Observations on the Natural History of the Royal Sunangel (Heliangelus regalis) in the Nangaritza Valley, Ecuador

Author(s) :Juan F. Freile, Paolo Piedrahita, Galo Buitrn-Jurado, Carlos A. Rodrguez, Oswaldo Jadn, and Elisa Bonaccorso Source: The Wilson Journal of Ornithology, 123(1):85-92. 2011. Published By: The Wilson Ornithological Society DOI: http://dx.doi.org/10.1676/10-054.1 URL: http://www.bioone.org/doi/full/10.1676/10-054.1

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The Wilson Journal of Ornithology 123(1):8592, 2011

OBSERVATIONS ON THE NATURAL HISTORY OF THE ROYAL SUNANGEL (HELIANGELUS REGALIS) IN THE NANGARITZA VALLEY, ECUADOR
N-JURADO,2 JUAN F. FREILE,1,4 PAOLO PIEDRAHITA,2 GALO BUITRO 2 3 CARLOS A. RODRIGUEZ, OSWALDO JADAN, AND ELISA BONACCORSO2
ABSTRACT.The Royal Sunangel (Heliangelus regalis) is endemic to sandstone ridges in southeast Ecuador and northeast Peru. This hummingbird is currently considered endangered, although little has been published on its natural history, distribution, and conservation. We found H. regalis in three habitat types, but abundance was higher in stunted shrubland, at ridgetops. It was observed to feed on seven plant species, mostly following regular feeding routes, between 0 and 2.5 m above ground. We describe six different vocalizations, as well as two flight displays, and observations on social interactions. We also discuss its current conservation status in Ecuador, where we estimate that ,2,500 individuals might persist. Received 5 April 2010. Accepted 8 September 2010.

The Royal Sunangel (Heliangelus regalis) occurs in ridgetop and adjacent stunted forests in the Cordillera del Co ndor, Cordillera de Cola n, Cordillera Azul, and other ridges with poor sandy soils in extreme southeastern Ecuador and northeastern Peru (BirdLife International 2009). It was considered endemic to Peru (its southernmost locality being Pauya, Cordillera Azul, Departan; Schulenberg et al. 2001) until mento San Mart recently (Schulenberg et al. 2007). Krabbe and Ahlman (2009) presented the first documented record for Ecuador from a shrubby forest on a sandstone mountain in the Nangaritza Valley, Zamora Chinchipe Province. Heliangelus regalis is currently ranked as globally Endangered because of its limited distributional range, where selective logging and forest clearing are increasing, and where largescale mining exploitation represents a major threat. Heliangelus regalis is seemingly rather numerous locally in the Cordillera del Co ndor, Cordillera de Cola n, and Cordillera Azul (Schulenberg et al. 2001, 2007). However, its global population is likely small, confined to unique forests in a limited center of endemism (Andean Ridgetop Forests; Stattersfield et al. 1998). Little is known about the ecology, distribution, and conservation status of the Royal Sunangel (Seddon et al. 1996).
1 Fundacio n Numashir, Casilla Postal 17-12-122, Quito, Ecuador. 2 a (QCAZ), Pontificia Universidad Museo de Zoolog Cato lica del Ecuador, Avenida 12 de Octubre 1076 y Roca, Quito, Ecuador. 3 Herbario Reinaldo Espinosa, Universidad Nacional de Loja, Ciudad Universitaria La Argelia, Loja, Ecuador. 4 Corresponding author; e-mail: jfreileo@yahoo.com

We undertook observations on the natural history of H. regalis at two different localities (above Miazi and above Yankuam) during field work for a rapid assessment of two sandstone, flat-top ridgetops (locally called tepuis because of resemblance to the Guianan table-top mountains) currently protected by the local comdeas (04u 139 58.80 S, 78u munity of Las Orqu 399 00 W, 900 m asl). We present our field observations to briefly assess its habitat preferences, comparing our results with previous habitat descriptions (Fitzpatrick et al. 1979, Seddon et al. 1996), contribute data on its diet, displays, vocalizations, and social interactions, and discuss its current conservation status. OBSERVATIONS Field Identification.Field identification of males was straight forward, as H. regalis is the only hummingbird entirely violet-blue, which looks mostly black in poor light conditions. Females were identified by pale tawny underparts with some green streaking-spotting in the throat and a plain tawny buff crescent in the chest with a rather long, deep blue forked tail (Schulenberg et al. 2007). There was no overlap with other Heliangelus species and identification of female plumage birds is considered accurate (Krabbe and Ahlman 2009). Habitat.We observed H. regalis in three different vegetation types (Table 1). On 8 April 2009, we observed a single female in the understory of stunted shrubland on a ridgetop above Miazi (04 u 15 9 0 0 S, 78 u 37 9 1.2 0 W; Fig. 1). Vegetation at this site was characterized by low stature, twisted canopy (28 m in height), many epiphytes and hemiepiphytes, and dense under-

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TABLE 1. Habitat types where H. regalis was recorded in two localities in the Nangaritza Valley, Zamora Chinchipe Province, southeast Ecuador.
Above Miazi Above Yankuam Elevation (m asl) Terrain

Dense foothill forest Dense lower montane forest Stunted shrubland and paramo-like habitat

1 female/1 male 1 female/2 males 1 female 1 female/4 males

1,250/1,300 1,5501,650 1,300 at Miazi, 1,800 1,850 at Yankuam

Slopes Slopes Ridgetop

story. This was the only individual observed despite considerable mist-netting and point-count sampling at this locality. A female was mist-netted at 1,250 m elevation on the slopes of a sandstone mountain above Yankuam Lodge (04u 159 7.20 S, 78u 409 1.20 W)

on 15 April 2009, and a male was observed feeding at mid-strata in dense foothill forest characterized by 1525 m canopy height, and dominated by several species of Rubiaceae, Euphorbiaceae, Melastomataceae, Myrsinaceae, Clusiaceae, Meliaceae, and Podocarpaceae with

FIG. 1. Area occupied by the three vegetation types inhabited by Royal Sunangel (Heliangelus regalis) in the Nangaritza Valley of southeast Ecuador. Map by Oswaldo Jada n and Elisa Bonaccorso.

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scattered Dictiocaryum lamarckianum, Iriartea spp., and Wettinia spp. palms. Two separate males and a single female were observed on 17 April 2009 in dense lower montane forest above Yankuam (Table 1). This forest type was characterized by a fairly discontinuous canopy with an average height of 10 m (max 5 12 m) with lower vegetation where limestone was exposed. Dominant families were Euphorbiaceae, Clusiaceae, Cunoniaceae, Rubiaceae, Humiriaceae, and Ericaceae. The understory was dense (, 80% ground cover in some areas) including patches of ground bromeliads with heavy loads of mosses, vine tangles, and epiphytes. Of the observed individuals, one male and one female were in a natural forest gap, while another male was in dense and tangled understory feeding at edges of trails, where vegetation was sparser than in areas away from trails. At least four males and one female were observed feeding at shrubby edges and in tangled interior on the ridgetop of the sandstone mountain above Yankuam (Table 1) on 17 and 1920 April 2009; three males were observed performing aerial displays near a tall rocky outcrop at the edge of the forest. Two distinctive but intermixed habitats occurred in this area: (1) stunted shrubland, and (2) an atypical paramo-like vegetation, despite the low altitude for paramos (Sierra 1999). These habitats were characterized by trees and bushes of low stature, a canopy at 28 m in height with emergent trees barely exceeding 5 m. The understory was dense, reaching 7580% cover in some areas; the ground cover was also dense with many ground bromeliads, acaulescent rosettes (those having or appearing to have no stem), paramo-like herbs, and terrestrial mosses. The density of epiphytes and mosses was low, but typical paramo families and genera were dominant, including Macleania and Cavendishia (Ericaceae), Macrocarpaea (Gentianaceae), Meriania and Miconia (Melastomataceae) bushes, ground Asteraceae, and Wienmania (Cunionaceae). Vegetation types described correspond to previous ecosystem classifications for the Nangaritza area. Detailed information about general vegetation types in the Nangaritza Valley is provided by Foster and Beltra n (1997), Palacios (1997), Neill (2007), and Jada n (2010). Feeding Behavior.A male H. regalis was feeding in dense foothill forest by hovering at an epiphytic Guzmania (Bromeliaceae) bromeliad ,3 m above ground. At least one other male fed

by hovering at several flowers of two ground bromeliads Tillandsia cf. asplundii, and one ericad shrub Disterigma alaternoides, both with fairly long (, 2 cm) corollas, in dense lower montane forest. Feeding heights ranged from 0.30 to 2.5 m. We observed 34 feeding visits of H. regalis to seven different plant species in stunted shrubland and paramo-like habitat above 1,800 m. Plants used for foraging included a small terrestrial yellow-flowered Guzmania gracilior (Bromeliaceae) with six feeding visits; a larger, green-andpink-flowered epiphytic G. garciaensis (4 feeding visits); the epiphytic, fuchsia-flowered Elleanthus ampliflorus (Orchidaceae) (1 visit); an unidentified small epiphytic bromeliad (1 visit); the stunted tree Macrocarpaea harlingii (Gentianaceae) was visited nine times; an epiphytic Cavendishia spp. and a shrub Macleania spp. (both pink-flowered Ericaceae) were visited six and three times, respectively. Feeding heights ranged from 0 to 2.5 m. H. regalis fed by hovering on most feeding visits (94%, n 5 34), but perched on a nearby twig at three of 10 flowers probed during a single visit to a ground-living small bromeliad. It perched on the ground at one of five flowers probed during another visit to a ground bromeliad. A single male was observed to return to the same flowering plants at 815 min intervals, following a somewhat similar route. It first visited a 3-m tall Macrocarpaea harlingii where it fed on several individual flowers (but not on the same flowers during consecutive visits), and then moved either to another Macrocarpaea or to an epiphytic Guzmania garciaensis, both ,5 m from the first Macrocarpaea. Subsequently, it visited a cluster of 10 ground G. gracilior, and left the area in the same direction from which it arrived. Another individual male was observed making regular visits to a patch of small G. gracilior and a patch of Macleania spp. shrub, returning to a perch of 2 m height. Social Interactions.Few interactions with other hummingbirds were observed. One male H. regalis was observed displacing, but not directly attacking or chasing, a male Ecuadorian Piedtail (Phlogophilus hemileucurus) at feeding sites (in ridgetop shrubland above Yankuam), and a male attacked and chased a male Green-fronted Lancebill (Doryfera ludovicae) when it hovered in front of ericaceous flowers in the same site; minutes later a D. ludovicae returned and hovered

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in front of the same flowers without being displaced. No other aggressive behaviors were reported although other hummingbird species occurred in feeding areas used by H. regalis. No interactions were observed in lower elevation forests above Yankuam (below 1,700 m). Intense disputes were observed among Brown Violetears (Colibri delphinae), Rufous-vented Whitetips (Urosticte ruficrissa), Violet-fronted Brilliants (Heliodoxa leadbeateri), and Blue-fronted Lancebill (D. johannae) at flowering forest trees and epiphytes, but no H. regalis was observed in those interactions. The only female found above Miazi was observed displacing an unidentified hummingbird and perching upright, with her neck stretched, possibly in territorial dispute. Vocalizations.We heard and tape recorded several types of vocalization. A tame and curious male approached one observer (J. F. Freile) and remained perched motionless for 1 min. He uttered a sharp, high-pitched, fast chichu p chu p! before take off with the last note more emphatic. A female in the same area perched in a natural p! We forest gap uttered a thin, high-pitched tzi failed to record both vocalizations. A feeding male uttered an emphatic tchu p or p! every 3.56 sec (deposited at www.xenocanto. ch org; XC 45910; Fig. 2A1), whereas other feeding calls were uttered at shorter intervals (22.5 sec), or in short two- to three-notes descending series tchu ptchup, tchUp-tchu p-tchup with the first note higher (XC 45911; Fig. 2A2). The duration of each note was ,0.1 sec for the feeding call (Fig. 2A2), while frequency ranged from 1.51.8 to 1719 kHz (call 2A1), and 1.5 to 21 kHz (call 2A2). Two displaying males produced a fast chattered series of very high tEEp or je e t notes lasting 2.5 4.5 sec (XC 45912; Fig. 2B). This vocalization contained 19 notes of 0.10sec mean duration (0.070.15 sec) with frequencies ranging from 1.7 to 16.69 kHz. Two males were observed in stunted shrubland and heard in intense dispute, constantly vocalizing an endless, thin, highpitched jumble jijijit9jijit9jijit9jijiji, notes uttered at a much faster rate than regular vocalizations. This dispute and chatter was only interrupted when a third male dashed towards them. Our recordings are uploaded to www.xentocanto.org (XC 4591045915) as only a single recording of H. regalis is currently deposited in a public audio library (Macaulay Library 18046; N. Krabbe, pers. comm.).

Display Flight.Three males were observed performing display flights in ridgetop stunted shrubland, one above a nectar source (Macrocarpaea harlingii) and two above a shrub edge and rocky outcrop. In display, one male ascended ,5 m in vertical flight, described one oval, possibly two, at the highest point and descended describing a semicircle to the same perch; the second male followed the firsts display flight constantly vocalizing (Fig. 3A). A somewhat similar display was also observed with a male ascending 1012 m and then descending in a semicircle, and diving out of sight (Fig. 3B). DISCUSSION Heliangelus regalis is generally regarded as locally fairly common (Seddon et al. 1996; Schulenberg et al. 2001, 2007; Dauphine et al. 2008). Schulenberg et al. (2001) suggested the Cordillera Azul might represent the center of abundance of H. regalis, and that it might be less threatened than currently believed. However, with a global range of only 2,100 km2, its global population has been roughly estimated at 2,500 9,999 individuals (BirdLife International 2009). Our observations suggest the abundance of H. regalis can vary across different vegetation types. However, in accordance to previous reports (Fitzpatrick et al. 1979, Davis 1986, Seddon et al. 1996, Dauphine et al. 2008), stunted shrubland appears as its preferred habitat, at least judging from relative abundances at the three vegetation types surveyed in this study. Higher numbers in stunted shrubland and paramo-like vegetation suggest these habitats provided plentiful food resources during the study season. Low detection in dense lower montane and foothill forests might also account for lower numbers in these forests (Poulsen et al. 1997). We cannot rule out seasonal movements along vegetation gradients as previously suggested by Seddon et al. (1996). Habitat suitability appears to be higher for stunted shrubland, but defining optimal habitats solely based on relative abundance has proven to be misleading (van Horne 1983, Morris 1987). H. regalis in stunted shrubland likely fed on most available nectar sources. Flowering was limited to a few individual plants of a few species. Our data suggest that H. regalis was not strongly dependent on one food plant species as found in the type locality, where the species was reported as highly dependent on Brachyotum quinquenerve (Melastomataceae) (Fitzpatrick et al. 1979); it

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FIG. 2. Spectrograms of three vocalization of male Royal Sunangel (Heliangelus regalis) in ridgetop habitats in the Nangaritza Valley of southeast Ecuador. A1 and A2 5 feeding calls; B 5 aerial displaying calls. Tape-recordings by Paolo Piedrahita and Juan F. Freile filed at www.xenocanto.org, XC 4591045912; figures by Paolo Piedrahita.

should be noted that B. quinquinerve was absent from our study area. Dauphine et al. (2008) also reported few food plants, but failed to identify them to the species level and provided no information on flowering at their study site. Males

and females, as previous authors (Seddon et al. 1996, Dauphine et al. 2008) have suggested, might feed at different elevations or different food plants; a suggestion we failed to prove as we observed few females.

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FIG. 3. Flight displays performed by male Royal Sunangel (Heliangelus regalis) at ridgetop shrubland in the Nangaritza Valley of southeast Ecuador. 3A has two males in successive flights (illustration by Juan F. Freile); 3B has a single displaying male (illustration by Galo Buitro n-Jurado).

Fitzpatrick et al. (1979) and Seddon et al. (1996) reported perching to be a more common foraging method than hovering, in contrast to our observations. We suggest these discrepancies indicate local or seasonal differences in feeding strategies, considering that Seddon et al. (1996) performed more prolonged observations (45 observations of males, 44 of females). Observations were limited to 6 hrs in stunted shrubland and paramo, but we suspect H. regalis followed regular feeding routes in fairly regular time periods within a fixed territory. Two events of territorial defense were observed which, in accordance to previous observations (Fitzpatrick et al. 1979, Seddon et al. 1996), suggest territoriality (Feinsinger and Colwell 1978). Other Heliangelus species are also reported to be territorial (Ortiz-Crespo 2003). Vocalizations are generally similar to those previously described (Schulenberg et al. 2007), but displaying notes were uttered in fast chattered series, contrary to the high teep note described by Schulenberg et al. (2007). The dispute calls we report are similar to a series of tick notes

described by Fitzpatrick et al. (1979) in malemale chases. To our ears, they appear harsher, more metallic, and more chattered than those described by Fitzpatrick et al. (1979). Displays differed from those observed at the type locality (Fitzpatrick et al. 1979) as we did not observe birds repeating the circular flight towards the opposite side of perches in a figure-8 pattern. More detailed observations are needed to elucidate if display flights differ locally. H. regalis is apparently fairly numerous in stunted ridgetop shrubland and paramo-like habdeas sandstone ridges. At least itats at Las Orqu five different birds were found (one female, four males) in a small sampled area at the ridgetop (, 0.02 km2; i.e., 500-m linear transect with a 40m width band), suggesting a healthy population. These observations provide a rough estimate of 250 individuals/km2 in the stunted ridgetop shrubland habitat that H. regalis seemingly preferred during our study. An estimate of the area covered by this forest type (Fig. 1) resulted in 10.11 km2 of the habitat where H. regalis was most abundant in the Nangaritza Valley. This

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suggests the total population in the Ecuadorian portion of H. regalis range might total ,2,500 individuals. These numbers are crude as better population size estimates and trends are needed. H. regalis is seemingly less numerous in lower elevation montane and foothill forests above Yankuam Lodge. Only three males and one female were observed during 7 days of pointcount surveys above Yankuam, in dense foothill and dense lower montane forests, despite a total of 25 10-min point counts. H. regalis was not found in similar habitats above Miazi during 21 10-min point counts, but one female was observed by random sampling on a small ridgetop. We sampled ,23 km between both study sites. This might indicate 11.5 birds/km in these two forest types, but dense habitat might reduce detection. CONSERVATION IMPLICATIONS Habitat loss is still incipient along the Ecuadorian range of the species, but mining concessions represent a serious forthcoming threat to the endemic biodiversity of the Cordillera del Co ndor. There is major interest by the Ecuadorian government to consolidate mining extraction in the Cordillera del Co ndor region because of apparently large deposits of gold and copper, as well as silver, silica, and other minerals and metals (Lo pez et al. 2003, Fontbote et al. 2004, Neill 2007, Drobe et al. 2008; see also www. aurelianecuador.com; www.corriente.com; www. kinross.com). Currently, several areas north of Nangaritza Valley are being prospected by mining companies, and access roads are being rapidly opened and improved. Populations of species confined to these poor-soil growing forests are imminently threatened as large sandstone ridges will potentially be opened to large-scale mining. Under this scenario, we consider accurate the status of globally Endangered (BirdLife International 2009) for H. regalis despite current population figures suggesting it is less threatened. Conservation initiatives are underway in the Nangaritza Valley, including private birdwatching tour operators, land protection, and an ongoing management plan developed by Fundacio n Arco Iris of Loja along with local communities. These initiatives benefit from biodiversity surveys that support the biological and hydrological importance of the Nangaritza Valley. We encourage others to undertake more specific studies in the region, particularly to assess populations and habitat of globally threatened species including

H. regalis . Easy access to the Nangaritza sandstone ridges facilitate biological surveys and bird studies of an avifauna generally regarded as difficult to reach. ACKNOWLEDGMENTS
Field work was part of a Rapid Assessment Program by Fundacio n Arco Iris of Loja, and supported by Conservation International. We especially thank Robert Jime nez for diligent assistance in field work. Oswaldo Jada n thanks var Merino of Herbario LOJA, Jose Zhofre Aguirre and Bol M. Manzanares, Charlotte Taylor, and Ron Liesner of Missouri Botanical Garden, and Alfonso Garmendia of Universidad Polite cnica de Valencia for assistance in plant identification. Niels Krabbe made thorough comments on the manuscript, T. S. Schulenberg provided information from Peru , and Alejandro Solano and Daniel Cadena kindly helped with bibliographic requests. We thank Leonardo Ordo n ez and Fundacio n Arco Iris staff for logistics and issuing plant research permits. This research was conducted under Ministerio del Ambiente research permits Numbers 006-IC-FLO-DBAP-VS-DRLZCH-MA and FAU-001-DNB/VS. Fieldwork by Galo Buitro n, Paolo Piedrahita, and Elisa Bonaccorso was supported by a Nacional de Ciencia y Tecnolog a (SENACYT) Secretar Project PIC-08-470. Comments by three anonymous reviewers greatly improved a previous version of this paper. Juan F. Freile dedicates this paper to the dearly loved memory of Tami Bueno.

LITERATURE CITED
BIRDLIFE INTERNATIONAL. 2009. The BirdLife checklist of the birds of the world, with conservation status and taxonomic sources, Version December 2009. BirdLife International, Cambridge, United Kingdom. http:// www.birdlife.org/datazone/species. , N., R. J. COOPER, AND A. TSAMAJAIN YAGKUAG. DAUPHINE 2008. A new location and altitudinal range extension for Royal Sunangel Heliangelus regalis. Cotinga 30: 8384. DAVIS, T. J. 1986. Distribution and natural history of some n and birds from the departments of San Mart Amazonas, northern Peru. Condor 88: 5056. DROBE, J., J. HOFFERT, R. FONG, J. P. HAILE, AND J. COLLINS. 2008. Mirador copper-gold project: 30,000 TPD feasibility study. Corriente Resources Inc., Vancouver, British Columbia, Canada. FEINSINGER, P. AND R. K. COLWELL. 1978. Community organization among neotropical nectar-feeding birds. American Zoologist 18: 779795. FITZPATRICK, J. W., D. E. WILLARD, AND J. TERBORGH. 1979. A new species of hummingbird from Peru . Wilson Bulletin 91: 177186. FONTBOTE, L., J. VALLANCE, A. MARKOWSKI, AND M. CHIARADIA. 2004. Oxidized gold skarns in the Nambija District, Ecuador. Society of Economic Geologists Special Publication 11: 341357. N. 1997. Vegetacio n y flora de la FOSTER, R. AND H. BELTRA Cordillera del Co ndor. Pages 4565 in The Cordillera

92

THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 123, No. 1, March 2011 Peru: a biological assessment (T. S. Schulenberg, and K. Awbrey, Editors). Conservation International RAP Working Papers 7. Washington, D.C., USA. POULSEN, B. O., N. KRABBE, A. FRLANDER, M. B. HINOJOSA, AND C. O. QUIROGA. 1997. A rapid assessment of Bolivian and Ecuadorian montane avifaunas using 20-species lists: efficiency, biases and data gathered. Bird Conservation International 7: 5367. SCHULENBERG, T. S., J. P. ONEILL, D. F. LANE, T. VALQUI, JAR. 2001. Aves. Pages 7584 in Peru AND C. ALBU , BiaboCordillera Azul (W. S. Alverson, L. O. guez, and D. K. Moskovits, Editors). The Field Rodr Museum Rapid Biological Inventories 2. Chicago, Illinois, USA. SCHULENBERG, T. S., D. F. STOTZ, D. F. LANE, J. P. . ONEILL, AND T. A. PARKER III. 2007. Birds of Peru Christopher Helm, London, United Kingdom. SEDDON, N., R. BARNES, S. H. M. BUTCHART, C. W. N. NDEZ. 1996. Recent observaDAVIES, AND M. FERNA tions and notes on the ecology of the Royal Sunangel Heliangelus regalis. Bulletin of the British Ornithologists Club 116: 4649. n remanente del Ecuador conSIERRA, R. 1999. Vegetacio tinental, circa 1996. 1:1,000.000. Proyecto INEFAN/ GEF-BIRF and Wildlife Conservation Society, Quito, Ecuador. STATTERSFIELD, A. J., M. J. CROSBY, A. J. LONG, AND D. C. WEGE. 1998. Endemic birds areas of the world. Priorities for biodiversity conservation. BirdLife International, Cambridge, United Kingdom. VAN HORNE, B. 1983. Density as a misleading indicator of habitat quality. Journal of Wildlife Management 47: 893901.

del Co ndor Region of Ecuador and Peru: a biological assessment (T. S. Schulenberg and K. Awbrey, Editors). Conservation International RAP Working Papers 7. Washington, D.C., USA. N, O. 2010. Plantas de los tepuyes de San Miguel de JADA deas, valle de Nangaritza, cordillera del Las Orqu Co ndor, sureste de Ecuador. In Evaluacio n ra pida de la o biodiversidad de los tepuyes de la cuenca alta del r Nangaritza, cordillera del Co ndor, Ecuador (J. M. Guayasamin, and E. Bonaccorso, Editors). Fundacio n Arco Iris and Conservation International, Quito, Ecuador. KRABBE, N. AND F. L. AHLMAN. 2009. Royal Sunangel Heliangelus regalis at Yankuam Lodge, Ecuador. Cotinga 31: 69. PEZ, F., M. TORRES, AND R. BELTRA N. 2003. La miner a LO en el Parque Nacional Podocarpus: consideraciones legales y ambientales. Fundacio n Ecolo gica Arco Iris, Loja, Ecuador. MORRIS, D. W. 1987. Tests of density-dependent habitat selection in a patchy environment. Ecological Monographs 57: 269281. nico de la regio n de la NEILL, D. 2007. Inventario bota cordillera del Co ndor, Ecuador y Peru : actividades y ficos del proyecto 20042007. Misresultados cient souri Botanical Garden, St. Louis, USA. http://www. mobot.org/MOBOT/Research/ecuador/cordillera/pdf/ EntireSpanishReport.pdf. es: historia natural de ORTIZ-CRESPO, F. 2003. Los colibr unas aves casi sobrenaturales. Imprenta Mariscal, Quito, Ecuador. o Nangaritza (cordillera PALACIOS, W. 1997. Cuenca del r del Co ndor), una zona para conservar. Pages 3745 in The Cordillera del Co ndor Region of Ecuador and