TbeCondor94:151-158 0 TheCooper Ornithological Society 1992

FORAGING BEHAVIOR AND DIET OF THE HELMETED ’

MIGUEL ANGELO MARIN? Departamento de Ecologia, Universidadede Brasilia, 70910 Brasilia, DF,

Abstract. I studied the foragingbehavior and diet of the ( galeata) in the gallery forestsin the cerrado (savanna)region of central Brazil. Observations of 21 color-marked were made from April 1988 to March 1989 on a 2.5-ha study plot in a gallery forest. They showed that Helmeted are highly frugivorous (85.7% of foraging observationswere on fruits; all stomach contentsanalyzed had fruits while arthro- pods were present in only 24.5%; and 96.2% of the fecal massescollected under a nest had fruits while 6 1.5% had arthropods);and take fruit mostly (46.5% of the observations)while in flight (sallies).Helmeted Manakins ate fruits of 17 speciesof 12 families of plants, as well as arthropods such as Araneae, Orthoptera, Coleoptera, Diptera, and Hymenoptera. One to six Helmeted Manakins attended 63.6% of the mixed- flocks observed. Green birds (females and immature males) foraged at lower heights (6.5 * 4.6 m) than adult and subadult males, which foraged at a height (8.2 f 4.4 m) closer to where they sang (10.2 + 3.8 m) or called (8.4 f 4.8 m). The Helmeted Manakin showedsimilarities to other manakins in the diet and foraging tactics used, even though it may not be promiscuous(Marini and Cavalcanti, in prep.) and has an unique geographicaldistribution among dichromatic man- akins. Key words: Helmeted Manakin; Antilophia galeata;foraging behavior:diet; frugivory; Pipridae; Brazil.

INTRODUCTION The natural history of the Helmeted Manakin The 52 speciesof Manakins (Pipridae: Sibley and is poorly known. It inhabits mostly dry and Monroe 1990) are mostly frugivorous and pro- flooded gallery forests in the cerrado region miscuous with strong sexual dichromatism. The (Meyer de Schauensee1970; Sick 1985, pers. ob- monotypic Helmeted Manakin (Antilophia gule- serv.), habitats structurally similar to the habitats ata), is also highly dichromatic (see de- of other manakins. Nests and eggswere described scription below), but may form long term pair- by Ihering (1900, 1902) and Marini (1992) ob- bonds, which leads Marini and Cavalcanti (in served its breeding biology. The only dietary de- prep.) to suggestthat it may not be promiscuous. scription is by Schubart et al. (1965) who found In addition to its unusual mating system for the fruit in stomach samples. Becauseof its possibly Pipridae, the Helmeted Manakin also has an unusual mating system, atypical range and poor- atypical geographical distribution in forests in ly known natural history, I studied aspectsof this the cerrado (savanna) ecosystemof central Bra- species’ foraging behavior and diet, and compare zil, between the Amazonian and Atlantic forests them with other manakins. (Pelzeln 187 1, Hellmayr 1929, Laubmann 1940, STUDY AREA AND METHODS Pinto 1944, Pinto and Camargo 1948, Fry 1970, Meyer de Schauensee 1970, Sick 1985, Sibley This study was conducted in the cerrado region and Monroe 1990, Willis and Oniki 1990). Here, of Brazil, in the gallery forest of the Corrego Ca- it is one of the most abundant (pers. petinga (a creek) of the Ecological Station of the observ.). Only two other manakins (the Band- University of Brasilia, Brasilia, Distrito Federal tailed Manakin [Piprufasciicuudu] and the Pale- (15’58’s; 47”56’W). The 2,300-ha station is lo- bellied Manakin [Neopelmu pullescens]) occur at cated within the Agua Limpa Farm and is subject lower densities than the Helmeted Manakin in to man-made and natural fires at irregular inter- these gallery forests (pers. observ.). vals. One large-scalefire burned most of the sta- tion in September 1987, including the understory of the gallery forest studied. Whether the fire I Received 20 May 1991. Accepted 16 October 1991. 2Present address:Department of Ecology, Ethology affected the plant community is unknown. and Evolution, University of Illinois, Champaign, IL The gallery forest of the Corrego Capetinga has 61820. emergent trees up to 20-30 m high, with contin-

[I511 152 MIGUEL ANGELO MARINI

TABLE 1. Per1centage of use of the foragingtactics by marked by a grid of 34 points at 30 m intervals. males (adults ana* SW‘radults) and greens (immature Observations were from April 1988 to March males and fern-a~>).‘--’ sample’ size (n) representsthe number of fora&;ing observations. 1989. Searcheswere made for birds at each point for 5 min, with 2-3 minutes between searching

:ens Total periods. I made approximately 41 visits/point - during the 12-month period of study. I made 450 Sally on fruit 45.5% 47.1% 46.5% (n = 15) (n = 24) (n = 39) manakin sightingsthat lasted from a few seconds Glean on fruit 39.4% 39.2% 39.3% to 5 min. The number of sightings/month at the grid ranged from 17 to 64 (X = 37.5). Sally on arthropod I consider one sighting independent from an- other sighting becauseI sampled an individual’s Glean and screen 0.0% 11.8% 7.1% on arthropod (n = 0) (n = 6) (n = 6) feeding height at the beginning of each obser- vation period, with a mean interval of 7-8 min Total (n = 33) (n = 51) (n = 84) between two consecutive samples. Only the first foraging tactic was recorded per sighting. Also, the height of the first vocalization emitted by a uous canopy between 10 and 15 m high (Ratter being observed was recorded. Foraging tac- 1980). Ratter recorded 120 plant species and tics were classified according to Remsen and found that the most common tree speciesinclude Robinson (1990): glean is “to pick food items Amaioua guianensis,an unidentified Lauraceae, from a nearby substrate, that can be reached Pouteria ramijlora, Salacia elliptica, Guatteia without full extention of legs or neck”; sally is sellowiiana and Pisonia gracilljlora. Detailed de- “to fly from a perch to attack a food item and scriptions of the study area and region are in then return to a perch”; and screen is to attack Ratter (1980), Eiten (1984) and Marini (1989). a food item in continuous flight. Additional ob- The Distrito Federal region has a well-defined servationsof foragingtactics and diet and of some dry season from May to September. Precipita- flocks were conducted in the 2.5-ha study plot or tion during the study period (Instituto Brasileiro on another study site 5 km away in the same de Geografia e Estatistica 1989) was similar but gallery forest. Additional observations of forag- lower than the mean precipitation from 1963 to ing and vocalization heightsalso were conducted 198 1 (Comissao de Planejamento Agricola-Dis- in the 2.5-ha study plot. trito Federal/Funda@o Zoobotanica do Distrito The 38 individually color-marked Helmeted Federal 1984). Manakins on the study plot during the 12-month Birds were mist-netted from September 1986 period of study included four adult males, three to April 1989, on 17 net sites inside the grid, subadult males, one immature male, six females mostly from sunrise to 13:OO.After capture, sex and seven green individuals of unknown sex ob- was determined by plumage (see below) and oc- served subsequently. I classified birds by plum- currence of brood patch. age and I considered them adult males if they Birds were collected from December 1987 to had black and red plumage. This included some April 1989, in six other gallery forests close to individuals with only a few green feathers. Sub- the main study area between 15’45’ and 16”OO’S adult males had green plumage mixed with black and 47”45’ and 48”05’W. Birds were sexed by and red feathers. Green birds (females and im- gonads and plumage, weighed and measured. mature males) had only a few reddish body feath- Their stomachs were fixed in 70% alcohol. The ers. Birds were classified as females if they had stomachcontents were analyzed for type and vol- brood patches. The reproductive season (July- ume of food present. Food was classified as an- December) was defined by gonadal development, imal (arthropods) or plant (fruits, seedsand plant nesting activity, brood patch, and male behavior parts). The relative volume was visually esti- (Marini 1992). No male maintained the green mated by comparing contents of different stom- plumage for two consecutive seasons,but none achs. The largest recorded stomach volume was was banded when in the nest. used as the standard comparison for estimating relative volume in the other stomachs. RESULTS Observations were made primarily in the Foraging behavior.Females and immature males morning (06:30-13:00) on a 2.5-ha study plot (green birds) used five foraging tactics: (1) sally FORAGING OF HELMETED MANAKIN 153

40 A _- - GREEN BIRDS m MALES

1-3 4-6 7-9 IO-12 ) 13 FEEDING HEIGHT (m)

“V

I CALLS (GREEN BIRDS)

50 IS8 CALLS (MALES)

0 SONGS (MALES)

: 40

8 5 30 i?

E 20

1-3 4-6 7-9 10-12 ) 13 VOCALIZATION HEIGHT (m) FIGURE 1. Top (A), percentageof feeding behavior by green birds (females and immature males) (solid bars) (n = 60) and by adult and subadult males (cross-hatchedbars) (n = 47) by height. Bottom (B), percentageof calls by green birds (females and immature males) (solid bars) (n = 24), and calls (cross-hatchedbars) (n = 33) and songs(open gray bars) (n = 6 1) by adult and subadult males by height. on fruit, (2) glean on fruit, (3) sally on , (4) (,qua”‘ notes) at significantly (t = 6.6, P < 0.00 1) glean on insect, (5) screen on insect on a sub- lower mean heights (4.7 f 2.2 m, mean + 1 SD; strate. Adult and subadult males used tactics 1, IZ = 17) than males sang (an eight note piercing 2 and 3. There were no differencesin the use of whistle) (10.2 t- 3.8 m, mean f 1 SD; n = 61) foraging tactics between males (adults and sub- or called (“qua” notes) (t = 3.5, P < 0.001) (8.4 adults) and immature males and females (Table f 4.8 m, mean k 1 SD; n = 33) (Fig. 1B). How- 1) (G = 0.04, df = 2, P > 0.975). Sallies on fruits ever, becausethe foraging heights were not dis- were more common (46.5%) than any other tac- tributed normally (Fig. 1A), comparison of means tic (Table 1). Birds ate more fruit (85.7% of 84 may not be valid and a G-test was used. No observations) than arthropods (14.3%). significant difference (G = 3.75, df = 4, P > 0.1) Green birds fed at significantly (t = 2.02, P < in foragingheights between green birds and males 0.025) lower mean heights (6.5 f 4.6 m, mean was detected. f 1 SD; IZ= 60) than males fed (8.2 f 4.4, mean Mixed-speciesflocks.I observed the Helmeted rf: 1 SD; n = 47) (Fig. 1A). Green birds also called Manakin participating in 14 out of 22 mixed-

FORAGING OF HELMETED MANAIUN 155

TABLE 3. Food types,foraging tactics and flockbehavior of manakin species.

Swcies Food tvw? Foraeinetactic” Autho6 aureola F 1 Pipra fasciicauda F, I P 2 Pipra filicauda F 3 Pipra mentalis F, I P, F (V P 4, 53% 7, 829 Pipra erythrocephala F, I P P 1, 3, 10 Pipra rubrocapilla F, A 11 Pipra chloromeros F, I P 2 Pipra pipra F, I P, F (S) P 1,9, 10, 12 Pipra coronata F, I F (S) P 2,3,9 Pipra serena F, I P P 10,13 Pipra villasboasi F, I 1 Pipra nattereri I 1 Pipra isidorei F 3 Antilophia galeata F, A F 6 Cl, P P 1, 14 linearis F, I D 9, 15, 16, 17, 18 Chiroxiphia lanceolata F I=(S, I-0 7, 9 Chiroxiphia pareola F F (3 3 Chiroxiphia caudata F, I F U-Q,P 19,20 Masius chrysopterus F, I F P 3, 21 Ilicura militaris F 19,22 gutturalis F, I F (W, I ’ P lo,23 Corapipo leucorrhoa F, A F (9 P I, 9, 24, 25 candei F F (3 9 Manacus aurantiacus F, I F 6) 9 Manacus vitellinus F F, P, J 627 Manacus manacus F, I F (S), P 1, 3, 10, 11, 12,26 Machaeropterusregulus F, I F (S) 3, 12 Machaeropterusdeliciosus I, F 3, 27 atronitens F, I 1, 3 flavicapilla 3 flavivertex 3 Heterocercuslinteatus F, 1 chrysocephalum F, I P, F (S) P : 10 Neopelma pallescens I 1’ Tyranneutesstolzmanni I, F P 1,2,28, 29, 30 Tyranneutesvirescens I, F P 10 Pipritesgriseiceps F, I P 9 Pipriteschloris

October 1988, 25 had fruit seeds, 15 also had two Coleoptera in two, one Diptera and one For- remains of arthropods and one had only arthro- micidae, and at least nine unidentified . pods. By far, the two most common fruits were Some clean seeds(apparently regurgitated) also Miconia hirtella with 450 seeds present in 12 found under this nest were Lauraceae (n = 90), fecal massesand Cecropiapachystachia with 760 Annonaceae (n = 31), Myrtaceae (n = 17), So- seedsin 11 fecal masses.Four to 35 seedsof five lanaceae (n = 9), and Virola sp. (n = 2). other specieswere present in five or fewer fecal masses,including two speciesof Solanaceae,and DISCUSSION three unidentified species. Among arthropods, The Helmeted Manakin is highly frugivorous: all there were 42 specimens of Orthoptera in four stomach samples had fruits (Fig. 2), while ar- fecal masses,six specimens of Araneae in four, thropods were present in only 24.5% (Table 2) 156 MIGUEL ANGELO MARINI

TABLE 4. List of families(n = 12) and species(n = is); Foster (1978), however, observed this species 17) of plantsconsumed by the HelmetedManakin. feeding on only three species of plants for five months during one year. Snow (1962) listed 105 Families Speaes speciesof plants in the diet of the White-bearded Annonaceae Guateria sellowiana Manakin (Manacus manacus) with Melasto- Lauraceae indet. tree sp. mataceaeand Rubiaceae,respectively, as the most Melastomataceae Miconia cuspidata,M. hirtella, common families eaten. The White-ruffed Man- M. pseudonervosa akin (Corapipo leucorrhoa) fed on several fruits, Meliaceae Guarea macrophylla Moraceae Cecropiapachystachia especially on Melastomataceae shrubs and trees Nyctaginaceae Guapira sp. (Skutch 1967). The Golden-winged Manakin Ochnaceae Ouratea castaneifolia (Masius chysopterus) was observed feeding on Phytolaccaceae Phytolaccadodecandra 10 speciesof four families of plants (Prum and Piperaceae Piper tectonifolium Rubiaceae Amaioua guianensis,two indet. Johnson 1987). shrubyspp. Arthropods eaten by manakins primarily are Smilacaceae Smilax siringoides flying insects and spiders. They include Orthop- Solanaceae indet. shrubysp. and indet. vine tera, Coleoptera, Diptera, Hymenoptera and SP. Araneae by the Helmeted Manakin, Coleoptera, Diptera and Odonata by the White-bearded Manakin (Snow 1962), Araneae by the White- in which the arthropods never represented more ruffed Manakin (Skutch 1967) and Orthoptera than 20% of the total stomach volume (Fig. 2); and Coleoptera by the Gray-headed Manakin 85.7% of the foraging observations were on fruits ( griseiceps) (Stiles and Skutch 1989). (Table 1); and 96.2% ofthe fecal massescollected Other studies did not specify which group of in- under a nest had fruits while 6 1.5% had arthro- sect was eaten (see Table 3). P.T.Z. Antas (pers. pods. There were only slight differences in for- comm.) observed the Helmeted Manakin eating agingtactics (Table 1) and diet (Tables 1,2) among alate termites in the border of a gallery forest and the sex and age classesanalyzed. The Helmeted Willis (1984) recorded one male following army Manakin’s foraging tactics and the kind of food ants to sally from twigs and leaves for small ar- taken are similar to other manakins (Table 3). thropods in Mato Grosso, Brazil. The high num- All manakin genera are known to eat fruits and ber of orthopterans in four fecal samples and of insects (Table 3). Diptera in one stomach content under the nest Green birds foraged more in the understory of the Helmeted Manakin, may be unusual among and males foraged more in the canopy (Fig. 1A). manakins. This difference may relate to males singing and The participation of the Helmeted Manakin calling significantly higher in the forest than green in mixed-species flocks, as well as its behavior birds (Fig. 1B). Alternatively, males and green of feeding on termites in the border of a gallery birds may be partitioning the fruits available in forest (P.T.Z. Antas, pers. comm.), and following the forest. Differences in foraging height between army ants (Willis 1984), suggeststhat its insect males and females have been shown by several foraging may be opportunistic because of the authors (see Holmes [ 19861 and references). small importance of insects on its diet. Several However, I can not conclude that foraging height other manakin specieshave also been observed in the Helmeted Manakin differs between the attending mixed-species flocks as non-perma- sexesbecause green birds included both females nent members (Table 3). Manakins join mixed- and immature males. For the same reason, I can speciesflocks temporarily, presumably to forage not correlatethe foragingheight of the green birds for insects. with nesting height. In conclusion, the Helmeted Manakin is sim- The Helmeted Manakin undoubtedly eatsfruits ilar to other manakins in relation to foraging of more speciesof plants than the 17 speciesof tactics used and diet, even though it has an atyp- 12 families that I could identify (Table 4). This ical range for a dichromatic manakin. Also, the broad use of plant speciesseems to be common proposed non-promiscuity of the Helmeted in the Pipridae. Wheelwright et al. (1984) re- Manakin (Marini and Cavalcanti, in prep.), as- corded 37 plant speciesof 22 families in the diet sociated with its highly frugivorous behavior, of the Long-tailed Manakin (Chiroxiphia linear- supports Beehler’s (1983) argument that frugiv- FORAGING OF HELMETED MANAKIN 157 ory “cannot, by itself, explain why some birds tivo dos ninhos e ovos das Aves do Brazil. Rev. are polygamous and others monogamous.” Mus. Paul. 4: 19l-300. IHERING,H. VON. 1902. Contribui@es para o co- nhecimentoda Omithologia de S. Paulo. Rev. Mus. ACKNOWLEDGMENTS Paul. 5:261-326. INSTITUTOBRASILEIRO DE GEOGRAFIAE ESTAT~STICA. This researchwas part of a M.Sc.thesiswhich was sup- 1989. Boletim agroclimatol6gico da Reserva nortedbv a BrazilianNational ResearchCouncil (CNPa) Ecol6gicado Roncador, Brasilia. scholarship, as well as grants from CNPq td R. g: LAUBMANN,A. 1940. Die Vogel von Paraguay. II. Cavalcanti and from the University-of Brasilia- Strecker and Schroder, Stuttgart, Germany. (UnB-DPP) to R. B. Cavalcanti and M.A.M. I would LEVEY,D. J. 1988. Spatial and temporal variation in like to thank R. B. Cavalcanti for advice. N. Burley, Costa Rican fruit-eating bird abundance. Ecol. D. Enstrom, S. Robinson, K. Sieving, and E. Willis for Monogr, 58:25l-269. comments on early drafts of the manuscript and dis- MARINI, M. A. 1989. 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