Parrot Behavior at A Rio Manu (Peru) Clay Lick: Temporal Patterns, Associations, and Antipredator Responses
Parrot Behavior at A Rio Manu (Peru) Clay Lick: Temporal Patterns, Associations, and Antipredator Responses
Parrot Behavior at A Rio Manu (Peru) Clay Lick: Temporal Patterns, Associations, and Antipredator Responses
DOI 10.1007/s10211-003-0080-y
ORIGINAL ARTICLE
Introduction
Many animals deliberately eat clay or earth (geophagy).
This occurs in a wide range of animals living in tropical
regions, including invertebrates, reptiles, birds, and
mammals, including humans (Sokol 1971; Arms et al.
1974; Emmons and Stark 1979; Kreulen 1985; Abrahams
and Parsons 1996; Gilardi et al. 1999). There are many
hypotheses for why animals eat soil, including mechanical enhancement of digestion, mineral supplementation,
acid buffering, adsorption of dietary toxins, and gastrointestinal cytoprotection (Gilardi et al. 1999). Based on a
comparison of the chemical composition of selected and
24
Methods
We studied the behavior of nine species of macaws, parakeets, and
parrots feeding at clay licks, or ccolpas, on the Manu River
Madre del Dios, Peru during August 2000. The main series of
licks (1148.930 S, 7125.400 W) is about 1 km upriver from
the Machiguenga Ccolpa Biological Station (1150.2920 S,
7125.5460 W; Fig. 1). We studied a second lick (1150.3900 S,
7125.2450 W) in a nearby quebrada, or creek, about 500 m
upstream from its mouth. The elevation is about 350 m, and the site
is about 12 h by boat (depending on water depth) upriver from the
junction of the Manu and the Alto Madre de Dios Rivers, near the
town of Boca Manu. The licks are the site of the main macaw
studies conducted by Munn (1992), who founded the biological
station.
The study area is within the Manu National Park, created in
1973 as part of the Manu Biosphere Reserve (designated in 1977) to
include the Manu Reserve Zone and the National Park, covering an
area of 1.5 million hectares. Ecological features are extensively
described in the book Manu (Wilson and Sandoval 1996) and in
many papers cited therein. Our study area was selected because it is
above the tourist zone, reducing the likelihood of human disturbance and ensuring that the behavior we were observing was not
unduly influenced by ecotourists (Munn 1992). Observation blinds
25
Fig. 1 Temporal patterns in
initiation of feeding bouts at the
Machiguenga clay licks in
Manu National Park, Peru
26
(1) 812 m from the central section of the river lick on the same
bank, (2) across the river from the river lick, and (3) across the
stream from the quebrada lick. Three or four observers were
required to watch both licks at the same time and record arrivals
and departures by species and interactions (about 3.5 person
months). We coordinated our activities among ourselves and with
our boatman using walkie-talkies to minimize disturbance to the
birds.
We made four kinds of observations: temporal patterns of lick
use, number of each species feeding at the lick, aggressive
encounters at the lick, and responses to predators and other
intruders. We also recorded when each species arrived in the trees
above the lick. From the close blind at the river, we could not see
all the tree tops, and arrival was noted by identifying vocalizations.
Temporal patterns were recorded by noting the time that the
first individual of each species landed on the lick to begin feeding,
and final departure of all individuals. We defined a feeding bout as
the time between when the first individual landed to feed and the
last one departed, with the lick used relatively continuously during
that time. We classified departures into three types. In a partial
departure, most birds left suddenly, but some birds remained and
were rejoined shortly, usually within 15 min. In a temporary
departure, all birds flew and either circled or landed in nearby trees
and returned, usually within 5 min. In a total departure or
abandonment, all birds left and none returned to the lick within
30 min. In the case of temporary departures, we did not identify the
end of the bout until 30 min had elapsed.
We also recorded the number of each species present every 1
5 min, depending on the changing species composition at the lick.
Since the number of individuals (and species) feeding at the lick,
particularly in the early morning, was very dynamic, we arbitrarily
analyzed the data as follows. To examine the number of
heterospecifics present for each species we analyzed the data by
noting how many of each species were present when the number of
any species being examined was the highest for that bout. For
example, if the maximum count for a given feeding bout for
yellow-crowned parrots was 12, we used the number of each other
species present the first time 12 was reached as the heterospecific
feeding flock size. The maximum counts for the various species
usually did not coincide.
Since the number of individuals feeding at the lick varied
markedly from minute to minute, and it was difficult to record all
aggressive interactions when the lick had over 35 individuals, we
recorded the number of aggressive interactions observed and the
number of birds we were observing. We compared aggression
among species using both a conspecific and heterospecific aggression index. We defined the conspecific index for species A as the
total number of intraspecific aggression interactions of A/number
of feeding bouts with A presentmean maximum number of species
A present for all the bouts. We defined the heterospecific index for
species A as the total number of heterospecific aggressive
interactions of A/number of feeding bouts of species Amean
maximum number of all heterospecifics present while species A
was feeding. These two indices provide a comparison for aggression rates among species.
We examined the responses of feeding birds to predators and
intruders by recording whether there was no response, partial,
temporary, or total departure, following certain visual or acoustic
stimuli. Many loud noises or sudden appearances of large birds
triggered a departure, often a total departure. By observing
simultaneously from different blinds we increased our likelihood
of identifying intruders. Raptorial birds (most of which are not
potential parrot predators) included hawk eagles (Spizaetus spp.),
roadside hawk (Buteo magnitorstris), great black-hawk (Buteogallus urubitinga), king vulture (Sarcoramphus papa), black vulture
(Coragyps atratus), greater yellow-headed vulture (Cathartes
melambrotus), and black caracara (Daptrius ater). Birds also
responded to the warning cries of russet-backed oropendola
(Psarcolius angustifrons), red howler monkey (Alouatta seniculus),
and dusky titi monkey (Callicebus moloch), to the voices of people,
the sound of motorized canoes, and the presence of white-lipped
peccaries (Tayassu pecari) and jaguar (Panthera onca).
Results
Daily temporal patterns
Observations on the daily patterns of feeding at two licks
were made in the dry season of August 2000. There were
three main temporal assemblages of species feeding at the
licks: (1) the parrots and chestnut-fronted macaws fed in
the early morning; (2) the large macaws fed from midmorning to early afternoon; and (3) parakeets fed
throughout the day, but peaked in the early afternoon
(Fig. 1). The clay licks served as a gathering place for
parrots and chestnut-fronted macaws in the early morning, and pairs flew to the trees above the lick from nearly
all directions. With first light (between 0530 and
0545 hours), chestnut-fronted macaws and parrots (mealy,
yellow-crowned) began to arrive in the treetops all along
the licks. Over a period of 520 min vocalizations
increased, and the birds coalesced in leafless trees above
the main central licks. Gradually, with increasing group
vocalizations, they began to move lower in the trees and
to land first in the shrubs above the lick, then on exposed
limbs hanging over the top of the lick; they then started
climbing down the vines toward the lick. Although the
noise of the flock increased, the first parrots to land on the
lick were usually silent. On most mornings, the first parrot
to land on the lick was a yellow-crowned parrot, followed
quickly by one or two conspecifics, and then usually by
blue-headed parrots.
Parrots (mealy, yellow-crowned) and chestnut-fronted
macaws always initiated feeding bouts in the early
morning, mainly between 0600 and 0700 hours (Fig. 1).
Most red and green macaws initiated feeding bouts
between 0900 and 1200 hours, although some bouts were
initiated as early as 0800 and as late as 1600 hours.
Although scarlet macaws usually fed with the red and
green macaws, some joined the early morning parrots and
some came to the lick alone. Of the 58 bouts of scarlet
macaws feeding, 19% were of one or two solitary
individuals, with no other parrots, macaws, or parakeets
around. In the quebrada, dusky-headed parakeets sometimes fed alone. The other species rarely fed alone, and
this only occurred when a whole group started to come
down on the vines and branches but were startled away by
the appearance of a predator or intruder.
27
Bouts
(n)
Time of day to
start feeding
Yellow-crowned parrot
36
06:500:07
C
6:470:06
C
6:590:09
C
7:010:09
C
7:040:14
C
10:460:22
B
10:2600:27
B
10:310:47
B
11:460:44
A
176 (0.0001)
Chestnut-fronted macaw
Blue-headed parrot
Mealy parrot
23
42
29
Orange-cheeked parrot
21
45
Scarlet macaw
58
Dusky-headed parakeet
45
Tui parakeet
40
KruskallWallis 2 comparison
of the lick, often piling on top of one another. Yellowcrowned parrots seemed intimidated by the appearance of
the mass of blue-crowns and either left or moved to the
edges of the lick to continue feeding.
Mean time to start feeding at the clay lick confirms the
relationships shown in Fig. 2 (Table 1). Not all species
started feeding at the beginning of the bout, although most
came down to the lick within about 10 min. Mean
duration of feeding bouts varied significantly by species
(Table 1). Table 1 also shows the order of appearance,
which is the mean number of appearances of the total
times that species visited. That is, each day, each species
was given a ranking for its order of appearance.
The order of appearance was as follows: yellowcrowned parrots (with the lowest number) usually landed
first, followed by blue-headed parrots, whereas orangecheeked parrots were usually last. Since the birds came in
three groups (parrots, parakeets, macaws), the numbers
reflect that pattern. Thus, yellow-crowned parrots came
first for the parrots, the macaws came at about the same
time, and the parakeets alternated which species landed
first (Table 1).
Species associations
Usually there were fewer than 20 individuals of each
species feeding in the early morning group, except for
blue-headed parrots (Table 2). The average number of
red and green macaws feeding at the lick was about 35,
while there were usually only about 6 scarlet macaws
Duration of
feeding
(min)
28.73.28
B
32.74.87
B
37.73.96
A, B
35.04.40
B
31.54.11
B
33.13.48
B
34.53.02
B
38.53.44
A, B
47.43.55
A
17.9 (0.02)
Order of
appearance
1.250.09
D
3.910.23
A
1.980.13
C
3.410.18
B
3.710.29
A, B
1.510.09
D
1.550.16
D
1.380.09
D
1.350.12
D
172 (0.0001)
Total number
of parrots and
macaws feeding
98.310.5
A, B
11211.5
A
94.79.16
A, B, C
11210.6
A
11215.2
A
44.84.23
D
42.14.91
D
77.27.14
B, C
72.14.83
C
83.6 (0.0001)
Bouts
(n)
36
23
42
29
21
45
58
45
40
Yellow-crowned parrot
Chestnut-fronted macaw
Blue-headed parrot
Mealy parrot
Orange-cheeked parrot
Scarlet macaw
Dusky parakeet
Tui parakeet
58.94.13
13.51.26
5.790.64
35.83.59
5.861.39
17.32.69
56.55.67
6.961.36
16.62.41
Mean number
of conspecifics
5.811.58
B, C
4.661.24
C
3.620.91
B
3.811.05
C
Chestnut
0.600.47 0.150.15
B
B
3.241.69 1.020.64
C
C
0.660.53 0.600.40
C
C
0.130.13 0.130.13
B
B
13.54.07
B
15.73.06
B
13.32.31
A
15.23.37
B
Yellow
Other species
13.53.50
B
12.62.18
A
15.82.94
B
13.62.44
B
Mealy
2.520.80
C
2.690.77
B
4.261.26
C
2.440.89
C
Orange
1.560.72 1.090.56
C
C
14.05.11
B
0.000.00
B
0.000.00
C
26.73.46
A
0.000.00
C
0.070.07
C
0.100.07
B
0.000.00
C
0.060.06
C
Red and
green
0.300.17
B
0.360.17
C
6.420.81
A
0.330.25
C
0.340.21
C
0.290.15
B
0.170.12
C
0.110.08
C
Scarlet
Tui
10.51.53
A
42.45.60
A
0.000.00 0.000.00
C
C
0.000.00 0.000.00
B
B
5.432.09 2.572.57
B, C
B, C
3.831.55 3.452.28
C
C
3.291.16 2.381.58
B
B
2.701.60 3.002.41
C
C
3.111.18 3.441.99
C
C
Dusky
172 (0.0001)
96.3 (0.0001)
196 (0.0001)
248 (0.0001)
85.7 (0.0001)
134 (0.0001)
120 (0.0001)
107 (0.0001)
152 (0.0001)
KruskalWallis 2 comparison
65.29.09
A
64.26.82
A
64.17.04
A
55.16.71
A
Blue
28
29
Fig. 3 Frequency distribution
of the maximum number of
each species feeding at the clay
lick. Shown are percentages for
each feeding group size
clay licks. It is clear from this table that some species (tui
parakeet) rarely fed with other species, while the parrots
always fed with other species.
Even at the river lick, they did not use the five faces
equally (Table 3). Only the parakeets used the first
section, and they mainly used the first and second sections
and did not come to the central faces. Similarly only the
macaws used the fifth section.
30
Table 3 Use of different faces of the river lick, given as percents of
bouts for each species on each face
River Licks
Yellow-crowned Parrot
Chestnut-fronted macaw
Blue-headed parrot
Mealy parrot
Orange-cheeked parrot
Scarlet macaw
Red and green macaw
Tui parakeet
46%
50%
50%
50%
67%
34%
27%
25%
25%
21%
20%
22%
37%
32%
47%
29%
25%
29%
30%
11%
7%
14%
53%
22%
27%
Aggressive interactions
The feeding assemblages of macaws, parrots, and parakeets at clay licks are very dynamic, with frequent shifts
in the species composition, the number of birds present,
and the relative proportion of different species. The large
numbers of birds at the lick resulted in displacements as
birds tried to land; there were also deliberate displacements, lunges, and fights with others on the lick. In
general, birds landed near conspecifics and defended
space against them, often supplanting them. The conspecific aggression index was higher for each species than
the heterospecific aggression index (Table 4). There were
also differences in the index among species: (1) in the
early-morning feeding group, blue-headed parrots were
the most aggressive, followed by mealy parrots; (2) red
and green macaws were more aggressive than scarlet
macaws; and (3) the two parakeets had similar aggression
indices (Table 4). The heterospecific index mirrored the
Early-morning group
Yellow-crowned parrot
Blue-headed parrot
Mealy parrot
Chestnut-fronted macaw
Orange-cheeked parrot
Mid-morning group
Red and green macaw
Scarlet macaw
Early-afternoon group
Dusky-headed parakeet
Tui parakeet
Conspecific
index
Heterospecific
index
0.065
0.391
0.204
0.037
0.022
0.006
0.063
0.021
0.003
0.0009
0.516
0.19
0.317
0.028
0.082
0.11
0.002
0.003
Table 5 Response of macaws, parrots, and parakeets to intruders and disturbances at a clay lick in Manu, Peru. Given is percent of bouts
the birds partially or permanently abandoned the lick (the remaining time they stayed)
Disturbance
Raptor
Large hawks/eagles
Roadside hawk
Black vulture
King vulture
Black caracara
Macaw/parrots
Large macaw call
Large macaw land
Small macaw call
Blue-headed landa
Oropendola callb
Monkeys
White-lipped peccaries
Jaguar
Machiguenga
Researchers
a
Over
b
Large macaws
Parakeets
Bouts
(n)
Partial or
temporary
%
Abandoned
%
Bouts
(n)
Partial or
temporary
%
Abandoned
%
Bouts
(n)
Partial or
temporary
%
Abandoned
%
28
13
8
4
14
54
0
25
100
14
38
0
50
0
14
33
22
20
8
23
33
0
20
0
26
55
0
60
100
70
24
12
10
10
26
17
58
10
0
19
83
25
80
100
79
66
21
26
65
50
45
38
5
12
22
28
79
18
196
35
55
59
95
88
0
54
13
43
18
20
0
6
26
46
87
0
25
30
66
64
0
79
89
50
26
46
56
56
21
49
18
65
53
6
55
33
6
28
16
16
60
58
33
36
13
25 landing at once
Loud social calls and alarm calls
14
27
25
6
18
19
57
82
80
100
94
11
31
lived about 6 km upriver sometimes passed by in nonmotorized canoes. The macaws, parrots, and parakeets
nonetheless responded differently regardless of the similar-looking boats. While the researchers usually moved
along the middle of the river, or were closer to the clay
lick, the Machiguenga usually went in the center or
toward the opposite side (near the sand bar where they
could look for turtle eggs). The greatest response, loud
calling and flying around, followed by complete abandonment of the lick and all the trees in the vicinity,
occurred when a boat of five Machiguenga stopped on a
mudflat across from the clay lick, and the people
disembarked to hunt for turtle eggs. When a similar
boatload of researchers landed, the birds were only
temporarily disturbed and returned to feeding on the lick.
Both groups of people were generally silent, and the noise
level did not appear to vary between groups.
Discussion
Competition
Fig. 4 Percent of times each group totally abandoned the feeding
bout because of interactions with different intruders and predators.
Large macaws include scarlet, and red and green; parrots include
chestnut-fronted macaw and the parrots; parakeets include tui and
dusky-headed. *P<0.01; **P<0.001
32
33
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