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Accepted by M.R. de Carvalho: 8 Sept. 2008; published: 29 Oct. 2008 59
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2008 · Magnolia Press
Zootaxa 1918: 59–68 (2008)
www.mapress.com/zootaxa/
A new Pamphorichthys (Cyprinodontiformes: Poeciliidae: Poeciliini)
from central Brazil
CARLOS AUGUSTO FIGUEIREDO
Universidade Federal do Estado do Rio de Janeiro – Escola de Ciências Biológicas – Departamento de Ecologia e Recursos Marin-
hos Av. Pasteur, 458/sala 408 CEP 22290-240, Rio de Janeiro, RJ, Brazil. E-mail. carlos.figueiredo@gmail.com
Abstract
A new species of Pamphorichthys is described from Preto river, tributary of the São Francisco river drainage. The new
species is the sister group of the remaining species of Pamphorichthys, showing synapomorphies of the genus, including
the reduced number of gonapophyses and gonapophyses tips parallel to the vertebral spine. The diagnosis is given by the
following unique features: outer row of teeth on pre-maxilar and dentarium with bilobed incisive teeth; distal segments
of the third ray of the gonopodium and an elongated anterior branch of the fourth ray with a putative origin in the fusion
of several distal segments; anterior and posterior branches of gonopodium ray 6 with distal segments fused into only one
segment; six dorsal-fin rays in males; and urogenital region of reproductive females with heavy dark pigmentation all
around the anus and urogenital opening.
Key words: Systematics, Taxonomy, Poeciliinae, Livebearer, Endemism
Resumo
Uma nova espécie de Pamphorichthys é descrita do Rio Preto, tributário da bacia do rio São Francisco. A nova espécie é
o grupo irmão das demais espécies do gênero, incluindo o reduzido número de gonapófises e a ponta das gonapofises
paralela à espinha vertebral. A diagnose é dada pelas seguintes características exclusivas: série externa de dentes no pré-
maxilar e dentário com dentes bilobados; segmentos distais dos raios 3 e ramo anterior do raio 4 alongados aparentando
ter sua origem na fusão dos segmentos mais distais; ramos, anterior e posterior do raio 6 do gonopódio com os segmentos
distais fusionados em um só segmento; seis raios na nadadeira dorsal de machos; região uro-genital de fêmeas reproduti-
vas com forte pigmentação escura a toda a volta do ânus e abertura urogenital.
Introduction
Although the Subfamily Poeciliinae, and especially the tribe Poeciliini, are well-known to scientists and
aquarium hobbyists, an expedition to the São Francisco river system in central Brazil in 1998 collected a small
(ca. 20 mm max.), unknown fish species of the tribe Poeciliini. Members of this subfamily are often used as
models in evolutionary and behavioral studies, as well as in many research projects that require sturdy, aquatic
organisms with short generation time. Studies on phylogenetic relationships of the genera composing Poecili-
inae have been conducted (Figueiredo, 2003; Lucinda & Reis, 2005) to allow comparative analysis in these
model systems. These studies made the generic placement of these species possible and brought to light an
unknown biodiversity, represented in part by the highly endemic species herein described.
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Methods
Specimens were examined under a Zeiss SV-11 stereomicroscope with transmitted and direct light, and vari-
able magnification between 6X and 60X. Measurements were taken with a digital caliper and recorded as per-
centages of standard length (SL), except for sub-units of head, which are presented as percentages of head
length (HL). Counts of external meristic structures were made on alcohol-preserved specimens. For internal
counts and osteological examination, cleared and stained specimens (C&S) were prepared using the technique
presented by Taylor and Van Dyke (1985) and further developed by Potthoff (1983), using water instead of
1% KOH with alizarin to stain bones and 8% of H2O2, instead of 30%, during the bleaching procedure, and
exposing the fish to direct sunlight until it got bleached.
Measurements and counts were made following Miller (1948, 1975) and Costa (1988) with some addi-
tions and adjustments to poeciliins, as follows.
Measurements
Standard length: from tip of snout, here always regarded excluding the pre-maxillaries to the posterior-
most limit of the hypural plate visualized through transmitted light. Head length: from tip of snout to posteri-
ormost limit of operculum. Pre-dorsal length: distance between tip of snout and anterior limit of insertion of
the first dorsal-fin ray. Dorsal-fin base: distance from the anterior limit of first dorsal-fin ray and posterior
limit of insertion of last dorsal-fin ray. Pre-pelvic length: distance from tip of snout to posterior limit of inser-
tion of outermost pelvic-fin ray. Pre-anal length (females): distance from tip of snout to anterior limit of inser-
tion of first anal-fin ray. Pre-anal length (males): distance from the snout tip to the anterior limit of the
urogenital papillae. Pelvic fin length: distance between proximal tip of first pelvic ray to distal tip of second
pelvic ray. Gonopodium length: distance from the anterior limit the urogenital papillae to the tip of gonopo-
dium, disregarding the gonopodium palp. Anal-fin lenght (females): distance between anterior limit of inser-
tion of first anal ray and tip of the longest anal-fin ray. Gape: as ‘Mouth, width’ in Miller (1948).
Counts
In every male specimen, seven counts on alcohol-preserved specimens and twelve counts of internal fea-
tures in cleared-and-stained specimens were taken. Females had nine counts on alcohol-preserved specimens
and four counts of internal features in cleared-and-stained specimens. Four additional counts to males and two
to females were taken relative to structure positioning. Counts listed in the text are followed by the number of
times it was observed in parentheses.
Counts differing from Miller (1948, 1975) are as follows. Alcohol-preserved specimens: number of dor-
sal-fin and anal-fin rays (counting each element as a single ray of anterior dorsal-fin for both males and
females and anal-fin of females). Branched caudal-fin rays. Scales in transversal series (counting on the line
starting from the scale on the origin of the dorsal fin, and going to the scale immediately posterior on next lon-
gitudinal series, often ending on the base of the anal fin). Cleared-and-stained specimens: unbranched supe-
rior rays in caudal fin. Unbranched inferior rays in caudal fin. Number of gonapophyses (modified hemal
arches in gonopodial suspensorium of males). Pectoral-fin rays. fin rays of gonopodium (males; counting all
elements). Dentary teeth on the outer row. Number branchiostegal rays. Positioning in relation to vertebrae:
vertebra anterior to insertion of first proximal radial of the dorsal fin. Vertebra on the vertical passing through
the insertion of the first pelvic-fin ray. Vertebra of the first gonapophysis (males). Vertebra on the vertical
passing trough the tip of the gonactinostal complex 2-3-4 (usually where the ligaments containing the ligo-
style attaches).
Institutional abbreviations are listed in Leviton et al. (1985). Nomenclature for general osteological fea-
tures is according to Rauchenberger (1989) and Weitzman (1962). The male anal fin is treated as gonopodium.
Gonopodium lepidotrichia segments are treated just as segments. Gonopodium ornamentation is named after
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NEW PAMPHORICHTHYS
Rosen and Gordon (1953) with the addition of one special structure, the subdistal anterior processes on the
anterior branch of ray 5, which are treated as keel-like segments.
Photographs of structures were taken under an Olympus SZH 10 microscope attached to a Sony CCD-
IRIS camera and a Sony Color video printer. SEM images were taken on the Leo 440 Scanning Electron
Microscope. The specimen prepared for SEM went through conventional clear and staining, then osteological
pieces were made clear of soft tissues and dehydrated after Lindquist and Dillaman (1986).
Pamphorichthys pertapeh, new species
(Fig. 1)
Holotype. MNRJ 32467, male. 16,3 mm SL; Brazil: State of Goiás: Municipality of Formosa: lagoa (lake)
Perta-Pé at the left side of rio Bezerra, tributary of Preto river, tributary of Paracatu river, São Francisco river
basin; Coord. 15°59'02”S 047°11'51”W (Fig. 2); C. A. Figueiredo and D. F. Moraes Jr. 18.Oct.1998.
Paratypes. MNRJ 18090, locality data as holotype, 49 exs. (10 exs. males 16,7–14,2 mm SL, 31 exs.
females 19,4–12,7 mm SL, 07 exs. young 13,7–7,4 mm SL); MZUSP 99750, locality data as holotype, 20 exs.
(09 exs. males 15,5–13,8 mm SL, 08 exs. females 17,2–13,8 mm SL, 3 exs. young 12,1–10,5 mm SL); MCP
41345, locality data as holotype, 20 exs. (09 exs. males 16,8–13,4 mm SL, 07 exs. females 17,4–13,7 mm SL,
04 exs. young 13,4–10,4 mm SL); USNM 393312, locality data as holotype, 20 exs. (08 exs. males 15,9–14,5
mm SL, 07 exs. females 17,9–14,1 mm SL, 05 exs. young 12,9–10,2 mm SL); UF 171129, locality data as
holotype, 20 exs. (09 exs. males 17,0–14,0 mm SL, 07 exs. females 18,7–13,1 mm SL, 04 exs. young 11,8–
09,3 mm SL); UMMZ 248719, locality data as holotype, 20 exs. (08 exs. males 16,5–14,6 mm SL, 08 exs.
females 16,2–14,2 mm SL, 04 exs. young 12,4–10,0 mm SL)
Diagnosis. Pamphorichthys pertapeh is distinguished from all other Poeciliinae species by two derived
characters, unique among poeciliins: (a) bilobed teeth present on dentary and external tooth row of the pre-
maxilar bone (Fig. 3) and (b) distal-most segments of the anterior branch of the fourth gonopodial ray and of
the third gonopodial ray are independently fused into elongated segments (Fig. 4). Other diagnostic characters
that are also found in other species of the tribe Poeciliini are: dark longitudinal stripe on female caudal pedun-
cle; urogenital region of females with heavy dark pigmentation; third pelvic-fin ray of males curved distally
(Fig. 5); and specialized pair of anal scales anterior to the urogenital opening in females.
Description. Morphometric and meristic data are given for males (table 1) and for females (table 2).
Females larger than males.
Dorsal profile of males gently convex from snout to dorsal fin origin, and slightly concave to caudal-fin
base. In females, dorsal profile straight on head, gently curved from occipital base to dorsal-fin origin and
slightly convex from origin of dorsal fin to caudal-fin base. Ventral profile of males convex on head, with a
slight notch between head and pelvis. Pelvic region convex, abruptly angled at the insertion of pelvic fins, pre-
senting a depression between its posterior margin and urogenital papillae, which is attached to the gonopo-
dium. Caudal peduncle convex from gonopodium base to caudal-fin base, with a discrete keel of scales in
some males. Ventral profile of females gently convex from head to anal opening with a slightly notch on the
opercular borders. A depression occurs between the anus and the base of the anal fin, which is thickened on its
first rays. Profile slightly convex from the first rays of the anal fin to the base of the caudal fin. Border of the
dorsal fin in both sexes gently curved on distal extremes. Border of caudal fin in both sexes rounded. Border
of anal fin is slightly rounded in females, with the third ray being longer than the remaining rays. Pectoral fins
rounded and symetrical. Pelvic fins in full-grown males have the second ray elongated and the third ray
curved at its subdistal end. Female pectoral fin rays, without any special mophological features, have the sec-
ond ray being the longest. Gonopodium with a gonopodium palp enveloping its tip and extending beyond the
tips of the gonopodium rays.
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TABLE 1. Morphometric data for males.
TABLE 2. Morphometric data for females.
Dorsal-fin rays i(7) or ii(10)-7(15) or 8(2) (males), i(6) or ii(9)-7(14) or 8(1) (females); anal-fin rays in
females iii(15)-9(15); pelvic-fin rays 5(5); caudal-fin branched rays 11(3); unbranched superior caudal-fin
rays 6(1) or 7(1); unbranched inferior caudal-fin rays 6(2); pectoral-fin rays 10(1) or 11(2); scales in longitu-
Measurements N Holotype Min Max Mean Stdv
Standard Lenght (mm) 17 16.14 14.48 17.2 15.92 0.88
In percents of standard length
Body depth 17 24.5% 23.8% 28.1% 25.9% 1.2%
Depth of caudal peduncle 17 17.7% 15.6% 19.7% 17.5% 1.0%
Lenght of caudal peduncle 17 55.8% 50.4% 57.6% 53.7% 1.9%
Predorsal lenght 17 55.2% 53.2% 57.0% 55.2% 1.2%
Length of dorsal-fin base 17 10.2% 9.5% 12.6% 10.6% 0.8%
Prepelvic lenght 17 33.7% 31.7% 36.2% 34.0% 1.2%
Pelvic-fin lenght 17 20.2% 20.2% 23.7% 22.5% 1.1%
Preanal lenght 17 39.5% 36.8% 44.6% 39.8% 2.1%
Lenght of gonopodium 17 33.5% 33.5% 38.2% 35.7% 1.3%
Head lenght 17 25.0% 24.5% 26.6% 25.8% 0.6%
In percents of head length
Head depth 17 71.3% 66.1% 75.6% 69.5% 2.7%
Head width 17 70.5% 67.5% 78.0% 72.6% 2.6%
Eye diameter 17 38.9% 36.4% 44.3% 39.3% 2.0%
Gape 17 38.9% 26.8% 38.9% 30.7% 3.2%
Measurements n Min Max Mean Stdv
Standard Lenght (mm) 15 15.37 20.0 18.04 1.18
In percents of standard length
Body depth 15 24.9% 29.6% 28.0% 1.3%
Depth of caudal peduncle 15 15.8% 17.7% 16.8% 0.5%
Lenght of caudal peduncle 15 30.8% 37.6% 34.5% 2.4%
Predorsal lenght 15 56.8% 59.7% 58.4% 0.9%
Length of dorsal-fin base 15 8.8% 10.6% 9.8% 0.5%
Prepelvic lenght 15 44.7% 49.4% 47.3% 1.4%
Pelvic-fin lenght 15 9.8% 12.5% 11.3% 0.9%
Preanal lenght 15 57.3% 65.4% 60.5% 1.7%
Lenght of anal-fin 14 17.1% 23.1% 19.1% 1.8%
Head lenght 15 27.2% 29.9% 28.3% 0.8%
In percents of head length
Head depth 15 62.6% 72.1% 67.5% 2.8%
Head width 15 73.2% 84.2% 77.2% 2.7%
Eye diameter 15 32.9% 38.8% 36.4% 1.6%
Gape 15 28.8% 34.0% 31.9% 1.8%
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NEW PAMPHORICHTHYS
dinal series 26(6) or 27(9) (males), 26(3), 27(3), 28(7), and 29(2) (females); scales in transversal series 7(32);
scales around caudal peduncle 15(1) or 16(32); pre-dorsal scales 15(5), 16(12), or 17(8). Pre-caudal vertebra
14(6); caudal vertebra 15(6); number of gonapophyses 2(2); first gonapophysis originating at the 14th(3) verte-
bra; number of gonactinosts 7(3). Premaxilary teeth on the outer row, males: 6(1), females: 8(1) or 10(1);
Dentary teeth on the outer row, males: 6(1) or 7(1); females: 8 (1) or 10(1); first proximal radial of dorsal fin
between 11th and 12th vertebrae (6); branchiostegal rays 5(5).
FIGURE 1. Pamphorichthys pertapeh lateral view. (A) Holotype, MNRJ 32467 male, 16,3 mm SL; (B) paratype, MNRJ
18090, female 19,0 mm SL.
Coloration in alcohol. Cream or pale yellow body background color. Scales with dark chromatophore on
posterior borders, except those from the abdomen, producing a reticulate pattern. Chromatophore density
increased on the dorsal portion of the body, presenting an overall pattern of a darker body seen from above.
Abdomen without any sign of scale reticulation. Ventral portion of caudal peduncle with reticulate pattern
abruptly lighter, but still present. Horizontal septum region is denser in chormatophores and show a thin irreg-
ular stripe more conspicuously posterior to a vertical line passing on the insertion of the first dorsal-fin ray.
Ventral portion of dorsal fin with one or two discontinuous dark transversal stripes consisting of horizontal
blotches present on the intermembranes of the fin. Dorsal-fin distal tip dark in males. Other fins hyaline,
including gonopodium of males. Female urogenital region covered with a very conspicuous dark patch (Fig.
6). Distribution and habitat notes. Known only from type location, a marginal lake on the left side of
Bezerra river, a tributary of the Negro river, tributary of Paracatu river at the São Francisco river basin. Speci-
mens were collected at the lakeshore, which has a sandy bottom, and in clear and shallow waters near grasses
and submerged bushes (Fig. 7).
Etymology. The specific epithet, pertapeh, is an allusion to the type locality, “Lagoa Perta-pé”, which can
be translated into “Squeeze-Foot Lake”. The name of the lake refers to the dense aquatic vegetation present in
the lake, except for its shore, that makes wading almost impossible. A noun in aposition
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FIGURE 2. Geographic position of type locality of Pamphorichthys pertapeh, Perta-Pé lake, Bezerra river, near the
region called “Águas Emendadas”, meaning Mended Waters, referring to the confluence of three major neotropical
watersheds: A. Tocantins river basin; B. Paraná river basin; C. São Francisco river basin. Dashed line represents water-
sheds limits.
Discussion. Species comprising the tribe Poeciliini need to be classified according to their phylogenetic
history, and an extensive phylogenetic analysis of the group is still lacking. Although there have been some
recent studies on the systematics of the tribe (Rodriguez, 1997; Ptacek & Breden, 1998; Breden et al., 1999;
Hamilton, 2001) its phylogenetic relationships, both internally and with sister groups, have yet to be clarified
and many taxonomic revisions are needed. Consequently, generic and supra-generic taxonomy, in many
groups of the subfamily, is still unstable.
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NEW PAMPHORICHTHYS
The species being described here is considered a member of the genus Pamphorichthys. There are cur-
rently no diagnostic characters that allow the inclusion of this species in other available Poeciliini genera, as
defined by Rodriguez (1997) or Rosen and Bailey (1963). Their definitions of the subgenera of Poecilia were
based mainly on superficially examined characters or characters with doubtful polarization.
FIGURE 3. SEM photography of teeth showing outer row of the premaxila with bilobed teeth.
FIGURE 4. Gonopodium of Pamphorichthys pertapeh sp. nov. full view. Scale bar = 1 mm; detailed view in the box of
gonopodium tip. Membranous cap represented on the tip of rays 3 and 4a. Scale bar = 1 µm.
Pamphorichthys pertapeh is proposed as the sister-group of the remaining species of Pamphorichthys and
this relationship is supported by ten shared derived characters: absence of parietal bones, which are also
absent in Poecilia reticulata; shape of the ventral component of the anterior process of the maxilar bone;
abrupt shape of the ventral profile of the premaxilar bone; dentary without a lateral process widening the
insertion area of the teeth, this process is also absent in Micropoecilia in the sense of Meyer (1993) and
Poecilia elegans (Trewavas, 1948); forward bending of the median-ventral process in the dentary; process at
the endopterigoid connecting to the lateral-ethmoid absent; reduced number of pectoral-fin rays (10-11); sec-
ond pelvic-fin ray separated from other rays by a deep notch; ventral longitudinal process of the pelvic bones
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almost imperceptible; number of gonactinosts reduced to seven; gonapophyses parallel to the vertebral spine;
Hollister foramen restricted to the anterior tip of the first gonapophysis; and all cephalic sensorial pores sub-
stituted by pits with exposed neuromasts.
FIGURE 5. Urogenital region of female of Pamphorichthys pertapeh showing the pigmented area around anus and uro-
genital opening. Scale corresponds to 1 mm.
At least five derived characters are shared by Pseudolimia heterandria, Pamphorichthys pertapeh, and the
remaning species of Pamphorichthys: Transverse processes of the parasphenoid not touching the roof of skull;
pre-orbital sensorial canal not completely ossified, as in Poecilia reticulata; reduced number of pectoral-fin
rays (< 12); anterior insertion of pelvic-fin rays; Hollister foramen on the anterior half of the first gonapophy-
sis. Four characters are shared by Limia and Pamphorichthys pertapeh: distal portion of the third pelvic-fin
ray curved, putatively reversed in Pseudolimia heterandria; first gonactinost elongated; median segments on
the fifth gonopodial ray modified with dorsal projections; a pair of modified scales between anal fin and uro-
genital opening of females.
Pamphorichthys pertapeh is known only from its type-locality and its highly endemic status places it as an
endangered, vulnerable species according to IUCN rules (criterion D2; known occurrence restricted to a small
location, i.e. the type-locality). Its description makes it known for biodiversity accounts and conservation
efforts, partly revealing an unsuspected morphological diversity and a high level of endemicity not previously
known among the species of the Pamphorichthys clade.
Comparative material. Poecilia vivipara: MNRJ 16024 (17 exs.), UFRJ 4091 (04 exs.); Poecilia elegans
23392 (04 ex.), UF 25035 (30 ex.); Limia grossidens: USNM 220524 (paratypes, 82ex.); Pseudolimia
heterandria: BMNH 1909.4.2:30-32 (syntypes) (3ex.), CAS 164187; Pamphorichthys araguaiensis: UFRJ
1519 (69ex.); Pamphorichthys hasemani: UFRJ 3646 (131ex.); Pamphorichthys hollandi: MZUSP 47356
(273ex.); Pamphorichthys minor: UFRJ 3944 (500ex.), USNM 120268 (02ex. females); Pamphorichthy
scalpridens: UFRJ 3914 (362ex.);Micropoecilia branneri: MZUSP 54512 (72ex.), MZUSP 76348 (03ex.);
Micropoecilia parae: MZUSP 65428 (117ex.), USNM 66113 (cotype of Acantophacelus melanzonus, 01ex.);
Micropoecilia picta: MZUSP 65423 (98ex.), USNM 151459 (04ex.).
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NEW PAMPHORICHTHYS
FIGURE 6. Pelvic-fin of sexually mature males Pamphorichthys pertapeh. Scale corresponds to 1 mm.
FIGURE 7. Collection site at type-locality.
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Aknowledgments
Thanks are due to Mário de Pinna and other members of the Fish Section at the Museu de Zoologia da Univer-
sidade de São Paulo (MZUSP), São Paulo; and Ricardo Campos-da-Paz of the Universidade Federal do
Estado do Rio de Janeiro, Rio de Janeiro, for their support; Flávio Lima and Paulino Souza for their construc-
tive criticism of earlier versions of the manuscript; Marcelo Pires kindly reviewed my English and made valu-
able comments. This study was partially supported by a Post-Doc grant from the Conselho Nacional de
Desenvolvimento Científico e Tecnológico (CNPQ; process number 155636/2006-5). This study originated
from doctoral work of the author at MZUSP (Doctoral grant FAPESP 99/01048-8)
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