Observations on the Andean-Patagonian Component of Southeastern Brazil's AvifaunaAuthor(s): Helmut SickSource: Ornithological Monographs, No. 36, Neotropical Ornithology (1985), pp. 233-237Published by: University of California Press for the American Ornithologists' UnionStable URL: http://www.jstor.org/stable/40168284 .

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OBSERVATIONS ON THE ANDEAN-PATAGONIAN COMPONENT OF SOUTHEASTERN BRAZIL'S AVIFAUNA

Helmut Sick Academia Brasileira de Ciencias, C. Postal 229, 20.000 Rio de Janeiro, Rio de Janeiro, Brazil

ABSTRACT. Several birds typical of southeastern Brazil's isolated and cold mountains have most probably been derived from Andean/Patagonian ancestors following Pleistocene glaciation. Typical species endemic to that area include Itatiaia Spinetail (Schizoeaca moreirae), Long-tailed Cinclodes (Cinclodes pabsti), and Mouse- colored Tapaculo (Scytalopus speluncae). Uniquely among Andean-Patagonian de- rivatives, Band-winged Nightjar (Caprimulgus longirostris) has also colonized low- lands, even reaching tropical, urban Rio de Janeiro. Secondary radiation has led to at least one species of Andean origin (Brasilia Tapaculo, Scytalopus novacapitalis) colonizing the central Brazil Plateau, apparently having followed the inland move- ment of coastal forests. Extension southward of Amazonian heat and northward of Patagonian aridity are advanced in explanation for the rupture of a formerly contin- uous band of vegetation linking the Andes and southeastern Brazil, with resulting isolation and speciation of landbirds.

RESUMEN. Varias aves tipicas de las montanas aisladas y frias del sureste de Brasil han derivado muy probablemente de ancestros andino-patagonicos luego de las glaciaciones del pleistocene Especies endemicas tipicas de esa area incluyen Schi- zoeaca moreirae, Cinclodes pabsti, y Scytalopus speluncae. La chotacabra sen-ana (Caprimulgus longirostris) es unica entre los descendientes andinopatag6nicos, ya que tambien ha colonizado las tierras bajas e incluso ha llegado a las areas urbanas y tropicales en Rio de Janeiro. La radiation secundaria ha hecho que al menos una especie de origen andino (Scytalopus novacapitalis) colonice la meseta central de Brasil, aparentemente habiendo seguido las extensiones tierra adentro del bosque atlantico. La extensidn del calor amaz6nico hacia el sur y de la aridez patag6nica hacia el norte brindan una hipdtesis para explicar la ruptura de una banda de vege- taci6n que en el pasado era continua y enlazaba los Andes con el sur de Brasil, dando como resultado la aislacion y especiacidn de aves terrestres.

The Itatiaia Highlands of southeastern Brazil (Fig. 1) were recognized early in the 20th century as having a peculiar flora composed largely of endemics whose closest relatives were in the Bolivian Andes (Dusen 1903). Shortly thereafter, Miranda Ribeiro (1906) extended the analysis to the area's fauna, confirming its Andean affinities. He noted among other obser- vations that the nearest relatives of certain catfishes (Siluroidei) and some birds were not in Brazil but in the Andes, a conclusion later reached for frogs and butterflies as well (Lutz 1951; Ebert 1960). In the present paper I offer observations on the taxonomic affinities of birds that occur generally in southeastern Brazil.

GEOLOGY AND CLIMATE

The Itatiaia Mountains appear to have been glaciated on several widely separated occasions (Ebert 1960), at least in the Wurm phase of the Pleistocene and possibly in pre-Devonian and Gondwanan times (Maack 1951), although not all experts concur that the area has ever been glaciated, and palynological evidence appears to be lacking (Oedman 1955; Beuerlen 1970). Nonetheless, most authors agree that U-shaped valleys 6-8 km long, typical glacial signs, are present, even though later water erosion has steepened their profiles to V-shapes. It is believed that vegetation covered the borders of the valleys, that the snow line was at ca. 1500 m, and that an ice-cap at ca. 2100-2600 m fed the glaciers (Ebert 1960).

Presumably, the same kinds of alternating warm-humid/cold-dry cycles that in the Andes facilitated the expansion and contraction of forests and other vegetation, with resulting spe- ciation and range changes in birds (cf. Haffer 1967, 1981; Vuilleumier 1969a), also occurred in southeastern Brazil.

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234 ORNITHOLOGICAL MONOGRAPHS NO. 36

Fig. 1 . Immigration of plants (horizontal arrow) from northern Argentina to southeast Brazil. Climate (heat, aridity; thick vertical arrows) and topography (stippled areas above 300 m) have been suggested as effecting disjunction of ranges between the Andes and southern Brazil. The square is the area shown in Figure 2. Adapted from Smith (1962).

REPRESENTATIVE TAXA Schizoeaca moreirae. Itatiaia Spinetail. This very local furnariid most clearly demonstrates

the faunal closeness of the Itatiaia Highlands and the Bolivian and Patagonian Andes. It is the most common species in the Chusquea bamboo above timberline (1800 m).

Hellmayr (1925) erected a new genus (Oreophylax) for this species, which is typically "orial" (=flora and fauna above timberline), in order to distinguish it from Synallaxis, but I believe that Vaurie (1971) was correct in allying it with the Andean thistletails {Schizoeaca). Its nest is a soft ellipsoid-shaped structure rather than an elongate one of rigid and spiny twigs (Sick 1970); this is typical of nests of Schizoeaca and in contrast to those of Synallaxis. Vuilleumier (1969b) described the nest of Schizoeaca harterti from Bolivia as very similar to that of moreirae. The vocalizations of the Itatiaia Spinetail are similar to those of tit-spinetails (Leptasthenura), also an Andean derivative, but not to those of Synallaxis.

Cinclodes pabsti. Long-tailed Cinclodes. This endemic, ground-dwelling furnariid is confined

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NEOTROPICAL ORNITHOLOGY 235

Fig. 2. Distribution of some Andean tapaculos (Rhinocryptidae) and furnariids (Furnariidae) in Brazil showing colonization of southeast and central Brazil.

to the high plateau (e.g., 1700 m, Campo dos Padres) of northern Rio Grande do Sul and adjacent Santa Catarina, the coldest region of Brazil (Sick 1973). Only recently described (Sick 1969), it is allopatric with its nearest relative, Bar- winged Cinclodes (C fuscus). This latter species is widespread throughout southern South America and in the Andes, although in winter it reaches lowland coastal areas in Rio Grande do Sul. Cinclodes is a well-defined genus, typical of the Andes and Patagonia, and occurs south to Tierra del Fuego.

Rhinocryptidae. Tapaculos. Tapaculos are widespread in the Andes and in Patagonia but have penetrated Brazil only in the southeastern mountains. Scytalopus speluncae, the Mouse- colored Tapaculo, is common on high, dense forest borders, even reaching peaks where, in Chusquea, it meets Schizoeaca moreirae. Most other tapaculos are scarce in southeastern Brazil.

Caprimulgus longirostris. Band- winged Nightjar. This typically Andean bird has invaded Brazil by way of the high, treeless parts of southeastern Brazil (Sick 1959). Unlike other montane derivatives, this species has continued its expansion in Brazil, moving into lower parts of Santa Catarina and Rio Grande do Sul. Most unexpectedly, and so far unique among Andean forms, Band-winged Nightjar has moved from the mountains to the tropical lowlands, and during the last 40 years has become a resident of the center city and suburbs of Rio de Janeiro (Sick 1963).

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236 ORNITHOLOGICAL MONOGRAPHS NO. 36

DISCUSSION The most plausible explanation for the origin of the peculiar montane flora and fauna of

southeastern Brazil is provided by Smith (1962), although a comprehensive biogeographic analysis of the area remains to be done. Smith argued that northern Argentina and southern Brazil were once linked floristically at ca. 27°S, the ranges of Andean plant genera having been constricted by the southward push of Amazonian heat (the 26° summer isotherm) and the northward push of Patagonian aridity (Fig. 1). This link has been broken many times. For example, the southeastern Brazil Araucaria forest now seems clearly an isolated relict of the Andean flora (Rambo 1951). Before rupture, however, there apparently was a band of con- tinuous vegetation extending the 1 500 to 2000 km distance between the Andes and south- eastern Brazil, and this served as the colonization corridor from the Andes. It is likely, however, that Andean colonizers such as the Itatiaia Spinetail, expanded their geographical ranges by jumping between isolated mountains (cf. Mayr and Phelps 1967 for a Venezuelan analogue).

Not all movement was toward southeastern Brazil from the Andes, however. An isolated rhinocryptid, the Brasilia Tapaculo (Scytalopus novacapitalis), has colonized the Brasilia Pla- teau in Central Brazil (1000 m). Floristic analysis of the forest where this bird occurs suggests that it followed plants invading that plateau from the Atlantic forest; typical of many coastal plants there are edible palm (Euterpe edulis) and the large fern Blechnum brasiliensis. Other Brazilian coastal birds, including furnariids (Sharp-tailed Stream-creeper, Lochmias nematura) and tyrants (Southern Antpipit, Corythopsis delalandi), have followed the same route to the Brasilia Plateau (Fig. 2).

The unravelling of zoogeographic patterns in Brazil, especially in the southeast, has barely begun; the field is ripe for investigation.

ACKNOWLEDGMENTS I thank C. T. Rizzini for information about Atlantic coastal plants that have invaded central

Brazil, and J. Haffer for suggestions on an earlier draft of this paper. P. A. Buckley, L. L. Short, and M. S. Foster assisted greatly in putting the manuscript into final form.

LITERATURE CITED

Beuerlen, K. 1 970. Geologie von Brasilien. Beitrag zur regionalen Geologie der Erde, Vol. 9. Borntrager, Berlin.

Dusen, D. K. H. 1903. Sur la flora de la Serra do Itatiaia. Arq. Mus. Nac. Rio de J. 13:1-1 19. Ebert, H. 1960. Novas observacoes sobre a glaciacao pleistocenica na serra do Itatiaia. An. Acad. Bras.

Cienc. 32:51-73. Haffer, J. 1967. Speciation in Colombian forest birds west of the Andes. Am. Mus. Novit. No. 2294. Haffer, J. 1981. Aspects of neotropical bird speciation during the Cenozoic. Pp. 37 1 -394, In G. Nelson

and D. E. Rosen (eds.), Vicariance Biogeography: A Critique. Columbia University Press, New York.

Hellmayr, C. H. 1925. Catalogue of Birds of the Americas. Pt. IV. Field Mus. Nat. Hist. Publ. 234, Zool. Ser. Vol. 13.

Lutz, B. 1951. Nota pre\ia sobre alguns anflbios anuros do Alto Itatiaia. Hospital: maio. Maack, R. 1951. Vestigios pre-devonianos de glacicao e a sequencia de camadas devonianos no Estado

do Parana. Ara. Biol. Tecnol. (Curitiba) V-VI O6):197-230. Mayr, E., and W. H. Phelps, Jr. 1967. The origin of the bird fauna of South Venezuelan highlands.

Bull. Am. Mus. Nat. Hist. 36:269-327. Oedman, H. O. 1955. On the presumed glaciation in the Itatiaia mountain, Brazil. Engenh. Mineral

Metal. Rio de J. 21:107-108. Rambo, B. 1951. O elemento andino no Pinhal Riograndesse. An. Bot. Herb. Barbosa Rodrigues 3:

7-39. Ribeiro, A. Miranda. 1906. Vertebrados do Itatiaia (Peixes, Serpentes, Saurios, Aves e Mamiferos).

Arq. Mus. Nac. Rio de J. 13:165-190. Sick, H. 1959. O redescobrimento no Brasil do Bacurau Caprimulgus longirostris. Bonaparte (Capri-

mulgidae, Aves). Bol. Mus. Nac, Rio de J. Zool. 204:1-15. Sick, H. 1963. O bacurau Caprimulgus longirostris e outras aves noturnas do Estado da Guanabara.

Vellozia I, 3:107-116. Sick, H. 1 969. Uber einige Topfervogel (Furnariidae) aus Rio Grande do Sul. Brasilien, mit Beschreibung

eines neuen Cinclodes. Beitr. Neotrop. Fauna 6:63-79. Sick, H. 1970. Der Strohschwanz, Oreophylax moreirae, andiner Furnariide Sudostbrasiliens. Bonn.

Zool. Beitr. 21:251-268.

This content downloaded from 130.60.206.43 on Thu, 8 Aug 2013 09:01:57 AMAll use subject to JSTOR Terms and Conditions

NEOTROPICAL ORNITHOLOGY 237

Sick, H. 1973. Nova Contribuicao ao conhecimento do Cinclodes pabsti Sick, 1969. Rev. Bras. Biol. 33:109-117.

Smith, L. B. 1962. Origins of the flora of Southern Brazil. Contrib. U.S. Natl. Herb. 35:215-249. Vaurie, C. 1971. Classification of the Ovenbirds (Furnariidae). Witherby, London. Vuilleumier, F. 1969a. Pleistocene speciation in birds living in the high Andes. Nature 223:1179-

1180. Vuilleumier, F. 1969b. Field notes on some birds from the Bolivian Andes. Ibis 1 1 1:599-608.

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