LEIDSE GEOLOGISCHE MEDEDELINGEN, Deel 49, Aflevering 3, pp. 507-.537, 1-3-1976

New Gadiform Otoliths from the Tertiary of the

North Sea Basin and a revision of some fossil and recent species

BY

P.A.M. Gaemers

Abstract

Gadidae from different localities in the North Sea Basin Pliocene) are Otoliths of 10 new species, includingnine (Middle Oligocene — Upper

described: Gadus parallelus, Trisopterus concavus, T. incognitus, Colliolus parvus, C. johannettae, C. schwarzhansi, Palaeoraniceps regularis, Molva primaeva, Brosme heinrichi and ? Macruridarum deurnensis.

established: Neocolliolus and Three new genera are Protocolliolus, Palaeoraniceps.

(1884, 1891) has been studied; as a result some systematic errors are Type material ofsome fossil otoliths ofGadidae described by Koken corrected.

Pollachius carbonarius is ascertained of The existence of two recent species of coal-fishes (Pollachius virens (L.) and (L.)) by study

their otoliths.

INTRODUCTION Diagnosis. - A slender Gadus species with a distinct and

regularpattern of knobs and furrows. Dorsal rim between

Otoliths of new species of the Gadiformes have been predorsal and postdorsal angle long and nearly straight.

discovered from different localitiesin the North Sea Basin. Otoliths only slightly bent along long axis, thus inner

The materialdescribedin this publication comes from West surface somewhat convex and outer surface slightly

and the Netherlands. The in the direction of the and East Germany, Belgium new concave length. species come from various Tertiary deposits ranging from

Middle Oligocene to and including Upper Pliocene. Description. - Otoliths strong and large. Outline oval and fossil It is necessary to revise the classification of some elongated. Rather thick ventral rim is weakly and regularly and recent species of the Gadidae on the ground of their bent and has clearly visible furrows and knobs. Thin and otoliths, because of the new material at my disposal. delicate caudal end with a blunt point. Dorsal rim thin;

furrows and knobs not as pronounced as on ventral rim.

Distinct predorsal and postdorsal angles. Dorsal rim

SYSTEMATIC DESCRIPTIONS between these angles long and nearly straight. Rostral rim

axis. straight and with oblique angle on long

PISCES Phylum Inner surface slightly convex and curved along long axis.

Superclassis TELEOSTOMI (OSTEICHTYES) Sulcus acusticus long, wide and almost entirely filled with Classis ACTINOPTERYGII colliculi. Cauda twice as long as ostium. Rostral end of Subclassis TELEOSTEI ostium vague and somewhat bent toward dorsal side. Ordo GADIFORMES Ostium slightly narrower than cauda. Collum rather short

Familia GADIDAE 1810 Rafinesque, and narrow. Small furrows on dorsalpart enter the colliculi; SubfamiliaGADINAE some are furcated. Small furrows on ventral part are Genus GADUS 1758 Linnaeus, distinctbut short; they do not traverse large ventralfurrow.

Area narrow and shallow.

Gadus parallelus n. sp. Outer surface slightly concave in lengthwise direction la, 2a, (PI. I, Figs, b, b) and weakly S-shaped in the direction of the height (ventral

part convex, dorsal part somewhat concave). Entire

Type. - Holotype: PI. I, Figs, b, Coll. RGM 175800. la, surface highly sculptured, especially ventralpart. Medially

a cluster of circular knobs. Peripheral knobs in obvious

Locus typicus. - Sand-filled ground of Ford factory, pattern nearly perpendicular to the rims. Antwerp-North, Belgium.

L: 12.85 mm, H: 5.09 mm, T: 1.96 mm (holotype)

Stratum typicum. - Pliocene, upper part.

Caudalend of holotype is slightly eroded, thus true length is

Derivationominis. -Parallèlos (Greek) = parallel. Because somewhat greater. the dorsal and ventral rims are nearly parallel.

- * Material. 5 sagittas, Pliocene, Ford Antwerp- Dept. of Stratigraphy and Palaeontology, Geological and factory, North, Coll. RGM 175800, 175801, 175802. 3 Mineralogical Institute, University of Leiden, Garenmarkt lb, specimens Leiden, the Netherlands. eroded and without caudal point, 2 broken and 508

considerably eroded sagittas, basal shell bed of Sands of the shape of their otoliths. Length-height and seasluice Kallo, Kallo, Prov. East Flanders, Belgium, Coll. length-thickness ratios rarely overlap, whereas the Gaemers, Vervoenen. ratio leg. length-curvature is less distinctive. Moreover, in most

cases the ratioof the lengthoffish to the length of otolithfor

Distribution. - Upper Pliocene, Sands of Kallo. adult specimens of both species is highly characteristic: P. virens has ratio which a ranges between 32 and 45.5, and P.

Discussion. - Variation to be restricted to carbonarius seems mainly a ratio which ranges between 44 and 54. When

bend in the ventralrim clear the and the predorsal and postdorsal the very difference in colour between the black

angles, which can be more or less pronounced. coalfish P. carbonarius and the white coalfish P. virens is

It is that the best which has also considered it is a pity preserved specimen, clear that Linnaeus was right in chosen been as holotype, could not be collected differentiating between the two species and that later

The exact ofthis fossil is biologists in the in stratigraphically. age unknown, were wrong putting two together one but it certainly belongs to the Pliocene; this could be species.

determinedby otherfinds ofotoliths in the same sand-filled

ground. It cannot be of Pleistocene age because at that time Genus TRISOPTERUS Rafinesque, 1814 Gadus morhua already existed (Gaemers, 1974b). Judging

from the of the and the other two preservation holotype Trisopterus concavus n. sp.

specimens, they can derive from the Sands of Kallo or the (PI. II, Figs, la, b, 3a, b)

Sands of Luchtbal, certainly not from the Lower Pliocene

Sands of Gadus cf. Koken, Leriche 320. Kattendijk. spectabilis 1891; 1906, p.

Gadus luscus Koken spectabilis 1891; Weiler, 1942, p. 79, pi. Ill, figs. 15, 17. Genus POLLACHIUS Bonaparte, 1845 Gadus luscus spectabilis Koken, 1891; Weiler, 1959, p. 97. Gadus luscus spectabilis Koken, 1891; Zilch, 1965, p. 457. Pollachius virens (Linnaeus, 1758) Gadus luscus spectabilis Weiler, 1942; Gaemers, 1969a,p. 8, pi. I, (PI. I, Figs. 3a, b; PI. II, Figs. 2a, b, 4a, b) figs. 7a, b.

luscus Trisopterus spectabilis Weiler, 1942; Gaemers, 1969b, p.

- A Diagnosis. Pollachius species with large, elongated 73.

and relatively thick otoliths, which are bent along Trisopterus luscus distinctly spectabilis Koken, 1891; Gaemers, 1971, p. the long axis. Ventral part conspicuously thicker than 242, pi. I, fig. 7; pi. VI, figs. 4, 5.

dorsal part. Sulcus acusticus wide and straight. Cauda

Type. - everywhere of nearly equal width. Ventral rim straight to Holotype: PI. II, Figs, la, b, Coll. RGM 175822.

somewhat convex in adult specimens. Caudalpart not (and

old Locus - Miste the in very specimens sometimes slightly) twisted typicus. near Winterswijk, Netherlands.

perpendicular to the long axis.

Stratum typicum. - Middle Miocene, Hemmoorian.

L: 21.77 mm, H: 8.91 mm, T: 4.34 mm (L of fish: 89 cm)

Derivatio - L: 18.49 nominis. Namedafter the concave caudal part mm, H: 7.68 mm, T: 3.36 mm (L of fish: 64 cm) of the dorsal rim. L: 18.49 mm, H: 7.42 mm, T: 3.56 mm (L of fish: 66 cm)

L: 18.79 mm, H: 7.00 mm, T: 3.06 mm (L of fish: 63 cm)

L: 16.90 H: - A thick and mm, 6.24 mm, T: 3.03 mm (L of fish: 56 cm) Diagnosis. high Trisopterus species of medium L: 15.48 mm, H: 6.38 mm, T: 2.98 mm (L of fish: 49 cm) size, which is distinctly bent along the long axis. Caudal part ofdorsal rim concave. Rostral rim with distinct

Knobs Pollachius carbonarius (Linnaeus, 1758) oblique angle on long axis. on outer surface of normal size. (PI. I, Figs. 4a, b, 5a, b, 6a, b)

Description. - Strong, thick and high otoliths of medium Diagnosis. - A Pollachius species with large, clearly size. Outline elongated. Ventral rim and rather thin thick, blunt, regularly elongated otoliths, which are distinctly bent bent and obviously sculptured. Caudal end clearly and along the long axis. Ventral part only slightly thicker than much dorsal sharply pointed. Dorsal rim thinner than ventralrim part. Sulcus acusticus very wide and bent with

but still rather blunt: dorsal rim s-shaped with a smooth convex side toward dorsal rim. Cauda widens distinctly at concave part near caudal end and a slightly sculptured caudal end. Ventral rim straight to concave in adult

convex part near rostral end. Rostral rim with distinct specimens. Caudal part of adult specimens conspicuously oblique angle on long axis; this rim is nearly with bent perpendicular to the long axis. straight two large, low knobs and a shallow depression in between. Inner surface L: markedly convex. Sulcus acusticus long, 21.27 mm, H: 6.85 mm, T: 2.58 mm (L of fish: 98 cm) L: 20.88 H: wide and shallow. Cauda somewhat than twice mm, 7.50 mm, T: 3.09 mm (L offish: 101 cm) very more

L: 20.74 H: 7.68 T: ostium. mm, mm, 3.29 mm (L offish: 103 cm) as long as Collum narrow and very short. Sulcus

L: 16.50 H: mm, 6.22 mm, T: 2.48 mm (L of fish: 82 almost filled with colliculi. cm) entirely Cristae very L: 17.78 mm, H: 6.29 mm, T: 2.64 mm (L offish: 78 cm) insignificant. Area very shallow and long. Ventral furrow clear. Small furrows and knobs from ventral rim up to

Discussion. - be Both species can easily distinguished by ventral furrow. 509

Outer surface distinctly concave, especially along dorsal Paralectotype . - 1 sagitta, Gadus spectabilis, Upper

rim. Thickest part of otolith near ventral and rostral rims. Miocene, Langenfelde, Ot. 55, Coll. Koken (PMHU).

thin. Middle of Ventral part very thick, dorsal part part

otolith s-shaped along height axis. Big knobs and deep Diagnosis. - A very thick and robust Trisopterus species.

to the Cauda furrows over the entire surface and perpendicular very wide along entire length and ostium much

rims. central knobs. obvious Medially some smaller, especially near collum; thus an jump is

visible on the ventral side of the sulcus near the collum.

L: 9.58 H: 4.41 T: 3.02 mm (holotype) mm, mm, Inner surface markedly convex along long axis. Outer

L: 9.76 mm, H: 4.62 mm, T: 3.28 mm surface in first sketch slightly concave along long axis (in

L: 9.55 mm, H: 4.46 mm, T: 3.07 mm juvenile specimens), but in the adult stage flat because of L: 8.99 mm, H: 3.94 mm, T: 2.91 mm the presence of large knobs. These knobs are much larger L: 8.42 mm, H: 3.70 mm, T: 2.36 mm than in other known 2.08 species. L: 7.55 mm, H: 3.24 mm, T: mm

L: 7.03 mm, H: 3.04 mm, T: 1.90 mm

- Otoliths thick and robust. Outlineoval L: 5.08 mm, H: 2.21 mm, T: 1.26 mm Description. very

L: 4.50 mm, H: 1.95 mm, T: 1.04 mm and elongated. Very thick ventral rim is regularly bent. Thinner, but still rather thick dorsal rim weakly and

Material. - 106 sagittas, Miste near Winterswijk, the regularly bent and with distinct predorsal angle. Ventral

Netherlands, Miocene, Hemmoorian, Coll. Gaemers, leg. rim with highly sculptured knobs and furrows. Dorsal rim

A. F. J. Janssen. 1 sagitta (holotype), Miste, Miocene, more or less smooth. Rostral rim straight and with oblique 175822. 23 axis. Hemmoorian, Coll. RGM sagittas, angle on long

'Flachbohrung 37', Oploo, the Netherlands, Upper Inner surface highly convex along long axis. Bend most

Miocene (Langenfeldian), Coll. RGM 175823, 175824, leg. pronouncedat caudalend. Sulcus acusticus long, wide and

RGD. 2 sagittas, Langenfeldian, Pinneberger Schichten, straight, almost completely filled with colliculi. Cauda 2 to

boringSchenefeld HWW9, 80mand88-90m, Coll. GLSH. 2.5 times the length of ostium. Cauda very wide; ostium

14 sagittas, bed of Königsbach creek near Königsmühle, narrower, especially near collum. Dorsal side of sulcus

Dingden, Western Germany, Dingdener Schichten, straight; ventral side with an obvious jump at collum.

RGM Feinsand, Reinbekian, Coll. 175876, leg. Collum limited to a narrow groove on dorsal side,

A. W. Janssen, 1970. somewhat wider on ventral side. Most fully developed cristae along ostium. Area narrow and long. Distinct

Distribution. - Miocene: Hemmoorian, Reinbekian and ventral furrow forms the utmost limit of the small furrows

Langenfeldian. and knobs. Sometimes these furrows are furcated in the

middle part of ventral rim.

- luscus Discussion. This species resembles Trisopterus Outersurface slightly concave along long axis injuvenile

and (L.) very closely. Length-height length-thickness specimens but flat inadult ones because ofthe big knobs; it

but T. ratios are almost the same, nevertheless concavus is convex in the directionof the height. The knobs are few

adult in appears more slender as far as specimens are number but large and very pronounced; they form a

concerned. T. concavus seems to be much less variable rough and rather irregular pattern. On the ventral side

of luscus than the recent species. Juvenile specimens T. knobs are the highest. Thickest part of otoliths often

caudal and have a very sharp point a relatively high rostral characteristically shifted to caudal part. Few or no central

part. Juvenile specimens of T. concavus have a more knobs.

well-balanced outline and a somewhat more compact

T. L: 9.60 H: 3.88 T: 2.79 mm appearance than the juvenile otoliths of luscus. mm, mm, (Deume)

L: 7.42 mm, H: 3.07 mm, T: 2.01 mm (Deume)

T. L: 7.27 mm, H: 3.03 mm, T: 1.66 mm (Oploo) - with Remarks. Up to now this species was confused T: L: 7.36 mm, H: 3.02 mm, 1.92 mm (Oploo) spectabilis (Koken) because the figures in the original L: 8.14 mm, H: 3.47 mm, T: 2.41 mm (Oploo) publication of Koken (1891) were misleading. L: 9.43 mm, H: 3.89 mm, T: 2.81 mm (Oploo)

L: 9.48 mm, H: 4.01 mm, T: 3.13 mm (Oploo) Trisopterus spectabilis (Koken, 1891) L: 9.50 mm, H: 3.98 mm, T: 2.82 mm (Oploo) (PI. II, 5, 6a, PI. Ill, la, b-4a, b, c) Figs. b; Figs, L: 9.62 mm, H: 3.90 mm, T: 3.02 mm (Oploo)

L: 9.68 mm, H: 4.11 mm, T: 3.06 mm (Oploo)

L: 9.85 mm, H: 4.12 mm, T: 3.07 mm (Oploo) Otolithus (Gadus) spectabilis Koken, 1891, p. 94, pi. Ill, figs. 3a, L: 10.09 mm, H: 4.25 mm, T: 2.91 mm (Oploo) b, 4. L: 10.15 mm, H: 4.70 mm, T: 3.43 mm (Oploo)

L: 10.30 mm, H: 4.56 mm, T: 3.46 mm (Oploo) Type. - Lectotype: PI. Ill, Figs. 3a, b, c, Coll. PMHU, Ot. L: 10.42 mm, H: 4.71 mm, T: 3.74 mm (Oploo) 55, Coll. Koken. L: 9.36 mm, H: 4.55 mm, T: 3.81 mm lectotype (eroded)

L: 8.99 mm, H: 3.89 mm, T: 2.46 mm paralectotype

Locus typicus. - Langenfelde, Holstein, Bundesrepublik

Deutschland. Material. - 2 sagittas. Upper Miocene, Langenfelde, Coll.

Koken (PMHU). 5 sagittas, Upper Miocene, Sands of

Stratum typicum. - Upper Miocene, Langenfeldian. Deurne, E-3 motorway, Borgerhout near Stenen Brug, 510

Antwerp, Belgium, Coll. RGM 175837, 175838, 175839. 46

sagittas, Upper Miocene, Langenfeldian, 'Flachbohrung

37', Oploo, eastern part of North Brabant, Netherlands,

Coll. RGM 175825, 175826, 175827, leg. RGD, Haarlem. 1

eroded sagitta, Ellewoutsdijk, Westerschelde, Zeeland,

Netherlands, Coll. Gaemers, leg. Cadée. 1 sagitta,

Langenfeldian, Pinneberger Schichten, boring Schenefeld

1 HWW 9, 88-90 m, Coll. GLSH. sagitta, Langenfeldian-

Gramian, PinnebergerSchichten, boring KatharinenhofIII

near Wedel, 161-164 m, Coll. GLSH.

Distribution. - Upper Miocene: Langenfeldian and

Gramian.

Remarks. - The specimen in the Westerschelde is the first

evidence of the occurrence of Upper Miocene otoliths in

this area. The specimens described as Gadus luscus

spectabilis Koken by Weiler(1942, 1959), Zilch (1965) and

Gaemers (1969a, b, 1971), and as Gadus cf. spectabilis by

Leriche (1906), belong to another Trisopterus species,

which is described here as the new species T. concavus.

Trisopterus luscus (Linnaeus, 1758)

(PI. Ill, Figs. 5a, b)

Otolithus (Gadus) anglicus Koken, 1891,p. 94, pi. Ill, figs. 8, 9.

Trisopterus luscus anglicus (Koken); Gaemers & Schwarzhans,

1973, p. 210, pi. IV, figs. 6a, b.

Fig. 1. a. Inner surface, b. Outer surface, c. Ventral side of right

- A thick and high Trisopterus species of Diagnosis. sagitta from Trisopterus incognitus n. sp. Magnification 10x. medium size, which is distinctly bent along the long axis.

Caudal part of dorsal rim convex or straight. Rostral rim

usually has slightly oblique angle on long axis. Knobs on - Heinrich Type. Holotype: Figs, la, b, c, Coll. (PMHU); outer surface of normal size. Inner surface markedly Heinrich (1969): pi. XI, fig. 2. to convex along long axis. Outer surface clearly concave Locus s-shaped along long axis. typicus. -Clay pit, Hohen Woos, Deutsche Demokratische Republik.

L: 11.36 mm, H: 5.38 mm, T: 3.25 mm (Gram) Stratum typicum. - Upper Miocene, Langenfeldian (see

Heinrich, 1969).

Material. - 5 sagittas,Gramian, Gram, Coll. RGM 175851,

= 175852, leg. A. W. Janssen, 13-8-1970. 2 sagittas, Sands of Derivatio nominis. - Incognitus (Lat.) unknown, un-

Deurne, Antwerp, Borgerhout near Stenen Brug, Coll. expected, not recognized. Named for the unexpected

RGM 175840, Sands of of leg. A. W. Janssen. 1 sagitta, occurrence a new Trisopterus species in the Upper

Deurne, Antwerp, Borgerhout, Foorplein, Coll. RGM Miocene.

175844, leg. F. J. Janssen. 1 sagitta, Gramian, boring Diagnosis. - A thick and high Trisopterus species of Flensburg-Feldmühle 3,69-65 m, Coll. GLSH. 13 sagittas, medium size, which is not or only very slightly bent along 1 Gramian, boring Pinneberg-Elz, 84-92 m, Coll. GLSH. the long axis. Distinct postdorsal angle. Part of dorsal rim sagitta, Gramian, boring Flensburg-Bommerlunder, between predorsal and postdorsal angle long and nearly 134-140m, Coll. GLSH. 3 sagittas, Pinneberger Schichten, straight. Rostral rim perpendicular to the long axis. Inner Gramian, boring Tornesch II, 19-25 m, Coll. GLSH. surface somewhat convex along longaxis. Outer surface

very slightly convex or flat along long axis. Distribution. - Upper Miocene (Gramian)-Recent.

Description. - Strong, thick and high otoliths of medium

Discussion. - Otoliths of equal thickness and height, size. Outline elongated. Ventral rim thick, blunt, regularly described Koken by (1891) as Gadus anglicus can be found bent and distinctly sculptured. Caudal end pointed. Dorsal among the recent specimens of T. luscus. There is no rim much thinnerthan ventralrim but still rather blunt. Part reason therefore to distinguish between two species. of dorsal rim between predorsal and postdorsal angle very

and nearly straight. Part of dorsal rim between Trisopterus incognitus n. sp. long

(Figs, la, b, c) postdorsal angleand caudalend slightly concave. Predorsal

angle very sharp, postdorsal angle not as fully developed.

2. Rostral rim and Merlangius cognatus (Koken); Heinrich, 1969, p. 25, pi. XI, fig. straight perpendicular to the long axis. 511

Inner surface somewhat convex along long axis. Sulcus Diagnosis. - A slender, rather thin Trisopterus species.

acusticus long, wide and shallow. Cauda twice as long as Sulcus acusticus wide. Cauda as wide as ostium. Ostium rims ostium. Collumsomewhatnarrowedand very short. Sulcus somewhat narrower near collum. Dorsal and ventral

almost completely filled with colliculi. Cauda with weak regularly and weakly bent. Rostral rim nearly straight with

s-shaped bend. Cristae very insignificant. Crista superior oblique angle on long axis. Inner surface convex along long

axis, somewhat more pronounced than crista inferior. Area axis. Outer surface slightly concave along long clearly

shallow and long. Ventral furrow clear and rather far from concave in juvenile specimens. rim ventral rim. Small furrows and knobs from ventral up to

ventral furrow. L: 10.79 mm, H: 4.64 mm, T: 2.84 mm (Oploo)

L: 10.57 mm, H: 4.57 mm, T: 2.95 mm (Gram) Outer surface very slightly convex orflatalong long axis.

Ventral part of otolithclearly thickest. Big knobs and clear

Material. - 2 sagittas, Upper Miocene, Gramian, Gram, furrows perpendicularto therims. Middlepart ofotoliths in Denmark, Coll. RGM 175848, 175849. 2 sagittas, Gramian, particular distinctly sculptured. Medially some central boring at Pinneberg-Elz, 84-92 m, Coll. GLSH. 3 sagittas, knobs. Upper Miocene, Langenfeldian, 'Flachbohrung 37',

Coll. RGM RGD, Haarlem. 1 2.05 Oploo, 175828, 175829, leg. L: 7.47 mm, H: 3.55 mm, T: mm (holotype) sagitta, Gramian, Pinneberger Schichten, boring Tornesch

Caudal end is broken off, thus true length is somewhat II, 19-25 m, Coll. GLSH.

greater.

Distribution. - Upper Miocene (Langenfeldian) - Recent.

Material. - Several sagittas (mostly fragments), clay pit,

Hohen Woos, Upper Miocene, Langenfeldian, Coll. Discussion. - The specimens of the Gramian>cannot be from their the Heinrich (PMHU). distinguished recent ones by shape. Because otoliths of this species have been differentiated from the

branch of luscus for a geological period Distribution. - Upper Miocene, Langenfeldian. Trisopterus long

and since they show even less resemblance to T. minutus,

T. Discussion. - The general form of the otoliths of T. capelanus has to be consideredas a separate species and of stated Svetovidov incognitus indeed resembles that of Merlangiogadus not as a subspecies T. minutus, as by (in Hureau & Monod, 1973). cognatus in many respects. Therefore it is not surprising

that Heinrich assigned this species to M. cognatus. The otoliths of T. capelanus of the Langenfeldianfrom show closer the otoliths from the Especially the long straight part between the predorsal and Oploo a similarity to T. concavus—T. luscus than the postdorsal angles causes confusion. Nevertheless the lineage geologically

entire shape of the sulcus, the straight rostral rim and the younger specimens. The splitting of that lineage very occurred before the Miocene. greatthickness are characteristic for the genus Trisopterus. probably shortly Upper

makes it clear that The Gaemers & The presence of a postdorsal angle species Trisopterus elongatus Schwarzhans introducedbefore I had the otoliths T. incognitus is an ancestor of T. minutus, which has a was seen of T. It is between the somewhat more compact and thicker appearance. capelanus. not possible to distinguish

two morphologically.

Trisopterus minutus (Linnaeus, 1758) 1973 (PI. IV, Figs. 2a, b, 4a, b) Genus MERLANGIOGADUS Gaemers,

Merlangiogadus cognatus (Koken, 1891) Diagnosis. - A thick and high Trisopterus species of (PI. V, Figs, la, b, 2a, b) medium size, which is distinctly bent along the long axis.

of dorsal rim between Distinct postdorsal angle. Part Otolithus (Merlangus) cognatus Koken, 1891, p. 89, pi. V, fig. 1 short and to predorsal and postdorsal angle straight slightly non pi. Ill, fig. 5.

Rostral rim to the axis. convex. nearly perpendicular long Merlangus cognatus Koken; Weiler, 1942, p. 79, pi. IX, figs. 7, 8. axis. Outer Merlangus Koken; Weiler, Inner surface markedly convex along long cognatus 1959, p. 97. Merlangus cognatus Koken; Brzobohaty, 1964, 275. surface slightly concave or s-shaped along long axis. p. Merlangus 461. cognatus Koken; Zilch, 1965, p. Gadus elegans sculptus Koken; Smigielska, Distribution. - Upper Pliocene-Recent. 1966, p. 241, pi. XV, fig. 3.

Merlangus? Koken; Trisopterus capelanus (Lacepède, 1800) cognatus Smigielska, 1966, p. 243, pi. XV, fig 4. (pi IV, Figs, la, b, 3a, b)

Merlangius cognatus (Koken); Weiier, 1968, p. 32. Merlangius Gadus planatus Bassoli & Schubert, 1906; Chaine & Duvergier, cognatus (Koken); Gaemers, 1969a, p. 8, pi. I, fig. 8.

14. Merlangius cognatus (Koken); Gaemers, 74. 1928, p. 195, pi. 6, figs. 13, 1969b, p. Merlangius Gadus cf.friedbergi Chaine & Duvergier, 1928; Heinrich, 1969, p. cognatus (Koken); Gaemers, 1971, p. 244, pi. I, fig. 9 80, pi. IV, figs. 3a, b. pi. II, fig. 3, pi. V, fig. 5, pi. VI, fig. 1.

Trisopterus elongatus Gaemers & Schwarzhans, 1973, 210, pi. I, Merlangiogadus cognatus (Koken); Gaemers, 1973, 63. p. p. figs. 5, 6a, b, pi. IV, figs. 3a, b.

Type. - Trisopterus minutus capelanus (Lacepède, 1800); Svetovidov, Lectotype: PI. V, Figs, la, b, Coll. PMHU, Ot. 46,

310. 1973, p. Coll. Koken. 512

Locus typicus. - Langenfelde, Western Germany. collum somewhat narrowed and clearly deeper. Dorsal rim

variable: nearly straight without indentations to clearly

Stratum typicum. - Upper Miocene, Langenfeldian. indented half-way between the caudal point and the

predorsal angle.

Paralectotype. - PI. V, Figs. 2a, b, Coll. PMHU, Ot. 46,

L: 9.79 H: 3.81 T: 1.76 Coll. Koken, same locality and age as lectotype. mm, mm, mm (lectotype)

L: 9.18mm, H: 3.59mm, T: 1.21 mm (pi. V, Figs. 3a, b)

Diagnosis. - An elongate, but relatively compact

Distribution. - Upper Oligocene or Lower Miocene. Merlangiogadus species. Adult specimens relatively shorter and than compacter juvenile ones.

Discussion. - The drawings of Koken (1891) are for misleading two reasons. Firstly the otoliths of Material. - 1 sagitta, Upper Miocene, Langenfeldian, Merlangus spatulatus and ofMerluccius attenuatus are too 'Flachbohrung 37', Oploo, Coll. RGM 175830, leg. RGD. slender. Secondly the strange indentationhas been greatly

exaggerated. In reality the otoliths of the two forms Distribution. - Lower Miocene-Upper Miocene. distinguished by Koken resemble one another much more

than is suggested by Koken's Also the sulcus of Remarks. - drawings. Very probably the lectotype is the same both A forms is very similar. specimen of G. spatulatus specimen as shown by Koken (1891, pi. V, fig. 1). Many with details a distinctly straight dorsal rim has the pointed rostral agree with this drawing. The only difficulty is that end of this thereforeis Merluccius attenuatus; specimen an the lectotype is a mirror image of the drawing. intermediate form. The specimen drawn by Schwarzhans

(1974, fig. 74) is another intermediate form with a weak Genus GADICHTHYS Gaemers & Schwarzhans, 1973 indentationin the dorsal rim. Because of these intermediate

forms I decided uniteboth to of Koken's species into one Gadichthys undosus Gaemers, 1973

single one.

Gadichthys undosus Gaemers, 1973, p. 60, pi. I, figs. 1, 2, 4, PI. II,

fig. 7. Gadichthys antwerpiensis (Gaemers, 1971)

Gadiculus undosus 139. (Gaemers); Gaemers, 1974b, p.

Gadus friedbergi Chaine & Duvergier; Weiler, 1942, pi. V, figs.

Distribution. - Upper Oligocene. 14a, b.

Merlangusspatulatus (Koken); Weiier, 1942,p. 79, pi. IX, figs. 6, 9. Remarks. - Up to now this species is the oldest known Merlangius spatulatus (Koken); Gaemers, 1969a, 9, pi. I, fig. 9. representative of the Gadichthys lineage. p. Trisopterus antwerpiensis Gaemers, 1971, p. 242, pi. I, fig. 8, pi. V, fig. 3. Gadichthys spatulatus (Koken, 1891) Gadichthys antwerpiensis (Gaemers); Gaemers & Schwarzhans, (pl.V, Figs. 3a, b-5a, b) 1973, p. 2B.

Gadiculus antwerpiensis (Gaemers); Gaemers, 1974b, p. 139. Otolithus (Merlangus)spatulatus Koken, 1891, p. 89, pi. II, figs. 6,

7, 10. Distribution. - Lower-Middle Miocene. Otolithus (Merluccius) attenuatus Koken, 1891, p. 85, pi. II, figs.

1,2. Gadichthys miocenicus (Heinrich, 1969) Merlangius spatulatus (Koken); Heinrich, 1969, p. 61, fig. 11.

Colliolus spatulatus (Koken); Schwarzhans, 1974,p. 101, fig. 74, miocenicus Merlangius spatulatus Heinrich, 1969, p. 25, pi. V, non figs. 4-7, 13, 14. figs. I, 2, pi. XVIII, 1. Colliolus fig. 2, pi. XIX, fig. attenuatus (Koken); Schwarzhans, 1974, p. 102, fig. 75 Gadichthys miocenicus (Heinrich); Gaemers, 1973, p. 62.

Gadiculus miocenicus 139. (Heinrich); Gaemers, 1974b, p. Type. - Lectotype: PI. V, Figs. 5a, b, Coll. PMHU, Ot. 43,

Coll. Koken. Material. - 1 sagitta, Upper Miocene, Langenfeldian,

'Flachbohrung 37', Oploo, Coll. RGM 175831, leg. RGD. Locus typicus. - Germany.

Distribution. - Upper Miocene, Langenfeldian.

Stratum typicum. - Upper Oligocene or Lower Miocene

(Sternberger Gestein). Gadichthys benedeni (Leriche, 1926)

- 1 Paralectotypes. sagitta, Merlangus spatulatus, Gadus benedeni Leriche, 1926, p. 457, pi. XLI, fig. 10.

Koken 2 benedeni 244. Sternberger Gestein, Ot. 42, Coll. (PMHU). Trisopterus (Leriche); Gaemers, 1971, p.

benedeni Gaemers & sagittas, Merluccius attenuatus, Sternberger Gestein, Ot. Gadichthys (Leriche); Schwarzhans, 1973,

213, pi. I, 13-17, pi. VII, 1, non pi. I, 8. 41, leg. Von Nettelblad (PMHU). p. figs. fig. figs. 7, benedeni Gadichthys (Leriche); Gaemers, 1973, p. 62.

Gadiculus benedeni (Leriche); Gaemers, 1974b, p. 139. Diagnosis. - A slenderand thin Gadichthys species with a

very sharp caudal point and a rounded to rather pointed

rostral end. Sulcus wide and shallow but near the long Distribution. - Lower Pliocene. 513

form of both Gadichthys verticalis (Gaemers & Schwarzhans, 1973) relatively long cauda), is the intermediate

Gadiculus species.

benedeni verticalis Gaemers & is the of Gadichthys Schwarzhans, 1973, p. Possibly ?G. venustus juvenile form G. 214, pi. I, fig. 12, pi. VI, fig. 1. both forms miocenicus. This is more probable, because are Gadichthys benedeni verticalis Gaemers & Schwarzhans; known from the same geological stage (Langenfeldian).

Gaemers, 1973, p. 62. Moreover small specimens of G. miocenicus and large - Gadiculus benedeni verticalis (Gaemers & Schwarzhans); unknown. specimens of ?G. venustus are The only Gaemers, 1974b, p. 139. intermediate size and difficulty is that specimens with an

length-height ratio have not been found up to now. Distribution. - Upper Pliocene. in the Should my hypothesis prove to be correct future,

G. miocenicus has to be consideredas a junior synonym of

Discussion. - In the first instance this species was

G. venustus. considered as a subspecies of G. benedeni. This was due

to mainly to the fact that specimens belonging a

Boreogadus species were assumed to be juvenile Genus GADICULUS Guichenot, 1850

specimens of G. benedeni. This fault will be corrected in

Gaemers & Schwarzhans (in prep.). All juvenile and adult Gadiculus ellipticus (Schubert, 1905)

G. specimens ofG. benedeni can now be distinguished from (PI. V, Figs. 6a, b; PI. VII, Figs. 3a, b, 4a, b)

to verticalis and therefore there is no longer any reason

consider the Upper Pliocene otoliths of Gadichthys as a Otolithus (Macrurus) ellipticus Schubert, 1905, p. 622, pi. XVI, subspecies of G. benedeni. figs. 31-33.

excisus Otolithus (Macrurus) Schubert, 1905, p. 623. thori Gadichthys (Schmidt, 1914) Otolithus (Macrurus) ellipticus Schubert; Schubert, 1906, p. 665,

pi. V, figs. 8-12.

Gadiculus Thori J. Schmidt, 1914, p. 1, fig. 1. Glyptorhynchus ellipticus (Schubert); Dieni, 1968, p. 269, pi. Ill,

Gadiculus thori 306. 5. argenteus Schmidt; Svetovidov, 1973, p. figs. 4,

Gadiculus thori Schmidt; Gaemers, 1974b, p. 139. Macrurus communis (Prochazka); Anfossi and Mosna, 1972, p.

105, pi. IV, fig. 6.

Distribution. - Recent.

Diagnosis. - A compact, small Gadiculus species with dorsal Discussion. - The outline, especially the rimand the indistinctpredorsal and postdorsalangles. Postdorsal angle caudal point, of G. thori is highly variable. Also the developed most fully in juvenile specimens. Caudal end variable. probably G. thori length-heightratio is quite Very Dorsal rim clearly but not very sharply pointed. fairly is a late ofthe Gadichthys lineage. Unfortunately, for stage regularly rounded, especially in adult specimens. Highest want of otoliths of this species it is still impossible to point of otolith more towards the middle. Ventral rim reconstruct the diagrams with length-height and normally bent in rostral part. Variability is greatest in the length-thickness ratios. When these diagrams become dorsal rim. identification be verified. In available, the genus can any

case Gadiculus and thori are argenteus Gadichthys L: 5.00 mm, H: 3.17 mm, T: 1.04 mm

for otoliths of both can separate species, the species always L: 4.85 mm, H: 3.27 mm, T: 0.99 mm

be distinguished. L: 5.18 mm, H: 3.52 mm, T: 1.06 mm

L: 2.51 mm, H: 1.84 mm, T: 0.56 mm

? Gadichthys venustus (Koken, 1891)

(PI. Figs. b) V, 7a, Material. - 4 sagittas (3 juveniles), Dingdener Schichten,

Glimmerton, Middle Miocene, bed of creek near pi. figs. 2, 3. Otolithus (Gadus) venustus Koken, 1891, p. 91, V, Königsmühle, Dingden, Western Germany, Coll. RGM Gadus venustus Koken; Weiler, 1942, 77, pi. V, figs. 59, 60. p. 175873, 175874, leg. M. van den Bosch. 1 sagitta, same Gadus 4. venustus Koken; Heinrich, 1969, p. 24, pi. Ill, figs. 2, locality and same bed, Coll. Gaemers, leg. Cadée. I sagitta, Gadiculus 139. venustus (Koken); Gaemers, 1974b, p. same locality and same bed, Coll. RGM 175872, leg. M.

Freudenthal. 1 sagitta, same locality, DingdenerSchichten, Distribution. -Upper Miocene, Langenfeldian. Feinsand, Middle Miocene, Coll. RGM 175875, leg. A. W.

Janssen.

Discussion. - Gaemers (1974b) considered this species to

belong to Gadiculus Guichenot as the ancestor of

Distribution. - Middle Miocene-Lower Pliocene. Gadiculus argenteus. Indeed ?G. venustus has some

In characteristics in common with Gadiculus. the first place

Remarks. - Possibly the Miocene Gadus communis of the otoliths are not bent along the long axis. Secondly the Prochazka is the height profile closely resembles that of Gadiculus. (1894) same species as Gadiculus the The Middle Miocene Gadiculus ellipticus Schubert ellipticus (Schubert). Very probably specimen shown -_ by Prochazka was eroded. This drawing resembles the recent G. argenteus so closely that it is highly shows a more

shown unlikely that the Upper Miocene ?G. venustus, which elongated form than the specimens by Schubert,

which resemble the material of differs in some respects (a very pointed caudal end and a Dingden very closely. 514

Gadiculus Guichenot, 1850 rim is argenteus caudal rounded; the sulcus is relatively narrow.

These characteristics of Protocolliolusagree with those of

Macrurus 24. sp. Gaemers&Schwarzhans, 1973, p. 218,pi. I,fig. Colliolus and Neocolliolus. Therefore Protocolliolus will

be the ancestor of the Colliolus lineage.

Diagnosis. - A compact Gadiculus species with rather

distinct predorsal and postdorsal Predorsal angle angles. Genus COLLIOLUS Gaemers & Schwarzhans, 1973

most conspicuous. Caudal end clearly but not very sharply

pointed. Dorsal rim before predorsal angle, between Species typicus. - Gadus friedbergi Chaine & Duvergier,

predorsal and postdorsal angles and behind postdorsal 1928. in and adult angle straight (except very large specimens).

Highest of otolith toward the front. point conspicuously - Diagnosis. A genus with oval to elongated otoliths. Ventral rim markedly bent in rostral Variability in part. Rostral end rounded and caudalpoint rather sharpto sharp. outline is small and the outline as a whole is regular. very Predorsal and postdorsal angles weakly to moderately

pronounced. Postdorsal angle sometimes absent. Parts

L: 6.27 mm, H: 4.23 mm, T: 1.54 mm between caudal point and postdorsal angle and between L: 5.35 mm, H: 3.93 mm, T: 1.16 mm predorsal and postdorsal angles straight to slightly

concave. Sulcus acusticus straight and rather narrow, Distribution. - Upper Pliocene-Recent. collum rather long and slightly narrowed. Cauda about

twice as long as ostium. Sulcus rather deep, colliculi not Remarks. — Macrurus from Ouwerkerk of sp. boring very thick. Ventral part of otoliths is thickest. Upper Pliocene age fits the diagnosis ofG. argenteus quite

well. Although the inner surface is totally destroyed, the Distribution. - Middle Oligocene-Lower Pliocene. outline and the profile of the otolith reveal, that this Colliolus parvus n. sp. Middle Oligocene. specimen belongs to this species. Colliolussculptus (Koken, 1891) Upper01igocene-?Lower

Miocene. Genus PROTOCOLLIOLUS n. gen. Colliolus Miocene. johannettae n. sp. Lower Colliolus friedbergi (Ch. & Duv., 1928) Middle Species typicus. - Gadus eocenicus Frost, 1931. Miocene-Lower Pliocene.

Colliolus schwarzhansi n. sp. Middle Miocene.

- A Diagnosis. genus with thick and compact otoliths.

Outlineoval with roundedrostral end and distinctly pointed Colliolusparvus n. sp.

caudal end. Predorsal and postdorsal angles moderately (PI. VI, Figs, la, b-8a, b)

pronounced. Parts between caudal point and postdorsal

angleand betweenpredorsal and postdorsal angles straight. Type. - Holotype: PI. VI, Figs, la, b, Coll. RGM 175879. Sulcus acusticus straight and rather wide, collum short and

clearly narrowed. Cauda somewhat less than twice as long Locus typicus. - Clay pit 'De Vlijt' near Winterswijk, the as ostium. Sulcus shallow and to a great extent filled with Netherlands.

colliculi. Thickest part of otoliths is at the median line or

slightly shifted dorsally. Stratum typicum. - Middle Oligocene, Rupelian, top of

Brinkheurne Member.

Distribution. - Lower Eocene.

Protocolliolus eocenicus Lower Eocene Derivatio - = (Frost, 1931) nominis. parvus (lat.) small. Namedafter the

(Lower Ypresian). small size of the otoliths.

- Discussion. Up to now only one species, P. eocenicus, Diagnosis. - A small, compact and barely sculptured be in can placed the genus Protocolliolus. Certainly more Colliolus species. Ventral rim slightly sculptured. Dorsal finds have be in the future. to expected rim smooth. Predorsal and postdorsal angles well Stinton (1965) considers the species P. eocenicus to be a developed. Caudal end pointed, rostral end angular to member of the genus Microgadus Gill, 1863, ofwhich only pointed. Outer surface with a few thick knobs and deep the Girard is recent Microgadus proximus known. Based furrows, especially in the middle part. Not bent along long on otoliths however this latter species is most closely axis. related to Merlangius, and to a lesser degree to

Micromesistius. Most important differences between Description. - Otoliths thick, small and compact. Outline

Microgadus and Protocolliolus are the following. The elongated. Only ventral rim slightly sculptured, otherwise otoliths of Microgadus are relatively thin with the highest smooth. Ventral rimthick, bluntandregularly bent. Caudal in the the rim has part nearly middle; caudal developed end distinctly pointed. Dorsal rim rather thick and sharper more or less as a straight line (when the sculpturing is not than ventral rim. Predorsal and postdorsal angles well taken into which forms consideration), a sharp angle with developed. Parts of dorsal rim between ends and angles the ventral rim and obtuse an angle with the dorsalrim; the straight or nearly straight. Postdorsal angle not far from sulcus is relatively wide. Protocolliolus has thick otoliths; caudal end. Rostral end angular to pointed. the highest part is definitely towards the caudal end; the Inner surface slightly convex. Sulcus acusticus long, 515

rather wide and nearly straight. Cauda straight or very angle; this angle dividesdorsal rim between predorsal angle

slightly bent withconvex side to dorsalrim. Cauda less than and caudal point into two equal to nearly equal parts, which

twice the lengthof ostium. Collum deepand narrowest part are straight to slightly concave. Distinctly bent along long

of sulcus. Distinct colliculi fill sulcus only partly. Cristae axis.

and Ventral insignificant. Area shallow large. furrow very

indistinct. Weak small furrows and knobs with regular Description. - Otoliths strong and large. Outline highly

pattern reach ventral furrow. elongated. Thick and blunt ventral rim is regularly bent,

Outer surface convex, highly sculptured in the middle, sharpest curve on rostral side. Caudalend with sharp point.

nearly smooth at the ends. Relatively big knobs and clear Dorsal rim thinnerand sharper than ventral rim. Predorsal

furrows form an irregular pattern. angle clearly visible. Postdorsal angle usually rather well

developed; this angle is located (nearly) midway between

L: 3.00 mm, H: 1.51 mm, T: 0.64 mm (holotype) two straight to slightly concave parts of the dorsal rim

L: 3.07 mm, H: 1.62 mm, T: 0.70 mm between predorsal angleand caudalend. Dorsal rim before

L: 2.72 mm, H: 1.38 mm, T: 0.67 mm predorsal angle straight to slightly convex.

L: 2.88 mm, H: 1.42 mm, T: 0.72 mm Inner surface markedly convex, especially in adult L: 3.08 mm, H: 1.62 mm, T: 0.71 mm specimens. Sulcus acusticus long with average width. L: 2.47 mm, H: 1.27 mm, T: 0.58 mm Caudal part of cauda slightly bent with convex side toward L: 2.13 mm, H: 1.24 mm, T: 0.53 mm rim. ventral Sulcus as a whole nearly straight. Cauda nearly

the twice as long as ostium. Sulcus rather deep, in spite of Material. - All sagittas: clay pit 'De Vlijt', Winterswijk, distinctly developed colliculi. Crista superior sharp and Netherlands,Middle Oligocene, Rupelian. 460 sagittas, top clear,especially ostium. Crista inferior of Brinkheurne Member, Coll. RGM 175879, 175880, along insignificant.

Area wideand large. Weak furrows traverse entire area 175881, leg. M. v. d. Bosch, 1973 (new pit). 9 sagittas, up crista Ventral furrow distinct. Small furrows Brinkheurne Member, Coll. RGM 85513, leg. De Groot (old to superior. to ventral in pit). 21 sagittas, Brinkheurne Member, Coll. RGM 127064, deep, extending furrow; juvenile specimens intersect ventral furrow. leg. M. Freudenthal. 13 sagittas, basal deposits of they normally Outer surface and Winterswijk Member, Coll. RGM 175822, leg. M. v. d. concave, totally distinctly sculptured. low and wide of knobs with the axis Bosch. 29 sagittas, basal deposits of Winterswijk Member, Medially a ridge long to the of the otolith. knobs Coll. RGM 175883, 175884. parallel height Peripheral to the rims. The otoliths from the Brinkheurne Member are foundin perpendicular

the Serpula septaria - Ancystrocyrinx volgeri assemblage

(van den Bosch et al., 1975). zone L: 9.62 mm, H: 3.79 mm, T: 1.74 mm (holotype)

L: 7.04 mm, H: 2.87 mm, T: 1.23 mm (PI. VI, Figs. 9a, b)

Distribution. - Middle Oligocene, Rupelian.

Material. - All sagittas: testpit for E-3 tunnel, Antwerp, Discussion. - The shape ofthe dorsal rim, the caudal and Miocene, Sands of Edegem. 133 sagittas, Coll. RGM rostral ends and the sculpture on the outer surface are

175855 (holotype), 175856, 175860, leg. v. d. Mark. 147 variable. This species can be consideredthe ancestor ofall sagittas, Coll. RGM 175858.42 sagittas, Coll. RGM 175859, other Colliolus species. leg. F. J. Janssen. 93 sagittas, Coll. RGM 175857, leg.

Haandrikman. 350 sagittas, Coll. RGM 155176, 155177. 81 Colliolus johannettae n. sp. sagittas, Coll. Gaemers, leg. Cadée and Coll. Cadée. (PI. VI, Figs. 9a, b, 10a, b)

Trisopterus friedbergi (Chaine & Duvergier, 1928); Gaemers, Distribution. - Lower Miocene. 3. 1971, p. 242, pi. I, figs. 5, 6; pi. V, fig. 2; pi. VI, fig.

Non Trisopterusfriedbergi (Chaine& Duvergier); Gaemers, 1971,

Sands of Antwerp. Discussion. - Variation is small in most respects. Highest

variability notedin dorsal rim. Most specimens have rather

Type. - Holotype: PI. VI, Figs. 10a, b, Coll. RGM 175855, distinct predorsal and postdorsal angles, but in some there

leg. D. v. d. Mark. is a smoothly curved dorsal rim without angles. The

length-height and length-thickness ratios show little

- bank of Locus typicus. Test pit for E-3 tunnel, right variability.

Scheldt River, Antwerp, Belgium.

Colliolus schwarzhansi n. sp.

Stratum typicum. - Miocene, Sands of Edegem. (PI. VI, Figs. 11a, b-13a, b)

nominis. - wife Johannetta Derivatio Namedafter my Anje

Type. - Holotype: PI. VI, 12a, b, Coll. Swen. Figs. RGM 175862, leg. A. W. Janssen, 1970.

- thick Colliolus Diagnosis. A very slender but normally

with caudal Predorsal Locus - Bed of the species a very sharp point. angle typicus. Königsbach Creek near

distinct. Generally a rather well developed postdorsal Königsmühle, Dingden, Western Germany. 516

Stratum - Middle Miocene, Reinbekian: typicum. Reinbekian, 165 sagittas, Coll. RGM 175862 (holotype), Dingdener Schichten, Feinsand. 175861, 175863, 175864, leg. A. W. Janssen; 35 sagittas

Coll. Gaemers, leg. Cadée.

Bank of creek near Königsmühle, Dingden, Western Derivatio nominis. - Named after Werner Schwarzhans, Germany, DingdenerSchichten, Glimmerton, Reinbekian: otolith specialist from Berlin, Western Germany 123 sagittas, Coll. RGM 175865, 175866, leg. v.d. Bosch; 23

sagittas, Coll. Gaemers, leg. Cadée. Diagnosis. - A slender and thin Colliolus species. Outer Caudal point of specimens is broken off. surface and outline distinctly and regularly sculptured. many

Dorsal rim from A predorsal angle up to caudalend straight.

Distribution. - Middle Miocene, Reinbekian. weakly developedpostdorsal angle can be present and then

the dorsal rim between the predorsal angle and the caudal

Discussion. - Variation is restricted mainly to differences end is divided into two straight or slightly concave parts. in the ofthe shape dorsal rim. Postdorsal angle is present or Caudal end sharply pointed. Rostral end angular. Weakly Dorsal rim before is not. predorsal angle convex to straight. bent along long axis. In contrast to other large Colliolus species the rostral

is end angular; as a result the rostral end ofC. schwarzhansi

- Otoliths thin and Outline Description. large. highly appears more robust than in other species of this genus.

and Rather thick and elongated beautifully sculptured. C. schwarzhansi and C. friedbergi are found togetherin

blunt ventral rim bent. Caudal end with regularly sharp the 'Dingdener Schichten' of Dingden. They can easily be Dorsal rim thinner and than ventral rim. point. sharper distinguished from one anotherbecause C. schwarzhansi is Predorsal angle rather well to well developed. A weakly much longer, thinnerand less bent along the long axis than divides the dorsal rim pronounced postdorsal angle C. friedbergi. Moreoverthe juvenile specimens of the two between and caudal end into predorsal angle two, nearly species differ conspicuously. C. johanettae can be

equal, straight to slightly concave parts. This part of dorsal distinguished from C. schwarzhansi because the former is rim also be then can absolutely straight; no postdorsal angle thicker and bends more along the long axis.

exists. Dorsal rim before predorsal angle straight to

convex. Rostral end is angular. Colliolus friedbergi (Chaine & Duvergier, 1928) Inner surface slightly convex. Sulcus acusticus long and

Gadus Chaine & nearly straight. Caudal part of cauda somewhat bent with (?)friedbergi Duvergier, 1928, p. 197, pi. VI, figs.

16-27. convex side toward ventral rim. Cauda somewhat more Gadus schuberti Smigielska, 1966, 240, pi. XV, figs. 2. than twice the length of ostium. Sulcus rather shallow. p. 1,

Ostium deeper than cauda. Colliculi clearly developed.

Remarks. - In contrast to the of Smigielska Cristae opinion insignificant. Crista superior is traversed by many it be stated length-height distinct (1966) can that the and furrows which cross the wide, large and shallow length-thickness ratios for Gadus schuberti are exactly the area. Thus the crista superior is knobbed. Ventral furrow same as those for C. friedbergi. The somewhat different distinct and rather far from ventral rim, which is clearly predorsal and postdorsal angles as well as the lesser degree incised by small furrows. These furrows are usually of sculpturing of the outer surface fall within the variation bifurcated in the middle of part the ventral rim. They cross of the C. friedbergi. ventral furrow many times. The specimens described as Gadus schuberti are Outer surface young slightly concave, totally and distinctly specimens of the same species as those adults which are sculptured. Mediallyan irregular and inconspicuous row of

correctly identified as C. friedbergi. This can also be circular to elongated knobs. Peripheral knobs deduced from the smaller size and the lesser curvature perpendicular to the rims. The highest part of the otolith along the long axis. distinctly shifted towards ventral rim; this characteristic is typical for Colliolus species in contrast to Gadiculus Genus NEOCOLLIOLUS species, in which the n. gen. highest part of the otolith is more in the middle.

Species typicus. - Gadus Esmarkii Nilsson, 1855.

L: 11.85 mm, H: 4.02 mm, T: 1.48 mm (holotype)

- A slender Diagnosis. genus with otoliths. Outline L: 10.62 mm, H: 3.72 mm, T: 1.47 mm more or less elongated with rounded rostral end and L: 9.43 H: 3.32 mm, mm, T: 1.18 mm L: sharply pointed caudal end. Predorsal and postdorsal 8.45 mm, H: 3.08 mm, T: 1.12 mm

L: 7.62 H: 2.87 angles weakly to moderately pronounced. Parts between mm, mm, T: 0.99 mm

L: 6.36 and and between mm, H: 2.34 mm, T: 0.78 mm caudal point postdorsal angle predorsal

L: 5.45 H: 2.04 mm, mm, T: 0.69 mm and postdorsal angles straight. Sulcus acusticus straight

L: 4.14 mm, H: 1.67 mm, T: 0.49 mm and rather wide, collum short and somewhat narrowed.

Cauda twice ostium. Sulcus and as long as very shallow

with colliculi. almost completely filled Ventral part of

otolith is thickest. Material. - Bed of creek near Königsmühle, Dingden,

Western - Germany, Dingdener Schichten, Feinsand, Distribution. Upper Miocene-Recent. 517

Neocollioluspliocenicus (Gaemers & Schwarzhans, 1973) ?Palaeoraniceps upnoriensis (Stinton, 1965) Lower

Upper Miocene-Lower Pliocene. Eocene.

Neocolliolus esmarki (Nilsson, 1855) Recent. Palaeoraniceps tuberculosus (Koken, 1884) Lower

Oligocene-Upper Oligocene.

n. Lower Discussion. - Biologists believe that the recent species Palaeoraniceps regularis sp. Miocene.

Neocolliolusesmarki belongs to the genus Trisopterus (see -The be Svetovidov, 1973) or the genus Boreogadus (see Muus, Discussion. genus Palaeoraniceps can easily form of the 1966). distinguished from Raniceps by the different

The fishes and the otoliths of the recent Boreogadus sulcus and the colliculi. Fedotov describes several skeletons of the saida differ so greatly from those of N. esmarki that the (1974) Upper

be distant. established in his relationship must very Thereforeit is impossible Oligocene species Pseudoraniceps sagus,

to place N. esmarki within the genus Boreogadus. The publication as a new species belonging to a new genus.

affinity with Trisopterus is much greater, but the shape of Unfortunately otoliths of this species are unknown. Two

P. P. the fishes as well as their otoliths still show many other Oligocene species, par cus (Daniltshenko) and

differences. We will restrict ourselves here to the otoliths oligocaenicus (Bogatshov), are also known by their

of wide sulcus skeletons from the Caucasus. It is not known whether alone. The sagittas Trisopterus have a very one of acusticus, are relatively thick and usually do not have of them is identical with Palaeoraniceps tuberculosus

dorsalangles Trisopterus minutus is the only exception; it Western Europe. It will be necessary to find either the

P. has a weakly developed postdorsal angle). These skeleton of, tuberculosus or the otoliths of the Russian

the otoliths in properties contrast with those ofN. esmarki. Other recent Pseudoraniceps species. Perhaps are present differ situ in the Russian It would be genera even more. species. exceedingly found in these skeletons. With N. esmarki and N. pliocenicus show the closest interesting if they were to be

the and of resemblance to the fossil genus Colliolus Gaemers & photograph description Raniceps upnoriensis

Schwarzhans. The main differences with this genus are the Stinton, 1965, it is not possible to determine whether this

Also other more compact form and the somewhat widerand shallower species belongs to the genus Palaeoraniceps.

sulcus Thus consider Colliolus described Stinton of Neocolliolus. we may species by as Raniceps species, namely

ancestral of R. R. Lower as the genus Neocolliolus, which is ex- papillosus and latidens from the Eocene

London cannot be with in the pressed in the names of these genera. Clay, placed certainty new

genus. The drawings which accompany the descriptions

Neocolliolus pliocenicus (Gaemers & Schwarzhans, 1973) are very rough and thus insufficient. In any case they all

cannot be members of the Raniceps lineage and have to

Gadus elegans Koken, 1884; Smigielska, 1973, p. 6, pi. I, figs, la, belong to another genus.

b.

- tuberculosus 1884) Remarks. The specimen shown by Smigielska closely Palaeoraniceps (Koken,

resembles the holotype of Neocolliolus pliocenicus (PI. IV, Figs. 5a, b)

(Gaemers & Schwarzhans, 1973; PI. IV, figs. 4a, b). The

shapeand depth ofthe sulcus are The outlineof both equal. Otolithus (Gadi) tuberculosus Koken, 1884, p. 540, pi. XI, fig. 1.

in 12. specimens is very similar. The absence of distinct angles Otolithus (Gadidarum)planusKoken, 1884, p. 545, pi. XI, fig.

the Tortonian otoliths of Smigielska is the only difference Otolithus (Gadidarum)latisulcatus Koken, 1884, p. 545, pi. XI, fig.

from the Pliocene holotype. 5. 88. Otolithus (Raniceps) tuberculosus (Koken); Koken, 1891, p.

Otolithus latisulcatus (Koken); Koken 1891, 86, SubfamiliaRANICEPSINAE (Raniceps) p. pi. IV, figs. 4, 4a, PI. Ill, figs. 2, 2a. Genus PALAEORANICEPS n. gen. Otolithus (Raniceps) planus (Koken); Koken, 1891,pi. Ill, figs. 7,

7a; pi. IV, figs. 5, 5a.

Species typhus. - Otolithus (Gadi) tuberculosus Koken, Macrurus latisulcus (Koken); Leriche, 1910, p. 352, fig. 152. 1884. Raniceps latisulcatus (Koken); Weiier, 1942, p. 83.

Raniceps planus (Koken); Weiier, 1942, p. 83, pi. V,.figs. 4a, b, 12,

Diagnosis. - A genus with thick and compact otoliths. non fig. 10.

Outline egg-shaped with rounded rostral end and bluntly Raniceps tuberculosus mut. supraoligocaena Weiier, 1942, p. 81, pi. 10, figs. 3-6. pointed caudal end. No angles or only very weakly tuberculosus 10, 1, Raniceps (Koken); Weiier, 1942, p. 81, pi. figs. developed dorsal angles. Sulcus acusticus very wide and 2. slightly bent with convex side toward ventral rim. Collum Raniceps latisulcatus (Koken); Weiier, 1957, p. 137 very short and only slightly narrowed. Cauda 1.6 to 2 times Raniceps latisulcatus (Koken); Weiier, 1958,p. 332, pi. 1, figs. 17, as long as ostium. Sulcus very shallow and to a large extent 18.

filled with colliculi which have an angular form at the Raniceps planus (Koken); Weiier, 1958, p. 332, pi. 1, fig. 19. collum. Thickest of otoliths is ventral. part slightly Raniceps latisulcatus (Koken); Martini, 1964, p. 53, fig. 1 (4).

Raniceps planus (Koken); Martini, 1964, p. 53, fig. 1 (5).

lalisulcatus Gaemers, Distribution. - Lower Oligocene-Lower Miocene. Lower Raniceps (Koken); 1972, p. 76, pi. 2, figs.

Eocene? 3a, b. 518

Coll. Type. - Lectotype: PI. IV, Figs. 5a, b, PMHU, Ot. slightly bent. Entire sulcus only slightly bent toward

67, Coll. Koken. ventral side.

Locus typicus. - Siildorf, Germany. Description. - Otoliths strong and medium-sized. Outline

egg-shaped. Numerous knobs and furrows all around.

Stratum typicum. - Middle Oligocene. Rather thick ventral rim is regularly bent. Caudalend with

blunt point, situated more towards dorsal rim. Dorsal rim

Paralectotypes. - 2 sagittas, Raniceps tuberculosus, thinner than ventral rim, regularly bent. Predorsal angle

Middle Oligocene, Siildorf, ot. 67, Coll. Koken; 3 sagittas, weakly developed.

Lower R. tuberculosus, Oligocene, Lattorf, ot. 68, Coll. Inner surface nearly flat, only very slightly convex.

7 R. tuberculosus, Koken; sagittas, Magdeburger Sand, Sulcus acusticus long, wide and shallow, somewhat convex

Middle Oligocene, Neustadt near Magdeburg, ot. 69, Coll. towards ventral side. Rims of cauda straight. Rims of Middle Koken; 15 sagittas, R. planus, Oligocene, ostium slightly curved. Ostium and cauda angular near

Koken. All Coll PMHU. ostium. collum Söllingen, ot.70,Coll. specimens: collum. Cauda 1.5 times as long as High

distinctly divides sulcus into ostium and cauda. Colliculi

thick and darkly coloured. Cristae low but sharp; crista

- P. tuberculosus Discussion. The unique shape of as superior more pronounced than crista inferior. Furrows in

shown by Koken (1884) is a very distorted and therefore dorsal part deeper and usually longer than in ventral part.

unserviceable drawing. I have studied Koken's type Nearly entire inner surface sculptured.

materialof Raniceps tuberculosus, R. latisulcatus and R. Outer surface convex, markedly and totally sculptured.

in the of cluster of circular knobs. planus, which is stored Palaontologisches Museum Medially a big to oval Peripheral

the Humboldt Universitat. It wasevident from this material knobs perpendicular to the rims.

that these three species distinguished by Koken belong to only one species. The specimens ofR. tuberculosus are all L: 7.10 mm, H: 3.73 mm, T: 1.66 mm (holotype)

T: adult forms. The best preserved specimen has been chosen L: 5.27 mm, H: 2.83 mm, 1.19 mm (eroded)

L: 3.41 mm, H: 1.78 mm, T: 0.90 mm as lectotype and is shown in this publication in the hope that

there will be in the future for no longer any problems

Material.- Totally 4 sagittas (one broken and one eroded), scientists. The specimens of.R. planus are all smallotoliths Sands of Edegem, bed 3, Miocene, right bank of Scheldt except one, which is medium-sized. This otolith cannot be River, test pit E-3 tunnel, Antwerp: 1 sagitta, Coll. RGM distinguished from the specimens of.R. tuberculosus. The 175869; 3 sagittas, Coll. RGM 175867 (holotype), 175868, small specimens are rather variable. They have a flat inner leg. Haandrikman. surface and a clearly convex outer surface. The widthofthe

sulcus as well as the length-height ratio of the otoliths is

Distribution. - Miocene; Sands of Edegem. highly variable. There is no doubt that they are juvenile

specimens of P. tuberculosus. I also have seen the type

Discussion. - Variability is small. Rims ofostium are more material of Raniceps latisulcatus. of or less curved. Most probably this species is variety èta From the drawingand the description (Koken, 1884) it is Raniceps latisulcatus (Koken, 1884) which is mentioned already clear that these specimens are considerably eroded from the black (glauconitic) Miocene Sands of Antwerp. It and therefore not suitable for the description of a new this describe is a pity thatKoken does not show variety, nor species. These specimens also belongto P. tuberculosus. it.

Palaeoraniceps regularis n. sp. 2 - ofthis Remarks. Together with specimens new species, (PI. VII, Figs, la, b) broken and eroded otoliths of Raniceps tuberculosus

(Koken, 1884) were found in the Sands of Edegem at Type. - Holotype: PI. VII, Figs, la, b, Coll. RGM 175867, d. Coll. RGM Antwerp (Coll. RGM 175871, leg. v. Bosch; leg. Haandrikman. 175870, leg. F. J. Janssen). The conservation of these

otoliths is very bad and also differs from that of the Locus typicus. - Test pit for E-3 tunnel, right bank of autochthonous ones. They are darker, more bluish in Scheldt River, Antwerp, Belgium. colour. They originate from the Middle Oligocene Clay of

Boom and have survived because of their strength. Stratum typicum. - Lower Miocene,Sands of Edegem, bed

3 (see Janssen and van der Mark, 1968).

Genus RANICEPS Oken, 1817

= Derivatio nominis. - regularis (lat.) regular. Namedafter

- with thick and the very regular outline. Diagnosis. A genus compact otoliths.

Outline egg-shaped with rounded rostral end and bluntly

- A No Diagnosis. highly sculptured and very regularly formed pointed caudal end. angles. Sulcus acusticus very wide with side Palaeoraniceps species. Weak predorsal angle. Blunt point and clearly bent convex toward ventral rim. at caudal end shifted to dorsal rim. Ostium and cauda Collum average in length and very narrow. Cauda 1 to 1.3 angular near collum. Rims ofcauda straight, rims ofostium times as long as ostium. Sulcus very shallow and to a large 519

extent filled with colliculi which have a rounded oval form. slightly and regularly bent and somewhat sculptured near

Thickest part of otoliths is slightly ventral. rostral and caudal ends; otherwise smooth. Dorsal rim

but irregularly sculptured with rather big not very

Raniceps raninus (Linnaeus, 1758) pronounced knobs. Mediodorsalangle clearly developed. rims (PI. VII, Figs. 2a, b) Rostral end more sharply pointed than ventralend. All

sharp and thin.

Gaemers & Inner Raniceps sp. Schwarzhans, 1973, p. 217, pi. II, fig. 5; surface slightly convex. Sulcus acusticus long,

4. PI. VII, fig. wideand shallow. Ratio ofthe length of ostiumto the length

of cauda is ca. 3/4. Collum narrow. Sulcus disappears at the

- A moderately and irregularly sculptured Diagnosis. rims. Colliculi with oval shape. Cristae distinguishable; with oval outline. Generally no angles. Raniceps species crista superior and inferior equally important. Clearly Ostium caudarounded collum. Rims ofostium and and near but visible area high not very long. Small furrows short and cauda bent. Entire sulcus distinctly bent toward ventral shallow. Ventral furrow very well developed. side. Outer surface slightly concave. Most conspicuous

furrows and knobs along dorsal rim and caudal end.

Material. - 1 broken sagitta, Sands of Kallo, Upper Sculpture pattern irregularly radiate. Thickest part of Pliocene, basal shell bed, pit for sea sluice near Kallo, otolith in the middle, just below mediodorsal angle. Belgium, Coll. Gaemers, leg. Vervoenen.

H: L: 8.72 mm, 3.59 mm, T: 0.96 mm

Distribution. - Upper Pliocene-Recent.

Material. - 2 sagittas (1 considerably eroded), bed of

Remarks. - is certain that raninus has lived It not Raniceps Königsbach creek near Königsmühle, Dingden, Western

in the neighbourhood of Antwerp during the formationof Germany, Middle Miocene, Dingdener Schichten, the Sands of Kallo. The specimen found is somewhat Feinsand, Coll. RGM 175877 (holotype), 175878, leg.

eroded; moreover there are many fossils in the basal bed of A. W. Janssen, 1970.

this location the Sands of Kallo which were transported to

older and which originate from Pliocene deposits. Of Distribution. - Middle Miocene: Hemmoorianand Rein-

the course minimum age is the age of the Sands of Kallo. bekian.

otolith that this The state of preservation of the proves

fossil from the Sands of Kallo the Sands of - comes or Discussion. This species can be considered as an ancestor

Luchtbal. of the recent Molva molva (Linnaeus) which is much larger

with greater curvature. Molva primaeva seems to be

Subfamily LOTINAE intermediatebetween the Palaeogadus lineage and Molva Genus MOLVA Lesueur, 1819 molva (L.). Therefore it seems very likely that the Molva

lineage has proceededfrom thePalaeogadus lineage which Molva primaeva n. sp. has become extinct. Palaeogadus species have a much 7a, (PI. VII, Figs. b) more asymmetrically triangular shape than Molva species;

moreover curvature is less.

Nemopteryx emarginatus (Koken, 1891);Weiler, 1942,pi. 6, figs,

la, b. - definite of Remarks. Weiier (1942) shows a example

Molvaprimaeva from the HemmoorianofHemmoor but he

Type. - Holotype: PI. VII, Figs. 7a, b, Coll. RGM 175877, has identified this specimen as Nemopteryx ( = leg. A. W. Janssen, 1970. Palaeogadus) emarginatus. The close resemblance of

these species is demonstrated by this error.

Locus typicus. - Bed of Königsbach creek near

Königsmühle, Dingden, Western Germany. Genus BROSME (Cuvier) Oken, 1817

Stratum typicum. - Middle Miocene, Reinbekian,

Brosme heinrichi n. Dingdener Schichten, Feinsand. sp. (Figs. 2a, b, c)

= Derivatio nominis. - primaevus, -a (Latin) in his (her) Otolithus (inc. sed.) 4 Heinrich, 1969, 46, pi. XI, fig. 3. first youth. Very probably this species represents the first sp. p.

appearance of the genus Molva.

Type. - Holotype: Figs. 2a, b, c, Coll. Heinrich (PMHU).

Heinrich, 1969: pi. XI, fig. 3. Diagnosis. - A small Molva species, which is only slightly bent along the long axis. Therefore innersurface is slightly

Locus - Hohen convex, outer surface somewhat concave. No torsion along typicus. Clay pit, Woos, Deutsche long axis. Demokratische Republik.

- Description. - Elongated, rather thin otoliths. Outline Stratum typicum. Upper Miocene, Langenfeldian (see nearly an isosceles triangle with a low height. Ventral rim Heinrich, 1969). 520

L: 8.42 mm, H: 3.88 mm, T: 1.06 mm

Material. - 1 sagitta, clay pit, Hohen Woos, Eastern

Germany, Upper Miocene, Langenfeldian, Coll. Heinrich

(PMHU).

Distribution. - Upper Miocene, Langenfeldian.

Discussion. - The recent species Brosme brosme

(Ascanius, 1772) has more compact, less pointed otoliths,

which have some torsion along the long axis. The part ofthe dorsal rim before the mediodorsalangle is convex, the part

behind the mediodorsal angle concave. The ventral rim is

not as regularly bent. On the basis of these differencesB.

heinrichi and B. brosme can easily be distinguished from

one another.

Familia ?MACRURIDAE

?Macruridarum deurnensis n. sp.

(PI. VII, Figs. 5a, b, 6a, b)

Type. - Holotype: PI. VII, Figs. 6a, b, Coll. RGM 175847,

leg. F. J. Janssen.

Locus typicus. - Foorplein, Antwerp/Borgerhout, Bel-

gium.

Ventral side of left Fig. 2. a. Inner surface, b. Outer surface, c.

Stratum - Miocene, Sands of Deurne. 10x. typicum. Upper sagitta from Brosme heinrichi n. sp. Magnification

Derivatie) nominis. - Named after the sands in which the

species is found.

Derivatio nominis. - Named after Dr. Wolf-Dieter

Heinrich, otolith specialist from Berlin, Eastern Germany, Diagnosis. - A species with small and compact otoliths. who first found and described this species. Inner surface flat. Cauda and ostium equal in length.

long. Entire sulcus acusticus clearly depressed. Diagnosis. - A rather small and slender Brosme species Collum Below collum distinct furrow. Distinct predorsal and with a clearly triangular outline which is practically Ventral furrow far from ventral rim. isosceles. Rostral and caudal points both sharp. Middorsal postdorsal angles. angle distinct. Parts between rostral point and mediodorsal

- otoliths. Outline angle and between caudal point and middorsalangle nearly Description. Small, compact and strong Ventral and dorsal rims rather thin and straight. Otolith slightly bent along long axis and without egg-shaped. sharp. Ventral rim bent. Dorsal rim consists of straight torsion along this axis. regularly and pieces between caudal and rostral ends, and predorsal

- otoliths. Outline Description. Rather thin and elongated Caudal end postdorsal angles. Both dorsal angles distinct. isosceles Ventral rim practically like an triangle. regularly bluntly pointed. Entire outline sculptured with low and and symmetrically bent and nearly smooth. Dorsal rim regular knobs and clear small furrows in between. distinct middorsal somewhat irregularly sculptured with a Inner surface flat. Entire sulcus acusticus distinctly angle. Parts of dorsal rim before and behind mediodorsal depressed and wide. Ostium and cauda equal in length. angle nearly Rostral end a little bit more sharply straight. Colliculi small and nearly circular. Collum long and pointed than caudal end. All rims sharp and thin. somewhat narrower than ostium and cauda. Below collum

Inner surface slightly convex. Sulcus acusticus very long and distinct furrow, slightly bent with convex side wide and rather long and deep. Ostium a little shorter than toward ventral rim. Area unimportant. Ventralfurrow very at cauda. Collum narrow and long. Sulcus disappears very obvious and far from ventral rim. the rims. Caudal colliculum fartherfrom the rim than ostial Outer surface somewhat convex and well-sculptured Cristae colliculum; both are roundedand oval in shape. not with a radiate pattern of distinct knobs and furrows. very pronounced; crista superior sharper than crista Furrows are shallowest near caudal and rostral ends. inferior. Area large but not clearly bordered along the

dorsal rim. Ventral furrow far from ventral rim. T: 0.58 L: 2.51 mm, H: 1.98 mm, mm (holotype)

Outer surface concave. Most slightly conspicuous L: 2.36 mm, H: 1.70 mm, T: 0.48 mm furrows and knobs along dorsalrim between mediodorsal

and caudal - of angle point. Sculpture pattern vague. Material. 2 sagittas, Sands Deurne, Antwerp- 521

Borgerhout, Foorplein, Coll RGM 175846, 175847, Naturkunde an der Humboldt-Universitat zu Berlin,

leg. F. J. Janssen. Deutsche Demokratische Republik (PMHU).

I am grateful to the following individuals who made this

- Miocene, Distribution. Upper Sands of Deurne. publication possible by placing at my disposal fossil otoliths

of codfishes: Mr. M. van den Bosch and Mr. A. W.

Discussion. - This species as well as the other small species Janssen (both RGM), Mr. M. C. Cadée (Leiden), Dr. W.

Hinsch described as Macruridae from the Middle Miocene and (GLSH), Mr. A. F. J. Janssen (Purmerend) and older deposits shows a striking resemblance to the otoliths Mr. M. Vervoenen (Aalst, Belgium).

of Gadiculus species. The sulcus acusticusof both groups Moreover I am grateful to Dr. W.-D. Heinrich and

is nearly the same. Small specimens of Gadiculus have a Dr. H. Jaeger(both PMHU), who lent me Dr. E. Koken's

shallower sulcus than the small Macruridae. Ostium and type material; with this material it was possible to eliminate

cauda of Gadiculus can differ considerably in length, some errors which had been introduced into the literature

whereas these are usually equally long in the Macruridae. mainly as a result of mistakes in the drawings in Koken's

in send Properly speaking the most striking difference the publications. The same persons were so kind as to me

otoliths is their size. The small Macruridae generally do not otoliths of Gadidae from Heinrich's collection of Hohen

exceed three millimeters in length, whereas otoliths of Woos. I wish to thank Mr. V. W. M. van Hinsbergh Gadiculus of millimeters. and Schwarzhans species often reach a length 10 (Nijmegen) Mr. W. (West-Berlin) for the

possibility of publishing some Gadidae from Winterswijk

and Dingden. I am thankful to Dr. N. Daan (RIVO,

ACKNOWLEDGEMENTS IJmuiden), Dr. J. E. Fitch (Long Beach, California), Dr.

D. Nolf (Gent, Belgium), Mr. W. Schwarzhans

I thankful to Prof. Dr. A. Brouwer for his critical Mr. C. von am very (West-Berlin), Vaupel-Klein (Leiden) and remarks. Mr. G. van der Velde (Nijmegen) for their friendly help in

In addition to some material from my own collection the obtaining recent otoliths of Gadidae.

I wish otoliths described in this publication were kindly placed at to thank Mr. W. C. Laurijssen and Mr. W. A. M. Devilé for my disposal by the Rijksmuseum van Geologie en preparing the good photographs, and Miss

Mineralogie at Leiden (RGM), the Rijks Geologische C. N. de Vries and Mrs. F. G. M. Madjlessi-Leenen for

Dienst at Haarlem (RGD), the Geologisches Landesamt typing a substantial part of the manuscript.

Schleswig-Holstein at Kiel, Bundesrepublik Deutschland Finally I am grateful to Mrs. G. P. Bieger-Smith for

(GLSH) and the Palaontologisches Museum, Museum fiir correcting the English text.

REFERENCES

G. & 1972. Otoliti del Pliocene Inferiore di Anfossi, Mosna, S., -, 1969b. Otolieten uit het Anversien van Antwerpen II. Meded.

1st. Lugagnano(Piacenza). Atti Geol. Univ. Pavia, 23, pp. 90-118. Werkgr. Ten. Kwart. Geol., 6/4, pp. 69-79.

fossili terziari dell' Emilia. Rev. Ital. Bassoli, G., 1906. Otoliti -, 1971. Bonefish-otoliths from the Anversian (Middle Miocene)

36-58. of Leidse Geol. Meded., 46, 237-267. Paleontol., 12, pp. Antwerp. pp.

Bosch, M. Cadée. M. C. & Janssen, A. W., 1975. 1972. Otoliths from the of the Sands of van den, -, type locality Berg

I .il host rati graphical and biostratigraphicalsubdivision of Tertiary (Middle Oligocene) at Berg, Belgium. Meded. Werkgr. Tert.

- Kwart. deposits (Oligocene Pliocene) in the Winterswijk-Almelo region Geol., 9/3-4, pp. 73-85.

(eastern part of the Netherlands). Scripta Geol., 29, pp. 1-167. -, 1973. New otoliths from the Tertiary of the North Sea Basin.

otolitü Meded. Brzobohaty, R., 1964. Poznamka k vyzkumu rybich z Werkgr. Tert. Kwart. Geol., 10/2, pp. 58-75. tertiéru 1974a. De betekenis taxonomische waarde otolieten. zapadnich Karpat. Zpravy o geolog. vyzkumech v. gr. -, en van

Het 1963, pp. 275-276. Zeepaard, 34/3, pp. 29-40.

J. & J., 1928. Sur les otolithes fossiles de la 1974b. Otolieten uit het Merksemien Icenien Chaine, Duvergier, -, en van boring

Pologne. Rocznik polskiego towarzystwa geologicz, 5, pp. Ouwerkerk (Zeeland, Nederland). Meded. Werkgr. Tert. Kwart.

190-204. 133-143. Geol., 11/3, pp.

Dieni, I., 1968. Gli otoliti del Pliocene Inferiore di Orosei Gaemers, P. A. M. & Schwarzhans, W., 1973. Fisch-Otolithen

(Sardegna). Mem. Accad. Patavina SS. LL. AA. Classe Sci., Mat. aus dem Pliozan von Antwerpen (Belgien) und Ouwerkerk

Nat., 80, pp. 243-295. (Niederlande) und aus dem Plio-Pleistozan der Westerschelde

Gadidae from the Fedotov, V. F., 1974. New Upper Oligocene (Niederlande). Leidse Geol. Meded., 49, pp. 207-257. Translated from 78-83. - & Fisch Otolithen aus der Caucasus. Paleontol. J., 1974/1, pp. -. Typuslokalitat der obermiozanen

Treskovyye iz verknego oligotsena Kavkaza. Paleontol. Zhur., Sylt-Stufe (Morsum Kliff, Insel Sylt, Nordwestdeutschland).

1974/1, pp.83-89. Leidse Geol. Meded. (in prep.).

Frost, G. A., 1931. Appendix 1. Fish otoliths from Eocene strata Heinrich, W.-D., 1969. FischotolithenausdemObermiozan von

faunas of below the London Clay. In White, E. I.: The vertebrate Hohen Woos. Geologie, 67, pp. 1-111.

A. W. & D. 1968. den the English Eocene, 1, pp. 105-109. Janssen, Mark, van der, Einleitung zu Anversien Gaemers, P. A. M., 1969a. Otolieten uit het van Beitragen zur Kenntnis der Molluskenfauna des jüngeren Tertiars

3-21. 76-82. Antwerpen. Meded. Werkgr. Tert. Kwart.Geol.,6/1-2,pp. im Nordseebecken. Basteria, 32/4-5, pp. 522

Koken, E., 1884. Über Fisch-Otolithen, insbesondere über 1906. Die Fischotolithen des -, oesterreich-ungarischen

der norddeutschen Z. Tertiars. III. Jahrb. diejenigen Oligozan-Ablagerungen. geol. Reichsanstalt, 56, pp. 623-706.

deutsch. geolog. Ges., 36, pp. 500-565. Schwarzhans, W., 1974. Die Otolithen-Fauna des Chatt Aund B Über 1885. Otolithen. In: A. von: eine von -, Koenen, palaeozane (Oberoligozan, Tertiar) Niederrhein, unter Einbeziehung

Abh. Ges. Wissensch. weiterer Fauna von Kopenhagen. kgl. Göttingen, 32, Fundstellen. Decheniana, 126/1-2, pp. 91-132.

113-116. pp. Smigielska, T., 1966. Otoliths of fishes from the Tortonian of Neue 1891. Untersuchungen an tertiaren Fisch-Otolithen II. South Poland. Ann. Soc. -, géol. Poiogne, 36, pp. 205-275.

Z. deutsch. geolog. Ges., 43, 77-170. 1973. Fish otoliths from the Lower Tortonian pp. -, deposits at

M., 1906. Note préliminairesur faluns Niskowa Sacz. Ann. Soc. 3-45. Leriche, lespoissonsdes near nowy géol. Poiogne, 43, pp.

néogènes de la Bretagne, de l'Anjou et de la Touraine. Ann. Soc. Stinton, F. C, 1965. Teleost otoliths from the Lower London

géol. du Nord, 35, pp. 290-321. Tertiaries. Senck. leth., 46a, pp. 389—425.

1908. Note sur les et éocènes des -, poissons paléocènes Svetovidov, A. N., 1973. Merlucciidae, Gadidae. In: Hureau, J.

Ann. environs de Reims (Mame). Soc. géol. du Nord, 37, pp. C. & Monod, Th. (ed.): Checklist ofthe fishes ofthe north-easlern

Atlantic 229-265. and of the Mediterranean (Clofnam), Part I. Unesco,

1910. Les de la Mém. Musée 300-320. -, poissons oligocènes Belgique. Paris, pp.

Roy. Hist, natur. Belg., 5, pp. 231-264. Voigt, E., 1926. Überein bemerkenswertes Vorkommen neuer

Martini, E., 1964. Ein in Stettiner Gestein Fischotolithen in Otolithenpflaster einem Senongeschiebevon Cöthen in Anhalt. Z.

Natur und 53-59. 2, 172-187. Museum, 94, pp. Geschiebeforsch., pp.

Posthumus, O., 1923. Bijdrage tot de kennis der tertiaire Weiier, W., 1942. Die Otolithen des rheinischen und

vischfauna van Nederland; oligoceene en mioceene otolieten uit nordwestdeutschen Tertiars. Abh. Reichsamts Bodenforsch., N. het Peelgebied en van Winterswijk. Verhandel, v. h. Geol.-Mijn- F., 206, pp. 1-140. bouwk. 1957. Zur Fischfauna des bei Genootschap voor Nederland en Koloniën, Geol. Ser., 7, -, Dobergs Bünde in Westfalen.

105-142. Palaeontol. 135-138. pp. Z., 31, pp.

VI. 1894. Miozan Kralice 1958. Fisch-Otolithen Prochazka, J., Das von nachst -, aus dem Oberoligozan und dem

Namêsti in Mahren. Sitz. k. Böhm. Gesellsch. d. Wissensch., Mittelmiozan der Niederrheinischen Bucht. Fortschr. Geologie

Rheinland und math.-naturw. Cl. (1893), pp. 1-71. Westfalen, 1, pp. 323-361.

1959. Fisch-Otolithen dem Hemmoor Schmidt, W., 1968. Vergleichend morphologische Studie über -, aus Schleswig-

Holsteins. die Otolithen mariner Knochenfische. Arch. Fischereiwiss., 19, Meyniana, 8, pp. 96-194.

1968. Fossilium I Animalia. Beiheft 1, pp. 1-96. -, Catalogus. (Ed. F. Westphal), Pars 117, Otolithi Piscium. Schubert, R. J., 1905. Die Fischotolithen des oesterreich- W. Junk, 's-Gravenhage, pp. 1-196. ungarischen Tertiars. II Macruriden und Beryciden. Jahrb. Zilch, A., 1965. Die Typen und Typoide des Natur-Museums geol. Reichsanst., 613-638. Senckenberg, 31: Fossile Fisch-Otolithen. Senck. 55, pp. leth., 46a, pp. 453^*90. 523

PLATES 524

PLATE I

Fig. 1a, b. Gadus parallelus n. sp., Pliocene, Antwerp, Ford factory, holotype (Coll. RGM

175800) 6x.

Gadus RGM Fig. 2a, b. parallelus n. sp.. Pliocene, Antwerp, Ford factory, paratype (Coll.

175801) 6x.

Fig. 3a, b. Pollachius virens (Linnaeus, 1758). Recent, North Sea (Coll. Gaemers, leg. RIVO)

4x.

Fig. 4a, b. Pollachius carbonarius (Linnaeus, 1758), Recent, North Sea (Coll. Gaemers, leg.

RIVO) 4x.

Fig. 5a, b. Pollachius carbonarius (Linnaeus, 1758), Recent, North Sea (Coll. Gaemers, leg.

RIVO) 4x.

Fig. 6a, b. Pollachius carbonarius (Linnaeus, 1758), Recent, North Sea (Coll. Gaemers, leg.

RIVO) 4x. 525/I 526

PLATE II

Fig. 1a, b. Trisopterus concavus n. sp., Middle Miocene, Hemmoorian, Miste, holotype (Coll. RGM 175822) 7.5x.

Fig. 2a, b. Pollachius virens (Linnaeus, 1758), Recent, North Sea (Coll. Gaemers, leg. RIVO)

4x.

b. Fig. 3a, Trisopterus concavusn. sp., Upper Miocene, Langenfeldian,Oploo, paratype (Coll.

RGM 175823, leg. RGD) 7.5x.

Fig. 4a, b. Pollachius virens (Linnaeus, 1758), Recent, North Sea (Coll. Gaemers, leg. RIVO)

4x.

Fig. 5. Trisopterus spectabilis (Koken, 1891), Upper Miocene, Langenfeldian, Oploo (Coll.

RGM 175825, leg. RGD) 7.5x.

Fig. 6a, b. Trisopterus spectabilis (Koken, 1891), Upper Miocene, Langenfeldian,Oploo (Coll.

RGM 175826, leg. RGD) 7.5x. 527/II 528

PLATE III

Fig. 1a, b. Trisopterus spectabilis (Koken, 1891), Upper Miocene,Gramian, Sands of Deurne,

juvenile specimen (Coll. RGM 175837) 10x.

Fig. 2a, b. Trisopterus spectabilis (Koken, 1891), Upper Miocene, Gramian, Sands of Deurne

(Coll. RGM 175838) 7.5x.

Fig. 3a, b, c. Trisopterus spectabilis (Koken, 1891), Upper Miocene, Langenfeldian,

Langenfelde, lectotype (Coll. PMHU, Ot. 55) 7.5x.

Fig. 4a, b, c. Trisopterus spectabilis (Koken, 1891), Upper Miocene, Langenfeldian,

Langenfelde, paralectotype (Coll. PMHU, Ot. 55) 7.5x.

Fig. 5a, b. Trisopterus luscus (Linnaeus, 1758), Upper Miocene, Gramian, Gram (Coll. RGM

175851) 7.5x. 529/III 530

PLATE IV

Fig. 1a, b. Trisopterus capelanus (Lacepède, 1800). Upper Miocene. Langenfeldian, Oploo

(Coll. RGM 175828) 7.5x.

Fig. 2a, b. Trisopterus minutus (Linnaeus, 1758), Pliocene, Antwerp, Ford factory (Coll. RGM

175808) 10x.

Fig. 3a, b. Trisopterus capelanus (Lacepède, 1800), Upper Miocene, Gramian, Gram (Coll.

RGM 175849) 7.5x.

Fig. 4a, b. Trisopterusminutus (Linnaeus, 1758), Upper Pliocene, Sands of Kallo, sea sluice near

Kallo (Coll. Gaemers, leg. Vervoenen) 10x.

Fig. 5a, b. Palaeoraniceps tuberculosus (Koken, 1884), Middle Oligocene, Süldorf, lectotype

(Coll. PMHU, Ot. 67) 6x. 531/IV 532

PLATE V

Fig. 1a, b. Merlangiogadus cognatus (Koken, 1891), Upper Miocene, Langenfeldian

Langenfelde, lectotype (Coll. PMHU, Ot. 46) 7.5x.

Fig. 2a, b. Merlangiogadus cognatus (Koken, 1891), Upper Miocene, Langenfeldian

Langenfelde, paralectotype (Coll. PMHU, Ot. 46) 7.5x.

Fig. 3a, b. Gadichthys spatulatus (Koken, 1891), Upper Oligocene or Lower Miocene,

Sternberger Gestein, paralectotype (Coll. PMHU, Ot. 41) 7.5x.

Fig. 4a, b. Gadichthys spatulatus (Koken, 1891), Sternberger Gestein, paralectotype (Coll.

PMHU, Ot. 42) 7.5x.

Fig. 5a, b. Gadichthys spatulatus (Koken, 1891), SternbergerGestein, lectotype (Coll. PMHU,

Ot. 43) 7.5x.

Fig. 6a, b. Gadiculus ellipticus (Schubert, 1905), Middle Miocene, Reinbekian, ‘Glimmerton’,

Dingden (Coll. Gaemers, leg. Cadée) 10x.

Fig. 7a, b. ?Gadichthys venustus (Koken, 1891), Upper Miocene, Langenfeldian,boring Städt.

Krankenhaus, Kiel, 96—99 m (Coll. GLSH) 10x. 533/V 534

PLATE VI

Fig. 1a, b. Colliolus parvus n. sp., Middle Oligocene, clay pit ‘De Vlijt’, Winterswijk, holotype

(Coll. RGM 175879) 10x.

b. Colliolus Middle Fig. 2a, parvus n. sp., Oligocene, clay pit ‘De Vlijt’, Winterswijk, paratype

(Coll. V. van Hinsbergh) 10x.

Fig. b. Colliolus n. 3a, parvus sp., Middle Oligocene, clay pit ‘De Vlijt’,Winterswijk, paratype (Coll. Gaemers, leg. Cadée) 10x.

Fig. 4a, b. Colliolus parvus n. sp., Middle Oligocene, clay pit ‘De Vlijt’, Winterswijk, paratype (Coll. Gaemers, leg. Cadée) 10x.

Fig. 5a, b. Colliolus parvus n. sp., Middle Oligocene, clay pit ‘De Vlijt’, Winterswijk, paratype

(Coll. V. van Hinsbergh) 10x.

b. Colliolus Middle Fig. 6a, parvus n. sp., Oligocene, clay pit ‘De Vlijt’, Winterswijk, paratype

(Coll. RGM 175880) 10x.

7a, b. Colliolus Middle Fig. parvus n. sp., Oligocene, clay pit ‘De Vlijt’, Winterswijk, paratype

(Coll. V. van Hinsbergh) 10x.

8a, b. Colliolus Middle Fig. parvus n. sp., Oligocene, clay pit ‘De Vlijt’, Winterswijk, paratype

(Coll. V. van Hinsbergh) 10x.

9a, b. Colliolus Fig. johannettaen. sp., Lower Miocene, Sands of Edegem,test pit E-3 tunnel,

Antwerp, paratype (Coll. RGM 175856) 7.5x.

Fig. 10a, b. Colliolus johannettaen. sp., Lower Miocene, Sands of Edegem,test pit E-3 tunnel,

Antwerp, holotype (Coll. RGM 175855) 7.5x.

Fig. 11a, b. Colliolus schwarzhansi n. sp., Middle Miocene, Reinbekian, ‘Feinsand’, Dingden,

paratype (Coll. RGM 175861) 7.5x.

b. Colliolus schwarzhansi Middle Fig. 12a, n. sp., Miocene, Reinbekian, ‘Feinsand’, Dingden,

holotype (Coll. RGM 175862) 7.5x.

Fig. 13a, b. Colliolus schwarzhansi n. sp., Middle Miocene, Reinbekian, ‘Feinsand’, Dingden,

paratype (Coll. RGM 175863) 7.5x. 535/VI 536

PLATE VII

1a, b. Lower Sands of E-3 Fig. Palaeoraniceps regularis n. sp., miocene, Edegem, test pit

tunnel, Antwerp, holotype (Coll. RGM 175867) 10x.

Fig. 2a, b. Raniceps raninus (Linnaeus, 1758), Upper Pliocene, Sands of Kallo, basal shell bed,

sea sluice near Kallo (Coll. Gaemers, leg. Vervoenen) 7.5x.

Fig. 3a, b. Gadiculus ellipticus (Schubert, 1905), Middle Miocene, Reinbekian, ‘Glimmerton’,

Dingden (Coll. RGM 175872) 10x.

Fig. 4a, b. Gadiculus ellipticus (Schubert, 1905), Middle Miocene, Reinbekian, ‘Glimmerton’,

Dingden (Coll. RGM 175873) 10x.

Fig. 5a, b. ?Macruridarum deurnensis n. sp., Upper Miocene, Gramian, Sands of Deurne,

paratype (Coll. RGM 175846) 20x.

Fig. 6a, b. ?Macruridarum deurnensis n. sp.. Upper Miocene, Gramian, Sands of Deurne,

holotype (Coll. RGM 175847) 20x.

Fig. 7a, b. Molva primaeva n. sp., Middle Miocene, Reinbekian, ‘Feinsand’,Dingden, holotype (Coll. RGM 175877)7.5x. 537/VII