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ORNITOWGIA NEOTROPICAL 8: 107-112, 199; @ The Neotropical Ornithological Society

Taxonomy

THE HYBRID ORIGIN OF A VENEZUELAN TROCHlLID, DISTANS WETMORE & PHELPS 1956

André-A. Weller & Karl-L. Schuchmann

Alexander Koenig Zoological Research Institute and Zoological Museum, Oept. of Ornithology, Working Group: Biology and Phylogeny of Tropical , Adenauerallee 160, 0-53113 Bonn, Germany.

words: Amazilia distans, Amazilia fimbriata, Hylocharis cyanus, hybridization, Táchira, Venezuela.

INTRODUCTION Institution (USNM) collection (Colección Amazilia di5tan5 was described as an endemic Ornitológica Phelps [COP # 60790], Caracas, from SW Táchira, Venezuela (Wetmore & Phelps presently on loan to USNM, as USNM # 1956), and it occupies an isolated position among 461965; cf. Deignan 1961) was examined and the 97 Venezuelan . The compared directly with skins of other taxa reasons for this are that the taxon was discovered (Amazilia, other genera). Data from Amazilia relatively recently and that only one skin specimens in other museums were also included specimen exists. Although the validity of this in the comparisons (see "Acknowledgments"). species has not been questioned by later authors Measurements, rounded to the nearest 0.1 mm (Mayr 1971, Meyer de Schauensee & Phelps (Fig. 2), were made using a digital caliper. Plum- 1978, Sibley & Monroe 1990 among others), and age colors were determined by visual inspection although subsequent, unconfirmed observations under natural light. Because of the presence of of A. di5tan5 have been made, almost nothing is iridescence, morphological evaluation of typical known of its biological or ecological character- hummingbird colors depends to some extent on istics (reviewed in Collar et al. 1992). As a result the subjective judgement of the taxonomist (see of its apparent extreme rarity and its relatively also Howell 1993); vernacular names were ac- threatened habitat, which can be described as cordingly used here to describe colors. With mixed tropical and semideciduous forest (Huber regard to the assessment of hybrid origin, the & Alarcón 1988), the species is classified as methods of geographical and phenological ex- critically endangered (Collar et al. 1992). How- clusion of potential parental speciesexplained in ever, as we point out below, the unusual colora- detail in Graves (1990) were employed. tion of the holotype does not allow it to be According to Wetmore & Phelps (1956) the assigned to any of the currently recognized type specimen of Amazilia distans carne from South American Amazilia species groups. We "El Salao (300 meters), near Burgua, Táchira, hypothesize that A. di5tan5 is a hybrid namely Venezuela," where it was collected by R. Urbano Hylochari5 cyanu5 x Amazilia fimbriata. in 1954. Paynter (1982) gives the site coordinates as 07°26'N, 72°00'W. The specimen was sexed MATERIALS AND METHODS as male. Collar et al. (1992: 492) state that "There In the course of biogeographical and taxonomic are apparently two specimens, a male in [the] studies of the hummingbird Amazilia, the USNM 'the type' and a male in [the] COP;" type specimen of Amazilia distans in the N ation- these are, in fact, one and the same specimen al Museum of Natural History, Smithsonian (M. Lentino, pers. comm.).

107 RESULTS AND DISCUSSION combination events (e.g., basic plumage color, dark face mask, differently colored throat, breast, Characteristic plumage features of the USNM and tail pattern) only the genera Amazilia, Chlo. male are its glossy turquoise blue crown and sim- rostilbon, Hylocharis and Lepidopyga remained. ilarly-colored throat and breast. The chin and The speciesin question are all relatively uniform upper throat show gray-white feather disks. The in possessing dark green dorsal plumage with uppertail coverts appear copper to faint red-vio- mostly contrasting tail feathers. let, the inner rectrices are blackish bronze green Since intermediate characters are produced by while the outer ones are more blue-black. On hybridization, then given the turquoise blue pat- the underparts, a small white patch on the breast tern of the head and breast plumage, one parent , and a gray abdominal coloration are conspicuous species can be postulated with a blue or violet in the otherwise faintly gold-green plumage. The coloration in that area. In the genera named, this undertail coverts have a dark center and a nar- feature is prominent only in the male Hylocharis row white border. The bill appears horn-colored cyanus.This speciesis strongly sexually dimorphic; with a dark brown tip. On the basis of its pheno- the female lacks the contrasting head and breast type Amazilia distans was originally judged by coloration and the green mid-belly of the male, Wetmore & Phelps (1956) and by Mayr (1971) to and has instead underparts dominated by grayish be closely related to A. fimbriata. Our own mor- tones. The disks of the male chin and uppermost phological comparisons disclosed few, if any, pa- throat feathers appear whitish to cream-colored. rallels with the plumage patterns of recognized Both sexes show a gold-bronze rump, grading Amazilia species groups (3-4 subgenera), or into the dark red-violet uppertail coverts, and with other trochilid genera native to northwest South America, and we were therefore led to hy- dark steel blue rectrices. The adult male under- pothesize that the specimen was a hybrid. tail coverts are the same color as the tail, while A number of Venezuelan trochilid genera can those of the subadult males and the females are be readily excluded as potential hybridization brownish. partners, based on the position and altitude of The characters described are found only in the type locality (Acestrura, Anthracothorax, Avo- one Hylocharis taxon in the type locality, the cettula, Boissonneaua, Calliphlox, Chalcostigma, subspecies H. cyanus viridiventris. This sub- Chrysolampis, Discosura, Ensifera, Eriocnemis, specieshas a disjunct distribution, with a north- western population from NE Colombia to W Heliangelus, Heliothryx, Hylonympha, Lafresnaya, Leucippus, Lesbia, Metallura, Oxypogon, Poly- Venezuela (Fig. 1) and a more easterly one (Vene-- plancta, Polytmus, Popelairia, Ramphomicron, zuela south of the Orinoco to SE Colombia, the Topaza). Since the characters of the type speci- Guianas, N Bolivia, and SE Brazil). men point unambiguously to it being a member The main criteria to be taken into account of the , the subfamily Phaethornith- when determining the hybridizing partner are inae (e.g., Phaethornis spp.) can be eliminated as the white breast patch, the gray abdominal a source of parental species. coloration, the blackish bronze green inner tail The morphometric results (Fig. 2) show that feathers and the slight forking of the tail. Because the specimen is a medium-sized trochiline with- of the absence of white breast markings and a out extreme lengthening ( Coeligena, Doryfera, clearly forked tail (the inner rectrices are much Heliomaster) or clear curvature of the bill shortened), Lepidopyga goudoti and the local (Campylopterus, Sternoclyta). In addition, the speciesC. stenura and C. mellisugus absence of ornamental plumage features such as cannot be considered as potential partners. Of lengthened crown, throat or tail feathers excludes the remaining genus Amazilia, two speciesoccur Aglaiocercus kingi and the Lophornis species. in southern Táchira, Venezuela, A. viridigaster Further narrowing of the candidate speciespool and A. fimbriata. However, the local viridigaster was possible only by examining plumage pat- population is characterized by rather purplish terns. tail feathers. While it is true that females have After consideration of Amazilia distans color grayish underparts, the white breast patch is characters that could not have resulted from re- lacking in both sexes. SHORT COMMUNICATIONS

Therefore, what remains to be determined is & Phelps 1978) ca~not be clearly distinguished whether Amazilia fimbriata could be the parent from the form elegantissima found further to the of A. distans. The species occurs sympatrically east. One character that all representatives of with Hylocharis cyanus in southern Táchira and A. fimbriata, including elegantissima, have in probably also at the eastern edge of the Vene- common with each other, is that the center of zuelan Andes between the deptos. of Mérida and the belly is pure white, and, above this, the lower Portuguesa (Fig. 1; Osés 1995). A. fimbriata breast bears a small white patch. As in other shows only slight subspecific variation through- Amazilia species there is no marked sexual out its range. Some of those in the Andes (fZuvia- dimorphism. Females possess a rather lighter- tilis, laeta) have a turquoise-iridescent throat, colored throat and broader white edges to the while those at the eastern limit of the range undertail coverts, as well as light gray tips to show, among other features, pure white undertail the outer tail feathers. The first two characters coverts (Brazil: nigricauda, tephrocephala). Two and the white breast patch are found also in subspecies have been described from northern A. distans. and western Venezuela, A. f elegantissima (Todd All of our measurements of the type speci- 1942) and A. f obscuricauda (Zimmer & Phelps men (Fig. 2) are intermediate between those 1951). However, notwithstanding possible dif- of female Amazilia fimbriata elegantissima and ferences in the colors of the uppertail coverts and male Hylocharis cyanus viridiventris. Comparing central rectrices (more strongly purplish), the the morphological results and the morpho- subspecies obscuricauda from the western states metric data, the conclusion can be drawn that from Cojedes to Táchira (Meyer de Schauensee A. distans is in reality a hybrid of both taxa. In

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( ( 10° ) í ~

..;)

5°

750 700 650 600

FIG. 1. Distribution of Amazilia fimbriata elegantissima and Hylocharis cyanus viridiventris (northern popula- tion), area of possible hybridization between these taxa, and type locality of Amazilia distans. The ranges shown incorporate localities of examined specimens (seeAppendix) as well as data in Meyer de Schauensee(1949), Meyer de Schauensee & Phelps (1978), Hilty & Brown (1986), and Osés (1995).

109 view of the fact that there have been several Further hybridizations involving Amazilia unconfirmed sightings of A. distans it must be species have only been confirmed at the intra- presumed that hybridization between the two generic level (Table 1; cf. Gray 1958). According species in SW Venezuela occasionally occurs. to our investigations, a Bogotá specimen known Interestingly, a hybrid of both species(H cyanus as 1haumatias caeruleicepsGould 1860 cannot be x A. f nigricauda) has already been described considered an Amazilia versicolor milleri x Chry- from Bahia, Brazil, by Berlioz (1929). According suronia oenone (Simon 1910) hybrid. The same to that author, its characters are even closer to applies to the Cyanomyia ( = Amazilia) salvini A. fimbriata than in the A. distans type, reflect- Brewster 1893 type specimen. This bird was ing in the coloration of its median lower belly obtained in N Mexico (Sonora) and was later , and undertail coverts the pure white abdominal identified by Griscom (1934) as an Amazilia parts of A. f nigricauda. In addition, the Berlioz violiceps conjuncta x Cynanthus latirostris hybrid, specime:n shows a similar turquoise pattern in an opinion with which Phillips (1965) con- the throat and upper breast, but apparently more curred. However, our examination of the type scattered, bluish-green crown feathers. Thus, its specimen yielded no intermediate character com- dorsal coloration strongly resemblesA. fimbriata. bination of those species.In fact, its morphologi-

H.c.v A.d A.f.e H.c.v A.d. A.t:e.

Bill Wing

H.c.v. A.d. A.f.e H.c.v A.d. A.t:e,

Rectrix 1 Rectrix 5

FIG. 2. Morphometric ranges and means for male Hylocharis cyanus viridiventris, male Amazilia distans, and female Amazilia fimbriata elegantissima. Measurements of the bill include the operculum. For statistical reasons, Hylocharis and Amazilia specimens from populations outside Táchirawere included in the analysis; localities are given in the Appendix.

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Amazilius Ocai Gould 1859 Berlioz (1932)

Amazilia bangsi Ridgway 1910 Bangs (1930) Berlioz (1929)

Amazilia distans Wetmore & this study Phelps 1956 Butler (1932) Berlioz (1937) Dickey & van Rossem (1938)

cal and morphometric characteristics indicate ready graphs and checked the English text. We that it is an aberrant individual of A. violiceps appreciate critical comments on the manuscript ellioti (Weller, in prep.), an observation sup- by W. Bock and A. Andors, AMNH, New York. ported by evidence of the relatively frequent This work was supported by grants from the occurrence of phenotypical mutants throughout Gesellschaft für Tropenornithologie, Germany, the range of this race (Friedmann et al. 1950). the Field Museum of Natura1 History, Chicago, Only one further intergeneric hybrid in- the Frank M. Chapman Memorial Fund, New volving Hylocharis is mentioned in the literature; York, and the Smithsonian ResearchOpportuni- intrageneric hybrids have apparently not yet ties Fund, Washington, D. C. been demonstrated. A series of three specimens of Eucephala ( = Hylocharis) pyropygia Salvin & REFERENCES Godman 1881 was identified by Berlioz (1938) as Bangs,O. 1930. Types of birds now in the Museum of consisting of Chlorostilbon aureoventris puchera- Comparative Zoology. Bull. Mus. Comp. Zool. 70: ni x Hylocharis cyanus hybrids. The potential 145-426. ability of the latter species to hybridize with Berlioz, J. 1929. Un cas nouveau d'hybridité chez les various genera can possibly be accounted for by Trochilidés. Oiseau 10: 340-343. its wide range in South America and by unspeci- Berlioz, J. 1932. Notes critiques sur quelques Trochili- fied biological (bioacoustic?) characteristics. dés du British Museum. Oiseau 2 (n. s.): 530-534. Berlioz, J. 1937. Three cases of presumed natural ACKNOWLEDGMENTS hybrids among Trochilidae. Ibis, XIV series, I: We thank the curat6rs and scientific staff of the 105-109. following museums for lending us specimens and Berlioz, J. 1938. Notes critiques sur des Trochilidés. Oiseau 8 (n.s.): 3-19. for otherwise facilitating our research: American Brewster, W. 1893. Description of a new hummingbird Museum of Natural History, New York; Field from northern Mexico. Auk 10: 214-215. Museum of Natural History, Chicago; Museu de Butler, A. L. 1932. Notes on humming-birds. Bull. Biologia Mello-Leitao, Santa Teresa; Naturmu- Brit. Orn. d. 52: 129-131. seum Senckenberg, Frankfurt/M.; National Mu- Collar, N. J., Gonzaga, L. P., Krabbe, N., Madroño- seum of Natural History, Washington, D. C.; and Nieto, A., Naranjo, L. G., Parker, T. A., III, & Zoologisches Forschungsinstitut und Museum D. C. Wege. 1992. Threatened birds of the Americas. A. Koenig, Bonn. In particular, we are grateful to Part 2. Third edition. Cambridge, U. K. Dr. G. Graves and P. Angle both of whom as- Deignan, H. 1961. Type specimens of birds in the sisted A. W. while at the USNM. M. Leutfeld, T. United States National Museum. Bull. U. S. Nat. Züchner, and B. Hillcoat helped prepare camera- Mus. 221: 1-718.

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Dickey, D. R., & A. J. van Rossem. 1938. The birds of Ridgway, R. 1910. Diagnosis of new forms of Micro- El Salvador. Field Mus. Publ. Chicago (Zool.) 23: podidae and Trochilidae. Proc. Biol. Soc. Wash. 23: 1-609. 53-55. Friedmann, H., Griscom, L., & R. T. Moore. 1950. Salvin, O., & F. D. Godman. 1881. On some new and Distributional check-list of the birds of Mexico. little-known species of Trochilidae. Ibis, IV series, Part 1. Pac. Coast Avifauna no.29: 1-102. V: 595-597. Gould, J. 1859. Descriptions of four new species of Sibley, C. G., & B. L. Monroe, Jr. 1990. Distribution humming-birds from Mexico. Ann. Mag. Nat. and taxonomy of birds of the world. New Haven. Hist., series 3; 4: 96-98. Simon, E. 1910. Notes critiques sur les Trochilidés. iii. Gould,J. 1860. Descriptions of twenty-two new species Sur quelques hybrides. Rev. Fr. d'Orn. 1: 177-178. of humming-birds. Proc. Zool. Soc. London 28: Todd, W. E. C. 1942. List of in the col- 304-312. lection of the Carnegie Museum. Ann. Carnegie Graves, G. R. 1990. Systematics of the .green-throated Mus. 29: 271-370. sunangels" (Aves: Trochilidae): valid taxa or hybrids? Wetmore, A., & W. H. Phelps, Jr. 1956. Further addi- Proc. Biol. Soc. Wash. 103: 6-25. tions to the list of birds of Venezuela. Proc. Biol. Gray, A. P. 1958. Bird hybrids. Farnham Royal, Bucks, Soc. Wash. 69: 1-12. England. Zimmer, J. T., & W H. Phelps. 1951. Four new sub- Griscom, L. 1934. The ornithology of Guerrero. Bull. species of birds from Venezuela. Amer. Mus. Mus. Comp. Zool. 75: 365-422. Novit. 1395: 1-10. Hilty, S. L., & W. L. Brown. 1986. A guide to the birds

of Colombia. Princeton, N. J. Accepted 28 June 1997. Howell, S. N. G. 1993. A taxonomic review of the Green-fronted Hummingbird. Bull. Brit. Orn. Cl. 113: 179-187. Huber, O., & C. Alarc6n. 1988. Mapa de vegetaci6n de Venezuela. Caracas: Ministério de Ambiente y de los Recursos Naturales Renovables (Divisi6n de APPENDIX Vegetaci6n). Mayr, E. 1971. New birds described from 1956 to 1965. Localities of specimens of Hylocharis cyanus viridiven- J. Orn. 112: 302-316. tris and of Amazilia fimbriata elegantissima examined Meyer de Schauensee,R. 1949. The birds of the Repu- in the study (from N to S; coordinates after Paynter blic of Colombia. Part 2. Caldasia, 5,23: 381-644. 1982, Paynter & Traylor 1981). Meyer de Schauensee,R., & W. H. Phelps, Jr. 1978. Hylocharis cyanus viridiventris. Colombia: Carrai- A guide to the birds of Venezuela. Princeton, N. J . pía, La Guajira (11°16'N, 72°22'W); El Bosque, La Osés, C. S. 1995. Distribuci6n geográfica de la familia Guajira (c. 11°09'N, 72 °20'W); Petrólea, Norte de Trochilidae (Aves: ) en Venezuela con Santander (08 °30'N, 72 °35'W); Villa Felisa, Norte de algunas implicaciones biogeográficas. Unpublished Santander (c. 07°45'N, 72°33'W); Venezuela: Cerro master thesis, Universidad Central de Venezuela, Yapacana, Amazonas (c. 03 °42'N, 66 °45'W); Tamata- Facultad de Ciencias, Escuela de Biología, Caracas. ma, Amazonas (03 °09'N, 65°50'W). Paynter, R. A., Jr. 1982. Ornithological gazetteer of Amazilia fimbriata elegantissima. Venezuela: Boca Venezuela. Museum of Comparative Zoology, Cambridge, Mass. Tocuyo, Falcon (11 °03'N, 68 °21'W); Caracas, Distrito Paynter, R. A., Jr., & M. A. Traylor, Jr. 1981. Ornitho- Federal (10°30'N, 66°55'W); San Esteban; Carabo- logical gazetteer of Colombia. Museum of Compa- bo (10°26'N, 68°01'W); Maracay, Aragua (10°15'N, rative Zoology, Cambridge, Mass. 67°36'W); Ocumare del Tuy, Miranda (10°07'N, Phillips, A. R. 1965. Notas sistematicas sobre aves 66 °46'W); Guanta, Anzoátegui (10° 14'N, 64°36'W); Mexicanas, III. Rev. Soc. Mex. Hist. Nat. 25: Maturín, Monagas (09°45'N, 63° 11'W); Hato Flores 217-242. Moradas, Guárico (c. 08 °25'N, 67°30'W).

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