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New evidence for complex climate change in MIS 11 from Hoxne, Suffolk,UK

Nick Ashtona,,Simon G Lewisb,Simon A Parfittc,d,Kirsty EH Penkmane,G Russell Coopef
aDepartment of Prehistory and Europe, British Museum, Franks House, 56Orsman Road, London N1 5QJ, UK
bDepartment of Geography, Queen Mary, University of London, Mile End Road,London E1 4NS, UK
cDepartment of Palaeontology, Natural History Museum, Cromwell Road,London SW7 5BD, UK
dInstitute of Archaeology, University College London, 31-34 Gordon Square,London WC1H 0PY, UK
eBioArCh, Departments of Biology, Archaeology and Chemistry, University ofYork, Heslington, York YO10 5DD, UK
fTigh-na-Cleirich, Foss, nr Pitlochry, Perthshire PH16 5NQ, UK

Corresponding author. Tel.: +44 2073238093; fax:+44 2073238101.nashton@thebritishmuseum.ac.uk

Received 2007 Jun 14; Revised 2008 Jan 2; Accepted 2008 Jan 4.

© 2008 Elsevier Ltd.

Open Access underCC BY 3.0 license

PMCID: PMC2748712  PMID:19777138

Abstract

The climatic signal of Marine Isotope Stage (MIS) 11 iswell-documented in marine and ice-sheet isotopic records and is known tocomprise at least two major warm episodes with an intervening cool phase.Terrestrial records of MIS 11, though of high resolution, are often fragmentaryand their chronology is poorly constrained. However, some notable exceptionsinclude sequences from the maar lakes in France and Tenaghi Philippon in Greece.In the UK, the Hoxnian Interglacial has been considered to correlate with MIS11. New investigations at Hoxne (Suffolk) provide an opportunity to re-evaluatethe terrestrial record of MIS 11. At Hoxne, the type Hoxnian Interglacialsediments are overlain by a post-Hoxnian cold-temperate sequence. Theinterglacial sediments and the later temperate phase are separated by theso-called ‘Arctic Bed’ from which cold-climate macroscopic plant andbeetle remains have been recovered. The later temperate phase was depositedduring an episode of boreal woodland and is associated with the artefacts, adiverse vertebrate fauna and molluscs. New amino acid geochronological data andbiostratigraphical considerations suggest that the post-Hoxnian sequencecorrelates with late substages of MIS 11. The paper further investigates thecorrelation of the sequence at Hoxne with the palynological sequences foundelsewhere in Europe and adjacent offshore areas.

1. Introduction

In recent years, the complexity and structure of Marine Isotope Stage (MIS)11 has been a focus of research, in part driven by the similarity of orbitallyforced insolation changes during MIS 11 and the Holocene (Oppo et al., 1998;Droxler et al., 2003;Loutre andBerger, 2003;Ruddiman,2005;Wu et al.,2007). MIS 11 is therefore important as an analogue forcurrent and future climate scenarios. An important aspect of this work is howglobal temperature changes affect terrestrial biota, which can be addressedthrough the correlation of marine and terrestrial records (Tzedakis et al., 1997;Desprat et al., 2005;Wu et al., 2007). It has become increasinglyclear that there is a much more complex relationship between the oftenfragmented terrestrial record and the marine and ice records. The complexity ofMIS 11 has now been shown through marine and ice-sheet isotope records (e.g.Bassinot et al., 1994;EPICA Community Members, 2004)and long palynological records from marine cores (Desprat et al., 2005). These all indicate a sharp warming atca 425 ka with what appears to be a relatively stable climatethrough to ca 390 ka. Thereafter, the records arecharacterised by a series of warm–cold oscillations until ca 360 ka with the onset of more extreme cold.

Two conventions have been established for the naming of isotopic andtherefore climatic fluctuations within numbered isotope stages. MIS 11 has beendivided into substages 11c, 11b and 11a (e.g.Tzedakis et al., 2001). However, in other records (e.g.MD900963,Bassinot et al., 1994) amore complex pattern can be seen with additional warm–cold oscillation(Fig. 1). Therefore, an alternative system identifies negative andpositive isotopic events, which are numbered using a decimal system(Imbrie et al., 1984;Bassinot et al., 1994;Desprat et al., 2005). This hasthe advantage of allowing additional isotopic events to be incorporated. Thesetwo conventions are different because the first denotes periods of time, whereasthe second identifies specific isotopic events, and therefore the terminologyshould not be used interchangeably. An additional short-lived warm episode hasbeen recognised in some records (e.g.Prokopenko etal., 2001) and referred to as 11e, but this is not prominentin either SPECMAP orMD900963. The structure and terminology for MIS 11 is usedin this paper are shown inFig.1.

Fig. 1.

Fig. 1

Structure and sub-division of MIS 11 as shown by isotopic recordsfrom ocean and ice cores. Isotope substages afterTzedakis et al. (2001), isotopic events afterBassinot et al. (1994). Sources: Insolation,Berger and Loutre (1991); EPICAdeuterium record,EPICA community members(2004); LR04 benthic stack,Lisiecki and Raymo (2005); SPECMAP stack,Imbrie et al. (1984);MD900963,Bassinot et al. (1994).

Only recently has it been possible to recognise these complex changes inthe terrestrial record, particularly in southern Europe where pollen sequenceshave enabled correlation with marine isotope substages (Reille and de Beaulieu, 1995;Tzedakis et al., 1997;de Beaulieu et al., 2001;Desprat et al., 2005;Tzedakis et al., 2006). In Britain, suchsequences have not been found, and most palynological records are of relativelyshort duration (cf.Thomas, 2001).Although there is now widespread agreement that the Anglian glaciationcorrelates with MIS 12 and the Hoxnian Interglacial broadly with MIS 11(Bridgland, 1994;Bowen, 1999;Rowe etal., 1999;Grün andSchwarcz, 2000;Preece andPenkman, 2005;Preece et al.,2007), individual substages of MIS 11 have not beenconvincingly identified (but seeSchreve(2001a, b)). The climatic record ofthe Hoxnian has been largely based on palynology (West, 1956;Turner,1970), but it is still not clear whether the Hoxnianencompasses the whole of MIS 11 or just one substage. One site that has thepotential to provide information of this kind is the site of Hoxne, Suffolk, UK(TM176767), where recent fieldwork has shown a more complex sequence of climatefluctuations that may be attributable to marine isotope substages.

The sequence at Hoxne forms the stratotype of the Hoxnian Interglacial(Mitchell et al., 1973) which,based on the analysis ofWest (1956)of the lacustrine sediments, spans pollen zones HoI-III. However, thestratigraphy also includes an important series of sediments that post-date thislacustrine sequence. These are of particular importance because they containabundant palaeoenvironmental evidence and also a rich Palaeolithicarchaeological record (Singer et al.,1993). Despite the long history of research at Hoxne, itremains unclear how the primary context human industries relate to theenvironmental record and the correlation of this part of the succession with themarine isotope sequence is uncertain (Bowen et al.,1989;Gladfelter et al.,1993;Turner and West1994;Ashton et al.,1995;West and Gibbard,1995; Grün and Schwarcz 2000;Schreve 2001a, b). Thecurrent research has focussed on these issues as part of a wider investigationof human presence during MIS 11 under the auspices of the Ancient HumanOccupation of Britain Project (Stringer and AHOBProject Members, 2003;Ashton etal., 2006).

Since the discovery at Hoxne of Lower Palaeolithic handaxes byFrere (1800), the site has beenthe subject of several investigations, providing often radicalre-interpretations of previous work. This work has focussed on two pits, eitherside of the Hoxne to Eye road (Figs. 2 and3). The Old Brick Pit to the east of the road was the subjectof the earliest investigations, and these were supplemented by work in theOakley Park Pit when this was first opened in the mid-19th century. The two pitswere opened for gravel and clay extraction from the infillings of a basin thatis now located on the interfluve between the Goldbrook stream and River Dove,which flow into the River Waveney. The current research has involvedinvestigation of archive material together with cutting and sampling of keysections (between 2000 and 2003;Ashton et al.,2003) and has enabled a re-evaluation of thepalaeoenvironmental and archaeological evidence from the site.

Fig. 2.

Fig. 2

Location of sites mentioned in the text.

Fig. 3.

Fig. 3

Site location and plan. The basin contours (mOD) and limits of thelake are based onWest(1956).

2. Previous interpretations

The basis for much of our understanding of the site comes from the work ofClement Reid (Evans et al., 1896). Heprovided detailed and clear descriptions from boreholes and open sections of allthe major sediment units and undertook the first palaeobotanical investigationsat the site. Reid's work demonstrated that a basin, formed in the surfaceof the ‘boulder clay’ (till), was infilled with interglaciallacustrine sediment (Bed E). Drying out of the lake was indicated by theformation of peat (Bed D), prior to the re-establishment of the lake under coldconditions (‘Arctic Bed’: Bed C). These lacustrine sediments wereoverlain by a fluvial, shelly, gravel (Bed B), and the artefact-bearing sedimentor ‘Palaeolithic Loam’ (Bed A). The latter extended beyond theconfines of the basin (Figs. 4 and5).

Fig. 4.

Fig. 4

Stratigraphical interpretation of (a) Reid (Evans et al., 1896) in comparison to that of(b)West (1956). Modified fromWest and McBurney (1954, Fig. 2)and fromWest (1956, Fig.15).

Fig. 5.

Fig. 5

The interpretations of the stratigraphy of Reid (Evans et al., 1896),West (1956),Singeret al. (1993) and current work. The handaxe symbols show thecontexts in which artefacts were thought to be located.

In the 1950s, detailed fieldwork was undertaken byWest (1956), who also examined the palynologyof the lake sediments. West argued for important modifications to thestratigraphy offered by Reid. Other than the addition of a cold lacustrinesediment (Stratum F) above the till (now Stratum G), he argued that the lowerpart of Reid's Bed A had been misinterpreted and was in fact decalcifiedBed (now Stratum) E. This implied that the human occupation of the site wasassociated with the lacustrine sediments of the newly defined HoxnianInterglacial, rather than in the later sediments at the site (Figs. 4 and 5). No archaeology wasdiscovered in Stratum E during West's work, other than two flakes fromsections 40 and 100 on the west of the Oakley Park Pit (West, 1956;Fig.3). However, it is clear from re-examination of the sectiondrawings and the heights of the objects in comparison to those found in the1970s and during the recent excavation that they were actually recovered fromsediments overlying Stratum D.West(1956) defined pollen zones at Hoxne, which were subsequentlymodified following work on the more complete sequence at Marks Tey, Essex. TheHoxne lake sediments (Strata E–D) were assigned to pollen zones HoIthrough to HoIIIa, with HoIIIb and HoIV being absent from the sequence(Turner, 1970).

Major archaeological excavations by the University of Chicago in the 1970s(Singer et al., 1993) in theOakley Park Pit and in the field to the west of this pit (Fig. 3), provided the first properly excavatedartefact assemblages from the site and led to further re-interpretation. It wasargued that there were two significant phases of human occupation. The first or‘Lower Industry’ consisted of primary context handaxes, cores andknapping debitage in association with a temperate faunal assemblage, andoccurred in a single horizon towards the base of Stratum C. A new bednomenclature was introduced for a sequence of fluvial, alluvial and solifluctionsediments (Beds 1–9) that lie above Stratum D (Gladfelter, 1993). These had formerly been attributed toStrata B and A. The key units were a chalky gravel (Bed 4), which was overlainby a fine-grained alluvial sediment (Bed 5) and a further gravel (Bed 6). The‘Upper Industry’, consisting of pointed handaxes, flake tools, coresand debitage in variable condition, was recovered from the top of Bed 5 and insecondary context in the lower part of Bed 6. The ‘Lower Industry’was associated with a temperate vertebrate fauna. However, this conflicts withthe palaeobotanical evidence of Reid (inEvans etal., 1896) andWest(1956) who attributed Stratum C (‘Arctic Bed’) toa cold stage.

The dating of the site and its correlation with the marine isotope recordwas also unclear from this work (Gladfelter et al.,1993;Turner and West,1994). Amino acid D/L ratios onValvata shells from Stratum E suggested a correlationof the lacustrine sequence with MIS 9 (Bowen et al.,1989), with the implication that either there is a majorhiatus between the Lowestoft Till (MIS 12) and the lake beds, or that theglacial sequence dates to MIS 10. Dating of the overlying Lower Industry(interpreted as lying in Stratum C) was assessed through thermoluminescencedating on two burnt flints, which yielded a mean date of 210±20 ka, suggesting an MIS 7 age. However, the dosimeter readings(taken several years after the excavation) were from different locations andalso demonstrated considerable variation across the site. This suggested thatthere were likely to be considerable errors in the TL age estimates(Bowman, 1993). Initial ESR dateson enamel from two horse (Equus ferus) teeth alsoassociated with the Lower Industry gave an average age of 319±38 ka, suggesting that Stratum C was attributable to MIS 9(Schwarcz and Grün, 1993).However, subsequent remodelling of the data has suggested an MIS 11 age withdates of 404±33/42 and 437±38 ka (Grün and Schwarcz, 2000). Finally,assessment of the mammalian fauna that is associated with the Lower Industry andfrom Bed 4 has been argued to show marked similarities with that from Swanscombe(Stuart et al., 1993), and bothfaunal assemblages have been attributed to the first post-Anglian warm stage andassigned to the Swanscombe Mammal Assemblage Zone (MAZ) bySchreve (2001a, b).

3. Stratigraphic re-interpretation

A revised stratigraphy is proposed, based on the curent re-investigation ofthe site that focussed on the relationship between Stratum C and the sedimentscontaining the Lower and Upper Industries (Fig.5;Table1). This has resolved theconfusion inSinger et al. (1993) intheir varied interpretations of the relationship between Stratum C and Bed 4(seeBridgland, 1994;Turner and West, 1994). The stratigraphicinterpretations and nomenclature ofSinger et al.(1993) can now be reconciled with that ofWest (1956). This paper uses the Reid/Westnomenclature, with some modifications based on the current research(Table 1).

Table 1.

New bed names and descriptions with interpretation of climate andcontext of archaeology

Bed nameBed descriptionPollen zoneClimatic interpretationArchaeology
Stratum A1CoversandCold
Stratum A2(i)Cryoturbated sand and gravelCold
Stratum A2(ii)Solifluction gravelColdDerived Upper Industry
Stratum A2(iii)Alluvial sandy clayWarmUpper Industry
Stratum B1Fluvial sand, silt and clayWarmLower Industry
Stratum B2Fluvial chalky gravelWarm
Stratum CLacustrine sands and siltsCold
Hiatus
Stratum DPeatHoIIIaWarm
Stratum ELacustrine clayHoI–IIcWarm
Stratum FLacustrine claylAnCool
Stratum GTillCold

Fig. 3 shows the location of therecently excavated sections (Areas I–VII) and boreholes. The mostsignificant information comes from Areas III and IV. The latter was a narrowtrench, previously excavated in 1978 that was re-opened and widened to allowdetailed investigation and sampling. The trench linked the locations of theLower and the Upper Industries in the field to the west of the Oakley Park Pit(Figs. 3 and 6). Thissection is critical to understanding the relationship of the Upper and LowerIndustries to Stratum C and Beds 4–6.

Fig. 6.

Fig. 6

Schematic cross-section through the Hoxne lake basin (afterWest, 1956) with a detailedsection through Area IV.

Stratum E was exposed at the base, overlain by a near-continuous horizon ofStratum D, which reached a maximum thickness of 35 cm at thesouthern end, but thinned to <1 cm black stained clay atthe northern end (Fig.6). Overlying this was a thinhorizon of Stratum C, 50-cm thick over most of the exposure, but cut out at thesouthern end by fine sand and chalky gravel of Stratum B.

Above this, a concave-up erosional lower bounding surface is incised intoStrata C and B, forming a broad (>30 m), shallow (ca 2 m) channel feature, infilled with bedded sands, silts and clays.These are interpreted as a lateral accretion facies and indicate lateralmovement to the north. The orientation of bone long axes and their distributionwith that of the artefacts also suggest a fluvial deposit and indicate aNE–SW orientation of the channel. This channel feature was not recognisedby previous workers and the sediment was thought to be part of the lacustrinesequence. It is here assigned to Stratum B because it is a fluvial deposit andis referred to as Stratum B1. The underlying chalky gravel is therefore assignedto Stratum B2 (Fig. 5).

Stratum A was sub-divided into A1 and A2 byWest (1956). As a result of the current work, Stratum A2 isfurther sub-divided into A2(i–iii). Stratum A2(iii), a sandy clay unit, isinterpreted as alluvium. Above this, a coarse flint gravel with a sandy claymatrix (Stratum A2(ii)) and a series of laminated sands and silts (A2(i)) arecapped by gravely sands (Stratum A1). Post-depositional disturbance anddownslope movement have affected the uppermost part of the succession.

4. Palaeoenvironmental context for human occupation atHoxne

The sedimentary succession at the site contains palaeoenvironmental dataindicating a fluctuating climate. The depositional environment, vegetational andfaunal character, and thermal conditions can be considered for each stratum inturn.

4.1. Stratum G

The ‘boulder clay’ of Reid was assigned to the LowestoftTill byWest (1956) andrepresents widespread glaciation of eastern England by a British-basedice-sheet depositing the characteristic chalk and flint-rich tills of theLowestoft Formation (Perrin et al.,1979;Bowen,1999;Clark et al.,2004). This glaciogenic unit is attributable to theAnglian Stage (MIS 12).

4.2. Stratum F

This lacustrine clay lies at the bottom of the basin and containspollen and beetles (West, 1956;Coope, 1993). The beetleremains indicate a rapid amelioration of climate to near interglacialconditions during the late Anglian.

4.3. Stratum E

These lake sediments form the major filling of the basin and contain apollen sequence that has been attributed to pollen zones HoI—HoIIc ofthe Hoxnian Interglacial. The pollen indicates development of fullytemperate deciduous woodland (West,1956;Turner,1970). The prominent non-arboreal pollen phase at the topof Stratum E is characteristic of a number of sites in the region(Turner, 1970;Thomas, 2001, 2002). Its origin isunclear, though it is not thought to show a cooling in climate(Turner, 1970).

4.4. Stratum D

This peat horizon indicates drying out of the lake basin andencroachment of terrestrial vegetation over the lake bed. The arborealpollen contains significant quantities of alder, suggesting an alder carrenvironment developed during pollen zone HoIIIa. (West, 1956). Beetles indicate mean Julytemperatures of between 15 and 19 °C (Coope, 1993).

4.5. Stratum C

A return to lacustrine deposition is shown by the laminated sedimentsof Stratum C, which record fluctuating flows, with influx of coarser sandsand silts, together with pellets of reworked lacustrine sediments andorganic material. These were well exposed in Area VII. This stratum wasoriginally assigned to pollen zone HoIIIb on the basis of the high counts ofAbies, which is characteristic of this pollenzone (Turner, 1970). Theoccurrence ofAbies and other thermophilous plantsis, however, at odds with the presence of leaves of Arctic/Alpine plants,notably dwarf birch (Betula nana) and three speciesof dwarf willow (Salix myrsinites,S.herbacea andS. polaris)(Evans et al., 1896). Leavesof these species were recovered during the current work and are almostcertainly contemporary with the unit, as they are fragile and would notsurvive reworking. This suggests that some of the pollen (includingAbies) has been reworked into this unit(West, 1995) and furthermoreindicates a hiatus between the deposition of Strata D and C.

The interpretation of a cold climate is also supported by the analysisof the beetles. Altogether, 72 coleoptran taxa have been recognised of which42 can be named to species. Of these, 10 do not now live in the BritishIsles. There is little change through the sequence, so the species have beengrouped together as a single assemblage inTable2. The local environmentsuggests a pool of standing water with much marginal emergent vegetationsuch as the aquatic grassGlyceria and a surfacewhich was at least in part covered withLemna. Theimmediate surroundings of the pool were dominated by sedges and other reedyplants. The low numbers of dung beetles suggests that there were few largeherbivorous mammals present at this time.

Table 2.

Coleoptera from Stratum C, Hoxne

Carabidae
 Notiophilus cf.aquaticus (L.)2
 Dyschirius globosus(Hbst.)3
 Trechus secalis(Payk.)1
 Bembidion hastiSahlb.a1
 Bembidion cf.mckinleyi Fall.a1
 Bembidion guttula(F.)/unicolor Chaud.2
 Bembidion sp.4
 Patrobus cfatrorufus (Ström)2
 Pterostichus nigrita(Payk.)1
 Amara sp.2


Dytiscidae
 Potamonectes griseostriatus(de Geer)3
 Agabus bipustulatus(L.)1
 Ilybius sp.2
 Rhantus sp.1
 Colymbetes dolabratus(Payk.)a2
 Colymbetes sp.5
 Graphoderus sp.1
 Dytiscus sp.1


Gyrinidae
 Gyrinus aeratusSteph.1
 Gyrinus sp.2


Hydraenidae
 Hydraena sp.5
 Ochthebius minimus(F.)7
 Helophorus obscurellusPopp.a6
 Helophorus cf.aquaticus (L.)1
 Helophorus smallspp.5


Hydrophilidae
 Cercyon convexiusculusSteph.4
 Enochrus sp.2
 Hydrobius fuscipes(L.)7


Orthoperidae
 Orthoperus sp.2


Ptiliidae
 Ptenidium sp.2
 Acrotrichis sp.2


Staphylinidae
 Pycnoglypta lurida(Gyll.)a1
 Olophrum fuscum(Grav.)7
 Olophrum boreale(Payk.)2
 Eucnecosum brachypterum(Grav.)24
 Geodromicus kunzeiHeer1
 Boreaphilus henningianusSahlb.a5
 Holoboreaphilusnordenskioeldi (Makl.)a6
 Trogophloeus sp.4
 Oxytelus rugosus(F.)1
 Bledius sp.1
 Stenus spp.7
 Euaesthetus laeviusculusMannh.1
 Lathrobium sp.1
 Tachyporus sp.1
 Tachinus rufipes (deGeer)1
 Tachinus cf.corticinus Grav.1
 Alaeocharinae gen. et sp. indet.59


Elateridae
 Agriotes sp.1


Throscidae
 Throscus sp.1


Helodidae
 gen. et sp. indet.7


Dryopidae
 Dryops sp.3


Byrrhidae
 Simplocaria metallica(Sturm.)a6


Coccinellidae
 Hippodamia arcticaa1


Tenebrionidae
 Eledona agaricola(Hbst.)1


Scarabaeidae
 Aphodius spp.2


Chrysomelidae
 Donacia dentata Hoppe2
 Donacia semicupreaPanz.5
 Donacia aquatica(L.)1
 Donacia thalassinaGerm.9
 Donacia cinereaHbst.2
 Plateumaris affinis(Kunze)4
 Chrysomela sp.1


Curculionidae
 Apion sp.1
 Sitona sp.1
 Stenoscelis(=Brachytemnus)submuricatus (Schönh.)a3
 Bagous sp.2
 Tanysphyrus lemnae(Payk.)9
 Notaris bimaculatus(F.)4
 Notaris acridulus(L.)1
 Notaris aethiops(F.)2
 Thryogenes sp.1
a

Indicates species not now native to the British Isles.

Taken as a whole, the coleopteran assemblage indicates very cold andcontinental climatic conditions with a number of species now found livingtoday no nearer than arctic Russia (e.g. the closest locality forHelophorus obscurellus is on the Kanin Peninsula,the closest locality forHoloboreaphilusnordenskioeldi is central Novaya Zemlya). However, there arethree species whose presence in this assemblage seems to be climaticallyanomalous. First,Stenoscelis submuricatus is aMediterranean beetle that lives in dead wood. It was very common in StratumD. This species could have been derived from Stratum D, having beenincorporated into Stratum C (sealed from agents of decomposition insidereworked pieces of wood). On a less extreme scale,Eledonaagricola lives in various fungi growing on deciduous trees,chieflyPolyporus sulphuraeus growing onSalix. In northern Europe, this beetle reachesonly as far north as latitude 60°N. Species ofThroscus inhabit leaf litter but again theirgeographical ranges only reach as far north as latitude 62°N. Both theserecords are based on single fragments and it is likely that they were alsoderived from the eroded deposits of the lacustrine sequence of Stratum Dimmediately beneath. Other than the probable derived elements, the insectsindicate mean temperatures in July of, or below 10 °Cand in January and February of about −15 °C.

4.6. Strata B2 and B1

The chalky sandy gravel of Stratum B2 is a fluvial sediment andcontains a rich vertebrate fauna. The sands, silts and clays of Stratum B1are also fluvial and rest in a channel cut into Stratum B2. The ‘LowerIndustry’ in association with the vertebrate fauna was recovered alongthe northwest margins of this channel.

The vertebrate faunal assemblages from Strata B2 and B1 are verysimilar in composition (Stuart et al.,1993). The larger mammalian fauna is dominated byEguus. ferus (horse),Cervuselaphus (red deer),Dama dama (fallowdeer), together with occasional remains ofMacacasylvanus (macaque),Ursus sp. (bear),Lutra lutra (otter),Pantheraleo (lion),Stephanorhinus sp.(extinct rhinoceros) andCapreolus capreolus (roedeer). Insectivores and rodents dominate the smaller mammals, which includeCastor fiber (European beaver),Trogontherium cuvieri (extinct giant beaver),Talpa minor (extinct mole),Microtus(Terricola) cf.subterraneus (pinevole) and lemming (identified asLemmus lemmus(Norway lemming) byStuart et al.,1993). Remains of birds, amphibians, reptiles and fishwere also recovered, including the articulated skeleton of a rudd(Scardinius erythrophthalmus) (Brian Irving,personal communication), the latter suggesting water temperatures wererelatively warm during the summer months (cf.Stuart, 1982).

The range of species suggests a mix of environments. The dominance ofhorse indicates areas of open landscape, whereas forest is indicated byfallow deer, beaver and macaque. Although lemmings are only found in cold,northern latitudes today, they may have had a different distribution andhabitat requirements in the Middle Pleistocene. For example, they occur atBoxgrove, West Sussex (Parfitt,1999), during the latter part of an interglacial wherethey are associated with mammals and Mollusca typical of temperate deciduouswoodland (e.g.Myotis bechsteinii (Bechstein'sbat),Muscardinus avellanarius (common dormouse),D. dama (fallow deer),Acanthinulaaculeata,Spermodea lamellata andAegopinella pura).

4.7. Strata A2 and A1

The alluvial silt of Stratum A2(iii) contains a sparse fauna includingan indeterminate species of elephant, extinct rhinoceros, horse, red deer,roe deer and fallow deer, again all suggesting a temperate climate. There isa possibility, however, that this fauna is derived from the lower units. Theoverlying sands and gravels of A2(ii), A2(i) and A1 contain no biologicalremains other than mixed pollen (Mullenders,1993;Turner and West,1994). Stratum A2(i) displays contemporaneous ice-wedgecasts, indicating a permafrost environment (Singer et al., 1993). Periglacial structures in StratumA1 also suggest a return to a cold climate.

4.8. Palaeoclimatic summary

The complete succession at Hoxne indicates a complex pattern ofclimatic fluctuations and changes in depositional regime. Followingdeglaciation, the lake basin began to infill under cool conditions (StratumF) followed by rapid amelioration to full interglacial conditions whichpersisted throughout the accumulation of Stratum E. After a phase ofnon-lacustrine conditions when peat formed across the former lake basin(Stratum D), there is a hiatus and then a return to lacustrine conditions isindicated by Stratum C. By this time, climate had deteriorated with plantmacrofossils and beetles indicating deposition under much colder conditions.In Area VII of the recent excavations, the top of Stratum C interdigitateswith and is overlain by sand and fine, chalky gravel indicating increasedflow into the basin and the establishment of a fluvial environment acrossthe site (Stratum B2). This is incised by a further fluvial channel, whichis infilled with fine-grained sediments (Stratum B1). The faunal elementswithin Stratum B suggest climatic amelioration, though probably not to thesame extent as indicated by Strata E and D. Temperate climate also prevailedduring deposition of Stratum A2(iii). The remainder of Strata A2 and A1accumulated under cold climate conditions.

The archaeological assemblages of the Lower and Upper Industries andtheir associated mammalian assemblages can now be placed within thisstratigraphic and environmental framework. No archaeological material can besecurely attributed to Strata F–D. The Lower Industry is associatedwith the base of the channel-fill represented by Stratum B1 (Fig. 6). The Upper Industry was recoveredfrom the upper part of Stratum A2(iii) and in a secondary context withinoverlying gravel, Stratum A2(ii). The critical consideration here is thatboth the Lower and Upper Industries can now be shown to post-date the‘Arctic Bed’ of Stratum C.

The cold event represented by Stratum C and the temperate eventrepresented by Stratum B have so far not been successfully dated orcorrelated with other terrestrial sequences or with the marine isotoperecord. Given the climatic complexity of MIS 11 (e.g.Bassinot et al., 1994;Tzedakis et al., 1997;Petit et al., 1999;Desprat et al., 2005) they could becorrelated with later cold and warm events in MIS 11, or alternatively witheven younger cold and warm episodes.

5. Amino acid geochronology

Amino acid racemization (AAR) analyses were undertaken on 12Bithynia tentaculata opercula using the methodsoutlined inPenkman (2005) andPenkman et al. (2008). The methodis based on the extent of protein decomposition, which increases with time,although there is an increased rate of breakdown during warm stages and aslowing in cold stages.

The samples were from Stratum E (NEaar 0498–0500, 2446–2447)and Stratum B2 (NEaar 3143–3150). The results show levels of proteindecomposition higher than those from sites correlated with MIS 9, but lower thanthose from sites of pre-Anglian age (Fig.7;Table 3). Furthermore, thelevels of protein decomposition are similar to those from sites correlated withMIS 11, including Elveden (Ashton et al.,2005), Beeches Pit (Preece etal., 2007), Barnham (Preece andPenkman, 2005), Clacton (Penkmanet al., in press), Woodston and Swanscombe (Penkman, 2005). This indicates an age for Hoxnebetween the Anglian (MIS 12) and early MIS 9. The opercula samples from StratumE tend to have slightly greater protein decomposition than those from Stratum B2and less degraded protein than found in opercula from the Lower Freshwater Bedat Clacton, which was deposited early in MIS 11 (Bridgland et al., 1999).

Fig. 7.

Fig. 7

D/L values of Asx, Glx, Ala, Val and [Ser]/[Ala] for the (A) Free(FAA;F) and (B) Total Hydrolysable amino acid (THAA;H) fractions of bleachedBithynia tentaculata opercula from Hoxne (Strata Eand B2), compared with shells from sites correlated with MIS 9 (Cudmore Grove,Grays, Hackney, Purfleet) and sites correlated with MIS 11 (Elveden, EbbsfleetSouthfleet Road, Swanscombe, Woodston, Clacton, Beeches Pit). For each group,the base of the box indicates the 25th percentile. Within the box, the solidline plots the median and the dashed line shows the mean. The top of the boxindicates the 75th percentile. Where more than nine data points are available,the 10th and 90th percentiles can be calculated (shown by lines at the bottomand the top of the boxes, respectively). The results of each duplicate analysisare included in order to provide a statistically significant sample size. They-axes for the [Ser]/[Ala] data are plotted in reverse, so that the direction ofincreased protein degradation for each of the indicators remains the same. Note:different scales on they-axes.

Table 3.

Amino acid data on opercula ofBithyniatentaculata from Strata E and B2 at Hoxne

NEaar no.Sample nameAsx D/LGlx D/LSer D/LAla D/LVal D/L[Ser]/[Ala]
0498bFHoBto1bF0.769±0.0000.366±0.0010.745±0.0000.480±0.0030.261±0.0000.359±0.001
0498bH*HoBto1bH*0.686±0.0010.293±0.0000.743±0.0020.424±0.0020.236±0.0020.308±0.001
0500bFHoBto2bF0.782±0.0010.369±0.0290.990±0.0020.531±0.0010.285±0.0080.283±0.004
0500bH*HoBto2bH*0.692±0.0020.296±0.0010.750±0.0040.444±0.0030.235±0.0020.281±0.003
2446bFHoBto3bF0.777±0.0020.374±0.0001.051±0.0060.483±0.0030.271±0.0030.302±0.002
2446bH*HoBto3bH*0.689±0.0020.283±0.0010.754±0.0060.413±0.0050.216±0.0020.287±0.000
2447bFHoBto4bF0.763±0.0140.395±0.0021.044±0.0050.485±0.0030.285±0.0030.296±0.004
2447bH*HoBto4bH*0.689±0.0000.290±0.0000.748±0.0130.412±0.0010.223±0.0140.282±0.008
3143bFHo64Bto1bF0.745±0.0060.332±0.0031.028±0.0040.434±0.0080.256±0.0020.310±0.006
3143bH*Ho64Bto1bH*0.689±0.0010.305±0.0000.805±0.0130.390±0.0000.220±0.0060.310±0.002
3144bFHo64Bto2bF0.752±0.0040.346±0.0041.030±0.0070.457±0.0020.287±0.0010.300±0.003
3144bH*Ho64Bto2bH*0.690±0.0020.311±0.0000.807±0.0010.395±0.0010.223±0.0030.315±0.001
3145bFHo64Bto3bF0.775±0.0050.334±0.0061.037±0.0010.498±0.0070.286±0.0030.286±0.002
3145bH*Ho64Bto3bH*0.696±0.0020.311±0.0000.783±0.0090.435±0.0000.236±0.0020.289±0.001
3146bFHo64Bto4bF0.749±0.0010.300±0.0051.013±0.0030.440±0.0010.269±0.0000.306±0.000
3146bH*Ho64Bto4bH*0.680±0.0020.279±0.0000.764±0.0010.383±0.0030.210±0.0020.300±0.000
3147bFHo50Bto1bF0.748±0.0050.342±0.0021.028±0.0110.439±0.0150.264±0.0030.298±0.008
3147bH*Ho50Bto1bH*0.709±0.0010.332±0.0010.833±0.0000.415±0.0000.240±0.0010.293±0.009
3148bFHo50Bto2bF0.767±0.0020.337±0.0051.012±0.0320.503±0.0010.303±0.0010.289±0.003
3148bH*Ho50Bto2bH*0.703±0.0030.321±0.0010.784±0.0000.450±0.0030.257±0.0020.286±0.000
3149bFHo50Bto3bF0.759±0.0060.327±0.0071.026±0.0000.482±0.0020.275±0.0010.297±0.001
3149bH*Ho50Bto3bH*0.679±0.0030.306±0.0030.754±0.0110.419±0.0040.223±0.0030.298±0.001
3150bFHo50Bto4bF0.771±0.0000.321±0.0020.771±0.0010.496±0.0000.294±0.0010.310±0.001
3150bH*Ho50Bto4bH*0.701±0.0030.312±0.0060.686±0.0050.437±0.0050.242±0.0030.286±0.002

Error terms represent 1 S.D. about the mean forthe duplicate analyses for an individual sample. Each sample was bleached (b),with the free amino acid fraction signified by ‘F’ and the totalhydrolysable fraction by ‘H*’. NEaar 0498-0500 and 2446-2447 arefrom Stratum E, and NEaar 3143-3150 are from Stratum B2.

The opercula from Stratum B2 have some of the lowest levels of proteindecomposition determined from MIS 11 sites. This therefore suggests an age forthe opercula between mid-MIS 11 and early MIS 9. While the values obtained fromthe opercula from Stratum B2 generally show higher levels of proteindecomposition than those obtained from the MIS 9 sites analysed, the separationbetween the Stratum B2 samples and those deposited early within MIS 9 is small.As so little decomposition occurs in the cold stages, and because of the extentof natural variability in biological samples, it can be difficult todiscriminate the end of one warm stage from the beginning of the next. Althoughan age late in MIS 11 is more likely, it is not possible to rule out an earlyMIS 9 age, given the level of resolution currently obtainable from thetechnique.

6. Biostratigraphy

The mammalian fauna from Strata B1 and B2 also provides an indication ofage. Three species are of possible biostratigraphic significance. The mostimportant of these isMicrotus (Terricola) cf.subterraneus. Although it is widespread in Europetoday, it appears to have been absent in Britain after MIS 11 (Parfitt, 1998). Of particular significance isits absence from the very rich faunal assemblages from Cudmore Grove, Grays andPurfleet, all of which have been attributed to MIS 9 and from any younger sites(Bridgland, 1994;Schreve et al., 2002).

Of lesser significance is the presence ofTrongontheriumcuvieri andTalpa minor. Although theyare thought to have become extinct after the Hoxnian in Britain, and possiblythe Holsteinian in Europe, their remains are so rare that any apparent absencein sites attributed to MIS 9 or later might be due to insufficientsampling.

7. Discussion

Both the amino acid geochronology and the biostratigraphy, together withthe reassessment of the ESR dates (Grün andSchwarcz, 2000; see above), suggest that Strata B1 and B2 aremost likely to be attributable to MIS 11. This therefore implies that theunderlying Strata E and D (Hoxnian) date to the first prolonged temperatesubstage in MIS 11, and that Strata C and B are later cold and warm substages,respectively, within MIS 11.

This correlation of the sequence at Hoxne with substages of MIS 11 haswider implications for its correlation with other terrestrial sites in the UKand further afield. Although the full Hoxnian Interglacial sequence is not foundat Hoxne, a complete succession is found at Marks Tey (Fig. 2), where pollen zones HoI-IV arerepresented (Turner, 1970).Furthermore, the palynology suggests that there is no evidence for a hiatusbetween the Anglian till and the lacustrine sediments at either Hoxne or MarksTey. The Hoxnian record at Marks Tey is an overlapping composite sequence fromtwo main cores. The interpretation of these cores is of a continuous temperatesequence through the Hoxnian without any indication of a cold event. Togetherwith the evidence from Hoxne of a later MIS 11 cold substage, this suggests thatthe Hoxnian Interglacial can be equated with the first major temperate substagewithin MIS 11.

Palynology has also been used to correlate the lacustrine deposits at Hoxneand Marks Tey with the organic channel-fills at Clacton and Tillingham, whichform part of the Thames/Medway sequence (Fig.2). At Clacton, the Freshwater Beds and Estuarine Bed(Pike and Godwin, 1953) have beencorrelated with HoIIb–HoIIIb (Kerney,1971;Bridgland et al.,1999), while at Tillingham, the silty sands and organic siltsare attributed to HoIII (Roe, 2001)(Fig. 8). On the basis of their lithology, terrace stratigraphy andmolluscan assemblages (Bridgland,1994;Roe, 2001;Preece et al., 2007), both thesesites are argued to be part of the same terrace aggradation as the Lower Gravel,Lower Loam and Middle Gravels at Swanscombe (Fig.2). All three sites record the immigration of the‘Rhenish’ fauna, probably in late HoII (indicating a confluence ofthe Thames with the Rhine). Furthermore, the presence of estuarine molluscsindicates a high sea-level stand, argued from the evidence at Clacton andTillingham to occur during HoIIIb (Fig.8).

Fig. 8.

Fig. 8

Suggested correlation between Hoxne, Marks Tey, Clacton, Tillinghamand Swanscombe with pollen zones. For Clacton, UFB=Upper Freshwater Bed andEB=Estuarine Bed. For Swanscombe, LMG=Lower Middle Gravel and UMG=Upper MiddleGravel.

These correlations therefore suggest that the sequences at Clacton,Tillingham and the Lower Gravel to Middle Gravels at Swanscombe can also beattributed to the first temperate substage of MIS 11. This is at variance withthe interpretation of Swanscombe proposed bySchreve (2001a, b) who attributed theMiddle Gravels to a later temperate substage within MIS 11.

The interglacial sequence at Quinton in the West Midlands has beeninterpreted as spanning the entire Hoxnian Interglacial on the basis of itspalynology (Horton, 1989). However,evidence from the Coleoptera indicates a more complex climatic picture with a‘cold interlude’ occurring during the latter part of theinterglacial (Coope and Kenward,2007). The beetle fauna from this ‘cold interlude’ isvery similar to that of the ‘Arctic Bed’ at Hoxne and suggests apossible correlation. Alternatively, since the uppermost samples of the Quintonsequence, attributed to the onset of the succeeding glacial, also yielded asimilar suite of cold-adapted beetle species, correlation of the Hoxne ArcticBed with these uppermost samples at Quinton is a possibility. The beetleevidence for the ‘cold interlude’ at Quinton is at odds with thepalynological evidence which “does not appear to show any response to thiscold episode” (p. 3284). A similar discrepancy in the evidence fromStratum C at Hoxne has been accounted for by reworking of temperate pollen intoStratum C (see above andTurner,1970). A comparable situation may have occurred at Quinton wherepollen of temperate character is reworked from the underlying deposits and foundin conjunction with an autochthonous coleopteran assemblage indicative of coolerclimatic conditions. A re-evaluation of the palynology of the Quinton successionmay help to resolve these problems associated with correlation of the Hoxne andQuinton sequences.

In Europe, significant advances have been made over the last decade inrelating the vegetational record from long, continuous sequences from sites insouthern Europe to the marine isotope record. Key to this success has been coreMD01–2447, near the northwest coast of the Iberian Peninsular(Fig. 2), where the marine isotoperecord can be directly compared to pollen that reflects vegetational changesinland (Desprat et al., 2005). Thiscore is argued to span the last 426 ka and has been comparedto other continuous or composite palynological sequences from Tenaghi Philipponin Greece, and Velay maar sites (Praclaux, Le Bouchet and Ribains;Fig. 2) in France (Reille and de Beaulieu, 1995;Tzedakis et al., 1997, 2001,2006). All these sequences show a similar pattern of vegetationand climate change with successive interglacial/glacial cycles. These can berelated to records of global climate change from deep-sea cores (Oppo et al., 1998;McManus et al., 1999), ice cores (Petit et al., 1999;EPICA Community Members, 2004) and the changes in thebiogenic silica content in the sequence from Lake Baikal (Prokopenko et al., 2001) (Fig. 9). Absolutedates from the terrestrial sites support these correlations with40Ar/39Ar dates on trachytictephra in deposits of the third interglacial at Velay (Le Bouchet Interglacial)suggesting an MIS 7 age (de Beaulieu et al.,2001), and palaeomagnetic analyses and U-series dates on thesequence at Tenaghi Philippon providing further tie-points to the marine isotoperecord (Tzedakis et al., 1997,2006).

Fig. 9.

Fig. 9

Correlation of MIS 11 sites across Eurasia with the Antarctic icecore based on data derived fromEPICA Communitymembers (2004),Desprat et al.(2005),Reille and de Beaulieu(1995),West(1956),Turner(1970),Nitychoruk et al.(2005) andProkopenko et al.(2001). The arboreal pollen (AP) curves for Velay, Hoxne,Marks Tey and Ossowka do not include pine. The EPICA, MD01–2447, Velay andLake Baikal records are plotted using the timescales published in the originalreports. The AP curves from Hoxne, Marks Tey and Ossowka have been converted toa timescale by correlation with the Velay and MD01–2447 records based onthree tie-points: the rapid increase of AP at the end of the Anglian/Elsterianglaciation; the NAP phase midway through the Hoxnian/Holsteinian Interglacial;and the sudden decrease in AP at the end of the interglacials.

For core MD01–2447, the first part of MIS 11 has been characterisedas showing a long, marked warm period from 426 to 394 kacalled the Vigo Interglacial, which has been correlated with the PraclauxInterglacial at the Velay sites. The Praclaux Interglacial has also been arguedto be similar to the Holsteinian palynological records of northern Europe(Reille and de Beaulieu, 1995;Turner, 1998;de Beaulieu et al., 2001;Desprat et al., 2005; though also seeGeyh and Müller, 2005;Geyh and Müller, 2006;Scourse, 2006) and to the HoxnianInterglacial (Turner, 1998). Theevidence from Hoxne would support this interpretation. Key features of theHoxnian pollen records are the early occurrence ofPicea,the development of mixed oak forest followed by a marked phase ofAbies, and the occurrence ofPterocarya in the later part of the interglacial. Wetherefore conclude that the Hoxnian, Holsteinian, Vigo and Praclauxinterglacials all correlate with the first part of MIS 11.

In the later part of MIS 11, three cold/warm cycles have been recognised incore MD01–2447. These cycles are similar to a series of short-livedcold/warm phases in the Velay sites, which occur after the Praclaux Interglacialand prior to the Bargette cold episode of MIS 10. At Velay, twostadial/interstadial cycles have been named (Chaconac stadial/Jagonas 1interstadial and Coucouron stadial/Jagonas 2 interstadial).

During the stadials, the pollen from core MD01–2447 indicates thateither heath or dry grassland dominated the local vegetation, while at Praclauxthe environment was open with an abundance of steppe taxa. The interstadialsindicate the re-emergence of forest cover with some deciduous woodland. In coreMD01–2447,Pinus andQuercusare prominent, with lesser quantities ofCarpinus andAbies. Pine, however, was argued to beover-represented due to better dispersal ability and buoyancy. The upland siteof Praclaux is characterised during these interstadials by the dominance ofPicea, but also by the presence ofCarpinus,Quercus,Buxus,Fraxinus andTilia. It is suggested that the presence ofCarpinus (up to 10%) may indicate that there was agreater abundance of this taxon at lower altitudes (Reille and de Beaulieu, 1995).

How far north this deciduous woodland stretched is difficult to gauge, dueto the paucity of sites that clearly correlate with these phases. Although thereis no unequivocal palaeobotanical information on the vegetation at Hoxne fromStratum B, the mammalian fauna includes obligate woodland species (e.g. beaver,fallow deer and macaque) providing strong evidence that there must have beensome forest cover.

The evidence from Hoxne, therefore, suggests that the ‘ArcticBed’ of Stratum C and the temperate phase of Stratum B correlate with oneof the cold/warm cycles in the later part of MIS 11, although because of thehiatus between Stratum D and C, it is not clear to which cycle they should beattributed. The problem of recycled pollen in both Strata C and B also makes itdifficult to reconstruct their vegetation histories, other than the survival ofleaves of dwarf birch and dwarf willow, in Stratum C. However, coreMD01–2447 and Praclaux provide clues about the vegetation that might havebeen present at Hoxne, despite differences in latitude, and in the case ofPraclaux in altitude (1100 m, compared to Hoxne at 30 m) between the sites.

Elsewhere in northern Europe, there is little agreement on the correlationof post-Holsteinian temperate events. Most authorities would now agree that theHolsteinian is attributable to MIS 11. If the interpretation favoured here iscorrect, that the Hoxnian and Holsteinian both correlate with the firsttemperate event of MIS 11, then this still leaves the question of whether laterMIS 11 interstadials can be recognised in northern Europe.

One of the best Holsteinian pollen records comes from the lacustrinesequence at Ossowka in eastern Poland (Nitychoruk etal., 2005;Fig. 2).This sequence has been constrained by TL dates of ca 430 ka atthe MIS 12/11 boundary, and the estimation of the duration of the sequence iscalculated from annual laminations in the interglacial part of the record. Likethe pollen sequences in southern Europe, after a stable temperate climate of anestimated 35–39 ka (the Holsteinian), there follows aseries of climatic oscillations with open, cold vegetation alternating with aboreal environment dominated by pine. If the estimated timescale ofNitychoruk et al. (2005) iscorrect, this would imply that the later temperate events in MIS 11 arecharacterised by boreal pine forest in central, northern Europe.

Correlation with other north European sites (e.g. Bilzingsleben andSchöningen) is as yet uncertain due to the varying interpretations that arecurrently put forward (cf.Mania,1995;Urban, 1995,2007;Turner,1998;Bridgland et al.,2006). However, it is worth noting some of the similaritiesbetween the Channel II, Level 4b deposits at Schöningen to Stratum B atHoxne. Level 4b, which includes most of the spears, is assigned to the ReinsdorfB Interstadial (Kolfschoten, 1993).The fauna is dominated by horse and the pollen indicates boreal forestpredominantly of pine, but with some spruce, birch and larch (Urban, 2007).

The differences in the vegetational records from southern to northernEurope would suggest quite a marked climatic gradient between 40° and50° latitude during the later interstadials of MIS 11. A similar pattern hasalso been identified for MIS 5, where the vegetational records for substages 5cand 5a at Grande Pile (France;Woillard,1978) show deciduous woodland, whereas those further north inthe Netherlands, Germany, Denmark and, to a lesser extent, the UK show that thevegetation was dominated by boreal forest (Behre,1989;Turner,1998). Turner suggests that either the phases were too short toallow for the immigration of thermophilous trees, or that there was a realclimatic barrier to the spread of deciduous woodland to the north. This may berelated to circulation patterns in the North Atlantic Ocean, with a southerlyshift in the Gulf Stream, leading to even cooler temperatures in northernEurope. There was also likely to have been a west–east gradient inclimate;Zagwijn (1990) has suggestedthat summer temperatures during substage 5c showed a marked decrease from thesouthwest to the northeast in Europe, unlike substage 5e, where the gradient wasfrom southeast to northwest. Although during substage 5c there seem to be fewvegetational differences between sites in Britain and those further east, whereforests of pine, birch and occasionally spruce were dominant (Behre, 1989), it has been suggested that Britainhad a more continental climate with cold winters, but warm summers(Coope, 1977). If this can beused as an analogue for the late MIS 11 interstadials, then Hoxne might have hadvegetation of boreal forest, but with warm summer temperatures. This conclusionis supported by the faunal evidence from Stratum B at Hoxne.

8. Conclusions

Hoxne is a key site for understanding the Middle Pleistocene sequence ofnorthern Europe and understanding how this correlates with sequences fromsouthern Europe. The site provides a stratigraphic sequence that includes twopost-Anglian temperate phases. The first of these (the Hoxnian) is argued tocorrelate with the first sustained temperate phase in MIS 11 between ca425–395 ka. The second, as represented by Stratum B,is correlated with a later interstadial in MIS 11. These two temperate phasesmay be tentatively correlated with substages 11c and 11a, respectively, whichare evident in the SPECMAP stack (Imbrie et al.,1984;Tzedakis et al.,2001). Alternatively, the acme of the Hoxnian may becorrelated with isotopic event 11.3 and Stratum B with either event 11.23 or11.1 ofBassinot et al. (1994). Theintervening cold episode, represented by Stratum C is correlated with marineisotope substage 11b and may equate to either event 11.24 or 11.22 in the LowLatitude Stack (Bassinot et al.,1994).

Lithostratigraphy, palynology and molluscan data suggest that the sequencesat Clacton, Tillingham and the Lower Gravels to Middle Gravels at Swanscombe canbe attributed to the first temperate event (the Hoxnian). Comparison with thecontinuous palynological records from southern Europe suggests that the Hoxniancorrelates with the Vigo Interglacial of northwest Iberia, the PraclauxInterglacial of the Velay maars sites and to the Holsteinian Interglacial ofnorthern Europe. Stratum B is argued to correlate with either the Jagonas 1 or 2Interstadial from the Velay sites. Reconstruction of the vegetation during theseinterstadials suggests that in northern Europe they were dominated by apine-birch boreal forest, which supported a diverse large mammalfauna.

Traditionally, these faunas have been interpreted as indicating fullyinterglacial conditions. The evidence from Hoxne, therefore, clearly indicatesthat similar faunal assemblages can also occur in environments of interstadialcharacter. This has implications for the biostratigraphical subdivision oftemperate episodes in the Middle Pleistocene based on mammalian evidence (cf.Schreve, 2001a, b).

Hoxne is also an important site for understanding the Lower Palaeolithicoccupation of northern Europe. The archaeological assemblages at Hoxne can nowbe shown to date to an interstadial that has not been previously recognised inBritain. Although there are several MIS 11 sites where fine-grained, organicsediments allow detailed environmental reconstruction, they have all suggestedthat human occupation was associated with deciduous woodland in fully temperateclimate (cf.Ashton et al., 2006). AtHoxne, however, humans can be demonstrated to have lived in a boreal forestenvironment and probably with distinctly cooler winters. This prompts questionsabout the technologies required (clothing, shelters, control of fire) orphysical adaptations needed in order to survive these cooler environments. Therange of environments that humans inhabited during the Middle Pleistocene haslong been the subject of debate (Gamble,1987, 1992;Roebroeks et al.,1992). Hoxne now adds to the small list of sites from theLower Palaeolithic where the human habitat can be reconstructed in more detailand indicates human adaptability to a range of different habitats.

Acknowledgements

We would like to thank the British Museum, the British Academy and the Societyof Antiquaries for funding the fieldwork, and NERC, English Heritage and theWellcome Trust for funding the AAR analyses. Thanks are also due to Andrew andMelanie Banham, Peter Whatling and Suffolk County Council for access to the sites.We are grateful to Robert Symmons and Silvia Bello for producing the illustrationsand to Richard Preece, Roger Jacobi, Charles Turner and Matthew Collins for helpfuldiscussions. The constructive comments of the referees is also acknowledged. Thispaper is a contribution to the Ancient Human Occupation of Britain project, fundedby the Leverhulme Trust.

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