Neotropical Ichthyology, 6(3):339-346, 2008
Copyright © 2008 Sociedade Brasileira de Ictiologia
A new species of Rineloricaria (Siluriformes: Loricariidae: Loricariinae)
from rio Daraá, rio Negro basin, Amazon, Brazil
Lúcia H. Rapp Py-Daniel1 and Ilana Fichberg2
Rineloricaria daraha, new species, is described from the rio Daraá, tributary of rio Negro, northwestern Amazonas State,
Brazil. The new species is diagnosed by having seven branched pectoral-fin rays, finger-like papillae on the lower lip, a large
multi-angular preanal plate, and at least four quadrangular plates of variable size surrounding the preanal plate. The new
species is known only from rio Daraá and its waterfalls.
Rineloricaria daraha, espécie nova, é descrita do rio Daraá, um afluente do rio Negro, noroeste do Estado do Amazonas,
Brasil. A nova espécie pode ser diagnosticada por apresentar sete raios ramificados na nadadeira peitoral, lábios inferiores com
papilas digitiformes, uma grande placa pré-anal multi-angular e, pelo menos, quatro placas de diferentes tamanhos circundando
a placa pré-anal. A nova espécie é conhecida apenas do rio Daraá e suas cachoeiras.
Key words: Armored catfish, Freshwater fish, Taxonomy, Neotropical fish.
Introduction
Rineloricaria Bleeker, 1862, is the most species-rich genus of the Loricariinae with approximately 60 species
(Rodriguez & Reis, 2008; Ghazzi, 2008), and widely distributed
from Panama in Central America to northern Argentina, on
both slopes of the Andes (Reis & Cardoso, 2001; Covain &
Fisch-Muller, 2007). Rineloricaria has been diagnosed by a
combination of characters, such as presence of a small postorbital notch, surface of the inferior lip bearing short buttonlike papillae, each hemi-maxilla carrying up to 15 short
mandibulary teeth, teeth strong and deeply forked, dark bands
on the dorsal region of the body with the first one at the origin
of dorsal fin, pectoral fin with one spine and six branched
rays, a distinctly polygonal preanal plate usually surrounded
by three to five other plates, and conspicuous sexual
dimorphism in mature males, consisting of numerous
developed odontodes along the sides of the head, on the
pectoral-fin spine and predorsal area (Isbrücker & Nijssen,
1992: figs. 34-36). In males, the pectoral-fin spine is often thick,
short, and curved when compared to females. Males of some
species of Rineloricaria have well-developed odontodes all
over the predorsal area.
The genus Hemiloricaria Bleeker, 1862 had been largely
considered as a synonym of Rineloricaria until, Isbrücker
et al. (2001) resurrected the genus and created two others
(Leliella and Fonchiiichthys) to accommodate some species
previously included in Rineloricaria, based on differences
on sexual dimorphism. Ferraris (2003) maintained
Hemiloricaria as a synonym of Rineloricaria, but in 2007,
he also resurrected the genus without comments, but did
not consider Leliella or Fonchiiichthys as valid. Covain &
Fisch-Muller (2007) followed Ferraris (2003) and did not consider Hemiloricaria, Fonchiiichthys and Leliella as valid
names.
Rodriguez & Reis (2008) partly accepted Isbrücker et al.
(2001) phenetic proposition of spliting Rineloricaria and
Hemiloricaria. Rodriguez & Reis, however, proposed that
Hemiloricaria would comprise a wide distributed group of
species (Amazon and non-Amazon species) whereas
Rineloricaria would be restricted to species occurring in rio
Paraná and its tributaries, and the coastal drainages from Uruguay to northeastern Brazil.
We describe herein a very distinctive new species from rio
Daraá, a tributary to the rio Negro, in northwestern of
Amazonas State, Brazil, in the genus Rineloricaria, following
Covain & Fisch-Muller (2007), but we expect that additional
information on the phylogeny and biogeography of this group
will eventually clarify the taxonomic limits and phylogenetic
relationships of that genus.
1
Programa de Coleções e Acervos Científicos, Instituto Nacional de Pesquisa da Amazônia (INPA), Av. André Araújo, 2936, Petrópolis,
69011-970 Manaus, AM, Brazil. rapp@inpa.gov.br
2
Museu de Zoologia da Universidade de São Paulo (MZUSP), Av. Nazaré, 481, Ipiranga, 04263-000 São Paulo, SP, Brazil. ilanafic@ib.usp.br
339
340
A new species of Rineloricaria from rio Negro basin
Material and Methods
Measurements follow Boeseman (1976), Reis & Cardoso
(2001) and Rodriguez & Miquelarena (2005). Straight-line distances were measured with digital calipers. The term lateral
abdominal plate follows Reis & Pereira (2000) terminology for
what is sometimes called thoracic plates. Dorsal and pectoral
spines refer to the first unbranched and thick ray of each fin.
This term is preferred to “unbranched or simple ray” due to
the presence of non-pungent spines on some loricariids that
are homologous to the unbranched first dorsal and pectoralfin ray in loricariines, and to avoid confusion with the
unbranched last rays of all fins. Measurements in Table 1 are
expressed as proportions of the Standard Length (SL), except
for subunits of the head, which are expressed as proportions
of Head Length (HL). Plate counts and nomenclature follow
schemes of serial homology proposed by Schaefer (1997).
Some paratypes were not measured due to reduced size or
bad state of preservation.
Several Rineloricaria species were not available for close
examination and had, therefore, their morphological conditions determined from literature accounts. Institutional ab-
breviations are INPA (Instituto Nacional de Pesquisas da
Amazônia), MZUSP (Museu de Zoologia da Universidade de
São Paulo), NMW (Naturhistorisches Museum Wien), and
MCZ (Museum of Comparative Zoology in Harvard).
Results
Rineloricaria daraha, new species
Figs. 1-3
Holotype. INPA 28579, 200.9 mm SL (male), Brazil, Amazonas,
rio Daraá, cachoeira do Aracu (00°25’00”S 064°46’59”W), 29 Nov
1991, R. Sotero & R. Ribeiro.
Paratypes. (20) Brazil, Amazonas, rio Daraá. INPA 6586, 4, 129.3186.8 mm SL, same data as holotype. MZUSP 31396, 1 c&s, 127.5
mm SL, female, cachoeira do Aracu, 10 Feb 1980, M. Goulding.
MZUSP 35094, 2, 40.7-46.6 mm SL, females, cachoeira do Aracu,
10 Nov 1980, M. Goulding. INPA 6587, 2, 57.7-65.9 mm SL,
cachoeira do Pacu, 1 Dec 1991, L. Aquino. INPA 12045, 7, 63.0117.4 mm SL, cachoeira do Pacu, 1 Dec 1991, L. Aquino. INPA
17939, 3, 41.3-74.5 mm SL, cachoeira do Aracu, 10 Nov 1980, M.
Goulding, INPA 28780, 1, 75.0 mm SL, juvenile, rio Daraá, cachoeira
do Panãpanã, 00°02’15.2”S 64°47’44”W, 4 Feb 2008, M. Rocha &
V. Masson.
Fig. 1. Rineloricaria daraha. holotype. INPA 28579; 200.9 mm SL; Brazil, Amazonas, rio Daraá, Aracu falls.
L. H. Rapp Py-Daniel & I. Fichberg
Diagnosis. Rineloricaria daraha is distinguished from all its
congeners by having seven branched pectoral-fin rays (vs.
six), long digitiform papillae on the ventral surface of the lower
lip (vs. button-like papillae) and by the presence of a large
and multi-angular preanal plate limited anteriorly by four or
more variably sized plates much smaller than the preanal plate
(vs. a quadrangular preanal plate surrounded by three to five
polygonal plates slightly smaller than the preanal plate).
Description. Morphometric data in Table 1. Body elongated
and depressed. Dorsal profile slightly convex from tip of snout
to origin of dorsal fin. Body slender along its whole extension, tapering softly towards base of caudal fin. Ventral profile straight from tip of snout to pelvic fin. Lateral profile from
pelvic fin to caudal fin becomes more depressed towards caudal-fin base where body becomes more flattened. Greatest
body depth at dorsal-fin base.
Head elongated and depressed. Snout long (53-59% of
head length) and pointed with small naked area. Naked area
not reaching most anterior pore of infraorbital sensory canal.
Orbit small and round, with deep and short lanceolate postorbital notch. Head variably keeled; when present, keel reduced
to low ridge in front of eye.
Predorsal area not keeled, except by weak pair of carinae
on last predorsal plate. Twenty seven lateral plates conspicuously carinated forming a keel. Two longitudinal and confluent
keels meeting at 14th plate of median series. From this point
on, remaining thirteen carinated plates of both lateral series
form single keel until base of caudal fin.
Mouth opening small and surrounded by short upper lip
and well-developed lower lip. Margin of lip fringed with small
filaments. Rictal region with well marked groove. Above
groove, line of button-like papillae cover maxillary barbel.
Papillae much longer at tip of maxillary barbel. Upper lip with
few button-like papillae, lower lip surface covered (densely
covered in some specimens) by long digitiform papillae, resembling short filaments. Digitiform papillae concentrated
around mouth cavity. Small specimens (smaller than 50 mm
SL) with round papillae on lip surface. Buccal papillae present
behind dentaries; few buccal papillae on premaxillaries. Most
of palate smooth, without papillae, except for straight line
with three to five papillae, longitudinally oriented, situated at
lateral tip of premaxillaries. Mandibular teeth short but robust, deeply forked: seven to eight teeth on premaxillaries
and five to seven on dentaries. Dentary teeth larger than
those on premaxilla.
Dorsal fin I,7, anterior and long, running along 8 or 9 dorsal plates; pectoral fin I,7 with spine almost straight on females or non-reproductive males, and thick and curved on
mature or almost mature males. Pectoral fin reaching third
plate of dorsal lateral series. Pelvic fin i,5 and anal fin i,5, welldeveloped. Pelvic fin reaching little beyond insertion of anal
fin mostly in females; pelvic fin of males not reaching anal-fin
insertion when adpressed. Pelvic unbranched ray thicker in
mature males. Anal fin running along seven plates. Caudal fin
341
i,10,i, bilobed to slightly emarginated, with upper lobe little
longer than lower lobe. Dorsalmost caudal-fin ray bearing
long and thick filament.
Abdominal surface completely covered by unorganized
small quadrangular plates in adults, except for naked gular
area. Five or six large lateral abdominal plates. Strong gradient of increasing plate size towards pelvic fin. Abdominal
plates tightly packed and forming shield. Anterior borderline
of this shield irregular, curved caudad in mid-line in some
specimens. Preanal plate present, conspicuously larger than
other plates and almost round in shape in some specimens.
Preanal plate surrounded by at least four quadrangular plates
of largely different sizes, with some much smaller than preanal plate itself. Abdominal plating development following
ontogenetic process with young specimens showing abdomen largely naked, with few odontodes spread all over abdominal surface. Young adults (approximately 130 mm) with
abdomen completely covered.
Color in alcohol. Light brown background densely covered
with conspicuous black round spots on head; faded and irregularly shaped dark blotches on body. Most spots on head
contain pores of cephalic sensory canal. Sphenotic and supraoccipital bordered by soft black connective tissue in recently preserved specimens. Two conspicuous black spots
just above naked area behind cleithrum, spots surrounding
one or two exits of lateral sensory canal. Five wide dark transverse bands on body: first just behind dorsal-fin base, three
bands along caudal peduncle and last one close to caudal-fin
base. Bands faded in some specimens. Ventral surface uniform. All fin rays irregularly stained of dark, forming large
bands of just spots on fins. Interradial membranes hyaline.
Dorsal, pectoral and caudal fins darkened at base.
Table 1. Morphometric characters for holotype and paratypes
of Rineloricaria daraha.
Hol
n
range
200.9 16 40.7-200.9
Percents of standard length
Head length
22.4 16
21.6-24.3
Predorsal length
33.6 16
31.1-34.4
Dorsal-spine length
16.7 16
15.5-20.0
Anal-spine length
13.2 16
12.6-18.8
Pectoral-spine length
12.8 16
12.8-19.1
Pelvic-spine length
12.2 16
12.2-17.4
Thoracic length
28.8 16
20.0-28.8
Abdominal length
13.9 16
11.6-14.5
Cleithral width
17.4 16
16.0-18.2
Body depth at dorsal-fin origin 9.7
16
8.6-11.1
Body width at anal-fin origin
12.7 16
8.2-13.2
Postanal length
62.2 16
53.4-63.7
Percents of head length
Head width
73.8 16
71.9-80.4
Snout length
59.0 16
53.3-59.9
Orbital diameter
13.7 16
13.7-17.5
Interorbital width
21.5 16
21.8-26.2
Head depth
38.0 16
33.7-90.6
Premaxillary ramus
7.3
16
6.1-11.2
mean
SD
22.5
33.4
18.5
16.4
17.5
15.6
26.8
13.0
17.2
9.5
11.1
58.3
0.66
0.93
1.53
1.69
1.83
1.26
2.09
0.71
0.69
0.74
1.44
3.42
77.0
57.4
15.9
22.9
59.8
9.1
2.87
1.74
1.29
1.51
24.7
1.69
Standard length (mm)
342
A new species of Rineloricaria from rio Negro basin
Fig. 2. Rineloricaria daraha. Paratypes, male (a and c = INPA 6586; 186.8 mm SL), female (b and d = INPA 6586; 131.2 mm SL).
L. H. Rapp Py-Daniel & I. Fichberg
Sex dimorphism. Mature males present the following features: large patch of thick enlarged odontodes on sides of the
head; pectoral-fin spine thick and strongly curved; pectoral
spine and rays heavily covered by enlarged series of
odontodes; shorter snout and fins when compared to young
343
and females (Figs. 2 and 3). Dorsal, pectoral, pelvic and anal
fins slightly longer on adult females than on mature males.
Distribution. The new species is only known from cataracts
of the rio Daraá (cachoeira do Aracu, Pacu and Panãpanã), a
Fig. 3. Rineloricaria daraha. Paratypes. Dorsal and ventral detail of head of sexually dimorphic male (a and c = INPA 6586;
186.8 mm SL) and female (b and d = INPA 6586; 131.2 mm SL).
A new species of Rineloricaria from rio Negro basin
344
tributary to the rio Negro, in the state of Amazonas, northwestern Brazil (Fig. 4).
Etymology. The specific epithet daraha (Daraá in Portuguese)
refers to the type locality.
Discussion
Several species of Rineloricaria have been described from
the Amazon basin. Most of these represent a taxonomic challenge due to poor original descriptions and unavailable type
material. Rineloricaria phoxocephala (Eigenmann & Eigenmann, 1889), described from Coary, and R. castroi Isbrücker
& Nijssen, 1984, are commonly found in big rivers and floating meadows (pers. obs. – LRP). Both species are slender and
with few abdominal series of plates. Rineloricaria
phoxocephala has a more acute, long snout and dark dots on
cephalic and lateral line pores that resemble R. daraha. However, the dots on R. phoxocephala are really reduced and
less dense than in R. daraha. Besides, R. phoxocephala has
well-organized abdominal plates, rather than the scattered
small plates on R. daraha. Rineloricaria castroi has conspicuous markings on the fins, with large alternating bands
of dark and light brown (Isbrücker & Nijssen, 1984: fig. 1).
Rineloricaria lanceolata (Günther, 1868) and R. heteroptera
Isbrücker & Nijssen, 1976, are often seen in small sandy-
bottom streams (pers. obs. – LRP) and can be identified by
color pattern. Rineloricaria lanceolata shows conspicuous
coloration consisting of black bands covering dorsal, anal,
pectoral and pelvic fins (Isbrücker, 1973: fig. 2). Rineloricaria
heteroptera resembles R. daraha on the shape of the head
and presence of densely marked dark blotches along the head
and body, even reaching the ventral region in some specimens (Isbrücker & Nijssen, 1976; figs. 1, 2). However, R.
heteroptera has a longitudinal band along the second and
third dorsal-fin rays and large quadrangular abdominal plates
organized in series, whereas R. daraha does not have any
longitudinal dark bands on fins and the abdomen is covered
by very reduced plates without any organization.
Rineloricaria beni (Pearson, 1924), R. hasemani Isbrücker &
Nijssen, 1979, R. konopickyi (Steindachner, 1879), R. mellini
(Schindler, 1959), R. microlepidota (Steindachner, 1907), R.
morrowi Fowler, 1940, R. teffeana (Steindachner, 1879) and R.
wolfei Fowler, 1940 were reported from the mainstem of the
Amazon (Reis et al., 2003) and are very poorly represented or
misidentified in fish collections. Rineloricaria eigenmanni
(Pellegrin, 1908), R. fallax (Steindachner, 1915), R. formosa
Isbrücker & Nijssen, 1979, R. platyura (Müller & Troschel,
1848) and R. stewarti (Eigenmann, 1909) have been reported
from the periphery of the Amazon, mainly Guyana Shield and
rio Negro basin and represent a group of slender species with
three to five organized series of abdominal plates.
Fig. 4. Type locality of Rineloricaria daraha, rio Daraá, Amazonas, Brazil.
L. H. Rapp Py-Daniel & I. Fichberg
Rineloricaria formosa, R. fallax, and R. platyura have a color
pattern that resembles R. morrowi and R. melini: the presence
of a conspicuous dark round spot on the supraocciptal or
predorsal area (Isbrücker & Nijssen, 1979: fig. 2, R. formosa).
Rineloricaria fallax is easily distinguished from R. daraha
by the presence of a large dark spot bordered by two well
defined lanceolate dark lines on the predorsal plate (Isbrücker
& Nijssen, 1979).
More than 30 species of Rineloricaria were recorded from
the Paraná, Uruguay and coastal drainages of northeastern,
southeastern and south Brazil. All of those species have the
abdominal area completely or partly plated (Rodriguez & Reis,
2008; Ghazzi, 2008). When completely covered, the abdominal plates are organized in series and the preanal plate is
surrounded by three large quadrangular plates. Otherwise,
the abdomen is almost completely naked, showing only large
quadrangular plates between the pelvis and urogenital opening (preanal plate and border plates). Despite the diversity of
plating exhibited by these sepcies, none has the pattern described for R. daraha.
Rineloricaria daraha also share some characters with
other loricariine genera. R. daraha, for instance, is unique in
showing digitiform papillae on the lower lip surface. The elongated papillae on the lower lip and the lack of organization of
the abdominal plates in R. daraha resemble the conditions in
species of Loricaria. Loricaria, however, differs from all
Rineloricaria by the presence of few and hypertrophied
mandibulary teeth (R. daraha 7-8 vs. Loricaria spp. 3-5), long
filaments covering almost the whole surface of both lips (vs.
digitiform papillae on the lower lip, more concentrated near
the mouth in R. daraha) and lack of a preanal plate (vs. a
conspicuous preanal plate in R. daraha). In addition, even
young specimens of Loricaria show long filaments on the
lips, whereas young specimens of R. dahara have buttonlike papillae.
Among the Loricariinae, only representatives of the genus Lamontichthys and R daraha have seven branched pectoral-fin rays. All remaining Loricariinae have only six
branched rays on the pectoral fin (Isbrücker & Nijssen, 1978).
Although the number of pectoral-fin rays and the lip structure found in the new species are unique within Rineloricaria,
sexual dimorphic features of the mature males are similar to
those found in several species of Rineloricaria (R. castroi,
R. phoxocephala, R. heteroptera, and others), and unlike
those found in species of Loricaria.
An interesting aspect of Rineloricaria daraha concerns
its distribution. Specimens of the new species were first collected by M. Goulding in 1980, then in 1991 by L. Aquino, R.
Sotero and R. P. Ribeiro, and again, in 2008, by M. Rocha and
Vitor Masson, all from the same locality. Several researchers
have been collecting around this region in the Amazon, but
no other sample has ever been obtained, suggesting that this
species might be restricted to the type locality, rio Daraá.
Despite the existing records, it seems unlikely that R. daraha
is confined only to rio Daraá and its waterfalls. Some
345
Rineloricaria species show a wide geographic area of occurrence (e.g.: R. heteroptera, R. lanceolata, R. castroi, R.
phoxocephala – pers. obs.) but we are not aware of any
Amazonian species of Rineloricaria with such a restricted
distribution. On the other hand, disjunct geographic range
has been registered on some representatives of Rineloricaria
from rio Paraná, Uruguay and southern Brazilian coastal drainages (Rodriguez & Reis, 2008; Ghazzi, 2008).
Material examined. Brazil: Rineloricaria castroi: MZUSP 15731,
holotype, 165.4 mm SL, Pará, Trombetas Biological Station, rio
Trombetas, 1º0’S 57º0’W; INPA 19997, 6, 98.8-122.0 mm SL,
Amazonas, Solimões, Paraná do Pirapora, Brasil; INPA 22123, 1,
122.97 mm SL, Amazonas, Manaus, rio Solimões, ilha da
Marchantaria. Rineloricaria fallax: NMW 44864, lectotype
(Steindachner, 1915), designated by Isbrücker & Nijssen, 1979,
igarapé do Carauná, near Boa Vista, rio Branco drainage, Roraima,
Brasil; paralectotypes: NMW 45046, 1, rio Branco, near Boa Vista,
NMW 44867, 2 (only one specimen listed as paralectotype by
Isbrücker & Nijssen, 1979), rio Branco, Boa Vista; NMW 44868, 1,
near Conceição, rio Branco, Boa Vista; NMW 46159, 2 (only one
specimen listed as paralectotype by Isbrücker & Nijssen, 1979),
Bem Querer, rio Branco. Rineloricaria formosa: MZUSP 38997, 5,
paratypes, 55.0-58.8 mm SL, Amazonas, Igarapé tributary of rio
Uaupes; MZUSP 38969, 2, paratypes, 71.3-81.4 mm SL,
Amazonas, igarapé Acaraposo, tributary of rio Tiquié; MZUSP
92367, 3, 113.80-127.39 mm SL, Amazonas, rio Negro drainage, rio
Tiquié, 00º10’S 069º07’W. Rineloricaria heteroptera: MZUSP
38954, 7, paratypes, 68.14-109.98 mm SL, rio Amazonas, Manaus,
reserve Ducke, 03º08’S 60º02’W; MZUSP 88996, 24, 56.92-120.79
mm SL, Amazonas, Manaus, rio Preto da Eva, 2º40’49.2”S
59º42’47.6”W; INPA 25862, 2, 107.0-139.7 mm SL, Pará, Oriximiná,
igarapé Periquito, below Saracá mine, lago Sapucuá. Rineloricaria
lanceolata: MZUSP 89303, 29, 36.9-86.8 mm SL, Goiás, Nova
Crixás, drenagem Araguaia, córrego Pitomba, 140 o 08’35”S
050º20’13”W; MZUSP 81379, 9, 33.3-79.8 mm SL, Amazonas, rio
Negro drainage, rio Tiquié, igarapé Onça, 00o13’52”N 69o51’5”W;
MZUSP 23445, 18, 42.4-76.6 mm SL, Amazonas, Fonte Boa,
Igarapé Tomé, Ati-Paraná, NW of Fonte Boa; MZUSP 93305, 1,
43.0 mm SL, Amazonas, rio Negro drainage, igarapé Cunuri (ou
Maracu), 00o13’00”N 069o36’00”W; MZUSP 24131, 56, 50.1-87.9
mm SL, Pará, Jatobal, rio Tocantins, 04 o 32’S 049 o 32’W;
Rineloricaria phoxocephala: MCZ 49057, 1, paratype, lago Coari,
Amazonas; INPA 25864, 1, 119.9 mm SL, Amazonas, Anavilhanas,
rio Negro; INPA 22074, 4, 91.6-105.1 mm SL, Amazonas, Manaus,
Paraná do Pirapora, rio Solimões. Rineloricaria platyura: NMW
44869 (paralectotype of Loricariichthys fallax), 1, Maguary (?),
near Pará. Loricaria cataphracta: INPA 8408, 2, 162.2-167.9 mm
SL, Amazonas, Coari, rio Solimões, mouth of lago Coari; MZUSP
57941, 1, 104.55 mm SL, Pará, rio Tapajós, below lago Azul.
Loricaria sp.: MZUSP 81257, 1, 171.01 mm SL, Amazonas,
drenagem do rio Negro, rio Tiquié, small beach in pond below
cachoeira do Caruru , 00 o 16’29”N 69º54’54”W. Guyana:
Rineloricaria fallax: NMW 44866, 5 (only one specimen listed as
paralectotype by Isbrücker & Nijssen, 1979), Rupununi River.
Acknowledgments
We want to thank José L. O. Birindelli and Hélio R. da
Silva, for their careful comments and suggestions on the manu-
346
A new species of Rineloricaria from rio Negro basin
script; Michael Goulding, Luis Aquino, Raimundo Sotero,
Roberval P. Ribeiro, Marcelo Rocha and Vitor Masson for
collecting the available specimens; Leandro M. Sousa and
Eduardo Baena for the illustrations; Edinho, Jander and Jandir
from the Secretaria de Meio Ambiente de Santa Isabel do Rio
Negro (Amazonas, Brazil) for providing the logistics in the
2008 field trip. MZUSP and INPA for the loan of specimens.
FAPESP grant no 04/01839-5 to IF and CNPq (Edital Universal
2004) grant for L.R.P. Field trip of M. R. and V. M. supported
by institutional projects of Jansen Zuanon and Lucia Rapp
Py-Daniel (INPA). Collecting license IBAMA no.11696-2.
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Accepted July, 2008
Published September 30, 2008