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Microtine remains from the Norwich Crag (Lower Pleistocene) of Easton Bavents, Suffolk
Microtine Remains from the Norwich Crag (Lower Pleistocene) of Easton Bavents, Suffolk by J. N. CARRECK Received 19 September, 1963; taken as read 2 December 1966 CONTENTS 1. 2. 3. 4.
page 491 493 494 495 495 496
INTRODUCTION DESCRIPTION DISTRIBUTION CONCLUSION ACKNOWLEDGMENTS REFERENCES
ABSTRACT: A small series of Microtine teeth from the Lower Pleistocene Norwich Crag of Easton Bavents, Suffolk, preserved in Ipswich Museum, is described and assigned to the vole Mimomys, some, if not all, being of M. pliocaenicus (ForsythMajor), a new record for this locality. Selected specimens are figured. The stratigraphical and geographical distribution of the species in Britain and on the Continent are discussed in relation to its presence at this site, and the local environment in which it I ived is considered.
1. INTRODUCTION the recent publication of Funnell & West's paper on the Early Pleistocene deposits of Easton Bavents, near Southwold, Suffolk (1962), it may be of value to place on record the occurrence of some Microtine remains in the Norwich Crag of that locality collected by the brothers D. and P. Long of Lowestoft, and Mr. R. J. Winyard of Walberswick, Suffolk, and preserved in Ipswich Corporation Museum. These were kindly submitted to the author for determination by Mr. H. E. P. Spencer, when geologist of that museum. The precise horizon from which the fossils were obtained is not certainly recorded, but all or most came from the Upper Shell Bed described by Larwood & Martin (1954, 166). Easton Bavents cliff is a significant locality for larger mammalian remains in the Norwich Crag (Larwood & Martin, 1954, 166-8; Spencer, 1963, 264-5 and 1964, 344; Spencer, in Funnell & West, 1962, 137). These authors record a Baleen whale, Balaena sp.; 'Mastodon' sp.: the Southern Elephant, Mammuthus (Archidiskodon) meridionalis (Nesti); cf. 'Rhinoceros' sp.; the zebra, Equus (Hippotigris) robustus Pomel; Hippopotamus sp.; and the deer Euctenoceros sedgwicki (Falconer), Eu. falconeri (Dawkins), and Megaceros verticornis (Dawkins). The only previously published record of Microtine remains from the Norwich Crag of Easton Bavents, IN VIEW OF
491
492
J. N. CARRECK
5
2
7
8 Fig. 1. Teeth of the vole Mimomys from the Norwich Crag of Easton Bavents, Suffolk 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.
Left first lower molar (rnr), of Mimomys pllocaenicus, Crown (occlusal) view, x 8.0. The same specimen, Inner (lingual) view, x 8.0. Another left mi (incomplete), M. pliocaenicus, Occlusal view, X8.5. The same specimen, Outer (labial) view, X 8.0. A further example of the left rni (incomplete), Mv pliocaenicus. Occlusal view, X 7.5. The same specimen. Inner view, x7.5. Right first upper molar (m-j, Mimomys sp, indet. Occlusal view, X8.5. The sarne specimen. Inner view, x8.5. Right lower incisor (incomplete), Mimomys sp, indet. Inner view, X 2.0. Left upper incisor (incomplete), Mimomys sp, indet. Inner view, X 2.0.
MICROTINE REMAINS, NORWICH CRAG, SUFFOLK 493
however, is of some incisors in the W. M. Crowfoot and Searles V. Wood collections at Norwich Castle Museum, and others obtained in 1951-2 by the brothers Long (Larwood & Martin, 1954, 168). These teeth are unfortunately not diagnostic. 2. DESCRIPTION The remains examined by the present writer consist of lustrous, mineralised molar and incisor teeth, the former showing dark brown enamel and light brown dentine and cement, and the incisors showing dark brown enamel and light brown dentine. Some are a little rolled, but Mr. Spencer (verbal information, May 1964) considers that they are probably not derived from a deposit other than the local Norwich Crag as he knows of no apparent source. Mimomys pliocaenicus (Forsyth-Major). Four left m, One of these molars (Fig. I, Nos. 5 and 6) is of an adult vole in which the posterior loop is broken away, one root is developed but partly broken off, and the other is lost. Maximum length preserved: 2.75 mm.; maximum width: 1.4 mm. The inner portion of the third outer enamel infold is completely insulated. Another example (Fig. 1, Nos. 3 and 4) is very similar as regards the form of the occlusal surface, but the upper part of the posterior loop is broken away, the root below is preserved and the anterior root is lost. Maximum length: 3.4 mm.; maximum width: 1.4 mm. In another specimen the roots have not yet appeared. The occlusal surface is deeply worn, and the posterior loop and second inner salient angle have been lost. The last specimen (Fig. I, Nos. 1 and 2) shows the distinctive enamel islet, large and completely developed, in the anterior loop. The tooth is perfect and the two completely formed roots are preserved. Therefore this tooth is of an adult individual. Maximum length: 3.3 mm.; maximum width: 1.4 mm.; maximum height: 4.3 rom. M. cf. pliocaenicus. One left m 3, one right m3 and a left m., In the left m3 root development has begun and an enamel islet is present. The right m 3 also shows root development and is very similar to the last. The left m I is very incomplete, being the whole base of the tooth but without roots and only possessing the anterior part of the occlusal surface (anterior loop), although this shows the enamel islet and a slight prism fold. Mimomys sp. indet. One left rn-, four right m! (Fig. 1, Nos. 7 and 8), one right m 2 , one right m, and eleven parts of incisors (Fig. I, Nos. 9 and 10). The upper molar shown in 7 and 8 of Fig. 1 has a maximum length of 2.9 mm., and a maximum width of 1.6 mm. The lower incisor shown in No.9 has a length of 14.25 mm. and a height of 1.6 mm., whilst the upper incisor in No. 10 has a length of 10.1 mm. and a height of 1.9 mm. These remains are not sufficiently diagnostic for precise determination, but may well be of M. pliocaenicus.
494
1. N. CARRECK
3. DISTRIBUTION This species of vole was originally described from the Late Villafranchian lacustrine deposits of the Val d'Arno, Tuscany, Northern Italy (ForsythMajor, 1902, 103) and also in that country remains are known from the Lower Pleistocene lacustrine marl at Bocchignano in Sabino (Tuccimei, 1893), probably referable to this species or a near ally. In France it is known from the Late Villafranchian of Seneze in Haute-Loire (Kormos, 1931, 1-5), and the Middle Villafranchian deposits of Sete, Herault (Friant, 1953; Hinton, 1954) have yielded remains then ascribed to this species, but now referred to a much more primitive species, M . occitanus Thaler. This author considers it to be of Pliocene age (Thaler, 1955). Other French occurrences are the 'Gunzien' (Villafranchian or earlier) of Perrier, Central France (Friant, 1954, 166-8), and the Villafranchian of Saint-Vallier (Viret, 1954). In Holland remains are known from the Tiglian (Middle Villafranchian) of Tegelen (Newton, 1908; Schreuder, 1949, 116-17), Gorinchem (Gorkum) (Riitten, 1909; Hinton, 1926, 358), Eindhoven (Van der Sluijs, 1957, 257) and Veghel (Van der Sluijs, op. cit., 258), and the Lower Taxandrian (Late Villafranchian) of Raamsdonkveer ? and Breda ? (Van der Sluijs, 1957, 257), and in Germany from the Early Gla ciat ion Interstadial/Earl y Glaciation II deposits of Gundersheim (Heller , 1936), the Lower Cromerian, s.l., of Deinsdorf (idem, 1963), and Lower Pleistocene deposits at Hohensiilzen (Weiler, 1952). In Poland others have been described from deposits probably of the Early Glaciation Interstadial at Kadzielnia Hill in Kielce (Kowalski, 1958, 29-33, and in lit., 1964). In Czechoslovakia Fejfar (1961) has recorded this species from lacustrine tuffaceous deposits of the latter part of the Villafranchian at the Hajnacka II site. In Hungary remains have been found in the Middle /Late Villafranchian deposit s of Beremend (Mehely, 1914), and the Antepenultimate Interglacial (Cro merian, s.l.) deposits at Berg NagyHarsany and Csarnota (Mehely, 1914), Villany-Kalkberg (Ko walski, 1958, 6), and in Rumania in deposits of the Early Glaciation II (Late Villafranchian) at Puspokfurdo (Kowalski, 1958). The English occurrences are in the Norwich Crag of Thorpe, near Aldeburgh, Suffolk (Forsyth-Major, 1902), Yam Hill, near Southwold, Suffolk (Hinton, 1926, 355 and 361), Covehithe, near Southwold (undescribed material in Ipswich Museum, determined by the present writer), and Bramerton, near Norwich, Norfolk (Hinton, 1926, 361) ; the Upper Freshwater Bed of Ostend, near Bacton, Norfolk (Hinton, 1926, 355), probably derived from the Norwich Crag; in derived pieces of clay in the middle or estuarine division of the Forest Bed Series (Antepenultimate Interglacial) at East Runton, Norfolk (Hinton, 1926, 358 and 376), and in Weybourne Crag at that locality (Hinton, 1926). In the Norwich Cra g it occurs alone or possibly in association with Mimomys newtoni (Forsythe
MICROTINE REMAINS, NORWICH CRAG, SUFFOLK 495 Major), but in the Weybourne Crag of East Runton it is found with M. savini Hinton, M. intermedius (Newton) and M. majori Hinton. 4. CONCLUSION From the occurrences here quoted it is seen that the recorded chronological range of this species is from the Middle Villafranchian to the Antepenultimate (Cromerian, s.l.) Interglacial, although Van der Sluijs (1957, 258) records it also from the Lower Villafranchian of Germany, without specifying a locality. This duration is supported by Kretzoi's statement (quoted by Oakley, 1966, 33) that in Hungary Mimomys was the dominant vole in the Villafranchian, s.1., stage, declined during the Cromerian and had disappeared by about the Elsterian. The latter has been correlated with the Lowestoft cold stage of East Anglia (West, 1963, table VII, 168). It should be noted, however, that there appears to be no certain evidence of M. pliocaenicus having lived in Britain during the Antepenultimate Interglacial since the remains from Ostend and the Forest Bed Series at East Runton are considered to have been derived from older deposits. In this connection it is significant that Hinton has stated (1954), "Mimomys pliocaenicus est, en verite, l'un des especes Qui caracterisent notre Norwich Crag anglais et sa duree dans le temps n'est, certes, pas considerable'. Funnell & West (1962, 126-9) consider that the Upper Shell Bed of Larwood & Martin was accumulated during the time represented by Zone L4a of the Baventian Stage of the Lower Pleistocene. The pollen evidence from this deposit, obtained by the former authors, indicates forests with Pine (Pinus) dominant, and Spruce (Picea), together with Alder (Alnus), grasses and open heath vegetation, under climatic conditions of oncoming cold, relative to the probably cool temperate climate of the preceding Zone L3 (Antian Stage). It is regrettably not yet possible to make any inferences concerning the ecology of M. pliocaenicus, but the voles may have lived in one or more of these plant communities, although the mammalian remains of the Norwich Crag appear to represent a long period of time and a number of biotopes. ACKNOWLEDGMENTS The present writer is indebted to Mr. H. E. P. Spencer for permission to examine these fossils, Professor K. Kowalski, Director of the Institute of Systematic Zoology, Cracow, Poland, for information and references concerning some foreign localities, Mr. P. E. Long, who has supplied additional information on the provenance of these specimens and Miss D. J. Mace of the Geology Department, Queen Mary College, for assistance in drawing the illustrations.
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REFERENCES FEJFAR, O. 1961. Review of Quaternary Vertebrata in Czechoslovakia. Wydaw.Juine, 34, 109-18. FORSYTH-MAJOR, C. J. 1902. On some Jaws and Teeth of Pliocene Voles (Mimomys gen. nov.) from the Norwich Crag at Thorpe, and from the Upper Val d'Arno. Proc. zoot. Soc. Lond., 1, 102-7. FRIANT, M. 1953. Presence d'un Rongeur du Quaternaire ancien, Ie Mimomys pliocaenicus Maj., en France mediterraneenne (Sete, Herault). c. r, hebd Seanc, Acad. sa., Paris, 236, 730--2. - - - - . 1954. Une faune du Quaternaire ancien, en France mediterraneenne (Sete, Herault). Annis Soc. geol. N., 73, 161-70. FUNNELL, B. M. & R. G. WEST. 1962. The Early Pleistocene of Easton Bavents, Suffolk. Q. JI geol. Soc. Lond., 118, 125-41. HELLER, F. 1936. Eine oberpliozane Wirbeltierfauna aus Rheinhessen. Nelles lb. Miner. Geol. Paldont, Bei/Bd., 76, B, 99-160. - - - - . 1963. Eine altquartare Wirbe1tierfauna des unteren Cromerium aus der nordlichen Frankenalb. Neues lb. Miner. Geol. Paldont, Abh., 118, 1-20. HINTON, M. A. C. 1926. Monograph of the Voles and Lemmings (Microtinae), Living and Extinct. Vol. I, Brit. Mus. (Nat. Hist.), London. - - - - . 1954. Note sue Ie Mimomys pliocaenicus Maj. de Sete, Annis Soc. geol. N., 73,170. KORMOS, T. 1931. Oberpliozane Wiihlmause von Seneze (Haute-Loire) und Val d'Arno (Toscana). Abh. schweiz. paldont, Ges., 51, 1-14. KOWALSKI, K. 1958. An Early Pleistocene Fauna of Small Mammals from the Kadzielnia Hill in Kielce (Poland). Acta palaeont, pol., 3, 1-47. LARWOOD, G. P. & A. J. MARTIN. 1954. Stratigraphy and Fauna of the Easton Bavents Cliff Sections near Southwold, Suffolk. Trans. Suffolk Nat. Soc., 8, 157-71. MEHELY, L. VON. 19/4. Fibrinae Hungariae. Die Tertiaren und Quaternaren Wurzelzahnigen Wiihlmause Ungarns. Annis hist.mat, Mus. natn. hung., 12, 155-243. NEWTON, E. T. 1908. Note relative il. des fragments fossiles de petits vertebres trouves dans les depots pliocenes de Tegelen-sur-Meuse. Bull. Soc. beige Geol. Paleont. Hydrol., 21, 591-6. OAKLEY, K. P. 1966. Discovery of Part of Skull of Homo erectus with Buda Industry at Vertesszollos, North-West Hungary. Proc, geol. Soc., 1630, 31-4. RUTTEN, L. M. R. 1909. Die diluvialen Sdugetiere der Nieder/ande, pp. 88-9 and Tafel I, fig. 18. Berlin. SCHREUDER, A. 1949. Nieuwe Zoogdierfossielen uit de Tegelse Klei. Geol.rmijnbouwk, Biblphie Ned.slndie, ll, 115-26. SPENCER, H. E. P. 1963. The Prehistoric Deer of the East Anglian Crag Deposits. Trans. Suffolk Nat. Soc., 12, 262-6. - - - - . 1964. The Contemporary Mammalian Fossils of the Crags. Trans. Suffolk Nat. Soc., 12, 333-44. THALER, L. 1955. Sur l'age pliocene de 1a faune des grottes du Lazaret (Sete, Herault). c. r. hebd. Seanc, Acad. Sci., Paris, 241, 433-5. TUCCIMEI, G. 1893. Resti di Arvicola nel Pliocene Lacustre della Sabina. Mem. Accad. pont. Nuovi Lincei, 9, 35-45. VAN DER SLUIJS, G. K. 1957. The Use of Voles for Stratigraphic Correlations. Geol.mijnbouwk, Biblphie Ned-Lndie, 19, 257-9. VIRET, J. 1954. La loess Ii banes durcis de Saint Vallier (Drome) et sa faune de Mammiferes viliafranchiens. Nouv. Archs Mus. Hist, nat. Lyon, 4, 1-200, 33 pis. WEILER, W. 1952. Pliozan und Diluvium im siidlichen Rheinhessen. 1 Teil. Notizbl. hess. Landesamt, Bodenforsch, Wiesbaden, 6, 3, 147-70. WEST, R. G. 1963. Problems of the British Quaternary. Proc, Geol. Ass., 74,147-86.
J. N. Carreck Department of Geology Queen Mary College Mile End Road, London E.!
page 491 493 494 495 495 496
INTRODUCTION DESCRIPTION DISTRIBUTION CONCLUSION ACKNOWLEDGMENTS REFERENCES
ABSTRACT: A small series of Microtine teeth from the Lower Pleistocene Norwich Crag of Easton Bavents, Suffolk, preserved in Ipswich Museum, is described and assigned to the vole Mimomys, some, if not all, being of M. pliocaenicus (ForsythMajor), a new record for this locality. Selected specimens are figured. The stratigraphical and geographical distribution of the species in Britain and on the Continent are discussed in relation to its presence at this site, and the local environment in which it I ived is considered.
1. INTRODUCTION the recent publication of Funnell & West's paper on the Early Pleistocene deposits of Easton Bavents, near Southwold, Suffolk (1962), it may be of value to place on record the occurrence of some Microtine remains in the Norwich Crag of that locality collected by the brothers D. and P. Long of Lowestoft, and Mr. R. J. Winyard of Walberswick, Suffolk, and preserved in Ipswich Corporation Museum. These were kindly submitted to the author for determination by Mr. H. E. P. Spencer, when geologist of that museum. The precise horizon from which the fossils were obtained is not certainly recorded, but all or most came from the Upper Shell Bed described by Larwood & Martin (1954, 166). Easton Bavents cliff is a significant locality for larger mammalian remains in the Norwich Crag (Larwood & Martin, 1954, 166-8; Spencer, 1963, 264-5 and 1964, 344; Spencer, in Funnell & West, 1962, 137). These authors record a Baleen whale, Balaena sp.; 'Mastodon' sp.: the Southern Elephant, Mammuthus (Archidiskodon) meridionalis (Nesti); cf. 'Rhinoceros' sp.; the zebra, Equus (Hippotigris) robustus Pomel; Hippopotamus sp.; and the deer Euctenoceros sedgwicki (Falconer), Eu. falconeri (Dawkins), and Megaceros verticornis (Dawkins). The only previously published record of Microtine remains from the Norwich Crag of Easton Bavents, IN VIEW OF
491
492
J. N. CARRECK
5
2
7
8 Fig. 1. Teeth of the vole Mimomys from the Norwich Crag of Easton Bavents, Suffolk 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.
Left first lower molar (rnr), of Mimomys pllocaenicus, Crown (occlusal) view, x 8.0. The same specimen, Inner (lingual) view, x 8.0. Another left mi (incomplete), M. pliocaenicus, Occlusal view, X8.5. The same specimen, Outer (labial) view, X 8.0. A further example of the left rni (incomplete), Mv pliocaenicus. Occlusal view, X 7.5. The same specimen. Inner view, x7.5. Right first upper molar (m-j, Mimomys sp, indet. Occlusal view, X8.5. The sarne specimen. Inner view, x8.5. Right lower incisor (incomplete), Mimomys sp, indet. Inner view, X 2.0. Left upper incisor (incomplete), Mimomys sp, indet. Inner view, X 2.0.
MICROTINE REMAINS, NORWICH CRAG, SUFFOLK 493
however, is of some incisors in the W. M. Crowfoot and Searles V. Wood collections at Norwich Castle Museum, and others obtained in 1951-2 by the brothers Long (Larwood & Martin, 1954, 168). These teeth are unfortunately not diagnostic. 2. DESCRIPTION The remains examined by the present writer consist of lustrous, mineralised molar and incisor teeth, the former showing dark brown enamel and light brown dentine and cement, and the incisors showing dark brown enamel and light brown dentine. Some are a little rolled, but Mr. Spencer (verbal information, May 1964) considers that they are probably not derived from a deposit other than the local Norwich Crag as he knows of no apparent source. Mimomys pliocaenicus (Forsyth-Major). Four left m, One of these molars (Fig. I, Nos. 5 and 6) is of an adult vole in which the posterior loop is broken away, one root is developed but partly broken off, and the other is lost. Maximum length preserved: 2.75 mm.; maximum width: 1.4 mm. The inner portion of the third outer enamel infold is completely insulated. Another example (Fig. 1, Nos. 3 and 4) is very similar as regards the form of the occlusal surface, but the upper part of the posterior loop is broken away, the root below is preserved and the anterior root is lost. Maximum length: 3.4 mm.; maximum width: 1.4 mm. In another specimen the roots have not yet appeared. The occlusal surface is deeply worn, and the posterior loop and second inner salient angle have been lost. The last specimen (Fig. I, Nos. 1 and 2) shows the distinctive enamel islet, large and completely developed, in the anterior loop. The tooth is perfect and the two completely formed roots are preserved. Therefore this tooth is of an adult individual. Maximum length: 3.3 mm.; maximum width: 1.4 mm.; maximum height: 4.3 rom. M. cf. pliocaenicus. One left m 3, one right m3 and a left m., In the left m3 root development has begun and an enamel islet is present. The right m 3 also shows root development and is very similar to the last. The left m I is very incomplete, being the whole base of the tooth but without roots and only possessing the anterior part of the occlusal surface (anterior loop), although this shows the enamel islet and a slight prism fold. Mimomys sp. indet. One left rn-, four right m! (Fig. 1, Nos. 7 and 8), one right m 2 , one right m, and eleven parts of incisors (Fig. I, Nos. 9 and 10). The upper molar shown in 7 and 8 of Fig. 1 has a maximum length of 2.9 mm., and a maximum width of 1.6 mm. The lower incisor shown in No.9 has a length of 14.25 mm. and a height of 1.6 mm., whilst the upper incisor in No. 10 has a length of 10.1 mm. and a height of 1.9 mm. These remains are not sufficiently diagnostic for precise determination, but may well be of M. pliocaenicus.
494
1. N. CARRECK
3. DISTRIBUTION This species of vole was originally described from the Late Villafranchian lacustrine deposits of the Val d'Arno, Tuscany, Northern Italy (ForsythMajor, 1902, 103) and also in that country remains are known from the Lower Pleistocene lacustrine marl at Bocchignano in Sabino (Tuccimei, 1893), probably referable to this species or a near ally. In France it is known from the Late Villafranchian of Seneze in Haute-Loire (Kormos, 1931, 1-5), and the Middle Villafranchian deposits of Sete, Herault (Friant, 1953; Hinton, 1954) have yielded remains then ascribed to this species, but now referred to a much more primitive species, M . occitanus Thaler. This author considers it to be of Pliocene age (Thaler, 1955). Other French occurrences are the 'Gunzien' (Villafranchian or earlier) of Perrier, Central France (Friant, 1954, 166-8), and the Villafranchian of Saint-Vallier (Viret, 1954). In Holland remains are known from the Tiglian (Middle Villafranchian) of Tegelen (Newton, 1908; Schreuder, 1949, 116-17), Gorinchem (Gorkum) (Riitten, 1909; Hinton, 1926, 358), Eindhoven (Van der Sluijs, 1957, 257) and Veghel (Van der Sluijs, op. cit., 258), and the Lower Taxandrian (Late Villafranchian) of Raamsdonkveer ? and Breda ? (Van der Sluijs, 1957, 257), and in Germany from the Early Gla ciat ion Interstadial/Earl y Glaciation II deposits of Gundersheim (Heller , 1936), the Lower Cromerian, s.l., of Deinsdorf (idem, 1963), and Lower Pleistocene deposits at Hohensiilzen (Weiler, 1952). In Poland others have been described from deposits probably of the Early Glaciation Interstadial at Kadzielnia Hill in Kielce (Kowalski, 1958, 29-33, and in lit., 1964). In Czechoslovakia Fejfar (1961) has recorded this species from lacustrine tuffaceous deposits of the latter part of the Villafranchian at the Hajnacka II site. In Hungary remains have been found in the Middle /Late Villafranchian deposit s of Beremend (Mehely, 1914), and the Antepenultimate Interglacial (Cro merian, s.l.) deposits at Berg NagyHarsany and Csarnota (Mehely, 1914), Villany-Kalkberg (Ko walski, 1958, 6), and in Rumania in deposits of the Early Glaciation II (Late Villafranchian) at Puspokfurdo (Kowalski, 1958). The English occurrences are in the Norwich Crag of Thorpe, near Aldeburgh, Suffolk (Forsyth-Major, 1902), Yam Hill, near Southwold, Suffolk (Hinton, 1926, 355 and 361), Covehithe, near Southwold (undescribed material in Ipswich Museum, determined by the present writer), and Bramerton, near Norwich, Norfolk (Hinton, 1926, 361) ; the Upper Freshwater Bed of Ostend, near Bacton, Norfolk (Hinton, 1926, 355), probably derived from the Norwich Crag; in derived pieces of clay in the middle or estuarine division of the Forest Bed Series (Antepenultimate Interglacial) at East Runton, Norfolk (Hinton, 1926, 358 and 376), and in Weybourne Crag at that locality (Hinton, 1926). In the Norwich Cra g it occurs alone or possibly in association with Mimomys newtoni (Forsythe
MICROTINE REMAINS, NORWICH CRAG, SUFFOLK 495 Major), but in the Weybourne Crag of East Runton it is found with M. savini Hinton, M. intermedius (Newton) and M. majori Hinton. 4. CONCLUSION From the occurrences here quoted it is seen that the recorded chronological range of this species is from the Middle Villafranchian to the Antepenultimate (Cromerian, s.l.) Interglacial, although Van der Sluijs (1957, 258) records it also from the Lower Villafranchian of Germany, without specifying a locality. This duration is supported by Kretzoi's statement (quoted by Oakley, 1966, 33) that in Hungary Mimomys was the dominant vole in the Villafranchian, s.1., stage, declined during the Cromerian and had disappeared by about the Elsterian. The latter has been correlated with the Lowestoft cold stage of East Anglia (West, 1963, table VII, 168). It should be noted, however, that there appears to be no certain evidence of M. pliocaenicus having lived in Britain during the Antepenultimate Interglacial since the remains from Ostend and the Forest Bed Series at East Runton are considered to have been derived from older deposits. In this connection it is significant that Hinton has stated (1954), "Mimomys pliocaenicus est, en verite, l'un des especes Qui caracterisent notre Norwich Crag anglais et sa duree dans le temps n'est, certes, pas considerable'. Funnell & West (1962, 126-9) consider that the Upper Shell Bed of Larwood & Martin was accumulated during the time represented by Zone L4a of the Baventian Stage of the Lower Pleistocene. The pollen evidence from this deposit, obtained by the former authors, indicates forests with Pine (Pinus) dominant, and Spruce (Picea), together with Alder (Alnus), grasses and open heath vegetation, under climatic conditions of oncoming cold, relative to the probably cool temperate climate of the preceding Zone L3 (Antian Stage). It is regrettably not yet possible to make any inferences concerning the ecology of M. pliocaenicus, but the voles may have lived in one or more of these plant communities, although the mammalian remains of the Norwich Crag appear to represent a long period of time and a number of biotopes. ACKNOWLEDGMENTS The present writer is indebted to Mr. H. E. P. Spencer for permission to examine these fossils, Professor K. Kowalski, Director of the Institute of Systematic Zoology, Cracow, Poland, for information and references concerning some foreign localities, Mr. P. E. Long, who has supplied additional information on the provenance of these specimens and Miss D. J. Mace of the Geology Department, Queen Mary College, for assistance in drawing the illustrations.
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J. N. Carreck Department of Geology Queen Mary College Mile End Road, London E.!