Garrulus glandarius - British Birds

(278) NOTES ON VOICE AND DISPLAY OF THE JAY. BY DEREK

GOODWIN

THE following observations on the voice and display of the Jay (Garrulus glandarius), whilst not to be considered fully comprehensive, seem worth recording in order to amplify the remarks in The Handbook on this subject. Most of the more detailed work has been with tame birds in confinement, but so far as possible all behaviour has been checked with wild birds and no notes have been included in the " innate " list without comment unless they have been heard from wild birds under circumstances that suggest that the correct interpretation has been put on their use by tame birds. As the sexes cannot be distinguished visually and no birds have been killed the terms " male " or " female " will be understood to refer with certainty only to tame specimens. Owing to the fact that certain posturings are characteristically associated with certain notes it seems best to give a numerical list of the former and to save repetition by simply referring to these numbers in the section on voice. POSTURING AND DISPLAY.

(I) Feathers of rump, upper tail-coverts, flanks, belly and (usually) crest and upper mantle erect; feathers of back and upper breast

FIG. I.

MALB IN POSTURE J, VERSION WITH CREST ONLY HALF-ERECT.

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depressed. Wings slightly drooped, but tips usually above or on level with tail and primaries tightly folded, but secondaries slightly spread and usually with the black inner greater coverts drooping in such a way as to interpose a solid black patch between the blue wing-coverts and the (visible) white on the secondaries. The whole effect is to increase and accentuate the black and white area of the wing and to alter profoundly the bird's outline. Used by male when in a state of sexual or aggressive excitement. In many cases it seems to be the outcome of any more or less generalized excitement (from which fear is absent) which finds an outlet in aggressive behaviour. Accompanied either by mimicry or the hissing note [q.v.) or both. When uttering notes in this posture the bird reaches head and body forward while remaining as it were " hinged " on its feet, or if on the ground may walk a few short tottering steps forward ; in either case the procedure is suggestive of some strained and agonized attempt at vomiting. In its less exaggerated forms only the feathers on rump, upper tailcoverts and malar stripe may be appreciably erected. Male of tame pair when uttering hissing note at times appears deliberately to present himself laterally towards the female, and the reaching forward and hissing at the moment that the female happens to fly past the male seems to occur too frequently to be fortuitious. This posturing is commonly associated with heavy formal hops or in flight with a slower and oddly " stilted " wing-beat. (2) Much as last but crest not, or only partially, erect and feathers on mantle somewhat raised giving a different outline, more " roachbacked " instead of the flat back with puffed out rump. Head usually inclined downwards to some extent. Used by female in what appear to be moments of sexual, aggressive or pleasurable excitement. Accompanied by mimicry or the " castanet-note " (q.v.). Both this and the preceding masculine equivalent commonly seen at the '' ceremonial gatherings " in spring. (3) Wings half spread, or sometimes fully spread out on either side of body, arched like a fan and quivered; body and head in somewhat crouching, horizontal position, but body usually well clear of perch; rump and tail somewhat raised above line of back and violently jerked and quivered with a rotary rather than a purely up and down or lateral motion. Seen from above the contrast between the still head and body and the brilliant quivering wings and spasmodically jerking white upper tail-coverts is most striking. May be accompanied by appeal note (q.v.). In less extreme forms bird may merely make slight " suggestion " of extending and quivering wings without tail movements and every intergradation occurs. Used by female towards male and by birds of either sex towards human owner, and was used by male of captive pair when fed by female. May accompany or anticipate a gift of food but appears primarily indicative of a feeling of submissiveness or inferiority or a desire to placate the object at which it is directed. Tame birds

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commonly use it towards their owner when they have been frightened (by some other agency) and are still in a worried state, if he catches another Jay and it screams when handled, or if he shouts angrily at them. In the case of one tame male still in my possession the presence or absence of his mate or other " subordinates " seems the chief factor which decides whether shouting at him in an angry tone elicits the wing-quivering display, or rising anger that culminates in a furious attack at my face. On October 29th alone with him in the lighted shelter after dark, by shouting at him I elicited the full submissive display. Then his mate was brought in as well and one minute after her entry similar shouting at the male evoked aggressive behaviour culminating in attack. I have never known this display to be used towards its owner by any Jay that feared him. Nervous birds do not react at all in the above manner until after they have become quite tame and accustomed to feed from the hand without fear. A Jay purchased in the winter of 1947 that was in a very poor condition at first used the wing-quivering

F I G 2. F E M A L E IN POSTURE 2. I N MORE INTENSE VERSION FEATHERS OF MANTLE, RUMP ANP CREST ARE MORE RAISED, EXAGGERATING THIS APPEARANCE.

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display every time another Jay perched near it or a human being entered the aviary, but as it improved in health it ceased to do so to the same extent. In May, 1947, the wing quivering display by the female of the captive pair before alluded to induced an abortive and unsuccessful attempt at copulation by the male (both hatched June, 1946), but the birds did not then appear to be mated and made no attempt to nest. In 1948 when they nested, although the female was observed greeting the male in this manner on scores of occasions over a period of several weeks, no attempt at coition on his part was ever seen, although the subsequent hatching of their eggs proved that it must have occurred.

F I G . 3. MALE OR FEMALE IN SUBMISSIVE POSTURE. W I N G S ARE ARCHED AND HELD OUT FROM BODY, NOT MERELY DROOPED STRAIGHT DOWN.

(4) Typical juvenile food-begging with flapping or fluttering wings and version of appeal call (q.v.) practically similar to that of juvenile. Both notes and movements of wings are perfectly distinct from (3) and I have seen no intergradations between the two. Observed on three occasions from female of captive pair when fed by male during incubation (out of scores of such feedings observed) and was regularly used towards me by the same bird in 1947 and 1948, but only during the summer moult. This point needs further investigation before one can be at all confident as to its significance, but it seems possible that bodily weakness through hunger or the moult may result in a temporary reversion to infantile behaviour. Similar behaviour in the female Rook (Corvus frugilegus) is well-known and I have observed parallel behaviour in the Carrion Crow (Corvus corone), Domestic Pigeon (Columba livia) and Palm-Dove (Stigmatopelia senegalensis).

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(5) Body feathers puffed out, but crest flat or only partially erect, a threat posture of both sexes. Used by tame birds when attempting to attack or drive off Jackdaws (Corvus monedula) perching on or near their aviary. In minor forms with feathers usually little puffed out may be used when attempting to drive other birds— Jays, doves, pheasants, etc., away from food. May be accompanied by bill-snapping, the anger-appeal note, the rare grating note or the anger-screech. (6) Crest fully erect, body feathers depressed ; head and body jerked violently downwards and sideways from an upright position to one with head well below level of perch and back to original position in one violent movement. Accompanied by alarm note (q.v.) and used by both sexes when seriously alarmed. The same movements in a much less violent form may be used—often silently —under conditions of mild alarm. (7) False bill-wiping, etc.: Exaggerated formal bill-wiping, jerky movements of body and tail, pecking at the perch and the tearing to pieces of any convenient inanimate object, commonly accompany nervous tension, particularly thwarted anger. I have only once seen the Jay hammer at its perch or other objects with closed bill as do crows and the Magpie (Pica pica) when in a state of nervous tension. MIMICRY.

The extent to which the Jay habitually mimics other birds and even non-avian sounds can hardly be over-emphasized. Not only do both sexes frequently utter copied notes, commonly interspersed with low-pitched conversational versions of the appeal note, when foraging or resting at ease, but they are also used in emotional situations, particularly extreme anger. In tame birds, and most likely in wild ones also, there is a tendency for a few stereotyped imitations to be constantly uttered in moments of aggressive excitement. In tame birds favoured items in the repertoire may be " played to death " with irritating persistence for some weeks, but later gradually dropped in favour of new accomplishments if the bird does not again hear the original. Most copied sounds are delivered with remarkable accuracy, differing if at all from the original only in being less loud and more soft and husky in tone, but in some cases where accuracy appears to be impossible a sort of "symbolic interpretation " is achieved. The hooting of the Tawny Owl (Strix aluco) seems to be one of the most popular items in the repertoire of wild Jays as I have heard it wherever I have had opportunities for observing them unseen, in various parts of Yorkshire and in Shropshire as well as in the south of England. The chattering of the Magpie (Pica pica) might be mentioned as one of the notes that the Jay can mimic with absolute fidelity so that no person not seeing the bird could possibly detect the fraud.

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INNATE NOTES.

The extensive repertoire of copied notes possessed by most Jays and their use in emotional situations—particularly at the spring gatherings—make great caution necessary in sifting out the innate notes. I have thought it safe to assume, however, that if a note is constantly used by different individuals in similar emotional situations and the same notes have been heard from wild birds under apparently similar stimuli, that such notes are innate. Even in association with mimicry the innate notes can be picked out as remaining more or less constant, while the copied notes vary from

Jay to Jay. Because of the difficulty of accurately conveying the Jay's notes by means of letter combinations (four people asked to transcribe the same note of the same Jay gave as many different versions) I have tried where possible to describe them in terms of notes likely to be familiar, and where these are helped out by letter combinations these follow the suggestions laid down in the introductory notes to The Handbook. (i) The " Appeal Note ": The most frequently used note, and in all its many variants still to a critical ear recognizably based on the hunger-call of the juvenile. Could perhaps be written " Aaa " or " Aaar " (the r not rolled), or perhaps approaching " Oor " or " Choork " for the softer variants which have a decidedly muted, " kissing," sound. Typically the tone is querulous and suggestive of the mew of a hungry domestic cat, but it is capable of an infinite number of inflections and modulations. Only the more constant variants are listed but it must be understood that every intergradation between them occurs. (a) Hunger call of juvenile, " Aa; Aah; " breaking into a squawking gabble when fed. A very similar version is sometimes used by adult female as previously stated. (b) " A a a " with a very querulous nagging tone. Used by tame birds when hungry, and in a quieter tone under conditions of frustration, as when a timid bird watches others taking mealworms from the hand but fears to do likewise. Is constantly used by wild birds and is probably an expression of a moderate degree of hunger and also of use as a contact note. (c) Very similar to above, but softer and more plaintive, constantly repeated with an indescribably pathetic tone. Used by birds collecting food for young, by a bird with an acorn or other booty whilst apparently " trying to decide " where to hide or bury it, by birds indulging in pursuit flights in early spring that appear connected with pair-formation, by unpaired captive birds (in spring) at sight of any strange Jay, and once by a single bird (wild) that was trailing a constantly displaying pair and hopping about them uttering this note without ceasing whenever they perched posturing at each other. (d) Short, husky, throaty version, deeper in tone but less loud than preceeding. Used by male offering to feed female and by

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female as she accepts food from male or to solicit courtship feeding. Was also used, typically in a softer more crooning tone, by both sexes of tame pair when offering to feed young in nest, and female of this pair at first (?) sight of newly hatched young half raised herself in nest and for about ten minutes alternately fed, or at least appeared to feed them, and uttered a very low, tender, almost " cooing " version of this note. (e) " Anger-appeal " note : A short husky panting note, rapidly repeated and barely recognizable as a variant of the appeal call. Used by tame pair (sometimes) when young were handled and always accompanied by violent physical attack on the offender. Heard twice from wild pairs dashing up in answer to alarm notes of fledglings when the latter were handled (but giving way to the alarm note on seeing man) and once by a pair as they chased a Sparrow-Hawk (Accipiter nisus) out of a spinney where they had young. (2) The Grating Note : A short hard grating note, rather like alarm note pf Mistle-Thrush (Turdus viscivorus). Used, but comparatively rarely, by tame birds when trying to drive away Jackdaws (Corvus monedula) alighting on their aviary. Only heard on three occasions from wild birds : once from one mobbing a Carrion Crow {Corvus corone), once from one as it turned at bay when attacked by a pair of Blackbirds (Turdus merula) and once by a bird as it flew at and struck (twice) a perching female SparrowHawk (Accipiter nisus) about which it and three other Jays, none of whom ventured to follow its example, were hopping silently in posture suggesting mild alarm. (3) The Hissing Note : A loud gasping hiss, with somewhat oi an " r " sound in it, that a friend of mine likened to a " circular saw being allowed to run a little and then suddenly cut off " and which is presumably the note that the Rev. C. A. Johns (British Birds in Their Haunts) describes as " a n accurate imitation of sawyers at work." Not unlike the hiss of a displaying male Golden Pheasant (Chrysolophus pictus) but much louder and harsher, and commonly prolonged into two or even three slurred " syllables." Used by displaying male (Posture 1) and I think primarily in the nature of sexual display although a tame bird habitually used it towards me in an apparently purely aggressive manner when his mate was incubating. (4) The Castanet Note : An odd castanet-like clicking, several clicks being commonly run together. By female in conditions of excitement (posture 2), This note also appears to be used as a " base " for the imitation of mechanical noises. In this connexion it seems worth noting that of a tame pair, which both freely imitate the sound of the lawn mower, the female's version is a clicking imitation of the running of the machine, whereas that of the male is a " symbolic interpretation " quite free from any clicking sounds. I have heard very similar clicking notes uttered in a similar posture

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to those of the female Jay from the Rook (Corvus fmgilegus) and Hooded Crow (Corvus comix). (5) The Chirruping Note : A stammering disconnected chirruping, almost a twitter, very soft and faint in tone, commonly prefaced and interspersed by very soft, tender, muted versions of the appealnote. Used by female (tame birds only heard, owing no doubt to poor carrying power of these notes) towards male, or towards owner in a variety of situations. Appears to be expressive of submissive affection and is also used in moments of apparent perplexity or relief from tension or fear, and sometimes, inter-mixed with mimicry, as a sort of sotto-voce self-directed conversation. Twice heard by male of tame pair. Once when he was moulting (during which time their normal roles were reversed and he was subordinate to his mate, as he was also during the period that she was broody) male was burying a peanut when female flew down and alighted on a spray a foot above him, obviously contemplating robbery. Male then looked up at her in a very meek manner and uttered these notes. On October 20th male in lighted shelter at night attempted to drive another Jay from its perch, but when it refused to be intimidated he seemed nonplussed and drew back a little from it. His mate then flew up and perched between them (quite fortuitously) and he uttered the plaintive twittering notes at her, to which she replied in kind. (a) A louder, less tender, and more " chinking " version of the preceding. Used by male of tame pair when hopping around female in display posture during early (pre-building) stages of breeding-cycle. Usually heard at spring gatherings of wild birds, and at one of these I observed three birds in posture 1 that all appeared to be uttering these notes as they hopped all about a fourth that perched immobile. As, however, there were many other Jays present that could not be clearly seen (as usual it was taking place in thick cover) it is not certain that the actions of these four birds had reference to each other. (6) The " very harsh far-carrying deliberate note . . . quite like heron's " frank " call of The Handbook. In what I think to be its " pure " state this note is a very loud crow-like " Kraah " or " Kraa-aah ", at its loudest, much louder and more arresting than any other corvine utterace known to mk, except possibly the Jay's most violent alarm notes. It has, however, endless subsidiary versions, and the bird constantly intergrades it with apparently genuine imitations of the Heron's (Ardea cinerea) call. Used at the spring gatherings and in its loudest forms associated with intense excitement of the participants. This is doubtless the " loud harsh note " said by W. H. Hudson (Birds and Man) to be used to call the birds together for these gatherings. It may, however, be used by odd birds without having this effect, and gatherings entailing a great deal of excitement and display may take place without this note being used.

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(7) The Alarm Note : The harsh rasping screech that most people naturally but incorrectly assume to be the Jay's most usual note. In serious alarm is delivered with great vehemence and the movements previously described (6). This usually induces similar reactions in other Jays near by. Under what might be called the minimum conditions of alarm that will, or rather may, elicit screeching, the bird makes no violent gestures and may continue to feed, preen or bathe between shrieks. In this form the notes commonly have no effect on other Jays, beyond putting them mildly on the " qui vive," and the difference in intensity of the notes is quite discernible to the human ear, although naturally every intermediate degree of urgency occurs. That fear, rather than anger or protest is—with the exception to be discussed later—the emotion inducing screeching, seems tolerably certain. For example when one handles a fledgling Jay from a wild brood and it cries in alarm the parents at once dash up (if they are within hearing, which is not always the case) and fly about uttering the alarm notes with great vehemence. Yet when I handled the young of a captive pair—on a great many occasions—although I was attacked furiously by both parents, neither uttered the alarm notes, nor was the anger-screech (q.v.) used except on two occasions. So it seems likely that the screeching of wild Jays under these circumstances is an expression not so much of their resentment at the human intruder as of the fear which alone prevents them from attacking him. If I catch one of my tame Jays at night it screeches loudly in alarm, but speaking gently to it soon calms it. In the case of one very tame hen bird it is only necessary to speak a single word or merely to shine the torch in my face for her instantly to cease screeching ; yet there can be no doubt whatever from their behaviour that all the birds hate and dislike being handled most intensely. (7a) The Anger-screech : A very harsh sharp single screech may be uttered by a Jay under conditions that suggest that anger is the emotion inducing it. If a peanut or some other titbit is thrown in the general direction of the subordinate bird of a couple of captive Jays, the dominant bird commonly dashes at it uttering this screech, which has the effect of deflecting the subordinate bird even if it is already diving to pick up the nut. A bird that by accident drops an acorn or other nut that it is eating also utters this note as it dives headlong to retrieve it. A Jay perched on a branch of an elm against the trunk in Kensington Gardens which dropped an acorn screeched in this manner as it—to use the words of a second witness—" literally fell down the trunk of the tree " to retrieve it. In a particularly harsh tone this note is sometimes used by a tame male bird when attacking me ; but it seems significant that this usually happens when the attacks are thwarted or more particularly if I behave in a way that would frighten a less bold and angry bird, such as tapping him on the breast with a finger or throwing a handkerchief over his head. Although at

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present it would seem necessary to consider this note on its face value as a threat, yet I think the emotion concerned might prove on further analysis to be closer akin to alarm than to anger. It is certainly never normally used to make subordinate birds vacate a perch, bath or food-dish. A single sharp screech not to my ears differing from the above may be uttered by a wild female Jay as she is flushed from the nest, although quite as often she flies off silently. (8) Bill-snapping : Loud snapping of the mandibles is used as a threat when attempting to drive other birds—of any species— from food, or as a prelude to attack. The accompanying drawings were done by Mr. R. A. Richardson from my own very poor sketches and are attempts to give a general idea of the postures represented. Once again it must be emphasized that although the drawing may suggest a similarity between the submissive (?) display and the food-begging of the juvenile in actual fact the actions used are so different that no close relation between the two is suggested. On re-reading this paper prior to publication it seems advisable to touch on a few later observations. (a) The Grating Note has since been heard many more times by birds attacking Carrion Crows, Jackdaws or Sparrow-Hawks and by birds attacked by thrushes (Turdus sp.) It appears only to be uttered as the Jay darts at its enemy, or as it turns to defend itself from attack. In some variants (where considerable fear is entailed?) this note approaches closely to a short hard version of the alarm. (b) The Hissing Note appears not to be used by many wild males when displaying. They commonly use various stereotyped phrases, apparently mainly of copied sounds, and usually some hissing sound occurs in this phrase. Therefore although the hissing note as described has been heard from at least three different wild birds it seems possible that it may not be entirely innate. (c) Clicking sounds may occur in the display phrases of (presumed) males, but they are quite different in character from the typical clicking notes of the female. This difference cannot well be described, but there is no danger of the ear failing to distinguish it. (d) Coition was observed in a captive pair on April 23rd, 1949, at 4 p.m. The female, slightly in display posture (2) was on a perch about four feet from the male, who stood very still watching her. Suddenly she went into a most extreme and ecstatic version of the submissive display (3). Her wings were arched and spread to their fullest extent and tilted forwards, as was the whole body, suggesting the display of an Argus Pheasant (Argusianus argus) or Sun-Bittern (Eurypyga helias), the tail was raised and both wings and tail violently quivered. The male flew on to her back and copulated with fluttering wings. The female maintained the wing-spread posture during coition and afterwards jerked herself in the manner that Jays and other passeres commonly do after defalcation, but more violently, for several seconds.