Blache’s flounder,
Se vuelve a describir el pez plano de Blache’s
The lefteye flounders of the family Bothidae are a large group of marine fishes living in tropical and temperate waters of all oceans. They are arranged in the order Pleuronectiformes (
The genus
Species | Depth range (m) | Geographical distribution | Remarks |
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shallow | EP: Juan Fernandez and Desaventurados Islands | |
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6-38 | EI, WP: Bass Strait and Tasmania, Australia | |
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20-30 | WP: East China Sea | |
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1-50 | MS: Mediterranean and Black Sea | Junior synonym: |
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20-40 | RS: probably Gulf of Suez. MS: southeastern Mediterranean | |
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26-50 | WP: New South Wales and Queensland, Australia | |
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5-60 | EI: Southwestern Australia | |
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10-70 | EI, WP: South Australia to New South Wales, Australia | |
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30-71 | EI, WP: Persian Gulf east to Philippines and New Guinea | |
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16-90 | EI, WP: Arafura Sea and Queensland (Australia) | |
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18-141 | RS, WI, EI: Central Red Sea, Persian Gulf and Madagascar east to Indonesia | Recorded from Eritrea, Red Sea by Dor (1970) as |
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80-100 | EI, WP: Northern Australia to southern Japan | |
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5-200 | EI, WP: Southwestern to southeastern Australia | |
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34-175 | EA: Guinea-Bissau south to Angola | Revalidated in the present paper; previously synonymized with |
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85-154 | EI, WP: Southern Japan to northern Australia | |
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45-200 | EA, MS: Mediterranean and Black seas; eastern Atlantic from Norway to Angola | |
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50-220 | WI, EI, WP: South and East Africa east to Loyalty Islands | |
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70-200 | EA, WI: Morocco to South Africa, Saint Helena and Ascension Is. | Synonym: |
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83-220 | RS, WI: southern Red Sea and northwestern Indian Ocean | |
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5-300 | EA, MS: Mediterranean and Black seas, and eastern Atlantic from Ireland to Sierra Leone | Synonym: |
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100-224 | EI, WP: Taiwan to Indonesia and northwestern Australia | |
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138-225 | WI: northwestern Indian Ocean | Synonym: |
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15-350 | EA, MS: western Mediterranean and eastern Atlantic from Scotland to northern Mauritania, including Azores and Madeira | |
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125-254 | WI: Saya de Malha Bank and Madagascar | |
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4-390 | EI, WP: India east to New Caledonia, north to Japan | |
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160-265 | EP: Nazca Ridge | |
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215 | WP: Japan; Chesterfield Islands | |
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187-292 | WP: Philippines to Vanuatu | |
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240-250 | WP: New Caledonia | |
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300 | WP: Chesterfield Islands | |
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150-440 | WP: Indonesia east to Hawaiian and Line Islands | |
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4-737 | WP: New Zealand | Junior synonym: |
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363-383 | RS: Central Red Sea | |
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85-897 | EA, MS: Mediterranean Sea and eastern Atlantic from Gibraltar to western Sahara | |
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? | EI: Tasmania, Australia | |
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? | EI, WP: Sumatra (Indonesia), China |
While examining the catch of R/V
Morphological descriptions. Counts and measurements follow
Comparative material.
Genetic analyses. A piece of the right pectoral fin was removed from fresh specimens and preserved in 96% ethanol. A total of 19 samples were used for molecular analyses including specimens of
Species | Sample ID | Deep | Latitude | Longitude | Area | COI | 12S |
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OP345057 |
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OP345058 |
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OP345059 |
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ArcaM77 | 118 | 10.5788 | 17.0778 | Atlantic | OP341523 | OP345065 |
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Arca77b | 118 | 10.5788 | 17.0778 | Atlantic | OP341524 | OP345066 |
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Arca77a | 118 | 10.5788 | 17.0778 | Atlantic | OP341525 | OP345067 |
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Aglm76 | 131 | 39.7978 | 4.4397 | Mediterranean | OP341518 | OP345060 |
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AgIm76_2 | 131 | 39.7978 | 4.4397 | Mediterranean | OP341519 | OP345061 |
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AgIm76_3 | 131 | 39.7978 | 4.4397 | Mediterranean | OP341520 | OP345062 |
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AgIm76 | 131 | 39.7978 | 4.4397 | Mediterranean | OP341521 | OP345063 |
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AgIm76_4 | 131 | 39.7978 | 4.4397 | Mediterranean | OP341522 | OP345064 |
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Agla11 | 61 | 40.3105 | 0.608 | Mediterranean | OP341526 | OP345068 |
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Agla11_2 | 61 | 40.3105 | 0.608 | Mediterranean | OP341527 | OP345069 |
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Agla71 | 112 | 39.9335 | 3.3058 | Mediterranean | OP341528 | OP345070 |
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AgRo73 | 276 | 40.1917 | 4.1132 | Mediterranean | OP341532 | OP345074 |
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AgRo73_2 | 276 | 40.1917 | 4.1132 | Mediterranean | OP341533 | OP345075 |
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Agth5 | 95 | 39.8313 | 0.7073 | Mediterranean | OP341529 | OP345071 |
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Agth5_2 | 95 | 39.8313 | 0.7073 | Mediterranean | OP341530 | OP345072 |
DNA was extracted from the tissue using the DNeasy Blood and Tissue Extraction kit (Qiagen, West Sussex, UK). Polymerase chain reaction (PCR) was used to amplify two partial mitochondrial genes (mtDNA):
Sequences were imported into BioEdit 7.0.5.2. (
Genetic distance (
The sequences were concatenated and used to determine the phylogenetic relationships between the
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Other material: CFM_IEOMA 2102 (1), 2832 (2), 2838 (2), 7766 (4), 7767 (5), 7768 (2); SMNS 27381 (2), 27382 (2), 27383 (1) | |||
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Spec. 1, BMNH 1930.5.6.34 | Spec. 2, BMNH 1930.5.6.35 | ||||
Sex | Male | Female | Female | Males (n = 11) | Females (n = 9) |
Dorsal-fin rays | 95 | 94 | 91 | 89-96 | 89-99 |
Anal-fin rays | 71 | 64 | 69 | 66-75 | 70-77 |
Pectoral-fin rays (ocular side) | 10 | 10 | 10 | 10-12 | 11 |
Pectoral-fin rays (blind side) | 9 | 7 | 7 | 9-10 | 9-10 |
Caudal-fin rays | iii,11,iii | iii,11,iii | iii,11,iii | iii,11,iii (1 specimen: iii,12,ii) | iii,11,iii |
Lateral-line scales | 55 | 52 | 55 | 53-56 | 53-57 |
Gill rakers on first arch, lower branch (ocular side) | 6 | 6 | 6 | 5-8 | 6-7 |
Vertebrae | 10 + 31 | 10 + 30 | 10 + 30 | 10 + 31-33 | 10 + 31-33 |
SL | 73.7 | 66.7 | 62.0 | 66.7-110.8 | 89.0-107.0 |
Head length (ocular side) | 24.6 | 26.1 | 26.0 | 23.0-27.8 | 22.9-25.1 |
Body depth | 39.6 | 42.4 | 41.4 | 39.9-45.6 | 39.7-43.8 |
Upper orbit diameter | 8.4 | 8.7 | 9.1 | 6.9- 9.6 | 6.5- 8.5 |
Lower orbit diameter | 8.2 | 8.5 | 9.3 | 6.9- 9.0 | 6.5- 8.3 |
Interorbital distance | 0.9 | 0.8 | 0.4 | 0.4- 0.7 | 0.3- 0.5 |
Upper jaw length (ocular side) | 9.4 | 8.4 | 9.6 | 7.0- 8.6 | 7.0- 8.1 |
Lower jaw length (ocular side) | 6.7 | 6.6 | 7.3 | 5.2- 7.0 | 5.1- 6.1 |
Lower jaw length (blind side) | 6.9 | 7.2 | 8.3 | 5.2- 6.8 | 5.3- 5.9 |
Caudal peduncle depth | 9.4 | 11.2 | 10.2 | 8.6-11.3 | 8.8- 9.5 |
Length of longest dorsal-fin ray | 22.8 | 12.4 | 13.6 | 23.9 -34.7 | 7.3-10.9 |
Length of longest anal-fin ray | 9.6 | 10.3 | 13.4 | 10.2-12.7 | 10.4-11.5 |
Pectoral-fin length (ocular side) | 11.4+ | 16.2 | 15.7 | 14.4-17.3 | 15.4-17.5 |
Pectoral-fin length (blind side) | 6.4 | 6.2 | 6.7 | 7.2- 8.3 | 6.7- 7.1 |
Pelvic-fin length (ocular side) | 13.9 | 12.7 | 10.3 | 14.2-15.6 | 11.5-16.2 |
Pelvic-fin length (blind side) | 8.2 | 8.0 | 10.8 | 8.5-11.3 | 8.1-10.5 |
Length of pelvic-fin base (ocular side) | 7.9 | 6.8 | 6.4 | 7.0- 9.4 | 6.5- 8.8 |
Length of pelvic-fin base (blind side) | 4.6 | 4.8 | 6.9 | 3.3- 4.5 | 2.4- 4.0 |
Length of longest caudal-fin ray | 14.2+ | 18.6 | 18.8 | 17.6-20.1 | 17.4-19.5 |
Photographs of preserved specimen taken by L. Goodayle. Above: Ocular side. Below: blind side. Scale indicated 5 cm.
Photographs of fresh specimens taken by Pere Oliver Reus. Ocular side.
Photographs of fresh specimens taken by Pere Oliver Reus. Blind side.
Body oval, laterally strongly compressed, eyes on the left side of body (“ocular side”), right side of body without eyes (“blind side”). Body depth 2.5 [2.2-2.5] in SL, 39.6 [39.7-45.6]% of SL. Head length of ocular side 4.1 [3.6-4.4] in SL, 24.6 [22.9-27.8]% of SL. Snout slightly pointed, not completely scaled, tip naked
Dorsal fin with 94 [89-99] soft rays, the second to fifth rays elongate and filamentous in the male. Anal fin with 73 [66-77] soft rays. Caudal fin distally rounded, caudal fin rays iii,11,iii [iii,11,iii]. Length of longest caudal-fin ray 7.0- [5.0-5.7] in SL, 14.2+ [17.4-20.7]% of SL, damaged in lectotype. Pectoral fin with 10 [10-12] soft rays on ocular side, with 9 [9-10] rays on blind side. Pectoral-fin length 8.7- [5.7-6.6] in SL, 11.4+ [14.4-17.5]% of SL on ocular side, damaged in lectotype, 12.2 [12.0-16.1] in SL, 8.2 [6.2-8.3]% of SL on blind side. Pelvic-fin length 12.6 [6.4-9.7] in SL, 7.9 [10.3-15.6]% of SL on ocular side, 12.2 [8.8-12.5] in SL, 8.2 [8.0-11.3]% of SL on blind side.
A specimen recorded as
This species is distributed from Morocco and the Canary Islands southward along the coast of West Africa. Populations in the northeastern Atlantic from the Strait of Gibraltar, Madeira and the Azores northward, including the Mediterranean Sea, belong to
The bright red colour of the blind side of males of
(
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Neotype of |
Other material: SMNS 27384 (2), 27385 (2), 27386 (4), 27390 (1), 27393 (1), 27396 (1) | ||
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Sex | Male | Male | Males (n = 9) | Females (n = 2) |
Dorsal-fin rays | 101 | 96 | 98-101 | 97-98 |
Anal-fin rays | 75 | 74 | 74-79 | 76-77 |
Pectoral-fin rays (ocular side) | 10 | 10 | 11 | 10-11 |
Pectoral-fin rays (blind side) | 10 | 7 | 9-10 | 9-10 |
Caudal-fin rays | ii,13,ii | ii,13,ii | ii-iii,11-13,ii-iii | iii,11,iii |
Lateral-line scales | 58 | 63 | 58-62 | 61 |
Gill rakers on first arch, lower branch (ocular side) | 8 | 8 | 7-9 | 8-9 |
Vertebrae | 10 + 31 | 10 + 32 | 10 + 31-32 | 10 + 33 |
Measurement: | ||||
SL | 118.7 | 110.8 | 92.0-146.9 | 92.0-104.0 |
Proportions (% of SL): | ||||
Head length (ocular side) | 24.8 | 25.4 | 24.7-25.7 | 24.8-26.3 |
Body depth | 38.7 | 40.8 | 36.1-43.9 | 36.6-39.0 |
Upper orbit diameter | 7.6 | 8.4 | 6.7- 8.4 | 7.1- 7.7 |
Lower orbit diameter | 7.5 | 8.2 | 6.7- 8.4 | 7.1- 7.6 |
Interorbital width | 0.7 | 1.0 | 0.4- 0.8 | 0.4 |
Upper jaw length ocular side) | 8.1 | 8.5 | 7.9- 8.7 | 8.5- 8.8 |
Lower jaw length (ocular side) | 5.7 | 7.4 | 5.7- 6.9 | 6.8- 7.0 |
Lower jaw length (blind side) | 5.5 | 6.3 | 5.5- 6.6 | 6.5- 7.1 |
Caudal peduncle depth | 9.3 | 8.7 | 8.5-10.2 | 8.8- 9.0 |
Length of longest dorsal-fin ray | 17.9 | 15.5 | 16.4-23.1 | 7.3- 7.9 |
Length of longest anal-fin ray | 11.5 | 9.5 | 11.0-11.5 | 9.1-12.9 |
Pectoral-fin length (ocular side) | 17.3 | 15.2 | 16.6-19.1 | 17.0-17.9 |
Pectoral-fin length (blind side) | 7.7 | 10.1 | 7.8-11.8 | 8.2 |
Pelvic-fin length (ocular side) | 19.1 | 15.3 | 13.6-19.1 | 15.2 |
Pelvic-fin length (blind side) | 11.9 | 12.4 | 10.2-11.9 | 10.1-11.0 |
Length of pelvic-fin base (ocular side) | 9.3 | 8.6 | 6.6- 9.9 | 8.0-8.8 |
Length of pelvic-fin base (blind side) | 4.4 | 7.4 | 3.4- 5.0 | 4.0- 5.0 |
Length of longest caudal-fin ray | 19.0 | 20.1 | 17.8-20.4 | 18.5-20.5 |
Photograph of preserved specimen. Ocular side.
Photograph of preserved specimen. Blind side.
Photographs of preserved specimen taken by L. Goodayle. Above: Ocular side. Below: blind side. Scale indicated 5 cm.
X-ray taken by J. Maclaine.
Photograph of fresh specimens taken by Francesc Ordines. Ocular side.
Photograph of fresh specimens taken by Francesc Ordines. Blind side.
Body oval, laterally strongly compressed, eyes on the left side of body (“ocular side”), right side of body without eyes (“blind side”). Body depth 2.6 (2.4) [2.3-2.8] in SL, 38.7 (40.8) [36.1-43.9]% of SL. Head length 4.0 (3.9) [3.8-4.2] in SL, 24.8 (25.4) [24.0-26.3]% of SL Snout slightly pointed, not completely scaled, tip naked
Dorsal fin with 101 (96) [97-101] soft rays, the second to fifth to seventh rays (male), second to third (female) elongate and filamentous. Anal fin with 75 (74) [74-79] soft rays. Caudal fin distally rounded, caudal fin rays ii,13,ii (ii,13,ii) [ii-iii,11-13,ii-iii]. Length of longest caudal-fin ray 5.3 (5.0) [4.9-5.6] in SL, 19.0 (20.1) [17.8-20.5]% of SL. Pectoral fin with 10 (10) [10-11] soft rays on ocular side, with 10 (7) [9-10] rays on blind side. Pectoral-fin length 5.8 (6.6) [5.2-7.3] in SL, 17.3 (15.2) [16.6-19.1]% of SL on ocular side, 13.0 (9.9) [8.4-12.9] in SL, 7.7 (10.1) [7.7-11.8]% of SL on blind side. Pelvic-fin length 5.2 (6.5) [5.1-7.4] in SL, 19.1 (15.3) [13.6-19.2]% of SL] on ocular side, 8.4 (8.0) [8.4-9.9] in SL, 11.9 (12.4) [10.1-11.9]% of SL on blind side.
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-- | 81.7 | 82.5 | 74.3 | 52.7 | 78.7 |
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15.62 | -- | 24.5 | 80.7 | 75.7 | 45.7 |
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15.78 | 4.69 | -- | 81.4 | 75.4 | 43.6 |
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14.21 | 15.42 | 15.56 | -- | 75.3 | 72.7 |
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10.07 | 14.47 | 14.42 | 14.40 | -- | 73.5 |
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15.04 | 8.73 | 8.34 | 13.89 | 14.05 | -- |
The phylogenetic reconstruction for concatenated fragments clearly separates
Posterior probabilities are shown as percentages and are indicated near the nodes. The samples of
During our studies on the synonymy of
This species apparently prefers cooler water; in the western Mediterranean, it is mainly found in regions with cool upwelling water, e.g. in southern France and Liguria, and off the northern coasts of Menorca or Sicily islands. In recent years, it has become rare in some of these habitats, so it may now be considered Near Threatened in the Mediterranean.
The specimens of
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Dorsal-fin rays | 89-99 |
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91 | 91-98 | 100-108 | 96-105 | 96-106 |
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Elongate dorsal fin rays | rays 2 to 5 | rays 2 to 6 |
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rays 1 to 8 (male), rays 1 to 2 (female) | rays 2 to 4 (male), none (female) |
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rays 2 to 5-7 (male), rays 2 to 3 (female) |
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Colour of filaments | light grey, rays with black margins, basally yellowish |
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brown | light brown | light brown |
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Anal-fin rays | 64-77 | 61-66 | 69 | 70-76 | 78-83 | 74-81 | 74-82 | 61-69 |
Pectoral-fin rays (ocular side) | 10-12 | 12-13 | 11 | 12-14 | 12 | 10-13 | 10-12 | 10-12 |
Pectoral-fin rays (blind side) | 7-10 | 7-9 | 9 | 8-11 | 10 | 9-12 | 7-10 | 8-9 |
Caudal-fin rays | iii,11,iii | iii,11,iii | iii,11,iii |
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iii, |
ii-iii,11-14,ii-iii | ii-iii, |
Lateral-line scales | 45-57 | 52-54 | 53 | 55-62 |
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56-67 | 58-66 | 49-56 |
Gill rakers | 0 + 5-8 (usually 0 + 6-7) | 0 + 4-5 | 0 + 8 | 0 + |
0 + 6-7 | 0 + |
0 +7-10 | 0 + 7-9 |
Condition of gill rakers | not serrated | not serrated | not serrated | not serrated | not serrated | not serrated | not serrated | not serrated |
Vertebrae | 10 + 30-33 | 10 + 30 | 10 + 30 | 10 + 31-32 | ? | 10 + |
10 + |
10 + 27-30 |
Head length (% of SL) | 23-28 | 23-25 | 23 | 23-28 | 27-28 | 22-28 | 22-28 | 26-31 |
Body depth (% of SL) | 40-46 | 40-44 |
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36-43 |
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40-45 | 36-45 | 45-62 |
Length of 2nd dorsal-fin ray in male (% of SL) (if filamentous) | 24-35 |
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-- | 27-50 | 17-26 |
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13-19 | 17-21 |
Length of 3rd dorsal-fin ray in male (% of SL) (if filamentous) | 24-37 |
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-- | 28-54 | 18-28 |
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16-23 | -- |
Interorbital | bony ridge, no scales | bony ridge, no scales | bony ridge, no scales | bony ridge, no scales | bony ridge, no scales |
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bony ridge, no scales | bony ridge, no scales in juveniles; |
Interorbital (% of upper orbit diameter) | 4-17 | 11-16 | 11 | 10-14 | 10-14 |
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4-11 | 13-25 |
Prevomer | small, weakly projecting into mouth cavity | small, weakly projecting into mouth cavity | small, weakly projecting into mouth cavity |
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small, weakly projecting into mouth cavity | small, weakly projecting into mouth cavity | small, weakly projecting into mouth cavity | small, weakly projecting into mouth cavity |
Distribution | eastern Atlantic from Morocco south to Namibia; Canary Islands | [Gulf of Suez?] eastern Mediterranean | Sumatra (Indonesia), China | central Red Sea and Persian Gulf east to Indonesia | Taiwan to Indonesia and northwestern Australia | Morocco to South Africa | Mediterranean and eastern Atlantic from Scotland to Strait of Gibraltar; Azores and Madeira | Mediterranean and Black seas, and eastern Atlantic from Ireland to Sierra Leone |
Depth | 34-175 | ca. 20-30 | ? | 18-141 | 100-224 | 70-200 | 15-350 | 5-300 |
Remark: This key has been updated on the basis of
1a. Dorsal-fin rays 110-118; anal-fin rays 86-94 ...
1b. Dorsal-fin rays less than 110; anal-fin rays less than 85 ... 2
2a. Gill rakers 0 + 4-5; dorsal and anal fins plain black ...
2b. Gill rakers 0 + 7-13; dorsal and anal fins pale, sometimes with dark spots or only anterior rays black ... 3
3a. Dorsal-fin rays 74-80; anal-fin rays 51-57 ...
3b. Dorsal-fin rays 81 or more; anal-fin rays 61 or more ... 4
4a. Dorsal-fin rays 81 to 93; anal-fin rays 61 to 74; at most one elongate dorsal-fin ray ... 5
4b. Dorsal-fin rays 95 to 106; anal-fin rays 74 to 82; two or more elongate dorsal-fin rays ... 6
5a. No elongate dorsal-fin rays; no dark markings on ocular-side pectoral fin; no series of dark spots present along base of caudal fin ...
5b. Second dorsal-fin ray of males elongate (more than 60% of head length) with dark-fringed, broad membrane (1st, 3rd and 4th dorsal-fin rays may also be elongate, but these are only about one-third as long as second ray); ocular-side pelvic fin often with dark spot or blotch on posterior rays; a series of dark spots often present along base of caudal fin ...
6a. Dorsal-fin rays 2 to 5 (-7) elongate (elongate rays nearly equal to head length in males); eyes separated by a bony ridge; lateral-line scales 52 to 66; 5 to 10 gill rakers on lower limb of first gill arch ... 7
6b. Anterior 3 or 4 dorsal-fin rays of males noticeably prolonged and of nearly equal length; eyes separated by a concave scaly space; lateral-line scales 56 to 67; 10 to 13 gill rakers on lower limb of first gill arch ...
7a. Lateral-line scales 58-66; dorsal-fin rays 96-106; anal-fin rays 74-82; gill rakers 0 + 8-10; ventral side of males pale in life; dorsal-fin filaments in male plain whitish ...
7b. Lateral-line scales 52-57; dorsal-fin rays 89-99; anal-fin rays 64-78; gill rakers 0 + 5-8 (usually 0 + 6-7); ventral side of males bright red in life; dorsal-fin filaments in male light grey, with black margins, distally yellowish ...
We would like to thank J. Maclaine (BMNH) for taking x-rays of syntypes of
The authors wish to thank all the participants in the BISSAU1219 and MEDITS surveys, as well as the crew of R/V