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Noqvtates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Noqvtates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 28 1 1, pp. 1-14, figs. 1-2, tables 1-4 February 20, 1985 Variation and Population Structure of the Tourmaline Sunangel, Heliangelus exortis exortis (Aves, Trochilidae) ROBERT BLEIWEISS' ABSTRACT The nominate form of the polytypic Andean tion. Geographic patterns of variability in the hummingbird the Tourmaline Sunangel, Helian- polychromatism do not appear to result from in- gelus exortis exortis, exhibits dramatic variation creased variability in putative zones ofhybridiza- in a female polychromatism of throat iridescence tion between differentiated populations. No char- within and among populations that has not here- acter breaks that might indicate noninterbreeding tofore been appreciated. An analysis of the geo- parapatric forms were noted. It is concluded that graphic population structure of samples referable H. e. exortis comprises a single biological entity to H. e. exortis was conducted to determine the and taxon within the polytypic H. exortis complex. biological status and historical process of differ- The evolutionary origins of the complex pattern entiation ofpolychromatically differentiated sam- of geographic variation in H. e. exortis are dis- ples. Geographic variation in additional color and cussed. I suggest that biological attributes, in ad- six mensural characters is discordant within and dition to potential geographic barriers, are nec- between the sexes. The polychromatism shows both essary for a full understanding of the geographic regular and mosaic patterns of geographic varia- differentiation seen in H. e. exortis. INTRODUCTION There are few detailed studies of the geo- categories ofisolates, continua (clines and re- graphic population structure ofAndean birds lated phenomena), and secondary contact. (Vuilleumier, 1968, 1980a, 1980b, 1984; There are also classes ofgeographic variation Graves, 1982). These studies have been aimed that do not fit these categories but neverthe- primarily at reconstructing the course ofgeo- less manifest striking patterns. This is espe- graphic speciation by describing geographic cially true in topographically complex moun- variation in terms of Mayr's (1959, 1963) tain systems and oceanic archipelagoes. ' Chapman Research Fellow, Department of Ornithology, American Museum of Natural History. Copyright © American Museum of Natural History 1985 ISSN 0003-0082 / Price $1.90 2 AMERICAN MUSEUM NOVITATES NO. 2811 Unusual forms of geographic variation re- this exceptional form. The present paper treats corded in these regions include changes in geographic variation in the female polychro- sexual dimorphism, sex-limited variation, matism in conjunction with variation in oth- and mosaic, nonclinal variation (Mayr, 1942). er color and mensural characters to examine These complex patterns suggest biological as these questions. well as extrinsic factors as causal agents of geographic variation, and are of general in- ACKNOWLEDGMENTS terest for understanding the full range of fac- I thank the curators of the collections used tors that promote geographic differentiation in this study for providing specimens under and speciation. Because of great interest in their nectar- their care: Dr. F. B. Gill, Academy ofNatural feeding habits, hummingbirds are becoming Sciences, Philadelphia; Dr. J. P. O'Neill, one of the better known avian families from Louisiana State University Museum of Zo- the standpoint ofecology and behaviof. This ology, Baton Rouge; Dr. R. A. Paynter, Jr., knowledge has additional application in stud- Museum of Comparative Zoology; Dr. L. L. ies of the biological basis of geographic dif- Short, American Museum of Natural His- ferentiation, of which hummingbirds in the tory; Dr. D. S. Wood, Carnegie Museum of topographically complex Andes provide Natural History; Dr. R. L. Zusi, National many dramatic examples (Zimmer, 1951a, Museum ofNatural History, Smithsonian In- 1951b, 1952). However, studies of hum- stitution. Drs. W. H. Bossert, L. L. Short, F. mingbird systematics have lagged far behind Vuilleumier, E. E. Williams and Mr. G. C. knowledge of their feeding ecology and be- Mayer made helpful comments on an earlier havior. draft of this paper. Laszlo Meszoly drafted While conducting a survey of geographic figure 1 and Lynn Winter drafted figure 2. variation and speciation patterns in Andean This work was funded in part by a grant from hummingbird genera, I found an unusual form the Frank M. Chapman Fund of the Amer- of color variation in members of the genus ican Museum of Natural History. Heliangelus. Females of several species ex- hibit dramatic variation in the development TAXONOMY OF THE of the highly iridescent throat, or "gorget," HELIANGELUS EXORTIS characteristic of males (Bleiweiss, MS). This COMPLEX "polychromatic" variation is particularly Hummingbirds of the genus Heliangelus pronounced within and among populations Gould, 1848, are endemic to mid-elevations ofthe nominate form ofthe polytypic species (1500-3000 m) ofthe Andes from Venezuela Heliangelus exortis, the Tourmaline Sunan- to Bolivia. Current taxonomic treatments gel. Samples from the central section of the recognize seven species in the genus, some of Central and Eastern Cordilleras of Colombia which are highly polytypic (Peters, 1945; show a range of variation from a nonirides- Meyer de Schauensee, 1966). The Tourma- cent to a fully iridescent gorget similar to line Sunangel, Heliangelus exortis, is cur- males, whereas samples from southern Co- rently treated as a polytypic species com- lombia have only discrete noniridescent and prised ofa nominate form with violet-chinned fully iridescent male-like morphs. In con- and rose-throated males occurring in the three trast, the polychromatism is rare or absent Andean ranges of Colombia south through in samples from the eastern slopes ofthe An- the eastern slope of the Ecuadorian Andes, des in Ecuador. and two apparently allopatric forms with or- Intraspecific sex-limited variation and ange-throated males, H. e. micraster and H. variation in sexually dichromatic features are e. cutervensis, in the Andes of southwestern both rare phenomena in birds. The geograph- Ecuador and northern Peru, respectively ic population structure of Heliangelus e. ex- (Meyer de Schauensee, 1966). A number of ortis is therefore of great interest for (1) as- characteristics support the conclusion that H. certaining whether H. e. exortis should be e. exortis and the orange-throated forms are considered a single species and (2) gaining closest relatives in Heliangelus (Zimmer, insight into the process of differentiation in 1951 b; Fitzpatrick, Willard, and Terborgh, 1985 BLEIWEISS: HELIANGELUS EXORTIS EXORTIS 3 TABLE 1 Distribution and Sample Size for Heliangelus exortis exortis Specimens Examined Cordillera Locality Pooleda West Central Eastb Adults Immatures 1. El Peii6n A West 2. La Aguadita A West 3. El Roble A West 6 4. FusugasugA-Silvania A West 4 5. Hacienda Zuliaba East 7 6. Santa Elena East 2 7. Paramo de Sons6n West 6 2 8. El Zancudo B West 17 1 9. Santa Isabel B West 1 1 10. Laguneta West 43 15 11. Toche; Tolima West 7 6 12. Purace West 1 13. Coconuco West 14. Paletara West 2 15. Almaguer West 6 3 16. Hacienda La Ilusi6n West 8 1 17. La Florida C East 1 18. Coast Range C East 3 19. Cerro Munchique C East 17 1 20. Munchique-El Tambo C East 2 21. Tijeras-Moscopan East 5 22. Guanderal D East 3 23. Cerro Pax D East 8 24. Papallacta E East 7 25. Cuyujua E East 2 26. Baeza E East 3 27. Volcan Sumaco Isolated peak 10 28. San Antonio F East .1 1 29. Ambato F East 1 30. Bafios F East 6 31. Volcan Tungurahua F East 4 32. Planchas East 2 33. Coraz6n West a Letters A-F designate pooled localities used for analysis of geographic variation. b East and west designations under each Cordillera refer to the slope of the range. 1979; Bleiweiss, in prep.). Each can be dis- AND tinguished from most sympatric congeners by MATERIALS METHODS the absence of a white pectoral band, and The present study is based upon 230 spec- uniquely, by white female throat feathers and imens from six museum collections; Amer- under-tail coverts in both sexes. The Purple- ican Museum of Natural History, New York throated Sunangel, H. viola, also lacks a white (AMNH), Academy of Natural Sciences, pectoral band and may be closely related to Philadelphia (ANSP), Carnegie Museum of H. e. exortis (Chapman, 1926). However, H. Natural History, Pittsburgh (CAR), Museum viola is sympatric with the orange-throated ofComparative Zoology, Cambridge (MCZ), members of the H. exortis complex and is Louisiana State University Museum of Nat- thus a distinct species. In my analysis, I define ural History (LSUMZ), and the National Mu- H. e. exortis as all rose-throated populations seum of Natural History, Washington D.C. of the H. exortis complex. (USNM). I used all specimens with adequate 4 AMERICAN MUSEUM NOVITATES NO. 2811 FIG. 1. Geographic distribution ofHeliangelus exortis exortis specimens examined. Locality numbers are identified in table 1. Letters A-F indicate localities pooled for the analysis of geographic variation (see Appendix 1). locality data from throughout the range ofH. on the labels of only two specimens. Imma- e. exortis (table 1, figure 1, see Appendix 1: tures were distinguished by the presence of Specimens Examined). corrugations on the culmen, lacking in adults Study skins were aged and sexed by exter- (Ortiz-Crespo, 1972). Older immatures, rec- nal characters
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