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Studies of the Guild of Ant-Following Birds at B~Lém, Brazil

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Studies of the Guild of Ant-Following Birds at B~Lém, Brazil Stud ies of the guild of ant-following birds at B~lém , Brazil c·) YOSHIKA ÜNIKI Dcpnrtmenl of Biology, Princeton University, Princeton, N . I. ABSTRACT Belgian Congo. In the New World tropics, birds of the family Formicariidae (antbirds) are the Studies of ant-following birds at Belém show that main ant-followers. Skutch (1969) and Slud some species are professional ant-followers, that is, (1960, 1964) note ant-following birds in Costa obtain more than 50% of their food near ants. Pro· fessional antAollowers at Belém are : Black-spotted Rica. Johnson (1954) briefly reports on ant­ Bare-Eye, Phlegopsis nigromaculata, White-backed following birds on Barro Colorado lsland, Pana­ Fire-Eye, Pyriglena leuconota, Scale-backed Antbird, má Canal Zone. In a long-term study, Willis Hylophylax poccilcnota ( Formicariidae); Barred (1967) considers one of the main species in Woodcreeper, Dcndrocohptes certbia, Plain-brown Woodcreepc"", Den.lrocincla fultginosa. Whit.e-chinned detail. He also reports in less detail on other Woodcreeper, D. merula; and Gray-headed Tanager specics and on ant-followers in other Neotropical Eucometis penicillata. A list of thirty-seven species 1egions (Willis, 1960, 1966a, 1966b, 1968a, 196Bb, of nonprofessional birds was obtained during the 1969). Oniki and Willis (in press) discuss ant­ study period Nonprofessional spxies only occasion­ following birds of the Guianas and nearby ally follow a.rmy ant ra1ds. The ~rmy ant (Eciíon llurchelli) wns the main species followed. Foraging regions. habits, perch heights and prey items were noted for Meerwarth ( 1904) reports that in Pará, most of the professional and nonprofessional species Braz i I, Pyriglena atra ( = Pyriglena /euconota) of birds. Also, their interactions at swarms of ants and a compa.rison with other ant-following guilds is c.:ld Ph/ogopsis nigromaculata (= Phlegopsis provided. nigromaculata) follow army ants constantly to feed on insects flushed by the ants. There have been no other studies of ant-following !NTRODUCTION birds in that region. This report discusses observations of ant-following birds at Belém . In tropical forests, as swarming carnivorous army ants approach, small animais run or fly Pará, Brazil. helter-skelter. Birds flock around these swarms to obtain arthropods flushed by the ants. Some STUDY AREAS birds are so persistent at feeding near the ants that they havc been termed "professional" ant­ The study was conducted mostly at the followers, that is, they obi:ain more than 50% Aurá Reserve (Novaes, 1970); occasional obser­ of their food near the ants (Willis, 1967). Other vations were made at the Mocambo Forest species of birds form part of the avian ant· Reserve. These reserves are part of the APEG following groups or • guilds" (Root, 1967) only lArea de Pesquisas Ecológicas do Guamá) of now and then, and constitute the nonpro· the Instituto de Pesquisas e Experimentação fessional" or "amateur" ant-followers. Agropecuárias do Norte (IPEAN) at Belém. Recent studies of ant-following birds show The ave r age rainfall of Belém (altitude 13m: that QJfferent species tollow ant raids in differ­ 1< 28' S lat., 48° 27' W long.) from 1923 to 1963 ent regions. Chapin (1932) reports thrushes as is shown in Figure 1 . The dry season is June the principal birds following driver ants in the to November. (1) - Based on M. S. thests, University of Kansas, U.S.A. -59 titute a lower stratum (20 to 30m). In an even 2837 500 r-- lower stratum plants of the families Rubiaceae, - Solanaceae, Myrtaceae, Sterculiaceae, Carir;a­ E r-- ceae, etc. are dominant (Kuhlman, 1959). How­ E r-- ever, várzea forest is poorer in species than c: t-- 300 terra-firme forest (Ducke and Black, 1954) r-- Capoeira or second-growth forest is charac­ o terized by low vegetation that grows up after ....c: t--t-- o l--r-- the primary or virgin forest is cut. Grasses 0:: ~ 100 and sedges are abundant in the capoeira. o Saplings, bushes and vines make the capoeir2 J F M A M J J, A S O N D very thick and difficult to penetrate. SELEM The Mocambo Forest Reserve is a terra­ FIGURE l - Avcragc montly and yearly raioiall al firme forest of 5. 5 hectares surrounded by Belém in mm. igapó forest and composed of dense and varied vegetation dominated by Ca/ophyllum brasíli• The Aurá Reserve vegetation consists mainly ensis, Macro/obium, acaciefolium, Nectandra of "várzea" or tidal-swamp forest, with a small amazonum, Piranhea trifoliata, Triplaris surina­ plot of "terra-firme" forest o r mature upland mensis, Bombax aquaticum, etc. (Kuhlmann, forest and a strip of "capoeira" or second 1959). The igapó forest is a forest of swampy growth bordering a small plantation of rubber areas permanently covered by water and differs trees. Dr. João Murça Pires, botanist at IPEAN, from the várzea forest by the absence of such has studied in detail the ecology and compo­ indicator plants as Euterpe oleracea. sition of these forests. MATERIAL AND METHODS The terra-firme forest is characterized by tall trees, some of them over 40 m high. They I studted ant-following birds by direct obser­ are slender and branch into a dense canooy. vation, using 7 x 35 binoculars. ldentification The leaves of these trees fali and decompose of birds, using the nomenclature of Meyer de rapidly on the humid surface of the ground. Schauensee ( 1966) was fac i I itated by the use o f Many bromeliads and mosses cover the tree study skins of the Museu Paraense "Emílio trunks. Leguminous trees are the most im­ Goeldi", Belém. portant species in this type of forest. Very Part of the Aurá and ali of the Mocambo often the shallow-rooted trees fali before a reserve are marked in a grid of 10m squares. storm or strong wind, opening clearings in the with numbered tags on stakes at corners and forest; shortly saplings develop there, replacing the old fal len trees. According to Kuhlmann with paths along major grid !ines; this facilitated (1959), Bertho/letia excelsa (castanheira) and rnapping movements of army ants and ant­ Castilloa u/ei (caucho) are characteristic of following birds. terra-firme forest. From April 1966 to April 1968, I followed Várzea or swamp forests occur in areas severa! colonies of the army ant Eciton burche/Ji regularly flooded by tides, which keep the to study the ant-following birds. Observations of ground wet and swampy. The tree species ant-followers were taken mostly in the morning mostly differ from those of the terra-firme when ants and birds were most active; I made forest. Hevea brasiliensis (seringueira) and few observations in the afternoon, from 12:00 Calycophyllum spruceanum (pau mulato) cons· tr 18:00. Ant-following birds were netted in titute the highest trees (40 m) and other genera front of a swarm only after at least two hours such as Sapium, Viro/a, Cordia, lnga, Reedia, of observation. Captured birds were color­ Triplaris, Cecropia, Cassia, Plumería, etc. cons· banded, measured, and weighed. 60- THE ARMY ANTS day when ants are very active and one on a statary day when ants are not swarming from The army ants the "tauócas" o r murupe­ their bivouac at ali. tecall of the Brazilian lndians, are carnivorous A Nomadic pay : I arrive at the bivouac in and hunt through the forest in large swarms, a patch of second-growth forest at 06:50. At called "correições" in Brazil. As a swarm 07:18 a mal e White-backed Fire-Eye (Pyriglena advances over the ground, ants examine logs, leuconota) arrives. Perching on low twigs and holes in the ground, dead fallen leaves, etc. hopping from one twig to another, he investi­ lmportant species of army ants for birds gates the colony. He wanders about for a at Belém are Eciton burchel!i and E. rapax, while as the ants start swarming; as soon as especially the former. According to Schneirla the ants spread out and start flushing insects (1957) and Rettenmeyer (1963), Eciton burchelli he starts dropping down for them has a life cycle with two distinct phases. In the Gradually the number of birds increases, 11 nomadic" phase, the ants raid actively ali day with female and other male Fire-Eyes calling and to provide food for their growing larvae, and sallying to the ground or low branches for prey move the "bivouac" o r temporary nest to over the advancing swarm front. Grunting another site almost every night. The bivouac sounds from Black-spotted Bare-Eyes (Ph/egopsis nigromaculata) indicate that they have arrived movement is usually in the direction of the si lently and are hiding behind low bushes. strongest branch of the swarm during the day. Plain-brown Woodcreepers (Dendrocincla fuligi­ unless ants swarmed in one direction in the nosa) move in and fly from one trunk to another morning and changed to another direction in as they forage. The three main professional the afternoon. The distances between con­ ant-followers are thus the first three species to secutive nomadic bivouacs at Belém averaged arrive this morning. 70.1 m (n = 274). The average length of the A Long-billed Gnatwren (Ramphocaenus nomadic phase was 12.6 days (n = 8). In melanurus), foraging 4 m up, flutters rapidly Panamá the length of the nomadic phase aver­ from one perch to another and calls "tch-tch­ ages 13-14 days (Johnson, 1954; Willis, 1967). tch 11 as it wags its thin, long tail like the In the other o r "statary" phase, the colony pendulum of a clock. Individual Silver-beaked stays in one bivouac for about three weeks. Tanagers (Ramphocelus carbo) wander about. In this phase, the queen lays eggs and the cal ling.
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