The sea bream Diplodus levantinus n. sp. is described from off the coasts of Israel in the eastern Mediterranean Sea, where it replaces Diplodus sargus (Linnaeus, 1758). The new species is characterized by 11-12 spines and 10-16 soft rays in the dorsal fin, 3 spines and 11-13 soft rays in the anal fin, 15-17 pectoral fin rays, 6-9 + 8-12 gill rakers on the first gill arch, upper and lower jaws with a single row of 4 incisors on each side, followed by a total of 16-19 molariform teeth in the upper jaw and 12-14 molariform teeth in the lower jaw, with the molariforms of the upper jaw separated from the incisors by a wide, toothless gap, and the sides of the body in adults with 8 vertical bars of equal width which are present even in large adults, followed by a broad bar on the caudal peduncle which usually nearly reaches the ventral margin of the caudal peduncle. An updated checklist of the species of the genus Diplodus, and a key to species of the Diplodus sargus species group from the eastern Atlantic and Mediterranean Sea, are presented.
RESUMEN
El sargo levantino Diplodus levantinus n. sp. se describe a partir de ejemplares de las costas de Israel, en el Mediterráneo oriental, donde reemplaza a Diplodus sargus (Linnaeus, 1758). La nueva especie se caracteriza por: 11-12 espinas y 10-16 radios blandos en la aleta dorsal, 3 espinas y 11-13 radios blandos en la aleta anal, 15-17 radios en las pectorales, 6-9 + 8-12 branquispinas en el primer arco branquial; las mandíbulas superior e inferior con una sola fila de 4 incisivos en cada lado, seguidos por un total de 16-19 molares en la mandíbula superior y 12-14 molares en la mandíbula inferior, con los molares de la mandíbula superior separados de los incisivos por una gran distancia sin dientes; los lados del cuerpo, en los adultos, con 8 bandas verticales de igual anchura, presentes incluso en adultos de gran tamaño, seguidas de una amplia banda en el pedúnculo caudal que, por lo general, casi alcanza el margen ventral del pedúnculo caudal. Se diferencia principalmente de D. sargus por el menor número de dientes molares en las mandíbulas superior e inferior, y por la amplia separación, sin dientes, entre molares e incisivos de la mandíbula superior. Se presenta una lista actualizada de las especies del género Diplodus y una clave para las especies del grupo de especies Diplodus sargus del Atlántico este y el Mediterráneo.
KEYWORDStaxonomyfishesnew speciesMediterranean Seaneotype designationchecklistkeyPALABRAS CLAVEtaxonomíapecesnueva especiemar Mediterráneodesignación de neotipolista de especiesclaveINTRODUCTION
The sparid fish genus Diplodus are distributed in tropical and warm temperate waters of the Atlantic Ocean and the western Indian Ocean including the Red Sea. They are mainly found on shallow water reefs. The genus includes a total of 22 valid subspecies and species (Paz et al. 1974, Paz 1982, Eschmeyer and Fricke 2015).
Diplodus was first described by Rafinesque (1810a: 26, 54) for Sparus annularis Linnaeus, 1758. The species of the genus have been known since ancient times; two species now assigned to the genus were described by Linnaeus (1758), namely Diplodus annularis and D. sargus. The latter species has been known since Aristotle, and its description by Linnaeus (1758) was based on multiple sources from localities in the Mediterranean Sea (Greece, France, Italy). Valenciennes in Cuvier and Valenciennes (1830: 14) unnecessarily renamed the species as Sargus rondeletii, a name that was used by many subsequent authors. Diplodus sargus may reach a total length of up to 40 cm (Man-Wai and Quignard 1982: 175), which equals a standard length of approximately 324 mm; Fischer et al. (1987) even give a maximum total length of 45 cm. Cadenat (1964) revised the genus Diplodus from West Africa, and revived the name Diplodus sargus; he described two subspecies, including the description of Diplodus sargus typicus, which included specimens from West Africa as well as the Mediterranean Sea. Partially based on his West African material, Paz et al. (1974) described a new subspecies Diplodus sargus cadenati from West Africa and the Canary Islands.
In his thesis project, Paz (1975) revised the genus Diplodus. He defined the Diplodus sargus species group as having 4-10 vertical bands along the sides of the body, plus sometimes additional smaller bands in the spaces between the main bands, 4 incisors each in the upper and lower jaws, and large molars in 2-4 rows. In the species group, he included the subspecies Diplodus sargus sargus (Linnaeus, 1758) from the Mediterranean, D. s. cadenatiPaz, Bauchot and Daget, 1974 from the eastern Atlantic, D. s. capensis (Smith, 1844) from South Africa, and D. s. lineatus (Valenciennes in Cuvier and Valenciennes, 1830) from the Cape Verde Islands. These results were previously summarized in the paper of Paz et al. (1974). In addition, he reported “Diplodus X” from Israel (Paz 1975: 57-61, Fig. 27), which he thought to be a questionable hybrid between D. sargus and D. annulatus. Paz (1982) expanded the Diplodus sargus species group to include two additional subspecies from the South Atlantic, four from the western Atlantic, and two from the Indian Ocean. The suspected hybrid status of the population in Israel was not mentioned again in this paper or in subsequent literature.
When we examined specimens of Diplodus from Israel, our attention was drawn to the identity of specimens previously named Diplodus sargus (non Linnaeus, 1758), which were found to be distinct from “typical” Diplodus sargus from other parts of the Mediterranean Sea. The analysis of these individuals demonstrated that the population from Israel represents a separate species, which is described in the present paper.
MATERIALS AND METHODS
Specimens were examined at the Academy of Natural Sciences of Philadelphia (ANSP); Hebrew University, Jerusalem (HUJ); Muséum National d’Histoire Naturelle, Paris (MNHN); Museo Civico di Storia Naturale “Giacomo Doria”, Genoa, Italy (MSNG); Staatliches Museum für Naturkunde, Stuttgart (SMNS); National Museum of Natural History, Washington, D.C. (USNM); and Museum für Naturkunde der Humboldt- Universität, Berlin (ZMB).
Descriptive methods follow Caldwell (1965). Proportional measurements and counts for the holotype of Diplodus levantinus n. sp. are given first, followed by those of the paratypes in parentheses. Following the method of Fricke et al. (2007), subspecies are no longer used: they are either raised to the species level or synonymized. The classification is based on Eschmeyer and Fricke (2015) and references follow Fricke (2015). The museum abbreviations follow Fricke and Eschmeyer (2015).
Holotype: HUJ 20535, 184.1 mm SL, eastern Mediterranean Sea, Israel, off Jaffa, ca. 32°03’N 34°45’E, collected by local fishermen, 30 Oct. 2015.
Paratypes: Eastern Mediterranean Sea, Israel: HUJ 2134 (1, 176 mm SL), near mouth of Wadi Hadera, Heinz Steinitz, 24 Oct. 1955. HUJ 6399 (7, 50-72 mm SL), Caesarea, Adam Ben-Tuvia, 25 Sept. 1951. HUJ 6402 (2, 50-76 mm SL), Tanturra, Adam Ben-Tuvia, 4 Apr. 1955. HUJ 6403 (2, 70-76 mm SL), Netanya, Adam Ben-Tuvia, 12 Dec. 1951. HUJ 7462 (1, 231 mm SL), Caesarea, Asher Ziv, 12 Aug. 1959. HUJ 14441 (1, 139.4 mm SL), Haifa, Daniel Golani, 5 June 1990. HUJ 19859 (3, 100-114 mm SL), Haifa, Daniel Golani, 28 July 2009. HUJ 20257 (8, 54-95 mm SL), Sdot Yam, 0.5-2.0 m depth, Daniel Golani, 21 Jan. 2014. HUJ 20282 (2, 22-30 mm SL), Jaffa, Daniel Golani, 11 Sept. 2013. HUJ 20344 (1, 128 mm SL), near mouth of Wadi Hadera, Israel, Heinz Steinitz, 24 Oct. 1955. MSNG 58305 (1, 107 mm SL), Haifa, Daniel Golani, 28 July 2009.
<italic>Diplodus levantinus</italic> n. sp., HUJ 20535, holotype, 184.1 mm SL, off Jaffa, Israel, 30 Oct. 2015. Lateral view; photograph of the fresh colouration taken a few hours after collection (Photograph: D. Golani).
Diagnosis. A species of Diplodus Rafinesque 1810 with 11-12 spines and 10-16 soft rays in the dorsal fin, 3 spines and 11-13 soft rays in the anal fin, 15-17 pectoral fin rays, 6-9 + 8-12 gill rakers on the first gill arch, upper and lower jaws with a single row of 4 incisors on each side, followed by a total of 16-19 molariform teeth in the upper jaw and 12-14 molariform teeth in the lower jaw, with the molariforms of the upper jaw separated from the incisors by a wide gap, and the sides of the body in adults with 8 vertical bars of equal width, which are present even in large adults, followed by a broad bar on caudal peduncle, which usually nearly reaches the ventral margin of the caudal peduncle.
Description. Dorsal fin-ray formula XII, 13 (XI to XII, 10 to 16, usually XII, 13 to 14). Anal fin ray formula III, 13 (III, 11 to 13). Pectoral fin ray formula, all elements, 16-16 (15 to 17). Gill rakers 6+9 (6-9+8-12) on first gill arch.
Selected body proportions, included in Table 1, are part of the description.
Selected body proportions of the holotype and selected paratypes of <italic>Diplodus levantinus</italic> n. sp., expressed in percentage of standard length.
<italic>Diplodus levantinus</italic> n. sp., HUJ 20257, paratype, specimen 1, 95 mm SL, Sdor Yam, Israel, 21 Jan. 2014. Lateral view; photograph of the fresh colouration taken a few hours after collection (Photograph: D. Golani).
<italic>Diplodus levantinus</italic> n. sp., HUJ 2134, paratype, 176.3 mm SL, near mouth of Wadi Hadera, Israel, 24 Oct. 1955. Lateral view; photograph of the colouration in preservative taken 60 years after preservation (Photograph: D. Golani).
Body completely scaled; scales ctenoid. Predorsal scales extending to a point above about centre of eye. Cheeks scaled. Snout scaleless. A long ventral axillary scale. Inter-ray membranes of dorsal, anal, pelvic and pectoral fins scaleless. Small scales extending about three-fourths the way out from base of caudal fin. Lateral line a smooth shallow convex curve from its origin to just beyond end of dorsal fin, then nearly straight to fold formed when tail is bent upward. Four scales (two to four) in a line angling upward, extending onto basal portion of caudal fin. Lateral line scales 68 (59-72). Eight (eight or nine) scales above lateral line to origin of dorsal fin, 16 (16-17) below to origin of anal fin, and seven (seven) above highest curve of lateral line to base of dorsal fin. Maximum observed standard length 231 mm.
Dorsal outline a regular curve from tip of snout to end (insertion) of dorsal fin, with only a slight concavity above eye and a slight convexity in front of eye. Snout slightly pointed. Ventral outline slightly convex (nearly straight) from tip of snout to pelvic fin origin, thence nearly straight to anal fin origin, and after angling upward at an angle of about 40°, convex to end (insertion) of anal fin. Dorsal and anal fins low. Pectoral fin long, reaching nearly to or past origin of anal fin, and increasing in relative length with increase in body length. Caudal fin forked. Mouth small, maxillary not reaching anterior margin of orbit in small specimens. Anterior nostril round; posterior nostril an elongate oval, the opening slit-like.
A single outer row of incisor teeth, four on each side both upper and lower, and none notched; upper portion of each tooth essentially rectangular in outline above a narrowed base bearing a posterior buttress on inner surface; teeth protrude and are rather strongly incurved towards cutting edge; anterior corner of each lateral tooth elevated. A mosaic of small molariform teeth in posterior part of mouth, 16 (16-19) in upper jaw, 12 (12-14) in lower jaw; anterior part immediately behind incisor teeth is naked; behind these, in lateral posterior part of mouth, there are three (two or three) rows of molariform teeth above, and three (two or three) rows below.
Vertebrae (Fig. 4): 10 precaudal plus 13 caudal plus one hypural, equalling 10 plus 14 as usually recorded. Three predorsal bones. Two dorsal spines situated on first pterygiophore, thereafter one spine on each pterygiophore (a count of 0-0-0-2 as discussed by Smith and Bailey, 1961).
<italic>Diplodus levantinus</italic> n. sp., HUJ 20535, holotype, 184.1 mm SL, off Jaffa, Israel, 30 Oct. 2015. Lateral view; x-ray (Photograph: I. Aizenberg).
Colour in alcohol. For pigmentation of body refer to Figures 1-2, which are part of the description.
Head and body silver, scales with horizontal rows of dark pigment. Head dorsally dark grey between the eyes, and a vertical dark streak below the eye. Posterior edge of opercle dark. A dark pectoral axillary blotch. Caudal peduncle with a broad dark grey streak that often reaches to the lower midline. Sides of body with about eight narrow vertical bars that extend across the upper three-fourths of the body; the posterior two bars are present but not very well visible on the photograph of the holotype due to light reflections (Fig. 1). Pectoral fins immaculate. Inter-spine and inter-ray membranes of dorsal, anal and caudal fins tinged with dark pigment, which is more intense towards bases of fins. Anterior inter-ray membranes of pelvic fins dark, more so than those of dorsal, anal and caudal fins; the spine and the first membrane are white, however, as are the posterior membranes. Caudal fin distally dark grey.
Etymology. The name of the new species, levantinus, refers to the Levant, a historical name for the coasts of the eastern Mediterranean.
Distribution and habitat. Known only from the coast of Israel, eastern Mediterranean Sea, between Haifa and Jaffa (Fig. 5). The species is found from shallow water to 50 m depth, usually on sand bottom near rocks; juveniles are found above sandy substrate near rocks at depths of 0.2-2 m.
Geographical distribution of the species of the <italic>Diplodus sargus</italic> group in the eastern Atlantic, Mediterranean Sea and southwestern Indian Ocean. Circles, <italic>Diplodus sargus</italic>; triangles, <italic>D. capensis</italic>; squares, <italic>D. cadenati</italic>; black stars, <italic>D. levantinus</italic> n. sp.; white stars, <italic>D. lineatus</italic>.
Comparisons. The dentition and colouration of the eastern Atlantic and Mediterranean species are compared in Figs 6-7 and Table 2. The new species is distinguished from D. sargus (Mediterranean Sea except Israel; Figs 8, 6A, 7A-C) by the number of molariform teeth in the upper and lower jaws in juveniles and adults (D. sargus: upper jaw 28-32, 4 rows; lower jaw 20-28, 4-5 rows. D. levantinus n. sp.: upper jaw 16-19, 2-3 rows; lower jaw: 12-14, 2-3 rows), the molariform teeth which do not extend to the base of the incisors in the upper jaw but leave a wide gap (nearly extending to the base of the incisors in D. sargus), the presence of vertical bars on the sides of the body in large specimens (usually absent in D. sargus), and sides of body in moderate and small adults with bars of equal width (usually alternating wide and narrow bars in D. sargus). It differs from D. cadenati (eastern Atlantic; Figs 9, 6C, 7G-I) in the number of molariform teeth in the upper and lower jaws in juveniles and adults (D. cadenati: upper jaw 39-43, 4-5 rows; lower jaw: 27-31, 4-6 rows. D. levantinus n. sp.: upper jaw 16-19, 2-3 rows; lower jaw: 12-14, 2-3 rows), the molariform teeth which do not extend to the base of the incisors in the upper jaw but leave a wide gap (extending to the base of the incisors in D. cadenati), sides of body in large and moderate adults with bars of equal width (usually alternating wide and narrow bars in D. cadenati), sides of body in small adults with 8 vertical bars (with about 5 vertical bars in D. cadenati), and the black blotch on the caudal peduncle usually nearly reaching the ventral margin (usually only covering the upper three-fourths in D. cadenati). It is distinguished from D. lineatus (Cape Verde Islands; Figs 10, 6E, 7M-O) by the number of molariform teeth in the upper and lower jaws in juveniles and adults (D. lineatus: upper jaw 8-10, 1-2 rows; lower jaw: 3-4, 2 rows. D. levantinus n. sp.: upper jaw 16-19, 2-3 rows; lower jaw: 12-14, 2-3 rows), sides of body in small adults with 8 vertical bars (with 4-5 vertical bars in D. lineatus), and the black blotch on the caudal peduncle usually nearly reaching the ventral margin (usually only covering the upper half to upper three-fourths in D. lineatus). It differs from D. capensis (South Africa; Figs 6B, 7D-F) in the number of molariform teeth in the upper and lower jaws in juveniles and adults (D. capensis: upper jaw 36-41, 4-5 rows; lower jaw: 19-22, 3-4 rows. D. levantinus n. sp.: upper jaw 16-19, 2-3 rows; lower jaw: 12-14, 2-3 rows), the molariform teeth, which do not extend to the base of the incisors in the upper jaw but leave a wide gap (extending to the base of the incisors in D. capensis), the presence of vertical bars on the sides of the body in large specimens (absent in D. capensis), and sides of body in moderate and small adults with 8 vertical bars (with 9 vertical bars in D. capensis). The new species is distinguished from D. argenteus (western Atlantic), D. ascensionis (Ascension Island), D. bermudensis (western Atlantic), D. caudimacula (western Atlantic), D. helenae (St. Helena), D. holbrookii (western Atlantic), D. kotschyi (northwestern Indian Ocean) and D. noct (Red Sea) by the presence of vertical bars on the sides of the body in large specimens (absent in D. argenteus, D. ascensionis, D. bermudensis, D. caudimacula, D. helenae, D. holbrookii, D. kotschyi and D. noct), sides of body in moderate and small adults with 8 vertical bars (with 9 vertical bars in D. argenteus, D. bermudensis, D. caudimacula, D. helenae, D. kotschyi, and D. noct), and the black blotch on the caudal peduncle usually nearly reaching the ventral margin (usually only covering the upper three-fourths in D. argenteus, D. ascensionis, D. bermudensis, D. caudimacula, D. holbrookii, D. noct and D. kotschyi, in large specimens restricted to a round blotch in the middle in D. kotschyi and D. noct).
Dentition of the species of the eastern Atlantic and Mediterranean species of the <italic>Diplodus sargus</italic> group and similar species; (above) upper jaw, (below) lower jaw. A, <italic>Diplodus sargus</italic>, SMNS 9153, specimen 1, 190 mm SL, Croatia, Istria, 9 June 1978. B, <italic>Diplodus capensis</italic>, SMNS 25672, 1 specimen, 176 mm SL, South Africa. C. Diplodus cadenati, HUJ 20534, specimen 1, 198 mm SL, Senegal, Dakar, Apr. 2015. D, <italic>Diplodus levantinus</italic> n. sp., HUJ 7462, 1 paratype, 231 mm SL, Israel, Caesarea, 12 Aug. 1959. E, <italic>Diplodus lineatus</italic>, HUJ 20368, specimen 1, 222 mm SL, Cape Verde Islands, Boa Vista Island, 9 Sept. 2014. F, <italic>Diplodus fasciatus</italic>, HUJ 20369, specimen 1, 199 mm SL, Cape Verde Islands, Boa Vista Island, 9 Sept. 2014. G, <italic>Diplodus prayensis</italic>, HUJ 20370, 199 mm SL, Cape Verde Islands, Boa Vista Island, 9 Sept. 2014. H, <italic>Diplodus annularis</italic>, SMNS 995, specimen 3, 181 mm SL, Italy, Trieste, Dec. 1862. Scales indicate 3 mm.
Generalised body colour patterns of the species of the <italic>Diplodus sargus</italic> group; large adults, medium adults and small adults. <italic>Diplodus sargus</italic>: A, MNHN A-8098 (255 mm SL), France, Marseille; B, MNHN 1961-0894 (192 mm SL), Monaco, 1961. C, MNHN uncat. (92 mm SL), France, Villefranche-sur-Mer. <italic>Diplodus capensis</italic>: D, MNHN A-8096 (315 mm SL), South Africa, Cape Province. E, MRAC 97894 (215 mm SL), South Africa, Elephant Bay. F, SAIAB 1638 (109 mm SL), South Africa, Cape Province. <italic>Diplodus cadenati</italic>: G, BMNH 1895.5.28.24 (310 mm SL), Madeira. H, BMNH 1858.8.3.7-8 (182 mm SL), Canary Islands. I, IFAN 867 (89 mm SL), Senegal, Dakar. <italic>Diplodus levantinus</italic> n. sp.: J, HUJ 7462 (281 mm SL, paratype), Israel, Caesarea, 12 Aug. 1959. K, HUJ 2134 (176.3 mm SL, paratype), Israel, Wadi Hadera, 24 Oct. 1955. L, HUJ 20257 (spec. 1, 95 mm SL, paratype), Israel, Sdot Yam, 21 Jan. 2014. <italic>Diplodus lineatus</italic>: M, HUJ 20368 (spec. 1, 222 mm SL), Cape Verde Islands, Boa Vista, 9 Sept. 2014. N, HUJ 20369 (spec. 2, 177 mm SL), Cape Verde Islands, Boa Vista, 9 Sept. 2014. O, MNHN 1971-0002 (98 mm SL), Cape Verde Islands. This figure follows <xref ref-type="bibr" rid="CIT49">Paz (1975</xref>: 47), but is corrected and updated; figures A-I and O based on <xref ref-type="bibr" rid="CIT49">Paz (1975</xref>: Fig. 23). Scales indicate 20 mm.
Comparison of eastern Atlantic and Mediterranean species of the <italic>Diplodus sargus</italic> species group.
D. levantinus n. sp.
D. sargus
D. cadenati
D. lineatus
D. capensis
Dorsal fin rays
XI-XII, 10-16
XI-XIII, 12-15
XI-XIII, 12-15
XI-XIII, 12-15
XI-XIII, 13-16
Anal fin rays
III, 11-13
III, 12-14
III, 12-14
III, 12-14
III, 13-14
Pectoral fin rays
15-17
15-17
16-17
15-18
15-17
Lateral line scales
59-72
57-68
58-67
57-66
61-69
Number of molariform teeth in upper jaw
16-19
28-32
39-43
8-10
36-41
Rows of molariform teeth in upper jaw
2-3
4
4-5
1-2
4-5
Number of molariform teeth in lower jaw
12-14
20-28
27-31
3-4
19-22
Rows of molariform teeth in lower jaw
2-3
4-5
4-6
2
3-4
Extent of molariforms towards incisors in upper jaw
Not reaching to base of incisors, leaving a wide gap
Nearly reaching to base of incisors
Reaching to base of incisors
Not reaching to base of incisors, leaving a wide gap
Reaching to base of incisors
Extent of molariforms towards incisors in lower jaw
Not reaching to base of incisors, leaving a narrow gap
Nearly reaching to base of incisors
Reaching to base of incisors
Not reaching to base of incisors, leaving a wide gap
Reaching to base of incisors
Large adults: vertical bars on body
8 of equal width
None
Alternatingly 5 wide, 3-4 narrow, total 8-9
4 of equal width, 1 narrow
None
Medium-sized adults: vertical bars on body
8 of equal width
Alternatingly 4-5 wide, 3-4 narrow, total 8-10
Alternatingly 4-5 wide, 3-4 narrow, total 7-9
4 of equal width, 1 narrow
9 of equal width
Small adults: vertical bars on body
8 of equal width
Alternatingly 4-5 wide, 3-4 narrow, total 9-10
5 of equal width
4-5 of equal width
9 of equal width
Extent of dark blotch on caudal peduncle
Nearly reaching ventral margin
Usually only on upper three-fourths
Only on upper three-fourths
Only on upper half to upper three-fourths
Only on upper three-fourths
<italic>Diplodus sargus</italic>, HUJ 20542, 2 spec., 220-225 mm SL, France, Marseille, 8 Dec. 2015. Lateral view; photograph of the fresh colouration taken a few hours after collection (Photograph: R. Fricke).
<italic>Diplodus cadenati</italic>, HUJ 20532, 3 spec (upper to lower: 151, 149, 140 mm SL), Tenerife, Canary Islands, 25 July 2015. Lateral view; photograph of the fresh colouration taken a few hours after collection (Photograph: R. Fricke).
<italic>Diplodus lineatus</italic>, HUJ 20368, 4 spec (Upper to lower: 222, 176, 166, 162 mm SL), Boa Vista, Cape Verde Islands, 9 Sept. 2014. Lateral view; photograph of the fresh colouration taken a few hours after collection (Photograph: R. Fricke).
DISCUSSION
The southeastern Mediterranean (Levantine) populations previously attributed to Diplodus sargus clearly belong to a different species, based on differences in the dentition and colouration. The only junior synonym of D. sargus listed in the Catalogue of Fishes online (Eschmeyer and Fricke 2015) that was described from the eastern Mediterranean, Sargus raucusGeoffroy Saint-Hilaire, 1809, was first considered as the valid name for these; the species was illustrated from Egypt by Geoffroy Saint-Hilaire (1809: pl. 18, Fig. 1), and no type material is available. The material was collected in Alexandria, Egypt in 1798-1799, and when Alexandria was conquered by Britain, Geoffroy Saint-Hilaire refused to hand over the materials and documents to the British general Hutchinson, and later sent the material to Paris (Bauchot et al. 1990: 88). However, Bauchot and Daget (1971: 322) found two specimens of Sargus raucus from Alexandria (MNHN 0000-5740), which were used for the description of Geoffroy Saint-Hilaire (1827: 340), and identified them as Diplodus cervinus (Lowe, 1838). None of the specimens agrees with the holotype illustrated by Geoffroy Saint-Hilaire (1809: Pl. 38, Fig. 1), which is a smaller specimen and clearly belongs to the Diplodus sargus species group, so the holotype is not included in MNHN 0000-5740. The material collected by E. Geoffroy Saint-Hilaire in Egypt was exclusively sent to Paris, but no other material of the Diplodus sargus species group collected during the Egyptian mission is extant in the MNHN collection (see also Bauchot and Daget 1972: 69). We therefore conclude that the holotype is lost. A specimen from Bardawil Lagoon, Egypt (HUJ 6047), very similar in colouration to the one illustrated by Geoffroy Saint-Hilaire (1809), turned out to have the typical dentition of Diplodus sargus with irregularly grouped small molars reaching right to the front teeth, without a naked area in between. The specimen has a dorsal fin count of XII+13, an anal fin count of III+13, pectoral fin rays 16, gill rakers 5+8. In order to avoid nomenclatural confusion, and to fix the classification of southeastern Mediterranean species in the Diplodus sargus group, the specimen (HUJ 6047) is hereby designated as the neotype of Sargus raucus Geoffroy Saint-Hilaire, 1809 (Fig. 11), which is thus a junior synonym of Diplodus sargus (Linnaeus, 1758). The neotype originates from Bardawil Lagoon, northern Sinai, Egypt, which is as close as possible to the original type locality (Egypt). A second specimen from Bardawil Lagoon (HUJ 6057) also agrees in the dentition and general characters with Diplodus sargus. A third specimen, ZMB 1055 (70.5 mm SL) from Alexandria, Egypt, collected during the expedition of Hemprich and Ehrenberg, has likewise a dentition typical of Diplodus sargus. It can be concluded that Diplodus sargus is found along the Mediterranean coast of Egypt, while the species is replaced by D. levantinus n. sp. in Israel.
<italic>Diplodus sargus</italic>, HUJ 6047, neotype of <italic>Sargus raucus</italic> Geoffroy Saint-Hilaire 1809, 64.3 mm SL, Bardawil Lagoon, Egypt (Photograph: D. Golani).
The difference of the Israeli population from other species of the Diplodus sargus species group were first recognized in the doctoral dissertation of Paz (1975: 57), who provisionally named the specimens from Israel “Diplodus X” and assumed that they were hybrids between D. annularis × D. sargus, due to the dentition, which seemed intermediate between those species. However, the hybrid theory can be ruled out, as no true Diplodus sargus is found in Israel, and genetically they are clearly part of the Diplodus sargus species group, without a component of another species of Diplodus (Yaron Tikochinski, personal communication). Except in the work of Paz (1975), the distinctiveness of Israeli populations previously misidentified as D. sargus was neither mentioned nor discussed again in the literature, including the later paper of Paz (1982).
The body shape and colouration changes during growth in Diplodus spp.; juveniles are more slender, adults more high-bodied, and in some species the number of stripes along the sides of the body increases. Allometric growth in Diplodus sargus compared with two other species of the genus was analysed by Loy et al. (2001). Allometric growth is also obvious in Diplodus levantinus n. sp., but the number of stripes along the sides of the body does not change from juveniles to adults in this species.
Considering the genetics of species in the Diplodus sargus species group, no genetic differences were found between Diplodus sargus and D. cadenati by Domingues et al. (2007). They agreed with Bargelloni et al. (2005) who considered two hypotheses to explain this lack of differentiation. Either D. sargus is a recent immigrant in the Mediterranean or historical bottlenecks and recolonization processes prevented strong differentiation of Atlantic and Mediterranean basins. In a genetic study of several species of Diplodus, including D. levantinus n. sp., D. sargus and D. lineatus, no significant differences were found among these species (Yaron Tikochinski, personal communication). On the other hand, the Diplodus sargus species group is well distinguished from other species of Diplodus (see Summerer et al. 2001, Casu et al. 2009). However, González-Wangüemert et al. (2006) detected diagnostic alleles differentiating between Diplodus sargus and D. cadenati; additional molecular analyses should be performed for the species group. Apart from potential genetical differences, the species in the Diplodus sargus species group are sufficiently distinguished by their dentition and colouration.
The restricted range of Diplodus levantinus n. sp. appears to represent a relict distribution; its particular dentition recalls that of D. lineatus from the Cape Verde Islands, which is also considered a relict distribution. Species with a stronger dentition (including D. sargus in the Mediterranean) seem to have spread over most of the range of the species group, leaving the species with weaker dentition in such small, restricted areas at the margin. The range of D. levantinus is separated from that of D. sargus by wide stretches of sandy shore without suitable habitat in the south; the situation in the north of its distribution range remains unknown.
Checklist of species of the genus <italic>Diplodus</italic>
Diplodus annularis (Linnaeus, 1758)
Sparus unicolor flavescens, macula nigra annulari ad caudam Artedi 1738a: 37, No. 12 (based on Artedi 1738b; Sparus of authors).
Sparus unicolor flavescens, macula nigra annulari ad caudam Artedi 1738b: 57, No. 1 (based on Ruàs of Aristoteles and Aelianus; Sparos of Athenaeus, Oppianus and Eustathius; Sparus of Plinius, Albertinus, Cubae, Bellonius, Rondeletius, Salviani, Aldrovandius, Jonston, Charleton, Willughby, Rajus; Marinus of Gesner; Sparulus of Ovidius; Fluta and Sargus of Gaza; Caspargus of Aelianus).
Sparus annularisLinnaeus, 1758: 278 (in mari Adriatico, infero/Adriatic Sea; no types known; based on ‘Art. gen. 37, syn. 57, Sparus unicolor flavescens ...).
Charax puntazzo var. ponticusNordmann, 1840: 387, pl. 4, Fig. 2 (Sevastopol, Crimea, Ukraine, Black Sea; no types known).
Puntazzo annularisBleeker, 1876: 284 (Mediterranean Sea; no types known; unneeded new name for Charax puntazzo of Cuvier and Valenciennes, =Sparus puntazzoWalbaum, 1792).
Distribution: Mediterranean Sea, Black Sea; eastern Atlantic from Sierra Leone north to southern Bay of Biscay, including Cape Verde and Canary islands.
Diplodus sargus (Linnaeus, 1758)
Sparus lineis transversis varius, macula nigra insigni ad caudam Artedi 1738a: 37, No. 13 (based on Artedi 1738b, No. 58; Sargus of authors).
Sparus lineis transversis varius, macula nigra insigni ad caudam Artedi 1738b: 58, No. 2 (Greece, France, Italy, Venice; based on O’Sargos of Aristoteles, Aelianus, Oppianus and Atheneus; Sargus of Ennius, Ovidius, Plinius, Juvenal, Bellonius, Rondeletius, Salviani, Gesner, Aldrovandius, Jonston, Charleton, Willughby and Rajus).
Sparus sargusLinnaeus, 1758: 278 (in M. infero, Adriatic Sea; no types known; based on ‘Art. gen. 37, syn. 58, Sparus lineis transversis varius ...).
Sparus cinctus Walbaum, 1792: 282 (Marseille, France; no types known; based on Sparus sp., ‘Sparus fascia subcaudali ...’ of Brünnich 1768: 39, No. 58).
Sargus raucusGeoffroy Saint-Hilaire 1809: Pl. 18, Fig. 1 (Alexandria, Egypt; neotype: HUJ 6047, as designated above.)
Distribution: southwestern Indian Ocean: Mauritius, Rodrigues.
Diplodus vulgaris (Geoffroy Saint-Hilaire, 1817)
Sargus noster primus ... Aldrovandi, 1638: Fig. [p. 174].
Sargus vulgarisGeoffroy Saint-Hilaire, 1817: 342, Pl. 18, Fig. 2 (Alexandria, Egypt; no types known).
Sparus aldrovandiNardo, 1827: 21, 38 (no. 117) (Adriatic Sea; no types known; based on Aldrovandi 1638, p. 174).
Sargus salviani Valenciennes in Cuvier and Valenciennes, 1830: 28 [Mediterranean Sea; part, based on Sargus of Salviani 1558, p. 179; Aldrovandi 1638, p. 174; Sargus vulgaris of Geoffroy Saint-Hilaire 1817, pl. 18, Fig. 2; syntypes: MNHN 8533 (1), 8534 (1), 8549 (2)].
Distribution: Mediterranean Sea, southwestern Black Sea; eastern Atlantic from Canary Islands and Madeira north to Brittany (France); Angola.
Key to eastern Atlantic and Mediterranean species of the <italic>Diplodus sargus</italic> group
1
Each side of upper jaw with a total of 28-43 molariform teeth in 4-5 principal rows; each side of lower jaw with a total of 20-31 molariform teeth in 3-6 principal rows; molariform teeth in upper jaw reaching to base of incisors
2
–
Each side of upper jaw with a total of 8-19 molariform teeth in 1-3 principal rows; each side of lower jaw with a total of 3-14 molariform teeth in 2-3 principal rows; molariform teeth in upper jaw not reaching to base of incisors, separated by a distance of at least 2 lengths of the largest molariform tooth
4
2
Each side of upper jaw with a total of 28-32 molariform teeth
Diplodus sargus
–
Each side of upper jaw with a total of 36-43 molariform teeth
3
3
Each side of lower jaw with a total of 27-31 molariform teeth in 4-6 principal rows; sides of body in large adults with alternating broad and narrow vertical dark bars, in addition to dark blotch on caudal peduncle
Diplodus cadenati
–
Each side of lower jaw with a total of 19-22 molariform teeth in 3-4 principal rows; sides of body in large adults without vertical dark streaks, in smaller adults with about 9 vertical dark bars of even width, in addition to dark blotch on caudal peduncle
Diplodus capensis
4
Each side of upper jaw with a total of 8-10 molariform teeth in 1-2 principal rows; each side of lower jaw with a total of 3-4 molariform teeth in 2 principal rows; molariform teeth in lower jaw not reaching to base of incisors, separated by a distance of at least 2 lengths of the largest molariform tooth; sides of body with 4 broad and one narrow vertical dark bars, in addition to dark blotch on dorsal half of caudal peduncle
Diplodus lineatus
–
Each side of upper jaw with a total of 16-19 molariform teeth in 2-3 principal rows; each side of lower jaw with a total of 12-14 molariform teeth in 2-3 principal rows; molariform teeth in lower jaw nearly reaching to base of incisors; sides of body with about 8 narrow vertical dark bars of even width, in addition to dark blotch nearly all across caudal peduncle
Diplodus levantinus n. sp.
ACKNOWLEDGEMENTS
We are grateful to Dr. I. Aizenberg (Veterinary Teaching Hospital, Koret School of Veterinary Medicine, Hebrew University of Jerusalem, Israel) for providing a radiograph of the holotype of Diplodus levantinus n. sp. We thank Rui Freitas (Universidade de Cabo Verde), who assisted in acquiring endemic Diplodus species from the Cape Verde Islands. We are grateful to Alberto Brito (Universidad de La Laguna, Tenerife, Spain) for his hospitality, and for collecting D. cadenati from Tenerife and Senegal. Dominique Didier (ANSP, Philadelphia), Marie-Louise Bauchot and Martine Desoutter (MNHN, Paris), Giuliano Doria (MSNG, Genoa), Jeffrey T. Williams (USNM, Washington DC) and Peter Bartsch (ZMB, Berlin) gave access to materials in their care.
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