INTRODUCTIONTop

The scorpionfishes of the genus Scorpaena Linnaeus, 1758 are mostly distributed in warm temperate seas, and occasionally also in tropical seas, around the world. They inhabit benthic habitats, mostly dwelling on rocky reefs. The genus includes a total of 61 valid species (Fricke et al. 2018Fricke R. (ed.). 2018. References in the Catalog of fishes, electronic version (4 Sept. 2018). California Academy of Sciences, San Francisco. Accessed on 7 Sept. 2018. http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp). It is characterized within the family Scorpaenidae by the dorsal rays normally XII, 9 (7-10, 8 or 10 normal for some species), anal rays normally III, 5; pectoral rays 16-21, some rays branched, the branching usually compound in larger specimens; swimbladder absent; vertebrae 24; scales on body cycloid or ctenoid; occipital pit usually present, never flat or convex; palatine teeth present; ventral margin of lacrimal bone usually with numerous spines; posterior lacrimal spine absent or not hooked forward; no slit behind fourth gill arch; scales on pectoral fin base reduced or absent; lateral line normal, continuing onto or near base of caudal fin; pored lateral-line scales forming relatively complete tubes; and peritoneum pale (Eschmeyer 1969Eschmeyer W.N. 1969. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces: Scorpaenidae). Occ. Pap. Calif. Acad. Sci. 79: i-iv + 1-143.; Poss 1999Poss S.G. 1999. Scorpaenidae (scorpionfishes, also: lionfishes, rockfishes, stingfishes, stonefishes, and waspfishes). Pp. 2291-2352, iii-v. In: Carpenter K.E., Niem V.E. (eds.), Species identification guide for fisheries purposes. The living marine resources of the western central Pacific. Bony fishes part 2 (Mugilidae to Carangidae), volume 4. FAO, Rome, 2069–2790, i–viii.; modified).

The genus Scorpaena has been known since ancient times (Aristotle, 4th century BC; see Artedi 1738bArtedi P. 1738b. Synonymia nominum piscium fere omnium; in qua recensio fit nominum piscium, omnium facile authorum, qui undam de piscibus scripsere: uti Græcorum, Romanorum, Barbarorum, nec non omnium insequentium ichthyologorum, una cum nominibus inquilinis variarum nationum. Ichthyologiae pars 4. Conradus Wishoff, Lugduni Batavorum, Leiden, i + 118 + xxi pp.); in modern ichthyology it was first described by Linnaeus (1758: 266)Linnaeus C. 1758. Systema naturae, Ed. X. (Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.) Laurentius Salvius, Holmiae, ii + 824 pp., with S. porcus Linnaeus, 1758 and S. scrofa Linnaeus, 1758 as the only known species at the time. The species description of S. porcus by Linnaeus (1758)Linnaeus C. 1758. Systema naturae, Ed. X. (Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.) Laurentius Salvius, Holmiae, ii + 824 pp. was based on multiple sources from localities in the Mediterranean Sea (see Artedi 1738bArtedi P. 1738b. Synonymia nominum piscium fere omnium; in qua recensio fit nominum piscium, omnium facile authorum, qui undam de piscibus scripsere: uti Græcorum, Romanorum, Barbarorum, nec non omnium insequentium ichthyologorum, una cum nominibus inquilinis variarum nationum. Ichthyologiae pars 4. Conradus Wishoff, Lugduni Batavorum, Leiden, i + 118 + xxi pp., “Scorpaena pinnulis, ad oculos et nares”), and it was subsequently designated by Bleeker (1876: 3)Bleeker P. 1876. Mémoire sur les espèces insulindiennes de la famille des Scorpénoïdes. Verh. Kon. Akad. Wet. Amsterdam 16: 1-100, pls. 1-5. as the type species. The genus has been placed on the Official List of Generic Names in Zoology by Opinion 77 (Anonymous 1922Anonymous. 1922. Opinion 77. Thirty-five generic names in Protozoa, Coelenterata, Trematoda, Cestoda, Cirripedia, Tunicata, and Pisces placed in the Official List of Generic Names. In: Opinions rendered by the International Commission on Zoological Nomenclature. Opinions 68 to 77. Smiths. Misc. Coll. 33: 71-73.).

When examining specimens of Scorpaena from the Gulf of Aqaba, our attention was drawn to the identity of specimens previously misidentified as Scorpaena porcus. We found these specimens to be distinct from populations in the Mediterranean Sea, prompting our examination of other previous records of S. porcus, allegedly from the Red Sea. The analysis of these individuals demonstrated that the Gulf of Aqaba population represents a separate species, which is described in the present paper.

MATERIALS AND METHODSTop

Specimens were examined at the Australian Museum, Sydney (AMS), the Natural History Museum, London (BMNH), the Hebrew University, Jerusalem (HUJ), the Muséum National d’Histoire Naturelle, Paris (MNHN), Tel Aviv University (SMNHTAU) and the Staatliches Museum für Naturkunde, Stuttgart (SMNS).

Descriptive methods follow Eschmeyer (1969)Eschmeyer W.N. 1969. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces: Scorpaenidae). Occ. Pap. Calif. Acad. Sci. 79: i-iv + 1-143.. In the description, the data of the holotype are presented first, followed by those of the paratype in parentheses. The classification is based on Fricke et al. (2018)Fricke R., Eschmeyer W.N., Laan R. van der (eds.). 2018. Catalog of fishes, electronic version (4 Sept. 2018). California Academy of Sciences, San Francisco. Accessed on 7 Sept. 2018. http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp and references follow Fricke (2018)Fricke R. (ed.). 2018. References in the Catalog of fishes, electronic version (4 Sept. 2018). California Academy of Sciences, San Francisco. Accessed on 7 Sept. 2018. http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp. The museum abbreviations follow Fricke and Eschmeyer (2018)Fricke R., Eschmeyer W.N. 2018. A guide to fish collections in the Catalog of fishes. Online version, updated 4 Sept. 2018. California Academy of Sciences, San Francisco. Accessed on 7 Sept. 2018. http://researcharchive.calacademy.org/research/ichthyology/catalog/collections.asp.

Comparative material: Parascorpaena aurita: BMNH uncat. (1, 80.7 mm SL, previously identified as Scorpaena erythraea), Red Sea; BMNH 1871.4.13.26 (1, 88.7 mm SL, previously identified as Scorpaena erythraea), Massaua, Eritrea, Red Sea. Scorpaena porcus: BMNH 1929.8.7.25 (1), Sevastopol, Black Sea; BMNH 1935.4.12.2 (1), Dorset, UK; BMNH 1938.11.15.50 (1, 124.1 mm SL), Philippeville [Skikda], Algeria; BMNH 1960.6.24.145-149 (5, 99.1-131.3 mm SL), Catalunya, Spain; BMNH 1963.5.14.655 (1, 118.4 mm SL), Banyuls-sur-Mer, France; BMNH 2015.3.12.9 (1, 233.4 mm SL), Plymouth, UK; HUJ 4962 (1), Bat-Galim, Israel; HUJ 5315 (1), Michmoret, Israel; HUJ 12060 (1), Ligurian Sea, Italy; HUJ 12258 (1), Jaffo, Israel; HUJ 14139 (2), Famagusta, Cyprus; HUJ 14223 (2), Cyprus, HUJ 14555 (1), Famagusta, Cyprus; HUJ 14573 (4), Rhodes, Greece; HUJ 14591 (1), Rhodes, Greece; HUJ 14674 (1), Cape Kiti, Cyprus; HUJ 17860 (1), Malta; HUJ 19126 (2), Cape Dolex, Cyprus; HUJ 19129 (3), Famagusta, Cyprus; HUJ 19394 (1), İstanbul, Turkey; HUJ 19437 (1), Yanai Beach, Israel; HUJ 20283 (6), east of Mallorca, Balearic Islands, Spain; MNHN 0000-6706 (2 syntypes of Scorpaena erythraea Cuvier in Cuvier and Valenciennes, 1829, 107.6-154.7 mm SL), Red Sea/locality probably erroneous; the larger specimen is the lectotype as designated below; SMF 35951 (9, 104.1-171.7 mm SL), France, Marseille; SMNHTAU P.2578 (1), Red Sea?; SMNS 1003 (4), Nice, France; SMNS 1673 (1), Alicante, Spain; SMNS 9186 (1), Mallorca, Baleares, Spain; SMNS 9196 (1), Mallorca, Baleares, Spain; SMNS 9407 (1), Menorca, Baleares, Spain; SMNS 9450 (1), Osor, Cres, Croatia; SMNS 9610 (3), Varkiza, Greece, SMNS 9613 (1), Varkiza, Greece; SMNS 9879 (2), Varkiza, Greece; SMNS 10044 (3), Porto Santo Stefano, Toscana, Italy; SMNS 10055 (6), Porto Santo Stefano, Toscana, Italy; SMNS 11519 (1), Bodrum, Muğla, Turkey; SMNS 11531 (1), Bodrum, Muğla, Turkey; SMNS 11534 (1), Bodrum, Muğla, Turkey; SMNS 11583 (2), Greece, Varkiza; SMNS 16399 (1), Beirut, Lebanon; SMNS 16696 (1), Karataş, Adana, Turkey; SMNS 16697 (1), Karataş, Adana, Turkey; SMNS 19044 (1), Girne, Northern Cyprus; SMNS 19046 (11), Girne, Northern Cyprus; SMNS 19056 (12), Girne, Northern Cyprus; SMNS 19062 (6), Girne, Northern Cyprus; SMNS 19078 (3), Girne, Northern Cyprus; SMNS 19099 (23), Girne, Northern Cyprus; SMNS 19209 (1), Mallorca, Baleares, Spain; SMNS 20360 (1), Tabarka, Tunisia; SMNS 24887 (1), Paradise Bay, Malta; SMNS 24896 (4), Paradise Bay, Malta; SMNS 24904 (2), Paradise Bay, Malta; SMNS 25514 (4), Monaco; SMNS 25515 (2), Palermo, Sicily, Italy; SMNS 25517 (1), Venice, Italy; SMNS 25518 (4), Venice, Italy.

TAXONOMYTop

Scorpaena decemradiata n. sp.
(Fig. 1, Table 1)

Scorpaena porcus (non Linnaeus 1758Linnaeus C. 1758. Systema naturae, Ed. X. (Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.) Laurentius Salvius, Holmiae, ii + 824 pp.): Frøiland 1972Frøiland Ø. 1972. The scorpaenids of the Red Sea (Pisces: Scorpaenidae). A taxonomical and zoogeographical study. Mimeographed PhD dissertation, University of Bergen, Norway, vi + 160 pp.: 23 (Eilat, Israel, Gulf of Aqaba, Red Sea; based on HUJ 2418). Dor 1984Dor M. 1984. Checklist of the fishes of the Red Sea. CLOFRES. The Israel Academy of Sciences and Humanities, Jerusalem, xxii + 437 pp., 1 map.: 82 (part). Goren and Dor 1994Goren M., Dor M. 1994. An updated checklist of the fishes of the Red Sea. CLOFRES II. The Israel Academy of Sciences and Humanities, Jerusalem, xii + 120 pp., 2 maps.: 22 (part).

Holotype: HUJ 2418, 123.1 mm SL, Red Sea, Gulf of Aqaba, Israel Eilat, Y. Berens, Sept. 1960.

Paratype: HUJ 20671, 1 specimen, 95.0 mm SL, Red Sea, Gulf of Aqaba, Israel Eilat, Y. Berens, Sept. 1960.

Diagnosis. A species of Scorpaena with dorsal fin spines XII, soft dorsal fin rays 10 (the last divided at base); pectoral fin rays 16, uppermost branched pectoral fin ray is the second; lacrimal with 2 spines over maxilla that point at nearly right angle from each other, the posterior pointing ventrally and slightly anteriorly; occipital pit well developed; anteriormost mandibular lateral-line pores small, separated; scales ctenoid; 59-62 scale rows in longitudinal series; scales absent on chest and pectoral fin base; and cirri developed over entire head and body, but no cirri on lower jaw.

Description. Dorsal fin-ray formula XII, 10 (XII, 10). Anal fin-ray formula III, 5 (III, 5). Pectoral fin-ray formula, all elements, 16 (16), upper 2nd-7th (2nd-7th) branched. Gill rakers 5+12, total 17 (4+12, total 16) on first gill arch.

Selected body proportions and counts, included in Table 1, are part of the description.

Body scaled; scales ctenoid. Chest, pectoral fin base and head naked. Predorsal scales 5 (6). Preorbital bone usually with 2 (2) spinous points over maxillary forming about a right angle; posterior spine pointing forward. Occipital pit present, well developed. Suborbital ridge with 3 (2-3) spinous points; first below ridge which runs under eye, second at end of this ridge, and third just before supplemental preopercular spine. Upper posttemporal spine present. Interorbital ridges present, diverging at rear. Supraorbital tentacle at most half of orbit diameter. Few small dermal flaps associated with preorbital, preocular, parietal, nuchal and preopercular spines; other tentacles at anterior nostril, below suborbital ridge, on eye, opercle flap, some body scales, and some pored lateral-line scales. Pores at symphysis small, separate. Lateral line a shallow convex curve from its origin to caudal fin base, with 29 (30) pored scales. Scale rows in longitudinal series 62 (59), vertical scale rows 53 (48). Maximum observed standard length 123 mm.

Colour in alcohol. For pigmentation of body refer to Figure 1, which is part of the description.

Head and body reddish brown, back with five indistinct darker saddles that continue irregularly across the upper two-thirds of the body. Head reddish brown; eye dark grey. Pectoral fin base and belly white. Dorsal fin marbled with brown, without a black blotch. Pectoral fins pale, with series of dark brown spots in their upper two-thirds. Pelvic fins pale. Anal fin light brown, with a central and a distal posterior bar of dark brown spots. Caudal fin whitish, with a basal, a central and a distal vertical dark brown bar.

Etymology. Decem (Latin) means ten; radiata (Latin) means rayed. The name refers to the ten soft rays in the dorsal fin of the new species, which clearly distinguish it from the closely related species, S. porcus.

Distribution and habitat. Known only from the Gulf of Aqaba (Eilat, Israel) (Fig. 2). The depth of collection and the habitat are unknown.

Comparisons. Scorpaena decemradiata n. sp. is very similar to S. porcus from the Mediterranean Sea and eastern Atlantic; these two species are distinguished from other congeners by the following combination of characters: presence of a deep occipital pit, pectoral fin base naked, lateral-line pores immediately posterior to symphysis of lower jaw widely separate, and more than 56 scale rows in longitudinal series. The new species is distinguished from S. porcus (see Table 2) by 10 dorsal fin soft rays (versus 7-9 in S. porcus), scales on body ctenoid (versus emarginate), and uppermost branched pectoral fin ray is the second (versus usually third, rarely second to fourth).

An identification key to the species of Scorpaena in the eastern Atlantic, Mediterranean and Red Sea is presented below, in order to easily identify this species.

In the Red Sea, the new species might be confused with Parascorpaena aurita (Rüppell, 1829), which among other characters, however, only has 8-9 dorsal fin soft rays (versus 10 in Scorpaena decemradiata n. sp.), and larger scales, with only 35-44 scale rows in longitudinal series (versus 59-62 in Scorpaena decemradiata n. sp.).

Scorpaena porcus Linnaeus, 1758
(Figs 3-4, Table 3)

Scorpaena erythraea Cuvier in Cuvier and Valenciennes 1829Cuvier G., Valenciennes A. 1829. Histoire naturelle des poissons. Tome quatrième. Livre quatrième. Des acanthoptérygiens à joue cuirassée. F. G. Levrault, Paris, xxvi + 2 + 518 pp, pls. 72-99, 97 bis. [Cuvier authored volume.]: 316 (Egypt, Red Sea; locality probably in error). Günther 1860Günther A.[C.L.G.] 1860. Catalogue of the fishes in the British Museum. Catalogue of the acanthopterygian fishes in the collection of the British Museum. Volume 2. Squamipinnes, Cirrhitidae, Triglidae, Trachinidae, Sciaenidae, Polynemidae, Sphyraenidae, Trichiuridae, Scombridae, Carangidae, Xiphiidae. British Museum, London, xxi + 548 pp.: 116. Klunzinger 1870Klunzinger C.B. 1870. Synopsis der Fische des Rothen Meeres. I. Theil. Percoiden-Mugiloiden. Verh. K.-K. Zool.-Bot. Ges. Wien 20: 669-834.: 803. Klunzinger 1884Klunzinger C.B. 1884. Die Fische des Rothen Meeres. Eine kritische Revision mit Bestimmungstabellen. I. Teil. Acanthopteri veri Owen. E. Schweizerbart’sche Verlagshandlung, Stuttgart, ix + 133 + [13] pp., pls. 1-13.: 70. Fowler and Steinitz 1956Fowler H.W., Steinitz H. 1956. Fishes from Cyprus, Iran, Iraq, Israel and Oman. Bull. Res. Counc. Israel 5B: 260-292.: 281. Smith 1957Smith J.L.B. 1957. The fishes of the family Scorpaenidae in the western Indian Ocean. Part I. The sub-family Scorpaeninae. Ichth. Bull., Dept. Ichth., Rhodes Univ. 4: 49-72, pls. 1-4.: 51 (uncertain identity; locality in error).

Scorpaena porcus: Dor 1984Dor M. 1984. Checklist of the fishes of the Red Sea. CLOFRES. The Israel Academy of Sciences and Humanities, Jerusalem, xxii + 437 pp., 1 map.: 82 (part). Eschmeyer and Dempster 1990Eschmeyer W.N., Dempster L.J. 1990. Scorpaenidae. Pp. 665-679. In: Quéro J.C., Hureau J.C., Karrer C., et al. (eds), Check-list of the fishes of the eastern tropical Atlantic (CLOFETA). Vol. 2. UNESCO, Paris, pp. 665-679.: 674-675 (part: synonymy of S. erythraea, type locality allegedly incorrect). Goren and Dor 1994Goren M., Dor M. 1994. An updated checklist of the fishes of the Red Sea. CLOFRES II. The Israel Academy of Sciences and Humanities, Jerusalem, xii + 120 pp., 2 maps.: 22 (part). Golani and Bogorodsky 2010Golani D., Bogorodsky S.V. 2010. The fishes of the Red Sea – reappraisal and updated checklist. Zootaxa 2463: 1-135.: 65 (Red Sea record incorrect, based on misidentifications).

Scorpaena klausewitzi: Frøiland 1972Frøiland Ø. 1972. The scorpaenids of the Red Sea (Pisces: Scorpaenidae). A taxonomical and zoogeographical study. Mimeographed PhD dissertation, University of Bergen, Norway, vi + 160 pp.: 25 (Eilat, Israel, Gulf of Aqaba, Red Sea; based on SMNHTAU 2578; locality in error; nomen nudum, appeared in an unpublished doctoral thesis).

Scorpaenopsis erythraea: Frøiland 1972Frøiland Ø. 1972. The scorpaenids of the Red Sea (Pisces: Scorpaenidae). A taxonomical and zoogeographical study. Mimeographed PhD dissertation, University of Bergen, Norway, vi + 160 pp.: 72 (Red Sea, locality in error).

Additional synonymy is provided in the checklist below.

Diagnosis. A species of Scorpaena with dorsal fin spines XII, soft dorsal fin rays 7-9 (the last divided at base); pectoral fin rays 16-18, uppermost branched pectoral fin ray is usually the third (rarely second to fourth); lacrimal with usually 2 spines over maxilla that point at nearly right angle from each other, the posterior pointing ventrally and slightly anteriorly; occipital pit well developed; anteriormost mandibular lateral-line pores widely separated; scales small, emarginate, without distinct ctenii; 56-60 scale rows in longitudinal series; scales absent on chest and pectoral fin base; cirri well developed over entire head and body, but no cirri on lower jaw.

Distribution and habitat. Mediterranean Sea, Black Sea, eastern Atlantic: British Isles to Morocco including Azores and Canary Islands (Fig. 2). The species dwells benthic habitats, from shallow water to 800m depth, in the shallows usually on rocks covered with algae.

DISCUSSIONTop

Scorpaena erythraea was originally described by Cuvier in Cuvier and Valenciennes (1829: 316)Cuvier G., Valenciennes A. 1829. Histoire naturelle des poissons. Tome quatrième. Livre quatrième. Des acanthoptérygiens à joue cuirassée. F. G. Levrault, Paris, xxvi + 2 + 518 pp, pls. 72-99, 97 bis. [Cuvier authored volume]., based on three specimens collected by Étienne Geoffroy Saint-Hilaire in Egypt. Geoffroy Saint-Hilaire’s fish material was collected during the French expedition to Egypt in 1798-1799; in 1799, the material was transported to Alexandria, Egypt, and when Alexandria was conquered by British troops, 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: 88Bauchot M.-L., Daget J., Bauchot R. 1990. L’ichtyologie en France au début du XIXe siècle. L’Histoire naturelle des poissons de Cuvier et Valenciennes. Bull. Mus. Nat. Hist. Nat., A (Zool.), 12 Suppl.: 1-142.). The specimens were neither described nor illustrated in the works of Étienne and Isidore Geoffroy Saint-Hilaire (1802aGeoffroy St. Hilaire E. 1802a. Description d’un nouveau genre de poisson, de l’ordre des abdominaux. Bull. Soc. Phil. Paris 3: 97-98, pl. 5., 1802bGeoffroy St. Hilaire E. 1802b. Histoire naturelle et description anatomique d’un nouveau genre de poisson du Nil, nommé polyptère. Ann. Mus. Nat. Hist. Nat., Paris 1: 57-68, pl. 5., 1809Geoffroy Saint-Hilaire E. 1809. Poissons du Nil, de la mer Rouge et de la Méditerranée. In: Description de l’Egypte ou recueil des observations et des recherches qui ont été faites en Égypte pendant l’expedition de l’Armée français, publié par les ordres de sa Majesté-L’Empereur Napoléon le Grand, v. 1 (part 1). Imprimerie Impériale, Paris, 1-52 [141-338], Poissons Pls. 1-17., 1817Geoffroy Saint-Hilaire E. 1817. Poissons du Nil, de la mer Rouge et de la Méditerranée. In: Description de l’Egypte ou recueil des observations et des recherches qui ont été faites en Égypte pendant l’expedition de l’Armée français, publié par les ordres de sa Majesté-L’Empereur Napoléon le Grand. Tome 1, (pt. 1). Imprimerie Impériale, Paris, pls. 18-27., 1827aGeoffroy St. Hilaire I. 1827a. Suite de l’histoire des poissons du Nil. In: Description de l’Egypte ou recueil des observations et des recherches qui ont été faites en Égypte pendant l’expedition de l’Armée français, publié par les ordres de sa Majesté-L’Empereur Napoléon le Grand. Tome 1, (pt. 1). Imprimerie Impériale, Paris, pp. 265-310., 1827bGeoffroy Saint-Hilaire I. 1827b. Poissons du Nil, de la mer Rouge et de la Méditerranée. In: Description de l’Egypte ou recueil des observations et des recherches qui ont été faites en Égypte pendant l’expedition de l’Armée français, publié par les ordres de sa Majesté-L’Empereur Napoléon le Grand. Imprimerie Impériale, Paris, pp. 311-343.).

The species was subsequently reported by several authors based on Cuvier in Cuvier and Valenciennes (1829)Cuvier G., Valenciennes A. 1829. Histoire naturelle des poissons. Tome quatrième. Livre quatrième. Des acanthoptérygiens à joue cuirassée. F. G. Levrault, Paris, xxvi + 2 + 518 pp, pls. 72-99, 97 bis. [Cuvier authored volume].; only Günther (1860: 116)Günther A.[C.L.G.] 1860. Catalogue of the fishes in the British Museum. Catalogue of the acanthopterygian fishes in the collection of the British Museum. Volume 2. Squamipinnes, Cirrhitidae, Triglidae, Trachinidae, Sciaenidae, Polynemidae, Sphyraenidae, Trichiuridae, Scombridae, Carangidae, Xiphiidae. British Museum, London, xxi + 548 pp. described an additional specimen from the Red Sea (BMNH 1871.4.13.26). This specimen, as well as another specimen identified as Scorpaena erythraea (non Cuvier in Cuvier and Valenciennes 1829Cuvier G., Valenciennes A. 1829. Histoire naturelle des poissons. Tome quatrième. Livre quatrième. Des acanthoptérygiens à joue cuirassée. F. G. Levrault, Paris, xxvi + 2 + 518 pp, pls. 72-99, 97 bis. [Cuvier authored volume].), turned out to be based on misidentified Parascorpaena aurita (Rüppell, 1829). Smith (1957: 51)Smith J.L.B. 1957. The fishes of the family Scorpaenidae in the western Indian Ocean. Part I. The sub-family Scorpaeninae. Ichth. Bull., Dept. Ichth., Rhodes Univ. 4: 49-72, pls. 1-4. erroneously reported Scorpaena erythraea from Mauritius.

Eschmeyer and Dempster (1990: 674-675)Eschmeyer W.N., Dempster L.J. 1990. Scorpaenidae. In: Quéro J.C., Hureau J.C., Karrer C., et al. (eds), Check-list of the fishes of the eastern tropical Atlantic (CLOFETA). Vol. 2. UNESCO, Paris, pp. 665-679. were the first authors to discuss the identity of S. erythraea; they noticed that the species was probably identical with S. porcus Linnaeus, 1758, and supposed that the type locality of S. erythraea was incorrect and the species was rather collected on the Mediterranean Sea shore of Egypt. This suspicion was followed by subsequent authors. Golani and Bogorodsky (2010: 65)Golani D., Bogorodsky S.V. 2010. The fishes of the Red Sea – reappraisal and updated checklist. Zootaxa 2463: 1-135. treated the Red Sea record of S. porcus as incorrect, and suggested that it was based on misidentifications. A confusion of the localities of material collected by Geoffroy Saint-Hilaire is highly likely, as the material was apparently not labelled originally, but subsequently after it arrived at Paris. There were just three major collecting localities of that expedition, the Nile, the Mediterranean Sea at Alexandria, and the Red Sea coast of Egypt. Assigning the locality was straightforward for freshwater fishes, and not a problem for material with manuscript notes by Geoffroy Saint-Hilaire, which could be easily identified, but for the other material which was not mentioned by Geoffroy Saint-Hilaire, the locality had to be guessed. The present study confirms the identity of the syntypes of Scorpaena erythraea Cuvier in Cuvier and Valenciennes, 1829 as conspecific with Scorpaena porcus Linnaeus, 1758, and the highly probable confusion of the type locality. All the characters examined agree well with eastern Mediterranean populations of S. porcus (see Table 3). The syntypes of S. erythraea (MNHN 0000-6706) were most probably collected near Alexandria, Egypt. The larger specimen (154.7 mm SL; Fig. 4, Table 2) of MNHN 0000-6706 is hereby designated as the lectotype of Scorpaena erythraea Cuvier in Cuvier and Valenciennes, 1829.

While we thus conclude that Scorpaena erythraea is a junior synonym of S. porcus and based on Mediterranean material, another specimen allegedly originating from the Gulf of Aqaba, Red Sea, SMNHTAU 2578, remains dubious. It was named Scorpaena klausewitzi in the unpublished doctoral dissertation by Frøiland (1972: 25)Frøiland Ø. 1972. The scorpaenids of the Red Sea (Pisces: Scorpaenidae). A taxonomical and zoogeographical study. Mimeographed PhD dissertation, University of Bergen, Norway, vi + 160 pp., and then again referred to as Scorpaenopsis erythraea (see Frøiland 1972: 72Frøiland Ø. 1972. The scorpaenids of the Red Sea (Pisces: Scorpaenidae). A taxonomical and zoogeographical study. Mimeographed PhD dissertation, University of Bergen, Norway, vi + 160 pp.). The former is a manuscript name that was supposed to be published but never was as the manuscript was first submitted to and then withdrawn from the journal Senckenbergiana biologica. However, the specimen also clearly belongs to S. porcus, and the locality is most probably in error, i.e. the specimen probably originated from the Mediterranean coast of Israel.

Frøiland (1972: 23)Frøiland Ø. 1972. The scorpaenids of the Red Sea (Pisces: Scorpaenidae). A taxonomical and zoogeographical study. Mimeographed PhD dissertation, University of Bergen, Norway, vi + 160 pp. also recorded Scorpaena porcus (non Linnaeus, 1758) from Eilat, Israel, based on two specimens numbered HUJ 2418; these are the only specimens that were really collected in the Red Sea and belong to the Scorpaena porcus complex, but were found to represent S. decemradiata n. sp., which is described in the present paper.

Summarizing these results, Scorpaena porcus was previously thought to be a case of anti-Lessepsian migration, but is obviously restricted to the eastern Atlantic and Mediterranean, and previous records from the Red Sea were based on material with incorrect localities or misidentifications. The Red Sea (Gulf of Aqaba) is inhabited by the closely related species S. decemradiata n. sp. This species has not yet been recorded from Jordan (Khalaf and Zajonz 2007Khalaf M., Zajonz U. 2007. Fourteen additional fish species recorded from below 150 m depth in the Gulf of Aqaba, including Liopropoma lunulatum (Pisces: Serranidae), new record for the Red Sea. Fauna Arab. 23: 421-433.).

An updated checklist of the species of the genus Scorpaena is presented here (Appendix 1). It now includes a total of 62 valid species. Most species are known from the eastern Atlantic, including the Mediterranean (18 species, which is 29.0% of the total species of the genus; 15 species or 24.2% are endemic to the region), followed by the eastern Pacific (14 species, 22.6%, all endemic) and the western Atlantic (14 species, 22.6%, with 12 species endemic, 19.4%), the western Pacific (12 species, 19.4%; 8 endemic, 12.9%), the eastern Indian Ocean (5 species, 8.1%; 2 endemic, 3.2%), the central Pacific (3 species, 4.8%; all endemic), the Red Sea (1 endemic species, 1.6%), and the western Indian Ocean (1 species, 1.6%; none endemic). From the Red Sea, S. decemradiata n. sp. is the only known species of this genus; it is probably endemic to the northern Red Sea, because it is not present in the German deep-sea expeditions to the central Red Sea, MESEDA I–III and MINDIK (Türkay 1996Türkay M. 1996. Composition of the deep Red Sea macroand megabenthic invertebrate fauna. Zoogeographic and ecological implications. In: Uiblein F., Ott J., Stachowitsch M. (eds), Deep-sea and extreme shallow-water habitats: affinities and adaptations. Biosyst. Ecol. Ser. 11: 43-59.; fish identifications and an unpublished faunal account by Uwe Zajonz).

As only two specimens of Scorpaena decemradiata n. sp. are known, little can be said about intraspecific variation. The intraspecific variation between specimens of S. porcus in the central Mediterranean Sea and the Black Sea was examined by Manilo and Peskov (2016)Manilo L.G., Peskov V.N. 2016. Comparative morphometric analysis of the small-scaled scorpionfish, Scorpaena porcus (Scorpaenidae, Scorpaeniformes), from the southern coast of the Crimea and eastern part of the Adriatic Sea. Vestn. Zool. 50: 533-538., who found some significant differences for some length proportions between specimens in the two regions. Boissin et al. (2016)Boissin E., Micu D., Janczyszyn-Le Goff M., et al. 2016. Contemporary genetic structure and postglacial demographic history of the black scorpionfish, Scorpaena porcus, in the Mediterranean and the Black Seas. Molec. Ecol. 25: 2195-2209. examined the population genetics of this species, and also found a weak genetic differentiation between populations in the Black Sea and the Mediterranean Sea. In the present study, western and eastern Mediterranean populations of S. porcus are compared (see Table 3), with no significant differences in fin-ray counts and scales, but again some differences in length proportions. We could not determine whether there are clines or rather a strict separation between the populations, but doubt that these differences are of taxonomic significance.

The restricted range of Scorpaena decemradiata n. sp. may be a relict distribution; the Scorpaena porcus complex that prefers a warm temperate climate may have had a wider distribution range during glacial periods, possibly all around the African continent, like S. scrofa. In this scenario, the S. porcus complex retreated to the north on both sides of the continent when the sea temperatures became warmer, but only on the western side was there room to spread out, while on the eastern side it was limited to the northernmost extent of the Red Sea in the Gulf of Aqaba. The subsequent continental barrier then facilitated speciation in this group, resulting in two different species on the western and eastern sides of the continent.

Key to eastern Atlantic, Mediterranean and Red Sea species of the genus Scorpaena

Remark. This key is based on Poss (2016)Poss S.G. 2016. Scorpaenidae. Scorpionfishes. Pp. 2250-2287. In: Carpenter K.E., Angelis N. de (eds.), The living marine resources of the Eastern Central Atlantic. Volume 3. Bony fishes part 1 (Elopiformes to Scorpaeniformes). FAO Species Identification Guide for Fishery Purposes. FAO, Rome, i-xiv, 1511-2342., but updated and expanded to cover the Red Sea species.

1	Pit or depression in occiput shallow or absent	2
–	A deep pit or depression in occiput (somewhat intermediate in Scorpaena elongata)	5
2	Chest (area anterior to pelvic fin) naked, without scales; membranes between dorsal spines 2 to 4 deeply incised more than half length of spine and nearly to base of fin in some specimens	S. normani
–	Chest with scales (sometimes deeply embedded and difficult to see); membranes between dorsal spines 2 to 4 incised about half length of spines or less	3
3	Suborbital ridge smooth, without spines; a shallow occipital pit; pectoral fin rays 20	S. ascensionis
–	Suborbital ridge with 1 or 2 small spines; no occipital pit; pectoral fin rays 15 to 18	4
4	Posterior lacrimal (preorbital) spine points anteriorly; second preopercular spine from above small, smaller than third and fourth below; 66 to 69 scales in longitudinal row above lateral line; no white specks in axil (inner surface) of pectoral fin	S. canariensis
–	Posterior lacrimal (preorbital) spine points posteroventrally; second preopercular spine from above small, large, equal to or larger than third and fourth below; 52 to 56 scales in longitudinal row above lateral line; white specks in axil of pectoral fin	S. maderensis
5	Base of pectoral fin and chest scaled; scales on flank cycloid	6
–	Base of pectoral fin naked, without scales (or with a few small deeply embedded scales); scales on flank ctenoid (or emarginate in S. porcus)	8
6	Fewer than 50 scales in lateral row behind supracleithral spine to base of caudal fin	7
–	More than 60 scales in lateral row behind supracleithral spine to base of caudal fin	S. mellissii
7	Medial surface of pectoral fin and pectoral axil with large brown spots on a relatively pallid background	S. laevis
–	Medial surface of pectoral fin and pectoral axil black, with large white spot	S. plumieri
8	Lateral-line pores immediately posterior to symphysis of lower jaw fused into a single median pore that is usually readily visible	9
–	Lateral-line pores immediately posterior to symphysis of lower jaw widely separate, although at times minute (S. decemradiata n. sp., S. scrofa) and difficult to locate	12
9	Maxilla with a ridge that runs along its length	S. loppei
–	Maxilla without ridge running along its length	10
10	A distinct, large spot on spinous dorsal fin between spines 6 and 9 (although sometimes more restricted); pectoral fin, soft part of dorsal fin and anal fin without numerous small, but distinct spots; spots when present on caudal fin confined primarily to fin rays	11
–	No distinct, large spot on spinous dorsal fin between spines 6 and 9; pectoral fin, soft part of dorsal fin, and anal fin with numerous small, but distinct spots; spots on caudal fin confined primarily to fin membranes	S. azorica
11	Dorsal soft rays usually 9 (last double); posteriormost lacrimal (preorbital) spine points ventrally or slightly to rear (not present or less distinct in juveniles)	S. angolensis
–	Dorsal soft rays 10 (last double); posterior lacrimal spines strongly curved to rear (may be less distinct or absent in juveniles)	S. annobonae
12	More than 56 scale rows in longitudinal series (counted from immediately behind supracleithral spine to base of caudal fin)	13
–	Fewer than 51 scale rows in longitudinal series (counted from immediately behind supracleithral spine to base of caudal fin)	14
13	Dorsal fin soft rays 7-9; scales on body emarginate, without distinct ctenii (small spines at posterior margin of scale); uppermost branched pectoral fin ray is usually the third (occasionally the second to fourth)	S. porcus
–	Dorsal fin soft rays 10; scales on body ctenoid; uppermost branched pectoral fin ray is the second	S. decemradiata n. sp.
14	Numerous cutaneous flaps and cirri on ventral side of head	S. scrofa
–	Ventral surface of head without flaps or cirri	15
15	Spinous part of dorsal fin without a black spot; pectoral fin rays usually 19 (sometimes 18)	S. elongata
–	Spinous part of dorsal fin with a distinct black spot; pectoral fin rays usually 17 or 18, rarely 19	16
16	More than 20 rows of scales anterior to anus at ventral midline; orbit diameter smaller than snout (ratio of snout/orbit 0.8 to 1.2), except in small specimens	S. stephanica
–	Fewer than 20 rows of scales anterior to anus at ventral midline; orbit diameter slightly larger than snout (ratio of snout/orbit 0.9 or less)	S. notata

ACKNOWLEDGEMENTSTop

We are grateful to James Maclaine and Oliver A. Crimmen (BMNH), Patrice Pruvost (MNHN, Paris), and M. Goren and N. Stern (SMNHTAU, Tel Aviv) who gave access to materials in their care, to D. Darom (HUJ, Jerusalem) for taking a photograph of Scorpaena porcus, and to N. Gluzman, Captain of the trawler F/V Bilu, for allowing us to examine the catch on his vessel. The research contributions of U. Zajonz to this article were supported by the Deutsche Forschungsgemeinschaft (German Research Council, grant KR 1758/1-1).

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