BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY VOL ii 1953-1956 PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM LONDON : 1953-1956 DATES OF PUBLICATION OF THE PARTS No. i. June IQ53 No. 2. November ...... 1953 No. 3. November ...... 1953 No. 4. March IQ54 No. 5. September ...... 1954 No. 6. January ...... 1955 No. 7. September ...... 1955 No. 8. February 1956 PRINTED IN GREAT BRITAIN AT THE BARTHOLOMEW PRESS DORKING BY ADLARD AND SON, LTD. CONTENTS GEOLOGY. VOLUME 2 Page No. i. The Deer of the Weybourn Crag and Forest Bed of Norfolk. A. AZZAROLI ........... i No. 2. A Coniferous Petrified Forest in Patagonia. M. G. CALDER . . 97 No. 3. The Solution of the Piltdown Problem, j. s. WEINER, K. p. OAKLEY & W. E. LE GROS CLARK ......... 139 No. 4. Some Upper Cretaceous and Eocene Fruits from Egypt. M. E. j. CHANDLER .......... 147 No. 5. The Carboniferous Flora of Peru. w. j. JONGMANS .... 189 No. 6. Further Contributions to the Solution of the Piltdown Problem. J. S. WEINER, W. E. LE GROS CLARK, K. P. OAKLEY and others . . 225 No. 7. The Schizaeaceae of the South of England in Early Tertiary Times. M. E. J. CHANDLER ......... 29! No. 8. The Brachyopid Labyrinthodonts. D. M. s. WATSON . . . 315 7 JUL 1953 THE DEER OF THE WEYBOURN CRAG AND FOREST BED OF NORFOLK A. AZZAROLI BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY Vol. 2 No. i LONDON : 1953 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY The following papers appeared in Volume I (1949-52) : Price No. i (1949). The Pterobranch Rhabdopleura in the English Eocene. H. D. Thomas & A. G. Davis 75. 6d. No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P. Oakley & M. F. Ashley Montagu 55. No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone of the Welsh Borders. E. I. White. Pteraspis leathensis White a Dittonian Zone-Fossil. E. I. White . . . 7 s. 6d. No. 4 (1950)- A New Tithonian Ammonoid Fauna from Kurdistan, Northern Iraq. L. F. Spath ....... ios. No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal- pellum. T. H. Withers ........ 55. No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae) . T. H. Withers 5 s. No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian of Podolia. W. N. Croft . .' IO s. No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle East. T. F. Grimsdale ........ ios. No. 9 (1952). Australian Arthrodires. E. I. White .... 155. No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 6s. THE DEER OF THE WEYBOURN CRAG AND FOREST BED OF NORFOLK BY AUGUSTO AZZAROLI (University of Florence) Pp. 1-96; 50 Text-figs. BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY Vol. 2 No. i LONDON: 1953 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical Series. Parts appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 2, No. i' of the Geological series. PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued June, 1953. Price Twenty-five Shillings. THE DEER OF THE WEYBOURN CRAG AND FOREST BED OF NORFOLK By A. AZZAROLI (University of Florence) SYNOPSIS The systematics of the Cervidae and the stratigraphy of the fossiliferous Pleistocene deposits of the Norfolk coast are clarified by a revision of material mainly in the A. C. Savin collection, in the British Museum (Natural History), and the John Gunn collection, in the Castle Museum, Norwich. Three successive faunas are distinguished in the Forest Bed Series. The nomen- clature and relationships of various other fossil deer are also considered. CONTENTS Page INTRODUCTION . .......... 4 GEOLOGY ............ 5 THE PHYLOGENY OF THE CERVIDAE ....... 7 DEVELOPMENT OF THE ANTLERS . . . . . . . .11 SYSTEMATIC DESCRIPTIONS ......... 15 Genus Libralces Azzaroli . . . . . . .15 Libralces gallicus Azzaroli . . . . . . .16 Libralces reynoldsi n. sp. ....... 19 Libralces minor n. sp. ........ 27 Libralces latifrons (Johnson) . . . . . .27 Libralces incertae sedis ....... 30 Genus Capreolus Hamilton Smith ...... 30 Capreolus capreolus (L.) ....... 32 Genus Cervus L. ......... 32 Cervus cf. elaphus L. . . . . . . . -35 Genus Euctenoceros Trouessart . . . . . . -37 Euctenoceros tetraceros (Dawkins) ...... 38 Euctenoceros ctenoides (Nesti) ...... 38 Euctenoceros sedgwicki (Falconer) . . . . . .40 Euctenoceros incertae sedis ....... 45 Genus Megaceros (Owen) ....... 46 The group of Megaceros giganteus ...... 48 The group of Megaceros verticornis . . . . .51 Megaceros verticornis (Dawkins) ...... 53 Megaceros dawkinsi (Newton) . . . . . .61 Megaceros savini (Dawkins) ....... 67 Megaceros, incertae sedis ....... 72 Cervidae incertae sedis . . . . . . . -73 " Cervus " obscurus n. sp. ....... 73 Cervid cf. Dama nestii nestii Forsyth Major .... 79 Cervid cf. Dama clactoniana Falc. . . . . . .81 Species incorrectly recorded from the Forest Bed . .83 CHRONOLOGY ........... 84 SUMMARY . . . . . . . . . . . .90 REFERENCES ........... 91 INTRODUCTION A REVISION of the Forest Bed deer has been needed for a long time. The descrip- tions of the most interesting specimens are scattered in the older literature : Falconer (1868), Dawkins (1887), Newton (1882), Gunn (1891), Harmer (1899). A few specimens were described more recently by Reynolds (1929-34), but these works refer mainly to the antlers, and no comprehensive study has been published. Moreover the series in the British Museum (Natural History) has been considerably enriched through the careful and long-continued work of A. C. Savin (1861-1948) of Cromer ; his first collection was acquired by the Geological Department in 1897, and his second in 1945. The material examined consists of several hundred specimens, mostly in the Savin collection. I have further studied the important collection made by John Gunn (1801 1890), now in the Castle Museum at Norwich. There are in addition a few fossils of various origins in the two museums mentioned, in the Museum of the Geological Survey, in the Natural History Museum at Ipswich and in the private collection of Mr. J. E. Sainty of West Runton. I have not seen the material of other private collections, described by Dawkins (1887). The fossils are highly mineralized, hard, and generally have a peculiar dark brown colour and a shiny surface; some of them, belonging to the older faunal elements (see later), are more ferruginous. Some are much worn, but for the most part they have a well-preserved surface. However, they consist almost exclusively of isolated bones. Only the skull of Megaceros verticornis, described as Cervus bel- grandi by Harmer (1899), was found in connection with the first two vertebrae. This specimen is also exceptional in that the upper portions of its antlers are present. In the other specimens only the more resistant lower parts have been preserved. Most of them consist of lower portions of antlers, frontals, horizontal rami of man- dibles, isolated teeth, limb bones and vertebrae. Brain cases are not uncommon, but no facial portions of skulls have been preserved, and maxillae are rare. Humeri and scapulae are represented only by the lower portions, except a heavy humerus of Libralces reynoldsi. In the text I have used the abbreviations B.M. (G.D.) for the British Museum (Natural History), Department of Geology; G.S. for the Museum of the Geological Survey of England. The registered numbers of the specimens, when quoted in the explanations of the figures have not been repeated in the text. British Museum numbers preceded by the name Savin refer to his second collection. A cknowledgments I am greatly indebted to Dr. A. T. Hopwood of the British Museum for having suggested to me the subject of this study and for his invaluable assistance in my DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 5 work. I wish to thank also Mr. W. N. Edwards, Keeper of the Geological Depart- ment of the British Museum ; Miss G. V. Barnard, formerly Curator of the Norwich Museum; Dr. C. J. Stubblefield, Chief Palaeontologist of the Geological Survey, and Mr. H. E. P. Spencer, Curator of the Museum at Ipswich, for having kindly given me facilities for studying their collections; Mr. J. E. Sainty of West Runton, who led me in a field trip on the Norfolk coast; and Dr. K. P. Oakley (London), Prof. W. B. R. King (Cambridge), Prof. F. E. Zeuner (London), Prof. I. M. Van der Vlerk (Leiden) and Dr. R. Lagaaij (Leiden) for their criticism of the chronological section of this work. GEOLOGY The Weybourn Crag and Forest Bed Series of the Norfolk Coast The Weybourn Crag and Forest Bed series, described in Reid's classical memoir (1890), is exposed at the foot of the cliffs along the Norfolk coast for some miles east and west of Cromer, and the Forest Bed is exposed also over a short distance around Pakefield, near Lowest oft. The cliffs consist of incompetent rocks marine and freshwater gravels, sands and clays overlain by glacial drift, the whole resting on a levelled surface of chalk, and erosion is extremely rapid. The exposures con- tinually change, and Reid's work is an invaluable record of many years of survey, at a time when the lesser development of coast defences made observation easier. Study of the stratigraphy is made difficult by rapid lateral changes, by breaks in the sequence, and by the discontinuity of outcrops. In many places the contorted glacial drift has disturbed this series, or cut it down to the chalk. The marine and freshwater series, to summarize Reid's data, rests on the chalk. At the junction the surface of the chalk is covered by a " stone band " of large unworn or little-worn flints; this stone band represents no stratigraphical horizon. In the country around Cromer it is overlain by the " Weybourn Crag," a false- bedded shelly sand alternating with laminated clay. This crag generally occurs in patches on the stone band, beneath high tide-mark, between Weybourn and Trimingham. It " has been traced continuously from Cromer to Weybourn, except for short distances, where it is cut out by channels of the estuarine Forest Bed, or has been ploughed out by glacial action, so that the Boulder Clay rests immediately upon the Chalk" (Reid, 1890: 139). In a section at East Runton it yielded a rich fauna of molluscs and some mammal bones. In isolated outcrops its distinction from the Forest Bed is sometimes difficult, so that many authors overlooked its occurrence. Towards the south-east the Weybourn Crag becomes more clayey and probably passes laterally into the Chillesford Clay, which in this section of the coast has been found only in borings, at Mundesley and possibly at Happisburgh. The Forest Bed series covers unconformably the Weybourn Crag and has always a more or less gravelly base. Probably there is a land surface between these two formations, " for in one place the Crag has a rather weathered appearance; but of this one cannot be certain" (Reid, 1890: 149). The Forest Bed series is in three divisions: a " Lower Freshwater Bed," composed of carbonaceous clays, silt and loam; an "Estuarine Bed" or Forest Bed sensu 6 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK stricto, composed of false-bedded carbonaceous sands and laminated clays with mammal bones and drifted tree-stumps; and an " Upper Freshwater Bed," com- posed of peaty clays and sands. The discontinuous Lower Freshwater Bed is very seldom preserved, and its observation to-day is still more difficult than at the time when Reid saw it on the shore north west of Cromer, where it cut through the Weybourn Crag (1890: 159), and at Trimingham (1890: 163). The Estuarine Bed is practically continuous. The Upper Freshwater Bed is somewhat variable in facies and its identification is not always easy. It also is discontinuous and is separated from the estuarine division by an eroded and deeply weathered surface. The Occurrence of Mammals Many of the mammalian remains were found loose on the beach, especially after storms. Some, however, were actually extracted from the rock in situ. As a rule they were said to come from the Forest Bed, sensu lato, which is still considered the chief mammal-bearing horizon; but in the course of the present work it has become apparent that they are of different ages, so that their occurrence needs more careful discrimination. Most of the fossils I have examined were obtained by A. C. Savin of Cromer, who collected for more than fifty years and gave accurate indications in his catalogue. The specimens I have determined are said to come from the Forest Bed at W. Runton, E. Runton, Cromer, Overstrand, Sidestrand, Trimingham, Mundesley, Bacton, and Pakefield; from the beach at Palling and from the Upper Freshwater Bed at West Runton. A single specimen, a fragment of a lower jaw of Megaceros dawkinsi, is said to come from the Weybourn Beds at Weybourn. The specimens from other collections were all stated to come from the Forest Bed, partly from the same localities as Savin's, partly from other locali- ties, such as Ostend, Happisburgh, the Walcot gap, and Hopton. However, not seldom there is only the indication " Forest Bed; Norfolk." A few specimens were dredged off the coast and on the Dogger Bank. It is remarkable that Savin did not distinguish an Upper Freshwater Bed in many localities where it has been recorded by Reid; possibly he was misled by differences in facies from the corresponding bed at West Runton, and as a matter of fact the distinction of these horizons in the field is not always easy. Savin dis- tinguished a Lower Freshwater Bed at East Runton, and Weybourn Beds at East Runton and West Runton, but recorded from them only fish remains. Here too he was probably misled by the common belief that mammal remains occurred only in the Forest Bed and Upper Freshwater Bed. But, as stated above, Reid found mammal bones also in the Weybourn Crag, at East Runton. Finally, Savin's attribution to the Weybourn Crag of a lower jaw from Weybourn might be based merely on the locality and might be equally incorrect. The mixed character of the " Forest Bed Fauna " led me to suppose that its members came from different horizons. The characters of the fossils, consisting of large and well-preserved antlers of deer, rules out any possibility that they were washed in. The distribution of the species I have studied shows that there is no sharp break DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 7 between the fauna of the estuarine Forest Bed and that of the Upper Freshwater Bed, which, according to my correlations, belong to the great Mindel-Riss inter- glacial (see later, section " Chronology "), whereas an older fauna may be sharply distinguished. There is no doubt that it came from an older horizon, separated from the estuarine bed by a remarkable gap. The widespread occurrence of this fauna, as well as the high ferruginous content of most of its specimens, rule out the Lower Freshwater Bed, whilst the Weybourn Crag corresponds to what may be expected to be their matrix. In fact, most of Savin's specimens are said to come from the gravel pans and sands on the shore, at some distance from the cliffs; and Mr. Sainty of W. Runton has in his private collection a beautiful lower jaw of Libralces gallicus, a typical representative of this earlier fauna, which he extracted from a patch of shelly crag directly overlying the stone band, on the shore between West Runton and East Runton, below high tide-mark. This crag, still occurring in patches but variously exposed owing to the rapidity of erosion, corresponds to Reid's description of the Weybourn Crag. Boyd Dawkins, who certainly was not influenced by prejudices about the strati- graphy, referred a skull of the same species from the Savin collection (1887, pi. i, fig. i) to the " Weybourn Beds (Forest Bed Series), East Runton, Cromer." Savin's catalogue gives for this specimen the sole indication " East Runton Forest Bed." These older species were found only in the western section of the coast, between Sidestrand and West Runton, where the Weybourn beds are exposed. Very few specimens came from Mundesley, where the Chillesford Clays are concealed beneath the beach. THE PHYLOGENY OF THE CERVIDAE The early history of the deer is little known. Probably they achieved the greater part of their evolution in Asia and North America, where the record is fragmentary and very scanty, during the Pliocene. In Europe their history is largely one of successive waves of immigration and replacement. Their classification is therefore vague. Simpson (1945) did not give an exact picture of the phyletic relationships: for instance, his Odocoileinae are not a phyletic unit, neither are his Cervinae except in so far as living species are concerned. In practice only restricted groupings of related genera can be established; the precise rank of these higher taxonomic units is largely a matter of taste. They may be called without prejudice subfamilies. Among the living deer seven sub- families are recognized: Cervinae, Muntiacinae, Odocoileinae, Capreolinae, Rangi- ferinae, Alcinae, Hydropotinae. As yet hardly any other taxonomic categories within or above these subfamilies can be established, and if we take into account also the fossil forms, things become more difficult. The present work is concerned only with the Cervinae, Capreolinae and Alcinae. The Capreolinae and Alcinae The history of the Capreolinae is quite obscure. Some Miocene forms have been tentatively referred to this group, but they are poorly known (Azzaroli, 1948: 8 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 46 ff.), and, so far as I am aware, there is no record of this subfamily during the Pliocene. The dental characters vaguely suggest some affinity with the Alcinae. The Alcinae are represented in the Quaternary by three genera, Alces, Cervalces and Libmlces, closely similar in the characters of the dentition and limb bones, but widely divergent in the skull. They are practically unknown in the Pliocene, but were already differentiated in the Pontian, where they are represented by Alces maeoticus. The Cervinae The problem of the Cervinae is less simple, and requires more explanation. In 1948 I stated that they were derived from the Pontian Pliocervinae of S.E. Europe and China; the definition of the Pliocervinae was based on Zdansky (1925, 1927). A study of the works of Alexejew and Khomenko on the Pontian deer of S.E. Europe necessitates some modification of this statement. (a) The Pontian deer from S.E. Europe In 1913 Alexejew named Procervus variabilis from Pontian deposits near Petro- vierovka, S. Russia. A more complete description followed in 1915. This species is distinctly smaller than a fallow deer; its teeth are brachyodont, the upper molars have a discontinuous cingulum, and the lower molars a well-developed Palaeo- meryx-iold. Varying with the individual, P 4 is either primitive or advanced; the upper premolars are primitive and the upper canines relatively strong. The skulls are badly damaged but seem to be of a primitive, rather unspecialized type. The brain cases are long, the supraoccipital crests weak. The pedicles are set obliquely and form two ridges at the sides of the forehead. The antlers, of various ages, are short, stout and much flattened; their form is irregular and extremely variable. They consist of a flattened, very strong brow tine, sometimes bifurcated at the top; and a short beam, gradually expanding upwards into a narrow palmation, ending in two to six small tines. The surface is deeply grooved. The first bi- furcation takes place near the burr and eventually a small accessory tine is given off from it. The antlers of younger individuals are forked, and in the first stage they consist of simple prickets. The limb bones are primitive. The shaft of the ulna is strong, the upper epiphyses of the radius and tibia are small, and the fore limb has complete lateral metacarpals, with a well-developed articulation for the corresponding toes. In 1914 Khomenko published a description of some Pontian deer from Taraklia, Bessarabia. His material was scanty and fragmentary, nevertheless he founded on them three new genera and species, Cervavitus tarakliensis, Cervocerus novo- rossiae, Damacerus bessarabiae, based on antler characters. Khomenko's specimens also included some teeth; they showed slight differences which he supposed to be specific characters. On these three genera Khomenko founded his new subfamily Pliocervinae. Khomenko's genera have been variously accepted. Zdansky (1925) supposed that Cervocerus novorossiae and Damacerus bessarabiae were synonymous; Simion- escu & Dobrescu (1941) accepted the specific distinction only, and determined other DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 9 specimens from Bessarabia as Cervocerus bessarabiae. Simpson followed these authors and recognized the genera Cervavitus and Cervocerus, and included Damacerus in the latter. However, there is little doubt that Khomenko's genera and species are all synony- mous. The differences between the antlers are due to growth stages, and the teeth do not differ by more than individual characters. The choice for the generic and specific names is thus open, and here we shall adopt Damacerus bessarabiae. Its holotype is the antler figured by Khomenko (1914, pi. 4, fig. 6), the only one of Khomenko's specimens that is fully grown. The deer from Petrovierovka is closely related to this. The teeth are indis- tinguishable; P 4 displays the same fluctuations. The f rentals, too, seem to be very similar, and the only marked differences lie in the antlers. In Damacerus bessarabiae they are less flattened, the brow tine is smaller and the first bifurcation is set high above the burr. It is consistent therefore to attribute these two species to the same genus. Procervus is preoccupied and the valid name is Damacerus, which will include therefore two species, D. bessarabiae (the genotype), and D. variabilis. (b) The Pontian deer from China In 1925 and 1927 Zdansky identified with Khomenko's Cervocerus novorossiae some deer of various Pontian localities in China. Other specimens from Shansi were referred to the same species by Teilhard & Trassaert (1937). These identi- fications are incorrect. The Chinese deer show only superficial similarities to those of S.E. Europe, but at the same time they display differences showing that they belong to completely different lineages. The Chinese so-called Cervocerus novorossiae, as defined by Teilhard & Trassaert [Zdansky 's description is sometimes vague and partly contradicted by his figures; moreover some of his attributions are questionable (see Teilhard & Trassaert, p. 38)], is a species of slightly larger size than the European Damacerus. Its skull is rather primitive; the pedicles form two prominent ridges on the sides of the forehead. The lacrimal pits are large and deep. The antlers are three-tined, cylindrical, and display a great variability in the position of the first bifurcation and in the length of the beam. Teilhard & Trassaert distinguished a " short type," with a straight beam, and a " long type," with a gently curved beam; they are linked by inter- mediate forms. The dentition is brachyodont. P 4 is primitive and there is no trace of the Palaeomeryx-iold on the lower molars. The upper premolars have the inner wall split, and a strong cingulum is stated to occur in the upper molars, but this is not to be seen in the figure. There are two complete but very thin lateral metacarpals, with rudimentary epiphyses. With the possible exception of some more flattened antlers (pi. 5, figs, i, 2), Zdansky's Cervocerus novorossiae corresponds on the whole with this species. Ac- cording to him the cingulum is variable in the upper molars, and in the lower molars he described what he regarded as the remnants of a Palaeomeryx-i old : " Es handelt sich meistens um eine, vorn mehr oder weniger scharf begrenzte Rinne. An nur io DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK ganz wenigen Zahnen ist eine solche nicht zu konstatieren." This is somewhat vague, but there is certainly no typical Palaeomeryx-iold, and no trace of it can be seen in the published photographs. Zdansky also included in the Pliocervinae the genus Procapreolus Schlosser, the validity of which has been questioned by Teilhard & Trassaert. The latter authors, on the other hand, described a Cervavitus demissus whose generic identity with the European forms is no less questionable. (c) The relationships of the Pontian Cervinae In conclusion, in the Pontian, apart from some smaller forms more or less closely related to the Muntiacinae, and from the Alcinae, the Cervidae were already repre- sented by two well-differentiated groups, namely the European Damacerus, with two species, and the Chinese three-tined deer, seemingly belonging to one species, for which new generic and specific names are required. They differed in the charac- ters of the antlers, of the dentition, and of the fore limbs. These characters enable us to make some partly hypothetical inferences about their relationships. In its dental characters the European Damacerus is closely related to the two- tined Miocene deer, and may have been derived from a genus with a well developed burr, e.g., Euprox (for full references on Miocene deer see Stehlin, 1939, with biblio- graphy). In my opinion the value of the Palaeomeryx-iold. has been underestimated. It is supposed to have gradually disappeared by the end of the Miocene, but there is no evidence for this assumption. Throughout the Miocene it does not show any ten- dency to become reduced or to fluctuate, and it is still uncertain whether it has really vanished, or has been wiped out by the extinction of the species that bore it. A solution to this question might possibly be afforded by Cervus ramosus Croizet & Jobert (1826-28, fasc. 5, pi. 5, fig. 2). The age of the type is uncertain, but the same species occurs in the Villafranchian of St. Vallier-sur- Rhone and of Villaroya ; there is no trace of the Palaeomeryx-iold in its lower molars. A more primitive variety, Cervus ramosus var. pyrenaicus, which might perhaps be the ancestor of the typical form, was described by Deperet (1897 : 99-112, 124-125) from the Upper Pliocene of the Roussillon. Among the many lower jaws which can be attributed to this variety, there are specimens with a distinct Palaeomeryx-iold and specimens without any trace of it. I am unable to state whether there are intermediate conditions, nor whether all these jaws belong to the same species ; this however is highly probable, because the other deer from the Roussillon (Deperet's Capreolus australis and Capreolus ruscinensis, but actually belonging to one species) are much smaller. If so, Cervus ramosus would afford a good example of the Palaeomeryx-iold disappearing by fluctuation. In the three-tined Pontian deer from China there was no Palaeomeryx-iold. P 4 was more primitive than in the European deer of the same age; the upper pre- molars were rather molarized, but this character always seems to be very variable and its significance is not quite clear. This is all that is known of the Upper Miocene ancestors of the Cervinae. A DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK n large part of them very probably originated from the Chinese species, and the lineage represented in the Pontian of S.E. Europe may possibly have survived with Cervus ramosus up to the Lower Pleistocene. (d) The Pliocene and Pleistocene Cervinae The history of the Cervinae during the Pliocene is obscure. Besides Cervus ramosus and its small relative from Roussillon, which might be related to Damacerus, several species of quite uncertain affinities are known. Most of them were found in the Montagne de Perrier and other localities of France. There is no evidence that any of them survived into the Pleistocene. It would be consistent to give each lineage a distinct generic name, but the use of the name Cervus, although improper, cannot be dispensed with as long as these species are so poorly known. However, the name Anoglochis, originally proposed by Bravard, Croizet & Jobert, might perhaps be adopted for Cervus ramosus. At the beginning of the Quaternary we find that the various genera which compose the subfamily are already differentiated, although in many cases represented by relatively primitive species. Euctenoceros and Megaceros make their first appearance in the Red Crag of East Anglia, Euctenoceros also in the Dutch Poederlian (see later), Dama in the Villafranchian of Tuscany (Azzaroli, 1948), Rucervus in the Pin j or stage of the Siwalik hills (to be described in a forthcoming paper by the author), Rusa in the Sanmenian of Nihowan (Teilhard & Piveteau, 1930), Sika in Chouk'outien and other Pleistocene localities (Young, 1932), Axis in the Lower Pleistocene of Shansi and Java (Teilhard & Trassaert, 1937; Stremme, 1911); Cervus s.str. makes its first appearance in the second interglacial of Europe. Ela- phurus is doubtful in the Sanmenian of Nihowan, but has been identified in the Pleistocene of Japan (Matsumoto, 1915), and is represented also by a subfossil species at Anyang, N. Honan (Teilhard & Young, 1936). Przewalskium alone is not known as a fossil. There are in addition some primitive and poorly known species which cannot be fitted into any of these genera: " Cervus " rhenanus from Tegelen, " Cervus " perolensis from the Auvergne (Bout & Azzaroli, 1953), " Cervus " punjabensis and another species from the Upper Siwaliks (to be described in a forthcoming paper), and " Cervus " philisi from Seneze (Schaub, 1942). Many of these genera and species may have been derived from the Pontian three- tined deer of China; but some highly divergent genera, as, e.g., Elaphurus and Megaceros, are very probably of a quite different origin. THE DEVELOPMENT OF THE ANTLERS (a) General Considerations Much has been written on the growth of the antlers, but our knowledge on this point is not entirely satisfactory. Each species and genus has its own peculiar pattern of antlers, obviously determined genetically, but the mechanism by which this pattern is brought into existence is practically unknown. Wislocki and others (1946, 1947) have studied the innervation and the process of ossification. It has 12 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK been established that the form of the antlers is not influenced by the course of nerve fibres, and attempts to explain the form of the antlers by the course of blood- vessels (see literature in Wezel, 1949) are equally unsatisfactory. Moreover the antlers display an individual variability of form and size that cannot be explained by the action of genetic factors alone. There is evidence that more factors, partly genetic, partly purely mechanical, influence the determination of their form. Huxley (1932) established that their size is controlled by a complicated allometric law, varying from species to species. It is known, too, that increase in size of the antlers, within a species or subspecies, is followed by a more complicated ramifica- tion. Inasmuch as this phenomenon occurs among individuals of the same genetic constitution, or even in successive growth stages of the same individual, it cannot be because of genetic differences, but rather because of purely mechanical factors : the antler tissue, growing from the top of the pedicle, seems to be unable to develop indefinitely in cross-section, but tends to divide as soon as it has reached a certain limit of bulk. A closer study of this phenomenon is obviously outside the province of palaeontology. In the recent deer, division generally takes place dichotomously, with the first bifurcation or bifurcations in a nearly longitudinal plane. This led Pocock (1933) to formulate his theory of dichotomous growth; but it may be shown that this law is not general. Each species obviously has its own range of variation, and eventually its parti- cular type of asymmetry, as, e.g., the reindeer. Broadly, the species with more complicated antlers are also the more variable. Very much has been written on the variability of antlers, especially of the red deer ; one of the most recent accounts has been given by Wezel (1949). Allowance must however be made for the un- natural conditions under which red deer very often live (inbreeding, artificial selec- tion, unfavourable or exceedingly favourable environment). In deer living under natural conditions the range of variation is not so wide. (b) History If we trace back the history of the deer we may form a mental picture of the primitive conditions of the antlers, and of the way the more advanced features gradually became established. No continuous phyletic line of an appreciable length has yet been ascertained. The principle of parallel evolution, recently exposed and discussed by Merla (1949 : 117 ff.), will be extensively applied here. This implies the more general principle of evolution controlled by internal factors, the validity of which has been ques- tioned by many recent authors who hold a purely " Darwinian " point of view. The reasons for the interpretation accepted here have been discussed at length by Merla (1949) and by Watson (1949). In my opinion the principle of evolution directed by internal factors is the only one to give a satisfactory explanation of the phyletic development of antlers. It could be argued that their development might be controlled by factors linked with characters under selective control ; this interpretation however gives no explanation DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 13 for the numerous instances of parallel evolution, nor for the polymorphism of antlers, which is in striking contrast with the uniformity of the other body characters. Apart from the evolution of antlers, there are not many clear examples of ortho- evolution of characters which are surely not under selective control. A spectacular exception is the pachyostosis of the skull of Megaceros. Lower and Middle Miocene: the primitive antlers True Cervidae, shedding their antlers, made their first appearance in Europe during the Burdigalian and remained very primitive during the Helvetian and Tortonian (for full reference see Stehlin, 1939). They form a polymorphic group. Complete and well-preserved skulls are not known, but great differences are dis- played in the position of the pedicles, and in many cases there is no typical burr. But one common feature is apparent : the antlers never grow in the form of prickets. Even in the earliest stages they tend to divide, taking the form of an irregular crown (Stephanocemas ; in fact the form is somewhat intermediate between a crown and a fork) , a fan (Palaeoplatyceros] or a longitudinal fork (Heteroprox, Dicroceros, Euprox) . In Asia some of these forms survived into the Pliocene, and may perhaps have given origin to the living Muntiacinae. Pontian. The emergence of modern types A trend towards the formation of a beam became apparent in the Pontian. Amphiprox, from Eppelsheim, was still very primitive, with short antlers consisting of a straight beam and a very small brow tine, branching off high above the burr; but at the same time, deer with more complicated antlers lived in the S.E. of Europe and in China. They all had a distinct burr and a branched beam. Damacerus is probably closely related to the earlier European deer. Damacerus variabilis seems to represent the most primitive condition. The differentiation between brow tine and beam is not very well established : they are both flattened, the brow tine is relatively large and bifurcated at the top. The general form of the antlers is very variable and irregular. The antlers of young individuals consist of prickets, but it is questionable whether the ontogenetic development actually reproduces the phyletic history. The formation of the pricket might be due to the action of genetic factors, normally leading to the formation of the beam, which in the youngster are not counterbalanced by a sufficiently strong tendency towards ramification, owing to the small size. Damacerus bessarabiae is more advanced. There is a typical beam and a much smaller brow tine. The first bifurcation is set rather high above the burr, and in my opinion this should be considered an advanced character. The point, however, is not quite clear, and palaeontological evidence of the evolution of later forms (see, e.g., Dama nestii nestii from Olivola and the Upper Valdarno; Azzaroli, 1948) may seem to contradict this assumption. As a matter of fact, genetic and mech- anical factors trending in opposite directions interact. The upwards shift of the first bifurcation is determined genetically, and the downwards shift during onto- genetic development is certainly due to a purely mechanical factor, the larger bulk i 4 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK of growing tissue inducing earlier bifurcation. This factor also may eventually act in phyletic evolution. Moreover, a third character, namely, the opening of the angle of the bifurcations, seemingly under genetic control, may also interfere; this, however, seems to have become felt only after the end of the Miocene. In the three-tined Pontian deer of China the antlers consisted of a cylindrical beam and cylindrical tines. They are rather variable, both in length and in the position of the first bifurcation. It has been stated in the preceding section that some palmated antlers figured by Zdansky might perhaps belong to another species. Pliocene and Quaternary. New evolutionary trends Although the antlers of these primitive deer were relatively small and simple, they were very variable. No attempt will be made here to explain this, but in later forms the patterns of antlers became more constant. New features appeared, as bifurcations at an obtuse angle, helicoidal tortion and undulation of the beam and tines, peculiar types of spatial arrangement of the branches (e.g., on a spherical surface in Rucervus, on a plane in Euctenoceros) , or branching of the tines. The flattening of the beam and tines is a very common feature. In some cases it may be a secondary appearance, but its phyletic origin is not always clear. This is true also of the palmation. In some instances, e.g., Alces dices, no sharp dis- tinction can be traced between palmated and non-palmated antlers. (c) On the Homologies of the Tines The discussion of the formation of antlers leads to the question of the homologies of the tines. Some words on this problem are necessary because too rigid an inter- pretation of the homologies would be misleading. After considering the work of other authors, Pocock (1933) proposed an inter- pretation of the homologies based on the theory of dichotomous growth. This can be summarized as follows. The primitive condition is assumed to be represented by the pricket. In the next stage it divides into an anterior and a posterior tine (a 1 and p 1 }. These two tines may have the same potentiality of growth and further division: this happens, e.g., in Elaphurus and Blastocerus. But generally p 1 is the stronger and divides into a 2 and p z ; then p 2 divides into a? and p 3 , etc. In accordance with this principle Pocock established the homologies between the tines of all the living genera of the Cervidae and many fossil forms. In the particular instance of the red deer the bez tine normally occurs but is not constant; this was interpreted as having originated from the division of the brow tine. In 1948 I fully accepted Pocock's views, but I now realize that rigid application of them is not possible. This theory however has the merit of having shown that there is no fundamental difference between beam and tines. It has been shown above that the pricket is not the primitive condition, and that in the earliest stages branching is more or less irregular. As a matter of fact at the beginning the tines have no morphological individuality, that is, they cannot be compared to an organ such as a tooth. What is inherited is the general pattern of antlers. As this pattern gradually evolves, the beam and the tines acquire a DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 15 certain degree of individuality, which, however, is not absolutely fixed. The evi- dence for this is that the eventual suppression of a tine, or the presence of accessory tines or even an accessory beam (a not uncommon occurrence in the red deer and reindeer), does not alter the fundamental pattern of the antler. The example of the tamin (Fig. i) is also instructive. In this species, the surface of the antlers is sometimes very scabby; the tines (except of course the brow tine) are small, and no sharp distinction can be traced between the smaller tines and the larger asperities of the surface. FIG. i. Rucervus eldi. B.M. (Zool. Dept.), no locality. natural size. True homologies can be established only between related forms, in which the evolution of the antlers has followed the same path. It is meaningless to argue whether the homologue of the brow tine of, say, a fallow deer is represented by the small inner tine of Odocoileus, or by its bifurcated anterior tine, which in turn is obviously homologous with the anterior tine of Blastocerus ; or to look for the homo- logue of the bez tine of the red deer in the reindeer and in the giant deer. SYSTEMATIC DESCRIPTIONS Genus LIBRALCES Azzaroli In another paper (1952) I have established the new genus Libralces, with genotype L. gallicus from the Upper Villafranchian of Seneze. This species is present also in the Lower Quaternary of the Norfolk coast, together with three other species of the same genus. These are more imperfectly represented, and are distinguished mainly by their size. The distinction of the dentitions is easy, but greater diffi- culties arise in determining other remains, for the most part imperfect fragments 16 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK of skulls and antlers. The attribution of some of them will therefore remain un- certain. The limb bones that can be attributed to this genus are scanty. I have denned the genus Libralces as follows (1952 : 134) : skull broad, depressed and heavily built; forehead broad; antlers palmated, with a long beam directed horizontally outwards and gently curved. Nasals long, articulated with the premaxillae. Upper canines presumably present in the genotype, molars and premolars brachyodont, lower molars with strong basal columns; traces of the Palaeomeryx-iold in M t , eventually also in M 2 ; P 4 as in Alces. Forelimb telemeta- carpal, size large. Libralces gallicus Azzaroli (Figs. 2, 3, 4 B, 5 A, 9 B, 10 D.) 1887. Alces latifrons (Johnson) Dawkins (pars), p. i, pi. i, figs, i, 3 ?, 5. 1931. Alces latifrons (Johnson) : Roman & Dareste, p. 1256. 1944. Alces latifrons (Johnson) : Schaub, p. 285. 1952. Libralces gallicus Azzaroli, p. 134. Summary Description of the Holotype and Paratype The type of this species, a complete skeleton of a full-grown male, and a second incomplete skeleton were found in the Upper Villafranchian of Seneze. The original description may be summarized as follows: A species of large size, but distinctly smaller than the living elks. The skull is broad and depressed, with a broad fore- head and a very thick roof. The face is proportionately less developed than in Alces and is characterized by the long nasals, which are articulated with the pre- maxillae. The lower molars bear well-developed basal columns, and the first lower molar has a distinct groove on the hinder surface of its antero-external crescent. The antlers consist of a long, slender beam, directed horizontally outwards and gently twisted, which ends in a small palmation, set obliquely to the axis of the body, with small tines on its edge like Alces alces. The limb bones are slender and differ from those of Alces only by their smaller size. The neck is relatively long. The Specimens from the Norfolk Coast OCCURRENCE. All the specimens that can be identified with certainty were found at East Runton and Sidestrand. Imperfect specimens, whose identification is somewhat doubtful, were found at West Runton, Cromer and Pakefield; the last are much rolled. DESCRIPTION. The best specimen is represented by a brain case with its left antler, originally figured by Dawkins (1887, pi. i, fig. i). In Azzaroli (1952, pi. 15, fig. 2) and in Fig. 2 the antler has been omitted. An adult, or nearly adult antler from Sidestrand (Dawkins, 1887, pi. i, fig. 5) and two antlers from East Runton can also be attributed to this species. All these specimens show a rather narrow range of variation. The more robust antlers (Fig. 5A, and Dawkins' fig. i) tend to develop shorter beams. A second brain case (Fig. 3), mentioned also by Dawkins, very probably belongs DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 17 FIG. 2. Libralces gallicus, skull from East Runton. B.M. (G.D.), M. 6101. natural size. (Specimen figured by Dawkins, 1887, pi. i, fig. i, and by Azzaroli, 1952, pi. 15, fig. 2.) A, occipital view ; B, lateral view ; c, basal view. FIG. 3. Libralces gallicus ? Skull of a young specimen, dredged off the Dogger Bank. B.M. (G.D.), 46108. | natural size. GEOL. II, I. 2 i8 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK to the same species. It was dredged off the Norfolk coast and is highly mineralized, like the other fossils from the Weybourn Crag and Forest Bed. This brain case is smaller than the skull from East Runton, and nearly of the same size as the holotype. The sutures are open, giving evidence that it belonged to a young animal. The correlation between the antlers, the brain cases and the teeth is made possible by comparison with the holotype. Two incomplete lower jaws from East Runton B FIG. 4. A. Libralces minor, holotype. Lower dentition, no locality. B.M. (G.D.), M. 6227. B. L. gallicus, lower dentition, East Runton. B.M. (G.D.), M. 6206. Natural size. (Figs. 4B, 10 D; Azzaroli, 1952, pi. 15, figs. 3, 4) and some isolated lower molars from Sidestrand may be attributed to this species. A fine lower jaw in the private collection of Mr. J. E. Sainty of West Runton, recorded also above (section " Geology "), was extracted in situ from the Weybourn Crag between East Runton and West Runton . The identification of more imperfect specimens from other localities will be dis- cussed later. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 19 MEASUREMENTS : E. Runton. Dogger Bank. B.M. (G.D.) M. 6101 B.M. (G.D.) 46108 Skull : Occipital breadth . 164 . 145 Occipital height ... 107 . 99 Minimal frontal breadth ca. . 212 Breadth of condyles . 82 . 69 Parietal breadth behind the pedicles ... 109 . 105 E. Runton B.M. (G.D.) M. 6206 Lower tooth-row : Total length .... 136 Length of the three molars . 80 Breadth of M 2 . . . . 19 Libralces reynoldsi n. sp. (Figs. 5 B, 6-8, 9 c.) 1891. Alces sp. f Gunn, pi. 5, fig. 4. 1934. Alces latifrons (Johnson) : Reynolds (pars), fig. 6c. HOLOTYPE. B.M. (G.D.) M. 6553: a nearly complete brain case, with the left beam (Figs. 6, 9) figured also by Reynolds, 1933, fig. 6c. Mundesley. ADDITIONAL SPECIMENS. In the British Museum: two very imperfect antlers, from Sidestrand and Mundesley; an imperfect humerus from Overstrand; a navi- culocuboid from Overstrand; two phalanges, from Mundesley and Trimingham; a fragment of a lower jaw from Overstrand. In the Norwich Museum: a lower jaw, from Cromer; two imperfect antlers, the one from Trimingham (Gunn, 1891, pi. 5, fig. 4), the second dredged off the Norfolk coast. DIAGNOSIS. A much larger species of Libralces than L. gallicus, remarkably exceeding also the size of Cervalces scotti. Skull and antlers more heavily built and relatively broader than in L. gallicus', other characters closely similar. DESCRIPTION. Libralces reynoldsi differs from L. gallicus in its much larger size, and in some features obviously correlated with this character. The skull is similarly broad and depressed, and is distinguished by the stronger development of the supraoccipital and supratemporal crests ; the latter have developed a distinct knob behind each pedicle. A close comparison between the skulls of L. reynoldsi and of L. gallicus shows that their proportions are not identical. The skull of the larger species is relatively broader: it exceeds that of the genotype by roughly I in length and height, and by ^ in breadth. The holotype of L. reynoldsi seems to have been an unusually large specimen ; the diameter of its beam above the burr is 96 mm. In comparison with the strongest specimen of L. gallicus from East Runton (52 mm.), this gives a cross-section three times as large. Apart from the strong development of the supraoccipital crest, no external features intended to counteract the weight of the antlers are seen in the skull, but the thickness of the 20 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK frontals, which are massive, measures 40 mm., exactly double that in the larger specimens of L. gallicus. The brain cavity is therefore remarkably small. In the holotype it is partly filled with the matrix, a hard sandstone impregnated with iron oxides. FIG. 5. A. Libralces gallicus, right antler from East Runton. B.M. (G.D.), M. 6554. B. L. reynoldsi, right antler from Mundesley. B.M. (G.D.), Savin 2223. J natural size. The remains of the antlers are very incomplete. Unless too much worn, they show a very strong burr and a deeply grooved surface; the beam is gently bent. In the holotype and in the antler of Fig. 5 B, which are the largest specimens I know and presumably belong to the same individual, the beam is short and the DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 21 FIG. 6. Libralces reynoldsi holotype, Mundesley. B.M. (G.D.), M. 6553. A, front view ; B, upper view ; c, lateral view (the pedicle has been broken off and the anterior part of the brain cavity is exposed). J natural size. (Specimen figured by Reynolds, 1933, fig. 6c). 22 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK flattening of the palmation is distinctly felt at 25 cm. from the burr. The fragments from Sidestrand and from the Norfolk coast are nearly of the same size, but very incomplete. The smaller antler, from Trimingham (Gunn, 1891, pi. 5, fig. 4), has a diameter of 70 mm. above the burr, and shows that the more slender specimens tended to develop a long beam, as in L. gallicus: the flattening is not felt until 35-40 cm. from the burr. The palmation is not preserved among the fossils of East Anglia, but fairly com- plete antlers, very probably of this species, have been found in Germany (see below). A maximal span of m. 2 50 has been recorded. The identification of other specimens is made possible by their size. It may be safely assumed that the muzzle of Libralces displayed the same positive allometry as the other ruminants. We can expect therefore that the .difference in size between the teeth of L. gallicus and those of L. reynoldsi is greater than between their brain cases. No doubt such a comparison is valid only between broad limits, as it has to allow for individual and sexual variations, but it can be used here, where the species under consideration differ very greatly in size. Only two imperfect lower jaws, from Cromer and from Overstrand (Fig. 7), can be attributed to L. reynoldsi. Their size exceeds that of L. gallicus by roughly \. The teeth are brachyodont, and display all the characteristic features of Libralces. A humerus showing the characters of the Alcinae (Fig. 8), evidently belongs to this species. The proximal epiphysis and half of the distal epiphysis are missing, but from the remaining part a total length of 445-450 mm. can be inferred. A naviculocuboid (Fig. 8) and two phalanges may also be attributed to this species. The naviculocuboid is very large and massive, and relatively higher than the cor- responding bone of the giant deer. The three cuneiforms are fused with it. The phalanges are long and slender as in Alces, but much larger. OTHER LOCALITIES. Several remains from Eastern and Central Europe, attributed to Alces latifrons by authors, very probably have to be identified with L. reynoldsi. Pavlow (1906: 7, pi. i, figs, i, 2) described an incomplete antler and a lower jaw from Tiraspol (Bessarabia). The teeth, although much worn, show the same size and characters as those of L. reynoldsi. The antler is intermediate in size between the antlers from Mundesley and that from Trimingham, and its beam is relatively long; this seems to confirm the inference drawn from the English specimens that more slender individuals tend to develop longer beams. Another lower jaw with the same characters has been figured by Soergel (1923, pi. 2, figs. 5, 6). It was found in the sands at Mauer, together with other teeth and limb bones, among them a metatarsal of 450 mm. total length. This, too, fits very well with the size of the humerus from Overstrand. The same author stated (1913, 1923) that an " Alces latifrons " of nearly the same size is rather common at Mosbach, whereas at Siissenborn an " Alces latifrons " of a still larger size has been found. But it is hard to imagine how an animal of this structure could have developed a size still larger than that of our species. Possibly Soergel's statement was based on com- parisons between the antlers. According to Vaufrey (1931 : 538) more specimens of "Alces latifrons " have been found recently at Mosbach. The skull reproduced in his fig. 5, although too much reduced to permit exact comparisons, seems to belong DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 23 24 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK to L. reynoldsi. This author recorded a maximal span of m. 2-50 among the fossils from Mosbach. Recently I have seen a fine lower jaw from Mosbach in the Museum of Natural History at Basel (D. 228), which in all its characters is identical with those of L. reynoldsi from the Forest Bed. FIG. 8. A. Libralces reynoldsi, right humerus, Cromer. B.M. (G.D.), M. 6464. J natural size. B & c. L. reynoldsi, left naviculocuboid, Overstrand. B.M. (G.D.), M. 6526. natural size. THE RELATIONSHIPS OF Libralces reynoldsi. Although the record of L. reynoldsi is not fully satisfactory, its affinities with L. gallicus are clearly seen. It is highly probable that L. reynoldsi is its direct descendant. These two species differ mainly in size, and increase in size is a common evolutionary trend among ungulates. The antlers of L. reynoldsi seem to have evolved following a trend already hinted in DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 25 26 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK L. gallicus, that is, increase in size together with a shortening of the beam. Other differences in the skull, as shown above, are evidently correlated with the increase in weight of the antlers. L. reynoldsi is distinctly larger than Cervalces scotti, and is the largest cervid so far known. It may be inferred that it stood m. 1-90 to m. 2-00 at the withers; its skull was very heavily built, but its limb bones were long and slender as in the other Alcinae. MEASUREMENTS : Skull of the holotype : Occipital breadth . . . . . 218 Occipital height . . . . . . 135 Minimal frontal breadth . . . . 278 Breadth of the brain case behind the pedicles . 171 Breadth of the condyles . . . . 132 Lower tooth row (Cromer, Norwich Museum. 2. 116.22) Length of the three molars . . . . 112 Breadth of M 2 . . . . . . 25 Humerus (Overstrand, B.M. (G.D.) M. 6462) Total length ...... 445-450 (inferred) First anterior phalanx (Mundesley, B.M. (G.D.) M. 6538) Total length . . . . . . 100 Proximal breadth ..... 42 First posterior phalanx (Trimingham, B.M. (G.D.) M. 6535) Total length ...... 101 Proximal breadth ..... 38 Naviculocuboid (Overstrand, B.M. (G.D.) M. 6526) Antero-posterior diameter .... 75 Transverse diameter ..... 80 Height 42 Libralces cf. reynoldsi From the Upper Freshwater Bed, West Runton. (Fig. 7 B.) A fragment of a lower jaw with the three molars, from the Upper Freshwater Bed at West Runton, is distinguished by the very poor development of the basal 2 7 columns. The groove on the anterior crescent of M x is well marked ; a shallow groove is present also on the anterior crescents of M 2 and M 3 . Lib r alee s minor n. sp. (Figs. 4 A; 10 A, E, F.) 1934. Alces latifrons (Johnson): Reynolds (pars), fig. 6a ?, 7. HOLOTYPE. A left lower jaw (B.M. (G.D.) M. 6227. No exact locality). Figs. 4 A, 10 A, 10 F. ADDITIONAL SPECIMENS. Five incomplete lower jaws (one figured by Reynolds, fig. 7), and a lower molar, from East Runton and Sidestrand. Possibly some fragments of antlers of a rather small size, from East Runton. The most complete of these fragments was figured by Reynolds, fig. 6a. Very doubtful specimens come from West Runton, Overstrand and Trimingham. (See also below.) DIAGNOSIS. A Libralces with teeth smaller and distinctly narrower than those of L. gallicus. DESCRIPTION. The teeth need no particular description. Apart from the smaller size and narrower form, they correspond in all respects to those of L. gallicus. The groove on the anterior crescent of M x is always clearly seen ; it is not very clear in Reynold's photograph, which seems to have been retouched. A similar groove in M 2 is present in three specimens. The cross-section of the mandibular ramus is also distinctly narrower than in L. gallicus (Fig. 10). The identification of the antlers is based on their size and is uncertain ; it cannot be excluded that the specimens tentatively attributed to L. gallicus are young indi- viduals of other species. The antler figured by Reynolds differs from those of L. gallicus in its smaller size, its much shorter beam and its relatively larger palmation. The other specimens from East Runton show the same characters but are still less complete. One of them (B.M. (G.D.) M. 6550), in Savin's opinion (note in the manu- script catalogue) is possibly the symmetrical part of Reynolds' specimen. The fragments of beams from Overstrand, Trimingham and West Runton are very imperfect and much worn. MEASUREMENTS : Holotype. East Runton. B.M. (G.D.) M. 6227 B.M. (G.D.) M. 6210 Lower tooth-row : Total length .... 128 . 125 Length of the three molars . 74 . 71 Breadth of M 2 .... 14 . 15 Libralces latifrons (Johnson) (Figs. 10 B, c ; u) 1874. Cervus latifrons Johnson, pi. i. 1887. Alces latifrons (Johnson) Dawkins (pars), pi. i, fig. 6. 28 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK HOLOTYPE. A left antler from Happisburgh (Norwich Museum). ADDITIONAL SPECIMENS. None surely identified, but possibly three lower jaws, from the Forest Bed of Mundesley (Figs. 10, n), Walcott (Norwich Museum), and Cromer (private collection of J. E. Sainty). For more doubtful specimens see also later. FIG. 10. A. Libralces minor, holotype. B.M. (G.D.), M. 6227. J natural size. (See also Fig. 4.) B. L. latifrons ? Lower jaw from Mundesley. B.M. (G.D.), Savin 168. J natural size. C-F. Cross-sections of lower jaws below the hinder lobus of M2. f natural size, c. L. latifrons? B.M. (G.D.), Savin 168. D. L.gallicus, East Runton. B.M. (G.D.), M. 6229. E. L. minor, East Runton. B.M. (G.D.), M. 6210. (Speci- men figured also by Reynolds, 1933, fig. 7.) F. L. minor, holotype. B.M. (G.D.), M. 6227. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 29 DESCRIPTION. This species is based on an incomplete left antler with a part of the frontal, and is ill denned. The pedicle is long and set horizontally, and the frontal is very thick, as in the other species of Libralces. The size is intermediate between those of L. gallicus and of L. reynoldsi. The antler differs from these species in having a short and straight beam; the palmation seems to have been very broad. The remaining portion of the frontal gives evidence of a larger animal than L. gallicus', on the other hand, the shortness of the beam makes it improbable that this fragment belonged to a young specimen of L. reynoldsi, but this evidence is by no means conclusive. In the collections I have seen there are also three lower jaws of a size intermediate between that of L. gallicus and that of L. reynoldsi. L. latifrons is the only species FIG. ii. Libralces latifrons? Lower dentition, Mundesley. B.M. (G.D.), Savin 168. Natural size. (See also Fig. 10.) with which I can tentatively identify them. The jaw from Mundesley (Figs. 10, n) is fairly complete. Its cross-section is remarkably narrow and deep. There is a distinct burr behind the third lobe of M 3 . A jaw from Walcott (Norwich Museum) and one found on the beach between Cromer and the Runtons (private collection of J. E. Sainty) are more imperfect. MEASUREMENTS : Mundesley. B.M. (G.D.) Savin 168 Lower tooth-row : Total length Length of the three molars Breadth of M, 88 20 30 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK Lib r alee s incertae sedis (FigS. 12 A, B, C.) As stated above, the distinction between the four species of Libralces is based mainly on the size. Many fragments therefore cannot be identified with certainty ; in some instances it is practically impossible to distinguish the smaller species from young individuals of the larger species. A brief account of the most interesting fragments will be given here. The fragments figured by Dawkins (1887, pi. i, figs. 2, 3), both much worn, may possibly belong to L. gallicus. His pi. I, fig. 4 is either L. latifrons or, more probably, L. reynoldsi. Gunn's type of Alces bovides (1891, pi. i A)may be either L. gallicus or L. minor. On his plate 5 Gunn figured many specimens attributed to Alces bovides and Alces sp. The specimen of fig. 5, much rolled, seems to belong to a small species ; fig. i is the specimen figured by Dawkins on his pi. i, fig. 2; fig. 2 is neither Alces nor Libralces; fig. 3 is Libralces gallicus (Dawkins' pi. i, fig. 5), and fig. 4 is L. reynoldsi (see above). The fragment figured by Reynolds, 1933, fig. 66, is very doubtful, and might possibly belong to Euctenoceros. Among the hitherto undescribed specimens, the following are the most inter- esting : Fig. 12 A: A fragment of a frontal with the base of the antler, showing a very strong development of the burr. It belonged to a young specimen, probably L. gallicus. The frontal is rather thin (14 mm.). Fig. 12 B : Another fragment of a frontal, with the base of the antler. The pedicle is very long and the flattening begins near to the burr. This is a young specimen of either L. latifrons or L. reynoldsi. B.M. (G.D.) Savin 417. A skull roof of a young specimen, very probably L. reynoldsi, from the Forest Bed at Sidestrand. The median suture is open and forms a median ridge. The supratemporal crests are well developed, the pedicles small, giving evidence that the antlers were not yet fully developed. In conse- quence, the frontal is very thin. Its thickness is 27 mm. on the median ridge, but on the sides of the ridge it is only 16 mm. and the thickness of the parietal is n mm. Fig. 12 c. An antler from the Forest Bed at Mundesley evidently belongs to an abnormal specimen. It bears a small anterior tine on its proximal portion and shows an abrupt bending at the middle. The characters of the surface sculpture, which shows very deep and large grooves, makes it probable that this fragment belongs to Libralces. Genus CAPREOLUS Hamilton Smith There is not much to say about this genus. As far as I know, only Capreolus capreolus has been recorded from the Middle Pleistocene. According to Soergel (1923) this species is fairly common in Germany. The specimens from Mauer, which are of a larger size than the living species, have been distinguished by this author as C. capreolus mut. prisca. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 31 ^ s ll .N O cn fl fl "* ^ T N pa n a |00 - xS| > g d S w ^ - be '^ C /s o -c "^H N d pq $ o o > ^ 5 $ SH' M O co M d DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 37 DESCRIPTION. The antlers consist for the most part of basal fragments, and differ greatly in size; some of them are very massive (Fig. 13). The bez tine is generally well developed, and in some instances is pushed upwards. In one young specimen it does not occur in its typical form, but we find here a small tine which may be interpreted indifferently as a bez or a trez (Fig. 13 D). Possibly however some fragments of this species without a bez have escaped my attention. The only fairly complete antler is of medium size and is very stout. Its upper portion ends with three tines originating from two successive bifurcations, and is damaged. This antler cannot be identified with the species of Mauer and Mosbach, but does not show the typical cup of C. elaphus. As it seems to belong to a not fully grown specimen the question of its identity is still open. FIG. 14. Cervus cf. elaphus, lower dentition, Trimingham. B.M. (G.D.), M. 6220. Natural size. Four lower jaws, from Trimingham and from the Upper Freshwater Bed at W. Runton, do not show any remarkable feature. P 4 is always advanced. The attribution of some limb bones will be discussed later. Genus EUCTENOCEROS Trouessart A definition of this genus and a discussion of its synonymy have been given in previous papers (Azzaroli, 1948; Bout & Azzaroli, 1953). In these papers I did not include in it " Cervus " falconeri Dawk., whose relationships were not clear to me. Kunst (1937) pointed out its affinities to E. teguliensis (= E. ctenoides; see below), but as a matter of fact these two species differ in many characters. On the contrary its affinities with E. sedgwicki are very close. I therefore include it in this genus. It is the oldest and most primitive of its species, but not their common ancestor. Unfortunately in 1948 I overlooked Nesti's description of the species from the Upper Valdarno, which is not quoted in Forsyth Major's papers and is recorded only in Sherborn's catalogue. Nesti exhibited the deer from the Upper Valdarno at a congress of Italian scientists held in Florence in 1841, and proposed for the larger species the names Cervus dicranios, Cervus ctenoides and Cervus orticeros. His verbal description was summarized in the report of the meeting by the secretaries, Savi & Sismonda (1841). The name Cervus orticeros has been subsequently ignored and is probably synonymous with C. ctenoides. These two species have been briefly described also by Cornalia (1858-71: 61). The incorrect spelling dicranius has been introduced by Forsyth Major. The species of this genus display very peculiar features in the antlers, which make 38 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK them easily distinguishable from other genera. The first tine branches from the outer anterior part of the beam ; the following tines arise from its anterior part and lie in a plane. They are numerous, three in E. falconeri, four or five in the other species, and are frequently more or less undulated. Beam and tines display a tendency to become flattened and in some species the tines subdivide further. The flattening is generally more pronounced in the upper portion of the antlers, but in E. sedgwicki reaches as far down as the brow tine. In the larger specimens of some species a small accessory tine, directed upwards, is given off from the first bifurca- tion. With the sole exception of E. falconeri all the species of this genus are of a large size. Together with the common features described above, they display a remark- ably wide range of variation which makes specific distinctions difficult. I do not feel satisfied with some of the determinations I published in 1948. An antler from Olivola (fig. 8) then attributed to E. ctenoides is actually of E. dicranios. The tines of this specimen do not show the strong backward bending of the lectotype, but in its general characters this antler is more similar to it than to E. ctenoides. Moreover, my reconstruction was wrong: the lower tine preserved in this antler corresponds to the second tine, not to the third tine of the lectotype of E. dicranios. E. ctenoides and E. teguliensis are very probably identical. The antlers do not afford any good basis for distinction. Eiictenoceros tetraceros (Dawkins) (Figs. 15, 16, 17 A, B, D.) 1878. Cervus tetraceros Dawkins, p. 416, figs. 14, 17. 1891. " Cervus tetraceros Mackie " : Newton, p. 32, pi. 4, fig. 2. 1953. Euctenocevos tetraceros (Dawk.) : Azzaroli in Bout & Azzaroli, p. 43, figs. 1-6. OCCURRENCE. Fairly common at East Runton, rare at Overstrand. DESCRIPTION. This species is represented by two fine antlers (Fig. 15) and several fragments. They all agree with the specimens from Peyrolles; one specimen only (Fig. 16) is distinguished by the abnormal inward bending of its brow tine. As at Peyrolles, all the antlers of this species have been naturally shed. Three lower jaws show the same characters as those from Peyrolles. Probably also a fourth lower jaw with a primitive P 4 belongs to this species. MEASUREMENTS : East Runton. East Runton. East Runton. Lower tooth rows : B.M. (G.D.) M. 6232 B.M. (G.D.) M. 6218 B.M. (G.D.) M. 6221 Total length . . . 131 . 134 . 133 Breadth of M 2 . 16 . 15 . 15 Euctenoceros ctenoides (Nesti) 1841. Cervus ctenoides Nesti (in Savi & Sismonda, p. 159). 1858-71. Cervus ctenoides Nesti : Cornalia, p. 61. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 39 B FIG. 15. Euctenoceros tetraceros, East Runton. J natural size. A. Left antler, B.M. (G.D.), M. 6369. B. Right antler, B.M. (G.D.), M. 6370. 40 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 1887. Cervits savini Dawkins (pars), pi. 3, fig. 4. 1948. Cervus (Euctenoceros) ctenoides Nesti : Azzaroli (pars), p. 62, figs. 6, 7, 9. 1947. Eucladoceros teguliensis Dub. : Hooijer, p. 34 (cum syn.). SYNONYMY. The distinction between E. ctenoides and E. teguliensis does not seem to be valid. The ranges of variation of these two species widely overlap. The antlers from Tegelen do not attain the size of the larger specimens from the Upper Valdarno, but this may be due merely to the incompleteness of the record. The dentition and limb bones are of the same size. FIG. 1 6. Euctenoceros tetraceros, abnormal right antler, East Runton. B.M. (G.D.), M. 6399. J natural size. Among the specimens from East Anglia a very fine antler from Sidestrand (Dawkins' pi. 3, fig. 4) so closely resembles the lectotype of E. ctenoides that no one would hesitate to identify it with this species. OCCURRENCE. The only good evidence of the occurrence of this species is given by the antler from Sidestrand figured by Dawkins, who incorrectly attributed it to Cervus savini, and by some basal fragments from East Runton, of a slightly smaller size. For the identification of other fragments of antlers and of dentitions see later. Euctenoceros sedgwicki (Falconer) (FigS. l8, IQ, 20 A, 21 A, F.) 1868. Cervus sedgwicki Falconer (ex Gunn MSS.), p. 472, pi. 37, figs. 1-3. 1882. Cervus sedgwicki Falc. : Newton, p. 60. 1891. Cervus sedgwicki Falc. : Gunn, pi. 3, figs. 99, 100. 1891. Cervus sp. Gunn, pi. 3, figs. 105, 106. 1891. Cervus sedgwicki Falconer : Newton, p. 31. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 41 OCCURRENCE. This species is fairly common, but is represented for the most part by very imperfect fragments. The only satisfactory specimen is the holo- type (Fig. 18). This and most of the other fragments were found at Bacton, a few specimens at Mundesley and a much rolled fragment at East Runt on. For doubtful fragments see also below. FIG. 17. Euctenoceros ; dentition, East Runton. Natural size. A. E. tetraceros, B.M. (G.D.), M. 6218 (advanced form). B. E. tetraceros, B.M. (G.D.), M. 6221 (primi- tive form), c. E. ctenoides ? B.M. (G.D.), M. 6241 (primitive form). D. E. tetraceros, B.M. (G.D.), M. 6206 (advanced form). E. E. ctenoides? B.M. (G.D.), M. 6213. DESCRIPTION. This species is of a large size and is characterized by the high position of the brow tine, 10-15 cm - above the burr, and by a very marked flattening of the upper portion of the beam and of the tines, which divide into three or four secondary tines. The brow tine is also much flattened, and in the type it divides into four points. The high position and the flattening of the brow tine are remark- ably constant, and are well displayed even in young specimens (Fig. 21 F). The pedicles are set near one another, as in E. boulei and E. dicranios. 42 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK FIG. 1 8. Euctenoceros sedgwicki, holotype, Norwich Museum, Gunn Colin. No. 99. T ^ natural size. FIG. 19. Euctenoceros sedgwicki, reconstruction of the holotype. Norwich Museum, Gunn Colin. No. 99. J natural size. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 43 The upper portion of the type antler is badly damaged and its reconstruction is rather puzzling. On Fig. 18 I have indicated the most probable position of the fragments. The small cross-section of the beam at the broken surface makes it unlikely that there were additional tines. On Fig. 19 I have attempted a recon- struction of this specimen. COMPARISONS. Euctenoceros falconeri (Dawkins, 1868; see also Kunst, 1937: 101) is a species of medium size. It does not occur in the Weybourn Crag nor in the Forest Bed, but has been found in the Red Crag and in the Norwich Crag. FIG. 20. Euctenoceros sedgwicki and E. falconeri, lower portions of the antlers. natural size. A. E. sedgwicki, Mundesley. Norwich Museum 323. B. E. falconeri, Thorpe, Norwich Crag. Norwich Museum 313. c. E. falconeri, Horstead, Norwich Crag. Norwich Museum 310. It occurs also in the Poederlian of the Netherlands and in deposits of the same age in Belgium. The most complete specimens were found in the Belgian Kempen. The remains from East Anglia are rather fragmentary ; the most complete are the holotype (a young individual), and a crushed full-grown antler from the Red Crag, now in the Museum at Ipswich. In the British Museum and at Norwich I have seen other fragments from the Norwich Crag, and at Ipswich fragments from the Red Crag. The remains of the Norwich Crag may possibly have been derived from the Red Crag. Those from the Red Crag bear no exact indication of the horizon, but their state of fossilization shows that they were not derived. Two specimens in the Norwich Museum were figured by Gunn (1891, pi. 2, fig. 44 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK *i-oo S ^ o o -g 'c o< R O, Q -g p ^ . ^ C5 ffl-g-gfeffl ,30 C .. > W O . I > < O r* f"5 )-i P^ 2I fig. 1-8) is also a Megaceros. I do not feel certain of this. The lower jaw (Gervais' fig. i) is certainly of a giant deer, but the antlers seem rather to belong to a relative of Euctenoceros sedgwicki. Megaceros messinae Pohlig (1909) from Sicily seems to be another dwarf island form. Megaceros verticornis (Dawkins) (FigS. 23 A, B, 24-27, 28 M, N, 29 E, F, 30, 32 B, C.) 1872. Cervus verticornis Dawkins (pars), p. 405, fig. 2. 1882. Cervus verticornis Dawkins : Newton, p. 61. 1887. Cervus verticornis Dawkins : Dawkins (pars), pi. 5, figs. 1-3 ; pi. 6, figs, i, 2 ; pi. 7, figs, i, 2. 1891. Cervus verticornis Dawkins : Gunn (pars), pi. 2, fig. 97. 1891. Cervus verticornis Dawkins : Newton, p. 32. 1899. Cervus belgrandi Lartet : Harmer, p. 97, pi. 21. SYNONYMY. The definition of this species has been discussed by Newton. Dawkins, in his original description, included in it also an antler on which Newton later based his Cervus dawkinsi. Newton chose as the type of C. verticornis the basal fragment of an antler from Pakefield, figured by Dawkins (1872, fig. 2). In his later description (1887) Dawkins again included in C. verticornis some remains of other species. On p. 23 he recorded seven specimens with a double basal tine; exact reference was made only to a specimen in the British Museum, registered B.M. (G.D.), 33471. This register number includes a small sample of very imperfect fragments, none of which belongs to Megaceros verticornis. A double basal tine is quite exceptional in this species: I do not know the other specimens quoted by Dawkins, but as yet I have seen only a fragment of a young individual from Trimingham with a well-developed accessory basal tine (Fig. 26). This specimen was found after the publication of Dawkins' works. The antler from the Red Crag at Trimley figured by Dawkins (1887, pi. 6, fig. 3) is imperfect and cannot, since it is older, be identified confidently with M. verticornis. The fragment from Kessingland figured by Gunn (pi. 7, fig. 2) is doubtful ; it does not seem to belong to M. verticornis. OCCURRENCE. This is the commonest species of Cervidae in the Forest Bed and has been recorded from nearly all the fossiliferous localities : Kessingland, Pakefield, 54 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK Mundesley, Overstrand, Happisburgh, Trimingham, Hopton, Sidestrand, Bacton, Cromer; it is common also in the Upper Freshwater Bed at West Runton, but is not recorded at East Runton. The remains of this species consist for the most part of basal fragments of the antlers, either naturally shed or with fragments of the frontals. There are in addition two complete brain cases, one in the Norwich Museum (Figs. 24, 25), the second in the British Museum (Fig. 27). The latter was found in connection with the atlas, the axis and the antlers, nearly perfect, and has been described by Harmer under the name Cervus belgrandi, of which Cervus verticornis was incorrectly supposed to be a synonym. A shed antler, in the Museum of the Geological Survey, figured by Dawkins (1887, pi. 5, fig. 2), is broken at the beginning of the palmation. The correlation between skulls and teeth is based on the size, on their occurrence together in the Upper Freshwater Bed at West Runton, where no other species of Megaceros has as yet been found, and on comparison with M. aff. verticornis from Siissenborn. The dentition is represented by some lower jaws and maxillae. Some limb bones may also possibly belong to this species. DESCRIPTION. Skull: There is very little to add to Harmer's description of the skull from Pakefield. Harmer pointed out its large size and heavy form, its flat forehead and its divergent and widely spaced pedicles. The supraorbital foramina are very large and round. Two very faint ridges, starting from the coronal suture, run along the inner margins of the pedicles and vanish on the forehead. In other specimens these ridges are better developed (Fig. 25). Harmer also pointed out the lack of a transverse swelling of the frontal between the pedicles. The other remains of skulls display little individual variations. The pedicles are generally widely spaced, but in a frontal from Pakefield (Figs. 23 A, 25 D) they are set near each other and less divergent than in the average specimens. The frontal is still flat. Antlers: The characters of the lower portion of the antlers are remarkably con- stant. The brow tine arises at some distance above the burr, from the upper portion of the beam, and is strongly bent dowwards. It is rounded in cross-section and very long in a skull from Kessingland (Dawkins, 1887, pi. 7, fig. 2), the only specimen where it is complete. In many specimens there is a more or less marked knob below it, on the anterior side of the beam; this may be present on both sides or on one side only and is very variable. In a frontal from Trimingham, very much worn, it is very strong on the left side and seems to be completely lacking on the right side, but only in a young antler (Fig. 26), also from Trimingham, has it developed into an accessory lower tine. Dawkins' statement that an accessory lower tine occurs in seven other specimens has not been fully checked, but is based partly at least on incorrect identifications. The beam is long and bears an anterior and a posterior tine, ovoidal in cross- section. The position of the anterior tine is somewhat variable (see Figs. 25 c, 27, and the figures published by Dawkins, 1887). An additional anterior tine has been recorded only in one instance (Dawkins, 1887, pi. 5, fig. 3). Above the posterior tine the beam is twisted upwards and becomes flattened, and after a short distance DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 55 56 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK FIG. 25. Megaceros verticornis, skulls. A. Inferior aspect, from Kessingland. Norwich Museum 364. (See also figs. 23 & 24.) B. Posterior aspect, same specimen. J natural size. c. Anterior aspect. West Runton, Upper Freshwater Bed. B.M. (G.D.), M. 6099. f natural size. D, Anterior aspect. Pakefield, Norwich Museum 365. $ natural size. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 57 it suddenly expands into a palmation. This palmation is preserved only in Harmer's specimen, so that we have no information on its variations. It is very broad and thin and of a quite unusual pattern : its margin does not give origin to long tines as in most of the giant deer, but is scalloped and gently bent inwards. The anterior edge was destroyed on both sides. The plane of the palmation is nearly vertical FIG. 26. Megaceros verticornis, young specimen with an abnormal lower tine, Trimingham. B.M. (G.D.), Savin 713. natural size. FIG. 27. Megaceros verticornis, reconstruction of the specimen described by Harmer (1899), Pakefield. B.M. (G.D.), M. 11352. T x natural size. and set at 45 with the median plane of the body. The total span in the reconstructed specimen is 228 cm. Harmer also pointed out the rectilinear course of the blood vessels. Harmer's specimen is rather large, but the lectotype and the skull from the Upper Freshwater Bed of West Runton (Fig. 25 c) slightly exceed it. The antler figured by Dawkins, 1887, pi. 5, fig. 2, the most complete beyond Harmer's speci- men, is on the contrary distinctly smaller. 58 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK Dentition: The remains consist of a maxilla and two lower jaws from the Upper Freshwater Bed at West Runt on, a maxilla from Trimingham and some imperfect lower jaws from Pakefield, in the British Museum; a maxilla from Kessingland and probably two lower jaws, from Copt on and Kessingland, in the Norwich Museum. M FIG. 28. Cross-sections of lower jaws, below hinder lobe of M2. External side to the left, f natural size. A-I. Megaceros dawkinsi. A. Mundesley, B.M. (G.D.), M. 6335 (young). B. Overstrand, B.M. (G.D.), Savin 533. c. Sidestrand, B.M. (G.D.), M. 6224. D. Overstrand, B.M. (G.D.), M. 6209. E. East Runton, B.M. (G.D.), Savin 304. F. Mundesley, B.M. (G.D.), M. 6207 (old). G. Overstrand, B.M. (G.D.), Savin 526. H. Mundesley, B.M. (G.D.), Savin 339. i. Pakefield, B.M. (G.D.), M. 6501. j. Cervid indet., Overstrand. B.M. (G.D.), Savin 421. K & L. Megaceros savini ? K. Trimingham, B.M. (G.D.), M. 6220. L. Mundesley, B.M. (G.D.), Savin 1008 (old). M & N. Megaceros verticornis. M. Overstrand, B.M. (G.D.), Savin 1198. N. West Runton, Upper Freshwater Bed. B.M. (G.D.), Savin 1069. The hyperostosis of the lower jaw is moderate (Fig. 28). The teeth are relatively small and narrow. The basal columns are moderately developed, and in the maxilla from West Runton the upper premolars show a very faint trace of a cingulum. P 4 is somewhat intermediate between the primitive and the advanced condition. The enamel is moderately thick, the height of the crown also moderate. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 59 B D FIG. 29. Megaceros, lower dentition, ^f natural size. A. M. dawkinsi, Overstrand. B.M. (G.D.), Savin 533. B. M. dawkinsi, Mundesley. B.M. (G.D.), M. 6231 (young), c. M. savini ? Trimingham, B.M. (G.D.), M. 6207. D. M. savini ? Trimingham, B.M. (G.D.), Savin 439. E. M. vevticornis, West Runton, Upper Freshwater Bed. B.M. (G.D.), Savin 1069. F. M. verticornis, West Runton, Upper Freshwater Bed. B.M. (G.D.), Savin 1265. 60 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK VERTEBRAE. Atlas and axis were found in connection with Harmer's specimen. They have already been described by Harmer and are slightly smaller than the corresponding bones of Megaceros giganteus, but very similar in shape. COMPARISONS. The giant deer from Siissenborn, figured by Soergel (1927, pi. 17, fig. 2 ; pi. 18, figs. 2, 4, 7), is closely related to M. verticornis. The skull, the denti- tion and the lower portion of the antlers are identical. The upper portion of the antlers in the specimen from Siissenborn is flattened and gives off two large tines from the anterior edge ; the posterior portion is destroyed. In Harmer's specimen from the Forest Bed the anterior edge of the palmation is missing on both sides, but what remains of it on the right side is very thin (15-20 mm.) and is not likely to have developed large tines, which moreover would be in contrast with the general pattern of the palmation. The systematic value of these differences is not known, as we have no idea of the range of individual variation of these forms, but no doubt they are very closely allied. FIG. 30. Megaceros verticornis, lower molars, Pakefield. B.M. (G.D.), M. 6223. Natural size. Megaceros mosbachensis, of which only a brain case with the lower portions of the antlers and some fragments of the dentition are known, also closely resembles M. verticornis in the characters of the skull, but differs in the antlers and teeth (see above). The affinities between M. verticornis and M . dawkinsi are also very close and will be discussed later. The lower jaw of M. dupuisi differs from that of M. verticornis by its smaller size and the more primitive form of P 4 . The affinities between M. verticornis and M. giganteus have been discussed by Harmer and by Soergel. It has already been shown that although these two species are related they belong to different lineages. MEASUREMENTS OF THE TEETH : Upper tooth row : Total length Breadth of M* B.M. (G.D.), Savin 1156 W. Runton, U. Freshw. Bed 124 22 6i B.M. (G.D.), B.M. (G.D.), Savin 1069 Savin 1265 Lower tooth row : W. Runton, U. Freshw. Bed Total length . 146 . 152 Breadth of M 2 . 17 . 18 Megaceros dawkinsi (Newton) (FigS. 23 C, 28 A-I, 29 A, B, 31 A, 32A, 33-36.) 1872. Cervus verticornis Dawkins (pars), p. 405, fig. i. 1882. Cervus dawkinsi Newton, p. 54. 1882. Cervus fitchii Gunn MS. : Newton, p. 56. 1882. Cervus gunni Dawkins MS. : Newton, p. 57. 1887. Cervus dawkinsi Newton : Dawkins, p. 7, pi. 2, figs, i, 2, pi. 3, fig. 3. 1891. Alces ? : Gunn, pi. 5, fig. 6. 1891. Cervus fitchii Gunn, pi. 6, fig. i. 1891. Cervus dawkinsi Newton : Newton, p. 26. 1891. Cervus fitchii Gunn : Newton, p. 28. SYNONYMY. This species was founded by Newton on a young antler, formerly attributed to Cervus verticornis by Dawkins. Other fragments determined as Cervus fitchii and Cervus gunni by Newton belong^to^the same species, as already pointed out by Dawkins. Newton (1891) rejected this view, but in my opinion he overestimated differences due to individual variation. OCCURRENCE. This species is nearly as abundant and widespread as Megaceros verticornis and has been recorded from Bacton, Cromer, Trimingham, Sidestrand, Mundesley, Overstrand, Pakefield, Walcot and East Runton (one specimen, much rolled). A specimen in the Savin collection was found at Weybourn and was supposed to come from the Weybourn Crag; its occurrence has been discussed in the previous section. M. dawkinsi has not been recorded from the Upper Fresh- water Bed. The specimens consist of several portions of antlers and fragments of the f rentals ; a specimen from the Walcot gap, in the museum at Norwich, bears also a part of the left parietal and temporal bones, but no complete brain cases have been found. The correlation between antlers and teeth is based on the size. The dentition is represented by several lower jaws and few fragments of the maxillae. DESCRIPTION. The skull is distinctly smaller than that of Megaceros verticornis, but is represented by rather unsatisfactory specimens, so that exact measurements are possible only for the frontals. Its most striking feature is given by the convex form of the forehead and by the backward inclination of the pedicles. In the adult the pedicles are short, very massive and strongly divergent. Two shallow ridges on their inner borders, as in M. verticornis, are always distinct. The skull roof of the specimen reproduced on Fig. 33 A is pierced by numerous small openings; its inner surface in this region has been filled with plaster. Similar openings have been described also by Soergel (1927) in the giant deer from Mosbach and Siissen- born ; they were interpreted as canals for blood-vessels. 62 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK The antlers, of which the upper portion is unknown, are stout, palmated, and very variable. The beam is short and rounded. Its base, unlike that of M. verticornis, does not grow in the same direction as the pedicle but is stretched outwards and backwards, and is thinner than the pedicle. This is well seen also in a young speci- men reproduced on Fig. 33 c. The brow tine and the accessory lower tine are very variable, but are always more or less reduced. The type affords the only instance of a well-developed lower accessory tine. Very often both these tines have dis- appeared. The upper tines are moderately flattened and gently twisted upwards. FIG. 31. Megaceros, lower jaws. natural size. A. M. dawkinsi, Overstrand. B.M. (G.D.), Savin 526. B. M. savini ? Trimingham, B.M. (G.D.), Savin 439. c. M. savini ? Mundesley, B.M. (G.D.), Savin 1008. There may be one or two anterior tines, and a posterior tine is not always present. The upper portion is more or less abruptly twisted upwards, sometimes resembling features of M. verticornis, sometimes as in Fig. 36, rather recalling M . giganteus. In the specimen shown on Fig. 23 c and 33 A the branching of the right antler takes place in two different planes: this is probably due to an unusual shortening of the beam between the anterior tine and the beginning of the palmation. The teeth I refer to this species consist of several lower jaws, a fragment of a maxilla and some isolated molars. The dental characters recall those of M. verti- cornis; the teeth are however smaller, narrower, very hypsodont, and have a thick enamel. The lower molars bear strong basal columns and anterior folds. P 4 is DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 63 always very primitive and is characterized by the poor development of its hinder portion ; the latter is bordered by a small burr near the base of its outer wall. The hyperostosis of the lower jaw is well marked (Fig. 28). AFFINITIES. Dawkins (1887) pointed out the general resemblance of the antlers with those of M. giganteus, but concluded that the affinities between these two species are not very close. As he limited his observations to the antlers, he failed to perceive the affinities between M. dawkinsi and M. verticornis, which moreover was imperfectly known until the discovery of Harmer's specimen, ten years later. FIG. 32. Megaceros, upper dentition. Natural size. A. M. dawkinsi, Trimingham, B.M. (G.D.), Savin 7. B. M. verticornis ? Trimingham, B.M. (G.D.), Savin 942. c. M. verticornisy West Runton, Upper Freshwater Bed. B.M. (G.D.), Savin 1156. The characters of the skull leave little doubt that Megaceros dawkinsi belongs to the group of M. verticornis. The antlers support this conclusion : it has been shown that a small brow tine, similar to that of M. verticornis, is not uncommon. This never occurs in the group of M. giganteus. However, Megaceros dawkinsi, in spite of its small size, is not primitive, and does not belong to the same lineage as M. verticornis. The dentition displays simul- taneously primitive and advanced features; the lower pre-molars are perhaps of the most primitive pattern yet known, but both the hypsodonty and the thickness of the enamel are no doubt advanced features. There is no hint of the broadening 64 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 65 GEOL. II, I. 66 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK of the molars, undergone by Megaceros giganteus and M. mosbachensis. Nor can the antlers be considered primitive. The pedicles are disproportionately strong for their size; moreover, in cross-section the beam is thinner than the pedicle and is FIG. 35. Megaceros dawkinsi, left antler, seen obliquely from the outer side. Trimingham, B.M. (G.D.), Savin 1820. natural size. FIG. 36. Megaceros dawkinsi, reconstructions of the antlers based on the specimens of Figs. 33 & 34. natural size. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 67 stretched backwards two common features in the deer with reduced antlers. The unusually high instability of the tines is also a character peculiar to regressed antlers. Whether this regression of M . dawkinsi was due to inner factors or to influence of the environment can hardly be stated. The characters of the dentition suggest habits of life in open, grassy lands, rather than in woodlands; this is a rather unusual habitat for deer. Megaceros belgrandi and several related forms from Central and Eastern Europe, listed above, are closely similar to M. dawkinsi and possibly identical with it, but their remains are very imperfect. A striking similarity with the antlers of M. dawkinsi is shown by an antler of Cervus cazioti from the cave of Nonza in Corsica, figured by Deperet (1897^), which, however, is less than half the size of the antlers of M. dawkinsi. Cervus cazioti is no doubt a dwarf island form. Also its dentition recalls M. dawkinsi. MEASUREMENTS : B.M. (G.D.) M. 6302 Frontal Sidestrand Minimal breadth beneath the pedicles . 164 B.M. (G.D.), B.M. (G.D.), B.M. (G.D.), B.M. (G.D.), Savin 526 M. 6224 Savin 339 Savin 533 Lower jaws Overstrand Sidestrand Mundesley Overs trand Total length of the tooth row . 128 . 134 . 125 . 123 Breadth of M 2 .... 15 . 15 . 15 14, 5 Megaceros savini (Dawkins) (FigS. 23 D, 28 K, L, 29 C, D, 37-40-) 1887. Cervus savini Dawkins (pars), p. n, pi. 3, fig. 3, 5 ? 1891. Cervus savini Dawkins : Gunn (pars), pi. 7, figs. 4, 5. 1891. Cervus sp. Gunn, pi. 4, fig. 101, pi. 6, fig. 2. 1891. Cervus savini Dawkins : Newton, p. 30. SYNONYMY. Dawkins included in this species also an antler of Euctenoceros ctenoides (1887, pi. 3, fig. 4) and a fragment of a young specimen whose identity is uncertain (pi. 3, fig. 2), but Newton pointed out that these identifications are incorrect. Gunn attributed to it a frontal (pi. 4, fig. 102), which might possibly belong to a red deer. OCCURRENCE. This species is fairly frequent and is represented by not less than twenty antlers in the British Museum and at Norwich. Eleven of them were found at Trimingham, the others at Sidestrand, Overstrand, Mundesley and Kessingland. A frontal from Kessingland (Figs. 23 D, 37) and possibly another frontal from Pake- field (Dawkins, 1887, pi. 3, fig. 5) represent all that is known of its skull. The remains of dentition I tentatively refer to this species consist of three lower jaws from Trimingham and Mundesley, in the British Museum. 68 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK DESCRIPTION. The frontal, which indicates an animal of about the same size as Megaceros verticornis, clearly shows the hollowed forehead of the group of Megaceros giganteus. The identification of the frontal (Figs. 23 D, 37) is made possible by the basal portion of the antler. The antlers include specimens of all ages. The type (Dawkins, 1887, pi. 3, fig. 3) is of medium size, but the largest specimens, represented by basal fragments (Fig. 38 A, and a more imperfect fragment in the Norwich Museum, from an unknown FIG. 37. Megaceros savini, frontal, Kessingland. Norwich Museum 321. (See also Fig. 23.) J natural size. locality), attained the size of Megaceros verticornis and M. giganteus. The beam is flattened and hollowed below and bears a prominent ridge on its anterior side. The brow tine (Fig. 38 c, D, and Gunn, 1891, pi. 6, fig. 2) branches off close to the burr; it is flattened, broadly expanded into a small palmation, and set hori- zontally. It is very often broken off at the base, and I do not know any specimen where it is complete. The section of its base is triangular, with the longer side above. The antlers are widely divergent, as in M. giganteus and in M. verticornis (Fig. 39). The anterior and posterior tines are much flattened; above the posterior tine the beam is bent upwards, becomes still more flattened and branches into two terminal tines, apparently of equal size. The teeth I tentatively refer to this species are intermediate in size between those of M. verticornis and those of M. dawkinsi. The distinction is not sharp, and in DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 69 their extreme variations these species, especially M. verticornis and M. savini, may perhaps overlap. The lower tooth row is identical with that of Megaceros dupuisi (Stehlin, 1912). P 4 is primitive. The teeth I have seen are rather worn, but they do not seem to be hypsodont. The enamel is of medium thickness. FIG. 38. Megaceros savini, antlers. J natural size. A & B. Trimingham, large speci- men. B.M. (G.D.), Savin 1422. c & D. Overstrand, medium sized specimen. B.M. (G.D.), Savin 1880. E. Trimingham, young specimen. B.M. (G.D.), Savin 1241. Two imperfect maxillae from Trimingham might perhaps belong to this species. MEASUREMENTS : B.M. (G.D.), M. 6220 Lower jaw : Trimingham Total length . . . 135 Breadth of M, 16 70 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK FIG. 39. Megaceros savini, reconstruction of the holotype, B.M. (G.D.), M. 6093, and of the specimens of Fig. 38. ^ natural size. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 71 FIG. 40. Megaceros savini ? Very young specimen. Bacton, B.M. (G.D.), Savin 910. \ natural size. FIG. 41. Megaceros, limb bones. natural size. A. M. verticornis or savini. Right radius. No locality, B.M. (G.D.), Savin 1462. B. M. verticornis or savini. Right metacarpal, Trimingham. B.M. (G.D.), Savin 1104. c. M. dawkinsi ? Right metacarpal, Mundesley. B.M. (G.D.), Savin 2024. D. M. verticornis or savini, Left metapodial, Trimingham. B.M. (G.D.), Savin 1450. 72 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK Megaceros incertae sedis (Fig. 41 A-D.) LIMB BONES. Fourteen complete limb bones may be attributed to the genus Megaceros for their large size and heavy form. They are: two radii, three meta- carpals and four metatarsals in the British Museum; a metacarpal and a tibia in the Norwich Museum ; a radius, a metacarpal and a metatarsal in the Museum of the Geological Survey. The metapodials are characterized by the broad form of their distal articulations and are easily recognizable. The larger bones are of the same size as Megaceros giganteus and may be attributed to M. verticornis or to M. savini ; the smaller ones probably belong to M. dawkinsi, or to females of the larger species. MEASUREMENTS : Radius : Total length Proximal breadth Distal breadth B.M. (G.D.), Savin 464. Trimingham 34 79 B.M. (G.D.), Savin 1462. (No locality) 349 82 73 G.S. 21688. Kessingland 320 Metacarpal : Total length Proximal breadth Distal breadth B.M. (G.D.), Savin 1104. Trimingham 304 64 68 B.M. (G.D.), Savin 376. Sidestrand 293 56 60 B.M. (G.D.), Savin 2024. Mundesley 286 63 Total length Proximal breadth Distal breadth G.S. TF/55. Mundesley 271 58 Norwich Museum 531- (No locality) 310 70 74 B.M. (G.D.), Savin 1885. Sidestrand 261 48 52 Tibia Metatarsal : Total length Proximal breadth Distal breadth Total length . Distal breadth G.S. 21684. Kessingland 348 57 65 Norwich Museum 526. (No locality) 482 81 B.M. (G.D.), M. 6491. Trimingham 358 59 67 B.M. (G.D.), Savin 1450. Trimingham 345 55 64 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 73 CERVIDAE incertae sedis " Cervus " obscurus n. sp. (Figs. 42-45-) SYNTYPES. B.M. (G.D.), Savin 195, Mundesley (Fig. 42 A, B); B.M. (G.D.), M. 2321, Forest Bed : no exact locality (Fig. 42 C-E) ; B.M. (G.D.), M. 6421, Overstrand (Fig. 43). ADDITIONAL SPECIMENS. Two lower portions of shed antlers and three basal fragments of antlers with the f rentals; two of these (Fig. 44 c, D) possibly belong to the same individual. OCCURRENCE. Bacton, Mundesley, Sidestrand and Overstrand. In addition, an antler of doubtful attribution from Trimingham. DIAGNOSIS. A cervid of large size characterized by a small lower basal tine directed downwards, a large upper basal tine strongly twisted outwards, a massive beam and an upper tine set far apart from the basal tines. DESCRIPTION. This is a species of very unusual features. The size is large and the forms are massive as in Euctenoceros dicranios and ctenoides. The forehead is marked by a prominent median suture and two shallow ridges along the inner borders of the pedicles, as in Megaceros verticornis and its relatives, but is narrower. A small tine is given off close to the burr, at a wide angle with the beam, and is inclined inwards and downwards. A second basal tine, much more robust, arises at some distance from the burr and is strongly bent outwards. It is rather long and is flattened at its end. The beam is massive, straight, and ovoidal in cross- section, with the major axis set vertically. A third tine, directed upwards, is given off at a very great distance from the burr. The grooves of the blood-vessels are shallow. The remaining parts of the frontals enable one to reconstruct the position of the antlers. The beams diverge at about 45 from the median plane of the body. The antler from Trimingham which I doubtfully identify with this species (Fig. 44 A, B) differs in the characters of the lower tines. The lower tine arises at some distance from the burr; a second tine arises from the inner side, near its base. This tine corresponds for its position to the small knob between the two basal tines of the syntype of Fig. 42 c, D, E. AFFINITIES. The systematic position of this species is quite uncertain. No doubt it does not belong to the genus Cervus in the restricted sense denned in the previous pages. The lower portion of the antlers recalls that of Megaceros verti- cornis, but the characters of the upper portion of the antlers and of the forehead rule out any relationship with Megaceros] they rather point to some affinity with Euctenoceros, but this question is still open. Specimens of Doubtful Attribution LOWER JAWS. Two lower jaws from Overstrand and Trimingham (Figs. 28 j, 46, 47) might belong either to " Cervus " obscurus or to Euctenoceros sedgwicki. They both belong to young but fully grown specimens. The teeth are massive and 74 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK FIG. 42. " Cervus " obscurus, two of the syntypes. J natural size. A & B. Mundesley, B.M. (G.D.), Savin 195. c, D & E. Forest Bed> B.M. (G.D.), M. 2321. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 75 FIG. 43. " Cervus " obscurus. Syntype, Overstrand. B.M. (G.D.), M. 6421. natural size. FIG. 44. A & B. " Cervus " cf. obscurus, Trimingham. B.M. (G.D.), M. 6400. c. " Cervus " obscurus, right antler of young specimen, Mundesley. B.M. (G.D.), M. 6307. D. " Cervus " obscurus, left antler of a young specimen, Mundesley. B.M. (G.D.), M. 6315, probably both belonging to the same individual. J natural size. 76 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK seem to be slightly more hypsodont than those from East Runton which I tentatively refer to Euctenoceros ctenoides. P 4 has a complete internal wall and the molars bear a strong anterior ridge. The ramus is depressed and thick, and its cross- section closely resembles that of the lower jaws of Euctenoceros from the Upper Valdarno (Azzaroli, 1948, fig. n, no. 3). The jaw from Trimingham, which is more complete, is distinguished by a large coronoidal process. The breadth of M 2 is 17 mm. FIG. 45. " Cervus " obscurus, tentative reconstruction, based on the syntypes. size. A. Anterior view. B. Dorsal view. natural BRAIN CASE. The identification of a brain case from Trimingham (B.M. (G.D.), M. 6303) is uncertain. The shortness of the pedicles gives evidence that it belonged to a fully grown specimen ; they are, however, too small for Euctenoceros ctenoides, E. sedgwicki or " Cervus " obscurus. Moreover, the forehead is hollowed, and bears no trace of the ridges which distinguish the latter species. Euctenoceros tetraceros is not recorded at Trimingham, and the supraoccipital crest of the specimen in question is much stronger than that of the skull roof from East Runton I tentatively identify with E. tetraceros. The brain case from Trimingham might possibly belong to Cervus cf. elaphus', however, it exceeds the size even of the largest specimens from Val di Chiana (Azzaroli, 1948). MEASUREMENTS : Occipital breadth . . . . . .156 Occipital height ...... 93 Minimal frontal breadth . . . . .152 Breadth of the brain case between the parietals . 104 Breadth of the condyles . . . , .89 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 77 FIG. 46. Cervid indet., lower dentition. Trimingham, Norwich Museum 45. Natural size. FIG. 47. Cervid indet., Trimingham, Norwich Museum 45. (See also Fig. 46.) natural size. 78 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK LIMB BONES. An account of the complete limb bones is given in the following table of measurements; some of them have been reproduced in Fig. 48. The metapodials are easily recognizable from those of Megaceros by the narrower form of their distal articulation, but a more precise determination is difficult and can be based only on the proportions. They might be attributed to Euctenoceros, to the red deer, or to " Cervus " obscurus. FIG. 48. Cervids indet., limb bones. J natural size. A. Left metatarsal, West Runton, Upper Freshwater Bed. B.M. (G.D.), Savin 1643. B. Left metacarpal, Trimingham. B.M. (G.D.), Savin 2168. c. Left metacarpal, Trimingham. B.M. (G.D.), M. 6473. D. Right metatarsal, Overstrand. B.M. (G.D.), Savin 773. E. Right metacarpal, East Runton. B.M. (G.D.), M. 6476. Some bones from Trimingham and from West Runton (B.M. (G.D.), Savin 2168, 1424, 1643) are long and slender like those of Libralces gallicus, but their shape is quite different and leaves little doubt that they belong to a cervine. Their identi- fication is puzzling; Euctenoceros ctenoides and E. tetraceros can be excluded, and an identity with the red deer seems extremely improbable, although only this DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 79 among the five species quoted above has been found in the Upper Freshwater Bed at West Runton. Euctenoceros sedgwicki was presumably as heavy an animal as the largest representatives of its genus. An identification with " Cervus " obscurus seems less improbable. MEASUREMENTS : Radius : Total length Proximal breadth Distal breadth . B.M. (G.D.), 1105. Trimingham 337 70 63 B.M. (G.D.),M. 6469. Sidestrand 325 Metacarpal : Total length Proximal breadth Distal breadth . B.M. (G.D.), Savin 2168. Trimingham 321 5i 50 B.M. (G.D.), Savin 1424. Trimingham 316 53 53 61 B.M. (G.D.), M. 6476. East Runton B.M. (G.D.), M. 6473. Trimingham Femur : Total length Proximal breadth ..... Antero-posterior diameter of the distal epiphysis . 267 283 . 306 291 45 50 55 52 45 50 57 53 B.M. (G.D.), Savin 1446. Trirningham 360 103 85 B.M. (G.D.), B.M. (G.D.), B.M. (G.D.), B.M. (G.D.), Metatarsal : Total length Proximal breadth Distal breadth Savin 1685. Trimingham Savin 1071. Overstrand Savin 773. Overstrand M. 6490. Sidestrand 310 307 311 265 42 4i 45 39 47 52 B.M. (G.D.), M. 6495. East Runton Total length . Proximal breadth Distal breadth 293 44 50 293 47 B.M. (G.D.), Savin 1643. West Runton, U. Freshw. Bed 347 46 54 Cervid cf . Dama nestii nestii (Fig. 49-) 1882. Cervus etueriarum ? Croizet & Jobert : Newton, p. 55. ? 1889. Cervus rectus Newton, p. 145, fig. i, la. ? 1891. Cervus rectus Newton : Newton, p. 30, pi. 4, fig. i. SYNONYMY. Newton's Cervus rectus is based on a very young specimen, probably of this species, as may be seen by comparison of the type with fig. i n i, 2 in Azzarpli, 1948. OCCURRENCE. Seven lower portions of antlers and a fragment of the upper portion, from East Runton; two basal fragments from Sidestrand and one from FIG. 49. Cervid cf. Dama nestii nestii, antlers. J natural size. A. Left antler, Mundes- ley. B.M. (G.D.), Savin 1164. B. Left antler, East Runton. B.M. (G.D.), M. 6365. c. Fragment of the upper portion of a left antler, East Runton. B.M. (G.D.), M. 6389. D & E. Left antler, East Runton. B.M. (G.D.), M. 6568. DEER OF WEYBOURN CRAG AND FOREST BED OF NO,RFOLK 81 Mundesley. Probably two isolated M 3 and some imperfect metapodials from East Runton. DESCRIPTION AND COMPARISONS. This species is poorly represented. The antlers correspond with those of Dama nestii nestii from the Upper Valdarno and Olivola (Azzaroli, 1948) for the position and development of the brow tine and the regular helicoidal torsion of the beam. Also the fragment of the upper portion agrees with the terminal fork of this subspecies ; the curvature of the beam and the characters of the fork give evidence that the bifurcation took place in a transverse plane. If so, the antler had four tines. The other species, with which this species can be compared are : Cervus philisi from Seneze (Schaub, 1942), Cervus rhenanus from Tegelen (Hooijer, 1947) and Cervus perolensis from the Auvergne (Bout & Azzaroli, 1953). Cervus philisi differs in the stronger development of the brow tine, the straighter form of the beam, and the characters of the upper portion of the antlers. Cervus rhenanus, whose antlers are more imperfectly known, differs mainly in the higher position of the brow tine, and Cervus perolensis, also imperfectly known, in the stronger development of the brow tine. These three species are rather primitive and their relationships are not yet clear. Cervid cf. Dama clactoniana Falc. (Fig. 50) I include here several specimens of the size of a fallow deer. Their identification is somewhat uncertain and they might possibly include more than one species. Their age is also uncertain, but at least the specimens from the Upper Freshwater Bed are certainly post-Villafranchian. ANTLERS (Fig. 50). Three basal fragments from Bacton, Trimingham, and West Runton (Upper Freshwater Bed). They are distinguished by a very stout basal bifurcation and a strong curvature of the brow tine. The pedicles are small. The antlers from Bacton and from West Runton are disproportionately large for their pedicles and probably belong to old individuals. TEETH. Several isolated teeth and a complete lower tooth row from the Upper Freshwater Bed at West Runton; two imperfect lower jaws, from Overstrand and Trimingham. The lower molars bear well-developed basal columns. The P 4 from Overstrand is primitive, two P 4 from West Runton are advanced. The total length of the lower tooth row from West Runton is 100 mm. LIMB BONES. Two perfect metacarpals, from Overstrand and Trimingham. Fragments of about the same size occur in the Upper Freshwater Bed of West Runton. MEASUREMENTS : B.M. (G.D.), B.M. (G.D.), M. 6475. Savin 1399. Metacarpal Overstrand Trimingham Total length . 232 . 235 Proximal breadth -35-35 Distal breadth . 35 . 35 GEOL. II, I. 6 82 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK COMPARISONS. In the Villafranchian fauna only Da-ma nestii eurygonos (Azzaroli, 1948) can be compared with the above specimens. However, the specimens from Bacton and West Runton are larger than this subspecies; moreover those from West Runton are certainly younger, and this is probably true also for those from Bacton and Trimingham. After the close of the Villafranchian, no deer of this size is known until the Clacton stage. FIG. 50. Cervid cf. Dama clactoniana, right antlers. J natural size. A. Abnormal specimen, Bacton, very old ? B.M. (G.D.), Savin 1283. B. Abnormal specimen, very old? West Runton, Upper Freshwater Bed. B.M. (G.D.), Savin 554. c. Trimingham, B.M. (G.D.), M. 6367. In this horizon we find in East Anglia Dama clactoniana (Bate, 1938, Oakley & Leakey, 1938), with which the fragments described above may be identified. Keilhack (1888) described a pair of antlers of a large fallow deer from Belzig, near Berlin, which he supposed to belong to the Lower Pleistocene or even to the preglacial. But the occurrence of Alces in the same deposit points to an Upper Pleistocene age. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 83 Species incorrectly recorded from the Forest Bed " Cervus " polignacus Robert 1882. Cervus polignacus Robert : Newton, p. 59. 1891. Cervus polignacus Robert : Gunn, pi. i, fig. 94, pi. 2, fig. 95. 1891. Cervus polignacus Robert : Newton, p. 29, pi. 4, fig. 12. Newton, on Falconer's authority, referred to Cervus polignacus two imperfect antlers characterized by a low brow tine, branching at a very wide angle from the beam. These two specimens have been figured by Gunn and are now in the Norwich Museum. Several more imperfect antlers in the same museum have been referred to this species, and in the British Museum I have seen other fragments of a similar type. One of them, of a rather small size, has been incorrectly identified with Megaceros savini by Dawkins (1887, pi. 3, fig. 2). All these fragments are very unsatisfactory and are susceptible of different interpretations. The name Cervus Dama polignacus was given by Robert (1829) to two incom- plete skeletons from Solilhac: one of them is identical with his Cervus solilhacus (now Megaceros solilhacus} from the same locality. The second is a small red deer with abnormal characters; Pomel (1853) named it Cervus roberti. " Cervus " carnutorum Laugel A fragment said to come from the Chillesford Clay at Aldeby, identified with Cervus carnutorum and figured by Dawkins (1872), may either belong to a large specimen of Euctenoceros falconeri, or to E. sedgwicki. Three specimens from the Norwich Crag quoted by Newton (1891, p. 26, pi. 4, fig. 3) possibly belong to E. falconeri. Other specimens from the Forest Bed determined as Cervus carnutorum in the collections belong to E. sedgwicki. The name carnutorum, like polignacus, should be dropped (Stehlin, 1912). Alces alces L. 1891. Alces ? Gunn, pi. 6, fig. 4. A much rolled antler of Alces alces, from an unknown locality, has been figured by Gunn. Its fossilization is less advanced than that of the other specimens, and no doubt it came from a younger horizon. Megaceros giganteus Blumenbach 1882. ? Cervus megaceros Hart : Newton, p. 58. Although this species was recorded from the Forest Bed by the earlier authors, Newton excluded it. An isolated P 4 from Sidestrand, in the Savin collection, registered B.M. (G.D.), Savin 1601, might possibly belong to this species, but its occurrence in the Forest Bed can be excluded. 84 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK Rangifer tarandus L. 1891. Cervus sp. Gunn, pi. 6, fig. 3. A much-rolled fragment of an antler, found on the beach at Pakefield, belongs to this species. Gunn questioned whether it belonged to the Forest Bed Fauna, and it can be excluded. CHRONOLOGY The Characters of the so-called Forest Bed Fauna The so-called Forest Bed Fauna is very abundant in species, and includes repre- sentatives of faunas met with at different horizons in continental Europe. This unusual assemblage has been explained in various ways. Whereas the older authors favoured an Upper Pliocene age (Reid, 1890, with bibliography), or assumed that this fauna was partially or totally derived (Dubois, 1905), more recent authors (Osborn, 1922, followed by Zeuner, 1945) attributed the whole fauna to the early Pleistocene (post-Villafranchian), and interpreted the archaic species as relics. Zeuner distinguished a " Cromer Forest Bed," where these archaic representatives are found, and a " Bacton Forest Bed " without archaic elements, which he thought to be a little younger. All these interpretations are untenable. A Pliocene age is ruled out by the presence of species which immigrated into Europe after the close of Villafranchian times. On the other hand, the older representatives, once attributed to the Pliocene but actually of Upper Villafranchian age, do not constitute isolated relics; an entire faunal assemblage characteristic of that period is present. Moreover, primitive species occur together with their more advanced descendants, and Zeuner's " Cromer Forest Bed " is richer in species than any other locality. The deer, according to the present revision, include 15 species, and we cannot imagine that they lived at the same time. A derived fauna can also be excluded, as the fossils consist for a large part of large antlers of deer, jaws and limb-bones of elephants, all of them heavy and brittle, but rather well preserved. This is true both for the older and for the younger representatives of the fauna. The alternative solution, namely, that the deposit includes several horizons, has been discussed in a previous section (" Geology "), and affords the only satis- factory explanation of the mixed character of the " Forest Bed Fauna." An attempt will be made here to determine the stratigraphical range of the 15 species of deer. It will be useful, however, first to illustrate the time scale we shall use for com- parison. Faunal changes during the Lower and Middle Pleistocene The Pleistocene continental sequence is marked by widespread faunal changes, consisting of extinctions of old forms, immigrations of new forms and evolution of primitive species into more advanced ones in situ. Following the resolution made by the International Geological Congress in London (1948), we shall place the Calabrian, and its continental equivalent, the Villafran- chian, at the base of the Pleistocene. However, it is now generally realized that DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 85 the so-called main Villaf ranchian fauna does not constitute such a uniform assemb- lage as former authors assumed, and that it ranges over a relatively long lapse of time. The well known faunas of the pumiceous sands and conglomerates of the Mon- tagne de Perrier in Auvergne (Ravin des Etouaires, Roca Neyra, Pardines) are older than any of the other faunas hitherto attributed to the Villafranchian, and their inclusion in this stage is misleading. These faunas are broadly contemporaneous with those of the Roussillon (Deperet, 1897) and of the first lacustrine phase of the Upper Valdarno (lignites with Tapirus arvernensis and laminated clays with plants ; see Merla, 1949 : 51-57). In the Montagne de Perrier two horizons can be distin- guished (Jung, 1946), but according to Prof. Bout of Le Puy (private information) there is no break in the sequence and the series seems to range over a short interval of time. In the three localities recorded above there is no trace of the cold climatic phase that marks the beginning of the Pleistocene. The fauna of the immediately following cold phase is perhaps represented in the solifluction deposits of Vialette, in the highlands of southern France (see Bout & Cailleux, 1951). In my opinion this cold phase might correspond to the Giinz glaciation and marks the beginning of the Pleistocene. In the Upper Valdarno basin the first lacustrine phase is overlain uncomformably by a second lacustrine series (Merla, 1949), and to this only is the term Villafranchian generally applied by Italian geologists. Its lower portion, not exposed, has been explored by borings and has yielded a flora that seems to indicate a cold climate. In the exposed section probably two horizons can be distinguished palaeontologi- cally, a lower horizon with Mastodon arvernensis and an upper horizon with Elephas meridionalis, although these two species may have lived together for a short period (Merla, 1949; Azzaroli, 1950). The fauna of Olivola (Azzaroli, 1950) is contemporary with the lower section with Mastodon, that of Tegelen (Hooijer, 1947) with the upper section with Elephas. The latter, according to Woldstedt (1950, " Tegelenschichten ") is of an imme- diately pre-Elster age, that is, it belongs to the Giinz-Mindel interglacial. The end of this period is probably represented by the faunas of some solifluction deposits of southern France (Bout & Cailleux, 1951 ; Bout & Azzaroli, 1953, and private information from Prof. Bout). Seneze (Schaub, 1944) is perhaps the most cele- brated of these deposits. Other localities are Sainzelles and the Creux de Peyrolles. Perhaps also the loessic deposit of St. Vallier (Viret, 1948) should be placed here. These faunas are decidedly younger than that of Vialette and may be correlated with the Mindel glaciation, that marks therefore the close of the Villafranchian. In England the Pleistocene starts with the Red Crag (Lagaaij, 1952). The mam- malian fauna of this horizon is represented by poor fragments, among the most interesting of which are Mastodon arvernensis, Elephas cf . planifrons, two species of Megaceros, and Euctenoceros falconeri. This horizon presumably corresponds to the unexposed section of the Villafranchian Valdarno series. The equivalent of the highest section of the same series, and of the clays of Tegelen, is represented in England by the Weybourn Crag of Norfolk (see also later). 86 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK It is no easy task to state the fate of all the species of the main Villafranchian fauna, as the relationships of many of them with the representatives of later faunas are not yet clear. Sus strozzii, Euctenoceros ctenoides, E. dicranios, E. tetraceros, Dama nestii, " Cervus " philisi and its allies, Meganthereon meganthereon, Brachy- prosopus vireti, Gazellospira torticornis, Nemorhoedus meneghinii, Procamptoceras brivatense, Nyctereutes megamastoides and Acinonyx pardinensis have left no des- cendants in the later faunas. This is possibly true also for Leptobos', a doubtful Leptobos, however, has been recorded from Siissenborn (Merla, 1949 : 49). The destiny of the horses is less clear, as this group still needs revision (Stehlin, 1932- 33:51 ff.). The same difficulty arises for many carnivores; Epimachairodus survived for some time, and the larger felids, canids and bears might perhaps have evolved into more modern species (Stehlin, 1932-33, Schaub, 1949). The voles, represented in the Villafranchian by the primitive Microtus, underwent a gradual evolution which sets them among the most useful leading fossils for the Lower and Middle Pleistocene (Hinton, 1926; Schreuder, 1936, 1943, 1950; Heller, 1933, 1939; Van der Vlerk & Florschutz, 1950). A similar evolution was carried out by the elephants, although things are here less simple than they were thought at first (Trevisan, 1953). Also Libralces seems to have evolved further (see section " Palaeontology "). Dicerorhinus etruscus has been recorded also from post- Villafranchian deposits, but is represented there by more advanced forms (Soergel, 1923). Hippopotamus is found in the Villafranchian and in later times; its history may be one of successive immigrations and retreats (Stehlin, 1932-33). Macacus and Castor also survived, but their remains do not indicate whether they underwent any great change. Trogontherium boisvilletti seems to have survived unchanged in France and the British Isles (Schreuder, 1951). The following phase is marked by the immigration of new faunal elements from Eastern Europe or from Asia: primitive red deer (Cervus acoronatus, C, benindei and other poorly known forms), the roe, wild boars of the group of Sus scrofa (pro- bably represented by a distinct variety; Soergel, 1923), Gulo gulo, Bos primigenius, Bison priscus. The voles evolved from Mimomys into primitive Arvicola (A. greeni, bactonensis, mosbachensis}. A primitive Microtus has also been recorded from Mosbach. Lib- ralces seems to have evolved from L. gallicus into L. reynoldsi, and the elephants from Elephas meridionalis into various races of E. antiquus and E. trogontherii. The group of Megaceros verticornis flourished in this period and possibly evolved in situ. Probably also the rhinoceroses and many carnivores are descendants of Villafranchian ancestors. The best representatives of this period are the main faunas of Mosbach and Siis- senborn, and the fauna of Mauer, in Germany; the fauna of Tiraspol in Bessarabia (Pavlow, 1906); the Needian of the Netherlands (Van der Vlerk & Florschutz, 1950), and some fossiliferous sands in the Upper Valdarno, around the village of Bucine (horizon 3 in Merla, 1949 : 51). These sands overlie comformably the Villa- franchian, with the interposition of a series of barren gravels. In England the equivalent of these deposits is represented by the estuarine section of the Forest Bed series. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 87 The dating adopted here is in contradiction with some current views and needs explanation. Soergel (1928) assumed an early Mindel, if not even a pre-Mindel age for the fauna of Mauer. This was deduced from the occurrence of five horizons of weathered loess above the fossiliferous sands a rather unsafe way of arguing, inasmuch as the age of the deposit depends on the number of glacial phases we admit. Soergel assumed two stages in the Wiirm (Weichsel) glaciation, two in the Riss (Saale) and two in the Mindel (Elster) ; but recently Woldstedt (1950) described four phases in the Weichsel glaciation and three in the Saale. These are sufficient to explain the occurrence of even more than five horizons of weathered loess above deposits of the great interglacial, and any evidence for attributing Mauer to the Mindel glaciation falls short. On the other hand, positive evidence for attributing these faunas to the Mindel-Riss interglacial is afforded by the Needian of the Netherlands, which corresponds to the Poludina-beds representing the great interglacial in North- ern Germany (Woldstedt, 1950). In 1932 Solomon placed the Forest Bed series of Norfolk in the first interglacial and assumed that the overlying glacial drift included representatives of the Mindel (North Sea Drift), Riss (Great Eastern) and Wurm (Little Eastern) glaciations. But some of the difficulties met with in identifying Solomon's North Sea Drift with the Mindel glaciation and with the Norwich Brickearth of southern Norfolk were stressed a year before by Boswell (1931), and were given again by Boswell and by Hazzledine Warren in the discussion following the presentation of Solomon's paper. Solomon's evidence of a Mindel-Riss horizon within the glacial drift, assumed to be afforded by his " mid-glacial sands," is not conclusive, as these sands are barren and form no continuous horizon. In the dating adopted here the second (Mindel) glaciation falls between the Weybourn Crag and the Estuarine Bed. Its deposits, represented in southern Norfolk by BoswelTs Norwich Brickearth, may have been cut out in north Norfolk by the erosion at the base of the Estuarine Bed. The distribution of the Bovidae seems to have been influenced by geographic or climatic factors. The bison is recorded at Mauer, Mosbach and Siissenborn, whereas the aurochs makes its appearance in Central Europe during the Riss glacia- tion (probably in a mild interstadial). On the other hand this species is not un- common in the sands near Bucine in the Upper Valdarno, with Cervus cf . ela-phus and Elephas antiquus, which, from their position, cannot be much younger than the Villafranchian, whereas the bison is recorded in Italy only from the Upper Pleisto- cene. Possibly Bos immigrated into Italy along the southern slope of the Alps, whereas for some unknown reason it avoided Central Europe, and Bison did the reverse. In the gravels of Tiraspol, with a faunal assemblage characteristic of the stage of Mauer and the upper sands of the Upper Valdarno, Bos and Bison occur together. Unfortunately Bos is represented only by limb bones (Pavlow, 1906). A later phase is marked by the appearance of more advanced species of Arvicola, evolved in situ, and by a widespread occurrence of Microtus, probably immigrants ; but there does not seem to be any well-marked break in the fauna. Dama clac- toniana, recorded only from the British Isles, belongs to this period. The incoming Riss (Saale) glaciation seems to have extinguished Dama clac- 88 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK toniana and many species of the group of Megaceros verticornis. Elephas primi- genius, or a related form, occurs for the first time at the beginning of this phase, but became common only after a mild oscillation, well marked at Steinheim a.d. Murr. In this mild interstadial the giant deer are represented here by Megaceros antecedens (Berckhemer, 1941) and the red deer by Cervus elaphus angulatus (Beninde, 1937). In the following cold phase the red deer is represented by a modern form and Coelodonta antiquitatis makes its very first appearance as an immigrant. The date of the immigration of the reindeer is less clear. Soergel (1943) recorded it from Steinheim, Mosbach and Siissenborn, but I have been unable to check the dating of these specimens. They are all said to belong to the tundra group (R. arcticus}. Also Ovibos moschatus has been recorded from Siissenborn (Soergel, 1941). At the end of the Riss glaciation (late Drenthian) the voles are represented by modern forms. A good guide fossil for the post-Rissian is Megaceros giganteus. The Distribution of the Deer of the Norfolk Coast and their value for Stratigraphy As stated above, the deer of the Forest Bed series sensu lato include representatives of different faunas. Two faunas are quite distinct, whereas the existence of a third fauna is vaguely indicated by the deer, and is demonstrated conclusively only by other mammals. The earliest fauna belongs to the upper section of the Villafranchian. It can be correlated with the horizon of Tegelen, Seneze, and the ferruginous sands of the Upper Valdarno, and is represented by Libralces gallicus, Euctenoceros tetraceros, E. ctenoides, cervid cf. Dama nestii nestii. This fauna occurs in the Weybourn Crag. The second fauna belongs to the second interglacial and corresponds stratigraphi- cally to the classic faunas of Mosbach (main fauna), Mauer, Siissenborn (main fauna) and Tiraspol. Its representatives are Libralces reynoldsi, Cervus cf. elaphus, Megaceros verticornis, Capreolus capreolus. This fauna occurs in the Estuarine Section of the Forest Bed. The third fauna, corresponding to a later horizon of the same interglacial, is probably indicated by Dama clactoniana, associated with Cervus cf . elaphus, Mega- ceros verticornis, Capreolus capreolus. This fauna occurs, partly at least, in the Upper Freshwater Bed. Possibly there is no true break between the two younger faunas. The distribution of these species is not uniform throughout the outcrop of the Forest Bed series s.l. The Villafranchian species are restricted to the western section of the outcrop; they are very common at East Runton, where they are not accompanied by later fauna! elements, and at Sidestrand, associated with younger species ; rare at West Runton, Overstrand and Mundesley. Some doubtful and much-rolled specimens have been found also at Pakefield, at the eastern extreme of the outcrop. The second fauna occurs in all the localities, with the exception of East Runton, where only a few much-rolled specimens have been found. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 89 The poorly represented fallow deer, supposed to represent the third fauna, is recorded from Bacton, Trimingham, and from the Upper Freshwater Bed at West Runton. Other representatives of this fauna have been recorded by Hinton from the Upper Freshwater Bed at Bacton and West Runton (see later), but this fauna might be more widespread. Many localities have therefore yielded a mixed fauna, whereas a few of them have yielded more uniform faunas. Of particular interest are East Runton, with a pure Villafranchian fauna, and the Upper Freshwater Bed of West Runton, which, also on geological grounds, can be assumed to include only representatives of the "third" fauna. This distribution may give a clue to an approximate dating of the remaining species, not known or doubtful in other districts. Libmlces minor is recorded only at East Runton and Sidestrand; it belongs therefore to the Villafranchian fauna. Libralces latifrons from Happisburgh (doubtful at Mundesley, Cromer and Walcot) may belong either to the second or the third fauna. Euctenoceros sedgwicki, from Bacton and Mundesley, probably belongs to the second fauna. The presence of the large fallow deer and the absence of Libralces reynoldsi at Bacton would rather suggest a later age, but the absence of E. sedg- wicki in the very fossiliferous Upper Freshwater Bed of West Runton is, perhaps, more significant. This species is possibly present also at St. Prest. Megaceros dawkinsi and M. savini are probably of the same age. They do not occur in the Upper Freshwater Bed of West Runton, nor at East Runton. M. dawkinsi has been recorded also from the Weybourn Beds at Weybourn by Savin, but the name of the horizon was possibly taken from the locality and it would be unsafe to rely on this statement. This species, or a related form, seems to have been widespread in continental Europe. The dating of " Cervus " obscurus is more doubtful, but this species probably belongs to the second interglacial. The Evidence Afforded by Other Mammals on the Age of the Forest Bed Series The list of the other mammals from the Forest Bed s.l. also gives the impression of a mixed fauna. We need only mention the occurrence of four species of elephants, a fact not recorded from any other locality in Europe. The elephants still need revision, and it would be unsafe to assume the alleged Elephas primigenius as evidence of the existence of an early Rissian horizon in the Forest Bed series. But the evidence afforded by the voles (Hinton, 1926; 19260.) is more conclusive, and demonstrates the existence of an horizon equivalent to the Swanscombe gravels. The " shelly crag " at East Runton yielded Villafranchian representatives (Mimomys pliocaenicus, M. intermedius, M. savini} , whereas the Upper Freshwater Bed at West Runton yielded a mixed fauna: Mimomys intermedius, M. savini, M. maiori, Evotomys sp., Pitimys arvaloides, P. gregaloides, Microtus arvalinus, M. nivalinus, M. nivaloides, M. ratticeppoides. This is certainly not evidence of 90 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK the presence of different horizons in the Upper Freshwater Bed, as the older species may easily have been washed in. The age of the deposit is indicated by the younger species and corresponds to the early Drenthian (Van der Vlerk & Florschiitz, 1950). The same species of Microtus have been found also in the middle gravels of Swans- combe (Schreuder, 1950). A correlation between the Upper Freshwater Bed and Swanscombe has been made also by Hinton (19260, 336-337). Shortly later Mochi (1929 : 179-181) proposed a correlation of the Norfolk sequence with the earliest glaciations that agrees substantially with that stressed in these pages and differs only in minor details. SUMMARY GEOLOGY AND CHRONOLOGY. The mammal bearing horizons of the Norfolk coast include three horizons separated by unconformities: the Weybourn Crag, the Estuarine Bed or Forest Bed s. sir. and the Upper Freshwater Bed. The whole series rests on a levelled surface of chalk and is capped by glacial drift. The so- called Forest Bed Fauna is actually made of three successive faunas. The oldest one, from the Weybourn Crag, is of upper Villafranchian age and is assumed to correspond to the first (Gunz-Mindel) interglacial. Its representatives are: Euc- tenoceros ctenoides, E. tetraceros, Libralces gallicus, L. minor, and a small species perhaps identical with Dama nestii nestii. The second fauna, from the Estuarine Bed, is contemporary with the main faunas of Mosbach, Siissenborn, Mauer, Tiraspol, and the horizon of Neede, and is assumed to correspond to the second interglacial. Its representatives are: Megaceros verticornis, M. dawkinsi, M. savini, Libralces reynoldsi, Cervus cf. elaphus, Capreolus capreolus, and possibly Libralces latifrons, Euctenoceros sedgwicki and " Cervus " obscurus. The third fauna, from the Upper Freshwater Bed and possibly also from the highest section of the Estuarine Bed, is contemporary with the gravels at Swanscombe and with the early Drenthian and corresponds to a later period of the same interglacial. It is represented by a species possibly identical with Dama clactoniana, together with Megaceros verticornis, Cervus cf. elaphus and Capreolus capreolus. Conclusive evidence of its age is afforded by small rodents. PALAEONTOLOGY. Alcinae and Capreolinae were differentiated from the Cervinae before the Upper Miocene. The Upper Miocene Cervinae belong to two quite dis- tinct lineages. In eastern Europe they are represented by Damacerus bessarabiae (= Cervocerus novorossiae} and D. variabilis (= Procervus variabilis), and may per- haps have evolved into the Villafranchian Cervus ramosus. In China they are represented by three-tined deer (incorrectly identified with Cervocerus novorossiae by former authors), from which probably most of the Pleistocene and living Cervinae took origin. The phyletic development of antlers is also discussed. The deer of the Weybourn Crag and Forest Bed s.l. belong to the genera Libralces, Capreolus, Cervus, Euctenoceros, Megaceros and possibly Dama; there is in addition a species of Cervinae of uncertain affinities. Libralces is a close ally of Cervalces. L. gallicus is smaller than living elks, but L. reynoldsi, which is probably its descendant, is the largest deer hitherto known. L. latifrons and L. minor are poorly represented. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 91 The roe also is poorly represented. The earliest representatives of Cervus s. str. were more primitive than the living European red deer; some of them were similar to living red deer of Central Asia. The red deer of the Forest Bed also seem to be primitive, but their remains are not satisfactory. Euctenoceros is distinguished by peculiar features of the antlers. Its most primi- tive representative is E. falconeri from the Red Crag. E. sedgwicki is possibly its descendant. E. ctenoides (= E. teguliensis) and E. tetraceros do not differ from the types from other Villafranchian localities. Megaceros can be divided into two groups, based on characters of the skull and antlers: the group of M. giganteus and the group of M. verticomis. M. savini is a primitive species of the group of M. giganteus and its antlers are not palmated. M. verticomis, the commonest species of deer in the Forest Bed, has large and widely palmated antlers. M. dawkinsi belongs to the same group; it is smaller and has secondarily reduced antlers, and a hypsodont dentition. " Cervus " obscurus is a large species of quite unusual features and of unknown affinities. Its remains are scanty. Two much smaller species may possibly be identified with Dama nestii nestii and with Dama clactoniana. REFERENCES ALEXEJEW, A. 1913. Nouvelle espece de cerfs fossiles des environs du village Petrovierovka. Zap. novoross. Obshch. Est. (M6m. Soc. Nat. Nouv. Russie) Odessa, 40 : 1-13, pis. i, 2. (Russian with French translation.) 1915. Animaux fossiles du village Novo Elisavetovka. xiv + 453 pp., 10 pis. Odessa. ARAMBOURG, C. 1938. Mammiferes fossiles du Maroc. Mdm. Soc. Sci. nat. Maroc, Rabat, 46 : 1-74, pis. 1-9. AZZAROLI, A. 1948. Revisione della fauna dei terreni fluvio-lacustri del Valdarno Superiore, 3. I cervi fossili della Toscana con particolare riguardo alle specie Villafranchiane. Palaeontogr. Ital., Pisa, 43 : 45-82, pis. 7-19. 1950. Osservazioni sulla formazione villafranchiana di Olivola in Val di Magra. Mem. Soc. tosc. Sci. nat., Pisa, (A) 57 : 104-111. 1952. L'Alce di Seneze. Palaeontogr. Ital., Pisa, 47 : 133-141, pis. 15, 16. I952a. The Geological Age of the Cromer Forest Bed. Proc. prehist. Soc. E. Angl., London, (N.S.) 17 (1951) : 168-170. BACHOFEN ECHT, A. 1941. Die Geweihe von Cervus elaphus aus Hundsheim a. d. Donau, nebst Bemerkungen uber Geweihbildung. Palaeobiologica, Wien, 7 : 249-260, 18 figs. BATE, D. M. A. 1938. Notes on recent finds of Dama clactoniana (Cervus browni auctt.) in London and Swanscombe. Proc. prehist. Soc. E. Angl., London, (N.S.) 3 (1937) : 460- 463. ng. BELGRAND, E. 1869. Histoire generale de Paris, La Seine, I. Le bassin Parisien aux ages antehistoriques. 56 pp., 80 pis. Paris. BENINDE, J. 1937. Uber die Edelhirschformen von Mosbach, Mauer und Steinheim. a. d. Murr. Palaont. Z., Berlin, 19 : 79-116, pis. 6-8. 1940. Die Krone des Rothirsches. Z. 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Geol. Soc. Lond. 24 : 511-516, pis. 17, 18. 1872. On the Cervidae of the Forest-bed of Norfolk and Suffolk. Ibid., 28 : 405-410, 3 figs. 1878. Contributions to the history of the Deer of the European Miocene and Pliocene Strata. Ibid. 34 : 402-420, 17 figs. 1887. The British Pleistocene Mammalia, 6. British Pleistocene Cervidae. 29 pp., 7 pis. Palaeontogr. Soc., London. DE ALESSANDRI, G. 1903. Sopra alcuni avanzi di Cervidi Pliocenici del Piemonte. A tti R. Accad. Sci. Torino, 38 : 573-586, pi. i. DEPERET, C. 1890-97. Les animaux pliocenes du Roussillon. Mem. Soc. geol. Fr., Paleont., Paris, 1-7 : 1-194, P^ s - I-I9- iSgja. fitude de quelques gisements nouveaux de Vertebres pleistocenes de 1'Ile de Corse. Ann. Soc. linn. Lyon, 44 : 111-128, pi. i. DIETRICH, W. 1909. Neue Riesenhirschreste aus dem schwabischen Diluvium. Jh. Ver. vaterl. Naturk. Wurttemb., Stuttgart, 65 : 132-161, pis. 3-5. DUBOIS, E. 1905. L'age des differentes assises englobees dans la serie du " Forest-Bed " ou Cromerien (i). Bull. Soc. beige Geol. Pal. Hydr., Bruxelles (proc. verb.), 19 : 263-278. FABIANI, R. 1919. I Mammiferi Quaternari della regione Veneta. Mem. 1st. geol. Univ. Padova, 5, 2 : 1-174, pl - I ~3- FALCONER, H. 1868. Palaeontological Memoirs and Notes, 2. Edited by C. Murchison. xiii + 675 pp., 38 pis. London. FLEROW, C. 1930. The white muzzle deer (Cervus albirostris Przew.) as the representative of a new genus Przewalskium. C. R. Acad. Sci. U.R.S.S., Leningrad, 1930 : 115-120, fig. FRENTZEN, K., & SPEYER, C. 1928. Riesenhirsche aus dem Diluvium des Oberrheingebietes. Mitt. bad. geol. Landesanst., Freiburg, 10 : 175-232, pis. 13-17. FREUDENBERG, W. 1914. Die Saugetiere des alteren Quartars von Mitteleuropa mit beson- derer Beriicksichtigung der Fauna von Hundsheim und Deutschaltenburg in Nieder- osterreich nebst Bemerkungen uber verwandte Formen anderer Fundorte. Palaont. 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Das norwegische und das schottische Rotwild (Cervus elaphus L.). Ein kranio- metrischer Vergleich. Nyt. Mag. Naturv., Oslo, 65 : 175-219, 4 figs. JOHNSON, R. 1874. Notice of a new species of Deer from the Norfolk Forest-Bed. Ann. Mag. Nat. Hist., London, (4) 13 : 1-4. JUNG, J. 1946. Geologic de 1'Auvergne et de ses confins bourbonnais et limousins. Mem. Carte geol. det. Fr., Paris, ix + 372 pp. KEILHACK, K. 1888. Ueber einen Damhirsch aus dem deutschen Diluvium. Jb. preuss. geol. Landesanst., Berlin, 8 : 283-290, pi. n. KHOMENKO, J. 1914. La faune meotique du village de Taraklia du district Bendery. Annu. geol. min. Russ., N. Alexandria, 15 : 107-143, pis. 1-4. (Russian with French summary.) KIRCHNER, H. 1936. Mainfrankische Riesenhirschreste nebst Untersuchungen iiber Rassen- unterschiede, Geweihform, Bedeutung der Gefassrillen bei Riesenhirschen . Paldont. Z., Berlin, 18 : 227-256, pis. 15-17. KLEINSCHMIDT, O. 1938. Der fossile Hirsch von Mauer. Falco, Leipzig, 3* : 13-14. 1938^. 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Die oberpliocaene Saugetierfauna von Seneze (Haute-Loire) und ihre verbreit- ungsgeschichtliche Stellung. Ibid., 36 : 270-289, 8 figs. 1950- Revision de quelques Carnassiers villafranchiens du Niveau des Etouaires (Mon- tagne de Perrier, Puy-de-D6me). Ibid., 42 : 492-506, 8 figs. SCHMIDTGEN, O. 1938. Abnorme Geweihe aus dem Mosbacher Sand. Palaeobiologica, Wien, 6 : 335-339, pis. 21, 22. SCHREUDER, A. 1936. Fossil Voles and a Lemmus out of well-borings in the Netherlands. Verh. Akad. Wet. Amst. (2) 35, 3 : 1-24, n figs. 1943- Fossil Voles and other Mammals (Desmana, Talpa, Equus, etc.) out of well-borings in the Netherlands. Verh. geol.-mijnb. Genoot. Ned. Kolon., 's Gravenhage (Geol.) 13 : 399-434, 7 figs. DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 95 SCHREUDER, A. 1950. Microtinae from the Middle Gravels of Swanscom.be. Ann. Mag. Nat. Hist., London, (12) 3 : 629-635. 1951- The three species of Trogontherium, with a Remark on Anchitheriomys. Arch. neerl. 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Kiew. 1935- Riesenhirsche (Cervus giganteus Blum.) im Tschernigiwer Polessje. Die Quartar- periode, 9 : 107-117. Acad. Sci. Ukraine, Kiew. ZDANSKY, O. 1925. Fossile Hirsche Chinas. Palaeont. sinica, Peking, (C) 3 : 1-190, pis. 1-16. 1927. Weitere Bemerkung uber fossile Cerviden aus China. Ibid. (C) 5, i : 1-19, pi. I. ZEUNER, F. E. 1945. The Pleistocene Period, x + 322 pp. Ray Society, London. PRESENTED 7 JUL 1953 PRINTED IN GREAT BRITAIN BY ADLARD AND SON, LIMITED BARTHOLOMEW PRESS, DORKING 2 6 NOV 1953 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA M. G. CALDER BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY Vol. 2 No. 2 LONDON : 1953 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY The following papers appeared in Volume I (1949-52) : Price No. i (1949). The Pterobranch Rhabdopleura in the English Eocene. H. D. Thomas & A. G. Davis . . . . . .75. 6d. No. 2 (1949)- A Reconsideration of the Galley Hill Skeleton. K. P. Oakley & M. F. Ashley Montagu ...... 55. No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone of the Welsh Borders. E. I. White. Pteraspis leathensis White a Dittonian Zone-Fossil. E. I. White . 75. 6d. No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan, Northern Iraq. L. F. Spath . . . . . . .105. No - 5 (i95i)- Cretaceous and Eocene Peduncles of the Cirripede Euscal- pellum. T. H. Withers ........ 55. No. 6 (1951)- Some Jurassic and Cretaceous Crabs (Prosoponidae). T. H. Withers 5 5. No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian of Podolia. W. N. Croft ios. No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle East. T. F. Grimsdale . . . . . . . .ios. No. 9 (1952). Australian Arthrodires. E. I. White .... 155. No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard 6s. A CONIFEROUS PETRIFIED FOREST IN PATAGONIA BY MARY GORDON CALDER (University of Manchester) Pp. 97-138 ; Pis. 1-7 ; 7 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY Vol. 2 No. 2 LONDON : 1953 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical Series. Parts appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 2, No. 2 of the Geological series. PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued November 1953 Price Twelve Shillings A CONIFEROUS PETRIFIED FOREST IN PATAGONIA By MARY G. CALDER SYNOPSIS The present paper contains the results of a detailed investigation of a large collection of silicified coniferous remains from the remarkable petrified forest of Cerro Cuadrado, Patagonia. Conclusive evidence of only two species (Araucaria mirabilis and Pararaucaria patagonica), based on seed cones, has been found. Araucaria mirabilis belongs to section Bunya of the genus Araucaria, and the affinities of Pararaucaria patagonica are probably with the Taxodiaceae : both species are extinct. The associated wood, branches, twigs and seedlings probably belonged to one or other of the species described. The age of the forest is uncertain, and is at present placed somewhere between Middle Jurassic and Wealden. INTRODUCTION THE existence of petrified coniferous plant remains in regions west of the mouth of the Rio Deseado in the province of Santa Cruz, Patagonia, appears to have been first noted scientifically by Dr. A. Windhausen of the Argentine Geological Survey. In 1919 he discovered silicified cones which were kept as curios in stores and farms in the country south-west of the Gulf of San Jorge ; and in 1923 he found abundant petrified material of conifers in situ in the region of the Cerro Alto (or Cerro Chato), near the Estancia Belgrano, in a locality about S. lat. 47 50' by W. long. 68 40' (Windhausen, 1924 : 203, footnote). His collection of silicified wood and cones was passed on to Professor Gothan of Berlin, who gave a short description of them (Gothan, 1925). Dr. Carlos Spegazzini had also (1924) briefly described silicified twigs and cones, the latter similar to those found by Windhausen, which he had received from various donors : these specimens were from the region of the Cerro Madre y Higa, a small volcanic peak about 30 miles north-east of Windhausen's locality (Spegazzini, 1924). Meantime, during 1923-24, a palaeontological expedi- tion from the Field Museum of Natural History, Chicago, had been collecting fossil mammals in Eastern Patagonia, and Dr. Elmer S. Riggs and others made an addi- tional expedition to the region of the Cerro Madre y Higa, and its neighbouring peak about 10 miles to the west, the Cerro Cuadrado, where they found " a considerable number of fossil trees, some with stumps standing, others lying prone with broken branches and cones scattered about them, revealing a forest of fossil Araucaria or Brazilian Pines preserved on the site where it had grown " (Riggs, 1926 : 544 ; Wieland, 1935 : 6). These discoveries made it clear that in the region of the volcanic peaks of the Cerro Cuadrado, Cerro Madre y Higa, and Cerro Alto, in a district about 10 south of the modern forest of Araucaria araucana (Molina) (= A. imbricata Pav.) in Chile and Argentina, and about 26 south of the forests of Araucaria angustifolia (Bertol.) ( = A. brasiliana Rich.) in Southern Brazil, there existed the petrified GEOL. II, 2 7 too A CONIFEROUS PETRIFIED FOREST IN PATAGONIA remains of an extensive araucarian forest. The large Field Museum collection made by Riggs and his co-workers was later reviewed by Wieland (1929, 1935) ; his vividly-presented memoir, however, was without detailed anatomical studies of critical features. A fuller account of one of the cones in the Field Museum, with special reference to the embryo, was given by Darrow (1936). After this, little further botanical investigation of the petrified forest was undertaken, subse- quent accounts by Darrah (1939 : 222), Florin (1940 : 36), Arnold (1947 : 313) and Andrews (1947 : 175) being recapitulations of earlier studies, though Florin's later work (1944 : 513) gave a partly revised interpretation of the structure of one type of cone. A non-botanical account of the petrified forest is presented in the memoir by Wehrfeld (1935) of a journey through Santa Cruz. The geology of the regions between the Rio Deseado and the Rio Chico where the forest is situated has also been recently reviewed by Feruglio (1949 : 118 ; 1951 : 35), with special reference to fossil floras of plant compressions found at other localities in this area. Important new collections of the conifers of the petrified forest have, however, been made by Dr. Franz Mansfeld in the region of the Cerro Alto whilst searching for fossil vertebrates, and in 1936 the British Museum (Natural History) acquired an extensive collection of some 600 specimens from him, some by presentation and some by purchase. Similar material is also widely distributed in other museum and departmental collections : a demonstration of some of it was given by W. T. Gordon at the Geological Society of London on behalf of Dr. Mansfeld (Gordon, 1936). The collection in the Geology Department of the British Museum is a repre- sentative one ; in it are included silicified woods, branches and twigs of various sizes, seedlings, and cones, in varying states of preservation and offering remarkable evidence on the nature of the conifers of the petrified forest. The present account is based mainly on this material, though reference has also been made to specimens in other collections : Professor Gothan has described some of this more recently- collected material from specimens in Berlin and elsewhere (Gothan, 1950) and reference is made to this description when the seedlings are discussed. A more detailed study of these South American petrified conifers, and in particular, of the cones, appears to be desirable, especially in view of recent developments in studies of the taxonomy and morphology of the living Araucariaceae (White, 1947 ; Wilde & Eames, 1948, 1952 ; Buchholz, 1949) and of the fossil Araucariaceae (Kendall, 1948, 1949; Cookson & Duigan, 1951). In addition, the remarkable extension and organization of knowledge of the taxonomy, morphology and distri- bution of fossil conifers in general provided by the work of Florin (1931, 1940, 1938-45) give a more critical basis for comparative interpretation than was pre- viously available. LOCALITY AND AGE The site of the petrified forest is in the neighbourhood of the small volcanic peaks of the Cerro Alto (Chato), Cerro Cuadrado, and Cerro Madre y Higa, to the south of the Rio Deseado in the region between lat. 47-48 S., long. 68-69 W. (cf. Gothan, 1925, pi. 10 ; Wieland, 1935 : 6, text-fig, i). More recent accounts of the area where the silicified plant remains are found have, however, shown that the A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 101 extent of the forest is considerably wider than suggested in earlier reports. Feruglio's survey (1949 : 128, map opposite p. 134 ; 1951 : 69, 74) indicates a number of additional localities where cones and branches occur, some 25-30 miles to the north- west of the Cerro Alto : these are, Estancia Los Toldos, Canadon de las Cuevas, and west of Cerro Botellon. According to Feruglio's maps, there are at least two localities in the region south of the Rio Deseado called " Cerro Cuadrado " : one lies between the Cerro Alto and the Cerro Madre y Higa in the area originally indi- cated for the petrified forest, and another some 70 miles to the south-west near to the Estancia La Guitarra. From the latter locality Feruglio has described a quite separate mixed flora consisting of compressions including some araucarian remains (1951 : 62-65, 69), to which reference will later be made. Feruglio's review of the region of the Cerro Alto where the petrified araucarian remains are found (1949 : 128) does not include the vicinity of the Cerro Cuadrado, after which Wieland named the forest, as one of the fossiliferous localities ; the Cerro Alto appears to be the central locality of the forest, and also to be one which has produced very valuable and representative collections, though no doubt much remains to be discovered in other areas.. It is clear, from the various descriptions, that the forest was overwhelmed in an outburst of volcanic activity, the plant remains being embedded in a rhyolitic volcanic ash. Some are found lying weathered out on the surface, showing varying degrees of erosion of surface features, and some of the tree stumps are still standing as noted, for example, by Riggs (1926 : 544), and illustrated by Frenguelli (1941, pi. 25, fig. i) ; Frenguelli's photograph shows widely scattered trunks, and not a close-set forest formation. Numerous seedling stems, to which reference will later be made, also tend to confirm the impression that the forest was preserved in situ, and was overwhelmed whilst regeneration was in progress. The exact geological age of the forest is still doubtful. Dr. Mansfeld suggested that the rocks from which his specimens were collected were not older than Cretace- ous, and that they might even be of Tertiary age ; these estimates appear to have been based on the comparatively modern appearance of the plant structures, rather than on geological data (Gordon, 1936). The horizon has been placed, in other accounts of the petrified forest, as widely apart as mid-Triassic (Windhausen, 1924 : 203 ; 1931 : 201 ; Gothan, 1925 : 197 ; Wieland, 1929 : 60 ; 1935 : 8), Cretaceous (Florin, 1940 : 36 ; 1944 : 513) and Eocene (Frenguelli, 1933, cf. Barrow, 1936 : 328, and Fossa-Mancini, 1941 : 68). Dr. A. F. Leanza, of the University Museum of La Plata, Argentina, very kindly informed me (in lilt,, igih June, 1948) that geologically it could then only be indicated that the forest belonged to some stage in the Mesozoic. Windhausen dated the forest as Triassic in the belief that the volcanic ashes which enclose the plant remains form part of an eruptive which, in the geological correlations then in vogue, was comparable in age with the eruptive cycle which in the province of Mendoza originated the porphyritic series found beneath sediments containing a Thinnfeldia flora ; eruptive cycles are, however, now known also to have occurred in Patagonia at later stages in the Mesozoic. Ferug- lio has reviewed again, in some recent publications (1949 : 118 ; 1951 : 35) to which Mr. W. N. Edwards very kindly drew my attention, the problem of the geologic 102 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA age of the forest. He describes a series of floras of plant remains in the form of compressions from a number of outcrops of the complex of volcanic sediments (the " porphyritic series of Bahia Laura ") in the region between the Rio Deseado and the Rio Chico ; and compares them in detail with floras of mid- Jurassic to Wealden age, with special reference to species of Hausmannia, Cladophlebis, Nils- sonia, and Ptilophyllum. He concludes that the age of the volcanic series is in large part between mid- Jurassic and Wealden ; and that the age of the petrified forest around the Cerro Alto, which occurs in the same complex (although its exact stratigraphic position in relation to the other floras has not been determined), must lie somewhere within the same range (1949 : 131 ; 1951 : 74). Feruglio also points out that a major unconformity separates this volcanic series from the Upper Cretaceous ; and also that the original determination of a species of Estheria from this series in this area as being of Rhaetic age is not now generally accepted. These latter points further support his delimitation of the age on the basis of the floras. PRESERVATION AND TECHNIQUE The plant remains are silicified : the replacement of the organic matter has been fairly complete, and the specimens have not been effectively sectioned by any adaptation of the cellulose peel technique. Thin petrological slices have accordingly been used (prepared by Mr. J. Fowler of Sheffield) where details of the tissues are required. However, in some specimens or parts of specimens the silicification has gone so far that little detail can be distinguished : in others the wall structure is well denned, and such details as the nature of pitting in the xylem, and the structure of the embryos in some of the cones, may be observed. The cut and polished faces of the cones may be used quite effectively for general morphology in many cases ; Wieland relied very largely on this means of examination in his description of the types (Wieland, 1935), which was accordingly lacking in histological details that have proved to be of considerable importance to the interpretation of the general morphology and taxonomy. Cut and polished specimens form very pretty curios, as the preservation is in rather light silica enhanced by a variety of chalce- donic shades of colour. Darrow (1936 : 330) has already commented on the ten- dency for the various tissues of the seeds to be preserved in distinctive shades of quartz. (i) FOLIATED AND DEFOLIATED TWIGS AND BRANCHES Araucarites sanctaecrucis n. sp. (PL i, figs. 4-6, 8-13) 1924. Araucarites ? Spegazzini, p. 133, text-fig. 4 (8). 1935. Proaraucaria mirabilis (Speg.) : Wieland, p. 27, pi. 13, fig. i. DIAGNOSIS. Woody branches of araucarian habit, radially symmetrical, 5-25 mm. in diameter, occasionally with lateral axillary branches in one row or two oppo- site rows. Evergreen : leaves or leaf-scars arranged in a spiral with angle of diver- gence 3/8 or 5/13, the latter on the thicker branches. Leaves imbricate and more or less appressed. Leaf rhomboidal, 8-14 mm. in length, 4-8 mm. in width at its A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 103 broadest region, tapering distally to a subacute, somewhat incurved apex, the mar- gins converging at the apex at an angle of 6o-8o. Abaxial surface sometimes showing parallel longitudinal ridges and grooves. Free part of leaf flat, about half the length of the entire leaf, and seated on a flat leaf-base cushion, from which on older branches it has been shed. Leaf-base cushions on older branches 20-30 mm. wide, 5-10 mm. high, with marked longitudinal fissures. Venation of leaves and structure of epidermis not known. Secondary xylem, seldom preserved, with uniseriate and contiguous bordered pits on radial walls of tracheids. LECTOTYPE. The specimen figured by Spegazzini, 1924, fig. 4 (8). HYPOTYPE. V. 30941. Geol. Dept., B.M. (N.H.). The specimen illustrated in this paper (PI. I, figs. 8, 10, n). LOCALITY AND HORIZON. Cerro Alto, Santa Cruz, Patagonia : Between mid- Jurassic and Wealden. DESCRIPTION. Many of the detached branches show one side of the specimen much weathered, with surface features nearly obliterated, and the other with excellent preservation of external morphology (PI. i, figs. 8, 10, n). No doubt the better preserved surface was that which lay downwards on the forest floor, whilst the upper surface was subject to processes of weathering as the surrounding rhyo- litic volcanic ash was worn away ; some specimens of leafy branches are still em- bedded in the ash (PI. i, fig. i at L), but most are detached. PI. i, figs. 4, 9, 10, n and 13 illustrate the habit of branches of varying age. PL i, fig. n shows an un- branched stem probably four years old, while figs. 4 and 9 represent thinner branches, probably penultimate branches of the shoot, and show axillary branches similar to that (? i year old) in fig. 13, which tend to be arranged in two opposite rows or one row on the parent branch. This condition is found in the penultimate branch sys- tems of many conifers where dorsiventrality in respect of the ultimate branching is common. The defoliated branch of lower order (PI. i, fig. 6) may represent the broken apical region of a terminal main branch, with 3 laterals in an apparent whorl. In the foliated shoots the leaves are arranged in spiral phyllotaxy with angle of divergence 3/8 or 5/13, the latter on the thicker branches : Church (1904 : 99) has noted such spirals of a higher order on the thicker leafy branches in the Recent Araucaria excelsa. PI. i, fig. 10, t, shows the free lamina of the leaf, and at b the leaf-base cushion, from the stem shown in PI. i, fig. n. This specimen demon- strates that these plants were evergreen : in transverse section as seen in the cut surface at the upper end (PI. i, fig. 8), it shows poorly preserved secondary xylem with 3-4 rather ill-defined growth rings, probably annual. It also shows in some patches of the poorly preserved secondary wood that the radial pitting of the tracheids is uniseriate and contiguous as illustrated in PI. i, figs. 3, 7. The trans- versely cut face (PI. i, fig. 8, t) also shows the free part of the leaf in sectional view, demonstrating its flat, unkeeled nature. No veins are preserved in the leaf, so that the broad-based leaf habit cannot be definitely associated with parallel venation : nor can the longitudinal striation which is seen on the abaxial surface of the leaf in some specimens (e.g., PI. i, fig. 5) be definitely associated with any anatomical feature such as ribs of sclerenchyma. The furrows in such cases may represent the position of lines of stomata, but it has not been possible to demonstrate these structur- 104 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA ally, either from sections or cuticles. The older defoliated branch seen in PI. i, fig. 12, probably about 5 years old, shows tangentially-extended leaf-base cushions without any clearly-defined scar of attachment of the free lamina of the leaf : it is doubtful whether the longitudinal fissuring in such specimens is entirely a natural condition following on the stretching of the bark with increase in secondary thicken- ing, but it is a common feature in some living araucarians (cf . A . araucana, Seward & Ford, 1906, pi. 23, fig. D). In none of the specimens figured or described in previous accounts of the forest is there any indication of foliated branches : those referred to by Spegazzini (1924) and by Wieland (1935) as Araucarites ? and Proaraucaria mirabilis respectively are similar to that figured in PI. I, fig. 12 of the present account, which has been interpreted as an older branch where the free lamina of the leaf has been shed. After comparison of the variation in morphology of branches of different age in, for example, the living Araucana araucana (cf. Seward & Ford, 1906, pi. 23, A-G), there would seem to be no reason for regarding these thicker defoliated branches as a separate species in the absence of any further criteria of difference : the structure of the secondary xylem is too infrequently and inadequately preserved to be used as such. Various references to similar branches (Darrow, 1936 : 333, 337 where they are identified with Proaraucaria mirabilis, and Arnold, 1947 : 314) have compared them with the living South American species of Araucana, A. araucana and A. angustifolia, which comprise the section Columbea Endlicher, emend. Wilde & Eames. However, from the locality Meseta de Baquero, about 70 miles to the south-west of the Cerro Alto petrified forest, Feruglio has recently described com- pressions of leafy branches as a new species of Araucana, A. grandifolia (Feruglio, 1951 : 62, pi. 3, fig. 5), and he suggests that these might belong to the cone species Proaraucaria mirabilis, from the petrified forest, whose vegetative organs, he says, are not known (1949 : 137). He adduces no botanical reasons for this suggestion of relationship, and it is presumably based on geographical propinquity. Comparison of the habit of A . grandifolia with that of the leafy branches here described certainly does not suggest specific identity between the sterile branches from the two localities, for A . grandifolia has much larger leaves, long, rather narrow, pointed and lanceolate, and of spreading habit. There are no cuticular studies of this species, and on the evidence of the external morphology alone one might even hesitate to assign the branches to the genus Araucaria. Although the general habit of the foliated shoots from the petrified forest suggests comparison with an araucarian type, they cannot be identified on the basis of their external morphology with any living species of Araucaria. It is true that the flat and rather broad laminae of the leaves more closely resemble those of the living species of the sections Columbea, Bunya and Intermedia of the genus Araucaria than they do those of section Eutacta (cf. Wilde & Eames, 1952 ; White, 1947) ; but the combination of characters represented by their relatively small size and flat, some- what scale-like habit, broad base and rhomboidal form and appressed and imbricate arrangement cannot be matched in any known living species. Indeed, in the absence of information on leaf venation and epidermal characters there is no sound justifica- tion for their inclusion in the genus Araucaria ; and the only other existing genus A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 105 in which they might conveniently be placed is Araucarites. Araucarites has been used to define fossil cones or shoots of araucarian habit, for various reasons not identifiable with the genus Araucaria itself (Seward, 1919 : 256), though Seward pointed out (pp. 265-6) that the use of this generic name for sterile shoots is not in accordance with sound principles. More recent detailed studies of " araucarian " sterile shoots (Brachyphyllum, Pagiophyllum : Kendall, 1948, 1948^, 1949) have demonstrated that fossil genera which may be distinguished from Araucaria, in cuticular structure may have this habit ; though in the case of Brachyphyllum mammillare (Kendall, 1949) it is almost certainly related to cone structure resembling that of Araucaria. Where, as in the present material, it is not possible to make cuticular investigations, or any definite assignation to fertile material, it is most convenient meantime to retain the artificial genus Araucarites to include these branches, which are certainly of araucarian habit. As will be noted in a later section of this paper, the sterile branches are associated with seed cones of two highly distinctive types, one of araucarian and the other probably of taxodiaceous affinity : and in the case of the latter the foliation of the cone pedicel may be superficially compared with that of the separate leafy shoots, which have not been found in connec- tion with cones. The foliated and defoliated shoots may even include more than one species amongst them, though at present these cannot be differentiated. The name Araucarites sanctaecrucis is instituted for these sterile shoots, after the province in which the petrified forest occurs. (2) WOODS (PI. i, figs. 2, 3, 7.) These are represented by a large number of unbranched woody stems varying from about 8 cm. to 0-5 cm. in diameter, a few being still partially embedded in the rhyoli- tic ash (PI. i, fig. i, w). Most of these specimens have a complete thick cylinder of secondary xylem, but no distinctive external features : PI. i, fig. 2 shows a stem where part of an outer zone, probably representing remains of bark, may be distin- guished (c). The preservation of the tissues, so far as examined, is so poor that extensive cutting does not appear to be warranted : but it has been possible to make a few observations on the wood structure. In the stem shown in transverse section in PI. i, fig. 2, there is a narrow pith (p), and endarch primary xylem groups surroun- ded by a wide zone of secondary xylem (x) with several somewhat indeterminate growth rings, probably 12 in number. In radial longitudinal section the pitting of the radial walls of the tracheids of the secondary xlyem can be observed only in a few isolated patches where the organic matter of the walls has probably not been entirely replaced during silicification : this is illustrated in PL i, figs. 3, 7. The bordered pits are uniseriate and contiguous with slight flattening of outline where they adjoin ; but the structure of the medullary rays has nowhere been observed. In another larger specimen (8-5 cm. in diameter : V 397 6 ), probably from a branch 16 years old as judged by the ill-defined growth rings, similar uniseriate pitting has been observed, but again without preservation of medullary ray structure. This wood type so far as it can be defined is not identical with that reported by Gothan 106 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA (1925 : 198, pi. i, figs, i, 2) and Wieland (1935 : 16) in other specimens from the same locality. Gothan's wood, named by him Dadoxylon (Araucarioxylon) sp., show- ed radial tracheid walls with uniseriate bordered pits, sometimes distantly separate and rounded, sometimes contiguous, but with the indication of the cross-field pitting too indefinite for satisfactory use. Uniseriate contiguous pitting of the tracheids such as is illustrated here was a type widely represented in the Mesozoic (cf. Seward, 1919 : 165 ; Krausel, 1949) by a variety of " genera " of fossil woods e.g., Mesembrioxylon and Brachyoxylon. It has from time to time been interpreted as one transitional phyletically between the " pinacean " and " araucarian " types as these were formerly more or less rigidly defined (e.g., Hollick & Jeffrey, 1909 : 75) : Wieland regarded the Cerro Cuadrado woods as representing such a transitional type (1935 : 16). However, it has more recently been demonstrated that this type of xylem pitting cannot be regarded as necessarily indicative of transitional or ancestral Araucariaceae, since it is found within the range of variability in xylem structure in individual plants in modern genera of different families of the Coniferales (e.g., Pinus : Bailey, 1933 ; Sequoia : Bailey & Faull, 1934 ; Araucariaceae : Pool, 1929). According to Pool (1929 : 599) it is more common in the stem wood of Araucariaceae than is usually recognized. Without more details of the wood structure, therefore, and in particular of the medullary rays on which keys for wood identification have laid stress (Gothan, 1905 ; Krausel, 1919, 1945 ; Phillips, 1941), it is not possible to make any effective com- parison or identification of these woods. As will presently be shown, they are asso- ciated with two types of cone, one araucarian and the other probably most nearly related to the Taxodiaceae ; and the wood just described might have belonged to either-it corresponds closely with that described for the cone axis of the latter type. Similar wood structure has also been found, very poorly preserved, in the sterile foliated branches of araucarian habit already described. But it cannot be identified more specifically than as coniferalean wood. It may be noted in conclusion that Wehrfeld (1935) referred to abundant remains of araucarian petrified wood in the forest, but he adduced no evidence of structural detail. His illustrations are of uncut woody trunks and branches, some showing " annual " rings (e.g., p. 120, pi. f), and some of the trunks were very large, one incomplete specimen being reported as 100 metres in length (p. 125). If this were confirmed, it would seem that some of the trees of the Cerro Cuadrado petrified forest were perhaps the tallest that ever lived, and certainly the tallest that have been found in a petrified state. One would like to have precise measurements, not only of length, but of diameter at intervals throughout the length. As Fossa-Mancini has pointed out (1941 : 75), the theoretical maximum height of about 300 feet propounded by Galileo for a tree (cf . Thompson, 1942 : 28) was no doubt considerably exceeded by such a tree as that mentioned by Wehrfeld : and other reports (Feruglio, 1949 : 129) have also recorded very wide trunk bases in the forest. Leanza is quoted by Feruglio (1949 : 129, footnote) as reporting the base of a fallen trunk of circumference 10-3 metres (i.e., diameter approx. 3-26 metres). These accounts of gigantism in the trees of the petrified forest refer to the trunks as araucarian ; A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 107 as there is no account of wood structure to corroborate this, the identification has no doubt been made on the evidence of associated cones. The wood structure of branches so far examined structurally, as has been pointed out, is indefinitive in character for purposes of identification ; and as will later be shown, while one of the cone types found in the forest is araucarian, the other is probably most nearly related to the Taxodiaceae, in which family at the present day occur the most striking exam- ples of gigantism in trees, comparable with those of the petrified forest. It is perhaps more likely that these giant petrified trunks belong to this family than to the Arau- cariaceae, but more information on the details of their wood anatomy is needed. Wehrfeld also reported finding, more rarely, wood of palms. His description of these mentions woods showing yellowish spots which he interpreted as the remains of the principal vascular bundles ; but no structural evidence was given to support this interpretation, nor has it so far been possible to obtain evidence of the presence of monocotyledons from any other woods examined. The " yellowish spots " which he described may represent variation of the mineral differentiation in the stems rather than the position of vascular bundles ; for the petrifactions in the forest tend to show differentiation of colours in the quartz. (3) SEEDLINGS (PI. 2, figs. 14-25 ; Text-fig, i, A-C.) There are numerous (about a couple of hundred) petrified structures, many of them looking like small corms or carrots, which vary in shape from ellipsoidal to turbinate (PI. 2, figs. 14, 15, 17,), with intermediate forms such as that shown in PI. 2, fig. 16. Most of these specimens are detached : but PI. 2, fig. 14 shows one (s) lying prone and partially embedded in the matrix. The turbinate forms vary in size from 4 cm. in height and 5 cm. in widest diameter to 1-5 cm. in height and 2-2 cm. in diameter ; the ellipsoidal or clavate forms are more uniform in size, averaging 4-5 cm. in height and 1-5 cm. in widest diameter. None of these structures shows any continuation at either end of the axis into a leafy shoot or a well-defined root : PI. 2, figs. 18 and 19 show one specimen with a prolongation of the broader end of the axis which bears no evidence of insertion of lateral organs. From external examination of the specimens and from examination of cut and polished surfaces it may be seen that most are partially or completely decorticated (PI. 2, figs, 15, 17), and that a few have what appears to be a fairly thick periderm (PI. 2, figs. 18-20, 22, pd.}. The core of such specimens is composed of a thick zone of poorly preserved tissue resembling secondary wood (PI. 2, figs. 19, 20, 22, x) and a narrow region in the centre probably comprising pith and primary xylem. The decorticated specimens show on their external surfaces longitudinal fissures and markings (PL 2, figs. 15-17), some of which might indicate the position of primary medullary rays in the xylem ; however, as such markings may also be found on corticated specimens (PI. 2, fig. 18), they may be due, at least in part, to cracking during decay and preservation. Some thin sections have been prepared from the more promising of these specimens with a view to studying the arrangement and detailed structure of the tissues for io8 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA comparison with the gross morphology. The preservation is unfortunately very poor in most of the specimens cut : but PI. 2, figs. 23-25 shows the centres of trans- verse sections taken at the top (broader) end, middle, and base (narrower end) of a specimen similar to that shown in PI. 2, fig. 15, where preservation was better than in most. The outlines of the whole sections are shown in Text-fig. I, A-c (correspon- ding with PI. 2, figs. 23-25), where it can be seen that there is a medullated stele with the pith region (p) widening from the base upwards, whilst it also changes from rounded to ovate in shape in transverse section. PI. 2, fig. 21 shows this pith in median longitudinal section of the broader end of the specimen ; the innermost TEXT-FIG, i. Seedling (cf. Araucaria mirabilis). Transverse sections through a seed- ling at top, middle and base of swelling (cf. PI. 2, fig. 15), showing pith (p) in centre surrounded by primary xylem groups (pr) and secondary xylem (sec.), (cf. PI. 2, figs. 23-25)- V. 30951. x 5. elements of the xylem ring are narrower than those forming the bulk of the xylem, and the pith is notable for its more or less isodiametric, thick-walled cells. The bulk of this specimen is composed of poorly preserved tissue resembling secondary xylem traversed by medullary rays, in which structure of pits cannot be distinguished. The primary xylem groups which abut on the pith increase in number from below upwards, and the condition appears to be endarch, with stem structure throughout, and with no evidence of traces passing out to lateral appendages. Other sections from specimens similar to those in PI. 2, figs. 16, 17 show the same general arrange- ment of the tissues (cf. PI. 2, fig. 22) ; the structure of the tissue resembling secondary xylem can only be distinguished in a few better-preserved patches in some of the A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 109 longitudinal sections, where coniferalean pitting, of the type illustrated in PI. i, figs. 3 and 7, may be distinguished on the radial walls. These curious bodies must therefore be interpreted as swollen intercalary organs, having stem structure but without any evidence of lateral appendages, and with marked development of a tissue resembling secondary xylem. Their identification is a matter of some difficulty ; but the structures are so numerous, and the inter- calary swelling is so striking in all of them, that they must have some regular signi- ficance in the vegetation of the forest. Wehrfeld figured a number of such structures (1935 : 129), and referred to them as " a scientific novelty a series of fossil arau- carias," without, however, substantiating in any way the reference to the Araucaria- ceae. Gothan, too, in a recent brief study (1950), has described such structures as those in PI. 2, fig. 17 the turbinate or fig-shaped forms and believes the most likely interpretation to be that they are of the nature of insect galls, terminal on branches, where hypertrophy of the wood of the branch has been caused with the larva living in the centre, and with the gall eventually dropping off the end of the branch. He also believed they might be araucarian, but had found no evidence of any such galls in living Araucariaceae. It has not, however, been possible to find any evidence that the structures are other than intercalary, or that the turbinate forms are different in kind from the ellipsoidal forms ; and in some of the specimens sectioned there is a definite pith including thick-walled cells and with no sign of any larval intrusion. There remains, however, the possibility that these might be fungal stem galls of an intercalary nature ; such intercalary stem swellings are known in some cases to be caused by fungi, as in the case of Peridermium galls on Pinus stems ; but no records of such galls on araucarian stems have been found, though Connold (1909 : 132, text-fig. 159) has described long, irregular tumours on the roots of Araucaria imbricata which do not resemble the present structures at all. In the case of either terminal or intercalary stem galls, however, one would expect to find some evidence of lateral appendages, and there is none. On the other hand the size, shape and internal structure of these bodies all tend to emphasize their similarity to broken first-year seedlings of the genus Araucaria in the sections Bunya and Columbea, where tuberous development in varying degree is found in the hypocotyl, which is of stem structure with early incidence of secondary thickening (Diirr, 1865 : 103 ; Hemsley, 1901 ; Seward & Ford, 1906 : 333 ; Shaw 1909 ; Hill & de Fraine, 1909 : 212 ; Hickel, 1911 : 160). Wieland has figured certain branch-like structures (1935 : 27, pi. 13, fig. 2) as bases of second-year seedling stems of Proaraucaria, with broken root bases : but if these are indeed older seedlings, they are not markedly tuberous, and there is no evidence of any relationship with the smaller structures here described. The tuberous habit in seedlings in general is sporadic : it appears to be rare amongst gymnosperms, occurring in the Cycadales in Encephalartos , and in the Coniferales in the two sections of the genus Araucaria just mentioned. The proximity of these numerous tuberous bodies to fertile cones belonging to the section Bunya of the genus Araucaria, which will be described in the next section, also tends to corroborate their interpretation as araucarian seedlings : and the fact that the embryos in the seeds of these cones are dicotyledonous might suggest that the markedly ovate form no A CONIFEROUS PETRIFIED FOREST IN PATAGONIA of the pith at the upper end of the seedlings is related to a stelar adjustment below the level of the insertion of the vascular supply of two cotyledons on opposite sides of the hypocotyl axis. So far as has been ascertained, this is the only record of petrified seedlings. Kendall (1949, text-fig, i, j.) has drawn attention to a curious specimen, found as a compres- sion, from the Mid- Jurassic Estuarine Series of Yorkshire which she has interpreted as a seedling stage of Amucarites phillipsi Carruthers, but no other such records are known to the writer. Mr. W. N. Edwards informs me that there is a specimen in the Sedgwick Museum, Cambridge, from the Lower Greensand of Upware, Cambridgeshire, which was figured by Keeping (1883 : 150, pi. 8, fig. 7) as " a small fruit (?) ferruginized," but which superficially at least is very similar to some of the smaller araucarian seedlings from Patagonia. (4) CONES The cones, which represent the best preserved part of the collection, are of especial interest, since petrifactions of reproductive organs of fossil conifers are seldom found, though petrified coniferous woods are abundant. Two distinct types of seed cone are represented, but no male cones or pollen-producing organs have been found. These two types of cone have already been partially described by Spegazzini (1924), Gothan (1925), and Wieland (1935) ; but detailed studies of the structure, which affect the interpretation of the morphology and consequent assessment of the relation- ships of the cones, have not been made, with the exception of Darrow's investigation (1936) of the structure of the embryo of Araucaria mirabilis. Araucaria mirabilis (Spegazzini) (Pis. 3, 4, 5 ; Text-figs. 2-4) 1924. Araucarites mirabilis Spegazzini, p. 126, text-figs. 1-3, 4 (1-7). 1925. Araucaria windhauseni Gothan, p. 200, pis. 2-7, pi. 8, fig. I. 1929. Proaraucaria mirabilis (Speg.) Wieland, p. 60. 1929. Proaraucaria elongata Wieland, p. 60. 1931. Araucaria mirabilis (Speg.) Windhausen, p. 201. 1935. Proaraucaria mirabilis (Speg.) : Wieland, p. 19, pi. i ; pi. 7, figs. 2, 3 ; pi. x ; pi. xi, figs. i, 2, 4 ; text-fig. 4. 1935. Proaraucaria mirabilis var. elongata Wieland, p. 26, pi. 8, fig. i ; pi. 9, fig. 2 ; pi. 12, fig. 4. 1935. Proaraucaria patagonica Wieland, p. 26, pi. 6, figs. B, D, E, F. 1935. Proaraucaria mirabilis var. minima Wieland, p. 26, pi. 6, fig. A ; pi. 12, fig. 3. 1936. Proaraucaria mirabilis (Speg.) : Darrow, p. 328, text-figs. 1-13. 1936. Araucarites mirabilis Speg. : Gordon, p. 14. 1939. Proaraucaria mirabilis (Speg.) : Darrah, p. 222, text-figs. 133, 134. 1940. Araucaria mirabilis (Speg.) : Florin, p. 36. 1944. Araucaria mirabilis (Speg.) : Florin, p. 513. 1947. Proaraucaria mirabilis (Speg.) : Arnold, p. 313, text-fig. 156. 1947. Proaraucaria mirabilis (Speg.) : Andrews, p. 177, text-fig. 121. 1948. Proaraucaria mirabilis (Speg.) : Wilde & Eames, p. 312. 1949. Proaraucaria mirabilis (Speg.) : Feruglio, p. 129. 1951. Proaraucaria mirabilis (Speg.) : Feruglio, p. 35. A CONIFEROUS PETRIFIED FOREST IN PATAGONIA in EMENDED DIAGNOSIS. Seed cones varying in shape from spherical to somewhat ellipsoid, and in size from about 8 to 4 cm. in height and 8 to 4 cm. in diameter. Cone-scales numerous, arranged in a close spiral, one-seeded, with thick woody wing and deciduous laminar tip, which on dropping reveals the large woody ligule on top of the rhomboidal apophysis of the bract ; the fertile scale is fused with the bract for about two-thirds of its length, so that the depth of the ligular sulcus is about one-third of the length of the fertile scale. Cone-scales of mature seed cones (de- laminated) 13-16 mm. long and 10 mm. wide including the wing, tapering somewhat towards the base. Ligule 4 mm. wide, 1-2 mm. high, about 5 mm. long. Seed 8 mm. long and 3 mm. wide and deep, inverted, albuminous, apparently embedded in the fertile scale tissue ; testa with a thick stony layer. Embryo large, dicotyle- donous, with numerous spirally twisted suspensors at base. Evidence on mode of seed dispersal lacking. Axis of cone wide, with a wide pith including branched sclereids, surrounded by a narrow ring of separate vascular bundles each with a wide outer sheath of fibrous extra-phloem tissue. Vascular supply of cone-scale double, the bract trace single and arising separately from the base of the leaf-gap in the axis stele, the fertile scale supply of two strands, arising from either side of leaf gap, which fuse with inversion : both bract and fertile scale supply fork in base of cone-scale to give a lower and an upper (inverted) series respectively, the lower series passing out to the apophysis and laminar tip of the bract accompanied by a series of resin canals, the upper series terminating in the large strongly vascularized ligule. DESCRIPTION. The species is represented by a large number of specimens in the collection ; most of these are isolated specimens, but a few are still partially em- bedded in the matrix (PI. 3, fig. 26). Some of the cones must have been mature, as they contain fully formed embryos in the seeds, while others have no embryos, though the cones may be as large as those which do. The cones vary in size from 8 cm. in length and diameter to 4-4 cm. and 4 cm. in length and diameter respectively, and in shape from nearly spherical to ovoid or ellipsoid. There is no evidence to suggest that the smaller cones are necessarily immature, for one about 5 cm. in length and diameter (V. 30975) has fully formed embryos in the seeds. Some specimens appear more markedly elongate, with an ellipsoidal section, than others (e.g., PI. 4, figs. 38, 42) : such cones were interpreted by Wieland (1935 : 26) as a distinct variety, Proaraucaria mirabilis var. elongata, but structurally it has not been possible to differentiate these types, and there is evidence from the cone figured here that the condition has been exaggerated by considerable crushing prior to petri- faction, as may be seen in the compressed and broken outer parts of the scales in PI. 4, figs. 40, 42, while the proximal part of the scales has withstood the compression, being reinforced, perhaps, by the woody tissues of the testa. The cones have been petrified before the shedding of the seeds in most cases ; no separate petrified seeds nor cone-scales have been found, but there are a few specimens, such as those figured by Wieland (1935, pi. 6, E) and interpreted by him as another species, Proaraucaria patagonica, which no doubt represent the naked axes of the cones after the cone- scales have been shed (e.g., V. 30969). They correspond in size with the measure- ments of the cone axes themselves ; one has been sectioned, and shows a wide pith 112 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA with sclereids, and peripheral vascular bundles. The surface features correspond closely with those found in cones of living araucarians after shedding of the cone- scales (cf. Seward & Ford, 1906 : 361), with a reticulum of low ridges outlining rhom- boidal areas no doubt representing the areas of attachment of the scale bases to the cone axis. The absence of any separate cone-scales, which Gothan (1950 : 153) has emphasized as a feature suggesting that these cones did not shed their scales at maturity, and as one which might separate them genetically from those of other species of Araucaria, may well be explained in terms of the season in which the forest was overwhelmed. The seeds in the cone-scales in the final year of growth had not been shed, and the previous year's had germinated and are probably repre- sented in the forest by the very numerous petrified seedlings to which reference has already been made. Various views of external features of cones are shown in PI. 3, figs. 26-31, PI. 4, figs. 38, 39, and PL 5, fig. 46. Inserted on the cone axes are a large number of close-set ligulate cone-scales, each of which bears a single large inverted seed sunken in its tissue on the upper surface. The distal, abaxial faces of the cone-scales are seen arranged in a close spiral forming some 55 orthostichies (so far as has been examined). The surface morphology of the cone-scales varies, probably according to the age and condition of the cone and also to some extent to the degree of weather- ing of the surface, as shown in PI. 3, figs. 28, 29, 31, 34, PI. 4, figs. 38, 39, and PI. 5, fig. 46. PI. 4, figs. 38, 39, 42 show a cone where in surface view the cone-scales have laminar, longitudinally striated tips (t). In earlier descriptions of the cones, Gothan (1925, pi. 2) and Spegazzini (1924, text-figs, ib, 4 (6)) have illustrated some with a similar triangular lamina on the cone-scales, and Wieland mentions this feature (1935 : 175, pi. 10, fig. 2). Most of the cones, however, show surface mor- phology of cone-scales as illustrated in PI. 3, figs. 28, 29, and PI. 5, fig. 46, with prominent apophysis of bract (a) with marked lateral wings (w), and a large ligule (/). Consideration of sections of the specimen in PI. 4, figs. 38, 42, leaves little doubt that this latter condition is the result of the dropping of a laminar tip, though erosion of surface features may occasionally account for it. PI. 4, figs. 40, 42, show this laminar tip t, covering the ligule /, in radial longitudinal section ; and Text-fig. 4, A-I, and PL 4, fig. 43, PI. 5, fig. 54, from a tangential longitudinal series, demon- strate the same relationship between the laminar tip and the ligule. There is, too, some structural evidence of an absciss zone at the base of the laminar tip, e.g., at A in PI. 4, fig. 40. The lateral wings of the cone-scales are also seen to be present in the series of tangential sections, though they tend to be obscured in surface view (PI. 4, fig. 39) as compared with the cones with delaminated scales. In any respect other than the presence or absence of a laminar tip it is difficult to separate the two cone types : in both, seeds with fully formed dicotyledonous embryos of the same type have been found. The much smaller cone illustrated in PI. 3, figs. 31-34, shows similar longitudinally striated laminae clothing the exterior of the cone ; in section (PI. 3, figs. 32, 33) no ovules can be distinguished, and it may be either a young cone or one with abortive or unpollinated ovules. The comparatively large ligule can again be distinguished behind the distal lamina of the cone-scale (PI. 3, fig. 33,/). There seems little doubt that the three cone types just described represent A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 113 different stages in the development of the same cone species ; in many of the living Araucariaceae there is a laminar tip or spine on the cone-scale which often drops off when the cone is ripe (cf. Pilger, 1926 : 255). The ligulate cone-scales are modified in form towards the base of the cone, where they are sterile. PI. 3, fig. 30 shows the transversely cut tip of a cone pedicel (pi), around which are a number of such sterile cone-scales of simplified form, which pass over gradually into the fully formed fertile cone-scales above. This is a charac- ter of araucarian cones which has been emphasized by Seward & Ford (1906 : 354, text-fig. 22A) as contrasting with the condition in the Pinaceae where the cone-scales b.s. ^m^i \ B b.s. TEXT-FIG. 2. Araucaria mirabilis (Spegazzini). Transverse sections through vascular bundles of cone axis stele, showing from base upwards a series in the detachment of the bract supply (b.s.) and fertile scale supply (/i, /2) of cone scales. The two traces of the fertile scale supply fuse with inversion to give a single bundle (f.s.). The pith lies to upper side, cortex to lower side, of the vascular bundles, where xylem is shown black, with protoxylem white, phloem finely stippled, and extra-phloem tissue coarsely stippled. V. 30958. x 27-5. are much more sharply differentiated in form from the foliage leaves, and succeed them more abruptly. In none of the fossil cones examined is there critical evidence of the foliation of the pedicel, or of attachment to any of the leafy branches which are found amongst the remains of the forest. The axes of the cones are thick (PI. 3, fig. 27), with a wide pith and narrow ring of separate vascular bundles, seen at p and v respectively on the transversely frac- GEOL. II, 2. 8 114 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA tured surface and in Text-fig. 2 in transverse section. The pith contains branched sclereids ; the tissues of the vascular bundles are ill-preserved, but in patches of better preserved tracheids the xylem pitting varies from uniseriate to biseriate and alternating, with the pits separate, and each bundle is endarch, with a narrow zone of phloem, outside which is a deep arc of extra-phloem tissue (? pericycle) as seen in Text-fig. 2, e.p. This extra-phloem tissue is much better preserved than is the phloem, and consists of large, thick-walled, sometimes branching cells resembling the sclereids of the pith rather than a typical fibrous tissue. Comparison of this type of cone axis with that of other conifers reveals the striking similarity with that of living araucarian cones. Thomson (1913 : 4) has drawn particular attention to the exceedingly large size of the pith in the cone axis of the Araucariaceae, especially TEXT-FIG. 3. Araucaria mirabilis (Spegazzini) (cf. cone in PI. 4). Tangential longitudinal sections of cone-scales from base outwards, showing splitting of the two bundles b.s., f.s., in base (cf. Text-fig. 2), to give the vascular supply of bract and fertile scale respec- tively. Xylem shown black, with protoxylem white, extra-phloem tissue coarsely stippled. V. 30957. x 27-5. in the seed cones, where it may reach i in. in diameter. The sclereids of the pith and the extra-phloem or fibrous pericyclic tissue are, too, characters associated with the axis anatomy of the living Araucariaceae (cf. Seward & Ford, 1906 : 337). A study of the detachment of the cone-scale vascular supply from the axis stele has been made in transverse sections of the cone axis ; Text-fig. 2, A-F, illustrates successive stages, from below upwards, in the detachment of the traces, as seen in different sectors of .t he wide cylindrical stele. This has proved more satisfactory than the use of a series of sections, as 1-5-2 mm. may be lost between successive sections in preparing a series, while the spiral succession of the numerous scales is so close-set that from one section a close series of stages can be obtained. The bract supply ; A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 115 b.s., comes off as a single trace from the base of the leaf-gap, and free from the fertile scale supply, f.s., which is derived from the fusion of two strands, /i, /2, detached separately from either side of the leaf-gap above the bract supply. These two strands rotate through 180 while fusing, so that the resultant fertile scale fusion bundle is inverted (Text-fig. 2, D-F). In the base of the cone-scale there are, there- fore, two bundles, the upper one inverted. Text-fig. 3 shows their subsequent behaviour in tangential sections of the cone illustrated in PI. 4, fig. 38, taken in series from the inside of the cone outwards : Text-fig. 4, from the same series, shows H TEXT-FIG. 4. Araucaria mirabilis (Spegazzini) (cf. cone in PI. 4, and Text-fig. 3). Tangential longitudinal sections of cone-scales from base outwards, to show vascu- larization of bract (b), its laminar tip (t], and fertile scale (/) with ligule (/). Xylem shown black, sclerotic tissue coarsely stippled. S seed, T.S. = stony layer of testa, e = embryo, d resin ducts, f.s. = vascular supply of fertile scale, b.s. = vascular supply of bract. V. 30957. x 5. the relationship of the bundles to the cone-scale morphology throughout the scale and into the laminar tip. Each of these two bundles in the scale base branches, the bract bundle first (Text-fig. 3, D-E), and this lower series, though incompletely pre- served in the region below the seed (Text-fig. 4, E, F), appears to supply the apophysis and the laminar tip of the bract (Text-fig. 4, G, H, I ; cf. PI. 4, fig. 43 ; PI. 5, fig. n6 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 54). In the outermost region of the cone-scale (Text-fig. 4, G, H, and PL 5, fig. 54) structures resembling resin canals (d) lie below the bundles. PI. 4, figs. 43, 45 show obliquely cut bundles (v.b.b.) supplying the laminar tip of the bract ; these bundles are accompanied on either side by a strand of cells with dark contents (d), which may represent the upward extension of the resin canals into the laminar tip. It has not been possible to demonstrate whether, as in Araucaria bidwilli, there is a distinct system of bundles ending blindly in the apophysis, with the laminar supply branching off farther back. The upper inverted fertile scale bundle branches farther out in the scale base (Text-fig. 3, E-G), and forms a strong inverted upper series, probably supplying the base of the single inverted seed (Text-fig. 4, F) as well as the large ligule (Text-fig. 4, G, H, and PL 5, fig. 54). PL 4, fig. 44 demonstrates a vascular strand in the ligule. PL 5, fig. 53, taken from a delaminated cone, also demonstrates vascularization of the large ligule (/), along with the distinct series of the bract (b). There are two notable features here for comparison with the living species of Araucaria : the separate origin of the bract and fertile scale supplies, and the strong vascularization of the ligule. Only in the section Bunya Wilde & Eames amongst the living araucarians are these two conditions found (Eames, 1913 : 24 ; Aase, 1915 : 297 ; Wilde & Eames, 1948 : 322 ; 1952 : 44). This is probably the first time that it has been possible, in a fossil araucarian cone, to demonstrate these characters, which are of critical importance in considering the taxonomy and interrelationships of living araucarians. Text-fig. 4 and PL 5, fig. 54, also serve to demonstrate the lateral winging of the cone-scale ; this " winging," already noted in the external morphology of the cones, extends from near its base out to the apophysis, as seen in Text-fig. 4, B-H. A similar condition is found in the delaminated cone shown in PL 5, figs. 46, 50, 53. The wing is about as wide as the body of the seed, and is relatively thick ; it is supplied by vascular bundles of the lower (bract) series (Text-fig. 4, G, H, and PL 5, fig. 54). The tissues of the wing have evidently been woody, as may be seen in the cone in PL 5, fig. 50, where a core of thick-walled pitted cells is found, possibly of the nature of a transfusion tissue, and in PL 5, figs. 53, 54 where there is a deep hypodermal zone of thick-walled cells. Cone-scales as thick and woody and as widely winged as these must have been are found amongst living araucarians only in the section Bunya (Wilde & Eames, 1952 : 44). The morphology of the ligule, already noted in surface view of the cones, is seen in tangential longitudinal sections in Text-fig. 4, G, H, PL 5, figs. 53, 54, and in radial longitudinal section in PL 4, fig. 40, PL 5, fig. 52. Its vascular bundles have already been noted ; its ground tissue is similar to that of the wings, and no doubt it was also woody. The depth of the sulcus between the ligule and the bract may be exaggerated by artificial splitting between the two, extending apparently deep into the scale, the space occupied by clear quartz. The cone in which the split is most clearly and perhaps most naturally seen is illustrated in PL 3, figs. 35-37 : it will be noted that, where the scale is not quite medianly cut as at A in the radial longitudinal section of the cone in PL 3, figs. 35, 36, the split extends deeper than the level of the base of the seed, but where a more median cut of the seed is found, as at B, the split stops A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 117 towards the base of the seed. This is also demonstrated in the tangential longitudi- nal series of another cone in Text-fig. 3, F-H, where in Text-fig. 3, G, just outside the base of the seed, the ligule has separated from the bract on the margin but not yet in the median plane. It will be seen from PI. 3, figs. 35, 42 that the ligular sulcus is about one-third the depth of the fertile scale, the fertile scale being fused with the bract for about two-thirds of its length ; and this is about the same proportion for the depth of the ligular sulcus as in the living Araucaria bidwilli in mature cones (cf. Wilde & Eames, 1948 : 325 ; Pilger, 1926 : 256, text-fig. 139, A). Wieland based his generic definition of these cones as Proaraucaria on the greater depth and size of the split separating ligule from bract, as well as on the larger size of the ligule, in comparison with the condition in living species of the genus Araucaria (Wieland, 1935 : 19), and he regarded the fossil species as representing an intermediate stage in the evolution of the cone-scale, with less complete fusion of bract and fertile scale. His figures, however, nowhere make clear the natural depth of the split separating ligule from bract. From the present examination there is no evidence to show that the difference in degree of size of ligule and depth of sulcus separating it from the bract in the fossil and in living species is such as to warrant the erection of a new genus ; the fossil is closely similar to Araucaria bidwilli in these respects. The relationship of the seed to the upper surface of the scale is shown in Text- fig. 4, B-G, PL 4, figs. 40-42, and PL 5, figs 47, 50-52. The single large inverted seed (s) appears as if embedded in the scale tissue ; this is probably best seen in the tangential sections in PL 5, figs. 47, 50, the conspicuous stony layer of the testa (T.S.) lying within an outer sheet of tissue (F) which is continuous with the surface tissue of the wing of the scale on either side of the seed. Within the stony layer the nucellus is imperfectly preserved, but in many seeds the embryo sac may be defined, enclosing the endosperm with an embryo embedded in it (PL 5, figs. 47, 50-52, en and E). The dicotyledonous embryo is seen entire in PL 4, fig. 41, where the polished radially cut surface of the cone is photographed ; and embryos are seen cut in trans- verse section of the cone at E in PL 5, figs. 48, 51, and Text-fig. 4, E, in tangential longitudinal sections of the cone in PL 5, figs. 47, 49, 50, and in radial longitudinal sections in PL 4, figs. 40-42 and PL 5, fig. 52. Plate 4, fig. 41 and PL 5, figs 48 49 demonstrate the two cotyledons (co), and PL 4, figs. 41, 42, show the orientation of the embryo with the thick radicle pointing towards the micropylar end of the em- bryo sac, where there is a tangled mass of suspensors (ss) (PL 5, fig. 51). The embryo need not be described in detail as this has been done by Darrow (1936) ; but dico- tyledonous embryos of similar type have been found in both the laminated and delaminated types of cone (cf. PL 4, fig. 42, PL 5, figs. 47-51), and the orientation of the plane in which the median lines of the two cotyledons lie varies from one more or less parallel to the surface of the fertile scale (PL 5, figs. 47, 49) to one at right angles to it (Text-fig. 4, E). RELATIONSHIPS. From the fuller account of these araucarian cones which it has now been possible to give and with particular reference to the critical characters of mode of origin of the vascular supply of the cone-scale from the axial stele, size and nature of wing of cone-scale and size and nature of ligule, two major conclu- sions emerge which contrast with many of the previously expressed views on the n8 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA relationships of this species. The first is that there are no grounds for its taxonomic separation into a new genus Proaraucaria, which was effected by Wieland (1935) because he believed the cleavage between ligule and bract to be deeper than in any living species of Araucaria. Gothan later (1950 : 153) suggested, without further description of the cones, that the fact that they did not appear to shed their cone- scales at maturity was a character of major importance in separating the genus Proaraucaria : this condition has been discussed in an earlier section, where it was concluded that there is no evidence that the cones did not shed their scales, but a certain amount to suggest that they did. The second major conclusion is that the nearest related living species of Araucaria is A. bidwilli, the only species so far inclu- ded in the section Bunya recently erected by Wilde & Eames (1952) in their taxo- nomic revision of the genus Araucaria, and found to-day only in Eastern Australia in Queensland. Most of the previous references to these fossil araucarians have compared them with the two living South American species, A. araucana and A. angustifolia ; though Darrow compared the fossil cones with A . bidwilli, this was not substantiated by any critical study of the vascularization or of the ligule, but she effectively demonstrated the structure of the embryo, which is similar to that in A. bidwilli. It is true that in the fossil species there is no evidence for dehiscence of the individual cone-scales, for no separate seeds nor empty scales have been found ; and this character is one of those on which Wilde & Eames differentiated their new section Bunya (1952 ; 44). But in general structure of the cone-scale and ligule, and in particular of their vascularization, there is a striking similarity between the living Bunya and the fossil species : indeed, the larger size of the cones of A . bid- willi, which may reach 30 cm. in diameter, appears to be the chief difference between the species as represented by the cones. The use of the generic name Araucarites for these cones, though it has taxonomic priority, is best set aside (cf. also Florin, 1940 ; 1944) : for this name is reserved for cones or branches of araucarian habit which, however, are lacking in preservation of structural details which might give final evidence justifying inclusion in the genus Araucaria itself. The name Arau- caria mirabilis (Speg.) is therefore used, and the species is assigned to the section Bunya Wilde & Eames, the only other species of this section being the living Araucaria bidwilli, which is found to-day only in Queensland. The proximity of these cones to the seedling structures already described is certain- ly significant, though there is no evidence of organic relationship. Tuberous seed- lings are known amongst living species of Araucaria only in the section Bunya and in the South American section Columbea. The rarity of such a seedling habit in gymnosperms has already been noted ; and so far as records have been examined it is not found in any of the Taxodiaceae, to which the only other fertile cones in this area where the seedlings are found probably belong. Similarly, the proximity of the cones to the leafy branches of araucarian habit which have been described as Araucarites sanctaecrucis may be of significance, though evidence of attachment of cones to such branches is missing ; but as will presently be shown, the foliation of the pedicel in the Pararaucaria patagonica seed cones does not rule out the possibility that the branches, or some of them, may have been related to this quite distinct type of cone. A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 119 Pararaucaria patagonica Wieland (Pis. 6, 7 : Text-figs. 5-7) 1929. Pararaucaria patagonica Wieland, p. 60. 1929. Pararaucaria elongata Wieland, p. 60. J935- Pararaucaria patagonica Wieland, p. 21, pis. 2-5. 1936. Pararaucaria patagonica Wieland 1937. Pararaucaria patagonica Wieland 1940. Pararaucaria patagonica Wieland 1944. Pararaucaria patagonica Wieland 1947. Pararaucaria patagonica Wieland 1949. Pararaucaria patagonica Wieland 1951. Pararaucaria patagonica Wieland Gordon, p. 14. Darrah, p. 223. Florin, p. 36. Florin, p. 513, pi. 184, figs. 23-26. Arnold, p. 314. Feruglio, p. 129. Feruglio, p. 35. EMENDED DIAGNOSIS. Seed cones, ovoid in shape, varying in length from 4-7 cm. to 2-3 cm. and in diameter from 2-4 cm. to 1-3 cm., borne on slender pedicels clothed with spirally arranged, imbricate, broadly lanceolate and somewhat acute leaves with longitudinal striation of the abaxial surface. Bracts bearing the axillary fertile scales usually about 38 in number in average-sized cones, large and conspicuous, probably woody, each subtending a thick fertile scale, and arranged in a closely imbricate spiral succession with angle of divergence 3/8. Bract 10 mm. long, 12 mm. wide, 1-5 mm. thick, free from fertile scale for greater part of length ; fertile scale 10 mm. long, 12 mm. wide, 2-5 mm. thick, showing longitudinal ridges on its pro- truding distal abaxial face, and bearing usually one large inverted seed, laterally inserted, flattened and winged, the seed separating from scale at maturity. Seed 6 mm. long, 6 mm. wide, 2 mm. thick, with wing 2 mm. wide tapering towards base and tip of seed ; testa with inner stony layer, and outer layers composing wing of a characteristic stellate sclerotic lacunar tissue. Embryo 4 mm. long, poly- cotyledonary ; seed probably exalbuminous. Cone axis slender, with narrow pith and thick endarch cylinder of xylem. Succes- sion of elements in primary xylem includes scalariform and reticulate tracheids, passing over into elements with biseriate bordered pits. Secondary xylem tracheids usually with uniseriate bordered pits, contiguous and flattened, the pit apertures rounded or ovate ; medullary rays with cross field pitting of cupressoid form. No resin ducts or cells present. Bract supply arises from axis stele as a single trace from base of leaf gap ; fertile scale supply arising as two traces, one from each side of leaf gap above bract trace, the two traces fusing to give inverted bundle, which is accompanied into the scale on the adaxial face by a large strand of sclerenchyma which forks into two. Bract and fertile scale supplies single in base, branching higher up, with an abundant transfusion tissue linking the bundles tangentially and persisting in tips of bract and scale when bundles die out. Seed supplied by a single lateral strand from supply of fertile scale. DESCRIPTION. This species is represented by a large collection of mature seed cones ; no younger stages appear to be present, though it has been possible to section only a few of the specimens for more detailed examination. (One specimen, V. 3974 smaller than average with length 24 mm. and width 12 mm., probably represents an abortive or unfertilized cone, but preservation is not good enough for a convincing 120 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA identification.) The species is redescribed here with special reference to a number of diagnostic characters which were not included in Wieland's original account (Wieland, 1935) ; it is the most interesting species in the collections from the Cerro Alto localities, and its relationships with other conifers have not so far been assessed by comparison of a sufficiently wide range of characters. It may be of interest to note here that Feruglio (1949 : 129) reports, in a review of the localities where the plants of the petrified forest are known to occur, that Pararaucaria. has been found only in two of the localities with petrified cones, viz., the Cerro Alto, where it is associated with the cones of Araucaria mirdbilis, and near the Estancia Los Toldos, where the latter species is missing. The general habit of the cones is illustrated in PI. 6, figs 55, 56, 63. They vary in length from 4-7 cm. to 2-3 cm., and in diameter from 2-4 cm. to 1-3 cm., the average size being about 4 cm. long and 2 cm. in diameter. There is no evidence that smaller cones represent younger stages ; they probably represent attenuated cone forms, bearing fewer appendages. This range in size corresponds with that observed by Wieland in his specimens. A longer cone type was originally separated by him (1929 : 60) as Pararaucaria elongata, but later (1935 : 21, pi. 5, fig. 6) he included this type in Pararaucaria patagonica. A similar long cone is shown here in PI. 7, figs. 74, 75 ; and no other distinctive characters have been noted in it. These long cones are often poorly preserved and much weathered, as in PI. 7, fig. 74, but the bract and fertile scale show the same proportions as in smaller specimens. Cones of average size as well as these long cones have been found to contain fully developed embryos in the seeds (PI. 6, fig. 58 ; PI. 7, fig. 75), a condition refuting Wieland's suggestion that the latter represent the mature seed cones, while the former are immature. Such a range in size is a feature quite common in the cones of, for example, living species of Pinus. The cones are somewhat ovoid in shape ; many show marked unilateral weather- ing of the surface features and some a varying degree of compression. The specimen in PI. 6, fig. 63, approaches most nearly to the condition of the surface features in the living plants. A number of large and conspicuous bracts (b), about thirty- eight in an average-sized cone, are arranged in a close-set spiral around the axis, with successive bracts overlapping and with an angle of divergence of 3/8 as in other cones observed. In the axil of each bract is the fertile scale (/), showing slight longitudinal ridging of the abaxial surface, where it protrudes distally. Towards the cone apex the bracts and fertile scales diminish rapidly in size (PI. 6, fig- 56), the topmost members being sterile (cf. series of sections V. 30965, V. 30968). In some cones there is a markedly acute tip (PI. 6, fig. 56), in others it is more blunt (PI. 6, fig. 55). Only two specimens among the cones have been found attached to their pedicels : PI. 6, fig. 62, shows one of these, and clothing the pedicel is an overlapping spiral series of broad, flat, lanceolate leaves, as seen in more detail in PI. 6, fig. 64. These leaves are regularly striated on the abaxial surface ; the furrows may represent the position of lines of stomata, but it has not been possible to verify this from structural detail in either sections or cuticles. Another feature of some diagnostic interest is also illustrated by this specimen : it shows a sharp transition from the sterile leaves to the bracts and fertile scales at the base of the cone, a A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 121 character noted, for example, by Seward & Ford (1906 : 354) as one offering a marked distinction between the Araucariaceae and the Pinaceae, the transition in the former being much more gradual. The narrow cone axis is seen in transverse section of the basal region of a cone in PI. 7, fig. 65, and shows a relatively narrow pith and thick cylinder of xylem : the xylem diminishes in thickness towards the tip of the cone. The pith includes large scattered cells with dark brown contents : the stele is endarch, with scanty primary xylem and a wide zone of secondary xylem with regular radial files of tracheids, without parenchyma and without resin ducts or cells, and with unseriate secondary medullary rays. The structure of the xylem is illustrated in more detail in PI. 7, figs. 66-72. The centrifugal succession of elements in the primary xylem is shown in radial longitudinal section from right to left in PL 7, figs 66-68. Annular tracheids cannot be identified with certainty ; but a succession of spiral elements, in which there is some evidence for the formation of bordered pits between the turns of the spiral band (PI. 7, fig. 67, s.p.), is followed by widely reticulate tracheids (PI. 7, fig. 67, ret.) and these are succeeded, in the base of the cone, by tracheids with bi- seriate alternating bordered pits (PI. 7, figs. 70, 71, bi.), probably belonging to the secondary xylem. Beyond these come the tracheids forming the bulk of the woody cylinder ; these have uniseriate bordered pits on the radial walls (PI. 7, figs. 68, 69, 70, sec.). The pits are usually contiguous, with flattening of the pit outline in the area of contact (fig. 69) : the form of the pit apertures varies from rounded to oval. Parts of the medullary rays are seen in radial longitudinal section in PI. 7, figs. 68 70, 72 (m.r.) The rays are 1-5 cells high and ray tracheids have not been observed. The cross- field pitting is fairly constant (PI. 7, fig. 72), there being 2-4 horizontal rows of rather small pits, which are of markedly " cupressoid " form (cf. Phillips, 1941: 267). These characters of the xylem, not noted by Wieland, may most conveniently be discussed here : they represent features which have been widely used in the past as a means of identifying conifer woods. Bailey has recently re-emphasized the comparative significance of the primary developmental succession in the xylem of gymnosperms (Bailey, 1949, 1925 ; cf. also Florin 1936, 1937) ; and in Pararaucaria the sequence which has just been noted appears to be intermediate between that found in the " lower " gymnosperms (Cycadales, Cordaitales) and that in the "higher" gymnosperms (Coniferales, Ginkgoales, Gnetales). In the former groups scalariform elements are present and are succeeded by reticulate elements, and these by elements with the typical bordered pits of the metaxylem and secondary xylem ; bordered pits are not formed in the spiral tracheids, where, however, they do appear in the shortened succession found in the xylem of the latter groups, where scalariform and reticulate forms are usually absent. In Pararaucaria the presence of reticulate tracheids in the succession is comparable with the similar condition noted by Bailey (1925 : 593) as occurring sporadically in Araucaria and Agathis, which he has interpreted as intermediate bet- ween " lower " and " higher " gymnosperms in this respect. Though it is true that Bailey's observations were made on stem wood and the condition in Pararaucaria is 122 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA reported from the cone axis only the comparison is worth recording, though fuller knowledge of developmental sequence in the cone axes of the various groups is not available for comparison. The structure of the secondary xylem, in particular the radial pitting of the tracheid walls and the cross-field pitting of the secondary medullary rays, has often been used as a comparative basis for assessing relationships in coniferales. The variability in form and arrangement of tracheid pitting which has been demonstrated in stem and root wood in individual plants in a variety of families of the Coniferales is, however, such that the xylem type in the Pararaucaria cones might be included within the range found, for example, in Pinus, Sequoia, or Araucariaceae (Bailey, 1933 ; Bailey & Faull, 1934 ; Pool, 1929). Cross-field pitting in stem and root wood is also a variable character (e.g., Bannan, 1944). Cupressoid form of the pits in the cross-field is found fairly regularly in the Araucariaceae and Cupressaceae, and it also occurs amongst the Podocarpaceae, Taxodiaceae, Taxaceae, and even occasionally in the Pinaceae (cf. Phillips, 1941 : 268, 274-277). The condition in the Araucariaceae and in Taxodium, where the cupressoid cross-field pits may be as numerous as in Pararaucaria, probably conforms most nearly with that in the fossil cones. But details of cone-axis wood are available for comparison in very few cases : in Araucaria the records given by Thomson (1913, pi. 4, fig. 34) are of multiseriate pitting of the radial tracheid walls of typical araucarian form. This, together with the very wide pith and narrow ring of xylem (Thomson, 1913 : 4, pi. i, figs. 5, 7) serves readily to distinguish araucarian cone axes from those of Pararaucaria. No satisfactory direct comparison with cone axis structure of other families of conifers can be made without further investigation of these. The evidence of the cross-field pitting taken from stem wood structure would, however, tend to favour a reference of the fossil cones to relationship with the Taxodiaceae, where too the same type of tracheidal pitting has been demonstrated in the root of Sequoia semper- virens (Bailey & Faull, 1934, pi. 103, fig. 29). [See note on p. 138]. The mode of attachment of the vascular supply of the bract and fertile scale to the axis stele is illustrated in Text-fig. 5, A-D, which is taken from a series of sections at intervals of approximately 2-5 mm. apart, in the basal half of a cone, and viewed from the base upwards. It has not been possible to follow out the course of the bundles from a series through a single leaf-gap : but the vascular supply may be interpreted from a number of gaps cut at varying levels. The bract supply (b) comes from the base of the leaf-gap as a single strand, being separate in origin from the supply of the fertile scale, which is derived as two separate strands (Text-fig. 5A, f l , f z ,) from the sides of the gap above the bract supply ; these two strands subse- quently fuse to give one large bundle (Text-fig. 5, B-D, /) after each has undergone torsion through 180, so that the protoxylem comes to lie on the outer (abaxial) face of the fusion bundle (/), which is thus inverted alongside the bract supply (b) in the cortex ; the phloem is not preserved. This is also illustrated in PL 6, figs. 57, 58, (b.s., f.s.). A striking feature, though of subsidiary comparative interest, is the bulky strand of sclerenchyma which accompanies the inverted vascular supply of the fertile scale on its adaxial face (PI. 6, figs 57, 58, sc. ; Text-fig. 5, stippling). This strand, which has its origin in the cortex in the angle above the leaf-gap, forks A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 123 into two as the fusion bundle passes out of the cortex into the base of the fertile scale, and the paired strands remain a prominent feature in the fertile scale for about half its length (Text-fig. 6). This account of the origin of the bract and fertile scale supplies does not agree with that of Wieland. He has described, from cut surfaces of cones, the origin of the bract-fertile scale supply as a " single heavy concentric strand " (1935 : 22, pi. 4, A B rm, -b TEXT-FIG. 5. Pararaucaria patagonica Wieland. Transverse sections of stele of cone axis from below upwards, to show detachment of bract supply (b) and fertile scale supply ( /i , /2) , from leaf-gaps. The two traces /i , /2 of the fertile scale supply fuse with inversion to give a single bundle (/). Xylem black, with protoxylem white; scleren- chyma of fertile scale in double stippling. V. 30964 c-f. X 6-7. fig- 3 ' pl. 5> fig 5)- He nevertheless interpreted this condition as being in exact agreement with that in Pinus, where however, according to the account of Aase (1915), the bract trace is normally in its origin quite free (except at the cone base) from the fertile scale supply, as indeed is the case in others of the Pinaceae which have been examined (e.g., Aase, 1915 ; Radais, 1894). According to the observa- 124 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA tions on which the present description is based, Paramucaria does resemble the Pinaceae in this respect, as also the section Bunya of Araucaria, and some of the Taxodiaceae (Aase, 1915 : 294-7) ; and it has not been possible to confirm Wieland's observations. In his pi. 4, fig. 3, the supplies to the bract and fertile scale are clearly separate but in his pi. 5, fig. 5, there is some evidence that a single strand is present in the cortex outside the leaf-gap : this latter condition, however, is in the cone base, and such a condition has been noted by Aase (1915 : 282-5, 296) as occurring in the base of Pinus and Cryptomeria cones, though the normal condition higher up is to find the bract and fertile scale supplies quite separate in origin. The external features of the bract and fertile scale at the cone surface have already been noted : PI. 6, figs. 57-60 and Text-fig. 6 show their structure in series of tangential longitudinal sections in sequence from inside outwards. The most conspicuous characters are the thick, persistent and probably woody bract (b), subtending the thick woody fertile scale (/), which bears on its upper surface a single large, inverted seed, which is flattened and winged. The bract is free from the fertile scale for the greater part of its length, and is fused with it only towards the base (PI. 6, figs. 57-60 and Text-fig. 6). Here the double vascular supply, with the upper bundle (/.s.) inverted and accompanied by the two strands of sclerenchyma, is a prominent feature. The lower bundle, which supplies the bract, is single below but forks about half way up the free part of the bract, and the thick, protruding tip of the latter is supplied by a number of small vascular strands linked tangentially by a bulky tissue composed of isodiametric pitted cells, which may be described as a transfusion tissue (PI. 6, figs. 59, 60, and Text-fig. 6, TT). A curious feature of the bract as described by Wieland was that it was " pleated " (Wieland, 1935, pi. 4, figs. 4, 5). His illustrations show this in respect of one flank only of the bracts, and not in all of them. Closer examination of his figures and of the sections of the series here described has provided evidence (cf. PL 6, fig. 57) that this apparent lateral forking or pleating is found only in the bracts cut in inner tangential planes just outside the region where they are attached to the outer cortex of the axis. Here they are so close-set that the margins of their bases tend to be laterally con- fluent where they overlap, so that a kind of reticulum is formed on the surface of the axis. This is not necessarily a characteristic feature, but one correlated with the closely imbricate succession of the bracts : it is certainly not one characteristic of the free distal region of the bract. The fertile scale (/) is of similar size and thickness to the bract, and is also supplied in its basal region by a single vascular strand (/.s.) which remains unbranched until a short distance behind the distal attachment of the seed to the scale. Here it forks into 3-4 strands, and one of these, a lateral strand, passes up to supply the seed in the region of the basal foramen (PL 6, fig. 59, and Text-fig. 6F,/.s./.). Above the insertion of the seed the vascular strands supply the prominent tip of the fertile scale ; and here too there is a well-marked transfusion tissue as in the bract. The conspicuous strands of sclerenchyma noted in the base of the fertile scale fade out about half-way up the scale : their function may have been connected with the separation of the seed scales at maturity to liberate the seeds. Though the epi- dermal and hypodermal tissues of the scale in this region are poorly preserved, there A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 125 is some evidence (PI. 6, fig. 60, and Text-fig. 6n) of a grooved abaxial surface on the exposed distal face of the scale, possibly a natural feature, and one which has been noted externally as giving a longitudinally ridged appearance in a few of the best preserved cones (PI. 6, fig. 63). H TEXT-FIG. 6. Pararaucaria patagonica Wieland (cf. PL 6, figs. 57-60). Tangential longitudinal sections of fertile scales (/) with seed, and bracts (6), taken from base outwards, (j = transverse section.) Xylem black, sclerenchyma of fertile scale double stippled, stellate sclerotic wing tissue (w) of testa single stippled, transfusion tissue (TT) stroked. For. = foramen of seed, fsl lateral bundle of fertile scale supplying seed, ts = stony later of testa, Er = radicle of embryo, EC = cotyledons of embryo, e.s. = ? embryo sac, n ? nucellus. (A, B, V. 30959 c, e ; C-G, V. 30961 ; H, i, V. 30960** ; j, V. 309450). x 5. 126 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA The seeds themselves are large and flattened ; the latter feature appears to be general and may be a natural character. The usual condition is to find one seed per scale, inverted and with its basal attachment and vascular supply lateral as shown in PI. 6, fig. 59 and Text-fig. 6r. Wieland reported one cone only in part of which he found 2 seeds per scale, but this condition appears to be exceptional (Wieland, 1935 : 22). All the cones examined appear to be mature ; fully formed embryos are found preserved in the seeds of some of the cones sectioned, and the micropyles, directed towards the cone axis, are sealed (PL 7, fig. 73). The seeds at first sight appear to be embedded in the tissue of the fertile scale on its upper surface ; but closer examination shows that this is not the case. There is a con- spicuous " stony " layer in the testa (PL 6, figs. 57-59, Text-fig. 6, t.s.}, consisting of a deep zone of elongate, thick-walled cells lying with their longer axes at right angles to the surface. This layer is continuous except for the basal foramen in the region where the seed is attached. The stony layer is surrounded by an outer layer, confluent on either side of the seed body with a flattened wing (PL 6, figs. 57-60 and Text-fig. 6, w), which consists of a characteristic lacunar tissue of stellate, thick-walled cells or sclereids (PL 6, fig. 61), like the " cellules etoilees " of Radais (1894 : 231) in his description of the wings on the seeds of Abies. This tissue, resembling an aerenchyma, is markedly different from the tissue of the fertile scale below, to which it is closely adpressed. It corresponds exactly with the tissue de- scribed by Radais (1894 : 231, pi. 4, fig. 47) and von Tubeuf (1892 : 205, text-fig. 18) in the wings of the seeds of the Pinaceae, where, in the earlier stages of develop- ment, the whole of the inner surface of the seed, including the wing, is attached to the upper surface of the fertile scale, and separation only takes place at a later stage as the seed matures. The evidence from the specimens of Pammucaria here examined, which have all been mature, suggests that the seed, including the wing, is separate from the surface of the scale, except in the region of attachment at the basal foramen (PL 6, figs. 57-59 and Text-fig. 6). Text-fig. 7 illustrates the relationship of the seed to the scale in this region : and it can be seen that there is an " absciss " zone here consisting of an aerenchyma-like tissue continuous into the foramen, suggesting that at maturity there is rupture of the tissue in this stalk-like region. This rupture would finally liberate the ripe seed. Without younger stages, it is hardly possible to decide whether the outermost " wing " of the seed coat is at an earlier stage attached to the scale, as in the living Pinaceae, or free from it, as in the living Agathis (where the single seed is median) and some of the Taxodiaceae with inverted winged seeds (Taiwania, Taxodium}. This point, though a critical one, must remain undecided. Wieland's specimens were probably also mature and clearly show the separation of this wing (Wieland, 1935, pi. 4, fig. 5, pi. 5, fig. 2) which he did not recognize when he interpreted the condition of the seeds as comparable with that in Araucaria, "inclosed by the drooping curtain-like, expanded, and flattened and coalesced tip of the seed scale " (p. 22). It was principally on the basis of this feature, associated with the single seed, that he drew his comparison with the genus Araucaria, and instituted the name "Pararaucaria" to emphasize the intermediate nature of the genus; for he interpreted the slender cone axis and narrow pith as pinacean characters, together with the more or less free bract (p. 23). Florin, however, A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 127 described Wieland's "curtain-like" structure as the sarcotesta of the seed coat (1944 : 513), though he did not recognize the seed as winged. The tissues of the seed inside the stony layer of the testa are poorly preserved in most specimens : but PI. 6, figs. 57, 58 and Text-fig. 6, c, E, illustrate a cone in B TEXT-FIG. 7. Pararaucaria patagonica Wieland. A. Longitudinal section of fertile scale (/) and bract (b) of cone, cut to one side of median plane to show foramen of seed (For.) in region of attachment to scale, ts = stony layer of testa, w wing tissue of testa : xylem of cone axis and fertile scale shown in black. V. 30966*3. x 6-7. B. Tissue shown at B in foramen of seed in fig. A. x 240. c. Tissue shown at C in fig. A. x 240. In B and c, intercellular spaces stippled, thick- walled 1 fibrous cells in solid black. which embryos were preserved. The remains of the embryo sac are probably represented at e.s., and of the nucellus at n. The embryo is seen cut transversely in the region of the radicle towards the micropylar end of the seed in PL 6, fig. 57. and Text-fig. 6, c, and in the region of the cotyledons, towards the base of the seed, in PI. 6, fig. 58, Text-fig. 6, E. There are several (probably eight) cotyledons. PI. 7, figs. 73, 75 show an embryo in radial longitudinal section in another cone, with the radicle tip (r) pointing towards the sealed micropyle (m). This is the only specimen in which it has been possible to demonstrate an embryo in median longitudinal section 128 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA to confirm the orientation. In Wieland's account (1935 : 22) he refers to a suggestion of a dicotyledonous embryo, but the preservation of his specimens left this point inconclusive. Butts & Buchholz (1940) have shown that in the Pinaceae, polycotyledonary embryos with cotyledons reaching nine in number are a characteristic feature, while in the Taxodiaceae, the number varies among the genera from 8 to 2. But in the remaining families of the conifers embryos with 3 or more cotyledons do occur in certain genera, though the usual number is 2, so that the character has a limited diagnostic value. RELATIONSHIPS The chief features of Pararaucaria patagonica here described which were not noted by Wieland (1935) in his account of the species are : the foliation of the pedicel ; the details of the wood structure ; the detachment of the bract trace from the cone stele separately from the two traces which form the supply of the fertile scale ; the winged, detachable nature of the inverted seed and its lateral position ; and the polycotyledonary embryo. Wieland's account established the characters of slender cone axis with narrow pith, and the conspicuous bract scales of the cone bearing in their axils the large fertile scales each normally with one large inverted seed, which he interpreted as being enclosed in the scale tissue. On this association of characters he assessed the systematic position of the genus as intermediate between Pinaceae and Araucaria (Wieland, 1935 : 23), and accordingly instituted the generic name Pararaucaria. A re-analysis of the relationships of the genus is attempted in the following table on the basis of the fuller set of criteria of comparison which can now be used. The available characters of the petrified cones which might be suitable for giving a quantitative index of aggregate intergeneric differences are too few in number for effective use : but the table summarizes the position qualitatively. Pararaucaria patagonica Wieland, emend. List of Characters used for Comparison with some other Coniferales, Numbered for Reference as in Table below. 1. Bract free from fertile scale for greater part of length. 2. Bract large and conspicuous at maturity. 3. Bract trace distinct in origin from traces of fertile scale. 4. Usually one seed per fertile scale. 5. Seeds inverted. 6. Seeds attached laterally. 7. Seeds winged. 8. Seeds free throughout development from fertile scale, except in region of basal foramen. 9. Several cotyledons (> 4) in embryo of seed. 10. Cone axis slender, with narrow pith in stele. 11. Secondary medullary rays of cone axis wood with pits of cross-field cupressoid and numerous. A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 129 TABLE SHOWING POSITIVE CORRELATIONS WITH SOME FAMILIES AND GENERA OF CONIFERALES. Characters compared numbered as in list above. + = positive correlation in all species described. = positive correlation in some species described. ? = doubtful or undescribed character. Characters of bract, fertile scale and seed. Cone axis I. 2. 3- 4- 5- 6. 7. i 5. 9. 10. > n. + + + + + + + ? + + + + + + ? + Pararaucaria Araucaria Agathis . - ... + + + + + 4- Pinaceae . . .+ + + + 4- +.+ Taxodiaceae . ... + + + . 4- *Romeroites . . ... + ? ? + ?.4- ? *Cheirolepidaceae . ? .. + + ?.? ? *Pseudo- Araucaria ... ? ? .. ? + ?.? ? * The data for these fossils, which superficially bear some resemblance to Pararaucaria, are taken from the accounts given by Spegazzini (1924 : Romeroites), Hirmer & H6rhammer (1934 : Cheirolepi- daceae), Fliche (1895: Pseudo- Araucaria). A revision of the characters in all these types, especially of the relationship between seed and fertile scale, is necessary before any critical systematic comparison with Pararaucaria can be made. It will be seen on summation of positive correlations that the nearest comparison is with the Taxodiaceae (sensu Pilger, 1926 : 342), which shows the greatest number of positive correlations. Even allowing for uncertainty on criterion (8) in Para- aucaria (freedom of seed from scale throughout development), the comparison with the Taxodiaceae is closer than with the Pinaceae, for on criteria (2) and (n) (large size of bract ; and cross-field pits numerous, cupressoid) the conditions found in Pararaucaria and amongst the Taxodiaceae do not characterize the Pinaceae : and on criteria (i) and (4) (degree of freedom of bract from fertile scale, and number of seeds per scale), which represent the chief differences between Pararaucaria and the Taxodiaceae, it may be noted on (i) that in Cryptomeria (and probably also in the Jurassic genus Elatides) in the Taxodiaceae there is a form intermediate in respect of degree of fusion of bract and fertile scale, and on (4), that in Taiwania in the Taxodiaceae a single seed instead of two may sometimes be found, while in Pararaucaria the two-seeded condition has abnormally been found. In these two respects Cryptomeria and Taiwania respectively may be regarded as intermediate between the Pararaucaria condition and a more typical taxodiaceous condition. Certain features of Pararaucaria have been omitted from the lists above as not affording sufficiently reliable distinctions : these are the foliation of the pedicel, the sharp transition from the foliation of the pedicel to the bracts and fertile scales of the cone itself, the size of the cones and the number of cone-scales per cone, the radial pitting of the tracheid walls, and the vertical ridging of the fertile scales. But it may be noted that in each of these features correspondence with characters occurring amongst the Taxodiaceae may be found. In the case of the radial pitting GEOL. II, 2. 9 130 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA of the tracheids, reference may be made to the range of variability in xylem structure demonstrated in the Redwood, Sequoia sempervirens , by Bailey & Faull (1934, pi. 103, fig. 29), where the condition in the fossil cones is found in secondary xylem of root. The vertical ridging of the fertile scale is, so far as I am aware, a feature characteristic of the Taxodiaceae in the genera Sequoia, Sciadopitys, and Taxodium, though in the first two of these it is also found to some extent in the bract. Finally, the general "habit" of the cones is taxodiaceous ; the proportion of bract to fertile scale, the size of the cones, the vertical ridging of the fertile scale, all combine to give a habit which on superficial examination one would tend to refer to the Taxodiaceae. In the analyses just given, the Taxodiaceae are referred to in the sense of Pilger's definition (Pilger, 1926 : 347) ; the family in this widely-defined sense shows con- siderable diversity in certain characters, e.g., the orientation of the seed, and the number of the cotyledons. The probably heterogeneity of the Taxodiaceae in this inclusive sense has, however, been generally recognized : Pilger himself separates the genera into two sub-families and a number of sub-groups, while Hayata (1932) arranged the genera of Pilger's group in a number of separate families. Florin has more recently re-examined the relationships of the Taxodiaceae, sensu Pilger, (Florin, 1931 : 484-491 ; 1940 : 78). He agrees (1931 : 490) with Pilger's classi- fication, whilst admitting it may be artificial, since it deals with a family inter- mediate in its general characters between others more sharply defined : but at the same time he has re-emphasized its diversity and probable heterogeneity. In particular, he draws attention, amongst other distinctive features, to the character- istic epidermal characters of the Tasmanian genus, Athrotaxis, which can readily be used to distinguish it from other genera of Taxodiaceae in the northern hemisphere. In view of this diversity in the Taxodiaceae, sensu latu, Pararaucaria may mean- time most conveniently be referred to this family. It most nearly resembles Tai- -wania in the one-seeded condition, Cryptomeria in the condition of the bract in relation to fertile scale, and Sequoia and Taxodium in the polycotyledonary embryo, but resembles no one living genus in its combination of characters. The possibility remains, however, that the seed and its wing may have been attached, in earlier stages of development, to the fertile scale as in the Pinaceae : and in this case the most natural position would be to place Pararaucaria in a separate new family of Coniferales, intermediate in position between the Taxodiaceae and Pinaceae. In a recent review by Takhtajan (1953) the phylogeny of the Taxodiaceae has again been discussed, and he suggests that they probably originated from the earliest and most primitive Pinaceae, though no fresh evidence is adduced in support of this view. Wieland (1935 ' 24) referred Pararaucaria to the fossil family Cheirolepidaceae as defined by Hirmer & Horhammer (1934 : 79) : but this reference was based on the assumption that the seed was borne in the scale tissue as in the genus Araucaria, and as has already been demonstrated, this is not the case. Florin (1940 : 36 ; 1944 : 513) suggested that Pararaucaria might be identical with the South American fossil genus Romeroites, from the province of Neuquen. This genus, as described by Spegazzini (1924 : 134), is undoubtedly taxodiaceous, but differs from Para- raucaria chiefly in the thin, many-seeded fertile scales with seeds upright and fleshy, A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 131 and fusion of bract with fertile scale for the greater part of its length. A more detailed description of this genus is undoubtedly desirable, but according to Spegaz- zini's account it does not correspond with Pararaucaria, though the size of the cones and the general external habit are somewhat similar. Gothan (1950 : 153) in a very brief note recognized that Pararaucaria had nothing to do with Araucaria, and accordingly indicated that the genus was misnamed: but he gave no further description of the cones nor indication of their true relationships, of which, as has been shown, no more satisfactory definition can be given unless younger stages in the development of the cones are found. DISCUSSION The chief interest of these conifers of the Cerro Cuadrado forest lies in their relationship with other conifers and their geographical distribution. Araucaria mirabilis is an extinct species, and appears to have its nearest living relation in the Queensland " bunya bunya," A. bidwilli, and not in A. araucana or A. angustifolia, the only living species of Araucaria in South America ; but it might be considered to represent an ancestral form of these, if the wings of the cone scales had become reduced in the evolution of the species. This view on the evolution of the South American living araucarians has already been expressed by Darrow (1936 : 333). The South American fossil records (cf. Florin's review, 1940 : 33-40) do not include any which furnish critical evidence of the wingless cone scales of species of the section Columbea Endlicher, emend. Wilde & Eames : the cone scale from the (?) Upper Cretaceous of the Cerro Guido in Santa Cruz, which was named Araucarites patagonica by Kurtz (1902 : 49) and referred to as comparable with the living Araucaria brasilien- sis, was not figured, and was an incomplete specimen described as representing the lower part of the scale. Other cone scales in the form of compressions, from Meseta de Baquero in Santa Cruz, have also been referred to the section Columbea (Feruglio, 1951 : 65). These are more or less incomplete and are not illustrated, but from the description and from their comparison by Feruglio with Berry's cone scales of Arau- caria from the nearby Gran Bajo de San Julian (Berry, 1924 : 480, text-figs. 2, 2a) it appears that they are winged scales. Berry refers to his as " obviously thick and woody," which suggests that they may have belonged to the section Bunya Wilde & Eames though they are much smaller than those of the living A. bidwilli. Other seed scales, which he also compared with Berry's, were described by Feruglio as Araucaria (Feruglio, 1951 : 39), from the Gran Bajo de San Julian, and they too might have belonged to the section Bunya. Both Berry's and Feruglio's seed scales come from localities in the region of the same volcanic complex (the porphyritic series of the Bahia Laura), which Feruglio has dated as of age mid- Jurassic to Wealden (1951 : 74), and which also contains the petrified forest around the Cerro Alto ; but it is unlikely, owing to the incomplete preservation of the compressed cone scales, that any close comparison can be made between them and the petrified cone scales of Araucaria mirabilis in the forest, though they are not dissimilar in size and general form. Feruglio does not believe that the San Julian specimens are identical with Araucaria mirabilis. GEOL. II, 2. 9 132 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA There is, therefore, as yet no critical record of wingless cone scales in South Ameri- can fossil floras. The evidence so far provided from sterile shoots with broad flat leaves might indicate araucarians of the sections Bimya Wilde & Eames, Intermedia White, or Columbea Endlicher emend. Wilde & Eames ; and petrified woods have not yet provided reliable evidence for distinction of these sections. The South American records are, however, relatively scanty, and the absence from them of critical evidence of fossil species of Columbea, sensu Wilde & Eames, is necessarily inconclusive for assessment of the age of this section on the South American conti- nent in comparison with that of species of the section Bunya, of which A . mirabilis is the only fossil form with critical structural evidence of relationship. Wingless araucarian cone scales are rare in the fossil records of the Southern Hemisphere as a whole (Florin, 1940 : 27), but they have been reported from the Jurassic of Australia and New Zealand (A. grandis Walkom, 1921 : 13, pi, 3, fig. 6 ; Edwards, 1934 : 100), suggesting that they may have been as ancient as the winged types, which are abundant in Jurassic rocks, and indicating that they did occur in Mesozoic times in Australasia, where they are now extinct The evidence, therefore, though scanty, does not support the attractive hypothesis of the South American evolution of Columbea ; and accordingly, A . mirabilis probably bears no direct relationship to the living South American species, but represents an extinct element of an earlier more widespread araucarian flora, some of the species of which have disappeared. The presence of such a type as A . mirabilis in South America, however, does provide another link between the floras of Australasia and South America. A. mirabilis may well have been a member of the parent stock from which the modern " bunya bunya " of Queensland sprang, a stock more widely spread than its descendents ; but in the development of the South American floras it eventually died out, while the species with wingless cone scales persisted there to the present day. Araucarians belonging to the Australasian sections Eutacta or Intermedia, with wide, thin, papery wings on the seed scales, were also represented in earlier South American floras ; perhaps the best evidence for this is provided by Araucaria pichileufensis , described by Berry from Rio Pichileufu in Territory of Rio Negro (Berry, 1938 : 59, pi. n, fig. i). This is a Tertiary species, though its exact age is doubtful (Florin, 1940 : 39), and might suggest that either section Eutacta or Intermedia persisted longer before extinction on the South American continent than did Bunya, though critical records are too scanty for such a generalization to be made at this stage. Florin has renewed the emphasis on the part probably played by Antarctica, whether as an intercontinental land bridge along which migration could take place, or as the place of origin of some of the southern genera (Florin, 1940 : 85-6, 92) : and the distribution of fossil and living species of Araucaria in South America and Austral- asia supports this hypothesis. Pararaucaria patagonica represents an extinct genus and presents a different problem in distribution. The Taxodiaceae, which probably represent the nearest living relations of this genus, are to-day confined to the northern hemisphere, with the exception of Athrotaxis, with three species living in Tasmania. The only other fossil records of this family from the southern hemisphere are three species of Athro- taxis A . ungeri (Halle) from the Mesozoic (possibly Lower Cretaceous) of Southern A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 133 Patagonia, and two other species from the Cretaceous of New Zealand and the Ter- tiary of Tasmania (Halle, 1913 : 40-44 ; Florin, 1940 : 35, 77) and Romeroites argentinensis Speg. (Spegazzini, 1924 : 134-9), f rom Neuquen, South America (? Upper Jurassic or Cretaceous). Florin has, however, drawn attention to the separate and peculiar nature of the genus Athrotaxis, and has suggested that it may represent a distinct, though probably remotely related, line of descent from the Taxo- diaceae. Paramucaria adds, then, to the evidence for an earlier more widespread distribution of taxodiaceous types of plant in the southern hemisphere ; along with Romeroites, it is now extinct, while Athrotaxis has disappeared from South America. Pamraucaria is the only genus related to the Taxodiaceae which shows freedom of bract from fertile scale for the greater part of its length, and so might be regarded as an earlier member of an evolutionary series of forms leading to some of the modern Taxodiaceae where fusion is more or less complete. Such an interpretation would place Pararaucaria in line with that of Florin for the evolution of the coniferalean ovulate strobilus, where he regards the cohesion of the bract with its axillary seed scale complex as a condition derivative from more primitive forms where they are free. However, the Taxodiaceae (sensu Pilger) are an ancient and widespread family in the northern hemisphere ; among living genera, Sciadopitys is already known in the Jurassic, and other genera are abundant in younger rocks, especially in the Tertiary. Hirmer has however (1936 : 65), also included the much older Upper Permian Pseudovoltzia, the Triassic Voltzia and the Rhaeto-Liassic Sweden- borgia in this family : in the two former genera, the bract and ovuliferous scale are more or less free, and in the latter are partially fused. Lack of critical evidence for the exact geological age of Pararaucaria precludes close phyletic comparison with other taxodiaceous genera : but as already indicated (p. 102), the most recent evidence places the age of the petrified forest as at some stage between mid- Jurassic and Wealden. The characters of Pararaucaria itself certainly support the assignation of Mesozoic rather than Tertiary age, for in younger floras the majority of the genera are modern, and Pararaucaria cannot be closely compared with any living genus. A reference made by Wehrfeld (1935) to wood of palms in the Cerro Cuadrado floras has never been substantiated ; and the genus Araucaria, as represented by the extinct A. mirabilis in the same flora, is itself an ancient one, dating back to the Jurassic. It is probable that Pararaucaria represents an extinct offshoot from some common earlier stock of the Taxodiaceae, no doubt of northern origin, which underwent separate evolution south of the equator, as suggested by Florin for the case of Athro- taxis, rather than a type to be regarded as directly ancestral to any of the living genera. The unique one-seeded fertile scale of Pararaucaria is a striking case of homeotic similarity with the condition found in many other conifers of the southern hemisphere Araucaria, Agathis, and the Podocarpaceae, where distribution of the single seed is associated with the entire woody cone-scale, separate winged seed and entire cone-scale with fleshy developments respectively. Biologically, the closest com- parison in this respect with Pararaucaria is with Agathis, where there is also a single winged seed, but with median insertion on a single cone-scale representing the completely fused bract and fertile scale. In the living northern Coniferales (ex- 134 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA eluding Taxales) the cone-scales are normally two to many-seeded, with detachable seeds, though in the taxodiaceous genus Taiwania, which usually has two winged seeds per scale, one of these is frequently missing and the condition is similar to that in Pamraucaria. This modern condition in the northern Coniferales is in contrast with that amongst the oldest northern conifer floras of the Upper Carboni- ferous and Permian, where two of the most widespread genera, Lebachia and Ull- mannia, had one-seeded seed-scale complexes. This condition disappeared relatively early in the history of northern conifers ; though the Araucariaceae are notable exceptions during the Mesozoic in northern lands, they too disappeared from northern floras during early Tertiary times. In any case, this family may have had its origins in the southern continent and have spread northwards, though final proofs of this are lacking (Florin, 1940 : 78-82, 90). However, little is known of the early history of the southern floras ; and in them the one-seeded cone scale may in some cases represent, as seems probable in Pamraucaria, the result of reduction in number of seeds on the fertile shoot, rather than the retention of a primitive character. There is no evidence that the possession of a single seed per seed-scale complex has conferred any special biological advantage amongst southern conifers, except in so far as it has made possible, in the genus Araucaria, the distribution of the single seed still associated with the protective cone-scale tissue in fruit-like fashion, simulating the samara or nut of angiosperms ; a somewhat similar condition is found in Podocarpaceae, in association with fleshy developments. In these two families a condition analogous in some respects with angiospermy has thus been attained, and the corresponding increase in degree of protection of the seed might in some measure account for the success of these two families. But in Agathis and Pararaucaria no such additional protection is associated with the single- seeded habit, though the larger size of the seeds ensured by the reduction in number may have had advantages in germination. CONCLUSIONS According to this interpretation of the remarkable petrified forest of the Cerro Cuadrado region, it is unique in a number of respects amongst other petrified forests. The abundant material of petrified and fertile seed cones of the two dominant types of tree gives a basis for a fairly satisfactory systematic comparison with other living and extinct conifers : the structure of the embryos, for example, and of the vascularization of the cone-scales and ligule in the araucarian cones, are characters which one can hope to demonstrate only very rarely in fossil conifers, and close comparison with living species is impracticable without them. Conclusive evidence of only two species, both based on seed cones, has been found in the forest Arau- caria mirabilis ; and Pararaucaria patagonica, whose affinities are probably with the Taxodiaceae. The detached portions of wood, branches and twigs most probably belonged to one or other of these species, though critical proof of relationship is lacking. The presence of the numerous petrified seedlings affords a unique demon- stration of regeneration, probably of the araucarian species, in the forest. Both the dominant species are extinct, but one, A. mirabilis, probably disappeared in the course of evolution of section Bunya of the genus Araucaria, while the other, A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 135 Paramucaria patagonica, left no descendants, the family of the Taxodiaceae having disappeared from South America without having played a prominent part in its floras so far as may be judged from the scanty fossil evidence. The characters of Pararaucaria in particular suggest an age not younger than Cretaceous for the forest : for no close taxonomic comparison with any living genus can be made. 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Contribution a 1'etude de 1'anatomie comparee du fruit des coniferes. Ann. Sci. nat. hot., Paris (7) 19 : 165-369, pis. 1-15. RIGGS, E. S. 1926. Fossil hunting in Patagonia. Nat. Hist. N.Y., 26 : 536-544. SEWARD, A. C. 1919. Fossil Plants, 4. xvi + 543 pp., 190 figs. Cambridge. & FORD, S. O. 1906. The Araucarieae, Recent and Extinct. Philos. Trans., London (B) 198 : 305-411, pis. 23, 24. SHAW, F. J. F. 1909. The seedling structure of Araucaria Bidwillii. Ann. Bot., London, 23 : 321-334, pi. 21. SPEGAZZINI, C. 1924. Coniferales Fosiles Patagonicas. An. Soc. dent, argent., Buenos Aires, 98 : 125-139, 6 figs. STILES, W. 1908. The anatomy of Saxegothea conspicua Lindl. New Phytol., Cambridge, 7 : 209-222, text-figs. 28-34. TAKHTAJAN, A. L. 1953. Phylogenetic principles of the system of higher plants. Bot. Rev., Lancaster, 19 : 1-45. THOMPSON, D'ARCY W. 1942. On Growth and Form. 2nd ed., 1116 pp., Cambridge. THOMSON, R. B. 1913. On the comparative anatomy and affinities of the Araucarineae. Philos. Trans., London (B) 204 : 1-50, pis. 1-7. TUBEUF, C. VON. 1892. Beitrag zur Kenntnis der Morphologic, Anatomie und Entwicklung des Samenfliigels bei den Abietineen. Ber. hot. Ver. Landshut., 12 : 155-196, pi. 3. WALKOM, A. B. 1921. Mesozoic Floras of New South Wales. I : Fossil Plants from Cocka- butta Mountain and Talbragar. Mem. Geol. Surv. N.S.W., Sydney (Palaeont.) 12 : 1-21, pis. 1-6. WEHRFELD. 1935. Patagonia, el gran acervo de fosiles de la argentina. Rev. geogr. amer., Buenos Aires, 4 : 117-130, i pi. WHITE, C. T. 1947. Notes on two species of Araucaria in New Guinea and a proposed new section of the Genus. /. Arnold Arbor., Lancaster, 28 : 259-260. WIELAND, G. R. 1929. The world's two greatest petrified forests. Science, New York, 69 : 60-63. 1935. The Cerro Cuadrado petrified forest. Publ. Carneg. Instn., Washington, 449 : ix + 1 80 pp., 33 pis. WILDE, M. H., & EAMES, A. J. 1948. The ovule and " seed " of Araucaria Bidwilli, with discussion of the taxonomy of the genus. I : Morphology. Ann. Bot., London (n.s.) 12 : 311-326, pi. 6. 138 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA WILDE, M. H., & EAMES, A. J. 1952. The ovule and "seed" of Araucaria Bidwilli, with discussion of the taxonomy of the genus. II : Taxonomy. Ann Bot., London (n.s.), 16 : 27-47. Pis. 2, 3. WINDHAUSEN, A. 1918. The problem of the Cretaceous-Tertiary boundary in South America and the stratigraphic position of the San Jorge-Formation in Patagonia. Amer. J. Set., New Haven, 45 : 1-53. 1924. Lineas generates de la constitucion geologica de la region situada al oeste del Golfo de San Jorge. Bol. A cad. Cienc. Cordoba, 27 : 167-320. 1931. Geologia Argentina, 2. 645 pp., 214 figs. Buenos Aires. WORSDELL, W. S. 1899. Observations on the vascular system of the female " flowers " of Coniferae. Ann. Bot., London, 13 : 527-548, pi. 27. Note for p. 122. The pith structure in the Pararaucaria cones resembles quite closely that of the stem in the living Taiwania cryptomerioides. Doyle M. H. & Doyle, J. (1948, Proc. R. Irish Acad., 52, B : 26, text-fig. 5) have demonstrated that the pith structure in stems of Taxodiaceae may be used as a diagnostic character for the genera, though corresponding observations for cone axis anatomy are not available.] PRESENTED 2 3 NOV 1953 DESCRIPTION OF PLATES All figures are from untouched photographs. PI. i, figs, i, 4-6, 8-13 ; PI. 2, figs. 14-20 ; PL 3, figs. 26-37 ; PL 4, figs. 38-42 ; PL 5, fig. 46 ; PL 6, figs. 55, 56, 62-64 ' pl - 7. fi g s - 74 75. are taken by reflected light. PL 4, fig. 43 and PL 5, figs. 50-54 are taken from thin sections against a black background. The photographs in PL i, figs. 3, 7 ; PL 5, figs. 48, 49 ; PL 6, fig. 63, and PL 7, figs. 66-73 were taken by Mr. E. Ashby, Department of Cryptogamic Botany, University of Manchester. PLATE i Araucarites sanctaecrucis n. sp. and Coniferous Wood. FIG. i. Leafy (L) and decorticated (w) branches embedded in matrix. V. 30936. x 0-6. Coniferous Wood. FIG. 2. Transverse section of a woody branch, probably 12 years old, showing pith (p], annual rings in the secondary xylem (x), and remains of cortical tissue (? periderm, c). V. 30937. x o'8. FIG. 3. Radial longitudinal section of branch seen in fig. 2, showing pitting on radial walls of tracheids. V. 309376. x 250. Araucarites sanctaecrucis n. sp. FIG. 4. Foliated ? 2-year-old branch, showing lateral branches. V. 30938. x 0-9. FIG. 5. Detail of abaxial leaf surface from specimen showing grooves and ridges (cf. branch seen in fig. 4). V. 30939. x 15. FIG. 6. Partially decorticated ? 3-year-old branch with apparent whorl of ? 2-year-old lateral branches. V. 30940. x o'8. Coniferous Wood. FIG. 7. Detail of pitting shown in fig. 3. V. 309376. x 500. Araucarites sanctaecrucis n. sp. FIG. 8. Transversely cut top end of branch shown in figs. 10, n, with (? 4) annual rings in the secondary xylem (x), and leaf bases (&) and leaf tips (t) transversely cut. V. 30941. X i. FIG. 9. Leafy 2-year-old branch with one-year-old branches arranged in two ranks. V. 30942. x 0-9. FIG. 10. Detail of leaves from the top of the branch shown in figs. 8, n. b leaf base, t = leaf tip. V. 30941. X 3-8. FIG. ii. Leafy branch (? 4-year-old). .30941. X 0-8. FIG. 12. Defoliated branch (? 5-year-old) showing rhomboidal leaf bases. V. 30943. X 0-9. FIG. 13. i-year-old leafy branch. V. 30944. x o'8. Bull. B.M. (N.H.) Geol. 2, 2 PLATE 1 10 13 ARAUCARITES PLATE 2 Seedlings (cf. Araucaria mirabilis). FIG. 14. Broken seedling (S), along with separate fragments of branches and twigs, embedded in matrix. V. 30936. x 0-6. FIGS. 15-17. Broken seedlings of various shapes (see text). V. 3094648. x 0-9. FIG. 18. External view of seedling showing periderm (pd) and woody core (x). V. 30949. X 0-9. FIG. 19. Median longitudinally-cut surface of half seedling shown in fig. 18. x 0-9. FIG. 20. Transversely-cut surface of swollen region of seedling similar to those shown in figs. 17, 18. V. 30950. x 0-9. FIG. 21. Radial longitudinal section of top end of seedling similar to that shown in fig. 15. p = pith. V. 30951^. x o'8. FIG. 22. Transverse section of central swollen region of seedling similar to those shown in figs. 17-20. pd = periderm, x = secondary xylem. V. 3095205. x o'8. FIGS. 23-25. Transverse sections through top, middle and basal end respectively of seedling similar to that shown in fig. 15. p = pith, pr primary xylem, x = secondary xylem (cf. Text-fig, i). V. 3og$ib-d. x 25. Bull. B.M. (N.H.) Geol. 2, 2 PLATE 2 14 20 Araucarian Seedlings PLATE 3 Araucaria mirabilis (Spegazzini). FIG. 26. Petrified cones embedded in matrix. V. 30953. x 0-4 FIG. 27. Cone broken transversely half way up axis, showing pith (p), vascular bundles (v), and cone-scales with seeds (s). V. 39054. x 0-9. FIG. 28. External features of cone shown in fig. 27. / = ligule, a = apophysis of bract. X 0-9. FIG. 29. Detail of external features of cone-scales shown in fig. 28. / = ligule, a = apo- physis of bract, w wing. x 4-1. FIG. 30. Base of cone shown in fig. 35 ; showing transversely cut pedicel (pi), and gradual transition from sterile to fertile scales at base of axis. V. 30955. x 0-8. FIG. 31. Small cone, showing overlapping laminar tips of bracts. V. 30956. x 0-8. FIG. 32. Cone shown in fig. 31, median longitudinal surface, showing wide pith (p) and immature or sterile cone-scales (c). x 0*8. FIG. 33. Detail of cone scales in fig. 32, showing ligule (/) and bract (b) ; no ovules have been formed. x 4*1. FIG. 34. Detail of fig. 31, showing laminar tips of bracts. x 4-1. FIG. 35. Polished longitudinally cut surface of half cone shown in fig. 30. A = cone-scale shown in detail in figs. 36-37 ; B = cone-scale cut in approximately median longitudinal section ; E = embryo in seed, x 0-9. FIG. 36. Detail of fig. 35 at A. / = ligule, a = apophysis of bract (artificially split), S = seed (cut to one side of median plane), l.s. ligular sulcus. x 4-1. FIG. 37. Surface view of half cone-scale shown in section in fig. 36. / ligule, a = apo- physis of bract, x 4-1. Bull. B.M. (N.H.) Geol 2, PLATE 3 s*4>p ~^:-^* ARAUCARIA MJRABILIS PLATE 4 Araucaria mirabilis (Spegazzini). FIG. 38. Half cone in surface view, showing persistent laminar tips of bracts of cone-scales. Structure illustrated in figs. 39-45, and PI. 5, fig. 54. V. 30957. x 0-8. FIG. 39. Detail of fig. 38, showing persistent laminar tips of bracts (t), with longitudinal striation. x 4-1. FIG. 40. Detail of median longitudinally cut face at mid left of half cone in fig. 42, with apophysis of bract (a), persistent laminar tip of bract (t), ligule (/), embryo (e), (A = ? absciss zone.) x 4*1. FIG. 41. Detail of embryo from fig. 42, showing two cotyledons, x 4'i. FIG. 42. Polished median face of half cone shown in fig. 38 ; p = pith, v.b. = vascular bundles of axis ; details of cone scales in figs. 40, 41. x 0-8. FIG. 43. Tangential longitudinal section of cone (cf. figs. 38-42), taken at level of per- sistent laminar tip of bract, a = apophysis of bract, t = laminar tip, v.b.b. = vascular bundles of bract. V. 30957. x 4-1. FIG. 44. Detail of vascular strand in ligule. tr = spiral tracheids. V. 30957. x 208. FIG. 45. Detail of vascular bundle from persistent laminar tip of bract shown in fig. 43. d = ? resin duct, v.b. = vascular bundle, x 55. Bull. B.M. (N.H.) Geol. 2, 2 PLATE 4 42 45 AKAUCARIA MIRABILIS PLATE 5 Araucaria mirabilis (Spegazzini). FIG. 46. Surface view of mature seed cone whose structure is illustrated in figs. 47-53. / = ligule, a apophysis of bract. V. 30958. x 4-1. FIG. 47. Detail of tangential longitudinal section of cone-scale from fig. 50 ; T.S. = testa of seed (stony layer), F = surface tissue of fertile scale, en = endosperm, E = embryo. V. 309580. x 13. FIG. 48. Detail of embryo seen obliquely cut in transverse section of cone in fig. 51 (E) ; co = cotyledon, en endosperm. V. 309586. x 25. FIG. 49. Transverse section of embryo from tangential longitudinal section of cone (cf. fig. 47), showing two cotyledons (co). V. 30958$. x 25. FIG. 50. Tangential longitudinal section of cone, showing cone-scales with seeds, c = cone- scale with wing, S = seed with endosperm and embryo, F = surface tissue of fertile scale. V. 309580. x 4-1. FIG. 51. Transverse section of cone ; T.S. = testa of seed (stony layer), en = endosperm, E = embryo, ss = suspensors. V. 30986. x 4*1. FIG. 52. Radial longitudinal section of cone ; lettering as in fig. 51. V. 3O958c. x 4*1. FIG. 53. Tangential longitudinal section of cone, taken at i mm. from outer surface ; / = ligule with vascular bundles, b = bract with vascular bundles. V. 30958^. x 8. , FIG. 54. Tangential longitudinal section of cone illustrated in PI. 4, taken at level of ligule (/) and showing vascular bundles of ligule (v.b.l.), and vascular bundles of bract (v.b.b.) with ? resiri ducts (d). V. 30957$. x 8. Bull. B.M. (N.H.) Qeol. 2, 2 PLATE 5 54 ARAUCARIA MIRABILIS PLATE 6 Pavaraucaria patagonica Wieland. FIG. 55. Cone showing fertile scales (/) and bracts (6), the surface features weathered. V. 30959. x i. FIG. 56. Cone showing less marked degree of weathering. V. 30960. x i. FIGS. 57-60. Tangential longitudinal sections (cf. also Text-fig. 6) through fertile scales and bracts, taken from base outwards. / = fertile scale, f.s. vascular supply of fertile scale, f.s. I. lateral bundle supplying seed in region of basal foramen, sc = sclerenchyma of fertile scale, b = bract, b.s. vascular supply of bract, T.T. = transfusion tissue, w = wing tissue of testa of seed (cf. fig. 61), t.s. = stony layer of testa of seed, F = basal foramen of seed, E.r. = radicle of embryo, E.G. = cotyledons of embryo, n nucellus, e.s. = embryo sac. Figs. 57-59, V. 3096ia-c. Fig. 60, V. 30960^ x 8. FIG. 61. Stellate sclerotic tissue of wing of seed (cf. fig. 59, w). t.s. = stony layer of testa of seed. V. 30961^. x 50. FIG. 62. Cone with surface features much weathered, showing foliated pedicel (pi.). V. 30962. x i. FIG. 63. Cone showing surface features of fertile scales and bracts, the fertile scales with vertical grooving of protruding abaxial surface. V. 30963. x 0-8. FIG. 64. Detail of leaves on pedicel from cone in fig. 62, showing longitudinally striated abaxial surface. V. 30962. x 3-3. Bull. B.M. (N.H.) Geol. 2, 2 PLATE 6 K v- j^f^ 'V;V 60 PARARAUCARIA PAT AGON 1C A PLATE 7 Pararaucaria patagonica Wieland. FIG. 65. Transverse section of stele of cone axis in lower half of cone, showing narrow pith (p) and thick cylinder of xylem (x), with bract trace at b. V. 30964^. x 11-7. FIGS. 66-72 illustrate structure of xylem of cone axis. V. 30959^. FIG. 66. Radial longitudinal section of transitional region of xylem, showing from right to left the sequence from protoxylem (px) through metaxylem (mx) to secondary xylem. x 166. FIG. 67. Detail of pitting in primary xylem from fig. 66, showing spiral pitted tracheids (s.p.) and reticulate tracheids (ret.). x 417. FIG. 68. Radial longitudinal section of xylem showing, on the right, reticulate tracheids of primary xylem, and on the left, the radial walls of secondary tracheids (sec.) with uniseriate bordered pits. m.r. = medullary ray. x 166. FIG. 69. Detail from fig. 68, showing pits on radial walls of secondary xylem tracheids, and part of medullary ray (m.r.) with cross-field pitting, x 333. FIG. 70. Radial longitudinal section of secondary xylem at base of cone axis, showing uniseriate and biseriate (bi) radial pitting of tracheids, and medullary ray with cross-field pitting (m.r.) x 166. FIG. 71. Detail of biseriate pitting on radial walls of secondary tracheids, from fig. 70. x 333- FIG. 72. Radial longitudinal section of secondary xylem in region of medullary ray, showing cupressoid cross-field pitting (c). (Longitudinal axis of cone runs across the photo.) x 417. FIG. 73. Radial longitudinal section of fertile scale with seed and embryo, seen (reversed) at E in fig. 75, showing orientation of embryo, c = cotyledons, r = tip of radicle, m = closed micropyle. V. 309650 x 8. FIG. 74. External features of long cone, showing weathered surfaces of fertile scales (/) and bracts (b). V. 30965. x 1-5. FIG. 75. Median longitudinally cut surface of cone in fig. 74, showing fertile scales bearing seeds with embryos (E). 5 = axis stele. x 1-5. Bull. B.M. (N.H.) Geol. 2, 2 PLATE 7 PA RA RA UCA RJA PA TA GONICA PRESENTED 2 6 NOV 1953 PRINTED IN GREAT BRITAIN BY A D L A R D AND SON, LIMITED BARTHOLOMEW PRESS, DORKING . _____ . Nov 1g53 'HE SOLUTION OF THE PILTDOWN PROBLEM J. S. WEINER, K. P. OAKLEY AND W. E. LE GROS CLARK BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY Vol. 2 No. 3 LONDON: 1953 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY The following papers appeared in Volume I (1949-52) : Price No. I (1949). The Pterobranch Rhabdopleura in the English Eocene. H. D. Thomas & A. G. Davis j s . 6d. No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P. Oakley & M. F. Ashley Montagu ...... ^s. No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone of the Welsh Borders. E. I. White. Pteraspis leathensis White a Dittonian Zone-Fossil. E. I. White j$. 6d. No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan, Northern Iraq. L. F. Spath ....... IDS. No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal- pellum. T. H. Withers 5$. No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae). T. H. Withers 55. No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian of Podolia. W. N. Croft IDS. No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle East. T. F. Grimsdale ........ IDS. No. 9 (1952). Australian Arthrodires. E. I. White .... 155. No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 65. THE SOLUTION OF THE PILTDOWN PROBLEM BY J. S. WEINER Department of Anatomy, University of Oxford K. P. OAKLEY Department of Geology, British Museum (Natural History) W. E. LE GROS CLARK, F.R.S. Department of Anatomy, University of Oxford Pp. 139-146 ; Pis. 8, 9 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY Vol. 2 No. 3 LONDON: 1953 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and, an Historical Series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 2, No. 3 of the Geological series. PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued November zist, 1953 Price Three Shillings and Sixpence THE SOLUTION OF THE PILTDOWN PROBLEM By J. S. WEINER, K. P. OAKLEY & W. E. LE GROS CLARK NOTE. The curator of a palaeontological collection, which may contain rare specimens of great scientific importance, is frequently faced with the problem of whether to allow such specimens to be reinvestigated by treatment with acids, sectioning, removal of fragments for chemical analysis, or other methods which might seem to involve damage to a unique object. The cautious attitude of a previous generation has undoubtedly preserved for their successors many fossils which, for example, might have been damaged by mechanical treatment in the past, but can now be developed in perfection by more recently devised chemical methods. In the case of the Piltdown relics, one can be certain that after they came into the late Sir Arthur Smith Woodward's possession they would not have been treated or tampered with either chemically or physically. The decision to submit these specimens to a rigorous re- examination was made some years ago ; the final result, unexpected at the time, will be found in the following pages. W. N. EDWARDS, Keeper of Geology. SINCE the report, some forty years ago, 1 of the discovery of several cranial fragments, a portion of a mandible and a canine tooth at Piltdown in Sussex, the problem of the " Piltdown skull " has been the subject of continuous controversy. Some authorities have accepted all the remains as those of an extinct type of hominid. But it is probably true to say that most anthropologists have remained sceptical or frankly puzzled by the contradictions which they present, for (apart altogether from other details) the combination of a cranium closely similar to that of Homo sapiens with a mandible and canine tooth of simian form seemed too incongruous. It has been suggested, indeed, that they really represent the fortuitous association of a Pleistocene human cranium with the remains of a fossil ape which had perhaps been secondarily derived from an earlier geological deposit. The application of the fluorine test (Oakley & Hoskins, 1950), however, made it quite clear that the mandible and canine were certainly not older than the cranium and, on the assumption that they were all genuine fossils, it naturally appeared to lend some support to those who held them to be contemporaneous and to belong to the same individual. It is also a fact that the remarkably flat wear of the molar teeth in the mandibular fragment is quite unlike that normally found in apes at a corresponding stage of dental attrition (though similar to the type of wear characteristic of the hominid dentition), while the mode of wear of the large canine tooth is also different from that which occurs in apes. But there is another possible explanation of the apparent contradictions shown by the Piltdown remains : that the mandible and canine tooth are actually those of a modern ape (chimpanzee or orang) which have been deliberately faked to simulate fossil specimens. It was not till one of us (J. S. W.) in the course of personal discussions put forward this proposition fairly and squarely as the only possible solution of the Piltdown puzzle, pointed out that the organic content of the mandible had never been examined, and moreover demonstrated experimentally that artificial abrasion of the teeth of a chimpanzee combined with appropriate staining produced an appearance astonishingly similar to the Piltdown 1 Dawson & Woodward, 1913. GEOL. ii, 3. 10 142 THE SOLUTION OF THE PILTDOWN PROBLEM molars and canine, that we decided on a critical re-study of all the Piltdown material with this specific possibility directly in view. The results of our in- vestigations have now demonstrated quite clearly that the mandible and canine are indeed deliberate fakes. The evidence 1 for this conclusion is briefly as follows : EVIDENCE OF THE ARTIFICIAL ABRASION OF THE PILTDOWN TEETH Molar Teeth of the Mandible (1) The occlusal surfaces (particularly of M 2 ) are planed down over almost their whole extent to a flatness which is much more even than that normally produced by natural wear (PI. 9, fig. 2). (2) The borders of the flat occlusal surfaces particularly the lateral borders are sharp-cut and show no evidence of the bevelling which is usually produced by natural wear (PI. 9, fig. i). (3) The centre of the talonid basin in M 2 is unworn, and is bounded by a sharp-cut and unbe veiled border of the planed surface of the crown. This appearance would be produced by artificial abrasion but would not be expected in natural wear (PL 9, fig. i). (4) The surface of the areas of dentine exposed on the antero-medial cusps of the two molars is quite flat and flush with the surrounding enamel, instead of forming a depression as would be expected in natural wear. (5) In both molars much more dentine has been exposed on the antero-internal than the antero-external cusps. But in the course of natural attrition the lateral cusps of lower molar teeth are normally worn down more rapidly (and thus usually show a greater exposure of dentine) than the medial cusps (PI. 9, fig. i). (6) The degree of wear in the two molars, M x and M 2 , is almost identical. But in early stages of natural attrition M t is commonly (though not always) more severely worn than M 2 (PI. 9, fig. 3). (7) The planes of the flat occlusal surfaces of the two molars are not congruous, i.e., they do not fit together to form a uniform contour. Unless the teeth have been displaced from their natural position after death (for which there is no evidence), this incongruity is difficult to explain by natural wear (PI. 8; 9, fig. 3). (8) Inspection of the isolated molar tooth (referred to the specimen called Pilt- down II) with a binocular microscope shows that the occlusal surface of the enamel has been finely scratched, as though by an abrasive. Canine Tooth (1) The mode of wear of this tooth is unlike that found normally either in ape or human canines, for the abraded surface has exposed the dentine over the entire lingual surface from medial to distal border and at one point actually reaches the apex of the pulp cavity (Pi. 9, fig. 5). (2) The condition of the apex of the root, and the wide and open pulp cavity seen in an X-ray photograph, indicate fairly certainly that the canine was still incom- pletely erupted or had only just recently completed its eruption. But this would be incompatible with the severe attrition of the crown if the latter were naturally produced (PI. 9, fig. 4). 1 The full evidence will be discussed in detail in a later number of this Bulletin. THE SOLUTION OF THE PILTDOWN PROBLEM 143 (3) X-ray examination shows no evidence of the deposition of secondary dentine (with a constriction of the pulp cavity) which might be expected if the severe abrasion of the lingual surface of the crown were the result of natural attrition (PL 9, fig. 4). (4) The abraded surface of the crown shows fine vertically disposed scratches (as seen under a binocular microscope) which suggest the application of an abrasive (PI. 9> %. 5). EVIDENCE OF THE FLUORINE CONTENT The fluorine method as applied in 1949 (and reported in full in 1950) served well enough to establish that neither the Piltdown cranium nor the mandible was Lower Pleistocene. It did not distinguish (nor at that time was it intended to distinguish) Upper Pleistocene from later material. The rate of fluorination at this site has prob- ably not been high enough to give a clear separation between Upper Pleistocene and, say, Early Post-glacial bones. Moreover the method of analysis used in 1949 was accurate only within rather wide limits when applied to samples weighing less than 10 milligrams, with the consequence that even the difference between the fluorine contents of fossil and modern specimens was obscured where the samples were of that order of magnitude. Improvements in technique have since led to greater accuracy in estimating small amounts of fluorine, and it therefore seemed worth while submitting further samples of the critical Piltdown specimens for analysis in the Government Laboratory. The new estimations, based mainly on larger samples, were made by Mr. C. F. M. Fryd. The following summary of the results leaves no doubt that, whereas the Piltdown cranium may well be Upper Pleistocene as claimed in 1950, the mandible, canine tooth and isolated molar are quite modern. %F o/ p ~V~P~Q : /o^ / O f 2 u 5 Minimum F-content of local U. Pleistocene bones . 0*1 0-4 Ditto. Upper Pleistocene teeth 1 . . . . o i 0-4 Piltdown cranium I. . . . . . o i 0-8 Piltdowncranium.il: frontal . . . . o-i 0-8 Piltdown cranium II : occipital . . . . 0-03 0*2 Piltdown mandible (bone) . . . i . d. No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan, Northern Iraq. L. F. Spath . . . . . . . IDS. No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal- pellum. T. H. Withers 55. No. 6 (1951)- Some Jurassic and Cretaceous Crabs (Prosoponidae). T. H. Withers . . 55. No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian of Podolia. W. N. Croft IDS. l?o. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle East. T. F. Grimsdale 10$. No. 9 (1952). Australian Arthrodires. E. I. White .... 15$. No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 6s. SOME UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT BY MARJORIE E. J. CHANDLER (With Appendices by M. Y. HASSAN and M. I. YOUSSEF) Pp. 147-187 ; Pis. 10-16 ; i Text-figure BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY Vol. 2 No. 4 LONDON: 1954 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series, corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 2, No. 4 of the Geological series. PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued March, 1954 Price Sixteen Shillings SOME UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT By M. E. J. CHANDLER SYNOPSIS The occurrence and preservation of fruits and seeds from the Dano-Montian and Eocene of Egypt is described, and previous work summarized and in some instances revised. These Egyptian fossils are clearly related to the London Clay Eocene plants on the one hand and to the Recent flora of South-Eastern Asia on the other as shown by the occurrence of Nipa in all three floras. The historical and geographical connexion of the three floras is considered, due attention being paid to recent work on plant migration by Chaney and Axelrod. The conclusion is reached that whereas all Tethyan (Indo-Malayan) types of flora may have their remote origin in a uniform palaeotropical plant belt in Early or Middle Cretaceous times, some secondary centre of colonization must be postulated for the Eocene tropical flora of Western Europe where the original palaeotropical angiosperm flora must have been exterminated by the Cenomanian transgression. The obvious source of recolonization lay in South-Eastern Asia, because the great East-West Tethyan ocean must have constituted a formidable barrier to mass migration from the African continent. Moreover the marked Malayan relationship of the London Clay flora supports its immediate Asiatic origin. Detailed systematic descriptions of the Egyptian fruits are given. INTRODUCTION A few fossil fruits recently discovered in Egypt were sent to the British Museum for identification. They are of great importance, for hitherto the bulk of our knowledge of the ancient flora of North Africa has been derived from the study of wood (Krausel, 1939), whereas fruits and seeds were almost unknown. The majority of the specimens came from the Dano-Montian Lower Esna Shales between latitudes 25 and 27 N. They were collected by two Egyptian geologists, Doctors M. Y. Hassan and M. I. Youssef, who in appendices to this paper give details of the deposits which yielded the fruits. (See also note on p. 187). Dr. Hassan's finds were from the Kharga Oasis of the Western Desert, Dr. Youssef's from the Kosseir area bordering the Red Sea. Like the London Clay, with which comparison will be made in the following pages, these plants occur in marine beds with a marine fauna, and must be presumed to have been derived from the nearby land surface of the African continent a little further south. By a curious coincidence a fruit from the Lutetian (or possibly slightly younger beds of Eocene age) of Egypt was also received for study at about the same time. It was collected by members of the Anglo-Egyptian Oil Company, Messrs. Thiebaud and Robson, at Wadi Rayan in the Western Desert, i.e., at about latitude 29 N. in the Fayum. It is an extremely puzzling specimen, and some points about its anatomy are rather obscure. GEOL. II, 4. IZ I 5 o UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT The preservation of these fruits and seeds is interesting and somewhat variable. Some of them, e.g., Palaeowetherellia, are reproduced cell by cell in limonite in part at least. Indeed they display the details of their structure almost as well as the pyritized fruits of the London Clay. They have the merit of being more stable and permanent in their preservation than pyritized specimens, but this advantage carries with it the disadvantage that they do not so readily disintegrate and fall to pieces along structural lines and natural planes of weakness as the London Clay pyritized fossils so frequently do. Some of the best and most delicate evidence from the London Clay fruits came from decaying specimens, not from those which were artificially sectioned. But some of the Egyptian fossils, e.g., Icacinaceae, have the appearance of being purely internal casts, for although they were already chipped, no cell-structure was exposed, or if visible was so indurated as to show little detail. Such specimens are unlikely to provide more information if broken or sectioned, and it therefore seems pointless to sacrifice them where clear and unmistakable evidence of internal structure along natural planes of weakness is lacking. It is highly probable that future collectors will in due course supplement the present meagre evidence from such tantalizing casts. Several of the new Dano-Montian specimens agree with one (.12985) which has been in the collection at the British Museum (Natural History) since 1912. This was described by Krausel (1939 : 106, pi. 2, figs. 9, 10 ; text-fig. 32) and was referred by him to Diospyros schweinfurthi Heer. It came from the Lower Danian of Farafra in the Western Desert at latitude 27 N. approximately. Heer's supposed Diospyros and another specimen which he named Royena desertorum were originally described by him in 1876 (see p. 168). The examination of the new material, and more especially of a newly fractured surface of .12985, leaves no doubt that Heer's " Diospyros " and " Royena " both belong to a single species which has no connection with the Ebenaceae. The species, named hereafter Palaeowetherellia schweinfurthi (Heer), is obviously closely related to Wetherellia variabilis Bowerbank from the London Clay, and to another species W. dixoni (Carruthers) to be redescribed from the Bracklesham Beds of Selsey. The genus is now referred tentatively to the family Euphorbiaceae. Apart from the above, only six Tertiary angiosperm fruits from Egypt have previously been recorded : Palmacites rimosus Heer from the Danian of Kharga (Heer, 1876 : n, pi. i, figs. 21, 22), an imperfect fruit of which no really distinctive characteristics are shown or described. The anastomosing " fibres " in fig. 22 may be nothing more than compression ridges such as are seen in places on the fruit of Icacinicarya youssefi. Heer's determination must be regarded as very doubtful pending re- examination of the specimen, Securidaca tertiaria Engelhardt (1907 : 213, pi. 19, fig. 6) from the Eocene of the Fayum. It would be unwise to comment upon this aceriform fossil without the most careful scrutiny of the original material combined with a comprehensive study of living fruits of this type in the various families in which they occur. Nipadites sickenbergeri Bonnet from the Middle Eocene Nummulitic Limestone UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 15* (Marine Lower Mokattam of Gebel Giuchi, Mokattam), a species which the available evidence identifies with Nipa burtini (Brongniart) (see pp. 159, 162). Rubiaceocarpum markgrafi Krausel from the same beds which is clearly the same Nipa represented by seeds with outer integument preserved (see pp. 161, 163). Nymphaeopsis bachmanni Krausel from the Lower Oligocene of Cairo (Krausel, I 939 : 39 pl- 2 fig s - 2-8 ; pi. 3, fig. 8 ; pi. 21, fig. 6 ; text-fig. 7) has features which suggest that puzzling material has been wrongly interpreted. Further reference is made to the matter on p. 183 when the structure of Thiebaudia rayaniensis is discussed. Teichosperma spadiciflorum Renner (1907 : 217 ; Krausel & Stromer, 1924 : 33, pl. i, fig. 2 ; text-figs. 1-3) from the Lower Oligocene of the Fayum, referred tenta- tively by Renner and with more certainty by Krausel & Stromer to Pandanaceae, needs re-investigation especially as regards the number of locules and form of the seed. If the seed is really curved as Krausel & Stromer's text-fig. 3 shows, the family Myrtaceae should be explored. But before making confident statements about this, it might be necessary to examine serial sections. Without further evidence no really satisfactory determination can be made, although relationship to Pandanaceae seems highly improbable. It is not surprising that fruits from a remoter period of angiosperm history than the Tertiary material hitherto studied are difficult to relate to living genera or to place in Recent plant families. But in spite of the fact that it has not been possible to identify all the specimens, the interest and significance of this tiny flora are out of all proportion to its size. The list of the newly found plants is given below. Family Nipaceae . Anonaceae Euphorbiaceae Euphor- biaceae ? Icacinaceae Flacourti- aceae ? Incertae sedis Horizon and locality Genus and species Nipa burtini (Brongn.) Anonaspermum aegypticum n. sp. Lagenoidea trilocularis Reid & Chandler Lagenoidea bilocularis Reid & Chandler Palaeowetherellia schweinfurthi (Heer) Icacinicarya youssefi n. sp. Icacinicarya sp. ? Thiebaudia rayaniensis n. gen. et sp. Carpolithus hassani n. sp. Carpolithus sp. (Icacinicarya ?) Carpolithus sp. Lower Danian Farafra Dano- Montian Kosseir Lutetian (approx.) Kharga and . Kosseir Kosseir . Wadi Rayan Kharga Kosseir Fuller details of localities are given under the descriptions of the species in the systematic part of this paper. No attempt has been made to determine a few small twigs, 152 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT These discoveries are of outstanding importance for a number of reasons : (1) As stated above, fossil fruits and seeds of any age are rare in Egypt. The recent finds suggest that careful collecting, persisted in over a long period, would reveal the presence of a considerable flora in that region. (2) Very little is known about pre-Tertiary angiosperm fruits and seeds from any area whatsoever. Knowledge derived from leaves or wood is usually comple- mentary to that based on fruits and seeds rather than strictly comparable with it. This was discussed by Reid & Chandler (1926 : 10-13) Plant organs such as leaves tend to be preserved in different deposits from fruits and seeds and to represent different elements in the parent plant-formations. But here in Egypt are data from pre-Tertiary Beds which are really comparable with the abundant Tertiary records of fruiting organs elsewhere. (3) The plants from Egypt are quite obviously related to the Tethyan type of flora found in the London Clay. For even in so small a group as this under dis- cussion similar or identical genera and even species occur. Note the presence of Palaeow ether ellia (a distinct genus and species representing Wetherellia), of a Nipa, of the two species of Lagenoidea actually found in the London Clay (so far as present information can demonstrate), of an Anonaspermum, and of the family Icacinaceae. Here is impressive evidence that the Malayan type of flora so characteristic of the London Clay was already present in Africa towards the end of Cretaceous times. It must be presumed to have flourished on the southern sea-board of the ancient Tethys ocean. (4) The presence in Egypt of a flora of strong tropical Malayan affinity, for such must be the relationship of any flora at all comparable with that of the London Clay, confirms the view supported by other lines of evidence that the climate of Africa to the north of the equatorial belt must have been remarkably different during the late Cretaceous and Eocene from what it is today. Nipa and its asso- ciates demand not only warmth but a high degree of humidity (Reid & Chandler, 1933:74^5^.). The discovery of this Tethyan flora in Egypt at the end of the Cretaceous must inevitably raise the question, " What was its historical and geographical connection with the biologically similar floras in the London Clay in Eocene times and in Indo- Malaya at the present day?" Did the Egyptian flora with the characteristic Nipa and its associates arise in remote times in South-East Asia and spread thence into the African and European continents as climatic conditions, combined with available migration routes, fav- oured its expansion, or had it some other origin ? From what primitive source or sources were these three related floras derived ? Professor Kryshtofovich expressed the opinion that in the tropics of South-East Asia the Malayan type of flora had remained " unmolested ever since its first descent from its Cretaceous ancestors " (1929 : 310, 311). Reid & Chandler (1933 : 82) stated that in their opinion the London Clay flora had its origin in Malaya, whence it migrated northwards and westwards along the shores of the Tethys ocean to Western Europe. The late Professor Seward criticized the views of these three authors on the UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 153 grounds that the early Tertiary flora of South-East Asia is but little known, and that the evidence concerning it, if it exists at all, has not yet been published (1934 : 23), so that there may, for all we know, have been no Tethyan type of flora in Indo- Malaya in the early Tertiary which could have served as a centre of dispersal. He propounded two alternative suggestions as to the origin of the Tethyan flora in Europe : (1) That like the later Tertiary floras it may have reached Europe " not from a South-Eastern home but from the North." (2) That the London Clay flora may have been part of a widespread flora " which in the course of the Tertiary period suffered progressive reduction and is now repre- sented by enfeebled relicts in Indo-Malaya." Before discussing these suggestions it may be of value to digress so as to take account of recent important American contributions to plant history set forth by Professor Chaney in two papers (1940, 1947). Chancy bases his views on successive American fossil floras. These show that there was gradual displacement of temperate to warm-temperate Cretaceous forests in low and middle latitudes in the United States at the end of the Cretaceous period. Their place was taken by warm tem- perate to sub-tropical vegetation (the Neotropical flora of Chaney) of Lower Eocene age derived largely from the Antillean region on the East, and from Mexico and Central America on the West. Evidently, he states, an early Tertiary northward movement of plant populations as a whole occurred, so that the temperate forest type of vegetation became concentrated in high latitudes, while tropical and sub- tropical vegetation, now characteristic of forests near the Equator, spread into middle latitudes, reaching 50 N. approximately on the western side of the continent, and 37 N. on the eastern side. Professor Chaney compares American plant history with that of the Old World, and emphasizes that in approximately the same period a parallel story of plant migration occurs there, modified to some extent by the extensive East-West barrier of the Tethys ocean, which caused some differentiation of vegetation to the North and South of it in Eocene and later times. He makes special mention of the tropical Malayan London Clay flora at latitude 50 N. on the west side of the Eurasian conti- nent whose modern equivalents (notably Nipa and mangrove) grow in the rain- forest of Indo-Malaya. He also lists and shows on a map (1940 : 482, 483, text-fig, i) numerous Eocene and a few Oligocene (palaeotropical) floras of the Old World which have similar tropical relationship, while localities for the corresponding (neotropical) floras in the New World are also plotted. He then recalls that since the Oligocene a reverse movement has taken place in both hemispheres, the sub- tropical and tropical forests gradually moving south again, giving way both in North America and in Eurasia to that temperate vegetation which he calls the Arcto- Tertiary flora. In America the tropical vegetation has survived in the Antilles, the North of South America, Central America and Mexico, with a few remains in South Florida. In Eurasia it survives in south-east Asia. Chaney perceives that such a wide-spread shifting of forest distribution must be due to factors which were world-wide in operation, affecting whole plant populations 154 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT (1947 : 141). If, as appears from the evidence available, he is right that migration from the end of the Cretaceous onwards to the Oligocene was northwards, so that plants from equatorial regions extended into middle latitudes, then the suggestion made by Seward that the Eocene tropical floras, like the later ones, may have reached Europe from the North does not fit into the picture and cannot be seriously enter- chalazo chalaza position of raphc micropylc hilum incurving of Utta to make a canal radicle chalaza chalazo TEXT-FIG, i. A-D Longitudinal sections through fruits showing placentation (diagram- matic). A. Diospyros sp. Recent. B. Wetherellia variabilis Bowerbank. London Clay ; Sheppey . c. Wetherellia dixoni (Carruthers) . Cuisian ? ; Selsey . D Palaeo- wetherellia schweinfurth i (Heer) . Upper Cretaceous ; Egypt. E. Longitudinal section through seed, Diospyros sp. Recent. UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 155 tained. There is certainly no evidence in Europe of a Malayan flora in more northerly stations than the London Basin. Reid & Chandler (1933 : 62) were convinced by its past distribution and present physiological requirements that the Nipa flora in the London Clay was living at the most northerly limit of the climatic conditions (both of temperature and humidity) which it could tolerate. Hence it is improbable that it could have come into being in yet more northerly regions which would have been still more unfavourable in these respects, regions which were, moreover, already occupied by the Arcto-Tertiary type of vegetation suited to them. For these reasons the origin of the Malayan flora to the North appears to be untenable. Before considering Seward's other alternative suggestion, it may be helpful to summarize briefly the facts (and deductions from them) which should be borne in mind and correlated in any further consideration of these problems. They are as follows : (1) A late Cretaceous Tethyan flora in Egypt found at approximately 25 to 30 N. probably derived from the African mainland at no great distance to the South. (2) An early Eocene Tethyan flora in Western Europe extending North to latitude 50 approximately. (3) The presence of similar tropical or sub-tropical floras in the Eocene and Oligo- cene situated apparently along the former northern boundaries of the Tethys ocean, as many of these floras yield Nipa. (4) A living Tethyan type of flora restricted to South-East Asia (the Indo-Malayan flora). (5) A northward extension of the equatorial tropical forest belt between Cretaceous and Oligocene times coinciding with the broadened latitudinal belt of increased temperature and humidity. With these facts and deductions in mind, we now turn to Seward's second alter- native theory, viz., that the Tethyan flora was part of a widespread flora which suffered progressive reduction, surviving in Indo-Malaya today. This is an un- questionably true statement, for on actual fossil evidence, so far as it goes, the flora of Indo-Malaya, or something closely akin to it, was once much more widely distri- buted (Chaney, 1940 : 482-485, text-figs, i, 2 ; Edwards, 1936 : 28, text-fig. 9, map showing the distribution of Nipa in relation to the approximate outlines of the Tethys ocean) . But Seward's statement does not carry the matter far enough ; it requires much more elucidation. Nor does it exclude the possibility of an imme- diate (early Eocene and post-Cretaceous) Malayan origin for the palaeotropical vegetation of Europe. In support of the theory that the various Tethyan floras originated within a uniform tropical belt of vegetation reference may be made to Axelrod (1952). He believes tfrat in the Lower Cretaceous, " long distance migration may have been more effective than at any later time," producing a more or less uniform type of vegetation within given climatic belts. During that period, he points out, the angiosperms were beginning to compete successfully with the older waning Mesozoic floras which they were in process of supplanting. He visualizes that by Middle Cretaceous times there must have been 156 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT basic, more or less world-wide, angiosperm floras with the minimum of regional differentiation, viz., a tropical to sub-tropical flora in lower and middle latitudes more uniform in type throughout the world than has existed at any time since that period, and a uniform temperate flora at higher latitudes. It is not unreasonable to suggest that Axelrod's uniform tropical angiosperm flora was the source from which the Recent tropical flora of South-East Asia and the Cretaceous-Oligocene tropical flora of North Africa were derived by direct and un- broken descent within these two regions. It may also have been the source of the Eocene European tropical flora, not by direct descent in Western Europe but by a more circuitous route. These suggestions are considered and explained in greater detail below. There is no reason to think that in Equatorial Asia there was ever a period during the Tertiary when tropical conditions did not persist. Therefore it is not unreason- able to postulate that the old palaeotropical flora probably survived over the equatorial belt in that part of the world throughout Tertiary times, even after the Oligocene, when it was driven south of its maximum Eocene extension. In that equatorial belt it would have retained or gradually evolved a character of its own, viz., that of the Indo-Malayan flora as we know it today. This is essentially the opinion expressed by Kryshtofovich and quoted on p. 152. In Africa the course of events may have been somewhat similar. Following upon the contraction of the Tethys sea in Oligocene and later times and the resulting climatic changes, the Tethyan flora must have been driven from the more northerly part of its former Cretaceous and Eocene territory by the pressure of unfavourable conditions. But it could probably have survived in the tropics of Central Africa. Desiccation combined with the southern trend of plant migration in post-Oligocene times would undoubtedly have prevented any later return to the more northerly latitudes it had previously occupied. Deserts and the Tethyan sea (or its shrunken remains) would probably have cut it off effectively from Asia. But within the African equatorial belt it may have persisted, possibly giving rise (or giving place ?) eventually to a distinctive African tropical vegetation as it evolved in isolation from the Asiatic stream of life. But what of the London Clay Eocene flora of Britain, and how was it related to the uniform palaeotropical flora of Lower Cretaceous times ? There can be no doubt that the Lower Cretaceous flora of this country must have been eliminated by the great marine transgression of the Upper Cretaceous. This transgression far overpassed the most northerly limits of tropical climate for which there is any evidence, viz., about latitude 50 N. If the European Lower Cretaceous palaeotropical flora persisted at all it must have been on islands, or on tracts of land on the western borders of the present European continent, perhaps on land margins now submerged beneath the sea. If Chaney is right about the world-wide character of climatic changes, the flora could not, in all probability, have survived unless land nearer the Equator was accessible for colonization during the late Cretaceous. For in America, as we have seen, temperate and warm-temperate forests occupied low and middle latitudes at this period, from which latitudes they were driven only at the end of the Cretaceous by the northward march of Equatorial vegetation. It UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 157 seems obvious that with the withdrawal of the Upper Cretaceous sea the re-exposed land surface (now having a sub-tropical or tropical climate) would have been available for recolonization by a flora of tropical type. This may have come either from southern and western plant " reserves " as suggested above, or from the only two alternative sources, viz., the African continent, or the south-east of Asia. In the former we know that a Tethyan flora was present in the late Cretaceous. In the latter there is no reason to doubt the existence of such a flora at that time as has been pointed out above. The theory of recolonization from the west offers no adequate explanation of the marked Indo-Malayan aspect of the incoming Eocene flora. It is unlikely that closely parallel development would have taken place independently in two such widely separated areas. It is equally improbable that the source of recolonization was the African continent. Had no serious physical barrier existed, the natural solution of the problem would be to assume that by Eocene times the Cretaceous Tethyan flora of Africa had migrated northwards and spread into Europe, as it followed the extension in latitude of the hot and humid climatic belt favourable to it. But the existence of the great ocean barrier of the Tethys over perhaps 1,000 miles or more from South to North must have had a marked deterrent effect. Axelrod (1952) discusses conditions affecting the migration of faunas and floras in geological times. He maintains that plant distribution at any particular time is " largely a reflection of the climates available for occupancy " (p. 177), i.e., climate is the chief factor controlling plant migration. But whereas he considers that a much higher degree of probability exists for plants than for animals in the matter of over- water dispersal, he is also sure that " within any given climatic region distance will finally impose a barrier to the migration of plants also." It is only fair to him to state that in spite of this, he regarded the presence of palaeotropical Tertiary vegetation on both sides of the Tethys as evidence that the sea in this particular case (as well as in others) did not prevent the distribution of the flora on both sides of it (1952 : 187). But this is a curious conclusion to have drawn, for he also explains that " The water barrier that a continental flora can transgress within an epoch of time (say, 10 to 15 million years) is not unlimited," while beyond 200 to 250 miles the probability of colonization by a whole flora is low. Hence what he designates the " waif assemblages " of ocean islands which lack balance, since whole classes and orders expected are absent, while others are only poorly represented. " It would appear," Axelrod writes, " that long-distance, over-water migration has not been generally effective in populating widely separated continents." Even during much longer periods of time, therefore, there may be good reason to doubt whether the Tethyan flora in Africa could have migrated directly across the great ocean which lay athwart its path to the North. Further, as Reid & Chandler (1933) pointed out, the London Clay flora shows far less affinity with the flora of Africa than with that of South-East Asia. To the present writer, therefore, it seems highly improbable that wholesale migra- tion took place across the Tethys. There remains the third suggested source for the post-Cretaceous colonization of 158 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT West Europe, namely the South-East Asiatic. On such evidence as is available this appears to be by far the most probable source for these reasons : (a) The Indo-Malayan region and Southern England both lay within the palaeo- tropical climatic belt of Eocene times. (b) The northern shores of the Tethys and the land adjoining were so situated as to provide a possible migration route from South-East Asia to West Europe within this greatly extended tropical belt of Eocene times which allowed the flora of equa- torial regions to occupy many degrees of latitude North of its present limits. (c) The London Clay flora is more closely linked with that of Indo-Malaya than with that of any other existing flora. The final answer to these speculations can only be provided by extensive re- search on fossil floras from many widespread regions. At present the evidence is insufficient to furnish incontrovertible proof, but the balance of probability seems in favour of a remote origin of the Tethyan type of flora within a uniform palaeo- tropical belt in the early Cretaceous, and of an immediate origin of the London Clay flora by migration from Indo-Malaya between Cretaceous and early Eocene times. The subsequent history of the London Clay flora and the reasons why it survived only in Indo-Malaya have been set forth by C. & E. M. Reid (1915), Reid & Chandler (1933), Chaney (1940, 1947) and others. When during the Oligocene the connection of the Tethys with the Indian Ocean was broken so that the former vast waterway became a huge land-locked Mediterranean, while the great transcontinental mountain barriers of Eurasia were being uplifted, progressive cooling of the climate in middle latitudes occurred, so that more temperate plants from the North gradually supplan- ted the former tropical ones of the Eocene. During this period final extermination was the fate of the Tethyan flora in Europe. With the shrinking of the Tethys a migration route to Asia no longer existed under favourable climatic conditions. The direct route to the tropics in the South was blocked by the combined barriers of East-West mountains and the East-West remains of the Tethys, as effectively as Northward migration had formerly been blocked by the vaster Tethyan ocean of late Cretaceous and Eocene times. But in South-East Asia, as already stated, survival of the palaeotropical flora and its descendants was possible, for there is no reason to think that equatorial latitudes in that continent have ever passed through a phase when it could not have retained its tropical flora. Even when the more northerly parts of the palaeo- tropical belt became cooler, so that the tropical flora could no longer live in them, migration to more southerly regions was possible, thanks to the absence of East- West barriers and to the existence of North-South valleys and coastal plains. Meanwhile it is likely that the Tethyan flora of Egypt, when more fully known, will reveal a far stronger African element than is found in the London Clay, cut off as this latter appears to have been from direct communication with the African continent throughout its history. The affinities of unknown plants in the Egyptian fossil flora should therefore be sought among tropical and sub-tropical African genera of the present day as well as among those of South East Asia. UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 159 I wish to express my warmest thanks to Mr. W. N. Edwards for helpful suggestions and criticisms, and to Mr. F. M. Wonnacott of the Geological Department for much help in the preparation of the manuscript for the press. SYSTEMATIC DESCRIPTIONS Note. The specimens, with two exceptions, are from the Dano-Montian Lower Esna Shales of Egypt. Those from Kosseir Area, Red Sea, were collected by Dr. M. I. Youssef, those from Kharga Oasis by Dr. M. Y. Hassan, to both of whom I am indebted for the opportunity to study these most interesting plants and for the generous gift to the British Museum of the figured specimens. The other two horizons and localities represented are the Lower Danian of Farafra (Palaeowetherellia) , and the Eocene beds of Wadi Rayan (Thiebaudia rayaniensis gen. et sp. nov.). MONOCOTYLEDONES Family NIPACEAE Genus NIP A Thunberg 1782 Nipa burtini (Brongniart) (PI. 10, figs. 1-5 ; pi. n, figs. 7-9) 1904. Nipadites sickenbergeri Bonnet, p. 499, figs, on pp. 500, 501. 1924. Nipadites sickenbergeri Bonnet : Krausel, p. 36. 1939. Nipadites sickenbergeri Bonnet : Krausel, p. 22, pi. I, figs. 1-18, text-fig. I. !939- Rubiaceocarpum markgrafi Krausel, p. 108, pi. i, figs. 19-24. For full synonymy see Reid & Chandler (1933 : 118). LOCALITIES AND HORIZONS. North slopes of Mokattam, near Cairo ; Middle Eocene. Gebel Atshan, Kosseir Area, Red Sea ; Dano-Montian Lower Esna Shales. DESCRIPTION AND REMARKS. The discovery of a small Nipa. fruit in the Dano- Montian of Kosseir and the re-investigation of fruits named Rubiaceocarpum mark- grafi Krausel from the Middle Eocene of Mokattam made it necessary to re-study Nipadites sickenbergeri Bonnet from the same beds as Rubiaceocarpum. As a result it can be stated with confidence that Rubiaceocarpum is a Nipa seed with outer integument preserved in part at least, and that it, and the seeds named Nipadites sickenbergeri, cannot be separated from Nipa burtini (Brongn.) in the present state of our knowledge. This conclusion was forced by the evidence upon the writer who had started with the conviction that N. sickenbergeri was distinguished beyond question from Nipa burtini by its ribbed and furrowed seed. Moreover the opinion expressed by Reid & Chandler (1933 : 122) that the fossils should be referred to Nipa rather than to Nipadites is now re-affirmed. 160 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT In order the better to understand the fossils a further examination of the living Nipa fruticans was carried out. The following details may now be added to those recorded by Reid & Chandler (1933 : 122). The hard compact endocarp when freed from the adherent pericarp (or sarcocarp) is more or less smooth but shows on its outer surface, in the parenchyma which covers it, impressions of stout longitudinal fibres. There is no conspicuous fluting or ribbing of this surface, but a tangled mass of finer fibres can be seen where the parenchyma is rubbed off. When abrasion is carried a stage further stout transverse fibres are exposed. These were observed in 1933. The locule shows conspicuous flattened transverse or oblique fibres and stout longitudinal ones overlying them which readily break away. When an endocarp is cut transversely, a nut-like seed may fall free if the material is in a suitable state of preservation. This bears upon its surface conspicuous flutings due to narrow deep furrows in a parenchymatous layer. In the furrows there are remains of fibres which probably belong to the endocarp. They have usually been torn away, leaving empty furrows only on the surface of the seed. Some of the fibres branch or unite so that the furrows and intervening ridges are not aways absolutely straight and longitudinal, although this is their general alignment. On the flat or slightly convex broad ridges between the furrows can be seen impressions of the transverse or oblique flattened fibres which belong to the locule wall described above (PI. n, fig. 10). In Reid & Chandler's description this coat of the testa was regarded as part of the endocarp because endocarp and testa are often fused. On the detached seed there is a deep narrow parallel-sided furrow (PL n, fig. 12). This is formed by an incomplete longitudinal ridge or septum which projects into the locule from the whole length of the carpel wall. The large sub-circular or circular basal aperture giving access to a cavity in the endosperm is a conspicuous feature also, and is associated with the embryo. The outer pulpy integument of the testa is fairly easily scraped away, exposing a close-textured rather brittle, gummy or resinous inner integument with a smooth transversely striate outer surface of fine cells. Embedded in the surface of this coat are a few flat fibres which branch and anas- tomose to form a coarse network. The fibres lie obliquely, transversely, or more infrequently longitudinally (PI. 10, fig. 6). On the inner side of the brittle inner integument is a raised network due to very fine fibres so arranged as to form fine transversely elongate meshes. The inner integument follows fairly closely the contours of the endosperm, but is separated from it by a thin soft film of fine cells. When the endosperm has rotted, as may happen in specimens which have drifted in the sea for some time, the empty integuments of the seed may still remain attached firmly to the endocarp as it lies within the drupe, but the film of cells just described has usually gone so that only the network of fibres with transverse meshes covers the exposed surface. It is the internal cast of this coat with its fibres which shows the ornamentation described on seed-casts of Nipa burtini from the London Clay (Reid & Chandler, 1933, pi. 2, fig. 6). No raphe or chalaza scar is visible on the interior of the inner integument in UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 161 Recent fruits, nor is their existence indicated by any change in the alignment of the fine fibrous network so far as I have been able to observe. When all the integuments are removed so that the actual endosperm is uncovered, a series of even finer furrows and ridges, like minute ruminations, are laid bare. At the apex and over the sides of the endosperm their orientation is clearly transverse (PI. 10, fig. 6 ; PI. n, fig. n). At the base they are coarser and the alignment more irregular, giving rise to small low rounded lobes of endosperm. Whatever the orientation, the small areas which they produce on the surface are more or less convex, especially when slightly weath- ered or abraded. The endosperm sometimes displays a few broad shallow longi- tudinal furrows or sinuosities at the lower end which gradually die out above (PI. II, fig. 12). They resemble the furrows sometimes seen in Nipa burtini and Nipadites sickenbergeri. On the exterior of the complete seed they are barely discernible, being obscured by the spongy outer integument. Nor are they usually apparent on the inner surface of the empty testa. The astute observer Hooker in an editorial note (Le Maout & Decaisne, 1876 : 822) comments that the seeds are erect and anatropous. But for this observation I should not have suspected their anatropous character, for there is no more indi- cation of a raphe on the outer surface of the testa than there is on the inner surface. Nevertheless examination of a detached endosperm-mass supports Hooker's state- ment, or at least gives evidence of comparable structures. After removal of the integuments a flat band of fibres was seen lying in one of the shallow longitudinal furrows of the surface. It extends upwards from the base almost throughout the length, follows a straight course, and is parallel-sided, quite different in character from the somewhat sinuous flat fibres in other furrows. That this is a definite structural feature, probably the raphe, is confirmed by the differentiation of a band of endosperm immediately beneath it. In this position there are small irregular rounded " ruminations," not transversely elongate ones as elsewhere, and some indication of longitudinal alignment is given by a tendency for longitudinal cracks to form (PI. n, fig. n). Somewhere in this band the fibres must enter the endo- sperm and pass to the chalaza. But the point of entry is so inconspicuous that it is hidden by the irregularities of the " ruminate " surface. Nevertheless a correspond- ing structure shows clearly in some fossil material, as will be described later. From the details given above it is fairly clear that the appearance of a fossil Nipa must vary considerably with the coat which happens to have been preserved or exposed. This depends partly on the mode of preservation of the fossil. Thus an internal seed-cast may show the structure of the endosperm as in the Egyptian Nipa, or it may merely be a cast of the inner integument formed after the endosperm had decayed as in many London Clay seeds. Any seed having the ridged outer integument preserved would have the dis- tinctive appearance of RuUaceocarpum, further details of which are given later. In a broken fruit some layer or layers of the endocarp may show, as in many fruits from the London Clay, and in a small Dano-Montian fruit from Egypt. In the London Clay fruits wear and tear has often exposed the inner layers of endocarp with flat transverse fibres described in 1933. By far the most common specimens at Sheppey and Herne Bay are fruits (perfect or imperfect), showing stout fibres i6z UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT embedded in the spongy tissue of the sarcocarp. Rarely is the epicarp preserved, and then only in fruits freshly removed from the matrix. It quickly cracks and falls to pieces on exposure to air. . It seems scarcely necessary to repeat detailed descriptions of the fossil fruits and seeds which have already appeared in published work (Reid & Chandler, 1933 : 119 ; Krausel, 1939 : 22 ; Seward & Arber, 1903, etc.). Other references will be found in the synonyms listed by Reid & Chandler (1933 : 118). The question inevitably arises once more, " How many fossil species can be dis- tinguished in the deposits with which we are now concerned ?" It must be remembered that whereas the London Clay fruits and seeds are rela- tively uncrushed, the Egyptian ones have often undergone an excessive amount of deformation by crushing and there appear to be no valid and consistent grounds for separating Nipa burtini and Nipadites sickenbergeri as distinct species. The longi- tudinal ridging and furrowing of the seed-casts, or its absence, although it was supposed by Bonnet to be of specific value, cannot really be used in diagnosis, for although smooth seed-casts are common in the London Clay, furrowed ones may also occur. Moreover both kinds are found in the Egyptian Eocene (Bowerbank, 1840, pi. 4, fig. i; pi. 5, fig. i ; Krausel, 1939, pi. i, figs. 11-13). Again in the Egyptian casts the furrows may be few and slight (V. 13695) or fairly conspicuous, or in some cases, overemphasized by crushing. The other chief difference which occurs in some cases between fossils from the two localities lies in the size. But differences in size are no more satisfactory than furrowing, as a reason for specific separation. It is true that the majority of Nipa burtini fruits and seeds from the London Basin are smaller than the majority of the seeds from Egypt, and that they vary greatly in Appearance. Yet large nuts do occur in the London Clay (Bowerbank, 1840, pi. 5, fig. i), while very large fruits and seeds occur in the Belgian strata (Seward & Arber, 1903). On the other hand the small fruits from the London Clay are indistinguishable from a Dano-Montian fruit from Kosseir (PL 10, figs, i, 2). Perhaps the predominance of small forms in one area and large in another may be due to the sorting effects of specific gravity rather than to systematic differences. The associated fruits of other families at Sheppey and Herne Bay are on the whole fairly small. . The following table shows minimum and maximum sizes of fossil Nipas from previously published records. It must be remembered, however, that a far larger number of specimens are known from the London Clay than from the other deposits and they include many immature or abortive fruits. London Clay Belgian Eocene Egyptian Eocene Fruits : Length, i to 18 cm. . 7-5 to 21 cm. Not known Breadth, i -3 to 12 cm. . 3 to 15 cm. . Seeds : Length, 2 to 8-9 cm. . 9 to 10 cm. (commonly 8-5 . 7-5 to n cm. (commonly 8-5 to 9 cm.) or 9 cm.) Breadth, 1-3 to 7 cm. . 9 cm. (commonly about 7 . 3-4 to 12-3 cm. (commonly cm.) about 5 to 7 cm.) UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 163 Thus whether furrowing or size be considered, there appear to be no sharp lines of demarcation between species from the localities with which we are concerned. If furrowing and size cannot be used as a basis for separating species and it is difficult to see how they can when every gradation may occur in these characters there are no other grounds obvious on which the diagnosis of more than one species can be based. Certainly the differences between the Nipas from the two areas under review are no greater than the differences among individual specimens in the London Clay itself (cf. Bowerbank, 1840, pi. 4, fig. i ; pi. 5, fig. i). Yet there is no reason whatso- ever for creating more than one species in this deposit. A few additional observations on the Egyptian nuts may not be out of place here. Krausel's figured specimens are largely seed-casts and might be expected to show endosperm structure. Unfortunately the small scale of the figures does not display such fine details except in his pi. i, fig. 8, where transversely aligned endosperm cells are clearly seen around the basal scar. Endosperm structure is also clearly visible in two casts from Mokattam in the British Museum (V. 13240 and .13695 ; PI. n, figs. 7, 8). The variable and considerable amount of compression of the seeds must be stressed, for some are almost reduced to thick concavo-convex lenticles (.13240). Inevitably this crushing increases the diameter. .13239 is obliquely distorted, the basal aperture having been forced into a basi-lateral position (PI. 10, fig. 3). As in the living, so also in the fossils, endosperm may show features not dis- cernible on the interior of the testa represented by internal casts. .13240, in addition to the typical transverse pattern of endosperm ridges (PL n, fig. 8), shows on its flatter surface a band of elongate, longitudinally aligned endosperm cells which by comparison with the living have structural significance, probably indicating the position of the raphe. Towards, but well below, the apex a radial arrangement of the endosperm on this band suggests the point of entry into the endosperm of the fibres from the raphe (PI. n, fig. 7). On the more convex surface of this specimen, the coarse network of fibres seen in the living between the two integuments of the testa are faintly impressed (PL 10, fig. 5). .13239 has a mosaic pattern all over the surface. This may be the impression of a cracked testa which has peeled. But the important feature of this specimen is that sub-apically the cast shows a deep, small, funnel-shaped opening towards which fibre-impressions converge, and into which they pass (PL 10, fig. 4). Here the entry of the raphe fibres is clearly indicated. .13695 shows a similar but more obscurely preserved radiating structure. Thus these specimens illustrate a well-established fact, that owing to the macera- tion that fossils have frequently undergone, they may display structures which are more difficult to demonstrate in Recent material. We must now return to the specimens named Rubiaceocarpum markgrafi Krausel. As already stated on p. 159 these appear to be seeds (abnormally short and broad) with outer integument preserved. They have undergone considerable lateral com- pression. Their form can be matched among the Nipa seed-casts illustrated by Krausel GEOL. II, 4. I2 164 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT (1939, pi. i, cf. figs. 8, 21 ; 18, 24 ; 16, 24 ; 17, 20). I have been able to examine the specimen illustrated in Krausel's fig. 24, which shows the base of the seed with its large aperture. A very interesting feature is a broad flat fibre band which half encircles the seed in the plane of symmetry (PI. n, fig. g/). This resembles the raphe fibre described above in Nipa fruticans, although seed and fibre are on a larger scale. The fact that it is visible at the surface of the specimen must mean that some measure of abrasion had removed part of the spongy parenchyma. The superficial impressions of the transverse fibres of the endocarp have also been des- troyed at the same time, traces only of them being visible in a few of the deeper furrows of the outer integument. We may ask what relationship does this ribbed entity bear to its living analogue? It appears to be an internal cast of a hollow mould left by a seed which subse- quently decayed. The filling of the hollow with limestone resulted in the characters impressed on the mould being reproduced on a cast, much as sealing-wax poured into the hollow mould of any fossil will reproduce its solid form. An actual replace- ment of the seed itself cell by cell would have given a far sharper representation of it, and in places at least would have provided evidence of cell-structure. The ribbed seed shown by Krausel (1939, pi. i, figs. 20, 21) perhaps retained even more of the outer integument, for there are suggestions of transverse fibre impres- sions at the base of fig. 21, and the raphe band appears to be only partly uncovered. The longitudinal striation on the ridges in Krausel's pi. i, fig. i, may indicate that part of the outer integument of the testa is still present on this cast, which he referred to Nipadites. But suggestions such as this can only be verified by examination of the specimens themselves. Reid & Chandler (1933 : 122) gave reasons for referring the London Clay fossils to Nipa rather than to Nipadites. They appear still to hold good. The most out- standing difference between living and fossil is the presence of the short longitudinal septum, which by partial subdivision of the locule in N. fruticans produces the only constant deep and conspicuous longitudinal furrow on the seed. This septum and furrow are absent in all the fossils. Nevertheless so closely identical with the living is the structure of the fossils in all other respects (even in smallest details) that the presence or absence of the septum still appears to the writer to have specific rather than generic value. Probably however, the real meaning of this character can only be decided by a detailed study of the developing ovary. It may, or may not, have a connection with the suppression of two out of the three original locules or ovules. In the meanwhile it is suggested that the name Nipa be used for the fossils. UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 165 DICOTYLEDONES Family ANONACEAE Genus ANONASPERMUM Ball emend Reid & Chandler, 1933 Anonaspermum aegypticum n. sp. (PL n, figs. 13-15) DIAGNOSIS. Seed-cast oval or oboval in outline, much compressed with slight median depression ; rumination ridges close and narrow, extending from the depres- sion to the margins, diverging from the depression at the distal end, occasionally forking near the margin, with a few short intermediate ridges. Four-partite in transverse section. Thickness only about half the greatest diameter in the plane of symmetry (15-5 mm.). HOLOTYPE. A seed-cast, distal end, with testa almost entirely abraded (PL n, figs. 13-15). Brit. Mus. (N.H.), No. .31106. LOCALITY AND HORIZON. Gebel Atshan, Kosseir Area, Red Sea ; Dano-Montian Lower Esna Shales. DESCRIPTION. The unmistakable internal cast of a seed of Anonaceae showing typical ruminate albumen and marginal encircling band of raphe and chalaza (PL n, figs. 13, 14). The hilar half of the seed is missing. The surviving fragment represents the distal end, which appears to have been severed from the proximal only a short distance above the middle of the specimen. The cast was originally oval or oboval in outline, bisymmetric, somewhat compressed at right angles to the plane of symmetry, anatropous with encircling raphe in the plane of symmetry. The raphe begins to broaden on one side where it merges into the linear chalaza (PL n, figs. 13, 14 ch.}. It completely fills the slight marginal groove at the edge of the albumen. Albumen having a median depression on each broad surface, ruminations forming narrow close ridges from the margin of the seed to the median depression. The ridges diverge from the end of the depression to the apex or distal end of the seed. The inner ends of opposite ridges do not unite at this point to form a series of curves as in Anonaspermum commune Reid & Chandler (1933 : 184, pi. 5, figs. 14-17). Short ridges of varying length sometimes arise at the margin and are interposed between the longer ones. Occasionally one of the longer ridges may divide into two towards the margin of the seed. The transverse section exposed on the fractured surface shows a four-partite arrangement of the albumen (PL n, fig. 15). In the slight depressions in the middle of the broad surfaces part of the fibrous layer of testa is preserved (PL n, fig. 13). On the four- partite sectioned surface oblong cells arranged in transverse rows at right angles to the plane of symmetry of the seed can be seen. The rows are about 0-028 mm. broad. The cells are partially obscured by sinuous fine striations lying parallel in a general way with the plane of symmetry. Parts of the ruminations as seen on the fractured 166 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT surface of the seed are obscured by a thin brown mineral film which bears the impression of a thin but rough coat, the cells of which lie at right angles to the plane of symmetry. It represents the filmy inner integument which penetrated between the plates of albumen as in living seeds. Actual length of seed preserved, 10 mm. ; estimated length of complete seed, about 22 mm. Maximum diameter actually preserved in plane of symmetry, 14 mm. ; probable measurement in the complete seed, about 15 to 15-5 mm. Dia- meter at right angles to plane of symmetry, 7 (at the centre) to 7-5 mm. (at each side). REMARKS. The seed was clearly larger than any described from the London Clay, about twice the size of the largest there recorded. It was also relatively thinner at right angles to the plane of symmetry than any London Clay species with albumen forming superficial ridges. Similar divergence of the ruminations at the distal end is seen in A. rugosum and A. pulchrum Reid & Chandler (1933 : 186, 187, pi. 5, figs. 21-27), but not in A. commune and A. rotundatum Reid & Chandler (1933 : 184-187, pi. 5, figs. 14-20). Although imperfect, this solitary specimen appears to be sufficiently distinctive to merit a specific name. It is described as Anonaspermum aegypticum. The family Anonaceae is .almost exclusively tropical today, occurring in both hemispheres. It is recorded from both hemispheres in Eocene times also. Family EUPHORBIACEAE Genus LAGENOIDEA Reid & Chandler emend. Since the first description of this genus was published (Reid & Chandler, 1933 : 493-497) a few small but important new facts have come to light which make it necessary to correct the diagnoses and descriptions then given. The information was derived from additional London Clay material of Lagenoidea trilocularis. The diagnosis should now read : Fruit a superior, syncarpous, loculicidal and septicidal capsule, two to four- loculed, locules single-seeded. Pericarp thick, formed of radially aligned cells, seeds pendulous, radially compressed, anatropous, raphe ventral, chalaza basi- ventral. TYPE SPECIES. L. trilocularis Reid & Chandler. The recent discovery that the seeds are pendulous with ventral raphe now makes it possible to refer the genus Lagenoidea to the family Euphorbiaceae. Formerly such a relationship was considered impossible, for the seeds were then believed to be erect and orthotropous owing to the fact that the raphe is slender and difficult to detect when it is preserved at all. The shape of the fruit and the combined loculicidal and septicidal dehiscence is typical of the Euphorbiaceae. This was recognized by Ettingshausen when he labelled one specimen (.23129) Euphorbiophyllum eocenicum (see Reid & Chandler, 1933 : 495, pi. 29, fig. 6). This specimen was never described or figured as such, hence the inappropriate name Euphorbiophyllum need not be retained for these UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 167 fruits. It has not yet been possible to connect them with a living genus, but the relationship must be sought among those sections of the Euphorbiaceae with single- seeded locules having locule-linings formed of complicated interlocking cells. Attenu- ated obovate seeds are rare in the family but obovate seeds may occur, e.g., in Chaetocarpus where the seeds may remain attached to the inner angle of the carpel wall when the external valves have fallen away. Lagenoidea trilocularis Reid & Chandler (PI. 12, figS. l6-20) 1933. Lagenoidea trilocularis Reid & Chandler, p. 493, pi. 29, figs. 1-18. The revised diagnosis of this species based on London Clay material as stated above should read : Fruit : Sub-globular, three- (rarely four-) loculed ; capsule splitting loculicidally and septicidally into six (rarely eight) segments. Length, 7 to 14 mm. ; diameter, 8 to 23 mm. ; length of locule-cast (= seed), 3-5 to 4-5 mm. ; greatest diameter, 2 mm. ; least diameter, i mm. LOCALITY AND HORIZON. Gebel Atshan and Gebel Durvi, Kosseir Area, Red Sea ; Dano-Montian Lower Esna Shales. MATERIAL. Two fruits, one with hollow interior. Brit. Mus. (N.H.), Nos. .31107-08. DESCRIPTION AND REMARKS. Two fruits, neither of which show internal structure. One (PI. 12, figs. 16-18) shows a thin wrinkled epicarp partly hiding the surface of the capsule, but the six segments can be clearly distinguished. A sub-circular basal scar indicates the former extent of the calyx (PI. 12, fig. 17). Length of fruit, 10 mm. ; transverse diameter, 9 by n mm. The second fruit (PI. 12, figs. 19, 20) also shows the six valves and basal scar, but its surface is somewhat encrusted and is therefore rather obscure. The interior is hollow, while there is a hole at the attachment through which the seeds may be presumed to have escaped. Length of fruit, n mm. ; diameter, 14 by 12-5 mm. Lagenoidea bilocularis Reid & Chandler (PL 12, figs. 21-23) 1933. Lagenoidea bilocularis Reid & Chandler, p. 496, pi. 29, figs. 19-27. LOCALITY AND HORIZON. Gebel Atshan, Kosseir Area, Red Sea. Dano-Montian Lower Esna Shales. MATERIAL. Two typical fruits. Brit. Mus. (N.H.), No. .31109. DESCRIPTION. Two sub-ovoid somewhat laterally compressed fruits showing six external longitudinal grooves delimiting the valves and septum. The narrow median opposite segments between the pairs of lateral segments mark the edges of 168 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT the septum. The pairs of end segments are bounded by planes of loculicidal and septicidal splitting. As in the London Clay fruits of this species the pairs of valves may be presumed to overlie the two locules. The stylar scar is seen at the narrow end in both fruits and the attachment scar is preserved at the broad end in the smaller specimen. Unfortunately the internal structure is not exposed. There can be no doubt however that the fruits are identical with the London Clay species Lagenoidea bilocularis Reid & Chandler. Length of the two endocarps, n and 9-5 mm. ; greatest diameters, n and 9 mm. ; least diameters, 6 and 5-5 mm. respectively. Family EUPHORBIACEAE ? Genus PALAEOWETHERELLIA nov. DIAGNOSIS. Fruit sub-globular, syncarpous, with more than five carpels ; dehis- cence loculicidal and septicidal, locules radially arranged, tangentially compressed ; placentation axile ; seeds solitary in each locule, occupying part only of the plane of the locule ; pendulous by long arched funicles which arise some distance below the apex of the fruit. Pericarp parenchymatous. Seeds slightly inflated, obovate in outline, beaked at the micropyle, anatropous with ventral raphe. Testa one cell thick, formed of equiaxial cells. TYPE SPECIES. P. schweinfurthi (Heer). Palaeowetherellia schweinfurthi (Heer) Chandler (PL 12, figs. 24-30 ; PL 13, figs. 31-38 ; PL 14, figs. 39-47 ; PL 15, fig. 48 ; Text-fig, i) 1876. Diospyros schweinfurthi Heer, p. 6, pi. i, figs. i-io. 1876. Royena desertorum Heer, p. 10, pi. i, figs. 11-16. 1889. Diospyros schweinfurthi Heer : Schenk, p. 745, text-fig. 384 7 ~ 13 . 1889. Royena desertorum Heer : Schenk, p. 745, text-fig. 384 14 ~ 18 . !939- Diospyros schweinfurthi Heer : Krausel, p. 106, pi. 2, figs. 9, 10 ; text-fig. 32. 1939. Royena desertorum Heer : Krausel, p. 106. DIAGNOSIS. Fruit shorter than broad, 6 to 9, 10 or even 12-loculed. Funicles arising from the axis at about one-third to one-half the length of the fruit from the apex, passing between the closely opposed surfaces of the locule above the seed. Seed so obliquely placed as to lie almost transversely with the hilum upwards to- wards the circumference of the fruit, and the ventral margin parallel with the arched funicle. Fine parenchyma which forms the bulk of the carpel wall enclosing numer- ous patches of coarser cells very liable to decay associated with a system of fibres near the surface of the endocarp. Diameter of endocarp, 17 to 22 (or ? 24) mm. ; length about 7-5 to 10 mm. Maximum length of fruit with exocarp preserved, 14 mm. UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 169 NEOTYPE. A nine-loculed fruit (PL 13, figs. 31-37). Figured Krausel, 1939, pi. 2, figs. 9, 10 ; text-fig. 32. Brit. Mus. (N.H.), No. .12985. LOCALITIES AND HORIZONS. Farafra, Egypt ; Lower Danian (Upper Cretaceous). Gebel Tarawan and Gebel Um-el-Ghanaim, Kharga Oasis, Egypt ; Dano-Montian Lower Esna Shales. Gebel Atshan, Gebel Durvi and Abu Tundub, Kosseir Area, Red Sea ; Dano-Montian Lower Esna Shales. DESCRIPTION. Fruit : Having an exocarp (leathery ? but usually abraded) about 1-6 mm. thick, its surface more or less smooth but the details of its structure obscured by mineral incrustation (PI. 12, fig. 27 ; PI. 14, figs. 42, 43). It encloses a syncarpous multilocular endocarp, circular or sub-circular in outline, sometimes slightly angled over the locules, depressed dorsiventrally, usually somewhat flattened at the apex, either slightly pointed below or somewhat excavated (PI. 12, figs. 24-26 ; PL 13, figs. 31, 32). The style base may be marked by a small inconspicuous scarcely prominent cir- cular scar (PL 13, fig. 31), but this is not usually apparent. External surface with radial ridges or sutures which correspond with the locules. Alternating with them are less conspicuous ones which overlie the septa ; they are not invariably seen, and are most marked in much abraded specimens or in those which have started to split septicidally (PL 12, figs. 25, 29, 30). Dehiscence conspicuously loculicidal, less obviously septicidal. Possibly the septicidal planes of weakness are cemented by infiltration of mineral substance as they are normally so inconspicuous. Carpels 6 to about 12, radially arranged about the axis of the fruit (PL 12, figs. 24-26 ; PL 13, figs. 31, 32), unequally developed in some specimens so that complete fruits may show perfect or imperfect radial symmetry. .12985, for example, shows unequal development at opposite ends of one diameter, two of the smaller carpels are evidently abortive, but their existence is clearly revealed by a longitudinal fracture of the fruit (PL 13, figs. 33-36). One of Heer's transversely sectioned specimens (1876, pi. i, fig. 9) in which he only recognized 8 locules clearly shows 9 in the figure, 2 being abortive or ill-developed. Up to the present the least number of carpels seen is six (PL 12, figs. 24, 25). Fibres of the fruit axis fused with the surrounding parenchyma not occupying a distinct central canal. Locules extend from the axis to the periphery of the fruit, but are tangentially compressed and so completely flattened around their edges that the two flat surfaces are contiguous. The seeds are neither as long nor as broad as the locules. It is only where the seed actually lies that there is a slightly inflated cavity equal to the thickness of the seed (PL 13, figs. 32-36 ; PL 15, fig. 48). Placentation axile. Seeds solitary, suspended by long arched funicles which spring from the axis at about one-third or one-half of the length of the fruit from the apex. From the point of emergence of the funicle from the axis (PL 13, figs. 33-37 ; Text-fig. ID) to the point where it passes into the seed at the hilum it lies within that part of the locule where the opposed walls are contiguous. The longer axes of the seeds lie at an angle of 60 or thereabouts to the axis of the fruit, hence the, position of the seeds is oblique or almost transverse in the fruits. As this is a consistent feature it must be original. Owing to this peculiarity, the dorsal surface of the seed lies towards the base of the fruit, the hilar end is directed outwards and upwards, the distal end lies close to the lower end 170 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT of the fruit axis, and the ventral margin is uppermost, lying parallel with the long arched oblique funicle. Pericarp of three layers : (i) a leathery (?) exocarp of fine parenchyma whose surface is obscure, but in section it can be seen in one place to be formed of rounded more or less equiaxial cells about 0-028 mm. in diameter. (2) A compact parenchy- matous coat which constitutes the main thickness of the endocarp. It is greatly thickened in the part of the fruit between the flattened margins of the locules. The outermost part of the endocarp is formed of close fine parenchyma which is rather readily abraded. Superficially, when the coat is weathered, the cells can be seen to be arranged in radiating groups. Possibly there is a fibre at the centre of each group. Differential weathering of this coat produces a series of rugosities having the effect of a coarse network, the centre of each radial group being sunk below the general level of the surface. Beneath the layers of radially grouped cells comes the main thickness of the endocarp. It is formed of equiaxial cells about 0-02 mm. in diameter. Within it are numberous patches of coarse angular paren- chyma, with cells about 0-05 to o-i mm. in diameter, developed most conspicuously near the outer surface of the coat. These patches also may have a thin fibrous core fed by fibres visible near the periphery on the flat surfaces of the loculicidally frac- tured carpels (PI. 12, fig. 30 ; PL 13, fig. 35). Here loops are seen from which branches are directed inwards to the centre of the endocarp, while more numerous and finer branches are directed outwards to its external surface. V.i 2985 had laid long exposed before fossilization, and the coarse parenchyma patches have decayed more readily than the more compact tissue which surrounds them. Their decay has produced deep funnel-shaped cavities with circular orifices which form a conspicuous feature at the surface of the endocarp. They are best shown on the lower surface, where they occur in more or less longitudinal rows (PI. 13, fig. 32). The whole surface of this specimen (as preserved) has been highly polished by the abrasion it has undergone. (3) The innermost carpellary coat is a smooth locule-lining formed of small cells arranged so as to produce straight or slightly sinuous or criss-cross lines or striations with a general transverse or oblique orientation. The distance between the striae is about 0-02 mm. The striate lining is seen to the left of the axis in PI. 13, fig. 36, and is more clearly visible on the specimen itself and on the surface of the abortive locule-cast. There is some reason to think that that part of the thick parenchymatous coat which was in contact with the locule-lining was rather spongy in texture, or that it was at least softer and less resistant to decay than the layers outside it. In places it is partially decayed, while in other places it appears obscurely columnar in trans- verse section (PI. 13, fig. 35, to the left of the fertile locule-cast). It clings in patches to the locule-cast, giving the appearance at first sight of a warty testa formed of fine angular, parenchymatous cells (PL 13, fig. 35 ; PL 14, fig. 41). But careful scrutiny reveals its true nature. The decay is most conspicuous around the axis at the base, exposing the locules and seeds in some specimens (PL 12, fig. 26 ; PL 13, fig. 32). Length of fruit, 7-5 to 9-5 mm. ; transverse diameter, 17 to at least 22 (? 24) mm. Length of .12985, 8 mm. ; diameter, 19 mm. Length of specimen shown in UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 171 PI. 14, figs. 42, 43, with exocarp preserved, 14 mm. ; estimated diameter, 24 by 17 mm. Length of another fruit, 10 mm. ; diameter, 22 mm. Seeds (PI. 13, figs. 33, 35, 37 ; PI. 14, figs. 40, 41, 43-47 ; PI. 15, fig. 48) : More or less obovate in outline, very slightly inflated, with a small beak-like projection at the hilar end which carries the micropyle. This organ is indicated by the radial alignment of the cells at its tip. That the radicle probably lay within the beak is suggested by its shape. The general symmetry indicates an anatropous seed with hilum in the concavity which delimits the " beak " from the main body of the seed, i.e., closely adjacent to the micropyle. Here the funicle is seen to enter the seed (cf. PI. 12, fig. 28 ; PI. 13, fig. 35). Raphe marginal, ventral, indicated by the form of the seed as well as by a shallow marginal groove at the rounded end opposite to the micropyle. The groove ends on the dorsal surface. Its termination probably indicates the position of the obscure chalaza. On the rounded end of the best developed seed exposed by abrasion within a locule-cast of .12985 the chalaza is probably marked by a small deep depression in the same position (i.e., at the lowest point of the seed as it lay in the fruit). If so this is the cast of the raphe fibres, where they turn sharply inwards to enter the seed or albumen. Testa apparently only one cell thick, since the cells of the external and internal impressions appear to agree exactly in size and character. It was formed of equi- axial polygonal angular cells about 0-025 to 0-03 mm. in diameter, convex externally (as shown by the concave external impressions on a small fragment of external cast seen in the specimen figured in PI. 14, figs. 42-47), concave internally (shown by the convex impressions on the internal casts of the seed figured in PI. 14, figs. 39- 47). Dimensions of seeds : Length, 5-5 to about 7 mm. ; breadth, 4 to 5 mm. ; thickness, i to 1-5 mm. REMARKS AND AFFINITIES. In examining and interpreting mineralized fruits it is necessary to remember that mineral substance in solution may percolate through all incipient fissures as well as into actual cavities. On evaporating and hardening it may then form a cement which prevents or hinders separation of parts along natural planes of weakness. But in that the film of cement may often be incom- plete, indications of natural dehiscence can usually be detected. Thus in Wether- ellia (Reid & Chandler, 1933 : 251), while many specimens were so cemented that their loculicidal dehiscence was concealed, the fruits were found in such large numbers that plenty of evidence as to their natural manner of splitting was available. In the fossil just described septicidal dehiscence may be obscured in a similar way. The mineral cement may cause adhesion of surfaces and tissues which in life, or at least after maceration, whether natural or artificial, would separate readily. For example, in Palaeowetherellia, a locule-cast may simulate a seed (which it covers and conceals), and patches of endocarp adhering to the cast but torn from adjacent tissues may look like testa (cf. PI. 13, fig. 35 ; PI. 14, fig. 41). Again thin films of cement between delicate tissues may reproduce as external or internal impression coats which could scarcely have escaped destruction, e.g., the delicate one-cell thick testa of W ether ellia and Palaeowetherellia. Palaeowetherellia is now known from more than a dozen specimens. In addition 172 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT to those figured by Heer (1876) as Diospyros, ten more, five of which are incomplete, are now available. These will be catalogued below with details of any special features they show and a note of the place of origin. The best preserved for study was described and figured by Krausel (1939 : 106, pi. 2, figs. 9, 10 ; text-fig. 32) as Diospyros schweinfurthi Heer. Certain of its characters, including the number of locules, were obscure at the time, but have since been clearly shown by the beautiful section (largely a natural fracture surface) now exposed (cf. PL 13, figs. 33-37). On this section most of the newly recorded details are based. Two other specimens from the Kharga Oasis have yielded addi- tional data, thanks to their broken condition, and have made it necessary to unite Diospyros schweinfurthi Heer and Royena desertorum of Heer as a single species (cf. p. 176). Other fruits or fragments from Kosseir throw light on the variation in the number of locules, the branching of fibres of the endocarp, mode of dehiscence, variations of size, and character of exocarp. The new evidence makes it impossible any longer to refer this species to the family Ebenaceae in spite of some superficial resemblances to Diospyros. In this genus the soft pulpy fruit breaks irregularly ; its radially arranged locules (with coarse striae) have no flattened area where the two opposed surfaces are in contact. The seeds are pendulous on short funicles which spring from the upper inner angles of the locules arising at the top of the fruit axis (Text-fig, i A) . The anatropous seeds have a conspicuous superficial dorsal marginal raphe which does not terminate at the chalaza (i.e., at the opposite end of the seed to the hilum) but is continued along the ventral margin so as to encircle the seed. It gradually tapers, dying out finally close to the hilum. The seeds lie vertically in the locules with their ventral margins parallel with and close to the axis. They are not beaked like those of the fossil but the terminal micropyle usually leads into a large conspicuous canal produced by incurving of the testa. The radicle of the embryo occupies this canal, which on an internal cast of the seed would appear as a conspicuous deep depression or pit (Text-fig. IE) with a projection from its base representing the cast of the radi- cular pocket. Similar features are characteristic of Royena, which resembles the fossil even less than Diospyros in the form and fewness of its seeds. One other characteristic may be noted, namely the cell-structure. In Diospyros the testa cells, while they may be similar in style to those of the Egyptian fossil, are convex intern- ally, so that on an internal seed-cast they would produce concave instead of convex impressions. Unfortunately it has not yet been possible to find any living closely allied genus, although a suggestion will be made below as to family relationship. But there can be no doubt at all that Palaeowetherellia closely resembles the fossil genus W ether ellia Bowerbank from the London Clay in its form, placentation, locule characters, seeds, and cell-structure, and in the tendency for dissolution of the carpel to occur at the centre which allows the seeds to escape through the gaps thereby produced a point noted by Bowerbank in describing Wetherellia. Similar decay may be seen in crab apples which have lain long on the ground in winter. A detailed description of Wetherellia variabilis was given by Reid & Chandler (1933, 251, pi. 9, figs. 7-22). A second species W. dixoni (Carruthers) has since been recognized from the Cuisian (?) or it may be Lutetian or Auversian of Selsey. UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 173 Its full description awaits publication. The characters of these two species of Wetherellia are summarized in a table below. W ether ellia variabilis Bowerb. Fruit : A syncarpous 2 to 5-loculed septicidal capsule or schizocarp, later splitting locu- licidally. Sub-globular to ovoid with length equal to, greater, or less than the diameter. Smooth, ribbed, or angled externally. Locules : Radially arranged around a central axis which extends throughout the length of the fruit. Tangentially compressed so that the opposed walls are contiguous except where the seed lies. Placentation : Axile ; solitary seeds suspended on long arched funicles which spring from the axis at a point about one-third or one-quarter of the length of the fruit from the apex. The funicles lie within that part of the locule where the opposed walls are contiguous. Seed: Slightly oblique, with the distal end nearer to the axis than the proximal. Pericarp : Of thick angular parenchyma. Locule- lining smooth and shining, obliquely or transversely and finely striate. mm. Diameter, 12 Dimensions of fruit : Length, 12 to 20 to 24 mm. Seed : Scarcely inflated, elongate obovate in out- line ; slightly beaked at the narrow hilar end, anatropous ; hilum terminal, raphe ventral, micropyle adjacent to the hilum ; chalaza small and inconspicuous at the opposite end to the hilum. Testa thin, a single layer of angular equi- axial cells convex externally, concave intern- ally, 0-03 to 0-05 mm. in diameter. Dimensions of typical seeds : 12 X 3-75 X 1-5 mm. Wetherellia dixoni (Carr.) A syncarpous 5 to 7 (or more ?) loculed capsule splitting septicidally and loculici- dally. Sub-globular but somewhat dorsiven- trally compressed. Longitudinally ribbed externally. Septal fibres apparently give rise to hollow external spines. As for W. variabilis. As for W. variabilis except that the funicles spring from the axis at a point about one-sixth to two-sevenths of the length of the fruit from the apex. As for W. variabilis, but the seeds some- what broader in proportion to their length. As for W. variabilis. Length, 4 to 7 mm. Diameter 13-5 to 15 mm. (somewhat dorsiventrally crushed). As for W. variabilis, but somewhat shorter and broader. Raphe fibres not actually seen. Testa not seen. (1) 5-5 x 4 x i mm. (2) 7 x 4-75 X 1-5 mm. UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT It may be noted that the parenchymatous tissues of both Wetherellia and of the Egyptian fruits is evidently very liable to decay, for partial destruction of the centre of the fruit has occurred in both so that the locules are exposed, allowing the seeds to escape. Consequently locules which have not dehisced in the normal way may be empty. While therefore Wetherellia and the Egyptian fossils show a striking measure of agreement such as suggests a family relationship, their characters are sufficiently distinctive to indicate a generic difference between them. The name Palaeowether- eUia is here suggested for the Egyptian fruits, which thus become Palaeowetherettia schweinfurthi (Heer). The distinctions so far as they are known at present may be summarized in tabular form. Wetherellia Septicidal capsule or schizocarp, also splitting loculicidally. Locules 2 to 7 (or more ?). Long .arched funicles springing from the axis at one-sixth to one-third of the length of the fruit from the apex. Seed lying slightly obliquely in the locule with the distal end nearer to the axis than the proximal. Seeds scarcely inflated, oval to elongate obovate in outline, beaked at the narrow end. Hilum terminal. Palaeowetherellia Loculicidal capsule also splitting septi- cidally. Locules 6 to about 12. Long arched funicles springing from the axis at one-third to one half the length of the fruit from the apex. Seed so obliquely placed as to be almost transverse, with the proximal end towards the outer edge of the fruit and the distal end near the lower end of the axis. Seeds slightly inflated, obovate in outline, beaked at the narrow end. Hilum almost terminal just below the beak on the ventral margin. In 1933 Reid & Chandler referred Wetherellia to the family Linaceae but it is necessary to correct this ascription on the following grounds : (1) More than five locules are now known to occur in undoubted Wetherellia (W. dixoni}. (2) As noted in 1933 the coat of Hugonia (Linaceae) with which Wetherellia was compared is fibrous, that of Wetherellia and of Palaeowetherellia is consistently parenchymatous with a few scattered fibres. (3) The seed in Hugonia occupies more of the locule than the seed of Wetherellia or Palaeowetherellia. (4) The funicle is short and straight in Hugonia, not long and arched as in Wetherellia and Palaeowetherellia. (5) Polygonal cells of the testa are much finer in Hugonia than in Wetherellia or Palaeowetherellia . (6) The chalaza of Hugonia forms a large conspicuous scar unlike the inconspicu- ous scarcely distinguishable chalaza of Wetherellia and Palaeowetherellia. Taken together these differences appear on maturer reflection to be such as to distinguish Wetherellia and Palaeowetherettia from Linaceae. UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 175 The true relationship of the fossils therefore remains to be discovered. They may belong to an extinct family whose nearest living allies have not yet been traced. But certain features point to Euphorbiaceae as a possible alliance. These features are the combination of loculicidal and septicidal dehiscence common in capsules of Euphorbiaceae, the number of radially arranged locules, and their tangential com- pression such an arrangement is present in Hura and the point of origin of the funicles from the axis which may be well below the apex of the fruit in many Euphor- biaceae. Pendulous seeds solitary in the locules with ventral raphe. At the same time it must be frankly admitted that no really comparable genus has yet been dis- covered, so that attribution to Euphorbiaceae must be regarded as doubtful. But no other known family shows so many of the characters of the fossil as the Euphor- biaceae. MATERIAL. Fruit (PI. 13, figs. 31-37). Also figured Krausel (1939, pi. 2, figs. 9, 10 ; text- fig. 32). Neotype. A nine-loculed fruit now fractured longitudinally to show one fertile and one abortive locule, the median axis and mode of placentation. Cell patches of the inner carpellary layers adhere to the fertile locule-cast giving it a roughened granular appearance which at first sight simulates a rough testa (PI. 13, figs. 33, 35). The elongate cells or striations of the locule surface are visible on the abortive locule-cast and the locule wall between it and the axis (PI. 13, figs. 34, 36). The base of the specimen has decayed probably through long exposure before fossilization, displaying locules, locule-casts and seeds (PI. 13, figs. 32-34). The carpel wall has also suffered from differential decay, which has produced deep superficial pits originally occupied by coarse-celled tissue especially on the lower surface (PI. 13, fig. 32). Incipient loculicidal splitting is visible on one radial rib at the apex of the fruit (PL 13, fig. 31). Diameter, 19 mm. ; height, about 8 mm. (somewhat reduced by decay at the base). From the Lower Danian (Upper Cretaceous) ; Farafra, Egypt. Brit. Mus. (N.H.), No. .12985. Part of a fruit (PL 13, fig. 38 ; PL 14, figs. 39-41)- A wedge-shaped loculicidal segment of a fruit bounded by the external surface on one side, and by two adjacent locules on its two lateral faces. The inner end of the wedge (fruit wall and axis) has broken or decayed, leaving exposed the locule and seed-casts projecting inwards (PL 14, fig. 40). Tangential breadth of segment, 6-5 mm. ; height, 7-5 mm. Radius from centre (as preserved) to outer edge, 7-5 mm. Estimated breadth of fruit, about 17 mm. Length of seed or locule-cast, 5-5 mm. ; breadth, 4 mm. ; thickness, i mm. The exocarp is not preserved, the surface of the capsule is rough, not differentially decayed before fossilization as in .12985. The substance is parenchyma with cells about 0-02 mm. in diameter in which patches of coarser parenchyma (cells 176 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT about 0-05 mm. in diameter) are embedded. The decay of such coarse patches produced the deep pits in .12985. The locule-casts narrow to the exterior and are directly slightly upwards, at first sight suggesting parietal placentation. One was more fully displayed by removal of a few adherent fragments of the opposed carpel wall. Some of the parenchyma cells of the wall adhere at this narrow end of the locule-cast producing the effect of a rough nodular testa (PI. 14, fig. 41). Nearer the inner end of the locule abrasion has removed first the parenchymatous cells of the carpel and then a layer of locule-cast, thereby exposing the seed-cast (PI. 14, fig. 41 sc.), with its angular equiaxial convex cell-impressions about 0-025 to 0-03 mm. in diameter (hence cells were concave inwards on the testa). The testa is also represented by fragments of its external impression showing the same cell-impressions which are concave on this surface (hence the external surface had convex cells). This fruit segment is clearly identical in character and size with Heer's figure of Royena desertomm (1876, pi. i, figs. 11-16). But the details of its cell-structure and the arrangement of its locules also unite it with .12985 and with another specimen figured in PI. 14, figs. 42-47 ; PI. 15, fig, 48. Hence this imperfect fruit constitutes a most important link in the evidence which unites Royena desertorum with Palaeowetherellia schweinfurthi (Heer) . From the Dano-Montian Lower Esna Shales ; Gebel Um-el-Ghanaim, Kharga Oasis, Egypt. Brit. Mus. (N.H.), No. .31114. A perfect six-carpelled endocarp (PI. 12, figs. 24, 25) with base intact. The locu- licidal sutures are clear, the septicidal more obscure except where abrasion has removed the outermost layers. The whole upper surface has been somewhat abraded showing the radiating groups of fine parenchyma. Diameter, 17 mm. ; length, 9-5 mm. Brit. Mus. (N.H.), No. .31110. A seven-partite endocarp, slightly asymmetrically developed. It is somewhat corroded on one side so that a seed (or locule-cast) is partially exposed. The exo- carp is almost entirely abraded, one small patch only persisting at the base. The surface of the endocarp is also much abraded. The whole specimen is encrusted with mineral deposit which forms small pimples over the exposed fibre ends and coarse cell patches. Diameter, 17 mm. ; length, 10-5 mm. M. I. Youssef Collec- tion, 1952. Two segments of an endocarp showing the cavities of two locules. The lines of loculicidal dehiscence can be detected. The surface is somewhat abraded, exposing the irregular rugosities due to the radiating clusters of cells just below. A seed- cast in the locule between the two segments shows clear evidence of the ventral raphe. M. I. Youssef Collection, 1952. The above three specimens are from the Dano-Montian Lower Esna Shales ; Gebel Atshan, Kosseir Area, Red Sea. An eight-loculed endocarp (PL 12, fig. 26), perfect except for decay at the centre of the base so that the inner angles of the locules are exposed. The septa show UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 177 median planes of weakness as for septicidal dehiscence. Five locules have retained their seeds, some of which are abortive ; from the others the seeds have fallen through the gap caused by the decay above described. No exocarp is preserved, and there is considerable mineral incrustation over the surface of the abraded endo- carp. Hollows for the fibre ends are visible on the lower surface ; a few encrusted rounded knobs probably indicate their position on the upper surface. Half of a fruit (PI. 12, figs. 27, 28) which has been fractured longitudinally through two of the locules. One has a locule-cast preserved, in the other the locule-cast is missing, but the funicle (now much encrusted with a mineral deposit) is seen. There appears to be evidence of seven locules on this fragment, so that the complete fruit must have had at least eleven or twelve locules. Adherent remains of exocarp are seen at the base and apex only and are shown in section where the coat has broken away from the endocarp. Exocarp and endocarp are much encrusted by a mineral deposit. On the endocarp the encrusted remains of the fibres described on p. 170 form small rounded prominences. Half of an endocarp which has split longitudinally (PL 12, figs. 29, 30). Three and a half carpels are preserved and the septicidal and loculicidal fracture lines are clearly visible. The half carpel has split both loculicidally and septicidally and has been pushed out of position. The surface is sufficiently abraded to expose the radial grouping of the small cell-clusters. The network of fibres is obvious on one loculicidal suture plane, although owing to the adherent parenchyma and secondary incrustation the fibres are not themselves exposed but their position is very apparent. No exocarp is preserved. Diameter of endocarp, 18 mm. ; length, 9 mm. A few small shell casts and impressions adhere to one fracture plane. The above three specimens are from the Dano-Montian Lower Esna Shales ; Abu Tundub, Kosseir Area, Red Sea. Brit. Mus. (N.H.), Nos. .31111-13. A fruit (PI. 14, figs. 42-47 ; PL 15, fig. 48) with exocarp preserved but cracked in such a manner that at first sight it simulates three perianth segments (PL 14, fig. 42). Actual diameter (incomplete because the specimen had been polished on one side at p in figs. 42, 43 thus showing a tangential section), 18 x 17 mm. Estimated complete maximum diameter (distorted), about 24 X 17 mm. Height, 14 mm. The whole fruit is cracked through desiccation and crushing, and is held together by calcite cement (white in the figures). On the polished surface two seeds can be seen, one in transverse and one in oblique section. The first shows the seed-cast (sc) surrounded fairly closely by the locule-cast (Ic) (PL 15, fig. 48), which is em- bedded in the pulp or parenchyma of the carpel. The base of the fruit is sunk, and broken along one radius so that a beautiful seed-cast was exposed, at first held in place by calcite cement, but later becoming detached (PL 14, figs. 43-47). The beaked end of the compressed ovate cast was directed outwards and upwards in the locule as in the section of .12985 shown in PL 13, figs. 33, 35, 37. 178 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT Length of seed-cast, 7 mm. ; breadth, 4-75 mm. ; thickness, 1-5 mm. Surface of cast with equiaxial slightly convex cell-impressions. Their divergence at the beaked end indicates the micropyle. A shallow marginal furrow at the opposite rounded end of the cast dies out on the dorsal surface (close to an acci- dental fracture line) . It indicates the inner end of the raphe, its termination marking the site of a small inconspicuous chalaza. From the Dano-Montian Lower Esna Shales ; Gebel Tarawan, Kharga Oasis, Egypt. Brit. Mus. (N.H.), No. V.3HI5. Two segments of an endocarp showing the cavities of three locules. The carpel wall between the two segments is abraded so that the seed-cast is exposed along its dorsal margin. When the segments are separated the chalaza can be detected at the lowest point of the seed-cast as it lies in the endocarp. The exterior of the endocarp is also abraded, so that in places the inner thick layer of fine parenchyma is exposed. Remains of the outer layer with radiating cell clusters occur hi patches, but are much obscured by mineral incrustation. Length of endocarp, 8 mm. ; maximum breadth across the two loculicidal segments, 14 mm. From the Dano-Montian Lower Esna Shales ; Gebel Durvi, Kosseir Area, Red Sea. M. I. Youssef Collection, 1952. Family ICACINACEAE Genus ICACINICARYA Reid & Chandler, 1933 Icacinicarya youssefi n. sp. (PI. 15, figs. 49-51) DIAGNOSIS. Fruit and endocarp almond-shaped or sub-obovoid in outline, lenti- cular in transverse section. External surface of endocarp with a series of more or less discontinuous irregular rugosities, some longitudinal, others transverse or oblique. Cells of endocarp markedly sinuous or coarsely digitate. Length of endocarp, 31 mm. ; breadth, 22 mm. ; thickness, 10 mm. Length of a fruit, 33 mm. ; breadth, 26-5 mm. ; thickness, 13 mm. HOLOTYPE. Endocarp figured PI. 15, fig. 51. Brit. Mus. (N.H.), No. .31117. LOCALITIES AND HORIZON. Gebel Atshan and Gebel Durvi, Kosseir Area, Red Sea ; Dano-Montian Lower Esna Shales. DESCRIPTION. Fruit (PI. 15, figs. 49, 50) : Bisymmetric, somewhat compressed, giving a lenticular transverse section, obovoid in outline, slightly mucronate at the apical style, attachment at the opposite extremity to the style. Surface much puckered as if by shrinkage of the mesocarp, but having a few rather ill-defined longitudinal ridges which may branch or anastomose halfway up. They may be due to fibres in the pulp or just beneath the epicarp. The actual epicarp itself is probably missing. One margin (overlying the funicle ?) thicker than the other with two or three marked longitudinal ridges. Whole surface with a pattern of UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 179 small, rounded or quadrangular depressions, about 0-5 to i mm. in diameter, ar- ranged in longitudinal rows. Finer structure of digitate or markedly sinuous cells. The convergence of the ornamentation at the two ends indicates the position of style and attachment described. Length of fruit, 33 mm. ; breadth, 26-5 mm. ; thickness, 13 mm. Endocarp (PI. 15, fig. 51) : Similar in form to the fruit, base narrowed to a point where the opening for the funicle is clearly seen. A ridge due to splitting followed by mineral infiltration is seen at the apical style. This tendency to split at the stylar end has been observed in other Icacinaceae, e.g., Natsiatum eocenicum Chandler from the Lower Headon of Hordle. As in genera of Icacinaceae one margin is much thicker and more rounded than the other. Experience has shown that the thick margin carries the funicle but no section is available, so this cannot be verified in the present instance. External surface with an obscure pattern of small mostly discontinuous rugosities, some elongate and longitudinally aligned. They do not produce a definite network of ridges and hollows as in Icacinicarya platycarpa Reid & Chandler (1933 : 345, pi. 16, figs. 11-18). Surface of endocarp formed of small digitate or conspicuously sinuous cells with a tendency to transverse alignment. They are about 0-114 nim. broad and 0-057 mm. in length. Length of endocarp, 31 mm. ; breadth, 22 mm. ; thickness, 10 mm. REMARKS AND AFFINITIES. Two specimens which by their general similarity may be presumed to belong to the same species although one shows the whole fruit, the other the endocarp only. One surface of the fruit shows the outline of a closely comparable endocarp owing to the contraction on drying which the pulpy exocarp has undergone (PI. 15, fig. 50). Although the most conclusive diagnostic characters of Icacinaceae can only be inferred because the unbroken character of the specimens conceals them, there can be little doubt of the relationship. The specimens are considerably larger than Icacinicarya platycarpa, which they resemble in their form, but as pointed out above, they can also be distinguished from that species by surface ornamentation. The cells of the endocarp are considerably larger than in that species and differ in their digitate form. Although further information about this species is much to be desired, it is suffi- ciently well defined to be recognizable again, so that the specific name, Icacinicarya youssefi, after the finder, has been given. Family ICACINACEAE ? ICACINICARYA sp. ? (PI. 15, figs. 52-54) LOCALITY AND HORIZON. Gebel Atshan, Kosseir Area, Red Sea ; Dano-Montian Lower Esna Shales. DESCRIPTION. Fruit : Sub-ovoid, bisymmetric about a plane which includes the two major axes and probably the funicle, although this was not actually seen, GEOL. II, 4. 13 i8o UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT somewhat sharply angled in the plane of symmetry on one side, rounded and much inflated on the other (funicular margin). Style terminal at the apex, marked by a slight prominence. Attachment at the opposite end of the major axis to the style where the fruit shows a slight flattening, marked clearly by a small circular scar from which the surface cells radiate. The broad margin of the fruit is semicircular in outline ; the narrow margin is slightly concave immediately below the style and for about one-third of the length, but is convex for the lower two-thirds. Surface puckered and wrinkled giving a leathery appearance. Surface cells obscure except around the attachment where they are equiaxial, rounded, slightly convex, and about 0-038 mm. in diameter. Length of fruit, 13 mm. ; breadth in plane of sym- metry, 11-5 mm. ; thickness at right angles to plane of symmetry, n mm. REMARKS. The solitary specimen looks like a drupe with epicarp and mesocarp preserved so that all the characters of the endocarp are hidden except so far as they can be inferred from the form and symmetry of the specimen. The peculiar asym- metric outline as viewed at right angles to the plane of symmetry and the occurrence of one broad rounded and one angled margin may indicate a fruit of Icacinaceae. But pending the discovery of further evidence the specimen can only be referred tentatively to the form-genus Icacinicarya. Family FLACOURTIACEAE ? Genus THIEBAUDIA nov. DIAGNOSIS. A large berry with about 36 parietal placentas, numerous seeds in two close set rows on the placentas, and a pulpy mass of tissue which occupies the whole of the fruit cavity. Seeds probably sub-ovoid. Length of fruit (compressed), 13 mm. ; breadth (much increased by compression but bereft of pericarp), 37 mm. Diameter of seeds, I to 1-5 mm. TYPE SPECIES. Thiebaudia rayaniensis n. sp. Thiebaudia rayaniensis n. sp. (PI. 16, figs. 58-63) DIAGNOSIS. As for genus. HOLOTYPE. A single fruit with most of the pericarp removed. Thiebaud & Robson Collection (Locality no. 604), 1951. Brit. Mus. (N.H.), No. .31120. LOCALITY AND HORIZON. Wadi Rayan, Western desert of Egypt ; Eocene (? Lutetian or slightly younger). DESCRIPTION. Fruit : A globular or sub-globular berry (now much compressed dorsiventrally, so that the wall has been buckled all around the equator of the specimen). Dehiscence probably by irregular breaking of the pericarp as no indi- cation of sutures for regular dehiscence can be seen. The pericarp is preserved only at the extreme base and apex (PI. 16, figs. 58, 59), elsewhere it has been broken away or abraded. The external surface shows a few obscure radial flutings at the upper end, but is UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 181 preserved in a matrix so coarse that it does not show the cell structure of the epicarp if still present. A slightly denned and very slightly sunk area at the apex, about 5 mm. in diameter, may mark the base of the style. Thickness of pericarp about 3 mm. Numerous broad, rather flat, stout, longitudinal bands of fibres can be seen (although all are now incomplete) in the lower half of the fruit lying in fragmentary remains of the pericarp, but above the equator they appear to have been abraded. Placentation parietal, the seeds arranged in two close-set rows on each of thirty-six longitudinal placentas. Sometimes the seeds in the two rows are opposite one another, sometimes they appear to be alternate. Between each pair of placentas there is a very slight longitudinal ridge which thickens at its extreme apical end. These ridges produce on the inner surface of the pericarp a series of broad shallow longitudinal channels or pockets into which the placentas and seeds fitted. Each placenta begins to produce its seeds at about 10 mm. from the apex of the fruit, where it springs from a pointed tongue of tissue with a narrow median furrow (PI. 16, figs. 58, 60). The whole of the interior of the fruit is filled by a pulpy mass which adheres closely to the pericarp. The shallow ridges of the pericarp give rise to grooves upon the surface of the pulpy-mass. These separate the broad flat-topped ridges opposite the placentas bearing the concavities caused by the seeds. It is the surface of this mass which is exposed intact on the upper surface of the specimen (PI. 16, figs. 58, 60). The pulp is formed of coarsely and deeply sinuous cells, and throughout its thickness there are numerous small cavities about the same size as these cells (i.e., about 0-057 to 0-114 mm. in diameter.) No tendency to split either radially or otherwise has been detected in the pulpy mass. Length of fruit (much reduced by dorsiventral compression), 13 mm. ; breadth (correspondingly increased by compression but reduced by the loss of the pericarp), 37mm. Seeds : Very obscure. Producing sub-circular or sub-ovoid hollows on the pulpy mass (PI. 16, figs. 58-61). Perhaps somewhat laterally compressed. The shallow convex external surface of the seed was ornamented with large inflated radially arranged cells or areoles diverging from a knob-like projection (PI. 16, fig. 62). This structure is visible on the impression of a detached seed (probably belonging to the fruit) which is preserved on the pericarp near the base. Obscure traces of similar cells were also seen on a few seed-impressions on the pulpy mass. Other seed-impressions merely show a rounded prominence which must represent a con- siderable depression (hilar, micropylar, or chalazal ?) on the actual seed surface (PI. 16, fig. 61, best shown in seed s). Internal casts of seeds (preserved in a few instances on the underside of the fruit but always incomplete, PI. 16, fig. 63) show a smooth shining surface with longitudinal striations due to very long narrow cells with stout longitudinal walls, 0-014 to 0-018 mm. broad, lying parallel with the long axis of the seed. There are also slight traces of a coat of transversely aligned cells or fibres. Diameter of seeds, i to 1-5 mm. An internal cast measured 1-5 mm. in length, i mm. in breadth. REMARKS. One fruit preserved in a ferruginous cement filled with quartz grains forming a hard mass. It appears to have been embedded in a cream-coloured fine 182 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT sand with Nummulites. The sand filled every crack and cranny of the cast, and a Nummulite (reported by Mr. C. D. Ovey as having a range from Lutetian to Middle Oligocene) was found in sand lying in the deep hollow between the lower surface of the pulpy mass and the remains of the pericarp at its base. There can be little doubt that the Nummulite belonged to the strata in which the fruit occurred, for little attempt had been made to clean the fossil of the adherent sand which clung with persistence. It has now been boiled and the sand brushed away. Other fossils from these beds are also said to be ferruginous, but this is the only plant as yet discovered. The age of the deposit is regarded as Lutetian, or possibly some- what younger, but definitely Eocene. As stated, most of the pericarp of this fruit was missing, revealing the central pulpy mass. It is a difficult specimen to understand and interpret. But probably, if its living counterpart were discovered, the description given above could be con- siderably simplified and shortened. The coarseness of the matrix is not well calculated to preserve fine details. Little can be seen of the seed structure, especially as the hollows on the central mass are largely merely cavities in the pulp due to the pressure exercised by the growing seeds. They do not therefore give much information apart from the size and, rather ob- scurely, the form of the close-set seeds which are themselves almost entirely absent. Even these impressions are partially confused by some measure of overlap. The bases of the stout fibres in the fruit wall persist in broken ragged fragments at the base of the specimen, where they project from the fragment of pericarp. Sometimes they are missing, and the openings from which they came can be seen in the edge of the broken wall. The upper side of the specimen shows in good condition the surface of the pulpy mass with impressions made by the seeds on the ridges opposite the placentas, and the shallow furrows between these ridges which are the impressions of the narrow ridges on the pericarp wall described above. The lower surface is less well preserved. It appears to have become torn and battered before or during fossilization, so that the actual lower surface of the pulpy mass together with the alignment and ridges on this surface is largely destroyed. Traces only of these features and of the collapsed and fragmentary pericarp wall with its fibres can be detected. Detached seeds represented by occasional external impressions or true internal casts have been pushed into the soft pulp and are visible here and there. Usually the casts are obscure. The true internal casts are rarer than the external impressions. A curious and at first misleading feature of this specimen is a coarse transversely elongate meshwork of angular ridges around the equator. A radial fracture of the pulpy mass demonstrates the purely secondary and inorganic character of these ridges, which are due to the infiltration and setting of a limonite cement in cracks caused by buckling of the walls and pulp. No organic structure is to be seen in the seams of limonite which fill these cracks. AFFINITIES. The parietal placentation of this multi-carpelled fruit limits relation- ship to very few families. At first sight the specimen recalls a Poppy capsule on account of its radial symmetry and rounded small seeds, while the remains of the fruit wall at the apex simulate the stigma-bearing disc of Papaver. Here, however, UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 183 the resemblance ends. The pulpy mass of the interior is wholly unlike anything in Papaveraceae. Parietal placentas associated with such a mass of pulp do, however, occur in Flacourtiaceae. But the difficulty here is that in no known genus of that family do as many as 36 placentas and carpels occur, while the details of seed structure in the fossil are too imperfectly known to be conclusive. No other living family appears to bear so close a relationship to this lovely fossil, and past experience has demonstrated the tendency for a larger number of locules to occur in fossil than in living forms, although not perhaps to the degree here recorded. In the absence of more satisfactory information the specimen has been referred doubtfully to the family Flacourtiaceae. It has been given a new generic name, Thiebaudia, after one of the finders, while the specific name rayaniensis indicates the place where it was found. The writer is of opinion that this specimen ought to be compared very carefully with Krausel's species Nymphaeopsis lachmanni from the Lower Oligocene of Cairo. The supposed placenta and unequal hammer-shaped involucral segments of that species may quite possibly be parts of a pericarp which has cracked and contracted into irregular segments on drying. Such a feature is displayed by one fruit of Palaeowetherellia from Kharga (PI. 14, fig. 42), in which the cracked berry with its mineral infiltrations simulates a trifid perianth. The fibres shown by Krausel (1939, pi. 2, figs. 2, 3, 6) on the surface exposed beneath the supposed involucral segments are not altogether unlike those of Thiebaudia. The direction in which they branch suggests that the supposed apex is in fact the base of the specimen. The buckling of the fruit at the equator due to dorsiventral compression has pro- duced a similar network of infiltration ridges to those described in Thiebaudia. It is not impossible that the apparent operculum of the seeds might be interpreted as a large chalazal scar. But these are merely suggestions for future consideration. They cannot be substantiated without the most careful study of the specimens themselves. Carpolithus hassani n. sp. (PI. 16, figs. 64, 65) LOCALITY AND HORIZON. Gebel-el-Ter, Kharga Oasis, Egypt ; Dano-Montian Lower Esna Shales. A beautiful but puzzling specimen suggests one valve of a bisymmetric endocarp. It appears to have been rubbed down artificially at the margins but to no great depth. The outline, as it exists at present, is broadly elliptical, 20 mm. long, 17-5 mm. broad, 5-5 mm. deep. The external surface is gently convex and the internal correspondingly concave. The thickness of the wall at the polished margin is 2 to 2-5 mm. Three deep external grooves must have some structural significance. They appear to be the outer edges of slits which pass through the whole thickness of the wall. The two nearest the margins of the valve slope inwards towards the centre of the fruit so that their inner edges, marked by deep grooves on the surface of the locule, lie slightly closer together than do their outer edges on the external surface 184 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT of the valve. These two grooves are parallel with the outline of the valve, but they do not meet, although their ends lie much closer together at one extremity (apex ?) of the valve than at the other. One groove is longer than the other. The third groove which pierces the endocarp also, lies midway between the other two but is shorter than either. It begins at about the middle of the valve and passes towards that end of it (base ?), where the other two grooves are most widely separated. It dies out before reaching the edge of the valve. Within the slits the surface of the endocarp is longitudinally striate. On the exterior, between the longer curved marginal groove and the short median one, a longitudinal slightly sinuous furrow can be seen which was evidently a channel for a fibre, impressions of which are seen. It gives off two or three short slender branches at its upper end. These diverge and taper upwards. Surface of valve rough, due to the convex angular walls of the parenchymatous cells of which it is composed. Cells 0-05 to o-i mm. in diameter. Locule surface smoother, formed of equally large equiaxial cells. A first glance at this specimen suggests that it may belong to Menispermaceae. But closer scrutiny shows that the relationship is impossible for the following reasons : (1) The curved marginal grooves do not unite at the apex to form a horseshoe, nor does one of their opposite ends curve appreciably outwards. (2) The external grooves correspond with internal grooves and not with internal ridges as in Menispermaceae, where the ridges form a cavity for the curved seed. (3) The endocarp of Menispermaceae is fibrous in structure, not parenchymatous. I have found no fruit with such peculiar slits which may be connected with ger- mination. Possibly the curved area enclosed by the slits may be associated with a curved embryo. Carpolithus sp. (Icacinicarya sp. ?) (PI. 16, fig. 66) LOCALITY AND HORIZON. Gebel-el-Ter, Kharga Oasis, Egypt ; Dano-Montian Lower Esna Shales. The internal cast of a valve of an endocarp 10-5 mm. long, 9-25 mm. broad, 1-8 mm. deep. The cast is formed of coarse crystalline ferruginous matter. Its surface shows an obscure network of ridges. The internal surface of the actual specimen would have shown corresponding grooves separated by shallow convexities. No cell-structure is visible. The form of the valve suggests Icacinaceae, but the evidence preserved is insufficient for certainty. Carpolithus sp. (PL 16, figs. 55-57) LOCALITY AND HORIZON. Gebel Atshan, Kosseir Area, Red Sea ; Dano-Montian Lower Esna Shales. A sub-globular fruit slightly laterally compressed, with a three-angled apex, the angles meeting in a slight prominence. They die out at about the middle of the fruit. Between these well-marked angles a few subsidiary inconspicuous longitu- UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 185 dinal ridges can be seen. Basal attachment small, slightly sunk, marked by a little projection at the middle of the hollow. At the extreme base of the fruit there are three short, very shallow, rounded furrows opposite the three apical ridges or angles. Surface, as now preserved, ornamented with numerous angular, more or less equiaxial contiguous concave areas, somewhat variable in size, up to I or 1-5 mm. in diameter but sometimes smaller. This cast is formed of radiating groups of fine cells, the groups often but not invariably coinciding with the concavities. Length of fruit, 13-5 mm. ; diameter, 10 by 13-5 mm. There is nothing to indicate the identity of this specimen. APPENDIX I NOTE ON THE OCCURRENCE OF THE FOSSIL FRUITS AND SEEDS COLLECTED FROM THE KHARGA OASIS, WESTERN DESERT OF EGYPT By M. YOUSSEF HASSAN The specimens were collected from a bed with an average thickness of 35 metres. It is composed of dark grey and greenish shales, well foliated and often densely seamed with gypsiferous and salt intercalations. Red nodules of botryoidal limonite, sometimes attaining large sizes, are abundant. Frequently the limonite appears in pseudocrystalline form, being cubes sometimes with interpenetration twinning. The bed is rich in fossils which are dwarfed and excellently preserved in limonite. The age of this bed is " Danian " or more probably Dano-Montian. The following is a complete list of the fossil fauna identified : Schizorhabdus libycus Zittel. Palaeopsammia multiformis Wanner. Bathypsammia cleopatrae Hassan MS. Trochocyathus epicharis Wanner. Trochocyathus deniseptatus Hassan MS. Dungulia libyca (Wanner). Pattalophyllia aegyptiaca (Wanner). Terebratulina sp. Pentacrinus sp. Nucula tremolate-striala Wanner. Nucula chargensis Quaas. Leda leia Wanner. Trapezium sp. Trapezium aff. acutangulum (Deshayes). Cardium cf . becksi Muller. Cardium cf. inaequiconvexum Cossman & Pissarro. Crassatella zitteli Wanner. Cucullaea schweinfurthi Quaas. Cucullaea sp. Limea sp. Chlamys mayer-eymari (Newton). Verticordia nova Hassan MS. Corbula striatuloides Forbes. Trochus cf. laryi D'Archiac & Haime. Scala calamistrata (Wanner) . Architectonica dachelensis (Wanner). Natica (Gyrodes) farafrensis Wanner. Natica (Euspira) terensis Hassan MS. Rissoa cf. crassistriata Wood. Campanile cf. brookmani Cox. Cerithium abictiforme Wanner. Cerithium bigeniculatum Wanner. Cerithium cf. periphractum Wanner. " Alaria " schweinfurthi Quaas. Cypraea cf. kayei Forbes. Tonna sp. Tudicla peroni Quaas. Sassia tuberculosa (Kaunnowen). Sassia farafrensis (Quass). Sassia chalmasi (Quaas). Sassia sp. Athleta (Volutilithes] daniensis (Quaas). Cyclichna cf. regulbiensis (Adams). Solidula chargensis (Quaas). Solidula pharaonum (Wanner) . Avellana cretacea Quaas. Pyrgopolon sp. Nautilus applanatus Wanner, ex Zittel MS. Nautilus desertorum Quaas, ex Zittel MS. 1 86 This horizon is resting on a series of alternating marls and impure limestones with occasional phosphatic bands of undoubtedly late Maestrichtian age. APPENDIX II NOTE ON THE OCCURRENCE OF THE FOSSIL FRUITS AND SEEDS COLLECTED FROM THE KOSSEIR AREA (RED SEA), EGYPT By MURAD I. YOUSSEF The fruits were collected from a single bed in different localities in the Kosseir Area (Red Sea). The bed in which these fossil fruits, as well as the fossils mentioned below, were found is a greyish-green shale varying in thickness between 50 and 125 metres. The concretions found throughout this bed, and the fossils occurring generally in a band near its middle part, are all limonitic. Cubic pseudomorphs of limonite, probably after pyrite, are also found. Irregular gypsum veins running in every direction, perhaps corresponding to the more or less pyramidal planes of jointing of the shales, are epigenetic, being secondary in origin. The fossils collected from this bed are considered to be Danian or Dano-Montian. Some 30 metres of shales and marls lying immediately below this bed are of the same age. These are underlain by Maestrichtian rocks containing many phosphate and phosphatic beds. The following is a list of the fossil fauna identified from the thick shale bed : Brachycyathus daniensis Wanner. Trochocyathus epicharis Wanner. Pattalophyllia aegyptiaca (Wanner). Caryosmilia granosa Wanner. Dungulia milneri (Gregory). Dungulia libyca (Wanner). Caryophyllia jasmundi Wanner. Palaeopsammia multiformis Wanner. Stephanophyllia (Microbacia) sp. Serpula cf . discoidea Wanner. Pentacrinus sp. Salenia sp. Cyphosoma sp. Hemiaster chargensis Wanner. Hemiaster (Leucaster) lamberti Cottreau. Terebratulina chrysalis (Schlotheim). Terebratulina sp. Nucula chargensis Quaas. Nucula lucida J. Boehm. Nucula tenera J. Miiller (non S. V. Wood). Nucula tremolate-striata Wanner. Leda leia Wanner. Thyasira cretacea (Wanner). Cardium cf. inaequiconvexum Cossmann & Pissarro. Cardium cf. becksi Miiller. Trapezium sp. Lucina dachelensis Wanner. Crassatella matercula Mayer-Eymar. Area modioloides Wanner. Cucullaea sp. Chlamys mayer-eymari (Newton). Verticordia nova Hassan MS. Scala cf . desertorum Wanner. Natica farafrensis Wanner. Campanile cf. brookmani Cox. Alaria schweinfurthi Quaas. Alaria sp. Sassia chalmasi (Quas). Neptunea zitteli (Quaas). Neptunea sp. Fusus sp. Athleta (Volutilithes) desertorum (Quaas). Solidula chargensis (Quaas). Avellana cretacea Quaas. Avellana sp. Cylichna cf. regulbiensis (Adams). Dentalium bicarinatum Wanner. Nautilus desertorum Quaas, ex Zittel MS. UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 187 REFERENCES AXELROD, D. I. 1952. Variables affecting the Probabilities of Dispersal in Geologic Time. Bull. Amer. Mus. Nat. Hist., New York, 99 : 177-188, 2 figs. BONNET, E. 1904. Sur un Nipadites de 1'eocene d'Egypte. Bull. Mus. Hist. nat. Paris, 10 : 499-502, 2 figs. BOWERBANK, J. S. 1840. A History of the Fossil Fruits and Seeds of the London Clay. 144 pp., 17 pis. London. CHANEY, R. W. 1940. Tertiary Forests and Continental History. Bull. Geol. Soc. Amer., New York, 51 : 469-488, pis. i, 2. 1947- Tertiary Centers and Migration Routes. Ecol. Monogr., Durham, N.C., 17 : 140-148. EDWARDS, W. N. 1936. The Flora of the London Clay. Proc. Geol. Ass. Lond., 47:22-31, 3 figs. ENGELHARDT, H. 1907. Tertiare Pflanzenreste aus dem Fajum. Beitr. Paldont. Geol. Ost.- Ung., Wien, 20 : 206-216, pis. 18, 19. HEER, O. 1876. Ueber fossile Friichte der Oase Chargeh. N. Denkschr. schweiz. Ges. Naturw., Zurich, 27 : i-n, pi. i. KRAUSEL, R. 1939. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wusten Agyptens, IV. Die fossilen Floren Agyptens, 3. Die fossilen Pflanzen Agyptens, E.-L. Abh. bayer. Akad. Wiss., Munchen (N.F.) 47 : 1-140, pis. 1-23. & STROMER, E. 1924. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wusten Agyptens, IV. Die fossilen Floren Agyptens, 3. Die fossilen Pflanzen Agyptens, A-C. Abh. bayer. Akad. Wiss., Munchen (N.F.) 30, 2 : 1-48, pis. 1-3. KRYSHTOFOVICH, A. N. 1929. Evolution of the Tertiary Flora in Asia. New Phytol., Cam- bridge, 28 : 303-312. LE MAOUT, E., & DECAISNE, J. 1876. A General System of Botany. 2nd edit. 1066 pp., 5500 figs. London. REID, C., & E. M. 1915. The Pliocene Floras of the Dutch- Prussian Border. Meded. Rijksopsp. Delfst., Amsterdam, 6 : 1-178, pis. 1-20. REID, E. M., & CHANDLER, M. E. J. 1926. The Bembridge Flora. Catalogue of Cainozoic Plants in the Department of Geology, 1. viii -+- 206 pp., 12 pis. Brit. Mus. (Nat. Hist.), London. 1933- The Flora of the London Clay, viii + 561 pp., 33 pis. Brit. Mus. (Nat. Hist.), London. RENNER, O. 1907. Teichosperma, eine monokotylenfrucht aus dem Tertiar Agyptens. Beitr. Palaont. Geol. Ost.-Ung., Wien, 20 : 217-220. SCHENK, A. 1889. Palaeophytologie. In Zittel, K. A., Handbuch der Palaeontologie, 2 : 958 pp., 433 figs. Munich & Leipzig. SEWARD, A. C. 1934- An Extinct Malayan Flora in England. Sci. Progr. Twent. Cent., London, 29, i : 1-24. & ARBER, E. A. N. 1903. Les Nipadites des couches Eocenes de la Belgique. Mem. Mus. nat. Belg., Bruxelles, 2 : 1-16, pis. 1-3. NOTE. While this paper was going through the press my attention was drawn to a memoir by L. W. LeRoy (Biostratigraphy of the Maqfi Section, Egypt. Mem. Geol. Soc. Amer., Washington, 54, 1953) m which, from a study of the Foraminifera, the author concludes not only that the " Esna Shales " are definitely Lower Eocene, but also that the Danian is probably unrepresented in Egypt. There may, however, be some unresolved confusion over the use of the term " Lower Esna Shales." GKOL. II, 4 I 4 PLATE 10 Nipa burtim (Brongniart) FIG. i. The domed apical end of a drupe which is incomplete below the middle. Length preserved, 25 mm. ; breadth, 27 mm. Estimated complete length, about 45-50 mm. It shows the typical longitudinal fibro-vascular bundles embedded in parenchyma, (u) the apical umbo. x 2-8. (.31105.) FIG. 2. The same, viewed from below, looking on to the fractured surface. The side shown in fig. i is towards the top of the figure. Part of the unridged smooth endocarp (e) is exposed. Transverse fibres of the endocarp are obscurely seen. The pericarp is in section at (p, p) and in surface view with its longitudinal fibres at (/). X 2-8. FIG. 3. An obliquely distorted seed-cast in which the basal aperture (a) occupies a basi- lateral position. Slight longitudinal flutings of the surface are visible. There is a mosaic pattern all over the surface, x approx. (V. 13239.) FIG. 4. The opposite surface of the same seed-cast, apical end, showing the sub-apical funnel-shaped opening (ch) and the raphe-fibre impressions converging towards it and passing into it. (m) marks the apical mucro of the nut. x 1-2. FIG. 5. A seed-cast so compressed as to be almost lenticular, (a) is the position of the basal aperture. The coarse network of fibres which lay between the two integuments of the testa is faintly impressed upon the cast to the left of the median line. X J approx. (.13240.) Nipa fruticans Thunberg FIG. 6. A seed for comparison with fig. 5 from which the outer coat of the testa has been entirely removed. Most of the inner coat has also been removed except beneath the slightly sunk coarse network of fibres ( /, / ) which lay between the two coats and which still adheres to the seed. In the meshes between the fibres (where the inner coat has gone) the transverse alignment of the small ruminations of the endosperm can be seen rather obscurely. The seed has been cut longitudinally. X 2 approx. Recent ; Singapore. Bull. B.M. (N.H.) Geol. 2, 4 PLATE 10 2 f NIPA BUKTINI, NIPA FRUTICANS PLATE 11 a burtini (Brongniart) FIG. 7. Opposite side of the seed-cast in PI. 10, fig. 5, showing detail of endosperm structure. The elongate alignment at (r) indicates the position of the raphe, the radial arrange- ment indicates the point of entry of the raphe fibres (ch) below the apex of the cast, x 1-8. FIG. 8 The same seed-cast, detail of the surface depicted in PI. 10, fig. 5, showing the more normal transverse alignment of the endosperm, x 1-8. FIG. 9. The base of a seed with its ribbed outer integument partially preserved but suffi- ciently abraded to expose the flat broad fibre band (/). The basal aperture is clearly seen. The specimen has undergone much lateral compression, x i. Figured by Krausel, 1939, pi. i, fig. 24 as Rubiaceocarpum markgrafi. Nipa fmticans Thunberg FIG. 10. A seed, side, showing the ribbed outer integument with clear impressions of the longitudinal, and more obscure impressions of the transverse, endocarp fibres, x 1-7. Recent; Singapore. FIG. ii. The seed in PI. 10, fig. 6, showing the flat raphe band (r) lying in a shallow longitudinal furrow of the endosperm. On each side of the band the transverse alignment of the endosperm is visible. Above where the band has been removed irregular arrangement of the endosperm is exposed at (ch) and just below longitudinal alignment can be detected. X 2 approx. FIG. 12. The same seed, base, showing the aperture for the radicle, the deep furrow (g) due to the incomplete septum which projected from the locule-wall, and shallow furrows or sinuosities of the surface. The raphe lay in the furrow at (r). x 1-75. Anonaspermum aegypticum n. sp. FIG. 13. The distal end of a seed-cast showing typical ruminate endosperm with encircling raphe seen at (r, r). The fibrous remains of testa still adhere in the depression at the middle of the broad surface. The specimen had been fractured transversely and the hilar end was missing, (ch) indicates the beginning of the chalaza, most of which has been broken away with the hilar end. x 3. (.31106.) FIG. 14. The same, end view. It shows the encircling raphe partly broken on the left so that the ruminations are exposed in section at rs. The chalaza begins at (ch). x 3. FIG. 15. The same, fractured surface, showing the four-partite endosperm, x 3. Bull. B.M. (N.H.) Geol. 2, 4 PLATE 11 IO NIPA BURTINI, NIPA FRUTICANS, ANONASPERMUM AEGYPTICUM PLATE 12 Lagenoidea trilocularis Reid & Chandler FIG. 16. Side view of a fruit showing three of the six loculicidal segments of the capsule. X 3. (.31107.) FIG. 17. Base of the same specimen, showing the attachment scar and wrinkled epicarp which partially obscures the segments. X 3. Fig. 1 8. Apex of the same, showing more clearly the six segments and lines of dehiscence. X 3- FIG. 19. Side view of a second specimen showing three segments clearly ; the edge of a fourth is just visible on the right-hand margin of the figure. X 2-8. (.31108.) FIG. 20. The same, base, showing six segments and an aperture where placenta and perianth have broken away. X 2-6. Lagenoidea bilocularis Reid & Chandler FIG. 21. A fruit, side, showing the narrow median segment representing the edge of the septum, and two of the broader lateral segments, x 3. (.31109.) FIG. 22. The same, apex, showing the narrow median segments which form the ends of the septum and the two pairs of segments which overlie the locules. Loculicidal splitting occurs at (I, I), septicidal splitting at (s, s). x 3. FIG. 23. The same, base. The septum is damaged on one side at this end of the fruit. Lettering as in fig. 22. x 3. Palaeowetherellia schweinfurthi (Heer) Chandler FIG. 24. Lower surface of a six-carpelled endocarp showing the six loculicidal suture lines (/, /). Slight abrasion of this surface has displayed the radiating groups of fine parenchyma which produce a pitted effect. Decay at the centre of the base has scarcely started. X 2-8. (.31110.) FIG. 25. The same, apex. The outer part of the carpel wall is corroded in places showing the finer-celled parenchyma of the layers beneath. Loculicidal sutures are clear ; the septicidal ones (sp, sp) between them are more obscure except where abrasion has partly removed the outer layers of the endocarp. x 2-8. FIG. 26. The under surface of an eight-loculed endocarp, perfect except for the charac- teristic decay at the centre of the base so that locules and septa (showing planes of weakness associated with septicidal splitting) are exposed. Several locules have shed their seeds. In others the seeds are ill-developed, x 2-8. (.31111.) FIG. 27. The base of half a fruit which retains a considerable patch of exocarp (ex) over the median area. Around the circumference the exocarp has been worn away exposing six seg- ments of endocarp representing seven locules (/, /). X 2-8. (.31112.) FIG. 28. The fractured surface of the same showing two locules. The one on the right is occupied by a locule- and seed-cast, (ch) indicates the position of the chalaza. The left-hand locule is empty but the much encrusted funicle (/) is visible. Its origin in the axis is obscured by mineral deposit, (ex) = exocarp in section, x 3. FIG. 29. Base of another incomplete endocarp with three and a half segments representing four locules (/,/). The half segment on the right has slipped out of position along the loculi- cidal fracture plane (/). The surface of this half segment shows the plane of septicidal fracture (sp). X 2-8. (V. 3 iii 3 .) FIG. 30. The same, looking on to the fractured surface, (sp) is the septicidal fracture plane. (/, /) indicates loculicidal surfaces. The network of fibres is seen on the right-hand loculicidal suture surface. The left locule surface is obscured by molluscan casts (out of focus in the figure). X 3. Bull. B.M. (N.H.) Geol. 2, 4 PLATE 12 LAGENOIDEA TRILOCULARIS, L. BILOCULARIS, PALAEOWETHERELLIA SCHWEINFURTHI PLATE 13 Palaeow ether ellia schweinfurthi (Heer) Chandler FIG. 31. The apex of a nine-carpelled fruit polished by abrasion. (I, 1} indicates planes of loculicidal dehiscence. Between them, but more obscure, are radial lines indicating planes of septicidal dehiscence (sp). At (Is) incipient loculicidal splitting can be seen at the middle of the ridge. The specimen has been fractured along the line (/,/). The small circular scar at the centre may be the axis or the style base, x 3. (.12985.) FIG. 32. The same, base, corroded at the centre so that the locules are exposed (some now empty). Note the conspicuous pits over the surface due to the decay of fibres surrounded by coarse parenchymatous patches. Lettering as in fig. 31. x 3. FIG. 33. The fractured surface (longitudinal section) of the above showing a locule-cast on the left with the funicle arising from the axis of the fruit and passing in the plane of the locule to the hilum situated near the upper outer angle. The right-hand locule is empty, x 3. FIG. 34. The counterpart half of the same, showing on the right the locule surface which overlay the cast in fig. 33, and on the left an abortive locule-cast which occupied the empty locule in fig. 33. x 3. FIG. 35. The left-hand part of the section in fig. 33. A larger magnification shows the funicle just above the locule-cast and the entry of the funicle into the cast marking the position of the hilum on the enclosed seed. This fertile locule lies at (fl) on the left of fig. 32 where the edge of the cast is seen. Around the hilar end of the cast adherent patches of tissue torn from the endocarp simulate a rugose testa, x 7. FIG. 36. The left-hand part of the section in fig. 34. The fibres of the axis show more clearly. Oblique fine striations on the surface of the abortive locule close to the axis are visible and the small abortive locule-cast is shown at (a). This locule lies at (fl) on the left of fig. 31. x 7. FIG. 37. The same as fig. 34, right-hand side of specimen, with a detached seed from another specimen laid in the locule to show the approximate position that the seed would have occupied. Axial fibres (a) and funicle (/) are well shown in the endocarp wall and on the flat surface of the locule respectively. The curved line at the rounded end of the seed is an acci- dental fracture, x 7. (cf. PI. 14, figs. 43-47.) FIG. 38. A segment of another fruit which has broken loculicidally, apex. Axis to left at (a), circumference to right, loculicidal surfaces above and below, x 6. (V. 31114.) Bull, EM. (N.H.) Geol. 2, 4 PLATE 13 37 32 PALAEOWETHERELLIA SCHWEINFURTHI PLATE 14 Palaeow ether ellia schweinfurthi (Heer) Chandler FIG. 39. Side view of the fruit segment in PI. 13, fig. 38. It shows the rounded inner end of a seed-cast (c) exposed by the breaking away in fossilization of the axis and inner angle of the carpel. Specimens with similar proportions were separated by Heer (1876) under the name Royena desertorum. x 6. FIG. 40. The same, looking on to the inner edge where the two locules and seed-casts can be seen, x 6. FIG. 41. Opposite side of the segment to that shown in Fig. 39. Part of the carpel wall still adheres to the cast at the outer end ; it conceals the hilar end of the seed and embracing locule-cast, and produces the false effect of a rough nodular testa. The smooth surface at (Ic) is the remains of the thin locule-cast closely adherent to the seed. To the left at (sc) the film of locule-cast has broken away, exposing the seed-cast. A narrow band of the smoothly finished loculicidal surface is preserved at (Is), but elsewhere this surface has been torn away, thereby exposing the angular cells of the carpel wall at (cw). x 10. FIG. 42. Apex of another fruit (incomplete as indicated by dotted lines) with exocarp preserved. Crushing and drying have caused the specimen to crack, white calcite has been deposited in the cracks which give a false appearance of three perianth segments. The speci- men had been rubbed down along a plane parallel with its axis at (p). X 1-8. ("^31115.) FIG. 43. The same, base. A crack on the right exposes a shining seed-cast. This cast is illustrated laid in the locule of another fruit in PI. 13, fig. 37. Calcite-filled cracks are again seen. Lettering as in fig. 42. x 1-8. FIG. 44. Part of the same, more highly magnified and tilted to show the seed-cast of which the lower margin is exposed. X 6. FIG. 45. The same seed-cast, removed from the fruit, showing the narrow ventral edge. The micropyle is at (m), the hilum at (h). The plasticene in which it was necessary to mount the specimen to obtain this view of the seed somewhat obscures its outline at the lower end of the figure. X 6-5. FIG. 46. The same seed-cast, side view, (m) indicates the position of micropyle and radicle, (h) that of the hilum. The chalaza lay near the lower end of the accidental fracture line. X 10. FIG. 47. The same cast, opposite surface, to show the bisymmetry of the seed, x 10. Bull. B.M. (N.H.) Geol. 2, 4 PLATE 14 42 PALAEOWETHERELLIA SCH WEINFURTHI GEOL. II, 4 15 PLATE 15 Palaeowetherellia schweinfurthi (Heer) Chandler FIG. 48. The partially polished surface (p) of the fruit in PL 12, figs. 41, 42, showing a tangential section through a locule and seed. The seed-cast (sc) is closely embraced by the locule-cast (Ic) lying between the two thick loculicidal valves of the carpel. The raphe (r) is probably indicated by the slightly constricted area at the narrow upper edge of the seed-cast. The white mass to the right is calcite filling the loculicidal split and now cementing together the two valves of the carpel (v, v). x 10. Icacinicaryayoussefi n. sp. FIG. 49. A fruit, broad surface, showing form and ornamentation, (st] indicates the posi- tion of the style. The funicle was situated in the right margin of the enclosed endocarp. x 2-6. (V. 3 ui6.) FIG. 50. The same, opposite surface. Contraction of the pericarp has brought out the outline of the endocarp within, x 2-6. FIG. 51. An endocarp bereft of the pericarp. Note the surface ornamentation, the basal aperture for the entry of the funicle at (/), and the short vertical fracture with infiltrated mineral substance at the sty lar end (st). x 2-6. (.31117). Icacinicarya sp. ? FIG. 52. Side view of a wrinkled drupe showing the asymmetry characteristic of Icacinaceae. The more convex left margin may be presumed to carry the funicle. (st) indicates the style. X 2-8. (V.3in8.) FIG. 53. Base of the same, showing the inflated funicular margin (below) and narrow opposite margin (at the top of the figure). The small circular scar of attachment, from which the cells radiate, is seen at the centre, x 3. FIG. 54. Apex of the same. Funicle bearing margin to the left. Style at (st). x 2-6. Carpolithus sp. FIG. 55. Fruit, side, showing an angle at the apical end and small angular concavities all over the surface, x 2-8. (.31119.) FIG. 56. The same, apex, showing the three angles or ridges and the surface concavities. X 2-6. FIG. 57. The same, base, showing the slightly sunk attachment and the surface concavities, as well as the shallow grooves which lie opposite the apical angles, x 3. Bull. B.M. (N.H.) Geol. 2, 4 PLATE 15 50 49 PALAEOWETHERELLIA SCHWEINFURTHI, ICACINICARYA YOUSSEFI, ICACINICARYA SP. ? CARPOLITHUS SP. PLATE 16 Thiebaudia rayaniensis n. gen. et sp. FIG. 58. Apex of fruit. The pericarp is preserved only at the centre. It shows shallow, inconspicuous, radial furrows diverging from a slightly sunk circular area (perhaps the style base). Where the pericarp is broken away, the pulpy mass which fills the fruit cavity is exposed. On the surface of this mass radial furrows mark the position of shallow longitudinal ridges which must have projected from the fruit-wall. Between them impressions of the placentae are seen each with two rows of seeds now represented by the hollows which they produced on the surface of the pulp. Each placenta arises from a short tongue of tissue with longitudinal median furrow. These tongues project from beneath the edge of the remaining pericarp, x 1-6. (V.3H20.) FIG. 59. The same, from below. Here the pericarp is preserved only at the very centre of the base. The radial furrows (so clear on the upper surface) are obscure, for this surface has been much battered and partially destroyed. The arrangement of the seeds has been disturbed, but a few external seed impressions and internal casts are preserved scattered somewhat irregularly. A small concavity, seen at the lower edge of the central fragment of the pericarp, may be the external impression of a detached seed (cf. fig. 62). x 1-6. FIG. 60. Part of the upper surface (indicated by (x) in fig. 58). It shows more clearly the seed pockets on each side of the placentas, and the impressions of the ridges described above. The margin of the fragment of pericarp lies near the base of the figure at (p). x 6. FIG. 61. Part of the under surface (indicated by (x) in fig. 59) showing a few of the hollow pockets for the seeds. They sometimes show a central projection (best seen in the hollow marked s) which suggests a concavity on the surface of the seed. A few broken fibre-fragments (remains of pericarp) adhere to the pulpy mass, x 6. FIG. 62. The external impression of the seed (?) on the pericarp in fig. 59 as described above. The surface of the pericarp was painted white around the impression to bring out its outline. A central depression must correspond with a prominence on the seed itself. Coarse cells or areoles are obscurely seen diverging from this central depression, x 16. FIG. 63. A somewhat imperfect internal cast of a seed from the lower surface of the pulpy mass. Striae due to cells on the surface of the cast lie parallel with its longer axis. Although the cast is obscure a sharp marginal angle suggests a bisymmetric form, x 20. (V.3U2O.) Carpolithus hassani n. sp. FIG. 64. The exterior of a valve showing three deep furrows and a branching fibre which is partially embedded in the surface between the right-hand furrow and the short middle one. X 1-8. (.31121.) FIG. 65. The internal aspect of the same, showing that the wall is pierced by the three external furrows. The thickness of the wall is also shown. The smooth marginal areas appear to have been artificially produced by polishing, x 1-8. Carpolithus sp. (Icacinicarya sp. ?) FIG. 66. The internal cast of a valve of a fruit showing form and ornamentation suggestive of Icacinaceae. x 1-8. (.31122.) Bull. EM. (N.H.) Geol. 2, 4 PLATE 16 THIEBAUDIA RAYANIENSIS, CARPOLITHUS HASSANI CARPOLITHUS SP. (ICACINICARYA SP. ? ) . PRESETED 1 6 MAR 1954 PRINTED IN GREAT BRITAIN BY ADLARD AND SON. LIMITED BARTHOLOMEW PRESS, DORKING 5 OCi 1954 THE CARBONIFEROUS FLORA OF PERU W. J. JONGMANS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY Vol. 2 No. 5 LONDON: 1954 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY The following papers appeared in Volume I (1949-52) : Price No. I (1949). The Pterobranch Rhabdopleura in the English Eocene. H. D. Thomas & A. G. Davis 75. 6d. No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P. Oakley & M. F. Ashley Montagu 5$- No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone of the Welsh Borders. E. I. White. Pteraspis leathensis White a Dittonian Zone-Fossil. E. I. White 75- 6d' No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan, Northern Iraq. L. F. Spath .... . IDS. No. 5 (1951)- Cretaceous and Eocene Peduncles of the Cirripede Euscal- pellum. T. H. Withers .55. No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae). T. H. Withers ... 5*- No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian of Podolia. W. N. Croft . . IDS. No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle East. T. F. Grimsdale . IDS. No. 9 (1952). Australian Arthrodires. E. I. White . . . 155. No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 65. THE CARBONIFEROUS FLORA OF PERU BY WILHELMUS JOSEPHUS JONGMANS Pp. 189-224 ; Pis. 17-26 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) GEOLOGY Vol. 2 No. 5 LONDON : 1954 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical Series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 2, No. 5 of the Geological series. PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued September 1954 Price Fifteen Shillings THE CARBONIFEROUS FLORA OF PERU By W. J. JONGMANS SYNOPSIS A reconsideration of all the older records of Carboniferous plants from Peru, together with an examination of two important newer collections, fully confirms the Lower Carboniferous age of the flora. Several new species are described, including representatives of Lepidoden- dropsis, and the world distribution of this genus is reviewed. INTRODUCTION THE Carboniferous flora of Peru has been the subject of several papers. The principal flora is found on the peninsula of Paracas, which, as Berry (1922) points out, is largely made up of continental Carboniferous sediments and constitutes one of the very few deposits of this character in South America, and the only known occurrence of rocks of this age on the west coast of South America. Somewhat to the north-east of the peninsula there are other Carboniferous localities at Huanuco and in the neighbourhood of Cerro de Pasco. Up to the present little has been known of the flora of these localities. Gothan (1928) mentions Rhacopteris circularis Walton from Vichaicoto and S. Huanuco and Knorria and a Calamites-like specimen from Cachama, between Cerro de Pasco and Huanuco. Fortunately one of my former assistants, Dr. N. de Voogd, sent me a good collection from Carhuamayo. This collection and another from Paracas, put at my disposal by the Trustees of the British Museum, will be described in this paper. REVIEW OF THE LITERATURE ON THE CARBONIFEROUS FLORA OF PERU Almost at the same time as Berry, Seward (1922) described another set of plants from Paracas. According to these papers the outcrop at Paracas was first discovered by Fuchs (1900). Seward states : "The coal occurs in a series of greenish sandstones and grey and black carbonaceous shales, which have a north-easterly strike and dip about 25 south-eastwards. These are overlain unconformably on the neck of the peninsula by Tertiary sandstones and impure limestones. There is no definite stratigraphical evidence of the age of the coal-bearing beds, and the plants are there- fore of special importance." Fuchs (1900) recorded the following species : Catamites suckowi Bgt., Sphenopteris hartlebeni Dkr., Baiera pluripartita Schl., Lepidodendron sternbergii Bgt., Sigillaria tessellata Bgt., and Stigmaria ficoides Bgt. GEOL. II, 5. j6 IQ2 THE CARBONIFEROUS FLORA OF PERU Fuchs assigned the beds to the Upper Coal Measures, but his list is scarcely pos- sible ; it contains common Carboniferous plants and Wealden species. The Carboniferous flora was mentioned subsequently by Fuchs (1905), Dorca (1909), Marsters (1909) and Lisson (1917), without any reference to the Wealden species. Lisson's paper is the only one which contains any new facts on this flora ; he mentions Lepidodendron rimosum and L. obovatum, determined by Zeiller. The age is given as Westphalian. Some critical remarks will be given when treating the papers by Berry (1922) and Gothan (1928). The next contribution to the Carboniferous flora of Peru was that by Steinmann (1911) who did not visit Paracas. He examined Fuchs' collection at the Cuerpo de Ingenieros de Minas in Lima, and listed the following forms for the Paracas flora : Archaeocalamites radiatus, Lepidodendron cf. veltheimi, L. cf. volkmanni, Sphenopteris affinis (furcata}, Rhodea filifera and Rhabdocarpus. The occurrence of these species would prove that the flora belongs to the Lower Carboniferous. It is interesting but somewhat remarkable that the plant-bearing Carboniferous should be of Lower Carboniferous age and the invertebrate-bearing Carboniferous of Upper Carboniferous age. Unfortunately no literature on the invertebrate fauna of the Peruvian Carboniferous is known to me, and so far as can be seen from the sections in Berry's papers (1922, 1922^) no invertebrate fauna had been met with on the peninsula of Paracas during his visit. Steinmann states that no marine fossils are known from Paracas. He visited a second Carboniferous locality near Huichaycota, some kilometres south of Huanuco on the Huallaga and found large stems of Lepidodendron and at different places many specimens which he records as Rhacopteris inaequilatera Goepp. The introductory paper by Berry (i 922*1) contains no descriptions of the plants, but discusses the locality and the history of the knowledge of the flora up to that date and records the following : Palmatopteris furcata (Bgt.), Eremopteris whitei Berry, E. peruianus Berry, Cata- mites suckowi Bgt., Calamostachys sp., Lepidodendron rimosum Sternb., L. obovatum Sternb., Lepidophyllum sp., Lepidostrobus sp., Stigmaria sp. and Knorria sp. Berry visited Paracas and was able to make large collections ; his remarks on the locality and the mode of occurrence of the plants are very valuable. According to Berry the fossil plants occur at different horizons in the sections he examined, and " there is no chronologic change in the flora from top to bottom although fossil plants are more varied in the lowermost horizon. The materials are relatively coarse throughout and would seem to indicate rapid deposition." Over half of the section " is described as sandstone, which is often coarse and arkosic. Of the 273 feet described as shale 192 feet are distinctly sandy, so that less than 14% of the total thickness [of 585 feet], including the so-called coal seams, is fine-grained shale and even the coal contains much silty impurities. No underclays with rootlets, or upright stems were observed and the coaly layers have every appearance of having been formed of drift material " (Berry, 1922^ : 191-192). The flora is " extremely limited, although some of the elements are exceedingly common," especially the plants recorded as Palmatopteris furcata, Eremopteris whitei, Catamites suckowi and Lepidodendron rimosum (Berry, 1922^ : 193). Berry THE CARBONIFEROUS FLORA OF PERU 193 imagines " that the coarseness of the sediments and the apparent drifting of the material are mainly responsible for the absence of a more representative flora." Thus his collections contained " no traces of Sigillaria, Cordaites, Sphenophyllum, Catamite foliage, nor of any Neuropterids, Pecopterids, Alethopterids or Lonchop- terids." He states that this feature of the flora is undoubtedly responsible for Steinmann's opinion (1911) that the Paracas flora is of Lower Carboniferous age. According to Berry the Paracas Carboniferous corresponds to the Westphalian stage. In the same year Seward (1922) described a collection of plants from Paracas made by J. A. Douglas in 1911. Seward's flora contains Sphenopteris sp., Lepidodendron sp., Sigillaria (or Lepidodendron) sp., Bothrodendron sp., and Planta incertae sedis. His determinations are considered in detail below : Sphenopteris sp. (Seward, 1922, pi. 13, figs. 1-3) is present in several fragments. " The branched axis is longitudinally striated and smooth ; the pinnules are more or less deltoid, deeply dissected, and the ultimate segments are obtuse or truncate." Seward compares it with Sphenopteris furcata Bgt., but in this species the segments are acute. He also compares it with Eremopteris missouriensis Lesquereux (White, 1899, pi. 5, figs. 1-30) which does possess obtuse or truncate ultimate segments. Several Lower Carboniferous species have pinnules which closely resemble the Peruvian specimens ; " the deeply dissected form of the lamina suggests comparison with pinnules of Rhodea and Sphenopteridium." Thus Nathorst (1920, pi. i, figs, 11-13) figures as Sphenopteridium norbergii a plant which he compares with Sphen- opteris affinis L. & H. ; the latter differs in its broader, thicker segments, and its stronger and more numerous nerves. It will later be shown that such comparison is important for the determination of the specimens and their age. Lepidodendron sp. (Seward, 1922, pi. 13, figs. 4-6). The most important specimen (fig. 4) shows a branch with attached leaves. It is preserved on a carbonaceous sandstone and most of the details cannot be seen. However, it is clear that the " cushions " of the leaves are separated by undulate lines, that succeeding " cushions" are connected at the upper and lower ends, and that the surface is striated, though not in the deeply impressed portions of the cushions. Seward's fig. 5 shows some details on the leaf-cushion and the leaf-scar. On pi. 13, fig. 6 Seward figures a Lepidodendron which he considers belongs to the same species as those illustrated in figs. 4, 5. The specimen is interesting as it shows the leaf-like organs very clearly. It somewhat resembles the figure published by Johnson (1913, pi. 41, fig. 3), but it is impossible to decide whether it is a portion of a cone or not. The long erect leaves are narrow and possess a distinct middle nerve. They are sharply pointed at their ends. Sigillaria (or Lepidodendron) sp. (Seward, 1922, pi. 13, figs. 7, 8). " Pieces of a stem having contiguous leaf-cushions which bear leaf-scars agreeing both with some types of Sigillaria (e.g., S. brardi Bgt.) and with certain species of Lepidoden- dron. On the upper part of several leaf -cushions there is a small circular scar, presumably a ligule-pit. No vascular bundle scars or parichnos-scars can be detec- ted." Seward compares his specimens with those named 5. brardi from South Africa (Seward, 1897 : 326) and Brazil (White, 1908), which, however, probably do not belong to Brnogniart's species, nor even to the genus Sigillaria. He also com- 194 THE CARBONIFEROUS FLORA OF PERU pares them with S. mutans Weiss & Sterzel (1893 : 84). It is very curious that Seward should compare his specimens with a true Stephanian species. Bothrodendron (?) sp. (Seward, 1922, pi. 13, fig. 9 ; text-fig.). Seward describes two specimens under this heading " although it is by no means certain that the specimens shown in pi. xiii, fig. 9, and in the text-figure belong to the same species." The specimen figured in pi. 13 shows " spirally-disposed and widely-separated, slightly prominent, transversely elongated, rhomboidal leaf-scars. " On the right- hand side, a thin carbonaceous layer probably represents the actual surface . . . On the partly decorticated surface there are discontinuous longitudinal ridges, and an irregular transverse wrinkling, but on the carbonized film no wrinkling is seen. There is no indication of any leaf-cushion, no ligular pit, and only a very faint suggestion in a few of the scars of a median vascular scar. The leaf-scars shown in the text-figure are rather more rounded and appear as slightly concave areas . . . In the small and widely-separated leaf-scars these fragments agree with Both- rodendron, Pinakodendron, and Asolanus. The form of the leaf -scar and the absence of a leaf-cushion are features more suggestive of Bothrodendron." The features mentioned by Seward are those which are shown on the stems and larger branches of Bothrodendron, whereas clear leaf-cushions and leaf-scars of a more lepidodendroid form occur on the smaller branches. It is known that the gap between these two extreme forms is filled by a whole series of transitions. It is somewhat tempting to consider the three forms described by Seward as smaller branches and stems of the same species. The last plant mentioned by Seward (1922, pi. 13, fig. 10) has not been named. This is represented by crowded branched filaments which are " probably portions of pinnules of a fern-like plant, such as some of the Lower Carboniferous species referred to Rhodea or Sphenopteridium." Seward was at first inclined to regard his material as Upper Carboniferous in age. Dr. Kidston, however, regarded the palaeobotanical evidence as more favourable to a Lower Carboniferous horizon. After a re-examination of the specimens Seward modified his first opinion and agreed with Kidston. He remarks, however, that the plants are too imperfect to serve as trustworthy guides and that " further research is greatly to be desired, since the available data are inadequate as a basis for any positive statement." Shortly after Seward's paper appeared, a fully illustrated account of the flora was published by Berry (1922). The determinations are discussed below : Palmatopteris furcata (Bgt.) Berry, 1922, pi. i, figs. 1-3. The plant he named Palmatopteris furcata is exceedingly common in the Paracas deposits. He unites with it Sphenopteris affinis (Steinmann, 1911 : 50) and Sphen- opteris hartlebeni (Fuchs, 1900 : 50). These records certainly represent the same species, but it is not certain that the specific determination given by Berry is right. He does not give a detailed description and an opinion can be based only on his figures (pi. i, figs. 1-3). The species has since been renamed Sphenopteris paracasica by Gothan (1928 : 293). THE CARBONIFEROUS FLORA OF PERU 195 Eremopteris peruianus and E. whitei (Berry, 1922, pis. 2-4). Berry describes two new species of Eremopteris. He points out the difficulties of demarcation between Eremopteris (especially as used by American authors) and Rhacopteris. He compares his Eremopteris peruianus (pis. 2, 3) with Rhacopteris transitions (Ett.) as recorded by Stur (1875, pi. 8, figs. 5-7) but there are other species of " Anisopteris " with which Berry's material may be compared. Accord- ing to his description and figures the size and form of the pinnules are very variable and it is quite likely that more than one species of Rhacopteris occurs in the material. According to Berry his second species, Eremopteris whitei, is identical with E. elegans Lesquereux (1880, pi. 53, fig. 7). Berry considers this species to be entirely different from the European Rhacopteris elegans (Ett.) and from R. (Sphenopteris} asplenites Gutb. These European species have nothing in common with the Peru- vian material. Berry describes his material as follows : " The pinnae are linear oblong, their divisions or pinnules are oblique, oblong or rhomboidal in form, narrowed to the somewhat decurrent base, deeply pinnately cut by narrow sinuses into cuneate divisions which are rounded or subcrenate distad. The venation is flabellate and largely immersed in the thick substance of the lamina." Eremopteris whitei Berry cannot be a Rhacopteris (sensu stricto) or Anisopteris. It might possibly be compared with some species of Sphenopteridium as figured by different authors (e.g., Walton, 1926) but Berry's figure alone is not sufficient to identify it with Eremopteris or Rhacopteris, and until further data are available it can be named Sphenopteris whitei (Berry). It must have been a rather robust plant as evidenced by the stout longitudinally striated rachises. The nervation, as far as can be seen in Berry's figure, does not seem to agree with that of Rhacopteris or Anisopteris nor with Eremopteris elegans as figured by Lesquereux (1880, pi. 53, fig. 7) ; it can much better be compared with that of Sphenopteridium. Another plant with which it may be compared is that figured by me (1940, pi. 4, fig. 9) from the Lower Carboniferous of Egypt and erroneously referred to Rhodea cf. hochstetteri Stur. Better material which I received later shows that the Egyptian specimens cannot belong to Stur's species, and I now have little doubt as to their identity with Berry's species. Calamites suckowi Bgt. (Berry, 1922, pis. 5-7). The plants figured by Berry as Calamites suckowi Bgt. are very fragmentary and broken. Most of them do not show a nodal line. The best example is seen in pi. 6. Here a nodal line is present on which the sharply pointed ribs are clearly alternating, so that a reference to Asterocalamites is precluded. The stems undoubtedly belong to Calamites, but the sharply pointed ribs show that it cannot be C. suckowi ; it looks more like C. undulatus Sternb. Lepidodendron rimosum Sternb. and L. obovatum Bgt. (Berry, 1922, pi. 8 ; pi. i, fig- 5). The two species of Lepidodendron, L. rimosum Sternb. (plate 8) and L. obovatum Bgt. (pi. i, fig. 5) described and figured by Berry agree with Zeiller's opinion on this ig6 THE CARBONIFEROUS FLORA OF PERU material (1917). It must be pointed out, however, that if Berry's figures agree with the originals there is a remarkable difference between pi. i, fig. 5, and pi. 8, figs, i, 2, on the one hand and pi. 8, fig. 3, on the other. Only in pi. 8, fig. 3, is a leaf- scar visible and it is likely that this drawing was not made from the figured specimens but included only to indicate Berry's conception of the species. His determ