Bull. Soc. belge Géol., Paléont., Hydrol. T. 78 fasc. 3-4 pp. 219-234 Bruxelles 1969 Bull. Belg. Ver. Geol., Pa!eont., Hydrol. V. 78 deel 3-4 blz. 219-234 Brussel 1969

A TELEOST OTOLITH FAUNA FROM THE SANDS OF LEDE, BELGIUM

F. C. STINTON and D. NoLF

The tertiary strata of Belgium is renowned ichthyologists in the past has united the for its rich fish fauna and t?e elasmobranch ophidioid fishes with the blennioids. However, remains have been adequat~ly described in the a study of the otoliths of members of each excellent works of Leriche, Casier and others. order immediately reveals the inaccuracies of The teleost remains have received a very this system and shows that the ophidioids are limited attention despite their being as com­ closely related to the gadoids. McALLISTER mon as those of the elasmobranchs but this (1968), proposed the orders Ophidiidiformes probably results from the difficulty of iden­ and Gadiformes, consecutively but the rela­ tifying such fragmentary specimens. However tionship shown by the overall characters pre­ it is possible to establish a teleost fauna on cludes their ordinal separation. The classific­ the evidence of the numerous otoliths which ation adopted in the present work is that occur in these strata. A few otoliths have been proposed by GREENWOOD, ROSEN, WEITZMAN described from the sands of Brussels and the & MYERS (1946), (see also ROSEN & PATTER­ Ieper formation by LERICHE (1905) and CASIER SON, 1969), which places each group at sub­ (1946) but all were referred to incorrect ordinal level within the one order Gadiformes. genera. None have so far been recorded from The family Antigoniidae is included with the the sands of Lede and it is proposed to berycoids as proposed by STINTON (1967). describe a series obtained by one of us (D.N.). These resulted from the processing of some Systematic Descriptions 300 Kg. of matrix from an outcrop of the Sands of Lede at Balegem. In the present Class Actinopterygii work the basement bed of the sands of Lede Subclass Teleostei is excluded because of its content of Superorder Elopomorpha reworked material although any forms known Ortler Anguilliformes to occur at this horizon will be mentioned in Suborder Anguilloidei the text. Family Anguillidae All the otoliths so far found have been Genus Anguilla Shaw, 1803 sacculiths, all were from fish which had 1803. Anguilla Shaw, G. 4, 15: type Muraena evidently not reached full maturity and all anguilla Linnaeus ( = Anguilla vulgaris show some degree of erosion. It is probably Shaw). these factors together with a lack of suitable ANGUILLA RECTANGULARIS sp. n. Pl. comparative material from extant fishes which figs. 1,2. lead to the misidentifications of LERICHE and Holotype: Registration No. Ht. p. 1588. CASIER, as mentioned by STINTON (1968). Dimensions: Length 3.00 mm (incomplete) 20 teleost species will be established of which Width 2.55 mm. 14 are undescribed. Material: The unique holotype. A thin, rather rectangular, right sacculith Classification which has lost the rostral area. Slightly un­ The scheme accepted by the majority of dulant, horizontal dorsal rim with a small

219 projection at the junction with the vertical, ly undulant sulcus opening on the anterior rounded posterior rim; rounded, somewhat rim and not reaching the posterior rim in its flattened ventral rim; details of anterior rim diagonal course. The sulcus shews little dif­ missing. Markedly concave outer face with ferentiation into ostium and cauda, being radial tuberosities indistinctly visible on ail equal in depth and width throughout. Crista areas. A convex inner face with a median, superior and crista inferior slightly thickened. curved sulcus ending near the posterior rim. Rostrum, antirostrum and excisura present The ostium is missing in this specimen but but no collicula. there is a long, narrow cauda present. No The features of this otolith are very similar collicula. to sacculiths of Gymnothorax ocellatus saxi­ More complete specimens have been reco­ cola JORDAN & DAVIS, btit the outline of the vered by one of us (F.C.S.) from the lower sulcus is more regular in the fossil form. Bartof! beds of the English eocene and one PRIEM (1906), considered this form (as Oto­ of these (fig. 2) shows a very short, wide, lithus (Congeris) sp.) to be synonymous with deep ostium and a long narrow cauda differ­ his Otolithus ( Congeris) papointi but the two entiated by a narrowing of the crista superior types are quite distinct. and crista inferior. A prominent rostrum and Sands of Lede, Balegem, Belgium. Common. excisura are present but no antirostrum or Basement Beds of the sands of Lede, Bale­ collicula. gem. This otolith compares closely with saccu­ Sands of Aalter, Aalter and Scherpenberg, liths of the living Anguilla reinhardti Stein­ Belgium. dachner, both in its peripheral contours and Upper Bracklesham Beds, England. Rare. the characters of the sulcus. Lutétien, France, Rare. Sands of Lede, Balegem, Belgium. Upper Bracklesham Beds and Lower Barton Family Moringuidae Beds, England. Genus Moringua Gray, 1831. Family Muraenidae 1831. Moringua Gray J.E., Zoo!. Mise., 95: Genus Gymnothorax Bloch, 1795. type Moringua lateralis Gray. MoRINGUA FISSURA sp. n. Pl. figs. 4a,b l 795. Gymnothorax Bloch, M.E. 9, 83: type Gymnothorax reticularis as first de­ Holotype: Registration No. Ht. p. 1590 .. signated by Bleeker, 1865. Material: The unique holotype. GYMNOTHoRAX DIAGONALIS sp. n. Pl. Dimensions: Length 4.21 mm. Width 2.06 mm. fig. 3. A somewhat eroded right sacculith. Slightly 1906. Otolithus (Congeris) sp. Priem F., Bull concave, almost horizontal dorsal rim; round­ Soc. géol. Fr., (4), 6, 276, figs. 44-45. ed posterior rim; irregularly rounded ventral rim, deepest towards the posterior end; short, Holotype: Registration No. Ht, p. 1589. oblique anterior rim. Moderately concave Material: 21 sacculiths. outer face which is smoothly rugose. Severa! Dimensions: Length 3.96 mm. Width 2.86 mm concentric growth lines are visible (fig. 4b) A biconvex, regularly rounded, left saccu­ through the effects of attrition. Convex inner lith with the rims coalescing. The dorsal rim face with a deep, somewhat diagonal, undu­ is short, slightly rounded and highest ante­ lant sulcus which opens on the anterior rim riorly; short, slightly obtuse posterior rim; and terminates well short of the posterior long, regularly rounded ventral rim; long, rim. Sulcus consisting of a short, rather oval oblique anterior rim which is notched at its ostium and a long, arcuate cauda with a ventral end. Outer face eroded but undulant truncated end. An obtuse upper angle at the with a depression from the antero-ventral junction of ostium and cauda. The crista corner towards the centre of the otolith. inferior is continuous and undulant. A feeble Smooth inner face with a conspicuous, slight- rostrum is present but no antirostrum, exci-

220 sura or collicula. The posterior end of the Bracklesham beds with N.laevigatus; upper otolith is expanded and somewhat up-turned Bracklesham beds with N. variolarius, Lower giving the otolith a moderate torsion. Barton beds, England. Common in the Mio­ This otolith shows a considerable resem­ cardia Bed (CURRY, 1958) = Fischer Bed 17, blance to the sacculith of the living Maringua Bracklesham Bay, Sussex. micrachir Bleeker, figured by KoTTHAUS (1968. Calcaire grossier and Auvers formation, fig. 112 p. 48) differing in its concave dorsal France. rim and irregular ventral rim. Superorder Scopelamorpha Sands of Lede Balegem Belgium. Ortler Myctaphiformes Family Congridae Su border Myctophaidei Genus Paraconger Kanazawa, 1961. Family Synadantidae 1961. Paracanger Kanazawa R.H .. Proc. U.S. Genus Synodus Gronow, 1763. Nat. Mus., 113, 4: type Eche/us caudi­ 1763. Synadus Gronow L.T., Zoophyl., 112: limbatus Poey. type Salma synodus Linnaeus. PARACONGER SAUVAGE! (Priem) Pl. fig. 5. SYNODUS RECTUS (Frost) Pl. fig. 6. 1906. Otalithus (Sparidarum) sauvagei Priem 1933. Otolithus (Elaps) rectus Frost G.A., F., Bull. Soc. géol. Fr., (4), 6, 274. figs. Ann. Mag. Nat. Rist., (10), 12, 389 pl. 36, 37. 12 fig. 2. 1932. Cangramuraena amaris Suie J., Rozpr. Material: 3 sacculiths. geol. Ûst. es!., Praze. 7, 62. pl. 1 figs. Registration: No. 24/181. 10 a,b. Dimensions: Length 4.25 mm. (incomplete) 1933. Otolithus (Platessa) sectar Koken ; Width 2.12 mm. Burton E.St.J., Proc. Geol. Ass., 44, The holotype of Frost is a very eroded 160. Name only. errore. specimen in which the peripheral contours 1933. Otalithus (Congramuraena) amaris Suie; have been considerably modified. It bas be­ Frost G.A., Ann. Mag. Nat. Hist., (10), come pointed posteriorly and has lost much 12, 394. pl. 12 fig. 19. of the detail of the sulcus. In the present 1968. Paracanger amaris (Suie); Stinton, P.C. specimen it will be seen that the dorsal rim Coll. Eoc. p. 157, pl. fig. 11. is crenulated while the posterior tim is round­ ed and bilobed. Also, the ostium is deeper Registration No. 24/181. than the cauda from which it is separated by Dimensions: Length 5.55 mm. Width 4.40 mm. distinct upper and lower angles. Material: 54 sacculiths. This specimen compares exactly with well­ This sacculith is less eroded than those preserved examples found by one of us figured by PRIEM (1906) and SuLc (1932). The (F.C.S.) in the lower Barton beds of Hamp­ posterior rim is seen to be feebly crenulated, shire, England. It is undoubtedly the species the dorsal rim is faintly crenulated and the described by Frost although it is evident that cauda has a bulbous extremity. he was mislead by the contours of his holo­ This otolith is remarkably similar to the type. The widely divergent features of the sacculiths of the living Paracanger macraps fossil form and sacculiths of Elaps confirm the (Günther), as noted in an earlier paper misidentification. However, sacculiths from (STINTON, 1968). living species of Synodus Gronow, show Localities: distinct similarities to the fossil which must Sands of Lede, Balegem, Belgium. Com­ be referred to this genus. mon. Localities: Basement beds of the sands of Lede, Bale­ Sands of Lede, Balegem, Belgium. gem. Upper Bracklesham Beds and Lower Bar­ Sands of Aalter, Aalter and Scherpenberg. ton Beds, England.

221 Superorder Paracanthopterygii Family Brotulidae Order Gadiformes Genus Hoplobrotula Gill, 1863. Su border Ophidioidei 1863. Hoplobrotula Gill T.N., Proc. Acad. Family Ophidiidae Nat. Sei. Philad., 15, 255: type Brotula Genus Lepophidium Gill, 1895. armata Temminck & Schlegel. 1895. Lepophidium Gill, T.N., Amer. Nat., HOPLOBROTULA LERICHE! sp. n. Pl. fig. 8. 29, 167: orthotype Leptophidium pro­ Holotype: Registration No. Ht. p. 1591. fundorum Gill. Substitute for Lepto­ Material: 48 sacculiths. phidium Gill, preoccupied in Snakes. Dimensions: Length 5.04 mm. Width 2. 75 mm. LEPOPHIDIUM AEQUALIS sp. Il. Pl. fig. 7. A rather worn, elongate right sacculith, Holotype: Registration No. Ht. p. 1592. produced to a blunt posterior point. Dorsal Material: 112 sacculiths. rim undulant, rising to an anterior point; Dimensions: Length 5.80 mm. Width 3. 79 mm. short, concave posterior rim; shalowly round­ ed ventral rim; obtuse, slightly undulant A slightly eroded left sacculith, pointed anterior rim. Concave outer face, smoothly posteriorly and bluntly rounded anteriorly. undulant but in occasional specimens radial Dorsal rim sloping slightly upwards towards ribbing to a median ridge is evident, indica­ the anterior end where it forms a marked ting that unworn examples would have prominence at its junction with the anterior crenulated rims. Convex inner face with a rim; short, oblique, slightly undulant poste­ median, wide sulcus which opens very nar­ rior rim which coalesces with the dorsal rim rowly on the anterior rim and does not reach to form a very slight rounded angle; rounded the posterior rim. Sulcus consisting of a long, ventral rim which is scalloped on the posterior oval ostium sharply delineated from the short, part; irregularly rounded anterior rim, slightly uncinate cauda which forms a marked lower produced centrally and feebly crenulated on angle at its junction with the ostium. Crista the antero-dorsal projection. Outer face pro­ superior continuous. Ostium filled with a col­ minently and irregularly tuberculated in un­ liculum. No rostrum, antirostrum or excisura, worn specimens. A convex inner face with a This form is remarkably similar to saccu­ median sulcus which is completely enclosed liths of the living Japanese species Hoplo­ apart from a very narrow, short shallow brotula armata (TEMMINCK & SCHLEGEL), groove to the anterior rim. It is elliptical and differing only in the deeper ventral rim in the undifferentiated, being rounded at the caudal fossil form. end in unworn specimens. Eroded specimens usually shew a tapered caudal end. Sulcus Localities: completely filled with a colliculum. The lower Sands of Lede, Balegem, Belgium. area is faintly ornamented with scalloped Basement bed of the sands of Lede, Bale­ ridges on the posterior part. gem. This otolith shews a marked similarity to Calcaire grossier, France. sacculiths of the living Lepophidium cervinum Genus Ogilbia (Jordan & Evermann) GoooE & BEAN, but differs in being relatively Evermann & Kendall, 1898. thinner while L. cervinum has a differentiated 1898. Ogilbia (Jordan & Evermann) Ever­ sulcus, the cauda being very short, bulbous mann B.W. & Kendall W.C., Bull. U.S. and indistinct. Fish. Comm., 17, (1897) 132: orthotype Localities: Ogilbia cayorum Evermann & Kendall. Sands of Lede, Balegem, Belgium. Very 0GILBIA SUBREGULARIS (Schubert) Pl. common. figs. 9,10. Basement bed of the sands of Lede. 1916. Otolithus (Ophidiidarum) subregularis Upper Bracklesham Beds, England. Very Schubert R.J., Jahrb. der k.k. geol. rare. Reichs., 65, 285, pl. 7 figs. 35-37.

222 1933. Otolithus (Ophidiidarum) cf. acutangu­ Basement bed of the sands of Lede. lum Koken; Burton E.St.J., 1933. Proc. Sands of Aalter, Aalter, Belgium. Geol. Ass., 44, 160. Name only. errore. Upper Bracklesham Beds and Barton Beds, Otolithus ( Ophidiidarum) subregu[aris England, where it is very common at some Schubert; Burton E.St.J., ibid., 44, 160. horizons. Name only. Calcaire grossier, France. Rare. 1934. Otolithus (Ophidiidarum) planus Frost Superorder Acanthopterygii G.A., Ann. Mag. Nat. Hist., (10), 14, Ortler Beryciformes 444. pl. 12 fig. 6. Suborder Berycoidei Family Trachichthyidae Materials: 13 sacculiths. Genus Trachichthodes Gilchrist, 1903. Dimensions: Left sacculith (Fig. 9) Length 4.86 mm. 1903. Trachichthodes Gilchrist J.D.F., Mar. Width 2.39 mm. Invest. S. Afr., 2, 203: type species Right sacculith (Fig. 10) Length 3.07 mm. Trachichthodes spinosus Gilchrist. Width 1.75 mm. TRACHICHTHODES CIRCULARIS (Shepherd) PL fig. 11. An eroded left sacculith (fig. 9). Smoothly 1916. Otolithus (Apogonidarum) circularis rounded dorsal rim which becomes concave Shepherd C.E., Knowledge, 39, (N.S. posteriorly; posterior rim very short, obtuse 13), 183. fig. 154 (3) top two figures only. and very faintly crenulate; long, gently rounded ventral rim which is slightly undulant 1933. Otolithus (Monocentris) lemoinei Priem; at the posterior end; oblique anterior rim Burton E. St.J., Proc. Geol. Ass., 44, backed by a rounded protuberance of the 160. Name only. errore. outer face. Outer face smooth, longitudinally Material: 3 sacculiths. thickened on the ventral periphery and a Dimensions: Length 2.80 mm. Width 2.50 mm. small central depression. Smooth, convex An eroded, circular left otolith. Dorsal rim inner face with a sulcus which is virtually faintly crenulate, rising to an anterior point enclosed apart from a very indistinct, tiny and notched at its posterior end; short, groove to the anterior rim. Sulcus consisting vertical posterior rim; rounded ventral rim; of a long, elliptical ostium and a very short, rounded anterior rim which becomes oblique uncinate cauda. A lower angle present at the in its upper part. Outer face smoothly rugose junction of ostium and cauda; crista superior with a central depression across the median continuous. Sulcus partially filled with colli­ area. A somewhat convex inner face with a cula. In more eroded specimens it ·appears prominent sulcus opening widely on the ante­ that the sulcus is completely filled. The rior rim and just opening on the posterior rim smaller, less eroded right sacculith (fig, 10) also, though this latter feature may result shews crenulated dorsal, anterior and ventral from slight erosion. The sulcus is situated rims with a denticulated posterior rim. The slightly above the mid-line of the otolith and outer face is convex, with irregular central consists of a wide, oval, rather deep ostium tuberculations and prominent radial ribs on with a narrower, slightly undulant cauda. The all peripheries. It is common to find specimens undulant crista superior is markedly accent­ modified to a sharp point where the posterior uated by a depression above it. There is no rim has been lost entirely. upper angle but a prominent lower angle is This is undoubtedly the species described by present at the junction of ostium and cauda. SCHUBERT (1916) from the Bartonian of There is no evidence of a rostrum, antirostrum England. or excisura in this specimen. No collicula. Localities: The figures of SHEPHERD (1916), are vir­ Sands of Lede, Balegem, Belgium. Com­ tually unrecognisable but comparison with the mon. original specimens in the collections of the

223 Department of Palaeontology, British Museum rostrum, antirostrum and excisura are present (Natural History) prove that the present spe­ but no collicula. A smooth, semicircular lower cimen is the same species. The redescription area with a suggestion of peripheral ribbing and present figure is intended to clarify the on the ventral rim. The juvenile sacculith situation. (fig. 12) shews a markedly crenulate periphery This species closely resembles sacculiths of with prominent ribbing on the outer face. the living Australian species Trachichthodes This otolith has a strong affinity with those a/finis (Günther), differing only in its more of the living Antigonia capros Lowe, both in circular outline. This feature also distinguishes its contours and the characters of the sulcus, it from the Yprésien species T. lemoinei and is undoubtedly referable to this genus. (Priem). Localities: Localities: Sands of Lede, Balegem, Belgium. Sands of Lede, Balegem, Belgium. Basement bed of the sands of Lede, Bale­ Basement bed of the sands of Lede, Bale­ gem. gem. Sands of Aalter, Aalter, Belgium. Lower Bartonian, England. Upper Bracklesham Beds, England. Lutétien, France. Order Scorpaeniformes Family Antigoniidae Su border Platycephaloidei Genus Antigonia Lowe, 1843. Family Platycephalidae 1843. Antigonia Lowe R.T., Proc. Zoo!. Soc., Genus Platycephalus Bloch, 1795. 11, 85: orthotype Antigonia capros 1795. Platycephalus Bloch M.E., Naturg. Lowe. Ausl. Fische., 9, 96: type Callionymus ANTIGONIA ANGUSTA sp. n. Pl. figs. indicus Linnaeus = Platycephalus spa­ 12, 13. thula Bloch = Cottus insidiator Fors­ Holotype: Left sacculith, fig. 13. Registration kâl. No. Ht. p. 1593. Non Platycephalus Ribeiro, A. de M. Material: 2 sacculiths. 1902., 7: type Sciaena undecimalis Dimensions: Fig. 12. Length 1.85 mm. Width Bloch. 2.51 mm. 13. Length 2.49 mm. Width PLATYCEPHALUS ACULEATUS sp. n. Pl. 2.90 mm. fig. 14. Holotype (fig. 13), an eroded, biconvex left Holotype: Registration No. Ht. p. 1594. sacculith. High, rounded dorsal rim which is Material: 3 sacculiths. irregularly crenulated; short, slightly rounded, Dimensions: Length 2.99 mm. Width 1.34 mm. obtuse, faintly crenulated posterior rim; A slender, elliptical left sacculith. Rounded, deeply rounded, angular, denticulated ventral irregularly crenulated dorsal rim continuous rim; rounded, notched anterior rim. Outer with the crenulated posterior rim; posterior face ornamented with irregular ridges. Smooth point slightly truncated and crenated; slightly inner face with a sulcus slightly above the undulant, almost horizontal, crenated ventral mid-line, opening on the anterior rim and rim; oblique, crenated anterior rim. Concave almost extending to the posterior point of the outer face, longitudinally thickened in the otolith. Sulcus consisting of a wide, spatulate, central area with short, tuberculated ribs on deep ostium set diagonally, and an horizontal, ail the peripheral areas. Convex inner face slightly arcuate, narrow cauda of moderate with a narrow, ahnost straight sulcus opening depth, tapering at its extremity. There is a obliquely on the anterior rim and following a very slight rounded upper angle and a marked slightly upward course a little above the mid­ lower angle at the junction of ostium and line of the otolith to terminate well short of cauda. Crista superior undulant and strongly the posterior rim. It consists of a long, narrow accentuated by a depression above it. A small ostium only slightly wider than the slightly

224 undulant cauda which turns downwards moderate lower angle at the junction of ostium slightly at its tip. A very indistinct, obtuse and cauda. A shallow depression above the upper angle and a small lower angle are crista superior, accentuating it, and this de­ present at the junction of ostium and cauda, pression is traversed by radial ribbing in these features being lost in eroded specimens. unworn specimens. A blunt rostrum, slight A small rostrum, very slight antirostrum and antirostrum and excisura are present and excisura are present but no collicula. Crista collicula partially fill both ostium and cauda, superior accentuated by a depression above the two being separated by a slight diagonal it. A smooth lower area with very short groove. There is a smooth, semicircular lower ribbing on the ventral periphery. Posterior area. end of otolith compressed and rugose. The rectangular ostium distinguishes this These otoliths have a strong affinity with form from other described species which all sacculiths of the living Platycephalus indicus have an oval ostium. (Linn.), and P. bassensis Cuv. & Val., in their Locality: peripheral contours and the characters of the Sands of Lede, Balegem, Belgium. Com­ sulcus. mon. Localities: Basement beds of the sands of Lede, Bale­ Sands of Lede, Balegem, Belgium. gem. Basement beds of the sands of Lede, Bale­ Family Ambassidae gem. Genus Ambassis Cuvier & Valencien­ Upper Bracklesham Beds, England. nes, 1828. Calcaire grossier, France. 1828. Ambassis Cuvier G. & Valenciennes A., Ortler Perciformes Hist. Nat. Poiss., 2, 175: type Centro­ Su border Percoidei pomus ambassis Lacepede ( = Ambassis Family Apogonidae commersoni Cuv. & Val. Genus Apogon Lacépède, 1802. AMBASSIS ELECTILIS sp. n. Pl. fig. 15. 1802. Apogon Lacépède B.G., Hist. nat. Holotype: Registration No. Ht. p. 1598 Poiss., 3, 411: type Apogon ruber Lacé­ Material: 3 sacculiths. pède. Dimensions: Length 2.34 mm. (incomplete), APOGON ARAMBOURGI Pl. figs. 16a,b. Width 2.01 mm. Holotype: Registration No. Ht. p. 1595. A thin, rather angular right sacculith with Material: 21 sacculiths. part of the rostral area missing. Short, cre­ Dimensions: Length 5.54mm. Width4.54mm. nulate dorsal rim; nearly vertical, slightly A slightly eroded left, oval sacculith with a concave posterior rim which forms a slight bluntly rounded, crenulated posterior end. rounded « boss » at its junction with the Dorsal, posterior and ventral rim continuous dorsal rim; relatively deep, angular, promi­ and crenulated; obtuse anterior rim with a nently denticulated ventral rim; dorsal part small indentation where the sulcus opens. of anterior rim crenulated. Outer face Outer face strongly ornamented with radial thickened centrally, with tuberosities on the ribbing on the peripheral areas, an irregul­ dorsal periphery and short, radial ridges on arly raised central umbo and several verrucose the ventral periphery. Slightly convex inner projections. Convex inner face with a pro­ face with a median sulcus opening on the minent median sulcus opening obliquely on anterior rim and terminating short of the the anterior rim and not reaching the poste­ posterior rim. Sulcus consisting of a short, rior point on the otolith. Sulcus consisting of deep, spatulate ostium and a straight, narrow, a rather rectangular, wide, shallow ostium shallower cauda which is rounded at its equal in length to the narrower cauda. There extremity. Crista superior continuous, almost is a concavity of the crista superior and a horizontal and somewhat accentuated by a

225 depression above it. A marked lower angle which is smoothly tuberculated in the central at the junction of ostium and cauda. Slight area and is delicately ornamented with short antirostrum and excisura. Rostrum probably radial ribbing on the peripheral areas. Convex present in entire specimens. No collicula. inner face with an undulant sulcus opening Ventral periphery ornamented with short narrowly on the anterior rim and ending near parallel ribs. the postera-ventral point of the otolith. This ornate sacculith has similar characters Sulcus consisting of a moderately long ostium, to those of the living Ambassis macleayi tapering towards the cauda from which it is (Castelnau), differing in its crenulated rims, separated only be a constriction of the crista more circular outline and truncated posterior superior and crista inferior, both of which rim. are continuous and undulant. Also, the ostium Localities: is slightly deeper than the cauda which is Sands of Lede, Balegem, Belgium. arcuate and tapers at its extremity. Crista Basement bed of the sands of Lede. superior accentuated by a depression above Upper Bracklesham Beds and Lower Bar- it which is traversed by indistinct radial, ton Beds, England. scalloped ribs to the dorsal rim. Moderate Family Pseudopriacanthidae rostrum, antirostrum and excisura present. Genus Pseudopriacanthus Bleeker, 1869 No collicula. A deep, triangular, smooth lower area with short, parallel ridges on the 1869. Pseudopriacanthus Bleeker P., Vers!. ventral periphery. In eroded examples the Akad. Amsterdam (2), 3, 241 : ortho­ contours of the otolith become much more type Priacanthus niphonius Cuv. et Val. rounded. PsEUDOPRIACANTHUS RUTOTI (Leriche) These sacculiths are almost identical with Pl. fig. 17. those of the living Pseudopriacanthus altus 1905. Sparidarum rutoti Leriche M., Mém. Gill, differing only in the more rounded and Mus. Roy. d'Hist. Nat. Belg., 3, 165. markedly denticulate dorsal rim with the Pl. 12, figs. 9,10. somewhat concave posterior rim seen in the 1916. Otolithus (Pagellus ?) gregarius Koken; living type. Leriche supposed this form to be Schubert R.J., Jahrb. der k.k. geol. referable to the group Sparidarum, as defined Reichsanst., 65, 278. Pl. 7 figs. 14, 14a. by Koken, but was obviously not familiar (errore). with the sacculiths of Pseudopriacanthus. 1933. Otolithus (Psetta) praemaxima She­ Localities: pherd; Burton E.St.J., Proc. Geol. Ass., Sands of Lede, Balegem and sands of Brus­ 44, 160. Name only. errore. sels, Belgium. Uncommon. Basement bed of the sands of Lede. 1934. Otolithus (Trigla) rhombicus Schubert; Upper Bracklesham Beds, England. Frost G.A., Ann. Mag. Nat. Rist., (10), Calcaire grossier, and Auvers formation, 14, 442. pl. 12 fig. 1. errore. France. Material: 12 sacculiths. Family Serranidae Dimensions: Length 2.96 mm. Width 2.45 mm. Genus Serranus Cuvier, 1817. A well-preserved left sacculith with several 1817. Serranus Cuvier G., Règne Anim., Ed. features not seen in the eroded holotype of 1, 2, 276: type Perca cabrilla Linn., by Leriche. This otolith is subquadrilateral and common consent. thin. The rounded dorsal rim is crenulated; subgenus Serranellus Jordan & Eigen­ posterior rim short, obtuse, with two crenu­ mann, 1888. lations on its upper half; the markedly angu­ lar, deep ventral rim is keeled and prominently 1890. Serranus (Serranellus) Jordan D.S. in denticulated; anterior rim notched and conti­ Jordan D.S. and Eigenmann C.H., Bull. nuous with the dorsal rim. Concave outer face U.S. Fish. Comm., 8, (1888), 399:

226 orthotype Perca scriba Linn. CENTROPRISTES EXSCULPTUS sp. n. Pl. SERRANUS (SERRANELLUS) DELICATULUS fig. 19. sp. n. Pl. fig. 18. Holotype: Registration No. Ht. p. 1597. Holotype: Registration No. Ht. p. 1596. Material: The unique holotype. Material: The unique holotype. Dimensions: Length 4.78 mm. (incomplete) Dimensions: Length 3.18 mm. (incomplete) Width 2.89 mm. Width 1. 72 mm. An elegant, thin, elliptical right sacculith. Strongly denticulated dorsal and posterior An elliptical, thin left sacculith with the tip rims which are continuous; long, curved, of the rostrum missing. Very slightly curved, denticulated ventral rim, the denticulations broadly bi-lobed dorsal rim which is notched being more strongly marked at the posterior at its junction with the obtuse, short, tri-lobed point; oblique, denticulate, notched anterior posterior rim; long, slightly rounded, denti­ rim. Concave outer face strongly ornamented culated ventral rim; long, oblique, undulant with an irregular network of internai ridges anterior rim. Strongly concave outer face and radial ribbing on all rims. Convex inner ornamented with broad, radial, pectinated face with a median, arcuate sulcus opening ribbing on the dorsal area. Convex inner face on the anterior rim and terminating near the with an arcuate sulcus opening obliquely on posterior end of the ventral rim. Sulcus the anterior rim and terminating before consisting of a relatively long, narrow, rec­ reaching the posterior point of the otolith. tangular ostium and a narrower, curvedcauda. Sulcus consisting of a rather narrow, mode­ A rounded upper angle and an obtuse lower rately deep ostium and a narrower cauda angle are present at the junction of ostium which is undulant and turns downwards near and cauda. The crista superior is accentuated its rounded extremity. An undifferentiated, by a depression above it and the dorsal area undulant crista superior which is accentuated is traversed by wide, parallel ridges. It is by a shallow depression above it. Two broad likely that the prominent rostrum is continued ridges traverse the depression to the dorsal to a sharp point but the tip has broken off. periphery. An obtuse lower angle is present A slight antirostrum and excisura are present at the junction of ostium and cauda. Promi­ but no collicula. The posterior end of the nent rostrum and excisura present but no otolith is compressed and prominently sculp­ visible antirostrum or collicula. The posterior tured. A smooth, semicircular lower area. end of the otolith is compressed and rugose. This otolith compares closely with sacculiths Smooth, semicircular lower area and a nar­ of the living Centropristes striatus (Linn.), in rowly keeled ventral periphery. its peripheral contours and characters of the This ornate otolith is typically serranid in sulcus, differing only in the more elegant character and compares closely with sacculiths sculpturing of the dorsal and ventral rims. of the living Serranellus scriba (Linn.), in its ornamentation and sulcus characters but is Locality: Sands of Lede, Balegem, Belgium. relatively shorter and broader in its contours. Family Lutianidae Genus Lutianus Bloch, 1790. Locality: 1790. Lutianus Bloch M.E., Naturg. Ausl. Sands of Lede, Balegem, Belgium. Fische, 4, 105 : type Lutianus lutianus Basement bed of the sands of Lede. Bloch. Also spelled Lutjanus. Genus Centropristes Cuvier, 1829. LUTIANUS KOKENI (Leriche) Pl. fig. 20. 1829. Centropristes Cuvier G., Règne Anim., 1905. Percidarum kokeni Leriche M., Mèm. Ed. II, 2, 145. (Cuv. et Val., 3, 56): Mus. Roy. d'hist nat. Belg., 3, 162. type Centropristes nigricans Cuv. et Val. pl. 12 figs. 1-8. (Labrus striatus Linn.). Later written 1906. Otolithus (Percidarus) concavum Priem Centropristis. F., Bull. Soc. géol. Fr., (4), 6, 269.

227 figs. 16-18 only. Localities: Non Bauza Rullan J., 1957. Sands of Lede, Balegem, Belgium. Rare. 1906. Otolithus (Percidarum) obtusus Priem Upper Bracklesham and Barton Beds, F., !oc. cit., 270, text figs. 21-23 only. England. 1911. Otolithus (Percidarum) concavus Priem Lutétien and Bartonien, France. F., Ann. Pal. Soc., suppl., 28-29. text­ Family Sparidae figs. 26, 27. Subfamily Denticinae 1912. Otolithus (Percidarum) concavus Priem Genus Dentex Cuvier, 1815. F., Bull. Soc. géol. Fr., (4), 12, 246. 1815. Dentex Cuvier G., Mem. Mus. Rist. Name only. Nat. Paris. 1 (6), March, 456: 1817. 1912. Otolithus (Percidarum) concavus Priem Regne Anim., II, 273: type Sparus F., !oc. cit., 248. text-figs. 7,8. dentex Linn. 1913. Otolithus (Percidarum) aff. concavus DENTEX NOTA (Shepherd) PI. figs. 21,22. Priem F., !oc. cit., (4), 13, 152. text­ 1915. Otolithus (Pagellus ?) gregarius Schu­ figs. 2,3. bert R.J., Jahrb. der k.k. Reichsanst., 1916. Otolithus (Percidarum) bartonensis 65, 278. Taf. 7 figs. 12,13. errore. Schubert R.J., Jahrb. der k.k. geol. Otolithus (Berycidarum) bouryi Schu­ Reichsanst., 65, 278. pl. 7 figs. 9-11. bert R.J., !oc. cit., 280. fig. 22. 1916. Oto/ithus (Serranidarum) bartonensis 1916. Otolithus (Apogonidarum) nota She­ Shepherd C.E., Knowledge, 39, pherd C.E., Knowledge, 39, n.s. 13, Otolithus (Serranidarum) concavus (N.S. 183. fig. 154 (6). Bottom two figures. 13), 184. fig. 154 (8). Otolithus (Pagel/us) praecentrodontus 1933. Otolithus (Serranus) bartonensis Schu­ Shepherd C.E., loc. cit., 180. fig. 155 bert; Burton E.St.J., Proc. (la). Otolithus (Serranus) concavus Priem, Otolithus (Sargus) pre-rondolettii She­ Geol. Ass., 44, 160. Name only. errore. pherd C.E., loc. cit., 180. fig. 155 (lb). 1934. Otolithus (Percidarum) concavus Priem; 1933. Otolithus (Serranus) concavus Burton Frost G.A., Ann. Mag. Nat. Rist., E.St.J., Proc. Geol. Ass., 44, 160. name (10), 13, 426. pl. 15 fig. 1. only. 1934. Otolithus (Percidarum) bartonensis 1934. Otolithus (Dentex) pu/cher Frost G.A., Priem; Frost G.A., ibid. 426. pl. 15, Ann. Mag. Nat. Rist., (10), 13, 380. fig. 2. pl. 14 fig. 1. 1965. Allomorone bartunensis (Priem); Friz­ Otolithus (Dentex) regufaris Frost G.A., zell D.L. & Dante J.H., Journ. Pal., loc. cit., 384. pl. 14, fig. 12. 39, no. 4, 703. Name only. Material: 2 specimens. 1968. Lutianus concavus Stinton F.C., Mèm. Dimensions: Fig. 21. Length 5.14 mm. Width B.R.G.M., No. 58, 158. pl. (unnum­ 3.80 mm. Fig. 22. Length 2.83 mm. Width bered) fig. 19. 1.97 mm. Material : One fragmentary sacculith. Figure 21 represents an eroded left saccu­ Although fragmentary, sufficient of the lith. Dorsal rim very faintly crenulated; specimen remains to identify it as typical of posterior rim missing; a somewhat deeply the species described by LERICHE (1904) 1905. rounded, crenulated ventral rim; obtuse It occurs in vast numbers in the upper Eocene anterior rim. A concave outer face with in­ of France and England. The many synonyms distinct tuberosities and a suggestion of radial result from the degrees of erosion or the ribbing on the dorsal area. A convex inner growth-stage of the various specimens cited face with a median sulcus opening on the as holotypes. anterior rim and terminatin_g near the postero-

228 ventral corner of the otolith. Sulcus consisting cauda. An undulant crista superior accentuat­ of a wide, spatulate ostium, slightly deeper ed by a depression above it and this is tra­ than the narrow, arcuate cauda which tapers versed by wide, tuberose ribs. A prominent to a point at its extremity. There is an obtuse rostrum is probably present in entire specimens upper angle and a marked lower angle at the but it is lost in this example. Antirostrum and junction of ostium and cauda. Crista superior excisura present but no collicula. A smooth, accentuated by a depression above it. Rostral semicircular lower area. Posterior end of area somewhat eroded. No collicula. Fig. 22 otolith compressed and traversed by three represents a juvenile specimen and shows an ridges to the posterior rim. oval outline with a suggestion of a blunt This sacculith is closely similar to those of rostrum but part of the rostrum is missing. the living species Lagodon rhomboides (Linn.), These otoliths show a strong resemblance differing only in the fewer, wider denticula­ to the form described by SHEPHERD (1916), as tions of the posterior rim and the widely far as can be judged from such eroded spe­ crenulated dorsal rim. cimens. Locality: Sands of Lede, Balegem, Belgium. Localities : Subfamily Pagellinae Sands of Lede, Balegem, Belgium. Gen'.ls Oblada Cuvier, 1829. Basement bed of the sands of Lede, Bale­ gem. 1829. Oblada Cuvier G., Règne Anim., Ed. II, Upper Brackleshem Beds and Lower Bar­ 2, 185: type Sparus melanurus Linn., ton Beds, England. monotypic. ÜBLADA JONETI sp. n. Pl. fig. 24. Subfamily Sparinae Genus Lagodon Holbrook, 1855. Holotype: Registration No. Ht. p. 1600. Material: The unique holotype. 1855. Lagodon Holbrook J.E., Ichthyology of Dimensions: Length 3.70 mm. (incomplete) S. Carolina. 59: orthotype Sparus rhom­ Width 2.58 mm. boides Linn. LAGODON PECTINOIDES sp. n. Pl. fig. 23. An eroded, elliptical, left sacculith. Round­ ed, faintly crenulate dorsal rim; short, obtuse Holotype: Registration No, Ht. p. 1601. posterior rim; rounded, smooth ventral rim; Material: The unique holotype. oblique, broken anterior rim. Concave outer Dimensions: Length 3.77 mm. (incomplete) face with an indistinct central umbo and Width 2.26 mm. radial ridges to the dorsal and posterior rims. A thin, left sacculith. An irregularly crenu­ The degree of ornamentation has been lated dorsal rim, concave posteriorly; vertical, markedly affected by erosion. Ventral peri­ tridenticulate posterior rim; rounded ventral phery thickened. Convex inner face with an rim, indistinctly crenulated posteriorly; obli­ arcuate sulcus opening widely on the anterior que, prominently notched anterior rim. rim and terminating near the postero-ventral Concave outer face with broad radial ribs on border. Sulcus consisting of a fairly deep, the dorsal and posterior areas and a conceu­ spatulate ostium and a narrower, arcuate tric growth line paralleling the periphery. cauda. The rims of the ostium become straight Convex inner face with a sulcus opening on towards the anterior end but, this feature is the anterior rim and extending somewhat largely lost in the present incomplete spe­ diagonally to terminate near the postero­ cimen. The crista superior is markedly accent­ ventral corner of the otolith. Sulcus consisting uated by a depression above it. Marked upper of a fairly deep, somewhat oval ostium and and lower angles at the junction of ostium a rather wide, undulant cauda which turns and cauda. Characters of rostral area missing. downwards at its extremity. A very slight, No collicula. Smooth lower area. rounded upper angle and an obtuse lower This eroded otolith somewhat resembles angle present at the junction of ostium and sacculiths of the living Oblada melanura

229 (Linn.), especially in the character of the extensive MS. species to make a tentative sulcus, but the fossil form is relatively wider correlation with the geological horizons in and shorter. other areas. Locality: Sands of Lede, Balegem, Belgium. From the below tables it is evident that the Basement beds of the sands of Lede, com­ teleosts of the sands of Lede have a close mon. affinity with those of the Upper Bracklesham beds of England. There is also evidence that All the described specimens are lodged in this fauna is comparable with that of the the collections of the Musée Royal d'Histoire Lower Barton beds, and of the Calcaire Naturelle de Belgique and bear the registra­ Grossier but this may be due to a more tion numbers. 24/181 of the general inventary extensive vertical distributions of some spe­ of that Institution. cies. A more intensive investigation of the sands of Brussels and Auvers would un­ Discussion doubtedly produce further evidence in this field of research. However, it is quite evident Although much work still remains to be done that few species are common to the N um­ on the teleost faunas of the zones of the mulites laevigatus zone of England and the Eocene of Belgium and France it is possible sands of Lede. Nevertheless it must be noted to establish certain relationships between them. that the sands of Aalter which are probably of the same age (or less older) as the N. Points oj Comparison laevigatus beds of England, have five species in common with the sands of Lede. Belgian Basin Sorne Biological Characteristics of the Teleosts from the Sands of Lede A sufficient number of species for comparative Dredging experiments today reveal that heavy purposes have been obtained only from the concentrations of otoliths are to be found at sands of Aalter. Only three species (as oto­ depths of not less than 40 metres and investi­ Iiths) are known from the sands of Brussels, gations of bottom deposits off the Woods two of which occur in the sands of Lede. Hole area of the east coast of the United No otoliths have so far been recorded from States revealed a marked concentration of the sands of Wemmel. sacculiths of the ophidioid Lepophidium cer­ vinum GoonE & BEAN, among the numerous Paris Basin otoliths. This parallels the abundance of the Otoliths have been described by Priem and sacculiths of Lepophidium aequalis sp. n., in Leriche from the Paris Basin and one of us the Lédien deposits. The absence of clupeoid otoliths from this ~F.C.S.) has collected from a number of horizons of the French Eocene. A reasonable assemblage is a common feature of Eocene fauna is known from the Sables de Cuise and strata. One of us (F.C.S.) has recovered from the Lutétien but otoliths are scarce in occasional sacculiths of several extant genera the sands of Auvers and Marines. This of clupeids from various horizons of the scarceness probably results from the very English Upper Bracklesham Beds, thus littoral or beach-like character of this facies. establishing the presence of this large group It is evident that further investigations are of teleosts in the seas of Lede. required before firm conclusions on correla­ tion with other areas can be made. Acknowledgements Hampshire and London Basins Dr. c. PATTERSON and Mrs. WELCH, of the Many otoliths have been described from the Department of Palaeontology, British Mu­ English Eocene and one of us (F.C.S.) has seum (Natural History) h;j-ve accorded most

230 Ledien species common to other 1 2 3 4 5 6 7 8 9 10 11 12 formations

Sables d 'Aalter Sables de Bruxelles • Calcaire grossier --· • Sables d 'Auvers • Bracklesham beds (with N.laevig?tus) • Bracklesham beds (with N.variolarius) • Lower Barton beds •

Table of the stratigraphical distribution of the otolith based teleosts from the sands of Lede

1

1 -ti 1 1 ,.... 1 -0"' -ti E: .... -ti f!l <::! -ti ...... -0 ~ ·21 ::: -0"' b'li ~ "' .::::: "1::! {! tl ,.. 1 ..:!: ::: <::!"' l E: ·~ ~ ,:: -0 "'::::: .;, ""'<: ~ "' -.:i"' ""'<: <:i ro ...... ~ ~~ ~ ~ ~ ::: ~· "<:! -0 ~ -~ ..:!: z ...... -ti ~ ~~ -ti <:l .;:: -t; -5 ::: ~ -.:i ::: ~ "' "' "' 1 tl ;,::;~ <:l ~ ~ ~~ 1 ~ a ""'<: ~~ -.:i"' 1 ! 1 1 1 A nguilla rectangularis n. sp. L Up B 1 Gymnothorax diagonalis n. sp. A (L) L G Up u oringua fissura n. sp. L p araconger sauvagei (PRIEM) A (L) L G A Lo Up B s ynodus rel!i.s (FROST) L Up B R oplobrotula lerichei n. sp. (L) 1 L G L epophidrum aequalis n. sp. (L) L 1 Lo Up Ogilbia ~ubregularis (SCHUBERT) A (L) L G L<. Up B Trachichthodes circularis (SHEPHERD) (L) L G B A ntigonia angusta n. sp. A (L) L Up p latycephalus aculeatus n. sp. (L) L G Up A pogon arambourgi n. sp. (L) L p seudopriacanthus rutoti (LERICHE) Br (L) L G A Up serranus delicatulus n. sp. (L) L c entropristes exsculptus n. sp. L A mbassis electilis n. sp. (L) L Up B L utianus kokeni (LERICHE) A Br L G A Up B D entex nota (SHEPHERD) (L) L Up B Oblada joneti n. sp. (L) L L agodon pectinoides n. sp. L valuable advice on the format and relevant throughout the world who have supplied the literature in this paper. Most grateful thanks necessary fish and otoliths, without which this are extended to all institutions and people paper could not have been prepared.

231 EXPLANATION OF PLATE

Fig. 1. Anguilla rectangularis sp. n. Right sacculith, inner face. Holotype x 8 2. Anguilla rectangularis sp. n. Right sacculith, inner face. Lower Barton Beds, Hor. A3, Highcliff, Hants. x 9 3. Gymnothorax diagonalis sp. n. Left sacculith, inner face. Holotype. x 8 4. a, b. Moringua fissura sp. n. Right sacculith, inner and outer faces. Holotype. x 9 5. Paraconger sauvagei (Priem). Left sacculith, inner face. x 8 6. Synodus reclus (Frost). Left sacculith, inner face. x 8 7. Lepophidium aequalis sp. n. Left sacculith, inner face. Holotype. x 9 8. Hoplobrotula lerichei sp. n. Right sacculith, inner face. Holotype. x 8 9. Ogilbia subregularis (Schubert). Left sacculith, inner face. x 8 10. Ogilbia subregularis (Schubert). Juvenile right sacculith, outer face. x 8 11. Trachichthodes circularis (Shepherd). left sacculith, inner faœ. x 8 12. Antigonia angusta sp. n. Juvenile Jeft sacculith, inner face. x 7 13. Antigonia angusta sp. n. Left sacculith, inner face. Holotype. x 8 14. Platycephalus aculeatus sp. n. Left sacculith, inner face. Holotype. x 10 15. Ambassis electilis sp. n. Right sacculith, inner face. Holotype. x 9 16. a, b. Apogon arambourgi sp. n. Left sacculith, inner and outer faces. Holotype. x 6 17. Pseudopriacanthus rutoti (Leriche). Left sacculith, inner face. x 9 18. Serranus (Serranellus) delicatulus sp. n. Left sacculith, inner face. Holotype. x 8 19. Centropristes exsculptus sp. n. Right sacculith, inner face. Holotype. x 8 20. Lutianus kokeni (Leriche). Left sacculith, inner face. x 8 21. Dentex nota (Shepherd). Left sa-::culith, inner face. x 8 22. Dentex nota (Shepherd). Juvenile right sacculith, inner face. x 8 23. Lagodon pectinoides sp. n. Left sacculith, inner face. Holotype. x 9 24. Oblada joneti sp. n. Left sacculith, inner face. Holotype. x 8

Except where indicated, all the specimens are from the sands of Lede, Balegem, Belgium.

BIBLIOGRAPHIE

BLEEKER, P. (1865). Systema Muraenorum revi­ ranée, et à leur occasion sur des Poissons sum. Ned. Tijdschr. Dierk., 2, pp. 113-122. d'autres mers plus ou moins liés avec eux. BLEEKER, P. (1869). Neuvième notice sur la faune Mém. Mus. d'hist. Nat., Paris. 1. ichthyologique du Japon. Vers!. Akad. Amster­ CUVIER, G.L. (1817). Le règne animal distribué dam. (2), 3, pp. 237-259. d'après son organisation pour servir de base à BLOCH, M.E. (1790). Naturgeschichte der Aus­ l'histoire naturelle des animaux:, et l'introduc­ landischen Fische. 4, xii + 128 pp. pls. 217-259. tion à l'anatomie comparée. 2, P6issons. 532 pp. BLOCH, M.E. (1795). Naturgeschichte der Aus­ Paris. landischen Fische, 9, 397-432. 192 pls. CUVIER, G.L. (1829). Le règne animal, 2. Poissons, BURTON, E.St.J. (1933). Faunal Horizons of the 2e éd., pp. 406. 120 pls. Paris. Barton Beds in Hampshire. Proc. Geol. Ass., CUVIER, G.L. & VALENCIENNES, A. (1828-1849). London, 44, pp. 131-167. Histoire Naturelle des Poissons. 22 vols. 650 pls. CASIER, E. (1946). La faune ichthyologique de Paris. l'Yprésien de la Belgique. Mem. Mus. Hist. nat. EVERMANN, B.W. & KENDALL, W.C. (1783). Belg., Bruxelles. 104, pp. 1-267, pls. 1-6. Descriptions of new or little known Genera CURRY, D. & WrsDEN, D.E. (1958). Geology of and Species of Fisches from the United States. the Southampton Area including the coast Bull. U.S. Fish. Comm., 17, 125-133 pp. sections at Barton, Hants and Bracklesham, FRIZZELL, D.L. & DANTE, J.H. (1965). Otoliths Sussex. Geol. Ass. Guides, London. No. 14. of some Cenowic F1shes of the Gulf Coast. CuvIER, G.L. (1815). Observations et Recherches Journ. Pal., 39, pp. 687-718. pls. 86-88. Critiques sur differents Poissons de la Méditer- FROST, G.A. (1933). Otoliths of Fishes from the

232 ~9

19 22 20 21

23

lower Tertiary Formations of Southern England. the Atlantic and Gulf Coasts. Proc. U.S. Nat· l. Isospondyli, Apodes, Berycomorphi. Ann. Mus., 8, pp. 589-605. Mag. Nat. Hist., (IO), 12, pp. 387-396. pl. 12. GRAY, J.E. (1831). Description of twelve new FROST, G.A. (1934). Otoliths of Fishes from the genera of Fish, discovered by Gen. Hard wicke lower Tertiary Formations of Southern England. in India. The Zoological Miscellany. XIII, III. Percomorphi, Scleroparei. !oc. cit., (10), pp. 7-10. 13, pp. 426-433. pl. 15. GRoNow, L.T. (1763). Zcophylacii Gronoviana ... FROST, G.A. (1933). Otoliths of Fishes from the Animalis quae in museo suo adservat, etc., lower Tertiary Formations of Southern England. 380 pp., 20 pls. IV. Scleroparei, Percomorphi. loc.

233 A review of the Genera and Species of Serra­ Vôge! und Fische. Arch. Naturg. Berlin. 23, nidae found in the waters of America and pp. 262-272. Europe. Bull. U.S. Fish. Comm., 8, pp. 329-441. PRIEM, F. (1906). Sur les Otolithes des Poissons KANAZAWA, R.H. (1961). Paraconger, a new genus éocènes du Bassin parisien. Bull. Soc. géol. Fr., with three new species of Eels (Family Congri­ (4), 6, pp. 265-280. dae). Proc. U.S. Nat. Mus., 113, pp. 1-14. 2 pls. PRIEM, F. (1911). Étude des poissons fossiles du KoKEN, E. (1891). Neue Untersuchungen an Bassin parisien (supplément). Ann. Pal., Paris. tertiiiren Fisch-Otolithen, Il. Z. dtsch. geai. 6, 1-44 pp. pis. 1-5. Ges., Berlin. 43, pp. 77-170. pls. 1-10. PRIEM, F. (1912). Sur des Otolithes éocènes de KoTTHAUS, A. (1968). Fische des Indischen France et d'Angleterre. Bull. Soc. géol. Fr., (4), Ozeans. Ergebnisse der ichthyologischen Unter­ 12, pp. 246-249. suchungen wahrend der Expedition des Forsch­ PRIEM, F. (1913). Sur des Otolithes de l'Eocène - ingsschiffes "Meteor" in den Indischen Ozean, du Cotentin et de Bretagne. Bull. Soc. géol. Fr., Oktober 1964 bis Mai 1965. Gebrüder Born­ (4), 13, pp. 151-158. traeger, Berlin. Stuttgart. 3, pp. 14-56. Abb. RoSEN, D.E. & PATTERSON, c. (1969). The 97-148. Structure and Relationships of the Paracan­ LACÉPÈDE, B.G. (1802). Histoire naturelle des thopterygian Fishes. Bull. Am. Mus. Nat. Hist., Poissons. 3, 415 pp. pls. 1-6. Paris. New York. 141, pp. 357-474. pls. 52-78. LERICHE, M. (1905). Les Poissons eocènes de la SCHUBERT, R.J. (1916). Obereocane Otolithen vom Belgique. Mém. Mus. Hist. nat. Belg., Bruxelles. Barton Cliff bei Christchurch (Hampshire). Jb. 3, pp. 49-228. pls. 4-12. geai. Reichsanst. (Bundesanst.) Wien. 65, pp. LERICHE, M. (1912). L'Eocène des bassins pari­ 277-288. pl. 7. sien et belge (Livret guide de la réunion extra­ SHAW, G. (1803). General Zoology, or Systernatic ordinaire de la société géologique de France à Natural History. 4. i-v + 1-632, pp. 92 pls. Laon, Reims, Mons, Bruxelles, Anvers, 27 SHEPHERD, C.E. (1916). Fossil Otoliths. Know­ août - 6 septembre 1912. Bull. Soc. géol. Fr., ledge, 39 (NS 13), pp. 177-184. 2 pis. 4e ser., T. 12, pp. 712-824. STINTON, F.C. (1967). The Otoliths of the tele­ LINNAEUS, C. (1758). Systema Naturae, etc. Ed. ostean fish Antigonia capros and their taxo­ 10, 1, Regnum Animale. Holmiae. 824 pp. nomie significance. Bocagiana, Mus. Mun., LOWE, R.T. (1843). Notice of Fishes newly ob­ Funchal, Madeira. pp. 1-8. pl. 1. served or discovered in Madeira during the STINTON, F.C. (1968). On the Study of Tertiary years 1840, 1841 and 1842. Proc. Zoo/. Soc., Fish Otoliths. Mém. B.R.G.M., Paris. 58, pp. London. 11, pp. 81-95. 153-162. 1 pl. McALLISTER, D.E. (1968). Evolution of Bran­ SULC, J. (1932). Otolity Paleogénu okoli Biarritz chiostegals and Classification of Teleostean (Les Otolithes du Paléogène des environs de Fishes. Bull. Nat. Mus. Canada, Ottawa. 221, Biarritz). Rozpr. geai. Ust. es!. Praze, 7, pp. Biol. ser. 77, i-xiv + 1-239 pp. 1-94. pis. 1-3. PHILIPPI, R.A. (1857). Über einige chilenische Communication présentée le 16 décembre 1969

234