WHY ARE NEOTROPICAL BIRDS MORE COLOURFUL THAN NORTH AMERICAN BIRDS?
By
First Published in The Avicultural Magazine Vol. 78 No. 3
Copyright © 1972 Avicultural Society, Published with Permission
When one of us (MFW) was in Costa Rica for a summer, a common question from non-ornithological colleagues was "Why are tropical birds more colourful than those in the US?" The present exercise was stimulated by that question since it was not clear that tropical birds are more colourful, much less why.
We have attempted to classify the avifauna of South America, North America, and Europe into categories of "colourful" or "not so", but we ignored degrees of colourfulness. Criteria for the division are necessarily subjective, but we tried to be consistent in their application. If the plumage of a species was described as having fairly large patches of bright yellow, orange, red, blue, purple or green, or any combination thereof, it was called "colourful". For example, the pileated and red-headed woodpeckers of North America were called colourful because of the amount of red on the head of at lease one sex, but the downy and red-bellied woodpecker were not. Although some people would undoubtedly categorise species by somewhat different means, we feel that the consistency of our usage validates the approach. Decisions about the colourfulness were based on descriptions and pictures in three sources: Meyer de Schauensee (1970), Robbins, et al. (1966), and Petersen, et al. (1966). Some species that could be considered strimming, flashy and conspicuous (such as American magpies) were not, however, called "colourful", for their conspicuousness is derived from e.g. black and white patterns, and we felt that this was not included in our colleagues' questions. We omitted consideration of primarily aquatic and nocturnal families for the same reason.
At the same time, we recorded the incidence of sexual dichromatism in each species and the habitat of each. Habitat, when given, usually referred to the breeding habitat, at least for North American and European birds. Habitats were classified into two general types: "open" (marshes, grassland, sand, etc.) and "wooded" (all woodlands and forests, scrub, etc.). Finer subdivisions were not practicable owing to the authors' differences in descriptions. The South American avifauna was split into "lowland tropical" and "non-tropical", which refers both to latitudinal and altitudinal zones that are not tropical according to Meyer de Schauensee (op. cit.), and to wide-ranging species occupying many zones including tropical.
Regional differences in frequency of colourfulness and dichromatism are clear. χ2 tests (P < 0.05) between pairs of regions indicate the following ranking:
DICHROMATISM:
North America (39%) = South American tropics (39%) > Europe (32%) = South American nontropics (26%)
COLOURFULNESS:
South American tropics (32%) > South American nontropics (27%) = North America (23%) > Europe (10%)
Birds of the South American lowland tropics are therefore more frequently colourful than those of North America or the South American nontropics; a consideration of degree of colourfulness or combination of colour would have emphasised the difference. European birds are relatively dull in colour. Colourfulness is not particularly associated with a tendency towards dichromatism, for North American and South American tropical birds have a higher proportion of sexually dichromatic forms than the others; Europe and the South American nontropical avifaunas tend toward sexual monomorphism.
In almost all cases the incidence of sexual dichromatism and colourfulness in open habitats is similar to their incidence in the entire regional avifauna. An exception is found in the South American non tropics, when colourfulness in open habitats is significantly (χ2, P < 0.05) less frequent than in the whole avifauna. Given that a species lives in an open habitat then, it usually has the same probability of being colourful as any species taken from that avifauna.
Although in all four regions the percentage of colourful and sexually dichromatic species that live in the "open" habitats is the markedly less than the percent that live in wooded habitats, the relative number of species living in the open is also less than that of wooded areas. Neither kind of habitat, then, is relatively more conducive to the development of dichromatism or colourfulness.
Certain families clearly contain relatively more dichromatic or colourful species than other families. In addition, for each region, certain families are the largest contributors of species to the dichromatic and colourful portions of the avifauna. However, seldom is an entire family dichromatic or colourful, which argues to some extent against genetic fixation of such a trait within a phylogenetic lineage, in most cases.
Seasonal sexual dichromatism in Nectariniidae, Parulidae, Icteridae (Hamilton 1961, Hamilton & Barth 1962, Skutch 1957), Central American Fringillidae and Thraupidae (Skutch 1940), and in African Ploceidae (Moreau 1960) was associated with duration and nature of the pair bonds, social behaviour in the non breeding season, degree of migratoriness, and, for some, habitat. For comparisons of regional avifaunas, however, the explanations do not seem to be the general ones. For instance, dichromatism (seasonal and permanent) is least developed in the European and South American nontropical avifaunas, but we expect that long distance migration or wandering is probably more common there than in the South American lowland tropics.
Perhaps a general explanation is unavailable. Hamilton and Barth (op. cit.) list several kinds of selective factors that influence the evolution of species-specific plumage patterns. These, and perhaps others, probably vary in their importance amongst species. Colourfulness is equally difficult to assess, especially in view of our ignorance of the possible cryptic nature of the bright plumages amidst brightly coloured flowers, fruits and leaves, and of our lack of information concerning relative predation rates.
Summary
Tropical birds of South America are more frequently colourful than those of North America or nontropical areas of South America; European birds are the least colourful of all. Assessment of degree of colourfulness or colour combinations would likely increase most of these differences. Sexual dichromatism is most common amongst members of the North American and South American tropical avifauna, and least in Europe and the South American nontropics. Habitat association of these characters are not generally evident, and taxonomic associations are not very helpful. At this time we seem to lack a general explanation for the evolution of the observed tendencies.
Literature Cited
- HAMILTON, T.H. 1961 On the functions and causes of sexual dimorphism in breeding plumage character of North American species of warblers and orioles. Am. Nat. 95 : 121-123
- HAMILTON, T.H. and BARTH, R.H. 1962 The biological significance of season change in male plumage appearance in some New World migratory bird species. Am. Nat. 96 : 129-144
- MEYER DE SCHAUENSEE, R. 1970 A Guide to the Birds of South America. Livingston Publ. Co., Wynnewood, Pa.
- MOREAU, R.E. 1960 Conspectus and classification of the Ploceine weaverbirds. Ibis 102 : 298-321
- PETERSON, R.T., MOUNTFORT, G. and HOLLOM, P.A.D. 1966 A Field Guide to the Birds of Britain and Europe. H.M. Co., Riverside Press, Cambridge, Mass.
- ROBBINS, C.S., BRUUN, B., and ZIM, H.S. 1966 A Guide to the Field Identification. Birds of North America. Golden Press, N.Y.
- SKUTCH, A.F. 1940 Some aspects of Central American birdlife. Sci. Monthly, 51 : 409-418; 500-511
- SKUTCH, A.F. 1957 The resident wood warblers of Central America. In Griscom, L., Sprunt, A., et al. The Warblers of America. Devin-Adair, N.Y. : 275-285
APPENDIX 1
Numbers of sexually dichromatic and colourful species in each family. Winter visitors from North America are included in the South American tallies because many of these live for several months in lower latitudes. Accidental and exotic species and occasional visitors are omitted. The systematic classification of each source book are retained, and brackets used to indicate correspondence between regions.
A | Europe | #spp. | #dichromatic (%) | #colourful (%) |
Accipitridae | 28 | 6 (21%) | 0 | |
Falconidae | 10 | 4 (40%) | 0 | |
Tetraonidae | 5 | 4 (80) | 0 | |
Phasianidae | 6 | 1 (17) | 0 | |
Turnicidae | 1 | 0 | 0 | |
Otididae | 3 | 1 (33) | 0 | |
Burhinidae | 1 | 0 | 0 | |
Glareolidae | 3 | 0 | 0 | |
Pteroclidae | 3 | 3 (100) | 0 | |
Columbidae | 6 | 0 | 0 | |
Cuculidae | 3 | 0 | 0 | |
Apodidae | 3 | 0 | 0 | |
Meropidae | 1 | 0 | 1 (100) | |
Coraciidae | 1 | 0 | 1 (100) | |
Upupidae | 1 | 0 | 0 | |
Picidae | 10 | 6 (60) | 1 (10) | |
Alaudidae | 11 | 1 (9) | 0 | |
Hirundinidae | 5 | 0 | 2 (40) | |
Motacillidae | 10 | 3 (30) | 2 (20) | |
Laniidae | 5 | 2 (40) | 0 | |
Bombycillidae | 1 | 0 | 0 | |
Cindidae | 1 | 0 | 0 | |
Troglodytidae | 1 | 0 | 0 | |
Prunellidae | 1 | 0 | 0 | |
Musicapidae | 77 | 29 (38) | 5 (6) | |
Paridae | 11 | 0 | 2 (18) | |
Sittidae | 4 | 0 | 0 | |
Certhiidae | 2 | 0 | 0 | |
Emberizidae | 14 | 10 (71) | 2 (14) | |
Fringillidae | 19 | 13 (68) | 8 (42) | |
Ploceidae | 5 | 2 (40) | 0 | |
Sturnidae | 3 | 0 | 1 (33) | |
Oriolidae | 1 | 1 (100) | 1 (100) | |
Corvidae | 12 | 0 | 1 (9) | |
TOTAL | 268 | 86 (32%) | 27 (10%) |
B | North America | #spp. | #dichromatic (%) | #colourful (%) |
Cathartidae | 3 | 0 | 0 | |
Accipitridae | 23 | 2 (9) | 0 | |
Pandionidae | 1 | 0 | 0 | |
Falconidae | 7 | 3 (43) | 0 | |
Meleagrididae | 1 | 0 | 0 | |
Cracidae | 1 | 0 | 0 | |
Tetraonidae | 10 | 9 (90) | 0 | |
Phasianidae | 6 | 4 (67) | 0 | |
Columbidae | 11 | 1 (9) | 0 | |
Cuculidae | 6 | 0 | 0 | |
Apodidae | 4 | 0 | 0 | |
Trochilidae | 15 | 14 (93) | 15 (100) | |
Psittacidae | 1 | 0 | 1 (100) | |
Trogonidae | 1 | 1 (100) | 1 (100) | |
Picidae | 22 | 15 (68) | 8 (36) | |
Cotingidae | 1 | 1 (100) | 0 | |
Tyrannidae | 31 | 1 (3)0 | 3 (9) | |
Alaudidae | 1 | 0 | 0 | |
HIrundinidae | 8 | 1 (13) | 2 (25) | |
Corvidae | 15 | 0 | 6 (40) | |
Paridae | 14 | 2 (14) | 0 | |
Chameidae | 1 | 0 | 0 | |
Cinclidae | 1 | 0 | 0 | |
Sittidae | 4 | 2 (50) | 0 | |
Certhiidae | 1 | 0 | 0 | |
Troglodytidae | 10 | 0 | 0 | |
Mimidae | 10 | 0 | 0 | |
Turdidae | 13 | 7 (54) | 6 (46) | |
Sylviidae | 5 | 4 (80) | 0 | |
Motacillidae | 4 | 0 | 1 (25) | |
Bombycillidae | 2 | 0 | 0 | |
Ptilogonatidae | 1 | 1 (100) | 0 | |
Laniidae | 2 | 0 | 0 | |
Coerebidae | 1 | 0 | 1 (100) | |
Vireonidae | 12 | 1 (9) | 1 (9) | |
Parulidae | 53 | 36 (68) | 25 (47) | |
Icteridae | 20 | 18 (90) | 11 (55) | |
Thraupidae | 4 | 4 (100) | 4 (100) | |
Fringillidae | 77 | 32 (42) | 16 (21) | |
TOTAL | 403 | 159 (39%) | 101 (25%) |
C | South America | #spp. | #dichromatic (%) | #colourful (%) | #spp. | #dichromatic (%) | #colourful (%) |
TROPICAL | NONTROPICAL | ||||||
Rheidae | 24 | 7 (29) | 0 | 2 | 0 | 0 | |
Cathartidae | 5 | 0 | 0 | 18 | 0 | 0 | |
Accipitridae | 36 | 3 (9) | 0 | 1 | 1 (100) | 0 | |
Pandionidae | 11 | 2 (18) | 0 | ||||
Falconidae | 11 | 0 | 0 | 1 | 0 | 0 | |
Cracidae | 25 | 7 (28) | 0 | 11 | 3 (27) | 0 | |
Phasianidae | 5 | 3 (60) | 0 | 6 | 0 | 0 | |
Opisthocomidae | 1 | 0 | 0 | 7 | 5 (71) | 0 | |
Psophiidae | 3 | 0 | 0 | ||||
Cariamidae | 2 | 0 | 0 | ||||
Burhinidae | 2 | 0 | 0 | ||||
Thinocoriidae | 4 | 3 (75) | 0 | ||||
Columbidae | 26 | 6 (23) | 0 | 15 | 2 (13) | 0 | |
Psittacidae | 82 | 11 (13) | 82 (100) | 26 | 3 (12) | 26 (100) | |
Cuculidae | 22 | 0 | 0 | 1 | 0 | 0 | |
Apodidae | 15 | 1 (7) | 0 | 6 | 0 | 0 | |
Trochilidae | 133 | 79 (59) | 84 (63) | 96 | 55 (57) | 72 (75) | |
Trogonidae | 11 | 11 (100) | 11 (100) | 3 | 3 (100) | 3 (100) | |
Motmotidae | 4 | 0 | 4 (100) | ||||
Galbulidae | 15 | 7 (47) | 10 (67) | ||||
Bucconidae | 32 | 0 | 0 | ||||
Capitonidae | 12 | 9 (75) | 12 (100) | ||||
Ramphastidae | 33 | 8 (24) | 33 (100) | 8 | 0 | 8 (100) | |
Picidae | 72 | 52 (72) | 8 (11) | 7 | 7 (100) | 2 (29) | |
Dendrocoloptidae | 42 | 0 | 0 | 4 | 0 | 0 | |
Furnariidae | 111 | 0 | 0 | 97 | 0 | 0 | |
Formicariidae | 195 | 171 (88) | 0 | 32 | 7 (22) | 0 | |
Rhinocryptidae | 11 | 6 (55) | 0 | 17 | 3 (18) | 0 | |
Cotingidae | 57 | 48 (84) | 16 (28) | 16 | 9 (56) | 8 (50) | |
Rupicolididae | 2 | 2 (100) | 2 (100) | ||||
Pipridae | 49 | 40 (82) | 25 (51) | 1 | 1 (100) | 1 (100) | |
Tyrannidae | 202 | 27 (13) | 15 (7) | 106 | 6 | 0 | |
Oxyruncidae | 1 | 1 (100) | 0 | ||||
Phytotomidae | 1 | 1 (100) | 0 | 2 | 2 (200) | 0 | |
Alaudidae | 1 | 0 | 0 | ||||
Hirundinidae | 11 | 2 (18) | 1 (9) | 9 | 0 | 2 (22) | |
Corvidae | 11 | 0 | 7 (64) | 2 | 0 | 2 (100) | |
Troglodytidae | 28 | 0 | 0 | 11 | 0 | 0 | |
Mimidae | 5 | 0 | 0 | 3 | 0 | 0 | |
Turdidae | 20 | 4 (20) | 0 | 12 | 5 (42) | 0 | |
Sylviidae | 8 | 3 (38) | 0 | ||||
Motacillidae | 2 | 0 | 0 | 5 | 0 | 0 | |
Vireonidae | 19 | 0 | 2 (11) | 3 | 0 | 0 | |
Icteridae | 48 | 11 (23) | 30 (63) | 15 | 6 (40) | 8 (53) | |
Parulidae | 27 | 8 (30) | 21 (78) | 21 | 7 (33) | 10 (48) | |
Coerebidae | 19 | 15 (79) | 74 (14) | 18 | 5 (28) | 4 (22) | |
Tersinidae | 1 | 1 (100) | 0 | ||||
Thraupidae | 126 | 69 (55) | 81 (64) | 49 | 13 (27) | 27 (55) | |
Catamblyrhynchidae | 1 | 0 | 0 | ||||
Fringillidae | 112 | 46 (41) | 20 (18) | 73 | 36 (49) | 19 (26) | |
TOTAL | 1678 | 657 (39%) | 539 (32%) | 722 | 186 (26%) | 194 (27%) |