.--During the fall and winter of 1954 and the spring
of 1955, I had an opportunity to observe some hostile behavior patterns in an Oriental
Pratineole (Glareola maldivarum or G. lratincola maldvarum) at the New York
Zoological Park.
This bird was kept in a small aviary, with many passefines (mostly estrildine
finches) and a few larger birds (such as sandgrouse and quail).
Various disputes and fights were not uncommon in the aviary. The pratineole
was remarkably sluggish when left alone; but it was vigorous in attempting to
maintain an "individual distance" area around itself, within which it would not
willingly allow any other bird to approach. It was sometimes forced to escape
before the larger birds; but it could usually induce the smaller trespassers to stop
or retreat. This was accomplished by attacking the trespassers, by directing un-
mistakably hostile displays toward them, or by alternating attack intention move-
ments and hostile displays in rapid succession.
One type of hostile display, particularly common in the spring, was a harsh, loud
"chirrup" note, or a brief series of two or three similar "chirrup" notes. These
sounds were usually accompanied by movements: the head was lifted and pointed
slightly upward, and the neck was simultaneously stretched upward and forward.
Such movements were variable, rapid, and almost always very brief.
A second type of hostile display, apparently threat (see Moynihan, Auk, 72:
247-259, 1955), was somewhat more elaborate and often more prolonged. It in-
eluded a characteristic twittering call and one or two distinctive postures.
The twittering call itself was a rapid series of high-pitched, faint, rather melodious,
monosyllabic notes (impossible to transcribe adequately). The ntnnber of notes
varied from two or three to more than a dozen. This utterance was quite different,
in quality, from any of the calls of the gulls, terns, and shorebirds with which I
am familiar. In rhythm, however, it was slightly reminiscent of some staccato
"long calls" and "crackling calls" of certain gulls (summarized in Moynihan, Be-
haviour, Supplement 4: 1-206, 1955), and also, probably, the "gakkering" call of
the Sandwich Tern (described by van den Assera, De Levende Natuur, 57: 1-9,
1954).
The typical posture associated with twittering was assumed as soon as the call
began. The bird stretched its head and neck downward and forward; and usually
maintained this posture, while the call continued, until its opponent escaped or a
fight developed.
The motivation of this performance, like that of most threat displays in other
birds, was obviously composed of conflicting attack and escape tendencies. These
tendencies, or drives, seemed to fluctuate in relative and actual strength; and these
fluctuations, in turn, seemed to be correlated with variations in the physical form
of the display.
Thus, for instance, the extent to which the bill was opened and the neck was
stretched seemed to depend upon the strength of both drives. When both were
weak, the bill was closed and the neck was short (Figure CO. The neck became
longer and the bill was opened more widely as the intensity of the two drives in-
creased (Figure B.).
The head and neck were sometimes held in an approximately straight line; but
this was relatively rare. The head was usually held at a slightly different angle,
pointing more strongly downward than the neck. The most extensive downward
inclination of the head seemed to be diagnostic of situations in which the pratincole
was "timid" or "frightened," i.e. obviously reluctant to attack with full force.
This would suggest that the extreme downward pointing of the head was produced
by a relatively strong escape tendency, an escape drive somewhat more strongly
activated than the attack drive.
The angle of the neck was usually rather stable. Occasionally, however, just
before attacking, the bird would suddenly stretch its neck upward (Figure A.)
and twitter more loudly. This would suggest that the upward movement occurred
when the attack drive was much stronger than the escape drive.
A very conspicuous movement was sometimes superimposed upon the twittering
posture. The bird would suddenly raise its arms, apparently without flexing or
POSTURES ASSOCIATED wITI-I TI-IE TWITTERING CALL OF TI-IE ORIENTAL PRATINCOLI.g.
From top to bottom. A. Occasional posture before attack. B. Typical moderate-
intensity posture. C. Typical low-intensity posture. D. High-intensity posture
with wings raised. E. Moderate-intensity posture with "crest" raised.
rotating the carpi (Figure D.), and then as suddenly lower them again. This
movement was most effective in revealing the rufous on the underside of the wing
as a brilliant flash of color. It was so rarely performed that the causal factors re-
sponsible for its production were difficult to determine; but it seemed to appear
when both types of hostile motivation were at their strongest possible (at their
strongest, that is, in this particular environment).
Both the twittering postures and wing-movements can be easily explained as
combinations of advance, retreat, and flying intention movements; movements
which have become more or less thoroughly ritualized (i.e. have become standardized
and acquired a function as social signals).
Another behavior pattern, sometimes superimposed upon the typical twittering
posture, was more obscure in function and derivation. The bird would sometimes
raise its crown feathers, slightly darker than the surrounding plumage, to form a
definite, if not very conspicuous, crest (Figure E.). This was also accompanied,
in some cases at least, by a slight raising or ruffling of the back and scapular feathers.
Such infrequent ruffling seemed to be a low-intensity indication of a more extreme
behavior pattern usually produced after the completion of these hostile displays
(both the "chirrup" and twittering). As soon as the pratincole had stopped dis-
playing and relaxed its posture, it would ruffle all its head, neck, and body feathers,
shake out its wings, and then smooth its whole plumage down again.
Ruffling and shaking are obviously "comfort movements" or derived from com-
fort movements; but their apparent standardization after hostile display might
suggest that they may also subserve a social signal function, in a more natural
environment, during intra-specific disputes.
The significance of any reaction by a caged bird, of course, is always difficult to
analyze; and the hostile behavior patterns described in the preceding paragraphs
may comprise no more than a small fraction of the total hostile behavior shown by
Oriental Pratincoles in the wild. (Yeates [Brit. Birds, 41: 301-303, 1948], has
described several apparently hostile patterns of the Common Pratincole [G. p.
pratincola] breeding in the Camargue. Some of these patterns may be related to
those described above; but the nature and extent of such possible relationships must
remain indeterminate until further studies have been made.)
If, therefore, the hostile behavior patterns of this captive Oriental Pratincole have
been discussed in some detail, it is only because they have a certain evolutionary
interest. They resemble certain hostile displays of gulls and terns. The rhythm
of the twittering call has already been mentioned in this connection, but the associated
movements and postures are even more suggestive. Certain threat displays of the
Laughing Gull and the Sandwich Tern include a ruffling component (Noble and
Wurm, Ann. N.Y. Acad. Sci., 45: 179-220, 1943; van den Assera, op. cit., 1954),
and so do other displays in such species as the Frankiin's Gull (personal observation).
More important, the various twittering postures are quite similar in form to some
of the "oblique," "low oblique," and "head-down" threat display postures that
have been found in many different species of gulls and skuas (see Moynihan, Be-
haviour, Supplement 4: 1-206, 1955); and this captive pratincole assumed these
postures and began twittering in situations very much like those in which many
captive gulls would assume an "oblique," "low oblique," or "head-down" posture
and begin the "long call."
This hostile behavior may thus provide further evidence of the close relationship
between the families Glareolidae and Laridae within the order Charadriiformes.--
M. MOYNIHAN, Department of Conservation, Cornell University, Ithaca, New York.