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Full Article Ardeola 50(2), 2003, 223-235 GEOGRAPHIC VARIATION IN THE GREAT PAMPA-FINCH EMBERNAGRA PLATENSIS COMPLEX: EVIDENCE FOR TWO SPECIES Floyd E. HAYES*1 SUMMARY.—Geographic variation in the Great Pampa-Finch Embernagra platensis complex: evidence for two species. Aims: The Great Pampa-Finch Embernagra platensis is represented by four subspecies belonging to two dis- tinct groups occurring in allopatry: (1) nominate E. p. platensis in the east; and (2) E. p. olivascens, E. p. gos- sei and E. p. catamarcanus in the west. Morphological variation within the complex was assessed to infer the taxonomic relationship between the two forms. Location: Southern South America. Methods: Several size, plumage and soft part traits were measured for 339 specimens and analysed with uni- variate and multivariate statistical methods. Results: I found no evidence for long-distance migration, conflicting support for Bergmann’s rule (body size negatively correlated with latitude and positively correlated with elevation), and more support for Gloger’s rule. The structure and colouration of the bill were the most diagnostic traits. The absence of intermediate spe- cimens and lack of clinal variation of bill colouration within each form demonstrates that intergradation eit- her does not occur or is potentially restricted to a narrow, still undiscovered contact zone. Conclusions: Given the likelihood that bill structure and colour, combined with other traits, may represent re- productive isolating mechanisms between the two groups, I tentatively propose the recognition of two species within the E. platensis complex: the monotypic Great Pampa-Finch, E. platensis, and the polytypic Olive Pampa-Finch, E. olivascens. Key words: Embernagra platensis, geographic variation, Great Pampa-Finch, South America, taxo- nomy. RESUMEN.—Variación geográfica en el complejo de especies del Verdón Embernagra platensis: evi- dencias de dos especies. Objetivos: El Verdón Embernagra platensis de Sudamérica esta representado por cuatro subespecies que per- tenecen a dos grupos distintos en alopatria: (1) la raza nominada E. p. platensis en el este; y (2) E. p. olivas- cens, E. p. gossei y E. p. catamarcanus en el oeste. Se estudió la variación geográfica para inferir la relación taxonómica entre los dos grupos. Localidad: El sur de Sudamérica. Métodos: Algunas caracteristicas del tamaño, plumaje, y partes blandas fueron medidas para 339 especime- nes y analizadas con métodos estadísticos univariables y multivariables. Resultados: No se encontró evidencias de la existencia de migración a largas distancias, ni evidencias para comprobar la ley de Bergmann (el tamaño del cuerpo está correlacionado negativamente con la latitud y po- sitivamente con la elevación), pero si se encontraron evidencias para comprobar la ley de Gloger. La estruc- tura y coloración del pico fueron las variables com mayor poder diagnóstico. La ausencia de especimenes in- termedios y la ausencia de variación clinal de la coloración del pico en cada forma, demuestran que la gradación no existe o potencialmente está restringida a un área de contacto bien estrecho y no conocido. Conclusiones: Dada la probabilidad que la estructura y color del pico, combinado con otras caracteristicas, puedan representar mecanismos del aislamiento reproductivo entre los dos grupos, se propone tentativamen- te el reconocimiento de dos especies dentro del complejo de Embernagra platensis: el monotípico Verdón Grande, E. platensis, y el politípico Verdón Olivo, E. olivascens. Palabras clave: Embernagra platensis, variación geográfica, Verdón, Sudamérica, taxonomía. * Department of Life Sciences, University of the West Indies, St. Augustine, Trinidad and Tobago. 1 Present address and correspondence: Department of Biology, Pacific Union College, Angwin, 1 Angwin Ave., CA 94508, USA. e-mail: [email protected] 224 HAYES, F. E. INTRODUCTION possibly being intermediate. Ridgely & Tudor (1989) suggested that platensis and olivascens An accurate assessment of phylogenetic re- might represent distinct species. lationships and speciation processes among sis- The purpose of this study is to evaluate the ter taxa is premised upon a thorough unders- relationships between sibling taxa of the Great tanding of geographic variation, invariably Pampa-Finch by examining sexual dimorphism based upon (but not limited to) an analysis of and geographic variation of morphometric cha- museum specimens (Zink & Remsen, 1986; racters. The results of these analyses are further Remsen, 1995). A primary goal of such an used to (1) evaluate quantitatively Bergmann’s analysis is to evaluate the extent of gene flow rule, which predicts that body size is inversely between adjacent taxa in order to determine or correlated with temperature and humidity (Ja- infer whether primary intergradation (clinal va- mes, 1970); (2) evaluate qualitatively Gloger’s riation), secondary intergradation (hybridisa- rule, which predicts that populations in more tion), or the absence of intergradation (potential humid areas are more heavily pigmented than or complete reproductive isolation) occurs (e.g. those in drier areas (Zink & Remsen, 1986); Mayr, 1963, 1970; Gould & Johnston, 1972; and (3) infer the extent of reproductive isolation Zink & Remsen, 1986). among the taxa and implications for taxonomy. The Great Pampa-Finch Embernagra pla- tensis, a large, olive ‘tanager-finch’ whose ta- xonomic affinities remain unresolved (e.g. Isler Taxonomic background & Isler, 1987; Sibley & Ahlquist, 1990; Sibley & Monroe, 1990; Burns, 1997; Lougheed et In an early revision of Embernagra, Hell- al., 2000), is widely distributed east of the An- mayr (1938) recognised three subspecies of E. des in southern South America. It is a fairly platensis, which had been described initially common and relatively conspicuous inhabitant as separate species: nominate platensis in the of damp grasslands, where pairs or small east, olivascens in the west, and gossei in the groups are territorial and often sing while per- southwest (Fig. 1). Nominate platensis is the ched on the tops of bushes and fence posts (e.g. most widely distributed, the smallest, and the Wetmore, 1926; Belton, 1985; Ridgely & Tu- only race with prominent dusky streaking on dor, 1989; Fjeldsa & Krabbe, 1990; Sick, the back. Olivascens differs from platensis by 1993). It appears to be a sedentary species in averaging larger in size, having a more strongly which long-distance migration has not been re- curved, deeper orange beak with less extensive ported (Ridgely & Tudor, 1989; Chesser, dusky colouration on the upper bill, lacking 1994). streaks or only faintly streaked on the back, A polytypic species, the Great Pampa-Finch and having a paler loral region, chin and abdo- is represented by four described subspecies be- men (Chubb, 1918; Wetmore, 1926; Hellmayr, longing to two distinct groups: (1) nominate E. 1938; Short, 1975; Contreras, 1980). The extent p. platensis, widely distributed in the east; and of streaking on the back is highly variable; (2) E. p. olivascens, E. p. gossei and E. p. cata- some individuals of olivascens, especially from marcanus in the west (Paynter, 1970; Short, Bolivia (pers. obs.), have faintly streaked backs 1975; Contreras, 1980; Nores, 1986; Ridgely & approaching that of platensis, and worn indivi- Tudor, 1989). But as with many species of Ne- duals of platensis have essentially unstreaked otropical birds, their geographic variation and backs resembling olivascens (Wetmore, 1926; phylogenetic relationships remain poorly do- Hellmayr, 1938). cumented. In a survey of the Chaco avifauna, Gossei resembles olivascens, with which it is Short (1975) stated that platensis and olivas- most closely related, but averages larger in size, cens intergrade in the east-central Chaco. Ho- has a paler grey throat, chest and abdomen, du- wever, such intergradation has never been de- ller rectrices, and more greyish-green upper- monstrated. An analysis of 37 specimens of E. parts (Chubb, 1918; Wetmore, 1926; Contreras, platensis in Argentinian museums by Contreras 1980). Olrog (1963) synonymised gossei with (1980) remains the most detailed analysis of olivascens and subsequent authorities (e.g. geographic variation within the species, in Paynter, 1970; Short, 1975) adopted Olrog’s which only two specimens were described as treatment. However, Contreras (1980) exami- Ardeola 50(2), 2003, 223-235 GEOGRAPHIC VARIATION IN THE GREAT PAMPA-FINCH EMBERNAGRA PLATENSIS COMPLEX 225 FIG. 1.—Distribution of Embernagra p. platensis, E. p. olivascens and E. p. gossei specimens examined in this study, and specimen localities for E. p. catamarcanus from Nores (1986). Localities indicated by numerals re- fer to the most southeasterly specimen of E. p. olivascens (1) and the nearest, possibly hybrid specimen of E. p. platensis (2) reported by Contreras (1980). [Distribución de los individuos de Embernagra p. platensis y E. p. olivascens examinados en este estudio y lo- calidades de los individuos de E. p. catamarcanus obtenidos de Nores (1986). Las localidades marcadas con un número representan al individuo más sureste de E. p. olivacens (1) y al más cercano y posible híbrido de E. p. plantensis (2) indicado en Contreras (1980).] ned 13 specimens of gossei in Argentinian mu- The Great Pampa-Finch’s closest relative ap- seums and found them readily separable from pears to be the Pale-throated Serra-Finch (E. olivascens. longicauda), which is locally distributed in eas- A fourth subspecies, catamarcanus, was sub- tern Brazil (O’Brien, 1968;
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