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1. Genus Coeligena Lesson, 1832

1.4 Coeligena bonapartei (Boissonneau, 1840)

A Males B Females

Merida-M Tama-M Las Vegas-M Cundinamarca-M Antioquia-M Tolima-M Monchique-M Moscopan-M Llorente-M East Pichincha-M Tungurahua-M Morona-M Zamora-F Chimborazo-F Pichincha-F San Martin-M Utcubamba-M Carpish-M Chilpes-M Ayacucho-F Vilcabamba-F

pools

30.0 40.0 50.0 60.0

Tail-length (mm)

A

B A

B A A

B A B A

A

B A

B A A

B A

B A

B

A A

B A

B A

B A

B A

B A

B A

B A

B A

B A

A Males

B Females

Merida-M Tama-M Las Vegas-M Cundinamarca-M Antioquia-M Tolima-M Monchique-M Moscopan-M Llorente-M East Pichincha-M Tungurahua-M Morona-M Zamora-F Chimborazo-F Pichincha-F San Martin-M Utcubamba-M Carpish-M Chilpes-M Ayacucho-F Vilcabamba-F

pools

0.0 5.0 10.0

Tail Fork-depth (mm)

A

B A

B A

A B

A A

B A

B A

B A A

B A B

A

B A

A

B A

B A

B A

B A

B A

B

A B

A

B A

B A

Figure 2.3.4. Geographic variation of the tail-length and tail fork-depth in the superspecies complex C. torquata. The circles and triangles represent the arithmetic mean in males and females, respectively; the vertical solid lines the standard error of the mean; the dashed lines represent the pool groups for each allospecies, from left to right: C. conradii, C. torquata , and C. insectivora.

1.4.1 Geographic distribution and taxonomy

Coeligena bonapartei occupies a relatively small geographic range, occurring in a patchy pattern on the Sierra de Perija, along the Colombian-Venezuelan border, in the East Andes of central Colombia, and with only one record from the Paramo de Frontino, in the northern Central Andes of Colombia. Three subspecies have been recognised: C. b. consita, in the Sierra de Perija, along the border of Venezuela and Colombia; C. b. bonapartei, on the western slope of the East Andes of Colombia, between Boyacá and Bogotá; C. b. orina, only known from the Páramo of Frontino, in Antioquia, northern central Colombia (Schuchmann 1999).

The species was first described by Boissoneau (1840) as Ornismia Bonapartei, its origin being assigned to Santa Fe de Bogotá and later placed in the genus Helianthea Gould, 1848 by Bonaparte (1850), there were no changes in the species’ taxonomy until Peters’ (1945) check-list, which included it in the genus Coeligena, Lesson 1832.

Wetmore & Phelps (1952) described the race Coeligena bonapartei consita, from the Perija Mountains of Venezuela and Colombia, and later Wetmore (1953) described the species Coeligena orina, from the Paramo Frontino, Antioquia, in the northwestern Colombian Andes, only known from the holotype (deposited in the National Museum of Natural History, Washington, D.C.). The taxon was considered by Bleiweiss (1988b) to be a subspecies of Coeligena bonapartei (Boissonneau, 1840), bringing to three the number of currently recognised subspecies: C. b.consita, C. b. bonapartei and C. b. orina (Fjeldså & Krabbe 1990, Schuchmann 1999).

1.4.2 Diagnosis

Coeligena bonapartei is sexually dimorphic. Males have a long black bill; head black with a slightly bluish green iridescent frontlet; upperparts shining dark Parrot Green 160, more golden Spectrum Orange 17 on lower back; upper tail-coverts metallic dark golden bronze.

The throat is glittering dark Emerald Green 163 with a bluish iridescent Spectrum Violet 72 patch; belly golden; under tail-coverts golden Olive-Green 47 fringed rufous; tail forked, dark golden Olive-Green 47. The females are similar to males, with longer bills; head Parrot Green 160; chin and throat rufous Amber 36 with some metallic green discs; underparts rufous Amber 36 with large green discs; belly golden; under tail-coverts bronzy olive green, largely

fringed rufous Amber 36; tail less forked, golden-yellowish Olive-Green 50, sometimes buff on tips. Immature resembles adult female.

1.4.3 Geographic variation analysis

A total of 28 specimens of Coeligena bonapartei (16 males, 10 females and 2 immatures) were analysed. Only 39% of the skins was of analytical value, due to the absence of precise information on the collecting sites or origin of the specimen (many of them are catalogued as

‘Bogotá skin trade’ or ‘Colombia’). Specimens with ‘good’ data were grouped into the following three geographic pools:

1. Perija: located on the border between Colombia and Venezuela, in the Perija Mountains, between 1800 and 3000 m a.s.l.

2. Cundinamarca: located on the western slopes of the East Andes of central Colombia, between 2500 and 3200 m.

3. Frontino: located in the Paramo Frontino, in the West Andes of northwestern Colombia, at 3890 m. Exceptional pool formed by only one specimen.

1.4.3.1 Morphometric variation

Males of Coeligena bonapartei have shorter bills, longer wings and longer and more deeply forked tails than females (Tab. 2.4.1). In males, no significant differences between the pools was found (Fig. 2.4.3-4); nevertheless, the only male from the Perija pool (wing-length: 78.6 mm; tail-length: 50.4) had wings and tail longer than the males from the Cundinamarca and Frontino pools (wing-length: 72.0-73.3 mm; tail-length: 44.3-45.3 mm). The female sample size did not allow statistical comparisons.

1.4.3.2 Plumage variation

The male from the pool Perija (1) exhibited a black head with a metallic Apple Green 61 frontal patch; nape black; upperparts and upper tail-coverts very bronzy dark Parrot Green 160. The chin is metallic Apple Green 61; throat yellowish Spectrum Green 62; gular patch very bluish True Violet 172A; underparts glittering reddish bronze; under tail-coverts Olive-Green 50 tipped Cinnamon 39. The rectrices are very bronzy Olive-Olive-Green 50; wing-coverts like back; primaries Raw Umber 223; secondaries Cinnamon 39. The females are similar to

green. The chin and throat are pale Cinnamon 39, with tiny metallic yellowish Parrot Green 260 discs; underparts more mixed with yellowish green. The rectrices are bronze Olive-Green 50, outer feather border white, the rest tipped white.

Figure 2.4.1. Geographic distribution of C. bonapartei. The polygons represent the geographic pools, dots the localities.

The males from the Cundinamarca pool (2) present an appearance similar to that described above for the Perija pool, differing in the more blackish coloration on the head; nape and upper back glittering dark Parrot Green 160 fringed golden; lower back and upper tail-coverts metallic golden olive Spectrum Orange 17. The chin and throat are bluish dark Parrot Green 160; gular patch True Violet 172A; underparts like throat; belly golden Spectrum Orange 17 mixed with some olive; under tail-coverts light Olive-Green 50 bordered Cinnamon 39. The rectrices are Olive-Green 50; Remiges Raw Umber 223. The only specimen from El Peñon, above Fusagasuga, at the northern extreme of the pool, has the inner base of the secondaries Cinnamon 39. No females were found in this pool.

The Frontino pool is formed by only one male specimen, probably an immature (presence of

males from pool 2, being much blacker on the head, upperparts and underparts; the lower back and belly more yellowish Parrot Green 160; all remiges Raw Umber 223.

Figure 2.4.2. Geographic distribution of C. bonapartei with the range of each subspecies indicated by polygons.

The specimens from Perija pool correspond with the subspecies Coeligena bonapartei consita (Wetmore and Phelps, Jr., 1952), which was described for the first time only from adult females. Only one male specimen was found in all the museums visited, which is deposited in the Colección Ornitológica Phelps, in Caracas. The taxon is distinguishable by the pale brown coloration on the secondaries, sharing this character with the species Coeligena eos, from the Venezuelan Andes, with the rest of the body coloration being similar to the nominate race (Fig. 2.4.2).

The Cundinamara pool corresponds with the nominate race, C. b. bonapartei (Boissonneau, 1840). The majority of the ‘Bogota’-labelled males show the same coloration pattern as observed for males from the Cundinamarca pool. As mentioned, there were no females that

labelled ‘Bogota’ belong to this pool. These ‘Bogota’ females are similar to females from Perija, except for the absence of the cinnamon spot on the secondaries and their tendency to be smaller in size (Fig. 2.4.3-4).

The Frontino pool (3) is represented by only one immature male that lacks the metallic green frontal spot and the pale brown spot on the secondaries. The specimen was described as Coeligena orina Wetmore, 1953 and recently Bleiweiss (1988), based on non age-related distinctive plumage features, considered the holotype as the immature male of a C. bonapartei subspecies different from C. b. bonapartei and C. b. consita. On this assumption, the name Coeligena bonapartei orina (Wetmore, 1953) has been assigned to the population occurring in the Paramo Frontino (Fjeldså & Krabbe 1990, Schuchmann 1999). Nevertheless, there are authors who prefer to consider the taxon as a valid species (e.g., Hilty & Brown 1986). In this study, the taxon is considered as a subspecies of C. bonapartei, but adult specimens of both sexes are still necessary to clarify the taxonomic and systematic status of the group.

Table 2.4.1. Descriptive statistics of the morphometric parameters of C. bonapartei. Pools are ordered geographically from north to south.

Mophometric parameters

Pools Perija Cundinamarca Unlocated Frontino

bill-length

males mean

±S.D.

Max.

Min.

N

- - - - -

35.6 1.27 36.5 34.7 2

34.5 1.36 34.5 32.5 12

35.5 - - - 1 (mm) females

mean

±S.D.

Max.

Min.

N

39.4 1.19 40.6 37.6 6

- - - - -

37.6 1.46 39.1 36.0 4

- - - - -

wing-length

males mean

±S.D.

Max.

Min.

N

78.6 - - - 1

73.3 1.56 74.4 72.2 2

70.4 4.37 70.4 62.7 12

72.0 - - - 1 (mm) females

mean

±S.D.

Max.

Min.

N

72.6 1.40 74.1 70.7 6

- - - - -

67.1 3.02 70.6 63.3 5

- - - - -

tail-length

males mean

±S.D.

Max.

Min.

N

50.4 - - - 1

44.3 0.70 44.8 43.8 2

44.8 1.86 44.8 40.1 12

45.3 - - - 1 (mm) females

mean

±S.D.

Max.

Min.

N

44.8 1.02 46.1 43.9 6

- - - - -

41.9 1.79 44.3 39.7 5

- - - - -

tail fork-depth

males mean

±S.D.

Max.

Min.

N

- - - - -

9.9 - - - 1

8.3 1.85 8.3 5.4 12

6.5 - - - 1 (mm) females

mean

±S.D.

Max.

Min.

N

7.0 1.70 8.2 5.8 2

- - - - -

4.7 2.24 7.2 1.5 5

- - - - -

A males B females

Perija-F CundinamarcaM UNLOCATED-F UNLOCATED-M Frontino

pools

34.0 36.0 38.0 40.0

bill-length (mm) A A

B

B

A

A males

B females

Perija-M Perija-F CundinamarcaM UNLOCATED-F UNLOCATED-M Frontino

pools

65.0 70.0 75.0 80.0

wing-length (mm)

A

A B

A

B A

Figure 2.4.3. Geographical variation of the bill- and wing-lengths in C. bonapartei. The circle and triangles represent the arithmetic mean of males and females, respectively; and the vertical lines the standard error of the mean. Pools are ordered from north to south. The group with the name ‘Unlocated’ is linked to the Cundinamarca pool (see text).

A males B females

Perija-M Perija-F CundinamarcaM UNLOCATED-F UNLOCATED-M Frontino

pools

40.0 42.5 45.0 47.5 50.0

tail-length (mm) A

A B

A

B A

A males

B females

Perija-F CundinamarcaM UNLOCATED-F UNLOCATED-M Frontino

pools

2.5 5.0 7.5 10.0 12.5

tail fork-depth (mm)

A

A B

B A

Figure 2.4.4. Geographical variation of the tail-length and the tail fork-depth in C. bonapartei. The circle and triangles represent the arithmetic mean of males and females, respectively; and the vertical lines the standard error of the mean. Pools are ordered from north to south. The group with the name ‘Unlocated’ is linked to Cundinamarca pool (see text).