Description of a new species of annual fish of the genus Neofundulus (Cyprinodontiformes: Rivulidae) from the upper rio Mamore basin, Bolivia.
Received: 03 March 2013--Accepted: 17 June 2013Abstract
Neofundulus splendidus n. sp. is described from Bolivia, departamento de Santa Cruz, rio San Miguel, upper Mamore basin. Neofundulus splendidus n. sp. is distinguished from the remaining Neofundulus species by a combination of characters: Large black spot at humeral region with diameter of approximately 1.5-2.0 of pupil in males and females (vs. black spot in humeral region, when present, small--half of pupil), 40-41 scales in longitudinal series (vs. 35-38 in remaining congeners). This is the first record for the genus Neofundulus in rio Mamore basin.
Resumo
Neofundulus splendidus n. sp. e desrcito da Bolivia, departamento de Santa Cruz, rio San Miguel, alto bacia Mamore. Neofundulus splendidus n. sp. destingue das demais especies de Neofundulus pela seguintes combi-nacoes de caracteristicas: Grande mancha preta na regiao de humeral com diametro de aproximadamente 1.5-2.0 do diametro da pupila em machos e femeas (vs. pequena ou ausente mancha preta na regiao de humeral), 40-41 escamas em series longitudinais (vs. 35-38 nas demais especies). Este e o primeiro regiscro para o genero Neofundulus na bacia do rio Mamore.
Zusammenfassung
Neofundulus splendidus sp. n. wird vom Fluss San Miguel aus dem Einzugsgebiet des Mamord-Oberlaufs in Bolivien, Departamento Santa Cruz, beschrieben. Die neue Art unterscheidet sich von anderen Arten der Gattung Neo-fiindulus durch eine Kombination folgender Merkmale: gro[ss]er schwarzer Fleck im Humeralbereich mit etwa dem 1,5- bis 2,0-fachen Durchmesser der Pupille bei Mannchen und Weibchen (im Gegensatz zu einem kleinen schwarzen Fleck im Humeralbereich, der--wenn uber-haupt vorhanden--etwa dem halben Pupillendurchmesser entspricht), 40-41 Schuppen in Langsreihen (35-38 bei den ubrigen Angehorigen der Gattung). Es handelt sich um einen Erstnachweis der Gaming Neofundulus im Rio-Marmore-Einzugsgebiet.
Resume
Neofindulus splendidus n. sp. est decrit, venant de Bolivie, Departamento de Santa Cruz, rio San Miguel, bassin superieur du Mamore. Neofundulus splendidus n. sp. se distingue des autres especes de Neofundulus par une combinaison de caracteristiques: une grande tache noire dans la region humerale avec un diametre d'environ 1,5-2,0 de la pupille pour les males et les femelles (contre une tache noire dans la region humerale, si elle est presente, d'une petite moitie de La pupille), 40-41 ecallies en series longitudinales (contre 35-38 pour les autres congeneres). Il s'agit de la premiere occurrence du genre Neofundulus dans le bassin du rio Mamore.
Sommario
Neofundulus splendidus n. sp. e descritto dalla Bolivia, Departamento de Santa Cruz, Rio San Miguel, bacino superiore del Mamore. Neofundulus splendidus n. sp. si distingue dalle altre specie di Neofundulus per una combinazione di caratteri: presenza sia in maschi sia in feminine di una grande macchia nera in regione omerale con diametro di circa 1.5-2.0 volte quello della pupilla (vs. macchia nera nella regione omerale, quando presente, pic-cola c di diametro meta della pupilla), 40-41 scaglie in serie longitudinale (vs. 35-38 nei congeneri). Ii genere Neofundulus non era stato ancora segnalato nel bacino del Rio Mamore.
INTRODUCTION
Even though the first Neofundulus species N paraguayensis, was described over a hundred years ago (Eigenmann & Kennedy, 1903), few species of the genus are so far known, when compared with other Rivulidae genera, and few studies have been conducted on this genus.
So far, only five Neofundulus species have been described: N. paraguayensis (Eigenmann & Kennedy, 1903) from the rio Paraguay basin, N. ornatipinnis Myers 1935 from the rio Paraguay basin, N parvipinnis Costa, 1988 from the rio Cuiaba basin (rio Paraguay basin), N guaporensis Costa 1988 from the rio Guapore basin, and N acutirostratus Costa 1992 from the rio das Velhas, rio Sao Francisco basin.
Myers (1924) erected Neofundulus for Fundulus paraguayensis, which has as its type locality a pool near the arroyo Trementina Paraguay. Aramburu et al. (1962) studied material of the genus collected in Argentina and verified through meristic analysis and ray fin counts that Neofundulus paraguayensis could not be distinguished from Neofundulus ornatipinnis, which they considered to be a synonym of the earlier.
Costa (1988) revalidated the two species, based on number of rays of dorsal fin, number of scales in transversal series and caudal peduncle and in the form of the fins he considered N. ornatipinnis e N. paraguayensis separate species and described two additional species Neofundulus parvipinnis and Neofundulus guaporensis.
The last Neofundulus species of the genus to be described, N. acutirostratus, was based on a single specimen collected in 1942 by A. L. Carvalho and G. S. Myers (Costa, 1992).
The genus Neofundulus is one of the several annual genera of Rivulidae, that is, they are considered annual fish for living in temporary habitats and resisting desiccation by laying their eggs on the substratum. Annual fish are unique in the fact that they develop their eggs very slowly and can go up through to three diapauses, depending on genetic and environmental factors (Myers 1952; Wourms 1972).
The new species described below was found in a roadside annual pool between the towns of San Miguel and San Juan, departamento de Santa Cruz, in the area near the rio San Miguel, a tributary of the right bank of the rio Mamore, Bolivia.
MATERIAL AND METHODS
Measurements were taken point-to-point under a stereomicroscope with a digital caliper to the nearest 0.1 mm on the left side of the specimen following Costa (1995; 2007). Measurements are expressed as percents of standard length (SL), except the measurement of the head, which are recorded as percents of head length (HL).
Counts of vertebrae and pleural ribs were taken from radiographs of the holotype, five males paratypes and six females paratypes. Terminology for frontal squamation follows Hoedeman (1958) and Costa (2006). For vertebral counts the caudal compound centrum was counted as a single element. Institutional abbreviations follow Sabaj-Perez (2010), with addition of UNITAU (Universidade de Taubate), MNKP-Museo Noel Kempff, Santa Cruz de La Sierra, Bolivia. Osteological features included in the description are those considered phylogenetically informative in studies on Neofundulus (1988, 1992).
Neofundulus splendidus, n. sp.
(Figs 1-2, Table I)
Table I. Morphometric and meristic data for the holotype (H) and
parauypes of Neofimdulus splendidus.
H Paratypes
Male Male n = 5 Females n = 6
Standard length (mm) 63.5 33.9-40.5 33.5-37.1
Percents of standard length
Body depth 66.4 22.1-23.6 18.5-20.5
Caudal-peduncle depth 14.3 13.6-14.5 12.7-14.4
Pre-dorsal length 67.1 73.7-77.0 71.4-75.4
Pre-pelvic length 53.5 52.3-54.0 53.7-57.0
Length of dorsal-fin base 17.8 10.8-13.3 12.7-15.7
Length of anal-fin base 21.2 17.7-23.9 17.1-22.3
Caudal-fin length 18.4 24.2-28.4 22.4-30.4
Pectoral-fin length 21.4 19.8-24.1 15.7-26.9
Pelvic-fin length 11.8 11.3-17.9 11.6-16.1
Head length 24.7 29.3-32.4 28.4-33.9
Percents of head length
Head depth 68.0 57.3-64.1 52.6-62.3
Head width 34.6 32.5-40.7 31.3-35.9
Lower jaw length 17.2 16.9-20.8 17.1-19.4
Eye diameter 28.0 27.9-29.2 26.5-29.4
Counts
Dorsal fin 10 9-10 11-12
Caudal fin 22 22 22-23
Anal fin 14 14-15 15-16
Pelvic fin 07 07 6-7
Pectoral fin 14 12-14 12-14
Meristic
Scales in longitudinal series 40 40-41 40-41
Scales in transversal series 09 09 09
Horizontal scales around 16 16 16
caudal peduncle
Holotype: MNKP-11205, 01 male, 63.5mm SL, Bolivia. Deposited at Museo Noel Kempff, road San Miguel to San Juan, Departamento Santa Cruz, San Miguel city, rio San Miguel, rio Mamore basin (16°46'22.0"S 61°11'12.6'W), altitude 440 m, 20 February 2012, Didier Pillet, Bruno Accorsi, Jean Marc Beltramon, Michel Beuchey& Christine Lambert.
Paratypes: MNKP-11206, five males 33.9-40.5 mm SL and six females, 33.5-37.1 mm SL, Bolivia. Deposited at Museo Noel Kempff, Santa Cruz de La Sierra, ZUEC-7303, one male, 35.7 mm SL and one female, 36.2 mm SL. Deposited at Museu de Zoologia da Universidade de Campinas, Campinas, Sao Paulo, Brazil, collected with the holotype.
Diagnosis: Neofundulus splendidus differs from the remaining species of the Neofundulus genus by the male body with large black spot at humeral region with diameter of approximately 1.5-2.0 of pupil in males and females (vs. black spot in humeral region, when present, small), 40-41 scales in longitudinal series (vs. 35-38 in remaining congeners), by color pattern, except N. guaporensis and N. acutirostratus, consisting in three to four yellow lines in the body, alternated with two to three slender red lines, that become thicker and dark towards the caudal peduncle (vs. yellow lines lacking in N. paraguayensis, a single yellow line in N. ornatipinnis, three yellow lines alternated with three red lines in N. parvipinnis), the color of pectoral fin in males, with finely delicate drawing with narrow white grading to yellow lines within a dark orange background (vs. pectoral fin with black spots in remaining congeners), presence of four to five transverse black bars in dorsal fin in males (vs. black bars absent or presence of small black spots, or else combinations of two black bars with small black spots in the remaining Neofundulus species), three transverse brown bars in the body of females (vs. bars lacking in females of the remaining congeners), transverse brown bars in dorsal and anal fins in females (vs. brown bars absent or presence of small black spots in dorsal fin of the females of remaining Neofundulus species).
Description: Morphometric and meristic data presented in Table I. Largest specimen examined 63.5mm SL. Body elongated, slender, subcylindrical, deeper than wide, compressed posteriorly. Greatest body depth at level of pelvic-fin base. Dorsal profile gently convex from snout to end of dorsal-fin base, approximately straight to slightly concave on caudal peduncle. Ventral profile slightly convex on head, almost straight from anterior portion of abdominal region to end of anal-fin base, slightly concave along caudal peduncle. Greatest body depth at level of pelvic-fin base. Snout very blunt. Jaws long.
Tip of dorsal and anal-fins slightly pointed in male, rounded in female, lacking filamentous rays. Anal fin approximately rectangular in males, longer than deep, reaching vertical through base of caudal fin. Anal-fin rounded, short, and without filamentous rays in females.
Caudal-fin rounded to subtruncated in males and females, without filamentous rays. Pectoral-fin elliptical. Pelvic-fin pointed, tip reaching between base of 3rd and 4th anal-fin rays in male and 1st and 2nd anal-fin rays in female. Pelvic-fin bases in close proximity in males, and separated by small interspace, in females. Dorsal-fin origin on vertical through base of 5th or 6th anal-fin ray, and between neural spines of 22nd and 24th vertebrae. Anal-fin origin between pleural ribs of 17th and 19th vertebrae. Dorsal fin rays 10-13; anal fin rays 14-16; caudal fin rays 2224; pectoral fin rays 12-14; pelvic fin rays 7.
Scales large, cycloid. Body and head entirely scaled, except antero-ventral surface of head. Body scales extending over caudal-fin base; no scales on dorsal and anal fins. Frontal squamation E-patterned; E-scales not overlapping medially; scales arranged in irregular circular pattern around A-scale without exposed margins. Longitudinal series of scales 40-41; transverse series of scales 9; scale rows around caudal peduncle 16. Total vertebrae 34-35. Contact organs absent.
Coloration' in life (Figs 1-2):
Males: Side of body with yellow, red and iridescent clear blue spots longitudinally aligned, becoming olive-green on the back, abdominal area whitish. Large black spot on humeral region with diameter of approximately 1.5-2.0 of pupil. Sides of head olive-green. Iris yellow, with dark, vertical brown bar crossing center of eye. Dorsal fin pale yellow, with four to five transverse dark bars. Anal-fin dark olive-green, large golden yellow bar surrounded by dark pigmentation near the basis, in some specimens seemingly to continue into the caudal-fin. Caudal-fin dark gray, with white and yellow spots irregularly distributed, and distal portion of the fin black. Pelvic fin dark orange, without spots or lines. Pectoral fin with finely delicate drawing composed by narrow white grading to yellow lines within a dark orange background.
Females: Side of body light brown, with three longitudinal brown stripes. Large black spot on humeral region with diameter of approximately 1.5-2.0 of pupil. Dorsum light brown, abdominal region whitish. Sides of head grey, with a pale metallic yellow tinge on opercle. Iris yellow, with pale brown horizontal bar crossing center of eye. Dorsal and anal fins hyaline, with longitudinal brown bars. Caudal-fin pale yellow, with transverse narrow, light brown bars. Pectoral fin yellow with small brown spots. Pelvic fin pale yellow.
Distribution: Neofundulus splendidus is currently only known from its type-locality.
Habitat (Fig. 3): The type-locality lies at the plateau area (440 ma.s.l.), just southeast of Llanos de Mojos, which occupy much of the rio Mamore basin in Bolivia (see Loubens et al., 1992). The pool is located beside the highway, between the cities of San Miguel to San Juan. Water temperature of the pool was 24.6°C, at the depth of 40 cm. In the marginal area of the pool, at the depth of 10 cm, water temperature was 29°C. Neofundulus splendidus was the only fish species present in the pool, where it was found over its entire area. The pool presented clean water, pH 6.4, 0.5 mg/l of dissolved iron (Fe) and electric conductivity 67 µS. Other animals recorded at the pool were Phylomedusa sp. tadpoles, clams and freshwaters crabs. A dense coverage of aquatic vegetation mainly formed by Echinodorus sp. and Nymphaea sp., was present at the pool.
Behavior in captivity: The male initiates the courtship expanding its unpaired fins next to the female, forming with their bodies, angles of 10° or 15°. Sometimes the male touch, with its mouth, the female abdominal region. After the male display, it touches the substratum with its snout, and the couple dives in the substratum. The female lay two or three large-sized (1.5 mm diameter) eggs per day. Males can be sometimes aggressive towards the female, which makes necessary to separate the couple after the end of courtship.
Etymology: In reference to the exuberant colors of the male. An adjective.
DISCUSSION
Neofundulus splendidus can be easily identified from its congeners by the color pattern of the pectoral-fin in males, and by the presence of a large black spot in the humeral region of males and females. Neofundulus was previously found in southern Bolivia in the Paraguay basin (Costa et al. 1996), but this is the first record of the genus for the tic) Mamore basin. Although the genus Neofundulus had already been found in another tributary of the rio Madeira basin, the rio Guapore basin, the new species extends the distribution of the genus and suggests that the genus might be more widespread in other hydrographic basins on the right bank of the Amazon basin.
Other species of annual fish of several genera, Papiliolebias, Spectrolebias, Trigonectes, Moema, Aphyolebias, and Pterolebias were found recently in Bolivia, at the do Mamore, rio Guapore and rio Paraguay basins (Costa 1996, 1998a, Costa et al. 1996, Costa et al. 1997). Since the area searched was relatively small when the total area of these hydrographic basins is considered, and also considering the remoteness of most of the area, it is very likely that further fieldwork will discover additional new rivulid species in this region.
Neofundulus is distinguished from the remaining genera of Rivulidae family by having the following synapomorphies: fourth pectoral radial not expanded ventrally, a subasal longitudinal white-yellowish stripe on the anal fin in males, transverse black bars on caudal fin of females, and black spots on pectoral fin of males (Costa 19986) Neofundulus splendidus presents very narrow yellow lines on the pectoral fins, instead of black spots present on the other Neofundulus species, thus an autapomorphy of this species. Neofundulus splendidus appears to be closer to Neofundulus parvipinnis, since because both species share 14 to 16 anal-fin rays and a color pattern of the males presenting longitudinal yellow stripes.
ACKNOWLEDGEMENTS
We are grateful to Didier Pillet, Bruno Accorsi, Jean Marc Beltramon, Michel Beuchey, and Christine Lambert for senting the type-material, Itamar Martins for support at the laboratory, Flavio C. T. Lima for useful suggestions to the manuscript, Guillaume Dethu for the pictures and Didier Pillet for informations about breeding behavior.
REFERENCES
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COSTA, W. J. E. M. 1992. Descricao de uma nova especie de Neofundulus (Cyprinodontiformes: Rivulidae), da bacia do rio Sao Francisco, Brasil. Revista Brasileira de Biologia 52 (4): 615-618.
COSTA, W. J. E. M. 1995. Pearl killifishes: The Cynolebiatinae. Systematics and Biogeography of a Neotropical annual fish subfamily (Cyprinodontiformes: Rivulidae). TFH, Neptune City, 128 pp.
COSTA, W. J. E. M. 1996. A new Specie of the annual fish genus Pterolebias (Cyprinodontiformes: Rivulidae) from the Rio Mamore basin, Bolivian Amazon. Neotropical Ichthyology, 7: 91-96.
COSTA, W. J. E. M. 1998a. Revision of the Neotropical annual fish genus Plesiolebias (Cyprinodontiformes: Rivulidae). Ichthyological Exploration of Freshwaters, 8: 313-334.
COSTA, W. J. E. M. 1998b. Phylogeny and classification of Rivulidae revisited: Origin and evolution of annualism and miniaturization in rivulid fishes (Cyprinodontiformes: Aplocheiloidei). Journal of Comparative Biology, 3: 33-92.
COSTA, W. J. E. M. 2006. Descriptive morphology and phylogenetic relationship among species of the Neotropical annual killifish genera Nematolebias and Simpsonichthys (Cyprinodontiformes: Aplocheiloidei: Rivuli-dae). Neotropical Ichthyology, 4: 1-26.
COSTA, W. J. E. M. 2007. Taxonomic revision of the seasonal South American killifish genus Simpsonichthys (Teleostei: Cyprinodontiformes: Aplocheiloidei: Rivuli-dae). Zootaxa, 1669: 1-134.
COSTA, W. J. E. M., BARRERA, S. & SARMIENTO, J. 1996. A new species of annual fish genus Pterolebias (Cyprinodontiformes: Rivulidae) from the Rio Mamore basin, Bolivian Amazon. Ichthyological Exploration of Freshwaters 7: 91-95.
COSTA, W. J. E. M., BARRERA, S. & SARMIENTO, J. 1997. Simpsonichthys filamentosus, une nouvelle espece des Lla-nuras Benianas, bassin du Rio Mamore, Bolivia. Revue Francaise d'Aquariologie, 24: 83-86.
EIGENMANN, C. H. & KENNEDY, C. H. 1903. On a collection of fishes from Paraguay with a synopsis of the American genera of cichlids. Proc. Acad. Nat. Sci. Philad., 477-537.
HOEDEMAN, J. J. 1958. The frontal scalation pattern in some groups of tooth carps. Bulletin of Aquatic Biology, 1: 23-28.
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Dalton Tavares Bressane Nielsen (1) and Roger Brousseau (2)
1) Laboratorio de Zoologia, departamento de Biologia, Universidade de Taubate, Pca Marcelino Monteiro 63, CEP: 12030-010, Taubate, SP, Brazil. E-mail: dnielsen@uol.com.br 2) 8345 Bull Mountain Circle, Elk Grove, California 95758, USA. Email: dr_rog@yahoo.com
Key to Neofundu1us species based on Costa 1992, with the addition
of Neofundulus splendidus:
la. Origin of dorsal fin on vertical through base 2
of first or second anal-fin rays
lb. Origin of dorsal fin on vertical through base 4
of fifth or sixth anal-fin rays
2a. Caudal-fin rounded; tip of pelvic fin in males N.
reaching fourth to sixth anal-fin ray; 16 paraguayensis
horizontal scales around caudal peduncle
2b. Caudal-fin truncated; tip of pelvic fin in 3
males reaching first or second ray of anal
fin; 18 to 20 horizontal scales around caudal
peduncle
3a. Lateral profile of the head pointed; caudal N.
peduncle depth 17.0 % SL acutirostratus
3b. Lateral profile of the head rounded; caudal N. ornatipinnis
peduncle depth 12.5-15.0 % SL
4a. 12 rays anal fin N. guaporensis
4b. 14 to 16 rays anal fin 5
5a. 35 to 36 scales in longitudinal series N. parvipinnis
5b. 40 to 41 scales in longitudinal series N. splendidus
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| Author: | Nielsen, Dalton Tavares Bressane; Brousseau, Roger |
|---|---|
| Publication: | aqua: International Journal of Ichthyology |
| Article Type: | Report |
| Geographic Code: | 3BOLI |
| Date: | Jul 19, 2013 |
| Words: | 3285 |
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