HANDBOOK OF THE BIRDS OF THE WORLD A new species of

HANDBOOK OF THE BIRDS OF THE WORLD

282

Holotype.— Museu de Zoologia da Universidade de São Pau-

lo (MZUSP) 92306, adult male from Brazil: Mato Grosso;

left bank of the Rio Roosevelt in the Municipality of Colniza

(09°07’54’’S/60°42’31’’W) at about 150 m elevation; collected

5 August 2011 by Fabio Schunck, prepared by Marcelo Félix.

Voice recorded by Bret M. Whitney, original numbers BMW

13977-79; Macaulay Library of Natural Sounds (ML) 169980;

Isler inventory (ISL) BMW C 0518. Pectoral muscle tissue

preserved in approximately 96% alcohol: MZUSP 92306,  eld

number ROO-09.

Diagnosis: Morphology.— As is typical of the several allospe-

cies in the Hypocnemis cantator complex, morphological and

plumage differentiation is weak. Based on the large series of

the Hypocnemis cantator complex housed in the collections of

MZUSP and the Museu Paraense Emílio Goeldi (MPEG), H.

rondoni shows, on average, stronger rufous edging on the bases

of the rectrices. Voice.— The “common call” (sensu Isler et al.

2007) is immediately distinguishable in the  eld from those of

all other members of the complex and, in spectrographic analy-

sis, by its structure and pace (described below). Male loudsongs

are distinguished from those of the closely related H. ochrog-

yna (Rondonia Warbling-Antbird) by three characters; female

loudsongs by one (Isler et al. 2007). Selected audio  les for

inter-taxon comparisons, including those used for spectrograms

in Isler et al. (2007) and this paper, are available for listening

to on the Internet Bird Collection (IBC) website. Genetic di-

vergence.— Separated from its sister-species H. ochrogyna by

approximately 4.2% sequence divergence in the mitochondrial

gene ND2 (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. Alphanumeric color

designations determined through direct comparison with Munsell soil color charts

(1994); colors in quotation marks are chart designations. Plumage fresh and un-

worn, tail and wing not in molt; skull 100% ossi ed. Forecrown blackish directly

over base of bill, becoming whitish to either side to blend with whitish lores. Crown

blackish with a pale central stripe imparted by whitish proximal webs on these tiny

feathers, appearance of stripe varying toward centralized spot-streaking depending

on feather arrangement. Whitish supercilium contrasts with blackish sides of crown

and prominent blackish line from lores posterior through eye; auriculars similarly

whitish and de ned along the lower edge by a uniformly narrow blackish malar

streak. Nuchal region blackish irregularly speckled whitish. Mantle and back feath-

ers blackish with whitish proximal and “olive brown” (2.5Y 4/4) distal margins pro-

A new species of antbird in the Hypocnemis cantator complex from the

Aripuanã-Machado interﬂ uvium in central Amazonian Brazil

Bret M. Whitney

, Morton L. Isler

, Gustavo A. Bravo

, Natalia Aristizábal

, Fabio Schunck

Luís Fábio Silveira

, Vítor de Q. Piacentini

, Mario Cohn-Haft

, and Marco Antonio Rêgo

In June, 2000, during a brief visit to the town of Manicoré on the right bank of the middle Rio Madeira in Amazonas, Brazil, BMW and

MCH were attracted to an unusual vocalization of Hypocnemis

cantator (Warbling Antbird) that appeared to be homologous to a call

commonly heard from other populations across the wide range of the complex. Fieldwork by BMW in July and October, 2001 on the

middle and lower Rio Aripuanã and at the mouth of the Rio Roosevelt, and in June, 2002 on the lower Rio Machado (or Ji-Paraná) in

Rondônia with MCH de ned the distribution of this unique call, restricted to the Aripuanã-Machado inter uvium. The taxonomic study

of the H. cantator complex presented by Isler et al. (2007) con rmed the diagnostic characteristics of this vocalization and mapped its

geographical distribution, but did not introduce a name for the population (which was referred to as “implicata 2, taxon novum”). Col-

lection of fresh specimens of most members of the H. cantator complex accompanied by recordings of their songs and calls has permit-

ted DNA-based phylogenetic analyses that corroborate the taxonomic revision proposed by Isler et al. (2007) including the signi cant

differentiation of “implicata 2”, which we are now prepared to name:

Hypocnemis rondoni

Manicore Warbling-Antbird

Cantador-de-rondon (Portuguese)

Louisiana State University, Department of Biological Sciences, Museum of Natural Science, 119 Foster Hall, Baton Rouge, Louisiana 70803, USA. ([email protected])

Department of Vertebrate Zoology, Birds, National Museum of Natural History, Smithsonian Institution, P. O. Box 37012, Washington, D. C. 20013-7012, USA.

Seção de Aves, Museu de Zoologia da Universidade de São Paulo (MZUSP), Avenida Nazaré 481, Ipiranga, São Paulo, SP, Brazil CEP 04263-000.

Departamento de Biodiversidade (CBio) e Coleções Zoológicas - Aves, Instituto Nacional de Pesquisas da Amazônia (INPA), Manaus, AM, Brasil CEP 69080-971.

Genus Hypocnemis 8: 645.

ORIGINAL SCIENTIFIC DESCRIPTIONS

283

ducing an irregularly streaked effect; scapulars more purely “olive brown”. Semi-

concealed, white interscapular patch formed by more extensively white feathers

underlying central mantle. Lower back without black thus contrasting with mantle,

slightly paler than “dark yellowish brown” (10YR 4/6) becoming paler and slightly

rustier through rump (nearest 10YR 5/8). Rectrices essentially concolor with lower

back, margined with same, slightly rustier hue of rump. Tail weakly graduated, each

rectrix marked with a restricted blackish subterminal smudge (nearly obsolete in

dorsal aspect) inside a slightly more extensive, 1-mm wide buffy-white tip reduced

to minuscule points on central rectrices. Throat whitish with weakly mottled ap-

pearance imparted by slightly darker feather tips, this effect abruptly heightened

through breastband by more extensive black central stripes of individual feathers

increasing in width towards sides of breast. The resulting blotchy pattern changes

slightly posteriorly, along the lower edge of the breast/upper belly, as feather webs

return to whitish and black marking on individual feathers shifts posteriorly to en-

compass only a fringe, producing a faintly scalloped effect. Center of belly whitish.

Sides and anks distinctly orangish or rusty, somewhat brighter than “brownish

yellow” (10YR 6/8), the anteriormost feathers bearing irregular vestiges of black-

ish markings described for lower breast; undertail coverts concolor with anks and

unmarked. Lesser upperwing coverts black with discrete white dots encompassing

both webs at the feather tip, median coverts blackish tinged olive with slightly larger

white tips. Greater coverts nearest “olive brown” (10 YR 2.5 Y 4/4) with a blackish

smudge on the distal web anterior to the pale tip, these tips being concentrated on

the distal webs and grading from whitish on the outer part of the wing to distinctly

buffy or weakly orangish (slightly paler than 10YR 5/8) where they overlie the

folded remiges which feathers are nely margined with this same color. Alula deep

black with narrow but bold white margin concentrated on distal web; overlying

feathers at bend of wing with similar margination concentrated toward the feather

tip on both webs. Primary coverts blackish with paler, weakly orangish margins

blending with stack of folded primaries. Innermost, overlying secondaries this same

color washed very slightly paler at the tips with margins of proximal webs weakly

integrated and contrastingly paler (near 10YR 8/4). Underwing coverts white with

irregular blackish blotches. Soft parts in life: maxilla black, mandible gray; tarsus

and feet yellowish-gray; iris brown. Standard measurements: total length (just

before specimen preparation) 113 mm; bill (culmen from base at skull) 17.1 mm;

bill from anterior edge of nares 9.5 mm; bill width at anterior edge of nares 4.4 mm;

wing (chord) 52.2 mm; tail 39.9 mm; tarsus 17.7 mm; mass 12.5 g.

Etymology.— Cândido Mariano da Silva Rondon (1865-1958)

must have been a truly remarkable person. Among his many ac-

complishments as a leader of men, Rondon worked with Ben-

jamin Constant to articulate the “proclamation of the republic”

of Brazil in 1889; he headed the “Brazilian Boundary Inspection

Agency” that largely delimited the borders of that vast coun-

try as well as the “Strategic Telegraphic Commission” of Mato

Grosso that laid more than 7,000 kilometers of telegraph line

across remote western Brazil; and he organized and led the fa-

mous “Roosevelt-Rondon Expedition” that charted the course

of the rio da Dúvida (“River of Doubt”) that he then christened

the Rio Roosevelt — but we admire him most for his outstand-

ing qualities as a standard-bearer for the well-being and respect

of indigenous cultures, founding and for many years directing

the Serviço de Proteção aos Índios (“Indian Protection Bureau”)

which became the modern federal agency Fundação Nacional

do Índio (FUNAI). Rondon died in Rio de Janeiro at the age of

92. We are privileged to honor Marechal Rondon with the name

of this distinctive little antbird.

The English name calls attention to the discovery of H.

rondoni at the old Madeiran town of Manicoré. The correct

pronunciation is: money-co-RAY, with the accent on the nal

syllable. We think “Rondon’s Warbling-Antbird” would be

confusingly similar to the name of H. ochrogyna, Rondonia

Warbling-Antbird.

REMARKS

Type series.— The allotype of Hypocnemis rondoni is MZUSP

92305, adult female, from the same locality as the holotype.

The remaining paratypes of H. rondoni are the following fteen

specimens: MZUSP 62284 male (AM, left bank rio Aripuanã at

Prainha), 80610 male, 80611 male, 80617 male, and 80615 fe-

male (AM, left bank rio Roosevelt at “trilha Esperança”); MPEG

31146 female (AM, left bank rio Aripuanã at the mouth of the

rio Guariba), 57661-57665 female, male, male, female, male re-

spectively (AM, Manicoré, “rodovia do estanho” km 126-137);

and Instituto Nacional de Pesquisas da Amazônia (INPA) 749

female (AM, left bank rio Aripuanã at rio Arauazinho), 1798

sex unknown (RO, right bank upper rio Machado in Reserva Bi-

ológica do Jaru); Louisiana State University Museum of Natural

Science (LSUMNS) 182835 male and 182836 female (MT, left

bank rio Roosevelt).

Sex for sex, there is minor variation among specimens in

the extent (mostly width) of the breastband and pattern of dark

markings on individual feathers comprising it, and also in size

and extent of pale markings on the crown and wing coverts and

width of dark streaking in the mantle, but none of this is perti-

nent to diagnosability of the taxon.

Ecology and behavior.— Hypocnemis rondoni, like other mem-

bers of the H. cantator complex, forages in the understory of ter-

ra rme forest and joins mixed-species ocks only when these

pass through its territory. It does not appear to be associated with

any microhabitat variance but tends to occupy borders of light

gaps, treefalls, road edges, and other places where sunlight pen-

etrates to the forest understory promoting locally denser vegeta-

tive growth. Two stomachs were examined, one of which held

insects (Coleoptera and Orthoptera) and one had material too

fully digested to identify; stomach contents are preserved at

the MZUSP. The nest and eggs of H. rondoni remain unknown.

High-denition video of Hypocnemis rondoni in habitat may be

viewed on the IBC website.

Vocalizations.— When compared to common calls of other

taxa in the Hypocnemis cantator complex (Fig. 2), the common

call of rondoni is unique (Isler et al. 2007). This vocalization

(n = 25 from localities indicated “V” on Fig. 1; see SI for a list)

typically consists of four, less often three or ve, notes. The rst

note is short but usually embellished with overtones, typically

rises in frequency, and has a unique, screechy quality. The sub-

sequent two, three, or four short notes are delivered at succes-

sively higher frequencies although the nal two notes are some-

times at the same frequency. Pace is rapid, the most rapid of any

Hypocnemis [cantator] population, and rondoni never ends its

call with raspy notes as do some other populations. Principal

Figure 1.

Geographic distribution of taxa

in the Hypocnemis cantator

complex in south-central

Amazonian Brazil. Some

symbols were shifted slightly

from georeferenced locations

to permit better clarity of

relative positions at this scale

(the map is viewable at larger

scale in online SI). Red dots

= H. rondoni. Red star = type

locality of H. rondoni. Black

squares = H. ochrogyna. Black

triangles = H. peruviana. Black

diamonds = H. striata implicata.

The white circle on the right

bank of the Rio Roosevelt

marks an area from which

we have recordings of both

striata and ochrogyna. A white

? indicates areas that have

not been inventoried where

range limits of taxa need to be

determined. Adjacent letters

provide documentation: S =

specimen; V = vocal recording.

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.

3.0 4.0 5.02.0

1.0

6.0

A B C D

Time (seconds)

Frequency (kHz)

0.04

6.8 %

5.4 %

5.3 %

5.2 %

4.2 %

Outgroups

H. hypoxantha

H. subflava

H. cantator

H. flavescens

H. peruviana

H. rondoni

H. ochrogyna

H. s. striata

H. s. affinis

H. s. implicata

HANDBOOK OF THE BIRDS OF THE WORLD

284

variations in rondoni calls involve the rst note which is some-

times shaped like an inverted “U” and less often twisted into

a squiggle. An individual may deliver several variations. For

example, in one recording from Nova Olinda on the left bank

of the Rio Aripuanã (ISL BMW.200:46), a series of calls begins

with a rising rst note that gradually changes into an inverted

“U” and nally is distorted into a twisted shape. We suspect

that observed inter-individual variation in the sound/structure

of the rst note of the common call relays individual-specic

identication information. Isler et al. (2007) demonstrated that

common calls were important species-level indicators in the H.

cantator complex.

Although indistinguishable from the neighboring striata

population to the north, the male and female loudsongs of ron-

doni were found in an earlier study (Isler et al. 2007) to differ

diagnosably from ochrogyna, the population to the south, by

three characters and one character respectively and from peru-

viana to the west in an even greater number of vocal characters.

Vocal analysis for this study documented loudsong types of both

striata and ochrogyna in close proximity on the right bank of the

Rio Roosevelt (Fig. 1, symbols circled). Whether this interesting

situation reects sympatry of two species or perhaps introgres-

sion between them remains a (fascinating!) subject for further

research.

Phylogenetic relationships.— DNA sequence data for the

mitochondrial gene NADH subunit 2 (ND2, 1041 base pairs)

were obtained for 63 individuals in the genus Hypocnemis

from scattered locations across its distribution, representing the

seven currently recognized species (see SI for a list of ingroup

and outgroup taxa). Phylogenies by maximum-likelihood and

Bayesian inference methods (see details in SI) were largely

consistent with a previous species-level phylogenetic hypoth-

esis of the genus Hypocnemis (Tobias et al. 2008; Fig. 3). We

identied, however, three well-supported lineages south of the

Amazon and east of the Madeira instead of two: H. striata, H.

ochrogyna, and H. rondoni. The latter two represent genetically

distinctive groups that replace each other across the Aripuanã/

Roosevelt and Machado rivers and are more closely related

to each other than they are to any other lineage in the genus.

These results conrm that H. striata (sensu Isler et al. 2007)

is paraphyletic; additional observed genetic structure within it

remains unresolved and awaits further analysis of a denser and

appropriately geographically distributed sample. For the time

being, at least, we recommend maintaining implicata and afnis

as subspecies.

Conservation.— Hypocnemis rondoni has the most restricted

global population of any member of the H. cantator complex

and a smaller population than most currently recognized spe-

cies of Amazonian birds, but it is not currently threatened by

anthropogenic alteration of its habitat or other sources. Of great

concern, however, is the fragmentation and outright destruc-

tion of forest in the narrow headwaters region of the Machado,

Roosevelt, and Aripuanã rivers, which is mostly unprotected,

and some of these watersheds will soon be altered drastically

by installation of massive hydroelectric dams. Unless this area

is adequately sampled very soon, we stand to lose much criti-

cal information on the processes involved in the differentiation

of Amazonian birds in general and the dynamics of secondary

contact in particular.

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 and

also to the Instituto Brasileiro do Meio Ambiente e dos Recur-

sos Naturais Renováveis (IBAMA – SISBIO) for collecting

permits. We are grateful to Robb T. Brumeld and Donna L.

Dittmann at LSUMNS for allowing access to tissue samples

under their care, and to Ingrid Macedo of INPA and Alexan-

dre Aleixo and Fátima Lima of MPEG for help with specimens

under their care. Molecular work was supported by grants to

Figure 2.

Common calls of taxa in the

Hypocnemis cantator complex

in south-central Amazonian

Brazil. (A) H. rondoni (holotype

MZUSP 92306): Mato Grosso;

left bank Rio Roosevelt (ISL:

BMW C 0518). (B) H. striata

implicata: Amazonas; Prainha

Nova (ISL: BMW 195 030).

right bank Rio Machado

opposite Palmeiras (ISL: BMW

205 012). (D) H. peruviana:

Amazonas; 18 km west Humaitá

(ISL: BMW 195 020).

Figure 3.

Maximum-likelihood tree

topology of the genus

Hypocnemis showing

that H. rondoni is sister to

H. ochrogyna and that this pair

is sister to the as-yet unresolved

H. striata complex. All resolved

nodes have bootstrap support

values based on 1000 replicates

>70 and posterior probability

values >0.95.

ORIGINAL SCIENTIFIC DESCRIPTIONS

285

GAB from the Frank Chapman Memorial Fund – AMNH, the

American Ornithologists’ Union, the LSUMNS Big Day Fund,

LSU Biograds, and NSF (DEB-1011435). We are grateful to

NASA for free and open access to the MODIS (EOSDIS) satel-

lite imagery used to produce the map image. Gabriel Bif of the

Entomology Department of the MZUSP graciously helped us

by identifying arthropod stomach contents. Phyllis Isler kindly

prepared the spectrograms. Richard Banks and Thomas Schu-

lenberg provided helpful comments on the manuscript. Hilary

Burn painted the gures of H. rondoni that accompany this de-

scription.

Literature Cited

Isler, M. L., P. R. Isler, and B. M. Whitney (2007). Species limits in

antbirds (Thamnophilidae): The Warbling Antbird (Hypocnemis can-

tator) complex. Auk 124: 11–28.

Munsell Soil Color Charts (1994). Macbeth Division of Kollmorgan In-

struments Corporation, New Windsor, NY.

Tobias, J. A., J. M. Bates, S. J. Hackett, and N. Seddon (2008). Comment

on “The latitudinal gradient in recent speciation and extinction rates

of birds and mammals.” Science 319: 901.