Levantamento da avifauna das savanas amazônicas e de habitats adjacentes da região de Monte Alegre (Pará, Brasil), com comentários sobre biogeografia e conservação

Marcelo Ferreira de Vasconcelos^I, Sidnei de Melo Dantas^II, José Maria Cardoso da Silva^III

^IPontifícia Universidade Católica de Minas Gerais. Belo Horizonte, Minas Gerais, Brasil
^IIMuseu Paraense Emílio Goeldi. Coordenação de Zoologia. Belém, Pará, Brasil
^IIIConservação Internacional do Brasil. Belém, Pará, Brasil

Autor para correspondência

ABSTRACT

Knowledge of the avifaunal composition of Amazonian savannas is important in facilitating a greater understanding of the patterns of evolution ofthe regional biota. Despite this, this vegetation type has been poorly sampled in comparison to adjacent rainforest habitats. The aim of this paper is to provide an avifaunal inventory of the savannas and other adjacent habitats of Monte Alegre, on the left bank of Amazon River, northwestern Pará, Brazil. Our survey is based both on our own fieldwork and on previously-collected specimens deposited in museums. A total of 300 bird species have been recorded in the region. Ninety-four species (31.3%) were restricted to a single habitat type, indicating that habitat heterogeneity is important to the overall species richness. Despite the fact that the Monte Alegre savannas present typical species restricted to this habitat, several open-country birds that occur in larger savanna areas located on the borders of Amazonia (along the Atlantic coast and close to the 'cerrado' boundaries) were not recorded in this area. Although Monte Alegre is species impoverished relative to other savanna regions in the Amazon basin, the Monte Alegre savannas shelter the Sulfur-breasted Parakeet (Aratinga maculata), a vulnerable species in Pará state that is only known from the Guianan area of endemism (Aleixo et a/., 2011). We also comment on biogeography, taxonomy, noteworthy records, natural history and conservation of the regional avifauna.

Keywords: Avifaunal survey. Amazonian savannas. Biogeography. Natural history. Conservation. Aratinga maculata.

RESUMO

O conhecimento sobre a composição da avifauna das savanas amazônicas é muito importante para o entendimento de padrões da evolução da biota regional. Entretanto, este tipo de vegetação foi pouco amostrado em comparação com áreas de florestas adjacentes. O objetivo deste trabalho é apresentar um levantamento da avifauna das savanas e de habitats adjacentes de Monte Alegre, na margem esquerda do rio Amazonas, noroeste do Pará, Brasil. Nossa pesquisa baseou-se em nossos trabalhos de campo e em exemplares coletados anteriormente e depositados em museus. Um total de 300 espécies de aves foi registrado na região. Noventa e quatro espécies (31,3%) estiveram restritas a um único tipo de habitat, indicando que a heterogeneidade de habitats é importante para a riqueza geral de espécies. Apesar de as savanas de Monte Alegre apresentarem espécies típicas deste habitat, faltam outras aves de ambientes abertos que ocorrem em áreas maiores de savana localizadas nas bordas da Amazônia, especialmente ao longo da costa Atlântica ou próximo aos limites do cerrado. Embora a avifauna de Monte Alegre seja relativamente pobre em comparação com outras áreas de savanas na bacia amazônica, as savanas de Monte Alegre abrigam o cacaué (Aratinga maculata), uma espécie vulnerável no estado do Pará, com distribuição restrita à área de endemismo Guiana (Aleixo et al., 2011). Também comentamos sobre biogeografia, taxonomia, registros notáveis, história natural e conservação da avifauna regional.

Palavras-chave: Levantamento de avifauna. Savanas amazônicas. Biogeografia. História natural. Conservação. Aratinga maculata.

Patches of savannas and other open vegetation biotypes are found nested in extensive areas of rainforest in Brazilian Amazonia (Prance, 1978; Pires & Prance, 1985; Sanaiotti et al., 1997; Miranda & Absy, 2000; Vicentini, 2004; Gottsberger & Silberbauer-Gottsberger, 2006; Pennington et al., 2006; Ratter et al., 2006; Magnusson et al., 2008). Amazonian savannas occur mainly on poor, sandy soils on both banks of the Amazon River (Gottsberger & Silberbauer-Gottsberger, 2006; Pennington et al., 2006; Ratter et al., 2006). Important sites include those in the states of Roraima, Amapá, Amazonas and Pará (Andrade-Lima, 1958; Miranda, 1993; Miranda & Absy, 2000; Gottsberger & Silberbauer-Gottsberger, 2006; Pennington et al., 2006; Ratter et al., 2006; Magnusson et al., 2008).

Knowledge of the avifauna composition of Amazonian open habitats is crucial to understanding patterns of distribution and evolution of birds in South America (E. Snethlage, 1909; Haffer, 1967, 1969, 1974, 1985, 1987; Silva, 1995; Silva et al., 1997; Silva & Bates, 2002; Bates et al., 2003; Robbins et al., 2004; Mittermeier et al., 2010). Amazonian savannas are considered present-day 'refugia' of a widespread open vegetation type that probably expanded across the Amazonian belt of low precipitation during dry phases of the Cenozoic (Haffer, 1967, 1969, 1974; Prance, 1978, 1987; Haffer & Prance, 2001; Ab'Saber, 2002). Despite their importance for the Amazonian biogeography, these areas have been poorly sampled in comparison to rainforest sites throughout the Amazonian region, and published avifaunal inventories have focused on savannas in the Sipaliwini region (Suriname) and in the Brazilian states of Roraima, Amazonas, Pará, Amapá and Rondônia (Henriques & Oren, 1997; Silva et al., 1997; Sanaiotti & Cintra, 2001; Naka et al., 2006; Aleixo & Poletto, 2007; Santos & Silva, 2007; Mittermeier et al., 2010; Aleixo et al., 2011).

Monte Alegre is located on the left (northern) bank of the Amazon River, northwestern Pará, Brazil. The region is located on the southern border of the Guianan area of endemism, a well known area of avian endemism (see Cracraft, 1985). Monte Alegre savannas are isolated from other large savanna areas (e. g., 'cerrado', Amapá, Llanos, Roraima-Rupununi, Sipaliwini) by hundreds of kilometers of Amazonian rainforest, but are only separated by the Amazon River from other savanna enclaves on the southern bank (Alter do Chão) (Figure 1).

Ornithological surveys of Monte Alegre started in the 19^th century, when F W. Sieber collected some specimens in this region, between 1801 and 1812 (Pinto, 1979). These specimens were deposited in the Museum für Naturkunde (ZMB), Berlin (Pinto, 1979). The famous British naturalist A. R. Wallace also explored the area in 1849 and collected a few specimens that have been deposited in the British Museum of Natural History (BMNH), Tring (Sclater & Salvin, 1867; Pinto, 1979). Later, more detailed ornithological collections were made by E. Snethlage, O. Martins, A. Costa, J. de Sá, J. Anthero and F J. José during the first two decades of the 20^th century (E. Snethlage, 1906, 1907, 1914; H. Snethlage, 1930; Miranda-Ribeiro, 1938). During these expeditions, over 720 specimens were collected and deposited in the Museu Paraense Emílio Goeldi (MPEG), Belém. Between 1917 and 1920, F Q. Lima and E. Garbe, working for the Museu Paulista (currently Museu de Zoologia da Universidade de São Paulo, MZUSP), also collected specimens in Monte Alegre (Pinto, 1944, 1945). Shortly after these expeditions, C. Lako collected birds in the Monte Alegre region (Maicuru River) in 1928 and 1929 (Paynter & Traylor, 1991; Silveira et al., 2005). These specimens were deposited in the Field Museum of Natural History (FMNH) and in the American Museum of Natural History (AMNH) (Paynter & Traylor, 1991; Silveira et al., 2005). A. M. Olalla also visited the region in 1936, when he collected specimens that were sent to the FMNH (Paynter & Traylor, 1991). Recently, a new species of parakeet (Psittacidae), Aratinga maculata (Sulfur-breasted Parakeet), was described from Monte Alegre (Silveira et al., 2005) (the species was described as Aratinga pintoi but this is now considered a junior synonym of Aratinga maculata, see Nemésio & Rasmussen, 2009), and the region became better known among ornithologists and birdwatchers (Silveira, 2005; Balchin, 2006). Although this region has been inventoried by generations of ornithologists, old collections from Monte Alegre have never been studied in detail and the area lacks an ornithological checklist. Thus, the aim of this paper is to provide an avifaunal inventory of the Monte Alegre region, based on our own fieldwork and on specimens held in museums. We also comment on biogeography, taxonomy, noteworthy records, natural history and conservation of the avifauna.

STUDY AREA

Our study was located in the Monte Alegre region (between 01^o 56' and 02^o 07' S, and 54^o 03'and 54^o 13' W), state of Pará, northern Brazil (Figure 1). The elevation ranges from 3 m to 400 m (Pastana, 1999) and the climate is hot and humid (type Awi of the Kõppen classification) (Ayoade, 1998). Temperatures vary between monthly averages of 25.7o and 27^o C (Oliveira-Júnior et al., 1999) and mean annual precipitation is 1,678 mm (Moraes et al., 2005). Rainfall peaks in April and the rainy season is concentrated between February and May, when monthly precipitation surpasses 200 mm. The dry season is between September and October, when monthly rainfall averages are lower than 50 mm (Oliveira-Júnior et al., 1999).

Savanna vegetation (locally named 'coberto') occurs generally on sandy or rocky soils, with the presence of a few species of grasses and herbs and a very inconspicuous litter layer (Figure 2A and Figure 2B). Shrubby-arboreal vegetation is primarily represented by sparse individuals of the following species: Curatella americana, Qualea grandiflora, Salvertia convallariodora, Sclerolobium paniculatum, Anacardium occidentale, Palicourea rigida and Tocoyena formosa, generally varying from 1 to 4 m in height. At Serra do Ereré (02^o 01' 16" S 54^o 10' 55" W), a mountainous locality in this region (elevation: 220 m), savanna occurs mainly over and between rock outcrops, with shrubs of Cereus sp., Phyllodendron sp. and Norantea guianensis. In this area there are also sparse shrubby-arboreal individuals of Aspidosperma sp., Byrsonima crassifolia, Q. grandiflora and S. convallariodora growing on sandy or rocky soils. Some of these trees are parasitized by a mistletoe (Psittachanthus sp.) with hummingbird-pollinated flowers (Vasconcelos & Silva, 2005). For a detailed botanical inventory of the Monte Alegre savannas, see Andrade-Lima (1958).

These savannas are intermixed with patches of 'campinarana', a low-canopy forest growing on sandy soil with trees reaching up to 10 m in height. These 'campinaranas' are very dense, with a developed understory with tangles of vines and shrubs (mainly Miconia sp.). At some places within the savannas, there are clumps of trees that resemble small patches of 'campinarana', with a denser layer of litter and shrubs of Eugenia sp., Miconia sp. and Cereus sp.

Original 'terra firme' forests ('floresta ombrófila') were extensively cleared and fragmented (Pereira et al., 2003). Several areas originally covered by this vegetation type are now occupied by clumps of an abundant pioneer species of palm tree (Attalea sp.), and the remnant fragments are subject to illegal logging (see below). Typical tree species of those forests are: Aspidosperma desmanthum, Bellucia sp., Geissospermum sericeum, Hymenaea parviflora, Inga micradenia, Licania spp., Myrcia paivae, Nectandra mollis and Protium paniculatum (Garcia et al., unpublished data 2008).

METHODS

We made three expeditions to Monte Alegre aiming principally to collect specimens in the savannas. The first expedition was performed by JMCS, MFV and D. C. Pimentel Neto between 2 and 20 January 1999. Specimens were collected with shotguns and mist-nets and have been deposited in MPEG and in the ornithological collection of the Department of Zoology, Universidade Federal de Pernambuco (UFPE). Localities sampled during this expedition were: savannas and 'campinaranas' at km 4 of road PA-423 (01^o 56' 38" S 54^o 03' 45" W); savannas of Campo do Desterro (01^o 57' 04" S 54^o 10' 20" W); savannas and 'terra firme' forest fragments at Colônia do Ererê (01^o 58' 14" S 54^o 10' 01" W); and savannas and 'campinaranas' at Serra do Ererê (02^o 01' 16" S 54^o 10' 55" W). The second expedition was made by SMD and J. N. Santa Brígida, between 21 and 28 September 2006, during a study for the management plan of the Monte Alegre State Park. Fieldwork was conducted in the following localities: a 'terra firme' forest fragment at Ilha Grande (02^o 01' 55" S 54^o 10' 39" W); flooded forest at Aruchi (02^o 01' 28" S 54^o 12' 33" W); flooded forest at Igarapé do Ererê (01^o 59' 20" S 54^o 08' 57" W); 'terra firme' forest and savannas at Colônia do Ererê (02^o 00' 06" S 54^o 10' 48" W); savannas at Santana (02^o 00' 30" S 54^o 09' 58" W); and savannas at Serra do Ererê (02^o 01' 32" S 54^o 11' 07" W). Specimens were collected with mist-nets and deposited in MPEG. The third expedition was made by SMD, between 19 and 23 March 2009, in the following localities: lake at Lago da Conceição (02^o 04' 54" S 54^o 10' 21" W); 'buriti' palm grove at Mutuacá (02^o 03' 18" S 54^o 12' 21" W); and flooded forests along the Amazon River (02^o 07' 08" S 54^o 09' 36" W). In all expeditions, birds were also photographed and tape-recorded. Recordings will be deposited at the Arquivo Sonoro Prof. Elias Coelho (ASEC), Departamento de Zoologia, Universidade Federal do Rio de Janeiro (UFRJ), Rio de Janeiro, Brazil. Some of these recordings have been archived on the online resource Xeno-Canto (2009).

We also studied old specimens collected in the Monte Alegre region deposited in MPEG and checked the literature (Pinto, 1944; Ruschi, 1951; Silveira et al., 2005) for additional specimens deposited in the following institutions: AMNH, FMNH, MZUSP and Museu Nacional do Rio de Janeiro (MNRJ).

We took notes in the field in which habitat each species was observed. For those recorded exclusively in museums, we classified them by habitat type based on our own field experience and following Ridgely & Tudor (1989, 1994), Stotz et al. (1996), Cohn-Haft et al. (1997), Borges et al. (2001) and Naka et al. (2006). Systematic order and nomenclature follow those of the Comitê Brasileiro de Registros Ornitológicos (CBRO, 2009).

RESULTS AND DISCUSSION

AVIFAUNAL SURVEY

A total of 300 bird species was recorded in the Monte Alegre region (Appendix). Flooded forests and 'terra firme' forests held the highest species richness (146 and 143 species, respectively), followed by 'campinarana' (122 species), savannas (109 species) and 'campos de várzea' (99 species) (Table 1). Disturbed areas and open waterbodies (lakes and rivers) had the lowest species richness (respectively 57 and 54 species, Table 1).

Ninety-four species (31.3%) were restricted to a single habitat type (Table 1). This shows that habitat heterogeneity is important for the overall species richness (see Cohn-Haft et al., 1997). Similar results were obtained in other surveys of Amazonian savannas intermixed with other vegetation types (Silva et al., 1997; Aleixo & Poletto, 2007; Santos & Silva, 2007; Aleixo et al., 2011). Open waterbodies (lakes and rivers) hold the highest percentage of exclusive species (48.1%), followed by savannas, which presented 18.3% exclusive species (Table 1). On the other hand, 'campinaranas' showed the lowest percentage of exclusive species (3.3%), probably because this habitat shelters several birds that also live in other forest types, such as 'terra firme' and flooded forests (Borges, 2004).

Aratinga maculata (Sulfur-breasted Parakeet) was the only species considered to be restricted to the Monte Alegre region (Silveira et al., 2005), although some specimens listed by these authors were taken from Suriname and considered mislabeled. Recently, Mittermeier et al. (2010) collected an additional specimen in the Sipaliwini savanna, confirming its presence in Suriname. More recent records were presented by Aleixo et al. (2011) from Trombetas State Forest, southern part of the Grão Pará Ecological Station and Paru State Forest. Lophornis ornatus (Tufted Coquette) was previously considered by Cracraft (1985) as a Guianan area of endemism endemic, but it also occurs in Cracraft's Parian Montane area of endemism (see map in Züchner, 1999) and on the right bank of the Amazon River, in the savannas of Alter do Chão (Sanaiotti & Cintra, 2001).

Three threatened species were recorded in the region (following Machado et al., 2005; SECTAM, 2006; BirdLife International, 2009): Anodorhynchus hyacinthinus (Hyacinth Macaw) - globally endangered, vulnerable in Brazil and in Pará state; Primolius maracana (Blue-winged Macaw) - near threatened globally and in Brazil, vulnerable in Pará state; and Aratinga maculata (Sulfur-breasted Parakeet) - vulnerable in Pará state, but not yet evaluated by BirdLife International (2009) and by the Brazilian red list.

COMMENTS ON THE BIOGEOGRAPHY OF THE MONTE ALEGRE SAVANNA AVIFAUNA

It is possible to make some biogeographical comments on the avifauna of the Monte Alegre savannas. For example, Colinus cristatus (Crested Bobwhite) is the only species restricted to the savannas north of the Amazon River found in Monte Alegre (Sick, 1997). On the other hand, 13 savanna species whose ranges in South America are mainly centered on regions located south of Amazonia ('caatinga', 'cerrado' and 'chaco'; following Silva et al., 1997) were recorded at Monte Alegre: Buteo albicaudatus (White-tailed Hawk), Aratinga aurea (Peach-fronted Parakeet), Chordeiles pusillus (Least Nighthawk), Colaptes campestris (Campo Flicker), Formicivora rufa (Rusty-backed Antwren), Lepidocolaptes angustirostris (Narrow-billed Woodcreeper), Elaenia cristata (Plain-crested Elaenia), Suiriri suiriri (Suiriri Flycatcher), Xolmis velatus (White-rumped Monjita), Tyrannus albogularis (White-throated Kingbird), Tyrannus savana (Fork-tailed Flycatcher), Mimus saturninus (Chalk-browed Mockingbird) and Ammodramus humeralis (Grassland Sparrow). These species exhibit the Peri-Atlantic distribution pattern (following Silva, 1995). This pattern suggests connections between the 'cerrado' region and the northern South American savannas through an Atlantic coast savanna corridor (Silva, 1995; Silva & Bates, 2002). However, other important species sharing this pattern of distribution occur in coastal savannas (e. g., Marajó island and Amapá state; see Silva, 1995; Henriques & Oren, 1997; Silva et al., 1997), but are absent in the Monte Alegre savannas. Examples are: Uropelia campestris (Long-tailed Ground-Dove), Guira guira (Guira Cuckoo), Neothraupis fasciata (White-banded Tanager) and Coryphaspiza melanotis (Black-masked Finch). Thus, it seems that the avifauna of the Monte Alegre savannas have been isolated from other large South American savannas ('cerrado' and Llanos) for a longer period in comparison to coastal savannas, which seem to have had more recent connections to the 'cerrado' (Silva, 1995; Silva et al., 1997; Silva & Bates, 2002; Bates et al., 2003; Mittermeier et al., 2010). This pattern does not support the presence of a broad savanna corridor across the Amazonian belt of low precipitation, one of the assumptions of the refuge theory (Haffer, 1969, 1974). However, in a recent paper, Haffer & Prance (2001) stressed that not just savannas were believed to expand and isolate the wet forest 'refugia'. They suggested that other vegetation types of seasonally dry climates (e.g., dry forest, liana forest and bamboo forest) could have been effective barriers to the dispersal of rainforest 'refugia' organisms (see also Pennington et al., 2000). Bonaccorso et al. (2006) modeled potential distributions of 17 species of plants and birds during the Last Glacial Maximum and suggested that past climate changes fragmented rainforest species' ranges but there was not a clear pattern for savanna species. These results also do not support the broad expansion of savannas into areas currently covered by Amazonian rainforests. Various other studies show a similar pattern; most notably those based on the pollen record (see Bush & de Oliveira, 2006 for a recent review).

Furthermore, the relatively small area of Monte Alegre savannas, coupled with their isolated location in central Amazonia relative to other larger savanna areas within the borders of the Amazon region, such as Amapá (Silva et al., 1997), Roraima-Rupununi (Robbins et al., 2004; Santos & Silva, 2007) and the upper Marmelos River (Aleixo & Poletto, 2007), can also explain the absence of several other typical South American savanna species recorded in those regions. Examples are: Heliactin bilophus (Horned Sungem), Melanopareia torquata (Collared Crescentchest), Polystictus pectoralis (Bearded Tachuri), Euscarthmus rufomarginatus (Rufous-sided Pygmy-Tyrant), Xolmis cinereus (Gray Monjita), Anthus lutescens (Yellowish Pipit), Cypsnagra hirundinacea (White-rumped Tanager), Emberizoides herbicola (Wedge-tailed Grass-Finch) and Sporophila bouvreuil (Capped Seedeater). Alter do Chão, another central Amazonian savanna site, also presents an impoverished savanna avifauna (Sanaiotti & Cintra, 2001). Thus, local extinction is another potential explanation for the lack of some species in Monte Alegre and other small and isolated central Amazonian savannas. These species may have colonized the region during past expansions of non-forest vegetation types into the Amazonian region (Haffer & Prance, 2001), but may have subsequently become extinct as either the habitat became unsuitable or habitat patches too small to support effective metapopulations.

SPECIES ACCOUNTS

Below, we present comments on distribution, taxonomy and natural history of selected species:

Colinus cristatus (Crested Bobwhite)

The only regional record for this northern South American savanna game bird is based on two specimens (a couple) collected by O. Martins in October 1916 (MPEG 13006­13007). This record was probably overlooked, since it represents a range extension for this species to the mid-lower left bank of the Amazon River (see maps in Carrol, 1994 and Infonatura, 2009). We failed to find the Crested Bobwhite during our fieldwork and it is possible that the species could have been extirpated from the region due to hunting pressure (see below).

Buteo albicaudatus (White-tailed Hawk)

This is an uncommon species in Amazonia, with sparse records for other savanna areas, such as Marajó island (Henriques & Oren, 1997), Amapá (Silva et al., 1997), Alter do Chão (Sanaiotti & Cintra, 2001), Roraima-Rupununi (Robbins et al., 2004; Santos & Silva, 2007), upper Marmelos River (Aleixo & Poletto, 2007) and Sipaliwini (Mittermeier et al., 2010). In Amazonia, the species was also recorded around 'terra firme' forests in the Alta Floresta region (Zimmer et al., 1997) and on the Serra dos Caiabis, in transitional forest (Lees et al., 2008). The first record for Monte Alegre is an adult bird photographed by MFV on 16 January 1999. The bird was perched on a rock outcrop atop Serra do Ererê and took flight some minutes later. On 19 January 1999, three individuals (one in dark phase) were seen flying over the same locality.

Aratinga maculata (Sulfur-breasted Parakeet)

This species is endemic to the Guianan area of endemism (Silveira et al., 2005; Mittermeier et al., 2010; Aleixo et al., 2011); records from Santarém, on the south bank of the Amazon River, are probably from captive birds or mislabeled old specimens (Silveira et al., 2005). We observed flocks of 3-8 individuals in five localities: Monte Alegre city, Campo do Desterro, Colônia do Ererê, Santana and Aruchi. Birds were observed principally in savannas but were also recorded close to forest fragments or flying over forested or urban areas. Two males that were used in the original description of A. pintoi (Silveira et al., 2005) were collected by JMCS, MFV and D. C. Pimentel Neto, at Colônia do Ererê on 9 January 1999 (MPEG 54330, UFPE 1792).

Chordeiles pusillus (Least Nighthawk)

 Lophornis ornatus (Tufted Coquette)

This species is known from Monte Alegre from a single male specimen (MPEG 4759) collected by E. Snethlage on 15 February 1906. This appears to be the only specimen from the mid-lower reaches of the north bank of the Amazon River (see maps in Zuchner, 1999 and Infonatura, 2009).

Notharchus ordii (Brown-banded Puffbird)

This poorly known puffbird is known from scattered records in the basins of the Amazon and Orinoco (Zimmer & Hilty, 1997; Zimmer et al., 1997; Borges et al., 2001; Rasmussen & Collar, 2002; Infonatura, 2009). The only record for the left bank of the mid-lower Amazon River appears to be a female (MPEG 4405) collected by A. Costa on 17 November 1905.

Formicivora rufa (Rusty-backed Antwren)

Silva et al. (1997) suspected that the subspecies of the Rusty-backed Antwren occurring in Monte Alegre would be F. r. chapmani, a taxon restricted to the Amazonian savannas other than Humaita and the upper Marmelos River (both in Amazonas state and close to the border of the 'cerrado'), where the nominate race has been collected (Silva et al., 1997; A. Aleixo & F Lima, personal communication). This species is known in Monte Alegre from a female (MPEG 4744) and a male (MPEG 6004), collected by E. Snethlage and O. Martins, on 14 February 1906 and on 21 July 1908, respectively. Between 10 and 19 January 1999, four specimens (one male and three females) were obtained in the Monte Alegre savannas by JMCS, MFV and D. C. Pimentel Neto (MPEG 54367­54370). The female specimens present heavy streaking on underparts (especially on throat) and blackish tails, typical features of F r. chapmani, following Zimmer & Isler (2003) (but see Pinto, 1940, 1947 for a critical point of view, suggesting that some of these characters may be related to individual variation).

Suiriri suiriri (Suiriri Flycatcher)

Suiriri Flycatcher is known from few sites in Amazonia (Silvaet al., 1997; Sanaiotti & Cintra, 2001; Robbins, 2004; Infonatura, 2009; Mittermeier et al., 2010). However, S. s. affinis is fairly common in the Monte Alegre savannas, where five specimens were collected by E. Snethlage (MPEG 5934) and by JMCS, MFV and D. C. Pimentel Neto (MPEG 54390-54391, UFPE 1770-1771).

Conopias trivirgatus (Three-striped Flycatcher)

Sicalis citrina (Stripe-tailed Yellow-Finch)

The Stripe-tailed Yellow-Finch is a species with two main populations in South America: one north of Amazonia and the other south of this region (the North-South disjunction pattern of distribution proposed by Silva, 1995). This species has only been previously found on the borders of Amazonia (Ridgely & Tudor, 1989; Infonatura, 2009). The first record of this species in a central Amazonian savanna is of a male (MPEG 54434), collected by JMCS, MFV and D. C. Pimentel Neto, at Serra do Ereré on 16 January 1999. This bird was associated with a flock of several other individuals that were vocalizing and performing aerial displays, similar to those described from populations south of the Amazon (Vasconcelos et al., 2007a). Small flocks of 2-5 birds were observed foraging on the ground, among clumps of grasses. Silva (1995) suggested that populations of this species might be able to disperse across Amazonia, since it has a tendency toward long-distance movements (Ridgely & Tudor, 1989; Sick, 1997).

CONSERVATION

Vegetation types in Monte Alegre have been severely degraded by human activities since the first colonizers arrived in the region, during the 17^th century (Pastana, 1999). Savannas provided a good opportunity for cattle ranching (Pastana, 1999), probably because this open physiognomy (with native grasses) facilitated this activity, and it is possible that non-natural fires have occurred in this open vegetation type at least since the 18^th century. To what extent fires caused faunal and floral impoverishment is unknown and it is possible that even the earliest naturalists that explored the region already found degraded savanna vegetation in Monte Alegre. Fires continue to be set in the savannas, and the herb layer is extremely depauperate and sparsely vegetated in recently burnt areas.

Intensive logging in Monte Alegre started toward the end of the 18^th century when the Royal Sawmill ("Serraria Real") exploited the timber for commercial purposes (Pastana, 1999). This logging activity and the cocoa plantation that followed during the first years of the 19^th century were probably two early regional drivers of deforestation (Pastana, 1999). Logging activities continue in the region today, and the remnant fragments of forest are isolated and impoverished. It is possible that several forest bird species have disappeared from Monte Alegre; for example, a number of woodcreepers and antbirds of the Guianan area of endemism have not been observed despite our intense inventory (Appendix, see Cracraft, 1985; Cohn-Haft et al., 1997; Alerxo et al., 2011).

An important area for bird conservation was recently identified in the 'várzeas' of Monte Alegre and adjacent regions, especially because it shelters restricted-range species such as Myrmotherula kJagesi (Klages' Antwren) and Cranioleuca muelleri (Scaled Spinetail) (De Luca et al., 2009).

Fortunately, the Monte Alegre State Park was decreed in November 2001. This important 5,800 ha reserve protects significant tracts of savannas and forests (Pereira et al., 2003; Garcia et al., unpublished data 2008), and will hopefully protect the regional biodiversity in this unique area of Amazonia.

CONCLUSIONS

Although the avifaunal community of the Monte Alegre savannas is typical of other savanna habitats, we failed to record several open-country birds that occur in larger savanna areas located on the borders of Amazonia such as along the Atlantic coast or close to the 'cerrado' boundaries (southern Amazonian savannas). The isolated position and small area of the Monte Alegre savannas could explain their impoverished aspect in comparison to other, larger Amazonian savannas. Nevertheless, Monte Alegre savannas shelter the Guianan endemic Sulfur-breasted Parakeet, a vulnerable species in Pará state (SECTAM, 2006). Furthermore, the high regional species richness associated with the mosaic of different vegetation types makes Monte Alegre a key area for avifaunal conservation in Brazilian Amazonia.

ACKNOWLEDGEMENTS

We are especially grateful to M PEG and Secretaria do Meio Ambiente (SEMA-PA) for the logistical and financial support of our fieldwork. We also thank D. C. Pimentel Neto and J. N. Santa Brígida for their invaluable help in collecting and preparing specimens. G. Pereira (UFPE), A. Aleixo and F Lima (MPEG) helped to obtain data from specimens deposited in those institutions. M. Maldonado-Coelho, M. S. Hoogmoed, M. Votaw and two anonymous reviewers revised early versions of the manuscript. D. Hoffmann kindly prepared Figure 1. K. -L. Schuchmann, L. FF Silveira, D. Serripieri, G. N. Maurício, V. T. Lombardi and A. Aleixo provided important bibliography. This paper is dedicated to the memory of J. Haffer for his important contributions to the biogeographical knowledge on the Amazonian avifauna.

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Autor para correspondência:
Marcelo Ferreira de Vasconcelos.
Pontifícia Universidade Católica de Minas Gerais.
Pós-graduação em Zoologia de Vertebrados e ]]> Museu de Ciências Naturais.
Avenida Dom José Gaspar, 500,
Prédio 41 Coração Eucarístico.
Belo Horizonte, MG, Brasil.
CEP 30535-610
(mfvasconcelos@gmail.com).

Recebido em 16/03/2010
Aprovado em 28/06/2011

Responsabilidade editorial: Marinus Hoogmoed