sm82n3-4824

Scorpaena decemradiata new species (Teleostei: Scorpaenidae) from the Gulf of Aqaba, northern Red Sea, a species distinct from Scorpaena porcus

Ronald Fricke 1, Daniel Golani 2, Brenda Appelbaum-Golani 3, Uwe Zajonz 4,5

1 Im Ramstal 76, 97922 Lauda-Königshofen, Germany.
(RF) (Corresponding author) E-mail: ronfricke@web.de. ORCID iD: http://orcid.org/0000-0003-1476-6990
2 National Natural History Collections and Department of Ecology, Evolution and Behavior, The Hebrew University of Jerusalem, 91904 Jerusalem, Israel.
(DG) E-mail: dani.golani@mail.huji.ac.il. ORCID iD: http://orcid.org/0000-0003-4575-3324
3 The Hebrew University of Jerusalem, 91905 Jerusalem, Israel.
(BA-G) E-mail: brendag@savion.huji.ac.il. ORCID iD: http://orcid.org/0000-0002-3237-6980
4 Senckenberg Biodiversität und Klima Forschungszentrum, Arbeitsgruppe Biogeografie, Senckenberganlage 25,
60325 Frankfurt am Main, Germany.
5 Senckenberg Forschungsinstitut und Naturmuseum, Sektion Ichthyologie, Senckenberganlage 25,
60325 Frankfurt am Main, Germany.
(UZ) E-mail: uzajonz@senckenberg.de. ORCID iD: https://orcid.org/0000-0001-6106-1411

Summary: The scorpionfish Scorpaena decemradiata n. sp. is described from off the coast of Israel in the Gulf of Aqaba, northern Red Sea. The new species is similar to S. porcus Linnaeus, 1758, but is characterized by 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. An updated checklist of the species of the genus Scorpaena Linnaeus, 1758 and a key to the species of the eastern Atlantic, Mediterranean Sea and Red Sea are presented.

Keywords: taxonomy; fishes; new species; Red Sea; distribution; checklist; key.

Scorpaena decemradiata nueva especie (Teleostei: Scorpaenidae) del golfo de Aqaba, mar Rojo del norte, una especie distinta de Scorpaena porcus

Resumen: En este trabajo se describe el escorpeniforme Scorpaena decemradiata n. sp. de la costa de Israel, golfo de Aqaba, norte del mar Rojo. La nueva especie es similar a S. porcus Linnaeus, 1758, pero se caracteriza por la presencia de XII espinas y 10 radios blandos (el último dividido en la base) en la aleta dorsal; 16 radios en la aleta pectoral, el radio ramificado más superior es el segundo; lacrimal con 2 espinas sobre el maxilar que apuntan casi en ángulo recto, la posterior apuntando ventralmente y ligeramente en dirección anterior; fosa occipital bien desarrollada; los poros más anteriores de la línea lateral mandibular son pequeños y separados; escamas ctenoides; de 59 a 62 filas de escamas en series longitudinales; escamas ausentes en el torax y la base de la aleta pectoral; y cirros desarrollados sobre toda la cabeza y el cuerpo, pero no en la mandíbula inferior. Se presenta una lista actualizada de las especies del género Scorpaena Linnaeus, 1758, y una clave para las especies del Atlántico este, mar Mediterráneo y mar Rojo.

Palabras clave: taxonomía; peces; nueva especie; mar Rojo; distribución; lista; clave.

Citation/Como citar este artículo: Fricke R., Golani D., Appelbaum-Golani B., Zajonz U. 2018. Scorpaena decemradiata new species (Teleostei: Scorpaenidae) from the Gulf of Aqaba, northern Red Sea, a species distinct from Scorpaena porcus. Sci. Mar. 82(3): 169-184. https://doi.org/10.3989/scimar.04824.17A

LSID: urn:lsid:zoobank.org:pub:473F6B7B-D02F-4FCB-9DD2-7B060DD3B61D

Editor: E. Macpherson.

Received: June 25, 2018. Accepted: September 13, 2018. Published: September 26, 2018.

Copyright: © 2018 CSIC. This is an open-access article distributed under the terms of the Creative Commons Attribution 4.0 International (CC BY 4.0) License.

Contents

Summary
Resumen
Introduction
Materials and methods
Taxonomy
Discussion
Acknowledgements
References

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

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Fig. 1. Scorpaena decemradiata n. sp., HUJ 2418, holotype, 123.1 mm SL, Red Sea, Gulf of Aqaba, Israel, Eilat, 1960. Lateral view; photograph of the colouration in preservative taken 56 years after collection (Photograph: D. Golani).

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.

Table 1. – Selected counts and body proportions of Scorpaena decemradiata new species; proportions expressed as percentage of standard length unless otherwise stated.

Range Red Sea HUJ 2148, holotype (123.1 mm SL) Red Sea HUJ 20671, paratype (1) (90.5 mm SL)
Counts:
Dorsal fin spines XII XII
Dorsal fin soft rays 10 10
Anal fin spines + soft rays III + 5 III + 5
Pectoral fin rays 16 16
Uppermost branched pectoral fin ray 2nd 2nd
Lowermost branched pectoral fin ray 7th 6th
Pored lateral-line scales 29 30
Scale rows in longitudinal series 62 59
Vertical scale rows 53 48
Circumpeduncular scales 42 40
Gill rakers (total) 17 16
Gill rakers on upper arch 5 4
Spines on suborbital ridge 3 2-3
Pectoral fin reaching to level of anus anus
Proportions:
Body depth 41 41
Head length 44 42
Horizontal eye diameter 11 12
Tip of snout to dorsal fin origin 35 36
Tip of snout to anal fin origin 69 71
Tip of snout to dorsal fin insertion 73 73
Tip of snout to anal fin insertion 75 70
Tip of snout to pectoral fin origin 38 38
Tip of snout to pelvic fin origin 41 38
Snout in orbit diameter 1.2 1.4
Orbit diameter in head length 4.0 3.5
Interorbital distance in orbit diameter 1.8 1.6
Dorsal fin origin to caudal fin base 75 75
Anal fin origin to caudal fin base 38 39
Pectoral fin origin to caudal fin base 68 66
Length of dorsal fin base 62 63
Length of anal fin base 16 18
Length of pectoral fin base 16 15
Length of pectoral fin 29 32
Length of pelvic fin 26 29
Length of pelvic fin spine 13 15
Dorsal fin insertion to caudal fin base 32 32
Anal fin insertion to caudal fin base 30 33
Least depth of caudal peduncle 12 12

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.

figure2

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Fig. 2. – Geographical distribution of the Scorpaena decemradiata n. sp. and S. porcus Linnaeus, 1758 in the eastern Atlantic, Mediterranean Sea and Red Sea. 1, S. decemradiata n. sp., type locality; 2, S. porcus; 3, S. porcus (probable lectotype locality of S. erythraea).

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).

Table 2. – Comparison Scorpaena decemradiata n. sp. and S. porcus; proportions expressed as percentage of standard length unless otherwise stated. Differences of S. porcus compared with S. decemradiata n. sp. are printed in bold face.

Scorpaena decemradiata n. sp. S. porcus
Dorsal fin spines XII XII
Dorsal fin rays 10 7-9
Anal fin spines + rays III, 5 III, 4-5
Pectoral fin rays 16 16-18
Uppermost branched pectoral fin ray 2nd usually 3rd
Lateral scale rows 59-62 56-60
Vertical scale rows 48-53 51-58
Circumpeduncular scales 40-42 32-40
Body scales ctenoid emarginate, without distinct ctenii
Gill rakers on upper arch 4-5 5-6
Body depth 41 34-40
Tip of snout to pectoral fin origin 38 39-45
Tip of snout to pelvic fin origin 38-41 40-48
Orbit diameter in head length 3.5-4.0 3.7-4.3
Length of pectoral fin base 15-16 12-15

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.).

figure3

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Fig. 3.Scorpaena porcus Linnaeus, 1758, HUJ 12258, 153 mm SL, Jaffa, Israel, Mediterranean Sea, 23 Apr. 1987. Lateral view; photograph of the fresh colouration taken a few hours after collection (Photograph: D. Darom).

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Fig. 4.Scorpaena porcus Linnaeus, 1758, MNHN 0000-6706, lectotype of Scorpaena erythraea Cuvier in Cuvier and Valenciennes, 1829, specimen 1, 154.7 mm SL, Egypt, Red Sea/probably erroneous, Mediterranean Sea, Alexandria, 1798-1799. Lateral view; photograph of the colouration in preservative taken ca. 218 years after preservation.

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.

Table 3. – Selected counts and body proportions of Mediterranean specimens of Scorpaena porcus Linnaeus, 1758; proportions expressed as percentage of standard length unless otherwise stated. Number of specimens in parentheses.

Range Eastern Mediterranean (?): MNHN 0000-6706 (2 syntypes of Scorpaena erythraea; larger specimen designated as lectotype, see above) Eastern Mediterranean: HUJ 4962 (1), 5315 (1), 12258 (1), 14139 (2), 14223 (2), 14555 (1), 14591 (1), 14573 (4), 14674 (1), 19126 (2), 19129 (3), 19394 (1), HUJ 19437 (1), SMNHTAU P2578 (1) (53.1-154.1mm SL; n=22) Western Mediterranean: BMNH 1938.11.15. 50 (1), BMNH 1960. 6.24.145-149 (5), BMNH 1963.5.14. 655 (1), HUJ 12060 (1), HUJ 17860 (1), HUJ 20823 (5), SMF 35951 (9) (74.5-171.7mm SL; n=23)
Counts:
Dorsal fin spines XII XII XII
Dorsal fin soft rays 9 (2) 8 (1), 9 (20) 7 (1), 9 (23)
Anal fin spines + soft rays III + 4 (1), III + 5 (1) III + 5 (20), III + 6 (1) III + 4 (1), 5 (23)
Pectoral fin rays 16 (3), 17 (1) 16 (12), 17 (20), 18 (1) 16 (23), 17 (18), 18 (5)
Uppermost branched pectoral fin ray 3rd (4th) 2nd (1), 3rd (32), 4th (1) 2nd (11), 3rd (32), 4th (3)
Lowermost branched pectoral fin ray 7th-8th 5th-8th 5th-7th
Pored lateral-line scales 18-31 28-31 28-30
Scale rows in longitudinal series 56-57 56-60 56-60
Vertical scale rows 52-53 51-58 51-58
Circumpeduncular scales 34-36 32-38 34-40
Gill rakers (total) 16 16-17 16-18
Gill rakers on upper arch 5 5- 6 5- 6
Spines on suborbital ridge ? 2- 3 2- 3
Pectoral fin reaching to level of anus – 2nd anal spine anus - 2nd or 3rd anal spine anus - 2nd anal spine
Proportions:
Body depth 36-37 36-44 34-40
Head length 42-44 42-47 42-47
Horizontal eye diameter 11 10-13 10-13
Tip of snout to dorsal fin origin 34 35-39 35-39
Tip of snout to anal fin origin 58-74 69-74 67-76
Tip of snout to dorsal fin insertion 71-73 72-79 67-72
Tip of snout to anal fin insertion 76-79 75-80 73-82
Tip of snout to pectoral fin origin 39 40-42 39-45
Tip of snout to pelvic fin origin 40-41 42-44 41-48
Snout in orbit diameter 1.1-1.3 1.2-1.7 1.0-1.4
Orbit diameter in head length 3.7-3.9 3.7-4.3 3.7-4.4
Interorbital distance in orbit diameter 1.5-1.8 1.0-1.8 1.3-2.1
Dorsal fin origin to caudal fin base 70-75 70-80 70-74
Anal fin origin to caudal fin base 34-35 35-39 33-37
Pectoral fin origin to caudal fin base 60-65 63-69 62-67
Length of dorsal fin base 60-64 56-64 57-63
Length of anal fin base 14-17 15-18 15-22
Length of pectoral fin base 12-13 14-15 14-17
Length of pectoral fin 29-31 29-34 28-30
Length of pelvic fin 26-31 25-30 23-27
Length of pelvic fin spine 12-17 14-17 11-16
Dorsal fin insertion to caudal fin base 30-32 30-33 30-33
Anal fin insertion to caudal fin base 29-30 27-32 26-33
Least depth of caudal peduncle 11 10-12 10-12

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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Motomura H., Senou H. 2008. A new species of the scorpionfish genus Scorpaena (Scorpaenidae) from Izu Peninsula, Pacific coast of Japan. J. Fish Biol. 72: 1761-1772.

Motomura H., Last P.R., Yearsley G.K. 2005. Scorpaena bulacephala, a new species of scorpionfish (Scorpaeniformes: Scorpaenidae) from the northern Tasman Sea. Zootaxa 1043: 17-32.

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Appendix 1. – Checklist of the species of the genus Scorpaena Linnaeus, 1758.

Species Original description Type locality Primary type(s) Junior synonym(s) Geographical distribution
S. afuerae Hildebrand, 1946 Hildebrand 1946Hildebrand S.F. 1946. A descriptive catalog of the shore fishes of Peru. Bull. U. S. Nat. Mus. 189: i-xi + 1-530.: 443, fig. 85 Lobos de Afuera Island, Peru Holotype: USNM 128130 E Pacific: Costa Rica to Peru and Cocos Island
S. agassizii Goode and Bean, 1896 Goode and Bean 1896Goode G.B., Bean T.H. 1896. Oceanic ichthyology, a treatise on the deep-sea and pelagic fishes of the world, based chiefly upon the collections made by the steamers Blake, Albatross, and Fish Hawk in the northwestern Atlantic, with an atlas containing 417 figures. Spec. Bull. U. S. Nat. Mus. 2: Text volume: i-xxxv + 1-26 + 1-553, Atlas volume: i-xxiii, 1-26, 123 pls.: 247, pl. 67 (fig. 243) Gulf of Mexico, 23°13’N 89°10’W Holotype: MCZ 27996 W Atlantic: North Carolina (USA) and Gulf of Mexico south to N Brazil
S. albifimbria Evermann and Marsh, 1900 Evermann and Marsh 1900Evermann B.W., Marsh M.C. 1900. The fishes of Porto Rico. Bull. U.S. Fish Comm. 20 (1): 49-350, pls. 1-49.: 275, fig. 82 off Culebra Island, southwest of Culebritas Lighthouse, Puerto Rico Holotype: USNM 49532 W Atlantic: Florida (USA) and Bahamas south to Netherlands Antilles
S. angolensis Norman, 1935 Norman 1935Norman J.R. 1935. Coast fishes. Part I. The South Atlantic. Discovery Rep. 12: 1-58.: (26) 28, fig. 10 Elephant Bay, Angola Holotype: BMNH 1910.2.24.1 E Atlantic: Mauritania to Angola, including Cape Verde Islands
S. annobonae Eschmeyer, 1969 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.: 75, Fig. 8c Annobón Island, Equatorial Guinea, Gulf of Guinea, 1°24’N, 5°38’E Holotype: CAS 14214 E Atlantic: Annobón Island
S. ascensionis Eschmeyer, 1971 Eschmeyer 1971Eschmeyer W.N. 1971. Two new Atlantic scorpionfishes. Proc. Calif. Acad. Sci. (Ser. 4) 37: 501-508.: 503, fig. 1 Ascension Island Holotype: BMNH 1935.5.2.33 SE Atlantic: at Ascension Island
S. azorica Eschmeyer, 1969 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.: 75, Fig. 8c : 80, fig. 8a Terceira, Azores Holotype: USNM 94463 NE Atlantic: Azores Islands
S. bergii Evermann and Marsh, 1900 Evermann and Marsh 1900Evermann B.W., Marsh M.C. 1900. The fishes of Porto Rico. Bull. U.S. Fish Comm. 20 (1): 49-350, pls. 1-49.: 276, fig. 83 Mayaguez, Puerto Rico Holotype: USNM 49533 W Atlantic: New York (USA) and Bermuda south to northern Brazil
S. brachyptera Eschmeyer, 1965 Eschmeyer 1965Eschmeyer W.N. 1965. Western Atlantic scorpionfishes of the genus Scorpaena, including four new species. Bull. Mar. Sci. 15: 84-164.: 111, figs 7a-b off Venezuela, 10°50’N, 66°55’W Holotype: USNM 198153 W Atlantic: Florida (USA) south to Panama and Venezuela
S. brasiliensis Cuvier in Cuvier and Valenciennes, 1829 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].: 305 Brazil Syntypes: MNHN 0000-6672 (1), 0000-6688 (1), 0000-7300 (1, dry); ZMB 747 (1) S. stearnsii Goode and Bean, 1882Goode G.B., Bean T.H. 1882. Descriptions of twenty-five new species of fish from the southern United States, and three new genera, Letharcus, Ioglossus, and Chriodorus. Proc. U. S. Nat. Mus. 5: 412-437.: 421.
S. colesi Nichols, 1914Nichols J.T. 1914. A new Scorpaena and a rare ray from North Carolina. Bull. Amer. Mus. Nat. Hist. 33: 537-538.: 537, fig. 1.
W Atlantic: Virginia (U.S.A.) and Gulf of Mexico south to northern Brazil
S. brevispina Motomura and Senou, 2008 Motomura and Senou 2008Motomura H., Senou H. 2008. A new species of the scorpionfish genus Scorpaena (Scorpaenidae) from Izu Peninsula, Pacific coast of Japan. J. Fish Biol. 72: 1761-1772.: 1762, figs. 1-5 off Futo, Ito City, Shizuoka, Japan, 34°52’N, 139°08’E Holotype: KPM-NI 16667 NW Pacific: Izu Peninsula, Japan
S. bulacephala Motomura, Last and Yearsley, 2005 Motomura et al. 2005Motomura H., Last P.R., Yearsley G.K. 2005. Scorpaena bulacephala, a new species of scorpionfish (Scorpaeniformes: Scorpaenidae) from the northern Tasman Sea. Zootaxa 1043: 17-32.: 19, figs. 1-3 south of Norfolk Island, Norfolk Ridge, 28°54’-55’S, 167°40’-41’E Holotype: CSIRO H 6009-05 SW Pacific: Norfolk Island and Lord Howe Island, New Caledonia, Vanuatu
S. calcarata Goode and Bean, 1882 Goode and Bean 1882Goode G.B., Bean T.H. 1882. Descriptions of twenty-five new species of fish from the southern United States, and three new genera, Letharcus, Ioglossus, and Chriodorus. Proc. U. S. Nat. Mus. 5: 412-437.: 422 Clearwater Harbor, Florida, USA Holotype: USNM 23556 S. russula atlantica Nichols and Breder, 1924Nichols J.T., Breder Jr. C.M. 1924. New Gulf races of a Pacific Scorpaena and Prionotus, with notes on other Gulf of Mexico fishes. Proc. Biol. Soc. Wash. 37: 21-24, pl. 7.: 21, pl. 7. W Atlantic: South Carolina (U.S.A.) and Gulf of Mexico south to northern Brazil
S. canariensis (Sauvage, 1878) Sauvage 1878Sauvage H.-E. 1878. Desription de poissons nouveaux ou imparfaitement connus de la collection du Muséum d’Histoire Naturelle. Famille des Scorpénidées, des Platycéphalidées et des Triglidées. Nouv. Arch. Mus. Hist. Nat., Paris (Sér. 2) 1: 109-158, pls. 1-2: 117, pl. 1 (figs. 1-2), as Sebastes canariensis Canary Islands Holotype: MNHN 0000-7031 S. patriarcha Berthelot [ex Valenciennes], 1840Berthelot S. 1840. De la pêche sur la côte occidentale d’Afrique, et des établissements les plus utiles aux progrès de cette industrie. Béthune, Paris, 302 + 1 pp., 1 map.:120 (not available, name only) E Atlantic: Canary Islands, Madeira and Azores
S. cardinalis Solander and Richardson in Richardson, 1842 Solander and Richardson in Richardson 1842bRichardson J. 1842b. Contributions to the ichthyology of Australia. Ann. Mag. Nat. Hist. (N.S.) 9: 207-218: 212 White Island, New Zealand, 37°30’S, 177°09’E Neotype: NMNZ P.044152 S. plebeia Solander and Richardson in Richardson, 1842bRichardson J. 1842b. Contributions to the ichthyology of Australia. Ann. Mag. Nat. Hist. (N.S.) 9: 207-218: 214.
S. cookii Günther, 1874Günther A.[C.L.G.] 1874. Andrew Garrett’s Fische der Südsee. Band I, Heft. III. J. Mus. Godeffroy, 2: 58-96, pls. 40-60.: 78, pl. 55
SW Pacific: New Zealand to Lord Howe, Norfolk and Kermadec islands
S. cocosensis Motomura, 2004 Motomura 2004Motomura H. 2004. New species of scorpionfish, Scorpaena cocosensis (Scorpaeniformes: Scorpaenidae) from the Cocos Islands, Costa Rica, eastern Pacific Ocean. Copeia 2004: 818-824.: 819, figs. 1-3 off Nuez Island, Cocos Island, 5°34’00”N, 86°59’20”W Holotype: CAS 219506 E Pacific: Galapagos Archipelago and Cocos Island
S. colorata (Gilbert, 1905) Gilbert 1905Gilbert C.H. 1905. II. The deep-sea fishes of the Hawaiian Islands. In: The aquatic resources of the Hawaiian Islands. Bull. U. S. Fish Comm. 23: 577-713, pls. 66-101.: 627, fig. 243 S of Molokai, Hawaiian Islands Holotype: USNM 51631 Central Pacific: Hawaiian Islands to Johnston Atoll
S. decemradiata n. sp. Present paper Red Sea, Gulf of Aqaba
S. dispar Longley and Hildebrand, 1940 Longley and Hildebrand 1940Longley W.H., Hildebrand S.F. 1940. New genera and species of fishes from Tortugas, Florida. Pap. Tortugas Lab., Carnegie Inst. Wash. 32: 223-285, pl. 1.: 246, fig. 12 S of Tortugas, Florida, USA Holotype: USNM 108867 S. similis Gunter, 1948Gunter G. 1948. Notes on fishes of the genus Scorpaena from the South Atlantic and Gulf coasts of the United States, with descriptions of two new species. Copeia 1948: 157-166.: 161, pl. 1. W Atlantic: South Carolina (USA) and Gulf of Mexico south to northern Brazil
S. elachys Eschmeyer, 1965 Eschmeyer 1965Eschmeyer W.N. 1965. Western Atlantic scorpionfishes of the genus Scorpaena, including four new species. Bull. Mar. Sci. 15: 84-164.: 114, fig. 7c N of Puerto Rico, 18°15’N, 67°33’W Holotype: USNM 198149 W Atlantic: Florida (U.S.A.) south to Panama
S. elongata Cadenat, 1943 Cadenat 1943Cadenat J. 1943. Les Scorpaenidae de l’Atlantique et de la Méditerranée. Première note. Le genre Scorpaena. Rev. Trav. Inst. Pêch. Marit. 13: 525-563.: 552, figs. 1(4), 9 Cap Blanc, Mauritania Syntypes: MHNLR P.798 (2) E Atlantic: Morocco to Namibia; Mediterranean Sea
S. fernandeziana Steindachner, 1875 Steindachner 1875Steindachner F. 1875. Ichthyologische Beiträge (II). I. Die Fische von Juan Fernandez in den Sammlungen des Wiener Museums. II. Über einige neue Fischarten von der Ost- und Westküste Süd-Amerikas. Sitzungsber. K. Akad. Wiss., Math.-Naturw. Cl. 71: 443-480, pl. 1.: 451 [9], pl. 1 (figs. 1, 1a) Juan Fernández Islands Holotype: whereabouts unknown SE Pacific: Juan Fernández and Desaventurados islands
S. gasta Motomura, Last andYearsley, 2006 Motomura et al. 2006Motomura H., Last P.R., Yearsley G.K. 2006. New species of shallow water scorpionfish (Scorpaenidae: Scorpaena) from the central coast of Western Australia. Copeia 2006: 360-369.: 361, figs. 1-3 Kalbarri, Western Australia, 27°30’S, 114°25’E Holotype: WAM P.27960-006 SE Indian Ocean: Western Australia
S. grandicornis Cuvier in Cuvier and Valenciennes, 1829 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].: 309, pl. 86 Martinique Island, West Indies; Puerto Rico; Havana, Cuba; Santo Domingo Syntypes: MNHN 0000-6681 (1), 0000-6689 (1), 0000-6900 (1); SMF 440 (1); ZMB 752 (1) W Atlantic: Bermuda and Florida (USA) south to southern Brazil; S Atlantic: St. Helena and Ascension
S. grandisquamis Ogilby, 1910 Ogilby 1910Ogilby J.D. 1910. On some new fishes from the Queensland coast. Endeavour Series, I. [Was to have appeared as Proc. Roy. Soc. Queensl., 23; withdrawn and privately published]. Brisbane, 85-139.: 107 North West Islet, Capricorn Group, Queensland, Australia Holotype: AMS E.1418 SW Pacific: Queensland (Australia)
S. grattanica Trunov, 2006 Trunov 2006Trunov I.A. 2006. Ichthyofauna of seamounts around the island of Ascension and St. Helena Island (Atlantic Ocean). Vopr. Ikht. 46: 471-477. [In Russian. English translation appeared in J. Ichth. 46:493-499.]: 472, fig. 2 Grattan Bank, near Ascension Island, 9°46’S, 12°48’W Holotype: ZIN 52148 S Atlantic: Grattan Bank near Ascension Island
S. guttata Girard, 1854 Girard 1854Girard C.F. 1854. Observations upon a collection of fishes made on the Pacific coast of the United States, by Lieut. W. P. Trowbridge, U. S. A., for the museum of the Smithsonian Institution. Proc. Acad. Nat. Sci. Philad. 7: 142-156.: 145 Monterey, California, USA Holotype: USNM 350 S. guadalupae Fowler, 1944Fowler H.W. 1944. Results of the fifth George Vanderbilt expedition (1941) (Bahamas, Caribbean Sea, Panama, Galápagos Archipelago and Mexican Pacific islands). The Fishes. Monogr. Acad. Nat. Sci. Philad. 6: 57-529, pls. 1-20.: 429, figs. 217-218.
S. microlepis Gunter, 1948Gunter G. 1948. Notes on fishes of the genus Scorpaena from the South Atlantic and Gulf coasts of the United States, with descriptions of two new species. Copeia 1948: 157-166.: 162, pl. 2.
E Pacific: central California (USA) to Gulf of California (Mexico)
S. hemilepidota Fowler, 1938 Fowler 1938Fowler H.W.1938. Descriptions of new fishes obtained by the United States Bureau of Fisheries steamer “Albatross”, chiefly in Philippine seas and adjacent waters. Proc. U.S. Nat. Mus. 85: 31-135.: 63, fig. 26 off Tubig Point, between Samar and Masbate, Philippines, 12°12’35”N, 124°02’48”E Holotype: USNM 98884 W Pacific: Philippines
S. histrio Jenyns, 1840 Jenyns 1840Jenyns L. 1840-1842. Fish. In: The zoology of the voyage of H. M. S. Beagle, under the command of Captain Fitzroy, R. N., during the years 1832 to 1836. Issued in 4 parts. Smith, Elder, and Co., London, i-xvi + 1-172, pls. 1-29. [Sherborn (1897) dates to Jan. 1840 (pp. 1-32), June 1840 (33-64), Apr. 1841 (65-96), Apr. 1842 (97-172).]: 35, pl. 8 San Cristóbal Island (Chatham Island), Galápagos Islands Syntypes: BMNH 1917.7.14.73, 1917.7.14.74 (1) S. fucata Valenciennes, 1846Valenciennes A. 1846. Ichthyologie. 1 table + pls. 1-10. In: Petit-Thouars A. du (ed.), Atlas de Zoologie. Voyage autour du monde sur la frégate “Vénus”, pendant les années 1836-1839. Gide, Paris.: Pl. 3 (fig. 2).
S. pannosa Cramer in Gilbert, 1897Gilbert C.H. 1897. Descriptions of twenty-two new species of fishes collected by the steamer Albatross, of the United States Fish Commission. Proc. U. S. Nat. Mus. 19: 437-457, pls. 49-55.: 446, pl. 52
E Pacific: Baja California (Mexico) S to Chile and some offshore islands, including the Galápagos Islands
S. inermis Cuvier in Cuvier and Valenciennes, 1829 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].: 311 Martinique Island, West Indies Holotype: MNHN 0000-0693 S. occipitalis Poey, 1860Poey F. 1858-1861. Memorias sobra la historia natural de la Isla de Cuba, acompañadas de sumarios Latinos y extractos en Francés. Tomo 2. Viuda de Barcina, La Habana, 1-96 (1858), 97-336 (1860), 337-442, (1861), pls. 1-19.: 171.
S. mercatoris Delsman, 1941Delsman N.C. 1941. Résultats scientifiques des croisières du Navire-école Belge “Mercator”, vol. III, No. 3. Pisces. Mém. Mus. Roy. Hist. Nat. Belg. (Sér. 2) 21: 47-82.: 74, fig. 11.
S. luckei Fowler 1941Fowler H.W. 1941. Notes on Florida fishes with descriptions of seven new species. Proc. Acad. Nat. Sci. Philad. 93: 81-106.: 87, figs. 1-2.
W Atlantic: Florida (USA) and Bahamas south to Dutch West Indies
S. isthmensis Meek and Hildebrand, 1928 Meek and Hildebrand 1928Meek S.E., Hildebrand S.F. 1928. The marine fishes of Panama. Part III. Field Mus. Nat. Hist., Publ., Zoöl. Ser. 15 (249): xxv-xxxi + 709-1045, pls. 72-102.: 842, pl. 80 Porto Bello, Panama, Caribbean Sea Holotype: USNM 81617 W Atlantic: Panama south to Rio de Janeiro (Brazil)
S. izensis Jordan andStarks, 1904 Jordan and Starks 1904Jordan D.S., Starks E.C. 1904. A review of the scorpaenoid fishes of Japan. Proc. U.S. Nat. Mus. 27: 91-175, pls. 1-2.: 134, fig. 10 Suruga Bay, Japan Holotype: USNM 50909 SE Indian Ocean and W Pacific: Western Australia north to S Japan
S. jacksoniensis Steindachner, 1866 Steindachner 1866Steindachner F. 1866. Über die Fische von Port Jackson in Australien. Anz. K. Akad. Wiss., Math.-Naturw. Cl. 3: 50-54.: 50 Port Jackson, New South Wales, Australia Holotype: NMW 75379 SW Pacific: Queensland to Victoria (Australia)
S. lacrimata Randall and Greenfield, 2004 Randall and Greenfield 2004Randall J.E., Greenfield D.W. 2004. Two new scorpionfishes (Scorpaenidae) from the South Pacific. Proc. Calif. Acad. Sci. 55: 384-394.: 391, fig. 3 Tahiti, Society Islands Holotype: BPBM 31706 S Pacific: Society Islands
S. laevis Troschel, 1866 Troschel 1866Troschel F.H. 1866. Ein Beitrag zur ichthyologischen Fauna der Inseln des Grünen Vorgebirges. Arch. Naturgesch. 32: 190-239, pl. 5.: 206 Cape Verde Islands Holotype: NMW 5632 S. senegalensis Steindachner, 1881aSteindachner F. 1881a. Beiträge zur Kenntniss der Meeresfische Afrikas’s (und Beschreibung einer neuen Sargus-Art von den Galapagos-Inseln). Anz. Akad. Wiss. Wien, Math.-Naturw. Cl. 18: 157-160.: 150; Steindachner 1881bSteindachner F. 1881b. Beiträge zur Kenntniss der Fische Afrika’s und Beschreibung einer neuen Sargus-Art von den Galapagos-Inseln. Denkschr. K. Akad. Wiss. Wien, Math.-Naturw. Cl. 44: 19-58, pls. 1-10.: 31, pl. 4 E Atlantic: Azores and Madeira S to Gulf of Guinea; Cape Verde Islands
S. loppei Cadenat, 1943 Cadenat 1943Cadenat J. 1943. Les Scorpaenidae de l’Atlantique et de la Méditerranée. Première note. Le genre Scorpaena. Rev. Trav. Inst. Pêch. Marit. 13: 525-563.: 541, figs. 1(3), 3 Gulf de Gascogne, France Syntypes: MHNLR P.797 (4); MNHN 1947-0002 to 0003 (2), 1887-0305 to 0306 (2), 1887-0307 to 0308 (2), 1887-0310 (1) Mediterranean Sea, E Atlantic: Bay of Biscay south to Western Sahara
S. maderensis Valenciennes in Cuvier and Valenciennes, 1833 Valenciennes in Cuvier and Valenciennes 1833Cuvier G., Valenciennes A. 1833. Histoire naturelle des poissons. Tome neuvième. Suite du livre neuvième. Des Scombéroïdes. F. G. Levrault, Paris, xxix + 3 + 512 pp., pls. 246-279. [Cuvier authored pp. 1-198, 330-359, 372-427; Valenciennes the balance].: 463 (as S. madurensis) Madeira Syntypes: MNHN 0000-6682 (4), 0000-6683 (4) S. rubellio Jordan and Gunn, 1898Jordan D.S., Gunn Jr. J.A. 1898. List of fishes collected at the Canary Islands by Mr. O. F. Cook, with descriptions of four new species. Proc. Acad. Nat. Sci. Philadelphia 50: 339-347.: 344. Mediterranean Sea, E Atlantic: Morocco to Senegal including Azores, Madeira, Canary and Cape Verde islands
S.melasma Eschmeyer, 1965 Eschmeyer 1965Eschmeyer W.N. 1965. Western Atlantic scorpionfishes of the genus Scorpaena, including four new species. Bull. Mar. Sci. 15: 84-164.: 109, fig. 6c off Brazil, 2°10’S, 41°33’W Holotype: USNM 198154 SW Atlantic: off Brazil
S. mellissii Günther, 1868 Günther 1868Günther A.[C.L.G.] 1868. Report on a collection of fishes made at St. Helena by J. C. Melliss, Esq. Proc. Zool. Soc. London 1868: 225-228, pls. 18-19.: 228, pl. 19 St. Helena, 15°58’S, 5°43’W Holotype: BMNH 1867.10.8.6 S Atlantic: Saint Helena
S. miostoma Günther, 1877 Günther 1877Günther A.[C.L.G.] 1877. Preliminary notes on new fishes collected in Japan during the expedition of H. M. S. ‘Challenger’. Ann. Mag. Nat. Hist. (Ser. 4) 20: 433-446.: 435 Yokohama, Japan Holotype: BMNH 1879.5.14.235 NW Pacific: Japan to China
S. mystes Jordan and Starks in Jordan, 1895 Jordan and Starks in Jordan 1895Jordan D.S. 1895. The fishes of Sinaloa. Proc. Calif. Acad. Sci. (Ser. 2) 5: 377-514, pls. 26-55.: 491, pl. 52 Mazatlán, Sinaloa, W Mexico Lectotype: CAS-SU 2919 (missing); established by Jordan and Evermann (1900:3288) Jordan D.S., Evermann B.W. 1900. The fishes of North and Middle America: a descriptive catalogue of the species of fish-like vertebrates found in the waters of North America, north of the Isthmus of Panama. Part IV. Bull. U. S. Nat. Mus. 47: i-ci + 3137-3313, pls. 1-392. Holoscorpaena didymogramma Fowler, 1944Fowler H.W. 1944. Results of the fifth George Vanderbilt expedition (1941) (Bahamas, Caribbean Sea, Panama, Galápagos Archipelago and Mexican Pacific islands). The Fishes. Monogr. Acad. Nat. Sci. Philad. 6: 57-529, pls. 1-20.: 277, figs. 214-215 E Pacific: southern California (USA.) south to Chile
S. neglecta Temminck and Schlegel, 1843 Temminck and Schlegel 1843Temminck C.J., Schlegel H. 1843. Pisces. In: Fauna Japonica, sive descriptio animalium quae in itinere per Japoniam suscepto annis 1823-30 collegit, notis observationibus et adumbrationibus illustravit P. F. de Siebold. Parts 2-4. A. Arnz et socios, Leiden, pp. 21-72.: 43 pl. 17 (fig. 4) Nagasaki, Japan Syntypes: RMNH D618-623 (6, stuffed) S. fimbriata Döderlein in Steindachner and Döderlein, 1884Steindachner F., Döderlein L. 1884. Beiträge zur Kenntniss der Fische Japan’s. (III.). Denkschr. K. Akad. Wiss. Wien, Math.-Naturw. Cl. 49: 171-212, pls. 1-7. [Also as separate, Wien, 1884.]: 195 [27] Indo-West Pacific: Andaman Islands east to Philippines and New Guinea, N to Japan, S to northern Australia
S. normani Cadenat, 1943 Cadenat 1943Cadenat J. 1943. Les Scorpaenidae de l’Atlantique et de la Méditerranée. Première note. Le genre Scorpaena. Rev. Trav. Inst. Pêch. Marit. 13: 525-563.: 539, fig. 2 Mauritania Syntypes: (17) MNHN B-2548 (2) E Atlantic: Mauritania south to Angola
S. notata Rafinesque, 1810 Rafinesque 1810Rafinesque C.S. 1810. Caratteri di alcuni nuovi generi e nuove specie di animali e piante della Sicilia, con varie osservazioni sopra i medisimi. Sanfilippo, Palermo. (Part 1 involves fishes, pp. [i-iv] 3-69; part 2 with slightly different title, pp. ia-iva + 71-105; pls. 1-20).: 33 Sicily, Italy No types known ?S. barbata Gronow in Gray, 1854Gray J.E. 1854. Catalogue of fish collected and described by Laurence Theodore Gronow, now in the British Museum. British Museum, London, vii + 196 pp.: 116.
S. ustulata Lowe, 1841Lowe R.T. 1841. Certain new species of Madeiran fishes ... Proc. Zool. Soc. London 8(89) (for 1840): 36-39.: 36.
S. maderensis (non Valenciennes, 1833): Lowe 1841Lowe R.T. 1841. Certain new species of Madeiran fishes ... Proc. Zool. Soc. London 8(89) (for 1840): 36-39.: 36.
S. teneriffea Jordan and Gunn, 1898Jordan D.S., Gunn Jr. J.A. 1898. List of fishes collected at the Canary Islands by Mr. O. F. Cook, with descriptions of four new species. Proc. Acad. Nat. Sci. Philadelphia 50: 339-347.: 345.
S. scrofa afimbria Slastenenko 1935Slastenenko E.P. 1935. The scorpionfishes Scorpaena of the Black Sea. Dokl. Akad. Nauk SSSR, Ser. A, 1: 74-80.: 75, fig.
Mediterranean Sea, Black Sea, E Atlantic: Bay of Biscay south to Senegal including Azores, Madeira and Canary Islands
S. onaria Jordan and Snyder, 1900 Jordan and Snyder 1900Jordan D.S., Snyder J.O. 1900. A list of fishes collected in Japan by Keinosuke Otaki, and by the United States steamer Albatross, with descriptions of fourteen new species. Proc. U.S. Nat. Mus. 23: 335-380, pls. 9-20.: 365, pl. 16 Misaki, Japan Holotype: USNM 49405 Indo-West Pacific: NWAustralia and Andaman Sea; Korea and Japan to Taiwan; E Australia and N New Zealand to New Caledonia
S. orgila Eschmeyer and Allen, 1971 Eschmeyer and Allen 1971Eschmeyer W.N., Allen G.R. 1971. Three new species of scorpionfishes (family Scorpaenidae) from Easter Island. Proc. Calif. Acad. Sci. (Ser. 4) 37: 515-527.: 517, figs. 1a, 2 Easter Island Holotype: CAS 24809 SE Pacific: Easter Island
S. papillosa (Schneider and Forster in Bloch and Schneider, 1801) Schneider and Forster in Bloch and Schneider 1801Bloch M.E., Schneider J.G. 1801. M.E. Blochii, Systema Ichthyologiae Iconibus cx Ilustratum. Post obitum auctoris opus inchoatum absolvit, correxit, interpolavit Jo. Gottlob Schneider, Saxo. Sander, Berolini, lx + 584 pp., 110 pls.: 196 New Zealand No types known – S. cottoides Forster in Bloch and Schneider, 1801Bloch M.E., Schneider J.G. 1801. M.E. Blochii, Systema Ichthyologiae Iconibus cx Ilustratum. Post obitum auctoris opus inchoatum absolvit, correxit, interpolavit Jo. Gottlob Schneider, Saxo. Sander, Berolini, lx + 584 pp., 110 pls.: 196.
S. miles Richardson, 1842aRichardson J. 1842a. Contributions to the ichthyology of Australia. Ann. Mag. Nat. Hist. (N.S.) 9: 15-31.: 18.
S. cruenta Solander in Richardson, 1842bRichardson J. 1842b. Contributions to the ichthyology of Australia. Ann. Mag. Nat. Hist. (N.S.) 9: 207-218.: 217.
S. ergastulorum Richardson, 1842bRichardson J. 1842b. Contributions to the ichthyology of Australia. Ann. Mag. Nat. Hist. (N.S.) 9: 207-218.: 217.
S. militaris Richardson, 1842cRichardson J. 1842c. Description of Australian fish. Trans. Zool. Soc. London 3: 69-131, pls. 4-6.: 90.
SW Pacific: SE Australia and New Zealand
S. pascuensis Eschmeyer and Allen, 1971 Eschmeyer and Allen 1971Eschmeyer W.N., Allen G.R. 1971. Three new species of scorpionfishes (family Scorpaenidae) from Easter Island. Proc. Calif. Acad. Sci. (Ser. 4) 37: 515-527.: 523, figs. 1b, 3 Easter Island Holotype: CAS 24812 SE Pacific: Easter Island
S. pele Eschmeyer and Randall, 1975 Eschmeyer and Randall 1975Eschmeyer W.N., Randall J.E. 1975. The scorpaenid fishes of the Hawaiian Islands, including new species and new records (Pisces: Scorpaenidae). Proc. Calif. Acad. Sci. (Ser. 4) 40: 265-333.: 320, figs. 24, 25a Oahu, Hawaiian Islands Holotype: USNM 214046 Central Pacific: Hawaiian Islands
S. pepo Motomura, Poss and Shao, 2007 Motomura et al. 2007Motomura H., Poss S.G., Shao K.-T. 2007. Scorpaena pepo, a new species of scorpionfish (Scorpaeniformes: Scorpaenidae) from northeastern Taiwan, with a review of S. onaria Jordan and Snyder. Zool. Stud. 46: 35-45.: 36, figs. 1, 2A NE Taiwan Holotype: ASIZP 65020 NW Pacific: Taiwan to Japan
S. petricola Eschmeyer, 1965 Eschmeyer 1965Eschmeyer W.N. 1965. Western Atlantic scorpionfishes of the genus Scorpaena, including four new species. Bull. Mar. Sci. 15: 84-164.: 107, fig. 6b off Brazil, 1°59’S, 42°05’W Holotype: USNM 198150 SW Atlantic: Brazil
S. plumieri Bloch, 1789 Bloch 1789Bloch M.E. 1789. Tvä utlaendska fiskar. Kongl. Vet. Akad. nya Handl. 10: 234-236.: 234, pl. 7 (fig. 1) Martinique No types known – S. bufo 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].: 306.
– S. scrofina Valenciennes in Cuvier and Valenciennes, 1833Cuvier G., Valenciennes A. 1833. Histoire naturelle des poissons. Tome neuvième. Suite du livre neuvième. Des Scombéroïdes. F. G. Levrault, Paris, xxix + 3 + 512 pp., pls. 246-279. [Cuvier authored pp. 1-198, 330-359, 372-427; Valenciennes the balance].: 465.
Apistes exul Gosse, 1851Gosse P.H. 1851. A naturalist’s sojourn in Jamaica. Longman, Brown, Green and Longmans, London, xxiv + 508 pp.: 207.
S. rascasio Poey, 1860Poey F. 1858-1861. Memorias sobra la historia natural de la Isla de Cuba, acompañadas de sumarios Latinos y extractos en Francés. Tomo 2. Viuda de Barcina, La Habana, 1-96 (1858), 97-336 (1860), 337-442, (1861), pls. 1-19.: 169.
S. albofasciata Metzelaar, 1919Metzelaar J. 1919. Report on the fishes, collected by Dr. J. Boeke in the Dutch West Indies 1904-1905, with comparative notes on marine fishes of tropical West Africa. F. J. Belanfante, ‘s-Gravenhage, 314 pp.: 145, fig. 43.
S. nuttingi Evermann and Seale, 1924Evermann B.W., Seale A. 1924. Report on the fishes collected by the Barbados-Antigua Expedition from the University of Iowa in 1918. Univ. Iowa Stud. Nat. Hist. 10: 25-40.: 39, pl. 1.
S. colonensis Meek and Hildebrand, 1928Meek S.E., Hildebrand S.F. 1928. The marine fishes of Panama. Part III. Field Mus. Nat. Hist., Publ., Zoöl. Ser. 15 (249): xxv-xxxi + 709-1045, pls. 72-102.: 844, pl. 81 (fig. 1).
W Atlantic including Ascension and Saint Helena
S. porcus Linnaeus, 1758 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.: 266 Mediterranean Sea; Atlantic Possible syntypes: NRM 22 (21), 23 (1), 123 (1) Cottus massiliensis Forsskål, 1775Forsskål P.S. 1775. Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in itinere orientali observavit. Post mortem auctoris edidit Carsten Niebuhr. Möller, Hauniae [Copenhagen], 20 + xxxiv + 164 pp., map.:x, 24; see Fricke (2008: 24)Fricke R. 2008. Authorship, availability and validity of fish names described by Peter (Pehr) Simon Forsskål and Johann Christian Fabricius in the ‘Descriptiones animalium’ by Carsten Niebuhr in 1775 (Pisces). Stuttg. Beit. Naturk. A, N. S. 1: 1-76.. – S. rascassa Lacepède, 1801Lacepède B.G.E. 1801. Histoire naturelle des poissons. Tome 3. Plassan, Paris, lxvi + 558 pp., pls. 1-34.: 275.
S. 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.
S. fasciata Costa, 1842-1853Costa O.G. 1842-53. Fauna del regno di Napoli, ossia enumerazione di tutti gli animali che abitano le diverse regioni di questo regno e le acque che le bagnano, etc. Pesci. v. 3 (pt 2). Napoli, 148 pp. [variously paginated], 69 pls.: Scorpaena p. 3, pl. 4.
S. scarpaena Nardo (ex Chiereghini), 1847Nardo G.D. 1847. Sinonimia moderna delle specie registrate nell’ opera intitolata: “Descrizione de’ crostacei, de’ testacei e de’ pesci che abitanno le lagune e golfo veneto rappresentati in figure à chiaro-scuro ed a colori.” Antonelli, Venezia, xi + 128 pp.: col. 121.
S. phorcus Gronow in Gray, 1854Gray J.E. 1854. Catalogue of fish collected and described by Laurence Theodore Gronow, now in the British Museum. British Museum, London, vii + 196 pp.: 117.
S. klausewitzi Frøiland in 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.
Mediterranean Sea, Black Sea, E Atlantic: British Isles to Morocco including Azores and Canary Islands
S. russula Jordan and Bollman, 1890 Jordan and Bollman 1890Jordan D.S., Bollman C.H. 1890. Descriptions of new species of fishes collected at the Galapagos Islands and along the coast of the United States of Colombia, 1887-’88. In: Scientific results of explorations by the U. S. Fish Commission steamer Albatross. Proc. U. S. Nat. Mus. 12: 149-183.: 165 off Pacific coast of Colombia Syntypes: BMNH 1900.9.29.189 (1); NMW 8864 (1); CAS-SU 384 (4); USNM 41138 (1), 41140 (1), 41146 (1), 41154 (1) 41160 (1), 41191 (1), 41208-09 (1, 1), 41366 (1), 41379 (1), 41487-88 (1, 1), 205651 (1) E Pacific: Gulf of California to northern Peru
S. scrofa Linnaeus, 1758 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.: 266 Mediterranean Sea No types known – Scorpaena tota rubens, cirris plurimis ad os: Artedi 1738aArtedi P. 1738a. Genera piscium. In quibus systema totum ichthyologiæ proponitus cum classibus, ordinibus, generum characteribus, specierum differentiis, observationibus plurimis. Redactis speciebus 242 ad genera 52. Ichthyologiæ pars 3. Conradus Wishoff, Lugduni Batavorum, Leiden, 84 + ii pp.: 47; 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.: 76.
– Scorpaena cavite cavernoso, cirris geminis in maxilla inferiore: Gronow 1754Gronow L.T. 1754. Museum Ichthyologicum, sistens piscium indigenorum et quorundam exoticorum, qui in Museo Laur. Theod. Gronovii adservantur, descriptiones, ordine systematico; accedunt nonullorum exoticorum piscium icones, aeri incisae. Tomus 1. Theodorus Haak, Leiden, viii + 70 pp., pls. 1-4.: 46 (no. 103).
?S. barbata Bonnaterre [ex Gronow], 1788Bonnaterre J.P. 1788. Ichthyologie. Tableau encyclopédique et methodique des trois règnes de la nature. Panckoucke, Paris, lvi + 215 pp., pls. A-B + 1-100.:70.
– ?S. gronovii Walbaum [ex Gronow], 1792Walbaum J.J. 1792. Petri Artedi sueci genera piscium. In quibus systema totum ichthyologiae proponitur cum classibus, ordinibus, generum characteribus, specierum differentiis, observationibus plurimis. Redactis speciebus 242 ad genera 52. Ichthyologiae pars III. Ant. Ferdin. Rose, Grypeswaldiae [Greifswald], [i-viii] + 723 pp., pls. 1-3.: 383.
S. lutea Risso, 1810Risso A. 1810. Ichthyologie de Nice, ou histoire naturelle des poissons du Département des Alpes Maritimes. F. Schoell, Paris, xxxvi + 388 pp., pls. 1-11.: 190.
S. istriensis Nardo [ex Chiereghini], 1847Nardo G.D. 1847. Sinonimia moderna delle specie registrate nell’ opera intitolata: “Descrizione de’ crostacei, de’ testacei e de’ pesci che abitanno le lagune e golfo veneto rappresentati in figure à chiaro-scuro ed a colori.” Antonelli, Venezia, xi + 128 pp.: col. 121.
S. natalensis Regan, 1906Regan C.T. 1906. Descriptions of new or little known fishes from the coast of Natal. Ann. Natal Gov. Mus. 1: 1-6, pls. 1-5.: 5, pl. 5.
?Scorpaenopsis marmoratus Bonde, 1923Bonde C. von 1923. Shallow-water fishes procured by the S.S. “Pickle.” Rep. Fish. Mar. Biol. Surv., Union of South Africa 3: 1-40, pls. 1-9.: 30, pl. 7.
Mediterranean Sea, E Atlantic: British Isles to Cape Verde Islands including Madeira; W Indian Ocean
S. sonorae Jenkins and Evermann, 1889 Jenkins and Evermann 1889Jenkins O.P., Evermann B.W. 1889. Description of eighteen new species of fishes from the Gulf of California. Proc. U. S. Nat. Mus. 11: 137-158.: 150 Guaymas, Sonora, W Mexico Holotype: USNM 39644 E Pacific: Mexico, Gulf of California and SW coast of Baja California
S. stephanica Cadenat, 1943 Cadenat 1943Cadenat J. 1943. Les Scorpaenidae de l’Atlantique et de la Méditerranée. Première note. Le genre Scorpaena. Rev. Trav. Inst. Pêch. Marit. 13: 525-563.: 550, figs. 1 (5), 8 Port-Etienne, Cap Blanc, Mauritania Syntypes: MNHN 1947-0004 (1), 1947-0005 (1) S. gaillardae Roux, 1954Roux C. 1954. Description de deux espèces nouvelles de poissons des côtes d’Afrique Équatoriale Française, Dentex polli et Scorpaena gaillardae. Bull. Mus. Nat. Hist. Nat. (Sér. 2) 26: 468-472.: 470. E Atlantic: Mauritania south to Angola
S. sumptuosa Castelnau, 1875 Castelnau 1875Castelnau F.L. 1875. Researches on the fishes of Australia. Philadelphia Cent. Exp. 1876, Intercol. Exhib. Essays 2: 1-52.: 17 Fremantle, Western Australia Syntypes: MNHN A-4409 (1, dry), B-2570 (1) SE Indian Ocean: Western Australia, Shark Bay south to Albany
S. thomsoni Günther, 1880 Günther 1880Günther A.[C.L.G.] 1880. Report on the shore fishes procured during the voyage of H. M. S. Challenger in the years 1873-1876. Report on the Scientific Res. Voy. H. M. S. Challenger 1873-76, Zool. 1: 1-82, pls. 1-32.: 24, pl. 12 Juan Fernández Islands Holotype: BMNH 1879.5.14.232 SE Pacific, Chile: Juan Fernández and Desaventuradas islands
S. tierrae Hildebrand, 1946 Hildebrand 1946Hildebrand S.F. 1946. A descriptive catalog of the shore fishes of Peru. Bull. U. S. Nat. Mus. 189: i-xi + 1-530.: 441, fig. 84 Lobos de Tierra Bay, Peru Holotype: USNM 128128 E Pacific: Peru and Chile
S. uncinata Buen, 1961 Buen 1961Buen F. de. 1961. Peces Chilenos. Familias Alepocephalidae, Muraenidae, Sciaenidae, Scorpaenidae, Liparidae y Bothidae. Montemar 1: 1-52.: 32, fig. 9 Isla de San Ambrosio, Chile Holotype: EBMC 10478 SE Pacific: Chile
S. wellingtoni Victor, 2013 Victor 2013Victor B.C. 2013. Scorpaena wellingtoni n. sp., a new scorpionfish from the Galápagos Islands (Scorpaeniformes: Scorpaenidae). J. Ocean Sci. Found. 8: 30-43.: 32, figs. 1-4 Galápagos Islands, Isla Isabela, Tagus Cove, 0.26°S, 91.37°W Holotype: SIO 13-2 E Pacific: Galápagos Islands (Ecuador)
S. aplodactylus Bleeker 18531 Bleeker 1853Bleeker P. 1853. Nieuwe bijdrage tot de kennis der ichthijologische fauna van Ceram. Nat. Tijds. Ned. Ind. 3: 689-714.: 698 Wahai, Seram, Indonesia Syntypes and/or Bleeker specimens: AMS B.8277 (1); RMNH 5861 S. bleekeri Day, 1878Day F. 1878. The fishes of India; being a natural history of the fishes known to inhabit the seas and fresh waters of India, Burma, and Ceylon. Part 4. Bernard Quaritch, London, xx + pp. 553-779, pls. 139-195: 747, pl. 36, fig. 2.
1 Questionable species


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