CRANIUM, 17, 2 - 2000

New herpetologicalrecords from the Middle Pleistocene Boxgrove Hominid Site, England

J.+Alan Holman

Summary

Additionalherpetological materialfrom the Middle Pleistocene Boxgrove hominidsite, West Sussex, England, has provided the first records from the site of two salamanders (crested newt, Triturus cristatus, and palmate newt,

and Rana and Triturus helveticus), two anurans (common parsley frog, Pelodytes punctatus, water frog ( [ridibunda] sp.),

well additional records of the nine one snake (smooth snake, Coronella austriaca) ; as as previously reported of herpetological species by Holman(1999). The total Boxgrove herpetofauna now consists three salamanders, seven of Pleistocene in Isles. anurans, two lizards, and two snakes, forming one the largest herpetofaunas the British

from but Pelobates All 14 herpetological species Boxgrove are presently living, three species: spadefoot, fuscus; that do common parsley frog, Pelodytes punctatus;and moor frog, Rana arvalis are continental species not presently the indicates climate that is similar the in the occur in Britain. As a whole, herpetofauna a to present one Boxgrove

The the of shallow wet a area. herpetological species indicate presence a pond; a or damp grassy area; moderately terrestrial and rather habitat. moist, well-vegetated, area; a open, sandy

Samenvatting

de Midden Pleistocene hominidsite in West heeft Nieuw herpetologisch materiaal van Boxgrove Sussex, Engeland,

zwemvoetsalamander Triturus twee Triturus cristatus, en twee nog watersalamanders (kamsalamander helveticus), waterkikker kikkers (de modderspringer of groengestipte kikker Pelodytes punctatus en de groene Rana [ridibunda] Coronella Verder aantal stukken sp.), en een slang (gladde slang austriaca) opgeleverd. zijn er nog een aangetroffen bestaat de al eerder doorHolman soorten. De van van negen (1999) gerapporteerde complete herpetofauna Boxgrove één de nu uit drie salamanders, zeven kikkers, twee hagedissen, en twee slangen, en vormt zo van grootste komen Pleistocene herpetofauna’s van de Britse eilanden. Alle veertien herpetologische soorten van Boxgrove

echter Pelobates kikke tegenwoordig nog voor; drie soorten (Europese knoflookpad fuscus, groengestipte rPelodytes het punctatus, heikikker Rana arvalis)zijn continentale soorten die tegenwoordig niet in Verenigd Koninkrijk

de klimaat dat overeenkomt met het klimaat van het voorkomen. Als geheel wijst herpetofauna op een huidige

gebied rond Boxgrove. De herpetologische soorten tonen de aanwezigheid aan van een ondiep water, een nat of

land vochtig grasland, een tamelijk vochtig, dicht-begroeid hoger gelegen en een vrij open, zanderig gebied.

Introduction Holman (1992,1998,1999) previously published the from reports on Boxgrove herpetofauna The Middle Pleistocene homonid locality (fig. 1) material collected to 1992. These fossils up repre- at Eartham Quarry, Boxgrove, West Sussex, of five sent at one species salamander, anurans, England, British National Grid Reference two lizards, andone snake. Since that time, much NumberSU920085, is one of the most publicized additional herpetological material has been Quaternary sites in the Old World because the collected at Boxgrove, and this has provided human artifacts and skeletal material suggest records of and two salamanders, two anurans, that it is the earliest human occupational site in one snake that are new to the fauna; as well as Europe (e.g. Roberts et ah, 1995; Roberts & Parfitt, additional material of previously reported 1999). Boxgrove is correlated with the Interglacial Boxgrove amphibians and reptiles. of the IV stage of the Cromerian Complex

et al., a Dutch/European sequence (Roberts 1995). This paper provides (1) systematic annotated In the site is correla- list of the material from a narrower sense, Boxgrove new herpetological

ted with the Miesenheim I Site in Germany (Kolf- Boxgrove, (2) a checklist of the entire herpeto- schoten known and & Turner, 1996), an important fauna as it is at present, (3) some continental herpetological site (Holman, 1998). comments on the significance of the herpeto- Roberts & Parfitt (1999) have provided the latest fauna.

information (previous to the present report) on Fossil elements are listed bybulk sample number, the fauna of Boxgrove. the contextual details of which are in Parfitt

(1999).

112 New herpetological records from the Middle Pleistocene Boxgrove HominidSite, England

Fig. 1 Location map of Amey’s Eartham Pit (A.E.P) where the Boxgrove herpetofauna was collected

Eartham verzameld Plattegrond van Amey’s Pit (A.E.P), waar de Boxgrove herpetofauna is

113 CRANIUM, 17, 2 - 2000

from and Annotated list of the new Boxgrove first record of the species Boxgrove only the fourth record from the Pleistocene of Britain herpetological material (Gleed-Owen, 1999; Holman, 1998). New mate- Order Caudata Scopeli, 1977 rial: two trunk vertebrae (54 and 690).

Triturus vulgaris Group Family Salamandridae Goldfuss, 1820 Triturus helveticus or vulgaris Genus Triturus Rafinesque, 1815 Palmate Newt or Smooth Newt Alpine Newts These small newt fossils do not belong to the T. This contains salamanders of small or genus cristatus but group, they are fragmentary enough moderately small size that usually have two so thatthey lack thecharacters that enable one to distinct phases: an eft phase that is terrestrial, has identify them as either T. helveticus or T. vulgaris. a rough skin, and lacks a dorsal crest or "tail fin"; Holman (1992, 1998) previously identified “T. and that has smooth a newt phase is aquatic, a from Site. helveticus or T. vulgaris” the Boxgrove fin." skin, and has a crest and sometimes a "tail New material: five vertebrae (40,78,485,590, and occurs Scandan- The genus presently in England, 773). avia, continental Europe, andAsia Minor around

Triturus the Black Sea and to the westward part of the vulgaris Group helveticus Caspian Sea eastward to the Ural Mountains. Triturus (Razoumowsky, 1789) Palmate Newt of Triturus Two species groups are presently

recognized: the Triturus cristatus group and the T. these New record from the site. The vulgaris group (Frost, 1985). Osteologically, species palmate

different that could be newt is a small that is smooth skinnedand groups are so they easily species Both has dark, webbedhind feet. The separated into two distinct genera. groups strongly palmate

have been recorded from the Pleistocene of newt is presently absent from Ireland but it

Europe (Holman, 1998). All of the three modern occurs in Great Britain, continental western

northern to southern species that presently occur in Britain have now Europe from Germany been recorded from the Boxgrove Site. France and northern Iberia, and east to Poland and the Czech Republic. Triturus helveticus IS Species group undesignated more terrestrial than T. cristatus, but it is more Triturus sp. aquatic than T. vulgaris. It is rather ubiquitous in Alpine newt its choice of breeding places. Holman (1998) gave

Damaged elements that cannot be referred with vertebral characters to separate T. helveticus from first certainty to either the Triturus cristatu s or the T. T. cristatus and T. vulgaris. This is the unequi-

T. here. Holman vocal record of this from the vulgari s group are included (1992, species Boxgrove from the Site and the second record for the Pleisto- 1998) previously reported Triturus sp. only of Boxgrove Site. New material: jaw fragment (182), cene Britain (Gleed-Owen, 1999). New mate- partial femur (621), eight fragmental vertebrae rial: two trunk vertebrae (679 and 741).

(235, 238, 278, 281, 663, and 741-743). Triturus vulgaris Group

Triturus cristatus Group Triturus vulgaris (Linnaeus, 1758)

Triturus cristatus Laurenti, 1758 Smooth Newt

Crested Newt The smooth newt is small, smooth-skinned newt

that lacks the webbed hind feet of T. helveticus.

Triturus New species record from the site. The crested vulgaris presently occurs in Ireland, that Great Britain, almost all of and the newt is a large rough-skinned taxon is Europe,

presently absent in Ireland but occurs in Great western part of Asia. The smooth newt is more

Britain and from Europe eastward to the central terrestrial than most other species of European

part of the former Soviet Union. Thecrested newt Triturus and tends to be a lowland species that

favors slow water where lives in a wide of moist habitats. It quiet or very flowing variety

thereis abundantvegetation. Theterrestrial form normally breeds in still, shallow water with an

the where it abundanceof Holman usually stays near breeding pond aquatic vegetation. (1998) characters that hides under rocks or other flattened objects. gave vertebral separate Triturus of the skull, from T. cristatus and T. helveticus. This Holman (1998) gave characters vulgaris

humeri, and vertebrae that separate T. cristatus newt has previously been reported from the from other species of European newts. This is the Boxgrove Site (Holman, 1992,1998) and has been

114 New herpetological records fromthe Middle Pleistocene Boxgrove Hominid Site, England

rather frequently reported from the Pleistocene New material: two right scapulae (772), right of from and Holocene Britain (Gleed-Owen, 1999; scapula (774), two scapulae among samples

Holman, 1998). Newmaterial: six trunk vertebrae 428-434, three more scapulae (286, 456, and 698), humeri (53, 91, 228, 297, 635, and 735). two left humeri (429), three left (280, 672, and 735), right humerus (799), five humeri (106, Family Pelodytidae Hogg, 1838 159,428,741, and 773), three left ilia (110,122, and Genus Pelodytes Bonaparte, 1838 279), and seven sacra (270,434,735, 742, 744, 749, Parsley Frog and 772). New material of probable true toad (cf.

bears two two humeri and and a tibiofibu- The genus only species, Pelodytes Bufo sp.): (73 78), punctatus and Pelodytes caucasicus (Frost, 1985; lar fragment (361). Duellman, 1993). Only the former species is Bufo bufo (Linnaeus, 1758) known as a Pleistocene fossil. The ilium of this Common European Toad is similar toPelobates of the genus generally fuscus Pelobatidae that the of the is the and of the family except edge Bufo bufo largest most common

dorsal acetabular expansion is slightly convex three European species of true toads. This taxon rather than slightly concave. This character tendsto be uniformly brownish in color. Bufo bufo

to be constant. is found in for Ireland and appears remarkably Europe (except some of the Mediterranean islands) east to Lake Baikal Pelodytes punctatus (Daudin, 1802) in southern Siberia; in the Caucasus; northern Common Parsley Frog Asia Minor and northern Iran. It also occurs in

northwestern Africa. This species is found in a

New species from the site. This species is a small, wider variety of habitats than other European with flat head and The toads. It has been recorded the agile frog a big eyes. previously from

occurs in Site Characters common parsley frog presently Boxgrove (Holman, 1992, 1998). Belgium through France to Spain and extreme for distinguishing the ilia of B. bufo from the other

in northwestern Italy. This taxon is principally a European species are given Holman (1998).

terrestrial form that rather New material: left ilium nocturnal, prefers (from among samples damp, well-vegetated habitats. Holman (1998) 428-434), nine left ilia (44, 238, 428, 438, 442, 476, discussed the of Pelobates 593, 639, and ilium separation punctatus 741), right (from among

from the other in the P. cauca- ten more ilia only species genus, samples 428-434), right (215, 428, and sicus on the base of ilial characters. This is only the 434,476, 548,581, 656, 665, 695, 739). second record of this species from the fossil Bufo calamita Laurenti, 1768 record of England, the first record being from the Natterjack Toad early Middle Pleistocene site at Westbury-Sub- Modern toads be Mendip Cave, Somerset, England (Gleed-Owen, natterjack may distinguished

1999; Holman, 1993, 1998). New material: left from B. bufo on the basis of the bright yellow ilium (131), two right ilia (434 and 778). stripe thatruns down the middleof the back. Bufo

calamita tends to move about by making mouse- Family Bufonidae Gray, 1825 like dashes rather than merely walking or Genus Bufo Laurenti other toads. hopping as in European Presently True Toads the natterjack occurs in western and northern

The consists of as far east western Russia and inclu- very large genus Bufo plump, Europe as that have of of and Ireland. In the warty, short-legged anurans a pair ding parts England nort- pronounced paratoid glands behind the head. hernpart of its range this species isusually found dunes the Two hundredand eleven living species are recog- in sandy areas including by sea. In nized The other of its it is in its (Duellman, 1993). genus is presently parts range more ubiquitous cosmopolitan except for the arctic regions, New habits. It has previously been recorded from Guinea, and Australia and nearby islands. The Boxgrove (Gleed-Owen, 1999; Holman, 1992, modern forms are mainly terrestrial, although 1998). New material: left ilium (221).

fossorial. The three modern some are European Family Ranidae Rafinesque-Schmaltz, 1814 species have all been identified from the Pleisto- Genus Rana Linnaeus, 1758 cene of the region. True Frogs

Bufo sp. The another genus Rana is extremely large taxon. True Toads They are typically long-legged frogs with smooth skins, webbed feet, and prominent tympanni.

115 CRANIUM, 17, 2 - 2000

Two-hundred and twenty-three living species rostrum. The present distribution of the moor The Rana is in northeastern France, Nether- are recognized (Duellman, 1993). genus frog Belgium, Finland is presently cosmopolitan except for southern lands, Germany, Denmark, Sweden, South America and most of Australia. In Europe south to the Alps, northern Yugoslavia, northern arvalis found the Rana contains referred to as Romania, and east toSiberia. Rana is genus species meadows and fields and wetland "water frogs" that are mainly aquatic and often in moist in have olive in their color and areas such and fens. Holman green or pattern as bogs (1992,1998) from the "brownfrogs" that are more terrestrial, brownish has previously identified R. arvalis

and often have dark face mask. Some Site. Holman out a in color, a Boxgrove (1998) pointed diag- of the nostic character the ilium of Rana New important osteological characteristics on arvalis. in Holman material: ilium genus Rana were given (1998). right (717). cf. Rana cf. Rana arvalis 1842 Rana sp. Nilsson, Probable True Frog Probable Moor Frog New material: partial femur (79). New material: two right ilia (78 and 263).

indet. Rana 1758 Rana sp. temporaria Linnaeus, True Frog Common Frog

The the It is not known whether these elements represent common frog is most widespread anuran

is water frogs or brown frogs. New material: in Britain and Europe today and often themost

two abundant anuran in terrestrial habitats in scapula (from among samples 428-434), moist, has scapulae (670 and809), six left humeri(72,75,112, central andnorthern Europe. This brown frog 118,129, and 696), five right humeri (75,129,156, a face mask and a small, soft metatarsal tubercle

773, and805), two left ilia (582), tworight ilia (45), that distinguishes it from the moor frog (Rana and often blade- fourteen left ilia (46, 77, 82, 91, 95,157,193, 200, arvalis) which has a large, hard,

287,349,578,603,662, and 841), eighteen right ilia like tubercle. Presently, Rana temporaria occurs (126,218,226,246,251,263,271,358,373,476,548, throughout Britain and Europe east to the Urals, 593, 596, 628-629, 679, 695, and 809), and three but excluding most of Iberia, much of Italy, and

the southern Balkans. It is the most sacra (141, 415, and 584). probably terrestrial of all of the European Rana species and Water Frog Species Group in found the many areas is in water only during

difficult to to It is of the most cold-tole- Waterfrogs are very identify species breeding season. one

(Sanchiz, 1998) and I have notbeen ableto deter- rant species in Europe. The common frog has mine the waterfrog species represented at the previously been identified from the Boxgrove Boxgrove Site. Sanchiz (1998) refers to these inde- Site (Gleed-Owen, 1999; Holman, 1992, 1998).

terminatewater frogs as follows. Holman (1998) gives ilial characters for the iden- tification of Rana temporaria. New material: two Rana (ridibunda) sp. left ilia (476 and 772), three right ilia (68,112, and Indeterminate European Water Frog 672).

Order Squamata Oppel, 1811 from Holman New Rana species group the site. (1998) has indicated how the ilia of European Lacertidae 1831 water frogs usually differ from those of brown Family Bonaparte, Genus Lacerta Linnaeus, 1758 frogs. Normally, one would expect that the Lizards of water would indicate the Common Eurasian presence frogs

of a body of freshwater near presence permanent has about This genus 43 living species (Frank & the fossil site. Water frogs as such have not previ- Ramus, 1995) andhas most of the general charac- ously beenreported from the Boxgrove Site. New ters of the family. Lacerta is known from most of material: four left ilia (35, 91, 250, and 580), and the Palearctic for cold Estes except very areas. two right ilia (519 and 644). (1983) has given important osteological charac- for the of the Brown Frog Species Group ters genus. Species genus Lacerta are

often divided two the into groups, larger, more Rana arvalis Nilsson, 1842 robust "green lizards" that tend to live in areas Moor Frog with dense vegetation and the "small lacertas"

the Podarcis that tend to be Moor frogs are a brown frogs with a face mask. (including genus ) habits. This taxon has relatively short legs and a pointed ubiquitous in their

116 New herpetological records from the MiddlePleistocene Boxgrove HominidSite, England

and Lacerta vivipara Jaquin, 1887 188, 286,476, 634, 672, 741, 743, 787, 809), six vertebrae and Viviparous Lizard fragmentary (91,126,287,591,654, 683), and three osteoscutes (471, 485, and 624). New positive identification. The viviparous

the in Colubridae 1811 lizard, a small Lacerta, is unique in genus Family Oppel,

Genus Coronella 1768 that it gives birth to living young, although some Laurenti, Coronellaaustriaca 1768 the South It is a Laurenti, populations in lay eggs. short-leg- rounded head that not Smooth Snake ged form with a is particu- larly distinct from the neck and has a rather dull brownish color normally with interrupted lines from The Coro- New species record the site. genus down the back andsides. This species is the most nella consists of rather small, smooth-scaled widespread lizard in the Palearctic, occurring snakes with a round body andan indistinct head. from northern Scandinavia, Ireland, and Britain and Both living species of the genus are secretive south to northern Spain, northern Italy, southern slow-moving. Both taxa are said to feed princi- Yugoslavia and Bulgaria, and then eastward differs pally on lizards. The small C. austriaca across much of Asia to the Pacific coast. the Coronella from the other snake in genus ( giron- of the dark that from the It is one of the most cold-tolerant reptiles dica) in having a stripe runs

and the Circle nostril to the This is in northern hemisphere enters Arctic eye. stripe usually lacking in localities in northern Europe. Unlike most C. girondica. Presently C. austriaca occurs in southern France, and northern Iberia, lizards of the genus Lacerta, the viviparous lizard England, requires a rather moist environmentand is one of east to southern Scandanavia and Russia and the few lizards that avidly eats earthworms. It south to Italy, Sicily, and Greece. It also occurs in

northern north to Iran. This snake is occupies a wide variety of habitats, ranging from Asia Minor heathlands to Alpine meadows in the south. This secretive, but it tendsto be active in the daylight habitats. viviparous lizard was previously tentatively hours, preferring dry and sunny Holman characters for the identifica- identified from Boxgrove as Lacerta cf. Lacerta (1998) gives This vivipara by Holman (1998). A partial right dentary tion of fossil vertebrae of Coronella austriaca. and from with four relatively slender teeth, including one is the first record of this genus species the Site. It is the second record tricuspid tooth, enables a positive identification Boxgrove only of (see Holman, 1998:85) of Lacerta viridis tobe made C. austriaca from the Pleistocene of Britain, the

from the Boxgrove Site. New material: partial first one being from the Early Middle Pleistocene right dentary with 4 teeth (188). of the Westbury-Sub-Mendip Cave Site inSomer- set. New Material: trunk vertebra (484). Family Anguidae Gray, 1825 Genus Anguis Linnaeus, 1758 Genus Natrix Laurenti, 1768 Anguis fragilis Linnaeus, 1758 Water Snakes

Slow Worm This snakes genus contains moderately stout

in with keeled scales. The is Anguis fragilis is the only modern species the strongly genus widely and a smooth-scaled, distributed in and southwestern genus. This taxon is very legless Europe also in Africa. lizard with a tail that is often longer than the central Asia, occurring In Europe,

of the maura and tesselata body, except when the tail has been broken off two species genus, N. N.

and regenerated. The color is usually brownish are aquatic, and the third, N. natrix, spends most

and males oftenhaveblue on the of the time on land. grey spots body. This all species occurs almost over Europe except cf. Natrix sp. for Ireland, southern Iberia, and the far north. It Probable Water Snake Species occurs in Britain andeast to the Urals and Cauca- New material: fragmentary vertebra (112) sus and parts of southwestern Asia as well as in

northwestern Africa. The slow worm prefers Natrix natrix (Linnaeus, 1758)

rather moist habitats with ample vegetation. It is Grass Snake

semifossorial and has slow and deliberate very The snake is a rather thick-bodied grass large, movements. Holman (1998) gives characters for snake with keeled scales. The color is variable, the identification of Pleistocene fossils of this but there is usually a yellow collar with a black species which has previously been reported from behind This border that occurs just the head. the Boxgrove Site (Gleed-Owen, 1999; Holman, all of species occurs in nearly Europe, ranging 1992, 1998). New material: five trunk vertebrae northto near theArctic Circle in Scandanavia and (32,44, 79,330 and 672), ten caudal vertebrae (53,

117 CRANIUM, 17, 2 - 2000

extending eastward to Lake Baikal. It also ocurs Natrix natrix (grass snake) in northwestern Africa. This taxon is diurnal and Natrix cf. Natrix natrix (probable grass snake) mainly terrestrial, although it is sometimes seen

in the water. swimming It prefers moist, grassy As far as can be determined, all of the Boxgrove habitats Holman over dry ones. (1998) gave This amphibian and reptiles are presently living. characters for the identification of fossil Natrix is in contrast to the temporally equivalent natrix vertebrae. This species has previously been Miesenheim 1 herpetofauna of the Neuwied reported from the Boxgrove Site (Gleed-Owen, Basin of Germany where Pliobatrachus cf. Plioba- 1999; Holman, 1992,1998). New material: partial trachus langhae Fejevary, 1917 has beenidentified. vertebra with complete hypapophysis (286). This permanently aquatic frog represents an Probable snake Natrix cf. Natrix natrix: new grass and and fact extinct family, genus, species, in material: partial vertebra (651). the extinct represents only anuran family ever described. Pliobatrachus has been identified from

other Middle Pleistocene localities in Central and Checklist of the Boxgrove herpeto- Eastern Europe (see Holman, 1998; Ratnikov, fauna 1997); thus one wonders, considering the large British Pleistocene herpetofaunas known from Following is an updated checklist of the amphibi- Cudmore Grove, and East Farm, Barn- that takes Boxgrove, ans and reptiles of the Boxgrove site ham, whether Pliobatrachus ever occurred in the into account the new herpetological records from in British Pleistocene, or for that matter, the the site reported in the present paper. Pleistocene of Western Europe. Newts Although no extinct amphibians and reptiles are Triturus species undeter- sp. (alpine newt, group known from the Boxgrove site, three extralimital mined) species are known that are presently absent from

Triturus cristatus (crested newt) - new record Britain, but occur at rather similar latitudes on the

Triturus helveticus or smooth or vulgaris (palmate continent(see Arnold & Burton, 1978). These taxa

newt) are: Pelobates fuscus (common spadefoot), Pelody-

Triturus helveticus (palmate newt) - new record tes punctatus (common parsley frog), and Rana Triturus vulgaris (smooth newt) arvalis (moor frog).

Anurans Paleoclimatically, there are no herpetological

that indicate that the climate in the Pelobatesfuscus (common spadefoot) species

area at the time of the of the Pelodytes punctatus (common parsley frog) - new Boxgrove deposition bones much different than it is record was at present.

The lack of such forms as Emys orbicularis (Euro- cf. Bufo sp. (probable true toad) and peanpond turtle) Elaphe longissima (Aescula- Bufo sp. (true toad)

species that are said to indicate Bufo bufo (common European toad) pian snake), climates than other British Bufo calamita (natterjack toad) warmer at present at and continental sites (e.g. Stuart, 1979; Holman, cf. Rana sp. (probable true frog) 1998; Bohme, 1996), are conspicuously absent at Rana sp. (true frog) the studied Site. Rana water - intensively Boxgrove (ridibunda) I sp. (European frog) new

record The Boxgrove amphibians and reptiles are yet to Rana arvalis (moor frog) be analyzed, if this is possible, on a microstrati- Rana cf. Rana arvalis (probable moore frog graphic basis. Nevertheless, as a whole, the Rana (common frog) temporaria herpetofauna generally indicates ponding and

moist habitats. All of the newt species Squamates adjacent prefer moist habitats and need at least temporary Lacerta vivipara (viviparous lizard) - new positive which breed. the water in to Among anurans, identification water frogs (Rana [ridibunda] sp.) prefer perma- Lacerta cf. Lacerta vivipara (probable viviparous nent aquatic situations. Parsley frogs (Pelodytes lizard) punctatus) prefer moist, well-vegetated areas. Anguis fragilis (slow worm) Both moor frogs (Rana arvalis) and common frogs Coronella austriaca (smooth snake) - new record (Rana temporaria) prefer marshy or damp grassy cf. Natrix water (probable grass or snake) areas. Common toads (Bufo bufo) are rather Natrix sp. (grass or water snake) ubiquitous in their habits, but always mustreturn

118 New herpetological records from the Middle Pleistocene Boxgrove Hominid Site, England

to water to breed. Both the spadefoot (Pelobates fuscus) Acknowledgements and both fuscus ) natterjack (Bufo calamita),, repre- I Mark Roberts and Simon am deeply grateful to sented by relatively few elements at Boxgrove, Parfitt for the opportunity to study the previous are more partial to sandy areas, but both need and new Boxgrove herpetofaunal material. water in which to breed. Teresa Peterson made the location of the map

Among the reptiles, both the viviparous lizard Boxgrove Site.

(Lacerta vivipara) and the slow worm (Anguis fragilis) prefer moist, at least moderately well-ve- The snake Natrix getated areas. grass ( natrix), as Adress of the author is in its name implies, usually most common J. Alan Holman moist grassy areas, usually fairly near an aquatic Michigan State University situation. The smooth snake (Coronella austriaca), East Lansing represented by only one vertebra from Boxgrove, Michigan prefers dry, sunny habitats. USA 48824-1045

119 CRANIUM, 17, 2 - 2000

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