Rissian, Eemian and Würmian Coleoptera assemblages from La Grande Pile (Vosges, France)

ELSEVIER

Palaeogeography, Palaeoclimatology,Palaeoecology114 (1995) 1-41

Rissian, Eemian and Wiirmian Coleoptera assemblages from La Grande Pile (Vosges, France) Philippe Ponel Laboratoire de Botanique historique et Palynologie (Bofte 451), UA CNRS 1152, Facultk des Sciences et Techniques de Saint Jbrdme, F-13397 Marseille Cedex 20, France

Received 31 January 1994; revised and accepted 24 August 1994

Abstract

The Grande Pile peat-bog sequence is one of the few west European sites that cover the entire time span of the last major climatic cycle (140,000 years). A recent program of coring has provided material for insect analysis. The aim of this palaeoentomological study is to interpret the environmental and climatic evolution from the end of the Rissian glaciation to the Holocene using subfossil Coleoptera. The studied samples yielded 394 taxa of Coleoptera, half of them identified to species level; 19 of which do not belong to the present-day French fauna. The large number of taxa suggests a wide variety of habitats and provides much detailed palaeoecological evidence for the period studied. The lowermost sediments of the sequence, corresponding to the end of the Rissian glaciation, were deposited under very cold conditions in a tundra environment. This is succeeded by a forest period in which two cool interludes of grassland environment occur. Although these periods are decidedly poor in tree-dependent Coleoptera they do not contain any really cold-adapted taxa. They divide the forest phase into three periods. The first one, corresponding to the Eemian Interglacial, shows an early stage in which the beetle fauna is characterized by species dependent on deciduous trees, a later stage in which this fauna is mixed with many conifer-dependent elements, some of which (e.g. Platypus oxyurus) suggest warmer and perhaps wetter climatic condition than today. The two later woodland periods yielded coleopteran assemblages rather similar to those recorded in the second part of the Eemian, i.e. with both deciduous- and conifer-dependent taxa. There is some evidence to suggest that these two periods were slightly cooler than the Eemian proper. Marked climatic deterioration becomes obvious in the upper half of the sedimentary sequence attributed to the last glacial period (Wiarm), with the reappearance of tundra beetle assemblages. Sediment and insect evidence suggest that the climate was extremely cold and continental at La Grande Pile at about 30,000 B.P. A comparison of the insect analysis with previous palynological works enables precise correlation between the results provided by these two independent approaches. However, large numbers of running-water Coleoptera in the forest periods, replaced by standing-water Coleoptera in cold periods, raise questions concerning the lacustrine origin of the sedimentation at La Grande Pile. 1. Introduction

The Grande Pile site shows an almost continuous sedimentary record, about 20 m thick, covering the last climatic cycle from the Riss glaciation through the whole of the Last Interglacial Complex, much of the Last Glaciation and the Holocene. It was the first west European 0031-0182/95/$9.50 © 1995 Elsevier Science B.V. All rights reserved SSDI 0031-0182(95)00083-2

site for the interpretation of the palaeoclimate of the last 140,000 years. Subsequent pollen analysis of two additional sites, the Echets mire near Lyon (Beaulieu and Reille, 1984) and the Massif Central maars (Reille and Beaulieu, 1990; Beaulieu and Reille, 1992b), have been shown to cover the same period. Since its discovery (Seret, 1967), La Grande Pile

2

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

has been the subject of intense palaeoecological research: pollen analyses by Woillard (1973, 1974a,b, 1975, 1977a,b, 1978a,b, 1979)then by Beaulieu and Reille (1992a); algal (diatoms) analyses by Louis and Smeets (1981), Louis et al. (1981), Louis and Ermin (1983), Louis and Peters (1983), Louis et al. (1983), Cornet (1988); palynology and sedimentology (Seret et al., 1992); palaeoclimatology (Guiot et al., 1989, 1992, 1993). Up to now, no palaeoentomological investigation had been undertaken at La Grande Pile, although it is now well established (Buckland and Coope, 1991) that in Quaternary ecology and climatology, insects and particularly Coleoptera can yield precise information and permit quantification of climatic reconstructions to be made (Atkinson et al., 1986, 1987). It must be emphasized that the present study is the first palaeoentomological analysis tracing the environmental and climatic evolution from the end of the penultimate glaciation to the Holocene (this paper dealt with this period up to the phase of maximum cold of the Wt~rm Glaciation).

2. Study area and site

The location of the Grande Pile peat-bog is so well known that only a brief summary will be

~

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PARIS

given here (Figs. 1 and 2). It occupies a closed depression located on an interfluvial plateau about 20 m above the Ognon valley. On a geological point of view, this peat-bog is located on the southern vosgian piedmont which culminates at Ballon de Servance (1216 m). It lies in the Saint Germain basin made up of Visean coal-bearing formations covered with Triassic sandstone sediments. These formations are buried under a thick fluvioglacial cover deposited during the Rissian and Wtirmian glaciations. The Triassic formations underlying the Grande Pile depression show an alternance of marl and sandstone sediments with local dolomitic facies (Th6obald et al., 1974). The peat-bog itself has a surface area of 25 ha at an altitude of 325 m above sea level. It is, at the present day, isolated from the surface hydrographic network (Seret et al., 1990) and no tributary feeds it (the Coleoptera provide evidence that, in the past, this isolation was not continuously present throughout the period studied). The mire has been recently drained by two channels, the first one northwestwards, the second one southwards, dug for peat extraction. The Ecromagny plateau (where La Grande Pile lies) was covered by the Linexert (or Rissian) ice sheet as evidenced by the presence of glaciolacustrine tills of the Linexert glaciation at the basis of the sedimentary sequence. The absence of

~ EPlNALI La Grande Pile

FRANCE

BESANCON

Fig. 1. Location of the Grande Pile peat-bog (47°44'N, 6°30'14"E).

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

1

2

3

3

Fig. 2. Topographic map of the Grande Pile peat-bog and its surroundings. 1 = The Grande Pile peat-bog. 2 = Lakes. 3 = Contour levels every 10 m, altitude in m a.s.l.

Wtirmian till deposits suggests that it was not covered by the Wtirmian ice sheets. Thus, La Grande Pile is located between the westward extension limits of the Rissian and Wtirmian moraines (Seret et al., 1990; Beaulieu et al., 1992). The Grande Pile mire is today surrounded by a

forest that belongs to the phytosociological community Quercion robori-petraeae; Carpinus betulus is also abundant and indicates the euro-siberian character of the local climate. Molinia coerulea occupies the peat-bog itself, concurrently with Sphagnum spp., Drosera rotundifolia, Oxycoccos

4

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

palustris, Eriophorum vaginatum, Menyanthes trifoliata and several Carex: C. fusca, C. canescens, C. vesicaria, C. stellulata. Several tree species are also colonizing the mire itself, e.g. Betula pubescens, Populus tremula, Frangula alnus, Salix spp. and Quercus robur (Woillard, 1975). The

stationary piston corer (O 8cm) (Aaby and Diggerfeldt, 1986). The first series of 8 corings about 1 m/1.5 m away from one another and about 20 m depth was made in the centre of the mire near to the site where Woillard's most complete diagram (1975, diagram X) was derived (Fig. 3, corings A, cores GP 90-1, GP 90-2, GP 90-3, GP 90-4, GP 90-5, GP 90-6, GP 90-7 and GP 90-8). The second series of cores (Fig. 3, corings B, cores GP 90-9, GP 90-10, GP 90-11, GP 90-12, GP 90-13 and GP 90-14) was bored nearer to the littoral area, 100 m southwards on the same line as the old GP XX coring site that yielded the diagram of Beaulieu and Reille (1992a). The cores were stored in PVC tubes. Only the core series from the central area (A) has been studied in detail so far.

mean temperatures for January and July are 0°C and 18.6°C, respectively (mean yearly temperature: 9.5°C). The yearly precipitation is 1040 mm.

3. Methods

3.1. Sampling In order to get sufficient sample volume, two series of parallel cores were undertaken using a

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P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

5

3.2. Lithostratigraphy The correlation of the stratigraphy of the cores took into account the nature, texture, colour and consistency of the sediment, the latter classifying into 4 main types: sand, silt, clay, and gyttja (Fig. 4). This permits the equivalent layers to be recognized from one core to another and the definition of more or less homogeneous sedimentological zones. This operating procedure has already been used at La Taphanel peat-bog (Massif Central, France) (Ponel and Coope, 1990; Ponel et al., 1991), This method enabled the subdivision of nearly all the cores GP 90-1 to GP 90-8 into 44 sedimentary slices, 8-9 kg each, numbered from 0 to 41 (layers 3 and 4 are split in two: 3A-3B, 4A-4B). The location of the 44 samples is shown on Fig. 4; it is an "average" location since the equivalent levels collected within a sample do not necessarily correspond to the same depth in each core. Good correlations can be established between our core stratigraphy and the stratigraphical logs published by Woillard (1975), but with some minor discrepancies. Of particular interest is a layer (sample 38 and 39) of an in situ intraclastic breccia which was not described in previous investigations at this site. This layer consists of fragments of bedded and disrupted sediments with no evidence of having been transported. It could indicate an episode of dessication or shallow water sedimentation within the range of winter freezing that could similarly disrupt the bedding. It is clear however that the conditions that lead to the disruption did not occur either above or below this particular horizon. The role of the insect fauna in this interpretation will be discussed later (study of GP-A7 faunal unit).

3.3. Correlation with palynology and chronology

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The chronology of this sequence (Fig. 5) is established by correlation with previous studies but is mainly derived from the interpretation proposed by Beaulieu and Reille (1992a); the latter is an attempt to correlate recent pollen diagrams and radiocarbon dates from La Grande Pile (Beaulieu and Reille, 1992a; Woillard, 1975, 1978a; Woillard

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Fig. 4. Synthetic lithostratigraphy of the central core series at La Grande Pile and location of samples for insect analysis. I = Bedded clay, silt and sand, 2 = silt, 3 = organic silt, 4 = silty breccia, 5 = minerogenic gyttja, 6 = organic gyttja, 7--Late Glacial gyttja. Numbers left: depth (m); numbers right: sampling levels.

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P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

and Moock, 1982) on the basis of palynological events.

4. Insect analysis

4.1. Extraction procedure for insect remains The sediment from each level was treated using the extraction method recommended by Coope (1986). The samples were first broken gently by hand in water in a bowl. Mild chemical treatment (sodium carbonate solution) was occasionally necessary. Water with macroremains mixed with disaggregated sediment was allowed to flow in a 300 #m sieve. The material collected in this way was largely made up of plant debris and in most cases an insect concentration had to be undertaken. The damp residue was mixed with kerosene, then excess oil poured off and water added to the bowl. After decanting the floating fraction, rich in insect remains, was then poured into the same sieve and washed, first with detergent then with water and with alcohol. The material obtained was sorted under a binocular microscope. The insect remains were eventually preserved in alcohol or glued on pieces of cardboard. Beetles macrofossils were identified by direct comparison with a modern reference collection. A total of 41 samples were analysed in this study (the insect assemblages from levels 24, 25 and 28 have been left out because of possible contamination during sampling). 394 taxa of Coleoptera were identified, more than half to species level (Table 1); 19 species present as fossils at La Grande Pile are not members of the present-day French fauna. Every sample yielded Coleoptera remains. Other insect remains such as chironomid larva heads and Trichoptera larval sclerites were observed but not identified. The sediment yielded also numerous remains of Crustacea such as Cladocera and Notostraca; the occurrences of the latter are briefly discussed in this paper in addition to beetle analysis.

4.2. The insect fauna The Coleoptera may be classified into several categories according to their ecological require-

7

ments, i.e. aquatic species (both running-water and standing-water species), terrestrial species, riparian species (that live on the shores of standing-water or on fiver banks), coprophagous and coprophilous species (directly or indirectly dependent on mammal faeces), necrophagous species (that live on dead animals), tree-dependent species (insects linked to leaves, wood, bark, etc) and plant detritus feeders. The wide variety of habitat requirements suggests that the assemblages obtained from La Grande Pile shows that the insect fauna was derived from a broad area of diverse environments. The main part of the Coleopteran record probably represents a fauna that lived in the close vicinity around the site (Lemdahl, 1990). However, most beetles fly readily and some specimens may have originated some kilometres away from the site. The list of taxa (Table 1) also indicates the occurrence of species with diverse climatic requirements. Their modern distribution range from boreal, boreo-montane to mediterranean. The succession of these species clearly reflects the largescale climatic changes that took place in this part of Europe during the last climatic cycle (Pons et al., 1992).

4.3. Analysis of the ma& ecological groups at La Grande Pile Total representation of Coleoptera (Fig. 6) The histogram of taxa shows some variations in the number of taxa per sample and in the variations of the total number of individuals per sample. It should be noted that no sample is totally devoid of beetle remains. Great variation is found in the total number of individuals which ranges from as low as 15 individuals to as many as 260 individuals per sample. Two drastic decreases occurred around samplel 0 and around samples 17-18.

Tree-dependent Coleoptera (Fig. 7) In this category are gathered all conifer and deciduous tree-dependent Coleoptera, the latter including willow and boreo-montane dwarfwillow-dependent beetles. The histograms of taxa and individuals (Fig. 7a) show clearly a dramatic change at the transition between sample 20 and

CARABIDAE Cicindela sp. Calosoma sycophanta (L.) Carabus clathratus L. C. cancellatus C. nitens L. C. arvensis Hbst. Carabus sp. Nebria gyllenhalli (Schrnh.) Nebria sp. Notiophilus sp. *Elaphrus lapponicus Gyll. Elaphrus riparius (L.) *Diacheila arctica (Gyll.) *D. polita (Fald.) Loricera pilicornis (F.) Dyschirius globosus (Hbst.) Perileptus areolatus (Creutz.) Trechus secalis (Payk.) T. rubens (F.) T. obtusus Er./ 4-striatus (Schrk.) Trechus sp. Bembidion bipunctatum (L.) B. nitidulum (Marsh.) *B. dauricum (Mots.) B. (Testediolum) sp. B. schappeli Dej. B. aeneum Germ. R unicolor Chaud. 1

1

11

2

1 1 1

1

1

11

1

1

1

1

1 1

1

1

121

1

1

1

11

1

1

1

1

1

1

2

3 1 1

1

1

1

1

1

1

1

1

10 11 12 13 14 15 16 17 18 19 20 21 22 23 26 27 29 30 31 32 33 34 35 36 37 38 39 40 41

1

1

1 2 3A 3B 4A 4B 5 6 7 8 9

1 1

0

Table 1 List of coleoptera showing the minimum number of individuals recovered from each sample. Nomenclature and taxonomic order from Lucht (1987). Species which are not members of the present-day French fauna are marked with an asterisk (*)

e~

DYTISCIDAE Hyphydrus ovatus (L.) Coelambus impressopunctatus (Schall.) Hydroporus palustris (L.)

Haliplus sp.

(Vanz.)

HALIPLIDAE Brychius elevatus

A. muelleri (Hbst.) A. fuliginosum (Panz.) Agonum sp. Amara lunicollis (Sehdte) A. quenseli (Sch~nh.) Amara sp. A. (Cyrtonotus) sp.

(Panz.)

P. diligens (Sturm) P. vernalis (Panz.) P. nigrita (Payk.) P. niger (Schall.) P. melanarius (Ill.) P. madidus (F.) Pterostichus sp. Abax sp. Synuchus nivalis (Panz.) Calathus melanocephalus (L.) ~4gonum ericeti

(Dej.)

B. guttula (F.) Bembidion sp. *Patrobus assimilis Chaud. Patrobus sp. Bradycellus ruficollis (Steph.) Poecilus lepidus (Leske) Poecilus sp. Pterostichus pumilio

1

1

1

1

1

1

1

2 2

4

1

2

113 1

1

1

1 1

1

1

1

7"

¢

H. gracilis Hydraena sp. Ochthebius gr. foveolatus Germ. Ochthebius sp.

HYDRAENIDAE Hydraena gr. riparia Kug.

(F.)

Rhysodes sulcatus

RHYSODIDAE

Gyrinus minutus F. G. aeratus Steph. G. aeratus Steph./marinus Gyll. Gyrinus sp.

GYRINIDAE

Colymbetes sp. Graphoderus sp, Acilius sp. Dytiscus sp.

(Payk.)

*C. dolabratus

(L.)

A. sturmi (Gyll.) *A. arcticus (Payk.) Agabus sp. llybius sp. Rhantus sp. Colymbetes fuscus

(L.)

Hydroporus sp. Potamonectes griseostriatus (Geer) P. assimilis (Payk.) Potamonectes sp. Agabus bipustulatus

Tablel(continued)

1

0

2

1 2

1

3A 3B 4A 4B 5

1

6

1

1

7 8

1

9

1

3 3 4 1

I1

1

2

1

2

1

?1

12

I

i

2

1

2

3

1

1

1

2223

1

1

11191

10 11 12 13 14 15 16 17 18 19 2 0 2 1

1

1

1

I

1

1

22

262729

1

1

1

1

43

1

1

2

43

t

1

1

1

22

1

1

2

I 1

1

511

1

1

1

I

3 4 1 4 1 1 1

I

30 31 32 33 34 35 36 37 38 39 4 0 4 1

4~

4~

,2

Necrophorus sp.

(Gyll.)

Thanatophilus sp. Pteroloma forsstroemi

SILPHIDAE

(Hbst.) Hister sp.

Paromalus flavicornis

(Panz.)

Plegaderus vulneratus

HISTERIDAE

lum (Hbst.)

Enochrus sp. Chaetarthria seminu-

(Gr.)

Anacaena sp. Laccobius sp. Enochrus affinis ( Thunb. )/coarctatus

(L.)

Hydrobius fuscipes

turn (F.)

Cercyon sp. Megasternum boletophagum (Marsh.) Cryptopleurum minu-

/us (L.)

Sphaeridium sp. Cercyon melanocepha-

(F.)

11. glacialis Villa 12 H. brevipalpis Bedel 13 H. ?flavipes F. H. gr. minutus F. Helophorus sp. 1 16 1 Coelostoma orbiculare 1

type

Limnebius sp. Hydrochus sp. Helophorus grandis IlL *H. sibiricus (Mots.) H. aquaticus (L.) *H. oblongus LeC.

1

1

1

1

1

1 22

2

1

2

1

1

2 1

11

11

22

1

34143

1

2 1

1

1

1

5 4 7 4

1

2

1

I 21

11

1

11

7

1

6

1

1

31

3

1

2

11

243

1

3 5 3 1

1 2 1 2 2 1

1

12

7

2

xt~

E"

.¢

G. plagiatus (F.) G. cf. kunzei Heer Anthophagus sp.

(Mall.)

Lesteva sp. Geodromicus nigrita

(Mannh.)

Acidota crenata (F.) A. cruentata

(Gray.)

O. fuseum (Grav.) *0. boreale (Payk.) Olophrum sp. Eucnecosum brachypterum

(Gyll.)

*Olophrum ?consimile

Grav.

Omalium caesura

(Steph.)

Acrolocha cf. sult'ula

(Gyll.)

Megarthrus sp. 1 Megarthrus sp.2 Proteinus spp. Eusphalerum spp. Acrulia inflata (Gyll.) Pycnoglypta lurida

Curt.

Micropeplus tesserula

STAPHYLINIDAE

Acrotrichis sp.

PTILIDAE

1

I

3A 3B 4A 4B 5 6

2

1

1

1

2 2 4

I

I

7 8 9

1

1

2

LIODIDAE G. sp. SCYDMAENIDAE Neuraphes sp.

l

1

0

CATOPIDAE Cholera sp. Catops sp.

Table 1 (continued)

2

2

1

8

I

1

2

6

1

3

2

1

1 1

2

I 21 15 10 1

1

2

I

I~112

3

1

I

2

3

1

1

1

I

132

1

2

1

1

I

1

1

1

13

1

2

8

1

1

1

1 1

1

1

1

2

1

1

1 1

2

1

1

1

2

1

1

1

1

1 1

l0 II 12 13 14 15 16 17 18 19 20 21 22 23 26 27 29 30 31 32 33 34 35 36 37 38 39 40 41

t~

e~

o~

e~

e~

G

Batrisodes sp. Bryaxis sp.

PSELAPHIDAE

T. laticollis Grav. T. corticinus Grav. T. elongatus Gyll. T. ?fimetarius Grav. Tachinus sp. Myllaena sp. Aleocharinae indet.

(Creer)

Mycetoporus sp. Bolitobius sp. Tachyporus chrysomelinus (L.) Tachyporus sp. Tachinus rufipes

sp.

Bledius sp. Stenus sp. Euasthetus bipunctatus ( L j u n g h ) Scopaeus sp. Lathrobium terminatum Gray. Xantholinus sp. sl. Staphylinus sp. Philonthus/Quedius

(Grav.)

O. piceus (L.) O. laqueatus ( M a r s h . ) O. sculpturatus Grav. *0. ?politus Er. O. nitidulus Grav. *0. gibbulus Epp. Oxytelus sp. Platysthetus cornutus

Grav.

Oxytelus insecatus

(Gray.)

*B. nordenskioeldi Mhkl. Aploderus caelatus

anus Sahib.

*Boreaphilus henningi-

1

10111

1

1

1

1

1

1

1

1

3

1 2

1 3

1

1

I1

8

2

1

1

1

5

1 5

12

1

1

3

1

6

4

15 4

2

2

1 19 10 5

5

1 5

1

1 2 1 4 2 1 1

1

1

2

8

2 1 2 1 2 1

1

1

2

4

111

1 7 4 2

1

1

1

1

126

1

1

134

1

1 1

1

8

1

11

1

3

1

2

2

2

14109

1

1

2

1

4

1

1

7

1

1 1

1

4

1

1

1

1

1 1

2 1

1

6

1

?11

1 1

1

7

1 1

I

1 3

26128

121

1

2 1

4

1

104

U,

-&

¢%

H E L O D I D A E indet.

G. sp,

Agrilus sp.

BUPRESTIDAE

Bonv.

Throscus carinifrons

THROSCIDAE

(Pill. Mitt.) G. sp.

?Hylochares dubius

EUCNEMIDAE

sp.

Fleutiauxellus maritimus (Curt.) Elateridae indet, pl.

(L.)

Denticollis linearis

(L.)

Ampedus cf. nigerrimus (Lac.) Adelocera murina ( L. I Ctenicera pectinieornis (L.) C. euprea (F.) Ctenicera sp. Prosternum tessellatum (L.) Selatosomus aeneus

ELATERIDAE

Haplocnemus sp. Dasytes sp.

MELYRIDAE

CANTHARIDAE Rhagonycha sp, Malthodes sp.

Table 1 (continued)

0

2

1

1 I

1

1

1 1

1 2

2

1

1

1

2

1

l

I

1

1

1

1

1

1

1

2

I

3A 3B 4A 4B 5 6

2

3

1

t

8 9

1 1

7

1

1

I

2

1 8 4 2

1

1

3

1

1

1

1

1

5

t

1

2

1

1

l0 II 12 13 14 15 16 17 18 19 20 21 22 23 26 27 29 30 31 32 33 34 35 36 37 38 39 40 41

7"

o~

.~

DRYOPIDAE

Airaphilus sp. Silvanoprus fagi (Gu6r.)

Aub6

Monotoma brevicollis

CUCUJIDAE

Rhizophagus depressus (F.) Rhizophagus spp.

RHIZOPHAGIDAE

(F.)

Pocadius ferrugineus

NITIDULIDAE Cateretes sp. Meligethes sp. Epuraea sp.

(L.)

Nemosoma elongatum

OSTOMIDAE

*Simplocaria metallica (Sturm)

BYRRHIDAE Cytilus sp. Byrrhus sp.

O. troglodytes (Gyll.) Oulimnius sp. Limnius volckmari (Panz.) L. opacus (MOll.) Limnius sp. Normandia nitens (M011.) Riolus sp.

(MOll.)

Oulimnius tuberculatus

(Moll.)

Dryops sp. Elmis t.aenea (MOll.) Esolus parallelipipedus 2

1

187 1 6 2

1 1 1

1

2

1

3

2

1

10

2

1

I1

1

1 2 1

3 1

2

8

1

1 111 1

2

1 1

3 7 2 3 7

1127144

1

1 5 2

1 356

1

3 3

4

2 1

1

1

5

2

7 9

2

4 4

8

3

8 6

1

4

2

5 2

1

8

1 2

1 1 4 5 1 2

1

1521104

1010183217171

3

2 6

1

10

3

3

5 3

1

1

7

3

1 1 2

1

11

6

8

1

1

3

1

1

9

8

1

3

2

?1 4

7 1 6

8

1 1

1 1 3

4

2 2

1

1

1

2

4

1

2

1

1

4 1 2 1 4 8 2 1 2 6 2

1 1

1

1

1

1

1

1

2

1

1 1

Chilocorus bipustulatus (L.) *Hippodamia 7-maculata ( Geer)

COCCINELLIDAE ?Epilachna sp. Scymnus sp.

ENDOMYCHIDAE ?Endomychus sp.

C filiforme (F.)

(F.)

Ditoma crenata ( F. ) Colydium elongatum

(oi.)

Pycnomerus terebrans

COLYDIIDAE

sp. type

Enicmus sp. Corticaria/Corticarina

LATHRIDIIDAE

G. sp.

Phalacrus sp. Olibrus sp.

Sturm

Phalacrus caricis

PHALACRIDAE

CRYPTOPHAGIDAE Cryptophagus sp. C (Micrambe) sp. A tomaria sp.

Laemophloeus bimaculatus ( Payk.) Laemophloeus sp.

(F.)

Pediacus dermestoides

Table I (continued) 0

1

1

1 2

I

1

1 1

7 8 9

2 2

3A 3B 4A 4B 5 6

1

2

5

1

6

I

I

2

1 3

12114

1

3

1

10 11 12 13 14 15 16 17 18 19 20 21 22 23 26 27 29 30 31 32 33 34 35 36 37 38 39 40 41

e~

SCARABAEIDAE Geotrupes sp. Onthophagus verticicornis (Laich.) Onthophagus sp.

SERROPALPIDAE Dircaea 4-guttata (Payk.)/australis Fairm.

MORDELLIDAE Anaspis humeralis (F.) Anaspis sp.

ANTHICIDAE Anthicus sp.

OEDEMERIDAE Oedemera lurida ( Marsh.)/virescens (L.)

PTINIDAE Ptinus fur (L.)

ANOBIIDAE Dryophilus pusillus (Gyll.) Ernobius cf. nigrinus (Sturm) Ernobius sp. Anobium punctatum (Geer) A. of. fulvicorne Sturm A. cf. pertinax (L.) Anobium sp. Dorcatoma sp.

CISIDAE Sulcacis ?bidentulus (Rosh.) C/s sp.

H. arctica (Schneider) Harmonia 4-punctata (Pont.)

1

1

1

2

1 1

3

4~

oa

E"

.¢

E-

Lema sp. Cryptocephalus ?moraei Cryptocephalus sp. Adoxus obscurus ( L.)

(Panz.)

Plateumaris sp. Maeroplea appendiculata

sp.

D, thalassina Germ, D. einerea Hbst. Donaeia sp. Donaeia/Plateumaris

(Brahm)

Donacia clavipes F. D. ?versieo[orea

CHRYSOMELIDAE

(Geer)

dlosterna tabacicolor

iF.)

Rhagium bifasciatum

CERAMBYCIDAE

(Hoch.)

Ceruchus chrysomelinus

LUCANIDAE

(L.)

*A. holdereri Reitter A. rufipes (L.) A. t. fimetarius (L.) Aphodius spp. Serica brunnea ( L.) ?Triodonta sp. Anomala sp. Valgus hemipterus

(L.)

Aphodius t. fossor

Table 1 (continued)

1

1

!

2 2112

0 1 2

1

6

1 1

5

1

2

1

4

1

1

1 1

2

2

1

1

1

1

3A 3B 4A 4B 5 6 7 8 9

1

l

1 1

1

3

1 1

1

1

l

2

l

1

1

1

!

1 1

I

1

1

1

1

1

1 1

1

1

2

2 11 2 7 4

I

1

1

5

1

4

106

2

10 11 12 13 14 15 16 17 18 19 20 21 22 23 26 27 29 30 31 32 33 34 35 36 37 38 39 40 41

4t~

e,

-m

,2

oc

viminalis

S. intricatus (Ratz.) S. cf. mali (Bechst.) S. cf. earpini (Ratz.) S. scolytus (F.) S. ratzeburgi Janson

(Gu&.)

Scolytus cf. amygdali

SCOLYTIDAE

G. sp.

Brachytarsus nebulosus ( Forst.)

ANTHRIBIDAE

(Gyn.)

B. debilis (GyU.) B. unicolor/debilis

(Ol.)

Bruchidius fasciatus

sp.

Bruchus/Bruchidius

BRUCHIDAE

Chaetocnema sp. Cassida sp.

(F.)

Phyllodecta sp. Galeruca tanaceti (L.) Luperus sp. Agelastica alni (L.) Phyllotreta sp. Aphthona sp. Haltica sp. Crepidodera sp. Chalcoides fulvicornis

Suffr.

Phytodecta sp. Phyllodecta laticollis

(L.)

Phytodecta

(Laich.)

Chrysomela sp. C. ?cerealis L. Chrysochloa sp. Phaedon sp. Plagiodera versicolora

4

1

2 8

6

5

4

2

6

4

1

2 2

2

1 4

1

3 6 1

2 3

3 5 2 2 1 1

1

2

4

2

1

1

1 1

1

1

I

1

E"

Kissophagus hederae (Schmitt.) Kissophagus/ Xylechinus sp. Pityophthorus pityographus (Ratz.) Pityophthorus sp. Pityogenes chaleographus (L.) P. trepanatus (N6rdl.) P. bMentatus (Hbst.) Pityophthorus sp. Pityokteines spinidens (Reitter) P. curvidens (Germ.) P. vorontzowi (Jacobs.) Orthotomicus suturalis (GyU.) O. laricis (F.) ?Orthotomicus sp. lps sexdentatus (Boerner) Xyleborus saxeseni (Ratz.) X. dryographus (Ratz.) Xyloterus domesticus (L.) Xyloterus sp.

(F.)

Leperisinus varius

(F.)

S. multistriatus (Marsh.) Hylastes ater (Payk.) Hylurgops palliatus (Gyll.) Polygraphus poligraphus (L,) P~ ?subopacus Thorns. Hylesinus oleiperda

Table 1 (continued)

0

l

1

2

1 1

1 ?1 3 6 3

8 9

2

1

I

1 2

4 10 12 8

1

I 2 2

5 8

1

I

2

1 1 5 3 6

1

3A 3B 4A 4B 5 6 7

2

I

1

I

5

I

5

!

2

5

8

3

1

1 2

I

1

2

2

!

2

1

l

2

4

~

2

1

1

4 1

39 11 4

8 6

3

1 2 5 4 2

2

1

I

3

1 2

10 11 12 13 14 15 16 17 18 19 20 21 22 23 26 27 29 30 31 32 33 34 35 36 37 38 39 40 41

"~

,~

~"

"~

t,,,,,

~.

"" ~

""

~-

~-

,~

Bagous sp. Notaris acridulus (L.) N. aethiops (F.) Anthonomus sp.

tus (Schfinh.)

Rhyncolus sp. Stenoscelis submurica-

(GyU.)

Dryophthorus corticalis (Payk.) Rhyncolus elongatus

(Marsh.) Sitona sp. Cleonus (sl.) sp.

Sitona of. flavescens

(F.)

O. rufifrons (GyU.) Otiorhynchus sp. Phyllobius/ Polydrusus sp. Polydrusus sp. Strophosoma capitatum (Geer) Barynotus obscurus

O. dubius (StrUm.)

Rhynchites sp. Byctiscus betulae (L.) B. populi (L.) Deporaus tristis (F.)/seminiger Rtt. 1). betulae (L.) Apion cerdo Gerst. Apion spp. Otiorhynchus gr. clavipes (Bonsd.)

(Scop.)

P. tomentosus (Gyll.) Rhynchites ?auratus

nanus (Payk.)

Pselaphorhynchites

CURCULIONIDAE

P. cylindrus (F.)

(Duf.)

Platypus oxyurus

PLATYPODIDAE

1

1

91

5

1

1

2

1

2 2

1

1

1

1

1

1

1

3

1

1

1

1

2 2 2 3 2 1 1

5 5 1 1

1 322

211

1

1

1

11

2112

141

1

1

231

11

1

1

1

7

1

3

1

11

1

2

1

1

1

1

1 1 3 1 3 5 3 1 1 4 2 1

1

21

1 1 1 1

1

7"

.¢

R. foBorum (Mfill.) R. (Pseudorchestes) sp. Rhynchaenus sp. Rhamphus pulicarius (Hbst.)

Phytobius muricatus Bris./granatus Gyll. Micrelus ericae (Gyll.) Gymnetron sp. Miarus sp, Anoplus plantaris (Naezen) Rhynchaenus quercus (L.) R_ avellanae (Oonov.) R. rusci (Hbst.)

Limnobaris T-album (L.)/pilistriata (Steph.) Eubrychius velutus

Brachonyx pineti (Payk.) Curculio venosus (Gray.) C. glandium Marsh. C. pyrrhoceras Marsh. Pissodes pini (L.) P. ?harcyniae (Hbst.) Magdalis nitida (Gyll.) Alophus triguttatus

Table 1 (continued)

0

2 1

1

3

2

3 1

2

2 2

l

1

2

1

1

1

1

t

2 2

8 2

l

I

I

1

3

I 2 3A 3B 4A 4B 5 6 7 8 9

1

1 1

2

1

2

-

l

5

1

2 1

1

!

1

1

1

1

1

1

1 1 3 4 1

2

2

I

3

1

2

1 6

1

1

1

1

2

1

1

~

1

1 3

1

1

1

l0 ll 12 13 14 15 16 17 18 19 20 2l 22 23 26 27 29 30 31 32 33 34 35 36 37 38 39 40 41

~-~

-Q

F

~"

~"

.~

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41 0

g.

I I I II

41 40 39 38 37 36 35 34 33 32 31. 30 28 2;' 26 23 22 21 20 19 18 17 16 15 14 13 12 11 10 9 8 ? 6 5 4B 4A 3B 3A 2 1 0

II

I I Ill

I I I r i l l

II

260 0

lO

I I I l l l l l t l l " l

I I

.

GP-A7b

l l .........

~m~mmm

m mmm m m m m m m

_ _

GP-A7a

GP-A6 GP-A5

GP-A4 -GP-A3 GP-A2b

23

sudden decreases during the predominantly forest period: one in sample 10, the other in samples 17-18. Many beetle species are dependent on deciduous trees. Fig. 7b shows the variations in numbers of taxa and individuals of such species. However, it is interesting to analyse the histograms relating to beetles specifically linked to particular host-plants (Fig. 7c). Comparisons with previously published pollen diagrams (Woillard, 1975, 1979; Beaulieu and Reille, 1992a) show an excellent agreement between pollen and beetle analysis, for example the agreement between oak pollen and oakdependent Coleoptera; the beetle taxa linked to deciduous Quercus appear in samples that are exactly equivalent to those from which deciduous oak pollen has been recorded. When compared with deciduous tree-dependent taxa, conifer-dependent taxa occur rather later in the sequence; the first individuals are not being recorded until sample 5 (Fig. 7d,e). Here again, the data obtained from palaeoentomological analysis agree perfectly with those obtained from pollen analysis, for example the bark-beetle Platypus oxyurus lives in the trunks of silver fir and its occurrence here matches the pollen curve for Abies (Fig. 8).

.-.I

x

--e

~o

133

$

$

(--

--i

Fig, 6. Total number of taxa (T) and individuals (/) of Coleoptera per sample.

sample 21: the tree-dependent Coleoptera which are common or very common from sample 1 to sample 20 disappear almost totally from sample 21 upwards; only a few isolated individuals of willow-dependent taxa persist. The basal sample 0 does not contain any tree-dependent Coleoptera. As described above, both histograms show two

Aquatic Coleoptera (Fig. 9) This histogram clearly indicates that numbers of aquatic taxa and individuals are remarkably stable throughout the sequence, implying that aquatic environments were continuously present on the site. It is much more informative however to split the frequency histogram into two, one for running-water Coleoptera and another one for standing-water Coleoptera. This reveals a striking contrast between the two categories. Runningwater species are markedly predominant (up to 71 individuals in sample 14) in the forest dominated part of the sequence. From the top of the Saint Germain period, standing-water species become relatively more abundant. It is worth noting that sample 1 (the Riss glacial period) with its cold-adapted species of Coleoptera is also characterized by a predominance of standingwater Coleoptera. Running-water Coleoptera being dependent on

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

24

10 0

60

10 20 0 30 0 110 ~'-~ I ' - ~ i 11 ,1"~ i i I I I [ ~ i 7 ,

i-~ i i [ i l ' l ,

41 40 39

38 37 36 35 34 33 32 31 30 28 27 26 23 22 21 20 19

18 17

16 15

~4 13 12 11 10

9 8 7

6 5

4B 4A 3B 3A 2 1 0

~.~ I'rl I'rl I'~ I'~

O

m03 "13 "X3

O

7-

:7

I'rl Z

m Z

O

O

mr~

I

II

,,,i

I

~1

II

I

Fig. 7. Tree-dependent Coleoptera, number of taxa (T) and individuals (I) per sample, a. Total representation of tree-dependent Coleoptera. b. Deciduous-dependent Coleoptera; c. Coleoptera (number of taxa) exclusively dependent on selected genera or families of trees; d. Conifer-dependent Coleoptea; e. Coleoptera (number of taxa) exclusively dependent on selected species or genera of trees.

highly oxygenated water are unable to survive in other types of environment. It is unlikely therefore that a lacustrine depositional environment prevailed in the interglacial part of the sequence.

Coprophagous Coleoptera (Fig. 10) On the whole this category is poorly represented in the assemblages from La Grande Pile. Dungbeetles are however more abundant in the

P. Ponel/Palaeogeography,Palaeoclimatology,Palaeoecology114 (1995) 1-41 20 m

41

10 0 r-.I

41 40 39 38 37 36 35 34 33 32 31 30 28 2;' 26 23 22 21 20 19 18 17 16 15 14 13 12 11 10 9 8 7 8 5 48 4A 3B 3A 2 1 0

40 39 38 3? 36 35 34 33 32 3t 30 28 21 26 23 Z2 21 20 19 17 16 15 14 13 12 11 10 9 B 7 6 5 4B 4A 3B 3A 2 1 0

700

40 10 0

i i i ~ i i i ~1 i i 1 i 1--,i

I I

25

I l |

II l |

,.n..

m m m m

l|

l,,

80

f i i i i ) i i I

' l•

GP-A6 GP-A~

' GP-A4

| |

|

GP-A3

"n ¢.rl ~10 -..t

¢o F i g . 8. C o m p a r i s o n

of the occurrences

of

Abies

pollen

~--4 t:~ H

E:J

g~g

g

~To

and

Platypus oxyurus.

lowermost and the uppermost levels (where Aphodius holdereri appears) of the sequence (samples 1 and 2, samPles 33-37), that is those corresponding to the glacial episodes, suggesting that many of the herbivorous mammals lived on the open grassland but avoided the thick forest. 4.4. The sequence of coleopteran assemblages and their palaeoenvironmental interpretation In order to make easier the description of the faunal and palaeoenvironmental changes through-

Fig. 9. Aquatic Coleoptera, number of taxa (T) and individuals (I).

out the sequence, the beetle assemblages are grouped into 7 main faunal units numbered from GP-A1 to GP-A7. They have been established on the basis of differences in the specific composition of the Coleoptera assemblage as a whole and not on the significance of certain indicator species. The term unit is preferred to zone because the latter

26

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41 20 rr7 41 40 39 38 37 36 35 34 33 32 31 30 28 2l 26 23 22 21 20 19 18 1? 16 15 14 13 12 11 10 9 8 7 6 5 4B 4A 3B 3A 2 1 0 "S o g 212

?, r-

Fig. 10. Coprophagous Coleoptera, number of individuals.

pearance of cold-adapted Coleoptera, the abundance of tree-dependent and running-water Coleoptera. This unit may be divided in two subunits GP-A2a (rich in deciduous tree-dependent Coleoptera but totally devoid of conifer-dependent Coleoptera) and GP-A2b (many conifer-dependent taxa, mixed with deciduous tree-dependent Coleoptera). --GP-A3: a small unit showing a significant decrease in tree-dependent Coleoptera and the dominance of standing-water Coleoptera over running-water Coleoptera. --GP-A4: this unit is very similar to GP-A2b, but with sporadic occurrence of isolated specimens of the relatively cold-adapted species Potamonectes assimilis. --GP-A5: fairly similar to GP-A3, with a pronounced decrease of tree-dependent Coleoptera, a slight rise of standing-water beetles and a rare occurrence of cold-adapted taxa. --GP-A6: this unit has similar beetle assemblages to that recorded in units GP-A4 and GP-A2b. Tree-dependent Coleoptera reappear but are less abundant in GP-A6 than in GP-A4 and GP-A2b. There is a predominance of runningwater Coleoptera. Cold-adapted Coleoptera are rare. --GP-A7: This large unit is made up of 18 samples. The beetle assemblage shows a great change compared with the lower samples. The tree-dependent taxa disappear almost totally, coldadapted and standing-water species increase in numbers and there is a corresponding decline in the numbers of running-water beetles. This unit may be divided into two subunits GP-A7a and GP-A7b, the latter is defined by an increase in cold-adapted species and an almost total loss of any running-water element.

has been used in a rather different manner to subdivide pollen diagrams.

4.5. Details of faunal units

Main faunal units --GP-AI: characterized by the occurrence of cold-adapted Coleoptera, the scarcity of treedependent Coleoptera and the high number of standing-water Coleoptera. --GP-A2: characterized by the complete disap-

GP-A 1 faunal unit This unit is characterized by the presence of a number of cold-adapted beetle species that do not occur in the Interglacial samples above. Many of these have arctic distributions today (e.g. Diacheila polita, Bembidion dauricum, Amara quenseli,

P. Ponel/Palaeogeography,Palaeoclimatology,Palaeoecology114 (1995) 1-41 Helophorus glacialis, Eucnecosum brachypterum, Hippodamia arctica). Diacheila polita is today widespread in the tundra of northwest North America and Eurasia. In Fennoscandia it is restricted to the eastern part of the Kola Peninsula in northern Russia. It usually lives on the peaty soil of open tundra, sometimes on the margin of pools with Carex or in drier places where Betula nana occurs (Lindroth, 1985). Bembidion dauricum is almost circumpolar. In North America it is restricted to the west of the Hudson Bay and to the Rocky Mountains. In Scandinavia it is known from a very few localities only, where it is limited to the birch zone and to the lower alpine region of the mountains. It occurs mainly on rather dry and sandy soils with sparse vegetation. It is found under stones among dry grass (Lindroth, 1985). Amara quenseli is a circumpolar species present also in Iceland and in Scotland. This rather xerophilous species inhabits open environments such as sandy or gravelly soils with scarce vegetation; it is characteristic of grasslands and heaths in alpine and subalpine regions (Lindroth, 1986). Eucnecosum brachypterum is more widely distributed: British Isles, northern Scandinavia, Central Europe from Germany to Russia, Alps (but not in the French Alps), Transylvania, Bulgaria, Caucasus, Siberia, north Mongolia, North America, mainly in subalpine and alpine regions (Zanetti, 1987). Helophorus glacialis is a boreo-alpine taxon present in Scandinavia and in the high mountains of southern and central Europe. It is the most stenotherm Helophorus species (Angus, pers. comm.). Restricted to glacial snow-melt water, usually in shallow ponds left on black soil behind the retreating ice, sometimes in rocky or clayey ponds (Hansen, 1987). In the southern Europe mountains, Helophorus glacialis is typical of glacier regions, at about 2700-2800 m (Mani, 1968). Hippodamia arctica is a very northern species that is not found south of latitude 65°N; in the southernmost part of its area it is restricted to mountains. According to Strand (1946), Hippodamia arctica was found on Salix scrub as well as on Betula, Empetrum and Arctostaphylos. Like many ladybirds, H. arctica probably feeds upon aphids

27

according to Strand (1946) and Coope and Sands (1966). With this beetle assemblage that corresponds clearly to an arctic tundra fauna are associated some species whose modern distribution and ecology do not fit with such an hostile environment, like Plagiodera versicolor and Gonioctena viminalis that feed upon willows (but not upon dwarf willows according to Koch, 1992), Galeruca tanaceti that feeds upon Compositae (Tanacetum vulgare, Achillea millefolium) or Perileptus areolatus and Bembidion iricolor whose geographical ranges are dominantly southern European today (Lindroth, 1985). Two hypotheses may be put forward to explain the anomalous presence of these two relatively southern species: ( 1) long-distance transport, or (2) the samples covered a climatic transition. The first hypothesis is improbable because La Grande Pile is located in a plain which does not favour long distance eolian transport, as it is the case with some mountain sites (Ponel et al., 1992; Tessier et al., 1993). The second hypothesis is more likely since the top of sample 1 certainly corresponds to a period of very sudden climatic improvement leading to the temperate conditions that prevail in the overlying faunal unit (GP-A2). During most of the GP-A1 unit, the local environment of La Grande Pile may be described as open and extremely cold, similar to an arctic tundra, with a dominantly herbaceous vegetation and scattered shrubs on which Hippodamia arctica may have hunted aphids. The presence of open water is suggested by the dytiscid Potamonectes griseostriatus. However, the occurrence of several xerophilous ground-dependent Coleoptera (Notiophilus, Bembidion dauricum, Amara quenseli) suggests that in places the surroundings of the site may have been rather dry. The top of GP-A1 unit shows a diversification of the willow-dependent fauna, with the leaf-beetles Plagiodera versicolor and Gonioctena viminalis, and the weevil Rhynchaenus saliceti. Running water and small streams are suggested by Perileptus areolatus and Linmius volckmari, the latter occurring as early as sample 1. This climatic improvement mentionned above was not followed by the immediate appearance of trees and tree-dependent Coleoptera suggesting a lag in their response time, possibly

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

28

because of unsufffcient soil maturity or slower rates of spread of woody plants (Coope and Angus, 1975).

GP-A2afaunal unit This unit is characterized by the abundance of deciduous tree-dependent Coleoptera and by the total absence of conifer-dependent and coldadapted Coleoptera. Curculio venosus, C. glandium and Rhynchaenus quercus feed exclusively upon oaks, so does Curculio pyrrhoceras that has its early live history in Cynips quercusfolii galls, a parasitic Hymenoptera living only on oaks. The occurrence of ash is attested by Hylesinus oleiperda and Leperisinusfraxini, while the presence of Acer pseudoplatanus is indicated by Deporaus tristis/ seminiger. This unit contains a vast number of willow, birch and poplar-associated beetles, e.g.

Phyllodecta laticollis, Plagiodera versicolor, Chalcoides fulvicornis, Byctiscus populi, Rhynchaenus rusci and Rhamphus pulicarius. Bark-beetles are represented by many species, including Scolytus multistriatus and S. scolytus that are mainly elmdependent.

Some click-beetle larvae such as

Adelocera murina and Ctenicera pectinicornis feed underground on various roots, whereas others develop in hollow trees. This is certainly the case for Denticollis linearis and Prosternum tessellatum, however the latter can also be found in alpine grasslands (Leseigneur, 1972). A true forest environment is clearly suggested by a number of species such as Pycnomerus terebrans, Colydium elongatum and Dryophthorus corticalis that live in old trees, or the anthribid Brachytarsus nebulosus, a parasite of Lecaninae (tree-dependent coccids) according to Hoffmann (1945). Herba layer taxa are poorly represented, with Bruchidae (the larvae of which develop within the seeds of Fabaceae) and Apion, including Apion cerdo which feeds on the genus

Vicia. Although the riparian Coleoptera are represented only by Stenus, Lathrobium and several species of Donaeia, truly aquatic taxa are very abundant. This group is dominated by runningwater insects such as Dryops, Elmis, Esolus, Oulimnius, Limnius, Normandia and Riolus. Lastly, it should be noted that present in this unit and nowhere else, there is a chafer not iden-

tiffed yet but probably belonging to the genus Triodonta. All the Triodonta species today are confined to southern Europe so the presence of this taxon in this unit suggests particularly mild climatic conditions that were more favourable to such thermophilous Coleoptera than those of the present. Thus the overall environment of the close surroundings during unit GP-A2a may be described as a forested landscape with various deciduous tree species and a poorly developed herbaceous stratum (probably due to the density of trees). The extraordinary abundance of runningwater beetles and the rarity of species of standing water suggests that the sediment was carried into the area by running water or even directly deposited by it.

GP-A2bfaunal unit Deciduous tree-dependent Coleoptera are still abundant, with many of the taxa already recorded in the underlying unit, but also Agelastica alni (specific to alder), Platypus cylindrus (mainly on oak) and Anoplus plantaris (specific to birch). The most important event is the addition of a rich xylophagous fauna made up of many coniferdependent species, mostly scolytids and weevils, such as Hylastes ater, Hylurgops palliatus, Ips

sexdentatus, Polygraphus polygraphus, Pityophthorus pityographus, Pityogenes chalcographus, P. bidentatus, P. trepanatus, Pityokteines spinidens, P. eurvidens, P. vorontzovi, Xyleborus dryographus, X. saxeseni, Rhyncolus elongatus, Magdalis nitida. Other

taxa

are

corticolous,

for

example

Rhizophagus spp, Colydium filiforme, Paromalus flavicornis and Plegaderus vulneratus. According to Lindroth (1985) the large Carabid Calosoma sycophanta also inhabits conifer and deciduous forests, since it is a predator species that exclusively feeds upon tree-dependent moth caterpillars (Lymantriidae, Thaumatopoeidae). Among treedependent Coleoptera, three other species

(Rhysodes sulcatus, Ceruehus ehrysomelinus, Platypus oxyurus) with relict modern distributions are extremely significant from a biogeographical and ecological point of view. Rhysodes sulcatus is today a very rare species recorded from very few localities between Europe and Asia Minor: Bohemia, Slovakia (Freude, 1971), Lombardy,

P. Ponel/Palaeogeography,Palaeoclimatology, Palaeoecology 114 (1995) 1-41

Tuscany, southern Sweden, Haute-Savoie (Ch~tenet, 1986), extreme south-west of France (western Pyr6n6es woodlands) (Tiberghien, 1969). According to Koch (1989), it lives under beech barks, but in the Pyr6n6es, Tiberghien (1969) reports that this species is exclusively found in decaying fir trunks (Abies alba) in which it seems to be localized in a wet and dense deep wood layer, underlying a rotten superficial layer. Ceruchus chrysomelinus is highly characteristic of wet forest in the European mountains, in France it is known from the Jura, the Alps and the Pyr6n6es. According to Paulian (1959), it lives within old decaying stumps of Abies and Picea. It was discovered by Ponel et al. (1992) above 2000 m in the alpine zone of the Taillefer Massif (Is6re, France), in a fossil assemblage of forest beetles in Holocene peat deposits. Platypus oxyurus is today much more restricted than the common species P. cylindrus. In France it is confined to the northern slopes of the Pyr6n6es (Pyr6n6es°Atlantiques, Corbi6res) where it lives at middle altitude. Its European distribution is discontinuous since it is also recorded from Corsica (For& d'Aitone), Calabria, Greece (island of Euboea) and Turkey (no precise locality). This insect is specific to Abies, in the trunks of which it digs deep ramified galleries (Sainte-Claire Deville, 1914; Balachowsky, 1949). The present-day restriction of its distribution suggests that some climatic factors may be involved, since Abies is today widespread in Europe. Furthermore, because this species is unable to develop in branches or in small-size trunks of young trees, it indicates that adult trees were involved. Platypus oxyurus was also discovered in deposits that have been dated from an earlier interglacial (Hoxnian Interglacial), in regions located as far away from its present-day area as Britain is today (Shotton and Osborne, 1965; Coope, 1990). From a purely climatic point of view the occurrence of Rhysodes sulcatus, Ceruchus chrysomelinus and Platypus oxyurus is significant: the known localities for these three species are characterized by a high humidity with heavy rainfall; it is especially the case for Rhysodes sulcatus and Ceruchus chrysomelinus (Herv6, 1951) whose humidity requirements seem to be very high. Thus the presence of these insects provides evidence for

29

the establishment in unit GP-A2b of a mature forest environment, with rather mild and wet climatic conditions.

GP-A3 faunal unit This unit is limited to sample 10 and is characterized by an almost complete disappearance of the tree-dependent insect fauna, which is here represented by only a single specimen of Platypus oxyurus and a single specimen of Rhyncolus elongatus. Bearing in mind the decidedly non-forest character of the assemblage as a whole, it is likely that these specimens really belong either to the overlying or the underlying faunal unit. Platypus oxyurus certainly appears to belong to the underlying unit GP-A2b in which it shows a major but short expansion spanning two samples, with up to at least 33 specimens. This problem may be attributed to inevitable imprecision in the cutting of the cores. This faunal unit shows a striking absence of any cold-adapted taxa in this unit in spite of the almost total disappearance of tree-dependent insects, and thus of the mature forest environment. Two hypotheses can be presented: first the climate deterioration was perhaps too short to allow the northern fauna to establish itself. Atkinson et al. (1987) and Coope (1987) demonstrated the exceptionally fast ability of beetle species to respond to climatic change, so this hypothesis is unlikely to be correct. Moreover the decline of tree-dependent taxa and of the total number of individuals clearly begins as early as the middle of unit GP-A2b, suggesting that, if declining temperatures were responsible for the diminution in the numbers of trees it was by no means a short sharp deterioration. Second, the climatic deterioration may not have been severe enough to permit the incoming of really coldadapted Coleoptera. This second hypothesis is supported by the occurrence of other taxa recorded in this assemblage, which include the riparian species Bembidion guttula, B. aeneum, Stenus, Platysthetus cornutus. Many aquatic beetles are also present, with a dominance of standing-water taxa such as Acilius, Hydroporus, Colymbetes, Hyphydrus ovatus, Helophorus spp. over runningwater taxa. One may conclude from these data that the community that lived around La Grande Pile throughout unit GP-A3 is consistent with an

30

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

open grassland environment, with open birch forest as suggested by pollen analysis. The beetles recorded in this unit do not provide any evidence for a real arctic tundra. The Grande Pile sediments at this time would appear to have been mainly deposited in standing water.

GP-A4 faunal unit This unit, which corresponds to 6 samples, is marked by the re-appearance of a beetle assemblage fairly similar to that described in GP-A2b unit. This assemblage is characterized by a rich fauna of tree-dependent species indicating the presence of both conifer and deciduous trees. Compared with unit GP-A2 some species have disappeared, e.g. Platypus oxyurus. Other species are recorded for the first time, for instance Alosterna tabacicolor (whose larvae develop in the dead wood of various deciduous trees), the scolytids Orthotomicus laricis and O. suturalis which are both parasite of many coniferous trees, Xyloterus domesticus that lives exclusively on various deciduous trees and Kissophagus hederae, another scolytid species that develops in large stems and dying twigs of ivy, Hedera helix (Balachowsky, 1949). Nemosoma elongatum is a today a very rare treedependent Ostomidae indirectly linked to trees. It is not a phytophagous insect but feeds upon larval exsuviae and excreta produced by scolytids. Pissodes pini has larvae that dig superficial galleries under the bark of Pinus sylvestris and Pinus uncinata. Brachonyx pineti feeds exclusively on Pinus sylvestris and has larvae that develop at the basis of the needles and not within the wood (Hoffmann, 1954). Other bark-beetles include Ditoma crenata, a predator on many scolytid species, Pediacus dermestoides, dependent on deciduous trees such as Quercus, Fagus, Acer and Laemophloeus bimaculatus, a rare Cucujid that hunts Dryocoetes villosus inside its own galleries under oak and beech barks (Lefkovitch, 1958). The last five species occurs for the first time in this unit. Most of the phytophagous taxa were already present in the lower units, but this is not the case for some weevils such as Rhynchites nanus (chiefly feeding on Salix, Betula, Alnus), Rhynchites tomentosus (feeding on Salix and sometimes on Populus) and Rhynchaenus avellanae (mainly feeding on

Quercus). Taxa belonging to the genera Phyllotreta, Apion and Bruchidius, provide evidence for the persistence of an herbaceous stratum, and an opening up of the environment is indicated by the ground-beetles Bradycellus ruficollis and Amara lunicollis which are usually found in moors or in forest clearings. The presence of marshes at the site is revealed by the occurrence of hygrophilous species such as Lathrobium terminatum and Stenus, and by many phytophagous taxa such as Eubrychius velutus (which feeds chiefly on Myriophyllum), Lirnnobaris and Donacia. Phalacrus caricis is a small phytophagous Coleoptera feeding on smutted Carex (Thomson, 1958), it is another typical marsh species. A few taxa indicative of standing water are recorded (Agabus,

Enochrus, Anacaena, Coelostoma, Helophorus). However, running-water taxa (Elmis, Esolus, Oulimnius, Limnius, Normandia) are again dominant and represented by large numbers of individuals. The occurrence of the water beetle Potamonectes assimilis is rather more difficult to interpret. Widespread today throughout northern and central Europe, it reaches the northeast of France, such as the lakes in Hautes-Vosges and Alsace (Guignot, 1947), Strasbourg, from where several recent findings are reported by Callot (1990). This insect does not occur in the underlying unit GP-A3 but appears for the first time in GP-A4 unit (in the lowermost and uppermost samples), where the abundance of tree-dependent Coleoptera suggest a temperate climate. The presence or absence of Potamonectes assirnilis may not depend only on thermal conditions, but also on the quality of water, this species being usually found in mountain lakes and springs according to most of the published data. Nevertheless the presence of this insect in GP-A4 unit probably denotes climatic conditions less temperate than those in GP-A2 unit.

GP-A5 faunal unit This unit, which is composed of only two samples, presents similarities to GP-A3 unit. The fall in the number of tree-dependent taxa obviously indicates a marked thinning out of the forest although pollen analysis suggests the continued presence of some birch. The tree-dependent beetle

P. Ponel/Palaeogeography,Palaeoclimatology, Palaeoecology114 (1995) 1-41

taxa do not disappear totally since Dircaea australis/quadriguttatus, Polygraphus polygraphus (on conifers), Deporaus betulae and Anoplus plantaris (on birch), Curculio pyrrhoceras (on oak) persist in very small numbers. The Melandryidae species belonging to the genus Dircaea may also reveal the presence of birch since several living specimens of this extremely rare insect were discovered at La Grande Pile during the coring operations, on erect dead birches covered with Polyporus (tree-dependent fungus). The cold-adapted species here are represented by Potamonectes griseostriatus only, a boreoalpine species whose distribution area expands from northern Europe to the Moroccan Atlas across the mountains of central and southern Europe. It is also known from northern Asia and boreal North America. In the Alps and the Pyrrnres it lives in high-altitude lakes (1800-2400 m) (Guignot, 1947). Some phytophagous genera such as Chaetocnema, Bruchidius, Apion, indicate the presence of herbaceous vegetation. The predator ground-beetle Synuchus nivalis that lives on dry and sandy soils (Koch, 1989) suggests an open environment also.

GP-A6faunal unit This unit is characterized by a moderate increase of tree-dependent Coleoptera assemblages but in rather less profusion than in GP-A2 and GP-A4. Typical forest species from this unit are Acrulia

inflata, Paromalusflavicornis, Laemophloeus bimaculatus, Colydium filiforme, Ceruchus chrysomelinus, Polygraphuspolygraphus, Hylurgops palliatus, Scolytus intricatus, S. ratzeburgi, Strophosoma capitatum, Curculiopyrrhoceras, Dryophthorus corticalis, Brachonyx pineti and Magdalis nitida. These species indicate a return of a truly forested landscape. However, the occurrence of a single individual of the northern swamp weevil Notaris aethiops is of interest here since this insect is only known in France from two localities in Puy-de-D6me (Massif Central) where it lives in peat-bogs and on the border of mountain-lakes and in cold marshes (Hoffmann, 1958), and probably develops on various Cyperaceae. This species possibly provides a hint that the climate was cooler than in either unit GP-A2 or GP-A4. Running-water aquatic genera such as Normandia, Esolus,

31

Limnius, Oulimnius, Normandia and Elmis once more return in abundance, both by the number of taxa and by the number of individuals present, and the still-water species are relatively rare, suggesting again that the sediment was washed in or even deposited by a stream. GP-A 7faunal unit This long and rather homogeneous unit extends over 18 samples. It is characterized by the almost complete disappearance of tree-dependent taxa except for the weevils Rhynchaenus foliorum and Anoplus plantaris, whose larvae mine the leaves of many willows (including dwarf-willows) and of birch (including Betula nana), respectively. Phytophagous taxa from the herb layer are extremely rare, they are represented, for example, by Micrelus ericae, exclusively feeding on Calluna vulgaris and Erica tetralix. The most striking feature of this faunal unit is the disappearance of the entire tree-dependent fauna and the appearance of a number of very cold-adapted species whose modern distribution is mainly restricted to northern Europe, Fennoscandia and the arctic regions of Russia, most of which are able to live above the tree-line or even obliged to do so. This cold fauna consists of ground Coleoptera (Carabidae) and aquatic Coleoptera (Dytiscidae, Hydraenidae, Hydrophilidae). Elaphrus lapponicus is only known today from northern England (one locality), Scotland and Fennoscandia. According to Lindroth (1985), this species occurs in the birch and the upper conifer region, and also in the lower alpine region. It inhabits small marshes with Carex, Eriophorum, mosses, etc. Also according to Lindroth, it is sometimes found associated with Diacheila arctica, probably because these species have similar ecological requirements; both species occur together in sample 36. This shows the extent to which the ecological requirements of Coleoptera remained stable despite the great climatic and biogeographical upheavals they have to withstood during the glacial/interglacial cycles. Unlike many alpine or subalpine species, E. lapponicus is occasionally found in relatively warmer places, which is consistent with its preference for sunny microhabitats (Lindroth, 1985). Diacheila arctica inhabits

32

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1 41

northern Scandinavia, the Kola peninsula and the north of Russia. It is an hygrophilous species living on lake shores and muddy depressions from the upper conifer zone to the lower alpine zone. Its favourite habitat seems to be fens with Carex, Eriophorum, Scirpus, Juncus and mosses (Lindroth, 1985). Diacheilapolita, last found in GP-A1, reappears in this unit. The circumpolar water beetles are represented by many taxa. Agabus arcticus is a carnivorous water beetle that lives in Sphagnum pools on peatbogs, and on mossy lake shores in the north of the British Isles and in Fennoscandia (BalfourBrowne, 1950). Colymbetes dolabratus is today known from the north of Fennoscandia, the north of Russia, Siberia, North America, Greenland and Iceland (Coope, 1968). It lives in standing water or in slowy moving streams. Helophorus sibiricus is an holarctic species, widespread in the palaearctic region extending from the north of Fennoscandia to Mongolia and China. In Fennoscandia it lives on river margins, but it can also be found in shallow grassy pools, especially in the eastern part of its area (Angus, 1973; Hansen, 1987). Helophorus oblongus is known today from North America and Siberia; according to Angus (1973) this species is probably widespread in the Siberian taiga and tundra. In the southern part of its area this species is apparently restricted to forest although it is common elsewhere in grassland pools (Angus, 1973). Helophorus glacialis, a highly stenothermic species whose ecological requirements were described above (GP-A1 unit), reappears in almost every sample of GP-A7 unit. Among the terrestrial insects, the silphid Pterolomaforsstroemi, is found in Scandinavia and also on high mountains of Central Europe. It lives in mosses, under gravel and pebbles, and along small streams and torrents where it preys upon molluscs (Chgttenet, 1986). The Omaliinae are common, with Euenecosum brachypterum previously found in GP-A1 unit, here accompanied by three other species characteristic of this type of assemblage (Coope, 1975; Taylor and Coope, 1985): Pycnoglypta lurida (northern Europe, in the south toward Denmark and northern Germany; Siberia, North America; in wetlands and marshes

according to Zanetti, 1987), Boreaphilus henningianus (Scandinavia, north Russia and the Harz mountains; under plant debris, in wet mosses and grassy marshes according to Coope, 1962) and Boreaphilus nordenskioeldi (isolated localities in the north of North America and Asia; its ecological requirements are similar to those of the previous species according to Coope, 1975). Another staphylinid, the coprophilous Oxytelus gibbulus, is found living today only in the Caucasus mountains and maybe also in eastern Siberia (Hammond et al., 1979). The oldest fossils for this species are dated from ca 600,000 yr B.P. (Waverley Wood, Warwickshire) (Shotton et al., 1993). It was sporadic in its occurrence at several sites in England dating from the mid-WOrm (Devensian, Weichselian). O. gibbulus was the most common staphylinid in England about 200,000 yr B.P. and was regularly associated with mammoths. Simplocaria metallica is widely distributed in Scandinavia and in few isolated localities in Central Europe. It is exclusively a moss feeder (Coope, 1961). Aphodius holdereri is today an asiatic species (a Tibetan endemic, known from lake Ko Ko Nor region in the north to the northern slopes of Himalaya, according to Coope, 1973). It appears in three samples only: 33, 36 and 37. The rare occurrences of these exotic species might be of use as stratigraphical markers permitting correlation of the Grande Pile sedimentary levels with the British sediments where similar fossil assemblages occur within a restricted time period (Coope et al., 1961; Coope, 1969). According to Coope (1979), Aphodius holdereri is "by far the most abundant large dung-beetle in fossil assemblages from the British Isles that date from the middle of the Last Glaciation". The occurrence of this and many other exclusively Asiatic species in western Europe at this time (Coope, 1994) indicates a climatic episode whose modern analogues may be found in Tibetan mountains, at altitudes from 3000 to 5000 m. As regards aquatic Coleoptera, running-water species decline gradually in the GP-A7 unit and are eventually replaced by standing-water species in the five upper samples, suggesting that the Glacial sedimentary environment was quite different from that of the Interglacial.

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

It is convenient to divide GP-A7 unit into two subunits: --GP-A7a (samples 21-35) is characterized by the almost complete disappearance of treedependent species, the first indications of the re-appearance of the most cold-adapted Coleoptera, the persistence of running-water beetles and of many taxa that are rather catholic in their ecological requirements. --GP-A7b (samples 36-41) is characterized by a rise in arctic-alpine Coleoptera, the complete disappearance of running-water beetles and an impoverishment of ubiquitous species. There can be little doubt that this fauna indicates a further increase in coldness. It is thus interesting to note that the breccia level contained in sample 38 and 39, thought to be the result of frost-shattering phenomena in shallow water, occurs just above the marked rise of many cold-adapted beetles, including the Tibetan dung-beetle Aphodius holdereri, in samples 36 and 37. This may well be the result of the progressive intensification of the cold in which the increasingly harsh conditions allow a rich cold-adapted fauna to become established. Later, the thermal conditions must have become so severe that even many of the cold-adapted species eventually succumbed. The environment was by now similar to a polar tundra with an impoverished fauna highly adapted to extremely cold conditions. This interpretation is also supported by the occurrence of the crustacean Notostraca Lepidurus arcticus Pallas which appears in huge numbers (223 mandibles were

33

recorded) in samples 38 and 39. The ecology and climatic requirements of this tadpole shrimp is summarized by Taylor and Coope (1985). It is today mostly found north of the Arctic circle in very shallow water of impermanent water bodies and is able to stand very cold conditions. The climatic interpretation of sedimentological and biological data does not correspond exactly to the timing of the early land-ice glacial maximum dated between 50,000 and 30,000 yr B.P. described by Seret et al. (1990). By comparison (Fig. 5) with the pollen zones and dates provided by Beaulieu and Reille (1992a: figs. 1 and 5) the cold maximum inferred from arthropod assemblages may be dated roughly just after ca 30,000 yr B.P. As a whole, the insects of the GP-A7b unit indicate extremely harsh climatic conditions, similar to those prevailing today in arctic tundra or in the highest mountains.

Summary of the correlation of the coleopteran succession with pollen and isotope stratigraphy The faunal units established here upon Coleoptera and those based on pollen zonation (Beaulieu and Reille, 1992a) and isotope zones (Woillard and Moock, 1982) may be readily correlated (Table 2). The two very cold periods GP-A1 and GP-A7 correspond to the termination of the penultimate glacial period (Linexert) and to the last glacial period (Pleniwiirm+Late Warm of Beaulieu and ReiUe, 1992a), respectively. Isotope stage 4 does not appear in the palaeoentomologlcal records; it may correspond to the non-analyzed

Table 2 Correlations between insect, pollen and extrapolated marine isotope zones Faunal units

Pollen chronozones Beaulieu and Reille (1992a)

Isotope stages Woillard and Moock (1982)

GP-A7b GP-A7a samples missing GP-A6 GP-A5 GP-A4 GP-A3 GP-A2b GP-A2a GP-A1

Wiirmian Pleniglacial Wttrmian Pleniglacial Lower Wiirmian Pleniglacial Saint Germain II Mrlisey II Saint Germain I Mrlisey I Eemian Interglacial Eemian Interglacial Riss (Linexert)

2 3 4 5a 5b 5c 5d 5e 5e 6

34

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

samples 24 and 25. The forested periods GP-A2, GP-A4 and GP-A6 correspond to the Eemian, to the Saint Germain I and to the Saint Germain II, respectively. They were interrupted by two treeless episodes that are equivalent to Mrlisey I and Mrlisey II and show that trees were greatly reduced. As observed from pollen analysis by Pons et al. (1989), there is no entomological evidence for an extremely cold phase at La Grande Pile during these two last episodes. These correlations are in broad agreement with the palaeoenvironment implications derived by Beaulieu and Reille (1992a) from the pollen analysis of core XX.

4.6. Discrepancy between water beetles occurrences and the possible lacustrine origin of La Grande Pile Fig. 9 clearly shows the great abundance of running-water Coleoptera in the samples attributed to the Eemian and to the Saint Germain I and II, with two declines in abundance in samples 10 and 17 corresponding to M~lisey I and M~lisey II, respectively, which are short treeless spells interpreted as reflecting moderate climatical deteriorations according to both pollen and insect analysis. Though occurring in low numbers, runningwater Coleoptera continue to be present from sample 22 to sample 36 and then disappear totally from sample 37 upwards (beginning of the Late WOrm). The lowermost samples 0 and 1 (Linexert, termination of the Rissian glaciation) also contain very few running-water beetles. The occurrence of standing-water beetles is virtually the reverse of this; large numbers in sample 1, small numbers in samples 2-23, quite large numbers in 26-37, then large numbers again in the uppermost sample 41. As regards Lepidurus arcticus, the abundance of this crustacean in samples 38 and 39 indicates that very shallow water was present during the Upper Wiirmian Pleniglacial. It is noteworthy that the insect data are supported by some pollen data. Thus Fig. 1 in Beaulieu and Reille (1992a) shows that shallow and calm water plants such as Isoetes are practically absent until the end of the Eemian. The Cyperacaeae, mostly shallow-water plants, are well represented at the end of the penultimate glacial period, then disappear until the Upper Wtirmian

Pleniglacial except two short occurrences in Mrlisey I and Mrlisey II (samples 10 and 17). Furthermore the isolated occurrences of Sphagnum in several points of the sequence indicate that exposed places were available and favoured the development of mosses. Lastly, the Ranunculus batrachium group appears just at the beginning of the Pleniglacial and increases gradually until the end of the sequence, probably in association with the infilling in of the site. With regards to the diatoms, Cornet (1988) states that sedimentological data suggesting a high water level (several metres) are not in agreement with algal data which indicated rather shallow water at the end of the Eemian. It is tempting to interpret the variations of running- and standing-water beetles as reflecting the hydrological regime that prevailed at La Grande Pile during the two last climatic cycles. The large numbers of running-water beetles combined with the scarcity of lake or pond taxa certainly indicate the presence or even the predominance of small streams during the Eemian and other forested periods on this site. These streams would have carried sediments sporadically into the depositional environment and the rates of accumulation are unlikely to have been constant. Estimations of rates of environmental changes based on the assumption of constant sedimentary input are thus unlikely to be reliable. The presence of such streams during these periods has further significant implications since the mire is located today on a slightly elevated interfluve plateau as stated by Cornet (1988) and by Woillard (1975): "Le bassin d'alimentation de la tourbi~re est tr~s limitr: aucun cours d'eau ne l'alimente et elle ne poss6de que deux exutoires de vidange (...)". The hydrology of the site must have been rather different in the past. La Grande Pile may not always have been in such a raised position above the alluvial plains of Ognon and Lanterne. Today the depression is only 20 m higher than the plains mentioned above, some geomorphological and hydrodynamical changes must have occurred contemporaneously with the climatical changes, especially during the Wiarmian deglaciation. The replacement of running-water Coleoptera by standing-water Coleoptera during cold episodes

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

(Riss, Mrlisey I, Mrlisey II, Pleniglacial) suggests a fundamental environmental change and the replacement of a running-water environment by a marsh or a very shallow lake environment. During Pleniglacial times the depth of the water must have been very slight (occurrence of Lepidurus arcticus) to permit the frost disturbance of the deposit. Maybe a decrease in temperature and precipitation caused the interruption in water flow. Analysis of group B core series, extracted close to the margin of the Grande Pile depression 150 m away from group A, might provide some new insight into this intriguing problem.

4. 7. Climatic reconstruction One of the most important aspects of modern Quaternary entomological analysis lies in the fact that many fossil sclerites can be identified to the species level. Furthermore these species appear to be identical to living specimens. A mass of biogeographical, ecological and climatical data can thus be obtained from the rich literature that has been devoted to Coleoptera during the two last centuries. Moreover a recent computerized palaeoclimate reconstruction method based on Coleoptera has been recently developed: the Mutual Climatic Range Method, or M.C.R.M. (Atkinson et al., 1986), used to obtain the climatic reconstruction presented in this paper. This method enables quantitative palaeoclimatic reconstructions to be made that are not matched by any other palaeoecological approaches to continental environments. It is based on the range of climates corresponding to the total geographical area occupied today by the species present in a fossil assemblage. The climatic estimate proposed for the whole assemblage is given by the overlap of the climatic ranges of the species identified in this assemblage. It is important to note that the precision of this reconstruction method relies in part on the identification of the taxa to species level. Moreover, the taxa included in the M.C.R.M. data-base only include Coleoptera that are thought to be not directly dependent of higher plants: these insects are especially significant from a palaeoclimatic point of view because their distribution is not likely to be linked to the occurrence of particular host-plants but

35

imposed by climatic factors. Usually, the food chains of such species ultimately depend on microorganisms (algae, unicellular fungi,...). At La Grande Pile this category is mainly represented by aquatic, riparian, terrestrial and coprophilous beetles belonging to the families Dytiscidae, Hydrophilidae, Staphylinidae, Carabidae and Scarabaeidae families (e.g. Elaphrus

lapponicus, Diacheila arctica, Diacheila polita, Bembidion dauricum, Amara quenseli, Potamonectes griseostriatus, Potamonectes assimilis, Agabus arcticus, Colymbetes dolabratus, Helophorus glacialis, Helophorus sibiricus, Helophorus oblongus, Pteroloma forsstroemi, Pycnoglypta lurida, Eucnecosum brachypterum, Boreaphilus henningianus, Boreaphilus nordenskioeldi, Simplocaria metallica, Hippodamia arctica, Aphodius holdereri, Notaris aethiops; Fig. 11). Many of these cold-adapted taxa are only able to complete their biological cycle under very harsh thermal conditions and are of great value for climatic reconstructions. For example, it is the case for the circumpolar groundbeetle Diacheila polita or the Tibetan dung-beetle Aphodius holderei which cannot tolerate average summer temperatures much higher than about 10°C; on the other hand they can withstand and even seem to prefer average winter temperatures as low as - 1 2 / - 2 4 ° C (Coope, pers. comm.). The Mutual Climatic Range Method has been used for each coleopteran assemblage obtained from the 41 Grande Pile samples (Fig. 12). This diagram shows clearly an early cold period represented by samples 1 and 2 during which the climate was also very continental. The Eemian warming took place from sample 2 upwards after which temperature fluctuates slightly but remains, on average, quite high until sample 22. After a warm maximum of summer temperature centred on samples 5, 6, 7 (broadly synchronized with the Taxus phase in Fig. 5 and overlapping the peak of Platypus oxyurus), the second half of the Eemian shows slowly declining temperature. At La Grande Pile therefore no indication is found for any cold periods "to levels more typical of the mid-glacial period" during the Eemian (as reported by the G.R.I.P. project, 1993); this result is in accord with Boulton (1993). The modest climatic deterioration of Mrlisey I is clearly visible in sample 10,

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

36

10 iTM

4, !I"i'

,o 39

I

10 i-~

10 i-'n

il

'l,ilE"

t I

i

38

36 35 a,t

.

33 a2 31 30 2~ 21 26 ~3 ~2

.

.

.

.

.

.

.

.

.

.

.

.

.

.

10 0 30 F-~II,

.

.

.

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.

.

.

.

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,

~0 19 18

16 15 13 12 11 10 9 ? 5 4B

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~ 2

m~

Fig. 11. Cold-adapted Coleoptera, number of taxa (T) and individuals (I): and occurrences of selected cold-adapted Coleoptera (number of individuals). it does not indicate a return to the same level of coldness as sample 0 and 1. In contrast, the M61isey II episode does not appear clearly. Samples 21-22 correspond to a fundamental climatic deterioration which can be attributed to the beginning of the Pleniglacial. Temperature remains low until sample 41 (probably final Pleniglacial), except in samples 23 and 34 that appears to represent short warming.

According to Fig. 11, these events would seem to be related to a reduction in the numbers of the cold-adapted elements rather than the reappearance of therrnophilous species. These events are not clearly recorded by pollen analysis (Fig. 5) and illustrate the rapidity of response of Coleoptera in comparison with the reaction time of the vegetation. Sample 34 corresponds to the

P. Ponel/Palaeogeography,Palaeoclimatology, Palaeoecology114 (1995) 1-41

37

'C

?ii i ? i ? \illii i? :ii? illllli ??/?Lil li????/????/??i • ° I ' 2 l,.[3bl,.14b 516 I' 1~ 1 9 ,o ,,I,2f,~l,,l,s[,~

,el,8,912o 2,12212~1~612,1291,o1,,1,21,31~,1,5 ~61~71,,1~91,o1,,

b G,,-,~,

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GP-A7b

M II

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LW

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I~

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E

E

M I

SG I

"C

+ 10'

iii iif:iiliiii:i-

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!!i/ii!ili!!!i!!!!

-10,

.20 ¸

- 30

............................

,..2122--

;21212;

,°o12222-2221h

.............................................

Fig. 12. "Mutual climatic range" climatic reconstruction for the Coleoptera from La Grand Pile (°C). The horizontal axis is approximatelyequivalentto time B.P.a. Insect samples,b, Entomologicalbiostratigraphicunits, c. Pollenchronozonesof Beaulieu and Reille (1992a), L= Linexert(or Riss), E= Eemian, M I= M61iseyI, M H= M61iseyII, SG I= Saint Germain I, SG H= Saint GermainII, PW= Pleniwllrm,LW= LateWOrm. upper half of palynozone 8f, the termination of which (sharp rise of Pinus curve) is dated ca 34,000 yr B.P. (Fig. 5). This short warming may be related to one of the episodes of relatively mild climate conditions described during the mid and late parts of the last glaciation by Coope and Angus (1975), Coope (1977) and more recently by Johnsen et al. (1992). The coldest part of the climatic reconstruction for the mean temperature of the warmest month corresponds to sample 38 (dated just after 30,000 yr B.P. according to Fig. 5) and (together with sample 39) to sediments that shows signs of frost activity, with an impoverished and extremely cold-adapted Coleopteran fauna (e.g. Diacheila polita, Boreaphilus nordenskioeldi) and the occurrence of the crustacean Lepidurus arcticus, just after the occurrence of the Tibetan element Aphodius holdereri in samples 36 and 37.

No other climatic episodes are clearly recorded, either because such episodes are too faint to be detected by the insect record, or because of possible sedimentary hiatuses. However, no such sedimentary gaps were recognised during pollen analyses (Woillard, 1975; Beaulieu and Reille, 1992a). An analysis of climatic reconstructions based on both Coleoptera and pollen evidence is proposed by Guiot et al. (1993); the climatic data obtained from La Grande Pile are discussed by these authors.

5. Conclusions This unique continuous beetle record in continental Western Europe gives a detailed picture of the drastic changes in the insect fauna that correlate with changes that affected the flora and the

38

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1 41

vegetation. These changes clearly reflect the major climatic changes that took place within the last glacial cycle, i.e. the last 140,000 years, in this part of the world. The steep warming of the transition RissEemian lead to the replacement of a cold-adapted fauna and of a tundra-type environment by a treedependent insect fauna made up dominantly of deciduous tree-dependent species in the first part of the Eemian interglacial. In the second part of the Eemian this tree-dependent fauna was largely dominated by conifer-dependent taxa. After a warm episode that overlaps partly the peak of the Mediterranean Platypus oxyurus, a progressive cooling ended in a change of the environment during the Mrlisey I event when the tree-dependent species disappeared but were not replaced by very cold-adapted taxa The landscape was then similar to an open grassland rather than a true tundra. The return of a forest environment is proved by the development of a rich tree-dependent fauna that thrived in the next period, namely Saint Germain I. This fauna showed some similarities with that of the second part of the Eemian (however Platypus oxyurus is now absent) and must be likened to a ta~ga-type insect fauna. After another cool event, namely Mrlisey II (which appears as a kind of replica of Mrlisey I), the Saint Germain II corresponds with a brief re-appearance of the forest fauna; it was abruptly interrupted by the faunal and environmental upheavals that marked the transition with the Pleniglacial and the establishment of a very cold-adapted insect fauna, concurrently with a tundra-like landscape. A period of severe cold occurred towards the top of the sequence (sample 38) just after 30,000 yr B.P., as indicated by biological and stratigraphical evidence. Concurrently with these fundamental climatic changes, the hydrological regime at the site seems to have undergone some important variations: the mild forested periods correspond to large numbers of running-water-dependent Coleoptera (Dryopidae) that live in highly oxygenated water only. Thus, this faunal community seems to indicate an hydrological regime characterized by the presence of streams. In contrast, the cold periods are characterized by the replacement of this fauna

by standing-water Coleoptera that denote the presence of a lake or pool environment. The exact mechanism of these hydrological variations is not yet fully understood but raises interesting questions about the steadiness of the sedimentation at La Grande Pile.

Acknowledgments First of all, I would like to thank G.R. Coope who took an essential part in the realization of this work. I am grateful to the "Commissariat h l'l~nergie Atomique" who supported this study and to V. Andrieu, J.-L. de Beaulieu, M. Campy, F. David, C. Goeury, J. Guiot, P. Guenet, M. Reille, W. Safar, G. Seret and his team for help and advice. Figs. 6-11 were drawn with the program GPAL3 created by Goeury (1988). Numerous helpful comments on an early version of the manuscript were made by H.J.B. Birks, G.R. Coope, P. de Deckker and G. Lemdahl; Michrle Pellet helped improve the final form of the English text.

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