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Bird Conservation International (2003) 13:319–349. BirdLife International 2003DOI: 10.1017/S0959270903003241 Printed in the United Kingdom
Avifauna of the Pilon Lajas BiosphereReserve and Communal Lands, BoliviaA. BENNETT HENNESSEY, SEBASTIAN A. HERZOG, MICHAEL KESSLERand DANIEL ROBISON
Summary
Pilon Lajas Biosphere Reserve and Communal Lands (Pilon Lajas) ranges from 300 m to1,400 m altitude in the departments of Beni and La Paz, Bolivia. It is located in the Bolivianand Peruvian lower yungas Endemic Bird Area (054) and within one of the regions ofhighest conservation priority in the Neotropics. Pilon Lajas includes primary evergreentropical lowland forest, foothill forest, and lower montane forest. Vegetation heterogeneityforms the basis for the high species richness of the area. A population of approximately600 Chiman, Moseten and Tacana indigenous people live within the reserve. Little hasbeen published regarding the avifauna of Pilon Lajas. Here we present the results of visualobservations and tape recordings of 22 ornithological surveys, recording 502 bird species.Serranıa Pilon is the most intensively studied and species-rich mountain range with 332species, 48 of which have been recorded only in this area within Pilon Lajas. We present26 new departmental records and evidence of breeding for 42 species. The reserve protectsthree globally Vulnerable and three globally Near Threatened species, four species ofHigh Conservation Priority and 32 of Medium Conservation Priority. Together withMadidi National Park and the adjacent Bahuaja Sonene National Park in south-easternPeru, Pilon Lajas forms a unique protected area system of more than 3.5 million ha andplays an important role in the establishment of biological corridors along the easternAndean slope in Bolivia.
Introduction
Pilon Lajas Biosphere Reserve and Communal Lands (Reserva de la Biosfera –Tierra Comunitaria de Orıgen Pilon Lajas in Spanish, hereafter referred to as PilonLajas) covers c. 400,000 ha on the Andean foothills in north-west Bolivia (Figure1). It is located in the Bolivian and Peruvian lower yungas Endemic Bird Area(EBA 054) of Stattersfield et al. (1998) and within one of the regions of highestconservation priority in the Neotropics (Stotz et al. 1996). Initially proposed in1974, Pilon Lajas was recognized as a Biosphere Reserve by UNESCO in 1977 andas a Biosphere Reserve and Indigenous Territory by the Bolivian Government in1992 (Sherholtz 1997). The area was legally designated but had no physical pro-tection until 1994, when Veterinarians Without Frontiers (VSF) received fundingfrom the European Union to design an operational plan for the reserve. Throughan agreement with the Bolivian National Protected Areas Service, VSF has beenco-administering Pilon Lajas since 1996.
Pilon Lajas is in an area with a moderate level of historic and current humanactivity and settlement. A population of approximately 600 Chiman, Moseten
A.B. Hennessey et al. 320
Figure 1. Map showing Pilon Lajas Biosphere Reserve, Bolivia.
and Tacana indigenous people live within the reserve. The Beni River, the west-ern border of Pilon Lajas (Figure 1), was the principal ‘‘highway’’ between theAndes and the rubber/brazil nut forests to the north for a century until a newroad was completed in 1989. This new road, forming Pilon Lajas’ eastern borderand connecting the area to the city of La Paz, was used for a government-sponsored colonization programme of families from other parts of the country.Many of these colonists believe they have a right to land within Pilon Lajas.Although parts of the reserve are extremely remote, requiring expeditions ofseveral days to be reached, other parts have long been accessible. It is estimatedthat only 1% of Pilon Lajas has been deforested for agriculture in the last century,but perhaps 80% of the lower tropical forest has been heavily hunted during thealligator hide and wildlife skin boom in the 1960s and 1970s. Between 1980 and1996, the mahogany lumber boom heavily affected the same area, particularlywithin 10 km of rivers and large streams. Up until 1996, five lumber companieswith dubious legal status and approximately 300 chain sawyers selectivelylogged and hunted in the reserve.
The Pilon Lajas management plan has subdivided the reserve into specificzones to combine indigenous community rights with conservation initiatives.Since 1996, a core of 20 park guards (many belonging to local indigenous groups)has been hired and trained and four park guard stations have been establishedin highly sensitive areas along borders to protect the reserve.
Avifauna of the Pilon Lajas Biosphere Reserve 321
Table 1. Survey areas visited by 22 ornithological expeditions in Pilon Lajas, Bolivia, between 1989and 1999.
Survey area Elevational range (m) Number of visits
Lowland forest areas 350–500 5Serranıa Bala 450–800 2Serranıa Beu 500–1,400 4Serranıa Cuchilla 450–1,000 2Serranıa Pilon 400–1,100 8Serranıa Zaguacales 400–800 1
The aim of this paper is to bring together unpublished avifaunal informationinto an extensive document that can be used for informed conservation decisionmaking and land protection priority assessments. This is important for the pro-tection and global attention of Pilon Lajas and the general Andean protected areacorridor of Madidi and Bahuaja Sonene National Parks.
Methods
This paper is based on 22 unpublished ornithological surveys by 15 researchersover 148 days between 1989 and 1999 (Table 1, Appendix 1) in Pilon Lajas. Theprincipal survey methods were non-systematic visual and acoustic observationsas well as tape recordings of vocalisations. Limited mist-netting was carried outby the first author (ABH) in Serranıa Cuchilla in 1999 and by F. Osorio andothers (Coleccion Boliviana de Fauna, La Paz; Appendix 1). A small number ofspecimens were collected by F. Osorio and others and deposited at the ColeccionBoliviana de Fauna.
Altitude ranges and abundance estimates are based on survey results by ABH.ABH surveyed different points at least 200 m apart, arriving before sunrise andidentifying and sound-recording vocalizations of the dawn chorus. Thereafter,different trails were surveyed, covering 1–3 km, often pausing work between12h00 and 15h00. ABH observed, sound-recorded and verified identificationswith the use of prerecorded tapes or playback, including rebound playback(broadcasting the first response to playback). Each evening ABH completed achecklist of the day’s observations, noting estimated abundances, field time, dis-tance walked and evidence (sight records, heard or tape-recorded). Abundanceestimates were classified as; common, recorded (visual observation or auditory)daily in small numbers (> 5); fairly common, recorded daily in small numbers(< 5); uncommon, recorded once in three days; and rare, recorded once duringthe survey. Tape-recording bird vocalizations is the most effective method forrapidly surveying neotropical forest avifaunas (Parker 1991; see also Haselmayerand Quinn 2000) and is an accurate means of species identification that can beeasily referenced. Over 400 recordings by ABH from Pilon Lajas have been arch-ived at the Cornell Laboratory of Ornithology, Macaulay Library of NaturalSounds (MLNS). Fieldwork covered all months except for January, March andMay, and included all habitat types within Pilon Lajas and as much area aspossible with the largest gap between sites being 15 km. For the purpose ofvegetation classification, we used the elevational belts of Stotz et al. (1996): lower
A.B. Hennessey et al. 322
tropical forest (< 500 m), hill tropical forest (500–900 m) and upper tropical forest(900–1,600 m). This was modified as necessary by reference to topographical loca-tion (level areas, slopes, ridges, etc.), geological substrate and state of vegetationsuccession. It should be noted that Parker (1989) erroneously gave 1,200 m as thehighest altitude of the road crossing Serranıa Pilon south of Yucumo (Figure 1).Rather, as indicated by Parker et al. (1991) and as confirmed by our measure-ments, the highest point of this road is located at c. 1,000 m.
Study Area
Pilon Lajas is situated at 14°26′ to 15°28′S and 68°42′ to 68°02′W in Beni and La Pazdepartments with an altitudinal range from 300 m to 1,400 m, although there prob-ably are a few peaks up to 1,600 m (detailed topographical maps of the area areunavailable). These mountain ranges are relatively isolated from themain Andeanridges. The nearest range of similar altitude is about 35 km to the south-west andthe closest peaks above 1,400 m are about 75 km to the south of Pilon Lajas.
The vegetation of Pilon Lajas is quite varied, changing in relation to humidity,elevation and geological substrate. At Rurrenabaque (14°26′S 68°31′W, 227 m),just north of Pilon Lajas (Figure 1), mean annual precipitation is 2,150 mm witha marked peak from November to April and a mean annual temperature of25.8 °C. The natural vegetation in about 38% of the reserve corresponds to ever-green tropical lowland forest. Higher elevations in Pilon Lajas, particularly thoseexposed to incoming clouds from the north-east, are somewhat cooler, receiveconsiderably more precipitation and, above c. 1,000 m, are frequently shroudedin mist. Here, forests are covered in epiphytic mosses, liverworts, ferns, brome-liads and orchids, and trees are often stunted.
Geologically, the area consists mainly of schists, loams and white sandstonesof Tertiary age. Flat valley bottoms with mixed alluvial, nutrient-rich soils andgood water supply are periodically flooded in the rainy season and support 30–40 m tall forest with abundant palms (e.g. Iriartea deltoidea, Socratea exorrhiza), figsand leguminaceous trees. Gentle to fairly steep slopes are covered with forest ofsomewhat lower stature with up to 150 tree species and about 200 species ofvascular epiphytes per ha (Seidel 1995, Smith and Killeen 1998, T. Kromer pers.com.). Above about 1,000 m the tree flora is dominated by the families Lauraceae,Leguminosae and Melastomataceae. Locally, there are dense stands of mahoganySwietenia macrophylla, many of which have been exploited in the last decades.Areas with white sand soils are very poor in nutrients and support a distinctand quite species-rich flora (Kessler 2001). Steep slopes, particularly at higherelevations, are characterized by unstable soils and frequent landslides and thusoften support vegetation in different stages of succession, ranging from low,dense thickets to young, 10–15 m tall forest with many slender trees. The highestridges often have shallow, leached soils and stunted, species-poor forest.
The overall picture emerging from the above combination of ecological factorsis a striking heterogeneity of the vegetation that is conspicuous in the field(particularly from the air) and that forms the basis for the high biological speciesrichness of the area. Botanically, individual vegetation types are of about averagediversity compared with other Neotropical areas, but the high number of distinctvegetation types leads to an exceedingly diverse overall flora that contains
Avifauna of the Pilon Lajas Biosphere Reserve 323
numerous elements not known elsewhere in the Bolivian Andes (Smith and Kil-leen 1998, Kessler 2001).
Results
Currently, 502 bird species are known from Pilon Lajas (Appendix 2), more thanone third of the 1,398 species reported in Bolivia (Hennessey et al. 2003). Webelieve that this includes at least 90% of the resident bird species in the reserve.Whereas 309 species (62%) were recorded in the approximately 150,000 ha oflower tropical forest in Pilon Lajas, only 79 species were exclusively found inthis zone. The lower tropical forest of Pilon Lajas appears quite species poor incomparison with other nearby lowland localities such as Alto Madidi,departamento (dpto.) La Paz, with 405 species (Parker and Bailey 1991) and the5,000 ha Tambopata-Candamo Reserved Zone, Peru, with 572 species (Parker etal. 1994). This relatively low diversity may well be a result of the geographicalposition of Pilon Lajas on the southern edge of the Amazon basin where tropicallowland forest is limited to a narrow, 10–30 km wide belt along the Andeanfoothills.
Serranıa Pilon is the most intensively studied and species-rich range withinPilon Lajas with 332 species, 48 of which have only been recorded in this area.Abundances of birds appear to be higher on Serranıa Pilon than on comparablenearby ranges. Daily mixed-species flock encounters on serranıas Bala and Cuch-illa (Figure 1) as well as on Serranıa Sadiri in Bolivia’s Madidi National Park(ABH pers. obs.) ranged between one and three per day, whereas encounter rateson Serranıa Pilon were between three and five per day. This is probably due toa higher level of precipitation in the area and the unusual topography of SerranıaPilon with several peaks, opposed to a typical single-peaked range, creating morehabitat area between 600 m and 900 m.
Upper tropical forest exists in Pilon Lajas only on serranıas Bue, Chepete andMuchanes. All surveys in this habitat were restricted to Serranıa Beu, where 246bird species have been recorded, 46 of which were only found in upper tropicalforest. Serranıa Beu is the only area in Pilon Lajas with large patches of bambooforest between 1,200 to 1,400 m; they cover an area of approximately 300 ha.
With respect to Arribas et al. (1995) and Remsen and Traylor (1989), the PilonLajas bird list includes 26 new records for dpto. Beni and three for dpto. La Paz(Appendix 2). However, these new records by no means represent true rangeextensions for the respective species. Bolivian departments are formed by arbit-rary political borders that have little or no value in a habitat or conservationcontext. For example, about 85% of the area of the department of Beni is coveredby pampas (savannah grasslands) whereas less than 1% contains hill tropicalforest, from where most of our new records originated.
Pilon Lajas is located approximately 25 km south of a large expanse of pampas,and its northern and eastern border is formed by a main road. Through the years,the area has seen the establishment of pampas bird species as colonists havecleared forest. In Rurrenabaque, each year more open-country species areobserved as the 30 km wide forest corridor between the town and the truepampas is being settled and destroyed. With continued forest clearing activitiesan increasing number of open-country species and austral migrants can be
A.B. Hennessey et al. 324
expected in open areas of Pilon Lajas with many of the former probably becom-ing resident along the edge of the reserve.
The majority of birds in Pilon Lajas increase their territorial singing in Sep-tember/October. The full onset of the breeding season is linked with the begin-ning of the rainy season in November. During the rainy season there is a markedincrease in insect abundance and more trees bear fruit. Local hunters claim thatlarge mammals have more fat from February to April. We found reproductiveevidence for 42 species (Appendix 2). Nest-building activity was observedbetween late July and mid-November (18 species), nesting activity (egg andhatchling stages) from mid-September to early November (nine species) andrecent fledglings (15 species) between early October and mid-February (with twoexceptions in mid-July).
Mixed-species flocks
ABH observed 45 mixed-species foraging flocks containing a total of 102 speciesin hill tropical forest (500–900 m). In the tropical lowlands, foraging flocks areoften strictly separated into canopy and understorey flocks (e.g. Munn 1985, Jul-lien and Thiollay 1998). Due to the variable nature of hill tropical forest withsteep slopes, landslides, cliffs, forest edges, stunted forest and tall forest, flockformations in Pilon Lajas were loosely categorized into three types: omnivoroustanager flocks, mid-canopy and understorey insectivorous mixed-species flocks.
The tanager canopy flocks were fast-moving, omnivorous flocks of the uppercanopy based around Tangara species with Paradise Tanager Tangara chilensis asthe nuclear species and Green-and-gold Tanager T. schrankii, Turquoise TanagerT. mexicana and Blue Dacnis Dacnis cayana as other characteristic species. Mid-canopy insectivorous flocks ranged from c. 5 m above ground to well inside thehighest canopy. These flocks were slower moving and sometimes large (c. 40individuals) with most species foraging for insects. Characteristic species wereSlate-throated Redstart Myioborus miniatus, Black-spotted Barbet Capito niger,Masked Tityra Tityra semifasciata, Guira Tanager Hemithraupis guira, Bronze-greenEuphonia Euphonia mesochrysa, Bay-headed Tanager Tangara gyrola and TropicalParula Parula pitiayumi. The nuclear species of understorey insectivorous flockswas Red-crowned Ant Tanager Habia rubica, with Buff-throated Foliage GleanerAutomolus ochrolaemus, Carmiol’s Tanager Chlorothraupis carmioli, Chestnut-tailedAntbird Myrmeciza hemimelaena, McConnell’s Flycatcher Mionectes macconnelliand Tawny-crowned Greenlet Hylophilus ochraceiceps as other characteristic spe-cies.
However, the majority of species in these flocks in hill tropical forest were notexclusive or even particular to any of the three flock types. This strongly contrastswith the rigidly organized flocks of Amazonian lowlands (Munn 1985, Jullienand Thiollay 1998) and supports the notion that conclusions on mixed-speciesflock structure and organization drawn from Neotropical lowland forests cannotbe generalized to all tropical forests (Jullien and Thiollay 1998, Herzog et al.submitted MS).
Species Accounts
Orinoco Goose Neochen jubataThis Near Threatened species has been observed only once in Pilon Lajas at an
Avifauna of the Pilon Lajas Biosphere Reserve 325
unknown date in 1994 by F. Osorio and others, although indigenous people saythe bird is sporadically seen on Rio Quiquebey. If a population still exists inPilon Lajas, it will be in isolated regions away from hunters.
Harpy Eagle Harpia harpyja
This globally Near Threatened species has never been sighted in Pilon Lajas bytrained observers. However, the Chiman and Moseten indigenous people arefamiliar with the bird and call it Bonij, but they do not have a name for thesimilar Crested Eagle Morphnus guianensis. ABH was shown a photo of a shotHarpy Eagle taken by an unknown tourist on the shore of the Beni River within Pilon Lajas around 1970, proving that it did exist in the area, but the exactlocation of the dead bird was unclear. A Moseten elder reported that a family ofHarpy Eagles exists east of S. Beu and described the species accurately as he saidthey ate one bird five years ago while hunting for White-bellied Spider MonkeysAteles belzebuth. A Chiman hunter reported the location of a nest in Pilon Lajassouth-west of the mouth of the Suapi River. We believe that a handful of HarpyEagles still exists within Pilon Lajas, but that this population is threatened andrestricted to areas with low human impact.
Southern Horned Curassow Pauxi unicornis
Parker (1989) gave the following account of this globally Vulnerable species froma 1989 survey in Serranıa Pilon along the Caranavi-Yucumo road at the south-eastern border of Pilon Lajas: ‘‘A man who has lived for one year near the roadpass on the Serranıa del Pilon accurately described this species and was certainthat he had seen (eaten?) it in ridgetop forest several times. Called ‘‘mutum’’.’’This record would appear to be a communication error as the name mutun loc-ally refers to Razor-billed Curassow Mitu tuberosa.
None of the 22 expeditions in Pilon Lajas, including two to inaccessible, pris-tine and apparently suitable habitat for this species between serranıas Beu andChepete, has resulted in a single observation of this species. Likewise, numerousinterviews including a survey of local indigenous knowledge by ABH and A.Perry have failed to produce any indication that the species has ever occurred inthe reserve. We believe that Pauxi unicornis does not exist in Pilon Lajas.
Oilbird Steatornis caripensis
Oilbird is a gregarious species, breeding colonially in caves. A single cave canhold up to 1,000 individuals (Thomas 1999; Duguid in litt. 2000). Only eight cavesites are known in Bolivia (ABH pers. obs.). Oilbirds could be threatened byhabitat loss and scarcity of isolated breeding areas. On 4 November 1998, ABHand park guards observed 350 individuals leaving an inaccessible cave at 600 mon the edge of Serranıa Chepete, beside the Beni River, during sunset and foundfresh eggshells below the entrance to the cave.
Lanceolated Monklet Micromonacha lanceolata
One immature and two adults were observed for approximately 15 min on 9December 1995 by ABH and A. Perry in the subcanopy 8 m above ground at
A.B. Hennessey et al. 326
1,150 m on Serranıa Beu, dpto. La Paz. One adult fed a small unidentified insectto the immature and then perched 30 cm away while the second adult landedbeside the immature and gave it a katydid-like insect longer than its bill. Theimmature clasped the insect in its bill for approximately 2 min before swallowingit. The immature’s breast had less clearly defined black streaks and greyer mot-tling than both adults. These birds were observed on a steep forested slope withmany successional stages from frequent landslides to dense forest patches withyoung trees. Another individual was observed in Pilon Lajas by A. Perry and R.Mitchell (pers. comm.) on the eastern edge of Serranıa Beu at 1,000 m on 3December 1996. These are the second and third observation of Lanceolated Mon-klet in Bolivia (Arribas et al. 1995). The first record was obtained by S. Cardiffand J.V. Remsen (in litt. 1998) at Cerro Asunta Pata (15°03′S 68°28′W), dpto. LaPaz, in 1993. The species was also observed in Carrasco National Park, dpto.Cochabamba, in July 1998 by A. Mee (in litt. 1998).
Scaled Fruiteater Ampelioides tschudii
Two birds, possibly a pair were noted repeatedly on Serranıa Cuchilla, dpto.Beni, at 900 m in 1998 and 1999, especially by voice (MLNS 101799, 101819,101874, 101877; Mayer 2000). On 17 November 1999 a male was observed byABH and M. Olivera feeding an immature an orange fruit of approximately 2.5cm in diameter and regurgitated half bill-sized black fruits. Previously known inBolivia only from Serranıa Bellavista (38 km N Caranavi), dpto. La Paz (Parkeret al. 1980), and Rıo Tuichi Valley, dpto. La Paz (Perry et al. 1997). ABH alsorecorded calls of one individual (MLNS 101845) at 900 m on Serranıa Sadiri(14°13′S 67°56′W) in Madidi National Park on 30 November 1999.
Subtropical Pygmy Owl Glacidium parkeri
C. Konig identified the first record of this species for Bolivia through recordingsmade by T.A. Parker during a 1979 expedition by the Museum of Natural Sci-ence, Louisiana State University, Baton Rouge, to Serranıa Bellavista, dpto. LaPaz (T.S. Schulenburg pers. comm.). B.M. Whitney (in litt. 2001) obtained thesecond record for Bolivia on Serranıa Eslabon in Madidi National Park in August1995 and the third in Pilon Lajas on Serranıa Pilon at 700 m on 9 August 1997.Subsequently the species was recorded by ABH (MLNS 101656, 101689) severaltimes on both the western (1,300 m) and eastern (1,200 m) slope of Serranıa Beuand once at 750 m on Serranıa Tequeje, Madidi National Park, on 6 April 2000.
Bolivian Recurvebill Simoxenops striatus
Until recently, this globally Vulnerable species was known from only three Boliv-ian localities (Parker et al. 1992), and Parker et al. (1996) considered it a highconservation and research priority. ABH observed and recorded several indi-viduals on Serranıa Beu at 800 m to 1,250 m from 17 to 24 September 1998 (MLNS101637) and on Serranıa Cuchilla at 800 m between 16 and 19 October 1998(MLNS 101778; Mayer 2000), which represent two new localities for the speciesin Bolivia. Additionally, it has been found at three other new sites in dptos. La
Avifauna of the Pilon Lajas Biosphere Reserve 327
Paz, Cochabamba and Santa Cruz by SKH and MK in 1996 and 1997. Furtherdetails will be published elsewhere.
Yungas Antwren Myrmotherula grisea
A single female-plumaged bird of this globally Vulnerable Bolivian endemic wasseen by SKH in an understorey mixed-species flock at c. 850 m on Serranıa Pilonon 4 August 1997, which represents the only record for Pilon Lajas. The bird wascharacterized by unmarked wing-coverts and warm olivish-brown upperpartswith a slight rufous tinge on wings and tail (in contrast to the all-grey upperpartsof the female Grey Antwren M. menetriesii) and it resembled a female YungasAntwren seen and tape-recorded by SKH together with a male (both singing) inAmboro National Park (dpto. Santa Cruz). Despite the presence of apparentlysuitable habitat within the species’s elevational range, Yungas Antwren appearsto be rare in Pilon Lajas.
Phyllomyias sp.
This undescribed species was first observed and tape-recorded in June 1989 at850–900 m on Serranıa Pilon along the Caranavi–Yucumo road (Parker et al.1991), who tentatively identified the birds as Planalto Tyrannulets Phyllomyiasfasciatus. However, they noted that their vocalizations were faster and higherpitched than those of birds in south-east Brazil and north-east Argentina andconcluded that ‘‘the Serranıa Pilon population may represent an undescribedform’’. On 2 June 1997 SKH tape-recorded three unidentified Phyllomyias tyran-nulets in the canopy of humid forest edge at 1,150 m at Cerro Asunta Pata, dpto.La Paz (15°03′S 68°28′W), and this recording was later identified with the helpof B.M. Whitney as belonging to the same undescribed Phyllomyias taxon asrecorded by Parker and Gell-Mann on Serranıa Pilon (see Mayer 2000). On 3September 1998, SKH and J.A. Balderrama collected two specimens at CerroAsunta Pata and rediscovered the Serranıa Pilon population at 850–1,000 m alongthe Caranavi–Yucumo road on 8 October 1998. Several tape-recordings from bothlocalities show that the vocal differences noted by Parker et al. (1991) betweenAndean and south-east Brazilian populations are both consistent and significant(recordings of P. fasciatus from south-east Brazil provided by J. Mazar Barnett). Aformal description of the species is currently in preparation (Herzog unpublisheddata).
It is noteworthy that despite the presence of apparently suitable hill tropicalforest habitat on Serranıas Beu, Cuchilla and Pilon the species has so far onlybeen found at a single locality in Pilon Lajas, indicating that it may have a natur-ally patchy distribution and a small range.
Shrike-like Cotinga Laniisoma elegans
This globally Vulnerable species was observed by ABH on 22 September 1999 15m above the ground on the northern side of Serranıa Beu in steep-sloped hilltropical forest (850 m). This is only the second observation for Bolivia of thispoorly known species (Hennessey et al. 2003).
A.B. Hennessey et al. 328
Scarlet Tanager Piranga olivacea
This Nearctic-Neotropical migrant has suffered population declines attributedto habitat loss in its wintering range (McNair and Escobar 1993). However, itsnon-breeding distribution in South America is poorly known (Parker et al. 1996).ABH observed this species on Serranıa Pilon in November 1996 and February1997 and 1998, on Serranıa Cuchilla in November 1999, and on Serranıa Sadiriin Madidi National Park in November 1999. Wintering males were identified bytheir contrasting black wings in November, and by the splotchy breeding plum-age in February. The identification of females was assumed through their pres-ence with Scarlet Tanager males, which in February consisted of associatinggroups of 5–8 individuals. The species was fairly common in all years and areas.For example, on a total of seven days on Serranıa Pilon during the months statedabove, ABH daily observed between two and seven individuals in non-breedingplumage in mid-canopy foraging flocks of hill tropical forest (600–900 m). Webelieve these records from serranıas Pilon, Cuchilla and Sadiri indicate that aconsiderable portion of the species’ population winters along the humid foothillsof the Bolivian Andes. This habitat is still largely unaltered in Bolivia; hence, adecline due to habitat loss is probably largely caused by the alteration of habitatin the breeding range or along the migration route of Scarlet Tanager.
Buff-rumped Warber Basileuterus fulvicauda
J. Ohlson (pers comm.) observed several individuals on Serranıa Bala from 25 to28 April 1999 and recorded one (Mayer 2000). Observations and recordings(MLNS 87814 and 101816) by ABH and SKH on serranıas Pilon and Cuchillahave exclusively been of the similar River Warbler Basileuterus rivularis. The songof both species is difficult to differentiate. Thus, both species occur within atmost 30 km of each other in Pilon Lajas, but the actual contact zone as well asthe degree of sympatry and hybridization (if any) remain to be determined.
Discussion
Recorded from Pilon Lajas are four species of Bolivian endemics, eight species ofrange-restricted endemics (Stattersfield et al. 1998), 28 zoogeographical regionalendemics (Stotz et al. 1996), three Vulnerable and three Near Threatened species(BirdLife International 2000), and four High Conservation Priority and 32Medium Conservation Priority species (Parker et al. 1996) (Appendix 2, Table 2).Of the High and Medium Conservation Priority species, eight have their centreof abundance in lower tropical (< 500 m), nine in hill tropical (500–900 m), 17 inupper tropical (900–1,600 m) and two in middle tropical (1,600–2,600 m) forest.All High Conservation Priority species in Pilon Lajas appear to be rare, with onlyone record per species, except for Simoxenops striatus. These apparently extremelylow population densities imply that optimal conditions for these species may notbe met in Pilon Lajas despite the presence of apparently suitable habitat and thatthe reserve alone cannot warrant their effective conservation in Bolivia.
Pilon Lajas protects a healthy part of the Bolivian and Peruvian lower YungasEBA (054) (Stattersfield et al. 1998). Eight range-restricted species have been
Avifauna of the Pilon Lajas Biosphere Reserve 329
Table 2. Species in Pilon Lajas, Bolivia, listed as High and Medium Conservation Priorities by Parkeret al. (1996) and Vulnerable (*) or Near Threatened (**) by BirdLife International (2000). Researchpriority and centre of abundance after Parker et al. (1996). Numbers in parentheses refer to thenumber of records.
Species Conservation Research Centre of Pilon Lajaspriority priority abundance abundance
Orinoco Goose**Neochen jubata High High Lower tropical Very rare (1)
Fasciated Tiger Heron Medium High Hill tropical Very rare (2)Tigrisoma fasciatum
Solitary Eagle Medium Medium Upper tropical Very rare (2)Harpyhaliaetus solitarius
Harpy Eagle** Medium High Lower tropical Very rare (?)Harpia harpyja
Red-throated Caracara Medium Medium Lower tropical Fairly commonIbycter americanus
Maroon-chested Ground Dove Medium Medium Middle montane Very rare (1)Claravis mondetoura
Blue-and-yellow Macaw Medium Medium Lower tropical Fairly commonAra ararauna
Military Macaw* Medium Medium Upper tropical UncommonAra militaris
Red-billed Parrot Medium Medium Upper tropical UncommonPionus sordidus
Rufous-vented Ground Cuckoo Medium Medium Lower tropical Very rare (1)Neomorphus geoffroyi
Band-bellied Owl Medium Medium Hill tropical Fairly commonPulsatrix melanota
Silky-tailed Nightjar Medium Medium Lower tropical Very rare (1)Caprimulgus sericocaudatus
White-browed Hermit Medium Medium Hill tropical Very rare (1)Phaethornis stuarti
Rufous-crested Coquette Medium Medium Upper tropical Very rare (1)Lophornis delattrei
Many-spotted Hummingbird Medium Medium Hill tropical Very rare (1)Taphrospilus hypostictus
Crested Quetzal Medium Medium Upper tropical Fairly commonPharomachrus antisianus
Ash-browed Spinetail Medium Medium Upper tropical RareCranioleuca curtata
Bolivian Recurvebill* High High Hill tropical UncommonSimoxenops striatus
Yungas Antwren* High High Hill tropical Very rare (1)Myrmotherula grisea
Rough-legged Tyrannulet Medium Medium Hill tropical UncommonPhyllomyias burmeisteri
Spectacled Bristle Tyrant Medium Medium Hill tropical UncommonPhylloscartes orbitalis
White-bellied Pygmy Tyrant Medium Medium Hill tropical Fairly commonMyiornis albiventris
Yungas Tody Tyrant Medium Medium Upper tropical CommonHemitriccus spodiops
Buff-throated Tody Tyrant** Medium Medium Upper tropical RareHemitriccus rufigularis
Shrike-like Cotinga High High Upper tropical Very rare (1)Laniisoma elegans
A.B. Hennessey et al. 330
Table 2. Continued.
Species Conservation Research Centre of Pilon Lajaspriority priority abundance abundance
Scaled Fruiteater Medium Medium Upper tropical UncommonAmpelioides tschudii
Andean Cock-of-the-Rock Medium Medium Upper tropical Fairly commonRupicola peruviana
Orange-eared Tanager Medium Medium Upper tropical Very rare (1)Chlorochrysa calliparaea
Pale-winged Trumpeter Medium Low Lower tropical RarePsophia leucoptera
Red-and-green Macaw Medium Low Lower tropical Fairly commonAra chloroptera
Upland Antshrike Medium Low Upper tropical Fairly commonThamnophilus aroyae
Scaled Antpitta Medium Low Upper tropical RareGrallaria guatimalensis
Slaty Gnateater Medium Low Upper tropical UncommonConopophaga ardesiaca
Slaty Tanager Medium Low Middle montane Very rare (1)Creurgops dentata
Bronze-green Euphonia Medium Low Upper tropical CommonEuphonia mesochrysa
Golden-eared Tanager Medium Medium Upper tropical Very rare (2)Tangara chrysotis
recorded of a potential 13 species found in Bolivia. The range-restricted endem-ism of Serranıa Beu demonstrated the area to be a potential Important Bird Area.Serranıa Beu and the surrounding area are sufficiently difficult to access throughclimbing, long hikes and the lack of road access, that the areas may be easilyprotected from habitat destruction through local population pressure.
Serranıa Pilon is an important area in Pilon Lajas to protect foothill-specializedspecies. The foothill forest, though not considered specialization enough todeserve EBA status, is a unique ecosystem that deserves conservation attention.Through high levels of precipitation and the mix between lowland and Yungasspecies, these areas tend to be the richest sites of avian diversity on the planet.With no avifaunal data from Serranıa del Tigre (13°38′S 68°40′W), mostly outsidethe Madidi National Park border, Serranıa Pilon is the richest area of foothillforest in Bolivia. The site, with a main road travelling through the area, alsocontains ecotourism-sustainable development possibilities.
Pilon Lajas has common borders with Madidi National Park to the north-westand with the Moseten Indigenous Area, which applies sustainable forestry prac-tices, to the south. However, the north- and south-eastern borders of Pilon Lajasare adjacent to major roads that have enabled human settlement on the parkboundaries, posing a serious future threat to the reserve. Pilon Lajas is trying tocounteract this encroachment with two park guard stations along both roadsand the implementation of public education programmes. Recently a sustainableecotourism project has been designed with a local indigenous group at the mouthof Rio Quiquebey.
Together with Madidi National Park and the directly adjacent Bahuaja Sonene
Avifauna of the Pilon Lajas Biosphere Reserve 331
National Park in south-eastern Peru, Pilon Lajas forms part of a unique protectedarea system of more than 3.5 million ha. It further represents a vital link towardsfoothill and montane forest areas further south and plays an important role in theestablishment of biological corridors along the eastern Andean slope in Bolivia.
Acknowledgements
We would like to thank Veterinarians Without Frontiers and the British, Dutchand Swiss Governments for their support and efforts to protect this special areaof the earth and to support our insistence for more field investigations. ABHwould like to especially thank Sjoerd Mayer for his meticulous review of PilonLajas sound recordings, the use of his pre-release CD-ROM Birds of Bolivia 2,and the hundreds of email messages on ‘‘mystery’’ sounds, queries and birdsound discussions. We are grateful to Bret M. Whitney for his assistance withPilon Lajas research and for his time and patience with his remarkable identifica-tions of ‘‘mystery’’ sounds that no one else could identify. ABH is most appreci-ative for the donations of tapes and equipment by Greg Budney at Cornell’sMacaulay Library of Natural Sounds. In addition, we thank Doug Stotz, TomSchulenberg and Jon Fjeldså for their informative answers to the many questionsinvolving Pilon Lajas research. Financial support was provided to ABH by Veter-inarians Without Frontiers as well as the British, Dutch and Swiss Governments,and to MK by the Deutsche Forschungsgemeinschaft.
Appendix 1. Ornithological surveys in Pilon Lajas, Bolivia.
Researchers Site Date Period(days)
T.A. Parker S. Pilon (600–1,000 m) June 1989 4F. Osorio, S. Barrera, Quiquibey and Suapi rivers and June 1993 10J.F. Guerra, J. Colorado (350–500 m)Sarmiento, L. VillalbaABH Suapi River (450 m) November 1995 4A. Perry, ABH S. Beu (western end, 700–1,300 m) December 1995 7F. Sagot Colorado (350–400 m) June 1996 6ABH, R. Silva Colorado to Quiquibey (350–400 m) July 1996 7ABH S. Bala (400–700 m) October 1996 3ABH, R. Mitchell S. Bala (500–900 m) November 1996 4ABH S. Pilon (600–1,000 m) November 1996 7A. Perry, R. Mitchell S. Zaguacales (350–700 m) November 1996 6A. Perry, R. Mitchell S. Beu (eastern end, 600–1,400 m) December 1996 11ABH S. Pilon (700–750 m) February 1997 1ABH S. Pilon (500–1,000 m) April 1997 5ABH S. Pilon (600–1,000 m) June 1997 7SKH, MK S. Pilon (450–1,000 m) July 1997 18B.M. Whitney, ABH S. Pilon (400–750 m) August 1997 3ABH S. Pilon (350–1,000 m) February 1998 11ABH S. Beu (western end, 1,200–1,450 m) September 1998 7ABH S. Beu (western end, 800–1,000 m) September 1998 5SKH, J.A. Balderrama S. Pilon (450–500 m) October 1998 4ABH S. Cuchilla (500–900 m) October 1998 6ABH S. Cuchilla (700–900 m) November 1999 12
A.B. Hennessey et al. 332
Ap
pen
dix
2:L
ist
ofB
ird
spec
ies
from
Pil
onL
ajas
Bio
sph
ere
Res
erve
and
Com
mu
nal
Lan
ds,
Dep
artm
ents
ofB
eni
and
La
Paz
,B
oliv
ia,i
ncl
ud
ing
evid
ence
,si
tes,
min
imu
man
dm
axim
um
elev
atio
nal
reco
rds
and
abu
nd
ance
esti
mat
es.
Nom
enclaturean
dtaxo
nomic
arrang
emen
taccord
withRem
senan
dTay
lor(1
989)
andArribas
etal
.(19
95).
Spec
ies:*,Zoo
geog
raph
ical
endem
ic(Stotz
etal
.199
6);**,Ran
gerestricted
endem
ic(Stattersfi
eld
etal
.199
8).***,P
olitical
endem
ic.
Evi
denc
e(inorder
ofpriority):SP
,spe
cimen
(Coleccion
Boliviana
deFa
una);T,soun
drecord
ing(C
orne
llLibrary
ofNatural
Soun
ds);O,v
isua
lob
servation;
A,
auditory;
L,local
inform
ation.
Site
s:1,
trop
ical
forest;2
,SerranıaZag
uacales;
3,S.
Bala;
4,S.
Cuc
hilla
;5,S
.Pilo
n;6S.
Beu
.M
in:M
inim
umelev
ation,
L,L
owland
(350
–500
m).
Max:M
axim
umelev
ation,
The
high
estviab
leforest
habitatin
Pilo
nLajas
is14
00m.
Abu
ndan
ce:C,common
,record
ed(visua
lob
servationor
auditory)
daily
insm
allnu
mbe
rs(>
5);F,
fairly
common
,record
eddaily
insm
allnu
mbe
rs(<
5);U,
uncommon
,recorded
once
in3day
s;R,rare,
record
edon
cein
6day
sor
less,V
R,recorded
4or
less
times
withinPilo
nLajas,n
umbe
rof
record
sin
quotes;M
n,Northernmigrant;M
s,So
uthe
rnmigrant;M
m;m
igrant
from
theAnd
es;+
,inc
ludingresiden
tpo
pulation
.B
reed
ing
info
rmat
ion:
NA,a
tten
dingne
st;N
B,b
uildingne
st;N
E,n
estwitheg
gs;N
M,c
ollectingne
stmaterial;F,
fled
gling.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Tin
amus
taoGreyTinam
ouT
1–6
L1,
000
CT
inam
usm
ajor
Great
Tinam
ouT
1–6
L60
0F
NE
17–1
0–98
Cry
ptur
ellu
sci
nere
usCinereo
usTinam
ouA
1,2,
6L
900
RC
rypt
urel
lus
soui
LittleTinam
ouT
1–6
L90
0C
Cry
ptur
ellu
sob
sole
tusBrownTinam
ouT
1,3–
6L
1,30
0C
Cry
ptur
ellu
sun
dula
tusUnd
ulated
Tinam
ouA
1,2,
4–6
L80
0C
Tac
hyba
ptus
dom
inic
usLeast
Grebe
O1
L35
0VR(1Mn)
Pha
lacr
ocor
axbr
asili
anus
Neo
trop
icCormoran
tO
1,2,
6L
500
FA
nhin
gaan
hing
aAnh
inga
O1
L50
0F
Tig
riso
ma
linea
tum
Rufescent
Tiger
Heron
O1,
3,5,
6L
700
UT
igri
som
afa
scia
tum
FasciatedTiger
Heron
O1,
4L
600
VR(2)
Pilh
erod
ius
pile
atus
Cap
pedHeron
O1
L40
0U
Ard
eaco
coiCocoi
Heron
O1
L35
0U
Ard
eaal
baGreat
Egret
O1
L35
0F
Bul
bucu
sib
isCattleEgret
O1
L40
0U
Egr
etta
thul
aSn
owyEgret
O1
L35
0F
Egr
etta
caer
ulea
LittleBlueHeron
O2
400
400
VR(1)
But
orid
esst
riat
usStriated
Heron
O1
350
350
VR(1)
Aga
mia
agam
iChe
stnu
t-be
llied
Heron
O2
350
350
VR(1)
Avifauna of the Pilon Lajas Biosphere Reserve 333
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Mes
embr
inib
isca
yenn
ensi
sGreen
Ibis
O1
L40
0R
Myc
teri
aam
eric
anaWoo
dStork
O1
L35
0VR(1)
Jari
bum
ycte
riaJabiru
O1
L35
0VR(1)
Cor
agyp
sat
ratu
sBlack
Vulture
O1,
3–5
L50
0C
Cat
hart
esau
raTurke
yVulture
O1,
3,5,
6L
900
CC
atha
rtes
burr
ovia
nusLesserYellow-headed
Vulture
O1
L35
0R
Cat
hart
esm
elam
brot
usGreater
Yellow-headed
Vulture
O1–
3,6
L35
0R
Sarc
oram
pus
papa
KingVulture
O1,
4,5
L60
0U
Cha
una
torq
uata
Southe
rnScream
erO
1L
350
VR(2)
Neo
chen
juba
taOrino
coGoo
seO
135
035
0VR(1)
Cai
rina
mos
chat
aMus
covy
Duc
kO
1,6
L60
0VR(3)
Mer
gane
tta
arm
ataTorrent
Duc
kO
670
070
0VR(1Mm)
Pan
dion
halia
etus
Osp
rey
O1
L40
0VR(1Mn)
Cho
ndro
hier
axun
cina
tusHoo
k-bille
dKite
O1,
3–6
L90
0U
Ela
noid
esfo
rfica
tusSw
allow-tailedKite
O1,
3,5,
6L
1,40
0F(+Mn)
Har
pagu
sbi
dent
atus
Dou
ble-toothe
dKite
T1,
3,5
L80
0R
Icti
nia
plum
beaPlumbe
ousKite
O4,
6L
1,40
0VR(2)
Acc
ipit
ersu
perc
ilios
usTinyHaw
kT
1,3,
5,6
L1,
250
VR(4)
F10
–10–
96A
ccip
iter
bico
lorBicolored
Haw
kO
1L
350
VR(1)
Leuc
opte
rnis
albi
colli
sWhite
Haw
kO
1,3–
5L
700
UB
uteo
gallu
sur
ubit
inga
Great
Black
-haw
kO
1,6
L60
0VR(3)
Har
pyha
liaet
usso
litar
iusSo
litaryEag
leO
5L
900
VR(2)
But
eom
agni
rost
risRoa
dsideHaw
kO
1,3,
5L
800
FB
uteo
brac
hyur
usSh
ort-taile
dHaw
kO
1,3,
5L
700
RB
uteo
albi
caud
atus
White-tailedHaw
kO
5L
600
VR(2)
Har
pia
harp
yjaHarpy
Eag
leL
1,6
L1,
000
VR(?)
Spiz
aetu
sty
rann
usBlack
Haw
kEag
leO
580
080
0VR(1)
Spiz
aetu
sor
natu
sOrnateHaw
kEag
leO
1,5
L1,
000
VR(2)
Dap
triu
sat
erBlack
Caracara
O1–
2L
500
UIb
ycte
ram
eric
anus
Red
-throa
tedCaracara
T1–
4,6
L70
0F
Her
peto
ther
esca
chin
nans
Lau
ghingFa
lcon
A1,
4L
500
U
A.B. Hennessey et al. 334
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Mic
rast
urru
ficol
lisBarredFo
rest
Falcon
T1,
4–6
L1,
350
FM
icra
stur
gilv
icol
lisLined
Forest
Falcon
T4–
670
01,
200
UM
icra
stur
mir
ando
lleiSlaty-ba
cked
Forest
Falcon
T1,
2,5
L70
0U
Mic
rast
urse
mit
orqu
atus
CollaredFo
rest
Falcon
T1,
3,5
L90
0U
Falc
oru
figul
arisBat
Falcon
O1,
3,5
L80
0U
Ort
alis
gutt
ataSp
eckled
Cha
chalaca
O1,
2,5
L50
0C
Pen
elop
eja
cqua
cuSp
ix’s
Gua
nT
1–6
L1,
300
FP
ipile
pipi
leBlue-throated
PipingGua
nO
1,2,
4–6
L80
0U
Mit
utu
bero
saRazor-billed
Curassow
T1–
6L
1,00
0F
Odo
ntop
horu
sgu
jane
nsisMarbled
Woo
dQua
ilT
5–6
L60
0U
Odo
ntop
horu
ssp
ecio
susRufou
s-breasted
Woo
dQua
ilT
6L
1,00
0R
Odo
ntop
horu
sba
llivi
aniStripe
-faced
Woo
dQua
il*,**
T6
1,30
01,
400
UF
15–0
9–98
Odo
ntop
horu
sst
ella
tusStarredWoo
dQua
ilA
1L
?LVR(1)
Ara
mid
esca
jane
aGrey-ne
cked
Woo
dRail
O1,
2,5,
6L
700
FE
uryp
yga
helia
sSu
nbittern
O1,
2,5
L80
0R
Pso
phia
leuc
opte
raPale-winge
dTrumpe
ter*
T2,
4–6
500
1,00
0R
Van
ellu
sca
yanu
sPiedLap
wing
O1
L50
0U
Cha
radr
ius
colla
risCollaredPlove
rO
1,2
L50
0F
Act
itis
mac
ular
iaSp
ottedSa
ndpipe
rO
1,2,
5L
500
FB
artr
amia
long
icau
daUplan
dSa
ndpipe
rO
61,
300
1,30
0VR(1Mn)
Pha
etus
asi
mpl
exLarge
-billed
Tern
O1
L50
0U
Ster
nasu
perc
iliar
isYellow-billed
Tern
O1
L50
0U
Ryn
chop
sni
gerBlack
Skim
mer
O1
L50
0U
Col
umba
spec
iosa
Scaled
Pigeo
nT
1–6
L1,
400
UC
olum
bapl
umbe
aPlumbe
ousPigeo
nT
1–6
L1,
400
CC
olum
basu
bvin
acea
Rud
dyPigeo
nO
1,2,
5L
800
UC
olum
bina
picu
iPicui
Groun
dDov
eO
1,5
L50
0C
Col
umbi
nata
lpac
otiR
uddyGroun
dDov
eO
1,5
L50
0C
Cla
ravi
spr
etio
saBlueGroun
dDov
eT
1,3,
6L
500
CC
lara
vis
mon
deto
uraMaroo
n-ch
estedGroun
dDov
eO
380
080
0VR(1)
Lept
otila
verr
eaux
iWhite-tippe
dDov
eT
2,5
L60
0U
Avifauna of the Pilon Lajas Biosphere Reserve 335
Ap
pen
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2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Lept
otila
rufa
xilla
Greyfron
tedDov
eT
1–6
L1,
000
CG
eotr
ygon
fren
ataWhite-throa
tedQua
ilDov
eT
61,
000
1,40
0F
Geo
tryg
onm
onta
naRud
dyQua
ilDov
eO
1,2,
6L
700
RA
raar
aura
naBlue-an
d-yellow
Macaw
O1–
4L
600
FA
ram
ilita
risMilitary
Macaw
T5,
670
01,
400
U-R
Ara
chlo
ropt
eraRed
-and
-green
Macaw
T1–
6L
900
FNA
08–1
0–96
Ara
seve
raChe
stnu
t-fron
tedMacaw
O1–
3,5
L60
0C
Ara
man
ilata
Red
-bellie
dMacaw
O1,
2L
900
RA
rati
nga
leuc
opht
halm
usWhite-eye
dParak
eet
O1–
6L
700
CA
rati
nga
mit
rata
MitredParak
eet
T4,
590
01,
400
FA
rati
nga
wed
delli
iDus
ky-headed
Parak
eet
O1–
6L
600
CP
yrrh
ura
mol
inae
Green
-che
eked
Parak
eet
T6
1,20
01,
400
FP
yrrh
ura
pict
aPainted
Parak
eet
T1,
4,5
L90
0U
Bro
toge
ris
cyan
opte
raCob
alt-winge
dParak
eet
O1–
3L
600
FP
ioni
tes
leuc
ogas
terWhite-bellie
dParrot*
O1,
2L
400
UP
iono
psit
taba
rrab
andi
Orang
e-ch
eeke
dParrot
O1
L50
0VR(1)
Pio
nus
men
stru
usBlue-he
aded
Parrot
T1–
6L
1,00
0F
Pio
nus
sord
idus
Red
-billed
Parrot
O5
L80
0U
Am
azon
afa
rino
saMealy
Parrot
T1–
6L
1,00
0C
Pia
yaca
yana
Squirrel
Cuc
koo
T1–
6L
1,40
0F
Cro
toph
aga
aniSm
ooth-billed
Ani
O1,
4,5
L50
0C
Tap
era
naev
iaStripe
dCuc
koo
O1
450
450
VR(1)
Dro
moc
occy
xph
asia
nellu
sPhe
asan
tCuc
koo
O1
L45
0R
Dro
moc
occy
xpa
voni
nusPav
onineCuc
koo
O1,
5L
500
RN
eom
orph
usge
offr
oyiRufou
s-ve
nted
Groun
dCuc
koo
O5
700
700
VR(1)
Otu
sgu
atem
alae
Vermiculated
ScreechOwl
T4–
660
080
0F
Otu
sch
olib
aTropicalScreechOwl
O1,
2L
500
FO
tus
inge
nsRufescent
ScreechOwl
T1,
4–6
900
1,30
0F
Otu
sw
atso
niiTaw
ny-bellie
dScreechOwl
T1,
2,5,
6L
900
FLo
phos
trix
cris
tata
Crested
Owl
O1–
6L
600
FP
ulsa
trix
pers
pici
llata
Spectacled
Owl
O1,
3,5
L60
0F
A.B. Hennessey et al. 336
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Pul
satr
ixm
elan
otaBan
d-bellie
dOwl
T4–
670
01,
200
FG
lauc
idiu
mha
rdyi
Amazon
ianPyg
myOwl
T1,
2,5
L70
0U
Gla
ucid
ium
park
eriSu
btropicalPyg
myOwl
T5,
670
01,
200
UG
lauc
idiu
mbr
asili
anum
Ferrug
inou
sPyg
myOwl
T1
L60
0F
Cic
caba
virg
ataMottled
Owl
T1,
5L
700
UC
icca
bahu
hula
Black-ban
ded
Owl
T1,
2,4–
6L
1,00
0F
Stea
torn
isca
ripe
nsisOilb
ird
T6
400
400
CN
ycti
bius
gran
disGreat
Potoo
T1,
2,6
L80
0R
Nyc
tibi
usae
ther
eusLon
g-taile
dPotoo
T1,
5L
900
RN
ycti
bius
gris
eusCom
mon
Potoo
O1
L50
0R
Cho
rdei
les
rupe
stri
sSa
nd-colored
Nightha
wk
O1
L50
0R
Nyc
tidr
omus
albi
colli
sParau
que
O1,
4L
500
RN
ycti
phry
nus
ocel
latu
sOcella
tedPoo
rwill
T5
L90
0R
Cap
rim
ulgu
sru
fusRufou
sNightjar
O5
L90
0VR(2Ms)
Cap
rim
ulgu
sse
rico
caud
atus
Silky-taile
dNightjar
O1
L50
0VR(1)
Hyd
rops
alis
clim
acoc
erca
Lad
der-tailedNightjar
SP1
L50
0VR(2)
Stre
ptop
rocn
eru
tila
Che
stnu
t-colla
redSw
ift
O1
L50
0VR(1)
Stre
ptop
rocn
ezo
nari
sWhite-collaredSw
ift
O1–
6L
1,40
0C
Cha
etur
aci
nere
iven
tris
Grey-rumpe
dSw
ift
O1,
5L
900
UC
haet
ura
brac
hyur
aSh
ort-taile
dSw
ift
O1,
5L
1,40
0F-U
Thr
enet
esle
ucur
usPale-taile
dBarbthroa
tO
1,4,
5L
900
RP
haet
horn
ism
alar
isLon
g-taile
dHermit
T1,
4–6
L1,
300
FP
haet
horn
ishi
spid
usWhite-beard
edHermit
O1,
3–6
L80
0U
Pha
etho
rnis
stua
rtiWhite-browed
Hermit*
O5
700
700
VR(1)
Pha
etho
rnis
rube
rRed
dishHermit
T1,
3–6
L1,
400
CC
ampy
lopt
erus
larg
ipen
nisGrey-breasted
Sabrew
ing
O1,
3–6
L1,
300
UFl
oris
uga
mel
livor
aWhite-necke
dJacobin
O1,
5–6
L85
0U
Ant
hrac
otho
rax
nigr
icol
lisBlack-throa
tedMan
goO
1,6
L90
0R
Kla
isgu
imet
iViolet-he
aded
Hum
mingb
ird
T3–
570
01,
000
ULo
phor
nis
dela
ttre
iRufou
s-crestedCoq
uette
O5
900
900
VR(1)
Loph
orni
sch
alyb
eusFe
stiveCoq
uette
O1,
5L
600
F
Avifauna of the Pilon Lajas Biosphere Reserve 337
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Chl
oros
tilb
onm
ellis
ugus
Blue-taile
dEmerald
O1,
6L
1,20
0R
Tha
lura
nia
furc
ataFo
rk-tailedWoo
dny
mph
O1,
5–6
L1,
400
UH
yloc
hari
scy
anus
White-chinn
edSa
pphire
O1,
5L
700
RC
hrys
uron
iaoe
none
Golden
-tailedSa
pphire
T1,
4–5
L90
0F
Tap
hros
pilu
shy
post
ictu
sMan
y-sp
ottedHum
mingb
ird
O5
875
875
VR(1)
Am
azili
ave
rsic
olor
VersicoloredEmerald
O1,
5L
900
UA
maz
ilia
lact
eaSa
pphire-spa
ngledEmerald
O5
875
875
VR(1)
Ade
lom
yia
mel
anog
enys
Speckled
Hum
mingb
ird
O6
1,20
01,
450
FP
olyp
lanc
taau
resc
ensGou
ld’s
Jewelfron
tT
1,4,
5L
900
UH
elio
thry
xau
rita
Black-eared
Fairy
O5,
687
590
0VR(2)
Cal
liphl
oxam
ethy
stin
aAmethy
stW
oodstar
O1,
5L
800
UP
haro
mac
hrus
anti
sian
usCrested
Que
tzal
T4,
690
01,
400
FT
rogo
nm
elan
urus
Black-tailedTrogo
nT
1–6
L1,
000
FT
rogo
nvi
ridi
sWhite-tailedTrogo
nT
2,5,
6L
700
UT
rogo
nco
llari
sCollaredTrogo
nT
1,2,
4,5
L1,
200
UT
rogo
ncu
rucu
iBlue-crow
nedTrogo
nT
1–6
L1,
400
FT
rogo
nvi
olac
eusViolaceou
sTrogo
nO
1L
450
UE
lect
ron
plat
yrhy
nchu
mBroad
-billed
Motmot
O1–
6L
1,00
0U
Bar
ypht
heng
usm
arti
iRufou
sMotmot
T1–
3,5,
665
01,
000
F-U
Mom
otus
mom
otaBlue-crow
nedMotmot
T1–
3,5
L50
0F
Cer
yle
torq
uata
Ringe
dKingfi
sher
O1,
2,4,
5L
600
FC
hlor
ocer
yle
amaz
onaAmazon
Kingfi
sher
T1,
2,5
L50
0U
Chl
oroc
eryl
eam
eric
anaGreen
Kingfi
sher
O1,
5L
500
UC
hlor
ocer
yle
inda
Green
-and
-rufou
sKingfi
sher
O2
L50
0R
Chl
oroc
eryl
eae
neaPyg
myKingfi
sher
O1,
6L
500
RN
otha
rchu
sm
acro
rhyn
chus
White-necke
dPuffbird
T4–
660
090
0U
Buc
com
acro
dact
ylus
Che
stnu
t-capp
edPuffbird
O1
L45
0VR(1)
Nys
talu
sst
riol
atus
Striolated
Puffbird
T1,
3–6
L80
0U
Mic
rom
onac
hala
nceo
lata
Lan
ceolated
Mon
klet
O6
1,15
01,
200
VR(2)
Mon
asa
nigr
ifron
sBlack-fronted
Nun
bird
O1,
2L
500
FM
onas
am
orph
oeus
White-fronted
Nun
bird
O1,
2,5,
6L
500
F
A.B. Hennessey et al. 338
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Che
lidop
tera
tene
bros
aSw
allow-w
ing
O1,
4,6
L50
0U
Gal
bula
rufic
auda
Rufou
s-taile
dJacamar
O1,
2,5,
6L
500
FC
apit
oni
gerBlack-spo
tted
Barbe
tT
1–6
L95
0C
Eub
ucco
rich
ards
oniLem
on-throa
tedBarbe
tO
1,4–
6L
850
RE
ubuc
cove
rsic
olor
VersicoloredBarbe
t*
T6
1,20
01,
400
FA
ulac
orhy
nchu
spr
asin
usEmeraldTou
cane
tT
4–6
L1,
000
FA
ulac
orhy
nchu
sde
rbia
nusChe
stnu
t-tipp
edTou
cane
tT
5,6
800
1,40
0F
Pte
rogl
ossu
sin
scri
ptus
LetteredAracari
O6
800
800
VR(1)
Pte
rogl
ossu
sfla
viro
stri
sIvory-bille
dAracari
O1,
5,6
L1,
000
UP
tero
glos
sus
cast
anot
isChe
stnu
t-earedAracari
T1,
2,5,
6L
1,00
0C
Pte
rogl
ossu
sbe
auha
rnae
siiCurl-crestedAracari
*T
1,5
L80
0R
Sele
nide
rare
inw
ardt
iiGolden
-collaredTou
cane
tT
1,2,
4–6
L80
0R
Ram
phas
tos
vite
llinu
sCha
nnel-billed
Tou
can
T1–
6L
1,40
0C
Ram
phas
tos
tuca
nusRed
-billed
Tou
can
T1–
6L
1,00
0C
Pic
umnu
sm
inut
issi
mus
Arrow
head
Piculet
O4,
5L
800
RP
icum
nus
auri
fron
sBar-breastedPiculet
*T
4,5
L90
0R
Mel
aner
pes
crue
ntat
usYellow-tuftedW
oodpe
cker
T1–
6L
700
CV
enili
orni
sfu
mig
atus
Smok
y-brow
nWoo
dpe
cker
T6
1,30
01,
400
UV
enili
orni
spa
sser
inus
LittleWoo
dpe
cker
O1,
3,4
L80
0U
Ven
ilior
nis
affin
isRed
-stained
Woo
dpe
cker
O1,
540
01,
000
FP
icul
usle
ucol
aem
usWhite-throa
tedWoo
dpe
cker
T1,
3–6
L1,
000
FP
icul
usch
ryso
chlo
rosGolden
-green
Woo
dpe
cker
T5,
650
070
0VR(3)
Pic
ulus
rubi
gino
susGolden
-oliv
eWoo
dpe
cker
T5,
650
01,
400
UC
eleu
sgr
amm
icus
Scale-breasted
Woo
dpe
cker
T4
900
900
VR(1)
Cel
eus
flavu
sCream
-colou
redW
oodpe
cker
O1
L50
0R
Dry
ocop
uslin
eatu
sLineatedWoo
dpe
cker
O1,
550
01,
000
RC
ampe
philu
sm
elan
oleu
cosCrimson-crestedWoo
dpe
cker
O5
400
400
VR(1)
Cam
peph
ilus
rubr
icol
lisRed
-necke
dW
oodpe
cker
T1–
6L
1,30
0F
F08
–10–
96D
endr
ocin
cla
fulig
inos
aPlain-brownW
oodcreepe
rT
1,2,
4–6
L1,
000
FD
econ
ychu
ralo
ngic
auda
Lon
g-taile
dWoo
dcreepe
rT
1,5,
6L
900
USi
ttas
omus
gris
eica
pillu
sOliv
aceo
usW
oodcreepe
rT
1,3–
6L
1,30
0F
Avifauna of the Pilon Lajas Biosphere Reserve 339
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Den
drex
etas
tes
rufig
ulaCinna
mon
-throa
tedW
oodcreepe
rT
4,5
L80
0R
Xip
hoco
lapt
espr
omer
opir
hync
husStrong
-billed
Woo
dcr.
T1,
4–6
L90
0F
Den
droc
olap
tes
cert
hiaBarredWoo
dcreepe
rT
1,4
L70
0F
Den
droc
olap
tes
picu
mnu
sBlack-ban
ded
Woo
dcreepe
rT
4–6
L90
0U
Xip
horh
ynch
uspi
cusStraight-billed
Woo
dcreepe
rT
1,6
L80
0R
Xip
horh
ynch
usoc
ella
tusOcella
tedWoo
dcreepe
rT
1,3,
5L
900
UX
ipho
rhyn
chus
spix
iiSp
ix’s
Woo
dcreepe
rT
1–6
L90
0F
Xip
horh
ynch
usgu
ttat
usBuff-throated
Woo
dcreepe
rT
1–6
L1,
000
FLe
pido
cola
ptes
albo
linea
tusLineatedWoo
dcreepe
rT
1,5,
6L
900
UC
ampy
lorh
amph
ustr
ochi
liros
trisRed
-billed
Scythe
bill
SP1,
5,6
L1,
300
UFu
rnar
ius
leuc
opus
Pale-legg
edHorne
roO
1L
350
USy
nalla
xis
azar
aeAzara’s
Spinetail*
T6
1,30
01,
400
FSy
nalla
xis
caba
nisiCab
anis’S
pine
tail*
T5,
680
01,
300
FSy
nalla
xis
guja
nens
isPlain-crowne
dSp
inetail
T1,
2L
500
FC
rani
oleu
cacu
rtat
aAsh
-browed
Spinetail
T5,
670
090
0R
Cra
niol
euca
gutt
urat
aSp
eckled
Spinetail
O1
L50
0VR(1)
Pre
mno
plex
brun
nesc
ensSp
ottedBarbtail
T6
1,30
01,
300
VR(1)
Hyl
octi
stes
subu
latu
sStripe
dW
oodha
unter
O5,
670
01,
300
VR(3)
Anc
istr
ops
stri
gila
tusChe
stnu
t-winge
dHoo
kbill
O5
800
800
VR(1)
Sim
oxen
ops
stri
atus
BolivianRecurve
bill*,**,***
T4,
670
01,
300
U–R
Phi
lydo
rer
ythr
ocer
cusRufou
s-rumpe
dFo
liage
-glean
erO
3,4
600
900
VR(3)
Phi
lydo
rer
ythr
opte
rusChe
stnu
t-winge
dFo
liage
-glean
erO
570
085
0VR(2)
Aut
omol
usoc
hrol
aem
usBuff-throated
Folia
ge-glean
erT
1,2,
4–6
L1,
400
FX
enop
sru
tila
nsStreak
edXen
ops
O1,
5,6
500
700
UX
enop
sm
inut
usPlain
Xen
ops
SP1,
5,6
600
900
USc
leru
rus
albi
gula
risGrey-throated
Leaftosser
T3,
5,6
700
800
RSc
leru
rus
mex
ican
usTaw
ny-throa
tedLeaftosser
T4,
570
080
0U
Cym
bila
imus
linea
tusFa
sciatedAntsh
rike
O1,
4,5
L1,
000
FT
arab
am
ajor
Great
Antsh
rike
O1,
2,5,
6L
1,00
0C
Tha
mno
philu
spa
lliat
usChe
stnu
t-ba
cked
Antsh
rike
T6
1,20
01,
400
RT
ham
noph
ilus
aeth
iops
White-sho
uldered
Antsh
rike
T1,
4–6
L1,
000
F
A.B. Hennessey et al. 340
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Tha
mno
philu
ssc
hist
aceu
sPlain-w
inge
dAntsh
rike
T1–
6L
1,00
0C
Tha
mno
philu
sar
oyae
Uplan
dAntsh
rike
*,**
T5,
61,
200
1,40
0F
Tha
mno
philu
sca
erul
esce
nsVariableAntsh
rike
T6
1,30
01,
300
VR(1)
Dys
itha
mnu
sm
enta
lisPlain
Antvireo
T1,
3,5,
645
01,
000
FT
ham
nom
anes
schi
stog
ynus
Bluish-slateAntsh
rike
*T
2,4,
560
01,
000
UM
yrm
othe
rula
brac
hyur
aPyg
myAntwren
T1,
4,5
L1,
000
FM
yrm
othe
rula
long
icau
daStripe
-che
sted
Antwren
O1,
560
090
0R
Myr
mot
heru
laha
uxw
elliPlain-throa
tedAntwren
O1
LL
VR(1)
Myr
mot
heru
lale
ucop
htha
lmaWhite-eye
dAntwren*
O5,
6L
900
VR(2)
Myr
mot
heru
laax
illar
isWhite-flan
kedAntwren
O1,
5.6
L1,
000
UM
yrm
othe
rula
gris
eaYun
gasAntwren*,**,***
O5
850
850
VR(1)
Myr
mot
heru
lam
enet
ries
iiGreyAntwren
T4,
5L
1,00
0F
Dic
hroz
ona
cinc
taBan
ded
Antbird
T4,
550
080
0U
Her
psilo
chm
usru
fimar
gina
tusRufou
s-winge
dAntwren
T2–
6L
1,00
0C-F
Dry
mop
hila
devi
lleiStriated
Antbird
T2,
5,6
700
1350
FC
erco
mac
raci
nera
scen
sGreyAntbird
T1,
2,4–
6L
1,00
0C
Cer
com
acra
nigr
esce
nsBlackishAntbird
O1
LL
RNM
07–1
0–98
Cer
com
acra
serv
aBlack
Antbird
T1,
4–6
L1,
000
UP
yrig
lena
leuc
onot
aWhite-backe
dFire-eye
T5,
670
01,
400
FM
yrm
obor
usle
ucop
hrys
White-browed
Antbird
O2
LL
RM
yrm
obor
usm
yoth
erin
usBlack-faced
Antbird
T1–
6L
1,40
0C
Hyp
ocne
mis
cant
ator
WarblingAntbird
T2,
5,6
L80
0C
Per
cnos
tola
leuc
osti
gmaSp
ot-w
inge
dAntbird
O1
LL
VR(2)
Myr
mec
iza
hem
imel
aena
Che
stnu
t-taile
dAntbird
T1,
2,4–
6L
1,20
0C
NE
21–0
9–97
Myr
mec
iza
atro
thor
axBlack-throa
tedAntbird
T1,
2,4–
6L
1,00
0C
Rhe
gmat
orhi
nam
elan
osti
ctaHairy-crested
Antbird
T4–
650
090
0R
Hyl
ophy
lax
naev
iaSp
ot-backe
dAntbird
SP1,
2,4–
6L
1,00
0U
Hyl
ophy
lax
poec
ilino
taScale-ba
cked
Antbird
O5
600
600
VR(1)
Form
icar
ius
colm
aRufou
s-capp
edAntthrush
T1
L50
0R
Form
icar
ius
anal
isBlack-faced
Antthrush
T1–
6L
1,00
0C
Cha
mae
zaca
mpa
niso
naSh
ort-taile
dAntthrush
T1,
5,6
1,20
01,
400
C
Avifauna of the Pilon Lajas Biosphere Reserve 341
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Gra
llari
agu
atim
alen
sisScaled
Antpitta
T5,
670
01,
000
RH
ylop
ezus
berl
epsc
hiAmazon
ianAntpitta*
O1
350
350
VR(1)
Con
opop
haga
arde
siac
aSlatyGna
teater
*T
61,
200
1,40
0U
Scyt
alop
usbo
livia
nusWhite-crowne
dTap
aculo
T6
1,20
01,
400
CP
hyllo
myi
asbu
rmei
ster
iRou
gh-leg
gedTyran
nulet
O5
800
1,00
0U?
Phy
llom
yias
scla
teriSclater’sTyran
nulet*
O5
480
1,20
0F
Phy
llom
yias
,und
escribed
species*,**,***
T1,
587
51,
000
U?
Zim
mer
ius
grac
ilipe
sSlen
der-foo
tedTyran
nulet
T1,
5,6
L1,
100
CO
rnit
hion
iner
meWhite-lored
Tyran
nulet
O1,
3–5
L90
0U
Cam
ptos
tom
aob
sole
tum
S.Beard
less
Tyran
nulet
O5
800
800
VR(1Ms)
Tyr
annu
lus
elat
usYellow-crowne
dTyran
nulet
O1
L50
0VR(1)
Myi
opag
isga
imar
diiFo
rest
Elaen
iaT
1,5
L95
0F
Myi
opag
isca
nice
psGreyElaen
iaO
4,5
L90
0R
Myi
opag
isvi
ridi
cata
Green
ishElaen
iaO
1L
500
?E
laen
iaal
bice
psWhite-crested
Elaen
iaT
5L
1,00
0R
Ela
enia
obsc
uraHighlan
dElaen
iaO
4,5
800
950
RSe
rpop
haga
cine
reaTorrent
Tyran
nulet
O6
1,10
01,
100
VR(1Mm)
Mio
nect
esst
riat
icol
lisStreak
-necke
dFlyc
atch
erT
61,
350
1,35
0VR(1Mm)
Mio
nect
esol
eagi
neus
Och
re-bellie
dFlyc
atch
erT
1,2,
5,6
L1,
100
FF
15–1
1–99
Mio
nect
esm
acco
nnel
liMcC
onne
ll’sFlyc
atch
erT
1,4–
6L
1,00
0U
Lept
opog
onam
auro
ceph
alus
Sepia-capp
edFlyc
atch
erT
1,5,
6L
1,00
0F
Lept
opog
onsu
perc
iliar
isSlaty-capp
edFlyc
atch
erT
5,6
850
1,30
0U
Phy
llosc
arte
sop
htha
lmic
usMarble-facedBristle
Tyran
tT
580
01,
000
RP
hyllo
scar
tes
orbi
talis
Spectacled
Bristle
Tyran
tO
5,6
700
1,00
0U
Phy
llosc
arte
sfla
vive
ntri
sRufou
s-loredTyran
nulet
O5
900
900
VR(1)
Cor
ytho
pis
torq
uata
Ringe
dAntpipit
T2,
5,6
L95
0U
Myi
orni
sal
bive
ntri
sWhite-bellie
dPyg
myTyran
t*
T2,
4–6
600
1,10
0F
Myi
orni
sec
auda
tusSh
ort-taile
dPyg
myTyran
tT
1,3–
6L
950
FNM
27–0
7–97
Hem
itri
ccus
flam
mul
atus
Flam
mulated
Tod
yTyran
t*
O5
L70
0R
Hem
itri
ccus
zost
erop
sWhite-eye
dTod
yTyran
tT
1,2,
5,6
L90
0F
Hem
itri
ccus
spod
iops
Yun
gasTod
yTyran
t*,**,***
T5,
61,
200
1,40
0C
A.B. Hennessey et al. 342
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Hem
itri
ccus
rufig
ular
isBuff-throated
Tod
yTyran
t*
T5,
680
01,
350
RT
odir
ostr
umpl
umbe
icep
sOch
re-faced
Tod
yFlyc
atch
erO
5L
500
UT
odir
ostr
umch
ryso
crot
aphu
mGolden
-browed
Tod
yFlyc
atch
erO
1,5,
60
1,00
0R
Ram
phot
rigo
nru
ficau
daRufou
s-taile
dFlatbill
O1
060
0VR(1)
Rhy
ncho
cycl
usol
ivac
eusOliv
aceo
usFlatbill
T1,
50
950
RT
olm
omyi
assu
lphu
resc
ensYellow-oliv
eFlyc
atch
erT
1,5,
6L
900
RNM
02–0
8–97
Tol
mom
yias
polio
ceph
alus
Grey-crow
nedFlyc
atch
erT
1,5
L?
?T
olm
omyi
asfla
vive
ntri
sYellow-breastedFlyc
atch
erT
480
080
0VR(2)
Pla
tyri
nchu
sco
rona
tusGolden
-crowne
dSp
adeb
illO
1,2
L60
0R
Ter
enot
ricc
user
ythr
urus
Rud
dy-taile
dFlyc
atch
erO
580
01,
100
RM
yiob
ius
villo
susTaw
ny-breastedFlyc
atch
erT
61,
200
1,35
0R
Myi
opho
bus
fasc
iatu
sBran-coloured
Flyc
atch
erT
1,5
L1,
000
FP
yrrh
omyi
asci
nnam
omea
Cinna
mon
Flyc
atch
erO
690
090
0VR(1Mm)
Con
topu
sfu
mig
atus
Smok
e-coloured
Pew
eeO
550
01,
000
RC
onto
pus
vire
nsEastern
Woo
d-Pew
eeT
2–5
L1,
000
C-F
(Mn)
Con
topu
sci
nere
usTropicalPew
eeO
5L
1,00
0R
Em
pido
nax
eule
riEuler’s
Flycatch
erT
1,4–
6L
1,00
0F
Cne
mot
ricc
usfu
scat
usFu
scou
sFlyc
atch
erT
560
060
0VR(1)
Sayo
rnis
nigr
ican
sBlack
Pho
ebe
O5,
6L
900
UP
yroc
epha
lus
rubi
nusVermilion
Flyc
atch
erO
1L
500
F(M
s)O
chth
oeca
litto
ralis
DrabWater
Tyran
tO
1,2,
5L
500
FM
usci
saxi
cola
fluvi
atili
sLittleGroun
dTyran
tO
1L
500
FK
nipo
legu
ssi
gnat
usAnd
eanTyran
t*
O5
1,20
01,
200
VR(1Mm)
Col
onia
colo
nusLon
g-taile
dTyran
tO
6L
500
RSa
trap
aic
tero
phry
sYellow-browed
Tyran
tO
544
044
0VR(1)
Att
ilabo
livia
nusWhite-eye
dAttila
T3,
445
070
0U
Att
ilasp
adic
eusBrigh
t-rumpe
dAttila
T1,
3–6
L1,
300
FF
21–1
1–99
Rhy
tipt
erna
sim
plex
Greyish
Mou
rner
T1–
6L
1,00
0F-U
Siry
stes
sibi
lato
rSirystes
O5
L70
0R
Myi
arch
ustu
berc
ulife
rDus
ky-cap
pedFlyc
atch
erT
1L
500
RM
yiar
chus
fero
xSh
ort-crestedFlyc
atch
erT
1,5
L70
0F
Avifauna of the Pilon Lajas Biosphere Reserve 343
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Myi
arch
usty
rann
ulus
Brown-crestedFlyc
atch
erT
1,5
L90
0U
Pit
angu
ssu
lphu
ratu
sGreat
Kiska
dee
O1
L50
0F
Meg
aryn
chus
pita
ngua
Boa
t-bille
dFlyc
atch
erO
1,5
L60
0F
Myi
ozet
etes
caya
nens
isRus
ty-m
argine
dFlyc
atch
erT
1,5
450
800
RM
yioz
etet
essi
mili
sSo
cial
Flycatch
erO
1,5
L70
0R
Myi
ozet
etes
lute
iven
trisDus
ky-che
sted
Flyc
atch
erO
1L
500
VR(1)
Con
opia
str
ivir
gata
Three-striped
Flyc
atch
erO
6L
900
RM
yiod
ynas
tes
chry
soce
phal
usGolden
-crowne
dFlyc
atch
erT
1,5
L1,
400
C-F
Myi
odyn
aste
sm
acul
atus
Streak
edFlyc
atch
erT
1–3,
5,6
L1,
000
FM
yiod
ynas
tes
lute
iven
trisSu
lphu
r-be
llied
Flyc
atch
erT
4–5
500
900
F(M
n)Le
gatu
sle
ucop
haiu
sPiratic
Flycatch
erT
1–6
L1,
000
FNA
12–1
0–96
Em
pido
nom
usva
rius
Variega
tedFlyc
atch
erO
360
060
0VR(1)
Em
pido
nom
usau
rant
ioat
rocr
ista
tusCrowne
dSlatyFlyc
atch
erO
570
070
0VR(1)
Tyr
annu
sal
bogu
lari
sWhite-throa
tedKingb
ird
O5,
670
095
0R
Tyr
annu
sm
elan
chol
icus
TropicalKingb
ird
T1,
3–6
L1,
200
CT
yran
nus
tyra
nnus
Eastern
Kingb
ird
O1,
4L
500
R(M
n)P
achy
ram
phus
cast
aneu
sChe
stnu
t-crow
nedBecard
O1
LL
VR(1)
Pac
hyra
mph
uspo
lych
opte
rusWhite-w
inge
dBecard
O1,
4,5
L80
0R
Pac
hyra
mph
usm
argi
natu
sBlack-cap
pedBecard
O1,
4–6
L1,
000
RP
achy
ram
phus
min
orPink-throated
Becard
T1,
3–5
L70
0U
NB
08–1
0–96
Pac
hyra
mph
usva
lidus
Crested
Becard
O1,
4L
1,00
0VR(2)
Tit
yra
caya
naBlack-tailedTityra
O1
LL
VR(1)
Tit
yra
sem
ifasc
iata
Maske
dTityra
T1–
6L
1,30
0F
NM
27–0
7–97
Lani
isom
ael
egan
sSh
rike
-likeCotinga
O6
850
850
VR(1)
Am
pelio
ides
tsch
udiiScaled
Fruiteater
T4
850
900
RLa
nioc
era
hypo
pyrr
aCinereo
usMou
rner
O1,
4L
500
RLi
paug
usvo
cife
rans
Scream
ingPiha
T1–
6L
900
CC
otin
gaca
yana
Span
gled
Cotinga
O1,
6L
800
VR(2)
Gym
node
rus
foet
idus
Bare-ne
cked
Fruitcrow
O1,
6L
600
VR(2)
Que
rula
purp
urat
aPurple-throated
Fruitcrow
T1,
3–6
L90
0F
Cep
halo
pter
usor
natu
sAmazon
ianUmbrellabird
O6
900
900
VR(1)
A.B. Hennessey et al. 344
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Rup
icol
ape
ruvi
anaAnd
eanCock-of-the
-Rock
T6
700
1,20
0U
Schi
fforn
istu
rdin
usThrus
h-lik
eMan
akin
T1,
2,4–
680
01,
400
FF
06–1
0–98
Pip
rite
sch
lori
sWing-ba
rred
Man
akin
T1–
660
01,
000
FT
yran
neut
esst
olzm
anni
DwarfTyran
tMan
akin
O1,
2,6
L80
0R
Mac
haer
opte
rus
pyro
ceph
alus
Fiery-capp
edMan
akin
SP1,
5L
400
VR(2)
Chi
roxi
phia
boliv
iana
Yun
gasMan
akin
*,**
T5,
61,
000
1,40
0C
Pip
raco
rona
taBlue-crow
nedMan
akin
O2,
660
01,
200
RP
ipra
fasc
iicau
daBan
d-tailedMan
akin
SP1,
2L
500
RP
ipra
chlo
rom
eros
Rou
nd-tailedMan
akin
T1–
6L
800
CT
achy
cine
taal
bive
nter
White-w
inge
dSw
allow
O1
L50
0C
Pro
gne
chal
ybea
Grey-breasted
Martin
O1
400
400
VR(1)
Not
ioch
elid
oncy
anol
euca
Blue-an
d-w
hite
Swallow
T3,
570
01,
400
FA
ttic
ora
fasc
iata
White-ban
ded
Swallow
O1,
2,4
L50
0C
Stel
gido
pter
yxru
ficol
lisS.
Rou
gh-w
inge
dSw
allow
O1–
5L
900
CC
ampy
lorh
ynch
ustu
rdin
usThrus
h-lik
eW
ren
T1,
3–5
L50
0F
Thr
yoth
orus
geni
barb
isMou
stache
dW
ren
T1–
6L
800
CT
hryo
thor
usgu
aray
anus
Fawn-breasted
Wren
T1
500
500
VR(1)
Tro
glod
ytes
aedo
nHou
seWren
T1,
3,5
L1,
300
FH
enic
orhi
nale
ucop
hrys
Gray-breasted
Woo
d-W
ren
T6
1,20
01,
400
CM
icro
cerc
ulus
mar
gina
tusNightinga
leW
ren
T1–
6L
900
CC
ypho
rhin
usar
adaMus
icianWren
T1
L40
0U
Don
acob
ius
atri
capi
llusBlack-cap
pedDon
acob
ius
O1,
5L
500
FC
incl
usle
ucoc
epha
lusWhite-cap
pedDippe
rO
670
070
0VR(1Mm)
Ram
phoc
aenu
sm
elan
urus
Lon
g-bille
dGna
twren
T1–
5L
900
FC
atha
rus
ustu
latu
sSw
ains
on’s
Thrus
hT
4–6
700
1,00
0F(M
n)P
laty
cich
lale
ucop
sPale-ey
edThrus
hT
61,
400
1,40
0VR(1)
Tur
dus
nigr
icep
sSlatyThrus
hO
5L
700
VR(2)
Tur
dus
amau
roch
alin
usCream
y-be
llied
Thrus
hT
1,3–
5L
700
FT
urdu
sig
nobi
lisBlack-billed
Thrus
hO
1,3–
5L
1,00
0F
Tur
dus
law
renc
iiLaw
renc
e’sThrus
hO
1,2
L50
0U
Tur
dus
haux
wel
liHau
xwell’s
Thrus
hO
1L
LR
Avifauna of the Pilon Lajas Biosphere Reserve 345
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Tur
dus
albi
colli
sWhite-necke
dThrus
hO
5L
900
FC
ycla
rhis
guja
nens
isRufou
s-brow
edPep
pershrike
T1,
3L
500
UV
ireo
lani
usle
ucot
isSlaty-capp
edSh
rike
-Vireo
T1,
3–6
L90
0U
Vir
eool
ivac
eusRed
-eye
dVireo
T1–
6L
1,00
0F
Vir
eofla
vovi
ridi
sYellow-green
Vireo
O5
700
900
R(M
n)V
ireo
leuc
ophr
ysBrown-capp
edVireo
O5
600
1,00
0R
Hyl
ophi
lus
thor
acic
usLem
on-che
sted
Green
let
O5,
660
01,
000
RH
ylop
hilu
shy
poxa
nthu
sDus
ky-cap
pedGreen
let
T1–
6L
900
CH
ylop
hilu
soc
hrac
eice
psTaw
ny-crowne
dGreen
let
T1–
6L
900
CA
mm
odra
mus
auri
fron
sYellow-browed
Sparrow
O1,
2,4–
6L
600
FC
yano
cora
xcy
anom
elas
PurplishJay
O1,
2,5
L70
0C
Cya
noco
rax
ynca
sGreen
Jay
T5,
61,
200
1,40
0U
Vol
atin
iaja
cari
naBlue-blackGrassqu
itO
1L
LVR(1Ms)
Spor
ophi
laco
llari
sRus
ty-collaredSe
edeater
O5
LL
VR(1Ms)
Spor
ophi
lalin
eola
Lined
Seed
eater*
O1
LL
VR(1Ms)
Spor
ophi
laca
erul
esce
nsDou
ble-colla
redSe
edeater
O1,
2,3
LL
R(M
s)Sp
orop
hila
cast
anei
vent
risChe
stnu
t-be
llied
Seed
eater
O1
LL
VR(2Ms)
Ory
zobo
rus
ango
lens
isLesserSe
edfinc
hO
5L
700
RA
rrem
onta
citu
rnus
PectoralSp
arrow
T1–
6L
1,00
0C
Par
oari
agu
lari
sRed
-cap
pedCardinal
O1
500
500
VR(1)
Phe
ucti
cus
aure
oven
tris
Black-backe
dGrosbeak
T1,
3,5,
6L
1,00
0C
Par
kert
hrau
stes
umer
alisYellow-sho
uldered
Grosbeak
T1,
4–6
L80
0R
Salt
ator
ross
usSlate-coloured
Grosbeak
T1,
4,5
L1,
400
RSa
ltat
orm
axim
usBuff-throated
Saltator
T1–
6L
1,40
0F
Salt
ator
coer
ules
cens
Greyish
Saltator
O1,
3,4
L60
0C
Cya
noco
mps
acy
anoi
desBlue-blackGrosbeak
T1–
6L
800
CC
yano
com
psa
bris
soni
iUltramarineGrosbeak
O3
700
700
VR(1)
Cis
sopi
sle
veri
anaMag
pieTan
ager
O1,
2,4–
60
1,30
0F
Chl
oros
ping
usop
htha
lmic
usCom
mon
Bus
hTan
ager
T5,
690
01,
450
CT
hlyp
opsi
sso
rdid
aOrang
e-he
aded
Tan
ager
O5
900
900
VR(1)
Hem
ithr
aupi
sgu
iraGuira
Tan
ager
T1,
5L
1,00
0C
A.B. Hennessey et al. 346
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Hem
ithr
aupi
sfla
vico
llisYellow-backe
dTan
ager
T1,
4–6
L1,
000
FN
emos
iapi
leat
aHoo
ded
Tan
ager
O1
L80
0R
Chl
orot
hrau
pis
carm
ioliCarmiol’s
Tan
ager
T1–
6L
1,20
0C
Lani
ove
rsic
olor
White-w
inge
dSh
rike
Tan
ager
*T
1,4–
6L
1,00
0F
Cre
urgo
psde
ntat
aSlatyTan
ager
*,**
O6
1,40
01,
400
VR(1)
Tac
hyph
onus
cris
tatu
sFlam
e-crestedTan
ager
O5
600
600
VR(1)
Tac
hyph
onus
rufiv
ente
rYellow-crested
Tan
ager
*O
1,4–
6L
1,00
0F
Tac
hyph
onus
luct
uosu
sWhite-sho
uldered
Tan
ager
T1,
3–6
L1,
000
FT
rich
othr
aupi
sm
elan
opsBlack-gog
gled
Tan
ager
O5,
690
01,
200
RH
abia
rubi
caRed
-crowne
dAnt-Tan
ager
T1–
6L
1,00
0C
Pir
anga
flava
Hep
atic
Tan
ager
T1,
3,5
700
1,00
0F
NA
15–1
1–99
Pir
anga
rubr
aSu
mmer
Tan
ager
O4,
5L
1,80
0U
(Mn)
Pir
anga
oliv
acea
ScarletTan
ager
O4,
570
01,
000
F(M
n)P
iran
gale
ucop
tera
White-w
inge
dTan
ager
T5,
670
01,
200
FR
amph
ocel
usca
rboSilver-beake
dTan
ager
T1–
6L
1,40
0C
Thr
aupi
sep
isco
pusBlue-grey
Tan
agar
O1,
4L
900
UT
hrau
pis
saya
caSa
yaca
Tan
ager
O1,
2,5,
6L
1,40
0C
Thr
aupi
spa
lmar
umPalm
Tan
ager
T1–
6L
1,40
0C
NB
20–1
1–99
Ani
sogn
athu
sfla
vinu
chaBlue-winge
dMou
ntainTan
ager
O6
1,30
01,
400
UE
upho
nia
chlo
roti
caPurple-throated
Eup
honia
T1,
3,5,
6L
700
CE
upho
nia
lani
iros
trisThick-billed
Eup
honia
T1,
3–6
L1,
200
FNM
20–1
1–99
Eup
honi
acy
anoc
epha
laBlue-ho
oded
Eup
honia
O1
LL
R(?Ms)
Eup
honi
ach
ryso
past
aWhite-lored
Eup
honia
T1,
4–5
L1,
000
UNM
02–0
8–97
Eup
honi
am
esoc
hrys
aBronz
e-greenEup
honia
T1,
4–6
600
1,20
0C
Eup
honi
am
inut
aWhite-ven
tedEup
honia
T1,
4L
1,00
0U
Eup
honi
axa
ntho
gast
erOrang
e-be
llied
Eup
honia
T4–
6L
1,40
0C
Eup
honi
aru
fiven
trisRufou
s-be
llied
Eup
honia
T1–
6L
1,00
0F
Chl
orop
honi
acy
anea
Blue-na
pedEup
honia
T1,
5,6
L1,
400
UC
hlor
ochr
ysa
calli
para
eaOrang
e-earedTan
ager
O6
1,30
01,
300
VR(1)
Tan
gara
mex
ican
aTurqu
oise
Tan
ager
T1,
4–6
L1,
300
CNM
06–1
0–98
Tan
gara
chile
nsisParad
iseTan
ager
T1,
3–6
L1,
300
CNM
17–1
1–99
Avifauna of the Pilon Lajas Biosphere Reserve 347
Ap
pen
dix
2:Con
tinu
ed.
Species
Eviden
ceSites
Min
Max
Abu
ndan
ceBreed
ing
Inform
ation
Tan
gara
schr
anki
iGreen
-and
-goldTan
ager
T1–
6L
1,30
0C
F05
–02–
98T
anga
raar
thus
Golden
Tan
ager
O5,
670
01,
400
RT
anga
raxa
ntho
ceph
alaSa
ffron-crow
nedTan
ager
O6
1,20
01,
400
UT
anga
raxa
ntho
gast
raYellow-bellie
dTan
ager
T1,
3–6
L1,
200
FT
anga
rach
ryso
tisGolden
-eared
Tan
ager
O6
1,20
01,
400
VR(2)
Tan
gara
gyro
laBay
-headed
Tan
ager
T1–
6L
1,20
0C
NM
17–1
1–99
Tan
gara
rufic
ervi
xGolden
-nap
edTan
ager
O6
1,30
01,
300
VR(1)
Tan
gara
cyan
icol
lisBlue-ne
cked
Tan
ager
O6
1,30
01,
350
VR(2)
Tan
gara
nigr
ocin
ctaMaske
dTan
ager
T1,
3–6
L1,
200
CF
16–1
1–99
Tan
gara
nigr
ovir
idisBeryl-spa
ngledTan
ager
O5
1,00
01,
000
VR(1)
Dac
nis
linea
taBlack-faced
Dacnis
T1,
3–6
L1,
250
FD
acni
sfla
vive
nter
Yellow-bellie
dDacnis
O1,
4,5
L80
0F
Dac
nis
caya
naBlueDacnis
T1,
3–6
L1,
300
CC
hlor
opha
nes
spiz
aGreen
Hon
eycreepe
rT
1,4–
60
1,40
0F
Cya
nerp
esca
erul
eusPurpleHon
eycreepe
rT
1.4–
60
1,00
0C
Cya
nerp
escy
aneu
sRed
-leg
gedHon
eycreepe
rT
1,5
01,
200
FF
07–0
2–98
Dig
loss
agl
auca
Deep-blue
Flow
erpiercer
O6
1,20
01,
200
VR(1)
Ter
sina
viri
disSw
allow
Tan
ager
T1,
3–6
L1,
400
UP
arul
api
tiay
umiTropicalParula
T1,
3–6
01,
300
CM
yiob
orus
min
iatu
sSlate-throated
Red
start
T2,
4–6
600
1,40
0C
NM
06–0
8–97
Bas
ileut
erus
bivi
ttat
usTwo-ba
nded
Warbler
T4–
670
01,
400
CB
asile
uter
usco
rona
tusRus
set-crow
nedW
arbler
T4
800
900
FB
asile
uter
uscu
liciv
orus
Golden
-crowne
dWarbler
T1–
4L
900
FF
19–1
0–98
Bas
ileut
erus
tris
tria
tusThree-striped
Warbler
T3,
680
01,
400
FB
asile
uter
usfu
lvic
auda
Buff-rumpe
dW
arbler
T3
600
600
R?
Bas
ileut
erus
rivu
lari
sRiver
Warbler
T1–
6L
900
F?F
20–0
7–97
Con
iros
trum
spec
iosu
mChe
stnu
t-ve
nted
Con
ebill
O1
450
450
VR(1)
Con
iros
trum
albi
fron
sCap
pedCon
ebill
O4
700
700
VR(1)
Psa
roco
lius
decu
man
usCrested
Orope
ndola
T1–
6L
1,30
0C
NA
20–1
0–98
Psa
roco
lius
atro
vire
nsDus
ky-green
Orope
ndola*
T6
400
1,40
0C
NB
04–1
1–98
A.B. Hennessey et al. 348
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SEBASTIAN K. HERZOGInstitut fur Vogelforschung ‘‘Vogelwarte Helgoland’’, An der Vogelwarte 21, 26386
Wilhelmshaven, Germany.
MICHAEL KESSLERAlbrecht-von-Haller-Institut fur Pflanzenwissenschaften, Abteilung Systematische Botanik,
Untere Karspule 2, 37073 Gottingen, Germany.
DANIEL ROBISONRurrenabaque, La Paz, Bolivia.
Received April 2002; revision accepted 9 June 2003