110 DAVID SNOW : MANAKIN DISPLAYS IBIS 103a

THE DISPLAYS OF THE MANAKINS PIPRA PIPRA AND TYRANNEUTES VIRESCENS*

DAVID SNOW Received on 6 June 1960

The displays of comparatively few of the 50-odd species of Pipridae have been recorded (see Sick (1959) for a general survey). Among those that have been studied the strangeness of the display movements is noteworthy, as well as their extreme ritualization and their great diversity between the species. The degree to which the displays are communal also varies. The best known displays are the communal ones in which each individual performs on one or more fixed perches, as in Manacus and some Pipra species.

During a recent visit to British Guiana I was able to see the undescribed displays of two species, Pipra p i p a and Tyranneutes virescm, as well as the display of Neopelma chrysocephalum which has been the subject of a short note by Davis (1949). The observa- tions were made in March, in forest near the Essequibo River between 15 and 30 miles south of Bartica.

I wish to acknowledge a grant from the National Science Foundation (United States) towards the cost of the expedition.

P@ra pipra. P. p i p a is a medium-sized manakin of typical Pipra build-compact, with short

legs, short tail and rather long wings. The male is completely black except for a snowy white cap; the female is olive-green. Males were distributed thinly through the forest, each bird apparently keeping to a limited area. The advertising call, a thin insect-like " chrrrrrrr " lasting about 14 seconds, is uttered rather regularly at intervals of about half a minute; 20 seconds was the shortest interval recorded. It is thus easily possible to walk past a calling bird without being aware of it.

For almost the whole of this time it was present within a limited area of about 50 yards by 50 yards which included a small clearing where a tree had fallen. Here, round the edges of the clearing, it had four or five main calling perches at heights of between 18 and 30 feet; but it also called elsewhere, and the main perches themselves consisted of a small part of a tree rather than one or two twigs only. Another male with a territory about 65 yards away (also including a small clearing) was just audible. The proximity of these two birds was almost certainly not due to chance as, in spite of a good search, no others were found in that area of forest.

The male under observation fed intermittently in its territory, picking insects from the leaves in flight. Occasionally he performed more or less stereotyped actions whose significance would probably have escaped me had I not been familiar with the displays of P. mentalis and P. erythrocephala (see below). These actions were of three sorts: (1) to-and-fro flights between adjacent perches at the same level, over distances of up to about 12 feet. During these flights a soft " flip-er-flip ", probably a wing-noise, could sometimes be heard. These flights were only occasionally seen and the perches used varied, as did the distances between them. (2) Occasionally the bird would quickly turn about on its perch, so as to face the other way, without any lateral movement along

No other call was heard. A single male was watched for 3& hours between 08.00 and 15.00 hrs.

*Contribution No. 1007, Department of Tropical Research, New York Zoological Society.

1961 DAVID SNOW : MANAKIN DISPLAYS 111

the perch. (3) When flying from one perch to another, over a distance of several yards, the bird would sometimes end its flight in a rapid swoop down below the level of the new perch, then up above it and down onto it, so that the trajectory was a shallow S-curve. This is not obvious unless the observer is looking approximately along the line of flight.

The female was first seen, silently approaching one of the male’s calling perches and peering about as she did so. A few seconds later the male, who had been some 30 yards away, appeared near her. He at once began flying round her with a slow flapping flight during which the wing-beat was deep and so slow that the wings were easily visible: the appear- ance was in fact very like that of a large black and white butterfly. Sometimes he flew towards her, sometimes to one side. When she flew to another perch he followed with the butterfly flight. After several seconds the female flew off into cover and the male followed. When they were seen again, he was continuing his butterfly flight round the female, and another male, presumably the neighbouring one, and another female were also present. The entire display was silent and lasted only 3-4 minutes.

The behaviour of Pipra pipra, as shown by these observations, has many important differences from that of P. mentalis and P. erythrocephala (Skutch 1949, Snow 1956 and unpublished). Males of the two latter species display in groups, each bird occupying a favourite perch to which it constantly returns and centred on which its main displays are performed. Display-perches may be several feet apart in the same tree or several yards apart in adjacent trees. At their display-perches males call persistently and at short intervals. The observer’s attention is far more easily attracted to them than it is to P . pipra. Further, the displays are more frequently performed: in my experience P. mentalis or P. erythrocephala could never be watched at their display-perches for as long as three hours without a considerable amount of display being seen. Their displays are also more highly ritualized. In particular the three subsidiary displays of P. p i p a , described above, have reached a more advanced state of development. Rapid and frequent to-and-fro flights between the main display-perch and an adjacent perch 3 or 4 feet away, accompanied by wing-noises, are an important element in their display, as also are rapid and repeated “ about-faces ”. At the height of their display both species fly 30 yards or more away from their display-perches, then return with very rapid flight, uttering a special call and landing on the display-perch with a sudden downward, then upward, then downward swoop. The display towards a female consists of these elements, with in addition a sideways or backwards movement along the display-perch, during which the bird takes such short and rapid steps that it appears to slide. During the slide the legs are fully stretched and the brightly coloured thigh-feathers are shown, and the conspicuousness of the display is further enhanced by various wing and tail movements. On the other hand there is no counterpart, in these two species, of P. pipra’s butterfly flight. Probably P. pipra has no counterpart of the sliding display of P. mentalis and P. erythrocephala: not only was no sign of it seen (P. mentalis and P. erythrocephala continually make little incipient slides when they are on their display-perches), but also the thigh-feathers of P. pipra are black like the rest of the body plumage, suggesting that there is no display in which these are exhibited.

Tyranneutes virescens. T. wirescens is a very small manakin, Pipra-like in shape but in plumage differing from

all the Pipra species in being olive-green with a concealed golden-yellow crest. Sick (1959) in his review mentions it only as one of the manakins which have an advertising call but no display.

Males of this species (presumed from behaviour, the sexes being alike in colour) were found scattered through the forest in small groups of two to four individuals.

A quite different display, directed towards a female, was seen once at 09.30 hrs.

Soon afterwards the display stopped and the birds dispersed.

112 DAVID SNOW : MANAKIN DISPLAYS IBIS 103a

Distances between the calling perches of adjacent males varied from about 30 to 50 yards. The four-syllabled call, a somewhat hoarse whistle aptly described by Beebe (1910) as “ chuckle-de-dee ”, carries sufficiently far for the calling birds to be within earshot of their immediate neighbours. It is repeated with great regularity approximately every six seconds.

A total of 12 hours 39 minutes, covering the entire day except the very late afternoon, with some periods duplicated, was spent watching one bird. This bird called mainly from the side branches of a small under-storey tree, 15 to 20 feet up, and less often from a variety of other perches within 25 feet of this tree, as high as 35 feet and occasionally as low as 5 feet. It went off for short periods to feed, not usually for more than 3 or 4 minutes, sometimes returning with a berry in its beak. It also, between calls, often made sallies to pick insects from the under-side of leaves within about 30 feet of its calling perch. The total time spent calling was remarkable, being 86% of the total observation time. In each of the hours from 10.00 and 17.00 (with one exception, 89%) it never called for less than 90% of the time. A rough calculation shows that it must have called nearly 6,000 times in the day. Other birds watched for shorter periods seemed to be equally persistent.

Six times during the period of observation the bird performed a remarkable and unex- pected display. It was immediately preceded by a sudden reduction in the volume of the call, which became very quiet and hoarse and was uttered rather rapidly, and by the partial erection of the crest feathers. The fist was a floating flight, with very rapid wing-beats, the body being held in a half upright position, the legs hanging down, the neck stretched and thin and the crest fully erected like a spiky golden coxcomb (Fig. la). In this position the bird would “ float ”

There were two elements in the display.

(4 (b) FIGURE 1. Display positions of Tyranneuutes wirescens. (a) The floating flight. (b) The head-

swinging display.

over short distances from one perch to another, or make little floating jumps sideways along the same perch, or occasionally make a long floating flight over a distance of up to ten yards. The longest floating flight seen was accompanied by a sharp mechanical trill which was never heard at any other time. When perched between the flights the soft version of the “ chuckle-de-dee ” was sometimes uttered rapidly, but far more striking was the bird’s posture, which formed the second element of the display. With the neck stretched to the full extent and crest raised, the bird slowly swung its head from side to side of the median line of the body, facing forwards the whole time and holding the position for a moment at the limit of each sideways movement (Fig. lb). It gave a curious impression of craning its neck to see round an obstacle in front of it, first trying one side then the other.

1961 DAVID SNOW : MANAKIN DISPLAYS 113

These displays appeared to be given spontaneously. No other individuals were seen near by, though if another bird had approached and stopped 30 or 40 feet away from the displaying bird it probably would not have been seen.

The display of Tyrannartes virescens bears some resemblance to that of Neopelma chrysocephalum, a considerably larger manakin with different proportions but very like P. oirescens in plumage. When displaying N. chrysocephalum erects its crest, jumps up a few inches into the air with rapidly beating wings, hovers a moment and descends to the same perch. Further, its buzzing call of three to five syllables, which is uttered during this display and at other times, is somewhat like the four-syllabled call of T. virescens.

The extreme development of the floating flight of T. virescens has probably been made possible by its very small size, which is no greater than that of a large humming bird such as Glaucis hirsuta: Haverschmidt (in Zitt.) has recorded weights of three specimens as 6, 6.7 and 8-5 gm. Its floating flight is in fact strikingly like the display flight of the humming bird Phaethornis longuemareus. Chiroxiphia pareola and C. linearis, the other manakins known to jump up and hover, are, like N . chrysocephalum, large species (weights around 20 gm.), and like N . chrysocephalum (weight 15-16 gm.) they hang for only a moment in the air.

Possibly these two species should be congeneric.

REFERENCES

BEEBE, W. 1910. Our Search for a Wilderness. New York. DAVIS, T. A. W. 1949.

SICK, H. 1959. SKUTCH, A. F. 1949. SNOW, D. W. 1956.

Field notes on the Orange-crested Manakin Neopelmn chrysocephalum (Pelz.). Ibis 91 : 349-350.

Der Balz der Schmuckvogel (Pipridae). Life history of the Yellow-thighed Manakin.

The dance of the manakins.

J. Om. 100 : 269-302. Auk 66 : 1-24.

Animal Kingdom (N.Y.) 59 : 86-91.

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