SHORT COMMUNICATIONS
ORNITOLOGIA NEOTROPICAL 19: 143–147, 2008
© The Neotropical Ornithological Society
BEHAVIOR OF THE TAWNY ANTPITTA (GRALLARIA QUITENSIS)
IN NORTHERN ECUADOR
Harold F. Greeney1 & Inka Harms1
1
Yanayacu Biological Station and Center for Creative Studies, c/o Foch 721 y Amazonas,
Quito, Ecuador. E-mail: revmmoss@yahoo.com
Comportamiento de incubación de la Gralaria Leonada (Grallaria quitensis) en el norte del Ecuador.
Key words: Nest, egg, incubation, Ecuador, Andes, paramo, Tawny Antpitta, Grallaria quitensis.
Recent years have witnessed rapid advances in
our knowledge of the basic breeding biology
of Grallaria antpittas (e.g., Freile & Renjifo
2003, Price 2003, Kofoed & Auer 2004, Martin & Dobbs 2004, Greeney et al. 2006).
Except for the Scaled Antpitta (G. guatimalensis) (Dobbs et al. 2001, 2003) and Moustached
Antpitta (G. alleni) (Freile & Renjifo 2003,
Londoño et al. 2004, Greeney & Gelis 2006),
however, we still know few details of such
aspects as incubation period, rhythms, and
behavior at the nest.
The Tawny Antpitta (G. quitensis) ranges
from northern Colombia to northern Peru,
inhabiting mostly elevations between 3000
and 4500 m (Krabbe & Schulenberg 2003). At
these high elevations it inhabits paramo, high
temperate woodlands, and areas dominated
by low scrubby vegetation (Krabbe & Schulenberg 2003). Unlike other antpittas, the
Tawny Antpitta is fairly easy to observe due
to its habit of running and calling in open
areas, yet its nest has only recently been
described (Greeney & Martin 2005) and noth-
ing further has been published on its breeding
behavior. Here we present the first detailed
observations on incubation behavior for the
Tawny Antpitta. In addition, we supplement
and amplify the nesting data previously
reported.
We studied the nesting behavior of Tawny
Antpitta from 26 September to 21 October
2005 in the paramo at the pass (00°22.4’S,
78°08.3’W), west of Papallacta, Napo province, Ecuador. On 26 September at 16:30 h
(EST), we flushed an adult from a nest containing a single egg, which we noted to be
warm but wet and slightly sticky, suggesting it
had just been laid (HFG pers. observ.). At
16:00 h on 28 September, the nest still contained a single egg, cold to the touch. The following morning at 06:00 h, the nest contained
two cold, dew covered eggs, suggesting that
the second had been laid the evening before
and that no adult had spent the night on the
nest. At this time we placed a video camera c.
4 m from the nest and began recording adult
behaviors. We continued to film the nest daily,
143
GREENEY & HARMS
FIG. 1. Incubation rhythms of the Tawny Antpitta at 4000 m. elev., Papallacta Pass, Napo, Ecuador.. The
period from laying of the last egg until hatching of the entire clutch, from 06:00 to 18:00 h, is shown. Periods of attendance are shown in black, absences in white. Stippled areas represent periods not observed.
White X’s designate instances where adults changed places at the nest, leaving the eggs uncovered for < 1
min. The day of incubation is figured on the left with the total daily % coverage shown on the right.
from roughly 06:00 h to 18:00 h, until 20
October, when one egg hatched at 10:15 h,
and the second at 10:45 h, giving an incubation period of roughly 21.5 days. We recorded
and transcribed a total of 252 h of video, capturing 207 h of incubation behavior (adults
present at the nest).
Daily coverage of the eggs ranged from
64 to 97%, except on the first day of incubation, when coverage was only 44% (Fig. 1).
For the entire incubation period, the eggs
were covered for 82% of daylight hours, or
86% if the first 3 days of irregular attendance
144
are excluded. Both adults incubated, with
average periods of attendance of 55 ± 37 min
(n = 209) and periods of absence of 12 ± 21
min (n = 164).
While incubating, adult Tawny Antpittas
spent 3.9% of their time engaged in frequent
(10.7 times per h) bouts of movement, including one or more of the following behaviors.
During 27% of 2158 movement bouts, adults
leaned over and rapidly thrust their bill in and
out of the nest lining at least once, as reported
for this (Greeney & Martin 2005) and other
antpittas (Dobbs et al. 2003, Greeney & Gelis
SHORT COMMUNICATIONS
2005, Greeney & Sornoza 2005). In total, we
observed 634 rapid probing events (3.1 times
per h). While the purpose of this behavior is
still unconfirmed, it is thought to be a means
of parasite removal in other birds (Haftorn
1994, Greeney 2004) and, as adults often
ingested something removed from the nest
after probing, we feel this is likely the case for
Tawny Antpitta. In some instances, rapid
probing appeared to facilitate rolling of the
egg, but we are unsure if this was deliberate.
On 28% of these movement bouts, adults
arranged material on the rim or in the cup of
the nest (3.2 times per h). During 5% of
movement bouts, adults briefly preened their
contour or wing feathers (0.5 times per h).
Other bouts of movement included only
standing and peering into the nest or shuffling
about to change position. While not moving,
adults frequently closed their eyes for brief
periods, apparently dozing. On 28% of adult
arrivals to the nest (n = 229), they brought
with them a pale grass fiber, which they added
to the lining of the nest before settling to
incubate. On one occasion, the arriving adult
passed the material to the second, which
added it before leaving. Also on one occasion,
an adult removed and carried away a stray
feather from the lining of the nest.
Adults frequently sang from the nest on
every day of observation, with the exception
of the first. Generally, singing bouts (interval
between songs > 30 s) began softly and
increased in volume, beginning with a one or
two-note partial song, and ending with a fullvolume three-note song. Bouts of singing
ranged in length from single songs to 4.2 min
(mean ± SD = 1.9 ± 1.3 min). At times, we
recorded adults singing at rates of up to 17
songs per min (mean ± SD = 10 ± 2 songs/
min.). If all adult time at the nest and all songs
are included, adults sang at an overall rate of
4.5 songs/h. In addition to these periods of
singing, on eight occasions adults made 3–7,
single note calls while sitting on the nest, usu-
ally in response to similar calls heard away
from the nest, and presumably made by their
mate. The calls were described by Ridgely &
Greenfield (2001) as a penetrating “keeyurr!,”
and from our experience appear to be given in
alarm.
Conclusions. An incubation period of 21.5 (22
days if laying and hatching times were not
known) is the longest documented for the
family Formicariidae. Holley et al. (2001)
estimate 17–21 days for the Ochre-breasted
Antpitta (Grallaricula flavirostris), Skutch (1996)
reports a 20 day period for Black-faced Antthrush (Formicarius analis), and the Peruvian
Antpitta (Grallaricula peruviana) has a 20 day
incubation period (Greeney et al. 2004). Other
documented periods are 16–17 days for
Rusty-breasted Antpitta (Grallaricula ferrugineipectus) (Schwartz 1957), 17 days for
Variegated Antpitta (Grallaria varia) (Erard
1982), 20 days for Scaled Antpitta (Dobbs et
al. 2003), and 15 days for Rufous-faced Antpitta (Grallaria erythrotis) (Bell & Bruning
1976).
The general patterns of nest attendance
for Tawny Antpittas are similar to those
reported for other antpittas. They show limited and irregular incubation for 1–3 days
after completion of the clutch and, once regular incubation begins, cover the eggs for a
high percentage of the day (Dobbs et al. 2003,
Greeney 2006). Also like other antpittas, and
given that they likely spent the entire night on
the nest, Tawny Antpittas tend to have
their longest period of inattendance around
dawn (Londoño et al. 2004, Greeney 2006).
Average periods of attendance during incubation are similar to those reported for Slatecrowned Antpitta (Grallaricula nana) (Greeney
& Sornoza 2005), higher than those for Peruvian Antpitta (Grallaria peruviana) (Greeney
2006), but considerably lower than the average for Scaled Antpitta (Dobbs et al. 2001,
2003).
145
GREENEY & HARMS
ACKNOWLEDGMENTS
We are very grateful to Luis Martínez and
the park guards of Cayambe-Coca Reserve
for their support and help. John V. and the
late Ruth Ann Moore, along with Matt
Kaplan, supported this work through the
Population Biology Foundation. This study
was also funded in part by a Rufford Award
and a Pamela & Alexander F. Skutch
Award. Field Guides, the PBNHS, the Humbolt Crew, and the Hertzberg Family Foundation continue to logistically support the work
of HFG. This is publication number 100 of
the Yanayacu Natural History Research
Group.
REFERENCES
Bell, J., & D. Bruning. 1976. Hatching and handrearing the Rufous-faced Antpitta. Notes on
antbirds kept at the New York Zoological Park.
Avicult. Mag. 82: 118–122.
Dobbs, R. C., P. R. Martin, C. Batista, H. Montag,
& H. F. Greeney. 2003. Notes on egg laying,
incubation and nestling care in Scaled Antpitta
Grallaria guatimalensis. Cotinga 19: 65–70.
Dobbs, R. C., P. R. Martin, & M. J. Kuehn. 2001.
On the nest, eggs, nestlings and parental care in
the Scaled Antpitta (Grallaria guatimalensis).
Ornitol. Neotrop. 12: 225–233.
Erard, C. 1982. Le nid et la ponte de Lipaugus vociferans, Cotingidé, et de Grallaria varia, Formicariidé. Alauda 50: 311–313.
Freile, J. F., & L. M. Renjifo. 2003. First nesting
records of the Moustached Antpitta (Grallaria
alleni). Wilson Bull. 115: 11–15.
Greeney, H. F. 2004. Breeding behavior of the
Bicolored Antvireo (Dysithamnus occidentalis).
Ornitol. Neotrop. 15: 349–356.
Greeney, H. F. 2006. Incubation behavior of the
Peruvian Antpitta (Grallaricula peruviana). Ornitol. Neotrop. 17: 461–466.
Greeney, H. F., & R. A. Gelis. 2005. A nest of the
Rufous Antpitta Grallaria rufula depredated by a
Turqoise Jay Cyanolyca turcosa. Cotinga 24: 110–
111.
146
Greeney, H. F., & R. A. Gelis. 2006. Observations
on parental care of the Moustached Antpitta
(Grallaria alleni) in northwestern Ecuador.
Ornitol. Neotrop. 17: 313–316.
Greeney, H. F., & P. R. Martin. 2005. High in the
Ecuadorian Andes: the nest and eggs of the
Tawny Antpitta (Grallaria quitensis). Ornitol.
Neotrop. 16: 567–571.
Greeney, H. F., & F. Sornoza. 2005. The nest and
egg of the Slate-crowned Antpitta (Grallaricula
nana), with observations on incubation behavior in southern Ecuador. Ornitol. Neotrop. 16:
137–140.
Greeney, H. F., R. A. Gelis, E. C. Hannelly & P. J.
DeVries. 2004. The egg and incubation period
of the Peruvian Antpitta (Grallaricula peruviana).
Ornitol. Neotrop. 15: 403–406.
Greeney, H. F., R. A. Gelis, C. Dingle, F. J. Vaca B.,
N. Krabbe, & M. Tidwell. 2006. The nest and
eggs of the Plain-backed Antpitta (Grallaria
haplonota) in eastern Ecuador. Ornitol.
Neotrop. 17: 601–604.
Haftorn, S. 1994. The act of tremble-thrusting in
tit nests, performance and possible functions.
Fauna Norv. Ser. C Cinclus 17: 55–74.
Holley, D. R., C. A. Lindell, M. A. Roberts, & K.
Biancucci. 2001. First description of the nest,
nest site, and eggs of the Ochre-breasted Antpitta. Wilson Bull. 113: 435–438.
Krabbe, N., & T. S. Schulenberg. 2003. Family Formicariidae (Ground antbirds). Pp. 682–731 in
del Hoyo, J., A. Elliot, & D. A. Christie (eds.).
Handbook of the birds of the world. Volume 8:
Broadbills to tapaculos. Lynx Edicions, Barcelona, Spain.
Kofoed, E. M., & S. K. Auer. 2004. First description of the nest, eggs, young, and breeding
behavior of the Great Antpitta (Grallaria
excelsa). Wilson Bull. 116: 105–108.
Londoño, G. A., C. A. Saavedra-R., D. Osorio, & J.
Martínez. 2004. Notas sobre la anidación del
Tororoi Bigotudo (Grallaria alleni) en la Cordillera Central de Colombia. Ornitol. Colomb.
2: 19–24.
Martin, P. R., & R. C. Dobbs. 2004. Description of
the nest, egg, and nestling of Watkins’s Antpitta
Grallaria watkinsi. Cotinga 21: 35–37.
Price, E. R. 2003. First description of the nest,
eggs, hatchlings, and incubation behavior of
SHORT COMMUNICATIONS
the White-bellied Antpitta (Grallaria hypoleuca).
Ornitol. Neotrop. 14: 1–5.
Ridgely, R. S., & P. J. Greenfield. 2001. The birds of
Ecuador. Cornell Univ. Press, Ithaca, New
York. Schwartz, P. 1957. Observaciones sobre
Grallaricula ferrugineipectus. Bol. Soc. Venez.
Cienc. Nat. 18: 42–62.
Skutch, A. F. 1996. Antbirds & ovenbirds. Univ.
Texas Press, Austin, Texas.
Accepted 22 November 2007.
147
View publication stats