Allen Et Al. (2009) - Palaeoenvironmetal Context of Woolly Mammoth at Candover, UK

GEOLOGICAL JOURNAL
Geol. J. 44: 414446 (2009)

Published online in Wiley InterScience
(www.interscience.wiley.com) DOI: 10.1002/gj.1161
Palaeoenvironmental context of the Late-glacial woolly mammoth

(Mammuthus primigenius) discoveries at Condover, Shropshire, UK
J. R. M. ALLEN 1,2*, J. D. SCOURSE 2, A. R. HALL 3 and G. R. COOPE 4,5
1
School of Biological and Biomedical Sciences, Durham University, Durham, UK
School of Ocean Sciences, College of Natural Sciences, Bangor University, Anglesey, UK
3
Department of Archaeology, University of York, The Kings Manor, York, UK
4
School of Geography Earth and Environmental Sciences, University of Birmingham, Edgbaston,
Birmingham, UK
5
Tigh-na-cleirich, Foss, by Pitlochry, Perthshire, UK
2
In 1986/1987 the remains of several mammoths, Mammuthus primigenius (Blumenbach), were discovered on the spoil heap of
an actively working gravel pit at Condover, Shropshire, England. The discovery of the remains posed two questions that could be
addressed by analyses of biological proxies. First, as none of the bones was found in situ it was necessary to confirm the stratum
in which the remains occurred. Second, what was the environment in which these animals lived and died? A range of biological
indicators was used to address these questions, including pollen, spore and algal, plant macrofossil, invertebrate, anuran and
biological mineral analyses. Multivariate statistical analyses of palynological and Pediastrum data, along with evidence from the
Coleopteran assemblages, support the attribution of the mammoth bones to a unit of dark grey clayey sandy silt, although they
may have lived at the time of the overlying green detritus mud. The palaeobiological data supports the correlation of these
sediments to the Devensian Late-glacial. The mammoths entered this basin at the start of the Late-glacial Interstadial (Greenland
Interstadial 1e) (ca. 14 8303930 cal. year BP; 12 300 110 14C year BP) and became mired in soft cohesive sediments.
Palaeotemperature reconstructions, based on the Coleopteran assemblages, from the time when the mammoths actually became
mired, show that the climate was temperate with mean July temperatures between 15 and 198C and mean January temperatures
between 13 and 68C. Biological indicators from the sediments encasing the mammoths indicate that the landscape
surrounding the basin was treeless and dry, contrasting with rich vegetation within the basin itself that had possibly attracted the
mammoths to the site. Evidence of sedimentary disturbance suggests that the mammoths caused large-scale bioturbation of the
deposits making palaeoenvironmental interpretations difficult. Fossils of terrestrial blowflies, carcass and dung beetles show that
some of the decaying corpses must have lain exposed on the land surface for sufficient time for the soft parts to have rotted away
and skin and bones to have become desiccated before many of them sank into the dark grey clayey sandy silt. Copyright # 2009
John Wiley & Sons, Ltd.
Received 12 February 2009; accepted 23 April 2009
KEY WORDS Devensian Late-glacial; palaeoclimate; pollen; Coleoptera; plant macrofossils; kettle-hole; Pediastrum
Supporting information may be found in the online version of this article (Supplementary Tables S1S10).
1. INTRODUCTION
In 1986 a spectacular set of remains of several Mammuthus primigenius (Blumenbach) (woolly mammoth) were
found at Norton Farm Pit, Condover, Shropshire, UK in the spoil heaps of overburden removed during quarrying
operations prior to extraction of the gravel (Coope and Lister 1987; Lister 2009). This sediment was derived from a
* Correspondence to: J. R. M. Allen, School of Biological and Biomedical Sciences, DurhamUniversity, South Road, Durham DH1 3LE, UK.
E-mail: j.r.m.allen@durham.ac.uk
Copyright # 2009 John Wiley & Sons, Ltd.
palaeoenvironmental context of condover mammoth site, uk
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largely infilled kettle-hole (Scourse et al. 2009). Over the next few years (19861990) the site was the subject of a
series of investigations (Lister 1993). The regional setting of the site (Latitude 528380 1500 N; Longitude 28450 4600 W;
UK NGR SJ494073, 80 m above sea level) has been described by Worsley (2005). Detailed investigations of the
mammoth bones (Lister 2009), which came from an adult male and several juvenile animals, and their local
stratigraphic context (Scourse et al. 2009) have also been undertaken. Biological investigations of the sediments
including pollen, spore and algal, plant macrofossil, insect and other invertebrate, anuran (frog and toad) and fish, as
well as biological minerals, are reported here.
Samples for biological analyses were taken on a number of occasions between 1986 and 1990, and from a
number of sedimentary units, with two main objectives. First, as none of the mammoth bones was discovered
in situ, a prime aim of the investigations was to identify as accurately as possible the sedimentary sequence infilling
the kettle-hole basin, so that sediment adhering to the bones could be compared with the in situ sedimentary units
(Scourse et al. 2009) and the source of the bones identified. The second aim was to provide evidence on the nature
of the changing environment in and around the Norton Farm Pit site before, during, and after the period when the
mammoths entered the basin.
A number of techniques can be used to provenance unstratified sediment such as that adhering to the mammoth
bones. This includes radiocarbon dating (i.e. by matching the radiocarbon-based age of the bones and the sediment
adhering to them to the radiocarbon-based age of the sediments (Scourse et al. 2009)), mineralogy, geochemistry
and grainsize analysis. Because the amount of sediment available for analysis is often very small, microfossils such
as pollen (with accessory spores and algae) are one of the most useful approaches and can provide evidence of the
regional vegetation of the locality, thus enabling broad stratigraphical correlations to be made. These can be
supplemented by more local information gained from macrofossil remains of plants, principally fruits and seeds,
but also leaves and wood. Evidence from faunal remains such as Coleoptera, Trichoptera, Chironomidae and
Anurans can also make unique contributions to the palaeoenvironmental interpretation.
2. MATERIALS AND METHODS

2.1. Pollen and Pediastrum
2.1.1. Field sampling
The samples used for pollen and Pediastrum analyses are shown in Supplementary Table S1 and the locations of the
various profiles from which the samples were taken are shown on the map of the site (Figure 1).
In 1987 Profile C (Figure 1; Face 1) was chosen as the reference section for the entire sequence infilling the
kettle-hole basin. This was because
(i) many of the identified sedimentary units (Scourse et al. 2009) could be observed in stratigraphic section in this
single profile and were accessible for sampling; and
(ii) this was where the stratigraphic unit hypothesized, during the JuneJuly 1987 sampling season, to be the
mammoth stratum (Unit C1), was thickest and most easily related to the other sedimentary units.
In order to maximize the sampled thicknesses of Units C1, D, E and F the sample sequence was divided into two
parts (Figure 2); a total of 27 samples was taken through Units C1, D, E and F at Profile C.
In addition to the samples from Profile C, a single sample of the overlying green detritus mud (Unit C3) was
collected in 1987 from Profile D (Figure 1; Face 1).
To relate the unstratified mammoth bones to the reference profile and to the other samples, three sediment
samples scraped from mammoth bones newly recovered from below the water level of the gravel pit during the
JuneJuly 1987 excavation were also prepared for pollen analysis. One of the samples was scraped from a juvenile
vertebra discovered close to the site of Borehole g (Figure 1); the other two were scraped from the juvenile jaw bones.
In January 1988, the water level in the pit was lowered by pumping and this allowed further sampling of Unit C1,
this time at Profile Z, closer to the location from where the bones had been removed during quarrying activities, to
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Figure 1. Site map showing location of Profiles at Norton Farm Pit 19861990.
the southsouthwest of Profiles C and D (Figure 1; Face 2). A vertical series of ten samples was taken through Unit
C1 at Profile Z; of these, only three, all situated towards the base of Unit C1, proved to contain palynomorph
concentrations in excess of 4000 grains g1 wet weight and were thus suitable for analysis. In addition, two samples
were taken from Unit C1 at Profile Y in Face 2, further to the north and closer to Face 1 than Profile Z (Figure 1).
During May 1990 a particularly thick sequence of Units E and F was exposed by quarrying activities in the
vicinity of Profile B (Figure 1). In order to confirm a Holocene age for the black humified peat of Unit F a further
series of 28 samples was taken; in this exposure Unit F was 2.0 m thick, Unit E 70 cm thick; the contact between
Unit E and Unit D also was covered in the sampling.
2.1.2. Laboratory methods
Samples for pollen analyses were initially prepared using standard treatment, including hydrofluoric (HF) acid
(Faegri and Iversen 1989), but the results were extremely disappointing. The pollen concentrations were low, and
the grains very badly corroded. An alternative method was subsequently used replacing HF digestion with a
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Figure 2. Profile C showing relative locations of the pollen sampling sequences. Unit Fblack humified peat; Unit Egrey clay; Unit D
brown sedge peat; Unit C1dark grey clayey sandy silt; Unit B2clast-supported gravel; Unit B1pink laminated clays and silts.
physical method for the removal of inorganic material. After disaggregation samples were washed over a 10 mm
monofilament nylon mesh (Cwynar et al. 1979; Lowe and Walker 1986) prior to heavy liquid separation using
saturated zinc chloride (ZnCl2: density 2.0 g cm3; (Bjorck et al. 1978; Watkins et al. 2007)). These preparations
were far superior, in terms of both the concentration and preservation of the palynomorphs, as has previously been
noted for highly minerogenic samples at other sites (Scourse et al. 1992). The addition of Lycopodium (clubmoss)
spore tablets (Benninghoff 1962; Matthews 1969; Bonny 1972) to weighed subsamples of the sediment prior to
pollen extraction enabled the calculation of pollen concentrations. The pollen extracts were mounted in silicone oil
and examined using a compound light microscope at a routine magnification of 400 with 1000 (oil immersion)
magnification being used for the examination of critical features. For each level at least 300 identifiable land pollen
and spores were counted. Unidentifiable pollen was recorded as crumpled, broken, corroded, degraded
(amorphous) or concealed (Cushing 1967). Plant names follow Clapham et al. (1962), and the pollen and spore
types follow the schemes of Faegri and Iversen (1989) and Birks (1993); the conventions used to indicate the
certainty of identification follow Birks (1993).
The use of density separation rather than HF also enhances the recovery of Pediastrum (class: Chlorophyceae,
order Chlorococcales). The colonies of this alga form from aggregating zoospores and the species differ in the
number and form of prongs on the peripheral cells, the amount of space between the cells, and the wall sculpture
(Millington et al. 1981). Pediastrum colonies were particularly rich in these samples and it was clear during routine
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pollen counting that a number of very distinct forms could be identified. The preservation was generally good,
although in some cases the wall sculpturing and the prongs were either eroded or broken, and there were also some
broken colonies of irregular shape. The identification of Pediastrum taxa found in these sediments follows
Jankovska and Komarek (1982) and Pascher (1913). The species/varieties identified include Pediastrum boryanum
typicum, P. boryanum longicorne, P. boryanum brevicorne, P. duplex, P. kawraiskyi, P. sturmii and P. tetras. An
indeterminable group included all broken and degraded colonies. Some of the P. boryanum brevicorne may include
small numbers of P. integrum. The frequencies are expressed as percentages of the total number of Pediastrum
counted. As different Pediastrum species have different responses to nutrients they can be used as indicators of the
trophic status of the water body. In general Pediastrum numbers increase as nutrients increase and changes in
species composition occur in connection with eutrophication or oligotrophication (Cronberg 1982).
The pollen diagram for Profile C was zoned using ZONATION (Gordon and Birks 1972) using 14 taxa (Betula
(excluding B. nana), Salix (excluding S. herbacea), Betula nana, Salix herbacea, Juniperus communis, Gramineae,
Cyperaceae, Solidago type (Compositae), Artemisia, Filipendula, Myriophyllum, Nymphaea, Potamogeton and
Pediastrum (total)). This Fortran IV package combines three zonation procedures, CONSLINK
(dendrogram for constrained single link analysis), SPLINF (dendrogram for binary division using an
information content criterion) and SPLITSQ (dendrogram for binary division using sum-of-squares
criterion). These procedures group similar samples together, and split or separate samples with few
similarities. The three levels within Unit C1 at the base of the upper part of the diagram were excluded from
this zonation procedure because they are possibly duplicated by samples from the lower part; a total of 24
levels was therefore used.
2.2. Plant macrofossils

The samples used for macrofossil analyses are shown in Supplementary Table S2.
Subsamples of raw sediment were broken down in water in a bucket and sieved (300 mm mesh) in a stream of
water to remove the fine sand, silt, clay and unidentifiable organic debris. In deposits such as these, where the
content of plant remains is generally quite small, it is most practicable to concentrate the organic fraction by means
of a washover, in which the disaggregated sediment is swirled in water in the bucket and the less dense material
decanted off before it settles out. The sieved plant remains were sorted under a binocular microscope and put aside
for identification, which relies on modern reference material and published accounts of modern and fossil
specimens such as Beijerinck (1947) and Katz et al. (1965). For this site a four-point scale of abundance was
adopted, from 1 (one or a few individuals or fragments) to 4 (an abundant component of the sample).
2.3. Insects
The samples used for insect analyses are shown in Supplementary Table S3.
The samples of sediment collected at the site were placed directly into polythene bags and kept in their fielddamp conditions until ready for laboratory processing. Standard techniques were adopted to recover insect fossils
(e.g. Coope 1968; Elias 1994). The sample was disaggregated by applying a gentle stream of water; the resultant
mixture was washed over a 300 mm aperture sieve; this is sufficiently fine to retain most insect remains. The
>300 mm fraction was washed with water and mixed with paraffin oil. Water was then added to permit a clear
separation between the two fractions. Floating material was decanted into a 300 mm sieve, washed in dilute
detergent and water to remove the oil and then in alcohol to remove the water. The insect fossils were sorted under
alcohol using a binocular microscope and stored in tubes of alcohol to prevent attack by fungi.
Identification of the fossil insect fragments was by direct comparison with specimens of modern animals. No
significant differences were found between the fossils and the modern material. By far the most common of the
recognizable fossils were of Coleoptera. Although remains of Diptera, Hymenoptera and other orders of insect were
also present, only the Trichoptera and Chironomidae have been studied in detail.
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2.4. Other environmental indicators

Anura, Trichoptera, Chironomidae, Ostracoda and biological mineralogical analyses all provided additional
evidence relating to the environment around the time of the demise of the mammoths; four brief reports are
included in the results section.
3. RESULTS
3.1. Pollen and Pediastrum
The pollen diagram for Profile C is shown in Figure 3. Sample depths are expressed as depths below the top of the
profile. The results of the ZONATION analysis applied to Profile C are shown in Figure 4; this supports the adoption
of pollen zones which coincide with the major lithostratigraphic boundaries. Zone NF1 coincides with sedimentary
Unit C1, NF2 with Unit D, NF3 with Unit E and NF4 with the base of Unit F (Figure 2). The Profile C pollen
diagram is divided into two sections, the separation being marked by the non-continuity of the pollen curves. The
sections in which the curves are joined represent continuous stratigraphic sequences. The lower part of the diagram
relates exclusively to Unit C1, whereas the upper part covers the stratigraphic sequence from the top of Unit C1
through Units D (brown sedge peat) and E (grey clay) to the base of F (black humified peat) (Figure 2). It is probable
that the upper part of the lower sequence overlaps the three samples of Unit C1 at the base of the upper sequence.
Samples taken from the very base of the upper sequence (Figure 3), and probably from Unit B1 (pink laminated
clays and silts), proved to be devoid of pollen and are not included on the pollen diagram.
3.1.1. Profile C: pollen assemblage zone NF1 (Unit C1): GramineaeCyperaceaeArtemisia
This pollen assemblage zone (paz) consists of ten samples taken from Unit C1 in Profile C (seven from the lower
section and three from the upper section). Pollen concentrations fluctuate widely from just under 6000 grains g1
wet weight in the basal level to over 109 000 grains g1 in the highest level; this increase is masked by saw-tooth
variations in the intervening levels.
The dominant taxa in this zone are Gramineae, Cyperaceae, and to a lesser extent, Artemisia. In the lower
part of the diagram Gramineae rises from just under 2% in the basal level to ca. 69% in the highest level against
which Cyperaceae displays a reciprocal relationship, falling from ca. 87% in the basal level to a little over 8%
in the highest. Artemisia uctuates between 1 and 11%, and occurs alongside a diverse range of herbs which
are all represented at low values (<2%). Paz NF1 also has a low but continuous curve for undifferentiated
Salix which rises to just over 8% at the top of the zone. High levels of reworked palynomorphs are indicated
by the pre-Pleistocene, dinoagellate cyst and indeterminable components of the spectra. Accordingly, the
sporadic counts of poorly preserved thermophilous tree taxa, such as Alnus, Carpinus, Quercus, Tilia and
Fraxinus, could possibly be attributed to reworking from the underlying glacigenic sediments. The
continuous but low values for Pinus can probably be attributed to long-distance aerial transport which is also
a more likely alternative explanation for the occurrence of some of the thermophilous tree taxa (cf. Cundill
and Whittington 1983).
Pediastrum colonies are very abundant in paz NF1, exceeding 50% of the pollen sum plus Pediastrum
(SP Pediastrum) in the highest level of the lower sequence (Figure 3). These are largely represented by
P. boryanum typicum (5085% of the Pediastrum sum) with subdominant P. boryanum longicorne (up to 20%) (Figure 5).
3.1.2. Profile C: pollen assemblage zone NF2 (Unit D): BetulaSalixJuniperus
Pollen concentrations for the nine samples in paz NF2 are generally higher, and more consistent, than in NF1
(38 000186 000 grains g1), as might be expected when comparing a highly organic sedge peat (Unit D) with a
largely minerogenic clayey sandy silt (Unit C1).
Paz NF2 is characterized by a quite different pollen assemblage from NF1. Declines in Gramineae, Cyperaceae
and Artemisia are accompanied by increases in Betula (both tree birch and B. nana (Berglund and Digerfeldt 1970;
Birks 1968; Birks 1973)), Salix (both shrub species and S. herbacea) and Juniperus communis. The Betula undiff.
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Figure 3. Pollen percentage diagram for Profile C. Taxa that never exceed 1% are included as other tree/shrub/herbaceous. Taxa between 1 and 2.5% are shown as dots. Pollen sum
(SP) TLP (Total Land Pollen); C Cryptogams; A Aquatics; I indeterminables; Ped Pediastrum; PP Pre-Pleistocene spores.
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Figure 4. Profile CResults of Zonation.
curve rises to a peak of over 46% in the centre of the zone; Salix undiff. ranges between 5 and 12% and Juniperus
between 1 and 11%. Juniperus and Salix cf. S. herbacea, which has a peak of over 9% in the basal level, decline
from maxima early in the zone as Betula undiff. (mostly tree birch) increases. As the latter taxon declines towards
the end of the zone so Betula cf. B. nana increases, reaching a peak of almost 18% towards the top. As these changes
occur within a highly organic sedge peat of telmatic character they probably reflect actual changes in the local
vegetation rather than any taphonomic artifact of sedimentation or preservation.
Figure 5. Pediastrum percentage diagram for Profile C.
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Accompanying these changes in the tree and shrub vegetation are changes in the herbaceous community. Other
than grasses and sedges, the most consistent herb taxa in paz NF1 are Artemisia, Compositae subfamily
Liguliflorae, Arenaria type, Caryophyllaceae undiff., Chenopodiaceae and Trifolium. These are replaced in paz
NF2 by Rosaceae undiff., Helianthemum, Liliaceae, Ranunculus repens type, Rubiaceae and Angelica type.
Changes also occur in the aquatic taxa. Pediastrum declines as macrophyte aquatic vegetation becomes
important. Pediastrum fluctuates between 0 and 4% in the lower part of the zone, but rises to 35% towards the top.
As in paz NF1 P. boryanum typicum dominates the Pediastrum assemblages, but P. duplex, which was entirely
absent in NF1, rises to exceed 40% of the Pediastrum spectra in the centre of the zone. Macrophyte aquatic taxa in
NF2 include Myriophyllum spp., predominantly M. spicatum (almost 12% at the base of the zone), but also
M. verticillatum and M. alterniflorum. Peaks of Potamogeton occur at the base (almost 2%) and top (26%) of the
zone, framing a continuous record of Nymphaea which reaches a peak of over 8% in the centre of the zone. Other
obligate aquatic taxa include Nuphar, Menyanthes trifoliata and Typha latifolia. NF2 is also rich in the usually
helophytic pteridophyte Equisetum which reaches almost 15% in the central part of the zone.
The low totals for indeterminate pollen, and the complete absence of dinoflagellate cysts and pre-Pleistocene
taxa, suggest minimal reworking in this zone.
3.1.3. Profile C: pollen assemblage zone NF3 (Unit E): BetulaGramineaeCyperaceaeArtemisia
Paz NF3 consists of seven samples taken from Unit E (grey clay) in Profile C. Pollen concentrations show a decline
from the high levels attained in NF2, rising from a minima of under 3000 grains g1 wet weight at the base of the zone
to over 70 000 grains g1 at the top. Reworking resumes in NF3 with increases in the occurrence of pre-Pleistocene and
indeterminable taxa. Though Betula undiff., B. cf. nana, Salix undiff. and S. herbacea decline in comparison with NF2,
they nevertheless maintain a presence throughout NF3. Apart from two isolated grains, Juniperus is absent in NF3.
Gramineae and Cyperaceae increase in frequency, Gramineae ranging between 41 and 20%, and Cyperaceae between
23 and 16%. Artemisia is well represented, rising to over 20% in the central part of the zone. The helophytic herb taxon
Ranunculus trichophyllus type rises to a peak of over 25% at the top of the zone. Other significant herb taxa include
Rumex acetosa type, for which there is a continuous curve, Cirsium/Carduus, Compositae subfamily Liguliflorae,
Arenaria type, Caryophyllaceae undiff., Potentilla type and Thalictrum.
The macrophyte aquatic taxa present in NF2 are almost completely absent in NF3, with the exceptions of
sporadic occurrences of Menyanthes trifoliata and Typha latifolia which may be re-worked from NF2. Pediastrum
increases in NF3 to reach a peak of over 71% in the top level, its highest value in the entire diagram. Once again
P. boryanum typicum dominates (up to 50% of the Pediastrum assemblage), P. boryanum longicorne reaches 25%
and there is a diverse range of other taxa, including P. boryanum brevicorne, P. duplex, P. kawraiskyi, P. sturmii and
P. tetras.
3.1.4. Profile C: pollen assemblage zone NF4 (base of Unit F): BetulaSalixJuniperusGramineae
Paz NF4 is represented by the single level from Unit F (black humified peat) at the top of Profile C (Figure 3). This
level has a pollen concentration of over 68 000 grains g1 wet weight and shows some significant changes from
zone NF3 below. These include an increase in Betula undiff. to almost 22%, in Salix undiff. to almost 17%, in
Salix cf. herbacea to almost 6%, and reductions in Artemisia, and Ranunculus trichophyllus type. Juniperus rises to
over 1%, and Equisetum reaches over 6%. Pediastrum declines from over 71% in the top level of NF3 to just over
1% in NF4.
3.1.5. Palynology of Unit C3 (green detritus mud)
Unit C3 (green detritus mud), which did not occur at Profile C, lies stratigraphically between Unit C1 (pollen
assemblage zone NF1) and Unit D (pollen assemblage zone NF2 (Scourse et al. 2009). The sample of Unit C3,
collected in 1987 from Profile D (Face Y; Figure 1) yielded a pollen spectrum dominated by Cyperaceae (43.7%),
Gramineae (37%) and Ranunculus repens type (7.7%), with traces of Betula, Artemisia, Armeria/Limonium (type
A), and Thalictrum (Figure 6). The pollen concentration exceeds 35 000 grains g1 wet weight. Pediastrum,
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Figure 6. Pollen percentage diagram for bone scraping and other samples from Units C1 and C3. Taxa that never exceed 1% are included as other tree/shrub/herbaceous. Taxa between
1 and 2.5% are shown as solid circles. Pollen sum (SP) TLP (Total Land Pollen); C Cryptogams; A Aquatics; I indeterminables; Ped Pediastrum; PP Pre-Pleistocene spores.
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dominated by P. boryanum typicum with subdominant P. boryanum brevicorne and P. boryanum longicorne,
reaches almost 49% (Figures 3 and 5).
3.1.6. Palynology of sediment scraped from mammoth bones
Three sediment samples scraped from juvenile mammoth bones were analysed. The sample from a vertebra
discovered close to the site of Borehole g (Figure 1) is the uppermost sample in Figures 6 and 7. The other two
samples were from scrapes of jaw bones. The two jaw samples are very similar to each other, with pollen
concentrations of 8000 grains g1 wet weight and almost 29 000 grains g1. The pollen assemblages (Figure 6) are
dominated by Cyperaceae (3844%), with subdominant Gramineae (1930%), Pinus (67%), Salix (45%) and
Trifolium cf. T. repens (610%). The assemblage in the vertebra sample (pollen concentration, 19 000 grains g1
wet weight) shows some differences. Cyperaceae is even more dominant (almost 80%) and Gramineae less so
(under 10%), with Pinus (5%) and Salix (2.5%); Trifolium cf. T. repens is absent. In all three samples levels of prePleistocene and indeterminable palynomorphs are high, and reworked dinoflagellate cysts occur in the vertebra
sample. Pediastrum totals are also high in all three samples (4175%; SP Pediastrum), dominated by
P. boryanum typicum with subdominant P. boryanum longicorne and P. sturmii (Figure 7).
3.1.7. Palynology of Unit C1 (dark grey clayey, sandy silt) from Profiles Z and Y, Face 2
The three samples from Unit C1 at Profile Z yielded assemblages extremely similar to zone NF1 from Profile C,
also from Unit C1. Cyperaceae dominates the spectra (up to 84%), with subdominant Gramineae (up to 31%) and
Pinus (up to almost 7%). As in NF1 there are isolated occurrences of thermophilous tree and shrub taxa, including
Alnus and Corylus/Myrica, and there are significant numbers of pre-Pleistocene, indeterminable and dinoflagellate
taxa. The pollen concentrations are low, ranging between 6000 and 13 000 grains g1 wet weight. Pediastrum
Figure 7. Pediastrum percentage diagram for bone scraping and other samples from Units C1 and C3.
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concentrations rise from less than 2% in the basal sample to over 66% (SP Pediastrum) in the top sample,
dominated by P. boryanum typicum with P. boryanum brevicorne (Figure 7).
A further two samples were taken in 1988 from Unit C1 at Profile Y in Face 2, further to the north and closer to
Face 1 than Profile Z (Figure 1). The pollen concentrations and assemblages here proved to be extremely similar to
both the samples from Profile Z (Figure 6) and zone NF1 from Profile C (Figure 3). The lower sample has a pollen
concentration of nearly 9000 grains g1 wet weight, rising to almost 23 000 in the upper sample. The pollen
assemblages in these two samples are dominated by Cyperaceae (5174%), with subdominant Gramineae (15
34%) and Salix (24%). High levels of reworking are indicated by significant quantities of pre-Pleistocene and
indeterminable taxa. Pediastrum rises from just over 11% (SP Pediastrum) in the lower sample to almost 65% in
the upper; these consist of dominant P. boryanum typicum with P. boryanum longicorne and P. boryanum
brevicorne (Figure 7).
3.1.8. Profile B pollen diagram
The base of this pollen diagram (Figure 8) overlaps with the top of the diagram from Profile C (Figure 3), and the
upper part, by comparison with other nearby well-dated pollen diagrams (e.g. Beales 1980), covers much of the
early Holocene. Important features in this respect include the immigration of Corylus, and later, Alnus. This
sequence is, however, difficult to interpret in detail because it is dominated by pollen taxa from very local sources;
many of the observed changes are therefore probably caused by small changes in the hydrology/water level of the
basin, as might be expected in a telmatic peat. The large fluctuations in the Cyperaceae curve, and the irregularity of
the curves for the major tree taxa, can probably be explained in this way. However, these tree taxa are very poorly
represented. In particular, the very low frequencies for the mixed woodland taxa Corylus, Ulmus and Quercus are
puzzling for most other pollen records covering the early Holocene from this area (e.g. Beales 1980) record high
frequencies for these taxa. Even accounting for the local pollen catchment, the record appears to indicate minimal
mixed woodland development in the early to mid-Holocene.
The diagram can be subdivided into five pollen assemblage zones (Figure 8). Zones NF2 and NF3 correlate with
these same zones in Profile C (Figure 3), and correspond to Units D and E respectively. Zone NF4, the base of Unit
F, correlates with the uppermost sample from Profile C (Figure 3). Zone NF5 is defined by the rise in Corylus/
Myrica, but is dominated by Betula, Salix and Cyperaceae, with subdominant Pinus and Gramineae. Zone NF6 is
defined by the rise in Alnus and is represented by only one level at the top of the profile.
3.1.9. Multivariate statistical analysis of the pollen spectra
In order to assess objectively the similarity between the pollen spectra from the bone scrapes and the spectra from
the stratigraphic samples the data were analysed using detrended correspondence analysis (DECORANA; Hill
1979). The same fourteen pollen taxa as used for the zoning of the main pollen diagram were used in the analysis.
The results are shown in Figure 9. The plot shows that the analysis separates the four pollen zones identified from
Profile C and that the pollen spectra from the bone scrapes are more similar to those from paz NF1 (Unit C1) than
any of the other zones. This supports the working hypothesis generated in the field, and supported by the
radiocarbon dating (Scourse et al. 2009), that most of the bones came from Unit C1. However one of the jaw bones
might derive from Unit C3 (Figure 9).
The elements which characterize and unify the pollen assemblages from Unit C1 include high Cyperaceae and
Gramineae with subdominant Pinus, Artemisia and Trifolium cf. T. repens. Indeterminate, pre-Pleistocene and
dinoflagellate levels are high, as are frequencies of Pediastrum; P. duplex is either entirely absent or present in
extremely low concentrations. Overall, pollen concentrations are low, and in some cases so low as to make the
samples effectively barren.
Comparison of the changes in the assemblages from NF1, Profile C (Figure 3) with the changes within Unit C1
from Profiles Z and Y (Figure 6) shows some common trends. Gramineae rises from the base to reach maximum
levels towards the top, whereas Cyperaceae shows a reciprocal pattern of decline upwards. Moreover, this is
matched by an increase in the total Pediastrum concentrations upwards. These detailed changes enable the bone
scrape samples to be more accurately placed within Unit C1. The juvenile vertebra sample (Figure 6) is rich in
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Figure 8. Pollen percentage diagram for Profile B. Taxa that never exceed 1% are included as other tree/shrub/herbaceous. Taxa between 1 and 2.5% are shown as solid circles. Pollen sum
(SP) TLP (Total Land Pollen); C Cryptogams; A Aquatics; I indeterminables; Ped Pediastrum.
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Figure 9. DECORANA Factor 1 and Factor 2. Samples are 124 the samples from Profile C ((117 are from the upper section (sample 1 from
Unit F, 28 from Unit E and 917 from Unit D) and 1824 from the lower section Unit C1), sample 25 is from the juvenile jaw scrape, samples 26
and 27 from vertebra scrapes, 28 is from Unit C3, samples 2933 are from Unit C1 samples in Profiles Y and Z.
Cyperaceae, relatively low in Gramineae with moderate Pediastrum concentrations, whereas the juvenile jaw
samples are less rich in Cyperaceae, but with more Gramineae and moderate to high levels of Pediastrum. This
suggests that the juvenile vertebra bone derives from a lower stratigraphic level within the sequence than either of
the jaw bones. This conclusion is supported by the correspondence analysis (Figure 9), one of the jaw bones
probably deriving from Unit C3.
The correspondence analysis demonstrates similarity of pollen assemblages in zones NF3 and NF1. Zone NF3
can be clearly separated, however, on the basis of the presence of Betula (tree and B. nana), increased significance
of Salix (including S. herbacea) and Artemisia, Ranunculus trichophyllus type and Rumex acetosa type.
3.2. Plant macrofossils
Nomenclature and taxonomic order for vascular plants follow Tutin et al. (19641990) and those for mosses follow
Smith (1978). Plant macrofossil abundance scores are shown in Supplementary Tables S4 and S5. Together with
identifiable plant remains, a few other components of the samples were noted, and these are also presented in the tables.
The assemblages from samples E, F, G and 58 (Supplementary Table S4) have some taxa in common, but they
cannot be compared too closely since sample size was so variable (samples E and 58 were large; samples F and G
were small). Sample F is noticeably richer than the others, and contains abundant remains of birch (Betula),
suggesting it has more in common with Unit D than C1. However, the possible origin of at least some of the bones
from Unit C3 (green detritus mud) on the basis of pollen spectra (see above), suggests that this particular sample
may have consisted of Unit C3 rather than C1. This might explain its somewhat richer assemblage, as C3 was much
more organic than C1; the stratigraphic position of C3 and its associated AMS 14C date (Scourse et al. 2009) would
be consistent with the start of the Betula rise (Figure 3). The sample from the top of C1 prepared for AMS
14
C dating (Scourse et al. 2009) was, however, also rich in Betula macrofossils.
There is greater consistency between one sample and the next in the series from Profile Z (Face 2)
(Supplementary Table S5), with a gradual decline in the overall diversity of taxa upwards through this sequence,
corresponding to an increase in the sand content and a decrease in the organic component of the samples
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represented by samples B8B1. Between the series from Profile Z, and the sample from Profile C and the bone
scrapes, there are some striking differences; the samples from Profile Z are much less rich in taxa, lacking most of
the aquatics (but the one pondweed, Potamogeton rutilus, found at Profile Z is not recorded from Profile C).
3.3. Coleoptera
Coleoptera lists are shown in Supplementary Table S6 (samples AG) and Supplementary Table S7 (samples from
Profile Z). The nomenclature and taxonomic order follows Lucht (1987), with species that do not occur today in
central Europe being inserted at the most suitable taxonomic level. The numbers indicate the minimum number of
individuals recovered from each sample; 134 taxa of Coleoptera were identified, of which 94 have been determined
to the level of species or species group. Eleven of the latter are now extinct in the British Isles. These exotic species
are indicated by in the Tables.
The sequence of samples shown in Supplementary Table S6 is subdivided into four sections.
(1) Samples A, B, D and E represent the dark grey clayey sandy silt (Unit C1), either as the direct matrix of the
bones or associated with them on the spoil heap.
(2) Sample C was collected in situ from the floor of the pit and well separated from the bone-bearing locality, but
included almost all the species found in the rest of this sub-group including many of the exotica. Sample C is
correlated with Unit C1.
(3) Sample F was obtained from the sinus cavities and lower jaw dental alveoli and is clearly different from the rest
of the samples.
(4) Sample G was taken from the green detritus mud (Unit C3) that overlay the dark grey clayey sandy silt (Unit
C1) and contains fauna so different from that of Unit C1 that it is here considered on its own.
Supplementary Table S7 shows the beetle fauna from the sequence of samples from Profile Z Face 2. It is dealt
with separately because of its stratigraphical integrity and because this series of samples shows progressive change
in the local environment.
3.3.1. Samples A, B, D and E
These four samples were of the dark grey clayey sandy silt (Unit C1) adhering to the mammoth bones
(Supplementary Table S3). A list of the taxa recorded is given in Supplementary Table S6 and the environmental
preferences of these taxa are outlined below.
3.3.1.1. Local environments indicated by the Coleopteran assemblage. Several families of Coleoptera provide
information about the aquatic environment. These include Haliplidae, Dytiscidae, Gyrinidae, Hydraenidae,
Hydrophilidae, Dryopidae and some of the Chrysomelidae and Curculonidae.
Three species of Dryopidae are characteristic of rapidly flowing, well-aerated water. They occurred only in
sample E. Most of the other water beetles live in slowly moving or stationary water. Both larvae and adults of
Haliplus are relatively poor swimmers and so live only in standing or very slowly moving water. In contrast, the
dytiscids are powerful swimming carnivores that can live in both flowing and standing water. Potamonectes
griseostriatus inhabits pools of clear water over a sandy or stony bottom, not extensively overgrown by vegetation.
It is a species of cold water, and in Britain it tends to be found in peaty pools at high altitude: in northern European
latitudes it is found at lower elevations. Potamonectes depressus is both a flowing and still-water species. At
Condover, it was represented by a very pale, largely yellow form that is often found in ponds with a clear sandy
bottom. Agabus bipustulatus is found in pools and puddles and is not a stream species, except where slowly moving
or almost stationary habitats occur. Agabus arcticus is a cold-water species. Agabus congener is here a
representative of a species complex of largely northern forms that includes A. lapponicus, A. thomsoni and others
from the eastern palaearctic region. This is a species group that prefers cold-water ponds that are not overgrown by
aquatic plants. Rhantus notatus shows a strong preference for ponds with a silty bottom with some aquatic
vegetation (Balfour-Brown 1950, p.235). Colymbetes dolabratus is adapted to very cold water and is now totally
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absent from the British Isles; its nearest occurrences today are in the alpine and subalpine zones of the Norwegian
mountains where it lives in small ponds and peaty pools. Dytiscus circumcinctus and Acilius, in this case probably
A. sulcatus, are both pond species and, being rather large beetles, they tend to inhabit rather larger bodies of water
than most of the other dytiscid species in this fossil assemblage.
Gyrinus aeratus and G. marinus are difficult to separate and it is possible that both species occurred at Condover.
The hydraenid species are almost all pond or puddle species. Most of the species here occur in grassy ponds with the
exception of Helophorus obscurellus which is one of the least aquatic species of its genus. Although little is known
about its precise ecological requirements, it is normally found under stones and vegetation in sandy places in the
tundra of northern Asia and on the high cold steppes of the mountains further south as far as the Tibetan plateau
(Angus 1992; p.29). By reference to its nearest relatives (e.g. H. nubilus), it is highly likely that its larvae are
phytophagous, feeding on grass shoots. H. sibiricus is a northern species found today from Siberia to Alaska, in
northern Canada and Fennoscandia, and in mountains further south in Scandinavia. It is found in grassy pools and at
the edges of streams and rivers (Angus 1992; p.41). Helophorus splendidus is one of the most cold-adapted species
of this genus. It is known from the tundra across arctic Siberia to the north coast of Canada.
Phytophagous aquatic Coleoptera provide information on the plant species present. Macroplea appendiculata
lives under water on various species of Potamogeton (pondweeds), Myriophyllium (milfoils), aquatic species of
Ranunculus and Nymphaea (water lilies). Donacia versicolorea feeds exclusively on Potamogeton natans (Koch
1992; p.52). The aquatic weevil Bagous lives on submerged water plants and its larvae feed on their stems and
leaves. It is a very poor swimmer and is thus confined to standing waters. Hydrobius fuscipes is a detritus pond
species tolerant of some brackishness in the water.
Hints of brackish habitats are provided by Berosus spinosus which is strongly halophilous being found in shallow
pools on salt marshes. Other species that are at least salt tolerant include Ochthebius lenensis and Cercyon marinus.
Bembidion aeneum is by far the most abundant species of carabid beetle in this assemblage. It is a very common
member of fully glacial and Late-glacial faunas in Europe and also tolerates brackish conditions (Lindroth 1985, p.173).
3.3.1.2. Terrestrial habitats indicated by the Coleoptera. Many species of the Hydrophilidae are not water
beetles; many species of Cercyon live in rotting plant detritus and sometimes in dung. Coelostoma orbiculare
occurs in marshy places where the margins of a pond are choked with dead or dying vegetation.
The carnivorous or scavenging ground beetles, the Carabidae, are represented by twenty species. Notiophilus
aquaticus is a xerophilous species that lives in dry open ground often on what would appear to be completely sterile
gravel and in montane environments it is associated with ericaceous heath (Lindroth 1985, p.78). Blethisa
multipunctata is a hygrophilous species often associated with Agonum versutum (Lindroth 1985) living in marshy
fens with Carex and Eriophorum but not where Sphagnum is predominant in the vegetation. The species have a
preference for sun-exposed ground. Elaphrus cupreus is a hygrophilous species that prefers eutrophic fens.
Elaphrus riparius prefers sparsely vegetated sun-exposed habitats on both clay and sandy soil. Dyschirius tristis is a
subterranean species that excavates burrows in firm clay soil beside water where the soil is bare in places otherwise
densely vegetated with Carex. This species prefers, but is not confined to, saline habitats. Trechus secalis is found in
both open country and woodland, under plant debris in meadows on clay-rich soils. Seven species of Bembidion
occur in this assemblage; these are highly active scavengers or predators living in damp habitats where they are
often dependent on particular types of soil, except Bembidion nigricorne which is xerophilous and lives on dry,
sandy, Calluna heaths in sparsely vegetated places. Bembidion bipunctatum occurs near water on a variety of soils
from stony gravel to silty sites with Carex or grasses. Bembidion obliquum occurs on soft moist ground with a
growth of Carex, grasses and Equisetum on clayish or peaty soils with sun-exposed patches. Bembidion
quadrimaculatum is a heliophilous species, living on clay-rich or sand clay soils with thin vegetation on the drier
banks of fresh water. Bembidion doris is a very hygrophilous species in wet highly vegetated places characterized
by, for example, Carex and Eriophorum. Poecilus cupreus inhabits moist meadow-like habitats with a dense
vegetation of sedges and grasses, and preferably clay soil. Pterostichus minor is found in wet habitats particularly
beside eutrophic ponds. Pterostichus adstrictus lives in open grassy places especially on gravelly soil that has a
mixture of clay and humus. Calathus melanocephalus is a very common species in open country, in dry grassy
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places, especially on sandy soils. Agonum versutum is also a species of open country, often where there is a
luxuriant growth of Carex and where the substrate is soft and muddy. Amara alpina is one of the most characteristic
carabid species of montane and tundra environments, ranging from shrub tundra to almost sterile ground with only
isolated patches of vegetation.
Species of Staphylinidae are mostly carnivores, preying on soil arthropods and worms. Many of the species in
this fossil assemblage live under plant debris in damp conditions, including Eucnecosum brachypterum, Acidota
crenata, Lesteva longelytrata, Geodromicus nigrita, Oxytelus rugosus, O. nitidulus and O. laqueatus. Platystethus
cornutus, the most abundant species here, is fossorial, digging tunnels in sandy banks. Bledius is not a carnivore but
feeds on algae in the superficial layers of sandy and silty soils Tachinus fimetarius prefers open, rather dry sandy
places where it lives under vegetable debris or dung.
Most species of byrrhids are strictly moss-feeders, both as adults and as larvae. Byrrhus, Cytilus, Curimopsis and
Simplocaria form a group that live in microhabitats with loose, well-drained granular soils which are often skeletal
and usually include a high proportion of sand, gravel, cobbles or broken rock. Typically the mosses Ceratodon
purpureus and Polytrichum juniperinum dominate these habitats in North America which also often contain many
weedy and pioneer plant species (Johnson 1987, p.126).
Other plant-eating beetles that feed exclusively on macrophyte vegetation belong to the Chrysomelidae and
Curculionidae. One of the most abundant is the weevil Sitona sp. whose larvae live exclusively on the roots of
Papilionaceae. The relatively high numbers of this weevil indicate pioneer vegetation invading an area of raw soils
with a poorly developed nutritional status. Several other species of beetle indicate the preponderance of weedy
species in the neighbourhood. Galeruca tanaceti lives on a variety of Compositae, but above all on Tanacetum
vulgare (tansy) and Achillea millefolium (yarrow). Adoxius obscurus feeds on Epilobium (willow herbs), its larvae
feeding on the roots of the same plant. Miarus campanulae is confined to Campanulaceae in rather dry places where
its larvae develop in galls on the seed heads. Hypera elongata is chiefly found on Myosoton, in particular the water
chickweed M. aquaticum, but it is also known on species of Stellaria (chickweeds and stitchworts) and on
Cerastium arvense (field mouse-ear). Otiorhynchus ovatus is a xerophilous weevil that feeds on a wide variety of
herbaceous plants. Barynotus squamosus is also polyphagous on low plants but in rather damper habitats. The only
evidence of shrubs is provided by three species, Melasoma collaris, Phyllodecta and Rhynchaenus folorium which
feed primarily on Salix. Of these only the latter species is at all common and it is an extremely small weevil that
likely fed on one of the dwarf willows. Chaetocnema hortensis feeds on grasses and is somewhat salt tolerant.
Species of Coccinellidae are predatory on aphids. Thus, although no aphids were recovered from the sediment,
no doubt because their frail cuticle failed to be preserved, it can be inferred with certainty that many plants were
infested with aphids. Hippodamia arctica is a species of the far north of Europe and is currently extinct in the
British Isles. It is found on small Salix bushes, Betula nana and Arctostaphylos alpinus (Strand 1946).
Specialist dung beetles are important in the context of these mammoth remains since they would be expected to
occur in great profusion had the mammoths been present at the time of sedimentation (e.g. Coope et al. 1961 and
Bosemier et al. 2003). Two species were present, Aphodius prodromus (Brahm.) and A. distinctus (Mull.), mostly
the former species but, since they are not always possible to separate from one another they are included in the
Tables as Aphodius spp. Both these species are obligate coprophages rarely found in rotting vegetable matter.
However, only in sample E is Aphodius prodromus at all common.
3.3.2. Sample C
A list of the Coleoptera recovered from sample C is given in Supplementary Table S6. The sample was obtained
from sediment in situ from sedimentary Unit C1. The fauna was almost identical to the assemblages from samples
A, B, D and E and thus suggests that Unit C1 was the mammoth stratum, as suggested also by the palynology and
the radiocarbon data (Scourse et al. 2009).
3.3.3. Samples from Profile ZFace 2; Unit C1
At this locality the dark grey clayey sandy silt (Unit C1) was arched into a sharp anticline (Scourse et al. 2009).
Detailed sampling of this stratum was undertaken because the sediment was similar to the matrix of the mammoth
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bones and it was one of the few opportunities to examine the stratigraphy in situ. The assemblage of Coleoptera
from the sequence of samples taken through this exposure of Unit C1 is shown in Supplementary Table S7 with
sample 010 cm taken from the top of the stratum and 9095 cm from the base. This suite of Coleopteran species
leaves no doubt that this outcrop of Unit C1 correlates with those samples from Unit C1 elsewhere and from the
sediment adhering to the mammoth bones, though no mammoth bones were found at this locality. Unfortunately,
the Coleopteran assemblages provided no evidence of any environmental changes though, again, the fauna includes
both cold-adapted species as well as relatively temperate ones.
Out of a total of 35 species recorded here, only six were not found in samples AE. Furthermore, the species that
are common to both assemblages include the exotic forms that are now absent from the British Isles. These include
Helphorus splendidus, whose occurrence in the British Late-glacial appears to be restricted to the period of cold
climate just prior to 15 440 cal. year BP (13 000 14C years BP) (e.g. Coope and Brophy 1972) i.e. to Greenland
Stadial 2 (Walker et al. 1999; Lowe et al. 2008).
Clivina fossor is a subterranean species that burrows in rather damp clay-rich soil in open country. Bembidion
virens is characteristic of sterile, gravelly or stony stream banks or similar lake margins and is always found near
water in company with Bembidion bipunctatum (Lindroth 1985, p.182). Bembidion quadripustulatum is a relatively
southern species compared with most other species in this assemblage, in Britain only reaching as far north as
Derby and in Scandinavia only occurring as accidental individuals on the south coast of Scania. It is found on damp
sun-exposed clay or sandy mud. Bembidion minimum is a halophilic species that lives on salt marshes on moist clay
where the vegetation is sparse and leaves bare patches where the beetle is often found in direct sunlight.
3.3.4. Sample F
This beetle assemblage, obtained from the mandibular canal in the lower jaw and from the sinus cavities in the
juvenile skull when these were being prepared in the laboratory, is very different from those described above
(Supplementary Table S6). The total weight of sediment involved was less than 1 kg yet the amount of insect
material recovered was prodigious. By far the largest quantity of insect fragments were the broken pupal cases of
the fly Phormia terraenovae, the larvae of which must have infested the body of the young mammoth (see
Erzinclioglu, in Lister 2009). Almost all the beetle fragments were of Aphodius prodromus, indicating that dung
was abundantly available at this time and, by the way the highly organic material had been forced into the jaw
cavities, it seems likely that the skull bones could have been trampled into the layer of dung. A smaller amount of
similar material was also recovered from the sinus cavities of the juvenile skull fragment. In this context it is notable
that the fly puparia were concentrated at the back of these cavities whilst the sediment rich in Aphodius remains was
more superficial, suggesting that the fly puparia were in place first before the dung beetles were forced into the
cavities. The large number of individuals of Aphodius (n 128) from this small sample is undoubtedly an
underestimation of their abundance (heads were the only skeletal element counted). This sample demonstrates just
how common the dung beetles must have been at the time when the mammoths were present at the Condover site
and contrasts markedly with their poor representation in the samples of dark grey clayey sandy silt matrix encasing
the majority of the mammoth bones.
Necrobia violacea is a corpse beetle that feeds on the larvae of flies and other insects in dried-out carcasses that
have been reduced to skin and bone. The occurrence of large numbers of Aphodius and the presence of Necrobia
shows that at this time, at least, both the dung and the bodies of juvenile mammoth must have lain about on the
surface for sufficient time for the carcass to be largely consumed by blow fly maggots and subsequently to have
become desiccated. This may account for the fact that some of the bones of the juvenile skull appeared to be rather
more decomposed than the rest of the skeleton, suggesting that they had been subjected to weathering prior to their
incorporation into the sediment.
The assemblage from sample E bears some similarities to that from sample F. Aphodius is moderately abundant
in sample E in marked contrast to samples A, B, C and D, also from the dark grey clayey sandy silt of the mammoth
bone matrix, and from Unit C1. Necrobia violacea was also present in both samples E and F, suggesting that these
comprised an admixture of the dark grey clayey sandy silt with some of the dung-rich material from the surface
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represented by sample F. Some evidence of this physical mixing of the grey silty clay with more organic material
was visible in the field, in the form of scattered clasts of darker mud scattered through the grey matrix.
3.3.5. Sample G
There are a number of important differences between the assemblage from samples from sedimentary Unit C1
(samples AE; Supplementary Table S6) and Sample G taken from the green detritus mud (Unit C3) directly
overlying it. There is a total absence in sample G of the suite of cold-adapted species that are a distinctive
component of the beetle assemblages in the dark grey clayey sandy silt of Unit C1. These absentees include Amara
alpina, Potamonectis griseostriatus, Agabus arcticus, Ochthebius lenensis, Helophorus obscurellus, Helophorus
sibiricus, Helophorus splendidus, Eucnecosum brachypterum, Simplocaria metallica, Curimopsis cyclolepidia and
Hippodamia arctica. The absence of some of these species could be attributed to sampling inadequacy, but the total
lack of them all is not so easily dismissed, especially since some of them were relatively abundant in samples AE.
In their place the fauna from sample G includes a number of relatively southern species such as Agonum
sexpunctatum, Coelambus impressopunctatus, Hygrotus inaequalis, Potamonectis assimilis, Plateumaris affinis
and Notaris scirpi, none of which are obligate northern species. A significant feature of this sample that
distinguishes it from the other samples is the large numbers of bones and spines of Gasterosteus aculeatus, the
three-spined stickleback, that were recovered from it as a by-product of the extraction of the insect fossils. No fish
remains of any sort were found in samples AE, in spite of the use of exactly the same extraction procedure. These
differences between the faunas of Unit C1 and Unit C3 indicate that the local environment and the regional climate
were also very different and would suggest that the two units are not just lateral variations of one another.
The occurrence in this sample of moderate numbers of elmid beetles indicates that here, as in sample E, there
must have been running water available. Potamonectes assimilis lives in streams or pools with a sandy bottom
where there is little vegetation (Nilsson and Holmen 1995). In contrast, Coelambus impressopunctatus and
Hygrotus inaequalis are species of well-vegetated ponds or very slowly moving water. The relatively large number
of individuals of Gyrinus indicates that the water surface was not completely covered with vegetation.
Bembidion nigricorne is a species of dry sand heathland, or moderately humid peaty soils where it occurs in
sparsely vegetated places. Agonum sexpunctatum is a strongly heliophilous species of open country on moist soils
that are exposed to the sunlight and sparsely covered with mosses or grasses.
3.3.5.1. Climatic interpretation of the beetle assemblages. In a fauna that is dominated by cold-adapted species,
the Coleopteran assemblages from Unit C1 and samples AE contain an admixed element of temperate forms that
are incompatible in terms of their present day geographical ranges; some high arctic species occur in the same
samples as southern English ones. The cold element in the beetle assemblage includes Bembidion virens,
Helophorus obscurellus, Amara alpina, Helophorus sibiricus, Potamonectes griseostriatus, Helophorus
splendidus, Agabus arcticus, Eucneiosum brachypterum/norvegicum, Agabus congener, Limplocaria metallica,
Colymbetes dolabratus, Curimopsis cyclolepidia, Ochthebies lenensis and Hippodamia arctica. The warm
element includes Bembidion minimum, Hygrotus inaequalis, Bembidion quadripustulatum, Chaetarthria
seminulum, Bembidion nigricorne, Berosus spinosus, Pterostichus minor and Oulimnius tuberculatus.
This mixture cannot be explained by invoking extreme variation in local microclimates, especially since the
topography of the Condover district is so flat that there is very little altitudinal contrast within the catchment area
from which the fossil insect assemblage was drawn. Theoretically no-analogue assemblages (i.e. assemblages of
species which do not live together today) are possible, because of lag effects at a time of rapidly changing climate
and the possibility that modern ranges are partly determined by competitive exclusion and not just individual
species physiological boundaries. However, this is thought an unlikely explanation for the mixed assemblage
because the known pattern of Late-glacial faunal change from other British sites indicates clear transitions from
cold to warm associations and not disharmonious faunas. For instance, at Glanllynnau (Coope and Brophy
1972), the northern and southern species were sequentially separated and clearly came from periods of quite
different climatic conditions. Thus, the mixture at Condover cannot be ascribed to a no-analogue situation. It is
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suggested that this mixture of species comprises animals that lived at different times in contrasting climates,
physically brought together into the same sediment.
As the beetle assemblage from the green detritus mud of sample G (Unit C3) showed no evidence of any mixing,
it was possible to apply the Mutual Climatic Range (MCR) technique (Atkinson et al. 1987) to derive atmospheric
palaeotemperatures. The mean July temperature is reconstructed to be 15198C and the mean January temperature
to 13 to 68C; this implies that the mean monthly temperatures lies somewhere between these values and not that
the temperatures varied between these limits.
3.4. Other environmental proxies
3.4.1. Anuran remains
(J. Alan Holman (deceased), Michigan State University Museum, East Lansing, Michigan, 48825-1045 USA.
Personal communication 1990)
Late-glacial amphibians have been very rarely reported from Britain (Stuart 1982) and have rarely been
specifically identified. Thus, the recovery of anuran (frogs and toads) bones that may be assigned to species from
the Condover mammoth site is of interest. The anuran bones were found in the green detritus mud (Unit C3) where
they were in association with fish remains, mainly Gasterosteids (sticklebacks). This anuran-bearing stratum was
directly above Unit C1, the hypothesized source of the mammoth remains.
The finds were a left femur (Figure 10A), a metatarsal (Figure 10B), and an omosternum (Figure 10D). The frog
bones are well preserved and mainly black in colour, with a very slight reddish tint. Measurements of the remains
are given in Supplementary Table S8.
All of the anuran bones have been assigned to the living species Rana temporaria Linnaeus, 1758, which occurs
today throughout Europe east to the Urals, but excluding most of Iberia, much of Italy, and the southern Balkans
(Arnold and Burton 1978; Frost 1985).
The femur: Fossil anuran femora have rarely been identified to either the generic or the specific level. This is
probably because it is difficult to distinguish left and right fossil specimens, and thus difficult to compare specific
structures on the femora of fossil and modern skeletons. Left and right ranid femora are usually distinguished on the
basis of a groove that occurs on the lateral side of the posteroventral surface of the bone (Figure 10A). At the generic
level the Condover femur may be identified as Rana as follows. It is distinguished from Bufo (Bufonidae) by having
Figure 10. (A) left femur (ventral view) of Rana temporaria from the Condover mammoth site; (B) posterior two-thirds of right femur (ventral
view) of modern Rana esculenta; (C) metatarsal of Rana temporaria from the Condover mammoth site; (D) omosternum of Rana temporaria
from the Condover mammoth site. All of the lines represent 1 mm.
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a more gracile femur and by lacking a posteroventral crest. It is distinguished from Alytes, Bombina, and
Discoglossus (Discoglossidae) by lacking a distinctly S-shaped shaft, having less swollen anterior and posterior
ends, and lacking a posteroventral crest. From Hyla (Hylidae) it is distinguished by having a less gracile femur and
in lacking a posteroventral crest. Finally, it is distinguished from Pelobates and Pelodytes (Pelobatidae) by having a
more gracile and much less strongly bowed femur.
At the specific level the Condover femur may be assigned to Rana temporaria on the basis of the deep lateral
groove that occurs on the posteroventral surface of the femur. This portion of the femur is smoother and more
rounded in Rana esculenta (Figure 10B) and Rana ridibunda. Moreover, femora of other European Rana appear to
be generally less gracile than those of Rana temporaria, especially British specimens.
The metatarsal: The fossil metatarsal (Figure 10C) represents a larger frog than the one represented by the femur,
thus at least two individual frogs are indicated by the fossil material. Although the bone appears to be indistinguishable
from modern Rana temporaria, it was not possible to tell whether it was from the left or the right foot.
The omosternum: The presence of a bony rather than a cartilaginous omosternum is an important diagnostic
character of the family Ranidae and the genus Rana (Duellman and Trueb 1986, p. 542). The family Bufonidae
usually lacks an evidence of an omosternum (as in Bufo bufo), and when it is present, it is cartilaginous and
normally would not fossilize. The other European anuran families (Discoglossidae, Hylidae, Pelobatidae) have
cartilaginous omosterna. The bony omosternum of Rana is normally subtriangular (Figure 10D) and has a wide
posterior articular portion and a constricted anterior portion. The fossil bone was indistinguishable from those of
modern Rana temporaria. The omosternum also represents a much larger specimen than the one represented by the femur.
Rana temporaria has previously been reported from the Middle Devensian site at Upton Warren, Worcestershire
(Coope et al. 1961). At this site, silt bands within the sands and gravels yielded pollen spectra that indicated
treeless, herb-dominated vegetation. One of the silt bands yielded a radiocarbon date of 42 100 800 14C year BP.
As at Condover, the Rana temporaria bones at Upton Warren were associated with those of the three-spined
stickleback (Gasterosteus aculeatus). The only other British Devensian herpetological records are Rana sp. and/or
Bufo sp. from the early Devensian site at Coston near Barnham Broom, Norfolk (Allison et al. 1952; Stuart 1982).
It seems noteworthy that the two Devensian herpetological species that have thus far been identified from Britain,
Rana temporaria and Lacerta vivipara, are forms that occur well within the Arctic Circle in Europe today (Arnold
and Burton 1978; maps 36 and 70). Moreover, of the 126 herpetological species reported in Europe by Arnold and Burton,
Rana temporaria and Lacerta vivipara are by far the most widely distributed herpetological species in far northern areas.
Today, Rana temporaria is very widespread in Britain and may be found in a variety of damp situations where
adequate herbage exists. Nevertheless, it is most often found near ponds (Smith 1964), and may stay in the water for
a month or more after the breeding season (Frazer 1983). It would seem likely that the fossil frogs lived in or near
the basin in which the mammoths became entrapped. The fact that the delicate anuran bones are well preserved
indicates that there was little or no post-mortem transport.
3.4.2. Trichoptera and Chironomidae
(N. E. Williams (retired), then Division of Life Sciences, Scarborough Campus, University of Toronto, 1265
Military Trail, Scarborough, Ontario. M1C 1A4 Canada. Personal communication, December 1991)
Two samples, one from Unit C1 (dark grey clayey sandy silt) and the other from Unit C3 (green detritus mud)
were examined. Trichopteran (caddisfly) and chironomid sclerites were extracted from the sediments, using the
same techniques as for the Coleoptera, and were mounted on glass slides in Berlese fluid. Totals of six caddisfly, and
twelve chironomid taxa were identified (Supplementary Table S9).
All of the aquatic insect taxa identified from Unit C1 are found in Britain today. There were no aquatic taxa
restricted to arctic, subarctic or boreal regions. Limnephilus stigma/nigriceps, L. vittatus and Agrypnia obsoleta are
all now widely distributed in northern and central Europe and European Russia, although L. vittatus does not reach
the northernmost parts of Scandinavia (Botosaneanu and Malicky 1978). The sand-cased L. vittalus is associated
with sandy and silty substrates of lakes, ponds and temporary pools, while both L. stigma and L. nigriceps occur
among dense emergent marginal vegetation in still and running waters, using plant materials for food and case
construction (Wallace et al. 1991). The predaceous Agrypnia obsoleta occupies all types of freshwaters but requires
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plant materials for case construction. In Britain it occurs mainly in upland areas of Scotland, northern England and
Wales, but it is more widespread in Ireland.
Two additional caddisfly species were present in the green detritus mud (Unit C3) sample. Both Lepidostoma
hirtum and Oecetis ochracea are now common and widespread throughout Britain, central and northern Europe and
European Russia (Botosaneanu and Malicky 1978).
The chironomids provide evidence for conditions at the time of deposition of Unit C3. Chironomini, excluding
Chironomus, Sergentia and Stictochironomus, are included by Walker et al. (1991) amongst the primarily
temperate, littoral taxa with distributions that do not extend beyond the tree line. They state that although most of
these thermophilous Chironomidae never individually compose a large proportion of the chironomid fauna of a
lake, their overall relative abundance declines rapidly with increasing latitude. In their study of Labrador lakes, all
sites where temperate littoral chironomids represented more than 10% of the total chironomid specimens counted
were south of 558N and mostly in the boreal spruce-fir forest vegetation zone; in tundra lakes this group comprised
05% of chironomid specimens. In the Unit C3 sample head capsules of members of this group associated with both
sediments and littoral plants were common. Of the remaining taxa, the detritivorous Chironomus sp. was abundant.
This genus has a worldwide distribution, but tends to be extremely abundant in very productive temperate lakes.
The considerable change in the chironomid assemblage between Unit C1 and Unit C3 would seem to indicate
warmer conditions during deposition of the upper sediments or a lag in arrival of thermophilous species caused by
very rapid warming. The latter is a definite possibility from the point of view of aquatic species since there were no
obligate northerners present in either sample.
3.4.3. Ostracoda
(Eric Robinson (retired), Department of Earth Sciences, University College London, now at Riverside Farm,
Whitehall, Watchet, Somerset, TA23 0BB, Personal communication, February 1988)
Investigations using standard techniques (e.g. Chapter 5 in Griffiths and Holmes 2000) were made of samples of
sediments from Unit C1 Profile Z (1988). These revealed a restricted species diversity of five species, of which one,
Limnocythere blankenbergensis (Diebel 1968), dominated, forming a species life assemblage comprising 85% of
the total assemblage. Originally described from Late Weichselian lake sediments in North Germany (Diebel
1968), it has not been found alive and this is its first record from Britain. It forms a dominant component of the
Late-glacial (Older Dryas) ostracod assemblage at Duvensee, North Germany (Gunther 1986) where, along
with Limnocythere inopinata (Baird, 1843) and Candona candida (Mueller, 1776), two species also found at
Condover, it has been interpreted as a characteristic shallow limnic, pioneer species capable of tolerating low
temperatures. Candona candida and Ilyocypris bradyi Sars, another of the Condover assemblage, are also
stenothermal, cold-water species (Klie 1938) often characteristic of pioneer assemblages of the Late-glacial
(Absolon 1973). The structure and context of the Condover ostracod assemblage from Unit C1 is therefore entirely
consistent with the age and environmental context of similar assemblages from the Weichselian Late-glacial of
North Germany.
3.4.4. Mineral magnetic measurements and the occurrence of greigite
(J. P. Smith (retired), Division of Environmental Science, School of Applied Sciences, University of
Wolverhampton. Personal communication, 1992)
Because of the disturbed nature of the Condover site, it seemed likely that simple, rapid and non-destructive
mineral magnetic measurements, that can be made on small samples of sediment, might be an aid to reconstructing
the general stratigraphy of the site, in providing a stratigraphic context for the excavated mammoth bones (cf.
Walden et al. 1992), and for gaining insights into the provenance of the basin infill.
Magnetic minerals can be both from primary (minerals formed in a rock at its time of formation) or secondary
(formed at the time or after deposition of the sediment by processes such as pedogenesis, fire or diagenesis/
authigenesis) (Oldfield and Thompson 1986). In sedimentary sequences such as that at Condover it might be
expected that the magnetic mineral assemblages would be dominated by the clastic detrital input to the basin.
Although the magnetic signature of sediment responds to the mixture of magnetic grain species, their size, shape
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and stoichiometry, in this environmental setting the likely contained magnetic minerals would be various
magnetites and haematites.
The values obtained for magnetic parameters can be used to characterize the contained magnetic mineral
assemblage. Supplementary Table S10 gives the data for three magnetic parameters for the sediment samples.
Magnetic susceptibility (x) is a measure of the concentration of ferrimagnetic minerals present, while the other two
parameters (IRM100mT / IRMSat and IRMSat/x) represent the magnetic mineral mixture and magnetic grain sizes
present. The same measurements for sediment samples scraped from the mammoth bones and for sediment believed to
be from the same stratum but not in immediate contact with the bone are also shown in Supplementary Table S10.
The parameter values for samples taken from the exposed faces are typical of Triassic substrates and Triassic
derived drift (Smith et al. 1990) and reflect differences in the mixture of detrital magnetites and haematites in the
deposits. However, the sediment in contact with the bone has a magnetic susceptibility value an order greater that of
the sediments in the basin and an IRMSat/x ratio much higher than that most other sediments. The only similarity is
with the sediment believed to come from the same stratum.
3.4.4.1. Magnetic extracts and XRD. The magnetic characteristics of the sediment attached to the bone
suggested a high concentration of a multidomain ferrimagnet, which was initially thought to be magnetite. A
magnetic extract was made from the sediment and the material analysed by XRD. The magnetic mineral was
identified as greigite, a ferrimagnetic spinel with the chemical formula Fe3S4.
Greigite is a metastable iron sulphide formed in aquatic environments normally as a member of a series between
amorphous FeS and pyrite. It is common in marine sediments but has also been recognized in freshwater systems
(Hilton 1990). The mineral is not stable in aerobic conditions and was not therefore part of the clastic input to the
deposit, but must have formed authigenically, as a result of the particular geochemical environment existing in the
deposit at the time of deposition or, diagenetically, soon after burial by overlying deposits. Adapting the findings of
Hilton (1990) and others, who have worked on the occurrence of greigite in limnic systems, the requirements for the
formation of greigite appear to be a source of labile iron, reducing conditions and a source of organic sulphur.
Such a geochemical environment must also be one that is consistent with the preservation of the mammoth bones
and other organic remains. The excess of greigite adjacent to the bones presumably reflects the availability of
organic sulphur. Some of the unusual effects reported by other scientists working on the remains (e.g. magnetic
insect remains, oxidized coatings on insect remains after separation and drying, and the corrosion of pollen grains
during chemical treatment) all reflect the coating of organic surfaces by the authigenic greigite. The greigite clearly
is a manifestation of the extreme geochemical environment responsible for the high quality of the preservation of
organic remains at the site. Presumably, the very high organic loading drove geochemical reactions towards the kind
of reducing conditions necessary for preservation.
Detailed comparisons of the magnetic signature of the sediments from the same stratum but not in immediate
contact with the bone gave similar characteristics, but the lower magnetic susceptibility suggested a much lower
concentration of the mineral. It therefore seems likely that the greigite was contained in the stratum containing the
mammoth bones and was selectively concentrated in and around the bones themselves.
4. ENVIRONMENTAL INFERENCES
Inferences based upon the range of environmental indicators are presented for the sedimentary units investigated.
The ages of these units are derived from a series of radiocarbon dates, calibrated using OxCal v 4.0 and calibration
data set IntCal04, reported in full in Scourse et al. (2009). The Devensian Late-glacial and early Holocene
environmental history of the basin at Norton Farm, Condover, is summarized in Table 1.
4.1. Units C1 and C3: Before 13 82014 830 cal. year BP (12 240 W 120
14
C year BP)
These units, which were the source of the mammoth bones, were the most intensively studied with data available for
all the proxies investigated.
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Characeae, (calcareous algae,
stoneworts) and pondweeds. Fringing
communities of sedge, club-rush
(Scirpus lacustris) and marsh cinquefoil
(Potentilla palustris). The presence
various species of Potamogeton,
Myriophyllium, Nymphaea and aquatic
species of Ranunculus are all indicated
by the aquatic plant-eating Coleoptera.
The Coleoptera also indicate a mosaic
of slow moving and stagnant water and
fast flowing well-aerated water. Frogs
(Rana temporaria) were present in the
vicinity and sticklebacks (Gasterosteus
aculeatus) were present in the water
body. The chironomids and caddis flies
indicate temperate conditions
Before and around 14 83013 930 cal. year BP

(12 300 110 14C year BP)
HIATUS AT PROFILE C
Fringing communities of sedge

(Carex) with helophytic Ranunculus.
Rich abundance of both submerged,
floating-leaved and submergent
waterplants, including water-milfoil
(Myriophyllum), marestail
(Hippuris vulgaris) and pondweeds
(Potamogeton). Diverse/abundant
helophytic assemblage
HIATUS AT PROFILE C
Aquatic/helophytic environment
within basin
BIOTURBATION
Eutrophic, nutrient rich,
low O2, calcareous
Oligotrophic, nutrient
poor, high O2
Moderately eutrophic,
calcareous
No information
Water in Basin:
trophic status
MAMMOTHS ENTER BASIN
14 83013 930 to 13 70013 250 cal. year BP

(12 300 100 to 11 590 110 14C year BP)
12 86012 380 to ca.11 500 cal. year BP

(10 640 100 to ca.10 000 14C year BP)
13 70013 250 to 12 86012 380 cal. year BP
(11 590 110 to 10 640 100 14C year BP)
ca.11 500 to 10 79010 250 cal. year BP

(ca. 10 000 to 9350 100 14C year BP
After 10 79010 250 cal. year BP

(9350 100 14C years BP)
Table 1. Summary of environmental history
Grass-dominated, but with

weed taxa including
Chenopodiaceae, Rumex and thrift
(Armeria maritima). Some dwarf
willow. No trees, but open birch
scrubland with dwarf birch
(Betula nana) towards 14 400 cal.
year BP (12 300 14C years BP).
A generally barren, sparsely
vegetated, landscape is indicated
by the Coloepteran assemblage.
The weevils indicate pioneer
vegetation. Ladybirds were present
indicating the presence of aphids,
and dung beetles indicate the
presence of some large
dung-producing animals
Grass-dominated, but with

Artemisia, Rumex and Thalictrum
Open birch woodland, with tree
and dwarf forms, juniper,
poplarprobably aspen (Populus)
and willow
Successive colonization by juniper

(Juniperus), willow (Salix), birch
(Betula), hazel (Corylus), pine
(Pinus) and alder (Alnus)
Dry land vegetation outside basin

437
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The plant macrofossil data from Unit C1 clearly indicate the local presence of a number of taxa recorded in the
pollen spectra, including Salix sp(p)., Betula sp(p)., Chenopodiaceae, Potentilla sp., Myriophyllum verticillatum,
Armeria maritima, and species of Gramineae and Cyperaceae, including Juncus sp(p)., Scirpus lacustris,
Eleocharis palustris and Carex spp. The decrease in abundance of plant macrofossils upwards through the series of
samples from Profile Z parallels the decline in pollen concentrations from the same profile. As with the
macrofossils, this profile was notably less rich (in terms of pollen concentrations) than Profile C. This may be
explained in terms of Profile Z lying closer to the centre of the fluvial channel (Scourse et al. 2009), Profile C being
situated in a marginal backwater in which the vegetation cover was more dense, and in which plant detritus
accumulated.
The plant macrofossil data from Unit C1 and the bone scrape samples indicate a water body possessing a modest
aquatic flora, possibly dominated by stoneworts (calcareous algae; family Characeae). These are typical colonizers
of newly created freshwater ponds and lakes of calcareous status. In contrast the high, and increasing, concentration
of Pediastrum (Figures 3 and 4), dominated by P. boryanum typicum, in Unit C1, is indicative of eutrophic water
with high nutrient levels and low oxygen content (Salmi 1963). The absence of P. duplex in Unit C1 may be
significant in that it is not tolerant of highly eutrophic conditions (Salmi 1963; Cronberg 1982).
There were probably fringing plant communities of limited extent with sedges (Carex), club-rush (Scirpus
lacustris) and marsh cinquefoil (Potentilla palustris). Terrestrial plants, whose remains must have been blown in or
washed into the basin through the fluvial channel which was active at this time (Scourse et al. 2009), include a few
taxa, which today would be considered weeds of cultivated and waste places, indicating unstable soils in the
vicinity. Thus, outside the basin itself the vegetation was dominated by grasses with a variety of taxa characteristic
of unstable minerogenic soils; these include allseed (Chenopodium polyspermum), goosefoots in C. Section
Pseudoblitum (probably C. glaucum or C. rubrum), orache (Atriplex sp.) and some of the Rumex sp(p). These
records, in association with thrift (Armeria maritima), are all indicative of a treeless environment. Lack of trees may
reflect cold climatic conditions though none of the taxa recorded are obligate arctic-alpine plants at the present day.
It is likely, however, that the remains of willows (Salix sp(p).) were all from dwarf forms typical of arctic-alpine
habitats. Alternatively, treelessness might be a response to extensive grazing pressure.
One of the plant taxa recorded is of interest from a palaeobotanical point of view. The pondweed from the
Profile Z samples, Potamogeton rutilus, appears not to have been recorded from the British fossil flora (there are
no records in Godwins (1975) compilation). It is commonly known as the Shetland pondweed, being recorded
within the British Isles only from those islands and the Outer Hebrides. Its distribution across Europe is in the
northern, central and eastern parts, as far north as northern Russia, but it is absent from Scandinavia. Such a
distribution lends support to the suggestion that Unit C1 accumulated in an essentially cool temperate to boreal
treeless environment.
The single pollen spectrum from Unit C3 indicates a very similar environment to that characterizing Unit C1;
in the correspondence analysis it plots very close to the C1 samples. The Unit C3 sample has relatively higher
values of Gramineae and lower values of Cyperaceae, and thus, shows a closer affinity to the spectra from the
upper part of the C1 sequence than the lower parts; this is consistent with the stratigraphical position of C3
overlying C1.
The reconstructions from units C1 and C3 provide a picture of the vegetation at Condover immediately before
and during the time the mammoths entered the basin. One could speculate that perhaps the mammoths were
tempted into the basin by the relatively rich vegetation growing in, and on the margins, of the backwaters and slowflowing channels, certainly far richer vegetation than the sparse grassland of the higher dry land surrounding the
basin.
In order to place the Condover record within a regional context it is important to compare the pollen stratigraphy
with well-dated sequences elsewhere. The nearest such record is from Llanilid in South Wales (Walker and
Harkness 1990; Walker et al. 2003), some 86 km to the southsouthwest of Condover. Here the chronology for the
Late-glacial section of the high-resolution pollen sequence is based upon 20 calibrated AMS 14C determinations.
The pollen spectra from zone NF1 are quite similar to the assemblages at Llanilid prior to ca. 14 200 cal. year BP,
corresponding to Llanilid local pollen assemblage zones L-1 and L-2 (Walker and Harkness 1990; Walker et al.
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2003). The major differences are the complete absence of Juniperus, and the reduced significance of Rumex and
Thalictrum at Condover; these can probably be attributed to local site differences and in the case of Juniperus,
which produces fragile pollen grains, to the largely minerogenic nature of the Unit C1 sediments. The Betula rise at
Llanilid starts around 13 600 cal. year BP and reaches levels in excess of 60% total land pollen between then and
13 200 cal. year BP (Walker et al. 2003). The Betula rise at Condover was underway by between 13 700 and
13 250 cal. year BP (11 590 110 14C year BP; OxA-2240), which is effectively synchronous with Llanilid given
the errors on the radiocarbon ages.
The mosaic of micro-habitats suggested by the terrestrial Coleoptera from Unit C1 (samples A, B, C, D, E and
from profile Z, Face 2) together suggest a barren, treeless, landscape, much of it sparsely vegetated with clayey, sandy
and gravelly soils exposed between plant clumps. Nearby in the poorly drained hollows the vegetation was more lush
and dominated by sedges and grasses surrounding eutrophic pools. The beetles provide also a hint of salinity.
Most of the aquatic Coloeptera species live in standing or slow moving water, except for a small number of
Dryopidae which require fast flowing well-aerated water. These flowing-water species occur only in sample E from
Uint C1, but they also occur in the sample from Unit C3green detritus mud (sample G). This probably indicates
that this stream habitat was very restricted at the Condover site, possibly confined to a relatively small channel or
several small channels (Scourse et al. 2009). Various phytophagous beetles indicate the presence of water plants,
such as species of Potamogeton, Myriophyllium, Nymphaea and aquatic species of Ranunculus. Most species of
byrrhids are strictly moss-feeders, both as adults and as larvae. The relatively high numbers of weevils that feed
exclusively on macrophyte vegetation indicate pioneer vegetation invading an area of raw soils with a poorly
developed nutritional status.
The Mutual Climate Range reconstruction for Unit C3 of Tmax (mean July temperature) of between 15198C and
Tmin (mean January temperature) between 13 to 68C is entirely consistent in range and timing with insect-based
MCR reconstructions from Llanilid (Walker et al. 2003) for the earliest part of the Late-glacial Interstadial. This
suggests that Unit C3 represents the initial warming represented by Greenland Interstadial 1e (Lowe et al. 2008) in
the NGRIP ice core record (Rasmussen et al. 2006).
The fossil insects were very corroded in the uppermost 30 cm of Profile Z Face 2 (Unit C1), to such an extent that
few could be recognized, and they were absent altogether from the topmost sample. These upper samples were also
rather brown in colour suggesting that they had at some time in the past been oxidized. It is likely therefore that the
corrosion of the insect fossils was due to weathering at some time after the emplacement of the dark grey clayey
sandy silt of Unit C1. Since the series of samples was taken from the steep limb of the anticline (created by kettlehole subsidence; Scourse et al. 2009) and well away from its crest, it would seem that the weathered top to this unit
was also involved in the fold; the period of weathering therefore pre-dated the formation of the anticline.
4.2. Unit D: 13 70013 250 to 12 86012 380 cal. year BP (11 590 W 110 to 10 640 W 100
14
C year BP)
The occurrence of significant numbers of woody taxa, including Betula, Salix, Populus and Juniperus in zone NF2
is evidence of a marked climatic improvement. The pollen stratigraphical changes at this level, in combination with
the radiocarbon dates, conclusively support the correlation of zone NF2 with the latter part of the Late-glacial
Interstadial (Greenland Interstadial 1a; Walker et al. 1999; Rasmussen et al. 2006; Lowe et al. 2008).
It is likely that there is an hiatus between Units C1 and D in Profile C. This is suggested by the difference between
the two 14C dates which lie either side of the contact; 14 72013 820 cal. year BP (12 240 120 14C year BP; OxA2265) below the contact and 13 70013 250 cal. year BP (11 590 110 14C year BP; OxA-2240) immediately
above. This hypothesis is further supported by the pollen record (Figure 3) in which a number of taxa exhibit abrupt
changes of frequency across the boundary, notably Cyperaceae. This would also account for the absence of Units
C2 and C3 at profile C.
The plant macrofossil samples from Unit D (Profile C; Supplementary Table S4) were larger than the samples
from Unit C1 and produced much larger and more diverse plant assemblages than Unit C1, with a prominent
component of tree birch (probably Betula pendula Roth/B. pubescens Ehrh.) though with some dwarf birch
(B. nana). This is entirely consistent with the pollen evidence (Figure 3) in which Unit D is characterized by large
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relative abundances of Betula, some attributable to B. nana (Birks 1968; Berglund and Digerfeldt 1970). The
presence of buds and fruits of Salix also suggests a local origin for the Salix pollen, some of which was attributable
to dwarf willow, S. herbacea (Birks 1973). Populus is a difficult pollen taxon to identify with certainty, but
the presence of Populus sp(p). bud scales (presumably from aspen P. tremula L.) from Unit D supports a local
source for the recorded pollen. The significant frequencies for Myriophyllum species in the pollen data are
supported by abundant Myriophyllum nutlets in this unit. The pollen records for Cirsium/Carduus and for
Potamogeton are also supported by macrofossil finds.
The appearance of Pediastrum duplex in Unit D suggests that conditions in the basin were slightly less eutrophic
than during the deposition of Unit C1 (Cronberg 1982; Figure 5). Though Salmi (1963) reports that P. duplex is
calciphobic, the abundance of Myriophyllum spicatum (a major component of the aquatics sum (Figure 3) suggests
fairly calcareous conditions (Clapham et al. 1962). There is a reduction in the overall productivity of Pediastrum in
Unit D which is consistent with a decrease in nutrient availability and a reduction in the level of eutrophication.
As at Llanilid (in zone L-2), the Betula peak at Condover is preceded by a distinctive peak of Juniperus; however,
this peak occurs earlier at Llanilid at ca. 14 800 cal. year BP (Walker et al. 2003) than at Condover where it is
registered at, or just after, 13 70013 250 cal. year BP (11 590 110 14C year BP; OxA2240). Similarly, the main
Betula peak at Llanilid appears to occur well before the main peak at Condover. Whether these differences are real,
or an artifact of the resolution of the radiocarbon method, is difficult to judge. A possible climatic deterioration
within the interstadial is associated with a Juniperus rise at Condover, but no such rise occurs at Llanilid. Though
the detailed Late-glacial pollen record from Hawes Water (Jones et al. 2002), 160 km north of Condover, lacks an
independent chronology, the early interstadial Juniperus phase here dates to between ca. 14 100 and 13 750 GRIP
year BP which is comparable to the timing at Llanilid. The arrival of Betula at Condover (ca. 13 70013 250 cal.
year (ca. 11 590 110 14C year BP)) and Llanilid (ca.13 800 cal. year BP (Walker et al. 2003)) coincides fairly
closely with records from sites in southwest England (Brown 1977), but neither site demonstrates the earlier Betula
phase prior to ca.13 990 cal. year BP (12 000 14C year BP) as recorded in north-west England (Pennington 1977;
Jones et al. 2002).
The overall form of the Betula curve within the Late-glacial Interstadial record from Condover is also similar to
the much more detailed sequence at Llanilid (Walker and Harkness 1990; Walker et al. 2003). This takes the form of
a double peak, with the earlier peak reaching the highest frequencies. A climatic oscillation within the Betula phase
has also been recorded, along with an earlier smaller second oscillation, at other sites across the UK including
Gransmoor in NE England (Mayle et al. 1999; Walker et al. 1993), Traeth Mawr (Walker 1982) and Borrobol
(Mayle et al. 1999), and is dated between 13 000 and 12 800 GRIP year BP at Hawes Water (Jones et al. 2002).
A cold event preceding the Younger Dryas has also been recognized in a number of other datasets (Siegenthaler
and Eicher 1986; Atkinson et al. 1987; Broecker et al. 1988; Lehmann and Keigwin 1992). It is possible that the
depression in Betula values at Condover between 60 and 65 cm in Profile C (Figure 3), associated with increases in
herbaceous taxa, might similarly be attributed to this event. It is now recognized that the Late-glacial Interstadial
was a time of inherently unstable climate during which there were at least two such deteriorations (Bjorck and
Moller 1987; Paus 1988, 1989). These events have been attributed to effluxes of very cold meltwater into the North
Atlantic from the rapidly disintegrating Laurentide Ice Sheet in North America (Lehmann and Keigwin 1992). Two
colder episodes, GI-1d and GI-1b, have been recognized within Greenland Interstadial 1 (GI-1) (Walker et al. 1999;
Rasmussen et al. 2006; Lowe et al. 2008); and the latter of these (GI-1b, 13 36113 149 ice core year BP based on
the GICC05 NGRIP timescale; Rasmussen et al. 2006) correlates with this cold event at Condover and Llanilid.
However, a preferred alternative is that the rise in the Betula curve just prior to the climatic deterioration of the
Loch Lomond Stadial, at just over 50 cm depth in Profile C (Figure 3), represents the influx of reworked pollen
associated with the onset of the climatic deterioration. This interpretation is supported by the marked influx of
indeterminable grains at this level. High frequencies of reworked and indeterminable pollen have been recorded
over this transition at a number of other Late-glacial sites in Britain (e.g. Lowe and Walker 1986; Walker and Lowe
1990) and similar features appear in continental pollen diagrams (e.g. Paus 1989).
The transition to stadial conditions at Condover occurs at 12 86012 380 cal. year BP (10 640 100 years BP
14
C year BP) which is effectively synchronous with the transition at ca. 12 600 cal. year BP at Llanilid at (Walker
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441
et al. 2003) and at Gransmoor in NE England (Lowe et al. 1995; Walker et al. 2003), and at Hawes Water at ca.
12 600 GRIP year BP (Jones et al. 2002). The onset of GS-1 in the NGRIP record is at 12 896 ice core year BP
(GICC05 timescale; Rasmussen et al. 2006).
4.3. Unit E: 12 86012 380 to 10 79010 250 cal. year BP (10 640 W 100 to 9350 W 100
14
C years BP)
The transition from the brown sedge peat of Unit D to the grey clay of Unit E is marked abruptly in the pollen
stratigraphy by the change from zone NF2 to NF3 (Figure 3). Unfortunately, no plant macrofossil data are available
from Unit E so it is not possible to separate conclusively local from extra-local or regional sources in the pollen
diagram. However, Pediastrum frequencies rise from the low levels of Unit D (Figure 3) and P. kawraiskyi, P.
sturmii and P. tetras, three species largely absent from the lower levels, become more abundant (Figure 5).
P. kawraiskyi and P. tetras, in particular, are associated with oligotrophic conditions with low nutrient status and
high oxygen content (Salmi 1963; Cronberg 1982). P. kawraiskyi is also an indicator of shallow, cool water, low in
calcium (Salmi 1963; Cronberg 1982). It is widespread in the rivers of Central Europe at the present day (Cronberg
1982), and was particularly abundant in the cold, clear waters of infilling lakes in the Weichselian Late-glacial
(Jankovska and Komarek 1982). This is entirely consistent with its occurrence at Condover in sediments
representing the climatic deterioration of the Loch Lomond Stadial.
This correlation is supported both by the pollen stratigraphy and the radiocarbon dates. The decline in woody
taxa (including Betula, Salix and Juniperus) and the increase in herbaceous taxa (including Gramineae, Artemisia
and Rumex) are consistent with this interpretation. The reduction in organic content from the underlying highly
organic sedge peat into the relatively inorganic grey clay can be attributed to a decline in the productivity of plants
both in and around the basin as a result of climatic deterioration.
The increases in Artemisia, Rumex and Thalictrum in the Loch Lomond Stadial sediments at Condover are
strikingly similar to those at Llanilid and at other sites. The high frequencies for Artemisia recorded from the Loch
Lomond Stadial at a number of sites in Britain have been widely regarded as reflecting not only colder, but also drier
climatic conditions (cf. Lowe and Walker 1986; Tipping 1991a, 1991b; Walker and Lowe 1990).
4.4. Unit F: After 10 79010 250 cal. year BP (9350 W 100
14
C year BP)
There is probably a condensed sequence (Scourse et al. 2009) or an hiatus at the base of Unit F because the date of
10 79010 250 cal. year BP (9350 100 14C year BP; OxA-2234) for this lithological boundary is considerably
later than for other sites in the British Isles, including Llanilid (Walker et al. 2003) and Llyn Cororion (Watkins
et al. 2007). Also, comparison with Crose Mere (Beales 1980) and Llanilid (Walker and Harkness 1990; Walker
et al. 2003) indicates that the distinctive early Holocene Juniperus peak (Tipping 1987) is missing at Condover.
Betula is already quite high at the base of Unit F, and the date of 10 79010 250 cal. year BP (9350 100 14C year
BP; OxA-2234) is consistent with dates for the Betula rise in the early Holocene at Llanilid and other sites in the
western British Isles (County Cork, Coxon 1986; south west Scotland, Moar 1969; County Tyrone, Pilcher 1973;
Pilcher and Smith 1979; Isle of Skye, Vasari 1977; north west Wales, Watkins 1990; Watkins et al. 2007). These
factors all suggest a period of erosion or non-deposition in the sequence at the base of this unit.
4.5. Stratigraphical attribution of the unstratified mammoth bones, environment at the time the mammoths
were present in the basin, and evidence for mammoth bioturbation
It is clear from radiocarbon analyses (Scourse et al. 2009), from the Coleopteran and from the pollen analyses
presented here, that the unstratified mammoth remains can be attributed to units C1 and/or C3. It is not possible to
separate units C1 and C3 on the basis of the radiocarbon determinations, but stratigraphically C3 was seen to overlie
C1 and the Coleoptera and chironomid data suggest much warmer conditions during C3 than in C1 times. Though
the sediment adhering to the majority of the mammoth bones can unequivocally be attributed to Unit C1, and the
radiocarbon data indicate that the bones and Unit C1 are of approximately the same age, there are elements in the
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ET AL.
palaeobiological data which might indicate that the bones are intrusive from above, and that the mammoths entered
the basin later than the deposition of Unit C1.
Crucial to understanding the relationship of the mammoth bones to their matrix are the dung beetles. These occur
in great profusion in deposits that are contemporary with large herbivorous mammals, for example at Upton
Warren, Worcestershire (Coope et al. 1961), and at Lynford, Norfolk (Bosemier et al. 2003). In the samples from
Unit C1 dung beetles were remarkably rare except for sample D and even here they were not as abundant as would
be expected had the mammoths been present at that time. However, the fauna from sample F, intimately associated
with the sinus cavities and dental alveoli of a juvenile skull and jaw, was almost entirely made up of species of the
dung beetle Aphodius leaving no doubt that these bones and their matrix were contemporary. The relative rarity of
dung beetles from samples of Unit C1 strongly suggests that the mammoths were not locally present at this time and
that the bones from these deposits are later intrusions into this layer. After death some of the mammoth carcasses
lay on a dry land surface surrounded by large quantities of dung. Because this dung was embedded in the bone
cavities, it is suggested that the bones themselves may have been subsequently trampled into the deposits in the basin.
It is clear from the sediments, the local stratigraphy (Scourse et al. 2009) and the palaeobiology that the
Condover basin is a Devensian Late-glacial kettle-hole basin. However, the gradients of the marginal slopes of the
basin at the time when the mammoths entered it would not have impeded their exit (Scourse et al. 2009) and some
other cause must be sought for their entrapment. Miring is the most likely explanation. The mechanism for this
miring is provided by the very soft cohesive sediments within, and on the margins of the basin. Biological and
sedimentological evidence for fluvial activity suggests that the basin was shallow and was probably flat-floored at
this time. This sedimentary sequence, containing the mammoth remains, subsided later in the Late-glacial
Interstadial through the melting of the buried ice block beneath the basin (Scourse et al. 2009). Even though the
mammoth bones were found in Unit C1, the paucity of dung beetles from this unit suggests that the abundant insect
fauna of Unit C1 cannot date from the time of the mammoths.
There was evidence both in the field and in the laboratory that the samples of dark grey clayey sandy silt (Unit
C1) were bioturbated. The deposit was far from homogeneous. Discrete clasts of organic matter a few centimetres
in diameter were scattered throughout, and in some cases the original bedding was disrupted. These features were
present in the sediments of Face 2 that was logged in situ. In places this sedimentary mixture involved the
underlying pink laminated clay (Unit B) which appeared to have been soft and plastic at the time, and occurred as
deformed wisps caught up in the mass of dark grey clayey sandy silt (Unit C1).
One possible cause of the physical mixing of the sediments could have been by slumping and sliding of the
deposits from the sides towards the middle of the basin. However, as noted above, at the time of the dark grey clayey
sandy silt (Unit C1), the basin floor was flat and its marginal gradients gentle, so inhibiting any gravitational
movement. A more plausible explanation for mixing of the sediments is that they were trampled by the mammoths
prior to, or even during, their entrapment in the mire. Some of the foot bones had pink clay adhering to them,
showing that at some time they had penetrated right through the layer of dark grey clayey sandy silt (Unit C1) into
the pink laminated clay (Unit B1) immediately below.
This explains the paucity of dung beetles within Unit C1. It is clear from the enormous number of dung beetles in
sample F (from the juvenile bones; Supplementary Table S3) that they had occurred in profusion when the
mammoths were present at the site. The species of dung beetle present in the cavities of the juvenile skull was also
scattered in low numbers throughout Unit C1. It is likely that these specimens were trampled into the mud together
along with the other obvious members of the temperate fauna from Unit C3 that may have been contemporaneous
with the mammoths. It is likely that much of these dung-rich sediment was washed out of the basin before the final
subsidence of its floor as the buried ice mass melted from beneath.
The hypothesis that the mammoth bones are intrusive into Unit C1 implies that the palaeoenvironmental data
from this unit relates to a period before the mammoths died at the site. In this context, it is probable that Unit C3 is
more nearly contemporary with the mammoths than Unit C1; the radiocarbon data would be consistent with this
explanation, although the two units cannot be separated on this basis alone. Whilst the vegetation inferred for the
two units is very similar, Unit C3 contains a more thermophilous, and unmixed, beetle assemblage and the
chronomid larvae suggest a warming between Unit C1 and Unit C3.
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5. CONCLUSIONS
1. Multivariate statistical analyses of the palynological data and similarities of the Coleopteran assemblages,
together with the radiocarbon dates (Scourse et al. 2009), support the attribution of the mammoth bones to Unit
C1, the dark grey clayey sandy silt, and to the immediately overlying green detritus mud, Unit C3.
2. The palaeobiological data supports the correlation of most of these sediments to the Devensian Late-glacial,
with Unit C1 accumulating early within the Late-glacial Interstadial and Units C3 and D accumulating later
during the Late-glacial Interstadial (Greenland Interstadial 1), Unit E within the Loch Lomond Stadial
(Greenland Stadial 1; Younger Dryas) and Unit F in the early Holocene.
3. Unit C1 times were characterized by a fluvial network within an actively subsiding kettle-hole basin with
ephemeral fast-flowing channels, pools and abandoned channels of eutrophic status choked with vegetation and
vegetative detritus.
4. At this time the basin was richly vegetated compared with the barren, treeless, and dry surrounding landscape,
possibly contributing towards the attraction of the mammoths to the site.
5. Towards the end of Unit C1 times, early within the Late-glacial Interstadial, a period of rapid warming
stimulated vegetation change and invertebrate ecosystem response. This time of transition (Unit C3) was almost
certainly when the mammoths entered the basin. Quantitative palaeotemperature reconstruction based on beetle
assemblages indicate mean July temperatures between 15 and 198C and mean January temperatures between
13 and 68C at this time.
6. At the time when the mammoths were present in the basin, the sides were too shallow to have prevented their exit
and it is likely that the mammoths became trapped by being mired in the soft cohesive sediments and that at
times their feet penetrated into the pink laminated clay beneath.
7. Trampling by the mammoths caused extensive mixing of the underlying sedimentary layers, leading to elements
of flora and fauna of different ages being found within a single sample.
8. The mammoth carcasses must have lain on the surface for sufficient time to have decomposed and to have
become desiccated after which their bones were further trampled into the sediment.
ACKNOWLEDGEMENTS
The authors thank ARC Western Ltd for access to the site and for help in exposing the profiles and retrieval of the
bones and sediment samples. Also Geoff McCabe and the staff of the Shropshire Museums service for their help
with the excavation and Eve Roberts for first discovering the bones.
J. Alan Holman was grateful to Dr Adrian Lister (then of the Department of Zoology, University of Cambridge),
for the privilege of studying these anuran fossils. Lisa Hallock made the drawings of the anuran fossils. His work on
British Pleistocene amphibians and reptiles was supported by United States National Science Foundation Grant
Number BSR 851 5665. James Scourse acknowledges a Royal Society-Leverhulme Trust Senior Research
Fellowship.
The authors thank Professor M.J.C. Walker and an anonymous reviewer for many helpful comments on an earlier
version of this paper.
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Allen Et Al. (2009) - Palaeoenvironmetal Context of Woolly Mammoth at Candover, UK

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GEOLOGICAL JOURNAL

Geol. J. 44: 414446 (2009)

Palaeoenvironmental context of the Late-glacial woolly mammoth

Copyright # 2009 John Wiley & Sons, Ltd.

palaeoenvironmental context of condover mammoth site, uk

2. MATERIALS AND METHODS

Geol. J. 44: 414446 (2009)

Geol. J. 44: 414446 (2009)

palaeoenvironmental context of condover mammoth site, uk

Geol. J. 44: 414446 (2009)

2.2. Plant macrofossils

Geol. J. 44: 414446 (2009)

palaeoenvironmental context of condover mammoth site, uk

2.4. Other environmental indicators

Geol. J. 44: 414446 (2009)

Copyright # 2009 John Wiley & Sons, Ltd.

Geol. J. 44: 414446 (2009)

palaeoenvironmental context of condover mammoth site, uk

Figure 4. Profile CResults of Zonation.

Figure 5. Pediastrum percentage diagram for Profile C.

Copyright # 2009 John Wiley & Sons, Ltd.

Geol. J. 44: 414446 (2009)

Geol. J. 44: 414446 (2009)

palaeoenvironmental context of condover mammoth site, uk

Copyright # 2009 John Wiley & Sons, Ltd.

Geol. J. 44: 414446 (2009)

Copyright # 2009 John Wiley & Sons, Ltd.

Geol. J. 44: 414446 (2009)

palaeoenvironmental context of condover mammoth site, uk

Geol. J. 44: 414446 (2009)

Copyright # 2009 John Wiley & Sons, Ltd.

Geol. J. 44: 414446 (2009)

palaeoenvironmental context of condover mammoth site, uk

Geol. J. 44: 414446 (2009)

Geol. J. 44: 414446 (2009)

palaeoenvironmental context of condover mammoth site, uk

Geol. J. 44: 414446 (2009)

Geol. J. 44: 414446 (2009)

palaeoenvironmental context of condover mammoth site, uk

Geol. J. 44: 414446 (2009)

Geol. J. 44: 414446 (2009)

palaeoenvironmental context of condover mammoth site, uk

Copyright # 2009 John Wiley & Sons, Ltd.

Geol. J. 44: 414446 (2009)

Geol. J. 44: 414446 (2009)

palaeoenvironmental context of condover mammoth site, uk

Geol. J. 44: 414446 (2009)

Geol. J. 44: 414446 (2009)

Copyright # 2009 John Wiley & Sons, Ltd.

Before and around 14 83013 930 cal. year BP

Fringing communities of sedge

MAMMOTHS ENTER BASIN

14 83013 930 to 13 70013 250 cal. year BP

12 86012 380 to ca.11 500 cal. year BP

ca.11 500 to 10 79010 250 cal. year BP

After 10 79010 250 cal. year BP

Table 1. Summary of environmental history

Grass-dominated, but with

Grass-dominated, but with

Successive colonization by juniper

Dry land vegetation outside basin

palaeoenvironmental context of condover mammoth site, uk

Geol. J. 44: 414446 (2009)