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A Taxonomic Revision of Sturisomatichthys Isbrücker and Nijssen, 1979 (Loricariidae: Loricariinae), with Descriptions of Three New Species

December 2019
Copeia 107(4):764

DOI:10.1643/CI-19-226

Authors:

Alejandro Londoño-Burbano

Pontifical Catholic University of Rio Grande do Sul

Roberto E. Reis

Pontifical Catholic University of Rio Grande do Sul

A taxonomic revision of Sturisomatichthys is presented. A total of 383 specimens including both type and non-type specimens of all valid species were examined. Sturisomatichthys was found to encompass ten valid species: Sturisomatichthys aureus, S. caquetae, S. citurensis, S. dariensis, S. festivus, S. frenatus, S. kneri, S. leightoni, S. panamensis, and S. tamanae. In addition, three new species are described: Sturisomatichthys reinae, new species, from the Baud´o River, in northwestern Colombia, belonging to the Caribbean slope of the continent; Sturisomatichthys guaitipan, new species, from the upper and middle Magdalena River basin, Colombia; and Sturisomatichthys varii, new species, from the San Juan River basin in western Colombia, on the Pacific slope. Sturisomatichthys caquetae, from the Morelia River, upper Amazon, is transferred back from Sturisoma, and S. leightoni was found to be distributed, besides the Magdalena-Cauca basin, in the Orinoco basin; thus, Sturisomatichthys is shown to be distributed in both trans- and cis-Andean drainages. A neotype is designated for S. aureus from the lower Magdalena basin. New records of localities in the northwestern region of South America, as well as the redescription and taxonomic comments on all valid species, are provided. Maps with the species distributions and a key for identification of the species are provided.

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A Taxonomic Revision of Sturisomatichthys Isbr¨

ucker and Nijssen, 1979

(Loricariidae: Loricariinae), with Descriptions of Three New Species

Alejandro Londo˜

no-Burbano

and Roberto E. Reis

A taxonomic revision of Sturisomatichthys is presented. A total of 383 specimens including both type and non-type

specimens of all valid species were examined. Sturisomatichthys wasfoundtoencompasstenvalidspecies:

Sturisomatichthys aureus,S.caquetae,S.citurensis,S.dariensis,S.festivus,S.frenatus,S.kneri,S.leightoni,S.panamensis,

and S.tamanae. In addition, three new species are described: Sturisomatichthys reinae, new species, from the Baud ´

o River,

in northwestern Colombia, belonging to the Caribbean slope of the continent; Sturisomatichthys guaitipan, new species,

from the upper and middle Magdalena River basin, Colombia; and Sturisomatichthys varii, new species, from the San Juan

River basin in western Colombia, on the Pacific slope. Sturisomatichthys caquetae, from the Morelia River, upper Amazon,

is transferred back from Sturisoma, and S. leightoni was found to be distributed, besides the Magdalena-Cauca basin, in

the Orinoco basin; thus, Sturisomatichthys is shown to be distributed in both trans- and cis-Andean drainages. A neotype

is designated for S.aureus from the lower Magdalena basin. New records of localities in the northwestern region of

South America, as well as the redescription and taxonomic comments on all valid species, are provided. Maps with the

species distributions and a key for identification of the species are provided.

LORICARIINAE currently includes 249 valid species

(Fricke et al., 2019a) classified in 31 genera (Covain et

al., 2016) and is distributed from the La Plata River in

southern Argentina to the northern Caribbean basins

(Ferraris, 2003). The species of this subfamily are character-

ized by possessing a long and depressed caudal peduncle and

the lack of an adipose fin. They usually live in the substrate

and show marked variations in the shape of their body due to

the different colonized environments, from lotic to lentic

systems, in organic or inorganic substrates such as rocks, logs,

sand, or mud (Covain et al., 2008).

Isbr¨

ucker and Nijssen (in Isbr¨

ucker, 1979) suggested a

classification of the Loricariinae that included several tribes

and subtribes, while describing several new genera and

species. Sturisomatichthys was described in that study as part

of the Harttiini along with Cteniloricaria,Harttia,Harttiella,

Lamontichthys,Metaloricaria, and Sturisoma. Sturisomatichthys

included four species, S. leightoni (type species) from the

Magdalena River basin, Colombia, S. caquetae from the

Morelia River, Amazon basin, Colombia, S. citurensis from

the Tuyra River, Panama, and S. tamanae from the San Juan

River basin, Colombia. The genus was compared to Sturisoma

from which, according to Isbr¨

ucker and Nijssen (in Isbr¨

ucker,

1979), it was distinguished by the absence of an enlarged

rostrum, which is present in Sturisoma, as the only diagnostic

character. The authors offered no further comments regard-

ing the genus.

Sturisomatichthys has remained as a poorly studied group

within the Loricariinae, and to date it has not received a

taxonomic revision. It was not until Rapp Py-Daniel (1997),

who carried out a morphology-based phylogeny of the

Loricariinae, that a representative of the genus was included

in an analysis of the subfamily. Sturisomatichthys citurensis

was found to be part of her Harttiini, as sister to the

remaining genera included in her Harttiina and Farlowellina,

more closely related to Farlowella.

Subsequently, Ghazzi (2003), in a morphology-based

phylogeny and taxonomic revision of Sturisoma, included

three species of Sturisomatichthys,S. citurensis,S. leightoni, and

S. tamanae. The author found S. citurensis and S. tamanae to

be closely related to a clade that included S. leightoni, while

three undescribed (trans-Andean) species revealed by the

author, plus S. panamensis and S. festivus, did appear in the

clade containing the type species of Sturisomatichthys. The

author proposed a ‘‘ ‘Trans’ New Genus’’ to include the

undescribed species plus S.festivus,S.frenatus,andS.

panamensis.

Covain et al. (2008) and Rodriguez et al. (2011) offered

DNA-sequence based phylogenies of the Loricariinae in

which S. citurensis was included. Both studies found S.

citurensis sister to Farlowella. Additionally, Covain et al.

(2008) included an analysis of morphological traits within

Loricariinae and tested their validity as informative charac-

ters. Regarding the only diagnostic character proposed for

Sturisomatichthys (absence of an enlarged rostrum), Covain et

al. (2008) found it to be not informative, and they suggested

that a deeper study of the genus was needed. They suggested

that the genus may represent a synonym of Sturisoma; this

was previously suggested by Covain and Fisch-Muller (2007).

Covain et al. (2016) included both S. citurensis and S.

leightoni in their DNA-based phylogeny of the Loricariinae.

These authors found S. leightoni nested within a clade

comprising S. aureus,S. citurensis,S. dariensis,S. festivus,S.

frenatus, and S. panamensis; those species, except S. citurensis,

belong to Sturisoma sensu lato and are all trans-Andean

species. For this reason, the authors suggested the above-

mentioned species also belong to Sturisomatichthys. Never-

theless, Covain et al. (2016) proposed that due to the trans-

Andean distribution of the new configuration of Sturisoma-

tichthys,S. caquetae, the only cis-Andean species until that

date, should be transferred to Sturisoma, assumed by the

authors to be fully cis-Andean. However, the authors did not

include S. caquetae in their analysis, nor did they examine its

holotype (ANSP 71719). They suggested a new generic

placement for the species only based on its distribution. In

the same way, the authors suggested S. kneri and S. tamanae

belong in Sturisomatichthys based on their distribution, but

none of the species were included in their study. Accordingly,

Covain et al. (2016) suggested Sturisomatichthys includes S.

aureus,S. citurensis,S. dariensis,S. festivus,S. frenatus,S. kneri,

S. leightoni,S. panamensis, and S. tamanae. Further, Sturisoma

comprises the remaining cis-Andean species of Sturisoma

PUCRS, Laborat´

orio de Sistema

´tica de Vertebrados, Av. Ipiranga, 6681, 90619-900 Porto Alegre, RS, Brazil; Email: (ALB) alondonoburbano@

gmail.com; and (RER) reis@pucrs.br. Send reprint requests to ALB.

Submitted: 26 April 2019. Accepted: 18 August 2019. Associate Editor: W. L. Smith.

Ó2019 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CI-19-226 Published online: 3 December 2019

Copeia 107, No. 4, 2019, 764–806

sensu lato plus S. caquetae. Roxo et al. (2019) proposed

Sturisomatichthys as sister of Farlowella, although only S.

tamanae was included in their molecular phylogeny; thus, no

further comments on the genus were offered by the authors.

The aim of the present study is to perform a taxonomic

revision of Sturisomatichthys. Since no taxonomic issues

regarding the genus have been addressed since its descrip-

tion, the analysis of the type series of all species was aimed to

be included, as well as an ample comparison to non-type

material of the species of the Loricariinae. Redescription of

species are presented, as well as updated diagnostic characters

for the genus and the species, and the descriptions of three

new species. The redescription of S. kneri is not included here

since the species was already described by Ghazzi (2005) and

no additional characters were found to rediagnose the

species. An identification key for the species and an update

on their distributions are also provided.

MATERIALS AND METHODS

Measurements and counts follow Ghazzi (2005), with the

addition of pre-pectoral length (from tip of snout to origin of

pectoral fin); distance between tip of snout and anterior

border of nostril; and internarial distance (taken at the

posteriormost margin of nares). Measurements were taken

point to point with digital calipers (0.1 mm precision).

Bilateral counts and measurements were taken from the left

side of the specimens except when the structure being

measured or counted was damaged, in which case data were

acquired from the right side.

Counts and nomenclature of lateral plate series follow

Ballen et al. (2016), nomenclature of abdominal plate series

follows Londo˜

no-Burbano and Reis (2016), and tooth cusp

names follow M¨

uller and Weber (1992). Number of speci-

mens in alcohol and cleared and stained are indicated by

‘‘alc’’ and ‘‘CS,’’ respectively. The abbreviation ‘‘St.’’ is used for

Sturisoma, in order to differentiate from Sturisomatichthys, for

which ‘‘S.’’ is used.

Institutional abbreviations follow Sabaj (2016). Osteolog-

ical nomenclature follows Paixa

˜o and Toledo-Piza (2009).

Specimens for osteological observations were cleared and

counterstained for bone and cartilage using the method of

Taylor and Van Dyke (1985). A dataset and phylogeny that is

in preparation for publication, including Sturisomatichthys

and allied genera, was used here for completeness of our

Discussion and support to our findings and hypotheses. The

construction of the distribution map for the species followed

the tutorial described by Calegari et al. (2016) using the

software and map packages in QGIS v. 2.18 and Google Earth.

Coordinates of localities that were not available in the

literature were calculated with Google Earth when possible.

RESULTS

Sturisomatichthys Isbr¨

ucker and Nijssen, 1979

Sturisomatichthys Isbr¨

ucker and Nijssen, in Isbr¨

ucker, 1979:91

(type species: Oxyloricaria leightoni Regan, 1912, by original

designation; gender: masculine).

Included taxa.—Sturisomatichthys aureus;S. caquetae;S. citur-

ensis;S. dariensis;S. festivus;S. frenatus;S. kneri;S. leightoni;S.

panamensis;S. tamanae;Sturisomatichthys guaitipan,new

species; Sturisomatichthys reinae, new species; Sturisomatich-

thys varii, new species.

Placement in Loricariinae.—Sturisomatichthys belongs to the

Farlowellina, along with Aposturisoma;Farlowella (type

genus); Lamontichthys;Pterosturisoma; and Sturisoma.

Diagnosis.—Sturisomatichthys can be distinguished from

members of the Loricariina by the possession of more than

30 teeth on both premaxilla and dentary (vs. less than 20);

absence of orbital notch (vs. notch present, except Fonchii-

loricaria); and 12 branched caudal-fin rays (vs. 10 branched

caudal-fin rays; except Fonchiiloricaria and Sturisomatichthys

guaitipan, new species). It is distinguished from genera

belonging to the Harttiini, encompassing Cteniloricaria,

Harttia, and Harttiella, by having abdominal plates always

present, developed, irregular, covering the entire abdomen

(vs. abdominal plates absent, or small and scarce, except

Cteniloricaria,Harttia absaberi,H. dissidens,H. duriventris,H.

fowleri,H. longipinna, and H. surinamensis); 15–18 plates on

the median plate series (vs. 19–21 plates on median plate

series); abdominal plates always reaching to the posterior

border of lower lip, covering cleithral region (vs. cleithral

region naked, except H. absaberi); and anterior abdominal

plates larger and more slender than central abdominal plates

(vs. plates absent, or the same size as central abdominal

plates). Sturisomatichthys is diagnosable from the remaining

Farlowellina (Aposturisoma,Farlowella,Lamontichthys,Ptero-

sturisoma, and Sturisoma) by having the nuchal plate at least

twice the size of predorsal plates (vs. nuchal plate the same

size or absent, except Sturisoma); from Lamontichthys it can be

distinguished by the possession of six branched pectoral-fin

rays (vs. seven branched pectoral-fin rays, which are also

present in Harttia leiopleura and Rineloricaria daraha); from

Farlowella it is distinguished by having irregular, numerous

central abdominal plates not arranged in series (vs. two or

three well-arranged series of large central abdominal plates)

and possession of a nuchal plate (vs. nuchal plate absent);

from Pterosturisoma it is distinguished by the possession of a

rostrum (vs. rostrum absent, except for S. caquetae,S.

citurensis,S. guaitipan, new species, and S. tamanae), S.

caquetae,S. citurensis, and S. tamanae are differentiated from

Pterosturisoma by having 19–21 coalescent plates in the

caudal peduncle (vs. 14–18 coalescent plates) and S.

guaitipan, new species, by having 12 branched rays on caudal

fin (vs. 10); from Sturisoma it is distinguished by the irregular,

numerous central abdominal plates not arranged in defined

series (vs. three clearly arranged series of developed central

abdominal plates); by the possession of dark spots on either,

or all, dorsal, pectoral, pelvic, or anal fins (vs. spots absent);

and by having 13–18 plates on the median plate series (vs.

20–21 plates on the median series).

Sexual dimorphism.—Adult males of Sturisomatichthys show

marked sexual dimorphism related to the development of

thin, short, and sometimes numerous odontodes on the sides

of the head. The development of odontodes can also be

perceptible on the first pectoral-fin ray and along the caudal

peduncle, although those odontodes are not as well devel-

oped as those on the head. No sexual dimorphism was

observed related to tooth cusps, urogenital pore, or thicken-

ing of the unbranched ray of the pectoral and/or pelvic fin.

Distribution.—Sturisomatichthys is distributed in the Tuyra,

Capeti, Aruza, and Bayano rivers in the Pacific slope of

Panama; Atrato, Sin´

u, Magdalena, Cauca, San Jorge, and

Catatumbo rivers on the Caribbean slope of Colombia;

Orinoco basin, Colombia; Maracaibo Lake basin in Venezu-

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 765

ela; Truando, San Juan, Baud´

o, and Pat´

ıa rivers on the Pacific

slope of Colombia; the Esmeraldas and Durango rivers on the

Pacific versant in northwestern Ecuador; and upper Amazon

River in Colombia (Figs. 1, 2, 3).

Remarks.—Isbr¨

ucker and Nijssen (in Isbr¨

ucker, 1979) includ-

ed as the only character to diagnose Sturisomatichthys the

absence of a rostrum, which differentiates it from Sturisoma.

That character was included by Covain et al. (2008), and it

was found to be uninformative and confirmed by Covain et

al. (2016).

Sturisomatichthys caquetae from the Caqueta

´River in

Colombia, Amazon basin, was included as part of the genus

representing the only cis-Andean species of the genus in the

original description. Covain et al. (2016) included most of

the species belonging to Sturisoma and some species of

Sturisomatichthys, including its type species, in their molec-

ular phylogeny of the Loricariinae. Those authors found

strong support for the inclusion of the trans-Andean species

traditionally assigned to Sturisoma in Sturisomatichthys.

Nevertheless, it was found here that S. caquetae does belong

to Sturisomatichthys, and S. leightoni, in addition to being

present in the Magdalena-Cauca basin, is also distributed in

the Orinoco basin (see below) which extends the distribution

of Sturisomatichthys out of the trans-Andean region.

As exposed by Covain and Fisch-Muller (2007) and

Londo˜

no-Burbano and Reis (2016), Dasyloricaria is the only

strictly trans-Andean genus in the Loricariinae. Based on the

results obtained in this study, this is not the case for

Sturisomatichthys as proposed by Covain et al. (2016).

Nevertheless, the distribution and occurrence of some of

the species of Sturisomatichthys (i.e., S. dariensis and S.

panamensis) in the Tuyra and Capeti basins of the Pacific

slope of Panama, and the Atrato River of the Caribbean slope

of Colombia, is the same as the distribution of Dasyloricaria

latiura (Londo˜

no-Burbano and Reis, 2016). This is also the

case for S. festivus, which is present in the Catatumbo River in

Colombia and the Maracaibo Lake basin in Venezuela, as is

D. filamentosa. A clear biogeographical component is ob-

served to explain the distribution of the species on both

genera (see Discussion here, and Londo˜

no-Burbano and Reis,

2016).

Sturisomatichthys aureus (Steindachner, 1900)

Figures 4, 5; Table 1

Loricaria aurea Steindachner, 1900: 206 (type locality:

Magdalena River at Bodega Central, Colombia; no type

assigned); 1902: 138, pl.5, figs. 1–1a (more detailed

description of the species; figures in dorsal and lateral

view of a specimen). Neumann, 2006: 276 (remarks of

holotype deposited at ZSM).

Sturisoma aureum (Steindachner, 1900).—Eigenmann, 1920a:

27 (present at upper Magdalena River, near Bogota

´); 1922:

96 (resurrection from synonymy with S. panamensis; short

description of the species). Fowler, 1942: 132 (distributed

at Magdalena basin; also at Meta River following Eigen-

mann [1922]). Miles, 1947: 104, fig. 68 (identification key

of the Loricariidae; drawings of dorsal view, and ventral

view of the head; distribution of the species; rejection of

Regan’s synonymy with S. panamensis). Dahl, 1971: 95

(present at Magdalena, San Jorge and Cesar rivers;

maximum size of 20 cm TL). Isbr¨

ucker, 1980: 93 (species

included in Sturisoma; no diagnosis); 1981a: 87 (listed as

valid species of Sturisoma). Roma

´n-Valencia, 1990: 205

(present at middle Atrato River). Isbr¨

ucker et al., 2001: 29,

32 (included in the catalog of the Loricariinae as valid in

Sturisoma, German version). Isbr¨

ucker, 2002: 29 (included

in the catalog of the Loricariinae as valid in Sturisoma,

English version). Ferraris, 2003: 346 (genus and species

distribution; synonymy; type material location; type as

missing; maximum body size; common name in Colom-

bia). Ghazzi, 2003: 223 (as incertae sedis within Loricar-

iinae). Lasso et al., 2004: 129 (not distributed in

Venezuela). Maldonado-Ocampo et al., 2006: 150 (distrib-

uted at Atrato River in Colombia; lot from ICNMHN).

Mojica et al., 2006a: 32 (distributed at the Magdalena-

Cauca Basin; catalog numbers at ICNMHN and IAvH);

2006b: 136 (distributed at Rancheria River, La Guajira,

Colombia; catalog numbers at ICNMHN); Villa-Navarro et

al., 2006: 13 (specimens in Colombian museums). Ferraris,

2007: 298–299 (type locality; distribution of the species).

Maldonado-Ocampo et al., 2008: 195 (distribution of the

species at Magdalena-Cauca basin, Pacific Slope rivers, and

Caribbean Slope rivers; type destroyed in World War II

[WWII], without citation of the source); 2013: 396 (present

at Atrato and Baud´

o rivers, Caribbean and Pacific versant of

Colombia, respectively; specimens at ICNMHN).

Sturisomatichthys aureus (Steindachner, 1900).—Covain et al.,

2016: 514 (valid as Sturisomatichthys aureum; phylogenetic

relationships of the Loricariinae subfamily using molecular

markers). DoNascimiento et al., 2018: 77 (as present and

endemic in Colombia, Magdalena-Cauca River basin).

Neotype.—ICNMHN 24054 (formerly ICNMHN 2414) 151.0

mm SL, Colombia, Cesar department, La Jagua de Ibiric ´

lower Magdalena River basin, Sororia Creek, tributary of

Tucuy River, 98410N, 73827 0W, by present designation.

Non-type material.—ICNMHN 2414, 25 alc, 110.2–141.7 mm

SL, same data as neotype; CIUA 365, 1 alc, 153.1 mm SL,

Colombia, Cesar department, lower Magdalena River basin,

Tucuy River, affluent of Calenturitas River, 98420N, 738160W;

CIUA 986, 1 alc, 118.1 mm SL, Colombia, Antioquia

department, Sons ´

on, middle Magdalena River basin, Samana

River, 58420300 N, 75818038 00 W; CIUA 1997, 1 alc, 140.8 mm SL,

Colombia, Santander department, Barrancabermeja, lower

Magdalena River basin, El Llanito swamp, San Silvestre

channel, 7840100 N, 738520100 W; ICNMHN 111, 2 alc, 162.3–

162.6 mm SL, Colombia, Choc ´

o department, Amparraid ´

district, Baud´

o River, 485701500N, 7782104900 W; ICNMHN 815,

1 alc, 156.9 mm SL, Colombia, La Guajira department,

Cerrejoncito Creek, 108570N, 728470W; ICNMHN 817, 1 alc,

Colombia, La Guajira department, Rancher´

ıa River; ICNMHN

7246, 2 alc, Colombia, Caldas department, Norcasia munic-

ipality, Middle Magdalena River, El Jagual Creek, tributary of

Manso River 98420N, 738160W; ICNMHN 8732, 5 alc,

Colombia, La Guajira department, Oreganal district, El

Cerrej´

on Mine, Rancher´

ıa River, downstream of Cerrej´

Creek, 11805022 00 N, 7284003100W; ICNMHN 8870, 4 alc,

Colombia, La Guajira department, Distracci ´

on municipality,

Chorreras district, Rancher´

ıa River at El Cercado, 10853048 00 N,

72853009 00 W; ICNMHN 8889, 13 alc, 115.8–165.2 mm SL,

Colombia, La Guajira department, Distracci ´

on municipality,

Chorreras district, Rancher´

ıa River at El Cercado sector,

108530N, 728530W; ICNMHN 9656, 4 alc, 143.9–152.9 mm

SL, Colombia, La Guajira department, Distracci ´

on munici-

pality, Chorreras district, El Cercado Bridge, Rancher´

ıa River,

108530N, 72853 0W; ICNMHN 9685, 2 alc, Colombia, La

Guajira department, Fonseca district, Rancheria River,

766 Copeia 107, No. 4, 2019

10853009 00 N, 7285005300W; ICNMHN 9697, 9 alc, 142.9–157.8

mm SL, Colombia, La Guajira department, Distracci ´

municipality, Chorreras district, Paso Ancho sector,

Rancher´

ıa River, 108530N, 728530W; ICNMHN 9842, 1 alc,

160.8 mm SL, Colombia, La Guajira department, Barrancas

district, La Quebrada Creek, tributary of Rancher´

ıa River,

10857021 00 N, 7284703100W; ICNMHN 10820, 4 alc, 127.0–

151.5 mm SL, Colombia, La Guajira department, Distracci ´

municipality, Chorreras district, El Cercado Bridge, Rancher´

ıa

River, 108530N, 728530W; ICNMHN 11652, 2 alc, Colombia,

La Guajira department, Riohacha City, Oreganal district,

Rancher´

ıa River, 1183105900N, 728540000W; ICNMHN 18166, 4

alc, Colombia, Cesar department, Pailitas municipality,

Lower Magdalena River, Hondo Creek tributary of Tunuma

channel, Bijao swamp, 88570N, 738370W; ICNMHN 18189, 8

alc, 118.9–151.2 mm SL, Colombia, Cesar department,

Pailitas district, Lower Magdalena River Raya Grande Creek,

tributary to Tunuma channel, Bijao swamp, 980007.7 00 N,

73839017.5 00 W, 62–93 m asl; ICNMHN 18408, 4 alc, 135.1–

146.0 mm SL, Colombia, Cesar department, Pelaya munic-

ipality, Lower Magdalena River, Sucio channel, Las Damas

and Orisn ´

o Creeks, tributaries of San Bernardo Swamp,

8843054.2 00 N, 73840050.500W, 42–86 m asl; ICNMHN 18504,

14 alc, 109.3–133.3 mm SL, Colombia, Cesar department,

lower Magdalena River basin, La Gloria municipality, El

Carmen district, Sima˜

na River, 8840026,700N, 7383802.400 W, 7 0

m asl; MPUJ 5796, 1 alc, 117.6 mm SL, Colombia, Santander

department, San Vicente de Chucur´

ı municipality, Lower

Magdalena River, 6848026.9400N, 73843024.0400 W, 111 m asl;

NRM 15150, 4 alc, 1 CS, 129.5–202.4 mm SL, Colombia,

Choc´

o department, Rio Baud ´

o basin, Boca de Pep´

e, various

tributaries and river close to village, 88590400N, 77830W.

Diagnosis.—Sturisomatichthys aureus is distinguished from its

congeners by the possession of a transverse bar at the dorsal-

fin origin, which is united to the dorsolateral stripe (vs.

transverse bar absent or, when present, not united to

dorsolateral stripe). From S. caquetae,S. citurensis,S. guaitipan,

new species, and S. tamanae, it is further distinguished by the

possession of a rostrum (vs. rostrum absent) and possession

of a dorsolateral stripe on the body, interrupted at caudal

peduncle (vs. stripe absent). Furthermore, it is distinguished

by the cleithral width 10.5–12.8% SL (vs. 9.9–10.5% SL in S.

dariensis; 8.5–10.5% SL in S. frenatus; 8.4–10.5% SL in S.

panamensis; 8.3–9.6% SL in S. reinae, new species; and 7.7–

9.2% SL in S. varii, new species). It is also distinguished from

S. dariensis and S. panamensis by the possession of a large,

dark blotch on the pelvic and anal fins occupying almost the

entire fin (vs. large, dark blotch absent or inconspicuous),

from S. leightoni by the possession of large, dark blotches on

the dorsal, pectoral, pelvic, and anal fins occupying almost

the entire fins (vs. dark blotches absent), and by the absence

of reticulations on predorsal, supraoccipital, and interorbital

regions (vs. reticulations present), and from S. festivus by the

absence of filaments on dorsal, pectoral, pelvic, and anal fins

(vs. filaments present). It is also distinguished by having

Fig. 1. Map of northwestern South America, including Colombia, Ecuador, Panama, and Venezuela, showing the distribution of Sturisomatichthys

aureus (circles); S. dariensis (white squares); S. festivus (diamonds); S. kneri (half white-half black square); S. panamensis (triangles);

Sturisomatichthys reinae, new species (pentagon); Sturisomatichthys guaitipan, new species (hexagon); and Sturisomatichthys varii, new species

(star). White symbols show type locality of each species. Each symbol can represent more than one locality.

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 767

more numerous premaxillary teeth, 47–69 (vs. 38–45 in S.

reinae, new species) and more numerous dentary teeth, 40–56

(vs. 29–36 in S. reinae, new species, and 26–39 in S. varii, new

species).

Description.—Morphometric data in Table 1. Body short and

slender completely covered by dermal plates with small

hypertrophied odontodes, poorly developed on sides of head,

and paired fins. Dorsal profile of head straight from tip of

snout to dorsal-fin base. Dorsal body profile slightly concave

posterior to dorsal fin, nearly straight on last plates of caudal

peduncle. Ventral body profile flat. Head and predorsal

region broader than remainder of body; postdorsal region

narrower towards caudal fin.

Head triangular. Rostrum short and broad along its entire

length. Ventral border of rostrum flat. Orbital rim circular,

without notch, dorsolaterally placed on head. Sphenotic

with short lateral expansion. Canal-bearing plate long, well

developed, not entirely in contact with anterior abdominal

plates, anterior to branchiostegal membrane, reaching tip of

rictal barbel. Upper lip covered with small, almost impercep-

tible plates; lower lip without such plates. Lower lip covered

with small papillae rounded or fringe-like. Rictal barbel short

and free from lip rim, not reaching to anterior border of

abdominal plates. Teeth slender, bifid, 47–69 on premaxilla,

40–56 on dentary; cusps strongly curved. Teeth on lower jaw

less than twice size of those on upper jaw.

First two predorsal plates with two low, inconspicuous

ridges. Nuchal plate twice size of predorsal plates. Median

series of lateral plates carinate, forming two longitudinal ill-

developed keels; coalescent plates 14–18, beginning at 13–16

median plates. Six to nine straight lateral abdominal plates,

between pectoral- and pelvic-fin origins. Preanal plate

bordered anteriorly by three well-developed plates of similar

size. Central abdominal plates unorganized; anterior abdom-

inal plates smaller and more numerous than posterior ones.

Posterior margin of dorsal fin straight to slightly concave,

unbranched and first branched rays longest; dorsal-fin rays

i,7. Pectoral-fin rays i,6, unbranched and first branched rays

longest, their tips reaching anterior third of pelvic fin. Pelvic-

fin rays i,5, unbranched and first branched rays longest, their

tips slightly passing base of anal fin. Anal-fin rays i,5, first

branched ray longer. Posterior caudal-fin margin deeply

concave, upper and lower ray bearing filamentous exten-

sions.

Coloration in alcohol.—Ground color of dorsum and head

light or dark brown to pale yellow; ventrally yellow.

Dorsolateral dark stripe starting on tip of snout. Stripe not

continuous, interrupted at predorsal plates, and present

again on caudal peduncle as discontinuous small stripe (Fig.

4). Broad transverse bar at origin of dorsal fin (Fig. 4). Several

thin, brown, sinuous lines between contralateral dorsolateral

stripes on snout, not continuing posterior to supraoccipital

(Fig. 4). Dark band along first two branched rays of dorsal fin;

other rays and membranes pale yellow. Dorsal-fin band with

one to several white ocelli on distal tip, and just anterior to

middle of fin (Fig. 5). Pectoral and pelvic fins with dark spot,

Fig. 2. Map of northwestern South America, including Colombia, Ecuador, and part of northern Brazil on its political border with Colombia, showing

the distribution of Sturisomatichthys caquetae (triangle; type locality) and S. frenatus (pentagon; white polygon is type locality). Each symbol can

represent more than one locality.

768 Copeia 107, No. 4, 2019

larger than that on dorsal fin (Fig. 5), on first two to three

rays on both fins, or spot occupying entire fin; several dark

spots easily visible mainly on unbranched ray. Edges of

caudal lobes with dark band with white ocellus on both

upper and lower rays, with same distribution as those on

dorsal fin (Figs. 4, 5).

Sexual dimorphism.—Males with numerous, thin, and short

hypertrophied odontodes on the sides of the head (Fig. 4).

Less conspicuous hypertrophied odontodes can also be

present on first ray of pectoral fin.

Distribution.—Sturisomatichthys aureus occurs in the lower

and middle Magdalena, Baud ´

o, Cesar, Sin ´

u, San Jorge, and

Rancher´

ıa rivers in Colombia (Fig. 1).

Remarks.—Steindachner (1900) described Sturisomatichthys

aureus originally as Loricaria aurea, from Bodega Central,

Magdalena River, Colombia. The author presented a short

description of the species focused on the general morphology

and counts of lateral plates. Subsequently, Steindachner

(1902) offered a second, more comprehensive description,

including measurements, general description of body shape,

and some meristic data. The description highlighted that S.

aureus had no sexual dimorphism of odontodes along the

sides of the head, which, contrarily, were observed in the

present study. Further on, the hypertrophied odontodes,

filaments on both upper and lower caudal-fin rays, and

curved teeth were also confirmed (see Description). It is

worth noting that Steindachner also reported, ‘‘the lateral

keels on the anterior part of the trunk are weakly developed

and unite on the left side of the body on the 13

plate, and

on the 15

plate on the right . . .’’ (Steindachner, 1902: 139;

our translation); Steindachner (1900: 207) also reported,

‘‘Both sides of the trunk [with plates of the median series

until] the 14

or 15

plate. [When] plates already together,

the simple side edge extends 17 plates’’ (our translation).

Those characteristics were verified on non-type material,

confirming that lower counts on median plates are diagnos-

tic for Sturisomatichthys (see genus Diagnosis). In addition,

Steindachner failed to mention any kind of coloration

pattern of the species, the only color remark being in the

etymology of the species, in that: ‘‘The name of the species is

chosen according to the golden glow spread over the whole

body’’ (Steindachner, 1902: 139; our translation). Addition-

ally, the author presented a drawing of a specimen in lateral

and dorsal view, and again, not showing any kind of

particular coloration (Steindachner, 1902: plate V., figs. 1–

1a). As described above, S. aureus does have dorsolateral

Fig. 3. Map of northwestern South America, including Colombia, Ecuador, Panama, and Venezuela, showing the distribution of Sturisomatichthys

citurensis (stars), S. leightoni (circles), and S. tamanae (squares). White symbols show type locality of each species. Each symbol can represent more

than one locality.

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 769

stripe, although discontinuous on the caudal peduncle, as a

diagnostic character. Finally, the poorly developed rostrum

presented in the drawings was confirmed on non-type

material, and furthermore, it was observed that there is

variation within the species regarding rostrum length.

Sturisomatichthys aureus has not been extensively studied

despite being described at the beginning of the 20

century.

The studies including the species deal with short descrip-

tions, few diagnostic characters to identify the species, and

distribution (e.g., Regan, 1904; Eigenmann, 1920a, 1922;

Miles, 1947; Dahl, 1971).

Fig. 4. Sturisomatichthys aureus, neotype, ICNMHN 24054, 151.0 mm SL, Colombia, Cesar department, La Jagua de Ibiric ´

o municipality, Lower

Magdalena basin, Sororia Creek tributary of Tucuy River. Photo by Henry Agudelo (ICNMHN).

Fig. 5. Sturisomatichthys aureus, ICNMHN 9842, 160.8 mm SL, Colombia, La Guajira department, Barrancas district, La Quebrada Creek, tributary of

Rancher´

ıa River. Note white ocelli on dorsal fin. Photo by Henry Agudelo (ICNMHN).

770 Copeia 107, No. 4, 2019

Regan (1904) was the first author to treat this species as

Oxyloricaria (¼Sturisoma), including it as a synonym of O.

panamensis. The author stated that after examining the type

of Loricaria panamensis (98 mm TL) and comparing it to the

type of L. aurea (169 mm SL) collected at the Magdalena River

basin, the latter was an adult of L. panamensis, described by

Eigenmann and Eigenmann (1889; see Remarks on S.

panamensis). After that, Eigenmann (1922) included the

species as Sturisoma aureum, following the synonymy of

Oxyloricaria and Sturisoma proposed earlier (Eigenmann,

1910). Eigenmann (1922) cited the synonymy between S.

aureus and S. panamensis proposed by Regan (1904), but

resurrected S. aureus discussing that the species could be an

intermediate between S. panamensis and S. leightoni.We

follow Eigenmann (1922) in assuming S. aureus as a distinct

species from S. panamensis.

In two studies of the ichthyofauna of the Choc ´

o region in

Colombia, Maldonado-Ocampo et al. (2006, 2013) reported

that S. aureus is present in the Atrato River basin, a river

emptying in the Caribbean versant of Colombia. It was

found here that the species is not present in that locality, but

instead, the records of those authors belong to other species

(see Remarks on S. panamensis). Nevertheless, Maldonado-

Ocampo et al. (2013) reported the species for the Baud´

River, which could be either S. aureus or S. reinae, new species.

On the other hand, Maldonado-Ocampo et al. (2008), in a

species checklist of the ichthyofauna of Colombia, reported

the species to be present in the Magdalena-Cauca basin, and

in the Caribbean versant of Colombia. This study agrees with

the report of the species in the Magdalena-Cauca basin, but

disagrees with the report that the species is present in the

Caribbean versant, except for the report of the Magdalena-

Cauca basin that belongs to that versant in Colombia.

Sturisomatichthys aureus is distributed in the Magdalena-

Cauca, Baud´

o, Sin ´

u, San Jorge, and Rancher´

ıa rivers of

Colombia (Fig. 1).

Sturisomatichthys aureus is sympatric with S. reinae, new

species, in the Baud ´

o River, but several characters (see

Diagnosis) can differentiate the species. Sturisomatichthys

aureus is also sympatric with S. leightoni in the middle portion

of the Magdalena River. Further on the distinguishing

features of the Diagnosis, the species have different body

size; while S. aureus can reach 200 mm SL, S. leightoni do not

surpass 120 mm SL. In general, both species possess different

characteristics which, despite being sympatric, may allow

them to occupy different niches and explore different types

of resources. In any case, ecological studies of these species

and of Sturisomatichthys in general are lacking.

Steindachner (1900, 1902) did not offer a catalog number

for the type specimen of Loricaria aurea, nor acronym of any

museum. Isbr¨

ucker (1980), in his catalog of the Loricariidae,

reported the holotype of Loricaria aurea as being the

Table 1. Morphometrics and meristics of Sturisomatichthys aureus and S. caquetae. Morphometric values as percents of standard length (SL) and

head length (HL). SD ¼standard deviation.

S.aureus (n¼51) S.caquetae

Min Max Mean SD Holotype

Standard length 109.3 202.4 138 90

Head length 16.7 22.5 20.9 1.3 22.2

Predorsal length 26.2 33.2 30.9 1.6 32.7

Abdominal length 11.0 13.9 12.2 0.7 14.4

Trunk length 11.2 17.4 14.5 1.3 14.4

Dorsal fin length 22.1 31.0 27.7 2.1 —

Pectoral fin length 17.6 23.5 20.8 1.6 —

Pelvic fin length 13.6 22.3 16.7 1.3 15.9

Anal fin length 16.7 22.8 20.1 1.4 18.8

Head depth at supraoccipital tip 8.0 10.5 9.4 0.7 8.4

Body depth at dorsal-fin origin 9.0 13.1 11.6 1.0 9.8

Caudal peduncle depth 1.1 2.0 1.5 0.2 1.3

Cleithral width 10.5 12.8 11.8 0.6 11.2

Body width at dorsal-fin origin 10.9 13.9 12.5 0.7 13.6

Body width at anal-fin origin 9.6 12.3 10.7 0.7 10.9

Body width at caudal-fin origin 1.5 2.8 2.2 0.4 2.4

Head width 60.4 79.6 64.9 4.5 60.0

interorbital distance 23.9 33.2 30.9 1.8 28.0

Orbital diameter 11.5 17.9 15.0 1.3 13.5

Snout length 53.4 64.8 61.4 2.0 56.0

Dentary length 8.7 15.0 11.5 1.3 12.5

Lower lip width 27.9 41.9 34.4 2.9 29.5

Plates on median series 13 16 14 14

Coalesced plates 14 18 16 14

Ventrolateral thoracic plates 6 9 7 7

Premaxillary teeth 47 69 57 33

Dentary teeth 40 56 42 30

Dorsal-fin rays i,7 i,7 i,7 i,7

Pectoral-fin rays i,6 i,6 i,6 i,6

Pelvic-fin rays i,5 i,5 i,5 i,5

Anal-fin rays i,5 i,5 i,5 i,5

Caudal-fin rays i,12,i i,12,i i,12,i i,12,i

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 771

specimen illustrated by Steindachner (1902: pl. V., figs. 1–1a).

That specimen is the only one included in Steindachner’s

description, and it is assumed to be the holotype, although

the author made no specific claim regarding the specimen.

Nevertheless, Neumann (2006) confirmed that the specimen

used by Steindachner in the original description was

deposited at ZMS, with the following information on the

label: ‘‘Holotype (unique): ZSM [Old Collection; without

registry of catalogue number], 16.9 cm SL, Rio Magdalena at

Bodega Central (Colombia); leg: local fishermen; coll. Th. v.

Bayern, 18.VI.1898.’’ and added that: ‘‘Holotype fixed by

monotypy (ICZN Art. 73.1.2)’’, confirming the type status of

the specimen. According to Dirk Neumann (pers. comm.),

there is no record of the lot number for the lost holotype of

the species.

According to Ferraris (2003, 2007), the holotype of

Loricaria aurea is housed at the NMW, but no catalog number

is included, and no explanation is given to demonstrate that

the specimen is housed in Vienna and not in Munich. Later,

Maldonado-Ocampo et al. (2008) reported the holotype as

destroyed during WWII, but again no reference is included to

back such a statement. Nevertheless, Fricke et al. (2019b) also

state that the type was destroyed during WWII: ‘‘Holotype

(unique): ZSM [old collection] destroyed in WWII’’. To clarify

the situation of the possible non-existence of the holotype, a

communication with Anja Palandacic (NMW) confirmed

that the specimen was not housed at NMW, but in ZSM.

Thus, and according to Dirk Neumann (ZSM, pers. comm.):

‘‘The type of Loricaria aurea was collected by Princess Therese

von Bayern and described by Steindachner. At the time of

description the specimen was still in the private collection of

Princess Therese, which ZSM received after her death in

1925. Only one specimen from her fish collection apparently

survived the war in ZSM, which is one of the two syntypes of

Pimelodella yuncensis.’’ Thus, the holotype of Loricaria aurea

was in fact destroyed during WWII, and, according to records

at both NMW and ZSM, there are no additional specimens

that Steindachner could have used to describe the species. In

this sense, a neotype is assigned to the species (ICZN Art.

75.3). The specimen catalogued under ICNMHN 24054 (ex

ICNMHN 2414), 151.0 mm SL (Fig. 4), from Colombia, Cesar

department, lower Magdalena basin, Sororia creek at Jagua de

Ibiric´

o, which is near the known type locality of the species

(Bodega Central, a small town in the lower portion of the

Magdalena River), is here designed as the neotype for S.

aureus.

Sturisomatichthys caquetae (Fowler, 1945)

Figure 6, Table 1

Harttia caquetae Fowler, 1945: 126, fig. 41–43 (type locality:

Morelia, Rio Caqueta

´, Colombia; holotype ANSP 71719,

length 90.0 mm SL).

Sturisomatichthys caquetae (Fowler, 1945).—Isbr ¨

ucker and

Nijssen, in Isbr¨

ucker, 1979: 88 (original description of

Sturisomatichthys). Isbr¨

ucker et al., 2001: 32 (included in

the catalog of the Loricariinae as valid in Sturisomatichthys,

German version). Isbr¨

ucker, 2002: 29 (included in the

catalog of the Loricariinae as valid in Sturisomatichthys,

English version). Ferraris, 2003: 347 (genus distribution;

synonymy; type material location at ANSP 71719, maxi-

mum length 19.3 cm); 2007: 300 (type locality; distribu-

tion of the species).

Sturisoma caquetae (Fowler, 1945).—Covain et al., 2016: 514

(phylogenetic relationships of the Loricariinae subfamily

using two mitochondrial and one nuclear markers; not

included in phylogenetic analysis). DoNascimiento et al.,

2018: 77 (as present and endemic in Colombia, Amazonas

River basin).

Diagnosis.—Sturisomatichthys caquetae is diagnosed from its

congeners, except S. citurensis,S. guaitipan, new species, and

S. tamanae, by the absence of a rostrum (vs. rostrum present)

and absence of dorsolateral stripe (vs. dorsolateral stripe

present). It is also distinguished from other congeners by

having 33 premaxillary teeth (vs. 47–69 in S. aureus; 61–124

in S. citurensis; 37–55 in S. festivus; 36–50 in S. frenatus; 37–53

in S. panamensis; 38–45 in S. reinae, new species; and 63–98 in

S. tamanae), and 30 dentary teeth (vs. 40–56 in S. aureus; 63–

100 in S. citurensis; 34–47 in S. dariensis; 33–42 in S. festivus;

31–46 in S. frenatus; 32–44 in S. panamensis; and 60–93 in S.

tamanae). From S. leightoni it is distinguished by having more

numerous and smaller central abdominal plates (vs. less

numerous and larger central abdominal plates), more than

six transversal bars not restricted to caudal peduncle (vs.

three to four transversal narrow bars restricted to caudal

peduncle), and lacking a dorsolateral discontinuous stripe on

both sides of the snout (vs. stripe present). Additionally, it is

distinguished by its head width (60.0% HL vs. 70.1–80.4%

HL in S. citurensis and 74.9–83.5% HL in S. tamanae) and

lower lip width (29.5% HL vs. 33.0–43.9% HL in S. citurensis

and 34.3–53.9% HL in S. tamanae). From S. festivus,itis

distinguished by the absence of filaments on dorsal, pectoral,

pelvic, and anal fins (vs. filaments present), and from S.

guaitipan, new species, by the trunk length (14.4% SL vs.

14.9–16.7% SL), body depth at dorsal-fin origin (9.8% SL vs.

10.3–12.1% SL), 12 branched rays on caudal fin (vs. 10), and

snout and head dorsally without spots (vs. spots present).

Description.—Morphometric data in Table 1. Both measure-

ments and description taken from holotype. Body short and

low, completely covered by dermal plates with few, short,

hypertrophied odontodes, ill developed, but perceptible on

sides of head. Dorsal profile of head straight from tip of snout

to predorsal region; slightly concave from that point to end

of dorsal-fin base, and straight from there until caudal-fin

origin. Ventral body profile flat.

Head triangular. Snout without rostrum. Orbital rim oval

without notch, dorsolaterally placed on head. Sphenotic

without lateral expansion. Canal-bearing plate broad and

well developed in contact with anterior abdominal plates,

anterior to branchiostegal membrane not reaching rictal

barbel. Upper lip covered with small, almost imperceptible

plates; lower lip without such plates. Lower lip covered with

round, small, almost imperceptible papillae, increasing in

size towards dentaries. Rictal barbel extremely short, free

from lip rim, never reaching anterior border of abdominal

plates, nor surpassing posterior border of lower lip. Teeth

slender, bifid, 33 on premaxilla and 30 on dentary; cusps

slightly curved. Teeth in lower jaw somewhat larger than

those in upper jaw.

Predorsal plates without ridges; nuchal plate twice size of

predorsal plates. Lateral plates carinate, forming two longi-

tudinal low subtle keels; 14 coalescent plates, beginning at

median plate. Eight straight lateral abdominal plates,

between pectoral- and pelvic-fin origins, longer than central

abdominal plates. Preanal plate bordered anteriorly by five

well-developed plates, central one larger than others, all in

contact with preanal plate. Central abdominal plates,

between lateral abdominal plates, without any kind of

772 Copeia 107, No. 4, 2019

arrangement. Anterior abdominal plates with almost same

size as central abdominal plates, most anterior ones enlarged,

not square.

Posterior margin of dorsal fin straight, unbranched ray

longest; dorsal-fin rays i,7. Pectoral-fin rays i,6, unbranched

ray longest (right fin, left broken), surpassing anterior third

of pelvic fin, reaching almost half of pelvic fin. Pelvic-fin rays

i,5, unbranched and first branched ray longest, their tips

reaching little beyond origin of anal fin. Anal-fin rays i,5,

unbranched ray longer. Posterior caudal-fin margin deeply

concave, filaments not observed in upper and lower caudal

rays due to damage. Dorsal, pectoral, pelvic, and anal fins

without filaments.

Coloration in alcohol.—Based on both holotype and holotype

drawing by Fowler (1945: 127, figs. 41–43). Ground color of

dorsum, head, and ventral side pale brown. Stripe on dorsal

portion of body not visible, nor specified by Fowler (1945).

Three to four visible narrow transverse stripes along caudal

peduncle, first at base of dorsal fin, last about three plates

anterior to origin of caudal fin. Transverse stripe somewhat

visible on holotype and included on drawings by Fowler

(1945), who reported eight transverse bars, that at dorsal-fin

origin being most distinct. Large, dark spots on membranes

and rays of pectoral, pelvic, and anal fins, scarce, not

occupying entire fins. Dorsal fin with dark brown spot at

distal tip of first two branched rays, and with conspicuous

dark spots along unbranched ray. Latter marks not men-

tioned on original description, nor drawing of type. Pectoral,

pelvic, and anal fin with spots along rays, without large spots

or stripe occupying entire fin. Caudal fin with same pattern

as pectoral, pelvic, and anal fin.

Sexual dimorphism.—The holotype, which is a male, possesses

few, short, hypertrophied odontodes on the sides of the head

(Fig. 6), which typically represents a sexually dimorphic

feature in species of Sturisomatichthys.

Distribution.—Sturisomatichthys caquetae occurs in the Caque-

´River, upper Amazon, Caqueta

´department, at Morelia

municipality, Colombia (Fig. 2).

Remarks.—Fowler (1945) described Sturisomatichthys caquetae

as Harttia caquetae, from Morelia, Caqueta

´River in the upper

Amazon, in Colombia. The author focused his description on

body measurements, counts of plates and teeth, and some

remarks on coloration. The species lacks the dorsolateral

stripe present in most species of Sturisomatichthys, but shows

some characteristics similar to those observed on S. leightoni,

consisting of transverse bars, beginning at the base of the

Fig. 6. Sturisomatichthys caquetae, holotype of Harttia caquetae, ANSP 71719, 90.0 mm SL, Colombia, Caqueta

´department, Morelia municipality,

Upper Amazon basin, Caqueta

´River. Photo by Kyle Luckenbill (ANSP).

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 773

dorsal fin (Fig. 6). On the other hand, the central abdominal

plates occur as small, irregularly arranged plates such as those

on S. citurensis,S. guaitipan, new species, and S. tamanae.In

addition, low counts on both plates on the median series and

coalescent plates agree with those found throughout the

genus (see below). Thus, the species shows characters that

diagnose it as a member of Sturisomatichthys. According to a

drawing of the type in its original description (Fowler, 1945:

127, figs. 41–43), both upper and lower caudal rays have

short filaments that do not reach half SL.

Sturisomatichthys caquetae is a poorly known species. After

its description, it was not included in any revision of the

Loricariidae, until Isbr¨

ucker and Nijssen (in Isbr¨

ucker, 1979)

included it as part of Sturisomatichthys due to the absence of a

produced rostrum. As shown above, Sturisomatichthys re-

mained as an obscure group, restricted to four species, until

the study of Covain et al. (2016). In that study, the number of

species was elevated from four to nine and restricted to trans-

Andean species. Nevertheless, the authors did not include S.

caquetae in their analysis or examine its holotype, and, based

on its cis-Andean distribution, they transferred it to Stur-

isoma.

Nevertheless, after examining the holotype and comparing

its characteristics to the remaining Sturisomatichthys (includ-

ing three new species described in this study), it was

determined that S. caquetae belongs to the latter genus. Even

though non-type material was not available for examination,

the species is transferred back from Sturisoma to Sturisoma-

tichthys based on the holotype. The most important

similarity, and the one that allows transferring the species,

is the low plate count of the median series (14), which is

diagnostic of Sturisomatichthys when compared to Sturisoma

(more than 18 plates in the median series; see Diagnosis of

the genus). Thus, Sturisomatichthys is not a fully trans-

Andean genus, as it will be discussed below (see Remarks

on S. leightoni), and S. caquetae remains the only fully cis-

Andean species of the genus.

It is worth noting that S. caquetae could be easily

considered as a junior synonym of Sturisoma brevirostre from

the I¸ca

´River, Brazil, near Leticia, Colombia, type locality of S.

caquetae. The I¸ca

´River is the same river as the Putumayo

River in Colombia, and receives the name I¸ca

´when crossing

the Brazilian political border; the Caqueta

´River is a tributary

to the Putumayo River. The main character Eigenmann and

Eigenmann (1889) relied upon for the description of St.

brevirostre is the absence of a rostrum, which is also

characteristic of S. caquetae. Non-type specimens of St.

brevirostre were not available for the present study, but,

through the analysis of the holotypes of both species (S.

caquetae, ANSP 71719 and St. brevirostre, MCZ 8095), it was

concluded that they belong to two different species. This

differentiation is proposed mainly based on the plate count

of the median series and the central abdominal plate array,

which distinguish Sturisomatichthys from Sturisoma.

Throughout the present study, plate counts (both median

and coalescent) were found not to vary along the ontogeny

of the specimens, and it is a constant character when it

comes to identifing species. The counts observed for juvenile

specimens are the same reported for adults. Both species are

scarce to non-existent in visited museums, and collection in

the type localities of both species and surrounding areas are

needed. Thus, the inclusion of non-type specimens are

extremely important to corroborate this hypothesis.

Material examined.—ANSP 71719, 90.0 mm SL, holotype,

Colombia, Caqueta

´department, Morelia municipality, upper

Amazon basin, Caqueta

´River, 182900900 N, 75843 02800 W, Mr.

Kjell von Sneidern.

Sturisomatichthys citurensis (Meek and Hildebrand, 1913)

Figure 7, Table 2

Oxyloricaria citurensis Meek and Hildebrand, 1913: 82 (type

locality: Cupe River, Cituro, Tuyra River basin, Panama;

holotype FMNH 7585, length 194 mm [TL]; original

description without specific remarks, focusing on measure-

ments and coloration of the specimens); 1916: 219

(identification key for Oxyloricaria, including O. panamen-

sis; ventral figure of one specimen; description of the

species; comparison with O. tamanae). Ibarra and Stewart,

1987: 63 (type series at FMNH; holotype missing or mixed

with paratypes; 150 paratypes reported from five different

localities).

Sturisoma citurense (Meek and Hildebrand, 1913).—Eigen-

mann, 1922: 96, plate XVI, fig. 3 (citation of the original

description; species distributed also at the Bayano River in

Panama; type specimens examined by the author).

Sturisomatichthys citurensis (Meek and Hildebrand, 1913).—

Isbr¨

ucker, 1979: 88 (original description of Sturisomatich-

thys; included in the genus as Sturisomatichthys citurensis);

1980: 92 (included in Sturisomatichthys as a member of the

Harttiina; no diagnosis). Isbr¨

ucker et al., 2001: 32 (included

in the catalog of the Loricariinae as valid in Sturisomatich-

thys, German version). Isbr¨

ucker, 2002: 29 (included in the

catalog of the Loricariinae as valid in Sturisomatichthys,

English version). Ferraris, 2003: 347 (genus distribution;

synonymy; type material location at FMNH 7585, 25 cm

SL; maximum body size). Covain and Fisch-Muller, 2007: 4

(specimen examined to describe Sturisomatichthys (MHNG

2676.04, 152.3 mm of SL, Panama, Darien, Rio Chucuna-

que near the village La Alba, de Rham); observations

regarding abdominal plates organization and size). Ferraris,

2007: 300 (type locality; distribution of the species).

Covain et al., 2008: 988 (belonging to Sturisomina as

sister group of Farlowella platoryncha; phylogenetic analysis

using two mitochondrial markers). Rodriguez et al., 2011: 9

(belonging to Farlowellina as sister group of Farlowella;

phylogenetic analysis using two mitochondrial and one

nuclear markers). Covain et al., 2016: 514 (valid as

Sturisomatichthys citurensis; phylogenetic relationships of

the Loricariinae subfamily using two mitochondrial and

one nuclear markers; appears as sister group to all

Sturisomatichthys).

Diagnosis.—Sturisomatichthys citurensis is diagnosed from its

congenersexcept S. caquetae,S. guaitipan, new species, and S.

tamanae, by lacking a rostrum (vs. rostrum present), lacking a

laterodorsal stripe on both sides of flanks (vs. stripe present),

absence of dark spots occupying the entire or almost entire

dorsal and pectoral fin (vs. dark spots present on either of

such fins), and snout circular in dorsal and ventral view (vs.

snout triangular). From S. caquetae and S. guaitipan, new

species, it is distinguished by the number of premaxillary

teeth (61–124 vs. 33 and 32–43, respectively) and dentary

teeth (63–100 vs. 30 and 30–44, respectively). It is distin-

guished from S. tamanae by having small spots on rays, never

occupying the entire fin, nor entering the membranes (vs.

large, dark spot occupying almost the entire dorsal, pectoral,

774 Copeia 107, No. 4, 2019

and pelvic fin), and less numerous central abdominal plates

(vs. more numerous central abdominal plates; Fig. 7).

Description.—Morphometric data in Table 2. Body slender,

completely covered by dermal plates with small hypertro-

phied odontodes, poorly or well developed on sides of head.

Dorsal profile of head straight from tip of snout to

supraoccipital plates; concave from that point to origin of

dorsal fin. Dorsal body straight posterior to dorsal fin until

origin of caudal fin. Ventral body profile flat. Head and

predorsal region broader than remainder of body.

Head rounded. Rostrum absent; head short and broad its

entire length. Orbital rim circular without notch, dorsolater-

ally placed on head. Sphenotic with thin, long lateral

expansion. Canal-bearing plate long, well developed, in

contact with anterior abdominal plates, anterior to bran-

chiostegal membrane, not reaching tip of rictal barbel. Upper

lip covered with small, almost imperceptible plates; lower lip

without such plates. Lower lip covered with small papillae,

which could be rounded or fringe-like extensions. Upper lip

well developed, almost twice as broad as lower lip, occupying

almost entire anterior ventral portion of snout, with straight

posterior border without covering premaxillary teeth (Fig. 7).

Rictal barbel short and free from lip rim, not reaching

anterior border of abdominal plates. Teeth slender, bifid, 61–

124 on premaxilla, 63–100 on dentary; cusps slightly curved.

Teeth in lower jaw somewhat larger than those in upper jaw.

Predorsal plates with two low, inconspicuous ridges.

Nuchal plate twice size of predorsal plates, also with

inconspicuous ridge. Lateral plates carinate, forming two

longitudinal sharp keels; coalescent plates 13–17, beginning

at 13–17 median plates. Six to 11 straight lateral abdominal

plates, between pectoral- and pelvic-fin origins. Preanal plate

bordered anteriorly by four well-developed and similarly

sized plates. Abdominal plates small and irregular, between

lateral abdominal plates, unorganized; anterior abdominal

plates smaller and more numerous than posterior ones.

Posterior margin of dorsal fin straight to slightly concave,

unbranched ray longest; dorsal-fin rays i,7. Pectoral-fin rays

i,6, unbranched ray longest, its tip reaching anterior third, or

just origin of pelvic fin. Pelvic-fin rays i,5, unbranched ray

longest, its tip surpassing anal-fin base. Anal-fin rays i,5, first

branched ray longer. Posterior caudal-fin margin deeply

concave, absence of filaments on both upper and lower rays,

maybe due to damage.

Coloration in alcohol.—Ground color of dorsum and head

dark brown, both dorsally and ventrally, becoming darker

ventrally from middle of abdomen posteriorly. No dorsolat-

eral stripe. Dark stripe or spots lacking on membranes of

pectoral and pelvic fins; membranes pale yellow. Dorsal,

pectoral, pelvic, and anal fins with conspicuous and

somewhat large brown spots along rays, without entering

membranes. Pigmentation on edges of caudal lobes poorly

visible, although darker pigmentation, not well defined as

spots or stripe, occurs along all of its rays and membrane.

Sexual dimorphism.—Males show developed but short hyper-

trophied odontodes along sides of head (Fig. 7). Odontodes

absent from fin rays.

Fig. 7. Sturisomatichthys citurensis, USNM 78361, 196.3 mm SL, Panama, Canal Zone, Darien Province at Aruza, Aruza River.

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 775

Distribution.—Sturisomatichthys citurensis occurs in the Baya-

no, Capeti, Chucunaque, Cituro, and Tuyra rivers, Panama,

and Atrato River basin in Colombia (Fig. 3).

Remarks.—As mentioned above, S. tamanae is morphologi-

cally more closely related to S. citurensis. Both species lack a

rostrum, although as it was observed by Meek and Hilde-

brand (1916): ‘‘[S. citurensis presents] head without ridges;

snout not produced, broadly obtuse’’, which is also different

from what was observed here on S. tamanae.InS. citurensis,

the anterior border of the head is rounded and short,

contrary to the triangular format in S. tamanae.Head

morphology in this species is characteristic within the genus

and can be useful to diagnose it from its congeners (see

Diagnosis and identification key). According to Ibarra and

Stewart (1987), the type series of S. citurensis comprises the

holotype (FMNH 7585) and 150 paratypes from five different

localities. In this study, 55 paratypes were examined, and it

was possible to verify those characters and the ones included

in the original description, as well as in non-type material,

across the samples of several populations of the species.

Covain and Fisch-Muller (2007), in a study of the

taxonomy of the Loricariinae, discussed the characteristics

of Sturisomatichthys. They highlighted that S. citurensis is

morphologically divergent from its congeners in ‘‘having an

abdominal plate cover consisting of small platelet without

any particular organization’’. According to the authors, this

could differentiate this species; this characteristic was found

to be useful as a diagnostic character for Sturisomatichthys

when compared to Sturisoma. The size of the plates present in

S. citurensis is similar to those observed in S. caquetae,S.

guaitipan, new species, and S. tamanae. Covain and Fisch-

Muller (2007) also pointed out the fact that the other species

(three others at the time) do have a short snout and could

represent a species complex as seen in Farlowella, specifically

the F. curtirostra group that possesses a short and broader

snout. In any case, the authors assumed that the weakness of

the diagnosis of Sturisomatichthys could lead to a synonymy

with Sturisoma.

Covain et al. (2016) proposed that S. citurensis is the sister

group of the remaining Sturisomatichthys (their fig. 3);

although it belongs to the genus, the morphology of the

species indeed differs from that of its congeners.

Material examined.—FMNH 29500–29536, 29542–29559

(paratypes all mixed in several lots), 55 alc, 146.2–211.7

mm SL, Panama, Darien Province, Tuyra River basin, Cupe

River at Cituro, 8859037 00 N, 7983101200W, S. E. Meek and S. F.

Hildebrand, 1912; STRI 1881, 2 alc, Panama, Darien

Province, Tuira River, Capeti River, 88302700 N, 7783405200 W;

STRI 8369, 2 alc, 134.8–159.7 mm SL, Panama, Chucunaque

River, 881704600N, 7784501800 W; USNM 78361, 6 alc, 177.6–

Table 2. Morphometrics and meristics of Sturisomatichthys citurensis and S. dariensis. Morphometric values as percents of standard length (SL) and

head length (HL). SD ¼standard deviation.

S.citurensis (n¼102) S.dariensis (n¼4)

Min Max Mean SD Min Max Mean SD

Standard length 95.2 225.4 174.2 169.5 209.3 104

Head length 16.8 21.1 18.8 0.8 20.0 20.4 20.2 0.2

Predorsal length 29.4 33.8 31.5 0.9 30.1 31.2 30.9 0.5

Abdominal length 13.8 18.0 15.8 0.7 11.8 12.6 12.0 0.3

Trunk length 13.2 17.5 15.4 0.7 12.9 15.4 14.0 1.0

Dorsal fin length 18.8 28.5 23.8 1.9 28.5 30.8 29.9 1.2

Pectoral fin length 16.5 21.5 19.5 1.1 20.5 22.3 21.6 0.8

Pelvic fin length 15.1 20.9 17.8 1.2 15.9 17.9 17.0 0.8

Anal fin length 16.6 21.3 19.0 1.0 20.8 24.0 22.8 1.5

Head depth at supraoccipital tip 7.6 10.8 9.0 0.6 7.6 9.0 8.2 0.6

Body depth at dorsal-fin origin 8.1 12.2 10.7 0.7 10.1 11.2 10.7 0.5

Caudal peduncle depth 1.1 2.0 1.5 0.1 1.2 1.3 1.3 0.1

Cleithral width 10.2 14.3 12.0 0.6 9.9 10.5 9.9 0.3

Body width at dorsal-fin origin 10.9 15.7 13.8 0.8 10.4 11.8 11.3 0.7

Body width at anal-fin origin 10.1 14.2 12.2 0.8 9.0 10.0 9.8 0.5

Body width at caudal-fin origin 1.8 2.7 2.2 0.2 2.2 2.6 2.4 0.2

Head width 70.1 80.4 75.5 2.5 57.4 62.4 59.2 2.4

Interorbital distance 25.7 35.5 30.9 1.8 26.9 30.3 29.3 1.4

Orbital diameter 11.8 16.7 14.2 1.1 11.5 13.5 11.8 0.9

Snout length 56.4 66.7 61.5 2.0 61.7 63.2 62.5 0.6

Dentary length 11.0 19.0 16.0 1.7 9.1 11.7 10.9 1.1

Lower lip width 33.0 43.9 40.3 2.5 27.1 28.5 27.6 0.6

Plates on median series 13 17 14 14 15 15

Coalesced plates 13 17 15 16 17 17

Ventrolateral thoracic plates 6 11 9 7 9 7

Premaxillary teeth 61 124 107 54 61 54

Dentary teeth 63 100 78 34 47 41

Dorsal-fin rays i,7 i,7 i,7 i,7 i,7 i,7

Pectoral-fin rays i,6 i,6 i,6 i,6 i,6 i,6

Pelvic-fin rays i,5 i,5 i,5 i,5 i,5 i,5

Anal-fin rays i,5 i,5 i,5 i,5 i,5 i,5

Caudal-fin rays i,12,i i,12,i i,12,i i,12,i i,12,i i,12,i

776 Copeia 107, No. 4, 2019

214.6 mm SL, Panama, Canal Zone, Darien Province at

Aruza, Aruza River, 98420N, 738160W; USNM 78364, 1 of 7 alc,

225.4 mm SL, Panama, Darien Province, Chepo district,

Mamon River, 98100N, 79860W; USNM 78365, 1 alc, 1 CS,

171.4–175.4 mm SL, Panama, Darien Province, Rio Tuyra at

Marrigante, 88190N, 788180W; USNM 78366, 1 alc, 95.2 mm

SL, Panama, Darien Province, Capeti River, 8830N, 778340W;

USNM 78367, 3 of 13 alc, 159.6–167.0 mm SL, Panama,

Darien Province, Boca de Cupe district, Tuyra River,

8803000 00 N, 7783500000W; USNM 78368, 18 of 30 alc, 176.0–

191.0 mm SL, Panama, Darien Province, Yape River, 8870N,

778340W; USNM 293596, 1 of 5 alc, 105.2 mm SL, Panama,

Darien Province, Pirre River above El Real, above Tidal Zone,

Rio Tuyra drainage, Pacific drainage, 8870N, 778440W; USNM

316292, 1 alc, 184.2 mm SL, Panama, Darien Province, El

Real de Santamar´

ıa district, Pirre River 3-5 miles above El

Real, Battelle Memorial Institute, Pacific Northwest National

Laboratory, 8870N, 778440W; USNM 341994, 1 alc, 177.7 mm

SL, Colombia, Choc ´

o department, Prado district, creek

emptying on Prado River, 10 min helicopter flight from

Prado district, 68130N, 778240W; USNM 342003, 2 alc, 134.0–

140.4 mm SL, Colombia, Choc ´

o department, Riosucio

municipality, Nercua River tributary of Truando and Atrato

rivers, unnamed creek into upper Necua River, 7810N,

778300W; ZMA 119.125, 3 alc, 170.4–175.2 mm SL, Panama,

´balo River tributary of Samb ´

u River, 8840N, 788170W; ZMA

139.628, 1 alc, 125.5 mm SL, Panama, Bayano district, Maj´

River, 9850N, 788450W; ZMA 140.798, 1 alc, 179.1 mm SL,

Panama, Bayano district, Castrigandi River, 78570N, 808190W.

Sturisomatichthys dariensis (Meek and Hildebrand, 1913)

Figure 8, Table 2

Oxyloricaria dariensis Meek and Hildebrand, 1913: 81–82

(type locality: Tuyra River, Boca de Cupe, Panama;

holotype FMNH 7584, length 245 mm [TL]; original

description without specific remarks, focusing on measure-

ments, counts, and coloration of the specimens); 1916: 262

(synonym of Oxyloricaria panamensis). Eigenmann, 1922:

95 (synonym of Oxyloricaria panamensis). Miles, 1947: 115

(synonym of Oxyloricaria panamensis). Ibarra and Stewart,

1987: 63 (type series at FMNH; holotype missing or mixed

with paratypes; 39 paratypes reported from seven different

localities).

Sturisoma dariense (Meek and Hildebrand, 1913).—Isbr¨

ucker,

1979: 88 (valid as Sturisoma dariense); 1980: 94 (included in

Sturisoma as a member of the Harttiina; no diagnosis).

Isbr¨

ucker et al., 2001: 32 (included in the catalog of the

Loricariinae as valid in Sturisoma, German version).

Isbr¨

ucker, 2002: 29 (included in the catalog of the

Loricariinae as valid in Sturisoma, English version). Ferraris,

2003: 346 (genus distribution; synonymy; type material

location at FMNH 7584, 25.4 cm SL; maximum body size);

2007: 299 (type locality; distribution of the species).

Ghazzi, 2003: 182 (synonym of S. panamense); 2005: 564

(paratype used for comparison with S. kneri).

Sturisomatichthys dariensis (Meek and Hildebrand, 1913).—

Covain et al., 2016: 514 (valid as Sturisomatichthys dariense;

phylogenetic relationships of the Loricariinae using two

mitochondrial and one nuclear markers).

Diagnosis.—Sturisomatichthys dariensis is diagnosed from S.

caquetae,S. citurensis,S. guaitipan, new species, and S. tamanae

by the possesion of a rostrum (vs. rostrum absent) and

presence of a dorsolateral stripe (vs. dorsolateral stripe

absent). It is distinguished from other congeners by the

number of premaxillary teeth (54–61 vs. 36–50 in S. frenatus,

24–50 in S. leightoni, 37–53 in S. panamensis, 38–45 in S.

reinae, new species) and number of coalescent plates (16–17

vs. 13–16 in S. leightoni). Furthermore, it is distinguished

from S. aureus by lacking a transverse bar at the origin of the

dorsal fin (vs. transverse bar present) and lacking a dark

blotch on pectoral, pelvic, and anal fins (vs. blotch present).

It is distinguished from S. panamensis by lacking a blotch on

the pectoral fin (vs. blotch present) and by having reticula-

tions on supraoccipital, interorbital, and predorsal regions

(vs. reticulations absent). It is distinguished by its cleithral

width (9.9–10.5% SL vs. 10.5–12.8% SL in S. aureus; 8.3–9.6%

SL in S. reinae, new species; 7.7–9.2% SL in S. varii, new

species). It is distinguished from S. festivus by lacking

filaments on the dorsal, pectoral, pelvic, and anal fins (vs.

filaments present).

Description.—Morphometric data in Table 2. Body short and

slender, completely covered by dermal plates with small

hypertrophied odontodes, ill developed on sides of head, and

paired fins. Dorsal profile of head straight from tip of snout

to supraoccipital; concave from that point to dorsal-fin base,

and straight from that point to caudal-fin origin. Dorsal body

profile slightly concave posterior to dorsal fin, nearly straight

on last plates of caudal peduncle. Ventral body profile flat.

Head and predorsal region broader than remainder of body;

postdorsal region narrower towards caudal fin.

Head triangular and pointed. Rostrum present, short and

somewhat broad on its entire length. Ventral border of

rostrum flat. Orbital rim circular, without notch, dorsolater-

ally placed on head. Sphenotic with short lateral expansion.

Canal-bearing plate long, well developed, in contact with

anterior abdominal plates. Upper lip covered with small,

almost imperceptible plates; lower lip without such plates.

Lower lip covered with round small papillae. Rictal barbel

short, although more developed than in congeners, and free

from lip rim, not reaching anterior border of abdominal

plates but reaching canal-bearing plate. Teeth slender, bifid,

54–61 on premaxilla, 34–47 on dentary; cusps slightly

curved. Teeth in lower jaw somewhat larger than those in

upper jaw.

First two predorsal plates with two low, inconspicuous

ridges; nuchal plate at least twice size of predorsal plates.

Lateral plates carinate, forming two longitudinal sharp keels;

16–17 coalescent plates, beginning at 14–15 median plates.

Seven to nine straight, very broad lateral abdominal plates,

between pectoral- and pelvic-fin origins. Preanal plate

bordered anteriorly by three well-developed and similarly

sized plates; anteriormost plate sometimes not in contact

with preanal plate. Central abdominal plates, between lateral

abdominal plates, irregularly arranged; anterior abdominal

plates smaller and more numerous than posterior ones.

Posterior margin of dorsal fin slightly concave, un-

branched ray longest; dorsal-fin rays i,7. Pectoral-fin rays

i,6, unbranched ray longest, reaching between anterior third

and half length of pelvic fin. Pelvic-fin rays i,5, unbranched

ray longest, its tip reaching further than anterior third of

anal-fin. Anal-fin rays i,5, unbranched ray longer. Posterior

caudal-fin margin shallowly concave, filaments lacking on

both upper and lower rays, maybe due to damage.

Coloration in alcohol.—Ground color of dorsum and head

dark or pale brown, ventrally yellow. Dorsolateral stripe on

both sides of body. Stripe broader just at posterior border of

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 777

orbital rim, until 9

–10

median plate (Fig. 8), becoming

less conspicuous and thin towards caudal peduncle. Remnant

of stripe sometimes present on ventral side of rostrum (Fig.

8). Large, dark spots on membranes and rays of pectoral fin.

Dorsal fin with dark brown band along first two branched

rays, and with conspicuous dark spots along unbranched ray.

Anal fin hyaline. Caudal fin with dark bands along first two

or three branched rays, both upper and lower, and connected

at base of fin, forming V-shaped pattern. Dark brown spots

on edges of caudal lobes.

Sexual dimorphism.—Males show developed and numerous

hypertrophied odontodes along sides of head (Fig. 8);

odontodes absent from fin rays. No sexual dimorphism was

observed on teeth or thickening of rays on pectoral and

pelvic fins.

Distribution.—Sturisomatichthys dariensis occurs in the Tuyra

and Capeti rivers in the Pacific slope of Panama (Fig. 1).

Remarks.—Meek and Hildebrand (1913) described Sturisoma-

tichthys dariensis mainly based on body and head propor-

tions. Nevertheless, the authors did describe the coloration of

the species, mentioning the presence of dorsolateral stripe on

the sides of body (Fig. 8). Even though both S. dariensis and S.

citurensis are sympatric in the Tuyra and Cupe River basins,

and small tributaries in the Pacific slope in Panama (Figs. 1,

3), they can be differentiated by S. dariensis having 34–47

dentary teeth (vs. 63–100); presence of a rostrum (vs. rostrum

absent); presence of a dorsolateral stripe (vs. stripe absent);

conspicuous and numerous spots on rays and membranes on

dorsal and pectoral fins (vs. few and light spots on fins).

Meek and Hildebrand (1916: 262) placed S. dariensis in the

synonymy of S. panamensis. The authors stated, ‘‘A study of

specimens from Colombia and Ecuador along with our

specimens shows that there are quite a number of synonyms

under this species [namely S. panamensis]. The fact that the

lateral keels in the young are not completely united while in

the adult they are, in part at least accounts for the numerous

names that have been proposed.’’ Eigenmann (1922), Miles

(1947), and Dahl (1971) followed this synonymy. Isbr¨

ucker

(1979) assumed S. dariensis as valid and as belonging to

Sturisoma, but did not elaborate on the reason for the

resurrection of the species. In several studies dealing with

the classification of the Loricariinae (and the Loricariidae), S.

dariensis continued to be assumed as valid (e.g., Isbr¨

ucker,

Fig. 8. Sturisomatichthys dariensis, STRI 8386, 206.0 mm SL, Panama, Darien Province, Chucunaque River basin, Tupisa River.

778 Copeia 107, No. 4, 2019

1980, 2002; Isbr¨

ucker et al., 2001; Ferraris, 2003, 2007;

Covain and Fisch-Muller, 2007; Covain et al., 2016).

On the other hand, Ghazzi (2003) followed the synonymy

of S. dariensis with S. panamensis proposed by Meek and

Hildebrand (1916). The author stated that when she

compared the juvenile paratypes of S. dariensis and the

holotype of S. panamensis (90.7 mm SL) examined in her

study, the specimens were really similar, and she assumed

them to be the same species. According to Ghazzi (2003),

there is only one species of the genus in the Tuyra River basin

in Panama (i.e., S. panamensis),afactthatwasnot

corroborated here. According to the material examined,

there are three species of Sturisomatichthys in that basin: S.

citurensis,S. dariensis, and S. panamensis (Figs. 1, 3).

According to Ibarra and Stewart (1987), there are 39

paratypes from seven different localities as part of the type

series of S. dariensis. The holotype (FMNH 7584) is reported

to be missing, which was corroborated by the records of the

FMNH collection website and personal observation by the

authors. A designation of a neotype is not included here due

to the existence of paratypes of the species, which are a good

sample for the ontogenetic variation of S. dariensis, as for its

distribution across localities in the Pacific versant in Panama.

Material examined.—STRI 8386, 1 alc, 206.0 mm SL, Panama,

Darien Province, Chucunaque River basin, Tupisa River,

881204.6 00 N, 8284102.400W; USNM 78373, 2 alc, 169.5–209.3

mm SL, Panama, Darien Province, Yape River, 8870N,

778350W; USNM 293273, 1 CS, 192.9 mm SL, Panama,

Darien Province, Rio Tuyra 2–3 km above Pinogana (Pacific),

8860N, 778470W.

Sturisomatichthys festivus (Myers, 1942)

Figures 9, 10; Table 3

Sturisoma festivum Myers, 1942: 100 (type locality: Rio

Monay, 35 km north of Trujillo, Motatan system, Mar-

acaibo basin, Venezuela; holotype SU 36505, length 169

mm SL; nine paratypes from the same locality; original

description). Schultz, 1944: 337 (distributed at Maracaibo

Lake basin; identification key for the Sturisoma of the

Maracaibo Lake basin). B¨

ohlke, 1953: 46 (citation of type

series). Isbr¨

ucker, 1980: 94 (included in Sturisoma as a

member of the Harttiina; no diagnosis). Ferraris and Vari,

1992: 31 (two paratypes at USNM reporting SL; remarks of

the former SU number and presence of the holotype in

that collection). Galvis et al., 1997: 97 (common name;

short description; habitat; distributed only at Maracaibo

Lake Basin; photo of a live specimen in lateral view).

Isbr¨

ucker et al., 2001: 32 (included in the catalog of the

Loricariinae as valid in Sturisoma, German version).

Isbr¨

ucker, 2002: 29 (included in the catalog of the

Loricariinae as valid in Sturisoma, English version). Ferraris,

2003: 347 (genus distribution; synonymy; type material

location at SU 36505, 16.9 cm SL). Ghazzi, 2003: 193

(belonging to a new genus, not to Sturisoma). Lasso et al.,

2004: 129 (as junior synonym of Sturisoma kneri [De Filippi,

in Tortonese, 1940]). Ghazzi, 2005: 564 (paratypes used for

comparison with S. kneri). Ferraris, 2007: 299 (type locality;

distribution of the species). Ortega-Lara et al., 2012: 91

(common name; specimens in Colombian and Venezuelan

museums). DoNascimiento, 2016: 534 (corrigendum of

Lasso et al., 2004, where the species was omitted; type

locality based on Ferarris, 2007).

Sturisomatichthys festivus (Myers, 1942).—Covain et al., 2016:

514 (valid as Sturisomatichthys festivum;phylogenetic

relationships of the Loricariinae using two mitochondrial

and one nuclear markers). DoNascimiento et al., 2018: 77

(as present in Colombia, Caribbean slope).

Diagnosis.—Sturisomatichthys festivus is diagnosed from its

congeners by the possession of long filaments, generally

exceeding half of SL, on leading ray of all fins, including

upper and lower rays on caudal fin (vs. filaments absent or

present only on pectoral and/or caudal fin), and dorsolateral

stripe not continuing on caudal peduncle (vs. dorsolateral

stripe absent or, when present, continuing onto caudal

peduncle as continuous or discontinuous stripe). From S.

caquetae,S. citurensis,S. guaitipan, new species, and S.

tamanae, it is distinguished by having a rostrum (vs. rostrum

absent) and a dorsolateral stripe (vs. dorsolateral stripe

absent).

Description.—Morphometric data in Table 3. Body short,

completely covered by dermal plates with small hypertro-

phied odontodes, ill developed, but perceptible on sides of

head. Dorsal profile of head straight from tip of snout to

origin of dorsal fin; concave from that point to caudal-fin

origin. Ventral body profile flat. Head and predorsal region

broader than remainder of body; postdorsal region narrower

towards caudal fin.

Head triangular. Rostrum present, short and broad on its

entire length. Ventral border of rostrum flat. Orbital rim

circular, without notch, dorsolaterally placed on head.

Sphenotic with short lateral expansion. Canal-bearing plate

broad and well developed, in contact with anterior abdom-

inal plates, anterior to branchiostegal membrane, not

reaching rictal barbel. Upper lip covered with small, almost

imperceptible plates; lower lip without such plates. Lower lip

covered with round, small, almost imperceptible papillae.

Rictal barbel short, free from lip rim, never reaching anterior

border of abdominal plates, nor surpassing posterior border

of lower lip. Teeth slender, bifid, 37–55 on premaxilla, 33–42

on dentary; cusps slightly curved. Teeth in lower jaw

somewhat larger than those in upper jaw.

Predorsal plates with two low, inconspicuous ridges;

nuchal plate twice as large as predorsal plates and more

developed, causing greater body depth (Figs. 9, 10). Lateral

plates carinate, forming two longitudinal sharp keels; 16–18

coalescent plates, beginning at 14–15 median plates. Seven to

eight straight lateral abdominal plates, between pectoral- and

pelvic-fin origins, with same size of central abdominal plates.

Preanal plate bordered anteriorly by three well-developed

and similarly sized plates; anteriormost plate not in contact

with preanal plate. Anterior abdominal plates smaller and

more numerous than posterior ones.

Posterior margin of dorsal fin straight, unbranched ray

longest; dorsal-fin rays i,7. Pectoral-fin rays i,6, unbranched

ray longest, reaching between half and distal tip of pelvic fin.

Pelvic-fin rays i,5, unbranched ray longest, its tip reaching

half length of anal fin. Anal-fin rays i,5, unbranched ray

longest. Posterior caudal-fin margin deeply concave, filament

only on upper ray (perhaps damaged in lower ray).

Coloration in alcohol.—Ground color of dorsum, head, and

ventral surface pale brown or yellow. Dorsolateral stripe

broader from anterior border of first predorsal plate to dorsal-

fin origin (Fig. 10), becoming less conspicuous and thin

towards caudal peduncle. Remnant of dark stripe sometimes

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 779

present on ventral side of rostrum, not separated as observed

on both sides of body but as single dark spot, occupying

entire ventral surface of rostrum (Fig. 10). Large, dark spots

on membranes and rays of pectoral, pelvic, and anal fins, not

occupying entire fin. Dorsal fin with dark brown band along

first two branched rays and with conspicuous dark spots

along unbranched ray. Caudal fin with dark bands along first

two or three branched rays, both upper and lower, and

connected at base of fin, forming V-shaped pattern. Dark

brown spots on edges of caudal lobes.

Sexual dimorphism.—The males examined showed sexually

dimorphic short hypertrophied odontodes on the sides of the

head (Fig. 9).

Distribution.—Sturisomatichthys festivus occurs in the Mara-

caibo Lake basin in Venezuela, and Catatumbo River in

Colombia (Fig. 1).

Remarks.—Myers (1942) described Sturisoma festivum from

the Monay River, Motatan System in the Maracaibo Lake

basin, Venezuela. The description highlights the fact that the

species has, in both juvenile and adult males and females,

filamentous extensions on the first ray of all fins (Figs. 9, 10).

The author also mentioned the presence of dorsolateral stripe

(Fig. 10); nevertheless, Myers (1942) stated that the colora-

tion was only observed in one juvenile specimen, but present

results confirm the stripe to be present in adults as well.

Finally, the author compared S. festivus to S. frenatus and S.

panamensis (assuming the former as synonym of the latter),

but differentiating both species by means of corporal

proportions and the fact that S. festivus has filamentous

extensions on the fins.

The taxonomic history of the species is not extensive.

Since its description, it has always been assumed as valid and

belonging to Sturisoma, until the study of Covain et al. (2016)

which placed it in Sturisomatichthys. Both paratypes cited by

Ferraris and Vari (1992) deposited at the USNM were

examined in this study. When compared to non-type

material, all of the characters offered by Myers (1942) were

observed across the samples.

Material examined.—AUM 22138, 8 alc, 151.8–161.3 mm

SL,Venezuela, Zulia state, Lake Maracaibo basin, Escalante

River basin, Onia River at El Padre Channel on road from

highway 2 to town of KM 35, 88450N, 718450W; AUM 22148,

1 alc, 115.7 mm SL, Venezuela, Zulia state, Escalante River,

Lake Maracaibo basin, Chamita River on road from highway

2 to town of Blanco channel, 88540N, 718490W; AUM 22184,

3 alc, Venezuela, Zulia state, Lake Maracaibo basin, Cata-

tumbo River basin, Zulia River, channel North of highway 6

bridge near Tachira-Zulia state line on small road that corsses

creek, 88300N, 728200W; CAS 136506, 4 alc, 121.8–183.8 mm

SL, paratypes, Venezuela, Trujillo state, Lake Maracaibo basin,

Monay River, 35 km north of Trujillo, 98220N, 708260W; CAS

Fig. 9. Sturisomatichthys festivus, paratype of Sturisoma festivum, USNM 130637, 160.0 mm SL, Venezuela, Trujillo state, Lake Maracaibo basin,

Monay River, 35 km north of Trujillo.

780 Copeia 107, No. 4, 2019

168512, 1 alc, 1 CS, 159.4– 162.9 mm SL, Venezuela, Lake

Maracaibo basin, Motatan River, 982303600N, 7083503000 W;

USNM 130637, 2 alc, 142.8–160.0 mm SL, paratypes,

Venezuela, Trujillo state, Lake Maracaibo basin, Monay River,

35 km north of Trujillo, 98220N, 708260W.

Sturisomatichthys frenatus (Boulenger, 1902)

Figures 11, 12; Table 3

Loricaria frenata Boulenger, 1902: 69–70 (type locality: San

Javier, Salidero, and Durango River, Pacific slope of

Ecuador; without designation of types). Meek and Hilde-

brand, 1916: 261 (junior synonym of Oxyloricaria pana-

mensis).

Oxyloricaria frenata (Boulenger, 1902).—Regan, 1904: 302,

plate VXIII, fig. 2 (description of the species; report of eight

specimens from northwestern Ecuador used for the

original description; total length of larger specimen, 250

mm TL; drawings of lateral view, and ventral and dorsal

view of anterior portion of the body). Meek and Hilde-

brand, 1916: 262 (junior synonym of Oxyloricaria pana-

mensis).

Sturisoma frenatum (Boulenger, 1902).—Eigenmann, 1910:

416 (identification key for Sturisoma; valid as Sturisoma

frenatum; reference to Regan, 1904). Isbr¨

ucker, 1979: 113

(designation of lectotype for the species: BMNH

1901.8.3:29, male, 236 mm SL); 1980: 94 (included in

Sturisoma as a member of the Harttiina; no diagnosis;

reference to Regan [1904] regarding specimens from the

original description, and Isbr¨

ucker [1979] where it is stated

that the lectotype is from Salidero, as well as five

paralectotypes, two from St. Javier, and three from

Durango River); 1981a: 88 (as valid in Sturisoma). Isbr¨

ucker

et al., 2001: 32 (included in the catalog of the Loricariinae

as valid in Sturisoma, German version). Isbr¨

ucker, 2002: 29

(included in the catalog of the Loricariinae as valid in

Sturisoma, English version). Ferraris, 2003: 347 (distribu-

tion; synonymy; type material location; maximum size;

common name in Ecuador); 2007: 299 (type locality;

distribution of the species). Ghazzi, 2003: 188 (belonging

to a new genus and not to Sturisoma); 2005: 564 (type series

used for comparison with S. kneri).

Sturisomatichthys frenatus (Boulenger, 1902).—Covain et al.,

2016: 514 (valid as Sturisomatichthys frenatum; phylogenet-

ic relationships of the Loricariinae subfamily using two

mitochondrial and one nuclear markers; appears as sister to

S. dariensis).

Diagnosis.—Sturisomatichthys frenatus is diagnosed from its

congeners by the possession of a longitudinal dark stripe on

the first two or three dorsal-fin rays, reaching their distal tip,

and extending to the base of remaining rays (vs. stripe absent

or not reaching distal tip of rays and restricted to first two

dorsal-fin rays, nor extended to the base of remaining rays),

and pelvic and anal fins with dark blotches occupying the

entire fin (vs. blotches absent or light spots not occupying

the entire fin). Moreover, it is distinguished from S. caquetae,

S. citurensis,S. guaitipan, new species, and S. tamanae by

Fig. 10. Sturisomatichthys festivus, AUM 22138, 161.3 mm SL, Venezuela, Zulia state, Lake Maracaibo basin, Escalante River basin, Onia River at El

Padre Channel on road from highway 2 to town of Km 35.

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 781

having a rostrum (vs. rostrum absent) and a dorsolateral

stripe (vs. dorsolateral stripe absent).

Description.—Morphometric data in Table 3. Body short and

low, completely covered by dermal plates with small, ill-

developed hypertrophied odontodes, not visible on sides of

head nor on paired fins. Dorsal profile of head slightly

concave from tip of snout to interorbital portion, straight

from that point to origin of dorsal fin; concave from that

point to caudal-fin origin. Ventral body profile flat. Head and

predorsal region broader than remainder of body; postdorsal

region narrower toward caudal fin.

Head triangular, elongated. Rostrum present, its ventral

border flat. Orbital rim circular, without notch, dorsolaterally

placed on head. Sphenotic with short lateral expansion.

Canal-bearing plate broad and well developed, in contact

with anterior abdominal plates, anterior to branchiostegal

membrane, not reaching to rictal barbel. Upper lip covered

with small, almost imperceptible plates; lower lip without

such plates. Lower lip covered with round, small, almost

imperceptible papillae. Posterior border of lower lip irregular,

with small papillae along its entire border. Rictal barbel short,

free from lip rim, never reaching to anterior border of

abdominal plates, nor surpassing posterior border of lower

lip. Teeth slender, bifid, 36–50 on premaxilla, 31–46 on

dentary; cusps slightly curved. Teeth in lower jaw larger than

those in upper jaw.

Predorsal plates with two low, inconspicuous ridges;

nuchal plate larger than predorsal plates. Lateral plates

carinate, forming two longitudinal weak keels; 16–18

coalescent plates, beginning at 14–17 median plates. Six to

nine straight lateral abdominal plates, between pectoral- and

pelvic-fin origins, longer than central abdominal plates.

Preanal plate bordered anteriorly by five or six well-

developed and similarly sized plates, all in contact with

preanal plate. Central abdominal plates, between lateral

abdominal plates, irregular and similar in size and shape as

anterior and posterior ones. Anterior abdominal plates of

similar size and number as posterior ones.

Posterior margin of dorsal fin straight, unbranched and

first and second branched rays longest; dorsal-fin rays i,7.

Pectoral-fin rays i,6, unbranched ray longest, reaching to or

slightly surpassing anterior third of pelvic fin. Pelvic-fin rays

i,5, unbranched ray longest, its tip reaching anterior third of

anal-fin. Anal-fin rays i,5, unbranched ray longest. Posterior

caudal-fin margin deeply concave, filaments absent on upper

and lower rays, maybe due to damage.

Coloration in alcohol.—Ground color of dorsum and head

pale or dark brown; pale yellow ventrally. Dorsolateral stripe

broader from posterior border of orbital rim to two plates

Table 3. Morphometrics and meristics of Sturisomatichthys festivus and S. frenatus. Morphometric values as percents of standard length (SL) and

head length (HL). SD ¼standard deviation.

S.festivus (n¼10) S. frenatus (n¼15)

Min Max Mean SD Min Max Mean SD

Standard length 115.7 183.8 159.7 163.8 239.7 200.4

Head length 18.3 20.4 18.9 0.6 19.1 23.1 20.6 0.9

Predorsal length 28.4 31.4 30.2 1.0 30.5 32.3 31.3 0.5

Abdominal length 9.9 11.3 10.7 0.5 10.2 12.9 12.2 0.7

Trunk length 13.4 15.7 14.7 0.8 13.8 15.7 15.1 0.6

Dorsal fin length 36.5 42.7 40.1 3.1 22.2 27.1 24.6 1.4

Pectoral fin length 24.6 39.3 32.6 5.4 17.4 23.3 19.2 1.3

Pelvic fin length 17.5 21.5 19.7 1.1 14.4 17.2 15.8 0.7

Anal fin length 24.8 33.5 32.1 3.7 17.2 20.5 19.5 1.0

Head depth at supraoccipital tip 8.6 10.3 9.5 0.5 8.0 9.3 8.6 0.3

Body depth at dorsal-fin origin 12.0 14.7 13.2 0.9 9.8 12.6 10.7 0.7

Caudal peduncle depth 1.3 1.7 1.4 0.2 1.0 1.7 1.3 0.2

Cleithral width 10.3 11.7 10.9 0.4 8.5 10.5 9.7 0.5

Body width at dorsal-fin origin 11.1 13.0 12.1 0.5 10.7 13.0 11.8 0.7

Body width at anal-fin origin 10.5 12.5 11.5 0.6 9.6 10.7 10.1 0.3

Body width at caudal-fin origin 2.2 3.1 2.7 0.3 1.9 2.5 2.1 0.2

Head width 65.3 73.7 69.3 2.7 48.4 62.3 59.5 3.8

Interorbital distance 33.3 40.2 37.2 2.4 25.8 33.3 30.3 2.0

Orbital diameter 11.7 16.0 13.6 1.1 9.6 13.8 11.6 1.0

Snout length 60.7 66.4 63.9 1.8 55.9 68.0 62.7 3.0

Dentary length 9.5 14.2 11.1 1.3 8.6 11.7 10.7 1.0

Lower lip width 30.9 34.5 32.4 1.2 25.1 33.6 28.3 2.7

Plates on median series 14 15 15 14 17 16

Coalesced plates 16 18 17 16 18 16

Ventrolateral thoracic plates 7 8 7 7 9 8

Premaxillary teeth 37 55 39 36 50 41

Dentary teeth 33 42 39 31 46 35

Dorsal-fin rays i,7 i,7 i,7 i,7 i,7 i,7

Pectoral-fin rays i,6 i,6 i,6 i,6 i,6 i,6

Pelvic-fin rays i,5 i,5 i,5 i,5 i,5 i,5

Anal-fin rays i,5 i,5 i,5 i,5 i,5 i,5

Caudal-fin rays i,12,i i,12,i i,12,i i,12,i i,12,i i,12,i

782 Copeia 107, No. 4, 2019

posterior to dorsal-fin origin (Figs. 11, 12), becoming less

conspicuous and thin towards caudal peduncle. Remnant of

dark stripe sometimes present on ventral side of rostrum,

separated as on both sides of body (Fig. 12). Large, dark

blotch on pectoral, pelvic, and anal fins, its length decreasing

from first to last ray on each fin (Figs. 11, 12). Dorsal fin with

dark brown stripe along first two branched rays, without

conspicuous dark spots along unbranched ray. Caudal fin

with dark stripe along first two or three branched rays on

upper and lower lobes, and connected at base of fin, forming

V-shaped pattern. Two or three white ocelli at distal portions

of two or three first upper and lower caudal-fin rays (Fig. 12).

Sexual dimorphism.—The specimens examined show sexual

dimorphism on the sides of the head, consisting of short, ill-

developed odontodes in males, as stated by Boulenger (1902;

Fig. 11).

Distribution.—Sturisomatichthys frenatus occurs in the Pacific

slopes of Ecuador, in the Esmeraldas, San Javier, Salidero, and

Durango rivers, and the Patia River in Colombia (Fig. 2).

Remarks.—The original description offered by Boulenger

(1902) highlights general aspects of the species, initially

described as Loricaria frenata. The author referred to the

dorsolateral stripe on the sides of the head, which was

corroborated on both type and non-type material (Figs. 11,

12). Additionally, Boulenger (1902) also described the dark

blotches on fins, which are present on some of the species of

Sturisomatichthys (see identification key).

Regan (1904) transferred the species to Oxyloricaria and

made a redescription including eight of the specimens used

by Boulenger (1902) on the original description. That second

description is similar to the original one, and Regan included

no additional characters. Nevertheless, Eigenmann (1910)

proposed Oxyloricaria as a junior synonym of Sturisoma and

assumed S. frenatus as valid and belonging to that genus.

Meek and Hildebrand (1916) did not follow Eigenmann’s

synonymy and continued to assume Oxyloricaria as valid; the

authors included S. frenatus as a junior synonym of S.

panamensis. Eigenmann (1922) did not consider the species

as valid and included the species as a synonym of Sturisoma

panamense.

The original description offered by Boulenger (1902)

indicated several specimens from St. Javier, Salidero and

Durango rivers, northwester Ecuador, Pacific versant. Never-

theless, the author indicated no holotype. Isbr¨

ucker (1979)

designated a lectotype for the species (BMNH 1901.8.3) and

assumed S. frenatus as valid and belonging to Sturisoma. The

type series was examined here, and even though the

specimens were collected more than 100 years ago, the

typical coloration on the fins and general characteristics

described by Boulenger (1902) are still visible (Fig. 11).

The species is distributed only in the Pacific versant of

northwestern Ecuador and southwestern Colombia (Fig. 2),

Fig. 11. Sturisomatichthys frenatus, lectotype of Loricaria frenata, BMNH 1901.8.3.29, 232.4 mm SL, Ecuador, Salidero, NW Ecuador.

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 783

contrary to the distribution of Eigenmann (1922) which

included the Atrato (northwestern Colombia) and Tuyra

basins (Pacific versant of Panama). Finally, this is the first

record of Sturisomatichthys frenatus in the Patia River,

southwestern Colombia contrary to Ortega-Lara et al.

(2006a) who did not assume the species to be present at

that basin.

Material examined.—BMNH 1901.8.3.29, lectotype, 232.4

mm SL, Ecuador, Salidero, NW Ecuador, elevation 350 feet,

1840N, 79820W; BMNH 1901.6.27:23–4, 2 alc, 179.9–227.8

mm SL, paralectotypes, same date as lectotype; BMNH

1901.8.3:30–31, 2 alc, 226.4–228.5 mm SL, paralectotypes,

same data as lectotype; CAS 13642, 2 alc, 202.3–239.7 mm

SL, Colombia, Nari˜

no department, Patia River basin, Magui

River, 184505700N, 7881005800 W; CAS 13643, 3 alc, 165.5–194.0

mm SL, Colombia, Nari˜

no department, Patia River between

Magui River and Telembi River, 18490N, 788160W; MHNG

2674.055, 1 alc, 198.5 mm SL, Ecuador, Esmeraldas state,

Bogota

´River at Carondelet, 1870N, 788450W; USNM 288089,

2 alc, 200.4–214.0 mm SL, Ecuador, Esmeraldas state, Bogota

River, Sabalera estuary, 600 mts. near campament La

Chiquita, road to Ricaurte, 1820N, 788500W; USNM 341993,

2 of 3 alc, 1 CS, 163.8–181.8 mm SL, Colombia, Nari˜

department, Teresita district, Salado River, 08570N, 75825 0W.

Sturisomatichthys kneri (Ghazzi, 2005)

Loricaria kneri De Filippi, in Tortonese, 1940.—Isbr¨

ucker,

1981a: 88 (listed as nominal species in Loricaria). Isbr ¨

ucker

et al., 2001: 29 (name listed as valid in Sturisoma, German

version). Isbr¨

ucker, 2002: 22 (name listed as valid in

Sturisoma, English version).

Loricaria kneri De Filippi, in Tortonese, 1940: 137 (type

locality: ‘‘Lago di Maracaibo,’’ Venezuela; name unavail-

able); Eschmeyer, 1998 (name unavailable, listed in type

catalog).

Sturisoma kneri (De Filippi, in Tortonese, 1940).—Isbr¨

ucker,

1979: 88 (unavailable name used in new combination);

1980: 94 (included in Sturisoma as a member of the

Harttiina; no diagnosis); 1981b: 88 (transferred from

Loricaria to Sturisoma).

Sturisoma kneri Ghazzi, 2005: 559, fig. 1 (type locality: Lago

de Maracaibo, Venezuela; holotype MZUT 764, 265.6 mm

SL; redescription of the species). Ferraris, 2007: 299 (type

locality; distribution of the species).

Sturisomatichthys kneri (Ghazzi, 2005).—Covain et al., 2016:

514 (valid as Sturisomatichthys kneri; phylogenetic relation-

ships of the Loricariinae subfamily using two mitochon-

drial and one nuclear markers; not included in

phylogenetic analysis).

Diagnosis.—Sturisomatichthys kneri is diagnosed from S.

caquetae,S. citurensis,S. guaitipan, new species, and S. tamanae

by possessing a rostrum (vs. rostrum absent) and a dorsolat-

eral stripe (vs. dorsolateral stripe absent). From the remaining

congeners, it is distinguished by the number of premaxillary

teeth (21–35 vs. 47–69 in S. aureus; 54–61 in S. dariensis; 37–

55 in S. festivus; 36–50 in S. frenatus; 37–53 in S. panamensis;

38–45 in S. reinae, new species) and dentary teeth (21–30 vs.

Fig. 12. Sturisomatichthys frenatus, MHNG 2674.055, 198.5 mm SL, Ecuador, Esmeraldas state, Bogota

´River at Carondelet.

784 Copeia 107, No. 4, 2019

40–56 in S. aureus; 34–47 in S. dariensis; 33–42 in S. festivus;

31–46 in S. frenatus; 32–44 in S. panamensis). From S. leightoni,

it is distinguished by the presence of a continuous dorsolat-

eral stripe (vs. dorsolateral stripe discontinuous) and interor-

bital distance (32.7–36.7% HL vs. 25.4–32.6% HL). From S.

varii, new species, it is distinguished by head depth (7.5–8.0%

SL vs. 9.3–11.3% SL), orbital diameter (12.2–13.2% HL vs.

13.6–18.1% HL), and premaxillary teeth (21–30 vs. 30–44).

From S. festivus it is further distinguished by lacking

filaments on dorsal, pectoral, pelvic, and anal fins (vs.

filaments present).

Description.—The reader is referred to the original description

of the species (in Ghazzi, 2005) for morphometrics and

meristics. Because the description and diagnosis of the

species is recent, it will not be redescribed here. The

diagnostic characters used here for the species were observed

from a CS paratype (see Material Examined for the species), a

single alcohol-preserved specimen, and characters of external

morphology provided by Ghazzi (2005) in the original

description.

Distribution.—Sturisomatichthys kneri occurs in Lake Maracai-

bo basin in Venezuela (Fig. 1).

Remarks.—Ghazzi (2005) redescribed Sturisoma kneri and

assumed the name Loricaria kneri as unavailable. The author

retained the specific epithet of the species and its holotype to

maintain taxonomic stability for the taxon (Ghazzi, 2005:

559) and included it in Sturisoma to make the new name

available. Ghazzi stated that even though Isbr¨

ucker (1980,

1981b) and Isbr¨

ucker et al. (2001) treated the name Sturisoma

kneri as valid, this mere citation is not enough (Article 11.5.2

of the ICZN, 1999). Thus, the validation of the name had to

be done by the redescription and diagnosis of the species

within the genus.

The only known specimens belonging to S. kneri are those

included in the type series in the original description (n¼5).

Only an additional specimen was available for this study.

Thus, data included in the diagnosis and comparison with

congeners of Sturisomatichthys and identification key were

taken from the original description by Ghazzi (2005).

Material examined.—AUM 22165, 1 alc (photo), 201.4 mm

SL, Venezuela, Zulia state, Escalante River on road from Santa

Barbara to La Solita, north of Las Casas, 98000N, 71854 0W;

MCNG 33535, 1 CS, paratype, Venezuela, Zulia state, Lake

Maracaibo basin, Urumana Creek, Cataneja farm, Goajira,

118050N, 718510W.

Sturisomatichthys leightoni (Regan, 1912)

Figure 13, Table 4

Oxyloricaria leightoni Regan, 1912: 669, plate LXXVII, fig. 2

(type locality: Honda, Colombia; no types assigned;

original description without specific remarks, focusing on

measurements and coloration of the specimens; two

specimens of 55 mm and 65 mm TL).

Sturisoma leightoni (Regan, 1912).—Eigenmann, 1920b: 11 (as

distributed at the San Juan River); 1922: 96, plate XVI, figs.

1 and 2 (as distributed on the upper Magdalena, San Juan

River and east of Bogota in Colombia; a total of 94

specimens from IUM and CM, from the upper Magdalena,

one of them from the San Juan River, which probably is a

specimen of S. tamanae; specimens from Cumaral, Meta,

Colombia which the author reported to be ‘‘slightly

different’’ from the rest). Fowler, 1942: 132 (distributed at

the San Juan, Cauca, Meta and Magdalena rivers, following

Regan [1912]). Miles, 1947: 114 (identification key of the

Loricariidae; distribution of the species; reproduction of

the distribution proposed by Regan [1912]). Dahl, 1971: 95

(present at upper Cauca and Magdalena; maximum size 18

cm of TL; identification key for Sturisoma along with S.

aureus and S. panamensis). Roma

´n-Valencia, 1995: 14

(distributed at La Vieja River, upper Cauca, Colombia).

Lasso et al., 2004: 129 (not distributed in Venezuela).

Sturisomatichthys leightoni (Regan, 1912).—Isbr¨

ucker, 1979:

88 (original description of Sturisomatichthys; designation as

type species of the genus; designation of lectotype by

subsequent designation: BMNH 1909.7.23:45, 57 mm SL);

1980: 92–93 (included in Sturisomatichthys as a member of

the Harttiina; no diagnosis; reference of Isbr¨

ucker [1979]

for the designation of lectotype). Isbr¨

ucker et al., 2001: 32

(included in the catalog of the Loricariinae as valid in

Sturisomatichthys, German version). Isbr¨

ucker, 2002: 29

(included in the catalog of the Loricariinae as valid in

Sturisomatichthys, English version). Ferraris, 2003: 347

(distribution; synonymy; type material location; maxi-

mum size; common name in Colombia). Maldonado-

Ocampo et al., 2005: 138, figs. 124 and 135 (common

names; reproduction of the original description; biology

and ecology of the species from previous reports; distribu-

tion from previous reports; localities from previous reports;

specimens in Colombian museums; drawing in lateral

view; distribution map). Mojica et al., 2006a: 32 (distrib-

uted in the Magdalena-Cauca basin; catalog numbers in

Colombian museums). Ortega-Lara et al., 2006b: 48

(distributed in the Cauca basin in Colombia; specimens

in Colombian museums). Villa-Navarro et al., 2006: 14

(distributed in the Magdalena River basin in Colombia;

specimens in Colombian museums). Maldonado-Ocampo

et al., 2008: 195 (distributed in the Magdalena and Pacific

basins in Colombia; citation of type series). Covain and

Fisch-Muller, 2007: 37 (type species of Sturisomatichthys;

citation of lectotype; remarks about the genus). Ferraris,

2007: 300 (type locality; distribution of the species).

Covain et al., 2016: 503 (valid as Sturisomatichthys leightoni;

phylogenetic relationships of the Loricariinae subfamily

using two mitochondrial and one nuclear markers; appears

as sister group to S. aureus). DoNascimiento et al., 2018: 77

(as present and endemic in Colombia, Magdalena-Cauca

River basin and Pacific slope).

Diagnosis.—Sturisomatichthys leightoni is diagnosed from its

congeners, except S. caquetae,S. citurensis,S. guaitipan, new

species, and S. tamanae, by having discontinuous, ill-marked,

and thin dorsolateral stripe (vs. continuous, well-marked,

and broad dorsolateral stripe). It is distinguished from the

above species by having a rostrum (vs. rostrum absent). It is

also distinguished from congeners by the number of

coalescent plates (13–16 vs. 16–17 in S. dariensis; 16–18 in

S. festivus; 16–18 in S. frenatus; and 16–19 S. varii, new

species). It is further distinguished by its cleithral width

(11.0–14.1% SL vs. 8.5–10.5% SL in S. frenatus; 8.4–10.5% SL

in S. panamensis; 8.3–9.6% SL in S. reinae, new species; and

7.7–9.2% SL in S. varii, new species). Furthermore, it is

distinguished from S. aureus by lacking large, dark blotches

on dorsal, pectoral, pelvic, and anal fins occupying almost

the entire fin (vs. blotches present) and by having reticula-

tions on predorsal, supraoccipital, and interorbital region (vs.

reticulations absent). It is distinguished from S. panamensis

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 785

by lacking a dark spot on the pectoral fin and a stripe along

first two rays of dorsal fin (vs. spot and stripe present) and by

having reticulations on supraoccipital, interorbital, and

predorsal region (vs. reticulations absent). From S. festivus,

it is distinguished by lacking filaments on dorsal, pectoral,

pelvic, and anal fins (vs. filaments present).

Description.—Morphometric data in Table 4. Body short and

low, completely covered by dermal plates with short

hypertrophied odontodes, visible on both sides of the head,

but not on paired fins. Dorsal profile of head straight from tip

of snout to origin of dorsal fin; concave from that point to

caudal-fin origin. Ventral body profile flat. Head and

predorsal region broader than remainder of body.

Head triangular. Rostrum present, its ventral border flat.

Orbital rim circular, without notch, dorsolaterally placed on

head. Sphenotic with reduced, almost absent lateral expan-

sion. Canal-bearing plate broad and well developed, in

contact with anterior abdominal plates, anterior to bran-

chiostegal membrane, not reaching rictal barbel. Upper lip

covered with small, almost imperceptible plates; lower lip

without such plates. Lower lip covered with round, small,

almost imperceptible papillae. Posterior border of lower lip

irregular, with small papillae present along its entire border.

Rictal barbel short, free from lip rim, never reaching anterior

border of abdominal plates, nor surpassing posterior border

of lower lip. Teeth slender, bifid, 24–50 on premaxilla, 22–44

on dentary; cusps slightly curved. Teeth in lower jaw larger

than those in upper jaw.

Predorsal plates with two low, inconspicuous ridges;

nuchal plate twice size of predorsal plates. Lateral plates

carinate, forming two longitudinal keels; 13–16 coalescent

plates, beginning at 13–17 median plates. Five to eight

straight lateral abdominal plates, between pectoral- and

pelvic-fin origins, longer than central abdominal plates.

Preanal plate bordered anteriorly by three, somewhat

developed plates of similar size, all in contact with preanal

plate. Central abdominal plates not organized; anterior

abdominal plates smaller and more numerous than posterior

Fig. 13. Sturisomatichthys leightoni, IUQ 2739, 87.9 mm SL, Colombia, Valle del Cauca department, San Antonio de Piedras municipality, Upper

Cauca River, Portugal de Piedras River. Photo by Aura Mar´

ıa Bastidas Quintero (IUQ).

786 Copeia 107, No. 4, 2019

ones. Central abdominal plates numerous and irregular,

smaller than anterior and posterior ones.

Posterior margin of dorsal fin straight, unbranched and

first and second branched rays longest; dorsal-fin rays i,7.

Pectoral-fin rays i,6, unbranched ray longest, barely reaching

origin of pelvic fin. Pelvic-fin rays i,5, unbranched ray

longest, its tip reaching origin of anal fin. Anal-fin rays i,5,

unbranched ray longest. Posterior caudal-fin margin deeply

concave, short filaments of less than half SL on both upper

and lower ray.

Coloration in alcohol.—Ground color of dorsum dark brown;

head and ventral surface light brown. Dorsolateral stripe on

snout (Fig. 13), not as visible posterior to orbits, yet present

and discontinuous. Dark, narrow stripe at beginning of dorsal

fin (Fig. 13) as continuation of dorsolateral stripe present on

snout. Dorsolateral stripe somewhat visible along caudal

peduncle, but not as visible as on other congeners. All fins

with dark and visible spots, no stripes (Fig. 13). Dorsal fin

with short band on distal tip of first two branched rays (Fig.

13).

Sexual dimorphism.—Males with small groups of thin and

short hypertrophied odontodes on the sides of the head, and

along the plates of the caudal peduncle (Fig. 13).

Distribution.—Sturisomatichthys leightoni occurs in the Mag-

dalena-Cauca and Orinoco basins in Colombia (Fig. 3).

Remarks.—Regan (1912) described Oxyloricaria leightoni from

Honda, a city in the upper portion of the Magdalena River,

Colombia; the author gave no specific location of the

collection of the types. In the description, the author cited

two juvenile specimens (55 mm and 65 mm TL) that were

examined here. Despite their juvenile state, the specimens

show the characteristics present in adults regarding colora-

tion of body and fins, the only difference being the presence

of short odontodes on the sides of the head in sexually

mature males (Fig. 13). Even though the specimens were

collected in 1909 (Regan, 1912), they are fairly well

preserved, and the characteristics cited in the original

description were identified.

Even though Eigenmann (1910) proposed Oxyloricaria as a

junior synonym of Sturisoma, Regan (1912) originally

described the species as Oxyloricaria ignoring the taxonomic

arrangement of Eigenmann. Nevertheless, Eigenmann

(1920a) referred to the species as Sturisoma leightoni and

mentioned its presence in the Magdalena-Cauca basin on

both sides of the eastern cordillera, taking Bogota

´as a

reference. In that same year, Eigenmann (1920b), in a study

of the fishes of the rivers of the western cordillera in

Colombia (San Juan, Atrato, Pat´

ıa, and Dagua rivers),

Table 4. Morphometrics and meristics of Sturisomatichthys leightoni and S. panamensis. Morphometric values as percents of standard length (SL)

and head length (HL). SD ¼standard deviation.

S.leightoni (n¼63) S.panamensis (n¼56)

Min Max Mean SD Min Max Mean SD

Standard length 47.4 128 98.8 102.3 251 191

Head length 20.1 24.2 22.2 0.9 19.0 21.6 20.3 0.7

Predorsal length 30.6 36.9 33.6 1.3 29.1 32.3 30.9 0.7

Abdominal length 12.1 15.0 13.5 0.8 9.3 12.5 11.4 0.7

Trunk length 13.3 18.4 15.8 1.1 12.4 16.2 14.3 0.9

Dorsal fin length 20.7 27.3 24.4 1.4 26.5 35.0 30.0 2.4

Pectoral fin length 17.0 22.9 20.2 1.3 19.3 26.1 21.3 1.4

Pelvic fin length 15.0 19.6 17.0 1.1 14.9 20.1 16.5 1.0

Anal fin length 15.9 20.2 18.6 1.0 18.0 24.7 22.1 1.6

Head depth at supraoccipital tip 7.0 12.2 10.3 1.0 6.8 9.8 8.6 0.6

Body depth at dorsal-fin origin 9.3 13.9 11.6 1.1 8.0 12.1 10.8 0.8

Caudal peduncle depth 1.3 2.1 1.5 0.2 1.0 1.8 1.3 0.1

Cleithral width 11.0 14.1 12.6 0.7 8.4 10.5 9.5 0.6

Body width at dorsal-fin origin 11.0 16.1 13.0 1.3 9.4 13.0 11.2 1.0

Body width at anal-fin origin 9.3 12.4 11.0 0.7 8.4 11.2 9.8 0.7

Body width at caudal-fin origin 1.6 3.0 2.4 0.3 1.5 2.7 2.2 0.3

Head width 58.4 75.0 68.2 3.2 50.0 63.1 56.8 3.5

Interorbital distance 25.4 32.6 29.9 1.4 25.6 34.1 28.7 1.9

Orbital diameter 10.9 16.6 13.3 1.2 11.1 15.3 12.5 1.1

Snout length 53.5 61.8 58.3 2.0 60.4 66.9 62.7 1.6

Dentary length 7.9 16.4 11.6 1.6 7.5 13.7 10.6 1.2

Lower lip width 26.2 37.5 33.8 2.6 21.6 30.5 27.9 1.7

Plates on median series 13 17 15 13 17 15

Coalesced plates 13 16 14 15 19 17

Ventrolateral thoracic plates 5 8 7 7 9 8

Premaxillary teeth 24 50 30 37 53 42

Dentary teeth 22 44 29 32 44 37

Dorsal-fin rays i,7 i,7 i,7 i,7 i,7 i,7

Pectoral-fin rays i,6 i,6 i,6 i,6 i,6 i,6

Pelvic-fin rays i,5 i,5 i,5 i,5 i,5 i,5

Anal-fin rays i,5 i,5 i,5 i,5 i,5 i,5

Caudal-fin rays i,12,i i,12,i i,12,i i,12,i i,12,i i,12,i

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 787

recognized the species as occurring in the San Juan River, and

as doubtfully occurring in the Atrato River, both records

being rejected here. Shortly after that, Eigenmann (1922)

offered records of the species from the upper Cauca and

Magdalena basins, San Juan River in the Colombian Pacific

slope, and Meta River in Cumaral, Colombia. The author

made a remark regarding those last specimens, stating that

‘‘The specimen from Cumaral in Eastern Colombia differs

slightly in having the fins a little shorter and the spines of

the males shorter’’ (Eigenmann, 1922: 96). The specimens

included by Eigenmann (1922; CAS 77279, three specimens,

and CAS 77280, two specimens) were examined here.

Nic´

eforo Maria collected both lots in February 1917, in

Cumaral, Meta department, Colombia. This locality is just on

the eastern border of the eastern Cordillera of the Andes, and

thus, draining to the cis-Andean Orinoco basin. The five

specimens analyzed have all characters that identify S.

leightoni, and the character discussed by Eigenmann (1922)

regarding ‘‘fins a little shorter and the spines of the males

shorter’’ was not found to be adequate for the separation of S.

leightoni as a distinct species from populations from Cumaral.

This locality belongs to the Orinoco basin, on one of its

western most points, which constitutes a new record for S.

leightoni in that basin, and a cis-Andean locality for the

species, being the only species of Sturisomatichthys present in

that basin (Fig. 3). Armbruster (2005), in a study of

Lasiancistrus, cited lots belonging to L. guacharote from

around Villavicencio. According to the author, Villavicencio

(same locality of lots belonging to S. leightoni collected by

Nic´

eforo Maria), is a town well connected with the aquarium

export business, and those specimens could be the result of

being introduced from the aquarium trade; this also could be

the case for the specimens examined here belonging to S.

leightoni. Nevertheless, since we do not have evidence of

introduction of these specimens, or a mistake in the labeling

of the localities by N. Maria, we consider S. leightoni to be

present at Cumaral (CAS 77279 and CAS 77280) due to the

characters observed in the specimens which agree with the

description and diagnosis of the species presented here.

As it was discussed above, according to Covain et al. (2016)

Sturisomatichthys is a strictly trans-Andean genus within the

Loricariinae. Nevertheless, through this record, and the fact

that Sturisomatichthys caquetae is a valid species (see above)

distributed in the upper Amazon Basin, the genus is

recognized as both cis- and trans-Andean, making Dasylor-

icaria the only strictly trans-Andean genus in the subfamily.

Subsequent studies to that of Eigenmann (1922) listed the

species as occurring, in addition to the Magdalena-Cauca

basin, in the Pacific and Eastern drainages in Colombia (i.e.,

Fowler, 1942; Miles, 1947; Dahl, 1971; Z´

u˜

niga et al., 2004;

Maldonado-Ocampo et al., 2005, 2008; Ortega-Lara et al.,

2006a; DoNascimiento et al., 2018), a fact that was not

corroborated in the present study. Here, it was found that S.

leightoni occurs in the Magdalena-Cauca and Orinoco basins

and not in western drainages (Fig. 3).

Isbr¨

ucker and Nijssen (in Isbr¨

ucker, 1979) described the

genus Sturisomatichthys assigning S. leightoni as the type

species. In that same study, a lectotype was designated for the

species (BMNH 1909.7.23:45). The type series was examined

in this study (see Material Examined), and the juvenile state

of the three individuals was verified. One of the paralecto-

types was less than 30 mm SL and it was not included in

Table 4, but the characters were identified, as in the lectotype

and the other paralectotype.

Material examined.—ANSP 84178, 1 alc, Colombia, Tolima

department, Honda municipality, Upper Magdalena River,

58120N, 748440W; BMNH 1909.7.23.45, 57.1 mm SL, lecto-

type, Colombia, Honda municipality, Magdalena River, alt.

300–400 feet, 5812025 00 N, 7484401200W; BMNH 1909.7.23.46-

47, 2 alc, 29.0–47.4 mm SL, paralectotypes, same data as

lectotype; CAS 77232, 7 alc, Colombia, Cundinamarca

department, Girardot municipality, Magdalena River basin,

48180N, 748480W; CAS 77279, 3 alc, Colombia, Meta

department, Cumaral municipality, Orinoco River basin,

Carneceria cannel, two days from Villavicencio city, 4880N,

738370W; CAS 77280, 2 alc, Colombia, Meta department,

Cumaral municipality, Orinoco River basin, Los Llanos,

northeast of Villavicencio city, 48160N, 738290W; CIUA 232,

1 alc, Colombia, Caldas and Antioquia departments, Magda-

lena River basin, La Miel River, 58460N, 748410W; CIUA 401, 2

alc, Colombia, Antioquia department, Bolombolo and Ven-

ecia municipalities, lower Cauca River basin, 58560N,

758460W; CIUA 402, 8 alc, Colombia, Valle del Cauca

department, Cartago City, upper Cauca basin, La Vieja River,

48440N, 758540W; CIUA 921, 5 alc, Colombia, Caldas,

Victoria, Magdalena basin, Guarin ´

o River, Quebrada Casan-

guillas, 58170N, 748440W; CIUA 987, 5 alc, 103.1–106.8 mm

SL, Colombia, Caldas department, Victoria municipality,

Magdalena River basin, Guarin ´

o River, Bocatoma Creek,

58170N, 748440W; CIUA 1059, 3 alc, Colombia, Caldas

department, Victoria municipality, Magdalena River basin,

Guarin´

o River, Bocatoma Creek, 58170N, 748440W; CIUA

1920, 1 alc, Colombia, Santander department, Betulia

municipality, Magdalena River basin, Sogamoso River, at La

Paz Bridge, 78120N, 738550W; CIUA 3162, 1 alc, Colombia,

Antioquia department, lower Cauca River, Middle Tenche

River, 7820N, 75850W; CIUA 3608, 4 alc, Colombia, Caldas,

Chinchina

´municipality, Cauca River, before discharge of San

Francisco dam, 58304300 N, 75843 05100 W, 851 m asl; CIUA 3818,

1 alc, Colombia, Caldas department, Chinchina

´municipal-

ity, Cauca River basin, before discharge of San Francisco dam,

5830600 N, 75844029 00 W, 861 m asl; CIUA 3826, 1 alc, Colombia,

Caldas department, Chinchina

´municipality, Cauca River,

before discharge of San Francisco dam, 5830300 N, 75844037 00 W,

855 m asl; CIUA 3858, 2 alc, Colombia, Caldas department,

Supia municipality, Supia River, near mouth at Cauca River,

5822041 00 N, 7583605500W, 748 m asl; CIUA 3917, 2 alc,

Colombia, Caldas department, Belalcazar municipality, Mag-

dalena River basin, Risaralda River, El Zancudo path,

48502700 N, 7585102100 W, 932 m asl; CIUA 3920, 1 alc,

Colombia, Caldas department, Anserma municipality, El

Tablazo path, Risaralda River, 5890600 N, 75849054 00 W, 1031 m

asl; CIUA 3992, 10 alc, Colombia, Caldas department,

Viterbo municipality, Magdalena River basin, Guarne River

tributary of Risaralda River, 58502900 N, 758520400W, 986 m asl;

CIUA 4800, 1 alc, Colombia, Antioquia department, Pue-

blorrico municipality, Cauca River basin, San Juan River, PCH

La Cristalina project, at discharge of dam, 5824025 00 N,

75853011 00 W; CIUA 4839, 7 alc, Colombia, Caldas depart-

ment, La Dorada municipality, Magdalena River basin, Do˜

Juana River, 58300300 N, 74841014 00 W; CIUA 4899, 2 of 6 alc,

88.0–93.7 mm SL, Colombia, Caldas, Magdalena River basin,

Purnio River, 582005700N, 7485004100 W; FMNH 55131, 30 alc,

Colombia, Tolima department, Honda municipality, upper

Magdalena, Bernal Creek, 58120N, 748440W; FMNH 55132, 6

alc, Colombia, Girardot municipality, 481801300 N,

74848014 00 W; FMNH 55134, 4 alc, Colombia, Gabi (?); FMNH

55136, 6 alc, 1 CS, Colombia, Paila municipality, 48230N,

76840W; ICNMHN 5290, 5 alc, Colombia, Boyaca

´, Puerto

788 Copeia 107, No. 4, 2019

Romero, middle Magdalena River, La Fiebre Creek, serran´

ıa de

los Quinchas, 58510N, 748190W; ICNMHN 6497, 48 alc,

Colombia, Boyaca

´, middle Magdalena River, Puerto Boyaca

´,

Puerto Romero, Las Quinchas, La Fiebre Creek, 58510N,

748190W; ICNMHN 15672, 1 alc, Colombia, Caldas, Norcasia,

Magdalena River basin, La Miel River, upstream from mouth

of Manso River, 48100N, 758160W; ICNMHN 16160, 3 alc,

Colombia, Caldas, Norcasia, Magdalena River basin, La Miel

River, downstream from mouth of Manso River, 48100N,

758160W; ICNMHN 16175, 4 alc, Colombia, Antioquia, San

Miguel, Magdalena River basin, La Miel River, 58460N,

748410W; ICNMHN 17733, 1 alc, Colombia, Santander, El

Carmen de Chucuri, El Top´

on farm, middle Magdalena,

Top ´

on River, main channel of river, 684504500 N, 7383405600 W,

250 m asl; IUQ 51, 1 alc, 102.1 mm SL, Colombia, Valle del

Cauca department, Cauca-Magdalena basin, upper Cauca

River, San Pablo Creek, tributary of La Paila River, 48210N,

76830W; IUQ 54, 1 alc, 102.7 mm SL, Colombia, Quind´

ıo

department, Alambrada district, upper Cauca River, Los

Kingos River tributary of La Vieja River, 48240600 N,

75852013 00 W, 1205 m asl; IUQ 58, 2 alc, 85.4 mm SL,

Colombia, Quind´

ıo department, Quimbaya municipality, La

Espa˜

nola district, Playa Azul farm, Upper Cauca River, Roble

River, 48340N, 758520W; IUQ 61, 2 alc, 107.3–109.3 mm SL,

Valle del Cauca department, upper Cauca River, San Pablo

Creek tributary of La Paila River, 48210N, 76830W; IUQ 100, 3

alc, Colombia, Quind´

ıo department, Quimbaya municipality,

Puerto Alejandr´

ıa district, upper Cauca River, La Vieja River,

48480N, 758560W; IUQ 146, 2 of 6 alc, 90.2–105.1 mm SL,

Colombia, Quind´

ıo department, Maravelez district, upper

Cauca River, La Vieja River, 48480N, 758560W; IUQ 157, 2 alc,

101.3–108.5 mm SL, Colombia, Valle del Cauca department,

road via Sevilla municipality, La India farm, La Paila River

tributary of Cauca River, 48190N, 76880W; IUQ 158, 2 alc,

108.5–118.5 mm SL, Colombia, Valle del Cauca department,

road via Sevilla municipality, La India farm, La Paila River

tributary of Cauca River, 48190N, 76880W; IUQ 216, 8 alc,

74.9–86.0 mm SL, Colombia, Quind´

ıo department, Puerto

Alejandr´

ıa district, upper Cauca River, La Vieja River, 48480N,

758560W; IUQ 286, 7 alc, 93.1–110.0 mm SL, Colombia, Valle

del Cauca department, road to Sevilla municipality, La India

farm, upper Cauca River, Totoro Creek tributary of Cauca

River, 28300N, 768240W; IUQ 672, 5 alc, 95.2–115.2 mm SL,

Colombia, Quind´

ıo department, La Tebaida municipality, La

Argentina district, upper Cauca River, La Vieja River, 48270N,

758470W; IUQ 829, 1 of 3 alc, 104.1 mm SL, Colombia,

Quind´

ıo department, Maravelez district, upper Cauca River,

48250100 N, 7584804100 W, 1074 m asl; IUQ 1555, 1 alc, 79.5 mm

SL, Colombia, Quind´

ıo department, Upper Cauca River, La

Vieja River, 48480N, 758560W; IUQ 1674, 4 alc, 74.2–92.7 mm

SL, Colombia, Valle del Cauca department, road via Zarzal

municipality, upper Cauca River, Las Ca ˜

nas creek tributary of

Cauca River, 48210N, 76840W; IUQ 1776, 1 alc, 128.0 mm SL,

Colombia, Quind´

ıo department, Quimbaya municipality,

Ocaso natural reserve, upper Cauca River, Roble River at

Playa Azul sector, 483506800N, 7585208100 W; IUQ 1812, 5 alc,

69.6–83.9 mm SL, Colombia, Quind´

ıo department, Quim-

baya municipality, Ocaso natural reserve, upper Cauca River,

Roble River, 100 mts of Playa Azul sector, 483506800N,

75852081 00 W; IUQ 1857, 8 alc, 96.6–115.1 mm SL, Colombia,

Quind´

ıo department, Quimbaya municipality, Ocaso natural

reserve, upper Cauca River, Roble River, near Playa Azul

sector, 483506800N, 7285208100 W; IUQ 2739, 1 alc, 87.9 mm SL,

Colombia, Valle del Cauca department, San Antonio de

Piedras municipality, upper Cauca River, Portugal de Piedras

River, 48304500 N, 76818045 00 W, 944 m asl; IUQ 2740, 1 alc, 68.6

mm SL, Colombia, Valle del Cauca department, San Antonio

de Piedras municipality, upper Cauca River, Portugal de

Piedras River, 48304500 N, 7681804600 W, 945 m asl; IUQ 3087, 1

alc, 118.7 mm SL, Colombia, Quind´

ıo department, upper

CaucaRiver,BoquiacreerktributaryofQuind

ıo River,

48380N, 758350W; IUQ 3089, 4 of 5 alc, 96.4–103.3 mm SL,

Colombia, Quind´

ıo department, La Tebaida municipality,

upper Cauca River, Espejo River, 48270N, 758570W; IUQ 3091,

1 of 2 alc, 103.4 mm SL, Colombia, Quind´

ıo department,

upper Cauca River, Boquia creerk tributary of Quind´

ıo River,

48380N, 758350W; MCP 41758, 1 alc, Colombia, Cauca

department, Buenos Aires municipality, Cauca River, Zanj ´

Bagazal River at Lomitas, 3800N, 768380W; MPUJ 7862, 2 alc,

Colombia, Cundinamarca, Villeta, Villeta River, Dulce River

upstream, 48590200 N, 74854052.200 W, 850 m asl; MPUJ 7865, 2

alc, Colombia, Cundinamarca, Villeta, Villeta River, road

Bogota

´-Villeta, 58400.800 N, 74827 01.600 W, 687 m asl; MPUJ

7868, 19 alc, Colombia, Cundinamarca, Villeta, Villeta River,

between Dulce River and Namay Creek, 4859031.500N,

74829011.0 00 W, 1200 m asl; USNM 121256, 1 alc, 105.6 mm

SL, Colombia, Tolima department, upper Magdalena River,

Luisa River, 4810N, 748520W; ZMA 139.631, 1 alc, 100.1 mm

SL, Colombia, Honda, upper Magdalena, 58120N, 74844 0W.

Sturisomatichthys panamensis (Eigenmann and Eigen-

mann, 1889)

Figure 14, Table 4

Loricaria panamensis Eigenmann and Eigenmann, 1889: 34–

35 (type locality: Panama; holotype MCZ 8126; length .95

m [?] SL). Eigenmann and Eigenmann, 1890: 365–366

(reported as closely related to Sturisoma rostratum; repro-

duction of the original description).

Oxyloricaria panamensis (Eigenmann and Eigenmann,

1889).—Regan, 1904: 301–302 (Sturisomatichthys aureus as

junior synonym; distributed in the Magdalena River and

Panama; holotype reported to be 98 mm TL). Meek and

Hildebrand, 1916: 229 (distributed at the Bayano and

Tuyra rivers, Panama, Atlantic and Pacific slopes in

Colombia, and southern Ecuador; key to Oxyloricaria

(¼Sturisoma); description of the species; Sturisomatichthys

aureus,S. dariensis, and S. frenatus as junior synonyms of

the species).

Sturisoma panamense (Eigenmann and Eigenmann, 1889).—

Eigenmann, 1910: 416 (identification key for Sturisoma;

valid as Sturisoma panamense; reference to Regan, 1904).

Regan, 1913: 470 (species as valid in Sturisoma; without

description). Eigenmann, 1920b: 11 (species distributed at

the Atrato, Dagua, San Juan, and lower Pat´

ıa rivers in

Colombia); 1920c: 14 (distributed at Chepo, Tuyra, and

Atrato rivers); 1922: 94-95 (synonymies; same distribution

as in Eigenmann, 1920b, 1920c). Miles, 1947: 114

(identification key for Sturisoma; synonymies including S.

dariensis and S. frenatus; distributed at lower Magdalena,

San Juan and Patia, Colombia, Bayano and Tuyra, Panama,

and northwest Ecuador). Dahl, 1955: 17 (distributed at the

Sin ´

u River, Colombia); 1971: 94 (as present at lower,

middle, and upper Magdalena, upper and middle Sin ´

River; maximum size of 20 cm TL; identification key of

Sturisoma along with S. aureus and S. leightoni). Isbr¨

ucker,

1980: 95 (included in Sturisoma as a member of the

Harttiina; no diagnosis); 1981a: 88 (as valid in Sturisoma);

Roma

´n-Valencia, 1990: 205 (present at middle Atrato

River). Isbr¨

ucker et al., 2001: 32 (included in the catalog

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 789

of the Loricariinae as valid in Sturisoma, German version).

Isbr¨

ucker, 2002: 29 (included in the catalog of the

Loricariinae as valid in Sturisoma, English version). Ferraris,

2003: 347 (genus distribution in Pacific slope rivers of

Panama, Colombia, and Ecuador, Caribbean slope rivers of

Colombia; synonymies; type material location at MCZ

8126; species with 26 cm SL maximum; common names in

Colombia and Ecuador). Ghazzi, 2003: 182 (belonging to a

new genus); 2005: 564 (holotype used for comparison with

S. kneri). Maldonado-Ocampo et al., 2005: 29, figs. 123 and

134 (synonyms; common names in Colombia; reproduc-

tion of the original description, in Spanish; records of

biology and ecology of the species; distribution and

distribution map; specimens in Colombian museums);

2006: 150 (distributed at Atrato River in Colombia,

specimens at Colombian museums). Mojica et al., 2006a

(distributed at Magdalena-Cauca Basin; catalog numbers in

Colombian museums). Ortega-Lara et al., 2006a: 182 (as

distributed at the Patia River, Colombia; specimens in

Colombian museums). Ferraris, 2007: 299 (type locality;

distribution of the species). Maldonado-Ocampo et al.,

2008: 195 (distributed at the Magdalena-Cauca basin, and

Caribbean and Pacific slopes in Colombia; specimens in

Colombian Museums); 2013: 396 (present at Atrato and

Le´

on river basins in the Caribbean slope, and Baud´

o, San

Juan, Anchicaya

´, Dagua and Patia River basin in the Pacific

slope, in Colombia; specimens in Colombian museums).

Sturisoma panamensis (Eigenmann and Eigenmann, 1889).—

Fowler, 1942: 132 (distributed at the Pat´

ıa, Mag¨

u´

ı, Telemb´

ı,

San Juan, Condoto, Atrato, Certegu´

ı, and Magdalena rivers,

following Regan [1913]).

Sturisomatichthys panamensis (Eigenmann and Eigenmann,

1889).—Covain et al., 2016: 514 (valid as Sturisomatichthys

panamense; phylogenetic relationships of the Loricariinae

subfamily using two mitochondrial and one nuclear

markers; appears as sister group to all Sturisomatichthys).

DoNascimiento et al., 2018: 77 (as present in Colombia,

Magdalena-Cauca River basin, and Pacific and Caribbean

slopes).

Diagnosis.—Sturisomatichthys panamensis is distinguished

from S. caquetae,S. citurensis,S. guaitipan, new species, and

S. tamanae by the possession of a rostrum (vs. rostrum absent)

and a dorsolateral stripe from the tip of snout to the caudal

peduncle (vs. dorsolateral stripe absent), and, from S.

caquetae,S. citurensis, and S. tamanae, by the number of

dentary teeth (32–44 vs. 30, 63–100, and 60–93, respective-

ly). From S. dariensis,S. leightoni, and S. varii, new species, it is

distinguished by lacking reticulations on the supraoccipital,

interorbital, and predorsal region (vs. reticulations present).

Furthermore, it is distinguished from S. aureus by having an

inconspicuous blotch on the pelvic and anal fins (vs. dark

blotch present on both fins) and lacking a transverse bar at

the origin of the dorsal fin united to the dorsolateral stripe on

Fig. 14. Sturisomatichthys panamensis, USNM 293412, 207.4 mm SL, Panama, Darien Province, Tuira River between Calle Larga and Pinogana,

above El Real, Pacific slope.

790 Copeia 107, No. 4, 2019

both sides of the body (vs. transverse bar present). From S.

reinae, new species, it is distinguished by having fewer and

smaller central abdominal plates (vs. more numerous and

larger central abdominal plates; Fig. 14). From S. dariensis, it is

distinguished by having a large, dark blotch on the pectoral

fin (vs. blotch absent). From S. festivus, it is distinguished by

lacking filaments on the dorsal, pectoral, pelvic, and anal fins

(vs. filaments present). From S. frenatus, it can be differenti-

ated by lacking dark blotches on the pelvic and anal fins (vs.

dark blotches present on all fins, occupying the entire fin)

and a stripe on first two rays of the dorsal fin not reaching

the distal tip of rays (vs. stripe present and reaching distal tip

of first two dorsal-fin rays). From S. leightoni, it is distin-

guished by having a dark blotch on the pectoral fin and a

stripe along the first two rays of the dorsal fin (vs. blotch and

stripe absent) and a continuous dorsolateral stripe (vs.

discontinuous dorsolateral stripe).

Description.—Morphometric data in Table 4. Body elongate,

completely covered by dermal plates with small, short

hypertrophied odontodes, perceptible on sides of head.

Dorsal profile of head slightly concave from tip of snout to

interorbital region; straight from that point to dorsal-fin

origin; slightly concave from dorsal-fin origin to end of

caudal peduncle. Ventral body profile flat. Head and

predorsal region broader than remainder of body; postdorsal

region narrower towards caudal fin.

Head triangular, pointed; rostrum present, broad on its

entire length. Ventral border of rostrum flat. Orbital rim

circular, without notch, dorsolaterally placed on head.

Sphenotic with short lateral expansion. Canal-bearing plate

well developed, in contact with anterior abdominal plates,

anterior to branchiostegal membrane reaching distal tip of

rictal barbel. Upper lip covered with small, almost impercep-

tible plates; lower lip without such plates. Lower lip covered

with round, small, almost imperceptible papillae. Rictal

barbel short, free from lip rim, not reaching to anterior

border of abdominal plates, nor surpassing posterior border

of lower lip. Teeth slender, bifid, 37–53 on premaxilla, 32–44

on dentary; cusps slightly curved. Teeth in lower jaw larger

than those in upper jaw.

Predorsal plates with two low, inconspicuous ridges;

nuchal plate twice as broad as predorsal plates. Median

plates carinate, forming two longitudinal sharp keels; 15–19

coalescent plates, beginning at 13–17 median plates. Seven to

nine straight ventral lateral plates, between pectoral- and

pelvic-fin origins, larger than central abdominal plates.

Preanal plate bordered anteriorly by three well-developed

and similarly sized plates; anteriormost plate sometimes not

in contact with preanal plate. Central abdominal plates

irregular, not as small as in other congeners (Fig. 14). Anterior

abdominal plates smaller and more numerous than posterior

ones.

Posterior margin of dorsal fin straight, unbranched and

first branched ray longest; dorsal-fin rays i,7. Pectoral-fin rays

i,6, unbranched ray longest, slightly surpassing anterior third

of pelvic fin. Pelvic-fin rays i,5, unbranched ray longest, its

tip slightly passing anterior third of anal fin. Anal-fin rays i,5,

unbranched ray longest. Posterior caudal-fin margin deeply

concave, long filaments, almost reaching half SL on some

specimens, on both upper and lower caudal rays. Un-

branched ray of pectoral fin elongated, not as long as

filaments on caudal fin of some species of Sturisomatichthys,

but which can significantly modify total extension of fin.

Coloration in alcohol.—Ground color of dorsum, head, and

ventral surface pale brown to yellow. Dorsolateral stripe on

both sides of body. Stripe broader from anterior border of first

predorsal plate to dorsal-fin origin (Fig. 14), becoming less

conspicuous and thin towards caudal peduncle. Remnant of

dark stripe sometimes on ventral side of rostrum, separated as

on both sides of body. Large, dark spots on membrane and

rays of pectoral fin. Dorsal fin with dark brown stripe along

first two branched rays, with conspicuous dark spots along

unbranched ray, not reaching its distal tips. Some specimens

with one to several white ocelli just anterior to middle of fin

(Fig. 14). Caudal fin with dark stripe along first two or three

branched rays, in both upper and lower lobes. Dark brown

spots on edges of caudal lobes.

Sexual dimorphism.—The specimens examined showed a

subtle sexual dimorphism consisting of short hypertrophied

odontodes on the sides of the head of males (Fig. 14).

Distribution.—Sturisomatichthys panamensis occurs in the

Aruza, Capeti, and Tuyra rivers in the Pacific versant,

Panama, and the Atrato, Sin ´

u, and Truando rivers in the

Caribbean versant, Colombia (Fig. 1).

Remarks.—Sturisomatichthys panamensis has been proposed as

senior synonym of several species of Sturisomatichthys.

Described by Eigenmann and Eigenmann (1889), the authors

only included ‘‘Panama’’ as the type locality of the species.

Their description is complete and included proportions,

general aspects of coloration, and plates and teeth counts.

Nevertheless, the holotype, the only type specimen, is a

juvenile of 95 mm SL (Eigenmann and Eigenmann, 1889),

and this could have been one of the reasons for it to be

assumed as a synonym to several others. Eigenmann and

Eigenmann (1890) pointed out that the species could be

related to Loricaria rostrata (¼Sturisoma rostratum).

Regan (1904) transferred the species to Oxyloricaria and

included St. rostratum and S. aureus as junior synonyms of S.

panamensis. The author discussed the fact that after examin-

ing the type of S. panamensis, there remained no doubt about

its conspecificity with S. aureus and thus the species was

distributed in the Magdalena River as well; the author did not

elaborate regarding the synonymy with St. rostratum.Ina

study of the fishes from Panama, Meek and Hildebrand

(1916) included the species, still as O. panamensis, ignoring

the synonymy of Oxyloricaria with Sturisoma proposed by

Eigenmann (1910). The authors included St. rostratum,S.

aureus,S. dariensis, and S. frenatus as junior synonyms of S.

panamensis, without any justification for this action, and

extended the distribution of the species to the Pacific slope of

Panama, both slopes of Colombia, and south to Ecuador.

Eigenmann (1920b, 1920c) included the species, as

Sturisoma panamense, as distributed in the Chepo and Tuyra

rivers belonging to the Pacific slope of Panama, the Atrato

River, Caribbean slope of Colombia, and the Dagua, San

Juan, and Pat´

ıa rivers, Pacific versant of Colombia. A little

after that, Eigenmann (1922) followed the same synonymies

proposed by Meek and Hildebrand (1916) for S. panamensis,

with the exception of S. aureus, which was assumed as valid

by the author, and did not include the Magdalena River as

part of the distribution of S. panamensis. In the same way,

Miles (1947) followed the taxonomic actions proposed by

Eigenmann. Fowler (1942) also cited the species as present

along the Pacific slope of Colombia in the Pat´

ıa, Mag¨

u´

ı,

Telemb´

ı, San Juan, and Condoto rivers, and the Caribbean

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 791

slope in the Atrato, Certegu´

ı, and Magdalena rivers, in

Colombia. Dahl (1955) was the first author to report the

species as distributed in the Sin ´

u River, on the Caribbean

slope of Colombia, and referred to the similarity of the

species with S. tamanae, differing, according to the author,

only on the counts of lateral plates, but assumed both species

as valid. Roma

´n-Valencia (1990) and Maldonado-Ocampo et

al. (2006) also included the Atrato River in the distribution of

the species. Ortega-Lara et al. (2006a) listed the species as

present in the Pat´

ıa River, but did not comment on the

synonymies of the species. On the other hand, and in

addition to records of distribution of S. panamensis in

localities of northwestern Colombia, Maldonado-Ocampo

et al. (2005, 2008) and Mojica et al. (2006a) reported the

species, using records of Colombian museums, in the upper

Cauca and Magdalena rivers. Maldonado-Ocampo et al.

(2013) included S. panamensis as present in the San Juan

River, Colombian Pacific, which is rejected in this study. The

specimens cited by the authors were not examined here (see

Remarks on S. tamanae and S. varii, new species), but it is

suggested that they do not belong to S. panamensis.

It was found here that S. panamensis is distributed in the

Aruza, Capeti, and Tuyra rivers in the Pacific versant,

Panama, and the Atrato, Sin ´

u, and Truando rivers in the

Caribbean versant, Colombia. This agrees only partially with

the distribution described by previous authors, rejecting the

presence of the species in the Magdalena-Cauca basin,

Mag¨

u´

ı, Telemb´

ı, Certegu´

ı, San Juan, Pat´

ıa, and Dagua rivers.

The synonymies proposed by Eigenmann and Eigenmann

(1889, 1890), Regan (1904), Meek and Hildebrand (1916),

Eigenmann (1922), Miles (1947), and Dahl (1971) are

rejected here.

Externally, S. panamensis is similar to S. frenatus, especially

regarding the coloration of the dorsal and pectoral fins (Figs.

11, 12, 14). Sturisomatichthys panamensis has a dark stripe on

the first branched dorsal-fin rays, but these do not extend

beyond those rays nor reach the distal tip of the rays,

contrary to S. frenatus. The blotches on the pectoral fin are

less conspicuous in S. panamensis, and pelvic and anal fins do

not present visible blotches, contrary to S. frenatus (see also

Diagnosis and identification key). Sturisomatichthys aureus

and S. leightoni are distributed in the Magdalena River, while

S. panamensis occurs in the Atrato and Tuyra rivers (among

others). As discussed by Schaefer (2011), the uplifting of the

central and western ranges of the Andes isolated the

Magdalena-Cauca basin from the western basins such as

the Atrato, San Juan, Pat´

ıa, Baud´

o (Colombia), and Tuyra

(Panama) basins (see also Londo˜

no-Burbano and Reis, 2016).

This shows a close relationship among populations distrib-

uted in the western drainages, on both the Pacific and

Caribbean slope, and the Magdalena-Cauca basin, which is

the case for other groups of the Loricariinae (see Discussion).

Material examined.—CIUA 4324, 2 alc, Colombia, Antioquia

department, Murind ´

o municipality, Atrato River basin,

Campo Alegre district, 6851034 00 N, 7684906900W; CIUA 4355,

1 alc, 183.1 mm SL, Colombia, Antioquia department,

Murind´

o municipality, Tad´

ıa district, Atrato River basin,

6851034 00 N, 768490900 W; CIUA 4357, 2 alc, Colombia, Anti-

oquia department, Vig´

ıa del Fuerte municipality, Buchad´

district, Guaguand´

o swamp, 6824013 00 N, 768450600 W; CIUA

4501, 3 of 5 alc, 142.7–203.2 mm SL, Colombia, Antioquia

department, Murind ´

o municipality, Atrato River basin,

Campo Alegre district, 6851034 00 N, 768490900 W; CIUA 4627, 2

alc, Colombia, Antioquia department, Vig´

ıa del Fuerte

municipality, Buchad´

o district, Atrato River basin,

6825032 00N, 76845 04800 W; CIUA 4784, 4 of 6 alc, 158.5–188.2

mm SL, Colombia, Antioquia department, Atrato River, Vigia

del Fuerte municipality, Buchado district, 68250N, 768450W;

NRM 36778, 1 alc, 142.2 mm SL, Colombia, C ´

ordoba

department, Sin ´

u River, Tierralta municipality, unnamed

Creek emptying about 300 mts downstream Urra

´Hydropow-

er Station, 78560N, 768170W; USNM 78370, 4 of 11 alc, 186.8–

209.4 mm SL, Panama, Darien Province, Capeti River, 8830N,

778340W; USNM 78371, 7 alc, 196.7–251.0 mm SL, Panama,

Darien Province, Aruza River, Aruza sector at Darien

Province, 8870N, 778350W; USNM 78372, 12 of 16 alc,

164.4–232.2 mm SL, Panama, Darien Province, Tuyra River

at Boca De Cupe, 88300000 N, 77835 00000 W; USNM 79145, 3 of 5

alc, 185.2–204.1 mm SL, Panama, Darien Province, Tuyra

River at Marrigante, 88190N, 788180W; USNM 293412, 10 of

14 alc, 1 CS, 186.4–229.0 mm SL, Panama, Darien Province,

Tuira River between Calle Larga and Pinogana, above El Real,

Pacific slope, 8830N, 778430W; USNM 316293, 9 alc, 102.3–

204.6 mm SL, Colombia, Choc ´

o department, Truando River,

Creek of Upper Nercua River, Large tributary of Truando

River, 58490N, 768370W; USNM 316295, 4 alc, 149.9–206.1

mm SL, Colombia, Choc ´

o deparment, Prado district, Creek of

Parado River, about 10 min helicopter flight from Village of

Parado; USNM 341993, 2 of 3 alc, 163.8–184.2 mm SL,

Colombia, Choc´

o department, Salado River near Teresita.

Sturisomatichthys tamanae (Regan, 1912)

Figure 15, Table 5

Oxyloricaria tamanae Regan, 1912: 669, plate LXXVII, figs. 1,

1a, and 1b (type locality: R´

ıo Tamana, R´

ıo San Juan, Choc ´

S.W. Colombia; two specimens, one male one female, 225

mm and 200 mm in TL; original description).

Sturisoma tamanae (Regan, 1912).—Regan, 1913: 470 (species

as valid in Sturisoma; without description). Eigenmann,

1920b: 11 (species distributed at the San Juan River in

Colombia); 1922: 95 (specimens from Istmina, Puerto

Negria, and the San Juan basin, Colombia). Fowler, 1942:

132 (distributed at the Tamana

´, Condoto, and San Juan

rivers, following Regan [1913] and Eigenmann [1922]).

Sturisomatichthys tamanae (Regan, 1912).—Isbr¨

ucker, 1979:

88, figs. 5–7 (original description of Sturisomatichthys;

species belonging to Sturisomatichthys; lectotype by subse-

quent designation [BMNH 1910.7.11: 133] female [?],

179.5 mm SL); 1980: 93 (included in Sturisomatichthys as

a member of the Harttiina; no diagnosis; reference of

Isbr¨

ucker [1979] for designation of lectotype). Isbr¨

ucker et

al., 2001: 32 (included in the catalog of the Loricariinae as

valid in Sturisomatichthys, German version). Isbr¨

ucker,

2002: 29 (included in the catalog of the Loricariinae as

valid in Sturisomatichthys, English version). Ferraris, 2003:

347 (genus distribution; synonymy; type material location

at BMNH). Maldonado-Ocampo et al., 2006: 150 (distrib-

uted at the Atrato River in Colombia; specimens at IAvH-

P). Ferraris, 2007: 300 (type locality; distribution of the

species). Maldonado-Ocampo et al., 2008: 195 (as distrib-

uted at Magdalena-Cauca basin, and Caribbean and Pacific

slopes in Colombia; citation of number lots of type series

in BMNH); 2013: 28 (distributed at Atrato, San Juan, and

Baud´

o rivers in Colombia; citation of type series at BMNH;

specimens at IAvH-P and IMCN in Colombia). Covain et

al., 2016: 514 (valid as Sturisomatichthys tamanae; phylo-

genetic relationships of the Loricariinae subfamily using

two mitochondrial and one nuclear markers; not included

792 Copeia 107, No. 4, 2019

in analysis). DoNascimiento et al., 2018: 77 (as present and

endemic in Colombia, Pacific slope).

Diagnosis.—Sturisomatichthys tamanae can be distinguished

from its congeners by the presence of a broad, almost square

spot at the origin of dorsal fin, occupying almost the entire

base of that fin (vs. spot absent), by lacking a dorsolateral

stripe from the tip of snout to the caudal peduncle (vs.

dorsolateral stripe present, except in S. caquetae,S. citurensis,

and S. guaitipan, new species), and lacking a rostrum (vs.

rostrum present, except in S. caquetae,S. citurensis, and S.

guaitipan, new species). It is distinguished from congeners by

the number of premaxillary teeth (60–93 vs. 40–56 in S.

aureus;33inS. caquetae; 54–61 in S. dariensis; 32–43 in S.

guaitipan, new species; and 38–45 in S. reinae, new species)

and dentary teeth (60–93 vs. 30 in S. caquetae; 34–47 in S.

dariensis; 33–42 in S. festivus; 31–46 in S. frenatus; 30–44 in S.

guaitipan, new species; 22–44 in S. leightoni; 32–44 in S.

panamensis; 28–36 in S. reinae, new species; and 26–39 in S.

varii, new species). Moreover, it is distinguished from S.

caquetae by the snout length (58.4–64.7% HL vs. 56.0% HL),

and from S. citurensis by having large, dark blotches

occupying almost the entire dorsal, pectoral, pelvic, and anal

fins (vs. small spots on rays, never occupying the entire fin,

nor entering the membranes), a triangular snout (vs. snout

rounded), and more numerous central abdominal plates (vs.

less numerous central abdominal plates).

Description.—Morphometric data in Table 5. Body short,

completely covered by dermal plates with well-developed

and numerous hypertrophied odontodes on sides of head of

males (Fig. 15). Dorsal profile of head straight from tip of

snout to origin of dorsal fin; concave from that point to

caudal-fin origin. Ventral body profile flat. Head and

predorsal region broader than remainder of body; postdorsal

region narrower towards caudal fin.

Head triangular and short; snout short, shorter in sexually

mature males (Fig. 15). Rostrum absent. Orbital rim circular,

without notch, dorsolaterally placed. Sphenotic with thin

and long lateral expansion. Canal-bearing plate broad and

well developed, in contact with anterior abdominal plates,

anterior to branchiostegal membrane, not reaching to rictal

barbel. Upper lip covered with small, almost imperceptible

plates; lower lip without such plates. Lower lip covered with

round small papillae, some almost imperceptible and others

well developed. Rictal barbel extremely short, free from lip

rim, never reaching to anterior border of abdominal plates,

nor surpassing posterior border of lower lip. Teeth slender,

bifid, 63–98 on premaxilla, 60–93 on dentary; cusps slightly

curved. Teeth in lower jaw larger than those in upper jaw.

Predorsal plates with two low, inconspicuous ridges;

nuchal plate twice size of predorsal plates (Fig. 15). Median

plates carinate, forming two longitudinal sharp keels; 16–19

coalescent plates, beginning at 13–15 median plates. Six to

nine straight lateral abdominal plates, between pectoral- and

pelvic-fin origins, larger than central abdominal plates.

Preanal plate bordered anteriorly by three well-developed

and similarly sized plates; anteriormost plate in contact with

the preanal plate at its most posterior tip. Central abdominal

plates between lateral abdominal plates, without defined

arrangement, as small, irregular plates (Fig. 15). Anterior

abdominal plates smaller and more numerous than posterior

ones.

Posterior margin of dorsal fin straight, unbranched ray

longest; dorsal-fin rays i,7. Pectoral-fin rays i,6, unbranched

ray longest, surpassing anterior third of pelvic fin. Pelvic-fin

rays i,5, unbranched ray longest, its tip reaching to anterior

third of anal fin. Anal-fin rays i,5, unbranched ray longest.

Posterior caudal-fin margin deeply concave, filaments on

Fig. 15. Sturisomatichthys tamanae, CAS 77238, 211.4 mm SL, Colombia, Choc´

o department, Puerto Negria and Istmina municipalities.

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 793

upper and lower caudal-fin rays. On pectoral fin, unbranched

ray with short but noticeable elongation, not as long as a

filament, but which can modify total extension of fin.

Coloration in alcohol.—Ground color of dorsum and head

dark brown or gray; pale brown or yellow ventrally.

Dorsolateral stripe absent. Anterior portion of body darker,

not as stripe, but as continuous spot from tip of snout to two

or three plates posterior to dorsal-fin origin (Fig. 15). Dark

stripe on dorsal fin, along first two branched rays. When

dorsal-fin stripe present, one to several white ocelli on distal

tip and just anterior to middle of fin (Fig. 15). Large, dark

spots from base but not reaching to distal tips on membranes

and rays of pectoral and pelvic fins, its length decreasing

from first to last ray on each fin, similar but not as

conspicuous as in S. frenatus. Well-developed dark spots

along unbranched ray of all fins. Caudal fin with dark spot at

base (Fig. 15).

Sexual dimorphism.—As shown by Regan (1912) in the

original description of S. tamanae, males of this species show

a marked sexual dimorphism (Fig. 15). Males appear to have a

shorter and broader snout and head (Fig. 15; Regan, 1912:

figs. 1a–1b) when compared to females. The males examined

show a well-developed sexual dimorphism consisting of

short and numerous odontodes on the sides of the head (Fig.

15).

Distribution.—Sturisomatichthys tamanae occurs in the Atrato

and San Juan rivers in the Pacific and Caribbean slopes of

Colombia (Fig. 3).

Remarks.—Sturisomatichthys tamanae lacks the dorsolateral

stripe characteristic of most species of Sturisomatichthys,a

feature shared with S. caquetae,S. citurensis, and S. guaitipan,

new species. Sturisomatichthys tamanae shares with S. citur-

ensis higher tooth counts on both the premaxilla (63–98 and

61–124) and the dentary (60–93 and 63–100). Nevertheless,

S. tamanae presents the following autapomorphies that

differentiate it from S. citurensis (see also Diagnosis and

identification key): cup-shaped portion of premaxilla relative

to cup-shaped portion of dentary smaller; and dorsal-fin

spinelet absent.

Eigenmann (1910) referred to Sturisoma tamanae as present

in the San Juan River, as did Eigenmann (1920b, 1922),

Fowler (1942), and Ferraris (2003, 2007); a record that is

corroborated here. On the other hand, Maldonado-Ocampo

et al. (2008, 2013), in addition to reporting the species in the

San Juan River, suggested that it occurs in the Magdalena-

Cauca basin, which is not corroborated here. The material

included by Maldonado-Ocampo et al. (2013) was not

Table 5. Morphometrics and meristics of Sturisomatichthys tamanae and Sturisomatichthys reinae, new species. Morphometric values as percents

of standard length (SL) and head length (HL). SD ¼standard deviation.

S. tamanae (n¼27) S.reinae (n¼9)

Min Max Mean SD Min Max Mean SD

Standard length 104 237.4 183.1 129.4 199 159.9

Head length 16.6 19.7 18.2 0.7 19.9 20.9 20.5 0.3

Predorsal length 26.7 30.0 28.5 0.8 29.4 32.3 31.1 0.9

Abdominal length 12.1 14.7 13.5 0.8 10.5 11.9 11.0 0.5

Trunk length 12.2 16.4 13.4 1.0 13.4 15.9 13.9 0.9

Dorsal fin length 23.3 33.4 26.8 2.6 26.1 30.3 29.3 1.7

Pectoral fin length 18.8 23.2 21.0 1.3 18.7 21.9 20.1 1.2

Pelvic fin length 15.4 18.4 16.6 0.8 14.8 17.1 15.6 0.8

Anal fin length 18.3 22.0 20.7 1.1 16.7 21.7 20.0 1.6

Head depth at supraoccipital tip 7.8 10.3 9.0 0.5 7.6 9.0 8.3 0.5

Body depth at dorsal-fin origin 8.8 13.0 11.2 1.0 9.6 11.5 10.4 0.7

Caudal peduncle depth 1.3 1.7 1.4 0.3 1.1 1.4 1.3 0.1

Cleithral width 10.9 13.4 12.5 0.6 8.3 9.6 9.1 0.4

Body width at dorsal-fin origin 11.1 14.9 13.5 0.9 9.8 12.4 11.0 0.8

Body width at anal-fin origin 9.5 12.5 11.4 0.7 8.3 10.6 9.3 0.7

Body width at caudal-fin origin 1.2 2.4 2.0 0.3 2.0 2.4 2.2 0.1

Head width 74.9 83.5 80.0 2.3 50.2 54.3 52.3 1.4

Interorbital distance 28.4 33.8 30.6 1.2 25.1 31.6 28.4 2.0

Orbital diameter 13.0 17.4 14.9 1.1 9.9 11.3 11.1 0.6

Snout length 58.4 64.7 60.7 1.3 60.4 64.8 63.7 1.3

Dentary length 12.1 16.9 14.5 1.1 7.6 9.8 9.2 0.7

Lower lip width 34.3 53.9 43.8 4.3 24.6 26.6 25.9 0.7

Plates on median series 13 15 14 14 16 15

Coalesced plates 16 19 17 15 18 17

Ventrolateral thoracic plates 6 9 8 7 8 8

Premaxillary teeth 63 98 65 38 45 42

Dentary teeth 60 93 63 28 36 33

Dorsal-fin rays i,7 i,7 i,7 i,7 i,7 i,7

Pectoral-fin rays i,6 i,6 i,6 i,6 i,6 i,6

Pelvic-fin rays i,5 i,5 i,5 i,5 i,5 i,5

Anal-fin rays i,5 i,5 i,5 i,5 i,5 i,5

Caudal-fin rays i,12,i i,12,i i,12,i i,12,i i,12,i i,12,i

794 Copeia 107, No. 4, 2019

examined and could not be attributed to either S. tamanae or

S. varii, new species (see also Remarks on S. varii, new

species). Maldonado-Ocampo et al. (2006, 2013) listed the

species as present in the Atrato and Baud ´

o rivers, which is

confirmed here. Thus, we conclude that S. tamanae is

distributed on both the Pacific and Caribbean versants of

Colombia (Fig. 3).

Isbr¨

ucker (1979) assigned as the lectotype for S. tamanae a

specimen of 177 mm SL, indicating that it could be a female.

That specimen was examined and confirmed as a female

when compared to the male paralectotype. In fact, the author

shows three figures (see figs. 5–7 in Isbr ¨

ucker, 1979) where

the sexual dimorphism of males is shown in detail,

corroborating the gender of the lectotype.

Material examined.—ANSP 198426, 1 alc, 137.3 mm SL,

Colombia, Choc´

o department, San Juan River basin, 4820N,

778260W; BMNH 1910.7.11.133, lectotype, 177.0 mm SL,

Colombia, Choc´

o department, Tamana River, San Juan River

system, 48590N, 768440W; BMNH 1910.7.11.134, 203.1 mm

SL, paralectotype, same data as lectotype; CAS 67414, 2 of 15

alc, 1 CS, 104.0–163.7 mm SL, Colombia, Choc ´

o department,

Istmina municipality, San Juan River basin, 5890N, 768410W;

CAS 77238, 6 alc, 169.4–211.4 mm SL, Colombia, Choc´

department, Puerto Negria and Istmina municipalities, 5890N,

768410W; CIUA 4628, 1 alc, 128.3 mm SL, Colombia,

Antioquia department, Vig´

ıa del Fuerte municipality,

Buchad´

o path, Atrato River basin, 6825032 00 N, 76845048 00 W;

CIUA 4788, 6 alc, 154.2–237.4 mm SL, Colombia, Antioquia

department, Murind´

o municipality, Atrato River basin,

Campo Alegre path, 6851029 00 N, 768490900 W; FMNH 55120,

4 of 8 alc, 184.4–198.4 mm SL, Colombia, Puerto Negria,

5890N, 768410W; FMNH 58332, 1 alc, 183.1 mm SL,

Colombia; FMNH 59507, 2 alc, 173.5–217.7 mm SL,

Colombia; USNM 167894, 1 alc, 177.1 mm SL, Colombia,

Choc´

o department, Istmina municipality, San Juan River,

5890N, 768410W; ZMA 115.842, 1 alc, 179.1 mm SL,

Colombia, Choc´

o department, San Juan River at Condoto,

58503000 N, 7683900000 W.

Sturisomatichthys reinae, new species

urn:lsid:zoobank.org:act:8F1FF661-827A-40E0-8CF2-

6C97B9B3BC17

Figure 16, Table 5

Holotype.—ICNMHN 24056 (formerly NRM 15155), 199.0

mm SL, Colombia, Choc´

o department, Baud ´

o River drainage,

Boca de Pep´

e, one of the various tributaries and river close to

village, 8859004 00 N, 778300000 W, S. O. Kullander and M. C.

Silvergrip, 12 February 1989.

Paratypes.—MCP 54152 (formerly NRM 15155), 2 alc, 160.1–

176.8 mm SL, same data as holotype; NRM 15155, 10 alc, 1

CS, 129.4–179.7 mm SL, same data as holotype.

Diagnosis.—Sturisomatichthys reinae is diagnosed from S.

caquetae,S. citurensis,S. guaitipan, new species, and S. tamanae

by possessing a rostrum (vs. rostrum absent) and a dorsolat-

eral stripe from the tip of snout to caudal peduncle (vs.

dorsolateral stripe absent). It is distinguished by the cleithral

width (8.3–9.6% SL vs. 10.5–12.8% SL in S. aureus; 9.9–10.5%

SL in S. dariensis; 10.3–11.7% SL in S. festivus; and 11.0–

14.1% SL in S. leightoni). It is also distinguished by the

number of premaxillary teeth (38–45 vs. 47–69 in S. aureus

and 54–61 in S. dariensis) and dentary teeth (28–36 vs. 40–56

in S. aureus). From S. festivus, it is distinguished by lacking

filaments on the dorsal, pectoral, pelvic, and anal fins (vs.

filaments present). From S. frenatus, it is distinguished by

lacking dark blotches on the pelvic and anal fins (vs. blotches

present). From S. leightoni, it is distinguished by having a

continuous, well-marked, and broad dorsolateral stripe (vs.

discontinuous and thin dorsolateral stripe), and by having a

large, dark blotch on the pectoral fin, occupying almost the

entire fin (vs. blotch absent). It is distinguished from S.

panamensis by having more numerous and larger central

abdominal plates (vs. fewer and smaller central abdominal

plates; Figs. 14, 16). Moreover, it is distinguished from S. varii,

new species, by lacking light brown blotches occupying the

entire pelvic and anal fins (vs. large light blotches present

and occupying entirely such fins) and dark reticulations on

the predorsal, supraoccipital, and interorbital region (vs. dark

reticulations present).

Description.—Morphometric data in Table 5. Body elongate,

slender, and somewhat depressed, completely covered by

dermal plates with short, scarce, developed, hypertrophied

odontodes on sides of head of males (Fig. 16). Dorsal profile

of head slightly concave from tip of snout to anterior border

of supraoccipital; straight from that point to origin of dorsal

fin. Concave posterior of dorsal fin to caudal-fin origin.

Ventral body profile flat. Postdorsal region narrower towards

caudal fin.

Head triangular and elongate in dorsal and ventral view.

Rostrum narrow, slender. Ventral border of rostrum flat.

Orbital rim circular, without notch, dorsolaterally placed.

Sphenotic without lateral expansion. Canal-bearing plate

broad and well developed, in contact with anterior abdom-

inal plates, anterior to branchiostegal membrane, not

reaching to rictal barbel. Upper lip covered with small,

almost imperceptible plates; lower lip without such plates.

Lower lip covered with round, small papillae, increasing in

size toward posterior border. Rictal barbel short, free from lip

rim, never reaching to anterior border of abdominal plates,

nor surpassing posterior border of lower lip. Teeth slender,

bifid, 38–45 on premaxilla, 28–36 on dentary; cusps slightly

curved. Teeth in lower jaw somewhat larger than those in

upper jaw.

Predorsal plates with low ridges, more pronounced on

sexually dimorphic males; nuchal plate at least twice size of

predorsal plates. Lateral plates carinate, forming two longi-

tudinal smooth keels, strong on dimorphic males, more

developed on caudal peduncle; 15–18 coalescent plates,

beginning at 14–16 median plates. Seven to eight lateral

abdominal plates, angled, between pectoral- and pelvic-fin

origins, larger than central abdominal plates. Preanal plate

bordered anteriorly by three well-developed plates; anterior-

most plate always in contact with preanal plate, and larger

than two other plates. Central abdominal plates, between

lateral abdominal plates, not arranged in rows; anterior

abdominal plates more numerous than posterior ones, and

most anterior ones longer than remaining (Fig. 16).

Posterior margin of dorsal fin straight, unbranched ray

longest; dorsal-fin rays i,7. Pectoral-fin rays i,6, unbranched

ray longest, reaching to anterior third of pelvic fin. Pelvic-fin

rays i,5, unbranched ray longest, its tip surpassing anterior

third of anal fin. Anal-fin rays i,5, unbranched ray longer.

Posterior caudal-fin margin deeply concave, with filaments

on lower unbranched ray. Unbranched ray on dorsal,

pectoral, pelvic, and anal fins with short elongation, not as

long as filaments present on caudal fin.

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 795

Coloration in alcohol.—Ground color of dorsum and head

pale or dark brown; yellow ventrally. Poorly defined

dorsolateral stripe on both sides of body. Stripe broader from

anterior border of first predorsal plate to origin of dorsal fin,

then becoming less conspicuous and thin towards caudal

peduncle (Fig. 16). On dorsal portion of head, several thin,

brown, sinuous lines between dorsolateral stripe on snout to

anterior border of first predorsal plate (Fig. 16). Remnant of

dark stripe sometimes present on ventral side of rostrum, not

separated as observed on both sides of body, but as consistent

dark blotch (Fig. 16). Small spots on rays of dorsal, pectoral,

pelvic, and anal fins. Dorsal and pectoral fins with dark

brown stripe along first two branched rays, with conspicuous

dark spots along unbranched ray. Caudal fin with dark stripes

along first two or three branched rays, both upper and lower,

connected at base of fin, forming V-shaped pattern. Dark

brown spots on edges of caudal lobes.

Sexual dimorphism.—The males examined show a subtle

dimorphism on the sides of the head, consisting of scarce

and short hypertrophied odontodes (Fig. 16).

Distribution.—Sturisomatichthys reinae is known from the

Baud´

o River basin, Pacific versant, in Colombia (Fig. 1).

Etymology.—The specific name honors Mrs. Ruth Gisela

Reina, late curator of fishes at STRI, for her contributions to

the knowledge of fishes and invaluable help and assistance

for several ichthyologists around the world during her time

at the STRI.

Remarks.—Sturisomatichthys reinae is sympatric with S. aureus

in the Baud´

o River, in Colombia. Regarding S. aureus,

Steindachner (1900, 1902) did not describe any coloration

pattern in his original description. In fact, the drawing

presented by Steindachner (1902, figs. 1 and 1a) shows no

evidence of the dorsolateral stripe characteristic of some

species of Sturisomatichthys. This could be due to the poor

preservation of color in the specimens collected in 1898 (see

Remarks on S. aureus), but observed across specimens

examined. Nevertheless, several characters were found to

distinguish S. reinae from S. aureus (see Diagnosis and

identification key).

Sturisomatichthys guaitipan, new species

urn:lsid:zoobank.org:act:03E476CF-45F7-4B60-9251-

1D46C6C4A840

Figure 17, Table 6

Holotype.—ICNMHN 24055 (formerly ICNMHN 1146), 90.7

mm SL, Colombia, Cundinamarca department, Apulo,

Magdalena River basin, Calandaima River, 48300N, 748340W,

P. Cala, 14 February 1970.

Paratypes.—ICNMHN 1146, 13 alc, 72.8–82.1 mm SL, same

data as holotype; ICNMHN 9237, 16 alc, 79.6–111.7 mm SL,

Fig. 16. Sturisomatichthys reinae, new species, holotype, ICNMHN 24056, 199.0 mm SL, Colombia, Choc ´

o department, Baud ´

o River drainage, Boca

de Pep´

e, various tributaries and river close to village.

796 Copeia 107, No. 4, 2019

MCP 54154, 3 alc, 91.2–102.8 mm SL, Colombia, Huila

department, Yaguara

´, Magdalena River basin, Yaguara

´River,

Valencia Bridge, 28400N, 758300W, P. Cala, 13 February 1991;

IUQ 1030, 1 alc, 70.2 mm SL, Colombia, Huila department,

Guadalupe, Magdalena River basin, Suaza River, La Viciosa

Creek, Villa Luz farm, around Guadalupe, 28100N, 758400W,

C. Roma

´n-Valencia and A. Roma

´n-Valencia, 18 December

1998.

Diagnosis.—Sturisomatichthys guaitipan is diagnosed from its

congeners by having ten branched rays on the caudal fin (vs.

12); having spots along the dorsal surface of snout and head

(vs. spots absent or present only on snout and rostrum in S.

leightoni); and, except from S. caquetae,S. citurensis, and S.

tamanae, by lacking dorsolateral stripes (vs. stripes present)

and a rostrum (vs. rostrum present). It is further distin-

guished from S. citurensis and S. tamanae by the number of

dentary teeth (30–44 vs. 63–100 and 60–93, respectively).

Sturisomatichthys guaitipan is distinguished from S. leightoni

by having ten branched rays on caudal fin (vs. 12); by lacking

dorsolateral stripes (vs. stripes present); by having spot

dorsally on snout and head (vs. only snout and rostrum

with few spots). Finally, S. guaitipan is externally more similar

to S. caquetae, from which it can be diagnosed by the trunk

length (14.9–16.7% SL vs. 14.4% SL); body depth at dorsal-

fin origin (10.3–12.1% SL vs. 9.8% SL); ten branched rays on

caudal fin (vs. 12); snout and head dorsally with spots (vs.

spots absent).

Description.—Morphometric data in Table 6. Body short and

low, completely covered by dermal plates. Males with few,

short, and poorly developed odontodes on sides of head and

unbranched rays of pectoral and pelvic fins. Dorsal profile of

head straight from tip of snout to origin of dorsal fin; slightly

concave from that point to caudal-fin origin. Ventral body

profile flat. Head and predorsal region broader than remain-

der of body.

Head triangular. Rostrum absent. Orbital rim circular,

without notch, dorsolaterally placed on head. Sphenotic

with short lateral expansion. Canal-bearing plate broad and

well developed, in contact with anterior abdominal plates,

anterior to branchiostegal membrane, and not reaching to

rictal barbel. Rictal barbel free from lip, usually reaching to

almost same length as lower lip. Upper lip covered with

small, almost imperceptible plates; lower lip without such

plates. Lower lip covered with round, small, almost imper-

Fig. 17. Sturisomatichthys guaitipan, new species, holotype, ICNMHN 24055, 90.7 mm SL, Colombia, Cundinamarca, Apulo, Magdalena River basin,

Calandaima River. Photo by Henry Agudelo (ICNMHN).

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 797

ceptible papillae, which decrease in size towards posterior

border of lip. Posterior border of lower lip irregular, with

small papillae present along its entire border. Teeth slender,

bifid, 32–43 on premaxilla, 30–44 on dentary. Teeth strongly

curved, cusps straight, on premaxillary and dentary teeth;

teeth equal size on both premaxillary and dentary bones.

Predorsal plates without conspicuous ridges; nuchal plate

twice size of predorsal plates. Lateral plates carinate, forming

two longitudinal keels; 12–15 coalescent plates, beginning at

13–16 median plates. Six to eight curved lateral abdominal

plates, between pectoral- and pelvic-fin origins, longer and

larger than central abdominal plates. Anterior abdominal

plates not reaching posterior border of lower lip; same size as

central abdominal plates. Preanal plate bordered anteriorly

by three, somewhat developed, and similarly sized plates, all

in contact with preanal plate. Central abdominal plates,

between lateral abdominal plates, without any kind of

organization; numerous, small, and irregular.

Posterior margin of dorsal fin straight, unbranched and

first and second branched rays longest; dorsal-fin rays i,7.

Pectoral-fin rays i,6, unbranched ray longest, reaching to

origin of pelvic fin. Pelvic-fin rays i,5, unbranched ray

longest, its tip surpassing origin of anal-fin. Anal-fin rays

i,5, unbranched ray longest. Caudal-fin rays i,10,i. Posterior

caudal-fin margin deeply concave, short filaments on both

upper and lower ray.

Coloration in alcohol.—Ground color of dorsum, head, and

ventral surface light brown. Snout and head with irregular,

small spots along entire dorsal surface, reaching to origin of

dorsal fin (Fig. 17). Dark, discontinuous, narrow stripe at

beginning of dorsal fin, similar to S. leightoni (Figs. 13, 17).

Short, discontinuous stripes connected to dorsolateral stripe

at beginning of dorsal fin (Fig. 17), reaching to seventh or

eighth plate on caudal peduncle (Fig. 17). Four to five

transverse, inconspicuous bands along caudal peduncle. All

fins with dark and visible spots, no stripes (Fig. 17).

Sexual dimorphism.—Males with short, poorly developed,

hypertrophied odontodes on sides of the head and along the

plates of the caudal peduncle (Fig. 17).

Distribution.—Sturisomatichthys guaitipan occurs in the upper

and middle Magdalena River basin in Colombia (Fig. 1).

Etymology.—The specific epithet guaitipan honors Gaitana,

an indigenous heroine from the 16

century, cacique of the

Timana

´tribe, who occupied the Colombian Andes in the

upper Magdalena valley in the Meta department. Gaitana, or

Guaitipan as she was also known, fought against the Spanish

invasion that occurred in South America between 1539 and

1540, ending in her defeat due to treason by the cacique

Matambo, who warned the Spanish army of the rebellion led

Table 6. Morphometrics and meristics of Sturisomatichthys guaitipan, new species, and Sturisomatichthys varii, new species. Morphometric values

as percents of standard length (SL) and head length (HL). SD ¼standard deviation.

S.guaitipan (n¼25) S. varii (n¼20)

Min Max Mean SD Min Max Mean SD

Standard length 70.2 111.7 86.2 114.7 194 157.9

Head length 22.0 25.5 23.4 0.9 18.3 20.6 19.4 0.7

Predorsal length 32.3 35.8 34.3 0.8 28.3 31.8 29.9 1.0

Abdominal length 12.7 15.5 13.8 0.8 10.7 13.0 11.6 0.6

Trunk length 14.9 16.7 16.2 0.5 12.5 15.1 13.6 0.8

Dorsal fin length 20.3 25.6 23.4 1.4 26.6 31.1 29.2 1.4

Pectoral fin length 17.6 23.0 19.8 1.4 17.8 21.7 19.9 1.0

Pelvic fin length 15.3 19.2 17.2 1.2 14.8 17.6 15.9 0.7

Anal fin length 15.6 20.5 18.4 1.4 17.7 22.8 20.4 1.2

Head depth at supraoccipital tip 9.3 11.3 10.2 0.5 7.1 9.0 8.0 0.5

Body depth at dorsal-fin origin 10.3 12.1 10.9 0.5 8.7 10.8 10.0 0.6

Caudal peduncle depth 1.4 1.9 1.6 0.1 1.1 1.3 1.2 0.1

Cleithral width 14.0 16.2 15.2 0.6 7.7 9.2 8.5 0.4

Body width at dorsal-fin origin 12.7 16.1 14.3 0.9 9.3 11.3 10.4 0.6

Body width at anal-fin origin 10.3 12.1 11.0 0.4 7.8 9.9 9.1 0.5

Body width at caudal-fin origin 1.3 2.4 2.0 0.3 1.8 2.3 2.1 0.1

Head width 66.5 80.6 74.8 3.9 48.1 57.5 54.1 2.3

Interorbital distance 27.3 33.5 30.4 1.6 27.0 33.1 29.6 1.7

Orbital diameter 13.6 18.1 15.0 1.1 10.0 13.5 11.8 0.9

Snout length 55.4 70.5 58.3 3.0 56.6 64.4 62.9 1.7

Dentary length 10.1 15.6 12.9 1.4 9.1 11.9 10.3 0.8

Lower lip width 38.2 48.3 41.4 2.8 24.9 31.9 29.4 1.8

Plates on median series 13 16 15 14 16 15

Coalesced plates 12 15 13 16 19 17

Ventrolateral thoracic plates 6 8 7 6 9 8

Premaxillary teeth 32 43 38 30 55 40

Dentary teeth 30 44 35 26 39 32

Dorsal-fin rays i,7 i,7 i,7 i,7 i,7 i,7

Pectoral-fin rays i,6 i,6 i,6 i,6 i,6 i,6

Pelvic-fin rays i,5 i,5 i,5 i,5 i,5 i,5

Anal-fin rays i,5 i,5 i,5 i,5 i,5 i,5

Caudal-fin rays i,10,i i,10,i i,10,i i,12,i i,12,i i,12,i

798 Copeia 107, No. 4, 2019

by Gaitana. Finally in 1626, joining forces with the Pijaos,

Panaes, and Pamaos, they obtained victory by expelling the

Spanish from their region.

Remarks.—Sturisomatichthys guaitipan has ten branched rays

on the caudal fin. According to several authors (i.e., Rapp Py-

Daniel, 1997; Covain and Fisch-Muller, 2007; Covain et al.,

2008, 2016; Rodriguez et al., 2011; Londo˜

no-Burbano and

Reis, 2016), species belonging to the Loricariina have ten

branched rays on the caudal fin (except Fonchiiloricaria

nanodon, which has 12 branched caudal rays), while species

belonging to the Harttiini and Farlowellina always have 12

branched rays. That character was always found to support

the separation of tribes within Loricariinae. Along with the

count of ten branched rays on the caudal fin, species of the

Loricariina always have less than 20 teeth on premaxilla and

dentary. On the other hand, species of Harttiini and

Farlowellina, always have more than 20 teeth on premaxilla

and dentary. Sturisomatichthys guaitipan have characteristics

of both the Loricariina and the Farlowellina (i.e., ten

branched rays on the caudal fin and more than 20 teeth on

premaxilla and dentary). The species has characteristics that

are diagnostic of Sturisomatichthys and falls within the

definition of the genus presented here. Even though the

allocation of the new species within Sturisomatichthys is well

supported, its inclusion in a phylogenetic context, with the

use of molecular markers, is extremely important to corrob-

orate its placement within Farlowellina.

The ichthyofauna present in the Magdalena basin has been

object of numerous studies (e.g., Eigenmann, 1920a, 1922;

Miles, 1947; Dahl, 1971; Zu˜

niga et al., 2004; Maldonado-

Ocampo et al., 2005; Mojica et al., 2006a; Villa-Navarro et al.,

2006) in which composition and taxonomic status of species

present in the basin were described. Throughout these

studies, species currently belonging to both Sturisomatichthys

and Sturisoma were included. As stated above, S. guaitipan

occurs in the upper and middle portions of the Magdalena

River basin, and the authors mentioned above could have

examined material belonging to that region. Nevertheless,

due to the scarce knowledge of Sturisomatichthys and allied

genera (even to date), the authors may have been overlooked

the fact that such material belonged to an undescribed

species and cited them as another species (most likely S.

leightoni).

Most of the material cited by the above authors was not

examined here, so any specimens can be unmistakably

assigned to the new species. However, by describing the

new taxon, the number of species of Sturisomatichthys present

in the Magdalena River increases to three and it shows how

unexplored that basin still is, even though it is the most

studied basin in Colombia. Further studies including material

already analyzed by authors from the 19

and 20

centuries

will likely yield new taxa for several groups present in the

Andean basins.

Sturisomatichthys varii, new species

urn:lsid:zoobank.org:act:7E985803-31B2-4E56-819C-

7EE871425079

Figure 18, Table 6

Holotype.—IAvH-P 22003 (formerly CAS 77225), 190.3 mm

SL, Colombia, Choc ´

o department, San Juan River, Condoto

River, an upper Eastern tributary of San Juan River, 5850N,

768390W, C. Wilson, 1913.

Paratypes.—CAS 77225, 2 alc, 89.4–137.9 mm SL, same data

as holotype; CAS 246603 (formerly CAS 67414), 6 alc, 1 CS,

85.8–166.7 mm SL, Colombia, Choc ´

o department, Istmina,

San Juan River basin, 58904800N, 7684101200 W, A. W. Henn and

C. Wilson, 1913; CAS 77224, 1 alc, 145.2 mm SL, Colombia,

Choc´

o department, San Juan River, Tado District, Tado River

emptying into San Juan River, elevation 316 feet, 581504800N,

76833036 00W, C. Wilson, 1913; CAS 77226, 8 alc, 142.8–194.0

mm SL, Colombia, Puerto Negria and Istmina municipalities,

5890N, 768410W, C. H. Eigenmann, 14–20 March 1912; FMNH

58333, 3 alc, 144.6–175.8 mm SL, Colombia, Choc´

department, Condoto, San Juan River basin, 585040 00 N,

768390100 W, C. E. Wilson, 1913; MCP 54153 (formerly CAS

77226), 2 alc, 163.1–166.2 mm SL, Colombia, Puerto Negria

and Istmina municipalities, 5890N, 768410W, C. H. Eigen-

mann, 14–20 March 1912; NRM 10683, 1 of 2 alc, 185.9 mm

SL, Colombia, Choc ´

o department, San Juan River, headwa-

ters, 48204200 N, 77826 02900 W, G. Gerring, 1939.

Diagnosis.—Sturisomatichthys varii is diagnosed from S.

caquetae,S. citurensis,S. guaitipan, and S. tamanae by having

a dorsolateral stripe (vs. dorsolateral stripe absent) and a

rostrum (vs. rostrum absent). It is further distinguished from

congeners by the cleithral width (7.7–9.2% SL vs. 10.5–12.8%

SL in S. aureus, 9.9–10.5% SL in S. dariensis, 10.3–11.7% SL in

S. festivus, and 11.0–14.1% SL in S. leightoni) and head width

(48.1–57.5% HL vs. 60.4–79.6% HL in S. aureus, 65.3–73.7%

HL in S. festivus, and 58.4–75.0% HL in S. leightoni). It is

distinguished from S. panamensis and S. reinae by having dark

reticulations on the predorsal, supraoccipital, and interorbit-

al regions (vs. dark reticulations absent). It is further

distinguished from S. reinae by having light brown blotches,

which do not occupy the entire pelvic and anal fin (vs.

blotch absent on both fins). From S. leightoni it is distin-

guished by the number of coalescent plates (16–19 vs. 13–

16), a shorter caudal peduncle depth (1.1–1.3% SL vs. 1.3–

2.1% SL), a continuous dorsolateral stripe (vs. dorsolateral

stripe discontinuous), and light blotches occupying almost

the entire pectoral and pelvic fins (vs. light blotch absent on

such fins). It is distinguished from S. frenatus by having light

brown blotches, which do not occupy the entire dorsal,

pectoral, pelvic, and anal fins (vs. dark blotches present and

occupying the entire such fins). It is distinguished from S.

aureus by lacking a transverse bar at the dorsal-fin origin

united to the dorsolateral stripe (vs. transverse bar united to

dorsolateral stripe present) and number of dentary teeth (40–

56 vs. 26–39), and from S. festivus, by lacking filaments on

dorsal, pectoral, pelvic, and anal fins (vs. filaments present

on such fins).

Description.—Morphometric data in Table 6. Body elongate,

slender, and depressed, completely covered by dermal plates,

males with small hypertrophied odontodes on sides of head

(Fig. 18). Dorsal profile of head slightly concave from tip of

snout to anterior border of supraoccipital; straight from that

point to origin of dorsal fin. Concave from dorsal fin to

caudal-fin origin. Ventral body profile flat. Postdorsal region

narrower towards caudal fin.

Head triangular and long. Rostrum present, slender; its

border flat. Orbital rim circular, without notch, dorsolaterally

placed on head. Sphenotic without lateral expansion. Canal-

bearing plate broad and well developed, in contact with

anterior abdominal plates, anterior to branchiostegal mem-

brane, not reaching to rictal barbel. Upper lip covered with

small, almost imperceptible plates; lower lip without such

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 799

plates. Lower lip covered with round, small papillae,

increasing in size toward anterior border of lower lip. Rictal

barbel short, free from lip rim, never reaching to anterior

border of abdominal plates, nor surpassing posterior border

of lower lip. Teeth slender, bifid, 30–55 on premaxilla, 26–39

on dentary; cusps slightly curved. Teeth in lower jaw larger

than those in upper jaw.

Predorsal plates without ridges; nuchal plate at least twice

size of predorsal plates. Lateral plates carinate, forming two

longitudinal ill-developed keels; 16–19 coalescent plates,

beginning at 14–16 median plates. Six to nine lateral

abdominal plates, angled, between pectoral- and pelvic-fin

origins, larger than central abdominal plates. Preanal plate

bordered anteriorly by three well-developed plates; anterior-

most plate sometimes not in contact with preanal plate, and

larger than two other plates. Central abdominal plates,

between lateral abdominal plates, without arrangement;

anterior abdominal plates more numerous than posterior

ones, and most anterior ones longer than remaining (Fig. 18).

Posterior margin of dorsal fin straight, unbranched and

first branched rays longest; dorsal-fin rays i,7. Pectoral-fin

rays i,6, unbranched ray longest, surpassing anterior third of

pelvic fin. Pelvic-fin rays i,5, unbranched ray longest, its tip

surpassing anterior third of anal fin. Anal-fin rays i,5,

unbranched and first branched rays longest. Posterior

caudal-fin margin deeply concave, with filaments on both

upper and lower unbranched rays. Unbranched ray on

dorsal, pectoral, pelvic, and anal fins with short elongation,

not as long as filaments on caudal fin.

Coloration in alcohol.—Ground color of dorsum and head

pale or dark brown; yellow ventrally. Dorsolateral stripe

broader from anterior border of first predorsal plate to origin

of dorsal fin (Fig. 18), becoming less conspicuous and thin

towards caudal peduncle. Remnant of dark stripe on ventral

side of rostrum, separated as observed on both sides of body

(Fig. 18). Large, light blotch on pectoral, pelvic, and anal fins.

Dorsal fin with dark brown stripe along first two branched

rays, and with conspicuous dark spots along unbranched ray.

Caudal fin with dark stripes along first two or three branched

rays, both upper and lower, and connected at base of fin,

forming V-shaped pattern. Dark brown spots on edges of

caudal lobes.

Sexual dimorphism.—The males examined show sexual

dimorphism consisting of short hypertrophied odontodes

on the sides of head (Fig. 18).

Distribution.—Sturisomatichthys varii occurs in the San Juan

River basin, in Condoto and Tado River, in Colombia (Fig. 1).

Etymology.—The specific name honors Richard P. Vari for his

important and numerous contributions to the knowledge of

Neotropical fishes, and for his interest in helping young

ichthyologists pursue careers in science to continue to

explore the diversity of such Neotropical fishes. Richard left

an important legacy which, with no doubt, will help present

and future ichthyologists in their research. The conclusion of

this study would not have being possible without the selfless

and extensive help received from Richard.

Remarks.—Sturisomatichthys varii is sympatric with S. tam-

anae. Both species can be easily differentiated externally by

the presence of a rostrum on S. varii (vs. rostrum absent), as

well as by the presence of dorsolateral stripe on the sides of

body (vs. stripe absent; Figs. 15, 18). Moreover, the following

body proportions are useful for their differentiation: shorter

cleithral width (7.7–9.2% SL vs. 10.9–13.4% SL), narrower

head (48.1–57.5% HL vs. 74.9–83.5% HL) and lower lip

(24.9–31.9% HL vs. 34.3–53.9% HL), and fewer dentary teeth

(26–39 vs. 60–93).

The new species is only known from the San Juan River

basin in Colombia. There is only one report that could be

related to additional specimens other than the type series of

the species (Maldonado-Ocampo et al., 2013). As pointed out

above (see Remarks on S. panamensis and S. tamanae), the

authors included material from Colombian museums as

belonging to S. panamensis. Nevertheless, that material was

not examined and could belong either to the new species or

to S. tamanae. In any case, the distribution of the new species

independent from the identity of that material remains

restricted to the San Juan basin.

Fig. 18. Sturisomatichthys varii, new species, holotype, IAvH-P 22003, 190.3 mm SL, Colombia, Choc ´

o department, San Juan River, Condoto River,

an upper Eastern tributary of San Juan River.

800 Copeia 107, No. 4, 2019

DISCUSSION

Isbr¨

ucker and Nijssen (in Isbr¨

ucker, 1979) described Sturiso-

matichthys as related to Sturisoma, diagnosing the genus by

the absence of a rostrum; presence or absence of a rostrum

has been demonstrated to be uninformative by Covain et al.

(2008). It was observed that the rostrum could be present or

absent on some species of Sturisoma and Sturisomatichthys.

For instance, S. caquetae,S. citurensis,S. guaitipan, and S.

tamanae lack an enlarged rostrum, as well as Sturisoma

brevirostre; the remaining species of both genera do have an

enlarged rostrum, whose development varies between species

(Fig. 19).

One character that was not mentioned by Isbr¨

ucker and

Nijssen (in Isbr¨

ucker, 1979) is the type of covering of the

abdominal plates. Species of Sturisoma have the abdomen

totally covered by large, square, central abdominal plates

arranged in three longitudinal well-defined series (Fig. 19A;

see also Covain and Fisch-Muller, 2007). On the other hand,

species of Sturisomatichthys have the abdomen totally covered

by small to medium, irregular, central abdominal plates,

which are not arranged in clear series (Fig. 19B, C). Even in

juveniles of species of both genera, that arrangement can be

seen. In addition, the anterior abdominal plates of Sturiso-

matichthys are not as enlarged as those in Sturisoma. Those

plates are generally of similar size to the central abdominal

plates in Sturisomatichthys, and generally reach the posterior

border of the lower lip. On the contrary, Sturisoma has

enlarged anterior abdominal plates, generally not reaching

the posterior border of the lower lip, and with a small naked

area on the cleithrum (Fig. 19A).

Originally, Sturisomatichthys encompassed three trans-An-

dean (S. citurensis,S. leightoni, and S. tamanae), and one cis-

Andean species (Sturisomatichthys caquetae). Covain et al.

(2016) proposed Sturisomatichthys to include only trans-

Andean species plus six species previously included in

Sturisoma (S. aureus,S. dariensis,S. festivus,S. frenatus,S. kneri,

and S. panamensis), and transferring S. caquetae to Sturisoma.

However, as discussed above, S. caquetae is a member of

Sturisomatichthys, and fully cis-Andean, and also there are

populations of S. leightoni present in the Orinoco basin, at

Cumaral, Meta, Colombia (Fig. 3; see Remarks on S. leightoni),

which was already been suggested by Eigenmann (1922).

Through the transfer of S. caquetae back from Sturisoma to

Sturisomatichthys and the confirmation of the specimens

collected by N. Maria and studied by Eigenmann (1922) as S.

leightoni from the Orinoco basin, Dasyloricaria remains the

only strictly trans-Andean genus in Loricariinae (Covain and

Fisch-Muller, 2007; Londo˜

no-Burbano and Reis, 2016).

Ghazzi (2003) performed a morphology-based phylogenet-

ic analysis of Sturisoma. One of the main results discussed by

that author is the inclusion of the trans-Andean species

previously included in Sturisoma, in her proposed new genus

called ‘‘ ‘Trans’ new genus.’’ Ghazzi (2003) included S.

leightoni in her analysis and found the species as sister to

her new genus. Because of that result, Ghazzi did not include

her new trans-Andean species in Sturisomatichthys and

proposed a new genus. The author diagnosed the new genus

by the presence of a dorsolateral stripe, assuming those as

unique for the new genus. As discussed above (see remarks on

S. leightoni), the type species of Sturisomatichthys does possess

that dorsolateral stripe, although in a somewhat different

format than its congeners (Fig. 13). In addition, the same

kind of stripe is also present in Farlowella and some species of

Sturisoma, rendering that character uninformative for a genus

diagnosis. As shown in the Diagnosis for Sturisomatichthys,

there are informative characters to maintain the species

proposed by Ghazzi as belonging to her new genus (i.e., S.

festivus,S. frenatus, and S. panamensis)inSturisomatichthys.

Therefore, we disagree with the necessity of a new genus, but

instead propose the inclusion of the species in Sturisomatich-

thys, corroborating the hypothesis presented by Covain et al.

(2016). Ghazzi (2003) also pointed out that the fact that S.

leightoni lacked a rostrum, which, according to her and the

original description of Sturisomatichthys (in Isbr¨

ucker, 1979),

Fig. 19. Anterior ventral view showing differences in central abdominal plate organization and rostrum length among (A) Sturisoma graffini, MUSM

58700, 187.6 mm SL; (B) Sturisomatichthys citurensis, USNM 78361, 196.3 mm SL; and (C) Sturisomatichthys festivus, AUM 22138, 161.3 mm SL.

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 801

is diagnostic of the genus, also differentiated Sturisomatich-

thys from her ‘‘ ‘Trans’ new genus.’’ Nevertheless, as men-

tioned above, that character is variable between Sturisoma

and Sturisomatichthys and thus not useful to diagnose either

of those genera. Two aspects highlighted by Ghazzi (2003)

while defining her new genus deserve mention. The first is

related to the inclusion of S. kneri in Sturisoma and not in her

new genus. This taxonomic decision disagrees with her

diagnosis of the new genus, since S. kneri also possesses a

dorsolateral stripe (Ghazzi, 2005), the main character used to

diagnose the new genus. On the other hand, Ghazzi (2003)

discussed that the position of S. kneri as part of Sturisoma was

not fully resolved (see fig. 42 in Ghazzi, 2003), thus

reinforcing the hypothesis that the species belongs to

Sturisomatichthys. The second issue is related to S. aureus,as

Ghazzi (2003) had no access to any material identified as that

species. She discussed that because of the absence of the

holotype of S. aureus (see Remarks on that species), it was

impossible to unequivocally identify specimens as S. aureus.

In any case, Ghazzi (2003) assumed the species to belong to

her new genus (here, Sturisomatichthys), but considering it as

incertae sedis. We were able to identify specimens as S. aureus

after the translation of the only two available descriptions of

the species (Steindachner, 1900, 1902), and based on the

localities described by the author and subsequent inclusion

of the species on distribution lists (see References for S.

aureus).

As described by Schaefer (2011), the collision of the

Panama-Choco island arc with the northwestern margin of

the South American plate from 12 to 6 Mya was primarily

responsible for the uplift of the Eastern Cordillera. The

narrow Cauca-Patia valley separates the Western and Central

cordilleras, while the broader and elongate Magdalena Valley

separates the Central and Eastern cordilleras. To the north

and continuing into the Caribbean Sea, the Eastern Cordil-

lera diverges to form the low Serrania de Perija

´range to the

west and the higher M´

erida Andes to the east of Lago

Maracaibo. This series of events illustrates the formation of

the diverse ecosystems present in northern South America.

Due to the orogenic formations, modifications in river flow

direction and isolation of river basins, new niches were

available for the already existing populations. Furthermore,

the barriers generated with the uplift of the cordilleras

resulted in isolation of those populations and created new

opportunities for diversification, which is likely what

occurred for the populations of species of Sturisomatichthys.

The north–south orientation of the cordilleras forms a

natural barrier to the prevailing atmospheric circulation

patterns, thereby creating major climatic and ecological

differentiation and a complex variety of ecosystems between

cis- and trans-Andean regions (Schaefer, 2011). This distinc-

tion of the ecosystems west and east of the central and

western cordilleras could be of great importance to the

biogeographical component observed for Sturisomatichthys.

Ecological studies involving the genus are needed to support

this statement.

The Magdalena and Cauca basins are generally considered

as a single drainage. Nevertheless, although the Cauca River

is a tributary of the Magdalena River, and consequently is

included within the latter basin in most classifications,

Schaefer (2011) regarded it as a separate and distinct

biogeographic unit because the two hydrosheds are indeed

distinct at higher elevations and, in their headwater regions,

separated by the Cordillera Central. This could explain the

absence of S. aureus from the Cauca basin. That species is

distributed in the lower Magdalena basin, but no records

were found on any of the Cauca localities. Sturisomatichthys

aureus is possibly restricted to lower elevations, characteristic

of the lower Magdalena portion, contrary to S. leightoni. The

latter is present in both the Magdalena and Cauca basins,

which could be related to ecological characteristics of the

species, and smaller size of adult individuals. Specimens of S.

aureus can reach to .200 mm SL, while S. leightoni do not

surpass 120 mm SL, and this allows the latter to explore

different, and perhaps, more diverse resources.

According to Rodr´

ıguez-Olarte et al. (2011), the trans-

Andean fish fauna has high species richness and endemism,

an ancestral relationship with the Amazon and Orinoco

biotas, and, for some families and genera, represents the

northern limit of their distributions, as is the case for

Sturisomatichthys. The authors estimated that among Pacific

drainages, the San Juan has the highest number of species

(95, 10 endemic), and in the Caribbean drainages, those

with the most species were the Magdalena (159, 66

endemic), Atrato (120, 19 endemic), and Maracaibo (115,

48 endemic), all located in humid regions. Within Stur-

isomatichthys, that assumption is met, since there are two

species present in the San Juan River (one of them

endemic), three in the Magdalena River (two of them

endemic), two in the Atrato, and one in the Maracaibo

Basin (endemic).

As noted above, there are similarities among trans-

Andean drainages regarding the presence of species of

Sturisomatichthys. There is evidence that before the uplift of

the Darien mountain range, the Atrato River emptied into

the Tuyra Gulf on the Pacific slope of Panama (Rodr´

ıguez-

Olarte et al., 2011). This is suggested by the large size of the

Tuyra River delta, which is disproportionate for a river of its

size. This also would explain the high proportion of species

shared between the Atrato and the Pacific drainages

(Rodr´

ıguez-Olarte et al., 2011). This is the case for S.

citurensis and S. panamensis, which are present in both the

Atrato and the Tuyra rivers. Relationships regarding

ichthyofaunal similarities between those basins, specific

to the Loricariinae, were also documented in Dasyloricaria

latiura, which is also present in both drainages (Londo ˜

no-

Burbano and Reis, 2016). Nevertheless, Rodr´

ıguez-Olarte et

al. (2011) also commented that the fish fauna of the Tuyra

province has low similarity with that of the Atrato,

suggesting rapid speciation and/or extinction associated

with the orogeny of the mountains separating these basins.

This is not the case for Sturisomatichthys,althoughweagree

that a rapid speciation could be occurring on both rivers,

especially the Tuyra River, suggested as a refuge from a

diversity point of view. There are three species of Sturiso-

matichthys in that river, S. citurensis,S. dariensis,andS.

panamensis. This is a single example of the great diversity

housed in that drainage, due to a possible unique internal

structure and conditions that resulted from the orogenic

history of the Pacific of Panama, which led to the current

geographical composition of that area. In addition, as

pointed out by Rodr´

ıguez-Olarte et al. (2011), a few other

possible refugia can be detected, such as the Atrato of

northern Colombia, where precipitation is among the

highest recorded for the world and produces the highest

water discharge. A similar situation exists in the watersheds

of the nearby Dari´

en region of Panama; this supports the

idea that the unique pattern of this area plays a key role in

such great diversity in such a small region.

802 Copeia 107, No. 4, 2019

KEY TO THE SPECIES OF STURISOMATICHTHYS

1a. Rostrum not enlarged; dorsolateral stripe on sides of

body lacking; numerous, small and irregularly

arranged central abdominal plates; stripe absent

along first two or three rays of dorsal fin ______________________ 2

1b. Rostrum enlarged; dorsolateral stripe on sides of

body present; few, large central abdominal plates;

stripe present along first two or three rays of dorsal

fin______________________________________________________________________________________________________ 5

2a. More than 60 premaxillary teeth; more than 60

dentary teeth________________________________________________________________________________ 3

2b. Fewer than 50 premaxillary teeth; fewer than 50

dentary teeth _______________________________________________________________________________ 4

3a. Dorsal, pectoral, pelvic, and anal fins without large,

dark blotch occupying almost entire fins; small

spots present on rays; snout rounded_____________________________

__________________________________________ Sturisomatichthys citurensis

Capeti, Tuyra and Bayano rivers, Panama,

and Atrato River basin, Colombia

3b. Dorsal, pectoral, pelvic, and anal fins with large,

dark blotch occupying almost entire fins; small

spots on rays absent; snout triangular ___________________________

__________________________________________ Sturisomatichthys tamanae

Atrato, San Juan, and Baud ´

o rivers,

Pacific slope, Colombia

4a. Twelve branched rays on caudal fin; transverse bars

present dorsally along body; spots absent from

rostrum and head dorsally; large distal spot present

on first branched rays of dorsal fin; short dorsolat-

eral, incomplete stripes absent from caudal pedun-

cle ______________________________________ Sturisomatichthys caquetae

Morelia, Caqueta

´River, Colombia

4b. Ten branched rays on caudal fin; transverse bars

absent dorsally along body; spots present on rostrum

and head dorsally; large distal spot absent from first

branched rays of dorsal fin; short dorsolateral,

incomplete stripes present on caudal peduncle___________

___________ Sturisomatichthys guaitipan, new species

Upper and middle Magdalena River, Colombia

5a. Large, dark blotch absent on pectoral fin; small

spots restricted to rays and/or membranes present

on pectoral fin; discontinuous or inconspicuous

dorsolateral stripe on sides of body __________________________________ 6

5b. Large, dark blotch present on pectoral fin, occupy-

ing or almost occupying entire fin; small spots on

pectoral fin absent; continuous and conspicuous

dorsolateral stripe on sides of body__________________________________ 7

6a. Conspicuous and discontinuous dorsolateral stripe

on sides of body; narrow transverse bar at origin of

dorsal fin, not occupying more than two rows of

plates; interorbital distance 25.4–32.6% in HL __________

____________________________________________ Sturisomatichthys leightoni

Magdalena-Cauca and Orinoco

basins, Colombia

6b. Inconspicuous but continuous dorsolateral stripe on

sides of body; transverse bar at origin of dorsal fin

absent; interorbital distance 32.7–36.7% in HL___________

_________________________________ _____________________ Sturisomatichthys kneri

Maracaibo Lake drainage, Venezuela

7a. Pelvic and anal fins without large, brown blotch

occupying or almost occupying entire fin_____________________ 8

7b. Pelvic and anal fins with large, brown blotch, dark

or inconspicuous, occupying or almost occupying

entire fin________________________________________________________________________________________ 9

8a. Enlarged spots on rays of pelvic and anal fins;

dorsolateral stripe not extending onto caudal pedun-

cle; long filaments on dorsal, pectoral, pelvic, and

anal fins; body depth at dorsal-fin origin 12.0–14.7%

SL; cleithral width 10.3–11.7% SL; head width 65.3–

73.7% HL; interorbital distance 33.3–40.2% HL;

orbital diameter 11.7–16.0% HL; lower-lip width

30.9–34.5% HL _________________ Sturisomatichthys festivus

Maracaibo Lake drainage, Venezuela, and

Catatumbo River, Colombia

8b. Small, almost imperceptible spots on rays of pelvic

and anal fins; dorsolateral stripe extending onto

caudal peduncle; filaments absent on dorsal, pecto-

ral, pelvic, and anal fins; body depth at dorsal-fin

origin 9.6–11.5% SL; cleithral width 8.3–9.6% SL;

head width 50.2–54.3% HL; interorbital distance

25.1–31.6% HL; orbital diameter 9.9–11.3% HL;

lower-lip width 24.6–26.6% HL_________________________________________

__________________ Sturisomatichthys reinae, new species

Baud´

o River, Pacific slope, Colombia

9a. Pectoral fin with stripe extending to almost all rays;

pelvic fin reaching more than first third of anal fin;

dorsal plates semitrapezoidal_____________________________________________ 10

9b. Pectoral fin with stripe restricted to first two to three

rays; pelvic reaching less than frst third of anal fin;

dorsal plates irregular or rectangular_____________________________ 11

10a. Dark stripe on first two or three dorsal-fin rays,

reaching their distal tip and extending to base of

remaining rays; dorsal-fin length 22.2–27.1% SL;

anal-fin length 17.2–20.5% SL; premaxillary teeth

36–50______________________________ Sturisomatichthys frenatus

Esmeraldas, San Javier, Salidero, and Durango

rivers, Ecuador, and Patia River, Colombia,

Pacific slope

10b. Dark stripe on first two or three dorsal-fin rays, not

reaching their distal tip nor extending to base of

remaining rays; dorsal-fin length 28.5–30.8% SL;

anal-fin length 20.8–24.0% SL; premaxillary teeth

54–61 ___________________________ Sturisomatichthys dariensis

Tuyra and Capeti rivers, Pacific slope, Panama

11a. Dark bar at dorsal-fin origin absent; dorsolateral

stripe continuous along its entire length; cleithral

width 7.7–10.5% SL ____________________________________________________________ 12

11b. Broad, dark bar at dorsal-fin origin, occupying

more than two rows of plates, and united to

dorsolateral stripe; dorsolateral stripe discontinu-

ous when entering caudal peduncle; cleithral

width 10.5–12.8% in SL _____________________________________________________

______________________________________________ Sturisomatichthys aureus

Lower and middle Magdalena Basin, Baud ´

o, Cesar,

Sin ´

u, San Jorge, and Rancher´

ıa rivers, Colombia

12a. Dark reticulations absent from predorsal, supraoc-

cipital, and interorbital region; leading ray on

pelvic fins produced, longer than other rays; pelvic

fin reaching or surpassing anterior third of anal fin

_________________________________ Sturisomatichthys panamensis

Aruza, Capeti, and Tuyra rivers, Pacific slope, Panama,

and Atrato, Sin ´

u, and Truando rivers, Caribbean slope,

Colombia

12b. Dark reticulations present on predorsal, supraoc-

cipital, and interorbital region; leading ray on

pelvic fins not produced, same size as other rays;

pelvic fin reaching only origin of anal fin ________________

____________________ Sturisomatichthys varii, new species

San Juan River, Pacific slope, Colombia

Londo˜

no-Burbano and Reis—Taxonomic revision of Sturisomatichthys 803

DATA ACCESSIBILITY

Supplemental material, including the full listing of the

comparative material examined, is available at https://www.

copeiajournal.org/ci-19-226.

ACKNOWLEDGMENTS

We thank Miriam Ghazzi for sending the specimens of

Sturisoma and Sturisomatichthys used in her study. For lending

material under their care and/or hospitality, and technical

assistance to the first author during visits to their museums

and collections, we are very grateful to David C. Werneke and

Jonathan W. Armbruster (AUM), Mark H. Sabaj and John G.

Lundberg (ANSP), James Maclaine (BMNH), Dave Catania

(CAS), Marcela Herna

´ndez Zapata and Luz Fernanda Jimenez

(CIUA), Caleb McMahan and Kevin Swagel (FMNH), Henry

Agudelo and Iva

´n Mojica (ICNMHN), C´

esar Roma

´n-Valencia

and Aura Mar´

ıa Bastidas Quintero (IUQ), Carlos Lucena and

Margarete Lucena (MCP), Raphael Covain and Sonia Fisch-

Muller (MHNG), Marcelo Britto (MNRJ), Marina Mendon¸ca

and Wolmar Wosiacki (MPEG), Alexander Urbano-Bonilla,

Alejandro M´

endez, and Saul Prada (MPUJ), Max Hidalgo

(MUSM), Alessio Datovo (MZUSP), Sven Kullander and Bodil

Kajrup (NRM), Ronald de Rutier (RMNH), Erling Holm and

Herna

´nL

opez-Ferna

´ndez (ROM), Ruth Reina (STRI), Rob

Robins (UF), Juliana Wingert and Luiz Malabarba (UFRGS),

Richard P. Vari, Jeff Clayton, Diane Pitassy, Sandra Raredon,

and Jeff Williams (USNM). Dirk Neumann (ZSM) made

valuable comments on the type of Loricaria aureum. Thanks

are due to L ´

ucia Rapp Py-Daniel (INPA), Tiago Pinto Carvalho

(UFRGS), and Carlos Lucena (MCP) for comments on early

stages of this study. The Pontif´

ıcia Universidade Cat´

olica do

Rio Grande do Sul (PUCRS), Porto Alegre, and the Museu de

Ciˆ

encias e Tecnologia da PUCRS, Porto Alegre (MCP)

provided work space and logistics for the completion of this

study. The first author was partially supported by a

Smithsonian Institution Short Term Visitor Academic Ap-

pointment fellowship, by CNPq through a Ph.D. scholarship

(process #140600/2014-0), and by financial support by

Colciencias. RER is partially financed by the Conselho

Nacional de Desenvolvimento Cient´

ıfico e Tecnol´

ogico

(CNPq, process #306455/2014-5).

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A new species of Sturisoma Swainson, 1838 (Loricariidae: Loricariinae), from the Madeira River basin, with a discussion of historical biogeography of western Amazonas and Paraguay River basins

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Sep 2023
NEOTROP ICHTHYOL

A new species of Loricaria is described from the upper Amazon River basin, Colombia. The new species is distinguished from its congeners primarily by having the dorsal portion of head with uniform black or dark brown coloration extending to three or four plates posterior to dorsal fin base, or with two longitudinal bands from tip of the snout to origin of dorsal fin; abdominal plates tightly joined and completely covering the median abdominal space and pectoral girdle; and pectoral and dorsal fins totally black or dark brown, without bands, spots, or blotches. The new species is further distinguished by plate counts, and body measurements. An analysis of genetic distances using the cytochrome oxidase c subunit 1 marker of the mitochondrial genome showed a clear differentiation between the new species and Loricaria cataphracta (5.8–7.6%), L. nickeriensis (5.7–6.1%), and L. simillima (2.7–7.0%). Species delimitation analyses were carried out, which further supported the new species as a divergent lineage within the genus. Fish species diversity of the upper Amazon River basin and taxonomic issues related to L. simillima are included as part of the discussion.

Estructura y diversidad de las comunidades ícticas a lo largo de un gradiente longitudinal en ríos costeros de la Sierra Nevada de Santa Marta

Article

Full-text available

Aug 2023

En el campo de la ecología de las comunidades ícticas uno de los principales desafíos ha sido entender los cambios longitudinales de la estructura y diversidad de los ensamblajes en ecosistemas loticos, y saber si estos están influenciados por factores abióticos y bióticos. En ese contexto, nos propusimos determinar la estructura y la diversidad de los ensamblajes ícticos en un gradiente longitudinal y su relación con algunas características del hábitat y las coberturas en ríos costeros de la Sierra Nevada de Santa Marta (SNSM). Se analizaron 13 estaciones distribuidas en siete ríos costeros de la SNSM; los peces se recolectaron con un equipo de electropesca en un transecto longitudinal de 100 m de largo, realizando bloqueos con una red de arrastre (5 m x 1,5 m). La estructura y composición se analizó mediante el escalamiento multidimensional no métrico (EMNM) usando el índice de Jaccard y modelos de distribución-abundancia. Para determinar la diversidad se utilizaron los números de Hill (1973) y un análisis de redundancia (AR) para evaluar la relación de las variables ambientales con los ensamblajes. El EMNM y los números de Hill evidenciaron que los ensamblajes estaban estructurados según la distancia hasta la desembocadura en cuatro grupos, tres de ellos ajustados al modelo de Zipf y uno al de preferencia de nicho. El AR evidenció que el patrón longitudinal observado se explicaba por el tipo de sustrato y la cobertura vegetal. Los resultados indicaron que los ensamblajes ícticos parecen obedecer a un gradiente longitudinal, lo que tendría implicaciones importantes para la conservación.

New information on Sturisomatichthys and Sturisoma

Article

Jan 2021

Detailed overview of the genera Sturisoma and Sturisomatichthys, their relationship and distinctive features.

The distribution of Cteniloricaria (Siluriformes: Loricariidae): known and new records in Brazil suggest headwater captures as drivers of disjoint distribution

Article

Full-text available

Jun 2021
NEOTROP ICHTHYOL

Cteniloricaria is a genus of Neotropical armored catfishes belonging to the Loricariinae, currently including two valid species: C. platystoma and C. napova. Cteniloricaria platystoma is presently recorded across the main coastal drainages of the Guiana Shield, from the Sinnamary River, French Guiana, to the Essequibo River basin, Guyana, and is considered to be restricted to the region. Cteniloricaria napova is only known from its type locality at the headwaters of the Paru de Oeste River, Amazon basin, Sipaliwini Savannah, Trio Amerindian territory in Suriname, close to the Brazilian border. Based on a specimen of C. napova, captured in the Cuminapanema River, a tributary to the Curuá River, within Brazilian territory, the geographic distribution of the species and the genus is extended, representing the first record of Cteniloricaria in Brazil. The genus shows a disjoint distribution, and divergence between populations in the north-flowing coastal rivers of the Guianas and the south-flowing Amazon tributaries, and more recent headwater capture between south-flowing Amazon tributaries, may have played a key role in shaping its current distribution. Illustrations, diagnostic characters, morphometrics, description of the habitat where the new specimen was captured, extinction risk assessment, and a discussion of the distribution of the genus are provided.

Peces de la subcuenca del río Catatumbo, cuenca del Lago de Maracaibo, Colombia y Venezuela

Article

Full-text available

Jul 2012

Registros de literatura, complementados con la revisión de ejemplares de colección de referencia de peces del Instituto Humboldt - IAvH-P (Colombia), Estación Biológica de Rancho Grande - EBRG y Museo de Ciencias Naturales de Guanare - MCNG (Venezuela), muestran que 109 especies nativas de agua dulce y 14 de origen marino y estuarino habitan la subcuenca el río Catatumbo, constituyéndose en el sistema con mayor diversidad de especies de peces de la cuenca del Lago de Maracaibo. Estas especies pertenecen a 10 órdenes, 39 familias y 91 géneros. Los órdenes con mayor número de familias, géneros y especies fueron Siluriformes con 12 familias, 42 géneros y 60 especies, y Characiformes 11 familias, 24 géneros y 32 especies. Las familias con mayor riqueza fueron Loricariidae (16 géneros y 25 especies), Characidae (13 géneros y 18 especies) y Pimelodidae (6 géneros y 8 especies); presentando las restantes 37 familias entre una y cinco especies. De las 123 especies registradas, 69 fueron capturadas y catalogadas como resultados de los muestreos durante el presente estudio y 11 corresponden a nuevos registros para Colombia. Para el río Catatumbo se registraron 61 especies de las 68 consideradas endémicas para la cuenca del Lago de Maracaibo y se establecieron como especies pesqueras 40, de las cuales seis fueron marino-estuarinas y 34 dulceacuícolas. Se discute las variaciones del número de especies registradas en los diferentes listados y sus implicaciones en la riqueza de especies para la subcuenca del Catatumbo.

Genera/species by family/subfamily in Eschmeyer's Catalog of Fishes.

Article

Full-text available

Jan 2024

Statistics of numbers of nominal and valid species described in families and subfamilies of recent fishes. Available under http://researcharchive.calacademy.org/research/ichthyology/catalog/SpeciesByFamily.asp

Revision of Loricaria Linnaeus, 1758 (Pisces, Siluriformes, Loricariidae).

Article

Full-text available

Jan 1982

Isaäc J. H. Isbrücker

Diagnose vierzehn neuer Gattungen der Familie Loricariidae Rafinesque, 1815 (Teleostei, Ostariophysi).

Article

Full-text available

Jan 2001

Nomenclator of the 108 genera with 692 species of the mailed catfishes, family Loricariidae Rafinesque, 1815 (Teleostei, Ostariophysi).

Article

Full-text available

Jan 2002

Isaäc J. H. Isbrücker

Classification and catalogue of the mailed Loricariidae (Pisces, Siluriformes)

Article

Full-text available

Jan 1980

Isaäc J. H. Isbrücker

Phylogenomic reappraisal of the Neotropical catfish family Loricariidae (Teleostei: Siluriformes) using ultraconserved elements

Article

Feb 2019

Neotropical freshwaters host more than 6000 fish species, of which 983 are suckermouth armored catfishes of the family Loricariidae – the most-diverse catfish family and fifth most species-rich vertebrate family on Earth. Given their diversity and ubiquitous distribution across many habitat types, loricariids are an excellent system in which to investigate factors that create and maintain Neotropical fish diversity, yet robust phylogenies needed to support such ecological and evolutionary studies are lacking. We sought to buttress the systematic understanding of loricariid catfishes by generating a genome-scale data set (1041 loci, 328,330 bp) for 140 species spanning 75 genera and five of six previously proposed subfamilies. Both maximum likelihood and Bayesian analyses strongly supported the monophyly of Loricariidae. Our results also reinforced the established backbone of loricariid interrelationships: Delturinae as sister to all other analyzed loricariids, with subfamily Rhinelepinae diverging next, followed by Loricariinae sister to Hypostominae + Hypoptopomatinae. Previous DNA-based relationships within Hypostominae and Loricariinae were strongly supported. However, we evaluated for the first time DNA-based relationships among many Hypoptopomatinae genera and found significant differences with this subfamily's current genus-level classification, prompting several taxonomic changes. Finally, we placed our topological results within a fossil-calibrated temporal context indicating that early Loricariidae diversification occurred across the Cretaceous-Paleogene boundary ∼65 million years ago (Ma). Our study lays a strong foundation for future research to focus on relationships among species and the macroevolutionary processes affecting loricariid diversification rates and patterns.

Eschmeyer's catalog of fishes: Genera, species, references.

Book

Feb 2019

Available online under http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp

Catalog of type specimens of recent fishes in the National Museum of Natural History, Smithsonian Institution

Article

Jan 1992

Check list of the Freshwater Fishes of South and Central America

Article

Jan 2003

A Taxonomic Revision of Sturisomatichthys Isbrücker and Nijssen, 1979 (Loricariidae: Loricariinae), with Descriptions of Three New Species

Abstract

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