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MAYA PROJECT RESEARCH
Description and Results
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Black Hawk-Eagle (Spizaetus tyrannus)
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The following account of our
research activities follows the outline just given. For each
topic, we describe the associated conservation challenges, give
some background knowledge on the topic, describe our research
activities, and give a brief synopsis of our results.
1. Basic Biology of Poorly-known Tropical Forest Birds of Prey
Many tropical forest raptors are biologically poorly known.
We know little in detail concerning the kinds of habitat these
raptors need, nor the amount of space they require. We also know
little about their diets, breeding biology, and population
ecology.
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Hector Madrid places a bal-chatri trap
in a tree-top,
in hopes of capturing
an Ornate Hawk-Eagle.
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Knowledge concerning these topics is basic to the development
of conservation efforts geared toward these species, should such
efforts prove necessary.
Because our organization is dedicated mainly toward the
conservation of birds or prey, 90% of our project effort was
devoted to answering these basic questions concerning tropical
forest raptors.
Including migrants and uncommon species, 50 species of
raptors (42 diurnal raptors and 8 owls) occur in the forests of
Petén, Guatemala. However, in our main study site, Tikal
National Park, about 21 species of raptors (not including the
vultures) are common enough to study. Of these 21 species, we
studied 20.
For each species, we studied the breeding biology and
behavior, diet, habitat use, and spatial needs. Our basic
strategy was to locate several breeding pairs of each species,
and conduct extensive observations at the nest--usually from a
platform in a nearby tree. In defining habitat needs, we also
made use of various kinds of census or population sampling
methods, described below.
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Ready for release, this White Hawk
(Leucopternis
albicollis) has been
fitted with a back-pack radio transmitter;
the
antenna wire is visible.
(Walter Martínez in photo)
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In many cases we also captured and placed radio transmitters
on several individuals over time. This allowed us to map out the
birds' movements, and to estimate the amount of space used,
i.e., home range size. This information, along with the
distance between neighboring nests, allowed us to estimate the
amount of habitat normally needed by a nesting pair (see below).
Using radio telemetry also facilitated our observing the birds'
behavior and habitat use.
From 1989 to 1996, we put in thousands of person-hours
studying these 20 raptor species. As a result, we have greatly
increased the sum total of scientific knowledge about many of
these raptors. Much of this information has been published in
scientific journals (see the Maya Project Bibliography given
here), and much awaits publication for the first time in a book
we are in the process of compiling.
The Ornate Hawk-Eagle (Spizaetus
ornatus) was one of our main study species. We studied
nine territories during 45 territory-years. Slightly
larger than a Red-tailed Hawk (Buteo jamaicensis),
females in this species weigh about 40% more than males.
Endowed with formidable talons, these handsome raptors hunt by
stealth within and below the canopy, stalking prey and then
attacking from a short distance. Among 325 identified prey
items, 56% were birds and 44% mammals, though bird contributed
70% of the diet by biomass, and mammals 30%. Birds taken were
largely toucan-sized but included some very large birds--guans,
curassows, and Ocellated Turkeys (Meleagris ocellata).
Most of the mammals taken were tree squirrels, but several
opossums, young coatimundis (Nasua narica), and a few
agoutis (Dasyprocta punctata)--large forest-floor
rodents--were also taken
(Whitacre et al. unpubl.).
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Atop Tikal's Temple 4, Julio Madrid
searches for a
signal from a
radio-tagged Ornate Hawk-Eagle .
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These birds built large stick-nests in tall, emergent trees
amidst mature forest. They laid a one-egg clutch, with most eggs
hatching late in the dry season and young fledging during the
first two-thirds of the rainy season. Incubation was mostly 44-46 days
in duration and chicks fledged 66-92 days after hatching. These
hawk-eagles displayed a prolonged period of juvenile dependence,
as verified in many cases by placing radios on juveniles prior
to fledging. Four young birds achieved independence on average
at 15.2 months of age and 12.4 months after fledging.
Adults advertised their territories by soaring prominently
above the canopy on warm, sunny mornings. Neighboring nests
averaged 3 km apart, for an average density of one territorial
pair per 10-12 km2 within local areas of suitable
habitat. Two adult males used home ranges estimated at 10-14 km2.
Early in the nesting period adult females ranged over quite
small areas--no more than 3 km2--whereas during the
period of fledgling dependency, they ranged over 11-19 km2,
depending on the method of home range calculation used
(Whitacre et al. unpubl.).
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This yearling Ornate Hawk-Eagle (Spizaetus ornatus)
was followed for the first three years of her life.
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In 45 territory-years, an egg was laid in 50-58% of
territory-years, depending on the method used. Our best estimate
was that 61% of eggs produced a fledgling and 53% of eggs
produced an independent adult. Hence we estimate that on average
each territory produced 0.305-0.35 fledglings per year and
0.265-0.306 independent adults. If the territory densities we
discovered (ca 10 territories per 100 km2) hold true
over large areas, then about 2.7-3.1 independent subadults were
produced yearly per 100 km2 of appropriate habitat.
Though these hawk-eagles nest principally among large
expanses of mature forest, we also studied a few nests in forest
fragments or peninsulas amidst the partly-deforested farming
landscape. Such instances, however, may represent cases of
adults clinging to traditional nest sites in the face of
progressive deforestation. It remains unknown precisely how much
forest fragmentation these hawk-eagles may be able to tolerate
while still maintaining a viable population.
The forest-falcons (genus Micrastur) are a strictly
Neotropical group of some half-dozen species. Though members of
the Falconidae, they are very different from "typical
falcons" of the genus Falco. These raptors are
secretive dwellers of the forest understory, rarely venturing
into the open or above the canopy. These fascinating raptors are
most active in the early morning and late evening hours. Rather
than soaring above the canopy, they maintain territorial spacing
by voicing loud calls that carry great distances through the
still pre-dawn forest. They have a distinct facial ruff like
that of an owl or harrier (Circus spp.), which probably
assists them in detecting prey by ear. About the size of a
merlin (Falco columbarius), these forest-falcons are
relatively size-dimorphic, with females averaging nearly 42%
heavier than males.
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The Barred Forest-Falcon
(Micrastur ruficollis)
was another of our
main study species.
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Russell Thorstrom studied Barred Forest-Falcons at Tikal,
gathering data on 20 territories during 98 territory-years (Thorstrom
2000, Thorstrom et al. 2000).
Of the more than 400 prey items identified at Tikal, reptiles
(mainly small lizards) made up 61.5% of the diet, birds 22%,
insects 8%, mammals 6% and frogs 2.5% (Thorstrom
2000). In
terms of biomass, birds made up 55% of the diet, lizards 31%,
mammals 10%, frogs 3%, and snakes and insects less than 1%
each. These raptors hunted stealthily below the canopy, and
sometimes frequented army ant swarms, preying on insects and
lizards fleeing from the ants.
Barred Forest-Falcons mainly utilized mature,
"upland" forest at Tikal, and 69 of 70 nests were
judged to occur in this forest type. Forest-falcons, like most
falconids, do not build a nest, and lay their eggs in cavities
in trees. Clutches were of two or
three eggs, averaging 2.6 eggs.
Laying occurred mainly late in the dry season, with hatching
taking place at the onset of the rainy season, a time of
increasing prey abundance. Eggs hatched 33-35 days after laying,
and nestlings fledged 35-44 days after hatching (Thorstrom et
al. 2000). Radio-tagged fledglings dispersed from their parents'
territories within four to seven weeks after fledging, presumably
achieving independence at that time.
Nesting territories were occupied year after year. On
average, occupied nests were 1.0 km apart, yielding an estimated
density of one territorial pair per 1 km2
of habitat.
For several radio-tagged breeding males, home ranges averaged
98-115 hectares in area, depending on the method used, with 2.8
to 5.5% overlap in adjacent home ranges. Russell's best
estimate of exclusive space per pair was 114 hectares, or one
pair per 1.14 km2 of habitat.
On average 1.1 young fledged per breeding attempt, and 0.8
young fledged per territorial pair-year. Annual survivorship of
adults was relatively high, estimated at 92.3% for females and
95.3% for males; several adults were at least nine years old at
the end of Russell's study.
Our studies showed these birds to be less common in the
fragmented and half-deforested farming landscape than within
Tikal's mature forest. Because of their modest home-range size,
these raptors may be more able to persist in farming landscapes
than some other forest raptors. However, forest remnants as
large as the mean home range (about 1 sq km) are uncommon in
many farming landscapes. Hence, we suspect that these small
raptors display a modest ability to persist in the face of
deforestation, but will probably decline or disappear in
excessively deforested landscapes.
Literature Cited, Basic Biology of Poorly-known Tropical
Forest Birds of Prey
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