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An apparent hybrid Heliodoxa hummingbird from the West Andes of Colombia.

Conservación Colombiana – Número 17 –octubre de 2012 1
Número 17 • octubre 2012
Aves de Colombia 2012
Birds of Colombia 2012
©2011 Fundación ProAves • Bogotá • Colombia • ISSN 1900–1592
2 Conservación Colombiana – Número 17 – octubre 2012
Conservación Colombiana
Journal for the diffusion of biodiversity conservation activities in Colombia.
Revista de difusión de acciones de conservación de la biodiversidad en Colombia.
ISSN 1900–1592. Non–profit entity no. S0022872 – Commercial Chamber of Bogotá
ISSN 1900–1592. Entidad sin ánimo de lucro S0022872 – Cámara de Comercio de Bogotá.
Edición Octubre 2012. Publicado 10 de noviembre de 2012.
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Conservación Colombiana – Número 17 –octubre de 2012 3
Contenidos –– Contents
Conservación Colombiana 17
Revision of the status of bird species occurring or reported in Colombia 2012.
Revisión del estatus de las especies de aves que han sido reportadas en Colombia 2012.
Thomas Donegan, Alonso Quevedo, Paul Salaman & Miles McMullan
Vocal differentiation and conservation of Indigo-crowned Quail-Dove Geotrygon
Diferenciación en la vocalización de Geotrygon purpurata y evaluación de su estado de conservación.
Thomas Donegan & Paul Salaman
An apparent hybrid Heliodoxa hummingbird from the West Andes of Colombia.
Un aparente híbrido del género Heliodoxa en la Cordillera Occidental de Colombia.
Thomas Donegan & Liliana Dávalos
Dos nuevas especies de aves para Colombia en el departamento del Guainía.
Two new bird species for Colombia from the department of Guainía.
Alonso Quevedo & Juan Carlos Luna
New records of Forster’s Tern Sterna forsteri for Colombia.
Nuevos registros de Sterna forsteri para Colombia.
Forrest Rowland & Bernard Master
Primera fotografía en su habitat y nuevo avistamiento del Cucarachero de Santa Marta
Troglodytes monticola, especie en Peligro Crítico.
First photograph in its habitat and new sighting of the Santa Marta Wren Troglodytes monticola, a
Critically Endangered species.
Juan Carlos Luna & Alonso Quevedo
Primer registro del Hornero del Pacífico Furnarius (leucopus) cinnamomeus en
First record of Pacific Honero Furnarius (leucopus) cinnamomeus in Colombia.
Juan Carlos Luna
Records of two escaped species of parrots for Colombia.
Registros de dos especies de loros exóticos en Colombia.
Oswaldo Cortés & Thomas Donegan 35-37
Corrigenda: Conservación Colombiana 15 37
Note on the identification of Lesser Nighthawk Chordeiles acutipennis in northern
Nota para la identificacion de Chordeiles acutipennis en el norte de Colombia.
Andrew R. Collins
A new group name for the Chachalacas (Aves: Cracidae: Ortalis).
Un nuevo nombre para el grupo de las chachalacas (Aves: Cracidae: Ortalis)
Thomas Donegan 41-44
First Record for the Black–and–white Tanager Conothraupis speculigera in Colombia.
Primer registro del frutero blanco y negro Conothraupis speculigera en Colombia.
Yojanan Lobo–y–HenriquesJC, John Bates & David Willard 45-51
Instrucciones para autores
Instructions for authors. 52-54
20 Conservación Colombiana – Número 17 – octubre 2012
An apparent hybrid Heliodoxa hummingbird
from the West Andes of Colombia
Un aparente híbrido del género Heliodoxa en la Cordillera Occidental de Colombia
Thomas Donegan1 & Liliana Dávalos2
1 ProAves Foundation, Southmead, The Vale, London N14 6HN. Email:
2 Department of Ecology and Evolution and Consortium for Inter-Disciplinary Environmental Research, 650 Life Sciences,
State University of New York at Stony Brook, Stony Brook, NY 11794, USA. Email:
Details and photographs are presented of an individual mist-
netted in the West Andes of Colombia in 1997, which seems
possibly to represent a combination between Fawn-breasted
Brilliant Heliodoxa rubinoides and Empress Brilliant H.
Presentamos medidas y fotografías de un aparente híbrido
capturado en la Cordillera Occidental de Colombia en
1997. Este individuo parece posiblemente representar una
combinación entre Heliodoxa rubinoides y H. imperatrix.
During August to September 1997, the authors studied birds
in Tambito Nature Reserve, El Tambo municipality, dpto.
Cauca in the Western Cordillera of Colombia (c. 02°30'N,
77°00'W), using mist-nets and observations (see Donegan &
Davalos 1999 for details). In a study site located in mature,
emergent secondary growth with abundant palms at 1,600 m
(Site 2 in Donegan & Dávalos 1999), the most abundant
genus during netting was Heliodoxa, with 30 Empress
Brilliant H. imperatrix and 9 Fawn-breasted Brilliant H.
rubinoides captured in four days. In addition, a further
hummingbird was captured which could not be identified. A
discussion of this individual was included in an unpublished
expedition report provided to donors and presence of a
hybrid Heliodoxa was briefly mentioned in Donegan &
Dávalos (1999), but no details have been published to date.
Hybrids are more frequent among the hummingbirds than
many other bird families. Many dubious taxa have been
described based on apparent hybrid types for both Colombia
(Hilty & Brown 1986) and South America (Remsen et al.
2012). The study and diagnosis of novel hummingbird
hybrids continues today (e.g. Banks & Johnson 1961, Graves
& Zusi 1990, Graves & Newfield 1996, Graves 1990, 1996,
1997a,b, 1998a,b, 1999a,b, 2001, 2003a,b,c, 2004, 2006,
2007a,b). Among reported study cases are both intergeneric
(Graves & Zusi 1990) and intrageneric (Graves 2003b)
hybrids considered to involve members of the genus
Heliodoxa. This paper discusses a previously undescribed
morphotype that apparently results from a novel intrageneric
combination in this genus.
Description of individual
The bird is illustrated in Figs. 1 to 5, with a morphological
description appearing in Table 2 and biometric data in the
Appendix. The photographs are of low quality, taken on a
late 1980s model SLR camera without flash and with print
film. As a result, they are not comparable to those taken by
ornithologists today, although the key features of the bird are
clearly visible. The individual is identified to the genus
Heliodoxa on account of its straight and black bill being
tapered to the skull, with feathers extending forwards on the
bill, unmodified remiges, unspotted rectrices, general size
(see Appendix), presence of a gorget, forked tail and
glittering crown. No morphological features of other
hummingbird genera were noted, so species of other genera
were not considered in detail here. Biometric measurements
are set out in the Appendix. The bird was in primary, tail and
body moult.
The most striking morphological feature of this individual
was its glittering silvery blue gorget (Fig. 1). This, combined
with the absence of any rufous on the malar (Fig. 5), make it
a presumed adult male and preclude the two Heliodoxa
species confirmed at the locality. Green-crowned Brilliant H.
jacula also occurs in the region, but at lower elevations, and
has a blue gorget. However, various features are inconsistent
for H. jacula, particularly the tail coloration (which is dusky,
not navy blue), light brown tail shafts (not concolor with the
tail), the shade of blue on the gorget and large gorget size
(cf. Figs. 9-10), some of which are more reminiscent of H.
rubinoides (cf. Fig. 8).
Differences from Heliodoxa species occurring on the Pacific
slope of Colombia are detailed further in Table 1.
New hummingbird species continue to be discovered (e.g.
Cortés-Diago et al 2007) and reported (Valdes-Velásquez &
Schuchmann 2009) from the West Andes. The presence of a
fourth sympatric Heliodoxa species at the same
locality would be unprecedented for the genus and seems
improbable. This individual is more likely to have had a
hybrid origin based on these considerations alone, and shows
various intermediate features between species present at the
study locality (Table 2).
Conservación Colombiana – Número 15 – octubre de 2012 21
Table 1. Differences between presumed hybrid Heliodoxa
and species occurring in the region.
Species Differences
Male Empress has a pink (not blue) gorget, a
longer and green tail, no brown in the wing or
tail, no white on the undertail coverts, no
speckling on the breast and a more tapered head
with more extensive bill feathering. The BMNH
collection includes two specimens of imperatrix
with a silver gorget, showing bluish or pinkish
sheens but these do not fully match that of the
presumed hybrid either.
Brilliant H.
Fawn-breasted has a pink (not blue) gorget, a
more extensively brown (less extensively green)
breast, a light brown tail which is much shorter
and lacks white undertail coverts. Its bill is more
tapered, downcurved and thicker. Tail, bill and
body lengths and mass of Fawn-breasted fall out
of range (the latter being too short or low).
Brilliant H.
Green-crowned’s blue gorget is smaller and
shaped as a crescent moon, rather than a larger
rotated D and is of a different shade of blue. It
has no brown in the wing. Mass of Green-
crowned is too low and other measurements are
at the extreme of variation in this species.
Other Colombian Heliodoxa (aurescens, leadbetteri,
schreibersi, gularis) are not in range and most show
distinctive marks not found on this individual, so are not
considered plausible parental species.
Key features for purposes of hybrid diagnosis are those
potentially referable only to a single species. The brownish
wing covert edgings, brownish leading primary (Fig. 1),
dusky tail (Fig. 4) and brown ventral tail shaft coloration
(Fig. 3) are exclusively H. rubinoides features among the
three possible parental species considered here. Brown and
reddish pigments are often expressed in hybrid
hummingbirds where they are available as a parental
character (Banks & Johnson 1961, Graves & Newfield 1996,
Graves 2004). The blue gorget is a feature solely of jacula.
However, the shade is lighter and more silvery. Some
imperatrix specimens at BMNH and photographs available
online show silver gorgets suggesting that silver coloration
may be a rare variation or polymorphy within the species.
The novel "bluing" of feathers is a particular feature of
hummingbird hybridisation (Graves 1998a, 1999b) and so
jacula cannot be considered as necessarily a parent.
Most biometrics fall within the range of all species, although
they are more typical of imperatrix, with various measures at
the extreme for both rubinoides and jacula (see Appendix).
No feather-by-feather measures of rectrices of the nature
undertaken by Graves (op. cit.) were undertaken due to a
lack of familiarity of the authors with this method for
assessing hybrid hummingbirds at the time of the study.
Although the central feathers were missing due to moult, the
two outer tail feathers on each side demonstrate a deeply
forked shape (Figs. 4-5), more like imperatrix (Fig. 7; males
have longer tails) or jacula (Fig. 9) and does not at all
resemble that of rubinoides (Fig. 6). The biometrics and
particularly long body and large mass fall only within the
range of imperatrix (Appendix).
Table 2. Morphological description of putative hybrid, with
an indication of possible parental species for each character.
Feature Possible species
Bill black and straight. Head
tapered into the bill. All.
Glittering green on upperparts All.
White postocular All.
Blue gorget on throat Closest to jacula's blue but
lighter / more silver. Close to
imperatrix's silver morph
gorget, but bluer.
Size of gorget on throat rubinoides or imperatrix (cf.
Figs. 9-10).
Fawn spotted breast coloration Most like rubinoides but more
extensively green. Male jacula
have a fawn base to glittering
green breast feathers, but fawn is
almost imperceptible unless a
bird is stretched or moulting.
White upper undertail coverts Unusual in both jacula (e.g. Fig.
10) and imperatrix.
Dull shade of glittering green on
lower underparts jacula.
Lower undertail coverts
glistening green jacula or imperatrix.
Thin white feathering on black
legs All.
Primaries, secondaries and
tertials “dusky” imperatrix or jacula.
Brownish / dusky tinge to some
wing coverts rubinoides.
Brown tinge on leading primary rubinoides.
Tail length jacula or between imperatrix and
Deeply forked tail jacula or imperatrix.
Coloration of tail feathers: dusky
with brownish tinge None; rubinoides has a brownish
tail, others have glittering
green/blue tails.
Central two tail feathers copper
brown rubinoides.
Copper rump jacula or rubinoides.
Tail feather shafts on underside
are brown rubinoides.
Body length imperatrix.
Wing length All.
Tail length imperatrix or jacula.
Bill length All.
Mass imperatrix.
22 Conservación Colombiana – Número 17 – octubre 2012
Figure 1. Putative hybrid Heliodoxa: frontal view. Photo: T.
Figure 2. Putative hybrid Heliodoxa: dorsal view. Photo: T.
Figure 3. Putative hybrid Heliodoxa: ventral view.
Note white undertail and brown tail shafts. Photo: T.
A male H. jacula x H. imperatrix specimen is to be found at
the British Natural History Museum (no. 1902.13.2211) and
was inspected by TMD in September 1997. The same
specimen was discussed in detail recently by Graves (2004),
whose diagnosis we agree with. Graves (2004) considered
the BMNH skin to represent "the only known instance of
intrageneric hybridisation in Heliodoxa". It differs from the
Tambito individual in having a green and relatively longer
tail and total body length, no brown in the wing coverts and
no noticeably paler base coloration or spotting of the
underparts (our measures: Tail 64 mm, Wing 77; Body 154
mm, broadly close to Graves (2004)'s measures in each
instance). Its blue violet gorget is noteworthy (given that
imperatrix has a pink gorget) and an instance of
predominating of blue coloration in a hybrid. Although not
all hybrid combinations would necessarily look similar (as a
result of meiosis in each parental species and sex-related
differences) both this and the Tambito bird are putative
males and specimen 1902.13.2211 is a rather different bird
from that captured at Tambito.
Figure 4. Putative hybrid Heliodoxa: dorsal view showing
tail shape. Photo: T. Donegan
Figure 5. Putative hybrid Heliodoxa: side view showing
head shape (out of focus). Photo: T. Donegan
We have no specimens or molecular data that could provide
definitive conclusions as to the hybrid or other origin of this
organism. The possibility of it representing a bizarre
aberration or even a new species cannot therefore be
Conservación Colombiana – Número 15 – octubre de 2012 23
excluded. A cross between H. jacula and H. rubinoides
would produce all observed characters except the large body
size and mass: a hypothesis for this combination would be
supported by the blue gorget and lack of brighter glittering
green plumage on the lower underparts. Given that H.
rubinoides and H. imperatrix were both common at this
locality and together can produce all observed features
except the blue gorget, the latter feature could represent an
example of 'bluing' of the silvery throat morphotype of
imperatrix in a hybrid combination between these two
species. We hope that with details included in this
publication, ornithologists and birders who visit the West
Andes can look out for other individuals demonstrating this
Figure 6. Tail view of male Fawn-breasted Brilliant H.
rubinoides from same expedition. Photo: T. Donegan
Figure 7. Tail view of female Empress Brilliant H.
imperatrix from same expedition. Photo: T. Donegan
Many thanks to Alex Cortés, the late Álvaro Negret, Luis
Alfonso Ortega, Gustavo Lacera, Mark Mulligan, Quintin
Lame and family, Ville Vepsäläinen and Paul Salaman for
support of our fieldwork. Fundación Proselva, Universidad
del Cauca and Corporación Autónoma Regional del Cauca
supported the project. Grants from the J. W. Bennet Award,
the G. R. N. Minchin Award and the Donald Robertson
Award assisted our research in Tambito. Data from San
Lucas and Yariguíes are based on research supported by
persons listed in Salaman et al. (2002), Donegan (2012) and
Donegan et al. (2010). Thanks to F. Gary Stiles, Á. Negret
and P. Salaman for discussing this specimen with us and the
latter for his comments on the MS and previous unpublished
expedition report extract on which this paper was partially
Figure 8. Frontal view of male Fawn-breasted Brilliant H.
rubinoides from same expedition. Photo: T. Donegan
Figures 9 a-b. Left: Moulting adult male nominate H.
jacula from Alto Honduras, Serranía de los Yariguíes (not
using flash). Note less extensive and dark blue gorget.
Photo: B. Huertas / Proyecto YARE, January 2006. Right:
Adult male nominate H. jacula from Cerro de la Paz,
Santander (using flash). Note less extensive and dark blue
gorget. Subspecies jamiesoni of the West Andes can have a
larger gorget but it does not approach that of the individual
illustrated in Figs. 1-6 in size. Photo: T. Donegan / Proyecto
EBA, January 2003.
24 Conservación Colombiana – Número 17 – octubre 2012
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Weller, A.-A. 2007. A new species of Eriocnemis
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C., Roa, C., Parra, R., Turner, C., Sharp, M. & Huertas,
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Conservación Colombiana – Número 15 – octubre de 2012 25
Appendix: biometric data
For each taxon, data are as follows: mean ± standard
deviation (lowest recorded value–highest recorded value) (n
= sample size). Data are based entirely on live mist-net
capture data. H. rubinoides and H. imperatrix data all
originates from the same study at Tambito by the authors. H.
jacula data comes from studies by TMD and others in
Serranía de San Lucas (see details in Salaman et al. 2002,
Donegan 2012) and Serranía de los Yariguíes, Colombia
(see details in Donegan et al. 2010). Live data and
photographs in this paper of H. jacula involve the nominate
subspecies rather than H. j. jamiesoni of the West Andes, but
inspection of specimens shows them to be only mildly
differentiated in biometrics.
Species /
Individual Body
(mm) Wing
(mm) Tail
(mm) Bill to
hybrid 138 78 55 26 9.2
144.6 ±
71.5 ±
3.7 (63-
59.6 ±
25.3 ±
1.4 (23-
9.3 ±
0.7 (8.5-
154.5 ±
74.6 ±
2.1 (72-
63.8 ±
24.3 ±
1.1 (23-
9.9 ±
1.1 (9.1-
121.8 ±
69.5 ±
3.7 (63-
42 (n=1) 23.4 ±
1.4 (21-
7.9 ±
0.6 (7.5-
122.0 ±
68.8 ±
4.1 (64-
42 (n=1) 24.3 ±
1.2 (23-
H. jacula
All 118.3 ±
67.6 ±
4.9 (57-
43.8 ±
4.9 (37-
26.4 ±
1.6 (23-
7.0 ±
0.6 (5.9-
H. jacula
Males 126.7 ±
72.3 ±
5.4 (62-
49.8 ±
4.1 (44-
25.9 ±
1.5 (24-
7.2 ±
0.7 (5.9-
Conservación Colombiana – Número 15 – octubre de 2012 55
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A new hummingbird species, the Gorgeted Puffleg (Eriocnemis isabellae, sp. nov.), is described from the Serranfa del Pinche, an unexplored isolated mountain massif in the Department of Cauca, southwest Colombia (02°16'04.18"N, 77°21'26.41"W, 2800 m a.s.l.). This species represents a distinct new member of the genus Eriocnemis and inhabits the cloud and temperate forest zone of the Serrania. Although it can be easily diagnosed as a member of Eriocnemis by the conspicuous white tibial tufts, violet blue under-tail coverts, and a bifurcated blue black tail, it widely differs in plumage from most other species of the genus, having the facial area, crown, and nape blackish tinged yellow olive green, and a distinctively bicolored, enlarged, iridescent throat patch with a violet blue centre and green sides. Some plumage characteristics are shared with other members of the genus (i.e., E. vestitus, E. nigrivestis). The new taxon is cologically associated with elfin forest, occupying a very small range at steep slopes along mountain ridges, The relative inaccessibility of this habitat, in combination with conservation measures supported by local authorities, organizations, and inhabitants, raises hope for the future protection of this unique and critically endangered trochilid.
Coeligena purpurea Gould, 1854 is shown to be a hybrid between Coeligena coeligena and Coeligena prunellei. The geographic distribution of the parental species suggests that the two hybrid specimens were collected in the Eastern Cordillera of the Colombian Andes. The hybrids exhibit a blended mosaic of plumage characters of the postulated parental species. External measurements of the hybrids fall within the cumulative ranges of characters of the parental species.
Eriocnemis soderstromi (Butler 1926) is shown to be a hybrid between E. nigrivestis, an endemic of the Volcan Pichincha region in Ecuador, and E. luciani, an inhabitant of timberline Andean forest from southern Colombia to Bolivia. This is the first report of intrageneric hybridization among the nine species of Eriocnemis. Plumage characters of the hybrid appear to be a blended intermediate of those of the parental species. External measurements of the hybrid are intermediate of those of parental species.
Amazilia distans Wetmore & Phelps, 1956, is believed to be a hybrid between Hylocharis cyamis and Amazilia fimbriata. The hybrid, collected in Estado Tâchira, Venezuela, exhibits a blended mosaic of plumage characters of the parental species. External measurements of the hybrid fall between the character means of the parental species which overlap in size.
An intergeneric hybrid hummingbird, Aglaiocercus kingi X Metallura tyrianthina, is described. External measurements of the hybrid are intermediate of those of the parental species. Back plumage iridescence is bluer (511 nm) in the hybrid than in either of the parental species (553-571 nm). This color shift is thought to be caused by a developmental aberrancy or mutation which affects melanin granules that produce iridescence in feather keratins.
Calliphlox iridescens Gould, 1860 is hypothesized to be a hybrid between Calliphlox amethystina and Chlorostilbon aureoventris. The hybrid, collected at Nova Friburgo, Rio de Janeiro, Brazil, exhibits a blended mosaic of plumage characters of the presumed parental species. External measurements of the hybrid fall between the character means of the parental species and approach the values expected from least squares regression of parental measurements.
Plumage pattern, plumage color, and external morphology of two specimens of hybrid hummingbird (Calypte anna X Stellula calliope) are described. The parental species differ substantially in size, and the hybrids are similar to the averages of character means of the parental species. Expression of parental plumage characters in the hybrids varies, particularly the color and configuration of the frontlet and gorget. The possible effects of egg volume on hybridization potential are discussed.
Calothorax decoratus Gould 1860 is shown to be a hybrid between Acestrura heliodor and Acestrura mulsant. Plumage characters of the hybrid are a blended combination of those of the parental species. External measurements of the hybrid approximate the values expected from least squares regression of parental measurements. This is the first conclusively documented case of hybridization among the diminutive woodstars (Acestrura).
Lesbia ortoni Lawrence, 1869, collected in the Quito Valley, Ecuador, is shown to be a hybrid between Lesbia victoriae and Ramphomicron microrhynchum, sympatric inhabitants of Andean forest edge and shrublands from Colombia to Peru. The hybrid exhibits a blended mosaic of plumage characters of the parental species. Although the parental species differ significantly in size, the external measurements of the hybrid approximate those predicted by least squares regression.
Cyanomyia salvini Brewster, 1893, collected in Sonora, Mexico, is shown to be a hybrid between Amazilia violiceps ellioti and Cynanthus latirostris magicus, whose breeding ranges overlap extensively in northwestern Mexico. This specimen represents the only known intergeneric hybrid between species currently placed in Amazilia and Cynanthus (Sibley and Monroe 1990).