<I>Eugerres</I> (Perciformes: Gerreidae)

BULLETIN OF MARINE SCIENCE, 80(1): 109–124, 2007

A REVIEW OF THE MARINE WESTERN ATLANTIC SPECIES OF THE GENUS EUGERRES (PERCIFORMES: GERREIDAE)

Adrián F. González-Acosta, José De la Cruz-Agüero, and José Luis Castro-Aguirre

ABSTRACT Taxonomy of three western Atlantic marine species of the genus Eugerres was assessed on the basis of examination of type series, and specimens representing all nominal species over their geographical distribution. Two specimens from type series (MNHN 0000-9447, 95 mm SL, and MNHN 0000-9452, 97 mm SL) of Eugerres plumieri (Cuvier) were found to be identical to the holotype (USNM 121721, 75 mm SL) of Eugerres awlae Schultz; so E. awlae is redescribed as a valid gerreid species. Likewise, the dry syntypes (MNHN A-5530, 249 mm SL, and MNHN A-5485, 140 mm SL) of E. plumieri were found to correspond to this species and are herein designated as lectotype and paralectotype, respectively. In the same way, the dry syntype (MNHN A-5510, 167 mm SL) of Eugerres brasilianus (Cuvier) is designated herein as lectotype, and other syntypes examined (MNHN A-5510, 94 mm SL, and MNHN 0000-0917, 160 mm SL) designated as paralectotypes. Anal formulae and the number of gill rakers on the lower limb of the first brachial arch were useful to discriminate E. brasilianus (A. III, 7; Gr. 9–11, mode 10) from E. awlae (A. III, 8; Gr. 14–17, mode 16) and E. plumieri (A. III, 8; Gr. 13–17, mode 14). Multivariate analysis sup- ports recognition of three western Atlantic marine species. Canonical variate analysis based on 21 box truss characters classified specimens into three distinct groups (P < 0.0001; 100% correct classification). Distances between the anterior margin of the isthmus and the dorsal extreme of the premaxilla, between the bases of the last dorsal ray and the first anal spine, and between the base of the last dorsal ray and the last anal ray were the most useful characters for group discrimination. Diagnosis and redescriptions are given for all taxa, along with a key for identification.

Three valid species of the gerreid genus Eugerres from the western Atlantic were recognized by Deckert (1973), Deckert and Greenfield (1987), and Gilmore and Greenfield (2002):Eugerres brasilianus (Cuvier, 1830) and Eugerres plumieri (Cuvier, 1830) from marine and estuarine habitats, and Eugerres mexicanus (Steindachner, 1863) restricted to freshwater habitats of southeastern Mexico and northern Guate- mala. However, despite these critical reviews, the identities and limits of distribution of the sympatric species E. brasilianus and E. plumieri have frequently been confused as a consequence of similarities in their external morphology (e.g., Aguirre- León et al., 1982; Báez-Hidalgo and Guevara-Carrió, 1983; Pérez-Hernández and Zavala-Hurtado, 1993). Based on their respective diagnoses, meristics, and biometric analysis using a multivariate approach, an extensive examination of recently collected and museum specimens of species nominally included in the western Atlantic genus Eugerres led us to validate the taxonomy of three marine and one freshwater species. Meristic and morphometric characters observed in some specimens were found to be identical to those of the E. awlae Schultz, 1949 holotype (USNM 121721, 74 mm SL), a species previously considered as synonym of E. plumieri by Deckert (1973), Deckert and Greenfield (1987), and González-Acosta (2005), because the original description was based on juveniles of the species. In this regard, E. awlae is redescribed here and

Bulletin of Marine Science 109 © 2007 Rosenstiel School of Marine and Atmospheric Science of the University of Miami 110 BULLETIN OF MARINE SCIENCE, VOL. 80, NO. 1, 2007 considered a valid gerreid species from the western Atlantic. The Mexican mojarra E. mexicanus is mentioned for comparative purposes only, because of its restricted freshwater habitat and peculiar morphology. The taxonomic status of this gerreid species is currently under review.

Methods

The meristic counts follow Hubbs and Lagler (1964), including the last two branched fin rays counted herein as one element. In total, 141 specimens were examined and listed after each species account. Institutional abbreviations are as listed in Leviton et al. (1985) with the additions of CICIMAR-CI for Centro Interdisciplinario de Ciencias Marinas, I.P.N at La Paz, Baja California Sur (Mexico), and ECOCH for El Colegio de la Frontera Sur, unidad Che- tumal, Quintana Roo (Mexico). The identification of individuals from the nominal species studied here was based on revisions and taxonomic keys by Schultz (1949), Deckert (1973), Randall and Vergara (1977), Deckert and Greenfield (1987), Gilmore and Greenfield (2002), and McEachran and Fechhelm (2005). Morphometric Procedures.—Standard stepwise Canonical Variate Analysis (CVA) was carried out using 21 box truss segments established by González-Acosta et al. (2005), which are defined by ten anatomical landmarks describing the body shape of each species (Fig. 1). Morphometric distances were measured with digital calipers of resolution 0.01 mm attached to a personal computer. All morphometric data were standardized to remove the size com- ponent from shape measurements, and to minimize possible bias due to variability of standard length, according to Elliot et al. (1995); likewise, measurements were transformed to logarithms to homogenize variances (Sokal and Rohlf, 1981; Bookstein, 1982). CVA was used to quantify the morphological shapes of the species E. awlae, E. brasilianus, and E. plumieri, by determination of linear combinations of variables with maximum separation among specimens of different species. The coefficients provided by CVA were used to determine the discrimination value of each original variable, and a scatter diagram based on the main canonical variables yielded a deployed image of the relative similarity of species. Statistical significance of the CVA results was determined by Wilks’ λ statistic (P < 0.0001). Biometric analyses were carried out using Statistica for Windows, version 6.0.

Results

Taxonomy of Western Atlantic Eugerres.—Meristic comparison revealed frequent overlap in the numbers of dorsal (IX, 10) and anal fin (III, 8) elements, and in the number of gill rakers (13–17) on the lower limb of the first branchial arch between E. awlae and E. plumieri (Table 1). On the other hand, E. brasilianus differs from those in having fewer gill rakers (9–11, mode 10) and anal spines and rays (III, 7). Eugerres mexicanus differs from the marine species (E. awlae, E. brasilianus, and E. plumieri) in having fewer gill rakers (12–15, mode 13), enlarged and ventrally flattened lips, and a distinct geographic distribution restricted to freshwater habitats. Canonical variate analysis, applied to test species assignment, produced two significant canonical variates (Wilks’ λ: 0.00001; F (42, 212) = 2070.5; P < 0.0001) account- ing for 100% of variation (CV1 = 74%; CV2 = 26%). The specimens were separated into three clearly distinct groups whose centroid distances were significant (Fig. 2); the Mahalanobis distance (D2) of E. awlae was greater than for any other species analyzed. Groups were classified individually at 100%. Examination of the canonical variates showed that CV1 and CV2 discriminated all groups of species. Distances between the anterior margin of the isthmus and upper point of the premaxilla, between GONZÁLEZ-ACOSTA ET AL.: REVIEW OF WESTERN ATLANTIC EUGERRES 111

Figure 1. Description of the box truss segments resulting from 10 external landmarks (points) used in the canonical variate analysis (21 interpoint distances or measurements): 1–2 (A) = upper tip of premaxillar to base of first dorsal spine; 2–3 (B) = base of first dorsal spine to base of last dorsal spine; 3–4 (C) = base of last dorsal spine to base of last dorsal ray; 4–5 (D) = base of last dorsal ray to dorsal posterior edge of caudal peduncle; 5–6 (E) = dorsal posterior edge to ventral posterior edge of caudal peduncle; 6–7 (F) = ventral posterior edge of caudal peduncle to base of last anal ray; 7–8 (G) = base of last anal ray to base of first anal spine; 8–9 (H) = base of first anal spine to base of pelvic fin spine; 9–10 (I) = base of pelvic fin spine to anterior margin of isthmus; 10–1 (J) = anterior margin of isthmus to upper tip of premaxillar; 1–9 (K) = upper tip of premaxillar to base of pelvic fin spine; 2–10 (L) = base of first dorsal spine to anterior margin of isthmus; 2–9 (M) = base of first dorsal spine to base of pelvic fin spine; 2–8 (N) = base of first dorsal spine to base of first anal spine; 3–9 O( ) = base of last dorsal spine to base of pelvic fin spine; 3–8 (P) = base of last dorsal spine to base of first anal spine; 3–7 Q( ) = base of last dorsal spine to base of last anal ray; 4–8 (R) = base of last dorsal ray to base of first anal spine; 4–7 S( ) = base of last dorsal ray to base of last anal ray; 4–6 (T) = base of last dorsal ray to ventral posterior edge of caudal peduncle; 5–7 (U) = dorsal posterior edge of caudal peduncle to base of last anal ray. Drawing modified from González-Acosta et al. (2005). the bases of the last dorsal ray and first anal spine, and between bases of the last dorsal ray and last anal ray, were most useful in discrimination among the species (Table 2). The CVA allowed us to validate morphological differences of three western Atlantic Eugerres marine species: E. awlae, E. brasilianus, and E. plumieri, based on the criteria of Schultz (1949), Deckert (1973), Randall and Vergara (1977), Deckert and Greenfield (1987), and Gilmore and Greenfield (2002). A key for identification of western Atlantic species of Eugerres, including E. mexicanus, is provided:

Key to Western Atlantic Species of Eugerres

1a. Enlarged lips, ventrally flattened and extending from anterior margin to behind the midpoint of orbit; origin of first dorsal fin behind the anterior margin of pectoral and pelvic fins; body elongated and thick; gill rakers on the lower limb of the first branchial arch, 12–15 (mode 13); pored lateral-line scales, 38–50 (mode 46). Species restricted to freshwater habitats from southeastern Mexico and northern Guatemala...... “Eugerres” mexicanus 112 BULLETIN OF MARINE SCIENCE, VOL. 80, NO. 1, 2007

Table 1. Distribution frequencies of meristic characters of three marine western Atlantic species of Eugerres.

Dorsal-fin spines (Roman) and rays (Arabic) Eugerres VIII, 10 IX, 9 IX, 10 N awlae 1 1 31 33 brasilianus 35 35 plumieri 1 60 61 Anal-fin spines (Roman) and rays (Arabic) Eugerres III, 7 III, 8 III, 9 N awlae 32 1 33 brasilianus 33 2 35 plumieri 59 2 61 Gill rakers on the lower limb of the first branchial arch Eugerres 9 10 11 12 13 14 15 16 17 n awlae 5 10 15 2 32 brasilianus 3 23 12 38 plumieri 11 28 17 3 1 60 Pored scales on lateral line Eugerres 34 35 36 37 38 39 40 41 n awlae 2 9 10 7 2 2 1 33 brasilianus 5 17 14 3 39 plumieri 2 12 18 17 11 1 61

Figure 2. Canonical variate scores for each individual of E. awlae (°), E. brasilianus (®), and E. plumieri (∆). Canonical variate axes, CV1 (74% of the variance of data) and CV2 (26%) produced by 129 specimens and 21 box truss distances. Classification accuracy, 100% on average and in all cases. Significant differences between the group centroids (see text). GONZÁLEZ-ACOSTA ET AL.: REVIEW OF WESTERN ATLANTIC EUGERRES 113

Table 2. Standardized coefficients of CVA of three marine western Atlantic species of Eugerres, according to geometric distances defined by González-Acosta et al. (2005).

Distance measurments CV1 CV2 Upper tip of premaxillar to base of first dorsal spine 1.6450 −1.0219 Base of first dorsal spine to base of last dorsal spine −0.3132 0.0635 Base of last dorsal spine to base of last dorsal ray −1.1482 0.8507 Base of last dorsal ray to dorsal posterior edge of caudal peduncle 0.677 −0.2329 Dorsal posterior edge to ventral posterior edge of caudal peduncle −0.1065 0.5733 Ventral posterior edge of caudal peduncle to base of last anal ray 0.1501 −0.3466 Base of last anal ray to base of first anal spine −1.3115 −0.0459 Base of first anal spine to base of pelvic fin spine −0.8218 0.9004 Base of pelvic fin spine to anterior margin of isthmus 0.3265 0.6832 Anterior margin of isthmus to upper tip of premaxillar 0.7077 0.7811 Upper tip of premaxillar to base of pelvic fin spine 1.1925 −0.1381 Base of first dorsal spine to anterior margin of isthmus −1.3535 −1.0761 Base of first dorsal spine to base of pelvic fin spine −1.2121 2.6617 Base of first dorsal spine to base of first anal spine −0.9243 −2.5803 Base of last dorsal spine to base of pelvic fin spine 0.5264 −3.9167 Base of last dorsal spine to base of first anal spine −3.3678 1.5479 Base of last dorsal spine to base of last anal ray 2.9492 −0.7239 Base of last dorsal ray to base of first anal spine 3.3113 −0.0990 Base of last dorsal ray to base of last anal ray −1.648 1.3548 Base of last dorsal ray to ventral posterior edge of caudal peduncle −1.648 1.3548 Dorsal posterior edge of caudal peduncle to base of last anal ray −0.5406 −0.0393 Eigenvalues 688.0133 244.3952 % accumulated 0.737883 1.0

1b. Small lips, not ventrally flattened and extending to anterior margin of the orbit; origin of first dorsal fin over anterior margin of pectoral and pelvic fins; body deep and compressed; gill rakers on the lower limb of the first branchial arch, 9–17 (mode 10, 14, or 16); pored lateral-line scales, 35–41 (mode 37 or 38). Species distributed in marine and estuarine habitats (occasionally enter continental waters) along the western Atlantic coast...... 2

2a. Anal fin III, 7 (occasionally III, 8); premaxillary groove partially or totally scaly, when scales are absent in the center of the groove they can be observed on the margins; gill rakers on the lower limb of the first branchial arch, 9–11 (mode 10); pored lateral-line scales 37–40 (mode 38)...... Eugerres brasilianus (Fig. 3)

2b. Anal fin III, 8; premaxillary groove not scaly; gill rakers on the lower limb of the first branchial arch, 13–17 (mode 14 or 16); pored lateral-line scales 35–41 (mode 37)...... 3

3a. Depressed second anal spine extending beyond the caudal-fin base; depressed second dorsal spine reaching base of 5th or 6th dorsal ray; predorsal profile ascending abruptly from the snout to the base of the first dorsal spine; posterior margin of first dorsal-fin concave; gill rakers on the lower limb of the first branchial arch, 14–17 (mode 16)...... Eugerres awlae (Fig. 4)

3b. Depressed second anal spine extending to behind midpoint of the caudal peduncle, but not past the caudal-fin base; depressed second dorsal spine reaching base of 1st to 4th dorsal ray; predorsal profile ascending gently from the snout to the base of the first dorsal spine; posterior margin of the first dorsal-fin not concave, almost straight; gill rakers on the lower limb of the first branchial arch, 13–16 (mode 14)...... Eugerres plumieri (Fig. 5) 114 BULLETIN OF MARINE SCIENCE, VOL. 80, NO. 1, 2007

Species Account

Eugerres brasilianus (Cuvier, 1830) (Fig. 3)

Gerres brasilianus Cuvier in: Cuvier and Valenciennes, 1830: 344, 458 (orig. descr.; type locality: Brazil). Gerres brasiliensis [sic] Cuvier. Nichols, 1912: 190 [lapsus calami]. Gerres patao Poey, 1860: 320 (orig. descr.; type locality: La Havana, Cuba). Diapterus brasilianus (Cuvier). Meek and Hildebrand, 1925: 392, 600 (new combination). Diapterus lineatus (Humboldt). Andreata, 1988: 66–67 (synon.) [non] Smaris lineatus Hum- boldt, 1821. Eugerres brasilianus (Cuvier). Jordan and Evermann, 1927: 506 (new combination).

Diagnosis.—Anal fin III, 7 (occasionally III, 8); second dorsal and anal spines mod- erately elevated; premaxillary groove totally or partially scaly, covered from the anterior margin by deciduous scales sometimes absent in the center of the groove but present on margins; gill rakers on the lower limb of the first branchial arch 9–11 (mode 10). Description.—Dorsal-fin elements IX, 10; anal-fin elements III, 7 (occasionally III, 8); pectoral-fin elements iii, 13; pelvic-fin elements I, 5; pored lateral-line (LL) scales 37–40 (mode 38); scales between dorsal-fin base and LL 3–4 (mode 3.5); scales between LL and anal-fin base 8–9 (mode 8). Predorsal profile smoothly convex, ascending gently from the snout to the first dorsal spine. Interorbital space with a central triangular depression. Maxillary fails to reach midpoint of pupil. Dorsal-fin origin on the anterior margins of the pectoral and pelvic fins. Second dorsal spine thicker than third, but almost equal in length; when depressed reaches the midpoint between 8th and 9th dorsal spines. Second anal spine thicker than third, and almost equal in length or occasionally longer; when depressed reaches the midpoint of the caudal- peduncle length. Pectoral fins lanceolate, extending to vent margin but not reaching the anal-fin origin. Caudal fin forked with equal length lobes. Lips small, molar-like pharyngeal teeth. Posterior and ventral preopercle margins serrated. Lower margins of preorbital (lachrymal) and jugal frequently have fine serrations. Subopercular and infraorbital margins often entire. Head covered by large cycloid scales, with ctenoid scales on the trunk. Meristic and morphometric data based on 35 specimens (Table 1). Measurements are presented in Table 3. Color in Alcohol.—Dorsum yellow, sometimes yellowish-brown; silvery reflections on body sides, and yellowish-silver belly; dark stripes along scales rows 9–11 (mode 10). Biological Notes.—Maximum size 400 mm TL (González-Acosta, 2005); inhabits sandy or rocky bottoms and penetrates continental waters 0.8 km upriver as was reported in Belize (Greenfield and Thomerson, 1997). Feeds on polychaetes (> 69% of its diet), crustaceans: amphipods, copepods, and ostracods (33%), and insects: larvae of Diptera and Odonate (< 3.3%) (Furtado, 1969; Teixeira and Helmer, 1997); other food includes foraminifera. This species is commercially important for local fisheries in coastal lagoons of the western Atlantic. In Brazil it is caught for sport (Ramires and Barrella, 2003), and due to its great abundance (> 1 t) is considered the most im- GONZÁLEZ-ACOSTA ET AL.: REVIEW OF WESTERN ATLANTIC EUGERRES 115

Figure 3. Eugerres brasilianus, Lectotype MNHN 5510, 167 mm SL, type locality: Brazil. portant resource in the “mojarra” fishery (Baisre, 1985). According to Báez-Hidalgo and Álvarez-Lajonchere (1980), E. brasilianus reaches 86% of total catch in Cuba. Other data on the ontogeny of this species is provided by Eiras-Stofella and Fanta (1991). Remarks.—Eugerres brasilianus differs from its congeners in distinctive anal formulae shared with G. cinereus (Walbaum, 1792), and in the presence of scales in the premaxillary groove that differs from the constrained groove described by Randall and Vergara (1977), Matheson and McEachran (1984), De la Cruz-Agüero (2001), and Gilmore and Greenfield (2002) forEucinostomus gula (Cuvier and Valenciennes in Quoy and Gaimard, 1824). Regan (1903) first reported scales in the premaxillary groove of Gerres patao Poey, 1860 (a synonym of E. brasilianus), as was confirmed in studies by Evermann and Meek (1886), Meek and Hildebrand (1925), and Guitart (1977), but this character was not established as diagnostic due to its ephemeral oc- currence. All the specimens of E. brasilianus examined herein were consistent in the presence of small scales covering the anterior margin of the premaxillary groove, as was asserted by Regan (1903). The analysis of syntypes (MNHN 0000-0917, 160 mm SL, and MNHN A-5510, 94–167 mm SL) of E. brasilianus revealed those characteris- tics. Likewise, observations carried out on specimens from Cuba and Brazil indicate that when the scales are absent in the center of the groove, they can be observed on the margins. The dry syntype (MNHN A-5510, 167 mm SL) ofE. brasilianus is designated herein as lectotype, other syntypes examined (MNHN A-5510, 94 mm SL, and MNHN 0000-0917, 160 mm SL) are designated as paralectotypes. Differences in the second dorsal spine and cephalic average lengths were used by Baéz-Hidalgo and Guevara-Carrió (1983) as criteria to discriminate between E. brasilianus and E. plumieri (sensu lato). However, these authors confused the identity of E. brasilianus, because the anal-fin counts they reported (III, 8) correspond to E. plumieri, a species morphologically similar to E. brasilianus. Based on a biometric study of the E. plumieri–E. brasilianus complex from sev- eral localities in the Gulf of Mexico, Pérez-Hernández and Zavala-Hurtado (1993: 128) established erroneously the presence of E. basilianus in the central and southern Gulf coasts. The specimens they reported as E. brasilianus might be E. plumieri 116 BULLETIN OF MARINE SCIENCE, VOL. 80, NO. 1, 2007 - E. brasil E. , EB: , EP (n = 61) EP 1.9–2.6 [2.2] 2.6–3.1 [2.9] 2.7–3.9 [3.4] 3.1–5.8 [4.0] 1.2–1.9 [1.5] 0.7–2.3 [1.5] 0.7–1.0 [0.9] 2.4–3.2 [2.8] 2.6–3.8 [3.3] 1.3–3.9 [3.5] 2.5–3.0 [2.7] 0.7–1.1 [0.9] 0.7–1.0 [0.9] 1.0–1.3 [1.2] 0.6–1.4 [1.1] 0.8–1.6 [1.2] E. awlae E. EB (n = 35) 2.0–2.5 [2.2] 2.8–3.3 [3.1] 3.2–6.3 [4.3] 3.5–6.1 [4.3] 1.4–2.9 [1.9] 0.9–1.6 [1.2] 0.6–0.9 [0.8] 2.5–3.9 [3.2] 2.8–3.8 [3.3] 3.0–3.6 [3.3] 2.5–3.1 [2.7] 0.5–1.0 [0.8] 0.5–0.9 [0.7] 0.9–1.2 [1.1] 0.7–1.2 [1.1] 0.9–1.6 [1.2] EA (n = 33) EA 1.8–2.5 [2.1] 2.5–3.0 [2.8] 2.2–3.7 [2.8] 2.7–4.1 [3.4] 1.1–1.6 [1.3] 1.5–2.6 [1.9] 0.7–1.9 [0.9] 2.4–3.3 [2.9] 2.9–3.9 [3.4] 3.3–4.0 [3.6] 2.5–3.2 [2.7] 0.8–1.4 [1.1] 0.8–1.4 [1.0] 1.0–1.3 [1.1] 1.0–1.4 [1.1] 1.2–1.7 [1.4] . Interval in millimeters, mean in brackets. EA: brackets. in mean millimeters, in Interval . Eugerres . E. plumieri , and EP: ianus Table 3. Morphometric relationships among marine western Atlantic species of species Atlantic western marine among relationships Morphometric 3. Table Body deep in standard length Head length in standard Length of second dorsal spine in standard length Length of second anal spine in standard length Length of second dorsal spine in greatest body depth Length of second anal spine in caudal peduncle length Distance between the tips of second dorsal and anal spines in standard length Orbit diameter in head length Snouth length in head Length of interorbital space in head length Maxillary length in head Head height in length second dorsal spine Head length in of second dorsal spine Length of second dorsal-spine in third dorsal spine length Length of second anal-spine in third anal spine length Length of second anal-spine in length anal fin base GONZÁLEZ-ACOSTA ET AL.: REVIEW OF WESTERN ATLANTIC EUGERRES 117

(sensu stricto). Unfortunately, the material they examined was never catalogued and therefore is not available for further analysis. Distribution.—Northern records of this species were reported by Evermann and Meek (1886) in Charleston, South Carolina (USA) and Darnell (1962) in Tampico, Tamaulipas (Mexico); but they have not been confirmed in further studies (e.g., Deckert, 1973; Deckert and Greenfield, 1987; Hoese and Moore, 1998; Poulakis et al., 2005) nor on the basis of specimens catalogued in biological museums (this study). Castro-Aguirre (1978) and Castro-Aguirre et al. (1999) mentioned its presence (not confirmed) in Mexican coastal lagoons of Veracruz (Alvarado and Tuxpan-Tam- pamachoco) and Campeche (Laguna de Términos). According to the specimen la- bels examined herein, E. brasilianus is distributed from the West Indies, including coastal waters of Belize and Costa Rica, to south of Brazil, with a disjunct distribution in Panama and Colombia. Material Examined.—CAS 122144 (4) [71.04–140.14], Rio Grande do Norte, Brazil; CAS 152251 (1) [147.9], Rio Bacanga, São Luis, Brazil; CAS 152253 (2) [45.4–46.12], CAS 161843 (2) [174.3–178.09], Pernambuco, Brazil; CAS 152260 (1) [79.09], CAS 15261 (1) [59.35], Fortaleza, Mucuripe, Brazil; CAS 161831 (6) [143.25–162.35], CAS 161834 (1) [212.52], Bahia Salvador, Brazil; CAS 161835 (3) [145.6–157.5], CAS 161841 (2) [169.21–222.23], CAS 161842 (1) [258.38], Espirito Santo, Brazil; CAS 161836 (2) [187.92–215.6], Lagoa de Conceição, Florianopolis, Brazil; CAS 161837 (2) [143.02– 144.62], CAS 17620 (1) [125.48], Sao Paulo, Brazil; CAS 161840 (1) [122.89], Bahia, Brazil; CAS 19702 (1) [208.29], CAS 102704 (2) [99.45–175.8], La Havana, Cuba; MNHN 1988-0087 (1) [145.0], Bahia, Brazil; Syntype MNHN 0000-0917 (1) [163.0], Rio de Janeiro, Brazil; MNHN A-0487 (1) [159.0], Syntype MNHN 0000-5510 (2) [94.0–165.0], Brazil; UCR 1808.005 (2) [113.6–159.7], Limón, Parque Nacional Tor- tuguero, Costa Rica.

Eugerres awlae Schultz, 1949 (Fig. 4)

Eugerres awlae Schultz, 1949: 146–148 (orig. descr.; type locality: lago Maracaibo, Venezuela). Gerres plumieri Cuvier. Evermann and Marsh, 1900: 210 (list; descr.; Puerto Rico) [non] Gerres plumieri Cuvier, 1830. Diapterus plumieri (Cuvier). Miller et al., 2005: 346 (descr.; distr.) [non] Gerres plumieri Cuvier, 1830. Eugerres plumieri (Cuvier). Deckert, 1973: 48. Deckert and Greenfield, 1987: 192. González- Acosta, 2005: 69–74 (synon.) [non] Gerres plumieri Cuvier, 1830.

Diagnosis.—Depressed second anal spine extending to beyond the caudal-fin base; depressed second dorsal spine reaching the base of the 5th or 6th dorsal ray; body deep; convex predorsal profile ascending abruptly from the snout to the first dorsal spine; posterior profile of the first dorsal-fin concave; pectoral fin extending behind the anal-fin origin. Description.—Dorsal-fin elements IX, 10 (occasionally VIII, 10 or IX, 9); anal-fin elements III, 8 (occasionally III, 9); pectoral-fin elements iii, 13 i; pelvic-fin elements I, 5; pored lateral-line (LL) scales 35–41 (mode 37); scales between dorsal-fin base and LL 3–3.5 (mode 3.5); scales between LL and anal fin base 8–9 (mode 8); gill rakers on the lower limb of the first branchial arch 14–17 (mode 16). Interorbital space 118 BULLETIN OF MARINE SCIENCE, VOL. 80, NO. 1, 2007

Figure 4. Eugerres awlae, Holotype USMN 121721, 74 mm SL, type locality: Lago Maracaibo, Venezuela. has a central oval depression; no scales in the premaxillary groove. Maxillary ex- tends to the anterior margin of pupil. Dorsal-fin origin on the anterior margin of the pectoral and pelvic fins. Second dorsal spine thicker and longer than third. Second anal spine thicker and longer than third (occasionally a little higher). Lanceolate pectoral fins extend beyond the vent or the anal-fin origin. Caudal fin forked with equal length lobes. Protractile and terminal mouth, small lips, and molar-like pharyngeal teeth. Serrated posterior and lower peopercle margins. Preorbital (lachrymal) and jugal have serrated margins. Subopercular and infraorbital margins frequently whole (occasionally serrated). Head covered by large cycloid scales, and ctenoid scales on the trunk. Meristic and morphometric data based on 33 specimens (Table 1). Mea- surements are presented in Table 3. Color in Alcohol.—Silvery on back, with brownish-yellow reflections; silvery-yellow belly; dark stripes along the scale rows 9–13 (mode 11). Yellowish-brown fins. Biological Notes.—According to collection data and other records (e.g., Schultz, 1949; Rodríguez, 1973; Miller et al., 2005), E. awlae inhabits shallow coastal waters over grassy areas and muddy bottoms, and brackish mangrove lagoons, and frequently enters the fresh water of river mouths. It is an euryhaline marine species that penetrates estuarine ecosystems to feed and nurse. Because of its buccal apparatus with pharyngeal molariform teeth, it probably feeds on benthic organisms (e.g., polychaetes, crustaceans, and mollusks) like the sympatric species E. plumieri. However, further studies are needed to distinguish between the biological information on the two species. Maximum size 256 mm SL. Commercial interest in E. awlae could be comparable to its congener E. plumieri. Remarks.—Eugerres awlae had been previously established by Deckert (1973), Deck- ert and Greenfield (1987), and González-Acosta (2005) as a synonym of E. plumieri because E. awlae was described using juvenile specimens (56.0–73.8 mm SL) characterized by an irregular anal formula (II, 8–9). We examined the X-ray photograph of the holotype (USMN 121721, 74 mm SL) of E. awlae and saw that the modification of anal-fin elements is due to the presence of a third anal spine branched on their upper margin. The meristic counts carried out for this specimen were: D. IX, 10; A. III, 8; Gr. 15 i; pored scales on LL 36. GONZÁLEZ-ACOSTA ET AL.: REVIEW OF WESTERN ATLANTIC EUGERRES 119

Two syntypes (MNHN 0000-9447, 95 mm SL, and MNHN 0000-9452, 97 mm SL) of E. plumieri [ascriptum], characterized by a depressed second anal spine extending to beyond the caudal-fin base, and a depressed second dorsal spine reaching the base of the 5th or 6th dorsal rays, were found to be identical to the description and holotype (USNM 121721, 74 mm SL) of E. awlae (Schultz, 1949). Therefore, E. awlae is considered herein as a valid gerreid species. Specimens with such traits, included in the “E. plumieri” complex, have been ommited frequently due to overlap in meristic characters used in the identification keys for the Gerreidae family of the western Atlantic (e.g., Randall and Vergara, 1977; Deckert and Greenfield, 1987; Gilmore and Greenfield, 2002). On this basis, the syntypes (MNHN 0000-9447) from the locality Antilles and from Martinique (MNHN 0000-9452) are herein designated as lectotypes of E. awlae. Distribution.—Based on verified records, the distribution ofE. awlae is established along the western coast of the Gulf of Mexico, the West Indies, and Central and South America (Venezuela). Further studies could confirm its presence in South Car- olina, southern Florida (USA), and probably southern Brazil, as has been reported for the E. plumieri complex. Material Examined.—ENCB 6456-A (1) [108.38], CICIMAR-CI s/n-A (1) [142.5], CICIMAR-CI s/n-B (4) [135.15–141.96], IBUNAM 256 (6) [81.24–125.06], IBUNAM 2476 (9) [79.07–115.21], IBUNAM 486 (1) [114.05], Veracruz, Mexico; IBUNAM 555 (4) [81.3–114.63], Campeche, Mexico; IBUNAM 975 (1) [123.19], Tabasco, Mexico; ECOCH 1422 (4) [70.12–84.55], ECOCH 1731 (5) [67.66–74.15], ECOCH 1940 (1) [73.4], ECOCH 1979 (4) [46.7–59.39], ECOCH 1999 (5) [73.41–90.17], ECOCH 3030 (2) [63.69–66.97], ECOCH 4608 (1) [74.73], Chetumal, Mexico; lectotypes MNHN 0000-9452 (1) [97.00], MNHN A-5485 (1) [137.0], Martinique, Antilles (West Indies) and MNHN 0000-9447 (1) [95.0], Antilles (West Indies); UCR 1418.003 (3)[68.91– 77.82], Rio Platano, Honduras; UCR 1808.005 (2) [69.79–129.4], Limon, Costa Rica. Eugerres awlae USNM 121721 [74 mm SL], Lago Maracaibo, Venezuela. Meristic counts and X-ray photograph from E. awlae USMN 121721 was provided by Dr. Jef- frey Williams from USMN.

Eugerres plumieri (Cuvier, 1830) (Fig. 5)

Gerres plumieri Cuvier in: Cuvier and Valenciennes, 1830: 340, 452, pl. 467 (orig. descr.; type locality: Antilles [sic]). Gerres brasilianus Cuvier. Evermann and Marsh, 1889: 209 [non] Gerres plumieri Cuvier, 1830. Gerres embryx Jordan and Starks, 1898: 1379 (orig. descr.; type locality: Charleston, South Carolina, USA) (synon.). Diapterus plumieri (Cuvier). Meek and Hildebrand, 1925: 592, 598 (new combination). Diapterus lineatus (Humboldt). Andreata, 1988: 67 (synon.) [non] Smaris lineatus Humboldt, 1821. Eugerres plumieri (Cuvier). Jordan and Evermann, 1927: 506 (new combination). Reséndez- Medina, 1970: 119 (notes; fig.; Laguna de Tamiahua, Veracruz, Mexico). Reséndez-Medi- na, 1973: 233 (notes; fig.; Laguna de Alvarado, Veracruz, Mexico). Schmitter-Soto, 1998: 143–145 (catalog.; notes; distr.; Bahia de Chetumal, Quintana Roo, Mexico). 120 BULLETIN OF MARINE SCIENCE, VOL. 80, NO. 1, 2007

Eugerres awlae Schultz. Deckert, 1973: 44, 48. Deckert and Greenfield, 1987: 192. González- Acosta, 2005: 69–74 (synon.). [non] Eugerres awlae Schultz, 1949. [?] Eugerres brasilianus (Cuvier). Castro-Aguirre, 1978: (cat.; distr.). Pérez-Hernández and Za- vala-Hurtado, 1993: 124,126 (biometry; Gulf of Mexico). Castro-Aguirre et al., 1999: 328 (cat.; distr.).

Diagnosis.—Depressed second anal spine passing the midpoint of the caudal peduncle, but not the caudal-fin base; depressed second dorsal spine reaching the base of the 1st to 4th dorsal ray; anterior portion of head blunt; predorsal profile ascending gently from the snout to first dorsal spine; posterior profile of the dorsal fin not concave; pectoral and pelvic fins extending beyond the vent but not past the anal-fin origin. Description.—Dorsal-fin elements IX, 10 (occasionally IX, 9); anal-fin elements III-8 (occasionally III, 9); pectoral-fin elements 16; pelvic-fin elements I, 5; pored lateral-line scales 37–40 (mode 37 and 38); scales between dorsal-fin base and lateral line 3–3.5 (mode 3.5); scales between lateral line and anal fin base 8–9 (mode 8); gill rakers in the lower limb of the first branchial arch 13 to 17 (mode 14). Interorbital space with a bell-shaped depression; no scales on the premaxillary groove. Maxillary surpasses the midpoint of the pupil. Dorsal-fin origin on the anterior margin of the pectoral and pelvic fins. Second dorsal spine almost equal in length to third. Second anal spine thicker and a little longer than the third. Lanceolate pectoral fins extend beyond the vent and the pelvic fins, but do not pass the anal-fin origin. Deeply forked caudal-fin, lobes equal in length. Protractile and terminal mouth, small lips, and molar-like pharyngeal teeth. Preopercular, lachrymal, and jugal bones finely serrated. Head has large cycloid scales, and the trunk is covered by ctenoid scales. Meristic and morphometric data based on 61 specimens (Table 1). Measurements are presented in Table 3. Color in Alcohol.—Yellowish-silver on back, with blue reflections; yellowish belly; dark stripes along the scales rows 9–14 (mode 12). Pale or yellowish dorsal and anal fins. Biological Notes.—Eugerres plumieri [s.s.] is an euryhaline species that inhabits coastal lagoons and estuaries with sandy and muddy bottoms bordered by mangroves, between salinities of 4 and 25 and 2.7 and 6.6 ml L−1 oxygen concentration. It is an iteroparous fish that spawns 70,000 ova per female (Schmitter-Soto, 1998). The morphology of the buccal apparatus appears adapted for consumption of benthic organisms with hard structures (e.g., polychaetes, amphipods, brachyura, cladocera, copepods, cumaceans, isopods, mysidaceans, ostracods, bivalves, and gasteropods, insects (larvae of quironomids), and detritus as has been mentioned for E. aff. plumieri (Springer and Woodburn, 1960; Austin, 1971; Austin and Austin, 1971; Odum and Heald, 1972; Amezcua-Linares and Yáñez-Arancibia, 1980; Vargas-Maldonado et al., 1981; Aguirre-León et al., 1982; Odum et al., 1982; Arenas-Granados and Ace- ro, 1992; Aguirre-León and Díaz-Ruiz, 2000). The species may be abundant in estuaries and coastal lagoons of the western Atlantic, and thus may have commercial importance for local fisheries, but further studies are needed to examine this. A very abundant but largely unexploited resource exists in some localities (Reséndez-Me- dina, 1970, 1973; Aguirre-León et al., 1982). In Colombia, however, E. plumieri [s.l.] is reported as a highly threatened species subject to excessive exploitation (Mejía and Acero, 2002). GONZÁLEZ-ACOSTA ET AL.: REVIEW OF WESTERN ATLANTIC EUGERRES 121

Figure 5. Eugerres plumieri Lectotype, MNHN A-5530, 249 mm SL, type locality: Puerto Rico, Antillas.

Remarks.—The taxonomy of E. plumieri [s.l.] was clarified in the critical reviews of the genus Eugerres by Deckert (1973), and Deckert and Greenfield (1987), in part to deal with the confusion with E. brasilianus and uncertainty in distribution limits that persists in the literature (e.g., Báez-Hidalgo and Guevara-Carrió, 1983) due to high similarity between them (sensu Aguirre-León et al., 1982; Pérez-Hernández and Zavala-Hurtado, 1993). The analysis, based on specimens of E. brasilianus and E. plumieri [s.s.] from museums and their distribution range, allowed us to corroborate differences between these species in anal formulae, numbers of gill rakers, and external morphology. Likewise, E. plumieri differs from its congenerE. awlae in the depressed second anal spine that passes the midpoint of the caudal peduncle, but fails to reach the caudal- fin base, and the depressed second dorsal spine reaching the base of the 1st to 4th dorsal ray. Due to frequent overlap in meristic traits and the synonym established between the two species, specimens of E. awlae were included in the E. plumieri complex. However, the morphological distinction of E. plumieri from E. awlae is herein cor- roborated by CVA applied to the data. In the same way, observations carried out on the dry syntype (MNHN A-5530, 249 mm SL) and other E. plumieri records from Mexico [sic] (MNHN 1883-0783, 280 mm SL, and MNHN 1883-0784, 194 mm SL) and Jamaica (MNHN 1883-0785, 164 mm SL) confirm the taxonomic identity of this species. Based on this criterion, the dry syntype (MNHN A-5530, 249 mm SL) of E. plumieri from Porto-Rico [sic] locality is designated herein as lectotype of this species. Other specimens examined from Guatemala (MNHN 1883-0783, 280 mm SL, and MNHN 1883-0784, 194 mm SL) and Jamaica (MNHN 1883-0785, 164 mmSL) are designated as paralectotypes. Distribution.—Eugerres plumieri [s.s.] exhibits a sympatric distribution with E. awlae along the coast of the Gulf of Mexico (Veracruz, Campeche, and Tabasco), and in the Mexican Caribbean (Bahia de Chetumal and continental waters of Quintana Roo). Records of specimens from the MNHN confirm its presence in localities of Mexico, Guatemala, and the West Indies (Jamaica and Puerto Rico). Further collection efforts could confirm its presence in South Carolina, southern Florida (USA), and Brazil, as was asserted in recent records (e.g., Deckert and Greenfield, 1987; Gilmore and Greenfield, 2002; McEachran and Fechhelm, 2005; Miller et al., 2005). 122 BULLETIN OF MARINE SCIENCE, VOL. 80, NO. 1, 2007

Material Examined.—ECOCH 1291 (2) [103.17–117.08], ECOCH 1294 (3) [61.51– 74.12], Canal Raudales [Laguna Guerrero], Chetumal, Quintana Roo, Mexico; ECOCH 1302 (2) [63.04–83.01], Laguna Xcalak, Quintana Roo, Mexico; ECOCH 1303 (3) [79.8–114.27], Rio Huach, Quintana Roo, Mexico; ECOCH 1709 (2) [69.41–94.95], Laguna Ubero, Quintana Roo, Mexico; ECOCH 3032 (2) [64.87–64.24], La Aguada, Quintana Roo, Mexico; ECOCH 3761 (10) [96.11–121.33], Isla Tamalkab, Chetumal, Quintana Roo, Mexico; ECOCH 3784 (2) [84.42–87.26], Punta Catalan, Chetumal, Quintana Roo, Mexico; ECOCH 3798 (2) [122.3–123.31], Punta Calentura, Che- tumal, Quintana Roo, Mexico; ECOCH 4553 (1) [80.22], Cayo Venado, Chetumal, Quintana Roo, Mexico; IBUNAM 2393 (5) [93.26–122.27], Isla Tamalkab [sic], Che- tumal, Quintana Roo, Mexico; IBUNAM 2922 (2) [124.24–127.34], Bahia Chetumal, Quintana Roo, Mexico; CICIMAR-CI s/n-C (3) [131.21–145.23], Veracruz, Mexico; ENCB 4900 (1) [162], Veracruz, Mexico; Paralectotype MNHN 0763 (1) [104.0], Isla Tamalcab, Chetumal, Quintana Roo, Mexico; Syntype MNHN A-5530 (1) [250.0], Puerto Rico; MNHN 1883-0785 (1) [164.0], Jamaica; MNHN 1883-0783 (1) [280.0], MNHN 0000-0784 (1) [194.0], MNHN A-9416 (1) [182.0], Mexico.

Acknowledgments

We thank W. Eschmeyer, D. Catania, and J. Fong (CAS); H. Espinosa Pérez and L. Huido- bro (IBUNAM); J. J. Schmiter Soto (ECOCH); E. Teniente (ENCB); P. Béarez, P. Pruvost, G. Duhamel, J. Gregorio, C. Ferrara, C. Guchereu, and L. Nandein (MNHN); S. L. Jewett, J. Wil- liams, and S. Rarendon (USNM); and W. Bussing (UCR) for loan and donation of specimens. A. F. G. -A. also thanks C. Myroth (ILD-NIU) for library assistance. The authors are grateful for grants from CGPI-IPN, EDI-IPN, COFAA-IPN, and SNI-CONACyT. T. Morey made cor- rections to the English manuscript. This study is part of the senior author’s doctoral disserta- tion. Our appreciation is also extended to three anonymous reviewers.

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Date Submitted: 8 September, 2005. Date Accepted: 19 September, 2006.

Addresses: (A.F.G.-A., J.C.-A., J.L.C.-A.) Centro Interdisciplinario de Ciencias Marinas, I.P.N., Colección Ictiológica, CICIMAR-IPN, Apartado Postal 592, La Paz, Baja California Sur, México, 23000. Corresponding Author: (A.F.G.-A.) Email: .

&lt;I&gt;Eugerres&lt;/I&gt; (Perciformes: Gerreidae)

<I>Eugerres</I> (Perciformes: Gerreidae)