'' BULLETIN DE L'fNSTITUT ROYAL DES SCfENCES NATURELLES DE BELGIQUE, BIOLOGfE, 76: 147-1 89,2006 BULLETIN VAN HET KONlNKLIJK BELGISCH LNSTITUUT VOOR NATUURWETENSCHAPPEN, BIOLOGlE, 76: 147-1 89,2006

Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fishes by Dirk NOLF & James C. TYLER

Abstract rine waters worldwide, with a few families absent from the Atlantic and eastern Pacific Oceans. They include the primi­ Otolith features are used to test osteology-based hypotheses about tive, deep-water, bottom-dwelling Triacanthodidae, but most interrelationships among caproid, zeiform and tetraodontiform families are more typical of shallow water and often are as­ fishes and intrarelationships of the taxa within each concerned sociated with coral reefs or surrounding sea grass beds and group. Otolith morphology suggests that caproids are the continental sand I mud flats: the Triacanthidae, Balistidae, plesiomorphic sister group of [zeiforms + tetraodontiforms]. In Monacanthidae, Aracanidae, Ostraciidae, Tetraodontidae zeiforms, otolith data are in reasonable agreement with the homoge­ and Diodontidae. One family, the Triodontidae, has a single neity of the families defined upon osteology, but they are of little use in supporting interrelationships among these families. Problem­ extant species found in deep Indo-western Pacific bottom atic cases are the plesiomorphic otoliths of Parazen and Zenion, waters, whereas the three species of Molidae are oceanic which would group these two taxa as a plesiomorphic sister group pelagic. of all other zeiforms. The placement of Parazen as sister to The phylogenetic relationships of the various clades of [Cyttopsis + Stethopristes] in parazenids and Zenion as sister to zeiform and tetraodontiform fishes and the affinities of both [Capromimus + Cytomimus] in zeniontids, however, is logically groups to other teleost taxa have been the subject of many consistent within both families. In tetraodontiforms, otoliths exhibit studies during the last 25 years. An overview of the pub­ a broad range of very different morphologies, but there are no ap­ lished work and a modern cladistic synthesis of the now parent contradictions with the osteology-based phylogenetic hy­ available data is presented in two recently published papers: potheses. The main interest of otolith morphologies among TYLER et al. (2003) and SANTINI & TYLER (2003), treating, tetraodontiforms is the evidence that strongly supports the recogni­ respectively, the zeiforms and the tetraodontiforms. Except tion of three clades within balistids, and the confirmation of tetraodontids and diodontids as, respectively, plesiomorphic and for some schematic drawings of extant triacanthodid otoliths apomorphic sister groups. in TYLER ( 1968) and a synoptic note by SCHWARZHANS (1996) in a paper on fossil otoliths, very little has been done Key words: otoliths, Caproidei, Zeiformes, Tetraodontiformes, on the systematic interpretation of the otoliths of these two phylogeny. orders of fishes. WINTERBOTTOM (1974) suggested that at least some Introduction zeiforms might be related to tetraodontiforms. ROSEN (1984) provided argumentation for a group comprising caproids, The order Zeiformes comprises mostly deep-bodied benthic tetraodontiforms and zeiforms, with the latter two being sis­ fishes found in marine waters worldwide. Many species of ter groups. TYLER et al. (2003) provided strong evidence for the order are deep-sea fishes, found at depths that may ex­ the monophyly of zeiforms and discussed a possible rela­ ceed 1000 m. Some species, however, especially in the fam­ tionship of zeiforms with tetraodontiforms and caproids, but ily Zeidae, live at depths of 100-300 m, and, in European they concluded that such relationships were ambiguous. waters, Zeus faber occurs usually between 50 and 150m but These putative relationships will be evaluated herein on the is occasionally caught as shallow as 10 m (NELSON, 2006, basis of otoliths. QUERO et al., 2003). There are six Recent families (some di­ We herein provide an overview of the previously published vided into subfamilies, see section on classification), with illustrations of zeiform and tetraodontiform otoliths and an about 16 genera and 32 species. Fig. 1 provides a pictorial extensive series of new figures, covering almost all existing overview of a representative of each of the recognized Re­ Recent subfamilial and higher clades. After a brief icono­ cent families and subfamilies. graphic and descriptive overview, otolith morphology is There are approximately 350 species of extant used to test the phylogenetic hypotheses based upon osteo­ Tetraodontiformes, grouped in ten families of great diversity logical features, and some new affinities are proposed on the in body form and osteological structure (Fig. 2). Most of basis of the otoliths. We believe that our coverage is very these families are found in warm and temperate shallow rna- satisfactory in providing an overall survey of otolith mor- I I

148 DIRK NOLF & JAMES C. TYLER phology in the various tetraodontiform clades, but the same ture has nothing "pseudo" about it), whereas "collicular cannot be said about our survey of the intraspecific variabil­ crest" is concise, standard language that says exactly what is ity of the otoliths of these fishes. intended. A significant problem is that tetraodontiform otoliths cannot be dissected from mu seum specimens without severe exter­ nal damage to the fishes because the classical dissection Classification of zeiforms and. tetraodontiforms; method through the gill apertures cannot be applied. Moreo­ iconography and brief descriptions of their otoliths ver, many tetraodontiform otoliths show fine salient struc­ tures and crests that are readily attacked by the preliminary Preliminary note concerning the iconography. -To make our formalin fixation that had been applied to most preserved figures as understandable and comparable as possible, we specimens. Also, only in a limited number of cases was it tried to represent all specimens by their left otoliths. Right possible to give even a small overview of the intraspecific otoliths were figured only if no left ones were available, or otolith variability. This is because, unfortunately, in many when the right otoliths provided some additional informa­ cases the otolith of only a single specimen was available. A tion. When a figure represents a right otolith, this is always single otolith provides some idea of the structure in that spe­ mentioned directly on the plate, and when both left and right cies and how that species may be related to other taxa, but it otoliths are figured, the left specimen is figured on the left does not adequately define the otolith morphology of that side and the right specimen on the right side, according to the species. The same is true for the illustration of ontogenetic CHAINE & DUVERGIER (1934) principle of representing changes. Therefore, the present study should be considered a otoliths. Locality and collection data are provided in an ad­ preliminary overview that still requires much additional in­ dendum. All scale bars represent one millimeter. The annota­ formation in such a speciose and rughly diversified group as tions Fig. a, b and care used to indicate, respectively, ventral, tetraodontiforms. inner(= mesial) and posterior views. Figures without a letter annotation show inner views.

Otolith terminology ZEIFORMES All the materials studied herein are saccular otoliths. Except Plesion: for some brief remarks on the lagaenar and utricular otoliths [CRETAZEIDAE, extinct; Late Campanian/Early of Balistes and Zeus in two papers by ASS IS (2003 and 2005), Maastrichtian; Nardo, Puglia, Italy; otoliths unknown] these two types of otoliths, ~v hich are very small in the groups studied herein, have scarcely been used in systematic studies and almost no collections of them are available. Fig. 3 CYTTOIDEI provides an overview of the terminology first proposed by CYTTIDAE CHAINE & DUVERGIER (1934) and now universally used for Available iconography: saccular otoliths; this generalized scheme is reproduced Cyttus australis (RICHARDSON, 1843) herein. Because many taxa of the groups studied herein have Pl. 1, Fig. 3 strongly modified otoliths, the figure of Chaine & Duvergier Cyttus novaezelandiae (ARTHUR , 1885) is followed by selected examples of some zeiform and Pl. 1, Figs. 4-6 tetraodontiform otoliths, where the homologies of their spe­ ciali zed structures are indicated with the standard terminol­ Cyttus traversi HUTION , 1872 ogy. Pl.l,Figs. l-2 We also figure an otolith of a myctophid and of a gadid fish SMAL~ et al., 1995, pl. 45, fig. D 1 because these otoliths have a well-marked collicular crest, a structure that is also visible in the otoliths of several zeiforms Cyttid otoliths are characterized by a very strong constriction and of some tetraodontiforms. Collicular crests are structures along the antero-posterior midline, and by the absence or that occur in the otoliths of many deep-water fishes with only rudimentary development of collicula. In the rather large otoliths, e.g., myctophids, beryciforms and gadiforms. small otoliths of Cyttus australis and C. novaezelandiae fig­ These ridges emerge near to and parallel to the crista inferior. ured herein, collicula are completely lacking and the distal In all groups (often unrelated), collicular crests are located portions of the ostium and the cauda are hollowed out. In the near the caudal crista inferior. When they occur in gadiform two figured large otoliths of C. traversi, rudimentary otoliths, they always occupy a central position (this is at the collicula are formed. In all cyttids, neither the ostium nor the transition of the ostial and caudal crista inferior), which fea­ cauda shows a ventral expansion. This readily allows cyttid ture constitutes a synapomorphy for gadiforms. Manifest otoliths to be distingui shed from those of oreosomatids, collicular crests with an ostial position are unknown in all whi ch otherwise have a very similar outline. In Cyttus groups of fishes. Although the collicular crests of zeiforms australis and C. traversi, the swollen upper portion of the often occupy a central position, they are mani festly caudal ventral area is small and elongate; in C. novaezelandiae, it is structures, which is evident when their otoliths are compared more extended ventrally. All examined otoliths have a well­ with those of Antigonia (Fig. 6). Collicular crests have often formed collicular crest. been called "pseudocolliculum" in various papers on otoli ths, a term of useless and inappropriate jargon (the struc- '' Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fishes 149

ZEOIDEI rior and a crista inferior that are clearly divided into ostial 0REOSOMATIDAE, Pseudocyttinae and caudal portions also is a plesiomorphic feature. For other Available iconography: features, see Cyttopsinae. Pseudocyttus maculatus (GILCHRIST, 1906) Pl. 1, Figs. 7-9 GON & HEEMSTRA, 1990, fig. 3, p. 227 PARAZENIDAE, Cyttopsinae SMALE et al., 1995, pl. 46, figs. D1-2 Available iconography: Cyttopsis cypho (FOWLER, 1934) For general features of oreosomatid otoEths, see Oreoso­ Pl. 3, Fig. 3 matidae, Oreosomatinae, herein. Pseudocyttus otoliths may RIVATON & BOURRET, 1999, pl. 144, figs. 7-11 be distinguished from those of the other oreosomatids by Cyttopsis roseus (LOWE, 1843) their somewhat stronger antero-posterior constriction and Pl. 3, Fig. 4 their rather wide dorsal area, but examination of a large series ASSIS, 2004, fig. 56, p. 118 of all oreosomatid species would be necessary to confirm CAMPANA, 2004, p. 277 (sac., utr., lag.) these differences. CHAINE, 1958, pl. p. 236 OHE, 1985, fig. 114, p. 76, as Zen itea 0REOSOMATIDAE, Oreosomatinae Stethopristes eos GILBERT, 1905 Available iconography: Pl. 3, Fig. 5 KOTLYAR & PARIN, 1990, fig. 5, p. 110 Allocyttus folletti MYERS, 1960 RIVATON & BOURRET, 1999, pl. 144, figs. 12-13 Pl. 2, Fig. 3 NOLF, 1985, fig. SSE, p. 73 Otoliths of the Cyttopsinae are rather similar to those of Allocyttus niger JAMES, lNADA & NAKAMURA, 1988 oreosomatids in their outline, but they do not exhibit the ven­ Pl. 2, Figs. 1-2 tral expansions of the ostium and of the posterior end of the Allocyttus verrucosus (GILCHRIST, 1906) cauda as in the latter family. Their collicula are marked, but Pl. 2, Fig. 4 they do not show as much hypertrophy as in oreosomatids. SMALE et al., 1995, pl. 46, figs. A1- 3 Swelling of the central upper part of the ventral area is visible Neocyttus helgae (HOLT & BYRNE, 1908) but is not as strongly marked as in parazenin otoliths. Pl. 2, Fig. 5, drawn by W. Sc~warzhans Neocyttus rhomboidalis GILCHRIST, 1906 ZENIONTIDAE Pl. 2, Fig. 6, redrawn after Smale et al., 1995 Available iconography: Smale et al., 1995, pl. 46, figs. B 1-3 Oreosoma atlanticum CUYlER, 1829 Capromimus abbreviatus (HECTOR, 1874) Pl. 2, Fig. 7, redrawn after SMALE et al., 1995 Pl. 3, Fig. 6 SMALE et al. , 1995, pl. 46, fi gs. C2- 3 Cyttomimus stelgis GILBERT, 1905 Pl. 3, Fig. 7 Oreosomatid otoliths are characterized by strong ventral ex­ KOTLYAR & PARIN, 1990, fig. 5, p. 110 pansions of the ostium and of the posterior end of the cauda. RIVATON & BOURRET, 1999, pl. 144, figs. 5-6 Both expansions are filled with well-developed collicular Zenion hololepis (GOODE & BEAN, 1896) formations of swollen aspect. The available material does not Pl. 4, Fig. 6 permit pertinent comments about differences among the taxa KOTLYAR & PARIN, 1990, fig. 5, p. 110 constituting the family. NOLF & CAPPETTA, 1989, pl. 14, fig. 12 SMALE et al. , 1995, pl. 45, fig. C2 Zenionjaponicus KAMOHARA, 1934 PARAZENIDAE, Parazeninae Pl. 4, Fig. I Available iconography: NOLF, 1985, fi g. 55B, p. 73 Parazen pacific us KAMOHARA , 1935 OH E, 1985, fi g. 116, p. 76, as Zenion pacificum Pl. 3, Figs. 1- 2 Zenion longipinnis KOTTHAUS, 1970 KOTLYAR, 2001, fi g. ld, p. 689 Pl. 4, Figs. 7- 8 NOLF, 1985, fig. 55A, p. 73 RIVATON & BOURRET, 1999, pl. 144, fi gs. 1-2 0HE, 1985, fig. 11 2, p. 76 SCHWARZHANS, 1980, fi g. 362, p. 110 Zenion sp. RIVATON & BOURRET, 1999, pl. 144, figs. 3-4 Among parazenids, the monotypic subfamily Parazeninae Zenion sp. displays the plesiomorphic otolith condition. The posterior SCHWARZHANS, 1980, fig. 361, p. 110 constriction is almost nonexistant in the specimen figured herein, but Ohe ( 1985) figured a specimen with a more Zeniontids can easily be divided into two groups on the basis marked posterior constriction. The presence of a crista supe- of their otolith morphology: Zenion and [Capromimus + II

150 DIRK NOLF & JAMES C. TYLER

Cyttomimus]. The most plesiomorphic condition (very rudi­ Zeus faber LINNAEUS, 1758 mentary antero-posterior constriction) is present in Zenion. Pl. 5, Figs. 5-8 Both groups have swollen collicula. In Zenion the collicula ASSIS, 2004, fig. 56, p. 11 8 are centrally located, whereas in Capromimus and CHAfNE, 1958, pl. p. 237, under the names of Cyttomimus the collicula are reduced to small swellings at Z. faber and Z. pungio the extreme anterior and posterior ends of the otoliths. In FROST, 1927, pl. 8, figs. 8-9 (sac., utr., lag.) Cyttomimus, the swollen central upper part of the ventral area HARKONEN , 1986, pl. 4 1, figs. A-B is more strongly developed and concentrated in the most cen­ NOLF, 1985, fig. 55C, p. 73 tral part of the otolith; the collicula protrude on the anterior SCHMIDT, 1968, pl. 4, fig. 58; pl. 18 and posterior sides of the otolith, and ventrally the beginning SMALE et al., 1995, pl. 45, figs. G 1-2 of the formation of an anterior and posterior lobe is visible. Zeid otoliths, with their tri-lobed outline, can be immediately distinguished from those of any other fish taxon. Other G RAMMICOLEPIDIDAE, Macrurocyttinae marked features are the protruding collicula that form addi­ Contained genus: Macrurocyttus; otoliths unknown. tional small anterior and posterior lobes, and the rim that is smooth along almost the entire outline of the otolith. In Zeus, GRAMMICOLEPIDIDAE, Grammicolepidinae the lobes of the ventral portion are directed further outward Available iconography: than in Zenopsis. Grammicolepis brachiusculus POEY, 1873 Pl. 5, Figs. 1- 2 TETRAODONTWORMES CAMPANA, 2004, pl. on p. 276, as Daramattus [PLECTOCRETACICOIDEI, extinct] america nus [CRETATRIACANTHIDAE, extinct; Upper Campanian­ OH E, 1985, fig. Ad 433, p. 75 Lower Maastrichtian; Nardo, Puglia, Italy; otoliths un­ Xenolepidichthys da/g/eishi GILCHRIST, 1922 known] Pl. 5, Fig. 3 [PLECTOCRETACICIDAE, extinct; Lower Cenoma­ NOLF, 1985, fig. 55D, p. 73 nian; Hake!, Lebanon; otoliths unknown] 0HE, 1985, fig. 111 , p. 75 [PROTRIACANTHIDAE, extinct; Upper Cenomanian­ RI VATON & BOURRET, 1999, pl. 144, figs. 14-18 Lower Turonian; Comen, Slovenia; otoliths unknown] SMALE et al. , 1995, pl. 46, figs. E l- 3

Otoliths of the Gramicolepididae are characterised by a TRIACANTHODOIDEI strong antero-posterior constriction and a sulcus with a hol­ TRIACANTHODIDAE, Hollardiinae low ostium and posterior caudal end. Collicula are either Available iconography: lacking or only developed as flat rudimentary structures. The Hollardia hollardi POEY, 1861 ostium and the posterior end of the ventral area are very sali­ TYLER, 1968, fig. 31G, p. 102 ent. There is a marked posterodorsal angle, which is mainly developed backward. This otolith type is very similar to that Parahollardia linea/a (LONGLEY, 1935) of cyttids; in fact, the only feature that seems to distinguish TYLER, 1968, fig. 18, p. 79 them is that in cyttids the posterodorsal angle is more ex­ Parahollardia schmidti WOODS, 1959 panded in the dorsal direction. The otolith represented on Pl. TYLER, 1968, fig. 250, p. 88 5, Fig. 2 shows a ventral rim with a manifest concave portion just in front of the salient posterior end. Such hollowing is Otoliths of hollardiin triacanthodids are known only by the not seen in the smaller otoliths figured by Campana (herein schematic drawings of TYLER (1 968), which depicted only reproduced on Pl. 5, Fig. I) and by Ohe, and it may be a fea­ the outlines of the otoliths and were figured upside down in ture that appears only at a certain size. all three cases. Unfortunately, our search (mostly at the Academy of Natural Sciences of Philadelphia) for these original otolith materials of cleared and stained specimens ZEIDAE was unsuccessful ; the otoliths probably have been dissolved Zenopsis conchifer (LOWE, 1852) or dispersed in the residue of the much dissected al izari ne Pl. 5, Fig. 4 preparation. These outline drawings, however, show a gen­ CAMPANA, 2004, p. 277 eral similarity with the triacanthodid otolith ill ustrations CHA INE, 1958, pl. p. 237 herein, except that they show a stronger constriction along SCHMIDT, 1968, pl. 5, fig. 59; pl. 18 the antero-posteri or midline. SMALE eta!., 1995, pl. 45, figs. E J- 2 Zenopsis nebulosus (Temrninck & Schlegel, 1845) Pl. 5, Figs. 9- 12 T RI ACANTHODIDAE, Triacanthodinae OHE, 1985, fi g. 11 3, p. 76 Avai lable iconography: Zeus capensis Y ALENCIENNES, 1835 Halimochirurgus centriscoides ALCOCK, 1899 SMALE et al. , 1995, pl. 45, figs. Fl -4 TYLER, 1968, fi g. 90G, p. 223 \I

Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fishes 151

]ohnsonina eriomma MYERS , 1934 Tripodichthys oxycephalus (BLEEKER, 1851) TYLER , 1968, fig.57G, p. 158 TYLER, 1968, fig. 132G, p.316 Macrorhamphosodes sp. Trixiphichthys weberi (CHAUDHURI , 1910) SMALE et al. , 1995, pl. 147, fig. A1 Pl. 6, Fig. 5 Macrorhamphosodes platycheilus FOWLER , 1934 TYLER, 1968, fig. 121G, p.288 TYLER, 1968, fig. 84G, p. 214 Triacanthid otoliths are comparable to th~~ e of triacan­ Macrorhamphosodes uradoi (KAMOHARA, 1933) thodids because of their rarher deeply incised sulcus, with at OHE, 1985, fig. 415, p. 173 least some rudiments of a crista superior, and because of their TYLER, 1968, fig. 78G, p. 202 antero-posterior constriction. Their dorsal area, however, is Paratriacanthodes retrospinis FOWLER, 1934 higher and narrower, and their rostrum is very salient relative Pl. 6, Fig. 3 to triacanthodids. TYLER, 1968, fig. 52G, p. 147 Triacanthodes anoma/us TEMM!NCK & SCHLEGEL, 1850 [MOCLAYBALISTOIDEA, extinct] OHE, 1985, fig.414, p.173 [MOCLAYBALISTIDAE, extinct; late Paleocene, Mo­ TYLER, 1968, fig. 2B, p. 8; fig. 39G, p. 120 Clay (Fur) Formation; Jutland, Denmark; otoliths un­ Triacanthodes ethiops ALCOCK , 1894 known] Pl. 6, Fig. 1 NOLF, 1985, fig. 79A, p. 105 [BOLCABALISTOIDEA, extinct] R!VATON & BOURRET, 1999, pl. 83, figs. 1-6 [BOLCABALISTIDAE, extinct; middle Eocene, TYLER, 1968, fig. 43G, p. 127 Lutetian; Monte Bolca, Italy; otoliths unknown] [EOSPINIDAE, extinct; late Paleocene, Danatinian For­ Triacanthodes sp. mation; Uylya-Kushlyuk, Turkmenistan; otoliths un­ R!VATON & BOURRET, 1999, pl. 83, figs. 7-8 known] Tydemania navigatoris WEBER, 1913 Pl. 6, Fig. 2 BALISTOIDEA RIVATON & BOURRET, 1999, pl. 83, figs. 9-10 BALISTIDAE TYLER , 1968, fig. 70, p. 187 Available iconography: Otoliths are unknown for the genera Atrophacanthus, Balistapus undulatus (MUNGO PARK, 1797) Bathyphylax and Halimochirurgus. Pl. 7, Fig. 4 Triacanthodids are characterized by round shaped otoliths NOLF, 1985, fig. 79B, p. 105 with a well-marked antero-posterior constriction. The outer SMALE et al., 1995, pl. 147, fig. B1 face is slightly concave and nearly flat; the inner face is Balistes capriscus GMELIN, 1789 strongly convex. The sulcus is well incised, and in at least Pl. 8, Figs. 1-6 some otoliths, there is a slight to distinct crista superior. The Assrs, 2004, fig. 91, p. 161 , as B. carolinensis presence of a crista superior is a plesiomorphic feature, as it CAMPANA , 2004, p. 273 is lacking in more derived tetraodontiform otolith types. FROST, 1930, pl. 23, fig. 3 Balistes polylepis STEINDACHNER , 1876 Pl. 8, Figs. 10-11 BALISTOIDEI TRIACANTHOIDEA Balistes vetula LINNAEUS , 1758 TRIACANTHIDAE Pl. 8, Figs. 7-9 Available iconography: Balistoides viridescens (BLOCH & SCHNEIDER, 1801) Pseudotriacanthus strigilifer (CANTOR , 1849) Pl. 8, Fig. 12 Pl. 6, Fig. 4 Canthidermis maculatus (BLOCH , 1786) SANTINI & TYLER, 2002, fig. 5B, p. 328 Pl. 8, Fig. 13 TYLER, 1968,fig. 124G,p. 296 OHE, 1985, fig. Ad 448, p. 171 Triacanthus biaculeatus (BLOCH, 1786) SMALE et al. , 1995, pl. 147, fig. C1 Pl. 6, Fig. 6 Melichthys indicus RANDALL & KLAUSEWITZ, 1973 OHE, 1985, fig. 413, p. 173, as T. brevirostris SMALE eta!. , 1995, pl. 147, fig. D1 SA NTTNI & TYLER, 2002, fig. SA, p. 328 Melichthys niger (BLOCH, 1786) TYLER, 1968, fi g. 113G, p. 266 Pl. 7, Fig. 1 Triacanthus nieuhofi BLEEKER , 1852 Melichthys vidua (RICHARDSON, 1845) TYLER, 1968, fi g. 117G, p. 278 Pl. 7, Figs. 5-6 Tripodichthys angustifrons (HOLLARD, 1854) Odonus niger (ROPPELL, 1836) TYLER, 1968, fi g. L30G , p. 310 Pl. 7, Figs. 2-3 Tripodichthys b/ochii (BLEEKER, 1852) RlVATON & BOURRET , 1999, pl. 84, fi gs. 5- 6 T YLER, 1968, fi g. 2A, p. 8; fig. 128G, p. 304 S MALEet al. , 1995, p1.1 47, fi g.E1 ''

152 DfRK NOLF & JAMES C. TYLER

Pseudobalistes fuse us (BLOCH & SCHNEIDER, 1801) A/uterus scriptus (OSBECK, 1765) Pl. 8, Fig. 14 Pl. 10, Fig. 4 Rhinecanthus aculeatus (LrNNAEUS , 1758) RlVATON & BOURRET, 1999, pl. 83, figs. 18-19 ? SMALE et al., 1995, pl. 147, fig. F1: probably a Anacanthus barbatus GRAY, 1830 Sufflamen otolith Pl. 10, Figs. 7-8 RIVATON & BOURRET, 1999, pl. 84, figs. 7-8 Brachaluteres ulvarum JORDAN & FOWLER , 1902 (only utricular) 0HE, 1985, fig. 410, p. 172 Rhinecanthus rectangulus (BLOCH & SCHNEIDER, 1801) Cantherhines dumerilii (HOLLARD, 1853) Pl. 8, Figs. 15-16 Pl. 11, Fig. 1 Sufflamen bursa (BLOCH & SCHNEIDER , 1801) Pl. 9, Fig. 4 Cantherhines macrocerus (HOLLARD, 1853) Pl. 11, Fig. 2 Sufflamen chrysopterus (BLOCH & SCHNEIDER, 1801) Pl. 9, Figs. 1-2 Cantherhines pardalis (ROPPELL, 1837) SMALE et al. , 1995, pl. 147, fig. G1 SMALE et al. , 1995, pl. 147, fig. Jl Sufflamen fraenatus (LASTR1CELLE, 1804) Cantherhines sandwichiensis (QUOY & GAIMARD, 1824) Pl. 9, Fig. 3 Pl. 11, Fig. 3 RlVATON & BOURRET, 1999, pl. 84, figs. 1-4 Chaetodermis penicilligerus (CUYlER, 1816) SMALE et al., 1995, pl. 147, fig. H1 Pl. 10, Fig. 12 Sufflamen verres (GILBERT & STARKS, 1904) OHE, 1985, fig. Ad 449, p. 172 Pl. 9, Fig. 5 Meuschenia australis (DONOVAN, 1824) Xanthichthys lineopunctatus (HOLLARD, 1854) Pl. 10, Fig. 11 0HE, 1985, fig. 408, p. 171 NOLF, 1985, fig. 79C, p. 105, as Navodon sp. Xant/zic/zthys mento (JORDAN & GILBERT, 1882) Meuschenia scaber (FORSTER, 1801) Pl. 7, Figs. 7-8 Pl. 10, Figs. 9-10

Otoliths are unknown for the genera Abalistes and Monacanthus chinensis (OSBECK, 1765) Xenobalistes. Pl. 11, Figs. 6-9 All balistid otoliths are characterized by a strong salient ros­ Monacanthus tuckeri BEAN, 1906 trum. The posterior incision is only very weakly incised. Pl. 11, Fig. 4 Based upon otolith morphology, one can distinguish three Oxymonacanthus longirostris (BLOCH & SCHNEIDER, easily recognizable groups among balistids. The first group 1801) includes the genera Balistapus, Melichthys, Odonus and Pl. 11, Fig. 5 Xanthichthys. It is characterized by very thick otoliths with a strongly convex inner face, a very deeply incised sulcus, a Paraluteres prionurus (BLEEKER, 1851) salient rostrum and antirostrum, and a rudimentary posterior Pl. 10, Fig. 13 constriction. The second group includes Batistes, Balis­ Paramonacanthus oblongus (TEMMINCK & SCHLEGEL, toides, Canthidermis, Pseudobalistes and Rhinecanthus, 1850) which has a very peculiar type of otolith. It is composed of Pl. 12, Figs. 9-12 two parts: a main otolith body with a more-or-less typical Pervagor janthinosoma (BLEEKER, 1854) otolith morphology, and a posteroventral accretional body SMALE et-a!. , 1995, pl. 148, figs. Al-3 that is about as large as the main otolith body. An accretional Pervagormelanocephalus (BLEEKER, 1853) body of that size is unique among all teleosts. The third group Pl. 11 , Fig. 10 is composed of the genus Sufflamen. It is characterized by a thick otolith with a strong expansion of the posterodorsal Pervagor spilosoma (LAY & BENNETT, 1839) area, a very deeply incised sulcus that is well delimited by a PI.11 , Figs. ll-12 salient crista superior and inferior, and a strongly protruding Pseudomonacanthus macrurus (BLEEKER , 1857) rostrum and antirostrum. Pl. 12, Figs. 2-5 Stephanolepis cirrhifer (TEMMINCK & SCHLEGEL, 1850) OHE, 1985,fig.409, p. 172 MONACANTHJDAE Stephanolepis hispidus (LINNAEUS , 1766) Available iconography: Pl. 12, Fig. 1 A/uterus monoceros (LINNAEUS , 1758) Thamnaconusfajardoi SM1TH, 1953 Pl. 10, Figs. 5-6 SMALE eta!., 1995, pl. 148, fig. B 1 OH E, 1985, fig. Ad 450, p. 172 Thamnaconus hypargyreus (COPE, 1873) SMALE eta!., 1995, pl. 147, figs. 11-4 Pl. 12, Fig. 6 A/uterus schoepfi (W ALBAUM , 1792) RIVATON & BOURRET, 1999, pl. 83, figs. 11-17 PI.10, Figs. 1- 3 (sac., utr. , lag.) ''

Otolith evidence concerning interrelationships of caproid, zeifonn and tetraodontiform fishes 153

Thamnaconus modestoides (BARNARD, 1927) Lactophrys trigonus (LLNNAEUS, 1758) SMALE et al., 1995, pl. 148, fig. C1 Pl. 14, Fig. 3 Thamnaconus modestus (GONTHER, 1877) Lactoriafomasini (BIANCONI, 1846) Pl. 12, Figs. 7-8 Pl. 14, Fig. 4 OHE, 1985, fig. 411, p. 172, as Navodon modest us SMALE et al., 1995, pl. 148, fig. D 1 Ostracion cubicus LlNNAEUS, 1758 Otolith morphology indicates that monacanthids are a rather Pl. 14, Figs. 5-6 homogeneous group. Monacanthid otoliths are characterized RIVATON & BOURRET, 1999, pl. 84, fig. 9 by a very strong antero-posterior constriction. The upper por­ SMALE et al. , 1995, pl. 148, fig. E1 tion of the otolith is always much narrower than the lower Ostracion meleagris SHAW, 1796 portion and often shows a well-marked posterodorsal angle. Pl. 14, Figs. 8-9 In many species, the lower portion of the otolith is about NOLF, 1985, fig. 79D, p. 105, as Ostracion twice as large as the upper portion, and sometimes it devel­ lentiginosum ops anterior, posterior and ventral expansions. Therefore, in several taxa, the anterior and posterior halves of the otolith Rhinesomus concatenatus (BLOCH, 1785) look almost symmetrical, which, for isolated otoliths, makes OHE, 1985, fig.419, p. 174 it sometimes difficult to judge if a left or a right one is con­ Tetrosomus gibbosus (LrNNAEUS, 1758) cerned. In such cases, the posterodorsal angle, if sufficiently Pl. 14, Fig. 7 developed, is often the best criterion to judge. The crista su­ FROST, 1930, pl. 23 , fig. 4, as Ostracion perior is lacking or obsolete in all the examined taxa. gibbosus

Otoliths are unknown for the genera Lactoria and Rhynchos­ 0STRACIOIDEA tracion. [SPINACANTHIDAE, extinct; middle Eocene, Lutetian; Ostraciid otoliths are very similar to those of aracanids. The Monte Bolca, Italy; otoliths unknown] only observable difference is the crista superior, which is ab­ [PROTOBALISTIDAE, extinct; middle Eocene, sent or tends to become obsolete in many of the examined Lutetian; Monte Bolca, Italy; otoliths unknown] ostraciids. In Acanthostracion, the otoliths are extremely thick, but all other examined species have a normal thick­ ness. For the rest, we judge the available material insufficient ARACANIDAE for any further conclusions. Available iconography:

Anoplocapros inermis (FRASER-BRUNNER, 1935) TETRAODONTOIDEI Pl. 13, Figs. 4-5 [EOPLECTOIDEA, extinct] Anoplocapros robustus (FRASER-BRUNNER, 1941) [EOPLECTIDAE, extinct; middle Eocene, Lutetian; Pl. 13 , Fig. 3 Monte Bolca, Italy; otoliths unknown] Aracana ornata (GRAY, 1838) Pl. 13, Fig. 6 TRIODONTOIDEA TRIODONTIDAE Kentrocapros aculeatus (HOUTTUYN, 1782) Available iconography: Pl. 13 , Fig. 7 OH ~ 1985,fig. 412, p. 173 Triodon macropterus LESSON , 1829 Pl. 13, Figs. 1-2 Otoliths are unknown for the genera Caprichthys, RIVATON & BOURRET, 1999, pl. 84, figs. 14-15 Capropygia, Polyplacapros and Strophiurichthys. Otoliths of the four taxa of aracanids that we examined are The otoliths of the single extant species of this family are characterized by a globally rounded outline, but with a very characterized by a high dorsal portion, a strong antero-poste­ strong antero-posterior constriction. The sulcus is rather rior constriction along the midline, and a ventral portion with strongly incised, and both the crista inferior and superior are a long prominent rostrum, a moderate posterior expansion, clearly developed. The presence of a well-developed crista and an extremely reduced ventral area, which makes up a superior can be considered a plesiomorphic feature relative concave ventral rim. to ostraciid otoliths, in which this crista often tends to be­ come obsolete. In most of the examined aracanid otoliths, the TETRAODONTOIDEA dorsal portion is manifestly narrower than the ventral por­ TETRAODONTIDAE tion. Available iconography: Amblyrhynchotes honkenii (BLOCH, 1795) Pl. 15 , Fig. 7, redrawn after SMALE et al., 1995 0STRACHDAE SMALE et al., 1995, pl. 148 , fi gs. FJ - 5 (series of Available iconography: variability) Acanthostracion quadricomis (LrNNAEUS, 1758) Arothron hispidus (LrNNAEUS, 1758) Pl. 14, Figs. 1-2 Pl. 15 , Fig. 1 I I

154 DlRK NOLF & JAMES C. TYLER

RIVATON & BOURRET, 1999, pl. 84, figs. 16-17 NOLF, 1985, fig. 79E, p. 105 SMALE et al. , 1995, pl. 148, fig. G 1 Takifuguniphobles (JORDAN & SNYDER, 1901 ) Arothron immaculatus (BLOCH & SCHNEIDER, 1801) OHE, 1985, fig. 421, p. 174 Pl. 15, Fig. 2 Takifugu oblongus (BLOCH, 1786) Arothronmappa (LESSON, 1831) PI.17, Figs.1-5 Pl. 15, Fig. 3 Takifugu porphyreus (TEMMINCK & SCHLEGEL, 1850) Arothron stellatus (BLOCH & SCH NEIDER, 1801) Pl. 17. Fig. 6 SMALE et al., 1995, pl. 149, fig. AI Takifugu rubripes (TEMMINCK & SCHLEGEL, 1850) Canthigaster amboinensis (BLEEKER, 1865) OHE, 1985, fig.420,p. 174 Pl. 16, Fig. 3 Takifugu xanthopterus (TEMMLNCK & SCHLEGEL, 1850) Canthigaster coronata (VAILLANT & SAUVAGE, 1875) Pl. 17, Figs. 7-8 Pl. 16, Fig. 2 Tetraodonfluviatilis HAMILTON, 1822 Canthigaster jactator (JENKJNS, 190 I) Pl. 17, Fig. 11 Pl. 16, Fig. 1 Canthigaster margaritata (ROPPELL, 1829) Tetraodonlineatus LINNAEUS, 1758 Pl. 16, Fig. 6 FROST, 1930, pl. 23, fig. 5, as Tetrodon fahaka Canthigaster punctatissimus (GUNTHER, 1870) Tetraodon palembangensis BLEEKER, 1852 Pl. 16, Fig. 4 Pl. 17, Fig. 9 Canthigaster rivulatus (TEMMINCK & SCHLEGEL, 1850) Tylerius spinosissimus (REGAN, 1908) Pl. 16, Fig. 5 Pl. 17, Fig. 10 0HE, 1985, fig. 418, p. 174 Otoliths are unknown for the genera Carinotetraodon, Canthigaster solandri (RICHARDSON, 1845) Colomesus, Feroxodon, Javichthys and Omegophora. Pl. 16, Fig. 7 Otoliths of tetraodontids show a considerable diversity. The SMALE et al., 1995, pl. 149, fig. B 1 crista superior is always lacking or obsolete. The central part Canthigaster valentini (BLEEKER, 1853) of the crista inferior is always very salient. Although only a RIVATON & BOURRET, 1999, pl. 84, fig. 13 limited number of specimens was examined (and in some Contusus richei (FREMINVILLE, 1813) cases, the various specimens of the species examined showed Pl. 15, Figs. 10- 12 considerable differences in size), some characteristic clusters could be determined. The outlines of the otoliths of Lagocephalus ?guentheri RIBEIRO, 1915 Amblyrhynchotes, Contusus, Sphoeroides and Takifugu are SMALE et al., 1995, pl. 149, figs. Cl-2 (growth similar to those of monacanthid otoliths, but they differ from series) monacanthids by having an extremely salient crista inferior. Lagocephalus inermis (SCHLEGEL, 1850) The outline and the very salient rostrum of Lagocephalus re­ SMALE et al., 1995, pl. 149, figs. 01- 2 semble that of Triodon. In Tetraodon, the crista inferior and Lagocephalus lunaris (BLOCH & SCHNEIDER, 1801) associated upper part of the ventral area becomes extremely Pl. 15, Figs. 8-9 distended, and the ventral part of the sulcus is deeply incised, OHE, 1985, fig. 416, p. 174 which has similarities to the morphology of the funnel-like Lagocephalus scleratus (GMELIN, 1789) sulcus of diodontids. A straight forward, recognizable mor­ Pl. 15, Figs. 4-6 phology is also present in Canthigaster. It has a strong RIVATON & BOURRET, 1999, pl. 84, figs. 18- 22 antero-posterior constriction, a very narrow and angular dor­ SMALE et al., 1995, pl. 149, figs. E 1- 3 sal portion, a widely extended ventral portion with a very sa­ lient rostrum and a round, thickened posterior end. Arothron Liosaccus cutaneus (GU nther, 1870) is only documented by very small otoliths, but these exhibit OHE, 1985, fig. 417, p. 174 some similarity to those of Canthigaster. Sphoeroides lobatus (STEINDACHNER, 1870) Pl. 16, Fig. 8 DIODONTIDAE Sphoeroides maculatus (BLOCH & SCHNEIDER, 1801) Available iconography: CAMPANA, 2004, p. 275 Allomycterus whitleyi PHILLIPPS, 1932 Sphoeroides nephelus (GOODE & BEAN, 1882) Pl. 18, Fig. I Pl. 16, Figs. 10- 11 Chilomycterus affinis GONTHER, 1870 Sphoeroides pachygaster (MULLER & TROSCHEL, 1848) Pl. 18, Fig. 3 SMALE et al. , 1995, pl. 149, figs. F 1- 3 Chilomycterus schoepfii (W ALBAU M, 1792) Sphoeroides spengleri (BLOCH, 1785) Pl. 18, Fig. 2 Pl. 16, Fig. 9 CAMPANA, 2004, pl. on p. 274 Sphoeroides testudineus (LINNAEUS, 1758) Cyclichthys orbicularis (BLOCH, 1785) PJ.16, Fig. 12 Pl. 18, Fig. 4 I I

Otolith evidence concerning interrelationships of caproid , zeiform and tetraodontiform fishes 155

Dicotylichthys punctulatus KAUP, 1855 KOTLYAR & PARJN, 1990, fig. 5, p. 110 Pl. 18, Fig. 5 0HE, 1985, fig. 110, p. 74 Diodon holocanthus LI NNAEUS , 1758 RI VATON & BOURRET, 1999, pl. 145, figs. 1-4 Pl. 18, Figs. 8-9 SCHMIDT, 1968, pl. 5, fig. 60; pl. 18 SCHWA RZHANS, 1980, fig. 376, p. 114 Diodon hysh·ix LrNNA EUS , 1758 STEURBAUT, 1979, pl. 6, figs. 12-1 3 Pl. 18, Fig. 6 STI NTON, 1978, fig. 26g, p. 136 RIVATON & BOURRET, 1999, pl. 84, figs. 10-12 SMALE et al., 1995, pl. 149, figs. G 1-3 Antigonia eos GILB ER7, 1905 NOLF, 1993, fig. 3N, p. 228 Diodon nicthemerus C UYlER, 18 18 Pl. 18 , Fig. 7 Antigonia malayana WEBER , 191 3 Fig. 7 Otoliths are unknown for the genera Lophodiodon and RJ VATON & BO URRET, 1999, pl. 145, figs. 5-9 Tragulichthys. Antigonia rubescens (GUNTHER, 1860) Diodontid otoliths, with their crystal-like salient needles and RI YATON & BO UR RET, 1999, pl. 145, figs. 10-15 their horn-like appearance, are among the strangest otoliths SMALE et al., 1995, pl. 47, figs. A1-3 known among fi shes. Their morphology can only be under­ stood if one considers them to be highly modified derivatives of the type observed in tetraodontids, such as Tetraodon. In CAPROIDAE, Caproinae diodontid otoliths, the salient central part of the crista inferior Capros aper LINNAEUS , 1758 is extremely protruding, and the ventral part of the sulcus Fig. 6 sinks deepl y, which together form a funnel-like structure. In ASS IS , 2004, fig. 57, p. 119 Diodon holocanthus, the homology with Tetraodon otoliths CHAINE, 1958, pl. p. 236 is still visible. Only by comparison with the transi ti onal oto­ FROST, 1927, pl. 8, figs. 11-13 (sac., lag., utr.) lith morphologies from Tetraodon to Diodon can the struc­ NOLF, 1985, fig. 55F, p. 73 ture of the otolith in Allomycterus be understood. NOLF, 1993, fig. 3, 0 , p. 228 NOLF & LAPIERRE, 1979, pl. 3, fig. 2 MOLOIDEA STI NTON, 1978, fig. 27d, p. 145 MOLIDAE Available iconography: Otolith-based fossil record of caproid, zeiform and Mola mola (LINNAEUS, 1758) tetraodontiform fishes NOLF, 1985, fig. 79F, p. 105 Ranzania laevis (PENNANT, 1776) Pl. 18, Fig. 10 CAPROIDAE - Antigonia angusta STLNTON & NOLF, 1970: middle In the labyrinth of Mola mola, a fish with very poor ossifica­ Eocene, Luteti an; Belgium. ti on of the skeleton, onl y some chalk dust has been collected - Antigonia capros LOWE, 1843: earl y Miocene, (figured by NOLF, 1985), which is not amenable to morpho­ Burdigalian; Aquitaine, France (NOLF & BRZOBOHATY, logical interpretation. For Ranzania laevis, a single very 2002). fragile small otolith is available, which shows some spine­ - Capros parvus (MENZEL, 1986): middle Miocene; Ger­ like crystal extrusions, as in diodontids. On the basis of many. homologies with other tetraodontiform otoliths, the small - "genus Capmidarum" exiguus (STINTON, 1978): middle Ranzania otolith can be interpreted as having an elongate Eocene, Luteti an; southern England. ventral portion, a strong but structureless cri sta inferior ac­ -"genus Caproidarum" sonodae NOLF & LAPIERRE, 1979): centuated by a deep ventral portion of the sul cus, and a very middle Eocene, Lutetian; Paris Basin . narrow, strongly salient dorsal area. We presume that the spine-like crystal extrusions of the Ranzania otolith are re­ Zeiformes incertae sedis lated to the crystalline needles of diodontids. - "genus Zeiformorum" janni SCHWARZHANS, 2003: Mid­ dle Paleocene, Selandian; Denmark - "genus Zeiformorum" tyleri NOLF, 2003: late Cretaceous, Examined out-group taxa Santonian, Pyrenees, France (Pl. 4, Figs. 11 -12). - "Bavarian zeifom1s" ("Isozen" sp. and "Isozen beatae," PERCIFORMES invalid names, see NOLF, 2003): Paleocene (Pl. 4, Fig. 5) and CAPROIDAE Maastrichtian (Pl. 4, Fig. 4); Bavaria (SCHWARZHANS, 1996). CAPROIDAE, Antigoniinae Available iconography: ZENIONTIDAE Antigonia capros LOWE, 1843 - Zenion hololepis GOODE & BEAN, 1896: early Pliocene, Fig. 6 Zancli an, SE France (NOLF & CAPPEITA , 1989, pl. 14, figs. H UYG HEBAERT & NOLF, 1979, pl. 4, figs. 11 - 12 13-14). 'I 156 DIRK NOLF & JAMES C. TYLER

- "genus aff. Zenion" sp. n. : early Eocene, Ypresian (Gan otolith apparently evolved independently in several taxo­ Clay) (Pl. 4, Figs. 2-3) and middle Eocene, Lutetian nomic groups that are not closely related. Tllis feature is ob­ (Miretrain marls); Aquitaine, France. served mainly in fishes with a mesopelagic or benthopelagic - Zenion sp.: early Oligocene, Rupelian; Piemonte, Italy habitat, and it must be considered a homoplasy. See Nolf, (NOLF & STEURBAUT, 2004, pl. 9, fig. 2: an eroded Zenion 1993, fig. 3, p. 228, for illustrations of this type of otolith in otolith, not identifiable at species level). myctophids, neoscopelids, five families of beryciforms and "genus Zeniontidarum" sulcifer (STINTON, 1966): early caproids. In Fig. 5, Polymixia and Sargocentron represent the Eocene, Ypresian, London Clay; southern England (Pl. 4, plesiomorphic condition, and Centroberyx affinis and BeJ) IX Figs. 9-10). represent the homoplasic apomorphic conditions. The Santonian (Upper Cretaceous) Centroberyx antiquus exhib­ its a more plesiomorphic level than in berycoid otoliths: this TRIACANTIDDAE species already has the round, rather deep-bodied form and - Triacanthus sp.: early Eocene, Ypresian (London Clay) the wide ostium, but it still has a narrow cauda, as in (Pl. 6, Fig. 7) (Coli. Rundle, new fossil record). perciforms, holocentrids and Polymixia. Both by its ancient age and morphology, the otolith of Centroberyx indicates a more plesiomorphic condition than in Recent berycids, and it 0STRACIIDAE helps to determine the polarity of otolith features. - Ostracion pergravis SCHWARZHANS, 2003: Paleocene, Selandien ; Denmark (Pl. 14, Fig. 10-11). 3. Caproid otoliths and the "berycid" aspect of those of Antigonia. Discussion Fig. 6 shows the plesiomorphic versus derived morphology Characterization of otoliths 111 caproid, zeiform and for, respectively, berycoids, caproids and percoids. In tetraodontiform fishes berycoids, the plesiomorphic condition is demonstrated by the holocentrid Sargocentron, and the derived condition is 1. Plesiomorphic otoliths and the plesiomorphic represented by Beryx decadactylus. Percoids, such as configuration of generalized perciform otoliths. Morone, demonstrate the plesiomorphic condition, which is very similar to that of Sargocen.tron, whereas the derived Before proceeding with the morphological analysis of condition is exhibited by Kurtus or Apogon. Both the zeiform and tetraodontiform otoliths, it is important to bear plesiomorphic and derived conditions also exist in caproids, in mind the configuration of plesiomorphic otoliths of a family that has been included in the Zeiformes in many acanthopterygian fishes in general, and especially of those of classifications, and STINTON (1967) even suggested that generalized perciforms. In Fig. 4, various types of Antigonia is a berycoid on the basis of its otolith morphol­ acanthopterygian otoliths (holocentids, atherinids, mugilids, ogy. Considering the deep-bodied otoliths with a straight sul­ polynemids) are compared with a number of other taxa, in­ cus as a homoplasic feature that has evolved independently cluding primitive teleosts, such as osteoglossids and elopids, in many groups, our interpretation is that in caproids the and selected euteleosteans, such as salmonids, aulopids, plesiomorphic sulcus condition is found in Capros, whereas synodontids and polymixiids. A first conclusion to be drawn the derived "berycid"-like otolith is found in Antigonia. The from this comparison is that generalized acanthopterygian antero-posterior constriction seen in Capros, however, is a (and percoid) otoliths exhibit a Bauplan that can be traced synapomorphy shared with zeiforms and tetraodontiforms. back as far as the elopids; even osteoglossids are not incom­ patible as an ancestral percoid otolith type. At the level of synodontids, it is even difficult to define, for that group, oto­ 4. Characterization of caproid, zeiform and lith features that are fundamentally different from those of tetraodontiform otoliths. percoids. If the series from Fig. 4 were presented in a cladogram, with osteoglossids as the most plesiomorphic In Capros, the otolith is deep-bodied, but the sulcus pattern is group and the acanthopterygians as the most apomorphic, the more like that of Sargocentron and Morone (Fig. 6), with a cladogram probably would meet with general agreement short ostium and with a long cauda that is slightly. bent among fish phylogenists, but it also would contain extremely ventrally in its posteri or portion. The otolith of Capros is in­ important lacunae. teresting in the sense that it already has a feature that can be taken as a synapomorphy for zeiform and tetraodontiform otoliths, which consists of a marked antero-posterior con­ 2. Polarity of the sulcus morphology in holocentrid and striction along the horizontal nlidline of the otolith. berycid otoliths. Importance of the Santonian Both zeiform and tetraodontiform otoliths belong to the Centroberyx antiquus. deep-bodied otolith type, but because this feature appears in­ dependently many times in the otoli ths of unrelated groups of Round and deep-bodied otoliths with a broad sulcus consist­ fishes, it cannot be utilized as a synapomorphy. ing of a rather long osti um and a straight, relatively wide Zeiform and tetraodontiform otoliths, however, are united by cauda, are considered to be derived; however, this type of a synapomorphy that consists of a swelli ng of the upper part I I

Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fishes 157

of the ventral area, just below the caudal crista inferior. This subfamily also constitutes a very homogeneous cluster, dif­ feature is lacking in caproid otoliths. Because in all taxa of fering from cyttids only by their less upward-protruding zeiforms and tetraodontiforms the cauda is markedly longer posterodorsal angle. Based only upon otolith morphology, than the ostium, this swelling occupies a central position. one would tend to group these two families closely together and also to relate the very homogeneous cluster of oreosomatids to them, as an apomorphic group, character­ 5. Polarity in zeiform otoliths. ized by a ventra] expansion of the anterior and posterior end of the sulcus and by hypertrophy of the collicula, but other­ In Fig. 7, a series ofberyciform (Monocentris and Diretmus), wise showing a very similar otolith type. caproid (Antigonia) and zeiform (Cyttus and Zeus) otoliths The parazenids clearly split into two clusters: Parazen, with are compared. The zeiforms Cyttus and Zeus represent, re­ a very plesiomorphic otolith type, and the Cyttopsinae spectively, a plesiomorphic and the most derived otolith (Cyttopsis + Stethopristes), which, based only upon otolith types among zeiforms. Both of these zeiforms have the con­ morphology, can be integrated perfectly in the cyttid­ striction along the midline and the swollen upper centra] part grarnmicolepidid-oreosomatid group. of the ventral area. Except for the constriction, the general In zeniontids, the situation is even more complicated. outline of the otolith in Cyttus still presents a berycid-like Zenion exhibits a plesiomorphic otolith type, whereas the appearance. In Zeus, the lower portion of the otolith has [Capromim.us + Cyttomimus] cluster can be defined as an strongly modified anterior and posterior lobiform expan­ apomorphic relative (characterized by reduction of the sions, but the upper portion remains comparable to that of collicula toward the anterior and posterior marginal zones) of Cyttus, and it still shows the development of a strong the cyttid-grammicolepidid-oreosomatid group. The situa­ posterodorsal angle, as in Cyttus and Diretmus. tion becomes even more complex because the otolith of Cyttomimus could function perfectly as a plesiomorphic sis­ ter group for the zeid family. The protruding collicula, the 6. Polarity in tetraodontiform otoliths. dorsal area and the swollen upper central part of the ventral area are almost identical in both groups. Moreover, the ante­ In Fig. 8, the otoliths from two taxa of the most phylo­ rior and posterior constrictions in the ventral margin of .genetically basal family (Triacanthodidae) of extant Cyttomimus strongly appear to be precursors of the ventral tetraodontiforms, and of a representative of each of three lobe-like expansions of zeid otoliths. more derived families (Monacanthidae, Tetraodontidae, In zeids, the otoliths of Zeus and Zenopsis, based upon their Diodontidae) of that order are compared with the otolith of a tri-lobed shape, comprise an unambiguous homogeneous caproid; all have a reasonable similarity.. The tetraodonti­ group, which is incontestably the most apomorphic zeiform forms, however, have a derived condition: the swollen upper clade. central part of the ventral area. Monacanthids, diodontids The most problematic cases are clearly the plesiomorphic and tetraodontids have even more strongly derived otoliths: otoliths of the genera Parazen and Zenion. Both of these gen­ monacanthids such as Chaetodermis and Pervagor exhibit an era were separately and relatively deeply nested in the osteol­ extreme antero-posterior otolith constriction; diodontids and ogy-based phylogeny of zeiforms by TYLER et al. (2003). tetraodontids have a funnel-like sulcus. Among other SCHWARZHANS (1996), using otolith information, proposed a tetraodontiform otoliths, very highly derived types can be cladogram in which he grouped Parazen and Zenion as the found, showing essentially autapomorphic features, which plesiomorphic sister group of all other zeiforms. Johnson and are discussed in the next section. PATTERSON (1993) previously had suggested that Zenion and Parazen might be the more morphologically primitive zeiform taxa; the otolith dat.a presented herein are in agree­ Phylogenetic evaluation of otolith features of zeiform ment with that proposal. and tetraodontiform fishes The phylogenetic placement of Parazen as sister to [Cyttopsis + Stethopristis] in parazenids and Zenion as sister In the following section, otolith-based data are compared to [Capromimus + Cytomimus] in zeniontids, however, is with osteological data from osteologically based logically consistent within both families. A plausible expla­ phylogenetic studies of zeiform (TYLER et al., 2003) (Fig. 9) nation is that in both of these families the genera with a and tetraodontiform (TYLER, 1980, and SANTINI & TYLER , plesiomorphic otolith type represent independent retention 2003) (Fig. 10) fishes. of the ancestral condition: they may be surviving taxa with In zeiforms, the otolith data seem to support reasonably well the plesiomorphic otolith type, whereas in other zeiform the homogeneity of the six defined families, but they are of families, such forms are extinct, or not yet discovered. We little use in interpreting the interrelationships among these note that TYLER and SANTINI (2005) analyzed all of the fossil families. and extant zeiform and zeomorph-like fishes (dating to the In cyttids, the otoliths suggest a very homogeneous cluster, Upper Cretaceous) using a large osteological data set. That with all species showing deep-bodied otoliths with a strong study also called into question several aspects of the posterodorsal angle, a strong antero-posterior constriction, monophyly and inclusiveness of the Parazenidae and and a hollow sulcus with no, or only rudimentary, collicular Zeniontidae. Additional study is necessary to resolve these formations. A similar otolith type is found in issues. grammicolepidids, among which the grammicolepidine In teb·aodontiforms, the otoliths exhibit a broad scope of ''

158 DIRK NOLF & JAMES C. TYLER very different morphologies. The most characteristic types whereas Tetraodon and Canthigaster represent two different are herein (Fig. 10) placed beside the phylogenetic hypoth­ groups that are easily distinguished by their otoliths. esis of SANT!Nl & TYLER (2003) for ease of comparison. Diodontid otoliths, with their deep funnel-like sulcus and All tetraodontiform otoliths have, at least in some measure, their salient crystal needles, represent an extremely derived an antero-posterior constriction along the midline of the apomorphic otolith type that is easy to recognize but is not otoliths, a synapomorphy shared with Capros and with easy to interpret. The diodontid cqndition can be best under­ zeiforms. stood by comparison with otoliths of the tetraodontid genus The position of triacanthodids and triacanthids as the most Tetraodon, which can be considered to be their plesiomorphic families of tetraodontiforms is confirmed by plesiomorphic counterpart. This confirms their position in their otolith morphology. Triacanthodid and triacanthid the cladogram, where tetraodontids and diodontids appear as, otoliths exhibit a round, deep body with a sulcus delimited by respectively, plesiomorphic and apomorphic sister groups. well-structured cristae, which are also found in many Molids are known only by the chalky dust in the Mala laby­ beryciforms and caproids. rinth and by a single Ranzania otolith; thus, one can only In balistids, three different clusters can easily be recognized conunent that it is highly modified but compatible with by otolith morphology (see descriptive section): (1) tetraodontiform otolith morphology, with similarities to Balistapus, Melichthys, Odonus and Xanthichthys; (2) diodontids in their crystalline needle-like extrusions. Batistes, Balistoides, Can.thidermis, Pseudobalistes and Rhinecanthus; and (3) Sufflamen. Among these, the "Balistapus group" apparently is the most plesiomorphic be­ Otolith-based evaluation of the affinities of the zeiform­ cause of their regular outline and their well-marked cristae; tetraodontiform group to other fish taxa however, the extreme thickening of their otoliths is an apomorphic condition. The other two groups have only very As previously stated under the heading "Characterization of marked autapomorphic features. caproid, zeiform and tetraodontiform otoliths," caproids (and We have more confidence in the phylogenetic validity of especially Capros) appear to be the best candidate for a these three groups than in the groups hypothesized with un­ plesiomorphic sister group of the zeiforms and certainty by TYLER (1980, p. 121), who stated that the ge­ tetraodontiforms, with which they share the synapomorphy neric relationships of balistids were not especially clear on of antero-posterior constriction of the otoliths. In addition to the basis of his osteological analysis. The otolith data also this, both groups share various similarities in their overall conflicts with the composition of the three groups into which appearance, such as the very high bodies of their otoliths, and MATSUURA (1979) divided the balistids. the often strongly developed posterodorsal angle, but such Monacanthid otoliths seem to indicate that this is a very ho­ features cannot be considered to be synapomorphies. mogeneous group. For example, otolith morphologies like Various other groups have also been suggested as relatives of that of Anacanthus can be considered to be plesiomorphic, zeiforms or tetraodontiforms. For example, HOLT (1894), whereas extremely slender otoliths, such as in Chaetodermis THILO (190 1) and BOULENGER (1 902) suggested that flat­ (Pl. 10, Fig. 12) and Stephanolepis (Pl. 12, Fig. 1), are con­ fishes were related to zeiforms. NORMAN (1 934) rejected sidered to be the most derived. The available otolith material such a relationship, and furthermore the otolith morphology does not allow for a detailed analysis that would lead to a does not provide any evidence for such a relationship. clustering of taxa within this large family (about 107 spe­ REGAN (1 910) proposed a close relationship between cies). zeiforms (his Zeidae) and caproids (his Caproidae), recog­ Similarities among aracanid and ostraciid otoliths indicate nized them as the order Zeomorphi, and related them to these taxa form a homogeneous group, but the material avail­ berycoids. PATTERSON (1968) regarded zeiforms (including able to us does not allow for a detailed analysis. The two caproids) as the apomorphic sister group to berycoids (in­ families may be distinguished, however, by the presence of a cluding stephanoberyciforms). WINTERBOTTOM (1974) sug­ well-developed crista superior in all of the examined gested a possible zeiform-tetraodontiform relationship. aracanids. This feature is considered to be plesiomorphic ROSEN (1984) provided evidence for a group comprising relative to ostraciids, in which the crista tends to become ob­ caproids, tetraodontiforms and zeiforms, with the latter two solete. being sister groups. JOHNSON AND PATTERSON (1993) sug­ The single extant species of triodontid is characterized only gested that zeiforms (excluding caproids) are the sister group by autapomorphic otolith features. One should note, how­ of berycifo rms (excluding stephanoberyciforms) plus ever, the considerable similarity in outline of Triodon otoliths percomorphs. and those of tetraodontids, such as Lagocephalus (Pl. 15, The recognition of caproids as the plesiomorphic sister group Figs. 4-6 and Figs. 8- 9). The otolith similarities agree with of zeiforms + tetraodontiforms is supported by their otolith their placement in the cladogram. morphology. Tett·aodontid otoliths, although united by a synapomorphy Concerning tetraodontifonns, the perciform acanthuroid (the presence of a very salient crista inferior) with fishes often have been considered to be likely relatives of diodontids, exhibit considerable diversity. On the basis of the tetraodontifonn s (e.g., WINTERBOTTOM, 1974, TYLER, 1968 li mited number of species examined (the family has about and 1980). Because there is no single reference for 185 species), at least three clusters can be recognized: rela­ acanthuroid otoliths, we provide figures of some in Pl. 19 and tively plesiomorphic tetraodontid otoli ths are found in Pl. 20. A glance at these plates indicates that acanthuroids Amblyrhynchotes, Contusus, Sphoeroides and Tak(fugu, have a rather conservati ve, plesiomorphic percoid otolith ''

Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fishes 159 type; compare these, for example, with Marone in Fig. 6. Literature Cited This is especially true for the genera Acanthurus, Ctenochaetus, Zebrasoma and Paracanthurus. The ASS IS, C.A., 2003. The lagenar otoliths of teleosts: their morphol­ acanthurid genus Naso (a phylogenetically basal lineage) and ogy and its application in spec ies identification, phylogeny and sys­ the siganids have a more specialized otolith type, but basi ­ tematics. Journal of Fish Biology, 2003 (3): 1268-1295. call y such otoliths remain of the percoid type. The ASS IS , C. A. , 2004. Guia para a identifica'Parazen pacificus: oste­ drafts of thi s manuscript; John Lundberg and Mark Sabaj for ology, systematics, and di stribution (Parazenidae, Zeiformes). Jour­ help in examining specimens at the Academy of Natural Sci­ nal of Ichthyology, 41 (9): 687-697. ences of Philadelphia; James DiLoreto, Smithsonian Institu­ KOTLYAR , A.N. & PARI N, N.Y., 1990. Otoliths and age of bottom tion, for help with transrrlitting and printing electronic illus­ and near bottom fishes of the Nazca and Sal a y Comez submarine trations and Anne Wauters (IRSNB), who helped in prepar­ rid ges. Akademia Nauk SSSR, Trudy In stitut Okeana!., 125:97-126. ing the drawings .. (i n Ru ss ian). The first author thanks the Smithsonian Institution, Washing­ MATSUUR A, K. , 1979. Phylogeny of the superfami ly Ba li sto idea ton , D.C., for two short-term visitor grants (in 1997 and (Pisces: Tetraodontiformes). Memoirs of the Faculry of Fisheries, 2000), during which stays the present study was initiated. Hokkaido Un iversit)• , 26 (1-2): 49-169. II

160 DIRK NOLF & JAMES C. TYLER

NELSON, J.S., 2006. Fishes of the world, Fourth edition. John Wiley SCHMIDT, W. , 1968. Vergleichend morphologische Studie tiber die & Sons, USA, 601 pp. Otolithen mariner Knochenfische. Archiv fiir Fischereiwissen­ NOLF, D., 1985. Otolithi Piscium. In: H.P. Schultze (ed.). Handbook schaft, 19 (!): 1-96. of Paleoichthyology, 10. Fischer, Stuttgart and New York, pp. 1- SCHWARZHANS, W. , 1980. Die tertiare Teleosteer-Fauna Neus­ 145. leeands, rekonstruiert anhand von Otolithen. Berliner geowissen­ NOLF, D., 1993. A survey of perciform otoliths and their interest for schaftliche Abhandlungen, A, 26: 1-2! 1. phylogenetic analysis, with an iconographical synopsis of the SCHWARZHA NS, W., 1996. Otoliths fro m the Maastrichtian of Ba­ Percoidei. Bulletin of Marine Science, 52 ( I): 220-239. varia and their evolutionary significance. In Aratia, G. & Viohl, G. NOLF, D., 2003. Fish otoliths from the Santonian of the Pyrenean (editors): Mesozoic Fishes- Systematics & Paleoecology, pp. 433- realm, and an overview of all otolith-documented North Atlantic 459. Verlag Pfeil, Mlinchen. Late Cretaceous teleosts. Bulletin de l'Institut royal des Sciences SMALE, M.J., WATSON, G. & HECHT, T. , 1995. Otolith atlas of naturel/es de Belgique, Sciences de La Terre, 73 : 155-173. South African marine fishes. Ichthyological Monographs, l: 1-253. NOLF, D. & CAPPETI'A, H., 1989. Otolithes de poissons pliocenes du STEURBAUT, E. ,I979. Les otolithes de Teleosteens des Marnes de Sud-Est de Ia France. Bulletin de 1'/nstitlll royal des Sciences Saubrigues (Miocene d' Aquitai ne meridionale, France). naturelles de Belgique, Sciences de Ia Terre, 58 (1988): 209-271. Palaeontographica, A, 166: 50-91 . NOLF, D. & LAPIERRE, H. 1979. Otolithes de poissons nouveaux ou STINTON, F.C. , 1967. The otoliths of the teleostean fish Amigonia peu conn us du Calcaire Grassier et de Ia Formation d' Auvers capros and their taxonomic significance. Bocagiana, Museu Mu­ (Eocene du Bassin parisien). Bulletin du Museum National nicipal do Funchal, 8: l-7. d 'Histoire Naturelle, Paris, 4, I, C, 2: 79-125. STINTON, F.C., 1978. Fish otoliths from the English Eocene, III. NOLF, D. & BRZOBOHATY, R., 2002 . Otolithes de poissons du Palaeontographical Society Monographs, 555: 127-189. Pah~ocanyon de Saubrigues (Chattien a Langhien), Aquitaine THILO, 0., 190 1. Die Vorfahren der Schollen. Bulletin de meridionale, France. Revue de Micropaleontologie, 45 (4 ): 261- l'Academie Imperiale des Sciences de St. Petersbourg, series 5, 14 296. (3): 3 15-350. NOLF, D. & STEURBAUT, E. , 2004. Otolithes de poissons de TYLER, J.C., 1968. A monograph on plectognath fishes of the !'Oligocene inferieur du Bassin liguro-piemontais oriental, ltalie. superfamily Triacanthoidea. The Academy of Natural Sciences of Rivista Piemontese di Storia naturale, 25 : 21-68. Philadelphia, Monograph, 16:1-364. NORMAN , J.R., 1934. A systematic monograph of the flatfishes TYLER, J.C., 1980. Osteology, phylogeny and higher classification (Heterosomata), vol. I: Psettodidae, Bothidae, Pleuronectidae. The of the fishes of the order Plectognathi (Tetraodontiformes). NOAA Trustees of the British Museum, London, 459 pp. Technical Report, National Marine Fisheries Circul(/1; 434: l-422. OHE, F. , 1985. Marine fish-otoliths of Japan. Special Volume ofBul­ TYLER, J.C. , O'TOOLE, B. & WI NTERBOTTOM , R., 2003. Phylogeny letin (Earth -Science), The Senior High School Attached to the Aichi of the genera and families of Zeiform fishes, with comments on University of Education, 184 pp. their relationships with tetraodontiforms and caproids. Smithsonian PATTERSON, C., 1968. The caudal skeleton in Mesozoic Contributions to Zoology, 618: 1-110. acanthopterygian fishes. Bulletin of the British Museum (Natural TYLER, J.C. & SANTINI , F., 2005. A phylogeny of the fossil and ex­ History), Geology, 17 (2): 47- 102. tant zeiform-like fishes, Upper Cretaceous to Recent, with com­ PATTERSON, 1993. An overview of the early fossil record of ments on the putative zeomorph clade (Acanthomorpha). Zoologica acanthomorphs. Bulletin of Marine Science, 52(!): 29-59. Scripta, 34 (2): 157- 175. QUERO, J. C. , PORCHE, P. & VAYNE, J.J., 2003. Guide des poissons WINTERBOTTOM, R., 1974. The familial phylogeny of the de I' Atlantique europeen. Delachaux et Nestle, Lonay (Suisse), 465 Tetraodontiformes (Acanthopterygii: Pisces) as evidenced by their pp. comparative myology. Smithsonian Contributions to Zoology, 155: 1-201. REGAN , C.T., 1910. The anatomy and classification of the teleostean fishes of the order Zeomorphi. Annals and Magazine of Natural History, series 8, 6: 48 1-484. RIVATON, J. & BOURRET, P., 1999. Les otolithes des poissons de !'Indo-Pacifique. Documents scientifiques et techniques, Institut de recherche pour le developpement, Noumea, 2 (2): 1-378. ROSEN , D.E., 1984. Zeiforms as primitive plectognath fi shes. Dirk NOLF American Museum Novitates, 2782: 1- 38. Koninklijk Belgisch Instituut voor Natuurwetenschappen SANTINI, F. & TYLER, J. , 2002. Phylogeny and biogeography of the Vautierstraat 29, B 1000 Brussel, Belgie · extant species of triplespine fi shes (Triacanthidae, emai l: diJ·[email protected] Tetraodontiformes). Zoologica Scripta, 31 (4): 321- 330. SANTINI, F. & TYLER, J.C., 2003. A phylogeny of the fami lies of James C. TYLER fossil and extant tetraodontiform fi shes (Acanthomorpha, National Museum of Natural History, Smithsonian Institu­ Tetraodontiformes), Upper Cretaceous to Recent. Zoological Jour­ tion (MRC- 106), Washington D.C. 20560, USA nal of the Linnean Society, 139: 565- 61 7. email: tyleij @si.edu '' Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fishes 161

CYTTIDAE (Cytlus travers!) ZENIONTIDAE (Cyllomimus stelgis)

ZEN IONTIDAE (Zenion hololepis) OREOSOMATIDAE, Pseudocyttinae (Pseudocytlus macula/us)

GRAMMICOLEPIDIDAE, Macrurocytinae Macrurocyttus acanthopodus

OREOSOMATIDAE, Oreosomatinae (Neocyttus rhomboidalis)

GRAMMICOLEPIDIDAE, Grammicolepidinae PARAZENIDAE, Parazeninae (Parazen pacificus) Xeno/epidichthys dalgleishi

PARAZENIDAE, Cytopsinae (Cyttopsis roseus) ZEIDAE (Ze usfabet)

Fig. 1. A pictorial overview of fis hes in the Recent fa milies and subfamilies of zeiform fi shes (after TYLER e/ a/. , 2003). I I

162 DIRK NOLF & JAMES C. TYLER

T 2 Tetraodontifonnes

OSTRACilDAE (Acanthostracion quadricoris)

TRIACANTHODIDAE (Paraho/ardia linea/a)

TRlODONTlDAE {Ihodon macropterus)

TRIACANTHlDAE (Pseudotriacanthus strigi/ifer)

TETRAODONTIDAE (Lagocepha/us /aevigatus)

l::l allsttdae (JJalistapus undulatus)

DIODONTIDAE (Diodon holocanthus)

MONACANTHIDAE (Monacanthus ci/iatus)

ARACANIDAE (Aracana aurita) MOLIDAE (Ranzania /aevis)

Fig. 2. A pictorial overview of fishes in the Recent families of tetraodontiform fishes (after TYLER, 1980). ,,

Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fishes 163

postcrodorsal angle caudal ~'....------< co lliculum caudal colliculum

rostrum

rim

ventral furrow

sulcus

CHAINE & D UVERGIER (1934) Symbo/ophorus boops (Myctophidae) otolith terminology

collicular crest

collicuar crest ventral swelling Cyttus traversi (Cyttidae) Gadiculus argenteus (Gadidae)

crista superior

crista inferior expansion crista inferior Sufjlamenfraenatus (Balistidae) Triacanthus biaculeatus (Triacanthidae) Batistes capriscus (Balistidae) funne l-shaped posterodorsal angle sulcus

inferior ventral swelling Monacanthus chinensis (Monaca nthidae) Canthigaster rivulatus (Tetraodontidae) Diodon. holocanthus (Diodontidae)

Fig. 3. Morphological nomenclature of otoliths, with examples in zeiforms, tetraodontiforms, myctophids and gad ids. II

164 DTRK NOLF & JAMES C. TYLER

Scleropages leichardti OSTEOGLOSSIDAE

Synodus foet ens Polymi.xia j aponica Sargocentron xantherythrinus SYNODONTIDAE POLYMIXIIDAE HOLOCENTRIDAE

Odontesthes bonariensis Tachystorna petardi ATHERLNIDAE MUGILIDAE Fig. 4. Various types of relati vely plesiomorphic teleostean otoliths.

polarity of sulcus evolution APOMORPHIC

plesiomorphic sulcus condition

Polymixia Sargocentron Centrobetyx antiquus japonica diadema (extinct, Santoni an)

Fig. 5. Polari ty of sulcus evolution in some selected lower acanthomorph fish taxa. I I

Otolith evidence concernjng interrelationships of caproid, zeif01m and tetraodontiform fishes 165

PLESIOMORPHIC APOMORPHI C Appendix: Geographic origin and depository of the figured specimens

Institutional abbreviations: ANSP = Academy of Natural Sciences of Philadelphia; IRSNB = ln stitut royal des Sciences naturelles de Belgique; MGUH =Geological Museum, University of Copenh age11 ; MHNB = Musee d' Hi stoire naturelle, Bordeaux; USNM = National Mu seum of Natural Hi story, Smithsonian Institution. S01gocentron Perusing the following list, one may be surprised that much of the locality data is vague. As previously mentioned, otolith di ssection in tetraodon­ tiforms causes severe external damage to the specimens; thus, it is rarely performed on mu seum materials. Consequently, almost all of the tetraodon­ tiform otoliths for this study were obtained from preserved fi shes from the aquarium trade, where usually no trustworthy locality data are available. In such cases, the general distribution of the species is provided in parentheses.

Acallfhostracion quadricornis, no locality data, Aquarium? (tropical W At- lantic and Caribbean), col!. LACM. Acallfllllrus 111onroviae, W Africa, col!. IRSNB, leg. Stinton. Acantllllrus nigrofuscus, New Caledonia, col!. IRSNB, leg. Seret. Allocyflusfollefli, off California, col!. IRSNB, leg. Fitch. Allocyflus nige1; off New Zealand, col!. IRS NB , leg. Linkowski. Allocytlus vermcosus, SW Atlantic, col!. lRSNB, leg. Karrer, via Fitch. Allo111yctems whitleyi, off New Zealand, col!. IRSNB, leg. Johnston. A/uterus 111onoceros, no locality data (c ircumtropical), col!. LACM 56211.00 I, leg. Fitch. Capros Antigonia A/uterus schoepji, no locality data (W Atl an tic, from Nova Scotia to Brazil), coli. LACM 56211.002, leg. Berry. A/uterus scriptus, Aquarium Noumea, New Caledonia, col!. IRSNB, leg. Seret. A111blyrhynchotes honckenii, off South Africa, redrawn after SM ALE et a/. , 1995. Anacallfillls barbatus, off Australia, col!. LACM 5621 1.003, leg. Coates. Anoplocapros inermis, off Australi a, col!. LACM 56233.00 I, leg. Laven berg. Anop/ocapros robustus, off Sydney, Australi a, col!. LACM 56233.002, leg. Fitch. Antigonia capros, off New Caledonia, col!. IRSNB, leg. Seret Antigonia malayana, off New Caledonia, col!. IRSNB, leg. Seret. Aracana omata, off Adelaide, Australia, col! . LACM. Arothron hispidus, Aquarium Noumea, New Caledonia, coli. IRSNB, leg. Seret. Arothron inunaculatus, Taiwan, col!. IRSNB , leg. Stinton. Arothronmappa, Aquarium Nancy (lndo-W Pac ific), col!. IRSNB. Marone Kurt us Au/opus cadenmi, off Congo, col!. IRSNB. Balistapus undulatus, Aquarium (lndo-W Pacifi c), col!. lRSNB. Batistes capriscus, off Dauphin Island, Alabama, col!. IRSNB, leg. Shipp. Batistes polylepis, off California, col!. LACM 5621 1.004, leg. Fitch. Batistes vetula . Caribbean, locality unknown (Caribbean), col!. LACM 562 11.005 (Pl. 8, Fi gs 7 and 9) and aquarium, col!. IRSNB (Pl. 8, Fig. 8). Balistoides viridescens, E Africa, col!. IRSNB. Beryx decadactylus (Fig. 6), Madeira, col!. IRSNB. Fig. 6. Plesiomorphic (to left) versus derived (to right) Cantherhines dwnerilii, locality unknown (from E Afri ca to Mexico; W Pa­ morphology for, respectively from top to bottom, beryciforms, cific), LACM 562 11.006. caproids and percoids. Cantherhines macrocerus, locality unknown: «tropical W Atlantic», col!. LACM 56211.007. Cantherhines sandwichiemis, off Philippines, col!. IRSNB, leg. Stinton. Canthidermis macula/us, off California, col!. LACM 562 11.016, leg. Fitch. Canthigaster amboinensis, East Indies, col!. IRSNB, leg. Stinton. Canthigaster coronata, off Hawaii , col!. lRSNB, leg. Stinton. Canthigaster jactato1; off Hawaii, col!. LACM 56226.00 I, leg. Baldwin. Canthigaster margarita/a, Aquarium (lnclo-W Pacific), col!. IRSNB, leg. Stinton. Canthigaster p11n ctatissim11s, Pacific coast of Mexico, col!. LACM 562 11 .008. leg. Fitch. Canthi gaster rivulatus, Hong Kong fish market, col!. lRSNB, leg. Stinton. Canthigaster solandri, Aquarium (M icronesia), col!. IRSNB. Capromimus abbreviatus, off Mahia Peninsula, New Zealand, col!. IRSNB. leg. John ston. Capros ape1: locality unknown (G ulf of Bi scaye?), col!. Chaine in tRSNB. I I

166 DIRK NOLF & JAMES C. TYLER

polarity plesiomorphic condition antero-posterior constriction

Monocentris Dire/nuts Ant igonia Cyttus travers! Zeus faber reedi argenteus malayana

Fig. 7. Comparison of beryciform and zeiform otoliths and illustration of zeiform synapomorphies: antero-posterior constriction along the midline and swelling of the central upperportion of the ventral area.

polarity from moderately modified to highly modified

Diodon holocanthus DIODONTIDAE

Capros aper Triacanthodes Tydemania Pervagor spilosoma CAPROIDAE ethiops navigatoris MONACANTHlDAE TRlACANTHODIDAE Canthigaster rivulatus TETRAODONTIDAE

Fig. 8. Polarity from moderately modified to highly modified otolith morphology in Capros and selected tetraodontiforms.

Caristius macropus, Atlantic, 45° 16 'N 48°08'W, coli. Schwarzhans. Cy11us rraversi, off New Zealand, coli. IRSNB. Cemroberyx a./finis, off New Zealand, coli. IRSNB. Elops affinis, off Cali fornia, coli. IRSNB, leg. Fitch. Cemroberyx antiquus, extinct, Upper Cretaceous, Santonian, S of Tremp, Dicotylichrhys puncrularus, Aquarium (Indian Ocean), coli. IRSNB, leg. Spain. Stinton. Chaetodermis penicilligerus, Aquarium Nancy (Malaysia, Indonesia, Aus­ Diodon holocanrhus, Hong Kong fish market, coli. IRSNB, leg. Stinton. tralia), coli . IRSNB. Diodon hystrix, off New Caledonia, coli. IRSNH, leg. Seret Chilomycterus affinis, Phi li ppines, coli. LRSNB, leg. Stinton. Diodon nicthemerus, off Adelaide, S Australia, coli. LACM 26225.00 I, leg. Chi!omycterus schoepfii, locality unknown (Florida?), LACM 56211.009, Fitch leg. Berry. Diretmus argenteus, Indian Ocean, 9°34 'S 59°55'E, coli. fRSNB from Contusus richei, off Gisborne, New Zealand, coli. IRSNB, leg. Johnston. USNM 308026 Coregonus nasus, N Canada, freshwater, coli. IRSNB. Grammico/epis brachiusculus, off New Caledonia, coli. IRSNB, leg. Ctaenochaetus strigosus, off Gisborne, New Zealand, coli. IRSNH, leg. Merrett. Stinton. Kenrrocapros aculeatus, Sagami Bay. Japan, coli. IRSNB, leg. Sti nton. Cyclichthys orbicularis, off Hawaii , coli. IRSNB, leg. Stinton. Kurt us guliveri, off Bivak Island. coli. IRSNB. Cyllomimus srelgis, of lie des Pins, New Caledonia, coli. IRSNB , leg. Seret. Lactophrys trigonus, locality unknown (tropical W Atlantic), coli. LACM Cyllopsis t-yphu, off NW Australia, coli. IRSNB, leg. Seret. 562 11 .01 2, leg. Fitch. Cyllopsis roseus, locality unknown (Gul f of Biscaye?), coli. Chai ne in Lacroria jomasini, off south Africa, coli. IRSNB, leg. Stinton. MHNB. Lagoceplwlus !tmaris, Hong Kong fish market. coli. IRSNB, leg. Stinton. Cyrrus ausrralis, off Auk land , new Zealand, coli. IR SNB, from USNM 177 Lagocephalus sclerarus, off New Caledonia, coli. IRSNB. leg. Serer. 084. Macrorhamplwsus scope/ax, off California, coli. LRSNB, leg. Fitch. Cyuus no va e~e /andia e . off Mahia Peninsula, New Zealand, coli . IRSNH, Melichrhys niger, Pacific, off Socorro, Mexico. coli. LACM 56229.0 II . leg. leg. Johnston. Fitch. '' Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fi shes 167

Neocyttus rhomboidales --

Allocyttus niger

Parazen pacific us ----~

Cyttopsis__ roseus

Capromimus abbreviatus --

cyttonum. us__ -·-' · stelg1s s'· , · ·uA !i;;-

Macrurocyttus : no data

ICOLEPIDIDAE Xe nolepid. ichthys da lgleishi 8 '

Fig. 9. Otolith-based testing of TYLER, O'TOOLE & WI NTERBOTrOM 's (2003) cladogram of zeiform interrelationships (thei r fi g. 4). I I .168 DIRK NOLF & JAMES C. TYLER

TR!ACANTHODIDAE

Triacanthodes Tydemania • TRJACANTHIDAE § [:5 Triacanthus Pseudotriacanthus ~~ ~ ~ Balistapus Sufflamen Batistes c£) Anacanthus Monacanthus Chaetodermis

ARACANIDAE

A rae ana Kentocapros OSTRACIIDAE § Acanthostracion Tetrosomus

-'- ;; -Triodon

Lag acephalus Canthigaster

Diodon

MOLIDAE

Ranzania

Fig. 10. Otolith-based testing of SANTINI & TYLER 's (2003) claclogram of Recent tetraoclo ntiform interrelationships (their fig. 5, only extant taxa shown here). Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fishes 169

·· ... ~~ ,;,:':::'>c. ~

lvlacrorhamphosus gracilis Platybe1yx opalescens Carislius macropus (LOWE, 1839) ZUGMA YER, 19 11 (BELLOTTI, 1903) MACRORHAMPHOSIDAE CARISTIIDAE

Fig. 11. Otoliths of Macrorhamphosidae (Gasterosteiformes, Syngnathoidei) and Caristiidae (Percoidei).

Melichthys vidua, off Philippine islands (Pl. 7, Fig. 5), coli. IRSNB and off Siga11us doliatus, off New Caledonia, coli. IRSNB, leg. Serer. John ston Island (Pl. 7, Fig. 6), coli. LACM 56230.00 I, leg. Bald win. Siga11us virgatus, Hong Kong fish market, coli. IRSNB, leg. Stinton. Meuschenia australis, off New Zealand, coli. IRSNB. Siga11us vu lpi11us, Aquarium (lndo-W Pacific), coli. IRSNB. Meuschenia scabe1; Tasman Bay, New Zealand, coli. IRSNB. Sphoeroides lobatus, tropical E Pacific, coli. LACM 562 11.01 3, leg. Fitch. Manacanthus chinensis, off Australia, LACM 56212.005, leg. Coates. Sphaeroides 11 ephelus, off Louisiana, coli. LACM 56232.00 I, leg. Fitch. MaJwcalltlw~· lu cke ri , Aquarium (West Indies), coli. IRSNB, leg. Stinton. Sphoeroides spe 11 gleri, locality unknown (tropical W Atlantic), coli. LACM Manacentris reedi, off San Felix Island (Chile), coli. IRSNB, leg. Fitch. 5621 1.014, leg. Berry. Mara11e /abrax, loc. unknown (Gulf of Biscaye?), coli. Chaine in !RSNB. Splweroides testudi11 eus, off Haiti, coli. IRSNB, leg. Moreaux. Nasa unicarnis, off New Caledonia, coli. LRSNB , leg. Seret. Steplwnolepis hispidus, off Lanzarote, coiL IRSNB. Nasa v/am ingii, Aquarium Nancy (Indo-W Pacific), coli. IRSNB. Stetlwpristes eos, off Papua New Guinea, coli. IRSNH, leg. Seret. Neacyllus he/gae, North atlantic, coli. Schwarzhans Sufjlamen bursa, Aquarium, (E Africa), coiL IRSNB, leg. Stinton. Neacyttus rhambaidalis, off South Africa, redrawn after SMALE eta/., 1995. Sufflamen chrysopterus, Aquarium (Red Sea to Phi li ppines), coli. IRSNB. Odanteslhes banariensis, off Uruguay, coli. IRSNH. leg. Stinton. Odanus nige1; aquarium Noumea, New Caledonia; coli. IRSNB, leg. Seret. Sujjlamen jrae1wtus, off New Caledonia, coiL IRSNB, leg. Serer. Oreasama at/anticum, off South Africa, redrawn after SMALE eta/. , 1995. Sujjlamen verres, Pacific coast of Mexico, coiL LACM 5621 1.015, leg. Ostracian cubicus, Aquarium Noumea, New Caledonia (Pl. 14, Fig. 5), Fitch. aquarium, leg. Stinton (Pl. 14, Fig. 6), coli. IRSNB. Synodusjoetens, off Georgia (Fig. 4e) and off Texas (Fig. 4f), coiL IRSNB. Ostracia11 me/eagris, Aquarium, E Africa (Pl. 14, Fig. 8), Hawaii (Pl. 14, Takifugu oblong us, Australi a, col i. LACM 562 12.004, leg. Coates. Fig. 9), coli. IRSNB, leg. Stinton. Takifugu parphyreus, off Japan, coli. IRSNB, leg. lizuka. Ostracian pergravis, Paleocene, Selandien, Denmark, coli. MGUH 26232 Takifugu xantlwptem s, Hong Kong fish market, coiL IRSNB, leg. Stinton. and 26233. Tetraodonjluviatilis, Aquarium (freshwater, South Asia), coli IRSNH. Oxyma11acanthus /angirastris, Aquarium (Indo-Pacific), coli. IRSNB, leg. Tetraadan pa/embange11sis, Thailand, coiL LRSNB , leg. Stinton. Stinton. Tetrosomus gibbosus, Aquarium (Indo-Pacific), coiL IRSNB, leg. Stinton. Paracamhurus hepatus, Aquarium (lndo-W Pacific), coli. IRSNB. Thamnaconus hypargyreus, NW Australia, coli. IRSNB, leg. Seret. Paralweres pria11ums, Aquarium, E Africa, coli. IRSNB, leg. Stinton. Tlwmnaconus modest us, Hong Kong fish market, coi L IRSNB, leg. Stinton. Paramanacanthus ob/angus, off Australia, coli . LACM 562 12.002, leg. Tra chysta111a petardi, off Townsville, Australia, coiL IRSNB, leg. Fitch. Coates. Triacanthodes ethiops, off Madagascar, coiL IRSNB, leg. Schwarzhans. Paratriaca11thades retrospinis, off Mozambique, coli. ANSP 103285. Triacanthus biaculeatus, Hong Kong fish market, coiL IRSNB, leg. Stinton. Parazen pacific us, off Miami, Florida, coli. IRSNB. Triacanthus sp., Ypresian , London Clay, coiL Rundle, new fossi l record. Pentanemus quinquariu~·. offW Africa, coli. IRSNB. Triodonmacrapterus, off New Caledonia, coli . IRSNB , leg. Seret. Pervagor melanoceplwlus, Aquarium, E Africa, coli. lRSNB, leg. Stinton. Trixipilichthys weberi, Andaman Sea, Thailand, coli . ANSP 102136. Pervagar spilasoma, Hawaii. coli. IR SNB, leg. Stinton. Tydemania 11a vigatoris, NW Australia, coiL IRSNB, leg. Seret. Platyberyx apalescens, Atlantic, 65°N 32°50' W, coli. Schwarzhans. Tylerius spi11osissimus, Hong Kong fish market, coli. IRSNB, leg. Stinton. Polymixiajaponica, off Japan, coli. IRSNB. Xamhichthys menlo, Socono Islands, coiL LACM 56234.00 I, leg. Fitch. Pseudobalistes juscus, Philippines, coli. IRSNB, leg. Stinton. Xenolepidichthys dalgleishi, W Africa, coiL !RSNB, leg. Fitch. Pseudacy11us maculatus, off New Zealand , leg. Linkowsk i (Pl. I, Figs. 7-8) Zane/us comwus, Aquarium Nancy (Atlantic, from E Africa till Mex ico; W and Antartica, El tanin station 1679 (Pl. I, Fig. 9), coli. IRSNB. Pacific), CoiL IRSNB. Pseudomoiwcantlws 1nacrurus, off Australia, col i. LACM 562 12.003, leg. Zebrasomajlavescens, Aquarium (Indo-Pacific), coi L LRSNB . Coates. Zenion hololepis, Caribbean, coli. IRSNB, leg. Fitch. Pseudotriacanthus strigilifer, off Bangkok, coli. LACM 56231.00 I, leg. Ze11i0 11 japonicus, off Queensland. Australia, coli. IRSNH, leg. Laven berg. Schwarzhans. Ran~ania /aevis, off New Caledonia, coli. IRSNB, leg. Seret. Zenionlongipinnis, off New Caledonia, coiL IRSNB , leg. Serer. Rhinecantlws rectallgulus, Aquari um, (Indo-Pacific), coli . JRSNB, leg. Ze11ops is conch({e1; locality unknown (Gulf of Biscaye?), coli . Chaine in Stinton. MHNB. Sargocentron diadema (Fig. 6), Aquarium (Red Sea to Chin a) , coli. IRSNB, Zenopsis nebulosus, off Mahia Peninsula, New Zealand, coli . IRSNB. leg. leg. Stinton. Johnston. Sargocentron xa11tlwyrhrinus. off Hawaii, coli. IRSNB, leg. Stinton. Zeusjabe1; loc. un known (G ulf of Bi scaye?). coiL Chaine in IRSNB. Sc!eropages /eichardti, Au stralia, fre shwater. coli. IRSNB, leg. Fitch. 11 170 DIRK NOLF & JAMES C. TYLER

Cyttus australis (RICHARDSON, 1843)

Cyttus traversi HUTION, 1872

Cyttus novaezelandiae (ARTHUR, 1885)

4c

7c

Pseudocyttus maculatus (GILCHRIST, 1906)

Plate 1. Otoliths of Cyttidae and Oreosomatidae (partim) Pseudocyttinae II

Otolith evidence concerning inten-elationships of caproid , zeiform and tetraodontiform fi shes 171

Al/ocyttus niger JAMES, INADA & NAKAMURA , 1988

Allocyttusfolletti M YERS, 1960

Allocyllus verrucosus (GILCHRIST, 1906)

Neocyttus helgae (HOLT & BYR NE, 1908) Neocyttus rhomboida/is GILCHRIST, 1906 Oreosom.a atlanticum CUY lER , 1829

Plate 2. Otoliths of Oreosomatidae. Oresomatinae II

172 DIRK NOLF & JAMES C. TYLER

Parazen pacific us KAMOHARA, 1935 Cyuopsis cypho (FOWLER, 1934) PARAZEN IDAE, Parazeni nae PARAZENIDAE, Cyttopsinae

Cyttopsis rose us (LOWE, 1843) Stethopristes eos G ILBERT, 1905 PARAZEN IDAE, Cyttopsinae PARAZENIDAE, Cyttopsinae

6c r

Capromimus abbreviatus (HECTOR, 1874) Cyttomimus stelgis GILBERT, 1905 ZEN IONTIDAE ZEN!ONTIDAE

Plate 3. Otoliths of Parazenidae and Zeniontidae (partim): Capromimus and Cyttomimus '' Otolith evidence concerning inteiTelationships of caproid, zeiform and tetraodontiform fishes 173

Ia ...... · ......

6b

Zenion hololepis (GOODE & BEAN, 1896)

Zenionjaponicus KAMOHARA , 1934 7c

Zenion longipinnis KOTTHAU S, 1970

·/ ~l Oa ~~·· . extinct ,~ ~· ,: .:~ · . ·· ·.: .. )_. ···· ,.. Ypresmn, ~ · ··:=: London Clay .··!".'- ' '~· '-~ ·~ -..::. . 2c 2b .·.··:.:(

right otolith "genus Zeniontidarum" sulcifer (STINTON, 1966) ex tinct, Gan, Santonian, Pyrenees Chateau Merce

3 .··._::::-. "gen us aff. Zen ion" sp. ··.-.::::·· ·.. ~ · · .... · ·. . . : . . ·: extinct, Ypresian, Gan Clay

:; . . ~ ~:• .. ·.·.

5 ·'Bava ri an zeiform·', "Bavarian zeiform", ex tinct, Maastri chti an ex tinct, Paleocene '·genu s Zeiformorum" tyleri NOLF, 2003

Plate 4. Otoli th s ofZeniontidae, Zenion and fo ss il relati ves+ incertae sedis fo ss il zeiforms I I

174 DIRK NOLF & JAMES C. TYLER

specimen figured by CAMPANA, as Daramaltus americanus

2c

Zeus f aber LTNNA EUS, 1758

3c

Zenopsis conchifer (LOWE, 1852) Zenopsis nebulosus (TEMMTNCK & SCHLEGEL, 1845)

Plate 5. Otoliths of Grammicolepididae and Zeidae ''

Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fishes 175

','. ~ . ;:, .

,::;.;:\;i.;:t.:i: .:Vz~·{::··:::.('/:; · .)· :; .-·- I b l lc

Triacanthodes ethiops ALCOCK , 1894 Tydemania navigatoris WEBER, 191 3

\

Paralriacanthodes retrospinis Trixiphichthys weberi Pseudotriacanthus strigi/ifer (CANTOR, 1849) fOWLER, 1934 (CHAUDHURJ, 191 0)

6c 7c \ / i i Triacanthus biaculeatus (BLOCH, 1786) Triacanthus sp. extinct, Ypresian, London lay

Plate 6. Otoliths ofTriacanthodidae and Tricanthidae II

176 DIRK NOLF & JAMES C. TYLER

~ ~ 3a

Melichthys niger (BLOCH, 1786) Odonus niger (RUPPELL, 1836)

Balistapus undulatus (MUNGO PARK, 1797) Melichthys vidua (RICHARDSON, 1845)

Xanthichthys menlo (JORDAN & GILBERT, 1882)

Plate 7. Otoliths of Balistidae (partim): Balistapus, Melichthys, Odonus and Xanthichthys II

Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fis hes 177

ri ght otolith

left otolith

Batistes capriscus GMELIN, 1789 Batistes vetula LINNAEUS, 1758

Balistoides viridescens 13b Balistes po{y{epis STEINDACHNER, 1876 (BLOCH & SCHNEIDER, 1801) Canthidermis macula/us (BLOCH, 1786)

left otolith Pseudobalisles fitscus (BLOCH & SCHNEIDER, 180 I ) Rhinecanthus reel angulus (BLOCH & SCHNEIDER, 180 I)

Plate 8. Otoliths of Balistidae (partim): Balistes, Camhidermis, Pseudobalistes and Rhinecamhus II

178 DIRK NOLF & JAMES C. TYLER

left otolith right otolith

Sufflamen chrysopterus (BLOCH & SCHNEIDER, 1801)

Sufflamenfraenatus LATREILLE, 1804 Sufflamen bursa (BLOCH & SCHNEIDER, 1801)

Plate 9. Otoliths of Balistidae (partim): Su.fflamen Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fishes 179

4c

A/uterus schoepji (WALBAUM, 1792) A/uterus scrip/us (OSBECK, 1765)

A/uterus monoceros (LINNAEUS, 1758) Anacanthus barbatus GRAY, 1830

Meuschenia scaber (FOSTER, 180 I) lvleuschenia australis (DONOVAN, 1824)

13 c

Chaetodermis penicilligerus (CUYlER, 181 6) Para/uteres prionurus (BLEEKER, 185 1)

Plate 10. Otoliths of Monacanthidae (partim): A/uterus, Anacanthus, Chaetodennis. Meuschenia, Paraluteres II

180 DIRK NOLF & JAMES C. TYLER

3a

Cantherhines macrocerus Cantherhines sandwichiensis Cantherhines dumerilii (1-!0LLARD, 1853 (1-!0LLARD, 1853) (QUOY & GA IMARD, 1824)

~ · right otoli th 6c 4a right otoli th

4c

7 J'vfonacanthus Iucken· B EA N, 1906

5c Monacanthus chinensis (OSBECK, 1765)

!I a Oxymonacanthus /ongirostris (BLOCH & SCHNEIDER, 180 1)

lOb

12b Pervagor me/anocepha/us (BL EEKER, 1853) Pervagor spilosoma (LAY & BENNETT, 1839)

Plate 11. Otoliths of Monacanthidae (partim): Cantherh.in.es, Monaca nth us, Pervagor and Stephanolepis II

Otoljth evidence concerning interrelationships of caproid, zeiforn1 and tetraodontiform fishes 181

lc

6c Pseudomonacanthus macrurus (BLEEKER, 1857) 6a I I

/

Th amnaconus hypargyreus (COPE, 1873)

7c

Paramonacanthus oblongus (TEMMJNCK & SCHLEGEL, 1850)

Thamnaconus modestus (GUENTHER. 1877)

Plate 12. Otoliths of Monacanth idae (partim): Paramonacanthus, Pseudomonacanthus, Stephanolepis and Th amnaconus I I

182 DIRK NOLF & JAMES C. TYLER

left otolith right otolith

Triodon macroplerus LESSON, 1829

3c

Anoplocapros robust us (FRASER- BRUNNER, 1941) Anop/ocapros inermis (FRASER - BRUNNER, 1935)

... :..... :.. .··.:

Aracana ornata (GR AY, 1838) Kentrocapros acu/ea/us (HOUTTUYN, 1782)

Plate 13. Otoliths ofTriodontidae and Aracanidae I I

Otolith evidence concerning interrelationshjps of caproid, zeiform and tetraodontiform fishes 183

3c

Laclophrys lrigonus (LINNAEUS, 1758)

~ ~a

Acanlhoslracion quadricornis (LINNAEUS, 1758) Lactoria fornasini ( B IANCONI, 1846)

;"'~~;~0,~~f;:i~*! ...... --.~-

5b

5c Ostracion cubic us LINNAEUS, 1758 Tetrosomus gibbosus ( L INNAEUS, 1758)

10 ' ' ~- ~ ....- ..... :· 8a lie

9 Ostracion pergravis S CH WA RZHANS, 2003 Ostracion meleagris S HAw , 1796 extinct. Paleocene. Seland ien, Denmark

Plate 14. Otoliths of Ostraciidae ''

184 DIRK NOLF & JAMES C. TYLER

4a

Arothron immaculatus (BLOCH & SCHNEID ER, 180 1)

Arothron hispidus (LINNAEUS, 1758)

Lagocephalus see/era/us (GMELIN, 1789) Arothron mappa (LESSON, 183 1)

Amblyrhynchotes honckenii (BLOCH , 1795)

Lagocephalus lunaris (BLOCH & SCHNEIDER, 180 I)

Con/usus richei (FREMINVILLE, 18 13)

Plate 15. Otoli th s ofTetraodontidae: Amblyrhynchotes, Arothron, Contusus, Lagocephalus II

Otolith evidence concerning interrel ationships of caproid, zeiform and tetraodontiform fishes 185

right otolith

3c 2c

Canthigaster corona/a Canthigasterjactator (JENK INS, 190 I) Canthigaster amboinensis (BLEEKER, 1865) (VAILLANT & SAUVAGE, 1875)

right otolith

6c

Canthigaster punctatissimus Canthigaster rivulatus (GUENTHER, 1870) (TEMMINCK & SCHLEGEL, 1850) Canthigaster margarita/a ( RUEPPELL, 1829)

7c 8a

Sphoeroides lobatus (STEINDACHNER, 1870) Sphoeroides spengleri (BLOCH, 1785)

lOb

Sphoeroides nephelus (GOODE & BEAN, 1882) Sphoeroides testudineus (LINNAEUS, 1758)

Plate 16. Otol iths ofTetradontidae: Canthigaster and Sphoeroides II

186 DIRK NOLF & JAMES C. TYLER

Takifugu porphyreus (TEMMINCK & SCHLEGEL, 1850)

Takifugu xanthopterus (TEMMIN CK & SCHLEGEL, 1850)

JOe

lOb

Tylerius spinosissimus (REGAN, 1908) Tetraodonjluviatilis 1-IAMIL TON. 1822

Plate 17. Otoliths ofTetraodontidae: Takifugu, Tetraodon and Tylerius II

Otolith evidence concerning interrelationshjps of caproid, zeiform and tetraodontjform fishes 187

Chilomycterus affinis G UENTH ER, 1870

4c A//omycterus whit/eyi PHILLIPPS, 1932 Chi/omycterus schoepjii (WALBAUM, 1792)

Cyc/ichthys orbicularis (BLOCH, 1785)

Diodon hystrix LINNA EUS, 1758 5

Diodon nicthemerus C UVI ER, 18 18 Dicotylichthys punctu/atus KAUP, 1855

8c

9

Diodon holocanthus LINNAEUS, 1758 Ranzania laevis (PENNA NT, 1776)

Plate 18. Otoliths of Di odontidae I I

188 DIRK NOLF & JAMES C. TYLER

6c

Acanthurus monroviae STEINDACHNER, 1876

Ctenochaetus strigosus (BENNETT, 1828)

Acanthurus nigrofitscus (FORSSKAL, 1775)

~, o, Zebrasoma xanthurum (BLYTH, 1852) Zebrasoma jlavescens (BENN ETT, 1828)

Plate 19. Otoliths of Acanthuridae (partime): Acanthurus, Ctenochaetus and Zebrasoma II

Otolith evidence concerning interrelationships of caproid, zeiform and tetraodontiform fi shes 189

Zane/us cornu/us (LINNAEUS, 1758)

Paracanthurus hepatus (LINNAEUS, 1766)

Siganus virgalus (VALENCIENNES, 1835)

Siganus doliatus CUYlER , 1855 Siganus vulpinus ( , 1845)

Plate 20. Otoliths of Acanthuridae (partim): Paracanthurus and Nasa; Siganidae: Zanclidae