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1. Genus Coeligena Lesson, 1832

1.12 Coeligena inca (Gould, 1852)

1.12 Coeligena inca (Gould, 1852)

taxon remains unclear, one example being the taxonomic list presented by Peterson (1996), where inca is omitted, presumably included in torquata.

For the purpose of this study, Coeligena torquata (Boissonneau, 1840) and Coeligena inca (Gould, 1844) were considered as separate entities (sensu Schuchmann 1999), but the close morphologic relationship is obvious, and it is very probable that C. inca and the species complex C. torquata represent a superspecies. This topic will be discussed in the section on phylogenetic relationships.

1.12.2 Diagnosis

Coeligena inca is a relatively large-sized, dimorphic species that shows a very high resemblance to Coeligena torquata in many plumage features. The males have long, straight, black bills. The head is blackish, fringed gold and green; crown with only one very yellowish Sky Blue 173 frontal spot. The upperparts are golden Parrot Green 160, becoming more shiny greenish on the lower back; upper tail-coverts yellowish Parrot Green 260. The throat is black with washed golden Spectrum Green 62 discs, that become more shiny on the anterior edge of the pectoral collar; pectoral collar feathers white at the base and Amber 36 on the tips.

Underparts glittering golden Parrot Green 160 with bluish fringes; under tail-coverts Parrot Green 160 tipped brownish white. The tail is forked with inner rectrices golden Parrot Green 160 and the others white with contrasting golden Parrot Green 160 tips varying in size.

Females have longer bills than males; head blackish Parrot Green 260 forming small discs;

greenish blue crown spot present; rest of the upperparts shining golden green. The chin and throat are Amber 36 with sides bluish Spectrum Green 62; pectoral collar like throat, without white. The underparts washed Cinnamon 39, sides with discs golden Spectrum Green 62. The tail is less deeply forked. Immatures are similar to adult females with head feathers fringed buffy (Buff 124) but the crown spot is absent.

1.12.3 Geographic variation analysis

A total of 39 skins of Coeligena inca (23 males and 16 females) was examined. The localities were grouped in four geographic pools (Fig. 2.12.1). There follows a list of these pools and the descriptions of the corresponding areas they occupy, ordered from north to south.

Figure 2.12.1. Geographic distribution of C. inca. The polygons represent the geographic pools, the dots the localities.

1. Urubamba: located north-east of Cuzco, on the east bank of the River Apurímac, southern Peru, between 1650 and 3300 m a.s.l.

2. Puno: located north-west of Lake Titicacaca, in southern Peru, near the border with Bolivia, between 1950 and 3100 m.

3. La Paz: located south of Lake Titicaca, in the Cordillera Real range, northern Bolivia, between 2400 and 2750 m.

4. Cochabamba: located in in the Bolivian Yungas, Cordillera Cochabamba range, northern Bolivia, between 2400 and 2750 m.

Table 2.12.1 Descriptive statistics of the morphometric parameters in C. inca. The pools are ordered geographically from north to south.

Morphometric

parameters Pools Urubamba Puno La Paz Cochabamba

bill-length

males mean

±S.D.

Max.

Min.

N

39.2 0.84 40.1 38.3 4

37.1 0.38 37.4 36.7 3

38.6 0.95 39.8 37.1 8

38.0 1.29 39.9 35.6 8 (mm) females

mean

±S.D.

Max.

Min.

N

39.8 1.48 40.8 38.7 2

40.0 1.52 41.4 37.5 6

39.2 1.56 40.3 38.1 2

40.1 0.64 40.8 39.7 3

wing-length

males mean

±S.D.

Max.

Min.

N

76.0 1.76 78.5 74.6 5

75.4 1.48 77.5 74.0 6

76.0 2.04 79.2 73.5 9

75.2 3.04 79.8 68.3 10 (mm) females

mean

±S.D.

Max.

Min.

N

69.6 1.06 70.3 68.8 2

70.4 1.44 73.5 69.3 7

70.2 0.56 70.6 69.8 2

70.9 0.99 72.1 70.1 4 1.12.3.1 Morphometric variation:

Coeligena inca has a marked morphometric dimorphism (Tab. 2.12.1, Fig. 2.12.3-4). Males have shorter bills, longer wings and longer, more deeply forked tails than the females (p<0.01, bill-length: males 38.2±1.14 mm, females 39.8±1.23 mm; wing-length: males 75.6±2.24 mm, females 70.4±1.18 mm; tail-length: males 46.2±2.65 mm, females 41.6±0.83 mm; tail fork-depth: males 4.6±1.92, females 1.4±1.17). None of the measured variables showed significant differences between pools.

Table 2.12.1. (Continued) Descriptive statistics of the morphometric parameters in C. inca. The pools are ordered geographically from north to south.

Morphometric

parameters Pools Urubamba Puno La Paz Cochabamba

tail-length

males mean

±S.D.

Max.

Min.

N

48.0 2.14 49.8 44.8 5

45.8 3.67 49.8 39.9 6

46.2 1.62 48.5 43.9 9

45.7 2.92 49.9 39.1 10 (mm) females

mean

±S.D.

Max.

Min.

N

40.9 0.98 41.6 40.2 2

42.1 0.62 42.9 41.0 7

41.4 1.27 42.3 40.5 2

41.4 0.80 42.6 40.7 4

tail fork-depth

males mean

±S.D.

Max.

Min.

N

5.6 2.04 8.4 3.5 4

4.2 1.71 5.8 1.6 6

5.2 0.65 6.4 4.4 8

4.0 2.53 8.6 -0.7 10 (mm) females

mean

±S.D.

Max.

Min.

N

0.2 0.07 0.2 0.1 2

2.0 1.38 4.1 0.4 6

0.9 0.98 1.6 0.2 2

1.6 0.32 1.8 1.2 3

1.12.3.2 Plumage variation

Coeligena inca has a homogeneous coloration pattern througout its range (see description in Diagnosis). The pools Urubamba (1) and Puno (2) did not show any detectable difference in plumage coloration. The males of both pools show a very blackish head fringed Dark Green 162A, with one frontal glittering greenish Turquoise Blue 65 crown spot. The nape is blackish Dark Green 162; upperparts golden Parrot Green 160 fringed turquoise. The innermost tail feather is yellowish Olive Green 50; the other rectrices white, with olive restricted to the tips.

The chin and throat are blackish-bronzy Parrot Green 160; collar Amber 36 with white;

are similar to the males. The head is dark Parrot Green 160 with one crown spot similar to the males; upperparts yellowish Parrot Green 260; upper tail-coverts yellowish Olive Green 50.

Tail similar to the males’. The chin and throat are scaly yellowish Parrot Green 160 laterally, and medially Amber 36; the pectoral collar has the same colour as the throat. The underparts are scaly blackish-bluish Parrot Green 260; under tail-coverts bluish Parrot Green 160.

The pools La Paz (3) and Cochabamba (4) showed very few differences from the two pools described above. The more evident variation was found in male crown, being more bluish, Cobalt Blue 68-Sky Blue 66, in these pools than in the Peruvian pools. The chin and throat have no bronze, being more golden-blackish Parrot Green 260. The pectoral collar and throat are separated by a glittering golden Spectrum Green 62 band; underparts and under tail-coverts more bluish Parrot Green 160. Females of both Peruvian and Bolivian pools are practically identical, presenting only minor variations, like less bluish on the underparts and more yellowish on the upperparts of the Bolivian pools.

Figure 2.12.2. Geographic distribution of C. inca. The subspecies ranges are represented by filled polygons.

1.12.4 Discussion

The Coeligena inca omissa holotype was collected in Huansampillo, south-eastern Peru, the specimens from the pools Urubamba and Puno belonging to this taxonomic group, whereas Coeligena inca inca was described from the north-east of Bolivia, corresponding with the pools La Paz and Cochabamba (Fig. 2.12.2). Zimmer’s (1948) description of C. i. omissa states that the most important differences found between the two races consisted in the amount of green on the throat; the more greenish tinges on the head; the amount of bluish hue in the frontal crown spot; the much slighter rufous hue on the breast, and the darker green on the back and underparts; the females are not distinguishable, but in the light of morphometric and plumage characteristics the validity of the race C. i. omissa is questionable.

Geographically it is not supported either, because the valley of the River Madre de Dios could hardly act as an effective barrier to the dispersal of these birds.

Unfortunately, there are not many specimens of this species in order to determine if the gap between the northern and the southern populations is not caused by poor sampling in the region. For the moment I suggest maintaining the name C. i. omissa for the Peruvian subpopulations until more evidence to the contrary is found.

A males B females

Urubamba-M Urubamba-F Puno-M Puno-F La Paz-M La Paz-F Cochabamba-M Cochabamba-F

pools

36.0 38.0 40.0 42.0

Bill-length (mm)

B

A B

A B

A B A

A males

B females

Urubamba-M Urubamba-F Puno-M Puno-F La Paz-M La Paz-F Cochabamba-M Cochabamba-F

pools

67.5 70.0 72.5 75.0 77.5

Wing-length (mm) B

A

B A

B A

B A

Figure 2.12.3. Geographic variation of the bill- and wing-length in C. inca. The circles and triangles represent the arithmetic mean of males and females, respectivela; and the vertical line the standard error of the mean. Pools are geographically ordered from north to south.

A males

B females

Urubamba-M Urubamba-F Puno-M Puno-F La Paz-M La Paz-F Cochabamba-M Cochabamba-F

pools

40.0 45.0 50.0

Tail-length (mm) B

A

B A

B A

B A

A males

B females

Urubamba-M Urubamba-F Puno-M Puno-F La Paz-M La Paz-F Cochabamba-M Cochabamba-F

pools

0.0 2.5 5.0 7.5

Tail Fork-depth (mm)

B A

B A

B A

B A

Figure 2.12.3. Geographic variation of the tail-length and tail fork-depth in C. inca. The circles and triangles represent the arithmetic mean of