Zoosyst. Evol. 98 (2) 2022, 327-343 | DO! 10.3897/zse.98.89413

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Redefinition of Heptapterus (Heptapteridae) and description of
Heptapterus carmelitanorum, anew species from the upper Parana
River basin in Brazil

Gabriel de Carvalho Depra!, Gast6n Aguilera*, Dario R. Faustino-Fuster?*°, Axel M. Katz°®,
Valter M. Azevedo-Santos’*?

Universidade Estadual de Maringd. Av. Colombo, 5790, 87020-900, Maringa, Paranda, Brazil

Unidad Ejecutora Lillo (CONICET)-Fundacion Miguel Lillo. Miguel Lillo 251, San Miguel de Tucuman (CP 4000), Tucumadn, Argentina
Departamento de Zoologia, Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil

Departamento de Ictiologia, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru

Escuela de Ingenieria y Gestion Ambiental, Facultad de Ingenieria y Gestion, Universidad Nacional Auténoma de Huanta, Huanta, Peru
Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil

Faculdade Eduvale de Avaré, Avaré, SGo Paulo, Brazil

Nucleo de Ecologia Aquatica e Pesca da Amazénia, Grupo de Ecologia Aquatica, Universidade Federal do Para, Belém, Para, Brazil

O ON DO FW DY

Programa de Pos-Graduagdao em Biodiversidade, Ecologia e Conservagdao, Universidade Federal do Tocantins, Porto Nacional, Tocantins, Brazil
https://zoobank. org/78 BD2D17-6B34-46D6-8 163-85C87B9652BF

Corresponding author: Valter M. Azevedo-Santos (valter.ecologia@gmail.com)

Academic editor: Nicolas Hubert # Received 23 June 2022 # Accepted 17 August 2022 Published 8 September 2022

Abstract

A new diagnosis and a new classification of Heptapterus are provided and a new species, H. carmelitanorum, is described.
Heptapterus is diagnosed by the following character combination: adipose fin confluent with the caudal fin; non-bifurcate caudal
fin; anal-fin insertion posterior to vertical through adipose-fin insertion; 10—23 anal-fin rays; anal fin not confluent with caudal fin;
and extremely elongate body, with a head length of 16.1-24.9%SL. Species included in Heptapterus are H. borodini, H. carmeli-
tanorum, H. carnatus, H. exilis, H. hollandi, H. mandimbusu, H. mbya, H. mustelinus, H. ornaticeps, and H. gengo. Some of the
character states diagnosing H. carmelitanorum among its congeners are the anal-fin insertion less than one eye diameter posterior
to a vertical through the adipose-fin insertion (vs. more than one eye diameter in all congeners); the isognathous mouth (vs. slight-
ly to moderately retrognathous, except H. borodini); and the keel formed by ventral procurrent caudal-fin rays shallow, far from
reaching anal-fin base (vs. keel formed by ventral procurrent caudal-fin rays deep, continuing almost to the anal-fin base, except in
H. borodini and H. hollandi).

Key Words

‘Chasmocranus’ brachynema, Grande River basin, Heptapterus mustelinus, Imparfinis borodini, Imparfinis hollandi, Minas Gerais,
Pariolius, Sapucai River basin, Siluriformes

Introduction there are an additional 1,120 species to be described in

this region (Ota et al. 2015). This hidden diversity is ex-
Siluriformes is one of the most species-rich actinopte- pected to be discovered within siluriform families that
rygian orders, with about 4,100 valid species (Fricke et already are species rich, but that have been subject to
al. 2022). Half of these species occur in the Neotropics _ little taxonomic effort. For example, the Neotropics en-
(~2,050) although diversity projections indicate that demic Heptapteridae has 231 valid species, is especially

Copyright Depra, G.d.C. et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use,
distribution, and reproduction in any medium, provided the original author and source are credited.

326 Depra, G.d.C. et al.: Redefinition of Heptapterus and description of a new species

common in low-order streams, and has received compar-
atively little recent attention from taxonomists (Fricke et
al. 2022). Notably, only 8.2% of these (19 species) were
described in the past ten years, compared to 22.2% of Lo-
ricariidae and 29.2% of Trichomycteridae (Fricke et al.
2022). This is despite both families also being endemic
to the Neotropics and frequently found in the same envi-
ronments as heptapterids.

Advancements in the alpha taxonomy of Heptapteri-
dae have been hampered by shortfalls in the classification
of those fishes. As presently understood, some heptapter-
id genera are highly heterogeneous assemblages resulting
from unjustified redefinitions and synonymies proposed
during the 20" century. Part of that heterogeneity results
from the inference that previously proposed generic char-
acters were insufficient to warrant distinction between
genera (e.g., Haseman 1911; Miranda-Ribeiro 1911;
Gosline 1941; Inger 1956; Mees 1967; Mees 1974; Mees
and Cala 1989). However, it can also be explained by the
fact that several of earlier species descriptions contained
errors regarding important diagnostic characters at spe-
cific and generic levels. For instance, Chasmocranus su-
rinamensis (Bleeker 1862) was described in Heptapterus
Bleeker, 1858, because the author thought it had the ad-
ipose fin continuous with the caudal fin, but it does not
(Inger 1956). Eigenmann and Norris (1900) diagnosed
Imparfinis Eigenmann & Norris, 1900, by having the eye
without free margin and vomerine teeth present, but the
type species /. piperatus Eigenmann & Norris, 1900, has
a free orbital margin and no vomerine teeth (Mees 1974);
and Borodin (1927) described Jmparfinis longicauda
Borodin, 1927 (= ‘I.’ borodini Mees & Cala, 1989) in
Imparfinis because that species has no free orbital mar-
gin and he understood that the adipose fin was not con-
fluent with the caudal. Through that series of misguided
taxonomic actions, /mparfinis came to include species
with lateral eyes with free rim, short body, long barbels,
and bifurcate caudal fin, not confluent with the adipose,
but also ‘J.’ borodini, a species with dorsal eyes without
free rim, elongate body, short barbels, and non-bifurcate
caudal fin, confluent with the adipose. Likewise, ‘/.’ hol-
landi Haseman, 1911, which shares with ‘J.’ borodini all
those character states in addition to other non-diagnostic
characters, was placed in Pariolius Cope, 1872, by Gos-
line (1941), and subsequently in Heptapterus by Mees
(1974), who synonymized the latter two genera. In con-
trast, Bockmann and Guazzelli (2003), in the most recent
complete classification of Heptapteridae, listed ‘J.’ hol-
landi and ‘I.’ borodini in their original genus /mparfinis,
even though both species fit an early definition of Hep-
tapterus by Bleeker (1864), except for a slightly different
position of the anal fin, relative to the adipose. When it
seemed that the confusion could not be further compli-
cated, Baumgartner et al. (2012) placed ‘/.’ hollandi in
Pariolius, following Gosline (1941), although Pariolius
(currently valid) includes only one very small, relatively
short-bodied species from western Amazon (Bockmann
and Slobodian 2017).

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What, then, can phylogeny say about the matter, if
pre-cladistic classificatory schemes seem to have failed?
A phylogenetic study (Silva et al. 2021) has recovered
both ‘1’ borodini and ‘I.’ hollandi as closely related to
H. mustelinus (Valenciennes, 1835), the type species of
Heptapterus. Apparently, the sole reason why they are
not currently assigned to Heptapterus is that they are
thought to belong to an undescribed genus, differing from
Heptapterus by a few characters (‘new genus D’ of Bock-
mann and Slobodian 2017). Recently, an undescribed
heptapterid species belonging to that lineage was collect-
ed in a small stream of the Grande River basin, in the
Upper Parana ecoregion (Azevedo-Santos et al. 2019).
Meanwhile, the re-examination of specimens deposited
in fish collections, as well as the sampling of previously
unknown populations, has been revealing other new spe-
cies belonging to that lineage — and their description can-
not wait for the description of the new genus. Until then,
those species must be described in the genus whose defi-
nition best matches the observed combinations of charac-
ters. As we will argue, in the case of the new species de-
scribed herein the most appropriate genus 1s Heptapterus.

Heptapterus has also been the subject of consider-
able taxonomic confusion. The type species, H. mus-
telinus, 1s most similar phenotypically to H. carnatus
Faustino-Fuster, Bockmann & Malabarba, 2019, H. exilis
Faustino-Fuster, Bockmann & Malabarba, 2019, H. man-
dimbusu Aguilera, Benitez, Teran, Alonso & Muirande,
2017, H. mbya Azpelicueta, Aguilera & Muirande, 2011,
H. ornaticeps Ahl, 1936, and H. gengo Aguilera, Miran-
de & Azpelicueta, 2011. All those species, except H. or-
naticeps, were described from the southern extreme of the
geographic range of Heptapteridae (Lower Parana, Sali
and Uruguay River basins, in Argentina and southern Bra-
zil) (Bleeker 1864; Ahl 1936; Aguilera et al. 2011, 2017;
Azpelicueta et al. 2011; Faustino-Fuster et al. 2019). In
addition, the type locality of H. ornaticeps, “Rio de Ja-
neiro”, is inaccurate (Bockmann and Guazzelli 2003). Be-
yond the aforementioned species and based on the defini-
tion of Heptapterus by Bockmann and Slobodian (2017),
the most recent complete classification of Heptapterus
by Faustino-Fuster et al. (2019) includes H. sympteryg-
ium Buckup, 1988, described from coastal drainages in
Rio Grande do Sul. However, prior to the publication of
those studies, the genus also included H. bleekeri Boese-
man, 1953 (= Chasmocranus bleekeri), H. fissipinnis Mi-
randa Ribeiro, 1911 (= Acentronichthys fissipinnis), ‘H.’
multiradiatus Thering, 1907, *H.’ stewarti Haseman, 1911,
and H. tapanahoniensis Mees, 1967 (= Chasmocranus
tapanahoniensis) (Bockmann and Guazzelli 2003, altered
by Bockmann and de Pinna 2004; DoNascimiento and
Milani 2008). All of those diverge from H. mustelinus by
one or more easily observable characters, and either can
be assigned to other valid genera or regarded simply as
incertae sedis within Heptapterini.

All that considered, a new diagnosis of Heptapter-
us was necessary, based on that provided by Bleeker
(1864), but with a few alterations — to reflect available

Zoosyst. Evol. 98 (2) 2022, 327-343

phylogenetic evidence and make the recognition of the
genus more straightforward. Thus, we propose the new
diagnosis for Heptapterus, present a list of the species
which should be regarded as belonging to Heptapterus,
and the description of a new species from the Grande
River basin in southeastern Brazil.

Material and methods

Morphological data

Measurements and counts were taken as in Faustino-Fus-
ter et al. (2019), with some alterations, as follows. Body
depth was measured both at the dorsal- and at the adi-
pose-fin origins. Body width was measured at dorsal-fin
origin and at cleithrum (cleithral width). Both bony and
fleshy interorbital distances were measured. Dorsal-, pec-
toral-, and pelvic-fin lengths were replaced by the lengths
of the first three rays of each fin. Also, the length of the
last dorsal-fin ray was added. Other additions include cau-
dal-fin depth; distance between snout tip and terminus of
dorsal-fin base; distance between snout tip and dorsal-fin
distal end, adpressed; length of stiffened part of first dor-
sal-fin ray; distance between snout tip and terminus of
pectoral-fin base; distance between snout tip and pecto-
ral-fin distal end, adpressed; length of stiffened part of
first pectoral-fin ray; distance between snout tip and ter-
minus of pelvic-fin base; distance between snout tip and
pelvic-fin distal end, adpressed; distance between pelvic
fins; distance between snout tip and terminus of anal-fin
base; first branched anal-fin ray length; distance between
snout tip and anal-fin distal end, adpressed; distance be-
tween snout tip and adipose-fin base end: snout-anus dis-
tance; snout-urogenital papilla distance; anus-urogenital
papilla distance; dorsal lobe of caudal fin length; ventral
lobe of caudal fin length; distance between snout tip and
posterior nare; anterior internarial width; and posterior
internarial width. The following measurements were ex-
cluded: dorsal-, pelvic-, and anal-fin insertion to hypural
plate; mandibular isthmus to lower and upper lips; postor-
bital distance; snout to anterior nostril distance; head
depth at interorbital; and head width at posterior nostril.

laterodorsal stripe

DB1

interorbital

pre-orbital stripe humeral mark

329

According to the relative position of the premaxil-
la and of the dentary, the mouth is classified in one of
the following categories: prognathous, when the dentary
projects anteriorly to the premaxilla; isognathous, when
the premaxilla and the dentary reach the same vertical an-
teriorly; and retrognathous, when the premaxilla projects
anteriorly to the dentary. Cephalic laterosensory canal
terminology follows Bockmann and Miquelarena (2008).
We propose a new terminology for the melanophoric col-
oration elements which is based on the examination of
the new species and comparative Heptapterini, mainly
Imparfinis minutus (Litken, 1874) and Rhamdioglanis
frenatus Ihering, 1907, which have the most developed
marks within the subfamily (Fig. 1). Dorsal bars (DBs)
are discrete transversal marks present along the dorsum,
from the top of the neurocranium to the end of the caudal
peduncle. The usual positions of those marks, in relation
to specific body structures, are as follows: DB8 is locat-
ed on the caudal peduncle, approximately at the base of
the anteriormost unbranched caudal-fin rays; DB7, at the
terminus of the adipose-fin base or slightly anterior to it;
DB6, approximately at adipose-fin insertion; DB5, mid-
way between dorsal and adipose fins; DB4, at dorsal-fin
base terminus; DB3, immediately anterior to dorsal-fin
insertion; DB2, slightly posterior to supraoccipital; DB1,
on supraoccipital and posterior portion of frontal. An in-
terorbital bar is a transverse mark between the eyes. A
pre-orbital stripe runs from the base of the maxillary bar-
bel or immediately behind to the anteroventral margin of
the eye. A humeral spot is located immediately posterior
to the pectoral girdle and either ventral to or crossing the
lateral line. A midlateral stripe runs along the lateral line,
and may extend from the humeral spot or behind, to the
caudal-fin base. A laterodorsal stripe may unite the lat-
eral extremities of DBs 2—5. A caudal spot is located at
the base of the upper caudal-fin lobe. Comparisons were
undertaken directly through examination of specimens,
including types, and original descriptions of valid species
of Heptapterus. The term ‘allomery’ is used in the same
sense as in Depra et al. (2021). For counts, we considered
all free vertebrae; the compound caudal centrum was ac-
counted as a single element and those in Weberian com-
plex were excluded.

caudal spot

ey midlateral stripe

Figure 1. Schematic representation of melanophoric coloration elements that may be present in Heptapterini species, based on the
new species described herein and comparative members of the same tribe (mainly /mparfinis minutus and Rhamdioglanis frenatus,

in which most of the elements are present).

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330 Depra, G.d.C. et al.: Redefinition of Heptapterus and description of a new species

The map with the species distribution was modified
from Azevedo-Santos et al. (2019) using the Software QGis
(Sherman et al. 2012). Biological collections where type
Specimens were deposited are LBP (Laboratorio de Biolo-
gia e Genética de Peixes, UNESP, Botucatu, Brazil) and
MNRJ (Museu Nacional, UFRJ, Rio de Janeiro, Brazil).
Definitions of all other institutional abbreviations men-
tioned in this study may be found in Sabaj (2020) — with
the exception of CICCAA (for this see Aguiar et al. 2022).

Results

Taxonomic accounts

Genus Heptapterus Bleeker, 1858

Type species. Pimelodus mustelinus Valenciennes, 1835.

Diagnosis. Heptapterus differs from all other Heptapter-
ini except Acentronichthys Eigenmann & Eigenmann,
1889, Nemuroglanis Eigenmann & Eigenmann, 1889,
Chasmocranus bleekeri, “Chasmocranus’ brachynema
Gomes & Schubart, 1958, ‘Heptapterus’ multiradiatus,
‘H.’ stewarti, and ‘H.’ sympterygium by the presence of an
adipose fin extensively fused with the caudal fin (Fig. 2).
Heptapterus is distinguished from Acentronichthys by
having a non-bifurcate caudal fin (1.e., caudal fin not di-
vided in two lobes; vs. bifurcate, with distinct dorsal and
ventral lobes), and from Nemuroglanis by having dark
bars and stripes on back of trunk (vs. absence of dark bars

adipose fin

(a)
anterior caudal-fin rays

(b)

pee ae

Figure 2. Schematic representation of the different degrees of
proximity and connection between the adipose and caudal fins
in Heptapterini. a. Adipose and caudal fins widely separate, as
in Imparfinis piperatus, for instance; b. Adipose fin reaching
the caudal fin, but not connecting to it (1.e., connective tissue
in which dorsal procurrent caudal-fin rays are imbedded is not
contiguous with connective tissue forming the adipose fin), as
in Chasmocranus longior, for instance; ¢. Adipose fin connect-
ing with caudal fin (i.e., connective tissue in which dorsal pro-
current caudal-fin rays are imbedded 1s clearly contiguous with
connective tissue forming the adipose fin), as in Heptapterus.

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and stripes on back of trunk) and 5-6 pairs of pleural ribs
(vs. 8—9 ribs). It is also distinguished from ‘C.’ brachyne-
ma by having an elongate body, with a head length of
16.1—24.9% (vs. 25.5—27.3%); the posterior extension of
mouth rim much shorter, with rictus barely reaching ver-
tical line through posterior nostril (vs. posterior extension
of mouth rim much longer, with rictus reaching vertical
line between posterior nostril and eye); and the premaxil-
lary tooth plate with no posterolateral extension, or with a
small one (vs. with a very long posterolateral extension). It
is further distinguished from C. bleekeri by having the pel-
vic-fin insertion posterior to vertical through insertion of
dorsal fin (vs. anterior) and anal-fin insertion posterior to
vertical through adipose-fin origin (vs. anterior). Addition-
ally, Heptapterus differs from ‘Heptapterus’ multiradiatus
and ‘H.’ stewarti by having fewer anal-fin rays (10-23 in
Heptapterus vs. 38-46 and 33-36 in ‘H.’ multiradiatus
and ‘H.’ stewarti, respectively). It is further distinguished
from ‘H.’ sympterygium by having the anal and caudal fins
separated (vs. anal fin confluent with the caudal fin), and
supraorbital pore 6 (s6) fused or closer to each other (vs.
separate and closer to the eye than to each other).

Species included. Heptapterus borodini (Mees &
Cala, 1989), H. carmelitanorum, H. carnatus Fausti-
no-Fuster, Bockmann & Malabarba, 2019, H. exilis Faus-
tino-Fuster, Bockmann & Malabarba, 2019, H. hollandi
(Haseman, 1911), H. mandimbusu Aguilera, Benitez,
Teran, Alonso & Mirande, 2017, H. mbya Azpelicueta,
Aguilera & Mirande, 2011, H. mustelinus (Valenciennes,
1835), H. ornaticeps Ahl, 1936, and H. genqo Aguilera,
Mirande & Azpelicueta, 2011.

Heptapterus carmelitanorum sp. nov., Azevedo-
Santos, Depra, Aguilera, Faustino-Fuster & Katz
https://zoobank.org/995EA984-A 1 D8-4A4E-8224-7D5D9637AFBF
Figs 3, 4; Suppl. material 1: Figs S1-S4; Table 1

‘Heptapterus’ sp.: - Azevedo-Santos et al. (2019) (listed in a survey).

Holotype. MNRJ 53174, 144.3 mm SL; Brazil: Minas
Gerais State: limit of Carmo do Rio Claro and Ilicinea
municipalities: Unknown named stream tributary of
Itaci stream, tributary of Sapucai River (stretch flood-
ed by Furnas reservoir), Grande River Drainage, Parana
River basin, ~20°54'57"S, 45°56'21"W, altitude about
830 m asl; A. M. Katz and V. M. Azevedo-Santos, 31
October 2021.

Paratypes. LBP 26570, 1, 95.7 mm SL; same locali-
ty as holotype; V. M. Azevedo-Santos and P. N. Coelho,
22 July 2017; LBP 26575, 1, 89.1 mm SL, same locality
as holotype; V. M. Azevedo-Santos and P. N. Coelho, 25
May 2018; LBP 23577, 1, 104.4 mm SL, same locality
as holotype; V. M. Azevedo-Santos and P. N. Coelho, 10
April 2017.

Diagnosis. Heptapterus carmelitanorum differs from
all congeners by possessing the anal-fin insertion less
than one eye diameter posterior to a vertical through the

Zoosyst. Evol. 98 (2) 2022, 327-343

(a)

331

Figure 3. Heptapterus carmelitanorum, new species, holotype, MNRJ 53174, 144.3 mm SL; a. Dorsal view; b. Lateral view;

c. Ventral view.

Figure 4. Heptapterus carmelitanorum, new species, holotype, coloration in life.

adipose-fin insertion (vs. more than one eye diameter pos-
terior). From all congeners, except H. borodini, by an isog-
nathous mouth (vs. slightly to moderately retrognathous).
It differs from all other congeners except H. borodini and
H. hollandi, by the keel formed by ventral procurrent cau-
dal-fin rays shallow, far from reaching anal-fin base (vs.
keel formed by ventral procurrent caudal-fin rays deep,
continuing almost to the anal-fin base, even though its
anterior portion is devoid of fin rays) (Fig. 5). It differs
from both H. borodini and H. hollandi by having an al-
most elliptical caudal fin (vs. lanceolate in H. borodini,
obliquely truncate to falcate in H. hollandi;, Fig. 6), the
length of its dorsal lobe 18.3-19.3% SL (vs. 24.4-43.3%

SL in H. borodini). Additionally, H. carmelitanorum
differs from all other congeners, except H. carnatus, H.
mbya, H. qenqo, and some specimens of H. hollandi, by
having inconspicuous dorsal bars (vs. conspicuous). From
H. borodini, H. carnatus, H. exilis, H. hollandi, H. mus-
telinus, and H. ornaticeps, by having 14—15 anal-fin rays
(vs. 10-12 in H. borodini and H. hollandi;, 18-21 in H.
carnatus, 16-19 in H. exilis, 18—23 in H. mustelinus; and
19 in H. ornaticeps). Differs from H. exilis by the com-
plete lateral line (in adults), continuous to base of hypural
plate (vs. incomplete, not reaching dorsal-fin insertion).
Heptapterus carmelitanorum further differs from H. hol-
landi by having 1,6 dorsal-fin rays (vs. 1,7).

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332

Description. General morphology (Figs 3—4, 7; Sup-
pl. material 1: Figs S1—S3). Available specimens (ho-
lotype and three paratypes) ranging from 89.1—144.3
mm SL; morphometric data in Table 1. General shape
of body presented in photographs of preserved and live
specimens. Dorsal profile convex from premaxillary
symphysis to end of dorsal-fin base; slightly convex
from that point to adipose-fin insertion; slightly convex
along adipose-fin base. Caudal-fin base rounded. Ven-

Depra, G.d.C. et al.: Redefinition of Heptapterus and description of a new species

tral profile convex from dentary symphysis to isthmus;
straight or slightly convex from that point to anal open-
ing; straight along anal-fin base; concave from its end to
caudal-fin base. In dorsal view, mouth rim gently arched,
convex; lateral profile of head convex due to well-de-
veloped adductor mandibulae muscle; lateral profile of
body straight to slightly convex along abdomen, tapering
gently to about half adipose-fin base, then tapering more
abruptly to caudal-fin base.

Table 1. Morphometric data of the type specimens of Heptapterus carmelitanorum sp. nov.

Paratype Paratype Paratype Holotype X SD
(LBP 23577) (LBP 26575) (LBP 26570) (MNRJ 53174)
Total length Neil ee) 106.6 114.1 144.3 121.6 16.296
Standard length 104.4 89.1 95.7 1204 102.5 13.670
As percentages of SL
Body depth at dorsalfin origin 12.5 9.8 9.6 11.5 10.8 0.014
Body depth at adipose-fin origin 12.3 11.0 10.6 11.1 11.2 0.007
Caudal-fin depth 14.1 12.5 14.4 10.6 12.9 0.017
Body width at dorsal-fin origin 12.5 11.4 11.4 11.7 MZ 0005
Cleithral width 15:8 14.9 15.4 16.9 1527 ~ £0:008
Head length to base of supra-occipital process 17.8 18.9 19.2 18.7 18.7. 0.006
Lateral head length (to posteriormost point of opercle) 20.5 20.5 20.6 21.3 20.7. 0.004
Maxillary-barbel length 82.2 78.1 92.4 724 81.3 0.084
Outer mental-barbel length 44.4 44.3 52.8 45.1 46.6 0.041
Inner mental-barbel length 29.9 29.0 27.9 31.1 29.5 0.014
Predorsal length 37.8 39.6 38.9 39.4 38.9 0.008
Distance between snout tip and terminus of dorsal-fin base 49.8 51.5 50.7 51.2 50.8 0.007
Distance between snout tip and dorsalfin distal end, adpressed 58.4 60.7 59.8 60.3 59.8 0.010
Dorsal fin to adipose fin 15.7 15.6 16.0 14.8 15.5 0.005
Dorsalfin base 11.8 11.6 11.1 11.4 11.5 0.003
Length of first dorsal-fin ray (unbranched) 9.5 10.2 10.9 9.9 10.1 0.006
Length of stiffened part of first dorsal-fin ray 3.3 4.5 4.9 4.1 4.2 0.007
Length of second dorsal-fin ray (first branched) 12.4 12.9 14.0 OS 12.4 0.015
Length of third dorsaLfin ray (second branched) 13.0 13.4 14.1 12.8 13.3 0.006
Length of last dorsalfin ray 9.0 9.1 10.0 9.4 9.4 0.005
Prepectoral length 20.1 20.0 19.5 20.0 19.9 0.002
Distance between snout tip and terminus of pectoral-fin base 22.8 22.1 23.0 23.0 22.7 0.004
Distance between snout tip and pectoralfin distal end, adpressed 31.9 32.2 33.4 32.7 32.6 0.007
Length of first pectoralfin ray (unbranched) 8.7 9.3 9.1 7.9 8.7 0.006
Length of stiffened part of first pectoraLfin ray 3.0 4.0 aul a3 3.4 0.005
Length of second pectoral-fin ray (first branched) 10.2 11.0 10.9 9.8 10.4 0.006
Length of third pectoralfin ray (second branched) 10.9 11.9 11.1 10.4 11.1 0.006
Pectoral to pelvic-fin distance 20.3 21.9 20.5 20.5 20.8 0.007
Prepelvic length 38.4 39.5 38.9 39.9 39.2 0.007
Distance between snout tip and terminus of pelvic-fin base 39.9 42.1 42.0 41.9 41.5 0.010
Distance between snout tip and pelvic-fin distal end, adpressed 50.9 54.2 54.8 54.3 53.5 0.018
Distance between pelvic fins 6.5 5.8 5.9 6.1 6.1 0.003
Length of first pelvic-fin ray (unbranched) 9.2 9.8 9.9 7.8 9.2 0.010
Length of second pelvic-fin ray (first branched) 10.8 10.1 12.0 12.6 11.4 0.011
Length of third pelvic-fin ray (second branched) 11.6 bai, 13.5 12.2 12.3 0.008
Pelvic to ana-fin distance 27.0 28.6 28.9 28.3 28.2 0.008
Anal-fin base 17.6 16.4 16.1 16.6 L627 0-007
Preanal length 64.9 69.6 68.7 68.7 68.0 0.021
Distance between snout tip and terminus of anal-fin base 84.1 84.7 84.1 85.0 84.5 0.005
First branched analfin ray length viel 6.4 Ji? O35 6.8 0.004
Distance between snout tip and anal-fin distal end, adpressed 90.5 90.8 91.0 91.7 91.0 0.005
Adipose-fin length 28.7 27.5 29.2 28.5 28.5 0.007
Preadipose length 65.3 67.0 66.0 66.6 66.2 0.007
Distance between snout tip and adipose-fin base end 93.3 94.6 96.7 95.4 95.0 0.014
Adipose-fin depth 2.7 22 2 1.8 2:3 0.004
Caudal-peduncle length 16.5 16.4 16.4 16.3 16.4 0.001
Caudal-peduncle depth at adipose-fin terminus 8.7 8.3 8.6 8.5 8.5 0.002
Snout-anus distance 44.4 46.8 46.0 45.8 45.8 0.010
Snout-urogenital papilla distance 47.9 49.9 48.4 49.7 49.0 0.010
Anus-urogenital papilla distance oe) 3:0 2.9 3.4 32 0.002
Dorsal lobe of caudal fin length 18.9 18.3 19.1 19.3 18.9 0.004
Ventral lobe of caudal fin length 15.2 15.4 16.2 16.6 15.8 0.006

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Zoosyst. Evol. 98 (2) 2022, 327-343

333

Paratype Paratype Paratype Holotype X SD
(LBP 23577) (LBP 26575) (LBP 26570) (MNRJ 53174)
As percentages of HL (lateral)
Head depth 46.3 41.5 43.1 43.6 43.6 0.020
Head width 76.6 Fost 74.6 78.6 75.6 0.026
Eye diameter 15.9 15:3 152 14.8 153 = «8005
Fleshy interorbital 15.9 - 16.2 19.1 LT) 0.01%
Bony interorbital 9.8 F135 10.2 10.1 10.4 0.007
Mouth gape 41.1 40.4 39.6 41.6 40.7. 0.009
Snout length 33:2 83.9 34.5 xX Hl 33.7. 0.007
Distance between snout tip and posterior nare 22.0 22.4 223 23:3 22.5 0.006
Distance between posterior nostril and eye 8.9 hd 9.1 ib; 9.2 0.015
Anterior internarial width 23.4 Boel, 19.3 21.0 22.2 0:026
Posterior internarial width 20.1 21.9 19.8 20.2 20.5 0.009
Intranarial length 22.4 20.2 21:3 24.9 eoa2? 9.020

(a)

(b)

keel

Figure 5. Schematic representation of the different degrees of
proximity between the anal and caudal fins in Heptapterus. a. Keel
formed by rigid connective tissue with imbedded ventral procur-
rent caudal-fin rays not much developed, its anterior end distant
from anal-fin base (Heptapterus borodini, H. carmelitanorum and
H. hollandi), b. Keel well developed, its anterior end reaching
or almost reaching anal-fin base (remaining Heptapterus species).

Head much depressed, flat dorsally and ventrally,
rounded laterally. Mouth isognathous. Mouth rictus
fleshy, folding ventrally, with large sub-labial groove be-
neath it (Fig. 7a). Lips double, 1.e., divided by deep la-
bial slit into outer and inner lip (Fig. 7b). Outer dorsal
lip thickly and abundantly plicate; outer lower lip thickly,
but scarcely plicate; inner dorsal and ventral lips finely
and abundantly plicate (Fig. 7b). Tubular anterior nos-
tril not reaching mouth rim. Deep skin fold surrounding
entire posterior nostril, but with deep posterior notch
(Fig. 7c). Maxillary barbel groove extending from base
of barbel almost to the eye; in dorsal view, rim of groove
almost parallel with body axis. Dorsal surface of snout
with shallow depression posteriorly to posterior nostril,
and elongate depression marking anterior cranial fonta-
nel (Fig. 7b). Bulging eyes covered in thick skin with no
free rim, almost completely dorsal. Base of inner mental
barbel slightly anterior to that of outer mental barbel, dis-
tinctly posterior to base of maxillary barbel. Maxillary
barbel reaching anterior margin of first pectoral-fin ray.
Shallow cleithral skin fold immediately posterior to bran-
chial aperture, posterior terminus medial to base of first
pectoral-fin ray (Fig. 7a). Abdominal region depressed,
distinctly broader than deep; in cross section, some-
thing between elliptic and rectangular. Cross section at

Figure 6. Different caudal-fin
a. H. borodini, NUP 14882, 74.3 mm; b. H. hollandi, young;
c. H. hollandi, adult, NUP 5978, 199.1 mm SL.

shapes in Heptapterus.

dorsal-fin base approximately as broad as deep, between
round and square. Body compressed from adipose-fin 1n-
sertion to caudal fin, cross-section distinctly deeper than
broad. Vertebrae 43. Ribs 9 (Suppl. material 1: Fig. S4).
Dorsal fin distal margin convex; 1,6*(4) rays (first ray
rigid only basally); each branched ray with, at least, tertia-
ry branches; thin membrane between rays. Pelvic-fin in-
sertion at same vertical as base of second (first branched)
dorsal-fin ray (2 specimens) or between bases of first and
second rays (2*). Adipose fin continuous (i.e., connected)
with the anteriormost ray of dorsal portion of caudal fin,
originating slightly anteriorly to vertical through anal-fin
insertion (distance less than one eye diameter); margin

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334 Depra, G.d.C. et al.: Redefinition of Heptapterus and description of a new species

Figure 7. Superficial structures in Heptapterus carmelitano-
rum, a. Sub-labial groove (blue arrowhead) and cleithral skin-
fold (black arrowhead); b. Labial slit (blue arrowheads) and
plicae on the outer (lemon arrowheads) and inner (pink arrow-
heads) lips; c. Posterior nostril, evidencing shape of posterior
notch (red arrowhead).

slightly convex. Caudal fin approximately elliptical, rays
of dorsal half little longer than ventral ones; xi1,8,8,x1*(1)
xv,7,8,xv(1), xvil,6,7,x1V(1), xvil,6,7,xvi(1) rays (Suppl.
material 1: Fig. S5); thin membrane between rays. Pec-
toral fin approximately elliptical, with anterior rays lon-
ger than posterior ones; i,7,1(2), 1,8*(2) rays on left side
(first ray rigid only basally); on right side, 1,7,1*(4); each
branched ray with, at least, tertiary branches; thin mem-
brane between rays. Pelvic fin approximately elliptical,
with anterior rays longer than posterior ones; 1,5 (4) rays

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on both sides; each branched ray with, at least, tertiary
branches; thin membrane between rays.

Premaxillary toothplate about twice as wide as long,
length of lateral margin slightly higher than symphyseal
margin; small posterolateral projection present; about six
rows of conical teeth (tooth plate virtually identical to the
one in Mees 1967, fig. Ic). External gill rakers on first
arch 1+6*(3), 1+7(1). Branchiostegal rays 8(2) (Suppl.
material 1: Fig. S5).

Laterosensory system. Cephalic laterosensory pores as
Bockmann and Miquelarena (2008) described for Rham-
della cainguae Bockmann & Miquelarena, 2008, except
in following details (Fig. 8): s2+12 pore much closer to
anterior nostril (vs. at about middle of the distance be-
tween anterior and posterior nostrils); s4 pore distinctly
more medial than s3 pore (vs. slightly more medial); s8
with two pores (s8a and s8p; vs. s8 with one pore); po3
with two pores (po3a and po3p; vs. po3 with one pore):
pm1 pore only slightly posterior to transversal line across
pm2 pore (vs. much posterior to it); pm1 directed me-
dially, facing antimere (vs. directed ventrally); pm2 and
pm3 pores facing anteroventrally (vs. posteroventrally
and ventrally, respectively); pm4 and pm5 pores antero-
medial to rictus (vs. posteromedial and posterior to it, re-
spectively); pm10 pore slightly closer to pol+pm11 pore
than to pm9 pore (vs. much closer to pm9 pore). Eye also
more distant from 15, 16, s6, s7, and s8 pores than in R.
cainguae, seemingly due to anterior displacement of eye
in Heptapterus carmelitanorum. Lateral line continuous
to hypural plate, with 43(1), 46(1), 63(1) pores, or ending
on hypural plate, but with large gap between anterior and
posterior portions, with 23(1) total pores (smallest speci-
men, LBP 26575).

Olfactory organ. One specimen (LBP 23577) dissect-
ed with two longitudinal series of flat, triangular lamel-
lae on right olfactory canal, each series with 32 lamellae
(Fig. 9).

Epidermal papillae. In LBP 23577, external surface of
body covered with densely packed, flexible, perpendicu-
larly protruding epidermal papillae (except lips; distal half
of barbels, tubular portion of anterior nostril and skin flap
of posterior nostril; center of eye; distal margin of bran-
chiostegal membrane; and nearly entire fins). Distance
between adjacent papillae ~0.15 mm, equal to their max-
imum length. Papillae slender, rod-like on most of body
(Fig. 10a, b); short, club-like, apparently with widened dis-
tal extremity on ventral surface of head (Fig. 10c; widened
portion possibly attached mucus). Very small papillae on
anterior face of first pectoral- and pelvic-fin ray; on base
of caudal-fin rays; on margin of eye; on base of tubular
portion of anterior nostril; on base of skin flap of posterior
nostril; on ventral half of adipose fin. Scarce, but well-de-
veloped papillae on urogenital papilla and anus. All epi-
dermal papillae visible only after removal of body mucus.

Color in alcohol (Fig. 3, Suppl. material 1: Figs S1,
S2). Background color greyish-brown, grading to white
towards belly and to white beige towards region between
anus and anal fin, and ventral side of head; transition

Zoosyst. Evol. 98 (2) 2022, 327-343

335

pol+pmil
a

Pe

poit+pmii |

po3a 112

po3p |I1
tA

po2
|

Figure 8. Cephalic laterosensory system of Heptapterus carmelitanorum, new species (based on LBP 23577); abbreviations as in

Bockmann and Miquelarena (2008).

Figure 9. Heptapterus carmelitanorum, LBP 23577, dissected
to show the olfactory organ (anterior side to the right). Some of
the lamellae in the outer (red arrowhead) and inner (green arrow-
head) series are outlined in blue to better evidence their shape.

between brown and light beige more abrupt on head than
in remainder of body. Caudal spot very faint, small, at base
of dorsalmost branched caudal-fin ray; DB8 and 7 absent;
DB6 through 4 inconspicuous, dark-brown (respectively,
at adipose-fin insertion; midway between dorsal and ad-
ipose fins; and terminus of dorsal-fin base); DB3 present
as roundish dark-brown spot immediately anterior to dor-
sal fin; DB2 very faint, little posterior to supraoccipital,
at vertical through posterior end of pectoral-fin base; DB1
dark brown, extending to opercle; interorbital bar indis-
tinct. Pre-orbital stripe very diffuse, dark-brown. Diffuse,
dark-brown humeral spot; faint midlateral stripe present
in LBP 26570 specimen; laterodorsal stripe absent.

Color in life (Fig. 4, Suppl. material 1: Fig. S3).
General pattern of body dark brown, yellowish in the
holotype (Fig. 4). Ventral region from isthmus to anal-
fin insertion paler than remainder of body and somewhat
pinkish, as well as cheek, branchiostegal membrane,
cleithrum and lateral line. All fin rays dark brown. Adi-
pose fin brownish yellow or dark yellowish brown. Inter-
radial membranes of pectoral, anal and caudal fins yellow.
Dorsal-fin interradial membrane hyaline, with scattered
melanophores on basal third. Barbels dark brown dorsally
and beige ventrally.

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336 Depra, G.d.C. et al.: Redefinition of Heptapterus and description of a new species

Figure 10. Epidermal papillae in Heptapterus carmelitanorum, LBP

- =

23577, paratype. a, b. Slender, rod-like papillae are distributed

on most of body, such as on the dorsum, between the head and dorsal fin (a) and on the head (b. arrow shows s6+s6 pore); c. Short,

club-like papillae are distributed on ventral surface of head.

Ontogeny. Strong positive allometry in cleithral width
(R? = 0.997), head length (0.742), fleshy interorbital dis-
tance (0.809), mouth width (0.633), and dorsal caudal-fin
lobe length (0.593; compare Fig. 3, Suppl. material 1:
Figs S1, $2); moderate positive allometry in ventral cau-
dal-fin lobe length (0.362); moderate negative allometry
in bony interorbital distance (0.392), maxillary-barbel
length (0.313), first dorsal-fin ray length (0.259), and
maximum adipose-fin height (0.317); strong negative
allometry in dorsal-adipose distance (0.656), first pecto-
ral-fin ray length (0.993), and first pelvic-fin ray length

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(0.918). Positive allomery present in the number of
branched rays in the dorsal caudal-fin lobe (R? = 0.5712)
and in the number of lateral-line pores (0.899).
Etymology. The specific name is a noun in apposition
derived from Carmelitanos (in Portuguese), the local ap-
pellation of people born or living in Carmo do Rio Claro
(Minas Gerais, Brazil), the city where the species was
discovered. The name is in honor of Carmelitanos, es-
pecially Ana Maria Vilela Soares, José Candido de Mel-
lo Carvalho, Moara Lemos, and Carlos Roberto Bueno
Junior, for their contributions to biological science.

Zoosyst. Evol. 98 (2) 2022, 327-343

Geographical distribution and ecological notes.
Heptapterus carmelitanorum is recorded only from a
single unnamed stream. The watercourse is a tributary of
Itaci stream — ribeirao Itaci, in Portuguese — which is an
affluent of Furnas reservoir (in the Sapucai River arm),
Grande River basin, in the upper Parana River system, in
Minas Gerais State, Brazil (Figs 11, 12).

The stream in which specimens of H. carmelitanorum
were collected has its source on a mountain known as
“Chapadao” (in Portuguese), approximately 1,300 me-
ters a.s.l. Its cannel crosses successive falls (forming
waterfalls), including one over 50 meters high. The type
locality lies downstream from the waterfalls. According
to the classification proposed by Strahler (1954), the
stream may be classified as third order. The water was
extremely clear (small characids readily observed) and
well oxygenated. The stream depth was shallow (not ex-
ceeding | meter), and its bed was completely formed by
rocks. Light penetration was low during samplings. In

SRY:

the reach, submerged tree roots and accumulated leaves
and fruits (especially Fabaceae) formed some microhab-
itats for some species, notably 7richomycterus candidus
(Miranda Ribeiro, 1949) and Cetopsorhamdia iheringi
Schubart & Gomes, 1959. The specimens of H. carmel-
itanorum were captured in environments that combined
rocks (generally juxtaposed) and a more turbulent flow
(see Fig. 12). Observation during sampling suggests that
the species 1s demersal.

Species collected with H. carmelitanorum include C.
iheringi, Hoplias malabaricus (Bloch, 1794), Knodus
moenkhausii (Eigenmann & Kennedy, 1903), Odontos-
tilbe weitzmani Chuctaya, Buhrnheim, & Malabarba,
2018, Oligosarcus argenteus Gunther, 1864, Pareiorhina
sp., Psalidodon sp., T: candidus, T: septemradiatus Katz,
Barbosa & Costa, 2013 (Azevedo-Santos et al. 2019). New
collections in the same reach resulted in the capture of ad-
ditional species, such as Apareiodon sp. (CICCAA06610)
and Rhamdiopsis sp. (CICCAA06611). In addition to

-20.900

Furnas Reservoir

-46.002

Furnas dam

Grande River

Sapucai River

45.951

Itaci stream

45.951 -45.900

Figure 11. Distribution of Heptapterus carmelitanorum.

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338 Depra, G.d.C. et al.: Redefinition of Heptapterus and description of a new species

Figure 12. Partial view (i.e., stretch) of the stream where the type specimens of Heptapterus carmelitanorum were sampled.

fishes, aquatic spiders (e.g., 7etragnatha sp.) and insects,
including specimens of the order Trichoptera in cases
formed by small gravels, were captured in the stretch.

Discussion

Definition of Heptapterus

We propose a new diagnosis for Heptapterus, aiming to
facilitate its recognition among members of Heptapterini,
based on external characters only. The new definition pur-
sued taxonomic stability by making the fewest possible al-
terations to the definition of Bleeker (1864), although that
means rejecting the definitions proposed by subsequent au-
thors, such as Gosline (1941) and Mees (1967). Our circum-
scription of Heptapterus also reflects that little knowledge
is currently available on Heptapterini phylogeny. Because
Silva et al. (2021) recovered ‘/mparfinis’ (= Heptapterus)
borodini as closely related to H. mustelinus, we opted pro-
visionally for including that species, along with the mor-
phologically similar H. carmelitanorum and H. hollandi in
Heptapterus [those species are considered by Bockmann
and Slobodian (2017) to form “new genus D”’].

About the fins of Heptapterus, Bleeker (1864:90) said:
“pinna dorsali anteriore ventralibus opposita” [dorsal fin

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longitudinally aligned with pelvic fin]; “adiposa elonga-
ta cum caudali unita, analis subelongata mediae adipo-
sae opposita” [adipose fin elongate, united with caudal
fin; anal fin subelongate, longitudinally aligned with the
middle of adipose fin]. He was vague about the degree
of elongation of the adipose fin in Heptapterus; the type
species, H. mustelinus, and most other congeners have an
adipose-fin base length greater than 40% SL (up to 58.5%
SL in H. mbya), which is a rare condition among Hep-
tapterini, found elsewhere only in Acentronichthys. How-
ever, H. mandimbusu, which is, in general terms, very
similar to the type species, may have a length as short
as 33.8% SL, which overlaps the values observed in H.
borodini (31.4—41.0% SL) and H. hollandi (29.1-35.1%
SL), and is only slightly higher than in H. carmelitano-
rum (27.4—29.1% SL). Thus, we had to adapt Bleeker’s
diagnosis to recognize that in Heptapterus the adipose-fin
base length presents a continuum of variation, ranging
from moderately to extremely elongate.

Bleeker (1864) was also unclear about the length of
the anal fin. Although H. mustelinus does have a large
number of anal-fin rays (18—23) lying about the middle
of the range observed in Heptapterini, other species pre-
viously included in Heptapterus, viz. H. mandimbusu,
H. mbya and H. gengo, may have lower numbers (respec-
tively, 14-18, 15-17 and 15-17). Heptapterus borodini,

Zoosyst. Evol. 98 (2) 2022, 327-343

H. carmelitanorum and H. hollandi have, respectively,
10-12, 14-15, and 10-12 anal-fin rays. The number of
anal-fin rays is useful to distinguish Heptapterus from the
incertae sedis *H.’ multiradiatus (38-46 rays), *H.’ stew-
arti (33-36) and ‘H.’ sympterygium (22-29). Our adapta-
tions of Bleeker’s (1864) diagnosis render the adipose-fin
base length and the number of anal-fin rays inadequate
to diagnose between Heptapterus and Chasmocranus
Eigenmann, 1912, a quite similar genus. That is because
C. bleekeri has an adipose-fin base length of up to about
35% (Mees, 1967), and the number of anal-fin rays in all
Chasmocranus species overlaps the lower portion of the
range observed in Heptapterus (which is 10—23). On the
other hand, the relative position between the adipose and
anal fins differs between the two genera. Some species,
such as H. mustelinus, do have the anal fin approximately
aligned with the center of adipose fin, as described by
Bleeker (1864). In other species, the anal-fin insertion
may be considerably closer to a vertical through the adi-
pose-fin insertion, such as in H. mandimbusu, H. borod-
ini, H. hollandi, and especially in H. carmelitanorum (in
which the anal-fin insertion is only slightly posterior to
adipose-fin insertion). This, too, makes our definition of
Heptapterus a little different from that of Bleeker (1864).
However, it is worth mentioning that this character state
is useful to distinguish Heptapterus from all species in-
cluded in Chasmocranus by Bockmann and Slobodian
(2017), and by us [viz. ‘C.’ brevior Eigenmann, 1912, C.
bleekeri Boeseman, 1953, C. chimantanus Inger, 1956,
C. longior Eigenmann, 1912, C. surinamensis (Bleeker,
1862) and C. tapanahoniensis|, all of which have the
anal-fin insertion anterior to a vertical through the adi-
pose-fin insertion. In fact, that character also diagnoses
Heptapterus from Pariolius, which was considered as a
Junior synonym of the former by Mees (1974).

In other aspects, our diagnosis of Heptapterus agrees
with that by Bleeker (1864). All species included here-
in in the genus have the pelvic-fin insertion at the verti-
cal through the base of the first dorsal-fin ray or slightly
posterior to it (up to third). By that character, they differ
from several members of Heptapterini, such as C. bleek-
eri, Horiomyzon Stewart, 1986, Nannoglanis Boulenger,
1887, Pariolius, and most Phenacorhamdia Dahl, 1961
(pelvic-fin insertion anterior to dorsal-fin insertion); and
Cetopsorhamdia Eigenmann & Fisher, 1916, and most /m-
parfinis sensu stricto (pelvic-fin insertion posterior to verti-
cal through fourth dorsal-fin ray). Still, that character is not
helpful to distinguish between Heptapterus and the most
similar genera, 1.e., Chasmocranus and Acentronichthys.

Used in combination, the adipose fin confluent with the
caudal fin and the non-bifurcate caudal fin distinguishes
Heptapterus from the remaining Heptapterini (except
C. bleekeri, some Nemuroglanis, and some of the spe-
cies recently removed from Heptapterus, viz. *‘H.’ mul-
tiradiatus, ‘H.’ stewarti and ‘H.’ sympterygium). More-
over, the extremely elongate body is found otherwise in
Heptapterini only in Acentronichthys, Chasmocranus,
some species of Phenacorhamdia, large specimens of

339

Rhamdioglanis thering, 1907, and the incertae sedis spe-
cies ‘I.’ longicauda (Boulenger 1887) and ‘I.’ microps
Eigenmann & Fisher, 1916.

One peculiar species of Heptapterini — the incertae se-
dis *C.” brachynema — does resemble Heptapterus species.
Whereupon the adipose fin is confluent with the caudal and
the pelvic-fin insertion is positioned between the verticals
through the insertion and through the middle of the dor-
sal-fin base, and the anal-fin insertion located posteriorly to
a vertical through the adipose-fin insertion (in which ‘C.’
brachynema differs from Chasmocranus). It is also quite
similar to H. borodini, H. carmelitanorum and H. hollandi
in having large, bulging, closely set eyes. However, ‘C.’
brachynema differs from all Heptapterus in having a dis-
tinctly shorter body (this character also distinguishes it
from all Chasmocranus, sensu stricto), bifurcate caudal fin
(although the notch between the two lobes is very shallow
and the dorsal one is distinctly longer than the ventral one);
and extremely long posterolateral extension of the premax-
illary toothplate (even longer than the extension present in
Chasmocranus, as can be seen in Gomes and Schubart
1958, and Mees 1967, fig. 1; this character is correlated
with the peculiar shape of the mouth of ‘C.’ brachynema).

Heptapterini in the southern Neotropics

The southern Neotropics (including the Parana-Paraguai
River system, the Sao Francisco River basin and all other
river basins that empty in the Atlantic Ocean between the
mouths of those major rivers) hold 35 valid Heptapteri-
ni species. These are: Acentronichthys leptos Eigenmann
& Eigenmann, 1889, A. fissipinnis, Cetopsorhamdia
iheringi, ‘Chasmocranus’ brachynema, ‘C.’ lopezae Mi-
randa-Ribeiro, 1968, °C.’ truncatorostris Borodin, 1927,
Heptapterus borodini, H. carmelitanorum, H. carnat-
us, H. exilis, H. hollandi, H. mandimbusu, H. mbya, H.
mustelinus, H. ornaticeps, H. qgenqo, ‘Heptapterus’ mul-
tiradiatus, “H.’ stewarti, ‘H.’ sympterygium, Imparfinis
minutus, I. mirini Haseman, 1911, 1 mishky Almiron,
Casciotta, Bechara, Ruiz Diaz, Bruno, d’ Ambrosio, Soli-
mano & Soneira, 2007, I. piperatus, I. schubarti (Gomes,
1956), ‘I.’ stictonotus (Fowler, 1940), Phenacorham-
dia roxoi Silva, 2020, P. tenebrosa (Schubart, 1964), P.
unifasciata Britski, 1993, P. hoehnei (Miranda Ribeiro,
1914), Rhamdioglanis frenatus, R. transfasciatus Miran-
da Ribeiro, 1908, Rhamdiopsis krugi Bockmann & Cas-
tro, 2010, R. microcephala (Litken, 1874), R. moreirai
Haseman, 1911, and 7Zaunayia bifasciata (Eigenmann &
Norris, 1900). This species richness comprises 39.3% of
the tribe. Considering genera, Acentronichthys, Rham-
dioglanis, Rhamdiopsis Haseman, 1911, and Taunay-
ia Miranda Ribeiro, 1918, are exclusively found in the
southern Neotropics, whereas Heptapterus is only mar-
ginally distributed in the Tocantins River basin.

Some species appear to be quite restricted in some
watercourses of southern Neotropics, such as ‘C.’
brachynema (Mogi-Guacu River), ‘C.’ /opezae (record-

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340 Depra, G.d.C. et al.: Redefinition of Heptapterus and description of a new species

ed from type locality in Cubatéo and from Ribeira de Ig-
uape River basin, but possibly restricted to the former),
H. carmelitanorum (restricted to type locality in the
Grande River basin), H. hollandi (Iguacu River basin),
‘H.’ multiradiatus (upper Tieté River basin), ‘H.’ stewarti
(upper stretches of Iguacu and Tibagi River basins), ‘H.’
sympterygium (Patos Lagoon basin), P. unifasciata (Pa-
ranaiba River basin), R. moreirai (upper stretches of Ig-
uacu and Tibagi River basins), Zaunayia bifasciata (up-
per stretches of Paraiba do Sul and Tieté River basins).
Others are thought to have a wider distribution, such as
H. borodini, I. mirini and I. schubarti (Ota et al. 2018;
Reis et al. 2020). However, populations assigned to those
Species, especially in State of Parana, seem to form spe-
cies complexes (G. C. Depra and V.M. Azevedo-Santos,
personal observation), which means that, instead of few,
widely distributed species, they may represent several rel-
atively restricted ones. The analysis of those populations,
which presently is being carried on by G. C. Depra, V.M.
Azevedo-Santos and collaborators, is likely to contribute
to an increase in the number of Heptapterini species, as
well as our knowledge of the biogeography of the Upper
Parana ecoregion — which already has been demonstrated
to comprise different sub-ecoregions, each with a peculiar
ichthyofauna (Reis et al. 2020).

Besides investigating the existence of species com-
plexes, we emphasize the importance of sampling poorly
known river basins, especially using different method-
ologies. For example, all the type specimens of H. car-
melitanorum were collected in a recent survey (Azeve-
do-Santos et al. 2019; and after new expeditions) carried
out in tributary streams of the Grande River system, in
southeastern Brazil (Azevedo-Santos et al. 2019). New
collections (one in 2020 and three in 2021) at the same
locality were performed and in streams with sources in
the same mountain (1.e., Chapadao) as the one in which
specimens of H. carmelitanorum were sampled, but no
additional specimens were captured. Publications (Alves
et al. 1998; Castro et al. 2004; Andrade and Braga 2005;
Ingenito and Buckup 2007; Pompeu et al. 2009; Belei and
Sampaio 2012; Casarim et al. 2012; Casatti et al. 2012:
Fernandes et al. 2012; Gandini et al. 2012; Fagundes et
al. 2015; Santos et al. 2017; Azevedo-Santos et al. 2018;
Thereza and Langeani 2019; Ribeiro et al. 2019; Lima
et al. 2021), as well as collections (e.g., LBP, DZSJRP)
with fishes from the Grande River basin were consulted,
but we were not able to find heptapterids (identified as
Imparfinis, Chasmocranus or Heptapterus) that represent
H. carmelitanorum. It is possible, therefore, that a new
survey may reveal undescribed heptapterids in single
streams or rivers of the upper Parana River basin.

Conclusion

Here we propose a new diagnosis of the genus
Heptapterus based on external characters. In addition, we
proposed H. carmelitanorum sp. nov. from the Grande

River basin, upper Parana River basin, in Minas Gerais,

zse.pensoft.net

Brazil. Heptapterus comprises ten valid species, viz.
H.. borodini, H. carmelitanorum, H. carnatus, H. exilis,
H. hollandi, H. mandimbusu, H. mbya, H. mustelinus,
H. ornaticeps, and H. qenqo. Additional surveys (as in
the case of H. carmelitanorum) and examination of hep-
tapterid specimens present in biological collections cer-
tainly will increase the diversity known to that genus.

Comparative material.

Material listed by Aguilera et al. (2011), Aguilera et al.
(2017), Faustino-Fuster et al. (2019) and individuals of
heptapterid species listed in Azevedo-Santos et al. (2019;
2020). In addition, the following lots were examined:

Acentronichthys leptos Eigenmann & Eigenmann, 1889:
ANSP 174017, 2, 76.1—78.2 mm SL, Macaé River, Rio
de Janeiro, Brazil.

Cetopsorhamdia iheringi Schubart & Gomes, 1959:
EEBP 368, 76.30 mm SL, Mogi Guacu River, Sao
Paulo, Brazil.

‘Chasmocranus’ brachynema Gomes & Schubart, 1958,
EEBP 629, 74.2 mm SL, Mogi Guacu River, Sao Pau-
lo, Brazil.

Chasmocranus longior Eigenmann, 1912: FMNH 53208,
holotype, 92.9 mm SL, Essequibo River, Potaro-Si-
paruni, Guyana.

‘Chasmocranus’ truncatorostris Borodin, 1927: AMNH
8640, holotype, 109.9 mm SL, Colonia Hansa, Santa
Catarina, Brazil.

Heptapterus borodini (Mees & Cala, 1989): AMNH
8639, holotype (examined by photograph); State of
Goias, Caldas Novas, Corumba River, Sao Paulo,
Brazil; NUP 5221, 6, 32.8-64.6 mm SL, Gamelei-
ra Stream, Goias, Brazil; NUP 6088, 1, 74.2 mm
SL. Piava Stream, Parana, Brazil; NUP 14882, 3,
44. 5—-85.9 mm SL, Araponga Stream, Mato Grosso
do Sul, Brazil.

Heptapterus hollandi Haseman, 1911: FMNH 54244,
holotype, 230.4 mm SL, Iguacu River, Porto Unido da
Victoria, Parana, Brazil; NUP 5978, 11, Caxias Reser-
voir, Capitao Leénidas Marques, Parana, Brazil.

‘Heptapterus’ multiradiatus \hering, 1907: FMNH
56901, 10, 34.3-86.8 mm SL, Tieté River, Sao Paulo,
Brazil. FMNH 93272, 73.1 mm SL, Upper rio Parana,
Sao Paulo, Brazil.

Imparfinis schubarti (Gomes, 1956): EEBP 391, Para-
type, 2, 80.3—90 mm SL, Mogi Guacu River, Sao Pau-
lo, Brazil.

Nemuroglanis lanceolatus Eigenmann & Eigenmann,
1889: FMNH 98306, 7, 14.5—-35.7 mm SL, Napo Riv-
er, Sucumbios, Ecuador.

Nemuroglanis mariai (Schultz, 1944): ANSP 139581, 1,
29.6 mm SL, Venturosa stream, Meta; ANSP 139582,
3, 20.0—23.1 mm SL, El Viento creek, Matazul, Meta,
Colombia; ANSP 139583, 1, 35.1 mm SL, unnamed
stream tributary to Mozambique lake, Hacienda Hu-
macita, Meta, Colombia.

Zoosyst. Evol. 98 (2) 2022, 327-343

Acknowledgments

We are grateful to Paula N. Coelho, Manoel T. Azevedo,
and Eugen E. Horvath for help with fieldwork. To Edmar
Bueno and Glicério Martins for allowed access to the type
locality of the new species. Maria Ines Borella and Marcos
Antonio Pereira de Godoy, who allowed access to the Mu-
seu de Historia Natural de Pirassununga. To Wagner M.S.
Sampaio and Patricia Giongo to allow us access to exam-
ine heptapterids of Instituto de Pesquisa em Fauna Neo-
tropical. To Isabel M. Soares, who helped the authors in
the analysis of some comparative specimens. To Claudio
de Oliveira (LBP), Cristiano Moreira (MNRJ), Nathan Lu-
jan (AMNH), Mark Sabaj and Mariangles Arce (ANSP),
C. McMahan, S. Mochel and K. Sagel (FMNH), Felipe P.
Ottoni (UFMA) and Luiz Malabarba and Juliana Wing-
ert (UFRGS) for curatorial assistance; and Marcelo Sales/
MNRJ for helping with X-rays images (Equipamento de
Radiografia Digital da Central Analitica Virtual do Museu
Nacional, Universidade Federal do Rio de Janeiro FINEP
Proc 14629-3; Proinfra 01/2007). To Josie South (Univer-
sity of Leeds) and three reviewers for the suggestions that
help us to improve the quality of this study. This manuscript
is dedicated to Valter M. Santos, Isabel C. Azevedo San-
tos, Valquiria Achcar M. Silva (in memoriam), M. Lourdes
Cabral, Manoel T. Azevedo Filho, Luiz Fernando de Aze-
vedo, and Joaquim da Silva Junior, some of the Carmeli-
tanos who had important role in the formation of the first
author of the species. AMK was supported by Funda¢aéo
Carlos Chagas Filho de Amparo a Pesquisa do Estado do
Rio de Janeiro (FAPERJ; grant E-26/202.005/2020); and
DRFF was supported by the Bohlke Award from the Acad-
emy of Natural Science of Drexel University (ANSP) and
by the Grainger Bioinformatics Center funding at the Field
Museum of Natural History (FMNH).

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Supplementary material |

Figures S1—-S5

Authors: Gabriel de Carvalho Depra, Gaston Aguilera,
Dario R. Faustino-Fuster, Axel M. Katz, Valter M.
Azevedo-Santos

Data type: Figures (docx. file)

Explanation note: Figure S1. Heptapterus carmelitano-
rum, new species, paratype, LBP 23577, 104.4 mm
SL, (a). Dorsal view, (b). Lateral view, (c). Ventral

343

view. Figure 82. Heptapterus carmelitanorum, new
species, paratype, LBP 26570, 95.7 mm SL. (a). Dor-
sal view, (b). Lateral view, (c). Ventral view. Figure
S3. Heptapterus carmelitanorum, new species, para-
type, color in life, LBP 23577, 104.4 mm SL. Figure
S4. X-ray of Heptapterus carmelitanorum, holotype,
MNRJ 53174, 144.3 mm SL. (a). Lateral view, (b).
Ventral view, (c). Dorsal view. Figure S5. X-ray
of Heptapterus carmelitanorum, holotype, MNRJ
53174, 144.3 mm SL. (a). Ventral view of head, (b).
Lateral view of caudal fin.

Copyright notice: This dataset is made available under
the Open Database License (http://opendatacommons.
org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow us-
ers to freely share, modify, and use this Dataset while
maintaining this same freedom for others, provided
that the original source and author(s) are credited.

Link: https://do1.org/10.3897/zse.98.89413.suppl1

zse.pensoft.net