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Courtship Behavior of the Greater Bird of Paradise

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Courtship Behavior of the Greater Bird of Paradise COURTSHIP BEHAVIOR OF THE GREATER BIRD OF PARADISE JA•s J. D•s•o• I• September1909, 48 Greater Birds of Paradise (Paradisaeaapoda apoda) of unknownsex were releasedon 280-acreLittle Tobago Island (11o18' N, 60ø30' W) in the southernWest Indies (Ingram, 1911). The birds had been obtained on th'e Aru Islands south of New Guinea and were placedon Little Tobagoin the hope of establishinga population safe from the plume hunting that threatenedtheir existencein their native home. Augmentedby three morebirds by 1912 (Ingram, 1913), the pop- ulation apparentlyfluctuated in numbersfor the next 20 years,but no one censusedthem carefully (Ingram, 1917, 1918; Baker, 1923; Guppy, 1931). The islandbecame a sanctuaryin 1928,but by 1955 only 11 apoda were th'oughtto survive (Ingram, 1956). E. T. Gilliard (19'58, and MS field notes) thoughtperhaps 35 birds were presentin 1958 but he was certain of only 15. A hurricanein 1963 may have causeda further re- ductionin the population. Thus althoughthe Little Tobago population has survived,it has never thrived as originallyhoped. I spent 9 months on Little Tobago studying this speciesin 1965-66. In that time I locatedonly 7 apoda; 4 full-plumagedmales, 1 subadult male, and 2 birds in the chocolatebrown plumage worn by both females and young birds. At least one of these was a female. I watchedapoda on Little Tobago almost daily from 23 September 19'65to 4 July 1966. Roldan George,a residentof nearby Tobago,helped in the field work and continuedobservations through 30 September1966. The period reportedon here is 1 October 1965 through 30 September 1966. We worked mainly at severaldisplay groundsthat the birds used extensivelyand in particular one that seemedto be the centerof activity. We usednatural hiding placesand took extensivefield noteson the court- ship behavior. As apoda are extremelywary, I made no attempt to captureor mark any of them, but I could distinguishadult males by differencesin the color and length of the side plumes. I actually watched the birds for about 364 hours and securedadditional information over many hours when I could only hear them. In all I watched304 courtshipdisplays and heard an additional 168. Knowledgeof the courtshipof apodahas heretoforebeen based largely on a few brief descriptions.Wallace in his classicalaccount (1869: 466) describeshow 12 to 20 full-plumagedmales on the Aru Islands assembled to play and dancein lofty, large-leavedforest trees, 305 The Auk, 87: 305-321. April 1970 306 JAMESJ. D•NS•ORE [Auk, Vol. 87 raise up their wings, stretch out their necks, and elevate their exquisite plumes, keeping them in a continual vibration. Between whiles they fly acrossfrom branch to branch in great excitement, so that the whole tree is filled with waving plumes in every variety of attitude and motion.... the wings are raised vertically over the back, the head is bent down and stretched out, and the long plumes are raised up and expanded till they form two magnificent golden fans. No one sinceWallace has describedthe displayof apoda from the Aru Islands. Baker's description(1923) of its display on Little Tobago is similar to that of Wallace. Crandall (1936) photographedand briefly describedthe displayof captivebirds and Iredale (1950) also has brief noteson the display. Gilliard (MS field notes) wrote the best previousdescription of the displayof apodaon Little Tobago. He noted that malesgathered at edges of a display area, each on a secondaryperch. Here each bird called and rapidly flappedhis wingsagainst his sidesin short, rapid wingbeats. From there the birds movedto the main display limbs where, with body held upright, they extendedtheir wings forward and erectedthe plumes over the back. Then abruptly the tail was depressed,the head,wings, and body were loweredso that the body was closeto the limb, and the plumes were still held erect over the back. In this positionthe bird movedalong the limb, sometimesprancing on stiff legs and at other times charging rapidly. The bird then stoppedabruptly and assumeda frozen posture, the bill pointing down, wings spread,and plumeserect. Gilliard called this the "flower" positionand consideredit the climax of the display. Gilliard observedthat when no femaleswere present,the malescharged back and forth• on the limbs, often in synchrony. However, when females were present,they assumedthe "flower" position. This positionwas only assumedon the centralor main dancingstages (courts). He also thought that the male with the longestplumes tended to assumethe centralposi- tion in the court and other males assembledaround him, each with a private perch. DIS•,•,^¾ G•ou•:Ds I located four main display grounds (the tree or trees used for dis- playing by one or more males) on Little Tobago. I occasionallysaw dis- plays at trees outsideof these four, but the use of these trees was too irregular and infrequentto ascertainany exact usagepattern. Three of the display groundswere within 63 m of each other on a woodedhillside. Each seemedto have one plumed male in regular attendance. The three birds could easily hear and probablyoften see each other. They often visited each other'sdisplay grounds,but the regular male at each used a centrallylocated court (the limbsof treesused repeatedly for displaying) April 1970] Greater Bird oJ Paradise 307 while the others seemedto use peripheralperches and courts. The fourth adult male had short sideplumes and probably had just reachedmaturity. This bird apparentlyhad no court he attendedregularly and was seldom seen. The fourth display ground was 365 m from the others and seemed to be used transiently. The two most frequently used display grounds were on the west slope of a hill and thus were not exposedto the early morning sun. Elsewhere I have described in detail the limbs used as courts and/or perches(Dinsmore, 1967). In general the principal requirementsfor a court seem to be a horizontal or gently sloping leaflessbranch, 8 cm or less in diameter, under a fairly thick canopy cover. Perches seem to re- quire thicker canopy. When deciduoustrees dropped their leavesin the dry season,the birds desertedperches there and establishednew perches in thick-canopiedevergreen trees. Most courtsand percheswere in under- story trees such as Eugenia sp. and Mayepea caribaea. Three of 13 courts had lateral scratch marks or small holes on them, apparently from bill-wiping and pecking, but none was worn smooth of bark. I never saw birds clearing leaves from the courts. Rather they seemedto chooselimbs or parts of limbs that were naturally clear of leaves. CALLS The basic call note of apoda is a deep loud "wauk" or "wonk." This note is deliveredin a number of ca]ling patterns. Severalof thesepatterns Mong with a few other ca]Is are associatedwith courtship display. The courtship display, describedin the next section, consistsof five main phases(wing pose,pump, bow, dance,and mounting). Rising calL--In this call four or sometimesfive deliberate"wauks" are repeatedat about 1-secondintervals, the first two notesof equal intensity and the rest successivelylouder and higher. Birds give this call, at the rate of about one per minute, most often in early morning from near their display grounds. They often give a similar call in late afternoon before they roost. Rapid wauk call.--This call is a series of rapid, equal pitch "wauk" notes deliveredin short bursts of severalper second,usually accompanied by synchronousshort wingbeats, the wing tips beingmoved about 2 inches from the bird's sides and then flapped back against the sides with' each note. It is usually given when a female is near the display ground and, with the followingcall, accompaniesthe wing posephase of the display. Wing pose calL--This call, given with the wing pose phase of the dis- play, is a shrill sliding"eee-ak" note, often repeatedseveral times. While calling the bird throws the wings forward into the wing pose position. Birds often alternate this and the precedingcall, deliveringrapid "wauk" 308 J^•r•s J. D•sz4oR• [Auk, Vol. 87 notes with the wings held forward rather than flapped against their sides as previously described. Pump call.--This consistsof a series of very rapidly given "wauk" notes of equal pitch, even faster than the rapid "wauk" call described above. The notes often are run together so that it sounds more like "wa-wa-wa-wa." Lasting up to about 10 seconds,the pump call is given without break during the pump phaseof the display. Baa calL--At the condusionof the pump call, the bird usually gives one or several drawn-out, harsh nasal "baa" notes as it tips the body forward and down. Click call.--During the dance phase of the display, the birds utter a slow and rhythmic "click" at about 1-secondintervals, much like the soundmade when a personclicks his tongueagainst the roof of his mouth. Between "clicks" I sometimes heard a faint nasal "bonk" note as the bird raisedthe wings for anotherdance step. Nasal call.--Similar to the "baa" call but more abrupt and nasal, this call is given most commonlyafter a display, either as the bird bill-wipes or moves from tree to tree tearing off leaves. Chugich call.--This call, heard only a few times, is a rather harsh, guttural "chug'-ich," "chug'-a," or "chug'-a-la" note. It is given in two quite different contextsand may be two distinct but similar calls. Males give it while bouncing along the court prior to giving the "dick" call and starting the rhythmic bouncing of the dance. I also heard it just after a bird left a display groundbut was still nearby, usually as he was about to move to his evening roost. COURTSHIP DISPLAY I recognizefive main phasesin the courtshipdisplay of apoda. They are describedin the sequencein which they usually were given. A more thorough'description of the sequenceis presentedlater in Table 1. Al- though the term "dance" often has been used in referenceto the entire display of apoda, in this discussiondance refers to only the fourth phase of the display.
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