A new species of Epinecrophylla antwren from the Aripuanã-Machado interfluvium in central Amazonian Brazil with revision of the “stipple-throated antwren” complex.

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ORIGINAL SCIENTIFIC DESCRIPTIONS 263
Holotype.— Museu de Zoologia da Universidade de São Paulo
(MZUSP) 80591, adult female from Brazil: Amazonas; left bank
of the Rio Roosevelt in the municipality of Novo Aripuanã at
“Esperança trail” (08°29’00’’S/60°59’22’’W); elevation about
110 m; collected 26 September 2007 by Vítor de Q. Piacenti-
ni and Luís Fábio Silveira, prepared by Emerson Boaventura.
Voice not recorded. Pectoral muscle tissue preserved in ethanol
(MZUSP 80591),  eld number 106. Hologenetype (Chakrabarty
2010) sequences of the mitochondrial gene NADH subunit 2
(ND2, 1041 bp) deposited in GenBank (KC768944).
Diagnosis: Morphology.— Alphanumeric color designations
determined through direct comparison with Munsell soil color
charts (1994); colors in quotation marks are chart designations.
Adult females of E. dentei are readily distinguished from adult
E. h. amazonica by the buffy (“very pale brown” [10YR 8/2])
contour feathers of the throat contra white in E. h. amazonica.
Both taxa present black in the basal (non-contour) feathers of
the throat, producing the distinctly streaked pattern, which is
lacking or inconspicuous in females of E. h. pyrrhonota. Adult
females have belly “yellowish brown” (10YR 5/6), notably dis-
tinct from the paler and grayer “very pale brown” (10YR 7/3) of
adult female E. h. amazonica from the left bank of the upper Rio
Madeira, but females from the region of the Rio Juruá appear to
be indistinguishable by color of the posterior underparts. Males
may be weakly distinguished from those of E. h. amazonica by
tail more nearly concolor with the upperparts (contra olive-gray
and contrasting more strongly with the reddish upperparts in E.
h. amazonica); and weakly from E. h. pyrrhonota by the overall
paler upperparts. Voice.— The loudsong differs diagnostically
(Isler et al. 1998) in pace from those of other taxa in the E. hae-
matonota complex and individual note structure is nearly 100%
diagnostic (described below). Selected audio  les for inter-taxon
comparisons, including those used for spectrograms in Krabbe
et al. (1999) and this paper, are archived at the Internet Bird Col-
lection (IBC) website. Genetic divergence.— Separated from
its sister-taxon E. haematonota amazonica by approximately
3.1% uncorrected sequence divergence in the mitochondrial
gene ND2. Also, ND2 uncorrected sequence divergence from
all other members of the haematonota complex is 5–6.3 % (see
Phylogenetic relationships, below).
Distribution.— Restricted to central Amazonian Brazil on the
right bank of the Rio Madeira in the Aripuanã-Machado inter-
 uvium: from the left bank of the Rio Aripuanã upriver to its
con uence with the Rio Roosevelt, from which point upriver it
is known only from the left bank of the Roosevelt in the state
of Amazonas and extending into northwest Mato Grosso south
and west to the right bank of the Rio Machado (or Ji-Paraná) in
the state of Rondônia; southern range limits unknown (Fig. 1).
Description of holotype.— See color illustration. Several photos (Fig.
2) and some high-de nition video of both sexes in the hand and are archived at
the MZUSP. Alphanumeric color designations determined through direct com-
parison with Munsell soil color charts (1994); colors in quotation marks are chart
designations. Plumage fresh and unworn, tail and wing not in molt; skull 100%
A new species of Epinecrophylla antwren from the Aripuanã-Machado
interfl uvium in central Amazonian Brazil with revision
of the “stipple-throated antwren” complex
Bret M. Whitney1, Morton L. Isler2, Gustavo A. Bravo1, Natalia Aristizábal1,
Fabio Schunck3, Luís Fábio Silveira3, and Vítor de Q. Piacentini3
Early in the focused investigation of the avifauna of the Aripuanã-Machado inter uvium (2000-2003), BMW determined that a form
of Epinecrophylla4 (then Myrmotherula) haematonota was restricted to this “speciation block,” later termed a “mini-inter uve” (Cohn-
Haft et al. 2007), and was apparently isolated from the nearest other members of the haematonota complex by the formidable barrier of
the Rio Madeira. The loudsong of this population was audibly distinct from that of E. h. amazonica on the opposite bank of the Madeira
and differed even more conspicuously from those of relatively distant E. h. haematonota and E. h. pyrrhonota, across the Rio Solimões/
Marañón and in upper Amazonia. Analysis of a suf ciently robust sample of recordings to document these observed vocal distinctions
in concert with results of morphological and DNA-based phylogenetic analyses showing appreciable differentiation in some of these
parameters has convinced us that this restricted Aripuanã-Machado population is most appropriately introduced at the species level
within a revised E. haematonota complex. We propose to name it:
Epinecrophylla dentei
Roosevelt Stipple-throated Antwren
Choquinha-do-rio-roosevelt (Portuguese)
1 Louisiana State University, Department of Biological Sciences, Museum of Natural Science, 119 Foster Hall, Baton Rouge, Louisiana 70803, USA. (ictinia@earthlink.net)
2 Department of Vertebrate Zoology, Birds, National Museum of Natural History, Smithsonian Institution, P. O. Box 37012, Washington, D. C. 20013-7012, USA.
3 Seção de Aves, Museu de Zoologia da Universidade de São Paulo (MZUSP), Avenida Nazaré 481, Ipiranga, São Paulo, SP, Brazil CEP 04263-000.
4 Genus Epinecrophylla - new genus (Isler et al. 2006), split from Myrmotherula 8: 577.

HANDBOOK OF THE BIRDS OF THE WORLD
264
ossied. Crown slightly redder than “dark yellowish brown” (10YR 3/4) with
tiny feathers at base of bill and above eye slightly bufer; facial and auricular
region mottled slightly paler, contrasting with crown. Nape slightly redder than
crown but contrasting strongly with uniformly “dark red” (brighter than 2.5YR
4/8) mantle, scapulars, back, rump, and uppertail coverts; innermost secondaries
(overlying folded wing) washed with this same color but slightly browner over-
all and narrowly tipped paler. Tail and primary stack concolor with crown, thus
contrasting equally strongly with red upperparts; central rectrices ever so slightly
paler tipped. Tail graduated, outer rectrices 6.4 mm shorter than central pair. Main
color of throat about the same as facial region, slightly whiter than “very pale
brown” (10YR 8/2) and marked with ve 1.5 mm wide black streaks formed al-
most entirely by contrasting basal feathering (i.e., contour feathers are essentially
unmarked), the central streak about 19 mm long and about 10 mm longer than the
shortest, outermost ones. Breast becoming slightly darker and browner than throat,
nearest “light yellowish brown” (10YR 6/4) blending to darker “yellowish brown”
(10YR 5/6) through the posterior underparts. Upperwing coverts nearest “very
dark brown” (10YR 2/2) with more olivaceous bases, each feather marked with
a bold, yellowish-buff (nearest 10YR 7/8) spot at the tip encompassing both webs
about equally. Alula and tiny feathers at bend of wing blackish with narrow but
sharply contrasting buff-white margins on distal webs becoming slightly wider to-
ward the feather tip. Primary coverts blackish with minute reddish tips. Underwing
coverts plain whitish blending toward color of breast at their margins. Soft part
colors: bill dark gray; tarsus and feet gray; iris gray. Standard measurements:
total length (just before specimen preparation) 108 mm; bill (culmen from base at
skull) 14.6 mm; bill from anterior edge of nares 9.6 mm; bill width at anterior edge
of nares 3.5 mm; bill depth at anterior edge nares 3.9 mm; wing (chord) 50.3 mm;
tail 36.7 mm; tarsus 15.5 mm; mass 9.0 g.
Etymology.— Emílio Dente (1919-1995) certainly was one of
the most important collectors and specimen preparators in the
history of Brazilian ornithology. He worked in several differ-
ent research institutions, including the Department of Zoology
of the Secretary of Agriculture (today the Museum of Zoology
of the University of São Paulo) and the Adolfo Lutz Institution
in São Paulo, collecting mostly ornithological material in many
different regions of Brazil, mainly in Amazonia. Outstanding
among the many collections made by Dente were the thousands
of specimens secured on the Rio Capim in Pará and in the state
of Roraima, all assiduously prepared and labeled with his im-
peccable penmanship. With the name of this antwren we honor
Emílio Dente not only for having been a productive collector,
but also for his highly rened specimen preparation technique
for both birds and mammals, a talent that he shared with many
up-and-coming taxidermists in Brazil, leaving a legacy in this
eld, and for being the upright, honest person he always was.
Dente died in Itapetininga, São Paulo, at the age of 76.
The English and Portuguese names reference the type local-
ity of E. dentei on the Rio Roosevelt, which forms the southeast-
ern distributional barrier.
REMARKS
Type series.— The allotype of Epinecrophylla dentei is MZUSP
80589, adult male from the same locality as the holotype. Re-
maining paratypes of E. dentei are the following twenty-seven
specimens: MZUSP 80587 male, 80588 male, 80590 male, and
80592 female (all from the type locality); 92307 female and
92308 male (MT, left bank Rio Roosevelt); Museu Paraense
Emílio Goeldi (MPEG) 30864 male (AM, right bank Rio Ma-
deira, km 969 on Jacareacanga-Humaitá highway, Rio 9 de
Janeiro); 58721 male, 58722 female, 58723 female, 58724 fe-
male, 58725 male, 58726 male, and 58728 male (AM, Humaitá
[right bank Rio Madeira], Parintintin Indigenous Territory);
59023 male and 59024 female (AM, Humaitá [right bank Rio
Madeira], Ipixuna Indigenous Territory); 71105 male and 71141
female (RO, Machadinho d’Oeste, right bank Rio Ji-Paraná); In-
stituto Nacional de Pesquisas da Amazônia (INPA) 707 female
and 751 male (AM, left bank rio Aripuanã, rio Arauazinho, iga-
rapé Três Jacus, 127 km south Novo Aripuanã); 1829 male and
1831 female (AM, left bank rio Aripuanã, igarapé Arauazinho,
130 km south Novo Aripuanã); 1788 male (RO, REBIO Jaru,
right bank rio Jí-Paraná [Machado], mouth of igarapé Tarumã);
1789 male (RO, REBIO Jaru, right bank rio Jí-Paraná, “trilha da
torre”). Louisiana State University Museum of Natural Science
(LSUMNS) 182832 male, 182833 male, and 182834 female
(MT, left bank Rio Roosevelt).
There is no appreciable variation in the series of adult fe-
males, but it appears that immatures have plainer throats (i.e.,
less obvious streaked pattern). Males show some variation in the
extent of black and white in the throat feathers, especially those
toward the posterior edge of the patterned patch: some individu-
als appear darker-throated owing to less extensive white tips on
individual feathers. It appears that iris color of adult females is
clear grayish or pale beige, and that of adult males, a much dark-
er brownish-red; bill shapes of the sexes may also differ slightly
but this requires further investigation (Fig. 2). Immature males
(skull ossications less than 50%) have grayish eyes.
Ecology and behavior.— Epinecrophylla dentei is, like all other
members of the genus, a dead-leaf foraging specialist as de-
ned by Rosenberg (1990); specic foraging behavior closely
matches that described in detail for E. spodionota by Whitney
(1994). One interesting difference in its foraging behavior is
an apparent association with Megastictus margaritatus (Pearly
Antshrike), interpreted by Whittaker (2009) as Epinecrophylla
following Megastictus, but perhaps only seasonally. The asso-
Figure 1.
Geographic distribution of
specimens examined in the
Epinecrophylla haematonota
complex in south-central
Amazonian Brazil. Red dots
= E. dentei. Red star = type
locality of E. dentei on the left
bank of the Rio Roosevelt.
Black squares = E. amazonica.
Adjacent letters provide
documentation: S = specimen;
V = vocal recording. A white
X marks places where BMW
has searched for E. dentei and
is confident that it is absent
although it occurs immediately
across the Aripuanã/Roosevelt
or Machado rivers at these
points; a white ? in the narrow
headwaters region of the
Machado and Roosevelt marks
an area that has not been
inventoried where the species
is expected to occur and range
limits need to be determined.
Black lines mark the boundaries
of Brazilian states as indicated
by their official abbreviations:
AM = Amazonas; RO =
Rondônia; MT = Mato Grosso.
The federal highway BR-230
(“Transamazônica”) is shown
in white.
Figure 2.
Epinecrophylla dentei female
(left) and male captured in a
mist-net on the left bank of the
Rio Roosevelt in Mato Grosso,
Brazil. Note the different iris
colors and bill shapes of the
sexes. Further sampling will be
required to confirm whether this
apparent sexual dimorphism is
constant in the species.
Photo by Fabio Schunck
(11 August 2011).

Frequency (kHz)
Time (seconds)
3.02.5 3.5
2
4
0
0 1.00.5 1.5 2.0
6
4.0
8
10
3.02.5 3.5
2
4
0
0 1.00.5 1.5 2.0
6
4.0
8
10
A B
CD
ORIGINAL SCIENTIFIC DESCRIPTIONS 265
ciation of these two thamnophilids has been observed by BMW
on a number of occasions in several different months of the
year, and it has not been clear who is following whom. It seems
plausible that they both benet, the antwren taking advantage
of the vigilance of the much more sedentary antshrike to avoid
predation while investigating complex dead-leaf structures, and
the watchful, “sit-and-wait” foraging strategy of the antshrike
(Whitney and Rosenberg 1993) perhaps aided by the antwren’s
invasive foraging behavior ushing arthropods from hiding in
dead foliage. Over the course of many years of eldwork, this
association has never been observed by BMW anywhere a mem-
ber of the haematonota complex occurs together with Megastic-
tus outside the range of E. dentei, and it seems to be a behavio-
ral characteristic of both species unique to Aripuanã-Machado
populations. BMW suspects that the overlap of widespread and
ecologically almost identical E. leucophthalma (White-eyed
Antwren) with E. dentei throughout the range of the latter in for-
est habitat with little heterogeneity in the understory may have
driven dentei to minimize niche overlap by associating less with
mixed-species ocks, where accompanied by vigilant Tham-
nomanes antshrikes, and more with non-ocking Megastictus.
In other regions of overlap of Epinecrophylla leucophthalma
and E. haematonota, such as southern Amazonia west of the Ma-
deira, greater habitat heterogeneity (in particular, the abundance
of bamboo and uvially perturbed forest) seems to account for
their slight segregation (Zimmer and Isler 2003, BMW pers.
obs.). No genetic mixing has been observed or is to be expected
between these two Epinecrophylla antwrens; they are in sister-
clades (GAB pers. obs.) and have highly differentiated songs
(BMW pers. obs.). Thus, previously established reproductive
barriers have been operative in fostering their overlap in second-
ary contact despite essentially identical foraging ecologies. The
stomach of one male (MZUSP 92308) contained only insects,
including fragments of Hymenoptera (Formicidae), Orthop-
tera, and larvae of either Coleoptera or Lepidoptera. The nest
and eggs of E. dentei remain unknown. High-denition video of
Epinecrophylla dentei in habitat may be viewed at IBC.
Vocalizations.— Our sample of 141 recordings of members of
the Epinecrophylla haematonota complex was segregated into
four study populations: (1) pyrrhonota north of the Rio Napo
and Amazon; (2) haematonota south of the Napo and Amazon
south to the Abujao region of Ucayali in Peru east to the Rio Ju-
ruá in Amazonas and Acre in Brazil; (3) amazonica in the Juruá-
Madeira interuvium in Brazil extending northeast to the Purús-
Madeira interuvium just south of the Amazon and extending
southwest to the Río Manu region of Peru and Pando, Bolivia;
and (4) dentei in the region east of the Rio Madeira between
the Rio Aripuanã/Roosevelt and the Rio Machado. Although we
employed the names of subspecies to refer to these populations,
the distributions of them are only partially consistent with cur-
rent subspecies’ range descriptions. We determined that the type
of amazonica was collected on the right bank of the Rio Juruá
(Ihering 1904, Pinto 1945).
All members of the complex deliver a large number of
types of vocalizations for a small thamnophilid, and nearly all
of them are consistent across populations. The only vocaliza-
tion with sufcient samples that differed diagnostically was the
loudsong (Fig. 3), which differed signicantly (following Isler
et al. 1998, 2007) in pace. Thus, haematonota (n = 15) and pyr-
rhonota (n = 22) loudsongs were delivered at the fastest pace
(means 13.0 notes/sec and 16.2 notes/sec, respectively); ama-
zonica (n = 14) at the slowest pace (5.6 notes/sec); and the pace
of dentei loudsongs (n = 15) was intermediate (7.9 notes/sec).
Except for pyrrhonota and haematonota, the average pace of the
four populations differed signicantly. Differences in note and
interval length between amazonica and dentei barely failed the
more stringent Isler et al. (1998) test that values would not be
likely to overlap with larger samples, and note shapes (rounded
crowns in dentei, at tops in amazonica) also separated dentei
and amazonica loudsongs almost perfectly in “blind tests” of
spectrograms. The different note shapes are, no doubt, responsi-
ble for the subtly different auditory quality of the songs of these
two. “Call-series” were a second vocal type with potential to
differ substantially between some of the populations, but sample
sizes were so minimal as to make them a concern only for future
recording efforts.
Phylogenetic relationships and taxonomy.— DNA sequence
data for the mitochondrial gene NADH subunit 2 (ND2, 1041
bp) were obtained for 50 individuals in the genus Epinecrophyl-
la, representing the eight currently recognized species, and all
known forms within the E. haematonota complex (see SI for a
list of ingroup and outgroup taxa).
Phylogenies by maximum-likelihood and Bayesian infer-
ence methods (see details in SI) showed that all individuals of E.
dentei (east of the Rio Madeira) belong, with high support, in a
clade that is sister to all individuals identied as E. h. amazonica
(west of the Madeira; Fig 4). Moreover, this clade is embedded
within a well-supported larger clade containing three other ge-
netically distinctive lineages. The rst is E. spodionota (Andean
foothills); the second contains E. h. haematonota with embedded
E. fjeldsaai (south of the Rio Napo south to the Rio Marañón/
Solimões); and the third E. h. pyrrhonota (north of the Rio Napo
and Rio Solimões). Relationships among these lineages are not
well resolved, which supports the idea that E. haematonota is
paraphyletic and genetically diverse (Fig. 4). As described by
Zimmer (1932), these taxa are clearly distinguished by plumage
differences which, when combined with the substantial levels
of genetic diversity and some signicant levels of vocal dif-
ferentiation, support raising them to the species level. We sug-
gest that the most satisfactory classication for these lineages is
Figure 3.
Loudsongs of taxa in the
Epinecrophylla haematonota
complex. (A) E. dentei: Brazil:
Mato Grosso; left bank Rio
Roosevelt (Whitney BMW
C 0462 in Isler inventory).
(B) E. amazonica: Bolivia:
Pando; Río Abuña (Parker TAP
034 006). (C) E. pyrrhonota:
Venezuela: Bolívar; Río Caura
(Whitney BMW 071 005).
(D) E. haematonota: Brazil:
Amazonas; Reserva Natural
Palmarí (Whitney BMW
185 041). See Krabbe et al.
(1999) for a representative
spectrogram of the loudsongs of
E. spodionota and E. h. fjeldsaai.

0.05
5.0 %
6.3 %
3.1 %
5.7 %
Outgroups
E. gutturalis
E. spodionota
E. dentei
E. amazonica
E. h. haematonota
+
E. h. fjeldsaai
E. pyrrhonota
HANDBOOK OF THE BIRDS OF THE WORLD
266
to recognize ve species within the complex: E. haematonota
(Napo Stipple-throated Antwren), E. pyrrhonota (Negro Stip-
ple-throated Antwren), E. amazonica (Madeira Stipple-throated
Antwren), E. dentei (Roosevelt Stipple-throated Antwren), and
E. spodionota (Foothill Stipple-throated Antwren).
Our analysis of ND2 revealed that E. fjeldsaai (2 specimens
from proximate localities whose identity was conrmed by
Krabbe et al. [1999]) is embedded within haematonota (0.5–
1.2 % sequence divergence between fjeldsaai and closer hae-
matonota), notwithstanding that we had no samples of haema-
tonota from very near the range of fjeldsaai (see SI). We do not
have a sufcient sample of loudsongs of fjeldsaai to determine
whether it is diagnosable from that of haematonota. However, in
its distinctive brown mantle, fjeldsaai presents obvious pheno-
typic differentiation that suggests a potential, as-yet unrecovered
genetic signature. Three distinct scenarios might explain this ob-
served pattern. First, it is possible that fjeldsaai and haematono-
ta are not reproductively isolated and that there is substantial
gene ow between them. Therefore, fjeldsaai’s distinctive phe-
notype would not represent a discrete change in plumage varia-
tion and intermediate phenotypes would be expected along their
contact zone. At the time of description of fjeldsaai as a species,
very few specimens (and no genetic analysis) were available.
Second, fjeldsaai and neighboring haematonota met in second-
ary contact, probably across most of fjeldsaai’s eastern distribu-
tional margin, before achieving complete reproductive isolation
but after attaining phenotypic differentiation, and now intro-
gression is taking place. Finally, it is possible that fjeldsaai and
haematonota speciated quite recently and became more or less
reproductively isolated but now are showing shared ancestral
polymorphism in mitochondrial genes that obscures their recent
history. Genetic divergences in the autosomal nuclear intron (b
brinogen intron 5 – bF5, 554 bp) between fjeldsaai and hae-
matonota also show that they are genetically indistinguishable
(0.1–0.4 % uncorrected sequence divergence; Bravo and Brum-
eld unpubl. data). At this time, because of the high similarity
in mitochondrial and nuclear haplotypes between fjeldsaai and
haematonota, we suggest that fjeldsaai is most appropriately
ranked as a subspecies of E. haematonota.
A robust analysis of population dynamics, including assess-
ment of the potential inuence of sexual selection for brown-
backed males in the range of fjeldsaai acting to stem introgres-
sion, will permit a much better-informed interpretation of their
evolutionary history and destiny. With acceptance of the above,
novel taxonomy would come a shift in English names; we pro-
pose the above to highlight signicant distributional attributes of
species within the stipple-throated antwren complex. We recom-
mend information-rich English names that unambiguously iden-
tify monophyletic lineages, even if they require as many or more
syllables as does the scientic name. "Roosevelt Antwren", for
example, strips away much of the utilitarian aspect of the name
in the sole interest of speaking/writing fewer syllables.
Conservation.— There are no imminent threats to the survival
of Epinecrophylla dentei, but the lack of protection of forest in
the narrow headwaters of the Machado/Roosevelt/Aripuanã riv-
ers, and the urgent need to secure adequate samples of specimens
from this region expressed by Whitney et al. (2013) in the de-
scription of Hypocnemis rondoni (this volume) is echoed here.
Acknowledgments.— Our sincere thanks to Waner Costa of
“Pousada Rio Roosevelt” in southern Amazonas state for his
hospitality to MZUSP personnel and permission to make im-
portant collections of birds on both banks of the river there in
September 2007. Marcelo Félix of MZUSP did an excellent job
of specimen preparation in Mato Grosso with BMW and FS in
August 2011. Thanks to the Fundação de Amparo à Pesquisa
no Estado de São Paulo (FAPESP) and Conselho Nacional de
Desenvolvimento Cientíco e Tecnológico (CNPq) for the con-
cession of grants (Evolução da Fauna de Vertebrados Terrestres
Brasileiros do Cretáceo ao Presente: Paleontologia e Filogenia,
CNPq 565046/2010-1), fellowships (LFS and VQP) and for the
authorization for collecting and Research by Foreigners (Carlos
Alberto Pittaluga Niederauer and Vânia Amaral Gurgel). The
Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais
Renováveis (IBAMA – SISBIO) and Instituto Chico Mendes de
Conservação da Biodiversidade for collecting permits. To Ener-
gia Sustentável do Brasil and ArcadisLogos, for personnel and
support during part of this study and Bruno Ehlers (United Par-
cel Services, UPS) for help with transport and eld equipment.
Molecular work was supported by grants to GAB from the Frank
Chapman Memorial Fund – AMNH, the American Ornitholo-
gists’ Union, the LSUMNS Big Day Fund, LSU Biograds, and
NSF (DEB-1011435). We are grateful to curators and collection
managers (see SI for list of museums) for allowing access to tis-
sue samples and specimens under their care, especially to Robb
T. Brumeld and Donna L. Dittmann (LSUMNS), Mark B. Rob-
bins (KU), F. Gary Stiles (ICN), and Ingrid Macedo (INPA). An-
drés M. Cuervo provided the DNA sequence from one specimen
housed at IAvH. Gabriel Bif of the Entomology Department of
the MZUSP graciously helped us by identifying arthropod stom-
ach contents. Our thanks to Phyllis Isler for kindly preparing the
spectrograms, and to Marco Antonio Rêgo for the map. We are
grateful to NASA for free and open access to the MODIS (EOS-
DIS) satellite imagery used to produce the map image. Fern-
ando Straube contributed some historical information on Emílio
Dente. Richard Banks and Thomas Schulenberg provided help-
ful comments on the manuscript. Hilary Burn painted the gures
of E. dentei that accompany this description.
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Figure 4.
Maximum-likelihood tree
topology of the Epinecrophylla
haematonota complex
within which E. dentei is
sister to E. amazonica, and
E. fjeldsaai is embedded within
E. haematonota. Relationships
among the four distinct lineages
in the complex remain poorly
resolved. All resolved nodes
have bootstrap support values
based on 1000 replicates >70
and posterior probability values
>0.95.

ORIGINAL SCIENTIFIC DESCRIPTIONS 267
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