A new area of endemism for Amazonian birds in the Rio Negro Basin.
|Abstract:||We describe a new area of endemism for Amazonian birds which we designate as the Jau Area of Endemism. This area of endemism in central-western Amazonia north of the Rio Solimoes was identified through congruent distributions of six avian taxa: Psophia crepitans ochroptera Pelzeln 1857, Nonnula amaurocephala Chapman 1921, Pteroglossus azara azara Vieilot 1819, Picumnus lafresnayi pusillus Pinto 1936, Synallaxis rutilans confinis Zimmer 1935, and Myrmoborus myotherinus ardesiacus Todd 1927. The southern and eastern limits of this area of endemism are the middle courses of the Solimoes and Negro rivers, respectively. The northern limits apparently coincide with sandy soil vegetation along the middle Rio Negro. The western boundary remains undefined, but could involve the Japura or Ica rivers north of the upper Solimoes. Taxonomic studies and expansion of ornithological collections are needed to more precisely delimit the Jau Area of Endemism. It is possible the avian taxa restricted to the Jag Area of Endemism are derived through parapatric or peripatric speciation events from taxa whose ranges were centered in the Imeri and Napo areas of endemism. Alternatively, tectonic events that affect the lower course of the Rio Negro could influence bird distribution in this region if they serve as vicariance mechanisms.|
Birds (Environmental aspects)
Borges, Sergio H.
Da Silva, Jose M.C.
|Publication:||Name: The Wilson Journal of Ornithology Publisher: Wilson Ornithological Society Audience: Academic Format: Magazine/Journal Subject: Biological sciences Copyright: COPYRIGHT 2012 Wilson Ornithological Society ISSN: 1559-4491|
|Issue:||Date: March, 2012 Source Volume: 124 Source Issue: 1|
|Geographic:||Geographic Scope: Brazil Geographic Code: 3BRAZ Brazil|
An area of endemism (AOE) is a "geographical region comprising
the distributions of two or more monophyletic taxa that exhibit a
phylogenetic and distributional congruence and having their respective
relatives occurring in other such-defined regions" (Harold and Mooi
1994:261). Areas of endemism are important for at least two reasons.
First, they represent the smallest geographical units for postulating
hypotheses about the history of their biotas (Cracraft 1988, 1994;
Morrone 1994; Morrone and Crisci 1995). Second, these areas are
considered priorities for establishment of conservation action because
they contain unique biotas (Fjeldsa 1993, Stattersfield et al. 1998).
There are modern approaches to identifying AOEs including parsimony analyses of endemicity (Morrone 1994, Morrone and Crisci 1995, Silva et al. 2004) and optimality criterion (Szumik et al. 2002, Szumik and Goloboff 2004). Traditionally, however, these areas have been identified through mapping the congruent geographical distribution of taxa based on extensive area of sympatry but not necessarily complete overlap of distributions (Platnick 1991).
Eight AOEs have been recognized for birds in the Amazonian lowlands (Fig. 1; Haffer 1974, 1978; Cracraft 1985; Silva et al. 2002). Most areas have boundaries coinciding with major rivers of the Amazon Basin (Wallace 1852, Haffer 1978, Cracraft 1985, Ayres and CluttonBrock 1992). We identified a number of avian taxa during a study of bird species distribution whose ranges apparently are restricted to the lower course of the Rio Negro (Borges 2004a, Borges 2007). Mapping the distributions of these taxa resulted in identification of an AOE not recognized in previous biogeographic analyses (Haffer 1978, Cracraft 1985). Our objective is to describe the new area of endemism based on congruent distributions of six avian taxa.
Study Area.--The study region encompasses the fight margin of the lower Rio Negro (Fig. 2). Most specimens analyzed were collected in Jau National Park (JNP), one of the largest (2,272,000 ha) protected areas in Brazil. The avifauna of JNP has been studied for the last 15 years including assessment of species diversity and general ecological requirements for most bird species (Borges and Carvalhaes 2000; Borges et al. 2001; Borges 2004a, b; Borges and Almeida 2011).
[FIGURE 1 OMITTED]
The new area of endemism is at proximities of two other areas of endemism, and we reviewed the geographical distribution of species considered endemic to the Napo and Imeri areas of endemism as originally proposed by Haffer (1978) and Cracraft (1985).
The taxonomic and geographical distributions of 383 bird species and subspecies were assessed in the study region. The geographic distribution of one species, Chestnut-headed Nunlet (Nonnula amaurocephala), and five subspecies, Greywinged Trumpeter (Psophia crepitans ochroptera), Ivory-billed Aracari (Pteroglossus azara azara), Lafresnayes's Piculet (Picumnus lafresnayi pusillus), Ruddy Spinetail (Synallaxis rutilans confinis), and Black-faced Antbird (Myrmoborus myotherinus ardesiacus) are restricted to the west of the lower Rio Negro and north of the Rio Solimoes. These taxa are considered endemic to the study region.
[FIGURE 2 OMITTED]
Grey-winged Trumpeter (Psophia crepitans ochroptera Pelzeln 1857). Pelzeln described this trumpeter as a distinct species, Psophia ochroptera. Ochroptera is treated as a subspecies of P. leucoptera in recent taxonomic literature (Pinto 1978, Sherman 1996). We agree with Haffer (1974) who proposed that ochroptera is a subspecies of P. crepitans, the trumpeter species distributed north of the Amazon River. P. crepitans has two other subspecies in addition to ochroptera: crepitans inhabiting the Guianan AOE and napensis, inhabiting the Napo and Imeri AOEs. P. c. ochroptera has been recorded mostly in white-sand woodland and terra firme forest in JNP (Borges et al. 2001, Borges and Almeida 2011), where an adult female was collected (INPA # 576). A recent molecular systematic study (Ribas et al. 2011), in addition to morphological assessment (Haffer 1974), found that ochroptera is diagnosable from the other subspecies in the Psophia crepitans complex.
Chestnut-headed Nunlet (Nonnula amaurocephala Chapman 1921). This rare puffbird was known only from specimens collected in Mana capuru used in the original description until it was rediscovered in JNP (Whittaker et al. 1995). It is associated with seasonally flooded blackwater forest or Igapo forest (Whittaker et al. 1995, Borges et al. 2001). The MZUSP has skins collected in Manacapuru (MZUSP 16561) and Codajas (MZUSP 16387), and one specimen was collected in JNP (MPEG 55855) in addition to specimens mentioned in Whittaker et al. (1995). The northernmost record of N. amaurocephala is the Unini River (Whittaker et al. 1995) and the easternmost is the Amana River (M. Cohn-Haft, pers. comm.) with no confirmed records from the upper portion of the Rio Negro (Haffer and Fitzpatrick 1985). It is suggested that N. amaurocephala forms a superspecies with N. ruficapilla and N. frontalis (Rassmussen and Collar 2002)
Ivory-billed Aracari (Pteroglossus azara azara Vieilot 1819). This subspecies of the Ivory-billed Aracari has the upper mandible brownish colored and is recorded from Castanheiro, Codajas, Igarape Cacau Pereira (Haffer 1974), Manacapuru (MZUSP 16833 and 16834), Maraa (MPEG 42505-42507), and JNP (Borges et al. 2001, Borges 2007). It is replaced by P. a. flavirostris in the Napo and Imeri AOEs and by P. a. mariae south of the Solimoes (Haffer 1974). There is some evidence of hybridization between mariae and azara near the mouth of the Purus River (Haffer 1974: 222-223). This Aracari has been recorded in both terra firme and flooded forests in JNP (Borges et al. 2001, Borges and Almeida 2011).
Lafresnaye's Piculet (Picumnus lafresnayi pusillus Pinto 1936). P. pusillus was described from Codajas on the left margin of the Rio Solimoes by Pinto (1936). This taxon was later reassigned as a subspecies of P. aurifrons (Peters 1948). Recent taxonomy considered pusillus as a subspecies of Picumnus lafresnayi (Winkler et al. 1995, Winkler and Christie 2002). P. lafresnayi forms a superspecies with P. pumilus that is apparently restricted to southern Venezuela and northwestern Brazil (Winkler and Christie 2002). There are six MZUSP specimens identified as P. l. pusilus collected in the Manacapuru region (MZUSP 16614-16619). The plumage of a single specimen collected in JNP and deposited in the INPA Bird Collection matched better with P. l. pusillus than with P. pumilus. The geographical distribution of pusillus appears to be restricted to the region west of the lower Rio Negro being replaced by P. pumilus in the Imeri AOE (Fig. 2). P. l. pusillus has been found mainly in flooded forests and secondary growth in JNP (Borges et al. 2001, Borges and Almeida 2011).
Ruddy Spinetail (Synallaxis rutilans confinis Zimmer 1935). This subspecies was described from specimens collected in Igarape Cacau Pereira in the lower portion of Rio Negro. It is replaced by S. r. dissors on the left margin of Rio Negro, which has been found in Manaus and in the upper Rio Negro at Sao Gabriel da Cachoeira and along the Casiquiare channel (Friedmann 1948, Pinto 1978). A better delimitation of the distribution of S. r. confinis will require specimen collections south of the Negro and Uaupes rivers. S. r. confinis is replaced in Napo AOE (eastern Ecuador and northeastern Peru) by S. r. caquetensis and in Inambari AOE (upper Amazonia south of the Solimoes River) by S. r. amazonica (Peters 1951, Pinto 1978).
Black-faced Antbird (Myrmoborus myotherinus ardesiacus Todd 1927). Haffer and Fitzpatrick (1985) evaluated the plumage variation in Myr moborus myotherinus and concluded that ardesiacus was a diagnosable taxon This subspecies was described from Manacapuru and has been recorded in Igarape Cacau Pereira, Santa Maria, Codajas (Zimmer 1932, Pinto 1978), Maraa (MPEG 42654-42672), and Jati National Park (MPEG 50614-50620), and is replaced in Imeri and Napo by M. m. elegans and in Inambari by M. m. myotherinus (Haffer and Fitzpatrick 1985). This antbird has been recorded in mature upland forest, white-sand campinaranas, and secondary growth forest in JNP (Borges et al. 2001, Borges and Almeida 2011).
Another seven avian taxa also are apparently restricted to central-western Amazonia: Amazona autumnalis diadema, Brachygalbula lugubris phaeonota, Nonnula rubecula simulatrix, Hylexestastes stresemanni stresemanni, Sclerurus rufigularis brunnescens, Myrmoborus lugubris stictopterus, and Hylophylax naevius obscurus. These subspecies, however, are known only from a few specimens and their taxonomic status and distribution needs additional evaluation.
Napo and Imeri Areas of Endemism.--The Imeri and Napo areas of endemism were supported by 21 and 56 avian taxa as originally described (Haffer 1978, Cracraft 1985). An updated distributional and taxonomic assessment decreased these numbers to six species (or subspecies) endemic to Imeri and 42 endemic to the Napo AOE (Appendix).
Several bird species previously thought to be restricted to the upper reaches of the Rio Negro have had their geographical distribution extended southward and to northern Peru (Borges et al. 2001, Alonso and Whitney 2003, Borges and Almeida 2011). Other Imeri birds are restricted to the northwestern portion of the Amazon Basin with no confirmed records in the lower reach of the Rio Negro or northern Peru. Most species restricted to Imeri AOE are specialists in white sand campinas (Cyanocorax helprini and Myrmeciza pelzelni), montane and pre-montane forests (Percnostola caurensis), and flooded forest (Thripophaga cherriei) (Zimmer 1999; Hilty 2003; SHB, unpubl, data).
Similarly, some birds previously thought to be endemic to the Napo region had their distributions extended east to the Rio Negro, as in the case of Myrmotherula ignota and Pteroglossus pluricinctus (Borges and Almeida 2011). Some species are even more widespread than previously described, but there are species endemic to northwestern (Imeri endemics), upper north Amazon (Napo endemics), and the central-western portion of the basin (this study).
We describe a previously undesignated area of endemism in central-western Amazonia that is supported by the congruent distribution of six avian taxa. Haffer and Fitzpatrick (1985) noted that a small number of strongly differentiated bird populations were restricted to this region but did not recognize it formally as an AOE due the reduced number of avian taxa (J. Haffer, pers. comm.). Conceptually and operationally, however, an AOE could be recognized with just two species (Platinick 1991, Harold and Mooi 1994).
Amorin and Pires (1999) also identified a small AOE bounded by the Rio Solimoes and Rio Negro without offering information about which species supported this biogeographic component (Amorim and Pires 1999: fig. 27). We suggest naming this new area of endemism as Jau in recognition of the importance of the National Park to biodiversity conservation in the Amazon. This area appears nominated as Rio Negro Area of Endemism in previous publications (Borges 2007, Ribas et al. 2011). However, we believe that Jail Area of Endemism is more appropriate to the geographical setting of the area (Fig. 2).
The precise geographical limits of this area are only partially identifiable. The southern and eastern limits probably coincide with middle portions of the Solimoes and Negro, large rivers known to isolate bird populations (Sick 1967, Haffer 1992). The northern boundary is more complicated, but from the middle Rio Negro northward the landscapes dominated by terra firme forests are extensively replaced by forests and fields growing in sandy soils called 'campinaranas' and 'campinas' (Anderson 1981, IBGE 1997). This more open vegetation occupies thousands of square kilometers between the middle and the upper Rio Negro. It is possible this discontinuity in vegetation serves as a barrier (or filter) to dispersal for some bird species, especially those that occur more frequently in terra firme forest on clay soils. The western boundary of Jau AOE also is difficult to identify due to the scarcity of bird collections from the region between Maraa to the Brazilian-Colombian border. We tentatively suggest the courses of the Japura or Ica rivers as the western boundary of Jail AOE, although additional fieldwork will be necessary (Fig. 2). We also note this region of faunal turnover could be not coincident with any fiver course as happens in the southern portion of the Amazon Basin (Haffer 1992).
One fundamental concern in recognizing and delimiting the Jau AOE is the current taxonomy of the birds considered endemics. The use of subspecies in biogeographic analyses is problematic because an unknown number of bird taxa described in the ornithological literature are not discrete evolutionary, units (Cracraft 1985, Haffer 1987). Some authors have been successful in using subspecies for biogeographic analysis (Bates et al. 1998, Borges 2007). Analyzing polytypical species as a single entity tends to over-estimate the geographical distributions of taxa and potentially ignores relevant units for conservation and biogeography (Bates and Demos 2001). We considered only taxa with accentuated morphological differentiation. However, the taxonomic status of Amazonian bird species, including those discussed here, needs to be continuously evaluated and their phylogenetic relationships clearly established. These taxonomic and biogeographic studies will require additional ornithological collections in the northern portion of the upper Rio Solitudes and in the middle to upper Rio Negro, principally along the fight margin of the latter, as these regions are likely contact zones.
The phylogenetic relationships of the species and subspecies endemic to the Jau AOE are not adequately known. However, three taxa (Pteroglossus azara azara, Picumnus lafresnayi pusillus, and Myrmoborus myotherinus ardesiacus) appear to have their closest relatives in the Imeri and Napo AOEs. This is supported by Borges (2007), who applied quantitative methods (parsimony analysis of endemicity and cluster analyses) to analyze the biogeographic relationships between avifaunas from different Amazonian AOEs. The avifauna of the JNP shares more species and subspecies with localities in the Imeri and Napo AOEs than with localities in the Inambari and Guiana AOEs (Borges 2007). This suggests the barrier effect of the Negro and Solimoes rivers is stronger than the barrier effect in the western border of the Jau AOE. A working hypothesis to the evolutionary history of the six taxa restricted to the Jau AOE is that they were derived through parapatric or peripatric speciation from ancient species whose ranges were once centered in the Imeri and Napo AOEs. Alternatively, tectonic events that affect the lower course of the Rio Negro (Forsberg et al. 2000, Almeida-Filho and Miranda 2007, Silva et al. 2007) could influence bird distribution in this region if they serve as vicariance mechanisms (Ribas et al. 2011).
We call attention to the urgent need to review the currently recognized AOEs for Amazonian birds. The last review of these areas was compiled over 20 years ago (Cracraft 1985), and taxonomic and geographical distribution data have continued to accumulate. The areas of endemism for Amazonian birds should be re-evaluated through modem biogeographic approaches (Morrone and Crisci 1995, Szumik and Goloboff 2004) facilitated by continuous progress increasing the data base of geographical distribution and systematics of neotropical birds.
SHB acknowledges Capes, WWF-Brazil, and Fundacao Vitoria Amazonica for financial and technical support. We thank Brazil's federal agency of environmental protection (IBAMA) for permission to work in Jail National Park. Luis Fabio Silveira, Maria Luiza, Alexandre Aleixo, and Mario Cohn-Haft kindly permitted us to examine bird collections under their care. Marcelo Moreira kindly prepared the maps. Morton Isler, Charles Zartman, and Camila Ribas helped in first versions of the paper. We appreciate the improvements in English usage by Phil Stouffer through the Association of Field Ornithologists' program of editorial assistance. We are also grateful to J. M. Bates and Jurgen Haffer for careful reviews of this paper and useful suggestions. We dedicate this paper to the memory of Dr. Jurgen Haffer for his inestimable contribution to the biogeography of neotropical birds.
Received 27 June 2007. Accepted 15 July 2011.
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SERGIO H. BORGES (1,3,4) AND JOSE M.C. DA SILVA (2)
(1) Departamento de Zoologia, Museu Paraense Emilio Goeldi, Belem, Para Brazil.
(2) Conservation International do Brazil, Avenue Nazare, 541/sala 310, 66035-170, Belem, Para, Brazil.
(3) Current address: Fundacao Vitoria Amazonica, Rua R/ S, casa 07, Quadra Q, Morada do Sol, 69060-080, Manaus, AM, Brazil.
(4) Corresponding author; e-mail: firstname.lastname@example.org
APPENDIX. Distribution of avian taxa in Imeri and Napo areas of endemism as originally proposed by Haffer (1978, marked with H) and Cracran (1985, marked with C. Bird taxonomy was updated following Gill and Donsker (2011). Species marked as E are endemics. Original name Current name Imeri Napo Jau (1) Crypturellus Crypturellus casiguiare--H casiquiare (2) x x C. duidae--C C. duidaes (3) x x x Aramides Aramides calopterus--H calopterus E Mitu salvini--H & C Mitu salvini E M. tomentosa--H M. tomentosum x x Neomorphus Neomorphus p. pucheranii--C pucheranii (4) x x Campylopterus Campylopterus villaviscensio--C villaviscensio E Leucippus Leucippus chlorocercus--H & C chlorocercus E Heliodoxa Heliodoxa gularis--H & C gularis E Topaza pyra--H & C Topaza p. x x x pyra (5) Phlogophilus Phlogophilus hemileucurus hemileucurus E Pyrrhura Pyrrhura albipectus--H & C albipectus E Galbula tombacea--H Galbula E tombacea tombacea (6) G. leucogastra G. chalco- chalcothorax thorax (7) E G. pastazae--H G. pastazae E Galbalcyrhynchus Galbalcyrhyn- leucotis--H chus leucotis E Nonnula ruficapilla Nonnula rufipectus--C ruficapilla E rufipectus N. brunnea--H N. brunnea E Notharchus ordii--H Notharchus x x x ordii (8) Pteroglossus f. Pteroglossus flavirostris--H azara x flavirostris P. pluricinctus--H P. pluricinctus x x x Selenidera r. Selenidera reinwardtii--H & C r. reinwardtii E S. nattereri--H & C S. nattereri x x Picumnus Picumnus pumillus--H & C pumillus E Celeus Celeus spectabilis--H & C spectabilis E spectabilis Hylexetastes Hylexetastes stresemanni stresemanni insignis--C insignis E Synallaxis Synallaxis moesta--H & C moesta E Thripophaga Thripophaga cherriei--H & C cherriei E Neoctantes niger--H Neoctantes x x niger (7) Myrmotherula Myrmotherula sunensis--H & C sunensis sunensi E M. obscura--H M. ignota obscura (11) x x x M. longicauda M. longicauda soderstromi--C soderstromi E M. cherriei--H & C M. cherriei (12) x x x M. ambigua--H & C M. ambigua x x Herpsilochmus sticturus Herpsilochmus dugandt--H & C dugandi E H. dorsimaculatus--H H. dorsima- culatus (13) Hypocnemis Hypocnetnis h. hypoxantha--C hypoxanthuta (14) x x x Pithys castanea--C Pithys castaneus (15) E Rhegmatorhina m. Rhegmatorhina m. melanosticta--C melanosticta E R. cristata--C R. cristata x x Myrmeciza Myrmeciza melanoceps--H melanoceps (16) E M. pelzelni--H & C M. pelzelni E M. disjuncta--H & C M. disjuncta (17) x x Gymnopithys Gymnopithys leucaspis leucaspis castanea--C castaneus E G. leucaspis G. leucaspis lateralis--C lateralis x x Thamnophius Thatnnophius praecox--H & C praecox E Percnostola Schistocichla caurensis--H caurensis (18) E Grallaria Grallaria dignissima--H & C dignissima E G. fulviventris Hylopezus f. --H & C fulviventris E Pipra coronata Lepidotrhix coronata--H coronata coronata E P. coronata L. coronata caquetae--C caquetae E P. pipra Dixiphia discolor--C pipra discolor E Heterocercus Heterocercus aurantiivertex--H & C aurantiivertex E H. flavivertex--H & C H. flavivertex (19) x x Attila Attila citri- citriniventris--H & C niventris (20) x x x A. torridus--C A. torridus E Myiophobus Myiophobus cryptoxanthus--H & C cryptoxanthus E Ramphotrigon Ramphotrigon fuscicauda--H fuscicauda (21) E Todirostrum c. Poecilotriccus calopterum--H & C calopterus E T. c. capitale--H P. capitalis (22) E Contopus n. Contopus n. nigrescens--C nigrescens E Microbates Microbates cinereiventris--H cinereiventris hormotus (23) E Cyphorinus aradus Cyphorinus salvini--C arada salvini E Microcerculus Microcerculus bambla albigularis--C bambla albigularis (24) E Cacicus sclateri--H & C Cacicus sclateri E Ocyalus Ocyalus latirostris--C latirostris E Hylophilus Hylophilus hypoxanthus hypoxanthus fuscicapillus--C fuscicapillus E H. b. brunneiceps--C H. brunneiceps (25) x x Cyanocorax Cyanocorax heilprini--H & C heilprini E Dolospingus Dolospingus fringilloides--H & C fringilloides (26) x x Taxonomic and distributional notes. (1) References for species records in the Jau Area of Endemism were Borges et al. (2001) and Borges and Almeida, (2011). (2) Recently recorded in northern Peru (Alonso and Whitney 2003). (3) Recently recorded in northern Peru (Alonso and Whitney 2003). (4) Subspecies N. p. lepidophanes occurs south of Rio Solimoes (Payne 1997). Haffer (1997) reported N. p. pucheroni in the upper Rio Negro (Imeri). (5) Recorded in southern Venezuela (Hilly 2003). (6) Subspecies G. t. menralis recorded south of Rio Solimoes (Inambari) (Tobias et al. 2002). (7) Also recorded in the Jurua River (Inambari) (Tobias et al. 2002). (8) Recently recorded in Inambari AGE (Guilherme and Borges 2011). (9) Records from Inambari, Imeri, and Tapajos AOEs (Zimmer and Isler 2003). (10) Subspecies M. s. yessupi recorded in Juruti River (Inambari) (Zimmer and Isler 2003). (11) Also recorded in Inambari AIDE (Zimmer and Isler 2003). (12) Recently recorded in northern Peru (Alonso and Whitney 2003). (13) Also recorded in Guiana AOE (Cohn-Haft et al. 1997, Zimmer and Isler 2003, Naka et al. 2006). (14) Also recorded in Inambari AOE (Zimmer and Isler 2003). (15) Recently rediscovered in field (Lane et al. 2006). (16) Also recorded in Inambari AGE (Zimmer and Isler 2003). (17) Recently recorded in Guiana AOE (Naka et al. 2006). (18) Recently rediscovered in field (Zimmer 1999). (19) Also recorded in Guiana AOE (Snow 2004, Naka et al. 2006). (20) Recently recorded in Inambart AIDE (Guilherme and Borges 2011). (21) Considered monotypic with several records south of the Amazonas River with an isolated population in the Napo region (Fitzpatrick 2004). (22) Considered monotypic with a population recorded in Rondonia AGE (Fitzpatrick 2004). (23) Distribution and taxonomy follow Restall et al. (2006). (24) Distribution and taxonomy follow Restall (2006). (25) Considered as monotypic by Ridgely and Tudor (1989). (26) Recently recorded in Guiana AGE (Robbins et al. 2005). The following taxa were not considered in the analysis. (1) Topaza pella pamprepta: considered as a synonym with T. p. smaragdula (Hu et al. 2000); (2) Taphrospilu.s hypostictus hypostictus: considered monotypic with widespread distribution through the Andes (Schuchmann 1999); (3) Pharomachrus pavoninus pavoninus: widespread distribution through the Amazon; (4) Nonnula amaurocephala: erroneously assigned as endemic to the upper Rio Negro by Cracraft (1985); (5) Phlegopsis barringeri: hybrid between Phlegopsis erythroptera and P. nigromaculata (Zimmer and Isler 2003); (6) Pipra filicauda: widespread distribution through the Amazon; (7) Mionectes oleagineus hauxwelli: race M. o. hauxwelit inseparable from nominate (Fitzpatrick 2004).
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