Articulo Nangaritza

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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/10.1676/10-054.1URL: http://www.bioone.org/doi/full/10.1676/10-054.1

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OBSERVATIONS ON THE NATURAL HISTORY OF THE ROYALSUNANGEL (HELIANGELUS REGALIS) IN THE NANGARITZA

VALLEY, ECUADOR

JUAN F. FREILE,1,4 PAOLO PIEDRAHITA,2 GALO BUITRON-JURADO,2

CARLOS A. RODRIGUEZ,2 OSWALDO JADAN,3 AND ELISA BONACCORSO2

ABSTRACT.The Royal Sunangel (Heliangelus regalis) is endemic to sandstone ridges in southeast Ecuador andnortheast Peru. This hummingbird is currently considered endangered, although little has been published on its naturalhistory, distribution, and conservation. We found H. regalis in three habitat types, but abundance was higher in stuntedshrubland, at ridgetops. It was observed to feed on seven plant species, mostly following regular feeding routes, between 0and 2.5 m above ground. We describe six different vocalizations, as well as two flight displays, and observations on socialinteractions. We also discuss its current conservation status in Ecuador, where we estimate that ,2,500 individuals mightpersist. Received 5 April 2010. Accepted 8 September 2010.

The Royal Sunangel (Heliangelus regalis)occurs in ridgetop and adjacent stunted forests inthe Cordillera del Condor, Cordillera de Colan,Cordillera Azul, and other ridges with poor sandysoils in extreme southeastern Ecuador and north-eastern Peru (BirdLife International 2009). It wasconsidered endemic to Peru (its southernmostlocality being Pauya, Cordillera Azul, Departa-mento San Martn; Schulenberg et al. 2001) untilrecently (Schulenberg et al. 2007). Krabbe and

Ahlman (2009) presented the first documentedrecord for Ecuador from a shrubby forest on asandstone mountain in the Nangaritza Valley,Zamora Chinchipe Province.

Heliangelus regalis is currently ranked asglobally Endangered because of its limiteddistributional range, where selective logging andforest clearing are increasing, and where large-scale mining exploitation represents a majorthreat. Heliangelus regalis is seemingly rathernumerous locally in the Cordillera del Condor,

Cordillera de Colan, and Cordillera Azul (Schu-lenberg et al. 2001, 2007). However, its globalpopulation is likely small, confined to uniqueforests in a limited center of endemism (AndeanRidgetop Forests; Stattersfield et al. 1998). Littleis known about the ecology, distribution, andconservation status of the Royal Sunangel (Sed-don et al. 1996).

We undertook observations on the naturalhistory of H. regalis at two different localities(above Miazi and above Yankuam) during fieldwork for a rapid assessment of two sandstone,flat-top ridgetops (locally called tepuis be-cause of resemblance to the Guianan table-topmountains) currently protected by the local com-munity of Las Orqudeas (04u 139 58.80 S, 78u399 00 W, 900 m asl). We present our field obser-vations to briefly assess its habitat preferences,

comparing our results with previous habitatdescriptions (Fitzpatrick et al. 1979, Seddon etal. 1996), contribute data on its diet, displays,vocalizations, and social interactions, and discussits current conservation status.

OBSERVATIONS

Field Identification.Field identification ofmales was straight forward, as H. regalis is theonly hummingbird entirely violet-blue, whichlooks mostly black in poor light conditions.

Females were identified by pale tawny underpartswith some green streaking-spotting in the throatand a plain tawny buff crescent in the chest with arather long, deep blue forked tail (Schulenberg etal. 2007). There was no overlap with otherHeliangelus species and identification of femaleplumage birds is considered accurate (Krabbe andAhlman 2009).

Habitat.We observed H. regalis in threedifferent vegetation types (Table 1). On 8 April2009, we observed a single female in the under-story of stunted shrubland on a ridgetop aboveMiazi (04u 159 00 S, 78u 379 1.20 W; Fig. 1).Vegetation at this site was characterized by lowstature, twisted canopy (28 m in height), manyepiphytes and hemiepiphytes, and dense under-

1 Fundacion Numashir, Casilla Postal 17-12-122, Quito,Ecuador.

2 Museo de Zoologa (QCAZ), Pontificia Universidad

Catolica del Ecuador, Avenida 12 de Octubre 1076 y Roca,Quito, Ecuador.

3 Herbario Reinaldo Espinosa, Universidad Nacional deLoja, Ciudad Universitaria La Argelia, Loja, Ecuador.

4 Corresponding author; e-mail: [email protected]

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

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story. This was the only individual observeddespite considerable mist-netting and point-countsampling at this locality.

A female was mist-netted at 1,250 m elevation

on the slopes of a sandstone mountain aboveYankuam Lodge (04u 159 7.20 S, 78u 409 1.20 W)

on 15 April 2009, and a male was observedfeeding at mid-strata in dense foothill forestcharacterized by 1525 m canopy height, anddominated by several species of Rubiaceae,

Euphorbiaceae, Melastomataceae, Myrsinaceae,Clusiaceae, Meliaceae, and Podocarpaceae with

TABLE 1. Habitat types where H. regalis was recorded in two localities in the Nangaritza Valley, Zamora ChinchipeProvince, southeast Ecuador.

Above Miazi Above Yankuam Elevation (m asl) Terrain

Dense foothill forest 1 female/1 male 1,250/1,300 SlopesDense lower montane

forest1 female/2 males 1,5501,650 Slopes

Stunted shrubland andparamo-like habitat

1 female 1 female/4 males 1,300 at Miazi, 1,8001,850 at Yankuam

Ridgetop

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

86 THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 123, No. 1, March 2011


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in front of the same flowers without beingdisplaced.

No other aggressive behaviors were reportedalthough other hummingbird species occurred in

feeding areas used by H. regalis. No interactionswere observed in lower elevation forests aboveYankuam (below 1,700 m). Intense disputes wereobserved among Brown Violetears (Colibri del-phinae), Rufous-vented Whitetips (Urosticte rufi-crissa), Violet-fronted Brilliants (Heliodoxa lead-beateri), and Blue-fronted Lancebill (D. johannae)at flowering forest trees and epiphytes, but no H.regalis was observed in those interactions. Theonly female found above Miazi was observeddisplacing an unidentified hummingbird and

perching upright, with her neck stretched, possi-bly in territorial dispute.

Vocalizations.We heard and tape recordedseveral types of vocalization. A tame and curiousmale approached one observer (J. F. Freile) andremained perched motionless for 1 min. Heuttered a sharp, high-pitched, fast chichup chup!before take off with the last note more emphatic.A female in the same area perched in a naturalforest gap uttered a thin, high-pitched tzip! Wefailed to record both vocalizations.

A feeding male uttered an emphatic tchup orchp! every 3.56 sec (deposited at www.xenocanto.org; XC 45910; Fig. 2A1), whereas other feedingcalls were uttered at shorter intervals (22.5 sec), orin short two- to three-notes descending series tchup-tchup, tchUp-tchup-tchup with the first note higher(XC 45911; Fig. 2A2). The duration of each notewas,0.1 sec for the feeding call (Fig. 2A2), whilefrequency ranged from 1.51.8 to 1719 kHz (call2A1), and 1.5 to 21 kHz (call 2A2).

Two displaying males produced a fast chattered

series of very high tEEp or jeet notes lasting 2.54.5 sec (XC 45912; Fig. 2B). This vocalizationcontained 19 notes of 0.10sec mean duration(0.070.15 sec) with frequencies ranging from 1.7to 16.69 kHz. Two males were observed instunted shrubland and heard in intense dispute,constantly vocalizing an endless, thin, high-pitched jumble jijijit9jijit9jijit9jijiji, notes utteredat a much faster rate than regular vocalizations.This dispute and chatter was only interruptedwhen a third male dashed towards them. Ourrecordings are uploaded to www.xentocanto.org(XC 4591045915) as only a single recording ofH. regalis is currently deposited in a public audiolibrary (Macaulay Library 18046; N. Krabbe,pers. comm.).

Display Flight.Three males were observedperforming display flights in ridgetop stuntedshrubland, one above a nectar source (Macro-carpaea 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 descendeddescribing a semicircle to the same perch; thesecond male followed the firsts display flightconstantly vocalizing (Fig. 3A). A somewhatsimilar display was also observed with a maleascending 1012 m and then descending in asemicircle, and diving out of sight (Fig. 3B).

DISCUSSION

Heliangelus regalis is generally regarded aslocally fairly common (Seddon et al. 1996;Schulenberg et al. 2001, 2007; Dauphine et al.2008). Schulenberg et al. (2001) suggested theCordillera Azul might represent the center ofabundance of H. regalis, and that it might be lessthreatened than currently believed. However, witha global range of only 2,100 km2, its globalpopulation has been roughly estimated at 2,5009,999 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 etal. 1996, Dauphine et al. 2008), stunted shrublandappears as its preferred habitat, at least judgingfrom relative abundances at the three vegetationtypes surveyed in this study. Higher numbers instunted shrubland and paramo-like vegetationsuggest these habitats provided plentiful foodresources during the study season. Low detectionin dense lower montane and foothill forests might

also account for lower numbers in these forests(Poulsen et al. 1997). We cannot rule out seasonalmovements along vegetation gradients as previ-ously suggested by Seddon et al. (1996). Habitatsuitability appears to be higher for stunted shrub-land, but defining optimal habitats solely based onrelative abundance has proven to be misleading(van Horne 1983, Morris 1987).

H. regalis in stunted shrubland likely fed onmost available nectar sources. Flowering waslimited to a few individual plants of a few species.Our data suggest that H. regalis was not stronglydependent on one food plant species as found inthe type locality, where the species was reportedas highly dependent on Brachyotum quinquenerve(Melastomataceae) (Fitzpatrick et al. 1979); it



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should be noted that B. quinquinerve was absent

from our study area. Dauphine et al. (2008) alsoreported few food plants, but failed to identifythem to the species level and provided noinformation 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 foodplants; a suggestion we failed to prove as weobserved few females.

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

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Fitzpatrick et al. (1979) and Seddon et al.(1996) reported perching to be a more commonforaging method than hovering, in contrast to ourobservations. We suggest these discrepanciesindicate local or seasonal differences in feedingstrategies, considering that Seddon et al. (1996)performed more prolonged observations (45observations of males, 44 of females).

Observations were limited to 6 hrs in stuntedshrubland and paramo, but we suspect H. regalisfollowed regular feeding routes in fairly regulartime periods within a fixed territory. Two eventsof territorial defense were observed which, inaccordance to previous observations (Fitzpatricket al. 1979, Seddon et al. 1996), suggestterritoriality (Feinsinger and Colwell 1978). OtherHeliangelus species are also reported to beterritorial (Ortiz-Crespo 2003).

Vocalizations are generally similar to thosepreviously described (Schulenberg et al. 2007),but displaying notes were uttered in fast chatteredseries, contrary to the high teep note described bySchulenberg et al. (2007). The dispute calls wereport are similar to a series of tick notes

described by Fitzpatrick et al. (1979) in male-male chases. To our ears, they appear harsher,more metallic, and more chattered than thosedescribed by Fitzpatrick et al. (1979).

Displays differed from those observed at thetype locality (Fitzpatrick et al. 1979) as we did notobserve birds repeating the circular flight towardsthe opposite side of perches in a figure-8 pattern.More detailed observations are needed to eluci-date if display flights differ locally.

H. regalis is apparently fairly numerous instunted ridgetop shrubland and paramo-like hab-itats at Las Orqudeas sandstone ridges. At leastfive different birds were found (one female, fourmales) in a small sampled area at the ridgetop(, 0.02 km2; i.e., 500-m linear transect with a 40-m width band), suggesting a healthy population.These observations provide a rough estimate of250 individuals/km2 in the stunted ridgetopshrubland habitat that H. regalis seeminglypreferred during our study. An estimate of thearea covered by this forest type (Fig. 1) resultedin 10.11 km2 of the habitat where H. regalis wasmost abundant in the Nangaritza Valley. This

FIG. 3. Flight displays performed by male Royal Sunangel (Heliangelus regalis) at ridgetop shrubland in theNangaritza 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 Buitron-Jurado).



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suggests the total population in the Ecuadorianportion of H. regalis range might total ,2,500individuals. These numbers are crude as betterpopulation size estimates and trends are needed.

H. regalis is seemingly less numerous in lowerelevation montane and foothill forests aboveYankuam Lodge. Only three males and onefemale were observed during 7 days of point-count surveys above Yankuam, in dense foothilland dense lower montane forests, despite a total of25 10-min point counts. H. regalis was not foundin similar habitats above Miazi during 21 10-minpoint counts, but one female was observed byrandom sampling on a small ridgetop. Wesampled,23 km between both study sites. This

might indicate 11.5 birds/km in these two foresttypes, but dense habitat might reduce detection.

CONSERVATION IMPLICATIONS

Habitat loss is still incipient along the Ecua-dorian range of the species, but mining conces-sions represent a serious forthcoming threat to theendemic biodiversity of the Cordillera del Condor.There is major interest by the Ecuadoriangovernment to consolidate mining extraction inthe Cordillera del Condor region because of

apparently large deposits of gold and copper, aswell as silver, silica, and other minerals andmetals (Lopez 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 ofNangaritza Valley are being prospected by miningcompanies, and access roads are being rapidlyopened and improved. Populations of speciesconfined to these poor-soil growing forests areimminently threatened as large sandstone ridges

will potentially be opened to large-scale mining.Under this scenario, we consider accurate thestatus of globally Endangered (BirdLife Interna-tional 2009) for H. regalis despite current popu-lation figures suggesting it is less threatened.

Conservation initiatives are underway in theNangaritza Valley, including private birdwatchingtour operators, land protection, and an ongoingmanagement plan developed by Fundacion ArcoIris of Loja along with local communities. Theseinitiatives benefit from biodiversity surveys thatsupport the biological and hydrological impor-tance of the Nangaritza Valley. We encourageothers to undertake more specific studies in theregion, particularly to assess populations andhabitat of globally threatened species including

H. reg alis. Easy access to the Nangaritzasandstone ridges facilitate biological surveys andbird studies of an avifauna generally regarded asdifficult to reach.

ACKNOWLEDGMENTS

Field work was part of a Rapid Assessment Program byFundacion Arco Iris of Loja, and supported by Conserva-tion International. We especially thank Robert Jimenez fordiligent assistance in field work. Oswaldo Jadan thanksZhofre Aguirre and Bolvar Merino of Herbario LOJA, JoseM. Manzanares, Charlotte Taylor, and Ron Liesner ofMissouri Botanical Garden, and Alfonso Garmendia ofUniversidad Politecnica de Valencia for assistance in plantidentification. Niels Krabbe made thorough comments onthe manuscript, T. S. Schulenberg provided information

from Peru, and Alejandro Solano and Daniel Cadena kindlyhelped with bibliographic requests. We thank LeonardoOrdonez and Fundacion Arco Iris staff for logistics andissuing plant research permits. This research was conductedunder Ministerio del Ambiente research permitsNumbers 006-IC-FLO-DBAP-VS-DRLZCH-MA andFAU-001-DNB/VS. Fieldwork by Galo Buitron, PaoloPiedrahita, and Elisa Bonaccorso was supported bySecretara Nacional de Ciencia y Tecnologa (SENACYT)Project PIC-08-470. Comments by three anonymousreviewers greatly improved a previous version of thispaper. Juan F. Freile dedicates this paper to the dearly lovedmemory of Tami Bueno.

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