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1. Genus Coeligena Lesson, 1832

1.8 Coeligena lutetiae (DeLattre & Bourcier, 1846)

1.8 Coeligena lutetiae (DeLattre & Bourcier, 1846)

39, sometimes with a few violet feathers; underparts yellowish Spectrum Green 62. The remainder is similar to the male. Immatures similar to the females.

Figure 2.8.1. Geographic distribution of C. lutetiae. The polygons represent the geographic pools, the dots the localities.

1.8.3 Geographic variation analysis

A total of 149 skins of Coeligena lutetiae (89 males, 54 females and 6 immatures) was examined. The localities were grouped into nine pools (Fig. 2.8.1). Below is the list of these pools, ordered from north to south, and the descriptions of the corresponding areas.

1. Quindio: located west of Quindio, in the Central Andes of western Colombia, between 2400 and 3900 m a.s.l.

2. Cauca: located from Nevado del Huila National Park southwards to the Puracé National Park, in the Central Andes of southwestern Colombia, between 1700 and 3800 m.

3. Llorente: located south of Pasto, on the eastern slopes of the southern Colombian

4. Carchi: located between Maldonado and Tulcan, on the western slopes of the northern Ecuadorian Andes, near the border with Colombia, between 3100 and 3300 m.

5. Pichincha: located in the vicinity of Quito and the Pichincha Volcano, on the western slopes of the northern Ecuadorian Andes, between 2700 and 4800 m.

6. Papallacta: located in the vicinity of Baeza and the Sumaco Volcano, on the eastern slopes of the northern Ecuadorian Andes, between 2500 and 3800 m.

7. Tungurahua: located on the Cordillera Llanganates, northeast of Tungurahua, on the eastern slopes of the Andes of central Ecuador, between 2500 and 4000 m.

8. Loja: located between Zamora and the headwaters of the River Jubones, on the eastern slopes of the southern Ecuadorian Andes, between 2800 and 3300 m.

9. Lagunillas: located in the border of Ecuador with Peru, on the eastern slopes of the Andes, between 2600 and 3000 m.

1.8.3.1 Morphometric variation

Specimens of Coeligena lutetiae showed significant sexual dimorphism in all the measured parameters, the males having shorter bills, longer wings and longer, more deeply forked tails than the females (Tab. 2.8.1). Significant morphometric variation was also found between pools and between sexes within the pools, the sexual morphometric differences tending to be more significant towards central Ecuador (Fig. 2.8.3-4). The shortest bills and the least significant sexual dimorphism (p<0.05) were found in the northernmost and southernmost pools of Quindio (pool 1, males: 36.8±0.61 mm; females: 39.1±1.19 mm) and Lagunillas (pool 9, males: 36.2±0.78 mm; female: 41.0 mm), whereas both sexes from the pools of Llorente (3), Carchi (4), Papayacta (5) and Pichincha (6) had the longest bills (p<0.01 bill-length males: 38.1-39.4 mm, females: 41.1-42.5 mm), with the most significant sexual dimorphism. The wing-length showed the same tendency, the specimens from Pichincha (5) and Papayacta (6) exhibiting the longest wings with the most significant sexual dimorphism (males: 76.8-77.3 mm; females: 72.5-73.4 mm). No other significant difference was found between the pools.

1.8.3.1 Plumage variation

The plumage coloration pattern in Coeligena lutetiae from Quindio, in central Colombia

(pools 6 to 9), showed very little geographical variation, the males with black heads; frontal spot large, metallic bluish Spectrum Green 62; nape and upperparts blackish Dark Green 262;

lower back less black; upper tail-coverts Olive-Green 46. The chin and throat are bluish metallic Parrot Green 260; gular spot metallic violet Spectrum Blue 69; underparts like the throat; belly more yellowish; under tail-coverts Parrot Green 160 occasionally bordered Warm Buff 118; tail feathers are Olive-Green 46 with no tips. The wings-coverts are less blackish than the upperparts; primary and secondary remiges Raw Umber 223; tertiary remiges completely Cinnamon 39, tipped Raw Umber 223. The females had head and upperparts very yellowish Dark Green 262, with scaly pattern on the head; upper tail-coverts bronzy Parrot Green 260. The chin and throat are dark Clay Color 123B, bordered with tiny glittering Spectrum Green 62 discs; underparts yellowish

Figure 2.8.2. Geographic distribution of C. lutetiae. The subspecies ranges is represented by the polygons. See text for details.

Dark Green 262 with very few brownish white mixture; under tail-coverts yellowish Parrot Green 260 bordered Clay Color 123B; rectrices Greenish-Olive 49 slightly tipped white. The wings are like in males, with paler tertiary remiges.

The pools located on the western slopes of the Ecuadorian Andes (Carchi, pool 4 and Pichincha, pool 5) presented strikingly constrasting differences, allowing a clear separation

eastern Andean slopes in the greener hue on the upperparts, the dark brownish olive on the upper tail-coverts and the conspicuous white coloration of the tertiary remiges, with the base very pale Cinnamon 39 and tipped Raw Umber 223. The females also differ from those of the eastern Andean, having the upperparts less yellowish; upper tail-coverts with no bronzy hue;

chin, throat and anterior chest pale Clay Color 123B very mixed with white, with only the sides being bordered with tiny glittering bluish Spectrum Green 62 discs; underparts very mottled with white on the chest and with Clay Color 123B on the belly; wings like males, with more Cinnamon 39.

1.8.4 Discussion

The degree of differentiation in the morphometry and plumage coloration and the isolation through the central Ecuadorian Andes from the eastern Andean groups indicate the presence of an unknown race of Coeligena lutetiae, which provisionally will be called C. lutetiae albimacualta ssp nov. (Sánchez Osés 2003,. submitted.).

Table 2.8.1. Descriptive statistics of the morphometric parameters in C. lutetiae. The pools are ordered geographically from north to south

Morphometric parameters

Pools Quindio Cauca Llorente Carchi Pichincha Papayacta Tungurahua Loja Lagunillas

Bill-length

males mean

±S.D.

Max.

Min.

N 36.8 0.61 37.6 36.2 5

37.7 1.30 39.6 36.2 5

38.1 2.32 40.8 32.5 9

39.4 0.15 39.6 39.3 3

39.4 0.98 42.1 37.7 24

39.3 1.40 41.8 36.1 21

40.1 - - - 1

- - - - -

36.2 0.78 37.1 35.6 3 (mm) females

mean

±S.D.

Max.

Min.

N 39.1 1.19 40.4 37.2 5

39.6 1.34 40.5 38.6 2

42.5 1.24 43.8 40.9 4

41.1 4.24 44.1 38.1 2

42.1 1.38 44.3 40.0 21

41.2 1.04 42.7 39.8 13

42.6 0.28 42.8 42.4 2

39.3 1.4 40.3 37.7 3

41.0 - - - 1

Wing-length

males mean

±S.D.

Max.

Min.

N 73.6 1.10 75.3 72.0 6

74.8 2.84 77.6 70.6 5

75.8 2.18 79.3 70.6 10

75.8 0.51 76.4 75.4 3

77.3 2.00 82.3 63.9 32

76.8 1.71 81.3 74.3 25

70.0 - - - 1

- - - - -

72.6 2.77 75.0 69.6 3 (mm) females

mean

±S.D.

Max.

Min.

N 68.6 2.80 70.6 63.8 5

69.0 1.13 69.8 68.2 2

69.6 2.54 71.7 65.9 4

73.0 2.12 74.5 71.5 2

73.4 2.50 79.4 68.6 22

72.5 2.21 78.2 70.7 13

70.7 - - - 1

70.6 0.78 71.1 69.7 3

71.7 - - - 1

tail-length

males mean

±S.D.

Max.

Min.

N 46.0 1.30 47.4 44.5 5

47.6 1.30 49.0 46.2 5

46.8 1.62 49.4 44.4 10

47.4 1.00 48.3 46.3 3

44.7 5.60 51.0 34.0 30

44.2 4.46 49.4 34.2 27

48.7 - - - 1

- - - - -

45.5 1.04 46.7 44.9 3 (mm) females

mean

±S.D.

Max.

Min.

N 43.5 1.92 46.2 41.7 4

41.7 1.31 43.1 40.5 3

42.8 2.21 45.4 40.1 4

43.5 0.14 43.6 43.4 2

43.6 2.36 48.4 39.1 22

42.0 1.76 44.6 38.7 13

43.1 - - - 1

42.6 1.04 43.3 41.4 3

44.2 - - - 1

tail fork-depth

males mean

±S.D.

Max.

Min.

N 11.2 1.28 12.7 9.7 5

12.2 1.27 13.7 11.4 3

9.6 1.66 11.7 7.1 7

10.1 0.40 10.5 9.7 3

6.6 6.62 14.2 -5.7 27

6.8 5.34 11.7 -6.6 25

6.8 - - - 1

- - - - -

11.2 2.00 13.3 9.3 3 (mm) females

mean

±S.D.

Max.

Min.

N 10.0 2.62 11.9 8.2 2

6.6 1.90 7.9 5.2 2

7.6 0.90 8.5 6.7 4

7.0 0.64 7.5 6.6 2

6.6 3.30 13.5 -4.2 21

6.2 2.10 9.7 2.5 12

7.3 - - - 1

5.9 1.42 6.8 4.3 3

8.6 - - - 1

A males B females

sex

Quindio-M Quindio-F Cauca-M Cauca-F Llorente-M Llorente-F Carchi-M Carchi-F Pichincha-M Pichincha-F Papallacta-M Papallacta-F Tungurahua-M Tungurahua-F Loja-F Lagunillas-M Lagunillas-F

pools

30.0 35.0 40.0 45.0

bill-length (mm)

B

B A

A

B

A B

A

B B

A B

B A

A

B

A

A males

B females sex

Quindio-M Quindio-F Cauca-M Cauca-F Llorente-M Llorente-F Carchi-M Carchi-F Pichincha-M Pichincha-F Papallacta-M Papallacta-F Tungurahua-M Tungurahua-F Loja-F Lagunillas-M Lagunillas-F

pools

50.0 60.0 70.0 80.0 90.0

wing-length (mm)

B A

B A

A B B

A

B B

A B A

B A

A B

Figure 2.8.3. Geographic variation of the bill- and wing-length in C. lutetiae. The circles and triangles represent the arithmetic mean of males and females, respectively; and the vertical line the standard error of the mean. The pools are geographically ordered, from north to south.

A males B females

sex

Quindio-M Quindio-F Cauca-M Cauca-F Llorente-M Llorente-F Carchi-M Carchi-F Pichincha-M Pichincha-F Papallacta-M Papallacta-F Tungurahua-M Tungurahua-F Loja-F Lagunillas-M Lagunillas-F

pools

40.0 42.5 45.0 47.5 50.0

tail-length (mm) B

A

B A

B A

B A

B B

A B A

B A

B A

A males B females

sex

Quindio-M Quindio-F Cauca-M Cauca-F Llorente-M Llorente-F Carchi-M Carchi-F Pichincha-M Pichincha-F Papallacta-M Papallacta-F Tungurahua-M Tungurahua-F Loja-F Lagunillas-M Lagunillas-F

pools

4.0 8.0 12.0 16.0

tail fork-depth (mm)

B A

B A

B A

B A

B B BA A B

A

B A

Figure 2.8.4. Geographic variation of the tail-length and tail fork-depth in C. lutetiae. The circles and triangles represent the arithmetic mean of males and females, respectively; and the vertical line the standard error of the mean. The pools are geographically ordered, from north to south.

1.9 Coeligena wilsoni (DeLattre & Bourcier, 1846)