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Breeding behavior, distribution, and conservation of the Sharp-tailed Tyrant Culicivora caudacuta (Vieillot, 1818) (Aves: Tyrannidae), a South American grassland specialist

Abstract

Culicivora caudacuta occurs in the Cerrado, Pampa and Chaco grasslands of Bolivia, Brazil, Paraguay, Argentina and Uruguay. Its breeding biology is poorly known. Here, I present a summary of the published information and new data gathered between 2003 and 2009 in southeast Brazil at Tapira, Minas Gerais. Breeding occurred during the rainy season (October to March), clutch size being three eggs. Juveniles and immatures show a different plumage from the adults, mostly brownish orange. All nests studied at Tapira showed evidence of cooperative breeding, with one helper engaged in incubation and provisioning the young. This is the first observations of this behavior for the species. The species has a wider range than currently understood and its presence in protected areas is similarly more common.

Keywords.
Nest description; Cooperative breeding; Eggs; Young plumage; Records

INTRODUCTION

The Sharp-tailed Tyrant Culicivora caudacuta (Fig. 1) is a small tyrant flycatcher (Tyrannidae: Elaeniinae) described in 1818 by Louis Jean Pierre Vieillot (1748-1830) as Muscicapa caudacuta (Le Moucherolle a queue en aiguille), based on the original description of the “Cola de agujas” (Nº 277) from Paraguay by Félix de Azara (1746-1821) (Vieillot, 1818Vieillot, L.J.P. 1818. Nouveau dictionnaire d’histoire naturelle. Paris, Chez Deterville. Tome XXI.). Later, in 1822, Coenraad Jacob Temminck (1778-1858) described the same bird under the name Muscicapa stenura (Gobe-Mouche a queue grêle), with an illustration (Fig. 2) based on specimens (Fig. 3) collected in Brazil (São Paulo state) during the expedition of the Austrian naturalist Johann Natterer (1787-1843) (Temminck, 1822Temminck, C.J. 1822. Nouveau recueil de planches colorieés d’oiseaux, pour servir de suite et de complément aux planches enluminées de Buffon. Paris, G. Levrault. Livr. 28, pl. 167, fig. 3.; Cory & Hellmayr, 1927Cory, C.B. & Hellmayr, C.E. 1927. Catalogue of birds of the Americas and the adjacent islands. Part V. Field Museum of Natural History, Zoological Series, 13:1-517. (Publication, 242)).

Figure 1
Adult Sharp-tailed Tyrant Culicivora caudacuta, (11 October 2011), Patrocínio, Minas Gerais, Brazil. Photo: RSS.

Figure 2
Illustration of Muscicapa stenura (Culicivora caudacuta), by Jean-Gabriel Prêtre, in Temminck’s work, with a wrongly drawn tail.

Figure 3
Specimen (RMNH 88812) of Culicivora caudacuta used in Temminck’s description. Photo: RSS.

Culicivora caudacuta occurs in central South America in habitats dominated by tall grasses and bushes in the Cerrado, Chaco and Pampa of Bolivia, Brazil, Paraguay, Argentina and Uruguay (Azpiroz, 1998Azpiroz, A.B. 1998. Avifauna del Valle del Lunarejo, Rivera. Achará, 1: 22-25.; Fitzpatrick, 2004Fitzpatrick, J.F. 2004. Family Tyraniidae (Tyrant-Flycatchers). In: del Hoyo, J.; Elliot, A. & Sargatal, J. (Eds.). Handbook of the Birds of the World. Volume 9: Cotingas to Pipits and Wagtails. Barcelona, Lynx Edicions. Pp. 170-463.; Lopes et al., 2009Lopes, L.E.; Malacco, G.B.; Alteff, E.F.; Vasconcelos, M.F.; Hoffmann, D. & Silveira, L.F. 2009. Range extensions and conservation of some threatened or little known Brazilian grassland birds. Bird Conservation International, 19: 1-11. http://doi.org/10.1017/S0959270909990190
http://doi.org/10.1017/S0959270909990190...
).

Brazil accounts for most of the species’ range, with records in the states of Amazonas, Maranhão, Tocantins, Bahia, Mato Grosso, Mato Grosso do Sul, Goiás, Distrito Federal, Minas Gerais, São Paulo, Paraná, Santa Catarina, and Rio Grande do Sul (Lopes et al., 2009Lopes, L.E.; Malacco, G.B.; Alteff, E.F.; Vasconcelos, M.F.; Hoffmann, D. & Silveira, L.F. 2009. Range extensions and conservation of some threatened or little known Brazilian grassland birds. Bird Conservation International, 19: 1-11. http://doi.org/10.1017/S0959270909990190
http://doi.org/10.1017/S0959270909990190...
; WikiAves, 2020WikiAves. 2020. WikiAves, a Enciclopédia das Aves do Brasil. Available: Available: http://www.WikiAves.com.br . Access: 01/06/2020.
http://www.WikiAves.com.br...
).

There are few studies of its biology and breeding behavior, the first data being gathered in Argentina in the early 20th century (Hartert & Venturi, 1909Hartert, E. & Venturi, S. 1909. Notes sur les oiseaux de la République Argentine. Novitates Zoologicae, 16(2): 159-267.), with little published since then. Most of the available information refers to breeding records and the presence of nests, young or immatures in the Brazilian states of Tocantins (Dornas & Pascoal, 2019Dornas, T. & Pascoal, W. 2019. Expansão de distribuição geográfica para a região do Jalapão de três espécies de Passeriformes encontradas no cerrado brasileiro. Cotinga, 41: 81-86.), Distrito Federal (Marini et al., 2012Marini, M.Â.; Borges, F.J.A.; Lopes, L.E.; Sousa, N.O.M.; Gressler, D.T.; Santos, L.R.; Paiva, L.V.; Duca, C.; Manica, L.T.; Rodrigues, S.S.; França, L.F.; Costa, P.M.; França, L.C.; Heming, N.M.; Silveira, M.B.; Pereira, Z.P.; Lobo, Y.; Medeiros, R.C.S. & Roper, J.J. 2012. Breeding biology of birds in the cerrado of central Brazil. Ornitología Neotropical, 23(3): 385-405.), Goiás (Hass & Silva e Silva, 2008Hass, A. & Silva e Silva, R. 2008. Culicivora caudacuta . In: Machado, A.B.M.; Drummond, G.M. & Paglia, A.P. (Org.). Livro Vermelho das Espécies Ameaçadas de Extinção. Brasília, MMA & Fundação Biodiversitas. v. 2, p. 628-629.), Minas Gerais (Ribon et al., 1995Ribon, R.; Estevão, G.R.M.; Simon, J.E.; Silva, N.F.; Pacheco, S. & Pinheiro, R.T. 1995. Aves do cerrado de Três Marias, estado de Minas Gerais. Revista Ceres, 42(242): 344-352.; Silveira, 1998Silveira, L.F. 1998. The birds of Serra da Canastra National Park and adjacent areas, Minas Gerais, Brazil. Cotinga, 10: 55-63.; Lombardi et al., 2010Lombardi, V.T.; Faetti, R.G.; D’Angelo-Neto, S.; Vasconcelos, M.F. & Gussoni, C.O.A. 2010. Notas sobre a nidificação de aves brasileiras raras e/ou pouco conhecidas. Cotinga, 32(1): 131-136.; Peixoto, 2014Peixoto, H.J.C. 2014. Ecologia e conservação de aves campestres ameaçadas de extinção no sul de Minas Gerais, Brasil. M.Sc. Dissertation. Viçosa, Universidade Federal de Viçosa. 100p.), Santa Catarina and Rio Grande do Sul (Fontana et al., 2003Fontana, C.S.; Bencke, G.A. & Reis, R.E. (Eds.). 2003. Livro vermelho da fauna ameaçada de extinção no Rio Grande do Sul. Porto Alegre, EDIPUCRS.; Rovedder et al., 2007Rovedder, C.E.; Reppening, M. & Fontana, C.S. 2007. Novos registros de ocorrência do papa-moscas-do-campo Culicivora caudacuta (Tyrannidae) para o estado do Rio Grande do Sul, Brasil. In: Congresso Brasileiro de Ornitologia, 15º. Resumos. Porto Alegre, PUCRS. Pp. 151-152.; Fontana et al., 2008Fontana, C.S.; Rovedder, C.E.; Repenning, M. & Gonçalves, M.L. 2008. Estado atual do conhecimento e conservação da avifauna dos Campos de Cima da Serra do sul do Brasil, Rio Grande do Sul e Santa Catarina. Revista Brasileira de Ornitologia, 16(4): 281-307.; Repenning et al., 2010Repenning, M.; Rovedder, C.E. & Fontana, C.S. 2010. Distribuição e biologia de aves nos campos de altitude do sul do Planalto Meridional Brasileiro. Revista Brasileira de Ornitologia, 18(4): 283-306.), and in Paraguay in Itapúa Department (Smith, 2017Smith, P. 2017. The juvenile plumage of Sharp-tailed Tyrant Culicivora caudacuta (Aves: Tyrannidae). Revista Brasileira de Ornitologia, 25(2): 122-124.).

More detailed data on the breeding biology of C. caudacuta were gathered at the Distrito Federal (Braz, 2008Braz, V.S. 2008. Ecologia e conservação das aves campestres do bioma cerrado. PhD Thesis, Departamento de Ecologia do Instituto de Ciências Biológicas, Universidade de Brasília, 178p.; Sousa & Marini, 2007Sousa, N.O. de M. & Marini, M.Â. 2007. Biologia de Culicivora caudacuta (Aves: Tyrannidae) no Cerrado, Brasília, DF. Revista Brasileira de Ornitologia, 15(4): 569-573.), Goiás (Braz, 2008Braz, V.S. 2008. Ecologia e conservação das aves campestres do bioma cerrado. PhD Thesis, Departamento de Ecologia do Instituto de Ciências Biológicas, Universidade de Brasília, 178p.) and Minas Gerais (Silva e Silva, 2006Silva e Silva, R. 2006. Biologia reprodutiva de Culicivora caudacuta (Tyrannidae) em Tapira, MG. In: Congresso Brasileiro de Ornitologia, 14º. Resumos. Ouro Preto, Universidade Federal de Ouro Preto, Sociedade Brasileira de Ornitologia.) in Brazil, and Formosa (Di Giacomo, 1996Di Giacomo, A.G. 1996. Biología reproductiva del Tachurí Coludo Culicivora caudacuta . In: Reunión Argentina de Ornitología, 9º. Libro de Resúmenes. Buenos Aires, Asociación Ornitológica del Plata. Pp. 16., 2005Di Giacomo, A.G. 2005. Culicivora caudacuta. In: Di Giacomo, A.G. & Krapovickas, S.F. (Eds.). Historia natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Inventario de la fauna de vertebrados y de la flora vascular de un área protegida del Chaco Húmedo. Aves Argentinas/Asociación Ornitológica del Plata. Pp. 342-343. (Temas de Naturaleza y Conservación 4); Di Giacomo et al., 2011Di Giacomo, A.G.; Di Giacomo, A.S. & Reboreda, J.C. 2011. Breeding biology and reproductive success of threatened Sharp-tailed Tyrant (Culicivora caudacuta) on the Chaco region of Argentina. In: Congreso de Ornitología Neotropical, 9º y Congreso Peruano de Ornitología, 8º. Libro de Resúmenes. Cusco, Unión de Ornitólogos del Perú. Pp. 442.) in Argentina.

Here I summarize all the available information on the distribution and breeding biology of C. caudacuta, and add new data mostly based on observations carried out at Tapira, Minas Gerais state, southeast Brazil, with the first observations of cooperative breeding by this species.

MATERIAL AND METHODS

Study area

The main study area is in the municipality of Tapira, southeast Minas Gerais, on the property of Fosfertil (now Mosaic Fertilizantes) known as Tapira Mining Complex (Complexo de Mineração de Tapira - CMT). This covers an area of 7,150.86 ha, including the largest phosphate mining operation in Latin America, with elevations from 950 to 1,300 m a.s.l. The area has several tailing dams (BDs) built to receive the discards from the mining process and protect the watershed downstream from the CMT (Santos et al., 2002Santos, R.L.C; Sobral, L.G.S.; Araújo, R.V.V. & Fosfertil, E.T. 2002. Produção de fosfato no Brasil: Complexo de Mineração de Tapira/Fosfertil. In: Encontro Nacional de Tratamento de Minérios e Metalurgia Extrativa, 19º. Anais. Recife.). One of these dams, BD-5 (Fig. 4) (19°49′36″S, 46°50′14″W, elevation 1,160 m), covers an area of 46 ha, and was formed by the damming of the Potreiro and Boa Vista creeks, showing varied aquatic habitats with cattails Typha domingensis (Typhaceae), muddy shores and open-water areas with different depths.

Figure 4
Partial view of BD-5 showing the study site at CMT, Tapira, Minas Gerais (18 June 2009). The arrow shows where nests were found. Photo: RSS.

This dam is surrounded by open Cerrado, with campos limpos (open grasslands) and campos sujos (grasslands with scattered bushes) showing a mix of invasive exotic grasses such as Melinis minutiflora (Poaceae) and Brachiaria decumbens (Poaceae), and native ones as Echinolaena inflexa (Poaceae), with scattered bushes, such as Baccharis dracunculifolia (Asteraceae) and Solanum lycocarpum (Solanaceae), among others.

Around the BD-5, as in the vicinity of Tapira and Araxá, there are also a few remnants of Atlantic Forest fragments, especially along watercourses, and a few plantations of Eucalyptus sp. (Myrtaceae).

The grasslands around BD-5 shelter several bird species typical of the Cerrado including Red-winged Tinamou (Rhynchotus rufescens), Spotted Nothura (Nothura maculosa), Dwarf Tinamou (Taoniscus nanus), Ocellated Crake (Micropygia schomburgkii), Red-legged Seriema (Cariama cristata), Collared Crescentchest (Melanopareia torquata), Crested Black-Tyrant (Knipolegus lophotes), Sedge Wren (Cistothorus platensis), Grassland Sparrow (Ammodramus humeralis), Blue Finch (Porphyrospiza caerulescens), Stripe-tailed Yellow-Finch (Sicalis citrina), Wedge-tailed Grass-Finch (Emberizoides herbicola), Plumbeous Seedeater (Sporophila plumbea), and Black-masked Finch (Coryphaspiza melanotis). This assemblage, as well as general habitat and setting, are very similar to those at Serra da Canastra National Park (Silveira, 1998Silveira, L.F. 1998. The birds of Serra da Canastra National Park and adjacent areas, Minas Gerais, Brazil. Cotinga, 10: 55-63.), only 25 km away.

Besides the BD-5, Culicivora caudacuta was also found and studied in another two areas in the CMT, one in the 1,430.20 ha legal reserve upstream from the dam (19°51′44″S, 46°47′52″W, elevation 1,250 m), an area with very similar habitat, and near Ribeirão do Inferno, another similar area covering 510 ha partially included in the municipality of Araxá (19°46′22″S, 46°52′48″W, elevation 1,240 m).

Dairy ranching is the main economic activity in the region where CMT is located but grazing animals are excluded from the dam area and the legal reserve. Grazing has transformed the native grasslands in open landscapes dominated by African grasses with some remnant native plants. Fires for “pasture renovation” are frequent and have a direct impact on the avifauna, especially grassland-dependent species.

Supplementary observations on the breeding biology of C. caudacuta were made in two additional areas. The first was in the municipality of Patrocínio, in the Triângulo Mineiro and Alto Paranaíba region of Minas Gerais, about 100 km from CMT. There the species was recorded more frequently in the headwaters of the Córrego Bebedouro (19°00′06″S, 46°46′07″W, elevation 1,012 m), an area belonging to Mosaic Fertilizantes, and in the headwaters of the Córrego do Mato (19°09′37″S, 46°55′28″W, elevation 1,005 m), Córrego do Fundão (19°06′49″S, 46°54′07″W, elevation 1,184 m), and Córrego Capoeira Grande (19°11′42″S, 46°55′56″W, elevation 1,049 m).

The second area, also in Minas Gerais, in the municipality of Paracatu, includes the headwaters of the Ribeirão Batalha (17°29′48″S, 47°15′48″W, elevation 880 m) at the border with Goiás state and the municipality of Catalão, about 260 km from CMT. The area has remnant veredas dominated by Mauritia palm swamps bordered by hydromorphic grasslands on undulated terrain and grasslands with mounds built by termites campos de murunduns in a landscape where most native vegetation has been replaced by soybean monoculture.

Sampling

The research done at CMT was part of a broader project to survey the avifauna in areas belonging to Fosfertil in Minas Gerais and Goiás. I made two-day visits every month totalling 158 days between June 2002 and October 2011. Observations were made ad libitum (Altmann, 1974Altmann, J. 1974. Observational study of behavior: sampling methods. Behaviour, 49(3-4): 227-267.) from sunrise to sunset over the entirety of the areas.

At Patrocínio I made monthly two-day visits between 2007 and 2012, 15 days in August 2020, and 16 days in May-June 2021 totalling 114 days, while at Ribeirão Batalha I made occasional visits between 2004 and 2009 with a total of 28 days.

Territorial groups at CMT were mist-netted in 2005, and as C. caudacuta would easily pass through the mesh I used a speaker to attract the birds while an assistant held the pole supporting one extremity of the net, closing it once a bird hit the net.

Netted birds received both metal rings from CEMAVE (Brazil’s official ringing scheme) and colour rings to allow individual identification. Bill, tail, tarsus, wing chord, nests and eggs were measured with a digital Mitutoyo® caliper; birds and eggs were weighted with Pesola® spring scales. All measurements are in mm, and when necessary other units are used.

Birds and their nests were found by active search with playback in suitable habitat inside CMT, especially near BD-5, where the birds were more habituated to human presence and, when nests were located, monitoring caused little interference in their behavior. Nests were not tagged with tape or other markers to avoid attracting predators.

Additional data on the breeding biology and localities were gathered from records at Wikiaves (http://www.wikiaves.com.br), xeno-canto (http://www.xeno-canto.org), Macaulay Library (http://www.macaulaylibrary.org), eBird (http://www.ebird.org), and from personal communications by fellow researchers.

RESULTS AND DISCUSSION

Breeding behavior

Breeding season

Culicivora caudacuta was first recorded at CMT on 26 November 2002 at BD-5, the same spot where nests were found, and in another two areas along the reservoir.

At CMT, the breeding season occurs from October to March (Silva e Silva, 2006Silva e Silva, R. 2006. Biologia reprodutiva de Culicivora caudacuta (Tyrannidae) em Tapira, MG. In: Congresso Brasileiro de Ornitologia, 14º. Resumos. Ouro Preto, Universidade Federal de Ouro Preto, Sociedade Brasileira de Ornitologia.), considering the time the first nests were found to the latest date fledglings were fed by their parents. The actual start of the nesting period is probably September since nest building take 10 to 15 days (Di Giacomo, 2005Di Giacomo, A.G. 2005. Culicivora caudacuta. In: Di Giacomo, A.G. & Krapovickas, S.F. (Eds.). Historia natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Inventario de la fauna de vertebrados y de la flora vascular de un área protegida del Chaco Húmedo. Aves Argentinas/Asociación Ornitológica del Plata. Pp. 342-343. (Temas de Naturaleza y Conservación 4)).

At Águas Emendadas Ecological Station, Central Brazil, the breeding season was bracketed between October and April, based on the presence of brooding patches, active nests and records of young being fed by adult birds (Sousa & Marini, 2007Sousa, N.O. de M. & Marini, M.Â. 2007. Biologia de Culicivora caudacuta (Aves: Tyrannidae) no Cerrado, Brasília, DF. Revista Brasileira de Ornitologia, 15(4): 569-573.; Marini et al., 2012Marini, M.Â.; Borges, F.J.A.; Lopes, L.E.; Sousa, N.O.M.; Gressler, D.T.; Santos, L.R.; Paiva, L.V.; Duca, C.; Manica, L.T.; Rodrigues, S.S.; França, L.F.; Costa, P.M.; França, L.C.; Heming, N.M.; Silveira, M.B.; Pereira, Z.P.; Lobo, Y.; Medeiros, R.C.S. & Roper, J.J. 2012. Breeding biology of birds in the cerrado of central Brazil. Ornitología Neotropical, 23(3): 385-405.). Not far away, at Chapada dos Veadeiros National Park, also in Central Brazil, breeding runs from mid-October to late March (Braz, 2008Braz, V.S. 2008. Ecologia e conservação das aves campestres do bioma cerrado. PhD Thesis, Departamento de Ecologia do Instituto de Ciências Biológicas, Universidade de Brasília, 178p.).

In Formosa, northwestern Argentina, nests were active between October and March, with no information on the time when fledglings were attended by adults (Di Giacomo, 1996Di Giacomo, A.G. 1996. Biología reproductiva del Tachurí Coludo Culicivora caudacuta . In: Reunión Argentina de Ornitología, 9º. Libro de Resúmenes. Buenos Aires, Asociación Ornitológica del Plata. Pp. 16., 2005Di Giacomo, A.G. 2005. Culicivora caudacuta. In: Di Giacomo, A.G. & Krapovickas, S.F. (Eds.). Historia natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Inventario de la fauna de vertebrados y de la flora vascular de un área protegida del Chaco Húmedo. Aves Argentinas/Asociación Ornitológica del Plata. Pp. 342-343. (Temas de Naturaleza y Conservación 4); Di Giacomo et al., 2011Di Giacomo, A.G.; Di Giacomo, A.S. & Reboreda, J.C. 2011. Breeding biology and reproductive success of threatened Sharp-tailed Tyrant (Culicivora caudacuta) on the Chaco region of Argentina. In: Congreso de Ornitología Neotropical, 9º y Congreso Peruano de Ornitología, 8º. Libro de Resúmenes. Cusco, Unión de Ornitólogos del Perú. Pp. 442.).

Culicivora caudacuta, as most species breeding at this time of the year, starts nesting at the end of the dry season when the first rains begin in September (Marini et al., 2012Marini, M.Â.; Borges, F.J.A.; Lopes, L.E.; Sousa, N.O.M.; Gressler, D.T.; Santos, L.R.; Paiva, L.V.; Duca, C.; Manica, L.T.; Rodrigues, S.S.; França, L.F.; Costa, P.M.; França, L.C.; Heming, N.M.; Silveira, M.B.; Pereira, Z.P.; Lobo, Y.; Medeiros, R.C.S. & Roper, J.J. 2012. Breeding biology of birds in the cerrado of central Brazil. Ornitología Neotropical, 23(3): 385-405.), as the rainy season is associated with greater abundance of food, especially insects, to feed the young.

The nest

The first nest (Nest 1) was found on 8 December 2003 at CMT, on the left bank of BD-5 (19°49′34″S, 46°50′28″W) when the fledglings were being attended by the parents (see below). It was a small and delicate, deep cup made of coarser plant fibres, flowers and cotton-like material bound with spider webs and lined with soft material. It was ca. 0.7 m above ground on the branches of a Baccharis dracunculifolia (Asteraceae) bush ca. 1.40 m high. The nest was 48.79 high, 40.55 deep, had an external diameter of 51.34. The empty nest was collected and is in the ornithological collection of the Museu de Zoologia da Universidade de São Paulo (MZUSP), São Paulo, Brazil.

Nest 2 (Fig. 5) was found on 8 November 2004, only 88 m from Nest 1, in the same area of BD-5 (Fig. 6). It had two very young nestlings and was also built on a B. dracunculifolia, at 1.30 m above ground.

Figure 5
Nest 2, 9 November 2004. Photo: RSS.

Figure 6
Nest site of Nest 1, 8 December 2003. Photo: RSS.

Nest 3 was found on 25 October 2005 in the same area, 48 m from Nest 1 and 40 m from Nest 2. It contained three eggs and was built 0.45 m above ground on a Campomanesia sp. (Myrtaceae) shrub about 0.90 m tall. Measurements were height 48.63, internal depth 40.93, external diameter at the rim 50.55, internal diameter 49.04.

Nest 4 was found still empty on 10 October 2008 (Fig. 7). It was built on a Vernonanthura cf. polyanthes (Asteraceae), also at BD-5 but was not monitored for long as on 27 October it looked unkempt and abandoned, apparently due to predation.

Figure 7
Nest site of Nest 4 in the same area, 11 October 2008. Photo: RSS.

At Ribeirão Batalha, on 28 November 2007, one adult was seen collecting the soft fibres (Fig. 8), likely for a nest that was not located. This strongly suggests the bird was breeding, as Di Giacomo (2005Di Giacomo, A.G. 2005. Culicivora caudacuta. In: Di Giacomo, A.G. & Krapovickas, S.F. (Eds.). Historia natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Inventario de la fauna de vertebrados y de la flora vascular de un área protegida del Chaco Húmedo. Aves Argentinas/Asociación Ornitológica del Plata. Pp. 342-343. (Temas de Naturaleza y Conservación 4)) points the adults continue to bring nest material throughout laying and incubation.

Figure 8
Adult Culicivora caudacuta collecting material for the nest. Photo: RSS.

At Chapada dos Veadeiros, another nest was found while still being built on 3 September 2007, at Fazenda Chapada das Almas (13°58′28″S, 47°27′03″W, 1,380 m), near the Rio das Almas, Alto Paraíso de Goiás, Goiás. This was in a wet grassland dominated by grasses some 40 cm tall with scattered bushes ca. 1 m tall by a riverine forest. Built ca. 85 cm above ground on a bush, the nest had grass seeds lining its structure, which was held together and attached to the branches with spider webs. (Dante Buzzetti pers. comm. May 2020).

Nests found at our study areas agree with descriptions from the Argentine Chaco (Hartert & Venturi, 1909Hartert, E. & Venturi, S. 1909. Notes sur les oiseaux de la République Argentine. Novitates Zoologicae, 16(2): 159-267.), Formosa Province (Di Giacomo et al., 2005Di Giacomo, A.G. 2005. Culicivora caudacuta. In: Di Giacomo, A.G. & Krapovickas, S.F. (Eds.). Historia natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Inventario de la fauna de vertebrados y de la flora vascular de un área protegida del Chaco Húmedo. Aves Argentinas/Asociación Ornitológica del Plata. Pp. 342-343. (Temas de Naturaleza y Conservación 4)), Central Brazil (Braz, 2008Braz, V.S. 2008. Ecologia e conservação das aves campestres do bioma cerrado. PhD Thesis, Departamento de Ecologia do Instituto de Ciências Biológicas, Universidade de Brasília, 178p.; Marini et al., 2012Marini, M.Â.; Borges, F.J.A.; Lopes, L.E.; Sousa, N.O.M.; Gressler, D.T.; Santos, L.R.; Paiva, L.V.; Duca, C.; Manica, L.T.; Rodrigues, S.S.; França, L.F.; Costa, P.M.; França, L.C.; Heming, N.M.; Silveira, M.B.; Pereira, Z.P.; Lobo, Y.; Medeiros, R.C.S. & Roper, J.J. 2012. Breeding biology of birds in the cerrado of central Brazil. Ornitología Neotropical, 23(3): 385-405.), Minas Gerais (Lombardi et al., 2010Lombardi, V.T.; Faetti, R.G.; D’Angelo-Neto, S.; Vasconcelos, M.F. & Gussoni, C.O.A. 2010. Notas sobre a nidificação de aves brasileiras raras e/ou pouco conhecidas. Cotinga, 32(1): 131-136.), and Rio Grande do Sul (Rovedder et al., 2007Rovedder, C.E.; Reppening, M. & Fontana, C.S. 2007. Novos registros de ocorrência do papa-moscas-do-campo Culicivora caudacuta (Tyrannidae) para o estado do Rio Grande do Sul, Brasil. In: Congresso Brasileiro de Ornitologia, 15º. Resumos. Porto Alegre, PUCRS. Pp. 151-152.). In all areas, nests were half-spherical, deep, solid cups made of fine plant material bound with spider silk and lined with soft material, especially fine fibres from Asteraceae seeds. They were built among the vertical branches of low bushes, mostly less than 1 m above ground.

The exception to the use of bushes seems to be the nest built close to the ground in a grass tussock found by Hass & Silva e Silva (2008Hass, A. & Silva e Silva, R. 2008. Culicivora caudacuta . In: Machado, A.B.M.; Drummond, G.M. & Paglia, A.P. (Org.). Livro Vermelho das Espécies Ameaçadas de Extinção. Brasília, MMA & Fundação Biodiversitas. v. 2, p. 628-629.) at Emas National Park, Central Brazil, suggesting some degree of plasticity. The same behavior was recorded by Lombardi et al. (2012Lombardi, V.T.; Santos, K.K.; D’Angelo-Neto, S.; Mazzoni, L.G.; Rennó, B.; Faetti, R.G.; Epifânio, A.D. & Miguel, M. 2012. Registros notáveis de aves para o sul do Estado de Minas Gerais, Brasil. Cotinga, 34: 32-45.) at Carrancas, Minas Gerais, in October 2009.

Peixoto (2014Peixoto, H.J.C. 2014. Ecologia e conservação de aves campestres ameaçadas de extinção no sul de Minas Gerais, Brasil. M.Sc. Dissertation. Viçosa, Universidade Federal de Viçosa. 100p.) also recorded an adult carrying cotton-like soft fibres to its nest on 12 October 2013 at Andrelândia, Minas Gerais, in grassland with shrubs.

The first published description of a C. caudacuta nest, from the Chaco of Santa Fe province, Argentina, was made in the early 20th century (Hartert & Venturi, 1909Hartert, E. & Venturi, S. 1909. Notes sur les oiseaux de la République Argentine. Novitates Zoologicae, 16(2): 159-267.). The nest, with three eggs, was built on a low spiny bush of a kind growing sparsely in the grasslands; it was solidly built with grass flowers and soft fibres neatly organised and lined with softer material. Its measurements were: height 70, diameter 50, internal depth 40, inner diameter 25-30 (Hartert & Venturi, 1909Hartert, E. & Venturi, S. 1909. Notes sur les oiseaux de la République Argentine. Novitates Zoologicae, 16(2): 159-267.).

A more detailed study was carried much later at El Bagual Ecological Reserve (26°10′S, 58°56′W), Formosa province, Argentina, from 1995 to 2010, with more than 500 nests (Di Giacomo, 1996Di Giacomo, A.G. 1996. Biología reproductiva del Tachurí Coludo Culicivora caudacuta . In: Reunión Argentina de Ornitología, 9º. Libro de Resúmenes. Buenos Aires, Asociación Ornitológica del Plata. Pp. 16., 2005Di Giacomo, A.G. 2005. Culicivora caudacuta. In: Di Giacomo, A.G. & Krapovickas, S.F. (Eds.). Historia natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Inventario de la fauna de vertebrados y de la flora vascular de un área protegida del Chaco Húmedo. Aves Argentinas/Asociación Ornitológica del Plata. Pp. 342-343. (Temas de Naturaleza y Conservación 4); Di Giacomo et al., 2011Di Giacomo, A.G.; Di Giacomo, A.S. & Reboreda, J.C. 2011. Breeding biology and reproductive success of threatened Sharp-tailed Tyrant (Culicivora caudacuta) on the Chaco region of Argentina. In: Congreso de Ornitología Neotropical, 9º y Congreso Peruano de Ornitología, 8º. Libro de Resúmenes. Cusco, Unión de Ornitólogos del Perú. Pp. 442.). These were built on annual herbs, the most important ones being Vernonia chamaedrys (Asteraceae) (more than 30% of the nests), Desmodium cuneatum (Fabaceae) (13%), Solidago chilensis (Asteraceae) (10%), and Eupatorium ivaefolium (Asteraceae) (8%), with additional 10 species accounting for the remainder.

As in our study area, nests were supported by several vertical or nearly vertical branches or, in some cases, inflorescences as in V. cognata, E. eburneum and E. elegans. Average nest height above ground was 1 m, ranging from 0.45 m to 1.8 m. Most nests were quite visible and similar to each other, built in the shape of compact, well-built half-spheres, straw or yellowish coloured with white inner lining. Nests are built with fine plant matter such as dry fibres, petioles, Poaceae and Asteraceae flowers bound with silk from spider webs or egg sacs. The same material is used to anchor the nest to supporting branches, which varied from three to eight. The nest chamber is lined with very soft material, mostly fine fibres from Asteraceae inflorescences (Di Giacomo, 2005Di Giacomo, A.G. 2005. Culicivora caudacuta. In: Di Giacomo, A.G. & Krapovickas, S.F. (Eds.). Historia natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Inventario de la fauna de vertebrados y de la flora vascular de un área protegida del Chaco Húmedo. Aves Argentinas/Asociación Ornitológica del Plata. Pp. 342-343. (Temas de Naturaleza y Conservación 4)).

Additional nests were found at Brasília National Park (15°47′S, 47°56′W), central Brazil, on 12 November 2004, and two nests were found at Chapada dos Veadeiros National Park (14°05′S, 47°40′W), Goiás state, in November 2005 and November 2006 (Braz, 2008Braz, V.S. 2008. Ecologia e conservação das aves campestres do bioma cerrado. PhD Thesis, Departamento de Ecologia do Instituto de Ciências Biológicas, Universidade de Brasília, 178p.). The nests were attached to vertical branches and shaped as deep cups. These were built with fine plant material including dried fibres, petioles, grass flowers and leaves woven with spider webs. Height above ground ranged from 64 cm to 1 m. Measurements of the nest found in November 2006 were inner diameter 20.9, outer diameter 44.4 and depth 54.8 (Braz, 2008Braz, V.S. 2008. Ecologia e conservação das aves campestres do bioma cerrado. PhD Thesis, Departamento de Ecologia do Instituto de Ciências Biológicas, Universidade de Brasília, 178p.).

Another study at Águas Emendadas Ecological Station (15°32′S, 47°36′W, 1,040 m), Distrito Federal, between 2004 and 2007, provided information on three nests. One, with three eggs, was found on 10 November 2005 at 0.34 m above ground on an Esenbeckia pumila (Rutaceae) bush 0.50 m tall. The other nests, one found on 22 and the other on 24 November 2007, were about 150 m apart in an open grassland. Both were on Eremanthus glomerulatus (Asteraceae) bushes about 0.55 m high, built 0.34 m and 0.53 m above ground (Sousa & Marini, 2007Sousa, N.O. de M. & Marini, M.Â. 2007. Biologia de Culicivora caudacuta (Aves: Tyrannidae) no Cerrado, Brasília, DF. Revista Brasileira de Ornitologia, 15(4): 569-573.). In the same area another nest was found on 4 October 2009 in a campo sujo area built 38 cm above ground and having three eggs (Marini et al., 2012Marini, M.Â.; Borges, F.J.A.; Lopes, L.E.; Sousa, N.O.M.; Gressler, D.T.; Santos, L.R.; Paiva, L.V.; Duca, C.; Manica, L.T.; Rodrigues, S.S.; França, L.F.; Costa, P.M.; França, L.C.; Heming, N.M.; Silveira, M.B.; Pereira, Z.P.; Lobo, Y.; Medeiros, R.C.S. & Roper, J.J. 2012. Breeding biology of birds in the cerrado of central Brazil. Ornitología Neotropical, 23(3): 385-405.).

The following additional accounts of nests of C. caudacuta found in Brazil provide more limited but important information as to localities and breeding dates for the species.

A nest found in late October and another in early November 2000 were cup-shaped and built near the ground inside grass clumps at Emas National Park (18°08′S, 52°56′W), Mineiros municipality, Goiás state (Hass & Silva e Silva, 2008Hass, A. & Silva e Silva, R. 2008. Culicivora caudacuta . In: Machado, A.B.M.; Drummond, G.M. & Paglia, A.P. (Org.). Livro Vermelho das Espécies Ameaçadas de Extinção. Brasília, MMA & Fundação Biodiversitas. v. 2, p. 628-629.).

Also at Emas National Park, but at Chapadão do Céu (18°17′48″S, 52°46′53″W, 850 m) a nest with three eggs attended by one incubating adult was found on 7 November 2004. The nest had been built 84 cm above ground on a Vernonia sp. (Asteraceae), in a campo sujo dominated by Tristachya leiostachya (Poaceae), and measured: total height 55, inner depth 30, external diameter 56, inner diameter at the rim 43 (Dante Buzzetti pers. comm. May 2020).

Southeast of Serra de Carrancas (21°27′S, 44°37′W, ca. 1,250 m), Carrancas, Minas Gerais state, a nest with three eggs was found on 21 October 2008. This was on a Diospyros hispida (Ebenaceae) bush in a small patch of campo sujo by a forest. The nest had been built with grass inflorescences and fine plant fibre (silk cotton), the latter mostly lining the incubation chamber. As other nests, the material was bound by spider webs, resulting in a soft and light, but resistant, construction. Measurements were: depth of the incubation chamber 30, external height 45, inner diameter 30 and external diameter 55 (Lombardi et al., 2010Lombardi, V.T.; Faetti, R.G.; D’Angelo-Neto, S.; Vasconcelos, M.F. & Gussoni, C.O.A. 2010. Notas sobre a nidificação de aves brasileiras raras e/ou pouco conhecidas. Cotinga, 32(1): 131-136.).

On 9 November 2006 a nest with two eggs was found at the headwaters of the Arroio Macena (28°30′55″S, 50°47′56″W, 940 m), Vacaria, Rio Grande do Sul. This was described as a delicate cup-shaped structure covered by silk cotton, built on a bush in a small wetland near a fallow dominated by Senecio sp. (Asteraceae) (Rovedder et al., 2007Rovedder, C.E.; Reppening, M. & Fontana, C.S. 2007. Novos registros de ocorrência do papa-moscas-do-campo Culicivora caudacuta (Tyrannidae) para o estado do Rio Grande do Sul, Brasil. In: Congresso Brasileiro de Ornitologia, 15º. Resumos. Porto Alegre, PUCRS. Pp. 151-152.).

One adult was photographed bringing nest material (silk cotton) to its nest at Itirapina Ecological Station (22°13′S, 47°54′W, 740 m), Itirapina, São Paulo state, on 27 November 2009. The nest had been built in an isolated bush in open grassland (Motta-Jr. et al., 2020Motta-Jr., J.C.; Ritter, A.C.; Ferrari, A.; Paschotto, F.R.; De Sordi, L.; Morandini, R.S.; Barros, F.M.; Ruffino, P.H.P. & Braga, A.C.R. 2020. Aves e seus ambientes na Estação Ecológica de Itirapina, SP. São Paulo, Instituto de Biociências da Universidade de São Paulo.).

Further nests found in different Brazilian localities have been documented with photographs. On 12 November 2019 a nest was found at Barbacena, Minas Gerais state, and on 10 January 2008 a nest with eggs was located at Indianópolis, Minas Gerais. Another was photographed at Piraí do Sul, Paraná state, on 18 January 2014 (WikiAves, 2020WikiAves. 2020. WikiAves, a Enciclopédia das Aves do Brasil. Available: Available: http://www.WikiAves.com.br . Access: 01/06/2020.
http://www.WikiAves.com.br...
).

Eggs, clutch size, incubation

On 25 October 2005, Nest 3 had three light cream eggs (Fig. 9), measuring 13.31 × 12.26, 13.56 × 11.75 and 13.51 × 11.58, respectively. The adults were seen taking turns at incubation (Fig. 10), but its length could not be determined.

Figure 9
Nest 3 of Culicivora caudacuta with eggs. Photo: RSS.

Figure 10
Nest 3 of Culicivora caudacuta, with adult incubating, 25 October 2005. Photo: RSS.

A nest found at Emas National Park, Goiás, on 7 November 2004 had three light cream eggs measuring 14.3 × 11.5, 14.2 × 11.3 e 14.0 × 11.4. When revisited on 13 November 2004 it was abandoned (Dante Buzzetti pers. comm. May 2020).

Our small sample precludes generalisations but a clutch size of three and egg colour agree with previous information from Argentina (Hartert & Venturi, 1909Hartert, E. & Venturi, S. 1909. Notes sur les oiseaux de la République Argentine. Novitates Zoologicae, 16(2): 159-267.; Di Giacomo, 2005Di Giacomo, A.G. 2005. Culicivora caudacuta. In: Di Giacomo, A.G. & Krapovickas, S.F. (Eds.). Historia natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Inventario de la fauna de vertebrados y de la flora vascular de un área protegida del Chaco Húmedo. Aves Argentinas/Asociación Ornitológica del Plata. Pp. 342-343. (Temas de Naturaleza y Conservación 4); Di Giacomo et al., 2011Di Giacomo, A.G.; Di Giacomo, A.S. & Reboreda, J.C. 2011. Breeding biology and reproductive success of threatened Sharp-tailed Tyrant (Culicivora caudacuta) on the Chaco region of Argentina. In: Congreso de Ornitología Neotropical, 9º y Congreso Peruano de Ornitología, 8º. Libro de Resúmenes. Cusco, Unión de Ornitólogos del Perú. Pp. 442.), Central Brazil (Sousa & Marini, 2007Sousa, N.O. de M. & Marini, M.Â. 2007. Biologia de Culicivora caudacuta (Aves: Tyrannidae) no Cerrado, Brasília, DF. Revista Brasileira de Ornitologia, 15(4): 569-573.; Braz, 2008Braz, V.S. 2008. Ecologia e conservação das aves campestres do bioma cerrado. PhD Thesis, Departamento de Ecologia do Instituto de Ciências Biológicas, Universidade de Brasília, 178p.) and Minas Gerais (Lombardi et al., 2010Lombardi, V.T.; Faetti, R.G.; D’Angelo-Neto, S.; Vasconcelos, M.F. & Gussoni, C.O.A. 2010. Notas sobre a nidificação de aves brasileiras raras e/ou pouco conhecidas. Cotinga, 32(1): 131-136.).

All published information puts egg size range at 14-18.7 × 10.3-12.4 mm, the eggs found at CMT being unusually short. The significance of this, if any, remains to be assessed.

Di Giacomo (2005Di Giacomo, A.G. 2005. Culicivora caudacuta. In: Di Giacomo, A.G. & Krapovickas, S.F. (Eds.). Historia natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Inventario de la fauna de vertebrados y de la flora vascular de un área protegida del Chaco Húmedo. Aves Argentinas/Asociación Ornitológica del Plata. Pp. 342-343. (Temas de Naturaleza y Conservación 4)) and Di Giacomo et al. (2011Di Giacomo, A.G.; Di Giacomo, A.S. & Reboreda, J.C. 2011. Breeding biology and reproductive success of threatened Sharp-tailed Tyrant (Culicivora caudacuta) on the Chaco region of Argentina. In: Congreso de Ornitología Neotropical, 9º y Congreso Peruano de Ornitología, 8º. Libro de Resúmenes. Cusco, Unión de Ornitólogos del Perú. Pp. 442.) state that incubation begins after the penultimate egg is laid and lasts 15 to 16 days. The pairs may have a second clutch during the same nesting season but this is raised in a newly built nest in a different part of the territory.

Nestlings, juveniles and parental care

Nest 1 was found on 8 December 2003 when the young had already left and were being attended by the parents. The short-tailed fledglings were orange-brown colour (Fig. 11). They remained in the nearby shrubs and were seen to be fed by three different adults (sometimes at the same time) with dragonflies, horseflies and other flying insects (Figs. 12 and 13) until the 11 December. This family group provided the first published photographs of young C. caudacuta (Silva e Silva, 2004Silva e Silva, R. 2004. Magia no cerrado: aves na imensidão. São Paulo, Editora DBA.).

Figure 11
Fledglings of Culicivora caudacuta near Nest 1. Photo: RSS.

Figure 12
Two adults feeding a nestling with dragonflies, 8 December 2003. Photo: RSS.

Figure 13
Adult feeding a nestling with a fly, 8 December 2003. Photo: RSS.

Later, on 16-17 January 2004, the young already showed longer tails and foraged by themselves, but always associated with the three adult birds.

On 8 November 2004 Nest 2 had two downy nestlings only a few days old attended by three adults that fed them beetles, dragonflies and flies and removed faecal sacs (Fig. 14), which were dropped 10 m away from the nest.

Figure 14
Adult removing a faecal sac from Nest 2 after feeding the nestlings. Photo: RSS.

On 15 December 2004 the young had already fledged and were foraging by themselves but were still followed by the three adults. A few times two adults were seen to harass the young to direct them back near to the nest.

On 7 and 8 November 2005 Nest 3 had one egg and one nestling (Fig. 15) already covered in orange-cream feathers, lighter on the underparts, and scattered down. It was also attended by three adults that fed it with horseflies, small grasshoppers, flies and dragonflies (Fig. 16) mostly caught 5-10 m from the nest. The nestling would produce a faecal sac soon after being fed and one of the adults would at once take it and drop five meters or so from the nest, closer than seen in the previous nest. The nestling was able to raise itself from the nest but kept still most of the time, calling when it heard the voice or wing sounds of adults nearby.

Figure 15
Culicivora caudacuta nestling. Photo: RSS.

Figure 16
Adult feeding the nestling. Photo: RSS.

Nests were not found the following nesting season at CMT but breeding was confirmed on 29 November 2006 when two adults, one previously banded at the site on 2 December 2005, and one short-tailed fledgling were seen. The young bird already showed a long tail and was following the adults when found again on 28 December 2006 and 24 January 2007.

The following day one adult pair followed by two young was found in the same area, with two adults and three young on 7 March 2007. One of these showed a whiter supercilium, a characteristic of an older bird than its siblings suggesting that the young from two consecutive clutches were following their parents.

Additional information on youngsters was gathered at other sites. At Patrocínio, around the headwaters of Córrego Bebedouro, two adults and one young were found on 6 February 2009, with one adult being captured and banded. Two adults and one young were also seen on 14 and 16 January 2012.

Also at Patrocínio, at the headwaters of Córrego Capoeira Grande, three adults and one young were seen on 1, 6 and 13 March 2011 (Fig. 17). The following season, two adults followed by two young were found on 12 January 2012.

Figure 17
Juvenile Culicivora caudacuta, with adult in the background. Photo: RSS.

At Paracatu, around Ribeirão Batalha, three adults and one young (Fig. 18) were found near a Mauritia palm swamp on 27 January 2009.

Figure 18
Juvenile of Culicivora caudacuta, Ribeirão Batalha. Photo: RSS.

The available literature provides further data on the presence of young C. caudacuta in other localities, mostly in Brazil. Although most mentions are just brief it is worth listing them in order to add more information.

In Brazil, the first data on young Sharp-tailed Tyrants are from 12 October 1988 at Fazenda Perdões (18°11′41″S, 45°25′17″W, 810-730 m), Três Marias, Minas Gerais. Two adults were seen feeding two juveniles showing a paler colour compared to the adults, lacking the black on the cap (Ribon et al., 1995Ribon, R.; Estevão, G.R.M.; Simon, J.E.; Silva, N.F.; Pacheco, S. & Pinheiro, R.T. 1995. Aves do cerrado de Três Marias, estado de Minas Gerais. Revista Ceres, 42(242): 344-352.).

On 17 February 1991 two fledglings were recorded calling at Itirapina Ecological Station (Parker III & Willis, 1997Parker III, T.A. & Willis, E.O. 1997. Notes on three tiny grassland flycatchers, with comments on the disappearance of South American fire-diversified savannas. In: Remsen-Jr., J. (Ed.). Studies in Neotropical Ornithology Honoring Ted Parker. Ornithological Monographs, 48: 549-555.).

In December 1996, a family group with two dependent young was found at Serra da Canastra National Park (20°15′S, 46°37′W), São Roque de Minas, Minas Gerais (Silveira, 1998Silveira, L.F. 1998. The birds of Serra da Canastra National Park and adjacent areas, Minas Gerais, Brazil. Cotinga, 10: 55-63.). Adults and three young were seen on 12 February 2002 (Dante Buzetti pers. comm. May 2020), and on 8 February 2006 three adults and one young were seen in the same park (RSS).

At Brasília National Park, a nest was found on 12 November 2004 with three nestlings being fed by two adults. The young had emerging wing, tail and body feathers and an average weight of 4.33 g, but were predated five days later (Braz, 2008Braz, V.S. 2008. Ecologia e conservação das aves campestres do bioma cerrado. PhD Thesis, Departamento de Ecologia do Instituto de Ciências Biológicas, Universidade de Brasília, 178p.).

At Chapada dos Veadeiros National Park a nest was found in November 2006 with two nestlings still showing a mostly naked body, with just some down on the cap, back and wings. These had an average weight of 1.5 g and were taken by a predator four days later (Braz, 2008Braz, V.S. 2008. Ecologia e conservação das aves campestres do bioma cerrado. PhD Thesis, Departamento de Ecologia do Instituto de Ciências Biológicas, Universidade de Brasília, 178p.).

During a study at Águas Emendadas Ecological Station, two young in different family groups were found in November 2004 and April 2005. They showed completely cinnamon-coloured faces, short tail feathers and limited flight but, two months later, one of them had a full black mask and made several vocalizations (Sousa & Marini, 2007Sousa, N.O. de M. & Marini, M.Â. 2007. Biologia de Culicivora caudacuta (Aves: Tyrannidae) no Cerrado, Brasília, DF. Revista Brasileira de Ornitologia, 15(4): 569-573.). In the same area, a nest was found on 4 October 2009. It had three eggs that hatched around 17 October, but the nestlings were preyed on about eight days later (Marini et al., 2012Marini, M.Â.; Borges, F.J.A.; Lopes, L.E.; Sousa, N.O.M.; Gressler, D.T.; Santos, L.R.; Paiva, L.V.; Duca, C.; Manica, L.T.; Rodrigues, S.S.; França, L.F.; Costa, P.M.; França, L.C.; Heming, N.M.; Silveira, M.B.; Pereira, Z.P.; Lobo, Y.; Medeiros, R.C.S. & Roper, J.J. 2012. Breeding biology of birds in the cerrado of central Brazil. Ornitología Neotropical, 23(3): 385-405.).

At Jalapão, Mateiros, Tocantins state, in a patch of cerrado and campo sujo, bordering a soy plantation (10°27′56″S, 46°08′34″W, 770 m), five birds, including a recently fledged young bird, were seen on 24 March 2018 (Dornas & Pascoal, 2019Dornas, T. & Pascoal, W. 2019. Expansão de distribuição geográfica para a região do Jalapão de três espécies de Passeriformes encontradas no cerrado brasileiro. Cotinga, 41: 81-86.).

At Carrancas, Minas Gerais, young were found in May 2009 and January 2010, and in the same region, at Serra da Chapada das Perdizes (1,500 m), on the border of Minduri municipality, further young were recorded in January 2010 (Lombardi et al., 2012Lombardi, V.T.; Santos, K.K.; D’Angelo-Neto, S.; Mazzoni, L.G.; Rennó, B.; Faetti, R.G.; Epifânio, A.D. & Miguel, M. 2012. Registros notáveis de aves para o sul do Estado de Minas Gerais, Brasil. Cotinga, 34: 32-45.).

On 7 October 2005, at Fazenda Indiana (20°25′S, 56°39′W, 120 m), Rio Salobra, Bodoquena, Mato Grosso do Sul, two adults with two young were observed in a recently burnt fallow near a rice plantation (Pivatto et al., 2006Pivatto, M.A.C.; Manço, D.G.; Straube, F.C.; Urben-Filho, A. & Milano, M. 2006. Aves do Planalto da Bodoquena, estado do Mato Grosso do Sul (Brasil). Atualidades Ornitológicas, 129: 28-53.).

At Fazenda Pai João (28°10′58″S, 50°38′28″W, 1,005 m), Capão Alto, Santa Catarina, one pair of adults followed by a juvenile was found on 13 January 2013 in a grassy area with sparse flowering bushes (RSS). At the same locality, on 22 January 2012, one young and one adult were photographed together, while on 30 January 2012, one young was photographed (Wikiaves, 2020WikiAves. 2020. WikiAves, a Enciclopédia das Aves do Brasil. Available: Available: http://www.WikiAves.com.br . Access: 01/06/2020.
http://www.WikiAves.com.br...
).

The nest found with two eggs at Vacaria, Rio Grande do Sul, on 9 November 2006 (Rovedder et al., 2007Rovedder, C.E.; Reppening, M. & Fontana, C.S. 2007. Novos registros de ocorrência do papa-moscas-do-campo Culicivora caudacuta (Tyrannidae) para o estado do Rio Grande do Sul, Brasil. In: Congresso Brasileiro de Ornitologia, 15º. Resumos. Porto Alegre, PUCRS. Pp. 151-152.) was photographed on 20 November 2006 with one nestling (Repenning et al., 2010Repenning, M.; Rovedder, C.E. & Fontana, C.S. 2010. Distribuição e biologia de aves nos campos de altitude do sul do Planalto Meridional Brasileiro. Revista Brasileira de Ornitologia, 18(4): 283-306.).

Near Ibirapuitã Biological Reserve, at Fazenda do Beto, Alegrete, Rio Grande do Sul, four individuals, including young, were found in early February 2001 (Fontana et al., 2003Fontana, C.S.; Bencke, G.A. & Reis, R.E. (Eds.). 2003. Livro vermelho da fauna ameaçada de extinção no Rio Grande do Sul. Porto Alegre, EDIPUCRS.).

In Paraguay, on 13 February 1995, at Aguara Ñu, Bosque Mbaracayú Natural Reserve (24°09′S, 55°17′W), Canindeyu departament, one adult and one immature were seen (Madroño Nieto & Esquivel, 1997Madroño Nieto, A. & Esquivel, E.Z. 1997. Noteworthy records and range extensions of some birds from the Reserva Natural del Bosque Mbaracayú (Mbaracayú Forest Nature Reserve), Departamento de Canindeyú, Paraguay. Bulletin of the British Ornithologists’ Club, 117(3): 166-176.).

Also in Paraguay, at Estancia Kanguery (26°30′S, 55°47′W), Itapúa department, San Rafael National Park, three recently fledged juveniles were seen, and photographed together with two adults on 5 April 2016 (Smith, 2017Smith, P. 2017. The juvenile plumage of Sharp-tailed Tyrant Culicivora caudacuta (Aves: Tyrannidae). Revista Brasileira de Ornitologia, 25(2): 122-124.).

In Argentina, at Estancia Virocay (28°15′S, 55°57′W), Corrientes, two young following a pair of adults were found on 16 December 2010 (Marino et al., 2013Marino, G.D.; Miñarro, F.; Zaccagnini, M.E. & López-Lanús, B. (Eds.). 2013. Pastizales y sabanas del cono sur de Sudamerica: Iniciativas para su conservación en la Argentina. Buenos Aires, Aves Argentinas/Asociación ornitológica del Plata, Fundación Vida Silvestre Argentina e Instituto Nacional de Tecnología Agropecuaria. (Monografia de Aves Argentinas n. 9).).

The young acquire full adult plumage five to six months after fledging and juvenile-plumaged birds were not recorded during the cold and dry winter months.

The juvenile plumage of C. caudacuta remained undescribed for a long time despite at least two juveniles having been collected by Johann Natterer in the early 19th century. Among the eleven C. caudacuta collected by Natterer in Brazil, including nine from São Paulo and two from Paraná (Pelzeln, 1868-1870Pelzeln, A. von. 1868-1870. Zur Ornithologie Brasiliens. Resultate von Johann Natterers Reisen in den Jahren 1817 bis 1835. Parts I-IV. Viena, A. Pichler’s Witwe & Sohn.), two specimens, NMW 17822 and NHMUK 1888.1.13.331 (Fig. 19), both collected on 11 February 1821 at Itararé, São Paulo state, are juvenile. The specimens are held at the collections of the Naturhistorisches Museum Wien (NMW), at Vienna, Austria (Hans Berg-Martin in litt., 2020), and Natural History Museum, Tring, UK (Hein van Grow in litt., 2021).

Figure 19
Immature specimen of Culicivora caudacuta at Natural History Museum, Tring. Photo: Hein van Grow.

Another specimen (ZSM 32700), which had probably just fledged judging from its short tail and colour, was collected in Paraguay on 4 December 1931 and is housed in the ornithological collections of the Zoologische Staatssammlung München (ZSM), Munich, Germany (Fig. 20), remained unknown until now (Markus Unsöld in litt., 2020). This is one of four specimens of C. caudacuta collected in Paraguay during the 1931 expedition (Laubmann, 1940Laubmann, A. 1940. Wissenschaftliche Ergebnisse der Deutschen Gran Chaco-Expedition: Die vögel von Paraguay. Sttutgart, Strecker und Schröder. v. 2.).

Figure 20
Specimen of young Culicivora caudacuta at Zoologische Staatssammlung München. Photo: Markus Unsöld.

Despite this, the juvenile plumage of C. caudacuta was first shown in a field guide by Sigrist (2004Sigrist, T. 2004. Aves do Brasil: uma visão artística. Vinhedo, Avis Brasilis., 2007Sigrist, T. 2007. Guia de campo: aves do Brasil oriental. Vinhedo, Avis Brasilis.), based on photographs taken at CMT by RSS.

As described in this guide, the juvenile plumage of C. caudacuta, from the time they fledge until acquiring adult plumage, is quite distinctive and unlikely to be confused with the non-breeding plumage or female of the Bearded Tachuri Polystictus pectoralis, as shown by Smith (2017Smith, P. 2017. The juvenile plumage of Sharp-tailed Tyrant Culicivora caudacuta (Aves: Tyrannidae). Revista Brasileira de Ornitologia, 25(2): 122-124.).

The female and young male of P. pectoralis show most of the head blackish, contrary to C. caudacuta which, after fledging and while still showing a short tail, has practically no black on the head.

As the tail grows and becomes as long as the head and body, the young start showing the first black streaks on the cap and show a very different jizz from the short, broad-tailed P. pectoralis.

Of 1,698 photos of C. caudacuta available at Wikiaves (2020WikiAves. 2020. WikiAves, a Enciclopédia das Aves do Brasil. Available: Available: http://www.WikiAves.com.br . Access: 01/06/2020.
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), up to 16 May 2020, only 47 show juvenile birds. These were taken at Tocantins, Goiás, Distrito Federal, Bahia, Minas Gerais, Paraná, Santa Catarina and Rio Grande do Sul, showing the confirmed breeding range includes most of the species’ range in Brazil. The presence of young in the months of October, November, December, January, February, March, April, and May agrees with the observations in the literature and the ones presented above, and suggest more than one brood can be raised per season.

Cooperative breeding and helpers

A common feature of family groups of C. caudacuta was the presence of three adult birds sharing the duties of incubation and provisioning of the young (Silva e Silva, 2006Silva e Silva, R. 2006. Biologia reprodutiva de Culicivora caudacuta (Tyrannidae) em Tapira, MG. In: Congresso Brasileiro de Ornitologia, 14º. Resumos. Ouro Preto, Universidade Federal de Ouro Preto, Sociedade Brasileira de Ornitologia.).

When Nest 1 and its fledglings were found in December 2003, three adults were found caring for the two young that remained next to the nest. Even when the young were already foraging by themselves, on 16 January 2004 and 2 February 2004, the three adults stayed near them, forming a family group, but only two of the adults would react aggressively to playback.

At Nest 2, found with two nestlings, three adults took turns feeding them and removing faecal sacs. The young continued to be attended by the three adults after they left the nest but, again, only two of the adult birds would approach in response to playback, the third remaining some distance away uttering softer and lower calls. Nest 3 also had three adults attending its single young from incubation until well after it had fledged.

The same behavior was also recorded in other sites we visited in Minas Gerais. At Ribeirão Batalha, Paracatu, one young bird attended by three adults was recorded on 9 November 2005 and 27 January 2009. The same grouping was seen on 1, 6 and 13 March 2011 at Patrocínio, near Córrego Capoeira Grande, when three adults and one young bird were attracted with playback.

The same grouping was found during a study on C. caudacuta at Águas Emendadas Ecological Station, where one young bird was attended by three adults, one of these providing food with higher frequency (Sousa & Marini, 2007Sousa, N.O. de M. & Marini, M.Â. 2007. Biologia de Culicivora caudacuta (Aves: Tyrannidae) no Cerrado, Brasília, DF. Revista Brasileira de Ornitologia, 15(4): 569-573.).

Cockburn (2006Cockburn, A. 2006. Prevalence of different modes of parental care in birds. Proceedings of the Royal Society B, 273: 1375-1383.) states that among the 1,097 New World suboscines, cooperative breeding is consistently rare, found in just 1% of the species. By contrast, a significantly larger proportion of all oscines are cooperative breeders (13%).

Although Fitzpatrick (2004Fitzpatrick, J.F. 2004. Family Tyraniidae (Tyrant-Flycatchers). In: del Hoyo, J.; Elliot, A. & Sargatal, J. (Eds.). Handbook of the Birds of the World. Volume 9: Cotingas to Pipits and Wagtails. Barcelona, Lynx Edicions. Pp. 170-463.) states no tyrant flycatcher is known to exhibit cooperative breeding on a regular basis, Griesser & Suzuki (2016Griesser, M. & Suzuki, T.N. 2016. Occasional cooperative breeding in birds and the robustness of comparative analyses concerning the evolution of cooperative breeding. Zoological Letters, 2(1): 1-11.) show that among the Tyrannidae, six species show cooperative breeding.

According to Griesser & Suzuki, (2016Griesser, M. & Suzuki, T.N. 2016. Occasional cooperative breeding in birds and the robustness of comparative analyses concerning the evolution of cooperative breeding. Zoological Letters, 2(1): 1-11.), most (93%) helpers are offspring that remain associated with their parents until the next breeding season and provide alloparental care at the nest of their parents or close relatives.

Following this pattern, it is likely the helpers among C. caudacuta are young from the previous nesting season that remained in the natal territory through the following season.

During the fieldwork recently carried out in Patrocínio, Minas Gerais, in August 2020, three groups of three individuals each of C. caudacuta were found, where it was noted that only two of these individuals responded promptly to playback, while the third individual was more distant and responded with a much softer and weaker vocalization, being possibly a young bird accompanying its parents.

Detailed studies, including genetic profiling, are necessary to further knowlege on this behavior in C. caudacuta and the few tyrant flycatchers also showing helper behavior like Streamer-tailed Tyrant Gubernetes yetapa (Wagener et al., 2019Wagener, T.L.S.; Santos, L.E.S.; Almeida, R.A.; Prestes, N.P. & Krügel, M.M. 2019. Gubernetes yetapa (Aves: Tyrannidae) as a nest helper: a report. In: Congresso Brasileiro de Ornitologia, 26º. Livro de Resumos. Vila Velha, ES. Pp. 130.).

Birds’ measurements

Eleven birds were captured for ringing in the study sites: nine, including one young, at Tapira between 1 and 2 December 2005, one at Paracatu on 28 December 2005, and another at Patrocínio on 6 February 2009. The measurements are in Table 1. These fall in the range of the 13 birds captured at Águas Emendadas Ecological Station (Sousa & Marini, 2007Sousa, N.O. de M. & Marini, M.Â. 2007. Biologia de Culicivora caudacuta (Aves: Tyrannidae) no Cerrado, Brasília, DF. Revista Brasileira de Ornitologia, 15(4): 569-573.).

Table 1
Measurements (mm) and body mass (g) of Culicivora caudacuta from Tapira (n = 9), Paracatu (n = 1) and Patrocínio (n = 1).

Distribution

The first information on the distribution of C. caudacuta was supplied in the 18th century by Félix de Azara and mentioned in the species’ description, where Paraguay was assumed as the type locality (Vieillot, 1818Vieillot, L.J.P. 1818. Nouveau dictionnaire d’histoire naturelle. Paris, Chez Deterville. Tome XXI.). Nevertheless, Azara’s observations also covered nearby parts of Argentina and Brazil (Beddall, 1975Beddall, B.G. 1975. “Un Naturalista Original”: Don Félix de Azara, 1746-1821. Journal of the History of Biology, 8(1): 15-66.) where C. caudacuta is known to occur, the presence of the species in Brazil having already been noted in the early 19th century (Temminck, 1822Temminck, C.J. 1822. Nouveau recueil de planches colorieés d’oiseaux, pour servir de suite et de complément aux planches enluminées de Buffon. Paris, G. Levrault. Livr. 28, pl. 167, fig. 3.). The assignment of Paraguay as type locality is accordingly best seen as tentative.

It took much longer for the actual distribution of C. caudacuta to be better understood as including parts of Brazil, Bolivia, Paraguay, and Argentina (Lopes et al., 2009Lopes, L.E.; Malacco, G.B.; Alteff, E.F.; Vasconcelos, M.F.; Hoffmann, D. & Silveira, L.F. 2009. Range extensions and conservation of some threatened or little known Brazilian grassland birds. Bird Conservation International, 19: 1-11. http://doi.org/10.1017/S0959270909990190
http://doi.org/10.1017/S0959270909990190...
). However, coverage was limited and the gaps that were shown may not be real.

Technological progress, especially the greater availability of digital cameras and sound recorders, with the rise of birdwatching and on-line platforms for ornithological data, such as Wikiaves, xeno-canto and eBird has increased coverage in a dramatic way and resulted in a jump in the known localities for the species.

On the other hand, greater coverage and the resulting increase in the number of localities mean that caution is required in considering claims that C. caudacuta may be expanding its range.

The recent data show that the area of occupancy of C. caudacuta has been underestimated. Using Minas Gerais state (58,652,800 ha), in Brazil, as an example, the species was previously known from just three localities (Lins, 1998Lins, L.V. 1998. Culicivora caudacuta. Pp. 337-338. In: Machado, A.B.M.; Fonseca, G.A.B.; Machado, R.B.; Aguiar, L.M.S. & Lins, L.V. (Eds.). Livro Vermelho das Espécies Ameaçadas de Extinção da fauna de Minas Gerais. Belo Horizonte, Fundação Biodiversitas.), later jumping to 20 (Lopes et al., 2009Lopes, L.E.; Malacco, G.B.; Alteff, E.F.; Vasconcelos, M.F.; Hoffmann, D. & Silveira, L.F. 2009. Range extensions and conservation of some threatened or little known Brazilian grassland birds. Bird Conservation International, 19: 1-11. http://doi.org/10.1017/S0959270909990190
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), and with the compilation presented here this jumps to 134 localities (Appendix 1; Fig. 21).

Figure 21
Records of Culicivora caudacuta. Yellow stars indicate museum specimens, dots indicate other categories of records. See Appendix 1 for details.

Culicivora caudacuta is mostly associated with open grassland and savanna of the Pampa and Cerrado biomes characterized by a dense ground cover of grasses and scattered, well-spaced low bushes, occasionally near watercourses, but it also occurs in areas where the original vegetation has already been contaminated by exotic grasses.

Records show it occurs from elevations of 60 m in the Chaco of Argentina and Paraguay to 1,550 m at Serra da Chapada das Perdizes, in Brazil (Lombardi et al., 2012Lombardi, V.T.; Santos, K.K.; D’Angelo-Neto, S.; Mazzoni, L.G.; Rennó, B.; Faetti, R.G.; Epifânio, A.D. & Miguel, M. 2012. Registros notáveis de aves para o sul do Estado de Minas Gerais, Brasil. Cotinga, 34: 32-45.).

The species has most of its range in Brazil, occurring in all major regions. Its presence in the states of Amazonas, Tocantins, Bahia, Santa Catarina and Rio Grande do Sul was confirmed only in the past two decades, showing the imperfect knowledge on its distribution. Regarding the last two states, Fontana et al. (2008Fontana, C.S.; Rovedder, C.E.; Repenning, M. & Gonçalves, M.L. 2008. Estado atual do conhecimento e conservação da avifauna dos Campos de Cima da Serra do sul do Brasil, Rio Grande do Sul e Santa Catarina. Revista Brasileira de Ornitologia, 16(4): 281-307.) point out that the lack of previous records is best considered a result of a lack of studies rather than a colonization event.

Among the new localities where the species has been found are savanna enclaves in the region of Humaitá and Canutama (MZUSP; Wikiaves, 2020WikiAves. 2020. WikiAves, a Enciclopédia das Aves do Brasil. Available: Available: http://www.WikiAves.com.br . Access: 01/06/2020.
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), in southern Amazonia (now being converted for soybean crops), and similar habitats at Campos Amazônicos National Park (961,317.77 ha) further east, where other grassland specialists have also been found (Cândido-Jr. & Dal’Maso, 2016Cândido-Jr., J.F. & Dal’Maso, A. 2016. Avifauna. In: Plano de Manejo Parque Nacional dos Campos Amazônicos. Brasília, ARPA. Pp. 204-209.). The Amazonian savannas are of great ecological interest and, in the case of the national park, may protect significant populations of the species.

The species has also been cited as occurring in Amapá state (BirdLife International, 2020bBirdLife International. 2020b. Species factsheet: Culicivora caudacuta. Available: Available: http://www.birdlife.org . Access: 19/06/2020.
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), another Amazonian region with extensive savannas, but the source of this record could not be traced and this locality was excluded here.

The current information shows that C. caudacuta has a much broader range than first thought, and besides at least 13 Brazilian states where it has been found, in Bolivia, in the departments of El Beni, La Paz and Santa Cruz; in Paraguay in the departments San Pedro, Presidente Hayes, Cordillera, Paraguarí, Concepción, Canindeyú, Caaguazú, Caazapá, Itapúa and Misiones; in Argentina in the provinces of Misiones, Chaco, Corrientes, Entre Ríos, Formosa and Santa Fe; and In Uruguay it is known from a single record from the department of Rivera.

Its presence is probable but still undocumented in southeast Peru, at Madre de Dios in the Pampas de Heath (12°53′S, 68°54′W), a seasonally flooded grassland on the Peru-Bolivia border making a northwestern extension of the Gran Chaco Boliviano-Paraguayo, where several grassland specialists such as Rhynchotus rufescens, Micropygia schomburgkii, Ammodramus humeralis, Sporophila plumbea and Coryphaspiza melanotis have been found (Graham et al., 1980Graham, G.L.; Graves, G.R.; Schulenberg, T.S. & O’Neill, J.P. 1980. Seventeen bird species new to Peru from the Pampas de Heath. The Auk, 97(3): 366-370.). A record of C. caudacuta from Pampa Moscoso was made just 13 km from Pampas de Heath, adding to the likelihood of the species occurring there.

Conservation

Culicivora caudacuta has long figured in threatened species lists: at global level it was considered as Threatened in 1988, Lower Risk/Near Threatened in 1994 and 2000, and Vulnerable from 2004 to the present (BirdLife International, 2020bBirdLife International. 2020b. Species factsheet: Culicivora caudacuta. Available: Available: http://www.birdlife.org . Access: 19/06/2020.
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).

Most countries in its range consider it to be threatened, as Argentina, where it was considered as Vulnerable (Fraga, 1996Fraga, R.M. 1996. Aves. Pp. 155-219. In: García Fernandez, J.J.; Fraga, R.M.; Díaz, G.B. & Baigún, R.J. (Eds.). Libro rojo de mamíferos y aves amenazados de la Argentina. Buenos Aires, Fundación para la Conservación de las Especies y Medio Ambiente.), and now as Threatened (Ministerio de Ambiente y Desarrollo Sustentable & Aves Argentina, 2017Ministerio de Ambiente y Desarrollo Sustentable & Aves Argentina. 2017. Categorización de las aves de la Argentina (2015). Buenos Aires, Ministerio de Ambiente y Desarrollo Sustentable & Aves Argentina.).

In Brazil, C. caudacuta is included in several state lists of threatened species, as in Minas Gerais (Lins, 1998Lins, L.V. 1998. Culicivora caudacuta. Pp. 337-338. In: Machado, A.B.M.; Fonseca, G.A.B.; Machado, R.B.; Aguiar, L.M.S. & Lins, L.V. (Eds.). Livro Vermelho das Espécies Ameaçadas de Extinção da fauna de Minas Gerais. Belo Horizonte, Fundação Biodiversitas.; COPAM, 2010Conselho Estadual de Política Ambiental (COPAM). 2010. Lista das espécies da fauna ameaçadas de extinção no estado de Minas Gerais. Deliberação Normativa COPAM Nº 147 de 30 de abril de 2010. Diário do Executivo - Minas Gerais, 04 de maio de 2010.), São Paulo (Bressan et al., 2009Bressan, P.M.; Kierulff, M.C.M. & Sugieda, A.M. (Eds.). 2009. Fauna ameaçada de extinção no estado de São Paulo: vertebrados. São Paulo, Secretaria do Meio Ambiente.), Paraná (Mikich & Bérnils, 2004Mikich, S.B. & Bérnils, R.S. (Eds.). 2004. Livro vermelho da fauna ameaçada de extinção no estado do Paraná. Curitiba, Instituto Ambiental do Paraná.), Santa Catarina (Occhialini, 2010Occhialini, D.S. (Ed.). 2010. Lista das espécies da fauna ameaçada de extinção em Santa Catarina. Relatório Técnico Final. Fundação do Meio Ambiente.) and Rio Grande do Sul (Fontana et al., 2003Fontana, C.S.; Bencke, G.A. & Reis, R.E. (Eds.). 2003. Livro vermelho da fauna ameaçada de extinção no Rio Grande do Sul. Porto Alegre, EDIPUCRS.).

On the other hand it has been dropped from the national list since 2014 (ICMBio, 2018Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio). 2018. Livro vermelho da fauna Brasileira ameaçada de extinção - Aves. Brasília, ICMBio. v. 3.), where it was previously listed as Vulnerable (Hass & Silva e Silva, 2008Hass, A. & Silva e Silva, R. 2008. Culicivora caudacuta . In: Machado, A.B.M.; Drummond, G.M. & Paglia, A.P. (Org.). Livro Vermelho das Espécies Ameaçadas de Extinção. Brasília, MMA & Fundação Biodiversitas. v. 2, p. 628-629.).

In Uruguay, where it is known from a single locality, it is considered as Vulnerable (Azpiroz et al., 2012Azpiroz, A.B.; Alfaro, M. & Jiménez, S. 2012. Lista roja de las aves del Uruguay. Una evaluación del estado de conservación de la avifauna nacional con base em los criterios de la Unión Internacional para la Conservación de la Naturaleza. Montevideo, Dirección de Medio Ambiente.), while in Bolivia it is not included in red lists (Balderrama, 2009Balderrama, J.A. 2009. Aves. Pp. 305-417. In: Aguirre, L.F. (Ed.). Libro rojo de la fauna silvestre de vertebrados de Bolivia. La Paz, Ministerio de Medio Ambiente.).

The main reason for C. caudacuta to be considered threatened is the widespread conversion of native grasslands all over South America into cultivation, mostly for soybeans, and pastures dominated by exotic grasses where the birds cannot persist (Codesido & Fraga, 2009Codesido, M. & Fraga, R.M. 2009. Distribution of threatened grassland passerines of Paraguay, Argentina and Uruguay, with new locality records and notes on their natural history and habitat. Ornitología Neotropical, 20(4): 585-595.; BirdLife International, 2020bBirdLife International. 2020b. Species factsheet: Culicivora caudacuta. Available: Available: http://www.birdlife.org . Access: 19/06/2020.
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).

In the study area the expansion of Eucalyptus monocultures for pulp and charcoal over the natural grasslands continues unchecked and is the main driver of habitat loss (Fig. 22), a trend also in southern Brazil, Uruguay and Argentina also (Modernel et al., 2016Modernel, P.; Rossing, W.A.H.; Corbeels, M.; Dogliotti, S.; Picasso, V. & Tittonell, P. 2016. Land use change and ecosystem service provision in the Pampas and Campos grasslands of southern South America. Environmental Research Letters, 11, 113002.).

Figure 22
Culicivora caudacuta habitat occupation by Eucalyptus sp. monoculture, Araxá, Ribeirão do Inferno (07 May 2014). Photo: RSS.

There are little data on population densities, and all come from Brazil. At Chapada dos Veadeiros National Park the estimated population is 1,475-4,022 individuals, with an average of 2,433 individuals, while at Brasília National Park there are estimated to be 440 to 2,374 individuals, with an average of 1,021 (Braz, 2008Braz, V.S. 2008. Ecologia e conservação das aves campestres do bioma cerrado. PhD Thesis, Departamento de Ecologia do Instituto de Ciências Biológicas, Universidade de Brasília, 178p.). Kanegae (2011Kanegae, M.F. 2011. Population size of threatened and endemic birds of the cerrado in Estação Ecológica de Itirapina, a fragmented area in the state of São Paulo, Brazil. Bird Conservation International, 22: 144-154.) found densities of 9.6 individuals/km² at Itirapina Ecological Station, with a total estimated population of 112 individuals.

Another population has recently been found in Tocantins state, in the Jalapão region including the Serra Geral do Tocantins Ecological Station and Jalapão State Park where a significant population seems to occur (Dornas & Crozariol, 2012Dornas, T. & Crozariol, M.A. 2012. Aves associadas a ambientes de veredas na Estação Ecológica Serra Geral do Tocantins com novos registros para a região e nota sobre a população local de Culicivora caudacuta. Atualidades Ornitológicas, 169: 54-65.; Dornas & Pascoal, 2019), especially along the wet grasslands bordering the Mauritia palm swamps veredas.

Culicivora caudacuta has been confirmed to occur at the following state-run protected areas in Brazil: Serra Geral do Tocantins Ecological Station, Jalapão State Park, Chapada Diamantina National Park, RESEX Recanto das Araras de Terra Ronca, Chapada dos Veadeiros National Park, Serra dos Pirineus State Park, Emas National Park, Águas Emendadas Ecological Station, APA Cafuringa, ARIE Cruls, Brasília National Park, Brasília National Forest, Brasília Botanical Garden, Guará Ecological Park, IBGE Ecological Reserve, Serra das Araras Ecological Station, Serra da Boa Esperança State Park, Serra do Cabral State Park, Tabuleiro Municipal Natural Park, Quedas do Rio Bonito Ecological Park, Serra do Intendente State Park, Sempre Vivas National Park, Mata do Limoeiro State Park, APA Morro da Pedreira, Serra do Rola Moça State Park, Serra do Cipó National Park, Serra da Canastra National Park, São Miguel Wildlife Sanctuary, Santa Bárbara do Rio Pardo Experimental Farm, Santa Bárbara Ecological Station, Itirapina Ecological Station, Botucatu State Forest, Itararé Unit of Research and Development, Vila Velha State Park, Guartelá State Park, APA Rio Iraí, APA Ibirapuitã, Ibirapuitã Biological Reserve, and the following private reserves (as recognised by Brazil’s environmental agencies): RPPN Fazenda Minnehaha, RPPN Lagoa do Formoso, RPPN Reserva Natural Serra do Tombador, RPPN SESC Pantanal e Entorno, RPPN Cisalpina, RPPN Fazenda da Barra, RPPN Aves Gerais, RPPN Mata do Jambreiro, RPPN Unidade de Conservação de Galheiros, RPPN Vale Encantado, and RPPN Foz do Rio Aguapeí.

In Bolivia it occurs at Madidi National Park, Barba Azul Nature Reserve, Beni Biological Station, and Noel Kempff Mercado National Park.

Protected areas in Paraguay where C. caudacuta has been recorded are Serranía San Luis National Park, Bosque Mbaracayú Natural Reserve, Kanguery Biological Station, and San Rafael National Park.

In Argentina it occurs in the following protected areas, many private: Puerto San Juan Private Reserve, Campo San Juan Reserve, Urutaú Reserve, Don Lorenzo Refuge, Santa Rosa Private Reserve, Iberá Provincial Reserve, Iberá National Park, Isla Apipé Grande Provincial Natural Reserve, Rincón de Santa María Natural Reserve, Mburucuyá National Park, El Bagual Reserve, Río Pilcomayo National Park, and Federico Wildermuth Provincial Reserve for Multiple Uses.

In Brazil it is likely that C. caudacuta will be found in additional protected areas since about half of the existing ones do not have a management plan with information on their avifauna while others have only preliminary data.

ACKNOWLEDGMENTS

Most information on the breeding biology of Culicivora caudacuta in Minas Gerais was gathered during bird surveys and monitoring in the properties of Fosfertil (currently Mosaic Fertilizantes), at Uberaba, Tapira, Patos de Minas, Patrocínio and Catalão, this last locality in Goiás. From 2001 to 2012 Fosfertil funded and supported my research and its publicising. My thanks to Paulo Eduardo Batista, communications sector, to the managers of each area, and those who assisted me in fieldwork: Romes do Nascimento, William Caixeta, Wilson Alves Barbosa Filho, Lucio dos Reis Oliveira, and especially, Caroline Favaro Oliveira Valera. Dr. Aristônio M. Teles (Universidade Federal de Goiás) identified the Asteraceae holding nests of Culicivora caudacuta at Tapira. Hans-Martin Berg (NMW), Marco Aurélio Crozariol (MNRJ), Hein van Grow (NHMUK), Eduardo Carrano (MHNCI), Markus Unsöld (ZSM), Luis Fábio Silveira (MZUSP), Maria de Fátima Cunha Lima and Lincoln Carneiro (MPEG), Leonardo Lima Bergamini (IBGE), Steven W. Cardiff (LSUMZ), Sylke Frahnert (ZMB), and Kathia Rivero (MNK), provided information on specimens held at their collections. Markus Unsöld, Hein van Grow, Hans-Martin Berg and Eduardo Carrano also kindly provided photographs of the specimens. Dante Buzzetti kindly supplied unpublished data on nests of Culicivora caudacuta. José Fernando Pacheco and Juvêncio Nunes da Costa shared information on localities. Fábio Olmos provided fundamental support for the English version of the manuscript and numerous suggestions and corrections, and Jeremy Minns revised the final English version of the manuscript.

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APPENDIX 1

Records of Sharp-tailed Tyrant Culicivora caudacuta. Record: O = observation, P = photography, S = specimen, V = vocalization recorded. Source: AMNH = American Museum of Natural History, New York, USA; ANSP = Academy of Natural Sciences, Philadelphia, USA; CM = Carnegie Museum of Natural History, Pittsburgh, USA; DZUFMG = Departamento de Zoologia da Universidade Federal de Minas Gerais, Belo Horizonte, Brazil; FMNH = Field Museum of Natural History, Chicago, USA; IBGE = Coleção Ornitológica da Reserva Ecológica do IBGE, Brasília, Brazil; LSUMZ = Louisiana State University, Museum of Natural Science, Baton Rouge, USA; MACN = Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina; MCNA = Museu de Ciências Naturais da Pontifícia Universidade Católica de Minas Gerais, Belo Horizonte, Brazil; MCP = Museu de Ciências da Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, Brazil; MCZ = Museum of Comparative Zoology, Cambridge, USA; MHNCI = Museu de História Natural Capão da Imbuia, Curitiba, Brazil; MHNSCP = Museo de Historia Natural de la Sociedad Científica del Paraguay, Asunción, Paraguay; MNK = Museo de Historia Natural Noel Kempff Mercado, Santa Cruz de la Sierra, Bolivia; MNRJ = Museu Nacional, Rio de Janeiro, Brazil; MPEG = Museu Paraense Emílio Goeldi, Belém, Brazil; MZUSP = Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil; NHMUK = Natural History Museum, Tring, UK; NMW = Naturhistorisches Museum Wien, Wien, Austria; RMNH = Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands; SMF = Senckenberg Naturmuseum Frankfurt, Germany; USNM = United States National Museum, Washington, USA; ZSM = Zoologische Staatssammlung München, Germany; EB = eBird database; ML = Macaulay Library database; WA = WikiAves database; XC = XenoCanto database. Personal observations: DB = Dante Buzzetti; FO = Fabio Olmos; JFP = José Fernando Pacheco; JNC = Juvêncio Nunes da Costa; RSS = Robson Silva e Silva.

Country, State/Province/Departament, Municipality, Locality Latitude Longitude Record Source BRAZIL Amazonas Canutama, Igarapé Assuã, BR 319 08º05'S 63º45'W S, P MZUSP, WA Humaitá 07º35'S 63º14'W O EB Maranhão Alto Parnaíba, Estiva 09º35'S 46º25'W S MPEG Tocantins Almas, RPPN Fazenda Minnehaha 11º08'S 47º08'W O (Dornas & Crozariol, 2012) Araguacema, Rio Piranha 08º48'S 49º25'W O JFP & FO Lizarda, 15 km East 09º27'S 46º35'W O (Pacheco & Olmos, 2010) Mateiros 10º22'S 46º08'W O (Dornas & Pascoal, 2019) Mateiros 10º23'S 46º03'W P WA, (Dornas & Pascoal, 2019) Mateiros 10º27'S 46º08'W O (Dornas & Pascoal, 2019) Mateiros, Jalapão State Park, lake near the dunes 10º34'S 46º40'W O RSS, (Pacheco & Olmos, 2010) Mateiros, Jalapão State Park, near Pousada do Jalapão 10º18'S 46º56'W O RSS & JFP Miracema do Tocantins 09º34'S 48º23'W P WA Ponte Alta do Tocantins 10º44'S 47º32'W P WA Rio da Conceição, Serra Geral do Tocantins Ecological Station 11º14'S 46º53'W P WA, (Dornas & Crozariol, 2012) Rio Sono 09º25'S 47º15'W P WA, (Dornas & Pascoal, 2019) Rio Sono, headwaters of Ribeirão Areias 09º48'S 47º34'W O (Dornas & Pascoal, 2019) Rio Sono, headwaters of Ribeirão Monte Santo 09º44'S 47º29'W P, V WA, XC, (Dornas & Pascoal, 2019) Rio Sono, Mansinha 09º20'S 47º16'W O (Dornas & Pascoal, 2019) Bahia Cocos 12º43'S 44º33'W P WA Correntina, Fazenda Jatobá 13º45'S 45º36'W O (Antas et al., 1993) Jaborandi, Pousada Trijunção, RPPN Lagoa do Formoso 14º47'S 45º56'W P WA Mucugê, Gerais of Chapada Diamantina 13º00'S 41º24'W P, V EB, WA, XC Goiás Água Fria de Goiás, Fazenda Nossa Senhora Aparecida 14º47'S 47º45'W P WA, EB Águas Lindas de Goiás, Lago do Descoberto 15º44'S 48º13'W O JNC Alto Paraíso de Goiás, Chapada dos Veadeiros National Park 14º05'S 47º40'W P, V (Reinert et al., 1998), (Braz, 2008), WA Alto Paraíso de Goiás, Fazenda dos Anões 14º19'S 47º30'W P EB Alto Paraíso de Goiás, Fazenda Chapadão das Almas, Rio das Almas 13º58'S 47º27'W P DB Alto Paraíso de Goiás, Fazenda Firmeza 14º17'S 47º29'W O EB Alto Paraíso de Goiás, Fazenda São Bento 14º06'S 47º28'W O EB Alto Paraíso de Goiás, Fazenda Volta da Serra 14º10'S 47º46'W O EB Alto Paraíso de Goiás, near Ribeirão São Miguel 14º09'S 47º47'W O EB Alto Paraíso de Goiás, Pontezinha 14º08'S 47º31'W P EB Alto Paraíso de Goiás, Road to Osho Lua 14º18'S 47º39'W O EB Alto Paraíso de Goiás, Road to Rio dos Couros 14º08'S 47º40'W O EB Alto Paraíso de Goiás, Templo Centelha Divina 14º08'S 47º34'W O EB Anápolis, Boa Vista, Chácara das Rosas 16º18'S 48º54'W O (Pereira & Silva, 2009) Anápolis, Trilha Ecológica do Tucano 16º17'S 48º56'W O (Pereira & Silva, 2009) Aparecida de Goiânia, Serra das Areias, Rio Dourados 16º51'S 49º18'W O JNC Campo Alegre de Goiás, Serra do Facão 17º39'S 47º43'W O JNC Catalão, headwaters of Ribeirão Batalha 17º32'S 47º27'W P RSS Cavalcante, Fazenda Piqui 13º29'S 47º33'W O EB Cavalcante, Road BR-10 13º56'S 47º27'W O EB Cavalcante, RPPN Reserva Natural Serra do Tombador 13º40'S 47º48'W P WA, (Antonelli-Filho, 2011) Chapadão do Céu 18º25'S 52º33'W P, V WA Chapadão do Céu, Emas National Park 18º17'S 52º46'W P (Rodrigues et al., 1999), DB, RSS Cocalzinho de Goiás 15º47'S 48º46'W P WA, JNC Cocalzinho de Goiás, headwaters of Rio Corumbá 15º51'S 48º45'W O JNC Cristalina 16º46'S 47º36'W P, V WA, XC Cristalina, Córrego Cristal 16º51'S 47º22'W O JNC Jataí 17º52'S 51º43'W P WA Mineiros, Emas National Park 18º08'S 52º54'W P (Rodrigues et al., 1999), RSS, WA, XC Ouro Verde 16º11'S 49º11'W O JNC Pirenópolis. Serra dos Pirineus State Park 15º48'S 48º53'W P WA, EB Planaltina 15º29'S 47º38'W S MNRJ Rio Quente 17º45'S 48º44'W O JNC São Domingos, RESEX Recanto das Araras 13º36'S 46º17'W O EB Taquaral de Goiás, Serra do Brandão 16º03'S 49º33'W O JNC Distrito Federal Brasília, Planaltina, Águas Emendadas Ecological Station 15º32'S 47º36'W O (Sousa & Marini, 2007) Brasília, Altiplano Leste 15º50'S 47º43'W P, V EB, XC, WA, RSS Brasília, ARIE Cruls 15º44'S 47º55'W O (Felizola & Azevedo, 2013) Brasília, Brazlândia and Sobradinho, APA de Cafuringa 15º32'S 48º04'W O (Bagno et al., 2006) Brasília, Cristo Redentor, RECOR 15º55'S 47º53'W S IBGE Brasília, Taguatinga, Brasília National Forest 15º45'S 48º04'W P, O EB, RSS Brasília, Lago Sul, Brasília Botanical Gardens 15º52'S 47º49'W O EB Brasília, Núcleo Bandeirante, Fazenda Água Limpa 15º57'S 47º56'W P, V EB, XC Brasília, Núcleo Bandeirante, Granja Modelo do Ipê 15º55'S 47º59'W O EB Brasília, Santa Maria 16º00'S 47º56'W O JNC Brasília, Brasília National Park 15º40'S 47º58'W S, O MNRJ, (Antas, 1995), EB Brasília, Guará, Guará Ecological Park 15º49'S 47º58'W S MNRJ Brasília, IBGE Ecological Reserve 15º55'S 47º52'W V (Braz & Cavalcanti, 2001), ML Mato Grosso Barão de Melgaço, RPPN SESC Pantanal e Entorno 16º35'S 56º15'W O (BirdLife International, 2020a) Chapada dos Guimarães, Chapada 15º25'S 55º48'W S AMNH, NHMUK Itiquira 17º17'S 53º49'W P, V WA Jaciara, Fazenda Santa Fé 15º59'S 55º02'W O (Petermann et al., 2001) Porto Estrela, Serra das Araras Ecological Station, Rio Salobo valley 15º39'S 57º13'W O (Silva & Oniki, 1988) Mato Grosso do Sul Bodoquena, Fazenda Indiana, Rio Salobra 20º25'S 56º39'W O (Pivatto et al., 2006) Bonito, RPPN Fazenda da Barra 21º06'S 56º13'W P WA, (Pellin et al., 2008) Brazilândia, RPPN Cisalpina 21º16'S 51º55'W O (Godoi et al., 2013), (Morante-Filho et al., 2014) Campo Grande, airport 20º28'S 54º40'W P EB, WA Costa Rica 18º18'S 54º54'W P WA Costa Rica, Emas National Park 18º18'S 52º57'W P, O WA, RSS Coxim, Fazenda Monte Verde 18º15'S 54º41'W S MZUSP, MCZ, (Pinto, 1944) Coxim, Fazenda Recreio 18º22'S 54º46'W S FMNH, (Pinto, 1944) Ribas do Rio Pardo 20º27'S 53º49'W P WA Terenos 20º25'S 54º53'W P, V EB, WA Minas Gerais Aiuruoca 21º47'S 44º42'W P EB, WA, (Peixoto, 2014) Andrelândia 21º44'S 44º11'W P XC, WA, (Peixoto, 2014) Antônio Carlos 21º18'S 43º45'W P WA Antônio Carlos, São Sebastião de Campolide 21º17'S 43º50'W P WA Araguari 18º39'S 48º11'W P WA Araxá, Horizonte Perdido 19º42'S 46º51'W O RSS Araxá, Ribeirão do Inferno 19º46'S 46º52'W O RSS Araxá, Serra da Bocaína 19º43'S 46º48'W P WA Barbacena, Galego 21º15'S 43º43'W P, V WA Belo Vale, Serra de Belo Vale 20º25'S 43º56'W V WA, EB Bias Fortes, Várzea de Santo Antônio 21º36'S 43º43'W P WA Boa Esperança, Serra da Boa Esperança State Park 20º58'S 45º40'W P EB, WA Bom Jardim de Minas, Fazenda Areão 21º52'S 44º07'W O (Pacheco et al., 2008), (Lopes et al., 2009) Brumadinho, Serra da Moeda 20º05'S 44º10'W P WA Buenópolis, Serra do Cabral State Park 17º51'S 44º15'W P WA, RSS Campos Altos 19º41'S 46º10'W P WA Capitólio 20º36'S 46º04'W V WA Carmo da Cachoeira 21º22'S 45º30'W P WA Carrancas 21º28'S 44º36'W P, V WA, (Peixoto, 2014) Carrancas 21º29'S 44º38'W O (Lombardi et al., 2012) Carrancas, Fazenda do Oswaldo 21º30'S 44º52'W P (Moura & Corrêa, 2012) Carrancas, Serra das Broas 21º36'S 44º36'W P (Moura & Corrêa, 2012), WA, RSS Carrancas, Serra de Carrancas 21º26'S 44º42'W P EB, XC, WA, RSS Conceição do Mato Dentro, Parque Natural Municipal do Tabuleiro 19º05'S 43º34'W O (IEF, 2016) Conceição do Mato Dentro, Parque Natural Municipal do Tabuleiro, headwaters of Ribeirão do Campo 19º04'S 43º37'W O (IEF, 2016) Conceição do Mato Dentro, Serra do Intendente State Park, hedwaters of Rio Cubas 19º07'S 43º33'W O (IEF, 2016) Conceição do Mato Dentro, Serra do Intendente State Park 18º58'S 43º39'W O (IEF, 2016) Congonhas 20º29'S 43º50'W O (Mazzoni et al., 2012) Coronel Xavier Chaves 21º03'S 44º09'W O EB Cruzeiro da Fortaleza 18º58'S 46º40'W P WA Cruzília 21º43'S 44º43'W V XC, (Peixoto, 2014) Curvelo 18º44'S 44º26'W P WA Curvelo, Fazenda Olhos D’Água 19º00'S 44º30'W O (Melo, 1997) Delfinópolis, Serra da Babilônia 20º21'S 46º34'W P WA, RSS Diamantina, Guinda 18º15'S 43º40'W P WA Diamantina, Sempre-Vivas National Park 17º48'S 43º46'W P WA Estrela do Sul, Fazenda Monte Carmelo, Ribeirão Piçarrão 18º48'S 47º53'W O (Lopes et al., 2009) Felixlândia 18º45'S 44º52'W P WA Francisco Dumont, Serra do Cabral, Vereda Córrego do Cachorro 17º23'S 44º15'W P WA Gouveia 18º34'S 43º54'W P, V WA Guapé, Serra da Rapadura/Serra dos Macacos 20º50'S 45º55'W P WA Ibertioga 21º24'S 43º56'W P WA, (Peixoto, 2014) Ibiá 19º29'S 46º32'W P WA Indianópolis 19º02'S 47º55'W P WA Indianópolis, Fazenda Monte Carmelo, Ribeirão Mandaguari 19º02'S 47º42'W O EB, (Lopes et al., 2009) Indianópolis, Fazenda Nova Monte Carmelo 18º55'S 47º40'W O (Ferreira et al., 2015) Itabira, Serra dos Alves 19º30'S 43º27'W P, V WA, EB Itabira, Mata do Limoeiro State Park 19º35'S 43º25'W P WA Itabirito 20º15'S 43º53'W P WA, EB Itajubá, near Frigorífico Frivasa 22º26'S 45º27'W P WA Itaverava 20º39'S 43º36'W P, V WA Itutinga 21º23'S 44º40'W O EB Jaboticatubas, Serra do Cipó National Park 19º30'S 43º36'W P WA Juiz de Fora, Paredão de Pedra, Condomínio do Sol 21º45'S 43º21'W P WA Juiz de Fora, Road of Torreões 21º49'S 43º28'W P WA Lavras, Quedas do Rio Bonito Ecological Park 21º19'S 44º58'W O EB Lavras, Serra do Carrapato 21º20'S 44º57'W P, V EB, WA Lima Duarte, Fazenda do Sr. G.C. Fonseca 21º43'S 44º00'W O (Lopes et al., 2009) Lima Duarte, São Sebastião do Monte Verde 21º55'S 43º50'W P WA Lima Duarte, Souza 21º48'S 44º03'W O (Lopes et al., 2009) Luminárias, Serra Grande 21º32'S 44º49'W P EB, XC, WA, (Peixoto, 2014) Madre de Deus de Minas 21º32'S 44º21'W P WA, (Peixoto, 2014) Medeiros 19º58'S 46º13'W P WA Minduri 21º40'S 44º36'W O (Lombardi et al., 2012) Minduri, Serra da Chapada das Perdizes 21º35'S 44º35'W O WA, (Lombardi et al., 2012) Morro do Pilar, APA Morro da Pedreira 19º17'S 42º39'W P WA Morro do Pilar, RPPN Aves Gerais, headwaters of Rio Picão 19º13'S 43º29'W P, V (Carrara & Faria, 2012) Morro do Pilar, Serra do Cipó National Park 19º15'S 43º31'W P, V WA, XC, (Rodrigues et al., 2011) Nepomuceno 21º22'S 45º14'W O EB Nova Lima, Estrada de Rio de Peixe 20º04'S 43º54'W P WA Nova Lima, Retiro das Pedras 20º06'S 43º59'W S, P, V DZUFMG, MCNA, WA Nova Lima, São Sebastião das Águas Claras 20º02'S 43º57'W P WA Nova Lima, Serra do Curral, RPPN Mata do Jambreiro 19º58'S 43º54'W P WA Nova Lima, Serra do Rola Moça State Park 20º03'S 44º00'W O (Hoffmann et al., 2007) Nova Ponte 19º09'S 47º41'W O EB Nova Ponte, Fazenda São Pedro 18º55'S 47º42'W O (SUPRAM, 2019) Olaria 21º52'S 43º55'W P WA Ouro Preto, Miguel Burnier 20º26'S 43º47'W P, V (Mazzoni et al., 2012) Paracatu, headwaters of Ribeirão Batalha 17º29'S 47º15'W P RSS Patos de Minas, Mosaic Fertilizantes 18º22'S 46º54'W P RSS Patrocínio, headwaters of Córrego Bebedouro, Mosaic Fertilizantes 19º00'S 46º46'W S, P RSS, MZUSP Patrocínio, headwaters of Córrego Capoeira Grande 19º11'S 46º55'W S, P, V RSS, XC, MZUSP Patrocínio, headwaters of Córrego do Fundão 19º05'S 46º53'W S, P MZUSP, RSS Patrocínio, headwaters of Córrego do Jacu 19º03'S 46º52'W P RSS Patrocínio, headwaters of Córrego do Mato 19º09'S 46º55'W P RSS Patrocínio, headwaters of Córrego Duas Pontes 19º09'S 46º56'W P RSS Patrocínio, near power line 1 19º04'S 46º52'W P RSS Patrocínio, Serra Chapadão de Ferro 18º55'S 46º49'W O (Mattos et al., 1991), RSS Perdizes 19º21'S 47º16'W P, V WA Perdizes, RPPN Unidade de Conservação de Galheiros 19º12'S 47º08'W O, V (Malacco et al., 2003), (Lopes et al., 2009) Piedade do Rio Grande 21º27'S 44º11'W P, V WA, XC Pimenta 20º28'S 45º48'W P WA Piranguinho 22º24'S 45º32'W P WA Piranguinho, Olegário Maciel 22º19'S 45º35'W O (Mattos et al., 1991) Piumhi, Serra de Piumhi 20º28'S 45º55'W P WA Pompéu 19º10'S 45º02'W P, V WA, (Souza et al., 2018) Pouso Alegre 22º07'S 45º51'W P, V WA, EB Prados, Pitangueiras 21º12'S 44º04'W P WA Prados, Ribeirão do Elvas 21º08'S 44º06'W P WA Quartel Geral, Quartel São João 19º15'S 45º47'W P, V (Moura et al., 2011) Resende Costa, Córrego da Cruz 20º54'S 44º15'W P, V WA, EB Sacramento, Serra da Canastra National Park 20º08'S 46º51'W P RSS, WA Santa Bárbara do Monte Verde 21º56'S 43º42'W P WA Santa Rita de Ibitipoca 21º33'S 43º55'W V XC Santa Rita do Sapucaí 21º12'S 45º42'W O EB Santa Vitória, São Simão Channel, Serra Negra 18º56'S 50º30'W O (Mattos et al., 1991) Santana do Garambéu 21º34'S 44º04'W P, V WA, XC Santana do Garambéu, Fazenda do Sr. Pp.S. Almeida 21º43'S 44º05'W O (Lopes et al., 2009) Santana do Riacho, Lapinha da Serra, Serra do Cipó 19º07'S 43º40'W P WA, EB Santana do Riacho, Cardeal Mota, Serra do Cipó 19º19'S 43º37'W P WA Santana dos Montes 20º47'S 43º41'W P WA, EB Santo Hilário 20º39'S 45º50'W O EB São Gonçalo do Abaeté, Fazenda São Francisco 4 and 10 18º18'S 45º48'W P, V WA São João Batista do Glória, Serra da Canastra National Park 20º32'S 46º24'W P WA, EB São João del Rei 21º15'S 44º20'W P, V WA, (Peixoto, 2014) São João del Rei 21º16'S 44º08'W O (Peixoto, 2014), RSS São João del Rei 21º16'S 44º04'W O (Peixoto, 2014) São João del Rei, Serra do Carvoeiro 21º09'S 44º11'W P EB São Roque de Minas, Serra da Canastra National Park 20º14'S 46º32'W P, V (Silveira, 1998), EB, ML, WA, XC, RSS São Thomé das Letras 21º43'S 44º58'W P WA São Tiago 20º54'S 44º30'W P WA São Vicente de Minas, Fazenda Bom Jardim 21º38'S 44º25'W P EB, WA Sete Lagoas, Serra de Santa Helena 19º26'S 44º16'W P WA Tapira, Mosaic Fertilizantes (BD-5) 19º49'S 46º50'W P, V RSS, XC Tapira, Mosaic Fertilizantes (Natural Reserve) 19º51'S 46º47'W P RSS Tapira, Ribeirão do Inferno 19º47'S 46º53'W P RSS Três Marias, Fazenda Araras 8 18º11'S 45º06'W P WA Três Marias, Fazenda Perdões 18º11'S 45º25'W O (Ribon et al., 1995) Uberaba, headwaters of Rio Uberabinha 19º21'S 47º54'W P WA, EB Uberaba, Fazenda Água Emendada 19º17'S 48º01'W O (Lopes et al., 2009) Uberaba, RPPN Vale Encantado 19º32'S 47º53'W P WA Unaí, Santuário da Vida Silvestre São Miguel, Fazenda São Miguel 15º50'S 46º30'W O (FUNATURA, 1994), (Lopes et al., 2008) Uruana de Minas, Cercado 16º17'S 46º15'W P WA Vargem Bonita 20º19'S 46º22'W P WA São Paulo Águas de Santa Bárbara, Fazenda Experimental Santa Bárbara do Rio Pardo 22º53'S 49º15'W O (Willis & Oniki, 2003) Águas de Santa Bárbara, Santa Bárbara Ecological Station 22º48'S 49º14'W O (Willis & Oniki, 2003) Batatais 20º53'S 47º37'W S MZUSP, (Pinto, 1944) Bofete 23º05'S 48º16'W P WA, EB Botucatu, Botucatu State Forest 22º56'S 48º27'W P, V WA, EB, XC Botucatu, Sítio São José 22º56'S 48º25'W P WA Broa, Rio Itaqueri 22º15'S 47º52'W O (Willis & Oniki, 2003) Castilho, RPPN Foz do Rio Aguapeí 21º06'S 51º44'W O (Miyaji, 2013) Franca 20º32'S 47º24'W S MZUSP, (Pinto, 1944) Iperó, Ipanema 23º26'S 47º36'W S SMF, NMW Itapetininga, Fazenda Campo Grande 23º38'S 47º58'W O (Willis & Oniki, 2003) Itararé 24º06'S 49º18'W S NMW, NHMUK, RMNH Itararé, Fazenda Santa Andrea 24º08'S 49º10'W O (Willis & Oniki, 2003) Itararé, Unidade de Pesquisa e Desenvolvimento de Itararé 24º16'S 49º13'W S MCP Itirapina, Itirapina Ecological Station 22º13'S 47º54'W P, V (Willis & Oniki, 2003), WA Ituverava, Rio das Pedras 20º11'S 47º56'W S NMW Santa Lúcia 21º41'S 48º04'W S ZMB São Carlos, Fazenda Santa Maria da Fábrica 22º10'S 47º55'W O (Willis & Oniki, 2003) Taquarivaí, Córrego Escaramuça (Scaramuza) 23º53'S 48º46'W S NMW Paraná Balsa Nova, São Luiz do Purunã 25º28'S 49º42'W P, V WA, EB, XC Campo Largo 25º27'S 49º29'W P WA Candói, Fazenda Rodeio Velho 25º38'S 52º07'W P, V WA, EB Castro 24º46'S 49º59'W P WA Curitiba 25º27'S 49º08'W S, V NMW, NHMUK, XC Jaguariaíva 24º15'S 49º42'W O, P, V (Santos, 2007), WA, EB Maringá, Campus do Centro Universitário de Maringá (CESUMAR) 23º26'S 51º55'W O (Esclarski et al., 2011) Palmeira, Fazenda Santa Rita 28º18'S 49º48'W O (Anjos & Graf, 1993) Piraí do Sul, Fazenda Cuiabá 24º24'S 50º02'W S, P MCP, WA Piraquara 25º26'S 49º04'W P, V WA Ponta Grossa, Vila Velha State Park 25º14'S 50º00'W O (Mikich & Bérnils, 2004) Porto Amazonas 25º32'S 49º53'W P WA Quatro Barras, APA Estadual do Rio Iraí 25º23'S 49º04'W S MHNCI, (Mikich & Bérnils, 2004) São José dos Pinhais, Banhados do Rio Miringuava 25º35'S 49º10'W P WA São José dos Pinhais, Várzea na estrada do Curralinho 25º33'S 49º03'W P EB Telêmaco Borba, Fazenda Monte Alegre 24º12'S 50º33'W O (Rocha et al., 2003) Tibagi, Guartelá State Park 24º40'S 50º13'W P, V EB, WA Santa Catarina Campo Belo do Sul 27º55'S 50º47'W P, V EB, WA Capão Alto, Fazenda Pai João 28º10'S 50º38'W P, V WA, RSS Lages, Coxilha Rica 28º17'S 50º17'W S, P, V MCP, (Repenning et al., 2010) Lages, Coxilha Rica 28º18'S 50º16'W S, P, V MCP, (Repenning et al., 2010) Lages, Fazenda Santa Maria, Coxilha Rica 28º15'S 50º18'W P, V (Repenning et al., 2010) Lages, Morrinhos, Coxilha Rica 28º16'S 50º17'W P WA Lages, Rio Lava-Tudo Valley, Coxilha Rica 28º18'S 50º14'W P WA Urupema, Morro do Combate 28º00'S 49º49'W O RSS Rio Grande do Sul Alegrete, Fazenda Cerro dos Porongos 30º04'S 55º31'W O EB Alegrete, Fazenda do Beto 29º54'S 55º49'W O (Fontana et al., 2003) Alegrete, Fazenda Sá Brito 29º58'S 55º45'W O EB Alegrete. Ibirapuitã APA 29º59'S 55º40'W O EB Alegrete, Ibirapuitã Biological Reserve 29º55'S 55º47'W V (Bencke, 2001) Alegrete, Serra do Caverá 30º23'S 55º24'W P WA Bom Jesus 28º42'S 50º24'W S, P, V MCP, XC, WA Bom Jesus, Arroio Água Branca 28º35'S 50º24'W S, P, V MCP, (Repenning et al., 2010) Bom Jesus, Banhado do Arroio Água Branca 28º36'S 50º23'W S, V MCP, EB Bom Jesus, Cachoeira dos Baggio 28º40'S 50º28'W P, V (Repenning et al., 2010) Bom Jesus, Fazenda da Ronda 28º28'S 50º42'W P, V (Repenning et al., 2010) Bom Jesus, northeast of Rio Santana 28º26'S 50º41'W P, V (Repenning et al., 2010) Bom Jesus, Road RS-110 28º35'S 50º22'W P, V (Repenning et al., 2010) Bom Jesus, Sanga José Luis 28º28'S 50º42'W P, V (Repenning et al., 2010) Bom Jesus, Várzea do Rio Santana 28º29'S 50º43'W P, V (Repenning et al., 2010) Campestre da Serra, Guacho 28º40'S 51º05'W P (Repenning et al., 2010), WA Cruz Alta 28º36'S 53º34'W P, V WA Manoel Viana, Assentamento Santa Maria do Ibicuí 29º29'S 55º37'W O EB Rosário do Sul 30º11'S 54º57'W P WA Rosário do Sul, Fazenda Schütz 30º18'S 54º51'W O EB Rosário do Sul, São Bento 30º17'S 54º46'W O EB Rosário do Sul, Vila Temp 30º14'S 54º51'W O EB Santa Margarida do Sul 30º21'S 54º04'W P WA Santa Maria 29º44'S 53º50'W P WA São Francisco de Assis 29º36'S 54º45'W O (Gressler & Krüger, 2005) São Francisco de Paula 29º09'S 50º24'W O EB São Gabriel 30º23'S 54º21'W P EB, WA São Gabriel, Suspiro 30º37'S 54º20'W P EB, WA São Gabriel, Horto Florestal Formosa (CMPC) 30º17'S 54º47'W P WA Tupanciretã, Porteira Encantada - Fazenda Moresco 28º57'S 53º46'W P WA Tupanciretã, old Road Tupã-Cruz Alta 28º56'S 53º45'W P, V EB, WA Vacaria, Arroio Pessegueiro 28º22'S 50º45'W P, V (Repenning et al., 2010) Vacaria, Banhado Rio Santana 28º28'S 50º42'W O EB Vacaria, Capão Alto 28º12'S 51º00'W P, V (Repenning et al., 2010) Vacaria, East of Rio Socorro 28º21'S 50º53'W P, V (Repenning et al., 2010) Vacaria, Estrada de Ferro 28º21'S 50º46'W P, V (Repenning et al., 2010) Vacaria, Estrada de Ferro, Fazenda Socorro 28º21'S 50º47'W P, V (Repenning et al., 2010) Vacaria, headwaters of Arroio Macena 28º30'S 50º47'W P, V (Repenning et al., 2010), WA Vacaria, Itacolomi 28º13'S 50º52'W P, V (Repenning et al., 2010) Vacaria, São Pedro, Capela do Caravaggio 28º08'S 50º54'W S, P, V MCP, (Repenning et al., 2010) Vacaria, Túneis da Estrada de Ferro 28º19'S 50º43'W P, V (Repenning et al., 2010) Vacaria, West of Rio Socorro 28º20'S 50º55'W P, V (Repenning et al., 2010) Vacaria, Várzea do Arroio Moema 28º30'S 50º48'W S MCP BOLIVIA La Paz 7 km East of Ixiamas 13º46'S 68º03'W O (Parker III et al., 1991) Franz Tamayo, Apolo, Madidi Savanna, Madidi National Park 14º43'S 68º21'W V (Soria-Auza & Hennessey, 2005), ML Ixiamas 13º45'S 68º05'W O EB Pampa Moscoso 13º02'S 68º50'W O EB El Beni 10 km Southwest of San Borja 14º53'S 66º51'W O (Parker III et al., 1991) 26 and 30 km East of San Borja 14º49'S 66º51'W V (Parker III et al., 1991), ML Barba Azul Nature Reserve 13º45'S 66º05'W P EB Beni Biosphere Station 14º38'S 66º17'W O EB Cerro San Simón 13º36'S 62º15'W O (Parker III & Rocha, 1991) Estancia El Porvenir, Beni Biological Station 14º50'S 66º17'W O (Brace et al., 1997) Estancia Motacuzal 13º45'S 64º37'W O EB Estancia Peñas Verdes 13º36'S 64º30'W O EB General Jose Ballivan, 3 km Southwest of San Borja 14º51'S 66º50'W S LSUMZ, (Schmitt & Schmitt, 1987) Las Palmiras 13º44'S 66º24'W O EB Llanos de Mojos, 7 Islas Camp 13º48'S 64º30'W O EB North of Trinidad 13º37'S 64º58'W O (Soria-Auza & Hennessey, 2005) Puerto Ustarez 12º44'S 64º38'W P EB Selva Blue, Laguna Larga 12º49'S 65º46'W O EB Upper Yata/Tapado 13º16'S 66º02'W O EB Santa Cruz Buena Vista 17º27'S 63º40'W S ANSP, CM Santa Rosa de la Roca 15º54'S 61º24'W P EB Serranía de Huanchaca I, Noel Kempff Mercado National Park 13º57'S 60º49'W S LSUMZ, (Bates & Parker III, 1998) Serranía de Huanchaca II, Noel Kempff Mercado National Park 14º31'S 60º44'W S LSUMZ, MNK, (Bates & Parker III, 1998), EB PARAGUAY San Pedro Establecimiento Laguna Blanca, Retiro Malvina 23º49'S 56º18'W O (Barnett et al., 2004) Estancia Villa Josefina 23º55'S 56º45'W O EB Forestadora Río Verde 23º46'S 56º20'W P EB Laguna Blanca 23º46'S 56º17'W P (Centrón, 2009a), EB North Yboty 23º43'S 56º18'W O EB Nueva Germania 23º54'S 56º34'W S ZSM, (Laubmann, 1940) Señorita 23º45'S 56º13'W O EB Presidente Hayes Benjamín Aceval, Villa Hayes 25º06'S 57º34'W S MHNSCP, (Bertoni, 1930) Estancia La Rafaela 24º53'S 57º27'W O EB Monte Sociedad 25º03'S 57º35'W S (Laubmann, 1940) Pirizal area, Catholic Mission 22º13'S 58º25'W O EB Ruta Transchaco, km 20 to 79 24º48'S 57º46'W O EB Ruta Transchaco, km 60 24º55'S 57º38'W O (Hayes, 1995) Cordillera Arroyos y Esteros, km 100 24º52'S 56º53'W O EB Estancia Sombrero 25º01'S 56º35'W O EB Paraguarí Agromonte 25º43'S 57º06'W O EB Estancia Barrerito 26º16'S 57º03'W O (Centrón, 2009c), EB Sapucái 25º40'S 56º55'W S NHMUK, (Chubb, 1910) Concepción Arroyo Tagatiya, Campos Cerrados 22º44'S 57º33'W O EB Cerrados de Concepción 22º25'S 57º10'W O EB Estancia San Luis de La Sierra, Arroyo La Paz 22º23'S 57º27'W O EB Serranía San Luis National Park 22º36'S 57º25'W O EB Zanja Moroti, Río Apa 22º30'S 57º00'W S ZSM, (Laubmann, 1940) Canindeyú Aguara Ñú 24º11'S 55º16'W P EB, XC Reserva Natural del Bosque Mbaracayú 24º06'S 55º14'W O (Madroño Nieto & Esquivel, 1997), EB Caaguazú Morombi 24º37'S 55º22'W O EB Upper Iguazú River 25º05'S 55º45'W S AMNH Caazapá Arrozal Codas 26º29'S 56º18'W O EB Estancia Roa Coé 26º33'S 56º06'W O (Codesido & Fraga, 2009) Estancia Tapyta 26º14'S 55º46'W O EB, XC San Juan Nepomuceno 26º08'S 55º56'W O EB Itapúa Carmen del Paraná 27º15'S 56º08'W O (Codesido & Fraga, 2009) Coronel Bogado, Fundación San Rafael 27º07'S 56º22'W O EB Dr. Blas Garay 26º46'S 56º16'W O EB Estero San José 26º55'S 56º04'W O (Bonzi et al., 2020) General Artigas, Estero Ñu Guasu 26º56'S 56º18'W O (Bonzi et al., 2020), EB Isla Yacyretá Natural Reserve 27º25'S 56º43'W O (Codesido & Fraga, 2009), (Bonzi et al., 2020), EB La Paz Fish Farm 27º00'S 55º54'W P EB Leandro Oviedo 26º43'S 56º16'W O EB San Miguel Potrero 27º03'S 56º08'W O (Bonzi et al., 2020) San Rafael, Guyra Reta Reserve, Kanguery Biological Station 26º30'S 55º46'W O (del Castillo & Centrón, 2010), EB San Rafael National Park, Estancia Kanguery 26º25'S 55º48'W O (Esquivel-M. et al., 2007), (Smith, 2017), EB Misiones Campo Llano 26º47'S 57º25'W O (Centrón & del Castillo, 2011), EB Estancia La Graciela 26º31'S 56º52'W O (Centrón, 2009b), (Ferreira, 2009), EB Santiago 27º05'S 56º50'W O EB Yabebyry 27º13'S 56º56'W O EB ARGENTINA Misiones Campo Prates, Barra Concepción 28º07'S 55º35'W O (Krauczuk, 2005) Candelaria, Estancia Santa Cecília 27º27'S 55º42'W P EB Candelaria, Puerto San Juan Private Reserve 27º21'S 55º37'W O (Krauczuk, 2005) Candelaria, Santa Ana, Campo San Juan Reserve 27º24'S 55º37'W P (Krauczuk, 1997), (Di Giacomo et al., 2007), EB Candelaria, Urutaú Reserve 27º29'S 55º46'W P EB Capital, Garupá, near Barrio Santa Helena 27º28'S 55º53'W O (Krauczuk, 2005, 2006) Capital, Posadas, airport 27º23'S 55º57'W O EB Capital, Posadas, Campus of Universidad Nacional de Misiones 27º26'S 55º53'W O (Krauczuk, 2005) Capital, Posadas, Don Lorenzo Refuge 27º25'S 55º52'W O (Krauczuk, 2005, 2006) Capital, Posadas, international bridge Posadas-Encarnación, El Zaimám creek confluence 27º22'S 55º53'W O (Krauczuk, 2006) Capital, Posadas, Itaembé Guazú 27º25'S 55º58'W P EB Capital, Posadas, mouth of Mártires river 27º23'S 55º57'W P EB Capital, Posadas, near the Ruínas de Mártires 27º25'S 55º56'W O (Krauczuk, 2005) Capital, Posadas, Nemesio Parma, Paraíso Costero 27º20'S 56º01'W P EB Capital, Posadas, Santa Rosa Private Reserve 27º24'S 55º53'W O (Krauczuk, 2005) Capital, Posadas, west access to Posadas 27º22'S 55º57'W O (Krauczuk, 2006) Concepción, Cerro Mártires y Barra Santa María 27º51'S 55º26'W O (Di Giacomo et al., 2007) Concepción de la Sierra, Barra Concepción 28º08'S 55º53'W S MACN, (Di Giacomo et al., 2007) Garupá creek basin 27º29'S 55º44'W O (Di Giacomo et al., 2007) Near Río Saimá, Itapua 27º22'S 55º58'W S (White, 1882) Chaco Bermejo, El Cachapé 26º50'S 59º00'W O (Di Giacomo et al., 2007) Bermejo, Sol de Mayo 26º59'S 58º42'W P EB Corrientes Capital, Estero Valenzuela 27º53'S 58º34'W O (Di Giacomo et al., 2007) Concepción and San Roque, Concepcíon - Chavarría 28º37'S 58º10'W O (Di Giacomo et al., 2007) Estancia Puerto Valle 27º43'S 56º29'W O (Parera, 2004) General Alvear, Bañado San Isidoro 28º54'S 56º36'W O EB General Paz, Lomas de Vallejos, Ruta Provincial 5 27º43'S 58º01'W O EB Gobernador General Virasoro, Estancia La Higuera 27º59'S 56º18'W O (Marino et al., 2013) Gobernador General Virasoro, Estancia Virocay 28º15'S 55º57'W O (Marino et al., 2013), EB Gobernador General Virasoro, Las Marías 28º06'S 56º03'W O (Krauczuk, 2005), EB Gobernador General Virasoro, Sosa Cué 28º00'S 56º02'W O EB Iberá Provincial Reserve 28º24'S 57º07'W O (Chebez et al., 1999) Ituzaingó, Estancia La Guayna 27º45'S 56º04'W O (Codesido & Fraga, 2009) Ituzaingó, Iberá National Park, Cambyretá 27º49'S 56º50'W O EB Ituzaingó, Isla Apipé Grande Natural Reserve 27º30'S 56º52'W O (Esteban, 1953) Ituzaingó, Lomada de San Alonso, Esteros del Iberá 28º14'S 57º24'W O EB Ituzaingó, Puerto Valle 27º43'S 56º30'W O EB Ituzaingó, Rincón de Santa María Natural Reserve 27º30'S 56º35'W P (Krauczuk, 2005), (Di Giacomo et al., 2007), XC, EB Ituzaingó, San Carlos, Rio Aguapey, Estancia San Joaquín 27º45'S 55º54'W S MACN, (Darrieu & Camperi, 1992) Isla Apipé Grande 27º30'S 56º52'W O (Esteban, 1953) Mburucuyá, Mburucuyá National Park 28º00'S 58º05'W O (Chebez et al., 1999), (Di Giacomo et al., 2007), EB Mburucuyá, Mburucuyá village and surroundings 28º02'S 58º13'W O EB Mercedes and San Martín, Rincón del Socorro and Iberá 28º32'S 57º10'W O (Di Giacomo et al., 2007), EB San Martín, Colonia Carlos Pellegrini 28º32'S 57º10'W O EB San Miguel, Estancia San Juan Poriahú 27º42'S 57º11'W O EB Santo Tomé, Estancia Mora Cué and around 28º18'S 56º10'W O (Di Giacomo et al., 2007) Santo Tomé, far northeast Corrientes 28º27'S 55º47'W O (Di Giacomo et al., 2007) Santo Tomé, Río Aguapey basin 28º36'S 56º56'W O (Di Giacomo et al., 2007) Santo Tomé, Ruta Provincial 40 28º22'S 56º07'W O EB Santo Tomé, Ruta Provincial 41 28º11'S 56º43'W O EB Entre Ríos Colón, Arroyo Caraballo 32º13'S 58º08'W O (Milat et al., 1985) Ibicuy, Ceibas 33º26'S 58º45'W O (Di Giacomo et al., 2007) Formosa Laishi, El Bagual Reserve 26º10'S 58º56'W P (Di Giacomo, 1996, 2005), EB Riacho Pilaga 26º05'S 57º59'W S USNM, (Wetmore, 1926) Río Pilcomayo National Park 25º04'S 58º07'W S (Chebez et al., 1999) Santa Fe Gral. Obligado y Vera, wooded wedge of Santa Fe 28º30'S 59º30'W O (Di Giacomo et al., 2007) Mocovi 28º24'S 59º42'W S AMNH, (Hartert & Venturi, 1909) San Martín, Provincial Reserve for Multiple Uses Federico Wildermuth 32º00'S 61º42'W O (Di Giacomo et al., 2007) Tacuarendi 28º25'S 59º18'W O (Lillo, 1909) Villa Ocampo 28º28'S 59º22'W S AMNH, (Hartert & Venturi, 1909) URUGUAY Rivera Near La Palma, Valle del Lunarejo 31º10'S 55º55'W P (Azpiroz, 1998)

APPENDIX 2

Specimens of Culicivora caudacuta.

BRAZIL

Amazonas:

MZUSP 92411, ♂, Igarapé Assuã, BR 319, Canutama, 07.xii.2011, Fábio Schunck & Bret Whitney.

MZUSP 92412, ♂, Igarapé Assuã, BR 319, Canutama, 07.xii.2011, Fábio Schunck & Bret Whitney.

MZUSP 92413, ?, Igarapé Assuã, BR 319, Canutama, 07.xii.2011, Fábio Schunck & Bret Whitney.

Maranhão:

MPEG 43519, ♂, Estiva, Alto Parnaíba, 03.iv.1989, Brígida, Rosemiro, J.M. Rosa, Raimundo & Dionísio.

Distrito Federal:

MNRJ 28988, ♀, Parque Guará, Brasília, 11.x.1963, Luiz Moojen.

MNRJ 28989, ♂, Parque Guará, Brasília, 11.x.1963, Luiz Moojen.

MNRJ 28990, ♂, Parque Guará, Brasília, 11.x.1963, Luiz Moojen.

MNRJ 28991, ♂, Parque Guará, Brasília, 11.x.1963, Luiz Moojen.

MNRJ 32662, ♂, Parque Nacional de Brasília, Brasília, 02.xii.1978, Helmut Sick.

MNRJ 32663, ♀, Parque Nacional de Brasília, Brasília, 02.xii.1978, Helmut Sick.

IBGE 719, ♂, Cristo Redentor, RECOR, Brasília, 04.vii.1984.

IBGE 720, ♀, Cristo Redentor, RECOR, Brasília, 04.vii.1984.

Goiás:

MNRJ 13218, ♂, Planaltina, 15.vi.1927, Emilie Snethlage.

MNRJ 13219, ♀, Planaltina, 15.vi.1927, Emilie Snethlage.

Mato Grosso:

AMNH 33317, ♀, Chapada, 13.iv.1883, Herbert Smith.

NHMUK 1889.1.10.299, ♀, Chapada, 17.x.1883, Herbert Smith.

Mato Grosso do Sul:

MZUSP 13211, ♀, Fazenda Monte Verde, Coxim, 29.vi.1930, João Leonardo Lima.

MZUSP 17391, ♀, Fazenda Monte Verde, Coxim, 08.viii.1937, José Leonardo de Lima.

MCZ 154615, ♀, Coxim, 29.vi.1930, João Leonardo Lima.

MCZ 154616, ♂, Coxim, 10.vii.1930, João Leonardo Lima.

MCZ 154617, ♀, Fazenda Monte Verde, Coxim, 10.vii.1930, João Leonardo Lima.

FMNH 17390, ♂, Fazenda Recreio, Coxim, 09.viii.1937, José Leonardo de Lima.

Minas Gerais:

DZUFMG 4511, ♂, Retiro das Pedras, Nova Lima, 02.vi.2005, Marcelo Ferreira de Vasconcelos, Diego Hoffmann, Leonardo Esteves Lopes & Éverton Vieira Ouriques.

DZUFMG 4512, ♀, Retiro das Pedras, Nova Lima, 02.vi.2005, Marcelo Ferreira de Vasconcelos, Diego Hoffmann, Leonardo Esteves Lopes & Éverton Vieira Ouriques.

MCNA 2211, ♂, Retiro das Pedras, Nova Lima, 02.vi.2005, Marcelo Ferreira de Vasconcelos, Diego Hoffmann, Leonardo Esteves Lopes & Éverton Vieira Ouriques.

MCNA 2277, ♀, Retiro das Pedras, Nova Lima, 02.vi.2005, Marcelo Ferreira de Vasconcelos, Diego Hoffmann, Leonardo Esteves Lopes & Éverton Vieira Ouriques.

MZUSP 115128, headwaters of Córrego do Fundão, Patrocínio, 22.x.2015, Luís Fábio Silveira & Robson Silva e Silva.

MZUSP 115129, Fazenda Todos os Santos, headwaters of Córrego Capoeira Grande, Patrocínio, 19.viii.2020, Robson Silva e Silva & Lucio dos Reis Oliveira.

MZUSP 115130, headwaters of Córrego Bebedouro, Patrocínio, 30.v.2021, Robson Silva e Silva, Paulo César Araújo dos Santos-Jr. & Lucio dos Reis Oliveira.

MZUSP 115131, headwaters of Córrego Bebedouro, Patrocínio, 30.v.2021, Robson Silva e Silva, Paulo César Araújo dos Santos-Jr. & Lucio dos Reis Oliveira.

MZUSP 115132, headwaters of Córrego Bebedouro, Patrocínio, 30.v.2021, Robson Silva e Silva, Paulo César Araújo dos Santos-Jr. & Lucio dos Reis Oliveira.

MZUSP 115133, headwaters of Córrego Bebedouro, Patrocínio, 30.v.2021, Robson Silva e Silva, Paulo César Araújo dos Santos-Jr. & Lucio dos Reis Oliveira.

São Paulo:

NMW 17817, ♂, Scaramuza, Itararé, 20.viii.1820, Johann Natterer.

NMW 17818, ♂, Scaramuza, Itararé, 20.viii.1820, Johann Natterer.

NMW 17820, ♀, Ipanema, 29.v.1819, Johann Natterer.

NMW 17821, ♂, Ipanema, 29.v.1819, Johann Natterer.

SMF 43938, ?, Ipanema, 29.v.1819, Johann Natterer.

NMW 17822, ♂, Itararé, 11.ii.1821, Johann Natterer.

NHMUK 1888.1.13.331, ♂, Itararé, 11.ii.1821, Johann Natterer.

RMNH 88812, ♂, Itararé, Johann Natterer.

NMW 17823, ♂, Rio das Pedras, 16.iv.1823, Johann Natterer.

MZUSP 1434, ♀, Batatais, 12.xii.1900, João Leonardo Lima.

MZUSP 7995, ♂, Franca, ix.1910, Ernest Garbe.

MCP 3269, Unidade de Pesquisa e Desenvolvimento de Itararé, Itararé, 26.i.2010, Márcio Repenning.

ZMB 2559, ♂, Santa Lúcia, between 1814 and 1831, Friedrich Sellow.

Paraná:

NMW 17819, ♂, Curitiba, 29.x.1820, Johann Natterer.

NHMUK 1888.1.13.330, ♂, 20.xi.1820, Curitiba, Johann Natterer.

MHNCI 5023, ♂, Quatro Barras, 07.iv.1999, Eduardo Carrano & Luiz Fernando Franco de Macedo.

MHNCI 5140, ?, Rio Iraí, Quatro Barras, 13.iv.1999, Eduardo Carrano & Luiz Fernando Franco de Macedo.

MCP 3583, Piraí do Sul, 29.i.2013, Tony Andrey Bichinski Teixeira.

MCP 3584, Piraí do Sul, 29.i.2013, Tony Andrey Bichinski Teixeira.

Santa Catarina:

MCP 2773, Coxilha Rica, Lages, 03.xii.2009, Márcio Repenning.

MCP 2793, Coxilha Rica, Lages, 12.i.2010, Mariana Lopes Gonçalves.

MCP 3294, Coxilha Rica, Lages, 01.ii.2009, Ismael Franz.

MCP 3689, Coxilha Rica, Lages, 10.ii.2010, Ismael Franz.

Rio Grande do Sul:

MCP 1845, Várzea do Arroio Moema, Vacaria, 22.xii.2006, Márcio Repenning.

MCP 2110, Banhado do Arroio Água Branca, Bom Jesus, 12.xii.2007, Cristiano Eidt Rovedder.

MCP 2111, Banhado do Arroio Água Branca, Bom Jesus, 24.xii.2007, Cristiano Eidt Rovedder.

MCP 2112, Banhado do Arroio Água Branca, Bom Jesus, 09.i.2008, Cristiano Eidt Rovedder.

MCP 2246, Banhado do Arroio Água Branca, Bom Jesus, 04.iv.2008, Márcio Repenning.

MCP 2265, Banhado do Arroio Água Branca, Bom Jesus, 03.xii.2007, Márcio Repenning.

MCP 2266, Banhado do Arroio Água Branca, Bom Jesus, 07.i.2008, Cristiano Eidt Rovedder.

MCP 3274, Bom Jesus, 08.i.2010, Cristiano Eidt Rovedder.

MCP 3275, Bom Jesus, 08.i.2010, Cristiano Eidt Rovedder.

MCP 3657, Coxilha Grande São Pedro, Vacaria, 16.i.2009, Márcio Repenning.

MCP 3683, Banhado do Arroio Água Branca, Bom Jesus, 14.xii.2009, Cristiano Eidt Rovedder.

MCP 3684, Banhado do Arroio Água Branca, Bom Jesus, 17.xi.2009, Cristiano Eidt Rovedder.

Unknown State

NHMUK 1888.1.1.433, ?, S.E. Brazil, P.L. Sclater.

ZMB 2560, ♂, between 1814 and 1831, Friedrich Sellow.

ZMB 2561, ♂, between 1814 and 1831, Friedrich Sellow.

ZSM, S. Brazil, ex. Mus. H. von Leuchtenberg.

BOLIVIA

El Beni:

LSUMZ 124493, ♂, General Jose Ballivan, 3 km SW San Borja, 06.x.1984, C. Gregory Schmitt.

Santa Cruz:

ANSP 143068, ♂, Buena Vista, 26.vii.1912, Joseph Steinbach.

CM P43912, ♀, Buena Vista, 25.vii.1911, Joseph Steinbach.

CM P43925, ♂, Buena Vista, 26.i.1912, Joseph Steinbach.

CM P51046, ♂, Buena Vista, 20.xi.1914, Joseph Steinbach.

CM P79126, ♂, Buena Vista, 16.iii.1917, Joseph Steinbach.

LSUMZ 150981, ♀, Velasco, Serranía de Huanchaca, 45 km E. Florida, 02.x.1989, Tristan J. Davis.

LSUMZ 150982, ♀, Velasco, Serranía de Huanchaca, 45 km E. Florida, 03.x.1989, John M. Bates.

LSUMZ 150983, ♀, Velasco, Serranía de Huanchaca, 45 km E. Florida, 03.x.1989, Gary H. Rosenberg.

LSUMZ 150984, ♀, Velasco, Serranía de Huanchaca, 45 km E. Florida, 04.x.1989, Abel Castillo.

LSUMZ 150985, ♂, Velasco, Serranía de Huanchaca, 45 km E. Florida, 06.x.1989, Tristan J. Davis.

LSUMZ 150986, ♂, Velasco, Serranía de Huanchaca, 45 km E. Florida, 12.x.1989, Tristan J. Davis.

LSUMZ 150987, ♂, Velasco, Serranía de Huanchaca, 45 km E. Florida, 12.x.1989, Gary H. Rosenberg.

LSUMZ 151790, ♀, Velasco, Serranía de Huanchaca, 45 km E. Florida, 03.x.1989, Abel Castillo (skeleton specimen).

LSUMZ 151884, ?, Velasco, Serranía de Huanchaca, 45 km E. Florida, 03.x.1989, Tristan J. Davis (alcoholic specimen).

MNK 701, ♀, Velasco, Serranía Huanchaca, 45 km E. Florida, 30.ix.1989, Abel Castillo.

PARAGUAY

Caaguazú:

AMNH 320548, ♂, Upper Iguazú River, 07.i.1931, Emil Kaempfer.

AMNH 320549, ♀, Upper Iguazú River, 07.i.1931, Emil Kaempfer.

San Pedro:

ZSM 32699, ♂, Col. Nueva Germânia, 01.ii.1932, E. Schunmacher.

Presidente Hayes:

MHNSCP, Benjamin Aceval, Villa Hayes.

? Monte Sociedad.

Paraguarí:

NHMUK 1905.10.12.364, ♂, Sapucái, 24.iii.1903, William Foster.

NHMUK 1905.10.12.365, ♂, Sapucái, 24.vii.1904, William Foster.

NHMUK 1905.10.12.366, ♂, Sapucái, 27.vii.1904, William Foster.

NHMUK 1905.10.12.367, ♂, Sapucái, 16.viii.1904, William Foster.

NHMUK 1905.10.12.368, ♀, Sapucái, 16.viii.1904, William Foster.

Concepción:

ZSM, Zanja Moroti, Río Apa.

ARGENTINA

Misiones:

MACN 44769, ♂, Concepción de la Sierra, Barra Concepción.

MACN 44771, ♂, Concepción de la Sierra, Barra Concepción.

Corrientes:

MACN 44770, ♀, Ituzaingó, San Carlos, Río Aguapey, Estancia San Joaquín.

MACN 44772, ♀, Ituzaingó, San Carlos, Río Aguapey, Estancia San Joaquín.

MACN 44773, ♂, Ituzaingó, San Carlos, Río Aguapey, Estancia San Joaquín.

Formosa:

USNM 227339, ♀, Kilometro 182, Riacho Pilaga, 10 mi NW, 14.viii.1920, Alexander Wetmore.

USNM 284426, ♀, Kilometro 182, Riacho Pilaga, 10 mi NW, 14.viii.1920, Alexander Wetmore.

? Río Pilcomayo National Park.

Santa Fe:

AMNH 498990, ♂, Mocovi, 02.xii.1903, Santiago Venturi.

AMNH 498991, ♀, Mocovi, 02.xii.1903, Santiago Venturi.

AMNH 435689, ♂, Villa Ocampo, 07.i.1904, Santiago Venturi.

AMNH 498989, ♂, Villa Ocampo, 01.xi.1905, Santiago Venturi.

Luís Fábio Silveira

Publication Dates

  • Publication in this collection
    17 Sept 2021
  • Date of issue
    2021

History

  • Received
    20 Apr 2021
  • Accepted
    17 June 2021
  • Published
    30 July 2021
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