Parental and Courtship Feeding in Red Jungle Fowl
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PARENTAL AND COURTSHIP FEEDING IN RED JUNGLE FOWL ALLEN W. STOKES TH•s paper describesparental and courtshipfeeding in a free-living, semidomesticatedpopulation of Red JungleFowl, Gallus gallus,and dis- cussestheir functionsand possibleorigin. Publishedbehavioral studies of unconfinedjungle fowl include those of Beebe (1918-1922), Johnson (1963), Collias et al. (1966), and Collias and Collias (1967). Kruijt (1964) and Lill (1966) studiedcaptive Red Jungle Fowl. Wood-Gush (1955) and Guhl (1962) review the behavior of domesticfowl. These authorssay little aboutparental or courtshipfeeding. The youngof gallinaceousbirds are precocial,but they needhelp from one or both parentsto obtain food during their first few weeks. Typically the parentlocates food and behaves in sucha way that theyoung will get it. The parent may let the chick take the morselfrom its beak, drop the food in front of the chick,or usespecific calls to alert the chick to.the presence of the food. All thesebehavior patterns I includeunder the term parental feeding. In many galliforms (Stokesand Williams, MS) the male may perform somewhatsimilar feedingbehavior toward the hen as part of his courtship. This is commonlycalled "tidbitting" (Domm, 1927) from the fact that the cockoften displayswith choicemorsels. The Red JungleFowl in the 40-hectareSan Diego Zoo are unconfined, but eachbird belongsto a small socialunit and remainswithin a territory the entireyear (Colliaset al., 1966). The birdsobtain some natural food, but rely primarily upon food providedfor other exhibit animalsby the zoo keepersand the generalpublic. I made intensivestudies largely on the birds of the Monkey Mesa from April throughJune 1969. I could recognizeindividually 16 cocksand 22 hens that used this area by means of comb configuration,scars, brood size, and other characteristics.They formedthree separateflocks, each dominatedby a singleadult cock,with little overlappingof homerange. The birdswere so tame I couldfeed them at closerange. The zoojungle fowl are normallysingle brooded with a peak of hatching in April. In 1964 and 1965 the midwinterpopulation ranged between 125 and 142,with cocksmaking up 56 per centof the total (Colliaset al., 1966). In 1969I estimatedthe populationat 222 birds,almost double that of 1964 and 1965. In a singleafternoon's census on 21 May of almostall the avail- able habitat I counted 124 cocks. These fell into two distinct age classes 21 The Auk, 88: 21-29. January 1971 22 ALLEN W. STOKES [Auk, Vol. 88 FEMALE ! SECONDS 1.0 Figure 1. Food ½a]]o:[ Red Jungle Fowl A, the :[ema]e;B, the male. on the basisof spur length and curvature. The 48 cockswith long, curved spursI judged to be at least 2 years old; 76 with shorter,blunt spurs I called yearlings. In their study of Red Jungle Fowl in northern India Collias and Collias (1967) found the breedingseason and sex ratio almostidentical with those at the San Diego Zoo, but the densityof jungle fowl in goodhabitat was only 6 birds/100 hectarescompared to the 555 birds/100 hectaresat the zoo in 1969. Feeding behaviorof the hen to her chicks.--During their first few days after hatchingjungle fowl chicksremained within a meter of the hen. When foraging the hen was ever on the move. She shifted the litter with a side- ways sweepof her beak and scratchedwith her feet. When she found a suitablepiece of food during thesefirst days shepicked it up, held it above or in front of the chick, and waited until it took the morsel from her beak. In good foragingareas she found food every few seconds.This movement of the beak near the chicksattracted their attention,and they soonbecame alert to her motions. At this stage the chickswere so.close to the hen she neededto move only her head to give them the food. In a week the chicks started to forage farther from the hen. Then when she locatedfood she sometimescarried the morsela few stepsto the chick and let it take the food from her beak or droppedit in front of the chick. Jan. 1971] Red Jungle Fowl 23 TABLE I INVENTORYOF RED Ju>mnE Fowl AT SAN DIEGO Zoo, 1969 27 April-I May 21 May Total Adult cock: With singlehen 16 16 32 With hen and chicks 5 2 7 Alone 19 ? ? Yearling cock: With single hen 9 5 14 With hen and chicks 23 6 29 Alone 31 ? ? Hen and chicks: With cock 28 8 36 Alone 26 13 39 Single hens: Alone 3 ? ? Total cocks observed: Adults 31 48 79 Yearlings 48 76 124 Per cent adult cocks 39 39 39 Cocks/100 hens (from Collias et al., 1966: 551) 137 Hens/adult cock 2: I More commonlyshe gavea food call (Figure 1A) which alerted the chicks and brought them on the run to her. By this time she usually dropped the morselbefore the chick, but often she merely pointed her beak at the morselwithout picking it up. As the chickscontinued to developand to forage still farther from her, the ben'sbehavior changed still more. Her food call becamelouder, faster, and longer. In addition she sometimesdabbled at the food with exag- geratedup and down head movements,two or three times per second. Chicksresponded to theseauditory and visualsignals when as far as 10 m from the hen. The hen stoppedsignaling as soonas the chicks arrived. When the broodforaged where food was plentiful, the hen neithercalled nor dabbled, but let the chicks feed for themselveswhile she also fed. Meal- wormsproved highly attractive,and henswith youngchicks never failed to tidbit with them. At 8 to 10 weeksthe chicksbecame fairly independentof the hen and vice versa; when given a worm the hen at this stage merely ate it, without callsor headmovements. The food call has the physical propertiesof soundseasy to. locate: abrupt start and stop, brevity, and wide frequencyrange. Food calls of other galliformshave the samecharacteristics and may be almost identical in form (Stokesand Williams, MS). Parental feeding by the cock.--About half o.f the hens with broodsac- quired a consortcock (Table 1, Figure 2) that helped her feed the chicks. When such a group foraged, the cock was usually farther than the hen from the chicks. When he found food he gave the same food call as the 24 ALLE• W. SToars [Auk, Vol. 88 24[ BROOD •ND BR INC. iBROOD] [YR • ? LA D • I AD • I •5 SINGLE INC 2] A D •' I •22[ BROOD--OFF AREA IND BR• .AD•i j ROOD9 I ADSINGLE 2 2 2 I II 2I I II 21 31 10 20 MARCH APRIL MAY JUN E Figure 2. Associationof male Red Jungle Fowl with females in relation to repro- ductive state of the female. hen (Figure lB) and the sameexaggerated head movements,at which the chicks and hen ran to him. I have seen a cock freeze with his beak a few cm above the grounduntil the chick cameand took the morsel,but more commonlythe hen ran up to him. As shecame to within 0.5 m of him he usually turned away from her, and she then in turn repeatedthe call over the food until her chicks arrived. Courtskip]eeding.--Male jungle fowlscommonly tidbitted to hensat the zoo. They did this to half-grownpullets, to hensready to lay, and to hens with chicks. The displaymovements and call were virtually identicalto thoseboth cock and hen usedwhile feedingchicks as describedabove, but the tidbitting cockusually picked up a greatervariety of items: the tiniest of morsels,whole peanut shells,palm nuts, wood chips, leaves, and leafy twigsup to 0.5 m long. Hence the feedingwas sometimessymbolic. Meal- worms releasedstrong tidbitting, but any novel or uncommonfood also served. If a hen did not run to the cockimmediately, he usuallyintensified his calling and motions. At times he carried the item a few steps toward the hen, especiallyif he found it partly or well-hidden,but only rarely wouldhe go right up to her. Occasionallyhe let the hen take food from Jan. 1971] Red Jungle Fowl 25 TABLE 2 SEXUAL BONDSIN RED JUNGLE FOWL IN RELATIONTO AGE O1*COCKS Monkey Mesa Remainder of zoo Adult Yearling No Adult Yearling males males males males males Single hens 8 1 0 38 14 Hens with brood 2 8 6 6 35 his beak. He wouldrepeatedly pick up the foodand dropit, dabblewith it on the ground,or more rarely freezewith his beak over it. A cock might continuesuch behavior for well over a minute,and then if no hen arrived, eat the morsel. The displayingcock partially flexed his legs, kept the wingsdose to the bodywith primarywing tips concealed, and heldthe. long tail feathershigh off the ground,but the only essentialcomponents of the displayseemed to be the call, the headmovements, and the actualdabbling with the morsel. The sight or soundof a tidbitting cock was a powerful stimulant to nearby cocksto begin tidbitting. If the latter were not near food, they picked up whateverwas at hand, often large leavesand small branches. The conspicuousmovements and associatedloud calling often attracted the henand her chicksto the secondcock, and sheand her broodmight then run back and forth betweenthe two cocksstanding perhaps 6-10 m apart. These rival cocksrarely fought, but often low-intensitydisplay and chasing ensuedthat madethe subordinatecock withdraw temporarily a meteror so and stop tidbitting. Sometimesafter such sessionsof rival tidbitting the hen shifted her attachment from one cock to the other and let him stand closeto her whileshe was feeding or resting. The sightof a mealwormmust be a strongstimulus for the cockto eat it, but he rarelydid so.while a henwas within about10 m of him. Exceptions were when a cock had been dominatedrecently by another cock standing nearby,after somegeneral disturbance in the vicinity,or whenthe hen and chickswere actively feedingwhere food was plentiful. A cock'sassociation with hensdepends upon his age and socialstatus (Table 2).