Systematics and biogeography of the genus Phalloceros Eigenmann , 1907 ( Cyprinodontiformes : Poeciliidae : Poeciliinae ) , with the description of twenty-one new species

The genus Phalloceros is revised. Phalloceros caudimaculatus (Hensel, 1868) and twenty-one new species are recognized in Phalloceros. The species and their distributions are: P. alessandrae, small coastal drainages of the Paraná State; P. anisophallos, small coastal drainages of the Rio de Janeiro State; P. aspilos, rio Parati-Mirim, Rio de Janeiro; P. buckupi, small coastal drainages of the Paraná State; P. caudimaculatus (Hensel, 1868), laguna dos Patos system, lower rio Uruguay, drainages of rio Tramandaí, rio Mampituba and coastal drainages of Uruguay and Argentina; P. elachistos, rio Doce drainage and small coastal drainages of the Espírito Santo State; P. enneaktinos, córrego da Toca do Boi, Rio de Janeiro; P. harpagos, rio Paraná-Paraguai basin and coastal drainages from Espírito Santo to Santa Catarina States; P. heptaktinos, rio Jacuí drainage; P. leptokeras, middle portions of rio Paraíba do Sul drainage; P. leticiae, upper rio Araguaia; P. lucenorum, rio Juquiá drainage; P. malabarbai, coastal drainage of the Santa Catarina State; P. megapolos, drainages of rio São João, rio Cubatão (North) and small adjacent drainages of the Paraná State; P. mikrommatos, rio João de Tiba basin, a coastal drainage of the Bahia State; P. ocellatus, coastal drainages of the Bahia and Espírito Santo States; P. pellos, small coastal drainages of the Paraná State; P. reisi, headwaters of rio Tietê, rio Paraíba do Sul, rio Ribeira de Iguape, and small coastal drainages of the São Paulo State; P. spiloura, coastal drainages of the Rio Grande do Sul and Santa Catarina States; P. titthos, coastal drainages of the Paraná State; P. tupinamba, rio Itamambuca and rio Macacu drainages, small coastal drainages of the São Paulo and Rio de Janeiro States, and P. uai, rio São Francisco basin. A lectotype for Girardinus caudimaculatus is designated. Diagnoses of intrageneric clades of Phalloceros are provided. Diagnoses and descriptions of distributions are provided for each species as well as a key for identification. Phylogenetic and biogeographical features of Phalloceros are discussed.

Phalloceros species are well known from several biological standpoints, being object of study for ecologists, anatomists, embryologists, and many others biology researchers.Despite of this, the genus Phalloceros is disappointingly illstudied from the perspective of systematics.Despite of its wide distribution, Phalloceros caudimaculatus is the only formally described species in the genus.
The intrageneric diversity and relationships remains unknown until now.Lucinda & Reis (2005: fig. 3) depicted a general hypothesis of phylogenetic relationships among several unnamed Phalloceros species and P. caudimaculatus, as part of a more inclusive phylogenetic study on the relationships among poeciliine genera, including, the phylogenetic position of the genus Phalloceros in the subfamily Poeciliinae (Lucinda & Reis, 2005: fig.1).Although the transformation series analysis was provided by Lucinda & Reis (2005), these authors did not present the diagnoses of intrageneric clades or describe the newly identified species.These clade diagnoses and species descriptions are provided herein (Fig. 1).
In species descriptions, numbers in square brackets following the counts indicate number of specimens for each count, whereas an asterisk indicates primary type counts.The entries under examined material and all geographic descriptors (including country) follow Lucinda (2005a and b).
The abbreviations E, W, S, and N are used respectively for East, West, South and North.Due to economy of space, several lots collected in the same locality or in very close localities were omitted.The whole list of examined specimens is available with the author and can be obtained upon request.
In the etymology of new specific epithets, the following abbreviations were used: adj.= adjective; n. = noun; num.= numeral; m. = masculine; f. = feminine; sg.= singular; pl.= plural; nom.= nominative; gen.= genitive; i.f.= indeclinable form.When the specific name is an adjective, the endings for the remaining genders are given in parentheses following the adjective.
Counts follow Lucinda (2005a and b), with the addition of the number of serrae on ray 4p of gonopodium; and with the modification on anal-fin ray counts for males, which also included the last, minute ray.Only branched rays were included in pectoral and caudal fin counts.All counts were made on the left side of adult specimens whenever possible.Adult males were considered those with a fully developed gonopodium.A fully developed gonopodium is that whose distal appendix is completely developed and lacks external coverage of skin.
Morphometric data were obtained as detailed in Lucinda ( 2005a and b).Clearing and staining followed the method of Taylor & Van Dyke (1985).Anatomical illustrations were prepared from sketches of structures from cleared and stained specimens as viewed through a camera lucida mounted on a dissecting stereomicroscope.Number and disposition of cephalic pores follow the nomenclature of Rosen & Mendelson (1960), Gosline (1949), andParenti (1981).Only adult individuals have been examined to avoid undesirable ontogenetic variation.Nomenclature of the gonopodium follows Rosen & Gordon (1953) modified by Lucinda & Reis (2005).Descriptions of gonopodial morphology are based on fully developed gonopodia of large adult males.Anatomical nomenclature, other than gonopodial, follows Rosen & Bailey (1963), Parenti (1981), and Rauchenberger (1989).
The phylogenetic species concept is followed: species are diagnosable and stable groups of organisms representing a stable lineage within a phylogenetic hypothesis.Therefore, species diagnoses are based on autapomorphies.Character state assignments, transformation series and clade numbers follow the phylogenetic analysis performed by Lucinda & Reis (2005).In the diagnoses uniquely derived and unreversed features are indicated by two asterisks (e.g.53-2**); uniquely derived features are indicated by one asterisk (e.g.24-1*).Character state illustrations are provided or referred to by Lucinda & Reis (2005).

Results
Twenty-two Phalloceros species are herein recognized: P. caudimaculatus (Hensel, 1868) and twenty-one new species, whose formal descriptions are provided below.Autapomorphies and synapomorphies supporting the monophyly of the genus and infrageneric clades are presented below as well.
Diagnosis.Phalloceros mikrommatos can be distinguished from its congeners by (1) the first gonapophysis angled 5-15 degrees relative to vertebral column ; and (2) second gonapophysis angled zero-15 degrees relative to vertebral column .Phalloceros mikrommatos can be further distinguished from its congeners but P. leticiae and P. ocellatus by the roundish to round and well-defined ocellated lateral spot [134-1] (Fig. 2).Phalloceros mikrommatos can be distinguished from P. leticiae and P. ocellatus by the number of anal-fin rays in females (11 vs. 10, respectively).
Description.Morphometric data in Tables 1 and 2. Range of SL: 18.1 to 27.0 mm (females), 17.4 to 20.4   .Female urogenital papilla straight along midline, located along midventral line and between anus and base of first anal-fin ray.Hooks of gonopodial appendix absent (Fig. 3).
Color in alcohol.Eye dark iridescent grey with greenish brown pupil.Ground pale yellow.Upper two thirds of body and head densely covered with brown chromatophores.In some individuals these chromatophores more concentrated on longitudinal line region forming faint brown discontinuous band along flanks.Lower third of flanks pale yellow contrasting with upper, darker region.Border of scales and subjacent skin replete with brown chromatophores.Chromatophores more concentrated at short distance from scale border conferring reticulate pattern to body sides, mainly on upper half.Brown chromatophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible.Ocellated lateral spot rounded or roundish encircled by light ring.Vertical bars along flanks inconspicuous or absent.Dorsal-fin membrane bearing faint band of brown chromatophores on distal border of dorsal fin.Faint band of chromatophores near dorsal-fin base.Pectoral-, pelvic-, and caudal-fin rays unpigmented.Dark brown line along R3.Anal fin of females hyaline; few chromatophores scattered through first three rays but not forming defined patch of pigmentation.
Distribution.Phalloceros heptaktinos is only known from tributaries of the arroio dos Ratos, Jacuí basin, in Mariana Pimentel, Rio Grande do Sul.Phalloceros heptaktinos is sympatric with P. caudimaculatus (Fig. 4).Description.Morphometric data in Tables 5 and 6.Range of SL: 18.9 to 45.0 mm (females), 14.3 to 28.8 mm (males Female urogenital papilla straight along midline, located along midventral line and between anus and base of first anal-fin ray.Hooks of gonopodial appendix absent (Fig. 11).Remarks.Among the original type series (Hensel, 1868: 362-364) only the seven specimens from lots ZMB 7425 and ZMB 7426 are surely syntypes of Girardinus caudimaculatus.It is not absolutely sure, whether the 25 specimens of lot ZMB 31496 belongs to the type series, since this number is secondarily based on an old label from the anatomical collection: "No, 25202, " Girardinus caudimaculata, C. da Serra ", -the remainder of the label is illegible (Paepke & Seegers, 1986).This is the collection of the former "Institute of Anatomy of the Humboldt-University" in the 19th century, which used to belong to the Medical Faculty.This is due to the fact that human-anatomists were traditionally doing much research work on comparative anatomy of vertebrates.Thus, collected material was often divided between Zoological (Philosophical) Faculty with the Zoological Collection and the Medical Faculty.Only from approximately 1890 onwards the collections were reunited.But, apparently also R. Virchow, the famous pathologist working at the Charity Hospital at Humboldt-University, took over some of the original anatomical collection for study (Peter Bartsch in litt., 2000).As there is no apparent evidence that Hensel has not examined specimens from lot ZMB 31496, it should be more advisable to label them as syntypes.Whether oncoming evidences demonstrate that these specimens have not been syntypes, they will lose their type status.Reinhold F. Hensel was a German geographer and naturalist who lived in Brazil from 1863 to 1866.He collected in Rio Grande do Sul, especially in Porto Alegre and the German colonies to the north of that city (Papavero, 1973).Further, when Hensel returned to Germany, he published the results of his works in Brazil.In 1867 he published a large geographical report of the Província of São Pedro do Rio Grande do Sul (presently the Rio Grande do Sul State) including a map of the region (Hensel, 1867).Later, he dealt with descriptions of new vertebrate taxa, including Girardinus caudimaculatus (Hensel, 1868;1870).The type-locality of G. caudimaculatus is mentioned as "an der Costa da Serra bei S. Leopoldo in Brunnen und Gräben" [= at the Costa da Serra at São Leopoldo in pounds and ditches].The 1867 map shows that Costa da Serra corresponds to the area located in the south margin of the rio Cadeia and arroio Feitoria nearby the Encosta da Serra, and is delimited to the south by the rio dos Sinos drainage.This region currently corresponds to portions of the counties of São Sebastião do Caí, Estância Velha, and Ivoti.Although it is not possible to precisely locate the type-locality it seems very probable that Hensel would have collected his syntypes in the drainage of the rio Cadeia.

Color in alcohol.
One of 26 specimens of USNM 309769, is not P. caudimaculatus (is P. harpagos).The overall coloration of this individual is different from the remaining and P. harpagos is not sympatric with P. caudimaculatus.This specimen likely came from another lot by mistake.

Phalloceros tupinamba, new species
Figs. 13, 14 Phalloceros sp.n.E Lucinda & Reis (2005).Diagnosis.Phalloceros tupinamba is readily distinguished from its congeners, except P. leptokeras and P. aspilos by (1) the presence a lateral ramus of the female urogenital papilla; (2) female urogenital papilla left turned; (3) large sickle like  hook on the gonopodial appendix located in its inner surface and close to its base; and (4) absence of inconspicuous vertical bars along body sides.Phalloceros tupinamba can be distinguished from P. aspilos by the presence of lateral spot and by the predorsal length of males respectively).Phalloceros tupinamba can be distinguished from P. leptokeras by the lateral spot normally pigmented, vertically elongated, and horizontally covering the length corresponding to one scale length and located on the 16 th , 17 th , 18 th or 19 th scale of longitudinal line (vs.densely pigmented, rectangle like, horizontally covering the length corresponding to two or three scales length located on the 14 th or 15 th (very rarely 16 th ) scale of longitudinal line.
Description.Morphometric data in Tables 7 and 8 Etymology.From the Tupi tupinamba alluding to the indigenous tribe that inhabited the region in Pre-Cabralian times.A noun in apposition.
Distribution.Phalloceros tupinamba is known from the rio Itamambuca and rio Macacu drainages, small coastal drainages of São Paulo and Rio de Janeiro States of Brazil (Fig. 12).
Remarks.The distribution of Phalloceros tupinamba is not continuous.The comparison between specimens from rio Macacu and rio Itamambuca has revealed no significant difference that could justify their recognition as distinct species.Until new evidence is available it is plausible to consider the distribution gap is probably due to the lack of collecting samples.Maybe new collecting efforts may reveal that the species also inhabits the coastal drainages between rio Macacu and rio Itamambuca.

Phalloceros aspilos + P. leptokeras
Clade 79, Lucinda & Reis (2005).Diagnosis.Phalloceros leptokeras can be diagnosed by the following uniquely derived autapomorphy: typical densely pigmented rectangle like lateral spot located on the 14 th or 15 th (very rarely 16 th ) scale of longitudinal line [134-5*] (Fig. 15).Furthermore, P. leptokeras is readily distinguished from its congeners, except P. tupinamba and P. aspilos, (1) by the presence a lateral ramus of the female urogenital papilla (Fig. 16); (2) female urogenital papilla left turned (Fig. 16); (3) large sickle like hook on the gonopodial appendix located in its inner surface and close to its base (Fig. 17); and (4) absence of vertical bars along body sides (Fig. 15).Phalloceros leptokeras can be distinguished from P. aspilos by the presence of lateral spot (vs.spot absent).Phalloceros leptokeras can be distinguished from P. tupinamba by the flattened female urogenital papilla (vs.not-flattened in P. tupinamba) and by the lateral spot densely pigmented, rectangle like, horizontally covering the length corresponding to two or three scales length (vs.normally pigmented, vertically elongated, and horizontally covering the length corresponding to the length of one scale in P. tupinamba).

Diagnosis
Description.Morphometric data in Tables 1 and 2   (3) large sickle like hook on the gonopodial appendix located in its inner surface and close to its base (Fig. 20); and (4) absence of inconspicuous vertical bars along body sides (Fig. 19).Phalloceros aspilos can be distinguished from P. leptokeras and P. tupinamba by the absence of lateral spot.Phalloceros aspilos can also be distinguished from P. tupinamba by the predorsal length of males (54.8-56.3 vs. 57.5-59.7 % SL, respectively).Distribution.Rio Parati-Mirim, Rio de Janeiro (Fig. 18).Diagnosis.Phalloceros elachistos can be distinguished from P. anisophallos, P. megapolos, P. spiloura, P. reisi, P. buckupi, P. alessandrae, P. lucenorum, P. uai, P. pellos, and P. malabarbai by the female urogenital papilla straight along midline and located between the anus and the base of first anal-fin ray (vs.curved to the right, located laterally; border of the anal aperture in contact with the first anal-fin ray).Phalloceros elachistos can be distinguished from P. tupinamba, P. leptokeras, and P. aspilos by (1) the female urogenital papilla straight along midline (vs.slightly left turned); (2) absence of a lateral ramus of the female urogenital papilla (vs.lateral ramus present); and (3) small and simple hook in gonopodial appendix (vs.large sickle like hook) (Fig. 22).Phalloceros elachistos can be distinguished from P. caudimaculatus, P. heptaktinos, P. ocellatus, P. mikrommatos, and P. leticiae by the possession of a hook in the gonopodial appendix (vs.hook absent).

Phalloceros elachistos
Phalloceros elachistos is readily distinguished from P. titthos by the absence of symphyseal papillae on mandible of large females (vs.symphyseal papillae present); and from P. enneaktinos by the possession of seven or eight dorsal-fin rays (vs.nine) (see remarks).Phalloceros elachistos can be distinguished from P. harpagos by the hook of gonopodial appendix, which is very small in adults, forming a minute lateral protuberance (Fig. 22) (vs.not forming a lateral protuberancesee remarks on P. harpagos description).In juveniles hook (usually) is absent or (rarely) minuscule.When present in juveniles hook never situated on the appendix corner (vs.present and located on the appendix corner).Juveniles never present skin covering the tip of appendix (vs.skin present).Distal half of appendix approximately as wide as (or slightly narrower than) proximal half (vs.narrower than proximal half).
Description.Morphometric data in Tables 9 and 10.Range of .Female urogenital papilla straight, located along midventral line and between anus and base of first anal-fin ray.Gonopodial appendix hook small sized.Appendix hook very small in adults, forming lateral minute protuberance (Fig. 22).Hook (usually) absent or minute (rarely) in juveniles.
When present in juveniles hook never situated on appendix corner.Juveniles never possess skin-covering tip of appendix.Distal half of appendix approximately as wide as (or slightly narrower than) proximal half (Fig. 22).Distribution.Drainages of rio Doce, Santa Maria da Vitória, Jucu, and Timbuí, coastal drainages of Espírito Santo State of Brazil (Fig. 18).

Color in alcohol
Remarks.One out of 48 studied specimens presented nine dorsal-fin rays.Color in alcohol.Eye black with greenish brown pupil.Ground color pale brown, darker in upper half.Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, conferring re-  Etymology.From the Greek εννεας [= enneás] (num.i.f.), meaning nine, and ακτινος [= aktinos] (n.f. nom.pl.), meaning rays.A noun in apposition alluding to the number of dorsalfin rays.
Distribution.Phalloceros enneaktinos is only known from the type locality in the Córrego da Toca do Boi, Rio de Janeiro (Fig. 4).
Description.Morphometric data in Tables 11 and 12   Etymology.From the Greek τιτθος [= titthós], n. m. nom.sg., a woman's breast or animal's teat.In allusion to the presence of small papillae at the mandibular symphysis of large adult females.Also alluding to the fact that this character is only present in adult females.
Remarks.The gonopodial tip of this species was not described and illustrated due to the lack of adult males available for study.Diagnosis.Members of this clade share the following not uniquely derived and/or reversed features: (1) dorsolateral process of basipterygium absent or small in adult males ; and (2) nine anal-fin rays in males .

Color in alcohol.
Eye black with greenish brown pupil.Ground Color brown, darker in upper half.Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, conferring reticulate pattern to body sides, mainly on upper half.Brown chromatophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible.Lateral dark brown spot located between 16 th and 19 th scale of longitudinal series.Dorsal-fin membrane hyaline bearing dark band of chromatophores on distal border of dorsal fin and another near dorsal-fin base.Pectoral, pelvic, and caudal fins hyaline.Dark brown line along R3.First three anal-fin rays of females covered of brown chromatophores forming patch of dark pigmentation.
Intraspecific Variability.Variation in some morphological features (e.g.color pattern, hook on gonopodial appendix, measurements) has been found among distinct populations of P. harpagos during this revisionary study.Although differences may be found between some population pairs, it has not been possible to assign any of them as diagnostically distinct from the remaining populations.So, these populations are referred to as the same species.Degree of development of spots varies enormously, ranging from large and wide dark patch through small and narrow spot to no apparent mark.Variation on color cannot be associated as exclusive of any local population, for all variants are generally present in any of the populations.Alternatively the frequencies of each of the color "types" can be correlated to populations.
Specimens from coastal drainages of Rio de Janeiro and São Paulo usually possess a roundish and short lateral spot, whereas southern populations in Santa Catarina usually show an elongate and wide lateral spot reaching dorsal and ventral profiles.Generally, specimens from the rio Paraná drainage possess a narrow and vertically elongate spot not reaching dorsal and ventral profiles.Individuals from some populations, e.g.rio Ribeira do Iguape, rio Iguaçu, present a very narrow and short lateral spot.
Particularly, the population in the rio Ribeira de Iguape is an interesting case.This river is formed by the junction of rio Ribeira and rio Juquiá.Specimens from the rio Ribeira have a very narrow and short lateral spot, while specimens from the rio Juquiá possess a large and wide lateral spot.They are quite different from each other.Despite this, none of them can be distinguished from the bulk of P. harpagos, i.e. each can not be diagnosed or recognized as a distinctive species.
Similarly, some specimens of the lower portions of rio Paraná differ from specimens from rio Paraguay.The former show narrow and vertically elongate lateral spot, whereas the later show large and roundish lateral spot.The two forms occur in the lower rio Paraná.This difference is congruent with differences in the depth of body and caudal peduncle of males.Again, both are clearly different from each other but cannot be distinguished from populations along the range of P. harpagos as a whole.
Degree of development of the hook in the gonopodial appendix may also vary from a large hook to lateral expansion, especially in specimens from coastal drainages in southern Santa Catarina, which usually present a shorter hook on gonopodial appendix.At the moment, it is advisable to treat these differences as intraspecific variation, unless new evidences become available and allow the recognition of new diagnosable taxonomic entities.
Etymology.From the Greek 'αρπαγος [= harpagos], n. m. nom.sg., meaning hook, alluding the presence of a hook in the gonopodial appendix.An noun in apposition.

Phalloceros ocellatus, new species
Figs .Female urogenital papilla straight along midline, located along midventral line and between anus and base of first anal-fin ray.Hooks of gonopodial appendix absent (Fig. 31).

Color in alcohol.
Eye dark iridescent grey with greenish brown pupil.Ground color pale yellow.Upper two thirds of body and head densely covered with brown chromatophores.In some individuals these chromatophores more concentrated on longitudinal line region forming faint brown discontinuous band along flanks.Lower third of flanks pale yellow contrasting with upper, darker region.Border of scales and subjacent skin replete with brown chromatophores, more con- Etymology.The noun ocellatus (-a, -um) is a Latin adj.(m.nom.sg.), meaning with little eyes, from ocellus, diminutive of oculus, eye, alluding to the lateral spot, which is rounded and has an external light ring.An adjective.
Distribution.Phalloceros ocellatus is distributed in coastal drainages of Bahia and Espírito Santo States between Prado and Sooretama (Fig. 12).Phalloceros alessandrae can be distinguished from P. megapolos by the gonopodial appendix normally developed (vs.greatly expanded in wing like expansions); and from P. spiloura, P. anisophallos, P. buckupi, P. uai, P. lucenorum, P. pellos, and P. reisi by (1) the halves of gonopodial paired appendix slender, not sickle like, and similar to each other (vs.sickle like and different from each other), bearing a medial corner (vs.medial corner absent); and (2) hooks on both halves of gonopodial paired appendix (vs.right half hook absent).Phalloceros alessandrae can be distinguished from P. malabarbai by (1) gonopodial appendix straight at the distal tip (vs.strongly arched at the distal tip); (2) distal portion straight and oblique forming a 45 º angle with the vertical axis (vs.distal portion bent upward); and (3) left and right halves coalescent at the base (vs.halves free not coalescent at the base) (Fig. 33). 5 and 6 Female urogenital papilla right curved, located laterally.Border of anal aperture in contact with first anal-fin ray or very close to it.Halves of gonopodial paired appendix slender, not sickle like, and similar to each other, bearing medial corner.Hook on both halves of gonopodial paired appendix.Hooks large downward directed and located at corner of gonopodial appendix.Gonopodial appendix long and straight at distal tip.Distal portion straight and oblique forming 45 º angle with vertical axis.Left and right halves coalescent at base (Fig. 33) Etymology.The specific epithet alessandrae (n.f. gen.sg.) comes from Alessandra.Phalloceros alessandrae is named after my wife Alessandra M. V. Lucinda.

Description. Morphometric data in Tables
Distribution.Phalloceros alessandrae is known from flooded areas on margins nearby Antonina (Paraná, Brazil) and rio Dois de Fevereiro, which flows into the Baía de Paranaguá (Fig. 18).

Phalloceros malabarbai, new species
Figs   Description.Morphometric data in Tables 3 and 4 Left and right halves free not coalescent at base (Fig. 35).

Color in alcohol.
Eye black with greenish brown pupil.Ground color cream, darker in upper half.Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, conferring reticu-late pattern to body sides, mainly on upper half.Brown chromatophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible.Lateral dark brown spot located over 17 th and 18 th scale of longitudinal series.Lateral spot covering approximately two scales on horizontal direction, and four scales on vertical.Dorsal-fin membrane hyaline bearing faint.band of brown chromatophores on distal border of dorsal fin and another near dorsal-fin base.Pectoral, pelvic, and caudal fins hyaline.Dark brown line along R3.
Etymology.The specific epithet malabarbai (n.m. gen.sg.) comes from Malabarba.The specific name malabarbai is a patronym for Luiz Roberto Malabarba, in recognition of his many contributions to Neotropical ichthyology.
Distribution.Drainages of the rio São João, rio Cubatão (Norte) and small adjacent drainages, which flows into the Baía de Guaratuba in Paraná State (Fig. 18).Diagnosis.Phalloceros lucenorum can be autapomorphically diagnosed by the rectangular tip of hook on left half of gonopodial appendix (Fig. 39) (vs.rounded tip).Moreover, P. lucenorum can be distinguished from its congeners but P. megapolos, P. spiloura, P. malabarbai, P. alessandrae, P. buckupi, P. uai, P. anisophallos, P. pellos, and P. reisi by (1) the female urogenital papilla curved to the right, located laterally (vs.slightly left turned and with a lateral ramus or straight located along midventral line) and (2) border of the anal aperture in contact with the first anal-fin ray or very close to it (vs.separated from first anal-fin ray by the female urogenital papilla).Phalloceros lucenorum can be distinguished from its congeners but P. spiloura, P. uai, P. anisophallos, P. pellos, and P. reisi by (1) the absence of the right hook and presence of the left one (vs.absence or presence of left and right hooks); and (2) gonopodial appendix asymmetrical, its halves different from each other; right half wider than left one (vs.gonopodial appendix symmetrical; its halves similar to each other).

Phalloceros lucenorum
Phalloceros lucenorum can be distinguished from P. spiloura by the absence of a caudal peduncle spot (vs.presence of caudal peduncle spot); from P. uai by the presence of a wide and square-shaped lateral spot in large specimens (vs.absence of such spot); from P. anisophallos by the lateral spot in adult females elliptical to roundish, but never forming a vertical bar (vs.lateral spot very narrow, forming a vertically elongated bar covering up to the length corresponding to two scales length in horizontal direction and 2-4 scales in vertical direction); from P. reisi by lateral spot always present and evident not covering the length corresponding to more than two scales length in horizontal or vertical directions (vs.lateral spot absent or small and discrete not covering the length corresponding to one or two scales length in horizontal or vertical directions).3 and 4. Range of SL: 26.9 to 37.0 mm (females), 18.5 to 28.0 mm (males).Dorsalfin rays: Hook on left half of gonopodial appendix with rectangular tip (Fig. 39).

Description. Morphometric data in Tables
Color in alcohol.Eye black with greenish brown pupil.Ground color pale brown, darker in upper half.Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, conferring reticulate pattern to body sides, mainly on upper half.Brown chromatophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible.Lateral spot large covering approximately two or three scales on horizontal direction, and three to five scales on vertical.Dorsal-fin membrane hyaline.Dark band of chromatophores on distal border of dorsal fin.Dark band of chromatophores near dorsal-fin base.Pectoral-, pelvic-, anal-, and caudal-fin rays unpigmented.Dark brown line along R3.
Etymology.The specific epithet lucenorum (n.m. gen.pl.) comes from Lucena.It is a patronym for Carlos A.S. Lucena and Zilda Margarete S. Lucena, in recognition of their many contributions to Neotropical ichthyology.
Distribution.Rio Juquiá, a tributary to the drainage of the rio Ribeira de Iguape in the State of São Paulo (Fig. 12).

Phalloceros uai + P. buckupi
Clade 71, Lucinda & Reis (2005) Diagnosis.Members of this clade share the following not uniquely derived and/or reversed features: (1) anterior cleft of anguloarticular small, not extending beyond posterior border of Meckel's cartilage ; (2) ventral process of anguloarticular long, extending anterior to where anguloarticular overlaps dentary ; and (3) first proximal radial of dorsal fin located between arches neural of 14 th and 15 th vertebrae in adult females .Diagnosis.Phalloceros uai is autapomorphically diagnosed by a lateral spot wide and square-shaped in large specimens [134-3*] (Fig. 40).Moreover, P. uai can be distinguished from its congeners but P. megapolos, P. spiloura, P. malabarbai, P. alessandrae, P. buckupi, P. anisophallos, P. lucenorum, P. pellos, and P. reisi by (1) the female urogenital papilla curved to the right, located laterally (vs.slightly left turned and with a lateral ramus or straight located along midventral line); and

Phalloceros uai, new species
(2) border of the anal aperture in contact with the first anal-fin ray or very close to it (vs.separated from first anal-fin ray by the female urogenital papilla).Phalloceros uai can be distinguished from its congeners but P. spiloura, P. anisophallos, P. lucenorum, P. pellos, and P. reisi by (1) the absence of the right hook and presence of the left one (vs.absence or presence of left and right hooks); and (2) gonopodial appendix asymmetrical; its halves different from each other; right half wider than left one (vs.gonopodial appendix symmetrical; its halves similar to each other).7 and 8 Color in alcohol.Eye iridescent grey with greenish brown pupil.Ground color pale brown, darker in upper half.Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, conferring reticulate pattern to body sides.Brown chro-matophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible.Lateral spot wide and square-shaped in large specimens.Lateral dark brown spot located over 14 th and 15 th , or 15 th and 16 th or 16 th and 17 th scale of longitudinal series.Lateral spot covering approximately two scales on horizontal direction, and three scales on vertical.Dorsal-fin membrane bearing dark band of chromatophores on distal border of dorsal fin.Dark band of chromatophores near dorsal-fin base.Pectoral, pelvic, and caudal fins hyaline.Dark brown line along R3.

Etymology.
Uai is an interjection typical of the natives of the State of Minas Gerais.It usually serves to express surprise, but also everything else.The epithet uai is a homage to Minas Gerais, my birthplace.A noun in apposition.
Distribution.Rio das Velhas, rio São Francisco basin (Fig. 7).Diagnosis.Phalloceros buckupi can be distinguished from its congeners but P. megapolos, P. spiloura, P. malabarbai, P. alessandrae, P. anisophallos, P. uai, P. lucenorum, P. pellos, and P. reisi by (1) the female urogenital papilla curved to the right, located laterally (vs.slightly left turned and with a lateral ramus or straight located along midventral line); and (2) border of the anal aperture in contact with the first anal-fin ray or very close to it (vs.separated from first anal-fin ray by the female urogenital papilla).Phalloceros buckupi can be distinguished from P. spiloura by the absence of a caudal peduncle spot (vs.presence of caudal peduncle spot); from P. megapolos by the gonopodium appendix normally developed (vs.greatly expanded in wing like projections); from P. uai by the absence of a wide and square-shaped lateral spot in large specimens (vs.presence of such spot).
Phalloceros buckupi can be distinguished from P. alessandrae and P. malabarbai by (1) halves of gonopodial paired appendix sickle like (vs.not sickle like); (2) medial corner absent (vs.present); (3) hook on left half of gonopodial paired appendix and right hook absent (vs.hook on both halves of gonopodial paired appendix); and (4) left hook small, directed downward or up-and forward, and located closer to the base of gonopodial appendix (vs.hooks large downward directed and located in the corner of gonopodial appendix) (Fig. 43).
Phalloceros buckupi can be distinguished from P. anisophallos, P. lucenorum, P. pellos, and P. reisi by the halves of the gonopodium slender and similar to each other (vs.halves of the gonopodium wide and different from each other; right half wider than left one).3 and 4. Range of SL: 21.7 to 32.9 mm (females), 17.8 to 19.9 mm (males).Dorsalfin rays:  Color in alcohol.Eye silvery grey with greenish brown pupil.Ground color pale brown, darker in upper half.Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, conferring reticulate pattern to body sides, mainly on upper half.Brown chromatophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible.Lateral dark brown spot located approximately over 16 th scale of longitudinal series.Lateral spot covering approximately one scale on horizontal direction, and three scales on vertical.Light brown vertical bars along flanks (absent in one specimen studied).Dorsalfin membrane bearing dark band of chromatophores on distal border of dorsal fin.Dark band of chromatophores near dorsal-fin base.Pectoral-, pelvic-, anal-, and caudal-fin rays unpigmented.Dark brown line along R3.

Description. Morphometric data in Tables
Remarks.Specimens from Caraguaçu (MCP 12584) are darker than remaining specimens, with the dark bands on dorsal fin more evident and the vertical bars inconspicuous or absent.
Etymology.The specific epithet buckupi (n.m. gen.sg.) comes from Buckup.It is a patronym for Paulo A. Buckup (collector of most specimens of this species) in recognition of his many contributions to Neotropical ichthyology.
Distribution.Phalloceros buckupi is known to occur in the rio Jacareí drainage and neighbouring, which flows into the Baía de Paranaguá in the coast of Paraná State of Brazil (Fig. 12).

Phalloceros anisophallos + P. pellos
Clade 72, Lucinda & Reis (2005) Diagnosis.Diagnosis.Phalloceros pellos can be distinguished from its congeners but P. megapolos, P. spiloura, P. malabarbai, P. alessandrae, P. buckupi, P. uai, P. lucenorum, P. anisophallos, and P. reisi by (1) the female urogenital papilla curved to the right, located laterally (vs.slightly left turned and with a lateral ramus or straight located along midventral line); and (2) border of the anal aperture in contact with the first anal-fin ray or very close to it (vs.separated from first anal-fin ray by the female urogenital papilla).Phalloceros pellos can be distinguished from its congeners but P. spiloura, P. uai, P. lucenorum, P. anisophallos, and P. reisi by (1) the absence of the right hook and presence of the left one (vs.absence or presence of left and right hooks); and (2) gonopodial appendix asymmetrical; its halves different from each other; right half wider than left one (vs.gonopodial appendix symmetrical; its halves similar to each other) (Fig. 47).
Phalloceros pellos can be distinguished from P. spiloura by the absence of a caudal peduncle spot (vs.presence of caudal peduncle spot); from P. uai by the presence of a wide and square-shaped lateral spot in large specimens (vs.absence of such spot); from P. anisophallos by the lateral spot in adult females elliptical to roundish, but never forming a vertical bar (vs.lateral spot very narrow, forming a vertically elongated bar covering up to the length corresponding to two scales length in horizontal direction and 2-4 scales in vertical direction); from P. lucenorum by a rounded tip of the hook on left half of gonopodial appendix (vs.rectangular tip).
Phalloceros pellos can be distinguished from P. reisi by (1) hook on left appendix very small (vs.larger), (2) lateral spot always present and evident not covering more than the length corresponding to two scales length in horizontal or vertical directions (vs.absent or small and discrete not covering one or two scales in horizontal or vertical directions).Color in alcohol.Eye black with greenish brown pupil.Background coloration dark brown, darker in upper half.Overall appearance dark.Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, conferring reticulate pattern to whole flanks.Brown chromatophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible.Margin of scales on prepelvic region clearly defined bordered by dark chromatophores.Lateral dark brown spot located over 15 th and 16 th or 16 th and 17 th scale of longitudinal series.Lateral spot covering approximately two scales on horizontal direction, and two to four scales on vertical, sometimes reaching dorsal and ventral profiles.Inconspicuous vertical bars present in some individuals.Dorsal-fin membrane dusky, bearing dark band of chromatophores on distal border and dark band of chromatophores near dorsal-fin base.Pectoral-, pelvic-and caudal-fin

Phalloceros spiloura + P. reisi
Clade 73, Lucinda & Reis (2005).Diagnosis.Phalloceros spiloura can be diagnosed by the following uniquely derived autapomorphies: (1) the possession of a rounded spot located on the lower half of the caudal peduncle close to the base of lowest caudal-fin rays (Fig. 48);

Diagnosis
(2) a patch of dark pigmentation on the last anal-fin rays of females (Fig. 48b); (3) halves of gonopodial paired appendix straight and perpendicular to R3 [96-2*] (Fig. 49).Moreover, P. spiloura can be distinguished from its congeners by ( 1   Diagnosis.Phalloceros reisi can be distinguished from its congeners but P. megapolos, P. spiloura, P. malabarbai, P. alessandrae, P. buckupi, P. uai, P. lucenorum, P. pellos, and P. anisophallos by (1) the female urogenital papilla curved to the right, located laterally (vs.slightly left turned and with a lateral ramus or straight located along midventral line) and ( 2) border of the anal aperture in contact with the first anal-fin ray or very close to it (vs.separated from first anal-fin ray by the female urogenital papilla).Phalloceros reisi can be distinguished from its congeners but P. spiloura, P. uai, P. lucenorum, P. pellos, and P. anisophallos by (1) the absence of the right hook and presence of the left one (vs.absence or presence of left and right hooks), and (2) gonopodial appendix asymmetrical; its halves different from each other; right half wider than left one (vs.gonopodial appendix symmetrical; its halves similar to each other).
Phalloceros reisi can be distinguished from P. spiloura by the absence of a caudal peduncle spot (vs.presence of caudal peduncle spot); from P. uai by the presence of a wide and square-shaped lateral spot in large specimens (vs.absence of such spot); from P. lucenorum by rectangular tip of hook on left half of gonopodial appendix (vs.rounded tip); from P. anisophallos by the lateral spot in adult females very narrow, forming a vertically elongated bar covering up to the length corresponding to two scales length in horizontal direction and 2-4 scales in vertical direction (vs.absent, small, or elliptical to roundish, but never forming a vertical bar lateral spot in adult females); from P. pellos by (1) the background coloration light brown or yellow (vs.dark brown) and (2) margin of scales on prepelvic region not visible, not bordered by dark chromatophores, forming a light, plain yellow region (Fig. 50) (vs.margin of scales on prepelvic region clearly defined and bordered by dark chromatophores).The lateral spot absent or small and discrete not covering one or two scales in horizontal or vertical directions differs P. reisi from P. lucenorum and P. pellos.
Description.Morphometric data in Tables 9 and 10  Etymology.The specific name reisi (n.m. gen.sg.) is a patronym for Roberto E. Reis, in recognition of his many contributions to Neotropical ichthyology.
Remarks.A single lot is known from the rio Juquiá drainage.Maybe this species is not sufficiently sampled in this area.

Discussion
Phalloceros is a well-diagnosed, monophyletic genus, whose phylogenetic position in the subfamily Poeciliinae was recently discussed by Lucinda & Reis (2005).According to these authors Phalloceros is the sister-group of Phallotorynus Henn, 1916 which together with Cnesterodon Garman, 1895 forms the tribe Cnesterodontini (see Lucinda, 2005b, Lucinda & Reis, 2005;and Lucinda et al. 2005 for further discussion).Lucinda & Reis (2005) were the first to attempt a solution for the recognition or diagnosis of a monophyletic Phalloceros.Previous diagnoses were non-cladistic.How-ever, Phalloceros was monotypic and the diagnosis of the species was the diagnosis for the genus.The lack of previous attempts to diagnose Phalloceros cladistically were partially explained by the monotypy of the genus.Definitions of monotypic genera are somewhat merged with the species definition.Eigenmann (1907) erected the genus diagnosing it by the presence of "antler-like processes" or "antler-like structures" [= paired appendix] at tip of R4a (Eigenmann, 1907: 427 & 431, respectively).However, the paired appendix is attached to R3 not to R4a (Rosen & Bailey, 1963;Lucinda & Reis, 2005).Rosen & Bailey (1963) also proposed this feature as diagnostic for Phalloceros.Lucinda & Reis (2005) regarded the presence of the paired appendix at the tip of R3 as an uniquely derived and unreversed synapomorpy shared by all Phalloceros species.
Eigenmann (1907) also mentioned teeth morphology as diagnostic for Phalloceros: "(…) the presence of outer series of spoon-oar-shaped teeth, somewhat expanded at tip and bent backward, close set, their margins in contact near their tips; much smaller teeth, triangular at tips, slightly contracted at base, forming one or more series behind the larger teeth."Nevertheless, this teeth morphology is not unique to Phalloceros and is widespread among poeciliine genera.Following Eigenmann (1907: 431), the length of the intestine, which is "twice as long as the entire fish," is another diagnostic feature for Phalloceros.This characther was not surveyed by Lucinda & Reis (2005).
The comparative examination of samples of Phalloceros allowed the recognition of twenty-one distinct, previously unrecognized or undiscovered species on the genus.The color pattern, meristic, and morphometric characters support the recognition of these new species as distinct from their congeners (see diagnoses).One of the reasons for this previously undetected diversity may be related to neglect of potential diagnostic features such as the morphology of the gonopodial appendix and the female urogenital papilla.Additionally, Phalloceros and other poeciliids have long been neglected by fish systematists, although this situation is changing.
The intrageneric phylogenetic relationships are far from satisfactory.At present, the cladogram which expresses the up-to-date state of knowledge of the relationships among Phalloceros species, exhibits many polytomies (Fig. 1).Discovery of new characters and/or species would be effective in defeating this polytomic situation.Unfortunately, the lack of resolution across much of the cladogram severely limits the historical biogeographic information available.Although the lack of resolution precludes unambiguous assertions, some tentative biogeogeophical remarks on Phalloceros distribution are outlined below.
Phalloceros species as well as almost all cnesterodontines (with rare exceptions) are found in southern South American drainages, one of the eight areas of endemism for poeciliines in the American continent identified by Lucinda & Reis (2005).Phalloceros species are also found in five of the eigth re-gions of endemism identified by Vari (1988) for the ichthyofauna of cis-Andean South America: (1) coastal drainages above the mouth of the rio Paraíba do Sul, (2) rio Tocantins drainage, (3) rio São Francisco drainage, (4) Paraguay (rio Paraguay, Uruguay, lower Paraná drainages, and coastal drainages south of São Paulo), and (5) upper rio Paraná.It is remarkable that most species inhabit the "Paraguay" and the "Coastal" regions of endemism.Few species are found in the São Francisco (P.uai), upper Paraná (P.harpagos), and Tocantins (P.leticiae).
It is also noteworthy that several species have a very limited distribution, e.g.P. enneaktinos, whereas P. harpagos exhibits a huge distribution range.This pattern of one widely distributed species plus several confined species is shared by some other neotropical fishes.
Most basal species of Phalloceros (Clade 95) inhabit the rio Tocantins and rio João de Tiba drainage.Remaining species (Clade 96) inhabit all coastal drainages from Bahia to the Republic of Uruguay as well as the upper Paraná.Most basal species of Clade 96, namely P. heptaktinos and P. caudimaculatus, are confined to the rio Jacuí, rio Uruguay and neighboring coastal drainages of Rio Grande do Sul and the Republic of Uruguay.Actually, the rio Uruguay drainage seems to represent a barrier to remaining species (Clade 83), for the only species found in the rio Uruguay drainage and neighboring coastal drainages to the south are P. heptaktinos and P. caudimaculatus.Remaining species (Clade 83) are found in upper Paraná and along coastal drainages to the north, i.e. from the rio Araranguá drainage (the austral boundary) to coastal drainages of Bahia State (the septentrional boundary).
Uniquely derived and unreversed features support very distinct monophyletic and endemic species-groups inside Clade 83.Members of Clade 81 are confined to the rio Paraíba do Sul drainage and neighboring coastal drainages and share hooks on paired gonopodial appendix large and sickle-like and female urogenital papilla left turned.Members of Clade 76 share a female urogenital papilla right turned and are split in two diferent well-coroborated monophyletic lineages: Clade 75, whose members are confined to small coastal drainages between the Baía de Paranaguá (Paraná State) and São Francisco do Sul (Santa Catarina) and share large hooks directed downward and located in the corner of gonopodial appendix, and Clade 74, whose members share hooks on gonopodial paired appendix present only on left half and are distributed in coastal drainages of Rio de Janeiro, São Paulo, and Paraná States as well as in the upper reaches of the rio São Francisco basin.
Most species of Phalloceros conform to a general pattern of allopatric distribution, although some species are sympatric or even syntopic.This is the case of Phalloceros harpagos, which is widely distributed and sympatric with many other species.Two other cases of sympatry (syntopy) are found: (1) Phalloceros heptaktinos and P. caudimaculatus and (2) Phalloceros leptokeras, P. reisi and P. harpagos.These distibution and sympatry/syntopy patterns can probably be explained by several past dispersion events concerning these species.
Phalloceros heptaktinos is sympatric and syntopic with P. caudimaculatus in the tributaries of the arroio dos Ratos (Jacuí basin), whereas P. leptokeras, P. reisi and P. harpagos are sympatric in the rio Paraíba do Sul drainage.Phalloceros leptokeras is even syntopic with P. harpagos, which could also require a dispersion explanation.However P. leptokeras and P. reisi are sympatric but not syntopic.Phalloceros leptokeras is restricted to the middle portions of rio Paraíba do Sul drainage, whereas P. reisi inhabits its headwaters.Phalloceros reisi also inhabits the headwaters of rio Tietê, the rio Ribeira de Iguape, and small coastal drainages in São Paulo State of Brazil.The presence of P. reisi in rio Paraíba do Sul and in rio Paraná drainages is congruent with the distribution pattern of Phallotorynus fasciolatus (Lucinda et al., 2005) and many other teleosts.As well, the presence of P. leptokeras in middle portions of rio Paraíba do Sul drainage and the presence of P. reisi in its upper portions is also congruent with the distibution pattern above.Both facts are probably related to and may be explained by the formation of the upper Paraíba do Sul by stream capture of rio Tietê headwaters (Malabarba, 1998).See Malabarba (1998), Weitzman &Malabarba (1999), andQuevedo &Reis (2002) for further discussion.
It is very likely that additional Phalloceros species will be discovered in the near future, owing to increasing collecting efforts on poorly sampled and unsampled South-American areas and to continuous increase of attention poeciliid fishes have received in the last years.Much effort must still be directed towards understanding of Phalloceros systematics and biogeography.

Diagnosis.
Members of this clade share the following not uniquely derived and/or reversed features: (1) ascending process of parasphenoids long and contacting pterosphenoids in adults [20-0]; (2) pelvic girdle of males not very anteriorly located; posterior border of basipterygium aligned with posterior border of chleitrum [35-2]; and (3) anterior tip of basipterygium in adult males clearly pointed [37-1].

.
Eye black with greenish brown pupil.Ground color pale brown, darker in upper half.Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, forming v-band conferring reticulate pattern to body sides, mainly on upper half.Brown chromatophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible.Lateral dark brown spot very small, inconspicuous or absent.When present located approximately on 18th scale of longitudinal series covering approximately one scale on both horizontal and vertical directions.Inconspicuous vertical bars along flanks.Dorsalfin membrane hyaline bearing dark band of chromatophores on distal border of dorsal fin and another near dorsal-fin base.Pectoral-, pelvic-and caudal-fin rays dusky.Dark brown line along R3.Etymology.Specific name from the Greek ελαχυς, −εια [= elachis, -eia], adj.m. nom.sg.], small, short, little, plus the superlative ending −ιστος [= -istos].Ελαχιστος, −η, −ον, [ = elachistos, -e, -on], meaning the least, which alludes to the small size of the specimens.An adjective, gender masculine.

Fig. 29 .
Fig. 29.Collection localities of Phalloceros harpagos.Some symbols represent more than one lot or locality.T represents type locality.The inset is the range of the genus.
Dark band of chromatophores near dorsal-fin base, more intense in posterior rays.Pectoral, pelvic, and caudal fins hyaline.Dark brown line along R3.Anal fin of females hyaline, except for patch of dark chromatophores on first three rays.

Table 7 .
Descriptive morphometrics of female specimens of Phalloceros tupinamba and P. uai.Measurements 1-10 are percents of standard length and measurements 11-13 are percents of head length.

Table 8 .
Descriptive morphometrics of male specimens of Phalloceros tupinamba and P. uai.Measurements 1-10 are percents of standard length and measurements 11-13 are percents of head length.H = holotype.

Table 12 .
Color in alcohol.Eye black with greenish brown pupil.Background coloration dark brown, darker in upper half.Overall appearance dark.Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, conferring reticulate pattern to whole flanks.Brown chromatophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible.Margin of scales on Descriptive morphometrics of male specimens of Phalloceros ocellatus, P. pellos, P. spiloura, and P. titthos.Measurements 1-10 are percents of standard length and measurements 11-13 are percents of head length.H = holotype.
Members of this clade share the following not uniquely derived and/or reversed feature: distal portion third and fourth gonactinosts completely fused [74-1].
Non-types.BRAZIL.Rio Grande do Sul.MZUSP 4481, 10 of 11, Torres, praia de Torres.Santa Catarina.MCP 10502, 18, Gravatal, lateral channel along rio Capivari, on road from Tubarão to Gravatal.MCP 10659, 7, Tubarão, 1984.MCP 22336, 16, Urubici, rio Urubici near bridge S of Urubici (upper rio Uruguai).MCP 23715, 2, Jacinto Machado, arroio affluent of rio Sertão, ca. 13 km SW from Jacinto Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, conferring reticulate pattern to body sides, mainly on upper half.Brown chromatophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible.Faint vertical bars along flanks.Dorsal-fin membrane hyaline.Band of dark chromatophores on distal border of dorsal fin.Band of dark chromatophores near dorsal-fin base, more concentrated posteriorly, forming distinct spot.Dark brown line along R3.Pectoral-, pelvic-, anal, and caudal-fin rays unpigmented.Rounded spot on lower half of caudal peduncle close to base of lowest caudal-fin rays.Patch of dark pigmentation on last anal-fin rays of females.Etymology.From the Greek σπιλος [= spílos], n. m. nom.sg., meaning spot + oura [= ourá], n. f. nom.sg., meaning tail, alluding to the presence of a rounded spot on close to base of lowest caudal-fin rays.A noun in apposition.