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101
Two new species of Curvianchoratus (Monogenoidea,
Dactylogyridae) parasitizing Psectrogaster amazonica
(Characiformes, Curimatidae) and a new record for
Curvianchoratus singularis in the Tocantins River, Maranhão, Brazil
Carine Almeida Miranda Bezerra1, Simone Chinicz Cohen2, Yuri Costa de Meneses2,
Helyab Gabriel Chaves Neres3, Diego Carvalho Viana3, Marcia Cristina Nascimento Justo2
1 Programa de Pós-Graduação em Ciência Animal, Universidade Estadual do Maranhão, Cidade Universitária Paulo VI, Avenida Lourenço Vieira da Silva 1000,
65055-310 São Luís, MA, Brazil
2 Laboratório de Helmintos Parasitos de Peixes, Instituto Oswaldo Cruz, FIOCRUZ, Avenida Brasil 4365, 21040-900 Rio de Janeiro, RJ, Brazil
3 NúcleodeEstudosMorfosiológicosAvançados,UniversidadeEstadualdaRegiãoTocantinadoMaranhão,CampusImperatriz,RuaGodofredoViana1300,
65901-480Imperatriz,MA,Brazil
Correspondingauthor:SimoneChiniczCohen(scohen@ioc.ocruz.br)
Copyright: © Carine A. M. Bezerra et al.
This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International –
CC BY 4.0).
Research Article
Abstract
Several studies have demonstrated parasitism by monogenoids in characiform sh
in the Neotropics. During studies on the helminth fauna of curimatids from the To-
cantins River, specimens of Psectrogaster amazonica Eigenmann & Eigenmann, 1889
were examined and species of Curvianchoratus Hanek, Molnar & Fernando, 1974
were found. Species of the genus are characterized mainly by the complex shape
of haptoral anchors with a modied dorsal anchor, composed by two subunits, dor-
sal-median and dorsal. To date, two species of Curvianchoratus are known to para-
sitize curimatid shes: the type species Curvianchoratus hexacleidus Hanek, Molnar
& Fernando, 1974 and Curvianchoratus singularis (Suriano, 1980). During examina-
tion of specimens of P. amazonica collected in the Tocantins River, Embiral, Imper-
atriz, Maranhão State, Brazil, two new species of Curvianchoratus were found and
are described herein. Curvianchoratus psectrogasteri sp. nov. and Curvianchoratus
dominguesi sp. nov. are characterized by possessing the male copulatory organ
formed by a long cirrus and a claw-shaped accessory piece, connected to the base
of the male copulatory organ by a ligament. The new species differs from the two
known congeneric species mainly by the morphology of the dorsal-median and dorsal
subunits of the dorsal anchor. Curvianchoratus psectrogasteri sp. nov. also differs
from other species of the genus by the absence of the ventral bar and Curvianchor-
atus dominguesi sp. nov. by the size and shape of the ventral bar. An amendment to
the diagnosis of Curvianchoratus is provided to accommodate the new species. The
present study increases the number of Curvianchoratus species to four and extends
the occurrence of the genus to the Tocantins-Araguaia Basin.
Key words: Curvianchoratus dominguesi sp. nov., Curvianchoratus psectrogasteri sp.
nov., ectoparasites, sh parasite, helminth fauna, Neotropical Region, taxonomy, Tocan-
tins-Araguaia Basin
Academic editor: Patrick M. Delgado
Received:
26 April 2023
Accepted:
26 May 2023
Published:
25 July 2023
ZooBank: https://zoobank.
org/6A0A780E-B78B-4E54-9237-
D01A82479F7B
Citation: Bezerra CAM, Cohen
SC, de Meneses YC, Neres HGC,
Viana DC, Justo MCN (2023) Two
new species of Curvianchoratus
(Monogenoidea, Dactylogyridae)
parasitizing Psectrogaster amazonica
(Characiformes, Curimatidae) and
a new record for Curvianchoratus
singularis in the Tocantins River,
Maranhão, Brazil. ZooKeys 1172:
101–116. https://doi.org/10.3897/
zookeys.1172.105500
ZooKeys 1172: 101–116 (2023)
DOI: 10.3897/zookeys.1172.105500
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ZooKeys 1172: 101–116 (2023), DOI: 10.3897/zookeys.1172.105500
Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Introduction
Characiformes is one of the main orders of Ostariophysi, which occur exclu-
sively in freshwater, with the greatest diversity and species richness concen-
trated in the Neotropical Region. These sh represent the historical connection
between Africa and South America, which can be proven through their distribu-
tion and the presence of fossil records (Malabarba and Malabarba 2010, 2014).
Among the families of the order, Curimatidae comprises eight extant genera
and more than 110 species, living in diverse freshwater habitats from Costa
Rica to Argentina (Melo et al. 2018).
Psectrogaster Eigenmann & Eigenmann, 1889 is a morphologically diverse
lineage of Curimatidae, distributed in streams, rivers, and still waters through-
out the drainage basins of the Orinoco River, Essequibo River, Amazon River,
Paraguay-Parana River, and some of the rivers of northeastern Brazil (Vari
1989). Psectrogaster amazonica Eigenmann & Eigenmann, 1889, popularly
known in Brazil as “branquinha” (Santos et al. 1984) is a freshwater sh that
lives in tropical waters in the Tocantins and Amazon River basins (Vari and
Röpke 2013; Froese and Pauly 2022). This species presents benthopelagic be-
havior, with the formation of large shoals, and has a maximum length of 19
cm, being one of the largest species of the genus (Vari 1989). The species is
one of the most common and abundant along the Tocantins River in midwest,
north and northeast Brazil and is the basis of an intensive shery in this area.
Individuals belonging to this species primarily feed on organic detritus, benthic
organisms, and periphyton (Santos et al. 1984).
Several studies have demonstrated parasitism by monogenoids in Curimati-
dae sh in the Neotropics (Table 1). However, research involving the helminth
fauna of P. amazonica is still incipient, with only two monogenoids described
for this host so far (Freitas et al. 2021). Other studies regarding this sh spe-
cies refer to larval development, the distribution of larvae in a natural breeding
area, hematological parameters, and branchial histopathological alterations
(Nascimento and Araújo-Lima 1993; Carvalho et al. 2009; Ponte et al. 2017;
Bezerra et al. 2020; Pereira et al. 2020).
Monogenoidea Bychowsky, 1937 are, in general, ectoparasites that mainly
infect sh species (Cohen et al. 2021). They have a simple and direct life cycle
and, consequently, may present a high reproduction rate in these hosts (Lapera
et al. 2017). Monogenoids are among the main components of the parasitic
fauna of freshwater sh that occur in the Neotropics, most of them belonging
to the family Dactylogyridae Bychowsky, 1933 (Cohen et al. 2013).
Curvianchoratus was proposed by Hanek, Molnar and Fernando (1974) to
accommodate Curvianchoratus hexacleidus Hanek, Molnar & Fernando, 1974, a
parasite of the characiform Steindachnerina argentea (Gill, 1858) (syn. Curima-
tus argenteus Gill, 1858) from the Arouca River, Trinidad. The main characteris-
tic of Curvianchoratus is the haptor with a modied dorsal anchor, composed of
two subunits, which is a unique shape within Dactylogyridae.
To date, two species of Curvianchoratus are known, both parasites of
curimatid shes: the type species C. hexacleidus and Curvianchoratus sin-
gularis (Suriano, 1980), originally described as Notodiplocerus singularis
by Suriano (1980) parasitizing Cyphocharax gilbert (Quoy & Gaimard, 1824)
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Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Table 1. Monogenoideans parasites of Curimatidae from South America.
Parasite Host Locality References
Anacanthoroides mizellei Kritsky &Thatcher, 1976 Steindachnerina insculpta Brazil Acosta et al. (2013)
Annulotrematoides amazonicus Kritsky & Boeger, 1995 Psectrogaster rutiloides Brazil Kritsky and Boeger (1995)
Annulotrematoides bonaerensis Rossim & Timi, 2016 Cyphocharax voga Argentina Rossin and Timi (2016)
Cacatuocotyle paranaensis Boeger, Domingues &
Kritsky,1997
Cyphocharax nagelii Brazil Boeger et al. (1997), Vieira et
al. (2013), Dias et al. (2017)
Characithecium chascomusensis (Suriano, 1981) Rossin &
Timi, 2014
Cyphocharax gilbert Argentina Suriano (1981)
C. voga Suriano (1997)
Curvianchoratus hexacleidus Hanek, Molnar & Fernando, 1974 Cyphocharax nagelii Brazil Vieira et al. (2013)
Curvianchoratus singularis (Suriano, 1980) Suriano 1986 C. gilbert Argentina Suriano (1980)
C. voga Rossin and Timi (2016)
Cyphocharax nagelii Brazil Vieira et al. (2013), Dias et
al.(2017)
Cyphocharax modestus Dias et al. (2017)
Steindachnerina insculpta Dias et al. (2017)
Diaphorocleidus kabatai Mendoza-Franco, Reina & Torchin,
2009
Stendachnerina insculpta Brazil Acosta et al. (2013)
Euryhaliotrema chaoi Kritsky & Boeger, 2002 Steindachnerina insculpta Brazil Acosta et al. (2013)
Paranaella luquei Kohn, Baptista-Farias & Cohen, 2000 Stendachnerina brevipinna Brazil Ceschini et al. (2010)
Philocorydoras margolisi (Molnar, Hanek & Fernando, 1974)
Yamada, Brandão, Yamada & Silva, 2015
Steindachnerina insculpta Brazil Dias et al. (2017)
Rhinoxenus guianensis Domingues & Boeger, 2005 Curimata cyprinoides French
Guyana
Domingues and Boeger (2005)
Urocleidoides curimatae Molnár, Hanek & Fernando, 1974 Steindachnerina argentea Trinidad Molnár et al. (1974)
Urocleidoides paratriangulus Freitas, Bezerra, Meneses,
Justo, Viana & Cohen, 2021
Psectrogaster amazonica Brazil Freitas et al. (2021)
Cyphocharax gouldingi
Urocleidoides surianoae Rossim & Timi, 2016 Cyphocharax voga Argentina Rossin and Timi (2016)
Urocleidoides tocantinensis Freitas, Bezerra, Meneses,
Justo, Viana & Cohen, 2021
Psectrogaster amazonica Brazil Freitas et al. (2021)
Urocleidoides triangulus (Suriano, 1981) Rossim &
Timi,2016
C. nagelii Brazil Vieira et al. (2013), Abdallah et
al. (2015), Dias et al. (2017)
Cyphocharax modestus Abdallah et al. (2015), Dias et
al. (2017)
C. gilberti Freitas et al. (2021)
Argentina Suriano (1981), Suriano (1997)
[syn. Pseudocurimata gilbert (Quoy & Gaimard, 1824)] from Argentina. Suriano
(1986) transferred the parasite to Curvianchoratus. Rossin and Timi (2016) re-
described this species from Cyphocharax voga (Hensel, 1870) from Argentina.
In Brazil, C. singularis has been reported in the rivers of the state of São Paulo
parasitizing Curimatidae shes (Vieira et al. 2013; Dias et al. 2017).
During studies on the helminth fauna of curimatids from the Tocantins
River, specimens of P. amazonica were examined and species of Curvianchora-
tus were found. The present paper presents an emended diagnosis of Curvian-
choratus, with the proposal of two new species and a new geographical record
for C. singularis.
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Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Material and method
From July 2018 to April 2022, 136 samples of P. amazonica (average 14 cm in
standard length and average 62 g in weight) were collected in the Tocantins Riv-
er, close to the urban perimeter of Imperatriz, in the village of Embiral, state of
Maranhão, Brazil (5°27'50’’S, 47°33'48’’W) (Fig. 1). Fish were caught with the aid
of local shermen using gill nets, or a hook and line and were examined asites
immediately after capture at the Nucleus of Advanced Morphophysiological
Studies at UEMASUL. The gills were removed and placed in vials containing
hot water (65 °C) in order to detach the parasites from the gill laments. Later,
absolute ethanol was added to reach a concentration of 70%. Monogenoids
were collected from the sediment and gill arches in the laboratory with the aid
of a stereoscopic microscope. Specimens were mounted in Hoyer’s medium
to study the hard parts, such as the copulatory complex and haptoral sclerites
and some were stained with Gomori’s trichrome to study the internal organs of
the parasite (Kritsky et al. 1978). Measurements are presented in micrometers;
range values are followed by mean and number of structures measured in pa-
rentheses. Sclerotized structures, such as bars and anchors were measured
according to the scheme shown in Fig. 2. The parasitological indices were cal-
culated as proposed by Bush et al. (1997), followed by the standard deviation.
The authorship of the taxa followed the recommendation of Article 50.1 of the
International Code of Zoological Nomenclature (ICZN), which deals with the
identity of the authors. The specimens were observed using an Olympus BX 41
microscope with phase contrast and Zeiss Axioskop 2 Plus microscope with
Figure 1. Map of Middle Tocantins River, showing the collection point of the hosts close to the urban perimeter of Imper-
atriz, in the village of Embiral, state of Maranhão, Brazil.
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Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Figure 2. Scheme of measurements of the sclerotized structures of haptoral anchors of Curvianchoratus spp. A dorsal-medi-
an subunit and B dorsal subunit of Curvianchoratus psectrogasteri sp. nov. C dorsal-median subunit D dorsal subunit of Cur-
vianchoratus dominguesi sp. nov. (sf) shaft, (pt) point, (dr) deep root, (sr) supercial root, (b) base, (tl) total length, (wd) width.
differential interference contrast, both equipped with a camera lucida for draw-
ings. The images were captured with a SONY MPEG Movie EX DSC-S75 digital
camera. Type specimens were deposited in the Coleção Helmintológica do In-
stituto Oswaldo Cruz (CHIOC) in Brazil.
Results
In the present study, a total of 87 monogenoid specimens were collected, being:
68 Curvianchoratus psectrogasteri sp. nov., eight Curvianchoratus dominguesi
sp. nov. and 11 Curvianchoratus singularis.
Taxonomic account
Class Monogenoidea Bychowsky, 1937
Subclass Polyonchoinea Bychowsky, 1937
Order Dactylogyridea Bychowsky, 1937
Dactylogyridae Bychowsky, 1933
Curvianchoratus Hanek, Molnár & Fernando, 1974
Emended diagnosis. Tegument smooth, thin. Body elongated; divisible into
cephalic area, trunk, peduncle, and haptor. Terminal and lateral cephalic lobes
developed or not; head organs and cephalic glands present. Eyespots present
or absent. Mouth subterminal, midventral, prepharyngeal. Pharynx muscular;
esophagus short. Intestinal caeca two, not presenting diverticula. Common gen-
ital pore midventral near the level of intestinal bifurcation. Gonads intercaecal,
overlapping or not; testis dorsal to germarium. Seminal vesicle large, dilated por-
tion of vas deferens; prostatic reservoir single. Copulatory complex comprising
male copulatory organ (MCO) and accessory piece; MCO consisting of sclero-
tized coiled tube with counterclockwise rings, which make a short turn on itself
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Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
near the base; cirrus and accessory piece articulated basally. Accessory piece
with bifurcated distal portion, pincer-shaped. Ovary pretesticular. Vagina medial
or sinistral. Vitellaria well developed, coextensive with intestinal ceca, absent
from regions of other reproductive organs. Peduncle short, haptor subquadrate.
Ventral bar present or absent. Dorsal bar absent. Ventral anchor of dactylogyrid
type. Dorsal anchor modied, composed by two subunits, dorsal-median and
dorsal, forming a full circle, the shaft of the dorsal subunit passes over the su-
percial root of the dorsal-median subunit anchor. Hooks 14; 5 pairs ventral, 2
pairs dorsal, with inated base and slender shank. Parasites of curimatid shes.
Remarks. Curvianchoratus was proposed to accommodate Curvianchoratus
hexacleidus Hanek, Molnár & Fernando, 1974, a parasite of the characiform S.
argentea from the Arouca River, Trinidad (Hanek et al. 1974). The main charac-
teristic of Curvianchoratus is the haptor with a modied dorsal anchor, com-
posed of two subunits, which is a unique shape for the genus. The absence
of bars in one of the species described here, made it necessary to expand the
original diagnosis of the genus to include the characteristics of the new spe-
cies described below, as well as to detail some morphological characters.
Curvianchoratus psectrogasteri Bezerra, Cohen, Meneses & Justo, sp. nov.
https://zoobank.org/CB6BBC35-0D07-4EC8-A9E6-1BD568F8899B
Figs 3A–H, 5A, B, E
Type host. Psectrogaster amazonica Eigenmann & Eigenmann, 1889 (Curimatidae).
Type locality. Embiral, rural zone (5°27'50"S, 47°33'48"W), municipality of Im-
peratriz, Maranhão State.
Infestation parameters. 22.06% prevalence; 68 total number of parasites;
2.27 ± 1.75 mean intensity; 1-15 range of intensity.
Deposited material. Holotype: CHIOC 39977a; paratypes: CHIOC 39977 b-d,
39978 a, b, 39979, 39980, 39981, 39982.
Etymology. The specic name of the species is derived from the generic
name of the host.
Description. Based on 43 specimens: 21 mounted in Gomori’s trichrome and
22 mounted in Hoyer’s medium: Body elongated, 425–883 (665; N = 30) long,
135–520 (308; N = 30) wide. Tegument smooth. Cephalic region with poor-
ly-developed cephalic lobes; ve pairs of head organs; cephalic glands lateral
to pharynx. Eyes 4; anterior pair smaller than posterior. Pharynx round, muscular,
32–75 (50; N = 18) long, 32–87 (50; N = 18) wide; esophagus short. Peduncle in-
conspicuous, followed by a small haptor compared to body size, 125–292 (225;
N = 21) wide. Ventral bar absent. Ventral anchor with inconspicuous deep root
and elongated supercial root, long and slightly curved shaft; straight tip, not
passing from the level of the tip of the supercial root, 21–30 (24; N = 43) long,
base, 10–13 (12; N = 43) wide. Dorsal anchor complex heavily modied, com-
posed by 2 subunits: dorsal-median and dorsal. Dorsal-median subunit strongly
curved, 52–87 (64; N = 34) long and base, 41–63 (49; N = 48), with the robust
shaft, 63–92 (78; N = 34) long, subdivided into two pieces, one anterior, with
sclerotized ridges, and one posterior, lightly sclerotized, point sickle-shaped,
articulated with the dorsal subunit, 4–7 (6; N = 36) long; deep and supercial
roots well developed, crested-shaped, 12–27 (16; N = 46) long and 10–20 (14;
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Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Figure 3. Curvianchoratus psectrogasteri sp. nov. from Psectrogaster amazonica from Tocantins River A whole specimen,
ventral view B copulatory complex, ventral view C vagina D egg E ventral anchor F hook G dorsal-median subunit H dorsal
subunit. Scale bars: 100 μm (A); 10 μm (B, C, E, F); 30 μm (D, G, H).
N = 37) long, respectively. Dorsal subunit large, irregular, 40–65 (48; N = 15) long
well-developed deep root 19–22 (20; N = 17), supercial root absent, and medi-
an region expands to form a rounded structure, 32–57 (40; N = 23) wide. Hooks
similar in shape, pairs 1 and 5 smaller than others, 11–14 (12; N = 27) long, pairs
2–4,6,7 with erected thumb, curved shaft, short point, shank with dilated bulb at
distal end, 13–18 (16; N = 95). Copulatory complex comprising male copulatory
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Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
organ (MCO) and articulated accessory piece. Male copulatory organ composed
by a tubular sclerotized cirrus that makes a short turn on itself near the base and
spirals halfway towards the tip of the cirrus, with sclerotized base possessing
delicate membrane, 47–60 (53; N = 5) in total length. Accessory piece, 25–30
(26; N = 5) long, claw-shaped, with dissimilar tip sizes, the smaller tip serves as
guide to MCO, hinged by a ligament to the base of the cirrus. Testis oval, dorsal
to germarium; seminal vesicle a distal enlargement of vas deferens; prostatic
reservoir single, posterior to MCO. Germarium pretesticular, 62–122 (102; N = 7).
Vaginal aperture midway between ovary and copulatory complex, sinistro-ven-
tral, with vaginal canal slightly sclerotized, followed by rounded seminal recepta-
cle, anterior to germarium. Oviduct, ootype, uterus not observed. Vitellaria dense,
extends over the entire body, except in the region of the reproductive organs and
haptor. Eggs operculated, 72–87 (82; N = 3) long by 60–70 (64; N = 3) wide.
Remarks. Curvianchoratus psectrogasteri sp. nov. is allocated in the genus
mainly by the morphology of the copulatory complex, composed by a tubular cir-
rus and a pincer-shaped accessory piece, connected to the base of the cirrus by
a ligament and by the presence of the greatly modied dorsal anchor, composed
by two subunits: dorsal-median and dorsal. The new species differs from the two
known species by the absence of the ventral bar and by the morphology of the
dorsal-median subunit (strongly curved, with the shank and tip visibly separated;
a robust shank ornamented with a thorn at the base and crested-shaped deep and
supercial roots in the new species vs. a stickle shaped end inserted in the cavity
of the shaft in C. hexacleidus and C. singularis) and by the morphology of the dor-
sal subunit (elongated, with a broadly circular central region in the new species
vs. elongated, with deep root in C. singularis and without in C. hexacleidus).
Curvianchoratus dominguesi Bezerra, Cohen, Meneses & Justo, sp. nov.
https://zoobank.org/E6B46BAD-9990-4E75-8781-D4DB18DE70D0
Figs 4A–G, 5C, D, F
Type host. Psectrogaster amazonica Eigenmann & Eigenmann, 1889 (Curimatidae).
Type locality. Embiral, rural zone (5°27'50"S, 47°33'48"W), municipality of Im-
peratriz, Maranhão State.
Infestation parameters. 5.15% prevalence; 8 total number of parasites; 1.14
± 0.26 mean intensity; 1–2 range of intensity.
Deposited material. Holotype: CHIOC 39984; paratypes: CHIOC 39985,
39986, 39987, 39988, 39989.
Etymology. The specic name is in honor of Dr Marcus Vinicius Domingues
for his contributions to the knowledge of neotropical monogenoidean fauna.
Description. Based on 7 specimens: 1 mounted in Gomori’s trichrome and 6
mounted in Hoyer’s medium: Body short, broad, 460–750 (575; N = 6) long, 225–
500 (362; N = 6) wide. Tegument smooth. Cephalic region with 3 pairs of lateral
lobes (two lateral and one terminal); ve pairs of head organs; cephalic glands
lateral to pharynx. Eyespots absent. Pharynx muscular, 50–63 (55; N = 3) long,
48–69 (58; N = 3) wide, esophagus short. Haptor small, compared to body size
length, 400 (N = 2) wide. Ventral bar short and robust, anvil-shaped, with slightly
jagged edges, 50–58 (52; N = 4) long. Ventral anchor with inconspicuous roots,
shaft with two subunits, 25–32 (30; N = 5) long, with a short base, 11–17 (14; N =
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Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Figure 4. Curvianchoratus dominguesi n. sp from Psectrogaster amazonica from Tocantins River A whole specimen,
ventral view B copulatory complex, ventral view C ventral bar D hook E ventral anchor F dorsal-median subunit G dorsal
subunit. Scale bars: 100 μm (A); 30 μm (B, F, G); 10 μm (C–E).
5). Dorsal anchor complex heavily modied composed by 2 subunits: dorsal-me-
dian and dorsal. Dorsal-median subunit robust curved and heavily sclerotized,
50–66 (58; N = 13) long, shaft, 41–47 (44; N = 14); point as a robust hooked
thorn, 6–10 (8; N = 14); supercial root, 5–8 (7; N = 3) and deep root, 13–17 (15;
N = 6), well developed with borders very close to each other; point, 6–10 (8; N
= 14). Dorsal subunit elongated, 65–100 (77; N = 9) long and 18–23 (19; N = 9)
wide, 30% larger than the dorsal-median subunit, posterior portion fan-shaped.
Hooks similar in shape, with erected thumb, curved shaft, and dilated bulb in the
end, pair 1 smaller than the others, 11–12 (11; N = 4) long; pairs 2–7, 14–17 (16;
N = 37). Copulatory complex comprising male copulatory organ and articulat-
ed accessory piece. MCO tubular, sclerotized, makes a short coil on itself, near
the cirrus base, 55–75 (69; N = 4) in total length. Accessory piece claw-shaped,
bifurcated, with different tip sizes; the smaller serves as a guide for the MCO,
42–70 (56; N = 5) long. Testis oval dorsal to the germarium; seminal vesicle
formed by a distal enlargement of the vas deferens; prostatic reservoir single, in
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Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Figure 5. Light photomicrographs of Curvianchoratus spp. A, B, E Curvianchoratus psectrogasteri sp. nov. A total, ventral
view B copulatory complex, ventral view E haptor C, D, F Curvianchoratus dominguesi sp. nov. C copulatory complex, ven-
tral view D total, ventral view F haptor. Scale bars: 100 μm (A); 20 μm (B); 30 μm (C, F); 100 μm (D); 40 μm (E).
form of half-moon. Germarium pretesticular; seminal receptacle anterior to ger-
marium; vaginal opening sinistro-ventral; vaginal canal wall sclerotized; oviduct,
ootype, uterus, and eggs not observed. Vitellaria dense extends over the entire
body, except in the region of the reproductive organs and haptor.
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Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Remarks. Curvianchoratus dominguesi sp. nov. is similar to the other species
of the genus mainly by the morphology of the dorsal anchor, the copulatory com-
plex, which is composed by a long cirrus and a pincer-shaped accessory piece,
connected to the base of the cirrus. The new species is similar to C. singularis and
C. hexacleidus by the presence of the ventral bar. The new species differs from
all congeneric species by the shape of the ventral anchor (formed by a shaft with
two subunits in the new species vs. well-developed roots in C.hexacleidus and
moderately developed deep root and well-developed supercial root in C. singu-
laris and C. psectrogasteri sp. nov.), the morphology of the ventral bar (short and
robust, anvil-shaped ventral bar in C. dominguesi sp. nov. vs. serpentine-shaped
in C. hexacleidus, M-shaped in C. singularis and absent in C. psectrogasteri sp.
nov.) and by the ratio of the subunits of the dorsal anchor (dorsal subunit is 30%
larger than the dorsal-median subunit in C. dominguesi sp. nov. vs. the approxi-
mately same size of the subunits in the other three species of the genus).
Curvianchoratus singularis (Suriano, 1980) Suriano, 1986
Notodiplocerus singularis Suriano, 1980. Syn.
Host. Psectrogaster amazonica Eigenmann & Eigenmann, 1889 (Curimatidae)
Site of infection. Gill.
Locality. Embiral, rural zone (5°27'50"S, 47°33'48"W), municipality of Imper-
atriz, Maranhão State.
Infestation parameters. 29.37% prevalence; 104 total number of parasites;
2,81 (±6,3) mean intensity; 1–38 range of intensity.
Type host and locality. Cyphocharax gilbert (syn. Pseudocurimata gilbert)
from Chascomus Lagoon, Buenos Aires Province, Argentina (Suriano 1980).
Other records. Cyphocharax voga from Chascomus Lagoon, Buenos Aires
Province, Argentina (Rossin and Timi 2016), Cyphocharax modestus, Cypho-
charax nagelii and Steindachnerina insculpta from Batalha and Peixe Rivers,
São Paulo State, Brazil (Vieira et al. 2013; Dias et al. 2017).
Specimens deposited. Voucher CHIOC 39990, 39991, 39992.
Remarks. Suriano (1980) described the species as Notodiplocerus singularis
and the original drawings are simplied lacking important details of the mor-
phology of these unique anchors that characterize the genus. In the description
provided by Rossin and Timi (2016), the morphology of the dorsal anchor com-
plex and copulatory complex was described in detail and illustrated by means
of drawings and photographs. The morphology of the present material is in
agreement with the material studied by Rossin and Timi (2016).
Discussion
Monogenoideans represent one of the main components of the helminth fauna
present in freshwater sh in the Neotropics, most of which are dactylogyridean
parasites of teleost sh (Boeger and Vianna 2006; Cohen et al. 2013; Braga
et al. 2014). The great heterogeneity of hydrographic basins present in Brazil
reects a rich and diversied monogenoid fauna in this region (Tavares-Dias et
al. 2022). Despite taxonomic efforts, the diversity of monogenoid parasites of
112
ZooKeys 1172: 101–116 (2023), DOI: 10.3897/zookeys.1172.105500
Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
freshwater shes from South America remains poorly known (Justo and Co-
hen 2019). According to Melo et al. (2018), most of the information regarding
monogenoidean diversity in the Neotropical Region is limited to medium-sized
sh species, while the parasitic fauna of small-sized species, especially Charac-
iformes and Siluriformes, is not yet described.
Studies on the parasite diversity of freshwater sh from the Tocantins-Ara-
guaia Basin have gradually increased in recent years [see Fehlauer and Boeger
(2005); Boeger et al. (2006); Kritsky et al. (2007); Domingues and Marques
(2011); Kritsky et al. (2013); Aguiar et al. (2017); Cárdenas et al. (2019, 2021,
2022); Cohen et al. (2020); Freitas et al. (2021)].
In this sense, the proposal of two new species of Curvianchoratus contrib-
utes to the knowledge of parasite fauna of the region and enlarges the morpho-
logical characters used to describe the unique species that ventral bar is ab-
sent (Curvianchoratus psectrogasteri sp. nov.), which is contrary to the known
species of the genus. Although the presence of the bar is an important charac-
teristic, we adopted a conservative approach when proposing a new species of
this genus, because C. psectrogasteri exhibit the complex morphology of the
dorsal anchor, characteristic of Curvianchoratus.
With the description of the two new species in the present study, Curvian-
choratus now encompasses four species, all parasitizing curimatid shes. Cur-
vianchoratus singularis is herein reported in P. amazonica from the Tocantins
River, expanding the occurrence of the species to the Tocantins-Araguaia Ba-
sin. The nding of three species of the genus in P. amazonica conrms the high
specicity of the genus to neotropical curimatid hosts. On the other side, Psec-
trogaster amazonica harbors ve monogenoidean species, two from a general-
ist monogenoidean genus, Urocleidoides (parasites from three different sh or-
ders) (Freitas et al. 2021) and three from the specialist genus Curvianchoratus.
Additional information
Conict of interest
No conict of interest was declared.
Ethical statement
Permission to collect the hosts was given by the Instituto Chico Mendes de Conser-
vação da Biodiversidade (ICMBio) through the Sistema de Autorização e Informação da
Biodiversidade (SISBIO) under license number 79538-1.
Funding
Bezerra, CAM was supported by a doctoral scholarship Fundação de Amparo à Pesquisa
e ao Desenvolvimento Cientíco e Tecnológico do Maranhão (FAPEMA) #154086/2021.
Author contributions
Conceptualization: SCC, MCNJ, CAMB. Formal analysis: SCC, MCNJ. Methodology:
CAMB, HGCN, YCM. Project administration: DCV. Writing – original draft: CAMB. Writing
– review and editing: SCC, MCNJ.
Carine Almeida Miranda Bezerra: Collected the hosts, and processed the methodology
of collect of parasites; substantial contribution in the concept; contribution to data anal-
ysis and interpretation; contribution to manuscript preparation; Simone Chinicz Cohen:
113
ZooKeys 1172: 101–116 (2023), DOI: 10.3897/zookeys.1172.105500
Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Substantial contribution in the concept and design of the study; contribution to data anal-
ysis and interpretation; contribution to data collection; contribution to manuscript prepa-
ration; contribution to critical revision, adding intelectual content. Yuri Costa de Meneses:
Processed the methodology to prepare of parasites; contribution to critical revision, add-
ing intelectual content. Helyab Gabriel Chaves Neres Collected the hosts, and processed
the methodology of collect of parasites. Diego Carvalho Viana: Project administration.
Marcia Cristina Nascimento Justo: Substantial contribution in the concept and design
of the study; contribution to data analysis and interpretation; contribution to manuscript
preparation; contribution to critical revision; Writing original draft; prepared gures.
Author ORCIDs
Carine Almeida Miranda Bezerra https://orcid.org/0000-0003-4842-1595
Simone Chinicz Cohen https://orcid.org/0000-0001-8204-336X
Yuri Costa de Meneses https://orcid.org/0000-0002-8737-1159
Helyab Gabriel Chaves Neres https://orcid.org/0000-0002-4454-0223
Diego Carvalho Viana https://orcid.org/0000-0002-3302-9892
Marcia Cristina Nascimento Justo https://orcid.org/0000-0002-0684-3389
Data availability
All of the data that support the ndings of this study are available in the main text.
References
Abdallah VD, Azevedo RK, Silva RJ (2015) First record of Palombitrema triangulum (Suriano,
1981) Suriano, 1997 (Monogenea: Dactylogyridae) from freshwater shes in Brazil. Bra-
zilian Journal of Biology 75(1): 229–233. https://doi.org/10.1590/1519-6984.12913
Acosta AA, Queiroz J, Brandão H, Carvalho ED, Silva RJ (2013) Helminths of Stein-
dachnerina insculpta in two distinct stretches of the Taqueri River, state of São Pau-
lo, Brazil. Revista Brasileira de Parasitologia Veterinária 22(4): 539–547. https://doi.
org/10.1590/S1984-29612013000400014
Aguiar JCC, Bueno GBF, Santos SMC, Adriano EA (2017) Supplementary taxonomic
description of Demidospermus pinirampi (Monogenoidea, Dactylogyridae), with a
new host record and an expansion of its distribution range. Acta Amazonica 47(4):
355–358. https://doi.org/10.1590/1809-4392201700112
Bezerra CAM, Sousa AL, Viana DC (2020) Histopathologic alterations of gill tissue in
Siluriformes and Characiformes from the Middle Tocantins River in the Brazilian
Amazon. Arquivo Brasileiro de Medicina Veterinária e Zootecnia 72(1): 285–289.
https://doi.org/10.1590/1678-4162-11467
Boeger WA, Domingues MV, Kritsky DC (1997) Neotropical Monogenoidea. 32. Cacatu-
ocotyle paranaensis n.g., n.sp. (Dactylogyridae, Ancyrocephalinae) from Characidium
spp. (Teleostei, Characidae) from the State of Parana, Brazil. Systematic Parasitology
36: 75–78. https://doi.org/10.1023/A:1005796027406
Boeger WA, Vianna RT (2006) Monogenoidea. In: Thatcher VE (Ed.) Amazon Fish para-
sites. Pensoft Publishers, Soa, 42–116.
Boeger WA, Fehlauer KH, Marques EE (2006) Neotropical Monogenoidea. 49. Four new
species of the Diplectanidae (Dactylogyrinea) from the gills of some pachyurines
(Teleostei: Sciaenidae) from the Rio Tocantins and Rio Doce Basins, with proposal
of Anoplectanum n.g. and Spinomatrix n.g. Systematic Parasitology 64(1): 57–68.
https://doi.org/10.1007/s11230-005-9021-7
114
ZooKeys 1172: 101–116 (2023), DOI: 10.3897/zookeys.1172.105500
Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Braga MP, Araújo SBL, Boeger WA (2014) Patterns of interaction between Neotropical
freshwater shes and their gill Monogenoidea (Platyhelminthes). Parasitology Re-
search 113(2): 481–490. https://doi.org/10.1007/s00436-013-3677-8
Bush AO, Lafferty KD, Lotz JM, Shostak W (1997) Parasitology Meets Ecology on Its
Own Terms: Margolis et al. Revisited. The Journal of Parasitology 83(4): 575–583.
https://doi.org/10.2307/3284227
Cárdenas MQ, Fernandes BMM, Justo MCN, Cohen SC (2019) A new species of Ichthy-
ouris Inglis, 1968 (Nematoda: Pharyngodonidae) parasitizing two Characiform shes
from Tocantins River, Maranhão State, Brazil. Comparative Parasitology 86(1): 5–9.
https://doi.org/10.1654/1525-2647-86.1.5
Cárdenas MQ, Justo MCN, Viana DC, Cohen SC (2021) New host record and geograph-
ical distribution of Nematoda parasitizing Hypophthalmus marginatus Valenciennes
(Siluriformes) from the Tocantins River, Brazil. Acta Scientiarum. Biological Sciences
43: e58014. https://doi.org/10.4025/actascibiolsci.v43i1.58014
Cárdenas MQ, Justo MCN, Reyes ARP, Cohen SC (2022) Diversity of Nematoda and Di-
genea from different species of characiform shes from Tocantins River, Maranhão,
Brazil. Revista Brasileira de Parasitologia Veterinária 31(3): e005122. https://doi.
org/10.1590/s1984-29612022038
Carvalho EG, Seibert CS, Coelho MS, Marques EE (2009) Parâmetros hematológicos de
espécies nativas do rio Tocantins, Auchenipterus nuchalis, Psectrogaster amazonica
e Squaliforma emarginata (Teleostei, Ostariophysi). Acta Scientiarum. Biological Sci-
ences 31(2): 173–177. https://doi.org/10.4025/actascibiolsci.v31i2.1024
Ceschini TL, Takemoto RM, Yamada FH, Moreira LHA, Pavanelli GC (2010) Paranaella
luquei (Monogenea: Microcotylidae), an ectoparasite from the gills of Steindachner-
ina brevipinna, in the tributaries Corvo and Guairaca, Paranapanema River, Parana,
Brazil. Helminthologia 47(1): 29–32. https://doi.org/10.2478/s11687-010-0005-z
Cohen SC, Justo MCN, Kohn A (2013) South American Monogenoidea Parasites of Fish-
es, Amphibians and Reptiles. Conselho Nacional de Desenvolvimento Cientico e
Tecnologico (CNPq), 659 pp.
Cohen SC, Justo MCN, Gen DVS, Boeger WA (2020) Dactylogyridae (Monogenoidea, Polyon-
choinea) from the gills of Auchenipterus nuchalis (Siluriformes, Auchenipteridae) from the
Tocantins River, Brazil. Parasite 27(4): 1–12. https://doi.org/10.1051/parasite/2020002
Cohen SC, Cárdenas MQ, Justo MCN (2021) Checklist de Monogenoidea parasitos de
peixes Siluriformes do Brasil. In: Melchior LAK, Meneguetti DUO, Camargo LMA, Oliveira
J (Eds) Atualidades em Medicina Tropical na América do Sul: Veterinária (1st edn).
Editora Stricto Sensu, 28–48. https://doi.org/10.35170/ss.ed.9786586283594.02
Dias KGA, Vieira DHMD, Camargo AA, Silva RJ, Azevedo RK, Abdallah VD (2017) Diversi-
ty of monogeneans from Characiformes shes in the Batalha River and Peixe’s River,
state of Sao Paulo, Brazil. Neotropical Helminthology 11(12): 317–330.
Domingues MV, Boeger WA (2005) Neotropical Monogenoidea. 47. Phylogeny and co-
evolution of species of Rhinoxenus (Platyhelminthes, Monogenoidea, Dactylogyri-
dae) and their Characiformes hosts (Teleostei, Ostariophysi) with description of four
new species. Zoosystema 27(3): 441–467.
Domingues MV, Marques FPL (2011) Phylogeny and taxonomy of Potamotrygonocotyle
Mayes, Brooks, Thorson, 1981 (Monogenoidea: Monocotylidae) with a description of
four new species. Journal of Helminthology 85(4): 353–380. https://doi.org/10.1017/
S0022149X10000684
Fehlauer KH, Boeger WA (2005) Neotropical Monogenoidea: Euryhaliotrema dontyko-
leos sp. nov. (Dactylogyridae) from the gills of the freshwater sciaenid, Pachyurus
115
ZooKeys 1172: 101–116 (2023), DOI: 10.3897/zookeys.1172.105500
Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
junki (Perciformes). The Journal of Parasitology 91(5): 1025–1027. https://doi.
org/10.1645/GE-486R.1
Freitas AJB, Bezerra CAM, Meneses YC, Justo MCN, Viana DC, Cohen SC (2021) Three
new species of Urocleidoides (Monogenoidea: Dactylogyridae) parasitizing charac-
iforms (Actinopterygii: Characiformes) in Tocantins River, states of Tocantins and
Maranhão, and new record for U. triangulus in Guandu River, state of Rio de Janeiro,
Brazil. Zoologia 38: e65001. https://doi.org/10.3897/zoologia.38.e65001
Froese R, Pauly D [Eds] (2022) FishBase. World Wide Web electronic publication. www.
shbase.org, version (08/2022). [Accessed on 5 April 2022]
Hanek G, Molnár K, Fernando CH (1974) Three new genera of Dactylogyridae (Mono-
genea) from freshwater frishes of Trinidad. The Journal of Parasitology 60(6):
911–913. https://doi.org/10.2307/3278510
Justo MCN, Cohen SC (2019) Monogenoidea: Implicações na aquicultura e revisão das
principais alternativas de tratamento. In: Ranzani-Paiva MJT, Takemoto RM, Lizama
MAP, Perazzolo LM, Rosa RD (Eds) Biotecnologia e Sanidade de Organismos Aquáti-
cos. Abrapoa, São Paulo, 371–398.
Kritsky DC, Boeger WA (1995) Neotropical Monogenoidea. 26. Annulotrematoides
amazonicus, new genus and species (Dactylogyridae: Ancyrocephalinae), from the
gills of Psectrogaster rutiloides (Kner) (Teleostei: Characiformes: Curimatidae) from
the Brazilian Amazon. Proceedings of the Biological Society of Washington 108(3):
528–532. https://www.biodiversitylibrary.org/part/44854
Kritsky DC, Boeger WA, Mendoza-Franco EF, Vianna RT (2013) Neotropical Monog-
enoidea. 57. Revision and phylogenetic position of Scleroductus Jara & Cone, 1989
(Gyrodactylidae), with descriptions of new species from the Guatemalan Chulin
Rhamdia guatemalensis (Günther) (Siluriformes: Heptapteridae) in Mexico and the
barred sorubim Pseudoplatystoma fasciatum (Linnaeus) (Siluriformes: Pimelodidae)
in Brazil. Systematic Parasitology 84: 1–15. https://doi.org/10.1007/s11230-012-
9387-2
Kritsky DC, Leiby PD, Kayton RJ (1978) A rapid stain technique for the haptoral bars
of Gyrodactylus species (Monogenea). The Journal of Parasitology 64(1): 172–174.
https://doi.org/10.2307/3279642
Kritsky DC, Vianna RT, Boeger WA (2007) Neotropical Monogenoidea. 50. Oviparous
gyrodactylids from loricariid and pimelodid catshes in Brazil, with the proposal
of Phanerothecioides n. g.; Onychogyrodactylus n. g. and Aglaiogyrodactylus n. g.
(Polyonchoinea: Gyrodactylidea). Systematic Parasitology 66(1): 1–34. https://doi.
org/10.1007/s11230-006-9053-7
Lapera IM, Silva AC, Canônico BM, Perezin GF, Tebaldi JH, Pala G, Manrique WG, Hoppe
EGL (2017) Metazoan parasites of Plagioscion squamosisimus, an invasive species
in the Tietê River, São Paulo, Brazil. Revista Brasileira de Parasitologia Veterinária
26(2): 143–151. https://doi.org/10.1590/s1984-29612017035
Malabarba MC, Malabarba LR (2010) Biogeography of Characiformes: an evaluation of
the available information of fossil and extant taxa. In: Nelson JS, Schultze HP, Wilson
MVH (Eds) Origin and Phylogenetic Interrelationships of Teleosts. Verlag Dr. Friedrich
Pfeilp, Munique, 317–336.
Malabarba L, Malabarba MC (2014) Filogenia e classicação dos peixes Neotropicais.
In: Baldisserotto B, Cyrino JEB, Urbinati EC (Eds) Biologia e Fisiologia de Peixes Neo-
tropicais de Água Doce. FUNEP, Jaboticabal, 12 pp.
Melo BF, Sidlauskas BL, Hoekzema K, Vari RP, Dillman CB, Oliveira C (2018) Molecular
phylogenetics of Neotropical detritivorous shes of the family Curimatidae (Teleostei:
116
ZooKeys 1172: 101–116 (2023), DOI: 10.3897/zookeys.1172.105500
Carine A. M. Bezerra et al.: Two new species of Curvianchoratus
Characiformes). Molecular Phylogenetics and Evolution 127: 800–812. https://doi.
org/10.1016/j.ympev.2018.06.027
Molnár K, Hanek G, Fernando CH (1974) Ancyrocephalids (Monogenea) from fresh-
water shes of Trinidad. The Journal of Parasitology 60(6): 914–920. https://doi.
org/10.2307/3278511
Nascimento FL, Araújo-Lima C (1993) Descrição das larvas de Psectrogaster amazonica
e Potamorhina altamazonica (Curimatidae, Pisces) da Amazonia central. Acta Ama-
zonica 23(4): 457–472. https://doi.org/10.1590/1809-43921993234472
Pereira NJ, Santos MM, Maião JPLS, Campos JSP, Silva ND, Mendes DCS, Lenz TM, San-
tos DMS (2020) Biomarcadores histológicos em brânquias de peixes na avaliação
da contaminação ambiental do Rio Mearim, Nordeste brasileiro. Journal of Develop-
ment 6(8): 56600–56616. https://doi.org/10.34117/bjdv6n8-175
Ponte SCS, Silva AJS, Zacardi DM (2017) Áreas de dispersão e berçário para larvas de
Curimatidae (Pisces, Characiformes), no trecho baixo do rio Amazonas, Brasil. Inter-
ciencia 42(11): 727–732. https://www.redalyc.org/journal/339/33953499004/html/
Rossin MA, Timi JT (2016) Dactylogyrid monogeneans parasitising Cyphocharax voga
(Hensel) (Teleostei: Curimatidae) from the Pampas region, Argentina: new and pre-
viously described species. Systematic Parasitology 93(7): 701–714. https://doi.
org/10.1007/s11230-016-9654-8
Santos GM, Jegu M, Merona B (1984) Catálogo de Peixes Comerciais do Baixo rio To-
cantins (1st ed.). Projeto Tucuruí, Eletronorte, INPA, Manaus, 82 pp.
Suriano DM (1980) Notodiplocerus singularis gen. et sp. nov. (Monogenea Ancyrocephali-
nae) parasita de las branquias de Pseudocurimata gilbert (Pisces Tetragonopteridae)
de la laguna de Chascomus, República Argentina. Neotrópica 26: 131–142.
Suriano DM (1981) Androspira gen. n. (Monogenea Ancyrocephalinae) parasito bran-
quial de Pseudocurimata gilbert (Quoy and Gaimard, 1824) Fernandez-Yepes, 1948
(Pisces Tetragonopteridae) de La Laguna de Chascomus, Republica Argentina. Neo-
trópica 27: 67–78.
Suriano DM (1986) Philocorydoras platensis gen. n. et. sp. n. (Monogenea: Ancyro-
cephalidae) from Corydoras paleatus (Jenyns) (Pisces: Callichthyidae) in Laguna
Chascomús – Republica Argentina. Helminthologia 23(4): 249–256.
Suriano DM (1997) Palombitrema heteroancistrium Price & Bussing, 1968 (Monogenea:
Ancyrocephalidae) from Astyanax (A.) fasciatus (Cuvier, 1819) (Pisces: Characidae)
in Chascomus Lake, Argentina: Anatomy and systematic position. Physis 53: 7–10.
Tavares-Dias M, Silva LMA, Oliveira MSB (2022) Geographic range, distribution pat-
terns and interactions of Monogenea Van Beneden 1858, with species of native host
freshwater shes from Brazil. Revista Brasileira de Parasitologia Veterinária 31(3):
e005722. https://doi.org/10.1590/s1984-29612022048
Vari RP (1989) Systematics of the Neotropical Characiform Genus Psectrogaster Eigen-
mann and Eigenmann (Pisces: Characiformes). Smithsonian Institution Press, Wash-
ington, 43 pp. https://doi.org/10.5479/si.00810282.474
Vari R, Röpke C (2013) Curimatidae. In: Queiroz LJ, Torrente-Vilara G, Ohara WM, Pires T,
Zuanon JA, Doria CR (Eds) Peixes do rio Madeira. Dialeto, São Paulo, 113–140.
Vieira DH, Caramello LE, Abdallah VD, Silva RJ, Azevedo RK (2013) Community ecology
of metazoan parasites of the sairú Cyphocharax nagelii from the Peixe River. Revista
Brasileira de Parasitologia Veterinária 22(4): 611–615. https://doi.org/10.1590/
S1984-29612013000400027
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