A New Area of Endemism for Amazonian in the Rio Negro Basin Author(s) :Sérgio H. Borges and José M. C. Da Silva Source: The Wilson Journal of Ornithology, 124(1):15-23. 2012. Published By: The Wilson Ornithological Society DOI: http://dx.doi.org/10.1676/07-103.1 URL: http://www.bioone.org/doi/full/10.1676/07-103.1

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A NEW AREA OF ENDEMISM FOR AMAZONIAN BIRDS IN THE RIO NEGRO BASIN

SE´ RGIO H. BORGES1,3,4 AND JOSE´ M. C. DA SILVA2

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 Solimo˜es 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, 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 Solimo˜es 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´orIc¸a´ rivers north of the upper Solimo˜es. 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 Jau´ 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 distribution in this region if they serve as vicariance mechanisms. Received 27 June 2007. Accepted 15 July 2011.

An area of endemism (AOE) is a ‘‘geographical 1978; Cracraft 1985; Silva et al. 2002). Most region comprising the distributions of two or more areas have boundaries coinciding with major monophyletic taxa that exhibit a phylogenetic and rivers of the Amazon Basin (Wallace 1852, distributional congruence and having their respec- Haffer 1978, Cracraft 1985, Ayres and Clutton- tive relatives occurring in other such-defined Brock 1992). We identified a number of avian regions’’ (Harold and Mooi 1994:261). Areas of taxa during a study of bird species distribution endemism are important for at least two reasons. whose ranges apparently are restricted to the First, they represent the smallest geographical lower course of the Rio Negro (Borges 2004a, units for postulating hypotheses about the history Borges 2007). Mapping the distributions of these of their biotas (Cracraft 1988, 1994; Morrone taxa resulted in identification of an AOE not 1994; Morrone and Crisci 1995). Second, these recognized in previous biogeographic anal- areas are considered priorities for establishment of yses (Haffer 1978, Cracraft 1985). Our objec- conservation action because they contain unique tive is to describe the new area of endemism biotas (Fjeldsa˚ 1993, Stattersfield et al. 1998). based on congruent distributions of six avian There are modern approaches to identifying taxa. AOEs including parsimony analyses of endemic- ity (Morrone 1994, Morrone and Crisci 1995, METHODS Silva et al. 2004) and optimality criterion (Szumik Study Area.—The study region encompasses the et al. 2002, Szumik and Goloboff 2004). Tradi- right margin of the lower Rio Negro (Fig. 2). Most tionally, however, these areas have been identified specimens analyzed were collected in Jau´ National through mapping the congruent geographical Park (JNP), one of the largest (2,272,000 ha) distribution of taxa based on extensive area of protected areas in . The avifauna of JNP has sympatry but not necessarily complete overlap of been studied for the last 15 years including distributions (Platnick 1991). assessment of species diversity and general eco- Eight AOEs have been recognized for birds in logical requirements for most bird species (Borges the Amazonian lowlands (Fig. 1; Haffer 1974, and Carvalhaes 2000; Borges et al. 2001; Borges 2004a, b; Borges and Almeida 2011). Biogeographic Analysis.—We compiled a list 1 Departamento de Zoologia, Museu Paraense Emı´lio of bird species and subspecies recorded in the Goeldi, Bele´m, Para´, Brazil. study area (Borges et al. 2001, Borges 2004a, 2 Conservation International do Brazil, Avenue Nazare´, Borges 2007, Borges and Almeida 2011). The 541/sala 310, 66035-170, Bele´m, Para´, Brazil. taxonomic status and distribution of birds re- 3 Current address: Fundac¸a˜o Vito´ria Amazoˆnica, Rua R/ S, casa 07, Quadra Q, Morada do Sol, 69060-080, Manaus, corded in the study region were evaluated AM, Brazil. through specimens housed at Museu Paraense 4 Corresponding author; e-mail: [email protected] Emı´lio Goeldi (MPEG), Museu de Zoologia da 15 16 THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 124, No. 1, March 2012

FIG. 1. Areas of endemism currently recognized for birds in the Amazonia lowlands based on Haffer (1974, 1978), Cracraft (1985), and Silva et al. (2002).

Universidade de Sa˜o Paulo (MZUSP), Bird Col- endemic to the Napo and Imeri areas of endemism lection of Instituto Nacional de Pesquisas da as originally proposed by Haffer (1978) and Amazoˆnia (INPA), Field Museum of Natural Cracraft (1985). History (FMNH), and American Museum of Natural History (AMNH). We also compiled RESULTS information on specimens collected in the Negro/ The taxonomic and geographical distributions of Japura´ interfluvium (e.g., Maraa˜, Barcelos, Mana- 383 bird species and subspecies were assessed in capuru) deposited in MPEG. All relevant informa- the study region. The geographic distribution of tion available from the taxonomic literature was one species, Chestnut-headed Nunlet (Nonnula also used (Pinto 1944, 1978; Sherman 1996; amaurocephala), and five subspecies, Grey- Winkler and Christie 2002; Borges 2004a). We winged Trumpeter (Psophia crepitans ochroptera), did not attempt a complete and detailed taxonomic Ivory-billed Aracari (Pteroglossus azara azara), revision of the bird taxa, but checked if the species Lafresnayes’s Piculet (Picumnus lafresnayi pusil- or subspecies were morphologically distinct from lus), Ruddy Spinetail (Synallaxis rutilans confinis), their closest relatives and if their ranges were and Black-faced Antbird (Myrmoborus myother- restricted to the study region. inus ardesiacus) are restricted to the west of the The new area of endemism is at proximities of lower Rio Negro and north of the Rio Solimo˜es. two other areas of endemism, and we reviewed the These taxa are considered endemic to the study geographical distribution of species considered region. Borges and Silva N A NEW AREA OF ENDEMISM FOR AMAZONIAN BIRDS 17

FIG. 2. Northwestern Amazonia, showing the study region and municipalities cited in the text. The light gray area indicates the suggested limits of the Jau´ Area of Endemism.

Grey-winged Trumpeter (Psophia crepitans capuru used in the original description until it ochroptera Pelzeln 1857). Pelzeln described this was rediscovered in JNP (Whittaker et al. 1995). trumpeter as a distinct species, Psophia ochrop- It is associated with seasonally flooded black- tera. Ochroptera is treated as a subspecies of P. water or Igapo´ forest (Whittaker et al. leucoptera in recent taxonomic literature (Pinto 1995, Borges et al. 2001). The MZUSP has skins 1978, Sherman 1996). We agree with Haffer collected in Manacapuru (MZUSP 16561) and (1974) who proposed that ochroptera is a Codaja´s (MZUSP 16387), and one specimen was subspecies of P. crepitans, the trumpeter species collected in JNP (MPEG 55855) in addition to distributed north of the Amazon River. P. specimens mentioned in Whittaker et al. (1995). crepitans has two other subspecies in addition to The northernmost record of N. amaurocephala is ochroptera: crepitans inhabiting the Guianan the Unini River (Whittaker et al. 1995) and the AOE and napensis, inhabiting the Napo and Imeri easternmost is the Amana˜ River (M. Cohn-Haft, AOEs. P. c. ochroptera has been recorded mostly pers. comm.) with no confirmed records from the in white-sand woodland and terra firme forest in upper portion of the Rio Negro (Haffer and JNP (Borges et al. 2001, Borges and Almeida Fitzpatrick 1985). It is suggested that N. 2011), where an adult female was collected (INPA amaurocephala forms a superspecies with N. # 576). A recent molecular systematic study ruficapilla and N. frontalis (Rassmussen and (Ribas et al. 2011), in addition to morphological Collar 2002) assessment (Haffer 1974), found that ochroptera Ivory-billed Aracari (Pteroglossus azara azara is diagnosable from the other subspecies in the Vieilot 1819). This subspecies of the Ivory-billed Psophia crepitans complex. Aracari has the upper mandible brownish colored Chestnut-headed Nunlet (Nonnula amauroce- and is recorded from Castanheiro, Codaja´s, phala Chapman 1921). This rare puffbird was Igarape´ Cacau Pereira (Haffer 1974), Manacapuru known only from specimens collected in Mana- (MZUSP 16833 and 16834), Maraa˜(MPEG 18 THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 124, No. 1, March 2012

42505–42507), and JNP (Borges et al. 2001, moborus myotherinus and concluded that arde- Borges 2007). It is replaced by P. a. flavirostris in siacus was a diagnosable taxon This subspecies the Napo and Imeri AOEs and by P. a. mariae was described from Manacapuru and has been south of the Solimo˜es (Haffer 1974). There is recorded in Igarape´ Cacau Pereira, Santa Maria, some evidence of hybridization between mariae and Codaja´s (Zimmer 1932, Pinto 1978), Maraa˜ azara near the mouth of the Purus River (Haffer (MPEG 42654–42672), and Jau´ National Park 1974: 222–223). This Aracari has been recorded in (MPEG 50614–50620), and is replaced in Imeri both terra firme and flooded in JNP (Borges and Napo by M. m. elegans and in Inambari by M. et al. 2001, Borges and Almeida 2011). m. myotherinus (Haffer and Fitzpatrick 1985). Lafresnaye’s Piculet (Picumnus lafresnayi This antbird has been recorded in mature upland pusillus Pinto 1936). P. pusillus was described forest, white-sand campinaranas, and secondary from Codaja´s on the left margin of the Rio growth forest in JNP (Borges et al. 2001, Borges Solimo˜es by Pinto (1936). This taxon was later and Almeida 2011). reassigned as a subspecies of P. aurifrons Another seven avian taxa also are apparently (Peters 1948). Recent considered restricted to central-western Amazonia: Amazona pusillus as a subspecies of Picumnus lafresnayi autumnalis diadema, Brachygalbula lugubris (Winkler et al. 1995, Winkler and Christie phaeonota, Nonnula rubecula simulatrix, Hylex- 2002). P. lafresnayi forms a superspecies with estastes stresemanni stresemanni, Sclerurus rufi- P. pumilus that is apparently restricted to gularis brunnescens, Myrmoborus lugubris stic- southern and northwestern Brazil topterus, and Hylophylax naevius obscurus. These (Winkler and Christie 2002). There are six subspecies, however, are known only from a few MZUSP specimens identified as P. l. pusilus specimens and their taxonomic status and distri- collected in the Manacapuru region (MZUSP bution needs additional evaluation. 16614–16619). The plumage of a single speci- Napo and Imeri Areas of Endemism.—The Imeri men collected in JNP and deposited in the INPA and Napo areas of endemism were supported by 21 Bird Collection matched better with P. l. and 56 avian taxa as originally described (Haffer pusillus than with P. pumilus. The geographical 1978, Cracraft 1985). An updated distributional distribution of pusillus appears to be restricted and taxonomic assessment decreased these num- to the region west of the lower Rio Negro being bers to six species (or subspecies) endemic to Imeri replaced by P. pumilus in the Imeri AOE and 42 endemic to the Napo AOE (Appendix). (Fig. 2). P. l. pusillus has been found mainly Several bird species previously thought to be in flooded forests and secondary growth in restricted to the upper reaches of the Rio Negro JNP (Borges et al. 2001, Borges and Almeida have had their geographical distribution extended 2011). southward and to northern (Borges et al. Ruddy Spinetail (Synallaxis rutilans confinis 2001, Alonso and Whitney 2003, Borges and Zimmer 1935). This subspecies was described Almeida 2011). Other Imeri birds are restricted to from specimens collected in Igarape´ Cacau the northwestern portion of the Amazon Basin Pereira in the lower portion of Rio Negro. It is with no confirmed records in the lower reach of replaced by S. r. dissors on the left margin of Rio the Rio Negro or northern Peru. Most species Negro, which has been found in Manaus and in restricted to Imeri AOE are specialists in white the upper Rio Negro at Sa˜o Gabriel da Cachoeira sand campinas (Cyanocorax helprini and Myrme- and along the Casiquiare channel (Friedmann ciza pelzelni), montane and pre-montane forests 1948, Pinto 1978). A better delimitation of the (Percnostola caurensis), and flooded forest (Thri- distribution of S. r. confinis will require specimen pophaga cherriei) (Zimmer 1999; Hilty 2003; collections south of the Negro and Uaupe´s rivers. SHB, unpubl. data). S. r. confinis is replaced in Napo AOE (eastern Similarly, some birds previously thought to be and northeastern Peru) by S. r. caque- endemic to the Napo region had their distributions tensis and in Inambari AOE (upper Amazonia extended east to the Rio Negro, as in the case of south of the Solimo˜es River) by S. r. amazonica Myrmotherula ignota and Pteroglossus pluricinc- (Peters 1951, Pinto 1978). tus (Borges and Almeida 2011). Some species are Black-faced Antbird (Myrmoborus myotherinus even more widespread than previously described, ardesiacus Todd 1927). Haffer and Fitzpatrick but there are species endemic to northwestern (1985) evaluated the plumage variation in Myr- (Imeri endemics), upper north Amazon (Napo Borges and Silva N A NEW AREA OF ENDEMISM FOR AMAZONIAN BIRDS 19 endemics), and the central-western portion of the necessary (Fig. 2). We also note this region of basin (this study). faunal turnover could be not coincident with any river course as happens in the southern portion of DISCUSSION the Amazon Basin (Haffer 1992). We describe a previously undesignated area of One fundamental concern in recognizing and endemism in central-western Amazonia that is delimiting the Jau´ AOE is the current taxonomy supported by the congruent distribution of six of the birds considered endemics. The use of avian taxa. Haffer and Fitzpatrick (1985) noted subspecies in biogeographic analyses is problematic that a small number of strongly differentiated because an unknown number of bird taxa described bird populations were restricted to this region in the ornithological literature are not discrete but did not recognize it formally as an AOE due evolutionary units (Cracraft 1985, Haffer 1987). the reduced number of avian taxa (J. Haffer, Some authors have been successful in using pers. comm.). Conceptually and operationally, subspecies for biogeographic analysis (Bates et al. however, an AOE could be recognized with just 1998, Borges 2007). Analyzing polytypical species two species (Platinick 1991, Harold and Mooi as a single entity tends to over-estimate the 1994). geographical distributions of taxa and potentially Amorin and Pires (1999) also identified a small ignores relevant units for conservation and bioge- AOE bounded by the Rio Solimo˜es and Rio Negro ography (Bates and Demos 2001). We considered without offering information about which species only taxa with accentuated morphological differen- supported this biogeographic component (Amorim tiation. However, the taxonomic status of Amazo- and Pires 1999: fig. 27). We suggest naming this nian bird species, including those discussed here, new area of endemism as Jau´ in recognition of the needs to be continuously evaluated and their importance of the National Park to biodiversity phylogenetic relationships clearly established. conservation in the Amazon. This area appears These taxonomic and biogeographic studies will nominated as Rio Negro Area of Endemism in require additional ornithological collections in the previous publications (Borges 2007, Ribas et al. northern portion of the upper Rio Solimo˜es and in 2011). However, we believe that Jau´ Area of the middle to upper Rio Negro, principally along the Endemism is more appropriate to the geographical right margin of the latter, as these regions are likely setting of the area (Fig. 2). contact zones. The precise geographical limits of this area are The phylogenetic relationships of the species only partially identifiable. The southern and and subspecies endemic to the Jau´ AOE are not eastern limits probably coincide with middle adequately known. However, three taxa (Pter- portions of the Solimo˜es and Negro, large rivers oglossus azara azara, Picumnus lafresnayi pusil- known to isolate bird populations (Sick 1967, lus, and Myrmoborus myotherinus ardesiacus) Haffer 1992). The northern boundary is more appear to have their closest relatives in the Imeri complicated, but from the middle Rio Negro and Napo AOEs. This is supported by Borges northward the landscapes dominated by terra (2007), who applied quantitative methods (parsi- firme forests are extensively replaced by forests mony analysis of endemicity and cluster analyses) and fields growing in sandy soils called ‘campi- to analyze the biogeographic relationships be- naranas’ and ‘campinas’ (Anderson 1981, IBGE tween avifaunas from different Amazonian AOEs. 1997). This more open vegetation occupies The avifauna of the JNP shares more species and thousands of square kilometers between the subspecies with localities in the Imeri and Napo middle and the upper Rio Negro. It is possible AOEs than with localities in the Inambari and this discontinuity in vegetation serves as a barrier Guiana AOEs (Borges 2007). This suggests the (or filter) to dispersal for some bird species, barrier effect of the Negro and Solimo˜es rivers is especially those that occur more frequently in stronger than the barrier effect in the western terra firme forest on clay soils. The western border of the Jau´ AOE. A working hypothesis to boundary of Jau´ AOE also is difficult to identify the evolutionary history of the six taxa restricted due to the scarcity of bird collections from the to the Jau´ AOE is that they were derived through region between Maraa˜ to the Brazilian-Colombian parapatric or peripatric speciation from ancient border. We tentatively suggest the courses of the species whose ranges were once centered in the Japura´orIc¸a rivers as the western boundary of Jau´ Imeri and Napo AOEs. Alternatively, tectonic AOE, although additional fieldwork will be events that affect the lower course of the Rio 20 THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 124, No. 1, March 2012

Negro (Forsberg et al. 2000, Almeida-Filho and BATES,J.M.AND T. DEMOS. 2001. Do we need to devalue Miranda 2007, Silva et al. 2007) could influence Amazonia and other large tropical forests? Diversity bird distribution in this region if they serve as and Distributions 7:249–255 BATES, J. M., S. HACKETT, AND J. CRACRAFT. 1998. Area- vicariance mechanisms (Ribas et al. 2011). relationships in the neotropical lowlands: a hypothesis We call attention to the urgent need to review the based on raw distributions of birds. Journal currently recognized AOEs for Amazonian birds. The of Biogeography 25:783–793. last review of these areas was compiled over 20 years BORGES, S. H. 2004a. Avifauna do Parque Nacional do Jau´: ago (Cracraft 1985), and taxonomic and geographical um estudo integrado em biogeografia, ecologia de distribution data have continued to accumulate. The paisagens e conservac¸a˜o. Dissertation. Museu Para- ense Emı´lio Goeldi, Para´, Brazil. areas of endemism for Amazonian birds should be re- BORGES, S. H. 2004b. Species poor but distinct: bird evaluated through modern biogeographic approaches assemblage in white sand vegetation in Jau´ National (Morrone and Crisci 1995, Szumik and Goloboff Park, Brazilian Amazon. Ibis 146:114–124. 2004) facilitated by continuous progress increasing BORGES, S. H. 2007. Ana´lise biogeogra´fica da avifauna da the data base of geographical distribution and regia˜o oeste do baixo Rio Negro, Amazoˆnia Brasileira. systematics of neotropical birds. Revista Brasileira de Zoologia 24: 919–940. BORGES,S.H.AND R. A. ALMEIDA. 2011. Birds of the Jau´ ACKNOWLEDGMENTS National Park and adjacent area, Brazilian Amazon: new species records with reanalysis of a previous checklist. SHB acknowledges Capes, WWF-Brazil, and Fundac¸a˜o Revista Brasileira de Ornitologia 19:108–133. Vito´ria Amazoˆnica for financial and technical support. We BORGES,S.H.AND A. M. P. CARVALHAES. 2000. Bird thank Brazil’s federal agency of environmental protection species of black water inundation forests in the Jau´ (IBAMA) for permission to work in Jau´ National Park. Luı´s National Park (Amazonas State, Brazil): their contri- Fa´bio Silveira, Maria Luiza, Alexandre Aleixo, and Mario bution to regional species richness. Biodiversity and Cohn-Haft kindly permitted us to examine bird collections Conservation 9:201–214. under their care. Marcelo Moreira kindly prepared the BORGES, S. H., M. COHN-HAFT,A.M.P.CARVALHAES, maps. Morton Isler, Charles Zartman, and Camila Ribas L. M. HENRIQUES,J.F.PACHECO, AND A. WHITTAKER. helped in first versions of the paper. We appreciate the 2001. Birds of Jau´ National Park, Brazilian Amazon: improvements in English usage by Phil Stouffer through the species check-list, biogeography and conservation. Association of Field Ornithologists’ program of editorial Ornitologia Neotropical 12:109–140. assistance. We are also grateful to J. M. 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APPENDIX. Distribution of avian taxa in Imeri and Napo areas of endemism as originally proposed by Haffer (1978, marked with H) and Cracraft (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 casiquiare—H Crypturellus casiquiare2 xx C. duidae—C C. duidae3 xxx Aramides calopterus—H Aramides calopterus E Mitu salvini—H & C Mitu salvini E M. tomentosa—H M. tomentosum xx Neomorphus pucheranii—C Neomorphus p. pucheranii4 xx Campylopterus villaviscensio—C Campylopterus villaviscensio E Leucippus chlorocercus—H & C Leucippus chlorocercus E Heliodoxa gularis—H & C Heliodoxa gularis E Topaza pyra—H & C Topaza p. pyra5 xxx Phlogophilus hemileucurus Phlogophilus hemileucurus E Pyrrhura albipectus—H & C Pyrrhura albipectus E Galbula tombacea—H Galbula tombacea tombacea6 E G. leucogastra chalcothorax G. chalcothorax7 E G. pastazae—H G. pastazae E Galbalcyrhynchus leucotis—H Galbalcyrhynchus leucotis E Nonnula ruficapilla rufipectus—C Nonnula ruficapilla rufipectus E N. brunnea—H N. brunnea E Notharchus ordii—H Notharchus ordii8 xxx Pteroglossus f. flavirostris—H Pteroglossus azara flavirostris xx P. pluricinctus—H P. pluricinctus xxx Selenidera r. reinwardtii—H & C Selenidera r. reinwardtii E S. nattereri—H & C S. nattereri xx Picumnus pumillus—H & C Picumnus pumillus E Celeus spectabilis—H & C Celeus spectabilis spectabilis E Hylexetastes stresemanni insignis—C Hylexetastes stresemanni insignis E Synallaxis moesta—H & C Synallaxis moesta E Thripophaga cherriei—H & C Thripophaga cherriei E Neoctantes niger—H Neoctantes niger9 xx Borges and Silva N A NEW AREA OF ENDEMISM FOR AMAZONIAN BIRDS 23

APPENDIX. Continued.

Original name Current name Imeri Napo Jau´ 1 Myrmotherula sunensis—H & C Myrmotherula sunensis sunensis10 E M. obscura—H M. ignota obscura11 xxx M. longicauda soderstromi—C M. longicauda soderstromi E M. cherriei—H & C M. cherriei12 xxx M. ambı´gua—H & C M. ambigua xx Herpsilochmus sticturus dugandi—H & C Herpsilochmus dugandi E H. dorsimaculatus—H H. dorsimaculatus13 xx Hypocnemis hypoxantha—C Hypocnemis h. hypoxantha14 xxx Pithys castanea—C Pithys castaneus15 E Rhegmatorhina m. melanosticta—C Rhegmatorhina m. melanosticta E R. cristata—C R. cristata xx Myrmeciza melanoceps—H Myrmeciza melanoceps16 E M. pelzelni—H & C M. pelzelni E M. disjuncta—H & C M. disjuncta17 xx Gymnopithys leucaspis castanea—C Gymnopithys leucaspis castaneus E G. leucaspis laterallis—C G. leucaspis lateralis xx Thamnophius praecox—H & C Thamnophius praecox E Percnostola caurensis—H Schistocichla caurensis18 E Grallaria dignı´ssima—H & C Grallaria dignı´ssima E G. fulviventris—H & C Hylopezus f. fulviventris E Pipra coronata coronata—H Lepidotrhix coronata coronata E P. coronata caquetae—C L. coronata caquetae E P. pipra discolor—C Dixiphia pipra discolor E Heterocercus aurantiivertex—H & C Heterocercus aurantiivertex E H. flavivertex—H & C H. flavivertex19 xx Attila citriniventris—H & C Attila citriniventris20 xxx A. torridus—C A. torridus E Myiophobus cryptoxanthus—H & C Myiophobus cryptoxanthus E Ramphotrigon fuscicauda—H Ramphotrigon fuscicauda21 E Todirostrum c. calopterum—H & C Poecilotriccus calopterus E T. c. capitale—H P. capitalis22 E Contopus n. nigrescens—C Contopus n. nigrescens E Microbates cinereiventris—H Microbates cinereiventris hormotus23 E Cyphorinus aradus salvini—C Cyphorinus arada salvini E Microcerculus bambla albigularis—C Microcerculus bambla albigularis24 E Cacicus sclateri—H & C Cacicus sclateri E Ocyalus latirostris—C Ocyalus latirostris E Hylophilus hypoxanthus fuscicapillus—C Hylophilus hypoxanthus fuscicapillus E H. b. brunneiceps—C H. brunneiceps 25 xx Cyanocorax heilprini—H & C Cyanocorax heilprini E Dolospingus fringilloides—H & C Dolospingus fringilloides26 xx

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 Solimo˜es (Payne 1997). Haffer (1997) reported N. p. pucherani in the upper Rio Negro (Imeri). (5) Recorded in sou- thern Venezuela (Hilty 2003). (6) Subspecies G. t. mentalis recorded south of Rio Solimo˜es (Inambari) (Tobias et al. 2002). (7) Also recorded in the Jurua´ River (Inambari) (Tobias et al. 2002). (8) Recently recorded in Inambari AOE (Guilherme and Borges 2011). (9) Records from Inambari, Imeri, and Tapajo´s AOEs (Zimmer and Isler 2003). (10) Subspecies M. s. yessupi recorded in Jurua´ River (Inambari) (Zimmer and Isler 2003). (11) Also recorded in Inambari AOE (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 AOE (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 Inambari AOE (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 Rondoˆnia AOE (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 AOE (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) Taphrospilus 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. hauxwelii inseparable from nominate (Fitzpatrick 2004).