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BEETLE RECORDS | Late Pleistocene of Europe
Late Pleistocene of Europe G Lemdahl, Linnaeus University, Kalmar, Sweden G R Coope,{ Royal Holloway, University of London, Egham, UK ã 2013 Elsevier B.V. All rights reserved.
Introduction In this article, the response of Coleoptera (beetles) to Upper Pleistocene climatic changes are summarized. This should cast light upon our understanding of the nature of these changes and thus enable us to predict the ways in which other animal and plant species might respond to any future climatic events. By using the changes in the range of beetle species during the Quaternary, it has been possible to quantify thermal climatic conditions and to estimate both the amounts and the rates of change involved. Furthermore, it has been possible to map the regional differences in thermal climates within northwestern Europe. The beetle assemblages described here include those from the Last (Eemian, Ipswichian) Interglacial, the Last (Weichselian, Devensian) Glaciation, and the transition to the present (Holocene) Interglacial. Most of the examples have been drawn from sites in the British Isles and northern Europe because it is there that these faunas have been most intensively investigated. The beetle record shows that the climate oscillated rapidly between temperate–oceanic and arctic–continental conditions. Where possible, the fossil insect assemblages have been correlated with marine isotope stages (MIS) or with the event stratigraphy of the Greenland ice cores. In the account that follows, paleotemperature estimates are given using the mutual climatic range (MCR) method (Atkinson et al., 1987) where Tmax indicates the mean temperature of the warmest month (July) and Tmin indicates the mean temperature of the coldest months (January and February; Figure 1).
Upper Pleistocene Beetle Assemblages Eemian Interglacial (MIS 5e) Extensive beetle assemblages of this age are known from seven localities in southeast England (Coope, 2000a). These faunas include a number of southern European species, often occurring in abundance (Coope, 1990). Quantitative estimates of the thermal climate of these sites have given the following figures: Tmax 18–24 C, Tmin 6 to 6 C. It is likely that the actual mean July temperatures may have been about 21 C and that mean January/February temperatures may have been about 4 C. Thus, the mean temperature of the warmest month was of the order of 3 C warmer than it is in southern England today. Winter figures are more uncertain, but they may not have been much different from those of the present day. Precipitation at that time was sufficient to maintain a vigorous flow in the rivers throughout the year. The termination of the Eemian Interglacial is difficult to investigate because this stage is apparently missing from all of {
Deceased.
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the British sequences. However, a borehole at Elsing in the valley of the River Wensum, about 22 km west of Norwich, penetrated Eemian deposits immediately overlain by sediments with arthropod faunas indicative of rapidly alternating climates ranging from temperate to arctic conditions (Taylor and Coope, 1985). At the Oerel site, in northwest Germany (Behre et al., 2005), beetle assemblages were recovered from sediments representing the late part of the Eemian interglacial. They were divided into two different faunal units, in which the older is rather species-rich and indicative of temperate conditions, compared with the younger sparser fauna, which lacked thermophiles. MCR estimates based on the older assemblages are: Tmax 14–24 C, Tmin 11 to 6 C. Further southwest, at the La Grande Pile site, Vosges, France, beetle remains were analyzed from ten samples assigned to Eemian deposits (Ponel, 1995). The beetle assemblages were divided into an early and a late faunal unit. The early unit is characterized by an abundance of beetles dependent on deciduous trees, and thermophilous predators and scavengers. MCR estimates based on this assemblage are: Tmax 16–22 C, Tmin 0–10 C. The faunal unit corresponding to the later part of the interglacial includes species feeding on both deciduous and coniferous trees. A number of species indicate a mild and humid climate: Tmax 17–26 C, Tmin 6 to 12 C.
Herning Stadial (MIS 5d) The insect assemblages from the Oerel and Gross Todtshorn sites in Germany (Walking, 1997), attributed to this period, are made up of only a few taxa. At Oerel, a few species confined to a marshy environment were identified. At Gross Todtshorn typical heath taxa were recovered. However, the faunas were too small to permit MCR reconstructions to be made. At La Grande Pile one sample included a number of species living on open ground, such as open grassland. However, the MCR estimates do not indicate arctic conditions during this time: Tmax 15–19 C, Tmin 15 to 9 C.
Brørup Interstadial (MIS 5c) Beetle faunas were described from four sites in central and northern Sweden that were originally thought to be interglacial in age (Lindroth, 1948) but which are now attributed to an early Weichselian interstadial. These faunas included several exclusively high northern and Asiatic species. More recently, Coleoptera and other insect remains were found and identified from deposits of Brørup age (Pera¨pohjola) at ten different sites in northernmost Sweden (Lemdahl, 1997). All the recorded taxa give a very consistent picture of a xeric to mesic tundra environment. A number of the identified species are not found within Fennoscandia today, but in northern Russia and
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Eemian and Weichselian sites
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Figure 1 Map showing Eemian to full-glacial Weichselian sites where fossil beetle assemblages were studied. (1) Early Weichselian sites in northern Sweden, (2) Dimlington, (3) Chelford, (4) Elsing near Norwich, (5) Latton, (6) Cassington near Oxford, (7) South Kensington, (8) Isleworth, (9) Peelo, (10) Oerel, (11) Gross Todtshorn, (12) Belchato´w, (13) La Grande Pile, (14) Gossau, (15) full-glacial site near Verona.
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(b)
Figure 2 (a) The Veiki moraine plateau at Outoja¨rvi, north easternmost Sweden. Here organic silty sediments dating from the Early Weichselian Bro¨rup interstadial (MIS 5c) were found between 1.5 and 5.5 m below the present soil surface. The sediments are covered by 1.5 m Holocene peat. (b) Sampling of the Early Weichselian sediments for plant macrofossil and insect analyses at Outoja¨rvi in an open section made by a dredger, during winter time when the upper meter of the ground was frozen.
Mongolia. MCR estimates indicate cold, harsh conditions: Tmax 9–11 C, Tmin 20 to 9 C (Figure 2(a) and 2(b)). Beetle faunas of this age indicate mixed conifer and birch woodland and are known from several sites in the English Midlands. The most extensively studied is from Chelford, Cheshire (Coope, 1959). MCR analysis of this fauna gives the following figures: Tmax 15–18 C, Tmin 11 to 1 C. It is likely
that the actual figure for Tmax was closer to 15 . Thus, the summer temperatures were similar to those in central England today, but the winters were substantially colder. Results from the Oerel site present two contrasting faunas from this interstadial. The earliest part, of low diversity, included northern species. This was followed by a striking increase in number of species, with terrestrial and aquatic
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beetles that today are found only in temperate regions where they inhabit fens and coniferous woodlands. Similar patterns are found in the results from Gross Todtshorn. MCR estimates from Oerel are as follows: Tmax 12–19 C, Tmin 21 to 6 C. At La Grande Pile the assemblage from this time is characterized by a rich fauna of tree-dependent species living on both deciduous and coniferous trees. Climate conditions were likely to have been less temperate than during the Eemian: Tmax 13–24 C, Tmin 13 to 10 C.
Rederstal Stadial (MIS 5b) and Odderade Interstadial (MIS 5a) Insect assemblages dating to the Odderade Interstadial (Ta¨rendo¨) were recovered from four Swedish sites (Lemdahl, 1997). They are characterized by arctic tundra faunas of low diversity. The climatic conditions seem to have been even more severe than those of the Brørup: Tmax 10–11 C, Tmin 20 to 17 C. There are no beetle faunas from Britain that can be unequivocally attributed to MIS 5b and 5a. The Rederstall assemblages from Oerel and Gross Todtshorn are made up of only a few beetle taxa and indicate the presence of shrub tundra. Tmax estimates from Gross Todtshorn are around 11 C. The Odderade Interstadial is well represented at Oerel where two different insect assemblages reflect two different climate regimes during the period. The first part of the interstadial is characterized by a thermophilus marsh fauna (Tmax 13–25 C, Tmin 8 to 10 C). During the latter part, there is a rather rapid change in which cold stenotherm species replace the temperate fauna (Tmax 13–14 C, Tmin 14 to 10 C). At Gro¨bern in eastern Germany, the beetle succession showed large scale post-Eemian climatic oscillations that probably belong to this period. These involved changes of at least 7 C in mean July temperatures (Walkling and Coope, 1996). Beetle assemblages dating to the Rederstall from La Grande Pile show a fall in the number of tree-dependent taxa and a cold-adapted species was also recorded (Tmax 13–20 C, Tmin 16 to 8 C). During the Odderade, the number of tree-dependent beetles increased again. The climate was probably slightly cooler than during the Brørup (Tmax 12–14 C, Tmin 0–6 C).
Middle Weichselian Stadials and Interstadials (MIS 4) This period was climatically complex. There are at least two separate middle Weichselian temperate interstadials in central and southern England separated from one another by a severely cold interval. Both of these temperate episodes postdate MIS 5 (Coope, 2000b) and yielded pollen indicating a treeless landscape. Beetle assemblages are known from three sites in England that date from the earlier of these two interstadials, namely that from Isleworth (Coope and Angus, 1975), the post interglacial deposits at Cassington (Maddy et al., 1998) and from Latton (Lewis et al., 2006). The interstadial beetle assemblages from these sites are rich and diverse and are made up wholly of temperate–oceanic species. Average MCR analyses based on the beetle faunas from these three sites gave the following estimates: Tmax 17–18 , Tmin 4 to 4 C. At Cassington, these temperate interstadial deposits were overlain by gravels in the middle of which organic silts filled a small paleochannel with a fauna made up of an unusual group
of exclusively arctic and continental beetle species. MCR analysis of the beetles from this deposit yielded the following estimates: Tmax 7–11 C, Tmin 30 to 10 C. An almost identical beetle assemblage was also found at South Kensington, London, suggesting that these two deposits are of the same age (Coope et al., 1997). At South Kensington the level containing this arctic fauna was overlain almost immediately by organic silty gravels which yielded a rich interstadial fauna composed entirely of temperate–oceanic beetle species. MCR climatic calculations of this upper temperate interlude gave the following estimates: Tmax 16–18 C, Tmin 6 to 1 C. This temperate period follows so closely on the arctic bed, both of them occurring within the same small channel deposit within the gravel sequence, that it seems likely that the climatic warming took place very rapidly. However, in the absence of reliable dates, the actual rate of climatic change cannot be determined. It is environmentally significant that, during both these temperate interludes, pollen analysis shows that the landscape in Britain was totally devoid of trees in spite of the climate being warm enough at these times to support mixed oak forest.
Middle Weichselian (MIS 3) Numerous organic deposits are known from England which have yielded beetle assemblages that can be attributed to the Middle Weichselian on stratigraphical grounds (Coope, 1987a,b), but it is not yet possible to allocate them precisely to their equivalent positions in the Greenland ice-cores chronology. Several sites that have been radiocarbon dated to about 42.0 ky BP have yielded beetle assemblages that include relatively southern species indicating warmer climates (Coope et al., 1961; Coope 1968, 2000b, 2002). Because of the unreliability of radiocarbon dates of this age, it is not possible at the moment to be sure if these isolated deposits represent just one warm episode; or several of them (i.e., the Upton Warren Interstadial Complex of Coope and Sands (1966)). A large number of beetle faunas from deposits with radiocarbon dates between 40.0 ky BP and 20.0 ky BP have remarkably consistent assemblages and include numerous species with exclusively arctic and/or Asiatic distributions today (Coope, 1995). Many of these exotic species are represented by large numbers of individuals. MCR estimates have been made on over 20 of these faunas (Coope, 1987b) and an approximate average may be summarized as follows: Tmax 9–11 C, Tmin 25 to 10 C. It is possible that these faunas represent several cold interstadials interspersed with even colder intervals as the Greenland Ice cores suggest (Johnsen et al., 1992) in which case biological activity in Britain may have fallen to a paleontologically invisible minimum. Although Scotland was totally overridden by ice sheets at the Last Glacial Maximum (LGM), insect-bearing sediments are occasionally found underneath the veneer of glacial deposits. Notable among these is the prolific site at Sourlie, near Irvine, western Scotland, which yielded a rich assortment of plant and animal fossils, including reindeer and wooly rhinoceros as well as abundant insect remains. Radiocarbon dates on plant remains and the collagen from a reindeer antler found in situ grouped around 30.0 ky BP (Bos et al., 2004). MCR paleoclimatic estimates based on the beetle fauna gave mean July temperatures of between 6 and 9 C and mean January/
BEETLE RECORDS | Late Pleistocene of Europe February temperatures of 34º and 11 C. A more recent study of a glaciotectonized organic deposit at Balglass Burn, high on the slopes of the Campsie Fells east of Glasgow, gave ages between 34.5 and 28.0 ky BP based on six radiocarbon dates on sedge fruits and beetle remains (Brown et al., 2007). Besides insects, pollen and plant macrofossil remains were also studied. All these indicate a small pond situated in an open tundra landscape. MCR estimates based on the beetle assemblage give Tmax 8–10 C and Tmin 26 to 10 C. The climatic implications of these beetle faunas demonstrate that much of central Scotland was not covered by glaciers during the middle pleniglacial at a time when thermal conditions were apparently sufficiently cold to permit ice accumulation in the highlands. Some other factor such as inadequate precipitation must also have been involved in maintaining ice-free conditions in northern Britain at this time. In the Netherlands pleniglacial, two extensive Middle Weichselian insect assemblages are known. Thus, a beetle fauna from Peelo included many of the exclusively arctic and/or Asiatic species that were also present in the British Isles at the same time (Coope, 1969). Excavations at Orvelte, near Drentse, to recover mammoth and wooly rhinoceros remains, enabled a large beetle fauna to be investigated that included an assemblage of cold-adapted species similar to the ones discussed above (Cappers et al., 1993). Both of these faunas yield MCR Tmax temperature estimates at or below 10 C and Tmin values below 20 C. A Middle Weichselian insect fauna from Poland included a very similar group of high arctic and Asiatic species (Kasse et al., 1998). MCR estimations based on this assemblage gave the following estimates: Tmax 8–12 C, Tmin 27 to 20 C. Insect remains were found in deposits referring to the Oerel interstadial both at the Oerel site and at Gross Todtshorn. The beetle assemblages are composed of northern Palaearctic species, which indicate a treeless environment and cold climate (Tmax 12–13 C, Tmin 9 to 2 C). At the Oerel site, insect remains were also recovered from a peat layer referred to the younger Glinde Interstadial. The beetle assemblage from this horizon is similar in composition to those from the earlier interstadial. Climate conditions were also likely to have been very similar (Tmax 9–14º, Tmin 17 to 2 C). In Switzerland, two insect assemblages of middle Weichselian age have been recorded from Gossau (Jost-Stauffer et al., 2005). They include several exclusively arctic and Asiatic species that do not occur anywhere in central Europe at the present day. MCR estimates gave the following figures: Tmax 8–13 C, Tmin 21 to 7 C. A second beetle assemblage from the Swiss plateau was obtained from samples taken in a peaty matrix of the mammoth skeleton at Niederweningen that was radiocarbon dated to around 45.0 ky BP (Coope, 2007). This fauna also includes many exclusively arctic/Asiatic species. The mammoths were probably mired in a reedy swamp in a landscape with scattered willows, birches, and conifers. An MCR estimate indicates Tmax around 10 C and Tmin below 10 C. Middle Weichselian beetle assemblages from La Grande Pile are characterized by the presence of almost entirely arctic and continental species. Obligate tree-dependent species were not recorded. The climate was cold in general through the period (Tmax around 12 C, Tmin approximately 10 C or below). A site of outstanding archaeological importance has recently been recorded at Lynford, Norfolk, where an extensive
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beetle fauna was found in close association with abundant evidence of Neanderthal exploitation of mammoths: smashed bones and numerous flint tools (Boismier et al., 2011). There can be little doubt that the beetle assemblage is contemporary with the human activities and provides an intimate view of the local environment (Coope, 2011). However, the beetle assemblage was unusual. It included abundant high northern and Asiatic species suggesting cold continental climatic conditions. But this fauna lacks several of the extreme cold-adapted species present in the typical full-glacial assemblages and includes rare species whose distributions do not extend into arctic regions today. These make MCR estimates difficult. Nevertheless, it is likely that the mean July temperatures were between 12 and 14 C and January/February figures at or below 10 C. The age of this important site is probably sometime late in MIS 4 or early MIS 3.
Last Glacial Maximum (MIS 2) Conditions during the LGM were extremely harsh and insect faunas from this period are consequently very rare. A small beetle assemblage obtained from Dimlington, East Yorkshire, has been radiocarbon dated to 18.5 ky BP (Penny et al., 1969). Given the smallness of this insect fauna, only a provisional MCR estimate is possible: Tmax 9–11 C, Tmin 27 to 7 C. These figures do not necessarily represent the typical climate at the LGM for England because the insects came from a single fossiliferous horizon in an otherwise barren sequence of laminated silts and may thus indicate a short period of atypically warm conditions in an otherwise even colder episode. In continental Europe, two beetle assemblages are known from the LGM. Beetle assemblages from La Grande Pile dating to this period indicate very cold and harsh conditions, with Tmax mostly around 10 C and Tmin well below 10 C. Fossil remains of beetles and oribatid mites, which are dated to around 18 800 BP, were recovered from a site close to Verona, northeastern Italy (Foddai and Minelli, 1994). They indicate climate conditions colder and wetter than today. Tmax may have been 8–9 C lower than present day.
Late Glacial Period This period falls well within the radiocarbon dating techniques. Dates are cited here in radiocarbon years BP and calibrated calendar years are shown in parentheses as cal years BP. Correlations are made with the climatic event stratigraphy of the Greenland Ice cores (Walker et al., 1999). As a large number of studies have been carried out on beetle assemblages from sites scattered all over Europe (cf. Coope et al., 1998), only a representative selection is discussed here (Figure 3).
Late Glacial stadial (Greenland stadial 2) Faunas that date from the cold period (Greenland stadial 2, GS-2) that immediately preceded the Late Glacial Interstadial, namely from just before 13.0 ky BP (ca. 15.4 cal ky BP), are rare. In Britain, the most stratigraphically secure faunal assemblage of this age is from the North Wales coast at Glanllynnau, where a sequence of insect-bearing deposits spans almost the whole of the early part of the Late Glacial Interstadial (Coope and Brophy, 1972, cited in Coope et al., 1998). The stadial
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Late Glacial sites
Figure 3 Map showing Late Glacial sites where fossil beetle assemblages were studied.
deposits yielded exclusively arctic and Siberian species. MCR calculations on this assemblage gave the following estimates: Tmax 10–11 C, Tmin 26 to 23 C. A second beetle assemblage of this age was associated with the recovery of the skeletons of an adult mammoth plus at least three juveniles from silty clay in a kettle hole at Condover, near Shrewsbury (Allen et al., 2009). The GS-2 stadial beetle fauna included many exclusively high arctic species, but unfortunately the wallowing of the mammoths in their efforts to extricate themselves from the adhesive clay had churned up the sediment, incorporating some of the more recent Late Glacial sediment into the stadial deposits, making the precise interpretation of the environment difficult. It is, however, clear that the stadial assemblage of beetles lacked the eastern Asiatic component so characteristic of the full-glacial faunas, suggesting that although the climate must have been of arctic severity at this time, it was probably less continental than during the mid-glacial (pleniglacial) episode. Only two beetle assemblages of this age are known from northern Europe. Both of these faunas are from Kullaberg, southwestern Sweden, and can be dated with confidence to this period (Lemdahl, 1988, cited in Coope et al., 1998). Although these include only a few species, they resemble those from Britain. Similarly, cold-adapted suites of beetle species are also known from Poland, and from the Swiss Plateau (Coope and Elias, 2000; Coope and Lemdahl, 2009; Elias and Wilkinson, 1983; Gaillard and Lemdahl, 1994; Hardon et al., 2000). MCR estimates gave the following figures: Tmax 10–13 C, Tmin 17 to 0 C.
Late Glacial interstadial Beetle faunas from the Late Glacial interstadial 1 (GI-1) are widespread across the whole of northwestern Europe. In Britain, important faunas have been described from Glanllynnau, North
Wales (Coope and Brophy, 1972, cited in Coope et al., 1998), St Bees, Cumbria (Coope and Joachim, 1980, cited in Coope et al., 1998), Gransmoor, East Yorkshire (Walker et al., 1993, cited in Coope et al., 1998), Hollywell Coombe, Kent (Coope, 1998) and from Llanilid, South Wales (Walker et al., 2003). Faunas from the early part of the GI-1e interstadial (the Bølling phase) from about 12.8 ky BP (15.1 cal ky BP) or slightly earlier, include an ecologically diverse variety of relatively southern species. At this time the vegetation was thin and made up chiefly of pioneer plant species. The only trees that were present were birches and willows and these were probably confined to the damp valley bottoms. At this time northern Europe as a whole was largely without a woodland cover in spite of the fact that for much of the region, the climate was warm enough to have supported mixed oak forest. Edaphic factors are thought to have been largely responsible for this open landscape. Average MCR estimates from these sites give approximate figures: Tmax 17–18 C, Tmin 7 to 1 C. Beetle faunas indicate that the initial warming at the start of the interstadial involved a rapid rise in mean July temperatures of at least 7 C and a corresponding rise in winter temperatures substantially greater than this. The rate at which the mean July temperature rose was about 1 C per decade and even higher (Coope and Brophy, 1972, cited in Coope et al., 1998). Because this change also involved a switch from continental to oceanic climatic conditions, the rise in mean annual temperature was considerably greater than these figures suggest, though the precise number of degrees involved is impossible to estimate with confidence. The beetle reconstruction that yielded this unexpectedly high rate of climatic change has subsequently been supported by evidence from the Greenland Ice cores, which show a dramatic climatic amelioration at about this time (Johnsen et al., 1992). It is interesting to observe that this episode of rapid climate warming at the start of the Late Glacial Interstadial is not recorded by beetle assemblages from southern Sweden (Lemdahl, 1988, cited in Coope et al., 1998) or from the coast of western Norway. During this period, the Scandinavian beetle assemblages are of an arctic or boreal character. It is likely that this reflects the local cooling effect provided by the close proximity of the waning Fennoscandian ice sheet. In contrast to the Scandinavian situation, faunas from Poland and the Netherlands (Geel et al., 1989, cited in Coope et al., 1998) indicate temperatures similar to those of the British Isles. At about the same time, warmth demanding beetle faunas are recorded from the Swiss Plateau (Coope and Elias, 2000; Coope and Lemdahl, 2009; Elias and Wilkinson, 1983; Gaillard and Lemdahl, 1994; Hardon et al., 2000). Similar faunal changes are traced in southern France (Ponel and Coope, 1990; Ponel et al., 2001). Shortly before 12.0 ky BP (ca. 14.0 cal ky BP), there was a sudden climatic cooling. Tmax figures suddenly fell to around 15 C or even lower and never recovered again during the rest of the interstadial; in other words for much of Western Europe the whole of the Allerød phase was substantially cooler than the preceding Bølling phase. The beetles indicate that during the Allerød there appears to have been several relatively minor climatic oscillations, well illustrated by the climatic curves from Gransmoor (Walker et al., 1993, cited in Coope et al., 1998) and Llanilid (Walker et al., 2003, and in summary by
BEETLE RECORDS | Late Pleistocene of Europe
Figure 4 Fieldwork at the small marsh (80 30 m) at He´re´mence situated at 2290 m.a.s.l. in the Swiss Alps. Four sites at different altitudes, including this site, were studied in a multiproxy project in order to record biotic responses to rapid climatic changes around the Younger Dryas in the Swiss Alps. Samples were recovered by using a Streif-piston corer of 8 cm diameter.
Lowe and Walker, 1997, cited in Coope et al., 1998). The frequency of these oscillations makes it difficult to give useful MCR estimates for the latter half of the Late Glacial Interstadial. Again, in contrast to the British Isles and much of Europe, the beetle assemblages from southern Sweden indicate that the highest temperatures were reached during the latter part of the Late Glacial Interstadial. However, it must be emphasized that the actual MCR temperature estimates were similar to the British figures at this time. In Norway arctic faunas are recorded for the Allerød phase (Birks et al., 1993, cited in Coope et al., 1998; Lemdahl, 2000a). In Poland species with a boreal distribution appear during this period (Lemdahl, 1991a, cited in Coope et al., 1998), which also seems to be the case in southern France. These beetle assemblages show that during the GI-1 there were distinct regional differences in climate and steep temperature gradients across much of northern Europe (Coope and Lemdahl, 1995; Coope et al., 1998).
Late Glacial stadial (GS-1) ¼ Younger Dryas The beetle evidence from the British Isles suggests that the beginning of the GS-1 (Younger Dryas) cold period appears as a culmination of the sequence of Allerød cooling events outlined above. Its beginning is conveniently dated at about 11.0 ky BP (ca. 13.0 cal ky BP). Many beetle assemblages are known from the Younger Dryas period (e.g., Coope et al., 1979, Coope and Joachim, 1980, cited in Coope et al., 1998; Coope, 1998; Walker et al., 1993, 2003) and include a large number of exclusively arctic species, many of which were previously encountered in the assemblages from full-glacial times. However, exclusively Asiatic species are very rare at this time. An average of MCR estimates based on faunas from this period indicates the following figures: Tmax 9–11 C, Tmin 10 to 20 C (Figures 4 and 5). In northern France, insect assemblages from Younger Dryas fluvial deposits recorded at St.-Momelin, yielded these MCR estimates: Tmax around 10 C and Tmin close to 12 C (Ponel
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Figure 5 Fossil head and thorax matching a modern specimen of the carrion beetle Thanatophilus dispar. Remains of this species were abundant in lake sediments dating to the Oldest Dryas (GS-2) at the site Hauterive Rouges-Terres, at the shore of the Lake Neuchaˆtel, Switzerland. A Paleolithic settlement was excavated adjacent to the site. Today T. dispar is mainly found on fish carrion where it probably feeds on fly larvae.
et al., 2007). MCR reconstructions show that, in contrast to the Late Glacial Interstadial, relatively similar climatic conditions seem to have prevailed in most of northern Europe at this time. In eastern Finland high arctic beetle faunas of Younger Dryas and early post glacial age colonized recently deglaciated areas (Bondestam et al., 1994). Although Younger Dryas beetle assemblages are very rare from the Swiss Plateau, they suggest cold, arctic conditions (Lemdahl, 2000b). Many sites that include the transition from the Late Glacial to the Holocene show signs of a hiatus, making it difficult to estimate the rate of change involved. However, all the available beetle evidence suggests that the change was abrupt and intense and probably similar in magnitude to the one at the start of the Late Glacial Interstadial (e.g., Ashworth, 1973; Lemdahl, 1991b, cited in Coope et al., 1998; Lemdahl, 2000; Ponel et al., 2007). The beetles indicate that throughout most of Europe at the start of the Holocene, temperatures reached present day levels in a geologically immeasurably short period of time.
See also: Beetle Records: Middle Pleistocene of Western Europe; Overview; Postglacial Europe.
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and coleopteran investigations. Palaeogeography, Palaeoclimatology, Palaeoecology 204: 65–100. Brown EJ, Rose J, Coope RG, and Lowe JJ (2007) An MIS 3 age organic deposit from Balglass Burn, central Scotland: Palaeoenvironmental significance and implications for the timing of the onset of the LGM ice sheet in the vicinity of the British Isles. Journal of Quaternary Science 22: 295–308. Cappers RTJ, Bosch JHA, Bottema S, et al. (1993) De reconstructive van het landschap. In: Sanden WAB, Cappers RTJ, Beuker JR, and Mol D (eds.) Mens en Mammoet, Archaeologische Monografieen van Drentes Museum Deel 5, pp. 27–41. Assen: Drentes Museum. Coope GR (1959) A Late Pleistocene insect fauna from Chelford, Cheshire. Proceedings of the Royal Society of London B151: 70–86. Coope GR (1968) An insect fauna from Mid-Weichselian deposits at Brandon, Warwickshire. Philosophical Transactions of the Royal Society of London B254: 425–456. Coope GR (1969) Insect remains from Mid Weichselian Deposits at Peelo, The Netherlands. Mededelingen Rijks Geologische Dienst 20: 79–83. Coope GR (1987a) The response of late Quaternary insect communities to sudden climatic changes. In: Gee JHR and Giller PS (eds.) Organisation of Communities – Past and Present, pp. 421–438. Oxford: Blackwell Scientific. Coope GR (1987b) Fossil beetle assemblages as evidence of sudden and intense climatic changes in the British Isles during the last 45,000 years. In: Berger WH and Labeyrie LD (eds.) Abrupt Climatic Change, Environment and Implications, pp. 147–150. Dordrecht: Reidel. Coope GR (1990) The invasion of Northern Europe during the Pleistocene by Mediterranean species of Coleoptera. In: di Castri F, Hansen AJ, and Debussche M (eds.) Biological Invasions in Europe and the Mediterranean Basin, pp. 203–215. Dordrecht: Kluwer. Coope GR (1995) Insect faunas in ice age environments: Why so little extinction? In: Lawton JH and May RM (eds.) Extinction Rates, pp. 55–74. Oxford: Oxford University Press. Coope GR (1998) Insects. In: Preec RC and Bridgland D (eds.) Late Quaternary Environmental Change in North-west Europe: Excavations at Holywell Coombe, South-East England, pp. 213–232. London: Chapman & Hall. Coope GR (2000a) The climatic significance of coleopteran assemblages from the Eemian deposits of southern England. Geologie en Mijnbouw 79: 257–267. Coope GR (2000b) Middle Devensian (Weichselian) coleopteran assemblages from Earith, Cambridgeshire (UK) and their bering on the interpretation of a ‘Full glacial’ floras and faunas. Journal of Quaternary Science 15: 779–788. Coope GR (2002) Changes in the thermal climate in Northwestern Europe during marine oxygen isotope stage 3, estimated from fossil insect assemblages. Quaternary Research 57: 401–408. Coope GR (2007) Coleoptera from the 2003 excavations of the mammoth skeleton at Niederweningen, Switzerland. Quaternary International 164–165: 130–138. Coope GR (2011) The insect remains from the mammoth channel at Lynford. In: Boismier WA, Gamble CS, and Coward F (eds.) Neanderthals Amongst Mammoths: Excavations at Lynford Quarry, Norfolk, UK. London: English Heritage. Coope GR and Angus RB (1975) An ecological study of a temperate interlude in the middle of the last glaciation, based on fossil Coleoptera from Isleworth, Middlesex. Journal of Animal Ecology 44: 365–391. Coope GR, Dickson JH, McCutcheon JA, and Mitchell GF (1979) The lateglacial and early postglacial deposit at Drumurcher, Co. Monaghan. Proceedings of the Royal Irish Academy B79: 63–85. Coope GR and Elias SA (2000) The environment of Upper Palaeolithic (Magdalenian and Azilian) hunters at Hauterive-Champre´veyres, Neuchaˆtel, Switzerland, interpreted from coleopteran remains. Journal of Quaternary Science 15: 157–175. Coope GR, Gibbard PL, Hall AR, Preece RC, Robinson JE, and Sutcliffe AJ (1997) Climatic and environmental reconstructions based on fossil assemblages from Middle Devensian (Weichselian) deposits of the River Thames at South Kensington, Central London UK. Quaternary Science Reviews 6: 1163–1195. Coope GR and Lemdahl G (1995) Regional differences in the Lateglacial climate of northern Europe based on coleopteran analysis. Journal of Quaternary Science 10: 391–395. Coope GR and Lemdahl G (2009) Insect analyses. In: Thew N, Hadorn P, and Coope GR (eds.) Hauterive/Rouges-Terres. Reconstruction of Upper Palaeolithic and Early Mesolithic Natural Environments, pp. 125–159. Arche´ologie neuchaˆteloise 44. Coope GR, Lemdahl G, Lowe JJ, and Walkling AP (1998) Temperature gradients in northern Europe during the last glacial–interglacial transition (14–9 14C kyr BP) interpreted from coleopteran assemblages. Journal of Quaternary Science 13: 419–433. Coope GR and Sands CHS (1966) Insect faunas of the last glaciations from the Tame Valley, Warwickshire. Proceedings of the Royal Society B 165: 389–412. Coope GR, Shotton FW, and Strachan I (1961) A Late Pleistocene fauna and flora from Upton Warren, Worcestershire. Philosophical Transactions of the Royal Society of London B244: 379–421.
Elias SA and Wilkinson B (1983) Lateglacial insect fossil assemblages from Lobsigensee (Swiss Plateau). Studies in the Late Quaternary of Lobsigensee 3. Revue de Pale´obiologie 2: 189–204. Foddai D and Minelli A (1994) Fossil arthropods from a full-glacial site in northeastern Italy. Quaternary Research 41: 336–342. Gaillard M-J and Lemdahl G (1994) Lateglacial insect assemblages from Grand-Marais, South-western Switzerland – Climatic implications and comparison with pollen and plant macrofossil data. Dissertationes Botaniceae 234: 287–308. Hardon P, Thew N, Coope GR, Lemdahl G, Hajdas I, and Bonani G (2000) A Late-Glacial and Early Holocene environment and climate history for the Neuchaˆtel region (CH). In: Richard H and Vignot A (eds.) E´quilibres et ruptures dans les ecosystems Durant les 20 derniers mille´naires en Europe de l’Ouest, Actes du colloque international de Besanc¸on, septembre 2000, pp. 75–90. Besanc¸on: Presses Universitaires Franc-Comtoises. Johnsen SJ, Clausen HB, Dansgaard W, et al. (1992) Irregular glacial interstadials recorded in a new Greenland Ice core. Nature (London) 359: 311–313. Jost-Stauffer M, Coope GR, and Schlu¨chter C (2005) Environmental and climatic reconstructions during Marine Oxygen Isotope Stage 3 from Gossau, Swiss Midlands, based on coleopteran assemblages. Boreas 34: 53–60. Kasse C, Huitzer AS, Krzyszkowski D, Bohncke SJP, and Coope GR (1998) Weichselian Late-Pleniglacial and Late-glacial depositional environment, Coleoptera and periglacial climate records from central Poland (Belchato´w). Journal of Quaternary Science 13: 455–469. Lemdahl G (1997) Early Weichselian insect faunas from northern Sweden: Climatic and environmental implications. Arctic and Alpine Research 29: 63–74. Lemdahl G (2000a) Late-glacial and Early Holocene Coleoptera assemblages as indicators of local environment and climate at Kra˚kenes Lake, western Norway. Journal of Paleolimnology 23: 57–66. Lemdahl G (2000b) Lateglacial and Early Holocene insect assemblages from sites at different altitudes in the Swiss Alps – implications on climate and environment. Palaeogeography, Palaeoclimatology, Palaeoecology 159: 293–312. Lewis SG, Maddy D, Buckingham C, et al. (2006) Pleistocene fluvial sediments, paleontology and archaeology of the upper River Thames at Latton, Wiltshire, England. Journal of Quaternary Science 21: 181–205. Lindroth CH (1948) Interglacial insect remains from Sweden. A˚rsbok Sveriges Geologiska Underso¨kning C42: 1–29. Maddy D, Lewis SG, Scaife RG, et al. (1998) The Upper Pleistocene deposits at Cassington, near Oxford, England. Journal of Quaternary Science 13: 205–231. Penny LF, Coope GR, and Catt JA (1969) Age and insect fauna of the Dimlington Silts, East Yorkshire. Nature 224: 65–67. Ponel P (1995) Rissian, Eemian and Wu¨rmian Coleoptera assemblages from La Grande Pile (Vosges, France). Palaeogeography, Palaeoclimatology, Palaeoecology 114: 1–41. Ponel P and Coope GR (1990) Lateglacial and early Flandrian from La Taphanel, Massif Central, France: climatic and ecological implications. Journal of Quaternary Science 5: 235–249. Ponel P, Andrieu-Ponel V, Parchoux F, Juhasz I, and de Beaulieu J-L (2001) Late-glacial and Holocene high-altitude environmental changes in Valle´e des Merveilles (Alpes Maritimes, France): Insect evidence. Journal of Quaternary Science 16: 795–812. Ponel P, Gandouin E, Coope GR, et al. (2007) Insect evidence for environmental and climate changes from Younger Dryas to Sub-Boreal in a river floodplain at St-Momelin (St-Omer basin, northern France), Coleoptera and Trichoptera. Palaeogeography, Palaeoclimatology, Palaeoecology 245: 483–504. Taylor BJ and Coope GR (1985) Arthropods in the Quaternary of East Anglia – Their role as indices of local Palaeoenvironments and regional Palaeoclimates. Modern Geology 9: 159–185. Walker MJC, Bjo¨rck S, Lowe JJ, et al. (1999) Isotopic ‘events’ in the GRIP ice-core: A stratotype for the Late Pleistocene. Quaternary Science Reviews 18: 1143–1150. Walker MJC, Coope GR, and Lowe JJ (1993) The Devensian (Weichselian) Lateglacial palaeoenvironmental record from Gransmoor, East Yorkshire, England. Quaternary Science Reviews 12: 659–680. Walker MJC, Coope GR, Sheldrick C, et al. (2003) Devensian Lateglacial environmental changes in Britain: A multi-proxy environmental record from Llanilid, South Wales, UK. Quaternary Science Reviews 22: 475–520. Walking AP (1997) Ka¨ferkundliche Untersuchungen an weiselzeitlichen Ablagerungen der Bohrung Gross Todtshorn (Kr. Harburg; Niedersachsen). Schriftenreihe derDeutschen Geologischen Gesellschaft 4: 87–102. Walkling AP and Coope GR (1996) Climatic reconstructions from the Eemian/early Weichselian transition in Central Europe based on the coleopteran record from Go¨bern, Germany. Boreas 25: 145–159.
Introduction In this article, the response of Coleoptera (beetles) to Upper Pleistocene climatic changes are summarized. This should cast light upon our understanding of the nature of these changes and thus enable us to predict the ways in which other animal and plant species might respond to any future climatic events. By using the changes in the range of beetle species during the Quaternary, it has been possible to quantify thermal climatic conditions and to estimate both the amounts and the rates of change involved. Furthermore, it has been possible to map the regional differences in thermal climates within northwestern Europe. The beetle assemblages described here include those from the Last (Eemian, Ipswichian) Interglacial, the Last (Weichselian, Devensian) Glaciation, and the transition to the present (Holocene) Interglacial. Most of the examples have been drawn from sites in the British Isles and northern Europe because it is there that these faunas have been most intensively investigated. The beetle record shows that the climate oscillated rapidly between temperate–oceanic and arctic–continental conditions. Where possible, the fossil insect assemblages have been correlated with marine isotope stages (MIS) or with the event stratigraphy of the Greenland ice cores. In the account that follows, paleotemperature estimates are given using the mutual climatic range (MCR) method (Atkinson et al., 1987) where Tmax indicates the mean temperature of the warmest month (July) and Tmin indicates the mean temperature of the coldest months (January and February; Figure 1).
Upper Pleistocene Beetle Assemblages Eemian Interglacial (MIS 5e) Extensive beetle assemblages of this age are known from seven localities in southeast England (Coope, 2000a). These faunas include a number of southern European species, often occurring in abundance (Coope, 1990). Quantitative estimates of the thermal climate of these sites have given the following figures: Tmax 18–24 C, Tmin 6 to 6 C. It is likely that the actual mean July temperatures may have been about 21 C and that mean January/February temperatures may have been about 4 C. Thus, the mean temperature of the warmest month was of the order of 3 C warmer than it is in southern England today. Winter figures are more uncertain, but they may not have been much different from those of the present day. Precipitation at that time was sufficient to maintain a vigorous flow in the rivers throughout the year. The termination of the Eemian Interglacial is difficult to investigate because this stage is apparently missing from all of {
Deceased.
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the British sequences. However, a borehole at Elsing in the valley of the River Wensum, about 22 km west of Norwich, penetrated Eemian deposits immediately overlain by sediments with arthropod faunas indicative of rapidly alternating climates ranging from temperate to arctic conditions (Taylor and Coope, 1985). At the Oerel site, in northwest Germany (Behre et al., 2005), beetle assemblages were recovered from sediments representing the late part of the Eemian interglacial. They were divided into two different faunal units, in which the older is rather species-rich and indicative of temperate conditions, compared with the younger sparser fauna, which lacked thermophiles. MCR estimates based on the older assemblages are: Tmax 14–24 C, Tmin 11 to 6 C. Further southwest, at the La Grande Pile site, Vosges, France, beetle remains were analyzed from ten samples assigned to Eemian deposits (Ponel, 1995). The beetle assemblages were divided into an early and a late faunal unit. The early unit is characterized by an abundance of beetles dependent on deciduous trees, and thermophilous predators and scavengers. MCR estimates based on this assemblage are: Tmax 16–22 C, Tmin 0–10 C. The faunal unit corresponding to the later part of the interglacial includes species feeding on both deciduous and coniferous trees. A number of species indicate a mild and humid climate: Tmax 17–26 C, Tmin 6 to 12 C.
Herning Stadial (MIS 5d) The insect assemblages from the Oerel and Gross Todtshorn sites in Germany (Walking, 1997), attributed to this period, are made up of only a few taxa. At Oerel, a few species confined to a marshy environment were identified. At Gross Todtshorn typical heath taxa were recovered. However, the faunas were too small to permit MCR reconstructions to be made. At La Grande Pile one sample included a number of species living on open ground, such as open grassland. However, the MCR estimates do not indicate arctic conditions during this time: Tmax 15–19 C, Tmin 15 to 9 C.
Brørup Interstadial (MIS 5c) Beetle faunas were described from four sites in central and northern Sweden that were originally thought to be interglacial in age (Lindroth, 1948) but which are now attributed to an early Weichselian interstadial. These faunas included several exclusively high northern and Asiatic species. More recently, Coleoptera and other insect remains were found and identified from deposits of Brørup age (Pera¨pohjola) at ten different sites in northernmost Sweden (Lemdahl, 1997). All the recorded taxa give a very consistent picture of a xeric to mesic tundra environment. A number of the identified species are not found within Fennoscandia today, but in northern Russia and
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Eemian and Weichselian sites
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3
2
6 7,8
9 4
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Figure 1 Map showing Eemian to full-glacial Weichselian sites where fossil beetle assemblages were studied. (1) Early Weichselian sites in northern Sweden, (2) Dimlington, (3) Chelford, (4) Elsing near Norwich, (5) Latton, (6) Cassington near Oxford, (7) South Kensington, (8) Isleworth, (9) Peelo, (10) Oerel, (11) Gross Todtshorn, (12) Belchato´w, (13) La Grande Pile, (14) Gossau, (15) full-glacial site near Verona.
(a)
(b)
Figure 2 (a) The Veiki moraine plateau at Outoja¨rvi, north easternmost Sweden. Here organic silty sediments dating from the Early Weichselian Bro¨rup interstadial (MIS 5c) were found between 1.5 and 5.5 m below the present soil surface. The sediments are covered by 1.5 m Holocene peat. (b) Sampling of the Early Weichselian sediments for plant macrofossil and insect analyses at Outoja¨rvi in an open section made by a dredger, during winter time when the upper meter of the ground was frozen.
Mongolia. MCR estimates indicate cold, harsh conditions: Tmax 9–11 C, Tmin 20 to 9 C (Figure 2(a) and 2(b)). Beetle faunas of this age indicate mixed conifer and birch woodland and are known from several sites in the English Midlands. The most extensively studied is from Chelford, Cheshire (Coope, 1959). MCR analysis of this fauna gives the following figures: Tmax 15–18 C, Tmin 11 to 1 C. It is likely
that the actual figure for Tmax was closer to 15 . Thus, the summer temperatures were similar to those in central England today, but the winters were substantially colder. Results from the Oerel site present two contrasting faunas from this interstadial. The earliest part, of low diversity, included northern species. This was followed by a striking increase in number of species, with terrestrial and aquatic
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beetles that today are found only in temperate regions where they inhabit fens and coniferous woodlands. Similar patterns are found in the results from Gross Todtshorn. MCR estimates from Oerel are as follows: Tmax 12–19 C, Tmin 21 to 6 C. At La Grande Pile the assemblage from this time is characterized by a rich fauna of tree-dependent species living on both deciduous and coniferous trees. Climate conditions were likely to have been less temperate than during the Eemian: Tmax 13–24 C, Tmin 13 to 10 C.
Rederstal Stadial (MIS 5b) and Odderade Interstadial (MIS 5a) Insect assemblages dating to the Odderade Interstadial (Ta¨rendo¨) were recovered from four Swedish sites (Lemdahl, 1997). They are characterized by arctic tundra faunas of low diversity. The climatic conditions seem to have been even more severe than those of the Brørup: Tmax 10–11 C, Tmin 20 to 17 C. There are no beetle faunas from Britain that can be unequivocally attributed to MIS 5b and 5a. The Rederstall assemblages from Oerel and Gross Todtshorn are made up of only a few beetle taxa and indicate the presence of shrub tundra. Tmax estimates from Gross Todtshorn are around 11 C. The Odderade Interstadial is well represented at Oerel where two different insect assemblages reflect two different climate regimes during the period. The first part of the interstadial is characterized by a thermophilus marsh fauna (Tmax 13–25 C, Tmin 8 to 10 C). During the latter part, there is a rather rapid change in which cold stenotherm species replace the temperate fauna (Tmax 13–14 C, Tmin 14 to 10 C). At Gro¨bern in eastern Germany, the beetle succession showed large scale post-Eemian climatic oscillations that probably belong to this period. These involved changes of at least 7 C in mean July temperatures (Walkling and Coope, 1996). Beetle assemblages dating to the Rederstall from La Grande Pile show a fall in the number of tree-dependent taxa and a cold-adapted species was also recorded (Tmax 13–20 C, Tmin 16 to 8 C). During the Odderade, the number of tree-dependent beetles increased again. The climate was probably slightly cooler than during the Brørup (Tmax 12–14 C, Tmin 0–6 C).
Middle Weichselian Stadials and Interstadials (MIS 4) This period was climatically complex. There are at least two separate middle Weichselian temperate interstadials in central and southern England separated from one another by a severely cold interval. Both of these temperate episodes postdate MIS 5 (Coope, 2000b) and yielded pollen indicating a treeless landscape. Beetle assemblages are known from three sites in England that date from the earlier of these two interstadials, namely that from Isleworth (Coope and Angus, 1975), the post interglacial deposits at Cassington (Maddy et al., 1998) and from Latton (Lewis et al., 2006). The interstadial beetle assemblages from these sites are rich and diverse and are made up wholly of temperate–oceanic species. Average MCR analyses based on the beetle faunas from these three sites gave the following estimates: Tmax 17–18 , Tmin 4 to 4 C. At Cassington, these temperate interstadial deposits were overlain by gravels in the middle of which organic silts filled a small paleochannel with a fauna made up of an unusual group
of exclusively arctic and continental beetle species. MCR analysis of the beetles from this deposit yielded the following estimates: Tmax 7–11 C, Tmin 30 to 10 C. An almost identical beetle assemblage was also found at South Kensington, London, suggesting that these two deposits are of the same age (Coope et al., 1997). At South Kensington the level containing this arctic fauna was overlain almost immediately by organic silty gravels which yielded a rich interstadial fauna composed entirely of temperate–oceanic beetle species. MCR climatic calculations of this upper temperate interlude gave the following estimates: Tmax 16–18 C, Tmin 6 to 1 C. This temperate period follows so closely on the arctic bed, both of them occurring within the same small channel deposit within the gravel sequence, that it seems likely that the climatic warming took place very rapidly. However, in the absence of reliable dates, the actual rate of climatic change cannot be determined. It is environmentally significant that, during both these temperate interludes, pollen analysis shows that the landscape in Britain was totally devoid of trees in spite of the climate being warm enough at these times to support mixed oak forest.
Middle Weichselian (MIS 3) Numerous organic deposits are known from England which have yielded beetle assemblages that can be attributed to the Middle Weichselian on stratigraphical grounds (Coope, 1987a,b), but it is not yet possible to allocate them precisely to their equivalent positions in the Greenland ice-cores chronology. Several sites that have been radiocarbon dated to about 42.0 ky BP have yielded beetle assemblages that include relatively southern species indicating warmer climates (Coope et al., 1961; Coope 1968, 2000b, 2002). Because of the unreliability of radiocarbon dates of this age, it is not possible at the moment to be sure if these isolated deposits represent just one warm episode; or several of them (i.e., the Upton Warren Interstadial Complex of Coope and Sands (1966)). A large number of beetle faunas from deposits with radiocarbon dates between 40.0 ky BP and 20.0 ky BP have remarkably consistent assemblages and include numerous species with exclusively arctic and/or Asiatic distributions today (Coope, 1995). Many of these exotic species are represented by large numbers of individuals. MCR estimates have been made on over 20 of these faunas (Coope, 1987b) and an approximate average may be summarized as follows: Tmax 9–11 C, Tmin 25 to 10 C. It is possible that these faunas represent several cold interstadials interspersed with even colder intervals as the Greenland Ice cores suggest (Johnsen et al., 1992) in which case biological activity in Britain may have fallen to a paleontologically invisible minimum. Although Scotland was totally overridden by ice sheets at the Last Glacial Maximum (LGM), insect-bearing sediments are occasionally found underneath the veneer of glacial deposits. Notable among these is the prolific site at Sourlie, near Irvine, western Scotland, which yielded a rich assortment of plant and animal fossils, including reindeer and wooly rhinoceros as well as abundant insect remains. Radiocarbon dates on plant remains and the collagen from a reindeer antler found in situ grouped around 30.0 ky BP (Bos et al., 2004). MCR paleoclimatic estimates based on the beetle fauna gave mean July temperatures of between 6 and 9 C and mean January/
BEETLE RECORDS | Late Pleistocene of Europe February temperatures of 34º and 11 C. A more recent study of a glaciotectonized organic deposit at Balglass Burn, high on the slopes of the Campsie Fells east of Glasgow, gave ages between 34.5 and 28.0 ky BP based on six radiocarbon dates on sedge fruits and beetle remains (Brown et al., 2007). Besides insects, pollen and plant macrofossil remains were also studied. All these indicate a small pond situated in an open tundra landscape. MCR estimates based on the beetle assemblage give Tmax 8–10 C and Tmin 26 to 10 C. The climatic implications of these beetle faunas demonstrate that much of central Scotland was not covered by glaciers during the middle pleniglacial at a time when thermal conditions were apparently sufficiently cold to permit ice accumulation in the highlands. Some other factor such as inadequate precipitation must also have been involved in maintaining ice-free conditions in northern Britain at this time. In the Netherlands pleniglacial, two extensive Middle Weichselian insect assemblages are known. Thus, a beetle fauna from Peelo included many of the exclusively arctic and/or Asiatic species that were also present in the British Isles at the same time (Coope, 1969). Excavations at Orvelte, near Drentse, to recover mammoth and wooly rhinoceros remains, enabled a large beetle fauna to be investigated that included an assemblage of cold-adapted species similar to the ones discussed above (Cappers et al., 1993). Both of these faunas yield MCR Tmax temperature estimates at or below 10 C and Tmin values below 20 C. A Middle Weichselian insect fauna from Poland included a very similar group of high arctic and Asiatic species (Kasse et al., 1998). MCR estimations based on this assemblage gave the following estimates: Tmax 8–12 C, Tmin 27 to 20 C. Insect remains were found in deposits referring to the Oerel interstadial both at the Oerel site and at Gross Todtshorn. The beetle assemblages are composed of northern Palaearctic species, which indicate a treeless environment and cold climate (Tmax 12–13 C, Tmin 9 to 2 C). At the Oerel site, insect remains were also recovered from a peat layer referred to the younger Glinde Interstadial. The beetle assemblage from this horizon is similar in composition to those from the earlier interstadial. Climate conditions were also likely to have been very similar (Tmax 9–14º, Tmin 17 to 2 C). In Switzerland, two insect assemblages of middle Weichselian age have been recorded from Gossau (Jost-Stauffer et al., 2005). They include several exclusively arctic and Asiatic species that do not occur anywhere in central Europe at the present day. MCR estimates gave the following figures: Tmax 8–13 C, Tmin 21 to 7 C. A second beetle assemblage from the Swiss plateau was obtained from samples taken in a peaty matrix of the mammoth skeleton at Niederweningen that was radiocarbon dated to around 45.0 ky BP (Coope, 2007). This fauna also includes many exclusively arctic/Asiatic species. The mammoths were probably mired in a reedy swamp in a landscape with scattered willows, birches, and conifers. An MCR estimate indicates Tmax around 10 C and Tmin below 10 C. Middle Weichselian beetle assemblages from La Grande Pile are characterized by the presence of almost entirely arctic and continental species. Obligate tree-dependent species were not recorded. The climate was cold in general through the period (Tmax around 12 C, Tmin approximately 10 C or below). A site of outstanding archaeological importance has recently been recorded at Lynford, Norfolk, where an extensive
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beetle fauna was found in close association with abundant evidence of Neanderthal exploitation of mammoths: smashed bones and numerous flint tools (Boismier et al., 2011). There can be little doubt that the beetle assemblage is contemporary with the human activities and provides an intimate view of the local environment (Coope, 2011). However, the beetle assemblage was unusual. It included abundant high northern and Asiatic species suggesting cold continental climatic conditions. But this fauna lacks several of the extreme cold-adapted species present in the typical full-glacial assemblages and includes rare species whose distributions do not extend into arctic regions today. These make MCR estimates difficult. Nevertheless, it is likely that the mean July temperatures were between 12 and 14 C and January/February figures at or below 10 C. The age of this important site is probably sometime late in MIS 4 or early MIS 3.
Last Glacial Maximum (MIS 2) Conditions during the LGM were extremely harsh and insect faunas from this period are consequently very rare. A small beetle assemblage obtained from Dimlington, East Yorkshire, has been radiocarbon dated to 18.5 ky BP (Penny et al., 1969). Given the smallness of this insect fauna, only a provisional MCR estimate is possible: Tmax 9–11 C, Tmin 27 to 7 C. These figures do not necessarily represent the typical climate at the LGM for England because the insects came from a single fossiliferous horizon in an otherwise barren sequence of laminated silts and may thus indicate a short period of atypically warm conditions in an otherwise even colder episode. In continental Europe, two beetle assemblages are known from the LGM. Beetle assemblages from La Grande Pile dating to this period indicate very cold and harsh conditions, with Tmax mostly around 10 C and Tmin well below 10 C. Fossil remains of beetles and oribatid mites, which are dated to around 18 800 BP, were recovered from a site close to Verona, northeastern Italy (Foddai and Minelli, 1994). They indicate climate conditions colder and wetter than today. Tmax may have been 8–9 C lower than present day.
Late Glacial Period This period falls well within the radiocarbon dating techniques. Dates are cited here in radiocarbon years BP and calibrated calendar years are shown in parentheses as cal years BP. Correlations are made with the climatic event stratigraphy of the Greenland Ice cores (Walker et al., 1999). As a large number of studies have been carried out on beetle assemblages from sites scattered all over Europe (cf. Coope et al., 1998), only a representative selection is discussed here (Figure 3).
Late Glacial stadial (Greenland stadial 2) Faunas that date from the cold period (Greenland stadial 2, GS-2) that immediately preceded the Late Glacial Interstadial, namely from just before 13.0 ky BP (ca. 15.4 cal ky BP), are rare. In Britain, the most stratigraphically secure faunal assemblage of this age is from the North Wales coast at Glanllynnau, where a sequence of insect-bearing deposits spans almost the whole of the early part of the Late Glacial Interstadial (Coope and Brophy, 1972, cited in Coope et al., 1998). The stadial
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Late Glacial sites
Figure 3 Map showing Late Glacial sites where fossil beetle assemblages were studied.
deposits yielded exclusively arctic and Siberian species. MCR calculations on this assemblage gave the following estimates: Tmax 10–11 C, Tmin 26 to 23 C. A second beetle assemblage of this age was associated with the recovery of the skeletons of an adult mammoth plus at least three juveniles from silty clay in a kettle hole at Condover, near Shrewsbury (Allen et al., 2009). The GS-2 stadial beetle fauna included many exclusively high arctic species, but unfortunately the wallowing of the mammoths in their efforts to extricate themselves from the adhesive clay had churned up the sediment, incorporating some of the more recent Late Glacial sediment into the stadial deposits, making the precise interpretation of the environment difficult. It is, however, clear that the stadial assemblage of beetles lacked the eastern Asiatic component so characteristic of the full-glacial faunas, suggesting that although the climate must have been of arctic severity at this time, it was probably less continental than during the mid-glacial (pleniglacial) episode. Only two beetle assemblages of this age are known from northern Europe. Both of these faunas are from Kullaberg, southwestern Sweden, and can be dated with confidence to this period (Lemdahl, 1988, cited in Coope et al., 1998). Although these include only a few species, they resemble those from Britain. Similarly, cold-adapted suites of beetle species are also known from Poland, and from the Swiss Plateau (Coope and Elias, 2000; Coope and Lemdahl, 2009; Elias and Wilkinson, 1983; Gaillard and Lemdahl, 1994; Hardon et al., 2000). MCR estimates gave the following figures: Tmax 10–13 C, Tmin 17 to 0 C.
Late Glacial interstadial Beetle faunas from the Late Glacial interstadial 1 (GI-1) are widespread across the whole of northwestern Europe. In Britain, important faunas have been described from Glanllynnau, North
Wales (Coope and Brophy, 1972, cited in Coope et al., 1998), St Bees, Cumbria (Coope and Joachim, 1980, cited in Coope et al., 1998), Gransmoor, East Yorkshire (Walker et al., 1993, cited in Coope et al., 1998), Hollywell Coombe, Kent (Coope, 1998) and from Llanilid, South Wales (Walker et al., 2003). Faunas from the early part of the GI-1e interstadial (the Bølling phase) from about 12.8 ky BP (15.1 cal ky BP) or slightly earlier, include an ecologically diverse variety of relatively southern species. At this time the vegetation was thin and made up chiefly of pioneer plant species. The only trees that were present were birches and willows and these were probably confined to the damp valley bottoms. At this time northern Europe as a whole was largely without a woodland cover in spite of the fact that for much of the region, the climate was warm enough to have supported mixed oak forest. Edaphic factors are thought to have been largely responsible for this open landscape. Average MCR estimates from these sites give approximate figures: Tmax 17–18 C, Tmin 7 to 1 C. Beetle faunas indicate that the initial warming at the start of the interstadial involved a rapid rise in mean July temperatures of at least 7 C and a corresponding rise in winter temperatures substantially greater than this. The rate at which the mean July temperature rose was about 1 C per decade and even higher (Coope and Brophy, 1972, cited in Coope et al., 1998). Because this change also involved a switch from continental to oceanic climatic conditions, the rise in mean annual temperature was considerably greater than these figures suggest, though the precise number of degrees involved is impossible to estimate with confidence. The beetle reconstruction that yielded this unexpectedly high rate of climatic change has subsequently been supported by evidence from the Greenland Ice cores, which show a dramatic climatic amelioration at about this time (Johnsen et al., 1992). It is interesting to observe that this episode of rapid climate warming at the start of the Late Glacial Interstadial is not recorded by beetle assemblages from southern Sweden (Lemdahl, 1988, cited in Coope et al., 1998) or from the coast of western Norway. During this period, the Scandinavian beetle assemblages are of an arctic or boreal character. It is likely that this reflects the local cooling effect provided by the close proximity of the waning Fennoscandian ice sheet. In contrast to the Scandinavian situation, faunas from Poland and the Netherlands (Geel et al., 1989, cited in Coope et al., 1998) indicate temperatures similar to those of the British Isles. At about the same time, warmth demanding beetle faunas are recorded from the Swiss Plateau (Coope and Elias, 2000; Coope and Lemdahl, 2009; Elias and Wilkinson, 1983; Gaillard and Lemdahl, 1994; Hardon et al., 2000). Similar faunal changes are traced in southern France (Ponel and Coope, 1990; Ponel et al., 2001). Shortly before 12.0 ky BP (ca. 14.0 cal ky BP), there was a sudden climatic cooling. Tmax figures suddenly fell to around 15 C or even lower and never recovered again during the rest of the interstadial; in other words for much of Western Europe the whole of the Allerød phase was substantially cooler than the preceding Bølling phase. The beetles indicate that during the Allerød there appears to have been several relatively minor climatic oscillations, well illustrated by the climatic curves from Gransmoor (Walker et al., 1993, cited in Coope et al., 1998) and Llanilid (Walker et al., 2003, and in summary by
BEETLE RECORDS | Late Pleistocene of Europe
Figure 4 Fieldwork at the small marsh (80 30 m) at He´re´mence situated at 2290 m.a.s.l. in the Swiss Alps. Four sites at different altitudes, including this site, were studied in a multiproxy project in order to record biotic responses to rapid climatic changes around the Younger Dryas in the Swiss Alps. Samples were recovered by using a Streif-piston corer of 8 cm diameter.
Lowe and Walker, 1997, cited in Coope et al., 1998). The frequency of these oscillations makes it difficult to give useful MCR estimates for the latter half of the Late Glacial Interstadial. Again, in contrast to the British Isles and much of Europe, the beetle assemblages from southern Sweden indicate that the highest temperatures were reached during the latter part of the Late Glacial Interstadial. However, it must be emphasized that the actual MCR temperature estimates were similar to the British figures at this time. In Norway arctic faunas are recorded for the Allerød phase (Birks et al., 1993, cited in Coope et al., 1998; Lemdahl, 2000a). In Poland species with a boreal distribution appear during this period (Lemdahl, 1991a, cited in Coope et al., 1998), which also seems to be the case in southern France. These beetle assemblages show that during the GI-1 there were distinct regional differences in climate and steep temperature gradients across much of northern Europe (Coope and Lemdahl, 1995; Coope et al., 1998).
Late Glacial stadial (GS-1) ¼ Younger Dryas The beetle evidence from the British Isles suggests that the beginning of the GS-1 (Younger Dryas) cold period appears as a culmination of the sequence of Allerød cooling events outlined above. Its beginning is conveniently dated at about 11.0 ky BP (ca. 13.0 cal ky BP). Many beetle assemblages are known from the Younger Dryas period (e.g., Coope et al., 1979, Coope and Joachim, 1980, cited in Coope et al., 1998; Coope, 1998; Walker et al., 1993, 2003) and include a large number of exclusively arctic species, many of which were previously encountered in the assemblages from full-glacial times. However, exclusively Asiatic species are very rare at this time. An average of MCR estimates based on faunas from this period indicates the following figures: Tmax 9–11 C, Tmin 10 to 20 C (Figures 4 and 5). In northern France, insect assemblages from Younger Dryas fluvial deposits recorded at St.-Momelin, yielded these MCR estimates: Tmax around 10 C and Tmin close to 12 C (Ponel
205
Figure 5 Fossil head and thorax matching a modern specimen of the carrion beetle Thanatophilus dispar. Remains of this species were abundant in lake sediments dating to the Oldest Dryas (GS-2) at the site Hauterive Rouges-Terres, at the shore of the Lake Neuchaˆtel, Switzerland. A Paleolithic settlement was excavated adjacent to the site. Today T. dispar is mainly found on fish carrion where it probably feeds on fly larvae.
et al., 2007). MCR reconstructions show that, in contrast to the Late Glacial Interstadial, relatively similar climatic conditions seem to have prevailed in most of northern Europe at this time. In eastern Finland high arctic beetle faunas of Younger Dryas and early post glacial age colonized recently deglaciated areas (Bondestam et al., 1994). Although Younger Dryas beetle assemblages are very rare from the Swiss Plateau, they suggest cold, arctic conditions (Lemdahl, 2000b). Many sites that include the transition from the Late Glacial to the Holocene show signs of a hiatus, making it difficult to estimate the rate of change involved. However, all the available beetle evidence suggests that the change was abrupt and intense and probably similar in magnitude to the one at the start of the Late Glacial Interstadial (e.g., Ashworth, 1973; Lemdahl, 1991b, cited in Coope et al., 1998; Lemdahl, 2000; Ponel et al., 2007). The beetles indicate that throughout most of Europe at the start of the Holocene, temperatures reached present day levels in a geologically immeasurably short period of time.
See also: Beetle Records: Middle Pleistocene of Western Europe; Overview; Postglacial Europe.
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