ORNITOLOGIA NEOTROPICAL 23: 367–374, 2012 © The Neotropical Ornithological Society

THE NEST, EGG, AND NESTLING OF THE STRIPE-HEADED ( ANDICOLUS) IN SOUTHERN

Harold F. Greeney

Yanayacu Biological Station & Center of Creative Studies, Cosanga, c/o Foch 721 y Amazonas, Quito, . E-mail: [email protected]

Resumen. – El nido, huevo, y pichón del Tororoí Andino (Grallaria andicolus) en el sur de Peru. – Describo por la primera vez el nido, huevo, y pichón del Tororoí Andino (Grallaria andicolus), una espe- cie que habita áreas alto Andinas desde el centro de Peru hasta el oeste de . En Noviembre de 2011 encontré ocho nidos, durante dos visitas a Abra Malaga, Región de Cuzco, Peru, 3300–4400 m s.n.m.. Los nidos eran grandes copas de musgos y palitos, con unas pocas pajas y raíces finas en la cámara interior. Parece que sitios de anidación pueden ser utilizados por varios intentos durante varios años con la construcción completa o parcial de nuevos nidos sobre los viejos. Fueron ubicados 1–1.8 m sobre el suelo en arboles de , en áreas de bosque tal como áreas activamente utilizados para la ganadería. Un nido contenía dos huevos frescos, de forma sub-elípticas y de color azul claro con pequeñas manchas cafés. Pichones poco desarrollados tenían piel de color rosada, con plumones grises y con el interior de la boca anaranjado y las comisuras de color amarillo cremoso. Ambos adultos alimentaban a los pichones. Según el estado de los nidos que encontré, parece que el periodo de repro- ducción para el Tororoí Andino en esta área comienza alrededor de los fines de agosto o principios de septiembre.

Abstract. – I describe, for the first time, the nest, egg, and nestling of the Stripe-headed Antpitta (Gral- laria andicolus), an inhabitant of the high from central Peru to western Bolivia. In November 2011, I found eight nests during two visits to Abra Malaga, Cuzco Region, Peru, 3300–4400 m a.s.l.. Nests were bulky, open cups of moss and sticks with an internal cup sparsely lined with pale fibers and rootlets. Nesting sites appear to be reused for multiple nesting attempts across multiple breeding seasons, with the complete or partial construction of new nests on top of old ones. Nests were placed 1–1.8 m above the ground in Polylepis trees, in well-forested areas as well as actively grazed . One nest con- tained a fresh clutch of two, pale-blue, subelliptical eggs with sparse brown spotting. Young nestlings had pink skin, sparse grayish natal down, bright orange mouth linings, and creamy-yellow rictal flanges. Both adults brought food to nestlings. Based on the condition of the nests I found, the reproductive period of the Stripe-headed Antpitta in this area likely begins in late August or early September. Accepted 27 August 2012.

Key words: Andes, egg, fledgling, Grallaria andicolus, nest, nestling, Polylepis, Stripe-headed Antpitta.

INTRODUCTION (Schäfer 2002, Greeney et al. 2006), one was (and still is) known only from captivity (Bell Only a decade ago, the nests of only seven of & Bruning 1976), and the nests of three other the 32 of Grallaria had been species were known only from brief notes described (Krabbe & Schulenberg 2003). Of (Sclater & Salvin 1879, Wiedenfeld 1982, de these seven described nests, the description Fabrègues 1991, Whitney 1992). Additionally, for one species proved to be unreliable at the time of Krabbe & Schulenberg’s (2003)

367 GREENEY review, the nests of only two species had been ing biology of the Stripe-headed Antpitta is described by several observers (Edwards & unknown. Recently, while high in the Peru- Lea 1955, Miller 1963, Rowley 1966, Érard vian Andes, I was provided with the opportu- 1982, Donahue 1985, Quintela 1987, Proto- nity to add this species to the growing list of mastro 2000, Dobbs et al. 2001). Our knowl- Grallaria antpittas with described nests. edge of this secretive and poorly studied genus changed drastically near the turn of the METHODS century, led by the first detailed study of Gral- laria reproductive habits (Dobbs et al. 2001), On 12 November 2001, I searched for nests based on observations of the Scaled Antpitta of the Stripe-headed Antpitta at Abra Málaga, (G. guatimalensis) in southern Ecuador. This Cuzco Region, Peru (13°08’S, 72°19’W). I work was quickly followed by papers from searched for nests on the western side of the these and other authors, adding nest descrip- pass, at elevations of 4000–4200 m a.s.l., tions for four species (Freile & Renjifo 2003, where habitat consisted of rocky grasslands Price 2003, Koefed & Auer 2004, Martin & interspersed with small patches of Polylepis- Dobbs 2004), and providing detailed natural dominated with an estimated maxi- history information for two (Dobbs et al. mum canopy height of 4–6 m. patches 2003, Londoño et al. 2004). This trend was were predominantly restricted to sheltered continued (Greeney & Gelis 2005, 2006; valleys and depressions. I located nests by fol- Greeney & Martin 2005, Greeney et al. 2006, lowing adults carrying nesting material or Martin & Greeney 2006, Greeney & Harms food, and by systematically searching patches 2008), and only five years after Krabbe & of Polylepis forest in the vicinity of visually Schulenberg’s (2003) publication, our under- detected adults. I returned and searched the standing of the nesting biology of Grallaria same area on 15 November. I present all lin- antpittas was well on its way to being demysti- ear measurements of eggs, nest dimensions, fied (Greeney et al. 2008). Enthusiasm for the and nest heights with ± SD. study of antpitta reproductive biology contin- ued (Greeney et al. 2009, Solano-Ugalde et al. RESULTS 2009, Greeney & Juiña 2010, Juiña et al. 2009, Greeney & Juiña 2011) and, to date, the nests Nests. I found eight nests, all of which were are known for more than half of the built in Polylepis trees. Six nests were sup- described species of Grallaria. ported by 4–6 small branches (2–3 cm diame- The Stripe-headed Antpitta (Grallaria andi- ter), and the remaining two were each colus) ranges from central Peru southward to supported by two larger (4–6 cm) horizontal western Bolivia (Krabbe & Schulenberg branches. One nest contained two young 2003). It inhabits low-canopied forests domi- nestlings (estimated 5–7 days old, Fig. 1), one nated by Polylepis (Rosaceae) and Gynoxys contained two eggs (undeveloped, Fig. 2), one (Asteraceae) trees, as well as adjacent boulder- was under construction (estimated 1/3 com- strewn grasslands at elevations from 3000 to pleted), and five were not currently in use. Of 4300 m (Schulenberg et al. 2007), and may be the inactive nests, one contained an addled locally common within some portions of its egg and one was only recently abandoned, range (Krabbe & Schulenberg 2003, Schulen- with fresh feces and several pieces of earth- berg et al. 2007). With the exception of a cur- worm still on the rim. Mean nest height was sory description of several nests found by Jon 1.6 ± 0.6 m (range = 1.0–2.8 m), while mean Fjeldså in Peru (Greeney et al. 2008), the nest- height of supporting trees was 2.8 ± 0.7 m

368 NESTING OF GRALLARIA ANDICOLUS

FIG. 1. Adult Stripe-headed Antpitta (Grallaria andicolus) provisioning two nestlings, 15 November 2011, Abra Malaga, Cuzco, Peru (photograph: H. F. Greeney).

(range = 2.0–4.0 m). Nests were bulky, open inner walls of the cup were partially decom- cups, composed primarily of moss, with only posed (although the sparse lining was fresh). sparse inner linings consisting of grass fibers The nest with eggs was composed of some and thin, flexible rootlets. In one nest (Fig. 2), freshly placed moss, but this had obviously the lining included pale, dried grasses, as well been added to a pre-existing nest, suggesting as fresh, green grass blades and straw. All that old nests are refurbished and reused, pos- nests also had short (5–15 cm) sticks incorpo- sibly for multiple years as I have observed rated into the outer portions of the mossy with the Tawny Antpitta (G. quitensis) in Ecua- cup. The numbers of sticks varied between dor (unpubl. data). The nest with nestlings nests, from 10–20 sticks mixed in with the appeared freshly constructed and the nest moss, to 30+ sticks forming a sparse platform under construction was not built on an old below the cup and (in two nests) a circle of nest. The rest of the inactive nests appeared sticks placed around the outer rim of the cup. to have been built at least one breeding season The recently abandoned nest had obviously previously, perhaps more. Based on compari- been originally built some time ago, presum- sons with an active nest of the Chiguanco ably at least a year prior to its most recent use, (Turdus chiguanco) at this site, which as most of the external moss was firmly contained one addled egg and one ca. 3-4 day- attached to the supporting branches and the old nestling on 15 November, and based on

369 GREENEY

FIG. 2. Clutch of the Stripe-headed Antpitta (Grallaria andicolus) provisioning two nestlings, 15 November 2011, Abra Malaga, Cuzco, Peru (photograph: H. F. Greeney). extensive personal experience with nests of completed nest, but dropped the stick while the Great Thrush (Turdus fuscater) in Ecuador, still below the nest, presumably because it I feel confident that the inactive nests I found detected my presence. Based on my examina- belonged to Stripe-headed Antpittas. Never- tion of this nest, I surmise that nests are built theless, the following measurements (cm) per- by placing a variable number of sticks on the tain only to the two active, completed antpitta supporting branches to form a loose plat- nests: maximum outer diameter, 24 and 24; form, upon which the mossy cup is built. It minimum outer diameter, 22 and 22; outer seems likely that additional sticks are added height, 13 and 15; inner diameter, 9.5 and 12; near the end of construction to create the inner depth, 6.5 and 8.5. indistinct ring around the rim of the inner On 15 November, an adult antpitta car- cup. The presence of sticks incorporated into ried a stick in the direction of the partially the mossy walls of nests likely represents

370 NESTING OF GRALLARIA ANDICOLUS sticks from the nest-rim which were added respectively. Contour feathers on all tracts had during previous occupancies and subse- broken their sheaths, with those on all upper quently covered with fresh moss. tracts and the ventral sterna tracts being tawny-buff with small black tips, and those on Eggs. All three eggs (including the addled one) the ventral abdominal tracts entirely buffy- were sub-elliptical and pale blue. The two cream colored. Primary feathers were 3-4 mm eggs in the active nest were sparsely blotched long and had not broken their sheaths, while with brown, with markings only slightly more secondary pin feathers were 5-7 mm long and concentrated on the larger end (Fig. 2). The just beginning to break their sheaths. Tail addled egg was too heavily stained to deter- feather development was still not visible. On mine if it bore markings. Mean linear mea- both visits, nestling bills were orange, with surements of all three eggs were 29.9 ± 0.5 by cream-colored rictal flanges and bright orange 23.3 ± 0.5 mm, and the two viable eggs both mouth linings. Egg teeth were still evident on weighed 8.3 g on 15 November. the second visit, and leg coloration had dark- ened only slightly. Nestlings. On 12 November, at 14:00 h (EST), the two nestlings weighed 17.0 and 16.5 g, and Adult behavior. All adults that I encountered left tarsal measurements were 18.4 and 18.1 away from nests were extremely wary, quickly mm, respectively. They were pink-skinned and disappearing from view when I approached to had sparse, dark grey or black natal down on within 10–15 m, making it difficult to follow the capital, spinal cervical, spinal dorsal, fem- them through the dense Polylepis. On two oral, and humeral feather tracts (all dorsal). occasions, however, while sitting quietly after Ventrally they lacked natal down except for a following adults into the forest, an adult reap- few sparse plumes on the sub-malar and ven- peared within 3–6 m from where I sat, period- tral cervical tracts. Contour pin-feather devel- ically hopping out of view and reappearing opment was most advanced on the spinal from a different direction, seemingly less shy dorsal, femoral, and humeral tracts, with pins once I stopped walking. At the nest under in these areas unbroken but emerged from the construction I was unable to determine how skin 1-2 mm. These pin feathers were pale, many adults participated in building, but at rusty-brown near the base and black apically. the nest with young two adults fed the nest- Contour pin feathers had emerged only < 1 lings, one of which was distinguished by an mm on the capital, spinal cervical, ventral cer- engorged tick attached to its fore-crown (Fig. vical, ventral sternal, and ventral abdominal 1). During my first attempt to photograph tracts. These pins were dark, except for those adults provisioning nestlings (12 November), of the lower ventral sterna and ventral I set my camera tripod only 5 m from the abdominal tracts which were pale yellow- nest, with no attempt at concealment. The white. The only contour pinfeathers to have first adult to arrive, upon detecting my pres- broken their sheaths where those of the lower ence while it was still several meters from the ventral abdominal tracts, which had < 1 mm nest, quickly ran into dense cover. Only 5 min of exposed white feather plumes. Primary later, an adult fed the nestlings and then set- flight feathers were just emerging from the tled to brood while clearly aware of my pres- skin and secondaries were broken through the ence. It left the nest only when I moved to skin ca. 2–3 mm. On 15 November, at 09:15 within 2 m of the nest. On 15 November, h, the nestlings weighed 24.3 and 24.7 g, with while I photographed the nest from only 3 m tarsal measurements of 23.4 and 22.9 mm, away, I detected no reluctance to approach the

371 GREENEY nest. After four adult visits an adult settled to haplonota) Antpittas in seeming to prefer loca- brood, flushing only when I approached to tions with multiple, small supports, as within 1 m. Of the five feedings I observed, opposed to the more firmly supported sites three were by the adult with the tick, which chosen by other species (e.g., tree trunks, was also the individual which brooded on stumps, banks). Sample sizes are still very low both occasions. for most species, however, and some species Adults approached the nest by running show variation in nest placement (Greeney et along the ground, hopping up to several al. 2008). The nestlings of Stripe-headed Ant- perches below the nest before arriving at the pittas are also similar to those described for rim from the same direction on all five visits. other Grallaria, being most similar to those of On all but one visit, the adults brought multi- the Tawny Antpitta, both of which also share ple prey items but fed to only one of the nest- similarities in habitat, nest placement, and egg lings. Food items were predominantly earth- coloration (Krabbe & Schulenberg 2003, worms, with a few larval insects similar to Greeney & Martin 2005, Greeney & Harms crane fly larvae (Tipulidae). Adult departures 2008). from the nest, both natural and when flushed, The secretive and elusive nature of Gral- were quick and silent and accomplished by laria antpittas has historically retarded our dropping straight to the ground below the understanding of their behavior, ecology, and nest and running rapidly away. During a total distribution. Numerous works by an expand- of 50 min spent at the nest with nestlings, as ing number of talented field workers have well as during 15 minutes at the nest with changed this trend during the past two eggs and 30 min at the nest under construc- decades, with four new species described tion, I heard no adult vocalizations. (Stiles 1992, Kattan & Beltrán 1997, Krabbe et al. 1999, Carantón-Ayala & Certuche- DISCUSSION Cubillos 2010) and novel information pub- lished on the biology, , distribution, The bulky, deep, open cup nests of Stripe- and conservation of many species (Krabbe & headed Antpittas are as described for other Coopmans 2000, Greeney et al. 2008, Salaman Grallaria (see references in introduction). Sim- et al. 2009, Freile et al. 2010). I sincerely hope ilarly, egg form and ground color are a match this trend continues, and encourage others with congeners. With regards to egg mark- to use this new body of information to docu- ings, only the eggs of Tawny Antpitta ment the life histories of the 15 Grallaria (Greeney & Martin 2005, Greeney & Harms species which have no published account of 2008), (G. squamigera; their breeding biology. Greeney & Juiña 2011), White-bellied Ant- pitta (G. hypoleuca; Price 2003), and Jocotoco ACKNOWLEDGMENTS Antpitta (G. ridgelyi; unpubl. data) are known to have sparse, dark flecking. This is variable Field work for this study was made possible in the Tawny Antpitta, however, and other by a travel grant from the Smithsonian Insti- species may prove to show similar variation tution and a Rubenstein Fellowship through once larger sample sizes have been examined. the Encyclopedia of Life. John V. and the late With regards to nest placement, Stripe- Ruth Ann Moore, as well as Matt Kaplan, and headed Antpittas join Tawny, Chestnut-naped Field Guides Inc. have generously supported (G. nuchalis), Chestnut-crowned (G. ruficapilla), my ornithological studies through donations Watkins’s (G. watkinsi), and Plain-backed (G. to the Population Biology Foundation. I thank

372 NESTING OF GRALLARIA ANDICOLUS

Glenn Seeholzer and members of the PBNHS Turqoise Jay Cyanolyca turcosa. Cotinga 24: 110– for help with field work, and Neander He- 111. ming and two anonymous reviewers for criti- Greeney, H. F., & R. A. Gelis. 2006. Observations cal comments which improved this paper. on parental care of the Moustached Antpitta (Grallaria alleni) in northwestern Ecuador. Orni- tol. Neotrop. 17: 313–316. REFERENCES Greeney, H. F., R. A. Gelis, C. Dingle, F. J. Vaca B., N. Krabbe, & M. Tidwell. 2006. The nest and Bell, J., & D. Bruning. 1976. Hatching and hand eggs of the Plain-backed Antpitta (Grallaria ha- rearing the Rufous-faced Antpitta. Notes on plonota) in eastern Ecuador. Ornitol. Neotrop. antbirds kept at the New York Zoological Park. 17: 601–604. Avic. Mag. 82: 119–122. Greeney, H. F., & I. Harms. 2008. Behavior of the Carantón-Ayala, D., & K. Certuche-Cubillos. 2010. Tawny Antpitta (Grallaria quitensis) in northern A new species of antpitta (Grallariidae: Gral- Ecuador. Ornitol. Neotrop. 19: 143–147. laria) from the northern sector of the western Greeney, H. F., & M. E. Juiña J. 2010. First descrip- Andes of . Ornitol. Colombiana 9: tion of the nest of Jocotoco Antpitta (Grallaria 56–70. ridgelyi). Wilson J. Ornithol. 122: 392–395. de Fabrègues, F. P. 1991. Die Einfarb-Ameisenpitta Greeney, H. F., & M. E. Juiña J. 2011. First descrip- Grallaria rufula. Trochilus 12: 67–69. tion of the nest of Undulated Antpitta Grallaria Dobbs, R. C., P. R. Martin, C. Batista, H. Montag, squamigera from south-west Ecuador. Bull. Br. & H. F. Greeney. 2003. Notes on egg-laying, Ornithol. Club 131: 67–69. incubation, and nestling care in the Scaled Ant- Greeney, H. F., M. E. Juiña J., S. M. I. Lliquin, & J. pitta Grallaria guatimalensis. Cotinga 19: 65–70. A. Lyons. 2009. First nest description of the Dobbs, R. C., P. R. Martin, & M. J. Kuehn. 2001. Yellow-breasted Antpitta Grallaria flavotincta in On the nest, eggs, nestlings, and parental care northwest Ecuador. Bull. Br. Ornithol. Club of the Scaled Antpitta (Grallaria guatimalensis). 129: 256–258. Ornitol. Neotrop. 12: 225–233. Greeney, H. F., & P. R. Martin. 2005. High in the Donahue, P. K. 1985. Notes on some little known Ecuadorian Andes: the nest and eggs of the or previously unrecorded of Suriname. Tawny Antpitta (Grallaria quitensis). Ornitol. Am. Birds 39: 229–230. Neotrop. 16: 567–571. Edwards, E. P., & R. B. Lea. 1955. Birds of the Greeney, H. F., E. T. Miller, & R. A. Gelis. 2009. Monserrate area, Chiapas, . Condor 57: Observations on parental care and fledging of 31–54. Watkins’s Antpitta (Grallaria watkinsi). Ornitol. Érard, C. 1982. Le nid et la pointe de Lipaugus voci- Neotrop. 20: 619–622. ferans, Cotingidé, et de Grallaria varia, Formica- Juiña J., M. E., J. B. C. Harris, & H. F. Greeney. ridé. Alauda 50: 311–313. 2009. Description of the nest and parental care Freile, J. F., & L. M. Renjifo. 2003. First nesting of the Chestnut-naped Antpitta (Grallaria nucha- records of the endangered Moustached Ant- lis) from southern Ecuador. Ornitol. Neotrop. pitta, Grallaria alleni. Wilson Bull. 115: 11–15. 20: 305–310. Freile, J. F., J. L. Parra, & C. H. Graham. 2010. Dis- Kattan, G. H., & J. W. Beltrán. 1997. Rediscovery tribution and conservation of Grallaria and and status of the Brown-banded Antpitta Gral- Grallaricula antpittas (Grallariidae) in Ecuador. laria milleri in the central Andes of Colombia. Conserv. Int. 20: 410–431. Bird Conserv. Int. 7: 367–371. Greeney, H. F., R. C. Dobbs, P. R. Martin, & R. A. Kofoed, E. M., & S. K. Auer. 2004. First descrip- Gelis. 2008. The breeding biology of Grallaria tion of the nest, eggs, young, and breeding and Grallaricula antpittas. J. Field Ornithol. 79: behavior of the Great Antpitta (Grallaria 113–129. excelsa). Wilson Bull. 116: 105–108. Greeney, H. F., & R. A. Gelis. 2005. A nest of the Krabbe, N., D. J. Agro, N. H. Rice, M. Jacome, L. Rufous Antpitta Grallaria rufula depredated by a Navarrete, & F. Sornoza M. 1999. A new spe-

373 GREENEY

cies of antpitta (Formicariidae: Grallaria) from Quintela, C. E. 1987. First report of the nest and the southern Ecuadorian Andes. Auk 116: 882– young of the Variegated Antpitta (Grallaria 890. varia). Wilson Bull. 99: 499–500. Krabbe, N., & P. Coopmans. 2000. Rediscovery of Rowley, J. S. 1966. Breeding records of birds of the Grallaria alleni (Formicariidae) with notes on its Sierra Madre del Sur, Oaxaca, Mexico. Proc. range, song and identification. Ibis 142: 183– West. Found. Vertebr. Zool. 1: 107–204. 187. Salaman, P., T. M. Donegan, & R. Prys-Jones. 2009. Krabbe, N., & Schulenberg, T. S. 2003. Family For- A new subspecies of Brown-banded Antpitta micariidae (ground-antbirds). Pp. 682–731 in Grallaria milleri from Antioquia, Colombia. Bull. del Hoyo, J., A. Elliott, & D. A. Christie (eds). Br. Ornithol. Club 129: 5–16. Handbook of the birds of the world. Volume 8: Schäfer, E. 2004. Die Vogelwelt Venezuelas und Broadbills to tapaculos. Lynx Edicions, Barce- ihre ökologischen Bedingungen. Band 3. Wir- lona, Spain. temberg-Verlag Lang-Jeutter & Jeutter, Berglen, Londoño, G. A., C. A. Saavedra R., D. Osorio, & Germany. J. Martínez. 2004. Notas sobre la anidación Schulenberg, T. S., D. F. Stotz, D. F. Lane, J. P. del Tororoi Bigotudo (Grallaria alleni) en la Cor- O’Neill, & T. A. Parker, III. 2007. Birds of dillera Central de Colombia. Ornitol. Colombi- Peru. Princeton Univ. Press, Princeton, New ana 2: 19–24. Jersey, USA. Martin, P. R., & Dobbs, R. C. 2004. Description of Sclater, P. L., & O. Salvin. 1879. On the birds col- the nest, egg and nestling of Watkin’s Antpitta lected by the late Mr. T. K. Salmon in the State Grallaria watkinsi. Cotinga 21: 35–37. of Antioquia, United States of Colombia. Proc. Martin, P. R., & Greeney, H. F. 2006. Description Zool. Soc. Lond. 1879: 486–550. of the nest, eggs, and nestling period of the Solano-Ugalde, A., A. Paz, & W. Paz. 2009. First Chestnut-crowned Antpitta Grallaria ruficapilla description of the nest, nest site, eggs, and from the eastern Ecuadorian Andes. Cotinga young of the (Grallaria gigantea). 25: 47–49. Ornitol. Neotrop. 20: 633–638. Miller, A. H. 1963. Seasonal activity and ecology of Stiles, F. G. 1992. A new species of antpitta (Formi- the avifauna of an American equatorial cloud cariidae: Grallaria) from the eastern Andes of forest. Univ. Calif. Publ. Zool. 66: 1–78. Colombia. Wilson Bull. 104: 389–399. Price, E. R. 2003. First description of the nest, Whitney, B. M. 1992. A nest and egg of the Rufous eggs, hatchlings, and incubation behavior of Antpitta in Ecuador. Wilson Bull. 104: 759– the White-bellied Antpitta (Grallaria hypoleuca). 760. Ornitol. Neotrop. 14: 535–539. Wiedenfeld, D. A. 1982. A nest of the Pale-billed Protomastro, J. J. 2000. Notes on nesting of Varie- Antpitta (Grallaria carrikeri) with comparative gated Antpitta Grallaria varia. Cotinga 14: 39– remarks on antpitta nests. Wilson Bull. 94: 41. 580–582.

374