SHORT CO,MMUNICATIONS

Wilson Bull., 11 l(3), 1999, pp. 437-439

Cooperative Foraging in the Mountain Caracara in Peru
Jason Jones ,2
437
ABSTRACT.-Cooperative foraging behavior is
rarely observed in ground-walking birds. I report on
observations of cooperative foraging behavior by
Mountain Caracaras (Phalcoboenus megalopterus) in
the puna region of Peru in September of 1995. On
several occasions, three individuals (two adults and
one immature) were observed working together to turn
over large rocks to obtain prey from beneath. These
cooperative foraging events are notable in that, unlike
cooperative foraging behavior observed in other
ground-walking birds, only one individual obtained
prey from a given cooperative effort. The presence of
the immature individual may be indicative of delayed
dispersal, a behavior not previously described for this
poorly known species. Received 14 Dec. 1998, ac-
cepted 28 March 1999.
Cooperative foraging involves two or more
individual organisms assisting one another in
obtaining a food item. Among vertebrates,
this behavior is well described in many social
mammals (Macdonald 1983, Serfass 1995)
and some fish (Dugatkin and Mesterton-Gib-
bons 1996) but is relatively uncommon in
birds (Sullivan 1984). Among birds, cooper-
ative hunting appears most frequently in sea-
birds (Parasitic Jaegers, Stercorarius parusi-
ticus, Pruett-Jones 1980; Brown and Ameri-
can White pelicans, Pelecanus occidentalis
and P. erythrorhynchos, J. Jones, unpub.
data). Examples from land birds include the
cooperative hunting behavior exhibited by
Harris Hawks (Parubuteo unicinctus; Mader
1979, Bednarz 1988), Golden Eagles (A&la
chrysaetos; Collopy 1983) and Crested Ca-
racaras (Caracara plancus; Morrison 1996).
Most of these instances of cooperative hunt-
ing involve highly mobile prey items; coop-
erative foraging for less mobile organisms is
uncommon (Sullivan 1984). In this report, I
4974 Lakeshore Rd., Kelowna, British Columbia,
Canada, VlY 7R3.
2 Present address: Dept. of Biology, Queen s Univ.,
Kingston, Ontario, Canada, K7L 3N6;
E-mail: [email protected]
detail observations of cooperative foraging
behavior in the Mountain Caracara (Phalco-
boenus megalopterus). This species is adept
at ground foraging and in non-urban areas
feeds on large arthropods, rodents, and birds
(Brown and Amadon 1968). Breeding usually
occurs between October and December with
two, rarely three, eggs laid (Brown and Ama-
don 1968).
STUDY AREA
The observations were made in the Peruvian puna
zone on the road between the towns of Quillabamba
and Ollantaytambo, Department of Cuzco (13 9 S,
72 14 W, 3750 m elevation). This region is charac-
terized by dry grasslands, dominated by genera such
as Calamagrostis and Fesfuca with interspersed shrubs
of the genera Astragalus, Berberis, and Lupinus (Park-
er et al. 1982). Mountain Caracaras are common in this
region (Parker et al. 1982) and are often found near
towns where they feed on refuse and carrion (White
and Boyce 1987).
RESULTS
I observed cooperative rock-turning on four
occasions from l-6 September 1995, as I
watched three individuals (two adults and one
immature) foraging together on the puna
grassland. The immature bird was easily dis-
tinguished by its plumage. On each occasion,
one of the adults approached a large rock,
walked around it, uttered a high-pitched
kieeer, and then stood by the rock. Apparently
responding to the vocalization, the other in-
dividuals joined the first at the rock and pro-
ceeded to work together to flip the rock from
its resting place, with each bird using one of
its talons. The bird that made the call partic-
ipated in the the turning but also appeared to
act as a watcher and was the individual re-
sponsible for prey capture. On one occasion,
the item was captured by an adult which then
gave it to the immature bird. No begging vo-
calizations were uttered by the younger bird
nor did it adopt any unusual posture. Each lift-
ing event took approximately 30 min from call
438 THE WILSON BULLETIN l Vol. 111, No. 3, September 1999
to prey capture. This species, although a
ground-foraging specialist, has weak legs
(Brown and Amadon 1968). After examining
the rocks (approximate dimensons 30 X 20 X
10 cm), I do not believe that one individual
could have turned over any of the rocks by
itself.
At no time during the days of observation
was the immature bird more than 100 m from
one or the other of the adults, although the
adults were often separated by distances up to
500 m. In a series of 12 one-hour watches (n
= 4 for each individual), I determined that the
two adults appeared to spend more time for-
aging than did the immature bird (adult =
68.6%, imm. = 52.3%) although the differ-
ence was not statistically significant (x2 =
3.334, df = 1, P = 0.067).
Subsequent investigation revealed that
worms or arthropods could be found under
most (19 of 24) of the rocks in the vicinity;
several of the rocks (5 of 24) also hid rodent
runways. Examination of the surrounding
grassland showed that prey items of a similar
size but different taxa (e.g., grasshoppers rath-
er than millipedes) were available without
rock lifting.
DISCUSSION
Unlike most land bird species, Mountain
Caracaras were not using cooperative foraging
behavior to pursue and subdue large, highly
mobile prey items. Rather, they required co-
operation to obtain access to otherwise inac-
cessible prey items. While the turning over of
small rocks was one of the main foraging be-
haviours exhibited by these individuals, the
cooperative rock-turning events did not occur
within the set watches and did not seem to
represent a major foraging strategy for these
individuals. The main difference between the
cooperative behavior observed in Mountain
Caracaras and that observed in other cooper-
atively foraging birds is that only one individ-
ual obtained food from a given foraging event;
Harris Hawks, for example, share large prey
that are cooperatively caught (Bednarz 1988).
This disparity in obtaining a food reward may
even out over time (e.g., one of the four prey
items was given to the immature bird) but
there apparently is often no immediate reward
for some of the individuals participating. That
individuals are willing to help without a re-
ward is perhaps indicative of the length of
time these birds remain together as a foraging
unit; that is, an individual is willing to help
today because its turn will come eventually
(see Tiivers 1971 for discussion of reciprocal
altruism).
Congeners of the Mountain Caracara hatch
their eggs in December and fledglings are usu-
ally independent by March (Brown and Ama-
don 1968). As my observations took place in
September, the immature member of the trio
was probably a chick from a previous breed-
ing effort and its presence, therefore, may rep-
resent delayed dispersal. Delayed dispersal is
fairly common in Neotropical raptors (Mader
1981). How common delayed dispersal is in
Mountain Caracaras and how it may affect the
incidence of cooperative foraging, is uncer-
tain.
ACKNOWLEDGMENTS
I thank .I. Barg, E. Carruthers, E Chavez-Ramirez,
D. Jones, W. Rendell and four anonymous reviewers
for providing helpful comments on earlier drafts of this
manuscript.
LITERATURE CITED
BEDNARZ, J. C. 1988. Cooperative hunting in Harris
Hawks (Parabuteo unicinctus). Science 239:
1525-1527.
BROWN, L. AND D. AMADON. 1968. Eagles, hawks and
falcons of the world. Vol. 2. McGraw-Hill, New
York.
COLLOPY, M. W. 1983. Foraging behavior and success
of Golden Eagles. Auk 100:747-749.
DUGATKIN, L. A. AND M. MESTERTON-GIBBONS. 1996.
Cooperation among unrelated individuals: recip-
rocal altruism, by-product mutualism and group
selection in fishes. Biosystems 37: 19-30.
MACDONALD, D. W. 1983. The ecology of carnivore
social behaviour. Nature 301:379-384.
MADER, W. J. 1979. Breeding behavior of a polyan-
drous trio of Harris Hawks in southern Arizona.
Auk 96:776-788.
MADER, W. J. 1981. Notes on nesting raptors in the
llanos of Venezuela. Condor 83:48-51.
MOFXISON, J. L. 1996. Crested Caracara (Caracura
plancus). In The birds of North America, no. 249
(A. Poole and E Gill, Eds.). The Academy of Nat-
ural Sciences, Philadelphia, Pennsylvania; The
American Ornithologists Union, Washington, D.C.
PARKER, T A., III, S. A. PARKER, AND M. A. PLENGE.
1982. An annotated checklist of Peruvian birds.
Buteo Books, Vermillion, South Dakota.
PRUETT-JONES, S. G. 1980. Team-hunting and food
sharing in Parasitic Jaegers. Wilson Bull. 92:524-
526.
SHORT COMMUNICATIONS 439
SERFASS, T L. 1995. Cooperative foraging by North TRIVERS, R. L. 197 1. The evolution of reciprocal altm-
American river otters, Lutra canadensis. Can. ism. Q. Rev. Biol. 46:35-57.
Field-Nat. 109:4X-459. WHITE, C. M. AND D. A. BOYCE. 1987. Notes on the
SULLIVAN, K. A. 1984. Cooperative foraging and court- Mountain Caracara (Phalcoboenus megalopte-
ship feeding in the Laughing Gull. Wilson Bull. rus) in the Argentine puna. Wilson Bull. 99:
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Wilson Bull., 11 l(3), 1999, pp. 439-440
Predation by Rufous Motmot on Black-and-Green Poison Dart Frog
Terry L. Master
ABSTRACT-I observed a Rufous Motmot (Bar-
yphthengus mart@ feeding a black-and-green poison
dart frog (Dendrobates auratus) to another motmot in
the Caribbean Slope lowland rainforest of northeastern
Costa Rica. Neither individual appeared to suffer any
ill effects from what was probably courtship feeding.
Small vertebrates are typical prey for the larger species
of motmots. Blue-crowned Motmots (Momotus mom-
ota) have been observed consuming several species of
poison dart frogs raised in captivity but captive reared
frogs either do not contain, or have reduced levels of,
the toxins that native frogs produce. Relatively little is
known about the effects of poison dart frog toxins on
predators. Presumably, the digestive system of the Ru-
fous Motmot is capable of neutralizing the potentially
toxic effects of such prey. Received 15 Sept. 1998,
accepted 1.5 Feb. 1998.
the black-and-green poison dart frog (Dendro-
bates auratus) and phantasmal poison dart
frog (Dendrobates tricolor). However, these
frogs were raised in captivity and either do not
produce or have relatively low levels of the
characteristic skin toxins &richer 1997; C.
Rowsom, pers. comm.).
Poison dart frogs have long been known to
possess toxic skin secretions, and, because of
their bright coloration, are thought to be apo-
sematic to visually hunting predators such as
Rufous Motmots (Baryphthengus martii)
which presumably have excellent color vision
(Brodie and Tumbarello 1977). Smith (1975)
demonstrated that hand-reared Torquoise-
browed Motmots (Eumomota superciliosa)
showed an innate avoidance of snake-shaped
models with patterns simulating those of coral
snakes. All other snake models were readily
attacked implying that aposematic coloration
is a deterrent to this species. Observations in-
dicate that Blue-crowned Motmots (Momotus
momota) at the National Aquarium consume
several species of poison dart frogs including
At approximately 9:30 CST on 26 March
1995, an adult Rufous Motmot was observed
in secondary lowland tropical forest from a
hiking trail located at Estacion Biologica La
Suerte, near Cariari, Limon Province, north-
eastern Costa Rica (10 26 N, 83 46 W).
The bird landed 25 m from the trail on an
exposed perch 3 m above the ground and was
easily observed for approximately 4 min. Af-
ter 4 min another individual landed on the
same branch next to the first individual. The
newly arrived motmot was carrying a black-
and-green poison dart frog in its beak which
it fed immediately to the first individual. It is
not possible to distinguish between sexes in
Rufous Motmots; however, this behavior was
interpreted as a male who was feeding the fe-
male as a courtship gesture. Both individuals
had diagnostic black breast marks and raquet-
tails indicative of adult birds, suggesting that
this was probably not a fledgling being fed.
The pair continued sitting on the branch for
approximately 30 min after which they flew
off together into the forest. Neither individual
appeared to suffer any ill effects from either
grasping or consuming the poison dart frog.
I Dept. of Biological Sciences, East Stroudsburg
Univ., East Stroudsburg, PA 18301;
E-mail: [email protected]
The typical diet of motmots varies somewhat
in conjunction with body size. Smaller species
prefer insects while larger species consume in-
sects along with other invertebrates, small ver-