Copeia, 2007(3), pp.

711–725

A New Schizodon (Characiformes: Anostomidae) from the Rı́o Orinoco

System, with a Redescription of S. isognathus from the Rı́o Paraguay System

BRIAN L. SIDLAUSKAS, JULÍO CESAR GARAVELLO, AND JENNIFER JELLEN

Schizodon scotorhabdotus is described from the Rı́o Orinoco system of Venezuela and
Colombia. Its most similar congener, S. isognathus, is redescribed from the Rı́o
Paraguay system of Brazil, Bolivia, and Paraguay, and a lectotype is designated for the
species. Schizodon scotorhabdotus and S. isognathus can be distinguished from each other
and from the most similar congener, S. knerii, by circumpeduncular and transverse
scale counts, jaw orientation and osteology, and details of coloration. A principal
components-based comparison of S. isognathus, S. knerii, and S. scotorhabdotus revealed
that the three cited species also differ in mean body shape, particularly in body width.
Comments are included on geographic distribution, life coloration, diet, and
ontogenetic variation, including notable shifts in mouth position and caudal-fin
morphology of these species.

Schizodon scotorhabdotus es descrito del sistema del Rı́o Orinoco de Venezuela y

Colombia. Su congener mas similar, S. isognathus, es redescrito del sistema del Rı́o
Paraguay de Brazil, Bolivia, y Paraguay, y un lectotipo de la especie es designado.
Schizodon scotorhabdotus y S. isognathus pueden ser diferenciados uno del otro y del
congener mas similar, S. knerii, por conteos circumpedunculares y de escamas
transversales, orientación y osteologı́a de la mandı́bula, y detalles de coloración. Una
comparación de S. isognathus, S. knerii, y S. scotorhabdotus basada en componentes
principales reveló que las tres especies citadas también difieren en forma promedio del
cuerpo, particularmente en ancho del cuerpo. Se incluyen comentarios sobre
distribución geográfica, coloración en vivo, dieta, y variación ontogenética, incluyendo
notables cambios en la posición de la boca y morfologı́a de la aleta caudal de las
especies.

HE 14 species of herbivorous characiforms mon application of the species-group name

T in the Neotropical genus Schizodon repre-
sent the second most species-rich genus within
the family Anostomidae. Only Leporinus possesses
more species. Though some members of Schizo-
don have been known to science for almost
200 years (Spix and Agassiz, 1829), study of the
genus continues to yield taxonomic discoveries.
Four species of Schizodon have been described in
the last twenty years (Bergmann, 1988; Garavello
and Britski, 1990; Garavello, 1994). Vari and
Raredon (1991) provided compelling evidence
for the distinctiveness of S. corti and elevated it
from the subspecies to species level.
Kner described Schizodon isognathus (1858; with
a more detailed description in 1859) from two
specimens originating in the ‘‘Rio Cujaba’’
(5Rio Cuiabá) of Brazil, a northern tributary of
the Rı́o Paraguay system. In the following century
and a half, this name was applied to specimens
collected in the Rı́o Paraguay system (Pearson,
1937; Géry et al., 1987; Willink et al., 2000), the
Rio São Francisco system of eastern Brazil (Gar-
man, 1890; Borodin, 1931), and the Rı́o Orinoco
system of Venezuela and Colombia (Peters, 1877;
Mago-Leccia, 1970; Taphorn, 1992). The com-
# 2007 by the American Society of Ichthyologists and Herpetologists
isognathus to Schizodon specimens that originated
in three widely separated river systems stemmed
from the common possession of a dark lateral
stripe in at least some of the specimens from
each basin. The distinctiveness of the Schizodon
population inhabiting the Rio São Francisco was
recognized by Steindachner (1875), who de-
scribed it as S. knerii (see also the redescription
in Bergmann, 1988). The purported equivalence
between the Rı́o Orinoco and Rı́o Paraguay
populations persisted (Eigenmann, 1910; Jepsen
and Winemiller, 2002), but was not evaluated
rigorously.
At least three recent authors (Vari and
Raredon, 1991; Antonio, in Taphorn, 1992:63)
have commented on the disjunct distribution of
cited specimens of S. isognathus and have
suggested that the population in the Rı́o Orinoco
may represent an undescribed species. Analysis
of meristic and morphometric characters, as well
as internal and external anatomy confirmed that
the Rı́o Paraguay and Rı́o Orinoco populations
of S. isognathus represent separate species. A
rediscovered syntype of S. isognathus in the NMW
collections matches recent material from the Rı́o
712 COPEIA, 2007, NO. 3
Paraguay system. This contribution describes the
striped form of Schizodon inhabiting the Rı́o
Orinoco system as a new species, restricts the
species-group name isognathus to the population
of striped Schizodon inhabiting the Rı́o Paraguay
system, designates the rediscovered syntype of S.
isognathus as the lectotype of that species and
provides an expanded description. This paper
also presents new observations on the diet and
morphology of S. knerii.
MATERIALS AND METHODS
Meristic counts are as in Sidlauskas and Santos
(2005), with the following change: unbranched
dorsal- and anal-fin-ray counts do not include
one (rarely two) small unbranched ray(s) that lie
under the fleshy covering at the fin origin. These
rays can be observed in radiographs or alizarin
preparations but are difficult to see in whole
specimens. Digital calipers were used to take
eighteen point-to-point measurements from the
left side of each specimen. These measurements
are from Winterbottom (1980), save that the
head length was measured to the posterior bony
margin of the opercle rather than the posterior
fleshy margin. These measurements were con-
verted to their natural logarithms and their
variation was analyzed by principal components
analysis (PCA) using the covariance matrix. The
three specimens in MZUSP 27969, which were
measured by JCG instead of JJ, were excluded
from PCA and subsequent statistical tests in
order to eliminate error associated with multiple
observers. The statistically unique eigenvectors
from the PCA were subsequently analyzed with
ANCOVA (allometric eigenvectors) or ANOVA
(non-allometric eigenvectors) to determine
whether the three species considered in this
analysis differed in shape. Cleared-and-stained
specimens (CS) were prepared according to
Taylor and Van Dyke (1985). Institutional ab-
breviations are as listed in Leviton et al. (1985).
Schizodon scotorhabdotus, new species
Figures 1–4; Tables 1, 2
Schizodon isognathus (not of Kner, 1858). Peters,
1877:472 (Venezuela, Apure).—Eigenmann,
1910:425 (in part, citation, Apuré).—Schultz,
1944:268, 271 (key to species).—Mago-Leccia,
1970:76 (species list, Venezuela).—Taphorn,
1992:63–65, fig. 25, fig. 26 (Venezuela, Apure
basin, diagnosis, diet, migration, suggestion of
undescribed status).—Winemiller and Jepsen,
1998:277 (western Venezuela, Caño Maraca,
diet).—Jepsen and Winemiller, 2002:53 (Ve-
nezuela, Apure).
Unidentified anostomid. Géry, 1977:161 (photo-
graph).
Schizodon sp. Vari and Raredon, 1991 (coloration,
dentition, meristics, suggestion of undescribed
status).
Holotype.—MBUCV-V-32950, 132.0 mm SL, Vene-
zuela, Guárico, Rı́o Aguaro, Rı́o Aguaro N of
Cabruta on cattle ranch on E side of Rı́o Guarico,
07u509280N, 66u309280W, 6 Feb. 1992, A. Ma-
chado and B. Chernoff.
Paratypes.—Colombia: Meta: ANSP 139254, 6,
23.8–38.9 mm SL, Lago Mozambique (on Mo-
zambique ranch) at shoreline on N side of lake,
Rı́o Meta drainage, 03u589N, 73u049W, 23 May
1969, J. E. Böhlke et al.; ANSP 131597, 3, 98.9–
108.1 mm SL, Hacienda Humacita, Caño Angos-
turas at and just below hacienda at Humacitas,
ca. 500 ft of stream, 22 Feb. 1972, J. E. Böhlke et
al.; ANSP 131599, 1, 242.1 mm SL, Quebrada
Venturosa, 1/4 creek mi (0.4 km) above road
between La Balsa and Puerto Lopez, drains into
Rı́o Meta, 04u059N, 72u589W, 15 March 1973, J. E.
Böhlke et al. Venezuela: Anzoategui: ANSP
166745, 3, 115.5–123.3 mm SL, Rı́o Orinoco
basin, Soledad, Lago Tineo, 08u119250N,
63u289200W, 14 March 1987, M. Rodriguez and
S. Richardson; Apure: ANSP 165322, 4, 84.1–
96.8 mm SL, Rı́o Cunaviche, ca. 20 km SW of
Cunaviche on San Fernando de Apure–Puerto
Paez highway, 07u209000N, 67u359000W, 6 Nov.
1989, S. Schaefer et al.; ANSP 165480, 1,
226.0 mm SL, Rı́o Capanaparo, backwater lagoon
(mouth of Caño Las Varitas) near San Fernando
de Apure–Puerto Paez highway, 07u029000N,
67u259000W, 7 Nov. 1989, S. Schaefer et al.;
FMNH 85478, 2, 33.9–49.4 mm SL, Rı́o Orinoco
drainage, 24 river km S of Biruaca on road to San
Juan de Payara? (label reads ‘‘Aparya’’), 7 Jan.
1975, J. E. Thomerson et al.; FMNH 85505, 2,
65.3–108.0 mm SL, Rı́o Orinoco drainage, Rı́o
Aruaca, 32.5 km S of Biruaca, 7 Jan. 1975, J. E.
Thomerson et al.; USNM 257987, 27 of 28
examined, 55.6–194.7 mm SL, 2 CS, 82.8 mm
SL immature female and 94.9 mm SL male, side
channel of Rı́o Apure, ca. 5 km W of San
Fernando de Apure, 07u539N, 67u299W, 21 Jan.
1983, technicians at Apure Fisheries Station;
Barinas: FMNH 104022, 1, 139.7 mm SL, Caño
Las Mulas on the S side of Rı́o Suripá, ca. 5 min
from mouth of Rı́o Caparo, 11 Jan. 1991, B.
Chernoff et al.; FMNH 104023, 3, 106.1–
123.7 mm SL, Caño Socopo, ca. 3.5 hrs upstream
from boat launch of Hato Mercedes in Rı́o
Suripá, 12 Jan. 1991, B. Chernoff et al.; INHS
35172, 2, 126.1–129.3 mm SL, Rı́o Masparro, Rı́o
Apure drainage, 3.5 mi NW of Libertad on road
 SIDLAUSKAS ET AL.—NEW SCHIZODON
Fig. 1. Schizodon scotorhabdotus, MBUCV-V-32950, 132.0 mm SL, holotype, left lateral view; Venezuela,
Guárico, Rı́o Aguaro N of Cabruta on cattle ranch on E side of Rı́o Guarico (07u509280N, 66u309280W).
to Barinas, 08u209180N, 69u409130W, 25 Jan. 1995,
L. Page et al.; Bolı́var: ANSP 135613, 1, 124.1 mm
SL, Caño Chuapo, ca. 20 min downstream from
Jabillal (opposite bank) on Rı́o Caura, 07u079N,
65u009W, 28 Jan. 1977, J. E. Böhlke et al.; Delta
Amacuro: ANSP 182897, ex DU F1152, 1, dry
skeleton, female, 224 mm SL, Rı́o Orinoco on N
shore at Isla Portuguesa, ca. km 402, in Caño
Anabata, 08u379200N, 61u479300W, 15 Nov. 1979,
D. J. Stewart et al.; MBUCV-V-32951, 3 of 11,
92.8–108.6 mm, Caño El Chano–Coporito above
vicinity of Tucupita, Padilla, and La Aguana, 22
Feb. 1978; Guarico: MBUCV-V-32970, 1, dry
skeleton, 271 mm SL, Rı́o Orinoco basin, Rı́o
Guárico system, Rı́o Orituco near Calabozo,
approx. 08u489N, 67u279W, 18 July 2001, W.
Dahdul et al.; FMNH 100121, 2, 77.7–106.6 mm
SL, Caño Las Mercedes near Las Guanotas, ca.
4 km from San Fernando de Apure, 13 Dec.
1988, B. Chernoff et al.; FMNH 100128, 1,
102.8 mm SL, Laguna Los Noreles, Caño Falcon,
11 Dec. 1988, B. Chernoff et al.; FMNH 100129,
1, 108.4 mm SL, ditch between antenna and
Caño Falcon Esteros, 11 Dec. 1988, B. Chernoff
et al.; FMNH 104024, 10, 76.4–112.8 mm SL, 1
CS, 86.7 mm SL male, Rı́o Aguaro N of Cabruta
on cattle ranch on E side of river (collected with
holotype); MZUSP 27969, 3, 83.0 –148.0 mm SL,
caño to W of road from Calabozo to San
Fernando, about 35 km to the S of Masaquaral
farm (Caño Falcon), 20 Jan. 1983, R. Vari et al.;
UMMZ 214743, 12, 127.7–151.1 mm SL, Caño
Falcon, near Camaguan, ca. 25 km N of San
Fernando de Apure, Rı́o Portuguesa drainage, 19
Feb. 1987, W. L. Fink et al.; UMMZ 214836, 1,
266.4 mm SL, Rı́o Caballo at Paso de Caballo,
Rı́o Guariquito drainage, 20 Feb. 1987, W. L.
Fink et al.; Monagas: UMMZ 211297, 4, 87.1–
108.1 mm SL, isolated pools on island, N side of
713
Rı́o Orinoco, 300 km from sea buoy, 08u319N,
62u269W, 10 Nov. 1979, L. Aguana; Portuguesa:
FMNH 96163, 3, 131.6–158.9 mm SL, Rı́o Mario,
tributary of Rı́o Portuguesa, E of Guanare, 26
Feb. 1976, D. C. Taphorn.
Non-type material.—Venezuela: Amazonas: AUM
41515, 1, Rı́o Manipiare at Laja Pelada landing,
27 km SSW of San Juan de Manapiare, 05u129N,
66u109W; AUM 44049, 1, Rı́o Orinoco at Macar-
uru landing, 03u969N, 67u039W.
Diagnosis.—Schizodon scotorhabdotus is distinguish-
able from all other species of Schizodon except S.
isognathus, S. knerii, and S. nasutus by possession of
a dark stripe running along the lateral-line-scale
row from the rear of the opercle to the caudal
peduncle (versus midlateral stripe entirely absent
in S. altoparanae, S. australis, S. borelli, S. corti, S.
dissimilis, S. fasciatus, S. intermedius, S. platae, and S.
rostratus, and midlateral stripe extending no
further anteriorly than to vertical drawn through
dorsal-fin origin in S. jacuiensis and S. vittatus).
Schizodon scotorhabdotus can be distinguished from
S. nasutus by a terminal or slightly upturned mouth
(versus a strongly subterminal mouth). It can be
distinguished from S. isognathus and S. knerii by
possession of 4 or 4.5 transverse scale rows above
the lateral line to the dorsal-fin origin (versus 6.5
scale rows [rarely 5.5 in S. knerii]), 4 or 4.5
transverse scale rows below the lateral line to the
anal-fin origin (versus 5.5 scale rows [rarely 4.5 in
S. knerii]) and seven branched anal-fin rays (versus
eight). It can be further distinguished from S.
isognathus by the presence of three or four vertical
dark blotches on the lateral surfaces of the body
intersecting the prominent lateral stripe (versus
vertical blotches absent) and a terminal mouth in
adults (versus a slightly subterminal mouth in
714 COPEIA, 2007, NO. 3
adults). It can be further distinguished from S.
knerii by having only 16 circumpeduncular scales
(versus 18–20).
Description.—Morphometric characters for holo-
type and paratypes provided in Table 1, meristic
characters provided in Table 2. Body compressed
fusiform, maximum depth averages 1.7 times
maximum width, range 1.3–2.2 in specimens
larger than 50 mm SL. Maximum depth and
width of body both occurring at vertical plane
through dorsal-fin origin. Head triangular in
lateral profile, dorsal slope of skull without
curvature in lateral view. Predorsal region of
body slightly convex; dorsal-fin base slanted
posteroventrally; profile from base of last dor-
sal-fin ray to procurrent rays of caudal fin straight
and sloped slightly posteroventrally. Ventral
margin of body gently convex, with point of
maximum inflection immediately anterior of
insertion of pelvic fin.
Mouth small; upper and lower jaws meet along
horizontal bisecting eye in specimens 90 mm SL
and larger and along horizontal through dorsal
portion of pupil in smaller specimens. Maxilla
slightly convex along ventral margin and widen-
ing posteriorly into small, rhomboidal plate.
Upper lip smooth, lower lip with barely apparent
ridges oriented anteroposteriorly. Premaxilla
and dentary with four asymmetrical, quadricus-
pid teeth. Teeth on each jaw graduated in size
with symphyseal teeth largest. Posterior tooth
much smaller than preceding three teeth. Poste-
rior (fourth) cusp smallest on each tooth, with
third cusp largest. Difference in size between
third and fourth cusp most pronounced on
second tooth of dentary. Single row of replace-
ment teeth embedded in soft tissue on medial
surface of premaxilla. Single row of replacement
teeth in crypt within dentary.
Anterior naris tubular, posterior naris rhom-
boidal with at least nine olfactory ridges visible.
Nares situated on horizontal through dorsal
border of pupil. Nasal large and flat with ossified
sensory canal with two intermediate pores, both
situated on anterior one-half of bone. Antorbital
triangular with long dorsal process. Infraorbital
one lying ventral to antorbital and with L-shaped
sensory canal. Infraorbitals two through five
bordering ventral and posterior margins of eye,
each with portion of sensory canal continuous
with canal of infraorbital one. Infraorbital six
large plate with tripartite sensory canal, lying
over dilatator fossa. Posterior branch of sensory
canal of infraorbital six communicates with canal
system of pterotic. Supraorbital flat, diamond-
shaped, and positioned along anterodorsal mar-
gin of orbit.
Fig. 2. Schizodon scotorhabdotus, ANSP 182897,
224 mm SL, paratype, left lateral view of caudal fin
and hypural plate showing pronounced ramification
and thickening of caudal-fin rays.
Four branchiostegal rays with medial smallest.
Medial three branchiostegal rays narrowing
anteriorly, lateral ray first widening anteriorly
then tapering. Fleshy opercular membrane fused
to branchiostegal rays and isthmus.
Anal fin preceded by one or two very small
additional unbranched rays, visible in radio-
graphs or cleared-and-stained material but not
visible externally and not included in counts.
First externally-visible unbranched ray of anal fin
one-half or less length of second unbranched ray.
Branched rays of anal fin articulated to second
through eighth proximal and distal pterygio-
phores.
Dorsal fin preceded by one very small addi-
tional unbranched ray, visible in radiographs or
cleared-and-stained material but not visible ex-
ternally and not included in counts. First
externally visible unbranched ray of dorsal fin
one-half to two-thirds length of second ray.
Branched rays of dorsal fin articulated to second
through eleventh proximal and distal pterygio-
phores. First dorsal pterygiophore typically lies
between neural processes of ninth and tenth
vertebrae (occasionally between processes of
eighth and ninth).
Shape of expanded pectoral fin triangular,
slightly falcate. Pelvic-fin insertion located at
vertical through base of first branched dorsal-
fin ray. Adipose-fin origin located on vertical
through base of third or fourth branched anal-fin
ray. Adipose fin shaped like half-moon. Profile of
caudal-fin lobes blunt and noticeably rounded.
Specimens over ca. 80 mm SL with first six or
seven dorsal caudal-fin rays and last six or seven
ventral caudal-fin rays repeatedly ramified,
broadened, and joined into thickened plates
(Fig. 2). Individual rays within caudal fin may
 SIDLAUSKAS ET AL.—NEW SCHIZODON
divide into as many as nine branches before
reaching distal margin of fin.
Body fully scaled. Fins not scaled except for
basal portion of caudal fin. Scales extend to
posterior margin of hypural plate and onto basal
portion of middle caudal-fin rays.
Coloration in alcohol.—Overall ground color
brown; background coloration darkest above
lateral line between opercle and dorsal fin,
lightest below lateral line, especially on caudal
peduncle. Dorsal surface of head dusky; ventral
surface of head, gular region, and branchiostegal
membrane pale. Distinct, dark, slightly poster-
odorsally-slanted midlateral stripe overlying ven-
tral one-half of lateral-line-scale row and dorsal
one-half of immediately ventral scale row. Stripe
shifting slightly dorsally at approximately twenty-
first lateral-line scale to include entire lateral-
line-scale row and ventral portion of scale row
immediately dorsal to lateral-line-scale row. Dark
stripe continuous anteriorly with horizontal
stripe on opercle and fourth and fifth infraorbi-
tal. Fleshy flap of opercle unpigmented. Lateral
surface of body with three to five dark, vertical
blotches intersecting lateral stripe in most speci-
mens. First blotch located just posterior to
opercle and covering lateral-line scales 4–6.
Second blotch situated ventral to dorsal fin and
covering lateral-line scales 11–15. Third through
fifth blotches highly variable in size and intensity
of pigmentation, but frequently with one or two
blotches located between verticals through dorsal
and adipose fins and one ventral to adipose fin.
Intensity of pigmentation of all vertical blotches
varies considerably among specimens of all sizes.
Lateral surface of body dorsal to lateral line with
four or five faint dark lateral stripes running
along dorsal and ventral portions of scale rows.
Lateral stripes most conspicuous between dorsal
and adipose fins. Melanophores scattered on
membranes between dorsal-fin rays. Other fins
with occasional melanophores outlining rays.
Some specimens with dark stripe on middle
caudal-fin rays continuous with dark midlateral
stripe on body. Adipose fin dusky with irregular
pattern of melanophores.
Coloration in life.—Color photo of living S.
scotorhabdotus appears in Géry (1977:161), labeled
as ‘‘Unidentified anostomid from Venezuela,
possibly of the genus Laemolyta’’. Though lateral
stripe of specimen in photo strongly resembles
stripe possessed by Laemolyta taeniata and L.
orinocensis, both of which are also found in
Orinoco basin, overall coloration, squamation,
and body shape of specimen clearly match S.
715
scotorhabdotus. Ground coloration silver; portion
above lateral line darkest; lateral stripe very dark,
almost black; vertical blotches posterior to
opercle and ventral to dorsal fin inconspicuous
but visible; dorsal and ventral margins of scale
rows above lateral line darker, creating appear-
ance of three jagged longitudinal lines along
dorsal one-half of body; fins hyaline. Coloration
in photograph of large (271 mm SL) specimen
taken just after death by W. Dahdul (MBUCV-V-
32970) largely agrees with description above,
except in that specimen four lateral blotches
darker and more distinct than lateral stripe.
Growth and ontogeny.—Position of mouth shifts
from upturned in young juveniles to terminal in
adults (Fig. 3). Specimen of 30.8 mm SL (ANSP
139254) has strongly upturned mouth reminis-
cent of adults of Laemolyta (Fig. 3A). In specimen
of 38.9 mm SL from same lot, mouth still
upturned but beginning transition to adult
terminal orientation (Fig. 3B). By 49.4 mm SL
(FMNH 85478), transition nearly complete
(Fig. 3C). Specimen of 88.4 mm SL (ANSP
165322) with fully adult condition (Fig. 3D).
Number of rami in principal rays of caudal fin
increasing considerably during ontogeny. Juve-
niles 38.9 mm SL (ANSP 139254) and smaller
have unramified, delicate caudal fins relative to
adult condition. In 49.4 mm SL specimen lon-
gest rays of caudal fin beginning to increase in
number of rami. By 88.4 mm SL, longest six or
seven rays in each caudal-fin lobe broadened,
increased greatly in number of rami (to as many
as nine), and joined tightly into semi-rigid plate.
At this stage caudal-fin lobes become truncate,
with rounded distal margins. All examined
specimens larger than 88.4 mm SL have highly
ramified, plate-like morphology of caudal fin
(Fig. 2).
Coloration largely constant over ontogeny,
with even smallest examined specimens exhibit-
ing dark lateral stripe on body. Contrary to adult
condition, smaller specimens have lateral stripe
continuing anterior of orbit onto snout and dark
anterior margin of adipose fin. Very small
juveniles of ca. 24–40 mm SL (ANSP 139254)
lack vertical blotches and caudal-fin stripe.
Vertical blotches and caudal-fin stripe vary in
intensity across all later stages of ontogeny.
Distribution.—Commonly occurring throughout
lowlands of Orinoco drainage, including rı́os
Aguaro, Apure, Aruapa, Caura, Meta, Suripá, and
Portuguesa (Fig. 4). Known from as far west as
Quebrada Venturosa in Rı́o Meta drainage in
central Colombia (ANSP 131599) and as far east
as Rı́o Orinoco delta (MBUCV-V-32951). No
716 COPEIA, 2007, NO. 3

Fig. 3. Ontogenetic change in mouth position among paratypes of Schizodon scotorhabdotus; specimens
smaller than 40 mm SL (A, ANSP 139254, 30.8 mm SL; B, ANSP 139254, 38.9 mm SL) with upturned
mouths; specimens of 90 mm SL and larger (D, ANSP 165322, 88.4 mm SL) with fully terminal mouths and
specimens between 40 and 90 mm SL (C, FMNH 85478, 49.4 mm SL) with mouths in intermediate position.

types originated further upriver on Rı́o Orinoco
than confluence with Rı́o Meta, though two non-
type specimens (AUM 41515 and AUM 44049
from rı́os Maniapare and Orinoco, respectively)
indicate that range also includes upriver portions
of Orinoco drainage in foothills of Guiana
Shield.
Diet and gut morphology.—Cited as herbivorous in
accounts from Venezuela (Taphorn, 1992; Jep-
sen and Winemiller, 2002). Winemiller and
Jepsen (1998) stated that macrophytes are pre-
ferred. Some specimens caught near cattle
pasture had consumed manure (Taphorn,
1992). Stomachs of cleared-and-stained speci-
mens contained large quantities of macrophytes,
small amount of filamentous algae and signifi-
cant amount of sand. Several large worms with
thick white integument found undigested in
lower intestines of all three CS specimens; these
appeared too large for fish to have swallowed at
present size; most likely these were parasitic.
Stomach connected to approximately twenty
pyloric caeca. Length of intestine proper 1.14
times SL; length of complete gastric tract in-
cluding stomach 1.55 times SL.
Habitat.—Most examined specimens of S. scoto-
rhabdotus originated in lagoons, pools, small
caños, and cattle pastures, all relatively lentic
environments. Apparent preference for lentic
habitats possibly sampling artifact, but dietary
preference for macrophytes that grow in slow,
shallow water, suggests that lentic habitat prefer-
ence is real. Taphorn (1992) listed species as
inhabiting lentic and lotic environments and as
accompanying prochilodontids on annual migra-
tions. As all but one of available specimens
collected during dry season (November–March),
preference for lentic environments may be
seasonal.
Etymology.—The species-group name, scotorhabdo-
tus, is derived from the Greek scotos meaning
dark, and rhabdos meaning rod or stick, in
reference to the prominent dark lateral stripe
that characterizes the species. An adjective in the
nominative singular.
 SIDLAUSKAS ET AL.—NEW SCHIZODON
Fig. 4. Distribution map for Schizodon isognathus
(black squares), S. knerii (open squares), and S.
scotorhabdotus (black dots). Holotype locality for S.
scotorhabdotus represented by a black star.
Schizodon isognathus Kner, 1858
Figures 4, 5; Tables 1, 2
Schizodon isognathus Kner, 1858:78 (original de-
scription and diagnosis, Cujaba 5 Rio
Cuiabá).—Kner, 1859:163 (more detailed
description).—Eigenmann et al., 1907:150
(species listing, Paraguay).—Eigenmann,
1910:425 (in part, citation, Brazil, Cu-
jaba).—Pearson, 1937:110 (Paraguay, species
list).—Bergmann, 1988:14 (Brazil, Rio Para-
guai).—Willink et al., 2000:291 (species list,
Pantanal of Brazil).
Anostomus isognathus. Günther, 1864:305 (change
in generic assignment, citation based on Kner,
1859).
Schizodon isognathum. Géry, 1977:154–155 (Rı́o
Paraguay basin).—Géry et al., 1987:375, 380–
381 (key, coloration, some meristics and
morphometrics, Rı́o Paraguay).
Lectotype.—NMW 58855, 1, approximately
234 mm SL, Brazil, Rio Cujaba (5Rio Cuiabá);
designated herein based on examination of
photos.
717
Non-type material.—Brazil: MZUSP 88603, 1 CS,
176.0 mm SL, female, Mato Grosso, Santo Antô-
nio do Leverger, Rio Cuiabá; ANSP 53825, 1,
234.1 mm SL, Mato Grosso, Descalvados; FMNH
59508, 5, 106.4–201.7 mm SL, San Luis de
Caceres, Mato Grosso. Brazil–Bolivia: FMNH
92292, 1, 82.8 mm SL, Bastos, Rio Alegre (Rio
Jauru) into Rı́o Guapore. Paraguay: FMNH 69725,
1, 50.6 mm SL, Villa Hays; UMMZ 206792, 2, 235.8
and 240.8 mm SL, Rı́o Apa, ca. 0.5 km upstream
(5E) of bridge between Brazil and Paraguay in
Bella Vista; UMMZ 207809, 1, 151.1 mm SL, Rı́o
Aquidaban at Paso Horqueta circa 24 km NNW of
Loreto; UMMZ 207885, 1, 129.3 mm SL, swamp
pond adjacent to Rı́o Aquidaban at Paso Hor-
queta, ca. 24 km NNW of Loreto.
Diagnosis.—Schizodon isognathus is distinguished
from all other species of Schizodon except S. knerii,
S. nasutus, and S. scotorhabdotus by a midlateral
stripe running along the lateral-line-scale row
from the opercle to the caudal peduncle (versus
midlateral stripe entirely absent in S. altoparanae,
S. australis, S. borelli, S. corti, S. dissimilis, S.
fasciatus, S. intermedius, S. platae, and S. rostratus,
and midlateral stripe extending no further
anteriorly than vertical through dorsal-fin origin
in S. jacuiensis and S. vittatus). It can be
distinguished from S. nasutus by a slightly sub-
terminal mouth (versus a strongly subterminal
mouth in S. nasutus). Schizodon isognathus is
distinguished from S. knerii and S. scotorhabdotus
by lack of vertical dark blotches on the lateral
surfaces of the body intersecting the midlateral
stripe (versus two to five vertical blotches typically
present). Schizodon isognathus is further distin-
guished from S. knerii by having only 16
circumpeduncular scales (versus 18–20 scales)
and prominent dark pigmentation of the lateral
stripe along its entire length (versus lateral stripe
faint and incomplete anteriorly in most speci-
mens of S. knerii). Schizodon isognathus is further
distinguished from S. scotorhabdotus by possession
of 6 to 6.5 scale rows above the lateral line (versus
4 to 4.5 rows), 5.5 scale rows below the lateral line
(versus 4 to 4.5 rows), eight branched anal-fin
rays (versus seven rays), and a slightly subtermi-
nal mouth in adults (versus a terminal mouth in
adults).
Description.—Morphometric characters presented
in Table 1, meristic characters presented in
Table 2. Body compressed fusiform, maximum
depth averages 2.1 times maximum width, range
1.7–2.6. Greatest depth and width of body at
dorsal-fin origin. Head triangular in lateral pro-
file, dorsal slope of skull without curvature or
very slightly concave in lateral view. Predorsal
718 COPEIA, 2007, NO. 3

TABLE 1. MORPHOMETRIC CHARACTERS OF Schizodon scotorhabdotus (n 5 97), S. isognathus (n 5 13), AND S. knerii (n 5 12).
Bent, damaged, or abnormal specimens (e.g., a specimen lacking an adipose fin) were not measured. HT 5
holotype. Standard lengths and statistics are given in millimeters (SD 5 standard deviation). Measurements 1–11
and statistics expressed as percentages of standard length, 12–17 as percentages of head length. Lectotypes of S.
isognathus and S. knerii unavailable for measurement.

S. scotorhabdotus S. isognathus S. knerii

Measurement HT Mean SD Range Mean SD Range Mean SD Range

Standard length 132.0 108.4 43.1 23.8–266.4 158.6 59.8 50.6–240.8 143.2 37.9 81.6–200.9
1. Lower jaw to anal-fin origin 82 83 2 77–86 81 1 80–83 81 2 79–84
2. Lower jaw to adipose-fin 85 86 1 83–89 87 1 86–89 85 2 83–88
origin
3. Lower jaw to dorsal-fin origin 45 46 2 42–52 46 1 45–50 48 1 45–50
4. Lower jaw to pelvic-fin origin 47 48 2 41–52 49 1 47–52 49 1 47–51
5. Dorsal- fin origin to caudal- 59 59 2 51–64 58 2 54–61 59 1 57–61
fin origin
6. Dorsal-fin origin to adipose- 43 44 2 38–48 44 1 41–46 42 2 39–45
fin origin
7. Caudal peduncle length 12 13 1 09–16 13 1 10–15 14 1 11–16
8. Body depth 25 26 2 22–31 27 2 25–30 30 1 27–33
9. Body width 16 15 2 08–22 13 2 11–15 15 2 13–19
10. Caudal peduncle depth 10 10 1 08–11 11 0 10–12 12 1 11–13
11. Head length 27 27 2 22–34 24 3 20–30 25 2 23–28
12. Lower jaw to preopercle 72 76 4 70–95 74 2 69–76 75 2 73–80
posterior margin
13. Snout length 37 37 3 29–45 37 3 27–40 39 3 32–45
14. Head depth 76 81 6 64–95 85 7 71–98 91 7 81–104
15. Snout depth 41 43 5 26–55 44 4 38–52 49 5 38–60
16. Eye diameter 29 31 4 24–41 30 3 24–36 30 4 24–35
17. Bony interorbital width 47 47 5 31–55 42 3 33–47 45 3 39–48

profile of body gently convex; base of dorsal fin
slanted posteroventrally; dorsal profile of body
between base of last dorsal-fin ray and procurrent
rays of caudal fin slightly convex to straight with
posteroventral slant. Ventral margin of body
gently convex, with point of maximum inflection
immediately anterior of insertion of pelvic fins.
Mouth small and very slightly subterminal;
upper and lower jaws meet along horizontal
running tangent to ventral margin of pupil.

TABLE 2. MERISTIC CHARACTERS AND STATISTICS FOR Schizodon scotorhabdotus (n 5 93), S. isognathus (n 5 14), AND S. knerii
(n 5 13). HT 5 holotype, LT 5 lectotype, SD 5 standard deviation. Incomplete series of small scales at bases of
dorsal and anal fins counted as 0.5 scales in transverse counts. Data for the lectotype of S. isognathus taken from
photos. Data for the lectotype of S. knerii taken from Bergmann (1988).

S. scotorhabdotus S. isognathus S. knerii

Count HT Mode SD Range LT Mode SD Range LT Mode SD Range

Unbranched dorsal-fin rays 2 2 0 2 2 2 0 2 2 2 0 2

Branched dorsal-fin rays 10 10 0.20 9–10 10 10 0 10 11 10 0.38 10–11
Unbranched anal-fin rays 2 2 0 2 2 2 0 2 2 2 0 2
Branched anal-fin rays 7 7 0 7 8 8 0 8 8 8 0.38 7–8
Pectoral-fin rays 17 17 0.54 15–18 ? 17 0.73 16–18 17 16 0.83 16–18
Pelvic-fin rays 9 9 0 9 ? 9 0 9 9 9 0.28 8–9
Branchiostegal rays 4 4 0 4 ? 4 0 4 4 4 0 4
Total vertebrae 41 41 0.55 40–42 ? 41 0.41 40–42 ? 40 0.39 39–40
Lateral-line scales 45 43 0.96 42–46 43 45 1.22 42–46 43 44 0.65 42–44
Scales above lateral line 4.5 4.5 0.07 4 or 4.5 6 6.5 0.13 6 or 6.5 6 6.5 0.50 5.5 or 6.5
Scales below lateral line 4.5 4.5 0.24 4 or 4.5 5 5.5 0.13 5.5 5 5.5 0.43 4.5 or 5.5
Predorsal scales 11 11 0.68 9–12 ? 12 0.73 11–13 12 11 0.52 11–12
Circumpeduncular scales 16 16 0 16 ? 16 0 16 ? 20 0.90 18–20
 SIDLAUSKAS ET AL.—NEW SCHIZODON
Maxilla slightly curved along ventral margin and
widening posteriorly into small, rhomboidal
plate. Upper lip smooth; lower lip with poorly
developed ridges oriented anteroposteriorly.
Upper and lower jaws with four teeth on each
side. Teeth graduated in size with symphyseal
tooth on each side largest and fourth much
smaller than other three teeth. All teeth multi-
cuspid with serrated edges; typically four cusps
on each tooth. Third tooth of dentary with two
cusps closest to symphysis much larger than
other two cusps, resulting in distinct slope and
notch of distal margin of tooth. Single row of
replacement teeth embedded in soft tissue on
medial surface of premaxilla. Single row of
replacement teeth in crypt within dentary.
Anterior naris tubular, posterior naris rhom-
boidal and revealing at least eight olfactory
ridges. Nares lie just ventral to horizontal
through dorsal margin of pupil. Nasal large
and flat with ossified sensory canal with two
intermediate pores, both situated on anterior
one-half of bone. Antorbital triangular with long
dorsal process. Infraorbital one lying ventral to
antorbital and with L-shaped sensory canal.
Infraorbitals two through five bordering ventral
and posterior margins of eye, each with portion
of sensory canal continuous with canal of
infraorbital one. Infraorbital two with small,
triangular, dorsomedially-directed flange at an-
terior extreme of bone, flange thickened on
medial surface into small bump. Infraorbital six
with tripartite sensory canal, infraorbital over-
lying dilatator fossa. Posterior branch of sensory
canal of infraorbital six communicates with
canal system of pterotic. Supraorbital flat, di-
amond-shaped, and positioned along antero-
dorsal margin of orbit. Small notch along
anteroventral margin of supraorbital. Four
branchiostegal rays, medialmost ray smallest.
Fleshy opercular membrane fused to branchios-
tegal rays and isthmus.
Very small unbranched ray preceding anal fin,
visible in radiographs or cleared-and-stained
material but not visible externally and not
included in counts. First externally visible un-
branched ray of anal fin one-half to two-thirds
length of second unbranched ray. Branched
anal-fin rays articulated to second through ninth
proximal and distal pterygiophores. First
branched anal-fin ray slightly longer than last
unbranched ray.
Very small unbranched ray preceding dorsal
fin, visible in radiographs or cleared-and-stained
material but not visible externally and not
included in counts. First externally visible un-
branched ray of dorsal fin one-half to two-thirds
length of second unbranched ray. Branched rays
719
of dorsal fin articulated to second through
eleventh proximal and distal pterygiophores.
First branched dorsal-fin ray slightly longer than
last unbranched dorsal-fin ray. First dorsal
pterygiophore lies between neural processes of
tenth and eleventh vertebrae.
Shape of extended pectoral fin triangular,
somewhat falcate. Pelvic-fin insertion located at
vertical through base of first or second branched
dorsal-fin ray. Adipose-fin origin located along
vertical through base of fifth or sixth branched
anal-fin ray. Adipose fin shaped like half-moon.
Lobes of caudal fin moderately falcate to truncate.
In specimens over 130 mm SL, first six or seven
dorsal caudal-fin rays and last six or seven ventral
caudal-fin rays heavily ramified, broadened, and
joined into thickened plates. Greatest degree of
ramification and broadening of rays observed in
largest individuals (ca. 230 mm SL). Individual
rays within caudal fin of large adults may divide
into as many as nine branches before approaching
distal margin of fin.
Body fully scaled. Fins not scaled except for
base of caudal fin where scales extend just
posterior of margin of hypural plate.
Coloration in alcohol.—Overall ground color tan to
brown, darker dorsal to lateral line. Dorsal
surface of head dusky; ventral surface of head
pale, particularly in gular region and membrane
covering branchiostegal rays. Distinct, dark,
midlateral stripe very slightly posterodorsally
slanted. Dark stripe overlies ventral one-half of
lateral-line-scale row, all of scale row immediately
ventral and dorsal portion of next ventral scale
row in area anterior of vertical through dorsal-fin
insertion. Stripe expands to include most or all
of lateral-line-scale row in region between verti-
cals through dorsal-fin insertion and anal-fin
origin. Lateral stripe shifts dorsally posterior of
adipose-fin insertion to include ventral half of
scale row immediately dorsal to lateral-line-scale
row but no longer include ventral half of scale
row immediately ventral to lateral-line-scale row.
Stripe continues posteriorly onto middle rays of
caudal fin. Dark lateral stripe continuous anteri-
orly with horizontal stripe on opercle and
infraorbitals four and five. Fleshy flap of opercle
unpigmented. Lateral surface of body dorsal to
lateral line with five or six faint dark lateral
stripes along dorsal portion of each scale row. In
some specimens, darkest pigmentation runs
along upper and lower margins of each scale,
giving impression of five horizontal zigzag lines
on each lateral surface of body. Melanophores
scattered over membranes between dorsal-fin
rays. Caudal fin almost entirely hyaline except
for dark stripe on middle fin rays. Distal margin
720 COPEIA, 2007, NO. 3
of adipose fin dusky. Pelvic, pectoral, and anal
fins hyaline.
Coloration in life.—Géry et al. (1987) wrote that
life coloration is ‘‘silver, with black longitudinal
band and zigzag lines, a yellow spot in the dorsal
half of the eye, the pectoral, pelvic and anal fins
yellow, a red caudal fin (during the spawning
period?)’’ (translation ours). Chernoff (pers.
comm., 2006, based on field notes) indicates
that dorsal half of eye yellow or orange.
Growth and ontogeny.—Position of mouth in very
young juveniles unknown. Mouth position shifts
ontogenetically from terminal in juveniles to
slightly subterminal in adults. Specimens of
50.6 mm SL (FMNH 69725) and 82.8 mm SL
(FMNH 92292) with mouth in terminal position;
specimen of 129.3 mm SL (UMMZ 207885) and
individuals of larger sizes with mouth in sub-
terminal position.
Caudal fin moderately falcate in juveniles and
young adults, more truncate in large adults.
Ramification and broadening of caudal-fin rays
increases ontogenetically. Specimen of 50.6 mm
SL with no noticeably increased ramification and
broadening of rays. Caudal-fin rays with limited
ramification at 82.8 mm SL, with noticeably
increased ramification at 129.3 mm SL, and with
greatest degree of thickening and ramification at
240.8 mm SL (UMMZ 206792).
Coloration apparently stable across post-larval
ontogeny, with lateral stripe on body well de-
veloped in smallest specimen examined
(50.6 mm SL; FMNH 69725). Coloration of
larvae and very young juveniles unknown.
Distribution.—Found throughout Rı́o Paraguay
system of Brazil, Bolivia, and Paraguay (Fig. 4).
Material examined comes from as far north as
Pantanal of Brazil in vicinity of Santo Antônio do
Leverger in extreme southwest Mato Grosso state
and as far south as Concepción and Loreto in
central Paraguay.
Diet and gut morphology.—Intestine exclusive of
stomach 1.44 times SL. Intestinal tract including
stomach 1.64 times SL. Posterior portion of
stomach thick walled. Pyloric caeca 14. Stomach
of cleared-and-stained specimen empty. Based on
similarity in dentition and gut morphology to S.
scotorhabdotus, S. isognathus presumably herbivo-
rous.
Lectotype designation for Schizodon isognathus.—A
series of photographs of a syntype of Schizodon
isognathus (NMW 58855), provided by H. Well-
endorf, was examined during this study. That
specimen (Fig. 5A), herein designated as the
lectotype of the species, is undoubtedly one of
the two specimens that Kner (1858, 1859) used in
the original description of the species. The
locality (painted on the specimen), meristics,
and external morphology match the data cited
for the syntypes and also match more recently
collected material that originated in the Rı́o
Paraguay system (Fig. 5, Table 2). The NMW
specimen measures approximately 260 mm TL
(510.24 inches) to the end of the damaged
caudal fin, a length which corresponds closely to
the published 10.5 inches for the smaller of the
two type specimens (Kner, 1859). Furthermore,
the original description indicates that the S.
isognathus syntypes originated with Natterer, are
dried and are kept in the ‘‘Imperial Museum.’’
The lectotype is a dried skin discovered in NMW,
which houses most of the other Kner types based
on Natterer’s collections (Vari, 1984). The other
syntype was not found during a search of the
NMW.
Additional notes on Schizodon knerii.—For com-
parative purposes and as a supplement to
Bergmann’s (1988) redescription of Schizodon
knerii, we present photos of the lectotype (NMW
62840:2; Fig. 6A) and a recently collected spec-
imen (Fig. 6B) of the species and provide some
additional observations. When he designated the
S. knerii lectotype, Bergmann (1988) cited the lot
number incorrectly as NMW 26840:2 (H. Well-
endorf, pers. comm., 2005).
Comparison of the color pattern of the
lectotype of Schizodon knerii (Fig. 6A) with that
of a recent specimen of the species (Fig. 6B)
reveals the variation in intensity of the midlateral
stripe that led previous researchers to cite
specimens of S. knerii as S. isognathus (Borodin,
1931:44). Most examined specimens possess
three or four faint blotches on the lateral
surfaces of the body and a faint lateral stripe
that darkens considerably over the caudal
peduncle to form a dark spot over the base of
the middle rays of the caudal fin (Fig. 6B). In
other specimens, including the lectotype
(Fig. 6A), the stripe is absent and only the
caudal spot remains.
Examination of the stomach of one of the
cleared-and-stained specimens of Schizodon knerii
(ANSP 171830) revealed large quantities of
shredded macrophytes and several large parasitic
worms. There were 19 pyloric caeca and an
overall gut morphology similar to that of S.
isognathus. The stomachs of the other two
cleared-and-stained specimens (MZUSP 88602)
contained large quantities of silt, suggesting that
detritus or algae may also form part of the diet.
 SIDLAUSKAS ET AL.—NEW SCHIZODON 721

Fig. 5. (A) Schizodon isognathus, NMW 58855, 234 mm SL, lectotype, right lateral view; Brazil, Rio Cuiaba;
right side of specimen presented to show painted collection number and locality information on tag pasted
to specimen; photo by H. Wellendorf. (B) S. isognathus, UMMZ 207885, 129.3 mm SL; left lateral view of
more recently collected specimen; Paraguay, swamp pond adjacent to Rı́o Aquidaban at Paso Horqueta,
circa 24 km NNW of Loreto.

The silt may also have been consumed inciden-
tally to eating macrophytes.
The larger CS MZUSP specimen of Schizodon
knerii (179.9 mm SL) is an adult female with very
ripe ovaries. This specimen was captured on 5
November near Três Marias in the upper São
Francisco, suggesting that the spawning season in
that region includes the spring.
Morphometric comparison.—The three examined
species of Schizodon possess very similar body
shapes, with considerable overlap among species
in the ranges of all measured body proportions
(Table 1). The species means differ slightly for
several measurements including head and snout
depth, body width, the width of the bony
interorbital, and the distance from the tip of
the lower jaw to the dorsal-fin origin (Table 1).
These differences suggest that mean body shape
of each species differs, despite the overall overlap
in morphometrics. Principal components anal-
ysis was used to further examine the morpho-
metric distinctiveness of these species and
returned two statistically distinct principal com-
ponents (Anderson’s test of eigenvalue equiva-
lence, P , 0.05). The first of these components
(PC1) correlates perfectly with standard length
(correlation coefficient r 5 20.998) and has
loadings of equal sign and approximately equal
magnitude for all 18 linear measures (Table 3).
Principal component 1 can be interpreted as
a loose index of the scale (size) of the speci-
mens, though because the loadings are not all
identical it is more accurate to state that PC1
describes the allometric trajectory of body
shape. The ANCOVA of the PC1 scores for each
species with SL as the covariate reveals that the
regressions for all three species have statistically
indistinguishable intercepts and slopes (P ..
0.05). The similarity of the regressions indicates
722 COPEIA, 2007, NO. 3

Fig. 6. (A) Schizodon knerii, NMW 62840:2, 235.0 mm SL, lectotype, left lateral view; Brazil, Rio São
Francisco; photo by H. Wellendorf; (B) S. knerii, ANSP 171830, 125.2 mm SL, left lateral view of more
recently collected specimen; Brazil, Minas Gerais, Rio Salinas, tributary of Rio Verde Grande, 51.0 km WSW
from Monte Azul on road to Jaı́ba.

that the growth allometry for all three species is
nearly identical.
Principal component 2 does not correlate
significantly with standard length (r 5 0.043)
and represents shape variation that is indepen-
dent of the scale or growth allometry of the
specimens. The strongest influences on PC2
scores are two measures of overall width (body
width and bony interorbital distance; Table 3). A
scatter of PC2 scores against PC1 scores (Fig. 7)
illustrates that while there is considerable overlap
in the range of PC2 scores for members of all
three species, the mean PC2 scores for each
species differ. Pairwise ANOVA for all three
between-species comparisons reveals that each
species possesses a unique mean PC2 score (P ,
0.05 in all three comparisons). Schizodon iso-
gnathus tends to have a narrower body and
shorter interorbital distance than do the other
two species, while S. scotorhabdotus tends to have
a wider body and longer interorbital distance
than do the other two species. While the overlap
of the range of PC2 scores means that morpho-
metrics cannot fully diagnose individual speci-
mens to species, the difference in PC2 means
confirms that the body morphologies of mem-
bers of each species are drawn from statistically
different distributions. The morphometric anal-
ysis upholds the recognition of all three species
as distinct.
Osteological comparison.—Schizodon scotorhabdotus
further differs from S. isognathus and S. knerii in
the morphology of the bones of the lower jaw
(Fig. 8). In Schizodon scotorhabdotus, the dentary is
a triangular bone that does not extend signifi-
cantly ventral to the ventral margin of the
retroarticular and anguloarticular (Fig. 8B).
Conversely, in S. isognathus (Fig. 8A) and S. knerii
the dentary is a rhomboidal bone that extends
significantly ventral to the ventral margin of the
retroarticular and anguloarticular. The latero-
sensory canal segment of the dentary runs nearly
horizontally in S. scotorhabdotus but exhibits an
 SIDLAUSKAS ET AL.—NEW SCHIZODON 723

TABLE 3. PRINCIPAL COMPONENT EIGENVALUES AND

LOADINGS CALCULATED FROM 18 LINEAR MEASUREMENTS
TAKEN FROM Schizodon scotorhabdotus (n 5 94), S.
isognathus (n 5 13), AND S. knerii (n 5 12). Loadings
with absolute magnitude greater than 0.3 on second
eigenvector shown in bold.

PC1 PC2
Eigenvalue 3.874 0.019
% Eigenvalue 98.209% 0.478%

Standard length 20.240 0.147

Lower jaw to anal-fin origin 20.243 0.079
Lower jaw to adipose-fin origin 20.241 0.139
Lower jaw to dorsal-fin origin 20.230 0.128
Lower jaw to pelvic-fin origin 20.237 0.189
Dorsal-fin origin to caudal-fin origin 20.248 0.121
Dorsal-fin origin to adipose-fin origin 20.251 0.119
Caudal peduncle length 20.259 0.187
Body depth 20.234 0.265
Body width 20.286 20.656
Caudal peduncle depth 20.256 0.257
Head length 20.200 20.038
Lower jaw to preopercle posterior 20.198 20.109
margin
Snout length 20.232 20.123
Head depth 20.225 0.082
Snout depth 20.239 20.178
Eye diameter 20.163 20.242
Bony interorbital width 20.234 20.380 Fig. 8. Lower jaw, right side, lateral view of (A)
Schizodon isognathus, MZUSP 88603, 176.0 mm SL
and (B) S. scotorhabdotus, FMNH 104024, 86.7 mm
SL, paratype. Images reversed to place anterior to
left. Dentition not illustrated.

anteroventral slope in S. knerii and S. isognathus.

The midlength pore of the sensory canal of the
dentary is positioned closer to the anterior pore
of the sensory canal in S. knerii and S. isognathus
than in S. scotorhabdotus, and the ascending
process of the anguloarticular is narrower in S.
scotorhabdotus than it is in S. isognathus and S.
knerii. The osteological differences in the lower
jaw support the recognition of S. scotorhabdotus as
a species separate from S. isognathus and S. knerii.

MATERIAL EXAMINED

Schizodon nasutus. FMNH 112960, ex USNM

Fig. 7. Scatter of scores on second principal
component (PC2) from linear morphometric anal-
ysis against first principal component (PC1) from
the same. Symbols represent Schizodon isognathus
(black squares; n 5 13), S. knerii (open triangles; n
5 12), and S. scotorhabdotus (black dots; n 5 94).
95% confidence ellipses and centroids (+ symbols)
are indicated for each species. Principal component
1 correlates strongly with standard length; the most
positive PC1 scores represent the smallest juveniles
in the analysis while the most negative PC1 scores
represent the largest adults.
326944, 1 CS, 106.7 mm SL, USNM 326944, 5, 1
CS, Brazil, Mato Grosso, Rio Paraná, Ilha Solteira;
USNM 317896, 1, Brazil, São Paulo, near Santa
Rosa de Viterbo, Barragem de Haipaiua, Usina
Amália, Rio Pardo drainage, Rio Pardo, main
channel of river.
Schizodon knerii. Lectotype: NMW 62840:2,
235.0 mm SL, Brazil, Rio São Francisco, only
photographs examined. Non-type material: Bra-
zil: Bahia: ANSP 171828, 1, 142.5 mm SL, Rio
Tatu (Bahia do Coco), tributary of Rio Itaguari/
Rio Carinhanha, 7.0 km S of Cocos, 14u149220S,
724 COPEIA, 2007, NO. 3
44u319420W; ANSP 171829, 3 of 5 examined,
109.3–143.1 mm SL, Riacho Santana (isolated
stagnant pool), 3.1 km S of Bom Jesus da Lapa
on road to Malhada, 13u319140S, 43u219280W;
UMMZ 216375, 1, 81.6 mm SL, Barra do Rio
Grande, Rio São Francisco, Minas Gerais; ANSP
171830, 2, 124.6 mm SL, CS male and 125.2 mm
SL, Brazil, Rio Salinas, tributary of Rio Verde
Grande, 51.0 km WSW from Monte Azul on
road to Jaı́ba, 15u129530S, 43u159490W; ANSP
171831, 3, 175.7–200.9 mm SL, Brazil, Rio
Verde Grande, on road from Montes Claros to
Janauba, 16u399010S, 43u429490W; MZUSP
88602, 2 CS, 127.2 male and 179.9 mm SL ripe
female, Três Marias, São Francisco, Rio São
Francisco.
ACKNOWLEDGMENTS
B. Chernoff and M. Westneat provided signif-
icant advice during all stages of this project.
Comments and suggestions from M. Foote, S.
Hackett, R. Vari, and L. Van Valen helped to
improve this manuscript considerably. J.
Armbruster, W. Dahdul, O. Oyakawa, M. de
Pinna, F. Provenzano, M. Retzer, M. Rogers, M.
Sabaj, K. Swagel, R. Vari, H. Wellendorf, and D.
Werneke graciously provided access to, images
of, or loans of specimens in their care. H.
Wellendorf also spent considerable time search-
ing through the NMW collection for the syntypes
of Schizodon isognathus, a service for which we are
deeply grateful. P. Woods provided translations
of the original German descriptions of S.
isognathus and S. knerii. N. Salcedo kindly trans-
lated the abstract into Spanish. M. Weitzman
prepared the baseline map that appears in Figure
4. The scatterplot in Figure 7 was prepared with
R. Strauss’ helpful MATLAB script ‘‘plotgrps,’’
available at http://www.biol.ttu.edu/Strauss/
Matlab/matlab.htm. S. Bartch, M. Payne, and J.
Paulas provided advice on Greek and Latin
vocabulary and usage. BLS was supported during
portions of this work by STAR graduate fellow-
ship #915987 from the U.S. Environmental
Protection Agency and an Armour Graduate
Fellowship from FMNH. NSF doctoral disserta-
tion improvement grant #DEB0412364 and
a Böhlke award from ANSP helped finance this
research.
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(BLS) UNIVERSITY OF CHICAGO, COMMITTEE ON
EVOLUTIONARY BIOLOGY, 1025 E. 57TH STREET,
CULVER HALL 402, CHICAGO, ILLINOIS 60637;
( JCG) DEPARTAMENTO DE ECOLOGIA E BIOLOGIA
EVOLUTIVA, UNIVERSIDADE FEDERAL DE SÃO CAR-
LOS—UFSCAR, RODOVIA WASHINGTON LUÍS, KM
235, P.O. BOX 676, 13565–905 SÃO CARLOS, SP
BRAZIL; ( JJ) UNIVERSITY OF CHICAGO, DEPART-
MENT OF MOLECULAR GENETICS AND CELL BI-
OLOGY, 920 E. 58 STREET, CHICAGO, ILLINOIS
TH
60637; AND (BLS, JJ) DIVISION OF FISHES, THE
FIELD MUSEUM, 1400 SOUTH LAKE SHORE DRIVE,
CHICAGO, ILLINOIS 60605 . E-mail: (BLS) bls16@
duke.edu; ( JCG) garavelo@power.ufscar.br; and
(JJ) jrjellen@uchicago.edu. Send reprint requests
to BLS. Submitted: 14 Sept. 2006. Accepted: 7
Feb. 2007. Section editor: C. J. Ferraris.