COASTAL FISHES OF THE WESTERN INDIAN OCEAN: A Longstanding Vision Realized

South African Institute for Aquatic Biodiversity Somerset Street, Private Bag 1015, Grahamstown, Eastern Province, South Africa, www.saiab.ac.za email: [email protected] Tel: +27 (0) 46 603 5800 Fax: +27 (0) 46 622 2403 COASTAL FISHES OF THE WESTERN INDIAN OCEAN:

Volume 1 Introductory chapters which include the oceanography of the WIO, the origins of coral reefs, A Longstanding Vision Realized an account of the people who laid the foundations of our knowledge of the WIO’s fishes, and more. The remainder of the volume covers the agnaths and condrichthyans of the area. Origins Volume 2 This volume includes a brief overview of the evolution of bony fishes, their anatomy and a key to the orders. Then follows accounts of families of the Orders Elopiformes to Mugiliformes. In 1949, not long after the publication of The Sea Fishes of Southern Africa, JLB Smith Volume 3 The remainder of the non-perciforms teleosts, from the Atheriniformes to the Scorpaeniformes. was offered the directorship of the Natal Museum in Pietermaritzburg. He turned the Volume 4 A key to the families of Perciformes, and the families Latidae to Chaetodontidae offer down, as he wanted to focus on a book on the fishes of the Western Indian Ocean, Volume 5 The key to the families, and the families Oplegnathidae to Draconettidae. which he said would take him a year or two. The vision was never fulfilled. Volume 6 The key to the families, and the families Gobiidae to Molidae, and the Latimeriidae. Each taxon has its appropriate keys and, for several of the families, a glossary of terms used in the account In 1986, the final successor to The Sea Fishes of Who put the book together? has been included. The volumes are well illustrated, with black-and-white illustration in keys and for species, Southern Africa, Margaret Smith and Phil Heemstra’s where appropriate, and colour plates of the species. The inside back and front covers of each volume will Smiths’ Sea Fishes, was published. Margaret died in Coastal Fishes of the Western Indian Ocean, with main have various maps of the Western Indian Ocean. 1987. Some 10 years later, Phil resurrected JLB’s idea editors Philip Heemstra, Elaine Heemstra, David of a book on the fishes of the Western Indian Ocean. Ebert, Wouter Holleman and John Randall, is to be Format and cost And now, 20 years after the re-birth of JLB Smith’s published early in 2018. It is the culmination of the The book will measure about 220mm wide x 280mm deep. Consideration is being given to both hard-cover idea, this is about to become a reality. work of more than 100 authors, photographers and illustrators, of editors, proofreaders, reviewers and soft-cover binding. The cost for a hard-cover will be in the region of R3500 per set ($270, €240) The Western Indian Ocean (WIO) is home to one and others, over a period of more than 20 years. and soft-cover R2000 ($150, €130). of the richest biotas of marine fishes on the planet, Amongst the major contributors are Phil Heemstra, comprising some 3600 species, or about 20% of the Dave Ebert, Dave Smith, Bruce Collette, Stuart Poss, Expression of interest world’s marine fish fauna. Although the WIO region Jack Randall, Gerry Allen, Helen Larson, Danny If you are interested in this publication, please 1) access the reply form on our website is the site for most of the earliest scientifically valid Hensley and Kunio Amaoka, Eric Anderson and http://www.saiab.ac.za/coastal-fishes-of-the-western-indian-ocean.htm and complete the online form, descriptions of Indo-Pacific fishes, the extent of its Keiichi Matsuura. OR 2) complete the form below, scan and mail it to [email protected], OR 3) post it to: fish fauna is still poorly known. Despite considerable effort by ichthyologists over the past two centuries, Coverage and contents WIO Fishes – Expression of Interest, SAIAB, Private Bag 1015, Grahamstown, 6140, South Africa many new species of fishes are described each year: This will help assess the number of copies to have printed. of the 329 new marine species described between The resulting publication, far more substantial Plate 20 Plate 202002 and 2012, 140 were from the WIO. than JLB Smith could have imagined, arranged in Your name: Family MyLIOBAtIDAE, Familyeaglerays; MyLIOBAtIDAE Family RhInOptERIDAE,, eaglerays; Family Cownose RhInOptERIDAE, Rays Cownosesix volumes, Rays covers species that occur in waters Institution:

Plate 20 generally shallower than 200 m. In this work, the Family MyLIOBAtIDAE, eaglerays; Family RhInOptERIDAE, Cownose Rays Western Indian Ocean includes the Red Sea and Address: Persian Gulf and extends to Cape Point, South Africa, and Kanyakumari, the southernmost tip of India. 1 1 2 Some authors2 have included fishes from the Gulf of Mannar and wider Sri Lanka. Email Address:

1 3 3 2 Binding preference: Hard cover Soft cover Nr. of sets Purpose: Institution Library Personal

3 4 4 5a 5a 6 6

5b 5b

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1 Aetobatis narinari, spotted eagleray, 1461 Aetobatis cm DW (Snarinari Mozambique)., spotted eagleray,2 Aetomylaeus 146 cm nichofi DW (S i, Mozambique).banded eagleray,5b 2 54Aetomylaeus cm DW, 92 nichofi cm TL ,i (Persianbanded eagleray,Gulf). 54 cm DW, 92 cm TL (Persian Gulf). 3 Aetomylaeus vespertilio, ornate eagleray3 Aetomylaeus (Maldives). vespertilio4 Myliobatis, ornate aquila eagleray, bullray, (Maldives). 128 cm DW 4 (Mozambique).Myliobatis aquila 5, Pteromylaeusbullray, 128 cm bovinus DW (Mozambique)., duckbill ray, 5 Pteromylaeus bovinus, duckbill ray, (a) 86 cm DW female; (b) 22 cm DW unborn(a) 86 pupcm DW (both female; South (b) Africa; 22 cm from DW Heemstra unborn pup & Heemstra (both South 2004). Africa; 6 Rhinoptera from Heemstra javanica & Heemstra, fl apnose 2004). ray, 128 cm 6 Rhinoptera javanica, fl apnose ray, 128 cm DW (N Mozambique). DW (N Mozambique).

1 Aetobatis narinari, spottedFamily eagleray, MOBULIDAE 146 cm DW (S Mozambique)., Mantas,Family 2devilrays Aetomylaeus MOBULIDAE nichofi i, banded, Mantas, eagleray, devilrays 54 cm DW, 92 cm TL (Persian Gulf). 3 Aetomylaeus vespertilio, ornate eagleray (Maldives). 4 Myliobatis aquila, bullray, 128 cm DW (Mozambique). 5 Pteromylaeus bovinus, duckbill ray, (a) 86 cm DW female; (b) 22 cm DW unborn pup (both South Africa; from Heemstra & Heemstra 2004). 6 Rhinoptera javanica, fl apnose ray, 128 cm DW (N Mozambique). Family MOBULIDAE, Mantas, devilrays

2 2 1 1

3 3 2 1

3

4 4 5 5

1 Manta birostris, giant manta, ~4 m DW1 Manta immature birostris male, giantdorsal manta, view (South ~4 m DWAfrica) immature and ventral male viewdorsal (Mozambique). view (South Africa) and ventral view (Mozambique). 2 Mobula eredoogootenkee, pygmy devilray,2 Mobula female eredoogootenkee (Madagascar). , pygmy3 Mobula devilray, kuhlii, female shortfi (Madagascar).n devilray, 110 3cm Mobula DW malekuhlii (Mozambique)., shortfi n devilray, 110 cm DW male (Mozambique). 4 Mobula thurstoni, bentfi n devilray, 1904 Mobula cm DW thurstoni (Oman)., 5bentfi Mobula4 n devilray, japanica 190, Japanese cm DW devilray (Oman). (Pakistan). 5 Mobula japanica, Japanese devilray (Pakistan). 5

1 Manta birostris, giant manta, ~4 m DW immature male dorsal view (South Africa) and ventral view (Mozambique). 296 2 Mobula eredoogootenkee296 , pygmy devilray, female (Madagascar). 3 Mobula kuhlii, shortfi n devilray, 110 cm DW male (Mozambique). 4 Mobula thurstoni, bentfi n devilray, 190 cm DW (Oman). 5 Mobula japanica, Japanese devilray (Pakistan).

296 (Bony Fishes) Order Beryciformes Family Monocentridae Osteichthyes Monocentridae Pineapple fishes Class Family by Phillip C� Heemstra

Body oval but somewhat compressed, with enlarged, bony, plate- Pineapple fishes like scales , bearing a sharp, carinate spine and fused to form a solid, bony armour. Two dorsal fins, first with 4–7 stout, ridged spines without interconnecting membranes; second dorsal fin by Phillip C� Heemstra with 11–12 soft (segmented) rays connected by a transparent membrane; anal fin similar to soft dorsal fin; caudal fin forked, his is a poorly understood group that currently is the lobes short, with rounded tips, branched rays 17; pelvic fins with a large spine and 3–4 tiny soft rays; the pelvic fin spines Monocentris japonica: 11 cm SL considered to include seven families: Berycidae, (O. Alvheim, IMR); ventral view Body oval but somewhat compressed, with enlarged, bony,can be locked plate- erect and and fit into a groove along sides of belly Holocentridae, Anoplogasteridae, Trachichthyidae, of lower jaw with light organs near Anomalopidae,T Diretmidae, and Monocentridae. They are when depressed. Snout short, bluntly rounded; light organ on symphysis shown by arrows (both 1a probably an artificial assemblage of unrelatedlike taxa. scales , bearing a sharp, carinate spine and fused lowerto form jaw. Solitary, a demersal. Mozambique). Most species live in deep marine waters and avoid bright Pineapple fishes live in 3–400 m. An Indo-Pacific family light, although may come closer to thesolid, surface at bonynight. The armour. Two dorsal fins, first with 4–7 stout,with 2 genera ridged and 4 species; 1 species in WIO. The monotypic Holocentridae, which have their greatestspines diversity onwithout coral interconnecting membranes; second Cleidopusdorsal gloriamaris fin is endemic to Australia. It has the jaws reefs, are largely or entirely nocturnal. The have relatively large 2a strongly curved, narrow suborbital space, and the light organ Family Anomalopidae eyes. During the day they tend to remain withhidden in crevaces,11–12 in soft (segmented) rays connected by a transparentis on the side of the lower jaw (hidden when mouth is closed). case or under ledges. Flashlight fishes Diagnosis: fin spines usually well developedmembrane; (rudimentary anal fin similar to soft dorsal fin; caudal fin Monocentrisforked, Bloch & Schneider, 1801 by John E� McCosker or absent in anoplogasterids, and in the dorsal and anal fins of Genus diretmids); pelvic fins with a strong spinethe and lobes6–12 soft short,rays with rounded tips, branched rays 17; pelvicJaws straight; fins light organ containing luminescent bacteria at A prominent luminous organ located beneath each eye. Body (only 3 or 4 rudimentary rays in Monocentridae): caudal fin front of lower jaw under chin. Three species, 1 in WIO. deep or somewhat fusiform, laterally compressed. Dorsal fin 3a Monocentris japonica: 11 cm SL with true (unpaired) small procurrent spineswith at base a of largecaudal spine and 3–4 tiny soft rays; the pelvic fin spines continuous or divided into separate spinous and soft-rayed fins. fin and 18 or 19 principal caudal-fin rays. Six families that can (O. Alvheim, IMR); ventralMouth viewoblique; snout blunt. Scales minute, strongly ctenoid. be recognised as coastal fishes occur in the canWIO. be locked erect and and fit into a groove along sidesMonocentris of belly japonica Teeth small, in bands on jaws and palatines; none on vomer. Pineapplefish of lower jaw with light organsGill-rakers wellnear developed. Light from the subocular luminous organ is continuously Key to Families when depressed. Snout short,4a. bluntly rounded; light organ on Gasterosteus japonicus Houttuynsymphysis 1782: 329, Pl. 2 (Nagasaki, shown Japan). by arrowsproduced (both by luminous bacteria and can be occluded using 1a. Pelvic fins with 1 huge spine (which can be locked erect) Monocentris japonicus Smith SFSA*; Smith 1986*; Kotlyar 1986*; rotational or lid-like mechanisms to block the light. Light is used and 3 or 4 rudimentary rays; scales enlarged, lower jaw. Solitary, demersal. Randal 1995*; Kotlyar 1996*; MichelMozambique). 1996*; Fricke 1999. with sharp ridges and spines, forming a bony armour to see by, to communicate, to confuse predators and to attract Heemstra & Heemstra 2004* resembling a pineapple � � � � � � � � � � � � � � � � � � � � MONOCENTRIDAEPineapple fishes live in 3–400 m. An Indo-Pacific family prey. Adults of most species live in deep water (below ~100 m) 1b. Pelvic fins with 1 spine and 6B12 soft rays; body not encased during the day, and ascend the reef face to feed during moonless in bony armour � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 2 with 2 genera4a. and 4 species; 1 species in WIO. The Dmonotypic IV–VII / 9–12; A 10–11; P 13–15; LL 12–17; GR 5–7 / 11–14. nights. (See McCosker, 1977, for a review of their biology.) 2a. Prominent luminous organ immediately BD 1.3–1.9, HL 1.8–2.4 in SL. Vertebrae 11+15. Jaws straight Rarely seen because of their nocturnal secretive behaviour, below eye � � � � � � � � � � � � � � � � � � � � � � � � � � � Cleidopus� � ANOMALOPIDAE gloriamaris is endemic to Australia. It haswith the villiform jaws teeth on jaws and palatines, none on vomer. Head, flashlight fishes occur in all tropical oceans.Anomalops Six genera katoptron with, 7 2b. No light organ beneath eye � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 3 body and fins yellow, the scales outlined in black; head naked. species, 1 in WIO, others likely (such as PLATE xx Anomalopidaewidely distributed from Indonesia to the south central Pacific) 3a. Jaws with large, fang-like teeth; dorsal-fin base strongly curved, narrow suborbital space, and the light organ Family A nocturnal species, found in caves and under ledges on coral to be found. Recently reviewed by Baldwin et al. (2007). 2 to 4 times as long as anal-fin base � � � � � � � � � � � ANOPLOGASTERIDAE Order Beryciformes 3b. Jaws with minute teeth; dorsal and anal-fin bases subequalis �on � � � � � the� � � 4 side of the lower jaw5a. (hidden when mouth is andclosed). rocky reefs, also on sand near wrecks and reefs. Juveniles in shallow water, adults have been trawled in deeper water (157 m). Key to Genera 4a. Dorsal fin with 10–13 strong spines � � � � � � � � � � � � � �HOLOCENTRIDAE Occasionally thrown ashoreFlashlight in rough seas. Light organs onfishes lower D single, continuous; mid-ventral scutes discontinuous; luminous organ 1a. 4b. Dorsal fin with 3–8 spines � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 5 jaw are used to locate prey (benthic crustaceans). Attains 17 cm. longer than eye, with a black eyelid-like flap that can be raised Photoblepharon to cover the organ � � � � � � � � � � � � � � � � � � � � � � � � � 5a. Anal fin with 4 spines and 12–30 soft rays, each maxilla with Indo-West Pacific: Red bySea toJohn South EAfrica� McCosker (south to Mossel Bloch & Schneider, 1801 1b. D divided into separate spinous and soft-rayed fins; 2 supramaxillae � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � Bay), India, Mauritius Oman, Saya de Malha Bank, Sri Lanka, BERYCIDAE Monocentris mid-ventral scutes continuous; light organ shorter 5b. Anal fin with 2 or 3 spines, 8–12 rays; Genus � � � � � � � Anomalops Kner, 1868 PLATE xx 5a. Somalia, Yemen to Indonesia, Australia, Philippines, Taiwan, than eye, without a black flap to cover it one supramaxilla � � � � � � � � � � � � � � � � � � � � � � � � museum specimens. Presently reported from Japan, Vietnam TRACHICHTHYIDAE China, Japan and New Zealand. Jaws straight; light organ containing luminescent bacteria at A prominent luminous organ located beneath each eye. Body and the northwestern Indian Ocean around Socotra and the front of lower jaw under chin. Three species, 1 in WIO. deep or somewhat fusiform, laterally compressed. Dorsal fin Monocentridae 61 Error Seamounts. Depth range: 200-900 m. (Bony Fishes) 59 Family Order Osteichthyes 5b. continuous or divided into separate spinous and soft-rayed fins. Mouth oblique; snout blunt. Scales minute, strongly ctenoid. 60 Order Beryciformes 2017/07/14 11:36 AM 2017/07/14 11:36 AM Monocentris japonica Teeth small, in bands on jaws and palatines; none on vomer. 1609 WIO Minion + Myriad2013.indd 61 1609 WIO Minion + Myriad2013.indd 59 2017/07/14 11:36 AM 1609 WIO Minion + Myriad2013.indd 60 Gill-rakers well developed. Pineapplefish 2017/07/14 11:36 AM 2a Light from the subocular luminous organ is continuously 1609 WIO Minion + Myriad2013.indd 58 Gasterosteus japonicus Houttuyn 1782: 329, Pl. 2 (Nagasaki, Japan). produced by luminous bacteria and can be occluded using Monocentris japonicus Smith SFSA*; Smith 1986*; Kotlyar 1986*; 1 rotational or lid-like mechanisms to block the light. Light is used Randal 1995*; Kotlyar 1996*; Michel 1996*; Fricke 1999. Beryx splendens: 24 cm (South Africa; from SSF). Family Anoplogasteridae PLATE xx to see by, to communicate, to confuse predators and to attract Heemstra & Heemstra 2004* PLATE xx Fangtooths prey. Adults Orderof most Beryciformes species live in deep water (below ~100 m) by M� Eric Andeson museum specimens. Presently reported from Japan, Vietnam during the day, and ascend the reef face to feed during moonless and the northwestern Indian Ocean around Socotra and the Gill, 1862 Body oval, thick anteriorly, compressed posteriorly with a Error Seamounts. Depth range: 200-900 m. Genus Centroberyx slender peduncle; head deeper than long. Mouth huge, oblique, D IV–VII / 9–12; A 10–11;2a P 13–15; LL 12–17;2b GR 5–7 / 11–14. nights. (See McCosker, 1977, for a review of their biology.) upper jaw protrusile, Juveniles with small depressible teeth; 1 Rarely seen because of their nocturnal secretive behaviour, Centroberyx druzhinini: ~14 cm SL (Mauritius; O. Alvheim, IMR). adult with long well-separated fangs on jaws; the longest pair at BD 1.3–1.9,1 HL 1.8–2.4 in SL. Vertebrae 11+15. Jaws straight Dorsal-fin base longer than anal-fin base. No ctenii on scales; lower jaw symphysis received into sockets of orbital cavity when flashlight fishes occur in all tropical oceans. Six genera with, 7 mouth is closed; tongue fills oral cavity when mouth is closed; with villiform teeth on jaws and palatines, none on vomer. Head, 2 Anomalops katoptron posterior margin with minute spinules. NoCentroberyx head spines. spinosus Dorsal- palatines with or without teeth. maxilla extending well past the eye; eye diameter subequal to greatest depth of maxilla. Gill- body and fins yellow, the scales2b outlined in black; head naked. species, 1 in WIO, others likely (such as Beryx splendens: 24 cm (South Africa; from SSF). Short alfinsino fin spines 5–7; anal soft rays 12–16; pelvic rays 7; LL 36–62 rakers long and slender in juveniles, but short, with 1–3 spinules widely distributed from Indonesia to the south central Pacific) Trachichthodes spinosus Gilchrist, 1903: 204, pl. 13 (type locality: off Kei in adults. Branchiostegal rays 8 or nine. Dorsal fin single, set 3 (total). Anterior Gill, 1862 edge of nasal bone withRiver spinules. mouth [Cape Morgan], Five South Africa);species, Barnard, 1925a: 363*; midway between head and tail, its base more than three times Genus Centroberyx 3 to be found. Recently reviewed by Baldwin et al. (2007). Gilchrist, 1903Fowler, 1935: and374; Smith, 1949:Austroberyx 152*. that of anal fin; caudal fin emarginate, with 17 branched rays; A nocturnal species, found in caves and under ledges on coral Dorsal-fin2 in base WIO. longer than anal-finTrachichthodes base. No ctenii on scales; Centroberyx spinosus: Busakhin, 1982: 14*; Heemstra, 1986: 410*; Smale et pectoral fins paddle-shaped, shorter than head; pelvic fins posterior margin with minute spinules. No head spines. Dorsal- al., 1995: 78. origin below rear end of pectoral-fin base; no fin spines except and rocky reefs, also on sand near wrecks and reefs.1 Juveniles in fin spines 5–7; anal soft rays 12–16; pelvic rays 7; LL 36–62 for procurrent caudal-fin rays at base of fin. Scales with a spiny 4 2 Beryx mollis: 21 cm SL (off Somalia; from Kotlyar 1993). (total).McCulloch, Anterior edge of nasal 1911 bone with are spinules. synonyms Five species, of Centroberyx. shallow water, adults have been trawled in deeper water (157 m). Gilchrist, 1903 and Austroberyx D V–VI, 14–15; A IV, 14–16; P 13; V I, 7; LL continuous, not saucer-like plate fused to a short pedicel connecting to a thin Key to Genera 2 in WIO. Trachichthodes extending onto caudal-fin rays, 40–47; GR 10–11/ 20–22. Depth plate embedded in the skin. Lateral line an open groove crossed 4 Beryx mollis: 21 cm SL (off Somalia; from Kotlyar 1993). McCulloch, 1911 are synonyms of Centroberyx. Occasionally thrown ashore in rough seas. Light organs on lower D single, continuous; mid-ventral scutes discontinuous; luminous organ 47–53% SL; head 35–40; length dorsal-fin base 38–43; length by well-separated bridges of scaly skin. Vertebrae without 1a. anal-fin base 32–34; predorsal 38–44; preanal 58–64; caudal- lateral projections; ribs not attached to centra; neural spines 5 io species io species jaw are used to locate prey (benthic crustaceans). Attains 17 cm. longer than eye, with a black eyelid-like flap that can be raised Photoblepharon Key toKey W to W fin 28–33; snout 15–20% HL; eye 31–47. Posterior margin of of abdominal vertebrae depressed. Adults bathypelagic in 700 � � � � � � � � � � � � � � � � � � � � � � � � � Beryx splendens 1a. LL 40–47; transverse scale rows between LL and anal-fin origin 15; total GR to 5000 m; juveniles epipelagic and mesopelagic. One genus, to cover the organ scales with denticles of varying lengths, no ctenii. Cheek scales Indo-West Pacific: Red Sea to South Africa (south to Mossel Beryx splendens 30–34 � � � � � � � � � � � � � � � � � � � �ansverse � � � � � � � � � � � � scale � � � C. spinosus rows between LL and anal-fin origin 15; total GR Anoplogaster Gill with 2 species, one in WIO. Slender alfonsino 1a. ansverseLL 40–47; scale rows between tr LL and anal-fin origin 18; total GR 5. Fulcral scale in pelvic axil well developed. No head spines. to separate spinous and soft-rayed fins; ) 1b. LL 53–62; tr C. druzhinini 6 1b. D divided in 26–29 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � Dorsal-fin origin on vertical through pelvic-fin origin. Anal fin Beryx splendens Lowe, 1834: 142 (text only, figure is of B. decadactylus 30–34 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � C. spinosus Bay), India, Mauritius Oman, Saya de Malha Bank, Sri Lanka, mid-ventral scutes continuous; light organ shorter Slender alfonsino (type locality: Madeira). origin on vertical through middle of dorsal fin. Pectoral fins Anoplogaster cornuta � � � � � � � Anomalops Kner, 1868 Beryx splendens: Lowe, 1843a: 49; Günther, 1887: 33; McCulloch,) 1929: LL 53–62; transverse scale rows between LL longerand thananal-fin pelvic fins. origin 18; total GR Somalia, Yemen to Indonesia, Australia, Philippines, Taiwan, 1b. C. druzhinini 5 7 than eye, without a black flap to cover it 130; Fowler, 1936a: 542*; Smith, 1949: 150*; Krefft, 1961: 24; Abe, 1972: � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � Fangtooth Beryx splendens Lowe, 1834: 142 (text only,525; figureKarrer, 1973: 221;is Woodsof B. & decadactylusSonoda, 1973: 286* Krefft, 1976: 74*; Centroberyx26–29 druzhinini Top of head, iris, back and caudal fin bright red, body golden. China, Japan and New Zealand. Busakhin, 1982: 6*; Masuda et al., 1984: 108* ; Heemstra, 1986: 410*; Busakhin, 1981: 1728* (type locality: Saya de Caudal fin with white margin and white base, the latter area D 16–20; A 7–9 ; P1 13–16 ; P2 7; BD 1.8–2.2, HL 2.5–2.9 in SL; Trachichthodes druzhinini 3 Maul, 1986: 741*; Kotlyar, 1987: 73; Ivanin, 989: 159; Dubochkin & wider in juveniles than adults. Dorsal, anal and paired fins eye diameter much smaller than interorbital width. Juveniles < 4 (type locality: Madeira). Malha Bank). 8 Kotlyar, 1989: 1; Mundy, 1990: 257*; Smale et al., 1995: 78*; Adam et al., 20 mm SL with a pair of horn-like spines on top of head, and Centroberyx druzhinini: Busakhin, 1982: 9, 14-15*; Masuda et al., 1984: unpigmented in young, paired fins becoming yellow or golden 1998: 11*. another pair (one on each preopercle) extending ventrally. Beryx splendens: Lowe, 1843a: 49; Günther, 1887: 33; McCulloch, 1929: 109* ; Kotlyar, 1993: 190; Masuda & Kobayashi, 1994: 60*. in adults. Known only from west of Mossel Bay (Cape Barracouta), South Africa, north to off Durban. Busakhin’s (1982) statement, Adult body, head, fins and inside of mouth dark brown to black. 130; Fowler, 1936a: 542*; Smith, 1949: 150*;D III–V, 12–15;Krefft, A IV, 25–32; 1961: P 14–19; 24; V I, 9–13;Abe, LL continuous,1972: D V–VII, 12–15; A IV, 15–17; P 13; V I, 7; LL continuous, not 9 6 mistakenly citing Fowler (1935 [not 1936]) as the source, that Attains 16 cm SL. Worldwide in tropical and temperate seas. No 65–70 (69–82 total); GR 5–7/16–21; pyloric caeca 23–34. Depth extending onto caudal-fin rays, 53–62; GR 5–9/ 16–23. Depth the species occurs north of Durban is probably correct, but commercial importance. 3 61 525; Karrer, 1973: 221; Woods & Sonoda,29–37% 1973: SL; head 286* 29–36; Krefft, length dorsal-fin 1976: base 15–22;74*; length 39–42% SL; head 31-36; length dorsal-fin base 34–35; length Family Monocentridae currently unverified by specimens. Depth range: 24–366 m. anal-fin base 26–34; predorsal 37–46; preanal 50–67; distance anal-finCentroberyx base 28–29; predorsal 42–47; preanaldruzhinini 58–64; caudal- 4 10 Busakhin, 1982: 6*; Masuda et al., 1984:dorsal-fin 108* insertion; Heemstra, to caudal-fin 1986: base 36-44; 410*; snout 20-26% HL; fin 39–42; snout 23–28% HL; eye 31–34. Posterior Busakhin, margin of 1981: 1728* (type locality: Saya de eye 29-40. Scales with wedge-shaped rows of triangular ctenii scalesTrachichthodes with uniform-sized denticles, druzhinini no ctenii. Cheek scale rows Maul, 1986: 741*; Kotlyar, 1987: 73; Ivanin,not formed 989: into ridges;159; disc-shaped Dubochkin pad on inner & surface; scales 5. Small fulcral scale in pelvic axil. No head spines, but rough 7 on D and A bases enlarged; fulcral scale in pelvic axil present. areas Malhaaround eyes Bank). and on cheeks. Dorsal-fin origin on vertical Kotlyar, 1989: 1; Mundy, 1990: 257*; Smale et al., 1995: 78*; Adam et al., through origin of pelvic fins. Anal-fin origin on vertical through 11 Cheek scales in 5–6 rows. Single pair of head spines set lateral 2017/07/14 11:36 AM to anterior nostril. Anal-fin origin on vertical with dorsal soft middleCentroberyx of dorsal fin. Pectoral druzhinini fins longer than: Busakhin, pelvic fins. 1982: 9, 14-15*; Masuda et al., 1984: 1998: 11*. rays 10–11. Pelvic-fin origin slightly anterior to vertical through Family Holocentridae 109* ; Kotlyar, 1993: 190; Masuda & Kobayashi, 1994: 60*. 5 pectoral-fin insertion. Pectoral fins longer than pelvic fins Top of head, back and caudal fin bright red, rest of body silvery- Anoplogaster cornuta 6 : 16 cm SL (off New Jersey; 2a Sargocentron diadema: 7 cm SL juvenile (Comoros). in adults. pink, other fins pink or red. Attains 25 cm. Woods & Sonoda 1973). : 18 cm SL (Red Sea; from CFSA). 1 Sargocentron caudimaculatum : 7 cm SL (Chagos). 4 Sargocentron ittodai: 13 cm SL (South 1 Monocentris japonica: 13 cm (South Africa; from CFSA). Tropical Indo-West Pacific, recorded from Japan, Philippines, 1609 WIO Minion +3 Myriad2013.indd Sargocentron inaequalis 61 2 Photoblepharon steinitzi Centroberyx spinosus: 6cm (South Africa; from SSF). 2b Sargocentron diadema: 15 cm SL (South Africa; from CFSA). 6 Sargocentron marisrubri: 12 cm SL (Red Sea). 3 Beryx decadactylus: 25 cm FL (Madeira; PCH). : 7 cm SL (Comoros; PCH.). Head, iris, back, fins, mouth and gill chambers bright red, sides the South China Sea and western Indian Ocean at Saya de Malha Africa). 5 Sargocentron macrosquamis: 7 cm SL (Comoros). 8 9 Sargocentron praslin: 14 cm SL (Solomon 4 Beryx splendens: 19 cm SL (Mozambique; PCH). 8 Sargocentron microstoma: 11 cm SL (Comoros). 5 Centroberyx druzhinini: 21 cm SL (South Africa; L. Scott). Bank and Madagascar. Depth range: 100–200 m. 7 : 11 cm SL (Seychelles). : 11 cm SL (Red Sea). of body silvery-pink. Attains 70 cm TL. D V–VII, 12–15; A IV, 15–17; P 13; V I, 7; LL continuous, not Sargocentron melanospilos : 10 cm SL (Mauritius). 11 Sargocentron rubrum 6 Centroberyx spinosus: 20 cm adult; 4 cm juvenile (South Africa; from CFSA). D III–V, 12–15; A IV, 25–32; P 14–19;Tropical V to I, subpolar 9–13; waters LL of all threecontinuous, major oceans. Depth Ids). 10 Sargocentron punctatissimum range: 25–1240 m. 65–70 (69–82 total); GR 5–7/16–21; pyloric caeca 23–34. Depth extending onto caudal-fin rays, 53–62; GR 5–9/ 16–23. Depth Family Monocentridae 65

29–37% SL; head 29–36; length dorsal-fin base 15–22; length 39–42% SL; head 31-36; length dorsal-fin base 34–35; length 66 1609 WIO Minion + Myriad2013.indd 65 64 Order Beryciformes 67 anal-fin base 26–34; predorsal 37–46; preanal 50–67; distance anal-fin base 28–29; predorsal 42–47; preanal 58–64; caudal- 2017/07/14 11:36 AM 2017/07/14 11:36 AM 2017/07/14 11:36 AM 9 1609 WIO Minion + Myriad2013.indd 67 dorsal-fin insertion to caudal-fin base 36-44; snout 20-26% HL; fin 39–42; snout 23–28% HL; eye 31–34. Posterior margin of 1609 WIO Minion + Myriad2013.indd 66 2017/07/14 11:36 AM eye 29-40. Scales with wedge-shaped1609 WIO Minion + Myriad2013.indd rows 64 of triangular ctenii scales with uniform-sized denticles, no ctenii. Cheek scale rows not formed into ridges; disc-shaped pad on inner surface; scales 5. Small fulcral scale in pelvic axil. No head spines, but rough on D and A bases enlarged; fulcral scale in pelvic axil present. areas around eyes and on cheeks. Dorsal-fin origin on vertical 10 Cheek scales in 5–6 rows. Single pair of head spines set lateral through origin of pelvic fins. Anal-fin origin on vertical through to anterior nostril. Anal-fin origin on vertical with dorsal soft middle of dorsal fin. Pectoral fins longer than pelvic fins. 5 rays 10–11. Pelvic-fin origin slightly anterior to vertical through 6 pectoral-fin insertion. Pectoral fins longer than pelvic fins Top of head, back and caudal fin bright red, rest of body silvery- 1 Monocentris japonica: 13 cm (South Africa; from CFSA). 2 Photoblepharon steinitzi pink, other fins pink or red. Attains 25 cm. 11 3 Beryx decadactylus: 25 cm FL (Madeira; PCH). : 7 cm SL (Comoros; PCH.). in adults. 4 Beryx splendens: 19 cm SL (Mozambique; PCH). Tropical Indo-West Pacific, recorded from Japan, Philippines, 5 Centroberyx druzhinini: 21 cm SL (South Africa; L. Scott).

Head, iris, back, fins, mouth and gill chambers bright red, sides the South China Sea and western Indian Ocean at Saya de Malha Family Holocentridae 6 Centroberyx spinosus: 20 cm adult; 4 cm juvenile (South Africa; from CFSA). Bank and Madagascar. Depth range: 100–200 m. of body silvery-pink. Attains 70 cm TL. 2a : 18 cm SL (Red Sea; from CFSA). Sargocentron diadema: 7 cm SL juvenile (Comoros). Tropical to subpolar waters of all three major oceans. Depth 1 Sargocentron caudimaculatum : 7 cm SL (Chagos). 4 Sargocentron ittodai: 13 cm SL (South 3 Sargocentron inaequalis range: 25–1240 m. 2b Sargocentron diadema: 15 cm SL (South Africa; from CFSA). 6 Sargocentron marisrubri: 12 cm SL (Red Sea). Africa). 5 Sargocentron macrosquamis: 7 cm SL (Comoros). 9 Sargocentron praslin: 14 cm SL (Solomon 8 Sargocentron microstoma: 11 cm SL (Comoros). 7 : 11 cm SL (Seychelles). : 11 cm SL (Red Sea). Sargocentron melanospilos : 10 cm SL (Mauritius). 11 Sargocentron rubrum Ids). 10 Sargocentron punctatissimum 67 64 Order Beryciformes 1609 WIO Minion + Myriad2013.indd 67 2017/07/14 11:36 AM 66 2017/07/14 11:36 AM

1609 WIO Minion + Myriad2013.indd 64

2017/07/14 11:36 AM

1609 WIO Minion + Myriad2013.indd 66 Key to Wio species caudal-fin base 30–37; snout 23–26% HL; eye 37–46. Scales 1a. Inner surface of scales with ovoid, cartilaginous pad near posterior margin; anal fin origin before a vertical with insertion of D; with irregular vertical rows of ctenii formed into low, parallel pyloric caeca 23–100 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 2 ridges; disc-shaped pad on inner surface of scales; scales on D and A bases enlarged; fulcral scale present in pelvic axil; cheek scales in 4–5 rows. Head spine lengths decrease with age. Anal fin origin on vertical with dorsal soft rays 8–9. Pelvic fin origin beneath pectoral fin insertion. Pelvic fins may be elongate in 1a Inner surface of scale from Beryx young. splendens with ovoid pad near posterior margin (redrawn from Busakhin 1982). Top of head, jaws, iris, back and fins bright red. Gill cover 1b. Inner surface of scales without pad near posterior margin; and sides of body silvery-pink. Attains 60 cm TL. anal fin origin on vertical with insertion of D; Found in coastal and open ocean seamount areas throughout Key to Wio species pyloric caeca 12–20 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � caudal-fin base 30–37; snout 23–26% HL; eye 37–46. Scales 1a. Inner surface of scales with ovoid, cartilaginous pad near the Atlantic, southern Indian, western and central Pacific

withB� mollis irregular vertical rows of ctenii formed into low, parallel

C  C  C  C  posterior margin; anal fin origin before a vertical with insertion of D; Weber, 1902 Family Berycidae Genus Photoblepharon pyloric caeca 23–100 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 2 ridges; disc-shaped pad onoceans. inner surface of Depth scales; scales rangeon D 7–1000 m. 2a. Four pairs of spines on head: 1) above eye; 2) in front of; and and A bases enlarged; fulcral scale present in pelvic axil; cheek Body deep. D single, continuous; 1 anal spine; pelvic spine Alfonsinos scales in 4–5 rows. Head spine lengths decrease with age. Anal absent; mid-ventral scutes discontinuous; LL scales enlarged; by M� Eric Anderson 3) below nostrils; 4) at base of lower jaw on vertical with spine abovefin eye; origin on vertical with dorsal soft rays 8–9. Pelvic fin origin 25-30 GR on first arch. Light organ large, longer than eye, with a beneath pectoral fin insertion. Pelvic fins may be elongate in black eyelid-like flap that can be raised to cover the organ. Two dorsal soft rays 16–20; LL 56–61 (not counting scales on C); (spelt palpebratus Body ovoid, compressed. Head about 3 in SL; eye large, rounded, 1a Inner surface of scale from Beryx young. species, 1 in WIO; the other, P. palpebratum splendens with ovoid pad near posterior snout short and steep. Nostrils usually large. Mouth large, oblique; Top of head, jaws, iris, back and fins bright red. Gill cover

by most authors), is widely distributed from central Indonesia depth 40–50% SL; pyloric caeca 74–100margin (redrawn from Busakhin 1982). CHONDRCHTHYES

CHONDRCHTHYES maxilla naked, 2 supramaxillae. Jaw teeth small, in bands; teeth � � � � � � � � � � � � and sides of body silvery-pink. Attains 60 cm TL. to the central Pacific. 1b. Inner surface of scales without pad near posterior margin; B. decadactylus present on vomer and palatines. Dorsal fin triangular, often with anal fin origin on vertical with insertion of D; Found in coastal and open ocean seamount areas throughout first to third soft ray greatly prolonged, with 3–7 spines and 12– pyloric caeca 12–20 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � the Atlantic, southern Indian, western and central Pacific pecies B� mollis Key to s 20 rays; anal fin with 3–4 spines and 12–30 soft rays. Pectoral fin oceans. Depth range 7–1000 m. 2a. F 1a. Anteriormost lateral-line pore preceded by a large white spot on dorsal wedge-shaped, rays 13–19. Pelvic fin with 1 spine and 7–13 soft our pairs of spines on head: 1) above eye; 2) in front of; and � � � � � � � � � � � � � P. palpebratum corner of opercle; pelvic rays typically 7 3) below nostrils; 4) at base of lower jaw on vertical with spine above eye; � � � � P. steinitzi rays. Caudal fin forked, with 16–17 branched rays. Scales large, dorsal soft rays 16–20; LL 56–61 (not counting scales on C); 1b. White spot on dorsal corner of opercle absent; pelvic rays 6 thin, ctenoid; fleshy disc often present on inner surface in Beryx. depth 40–50% SL; pyloric caeca 74–100 � � � � � � � � � � � � B. decadactylus Branchiostegal rays 7–8. Lateral line single, complete,2a poredBeryx LL decadactylus with four scales 36–82. Anterior part of head with few spines (Beryx), or spines absent (Centroberyx). Gill rakers well developed,pairs 22–34 of spines on head. Photoblepharon steinitzi on first arch. Vertebrae 10 + 14–15. Pyloric caeca 20–100. Gas 2a Beryx decadactylus with four Flashlight fish bladder present. pairs of spines on head. Medium-sized fishes of the shelf-slope break and upper Photoblepharon sp. Fridman 1972: 1 (notes on the ecology of a “new species” from the Gulf of Eilat). slope (about 200–600 m), may shoal up in shallower depths, Abe & Haned (sic) 1973: 57, Figs. 1-4 especially at night. Two genera with eight species. Two species Photoblepharon palpebratus steinitzi 2b. One pair of spines on head in front of eye; dorsal soft rays 13–15; LL 65–70; of Beryx are locally abundant and taken commercially in the depth 29–40% SL; pyloric caeca 23–34 (type locality, Ras Burka, Red Sea). McCosker, 1986: 412*. � � � � � � � � � � � � � � � B. splendens Photoblepharon palpebratus: non Boddaert: McCosker & Lagios 1975: WIO in moderate quantities and reduced to fishmeal2b. andOne oil or pair of spines on head in fr 2 (biology of Comoro Island specimens). sold fresh. More highly prized as food fish in the orient. ont of eye; dorsal soft rays 13–15; LLBeryx 65–70; decadactylus : 26 cm (Madeira; from Woods & Sonoda 1973). : McCosker & Rosenblatt 1987: 161 depth 29–40% SL; pyloric caeca 23–34 Photoblepharon steinitzi Beryx decadactylus� � � � � � � � � � � � � � � Plate 7 (steinitzi elevated to specific rank). Key to Genera B. splendens 1a. Dorsal-fin base shorter than anal-fin base; ctenii cover exposed Alfonsino Beryx mollis surface of scale; no V-shaped scales (ventral scutes) on belly Beryx decadactylus Cuvier, 1829: 151 (type locality: Lisbon, Portugal?); Beryx mollis Abe, 1959: 157* (type locality: Sagami, Bay, Japan); Zama & D II–III, 16–18; A I–II, 13–14; P 15–17; V I, 5–6; GR (7–8) + between pelvic fins and anus; head with 1–4 pairs of spines; Cuvier, in Cuvier & Valenciennes, 1829: 222. Yasuda, 1979: 149; Busakhin, 1982: 8*; Kotlyar,1993: 187–190*. 21–22). Depth 2.3–2.6; head 1.9–2.1; eye 3.4 in head. Body and dorsal-fin spines 3–5; anal fin spines 3–4 (usually 3), soft rays 25–30; � � � � � � � � � � Beryx Beryx decadactylus: Lowe, 1843a: 49; Günther, 1887: 33; Fowler, 1911: 173; Beryx decadactylus: 26 cm (Madeira; from Woods & Sonoda 1973). , Megamouth shark fins dark brown to grey-black. Attains 10 cm TL. pectoral-fin rays 14–19; pelvic-fin rays 9–13; LL 61–82 MEGAchASMIDAE McCulloch, 1914: 96; Fowler, 1928: 96*; Maul, 1954: 12*; Krefft, 1961:

Family 1b. Dorsal-fin base longer than anal-fin base; no scale ctenii, D IV, 12–14; A IV–V, 24–31; P 15–19; V I, 10–11; LL continuous,   , Basking shark  posterior margin of scale with spinules only; row of midventral scutes 24*; Penrith, 1967: 528; Abe and Maruyama, 1971: 1* ; Woods & Sonoda, 64–77 (total); GR 6–7/ 16–20; pyloric caeca 12–20. Depth 35– Family cEtORhInIDAE Known from Red Sea and Isle Grande Comore, occupies Beryx decadactylus on belly between pelvic fins and anus; head without spines; 1973: 282* Krefft, 1976: 73; Busakhin, 1982: 3* ; Masuda et al., 1984: 109* 45% SL; head 31–38; length dorsal-fin base 16–22; length anal- crevices and caves of deep reef diurnally and transits to shallow dorsal-fin spines 5–7; anal-fin spines 4, soft rays 12–16; � � � � � � � � � Centroberyx Heemstra, 1986: 409*; Maul, 1986: 740*; Mundy, 1990: 257.

pectoral-fin rays 13–14; pelvic fin rays 7; LL 36–62 fin base 28–37; predorsal 45–52; preanal 56–64; distance dorsal-

Cartilagenous Fishes Cartilagenous Cartilagenous Fishes Cartilagenous water nocturnally to feed, particularly during new moon. Most Beryx delphini Valenciennes, in Cuvier & Valenciennes, 1833: 454 (type fin insertion to caudal-fin base ca. 40-45; snout 16-25% HL; eye probably more widely distributed but undiscovered elsewhere Alfonsino locality: SE of Madagascar); Krefft, 1961: 26. 33-47. Scales with rows of triangularBeryx ctenii andmollis no disc-shaped in western Indian Ocean. Beryx borealis Düben & Koren, 1846: 33 (type locality: near Bergen, pad on inner surface; scales on D and A bases enlarged; fulcral Beryx decadactylus Cuvier, 1829:Norway); Krefft, 151 1961: (type 26. locality: Lisbon, Portugal?);scales in pelvic axil well developed.Beryx Cheek mollis scales Abe, in 6–7 rows.1959: 157* (type locality: Sagami, Bay, Japan); Zama & Genus Beryx Cuvier, 1829 Actinoberyx jugeati Roule, 1923: 1027 (type locality: Morocco?); Roule, Single pair of head spines set lateral to anterior nostril. Anal 1924: 68*. Head spines present; ctenii cover exposedCuvier, surface of inscales; Cuvier & Valenciennes, 1829: 222. fin origin on vertical with dorsalYasuda, fin insertion. 1979: Pelvic 149;fin origin Busakhin, 1982: 8*; Kotlyar,1993: 187–190*. Beryx longipinnis no enlarged scales with ridges on abdomen; usually a fleshy, Plate 8 Barnard, 1925: 504 (type locality: South Africa); Barnard, slightly anterior to vertical through pectoral fin insertion. Pelvic 1927: 363*; Fowler, 1936: 1268. 2 disc-shaped pad on inner surface of scales;Beryx dorsal decadactylus base shorter : Lowe, 1843a: 49; Günther, 1887: 33; Fowler, 1911:fin about 173; as long as pectoral fin in adults. than anal base; some specimens with prolonged dorsal-fin ray; Actinoberyx longipinnis : Smith, 1949: 151. PLATE xx 1 anterior edge of nasal bone without spinules;McCulloch, D III–V, 12-20; 1914: 96; Fowler, 1928: 96*; Maul, 1954: 12*; Krefft, 1961: D III–V, 16–20; A III–IV, 25–30; P 14–18; V I, 9–10; LL Body and fins pinkish redD with IV, silvery 12–14; tones on side, A general IV–V, 24–31; P 15–19; V I, 10–11; LL continuous, Family OphIchthIDAE, Snake-eels, sand-eelsA III, 25–30; V I, 9–13; total LL 61–82. Three species, all in WIO. 24*; Penrith, 1967: 528; Abecontinuous, and 56–61 Maruyama, (60–73 total); GR 1971: 5–7/16–19; 1* pyloric; Woods caeca &colour Sonoda, lighter than the other two species of Beryx; caudal fin 74–100. Depth 40–50% SL; head 30–38; length dorsal-fin base lobes darker than body. Attains64–77 41 cm TL. (total); Probably widespread GR 6–7/ 16–20; pyloric caeca 12–20. Depth 35– 1973: 282* Krefft, 1976:19–24; 73; lengthBusakhin, Orderanal-fin base1982: 35–36; Beryciformes 3* predorsal ; Masuda length 44–59;et al., in tropical1984: and 109* temperate Indo-West Pacific. This species was lost , megamouth shark, 1.9 m TL juvenile preanal length ca. 55–60; distance from dorsal fin insertion to in synonymy with B. splendens45% for 20 SL; years (Zamahead and Yasuda,31–38; length dorsal-fin base 16–22; length anal- 2 Megachasma pelagios Heemstra, 1986: 409*; Maul, 1986: 740*; Mundy, 1990: 257. 1979; Busakhin, 1982), so there may be many misidentified 1 Cetorhinus maximus, basking shark (no size or locality). Photoblepharon steinitzi: ~10cm; the white area below the eye is fin base 28–37; predorsal 45–52; preanal 56–64; distance dorsal- (Eastern Indian Ocean). luminous (Comoros; from SSF). Beryx delphini Valenciennes, in Cuvier & Valenciennes, 1833: 454 (type fin insertion to caudal-fin base ca. 40-45; snout 16-25% HL; eye Family Monocentridae 63 WIOF_TEMPLATE_Openers_EM1.indd 2-3 locality: SE of Madagascar); Krefft, 1961: 26. 33-47. Scales with rows of triangular ctenii and no disc-shaped 62 Order Beryciformes Family LAMnIDAE, Mackerel sharks 223 Beryx borealis Düben1609 &WIO MinionKoren, + Myriad2013.indd 1846: 63 33 (type locality: near Bergen, pad on inner surface; scales on D and A bases enlarged; fulcral 2017/07/14 11:36 AM Norway); Krefft,2017/07/14 11:361961: AM 26. scales in pelvic axil well developed. Cheek scales in 6–7 rows. 1609 WIO Minion + Myriad2013.indd 62 Actinoberyx jugeati Roule, 1923: 1027 (type locality: Morocco?); Roule, 2017/07/14 11:03 AM Single pair of head spines set lateral to anterior nostril. Anal 1924: 68*. Plate 7 fin origin on vertical with dorsal fin insertion. Pelvic fin origin Beryx longipinnis Family MEGAchASMIDAE, Megamouth shark Barnard, 1925: 504 (type locality: South Africa); Barnard, cEtORhInIDAE, Basking shark Plate 8 slightly anterior to vertical through pectoral fin insertion. Pelvic Family 1927: 363*; Fowler, 1936: 1268. PLATE xx Family OphIchthIDAE, Snake-eels, sand-eels fin about as long as pectoral fin in adults. 3 Actinoberyx longipinnis Order Beryciformes 1 : Smith, 1949: 151. 2 4 1 Apterichtus flavicaudus, yellowtail sand-eel, Brachysomophis cirrocheilos, smudged snake- D III–V, 16–20; A III–IV, 25–30; P 14–18; V I, 9–10; LL Body and fins pinkish red with silvery tones on side, general Brachysomophis crocodilinus, crocodile snake- 2 2 Megachasma pelagios, megamouth shark, 1.9 m TL juvenile 80 cm TL (Hawaii; JER). eel, 125 cm TL (Indonesia; JER). colour lighter than the other two species of Beryx; caudal fin 1 Cetorhinus maximus, basking shark (no size or locality). (Eastern Indian Ocean). eel, 44 cm TL (Rodrigues;continuous, PCH). 56–61 (60–73 total); GR 5–7/16–19; pyloric caeca 74–100. Depth 40–50% SL; head 30–38; length dorsal-fin base lobes darker than body. Attains 41 cm TL. Probably widespread Family LAMnIDAE, Mackerel sharks in tropical and temperate Indo-West Pacific. This species was lost 19–24; length anal-fin1 base 35–36; predorsal length 44–59; Apterichtus flavicaudus, yellowtail sand-eel, Brachysomophis cirrocheilos, smudged snake- 2 5 Brachysomophis crocodilinus, crocodile snake- in synonymy with B. splendens for 20 years (Zama and Yasuda, 80 cm TL (Hawaii; JER). eel, 125 cm TL (Indonesia; JER). preanal length ca. 55–60; distance from dorsal fin insertion to 6 eel, 44 cm TL (Rodrigues; PCH). 1979; Busakhin, 1982), so there may be many misidentified 3 4

5 6

4 4 Isurus oxyrinchus, shortfi n mako, 3 Family Monocentridae 63 Brachysomophis henshawi, reptilian snake-eel, , white shark, ~4 m TL mature male (South Africa; from Heemstra & Heemstra 2004). Callechelys catostoma, striped snake-eel, Callechelys marmorata, marbled snake-eel 3 Carcharodon carcharias 4 Isurus oxyrinchus, shortfi n mako, 78 cm TL (Oman; JER). 54 cm TL (Marshall Ids; JER). , white shark, ~4 m TL mature male (South Africa;5 from Isurus Heemstra & paucus Heemstra 2004)., longfi n mako, 2.3 m TL (Japan). 6 Lamna nasus, porbeagle, (Red Sea; SV Bogorodsky). 3 Carcharodon carcharias 5 Isurus paucus, longfi n mako, 2.3 m TL (Japan). 6 Lamna nasus, porbeagle, ~2.5 m TL female (South Africa; from~2.5 m TL femaleHeemstra (South Africa; from& Heemstra & Heemstra 2004). 2004). 4 94 cm TL immature male (NE Atlantic).94 cm TL immature male (NE Atlantic). 3 1609 WIO Minion + Myriad2013.indd 63 Family SphyRnIDAE, Hammerhead sharks Brachysomophis henshawi, reptilian snake-eel, Callechelys catostoma, striped snake-eel, Callechelys marmorata 2017/07/14 11:36 AM Family SphyRnIDAE, Hammerhead sharks 78 cm TL (Oman; JER). , marbled snake-eel 54 cm TL (Marshall Ids; JER). (Red Sea; SV Bogorodsky). 6 8 5

Callechelys marmorata,marbled snake-eel, Cirrhimuraena playfairii, fringelip snake-eel, Ichthyapus vulturis, vulture snake-eel, 32 cm TL (Maldives; JER). 24 cm TL (South Africa; JER). 21 cm TL (Hawaiian Ids; JER). 7

10

9 8

8 Sphyrna lewini, scalloped hammerhead, 47 cm TL immature female (Mozambique). , winghead shark, 1.8 m TL (N Australia). , 8 7 Eusphyra blochii 7 9 Sphyrna zygaena, smooth hammerhead, ~70 cm TL immature female (South Africa; from Heemstra & Heemstra 2004). 10 Sphyrna mokarran great hammerhead, 2.4 m TL (Red Sea). Family Holocentridae : 18 cm (Gulf of Aqaba; Lamnostoma orientalis, oriental snake-eel, Leiuranus semicinctus, halfbanded snake-eel, 2 Ostorhinchus acanthorhinus Muraenichthys schultzei, bluntnose worm-eel, 1 Neoniphon sammara: M43 mangrove 1-12-09 (Mozambique; R. Bills). 21 cm TL (Indonesia; JER). 42 cm TL (Maldives; JER). : 21 cm (Red Sea; JER). 4 Sargocentron seychellense: 15 cm SL (Mauritius; O. Alvheim, 9 cm TL (Eritrea; SV Bogorodsky). D. Golani). 3 Ostorhynchus hypsipterygion 6 Sargocentron tiere: 22 cm (Maldives; JER). 7 5 Sargocentron spiniferum: 30 cm (Marshall Ids; JER). IMR/ASCLME). 8 Sargocentron violaceum: 34 cm (Maldives; JER). 58 7 Sargocentron tiereoides: 15 cm (Maldives;6 JER). 5 10 57 69 Callechelys marmorata,marbled snake-eel, Cirrhimuraena playfairii, fringelip snake-eel, Ichthyapus vulturis, vulture snake-eel, 32 cm TL (Maldives; JER). 24 cm TL (South Africa; JER). 2017/07/14 11:36 AM 21 cm TL (Hawaiian Ids; JER). 1609 WIO Minion + Myriad2013.indd 69

9

8 Sphyrna lewini, scalloped hammerhead, 47 cm TL immature female (Mozambique)., 7 Eusphyra blochii, winghead shark, 1.8 m TL (N Australia). 9 Sphyrna zygaena, smooth hammerhead, ~70 cm TL immature female (South Africa; from Heemstra & Heemstra 2004). 10 Sphyrna mokarran great hammerhead, 2.4 m TL (Red Sea).

8 58 7 Family Holocentridae Lamnostoma orientalis, oriental snake-eel, Leiuranus semicinctus, halfbanded snake-eel, Muraenichthys schultzei, bluntnose worm-eel, 2 Ostorhinchus acanthorhinus: 18 cm (Gulf of Aqaba; 21 cm TL (Indonesia; JER). 42 cm TL (Maldives; JER). 1 Neoniphon sammara: M43 mangrove 1-12-09 (Mozambique; R. Bills). 9 cm TL (Eritrea; SV Bogorodsky). : 21 cm (Red Sea; JER). 4 Sargocentron seychellense: 15 cm SL (Mauritius; O. Alvheim, D. Golani). 3 Ostorhynchus hypsipterygion 6 Sargocentron tiere: 22 cm (Maldives; JER). 5 Sargocentron spiniferum: 30 cm (Marshall Ids; JER). IMR/ASCLME). 8 Sargocentron violaceum: 34 cm (Maldives; JER). 7 Sargocentron tiereoides: 15 cm (Maldives; JER).

57 69

2017/07/14 11:36 AM

1609 WIO Minion + Myriad2013.indd 69 Plate 5

Family SynGnAthIDAE, Seahorses, pipefishes

Glossogobius giuris : Smith 1959b (in part)*; Smith & Smith 1963 Head and body brownish with darker brown spots and (in part); Hoese & Winterbottom 1979; Hoda 1980*; SSF No. 240.44*; mottling; mid-side of body with five X-shaped (some doubled) Maugé 1986a; Keith et al. 1999*. dark brown blotches; 1st dorsal fin with rows of dark spots forming indistinct bands; base of caudal fin with short vertical Second dorsal fin 8 or 9 rays; anal fin 7–9 rays; pectoral fins dark bar or crescent, which may coalesce with posteriormost 17–23 rays; LS 29–35; TRB 8½–11. Predorsal scales 14–25, mid-side X-shaped blotch. Attains 95 mm SL. reaching forward to or just behind eyes. Opercle with patch of cycloid scales dorsally; preopercle naked or with small patch of Distribution WIO: a poorly known species (only type scales behind eye in large adults. In adults, 1st dorsal fin may specimens examined), recorded only from Mauritius Hippocampus borboniensis, Hippocampus cameleopardalis, Hippocampus capensis, Knysna (20 syntypes). Hippocampus debelius, softcoral have at least 2nd spine filamentous, but not greatly so. Réunion seahorse, 60 mm TL giraffe seahorse, 52 mm SL seahorse (South Africa; male (Seychelles; JER). seahorse, female (Red Sea; Remarks Two syntypes from India are a different species. (S Mozambique; PCH). J Lockyear). S Kahlbrock). Head and body light brown to yellowish brown, with 4–7 rounded to elongate dark brown blotches along side and large dark brown triangular to elongate blotch on caudal-fin base, partly extending onto fin; 1st dorsal fin with distinct dark brown to blackish spot near base of 1st spine, specimens about 55 mm SL or smaller with black band across tips of 1st dorsal-fin spines or black blotch over top of fin, black pigment , syntype (Mauritius). remains on spines only in adults. Attains 35 cm SL. Glossogobius kokius

Distribution Indo-West Pacific: East African coast to South (Fowler 1934) Africa (Umtata River mouth), Aldabra, Madagascar, Pakistan, Glossogobius tenuiformis Hippocampus debelius, softcoral Hippocampus fuscus, drab Hippocampus jayakari, spiny Hippocampus kuda, yellow PLATE 13 seahorse, male (Red Sea; Red Sea, Réunion and Seychelles to Japan to northern Australia. Natal flathead goby seahorse, 55 mm TL (Red Sea; seahorse, 73 mm TL (Gulf of seahorse, 137 mm TL S Kahlbrock). SV Bogorodsky). Aqaba; JER). Remarks Widely distributed, enters estuaries and freshwater Glossogobius tenuiformis Fowler 1934b: 496–497, Fig. 49 (St Lucia Lake, 32 (Mozambique; O Alvheim, IMR). river systems, extending well inland. Several species are likely km from mouth, northern KwaZulu-Natal, South Africa). confused under this name. Glossogobius giuris : Hoese 1986b (in part). Glossogobius callidus : Whitfield 1998 (in part)*.

Second dorsal fin 9 rays; anal fin 8 rays; pectoral fins 18–21 rays; LS 28–30; TRB 9–10. Predorsal scales cycloid, 11–16, reaching above, or just in front of, rear preopercular margin. Cheek and opercle naked. First dorsal fin with 2nd spine filamentous. Plate 31 , 23 cm SL (South Africa; from SSF). Hippocampus natalensis, cryptic Glossogobius giuris Hippocampus suezensis, Hippocampus tyro, 61 mm TL Hippocampus sp. 1, African FamilyHead MOnAcAnthIDAE and body yellowish brown to light brown, with 4–8 seahorse (S Mozambique; Egyptian seahorse, 35 mm SL female holotype (Seychelles; V Fraser). prickly seahorse (South Africa; (usually 4) pairs of small dark brown vertically elongate , Filefi shes (Persian Gulf; JER). JER). G de Valle). Glossogobius kokius (Valenciennes 1837) blotches along side, may be smaller brown spots interspersed, and small dark brown blotch at caudal base and two similar Mauritius flathead goby spots on base of fin, may coalesce to form Y-shaped blotch; 1st Gobius kokius Valenciennes in Cuv. & Val. 1837: 68 (Mauritius, Malabar dorsal fin without black blotch or band distally; irregular dusky and Pondicherry, India). band along base which may form dark spot at base of 1st spine. Cuv. & Val., 1837: 78 (Mauritius); Gobius filosus Valenciennes in When alive, dorsolateral part of head and body covered in fine Smith 1959b. orange to red speckles. Attains 74 mm SL. Glossogobius kokius: Akihito & Meguro 1975*; Maugé 1986a; Fricke 1999. Distribution WIO: endemic to South Africa (Zotsha River Second dorsal fin 9–10 rays; anal fin 8 or 9 rays; pectoral fins mouth to Richards Bay, KwaZulu-Natal). Hippocampus sp. 3, Sodwana Hippocampus sp. 3, Sodwana puff Kyonemichthys rumengani, 17–20 rays; LS 24–33; TRB 9–10. Predorsal scales cycloid, Remarks Found in estuaries and freshwaters. Has been puff seahorse (South Africa; Lissocampus bannwarthi, Suez in South African river seahorse (South Africa; pygmy pipedragon (Red Sea; 14–17, reaching to just in front of rear edge of preopercle. confused with G. callidus and G. giuris N Ayliffe). pipefish, 134 mm SL (Egypt; C van Jaarsveld). SV Bogorodsky). JER). Cheek naked; opercle naked or with very small patch of systems. A specimen resembling this species has been collected2 cycloid scales anterodorsally. Second dorsal-fin spine longest, from Réunion. filamentous in large males. 1 100 Order Perciformes

FAMILY Gobiidae 61 3

4 Plate 31 Plate 15 Family MOnAcAnthIDAE, Filefi shes Family GOBIIDAE, Gobies

5 2 2 1 3 3 1 6a 4 4 6

6b 5 5

7

6a 9

6b 8

7 10

7 12

11a

8 9 11b 8 9 15

13 10

14 11 16 18 12 10 17

14 13 20 19 1 Paramonocanthus arabicus11 , Gulf fi lefi sh, 7 cm SL (Oman). 3 Paramonacanthus nematophorus 2 Paramonacanthus frenatus, wedgetail fi lefi sh, 6 cm SL (S Mozambique) 2 Trimma annosum, greybeard pygmy goby, 23 mm SL male paratype5 Trimma (Vietnam). bisella, twosaddle pygmy , seagrass fi lefi sh, 9 cm SL (Red Sea). , pallid pygmy goby (Maldives). 5 Pervagor aspricaudus, orangetail fi lefi sh, 9 cm SL male (Mauritius). 4 Paramonocanthus pusillus, blackstriped fi lefi sh (Mozambique). 1 Trimma anaima 4 Trimma barralli, Barrall’s pygmy goby7 (Red Trimma Sea). dalerocheila , hotlips pygmy goby, (b) 5 cm SL (S Mozambique). 3 Trimma avidori, Avidor’s pygmy goby, 19 mm SL (Yemen). , longspine pygmy goby 7 Pervagor randalli, Randall’s fi lefi sh,6 Pervagor4 cm SL (Redjanthinosoma Sea). , redtail fi lefi sh, (a) 8 cm SL (Mauritius), 6 Trimma corallinum, polkadot pygmy goby, 20 mm SL (South Africa).9 Trimma filamentosus 9 Stephanolepis diaspros goby, 24 mm SL (Mauritius). , yellowtail pygmy goby (a) 24 mm SL, , reticulated fi lefi sh, ~11 cm SL (Oman). 8 Stephanolepis auratus, porky, 19 cm SL (South Africa).12 8 Trimma emeryi, Emery’s pygmy goby, 17 mm SL11 (New Trimma Caledonia). flavicaudatum 11 Th amnaconus erythraeensis, Aqaba fi lefi sh, 19 cm SL (Red 10Sea). Th amnaconus arenaceus, sandy fi lefi sh, 31 cm TL (South Africa). 17 mm SL paratype (Chagos). 13 Trimma griffithsi, Griffith’s pygmy goby, 13 Th amnaconus melanoproctes 12 Th amnaconus fajardoi (Red Sea). 10 Trimma fishelsoni, Fishelson’s pygmy goby, 27 mm SL (Red Sea). , pimple pygmy goby , Oman fi lefi sh, ~19 cm TL (Oman). , spotted fi lefi sh, 16 cm TL (Mauritius). 12 Trimma flammeum, flame pygmy goby, 20 mm SL (South Africa). 15 Trimma halonevum 14 Th amnaconus modestoides (b) 24 mm SL (both Red Sea). , cutface pygmy goby, 21 mm SL (Rodrigues). 14 Trimma haima 17 Trimma mendelssohni, Mendelssohn’s pygmy goby, , drab fi lefi sh, 32 cm SL (Kenya). 17 mm SL female paratype (Chagos). 16 Trimma hoesei, Hoese’s pygmy goby, 21 mm SL female holotype (Chagos). 19 Trimma omanensis, crescentwing pygmy goby (Maldives). , Naude’s pygmy goby, 27 mm SL male (Comoros). 22 mm SL (Rodrigues). 18 Trimma naudei , Sheppard’s pygmy goby (Red Sea). (Oman). 20 Trimma sheppardi

307 59 14 13

1 Paramonocanthus arabicus , Gulf fi lefi sh, 7 cm SL (Oman). 3 Paramonacanthus nematophorus 2 Paramonacanthus frenatus, wedgetail fi lefi sh, 6 cm SL (S Mozambique) , seagrass fi lefi sh, 9 cm SL (Red Sea). 5 Pervagor aspricaudus, orangetail fi lefi sh, 9 cm SL male (Mauritius). 4 Paramonocanthus pusillus, blackstriped fi lefi sh (Mozambique). (b) 5 cm SL (S Mozambique). 7 Pervagor randalli, Randall’s fi lefi sh,6 Pervagor4 cm SL (Redjanthinosoma Sea). , redtail fi lefi sh,(a) 8 cm SL (Mauritius), 9 Stephanolepis diaspros , reticulated fi lefi sh, ~11 cm SL (Oman). 8 Stephanolepis auratus 11 Th amnaconus erythraeensis, Aqaba fi lefi sh, 19 cm SL (Red 10Sea). Th amnaconus arenaceus, sandy fi lefi sh, ,31 porky, cm TL 19 (South cm SL Africa).(South Africa). 13 Th amnaconus melanoproctes 12 Th amnaconus fajardoi , Oman fi lefi sh, ~19 cm TL (Oman). , spotted fi lefi sh, 16 cm TL (Mauritius). 14 Th amnaconus modestoides , drab fi lefi sh, 32 cm SL (Kenya).

307