Printer Friendly

A new papiliokbias species teleostei: cyprinodontiformes: rivulidae from bolivian amazon.


Papiliolebias francescae, new species, is described based on specimens collected in a seasonal pool in Rio San Pablo drainage, Departamento Santa Cruz, central Bolivia. It differs from its congeners by the following unique features in males: dorsal and anal fins pointed; oblique, chevron-shaped, blue-gray and dark red stripes in close proximity on flanks; proximal half dorsal and anal fin whitish with broad dark red elongated spots; caudal Fin with two or three transverse rows of dark red spots; pupil with anterior pointed extension.


Die neue Art Papiliolebias francescae wird anhand von Ex-emplaren beschrieben, die in einem zeitbegrenzten Turn-pel im Einzugsgebiet des Flusses San Pablo im Departa-mento Santa Cruz in Zentralbolivien gesammelt wurden. Sic unterscheiden sich von den anderen Vertretern der Gattung durch folgende Kennzeichen der Mannchen: gepunktete Riicken- und Afterflosse; dicht stehende schrage, angewinkelte blaugraue und dunkelrote Streifen auf den Seiten; die korpernahe Halfte der Riicken- und Af-terflosse weiglich mit breiten dunkelroten langlichen Flecken; Schwanzflosse mit zwei oder drei Querreihen dunkelroter Flecken; nach vorne spitz geformte Pupille.


Papiliolebi as francescae, nouvelle espece, est decrit sur base de specimens collectes dans une mare tern poraire dans le drainage du Rio San Pablo, Departamento Santa Cruz, en Bolivie centrale. II se distingue de ses congeneres par les caracteristiques uniques suivantes chez les males: dorsale et anale pointues; des lignes obliques, en forme de chevrons, gris bleu et rouge fonce en patron serre sur les flancs; la moitie environ de la dorsale et de l'anale blanchatre avec de larges taches allongees rouge fonce; la caudale avec deux ou trois rangees transversales de taches rouge fonce; la pupille pourvue d'une extension anterieure pointue.


Papiliolebias francescae, nuove specie, e descritta sulla base di esemplari raccolti in una pozza stagionale del bacino del Rio San Pablo, Dipartimento di Santa Cruz, Bolivia centra-lc. Si differenzia dai congeneri per le seguenti caratteristiche uniche nei maschi: pinna dorsale ed anale appuntite; strisce rosso scuro e blu-grigio oblique, ravvicinate c a forma di spina di pesce sui fianchi; meta prossimale della pinna dorsale e pinna anale biancastra con ampie macchic allungate rosso scuro; pinna caudale con due o tre file trasversali di macchie rosso scum; pupilla con appuntita estensione anteriore.


Annual fishes are specialized members of the killifish suborder Aplocheiloidei, with their entire biological cycle occurring in seasonal freshwater swamps of tropical and subtropical areas of Africa, Central and South America (Costa 1995; Wildekamp 2004; DomInguez-Castanedo et al. 2013). Typical annual fish habitats are temporary pools or swamps; the adults die when the habitat dries out and the next generation survives in the form of desiccation-resistant eggs encased in dry mud (Costa 1995; Watters 2009). The eggs hatch after the onset of the rainy season, develop and grow rapidly (Blaiek et al. 2013, Costa 1998a).

Plesiolebiasini is a clade of small annual fishes, inhabiting temporary swamps of central and northeastern South America (Brazil, Bolivia, Paraguay, and Argentina) (Costa 1998a-b; 2007; 2011; Azpelicueta et al. 2011; Nielsen 8z Brousseau 2014). It comprises five genera, namely, Maratecoara Costa, Papiliolebias Costa, Pituna Costa, Plesiolebias Costa, Plesiokbias Costa and Stenolebias Costa, with a total of 22 species. Molecular data are available for only five plesi-olebiasine species (Hrbek 8c Larson 1999; Murphy et al. 1999) but a phylogenetic analysis based on a detailed survey of morphological characters of almost all species of the Plesiolebiasini was presented recently by Costa (2011). The genus Papiliolebias comprises P bitten, from the Chaco region of the Paraguay River basin, Paraguay and Bolivia, P hatinne from Rio Bermejo basin, Salta, Argentina, and P ashIcyae from upper Rio Mamor6 basin, Bolivia.

In February 2014 specimens of a Papiliolebias species was collected in the Rio San Pablo drainage in the Rio Madeira River basin. It became apparent that these specimens represent a species unknown to science which is herein formally described.


Measurements and counts were taken as described in Costa (1995). Measurements were made with a digital calliper, partly under a dissecting microscope, and rounded to the nearest 0.1 mm. If not stated otherwise, measurements are presented as percentages of standard length (SL), except for eye diameter and snout length, which are given as percentage of head length (HL).

Osteological preparations were made according to Taylor & Van Dyke (1985). Nomenclature for cephalic laterosensory series follows Costa (2001). Terminology for frontal squamation follows Hoedeman (1958); the E- scale was determined as that scale adjacent to the anteriormost neuromast of the posterior section of the supraorbital series.

Type material was deposited at the Museo Civico di Storia Naturale "Giacomo Dona"), Genova (MSNG). Papiliolebias francescae, n. sp. (Figs 1-2; Table I)

Holotype: MSNG 57820, male, 21.3 mm SL: Bolivia, Departamento Santa Cruz, N- uflo de Chavez Province, south of San Ramon, seasonal pools in the Rio San Pablo drainage, Rio Madeira River basin, 16[degrees]39,47''S 62[degrees]3119'W, altitude 240 m, Roger Brousseau, Joe Budterman, George Funkner, & Stefano Valdesalici, 20 February 2014.

Paratypes: MSNG 57801, four males, 19.4-21.8 mm SL, three females, 17.8-19.1 mm SL; one male, 20.1 mm SL, C8cS, one female, 19.1 mm SL, C&S, collected with the holotype.

Diagnosis: Papiliolebias francescae differ from the others member of this genus by the following unique features: dorsal and anal fins pointed in males (vs. rounded); oblique, chevron-shaped, blue-gray and dark red stripes in close proximity on flanks of males (vs. never a similar color pattern); proximal half dorsal and anal fin whitish with broad dark red elongated spots (vs. proximal portion black with white spots); caudal fin with two or three transverse rows of dark red spots (vs. never a similar color pattern); pupil with anterior pointed extension (vs. rounded pupil).

Description: Morphometric data are presented in Table I. Largest male examined 21.8 mm SL; largest female examined 19.1 mm SL. Dorsal profile slightly convex from snout to end of dorsal-fin base, slightly concave on caudal peduncle. Ventral profile gently convex from lower jaw to anal-fin origin, nearly straight to slightly concave along caudal peduncle. Body moderately slender, compressed. Greatest body depth at level of pelvic-fin base. Jaws short, snout blunt. Dorsal and anal fins pointed in males, tip reaching to vertical through base of caudal fin in males, dorsal fin slightly pointed, anal fin rounded in females. Caudal fin rounded. Pectoral fins elliptical, posterior margin reaching urogenital papilla in males, reaching pelvic fin base in females. Pelvic fins pointed, with tip reaching between base of 5th and 8th anal-fin ray in males; tip of each pelvic fin reaching base of 1st and 3rd anal-fin rays in females. Pelvic-fin bases separated. Dorsal-fin origin on vertical through base of 7th or 9th anal-fin rays, and between neural spines of 13th and 14th vertebrae. Anal-fin origin between pleural ribs of 11th and 12th vertebrae. Dorsal-fin rays 10-12; anal-fin rays 18-19; caudal-fin rays 22-25; pectoral-fin rays 12-14; pelvic-fin rays 7. Scales large, cycloid. Body and head entirely scaled, except anterior ventral surface of head. Body squamation extending onto anterior 20% of caudal fin; no scales on dorsal and anal-fin bases. Frontal squamation F-patterned; E-scales not overlapping medially; frontal scales circularly arranged around A-scale. Longitudinal series of scales 23-27; transverse series of scales 11-12; scale rows around caudal peduncle 12-15. Cephalic neuromasts: supraorbital 4 + 4, parietal 2, anterior rostral 1, posterior rostral 1, infraorbital 1 +1 + 1, preorbital 1, otic 1, postotic 1, preopercular 1, mandibular 2 or 4. One neuromast on center of each scale of lateral line of trunk, sometimes absent in few scales. Basihyal subtriangular. Urohyal dorsal process branched. Six branchiostegal rays. First branchial arch gill-rakers 1+7-8. Ventral process of posttemporal present. Total vertebrae 26.

Table I. Morphornetric data of the type series of Papiliolebias

                      Holotype  Males (N = 5 including    Female
                                      holotype)           (N=3)

Standard length (mm)    21.3          19.4-21.8         17.8-19.1

Percents of standard

Body depth              32.8          29.2-32.9         25.8-28.4

Caudal peduncle         15.9          14.6-15.9         13.0-15.8

Pre-dorsal length       66.6          65.9-71.1         65.1-67.2

Pre-anal lenght         58.6          55.1-60.9         58.1-60.6

Pre-pelvic lengrh       47.8          43.8-48.6         42.4-47.1

Length of dorsal-fin    17.8          15.9-22.9         15.7-16.2

Length of anal-fin      28.6          27.8-31.6         26.2-27.5

Caudal-fin length       30.0          26.6-32.1         26.9-30.0

Pectoral-fin length     20.6          13.9-20.6         10.6-15.3

Pelvic-fin length       15.0          14.1-16.0         12.3-13.1

Head length             27.6          26.6-30.7         25.6-29.5

Percents of head

Head depth              91.5          75.0-91.5         73.0-93.8

Head width              40.6          34.3-40.6         32.6-36.7

Eye diameter            40.6          31.6-40.6         33.3-36.7

Color in life: Males: Sides of body light blue, with oblique, chevron-shaped, bluish-gray and dark red alternating stripes; dark metallic golden-green blotch on humeral region. Dorsum red brown with small black dots. Sides of head yellowish brown, opercle golden yellow, scale borders brownish; black suborbital bar. Jaws orange. Iris orange, with a black vertical bar through center of eye. Ventral part of the head light blue. Bran-chiostegal membranes red. Dorsal fin white to yellowish white on the basal half, with 4-5 dark red brown broad bars on basal and posterior portion of fin; distal portion dusky grey. Anal fin white to yellowish white on the basal half, with 5-6 dark red brown broad bars on basal and posterior portion of fin; median area reddish; distal anterior portion dark grey blue. Caudal fin yellowish hyaline on proximal portion with two or three transverse rows of pale reddish spots, often fused to form bars; gray hyaline on distal portion. Pectoral fins hyaline. Pelvic fins dusky gray, reddish on proximal portion.

Females: Sides of head and trunk light gray brownish. Dorsum light brown. Lower surface of head light gray to whitish. Opercular region pale golden. Humeral region golden. Black suborbital bar. Jaws light brown. Iris yellow orange, with black bar through center of eye. Unpaired fins hyaline, yellowish medio-basal portion. Paired fins hyaline.

Distribution: Known only from the type locality, seasonal pools in the Rio San Pablo drainage, Rio Madeira basin, central-eastern Bolivia (Fig. 3).

Habitat: Papiliolebias francescae was found in a quite shallow (about 0.40-1.0 m deep), temporary pool about 60 meters long and 10 meters wide, with clear water and litter at the bottom.

Other fish present were an undescribed Trigonectes species and unidentified catfishes and tetras (Fig. 4).

Etymology: Named in honor of Francesca Fontana, wife of one of the discoverers and authors (SV), in recognition to her love, friendship and support over the years.


The genus Papiliolebias was defined by four synapomorphies (Costa 1998a): unpaired fins dark blue and with a white line along distal margin of anal fm in males, nine pelvic fin rays, and a metallic bluish green humeral blotch. Based on characteristic on recently described species the number of pelvic fin rays and unpaired fins coloration are au-tapomorphic for P bitten. We indeed preferred to assign the new species to Papiliolebias instead of creating a new genus, since we consider this choice more conservative.

Additionally to the above the genus Papiliolebias is characterized by: urohyal, dorsal process branched (also in some species of Pituna); third epibranchial, uncinate process, forming an angle of 45-60[degrees] with distal process (also in rivulines but not in plesiolebi-asines); posttemporal, ventral process present; opercular region with regular reticulate pattern formed by iridescent colour on middle of scales contrasting with dark orangish brown pigment along entire scale margins (also in Pituna); and humeral region with iridescent blotch (also in Pituna) (Costa 2011).

Osteologically Papiliolebias francescae differs from 1? bitten P. and P. hatinne by a triangular metapterygoid (vs. rectangular) (condition not known in P. ash-leyae). This unique conditon must be better investigated, because the two analyzed cleared and stained specimens were 19.1-20.1 mm SL, and perharps the anterior vertical portion of this bone that gives its triangular shape in the remaining congenersmight be present in large specimens. Additionally, Papiliolebias francescae differ from P. bitten i by possessing a reduced pattern of cephalic sensory canals.

As indicated in the Diagnosis, Papiliolebias francescae is readily distinguished from its congeners by several unique color pattern features. Additionally, Papiliolebias francescae can be diagnosed from its congeners by the number of caudal-fin rays (22-25 vs. 19-20 in P ashleyae, 25-28 in P. bitten, 20-23 in P hatinne), fewer number of vertebrae (26 vs. 25-28 in I? ashleyae, 27-30 in P. bitten, 26-28 in P. hatinne), more scales in transverse series (11-12 vs. 9 in P. ash-kyae), and more anal-fin rays (18-19 vs. 15-16 in P. ashleyae, 17-18 in P. bitten).

An additional, undescribed Papiliolebias species with pointed margin of dorsal and anal fin is known in the region around Trinidad in the Rio Mamore drainage but its description is not the aim of the present work (see Habliitzel, 2012 for details on this un-described species).


We thank Joe Bulterman and George Funkner for their assistance with the field work, Jacopo Valdesali-ci and Chiara Valdesalici for help with the pictures, and Pascal Hablatzel for the many suggestions. Flavio C. T. Lima and the anonymous reviewers provided constructive comments on the manuscript. We are grateful to Giuliano Doria (MSNG), for access to the collections under his care.


AZPELICUETA, M. M., BUT1, C. & GARCIA, G. 2009. Papil-iolebias hatinne, a new annual fish species (Cyprinodontif-ormes: Riyulidae) from Salta, Argentina. Revue Suisse de Zookie 116 (3-4): 313-323.

BLAZEK, R., POLA IK, M. 8c REICHARD, M. 2013. Rapid growth, early maturation and short generation time in African annual fishes. EvoDevo 4: 1-7.

COSTA, W. J. E. M. 1995. Pearl killifishes, the Cynolebiatinae: systematics and biogeography of the neotropical annual fish subfamily (Cyprinodontifirmes: Rivulidae). Neptune City, TFH, 128 pp.

cosTA, W. J. E. M. 1998a. Phylogeny and classification of Rivulidae revisited: origin and evolution of annualism and miniaturization in rivulid fishes. Journal of Comparative Biology 3: 33-92.

COSTA, W. J. E. M. 1998b. Revision of the neotropical annual fish genus Plesiolebias (Cyprinodontiformes: Rivuli-dae). Ichthyological Exploration of Freshwaters 8: 313-334.

COSTA, W. J. E. M. 2001. The neotropical annual fish genus Cynalebias (Cyprinodontiformes: Rivulidae): phylogeneti-crelationships, taxonomic revision and biogeography. Ichthyological Exploration of Freshwaters 12: 333-383.

COSTA, W. J. E. M. 2007. Taxonomy of the plesiolebiasine killifish genera Pituna, Plesiolebias and Maratecoara (Teleostei: Cyprinodontifomes: Rivulidae), with descriptions of nine new species. Zoottaa 1410: 1-41.

COSTA, W. J. E. M. 2011. Comparative morphology, phylogenetic relationships, and historical biogeography of plesi-olebiasine seasonal killifishes (Teleostei: Cyprinodontif-ormes: Rivulidae). Zoological Journal of the Linnean Society 162: 131-148.

DOMINGUEZ-CASTANEDO, 0., MOSQUEDA-CABRERA, M. A. & VALDESALICI, S. 2013. First observations of annualism in Millerichthys robustus (Cyprinodontiformes: Rivulidae) Ichthyological Exploration of Freshwaters 24: 15-20.

HABLOTZEL, P.I. 2012. Itenez- und Moxos-Karpfling, zwei neue Killifische aus Bolivien. Die Aquarien- und Terrarien-zeitschrift 3: 62-65.

HOEDEMAN, J. J. 1958. The frontal scalation pattern in some groups of toothcarps (Pisces, Cyprinodontiformes). Bulletin of Aquatic Biology 1: 23-28.

HRBEK, T. & LARSON, A. 1999. The evolution of diapause in the killifish Family Rivulidae (Atherinomorpha, Cyprinodontiformes): A molecular phylogenetic and biogeographic perspective. Evolution 53: 1200-1216.

MURPHY, W. j., THOMERSON, J. E. & COLLIER, G. E. 1999. Phylogeny of the neotropical killifish family Rivulidac (Cyprinodontiformes, Aplocheiloidei) inferred from mitochondrial DNA sequences. Molecular and Phylogenetic Evolution 13: 289-301.

NIELSEN, D. B. T. & BROUSSEAU, R. 2014. Description of a new annual fish, Papiliolebias ashleyae (Cyprinodontifor-mes: Rivulidae) from the upper Rio Mamore basin, Bolivia. aqua. International Journal of Ichthyology 20 (1): 53-59.

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: 107-109.

WATTERS, B. R. 2009. The ecology and distribution of Nothobranchius fishes. Journal of the American Killifish Association 42: 37-76.

WILDEKAMP, R. H. 2004. A world of killies. Atlas of the oviparous cyprinodontOrm fishes of the world Volume 4. The American Killifish Association, Elyria, 398 pp.

Stefano Valdesalici (1) and Roger Brousseau (2)

(1.) Via Ca Bertacclii 5, 42030 Viano (RE), Italy. E-mail:

(2.) 8345 Bull Mountain Circle, Elk Grove, California 95758, USA. E-mail:
COPYRIGHT 2014 Aquapress Publisher
No portion of this article can be reproduced without the express written permission from the copyright holder.
Copyright 2014 Gale, Cengage Learning. All rights reserved.

Article Details
Printer friendly Cite/link Email Feedback
Author:Valdesalici, Stefano
Publication:aqua: International Journal of Ichthyology
Date:Jul 29, 2014
Previous Article:Guidelines for authors.
Next Article:Cirrhilabrus squirei, a new wrasse perciformes; labridae from the great barrier reef and coral sea, australia.

Terms of use | Privacy policy | Copyright © 2021 Farlex, Inc. | Feedback | For webmasters