Revista Brasileira de Ornitologia 14 (3) 197-238 setembro de 2006 The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin

Luciano Nicolás Naka1,4,5, Mario Cohn-Haft1, Francisco Mallet-Rodrigues2, Marcos Pérsio Dantas Santos3, and Marcela de Fátima Torres1

1 Instituto Nacional de Pesquisas da Amazônia (INPA), Coleções Zoológicas e Coordenação de Pesquisas em Ecologia, C.P. 478 (69011-970), Manaus, Amazonas, Brazil.

2 Laboratório de Ornitologia, Departamento de Zoologia, Instituto de Biologia, UFRJ (21941-590), Rio de Janeiro, Rio de Janeiro, Brazil.

3 Universidade Federal do Piauí, Departamento de Biologia. (64049-550), Teresina, Piauí, Brazil.

4 Current Address: Museum of Natural Science, 119 Foster Hall, Louisiana State University, Baton Rouge, LA 70803, USA.

5 Corresponding author: Luciano Nicolas Naka, e-mail: [email protected]

Recebido em 30 de dezembro de 2005; aceito em 22 de maio de 2006

RESUMO. A Avifauna do Estado de Roraima: distribuição e biogeografi a na bacia do Rio Branco. O Estado de Roraima representa uma impor- tante unidade biogeográfi ca, que inclui toda a bacia do Rio Branco, um dos principais rios Amazônicos. Roraima é uma área de alta diversidade, onde fl orestas de terra fi rme, fl orestas alagadas, campinas, savanas, bosques secos e em galeria, e vegetação associada com os tepuis ocorrem em uma área relativamente pequena para padrões Amazônicos (225.000 km2). Neste trabalho, apresentamos a primeira compilação em quarenta anos, das 741 espécies de aves registradas para o Estado de Roraima, 38 das quais representam as primeiras menções para a região. Os dados foram compilados a partir de informações obtidas durante nossas expedicões de campo (512 espécies), complementadas por informacões obtidas de material de museu e da literatura. Apresentamos uma lista com 15 espécies citadas para Roraima no passado, que não consideramos como registros válidos e achamos que sua presença em Roraima ainda deve ser confi rmada, e uma outra lista com 69 espécies que acreditamos venham ser registradas em Roraima no futuro. Apresentamos também detalhes de alguns dos nossos registros de maior interesse, incluíndo o primeiro espécime para o Brasil de Atalotriccus pilaris, e a segunda localidade para o Brasil de Myrmeciza disjuncta. Uma análise dos padrões de distribuição da avifauna, em relação aos diferentes ambientes, revelou que a heterogeneidade ambiental (diversidade beta) é responsável por 60% da diversidade de Roraima, enquanto que o endemismo regional (diversidade gamma) responde por quase um terço das espécies de aves exclusivas de fl orestas de terra fi rme. Apresentamos também uma análise dos padrões biogeográfi cos das aves em Roraima, em relação aos diferente hábitats. E fi nalmente, discutimos os problemas de conservação desta região que encontra-se ameaçada pela crescente expansão de monoculturas (soja, acácias, arroz) principalmente nas áreas de savanas, fl orestas de galeria, e bosques secos, onde habitam algumas das espécies de aves endêmicas e mais ameaçadas da região.

PALAVRAS-CHAVE: Amazônia, Roraima, Rio Branco, avifauna, distribuição, biogeografi a.

ABSTRACT. The Brazilian state of Roraima represents a meaningful biogeographical unit, enclosing the entire basin of the Rio Branco, a major river in the Amazon basin. Roraima is also an area of high habitat diversity where terra fi rme forest, seasonally fl ooded forest, white-sand forest, savanna, gallery and dry forest, and the vegetation associated with the tepuis, all occur within a relatively small area (225,000km2) by Amazonian standards. We provide the fi rst comprehensive compilation of the avifauna of the state in forty years. We integrated the results of our own fi eld work, where we recorded 512 bird species, with data gathered from museum specimens and from published sources. We present a list of 741 bird species for the state of Roraima, 38 of which are mentioned for the fi rst time for the state. We also provide a list of 15 species previously cited for Roraima that we consider hypothetical, and predict the presence of another 69 species that we consider likely to be recorded in the state with further fi eld work. We also present details on some of our most interesting records, which include the fi rst Brazilian specimen of Atalotriccus pilaris, and the second locality for Brazil of Myrmeciza disjuncta. We conclude that habitat heterogeneity (beta-diversity) is responsible for 60% of the avian diversity of Roraima, whereas regional endemism (gamma-diversity) accounts for about one third of the bird species occurring exclusively in terra fi rme forests. We analyze the biogeographical patterns of avian distributions within the state, in relation to the different habitats present in Roraima, and discuss some of the main conservation problems in the state of Roraima, driven by the expansion of monocultures such as soybean, acacia, and rice plantations which are putting at risk large expanses of savannas, wetlands, dry and gallery forests.

KEY WORDS: Amazonia, Roraima, Rio Branco, avifauna, distribution, biogeography.

At the northern edge of Amazonia, the Brazilian state of Ro- Amazonian standards. Avian diversity in Roraima refl ects this raima represents one of the most heterogeneous regions in high habitat heterogeneity, as each major habitat contributes the Neotropics. Distinct habitats, including terra fi rme forest, with unique species. Regional diversity is also increased by seasonally fl ooded forest (várzea and igapó), white-sand for- the fact that different areas within Roraima are, to varying de- est (campina and campinarana), savanna, gallery and dry for- grees, biogeographically distinct; at least three proposed areas est, and various types of montane forests, all occur in an area of endemism extend into the state (Cracraft 1985). roughly the size of the United Kingdom, relatively small by Unlike most political units, the state of Roraima represents 198 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres

a meaningful biogeographical unit, enclosing the entire basin and some of its tributaries, the white-sand forest (campina and of the Rio Branco, the most important tributary of the Rio campinarana) in central Roraima, and the terra fi rme forest Negro, which itself is the second largest tributary of the Ama- east of the Rio Branco. We integrated our results with data ob- zon River (Goulding et al. 2003). The Rio Branco basin is tained independently by MPDS, and with data from previous unique in Amazonia because its main river, the Rio Branco, is workers available from published sources and museums (data a white-water river surrounded by black-water rivers, which compiled by FMR). differ in the amount of sediments, type of vegetation, and the In this paper, we provide a brief history of ornithologi- structure of the plant community (Worbes 1997). These dif- cal research in Roraima, present our fi eld records, including ferences between the two should also infl uence the abundance 38 bird species recorded for the fi rst time in Roraima, and and composition of the bird species that inhabit each type of provide the fi rst comprehensive compilation of the avifauna river. Therefore, a study of the avifauna of Roraima not only of the state since Pinto (1966). We also make predictions on sheds light on one of the least-studied areas of Brazilian Ama- the species likely to be recorded in the state with further fi eld zonia (Oren and Albuquerque 1991), but can also improve our work and describe the broad patterns of avian distributions understanding of Amazonian avian biogeography, generating within the Rio Branco basin in relation to the different habi- insights on ecological and geographical patterns. tats, discussing the implications of these patterns in a biogeo- Until the mid-20th century, most of the ornithological mate- graphical perspective and giving special attention to the role rial available for Roraima was that obtained by Johann Nat- of the Rio Branco as a biogeographical barrier. terer in the early 1800s. During the second half of the 20th cen- tury, several ornithologists visited or analyzed bird specimens HISTORY OF ORNITHOLOGICAL STUDIES from the state, but these studies concentrated on a few areas, IN RORAIMA including savannas and forests near Boa Vista, the upper Rio Branco, a handful of tepuis on the Brazilian-Venezuelan bor- The study of the avifauna of Roraima began with the ex- der, and the Maracá Ecological Station (see History of orni- plorations of Natterer almost two centuries ago. Since then, a thological studies in Roraima, below). Entire ecosystems and number of ornithologists have visited the region, mostly dur- biogeographic areas remained virtually unexplored until now. ing the last 50 years, defi ning important periods of high pro- In 2001, three of us (LNN, MCH, and MFT) initiated a ductivity in terms of accumulation of ornithological data, and study of the avifauna of Roraima and made fi ve fi eld trips periods of relative stagnation (Figure 1). to the state. We explored poorly sampled habitats and previ- The fi rst naturalist to visit the Rio Branco basin appears ously neglected biogeographical areas, such as the seasonally to have been Alexandre Rodrigues Ferreira, a Brazilian ex- fl ooded forest (várzea and igapó) along the lower Rio Branco plorer sent by the Portuguese crown in 1786 to investigate

Figure 1. Cumulative number of bird species known from Roraima since the fi rst related publication in 1850. Decades represent time of publication of lists of specimens or fi eld records. Arrows indicate the explorers or ornithologists responsible for most of the additions during that decade. The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 199

the economic potential of the region (Barbosa and Ferreira During the following years, most of the attention given to 1997). Ferreira sailed the Rio Branco and some of its tributar- Roraima came from Venezuela, mostly from explorers inter- ies, collecting plants and animals. These valuable specimens ested in the endemic avifauna associated with the tepuis. Félix were taken from Portugal by Saint-Hilaire in 1808 during the Cardona, a Venezuelan Captain, explored the Brazilian-Venezu- Napoleonic invasions and were purportedly deposited in the elan border, collecting several bird species, four of which were Paris Museum (Pinto 1966), but none of the material is pres- new to Brazil (Phelps and Phelps 1948). William H. Phelps ent in Paris today. and William H. Phelps, Jr. (1962) surveyed another Brazilian Specimens taken by Natterer, who navigated the Rio Bran- tepui, Cerro Uei-Tepui, adding 49 new species to Roraima and co during 10 months in 1831-1832, formed the basis of study Brazil. The study of the tepuis cotinued by José Hidasi, who of the avifauna of Roraima for many years. Natterer explored visited Serra Parima in 1962, collecting almost 30 bird species the upper Rio Branco and some tributaries, and its associated (Novaes 1965). However, with a later revision of the interna- savannas and forests, where he collected 157 bird species, tional limits of Brazil and Venezuela, the locality sampled by held at the Naturhistoriches Museum of Vienna (Pelzeln 1956, Hidasi (Posto Parima B) was confi rmed to be in Venezuela. 1859, 1861, 1862, 1863, 1868-1871). Two decades later, Robert Dickerman and William H. Phelps, The ornithologist Newton Dexter, a member of an expedition Jr. (1982), made an expedition to Cerro Urutani, collecting 35 funded by Nathaniel Thayer from Harvard University, visited the bird species, many of which were new to Roraima. This was Rio Branco and the Rio Negro in 1865 and 1866 (Barbosa and the last ornithological expedition to any tepui in Roraima, and Ferreira 1997). Unfortunately, the material collected by Dexter we believe that several other species are likely to be recorded (held at the Museum of Comparative Zoology in Cambridge) is in the tepuis with further fi eld work (see Discussion). of little use, as specimens lack precise locality data. In 1959, the Museu Paraense Emílio Goeldi (MPEG) or- In 1912, M. P. Anderson visited Roraima, where he ex- ganized an ornithological expedition to the Rio Mucajaí and plored the savannas around Boa Vista and the forests of Serra to Boa Vista, led by C. Carvalho and M. Amaral. However, it da Lua, Serra Grande, and Conceição, collecting over 500 wasn’t until 1966 that a comprehensive study of the avifauna specimens of almost 200 bird species, presently held at the of Roraima was published, improved signifi cantly by an ex- Field Museum of Natural History (FMNH) and analyzed by pedition sponsored by the Instituto Nacional de Pesquisas da Cory (1918, 1919, 1920), Cory and Hellmayr (1924, 1925, Amazônia (INPA) and the Museu de Zoologia da Universi- 1927), Hellmayr (1929, 1934, 1935, 1936, 1937, 1938), and dade de São Paulo (MZUSP) to the Rio Mucajaí. During this Hellmayr and Conover (1942, 1948a, 1948b, 1949). A decade expedition in 1962, Olivério Pinto collected more than 250 later, George C. Shattuck published the results of the 1924- bird species (held at the MZUSP), and he later compiled the 1925 Hamilton Rice expedition, where he made brief com- ornithological knowledge available until then on the avifauna ments on the avifauna encountered during the trip, which in- of Roraima (Pinto 1966). cluded the villages of Boa Esperança and Vista Alegre along Emílio Dente, a professional collector who came to Rorai- the Uraricoera River (Shattuck 1926). ma to join Pinto, decided to stay in Roraima after that trip and In 1927, an expedition led by George H. Tate, sponsored collected more material from the Rio Mucajaí. Dente obtained by the American Museum of Natural History (AMNH), and over 1200 specimens, most of which are held at the Natural His- another one led by General Mariano Rondon (supported by tory Museum of Los Angeles County (LACMNH), but also in the Brazilian government), independently explored the sur- at least seven other institutions, including the MPEG, MZUSP, roundings of Mt. Roraima, collecting a small number of bird the FMNH; the Museum of Comparative Zoology Harvard specimens from the savannas of northeastern Roraima. Much (MCZ); the Smithsonian National Museum of Natural History of the material collected by the AMNH expedition was includ- (USNM); the Academy of Natural Sciences (ANSP); and the ed in studies by Zimmer (1933, 1936, 1937a, 1937b, 1938, University of Michigan Museum of Zoology (UMMZ). 1939a, 1939b, 1940, 1941a, 1941b, 1941c, 1942a, 1942b, During the 1980s and 1990s, more intensive fi eldwork, 1943a, 1943b, 1944), whereas the material from the Brazilian including tape recording of vocalizations, behavioral observa- expedition (held at the Museu Nacional [MNRJ]) was listed tions, and use of newly available fi eld guides as identifi cation by Miranda-Ribeiro (1929). aids, was initiated at a few sites. In particular, the island of In January 1939, Albert Pinkus, a professional collec- Maracá was the center of intensive studies (Moskovits et al. tor from New York, and P.S. Peberdy from the Museum de 1985, Silva and Oren 1990, Stotz 1997, Silva 1998), but other Georgetown, Guyana, made an expedition to Mt. Roraima. areas were also explored along an improved network of roads Although most of the trip was devoted to Guyana, the explor- (Borges 1994, Forrester 1995, Zimmer et al. 1997). ers spent a few days in Brazil, reaching the valley of the Rio Of special importance was the work of D. Stotz, who cov- Cotingo (Barbosa and Ferreira 1997), obtaining an important ered a range of habitats in northern Roraima. In 1987, he vis- bird collection, most of which was deposited at the Museum ited Maracá, Pacaraima, Apiaú, and the upper Rio Branco, col- of Georgetown. Part of this collection was later purchased lecting nearly 150 bird species, held at the FMNH and MZUSP by the Phelps Collection of Venezuela, where more than 140 (Stotz 1997). In 1992, he explored areas around Bonfi m and specimens collected in Brazil are held. Cantá, collecting over 350 bird specimens, held at the FMNH. 200 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres

by the presence of rocky outcrops known as ‘inselbergs’, such as Serra da Lua, Serra de Mucajaí, and Serra Grande, which reach up to a 1000 m in elevation (Ab’Saber 1997). The climate of lowland Roraima is warm (with annual mean temperatures of 26-27ºC) but varies considerably at higher el- evations (Eden and McGregor 1998). As a general rule, annual rainfall increases from the northeast (1,100-1,400 mm/year) to the southwest, where rainfall is more abundant (2,000-2,300 mm/year) and less seasonal (Barbosa 1997) (Figure 3). Roraima can be divided in two main ecological domains: the savanna/dry forest domain and the humid forest or Ama- zonian domain; these are separated approximately at the 1700 mm/yr rainfall isocline, where there is a broad belt of transi- tion between the two (Figure 3). As a general rule, savannas occur in areas with annual rainfall below 1700 mm, usually concentrated during a period of 100-130 days (Barbosa 1997). Within the savannas, some areas are covered by dry and gal- lery forests, whereas poorly drained areas support stands of Figure 2. Map of the state of Roraima showing surrounding areas, Moriche palms (Mauritia fl exuosa), locally known as buri- major villages and towns, most important rivers, and study locali- tizais. Humid forests generally occur in areas with more than ties. Gray represents neighboring countries, light gray other Brazi- 2,000 mm/year, but include open vegetation areas known as lian states. Numbers represent the following localities: 1= Conceição campinas on sandy soils (see Habitats). do Maú, 2= Serra da Onça, 3= Serra da Malacacheta, 4= Serra do Tracajá, 5= Serra Grande, 6= Vila União, 7= Caracaraí Ecological Station, 8= Viruá National Park, 9= Niquiá Ecological Station, 10= HABITATS islands of Carneiro, 11= island of Inajatuba, 12= São Luiz, 13= São João da Baliza, 14= Caroebe, 15= Samaúma, 16= island of Cota, 17= Terra fi rme forest. Dominant habitat type in Amazonia. Remanso, 18= Floresta. Map based on data from Ministério do Meio In contrast to fl ooded forests, terra fi rme or upland forests, Ambiente (2001). never fl ood and generally occur at low elevations (below 600 m), in association with latosols and red-yellow podzols, well In the mid 1990s Fernando Pacheco (1995a) made a brief drained and poor in nutrients (Pires and Prance 1985). Terra visit to the mouth of the Rio Branco, adding the fi rst published fi rme forests are rich in plant species and have high tree bio- bird records from the lower section of the river. Finally, in the mass; the canopy reaches 20-25 m, with emergent trees of 21st century, Trolle and Walther (2004) made observations along 35-40 m. There is considerable variation in plant species rich- the Rio Jauaperí, and one of us (MPDS) made intensive col- ness described, with forests at Maracá, northern Roraima, be- lections near Caracaraí, Alto Alegre, and Boa Vista, obtaining ing less diverse than similar habitats elsewhere in Amazonia almost 500 specimens (held at the MPEG), some of which rep- (Nascimento 1997). However, plant species richness in other resented the fi rst records for the state (Santos 2004). Our fi eld forested areas within Roraima has not been studied in detail. work began in 2001, and is presented in the following sections. On and around rocky outcrops within the range of terra fi rme forest in southern and central Roraima (such as near Vila União), the vegetation resembles that of the dry forests further STUDY AREA AND METHODS north, containing large cacti and bromeliads and lower-stature THE STATE OF RORAIMA forest within a mosaic of more typical tall, moist forest away from rocks. The Brazilian state of Roraima, with an area of 225,116 Várzea. Typical fl ooded forest and other successional hab- km2, is located in northwestern Brazil, fl anked by Guyana to itats occurring along rivers and their fl oodplains in Amazonia. the east, and Venezuela to the north and west, enclosing the en- Várzeas are seasonally fl ooded by “white-water” rivers, which tire basin of the Rio Branco (Figure 2). Geologically, Roraima are rich in suspended sediments and have a muddy appear- lies on the ancient soils of the Guianan Shield, and most areas ance (Prance 1979, Goulding et al. 2003). Within Roraima, within the state are relatively fl at and low, especially in cen- várzeas are associated exclusively with the lower Rio Branco tral and southern Roraima, where low-lying depositional areas (Figure 3). The sediments carried by the river supply enough form the Rio Branco fl oodplains (Eden and McGregor 1998). In nutrients to support large trees and a well-developed canopy, northern and western Roraima, elevated terrain, ranging from reaching 20-25 m; the understory is generally open and poorly 1000 to almost 3000 m at Mt. Roraima, forms the divide be- developed. On white-water river islands, tree species diversity tween the Orinoco and Amazon basins. Northeastern Roraima and complexity of vertical structure increase progressively is covered by large expanses of fl at savannas, regularly broken through time (Robinson and Terborgh 1997), creating a se- The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 201

crowns, and thin and twisted branches (Anderson 1981). The fl oor is often covered by bromeliads, epiphytes, and lichens, whereas mosses are common on trunks and branches. Campi- nas are shrubbier and lower in stature (4-5 m) than campi- naranas and often contain open areas dominated by sedges (Cyperaceae) and can be devoid of woody vegetation. Despite their relatively low plant species diversity, campinas and campinaranas have a highly specialized plant community that includes several endemic taxa (Pires and Prance 1985). Savannas. The savannas of Roraima are part of the “Rio Branco-Rupununi” formation, shared by Brazil and Guyana, covering over 50,000 km2, and representing the largest con- tinuous Amazonian savanna (Barbosa et al. 2005). Locally known as lavrados, the savannas of northeastern Roraima oc- cur on well-drained soils and where rainfall is seasonal (con- centrated between May and August) and less abundant (below 1,700 mm/year). These savannas are not uniform and include several vegetation types (Eden and McGregor 1998, Barbosa Figure 3. Map of the state of Roraima and neighboring areas in Bra- and Miranda 2005), being fl atter to the south and more hilly zil, with main habitats and rainfall isoclines. Map and distribution to the north, where the landscape is covered by volcanic rocks of habitats based on data from Ministério do Meio Ambiente (2001); rainfall isoclines taken from Barbosa (1997). (Pires and Prance 1985). Savannas are xerophytic formations dominated by herbaceous plants and scattered low trees of ries of successional stages that range from grassy sandbars to only a few fi re-resistant species, such as Curatella americana river-edge forest (dominated by Cecropia), to mature várzea and Byrsonima crassifolia. The main plant families in terms forest, which has the highest number of tree species (Remsen of number of species are Poaceae, Fabaceae, Cyperaceae, Ru- and Parker 1985). biaceae, Caesalpinaceae, Melastomataceae, and Polygalaceae Igapó. Term generally applied to forests seasonally fl ood- (Miranda and Absy 1997). Although the savannas of Roraima ed by “black-water” and nutrient-poor rivers (Prance 1979). resemble the cerrados of central Brazil, they differ in their fl o- These rivers have high acidity, low sediment loads and drain ristic composition (Ratter and Dargie 1992), being more close- areas of lowland forest or campinas (see below). We distin- ly related to the vegetation of the Gran Sabana in Venezuela guish two types of igapó: tall, which resembles mature várzea (Berry et al. 1995). Within the savannas, poorly drained areas forest, and low, sometimes referred to as chavascal, which has support stands of Moriche palms and grassy wetlands, whereas a dense growth of relatively thin trunks and a much more uni- humid areas on better-drained and seasonally drier soils sup- form canopy height, with physiognomy and fl oristic composi- port gallery and dry forests (see below). tion resembling those of campinarana (Anderson 1981). Low Gallery forests. Riparian woodlands associated with rivers igapó is associated to sandy soils and has fewer tree species fl owing through the savannas of northeastern Roraima, such than tall igapó or várzea forests. In Roraima, igapó is associ- as the Surumu, Cotingo, Uraricoera, Tacutu, and the upper ated with most tributaries of the lower Rio Branco, such as the Rio Branco itself. Gallery forests usually cover a narrow (ca. Catrimani, Água Boa do Univini, Ajarani, Anauá, and Jaua- 50 m wide) band along the margins of watercourses, tending perí rivers (the latter being a tributary of the Rio Negro). to be low in stature (canopy reaching 10-15 m) and choked Campina and campinarana. These habitats, also known as with vines. These periodically fl ooded forests differ from white-sand savanna and white-sand forest, respectively, de- várzea and igapó of the humid Amazonian forest domain in velop in climates suitable for forests, but are associated with that gallery forests fl ood for shorter periods and with less pre- podzolized and oligotrophic quartzitic soils of extremely low dictable seasonality. nutrient content (Anderson 1981). They occur in areas with Dry forests. Semi-deciduous forests that occur within the seasonally variable edaphic conditions, caused either by soil savannas domain in areas with higher humidity conditions. fl ooding as a result of water table rise during the rainy season, They are particularly common at the edges of Amazonia, or by extreme dry soil conditions caused by the porosity of the where precipitation seasonality causes many trees to lose their sand (Pires and Prance 1985, Huber 1995). These habitats oc- leaves (Pires and Prance 1985). In Roraima, dry forests are cur in low-lying areas in central Roraima, and extend further patchily distributed (Silva 1997), occurring mainly around west towards the upper Rio Negro (Figure 3). The canopy of rocky outcrops along the transition between savanna and terra the campinarana is generally uniform and continuous, reach- fi rme domains, but also on hills in Curatella-dominated sa- ing 20-25 m in height (most often 10-15 m), having a distinc- vannas. Dry forests are physiognomically similar to gallery tive physiognomy composed by shrubs and small trees, dwarf forests but are not necessarily linked to rivers. and rachitic in aspect, with reduced foliage volume, small Sub-montane and montane humid forests. Areas above 202 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres

600 m in western Roraima are mostly covered by these habi- to access river-created habitats (várzea, igapó, and gallery for- tats. As a general rule, areas above 1000 m can be considered ests) we walked along the margins in times of low water, and montane, and are concentrated where Roraima borders Ven- used speed boats when these habitats were fl ooded. During ezuela and Guyana (Figure 3). In these regions the landscape the two expeditions in which MCH participated, he collected is dominated by steep and mountainous terrain, ranging from specimens, which are housed at the INPA Bird Collection in 600 to 2810 m on Mt. Roraima. Tepuis are spectacular, iso- Manaus. MPDS also did extensive collecting in Roraima, and lated, fl at-topped mountains with humid and densely forested these specimens are housed at the MPEG (details on the sites slopes and vertical sandstone cliffs that rise over the lowlands, and most important records obtained during MPDS’s expedi- with a particularly rich and endemic fl ora (Berry et al. 1995). tions will be published elsewhere). Tape-recordings by MCH Small treelets and bushes usually cover the plateaus, sur- and LNN are archived at the INPA Bird Collection. rounded by large barren areas. 13-19 March 2001. LNN and MFT surveyed areas close to Boa Vista, gallery forests along the lower Mucajaí and Anauá AREAS OF ENDEMISM FOR BIRDS IN RORAIMA rivers, the island of São José on the upper Rio Branco, and secondary terra fi rme forest near Alto Alegre (80 km west of Many bird taxa in the Neotropics (species or subspecies) Boa Vista), São Luiz, São João da Baliza, and Caroebe. share well-defi ned and congruent distributions (Haffer 1969, 14-28 May 2001. LNN, MCH, and MFT surveyed savan- 1974; Muller 1973). This spatial congruence was used to de- nas, dry forests, and gallery forests between Boa Vista and fi ne centers of dispersal (Haffer 1969, 1974) or areas of ende- Normandia, including Bonfi m, Serra do Tracajá, Serra da mism (Cracraft 1985), seven of which are located in the Ama- Onça, Contão, Conceição do Maú, Serra da Malacacheta, and zon. These areas are generally delimited by major Amazonian terra fi rme forest and campina/campinarana at Serra Grande rivers, such as the Amazon, Negro, Madeira, and Tapajós and Viruá National Park. rivers. The state of Roraima lies at the confl uence of three 6-11 July 2001. LNN and MFT joined an expedition of the such areas: Guianan, Imeri, and Pantepui. The Guianan area Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais of endemism roughly includes the lowland tropical forests of (IBAMA) covering by boat ca. 100 km of the lower Rio Branco, Guyana, Surinam, French Guiana, the state of Bolivar in Ven- from Santa Maria do Boiaçu to the mouth of the river, and also ezuela, and Brazilian Amazonia north of the Amazon River visiting Paraná da Floresta, and a portion of the Rio Jauaperí. and east of the Rio Negro (Cracraft 1985). Although its west- 8-9 August 2001. LNN and P. Coopmans made a short ern limit was defi ned by the Rio Branco by Cracraft (1985), visit to Viruá National Park and the Rio Uraricoera, close to this boundary is not explicit and is rather ambiguous, because the BR 174 highway. many contact zones of birds are located between the Rio 5-18 October 2001. LNN and J. Mazar Barnett joined an- Negro and the Rio Branco valleys (Haffer 1974). The Imeri other expedition sponsored by IBAMA, surveying the Eco- area of endemism is located in the vicinity of the Brazilian, logical Stations of Niquiá and Caracaraí, and the Viruá Na- Colombian, and Venezuelan borders, including Guainía and tional Park. This expedition covered by boat ca. 170 km of Vaupés in Colombia, the upper Rio Negro in Brazil, and the the lower Rio Branco, from Caracaraí south to the mouth of state of Amazonas in Venezuela (Cracraft 1985). Its endemic the Rio Catrimani, and also the rivers Água Boa do Univini, species and limits have been subject to criticism, mostly be- Viruá, Anauá, and Ajarani. cause several of the species that defi ne this area are taxa with 3-17 April 2003. LNN and MCH, accompanied for several poorly known distributions, and in many cases have proven days by M. J. Braun, revisited some areas explored in May to represent habitat specialists rather than geographic endem- 2001, particularly Serra do Tracajá. They also surveyed areas ics (Borges et al. 2001). The third area of endemism, which along the BR 174 highway and the Tacutu and Uraricoera riv- is characterized by birds associated with the tepuis, includes ers. LNN and MCH also dedicated four days to study the terra the Gran Sabana and Duida Subcenters, both of which have fi rme forests east of the Rio Branco, visiting Serra da Malaca- elements present in the mountains of northern Roraima. Addi- cheta and other forest patches along the RR 207 road (Boa tionally, several bird species from western Amazonia (gener- Vista to Serra do Tracajá), and areas near Vila União. ally absent from the core of the Guianan Shield) are present in western Roraima and their easternmost distributions seem to STUDY SITES coincide with the Rio Branco (see Discussion). We visited the following localities, which are in geograph- FIELD WORK ic order from north to south; numbers refer to the correspond- ing number of each locality on the map (Figure 2). During our expeditions to Roraima we conducted bird sur- Contão (4° 12’ N, 60° 32’ W). Amerindian village along the veys in the early morning (0500-1200), late afternoon (1500- Rio Cotingo, located at the southern end of hilly elevated lands, 1830) and, sporadically, at night, using binoculars and tape-re- which rise from the lowlands and extend north towards the Gran corders. We explored most areas using roads, which we trav- Sabana, across the Venezuelan border. We focused primarily on eled by foot or car. In forested areas we generally used trails; a boulder-strewn hill covered by dry forest, rising out of a Cu- The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 203

ratella-dominated savanna, about 2 km east of the village. We We worked along a 2-km trail within moderately disturbed also visited similar habitats along the fi rst two km along the primary forest with rock outcrops (see “Habitats” above) and road heading from this village to Mt. Roraima, and along the along the road to Cantá. road heading towards Boa Vista, where we made quick surveys Caracaraí Ecological Station (7). Protected area of 80,560 at Serra da Memória, Surumu, and Pedra Pintada. ha, situated to the west of the Rio Branco. The reserve is lo- Normandia (3° 53’ N, 59° 37’ W). Area covered by wet cated in a savanna-campinarana-terra fi rme transitional area, grasslands and savannas characterized by extensive colonies with patches of secondary terra fi rme forests surrounded by of tall, terrestrial termite mounds. agricultural lands and abandoned pastures. We focused our Bonfi m (3° 17’ N, 59° 53’ W). Town on the west bank of activities along the BR 210 road, and surveyed areas covered the Rio Tacutu, opposite Lethem in Guyana. We focused on with campinarana and secondary terra fi rme forest. the Curatella-dominated savannas surrounding Bonfi m, char- Viruá National Park (8). Protected area of 227,011 ha, sit- acterized by treeless hills to the west, and grassy wetlands to uated to the east of the Rio Branco and north of the Rio Anauá. the north, and on the gallery forests along the Rio Tacutu. The area, formerly planned for human settlement, became a Conceição do Maú (1) (3° 34’ N, 59° 53’ W). Ranch national park because of its poor fertility and inappropriate located at the confluence of the Tacutu and Maú rivers, quality for rural activities. Most of the area is located on sandy tributaries of the Rio Branco, and covered with gallery soils with poor drainage, covered by large expanse of campina forest. We focused on these forests along both rivers, but and campinarana. It also contains an isolated ridge of low also explored the extensive Curatella-dominated savannas hills with terra fi rme forest. We surveyed all main habitats around the ranch. available at the park, including campina and campinaranas Serra da Onça (2) (3° 08’ N, 60° 10’ W). Small chain of iso- (along the park’s main road), terra fi rme (next to the park’s lated hills covered by dry forests, and surrounded by expanses headquarters [1° 36’ N, 61° 13’ W]), várzea (along the Rio of fl at Curatella-dominated savannas and treeless rocky hills. Branco), and igapó (along the Rio Anauá). Boa Vista (2° 50’ N, 60° 40’ W). Economical and adminis- Niquiá Ecological Station (9). Protected area of 286,600 trative capital of the state, inhabited by nearly 200,000 people. ha, limited by the Rio Branco to the west, Rio Água Boa do We surveyed areas in a 15-20 km radius around the city, giving Univini to the east, and Rio Ajarani to the north. We surveyed special attention to the savannas south of the city, which in- all main habitats available at the reserve, including várzea clude a mosaic of grasslands, Moriche palm stands, wetlands, (along the Rio Branco), igapó (along the Água Boa do Uni- and agricultural fi elds. vini and Ajarani rivers), and terra fi rme. We concentrated our Serra da Malacacheta (3) (2° 41’ N, 60° 31’ W). For- surveys at Ecotur Park (00° 47’ 32’’ N, 61° 40’ 01’’ W), a ested hills at the interface of Amazonian humid forest and privately-owned lodge located opposite the reserve along the savanna realms, where partially disturbed humid forest are Rio Água Boa do Univini, from where we explored savannas, surrounded by Curatella-dominated savannas (apparently campinaranas and partially disturbed terra fi rme forest. infl uenced by grazing and burning). We include under these Upper Rio Branco. Section of the river delimited by the name similar lowland second-growth humid forests (2° 45’ confl uence of the Uraricoera and Tacutu rivers and Mucajaí. N, 60° 18’ W) present along the road from Boa Vista to The margins of the upper Rio Branco are covered by narrow Serra do Tracajá. strips of gallery forest. Among other places, we visited the Serra do Tracajá (4) (2° 36’ N, 60° 03’ W). Rocky hills island of São José (02° 56’ N, 60° 31’ W), which is covered by cloaked in dry forest, which rise abruptly out of extensive Cu- a mosaic of gallery forest and Cecropia-dominated forest. ratella-dominated savannas. With this name we refer to Serra Lower Rio Branco. Section of the river south of Caracaraí. do Tracajá itself, and other possibly unnamed nearby hills The margins of the lower Rio Branco are covered by várzea, with similar vegetation and physiognomy, located to the west including different successional stages, such as sandbars, low (2° 35’ N, 60° 07’ W) and southeast (2° 31’ N, 60° 00’ W and riparian scrub, Cecropia-dominated forest, and mature várzea 2° 28’ N, 60° 01’ W). forest. We intensively surveyed the whole extent of this part Rio Mucajaí (02° 28’ N, 60° 54’ W). Tributary of the Rio of the river, exploring by boat both margins and several river Branco that we explored close to the village of Mucajaí. We islands, such as the islands of Carneiro (10) (01° 24’ N, 61° visited both margins, covered by narrow stretches of disturbed 16’W), Inajatuba (11) (01° 18’ N, 61° 18’ W), and Cota (16) gallery forest. (01° 16’ S, 61° 50’ W). Serra Grande (5) (2° 34’ N, 60° 46’ W). Isolated mountain São Luiz (12), São João da Baliza (13), and Caroebe (14). (898 m) located at the interface of Amazonian humid forest Villages located along the BR 210 road, with disturbed terra and savanna realms. Covered with partially disturbed humid fi rme forest. We focused our surveys on forests present along forest in which we climbed to about 300 m. secondary roads. We explored relatively well-preserved terra Vila União (6) (02° 0’ N, 60° 38’ W). Small village sur- fi rme forests at the edge of the Wai-Wai Indian Reservation rounded by primary and secondary terra fi rme forest along (01° 05’ N, 59° 56’ W) and around the village of São Luiz. the RR 170 road. Most areas around the village were logged Rio Jauaperí. Black-water tributary of the Rio Negro that or recently occupied by government-sponsored settlements. forms the boundary between the states of Amazonas and Rorai- 204 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres

ma. We navigated and explored the lower part of the river, from recorded (604 species); 6% of the species (40 species) lack its mouth to the village of Samaúma (15). Most of the river was voucher specimens, but are documented by our tape-record- sparsely inhabited and covered by large expanses of tall igapó. ings. The remaining 12% (93 species) are listed based only on Paraná da Floresta. Narrow stretch of the Rio Negro, in sight or auditory records, but we believe that their inclusion is southernmost Roraima, covered by tall igapó. We explored acceptable because they are not especially diffi cult to identify the villages of Remanso (17) and Floresta (18), and neighbor- and are likely to occur in Roraima. Undocumented species for ing river islands. which we judge records to be controversial were regarded as hypothetical (see below). SPECIES LIST In terms of species richness, habitat heterogeneity (beta- diversity) is responsible for 60% of the avian diversity of Ro- We present a list of all bird species recorded in the state of raima, given that 446 bird species are restricted to single habi- Roraima, which include data obtained during our fi eldwork in tats (Table 1). Terra fi rme forests have the highest number of Roraima in 2001 and 2003, data gathered from museum speci- species among all habitats, contributing with nearly 300 spe- mens, and published and unpublished sources, all of which are cies (348 including predicted species, see Discussion). Flood- acknowledged in the Appendix. Species are assigned to their ed forests, montane forests and tepuis, and savannas each ac- main habitats, based on our experience in Roraima and comple- count for 11% of the avifauna, whereas campinas (3%), gal- mented by published data from the state. We include notes on lery and dry forests (4%), and wetlands (9%) are responsible distribution and seasonality, which refer to general areas of en- for a minor proportion of the avifauna (Table 1). demism and to migratory species, respectively. We also present Forty-fi ve species are migratory in the state of Roraima. the physical evidence used for the inclusion of each species in Twelve of these reach Roraima during the austral winter (aus- the list (specimen or tape-recordings; species lacking objective tral migrants), between April and August, whereas 29 species evidence refer to sight or auditory records). Tape-recordings are nearctic migrants, occurring in Roraima between Septem- are exclusively of our own, because we did not have access to ber and April (Appendix). Four species seem to have both other recordings from the state. Taxonomy and species nomen- resident and migratory populations, the latter coming either clature follow the Brazilian Ornithological Records Committee from the south (Tyrannus savana, Progne tapera, and Myio- (Comitê Brasileiro de Registros Ornitológicos 2006). dynastes maculatus) or the north (Vireo olivaceus). The record of Stercorarius parasiticus and Pelecanus occidentalis prob- ably represent vagrants. Another two species, Coccyzus euleri RESULTS and Tersina viridis, may also be migratory, but little data are AVIAN DIVERSITY available for these species in Roraima.

We recorded 512 bird species, 38 of which were the fi rst re- HYPOTHETICAL SPECIES cords for the state (see below). We list for the state of Roraima a total of 741 bird species, recorded as of the writing of this Fifteen species previously reported for Roraima are here in- paper (Appendix). Specimens document 82% of the species cluded as hypothetical and removed from the state’s list because

Table 1. Relative importance of habitats in terms of bird species richness in Roraima.

Terra fi rme forest Flooded Montane Campina & Gallery & Wetlands Habitata forest & Savanna Generalistsf Migr.g c d e Wam General Gui forest tepui campinaranas dry forests & rivers

Number of 31 190 75 87 85 85 21 32 70 taxab 45 42 (exclusive spp.) (25) (117) (53) (41) (75) (41) (12) (19) (63)

% of total 4 26 10 12 11 11 3 4 9 5 6 a Information derived from Appendix. Taxa occurring in more than one habitat were assigned to their preferred one (fi rst one mentioned in their respective column). b Includes different subspecies, thus the sum of the number of taxa is higher that the number of species recorded in Roraima. c Western Amazonian taxa (species or subspecies typical of western Amazonia, generally absent east of the Rio Branco). d Widespread Amazonian taxa. e Guianan taxa (species or subspecies typical of the Guianan area of endemism, generally absent south of the Amazon River and west of the Rio Negro). f Species found in numerous habitat types without clear preference. g Migratory species using a variety of habitats. The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 205

they are either unlikely to occur in the state or species diffi cult agree with Silva’s conclusions and exclude it from the list. On to identify in the fi eld without physical documentation. the other hand, it is plausible (and even likely) that C. undatus Anas georgica (Yellow-billed Pintail). Included in Roraima will eventually be recorded in the state, but probably to the based on a sight record made by G. Shattuck (1926) near Boa east of the Rio Branco. Two species previously dismissed by Esperança along the Uraricoera River. This record was admitted Silva (1998) for Maracá based on the same criteria (Ptero- by Pinto (1966, 1978). However, we suspect that these records glossus aracari and Tyranneutes virescens) have subsequently may represent a misidentifi cation. This duck is not known either been recorded in Roraima east of the Rio Branco (see species from Venezuela (Hilty 2003) or Guyana (Braun et al. 2000), and accounts for T. virescens). breeds thousands of km to the south of the study area. However, Campylorhamphus sp. (Scythebill). Birds belonging to it does migrate north from southern South America to winter in this genus were observed by Stotz (1997) in Sorocaima and large numbers in southern Brazil, and there is always a slight by Trolle and Walther (2004) along the Rio Jauaperí, but none possibility of a vagrant occurring in Roraima, but this hypoth- was either identifi ed to species or tape-recorded. The most esis should be supported with specimens. plausible species at Sorocaima is C. trochilirostris, which has Phaethornis augusti (Sooty-capped Hermit), Colibri thalas- been collected very close to that locality in Venezuela (Hilty sinus (Green Violet-ear), Avocettula recurvirostris (Fiery-tailed 2003; A. Whittaker, unpubl. data). We believe that records Awlbill), and Klais guimeti (Violet-headed Hummingbird). from Rio Jauaperí may represent C. procurvoides, the species These four species of hummingbirds were cited for Roraima typically present in terra fi rme forest and igapó. based on data from Augusto Ruschi’s collection, held at the Grallaria guatimalensis (Scaled Antpitta). Cited for Ro- Museu de Biologia Melo Leitão (Ruschi 1961). The general raima by Sick (1997), based on a specimen from “Serra do accuracy of the data on the specimens at this collection has Curupira, Estado de Roraima”. The specimen was fi nally been heavily criticized (Pacheco 1995b, Stotz 1998, Pacheco re-identifi ed as Grallaria varia cinereiceps, and its locality and Bauer 2001), and the source of those specimens remains corrected to Amazonas state (Mallet-Rodrigues and Pacheco dubious. Note that P. augusti and C. thalassinus would rep- 2003); therefore, there are no known documented records for resent the fi rst documented Brazilian records, but have been G. guatimalensis in Brazil, or for G. varia in Roraima. dismissed by Pacheco (1995b) and the Comitê Brasileiro de Tyrannus dominicensis (Gray Kingbird). Included in Ro- Registros Ornitológicos (2006), who we follow in excluding raima by Moskovits et al. (1985), based on a sight record at them from Roraima. Three other species (Doryfera johannae, the island of Maracá. However, this would represent the only Colibri delphinae, and Lophornis ornata) are also present in known record for Brazil, and has been dismissed by the Co- Ruschi’s collection, but were included in the list based on docu- mitê Brasileiro de Registros Ornitológicos (2006) until docu- mented records from other sources. mented records are available. We expect this species to occur Colibri coruscans (Sparkling Violet-ear). Cited for Ror- in the area as a migrant, but here follow the Committee in aima by Sick (1997) based on an alleged record from Cerro excluding it from the main list. Urutaní by Dickerman and Phelps (1982). This citation ap- Progne elegans (Southern Martin) and Progne subis (Purple pears to be erroneous, as this species is not mentioned in the Martin). More than 20 adult males and one female-plumaged original article. bird believed to be P. elegans (a taxon previously considered Electron platyrhynchum (Broad-billed Motmot). Cited for a subspecies of P. modesta) were observed by A. Whittaker Maracá Ecological Station by Moskovits et al. (1985), but (pers. comm.) on 20 August 1993 along the lower Rio Branco. dismissed by Silva (1998), who suggested an identifi cation He noted deep-forked tails on all birds as well as the distinct problem. This species is not known from anywhere near Ro- dark ventral area of the female. Subsequently, on 9 June 1995, raima; its distribution being mostly restricted to the south of A. Whittaker also found four adult males perched with Progne the Amazon River (Snow 2001). tapera on a dead tree along the lower Rio Branco. These birds Celeus fl avescens (Blond-crested Woodpecker). Listed for probably represent austral migrants from southern South Amer- Roraima by Pinto (1966) based on an alleged Boa Vista speci- ica, but distinguishing between this taxon and Progne subis men mentioned by Cory (1919). However, this specimen refers can be challenging in the fi eld. Likewise, Trolle and Walther to another locality formerly known as Boa Vista in the Brazil- (2004) recorded birds that they identifi ed as P. subis, but lack ian state of Maranhão and now called Santo Amaro (Paynter any objective evidence. Although we believe that both species and Traylor 1991). According to D. Willard (in litt.) the original are likely to be present in Roraima, we prefer to keep their pres- label refers simply to Boa Vista, but a secondary label assumed ence as hypothetical until specimens are obtained. that the specimen was from Roraima; thus, the species was sub- Tachyphonus rufus (White-lined Tanager) and Euphonia sequently reported as such by Cory (1919) and Pinto (1966). laniirostris (Thick-billed Euphonia). Both species were cited The label has since been corrected to Boa Vista, Maranhão. for Roraima by Borges (1994) near Boa Vista, but the author Celeus undatus (Waved Woodpecker). Included for Ma- suggested that they may have been misidentifi ed (S. Borges, racá Ecological Station by Moskovits et al. (1985), but dis- pers. comm.). E. laniirostris has also been mentioned by Mos- missed by Silva (1998), based on the confi rmed presence kovits et al. (1985) for Maracá, but that record was also dis- (specimens) of its allospecies C. grammicus at that site. We missed by Silva (1998). 206 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres

NEW RECORDS naevia, Inezia subfl ava, Hemitriccus minor, Schiffornis ma- jor, Hylophilus semicinereus, Ammodramus aurifrons, and We obtained 38 new state records during our fi eld work in Sicalis columbiana. Most of these species are widespread and Roraima. As expected, most of our additions are represented common in várzea forests elsewhere in Amazonia, and all are by species occurring in habitats or biogeographical regions present along the lower Rio Negro or the Solimões-Amazonas poorly sampled (or not sampled at all) previously. Among river systems. In many cases our records extend the known these are the terra fi rme forests east of the Rio Branco, the distribution of várzea species approximately 300 km north- várzeas of the lower Rio Branco, and the igapó, campinas and ward, from the mouth of the Rio Branco up to Caracaraí (de- campinaranas of central Roraima. tails of these records will be published elsewhere, Naka et al. Within terra fi rme forests we include 13 species previ- in prep). ously unknown from the state: Forpus modestus, Trogon ru- The campinas, which were previously unexplored in Ror- fus, Selenidera piperivora, Piculus chrysochloros, Sclerurus aima also presented new species for the state: Myrmeciza dis- mexicanus, Myrmotherula gutturalis, Formicarius analis, juncta, Hemitriccus inornatus, and Rhytipterna immunda. Myiopagis caniceps, Hemitriccus zosterops, Cotinga cotinga, Apart from unexplored habitats, we made nine additions in Tyranneutes virescens, Hylophilus thoracicus, and Euphonia relatively well-sampled areas such as the savannas of north- cayennensis. With the exception of F. modestus, all other spe- eastern Roraima, including Buteo brachyurus, Micropygia cies occur both in Manaus and Guyana (Cohn-Haft et al. 1997, schomburgkii, Calidris melanotos, Berlepschia rikeri, Pi- Braun et al. 2000) and were expected to occur in Roraima. cumnus cirratus, and Basileuterus fl aveolus. Near Boa Vista Taxa such as S. piperivora, M. gutturalis, T. virescens, and in heavily degraded areas, we recorded Podilymbus podiceps, Euphonia cayennensis represent eastern elements, being re- Gallinula chloropus, and Tyto alba. stricted to the Guianan area of endemism, and we suggest that they are restricted to the east of the Rio Branco (Table 2). SPECIES ACCOUNTS We found 13 new species for the state in riverine habitats Here we present details on 22 species which present spe- (either in várzea or igapó): Leucopternis schistaceus, Monasa cial interest, either by representing new records for Roraima nigrifrons, Cranioleuca gutturata, Myrmotherula klagesi, with some biogeographical signifi cance, poorly-known and Myrmotherula assimilis, Cercomacra nigrescens, Sclateria restricted species, or bird species of conservation concern.

Table 2. Pairs of taxa (allospecies or subspecies) whose distribution boundaries apparently coincide with the Rio Branco. Taxon west of the Rio Branco Taxon east of the Rio Branco Tinamus major zuliensis T. m. major Gypopsitta barrabandi G. caica Notharchus macrorhynchos N. hyperrhynchus Capito auratus C. niger Ramphastos tucanus cuvieri R. t. tucanus Ramphastos vitellinus culminatus R. v. vitellinus Selenidera nattereri* S. piperivora Celeus elegans jumanus C. e. elegans Cymbilaimus lineatus intermedius C. l. lineatus Myrmotherula haematonota M. gutturalis Myrmotherula menetriesii pallida M. m. cinereiventris Hypocnemis cantator fl avescens H. c. cantator Percnostola rufi frons minor P. r. subcristata Schistocichla leucostigma infuscata S. l. leucostigma Dendrocincla fuliginosa phaeochroa D. f. fuliginosa Sittasomus griseicapillus amazonus S. g. axillaris Glyphorynchus spirurus rufi gularis G. s. spirurus Lepidocolaptes albolineatus duidae L. a. albolineatus Phoenicircus nigricollis* P. carnifex* Tyranneutes stolzmanni T. virescens Hylophilus ochraceiceps ferrugineifrons H. o. luteifrons Euphonia rufi ventris E. cayennensis * Species not known from Roraima, but likely to be recorded with further sampling. The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 207

Details of those species found in fl ooded forest will be dealt has also only been found in areas where G. galbula was absent with elsewhere (Naka et al., in prep). (Robbins et al. 2004), suggesting that both species may be Micropygia schomburgkii (Ocellated Crake). MCH, LNN, excluding each other ecologically and geographically. These and MFT heard at least two individuals vocalizing in season- represent the fi rst records for the state of Roraima. ally inundated savannas near Serra do Tracajá on 22 May Capito niger (Black-spotted Barbet). MCH and LNN heard 2001. Subsequently, in November 2004, A. Whittaker and K. the distinctive vocalizations of nominate C. niger in terra Zimmer saw and tape-recorded three different birds 22 km fi rme forest near Vila União on 17 April 2003. Until recently, south of Pacaraima (A. Whittaker, in litt.). This secretive spe- most classifi cations in the last half-century treated C. niger as a cies is known from several sites in neighboring Venezuela widespread polytypic Amazonian species. Current authors (e.g. (Hilty 2003), French Guiana (Tostain et al. 1992), and Guy- Hilty 2003, Dickinson 2003, Comitê Brasileiro de Registros ana (Robbins et al. 2004), but these are the fi rst records for Ornitológicos 2006, but not Short and Horne 2002), however, Roraima, suggesting that its presence has been overlooked in have followed Haffer (1997) in reverting to an older classifi - the past. cation that restricts the name niger to the nominate form and Forpus modestus (Dusky-billed Parrotlet). MCH observed elevates auratus to species level, including most of the other and tape-recorded two individuals in terra fi rme forest with subspecies in the group. With the exception of Pinto (1966) and interspersed rocky outcrops near Vila União on 17 April 2003. Dickerman and Phelps (1982), who specifi cally mention either Although this parrotlet is widespread in Amazonia, its distri- C. auratus or a subspecies currently considered to belong to butional patterns are not yet well understood (Collar 1997). the auratus group, authors that mention C. niger for Roraima We believe that its distribution may be linked to areas with (Borges 1994, Moskovits et al. 1985, Stotz 1997, Silva 1998) considerable topographical relief and rocky soils, which may may be assumed to be referring to the broad concept of the explain its presence in Roraima and its apparent absence near species. All these records are from west of the Rio Branco, Manaus. Although new to Roraima, its presence was expect- which we believe represents the divide between C. niger and ed, as it occurs in neighboring Venezuela (Hilty 2003) and auratus, an idea also championed by Haffer (1997). Therefore, Guyana (Braun et al. 2000). we believe that all previous records from Roraima refer to C. Gypopsitta caica (Caica Parrot). A fl ock of nine individu- auratus, and ours (the only one east of the Rio Branco) should als was tape-recorded and one of them collected by MCH represent the fi rst record of C. niger for the state. on 17 April 2003 in terra fi rme forest near Vila União. The Selenidera piperivora (Guianan Toucanet). A single in- only other specimen available from the state was collected dividual was heard by MCH in terra fi rme forest near Vila by D. Stotz near fazenda Santa Cecilia (FMNH #389173). União on 17 April 2003. This represents the fi rst record for the Both specimens were collected east of the Rio Branco. Its state of Roraima. We agree with Haffer (1974) envisioning the allospecies, G. barrabandi, seems to replace G. caica west distribution of this species as being restricted in Roraima to of this river. However, MCH also observed a fl ock of G. bar- the east of the Rio Branco, being replaced by S. nattereri fur- rabandi on the east (left) bank of the lower Rio Branco, near ther west. Note that S. nattereri has not yet been recorded in the mouth of the river in July 1999. Throughout most of their Roraima, but we suspect that its absence may represent poor distribution, species in the genus Gypopsitta are parapatri- sampling in western Roraima (see Discussion). cally distributed; therefore having both species in possible Picumnus cirratus (White-barred Piculet). MCH, LNN, contact on the lower Rio Branco is an interesting issue re- and MFT tape-recorded and collected pairs in gallery forest quiring further study. The presence of G. barrabandi east of along the Rio Tacutu, near Conceição do Maú, and in dry for- the Rio Branco also represents an exception to the observed est at Serra do Tracajá and north of Boa Vista. These birds pattern of western elements being restricted to west of the may refer to the form confusus, known from southwestern Rio Branco (see Discussion). Guyana, French Guiana, and extreme northern Brazil (Win- Chordeiles pusillus (Least Nighthawk). We found and col- kler and Christie 2002). Although the latter authors include lected two morphotypes with distinct vocalizations, within the Roraima as the range of this race, we could fi nd no previous boundaries of the state. One appears to be resident in the cer- record of this species for the state. rado-like vegetation in the north and the other in the campinas Lepidocolaptes albolineatus (Lineated Woodcreeper). LNN of the south. Cohn-Haft will present a taxonomic revision of and MCH tape-recorded several individuals in terra fi rme for- Amazonian Chordeiles pusillus elsewhere. est near Vila União on 17 April 2003. There are two distinct and Galbula leucogastra (Bronzy Jacamar). Two individuals well-marked allopatric subspecies north of the Amazon River, were tape-recorded by LNN and J. Mazar Barnett in tall camp- which differ in morphology and vocalization (Marantz et al. inarana forest at Viruá National Park on 16 October 2001. Al- 2004). Our observations and tape-recordings in Roraima match though we have found this species to be common in campinas specimens and recordings from Manaus, where the nominate and campinaranas elsewhere in Amazonia, it appears to be race occurs (Marantz et al. 2004). All previous records from rare and local in Roraima. Surprisingly,Galbula galbula, on Roraima are from west of the Rio Branco (Cory and Hellmayr the other hand, was fairly abundant in campinas, savannas, 1925, Pelzeln 1868-71, Stotz 1997, Silva 1998, Silva and Oren and várzeas throughout the state. In Guyana, G. leucogastra 1990), and whenever subspecies were mentioned, authors refer 208 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres

to L. albolineatus duidae. Therefore, we believe that the Rio Branco was somewhat surprising, as this species is apparently Branco separates the range of these taxa, and our records ap- absent from other white-water rivers in Amazonia. pears to be the fi rst of the nominate race in the state. Myrmeciza disjuncta (Yapacana Antbird). MCH, LNN, and Synallaxis kollari (Hoary-throated Spinetail). A poorly MFT tape-recorded and collected a pair, but noted at least anoth- known, local, and endemic species found exclusively in gal- er three different individuals in a campina covered by clumps of lery forests from northeastern Roraima and adjacent Guyana, tall sedges interspersed with low bushes at Viruá National Park along the upper Rio Branco and some of its tributaries (Tacu- on 25 March 2001. In October 2001, LNN and J. Mazar Bar- tu, Surumu, and Cotingo rivers) (Ridgely & Tudor 1994, Col- nett located another pair at a nearby site within Viruá. The pair lar et al. 1992, BirdLife International 2000). Described from collected represents the fi rst record for Roraima, and the sec- fi ve specimens collected by Natterer in 1831-1832 along the ond Brazilian locality. M. disjuncta is a rare and poorly known Rio Branco near Boa Vista (Pelzeln 1956), this species was species, until recently only known from two sites in Venezuela rediscovered over a century later on the Rio Surumu (Pinto and one in Colombia (Zimmer 1999). The only other Brazilian 1966). Recent records come from Conceição do Maú (Forrest- locality is Jaú National Park (Borges and Almeida 2001), where er 1995) and the Rio Uraricoera (Grosset and Minns 2002). In this species was found in a campina located more than 500 km 27 May 2001 MCH, LNN, and MFT found and tape-recorded southwest of our site. Despite the large distance between sites, one bird along the Rio Cotingo at the village of Contão (one the campinas of Viruá and Jaú are physiognomically similar. of the historical localities mentioned by Pinto 1966). Subse- Not only do our records represent a considerable range exten- quently, in August 2001 and April 2003, LNN and P. Coop- sion on the north side of the Rio Negro, suggesting that the spe- mans, and MCH and LNN, respectively, found several birds cies occurs throughout the upper Negro basin in appropriate at the Rio Uraricoera at the site reported by Grosset and Minns habitat, but they are the fi rst from east of the Rio Branco, sug- (2002). At all sites, birds were found in or near vine tangles in gesting that the river is not a barrier to its distribution and that gallery forests, and were highly responsive to playback. At the the species may occur considerably further east still. Rio Uraricoera, birds were common, and at least fi ve differ- Sublegatus modestus (Southern Scrub-Flycatcher). MCH ent individuals were counted within 200 m of gallery forest. and LNN, accompanied by M. Braun, tape-recorded and col- Given its restricted distribution and the very low number of lected a pair of Sublegatus fl ycatchers on 9 March 2003 in records, S. kollari is presently considered threatened (BirdLife the savannas of Serra do Tracajá, fi nding several others on International 2000), but was excluded based on defi ciency of subsequent days. These birds were morphologically and vo- data from the Brazilian list of threatened birds (Ministério do cally similar to birds collected from the Rupununi savannas in Meio Ambiente 2003). An accurate assessment of its distribu- Guyana (M. Braun pers. comm.), which may represent a resi- tion and ecological requirements its being undertaken at pres- dent form of S. modestus (M. Robbins, in litt.). The race brevi- ent (Vale et al., in prep.). This new information is important rostris, which breeds from southern Bolivia to Argentina, is an to assess the level of threat to which this species is subject austral migrant in Amazonia, but is mostly recorded south of throughout its small range (see Discussion). the Amazon (Fitzpatrick 2004). Three specimens from the Ro- Myrmotherula gutturalis (Brown-bellied Antwren). MCH raima savannas held at the FMNH were identifi ed as S. modes- and LNN observed and tape-recorded a pair in a mixed-species tus and were taken in March and December, outside the main fl ock in the understory of terra fi rme forest near Vila União on austral migrant season. These records, along with those of the 17 April 2003, representing the fi rst record for Roraima. We Rupununi (Robbins et al. 2004), suggest the existence of a suspect that this species is limited in Roraima to terra fi rme resident form in these savannas, extending the former breed- forests east of the Rio Branco, being replaced by M. haema- ing area several hundred km northward. Further fi eld work is tonota further west. being conducted to assess the proper taxonomic status of these Myrmotherula cherriei (Cherrie’s Antwren). MCH, LNN, populations (Robbins et al., unpubl. data). and MFT encountered and tape-recorded several individu- Polystictus pectoralis (Bearded Tachuri). LNN and MFT als in a campina at Viruá National Park on 25 May 2001. In found one bird on March 2001 in natural grasslands 50 km July 2001, LNN and MFT saw and tape-recorded a female in south of Boa Vista, and in May 2001, MCH, LNN, an MFT várzea forest near Santa Maria do Boiaçu. Subsequently, in tape-recorded several individuals in the savannas near Serra do October 2001, LNN and J. Mazar Barnett found this species Tracajá. This poorly known species is apparently undergoing to be fairly common in tall and low igapó along the Água Boa severe population declines both in northern and southern South do Univini and Jauaperí rivers, and in a campinarana at the America as a result of grassland conversion into pastures and Caracaraí Ecological Station. This antwren is known to occur crop fi elds (Parker and Willis 1997). At present, this species is locally in eastern Colombia, southern Venezuela, and north- considered near-threatened (BirdLife International 2000) and eastern Peru, whereas in Brazil it was only known from the the drastic conversions of natural grasslands in Roraima repre- Rio Negro basin (Isler et al. 1999, Zimmer and Isler 2003), sent a clear threat for these populations (see Discussion). but not from the Rio Branco basin. Although its presence in Atalotriccus pilaris (Pale-eyed Pygmy-Tyrant). In May campina, campinarana, and igapó was not unexpected in Ro- 2001, MCH, LNN, and MFT tape-recorded and collected raima, its appearance in várzea forests along the lower Rio one individual in dry forest on a boulder-strewn hill near the The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 209

village of Contão, representing the fi rst Brazilian specimen. (Moskovits et al. 1985), but this record was dismissed by Sil- This species seems to be restricted in Brazil to a savanna type va (1998), arguing that a specimen of T. stolzmanni from the found only in northeastern Roraima, characterized by rocky same site was enough evidence to suspect a misidentifi cation. boulders, barren soil, and large termite mounds. Atalotriccus We agree with Silva, and believe that the presence of both al- pilaris was fi rst observed in Brazil by Forrester (1993) along lospecies in syntopy would be, at best, unlikely. We suspect the Rio Cotingo and subsequently tape-recorded by A. Whit- that the Rio Branco represents the distributional limit for both taker (pers. comm.) on a hill with low dry forests along the BR species, and therefore our records east of the Rio Branco must 174 highway, 35 km south of the Venezuelan border, and by represent the fi rst for Roraima. Willis (2003) at a nearby locality. Basileuterus fl aveolus (Flavescent Warbler). MCH, LNN, Hemitriccus zosterops (White-eyed Tody-Tyrant). LNN and MFT found and tape-recorded one bird in gallery forest and MFT saw and tape-recorded one bird in disturbed terra along the Rio Tacutu, near Conceição do Maú, on 20 May 2001. fi rme forest near Samaúma on 8 July 2001. In April 2003, Subsequently, they found this species in several sites domi- MCH and LNN heard another individual vocalizing in terra nated by dry forests, tape-recording and collecting specimens fi rme forest near Vila União. The vocalizations recorded at at Serra da Onça, Rio Uraricoera, and along the BR 174 high- both sites are similar to those known from Manaus (Cohn- way, 40 km north of Boa Vista. In 1989 this species was found Haft et al. 1997) and presumably refer to the taxon rothschildi at Dadanawa, along the Rupununi River in Guyana (Mees (Cohn-Haft 2000; MCH will revise the taxonomy of this and and Mees-Balchin 1990), which represented an extraordinary other Hemitriccus spp. elsewhere). range extension of over 1000 km from previously described Hemitriccus inornatus (Pelzeln’s Tody-Tyrant). LNN and populations in northern Venezuela and central Brazil (Ridgely J. Mazar Barnett found and tape-recorded two individuals on 9 and Tudor 1989). Ours are the fi rst records for Roraima but are and 10 October 2001 in tall campinarana along the Rio Agua likely to represent the same population sampled in Guyana. Boa do Univini at the Niquiá Ecological Station, and up to four birds on 14 October 2001 within the Caracaraí Ecological DISCUSSION Station. Until recently, this species was only known from the type specimen (Ridgely and Tudor 1994), but recent fi eld work PREDICTED AVIFAUNA OF RORAIMA has led to its rediscovery (Whittaker 1994) and ample range extensions (Zimmer, Whittaker, and Cohn-Haft, in prep.) A complete avifaunal survey of an area the size of Ror- Cnemotriccus fuscatus (Fuscous Flycatcher). We found aima, with its many ecological and biogeographical complexi- two distinct forms within the state, one associated with gallery ties, represents an overwhelming task. If we are to fully under- forests and the other with campinas. In March 2001, MCH, stand the distribution patterns and biogeographical aspects of LNN, and MFT encountered and collected what apparently all birds occurring in the state, we need much more intensive represented Cnemotriccus f. fumosus (a taxon well document- fi eld work, especially in remote regions that continue to re- ed by specimens from the savannas of Roraima) in gallery ceive little or no coverage, and more intensive collecting (with forests along the upper Rio Branco and its tributaries, the Ura- tape recordings and tissue samples) to allow recognition of the ricoera and Urubú rivers. A few days later, MCH, LNN, and true taxonomic or population-level diversity present. MFT tape-recorded a second vocal type in a campina at the Although we are far from having a complete list of all bird Viruá National Park. The latter bird appears to be closest to species occurring in Roraima, we can make inferences and pre- individuals found in other campinas, which apparently repre- dictions about the species that are expected to occur there. We sent the form C. f. duidae (Whitney, unpubl. data), not previ- have a good idea of the species present in neighboring regions, ously recorded in Roraima. such as Manaus (Cohn-Haft et al. 1997), the states of Amazo- Rhytipterna immunda (Pale-bellied Mourner). LNN and nas and Bolivar in Venezuela (Hilty 2003), Guyana (Braun et J. Mazar Barnett observed and tape-recorded individuals in a al. 2000), the Rupununi savannas (Robbins et al. 2004), the up- campinarana at the Caracaraí Ecological Station on 14 Octo- per Rio Negro (MCH, LNN, and colleagues, unpubl. data), and ber 2001. A few days later, they tape-recorded another bird in the right bank of the middle Rio Negro (Borges et al. 2001). a campina at the Viruá National Park. The distribution of this Taking into account our results and the biogeographical species seems to be tied to sandy soils (Sick 1997, Zimmer and affi nities of the avifauna of Roraima, we estimate conserva- Hilty 1997), and although these represents the fi rst records for tively that over 800 species occur within the state boundaries. Roraima, we were expecting to fi nd this species in the state. Based on a direct comparison with Cohn-Haft et al. (1997), Tyranneutes virescens (Tiny Tyrant-Manakin). MCH and Hilty (2003), Braun et al. (2003), and Robbins et al. (2004), LNN tape-recorded several individuals in terra fi rme forest we include a list of 69 species that are likely to be recorded in near Vila União on 17 April 2003. This species seems to be Roraima with further fi eld work (Table 3). Among these are limited to terra fi rme forests east of the Rio Branco, possi- 39 species that inhabit terra fi rme forests (20 species widely bly being replaced by T. stolzmanni to the west of this river. distributed in Amazonia, 12 species representing western ele- The only previous record of T. virescens within Roraima was ments present on the upper Rio Negro and extreme southern represented by a sight record from Maracá Ecological Station Venezuela, and 7 species known from the Guianan area of en- 210 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres

Table 3. Bird species predicted to occur in Roraima. Includes species known from areas near Roraima and not separated by any apparent biogeographical barrier; based on Mayr and Phelps (1967), Stotz et al. (1992), Cohn-Haft et al. (1997), Zimmer and Hilty (1997), Braun et al. (2000 and 2003), Borges et al. (2001), Hilty (2003), and Robbins et al. (2004).

Bird species from terra fi rme forest Amazona autumnalis c Celeus undatus c E. aurantioatrocristatus b Nyctibius leucopterus b Megastictus margaritatus a Phylloscartes virescens c N. bracteatus b Schistocichla caurensis ae Lophotriccus vitiosus b Lurocalis semitorquatus b Myrmeciza pelzelni a Neopipo cinnamomea b Chaetura chapmani b Dichrozona cincta a Iodopleura isabellae a Phaethornis malaris a Phlegopsis nigromaculata a Phoenicircus nigricollis a Topaza pyra a Grallaria varia b Phoenicircus carnifex c Avocettula recurvirostris b Deconychura stictolaema b Haematoderus militaris c Discosura longicauda b Campylorhamphus procurvoidesb Lepidothrix serena c Pharomachrus pavoninus b C. trochilirostris b Turdus lawrencii a Notharchus ordii b Philydor erythrocercum b Lamprospiza melanoleuca b Malacoptila fusca b P. erythropterum b Periporphyrus erythromelas c Selenidera nattereri a Xenops milleri b Schistocichla caurensis ae

Bird species associated with tepuis adjacent to Roraima Amazona dufresniana*d Grallaria guatimalensis* Xenopipo uniformis Campylopterus duidae Grallaricula nana* Cichlopsis leucogenys Colibri coruscans Elaenia dayi* Diglossa duidae Polytmus milleri* Pachyramphus viridis Euphonia cyanocephala Heliodoxa aurescens Pipreola whitelyi*f

Nearctic migrants that regularly visit Guyana, southern Venezuela, or Manaus Calidris himantopus Tyrannus tyrannus Catharus ustulatus Falco columbarius Tyrannus dominicensis Dolichonyx oryzivorus Buteo platypterus Vireo altiloquus

Savanna bird species found in the southern Rupununi Oxyura dominica Bubo virginianus Carduelis cucullata* Phaethornis augusti* Neopelma pallescens

Bird species associated with campina/campinarana Crypturellus duidae Attila citriniventris Dolospingus fringilloides

* Species not yet recorded in Brazil. a Western taxa present on the upper Rio Negro and known from extreme southern Venezuela, but generally absent from Guyana or Manaus. b Widespread taxa present throughout Amazonia, including the state of Amazonas, Brazil, and southern Venezuela. c Eastern elements, or Guianan endemics, present in Guyana and/or Manaus. d Although not especially linked to tepuis, Venezuelan records in Bolivar state come from the tepui foothills. e Associated to rocky areas, specially in sub-montane humid forest. f Reported twice near Manaus (Cohn-Haft et al. 1997), but lacking documentation for Brazil. The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 211

demism). Other species that will probably be recorded include zones, the valley of the Rio Branco seems to be an important 5 species reported from the southern Rupununi savannas in biogeographical barrier for Amazonian birds. Guyana; 3 species typical of campina/campinarana; 14 spe- cies associated with the tepuis and reported from mountains As a general rule, terra fi rme forests east of the Rio Branco adjacent to Roraima in Venezuela or Guyana; and 8 Nearc- show avian biogeographical affi nities with the Guianan area tic migrants that regularly visit southern Venezuela, southern of endemism, similar in terms of species composition to the Guyana, or Manaus. Note that seven of the species included in avifauna of Manaus (see Cohn-Haft et al. 1997) and Guyana Table 3 would also represent new Brazilian records, showing (see Braun et al. 2000). Forests west of the Rio Branco are the need for general collections in this poorly sampled region. more closely related to western Amazonia (with elements of Imeri and Napo areas of endemism), and have species com- AVIAN DIVERSITY position similar to that of areas in the state of Amazonas in Venezuela (Zimmer and Hilty 1997, Hilty 2003) or the upper The diversity of birds in the state of Roraima is a result of Rio Negro (MCH and LNN, unpubl. data). high habitat heterogeneity (beta-diversity) and regional ende- mism (gamma-diversity), and understanding both these factors Savanna-like open habitats. Two distinctive but physiog- is central to recognizing the patters of avian diversity within nomically similar habitats dominated by herbaceous commu- the Rio Branco basin. Habitat heterogeneity is responsible for nities are present in Roraima. Savannas and campinas struc- 60% of the bird species found in Roraima (446 species occur turally resemble one another, but have a radically different exclusively in single habitats; Table 1). As expected, beta–di- geological history and plant species composition (Brown and versity is highest in ecotone areas, where major habitats en- Prance 1987). Although both habitats have several species of counter one another such as within the savanna-dry forest-terra birds in common, which do not occur in any other habitats, fi rme interface. Regional endemism (species in similar habitats such as Formicivora grisea, Schistochlamys melanopis, and related to different biogeographical regions) plays a major role Emberizoides herbicola, they also have many species that within terra fi rme forests, accounting for more than a third of seem to occur in one habitat but not in the other. Examples the species richness found in this habitat (see Table 1). of bird species that occur in savannas but not in campinas The diversity of birds in a given site (alpha-diversity) is include: Cercibis oxycerca, Theristicus caudatus, Rampha- also highest within terra fi rme forests. Although we lack in- stos toco, Polystictus pectoralis, Turdus nudigenis, Anthus tensive surveys at single terra fi rme forest localities, we esti- lutescens, Ammodramus humeralis, Parula pitiayumi, Icterus mate that between 260 and 280 bird species probably occur at croconotus, Sturnella magna, and Euphonia fi nschi. Whereas any given site with continuous forest (Table 1 and 2), a value species typical of campinas, but not present in savannas in- similar to other terra fi rme forests throughout the Amazon clude: Crypturellus erythropus, Polytmus theresiae, Elaenia (Cohn-Haft et al. 1997). rufi ceps, Thamnophilus punctatus, Myrmotherula cherriei, Myrmeciza disjuncta, Heterocercus fl avivertex, and Tachy- BIOGEOGRAPHICAL AFFINITIES: phonus phoenicius. PATTERNS OF AVIAN DISTRIBUTION IN RORAIMA The savannas of Roraima-Rupununi represent the largest continuous savanna in Amazonia. Its fl ora and avifauna are re- Below, we present the patterns of avian distribution in Ro- ported to be more closely related to the llanos of Colombia and raima associated with major habitats (we excluded montane Venezuela than to other Amazonian savannas (Huber 1995, forests and tepuis because we did not sample those habitats Silva 1995), such as Sipaliwini, Amapá, or Alter do Chão. and have little to add to the existing literature). Robbins et al. (2004) presented a list of bird species from these Terra fi rme forest. The Rio Branco, which dissects Roraima areas, including the cerrados of central Brazil, showing that in an eastern and a western half, also seems to separate many the Gran Sabana represents a subset of the Roraima-Rupununi closely related taxa of forest birds on its opposite banks (Table savannas, whereas Sipaliwini is a subset of the savannas of 2). Although the general pattern suggests the importance of Amapá, more closely related to the cerrados of central Brazil. the Rio Branco as a biogeographical barrier, lack of studies on opposite sides of the river prevents us from specifying if the South of Caracaraí, at about the same latitude where sa- river itself represents the barrier, or whether large expanses of vannas are replaced by forests, large areas of poorly drained habitats other than terra fi rme forests also function as barri- sandy soils are covered by campina/campinarana. These hab- ers for those birds. The margins of the lower Rio Branco are itats extend westwards reaching the upper Rio Negro, creating mostly covered by várzea, and further north large expanses of the largest continuous area of white-sand forests in the Neo- campina, campinarana, and savanna also prevent terra fi rme tropics. In general, campinas are patchily distributed within forests from opposite sides of the Rio Branco from coming Amazonia, but their avifaunas seem to remain surprisingly into direct contact one with another (Figure 3). Whether the uniform. The campinas of Roraima present clear avian simi- Rio Branco itself or the sum effect of the river in conjunction larities with other well studied, isolated campinas elsewhere with other habitats are responsible for the presence of contact in Amazonia, such as those present near Manaus (MCH and 212 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres

LNN, unpubl. data), Jaú National Park (Borges 2004), and (45%) or natural reserves (7%) (Ministério do Meio Ambiente those of southern Venezuela (Zimmer and Hilty 1997). 2001). Indian Reservations are concentrated in western and ex- treme southeastern Roraima, protecting areas of montane and Riverine forests. Soil composition, fl ooding regimes, and sub-montane forests and lowland terra fi rme forests, respec- rainfall are important in defi ning habitats along the margins of tively. Fortunately, these areas are well protected from immedi- the Rio Branco. While the upper Rio Branco, located within ate development. Natural reserves are mostly restricted to areas the savanna domain, is margined by narrow stretches of gal- near Caracaraí, but entire habitats such as savannas, dry forests, lery forests, the lower Rio Branco, south of Caracaraí, have gallery forests, and várzeas, are inadequately protected. extensive tracts of várzea forests along the river. Gallery and várzea forests differ in bird species composition. Examples Outside protected areas, the state of Roraima is experienc- of bird species that inhabit várzea forests but are absent from ing serious environmental problems. At present, savannas, gallery forest include Leucopternis schistaceus, Amazona fes- natural grasslands, and gallery forests are being converted to tiva, Phaethornis rupurumii, Thamnophilus nigrocinereus, soybean (Gianluppi and Smiderle 2005), acacia (Arco-Verde Myrmmtherula klagesi, Cercomacra nigpescens, Myrmoborus et al. 2005), and rice plantations (Cordeiro 2005) at alarming lugubris, Xiphorhynchus kienerii, Synallaxis propinqua, Cra- rates. The conversion of gallery forests into rice fi elds is of nioleuca vulpina, Serpophaga hypoleuca, Stigmatura napen- special concern to the survival of Synallaxis kollari, a gallery sis, Schiffornis major, and Conirostrum bicolor. On the other forest specialist endemic to northeastern Roraima. hand, among the species that occur in gallery forests but are absent from várzea we can mention Phaethornis hispidus, In addition, natural savannas are subject to anthropogen- Picumnus spilogaster, Picumnus cirratus, Lepidocolaptes ic fi res, which affect 38% of the entire area of the savannas souleyetii, Synallaxis kollari, Myiopagis viridicata, Inezia every year (Barbosa and Fearnside 2005). An increase in the caudata, Poecilotriccus sylvia, Conirostrum speciosum, and frequency of burning in areas well adapted to fi re, such as Basileuterus fl aveolus. The avifauna of the gallery forests is the savannas, can have strong effects on the structure and more closely related to the avifauna present in dry forests, dynamics of the plant community (Miranda and Absy 1997), whereas the avifauna of the várzeas is similar to that found in and has been responsible for the spread of fi res in habitats not fl ooded forests along white-water rivers elsewhere in Amazo- adapted to natural burnings, such as humid forests (Barbosa nia, such as the Amazon and Madeira rivers. and Fearnside 1999). The effects of anthropogenic fi res on the The Rio Branco is unique in representing a white-water avian communities of the savannas have not been studied in river surrounded by black-water rivers (for ex., the Negro, Roraima, and deserve investigation. Jauaperí, Anauá, Agua Boa do Univini, and Catrimani riv- ers). In general, bird communities found in várzea along the Environmental problems in the southern part of the state Rio Branco are similar in species composition to the avifau- are linked to human settlements and state-sponsored coloniza- na inhabiting tall igapó on the lower Rio Negro (such as the tion programs in forested areas. As a general rule, peasants Anavilhanas archipelago). On the other hand, and somewhat are allocated to small areas of land, which are logged and surprisingly, the lower Rio Branco has several elements typi- subsequently burned to promote subsistence agricultural ac- cal of white-water rivers, absent from the entire Rio Negro tivities for a couple of years (Fearnside 1997). However, the basin. Species such as Synallaxis propinqua, Cercomacra ni- unsuitability of these areas for agriculture has been known for grescens, Stigmatura napensis, Serpophaga hypoleuca, and decades (Fearnside 1985), and after a few years of slash-and- Conirostrum bicolor seem to have isolated populations along burn agriculture, these now sterile lands are abandoned, pro- the Rio Branco (Pacheco 1995a, Naka et al. in prep.). ducing a vicious cycle creating social unrest that the govern- ment attempts to solve by encouraging new settlements under Apart from gallery forest and várzea, riverine environ- similar conditions. The situation in southeastern Roraima is ments fl ooded by black-water rivers are covered by igapó, critical, where large areas of abandoned pastures dominate the which occurs along most Rio Branco tributaries. Tall igapó landscape. In fact, our own research activities near São Luiz, has an avifauna similar to that of mature várzea forest, where- São João da Baliza, and Caroebe were obstructed by diffi culty as low igapó, locally known as chavascal, has bird species accessing undisturbed primary forests. in common with white-sand forests (campina/campinarana), The present development model in Roraima is clearly un- such as Polytmus theresiae, Thamnophilus amazonicus, Myr- sustainable, and the lack of concerted public policies is exac- motherula cherriei, Hemitriccus inornatus, Heterocercus fl a- erbating the situation. Environmental agencies should work vivertes, and Hylophilus brunneiceps. together with development organizations, focusing on the long-term sustainability of economic activities throughout the CONSERVATION state. We suggest creating natural reserves in neglected habi- tats, such as savannas, gallery and dry forests, and the creation The state of Roraima is unique in Brazil in that more than of large areas of National Forests (FLONAs) or Sustainable half of its territory is protected either by Indian Reservations Development Reserves (RDSs) in the várzeas and forests in The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 213

the southeastern corner of the state, where sustainable activi- Arco-Verde, M. F., H. Tonini, and M. Mourão Júnior (2005) A ties could be developed and controlled, as an alternative to SILVICULTURA NAS SAVANNAS DE RORAIMA, P. 195-200. IN: uncontrolled logging and disordered human settlement. If the R. I. BARBOSA, H. A. M. XAUD, AND J. M. COSTA E SOUZA government of Roraima fails to work toward more sustainable (EDS.) SAVANAS DE RORAIMA: ETNOECOLOGIA, BIODIVERSIDADE E economic activities, this region may lose an important part of POTENCIAIS AGROSSILVIPASTORIS. BOA VISTA: FEMACT-RR. its biodiversity in the next few years. Barbosa, R. I. (1997) Distribuição das Chuvas em Roraima, p. 325-335. In: R. I. Barbosa, E. J. G. Ferreira, and E. G. ACKNOWLEDGMENTS Castellón (eds.) Homem, ambiente e ecologia no estado de Roraima. Manaus: INPA. We are especially indebted to the Instituto Nacional de Pesquisas da Amazônia (INPA) in Roraima; in particular to _____ and P. M. Fearnside (1999) Incêndios na Amazônia R. I. Barbosa, V. Lauriola, and S. P. do Nascimento, for pro- Brasileira: estimativa da emissão de gases do efeito es- viding logistical support during our expeditions, including the tufa pela queima de diferentes ecossistemas de Roraima use of the Institute’s vehicle and lodgings. We are also grateful na passagem do evento “el niño” (1997/98). Acta Ama- to the Instituto Brasileiro do Meio Ambiente e dos Recursos zonica 29: 513-534. Renováveis (IBAMA) in Roraima for permitting collecting _____ and P. M. Fearnside (2005) The frequency and area and access to parks and other reserves under its jurisdiction. burned in the Roraima savannas of Brazilian Amazonia. In particular, we want to extend our most sincere gratitude to Forest Ecology and Management 204: 371-384. A. Galdino for his constant support in Caracaraí and for invit- ing us to participate in two expeditions sponsored by the Insti- _____ and E. J. G. Ferreira (1997) Historiografi a das expedi- tute. We would also like to thank A. F. Coutinho Mello, O. de ções científi cas e exploratórias no vale do Rio Branco, p. Souza Neves Jr., I. A. Rocha Santos, G. Pallazi, M. Kluppel, 193-216. In: R. I. Barbosa, E. J. G. Ferreira, and E. G. and C. Galvão da Silva for allowing and supporting our re- Castellón (eds.) Homem, ambiente e ecologia no estado search activities. The National Aviary provided funds for two de Roraima. Manaus: INPA. of our expeditions in northeastern Roraima. MCH, LNN, and MFT are grateful to Mr. Chagas, at Bar Tesão, for providing _____ and I. S. Miranda (2005) Fitofi sionomias e diversida- shelter during our work at Serra do Tracajá. LNN and MFT are de vegetal das savannas de Roraima, p. 61-78. IN: R. I. also grateful to M. O. de Albuquerque for introducing them to BARBOSA, H. A. M. XAUD, AND J. M. COSTA E SOUZA (EDS.) Roraima in March 2001. J. Mazar Barnett, L. Mestre, P. Coop- SAVANNAS DE RORAIMA: ETNOECOLOGIA, BIODIVERSIDADE E PO- mans, and M. Braun joined us during some of our expeditions, TENCIAIS AGROSSILVIPASTORIS. BOA VISTA: FEMACT-RR. and we thank them for their companionship in the fi eld. We _____, J. M. COSTA E SOUZA, AND H. A. M. XAUD (2005) Sa- are thankful to a S. Kenney (AMNH), D. Willard (FMNH), vanas de Roraima: referencial geográfi co e histórico, P. E. Bauerfeind and A. Gamauf (NMW), M. A. Raposo and J. 11-19. IN: R. I. BARBOSA, H. A. M. XAUD, AND J. M. B. Nacinovic (MNRJ), A. Aleixo (MPEG), and L. F. Silveira COSTA E SOUZA (EDS.) SAVANNAS DE RORAIMA: ETNOECOLO- (MZUSP), for providing information on specimens under their GIA, BIODIVERSIDADE E POTENCIAIS AGROSSILVIPASTORIS. BOA care. Thanks to I. Macedo for specimen preparation and manu- VISTA: FEMACT-RR. script corrections. C. Bechtold, S. Claramunt, G. Ferraz, D. Lane, J. V. Remsen Jr., and two anonymous reviewers made Berry, P. E., O. Huber, and B. K. 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fl oristic composition of 26 cerrado areas in Brazil. Ed- Sick, H. (1965) Jacus (Penelope) da região amazônica (Aves, imburgh Journal of Botany 49: 235-250. Cracidae). Papéis Avulsos de Zoologia 17: 9-16.

Remsen, J. V., Jr., A. Jaramillo, M. A. Nores, J. F. Pacheco, _____ (1997) Ornitologia brasileira. Ed. Nova Fronteira, M. B. Robbins, T. S. Schulenberg, F. G. Stiles, J. M. C. Rio de Janeiro, Brazil. da Silva, D. F. Stotz, and K. J. Zimmer (2005) A clas- Silva, E. L. S. (1997) A vegetação de Roraima, p. 401-415. sifi cation of the bird species of South America. Ameri- In: R. I. Barbosa, E. J. G. Ferreira and E.G. Castellón can Ornithologists’ Union. . Disponível em http://www. (eds.) Homem, ambiente e ecologia no estado de Rorai- museum.lsu.edu/~Remsen/ SACCBaseline.html (acesso ma. Manaus: INPA. em 09/06/2006). Silva, J. M. C. (1995) Biogeographical analysis of the South _____ and T. A. Parker, III (1985) Contribution of river-cre- American cerrado avifauna. Steenstrupia 21: 49-67. ated habitats to bird species richness in Amazonia. Bio- tropica 15: 223-231. _____ (1998) Birds of the Ilha de Maracá, p. 211-229. In: W. Milliken and J. Ratter (eds.) Maracá: the biodiversity Ridgely, R. S. and G. Tudor (1989) The Birds of South Amer- and environment of an Amazonian rainforest. London: ica, Vol. 1. The Oscine Passerines. Austin: University John Wiley and Sons. of Texas Press. _____ and D. C. Oren (1990) Resultados de uma excursão or- _____ and G. Tudor (1994) The Birds of South America, Vol. nitológica à ilha de Maracá, Roraima, Brasil. Goeldiana 2. The Suboscine Passerines. Austin: University of Tex- Zoologia 5: 1-8. as Press. _____ and E. O. Willis. (1986) Notas sobre a distribuição de Robbins, M. B., M. J. Braun, and D. W. Finch (2004) Avifauna quatro espécies de aves da Amazônia brasileira. Boletim of the Guyana southern Rupununi, with comparisons to Museu Paraense Emílio Goeldi, Série Zoologia 2: 151- other savannas of northern South America. Ornitologia 158. Neotropical 15: 173-200. Snethlage, E. (1914) Catálogo das aves amazônicas. Boletim Robinson, S. K. and J. Terborgh (1997) Bird community dy- Museu Paraense Emílio Goeldi 8: 1-465. namics along primary successional gradients of an Ama- zonian whitewater river. Ornithological Monographs Snow, D. W. (2001) Family Momotidae (motmots), p. 264- 48: 641-672. 285. In : J. del Hoyo, A. Elliot, and J. Sargatal, (edq.) Handbook of the Birds of the World, vol. 6: mousebirds Ruschi, A. (1961) A coleção viva de Trochilidae do Museu to hornbills. Barcelona: Lynx Edicions. de Biologia Prof. Mello Leitão, nos anos de 1934 até 1961. Boletim do Museu de Biologia Professor Mello Spix, J. B. von (1824-25) Avium species novae, quas in iti- Leitão no. 30. nere per Brasiliam annis 1817-20 collegit et descripti. 2 vols. Monachii: Hubschmann. Santos, M. P. D. (2005) New records of birds from the Brazi- Stotz, D. F. (1997) Levantamento preliminar da avifauna em lian state of Roraima. Bulletin of the British Ornitholo- Roraima, p. 581-608. In: R. I. Barbosa, E. J. G. Ferreira, gists’ Club 124: 223-226. and E. G. Castellón (eds.) Homem, ambiente e ecologia Schattuck, E. C. (1926) Observation on the Rio Branco, the no estado de Roraima. Manaus: INPA. Uraricoera and Parima rivers, p. 261-283. In: Medical _____, R. O. Bierregaard, M. Cohn-Haft, P. Petermann, J. Report of the 5th. Hamilton Rice Expedition to the Ama- Smith, A. Whittaker, and S. V. Wilson (1992) The status zon. Cambridge: Harvard University Press. of North American migrants in central Amazonian Bra- Schlegel, H. (1864) Muséum d’Histoire Naturelle des Pays- zil. Condor 94: 608-621. Bas: revue méthodique et critique des collections dépo- Teixeira, D. M., D. C. Oren and R. C. Best (1986) Notes on sées dans cet etablissemelt, 3 (Monographie Psittaci). Brazilian Seabirds, 2. Bulletin of the British Ornitholo- Leiden: E.J. Brill. gists’ Club 106: 74-77.

Sclater, P. L. (1874) On the species of the genus Synallaxis of Tostain, O., J. L. Dujardin, C. Érard, and J. M. Thiollay. 1992. the family Dendrocolaptidae. Proceedings of the Zoo- Oiseaux de Guyane. Societé d’Études Ornithologiques. logical Society of London: 2-28. Brunoy: Muséum National d’Histoire Naturelle, Labora- toire d’Écologie Générale. Short, L. L. and J. F. M. Horne (2002) Family Capitonidae (barbets), p. 140-219. In: del Hoyo, J., A. Elliot, and J. Trolle, M. and B. A. Walther (2004) Preliminary observations Sargatal (eds.) Handbook of the Birds of the World, vol. 7: in the Rio Jauaperí region, Rio Negro basin, Amazonia, jacamars to woodpeckers. Barcelona: Lynx Edicions. Brazil. Cotinga 22: 81-85. 218 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres

Vaurie, C. (1980) Taxonomy and geographical distribution _____ (1940) Studies of Peruvian Birds, no. 34. The genera of the Furnariidae (Aves, Passeriformes). Bulletin of the Todirostrum, Euscarthmornis, Snethlagea, Poecilotric- American Museum of Natural History 166: 1-357. cus, Lophotriccus, Myiornis, Pseudotriccus, and Hemit- riccus. American Museum Novitates 1066: 1-23. Whittaker, A. (1995) First record of Coccyzus pumilus for Brazil (Cuculiformes: Cuculidae). Ararajuba 3: 81. _____ (1941a) Studies of Peruvian Birds, no. 36. The genera Elaenia and Myiopagis. American Museum Novitates _____ (1996) First records of Orange-breasted Falcon Falco 1108: 1-23. deiroleucus in central Amazonian Brazil, with short be- havioral notes. Cotinga 6: 65–68. _____ (1941b) Studies of Peruvian Birds, no. 37. The genera Sublegatus, Phaeomyias, Camptostoma, Xanthomyias, Willis, D. (2003) Evidence for the occurrence of Pale-eyed Phyllomyias, and Tyranniscus. American Museum No- Pygmy-Tyrant Atalotriccus pilaris in Brazil. Ararajuba vitates 1109: 1-25. 11: 145. _____ (1941c) Studies of Peruvian Birds, no. 39. The genus Winkler, H. and D. A. Christie (2002) Family Picidae (wood- Vireo. American Museum Novitates 1127: 1-20. peckers), p. 296-558. In: del Hoyo, J. del, A. Elliott and J. Sargatal (eds.) Handbook of the Birds of the World, _____ (1942a) Studies of Peruvian Birds, no. 40. Notes on vol. 7: jacamars to woodpeckers. Barcelona. Lynx Edi- the genus Veniliornis. American Museum Novitates tions 1159: 1-12.

Worbes, M. (1997) The forest ecosystem of the Floodplains, _____ (1942b) Studies of Peruvian Birds, no. 42. The genus p. 223-265. In: W.J. Junk, (ed.) The Central Amazon Polioptila. American Museum Novitates 1168: 1-7. Floodplain: ecology of a pulsing system. Berlin: Spring- er. _____ (1943a) Studies of Peruvian Birds, no. 45. The gen- era Tersina, Chlorophonia, Tanagra, Tanagrella, Chlo- Zimmer, J. T. (1933) Studies of Peruvian Birds, no. 9. The rochrysa, and Pipraeidea. American Museum Novitates formicarian genus Thamnophilus. Part I. American Mu- 1225: 1-24. seum Novitates 646: 1-22. _____ (1943b) Studies of Peruvian Birds, no. 47. The genus _____ (1936) Studies of Peruvian Birds, no. 24. Notes on Tangara. Part II. American Museum Novitates 1246: Pachyramphus, Platypsaris, Tityra, and Pyroderus. 1-14. American Museum Novitates 894: 1-26. _____ (1944) Studies of Peruvian Birds, no. 48. The genera _____ (1937a) Studies of Peruvian Birds, no. 27. Notes on Iridosornis, Delothraupis, Anisognathus, Buthraupis, the genera Muscivora, Tyrannus, Empidonomus, and Compsocoma, Dubusia, and Thraupis. American Mu- Sirystes, with further notes on Knipolegus. American seum Novitates 1262: 1-21. Museum Novitates. 962: 1-28. Zimmer, K. J. and S. Hilty (1997) Avifauna of a locality in the _____ (1937b) Studies of Peruvian Birds, no. 28. Notes on Upper Orinoco drainage of Amazonas, Venezuela. Or- the genera Myiodynastes, Conopias, Myiozetetes, and nithological Monographs 48: 865-885. Pitangus. American Museum Novitates 963: 1-28. _____ and M. L. Isler (2003) Family Thamnophilidae (typical _____ (1938) Studies of Peruvian Birds, no. 29. The genera antbirds) p. 448-681. In: del Hoyo, J., A. Elliott and J. Myiarchus, Mitrephanes, and Cnemotriccus. American Sargatal (eds.) Handbook of the Birds of the World, vol. Museum Novitates 994: 1-32. 8: broadbills to tapaculos. Barcelona: Lynx Editions.

_____ (1939a) Studies of Peruvian Birds, no. 33. The gen- _____, A. Whittaker, and D. F. Stotz (1997) Vocalizations, be- era Tolmomyias and Rhynchocyclus with further notes havior and distribution of the Rio Branco Antbird. Wil- on Ramphotrigon. American Museum Novitates 1045: son Bulletin 109: 663-678. 1-23.

_____ (1939b) A new subspecies of Inezia subfl ava from the neighborhood of Mt. Duida, Venezuela. Proceedings of the Biological Society of Washington 52: 167-170. The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 219 References Ornithological collections Evidence TFTF TFVZTF SF DF CAM TF SF CAMVZ DF GF CAMTFCAM TF (WAM)/(GUI?) WAM RW S RW WAMR R W TW #, 2, 28, 42, 50, 54, 57 S T SVZ CAMTF SF DF #, 2, 28, 50, 54 TF T 28, 50 TTF MF S #, 28, 37, 42, 50, 54, 57VZ 54 28, 50, 54, 57 #, 2, 11, VZ IG CAM TTF DF S MPEG, MZUSP #, 28, 36, 37, 50, 54 S CAMSAV T STF MF S #, 2, 28, 50, 54 S S MPEG, MZUSP FMNH, LACMNH, MNHW, GUIW CP T #, 2, 23, 28, 37, 42, 50, 54W# S #, 2, 28, 37, 45, 50, 54 21, 28, 37, 42, 50, 54, 57 #, 2, 11, GUI S #, 28, 45, 50, 54, 57vagrant #, 42, 45, 54, 57 MPEG S MNHW, S 2, 22, 37, 42, 50 S RW S MPEG, MZUSP FMNH, LACMNH, MNHW, T CP, MPEG, MZUSP FMNH, LACMNH, MNHW, 28, 42, 50, 54 S #, 21, 37, 50, 51, 57 MPEG WR 48, 57 11, MNHW 21, 28, 37, 42, 50, 57 #, 2, 11, S 21, 28, 42, 50, 54, 57 T #, 11, RW R MPEG #, 2, 27, 28, 37, 42, 49, 50, 54 T RW MZUSP VZ S 28, 41, 50, 54, 57 #, 11, W MPEG, MZUSP FMNH, LACMNH, MNHW, MPEG, MZUSP FMNH, MNHW, CP, W FMNH, MPEG, MZUSP MNHW CP, LACMNH MPEG, LACMNH, MZUSP MPEG, MZUSP, FMNH, MNRJ, MNHW, #, 37 S MPEG CP, 45 CP #, 28, 45, 50, 54, 57 S S S #, 28, 42, 50, 54, 57 S T #, 28, 42, 50, 54, 57 #, 42 13, 28, 37, 50, 54 MNHW P MPEG 28, 50, 54 #, 28, 50 28, 50 FMNH, MZUSP MNHW MZUSP MPEG Family and bird species TINAMIDAE tao Tinamus Habitat Dist/Migr. Tinamus major Tinamus guttatus Tinamus cinereus Crypturellus soui Crypturellus undulatus Crypturellus erythropus Crypturellus variegatus Crypturellus ANATIDAE viduata Dendrocygna autumnalis Dendrocygna Neochen jubata Cairina moschata Amazonetta brasiliensis CRACIDAE Ortalis motmot Penelope marail Penelope jacquacu Aburria cumanensis Mitu tomentosum Crax alector ODONTOPHORIDAE Colinus cristatus Odontophorus gujanensis PODICIPEDIDAE dominicus Tachybaptus Podilymbus podiceps PELECANIDAE Pelecanus occidentalis PHALACROCORACIDAE brasilianus Phalacrocorax ANHINGIDAE Anhinga anhinga ARDEIDAE lineatum Tigrisoma Agamia agami Cochlearius cochlearius Zebrilus undulatus Botaurus pinnatus Ixobrychus exilis Appendix 2. List of bird species recorded in the state Roraima. 220 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres References Ornithological collections Evidence W RW RW AA W RW RW RW RW RW SAVVZ IGW45 SAV RW S RW S RW S RW 28, 50 #, 28, 50 S #, 2, 28, 45, 50, 54, 57 S #, 2, 28, 49, 50, 54 X S #, 2, 28, 37, 45, 50, 54, 57SAV #, 2, 28, 42, 50, 54, 57TF DF #, 28, 50, 54, 57 SX S #, 2, 28, 37, 45, 50, 54TF DF SAV #, 2, 28, 42, 45, 50, 54 T T SR MPEG S #, 23, 28, 37, 49, 50, 54 #, 28, 42, 50, 54, 57 FMNH, MNHW TTF MF S FMNH #, 2, 28, 42, 45, 50, 54DF S #, 2, 37, 42, 57 S MNHW FMNH MZUSP #, 12, 13, 50, 54 FMNH, MNHW #, 2, 28, 37, 42, 50, 54, 57 S S S #, 2, 28, 42, 45, 50, 57 FMNH, MPEG, MZUSP NEA FMNH #, 28, 37, 42, 50, 54, 57 #, 28, 34, 37, 42, 50, 54 S #, 28, 37, 45, 50, 54, 57 #, 2, 28, 50, 54, 57 MNHW MNHW S #, 2, 28, 42, 50, 54, 57 FMNH FMNH #, 28, 37, 50, 54, 55, 57 S 28, 50, 57 MNHW LACMNH, MNHW MNHW 37 #, 11, MNHW MPEG MNHW CP, SAV TF MF SAV RAA SAV AA SAV RW TFTFVZ DF SAV NEASAVTFTFTFVZDF VZ NEA STF S #, 28, 50, 57 T S #, 2, 28, 37, 42, 50, 54, 57 #, 28, 37, 50 S #, 28, 50, 54, 57 #, 2, 28, 50, 54 #, 28, 50, 54, 57 MPEG, MZUSP FMNH, LACMNH, MNHW, 37 S S 12, 13 28, 50 #, 54 #, 37, 42, 50, 51, 57 28, 50 T MPEG, UMMZ LACMNH, MNHW, MPEG # 28, 50 MZUSP MNHW, MNHW MPEG catus fi Family and bird species Ixobrychus involucris Habitat Dist/Migr. Nycticorax nycticorax Butorides striata Bubulcus ibis cocoi Ardea alba Ardea pileatus Pilherodius thula Egretta caerulea Egretta THRESKIORNITHIDAE oxycerca Cercibis Mesembrinibis cayennensis Phimosus infuscatus Theristicus caudatus Platalea ajaja CICONIIDAE Ciconia maguari Jabiru mycteria Mycteria americana CATHARTIDAE Cathartes aura Cathartes burrovianus Cathartes melambrotos Coragyps atratus papa Sarcoramphus PANDIONIDAE Pandion haliaetus ACCIPITRIDAE Leptodon cayanensis uncinatus Chondrohierax Elanoides for Gampsonyx swainsonii Elanus leucurus sociabilis Rostrhamus Harpagus bidentatus Harpagus diodon Ictinia plumbea buffoni Circus Accipiter poliogaster Accipiter superciliosus Accipiter bicolor Geranopsiza caerulescens Leucopternis schistaceus Leucopternis melanops The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 221 .) in litt J. Mazar Barnett ( References Ornithological collections Evidence TFTFTF CAMVZTF CAM GF SAVVZ CAMTF SF VZSAVSAV S S S T T NEA T #, 28, 37, 50, 54 S #, 37 #, 2, 28, 50, 54 #, 2, 28, 50, 54, 57 50 S #, 2, 24, 37, 42, 50, 54 60 #, 24, 28, 37, 42, 50, 54 MNHW MPEG MPEG, MZUSP FMNH, LACMNH, MNHW, MNHW MZUSP FMNH, MNHW, W RW TFSAVWWW SAVW W# RW (GUI)W R S S #, 2, 28, 37, 42, 50, 54, 57 #, 2, 28, 37, 42, 50, 54, 57 S T T # #, 2, 28, 42, 50, 54, 57 T S MZUSP MNHW, T MPEG, MZUSP LACMNH, MNHW, #, 28, 50, 54 #, 28, 50, 54 #, 50 #, 28, 50, 54 50 LACMNH, MPEG, MZUSP MPEG TF RSAV RSAV RW XSAVAAVZ SF SAV TF SFDFSAVTFTFVZTF DFTF S S S NEA S T SVZ IGTF TVZTF #, 2, 28, 35, 37, 42, 50, 54, 57 S S 28, 37, 50, 54X #, 2, 28, 37, 42, 50, 54 TXVZ DF #, 2, 28, 50, 54 #, 24, 28, 35, 37, 50, 54, 57SAV T #, 2, 24, 28, 37, 42, 50, 54, 57 #, 2, 24, 28, 37, 49, 50, 54 MZUSP 54, 55 MNHW, S MZUSP FMNH, MNHW, # MPEG FMNH, MNHW, #, 28, 49, 50, 54 MPEG, MZUSP S FMNH, LACMNH, MNHW, S 24, 28, 50 MNHW MPEG FMNH, MNHW, #, 28, 42, 50 T #, 24, 37 FMNH #, 28, 50 T #, 28, 37, 50, 54 S #, 2, 28, 50, 54, 57 S T S T #, 28, 45, 50, 54, 57 T #, 28, 42, 50, 54, 57 T #, 2, 28, 37, 42, 50, 54 #, 2, 28, 50, 54 MNHW FMNH FMNH, MNHW MZUSP MPEG, MZUSP FMNH, MNHW, FMNH, MPEG collis fi avirostris fl gularis fi Micrastur gilvicollis Micrastur mirandollei Micrastur semitorquatus Falco sparverius Falco ru Falco deiroleucus Falco femoralis Falco peregrinus ARAMIDAE Aramus guarauna PSOPHIIDAE Psophia crepitans RALLIDAE schomburgkii Micropygia Aramides cajanea Laterallus viridis Laterallus exilis Porzana albicollis Gallinula chloropus Porphyrio martinica Porphyrio Family and bird species Leucopternis albicollis Habitat Dist/Migr. Buteogallus urubitinga meridionalis Heterospizias nigricollis Busarellus Rupornis magnirostris Buteo albicaudatus Buteo nitidus Buteo swainsoni Buteo brachyurus Buteo albonotatus Morphnus guianensis Harpia harpyja Spizaetus tyrannus Spizaetus melanoleucus Spizaetus ornatus FALCONIDAE Daptrius ater Ibycter americanus Caracara cheriway Milvago chimachima cachinnans Herpetotheres Micrastur ru 222 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres References Ornithological collections Evidence W RW AA R W vagrantR RW NEARXAA SAV S AA SAV DF GF 54, 55 AA SCAM DF #, 2, 21, 28, 42, 50, 54MF S S 56 T S #, 2, 23, 28, 45, 50, 54, 57 #, 28, 42, 50, 54, 57 S FMNH, MPEG, MZUSP S S TEP S #, 23, 42, 50, 54, 57 S #, 2, 28, 50, 54 #, 2, 21, 28, 37, 42, 50, 54 S #, 2, 21, 27, 28, 37, 42, 50, 54, 57 FMNH T #, 28, 42, 50, 54 #, 2, 21, 42, 45, 54, 57 MZUSP MPEG #, 54 41 11, MPEG, MZUSP FMNH, LACMNH, MNHW, FMNH, LACMNH, MZUSP MPEG, MZUSP FMNH, MNHW, FMNH, MPEG FMNH, MPEG, MZUSP CP, MZUSP CP W RW RW RW RW RW RW NEA NEA NEA NEA S NEA S NEA S S T #, 28, 42, 45, 50, 55 #, 23, 28, 50, 54, 55, 57 #, 2, 28, 42, 50, 54, 55 54, 55 54, 55 # FMNH, MPEG MPEG, MZUSP LACMNH, MPEG, MZUSP FMNH, LACMNH, MPEG RVZ IG GFWR AA R W SAV RW RW SAVW SAV W SAV S S RW NEA RW S R TW #, 21, 28, 42, 50, 54, 57 T S #, 37, 50, 54, 57 #, 2, 23, 28, 42, 50, 54, 57 T P T, S #, 54, 55 #, 2, 23, 28, 42, 50, 54, 57 NEA S NEA S NEA #, 23, 42, 54, 57 FMNH, LACMNH, MPEG, MZUSP S FMNH, LACMNH, MPEG, MZUSP S T #, 13, 23, 27, 28, 37, 42, 49, 50 MPEG MNHW, S T FMNH, LACMNH, MPEG, MZUSP #, 23, 28, 42, 50, 54 #, 28, 37, 50 54, 55 42, 49 #, 28, 45, 50, 54, 55 MNRJ, MPEG, MZUSP FMNH, LACMNH, MNHW, FMNH, LACMNH, MPEG, MZUSP FMNH, LACMNH, MZUSP MNHW LACMNH MZUSP collis fi avipes fl JACANIDAE Jacana jacana STERCORARIIDAE parasiticus Stercorarius STERNIDAE Sternula superciliaris Phaetusa simplex RINCHOPIDAE niger Rynchops COLUMBIDAE Columbina passerina Columbina minuta Columbina talpacoti Claravis pretiosa Columba livia Patagioenas speciosa Patagioenas fasciata Tringa solitaria Tringa Actitis macularius Calidris minutilla Calidris fuscicollis Calidris melanotos subru Tryngites Family and bird species HELIORNITHIDAE Heliornis fulica Habitat Dist/Migr. EURYPYGIDAE Eurypyga helias CHARADRIIDAE cayanus Vanellus chilensis Vanellus Pluvialis dominica Charadrius collaris BURHINIDAE bistriatus Burhinus SCOLOPACIDAE Gallinago paraguaiae Gallinago undulata Limosa haemastica Bartramia longicauda melanoleuca Tringa Tringa The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 223 .) in litt References Ornithological collections P. Coopmans ( P. Evidence XVZ IG CAMTF TF IG CAM DFXVZ CAM DF GF SF SAV DF GF SFTFVZ IG TF VZ IG TF VZ IG TF SVZ S DF GF CAMSAV DF CAMSAV T SGF SAV T #, 2, 21, 42, 50, 51, 54 SAV #, 54 #, 2, 21, 28, 42, 50, 54, 57VZ GF CAM IG SAV T S STF MF #, 21, 28, 37, 41, 42, 50, 54TF T S GFSAV S 28, 42, 50, 54 #, 11, TF #, 2, 28, 37, 42, 50, 54 S SVZ IG TF S T FMNH, LACMNH, MPEG, MZUSP FMNH, LACMNH, MPEG, MZUSP 54 TF MF T SMF TEP T 28, 50, 54, 57 #, 11, #, 28, 37, 42, 50, 54, 57 MPEG, MZUSP FMNH, LACMNH, MNHW, CP, T STF #, 28, 37, 45, 50, 54, 57 S #, 2, 5, 13, 28, 37, 42, 50, 54 28, 45, 50, 54, 57 T #, 2, 11, T TEP T #, 2, 5, 28, 37, 42, 50, 54 WAM S #, 2, 5, 13, 28, 37, 42, 49, 50, 54, 57 MZUSP FMNH, LACMNH, MNHW, #, 2, 28, 42, 50, 54, 57 GUI LACMNH, MPEG, MZUSP CP, WAM S T WAM S S MPEG 28, 50, 57 MNHW, MPEG, MZUSP FMNH, LACMNH, MNHW, 49, MPEG, MZUSP FMNH, LACMNH, MNHW, MNHW S #, 2, 13, 14, 25, 37, 50, 51, 53 FMNH, MPEG S CP, CP S MPEG, MZUSP FMNH, LACMNH, MNHW, T T 41, 49 28, 50, 54 11, T #, 2, 5, 28, 37, 46, 49, 50, 54 #, 2, 28, 50, 54, 57 MPEG, MZUSP # MNHW T FMNH, MNHW FMNH, MPEG FMNH, MPEG CP, CP MPEG CP TF VZTFVZTF TFVZ DFTF VZTF VZVZ DF SAV VZDF TFTF WAM (GUI) GUIVZ IG GF S S S50 GF NEA T S SDFVZ SFDF GF #, 2, 28, 38, 42, 50, 54 #, 44 #, 37, 50, 54, 57 SX T T S # #, 2, 3, 28, 50, 54, 57 S T S #, 5, 27, 28, 37, 42, 50, 54 T #, 37, 54 S 28, T AUS #, 14, 37, 57 S S #, 2, 5, 28, 37, 42, 50, 54 FMNH, LACMNH, MPEG, MZUSP AUS? T T S #, 2, 28, 45, 50, 54 MPEG MNHW, MPEG #, 2, 37, 50, 54, 57 S MNRJ, MPEG, MZUSP FMNH, MNHW, #, 28, 42, 45, 50, 54, 57 MPEG #, 28, 42, 50, 54, 57 MPEG, MZUSP FMNH, LACMNH, MNHW, S #, 42, 54 FMNH, INPA MNHW MNHW MPEG LACMNH, MZUSP MPEG MNHW, #, 2, 5, 28, 41, 42, 50, 54, 57 MPEG LACMNH, MZUSP, LACMNH, FMNH, MPEG, MZUSP CP, FMNH, MZUSP Family and bird species Patagioenas cayennensis Habitat Dist/Migr. Patagioenas plumbea Patagioenas subvinacea Zenaida auriculata Leptotila verreauxi Leptotila rufaxilla Geotrygon montana PSITTACIDAE Ara ararauna Ara macao Ara chloropterus Ara severus Orthopsittaca manilata Diopsittaca nobilis Aratinga leucophthalma Aratinga solstitialis Aratinga pertinax picta Pyrrhura egregia Pyrrhura melanura Pyrrhura Forpus passerinus Forpus modestus chrysoptera Brotogeris cyanoptera Brotogeris Nannopsittaca panychlora huetii Touit Touit purpuratus Touit Pionites melanocephala Gypopsitta barrabandi Gypopsitta caica Pionus menstruus Pionus fuscus Amazona festiva Amazona ochrocephala Amazona amazonica Amazona farinosa accipitrinus Deroptyus OPISTHOCOMIDAE Opisthocomus hoazin CUCULIDAE Coccyzus americanus Coccyzus euleri Coccyzus melacoryphus Piaya cayana 224 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres References Ornithological collections Evidence TF CAMVZGFVZ IGXVZ DF SAV TF S S S S #, 28, 42, 49, 50, 54 T T S T #, 6, 28, 42, 50, 54 #, 28, 42, 50, 54, 57 S #, 2, 28, 42, 50, 54 49, 59 #, 2, 27, 28, 42, 50, 54, 57 37 MPEG, MZUSP FMNH, LACMNH, MPEG, MZUSP LACMNH, MPEG, MZUSP LACMNH, MPEG, MZUSP FMNH, LACMNH, MNRJ, MPEG, MZUSP MNHW TF AASAV VZ IG MFDF GF TFMFTFTFTFTFTFDF GF MF SAA SAV SSAVVZ TEP T T 2, 28, 37, 42, 49, 50MF #, 2, 28, 37, 50, 54 # S SVZ TF TF T SVZ IG DF GFTF 39, 49 11, S #, 28, 50 S TAA CAM SAV S S #, 54 MPEG, MZUSP MNHW, TR MPEG FMNH, LACMNH, MNHW, #, 28, 42, 50, 54AA CAM SAV T 28, 37, 50 #, 28, 50, 54VZ SAV #, 28, 50 VZ IG R #, 28, 37, 50 #, 2, 5, 28, 42, 50, 54SAVX S CAM SAV 28, 50 MFWhittaker (in litt) A. SAV CP DF SAV T MPEG S NEATF 49 11, MPEG, MZUSP #, 28, 50, 54, 57 S FMNH, MPEG, MZUSP T 28, 50, 54 LACMNH AUS MNHW #, 5, 28, 37, 50, 54 MNHW T 28, 50 S #, 5, 28, 37, 42, 50, 54 S TEP T #, 28, 50 #, 50, 54, 57 28, 50 S S T #, 5, 28, 37, 42, 50, 54 S S T #, 2 MNHW FMNH, INPA, CP MPEG, MZUSP LACMNH, MNHW, FMNH, INPA, #, 2, 5, 28, 37, 42, 50, 54, 57 T T 41, 49 S #, 50 #, 5, 28, 37, 41, 42, 49, 50, 54 T #, 54, 57 MPEG, MZUSP LACMNH, MNHW, FMNH, INPA, MPEG, MZUSP FMNH, LACMNH, MNHW, FMNH MPEG, MZUSP FMNH, LACMNH, MNHW, CP, CP INPA MPEG FMNH, INPA, pennis fi Family and bird species Piaya melanogaster Habitat Dist/Migr. Coccycua minuta Coccycua pumila major Crotophaga ani Crotophaga naevia Tapera pavoninus Dromococcyx Neomorphus ru TYTONIDAE alba Tyto STRIGIDAE Megascops choliba Megascops watsonii Megascops guatemalae Lophostrix cristata Pulsatrix perspicillata Bubo virginianus Strix virgata Glaucidium hardyi Glaucidium brasilianum Athene cunicularia Rhinoptynx clamator Asio stygius STEATORNITHIDAE Steatornis caripensis NYCTIBIIDAE Nyctibius grandis Nyctibius aethereus Nyctibius griseus CAPRIMULGIDAE pusillus Chordeiles rupestris Chordeiles acutipennis Chordeiles minor Chordeiles leucopyga Nyctiprogne Podager nacunda albicollis Nyctidromus Caprimulgus rufus Caprimulgus longirostris Caprimulgus cayennensis Caprimulgus maculicaudus Caprimulgus nigrescens The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 225 ) in litt. References Ornithological collections J. F. Pacheco and A. Carvalhães ( Pacheco and J. F. Evidence SAV GF DF AA GF DF SAV MFMFTF S TEP T #, 5, 28, 42, 50, 54 S S S 41, 49 11, 41, 43, 49, 54 2, 11, #, 5, 28, 43, 50, 54, 57 FMNH, LACMNH, MPEG, MZUSP FMNH CP, FMNH, MPEG CP MFVZ IGMFTF MFTF CAM VZ TEP S TEP S S T 49 11, S #, 42, 50, 51, 54 49, 54 11, #, 28, 42, 50, 54 #, 2, 50, 54, 57 MPEG, MZUSP CP MPEG, MZUSP CP SAVXMFX DF SFSAV SFTFVZTF SF AUS?MFVZTF SF CAM GF SF TEP SVZ CAMTF TF DF MFMF STF S #, 42, 49, 50MFVZ IG CAMTF DF S TMF TEP (WAM?)/(GUI) 49 11, VZ #, 2, 42, 50, 54TF #, 54 S S S SCAM S #, 2, 28, 42, 50, 54 SMF TMF S TEPMF #, 28, 50, 54, 57 2, 5, 28, 43, 50, 54 #, 2, 5, 28, 50, 54 43, 54VZ IG CAM TEP 2, 28, 42, 49, 50, 54GF DF SAV S 43, 54 MZUSP INPA, STF TEP 41, 43, 54 #, 11, TF CAM STF CAM 49, 54 MZUSP GUISAV FMNH, MZUSP #, 2, 42, 54 SCAM IG 40 11, TEP CP IG SAV TEPSAV S 41, 49 TEP FMNH, MPEG FMNH, MPEG FMNH, MPEG #, 54, 57 MPEG, MZUSP S S 40, 43, 49 11, S S T S MPEG CP, FMNH, MPEG #, 2, 5, 28, 50, 54 49, 54 13, 28, 50 #, 11, CP 43, 54 49 11, #, 28, 37, 42, 50, 54 49 S S 28, 50, 54 #, 11, FMNH, MPEG, MZUSP T S S T CP S #, 54 #, 43, 54 CP 40, 41, 43, 50, 54, 57 #, 11, T CP S FMNH, MPEG # FMNH, LACMNH, MPEG #, 28, 54 CP, 28, 50, 54 #, 2, 11, MPEG, MZUSP FMNH, MNHW, 42, 49, 50, 54 CP FMNH, MPEG FMNH, MPEG CP, CP FMNH, MPEG FMNH, MPEG CP, MPEG FMNH, INPA, MZUSP FMNH, LACMNH, MPEG mbriata fi Amazilia viridigaster Heliodoxa xanthogonys Heliothryx auritus Family and bird species Caprimulgus whitelyi Habitat Dist/Migr. Hydropsalis climacocerca Hydropsalis APODIDAE phelpsi Streptoprocne zonaris Streptoprocne Chaetura spinicaudus Chaetura cinereiventris Chaetura meridionalis Chaetura brachyura montivagus Aeronautes squamata Tachornis Panyptila cayennensis TROCHILIDAE Glaucis hirsutus leucurus Threnetes Phaethornis rupurumii Phaethornis griseogularis Phaethornis ruber Phaethornis hispidus Phaethornis bourcieri Phaethornis superciliosus Doryfera johannae Campylopterus largipennis Campylopterus hyperythrus Florisuga mellivora Colibri delphinae Anthracothorax nigricollis pella Topaza Chrysolampis mosquitus Lophornis ornatus Lophornis chalybeus Lophornis pavoninus notata Chlorestes mellisugus Chlorostilbon Thalurania furcata Hylocharis sapphirina Hylocharis cyanus Polytmus guainumbi Polytmus theresiae Amazilia versicolor Amazilia brevirostris Amazilia 226 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres CP, MPEG, MZUSP CP, b a References Ornithological collections Evidence SF SF MFSAV VZ CAMTF DF TFMFMFTFTFWR SRWR SVZR GF VZR GF TEP T 54 11, (GUI) #, 2, 5, 28, 37, 42, 50, 54, 57VZ DFTF S #, 2, 28, 50, 54 S SGF SF TTFGF T S #, 28, 50, 54, 57 MPEG, MZUSP FMNH, LACMNH, MNHW, VZ DF GF IG 39, 41, 49 11 # CAM S (GUI)TF STF S S #, 2, 28, 50, 54, 57 T S S #, 2, 28, 42, 50, 54, 57 CP TFTF 28, 42, 45, 50, 54, 57 (GUI) #, 2, 5, 28, 42, 50, 54, 57TF SF #, 2, 5, 28, 42, 50, 54, 57 #, 28, 50, 54TF FMNH #, 2, 28, 37, 42, 50, 54, 57TF IG S STF S (GUI) CP TF (GUI) MPEG TTF DF T FMNH, MPEG, MZUSP S TVZ IG MPEG CP, #, 2, 6, 37, 42, 54 S #, 54, 57X FMNH, LACMNH, MZUSP FMNH, LACMNH, MPEG, MZUSP #, 2, 6, 13, 28, 37, 42, 50, 54, 57 WAM S FMNH, MPEG, MZUSP T MPEG, MZUSP FMNH, LACMNH, MNHW, GUI TTF T #, 37, 49VZTF S #, 28, 50, 57 #, 57 MPEG MPEG, MZUSP FMNH, LACMNH, MNHW, #, 28, 50, 54 (WAM) S GUI T (50, 54) S MPEG, MZUSP FMNH, LACMNH, MNHW, S #, 37 S T #, 2, 6, 42, 50, 54 WAM FMNH, MPEG GUI #, 6, 37, 42, 57 28, 50, 54 54 #, 2, 6, 28, 37, 42, 50, 54, 57 MNHW FMNH S S T 54 FMNH, MPEG # T T #, 2, 6, 28, 37, 42, 50, 54, 57 42 (2, 28, 50, 54) #, 11, FMNH, LACMNH, MPEG, MZUSP MPEG, MZUSP FMNH, LACMNH, MNHW, #, 57 MPEG, MZUSP FMNH, LACMNH, MNHW, MNHW MPEG, MZUSP FMNH, LACMNH, MNHW, SAV DFSAV S T #, 28, 37, 42, 50, 54 MPEG, MZUSP LACMNH, MNHW, cauda fi Family and bird species Heliomaster longirostris Habitat Dist/Migr. macrorhynchos Notharchus Calliphlox amethystina TROGONIDAE viridis Trogon violaceus Trogon collaris Trogon personatus Trogon rufus Trogon melanurus Trogon ALCEDINIDAE Ceryle torquatus amazona Chloroceryle americana Chloroceryle inda Chloroceryle aenea Chloroceryle MOMOTIDAE Momotus momota GALBULIDAE Brachygalba lugubris Galbula albirostris Galbula ru Galbula galbula Galbula leucogastra Galbula dea aureus Jacamerops BUCCONIDAE hyperrhynchus Notharchus tectus Notharchus Bucco macrodactylus Bucco tamatia Bucco capensis Nonnula rubecula Monasa atra Monasa nigrifrons Chelidoptera tenebrosa CAPITONIDAE Capito niger Capito auratus RAMPHASTIDAE Ramphastos toco The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 227 References Ornithological collections Evidence TF VZ IG DFTF TFMFTFTF (WAM)/(GUI)TF SF GFVZTF STF SF GF T (WAM)/(GUI)TF SFDF GF #, 2, 28, 37, 42, 50, 54, 57 SGF DF TEP WAMVZ IG TF SF GUI TVZ GF DF GUI (GUI) WAMMF S #, 2, 28, 50, 54, 57 STF WAMTF S S S MPEG, MZUSP LACMNH, MNHW, S 41 11, T 28, 50, 54 # #, 2, 28, 37, 42, 50, 54, 57 28, 50, 54 #, 28, 54 S 51 S S TEP LACMNH, MPEG S S T T GUI T S 28, 41, 42, 50, 54 #, 6, 11, #, 2, 28, 42, 50, 54 #, 6, 28, 37, 49, 50, 54, 78 MPEG, MZUSP T MNHW, # #, 28, 42, 50, 54, 73 S MPEG 41, 49 CP FMNH, MPEG MPEG FMNH, LACMNH, MPEG, MZUSP #, 41, 50, 54 CP, MNHW FMNH, INPA, LACMNH, MPEG, MZUSP MPEG AMNH, FMNH, LACMNH, MZUSP INPA CP FMNH, LACMNH, MPEG TF VZTF TFMFVZ GFVZ IGTF VZ CAMTF VZ DFTF TF CAM DFVZ IGTF SF DF GF VZ IG MFTF VZ IG CAM DFTF SF WAM TEPTF SF STF SVZ DF SF SVZ IG DF GF CAM SAV S SVZ GF DF T SVZ IG #, 2, 6, 28, 42, 50, 54, 57 S S S #, 50, 54TF T T 39, 49, 54 11, TF S # #, 6, 28, 37, 50, 51, 54, 57TFDF GF CAM T #, 42, 54, 57 T #, 2, 6, 28, 37, 42, 50, 54, 57TF IG CAM #, 6, 28, 42, 50, 54, 57 T #, 28, 50, 54MF #, 28, 42, 50, 54, 57 S GUIMF 28, 42, 50, 54, 57 #, 2, 6, 11, TF FMNH, MPEG, MZUSP TTF S GUIVZTF S S MPEG, MZUSP FMNH, LACMNH, MNHW, #, 2, 8, 13, 28, 37, 42, 50, 54, 62, 57, 78 MPEG FMNH, LACMNH, MNHW, TF WAM (WAM) T S T S UMMZ MPEG, MZUSP, FMNH, LACMNH, MNHW, ANSP, FMNH CP, #, 2, 54 FMNH, LACMNH, MPEG, MZUSP CP, FMNH, LACMNH, MPEG, MZUSP MPEG S #, 8, 28, 37, 42, 50, 54 S S T LACMNH, MPEG, MZUSP FMNH, LACMNH, MPEG, MZUSP T TEP #, 8, 28, 37, 42, 50, 54, 62 TEP T T T FMNH, MPEG #, 8, 28, 37, 41, 42, 50, 54, 57 S #, 57 WAM #, 44 #, 28, 42, 50, 54 S T S GUI #, 2, 28, 50, 54 MPEG, MZUSP FMNH, LACMNH, MNHW, S MPEG, MZUSP FMNH, LACMNH, MNHW, CP, MPEG, MZUSP AMNH, FMNH, LACMNH, MNHW, 41, 49 S T 41, 54 11, T MPEG #, 28, 50, 54 T 2, 28, 37, 42, 50, 54 #, 2, 8, 42, 54 MPEG, MZUSP FMNH, MNHW, # MPEG FMNH FMNH, MPEG MPEG MPEG, MZUSP FMNH, LACMNH, MNHW, CP CP FMNH, MPEG, MZUSP nis fi avigula fl avus fl Family and bird species Ramphastos tucanus Habitat Dist/Migr. Ramphastos vitellinus derbianus Aulacorhynchus Selenidera piperivora viridis Pteroglossus azara Pteroglossus aracari Pteroglossus pluricinctus Pteroglossus PICIDAE Picumnus exilis Picumnus spilogaster Picumnus cirratus Melanerpes cruentatus passerinus Veniliornis kirkii Veniliornis af Veniliornis cassini Veniliornis Piculus Piculus chrysochloros Piculus rubiginosus Colaptes punctigula Celeus grammicus Celeus elegans Celeus Celeus torquatus Dryocopus lineatus Campephilus rubricollis Campephilus melanoleucos THAMNOPHILIDAE Cymbilaimus lineatus viridis Frederickena major Taraba Sakesphorus canadensis Thamnophilus doliatus Thamnophilus nigrocinereus Thamnophilus aethiops Thamnophilus murinus Thamnophilus punctatus Thamnophilus amazonicus Thamnophilus insignis Dysithamnus mentalis Thamnomanes ardesiacus Thamnomanes caesius Pygiptila stellaris Myrmotherula gutturalis 228 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres References Ornithological collections Evidence TF548VZ IGTF TF WAMSF GUICAM IGVZ STF STF DF GF CAM TF TMF WAM 49, TF #, VZ IGTF CAM (GUI) SMF FMNH S FMNH S GUI T 49, 50, 54 #, 28, 50, 54 T (WAM)/(GUI)? TEP S #, 2, 8, 42, 50, 54, 57 T S #, 57 S T T TEP S 2, 28, 50, 54, 57 #, 50, 54 # S T S 11 #, 54 T T FMNH, LACMNH, MPEG, MZUSP FMNH #, 54 # FMNH, MPEG 40, 41, 49 11, FMNH, MPEG FMNH, MPEG FMNH, MPEG CP CP MPEG FMNH, INPA, TF VZTF TF GF GF DF CAM SAV TF1VZ IGTF SF SF CAMSF (GUI)VZVZGF TF GF SVZTF T STF (GUI)VZ IG GF CAM #, SIG SF T S S #, 28, 29, 37, 42, 50, 54 T S T FMNH #, 28, 50, 51, 54 S T T # , 57 #, 2, 50, 54 #, 2, 28, 50, 54 T (WAM)/(GUI) S S #, 54 S S MPEG, MZUSP S FMNH, LACMNH, MNHW, T T S T T #, 4, 12, 13, 37, 42, 44, 49, 54, 78 S # T #, 2, 8, 28, 40, 42, 50, 54, 57 #, 8, 28, 42, 50, 54, 57 T MPEG #, 8, 42, 50, 54, 57 FMNH, INPA, #, 44 MPEG, MZUSP LACMNH, MNHW, FMNH, INPA, 54 LACMNH, MPEG T FMNH, MPEG FMNH, MPEG # FMNH, MPEG, MZUSP FMNH, MPEG, MZUSP FMNH FMNH, LACMNH, MPEG, MZUSP MPEG MPEG FMNH, MPEG TF (WAM)?/(GUI) S T #, 57TF (WAM)?/(GUI)TF SMFTF DF GFTF TTF SF #, CAMTFTF (WAM)?/(GUI) S MPEG TEP (GUI) GUI S S 41 S S (GUI) T T #, 8, 13, 28, 42, 49, 50, 54 T 41 S #, 2, 28, 42, 50, 54 S #, 57 S T # 50, 54, 57 2, 57 FMNH, LACMNH, MPEG, MZUSP FMNH CP, LACMNH, MPEG, MZUSP CP FMNH, MPEG FMNH, MPEG INPA MPEG TF CAMTFVZTF CAMTF GUITF (GUI) S T S #, 2, 28, 37, 42, 50, 54, 57 S T S 28, 50, 54 #, 2, 11, T S 42, 54 #, 54 MPEG, MZUSP FMNH, MNHW, 44 FMNH, MPEG CP, 57 FMNH, MPEG, MZUSP FMNH, MPEG MPEG marginatus fi gula fi frons fi Myrmotherula brachyura Myrmotherula ambigua Myrmotherula surinamensis Myrmotherula cherriei Myrmotherula klagesi Myrmotherula guttata Myrmotherula axillaris Myrmotherula longipennis Myrmotherula behni Myrmotherula menetriesii Myrmotherula assimilis Herpsilochmus dorsimaculatus Herpsilochmus roraimae Herpsilochmus ru Family and bird species Myrmotherula haematonota Habitat Dist/Migr. Microrhopias quixensis Microrhopias Formicivora grisea spodioptila Terenura cinerascens Cercomacra tyrannina Cercomacra laeta Cercomacra nigrescens Cercomacra carbonaria Cercomacra Myrmoborus leucophrys Myrmoborus lugubris Myrmoborus myotherinus Hypocnemis cantator Hypocnemoides melanopogon Sclateria naevia ru Percnostola Schistocichla leucostigma Schistocichla saturata Myrmeciza longipes Myrmeciza ferruginea Myrmeciza atrothorax Myrmeciza disjuncta Myrmornis torquata Pithys albifrons Gymnopithys ru Hylophylax naevius Hylophylax punctulatus Hylophylax poecilinotus CONOPOPHAGIDAE Conopophaga aurita GRALLARIIDAE Hylopezus macularius The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 229 FMNH, MNHW, MPEG FMNH, MNHW, c References Ornithological collections Evidence TFTFTF CAMTF CAMTFTF MFTFVZ IG GFTF (WAM)/(GUI)TF (WAM)?/(GUI)TF MF S STF CAM (WAM)/(GUI)?TF MF (WAM)?/(GUI) T SVZ IG DF GF S S SVZ IG #, 9, 28, 37, 42, 50, 54, 57 (WAM)/(GUI)VZ IG CAM #, 2, 28, 50, 51, 54 TTF STF CAM DF GF MF 54 28, 41, 50, 51, 54 #, 9, 11, GF DF 28, 50 11, 28, 50, 54 T (GUI) GUI S #, 2, 9, 28, 42, 50, 54, 57 (GUI) MPEG, MZUSP FMNH, LACMNH, MNHW, T S S #, 42, 57 S S T FMNH, MPEG CP, LACMNH, MPEG T T GUI #, 9, 28, 37, 50, 54 T S #, 28, 50, 54 T S FMNH, LACMNH, MPEG, MZUSP S 2, 28, 50 #, 9, 28, 37, 42, 49, 50, 54 #, 28, 50 #, 2, 9, 28, 37, 42, 50, 54, 78 T T S MPEG T CP, MPEG #, 9, 28, 42, 50, 54 #, 9, 42, 54 FMNH S #, 44 T 41, 42, 50, 54 #, 2, 11, T MPEG, MZUSP MPEG, MZUSP FMNH, LACMNH, MNHW, MNHW, MPEG FMNH, LACMNH, MNHW, LACMNH, MPEG, MZUSP #, 9, 13, 49, 54 INPA LACMNH, MPEG, MZUSP FMNH, INPA, FMNH, LACMNH, MPEG CP, FMNH, LACMNH, MPEG, MZUSP MPEG FMNH TFTFVZGF DF GF CAMSAV TFVZMF (WAM/GUI)VZGF SVZ IG MFVZ TWVZ S #, (9, 37, 50, 51, 54) MF (GUI) T S TEP S #, 9, 37, 41, 42, 54 T 54, 57 S T #, 9, 27, 28, 29, 37, 42, 50, 54, 78 TEP #, 2, 9, 28, 42, 50, 54, 57 S S S TEP S 39, 41 11, MNRJ, MPEG, MZUSP T LACMNH, MNHW, FMNH, INPA, 31 T T #, 28, 50, 54 S MPEG, MZUSP FMNH, MNHW, CP, S #, 33, 37, 42 #, 4, 9, 12, 13, 32, 37, 42, 47, 49, 58 41, 49 11, T FMNH, MPEG, MZUSP T #, 28, 33, 37, 50, 54 # MPEG, MZUSP INPA, 41, 49 11, CP MPEG, LACMNH MPEG, MZUSP LACMNH, MNHW, MPEG MNHW, CP CP TF CAMMFTFTFMFTF TEP S S T TEP #, 2, 28, 54 S 41, 49 11, S T #, 2, 8, 28, 42, 50, 54, 57 S 50 11, # # FMNH, MPEG FMNH, LACMNH, MPEG, MZUSP CP CP MPEG c d gula fi gularis fi Sclerurus caudacutus DENDROCOLAPTIDAE fuliginosa Dendrocincla merula Dendrocincla Deconychura longicauda Sittasomus griseicapillus Glyphorynchus spirurus Nasica longirostris ru Dendrexetastes Hylexetastes perrotii Xiphocolaptes promeropirhynchus certhia Dendrocolaptes picumnus Dendrocolaptes picus Xiphorhynchus kienerii Xiphorhynchus obsoletus Xiphorhynchus pardalotus Xiphorhynchus guttatus Xiphorhynchus Lepidocolaptes souleyetii Lepidocolaptes albolineatus FURNARIIDAE Furnarius leucopus Synallaxis albescens Synallaxis rutilans Synallaxis propinqua Synallaxis macconnelli Synallaxis gujanensis Synallaxis kollari Cranioleuca vulpina Cranioleuca demissa Cranioleuca gutturata Certhiaxis cinnamomeus Roraimia adusta Family and bird species Myrmothera campanisona Habitat Dist/Migr. Myrmothera simplex FORMICARIIDAE Formicarius colma Formicarius analis Chamaeza campanisona SCLERURIDAE Sclerurus mexicanus Sclerurus ru Campylorhamphus sp. Campylorhamphus 230 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres FMNH, INPA, LACMNH, MPEG, MZUSP FMNH, INPA, e References Ornithological collections Evidence VZ GFSAVTFVZVZ TF GF MFMFTFTF MFMFSF SAV (GUI) S S TEP S TEP T S T T #, 10, 54 S #, 2, 42, 54 S #, 30, 42 2, 42, 54 S S A. Whittaker (in litt.) 39, 49 11, S 49 11, T S S #, 10, 28, 50, 54, 57 54 #, 11, T 54 # 54 FMNH, LACMNH, MPEG FMNH, MPEG, MZUSP FMNH, MPEG, MZUSP FMNH, LACMNH, MPEG, MZUSP CP FMNH, MPEG CP CP FMNH, MPEG FMNH, INPA FMNH SAV SAV GF DF SFSAVSAVCAMMFTF AAVZ GF DF SAV MF AUSVZAA GF DF SAV S S TEP S S T #, 37, 42, 54 S TEP S #, 2, 10, 28, 42, 50, 54, 71 S T T T #, 10, 37, 41, 42 T #, 2, 28, 42, 50, 54 S S S #, 10, 28, 37, 50, 54 28, 50 #, 11, 40, 41, 49 11, T T #, 2, 10, 28, 37, 42, 50, 54, 70 #, 28, 50, 54 AMNH, FMNH, LACMNH, MZUSP 41, 49 MZUSP MNHW, #, 31 MPEG, MZUSP MPEG MNHW, MZUSP FMNH, INPA, FMNH, MNHW, CP, MPEG, MZUSP LACMNH, MNHW, AMNH, FMNH, INPA, FMNH CP, CP FMNH, MPEG CP VZ IGGF DF S S T T #, 54 #, 54 FMNH FMNH, INPA VZ TFVZ MFTFTFSFVZ IG CAMTF VZ IG DF CAMTF TF TEP (GUI) S S S T S S S #, 28, 50, 54 41 T T 28, 42, 50, 54, 57 #, 2, 9, 11, S #, 2, 28, 50, 51, 54, 78 #, 2, 28, 50, 54, 57 54 T FMNH, MPEG, MZUSP CP, # MPEG, MZUSP FMNH, LACMNH, MPEG LACMNH, MPEG CP FMNH FMNH TFTFTF CAMTFMFTF (GUI) TEP S S S S S T # 42, 54 S 54 50, 54 39, 49 11, 54 FMNH, MPEG, MZUSP FMNH, MPEG FMNH, LACMNH, MPEG CP FMNH FMNH, MPEG CP, VZ IGGF S T T # #, 10, 42, 68 (13, 28, 50, 54, 78) SAVTF MF WAM S T # CP TFTF DF SF CAM S T #, 28, 42, 50, 54 T #, 28, 50, 54 FMNH, LACMNH, MPEG, MZUSP e pileatus fi aveola fl caudatum avivertex fi ceps fl ava fi fl avogaster fl Polystictus pectoralis Corythopis torquatus Stigmatura napensis Zimmerius gracilipes Phylloscartes chapmani Phylloscartes nigrifrons Mionectes oleagineus Mionectes macconnelli Leptopogon amaurocephalus Sublegatus obscurior Sublegatus modestus Inezia sub Elaenia parvirostris Elaenia cristata Elaenia chiriquensis Elaenia ru Elaenia pallatangae Ornithion inerme Camptostoma obsoletum leucophrys Mecocerculus Serpophaga hypoleuca Phaeomyias murina Capsiempis Myiopagis viridicata Elaenia Lochmias nematura Xenops tenuirostris Xenops minutus TYRANNIDAE Phyllomyias griseiceps elatus Tyrannulus Myiopagis gaimardii Myiopagis caniceps Myiopagis Philydor pyrrhodes Automolus ochrolaemus Automolus infuscatus Automolus roraimae Automolus rubiginosus Automolus ru Inezia caudata Family and bird species Berlepschia rikeri Habitat Dist/Migr. Hyloctistes subulatus Philydor ru Myiornis ecaudatus Lophotriccus galeatus The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 231 References Ornithological collections Evidence DFVZ IGTFCAMCAMMFGF DFGFAAVZ IG DF DF GFSAV TFTF (GUI)VZ IG GFTFVZ SFTF S TEPVZ IG CAMDF GF TF S TTF T T STF #, 12, 13, 49, 61 # STF # S GUIMF T SSF #, 11 (GUI) T S TTF # 41, 49 (GUI) TTF #, 10, 28, 37, 42, 50, 54, 69, 57, 78 S #, 5, 8, 24, 44, 48TF S S S #, 10, 37, 41, 42, 50, 51, 54MF S T S 7, 10, 28, 44, 50, 54VZ SF T T TGF CAM MPEG, MZUSP AMNH, FMNH, LACMNH, MNHW, #, 28, 50, 54 TSF INPA #, 10, 28, 29, 42, 50, 54, 67 #, 28, 42, 50, 51, 54 (GUI) #, 2, 28, 50, 54MF TEP #, 2, 54MF TF S MPEG, MZUSP FMNH, LACMNH, MNHW, SAV CP, S SVZ IG GF S LACMNH, MPEG FMNH, INPA, SIG MF (GUI) TVZ R CP MPEG CP, TEP AMNH, FMNH, LACMNH, MPEG, MZUSP 54 RW VZ CP #, 10, 37, 42 42, 54 R W VZ S S FMNH, MPEG CP, 10, 37, 54MF SF 39, 49 11, S NEA SX FMNH NEA TEP MPEG, LACMNH TX S S AUS? AA VZ SF #, 28, 50, 54 #, 2, 10, 50, 51, 54 #, 28, 50, 54SF TEP T T S 50SF FMNH, MPEG 41 SX #, 28, 42, 50, 54 #, 37, 42, 54WVZ IG R S MPEG STF IG CAM T 54, 55 49 11, MPEG, MZUSP FMNH, MNHW, #, 2, 10, 27, 28, 42, 50, 54 S S MPEG FMNH, MNHW, S MPEG, MZUSP 10, 37, 42 MPEG CP 41, 49 FMNH, MPEG T S #, 10, 28, 49, 50 FMNH, MPEG S #, 10, 28, 37, 42, 50 S #, 2, 28, 50, 54 MPEG, MZUSP S FMNH, LACMNH, MNRJ, MPEG, MZUSP MPEG, MZUSP LACMNH, MNHW, FMNH, INPA, 54 T S MPEG T CP 28, 50, 54 S #, 2, 10, 28, 50, 54, 57 S #, 2, 10, 28, 42, 50, 54, 57 T T CP #, 2, 28, 37, 42, 50, 54, 57 T MPEG, MZUSP FMNH, LACMNH, MNHW, 49, 54 28, 50, 54 #, 2, 11, MPEG, MZUSP FMNH, LACMNH, MNHW, CP MPEG 54 FMNH, LACMNH, MNHW, #, 2, 10, 27, 28, 37, 42, 50, 54, 65, 57 CP FMNH, LACMNH, MPEG FMNH, LACMNH, MPEG, MZUSP FMNH, MPEG MNRJ, MPEG, MZUSP AMNH, FMNH, LACMNH, MNHW, MPEG, MZUSP FMNH, LACMNH, MNHW, MPEG MPEG FMNH CP, aviventris fl Family and bird species Atalotriccus pilaris Habitat Dist/Migr. Hemitriccus minor Hemitriccus zosterops Hemitriccus margaritaceiventer Hemitriccus inornatus Poecilotriccus russatus Poecilotriccus sylvia andrei Taeniotriccus maculatum Todirostrum cinereum Todirostrum pictum Todirostrum Rhynchocyclus olivaceus sulphurescens Tolmomyias assimilis Tolmomyias poliocephalus Tolmomyias Tolmomyias Platyrinchus saturatus Platyrinchus coronatus Platyrinchus platyrhynchos coronatus Onychorhynchus Myiophobus roraimae Myiophobus fasciatus Myiobius barbatus Myiobius atricaudus erythrurus Terenotriccus Hirundinea ferruginea euleri Lathrotriccus Cnemotriccus fuscatus Contopus cooperi Contopus fumigatus Contopus virens rubinus Pyrocephalus Knipolegus poecilocercus Knipolegus poecilurus Ochthornis littoralis Fluvicola pica Arundinicola leucocephala Colonia colonus Legatus leucophaius Myiozetetes cayanensis Myiozetetes similis Myiozetetes granadensis Myiozetetes luteiventris Pitangus sulphuratus Philohydor lictor Conopias parvus 232 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres .) in litt References Ornithological collections A. Whittaker ( # Evidence VZ T TFTF MFMFTFTFMFTF SF GUI WAM S WAM S TEP S T 40, 54 S S 54 #, 54 S 50, 54 28, 50 41, 49 #, 28, 42, 50, 54, 57 MPEG CP, FMNH, LACMNH, MPEG, MZUSP FMNH FMNH, MPEG MPEG CP DF GF VZ IG DF GF XSAVXAAVZ SAV XX AUS?SAVTF IGCAM STF AUS GF DFSAV TSAVX AUS #, 2, 10, 28, 37, 42, 50, 54, 57 DF GF SAV STF CAM AUS AUS/RES?TF CAMVZ IG S S NEA T STF S #, 50, 54 T T S T MPEG, MZUSP FMNH, LACMNH, MNHW, MF T #, 2, 10, 28, 41, 50, 54 #, 2, 28, 42, 50 #, 2, 10, 28, 42, 50, 54 T AUS S #, 2, 10, 28, 41, 42, 50, 54, 57, 64MF #, 2, 10, 28, 41, 42, 50, 54, 57, 64 STF # TF T S 28, 49 SMF TF T S T #, 2, 28, 42, 50, 54MF LACMNH, MPEG, MZUSP FMNH, INPA, AMNH, CP, S #, 10, 28, 50, 54, 66 TTF LACMNH, FMNH, MPEG, MZUSP AMNH, CP, # S T SMF FMNH, MPEG CP, 41, 42, 66 #, 10, 11, FMNH, MPEG, MZUSP TTF TEP #, 2, 54 #, 10, 37, 42, 54, 66 TTF DF VZ T FMNH, MPEG, LACMNH MZUSP #, 28, 42, 50, 54TF S #, 28, 42, 50, 54 #, 28, 42, 50, 54, 66TF SVZ GF T MPEG, MZUSP AMNH, FMNH, MPEG #, 2, 28, 50, 54CAM TEP STF 39 11, MZUSP FMNH, INPA, AMNH, CP, TF TEP MZUSP MNHW, AMNH, FMNH, INPA, S S S MPEG, MZUSP AMNH, LACMNH, MPEG, MZUSP 27, 42, 54 2, 11, S S FMNH, MPEG, MZUSP FMNH GUI #, 54 S 41, 49, 54 11, T S #, 2, 15, 28, 42, 50, 54, 57 41, 49 S MPEG WAM S #, 28, 50, 57 GUI T 2, 54, 57 T S S 28, 37, 49, 50, 54, 57 #, 11, 15, 28, 37, 50, 54 #, 2, 11, CP MNRJ, MPEG, MZUSP CP, 15, 28, 37, 50 T FMNH, LACMNH, MPEG, MZUSP #, 54 T 42, 50, 54 MPEG CP, MNHW CP, FMNH, MPEG MPEG FMNH, MNHW, CP, MPEG CP FMNH FMNH, MNHW FMNH, MPEG, MZUSP FMNH cauda fi Family and bird species Conopias trivirgatus Habitat Dist/Migr. Piprites chloris Corapipo gutturalis regulus Machaeropterus pyrocephalus Machaeropterus Lepidothrix coronata Lepidothrix suavissima Manacus manacus Myiodynastes maculatus Megarynchus pitangua sulphurea Tyrannopsis Empidonomus varius albogularis Tyrannus melancholicus Tyrannus savana Tyrannus dominicensis Tyrannus Rhytipterna simplex Rhytipterna immunda Sirystes sibilator tuberculifer Myiarchus swainsoni Myiarchus ferox Myiarchus tyrannulus Myiarchus Ramphotrigon ru Attila cinnamomeus Attila spadiceus OXYRUNCIDAE Oxyruncus cristatus COTINGIDAE Rupicola rupicola Cotinga cotinga Cotinga cayana albus Procnias averano Procnias Lipaugus vociferans Lipaugus streptophorus Xipholena punicea Gymnoderus foetidus Querula purpurata Perissocephalus tricolor Cephalopterus ornatus PIPRIDAE Neopelma chrysocephalum stolzmanni Tyranneutes virescens Tyranneutes The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 233 AMNH e (CP, FMNH, LACMNH, MPEG, MZUSP) (CP, f References Ornithological collections Evidence X RES/NEA S T # (2, 17, 28, 41, 42, 50, 54, 72) DF GF TF CAMDF GF MFCAMCAM IGTF SF CAM (GUI) S TEP T #, 15, 28, 37, 42, 50, 54 S S S S T T T 41, 49 11, #, 2, 15, 42, 50, 54, 57 #, 37 #, 42, 49, 57 MPEG, MZUSP FMNH, LACMNH, MNHW, FMNH, LACMNH, MPEG, MZUSP CP MPEG, MZUSP FMNH, MNHW VZ GFMFTF SF CAM MFVZTF CAMTFTF TFVZ VZ IG TF SF VZ SF TEPTF SFVZ DF GF (WAM)?/(GUI) STF S STF STF TSAV T #, 2, 15, 27, 28, 42, 50, 54, 57 GUI #, 54 #, 15, 28, 42, 50, 54 41, 49 11, X S STF S T S T FMNH, LACMNH, MNRJ, MPEG, MZUSP # S T #, 2, 28, 29, 37, 42, 50, 54, 57 S T #, 2, 28, 42, 50, 54, 57 2, 49, 50, 54 #, 15, 28, 37, 42, 50, 54, 63 T FMNH, MPEG, MZUSP S 2, 28, 50, 54 #, 37, 42, 50, 54 S S MPEG, MZUSP FMNH, LACMNH, MNHW, S CP T FMNH, MPEG #, 42, 54 T MPEG, MZUSP AMNH, FMNH, MNHW, # FMNH, MPEG, MZUSP S #, 49, 50, 51 #, 2, 54 T MPEG #, 28, 37, 41, 42, 50, 54, 78 MPEG, MZUSP LACMNH, MNHW, MPEG 28, 50, 54 FMNH, MZUSP MPEG, MZUSP FMNH, LACMNH, MNHW, CP, MPEG FMNH, MPEG FMNH CAM S 49, 54FMNHTF SFVZ IG GF DFMFCAM STFTF 49, TF SFTF W R TEPAA W R SAV (WAM)/(GUI)AA R SAV GUI SMF S S TR TR T # STF # AUS/RES #, 17, 28, 37, 42, 50, 54, 78 41, 49 11, T 54 S S #, 2, 50, 54 S TEP MPEG, MZUSP FMNH, LACMNH, MNHW, #, 28, 42, 50, 54, 57 S T S #, 28, 49, 50, 54 #, 2, 37, 49 S T #, 2, 17, 28, 42, 50, 54, 57 #, 2, 17, 28, 42, 50, 54, 57 S CP 41 S FMNH, MPEG T FMNH, MPEG LACMNH, MZUSP #, 17, 28, 42, 50, 54, 57 MPEG FMNH, LACMNH, MPEG, MZUSP #, 50 FMNH, LACMNH, MPEG, MZUSP MNHW FMNH, LACMNH, MPEG, MZUSP CP MPEG f avivertex fl licauda fi Family and bird species pareola Chiroxiphia Habitat Dist/Migr. Xenopipo uniformis Xenopipo atronitens Heterocercus Dixiphia pipra Pipra Pipra cornuta Pipra erythrocephala TITYRIDAE Schiffornis major Schiffornis turdina Laniocera hypopyrra Iodopleura fusca inquisitor Tityra cayana Tityra Pachyramphus rufus Pachyramphus polychopterus Pachyramphus marginatus Pachyramphus surinamus Pachyramphus minor Xenopsaris albinucha VIREONIDAE gujanensis Cyclarhis leucotis Vireolanius olivaceus Vireo Hylophilus thoracicus Hylophilus semicinereus Hylophilus pectoralis Hylophilus sclateri Hylophilus brunneiceps Hylophilus muscicapinus Hylophilus ochraceiceps CORVIDAE Cyanocorax violaceus Cyanocorax cayanus HIRUNDINIDAE albiventer Tachycineta tapera Progne chalybea Progne Pygochelidon cyanoleuca Atticora fasciata Atticora melanoleuca Neochelidon tibialis 234 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres References Ornithological collections Evidence SAV S 41, 50, 54WVZ SFSF SAV CP NEA NEA S S #, 16, 28, 37, 42, 50, 54 50 50, 51, 54 MPEG, MZUSP FMNH, LACMNH, MNHW, MPEG SAV RSAV R RW CAM SAV MFTFX NEAMFMF DF GFSAV SVZTF NEAVZ IG GF DFTF STF TEP #, 17, 28, 42, 50, 54, 55 (GUI)TF TEP #, 28, 42, 50, 54, 57 STF CAM TEPVZ IG DF GF SAV #, 28, 50, 54, 55 TF CAM S S S S 39, 41, 49 11, FMNH, LACMNH, MPEG, MZUSP S (GUI) 28, 50, 54 T T S 41, 49 T #, 2, 16, 27, 28, 37, 42, 49, 50, 54 #, 2, 16, 28, 37, 41, 42, 50, 54, 57 LACMNH, MPEG, MZUSP 41 S T #, 2, 16, 28, 37, 42, 50, 54, 57, 78 SMF S 41, 50, 54, 57 #, 2, 11, MFMF MNRJ, MPEG, MZUSP FMNH, LACMNH, MNHW, MPEG, MZUSP FMNH, LACMNH, MNHW, CP, S TAA GF DF SAV 28, 50, 57 S MPEG,MZUSP FMNH, LACMNH, MNHW, 49, 54 11, CAM MF #, 2, 16, 27, 28, 37, 42, 50, 54, 74, 78, 57 S CP TVZDF GF MNRJ, MPEG, MZUSP AMNH, FMNH, MNHW, #, 28, 50, 54VZ IG CAMTF 50, 54 MPEG CP, 54 TEPAA SAV CP TEP TEP CP SSAV S S MPEG S FMNH, MPEG CP, TX S #, 2, 16, 27, 28, 37, 42, 50, 54 MPEG S 41 CP, 40, 49 S S 28, 41, 49, 50 T FMNH, MPEG 41, 54 #, 11, #, 2, 16, 28, 42, 50, 54 T T S FMNH MNRJ, MPEG, MZUSP FMNH, MNHW, #, 16, 28, 37, 42, 50, 54 #, 42, 50, 51, 54, 57 T #, 2, 16, 27, 28, 37, 42, 49, 50, 54 S FMNH, MPEG, MZUSP CP MPEG, MZUSP S MNHW, MNRJ, MPEG, MZUSP LACMNH, MNHW, CP FMNH, INPA, CP #, 17, 28, 41, 42, 50, 54 CP MPEG, MZUSP T 17, 28, 41, 42, 50, 54, 57 #, 2, 11, FMNH, MPEG, MZUSP CP, FMNH, LACMNH, MPEG, MZUSP CP, g collis fi avipes fl aveola fl Family and bird species Alopochelidon fucata Habitat Dist/Migr. Donacobius atricapilla Stelgidopteryx ru TURDIDAE Catharus fuscescens Catharus minimus Platycichla Riparia riparia Hirundo rustica TROGLODYTIDAE ustulatus Microcerculus bambla Microcerculus musculus Troglodytes rufulus Troglodytes Cistothorus platensis griseus Campylorhynchus Thryothorus coraya Thryothorus leucotis leucosticta Henicorhina arada Cyphorhinus POLIPTILIDAE collaris Microbates Ramphocaenus melanurus Polioptila plumbea Polioptila guianensis SEDIS INCERTAE Platycichla leucops olivater Turdus leucomelas Turdus ignobilis Turdus fumigatus Turdus nudigenis Turdus albicollis Turdus MIMIDAE Mimus gilvus MOTACILIIDAE Anthus lutescens COEREBIDAE Coereba The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 235 References Ornithological collections Evidence CAM SAVTF SFVZ GF DF SAV VZ GFTFVZ GF DFTF SFCAMTFXXX STF CAM SMF (GUI)VZ GFTF SF MF T S TTFMF #, 28, 41, 50, 54 S #, 2, 37, 50, 54 STFMF S GUITF MF T #, 2, 42, 50, 54MF STFMF #, 18, 42, 54 STF MF S #, 27, 50, 54 #, 2, 18, 28, 42, 50, 54, 57 T GF DFSAV TEPTF #, 50, 54 S SMF (GUI) STF 41, 54 #, 11, WAM WAM FMNH, MPEG CP, S SSF T MPEG FMNH, LACMNH, MNHW, VZ IG CAM GF DF TF SAV T S #, 2, 18, 28, 42, 50, 54, 57TF S FMNH, LACMNH, MPEG, MZUSP #, 2, 28, 42, 50, 54 TEP #, 2, 18, 27, 28, 41, 42, 50, 54, 57, 77VZ MZUSP GUI 54 TF #, 2, 18, 28, 37, 42, 50, 54, 57 40TF DF FMNH, MPEG, MZUSP S S MNRJ, MPEG TF DF 2, 27, 50, 54 FMNH, LACMNH, MNRJ, MPEG, MZUSP AMNH, CP, WAM 49, 50, 54 STF S 54 S S TEPTF DF (GUI) FMNH, LACMNH, MPEG, MZUSP TF FMNH, MPEG S AUS? #, 2, 54 MPEG, MZUSP T FMNH, LACMNH, MNHW, CP GF DF 41, 49, 54 SVZ FMNH, LACMNH, MPEG, MZUSP #, 2, 18, 27, 41, 42, 50, 54, 76 S 40, 41, 54 #, 2, 28, 37, 42, 50, 54, 57MF S (GUI) 2, 49, 54 MPEG CP, DF MFSAV 27, 41, 49 11, SAV #, 50, 54 S S FMNH, MNRJ, MPEG MF #, 28, 50, 54 AMNH, FMNH, LACMNH, MNRJ, MPEG, MZUSP CP CP S MPEG, MZUSP FMNH, LACMNH, MNHW, CP, 50, 54, 57 #, 2, 42, 50, 54, 57 S S TEP MPEG CP, FMNH, MPEG T 57 #, 54 S CP S #, 28, 50, 54 S NEA T #, 2, 28, 42, 50, 54 MNRJ CP, CP #, 2, 28, 42, 50, 54, 57 FMNH, MPEG S TEP #, 37, 42, 54 #, 28, 50, 54 MPEG T FMNH, MPEG T 27, 41, 49 #, 31 S FMNH, MPEG, MZUSP #, 2, 18, 27, 28, 41, 42, 50 MPEG 55 MPEG, MZUSP 41, 49 FMNH, MZUSP FMNH, MPEG FMNH, LACMNH, MNRJ, MPEG, MZUSP CP, MPEG, MZUSP FMNH, MNHW, FMNH, MPEG MNRJ CP, CP h avicollis fl ava fl aviventer fl Family and bird species THRAUPIDAE Schistochlamys melanopis Habitat Dist/Migr. Piranga Cissopis leverianus Nemosia pileata Eucometis penicillata cristatus Tachyphonus luctuosus Tachyphonus surinamus Tachyphonus phoenicius Tachyphonus Lanio fulvus Ramphocelus carbo Thraupis episcopus Thraupis palmarum Cyanicterus cyanicterus Pipraeidea melanonota mexicana Tangara chilensis Tangara schrankii Tangara xanthogastra Tangara punctata Tangara guttata Tangara gyrola Tangara varia Tangara cayana Tangara nigrocincta Tangara cyanoptera Tangara velia Tangara viridis Tersina Dacnis cayana Dacnis lineata Dacnis Cyanerpes nitidus Cyanerpes caeruleus Cyanerpes cyaneus spiza Chlorophanes Hemithraupis guira Hemithraupis speciosum Conirostrum bicolor Conirostrum Diglossa major SEDIS INCERTAE Piranga rubra Piranga leucoptera 236 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres References Ornithological collections Evidence MFMF CAMSAVVZW SAV VZ TSAV # CAM SAV TEPAAVZ SF SAV WVZ W SAV W SSAV SAV WSAV SWVZ SF 41, 49 T S WSAV W SF SSAV 41, 54 #, 11, W VZ SAV T S CAM SF SSAV S #, 2, 20, 28, 37, 41, 42, 49, 50, 54SAV T TMF T 28, 41, 50, 54 # VZ T S #, 2, 28, 42, 50, 51, 54 STF #, 28, 41, 50, 54 #, 20, 28, 37, 42, 50, 54 SMF MPEG, MZUSP FMNH, LACMNH, MNHW, CP, TAAVZ IG GF R S T #, 28, 50, 54 S #, 2, 13, 49, 50, 51, 52, 54 #, 28, 49, 50, 51, 54TF CP FMNH, MPEG CP, STF CAM S STF SF LACMNH MPEG, MZUSP TEP 50 VZ GF DF SF MPEG, MZUSP LACMNH, MNHW, FMNH, INPA, CP #, 2, 28, 42, 50, 54TF SF FMNH, MPEG CP, SAV #, 2, 28, 41, 42, 50, 51, 54 TEP 20, 41, 54 2 S FMNH, MPEG #, 2, 20, 28, 42, 50, 51, 54 28, 50 S DF MFSAV S FMNH, MPEG SGF SF FMNH, LACMNH SGF DF SF 41, 49 T 28, 50, 54 TF SF DF MF #, 2, 20, 28, 42, 50, 54, 57 TTF SF MFDF FMNH, LACMNH, MPEG, MZUSP CP, #, 28, 42, 49, 50, 54WVZ FMNH, MZUSP SAV FMNH, LACMNH, MPEG,MZUSP 49 11, 28, 37, 41, 42, 50, 54 #, 2, 11, MF S NEA SMF S T T LACMNH NEA S CP S NEA NEA T #, 20, 27, 28, 37, 42, 50, 54 NEA #, 50, 54 #, 2, 54 FMNH, LACMNH, MPEG, MZUSP MPEG, MZUSP FMNH, LACMNH, MNHW, CP, S #, 2, 28, 42, 50, 54 T S S S LACMNH, MZUSP ANSP, #, 28, 50, 54 S T 30, 49 CP 37, 41, 49 #, 11, TEP MNRJ, MPEG, MZUSP LACMNH, MNHW, TEP S 39, 49 #, 2, 28, 37, 42, 49, 50, 54 #, 17, 28, 37, 49, 50, 54, 55 40, 49, 50, 54, 55 11, CP S S MPEG, MZUSP 28, 37, 50, 54 FMNH, MPEG 49 11, MPEG FMNH, LACMNH, MNHW, MZUSP MNHW, FMNH, MPEG 41, 49 MNHW CP, LACMNH CP, MPEG CP FMNH, MNHW CP CP Family and bird species oleagineus Mitrospingus Habitat Dist/Migr. EMBERIZIDAE Zonotrichia capensis Ammodramus humeralis Ammodramus aurifrons Sicalis citrina Sicalis columbiana Sicalis luteola Emberizoides herbicola jacarina Volatinia schistacea Sporophila intermedia Sporophila plumbea Sporophila americana Sporophila bouvronides Sporophila lineola Sporophila nigricollis Sporophila leucoptera Sporophila minuta Sporophila castaneiventris Sporophila angolensis Sporophila crassirostris Sporophila Catamenia homochroa conirostris Arremonops taciturnus Arremon Atlapetes personatus gularis Paroaria CARDINALIDAE Caryothraustes canadensis Saltator grossus Saltator maximus Saltator coerulescens Cyanocompsa cyanoides Spiza americana PARULIDAE Parula pitiayumi petechia Dendroica striata Dendroica fusca Dendroica Setophaga ruticilla Geothlypis aequinoctialis Myioborus miniatus Myioborus castaneocapillus The avifauna of the Brazilian state of Roraima: bird distribution and biogeography in the Rio Branco basin 237 k References Ornithological collections Evidence SAVTF SFTFTFMFTFTFMF S S WAM GUI T S TEP T S S #, 18, 37, 49, 50, 54, 75 S #, 2, 27, 28, 42, 50, 54 T S 2, 50, 54 #, 54 54 28, 41, 50 11, # 41 11, MPEG AMNH, FMNH, MNHW, LACMNH MNRJ, MPEG, MZUSP MPEG CP FMNH MPEG CP MFMF DFDF GFTF TEP S S S 40, 41, 49 11, S T 41, 54 # #, 17, 28, 42, 50, 54 CP FMNH, MPEG, MZUSP CP INPA TF CAMTF MFVZ IG CAMTF VZVZ GFXTFGF DF CAMSAV DF GFSAV MF SGF VZVZ W S TVZ RVZ IG GF S #, 17, 54 TVZ IG SF SFW SAV #, 28, 42, 50, 54, 57SAV S 2, 19, 28, 37, 50, 54, 57 TEP S SMF S S SCAM #, 54TF SF #, 28, 50 T T T S T #, 2, 19, 28, 37, 42, 50, 54, 57 #, 2, 19, 27, 28, 42, 49, 50, 54, 57 #, 27 #, 2, 19, 27, 28, 37, 42, 49, 50, 54 #, 28, 50, 54, 57 LACMNH, MPEG, MZUSP S MPEG FMNH, MNHW, S FMNH S 41, 49, 54 S FMNH, LACMNH, MNRJ, MPEG, MZUSP MNRJ, MPEG, MZUSP 28, 50 FMNH, LACMNH, MNHW, MPEG, MZUSP FMNH, LACMNH, MNHW, TEP 28, 50 T #, 2, 28, 37, 42, 50, 54 S 28, 50 #, 2, 19, 50, 54, 57 #, 2, 19, 28, 37, 42, 50, 54 S T S MPEG #, 2, 26, 27, 28, 37, 41, 42, 49, 50, 54 FMNH, MPEG T FMNH, MNRJ 41, 49, 54 MNRJ, MPEG, MZUSP T FMNH, LACMNH, MNHW, CP, MPEG, MZUSP FMNH, LACMNH, MNHW, #, 54 MPEG, MZUSP FMNH, LACMNH, MNHW, CP 50 (2, 28) FMNH, MPEG MPEG CP FMNH i j aveolus ventris fl fi nschi fi Euphonia violacea Euphonia chrysopasta Euphonia minuta Euphonia xanthogaster Euphonia ru Euphonia ceyennensis cyanea Chlorophonia Basileuterus culicivorus Basileuterus Phaeothlypis rivularis SEDIS INCERTAE Family and bird species Basileuterus bivittatus HabitatGranatellus pelzelni Dist/Migr. Euphonia ICTERIDAE viridis Psarocolius decumanus Psarocolius bifasciatus Psarocolius solitarius Procacicus Cacicus cela Cacicus haemorrhous Icterus croconotus Icterus chrysocephalus Icterus nigrogularis imthurnii Macroagelaius tanagrinus Lampropsar Chrysomus icterocephalus Gymnomystax mexicanus Molothrus bonariensis Molothrus oryzivorus Sturnella militaris Sturnella magna FRINGILLIDAE magellanica Carduelis Euphonia plumbea Euphonia chlorotica 238 Luciano Nicolás Naka, Mario Cohn-Haft, Francisco Mallet-Rodrigues, Marcos Pérsio Dantas Santos, and Marcela de Fátima Torres

a Previous sight records from Maracá are likely to represent N. hyperrhynchus, which should be present west of the Rio Branco. b Previous records mention Capito niger, but are likely to represent C. auratus. c Our records represent L. a. albolineatus; previous records likely to represent L. a. duidae. d Unidentifi ed to species by Stotz (1997) and Trolle and Walther (2004). e Previous records mention Inezia subfl ava, but are likely to represent I. caudata. f Our records are likely to refer to the resident form Vireo olivaceus chivi (as well as a skin at AMNH); previous records referred as belonging to the nominate migratory form. g Included within the Troglodytidae by the CBRO, but we follow the SACC (Remsen et al. 2006) and place it as Incertae sedis until its taxonomic relationships are better understood. h Included within the Thraupidae by the CBRO, but we follow the SACC (Remsen et al. 2006) and place it as Incertae sedis until its taxonomic relationships are better understood. i The records for the state represent the form roraimae, which has been suggested to represent a valid species (Hilty 2003) j Included within the Parulidae by the CBRO, but we follow the SACC (Remsen et al. 2006) and place it as Incertae sedis until its taxonomic relationships are better understood. k Original records mention E. laniirostris, but are likely to represent E. chlorotica (see Hypothetical species)

Habitat TF: terra fi rme forest; SF: secondary forest, forest edge, and clearings within terra fi rme forest; MF: montane forest (includes humid forests above 600 m, and all montane habitats present in the Tepuis); VZ: varzea forest; IG: igapo forest; CAM: campina and campinarana; SAV: savanna; GF: gallery forest; DF: dry forest; R: associated to rivers; W: wetlands; AA: antropic areas; X: virtually all habitats.

Distribution/migration GUI: species restricted to the Guianan area of endemism; (GUI): subspecies restricted to the Guianan area of endemism; WAM: species with distributions typical of western Amazonia, and generally absent from the Guianan area of endemism; (WAM) subspecies with distributions typical of western Amazonia, and generally absent from the Guianan area of endemism; ?: taxon possibly present (but not confi rmed) in Roraima; TEP: species or subspecies restricted to the Tepuis; NEA: Nearctic migrant (species breeding in North America that reach Roraima during the boreal winter [Sept-April]); AUS: Austral migrant (species breeding in southern South America that spend the Austral winter [April-August] in Amazonia and northern South America).

Evidence S: specimen available (see Ornithological collections); T: tape-recordings available at the INPA Archive of Bird Sounds; P: photograph available.

References 1: Bierregaard et al. (1997), 2: Borges (1994), 3: Cohn-Haft et al. (1997), 4: Collar et al. (1992), 5: Cory (1918), 6: Cory (1919), 7: Cory (1920), 8: Cory and Hellmayr (1924), 9: Cory and Hellmayr (1925), 10: Cory and Hellmayr (1927), 11: Dickerman and Phelps (1982), 12: Forrester (1993), 13: Forrester (1995), 14: Hellmayr (1906), 15: Hellmayr (1929), 16: Hellmayr (1934), 17: Hellmayr (1935), 18: Hellmayr (1936), 19: Hellmayr (1937), 20: Hellmayr (1938), 21: Hellmayr and Conover (1942), 22: Hellmayr and Conover (1948a), 23: Hellmayr and Conover (1948b), 24: Hellmayr and Conover (1949), 25: Joseph (1992), 26: Joseph (2001), 27: Miranda-Ribeiro (1929), 28: Moskovits et al. (1985), 29: Naumburg (1930), 30: Novaes (1967), 31: Pacheco (1995a), 32: Pelzeln (1856), 33: Pelzeln (1859), 34: Pelzeln (1861), 35: Pelzeln (1862), 36: Pelzeln (1863), 37: Pelzeln (1868-71), 38: Peters (1937), 39: Phelps (1973), 40: Phelps and Phelps (1948), 41: Phelps and Phelps (1962), 42: Pinto (1966), 43: Ruschi (1961), 44: Santos (2004), 45: Schattuck (1926), 46: Schlegel (1864), 47: Sclater (1874), 48: Sick (1965), 49: Sick (1997), 50: Silva (1998), 51: Silva and Oren (1990), 52: Silva and Willis (1986), 53: Spix (1824-25), 54: Stotz (1997), 55: Stotz et al. (1992), 56: Teixeira et al. (1986), 57: Trolle and Walther (2004), 58: Vaurie (1980), 59: Whittaker (1995), 60: Whittaker (1996), 61: Willis (2003), 62: Zimmer (1933), 63: Zimmer (1936), 64: Zimmer (1937a), 65: Zimmer (1937b), 66: Zimmer (1938), 67: Zimmer (1939a), 68: Zimmer (1939b), 69: Zimmer (1940), 70: Zimmer (1941a), 71: Zimmer (1941b), 72: Zimmer (1941c), 73: Zimmer (1942a), 74: Zimmer (1942b), 75: Zimmer (1943a), 76: Zimmer (1943b), 77: Zimmer (1944), 78: Zimmer et al. (1997).

Ornithological collections AMNH: American Museum of Natural History; ANSP: Academy of Natural Science of Philadelphia; CP: Colección Phelps; FMNH: Field Museum of Natural History; INPA: Instituto Nacional de Pesquisas da Amazônia; LACMNH: Los Angeles County Museum of Natural History; MNHW: Museum of Natural History of Wien; MNRJ: Museu Nacional de Rio de Janeiro; MPEG: Museu Paraense Emílio Goeldi; MZUSP: Museu de Zmologia da Universidade de São Paulo; UMMZ: University of Michigan Museum of Zoology.