200 Accepted by M. R. de Carvalho: 19 Jul. 2013; published: 29 Jul. 2013

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2013 Magnolia Press

Zootaxa 3693 (2): 200–206 www.mapress.com/zootaxa/ Article

http://dx.doi.org/10.11646/zootaxa.3693.2.6http://zoobank.org/urn:lsid:zoobank.org:pub:0EB0518C-BDFC-4EFC-9FEE-35B535579E90

A new species of Lycengraulis Günther, 1868 (Clupeiformes: Engraulinae) from the Amazon basin, Brazil, with comments on Lycengraulis batesii (Günther, 1868)

MARINA VIANNA LOEB1 & AYDA VERA ALCÂNTARA2

1Museu de Zoologia da Universidade de São Paulo. Avenida Nazaré, 481, 04263-000. São Paulo, SP, Brazil. E-mail: loeb.mv@gmail.com2Universidade Federal de Sergipe. Cidade Universitária, 49100-000. São Cristovão, Sergipe, Brazil. E-mail: ayda.ufs@gmail.com

Abstract

A new species of Lycengraulis from the Amazon basin is described. Lycengraulis figueiredoi can be distinguished from L. grossidens by a short upper maxilla, its posterior margin not reaching the lower maxilla joint (vs. upper maxilla longer, its posterior margin reaching to or a little past of mandible joint). Lycengraulis figueiredoi can be distinguished also from L. poeyi by 26 to 31 anal-fin rays (vs. 21 to 23 anal-fin rays) and by 44 to 46 vertebrae (vs. 42 vertebrae), and from L.batesii by having the anal-fin origin at vertical through base of second to fifth dorsal-fin ray (vs. anal-fin origin at vertical through base of sixth to 10th dorsal-fin ray) and by 17-21 gill rakers on the lower branch of first gill arch (vs. 12–15 gill rakers on the lower branch of first gill arch). The new species occurs in the rio Purus, Negro, Trombetas and Solimões, in the Amazon basin, Brazil.

Key words: Engraulinae; Neotropical fish fauna; Freshwater fish; Systematics; Sardines

Resumo

Uma nova espécie do gênero Lycengraulis da bacia Amazônica é descrita. Lycengraulis figueiredoi distingue-se de de L. grossidens pela presença de maxila superior curta, sua margem posterior não alcançando a articulação mandibular (vs. maxila superior longa, sua margem posterior alcançando ou ultrapassando a articulação mandibular). Lycengraulis figueiredoi pode ser distinguida de L. poeyi por de 26 a 31 raios na nadadeira anal (vs. 21 a 23 raios na nadadeira anal) e por de 44 a 46 vértebras (vs. 42 vértebras) e de L. batesii pela origem da nadadeira anal em vertical sob a base do segundo ao quinto raio da nadadeira dorsal (vs. origem da nadadeira anal sob a base do sexto ao décimo raio da nadadeira dorsal) e por 17–21 rastros branquiais no ramo inferior do primeiro arco branquial (vs. 12–15 rastros branquiais no ramo inferior do primeiro arco branquial). A nova espécie ocorre nos rios Purus, Negro, Trombetas e Solimões, na bacia Amazônica, Brasil.

Introduction

Lycengraulis Günther, 1868 comprises medium to large sized fishes (150–200 mm standard length) of marine, estuarine or freshwater habits distributed in South and Central America (Whitehead et al. 1988). According to Whitehead et al. (1988) the genera is distinguished in the Engraulinae sensu Nelson (2006) by the presence of enlarged, well-spaced canine-like teeth in the lower jaw. Species of the genus also have 12–27 gill rakers on the lower branch of first gill arch and 21–30 branched anal-fin rays.

According to Eschmeyer and Fricke (2013), four species of Lycengraulis are valid: L. grossidens (Spix & Agassiz, 1829), a marine species that occurs along Atlantic coast of Central and South America, L. batesii (Günther, 1868), that is widespread in the Amazon and Orinoco basins, rio Manacacias in Colombia and coastal drainages of the Guianas, L. poeyi (Kner, 1863), a marine species that occurs on the Pacific coast of Central America, and L. limnichthys Schultz, 1949, a species whose distribution is restricted to Lake Maracaibo basin,

Zootaxa 3693 (2) © 2013 Magnolia Press · 201A NEW SPECIES OF LYCENGRAULIS FROM BRAZIL

Venezuela (Whitehead et al., 1988). However, Lycengraulis has not been thoroughly reviewed recently and the number of valid species of the genus is uncertain, with some notes in Whitehead et al. (1988) suggesting several possible synonymies.

Alcântara (1973) described a new species of Lycengraulis from Amazon basin in her unpublished Msc. thesis. Based on additional material collected since 1973, the new species is herein described.

Material and methods

Measurements and counts were taken as described by Loeb (2012), except cheek length, measured from the midpoint of vertical through posterior margin of orbit to mandibular joint; maxilla length, measured from its anterior to its posterior tip; postorbital distance, the shortest distance between the vertical through posterior margin of orbit and the vertical through the posterior margin of the head; gill raker length, measured from base to distal tip of the raker inserted in the angle of first gill arch.

Standard length (SL) is expressed in mm, and other measurements are expressed as percentage of SL or head length. Measurements and counts were taken on the left side of the specimen whenever possible. Vertebral counts were performed according to Whitehead & Teugels (1985) on eight radiographed specimens.

Institutional abbreviations are ANSP, Academy of National Sciences, Natural History Museum (Philadelphia, USA); INPA, Instituto Nacional de Pesquisas da Amazônia (Manaus, Brazil); MNRJ, Museu Nacional, Universidade Federal do Rio de Janeiro (Rio de Janeiro, Brazil) and MZUSP, Museu de Zoologia da Universidade de São Paulo (São Paulo, Brazil).

Results

Lycengraulis figueiredoi, sp. n.(Fig. 1, Tab. 1)

Lycengraulis amazonensis Alcântara, 1972:19 [not a valid name, described as new in Alcântara's unpublished master thesis].—Whitehead et al., 1985:391 [erroneously considered a junior synonym of Lycengraulis grossidens].

Holotype. MZUSP 6032, 129.80 mm SL. Brazil, Amazonas, lago Beruri, rio Purus, 3°50'S 61°20'W, 7–8 April 1967, Expedição Permanente da Amazônia.

Paratypes. All from Brazil. ANSP 193934 (2, 64.9–78.8 mm SL), INPA 39522 (2, 63.6–77.0 mm SL), MNRJ 40719 (2, 66.32–81.86 mm SL): Pará, rio Trombetas, Oriximiná, 1°46'S 55°52'W, 12–15 October 1967, Expedição Permanente da Amazônia. MZUSP 6363 (1, 101.2 mm SL), Amazonas, lago Castro, rio Purus at mouth, 7–8 April 1967, Expedição Permanente da Amazônia. MZUSP 9815-21 (5, 112.11–123.24 mm SL), Amazonas, lago Beruri, rio Purus, 7–8 April 1967, Expedição Permanente da Amazônia. MZUSP 6749 (1, 118.6 mm SL), Amazonas, rio Negro at Manaus neighborhood, 15–23 November 1967, Expedição Permanente da Amazônia. MZUSP 7658 (1, 152.4 mm SL), Amazonas, lago José Açu at mouth, Parintins, 11–12 December 1967, Expedição Permanente da Amazônia. MZUSP 9822 (15, 52.2–78.6 mm SL), Pará, rio Trombetas, Oriximiná, 1°46'S 55°52'W, 12–15 October 1967, Expedição Permanente da Amazônia. MZUSP 5537 (1, 169.6 mm SL), Pará, Lago Jacupá, Oriximiná, February 1967, Expedição Permanente da Amazônia.

Diagnosis. Lycengraulis figueiredoi can be distinguished from L. grossidens by a short upper maxilla, its posterior margin not reaching the lower maxilla joint (vs. upper maxilla longer, its posterior margin reaching to or a little past of mandible joint). Lycengraulis figueiredoi can be distinguished also from L. poeyi by 26 to 31 anal-fin rays (vs. 21 to 23 anal-fin rays) and by 44 to 46 vertebrae (vs. 42 vertebrae), and from L. batesii by having the anal-fin origin at vertical through base of second to fifth dorsal-fin ray (vs. anal-fin origin at vertical through base of sixth to 10th dorsal-fin ray) and by 17–21 gill rakers on the lower branch of first gill arch (vs. 12–15 gill rakers on the lower branch of first gill arch).

Description. Morphometric data of the holotype and 30 paratypes are presented in Table 1. Body elongate, laterally compressed; greatest body depth at vertical through dorsal-fin origin. Dorsal body profile gently convex from tip of snout to vertical through posterior margin of posterior nostril, convex from latter point to the top of the

LOEB & ALCÂNTARA202 · Zootaxa 3693 (2) © 2013 Magnolia Press

head, approximately straight from the top of the head to dorsal-fin origin, straight and downward along dorsal-fin base, approximately straight along caudal peduncle. Ventral body profile gently convex from anterior tip of lower jaw to posterior margin of orbit, convex from latter point to the posterior end of anal-fin base, nearly straight along caudal peduncle. Caudal peduncle longer than deep. Vertebrae 44–46 (n = 8).

TABLE 1. Morphometrics and meristics of Lycengraulis figueiredoi, new species. SD = Standard Deviation, frequency in parenthesis.

Morphometrics Holotype n Range Mean SD

Standard length (mm) 129.8 30 52.2 169.6 85.6 29.5

Head length (mm) 30.2 30 12.5 41.1 20.9 6.8

Percentages of standard length

Body depth 20.6 30 19.1 24.4 21.7 1.3

Caudal peduncle depth 8.8 30 8.2 11.2 9.9 0.6

Dorsal-fin base length 11.2 30 10.0 13.9 12.0 0.8

Anal-fin base length 22.8 30 25.6 31.0 27.3 1.1

Pectoral fin length 19.7 30 13.4 21.6 18.2 2.3

Pelvic fin length 10.5 30 9.0 13.5 10.7 1.2

Prepelvic length 40.9 30 38.0 45.2 42.4 1.7

Prepectoral length 24.3 30 21.7 26.5 24.7 1.2

Predorsal length 55.3 30 55.2 86.2 59.0 5.3

Preanal length 56.8 30 57.5 62.6 60.5 1.5

Head length 23.3 30 22.5 26.8 24.6 0.9

Percentages of head length

Cheek length 47.7 30 35.3 48.8 39.8 3.7

Snout length 14.6 30 11.0 15.5 12.7 1.2

Orbital diameter 27.8 30 19.2 27.9 24.6 2.5

Upper jaw length 74.8 30 67.7 78.2 71.3 2.8

Postorbital distance 63.0 30 57.1 66.8 62.7 2.6

Gill raker angle on1st gill arch length

11.9 30 9.0 15.7 11.9 2.0

Meristics Holotype n Range

Dorsal-fin rays ii+11 30 13(2),14(16),15(11),16(1)

Anal-fin rays iii+23 25 26(1),27(1),28(7),29(6),30(8),31(2)

Pectoral-fin rays i+14 30 14(22),15(8)

Pelvic-fin rays i+6 30 7(30)

Vertebrae - 8 44(1),45(6),46(1)

Gill rackers on1st gill arch

Upper branch 14 30 13(2),14(21),15(7),16(1)

Lower branch 20 30 17(4),18(9),19(13),20(4),21(1)

Total 34 30 31(5),32(4),33(14),34(6),35(1),36(1)

Zootaxa 3693 (2) © 2013 Magnolia Press · 203A NEW SPECIES OF LYCENGRAULIS FROM BRAZIL

FIGURES 1. Lycengraulis figueiredoi, holotype, MZUSP 6032, 67.6 mm SL, Amazonas, lago Beruri at rio Purus, Brazil.

FIGURES 2. Lycengraulis batesii, syntype, BMNH 1849.11.8.73-75, approximately 120 to 190 mm SL, Pará, rio Pará, Brazil.

Conical-shaped head. Snout short, conical in lateral view. Mouth sub-terminal, inclined downward relative to body axis. Upper maxilla short and thin, posterior margin rounded, extending beyond the vertical through posterior margin of orbit, not reaching the mandible joint. Premaxilla, maxilla and dentary bearing one row of teeth. Teethenlarged, well-spaced canine-like and slightly recurved at premaxilla and maxilla. Dentary with larger teeth intercalated with smaller ones.

Eye lateral on head, located dorsal to horizontal through pectoral-fin insertion; eye visible in dorsal and ventral views. Two confluent nostrils on each side of the head, the anterior elliptical and the posterior crescent-sharped. Perforated scale on the lateral line absent. Cicloid scales higher than large, deciduous.

Dorsal-fin rays ii–iii+11–13 (n = 31), its origin closer to caudal-fin base than to tip of snout; dorsal-fin distal margin approximately straight in juveniles, slightly convex in adults. Anal-fin rays ii–iii+24–28 (n = 26), its origin at vertical through base of second to fifth dorsal-fin rays, anterior rays longer. Pectoral-fin rays i+13–14 (n = 31);

LOEB & ALCÂNTARA204 · Zootaxa 3693 (2) © 2013 Magnolia Press

distal tip of longest pectoral-fin ray reaching pelvic-fin origin; pectoral-fin distal margin slightly convex. Pectoral-fin axillary scale longer than half of pectoral-fin length, absent in most specimens, probably lost during collecting or storage. Pelvic fin-rays i+6 (n = 31), its pelvic-fin origin in midpoint of distance from pectoral-fin origin to anus; distal tip of longest pelvic-fin ray not reaching anal-fin origin; pelvic-fin distal margin slightly convex. Pelvic-fin axillary scale smaller than pectoral fin, absent in most specimens, probably lost during collecting or storage. Pelvic-fin origin closer to the base of the anteriormost pectoral-fin ray than to anal-fin origin. Caudal-fin rays 8,10,9,7, forked, lobes similar in size. Dorsal and ventral procurrent caudal-fin rays 8,7 respectively (n = 3 C&S).

Pseudobranch present, shorter than eye. Gill rakers on the first branchial arch long and thin in juveniles, short and thick in adults, 13–16 (n = 31) on the upper branch, 17–21 (n = 31) on the lower branch, 31–36 (n = 31) in total. Length of raker inserted in the angle of first gill arch smaller than orbital diameter.

Coloration in alcohol. Body coloration pale to light yellow, juveniles with lighter coloration. Longitudinal stripe brownish to silver, absent in most preserved specimens, when present extending from the posterior margin of head to caudal peduncle; width of the stripe of about 20% of body depth at vertical through pectoral-fin origin. Caudal-fin borders and gill cavity darkened. Dark spots dorsally on the head, along dorsal portion of body, along dorsal margin of longitudinal line and along base of dorsal and caudal fins. Orbit and middle portion of the postorbital region translucent to pale.

Etymology. The specific name was given in honor of José Lima de Figueiredo, a Brazilian ichthyologist from the Museu de Zoologia da Universidade de São Paulo who contributed significantly to the study of fish systematics in Brazil.

Distribution. Lycengraulis figueiredoi is known only from lago Castro and lago Beruri, in the drainage of rio Purus; rio Negro; lago José Açu at Parintins; lago Jacupá and rio Trombetas at Oriximiná, middle Amazon basin, Amazonas and Pará State, Brazil.

FIGURES 3. Distribution map of Lycengraulis figueiredoi. Holotype indicated by a star.

Zootaxa 3693 (2) © 2013 Magnolia Press · 205A NEW SPECIES OF LYCENGRAULIS FROM BRAZIL

Discussion

No consistent hypotheses on the relationships among the species of Lycengraulis have been published so far. The genus also has a large number of nominal species and synonymies. Nevertheless, some similarities between L. figueiredoi and L. batesii suggest that they are related. Both species occur in sympatry in the Amazon basin.

Lycengraulis batesii was described from the lower Amazon basin in Pará, Brazil, and occurs in freshwater or brackwish environments at Amazon and Orinoco basins. Lycengraulis figueiredoi and L. batesii share similarities in dorsal and ventral body profiles, body and head shape, teeth number and shape, dorsal, anal, pectoral and pelvic-fin shapes and number of rays, positioning of anal-fin origin and coloration in alcohol. However, both species can be differentiated by the number of gill rakers in first gill arch, a widely used character in Engraulidae systematics (Carvalho, 1950; Alcântara, 1973; Whitehead et al. 1988).

Alcântara (1973) believed that the species that is herein described as L. figueiredoi was also diagnosed by short maxilla, not reaching the lower jaw joint (vs. long maxillar, reaching the mandibular joint in L. batesii). However, analysis of a greater amount of specimens, revealed that this feature is highly variable in L. batesii.

Comparative material

Lycengraulis batesii: MZUSP 9752-6 (5, 127.2–132.8 mm SL), Amazonas, Coari, rio Solimões, 28 September 1968, Expedição Permanente da Amazônia. MZUSP 6750 (1, 113.2 mm SL), Amazonas, rio Negro at Manaus neighborhood, 15–23 November 1967, Expedição Permanente da Amazônia. MZUSP 5933 (1, 116.4 mm SL), Amazonas, lago Jacaré at rigth margin of rio Solimões upstream Manacapuru, 29–31 March 1967, Expedição Permanente da Amazônia. MZUSP 5494 (1, 107.6 mm SL), Pará, Oriximiná, rio Trombetas, February–March 1967, Expedição Permanente da Amazônia. MZUSP 9746-7 (2, 96.0 and 97.9 mm SL), Amazonas, rio Solimões close to Ilha de Bururuá, upstream mouth of rio Jutaí, 15 November 1968, Expedição Permanente da Amazônia. MZUSP 9769-75 (7, 37.5–81.0 mm SL), Pará, Oriximiná, rio Trombetas, 12–15 October 1969, Expedição Permanente da Amazônia. MZUSP 9749-50 (7, 151.1–152.5 mm SL), Amazonas, Fonte Boa, rio Solimões, 25 October 1968, Expedição Permanente da Amazônia. MZUSP 9742 (1, 124 mm SL), Roraima, rio Branco em Boa Vista, 12 February 1969, Expedição Permanente da Amazônia. MZUSP 9813-4 (1, 136.7–138.4 mm SL), Pará, Vila Santana, rio Capim, 14–23 August 1970, Expedição Permanente da Amazônia. MZUSP 9744-5 (2, 119.4 and 129.2 mm SL), Amazonas, rio Solimões, 21 September 1968, Expedição Permanente da Amazônia. MZUSP 9748 (1, 145.5 mm SL), Amazonas, Fonte Boa, rio Solimões, 6 October 1968, Expedição Permanente da Amazônia. MZUSP 9743 (1, 115.8 mm SL), Amazonas, Santo Antônio do Içá, rio Solimões, 17 October 1968, Expedição Permanente da Amazônia. Lycengraulis grossidens: MZUSP 9845-51 (7, 121.3–153.7 mm SL), Rio de Janeiro, Baía da Ilha Grande, May 1966, Expedição do Departamento de Zoologia (Museu de Zoologia)/IOUSP. Lycengraulis poeyi: CAS 233697 (1, 145.1 mm SL), Costa Rica, Gulf of Nicoya, Caballo Is., Erdman Cove, 22 Nov 1952, Peterson C. et al. CAS 233699 (2, 80.5–143.7 mm SL), Costa Rica, Gulf of Nicoya, Chira Flats., 06 Mar 1952, Peterson C. et al..

Acknowledgments

We thank the staff and studentes of the ichthyology lab at MZUSP for their help and collaboration. We also thank E. Baena, M. Marinho and J. Birindelli for helping with illustrations and comments on the manuscript. We owe special thanks to J. Maclaine (BMNH-NHM) for provide the photograph of the syntype of L. batesii, to M. Pastana and F. Bockmann (USP-RP) for taking the radiographs of Lycengraulis figueiredoi, to M. Marinho for providing measurements and counts of L. poeyi at CAS and to D. Catania and J. Fong (CAS) for taking the radiographs of L. poeyi. Marina Loeb was funded by Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP 2011/06830-0).

LOEB & ALCÂNTARA206 · Zootaxa 3693 (2) © 2013 Magnolia Press

Literature cited

Alcântara, A.V. (1973) Espécies amazônicas do gênero Lycengraulis Günther (Pisces, Engraulidae). Dissertação de Mestrado. Universidade de São Paulo, São Paulo, 58 pp.

Carvalho, J.P. (1950) Engraulídeos brasileiros do gênero Anchoa. Boletim do Instituto Paulista de Oceanografia, 1(2), 43–71. http://dx.doi.org/10.1590/s0100-42391950000200004

Di Dario, F. (2009) Chirocentrid as engrauloids: evidence from suspensorium, branchial arches, and infraorbital bones (Teleostei, Clupeomorpha). Zoological Journal of the Linnean Society, 156, 363–383. http://dx.doi.org/10.1111/j.1096-3642.2008.00472.x

Eschmeyer, W.N. & Fricke, R. (2013) Catalog of Fishes electronic version. Available from: http://research.calacademy.org/research/ichthyology/catalog/fishcatmain.asp (Accessed 15 February 2013).

Fowler, H.W. (1915) The fishes of Trinidad, Grenada and St. Lucia, British West Indies. Proceedings of the Academy of Natural Sciences of Philadelphia, 67, 520–546.

Fuster de Plaza, M.L. (1962) Una nueva especie de anchoa de las aguas argentinas Lycengraulis simulator (Pisces: Engraulidae). Physis, 23, 1–9.

Günther, A. (1868) Catalogue of the fishes in the British Museum. Trustees, British Museum, London, 512 pp. http://dx.doi.org/10.5962/bhl.title.8314

Günther, A. (1874) Descriptions of new species of fishes in the British Museum. Annals and Magazine of Natural History, 14, 453–455. http://dx.doi.org/10.1080/00222937408681007

Hildebrand, S.F. (1943) A review of the American anchovies (family Engraulidae). Bulletin of the Bingham Oceanographic Collection of Yale University, 8, 1–165.

Kner, R. (1863) Eine Uebersicht der ichthyologischen Ausbeute des Herrn Professors Dr. Mor. Wagner in Central-Amerika. Sitzungsberichte der Koningl. Bayerischen Akademie der Wissenschaften, München, 2, 220–230.

Kullander, S.O. & Ferraris-Jr., C.J. (2003) Family Engraulididae. In: Reis, R.E., Kullander, S.O. & Ferraris-Jr., C.J. (Eds.), Check List of Freshwater Fishes of South and Central America. Edipucrs, Porto Alegre, pp. 39–42.

Loeb, M.V. (2012) A new species of Anchoviella Fowler, 1911 (Clupeiformes, Engraulidae) from the Amazon basin, Brazil. Neotropical Ichthyology, 10(1),13–18. http://dx.doi.org/10.1590/s1679-62252012000100002

Nelson, J.S. (2006) Fishes of the world. John Wiley & Sons, New York. i–xix + 1–106 pp.Nizinski, M.S.T. & Munroe, T.A. (2002) Engraulidae. In: Carpenter, K.E. (Ed.), The living marine resources of the Western

Central Atlantic. v. 2: Bony fishes. Part 1. FAO Species Identification Guide for Fisheries Purposes and American Society of Ichthyologists and Herpetologists Special Publications No.5, Rome, pp. 764–794.

Schultz, L.P. (1949) A further contribution to the ichthyology of Venezuela. Proceedings of the United States National Museum, 99, 1–211. http://dx.doi.org/10.5479/si.00963801.99-3235.1

Whitehead, P.J.P., Nelson, G.J. & Wongratana, T. (1988). FAO species catalogue. Clupeoid fishes of the world (Suborder Clupeoidei). Part2 - An annotated and illustrated catalogue of the herrings, sardines, pilchards, sprats, shads, anchovies and wolf-herrings. FAO Fisheries Synopsis vol.7, Rome, 274 pp.

Whitehead, P.J.P. & Teugels, G.G. (1985) The west African pygmi herring Sierrathrissa leonensis: general features, visceral anatomy, and osteology. American Museum Novitates, 2835, 1–44