A new species of the genus Lile (Clupeiformes: Clupeidae) of the eastern tropical Pacific.
Resumen.--Se describe una especie nueva de sardina tropical que pertenece al genero Lile. Difiere de los otros miembros de este genero por su morfometria, morfologia, numero de branquiespinas, coloracion y patron de distribucion geografica. Esta es la tercera especie, de este genero, que se conoce en los litorales del Pacifico oriental tropical.
The Clupeidae, order Clupeiformes, is divided into five subfamilies. One of them, Clupeinae, contains 15 genera and almost 80 species widely distributed through temperate, subtropical and tropical seas of the world's oceans (Whitehead 1985). The genus Lile Jordan and Evermann 1896 of subfamily Clupeinae, is exclusive to tropical and subtropical waters of Middle and South America. Besides the new species described here, three other Lile species are known: (1) L. stolifera (Jordan and Gilbert 1882), with a disjunct distribution from Bahia Magdalena, Baja California Sur (BCS) and Golfo de California to northern Nayarit, Mexico, and from Costa Rica to Peru; (2) L. piquitinga (Schreiner and Miranda-Ribeiro 1903) which inhabits the fresh and brackish waters of the Atlantic drainage of Venezuela and Brazil; and (3) L. gracilis Castro-Aguirre and Vivero 1990, known from Jalisco, Mexico to Golfo de Fonseca, Honduras.
All body measurements and counts of fin rays, gill rakers, ventral scutes, and midlateral scales were taken based on Whitehead's methods (1985). Original morphometric data were transformed to thousandths of the cephalic length or standard length and descriptive statistics (mean, standard deviation, mode, and range) were computed. An analysis of discriminant function (Statistica 4.2 program) was used to identify the most important biometric characters to classify Lile nigrofasciata and L. stolifera.
Abbreviations of the fish collections where the studied specimens are deposited are as follows: CI, Centro Interdisciplinario de Ciencias Marinas (CICIMAR, I.P.N.), La Paz, BCS; UABC, Facultad de Ciencias, Universidad Autonoma de Baja California, Ensenada, Baja California; ENCB-I.P.N., Departamento de Zoologia, Escuela Nacional de Ciencias Biologicas, Mexico, D.F.; CIB, Centro de Investigaciones Biologicas del Noroeste, S.C., La Paz, BCS; IBUNAM, National Fish Collection at the Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico, D.F.; CAS, California Academy of Sciences, San Francisco, California, U.S.A.; and LACM, Natural History Museum of Los Angeles County, Los Angeles, California (cf. Other material examined in Appendix 1). Number of specimens examined is indicated in parenthesis.
Lile nigrofasciata sp. Nov. (Fig. 1).
Holotype. --CI: 2032, a female of 104 mm standar length (SL) from Bahia de Guasimas, Sonora, Mexico, collected by E.F. Balart and A. Arreola Lizarraga, 23 Feb. 1996, at a depth of 1.5 m.
Paratypes.--CI: 2039(1), 98 mm SL collected along the holotype: IBUNAM: P9520 (2), 96.0-100.0 mm SL, idem; CIB: 3001 (2), 93.1-94.5 mm SL, idem. CI: 1886 (50), 56.6-77 mm SL, Santispac, Bahia Concepcion, BCS, collected by P.J.P. Whitehead, 29 Nov. 1988. CI: 4922 (35), 74.1-86.0 mm SL, Estero El Conchalito, Ensenada de La Paz, BCS, Mexico, collected by A.F. Gonzalez-Acosta, 18 Aug. 1997.
The results of the morphometric ane meristic analysis are listed in the Table 1.
Diagnosis.--A species belonging to the genus Lile, with the following differential characteristics: body elongate, very compressed and high, its maximum depth 3.3 to 4.4 (mean = 3.7) times in standard length; dorsal and ventral profiles convex; 36 to 40 (mode = 38) gill rakers on lower limb of the first gill arch; 34 to 49 (mode = 40) midlateral scales; 14-18 (mode = 16) prepelvic scutes; 10-16 (mode = 12) postpelvic scutes; dorsal and anal fins with 13-17 (mode = 15) and 14-16 (mode = 16) rays, respectively; head relatively small, its maximum length 3.7 to 5.4 (mean = 4.2) times in standard length; body with a dark green (in life) or black (in alcohol) band extended from the supraposterior edge of the opercle to the final part of the caudal peduncle; iris black; pupil translucent, whitish or lightly gray; most of individuals examined have both tips of their caudal lobes with a black stain; color of the opercle in specimens <70 mm SL is gray-silver, and in those >80 mm SL with a dark gray or black stain.
Description.--Body elongate, fusiform, compressed, with dorsal profile arched and the ventral profile notably convex. Cephalic region spindle-shaped; mouth oblique, almost vertical. Head relatively small, its length 3.7 to 5.4 (mean = 4.2) times in standard length; eye large, its diameter 2.4 to 4.1 (mean = 3.0) times in cephalic length; preorbital distance shorter than eye diameter and contained 3.1 to 5.2 (mean = 3.9) times in cephalic length; postorbital distance longer than eye diameter and contained 1.9 to 3.2 (mean = 2.6) times in cephalic length. Posterior edge of opercle rounded and flat, not serrated, but with a concave notch on its post-distal part. Preopercle flat, its posterior edge straight, shallowly oblique and directed anteriorly where it connects with the rounded suborbital. A distinctive membranous structure on anterior edge of cleithrum that appears juxtaposed to first lateral scales, and covered with a great number of highly ramified complex tubules. Tubular system extended to nuchal regi on where it connects to other side of body by means of a tubular system. Two anterior fontanels located between supraocular and nuchal regions, distinguishable as two dark areas, also identified as a complex tubular system connected with those of the preopercle, opercle, and membranous structure. Mouth terminal, protrusible, forming a tubular structure composed of two supramaxillary bones, which are associated with the maxillary bone. Both sides of symphysis of lower jaw with several series of conical, blunt, minute teeth. Inner border of jaws without teeth; conical and unicuspidated teeth forming a conglomerate on the palatines. Tongue spatulate and lacking teeth. Six branchiostegal rays supported by flexible ligaments. Lower limb of the first gill arch with 36 to 40 (mode = 38) thin elongate gill rakers, closely approximated. The pseudobranch has from 13 to 14 relatively thick filaments. Body elongate, compressed, deep, maximum depth 3.3 to 4.4 (mean = 3.7) times in standard length; dorsal profile arched, v entral profile notably convex. The ventral scutes originate from the first scale behind the isthmus and finish before the anal pore; their numbers range from 14 to 18 (mode = 16) prepelvic and 10 to 16 (mode = 12) postpelvic or preanal. Origin of dorsal fin situated before that of the pelvic fins and closer to tip of mouth than to base of caudal peduncle. Dorsal fin length is almost equal to the cephalic length; dorsal rays from 13 to 17 (mode = 15), the last two rays highly ramified. Pectoral fins relatively small, falciform, their length almost equal as the cephalic length; insertion of pelvic fins begins at post-distal part of gill opening and above the thoracic border. Pelvic fins small, their maximum length 1.6 to 2.5 (mean = 1.9) times in cephalic length, and 8 branched rays. Anal fin short, not very high, with a scaly sheat, and 14 to 16 rays (mode = 16), the last two rays branched; anal fin situated at posterior one-third of the body. Caudal fin forked, 5-11 upper procurrent rays, 10 upper principal r ays (9 of them forked), 9 lower principal rays (8 of them forked), and 6-9 lower procurrent rays. Scales large, somewhat deciduous, with smooth border, four continuous grooves and one discontinuous of central position, the nearest groove to the lower border has the form of an arch, the other cross the entire scale. Number of scales in lateral series varies from 34 to 49 (mode = 40); predorsal scales 11- 13. Total vertebrate 40-41 (mode = 41), 14 or 15 (mode = 15) caudal. Predorsal bones (=upperneurals) between cranium and the first dorsal finrays 9-11 (mode = 11). Caudal complex with two (60%) or three (40%) epurals.
Biometric Analysis.--Twenty-one biometric variables of 85 examined specimens of Lile nigrofasciata and 71 L. stolifera were analyzed. Wilks' Lambda values varied from 0.0027 to 0.0204, suggesting a strong discrimination between the species (Wilk's Lambda for all variables combined: 0.00273, approx. F (63, 394) = 39.010, p < 0.0000). The values of tolerance ranged between 0.4974 and 0.9002. The classification matrix obtained here indicates that 100% of the examined individuals were correctly classified for the populations of Lile nigrofasciata (Bahia Concepcion and El Conchalito, BCS), as well as that of L. stolifera from El Conchalito, BCS. The specimens of L. stolifera from Bahia Concepcion were correctly classified with an accuracy of 97.22%.
Only three of the 21 biometric variables analyzed contributed significantly to the discrimination of both species of Lile: interorbital distance (p < 0.00001), number of midlateral scales (p < 0.00003), and number of gill rakers on the lower limb of the first gill arch (p < 0.00000).
The squared Mahalanobis distances were significantly greater between populations of L. nigrofasciata and L. stolifera, but were not significant between populations of the same species, as shown in the graph of the discriminant function analysis (Fig. 2).
Coloration.--Living specimens, with the exception of the olive-green or black midlateral band, are yellowish-green and somewhat translucent in their predorsal region. The head is also yellowish-green, with a gray-silver spot on the preopercle and opercle as result of a great concentration of black pigment inside the branchial chamber. Gums and distal part of the lower jaw are black. The upper lip and internasal area are black or dark gray. The gill filaments are whitish or yellowish. The oral chamber is whitish. A distinctive black or dark line travels from the nape to the dorsal fin origin and to the precaudal area. The iris is black or dark gray, the pupil is whitish and translucent. In most individuals both caudal lobes are tipped black, although some of them have only the upper lobe with a faint black tip. Anal, pectoral, and pelvic fins are yellowish or whitish.
Habitat.--The holotype and five paratypes were caught in 1.5 m in Bahia de Guasimas, Sonora, near the town of the same name. This coastal lagoon, with anti-estuarine circulation and muddy-sandy bottom, is located at the geographical coordinates, 27[degrees]54' and 27[degrees]59' N, and 1 10[degrees]48' and 110[degrees]55' W. Water conditions at capture were: salinity, 45 ppt (annual range for 1996, 35-45 ppt), temperature, 20.5[degrees]C (annual range for 1996, 14-39[degrees]C), and Secchi disc visibility, 0.72 m. The dominant fish species at the type locality were Aropsis seemani, Anchovia macrolepidota, Mugil cephalus, Diapterus peruvianus, Anchoa spp., Eucinostomus spp., and Colpichthys regis. The other paratypes from Bahia Concepcion and Ensenada de La Paz (Manglar El Conchalito), BCS, were collected in shallow littoral areas with soft silt, and were usually associated with mangrove roots that border these coastal lagoons. The fish communities at both BCS sites were represented by the following dominant s pecies: Eucinostomus spp., Diapterus peruvianus, Anchovia macrolepidota, Mugil cephalus and Lile stolifera. In addition, voucher specimens captured at oligohaline conditions (5.0-15 ppt) in the coastal lagoon of Tres Palos, Guerrero, Mexico. were also examined. Other specimens of the new species were collected in the Oriental, Occidental, Inferior and Superior coastal lagoons, Oaxaca, as well as in the Mar Muerto, Chiapas, in salinities of [greater than or equal to]35.5% In these last five localities the fish composition is very similar, but Lile gracilis replaces to L. stolifera.
Etymology: The name nigrofasciata alludes to the obvious dark or black band that travels the midlateral part of the body, from the posterior edge of the operculum to the end of the caudal peduncle.
Key to the species of Lile of the eastern topical Pacific
1. Midlateral band dark or black green; iris black; pupil translucent, dark gray or black; 36-40 (mode = 38) gill rakers on lower limb of the first gill arch ........... Lile nigrofaciata (Fig 3A). Midlateral band silvery; iris yellow or golden-silvery; pupil back, not translucent; 24-32 gill rakers on lowr limb of the first gill arch ..... 2
2. origin of dorsal fin located ahead of or on a vertical through the insertion of the pelvic fins; 27-42 midlateral scales in a longitudinal series; ventral profile notably convex; body depth 2.8-3.7 times in standard length; tips of both caudal lobes with conspicuous black blotches ... Lile stolifea (Fig. 3B). Origin of the dorsal fin located behind the insertion of the pelvic fins; 34-36 midlatrnal scales in longitudinal series; ventural profile slightly convex or straight; body depth 4.0-5.5 times in standard length; only the upper tip, if any, of the caudal fin with a faint dark mark ....... Lile gracilis (Fig. 3C).
Geographical distribution.--Based on material examined deposited in the ichthyological collections previously mentioned, Lile nigrofasciata is distributed in the Bahia Magdalena-Bahia Almejas lagoon complex in the southwestern Baja California Sur, Mexico, the Golfo de California (Bahia de Guasimas, Sonora; Bahia Concepcion and Manglar El Conchalito at Ensenada de La Paz, BCS), and along the western coast of Mexico and Central America, to Colombia, Ecuador and northern Peru.
The recognition of this new species is interesting from a systematic and bio-geographic point of view. Its closest relationship is with Lile stolifera, based on the position of the dorsal fin in relation to that of the pelvic fins, both species having the origin of the dorsal fin ahead of the insertion of the pelvics. This characteristic is also observed in the Atlantic species L. piquitinga (distributed from Venezuela to Brazil) but not in L. gracilis (distributed from the central Pacific of Mexico to Honduras) whose individuals have the dorsal fin origin slightly behind the origin of pelvic fins (Castro-Aguirre and Vivero 1990). L. stolifera and L. nigrofasciata are distinguished by gill-raker numbers on the lower limb of the first gill arch, the first species with 24 to 31 (mode = 27), the second with 36 to 40 (mode = 38).
The size and form of the scales also different; L. stolifera has small scales (average maximum length 3 mm) in the form of a shield with their posterior edges smoothly convex; in contrast, the scales of L. nigrofasciata measure on average 5 mm in maximum length, and their posterior edges are angular with the sharp tips. The most conspicuous diagnostic characteristic is coloration; L. stolifera has a brilliant silvery midlateral band, golden-yellow or silvery-golden iris, black (no translucent) pupil. In comparison, L. nigrofasciata has a dark or blackgreen midlateral band, dark or black gray iris, and the translucent gray pupil. Both species share characters such as the numbers of scales in a longitudinal series, ventral scutes, and anal rays (Table 1), and filaments of the pseudobranch (12 to 14). Based on five cleared-and-stained specimens of both species, a preliminary osteological study revealed some interesting facts: whereas the total vertebrae number are almost the same in L. stolifera and L. nigrofasc iata (41-42 vs. 40- 41), the caudal vertebrae number is not. L. stolifera has 16 to 17 (16) caudal vertebrae and L. nigrofasciata has 14 to 15 (15). On the other hand, L. gracilis has 38 to 40 (38) total vertebrae number and 17 to 18 (17) caudal vertebrae number. In comparison, the Atlantic species L. piquitinga, has 38 to 41 total vertebrae and 14 to 15 caudal vertebrae (Gomez-Gaspar 1976). L. stolifera has 8 to 10 (9) predorsal bones, L. nigrofasciata 9 to 11 (11), and L. gracilis 8 to 9 (9), very near to L. stolifera, but the first bone of the series is larger than in the other species (Fig. 4). L. piquitinga has 7 to 8 predorsal bones (Gomez-Gaspar 1976; McGowan and Berry 1984). Two or three epural bones were observed in the caudal skeleton of L. nigrofasciata (Fig. 5), whereas in L. stolifera two epural bones is the most frequent situation (80%) and rarely three. Vivero and Romero-Castillo (1990) found two epural bones in L. gracilis; however, in very small specimens (e.g. 35.6 mm) three epural bones can be observed; in some the double character of the second epural bone is still evident. Some osteological studies have shown that in some species of Clupeiformes the fusion of one pair of epural bones is relatively common during larval development (Balart 1985; Matsuoka 1998).
The geographical ranges of both species also differ (Fig. 6): Lile nigrofasciata has a widespread distribution in the eastern tropical Pacific, while L. stolifera has a quasi-antitropical distribution. Both species are sympatric from Bahia Magda-lena-Bahia Almejas coastal lagoon complex in the northwestern Baja California Sur and both coasts of Golfo de California to Bahia Banderas, Jalisco, Mexico. The distribution of Lile stolifera is interrupted from northern Bahia Banderas, appearing again as sympatric with L. nigrofasciata from Costa Rica to Peru (Castro-Aguirre et al. 1999). L. nigrofasciata is also sympatric with L. gracilis from the Mexican central Pacific to Golfo de Fonseca, Honduras (S. Contreras-Balderas, unpublished data).
Photographs of specimens identified as "Lile stolifera" that were provided by Chirichigno (1963: 14, fig. 5) and Yanez-Arancibia ("1978" : 266, plate 12, fig. 4) correspond to L. nigrofasciata.
It is not feasible to present a probable phylogeny of the genus because more detailed anatomical studies of the four nominal species are needed, particularly the osteology of the caudal complex and the neurocranium, as well as a comparative study of their early ontogenic stages. Based on some osteologic aspects of Lile gracilis, Vivero and Romero-Castillo (1990) proposed that the allopatrical processes led to the persistence of neotenic characters. It is necessary, however, to carry out finer studies to understand the evolutionary processes that gave origin to L. gracilis, the species that replaces L. stolifera in the central part of the tropical eastern Pacific.
Appendix 1. Other Material Examined:
Lile nigrofasciota sp. nov. Baja California Sur, Mexico: Ensenada de La Paz at Manglar [Estero de] El Conchalito, CI: 4954, 30(57.5-85.0 mm), 16-1711X/1997, A.F. Gonzalez Acosta; Bahia Magdalena at Estero San Buto [24[degrees] 46' N, 112[degrees] 05' W], CI: 4560, 1(110.0 mm), 4/VI/1984, coll.?; Bahia Magdalena at Estero Las Botellas [24[degrees] 26' N, 111[degrees] 07' W], CI: 778, 1(77.0 mm), 10/XI/1985, U. McGregor; Bahia Magdalena at Estero San Carlos [24[degrees] 47' N, 112[degrees] 05' WI, CI: 2279, 2(77.0-78.0 mm), 8/XI/1990, R. Rodriguez; Bahia Magdalena at Puerto Chale [24[degrees] 43' N, 111[degrees] 32' W], CL: 1861, 1(98.3 mm), 14/1V11986, F. Garcia; Bahia Magdalena at Estero Medano Amarillo [24[degrees] 43' N, 112[degrees] 02' W], CI: 1894, 4(87.0-96.0 mm), 11/II/1981, J.L. Castro Ortiz. Guerrero, Mdxico: Laguna de Tres Palos [15[degrees] 48' N, 98[degrees] 46' W], CI: 2282, 63(33.0-61.0 mm), 2411V/1991, P.J.P. Whitehead et al. Tumaco, Colombia, CAS: 213293, 20 (49.0-82,5 mm), 1913, A. Henn & C. Wilson.
Lile stolifera. Baja California Sur, Mexico: Ensenada de La Paz at Manglar El Conchalito, CI: 4738, 7(79.0-86.0 mm), 22-23/IV/1997, A.F. Gonzalez Acosta; Estero de San Jose del Cabo [22[degrees] 55' N, 109[degrees] 46' W], CI: 3279, 58 (56.0-71.0 mm), 17/V/1989, P.J.P. Whitehead & R. Rodriguez; Bahia San Juanico at Punta Pequefia, LACM: 32084, 73 (45-64 mm), 28/X/1971, C. Swift; Bahia Concepcion at Isla Anegada, CI: 1011, 36(53.0-74.0 mm SL), 12/VIII/1990, J. Rodriguez et al.; Rio Santa Rosalia [San Luciano], LACM: 50296, 4 (47-62 mm), 12/III/1957, G.W. Barlow; Rio Mulege at 2.1-3.0 km above the mouth, UABC: 0180, 2(47.7-61.7 mm), UABC: 0191, 3(46.2-50.7 mm), UABC: 0204, 1(47.7 mm), 14/X/1995, G. Ruix-Campos. Nayarit, Mexico: Estero San Blas, LACM: 50485, 1(86 mm), 30/I/1958, B.W. Walker et al. Puerto Parker, Costa Rica, CAS: 146862, 10(54.0-73.5 mm), 1222/111938, W. Beebe et al.
Lile gracilis. Guerrero, Mexico: Mouth of the Rio Balsas (La Barra de San Francisquito), ENCB-IPN: 7134, 6(53.0-74.0 mm), 24/II/1988, A. Marmolejo; and Rio Balsas at Brazo de San Francisquito, CIB: 2501, 5(35.6-67.6 mm), 6/X/1993, F de Lachica-Bonilla et al.
[Figure 2 omitted]
Table 1 Morphometry of Lile nigrofasciata sp. nov. and L. stolifera. Measurements according to Whitehead (1985). Lile nigrofasciata sp. nov. N = 85 Mean Mode Meristic measures Scales in lateral series 40 40 Gill rakers on lower limb of the 38 38 first gill arch Prepelvic scutes 16 16 Postpelvic or preanal scutes 12 12 Dorsal fin rays 15 15 Anal fin rays 15 16 Morphometric measures in thousandths of the standard length Cephalic length 239.3 225.4 Body depth 273.7 271.4 Predorsal length 442.5 438.6 Preanal length 754.1 746.5 Caudal peduncle length 122.9 121.2 Caudal peduncle length 114.4 113.6 Pelvic fin length 127.4 136.4 Pectoral fin length 183.3 183.1 Preorbital length 62.2 70.4 Postorbital length 94.0 100.0 Eye diameter 81.0 85.7 Pupil diameter 43.3 45.5 Interobital distance 62.3 57.1 Upper jaw length 105.6 100.0 Lile nigrofasciata sp. nov. N = 85 Min Max s Meristic measures Scales in lateral series 34 49 2.7 Gill rakers on lower limb of the 36 40 1.1 first gill arch Prepelvic scutes 14 18 0.8 Postpelvic or preanal scutes 10 16 1.1 Dorsal fin rays 13 17 0.8 Anal fin rays 14 16 0.7 Morphometric measures in thousandths of the standard length Cephalic length 185.7 271.2 14.4 Body depth 228.6 307.6 16.9 Predorsal length 338.7 491.6 20.7 Preanal length 677.4 885.7 34.6 Caudal peduncle length 101.6 157 11.5 Caudal peduncle length 97.4 128.8 7.1 Pelvic fin length 103.7 145.2 9.2 Pectoral fin length 134.3 213.1 13.4 Preorbital length 46.3 78.7 6.7 Postorbital length 64.0 136.0 10.3 Eye diameter 59.2 99.4 7.3 Pupil diameter 35.1 57.1 5.0 Interobital distance 40.3 85.4 10.7 Upper jaw length 61.4 121.4 9.3 Lile stolifera N = 71 Mean Mode Min Meristic measures Scales in lateral series 36 40 27 Gill rakers on lower limb of the 27 27 24 first gill arch Prepelvic scutes 16 16 13 Postpelvic or preanal scutes 12 13 10 Dorsal fin rays 15 15 14 Anal fin rays 15 15 13 Morphometric measures in thousandths of the standard length Cephalic length 237.7 250.0 208.3 Body depth 298.1 300.0 270.3 Predorsal length 440.2 423.7 400.0 Preanal length 749.4 721.3 694.9 Caudal peduncle length 128.4 125.0 101.7 Caudal peduncle length 123.1 125.0 106.7 Pelvic fin length 130.5 125.0 117.6 Pectoral fin length 179.6 169.5 157.8 Preorbital length 63.8 67.8 51.3 Postorbital length 95.5 100.0 77.4 Eye diameter 84.9 84.8 70.6 Pupil diameter 47.4 50.9 36.5 Interobital distance 73.6 67.8 61.0 Upper jaw length 105.9 101.7 87.7 Lile stolifera N = 71 Max s Meristic measures Scales in lateral series 42 4.0 Gill rakers on lower limb of the 31 1.5 first gill arch Prepelvic scutes 18 0.9 Postpelvic or preanal scutes 14 1.0 Dorsal fin rays 18 0.8 Anal fin rays 19 1.0 Morphometric measures in thousandths of the standard length Cephalic length 269.2 10.6 Body depth 356.6 12.6 Predorsal length 475.0 18.9 Preanal length 902.8 30.7 Caudal peduncle length 151.4 10.4 Caudal peduncle length 137.4 6.7 Pelvic fin length 146.2 7.3 Pectoral fin length 202.7 10.2 Preorbital length 75.5 6.2 Postorbital length 109.1 7.0 Eye diameter 105.3 6.3 Pupil diameter 60.0 6.4 Interobital distance 90.9 5.4 Upper jaw length 121.1 6.1
We wish to thank the following persons and institutions for the facilities and assistance provided during the present study: Jose De La Cruz-Aguero and Adrian Gonzalez-Acosta from Ichthyological Collection of CICIMAR-IPN, for providing help during the examination of specimens utilized in the description of this new species. Liduvina Perezgomez from Laboratory of Morphophysiology (CICIMAR-IPN), for assisting in the process of clearing and staining of the specimens utilized in the osteological analysis. CIBNOR's Arachnology & Entomology Laboratory for providing the camera lucida for drawing the fish bone sketches and to the Ichthyological Collection of CIBNOR for providing part of the type material supporting this study. Oscar Armendariz (CIBNOR) for making the final edition of the fish bone drawings. William N. Eschmeyer and Tomio Iwamoto from the Department of Ichthyology of the California Academy of Sciences in San Francisco for the loan of specimens for this study. Jeffrey A. Seigel from the Natural History Museum of Los Angeles County, in Los Angeles for the access to database and specimens. Salvador Contreras-Balderas from Bioconservacion A.C., in Monterrey (Mexico) for providing the southernmost record of L. gracilis. Daniel A. Guthrie and two anonymous reviewers provided useful comments and editorial help on the manuscript. Finally, Faustino Camarena-Rosales (UABC) for assisting in the statistical analysis of morphometric data.
Accepted for publication 16 March 2001
Balart, E.F. 1985. Development of the vertebral column, fins and fin supports in the Japanese anchovy, Engraulis japonicus (Clupeiformes, Engraulididae). Bull. Mar. Sci., 56: 495-522.
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Yanez-Arancibia, A. "1978" (1980). Taxonomia, ecologia y estructura de las comunidades de peces en lagunas costeras con bocas efimeras del Pacifico de Mexico. Centro Cien. Mar y Limnol., Univ. Nac. Auton. Mexico, Pubi. Esp., 2:1-306.
Jose Luis Castro-Aguirre (1), Gorgonio Ruiz-Campos (2) and Eduardo F. Balart (3)
(1.) Centro Interdisciplinario de Ciencias Marinas, I.P.N. Departamento de Pesquerias y Biologia Marina, Apdo. Postal 592, La Paz, Baja California Sur 23001 Mexico.
(2.) Universidad Autonoma de Baja California. Facultad de Ciencias, Apdo. Postal 1653, Ensenada, Baja California 22800 Mexico.
(3.) Centro de Investigaciones Biologicas del Noroeste, S.C. Laboratorio de Ictiologia, Apdo. Postal 128, La Paz, Baja California Sur 23001 Mexico.
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|Author:||Castro-Aguirre, Jose Luis; Ruiz-Campos, Gorgonio; Balart, Eduardo F.|
|Publication:||Bulletin (Southern California Academy of Sciences)|
|Date:||Apr 1, 2002|
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