Parrots of the Tambopata

Ecology of the Parrots of the Tambopata Alan Lee, March 2007. Introduction The following are summaries of accounts of 16 species of parrot, macaw and parakeet that can reliably be found at one of the three Rainforest Expeditions lodges (Posada Amazonas, Refugio Amazonas and Tambopata Research Centre), taken from the leading specialist field guide “Parrots – A guide to the Parrots of the World” by Tony Juniper and Mike Parr, published in 1998 and reprinted in 2003 with corrections. It is a substantial book 584 pages long, with 88 well illustrated plates, but reading it made me realise how many holes still exist in our knowledge of one of the largest and most uniformly distinctive groups of birds in the world. All parrots belong to the order Psittaciformes, and many consider the entire order also as a single family – Psittacidae. The most recognisable feature that places individual species in the parrot family is the characteristic bill – broad based with a chisel-shaped cutting edge on the lower mandible and with a wide curve and a sharp point on the upper. When walking on a perch or the ground, parrots show a characteristic swaggering gait. The zygodactyl feet (2 toes pointing forward, 2 pointing backward) provide a very strong grip and used with the bill as a kind of grappling hook, enable this largely arboreal grip of birds to climb with great agility. Uniquely among birds, the structure of the feet enables many of the arboreal parrots to manipulate food, usually with the left foot. Parrots undergo a complete moult annually with young birds replacing their juvenile feathers before they are one year old, although adult plumage in some species is only attained by the third year. Adults generally undertake a post breeding moult. Social behaviour Most species of parrot are gregarious for at least part of their year and are encountered in small flocks or at least in pairs. The adaptive significance of the various levels of gregariousness in parrots may be related to foraging effectiveness and anti-predator defence. Some species ‘appoint’ sentinels that watch over feeding flocks, and it is probably that communal roosts have at least some anti-predator benefits. Various birds of prey appear to be the principal predators of parrots, most notably Ornate-hawk Eagle and Orange-breasted Falcon in the Tambopata region. Monkeys, other arboreal mammals and snakes undoubtedly take a lot of eggs and nestlings. Communal activities, including those at clay licks may offer foraging benefits either by exchanging information on information on fruiting trees or by dispersing more effectively to reduce intraspecific competition. Breeding Parrots are mainly monogamous and in the case of larger species pair for life. The bond between pairs is constantly reinforced by allopreening and feeding. This strategy is perhaps adaptive, because of the high proportion of learned as compared to instinctive behaviour : pairs that know each other well and have experience of one another breed more successfully. Most species show no sexual dimorphism and where this does occur it is sligh. A few exceptional species are polygamous – the Kea and Kakapo males of New Zealand sometimes have several female breeding partners. Some parrots are territorial whilst breeding and defend at least a small area around their nest. Species living outside the tropics have more defined breeding seasons, as is the case along the Tambopata River (Brightsmith 2005). Most parrots are cavity-breeders, with nests located in hollows in trees and palms or termitaria. Young parrots are altricial and born either naked or with only sparse down. They are wholly dependent at first on their parents for food and often for warmth. Most species reach sexual maturity between their second and forth year with, as a general rule, maturity reached earlier in smaller species. Parrots of the Tambopata Although standard Rainforest Expedition literature lists 19 species as visible at their lodges, during a year of surveys in 2006 conducted by trained observers totalling 1400km of transect, we did not encounter 2 of these species at any level – Painted Parakeet and Tui Parakeet. These two species are regularly seen in Manu to the north of the Tambopata region, but if we are to include these on our list we may as well also include Peach-fronted Parakeet, Orange-winged Amazon and Red-shouldered Macaw (all found just to the south of the Tambopata region in the Heath area) and Military Macaw (found in the foothills to the west of the Tambopata Research Center and has been observed by the experienced bird guide Renzo Zeppilli). These species were also not seen at the 4 clay licks monitored, and were not viewed from towers and lookouts. Two other species, Scarlet-shouldered Parrotlet and Amazonian Parrotlet were detected, but are exceedingly rare, bringing the effective list down to 15 from a potential 23 species in the region. According to this reference the Blue-winged Parrotlet (Forpus crassirostrus) should also be found in the area, but no records exist for encounters with this species at any site along the Tambopata River, Manu or Heath. Listing issues aside, the Tambopata is of course known as the best destination in the world to visit clay licks (also known as mineral licks, salt licks, avian geophagy sites, saladeros and barrieros). The clay licks experience higher total numbers of birds and a greater number of species than any comparable site in the region. Although some discussion is still ongoing as to the reason for the incredible daily spectacles that can be observed, the long lasting benefits are not clearly understood. However, it is emerging that species that do engage in this activity show an ecological advantage over species in the area that do not engage in this activity. At Posada and Refugio Amazonas, the Black-capped parakeet is one of the most common forest species, and is fairly regularly observed at 2 clay licks and occasionally on a third. However, this species is not recorded at the clay lick at the Tambopata Research Center, and there are significantly lower group encounter rates with this species in the surrounding forest. White-bellied Parrots are not commonly seen at any of the clay licks but are less frequently encountered at TRC where other species enjoying the benefits of more regular use of the biggest clay lick in the area may be out-competing them, while at Posada and Refugio where clay lick use is not observed on a similar scale the White-bellied Parrot is more common. What repercussions does this have for the endangered species in the region, the Blue-headed Macaw? Very little is known about this species – until now no-one has encountered a breeding site and there is nothing published on its general ecology. This species has been observed at all three sites, but appears to be more common at TRC, where it is more reliably seen on the clay lick than the other two sites. It has also been observed in large numbers at other clay licks along the Alto Madre-de-Dios River where it appears to be more common along the foothills of the Andes. This species range is unerringly restricted to the geographical region where clay licks are most common, although undoubtedly this is not the only ecological element at play in determining its current status – almost a fifth of the world’s 350 species of parrot are under threat and most of the world’s parrots do not have access to clay licks. Blue-and-yellow macaw (Ara ararauna)     Identification: The only large, blue backed and yellow bellied macaw in Peru. It is locally common in Amazonian Peru but in drastic decline around major trapping centres (e.g. Iquitos and Pucallpa). It has suffered recent range contraction and decline in accessible areas owing to often illegal trapping for trade, hunting and habitat loss. It occurs in many protected areas and common pet, widespread in captivity. Ecology: Wooded country, often near water, including edge of lowland humid forest (mostly varzea, avoiding lowland terra firme forest), gallery forest in savanna, swamp forest and Mauritia palm swamp. Sometimes forages in more open country, coming to ground for palm fruits in Brazil. Gregarious, seen usually in pairs, family parties or flocks of up to 25 (sometimes many more, especially near roosts). Birds are most often seen in the morning and evening en route between roost and feeding areas. There are thought to be regular communal roosting sites in trees. Diet consists of a variety of locally available fruits (especially of various palms), nuts, leaf buds, etc., including Astrocarium, Mauritia (locally known as Aguaje) and Acrocomia palms, seeds of Platypodium grandiflorum, Sloanea, Brosimum, Sorocea, Spondias, Inga, Parkia, Hura crepitans and Enterolobium, nectar of Combretum laxum and aril of Hymenaea courbaril. Birds feed quietly high in canopy, often near clearings. Large numbers may congregate at favoured riverbank locations (such as Tambopata Research Center), often with other parrot species to take the mineral-rich clay soils that are exposed. Blue-and-yellow macaws nest in holes in dead palms. Breeds from November to January with 2 eggs per clutch. The sexes are similar. Immature has a brown iris. Scarlet Macaw Identification: A large red macaw that can be distinguished from the Red-and-green Macaw by the yellow on the shoulder (compared to the later species green). Slightly smaller with a longer tail and no red feathers on the bare, white skin around the eyes. Distribution and status: Found from Central America to Brazil, it is wide ranging. Scarlet macaws make seasonal movements in search of fruit and they are regular seasonal visitors to some areas. Locally common but declining throughout range, especially around developing centres, owing to habitat loss, trade and hunting for food and plumes. CITES Appendix 1. Ecology: Lowland tropical forest and savanna. Often near rivers throughout the range. Generally seen in pairs, but also parties of 3-4 or flocks of up to 30 with up to 50 roosting communally in tall trees (including mangroves). Diet: Fruit of Inga, Micropholis, Sterculia, Bursera, Dipteryx, Ficus, Spondias mombiu, Hura, Eschweilera and Terminalia and fruits and nuts of various palms, seeds of Jacaranda, Dialium, Caryocar, Hevea, Euterpe, Cedrela, and Sapium, flowers and nectar of Virola and Erythrina. Feeds in the upper canopy, usually silently, associating with other parrot species where food is abundant. Nests in large cavities old dead trees. Prefers soft wooded species so that the hollow can be modified. Breeds November to April in Peru. Sexes are similar, immature with shorter tail and brown iris. There are two races: Am.macao from Costa Rica south through South America and Peru, and A.m.cyanoptera which is larger with blue tips to yellow wing coverts found from Mexico to Nicaragua. Red-and-green Macaw (Green-winged Macaw) Identification: Told from sympatric Scarlet Macaw by green, not bright yellow, upperwing coverts, darker red plumage and slightly larger size, but immature may show some yellowish-green on upperwing-coverts. Generally in pairs, small flocks of several pairs or family parties. Distribution and status: Generally uncommon following decline caused by capture for trade and habitat loss – widespread through the Amazon basin, but usually less common than Scarlet and Blue-and-yellow Macaws (reported as rare in eastern Ecuador) although this situation reversed in Venezuela, Guiana, and Peru. Absent near population centres and declining or already disappeared at peripheries of range owing to habitat loss, trade and hunting. Widespread in captivity. Ecology: Occupies terra firme rain-forest in north of range, apparently avoiding swampy areas while in the south and east of the range they are more often in more open drier formations including floodplain forest, upland forest and dry woodland (Bolivia), entering savanna in Paraguay. Requires large trees for nesting. Birds roost in rock crevices where available and such sites are also used for nesting especially in Gerais region of Brazil. Diet includes fruits of Acrocomia and Caryocar (sw. Brazil) and fruits and seeds of Copaifera lansdorfii and Hymenaea corbaril (n.e Brazil). Breeding period from November to April in Peru. Clutch 2-3 eggs. Sexes similar, immature with shorter tail and brown iris, sometimes also with yellow on upperwing coverts. Chestnut-fronted Macaw Identification: Small green macaw with long pointed tail, bluish-green crown and inconspicuous chestnut forehead. Red underwing-coverts and blue flight feathers. Generally in pairs or small flocks. Large gatherings at roosts. Flight pattern is fast and direct on wingbeats faster than larger macaws. Distribution and status: Resident with some seasonal altitude movements in areas (western Ecuador and northern Venezuela). Fairly common to common over much of range. Although severe habitat causes local declines, moderate deforestation has led to increases in some areas. Ecology: Found in forest including partly cleared and secondary formations, forest edge and more open country with trees, including humid lowland forest with clearings, varzea, swamp-forest with dead trees, palm groves, gallery forest and savanna with pastures. Generally avoids continuous terra firme forest but transient birds may stop off to feed. Roosts communally, often between fronds of palm trees. Diet includes seeds of Hura crepitans, Sapium aereum, Cedrela odorata and Cupania cinerea, pulp and seeds of Inga laterifolia, Micropholus melinoneana, Euterpe precatoria and Gulielma and fruit of Ficus, Cecropia feulla, Caraniana and various palms (especially with smaller fruits); also flowers of e.g. Virola and Erythrina. Birds feed quietly in canopy where they may be hard to detect. Nests in cavity in dead palm or tree, often over water, and unusually in rock crevices. Clutch 2-3. Sexes are similar although some females may show paler, less prominent chestnut band on the forehead. Red-bellied Macaw Identification: Small, mostly green macaw with long pointed tail and swept-back pointed wings, giving birds streamlined appearance in their fast flight. The small bill, flat back to head and rounded chest tapering sharply back to tail-tip all increase the distinctiveness of the flight silhouette. Told from other small green macaws by combination of yellowish underwings, large yellow facial patch and reddish belly (that is not always easy to see) and blue forehead. May appear rather blue in evening light. Superficially similar Aratinga parakeets have relatively shorter wings and no bare yellow facial skin. Unobtrusive when feeding in Mauritia palms, otherwise generally seen in flight. Very Gregarious. Distribution and status: Locally common but their distribution is closely linked to Mauritia palms which are felled to access their fruits. However, populations are probably stable. Red-bellied macaws are most likely resident in some areas, but at least partially migratory elsewhere e.g. north western Bolivia where birds were reported common from September to November but absent November to January of another year. Ecology: Dependent on Mauritia palms, stands of which occur locally in swampy or seasonally flooded terrain from dense moist forest to gallery forest and more open savanna and grasslands. Gregarious with flocks of over 100 reported. Usually roosts in Mauritia palms where birds cling to inside of frond clusters. Feeds mostly on palm Mauritia flexuaosa fruits but berries and the fruits of Roystonea oleracea and Euterpe are also taken. Nests in dead palms (usually Mauritia), often over water. Sexes similar with males possibly on average larger. Immature with pale tip to the bill and smaller dark red patch on belly. Blue-headed Macaw Identification: Small, mostly green macaw with blue head - the contrast of these colours sometimes making a good field mark. Distinguished from sympatric Chestnut-fronted and Red-bellied Macaws by dark bare skin on the face, more extensive and obvious blue feathers on head and a pale bill. Generally seen in flight and probably frequently overlooked. Distribution and status: Western Amazon basin in extreme western Brazil (only from Acre sporadically), eastern Peru in the south-east and extreme north-western Bolivia. In Peru it is known from the upper Rio Huallaga Valley in Loreto, San Martin and Huanuco (including the outskirts of Tingo Maria), from one locality on the eastern slope of Serra de Divisor in the Rio Ucayali catchment, on the Rio Curanja, Rio Purus catchment, Rio Apurimac in Cuzco and in Madre de Dios west of Puerto Maldonado, around Puerto Maldonado and on the Rio Tambopata 50km from the Bolivian border; occurs in Manu National Park. In Bolivia It has been found in La Paz and perhaps south to Beni with indications that birds regularly occur south on the eastern Andean foothills in Boliva south to 14’30’S. Local and erratic in occurrence, but apparently fairly common in places. Perhaps expanding its rather limited range in south-western Amazonia owing to forest degradation. Rare in captivity. Ecology: Found in upper tropical forests between 150 and 1 550m. Prefers disturbed or partly open habitats with birds mostly occurring at forest edge along rivers, in clearings and around partly forested settlements, also reported from swampy areas with Mauritia palms in forest. This species possibly prefers foothill to lowland forest. Not very gregarious; generally in pairs or groups of three (flocks of 20 or more have been reported – AL), apparently not associating with Chestnut-fronted Macaw. Diet and breeding undescribed. Young birds seen with adults in April. Sexes similar, male possibly larger on average. Immature with dark iris. White-eyed Parakeet (Conure) Identification: Mainly green with conspicuous yellow and red underwing coverts and bright red on the bend of the wing. Often seen in small parties in swift straight flight at some height over trees and rivers. Distribution and status: The most widespread Aratinga parakeet, found throughout south America east of the Andes to northern Argentina and the Brazilian coast. Generally resident but some seasonal movements seem to occur in some areas. Locally abundant in Amazonia and eastern Peru. Ecology: This species occurs in a range of mainly lowland forest and woodland habitats, but reaches 1700m in Ecuador and 2500m in Bolivia. In Amazonia it is found in tropical rain-forest mostly adjacent to rivers (including varzea forest), scattered woodland and cultivated land in the pantanals, mangroves, savannas and palms in Guiana, chaco-type woodland, gallery and floodplain forests in Bolivia. Highly gregarious, commonly forming flocks of several hundred birds (more usually up to 30) which range widely between roosts and feeding areas. In Amazonia, birds generally travel along banks of rivers but also cross large tracts of forest to reach hidden oxbows and marshes. Roosts in large communal gatherings in trees, canefields and caves. Associates with other parrots and parakeets while foraging. Diet: Depends on habitat, but includes wide variety of nuts, seeds, fruits, berries, flowers and insects. Generally forages in canopy but grass seeds also taken. Reported foods include fruits of palms and Goupia glabra, Tetragastris altissima and Allantoma lineate, and Erythrina and Ficus flowers. Nests usually in a high tree cavity (often a palm) but also in limestone caverns in Mato Grosso, Brazil. Breeding in eastern Peru from July to August. Sexes similar. Immature has few red feathers on head, no red at bend of wing and olive outermost greater underwing-coverts. Dusky-headed Parakeet Identification: The only Aratinga parakeet with a grey head, otherwise mainly green, darker above, more yellowish beneath. There is blue in the flight feathers and upperwing-coverts. Conspicuous large bare white ocular patch. Usually seen in small flocks at forest edges along rivers. Distribution and status: Generally common throughout the western Amazon basin. Apparently nomadic in some parts of its range. Perhaps increasing owing to clearance and fragmentation of dense forest. Ecology: Lowland, mainly water-associated forest, reported to 750m in Bolivia. Also found in forest remnants in savanna and cleared areas with remnant woodland patches. Apparently seldom flies over dense forest. Usually in pairs or small parties but flocks of 75 reported where food is abundant. Diet: Forages in canopy for a wide range of seeds, fruits of palms, flowers of Erythrina and Dioclea, berries, and on rotting branches for insects and their larvae. Visits exposed clay banks. Nest in a cavity in trees such as old woodpecker nests or arboreal termite nests. Breeding in eastern Peru from June to September. Sexes are similar. Immature like adult but iris dark. Black-capped Parakeet Identification: Mainly green, long-tailed parakeet with black cap and red outer primary coverts and leading edge of wing. Usually remains in tops of tall trees, rarely descending below 30m. Gregarious in flocks in fast, shallowly undulating flight generally at canopy height. Distribution and status: Western Amazon basin, through eastern Peru to northern Bolivia. Generally common but may be declining due to habitat disturbance although status is unlikely to alter in the future as large areas of its range are protected. Ecology: Humid lowland tropical forest, including both varzea and terra firme formations, is the preferred habitat, with less frequent occurrence in the east Andean f