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Vertebrate faunas from the early Middle Pleistocene of East Anglia
... The extinct water vole, Mimomys savini, with rooted teeth, is thought to have given rise to Arvicoia terrestris cantiana, whose cheek teeth lack roots. The former occurs, together with a rich associated fauna (Stuart 1975(Stuart , 1996, at West Runton, whereas it is replaced by A t. cantiana in other faunas such as those from Ostend, Norfolk (Stuart & West 1976), Westbury-sub-Mendip, Somerset (Bishop 1982) and Boxgrove, Sussex (Roberts 1986). Opinion is divided as to whether the faunas with At. cantiana should be assigned to the late Cromerian (Stuart & West 1976) or belong to an additional stage between the Cromerian and the Hoxnian (Bishop 1982). ...
... The occurrence of Macoma balthica demonstrates a post-Middle Tiglian (post-Norwich Crag) age, whereas the absence of other species, for example Acila cobboldiae, Macoma obliqua and Mya arenaria (the last two known only as derivatives), points to a post-Late Tiglian (post-Weyboume Crag age). A rich vertebrate fauna is also known from the West Runton Freshwater Bed (Stuart 1975(Stuart , 1996. Stratigraphically important taxa include Mimomys savini, Pitymys gregaloides, Pliomys episcopalis, Sorex savini, S. runtonensis, Dicerorhinus etruscus, Megaloceros verticomis and M. savini. ...
... Blake (1890) recovered two vole teeth from black silt in a "basin-shaped hollow in the lower part of the Rootlet Bed". Re-examination of these specimens, now in the collection of the British Geological Survey, has confirmed that at least one is Mimomys savini (Stuart 1996). ...
Non-marine molluscan faunas from sites of late Lower and early Middle Pleistocene age in Britain, The Netherlands, France, Germany and Denmark are reviewed. The fauna from the Cromerian stratotype at West Runton shares two extinct prosobranchs (Valvata goldfussiana and Tanousia runtoniana) with that of the Bavel Interglacial at its type locality. Conversely, two other prosobranchs, Lithoglyphus jahni and Fagotia wuesti, characteristic of the Bavelian and earlier periods, are absent at West Runton. These facts suggest that the two sites are close in age but that West Runton is the younger. Moreover, the stratigraphical range of V. goldfussiana in the Netherlands (Middle Tiglian - Cromerian Interglacial I) suggests that West Runton should be placed early in the 'Cromerian Complex'.
British sites assigned to the 'Cromerian' Stage fall loosely into three groups. Group I is characterized by V. goldfussiana, T. runtoniana, Bithynia troscheli and Viviparus viviparus gibbus, and includes West Runton, Kessingland and Sugworth. The fauna of Group 2 with Valvata naticina, B. troscheli, Tanousia cf. stenostoma and Belgrandia marginata is known only from Little Oakley. Group 3 faunas, such as those from Sidestrand and Trimingham, with V. naticina, Bithynia tentaculata and B. marginata, lack distinctive 'Cromerian' elements and have close affinities with certain Hoxnian assemblages. The extinct water vole Mimomys savini has been recorded from sites assigned to each of these three Groups. At Ostend, Norfolk, an undiagnostic molluscan fauna has been recovered from beds that have yielded the vole Arvicola t. cantiana. It would appear from the malacological evidence that two, or more likely several, interglacials, currently assigned to the British Cromerian, are represented.
It is not possible to assign with confidence any of the continental sites discussed to these groups but the faunas from Noordbergum and Exmorra have closest affinities with Group 3, whereas those from I1sselstein and from North Sea borehole E8-4, both with v. goldfussiana. are closest to Group I. The malacological data do not support correlation of Noordbergum ('Interglacial IV') with West Runton but would support a recent suggestion, based on vertebrate evidence, that West Runton is older. It is not possible to correlate West Runton with any known parts of the four Dutch 'Cromerian' interglacials.
... Deposits of the early Middle Pleistocene can be identified on both biostratigraphic and lithostratigraphic grounds. The mammalian and molluscan assemblages of early Middle Pleistocene deposits in Britain and western/central Europe are relatively distinctive and can be used to identify deposits of this age (Stuart, 1996;Preece and Parfitt, 2000;Preece, 2001;Stuart and Lister, 2001). In particular, the evolution of a range of small mammal species is important in identifying early Middle Pleistocene deposits (Stuart, 1996;von Koenigswald and van Kolfschoten, 1996;Preece and Parfitt, 2000). ...
... The mammalian and molluscan assemblages of early Middle Pleistocene deposits in Britain and western/central Europe are relatively distinctive and can be used to identify deposits of this age (Stuart, 1996;Preece and Parfitt, 2000;Preece, 2001;Stuart and Lister, 2001). In particular, the evolution of a range of small mammal species is important in identifying early Middle Pleistocene deposits (Stuart, 1996;von Koenigswald and van Kolfschoten, 1996;Preece and Parfitt, 2000). In eastern England, the key stratigraphic unit for the correlation of the early Middle Pleistocene is the Lowestoft Formation (Bowen, 1999;Lee et al., 2004b), deposited during the Anglian glaciation and correlated with MIS 12, ca. ...
... The deposits with the most convincing evidence for climates consistent with modern-day conditions come from the Cromerian stratotype at West Runton on the north Norfolk coast (Reid, 1890;West and Wilson, 1968;West, 1980;Stuart, 1996Stuart, , 2000. In these deposits (referred to as the 'upper freshwater bed'), a wide range of vertebrate (Stuart, 1996;Preece and Parfitt, 2000;Parfitt, 2008), invertebrate (West, 1980;Coope, 2000, in press;Preece and Parfitt, 2000;Preece, 2001) and isotopic (Davies et al., 2000;Rose et al., 2008;Candy et al., 2010) evidence is indicative of a temperature regime that is consistent with modern-day British climates. ...
Long-term climate records such as SPECMAP and EPICA imply that the early Middle Pleistocene (Marine Isotope Stages 19–13, 780–450ka) was characterised by low magnitude climate cycles relative to the extreme glacial/interglacial cycles of the last 450ka. As the early Middle Pleistocene is the period during which the first known occupation of Britain occurred, understanding the nature of climate cycles in northwest Europe during this period is important. In order to develop a clearer understanding of the pattern of climate change during the early Middle Pleistocene, deposits of this period are divided into four groups that are based on the climatic proxy data they contain. Group 1 deposits are characterised by evidence for interglacial climates that were warmer than the present day. Group 2 deposits are characterised by evidence for interglacial climates that were consistent with the present day with respect to their degree of warmth. Group 3 deposits contain evidence for temperate climates that were cooler than the present day; such deposits possibly reflect the end of an interglacial or interstadial. Group 4 deposits record evidence for extreme climate cooling and widespread permafrost development. This categorisation indicates that during multiple glacial/interglacial cycles the climate of eastern England oscillated between periods that were warmer than the present day, sometimes ‘Mediterranean’ in character, through to periods that were characterised by extreme climate cooling and widespread periglaciation. Despite the climate patterns suggested in the SPECMAP and EPICA records, there is no recognisable difference between the pattern of climate forcing observed in Britain during the early Middle Pleistocene relative to that which occurred during the late Middle and Late Pleistocene. Early human colonisers in Britain during the early Middle Pleistocene were, therefore, subjected to the same extremes of climate as humans during the last 450ka. Consequently, it is probable that the pattern of depopulation during glacials and recolonisation during interglacials, proposed for the last four glacial cycles, is also likely to be true for the period 780–450ka. It is also important to recognise that lithic artefacts are found in association with all four climatic groups, indicating that the presence of humans during the early Middle Pleistocene was not restricted to the climatic peaks of interglacials.
... They contain a molluscan assemblage with Bithynia and other taxa characteristic of moving-water (West, 1980: Table 19), very different from the assemblage reported here from the 'Arctic Fresh-water Bed'. Similar pollen spectra, assigned to Cr IVa and Cr IVb, were obtained from sediment adhering to some of the bones (principally Arvicola) collected by C. Green from a nearby location, now lost (Stuart and West, 1976;Stuart, 1996). The vertebrates from this bed, like the molluscs just described, are completely different from those of the 'Arctic Fresh-water Bed', despite the boreal character of the pollen spectra (Table 5). ...
... Despite evidence from the pollen for boreal forest, the plant macrofossils from the interglacial channel deposits include several strongly thermophilous elements, such as Acer campestre, Cornus sanginea, Trapa natans and Valeriana dioica, indicating considerable summer warmth (West, 1980: 76). The vertebrates from Ostend in the Green/Gurney collections are thought to be penecontemporaneous and similarly indicate temperate conditions (Stuart and West, 1976;Stuart, 1996). From the occurrence of Arvicola (water vole), these seem to date from an interglacial occurring late in the 'Cromerian Complex', possibly equivalent to MIS 13 (Preece and Parfitt, 2000;Preece et al., 2009). ...
... Table 5 Comparison of the small mammal fauna collected or acquired before 1843 by C. Green and A. Gurney from 'Ostend' with that from the 'Arctic Fresh-water Bed' at the same locality. Sediments containing some of the voles collected by Green were subsequently attributed on the basis of palynology to the post-temperate stage of the 'Cromerian' (Stuart and West, 1976;Stuart, 1996). ...
At three sites along the North Norfolk coast, freshwater sediments containing arctic plants have been reported to occur immediately beneath the glacial deposits. The sections at Mundesley and Beeston are no longer visible but a small exposure of this ‘Arctic Fresh-water Bed’ has recently been rediscovered at Ostend near Bacton. The fossils from this deposit are distinctly high boreal/arctic in character, indicating a small, possibly ephemeral, pool in a generally open landscape supporting some dwarf shrubs and conifers. Noteworthy records include three beetle species (Pterostichus brevicornis, P. middendorffi and Helophorus obscurellus) that today have their closest occurrences on either the Kanin or Kola peninsulas in arctic Russia. A Mutual Climatic Range reconstruction on the limited beetle fauna suggests that the mean temperatures of the warmest and coldest months were between 9°C and 11°C and between −36°C and −10°C, respectively. Other noteworthy occurrences include the aquatic pulmonate gastropod Gyraulus rossmaessleri (second and earliest British record), the ostracod Amplocypris tonnensis (earliest British record), ground squirrel (Spermophilus sp.) and the first record of arctic lemming (Dicrostonyx sp.) from the Cromer Forest-bed Formation (CF-bF). The ‘Arctic Fresh-water Bed’ has been assigned to the Bacton Member of the CF-bF and formed during the early Anglian, correlated with Marine Isotope Stage (MIS) 12 of the deep sea record. The Ostend assemblage is therefore important in providing a glimpse of conditions that occurred during a major cold stage in the early Middle Pleistocene.
... Although many hippo fossils are known from this region, very few have secure stratigraphical context. Most occurrences represent isolated bones or teeth, or fragments thereof, found loose on the beach, having eroded out of the cliffs or foreshore (Stuart, 1996). ...
... Although pollen analysis of sediment adherent to museum hippo specimens from Trimingham, Mundesley and Bacton was argued to support a 'Cromerian' age (Stuart, 1986), Stuart (1996) admitted that the later recognition of multiple interglacials within the 'Cromerian Complex' prevented direct correlation to a single temperate episode. In addition, the high frequency of Abies pollen was used to rule out a 'Pastonian' age, since Abies was unknown from the British Early Pleistocene at the time (Stuart, 1986). ...
Although fossil assemblages from the late Early Pleistocene are very rare in Britain, the site of Westbury Cave in Somerset, England, has the potential to address this gap. The mammal fossils recovered previously from the Siliceous Member in Westbury Cave, though few in number, have hinted at an age for the deposits that is as yet unparalleled in Britain. Here, we describe the first bona fide occurrence of Hippopotamus in the British Early Pleistocene, discovered during recent reinvestigation of the Siliceous Member. The hippo fossil indicates a refined biochronological age of ca. 1.5-1.07 Ma for the Siliceous Member and a palaeoclimate that was warm and humid, which accords well with previous palaeoenvironmental inferences. A synthesis of late Early Pleistocene hippo occurrences suggests that the Siliceous Member hippo may have been part of an early colonization of northwest Europe by these megaherbivores, possibly during MIS (Marine Oxygen Isotope Stage) 31. Alternatively, it evidences a currently cryptic northward migration during an even earlier temperate phase. In either case, the Siliceous Member is likely to represent a warm period that has not been recognized previously in the British Quaternary record.
... At other pre-Anglian Arvicola sites, such as Ostend (Stuart and West, 1976;Stuart, 1996), Waverley Wood (Shotton et al., 1993) and the Sidestrand Unio-bed (Preece et al., 2009), the yield of small vertebrate remains is far lower, rendering attribution to either of these Arvicola groups much more difficult. In the case of Ostend, the vertebrates were collected in the 1840s and assigned to the 'Cromerian' on the basis of their pre-glacial context and pollen spectra from sediment adhering to critical specimens (Stuart and West, 1976;Stuart, 1996). ...
... At other pre-Anglian Arvicola sites, such as Ostend (Stuart and West, 1976;Stuart, 1996), Waverley Wood (Shotton et al., 1993) and the Sidestrand Unio-bed (Preece et al., 2009), the yield of small vertebrate remains is far lower, rendering attribution to either of these Arvicola groups much more difficult. In the case of Ostend, the vertebrates were collected in the 1840s and assigned to the 'Cromerian' on the basis of their pre-glacial context and pollen spectra from sediment adhering to critical specimens (Stuart and West, 1976;Stuart, 1996). The molluscan assemblage from Sidestrand differs markedly from other sites in the British 'Cromerian Complex' (Preece et al., 2009). ...
This paper reviews the stratigraphical and environmental contexts of early human occupation and lowland glaciation in Britain and northern Europe. It highlights the fragmentary nature of most of the critical sequences and the inadequacy of existing stratigraphical frameworks. The earliest archaeology known from NW Europe has been reported from Happisburgh 3, a site on the Norfolk coast. The flint artefacts from here were recovered from sediments previously assigned to the Pastonian Stage of the Early Pleistocene. However, the mammalian assemblage from Happisburgh 3 differs from other local sites attributed to the ‘Pastonian’, indicating a younger age within the Early Pleistocene. The sediments have reversed palaeomagnetic polarity and have palynological features inviting correlation with the Leerdam interglacial in the Dutch ‘Bavelian Complex’. The interglacials of the British ‘Cromerian Complex’ are compared and their archaeological record reviewed. At most of these sites, the archaeology includes hand-axes associated with a mammalian assemblage containing the water vole Arvicola but at Pakefield, Suffolk, a core-and-flake industry is associated with the ancestral Mimomys savini. This, together with the even older record from Happisburgh 3, indicates earlier periods of human occupation than previously known from NW Europe. At least five temperate episodes with distinct faunal assemblages are now recognized in the British early Middle Pleistocene but these are unlikely to equate directly with odd-numbered stages of the marine isotope record. This biostratigraphical framework conflicts with a ‘New glacial stratigraphy’ for East Anglia, which proposes glaciations during MIS 16, 12, 10 and 6. Evidence is reviewed from Sidestrand and Trimingham, where datable organic sediments occur directly beneath and above the Middle Pleistocene glacial succession, indicating that it was emplaced entirely during the Anglian glacial stage (MIS 12). A new stratigraphical framework is provided ranking British sites in order of relative age, which is then compared with a revised scheme for the Dutch late Early and early Middle Pleistocene succession. The revisions involve the relocation of the Waardenburg interglacial (‘Interglacial 1’) to the ‘Bavelian Complex’ and the use of the Brunhes-Matuyama palaeomagnetic reversal to define both the base of the Dutch ‘Cromerian Complex’ and the Early/Middle Pleistocene boundary. Moreover, following recent re-interpretation, the Noordbergum interglacial (‘Interglacial IV’) and ‘Glacial C’ are removed from the ‘Cromerian Complex’ and are correlated with MIS 11 and MIS 12 respectively. This scheme imposes greater consistency between the British and Dutch successions, as well as conformity with guidelines recommended by the International Stratigraphic Commission.
... The 'Crag' and 'Forest Bed' deposits have been famous for their fossil mammal remains since the early part of the 19th century (Owen, 1846;Newton, 1882aNewton, ,b, 1886Newton, , 1891Gunn, 1891;Lister, 1996;Stuart, 1982Stuart, , 1996. However, a relatively small proportion of mammalian remains has been collected in situ. ...
... The numerous previous palaeontological studies on the WRFB include: Azzaroli (1953), De Deckker (1979, Harrison (1979), Hinton (1908Hinton ( , 1911Hinton ( , 1926, Holman (1989Holman ( , 1998 Holman et al. (1988), Lister (1993Lister ( , 1996Lister ( , 1998, Meijer and Preece (1996), Newton (1880Newton ( , 1882aNewton ( , 1909, Owen (1846), Parfitt (2008), Preece (2001), Preece and Parfitt (2000), Reid (1882Reid ( , 1890, Rose et al. (2008), Stuart (1975Stuart ( , 1981Stuart ( , 1992Stuart ( , 1996, and West (1980). ...
... In this paper, the taxonomy used differs from that in recent discussions of the WRFB micromammals (e.g. Stuart, 1996; Preece and Parfitt, 2000). The previous taxonomies (Table 1, columns 1 and 2) group successive fossil 'populations' into singlespecies lineages based on the identification of phyletic sequences. ...
... These differences of approach have important implications. For example, the species lists of Stuart and Parfitt (Stuart, 1996; Preece and Parfitt, 2000) have a modern 'flavour', whereas in the alternative taxonomy nearly all of the WRFB species are considered to be extinct (Table 1, column 3). While it is beyond the scope of this paper to resolve the taxonomic issues, we nevertheless provide detailed descriptions, which are illustrated where possible. ...
In this paper, the small mammals recovered from sediments associated with the West Runton Elephant have been analysed and compared with sites in other parts of Europe. Major taxonomical problems are indicated and we suggest ways of utilising such morphological complexity to refine biostratigraphical and chronostratigraphic attributions.The micromammal assemblage from the West Runton Elephant Site so far totals 16 species from the West Runton Freshwater Bed (WRFB). There are strong similarities with the arvicolid spectrum of basal layer H8 in the Koněprusy C 718 profile (Czech Republic), which accumulated under cooler conditions preceding the thermal maximum of an early Middle Pleistocene interglacial. Rare steppic indicators (e.g. Cricetus) at West Runton also imply a somewhat cooler and more continental climate than the present day, but overall the fauna is fully temperate in character. West Runton shares several stratigraphically significant small mammal taxa with Voigtstedt (Central Germany) and morphometric comparisons suggest that Voigtstedt may be slightly younger than West Runton. The presence of Mimomys savini, with a latest occurrence in the early part of Marine Isotope Stage (MIS) 15, together with the palaeomagnetic evidence suggests the WRFB may have been emplaced during the early part of MIS 17. However, given the complexity of the marine isotope curve during the early part of the Brunhes Chron and differences between global and regional climatic evolution, it is difficult to assign the normally magnetised WRFB to a particular Marine Isotope Stage (MIS).
... Recent fieldwork at Sidestrand/Trimingham has shown that biostratigraphically distinct large and small mammal assemblages are present in the Sidestrand Hall Member, as well as the more extensive underlying 'Weybourne Crag' (Table 5). In an earlier account (Stuart, 1996) the water vole molars then available included some that were clearly rooted and therefore attributable to Mimomys. Re-examination of the original specimens and rare Mimomys molars in the new samples has confirmed the initial diagnoses as Mimomys, including Mimomys pliocaenicus, the dominant vole in the 'Weybourne Crag' (Mayhew and Stuart, 1986), but their preservation is quite unlike that of other fossil vertebrates from the Sidestrand Hall Member. ...
... The water vole (Arvicola) is the dominant small mammal in the sieved assemblage. None of the 22 water vole molars show a narrowing of the re-entrant angles at the base of the crown or (Stuart, 1996) included Pliomys episcopalis but this has proved to be a misidentification. Reworked taxa included undetermined shark, Raja clavata (thornback ray), a labrid or sparid (wrasse or sea bream) and the vole Mimomys pliocaenicus. ...
Considerable debate surrounds the age of the Middle Pleistocene glacial succession in East Anglia following some recent stratigraphical reinterpretations. Resolution of the stratigraphy here is important since it not only concerns the glacial history of the region but also has a bearing on our understanding of the earliest human occupation of north-western Europe. The orthodox consensus that all the tills were emplaced during the Anglian (Marine Isotope Stage (MIS) 12) has recently been challenged by a view assigning each major till to a different glacial stage, before, during and after MIS 12. Between Trimingham and Sidestrand on the north Norfolk coast, datable organic sediments occur immediately below and above the glacial succession. The oldest glacial deposit (Happisburgh Till) directly overlies the ‘Sidestrand Unio-bed’, here defined as the Sidestrand Hall Member of the Cromer Forest-bed Formation. Dating of these sediments therefore has a bearing on the maximum age of the glacial sequence. This paper reviews the palaeobotany and describes the faunal assemblages recovered from the Sidestrand Unio-bed, which accumulated in a fluvial environment in a fully temperate climate with regional deciduous woodland. There are indications from the ostracods for weakly brackish conditions. Significant differences are apparent between the Sidestrand assemblages and those from West Runton, the type site of the Cromerian Stage. These differences do not result from contrasting facies or taphonomy but reflect warmer palaeotemperatures at Sidestrand and a much younger age. This conclusion is suggested by the higher proportion of thermophiles at Sidestrand and the occurrence of a water vole with unrooted molars (Arvicola) rather than its ancestor Mimomyssavini with rooted molars. Amino acid racemisation data also indicate that Sidestrand is significantly younger than West Runton. These data further highlight the stratigraphical complexity of the ‘Cromerian Complex’ and support the conventional view that the Happisburgh Till was emplaced during the Anglian rather than the recently advanced view that it dates from MIS 16. Moreover, new evidence from the Trimingham lake bed (Sidestrand Cliff Formation) above the youngest glacial outwash sediments (Briton's Lane Formation) indicates that they also accumulated during a Middle Pleistocene interglacial – probably MIS 11. All of this evidence is consistent with a short chronology placing the glacial deposits within MIS 12, rather than invoking multiple episodes of glaciation envisaged in the ‘new glacial stratigraphy’ during MIS 16, 12, 10 and 6. Copyright © 2009 John Wiley & Sons, Ltd.
... The West Runton coprolite had the highest loss on ignition (Table 1), probably due to plant macrofossils and large amounts of pollen found within it. The calcium content of both wolverine (Gulo gulo) and wolf (Canis lupus) droppings (the latter species being present at West Runton; Stuart, 1996) were similar to those of C. crocuta at 21·91% and 25·90% respectively. ...
... Carnivore species recorded from other parts of the West Runton Freshwater Bed (Stuart, 1996) had to be considered as potential sources of the coprolite material. Although containing similar levels of calcium as C. crocuta droppings, the gross morphologies of the wolf and wolverine droppings are different from those of C. crocuta in that they contain sharp fragments of undigested bone. ...
Coprolites (fossilized droppings) found during the excavation of the West Runton Elephant in 1992 and 1995 were studied and attributed to the spotted hyaena (Crocuta crocuta) on the basis of their distinctive morphology and content. Recent droppings of captive spotted hyaenas were compared with the coprolites in terms of size, morphology and content and were used in simple laboratory experiments devised to improve understanding of their behaviour in depositional environments generally and their mode of burial at this locality specifically. The coprolites are 22–32% larger than the recent droppings, consistent with the interpretation that the European spotted hyaenas of the Cromerian Interglacial were larger than modern African equivalents. The fresh spotted hyaena droppings are hard and durable, sink rapidly through water and are able to withstand considerable trampling into sediment whilst maintaining a coherent form. The droppings have a high threshold entrainment velocity in water, far higher than that for silt or fine sand and consequently it is unlikely that the West Runton coprolites were transported to the site with the host sediment. This suggests three possible histories of deposition. (1) The droppings were deposited around the elephant skeleton (possibly by the same animals which caused damage to the skeleton) and the fine sediment was then washed in around them. (2) They were deposited by hyaenas while fine sediment was being deposited. (3) They were deposited after the sediment was deposited and then they were subjected to faunal disturbance by trampling and scattering. This approach studying the behaviour of, in this case, modern fresh droppings in experimental situations to answer specific questions may be of use in investigating other sites with similar natural depositional settings, not necessarily involving faecal material.
... Moose remains, from CF-bF of Dogger Bank (5) , East Runton (6) , Sidestrand (7) (Azzaroli, 1953 ) and Overstrand (8) (Lister, 1996) can be assigned to C. gallicus. They were collected in or are believed to originate from the exposed Pre-Pastonian and Pastonian levels of the CF-bF, as other mammal remains, such as Megaloceros obscurus, Eucladoceros and typical M. meridionalis (Azzaroli, 1953; Stuart, 1974 Stuart, , 1982 Stuart, , 1996 Lister, 1996 Lister, , 1998 Forsten, 1998). This fauna is here considered to be linked to the Olivola FU. ...
... The mammal assemblage on the whole (M. savini, Mimomys pusillus, Microtus gregaloides , M. meridionalis a.f., M. trogontherii, Palaeoloxodon antiquus, etc.) (Stuart, 1974Stuart, , 1996 Lister, 1996; Lister and Sher, 2001; Stuart and Lister, 2001; Stuart, pers. comm.) is assigned to the Slivia FU. ...
Systematics, taxonomy and phylogeny of Eurasian fossil moose are discussed in order to analyse their distribution in space and time. The largest European collections were studied. We recognise the genus Cervalces, including the chronospecies C. gallicus, C. carnutorum and C. latifrons, as well as the genus Alces, with the species A. alces. Cervalces differs from Alces in the facial area, in the length of the antlers and in the orientation of the palmation. Taking into account as more bibliography as possible, we suggest that the Siberian remains, due to their distance from the type localities, have size ranges and beam proportions a little different from the coeval European ones, so they are regarded as different geographic populations. Cervalces latifrons postremus systematics and chronology have been reconsidered. It results that it was present only in Siberia during the penultimate glaciation and was of the same body size as typical C. latifrons. It is likely that A. alces is not the direct descendant of the last European Cervalces, but its origin is still an open question. The present analysis provides a clearer picture of the geographical and chronological distribution of Cervalces and Alces.
... Four key localities form the core of the present study, between them yielding the richest stratified mammal faunas from the early Middle Pleistocene of Britain. Two are sites of the Cromer Forestbed Formation: Pakefield (Suffolk) (Stuart and Lister, 2001; Parfitt et al., 2005), and the type Cromerian West Runton Freshwater Bed (Norfolk) (Stuart, 1975Stuart, , 1996). The others are the Calcareous Member of the cave infill at Westbury-sub-Mendip (Somerset) (Bishop, 1982; Andrews et al., 1999), and the hominid locality of Boxgrove (West Sussex) (Roberts and Parfitt, 1999) (Fig. 1). ...
... At West Runton, Stuart (1974 Stuart ( , 1982 Stuart ( , 1996) and Sala (1986) recorded B. schoetensacki. In size, dental remains are generally within the main Westbury and Pakefield ranges (Figs. ...
Rhinoceroses (Stephanorhinus) and large bovids (Bison, Bos) from the early Middle Pleistocene of Britain are analysed with regard to their taxonomy, biometric variation, and possible biochronological significance. The localities considered are West Runton (type Cromerian), Pakefield, Westbury-sub-Mendip, and Boxgrove. The samples include important, previously undescribed material. Among the rhinos, in addition to the common Stephanorhinus hundsheimensis, upper dentitions from Pakefield and West Runton provide hints of undescribed taxa with affinity to, but distinct from, S. etruscus and S. hundsheimensis. At Boxgrove, a further rhino cf. S. megarhinus occurs, corroborating the chronological extension of this ‘Pliocene’ species into the Middle Pleistocene. A small bison referable to B. schoetensacki occurs at all sites, but in the Westbury Yellow Breccia (upper interglacial level) it is accompanied by a larger bison which may be B. priscus. Dental material from Pakefield provides evidence of additional, smaller bovid species, with features recalling Leptobos and Bubalus, but their identity is unclear; there is also larger material probably representing Bos, which if confirmed would be the oldest record of aurochs in the British Isles.
... Simultaneously, all those authors (García, 2003;García and Arsuaga, 2011;García et al., 2023) (Stuart and Lister, 2010). Originally classified as Martes sylvatica (Newton, 1880), it was soon reidentified as M. martes (Newton, 1880(Newton, , 1882(Newton, , 1891Reynolds, 1912;Kurtén, 1968;Stuart, 1974Stuart, , 1975Stuart, , 1981Stuart, , 1982Stuart, , 1988Stuart, , 1992Stuart, , 1996. It was never, however, described in detail, and even a provisional look at the relatively narrowly spaced mental foramens or the weak metaconid of m1 allows this determination to be seriously doubted. ...
... The speleoid lineage begun with the Plio-Pleistocene Etruscan bear (Ursus etruscus Cuvier, 1823), which lived between 5.3 Myr to 100 kyr (Kurtén, 1976;Argant and Crégut-Bonnoure, 1996;Rabeder et al., 2000;Petrucci and Sardella, 2009;Wagner, 2010) and continued with U. savini Andrews, 1922, the Early Pleistocene ancestor of cave bears and brown bears (Kurtén, 1976;Loreille et al., 2001). The immediate precursor of the cave bear was Deninger's bear (Ursus deningeri von Reichenau, 1904), a species restricted to Pleistocene Europe between 1.8 Myr to 100 kyr (Stuart, 1996;Königswald and Heinrich, 1999). Cave bears disappeared before the onset of the LGM (Pacher and Stuart, 2009;Bocherens et al., 2014;Baca et al., 2016) due to a poorly understood combination of factors, such as competition with and predation by other mammals, climatic deterioration, and loss of genetic diversity. ...
A recent palaeontological excavation in Muierilor Cave, southern Carpathians (Romania), has recovered one of the largest populations of Late Pleistocene wolves (Canis lupus Linnaeus, 1758; MNI =25) documented in Eurasian cave deposits. Many individual nests with wolf bones, including fully articulated skeletons, were found in a remote passage of the cave suggesting a communal behavior similar to that of modern conspecifics. Radiocarbon dates indicate constant use of the cave by these canids throughout the Marine Isotopic Stages 3 (MIS 3; 59‒29 cal ka BP). This deposit affords an opportunity to study the extraordinary resilience and adaptability of C. lupus following the extinction of other large carnivores at the onset of the Last Glacial Maximum (LGM). Using taphonomic, stable isotope and radiocarbon data, we make a case that these canids preyed/scavenged upon the cave bears (Ursus spelaeus Rosenmüller, 1794; MNI =82) that used the same cave for hibernation, along with other Ice Age megafauna such as cave lions (Panthera spelaea Goldfuss, 1810; MNI =4) and cave hyenas (Crocuta spelaea Goldfuss, 1823; MNI =3), which were also found in the same bone assemblage.
... The evolutionary characteristics of the small mammals from the Pekla section suggests that this fauna could be correlated with faunas from the fluvial deposits of the VI Dniester terrace in Kolkotova Balka, the stratotype section of the Tiraspolian faunal assemblage, situated near the city of Tiraspol (Moldova) and in the Levada section (VI terrace of Dniester River) (Alexandrova 1976;Markova 2016;Markova et al. 2021;Markova and Puzachenko 2018). The Novokhopersk and the Ilyinka 5, 4, 2 faunas (Don basin) possess a similar species composition (Stuart 1996;Maul and Parfitt 2010). Maul and Parfitt (2010) correlate this fauna with the beginning of the Cromer Interglacial II. ...
The history of the early Middle Pleistocene small mammal faunas of Eastern Europe is very complicated. The early Middle Pleistocene which spanned from the Brunhes-Matuyama transition (772.9 ka BP, within MIS 19) till the beginning of the Likhvin Interglacial (424 ka BP, MIS 11) includes a number of interglacials and glaciations. Rodent species of the Tiraspolian faunal assemblage were found in the Chaudian fluvial deposits of the Cape Pekla section (northern coast of the Taman Peninsula). The evolutional level of the Pekla rodents are similar to those from the stratotype section of the Tiraspolian faunal assemblage in the Kolkotova Balka in Moldova (MIS 17) , which includes Eolagurus sp., Mimomys savini, Microtus (Terricola) arvalidens, Microtus (Alexandromys) ex gr. oeconomus and other species. The Pekla fauna also resembles the rodent fauna from famous English West Runton Freshwater Bed locality formed during the Cromerian Interglacial II and some other East and West European faunas. In the current work the entire loess-paleosol sequence of the Pekla section was described with five paleosols from the Middle to the Late Pleistocene. The sequence reflects the complexity of climatic fluctuations from the early Middle Pleistocene to the Holocene.
... The CB bone remains are the most numerous of the large mammal species. Bones, teeth, and genetic research have shown that at least four different CB lineages existed before their extinction in the middle Pleistocene (11). ...
The extinct cave bear (CB) is often depicted as a large heavy animal with a prominent, massive skull and severely shortened pelvic limbs. The size and robustness of the CB are comparable to those of the largest living ursids. Great similarities between CB and the brown bear (BB) species prompted our morphometric comparison. The goal of this study was to elucidate the potential differences in the morphometric skull characteristics of CB and BB species. Craniometric measurements were performed on the skulls of both bear species and were compared to identify craniometric features indicative of possible adaptations in both bear species. The results revealed a marked difference in skull size; however, the shape of the CB cranium is generally known to be similar to that of the BB. In CB, the total length of the skull was approximately 1.5 times longer, and the external bony nasal aperture was larger due to the relatively shorter nasal bones. The nose length and median palate length were relatively longer in the CB, and the infraorbital foramen was located more caudally and closer to the zygomatic process. The infraorbital channel of the CB was located over to the second superior molar roots whereas in BB the latter extends beyond the roots of the first to the second superior molar roots. A marked difference was the non-existence of the three most anterior premolars in the maxilla and mandible of the CB, Higher body of the mandible, which, together with the extensive biting surfaces of the cheeks teeth, indicates a predominantly plant-based diet of the CB. The braincase length was considerably shorter in the CB, resulting in a relatively small neurocranium volume. The pronounced frontal fossa in the CB skull continues caudally into a strongly developed frontal area, which gives the CB skull a prominent steep profile. In summary, comparative craniometry showed that CB had a smaller neurocranial volume and had herbivore-adapted jaws and teeth. These metric features of the head skeleton may be related to a lower adaptability to extreme climatic conditions to which they were exposed to during the last Pleistocene glacial period, which may have contributed to their extinction. Key words: head skeleton; skull; mandible; craniometrical features; bearMORFOMETRIČNE ZNAČILNOSTI SKELETA GLAVE JAMSKEGA IN RJAVEGA MEDVEDA: PRIMERJALNA ŠTUDIJAIzvleček: Izumrli jamski medved je pogosto upodobljen kot robustna, težka žival z masivno lobanjo in izrazito skrajšanima medeničnima okončinama. Po velikosti ga lahko primerjamo z največjimi vrstami današnjih medvedov. Domnevne podobnosti med jamskim in rjavim medvedom so spodbudile našo primerjalno študijo, s katero želimo pojasniti morebitne razlike v morfometričnih značilnostih lobanj, iz česar je mogoče sklepati na možne prilagoditve pri obeh vrstah. Oblika lobanje je pri jamskem medvedu precej podobna obliki pri rjavem medvedu, ugotovili pa smo izrazito razliko v velikosti. Pri jamskem medvedu je celotna dolžina lobanje približno 1,5-krat daljša, zunanja koščena odprtina vhoda v nosno votlino pa je zaradi razmeroma krajših nosnih kosti obsežnejša. Dolžina nosu in mediana nebna dolžina sta pri jamskem medvedu daljši. Podočnična odprtina leži nekoliko kavdalneje in bližje ličnemu loku. Posledično se podočnični kanal pri jamskem medvedu nahaja le nad koreninami drugega molarja, medtem ko slednji pri rjavem medvedu sega še nad korenine prvega molarja. Razlika je bila opazna tudi v neizraslih prvih treh premolarjih tako v zgornji kot spodnji čeljusti jamskega medveda ter izrazitejšem telesu spodnje čeljustnice. Vse to, vključno z obsežnimi griznimi ploskvami molarjev, nakazuje pretežno rastlinsko prehrano. Dolžina možganske vot- line je pri jamskem medvedu opazno krajša, kar se odraža v sorazmerno majhni prostornini. Globoka čelna jama se kavdalno strmo nadaljuje v močno razvito čelno področje. Manjša prostornina možganske votline, rastlinski hrani prilagojene čeljusti in površine zob ter nekatere druge metrične značilnosti okostja glave bi lahko bile vzročno povezane z nižjo prilagodljivostjo ekstremnim podnebnim razmeram, katerim je v zadnjem pleistocenskem ledeniškem obdobju jamski medved neuspešno kljuboval in kar bi lahko znatno prispevalo k njegovemu izumrtju.Ključne besede: okostje glave; lobanja; spodnja čeljustnica; kraniometrične lastnosti; medved
... There is a commonly accepted opinion concerning the descendance of U. spelaeus from Ursus etruscus Linnaeus, 1758 (Bombiţă, 1954;Viehmann, 1973;Domşa and Popa, 1993), whereas there is one reference in Romania (Maniu, 1951) regarding U. spelaeus's direct ancestor, Ursus deningeri Richenau, 1904, as several studies from other regions revealed (Trimmel, 1968;Mazza and Rustioni, 1994;Kurten, 1976;Stuart, 1996;Pacher and Stuart, 2009). The reason could be the rarity of U. deningeri in Romanian sites (cf. a single fossil mentioned in Constantin et al., 2001). ...
Cave bears are among the best studied Upper Pleistocene extinct
species due to the high abundance of their fossils in caves across Europe. Ursus minimus (Pliocene) was the common ancestor of the the cave, brown and Asiatic black bears. The speleoid and the arctoid lineages split 1.6 million years ago (Bon et al., 2008) and cave bears became extinct around 27,500 years ago (Pacher and Stuart, 2009). Until recently, the cave bear’s distribution was thought to be restricted to the European continent, but the
study of Knapp et al. (2009) confirmed the existence of cave bears on the Asian continent. Morphological studies based on dental features (Rabeder, 1995) that later were confirmed by molecular results (Hofreiter et al. 2004), proposed two distinct species within the European cave bear group: U. spelaeus and U. ingressus (Rabeder et al., 2004). The first species corresponds to the western clade of the group, while the second species has
an Eastern European distribution. For reasons of consistency and lack of molecular data for the Romanian cave bears, the term U. spelaeus (sensu Rosenmüler and Heinroth, 1794) will be used throughout this study.
... These studies have defined a series of at least five separate biostratigraphical assemblages, based primarily on mollusc and small mammal fossils, which have been proposed to represent five separate temperate episodes (Preece and Parfitt, 2012), not all of full interglacial status. The separation of these assemblages is based upon the presence and absence of several species, but the clearest (and most widely cited) distinction is between those deposits that have yielded the extinct water vole Mimomys savini, which pre-date deposits that contain its descendant species Arvicola terrestris cantiana (Stuart, 1996;Preece andParfitt, 2000, 2012;Stuart and Lister, 2001). Amongst other features, Mimomys is characterized by rooted molars whereas Arvicola has continuously growing molars. ...
Research over the last two decades has revealed a rich record of Lower Palaeolithic occupation in Britain before 450 000 years. Acheulean industries (Mode II) first appear in the later part of the early Middle Pleistocene [Marine Isotope Stage (MIS) 19-13]. This paper reviews: (i) the age of the earliest Acheulean in Britain; (ii) the climates under which the earliest Acheulean industries occur, as recorded by the palaeoecological assemblages that are found at key archaeological sites; and (iii) the spatial and temporal pattern of regional climate change, i.e. the magnitude of glacial/interglacial cycles that occurred as a backdrop to the first arrival of Mode II archaeology in Britain. This review suggests that in Britain, the earliest Acheulean populations arrived during MIS 13 and that early occupation occurred under a range of climatic/environmental settings but frequently under post-temperate late interglacial or interstadial-type cool boreal environments. Furthermore, this review shows that the pattern of climate forcing in Western Europe during MIS 13-12 was not analogous to the interglacial cycles of the past 450 000 years as latitudinal climate gradients appear to have been less pronounced. The paper concludes by discussing the significance of these observations for understanding the arrival of the earliest Acheulean in Britain. Copyright (C) 2015 John Wiley & Sons, Ltd.
... The Donian till is overlain by a complicated loess-soil series with 4 paleosols. (Rink et al., 1996;Stuart, 1996;Maul and Parfitt, 2010). The locality is the stratotype of the Cromerian complex. ...
By now the abundant European small mammal data have been collected from the deposits of the first part of the Middle Pleistocene (MIS 18 e MIS 11). The significant number of global climatic events (glaciations and interglacials) corresponds to this period. We analyzed the evolutional changes of small mammals of the Middle Pleistocene, revealed the reactions of small mammals on the different climatic events (glaciations, interglacials) and correlated the Western and the Eastern European faunas. The collected data also help us to divide the geological deposits age and permit to reconstruct the palae-ogeographic picture of the past. The obtained results may serve as an important component for compiling biostratigraphic schemes of the Middle Pleistocene and the Pleistocene in general.
... comm.; Stuart and Lister, 2010). Along with the mammoth bones, some of which had been gnawed by hyaenas, were a number of associated hyaena coprolites scattered on and around the skeleton (Stuart, 1996;Larkin et al., 2000), one of which was used for this analysis (see Larkin et al., 2000, for location of coprolites). An edited volume summarizes much of the work done on the CF-bF, including the WRFB. ...
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... As originally conceived, palynological analysis of interglacial deposits could be used to link any deposit into the vegetation history of a specific interglacial and, therefore, to allow correlation of that unit with a specific warm stage (West, 1970). Although pioneering in its attempt to produce an over-arching terrestrial stratigraphy, it is now recognised that the scheme it produced was too simplistic and that some interglacials had such similar vegetational histories that they could not be readily differentiated (see Sutcliffe, 1975;Green et al., 1984;Bridgland, 1994;Stuart, 1996;Bowen, 1999;Keen, 2001;Schreve, 2001aSchreve, , 2001bThomas, 2001). Faunal biostratigraphy (Keen, 2001;Schreve, 2001a), radiometric dating (Gasgoyne et al., 1981;Rowe et al., 1999;Candy and Schreve, 2007;Preece et al., 2007) and relative dating (Bridgland, 1994(Bridgland, , 2000Penkman et al., 2007Penkman et al., , 2011 have now shown that the interglacial stages that were proposed on the basis of pollen biostratigraphy conflated deposits of two or more distinct interglacials into a single pollen-defined interglacial episode. ...
Interglacials of the Quaternary Period are currently the focus of a great deal of attention within the scientific community. This is primarily because they play a vital role in distinguishing between “natural” and “human” climate change in the current interglacial and in understanding how the Holocene would evolve in the absence of anthropogenic greenhouse warming. In this respect, Marine Isotope Stage 11 (MIS 11, ca 410,000 yr BP) is one of the key interglacial stages of the past 450,000 yr. The pattern of insolation variability that occurs during MIS 11 matches that which occurs in the Holocene more closely than in any other warm stage of the past half a million years. In addition there is now an extensive range of evidence for MIS 11 palaeoclimates and palaeoenvironments from marine, ice core, lacustrine and terrestrial sequences. The aim of this paper is to provide a comprehensive review of the current state of our understanding of MIS 11. This is the first paper to provide a detailed review of MIS 11 that incorporates the wide range of marine, ice core, long lacustrine and terrestrial records that have been generated over the last ten years since the last major overview. Crucially, it is the first review of MIS 11 that incorporates a detailed synthesis of the high-resolution terrestrial sequences of western and central Europe. This paper, therefore, provides a holistic integration of a diverse range of proxies and archives to provide a detailed understanding of the expression of MIS 11 in the Earth system. In particular the review focuses on: (1) the climatic background of MIS 11, (2) the robustness of the identification of MIS 11 in a diverse range of sequences, (3) the climatic structure of MIS 11, (4) the magnitude of warmth that occurred in this warm stage, (5) MIS 11 sea level magnitude and variability, (6) the duration of MIS 11, (7) evidence for abrupt climatic events within the interglacial of MIS 11 and (8) precipitation patterns and trends during this interglacial. The paper concludes by considering how useful MIS 11 is as an analogue for Holocene climates and compares it with other proposed analogues, such as MIS 19, with particular reference to the “early anthropogenic” hypothesis.
... The ectothermic vertebrate fauna (fishes, amphibians, reptiles) lacks southern thermophiles and xeric indicators (Boehme, 2010). For example, the European pond tortoise (Emys orbicularis), present in many interglacial faunas from southern Britain (Stuart, 1979(Stuart, , 1996, is absent from the WRFB. Boehme estimates mean July temperatures between 16 and 17 C, similar to the present day, but with cooler mean annual temperatures, c. 6e8 C (due to cooler winters), compared to 9.9 C today. ...
... The site lies on the north coast of Norfolk in (SE England) and the sediments, which are exposed at the base of steep gravel cliffs, are approximately 600,000-700,000 years old. During 1995 the skeleton of a steppe mammoth (Mammuthus trogontherii) was excavated by the Norfolk Archaeological Unit and the Castle Museum, Norwich (Ashwin and Stuart 1996;Stuart 1996). The WFRWB is comprised of a layer of organic rich detrital silt, approximately one metre thick, overlain by a 19 metre sequence of marine and freshwater sands and silts that make up the cliffs. ...
The long‐term survival of many material types in the archaeological record is contingent on the objects’ burial in favourable environments with which they can achieve some degree of equilibrium. This is especially true of organic artefacts. However, many burial environments evolve over time so that the sediments from which objects are eventually excavated may differ from those they experienced in the past. Many organic finds are so sensitive to changes in their chemical environment that their mere survival indicates that conditions have not changed dramatically since they entered the archaeological record. These materials are of limited use in understanding the history of an archaeological site’s development. Skeletal materials, however, are composites comprising organic and inorganic phases in intimate association. Although subject to microbial and chemical degradation processes they differ from non‐mineralized organics in that they persist in the archaeological record for thousands of years in a wide range of environmental conditions. These different degradation pathways leave very well‐characterized changes in the mineralized tissues that can be used to identify different “diagenetic trajectories” and thus different environmental conditions. This work presents an SEM and chemical examination of bone, dentine and antler from three very different wet sites and compares these to both experimentally buried bones and skeletons with known burial histories. It is thus possible to reconstruct the burial histories of the finds from unknown environments.
... The direct precursor of the Late Pleistocene cave bear was very probably Ursus deningeri, known from many localities of Middle Pleistocene age (e.g. Reichenau 1906;Stuart 1996;K¨onigswald & Heinrich 1999). The transition between Ursus deningeri and the cave bear Ursus spelaeus is generally considered to have occurred at around the beginning of the last Interglacial (Rabeder et al. 2000). ...
The cave bear (Ursus spelaeus) was one of several spectacular megafaunal species that became extinct in northern Eurasia during the late Quaternary. Vast numbers of their remains have been recovered from many cave sites, almost certainly representing animals that died during winter hibernation. On the evidence of skull anatomy and low δ15N values of bone collagen, cave bears appear to have been predominantly vegetarian. The diet probably included substantial high quality herbaceous vegetation. In order to address the reasons for the extinction of the cave bear, we have constructed a chronology using only radiocarbon dates produced directly on cave bear material. The date list is largely drawn from the literature, and as far as possible the dates have been audited (screened) for reliability. We also present new dates from our own research, including results from the Urals. U. spelaeus probably disappeared from the Alps and adjacent areas – currently the only region for which there is fairly good evidence –c. 24 000 radiocarbon years BP (c. 27 800 cal. yr BP), approximately coincident with the start of Greenland Stadial 3 (c. 27 500 cal. yr BP). Climatic cooling and inferred decreased vegetational productivity were probably responsible for its disappearance from this region. We are investigating the possibility that cave bear survived significantly later elsewhere, for example in southern or eastern Europe.
... Laplana, pers. comm., 2000), as occurs in the fossil species Mimomys savini (Sánchez, 1996;Stuart, 1996;Cuenca-Bescós, Laplana & Canudo, 1999). Palaeontological data strongly suggest that M. savini is the sister group of the genus Arvicola (see Chaline & Graf, 1988 and references therein). ...
The Palaearctic genus Arvicola includes two species: the south-western water vole A. sapidus, and the northern water vole A. terrestris. The latter has semiaquatic and/or subterranean populations, while populations of A. sapidus are always semiaquatic. According to the current phylogenetic and palaeontological data, adaptation to semiaquatic life is plesiomorphic for the genus Arvicola. We studied the ontogenetic allometry of skull and long bones of the semiaquatic A. sapidus, a semiaquatic population of A. terrestris (A. t. italicus), and two fossorial populations of A. terrestris (A. t. scherman and A. t. monticola). Animals from fossorial populations were smaller than were those from semiaquatic populations. We found that most of the ontogenetic allometric exponents of characters linked to digging in the skull and in the long bones were significantly higher in A. t. monticola, a fossorial clade, than they were in the semiaquatic populations. On the other hand, there may have been an evolutionary lag between invasion of the hypogeic habitat and the acquisition of fossorial adaptations in A. t. scherman. We showed statistically that the morphological differences linked to the invasion of a hypogeic habitat are already present in juvenile animals and, according to these results, suggest that these morphological differences are the direct expression of genetic changes rather than the outcome of epigenetic factors of mechanical origin. Moreover, we tried to ascertain whether the apomorphic shape of the skull and long bones in the fossorial populations of A. terrestris (compared with the primitive condition that would have been retained by the semiaquatic A. sapidus) are the outcome of a heterochronic process. Optimization by squared change parsimony supported the hypothesis of an apomorphic reduction of body size linked to the invasion of the subterranean habitat. The comparison of the ontogenetic trajectories of both skull shape and long bone shape suggested that a heterochronic process was involved in this morphological transformation. By using the ‘clock model’ method, this mechanism was identified as ‘accelerated dwarfism’ affecting both the skull and long bones. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 87, 381–391.
... Palaeoloxodon antiquus (straight-tusked elephant) X (X) X X X X Mammuthus trogontherii (mammoth-steppe) X X (X) X Total 12 12 11 12 11 12 10 5 'OIS 17' is based on the West Runton Freshwater Bed, Norfolk (Stuart, 1996); 'OIS 15' on Pakefield/Kessingland, Suffolk (Stuart and Lister, 2001); 'OIS 13' on Boxgrove, Sussex and Westbury-sub-Mendip, Somerset (Andrews et al., 1999;Roberts and Parfitt, 1999); OIS 11 on Swanscombe, Kent and Hoxne, Suffolk (Sutcliffe, 1964;Stuart et al., 1993;Schreve, 2001); OIS 9 on Purfleet, Essex (Schreve, 2001); OIS 7 (later part) on Aveley, Essex and correlated deposits (Schreve, 2001); OIS 5e on Barrington, Cambridgeshire, Trafalgar Square, London and correlated deposits (Stuart, 1976); OIS 1 on Star Carr, 'Yorkshire', ca. 9500 BP (Fraser and King, 1954;Legge and Rowley-Conwy, 1988). ...
Interactions between grazing animals and vegetation are assessed from three temporal perspectives: millions, thousands and hundreds of years. Data abundance and quality are highest for recent time periods, but geological data provide a background to the understanding of present-day grazing–vegetation interactions. The Quaternary glaciations and recent anthropogenic influences have contributed to the loss of European mega-herbivores. The geological record from the Eemian interglacial in Denmark suggests that presence of elephant and rhinoceros did not create widespread openings in forest cover. Large populations of giant deer in Ireland became extinct 11 000 years ago. We propose a theory that the giant deer were sufficiently abundant to convert juniper scrub communities into open grassland at a regional scale. The balance between grazers and browsers has undergone continuous change during the last 10 000 years with significant consequences for forest composition and structure. Hunting statistics and archival records permit crude reconstructions of population dynamics for certain ungulate species. High resolution pollen analysis and long-term monitoring generate reconstructions of vegetation that can be compared with fluctuating grazing pressure during the last few hundred years. Such data can be used to validate simulation models of grazing–vegetation interactions.
... British early Middle Pleistocene localities with important largemammal faunas in stratified context are the type Cromerian West Runton Freshwater Bed (Norfolk) (Stuart, 1975(Stuart, , 1996, Pakefield, including Kessingland and Corton (Suffolk) (Stuart and Lister, 2001;Parfitt et al., 2005), the Calcareous Member at Westbury-sub-Mendip (Somerset) (Bishop, 1982;Andrews et al., 1999), and Boxgrove (West Sussex) (Roberts and Parfitt, 1999) (Fig. 1). Smaller assemblages are known from Sugworth (Oxfordshire) (Stuart, 1980) and Little Oakley (Essex) . ...
The taxonomy and biostratigraphic significance of horses (Equus), deer (Capreolus, Dama, Cervus, Cervalces, Praemegaceros, Megaloceros) and pigs (Sus) from key British early Middle Pleistocene sites are investigated using a biometric approach. The sites compared are West Runton (type Cromerian), Pakefield, Little Oakley, Westbury-sub-Mendip, and Boxgrove. The samples include a significant amount of previously undescribed material. Stenonid horses dominate at Pakefield and West Runton, caballines at Westbury and Boxgrove, corresponding to the Mimomys/Arvicola division. The deer of Boxgrove include relatively large roe and small red deer, the latter distinguishing it from the Westbury Pink Breccia. The poorly-known giant deer Praemegaceros dawkinsi spans both the Mimomys (Little Oakley, Pakefield) and Arvicola (Boxgrove) zones of the Cromerian Complex. There is a marked shift in cervid abundance from predominant megacerines (Mimomys zone) to predominant red deer (Arvicola zone).
... However, our understanding of the sequence of temperate stages between the Matuyama/Brunhes boundary and the Anglian (¼Elsterian) glaciation in Britain (the 'Cromerian Complex', c. 0.78e0.5 Ma) has been greatly refined over the past decade ( Parfitt, 2000, 2008; Stuart and Lister, 2001). Extensive large mammal faunas are known from the type Cromerian West Runton Freshwater Bed (Norfolk), and the Rootlet Bed and associated deposits at Pakefield, Kessingland and Corton ('Pakefield'), Suffolk (Stuart, 1996; Stuart and Lister, 2001; Parfitt et al., 2005; Breda et al., in press; Lewis et al., in press; Lister et al., in press). A recent synthesis of the palaeoecology of the type Cromerian interglacial at West Runton (Stuart and Lister, in press) indicates temperate climate, high precipitation and low seasonality, typical of oceanic, mid-latitude Europe, supporting a diverse ecosystem dominated by forest but with productive open areas as well. ...
Large-scale fluctuations in global climate and resulting changes in ecology had a profound effect on human evolution and dispersal. Though hominin remains are scarce, studies focussing on the more abundant records of fossil land mammal communities can contribute greatly to our knowledge of the palaeoenvironmental circumstances that influenced and directed the global spread of hominins. To produce a comprehensive and accurate account of the evolution of western Palaearctic habitat diversity between 2.6 and 0.4 Ma BP, information generated from large mammal communities from 221 key sites has been included in this study.
The giant, short-faced hyena Pachycrocuta brevirostris was the largest Hyaenidae ever existed and the one that perfectly embodied the distinctive bone-cracking adaptations of this mammal family. Its dispersal into Europe is regarded as a biochronological marker of the Late Villafranchian at ~2.0 Ma, and its potential ecological interactions with other carnivorans and early Homo populations diffusing Out of Africa have given rise to extensive discussions. Nevertheless, our comprehension of the extinction of P. brevirostris remains vague. Here, we first critically evaluate the European fossil record of the species and then we review the whole Epivillafranchian and Galerian Hyaenidae record, including P. brevirostris, Crocuta crocuta and "Hyaena" prisca. Biometric comparisons with other extinct and extant bone-cracking hyenas are carried out.
In contrast to a common view, we recognize that there is neither evidence of a persistence of P. brevirostris beyond the Early-Middle Pleistocene boundary, nor of a coexistence between the giant hyena and C. crocuta. The replacement between the two species, which was also accompanied by the arrival of "H." prisca, occurred at ~0.8 Ma and can serve as a marker of the Epivillafranchian–Galerian turnover, part of the faunal renewal that reflects the response of mammal communities to the Early–Middle Pleistocene Transition. Moreover, we clarified that Pliocrocuta perrieri and "H." prisca were different species, and that the latter was relatively more widespread than often assumed, being recorded from localities spanning in age almost the whole Middle Pleistocene and even the early Late Pleistocene.
Sackdilling Cave is a karst fissure filled with a fossiliferous breccia including abundant fossil remains, mostly snails and small mammals. The taxonomic status of mustelid remains from this cave was previously regarded as controversial and, consequently, different authors listed different species from the locality. A detailed revision of the material shows the presence of five mustelid species: Meles sp., Martes vetus, Mustela strandi, Mustela palerminea, and Mustela praenivalis. Among them, especially noteworthy is the presence of the type specimen of Martes vetus. The Sackdilling holotype resembles Martes foina and differs from that of Martes martes in a short and broad viscerocranium; wide and less extended forward incisor row; compressed, short and wide temporal region; large, strongly inflated and convex tympanic bullae; and narrow P3 with weak lingual bulge. Some dental characters are specific for Martes vetus and show some intermediate values between Martes foina than Martes martes. Among them are the P4 protocone length and the M1 trigon length. The revised material from Sackdilling Cave of Mustela palerminea and Mustela praenivalis was compared with that from other Early and early Middle Pleistocene sites of Europe and showed the presence of intermediate characters. Subsequently, based on the entire mustelid assemblage and other faunal elements (mainly rodents), the age of this fauna was estimated as ca. 0.9-0.7 Ma.
The main evolutionary stages of formation of the faunas of small mammals of the Middle Pleistocene in Europe, belonging to the interval ~0.76−0.42 mln years, are described. Based on the analysis of the species composition of the faunas of this interval, six main phases of development of small mammals, corresponding to the main climatic events (interglacials, glaciations) during MIS 18 – MIS 12 are identified. Availability of data for some intervals of the first half of the Middle Pleistocene is not too large. Especially this applies to the cold stages of the first half of the Middle Pleistocene – the Don and Oka (or, that is the same, the Elster and Anglian) glaciations. In several cases the difference in time of the appearance of a number of taxa in Eastern and Western Europe was revealed. So, Arvicola cantianus appeared in Western Europe earlier than in Eastern Europe. This phenomenon can be explained by different rates of evolution in different parts of Europe, and, possibly, insufficient number of related data (geological, geochronological, and paleontological), which subsequently allow to determine the age of a number of localities. The analysis of correlation of the East European and West European faunas of small mammals was carried out. The biostratigraphic scheme and the map of localities of small mammals of the Middle Pleistocene during MIS 18 – MIS 12, i.e. in the first half of the Middle Pleistocene on the international scale (early Neopleistocene according to the scale RISK), were built. The obtained data allows to substantiate more accurately and to clarify the geological age of the deposits, and to reconstruct the palaeogeographical setting of the considered periods. They are also an important component in the establishment of biostratigraphic schemes of the Middle Pleistocene and the Pleistocene in general.
В статье рассмотрены основные эволюционные этапы становления среднеплейстоценовых фаун
мелких млекопитающих Европы, относящихся к интервалу ~0.76−0.42 млн лет. На основании
анализа видового состава фаун этого интервала удается выделить шесть основных фаз развития
мелких млекопитающих, отвечающих основным климатическим событиям (межледниковьям, оле-
денениям) на протяжении МИС 18 – МИС 12. Обеспеченность данными некоторых интервалов
первой половины среднего плейстоцена невелика. Особенно это относится к холодным этапам
первой половины среднего плейстоцена – донскому и окскому (=эльстер = англий) оледенениям.
В нескольких случаях выявляется разновременность появления ряда таксонов в Восточной и За-
падной Европе. Так, Arvicola cantianus появилась в Западной Европе раньше, чем в Восточной
Европе. Это явление может быть объяснено как разными темпами эволюции в разных частях Ев-
ропы, так и, возможно, недостаточным количеством сопутствующих датирующих данных (геоло-
гических, геохронологических, палеонтологических), которые впоследствии позволят уточнить
возраст ряда местонахождений.
Проведена корреляция восточно-европейских и западно-европейских фаун мелких млекопитаю-
щих. Построена биостратиграфическая схема и карта среднеплейстоценовых местонахождений
мелких млекопитающих, коррелируемых с МИС 18 – МИС 12, т.е. с первой половиной среднего
плейстоцена по международной шкале (ранним неоплейстоценом по шкале РМСК).
Полученные данные позволяют более корректно обосновать и уточнить геологический возраст
отложений и реконструировать палеогеографическую обстановку рассматриваемых периодов.
Они также являются важной составляющей при создании биостратиграфических схем среднего
плейстоцена и плейстоцена в целом.
The British Palaeolithic provides the first academic synthesis of the entire British Palaeolithic, from the earliest occupation (currently understood to be around 980,000 years ago) to the end of the Ice Age. Landscape and ecology form the canvas for an explicitly interpretative approach aimed at understanding the how different hominin societies addressed the issues of life at the edge of the Pleistocene world.
Voles of the genus Microtus underwent a rapid diversification during the Early and early Middle Pleistocene in Europe and have provided the basis for detailed stratigraphical subdivision during this time period. Pronounced evolutionary changes in their dental morphology include: 1) a progressive increase in complexity of the occlusal surface in the first lower and third upper molar, 2) an increase in the relative length of the anteroconid of the first lower molar, and 3) changes in enamel thickness. Evolutionary stages recognised in Microtus can be correlated with the palaeomagnetic time scale. Microtus presumably first appeared before the Olduvai Subchron > 1.95 Ma BP. Microtus assemblages dating to the Jaramillo Subchron (0.99-1.07 Ma) display considerable morphological variability, suggesting an important phase of speciation took place before this subchron. At least two distinct lineages can be recognised within the Jaramillo Subchron: the Stenocranius and the Pallasiinus lineages. The former is characterised by progressive narrowing of the connection between triangles T5-T6, the latter by progressive narrowing of the connection between triangles T4-T5. Morphological differences between the Microtus samples of the Jaramillo Subchron (from Colle Curti and Korotoyak on the one hand and those of Le Vallonnet, Untermaßfeld and Castagnone on the other) could be explained by the parallel occurrences of primitive members of these two lineages at this time. Morphotypes of Microtus samples older than the Jaramillo Subchron foreshadow the development of these two different lineages. Later on, during the early Brunhes Chron, more evolved species of the Microtus lineage can be recognised, including the main groups of the genus (i.e. Microtus, Pallasiinus, Stenocranius, Terricola, and Iberomys), which are widely considered as subgenera. These faunas can be distinguished from assemblages of the Jaramillo Subchron by the evolutionary level of Microtus, Mimomys savini and Drepanosorex as well as the occurrence of other taxa (e.g., Hypolagus). The detailed regional correlation of the different stages of the evolution of Microtus with the palaeomagnetic time scale requires further data and study. © E. Schweizerbart'sche Verlagsbuchhandlung (Nägele u. Obermiller), 2007.
The genus Macaca (Primates: Cercopithecidae) originated in Africa, dispersed into Europe in the Late Miocene and resided there until the Late Pleistocene. In this contribution, we provide an overview of the evolutionary history of Macaca in Europe, putting it into context with the wider late Miocene, Pliocene and Pleistocene European monkey fossil record (also comprising Mesopithecus, Paradolichopithecus, Dolichopithecus and Theropithecus). The Pliocene and Pleistocene European Macaca fossil material is largely regarded as Macaca sylvanus, the same species as the extant Barbary macaque in North Africa. The M. sylvanus specimens found at West Runton in Norfolk (53°N) during the Middle Pleistocene are among the most northerly euprimates ever discovered. Our simple time-budget model indicates that short winter day lengths would have imposed a significant constraint on activity at such relatively high latitudes, so macaque populations in Britain may have been at the limit of their ecological tolerance. Two basic models using climatic and topographic data for the Last Interglacial and the Last Glacial Maximum alongside Middle and Late Pleistocene fossil distributions indicate that much of Europe may have been suitable habitat for macaques. The models also indicate that areas of southern Europe in the present day have a climate that could support macaque populations. However, M. sylvanus became locally extinct in the Late Pleistocene, possibly at a similar time as the straight-tusked elephant, Palaeoloxodon antiquus, and narrow-nosed rhinoceros, Stephanorhinus hemitoechus. Its extinction may be related to vegetation change or increased predation from Homo, although other factors (such as stochastic factors occurring as a result of small population sizes) cannot be ruled out. Notwithstanding the cause of extinction, the European macaque may thus be a previously overlooked member of the Late Pleistocene faunal turnover.
Usage of the term ‘Cromerian’ differs between countries. In the Netherlands, four interglacial stages (and associated cold stages) are included within the so-called ‘Cromerian Complex’. In Britain, where the Cromerian stratotype is located, all ‘Cromerian’ sites have, until fairly recently, been attributed to different parts of a single interglacial stage. The first suggestion that this view might over-simplify reality was based on vertebrate evidence, in particular the occurrence of Arvicola terrestris cantiana, rather than Mimomys savini, in some deposits. Molluscan assemblages also exhibit significant differences between sites that have been attributed to the same interglacial stage. Valvata goldfussiana appears to characterize fluvial assemblages in the early part of the Cromerian Complex, whereas Valvata naticina typifies several of those that occur later. Combining molluscan and vertebrate evidence, it seems likely that as many as five distinct stages have been conflated and that the stratotype at West Runton occurs early in the Cromerian Complex, rather than immediately preceding the Anglian Stage.
Spotted hyaena (Crocuta crocuta) coprolites from four British Pleistocene sites were analysed for their pollen content. At the two open sites, the palynology was compared to that of the surrounding sediments. The results provide palynological data, supported by other lines of evidence, enabling reconstruction of human environments as well as providing insights into the taphonomic complexities of incorporation of pollen into coprolites. Pollen presence and preservation appear to be closely related to mammalian behaviour and post-depositional processes. Geological age does not seem to be a significant factor, as samples from two of these sites are amongst the earliest known from the Pleistocene to provide viable coprolite-derived pollen counts.
Intraspecific variability of the Pleistocene speciesSorex runtonensis Hinton, 1911 from different Polish and Russian (Caucasus Mts.) localities and its relationships to Recent red-toothed shrews
of similar body size and mandibular morphology,S. caecutiens Laxmann, 1788 andS. tundrensis Merriam, 1990, are explained on the grounds of multivariate statistics. The remains ofS. runtonensis from different localities form a single group and differ fromS. caecutiens. They resembleS. tundrensis by the first canonical root related to a mandibular size and proportions. This may indicate thatS. runtonensis andS. tundrensis are closely related but separated components of the species complextundrensis.
Terrestrial vertebrate faunas (mammals, reptiles and amphibians) from the Middle and Upper Pleistocene of Britain, Ireland and adjacent continental Europe are compared. Problems arise from uncertainties of status and correlation of stages. Evidence of insularity is provided by reduced diversity in (a) Britain versus the Continent, and (b) Ireland versus Britain. British temperate/interglacial faunas older than the Last Interglacial are very similar to continental faunas, notably the Cromerian faunas of West Runton, England, and Voigstedt, Germany. The earliest indications of insularity are in Last Interglacial faunas, in which, for example, pine voles Pitymys spp., extinct rhinoceros Stephanorhinus kurchbergensis, horse Equus ferus and humans, present in equivalent continental faunas, are absent from Britain. -from Author
The record of river terraces and related fluvial deposits from the Vltava and Labe (Elbe) rivers in the Czech Republic is reassessed. Field relations between the Labe terraces and sediments derived from Scandinavian ice sheets, together with biostratigraphy, provide the basis for correlation with the oxygen isotope record from the oceans. The resulting scheme allows an internally consistent correlation of these terraces with their more fragmentary counterparts downstream in Germany. It is also possible to use the terraces to estimate Late Cenozoic uplift in this region, which appears to have followed a similar pattern to other parts of Europe, in particular with a marked increase in uplift rates after ∼0.9 Ma. Previous schemes may, thus, have overestimated the ages of earlier parts of this Middle Pleistocene fluvial sequence in the Bohemian Massif. Analysis of the biostratigraphy reveals many similarities between interglacial faunas from southern England and the Czech Republic and raises questions about some of the previously published correlations.
Recent refinements to the dating of a number of Middle and Late Pleistocene localities and fossil mammal assemblages from the British Isles, and their correlation with Marine Isotope Stages (MIS), permit a more detailed assessment of the evolution of the guild of larger Carnivora. By around 0.4–0.5 Ma (MIS 13–11) or just after, the more archaic elements of the guild, such as the sabretoothed cat Homotherium latidens, the medium-sized pantherine Panthera gombaszoegensis and the large dog Canis lycaonoides, had disappeared and had been replaced, or were being replaced, by extant species such as lion (Panthera leo), spotted hyaena (Crocuta crocuta) and the wolf (Canis lupus), although claims for a last-glaciation appearance – or reappearance – of Homotherium require assessment. During MIS 5a this guild seems to have been temporarily replaced by a more restricted one of wolf in conjunction with a very large brown bear (Ursus arctos), together with a reduced prey fauna of bison and reindeer. Final extinction of Crocuta and lion in these islands took place towards the end of MIS 3, arguably prior to the Last Glacial Maximum. Copyright © 2009 John Wiley & Sons, Ltd.
A section close to the Mammoth (Mammuthus trogontherii Pohlig) remains at West Runton was sampled to allow both plant macrofossil and palynological investigations. The samples analysed yielded diverse and well preserved palaeobotanical assemblages. The plant macrofossil assemblages indicate that weathering probably occurred at the top of the sequence. Assemblages from lower in the profile suggest that sedimentation was either relatively rapid, some mixing of sediments occurred, or that both these factors operated, contributing to the uniformity of the assemblages. Waterside and damp ground taxa together with aquatic taxa are dominant and indicate that fresh-water in the channel was shallow, slightly basic, mesotrophic, and still or slow moving. The presence of taxa characteristic of open, disturbed or bare ground may point to the movement of animals to and from the water’s edge. Palynological data show that further afield, temperate mixed woodland composed of deciduous and coniferous taxa existed. A component of the pollen spectra suggest that the coastline was nearby. A correlation with West’s (1980) Cromerian IIa biozone can be made using the palynological data.
Deer species provide a valuable biostratigraphical tool through Cromer Forest-bed times, due particularly to species turnover between the Early and early Middle Pleistocene. This study is based on the reidentification of 348 fossil antlers. The provenance of most large mammal fossils from the CF-bF, collected over 150 years, was recorded only by the nearest coastal village. None the less, analysis of the cervid taxa by these ‘localities’ reveals interesting patterns. The fauna of the West Runton Freshwater Bed is of early Middle Pleistocene complexion, and that from the foreshore at East Runton is of Early Pleistocene. Pure or nearly pure early Middle Pleistocene assemblages also occur at Kessingland-Pakefield and at Trimingham. At Overstrand, Sidestrand, Mundesley, Bacton and Happisburgh, there is a mixture of Early and early Middle Pleistocene elements. Analysis of Savin's data shows that fossils of earlier species were generally found further down the beach than those of later ones. Late nineteenth century geological surveys, made when bone collecting was at its peak, give additional information about fossil horizons, which in several cases can be related to modern stratigraphical units. Most Early Pleistocene large-mammal bones came from Pastonian conglomerates, in contrast to small-mammal and molluscan assemblages mostly extracted from Pre-Pastonian Crag. However, the diversity of Early Pleistocene CF-bF cervid species in comparison with continental faunas, and their pattern of distribution between sites, suggests they may span more than one chronostratigraphic stage. Early Middle Pleistocene assemblages came from strata now referred to the Cromerian, and the differing proportions of taxa between sites provide limited evidence of time-transgression.
A thorough review of available fossil material of the larger-sized carnivores from the West Runton Freshwater Bed (WRFB) is presented. Both previously-collected and new fossil materials, some from the 1995 mammoth excavation, are taken into consideration, with particular emphasis on taxonomic assignation. Measurement and morphological data on the fossil material are viewed in the light of the evolutionary history of the various lineages, with some interesting results. The ursid remains show a mosaic of both brown and cave bear characteristics, and for the present are classified as Ursus sp. The dog and hyaenid are identified as Canis mosbachensis and Crocuta crocuta, respectively, with possible additional presence of Pachycrocuta (see Appendix). Dental remains of a small cat are assigned to the wildcat lineage but differ from modern Felis silvestris and are tentatively referred to Felis lunensis. There is also evidence for a serval-like cat and an early record of lynx. Within the larger Felidae, fossil evidence of Homotherium latidens and Panthera gombaszoegensis is presented. The large size of the lion, Panthera leo, is confirmed but is considered unexceptional in the context of European Pleistocene lions. The palaeoecology and changing composition of the larger carnivore guild is discussed. The diverse WRFB fauna exemplifies a period of overlap between more archaic and modern assemblages.
The Sand Hole Formation is a Pleistocene seismic–stratigraphic unit of restricted distribution in the Inner Silver Pit, southern North Sea. Pollen and dinoflagellate cyst analyses of this formation, penetrated by British Geological Survey borehole 81/52A and three juxtaposed vibrocores, enables division of the formation into two sequences of contrasting palaeoclimatic affinity. The lower sequence contains poorly-preserved pollen assemblages of low concentration dominated by residual reworked elements, and is coincident with cold, shallow marine dinoflagellate cyst and foraminiferal assemblages. This sequence overlies till and glacimarine sediments of the Anglian/Elsterian Swarte Bank Formation, and is also interpreted as late Anglian/Elsterian in age. The upper sequence contains well-preserved, high concentration, pollen spectra recording the latter half of an interglacial stage with Hoxnian affinities. The strong terrigenous signal preserved in this sequence suggests a subtidal depositional setting within a region of freshwater influence; correlation of the vibrocores with the longer record preserved in borehole 81/52A indicates that interglacial sedimentation rates increase southwards towards a suggested terrigenous source in the Wash area. The first half of the interglacial cycle is missing, indicating that a hiatus is present between the two sequences preserved within the Sand Hole Formation, but the termination of the interglacial is clearly preserved. The spatial and temporal distribution of the sediments and bounding unconformities in the region are consistent in recording the transgressive ravinement surface, highstand systems tract and subsequent regression within an interglacial eustatic cycle. The character, age and elevation of the preserved facies is typical of a glacio-isostatically controlled emergence cycle and lends support to the notion that this temperate stage (oxygen isotope stage 9) immediately followed extensive glaciation during the late Anglian/Elsterian.
The individualistic response of species to climate change is accepted by many although how this process works across several climate oscillations has not been widely considered. A cluster analysis using the Bray–Curtis metric with single linkage to show relative faunal similarity was performed on successively older British mammalian faunas to investigate whether they become progressively different compared to the present day (Holocene). British mammalian faunas from MIS 3, 5, 11, 13 and 17 were compared with the Holocene revealing that the last glaciation (MIS 3) is more different than are any of the interglacials (MIS 5, 11, 13, 17). Furthermore, the interglacials generally become more distinct from the Holocene with age. This difference relates to the fact that interglacial faunas have greater proportions of extinct and extirpated species with increased age. The increase in extirpated taxa in turn relates to faunas becoming more non-analogue with greater age. The increase in extirpated elements with age probably relates to the individualistic response to climate change which appears to be progressing with time. The implications of this progressive process are considered in relation to refugia, extinction and evolution.
The fossil record of vegetation and ungulates places present conditions and trends in a temporal perspective. Ungulate–vegetation interactions during the last 500 000 years were primarily driven by the climatic variation of the glacial–interglacial cycle. There were distinctive faunas associated with each temperate period and a loss of species diversity only in the present interglacial. Climate change and human activities have interacted during the most recent glacial cycle, accelerating extinction rates. This unique course of events has the consequence that no stable, ‘base-line’ conditions can be recognised. A review of the full-glacial ‘mammoth-steppe’ debate suggests that ungulate populations were limited by available forage, but a mosaic of habitat supported a diverse fauna in Beringia. In the debate over early–mid Holocene ‘wood pasture’, past ungulate populations are one of a range of disturbance factors, including burning, that influenced regional vegetation composition and structure in northern Europe. These debates concerning the scale and impacts of past ungulate–vegetation interactions will not be fully resolved until more is known about past ungulate population sizes. Modelling past scenarios would enhance the value of retrospective studies and help provide goals for management of near-natural ecosystems.
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