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NESTING IN SOUTHEASTERN : SEASONAL PATTERNS AND KEYSTONE Author(s): DONALD J. BRIGHTSMITH Source: The Wilson Bulletin, 117(3):296-305. 2005. Published By: The Wilson Ornithological Society DOI: http://dx.doi.org/10.1676/03-087A.1 URL: http://www.bioone.org/doi/full/10.1676/03-087A.1

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Wilson Bulletin 117(3):296±305, 2005

PARROT NESTING IN SOUTHEASTERN PERU: SEASONAL PATTERNS AND KEYSTONE TREES

DONALD J. BRIGHTSMITH1

ABSTRACT.ÐParrots that inhabit tropical lowland forests are dif®cult to study, are poorly known, and little information is available on their nesting habits, making analysis of community-wide nesting patterns dif®cult. I present nesting records for 15 of psittacids that co-occur in southeastern Peru. The psittacid breeding season in this area lasted from June to April, with smaller species nesting earlier than larger species. Why smaller species bred earlier is uncertain, though it may be related to interspeci®c competition for nest sites or variations in food availability. This study identi®ed two keystone resources used by nesting : Dip- teryx micrantha () and Mauritia ¯exuosa (). Local threats to these plant species are discussed. Received 25 August 2003, accepted 14 April 2005.

Nesting is a critically important stage in the est areas shrink, conservationists must priori- natural history of all species. Reproduc- tize their conservation efforts. Large, old trees tive failure has caused numerous conservation and the cavities they contain are vital for the crises, so knowledge of nesting habits is crit- persistence of many cavity-nesting ical (Ratcliffe 1967, Herkert et al. 2003). The (Mawson and Long 1994, Poulsen 2002). nesting ecology of many tropical species re- However, cavity nesters usually do not use mains poorly documented, especially for can- trees in proportion to their abundance, sug- opy nesters in dense, lowland tropical forests. gesting that some species are more im- The family is the most endangered portant than others to these birds (Martin and large avian family in the world, making its Eadie 1999, Monterrubio and Enkerlin 2004). study a conservation priority (Bennett and In this study, I compiled nesting records for Owens 1997, Collar 1997). Most of our 15 species to determine which trees were most knowledge of parrot nesting comes from an- important to the nesting parrot community in ecdotal accounts by early collectors (Huber southeastern Peru. 1933), regional avifaunal compendia (Cherrie 1916, Havershmidt 1968), detailed studies of METHODS individual taxa (reviewed in Masello and Study area.ÐI studied parrot nesting in the Quillfeldt 2002), and the monumental com- Departamento de Madre de Dios in south- pendium of Forshaw (1989). eastern Peru. The primary site was the Tam- parrot diversity is highest in the bopata Research Center (13Њ 07Ј S, 69Њ 36Ј W; western , where communities 250 m in elevation) on the border between the commonly include more than 15 species (Roth Tambopata National Reserve (275,000 ha) and 1984, Montambault 2002). This diversity Bahuaja-Sonene National Park (537,000 ha). peaks in southeastern Peru, where 18 to 20 The center is located in a small (Ͻ1 ha) clear- species have been reported at various sites ing surrounded by a mix of mature ¯oodplain (Terborgh et al. 1984, Foster et al. 1994). forest, riparian successional forest, Mauritia However, the nesting season for all but ®ve ¯exuosa (Arecaceae) palm swamps, upland species in the region remains undocumented, forest, and bamboo (Foster et al. 1994, Gris- making community-level analyses impossible. com and Ashton 2003; DJB pers. obs.). The Here, I report on the nesting season for 15 forest is classi®ed as tropical moist forest species of sympatric parrots inhabiting low- (Holdridge 1967). The site is adjacent to a lands of the western Amazon Basin in south- 500-m-long, 30-m-high riverbank clay lick, eastern Peru. where up to 1,000 and parrots gather Land clearing and pressures on global for- daily, resulting in high parrot densities in the est resources are constantly increasing. As for- area (Brightsmith 2004a). Annual rainfall is 3,200 mm. The dry season extends from April 1 Duke Univ., Dept. of Biology, Durham, NC 27708, to October, during which monthly rainfall av- USA; e-mail: [email protected] erages 90±250 mm (Brightsmith 2004a). Ad- 296 Brightsmith • COMMUNITY-WIDE PARROT NESTING IN PERU 297 ditional nesting records come from Posada tree. One pair, consisting of a wild bird and a Amazonas Lodge in the Native Community of hand-raised bird released to the wild, nested In®erno (12Њ 48Ј S, 69Њ 18Ј W; 195 m in ele- in wooden nest boxes in 2 years (see Nycan- vation; 2,800 mm annual rainfall; Pearson and der et al. 1995 for a description of the nest Derr 1986, Brightsmith and Aramburu 2004) boxes). Three nests were in ¯oodplain forest and Cocha Cashu Biological Station in Manu (one Ͻ10 m from the river edge) and nine National Park (11Њ 54Ј S, 71Њ 18Ј W;400min were in terra ®rme forest. elevation; 2,000 mm annual rainfall; Terborgh Blue-and-yellow ( ararauna).Ð 1983, Terborgh et al. 1984). These two sites Seventy-two nests in at least 50 different cav- are characterized by similar vegetation and ities were recorded. Most cavities (47 of 50) dry seasons, and they are located 50 km north- were in dead Mauritia ¯exuosa palms. Be- northeast and 250 km northwest of Tambopata cause it is dif®cult to climb dead palms, only Research Center, respectively. 21 nest trees were climbed, and nest contents Nesting records.ÐNesting records consist- were checked infrequently. I estimated that ed of two types: con®rmed nests and birds at egg laying occurred in November (n ϭ 9), De- cavities. Con®rmed nests were locations cember (n ϭ 2), and January (n ϭ 2). I con- where I observed eggs or chicks. Observa- ®rmed ¯edging in late February (n ϭ 1), tions of birds at cavities, where contents were March (n ϭ 4), and May (n ϭ 1). Fifty of not checked, were included only when birds these nests were in a 3-ha section of naturally were observed repeatedly at the cavity and dying Mauritia ¯exuosa palm swamp, where where behavioral cues suggested incubation or dead palms occurred at a density of Ͼ200 per feeding of young. Single observations of birds ha (A. Bravo and DJB unpubl. data). Sixteen at cavities were not included, as parrots may nests were in a small (Ͻ0.25 ha) section of a visit cavities when not breeding. swamp being managed to encourage nesting Most of the nesting records were from July of Blue-and-yellow Macaws (Nycander et al. to August (1998) and November to April 1995). Three other nests were in tall, dead (1999 to 2003) in Tambopata and September palms that rose above the surrounding vege- to November (1993, 1995, 1996, and 1997) in tation in an otherwise healthy palm swamp. Manu. I collected additional unpublished nest- Two nests were in ¯oodplain forest in dead ing records from researchers and guides with palms Ͻ10 m from the river edge. experience working in southeastern Peru. One additional nest was in an unidenti®ed These other observers were stationed at Tam- dead, hollow tree in terra ®rme forest, 10 m bopata year-round. from the edge of a cliff that overlooked the Data analysis.ÐI tested the relationship be- Tambopata River. The cavity was a deep, tween body size and the onset of breeding us- open-topped tube, similar in structure to a hol- ing a rank correlation of body mass versus low palm. month of ®rst breeding and a t-test (␣ϭ0.05) (Ara macao).ÐI studied 55 of month of ®rst breeding for large (Ͼ250 g) clutches at 26 different nest sites. I was certain versus small (Ͻ250 g) psittacids (Gibbons of ®rst-clutch initiation for 40 nests: late Oc- 1985). Body-mass data are from Dunning tober (n ϭ 1), November (n ϭ 32), and De- (1993). cember (n ϭ 7). When the ®rst clutch was lost or did not hatch, 35% (7 of 20) of the birds RESULTS re-laid in the same nest. Second clutches were Red-and-green Macaw (Ara chlorop- initiated in late December (n ϭ 4) or early tera).ÐTwelve nests of this species were January (n ϭ 3). Fledging took place in Feb- monitored in Tambopata between 1993 and ruary (n ϭ 4), March (n ϭ 14), and April (n 2003. I determined laying date for nine nests: ϭ 2). No eggs from second clutches hatched. September (n ϭ 1), November (n ϭ 7), and Nests were found in natural cavities of live December (n ϭ 1). Fledging was con®rmed in micrantha (n ϭ 6), live Hymenaea January (n ϭ 1) and March (n ϭ 5; Table 1). oblongifolia (Fabaceae; n ϭ 1), dead Iriartea Most nests were in cavities of live, emergent palm (n ϭ 1), and in arti®cial nest boxes made Dipteryx (Fabaceae) trees (n ϭ 7), although of wood or PVC pipe (n ϭ 18). No nests were one nest was in a cavity of an unidenti®ed found in dead Mauritia palms. Nests were in 298 THE WILSON BULLETIN • Vol. 117, No. 3, September 2005

¯oodplain forest (n ϭ 15), terra ®rme forest Scarlet-shouldered ( hue- (n ϭ 9), and Mauritia palm swamp (n ϭ 1); tii).ÐThis species is rare at the study sites. the habitat for one nest was not recorded. From August to September 1998, guides and Chestnut-fronted Macaw (Ara severa).Ð guests at Tambopata Research Center repeat- Birds were observed attending seven cavities edly saw two birds at a hole in an arboreal from November to February (Table 1). Ob- termite mound 3.5- to 4-m above the ground. servers saw the nest contents in only one cav- The site was in terra ®rme forest with a mix ity; the nest was in a dead Mauritia ¯exuosa of trees and bamboo (Guadua sarcocarpa). palm in the dying section of swamp discussed The birds were seen regularly attending the under Blue-and-yellow Macaw. It contained cavity over a period of a few weeks. chicks in February. The other nests were in Red-crowned Parakeet ( roseif- canopy branches of emergent Dipteryx mi- rons).ÐI located one nest of this species dur- crantha trees (n ϭ 6). All of the nest cavities ing October 1997 in Manu National Park. The were in trees that were relatively isolated from nest was approximately 9 m high in a live tree the surrounding vegetation in terra ®rme for- in late-successional ¯oodplain forest. One est (n ϭ 4), ¯oodplain forest (n ϭ 2), or Mau- newly hatched chick and three eggs were seen ritia palm swamp (n ϭ 1). on 4 October. A total of four birds attended Red-bellied Macaw (Orthopsittaca manila- this nest. They appeared to be adults, although ta).ÐI observed birds attending 26 cavities in two of the birds had less red on the head and Tambopata from October to February. Four may have been young from the previous year. nests contained eggs or chicks, and I estimat- During my last visit to the nest (4 November), ed that eggs were laid in October (n ϭ 2) and I heard young birds begging inside the cavity. November (n ϭ 2). All nests were in dead White-bellied Parrot (Pionites leucogas- Mauritia ¯exuosa palms; three nests were in ter).ÐTwo live featherless chicks were found the small (Ͻ0.25 ha) section of managed palm at the base of a suspected nesting tree in Oc- swamp, and 22 nests were in the 3-ha section tober, indicating that eggs were laid in August; of naturally dying Mauritia ¯exuosa swamp. ¯edging would have occurred in November or Both habitats are described above in the sec- early December. Birds were seen attending tion on Blue-and-yellow Macaw. three additional cavities in Tambopata and White-eyed Parakeet ( leucopthal- Manu from September to February. The nest mus).ÐIn January, my assistants observed cavities were in canopy branches of live trees birds repeatedly attending a cavity in a dead (two and one unknown palm in the center of a farm ®eld. The pair species). was likely incubating or brooding because one Blue-headed Parrot ( menstruus).Ð bird entered the palm and refused to leave, Five nests of this species were found, all from even when observers knocked on the trunk of June to November (Table 1). Laying dates the palm. were calculated for two nests: late May or ear- Dusky-headed Parakeet (Aratinga weddel- ly June (n ϭ 1) and September (n ϭ 1). lii).ÐNo chicks of this species were seen; all Chicks were seen in three nests. Fledging in three records reported here are of birds at- November was con®rmed at one nest. Nest tending cavities in dead trees or dead branches sites were dead palms (n ϭ 2) and PVC nest in live trees. The reports from Tambopata boxes (n ϭ 2). All nests were near some sort come from July, December, and January. of forest edge: river edge (n ϭ 2), clearing These observations are congruent with a re- edge (n ϭ 1), and a steep drop-off in terra port from local residents who say that the spe- ®rme forest (n ϭ 1). Habitat was not recorded cies nests ``year-round'' (Sixto Duri pers. for one nest. comm.). Nest cavities were in a dead tree of Yellow-crowned Parrot (Amazona ochro- an unknown species (n ϭ 1), a dead palm (n cephala).ÐA member of this species was seen ϭ 1), and a dead branch in a live Cecropia attending a dead palm from at least December (Cecropiaceae) tree (n ϭ 1). The nesting hab- to March in mature ¯oodplain forest (A. del itats included river edge (n ϭ 1), small farm Campo pers. comm.). However, local people (n ϭ 1), and a large natural gap in terra ®rme report that the chicks of this species ¯edge in forest (n ϭ 1). October. Birds were also seen brie¯y at three Brightsmith • COMMUNITY-WIDE PARROT NESTING IN PERU 299 observed Dry season Apr May Wet season XXXXX XXXXX o o XXXXX o Nov Dec Jan Feb Mar Dry season Jun Jul Aug Sep Oct a ) 67 XXX X ) 510 CCCCC ) 157 C ) 370 X XXXX )65XXX ) 108 C C C ) 155 X X X C C C )60CC ) 1,250 o o ) 1,125 ) 59 XXX X ) X 247XXXXX ) 430 XXXX ) 610 XXXX X Touit huetii ) 1,015 o Ara severa cyanoptera Ara ararauna ( Aratinga weddellii Amazona ochrocephala Ara chloroptera Pyrrhura roseifrons b Aratinga leucopthalmus Pionites leucogaster Species Mass Orthopsittaca manilata Pionus menstruus Ara macao Brotogeris sanctithomae Amazona farinosa ( b Nest data from Brightsmith (2004b). Mass data from Dunning (1993). TABLE 1. Parrot nesting phenology, by month and by season (dry or wet), in southeastern Peru (Departamento de Madre de Dios), 1993±1999. An ``X'' a b Chestnut-fronted Macaw ( Blue-headed Parrot ( Red-bellied Macaw ( Dusky-headed Parakeet ( Cobalt-winged Parakeet Yellow-crowned Parrot ( White-eyed Parakeet ( White-bellied Parrot ( Blue-and-yellow Macaw ( Scarlet Macaw ( Mealy Parrot ( Tui Parakeet Scarlet-shouldered Parrotlet ( regularly attending cavities (but no eggs or nestlings observed). indicates peak breeding season (and chicks or eggs seen in the nest), ``o'' indicates periods when few birds breed, and ``C'' indicates that birds were Red-and-green Macaw ( Red-crowned Parakeet ( 300 THE WILSON BULLETIN • Vol. 117, No. 3, September 2005 other cavities in ¯oodplain forest (Dipteryx September to mid-November (Brightsmith micrantha, Erythrina sp. and an unknown spe- 2000, 2004b). Amazonian (Nannop- cies; A. del Campo and J. I. Rojas pers. sittaca dachilleae) in Tambopata attended a comm.). tree cavity in July and September (O'Neill et Mealy Parrot (Amazona farinosa).ÐI ob- al. 1991). Of the remaining four species, none served two nests of this species. Eggs were has been found nesting in the region: Dusky- laid in December (n ϭ 1) and January (n ϭ billed Parrotlets ( sclateri) were seen 1). Both nests were in ¯oodplain forest; one going in and out of a tree cavity in July in had two eggs, the other one egg and one northern Peru (Forshaw 1989). No nests have chick. One was in a dead Iriartea palm, the been reported for Orange-cheeked Parrots other in a natural cavity in a live emergent (Pionopsitta barrabandi), but sightings of re- Dipteryx micrantha tree. cently ¯edged young of this species at the Relationship between body size and breed- Tambopata clay lick in December and Febru- ing season.ÐMost parrots and parakeets bred ary suggest that the birds may lay eggs in Oc- from June to November, whereas the macaws, tober or earlier (DJB pers. obs.). In , Mealy Parrot, and White-eyed Parakeet nested recently ¯edged Orange-cheeked Parrots also from November to March (Table 1). Smaller were seen during February and early March parrot species initiated nesting signi®cantly (Roth 1984, Forshaw 1989). No nesting in- earlier in the season than larger parrots. This formation is available for Black-capped Par- trend held for all 15 species (rank correlation: akeet (Pyrrhura rupicola) or Blue-headed Ma- r ϭ 0.70; t-test: t ϭ 3.08, df ϭ 13, P ϭ 0.009; caw (Propyrrhura couloni; Forshaw 1989, species listed in Table 1) and for the 11 spe- Collar 1997, Juniper and Parr 1998). cies of which I observed chicks or eggs (rank The ®nding that smaller species bred earlier correlation: r ϭ 0.81; t-test: t ϭ 5.25, df ϭ 9, was unexpected, but it could be related to in- P Ͻ 0.001). I was unable to analyze body size terspeci®c competition for nest sites or vari- versus nesting season using only phylogenet- ations in the availability of food resources ically independent contrasts because there is (Roth 1984). Competition between species is no complete phylogeny of New World parrots potentially important because Chestnut-front- (Wolf et al. 1998, Tavares et al. 2004). How- ed Macaws, Mealy Parrots, Yellow-crowned ever, the available partial phylogenies show Parrots, (Ramphastos spp.), Scarlet that there are at least two independent com- Macaws, and Red-and-green Macaws all over- parisons in the data set: Red-bellied Macaw lapped in their nest-site preferences (DJB un- versus Blue-and-yellow Macaw and Blue- publ. data). However, most of the smaller spe- headed Parrot versus Mealy Parrot (Russello cies that nested early in the season used sub- and Amato 2003, Ribas and Miyaki 2004). In strates and cavities ignored by larger birds both cases, the smaller species nests ®rst. (e.g., termite mounds, thin dead palms, and small cavities; DJB pers. obs.) suggesting that DISCUSSION something other than just competition drives During this study, nests or suspected nest the nesting phenology patterns I observed. sites were found for 13 of the 20 species of Seasonal differences in nesting may be due psittacids. This includes the ®rst nest-site de- to differences in diet and food availability. scription for Scarlet-shouldered Parrotlet and The smaller parrots that nest in the dry season the second for White-bellied Parrot. For usually eat more nectar, ¯owers, and small White-bellied Parrot, the only previous nest- seeds than larger species (Roth 1984, Desenne ing record is of an incubating female in a tree 1994; see also Terborgh 1983 for similar pat- cavity in eastern Brazil (Forshaw 1989). Of terns exhibited by primates). Flowering in the seven psittacid species for which nests many tropical communities peaks in the dry were not discovered, previous nest data are season (van Schaik et al. 1993, Fenner 1998) available for three. In Manu National Park, and many wind-dispersed in the Cobalt-winged Parakeet (Brotogeris cyanop- dry season, when deciduous canopy trees lose tera) and Tui Parakeet (B. sanctithomae) both their and higher wind velocities pro- nest in termite mounds, lay eggs in August duce ideal dispersal conditions (Fenner 1998). and September, and have chicks from mid- Because ¯owers and small wind-dispersed Brightsmith • COMMUNITY-WIDE PARROT NESTING IN PERU 301 seeds are relatively low-quality foods that re- can live for over 1,000 years, cavities in these quire a large energy investment to harvest, trees may remain useable by macaws for de- smaller-bodied parrots should have an advan- cades or centuries (Chambers et al. 1998; but tage when exploiting these resources (Ter- see Fichtler et al. 2003). As a result, hundreds borgh 1983). As a result, larger species should of macaw chicks may be produced from a sin- incur comparatively greater food shortages in gle tree during its lifetime. The of Dip- the dry season than smaller species, explain- teryx species are also a keystone resource for ing the wet season breeding of larger parrots a variety of tropical frugivores and granivores, found in Tambopata. Notably, the earliest- including Great Green Macaws (Ara ambigua; breeding species was the mid-sized Blue- G. Powell unpubl. data), ( spp.; headed Parrot; members of its (Pionus) Romo 1997), squirrels (Sciurus spp.), and are known to eat many ¯owers (Galetti 1993). agoutis (Dasyprocta spp.; Emmons 1984, For- Nest searching was not conducted with get 1993) equal intensity in all months. Although sam- Unfortunately, Dipteryx trees are increas- pling efforts were more intense later in the ingly logged throughout their range. Dipteryx season, most of my nesting records for small wood is in high demand for hardwood ¯ooring species come from the early part of the sea- (Toledo and RincoÂn 1999, Wood Flooring In- son. I have had crews observing macaws con- ternational 2003) and, in Peru, people use the tinuously from November 2000 to May 2004, wood to make charcoal. Landowners sell trees and they did not witness large macaws nesting Ͼ1 m in diameter for as little as US $30 (A. earlier in the season. Conducting more nest Hepworth unpubl. data). The recent increased searches from May through August would harvest of Dipteryx panamensis in Costa Rica likely reveal additional small species breed- is the most probable cause for the precipitous ing, corroborating the trend we found. decline of Great Green Macaws in that coun- Few studies have addressed parrot nesting try (Bjork and Powell 1995, Chassot and seasonality at the community level. Roth Monge 2002). Management schemes involv- (1984) hypothesized that congeners staggered ing planting of Dipteryx trees are underway in breeding to avoid competition for food. His Costa Rica. This can produce fruiting trees, data support the pattern for Aratinga and but large, gnarled adult trees riddled with use- Amazona, where smaller species did nest ear- able cavities are practically irreplaceable, as lier, but Pyrrhura and Ara overlapped exten- they take hundreds of years to grow. sively. My data, however, do not support tem- Palms have long been recognized as vital poral spacing by congeners, and my analysis to the survival of tropical frugivores and gran- of Roth's (1984) data shows that smaller spe- ivores (Emmons 1984, Henderson 1995). In cies tended to nest earlier, but not signi®cantly fact, several New World parrots are thought to so (rank correlation: r ϭ 0.34; t-test: t ϭ 0.94, be almost completely dependent on palms for df ϭ 12, P ϭ 0.37). Future studies should in- food, nesting sites, or both (Yamashita 1987, vestigate the interplay of competition for nest Forshaw 1989, Yamashita and Valle 1993, Ya- sites, diet, and phenological cycles in deter- mashita and Barros 1997, Salaman et al. mining the seasonality of parrot breeding. 2001). Eight parrot species were observed My study highlighted two types of sites that nesting in palms during this study, and reports are very important to nesting parrots: emer- from the literature show that two additional gent Dipteryx micrantha trees and dead palms. species also use palms (Red-and-green Macaw Six species were recorded using large, emer- and Dusky-headed Parakeet; Forshaw 1989, gent Dipteryx micrantha trees, and 75 and Nycander et al. 1995). In sum, half of the par- 88% of the natural nests used by Scarlet and rot species in this community nest in palms Red-and-green macaws, respectively, were in and palms are important not only for special- these trees (see also Nycander et al. 1995). ists, but for many generalists as well. Large emergents of this species often con- Mauritia ¯exuosa palms are particularly tained dozens of cavities, and individual trees valuable resources for parrots (Forshaw 1989, often had multiple pairs of macaws nesting in Bonadie and Bacon 2000). In Peru, at least them simultaneously (A. Hepworth and DJB seven species of psittacids nest in Mauritia unpubl. data). Because Dipteryx micrantha palms (GonzaÂlez 2003; this study), and studies 302 THE WILSON BULLETIN • Vol. 117, No. 3, September 2005 on Trinidad show that palm swamps are key the region alone, but they are threat- to the maintenance of parrot populations ened because local people commonly cut en- (Bonadie and Bacon 2000). Dying palm tire trees to harvest weevil larvae (Dry- swamps are particularly valuable because par- ophthoridae: Rhynchophorus palmarum) and rots nest at high densities in these sites. Gon- fruit (Peters et al. 1989, Vasquez and Gentry zaÂlez (2003) reported aggregations of macaws 1989). However, many psittacids, game spe- in dying sections of Mauritia palm swamps in cies, and large-bodied seed dispersers that northern Peru, similar to what I report for move between the swamps and the surround- Tambopata. Nesting densities there (0.075 ing landscape also depend on these fruits nests/ha; range ϭ 0.038±0.128/ha) were 100ϫ (Bodmer 1990, Bonadie and Bacon 2000). As smaller than those in Tambopata (Ͼ29 nests a result, the loss of these swamps would have in3haorϾ9.7/ha). In northern Peru, the par- great impacts on the ecosystem. rots spread out over larger areas of dying The two primary nesting resources exploit- swamp, and parrot collectors regularly re- ed by parrots in southeastern Peru are struc- duced the nest densities (GonzaÂlez 2003). In turally different. In fact, they represent op- Tambopata, the birds were concentrated in a posite ends of the tree-cavity spectrum. The small, protected area near a large clay lick dead palms are hollow tubes with open tops (Brightsmith 2004a). Breeding near the clay that allow rain to enter. They are thin-walled, lick may be particularly important because poorly insulated, and ¯imsy; also, they last for adult parrots feed their nestlings clay and con- only a few years before collapsing. In com- centrate their use of the lick during the breed- parison, Dipteryx cavities have thick walls of ing season (DJB unpubl. data). hard wood, full roofs that provide protection Areas of palm swamp can die synchro- from the rain, and are usually in live sections nously in response to ¯ooding and other of solid trees that live for centuries (Chambers events that result in depositions of large sed- et al. 1998; DJB pers. obs.). It is surprising iment loads (Kahn 1988, GonzaÂlez 2003; DJB that two such different substrates attracted the pers. obs.). How long the dead palms remain majority of nesting parrots. The only charac- useable for nesting parrots under natural con- teristic they shared was their isolation from ditions is unknown. However, Mauritia palms the surrounding vegetation. Dead-palm nest that died when their tops were cut off re- sites were almost always in the open: along mained upright an average of 4 to 5 years be- river edges, in dead swamps, above the sur- fore they fell (Nycander et al. 1995; DJB un- rounding canopy, or in forest openings. The publ. data). The short, useful lifetime of in- dividual dead palms suggests that dying palm Dipteryx cavities were far from heavy epi- swamps slowly shift in the landscape as new phyte and vine cover, distant from adjacent areas die and old areas become unusable. Like trees, and high above the ground. This sug- most parrots, those that nest in these dead gests that protection from non-volant preda- swamps probably ``wander,'' tracking shifts in tors has a great in¯uence on parrot nest-site food resources and nest-site availability (Col- selection (Massello and Quillfeldt 2002, lar 1997, Renton 2002). As a result, dying Brightsmith 2005a, 2005b). palm swamps probably serve as breeding The availability of suitable nest sites limits ``source'' areas in landscape-level source-sink the reproductive output of many cavity-nest- dynamics and play a disproportionately large ing species, especially in anthropogenically role in maintaining regional populations of modi®ed landscapes (Newton 1994). This these long-lived and highly mobile psittacids. study suggests that Dipteryx micrantha and Palms are valuable to local people, and doz- Mauritia ¯exuosa are keystone tree species for ens of species are used for food, ®ber, con- parrots nesting in southeastern Peru. Clearing struction materials, fuel, and medicines (VaÂs- for agriculture, targeted destruction of parrot quez and Gentry 1989, Henderson 1995). nests by collectors, and selective felling of Overexploitation is common and its potential key tree species will reduce the density of impact on the ecosystem is great (Johnson suitable nest cavities. Future studies should 1986, Bonadie and Bacon 2000). Mauritia continue to identify key nesting resources for ¯exuosa swamps cover at least 2 million ha in parrots and other cavity-nesting species so Brightsmith • COMMUNITY-WIDE PARROT NESTING IN PERU 303 that these important habitat features can be (Psittaciformes) and trogons (Trogoniformes). conserved in tropical landscapes. Journal of Avian Biology 36:64±73. BRIGHTSMITH, D. J. 2005b. Competition, predation and nest niche shifts among tropical cavity nesters: ACKNOWLEDGMENTS ecological evidence. Journal of Avian Biology 36: I thank the following people for contributing nest 74±83.  records, library work, and support in the ®eld: G. Ar- BRIGHTSMITH,D.J.AND R. ARAMBURU. 2004. Avian rospide, D. Brooks, A. del Campo, C. Carrasco, D. geophagy and soil characteristics in southeastern Combs, Sixto Duri, Silverio Duri, J. GonzaÂlez, A. Hep- Peru. Biotropica 36:534±543. worth, H. Lavado, R. Masias, R. von May, A. Mishaja, CHAMBERS, J. Q., N. HIGUCHI, AND J. P. SCHIMEL. 1998. J. Moscoso, E. Nycander, R. Olivera, J. Pesha, R. Pi- Ancient trees in Amazonia. Nature 391:135±136. ana, J. I. Rojas, A. Valdez, and L. Zapater. Thanks also CHASSOT,O.AND G. MONGE. 2002. Great Green Ma- to my assistants in Manu and Tambopata, the staffs of caw: ¯agship species of Costa Rica. PsittaScene Cocha Cashu Biological Station and Tambopata Re- 53:6±7. search Center, and the Native Community of In®erno CHERRIE, G. K. 1916. A contribution to the ornithology for permission to work on their lands. Thanks also to of the Orinoco region. Museum of Brooklyn In- the of®ces of the Instituto Nacional de Recursos Na- stitute of Arts and Sciences Bulletin 2:133±374. turales (INRENA) in Lima, Cuzco, and Puerto Mal- COLLAR, N. J. 1997. Family Psittacidae. Pages 280± donado. Earlier drafts of this work were improved by 479 in Handbook of the birds of the world, vol. the comments of E. Villalobos, J. Wiley, L. Pautrat, and 4: sandgrouse to cuckoos (J. del Hoyo, A. Elliott, two anonymous reviewers. This work was funded by and J. Sargatal, Eds.). Lynx Edicions, Barcelona, the National Science Foundation (grant number DEB± Spain. 95±20800), Conservation Food and Health Foundation, DESENNE, P. 1994. Estudio preliminar de la dieta de 15 EarthWatch Institute, Rainforest Expeditions (www. especies de psitaÂcidos en un bosque siempreverde, perunature.com.pe), Raleigh-Durham Cage Bird Socie- Cuenca del RõÂo Tawadu, Reserva Forestal El Cau- ty, Amigos de las Aves USA, and W. and L. Smith. ra, Edo. Bolivar. Pages 25±42 in BiologõÂa y con- servacioÂn de los psitaÂcidos de Venezuela (G. Mo- rales, I. Novo, D. Bigio, A. Luy, and F. Rojas- LITERATURE CITED Suarez, Eds.). Gra®cas Giavimar, Caracas, Vene- BENNETT,P.M.AND I . P. F. O WENS. 1997. Variation in zuela. risk among birds: chance or evolution- DUNNING, J. B. 1993. CRC handbook of avian body ary predisposition? Proceedings of the Royal So- masses. CRC Press, London, United Kingdom. ciety of London, Series B 264:401±408. EMMONS, L. H. 1984. Geographic variation in densities BJORK,R.AND G. V. N. POWELL. 1995. Buffon's Ma- and diversities of non-¯ying mammals in Ama- caw: some observations on the Costa Rican pop- zonia. Biotropica 16:210±222. ulation, its lowland forest habitat and conserva- FENNER, M. 1998. The phenology of growth and re- tion. Pages 387±392 in The large macaws: their production in plants. Perspectives in Plant Ecol- care, breeding and conservation (J. Abramson, B. ogy, Evolution and Systematics 1:78±91. L. Spear, and J. B. Thomsen, Eds.). Raintree Pub- FICHTLER, E., D. A. CLARK, AND M. WORBES. 2003. lications, Ft. Bragg, California. Age and long-term growth of trees in an old- BODMER, R. E. 1990. Responses of ungulates to sea- growth tropical rain forest, based on analyses of sonal inundations in the Amazon ¯oodplain. Jour- tree rings and 14C. Biotropica 35:306±317. nal of Tropical Ecology 6:191±201. FORGET, P. M. 1993. Post-dispersal predation and scat- BONADIE,W.A.AND P. R. BACON. 2000. Year-round terhoarding of Dipteryx panamensis (Papiliona- utilization of fragmented palm swamp forest by ceae) seeds by rodents in Panama. Oecologia 94: Red-bellied Macaws (Ara manilata) and Orange- 255±261. winged Parrots (Amazona amazonica) in the Na- FORSHAW, J. M. 1989. Parrots of the world, 3rd ed. riva Swamp (Trinidad). Biological Conservation Landsdowne Editions, Melbourne, Australia. 95:1±5. FOSTER, R. B., T. A. PARKER,A.H.GENTRY,L.H. BRIGHTSMITH, D. J. 2000. Use of arboreal termitaria by EMMONS,A.CHICCHO N,T.SCHULENBERG,L.RO- nesting birds in the Peruvian Amazon. Condor DRÂõGUEZETAL. 1994. The Tambopata-Candamo 102:529±538. Reserved Zone of southeastern Peru: a biological BRIGHTSMITH, D. J. 2004a. Effects of weather on avian assessment. RAP Bulletin, no. 6. Conservation In- geophagy in Tambopata, Peru. Wilson Bulletin ternational, Washington, D.C. www.conservation. 116:134±145. org/xp/CABS/publications/cabs࿞publ࿞research/rap࿞ BRIGHTSMITH, D. J. 2004b. Nest sites of termitarium bulletins/ terrestrial࿞ rap࿞bulletins/terrrapbulletins. nesting birds in SE Peru. Neotropical Ornithology xml (accessed May 2004). 15:319±330. GALETTI, M. 1993. Diet of the Scaly-headed Parrot BRIGHTSMITH, D. J. 2005a. Competition, predation and (Pionus maximiliani) in a semideciduous forest in nest niche shifts among tropical cavity nesters: southeastern Brazil. Biotropica 25:419±425. phylogeny and natural history evolution of parrots GIBBONS, J. D. 1985. Nonparametric methods for quan- 304 THE WILSON BULLETIN • Vol. 117, No. 3, September 2005

titative analysis. American Sciences, Columbus, NEWTON, I. 1994. Experiments on the limitation of bird Ohio. breeding densities: a review. Ibis 136:397±411. GONZAÂ LEZ, J. A. 2003. Harvesting, local trade, and NYCANDER, E., D. H. BLANCO,K.M.HOLLE,A.DEL conservation of parrots in the northeastern Peru- CAMPO,C.A.MUNN,J.I.MOSCOSO, AND D. G. vian Amazon. Biological Conservation 114:437± RICALDE. 1995. Manu and Tambopata: nesting 446. success and techniques for increasing reproduc- GRISCOM,B.W.AND P. M. S. ASHTON. 2003. Bamboo tion in wild macaws in southeastern Peru. Pages control of forest succession: Guadua sarcocarpa 423±443 in The large macaws: their care, breed- in southeastern Peru. Forest Ecology and Man- ing and conservation (J. Abramson, B. L. Spear, agement 175:445±454. and J. B. Thomsen, Eds.). Raintree Publications, HAVERSHMIDT, H. 1968. Birds of Surinam. Oliver and Ft. Bragg, California. Boyd, London, United Kingdom. O'NEILL, J. P., C. A. MUNN, AND I. FRANKE. 1991. Nan- HENDERSON, A. 1995. The palms of the Amazon. Ox- nopsittaca dachilleae, a new species of parrotlet ford University Press, New York. from eastern Peru. Auk 108:225±229. HERKERT, J. R., D. L. REINKING,D.A.WIEDENFELD, PEARSON,D.L.AND J. A. DERR. 1986. Seasonal pat- M. WINTER,J.L.ZIMMERMAN,W.E.JENSEN,E.J. terns of lowland forest ¯oor arthropod abundance FINCK ET AL. 2003. Effects of prairie fragmenta- in southeastern Peru. Biotropica 18:244±256. tion on the nest success of breeding birds in the PETERS, C. M., M. J. BALICK,F.KAHN, AND A. B. AN- midcontinental United States. Conservation Biol- DERSON. 1989. Oligarchic forests of economic ogy 17:587±594. plants in Amazonia: utilization and conservation HOLDRIDGE, L. R. 1967. Life zone ecology, revised ed. of an important tropical resource. Conservation Tropical Science Center, San Jose, Costa Rica. Biology 3:341±349. HUBER, W. 1933. Birds collected in northeastern Nic- POULSEN, B. O. 2002. Avian richness and abundance aragua in 1922. Proceedings of the Academy of in temperate Danish forests: tree variables impor- Natural Sciences Philadelphia 84:205±249. tant to birds and their conservation. Biodiversity JOHNSON, D. V. 1988. Worldwide endangerment of use- and Conservation 11:1551±1566. ful palms. Pages 268±273 in The palm±tree of RATCLIFFE, D. A. 1967. Decrease in eggshell weight in life: biology, utilization, and conservation (M. J. certain birds of prey. Nature 215:208±210. Balick, Ed.). New York Botanical Garden, Bronx, RENTON, K. 2002. Seasonal variation in occurrence of New York. macaws along a rainforest river. Journal of Field JUNIPER,T.AND M. PARR. 1998. Parrots: a guide to Ornithology 73:15±19. parrots of the world. Yale University Press, New RIBAS,C.C.AND C. Y. MIYAKI. 2004. Molecular sys- Haven, Connecticut. tematics in Aratinga parakeets: species limits and KAHN, F. 1988. Ecology of economically important historical biogeography in the `solstitialis' group, palms in the Peruvian Amazon. Pages 42±49 in and the systematic position of Nandayus nenday. The palm±tree of life: biology, utilization, and Molecular Phylogenetics and Evolution 30:663± conservation (M. J. Balick, Ed.). New York Bo- 675. tanical Garden, Bronx, New York. ROMO, M. 1997. Seasonal variation in fruit consump- MARTIN,K.AND J. M. EADIE. 1999. Nest webs: a com- tion and seed dispersal by canopy bats (Artibeus munity-wide approach to the management and spp.) in a lowland forest in PeruÂ. Vida Silvestre conservation of cavity-nesting forest birds. Forest Neotropical 5:110±119. Ecology and Management 115:243±257. ROTH, P. 1984. ReparticËaÄo do habitat entre psitacõÂdeos MASELLO,J.F.AND P. Q UILLFELDT. 2002. Chick growth simpaÂtricos no sul da AmazoÃnia. Acta Amazonica and breeding success of the . 14:175±221. Condor 104:574±586. RUSSELLO,M.A.AND G. AMATO. 2003. A molecular MAWSON,P.R.AND J. L. LONG. 1994. Size and age phylogeny of Amazona: implication for Neotrop- parameter of nest trees used by four species of ical parrot biogeography, and conser- parrot and one species of cockatoo in southwest vation. Molecular Phylogenetics and Evolution Australia. Emu 94:149±155. 30:421±437. MONTAMBAULT, J. R. 2002. Informes de las evalua- SALAMAN, P., A. CORTEÂ S,P.FLOREZ,J.C.LUNA,O. ciones bioloÂgicas Pampas del Heath, PeruÂ; Alto NIETO,J.F.CASTANÄ O, AND G. SUAREÂ Z. 2001. Proy- Madidi, ; y Pando, Bolivia. RAP Bulletin ecto Ognorhynchus progress report IV. www. 24. Conservation International, Washington, ognorhynchus.com/report4.htm (accessed August D.C. www.biodiversityscience.org/xp/CABS/ 2003). publications/cabs࿞publ࿞research/rap࿞bulletins/ TAVARES, E. S., C. YAMASHITA, AND C. Y. MIYAKI. terrestrial࿞rap࿞bulletins/terrrapbulletins.xml (ac- 2004. Phylogenetic relationships among some cessed May 2004). genera (Psittacidae) based on MONTERRUBIO,T.AND E. C. ENKERLIN. 2004. Present mitochondrial sequences. Auk 121:230±242. use and characteristics of Thick-billed Parrot nest TERBORGH, J. 1983. Five New World primates: a study sites in northwestern Mexico. Journal of Field Or- in comparative ecology. Princeton University nithology 75:96±103. Press, Princeton, New Jersey. Brightsmith • COMMUNITY-WIDE PARROT NESTING IN PERU 305

TERBORGH, J., J. W. FITZPATRICK, AND L. H. EMMONS. dent contrasts. Biological Conservation 86:243± 1984. Annotated checklist of birds and mammals 255. species of Cocha Cashu Biological Station, Manu WOOD FLOORING INTERNATIONAL. 2003. Species hard- National Park, Peru. Fieldiana Zoology 21:1±29. ness. www.w¯ooring.com/Technical࿞Info/Species࿞ ࿞ ࿞ TOLEDO,E.AND C. RINCOÂ N. 1999. UtilizacioÂn indus- Tech Info / species hardness. htm (accessed May trial de nuevas especies forestales en el PeruÂ. Ca- 2004). mara Nacional Forestal, Lima, Peru. YAMASHITA, C. 1987. Field observations and com- VAN SCHAIK, C. P., J. TERBORGH, AND S. J. WRIGHT. ments on the Indigo Macaw ( 1993. The phenology of tropical forests: adaptive leari), a highly endangered species from north- eastern Brazil. Wilson Bulletin 99:280±282. signi®cance and consequences for primary con- YAMASHITA,C.AND Y. M . DE BARROS. 1997. The Blue- sumers. Annual Review of Ecology and System- throated Macaw Ara glaucogularis: characteriza- atics 24:353±377. tion of its distinctive habitats in savannahs of the VAÂ SQUEZ,R.AND A. H. GENTRY. 1989. Use and misuse Beni, Bolivia. Ararajuba 5:141±150. of forest-harvested fruits in the Iquitos area. Con- YAMASHITA,C.AND M. DE P. V ALLE. 1993. On the servation Biology 3:350±361. linkage between Anodorhynchus macaws and WOLF, C. M., T. GARLAND, AND B. GRIFFITH. 1998. palm nuts, and the extinction of the Glaucous Ma- Predictors of avian and mammalian translocation caw. Bulletin of the British Ornithological Club success: reanalysis with phylogenetically indepen- 113:53±59.