyphaps and Lophophaps,the latter solelyterrestrial in habitsand a very fast runner, both have a rapid, dancing,jumping- jack mating displaywith tail elevatedand spreadagainst the par- tially openedwings, resulting in the iridescentspeculum of the wings showingbrightly. This 'dance' is nmch more rapid than the slow jumping-jackmovement of Chalcophaps.It is accompaniedby a marked brighteningof the iris. In flying, Ocyphapshas whistling wings,and flips its tail up, then down, when alighting. Sub[amily Geotrygoninae.-Quail-doveand Pigeon are terms which correctlyexpress the galline appearanceof thesebirds with their long legs, short tail, striding gait, and bobbing tails. Otidiphapsis quite pheasant-likein walk and actions;the othersare quail-like. When frightened,, Phlogoenas (except Terricolumba), and Leucosarciahave the habit of putting the breast against the ground,often in a cornerand elevatingthe tail sohigh that the under tail-covertsare toward the observer. The group of Phlogoenasor Gallicolu•nba,which Hachisuka (1931) has separatedas the new genus Terricolumba, lacks this habit. However, it possessesdis- tinctive positivehabits, viz., a barking call and a disheveled,loose- wingedappearance when courting or fighting. It wouldseem that the separationis madeon goodgrounds so far ashabits are concerned. Hachisuka'sdelimited genus (formerly part oœ Phlogoenas)embraces the so-calledBleeding-hearts (G. luzonica, G. crinigera,G. rufigula,etc.), birds that are characterizedby a car- mine, orange,or yellow patch in the centerof the breast. Besides the tail-elevationwhen frightened,other very distinctiveacts of these birds are: noddingwhile walking, and throwing the head back and expandingthe breastwhen cooing,so that the coloredspot shows to greatestadvantage, an act apparentlycomparable in purposeto the wing-spreadingin Phaps,Ocyphaps, and Lophophapsto displaythe speculumand iridescence. I have observedin both Geotrygon montana and Gallicolumba rufigulathe habit of raisingboth wingshigh abovethe back while standing,or while sittingin the nest. This is part of the courtship, malesdoing it more frequentlyand raising the wings higher than females. It is done when the mates are apart. Another habit of thesetwo species,and probablyof othersof thesegenera, is stamp- Vol.1•41 58] .I GIFFORD,Taxonomy and Habits oJ Pigeons 245

ing hard on the groundby the maleswhen courting. This is evi- dently analogousto the high steppingof Claravispretiosa men- tioned above. Starnoenashas the curioushabit of stoppingshort with head up when walking, then in a few secondsgradually lowering the head and resumingits walk. Mealworms and earthwormsare especiallyrelished by Galli- ,Geotrygon, Starnoenas, and Otidiphaps. Also my Phaps chalcopterahunt earthwormswhenever damp groundis spadedfor them. One day I observeda male Geotrygonmontana attempting to swallowa newly hatchedbut dead Chamaepeliasquab which lay on the ground. Evidently food must form a considerable part of the diets of the Geotrygoninaein their native .

LITEP.ATUP. X CITED BAKER, E. C. STUART 1913. Indian pigeons and doves. 8vo, xvi 4-260 pp., 27 pls.; London, H. F. Witherby & Co. HACHISUKA, HON. M. 1931. Notes sur les oiseaux des Philippines. L'Oiseau et la Revue Fran•aise d'Ornithologie, (2) 1: MATHEWS, G. M. 1913. A list of the birds of . 8vo, xxvii 4-453 pp.; London, H. F. Witherby & Co. PETEVa,J. L. 1937. Check-listof birds of the world. See vol. 3, xiii 4- 311 pp.; Cambridge, Mass.,Harvard University Press. RtCnARDSON,JANE, AND K•O•ER, A. L. 1940. Three centuriesof women'sdress fashions: a quantitativeanalysis. Univ. of California AnthropologicalRecords, 5: 111-154. R1DGWAYs ROBERT 1916. The birds of North and Middle America: etc. Bull. U.S. Nat. Mus., no. 50, pt. 7, xiii-{-543 pp., 24 pls. SALVADORI,T. 1893. Catalogue of the Columbae, or pigeons, in the collection of the . Cat. Birds British Mus., 21: xxx 4- 676 pp., 15 pls. SCLATER,W. L. 1924-30. SystemaAvium Aethiopicarum. A systematiclist of the birds of the Ethiopian region. Part 1, iv 4- 304 pp., 1924; Part 2, xi 4- 305-922 pp., 1930; London, Taylor & Francis. Universityof Cali[ornia Berkeley,California