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J. A. Moy-Thomas and T. S. Westoll On the Permian Coelacanth, Coelacanthus granulatus, Ag.

By J. A. MOY-THOMAS, M.A., University Museum, Oxford, and T. S. WESTOLL, B . S C , University College, London.
I. INTRODUCTION.

A LTHOUGH the Permian species of Coelacanth, Coelacanthus - ^ granulatus, has been known since 1829, and although many authors have described specimens of it, it has remained unfortunately one of the lesser known of all Coelacanths. Since C. granulatus is the genotype of Coelacanthus, and since all the Carboniferous and some Triassic species have been assigned to this genus, it is surprising that further description has not been forthcoming. One of us (J. A. M. T.) has for some time been working on the Carboniferous Coelacanths, and it has been impossible to determine whether these really belong to the genus Coelacanthus. Accordingly it was hoped that by re-examining all the available material a more diagnostic description might be given. In doing this the authors have only been partly successful, as some of the most important generic charactersthe dermal bones of the cheek and snoutwere not preserved in any of the material examined. However, sufficient diagnostic characters have been provided by this research to distinguish the Carboniferous forms from this Permian genus. Another reason for the re-examination of C. granulatus is that the other author (T. S. W.) has recently been revising the Permian fishes of England. As it will still be some time before this work can be published, it will be mutually convenient to write the present note. During this work the authors have had to borrow and examine a great deal of material from various museums. They have examined material in the British Museum (Natural History), including the types of C. granulatus and " C. caudalis " ; in the Royal Scottish Museum ; the Yorkshire Museum ; the Hancock Museum of Newcastle-upon-Tyne ; the Staatssammlung of Munich, including the types of " C. hassiae " and " C. macrocephalus " ; and in the Sorbonne. The authors would like to express here their gratitude to Dr. E. I. White of the British Museum (Natural History), Dr. A. C. Stephen of Edinburgh, Dr. Collinge of York, Mr. T. Russell Goddard of Newcastle-upon-Tyne, Professor F. Broili of Munich, and Professor M. Jacob of Paris for their kindness and generosity in lending material. The authors are also indebted to Professor E. S. Goodrich and to Professor D. M. S. Watson for reading this manuscript and for many helpful suggestions. One of the authors (T. S. W.) is indebted to the Department of Scientific and Industrial Research for a Senior Research Award.

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The earliest mention of a Permian Coelacanth is by Sedgwick (1829), who figured an almost complete specimen and a separate pterygoquadrate from the Marl Slate, but did not name or describe them. Agassiz (1839) figured two caudal regions from the Marl Slate as Coelacanthus granulatus without description. Miinster (1842) described a specimen from the Kupferschiefer as C. hassiae. In 1844 Agassiz published the description of his figures under the name of C. granulosus. Egerton (1850) described two more Permian fishes, one as C. granulatus and the other as C. caudalis. Geinitz (1861) described a few bones, easily identifiable from his figures as those of a Coelacanth, as Pygopterus humboldti. Huxley (1866) in his review of the Coelacanths figured and described Egerton's type specimen of C. caudalis anew. Willemoes-Suhm (1869) described and figured a specimen as C. macrocephalus, and redescribed some of Miinster's types of C. hassiae. Reis (1888) suggested that C. macrocephalus is not distinct from C. hassiae, to our knowledge of which species he added many valuable details and figures. Woodward and Sherborn (1890) suggested that C. caudalis is an immature form of C. granulosus, and in 1891 Woodward classified all the previously described Permian species into the single species C. granulatus. This conclusion of Woodward's has been confirmed by the authors beyond doubt after re-examination of all the types. Stensio (1932) defined the genus Coelacanthus, basing his definition entirely on C. granulatus.
III. COELACANTHUS GRASULATVS AGASSIZ. " Fossil Fish." Sedgwick (1829), p. 118, pi. xi, ix, fig. 3. Coelacanthus granulatus, Agassiz (1839), pi. lxii. Coelacanthus hassiae, Miinster (1842), p. 49. Coelacanthus granulosus, Agassiz (1844), p. 172. Coelacanthus granulatus, Egerton (1850), p. 235, pi. xxviii.* Coelacanthus caudalis, Egerton (1850), p. 236, pi. xxviii. Pygopterus humboldti, Geinitz (1861), pi. viii, figs. 1-3. Coelacanthus caudalis, Huxley (1866), pp. 14, 21, pi. v, fig. 5. Coelacanthus macrocephalus, Willemoes-Suhm (1869), p. 74, pi. xi, fig. 2. Coelacanthus hassiae, Willemoes-Suhm (1869), p. 76, pi. x, fig. 1 ; pi. xi, fig. 1. Coelacanthus macrocephalus, Reis (1888), p. 68. Coelacanthus hassiae, Reis (1888), p. 69, pi. iii, fig. 22 ; pi. iv, figs. 7,12,15,16,19 Coelacanthus granulosus, Woodward & Sherborn (1890), pp. 39, 40. Coelacanthus granulatus, Woodward (1891), pp. 400-1. Coelacanthus granulatus, Stensio, 1921, p. 118. Coelacanthus granulatus, Stensio, 1932, p. 40.

Type.Caudal region (British Museum (Natural History), 3338). Locality.Marl Slate of Durham and Northumberland and Kupferschiefer of Germany.
Description, (a) General Body Form.

The species varies from medium to large forms, in which the body is moderately slender and the head measures one-fifth of

TEXT-FIG. 1.Reconstruction of Coelacanthus granulatus. X J. The squamation is omitted. Only the known dermal bones are shown on the head. None of the bones of the palate or cranium is indicated.

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the total body-length. The pelvic fins lie slightly posteriorly to the anterior dorsal fin. The anal fin is almost opposite the posterior dorsal fin. The supplementary caudal fin is well marked but probably rather small.
(b) The Head.

The head was almost as broad as deep, unlike that of many Coelacanths. The anterior bones of the skull roof are not preserved in any of the material at the disposal of the authors. Parts of the frontals are present in one of the English specimens (B.M., 3340),

s.tem.

TEXT-FIG. 2.Skull of Coelacanthus granulatus. x $. Mostly from specimen P 3340 in the British Museum (Natural History), e.sc. = extrascapulars. fr. = frontal, pa. = parietal, s.tem. = supratemporal.

and are somewhat longer than wide, but probably incomplete. The growth centre is laterally placed. The supraorbitals and dermosphenotics are badly preserved, but there is no doubt that the latter is separate from the frontals. The parietals are known from several specimens and are elongated bones ; they are separate from the supratemporals, which are somewhat triangular bones
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extending anteriorly to a point. The supratemporals bear laterally a very well-marked facet for the articulation with the antero-dorsal margin of the opercular. There is a continuous internal flange on the lateral margin of the supra temporal and parietal. The extrascapulars are poorly represented but must have numbered about six, and would seem to have been in contact with one another, and with the parietals and supratemporals. All the bones of the skull roof are sparsely ornamented with feebly marked tubercles. Only the most obscure fragments of dermal bones of the cheek are visible, but the lacrimo-jugal seems to have been well ossified. The lower jaw was strongly ossified and consisted of an angular, splenial, dentary, coronoid (possibly more than one), and a prem.pt.

au.pal.

p.vo

ang.
den

TEXT-FIG. 3.Pterygoquadrate complex of Coelacanthus gramdalus from a specimen in the Hancock Museum, Newcastle. X J. ang. = angular, au.pal. = autopalatine. cor. = coronoid. den. dentary. d.pal. = dermal palatine. ec.pt. = ectopterygoid. gu. = gular. m.pt. = metapterygoid. pt. = pterygoid. p.vo. = prevomer. qu. = quadrat?.

articular, and in addition to these Meckel's cartilage was ossified at least in the articular. The coronoid is an oblong bone externally ornamented with vertically elongated tubercles. The internal surface of this bone is also ornamented with fine vertical striae. Anterior to the coronoid there are traces of small tooth-bearing bones which are probably additional coronoid elements as in Macropoma (Watson, 1925). The other bones of the lower jaw are, as far as can be determined, similar to those of other Coelacanths and call for no further comment.

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The neurocranium was comparatively well ossified, but unfortunately this region is only known in a recognizable condition from a single specimen in the Hancock Museum in Newcastle. The ethmosphenoid moiety is apparently represented by more than one ossification. The ethmoid region is strongly ossified but badly preserved in the known material, but it seems as though there were paired ossifications. The sphenoid portion is very well ossified and presents several interesting features. The posterior face has a large hollow, on either side of the upper margin of which there is a " condyle ". Between these " condyles " and slightly anterior to the dorsal margin of the hollow is a pair of tiny pits lying very close together. Antero-dorsally to this is a well-marked transverse groove. It is rather difficult to interpret these structures. Immediately dorsal to this region is the ventral margin of the cavum cerebrate, which is

TEXT-FIG. 4.Xeurocranium of Coelacanthus granulatus, from a specimen in the Hancock Museum, Newcastle. X 1. " eondyle " = condyle on basisphenoid. pa. = parietal, psph. = parasphenoid. s.tera. = supratemporal. 4 a.Posterior view of basisphenoid of this specimen.

too mutilated for further description. Only a pair of well-marked antotic processes is visible. It seems fairly certain that the basipterygoid process was not developed. The otic region is disappointing, but considerable ossification can be seen. The parasphenoid is a fairly slender spatulate bone, covered by numerous closely set teeth. It is very closely connected with the basisphenoid. The prevomer is a small bone bearing large conical teeth. The autopalatine is a triangular bone applied to the outer surface of the pterygoid, which has a narrow anterior shank, but becomes much broader posteriorly, and passes into the very strong vertical shank which bears a pair of strong ridges on its outer surface. These ridges are prolonged into short blunt points

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dorsally. Only traces of the metapterygoid have been seen. The quadrate is a large bone closely applied to but perfectly distinct from the pterygoid, and bears a large transverse condyle for the lower jaw. The dermopalatine is a small narrow bone bearing both granular, and a few larger conical, teeth. The ectopterygoid is a small slender bone with large conical teeth.

TEXT-FIG. 5.Urohyal of Coelacanthus granulatus from a specimen in the Hancock Museum, Newcastle. X 1.

The branchial arches are always too crushed for any description. The urohyal is large and bears a strong median ventral groove leading to the posterior cleft. It tapers anteriorly.

TEXT-FIG. 6.Left operoulum of Coelacanthus granulatus. Internal view from specimen 43472 in the British Museum (Natural History). X f.

The opercular is a triangular bone somewhat deeper than it is long, its posterior margin presenting a sigmoid curve. It is covered with characteristic ornament of very sparse feeble tubercles. The

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gular plate is about a quarter as broad as long, and has no ornament preserved in any of the material examined. There is a series of small unornamented plates surrounding the eye ; these are usually termed sclerotic plates, but have been recently shown (Moy-Thomas, 1935) to be ornamented in some forms and therefore, being superficial, cannot be strictly comparable to true sclerotic plates. The distribution of the sensory canals in the bones of the skull roof is difficult to make out, and the rarity of material makes it inadvisable to prepare the system. So far as can be seen there is a canal passing through the lateral parts of the supratemporal and parietal. In the lower jaw the course of the mandibular canal is shown in several specimens ; the canal is rather wide and opens on the posterior part of the angular by several wide pores. The canal is also visible in the splenial. (c) Axial Skeleton. The axial skeleton consists of about seventy vertebrae. The neural arches are of the type usually found in Coelacanths, and the spines are short behind the head as far as the anterior dorsal fin.

TEXT-FIG. 7.Axial skeleton of posterior abdominal region of Coelacanthus granulatus, showing pleural ribs passing into haemal spines. From Munich specimen A.S. 239. X 1.

The ventral elements of the more anterior segments are small paired ossifications somewhat triangular in shape, the basiventrals. Articulating with these are short free pleural ribs, which can be seen posteriorly to come gradually together at their distal ends and pass continuously into haemal arches. This fact is particularly interesting because Stensio (1932) has suggested that similar ribs in Laugia are dorsal ribs. The haemal arches and spines call for no special comment. (d) Paired Fins and their Girdles. The pectoral girdle is of considerable interest, being well preserved. The supracleithrum is a small triangular bone with a thickened ridge running to each angle on the internal face. The region

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usually occupied by the cleithrum in other Coelacanths is in this form occupied by two separate ossifications. The larger of these forms the whole blade of the bone down to the backwardly projecting angle and is then prolonged ventrally into a narrow anterior shaft. Behind this lies the second ossification, which is an elongated oval bone pointed at each end. This bone is entirely ventral to
s.cl.

e.cl.

TEXT-FIG. 8.Pectoral girdle of Coelacanthus granulatus, mainly from Munich specimens, cl. = cleithrum. clav. = clavicle, e.cl. extracleithrum. s.cl. = supracleithrum. The arrow indicates the point of attachment of the fin skeleton. X .

the origin of the pectoral fin and therefore cannot be homologized with actinopterygian postcleithrum. It must, therefore, be a new ossification which has so far not been described in other fishes, for which the name extracleithrum is proposed. The clavicle is a small curved pointed bone which meets the lower end of both cleithrum and extracleithrum. There are about sixteen lepidotrichia in the

TEXT-FIG. 9.Pelvic bone of Coelacanthus granulatus from Munich specimen A.S. 138. x 2.

pectoral fin, jointed distally for about two-thirds of their length. The pelvic girdle has already been quite well figured by Reis (1888, pi. 4, fig. 7). The small inaccuracies in his figure have been corrected in Text-fig. 9. The two plates meet one another and

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possibly overlap slightly in the middle line. The lepidotrichia of the pelvic fin are fifteen to sixteen in number and are jointed for about half their length.
(e) Unpaired Fins.

The internal skeleton of the anterior dorsal fin is generally an oval bone with somewhat angular rounded outlines. The fin consists of from eleven to thirteen strong lepidotrichia which are jointed distally for about two-thirds of their length except in the anterior short rays.

TEXT-FIG. 10.Anterior dorsal fin skeleton of Coelacanthus granulatus, slightly restored, from specimen P 3342 in the British Museum (Natural History). X 1.

TEXT-FIG. 11 a.Posterior dorsal fin skeleton of Coelacanthus granulatus from specimen 43426 in the British Museum (Natural History). X 2. 116.Anal fin skeleton of specimen in the Hancock Museum, Newcastle. X 1.

The basal plate of the posterior dorsal fin has a slender anteroventral process extending between the neural spines, and a broader anterior process ; posteriorly the plate is produced into an angle which must represent some of the postero-ventral process of the basal plate of Laugia. The fin is considerably lobed, and has eighteen to twenty rays, the larger of which are jointed for about half their length. The internal skeleton of the anal fin is a small narrow plate lying some distance ventrally to the posterior ribs. The fin consists of about fifteen rays, the longer jointed for about half their length.

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In the caudal fin the radials articulate with the slightly expanded distal ends of the neural and haemal spines, and distally bear stout lepidotrichia, except in the case of the most anterior two or three. The lepidotrichia number about twenty above and twenty below ; there is usually one more ray dorsally than ventrally. One or two anterior rays are unjointed, the remainder are distally segmented for a little over half their length. The supplementary caudal fin is rather small and consists of about twelve rays both dorsally and ventrally. The lepidotrichia of all the fins show no trace of ornament. (/) Squamation. The scales are fairly large and rather thin. The whole scale (of the anterior flank) is a little longer than high. The exposed

TEXT-FIG. 12.Scales of Coetacanthus granulatus from specimen P 3338 in the British Museum (Natural History). X 2.

part of the scale is distinctly higher than long and is ornamented with numerous well-marked slightly elongated tubercles. The scales carrying the lateral line are obvious and must have been more strongly ossified than the rest of the scales. (g) The Swim-bladder. This is well marked and extends back behind the pelvic fins.
IV. SUMMARY.

The following diagnosis of the genus Coelacanthus can now be given in a form comparable with that used by Stensio (1932, pp. 39-45). Medium to large Coelacanths, moderately slender. Head onefifth of total length, almost as broad as deep. Ethmosphenoid part

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of neurocranium probably ossified in three parts, probably also with granular ossifications. Basisphenoid with antotic process, probably without distinct basipterygoid process. Otico-occipital region well ossified. Supratemporal separate from parietal, dermosphenotic separate from frontal. Unornamented " sclerotic plates ". Cheek not yet known. Pterygoid with long low anterior shank, broad vertical shank. Coronoid almost rectangular, strongly ornamented on both faces. Opercular large and triangular, the posterior margin sigmoidally curved. Palatal teeth granular, few conical teeth on prevomer, dermopalatines, and ectopterygoid. Ornament of head bones sparse feeble tubercles. Ossified basiventral and short separate pleural ribs. Pectoral girdle with cleithrum and a separate extracleithrum : pectoral fin not situated low down. Pelvic girdle of characteristic shape : pelvic fins situated just behind the anterior dorsal fin. Anterior dorsal fin arising from somewhat oval basal plate. Posterior dorsal fin arising from basal plate with long anterior and antero-ventral processes and trace of postero-ventral process. Basal plate of anal fin a small narrow plate. Caudal fin well-developed with supplementary caudal fin. Lepidotrichia of all fins unjointed for some distance proximally and always without ornament. Scales somewhat longer than high, exposed part higher than long, ornamented with tubercles arranged somewhat rostrocaudally.
V. REFERENCES.

AGASSIZ, A., 1833-1844. " Recherches sur les Poissons Fossiles," Neuchatel. EGERTON, P. M. G., 1850. In W. King, " A monograph of the Permian fossils of England," , Palaeoni. Soc, London. GEINITZ, J. B., 1861. " Dyas, etc.," Leipzig. HUXLEY, T. H., 1866. " Illustration of the structure of Crossopterygian Ganoids. 2. Coelacanthini," Mem. Geol. Surv. Unit. King., Figs. and Desc. Brit. Org. Rent., dec. 12. MOY-THOMAS, J. A., 1935. " The Coelacanth Fishes of Madagascar," GEOL. MAG., LXXII, 213. MUNSTER, G., 1842. " Beschreibung einiger merkwiirdigen Fische, etc.," Beitrage zur Petrefahten-Kunde, Bayreuth, vol. 5. REIS, 0 . M., 1888. " Die Coelacanthinen mit besonderer Beriicksichtigung der im Weissen Jura Bayerns vorkommenden Arten," Palaeontographica, 35, 1. SEDGWICK, A., 1829. " On the geological relations and internal structure of the Magnesian Limestone, etc.," Trans. Geol. Soc, (2), 3, 37. STENSIO, E. A., 1921. Triassic Fishes from Spitzbergen, pt. 1, Vienna. 1932. " Triassic Fishes from East Greenland," Medd. om Gronland, Ixxxiii, Nr. 3. WATSON, D. M. S., 1921. " On the Coelacanth Fish," Ann. Mag. Nat. Hist., (9), 8, 320. WILLEMOES-SUHM, R., 1869. " Ueber Coelacanthus und einige verwandte Gattungen," Palaeontographica, 17, 73. WOODWARD, A. S., and SHERBORN, C. D., 1890. Catalogue of British Fossil Vertebrata. WOODWARD, A. S., 1891. Catalogue of the Fossil Fishes in the British Museum, pt. 2.