Received: 1 November 2022 Accepted: 26 April 2023

DOI: 10.1111/jfb.15422

REGULAR PAPER FISH

s basin
Knodus borari, a new species from the lower Rio Tapajo
(Characiformes: Characidae)

Cárlison Silva-Oliveira 1 | André L. Colares Canto 2 | Frank Raynner V. Ribeiro 2

1

s-Graduação em Sociedade, Natureza e Desenvolvimento, Universidade Federal do Oeste do Pará, Santarém, Brazil
Programa de Po
2
Instituto de Ciências e Tecnologia das Águas, Universidade Federal do Oeste do Pará, Santarém, Brazil

Correspondence

s-
Cárlison Silva-Oliveira, Programa de Po Abstract
Graduação em Sociedade, Natureza e
A new species of Knodus from the Rio Cupari, a right-bank tributary of the lower Rio
Desenvolvimento, Universidade Federal do
Oeste do Pará, Rua Vera Paz, s/n, 68040-255
s, is described. Among its congeners, the new species shares exclusively with K.
Tapajo
Santarém, Pará, Brazil.
figueiredoi the presence of only uni-tricuspid teeth in the premaxilla, maxilla and den-
Email: carlison3@gmail.com
tary. Knodus borari differs from K. figueiredoi by the colour pattern and morphological
Funding information
characteristics. The new species is the fifth species of the genus described from the Rio
Conselho Nacional de Desenvolvimento

gico, Grant/Award
Científico e Tecntolo
s basin, and the second restricted to the lower portion of the basin. This is addi-
Tapajo
Numbers: 436763/2018-4, 310480/2022-1,

s basin is a region of ichthyofaunal endemism.
tional evidence that the lower Rio Tapajo
317781/2021-9; FAPESPA (ICAAF 024/2014);
Instituto de Conservação Ambiental The

s
Nature Conservancy do Brasil (Águas Tapajo KEYWORDS
Project, Cooperation BR FY20 104) Amazonian fishes, biodiversity, endemism, Stevardiinae, taxonomy

1 | I N T RO DU CT I O N freshwater drainages in South America (Ferreira & Ohara, 2023;

Knodus Eigenmann has been traditionally defined based on homoplastic
features of external morphology. Among the Stevardiinae, it has been
differentiated from Bryconamericus Eigenmann only by the presence of
scales covering part of the caudal fin (sensu Eigenmann, 1918). None-
theless, based on molecular data, Thomaz et al. (2015) obtained the
most comprehensive phylogeny of Stevardiinae, in which a Knodus
sensu stricto is recognized as more closely related to Rhinobrycon Myers,
and the clade composed by these two genera is the sister group of a
more inclusive clade composed of other genera of Diapominii, including
Bryconamericus sensu stricto. Subsequent studies combining both mor-
phological and molecular data evidenced congruent results, supporting
the recognition of Knodus (sensu Thomaz et al., 2015) being closely
related to Rhinobrycon (e.g., Ferreira et al., 2021; Mirande, 2019).
Results of Thomaz et al. (2015) suggest for the first time that Bry-
conamericus could have a more restricted distribution, demonstrating
that species from the Amazon and Orinoco basins attributed to Bryco-
namericus could be more closely related to Knodus s.s. Thus, Thomaz
et al. (2015) proposed the relocation of some species of Bryconameri-
cus to Knodus and the synonymy of Bryconadenos Weitzman,
Menezes, Evers & Burns. Therefore, as currently defined, Knodus com-
prises 40 valid species distributed throughout most cis-Andean
Fricke et al., 2023). Despite the recent increase in alpha taxonomy
knowledge of the genus, the real diversity of Knodus remains underes-
timated (Aguiar et al., 2022; García-Melo et al., 2019).
Four valid species of Knodus are currently known to inhabit the

s drainage: Knodus cupariensis Sousa, Silva-Oliveira, Canto &
Rio Tapajo
Ribeiro, Knodus heteresthes Eigenmann, Knodus dorsomaculatus Fer-
reira & Netto-Ferreira and Knodus ytuanama Ferreira & Ohara. None-
theless, the number of species related to the genus Knodus is

s basin (e.g., Guimarães et al., 2018;
underestimated in the Rio Tapajo
Ohara et al., 2017; Ohara & Loeb, 2016; Silva-Oliveira et al., 2016).
During recent expeditions in Rio Cupari, a right-bank tributary of the

s, an undescribed species of Knodus was collected.
lower Rio Tapajo
2 | M A T E R I A L S A N D M ET H O D S
2.1 | Ethics statement and specimen preservation
All material used in this study was collected in accordance with Brazilian
law, under scientific collection licence number 44215-2 granted by Insti-
tuto Chico Mendes de Conservação da Biodiversidade (ICMBio). No
experimental procedures were performed with live fish. Specimens were

J Fish Biol. 2023;1–7. wileyonlinelibrary.com/journal/jfb © 2023 Fisheries Society of the British Isles. 1
2
FISH SILVA-OLIVEIRA ET AL.

anesthetized in a solution containing eugenol (clove oil), fixed in 10% 3.3 | Paratypes
formalin solution and subsequently transferred to 70% ethanol.
All collected with the holotype: INPA 59900, 5, 28.1–32.2 mm Ls.
UFOPA 1365, 8, 22.8–35.4 mm SL, 2 c&s 32.2–33.1 mm Ls.
2.2 | Morphological analysis

Counts and measurements were taken according to Fink and Weitzman 3.4 | Diagnosis
(1974), except for the number of rows of scales below the lateral line,
which were counted as far as the pelvic-fin origin. Measurements are Knodus borari differs from all its congeners, except K. figueiredoi Esguí-
expressed as percentages of standard length (LS), with the exception of cero & Castro, by possessing only conical to tricuspid teeth in the
subunits of the head, which are expressed as percentages of head length
(LH). Counts of vertebrae, supraneurals, procurrent caudal-fin rays, gill
rakers on the first branchial arch and branchiostegal rays were taken from
cleared and stained specimens (c & s) prepared according to the method
of Taylor and van Dyke (1985). Vertebral counts included the four
vertebrae in the Weberian apparatus. The compound caudal centrum
(PU1 + U1) was treated as a single element. In the description, counts
are followed by their absolute frequency in parentheses, and holotype
counts are indicated by an asterisk (*). Institutional abbreviations follow
Sabaj (2020). In the list of comparative material examined, museum abbre-
viations and catalogue numbers are followed by the total number of spec-
imens in each lot, range of standard length and abbreviated collection
data. The map was generated using QGis 2.4.0 Chugiak©.

3 | RESULTS

3.1 | Knodus borari, new species

(Figures 1–3; Table 1).

urn:lsid:zoobank.org:pub:6EEB7A7A-2D20-4077-8FF9-F89C55EB8CD2;
urn:lsid:zoobank.org:act:7B6032CA-7AE8-4870-9674-9ECFFB739858;

3.2 | Holotype F I G U R E 2 Lateral view of left side of upper and lower jaws of
Knodus borari, UFOPA-I 1365, paratype, 33.4 mm Ls. Scale bar = 1 mm
UFOPA-I 1364, 33.8 mm Ls; Rio Braço Leste, tributary of Rio Cupari,

s basin, Ruro
Rio Tapajo
polis, Pará, Brazil, 04
070 12.100 S 55
000 12.000
W; F. Ribeiro, C. Silva-Oliveira, D. Sousa, H. Napoleão, J. Sousa,
R. Caroline & T. Torres, 5 May 2015.

F I G U R E 1 Holotype of Knodus borari, UFOPA-I 1364, 33.8 mm
Ls. Rio Braço Leste, Rio Cupari, right-bank tributary of lower Rio
Tapajo
s basin, Ruro

polis, Pará State, Brazil
F I G U R E 3 Coloration in life of Knodus borari. UFOPA-I 1365:

s
(a) 35.2 mm Ls; (b) 32.3 mm Ls. Rio Cupari drainage, lower Rio Tapajo
basin, Ruro
polis, Pará State, Brazil
SILVA-OLIVEIRA ET AL.
FISH 3

TABLE 1 Morphometric data of holotype and 14 paratypes of Knodus borari

H Range Mean S.D.

Standard length (Ls, mm) 33.8 22.8–35.4 30.2 –

Percentages of Ls
Snout to dorsal-fin origin 51.2 51.2–52.8 51.9 0.6
Snout to pectoral-fin origin 23.4 23.4–26.2 25.3 1.1
Pectoral-fin length 21.3 21.3–23.2 21.8 0.8
Snout to pelvic-fin origin 45.3 44.7–47.9 46.3 1.3
Pelvic-fin length 15.1 14.2–16.4 15.4 0.8
Snout to anal-fin origin 60.1 60.1–62.7 61.3 1.1
Depth at dorsal-fin origin 29.0 26.8–30.7 29.0 1.4
Dorsal-fin height 24.3 24.3–25.2 24.7 0.4
Dorsal-fin base length 11.5 10.9–11.8 11.4 0.4
Dorsal-fin origin to caudal-fin base 50.6 50.6–51.4 51.1 0.3
Anal-fin base length 25.4 25.4–28.5 27.4 1.3
Anal-fin lobe length 18.3 17.1–19.2 18.0 0.9
Caudal peduncle length 13.6 13.6–18.9 16.6 1.9
Caudal peduncle depth 10.7 10.7–11.7 11.4 0.4
Eye to dorsal-fin origin 39.9 38.6–41.2 39.8 1.0
Head length 22.5 22.5–24.6 23.4 0.8
Upper-jaw length 7.1 7.0–7.7 7.3 0.3
Horizontal eye diameter 8.3 8.3–10.5 9.1 0.9
Snout length 5.0 4.8–5.3 5.1 0.2
Least interorbital width 8.6 8.6–9.6 9.0 0.4
Percentages of HL
Upper-jaw length 28.6 28.6–33.3 31.2 1.7
Horizontal eye diameter 36.6 36.6–42.9 39.0 2.8
Snout length 19.6 19.6–22.5 21.6 1.2
Least interorbital width 37.7 37.7–39.4 38.7 0.8

Note: Holotype (H) is included in the ranges and means.

Abbreviations: S.D., standard deviation.

inner premaxillary row and dentary teeth (vs. 3–5 cusps in K. rufford
Deprá, Ota, Júnior & Ferreira, and 5–8 cusps in the other congeners).
It can be distinguished from K. figueiredoi and K. rufford by having the
shorter maxilla, not exceeding (vs. exceeding) the anterior margin of
the orbit in a vertical plane. Additionally, K. borari differs from
K. figueiredoi by the presence (vs. absent) of a black blotch at the base
of caudal fin and by possessing i, 6, i (vs. i, 5, i) pelvic-fin rays, and from
K. rufford by having 6 (vs. 4–4½) scales between dorsal-fin origin and
lateral line, 13–14 circumpeduncular scales (vs. 11–12) and shorter
snout (19.6%–22.5% LH vs. 24.2–26.7 LH).
3.5 | Description
Morphometric data in Table 1. Body compressed; greatest body depth
just anterior to dorsal-fin origin. Dorsal profile of head distinctly con-
vex from upper jaw to posterior nostril, slightly convex from latter
point to posterior margin of the eye; slightly concave from this point
to tip of supraoccipital spine; slightly convex from this point to dorsal-
fin origin; dorsal-fin base straight; slightly convex to straight from end
of dorsal-fin base to adipose fin. Ventral profile of body convex from
lower lip to anal-fin origin; straight, posterodorsally inclined along
anal-fin base. Dorsal and ventral profile of caudal peduncle slightly
concave.
Mouth sub-terminal; jaws isognathus. Posterior extension of max-
illa not extending beyond anterior margin of orbit. Premaxillary teeth
in two rows; outer row with 3 (15), conical or tricuspid teeth, central
cusp when present longest; inner row with 4* (16) tricuspid teeth,
central cusp longest. Maxilla with 2 (4) or 3* (12), tricuspid teeth. Den-
tary with 3 (4) or 4* (11) large tricuspid teeth, followed by 3–4 smaller,
uni- to tricuspid teeth (Figure 2). Scales cycloid, moderately large, with
1–6 well-marked radii; circuli only present proximally. Lateral line com-
plete, slightly curved ventrally along its anterior third, with 36 (3), 37*
(9) or 38 (4) perforated scales. Longitudinal series of scales between
4
FISH
dorsal-fin origin and lateral line, 6* (16). Longitudinal series of scales
between lateral line and pelvic-fin origin, 3* (16). Predorsal scales
10 (2), 11 (4) or 12* (9), arranged in a single regular row. Scales around
caudal peduncle 13* (10) or 14 (15). Single series of 15–16 scales cov-
ering proximal third of anal fin. Base of caudal fin scales smaller than
of caudal peduncle. Scales covering proximal fourth of upper and
proximal third of lower caudal-fin lobe.
Dorsal fin rays ii, 8 (16). Dorsal-fin origin near middle of body, and
posterior to vertical through pelvic-fin origin. First unbranched dorsal-fin
ray approximately half length of second unbranched ray. Adipose-fin ori-
gin approximately at vertical through base of 16th to 17th branched
anal-fin rays. Pectoral-fin rays i, 9 (10); i, 10* (5); or i, 11(1); tip of pecto-
ral fin reaching pelvic-fin origin when adpressed. Pelvic-fin rays i, 6, i*
(16); tip of pelvic fin usually not reaching first anal-fin rays when
adpressed. Anal-fin origin located anterior to vertical through posterior
terminus of dorsal-fin base. Anal-fin rays v, 17* (9); 18 (4); or 19 (3); last
unbranched ray and first three or four branched anal-fin rays longest,
subsequent rays decreasing gradually in length. Caudal fin forked; lobes
equal in size. Principal caudal-fin rays i, 9, 8, i* (16); dorsal procurrent
caudal-fin rays 12 (2); ventral procurrent caudal-fin rays 10 (2).
Gill rakers on first gill arch 15 (1) or 16 (1): hypobranchial 2 (2),
ceratobranchial 6 (2), cartilage between epibranchial cartilage and cer-
atobranchial 1 (2), epibranchial 5 (1) or 6 (1). Gill rakers setiform. Bran-
chiostegal rays 4 (2): 3 (1) on anterior ceratohyal and 1 (2) on
posterior ceratohyal. Supraneural 5 (2). Precaudal vertebrae 16 (2);
caudal vertebrae 19 (2); total vertebrae 35 (2). First dorsal-fin ptery-
giophore inserted between 11th and 12th vertebrae.
3.6 | Sexual dimorphism
None of the examined specimens showed secondary sexual dimor-
phism in morphology, colour patterns or presence of bony hooks.
3.7 | Geographical distribution
Knodus borari is known from the Rio Cupari drainage, a tributary of

s, Pará State, Brazil (Figure 4).
the right margin of the lower Rio Tapajo
3.8 | Colour in alcohol
General body coloration light brown to pale yellow. Infraorbital series,
opercular regions and gular region silvery. Snout, dorsal region of
head, sixth infraorbital, opercle and anterior portion of maxilla dusky
with scattered melanophores. Infraorbital bones 1–5, posteromedial
portion of the maxilla and gular region yellowish. Melanophores more
concentrated on the scales edge of middorsal and dorsolateral
regions, forming slightly reticulate patterns; ventral and ventrolateral
scales of body with few scattered dark chromatophores or uniformly
pale. Rectangular humeral blotch, vertically elongated, extending
SILVA-OLIVEIRA ET AL.
vertically through two longitudinal scales series above the lateral line
and horizontally encompassing two lateral line scales. Dusky mid-
lateral stripe present from immediately posterior to humeral blotch to
caudal fin base. Dorsal fin with chromatophores more concentrated
along the inter-radial membrane. Hyaline adipose fin with few scat-
tered chromatophores. Hyaline pectoral and pelvic fins with few scat-
tered chromatophores along anterior and posterior borders of the
rays. Anal fin slightly darkened with chromatophores at the interradial
membranes, more concentrated in the distal portion, forming a dark
band. Caudal fin with melanophores concentrated in the basal portion
of the central rays forming a black blotch extending posteriorly to
midlength of central caudal-fin rays.
3.9 | Colour in life
General colour of body grey to yellowish. Lips, snout, anterior por-
tion of maxilla dark. Upper portion of the head and dorsal portion of
the body dark grey to yellowish. Dorsal region of orbit yellow. Ven-
trolateral region of orbit, infraorbital series, posteromedial portion
of maxilla, opercular regions, gular region and ventrolateral portion
of body silvery. Dorsal region of caudal peduncle with orange spots.
Base of dorsal, adipose, pectoral and caudal fins yellow. Medial por-
tion of anal fin yellowish; pelvic fins similar to those of preserved
specimens.
3.10 | Conservation status
Although K. borari is currently known from its type locality in the Rio
Cupari basin, no specific threats have been identified that put their
population at risk. Thus, following the IUCN categories and criteria
(IUCN, 2022), K. borari can be categorized as Least Concern (LC).
3.11 | Ecological notes
The Rio Braço Leste, the type locality of K. borari, is a fourth-order
stream (sensu Strahler, 1957), c. 13 m wide and 1–3 m deep, with
moderate to fast-flowing waters, substrate composed mainly of rocks,
gravel and sand, with the following physicochemical parameters:
pH 6.8; dissolved oxygen 6.5 mg l1; conductivity 27.4 μS cm1; and
temperature 25.8
C (Figure 5). Specimens of K. borari were cap-
tured in a section of the margin in more lentic environments, with
depths ranging from 10 to 80 cm. Specimens of K. borari were cap-
tured alone and together with other species of Characiformes such
as Astyanax aff. bimaculatus (Linnaeus), Bryconops sp., Creagrutus
petilus Vari & Harold, Cyphocharax spilurus (Günter), Hemigrammus
sp., Hyphessobrycon pulchripinnis Ahl, Jupiaba atypindi Zanata, Kno-
dus sp., Microschemobrycon callops Böhlke, Moenkhausia hasemani
Eigenmann, Moenkhausia lepidura (Kner), Moenkhausia sp., and Rhi-
nopetitia sp.
SILVA-OLIVEIRA ET AL.
FISH 5

F I G U R E 4 Map of South America,

showing the geographical distribution
of Knodus borari (yellow circle). Red
area represents the lower Rio Tapajo
s.
White squares represent the Pimental
(1) and São Luiz rapids (2)

F I G U R E 5 Type locality of Knodus

borari, Rio Braço Leste, Rio Cupari
drainage, Rio Tapajo
s basin, Ruro

polis,
Pará State, Brazil

3.12 | Etymology Knodus is not yet available. Thus, new species have been assigned to
Knodus following traditional classification, mainly due to the pres-
The species epithet borari refers to the Boraris, indigenous people ence of scales on the proximal half of each caudal fin lobe (e.g.,

s. A noun in apposition.
who inhabit the lower Rio Tapajo Deprá et al., 2021; Menezes & Marinho, 2019; Sousa et al., 2020),
but also by the evidence provided by Thomaz et al. (2015) (e.g.,
Menezes et al., 2020:8).
4 | DISCUSSION Among its congeners, K. borari shares exclusively with
K. figueiredoi the presence of only unicuspid or tricuspid teeth in the
Despite recent advances in the understanding of phylogenetic rela- premaxilla, maxilla and dentary. Nonetheless, they differ from each
tionships in Stevardiinae using both molecular and morphological other by morphometric and meristic characters and colour pattern
data (e.g., Ferreira et al., 2021; García-Melo et al., 2019; (see Diagnosis). Furthermore, K. borari presents an unusual in vivo col-
Mirande, 2019; Thomaz et al., 2015), a morphological diagnosis of our pattern not recorded for another described congener, which
6
FISH
consists of a circular patch of orange pigmentation at the base of the
main rays of the dorsal lobe of the caudal fin (Figure 3).

s are
Most of the fish species described from the lower Rio Tapajo
widely distributed in the Amazon basin (Silva-Oliveira et al., 2021;
Sousa et al., 2020). Nonetheless, as a result of a recent increase in sam-
pling in that region (mostly in the past 10 years), several new species

s drainage
with distributions restricted to this portion of the Rio Tapajo
have been recognized and described [i.e., Ancistrus krenakarore de Oli-
veira, Rapp Py-Daniel, Zawadzki & Zuanon; Apistogramma arua Römer
& Warzel; Bryconops allisoni Silva-Oliveira, Canto & Ribeiro; B. durbinae
(Eigenmann); B. munduruku Silva-Oliveira, Canto & Ribeiro; Characidium
papachibe Peixoto & Wosiacki; Cyphocharax muyrakytan Bortolo,
Lima & Melo; Cyphocharax cramptoni Bortolo & Lima; Geophagus pyro-
cephalus Chuctaya, Nitschke, Andrade, Wingert & Malabarba; Gymnotus
arapiuns Kim, Crampton & Albert; Hyphessobrycon cantoi Faria,
Guimarães, Rodrigues, Oliveira & Lima; Hypostomus labyrinthus Oliveira,
Ribeiro, Canto & Zawadzki; Leptophilypnion pusillus Robert; and Knodus
cupariensis Sousa, Silva-Oliveira, Canto & Ribeiro].
K. borari is the fifth species of the genus described from the Rio

s basin, and the second with restricted distribution to the lower
Tapajo

s basin (see Ferreira & Ohara, 2023; Sousa et al., 2020).
Rio Tapajo
Nonetheless, other undescribed fish species occurs only in this area
(e.g., Guimarães et al., 2018; Silva-Oliveira et al., 2016; Sousa

s is a region
et al., 2020; Per. obs.), indicating that the lower Rio Tapajo
of high endemism for freshwater fish (sensu Platnick, 1991; Mor-
rone, 1994). Nonetheless, there is currently no consensus on the bio-
geographical delimitation of this region. Dagosta and de Pinna (2017)

s basin into four areas of endemism
subdivided the Rio Tapajo

s). In their study, Dagosta
(Juruena, Teles Pires, Jamanxim and Tapajo

s” as the stretch between the
and de Pinna (2017) considered “Tapajo
confluence of the Rio Juruena and Rio Teles Pires to the confluence
with Rio Amazonas, including the Rio Curuá-Una.
In contrast, Jézéquel et al. (2020) subdivided the basin into seven

s being considered the stretch below
sub-basins, with the lower Tapajo
the confluence of Rio Jamanxim to the confluence with Rio Amazo-
nas. In the present study, based on the distribution pattern of the spe-

s as the
cies listed earlier, the authors delimit the lower Rio Tapajo
stretch downstream of the São Luiz rapids to the confluence with Rio
Amazonas.
Compared with its upper and middle portions, a large portion

s is an alluvial plain, characterized mainly
of the lower Rio Tapajo
by many lentic environments and sandy beaches along its main
course (Buckup & Santos, 2010; Espirito-Santo et al., 2005). None-
theless, many of its tributaries provide a heterogeneous fast-
flowing environment below the rapids of Pimental and São Luís do

s, which are the major rapids of the lower Rio Tapaj o
Tapajo
s,
located downstream of the mouth of the Rio Jamanxim and ca.

s in the mainstream Amazon
300 km from the mouth of the Tapajo
(Goulding et al., 2003). As waterfalls are potential barriers to fish
dispersal (Barthem et al., 2016; Britski & Lima, 2008; Goulding
et al., 2003), these environments may be a contributing factor to
the notable level of endemism among fish species inhabiting the

s basin.
different sections and tributaries of the Rio Tapajo
SILVA-OLIVEIRA ET AL.
5 | C O M P A R A T I V E M A T E R I A L EX A M I N E D
All from Brazil except if otherwise noticed: Knodus angustus: INPA
40989, 15, 20.9–25.9 Ls, Rio Aripuanã. Knodus cinarucoense: INPA
55110, 3, 17.2–29.5 mm Ls, Rio Branco basin. Knodus breviceps: INPA
2478, 12, 26.5–39.7 mm Ls, Rio Tocantins basin; MCP 42274,
16, 16.8–43.6 mm Ls, Rio Tocantins basin, Goiás; INPA 41033,
1, 74.4 mm Ls, Rio Tocantins basin. Knodus cupariensis: INPA 59396,
40.5 mm Ls, holotype; INPA 59397, 11, 31.6–45.5 mm Ls (2 c&s,
34.8–39.5 mm Ls), paratypes, Rio Cupari. Knodus deuterodonoides:
INPA 22960, 10, 16.2–18.3 Ls, Rio Negro. Knodus dorsomaculatus:
INPA 45307, 6, 33.1–41.0 mm Ls, Rio Teles Pires basin; INPA 44974,
6, 37.2–52.2 mm Ls, Rio Teles Pires basin; MZUSP 98299, 2, para-
types, 39.2–41.8 mm Ls, Rio Teles Pires basin. K. figueiredoi: INPA
41243, 4, paratypes, 22.9–26.6 mm Ls, Rio das Garças basin. Knodus
geryi: MZUSP 83354, holotype, 63.4 mm Ls, upper Rio Paraguai basin.
Knodus heteresthes: INPA 44830, 14, 20.8–24.8 mm Ls, Rio Teles Pires
basin; INPA 45065, 50, 18.3–27.5 mm Ls, Rio Teles Pires basin; INPA
44626, 99, 24.3–32.3 mm Ls, Rio Teles Pires basin; MZUSP 25542,

s basin; UFOPA-I 152, 8, 13.7–
136, 19.1–32.5 mm Ls, Rio Tapajo

s; UFOPA-I 153, 5, 19.3–34.7 mm Ls,
36.6 mm Ls, lower Rio Tapajo

s. Knodus hypopterus: MCP 46397, 5, 36.5–62.6 mm
lower Rio Tapajo
Ls, Peru, Río Madre de Dios basin. Knodus megalops: MCP 37416,
112, 23.9–48.7 mm Ls, Peru, Río Ucayali basin. Knodus meridae:
MZUSP 99147, 2, 29.2–36.3 mm Ls, Venezuela, Río Zulia; MZUSP
99146, 4, 37.7–41.3 mm Ls, Venezuela, Río Chama. Knodus moen-
khausii: MCP 47157, 6, 31.3–40.1 mm Ls, Rio Paraná basin; MZUSP
93714, 80, 15.9–34.9 mm Ls, Rio Jequitinhonha basin. Knodus orte-
guasae: ANSP 70504, holotype, (only photo and X-ray), Colombia, Río
Orteguasa. MCP 41933, 38.7–50.5 mm Ls, Rio Acre basin, Acre; MCP
46390, 3, 31.4–43.8 mm Ls, Peru, Loreto, Río Madre de Dios basin.
Knodus savannensis: INPA 24270, 34, 16.1–35.8 mm Ls, Rio Tocantins
basin; INPA 39927, 4, 28.4–40 mm Ls, Rio Tocantins basin. Knodus
smithi: INPA 53987, 2, 25.2–29.0 mm Ls, Rio Madeira basin. Knodus
tiquiensis: INPA 40924, 26, 26.5–54.5 mm Ls, Rio Negro basin; INPA
049392, 15, 26.4–51 mm Ls, Rio Negro basin. Knodus victoriae: MCP
23435, 14, 27.7–33.5 mm Ls, Rio Parnaíba basin. Knodus ytuanama:
INPA 59847, paratypes, 9, 49.2–82.6 mm Ls, Rio Juruena basin.
AUTHOR CONTRIBU TIONS
Cárlison Silva-Oliveira contributed to the original idea, development
of the study and writing the manuscript. André L. Colares Canto and
Frank Raynner V. Ribeiro contributed with development of the study
and writing the manuscript.
AC KNOW LEDG EME NT S
The authors are grateful to Hugo Napoleão, Deise A. Sousa, Jordson
Souza, Raiany Oliveira and Thaís Torres (UFOPA) for their help and
assistance during fieldwork; and to Aléssio Datovo, Mario de Pinna,
Michel D. Gianeti (MZUSP), Lúcia Rapp Py-Daniel (INPA) and Carlos
Lucena (PUCRS) for curatorial assistance and Marcos Mirande for
valuable comments and suggestions on the manuscript. Cárlison
S. Oliveira and Frank R. V. Ribeiro are funded by the Conselho
SILVA-OLIVEIRA ET AL.

gico (CNPq pro-
Nacional de Desenvolvimento Científico e Tecntolo
cesses number 317781/2021-9 and 310480/2022-1, respectively).
André L. C. Canto and Frank R. V. Ribeiro were partly supported by
FAPESPA (ICAAF 024/2014), Conselho Nacional de Desenvolvimento

gico (Process 436763/2018-4) and Instituto de
Científico e Tecnolo
Conservação Ambiental The Nature Conservancy do Brasil (Águas

s Project, Cooperation BR FY20 104).
Tapajo
ORCID
Cárlison Silva-Oliveira https://orcid.org/0000-0002-8816-5218
RE FE R ENC E S
Aguiar, R. G., Guimarães, E. C., Brito, P. S., Santos, J. P., Katz, A. M.,
Dias, L. J. B. S., … Ottoni, F. P. (2022). A new species of Knodus
(Characiformes: Characidae), with deep genetic divergence, from the
Mearim and Munim river basins, northeastern Brazil, and evidence for
hidden diversity in adjacent river basins. Neotropical Ichthyology, 20,
e210173.
Barthem, R., Ferreira, E.J.G. & Goulding, M. (2016) As migrações do jaraqui
e do tambaqui no rio Tapajo
s e suas relações com as usinas hidrelétri-
cas. In: Alarcon, Daniela Fernandes; Millikan, Brent; Torres, Mauricio.
(Org.). Ocekadi: Hidrelétricas, conflitos socioambientais e resistência na
Bacia do Tapajo
s (pp. 479-493.). 1ed. Brasília: International Rivers Brasil.
Britski, H. A., & Lima, F. C. T. (2008). A new species of Hemigrammus from
the upper rio Tapajo
s basin in Brazil (Teleostei: Characiformes: Chara-
cidae). Copeia, 2008, 565–569.
Buckup, P. A., & Santos, G. M. (2010). Ictiofauna da ecorregião Tapajo
s-
Xingu: fatos e perspectivas. Boletim da Sociedade Brasileira de Ictiolo-
gia, 98, 1808-1436.
Dagosta, F. C. P., & de Pinna, M. (2017). Biogeography of Ama4zonian
fishes: Deconstructing river basins as biogeographic units. Neotropical
Ichthyology, 15(3). https://doi.org/10.1590/1982-0224-20170034
Deprá, G. C., Ota, R. R., Vitorino Júnior, O. B., & Ferreira, K. M. (2021).
Two new species of Knodus (Characidae: Stevardiinae) from the upper
rio Tocantins basin, with evidence of ontogenetic meristic changes.
Neotropical Ichthyology, 19, e200106.
Espírito-Santo, F. D. B., Shimabukuro, Y. E., de Aragão, L. E. O. e C. &
Machado, E. L. M. (2005). Análise da composição florística e fitosso-
ciolo
gica da floresta nacional do Tapajo
s com o apoio geográfico de
imagens de satélites. Acta Amazonica, 35, 155–173.
Eigenmann, C. H. (1918). The American Characidae. Memoirs of the
Museum of Comparative Zoology, 43, 103–208.
Ferreira, K. M., Mirande, J. M., Quagio-Grassiotto, I., Santana, J. C. O.,
Baicere-Silva, C. M., & Menezes, N. A. (2021). Testing the phylogenetic
hypotheses of Stevardiinae gill, 1858 in light of new phenotypic data
(Teleostei: Characidae). Journal of Zoological Systematics and Evolution-
ary Research, 00, 1–26.
Ferreira, K. M., & Ohara, W. M. (2023). A new rheophilic species of Knodus
Eigenmann (Characiformes: Characidae: Stevardiinae) from the upper
rio Juruena, rio Tapajo
s basin, Chapada dos Parecis, Mato Grosso,
Brazil. Zootaxa, 5227, 365–377.
Fink, W. L., & Weitzman, S. H. (1974). The so-called cheirodontin fishes of
Central America with description of two new species (Pisces: Characi-
dae). Smithsonian Contributions to Zoology, 172, 1–46.
Fricke, R., Eschmeyer, W. N., & Van der Laan, R. (2023). Eschmeyer's catalog
of fishes: Genera, species, references. San Francisco, California. Available
from: https://researcharchive.calacademy.org/research/ichthyology/
catalog/fishcatmain.asp: California Academy of Sciences accessed
February 17, 2023.
García-Melo, J. E., Oliveira, C., Costa Silva, G. J., Ochoa-Orrego, L. E.,
Pereira, L. H. G., & Maldonado-Ocampo, J. A. (2019). Online species
FISH 7
delimitation of neotropical characins (Stevardiinae): Implications for
taxonomy of complex groups. PLoS One, 14, 1–22.
Goulding, M., Barthem, R., & Ferreira, E. (2003). The Smithsonian atlas of
the Amazon. 44 Washington, DC, USA: Smithsonian Books.
Guimarães, K. L. A., de Sousa, M. P. A., Ribeiro, F. R. V., Porto, J. I. R., &
Rodrigues, L. R. R. (2018). DNA barcoding of fish fauna from low order
streams of Tapajo
s River basin. PLoS One, 13, e0209430.
IUCN. (2022). The IUCN Red List of Threatened Species. Version 2022-1.
Available from: http://www.iucnredlist.org.
Jézéquel, C., Tedesco, P. A., Bigorne, R., Maldonado-ocampo, J. A.,
Ortega, H., Hidalgo, M., et al. (2020). A database of freshwater fish
species of the Amazon Basin. Scientific Data, 7, 1–9.
Menezes, N. A., Ferreira, K. M., & Netto-Ferreira, A. L. (2020). A new spe-
cies of Knodus (Characiformes: Characidae: Stevardiinae) from the rio
Aripuanã, rio Madeira basin, Brazil. Neotropical Ichthyology, 18,
e190139.
Menezes, N. A., & Marinho, M. M. F. (2019). A new species of Knodus
Eigenmann (Characiformes: Characidae: Stevardiinae) with comments
on nuptial tubercles and gill gland in characiform fishes. PLoS One, 14,
1–18.
Mirande, J. M. (2019). Morphology, molecules and the phylogeny of Char-
acidae (Teleostei, Characiformes). Cladistics, 35, 282–300.
Morrone, J. J. (1994). On the identification of areas of endemism. System-
atic Biology, 43, 438–441.
Ohara, W. M., Lima, F. C. T., Salvador, G. N., & Andrade, M. C. (2017).
Peixes do Rio Teles Pires (p. 408). Gráfica e Editora Amazonas, Goiânia:
Diversidade e Guia de Identificação.
Ohara, W. M., & Loeb, M. V. (2016). Ichthyofauna of the upper Juruena
river on Chapada dos Parecis, Mato Grosso, Brazil. Biota Neotropica,
16, e20160224.
Platnick, N. I. (1991). On areas of endemism. Australian Systematic Botany,
4, xi–xii.
Sabaj, M. H. (2020). Codes for natural history collections in ichthyology
and herpetology. Copeia, 108, 593–669.
Silva-Oliveira, C., Canto, A. L. C., & Ribeiro, F. R. V. (2016). Stream ichthyo-
fauna of the Tapajo
s National Forest, Pará, Brazil. ZooKeys, 580,
125–144.
Silva-Oliveira, C., Ota, R. P., Lima, F. C. T., & Rapp Py-Daniel, L. (2021).
Rediscovering species: Redescription of Bryconops gracilis
(Characiformes: Iguanodectidae), an often-misidentified species. Neo-
tropical Ichthyology, 19, e210054.
Sousa, D. J. A., Silva-Oliveira, C., Canto, A. L. C., & Ribeiro, F. R. V. (2020).
A new species of Knodus (Characiformes: Characidae) from the Rio
Cupari drainage, lower Rio Tapajo
s basin, Brazil. Zootaxa, 4747,
575–584.
Strahler, A. (1957). Quantitative analysis of watershed geomorphology.
Transactions, American Geophysical Union, 38, 913–920.
Taylor, W. R., & van Dyke, G. C. (1985). Revised procedures for staining
and clearing small fishes and other vertebrates for bone and cartilage
study. Cybium, 9, 107–119.
Thomaz, A. T., Arcila, D., Ortí, G., & Malabarba, L. R. (2015). Molecular phy-
logeny of the subfamily Stevardiinae gill, 1858 (Characiformes: Chara-
cidae): Classification and the evolution of reproductive traits. BMC
Evolutionary Biology, 15, 1–25.
How to cite this article: Silva-Oliveira, C., Colares Canto, A. L.,
& Ribeiro, F. R. V. (2023). Knodus borari, a new species from

s basin (Characiformes: Characidae).
the lower Rio Tapajo
Journal of Fish Biology, 1–7. https://doi.org/10.1111/jfb.15422