Breeding biology of the Sparkling Violetear Colibri coruscans in Quito

Abstract

Sparkling Violetear Colibri coruscans has a wide geographic distribution and is generally abundant. However, knowledge about its breeding biology is scarce and often limited to anecdotal observations. Therefore, in this study we present new breeding data in Quito and surrounding valleys, based on descriptions and observations of 45 broods in 39 nests (some nests were reused) in urban and suburban habitats, and 11 broods in woodland patches near Quito. We provide data on nest and egg measurements, maternal care, nestlings' development, and reproductive behavior. We studied nests reported on social media or discovered by the authors between, with binoculars, photo and video cameras. Nests were open cups built in native and exotic vegetation or on artificial structures, protected by a roof, at 1.2-8 m above the ground. Nests under roofs suffered little deterioration, allowing their reuse. Nest materials in the outer layer included vegetal fibers and moss, whereas the inner chamber had soft vegetal material such as hairy achenes. Nests in woodlands were covered in lichens in the outer layer. Clutch comprised two white ellipsoidal eggs. Incubation lasted 14-16 days, and offspring left the nest after 20-31 days. Only the female incubated and reared the nestlings. In some cases, males' courtship and/or feeding territory overlapped the females' nesting territory. We observed breeding activity year-round, with a peak from November to April, corresponding to the second half of the short rainy season, the short dry season and the first half of the long rainy season.

Full text

ARTÍCULO/ARTICLE
Musschenga, M.A., Juiña, M., & Cadena-Ortiz, H. (2022). Breeding biology of the Sparkling Violetear
Colibri coruscans in Quito. Revista Ecuatoriana de Ornitología, 8, 51–66.
Breeding biology of the Sparkling Violetear Colibri coruscans in Quito
Maartje A. Musschenga1,2,*, Mery Juiña3,4, Héctor Cadena-Ortiz2,3
1 Universidad Central de Ecuador, Facultad de Filosofía, Carrera de Pedagogía de las Ciencias Experimentales
Química y Biología, Gaspar de Carvajal y Avenida La Gasca, Ciudadela Universitaria, Quito, Ecuador.
2 Pajareando Ando Ecuador (Colectivo de Observadores de Aves).
3 Instituto Nacional de Biodiversidad (INABIO), Calle Rumipamba 341 y Av. de Los Shyris, Quito, Ecuador.
4 Fundación COFIVAL, 24 de Agosto s/n y esquina, Lumbisí, Quito, Ecuador.
*Corresponding author, e-mail: maartmus@hotmail.com
Editado por/Edited by: Harold F. Greeney
Recibido/Received: 3 March 2021 Aceptado/Accepted: 23 August 2022
Publicado en línea/Published online: 13 December 2022
Biología reproductiva del Orejivioleta Ventriazul Colibri coruscans en Quito
Resumen
El Orejivioleta Ventriazul Colibri coruscans tiene distribución geográfica extensa y es generalmente abundante. Sin
embargo, el conocimiento sobre su biología reproductiva es escaso y en su mayoría limitado a observaciones anecdóticas.
Por ello, en este estudio presentamos nuevos datos sobre su reproducción en la ciudad de Quito y valles circundantes, a
partir de descripciones y observaciones de 45 nidadas en 39 nidos urbanos y sub-urbanos (algunos nidos fueron reusados)
y 11 nidadas en bosques cerca de Quito. Presentamos medidas de los nidos y huevos, e información sobre cuidado maternal,
desarrollo de los polluelos y comportamiento reproductivo. Monitoreamos nidos que fueron reportados en redes sociales o
descubiertos por los autores entre 2009–2015 y enero 2018–mayo 2020, con ayuda de binoculares, cámaras fotográficas y
de video. Los nidos tuvieron forma de taza y fueron construidos en vegetación nativa y exótica, o en estructuras artificiales
protegidos por un techo, y se ubicaron entre 1,2–8 m sobre el suelo. Los nidos construidos bajo techos artificiales se
mantuvieron en buen estado y fueron reusados. Los materiales del nido incluyeron musgo y fibras vegetales en la capa
exterior y material vegetal suave como aquenios peludos en la cámara interior. Los nidos en bosques tenían además una
cobertura de líquenes en la superficie exterior. Las puestas fueron de dos huevos blancos, de forma elipsoidal. El periodo
de incubación fue de 14–16 días, y los polluelos abandonaron el nido después de 20–31 días. Solamente la hembra se ocupó
de la incubación y crianza de los polluelos. En algunos casos, el territorio de anidación de la hembra coincidió con el
territorio de alimentación y/o de cortejo de un macho. Encontramos actividad reproductiva durante todo el año, con un pico
entre noviembre y abril, correspondiente a la segunda mitad de la temporada lluviosa corta, la temporada seca corta y la
primera parte de la temporada lluviosa larga.
Palabras clave: colibrí, Colibri coruscans, comportamiento reproductivo, cuidado parental, hábitats urbanos, huevos,
incubación, nido, pichones.
Abstract
Sparkling Violetear Colibri coruscans has a wide geographic distribution and is generally abundant. However, knowledge
about its breeding biology is scarce and often limited to anecdotal observations. Therefore, in this study we present new
breeding data in Quito and surrounding valleys, based on descriptions and observations of 45 broods in 39 nests (some
nests were reused) in urban and suburban habitats, and 11 broods in woodland patches near Quito. We provide data on nest
and egg measurements, maternal care, nestlings’ development, and reproductive behavior. We studied nests reported on
social media or discovered by the authors between 2009–2015 and January 2018–May 2020, with binoculars, photo and
video cameras. Nests were open cups built in native and exotic vegetation or on artificial structures, protected by a roof, at
1.2–8 m above the ground. Nests under roofs suffered little deterioration, allowing their reuse. Nest materials in the outer
layer included vegetal fibers and moss, whereas the inner chamber had soft vegetal material such as hairy achenes. Nests
in woodlands were covered in lichens in the outer layer. Clutch comprised two white ellipsoidal eggs. Incubation lasted
14–16 days, and offspring left the nest after 20–31 days. Only the female incubated and reared the nestlings. In some cases,
males’ courtship and/or feeding territory overlapped the females’ nesting territory. We observed breeding activity year-
round, with a peak from November to April, corresponding to the second half of the short rainy season, the short dry season
and the first half of the long rainy season.
Keywords: Colibri coruscans, eggs, hummingbird, incubation, nest, nestlings, parental care, reproductive behavior, urban
habitats.

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INTRODUCTION
Sparkling Violetear Colibri coruscans (Trochilidae) has a broad geographic distribution from Venezuela to
northern Argentina, including Guyana and northern Brazil (Züchner et al., 2020). Throughout this range, it
occupies forest edges, open woodland, gardens, cropland and páramo, between 1700–4500 m a.s.l.; sometimes
descending to 200 m a.s.l. (Züchner et al., 2020). In Ecuador, it is common in the inter-Andean valleys at 1000–
3500 m a.s.l., in open and semi-open areas (Freile & Restall, 2018). In Quito, it is the fourth most abundant bird
species after Eared Dove Zenaida auriculata, Rufous-collared Sparrow Zonotrichia capensis, and Great Thrush
Turdus fuscater (Cisneros-Heredia et al., 2015).
Despite being fairly common, the breeding biology of C. coruscans is generally poorly understood and known
only from few anecdotal descriptions (Moore, 1947; Hainsworth, 1977; Schmidt-Marloh & Schuchmann, 1980;
de la Peña, 2001; Gonzáles & Castañeda, 2020). Ruschi (1965) studied captive C. coruscans in Río de Janeiro,
Brazil, and concluded that the female builds the nest, the eggs weigh 8–9 gr, incubation lasts 15 days, the two
eggs hatch the same day or on consecutive days, and nesting period is 22 days. Zerda-Ordóñez (1994) performed
an extensive study on its breeding biology in the Botanical Gardens of Bogotá, Colombia, and found that C.
coruscans establishes three types of territories: a display (males), nesting, and feeding territory (Zerda-Ordóñez,
1994). Nest building lasted c. 7 days, incubation of two eggs 16 days, incubation constancy (i.e., percentage of
time spent on the nest during the day) was 69%, and the offspring left the nest after 21 days (Zerda-Ordóñez,
1994). Nests described contained moss (Schäfer, 1954; Ruschi, 1965; Zerda-Ordóñez, 1994), were externally
covered in lichens (Schäfer, 1954), and had soft materials in the inner chamber (Schäfer, 1954; Zerda-Ordóñez,
1994).
In hummingbirds, the female carries out alone nest building, incubation and nestling care (Ortiz-Crespo, 2011).
However, Gonzáles & Castañeda (2020) observed a male C. coruscans perching around a nest, and there are
two additional anecdotal but controversial observations of incubating male C. coruscans (Moore, 1947; Schäfer,
1954; see Ortiz-Crespo, 2011). Further, no evidence of male involvement in breeding duties was found in other
studies of the species’ reproductive behavior; neither in the wild (Zerda-Ordóñez, 1994), nor in captivity
(Ruschi, 1965; Stoppelmoor, 2000).
Given the deficient knowledge about the breeding biology of C. coruscans, we present extensive information
on the species’ breeding biology and mating behavior, nests, eggs, nestlings’ development, and reproductive
success in Quito and surrounding valleys. Our field information was collected between 2009–2015 and 2018–
2020, and includes data about 45 active broods in urban and suburban habitats. To compare these nests with
those in a more natural landscape, we also collected data at montane forest patches near Quito, on the slopes of
Pichincha volcano. This study fills several gaps in our knowledge of C. coruscans breeding biology, especially
in suburban habitats. Further, we compare our results with previous breeding studies of the species and of other
hummingbirds.
METHODS
Study area
Between March 2009–November 2015, we studied nests near the residence of one of the authors in San Vicente,
Nayón (-0.173728, -78.446581), northeast of Quito, Pichincha province (Fig. 1). Between January 2018–May
2020, we visited nests reported on social media or discovered casually by the authors, in the following locations:
north Quito (-0.18323, -78.48445; c. 2800 m a.s.l.); San Rafael (-0.337207, -78.47705), La Armenia (-0.27052,
-78.4695), and Sangolquí (-0.33016, -78.4511) in Los Chillos valley, south-east of Quito; Cumbayá (-0.210725,
-78.44109), east of Quito; and San Antonio de Pichincha (0.004568, -78.44677), north of Quito (Fig. 1). Study
sites in urban and suburban areas ranged from 2350–2800 m a.s.l. Sampling effort was not standardized over
the study period.
The Quito area has an average annual temperature of 13.4 °C and a total annual rainfall of 1,175.2 mm, over
two rainy periods, a short one in October–November and a longer one in February–May (Pourrut et al., 1995).
All localities described in the valleys around Quito are moderately to highly urbanized, but contain remnants of

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native vegetation. Dry forest remains along creeks and steep slopes in the valleys of Cumbayá, Nayón, and San
Antonio (Museo Ecuatoriano de Ciencias Naturales, 2009), which is characterized by low-stature trees such as
Acacia macracantha (Fabaceae), dry shrubs of several species, succulents such as Agave americana
(Asparagaceae), and Opuntia sp. (Cactaceae). La Armenia, San Rafael, and Sangolquí are currently dominated
by exotic trees and planted gardens, with the introduced Eucalyptus sp. (Myrtaceae) being abundant; native
shrubland patches of high Andean shrub occur mostly along creeks and slopes, in which Baccharis sp.
(Asteraceae) and Mimosa sp. (Leguminoseae) are common (Museo Ecuatoriano de Ciencias Naturales, 2009).
Complementarily, in 2015 and 2020 we studied nests close to Yanacocha reserve (c. -0.11174, -78.58486),
northwest of Quito, in an area covered in montane forest and páramo at 3200–4400 m a.s.l., and along the dirt
road from Yanacocha to Alambi (c. -0.087288, -78.599637, c. 3100 m a.s.l.) (Fig. 1). In this area, forest patches
remain among cattle fields, separated by living fences of native shrubs, in which families like Melastomataceae,
Ericaceae, and Asteraceae are common.
Field sampling
Nests. Individual nests were assigned a number (N1–N50; Appendix 1, Supplementary Material). Consecutive
broods in the same nest are indicated with a letter (a-f) after the nest number (Appendix 2, Supplementary
Material). Coordinates were taken from Google Earth Pro.
For each nest, we recorded at least the location, substrate (human made or vegetation), date and nest stage.
Depending on time availability, access to the nest, and stage of occupancy, we further collected as much data
suggested by Cadena-Ortiz (2018): nest and egg measures, clutch size, nest height, nest materials, observations
on incubation and brooding sessions, provisioning rates, development of nestlings, and notes on behavior (see
below). Observations were made with binoculars (Vogelbescherming Buizerd 8x42), photo cameras (Panasonic
Lumix DMC-TZ60, Nikon Coolpix P900, Canon 5D Mark III), and video cameras (Sony Handycam HDR-
PJ200), placed in strategic locations at a distance of 2–5 m. We recorded time sessions with a stopwatch on a
smartphone Redmi M2003J15SS. We took measurements of nest, nestlings and eggs with a Pretul Vernier 1-
mm precision caliper. We measured nest and substrate height with a measuring tape or we estimated it when
height exceeded maximal length of the measuring tape.
When nests were accessible, we took the following measurements: inner diameter and outer diameter, outer
height and inner depth. Samples vary because measures of three nests (N6, N20, N21) are incomplete. Further,
two eggs were measured in N5, as well as a non-hatched egg in N17a. The latter was also checked for
irregularities and microbial infections, following Van der Burg (2017). Three nests (N2, N4, N5) were dried at
ambient temperature and the materials (seeds, feathers, moss, vegetal fibers, synthetic fibers, dry flowers)
carefully disaggregated and separately weighed on a digital balance with a precision of milligrams.
Nesting activity. At five nests (N1, N4, N5, N17a, N19) we took notes on the females’ and nestlings’ behavior,
as well as incubation, brooding and provisioning rates. Incubation sessions were observed with binoculars at
N5 from a c. 3 m lookout. To calculate incubation constancy, we used Skutch (1962) formula: T=100S/(R+S),
where S is the mean of sessions on the nest incubating and R the average of recesses (periods outside the nest).
At N4, brooding sessions were observed with binoculars and at N17a sessions were video-recorded.
Provisioning rates during and after brooding were video-recorded at a single nest (N19) and observed at four
nests (N1, N4, N5 and N17a).
The nestling period was determined at 15 nests (Appendix 1 in Supplementary Material). Nestlings of six nests
(N1, N4, N5, N17a, N19, N33) were photographed every 1–3 days to document their physical development.
Body, bill, wing, tarsi, and tail length were measured in six nestlings at N1, N4, N5, N17a, and N17b at least
once every week.
We calculated egg success (fledglings/total number of eggs) as an indicator for breeding success (Murray, 2000)
in 37 broods. The remaining broods were found at a later stage of occupancy or monitored over a short period,
and therefore were only included in analysis of hatching success (total number of eggs/number of hatchlings)
or fledging success (total number of nestlings/number of fledglings).
Mating and territorial behavior. We video-recorded copulation behavior at one nest (N17). We also observed
with binoculars five nests (N34–38) located in the courtship territory of three males and took notes about
territorial behavior of males, females and fledged young. We distinguished males from females by their behavior

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and plumage; only males perform aerial dive displays from a perch, and are slightly brighter and have larger ear
tufts (Zerda-Ordóñez, 1994; Freile & Restall, 2018; Clark et al., 2018).
Figure 1: Study area of the breeding biology of Sparking Violetear Colibri coruscans in Quito, surrounding valleys and a
montane rainforest reserve northwest of Quito. A) San Antonio de Pichincha; B) Yanacocha Reserve; dashed line B-C)
Yanacocha-Alambi road; D) north Quito; E) Nayón; F) Cumbayá; G) La Armenia; H) Sangolquí; I) San Rafael. Adapted
from Google, n.d., https://www.google.com/maps/place/Quito,+Ecuador/@-0.1865938,-78.5706257,11z. All rights
reserved by Google 2020. Adapted with permission of the author.
RESULTS
Mating and territorial behavior
We observed one copulation in a small urban garden in north Quito (N19). Before copulation, the male sat on a
clothesline and emitted a chip call. Suddenly, while still vocalizing, he turned his head to the opposite direction,
and later turned it completely at a 45-degree angle. Meanwhile, he spread his wings while vibrating them, and
extended his ear tufts. Once his ear tufts were fully extended, he held his tail and head in an upright position
and vibrated his wings close to the body; vocalization stopped. Then, while turning completely at c. 100-degree
angle, he spread his wings once more, and vibrated them again close to the body. He produced a few soft calls.
Then the female perched on the clothesline, with her body at a 90-degree angle to the male’s head, but her head
turned towards his. The male approached her flying and landed on her. During copulation, which lasted c. 2 s,
the female spread her wings. Afterwards, the male flew away while vocalizing. The female stayed on the
clothesline, and briefly lifted her tail four times. Then she vibrated her wings close to her body for c. 1 s, and
lifted her tail twice again.
We also observed five nests (N34–38) located in the courtship territory of three males in La Armenia. Nests
were 13–18 m from each other, in native Baccharis sp. (Asteraceae) and Mimosa sp. (Fabaceae) shrubs,
bordering a quiet street. Among the native shrubs, many introduced Leonotis sp. (Lamiaceae) grew. The males
usually spent their time singing from a perch. When foraging near the nests, they were tolerated by the nesting

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females. During c. 20 h of observation in a 3.5-month period (November 2019–March 2020), we observed twice
a joint defense of the nesting territory by a female and a male against another C. coruscans individual. Usually
only the female or only the male chased away conspecifics or other hummingbird species, such as Black-tailed
Trainbearer Lesbia victoriae. In Nayón (N22–30), males also had their courtship territory 10–30 m from active
nests, and here in one occasion we observed a male and a female defending a nest, for c. 5 min, against a Turdus
fuscater. We did not observe males in the immediate surroundings of the remaining nests studied.
Nests in urban and suburban areas
All 39 nests were open cups; 6 were located in north Quito, 6 in San Antonio de Pichincha, 2 in San Rafael, 1
in Sangolquí, 5 in La Armenia, 10 in Nayón, and 9 in Cumbayá (Fig. 1). Nineteen nests were found in gardens
and parks, 17 along secondary roads and streets, and 3 inside buildings.
Thirty-four nests had vegetation as substrate: 11 nests in native shrubs and trees of four different families, and
23 nests in ornamental (introduced) species of nine families (see Table 1). Five nests were located on human-
made structures under a protecting structure or roof (Fig. 2): the engine of an electrical garage door; hot water
tubes; security bars at a door; a metal wire clothesline; and a hanging flower pot holder made of vegetal fibers
and metal. Mean heights and measures of the nests are available in Table 2.
In general, the outer layer of the nests contained moss (Fig. 2), but two nests contained winged seeds of Tecoma
stans (Bignoniaceae). The inner chamber of the nests was covered with soft material such as Taraxacum
officinale (Asteraceae) seeds. Materials of three nests (N2, N4, N5) analyzed included T. officinale seeds, moss,
vegetal fibers, dried Callistemon citrinus (Myrtaceae) flowers, feathers, synthetic fibers, and unidentified seeds
(Table 3). Another nest (N36), which was found deteriorated and therefore not included in Table 3, contained
human hair (0.01 g) and spider web (0.04 g). In two other nests (N1, N4), spider web was used to attach them
to branches.
Table 1: Mean measures of Sparkling Violetear Colibri coruscans nests in Quito and surrounding valleys (Nayón,
Cumbayá, Los Chillos, San Antonio). Outer height refers to the vertical length of the cup. Substrate plant (Subst.) height
and nest height in m, remaining values in cm; diam. = diameter.
Subst. height
Nest height
Inner diam.
Outer diam.
Outer
height
Inner depth
Mean ± SD,
n
3.9 ± 2.8,
n = 21
2.1 ± 1.3,
n = 27
4.2 ± 0.4,
n = 13
6.2 ± 0.8,
n = 14
5.1 ± 1.6,
n = 15
2.7 ± 0.7,
n = 13
95% conf.
interval
2.7–5.1
1.6–2.6
3.9–4.4
5.7–6.7
4.2–6.0
2.2–3.1
range
1.6–15
1.2–8
3.5–5
5–8
2–8
1–4
One nest (N41), located in a suburban garden, was used four consecutive times in the same breeding season;
another (N17) was used 11 times in 30 months. Before reusing the nest, the female inspected it and added new
material. Since individuals were not marked, we do not know if it was the same female that reused the nest.
Nine nests (N2–N30) were built in the same location on the same substrate: a stolon of Cynodon dactylon
(Poaceae) hanging from the embankment of a secondary road in Nayón, between 2009–2015. See Appendix 1
in Supplementary Material for details on nest locations, substrate type, nest measures and nest materials.
Breeding season
We observed active nesting periods for 45 broods in 39 urban and suburban nests (Fig. 3). We found active
nests year-round, either in incubation, brooding and/or nestlings’ provisioning. Only 11 nests were active in
May–October, two of these were still active until November, while 35 were active exclusively in November–
April. March and April were the most active months with 10 and 11 active nests, respectively (see Appendix 2
in Supplementary Material).

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Figure 2: Eggs and nests of Sparkling Violetear Colibri corucans in Quito and surrounding valleys. N4 in a natural substrate
in Cumbayá, February 2018; substrate plant is a Ficus benjamina (Moraceae). N36 in a Baccharis sp. (Asteraceae) shrub
in La Armenia, December 2019. N25 in an artificial substrate: security bars at an entrance door in Cumbayá, April 2018.
N17a in a clothesline in north Quito, December 2018 (Maartje Musschenga).
Eggs
We observed eggs in 41 out of 45 broods studied. Thirty-eight of these clutches contained two eggs and two
clutches had only one egg. In the remaining brood, the first egg laid was depredated before the second was laid
(Appendix 2, Supplementary Material). Egg laying occurred at 24–48 h intervals (observed in N17a, N41a,
N41b). Incubation started when the second egg was laid, as observed in three broods (N40, N41a, N41b).
Incubation period observed in eight broods was 14–16 days (Appendix 2 in Supplementary Material for details).
All observed eggs were ellipsoidal and immaculate white (Fig. 2). Mean egg width was 0.9 cm (SD = 0.05; n =
3; range 0.9–1.0 cm), and mean length 1.5 cm (SD = 0.1; n = 3; range 1.4–1.6 cm). An unhatched egg had an
intact egg shell and contained healthy-looking yolk and albumen, not presenting any signs of microbial
infections.

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Figure 3: Breeding activity of 45 Sparkling Violetear Colibri coruscans broods in Quito and adjacent valleys in 2009–2015
and 2018–2020. Light gray: incubation; darker gray: brooding and nestlings provisioning. Not all broods were monitored
from incubation to fledging.
Table 2: Plant species used as nest substrate in 34 urban and suburban nests of Sparkling Violetear Colibri coruscans in
Quito and surrounding valleys. Five nests not included in this table had artificial structures as substrate. N refers to plant
species native to Ecuador, I to introduced species. Nine nests (N22–N30) in Cynodon dactylon (Poaceae) row were attached
to the same C. dactylon stolon, hanging from the embankment of a road.
Plant species
Family
No. of nests
Mimosa quitensis (N)
Fabaceae
3
Baccharis sp. (N)
Asteraceae
6
Citharexylum ilicifolium (N)
Verbenaceae
1
Chionanthes pubescens (N)
Oleaceae
1
Campsis radicans (I)
Bignoniaceae
1
Codiaeum variegatum (I)
Euphorbiaceae
2
Ficus benjamina (I)
Moraceae
1
Eucalyptus sp. (I)
Myrtaceae
1
Callistemon citrinus (I)
Myrtaceae
1
Bougainvillea sp. (I)
Nyctaginaceae
1
Rosa sp. (I)
Rosaceae
1
Hedera helix (I)
Araliaceae
1
Schefflera sp. (I)
Araliaceae
4
Dracaena sp. (I)
Asparagaceae
1
Cynodon dactylon (I)
Poaceae
9

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Maternal care
We only observed females attending their offspring. In general, before approaching the nest, females perched
on a branch or electrical wire at c. 15 m from the nest. During incubation, the female changed her position on
the nest and also moved her underbody periodically from side to side. During 13 h of observation, from 9h00–
16h00, spread over 5 days, one female (N5) remained 9 h 50 min incubating. The incubation constancy using
Skutch’s formula was 74.8%.
Siblings in the same brood hatched the same day in two nests (N1, N40) or 24 h after each other in two other
nests (N41a, N41b). In N4, the female stayed 48% of 6 h 27 min brooding on day 5 since hatching, whereas in
N17a the female spent 51% of 6 h on day 6 since hatching. In N41a, the female brooded the nestlings during
daytime until day 8 since hatching and at night until day 9.
Nestling provisioning occurred every 55 min on day 4 since hatching in N17a; totaling seven visits in 6 h 28
min (Table 4). In N21, nestling provisioning occurred every 30 min on day 5 since hatching; totaling eight visits
in 3 h 57 min. After the brooding period –i.e., when female did not cover nestlings anymore with her body, at
c. day 9 since hatching–, nestling provisioning occurred every 20–45 min in five different broods (mean 33.8;
SD = 9.3; n = 5); in N1, N4, N5, N17a, and N19. Both nestlings were fed during every feeding session.
Table 3: Nest material weight (in g) of three Sparkling Violetear Colibri coruscans nests in Quito and Cumbayá. Seeds
belong to Taraxacum officinale (Asteraceae); fibers are vegetal; flowers belong to Callistemon citrinus (Myrtaceae).
Table 4: Nestling provisioning in seven Sparkling Violetear Colibri coruscans broods, monitored from 2018 to 2020 in
Quito; in brooding (br) and after brooding (af) period. Observ. time = total observation time per nest; Number = total
number of times nestlings were fed during observation time; Rate = average provisioning rate per hour (observation
time/total provisioning).
Brood
Observ. time
Number
Rate
N17a: br
6 h 28 min
7
55
N21: br
3 h 57 min
8
30
N17a: af
4 h 30 min
7
38
N1: af
13 h 52 min
41
20
N4: af
8 h 13 min
11
45
N5: af
11 h 45 min
20
35
N19: af
35 h 6 min 11 s
67
31
Nestling development
We analyzed photographs of nine nestlings at N1, N4, N17a, N19, and N33, from 0–19 days since hatching,
and until day 28 in N5. Nestlings hatched featherless, with orange bill, dark gray/black dorsal skin, dark pink
ventral skin, and two tracts of orange neossoptiles on the back (Fig. 4). On day 5 since hatching, feather shafts
on the dorsal side started to emerge, and 1–2 days later, feathers appeared on the head and wings. Ventral
feathers lagged behind throughout development (the ventral apterium). From day 8 since hatching onwards, the
bill became blacker, and feather growth on the dorsal side was in full development. Nestlings in four different
broods opened their eyes at days 8, 9, 11 and 12. On days 14–15 since hatching, feathers on the dorsal side were
green with a golden edge, and showed almost fully broken shafts. On day 16 since hatching the tail was still
very short. On day 19, nestlings showed the diagnosable blue-purple malar stripe, as well as the blue-purple
patch on the belly. However, the ventral apterium had not been covered yet, and feather sheaths in underwings
and tail coverts had not disappeared. After fledging, wings and tail continued to grow and juveniles were
recognizable even after 6 weeks by a golden edge on body feathers, a faint white malar stripe, and a white
postocular spot. Measures of six nestlings of different ages are shown in Table 5.
Total nest
Seeds
Moss
Fibers
Flowers
Feathers
Mean ± SD,
n
2.9 ± 0.38,
n = 3
0.7 ± 0.35,
n = 3
0.7 ± 0.52,
n = 3
0.4 ± 0.24,
n = 3
1.2 ± 1.03,
n = 2
0.05 ± 0.06,
n = 2
95%
confidence
interval
2.6–3.4
0.4–0.9
0.4–1.1
0.1–0.4
0.5–0.9

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Observation of 10 nestlings in six broods from hatching to fledging showed that on day 1 since hatching,
nestlings started opening their bills after detecting any movement in the nest. From day 7 to 14 since hatching,
nestlings usually rested, responding only to their mothers’ presence by active begging (Fig. 4). From day 14 to
21 since hatching, nestlings became more active repeatedly preening, stretching and flapping their wings, and
pecking at the nest. They also started to produce high-pitched sounds in the presence of their mother. Nestlings
stayed for 20–31 days in the nest (mean 24.6 days; SD = 3.73; 95% confidence interval = 22.5–26.7; n = 12).
We observed fledging events in seven broods. In four broods, one nestling left the nest two days after its sibling
(N17a, N17b, N21, N40). In two broods, there was a 1-day difference in fledging (N41a, N41b) and in another
brood, the second nestling left the nest 5 days later (N17e). Juveniles started feeding by themselves 14–18 days
after fledging in N5 and N19, but in one case they were fed by the female up to 24 days after nest abandonment
(N5). In another brood (N17b), the female started incubating again one week after fledging of the previous
brood. Some days later, these fledglings approached the nest again but the mother did not interact with them.
After fledging, two juveniles of two different nests (N36, N37) stayed for some time in the nest territory –in a
suburban habitat– feeding and even singing near their nest and within a male’s courtship territory. No negative
interactions were seen between males and juveniles. However, juveniles and resident females extended their ear
tufts when they approached each other closely while foraging. Fledged juveniles in N37 frequently perched on
Leonotis sp. stems while sipping nectar, in contrast to adults, which usually fed by hovering.
Breeding success
Hatching success was 81%; 55 out of 68 eggs hatched. The two eggs of N33 were preyed upon; N24 and N26,
containing both two eggs, were abandoned during the incubation phase, possibly due to drought and disturbance
by children, respectively. The remaining seven eggs (N5, N17a, N17b, N17f, N22, N39a) were found complete
but unhatched in the nest.
Fledging success was 77%; 48 hatched eggs that were monitored until fledging, produced 37 fledglings. Four
nestlings were preyed upon by Turdus fuscater, in N9, N19, and N30; one nestling disappeared from N4; nest
N36 was found destroyed 6 days after hatching with no trace of its two nestlings; three nestlings were found
dead inside their nest (N16, N39b) and the nest of N22a was removed by an unidentified person; it is uncertain
what happened to its nestling.
Overall breeding success was 60% (62 eggs in 37 broods followed-up from incubation to fledging, produced 37
fledglings). See Appendix 2 in Supplementary Material for details on eggs, incubation, nest periods, hatching
intervals, surviving nestlings, and fledging intervals.
Nests in natural areas
Eleven nests (N32, N33, N42–50) were located in and around Yanacocha reserve. Two nests (N32, N33), found
in January 2015, were 50 m apart from each other. N32 was located at the bottom of a hanging vertical branch
in a living fence of native vegetation (substrate species unidentified), at the embankment of a secondary road
near the entrance of the reserve. It had the following measures (cm): inner diameter 3.8, outer diameter 6.2,
outer height 7.4, and inner depth 2.9. The inner chamber included pieces of white and green lichens and the
outer surface had moss. It was under construction on 28 January 2015; hatching of two eggs occurred between
26–28 February; on 16 March two fledglings were observed. N33 was under a roof, attached to a light cord
connected to an outdoor restroom at the entrance of the reserve. It was found on 28 January 2015 under
construction; on 10 February it contained two eggs, and on 8 March broken eggshells were observed, possibly
caused by depredation.
The remaining nine nests (N42–N50), found on 13 May 2020, were in vegetation bordering the road from
Yanacocha to Alambi. Three nests were attached to a stolon of Cynodon dactylon, hanging vertically from the
embankment towards the ground, whereas six nests were in native shrubs: one in an unidentified Rosaceae
species, another in an unidentified Asteraceae species, and the remaining in unidentified plants. Eight nests were
in incubation stage of two eggs, and one nest contained a single egg. All nests contained moss, and six nests
had lichens in the outer layer.

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Figure 4: Development of Sparkling Violetear Colibri coruscans nestlings in Quito from hatching day through day 28. Day
16 and 19 photos from brood N1; remaining photos from brood N5 (all photos by Maartje Musschenga, excepting days 0,
5, 7, 23 and 28 by Oswaldo Ponce).
DISCUSSION
Our observations of 45 broods in 39 urban and suburban nests, including some reused nests, and 11 nests in
natural woodlands in Quito and adjacent valleys, represent the first extensive effort to document the breeding
biology of C. coruscans in Ecuador. Generally, data fit into previous descriptions of the species’ breeding
biology (Ruschi, 1965; Hainsworth, 1977; Schmidt-Marloh & Schuchmann, 1980; Zerda-Ordóñez, 1994;
Stoppelmoor, 2000; de la Peña, 2001; Gonzáles & Castañeda, 2020). Additionally, we present data about nests
adapted to urban habitats, the use of artificial structures as nest substrates, and nesting under roofs, which
provide protection to nests and favor consecutive use of the same nest site.

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Copulation, as observed in this study, was different from that described by Zerda-Ordóñez (1994), because we
observed the female approaching the perching and vocalizing male, contra this author’s observation. Further,
Zerda-Ordóñez did not describe the male’s wing vibration and upheld position of tail that we observed.
We distinguished males and females by behavior and subtle plumage differences (Zerda-Ordóñez, 1994; Freile
& Restall, 2018; Clark et al., 2018). Even though sex identification was not always straightforward due to rapid
movements of the hummingbirds or inadequate light conditions, we never observed two individuals
simultaneously at any single nest, suggesting that females alone attend their nests, as occurs in most
hummingbird species (Ortiz-Crespo, 2011). Nevertheless, our observations suggest that in some cases –15
observed broods in suburban habitats; for instance, in areas with high density of nectar rich flowers like Leonotis
sp.– females’ nesting and/or feeding territories overlap with males’ courtship and/or feeding territories (see
Ruschi, 1965; Zerda-Ordóñez, 1994; Gonzáles & Castañeda, 2020). We observed males being tolerated by the
breeding females and vice versa, whereas other individuals of C. coruscans were chased away, as also observed
by Ruschi (1965) in captive C. coruscans. Wolf & Stiles (1970) suggested that it is a reproductive advantage
for a male ‘helping’ the female he has mated with, by giving her access to a secure nectar source and tolerating
her in his territory.
We observed three instances of joint defense of the nesting area by male and female. Given that not all broods
received extensive monitoring, we cannot determine the frequency of this behavior. Whether nest defense by
males is intentional or is a by-product of males’ defense of their own feeding and/or courtship territories needs
further investigation. Besides these observations of males overlapping territories with nesting females, we did
not observe males around or nearby other 30 nests, consistent with previous observations of nesting C.
coruscans (Hainsworth, 1977; Schmidt-Marloh & Schuchmann, 1980). Further, we did not find evidence of
males incubating, as was suggested by Moore (1947) and Schäfer (1954). The conditions under which females
nest and forage in males’ territories should be studied further.
The peak of C. coruscans’ breeding season in Quito and adjacent suburban valleys is November–April; we
found lower nesting activity in May–October. However, as 10 nests were found in May in Yanacocha (northwest
of Quito), the breeding season in the Quito area might extend into May or vary locally. The peak of breeding
season in our study area covers the second half of the short rainy season (October–November), the short dry
season (December–January), and the majority of the long rainy season February–June (Pourrut et al., 1995;
Ortiz-Crespo, 2011). Study effort was not equal throughout the year, as we visited nests reported by people in
social media or nests we incidentally discovered. Yet, the breeding seasonality we suggest for C. coruscans has
already been described for this species by Ortiz-Crespo (2011) and broadly overlaps with breeding season of L.
victoriae in Quito (September–April; Narváez-Izurieta, 2022). Zerda-Ordóñez (1994) found nests of C.
coruscans in April, May and October in Bogotá, which corresponds to the first rainy season (late March–mid
June; López-Jiménez, 2012) and the second rainy season (late September–early December). Given that C.
coruscans forages on a wide range of flowers (Züchner et al., 2020), nectar availability probably has little
influence on breeding activity as in other more specialist species (Stiles, 1985). There might be a relation with
insect abundance, necessary for the nestlings' growth, but this relation needs further investigation. Quito is
experiencing increasing temperatures (Cáceres et al., 1998), and projections for 2023 predict more extreme
minimum and maximum temperatures, heavier rainfall, and more droughts in some places (Serrano et al., 2017).
This climatic change might influence C. coruscans breeding season and reproductive success in the future.
All nests were cup shaped, as is typical for Trochilidae (Ortiz-Crespo, 2011), and were attached at their base or
sides to the substrate (branches or artificial structures). The majority of nests we observed had green moss on
the outer layer, as observed previously in this species and other hummingbirds (Ruschi, 1965; Schmidt-Marloh
& Schuchmann, 1980; Zerda-Ordóñez, 1994; Stoppelmoor, 2000; de la Peña, 2001; Fierro-Calderón & Martin,
2007; Gonzáles & Castañeda, 2020). One nest contained spider web as part of the nest binding material and in
two nests spider web was used to adhere the nest to supporting branches (see Ruschi, 1965; Zerda-Ordóñez,
1994; Stoppelmoor, 2000; de la Peña, 2001; Fierro-Calderón & Martin, 2007; Ortiz-Crespo, 2011; Gonzáles &
Castañeda, 2020). It seems plausible that we overlooked this material in other nests. The inner chamber of our
nests was always covered with soft, whitish material, as noted in earlier studies of the species (Schmidt-Marloh
& Schuchmann, 1980; Zerda-Ordóñez, 1994; Ortiz-Crespo, 2011; Gonzáles & Castañeda, 2020), likely to
protect and insulate the eggs and nestlings (Winkler, 2016).

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Table 5: Measures (cm) of six Sparkling Violetear Colibri coruscans nestlings in five different broods in Quito and
adjacent valleys, between day 4 and 20 since hatching. In day column, a and b indicate different nestlings in the same
brood. In the remaining broods only one nestling was alive during the study period.
Brood
Day
Body
Bill
Left wing
Right wing
Tail
N1
13a
1
3.1
3.4
1
17a
1.3
4.6
4.5
2
19a
1.4
5.1
5
2.4
20a
1.4
5.3
5.3
13b
1.1
3.2
3.1
1
17b
1.3
4.3
4.2
2
19b
1.3
4.5
4.6
2
20b
1.3
4.9
5.0
2.1
N4
4
3.5
8
4.8
1.9
1.9
14
3.6
3.5
1.3
N5
6
3.5
0.5
13
7.4
0.7
2.8
2.8
0.7
20
4
4.1
2.9
N17a
3
3
10
4.5
0.9
2.1
1.9
20
8
1.5
5.1
5.3
3.7
N17b
11
0.8
2.5
2.5
0.5
15
1.3
4.6
4.6
1.3
We also studied nests in montane woodlands around Yanacocha reserve, where lichens were present in 6 out of
10 nests. Lichens were also reported by other authors in nests of C. coruscans (de la Peña, 2001; Gonzáles &
Castañeda, 2020) and other hummingbirds in natural habitats (Fierro-Calderón & Martin, 2007; Ortiz-Crespo,
2011). The absence of lichens in suburban and urban nests in our study area may reflect a low abundance of
lichens in cities, due to drier climates (Brodo, 1966), a lack of wooded areas, or air pollution (Coffey & Fahrig,
2012). The whitish winged seeds of Tecoma stans we observed in the outer layer of two nests in a suburban
area could be an alternative to the camouflage function of lichens (Hansel, 1996), as C. coruscans has also been
reported using other whitish material on the outer layer, such as pieces of newspaper (Ortiz-Crespo, 2011).
Colibri coruscans nests in a variety of native and exotic plants and shrubs, but also takes advantage of human-
made structures; even in natural areas such as Yanacocha reserve, we observed a nest located on a light cord
under a roof. Several hummingbird species nest occasionally on bridges, under roofs, and in empty (Freeman
& Arango, 2012) or even occupied buildings (Ward et al., 2020), which protect nests from rain and predators
(Triana & Sandoval, 2011; Freeman & Arango 2012; Ward et al., 2020). Nesting in or under artificial ‘refuges’
that prevent nest deterioration might benefit breeding hummingbirds because a single nest can be used several
consecutive times in a same breeding season –up to 11 times in subsequent seasons as observed in our study–,
often by adding new materials to the nest (see Ortiz-Crespo, 2011; Ward et al., 2020). We also observed nests
attached to a vertical C. dactylon stolon hanging from a road embankment. In this case, nests were not reused,
but rebuilt from zero at least 10 times between 2009 and 2015.
Thirty-six out of 41 clutches studied consisted of two eggs, which is common in hummingbirds (Ortiz-Crespo,
2011). In two broods, the second egg was laid 24 h after the first, and in one brood between 24 and 48 h
afterwards. Incubation in three nests started when the second egg was laid. Ruschi (1965) observed a 2-day
interval between egg laying in C. coruscans and incubation started one day after the second egg was laid. Other
hummingbird species typically lay the second egg within 24–48 h after the first egg (Ortiz-Crespo, 2011; Fierro-
Calderón & Martin 2007; Ornelas, 2010), and incubation started after laying of the second egg in Azure-
crowned Hummingbird Saucerottia cyanocephala (Ornelas, 2010). More data are needed to confirm timing of
egg laying and start of incubation in C. coruscans. In our study, hatching in two broods occurred on the same
day and in two other broods occurred on consecutive days. Zerda-Ordóñez (1994) observed a typical 2-day
difference in hatching with exceptions of hatching on the same day, whereas Ruschi (1965) recorded a hatching
interval of 0–1 day.

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We found an incubation constancy of 75%. Skutch (1962) mentions at least 60% for hummingbirds and Zerda-
Ordóñez (1994) recorded a constancy of 60–72% on different days, increasing towards the end of incubation,
in C. coruscans. Brooding constancy, on the contrary, was 48–51% at the end of the first week (day 5–6) in two
nests studied, which is well within the range of tropical hummingbirds (Fierro-Calderón & Martin, 2007). Zerda-
Ordóñez (1994) observed a constancy of 39% on day 5 since hatching in the morning, 22% on day 5 since
hatching in the afternoon, and 11% on day 6 since hatching. The nestlings’ provisioning rate we found was
comparable to other hummingbird species (Fierro-Calderón & Martin, 2007).
The physical and behavioral development of nestlings in our study agree with observations of Zerda-Ordóñez
(1994) in Bogotá and Ruschi (1965) in captive C. coruscans in Brazil. In another species observed in Quito (L.
victoriae), down feathers were also present at hatching but later covered the whole body, contour feathers and
rectrices developed some days earlier than in C. coruscans, but no lag in the development of the ventral
pteryllium was mentioned, as we found in C. coruscans (see also Ruschi, 1965; Zerda-Ordóñez, 1994). Lesbia
victoriae nestlings opened their eyes on day 17, while in C. coruscans they opened eyes between days 8–12.
The nesting period of 20–31 days found in our study was highly variable compared to previous studies about
the same species: 21–25 days (Ruschi, 1965; Zerda-Ordóñez, 1994; Schmidt-Marloh & Schuchmann, 1980,
Stoppelmoor, 2000). However, it lies within the range reported for other hummingbird species in Quito (25–28
days in Western Emerald Chlorostilbon melanorhynchus, Ortiz-Crespo 2011; 25–27 days in L. victoriae,
Narváez-Izurieta, 2022). Similarly, the closely related Mexican Violetear Colibri thalassinus had a nesting
period of 19–28 days in Mexico (Wagner, 1945), while a nesting period of 19–40 days is reported for
hummingbirds in general (Ortiz-Crespo, 2011). The length of the nesting period might be related to food
availability and weather conditions (Ortiz-Crespo, 2011). We also observed that one juvenile started looking
for food on its own almost two weeks after fledging, but was fed by its mother for two more weeks (see also
Ruschi, 1965; Hainsworth, 1977; Stoppelmoor, 2001). Hainsworth (1977) concluded that by provisioning
fledglings with food, independence is delayed and juveniles can practice food search by themselves before
abandoning the nesting area. On the contrary, failure to provide the necessary energy by the mother may force
early fledglings’ independence. In one nest we studied, the female laid a new brood one week after fledging of
the previous brood, which forced the two fledglings to abandon the nesting area earlier.
We conclude that C. coruscans in Quito and surroundings is versatile in its choice of nesting sites and nest
materials. In urban and suburban habitats, the use of human-made structures aids in preservation of the nests;
therefore, the same nest can be reused several times during the same or over consecutive breeding seasons. We
also found that females’ nest and males’ feeding and/or courtship territories might overlap. In this case, males
are indirectly involved in the defense of the nest area, but are not involved in other breeding duties. In future
studies, territories of males and females should be mapped in order to determine how and under which
circumstances they overlap. Also, in further investigations sampling effort should be equivalent over the whole
year, in order to define the breeding season more precisely, as well as to monitor more nests from egg laying to
fledging.
ACKNOWLEDGEMENTS
We would like to thank Oswaldo Ponce, Jarvin Grain, Adela Espinosa, Elena Raza, Luis Sabino, Fernanda
Salazar, Juan Manuel Carrión, and Henry López for sharing data and/or allowing us to monitor the nests at their
homes and workplaces. Oswaldo Ponce also provided daily pictures of nestlings’ development. Thanks to Joel
Gordón for his help in analyzing videos and to José Luis Paucar for the graphics. An earlier version of this
manuscript received useful suggestions from Fernanda Duque and two anonymous reviewers. Michael Seager
and Julie Watson helped improve English grammar and sentence structure.
REFERENCES
Brodo, I.M. (1966). Lichen growth and cities: A study on Long Island, New York. Bryology, 69(4), 427–449.
DOI: https://doi.org/10.2307/3240577
Cáceres, L., Mejía, R., & Ontaneda, G. (1998). Evidencias del cambio climático en Ecuador. Bulletin de l'Institut
Français d' Études Andines, 27(30), 547–556. URL: https://www.redalyc.org/pdf/126/12627319.pdf

Revista Ecuatoriana de Ornitología, 8, 51–66
64
Breeding of Colibri coruscans in Quito
Musschenga, M.A. et al. (2022)
Cadena-Ortiz, H. (2018). Sugerencias para la toma de datos en eventos de biología reproductiva de aves.
Avances en Ciencias e Ingenierías, 10(1), 25–35. DOI: https://doi.org/10.18272/aci.v10i1.297
Cisneros-Heredia, D., Amigo, X., Arias, D., Arteaga, J., Bedoya, J., Espinoza, S., Montenegro, E., Nazati, G.,
& Carrión, J. M. (2015). Reporte del 1er conteo navideño de aves de Quito, Ecuador. Avances en Ciencias e
Ingenierías, 7(2), B37–B51. DOI: https://doi.org/10.18272/aci.v7i2.256
Clark, C., McGuire, J., Bonaccorso, E., Berv, J., & Prum, R. (2018). Complex coevolution of wing, tail, and
vocal sounds of courting male bee hummingbirds. Evolution, 72(3), 630–646. DOI:
https://doi:10.1111/evo.13432
Coffey, H., & Fahrig L. (2012). Relative effects of vehicle pollution, moisture and colonization sources on urban
lichens. Journal of Applied Ecology, 49, 1467–1474. DOI: https://doi.org/10.1111/j.1365-2664.2012.02208.x
de la Peña, M. (2001). Nidificación de algunas especies de aves en el este de la provincia de Catamarca,
Argentina. El Hornero, 16(1), 17–21. URL: https://www.avesargentinas.org.ar/el-hornero-volumen-16-
n%C3%BAmero-1
Fierro-Calderón, K., & Martin, T. (2007). Reproductive biology of the Violet-chested Hummingbird in
Venezuela and comparisons with other tropical and temperate hummingbirds. The Condor, 109, 680–685. DOI:
https://doi.org/10.1093/condor/109.3.680
Freeman, B., & Arango J. (2012). Notes on the nesting biology of the Empress Brilliant (Heliodoxa imperatrix)
in western Colombia. Boletín de la Sociedad Antioqueña de Ornitología, 21(2), 67–71. URL:
http://sao.org.co/publicaciones/boletinsao/20%282%29/NC520%282%29Freeman&Arango.pdf
Freile, J., & Restall, R. (2018). Birds of Ecuador. London, U.K.: Helm Field Guides.
Hainsworth, F. (1977). Foraging efficiency and parental care in Colibri coruscans. The Condor, 79, 69–75.
URL: https://www.jstor.org/stable/1367532?seq=1
Hansel, M. (1996). The function of lichen flakes and white spider cocoons on the outer surface of birds’ nests.
Journal of Natural History, 30, 303–311. DOI: https://doi.org/10.1080/00222939600771181
Gonzáles, P., & Castañeda, E. (2020). Aspectos sobre la biología reproductiva del colibrí Oreja-Violeta de
Vientre Azul (Colibri coruscans) en el departamento de Lima, con notas sobre su dieta. Boletín de la Unión de
Ornitólogos del Perú (UNOP), 15(2), 30–39. URL:
https://boletinunop.weebly.com/uploads/6/2/2/6/62265985/boletin_unop_vol._15_n%C2%B02_2020_-
_gonzales.pdf
López-Jiménez, V.L. (2012). Condiciones atmosféricas predominantes en el piedemonte de la cordillera
oriental, determinantes del régimen climático en las sedes de la universidad de la Salle, Bogotá. Épsilon, 1(18),
87–104. URL: https://ciencia.lasalle.edu.co/cgi/viewcontent.cgi?article=1140&context=ep
Moore, R. (1947). Habits of male hummingbirds near their nests. The Wilson Bulletin, 59(1), 21–25. URL:
https://sora.unm.edu/sites/default/files/journals/wilson/v059n01/p0021-p0025.pdf
Murray, B. (2000). Measuring annual reproductive success in birds. The Condor, 102(2), 470–473. DOI:
https://doi.org/10.2307/1369665
Museo Ecuatoriano de Ciencias Naturales. (2009). Ecosistemas del Distrito Metropolitano de Quito (DMQ).
Quito, Ecuador: Publicación Miscelánea 6. Serie de Publicaciones del Museo Ecuatoriano de Ciencias Naturales
- Fondo Ambiental del MDMQ.

Revista Ecuatoriana de Ornitología, 8, 51–66
65
Breeding of Colibri coruscans in Quito
Musschenga, M.A. et al. (2022)
Narváez-Izurieta, R. (2022). Notas reproductivas del Colacintillo Colinegro Lesbia victoriae (Apodiformes:
Trochilidae) en Quito, Ecuador. Revista Ecuatoriana de Ornitología, 8(1), 24–30. DOI:
https://doi.org/10.18272/reo.v8i1.1808
Ornelas, J.F. (2010). Nests, eggs, and young of the Azure-crowned Hummingbird (Amazilia cyanocephala).
The Wilson Journal of Ornithology, 122(3), 592–597. DOI: http://dx.doi.org/10.1676/09-155.1
Ortiz-Crespo, F. (2011). Los colibríes: historia natural de unas aves casi sobrenaturales. Quito, Ecuador:
Imprenta Mariscal.
Pourrut, P., Róvere, O., Romo, I., & Villacrés, H. (1995). Clima del Ecuador. In P. Pourrut (Ed.), El agua en el
Ecuador. Clima, precipitaciones, escorrentías (pp. 13–26). Quito, Ecuador: RR Editores Asociados. URL:
https://horizon.documentation.ird.fr/exl-doc/pleins_textes/pleins_textes_7/divers2/010014823.pdf
Schmidt-Marloh, D., & Schuchmann, K. (1980). Zur Biologie des Blauen Veilchenohr Kolibris (Colibri
coruscans). Bonner Zoologische Beiträge, 31(1–2), 61–77. URL: https://zoologicalbulletin.de/articles/bzb-
1980-1989/206-volume-31-12-august-1980
Ruschi, A. (1965). Observações sobre a nidificação, incubação e cuidados com a prole em Colibri coruscans
coruscans (Gould) realizado unicamente pela fêmea. Boletim Museu de Biología Profesor Mello Leitao, 45, 1–
9. URL: http://boletim.sambio.org.br/pdf/bi_45.pdf
Schäfer, E. (1954). Sobre la biología de Colibri coruscans. Separata del Boletín de la Sociedad Venezolana de
Ciencias Naturales, 82, 153–162.
Serrano, S., Ruiz, J., & Bersosa, F. (2017). Heavy rainfall and temperature projections in a climate change
scenario over Quito, Ecuador. LA GRANJA: Revista de Ciencias de la Vida, 25(1), 16–32. DOI:
https://doi.org/10.17163/lgr.n25.2017.02
Skutch, A. (1962). The constancy of incubation. The Wilson Bulletin, 74(2), 115–152. URL:
https://sora.unm.edu/sites/default/files/journals/wilson/v074n02/p0115-p0152.pdf
Stiles, F. G. (1985). Seasonal patterns and coevolution in the hummingbird-flower community of a Costa Rican
subtropical forest. Ornithological Monographs, 36, 757–787. DOI: https://doi.org/10.2307/40168315
Stoppelmoor, G. (2000). Captive breeding of the Sparkling Violet-ear Hummingbird. The American Federation
of Aviculture Magazine Watchbird, 27(1), 52–54. URL: https://watchbird-ojs-
tamu.tdl.org/watchbird/index.php/watchbird/article/view/1498
Triana, E., & Sandoval, L. (2011). Nest reuse by the Scintillant Hummingbird (Selasphorus scintilla). The
Wilson Journal of Ornithology, 123(3), 635–638. DOI: https://doi.org/10.1676/10-181.1
Van den Burg, A. (2017). Het onderzoeken van niet uitgekomen vogeleieren. Otterlo, The Netherlands:
Biosphere Science Productions.
Wagner, H. (1945). Notes on the life history of the Mexican Violet-ear. The Wilson Bulletin, 57(3), 165–187.
URL: https://www.jstor.org/stable/4157430
Ward, J., Meek, N., Johnson, T., Johnson, K., Batchelder, N., & Marks, J. (2020). Ace is the place: Black-
chinned Hummingbirds (Archilochus alexandri) have high nesting success and productivity inside a hardware
store. The Wilson Journal of Ornithology, 132(4), 1053–1058. DOI: https://doi.org/10.1676/1559-4491-
132.4.1053
Winkler, D. (2016). Breeding biology of birds. In: I. Lovette & J. Fitzpatrick (Eds.), Cornell Lab of
Ornithology’s handbook of bird biology (pp. 407–452). Chichester, U.K.: John Wiley & Sons.

Revista Ecuatoriana de Ornitología, 8, 51–66
66
Breeding of Colibri coruscans in Quito
Musschenga, M.A. et al. (2022)
Wolf, L., & Stiles G. (1970). Evolution of pair cooperation in a tropical hummingbird. Evolution, 24, 759–773.
URL: https://onlinelibrary.wiley.com/doi/pdfdirect/10.1111/j.1558-5646.1970.tb01811.x
Zerda-Ordóñez, E. (1994). Historia natural del Tominejo (Colibri coruscans coruscans) (Gould) (Aves,
Trochilidae). Universitas Scientiarium, 2(1), 65–85.
URL: https://repository.javeriana.edu.co/handle/10554/31277
Züchner, T., Boesman, P.F.D., & Kirwan, G.M. (2020). Sparkling Violetear (Colibri coruscans), version 1.0.
In: J. del Hoyo, A. Elliott, J. Sargatal, D.A. Christie & E. de Juana (Eds), Birds of the World. Ithaca, NY: Cornell
Lab of Ornithology. DOI: https://doi.org/10.2173/bow.spvear1.01