Parental and Courtship Feeding in Red Jungle Fowl

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 lengthand 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

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ROOD9 I AD SINGLE 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). Adult cockstend to escortsingle hens; yearlings to escorthens with broods.The data for Monkey Mesa are moreaccurate because they are basedon repeatedobservations of knownindividuals. On MonkeyMesa only one yearling acquireda singlehen as mate, and his intermediate length spurssuggested he hatchedvery early the precedingyear. The two hens with broods that had adult cocks as consorts had broods that were virtually independent;as both hensnested again shortly,in effect they 26 Ar•.r• W. Stox•s [Auk, Vol. 88 were physiologicallyidentical with single hens. Thus apparently adult cocksnormally court only singlehens; when this resourceruns out the adult cock may court a hen with well-grownchicks who might yet renest the sameseason. Likewise yearling cocks rarely acquirea singlehen, although they may court them when the adult cock is temporarily preoccupied;in- stead they court henswith chicksso their only competitorsare other yearlings. Per•nanenceof the social bon.ds.--In April the relationshipsbetween cocksand hens were already well-established.Bonds between an adult cockand singlehens normally formed well beforeegg laying (See Figure 2, Hens 5 and 5A). But an adult cock might form a bond with a hen almost immediatelyfollowing loss of her brood (as with Hen 6), or at the time her brood reachedindependence at about 8-10 weeksof age (Hens 4 and 22). An adult cocknever stayed with a hen after shehad startednesting, nor did he resumethe bond after her chickshatched. AlthoughI have no evidence, I suspectthat a hen about to,renest, after either lossor independenceof her chicks,will be courtedagain by her originalconsort. Bondsbetween yearling cocks and henswith chicksat first alsoseemed stable. I often found the same cock and hen togetherhour after hour and from day to day, althoughone or more other yearlingcocks often tidbitted and waltzed up to this samehen. Over a 2-month period a hen might have a successionof two or three different consortsas one cock gradually dom- inated his predecessor.Thus Cock 9, a yearling, escortedHen 6 and her chicksfrom 17 April through2 May, whenshe lost all her brood. She then left her usual range and began going with an adult cock. Cock 9 then started courtingHen 18 whosechicks hatched on 2 May; by 7 May he was being challengedand was gradually displacedby yearling Cock 12; Cock 9 still stayed in Hen 18's vicinity, but only rarely did he court her. Such changesin attachment between cock and hen were accompaniedby an increase in the area where the new consort held dominance over his rival. Sometimes a cock maintained his consort role with the same hen success- fully for at least 6 weeks. Thus the yearlingcocks on Monkey Mesa had threepossible roles during the breedingseason. One cockwas able to acquirea singlehen and presum- ably breedwith her; severalstayed close to, an adult cockand courtedhis singlehens when the adult momentarilystrayed from them; by far the most attached themselvesto a hen with chicks,either as consortor as a sub- ordinate to the consort. In contrast all three adult cockson Monkey Mesa acquired one or more single hens, and rarely paid any attention to hens with broods unless the chicks were reaching independence. The roles of juvenile cocks.--I was fortunate to find one juvenile cock as he began to consortwith a hen. This cock was about 3 months old in Jan. 1971] Red Jungle Fowl 27 early April whenI first notedhim with a brother,both alreadyindependent of their motherand sister. I periodicallythrew mealwormsto. this cock over the next 2 months. At first he ate them silently without sharing them with nearbychicks or hens. By late May he was spendingmore and more time near Hen 15 and her youngchicks. On 3 June he was not tidbitting to. her even when within 3 m of her. On 5 June he made a few half-heartedfood calls,but got no response.On 10 June he tidbitted and calledmuch more vigorously and this time Hen 15 cameup and took the worm. Thereafter he tidbitted to her regularly. In addition he largely severedhis ties with his brother. When he started courtingthis cock was about 5 monthsold, and his combwas about the size of a yearling's,but he had no spur.

DISCUSSION The functionof courtshipfeeding.--It is clear that tidbitting and calling serveas powerfulstimuli for a hen to approachthe displayingmale. The hen'sapproach permits the cockto proceedwith other courtshipdisplays suchas waltzingand actualmounting. Furthermore the repetitionof this displayweek after weekmay createso, strong a bondbetween cock and hen that the cockmay wanderaway from her for brief periodswithout danger of her strayingto anothercock. Thus the displayin adult cocksas given towardsingle hens seems to enhancetheir chancesof fosteringprogeny. The value of courtship.feeding to yearlingcocks is lessclear. I first thoughtthe displaycould lead to a bond that would give the cockan ad- vantage should that hen lose her brood and renest. However adult cocks quickly identifiedsuch hens, and supplantedthe yearling cocks. It now seemsmore probablethat the associationof yearling cockswith hens that had chicksprovides a trainingground where they can matchtheir strength and experienceagainst other yearlings.Such yearlings may then be more successfulin acquiringone or more matesthe followingyear. Analogous behavioroccurs in turkeysand variouslek speciesof grousein whichthe yearlingmales display toward the hensand to eachother, but are rarely successfulin mating (Patterson,1952: 152-154; Watts, 1968). The origin of courtshipfeeding.--In monogamousgalliforms the male commonlyplays an importantrole in feedingthe young (Hume and Mar- shall, 1878-1881; Beebe,1918 1922; and numerousreferences in the Avi- culturalMagazine). But evenin polygynousspecies the male may assume parental duties should the hen be killed (ibid.). This evidencesuggests that malesof polygynousas well as monogamousgalliform specieshave some motivation to feed chicks. In the Red Jungle Fowl the form of tidbittingand callingby the maleis virtuallyidentical to that of the female, whetherhe be courtinga singlehen or attractinga hen and chicksto food. 28 ALLE• W. STO•CES [Auk, Vol. 88

I believe, therefore, that the co.urtshipfeeding of Red Jungle Fowl origi- nated as functionalparental feedingwhen the speciesor its progenitorwas monogamous.Polygyny in the Red JungleFowl presumablyevolved when seasonallyabundant food enabled the hen to rear a brood with little or no assistancefrom her mate. A dominant cock could then pass on more progenyto the next generationby acquiringtwo or more hens,even though this meant foregoinghis role of providing food for the chicks. Yet the parental role still persistsin yearling cocks, but even here feeding has becomelargely a deviceto attract hensand to.gain experienceagainst rival cocks. At times it can even be entirely symbolic feeding. Courtship feeding in Red Jungle Fowl has not moved far along the evolutionarypath from functionalparental feeding. With the exceptionof the cock'ssomewhat exaggerated head movements,his turning asideas the hen approaches,and his use of symbolicfood, the behavioris identical in male and female. This is in contrast to certain other galliforms where courtshipfeeding has becomeinseparable. from primary courtshipdisplay, as in the peacockpheasants, Polyplectronspp. (Beebe, 1918-1922). This study has revealedthe possibilityo.f misclassifying breeding patterns of birds from field observations.A hen seenin companywith a cock, both beforeand after she has hatcheda brood, has been usedas evidencethat a speciesis monogamous.I observedessentially this relationshipamong the Red Jungle Fowl at the San Diego.Zoo, but the cock that associatedwith a hen after she hatchedher broodwas always different from her previo.us one,a fact determinedonly by beingable to identify individualbirds. The Red Jungle Fowl at the San Diego Zoo. must be largely polygynous,for yearling cocksseemed rarely able to acquire a mate; the population had about two adult hens per adult cock (Table 1), and many adult cocks were seen in close association with two. or more hens. The zoo population, even though free-living, may not be typical of jungle fowl elsewhere.The jungle fowl begin to breed in spring,when the warmer weather also brings a greater zoo attendanceand thus more food for the birds. This seasonallyrich sourceof food could be conduciveto polygyny and perhaps also to a higher occurrenceof secondbroods than occursin the species'native habitat.

A c KN'OWLEDG2V•ENTS

I am grateful to Kenton C. Lint, Curator of Birds at the San Diego Zoo for per- mission to use the facilities of the Zoo; to H. Warrington Williams for the preparation of sonograms; and to Keith L. Dixon for his constructive suggestionsfor the manu- script.

An unconfinedpopulation of Red Jungle Fowl was studied at the San Jan. 1971] Red Jungle Fowl 29

Diego Zoo from April through June 1969. The male's courtship feeding servesto appeaseand attract the female to him and in this way leads to a pair bond and eventualcopulation. Usually only adult malessucceed in attractingbreeding females through courtship feeding. Yearling males,in contrast, courtship feed hens with chicks. Consortingwith hens that are rearing chicksseems to function as a meansof gaining experienceagainst rival yearling cocksand with potential breedinghens that could enhance their chancesof acquiringa mate the followingyear. Yearling jungle fowl cocksregularly feed the chicksof the hensthey are courting. In addition cocksmay take over the entire role of raising chicks. Perhapsbecause of this retentionof the parental role in jungle fowl cocks,courtship feeding has becomeonly slightly ritualized as a form of display in contrastto its greaterritualization in someother galliforms.

LITERATURE CITED

BEEBE,W. 1918-1922. A monograph of the pheasants (4 vols.). London, H. F. Witherby & Co. CoLics, N. E., ^•I• E. C. CoLL•s. 1967. A field study of the Red Jungle Fowl in north-central India. Condor, 69: 360-386. COLL•AS,N. E., E. C. COLLI^S,D. I'IrYNS^•CEI•,^NI• L. MINmNC. 1966. Locality fixation, mobility, and social organization within an unconfined population of Red Jungle Fowl. Anim. Behav., 14: 550-559. Do2X•M,L.V. 1927. New experiments on ovariotomy and the problem of sex inver- sionin the fowl. J. Expt. Zool., 48: 31-171. GulL, A.M. 1962. The behavior of chickens.Pp. 491-530 in The behavior of do- mestic animals (E. S. E. I-Iafez, Ed.). Baltimore, Williams and Wilkins. Htu•vrE,A. O., ANDC. I'l. MARSHALL.1878--1881. The game birds of India, Burmah, and Ceylon. (2 vols.). Calcutta, published by the authors. JOHNSON,R.A. 1963. Habitat preference and behaviour of breeding jungle fowl in central western Thailand. Wilson Bull., 75: 270-272. KRmJT, J.P. 1964. Ontogeny of social behavior in Burmese Red Jungle Fowl (Gallus gallus spadiceus). Behav. Suppl., 12: 1-201. L•LL, A. 1966. Some observationson social organization and non-random mating in captive Burmese Red Jungle Fowl (Gallus gallus spadiceus). Behaviour, 26: 228-242. P^TT•RSO•,R. L. 1952. The Sage Grouse in Wyoming. Denver, Colorado, Sage Books, Inc. WATTS,C.R. 1968. Rio Grande Turkeys in the mating season. Trans. North Amer. Wildl. Natl. Res. Conf., 33: 205-210. WOoD-Gus•, D. G.M. 1955. The behaviour of the domesticchicken: review of the literature. Brit. J. Anim. Behav., 3: 81-110.

Utah State University, Logan, Utah 84321. Accepted 19 October 1969.