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Observations on the Natural History of the Royal Sunangel(Heliangelus regalis) in the Nangaritza Valley, EcuadorAuthor(s) :Juan F. Freile, Paolo Piedrahita, Galo Buitrón-Jurado, Carlos A.Rodríguez, Oswaldo Jadán, and Elisa BonaccorsoSource: The Wilson Journal of Ornithology, 123(1):85-92. 2011.Published By: The Wilson Ornithological SocietyDOI: 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 ROYAL

SUNANGEL (HELIANGELUS REGALIS) IN THE NANGARITZAVALLEY, 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 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 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 Martın; Schulenberg et al. 2001) untilrecently (Schulenberg et al. 2007). Krabbe andAhlman (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 Orquıdeas (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 (2–8 m in height), manyepiphytes and hemiepiphytes, and dense under-

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

Ecuador.2 Museo de Zoologıa (QCAZ), Pontificia Universidad

Catolica 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

The Wilson Journal of Ornithology 123(1):85–92, 2011

85

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 15–25 m canopy height, and

dominated 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 Chinchipe

Province, southeast Ecuador.

Above Miazi Above Yankuam Elevation (m asl) Terrain

Dense foothill forest 1 female/1 male 1,250/1,300 Slopes

Dense lower montane

forest

1 female/2 males 1,550–1,650 Slopes

Stunted shrubland and

paramo-like habitat

1 female 1 female/4 males 1,300 at Miazi, 1,800–

1,850 at Yankuam

Ridgetop

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 Jadan and Elisa Bonaccorso.

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

scattered Dictiocaryum lamarckianum, Iriarteaspp., and Wettinia spp. palms.

Two separate males and a single female wereobserved on 17 April 2009 in dense lowermontane forest above Yankuam (Table 1). Thisforest type was characterized by a fairly discon-tinuous canopy with an average height of 10 m(max 5 12 m) with lower vegetation wherelimestone was exposed. Dominant families wereEuphorbiaceae, Clusiaceae, Cunoniaceae, Rubia-ceae, Humiriaceae, and Ericaceae. The understorywas dense (, 80% ground cover in some areas)including patches of ground bromeliads withheavy loads of mosses, vine tangles, and epi-phytes. Of the observed individuals, one male andone female were in a natural forest gap, whileanother male was in dense and tangled understoryfeeding at edges of trails, where vegetation wassparser than in areas away from trails.

At least four males and one female wereobserved feeding at shrubby edges and in tangledinterior on the ridgetop of the sandstone mountainabove Yankuam (Table 1) on 17 and 19–20 April2009; three males were observed performingaerial displays near a tall rocky outcrop at theedge of the forest. Two distinctive but intermixedhabitats occurred in this area: (1) stunted shrub-land, and (2) an atypical paramo-like vegetation,despite the low altitude for paramos (Sierra 1999).These habitats were characterized by trees andbushes of low stature, a canopy at 2–8 m in heightwith ‘emergent’ trees barely exceeding 5 m. Theunderstory was dense, reaching 75–80% cover insome areas; the ground cover was also dense withmany ground bromeliads, acaulescent rosettes(those having or appearing to have no stem),paramo-like herbs, and terrestrial mosses. Thedensity of epiphytes and mosses was low, buttypical paramo families and genera were domi-nant, including Macleania and Cavendishia (Eri-caceae), Macrocarpaea (Gentianaceae), Merianiaand Miconia (Melastomataceae) bushes, groundAsteraceae, and Wienmania (Cunionaceae).

Vegetation types described correspond toprevious ecosystem classifications for the Nan-garitza area. Detailed information about generalvegetation types in the Nangaritza Valley isprovided by Foster and Beltran (1997), Palacios(1997), Neill (2007), and Jadan (2010).

Feeding Behavior.—A male H. regalis wasfeeding in dense foothill forest by hovering at anepiphytic Guzmania (Bromeliaceae) bromeliad,3 m above ground. At least one other male fed

by hovering at several flowers of two groundbromeliads Tillandsia cf. asplundii, and oneericad shrub Disterigma alaternoides, both withfairly long (, 2 cm) corollas, in dense lowermontane forest. Feeding heights ranged from 0.30to 2.5 m.

We observed 34 feeding visits of H. regalis toseven different plant species in stunted shrublandand paramo-like habitat above 1,800 m. Plantsused for foraging included a small terrestrialyellow-flowered Guzmania gracilior (Bromelia-ceae) with six feeding visits; a larger, green-and-pink-flowered epiphytic G. garciaensis (4 feedingvisits); the epiphytic, fuchsia-flowered Elleanthusampliflorus (Orchidaceae) (1 visit); an unidenti-fied small epiphytic bromeliad (1 visit); thestunted tree Macrocarpaea harlingii (Gentiana-ceae) was visited nine times; an epiphyticCavendishia spp. and a shrub Macleania spp.(both pink-flowered Ericaceae) were visited sixand three times, respectively. Feeding heightsranged from 0 to 2.5 m.

H. regalis fed by hovering on most feedingvisits (94%, n 5 34), but perched on a nearbytwig at three of 10 flowers probed during a singlevisit to a ground-living small bromeliad. Itperched on the ground at one of five flowersprobed during another visit to a ground bromeliad.

A single male was observed to return to thesame flowering plants at 8–15 min intervals,following a somewhat similar route. It first visiteda 3-m tall Macrocarpaea harlingii where it fed onseveral individual flowers (but not on the sameflowers during consecutive visits), and thenmoved either to another Macrocarpaea or to anepiphytic Guzmania garciaensis, both ,5 m fromthe first Macrocarpaea. Subsequently, it visited acluster of 10 ground G. gracilior, and left the areain the same direction from which it arrived.Another individual male was observed makingregular visits to a patch of small G. gracilior and apatch of Macleania spp. shrub, returning to aperch of 2 m height.

Social Interactions.—Few interactions withother hummingbirds were observed. One maleH. regalis was observed displacing, but notdirectly attacking or chasing, a male EcuadorianPiedtail (Phlogophilus hemileucurus) at feedingsites (in ridgetop shrubland above Yankuam), anda male attacked and chased a male Green-frontedLancebill (Doryfera ludovicae) when it hovered infront of ericaceous flowers in the same site;minutes later a D. ludovicae returned and hovered

Freile et al. N NATURAL HISTORY OF THE ROYAL SUNANGEL 87

in front of the same flowers without beingdisplaced.

No other aggressive behaviors were reportedalthough other hummingbird species occurred infeeding 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 andperching 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 tziıp! Wefailed to record both vocalizations.

A feeding male uttered an emphatic tchup orchıp! every 3.5–6 sec (deposited at www.xenocanto.org; XC 45910; Fig. 2A1), whereas other feedingcalls were uttered at shorter intervals (2–2.5 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 1.5–1.8 to 17–19 kHz (call2A1), and 1.5 to 21 kHz (call 2A2).

Two displaying males produced a fast chatteredseries of very high tEEp or jeet notes lasting 2.5–4.5 sec (XC 45912; Fig. 2B). This vocalizationcontained 19 notes of 0.10–sec mean duration(0.07–0.15 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 45910–45915) 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 edgeand 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 first’s display flightconstantly vocalizing (Fig. 3A). A somewhatsimilar display was also observed with a maleascending 10–12 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 2,500–9,999 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 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 mightalso 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

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

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.

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 45910–45912; figures by Paolo Piedrahita.

Freile et al. N NATURAL HISTORY OF THE ROYAL SUNANGEL 89

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 discrepanciesindicate 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 eventsof 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 male-

male 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 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 in

stunted ridgetop shrubland and paramo-like hab-

itats at Las Orquıdeas sandstone ridges. At least

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 40-

m 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 seeminglypreferred 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

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 Buitron-Jurado).

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

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 ,2–3 km between both study sites. Thismight indicate 1–1.5 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 ofapparently 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 ridgeswill 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. regalis. 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 by

Fundacion Arco Iris of Loja, and supported by Conserva-

tion International. We especially thank Robert Jimenez for

diligent assistance in field work. Oswaldo Jadan thanks

Zhofre Aguirre and Bolıvar Merino of Herbario LOJA, Jose

M. Manzanares, Charlotte Taylor, and Ron Liesner of

Missouri Botanical Garden, and Alfonso Garmendia of

Universidad Politecnica 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

Ordonez and Fundacion 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 Buitron, Paolo

Piedrahita, and Elisa Bonaccorso was supported by

Secretarıa Nacional de Ciencia y Tecnologıa (SENACYT)

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.

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