616 NOTES FROM T H E MAMMAL SOCIETY-NO. 53

REFERENCES Brown, L. E. (1969). Field experiments on the movements of Apodemus sylvaricus L. using trapping and tracking

Emlen, J. T., Hine, R. L., Fuller, W. A. & Alfonso, P. (1957). The use of dropping boards for population studies of

Randolph, S. E. (1973). A tracking technique for comparing individual home ranges of small mammals. J. Zool., Lond.

Wolton, R. J. (1985~). The ranging and nesting behaviour of Wood mice, Apodemus sylvaticus (Rodentia: Muridae), as revealed by radio-tracking. With an addendum by R. J. Wolton and B. J. Trowbridge: The effects of radio-collars on Wood mice, Apodemus sylvaticus. J . Zool., Lond. ( A ) M6: 203-224.

Wolton, R. J. (19856). A possible role for faeces in range-marking by the Wood mouse, Apodemus sylvaricus. Notes from the Mammal Society, No. 50. J. Zool., Lond. ( A ) 206: 286-291.

techniques. Oecologia 2: 198-222.

small mammals. J. Wild/. Mgmr 21: 300-314.

170 509-520.

Neolithic bats from Dowel Cave, Derbyshire D, W. Y ALDEN, Department of Zoology, University of Manchester, Manchester M 13 9PL

If the post-glacial history of British mammals is poorly documented (Yalden, 1982), then that of the bats is particularly poorly understood. Yet the history of the bat fauna is potentially very interesting; woodland species such as Myotis bechsteini and Plecotus spp. should have been much more numerous when woodland was more extensive, and various southern species such as P . austriacus and Rhinolophus ferrumequinum might have spread further north in warmer periods. In Poland, Woloszyn (1970) has documented numerous cave faunas, probably of Neolithic age, in which M. bechsteini was indeed very numerous (it is now as rare in Poland as in Britain). Myotis bechsteini is also reputed to be numerous in the Neolithic flint mines of Grimes Graves, Norfolk (Clarke, 1963), though no details of numbers seem to have been published.

A further site at which M. bechsteini has been reported is Dowel Cave, in the Carboniferous limestone of the Peak District (Bramwell, 1960a, 6 , 1977), but again no details have been published. This note concerns a re-examination of the chiropteran remains in the Dowel Cave fauna.

Site Dowel Cave lies near the village of Earl Sterndale in a dry valley, Dowel Dale, which is a side-

valley of the River Dove, at Nat. Grid. Ref. SK076676. The cave was excavated by the Peakland Archaeological Society in 1958-59, under the direction of Dr D. Bramwell. The main archaeo- logical interest was the presence of 10 or more human skeletons of Neolithic age (Bramwell, 1960a), but the deposits were up to 20 ft thick in places, and spanned a period from late Pleistocene onwards.

Material The vertebrate fauna recovered from the cave is extensive; Bramwell (1960b) lists 42 mammal species,

and a t least 51 bird species. The best-preserved material, including 10 bat skulls, comes from the Neolithic layer, a fine-grained silt. There are also 12 mandibles, and numerous post-cranial elements including 52

N O T E S F R O M T H E M A M M A L SOCIETY-NO. 5 3 617

humeri and 60 radii, as well as numerous metacarpals and phalanges. A few of the post-cranial remains come from earlier deposits, of Late Glacial and Mesolithic Age. Skulls and mandibles have been identified by reference to Miller (1912), and the Myotis have been checked against the reference material in the British Museum (Natural History). The humeri have been compared with Felten, Helfricht & Storch (1973), while the radii can readily be identified to family, and thereafter can only be roughly identified on size (Yalden, 1985).

Rhinolophus hippideros, Lesser horseshoe bat Three partial skulls, lacking the front teeth but with p4-m3 lengths of 4.28, 4.25 and 4.31 mm;

two right and one left mandible; 21 humeri, with the very characteristic distal articulation (see Felten et al., 1973; Yalden, 1985); and 19 radii, including 12 complete examples 34.0-36.3 mm long (2 = 35.4, S.D. 0.7).

All these specimens are unmistakably rhinolophid, but are far too small to belong to R. ferrumequinum.

Myotis nattereri, Natterer’s bat Seven partial skulls, with c-m3 lengths of approximately 5.63-6.08 mm (2 = 5.90, S.D. 0.14);

six lower jaws 6.33-c. 6.50 mm long; perhaps 28 humeri and 40 radii also belong to this species. Some of the skulls are lacking canines and other teeth, and the tooth row measurements for

them are taken from the alveoli, hence ‘approximately’ 5.63-6.08 mm. These skulls all have the ‘crowded’ third premolar which distinguishes M. nattereri from M . bechsteini (Miller, 1912; Yalden, 1985) and are also appreciably smaller than M. bechsteini. The lower jaws belong, on size, to M. nattereri as well. Attribution of the post-cranial remains is less certain. On the form of the elbow joint, the humeri all seem to be referable to Myotis (see Felten et al., 1973), but could, on size, be M. nattereri or M . daubentonii. Similarly, the radii could belong to the slightly smaller M. daubentonii or to the slightly larger M. bechsteini, but are most parsimoniously referred to M. nattereri. These three species overlap considerably in forearm length-33-40 mm, 38-45 mm and 36-43 mm, respectively; for 17 intact radii, the range of 35.3-39.8 (3 = 37.7, S.D. 1.4) clearly fits well with the forearm size of M. nattereri, allowing that the radius will be slightly shorter than the forearm.

Myotis daubenton& Daubenton’s bat Two right mandibles, c-m3 5.70, 5.80 mm. These mandibles are appreciably smaller than those of M. nattereri, but match M . daubentonii.

?Myotis mystacinus, Whiskered bat One radius, 30.5 mm long; two smaller humeri. It would be unrealistic to claim the identification of this species on such slender evidence. The

radius is apparently too small to belong to M. daubentonii; M. brandtii cannot be ruled out. Two of the humeri are appreciably shorter and more slender than those referred to M. nattereri, but could be any of the smaller Myotis sp.

618 NOTES FROM THE MAMMAL SOCIETY-NO. 5 3

?Plecoius sp., Long-eared bat One humerus, 24.1 mm long. The elbow joint of this bone appears to differ from those of Myotis, and matches the features

of Plecotus as shown by Felten et al. (1973). It is also slightly larger than the majority of specimens referred to M. nattereri from this collection.

Nyctalus leisleri, Leisler’s bat One humerus, 25.00 mm long. This humerus is stouter than any of the others, and has the prominent styloid process character-

istic of Nyctalus (Felten et al., 1973). However, it is much smaller than my reference specimen of N . noctula (30.04 mm long), and must be referable to N . leisleri.

Discussion Bramwell(196Ob) listed R. hipposideros, Plecotus auritus, Myotis bechsteini and M . daubentonii

as present in the Dowel Cave fauna, along with M. nattereri and Pipistrellus pipistrellus as probably present. There is no doubt about the presence of R. hipposideros, with three unmistak- able skulls and indications on postcranial elements of 11 individuals represented in total. It has not been possible to confirm the presence of M. bechsteini; two skulls, labelled ‘M. bechsteini’ and ‘ M . bechsteini? by Bramwell are, on checking, M . nattereri. Probably Bramwell was misled by the tables of measurements in Miller (1912), as initially I was myself. Miller gives measure- ments for maxillary and mandibular toothrows of Myotis which appear to span 1 z-m3 and 1 ,- m3 (because the tables for Barbastella and Plecotus specifically say ‘excluding incisors’, but none of the other genera are so labelled). Thus the range of maxillary toothrow given by Miller for M. bechsteini, 6-8-7.0 mm, corresponds to the estimated 1 2-m3 measurement of the Dowel Cave Myotis. In fact, checking back to the skulls which Miller himself measured, it is clear that the range 6.8-7.0 mm for Myotis bechsteini is for c-m3, and the Neolithic skulls are smaller than that. Myotis daubentonii is evidently also present in the Dowel Cave fauna, on the basis of the two mandibles. Myotis mystacinus and Plecotus sp. are possibly present. I am not clear on what grounds Pipistrellus was thought possibly present, but the small radius, here tentatively referred to M. mystacinus (a more likely inhabitant of caves) could have been the reason.

The contrast between this Neolithic fauna and the present fauna of the Peak District caves is notable. Recent records of hibernating bats, accumulated over 20 years (1966-1985) are com- pared with the apparent Neolithic fauna in Table I. At present, cave bats include four species of

T A B L E I Comparison of the Neolithic and Recent bat faunas in the

Peak District

Neolithic Author’s records, Dowel Cave 1966-85

Rhinolophus hipposideros 1 1 0 Myotis nattereri 20 21

M. mystacinuslbrandtii l? 44

Nyctalus leisleri I 0

M. daubentonii 2 17

PIecotus auritus I? 16

NOTES F R O M T H E M A M M A L SOCIETY-No. 53 619

Myotis and Plecotus auritus (cf. also Yalden, 1977), whereas the fossil fauna is dominated by R. hipposideros and M . nattereri. Coward & Oldham (1910) claimed that R . hipposideros was present in Dovedale, the Matlock area and in the nearby Alderley Edge mines, but gave no details of numbers of precise dates; there have been no records since their account (Ellison, 1959; Yalden, 1977; Clinging & Whiteley, 1980). Surprisingly, R. hipposideros has not been found in the Mesolithic fauna of Dog Hole Fissure, on the Derbyshire/Nottinghamshire border, reported recently by Stebbings in Briggs, Griffin, Stebbings & Watts (1984).

The predominance of R. hipposideros and M . nattereri in the fossil fauna may reflect the more wooded nature of the Peak District in Neolithic times (Hicks, 1971; Tallis & Switsur, 1983), but one might expect Plecotus also to have been common if that were the explanation.

The presence of N. leisleri in the fossil fauna is slightly anomalous, in that all the other species regularly hibernate in caves, which N. leisleri does not. However, it was presumably introduced into the cave as the prey of an owl, as were the very abundant small birds which also occur in the cave fauna. There are a few recent records of N. leisleri in the Sheffield area (Whiteley, 1984).

I thank J. E. Hill (B.M.(N.H.)) for his very helpful correspondence on the measurements of Myoris, and for access to the reference collection. Dr D. Bramwell kindly passed the material on to me, and discussed my findings. It is intended that the material be deposited in Stockport Museum, Vernon Park, Stockport, when re-examination and cataloguing are completed.

R E F E R E N C E S Bramwell, D. (1960~). The excavation of Dowel Cave, Earl Sterndale, 1958-59. Derbys. Arch. J . 79 (for 1959): 97-109. Bramwell, D. (19606). The vertebrate fauna of Dowel Cave. Peakland Arch. SOC. Newsl. 17: 9-12,23. Bramwell, D. (1977). Archaeology and palaeontology. In Limestones and caves of the Peak Districr: 263-291. Ford,

T. D. (Ed.). Norwich Geo. Abstracts. Briggs, D. J., Griffin, C. M., Stebbings, R. E. & Watts, C. M. (1984). Death, disarticulation and decay in an eight

thousand year old rockfall. In In rhe shadow of extinction: 55-74. Gilbertson, D. D. & Jenkinson, R. D. S. (Eds). Univ. of Sheffield: Dept. of Prehistory and Archaeology.

Clarke, R. R. (1963). Grimes Graves, Norfolk. London: HMSO. Clinging, V. & Whiteley, D. (1980). Mammals of the Sheffield area. Sorby Rec., Spec. Ser. No. 3: 1-48. Coward, T. A. & Oldham, C. (1910). The vertebrate fauna of Cheshire and Liverpool Bay. London: Witherby. Ellison, N. F. (1959). A checklist of the fauna of Luncushire and Cheshire: Mammalia, Reptilia, Amphibia. Liverpool:

Felten, H., Helfricht, A. Kc Storch, G. (1973). Die Bestimmung der europaischen Fledermause nach der distalen Epiphyse

Hicks, S. P. (1971). Pollen-analytical evidence for the effect of prehistoric agriculture on the vegetation of north

Miller, G. S. (1912). Catalogue of rhe mammals of western Europe. London: British Museum (Nat. Hist.). Tallis, J. H. & Switsur, V. R. (1983). Forest and moorland in the south Pennine uplands in the mid-Flandrian period. I.

Whiteley, D. (1984). Leisler’s bat in the Sheffield area-a second record. Sorby Rec. 2 2 27-28. Wotoszyn, B. (1970). [The Holocene chiropteran fauna from the Tatra caves]. Folia quatern. 3 5 1-52, [In Polish]. Yalden, D. W. (1977). Mammals. In Limestone and caves of the Peak District: 250-251. Ford, T. D. (Ed.). Norwich:

Yalden, D. W. (1982). When did the mammal fauna of the British Isles arrive? Mumm. Rev. 1 2 1-57. Yalden, D. W. (1985). The identification of British bats. London: Occ. Publ. Mammal Soc.

Lancs. Chesh. Fauna Committee.

des Humerus. Senckenberg. biol. 54: 291 -297.

Derbyshire. New Phytol. 7 0 647-667.

Macrofossil evidence of the former forest cover. J . Ecol. 71: 585-600.

Geo. Abstracts.