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Bird Speciation in Subtropical South America in Relation to Forest Expansion and Retraction

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Bird Speciation in Subtropical South America in Relation to Forest Expansion and Retraction The Auk 109(2):346-357, 1992 BIRD SPECIATION IN SUBTROPICAL SOUTH AMERICA IN RELATION TO FOREST EXPANSION AND RETRACTION MANUEL NORES ConsejoNacional de InvestigacionesCientfficas y Tgcnicas,Centro de ZoologfaAplicada, Casillade Correo122, 5000 Cdrdoba,Argentina ABS?RACT.--Ianalyzed two patternsof bird distribution in subtropicalSouth America. The first was the disjunctdistribution of pairs of speciesand subspeciesbetween the southern Yungas and the Paranenseforests, which are separatedby 700 .kin of xerophytic Chaco woodland.Forest birds penetrateup to 200 km into the Chacoalong gallery forests,but do not crossthe rest of the Chaco, which constitutesan effective barrier. The secondpattern involved several zones of secondarycontact located in the Chaco lowland, where several woodlandand grasslandspecies and subspeciesinteract. I concludethat both bird-distribution patternswere producedby forest expansionsalong the Bermejoand Pilcomayorivers that connectedthe southernYungas to the Paranenseregion and interrupted the arid vegetation in the center of the Chaco.Vegetational fluctuations probably occurredseveral times during the Quaternary and producedphenotypic differentiationof sistertaxa, both in the now- disjunctforests and in the currently continuousChaco. Received 26 June1990, accepted10 January 1992. THE FORESTand woodland area located around cluded that the Paranense forest of Misiones the Tropic of Capricorn in South America is was once continuous, through Paraguay, with biogeographicallycomplex. The region shows the Yungasof Tucum•n-Bolivia.However, O1- two distinct features: (1) the southernmost ex- rog (1984) reversedthis idea when he presented tension of tropical South American forests(Pa- a present-day ornithogeographic map of Ar- ranenseand southernYungas), with a high de- gentina in which he connectedboth regionsby gree of avian endemism (Cracraft 1985, Nores a continuousgallery forestalong the Pilcomayo 1989); and (2) the central Chaco woodland, with River. With regard to the secondpattern, Short low avian endemism (Short 1975). Relatively (1975) identified the area of the Bermejo and little is known about the climatic history of the Pilcomayorivers as a zone that marks a disjuc- area and the effects of Quaternary climatic tion in bird distribution in South America. changeson the dispersaland differentiation of Research carried out in South America over bird species.Understanding shifts in the re- the last 30 years has shown that, during the gion's vegetation during the Quaternary could Quaternary,semiarid conditions alternated with help to provide clues about events in the rest humid phases.These cyclescaused shrinkages of South America (B. Simpson,pers. comm.). of formerly widespread forests into humid Two separatepatterns in the distribution of pockets(refuges), followed by their subsequent the avifauna probably resultedfrom large-scale reexpansion (Garner 1959, Bigarella and An- vegetational dynamicsduring the Quaternary. drade 1965, Haffer 1969, 1974,Mayr 1969,Mayr First, there are several species and subspecies and O'Hara 1986, Vanzolini and Williams 1970, pairs that have disjunct populations in the Vuilleumier 1971, Prance 1974, Van der Ham- southern Yungas and Paranenseforests, sepa- men 1974, 1982,Tricart 1974, 1975, Simpsonand rated by 700 km of xerophyticChaco woodland Haffer 1978, Absy 1979, 1982, Schubert 1988). (Fig. 1). Second,a zone of disjunction occursin Paleoecologicaldata and present distribution the Chaco, along the Bermejo and Pilcomayo patterns of animals and plants suggest that rivers, where the distribution of several species during interglacial phasesthere were periods and subspeciespairs of woodland and grassland when temperature and humidity were higher birds meet (Fig. 1). and forestsmore widely distributedthan today The first pattern was previously pointed out (Groeber 1936, Smith 1962, Vanzolini 1968, 1974, by Olrog (1963), who noted that several bird 1981, Haffer 1974, 1985, Van der Hammen 1974, speciesof the southern Yungasalso occur in the 1982,1983, Fitzpatrick 1980,Markgraf 1985,No- Paranenseforest, or are representedthere by res and Cerana 1990). Some authors consider anothersubspecies. Later, he (Olrog 1979a)con- the evidence of aridity during the glacial pe- 346 April1992] BirdSpeciation inSouth America 347 Jill • IllIll[[ I• • ß IIIIIIll I1•_ ß ß JillIfil ß IIIIIlll ß • IIIIflll IIIIIIII b • Amazonian • Paranense L• Serra do Mar •1Yungas and Andes eastern slopes O 1OOO 2000 Km Fig. ! Biogeographicprovinces of South America, excluding the Pacific coast,most of the Andes, and southernand centralArgentina (adaptedfrom Cabreraand Willink 1973).Three regionsthat form the "arid diagonal" (Vanzolini 1974, Fitzpatrick 1980): (a) Chaco, showing the area of interaction of woodland and grasslandbirds (box with oblique lines); (b) Cerrado;and (c) Caatinga. From south to north, the breaks in the black band (representingYungas and Andes slopes)mark the limit of the southernYungas, northern Yungas,and easternslopes of the northern Andes. riods to be insufficient to indicate that the fauna are relatedto Quaternaryclimatic changes wet tropical lowlands changed substantially and if they have common histories, I studied (Connors 1986, Colinvaux 1989). Endlet (1977, the distributionof forestbirds and plantsin the 1982a,b) pointed out that current patterns of Chaco lowland. I then comparedthe levels of distribution are consistentwith geographical phenotypic differentiation of the birds that in- divergenceand adaptationsto present-daycon- habit the southernYungas with thoseoccurring ditions.Consequently, it is not necessaryto pos- in the Paranense region, and examined the tulate refuges to explain the observed distri- ranges of woodland and grasslandbirds that butions. currently interact in the center of the Chaco. To determine if the two patterns of the avi- Finally, I relatedthe extentof phenotypicdif- 348 MANUEL NORES [Auk, Vol. 109 -[] Yungasforest ] Paranenseforest ..,[]Galleryforest I • I I I 0 100 200 Km Fig. 2. Vegetationdistribution in studyarea. Solid dots indicate location of dry riverbedswith forest patches.Triangles indicate presence of foresttrees related to present-daywatercourses. (a, b and c) The most extremelocations of galleryforests in Chacolowlands. (d) Paranenseforest wedge reaching Paraguay River. Galleryforests located east of Paraguayand Paran•rivers, in floodedarea, are not represented(from Notes 1989). ferentiation of these birds with vegetation proximately700 km wide,this area slopes gently from changesthat apparently occurredin South west to east,and extendsinto Argentina,Bolivia and America during the Pleistoceneand Holocene. Paraguay.Rainfall increasesmarkedly from 500 mm in the west to 1,300 mm in the east. The Chaco, to- STUDYAREA getherwith the "Cerrado"and "Caatinga"in Brazil, formsthe "arid diagonal"(Vanzolini 1974, Fitzpatrick The studyarea is subtropicalSouth America,spe- 1980; Fig. 1). cificallythe region drainedby the Bermejo,Pilco- The third componentis the forest and "campos" mayo,Paraguay, and Paran•rivers (Fig. 2). Fromwest (Paranenseregion) of southeasternBrazil, eastern to east it includes three components. Paraguayand northeasternArgentina, around the up- The first component is the forests on the eastern per coursesof the Paran•and Uruguayrivers. Rainfall slopesand foothills of the Andes,and on the Sierras decreases towards the west from 1,700 mm to 1,300 Pampeanassouth of 18øS(southern Yungas). These mm. As a result,the vegetationbecomes increasingly forests stretch from Santa Cruz and Cochabamba de- impoverishedand it ends in a narrow wedge on the partmentsin Bolivia to CatamarcaProvince in Ar- ParaguayRiver (Fig. 2, point d). Detailed descriptions gentina.I definedthree altitudinal belts: (a) basalfor- of the study area are given in Hauman et al. (1947), est between 350 m and 600 m; (b) cloud forest between Ad•rnoli et al. (1972),Short (1975), and Cabrera(1976). 600 and 1,500 m; and (c) montane deciduousforest between 1,400 and 2,500 m. MATERIALS AND METHODS The secondcomponent includes the Chacolowland drained by the Bermejoand Pilcomayorivers and Avifauna.--I carried out field work from 1973 to other small tributaries of the ParaguayRiver. Ap- 1988 in the forests and Chaco woodland of northern April1992] BirdSpeciation inSouth America 349 Argentina, southern Bolivia, eastern Paraguay, and river and stream banks. Outside the flooded ar- southern Brazil (Nores 1989). From 1986 to 1988 I eas,the gallery forestswiden considerably.On concentratedon areas of particular biogeographic the Pilcomayoand Bermejorivers, the gallery complexity. These included river and stream banks, forests extend about 200 km to the west, reach- forest patcheson dry riverbeds, Paranenseprairies, and the interaction zone in the Chaco lowland. I cov- ing Fortln SargentoPrimero Leyes (24ø30'S, ered Argentine National Highway 81, which crosses 59ø23'W)and Puerto Bermejo(25ø39'S, 60ø08'W; the Chaco lowland from east to west, and most of the see Fig. 2, points a and b). On the streams,the secondaryroads that give accessto the Bermejoand gallery forestsare somewhatshorter. Pilcomayorivers and other small water courses(Fig. In the central and western Chaco rainfall is 2). At each study site, I identified all bird species lower and forest trees occuras small patches(of presentthrough a combinationof visualobservations, one to five species)on dry riverbeds,or in small songidentification, song playback, and capturewith groups(of one to three species)on river banks. mist nets.
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