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1350
Accepted by C. Gilbert: 20 Sept. 2006; published: 2 Nov. 2006 21
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2006 Magnolia Press
Zootaxa 1350: 21–31 (2006)
www.mapress.com/zootaxa/
Cnesterodon holopteros (Cyprinodontiformes: Poeciliidae:
Poeciliinae), a new species from the Republic of Uruguay
PAULO H. F. LUCINDA1, THOMAS LITZ2 & ROBERTO RECUERO3
1Laboratório de Ictiologia Sistemática, Universidade Federal do Tocantins, Campus de Porto Nacional, Rua
3, Quadra 17, s/n, Jardim dos Ipês, Caixa Postal 136, 77.500-000 Porto Nacional, TO, Brazil.
E-mail: lucinda@uft.edu.br.
2Krumpfhalde 47, 88448 Attenweiler, Germany. E-mail: TCLitz@aol.com.
3Senen Rodríguez 773, Las Piedras 23000, Canelones, Uruguay. E-mail: titorecu@hotmail.com.
Abstract
Cnesterodon holopteros new species is described from the río Uruguai drainage. The new species is
distinguished from its congeners by: (1) a series of small dark brown dots along predorsal portion of
first, second or third lateral scale row, associated with vertical bars along body sides; (2) eight or
nine (rarely seven) dark brown vertical bars (from one to three scales in depth) along lateral surface
of body in females, which may extend nearly down to belly (i.e., at level of ventral profile); (3)
bony style at male gonopodium tip long and gently arched, narrowing towards tip to form a distal
filament and bearing a V-shaped membrane restricted to its proximal third; (4) four branchiostegal
rays in females; and (5) unbranched pelvic-fin rays.
Key words: Neotropical Ichthyofauna, taxonomy, phylogeny, live-bearing killifishes
Resumen
Cnesterodon holopteros nueva especie es descripta de la cuenca del río Uruguay. La nueva especie
se distingue de sus congéneres por: (1) una serie de pequeñas manchas pardas oscuras a lo largo de
la porción predorsal de la primera, segunda o tercera linea de escamas relacionadas con barras
verticales a los costados del cuerpo; (2) ocho o nueve (raramente siete) barras verticales pardas
oscuras en los flancos de las hembras (ocupando una a tres hileras de escamas), que pueden
alcanzar casi hasta el vientre (p.ej. en el nivel del perfil ventral); (3) proceso óseo en el extremo del
gonopodio masculino largo, suavemente arqueado, angostándose hacia el extremo para formar un
filamento distal, con una membrana en forma de V, restringida a su tercio proximal; (4) 4 radios
branquiostegos en hembras; (5) radios pélvicos no ramificados.
LUCINDA
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ZOOTAXA Introduction
Cnesterodon Garman, 1895 is a well-diagnosed, monophyletic genus of poeciliid fishes
whose nomenclature, taxonomic history, and phylogenetic relationships were discussed in
detail by Lucinda & Reis (2005) and Lucinda (2005), respectively. Fishes of the genus
Cnesterodon are small poeciliids, diagnosable by the possession of the following uniquely
derived and unreversed features: (1) anterior tip of basipterygium sinuous in adult males;
(2) outer surface of basipterygium base narrow in adult males; (3) haemal arch and spine
of vertebrae 13–17 in adult males modified in rudimentary gonapophyses; (4)
gonapophysis of vertebra 14 rudimentary in adult males; (5) distal portion of pleural ribs 6,
7, and 8 in adult males expanded; (6) large bony basal process on first anal-fin proximal
radial in adult males; (7) unpaired appendix at tip of anal-fin ray 3 (R3); (8) distal segment
at tip of anterior ramus of anal-fin ray 5 (R5a) transformed into a retrorse triangular spine;
(9) dark brown spot posterior to anal-fin base of males continuous ventrally side by side
and meeting ventral median line of caudal peduncle (Lucinda, 2005). Lucinda & Reis
(2005) recognized the clade Phalloceros Eigenmann, 1907 + Phallotorynus Henn, 1916 as
the sister-group of Cnesterodon, which together comprise the tribe Cnesterodontini.
Despite the well-corroborated monophyletic diagnosis and phylogenetic position of the
genus Cnesterodon, its intrageneric relationships, intrageneric diversity, and species
distribution patterns are not well understood.
Cnesterodon comprises nine recognized valid species inhabiting coastal and inland
drainages of Argentina, southern Brazil, Paraguay, and Uruguay, with most of the known
species restricted to the headwaters or upper portions of the river systems throughout their
range (Lucinda, 2003; 2005). The first described Cnesterodon species is the generic type,
C. decemmaculatus (Jenyns, 1842). A second species, C. carnegiei Haseman, 1911, was
described 69 years later. A gap of 82 years separates the description of C. carnegiei from
the first taxonomic revision of Cnesterodon by Rosa & Costa (1993). The number of
species of Cnesterodon has grown significantly in little more than a decade, with four
species recognized in Rosa & Costa’s (1993) revision, and increasing to nine by the time
Lucinda’s (2005) study appeared. The most recent species described include C. hypselurus
Lucinda & Garavello, 2001; C. omorgmatos Lucinda & Garavello, 2001; C. raddai Meyer
& Etzel, 2001; and C. iguape Lucinda, 2005; with another new species currently being
described by Anza, Lucinda & Malabarba (referred to as Cnesterodon sp. n. B in Lucinda,
2005, Lucinda & Reis, 2005, and herein). Recent collections in the río Uruguay basin in
the department of Artigas, northern Uruguay (Fig. 1) have revealed yet another new
species of Cnesterodon, which is described herein.
Material and methods
Methods of counting, catalogue entries under material examined, geographic descriptors,
and comparative material follow Lucinda (2005). Measurements are those listed by
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Lucinda (2005), except for the snout-occipital measurement, which was excluded from the
present study. Measurement procedures are those listed by Rosa & Costa (1993). In
species descriptions, numbers in parentheses following the counts indicate number of
specimens for each count. Holotype counts are indicated by an asterisk. Tables of
descriptive morphometrics were elaborated with “Datax vers. 4.2” by R. Reis & N.
Fontoura. Cleared and stained specimens (c&s) were prepared according to the method
enumerated by Taylor & Van Dyke (1985). Nomenclature of the gonopodium follows
Rosen & Gordon (1953) and Lucinda & Reis (2005), which refer to first to ninth anal-fin
rays, respectively, as R1, R2, R3, ……and R9. Anterior and posterior rami of fourth and
fifth anal-fin rays are referred to, respectively, as R4a, R4p, R5a, and R5p. Descriptions of
gonopodial morphology are based on fully developed gonopodia of large adult males.
FIGURE 1. Distribution map showing approximate distribution of Cnesterodon species in Rio
Grande do Sul State of Brazil and in the Republic of Uruguay. Squares: collection localities of
Cnesterodon holopteros (T indicates type locality); shaded gray area: Cnesterodon brevirostratus;
shaded green area: Cnesterodon decemmaculatus.
LUCINDA
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ZOOTAXA Cladistic procedures and character state assignments in transformation series are based
on those presented by Lucinda & Reis (2005). Data gathered from the new taxon were
added to the phylogenetic data presented in Lucinda & Reis (2005) in order to evaluate its
phylogenetic position.
Museum aconyms are: MCP — Museu de Ciências e Tecnologia, Pontifícia
Universidade Catolica do Rio Grande do Sul, Porto Alegre; MUNHINA — Museo
Nacional de Historia Natural y Antropologia, Montevideo, Uruguay; and UNT —
Laboratório de Ictiologia Sistemática, Universidade Federal do Tocatins, Porto Nacional,
Brazil.
Cnesterodon holopteros, n. sp.
(Figs 2–4; Table 1)
Holotype (Fig. 2). MUNHINA 3218 (male, 19.8 mm SL), Arroyo Catalancito, Río
Cuareim drainage, Ruta 30, km 156.4, 30°37.87’S, 56°22.75’W, Departmento Artigas,
Uruguay, P. Laurino, T. Litz, E. Perujo, I. Perujo and H. Salvia, 16 Aug 2002.
Paratypes. Uruguay. Departmento Artigas. MCP 39803 (2, 19.5–23.4 mm SL),
MUNHINA 3219 (2, 17.9–20.3 mm SL) and UNT 1946 (4 [2 c&s], 19.4–28.1 mm SL),
paratopotypes, collected with the holotype. UNT 1964 (3, 11.4–14.5 mm SL), lagoon near
Franquia, 30°13.05’S, 57°37.29’W, P. Laurino, T. Litz, E. Perujo, F. Prieto and H. Salvia,
18 Mar 2003.
Diagnosis. Cnesterodon holopteros is distinguished from its congeners by (1) a series
of small dark brown dots present along predorsal portion of first, second or third lateral
scale rows, which are associated with vertical bars along lateral surface of body (Figs.
2–3); (2) eight or nine (rarely seven) dark brown vertical bars along lateral surface of body,
which may reach lower part of belly (i.e., at level of ventral profile) in females, the length
of each bar corresponding approximately to the vertical depths from one to three scales
(Fig. 3); (3) bony style at male gonopodium tip long and gently arched, narrowing towards
tip, forming a distal filament and bearing a V-shaped membrane restricted to its proximal
third (Fig. 4); (4) four branchiostegal rays in females; and (5) unbranched pelvic-fin rays
(branched pelvic-fin rays [except first one] in remaining Cnesterodon species).
Description. Morphometric data appear in Table 1. Ranges of standard length: 19.4 to
28.1 mm (females); 11.4 to 22.7 mm (males). Body compressed laterally. Body width in
predorsal region uniform, about half body depth. Post-dorsal region compressed near
caudal peduncle. Predorsal profile convex. Dorsal-fin base convex. Postdorsal profile
slightly concave. Pre-anal profile convex. Anal-fin base oblique. Post-anal profile almost
straight. Dorsal fin with semicircular border, located posterior to mid-body. Origin of
dorsal fin in females on vertical passing through base of third anal-fin ray; in males, origin
of dorsal fin posterior to vertical passing through origin of anal fin. Pectoral fin inserted
high on each side of body, at level of center of orbit. Pelvic fins small, pointed and not
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reaching origin of gonopodium in adult males; not reaching origin of anal fin in females.
Anal fin of females with straight ventral border. Origin of anal fin of females closer to
caudal peduncle than to snout tip. Origin of anal fin of males closer to snout tip than to
caudal peduncle. Mouth superior, aligned almost with upper border of pupil.
TABLE 1. Descriptive morphometrics of holotype and type series of Cnesterodon holopteros.
Measurements 1–9 are percentages of SL and measurements 10–13 are percentage of HL ( n =
number of specimens).
Dorsal-fin rays: 8* [10], 9 [2]. Pectoral-fin rays: 8 [2], 9* [4], 10 [4]. Pelvic-fin rays
(all unbranched): 4* [2] (males); 5 [4] (females). Anal-fin rays (females): 10 [5]. Anal-fin
rays (males): 8 [3], 9* [4]. Caudal-fin rays: 22 [2], 24 [4], 25 [3], 26* [3]. Predorsal scales
(females): 13 [1], 14 [3]. Longitudinal series scales: 29 [2], 30* [6], 31 [3]. Scales around
caudal peduncle: 16* [12]. Scales in transverse row: 8* [11]. Pleural ribs: 16 [2].
Epipleural ribs: 7 [1, female], 9 [1, male]. Vertebrae: 33 [2].
Preorbital ramus of cephalic sensory system represented by two or three grooved
neuromasts. Preorbital canal absent. Anterior portion of supraorbital ramus (pores 1 and
2a) parallel to upper lip, with five inconspicuous neuromasts on each side. Posterior
portion of supraorbital ramus (pores 2b, 3, 4a) composed of three grooved neuromasts.
Posterior remnants of infraorbital ramus represented by three neuromasts (pores 4b, 5, 6a)
and by one short groove (pores 6b and 7). Preopercular ramus represented by large groove
along preopercular posterolateral border and by prolonged canal along preopercle ventral
border by four neuromasts. Opercular canal absent. Mandibular ramus composed of two or
three superficial neuromasts (pores Z, Ya, and Yb) on anterior border of ventral surface of
mandible and by one superficial neuromast near maxillary distal end (pore W).
Males (n = 5) Females (n = 4)
Character Holotype Range Mean Range Mean
Standard length (mm) 19.8 17.9–22.7 20.1 19.4–28.1 22.8
1. Head length 24.2 19.0–24.2 21.8 22.8–27.8 26.2
2. Predorsal distance 51.5 45.8–53.6 50.4 53.7–74.4 61.4
3. Dorsal-fin base lenlellength 10.1 8.8–11.2 9.8 11.8–12.8 12.2
4. Anal-fin base length 10.1 6.2–13.8 8.7 9.6–10.7 10.2
5. Body depth 32.3 23.4–33.5 30.1 15.5–26.9 23.2
6. Pre-pelvic length 26.8 22.0–26.8 24.8 45.3–55.2 48.8
7. Pre-anal length 33.8 30.8–35.2 33.0 55.6–63.7 58.9
8. Postanal length 66.2 59.5–70.3 65.5 36.3–57.2 44.1
9. Caudal peduncle depth 13.6 11.9–15.6 13.8 13.5–15.5 14.3
10. Snout length 12.5 12.5–28.3 19.4 17.2–26.2 21.9
11. Orbital diameter 37.5 37.5–43.6 40.0 35.4–38.9 37.2
12. Postorbital length 50.0 32.6–50.0 42.1 38.5–42.2 40.1
LUCINDA
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FIGURE 2. Cnesterodon holopteros. Holotype, male, MUNHINA 3218, 19.8 mm SL, Arroyo
Catalancito, Departmento Artigas, Uruguay.
FIGURE 3. Cnesterodon holopteros. Paratype, female, UNT 1946, 28.1 mm SL, collected with the
holotype.
FIGURE 4. Gonopodial appendix of Cnesterodon holopteros, paratype, UNT 1946. Bar 1 mm.
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Gonopodial complex and male gonopodium. Gonopodial complex composed of
nine gonactinosts. Functional gonapophyses absent. Gonactinosts 2, 3, 4 fused.
Gonactinost 4 with wing-like expansions. Ligastyle absent. Gonopodium symmetrical.
Eight or nine gonopodial rays. Anal-fin ray (R1) and anal-fin ray 2 (R2) unbranched and
short. R1 with six segments. R2 with nine segments. R3 with 26 segments. At tip, long
slender bony style bearing narrow membrane produced in terminal filament. Four to six
paired retrorse spines on distal segments of posterior ramus of anal fin of ray 4 (R4p).
Anterior ramus of anal ray 5 (R5a) with subdistal and discrete dorsal curvature and
terminating in a retrorse claw. Anal rays 6, 7, 8 (R6, R7, and R8) branched. Distal
segments of R6 and R7 partially ankylosed. Bony style at tip of gonopodium in adult
males comparatively long and very slightly and gently arched, narrowing towards tip and
forming a short distal filament. Membrane on bony style V-shaped and restricted to
proximal third of bony style (Fig. 4).
Color in alcohol. Eye black, with greenish-brown pupil. Background color cream
yellow. Borders of scales and subjacent skin with numerous brown chromatophores,
conferring a reticulate pattern to body sides. Dorsum darker than ventral region. Head
dorsum dark brown. Median faint brown line along predorsal surface. Median dark brown
line along preanal surface. Fins hyaline. Fin rays with two rows of brown chromatophores
on each side, along entire ray. Males bearing eight or nine (rarely seven) dark brown
vertical bars along sides of body, corresponding approximately to vertical depth of one
scale and mostly confined to midline. Females bearing eight or nine (rarely seven) dark
brown vertical bars along sides of body, corresponding approximately to vertical depth of
one to three scales and sometimes reaching belly (at level of ventral profile). Series of
small dots present along predorsal portion of first, second, or third lateral scale row. Large
dark brown blotch on each side of ventral portion of body near male gonopodium, meeting
midventral post-anal line.
Comparisons. Characters cited in the “Diagnosis” support the recognition of this
species as distinct from its congeners. Cnesterodon holopteros can also be distinguished
from its congeners as follows: It is distinguished from C. omorgmatos and C. raddai by
having dark brown blotches along sides of body (forming bars vs. circular or irregular,
respectively); from C. hypselurus by the absence of a longitudinal dark brown stripe along
flanks (vs. presence of such a stripe in C. hypselurus); from C. septentrionalis Rosa &
Costa, 1993 in having a greater number of longitudinal scales (29–31 vs. 25–26,
respectively) and more transverse scale rows between dorsal and anal fins (8 vs. 7,
respectively). The absence of small scales covering the lateral and ventral region of body
below the pectoral fin in adult females and the pointed snout distinguishes C. holopteros
from C. brevirostratus Rosa & Costa, 1993. Cnesterodon holopteros is readily
distinguished from C. iguape (Lucinda, 2005:131; Fig. 5a) by the smaller post-gonopodial
blotch at the level of the ventral profile in adult males. Cnesterodon holopteros is
distinguished from C. carnegiei by the number of epipleural ribs (7–9 vs. 10–12) and by
the colour pattern.
LUCINDA
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ZOOTAXA Ecological notes. This species is known from the Arroyo Catalancito and a lagoon
near Franquia, both locations of which are located in the río Uruguay basin, within the
department of Artigas, Uruguay (Fig. 1). The type locality is a small, shallow, sidewater
creek tributary of the río Cuareim (= rio Quaraí), and is characterized by rocks, loose
stones, and gravel bottom with clear rapidly-flowing water. Grass and other vegetation
were present in the margins, and dense stands of Echinodorus uruguayensis grew in some
areas. Cnesterodon holopteros was found exclusively in shallow, densely vegetated areas
along the stream margins. The fish fauna in the arroyo Catalancito was equivalent to that
described by Pessano et al. (2005). However, the second location was a large lagoon of the
río Uruguay, near Franquia, with a diameter of approximately 100 meters. This location is
situated only a few kilometres south of the mouth of the río Cuareim, which is tributary to
the río Uruguay. Cnesterodon holopteros was also collected in shallow, densely vegetated
areas along the margins of the lagoon. Ecological data at the collecting site (18 March
2003) were air temperature: 26°C, water temperature 28°C, pH 6.8, and conductivity 78
µS/cm. About 30 fish species may be present in this lagoon, based on Sverlij et al’s (1998)
report for the río Uruguay.
It is noteworthy that C. decemmaculatus was also collected at a few locations in this
hydrographic region, but compared to other hydrographic regions in Uruguay fishes of the
genus Cnesterodon are rare, and other poeciliod species such as Phalloceros
caudimaculatus (Hensel, 1868) and Jenynsia onca Lucinda, Reis & Quevedo, 2002
apparently are absent. These results are in accord with Pessano et al. (2005) and Sverlij et
al. (1998), who did not report P. caudimaculatus or J. onca from either the río Cuareim or
río Uruguay drainages. In the description of J. onca, the distribution was reported to
include the río Ibicuí and río Negro drainages (which are situated, respectively, north and
south of the area under discussion) (Lucinda et al., 2002), but not the area in between. This
supports the hypothesis that the río Cuareim region differs hydrographically in some
respects from northern and southern drainages of the río Uruguay. However, Devincenzi &
Teague (1942) reported the collection of Cnesterodon decemmaculatus and Jenynsia
lineata (Jenyns, 1842) (the latter probably a misidentification of Jenynsia multidentata
(Jenyns, 1842)) from the Laguna de Curtiembre, tributary to the middle río Uruguay,
situated a few kilometres north of the city of Paysandú; about 200 km south of the
locations we investigated.
Distribution. Cnesterodon holopteros is currently known to inhabit the Arroyo
Catalancito and a lagoon near Franquia, Río Cuarein basin, Departmento Artigas, Uruguay
(Fig. 1).
Etymology. From the masculine, nominative, singular, Greek adjective olopteroj [=
holopteros], meaning the whole (i. e. undivided) fins, in reference to the unbranched
pelvic-fin rays.
Phylogenetic relationships. A total of 2209 equally most parsimonious cladograms
were found (length = 755, CI = 0.35, RI = 0.75), with three equally most parsimonious
© 2006 Magnolia Press 29
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trees describing variation in topology concerning Cnesterodon species relationships.
Cnesterodon topologies obtained are partially isomorphic with those discussed in Lucinda
(2005). However, the inclusion of C. holopteros resolved the basal polytomy of Lucinda´s
hypothesis (2005: 262; Fig. 2). All most parsimonious cladograms indicate: (1) C. raddai
is sister to all other Cnesterodon species; (2) Cnesterodon sp. n. B is sister to all remaining
Cnesterodon species; and (3) Cnesterodon holopteros is sister to a clade composed of C.
decemmaculatus, C. carnegiei, C. omorgmatos, C. brevirostratus, C. septentrionalis, C.
hypselurus, and C. iguape. A strict consensus cladogram appears in Fig. 5.
FIGURE 5. Strict consensus cladogram depicting hypothesis of phylogenetic relationships among
Cnesterodon species. Consensus tree resultant from 2209 equally most parsimonious cladograms
for the ingroup (Poeciliinae). Length = 755, Consistency Index = 0.35, Retention Index = 0.75.
Numbers on branches are node numbers. Support for nodes are as follows: Unique and unreversed
character states indicated with an asterisk. 1: 3.2, 4.3, 21.1, 32.1, 34.1, 36.3, 38.2, 40.1*, 42.2,.43.1,
45.1, 47.0, 48.3*, 49.0, 50.0, 53.3, 57.1*, 58.1, 60.0, 63.0, 65.2, 69.0, 70.2, 78.1, 80.2, 86.4, 91.1*,
92.1*, 93.1*, 102.1*, 111.1, 112.1, 117.2, 123.1, 125.1, 129.1, 132.0, 140.1*. 2: 60.2, 144.0. 3:
34.2, 62.1, 64.0. 4: 75.2. 5: 28.0, 63.1, 104.1*. 6: 12.0. 7: 22.0. 8: 34.1, 64.1.
Discussion
For the phylogenetic analyses, states for the new species are the following for Lucinda &
Reis’ (2005) characters 1 to 144: 002300011? 11-1010000 11-0101111 0112030201
0210100300 003---1102 1100210102 0020000112 0110041000 1110-----0 01000---00
1100102000 0010101210 0010000001 0100. The character states exhibited by
Cnesterodon holopteros allow its recognition as a member of the monophyletic genus
Cnesterodon, which is diagnosed by the nine uniquely derived and unreversed features
(Lucinda, 2005: 261). Global parsimony of character states in the consensus tree also
supports C. raddai as sister to a clade composed of all other Cnesterodon species.
Cnesterodon sp. n. B is sister to all remaining Cnesterodon species; and Cnesterodon
holopteros is sister to a clade composed of C. decemmaculatus, C. carnegiei, C.
omorgmatos, C. brevirostratus, C. septentrionalis, C. hypselurus, and C. iguape. The
generic phylogenetic relationships are not completely satisfactory given the polytomy (C.
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ZOOTAXA carnegiei, C. omorgmatos, (((C. brevirostratus, C. septentrionalis), (C. hypselurus, C.
iguape)))), but the cladogram expresses the up-to-date state of knowledge of the
relationships among Cnesterodon species. However, it represents an improvement on the
state of knowledge concerning Cnesterodon infrageneric relationships, given that
inclusion of C. holopteros resolved the basal polytomy of Lucinda’s (2005) hypothesis
(2005: 262; Fig. 2). This is an example on how the discovery of new characters and/or
species can help in the clarification of phylogenetic relationships.
It is likely that additional undescribed Cnesterodon species will be discovered as a
result of ever-increasing collecting efforts in scarcely sampled and unsampled South
American regions, and to a marked increase in attention to poeciliid fishes in recent years.
Acknowledgments
We are deeply indebted to the various curators for loan of specimens under their care. This
manuscript benefited from the critiscims of Roberto Reis (MCP), Lynne Parenti (USNM),
Carter Gilbert (FMNHL), and three anonymous reviewers. We thank the Laboratório de
Ictiologia Sistemática da Universidade Federal do Tocantins (UNT), the Núcleo de
Estudos Ambientais da Universidade Federal do Tocantins (NEAMB-UFT), for financial
and technical support during this study. We thank Stefan Körber, Mulheim, Germany, for
translation of the Spanish abstract. We are grateful to Sérgio Dias-da-Silva (UFT) for help
with the phylogenetic analysis, and we thank Anderson Soares (UNT) for help with the
figures. Thomas Litz thanks Elbio Lartigau, Pablo Laurino, Enrique Perujo, Francisco
Prieto, and Heber Salvia and their families for company, hospitality, and friendship during
various collecting trips in Uruguay. We also thank Hebert Nion (DINARA) for various
discussions and for arranging permissions. Partial financial support for Paulo Lucinda
came from a Research Grant from the Brazilian Federal Government (CNPq process
304838/2005-5).
Literature cited
Devincenzi, G.J. & Teague, G.W. (1942) Ictiofauna del Río Uruguay Medio. Anales del Museo
Nacional de Historia Natural de Montevideo, 5, 1–103, pls. 1–6.
Lucinda, P.H.F. (2003) Family Poeciliidae. In: Reis, R.E., Kullander, S.O. & Ferraris, Jr., C.J.
(Eds.), Check List of the Freshwater Fishes of South and Central America. Edipucrs, Porto
Alegre, pp. 555–581.
Lucinda, P.H.F. (2005) Systematics of the genus Cnesterodon Garman, 1895 (Cyprinodontiformes,
Poeciliidae, Poeciliinae). Neotropical Ichthyology, 3, 259–270.
Lucinda, P.H.F. & Reis, R. E. (2005) Systematics of the subfamily Poeciliinae Bonaparte (Cyprin-
odontiformes, Poeciliidae). Neotropical Ichthyology, 3, 1–60.
Lucinda, P.H.F., Reis, R.E. & Quevedo, R. (2002) Jenynsia onca, a new species of anablepid fish
(Teleostei: Cyprinodontiformes) from southern Brazil and its phylogenetic position. Ichthyo-
© 2006 Magnolia Press 31
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logical Exploration of Freshwaters, 13, 33–40.
Pessano, E.F.C., Azevedo, C.L.O., Querol, M.V.M., Querol, E., Brasil, L.G., Castro, L.R.B., Pinto,
T.B. & Corrêa, F.V. (2005) Ictiofauna do arroio Quaraí-Chico, bacia do médio rio Uruguai, no
interior do Parque Estadual do Espinilho, Rio Grande do Sul, Brasil. Biotemas, 18, 143–153.
Rosa, R.S. & Costa, W.J.E.M. (1993) Systematic revision of the genus Cnesterodon (Cyprinodonti-
formes, Poeciliidae) with the description of two new species from Brazil. Copeia, 1993,
696–708.
Rosen, D.E. & Gordon, M. (1953) Functional anatomy and evolution of male genitalia in poeciliid
fishes. Zoologica, 38, 1–47.
Sverlij S.B., López, H.L., Schenke, R.L.D. & Ros A.E. (1998) Peces del Río Uruguay. Comision
Administradora del Río Uruguay — Rolypel Ltda, Paysandú, 89 pp.
Taylor, W.R. & Van Dyke, G.C. (1985) Revised procedures for staining and clearing small fishes
and other vertebrates for bone and cartilage study. Cybium, 9, 107–119.
