Journal of Fish Biology (2016)

doi:10.1111/jfb.13184, available online at wileyonlinelibrary.com

A new species of the leopard pleco genus Pseudacanthicus

(Siluriformes: Loricariidae) from the Rio Xingu, Brazil

C. C. Chamon*† and L. M. Sousa‡

*Laboratório de Ictiologia Sistemática, Universidade Federal do Tocantins – UFT, Campus
Universitário de Porto Nacional, Setor Jardim dos Ipês, Rua 3, Quadra 17, s/no, Caixa
Postal 136, CEP: 77500-000, Tocantins, Brazil and ‡Laboratório de Ictiologia de
Altamira, Universidade Federal do Pará - UFPA, Campus Universitário de Altamira,
Faculdade de Ciências Biológicas, 68372-040, Altamira, PA, Brazil

(Received 20 August 2015, Accepted 16 September 2016)

The genus Pseudacanthicus comprises six valid species distributed in the Amazon basin and Caribbean
coastal drainages from Guyana to French Guiana: P. serratus, P. fordii, P. histrix, P. spinosus, P. leop-
ardus and P. pitanga. A new species of Pseudacanthicus is described from the Rio Xingu Basin,
distinguished from its congeners by the following combination of characters: presence of anastomose
dark blotches forming continuous zigzag bands alongside longitudinal keels; presence of dark blotches
on ventral surface of body and head; all fins with orange to red colour on unbranched rays and some-
times subsequent branched rays. Brief comments on ornamental fisheries and conservation of the new
species are also provided.
© 2016 The Fisheries Society of the British Isles

Key words: Amazon Basin; conservation; fish trade; Neotropical; taxonomy.

INTRODUCTION
The Loricariidae is the largest family among Siluriformes with >900 species described
(Eschmeyer & Fong, 2015), representing about a quarter of all catfish species (Reis
et al., 2003; Ferraris, 2007). Species of the family are widely distributed in the Neotrop-
ical region, from south-eastern Costa Rica to north-eastern Argentina (Isbrücker, 1980)
and occur in a large variety of freshwater environments.
Pseudacanthicus Bleeker 1862 is a member of the Ancistrini and included in the
Acanthicus Agassiz 1829 group (sensu Armbruster, 2004) or Acanthicus clade (sensu
Lujan et al., 2015), with Acanthicus, Megalancistrus Isbrücker 1980 and Leporacan-
thicus Isbrücker & Nijssen 1989 (clade 73). Members of this group are easily distin-
guished from other Loricariidae by the presence of rows of keels along the body formed
by sharply pointed odontodes. Other characteristics useful for identifying members
of the Acanthicus group are: eight or more dorsal-fin rays (v. six or seven in most
other loricariids except Pterygoplichthys Gill 1858, Chaetostoma Tschudi 1846 and
Pseudancistrus pectegenitor Lujan, Armbruster & Sabaj Pérez 2007); seven to eight

†Author to whom correspondence should be addressed. Tel.: +55 63 33630572; email:

chamon.carine@gmail.com

1
© 2016 The Fisheries Society of the British Isles
2 C. C. CHAMON AND L. M. SOUSA

infraorbitals (v. usually five or six); five anal-fin branched rays (v. usually four); and
presence of hypertrophied odontodes along the snout margin (v. hypertrophied odon-
todes absent in other loricariids except Panaque Eigenmann & Eigenmann 1889 and
Pseudancistrus Bleeker 1862). Within the Acanthicus group, Pseudacanthicus can be
distinguished by the presence of two small plates in the posterior area of the compound
pterotic (v. one median plate or plate absent), by the presence of narrow premaxillae
with a small number of elongate and curved teeth and other characters detailed below.
Pseudacanthicus comprises six valid species distributed in the Amazon Basin and
Caribbean coastal drainages from Guyana to French Guyana: Pseudacanthicus fordii
(Günther 1868) and Pseudacanthicus serratus (Valenciennes 1840), from coastal
drainages of Suriname; Pseudacanthicus histrix (Valenciennes 1840) and Pseudacan-
thicus spinosus (Castelnau 1855), in the River Negro and Amazon Basins; Pseudacan-
thicus leopardus (Fowler 1914), described from the River Rupununi Basin in Guyana;
Pseudacanthicus pitanga Chamon 2015, described from the Rio Tocantins Basin.
Chamon (2015) reported that the diversity of Pseudacanthicus is probably underes-
timated, with several undescribed species in Amazon tributaries. Accelerated growth
of hydroelectricity in the Amazon region, especially the Belo Monte Dam in Volta
Grande do Xingu, provided an impetus for describing one new species and discussing
its importance to ornamental fisheries and conservation.

MATERIALS AND METHODS

Measurements were made with digital calipers, point-to-point, on the left side of specimens
to the nearest 0·1 mm, following Armbruster (2003). Osteological nomenclature and vertebral
counts follow Schaefer (1987), Armbruster (2004) and Geerinckx & Adriaens (2006). Speci-
mens were cleared and stained (CS) according to the protocol of Taylor & Van Dyke (1985).
Vertebral counts include the first five vertebrae modified into the Weberian apparatus and fused
pre-ural centrum counted as a single element according to Lundberg & Baskin (1969). Terminol-
ogy for the laterosensory system of the head follows Arratia & Huaquin (1995); homologies of
the pre-opercular sensory canal follow Schaefer (1987). Data for material examined are provided
in the following order: institution and catalogue number, number of specimens examined, mini-
mum and maximum standard length (LS ), locality, geographical co-ordinates, date of collection
and collectors. Some lots lack data concerning date of collection and, or collectors. Specimens
examined belong to the following institutions: Academy of Natural Science of Philadelphia
(ANSP); Auburn University Museum (AUM); British Museum (Natural History) (BMNH);
Instituto Nacional de Pesquisa da Amazônia (INPA); Laboratório de Ictiologia de Altamira, Uni-
versidade Federal do Pará, Campus de Altamira (LIA); Muséum National d′ Histoire Naturelle
(MNHN); Museu de Zoologia da Universidade de São Paulo (MZUSP); Laboratório de Icti-
ologia Sistemática da Universidade Federal do Tocantins (UNT); Naturalis Biodiversity Center,
Leiden, which includes the collections of Rijksmuseum van Natuurlijke Historie (RMNH) and
Zoölogisch Museum Amsterdam (ZMA).

RESULTS

P S E U DAC A N T H I C U S P I R A R A R A N E W S P E C I E S
See Figs 1 and 2 and Table I.

Holotype
MZUSP 116932, 234·1 mm LS ; Rio Xingu, Senador José Porfírio, Pedral do Caitucá,
3∘ 34′ 24′′ S; 51∘ 51′ 33′′ W, 22 October 2013, A. Gonçalves, T. Albuquerque.

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
 N E W S P E C I E S O F P S E U DAC A N T H I C U S F RO M R I O X I N G U , B R A Z I L 3

Fig. 1. Holotype of Pseudacanthicus pirarara, Museu de Zoologia da Universidade de São Paulo – MZUSP
116932, 234·1 mm standard length (LS ), in lateral, dorsal and ventral view.

Paratypes
All from Brazil, Pará, Rio Xingu Basin: ANSP 194906, 2, 75·7–119·3 mm LS ;
Vitória do Xingu, along left bank of river, c. 8 km east of Vitória do Xingu (one of
presumably two dive sites in close proximity), 2∘ 53′ 18·70′′ S; 51∘ 56′ 24·54′′ W, 22
September 2013, M. H. Sabaj Pérez, L. M. Sousa, A. Gonçalves, N. K. Lujan, D.
B. Fitzgerald, P. Madoka Ito, A. Oliveira & R. Robles. ANSP 194992, 1, 208·5 mm
LS ; Altamira, channel of right major braid of Xingu, c. 15 km south-southeast of
Altamira, 3∘ 20′ 35·28′′ S; 52∘ 11′ 11·10′′ W, 12 September 2013, M. H. Sabaj Pérez,
L. M. Sousa, A. Gonçalves, N. K. Lujan, D. B. Fitzgerald, P. Madoka Ito, A. Oliveira
& R. Robles. ANSP 197624, 1, 58·6 mm LS ; Rio Xingu, central channel c. 4·5 km
upstream of campsite 3, 3∘ 24′ 58·5′′ S; 51∘ 42′ 38·7′′ W, 9 November 2014, M. H.

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
4 C. C. CHAMON AND L. M. SOUSA

(a) (b)

(c) (d)

(e) (f)

Fig. 2. Pseudacanthicus pirarara, live specimens showing variation in colour: (a) Laboratório de Ictiologia de
Altamira, Universidade Federal do Pará, Campus de Altamira – LIA 1466, 58·0 mm standard length (LS ),
Rio Xingu near São Félix do Xingu; (b) ANSP 197879, 137 mm LS , Rio Irini; (c) LIA 1251, 122·5 mm
LS , Rio Xingu near São Félix do Xingu; (d) Academy of Natural Science of Philadelphia – ANSP 193003,
193 mm LS , Rio Irini; (e) LIA 1297, 141·0 mm LS , Rio Xingu near São Félix do Xingu; (f) LIA (no number),
173·75 mm LS , Rio Xingu near Altamira.

Sabaj Pérez, L. M. Sousa, A. P. Gonçalves, D. B. Fitzgerald, V. Machado, P. M. Ito,

A. Oliveira, H. Gimênes Jr., M. Hardman, J. Tiemann, K. S. Cummings & M. C.
Dreher Mansur. AUM 65868, 1, 174·0 mm LS ; same data as ANSP 194906. INPA
4037, 1, 310·0 mm LS ; 1 October 1990, L. Rapp Py-Daniel & J. A. Zuanon. INPA
4038, 1, 169·56 mm LS ; 4 October 1990, L. Rapp Py-Daniel & J. A. Zuanon. INPA
31466, 4, 115·46–210·5 mm LS ; Vitória do Xingu, Itaobinha, bedrock on Rio Xingu,
3∘ 23′ 45′′ S; 52∘ 12′ 16′′ W, 4 November 2008, L. Rapp Py-Daniel et al. INPA 31776,
1, 230·5 mm LS ; Altamira, Ilha do Davi, 3∘ 32′ 39′′ S; 51∘ 57′ 29′′ W, 10 November
2008, L. Rapp Py-Daniel et al. INPA 31807, 2, 130·57–188·25 mm LS ; Babaquara,
Gorgulho da Rita, 3∘ 20′ 14′′ S; 52∘ 11′ 18′′ W, 7 November 2008, L. Rapp Py-Daniel
et al. LIA 1251, 1, 122·5 mm LS ; São Félix do Xingu, Remansinho, 6∘ 46′ 33′′ S;
51∘ 59′ 52′′ W, 23 September 2014, L. Sousa, A. Gonçalves & C. Martins. LIA 1341,
1, 132·9 mm LS ; São Félix do Xingu, Remansinho, 6∘ 46′ 33′′ S; 51∘ 59′ 52′′ W, 23
September 2014, L. Sousa, A. Gonçalves and C. Martins. LIA 1405, 1, 90·2 mm LS ;
Xadai, upstream São Félix do Xingu, 6∘ 53′ 50′′ S; 52∘ 02′ 10′′ W, 20 September 2014,
L. Sousa, A. Gonçalves & C. Martins. LIA 1466, 2, 52·3–57·9 mm LS ; São Félix
do Xingu, Araraquara, 6∘ 33′ 16′′ S; 52∘ 06′ 43′′ W, 25 September 2014, L. Sousa,
A. Gonçalves & C. Martins. LIA 1650, 4, 86·6–205·0 mm LS ; Vitória do Xingu,

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
 N E W S P E C I E S O F P S E U DAC A N T H I C U S F RO M R I O X I N G U , B R A Z I L 5

Table I. Morphometric and meristic characteristics of a sample of 43 Pseudacanthicus

pirarara

Holotype Range Mean s.d.

LS (mm) 234·1 55·7–310·0 – –
Percentage of LS
Predorsal length 40·8 38·8–46·0 43·0 1·5
Head length (LH ) 34·0 19·4–38·8 35·5 2·9
Body depth at dorsal origin 20·3 14·1–26·5 20·9 2·7
Cleithral width 28·6 19·6–32·4 29·8 1·9
Thorax length 22·4 19·0–29·7 24·7 2·3
Pectoral-spine length 33·4 29·2–37·4 32·5 2·5
Abdominal length 25·6 21·4–29·5 26·0 1·6
Pelvic-spine length 29·1 25·4–33·9 30·3 1·7
Postanal length 25·2 23·9–34·6 29·2 3·7
Anal-fin spine length 19·9 14·5–24·2 20·1 1·8
Dorsal-spine length 32·4 26·6–38·1 33·6 2·7
Dorsal-fin base length 25·6 24·0–29·3 26·8 1·3
Caudal-peduncle depth 13·2 11·5–15·2 13·6 0·8
Percentage of LH
Orbit diameter 12·1 7·6–22·0 12·4 2·4
Snout length 65·7 56·7–68·2 62·4 2·6
Internares width 15·0 10·9–23·5 14·3 2·0
Interorbital width 32·1 28·5–55·3 31·5 4·1
Head depth 57·6 33·4–62·3 52·2 5·2
Dentary tooth crown length 7·9 3·5–14·5 8·6 2·0
Premaxillary tooth crown length 4·3 2·2–8·5 5·0 1·4
Meristics Mode
Dorsal-plate series 26 24–27 26 –
Mid-dorsal plate series 26 24–27 25 –
Median-plate series 25 22–27 26 –
Mid-ventral-plate series 25 24–26 25 –
Ventral-plate series 18 11–19 18 –
Plates base of caudal 8 7–8 7 –
Premaxillary teeth 10 4–12 8 –
Dentary teeth 14 7–21 14 –
LS , standard length.

Ensecadeira do Pimental, 3∘ 22′ 53′′ S; 51∘ 56′ 38′′ W, 1 October 2012, Biota ichthy-
ological team. LIA 1651, 2, 128·8–133·9 mm LS ; Vitória do Xingu, Ensecadeira do
Pimental, 3∘ 22′ 53′′ S; 51∘ 56′ 38′′ W, 1 December 2012, Biota ichthyological team.
LIA 1652, 1, 195·7 mm LS ; Altamira, Gorgulho da Rita, 3∘ 20′ 23′′ S; 52∘ 11′ 25′′ W,
13 July 2013, LIA ichthyological team. LIA 1653, 1, 182·7 mm LS ; Senador José
Porfírio, Caitucá, 3∘ 34′ 25′′ S; 51∘ 51′ 34′′ W, 21 July 2013, A. Gonçalves, Alejandro
& T. Barbosa. LIA 1654, 1, 131·6 mm LS ; Altamira, Gorgulho da Rita, 3∘ 20′ 35′′ S;
52∘ 11′ 12′′ W, 14 October 2013, A. Gonçalves. LIA 1655, 257·0 mm LS ; Altamira,
Gorgulho da Rita, 3∘ 20′ 13′′ S; 52∘ 11′ 02′′ W, 5 July 2012, T. Barbosa. LIA 1656, 1,
169·2 mm LS ; Altamira, Gorgulho da Rita, 3∘ 20′ 20′′ S; 52∘ 11′ 21′′ W, 19 November
2012, T. Barbosa. LIA 1657, 1, 253·1 mm LS ; Senador José Porfírio, Ilha da Fazenda,

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
6 C. C. CHAMON AND L. M. SOUSA

3∘ 35′ 31′′ S; 51∘ 53′ 01′′ W, 19 September 2012, T. Barbosa. LIA 1659, 1, 132·6 mm
LS ; Altamira, Cajueiro, 3∘ 16′ 34′′ S; 52∘ 05′ 08′′ W, 18 September 2012, D. Bastos &
J. Santos. LIA 1660, 1, 161·1 mm LS ; Vitória do Xingu, bedrock in front of Garimpo
Galo, 3∘ 34′ 51′′ S; 51∘ 54′ 29′′ W, 24 November 2012, A. Gonçalves. MZUSP
107172, 1, 246·5 mm LS ; Paratinzinho, Rio Xingu, Altamira, Pará, Brasil (collected by
ornamental fishermen), 3∘ 14′ 59′′ S; 52∘ 5′ 56·9′′ W, 8 July 2010; O. T. Oyakawa, J.
Muriel-Cunha, C. C. Chamon, I. Fichberg, L. Rossi & A. Sawakuchi. MZUSP 107175,
7, 38·24–184·15 mm LS ; Altamira, Gorgulho da Rita, 3∘ 20′ 26·6′′ S; 52∘ 11′ 4·5′′ W,
6 July 2010, O. T. Oyakawa, J. Muriel-Cunha, C. C. Chamon, I. Fichberg, L. Rossi
& A. Sawakuchi . MZUSP 108566 (two skeletons), 122·2–127·7 mm LS ; Altamira,
Morro do Forte, 3∘ 12′ 18′′ S; 52∘ 11′ 29·1′′ W, 8 July 2010, O. T. Oyakawa, J.
Muriel-Cunha, C. C. Chamon, I. Fichberg, L. Rossi, A. Sawakuchi & ornamental fish-
ermen. MZUSP 108567, 1, 261·7 mm LS ; Altamira, near Morro do Forte, 3∘ 12′ 18′′ S;
52∘ 11′ 29·1′′ W (approximate co-ordinates), 8 July 2010, purchased from ornamental
fisherman by O. T. Oyakawa, J. Muriel-Cunha, C. C. Chamon, I. Fichberg, L. Rossi
& A. Sawakuchi. MZUSP 108572, 1, 138·98 mm LS ; Belo Monte, Pontoon in front
of D. M. and Sr. Valdomiro’s property, 3∘ 06′ 45·5′′ S; 51∘ 43′ 220·9′′ W, 12 July
2010, O. T. Oyakawa, J. Muriel-Cunha, C. C. Chamon, I. Fichberg, L. Rossi & A.
Sawakuchi. UNT 14635 (ex LIA 1297), 1, 140·9 mm LS ; São Félix do Xingu, Pedra
Preta, 6∘ 57′ 31′′ S; 52∘ 15′ 27′′ W, 22 September 2014, L. Sousa, A. Gonçalves &
C. Martins.
Rio Iriri: ANSP 193003, 1, 193·0 mm LS ; Rio Iriri, Cachoeira Grande do Iriri, c.
15 km upstream from confluence with Rio Xingu, 3∘ 50′ 35·5′′ S; 52∘ 44′ 8·3′′ W, 10
October 2012, M. H. Sabaj Pérez, M. Arce & L. Sousa. ANSP 197879, 1, 137·0 mm LS ;
Rio Iriri, cachoeira Grande do Iriri, c. 15 km upstream from confluence with Rio Xingu,
3∘ 50′ 32·3′′ S; 52∘ 44′ 03·9′′ W, 2 November 2014, M. H. Sabaj Pérez, L. M. Sousa, A.
P. Gonçalves, D. B. Fitzgerald, V. Machado, P. M. Ito, A. Oliveira, R. Oliveira, H.
Gimênes Jr., A. Carvalho-Batista, M. Hardman & K. Winemiller.
Rio Bacajá: LIA 1658, 1, 200·5 mm LS ; Rio Bacajá, região do Pariaxá, 3∘ 34′ 37′′ S;
51∘ 35′ 36′′ W, 24 September 2012, D. Bastos & J. Santos.
Non-type: LIA (GEPE2011102801), 3, 95·5–203·7 mm LS . LIA (GEPE201110
2001), 1, 239·3 mm LS . ZMA no catalog, 3, 233·88–248·41 mm LS .

Diagnosis
Pseudacanthicus pirarara is distinguished from congeners (except P. leopardus and
P. pitanga) by having fins with intensely orange to red colouration (v. fins with dark
background colour with white spots in P. serratus and P. fordii or grey background
colour with black blotches in P. histrix and P. spinosus). Pseudacanthicus pirarara is
distinguished from P. leopardus by the presence of anastomose dark blotches forming
continuous zigzag bands alongside longitudinal keels and having all fins with strong
red colour on unbranched rays and sometimes in subsequent branched rays (v. dark
blotches distinct, never connected and red colour restricted to dorsal and caudal-fin
rays). From P. pitanga, P. pirarara is distinguished by the presence of conspicuous
dark blotches on head and abdomen (v. dark blotches usually absent or otherwise faint
on head and always absent on abdomen). Pseudacanthicus pirarara is further diag-
nosed from congeners by the following combination of osteological characteristics:
nasal bone L-shaped (v. rectangular); ventral crest of basipterygii straight (v. rounded

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
 N E W S P E C I E S O F P S E U DAC A N T H I C U S F RO M R I O X I N G U , B R A Z I L 7

or triangular); internal crest of posterior process of basipterygii well-developed (v. crest

absent or weak).

Description
Morphometric and meristic data are summarized in Table I. Dorsal profile of body
slightly convex from tip of snout to vertical through dorsal-fin origin; concave, nearly
straight from that point to caudal-fin origin. Ventral profile of body straight from snout
tip to caudal-fin origin. Ventral surface from tip of snout to urogenital papilla lack-
ing plates except for a few small plates at pectoral and pelvic-fin origins. Greatest
body width at pectoral girdle. Trunk strongly keeled; five rows of longitudinal keels;
one keel along each series of body plates. Body depth greatest at dorsal-fin origin,
shallowest at caudal peduncle, between adipose fin and first procurrent caudal-fin ray.
Head deep, pointed anteriorly; snout and cheek completely covered by numerous small
plates, except for small naked area on tip of snout. Snout very pointed in dorsal view.
Nasal elongate, L-shaped. Frontal short, broadly contacting nares anteriorly and orbit
posteriorly. Anterior margin of frontal short, reaching posterior margin or half length,
of nares. Parieto-supraoccipital elongate, posterior edge narrow, with V-shaped crest
formed by conspicuous odontodes. Sphenotic short, not contacting sixth infraorbital
and lacking conspicuous odontodes. Orbit dorsolateral, moderate in size (7·6–22·0%
head length, LH ). Iris with small dorsal flap over pupil. Compound pterotic short with
few fenestrae; anterior process briefly contacting posterior margin of orbit. Posterior
area of compound pterotic with one or two small plates. Infraorbital series with eight
pores. Fourth infraorbital widely contacting posterior margin of orbit. Sixth infraorbital
contributing only to the posteroventral margin of orbit. Lateral-line pores restricted to
hypural plate.
Mouth moderate in size, nearly as long as wide. Lips large, covered with papillae;
size of papillae decreasing towards posterior margin of lower lip; central buccal papilla
absent or weakly developed. Upper lip folded over itself. Maxillary barbel short; base
of barbel united to lips, tip free. Lower lip not reaching anterior margin of coracoid.
Medial end of premaxillary teeth series curved inwards. Premaxillae and dentary nar-
row and elongate. Dentary strongly curved inwards. Teeth well-developed, slightly
thickened with long crown and large lateral cusp; distal edge slightly curved inward.
Four to five pairs of well-developed predorsal plates. Cheek plates evertible with hyper-
trophied odontodes.
Dorsal-fin rays I,8; spinelet v-shaped with locking mechanism. Pectoral and pelvic
fins well-developed, medial portion much expanded relative to base; distal margin
rounded. Pectoral-fin rays I,6; unbranched ray covered with conspicuous odontodes.
Tip of adpressed pectoral fin almost reaching vertical through medial, unbranched
pelvic-fin ray. Pelvic-fin rays i,5; pelvic-fin spine reaching vertical through anal-fin
base when adpressed. Anal-fin rays i,5. Caudal fin i,14,i, truncate; supracaudal plates
seven to eight. Usually four procurrent caudal-fin rays, on each dorsal and ventral lobe.
Caudal peduncle deep. Total vertebrae 29, precaudal 11. Sixth rib strongly thickened,
remaining ribs slender.

Colour in life
Colour pattern highly variable in life (Fig. 2). Most individuals have light brown
background with dark blotches covering head, entire body including abdomen and fins;

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
8 C. C. CHAMON AND L. M. SOUSA

dark blotches forming continuous zigzag bands alongside longitudinal keels. Some
specimens with dark background colour on dorsal surface and blotches delimited only
on ventral surface and on dorsal and caudal fins. All fins usually with intense orange
or almost red colouration that is most evident on unbranched rays. In some specimens
the caudal fin is almost completely orange to red with few irregular blotches.

Colour in alcohol
Colour pattern similar, but less intense in alcohol; reddish-orange colour in fins
largely faded, more so over time.

Etymology
The specific name (a noun in apposition) is an allusion to Pirarara, the Brazilian com-
mon name of the red tail catfish, Phractocephalus hemioliopterus (Bloch & Schneider
1801), which was incorporated by local fishermen in the common name of the species
(i.e. assacu-pirarara) due to its reddish fins. Assacu is a common name for the Amazon
tree Hura crepitans (Euphorbiaceae), which has bark covered with conical spines.

Distribution
Pseudacanthicus pirarara is endemic to the Rio Xingu Basin and has been collected
in the main channel from Vitória do Xingu to São Félix do Xingu and major tributaries
to the Xingu like Rio Bacajá and Rio Iriri (Fig. 3).

Ecological notes
The species can be found in the main channel of the river, in moderate to strong
currents and shallow to deep environments down to 15 m. It is usually collected at
depths of 2–3 m, often hiding beneath bedrock or conglomerate plates of rocks and
gravel (Figs 4 and 5). Pseudacanthicus pirarara is not a gregarious species and it is
not usually found with conspecific individuals or other species. They are carnivorous
and feed on small invertebrates. According to local fishermen, they are more easily
found during the high-water season (December to April), probably due to reproductive
behaviour.

Fisheries and economical importance

Pseudacanthicus pirarara is highly appreciated by advanced hobbyists in the inter-
national aquarium trade because it is a large and colourful species that requires special
tank set-ups to best suit their needs. It is also an important resource for local fisher-
men in the Rio Xingu, being one of the more valuable species depending on season.
Local fishermen refer to it as assacu-vermelho, assacu-pirarara or L025 (Schraml &
Schäfer, 2004; Stawikowski et al., 2004; Werner et al., 2005). It is usually captured
with the aid of an air compressor and hoses that supply breathable air to the fishermen
during 20–30 min dives. The safety procedures for such dives, however, are far from
satisfactory.

DISCUSSION
The genus Pseudacanthicus is a member of the Acanthicus group (sensu Arm-
bruster, 2004: clade 73), wherein Leporacanthicus is sister to a clade composed of

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
 N E W S P E C I E S O F P S E U DAC A N T H I C U S F RO M R I O X I N G U , B R A Z I L 9

1°

2°

gu
Xin
Rio
Vitória do Xingu
3°
Altamira
i

4°
rir
oI
Ri

u
Xing
Rio

5°

6°

São Félix do Xingu

7°

8°

55° 54° 53° 52° 51°

Fig. 3. Distribution map of Pseudacanthicus pirarara, , collecting sites; , type-locality; , towns.

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
10 C. C. CHAMON AND L. M. SOUSA

Fig. 4. Typical habitat where Pseudacanthicus pirarara occurs: (a) stretch of the Rio Xingu at Gorgulho da Rita
region, a shallow, fast-flowing portion of the river with gravel and conglomerate rocks on the bottom; (b)
and (c) Pseudacanthicus pirarara in natural habitat at Gorgulho da Rita.

Pseudacanthicus and Acanthicus + Megalancistrus. More recently, in a molecular

phylogenetic analysis, Lujan et al. (2015) found these genera in their Acanthicus
clade, in which Pseudacanthicus (including P. pirarara, L025) was sister to Lepora-
canthicus, with Acanthicus and Megalancistrus forming respective sister groups. In
this study, Pseudacanthicus pirarara exhibits >9·8% cytb sequence divergence from
P. leopardus (N. Lujan, pers. comm.). The Acanthicus group is very well supported
in both molecular (Lujan et al., 2015) and morphological analyses (Armbruster,
2004). According to Armbruster (2004), the Acanthicus group is supported by 11
morphological synapomorphies, the most remarkable of which is the presence of
well-developed keels with sharp odontodes along the body. Pseudacanthicus can be
distinguished from the remaining genera of this Acanthicus group by the presence of
two small plates in the posterior area of the compound pterotic (v. just one median
plate or plate absent). It can also be distinguished from Acanthicus by the presence
of an adipose fin; a much smaller compound pterotic, a shorter dentary and narrower,
more longitudinally elongate premaxillae. It is distinguished from Megalancistrus
by fewer dorsal-fin rays (8–9 v. 10–11); from Leporacanthicus by having more
premaxillary teeth (up to 12 v. up to 4) and by having caudal filaments at least in
juveniles (Chamon, 2015).

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
 N E W S P E C I E S O F P S E U DAC A N T H I C U S F RO M R I O X I N G U , B R A Z I L 11

Fig. 5. Rapid at Cachoeira Grande do Iriri, Rio Iriri, near confluence of Rio Xingu showing fast-flowing habitat
of Pseudacanthicus pirarara.

Species of Pseudacanthicus are very conservative in shape and meristic data. All
measurements and meristic data of species of Pseudacanthicus are overlapping; how-
ever, colour patterns serve to distinguish the species. Pseudacanthicus pirarara is dis-
tributed along the middle and lower Rio Xingu and in the Rio Bacajá and Rio Iriri
Sub-basins. According to Camargo et al. (2004), the middle course of the Rio Xingu
extends from the Xingu–Irirí confluence to the Belo Monte waterfalls while the lower
course extends from Belo Monte falls downstream to the mouth of the Rio Xingu.
The colour pattern is highly variable among specimens, perhaps due to different envi-
ronmental conditions throughout its distribution. The majority of specimens have a
light brown background with dark blotches covering the head, entire body surface and
fins; dark blotches connect to form continuous zigzag bands alongside longitudinal
keels; all fins have orange to red colouration at least at the unbranched and subse-
quent rays. This pattern occurs often in specimens from Gorgulho da Rita, Babaquara,
Ilha da Fazenda and Vitória do Xingu. Specimens from Cotovelo, Belo Monte and Rio
Bacajá usually have a dark background colour and blotches delimited only on the ven-
tral surface and on the dorsal and caudal fins. Specimens from São Félix do Xingu
often possess dorsal and caudal fins that are almost completely orange to red with few
dark blotches, whereas specimens from the Rio Iriri show the opposite: all fins almost
completely covered with dark blotches and the orange to red colour is faint, restricted
to the unbranched dorsal and caudal-fin rays. Although there seems to be a correlation
between colour pattern and some localities, all colour types can be found at all locali-
ties; only the frequency of a particular pattern varies for a given place. For example, it
is possible to find specimens with dorsal and caudal fins almost completely orange to
red in Belo Monte and specimens having body surfaces with a dark background colour
and faint orange fins in São Félix do Xingu.

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
12 C. C. CHAMON AND L. M. SOUSA

Pseudacanthicus pirarara is an important ornamental fish and commercial exploita-

tion has been common practice in the middle to lower Rio Xingu since the 1980s.
Such activities are economically profitable in some local communities and in many
cases, constitute the entire source of income for multiple families (Camargo et al.,
2011; Chamon & Rapp Py-Daniel, 2014). Contrary to the understanding of most peo-
ple, ornamental fishing activities do not exert a direct effect on the health of such fish
communities (Andrews, 1990; pers. obs.). According to Camargo et al. (2011), the
ornamental fishery is less detrimental than more serious environmental problems that
occur in the region, such as mining, deforestation and the construction of hydropower
dams.
In the Volta Grande do Xingu rapids, located below Altamira, the river falls 96 m,
quadruples its width and flows through several waterfalls and around countless islands.
In this region is concentrated much of the hydroelectric potential of the river (Goulding
et al., 2003; ISA, Instituto Socioambiental, 2015). The rapids of the Volta Grande are
home to several endemic species of loricariids, almost all with ornamental interest,
such as Leporacanthicus heterodon Isbrücker & Nijssen 1989, Hypancistrus zebra
Isbrücker & Nijssen 1991 and Scobinancistrus aureatus Burgess 1994, as well as the
more recently described Parancistrus nudiventris Rapp Py-Daniel & Zuanon 2005,
Baryancistrus xanthellus Rapp Py-Daniel, Zuanon & Ribeiro de Oliveira 2011 and
Baryancistrus chrysolomus Rapp Py-Daniel, Zuanon & Ribeiro de Oliveira 2011,
Ancistomus feldbergae (de Oliveira, Rapp Py-Daniel, Zuanon & Rocha 2012), Spec-
tracanthicus zuanoni Chamon & Rapp Py-Daniel 2014; and other still undescribed
species.
According to Winemiller et al. (2016), in the tropical region, the construction of large
dams is directly correlated with the loss of fish fauna diversity by blocking genetic flow
between populations and by the difficulty for migratory species to complete their life
cycles. Dam construction is a danger to the majority of the ornamental loricariids in
the Amazon Basin. The Rio Xingu, like other Amazon tributaries, is suffering from
an accelerated rate of dam construction and species that are naturally rare or endemic
to specific habitats may suffer drastic population declines and even become extinct
in nature (Rapp Py-Daniel et al., 2011; Chamon & Rapp Py-Daniel, 2014). Scientific
understanding of different aspects of the fish fauna of Amazon drainages, including
their taxonomy and biology, has not kept pace. In the case of the Rio Xingu, the Belo
Monte Dam project includes a major impoundment dam located 40 km downstream
of the city of Altamira that will form the Xingu reservoir. As a result, rapids in the
main channel will be flooded and transformed to a lentic environment. Fishes that live
in lotic environments, like P. pirarara, are not adapted to slow flowing or still waters,
which are typically lower in oxygen (Chamon & Rapp Py-Daniel, 2014; L. M. Sousa,
pers. obs.).
In the lower course of the Rio Tocantins, Santos et al. (2004) reported declines in the
populations of 22 species of fishes after construction of the Tucuruí dam. Pseudacan-
thicus pitanga was among the loricariids included in their report. In the Marabá region,
however, P. pitanga is still considered a relatively common ornamental resource, sug-
gesting that the Tucuruí dam did not affect this species in areas upstream, remote from
the affected area. There is no information on the occurrence of P. pitanga in upper and
middle Rio Tocantins after 2008 (Chamon, 2015).
Sub-populations of P. pirarara are predicted to decrease or even disappear in Volta
Grande, especially in the area that will be flooded. Subpopulations from Rio Iriri and

© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184
 N E W S P E C I E S O F P S E U DAC A N T H I C U S F RO M R I O X I N G U , B R A Z I L 13

Rio Xingu around São Félix do Xingu, however, may suffer fewer negative influences
due to the Belo Monte dam project. Future monitoring of this and other loricariid
species is necessary to test those predictions.

C O M PA R AT I V E M AT E R I A L
Pseudacanthicus leopardus: ANSP 39345, holotype 71·6 mm LS ; AUM 35738, 1 C
S; 41·1 mm LS ; INPA 25865, 1 C S; 54·1 mm LS . Pseudacanthicus serratus: RMNH
3125, holotype, 148·7 mm LS . Pseudacanthicus fordii: BMNH 1866.8.14.148–152,
syntypes, 4, 117·1–145·0 mm LS . Pseudacanthicus spinosus: MNHN A-9577, holo-
type, 262·6 LS ; MZUSP 23992, 1 C S; 83·3 mm LS ; MZUSP 24010, 1 C S; 60·2 mm
LS ; MZUSP 108573, 1 skeleton, 119·8 mm LS . Pseudacanthicus pitanga: INPA 4502,
1, 164·1 mm LS ; INPA 4558, 4, 91·4–147·5 mm LS ; INPA 4559, 1, 148·1 mm LS ;
MZUSP 24135, 1, 110·7 mm LS ; MZUSP 34295, 6, 170·3–256·9 mm LS ; UNT 857,
1, 170·1 mm LS ; UNT 960, 1, 200·7 mm LS ; UNT 967, 1, 87·3 mm LS ; UNT 8505, 1,
259·0 mm LS ; UNT 9061, 1, 224·2 mm LS ; UNT 1109, 1, 146·8 mm LS ; UNT 10297,
1, 228·5 mm LS ; ZMA 119·395, 3, 88·5–89·2 mm LS ; ZMA 119·829, 1, 82·8 mm LS .
Pseudacanthicus cf . histrix (Xingu): INPA 31804, 1 C S; 71·1 mm LS . Pseudacanthicus
sp. Tapajós: MZUSP 108195, 1 skeleton, 109·2 mm LS .
Authors wish to thank M. S. Pérez and J. Lundberg (ANSP), J. Armbruster (AUM), P. Camp-
bell (BMNH), L. Rapp Py-Daniel and J. Zuanon (INPA), R. Causse (MNHN), M. Gianetti and
O. Oyakawa (MZUSP), P. Lucinda (UNT), R. de Ruiter (RMNH) and H. Praagman (ZMA) for
hospitality during visits to the fish collections under their care, for loan of specimens and for col-
lection data. Thanks are also due to M. S. Pérez for suggestions, comments on the manuscript and
permission to use his image (Fig. 5) from the environment. Fieldwork was supported in part by
iXingu Project, NSF DEB-1257813 (M. Sabaj Pérez, J. Lundberg and K. Winemiller, Co-PIs),
Fundação de Amparo à Pesquisa do Estado de São Paulo, FAPESP, process 2007/07770-5
(C.C. Chamon) and by CNPq process 486376/2013-3 (L. M. Sousa).

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© 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.13184