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TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
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TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
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TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
SANDOVAL LAKE RESERVE
TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
SANDOVAL LAKE RESERVE
TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
SANDOVAL LAKE RESERVE
TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
SANDOVAL LAKE RESERVE
TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
SANDOVAL LAKE RESERVE
TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
SANDOVAL LAKE RESERVE
TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
SANDOVAL LAKE RESERVE
TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
SANDOVAL LAKE RESERVE
TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE
SANDOVAL LAKE RESERVE
TAMBOPATA TOURS - MANU NATIONAL PARK - LODGE

Parrot Behavior at a Peruvian Clay Lick: We documented the behavior of 13 parrot species at a geophagy site along the Tambopata River in southeastern Peru. These species use the lick in one or more multi-species aggregations composed predominantly of (1) large parrots and small macaws (81% of lick use), (2) large macaws (5%), or (3) parakeets and small parrots (5%). Monospecific flocks accounted for only 8% of lick use and lone individuals ,1% of lick use. The multi-species aggregations sorted by body size and were generally composed of species with similar coloration suggesting that group composition was driven by a mix of competition and predation. Three species regularly used the lick in monospecific groups and these had the largest group sizes away from the lick, suggesting a causal relationship between intraspecific sociality and lick use in monospecific groups. All groups were wary when approaching the lick, probably due to the risk from landslides and predators. We suggest that clay lick use strategies are molded by predation risk and competition acting on a suite of species with varying body size, coloration, and gregariousness (Tambopata Macaw Clay Lick).

Chuncho Macaw Clay Lick – Geophagy, the intentional consumption of soil, has been documented for a wide range of mostly herbivorous mammals, reptiles, and birds (Sokol 1971, Klaus and Schmid 1998, Diamond et al.1999, Brightsmith 2004). Hundreds of birds (up to 17 species) gather daily at river-edge ‘clay licks’ to consume soil throughout the western Amazon Basin (Emmons and Stark 1979, Burger and Gochfeld 2003, Brightsmith 2004). The birds, mostly psittacines, apparently consume soil for its high concentration of sodium (Brightsmith and Aramburu´ 2004, Brightsmith et al. 2008), but may also receive protection from dietary toxins (Gilardi et al. 1999) Chuncho Lodge. Aggregations of birds which use clay licks vary greatly in species composition and patterns of lick use, and much of this variation remains unexplained (Burger and Gochfeld 2003, Brightsmith 2004, Brightsmith and Aramburu´ 2004, Lee et al. 2009) chuncho macaws clay lick tambopata.

Chuncho Macaw Clay Lick – Observations suggest the birds’ behavior at clay licks has been molded by predation and competition, but few detailed studies have been conducted (Burger and Gochfeld 2003, Brightsmith 2004, Brightsmith and Aramburu´ 2004). Social forces such as information exchange, search for mates, and parental care favor group formation (Ward and Zahavi 1973, Wright et al. 2003). However, there are many costs to group membership including competition for resources (Grand and Dill 1999, Krause and Ruxton 2002) and disease transmission (Hoare et al. 2000). Clay licks and other geophagy sites provide good opportunities to study mixed species aggregations. We studied the behavior of parrots using a large clay lick along the upper Tambopata River in southeastern Peru in an effort to document lick use strategies for comparison with research at other sites in the region (Burger and Gochfeld 2003) chuncho macaws clay lick tambopata.

METHODS DE TAMBOPATA – CHUNCHO MACAW CLAY LICK:

Study Area.—Tambopata Research Center (13u 089 S, 69u 369 W) is in the Department of Madre de Dios in southeastern Peru in the Tambopata National Reserve (275,000 ha) near Bahuaja Sonene National Park (1,091,000 ha). The area is tropical moist forest near the boundary with subtropical wet forest. The elevation is 250 m asl with 3,200 mm of rain per year and a wet season from October to March (Tosi 1960, Brightsmith 2004). The area contains a mix of mature floodplain forest, successional floodplain forest, Mauritia flexuosa palm swamps, and upland forest (Foster et al. 1994) chuncho macaw clay lick tambopata.

The clay lick studied was a 500-m long, 25–30 m high, cliff on the right bank (west side) of the upper Tambopata River – Chuncho. The lick was apparently formed by the river’s erosion of recently uplifted Tertiary age alluvial sediments (Ra¨sa¨nen and Salo 1990, Foster et al. 1994, Ra¨sa¨nen and Linna 1995). It consists of two large exposed areas ,150 m in length on the south end and 200 m in length on the north end. The two are separated by a landslide of ,150 m in width chuncho macaws clay lick tambopata. The south end contains a clay layer ,15–17 m high, topped by a band of sand and cobble about 5 m thick. The north end has clay about 8 m high topped by 8 m of sand and cobble – chuncho macaws.

The soils of the clay layer are rich in high cation exchange capacity clays with high sodium levels (Gilardi et al. 1999, Brightsmith et al. 2008) chuncho macaws clay lick tambopata. The slope of the lick face ranges from moderate (,30u) to nearly vertical (80u).

Lick Counts.—Data were collected during December 2001 and 2002, and January 2000 and 2003 from a point ,200 m from the clay lick. Observers arrived before sunrise and stayed until the end of the early morning activity (0700 to 0730 hrs). Observers watched the staging birds and recorded when the first group of birds began to fly in slow circles in front of the lick in anticipation of landing – chuncho macaws clay lick tambopata. Observers counted all birds perched on each section of the clay lick every 5 min (Brightsmith 2004). More detailed location data were collected in December 2002 and January 2003 (n 5 20 mornings) for each bird on the lick to quantify the social group membership of each species using the lick.

Arrivals and Disturbance.—Observers recorded the numbers and species of parrots as they arrived in the area from a point on the opposite river bank, 400 m to the east of the clay lick. It was not possible to record the birds that arrived from forests behind the lick (to the west) – chuncho macaws clay lick tambopata.

Observers recorded the cause of the disturbance whenever .25% of the birds simultaneously flew from the clay lick or surrounding trees – chuncho macaw clay lick tambopata.

Data Analyses.—The clay lick use by each species was calculated as the total number of ‘bird minutes’ on the lick (Brightsmith 2004). Bird minutes were defined as the number of birds on the lick multiplied by the number of minutes they stayed on the lick (i.e., 4 birds for 15 min each 5 60 bird min). We conducted principal component analysis of the data for birds which simultaneously shared each section of the lick to identify the groups of species which used the lick together – chuncho macaw clay lick tambopata.

Only principal components with eigenvalues $1 are reported. We tested differences among species for group sizes arriving at the lick using KruskalWallace and Mood’s median test with 95% confidence intervals around the medians using StatGraphics Centurion XV. Normal variables are presented as mean 6 SD, while those that failed Shapiro-Wilks’ test for normality are presented as medians with 95% confidence intervals. Alpha 5 0.05 for all statistical tests chuncho macaw clay lick tambopata.

TABLE 1. Sociality of parrot species on the clay lick at Tambopata Research Center, Peru during 20 mornings from December 2002 to January 2003. The species are arranged by body mass. ‘Green macaws’ were recorded when observers could not distinguish between Chestnut-fronted and Red-bellied Macaws. Monospecific 5 percent of counts for which the species was recorded on the lick in a monospecific group. Large Parrot 5 percent of counts when the species was part of the large parrot aggregation – chuncho macaw clay lick tambopata. Parakeet and Large Macaw 5 percent of counts for which species were part of the parakeet and large macaw aggregations. Total 5 number of bird minutes recorded for the species. Body masses are from Dunning (1993) and Terborgh et al. (1990). 

Species Mass (g) – Large parrot – Large macaw – Parakeet – Mono-specific – Other – Totals

  • birds Red-and-green Macaw 1,250 40 52 2 7 0 124
  • birds Blue-and-yellow Macaw 1,125 49 45 1 6 0 1,137
  • birds Scarlet Macaw 1,015 56 42 0 2 0 574
  • birds Mealy Amazon 610 96 1 0 4 0 12,471
  • birds Yellow-crowned Amazon 510 79 3 2 9 6 130
  • birds Chestnut-fronted Macaw 430 95 3 0 2 0 2,267
  • birds ‘Green Macaw’ 100 0 0 0 0 336
  • birds Red-bellied Macaw 370 96 0 4 0 0 1,076
  • birds Blue-headed Macaw 250 72 0 28 0 0 18
  • birds Blue-headed Parrot 247 83 13 2 0 2 1,210
  • birds White-eyed Parakeet 157 68 5 8 17 2 11,363
  • birds White-bellied Parrot 155 0 0 43 58 0 40
  • birds Orange-cheeked Parrot 140 69 1 22 1 7 259
  • birds Dusky-headed Parakeet – 108 –  1 – 0 – 92 – 7 – 0 – 537

Totals Macaws (bird minutes) – – 25,400 – 1,727 – 1,555 – 2,580 – 280 – 31,542

RESULTS TAMBOPATA – CHUNCHO MACAW CLAY LICK:

Thirteen species of psittacines used the clay lick in the early morning period (before 0730 hrs, Table 1). Over 99% of all lick use was in groups. Mixed species aggregations accounted for 92% of the total lick use, monospecific groups 8%, and single individuals ,1% (Table 1). Five principal components (eigenvalue . 1) together explained 58% of the variance in group composition on the clay lick (Table 2) chuncho macaw clay lick tambopata. These principal components represent three mixed species aggregations which use the lick as distinct entities. The large parrot aggregation was composed of three abundant species: Chestnut-fronted Macaws (Ara severus), Mealy Amazons (Amazona farinosa), and Red-bellied Macaws (Orthopsittaca manilata). These were regularly joined by up to seven additional species: White-eyed Parakeet (Aratinga leucophthalma), Yellowcrowned Amazon (Amazona ochrocephala), Blue-headed Parrot (Pionus menstruus), Blueand-yellow Macaw (Ara ararauna), Scarlet Macaw (A. macao), Red-and-green Macaw (A. chloropterus), and Orange-cheeked Parrot (Pyrilia barrabandi). This aggregation, represented by PC I, accounts for 18% of the variance in lick use. The large macaw aggregations contained three common species: Red-and-green Macaws, Scarlet Macaws, and Blue-and-yellow Macaws (PC II, 14% of the variance) which were rarely joined by Blue-headed Parrots, Mealy Amazons, Whiteeyed Parakeets, and Chestnut-fronted Macaws.

The principal components analysis identified three parakeet and small parrot aggregations, one with Dusky-headed parakeets (Aratinga weddellii), Orange-cheeked Parrots, and Blueheaded Parrots (PC III, 9% of the variance), one
with White-bellied Parrots (Pionites leucogaster), Blue-headed Macaws (Primolius couloni), and Dusky-headed Parakeets (PC IV, 9% of the variance), and one with Dusky-headed Parakeets, Orange-cheeked Parrots, and White-eyed Parakeets (PC V, 8% of the variance). These three groups were functionally similar: both formed around flocks of Dusky-headed Parakeets or occasionally White-eyed Parakeets and used the same part of the lick. Thus, these groups were considered collectively as the ‘parakeet aggregation’ chuncho macaw clay lick tambopata.

Ten species were recorded using the lick in monospecific groups, but most were monospecific remnants of the mixed species aggregations. Only three species regularly used the lick in coherent monospecific groups: White-eyed Parakeets chuncho macaw clay lick tambopata,

chuncho macaw clay lick tambopata – Dusky-headed Parakeets, and White-bellied Parrots (Table 2). Single psittacines were recorded on the lick 58 times and these birds were often leading larger groups of birds to the lick (36%) or remained when larger groups abandoned the lick (31%) leaving only 19 instances of single birds using the lick.

We focused on the three mixed-species aggregations as they accounted for .90% of the clay lick use. The three mixed-species aggregations were independent, as they arrived, staged, and descended to the lick separately, and used different areas of the lick. They also rarely reacted to each other’s alarm calls. The behavior of the birds at clay licks can be divided into three distinct phases: arrival in the area, descent to the lick, and lick use – chuncho macaw clay lick tambopata.

Arrival in the Area.—All birds arrived in monospecific flocks. Multiple species, when seen together, did not perch or stage together indicating they were just casual associations. Observers could not usually detect arrivals of White-bellied Parrots or Orange-cheeked Parrots as they flew lower than other species and arrived quietly. However, all other species regularly arrived flying high above the canopy and were readily detected – chuncho macaw clay lick tambopata.

Table 2.- Weights for the five principal components which explain the most variance in group composition of psittacines at an avian geophagy site in southeastern Peru. All principal components have eigenvalues .1. Each principal component is identified with a text label which describes the bird aggregation it represents. The most abundant species in each aggregation are shown in bold – chuncho macaw clay lick tambopata.

  •                            PC I                 PC II              PC III               PC IV         PC V
  • Species – Large parrot – Large macaw – Parakeet 1 – Parakeet 2 – Parakeet 3
  • birds Red-and-green Macaw 0.09 0.56 20.03 0.09 0.12
  • birds Blue-and-yellow Macaw 0.18 0.47 0.04 20.01 0.00
  • birds Scarlet Macaw 0.17 0.54 0.11 0.03 0.03
  • birds Chestnut-fronted Macaw 0.55 20.09 20.08 0.16 0.12
  • birds Red-bellied Macaw 0.45 20.15 20.08 0.22 0.21
  • birds Mealy Amazon 0.51 20.20 20.04 0.10 20.05
  • birds Yellow-crowned Amazon 0.15 20.13 0.17 20.14 20.69
  • birds Blue-headed Parrot 0.21 0.13 0.53 20.14 20.34
  • birds Orange-cheeked Parrot 20.03 20.12 0.66 20.15 0.31
  • birds White-eyed Parakeet 0.23 20.22 0.08 20.27 0.21
  • birds Dusky-headed Parakeet 20.15 20.10 0.42 0.43 0.31
  • birds White-bellied Parrot 20.16 20.07 20.04 0.49 20.22
  • birds Blue-headed Macaw 20.01 0.02 0.16 0.59 20.22
  • Percent variance explained – 18 – 14 – 10 – 9 – 8

The members of the large parrot aggregation began to arrive 8.4 6 5.8 min before sunrise (n 5 70 mornings) and usually perched in trees immediately above and behind the clay lick. The median arriving group size was two except for Red-bellied Macaws which was three (Table 3).

The first large macaws began arriving at about the same time as members of the large parrot aggregation (9.3 6 11.7 min before sunrise, n 5 70 mornings). The large macaws continued to arrive throughout the morning at a slow steady rate (1.1 6 0.3 individuals/min, n 5 577 birds over 6 days). Macaws arrived in pairs (61%), singles (30%), and rarely groups of three or four (7%, n 5 291 groups; Table 3).

The members of the parakeet aggregation began to arrive 21.7 6 15.6 min after sunrise (n 5 68 mornings) and staged in short trees at the lick’s left edge. Both common parakeets arrived in large groups: Dusky-headed Parakeet median 5 10, n 5 34 groups, White-eyed Parakeet median 5 22, n 5 65 (Table 3). The arriving groups of parakeets were relatively large, but waited and joined with other conspecifics before moving to the lick.

Our observations suggest many birds spend hours socializing in the trees around clay licks without descending to eat soil. We focus in this paper on birds that consumed soil and do not address the social aspects of gathering near clay licks.

Descent to the Lick.—Most species were able to join more than one type of aggregation, but the three aggregations commonly approached the lick independently and in stereotypical patterns. The members of the large parrot aggregation began to move towards the clay lick by 15.7 6 11.5 min (n 5 66 mornings) after sunrise. There were at least 424 6 152 birds in the area (n 5 6 mornings) at this time. One or more small groups of birds (usually, 20) led the descent by flying in large circles in front of the lick. Birds from the trees joined these groups until there were up to 100 birds in flight. These flights lasted 3.4 6 4.3 min (n 5 62 mornings). The birds flew in slow circles in front of the lick, apparently choosing where to land. Detections of predators or landslides during these flights often caused the birds to choose an alternative section of the lick or break off approach completely.

The large macaw aggregations often formed as groups of 6–29 birds flew to the lick to join the tail end of the large parrot aggregation (19% of 69 mornings). Groups of up to 30 large macaws also initiated lick use on unoccupied sections of the clay lick (29% of 69 mornings). The latter occurred ,50 6 23 min after sunrise (n 5 26 mornings) when they staged and flew to the lick in a manner similar to that described for the large parrot aggregation.

Members of the parakeet aggregation descended to the left edge of the lick starting ,101 6 21 min (n 5 34 mornings) after sunrise. These groups did not engage in exploratory flights like the large parrot aggregation or large macaw aggregation, and instead moved deliberately through the trees progressively closer to the lick and then flew directly from the trees to the lick (usually a distance of ,20 m). There were at least 217 6 120 parakeets in the area (n 5 5 mornings) by the time the first parakeet flocks descended to the lick.

Lick Aggregation Dynamics.— Groups of birds on the lick were fluid; large numbers of birds flew regularly between the surface of the clay lick and the adjacent trees. Some birds took pieces of clay and carried them to the trees for consumption (chuncho clay macaw).

Thus, the maximum number of birds on the lick at any one time was substantially less than the total number of birds in the area. Entire feeding aggregations often abandoned the lick in response to alarm calls (chuncho clay macaw). No apparent cause for the alarm (n 5 1,060 disturbances) was detected in 90% of cases and the birds usually returned to the lick almost immediately. Documented causes of disturbance were rockslides (4%), raptors (2%), other large birds (2%), and people or boats (1%).

The large parrot aggregations formed on 97% of mornings (n 5 71) and accounted for 80% of the total lick use. Additional birds flew directly to the lick once the first birds landed, and numbers on the lick increased rapidly (152 6 85.2 individuals on the lick 10 min after start, n 5 65 mornings). Birds continued to arrive in the area and perch in the trees even after the first birds began to use the lick. At least 951 6 262 birds (range 5 791–1,428; n 5 6 mornings) arrived per morning of which 92% were species that joined the large parrot aggregation (874 6 260 birds, range 5 621–1,336; n 5 6 mornings). The daily maximum number of birds simultaneously on the lick in the large parrot aggregation averaged 192 6 86 (range 5 24 to 497, n 5 69 mornings).

The large parrot aggregation used the lick for 59.6 6 19.2 min (n 5 46 mornings). The three large macaws used the lick in the early morning as part of the large parrot aggregation (50% of total early morning lick use) or in aggregations dominated by large macaws (49% of total early morning lick use, Table 1). The aggregations dominated by macaws formed on 46% of 71 mornings and accounted for 5% of the total lick use. The number of birds on the lick increased within the first 5 min (12 6 9 at first detection, n 5 28 mornings) and remained fairly stable thereafter (14 6 7, n 5 11, 10 min after first detection). The average maximum number of individuals was 17 6 10 (n 5 33 mornings). About 10% of the total birds that arrived in the vicinity of the lick were large macaws (96 6 24 birds, range 5 66–122, n 5 6 mornings). The large macaw aggregations lasted 19 6 13 min (n 5 33 mornings).

The parakeet aggregations formed on 47% of 71 mornings and accounted for 5% of the total lick use. The majority of the birds in the parakeet aggregation were flocking parakeets and these flocks were restless, usually remaining on the lick for only a few minutes before taking flight and returning to the lick or adjacent trees. The average number of the birds on the lick, despite these fluctuations, remained fairly stable with time (27 6 25 birds, n 5 35 mornings, ,5 min after descending to the lick vs. 23 6 21 birds, n 5 8 mornings, 10 min later). About 36% of all birds arriving at the lick were species that joined the parakeet aggregation (340 6 240 birds, range 5 95–653, n 5 6 mornings). The maximum number of birds on the lick in parakeet aggregations averaged 40 6 26 (range 5 3 to 138, n 5 34 mornings). The parakeet aggregation used the lick for 16.2 6 11.4 min (n 5 34 mornings) before they dispersed.

Spatial Distribution parrot.—The clay lick was .1 km in length. However, 85% of the total clay lick use occurred on only four small areas, totaling only 18% of the exposed cliff. Each aggregation regularly used the same few lick areas (chuncho clay macaw parrot). The large parrot aggregation used two sections with exposed clay 9.8 to 15.2 m and 1.4 to 8.3 m above the cliff base. Neither section had vegetation immediately adjacent to the area used by the birds. The large macaw aggregation used two tall center sections of the lick with exposed clay 7.8 to 15.7 m high (chuncho clay macaw parrot).

Both were isolated from surrounding vegetation. Large macaw aggregations did not form on the lower portion of the lick. The parakeet aggregation used the far left edge of the lick almost exclusively (chuncho clay macaw parrot). This section had exposed clay 8.6 to 16 m high and trees immediately adjacent to it (chuncho clay macaw parrot).

Lick Use by Other Pssitacines.—The Whitebellied Parrot was uncommon on the lick (Table 1). It was difficult to detect when arriving, but apparently arrived in groups of up to 10 (Gilardi and Munn 1998; DJB, pers. obs.). This species did not depend on joining with other birds to use the lick. Small groups perched in the trees on the left edge of the lick and remained vigilant while a few individuals at a time descended to the lick (2.8 + 2.0 individuals, n 5 23). This species usually used the lick in monospecific groups (47%) or with the parakeet aggregation (36%, Table 1). This species also use the lick until ,1000 hrs, well after termination of the early morning activity (Brightsmith 2004).

 

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