Diodontidae

Leis, Jeffrey M., 2006, Nomenclature and distribution of the species of the porcupinefish family Diodontidae (Pisces, Teleostei), Memoirs of Museum Victoria 63 (1), pp. 77-90 : 78-79

Family Diodontidae

Diagnosis. Small to medium-sized fishes to 1 m in length, commonly 20–50 cm. Body wide and capable of great inflation, covered with massive spines that may be quite long; spines with large bases, or roots, under the skin; long spines usually erectile and two-rooted, short spines usually fixed in erect position by their three-rooted bases. Head broad and blunt; gill opening a relatively small, vertical slit immediately before pectoral-fin base; nasal organ usually in small tubes located in front of large eyes; mouth large, wide and terminal; teeth fused to form a strong, beak-like crushing structure without a median suture dividing the upper and lower jaws into left and right halves. Dorsal and anal fins without spines, set far back on body, and, like caudal fin, generally rounded; most fin rays branched; bases of fins often thick and fleshy; no pelvic fins. Lateral line inconspicuous. No normal scales.

Genera. There is no generally agreed-upon allocation of species to the nominal genera, nor is there any cladistic analysis of the family or any subset of it. Most authors recognize Diodon (five species, revised by Leis, 1978, ref. 5529) for species in which nearly all the dermal spines are erectile. Four monotypic Indo-Pacific genera, three of which are confined to Australasia, contain species that have a mixture of fixed and erectile spines – Allomycterus , Dicotylichthys , Lophodiodon and Tragulichthys – are usually recognized (see Gomon, 1994, ref. 22532; Leis, 2001, ref. 26318) and are in this paper. Chilomycterus sensu lato (about ten species with nearly all dermal spines fixed and immovable) has been more problematical. Tyler (1980, ref. 4477) recognized three groupings of Chilomycterus : 1) ʻAtlantic Chilomycterusʼ (five species confined to the Atlantic, and called by him antennatus , antillarum , mauretanicus , schoepfii and spinosus ); 2) what I call herein ʻCircumtropical Chilomycterusʼ (a circumtropical group considered by Tyler to consist of four species called by him affinis , atinga , reticulatus and tigrinus ); and 3) ʻIndo-Pacific Chilomycterusʼ (considered by Tyler to consist only of orbicularis ). I agree with Tyler (1980, ref. 4477) that nominal species in each of these three groups are morphologically more similar to each other than they are to species in the other groups. The type species of Chilomycterus is Diodon reticulatus Linnaeus (1758, ref. 2787); thus, if these groupings prove to be valid at the generic level,the circumtropical group becomes Chilomycterus , and I use it in that sense herein. Cyclichthys typically is used for several Indo-Pacific species (including two not mentioned by Tyler [1980, ref. 4477]), and the type of Cyclichthys is the Indo-Pacific orbicularis ; thus, the Indo-Pacific grouping can be considered Cyclichthys for the purposes of this paper. The Atlantic group of species, regarded by Tyler to be the most phylogenetically basal, is nearly always included in Chilomycterus . If these Atlantic species were removed from Chilomycterus , the generic name available for them is Lyosphaera , based on the unique pelagic stage found in at least some members of this group. Unfortunately, the identity of the type species is not clear (on the basis of distribution, schoepfii seems most likely). Lyosphaera has not been used as a generic name for these five Atlantic species, and until a full cladistic analysis is performed on the group, its use is not recommended. For the purposes of this paper, I use Tylerʼs term, “Atlantic Chilomycterus ”, to identify this grouping.

Chilomycterus (ex Bibron) Brisout de Barneville, 1846 (sensu stricto)

Chilomycterus (ex Bibron) Brisout de Barneville , 1846 (type species, Diodon reticulatus Linnaeus )

Cyanichthys Kaup, 1855 (type species is D. coeruleus [non- D. caeruleus Quoy and Gaimard ] Kaup = D. reticulatus Linnaeus, 1758 )

Diagnosis. All spines fixed, with long subdermal bases but short or absent external spines (relatively smaller in larger individuals); some spines on top of head with 4 bases; 10 C rays; 21–23 vertebrae; heavy jaw teeth, but trituration teeth few; no tentacles; nostril in adult an open, cup-shaped organ with reticulations; 1 or more spines wholly on dorsal surface of caudal peduncle; fins spotted; no large blotches on dorsal surface of head or trunk. Some additional osteological characters are given by Tyler (1980, ref. 4477).

The type species of Chilomycterus is Diodon reticulatus Linnaeus (1758, ref. 2787). Cyanichthys coeruleus Kaup (1855, ref. 2571) was based on an unregistered BMNH specimen of 43 mm SL (see also Günther, 1870, ref. 1995). Although Kaup Nominal species Current identity Nominal species Current identity affinis Günther 1870 Chilomycterus reticulatus (Linnaeus 1758) mauretanicus Le Danois 1954 Chilomycterus spinosus mauretanicus (Le antennatus Cuvier 1816 Chilomycterus antennatus (Cuvier 1816) Danois 1954 ) antillarum Jordan & Rutter 1897 Chilomycterus antillarum Jordan & Rutter melanopsis Kaup 1855 Diodon eydouxii Brisout de Barneville 1846

1897 meulenii Walbaum 1792 Chilomycterus schoepfii (Walbaum 1792) areolatus Gronow in Gray 1854 Chilomycterus schoepfii (Walbaum 1792) multimaculatus Cuvier 1818 Diodon holocanthus Linnaeus 1758 armillatus Whitley 1933 Diodon hystrix Linnaeus 1758 myersi Ogilby 1910 Dicotylichthys punctulatus Kaup 1855 atinga Bloch 1785 Diodon hystrix Linnaeus 1758 nicthemerus Cuvier 1818 Diodon nicthemerus Cuvier 1818 atringa Linnaeus 1758 not identifiable nigrolineatus Ayres 1842 Chilomycterus schoepfii (Walbaum 1792) bertolettii Lema Lucena Saenger & Diodon eydouxii Brisout de Barneville 1846 nigropunctatus Smith 1957 Lophodiodon calori (Bianconi 1854) Oliveria 1979 novemmaculatus Cuvier 1818 Diodon holocanthus Linnaeus 1758 bleekeri Günther 1910 Diodon liturosus Shaw (ex Lacepède) 1804 nudifrons Jenkins 1904 Diodon hystrix Linnaeus 1758 blochii Castelnau 1872 Diodon nicthemerus Cuvier 1818 ? orbe Lacepède 1798 Chilomycterus spinosus spinosus (Linnaeus bocagei Steindachner 1866 Diodon holocanthus Linnaeus 1758 1758) brachiatus Bloch and Schneider 1801 Diodon hystrix Linnaeus 1758 orbicularis Bloch 1785 Cyclichthys orbicularis (Bloch,1785) briareos Metzelaar 1919 Chilomycterus antennatus (Cuvier 1816) orbitosus Poey 1875 Chilomycterus antillarum Jordan & Rutter caeruleus Quoy & Gaimard 1824 Cyclichthys orbicularis (Bloch,1785) 1897 ? californiensis Eigenmann 1891 Chilomycterus reticulatus (Linnaeus 1758) paraholocanthus Kotthaus 1979 Diodon holocanthus Linnaeus 1758 calori Bianconi 1854 Lophodiodon calori (Bianconi 1854) parcomaculatus von Bonde 1923 Cyclichthys orbicularis (Bloch,1785) cornutus Kaup 1855 Chilomycterus spinosus spinosus (Linnaeus pentodon Atkinson in Bryant 1888 Chilomycterus schoepfii (Walbaum 1792)

1758) ? pilatus Whitley 1931 Allomycterus pilatus Whitley 1931 digitalis Breder 1927 Chilomycterus antennatus (Cuvier 1816) pilosus Mitchill 1815 Diodon holocanthus Linnaeus 1758 diversispinis Bleeker (ex Verreaux) 1865 Dicotylichthys punctulatus Kaup 1855 plumierii Lacèpede 1798 Diodon hystrix Linnaeus 1758 ? echinatus Linnaeus 1758 Chilomycterus reticulatus (Linnaeus 1758) punctatus Cuvier 1818 Diodon hystrix Linnaeus 1758 eydouxii Brisout de Barneville 1846 Diodon eydouxii Brisout de Barneville 1846 punctulatus Kaup 1855 Dicotylichthys punctulatus Kaup 1855 fuliginosus deKay 1842 Chilomycterus schoepfii (Walbaum 1792) quadradicatus Waite 1900 Chilomycterus reticulatus (Linnaeus 1758) galapagosensis Klausewitz 1958 Chilomycterus reticulatus (Linnaeus 1758) quadrimaculatus Cuvier 1818 Diodon holocanthus Linnaeus 1758 geometricus Bloch & Schneider 1801 Chilomycterus antillarum Jordan & Rutter reticulatus Linnaeus 1758 Chilomycterus reticulatus (Linnaeus 1758)

1897 rivulatus Cuvier 1818 Chilomycterus schoepfii (Walbaum 1792) grandoculis Ogilby 1910 Tragulichthys jaculiferus (Cuvier 1818) schoepfii Walbaum 1792 Chilomycterus schoepfii (Walbaum 1792) hardenbergi de Beaufort 1939 Cyclichthys hardenbergi (de Beaufort 1939) sexmaculatus Cuvier 1818 Diodon holocanthus Linnaeus 1758 holocanthus Linnaeus 1758 Diodon holocanthus Linnaeus 1758 spilostylus Leis & Randall 1982 Cyclichthys spilostylus (Leis and Randall hystrix Linnaeus 1758 Diodon hystrix Linnaeus 1758 1982) jaculiferus (non-Cuvier) McCulloch 1921 Allomycterus pilatus Whitley 1931 spinosissimus Cuvier 1818 Diodon hystrix Linnaeus 1758 jaculiferus Cuvier 1818 Tragulichthys jaculiferus (Cuvier 1818) spinosus Linnaeus 1758 Chilomycterus spinosus spinosus (Linnaeus lineatus Bloch & Schneider 1801 Chilomycterus schoepfii (Walbaum 1792) 1758) lissogenys Günther 1910 Chilomycterus reticulatus (Linnaeus 1758) tigrinus Cuvier 1818 Chilomycterus reticulatus (Linnaeus 1758) liturosus Shaw (ex Lacepède) 1804 Diodon liturosus Shaw (ex Lacepède) 1804 torosus Larrañaga 1923 Chilomycterus spinosus spinosus (Linnaeus maculato-striatus Mitchill 1815 Chilomycterus schoepfii (Walbaum 1792) 1758)? maculatus Duméril 1855 Diodon liturosus Shaw (ex Lacepède) 1804 totara Curtiss 1938 Diodon hystrix Linnaeus 1758 maculatus Hermann 1804 not identifiable triedricum Cuvier 1818 Diodon hystrix Linnaeus 1758 maculifer Kaup 1855 Diodon holocanthus Linnaeus 1758 ? verrucosus deKay (ex Mitchill) 1842 Chilomycterus schoepfii (Walbaum 1792)

whiteleyi Phillipps 1932 Allomycterus pilatus Whitley 1931 ? identified this specimen as D. coeruleus Quoy and Gaimard (an alternate spelling of caeruleus ), it is clearly a juvenile, pelagic specimen of Chilomycterus reticulatus (Linnaeus) . Kaup (1855, ref. 2571) asserted the specimen was from New Guinea, but according to Günther (1870, ref. 1995) the locality of the specimen “was never known at the British Museum”. A single species (see below).