E L S E V I E R Palaeogeography, Palaeoclimatology, Palaeoecology 114 (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 con- tinuous 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) analy- ses by Louis and Smeets (1981), Louis et al. (1981), Louis and Ermin (1983), Louis and Peters (1983), Louis et al. (1983), Cornet (1988); paly- nology 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 pre- cise information and permit quantification of cli- matic 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

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 sedi- ments. 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 hydro- graphic network (Seret et al., 1990) and no tribu- tary 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 south- wards, 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 glaciolac- ustrine tills of the Linexert glaciation at the basis of the sedimentary sequence. The absence of

~ "~'-~ PARIS ~

EPlNAL I

FRANCE

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

La Grande Pile

BESANCON

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41 3

1 2 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 exten- sion 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 com- munity 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 tri- foliata 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 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

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 com- plete 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.

~ drain LINEXERT " ~ \

,/ i'

" i "I\

N ~

o E }

$ i i

lOOm / / /"

,,,,, .,,.._.~f"~"~-?~ ~.

I I ,

i~.,,~, I ~ '"'" /7';' /I ......... ~.~ ................ ' I A l/ '~

'

o 5o ", % t','.

"' I~ J " , '< t I

', . . . . . - t/ , > , 7

Ili ditch/~', ..-~#

- - 4 .......... S';' ...// © location of the cores ana l ys~y - -W-~LARD ( 1 9 7 ~ - - - ~ % ] I ~ D > ~ . - ~ "~'~ " ,lir location ofthe core XX analysed by BEAULIEU & REILLE (1992a III ~ ' ,~ i ST GERMAIN

I palaeoentomological corings of June 1990 = ~-~ (A : cores 90-1 to 90-8 ; B : cores 90-9 to 90-14)

- - - peat-bog limit • boundary stones

forest tracks

I ' ' '¸' t t . . . . . .

ST GERMAIN

Fig. 3. Location of the cores on the Grande Pile peat-bog.

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 sedimento- logical 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 sedimenta- tion 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

The chronology of this sequence (Fig. 5) is established by correlation with previous studies but is mainly derived from the interpretation pro- posed 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

10.

11.

12-

13-

14-

15,

16.

17.

18.

19.

6 - ~ 4 1

7 - 3 8

" 3 7

35

3 4 9-

:~,~= 33

........... 2 9

...... 2 6

2 5

i;';'-'S' ¢ 2~

;;,; ...... 16

.... 15

, ,v 14 13:

. . . . 12 1'I

. . . . . 1 0

9 8 \M /

7

; ' , ; ( . . . . . . 6

v~ 4b . . . . . . . . . 4a:

~, 3a ~'-"-- - . 2 " - - - - - ' 1 : _- -_~

~ . ~ _ ~ c)

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.

,-? ,.c 1£ r~ m g g g o o o

m m m m m g g g g g g o o o o o o g g g ~ ~ g

I~ ~I 0 a'l" I N ~ ' I d i ' I 'O'S I I i ,

1 NlVMI~l~9 L S

p i ~ ~t

i I i i i i

,i

i ,

R a l l ~ l a x

-+++

g 8 ~ o "o®

o ~ +-

t . . . .

I

I! ........ i . . . . . . . ~+

+ I +++ + + ....... i

, , t r I i

I[ '

j,.,,LII , ,~ . . . . J . . . . . ill, + i I - j J r +~I.II~I. I h+l

I ' ' ' [+ , ,

+ + . ~ + ' , +, , I +

, . + -+ : ~ . . . . . , + + . ~ . . . - + ~ ~ + - + - - - J + ~ - - ~ + + - , . . . . . . +.+-4~+_..

g

, i

i

B 0

× ×

~ , .0

0

J j I [ [ J ! t ] ] .r ! ] I I [ ! I ! U I I i I+ ~ ! [ ! I + I J J f ! r itHitt ~ ' ~ .... ......../ . . - .............................+i...........'./.....~............i/ .........+...~:::i~..-.............../ ...- /. / ./-. . .- ..- / : . / ...- .... /..../~-.~..~>. + >

i i ii ++i iiiiiiii?.i.. !:.:.!!i!i))))) !! :: iiiiiiiiiiiiiii iiii~ iiiii .... / / / / ~:.:~:: • /::/." //' / : : / : / :

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41 7

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 nec- essary. Water with macroremains mixed with dis- aggregated 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 contamina- tion 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-

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 coprophi- lous 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 require- ments. Their modern distribution range from boreal, boreo-montane to mediterranean. The suc- cession of these species clearly reflects the large- scale 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 dwarf- willow-dependent beetles. The histograms of taxa and individuals (Fig. 7a) show clearly a dramatic change at the transition between sample 20 and

Tab

le 1

L

ist

of c

oleo

pter

a sh

owin

g th

e m

inim

um n

umbe

r of

ind

ivid

uals

rec

over

ed f

rom

eac

h sa

mpl

e. N

omen

clat

ure

and

taxo

nom

ic o

rder

fro

m L

ucht

(19

87).

Spe

cies

whi

ch

are

not

mem

bers

of

the

pres

ent-

day

Fre

nch

faun

a ar

e m

arke

d w

ith

an a

ster

isk

(*)

0 1

2 3A

3B

4A

4B

5

6 7

8 9

10

11

12

13

14

15

16

17

18

19 2

0 21

22

23

26 2

7 29

30

31

32

33

34

35

36

37

38

39 4

0 41

CA

RA

BID

AE

C

icin

dela

sp.

C

alos

oma

syco

phan

ta

(L.)

C

arab

us c

lath

ratu

s L

. C

. ca

ncel

latu

s C

. ni

tens

L.

C.

arve

nsis

Hbs

t.

1 C

arab

us s

p.

Neb

ria

gyll

enha

lli

(Sch

rnh.

) N

ebri

a sp

. N

otio

phil

us s

p.

1 1

*Ela

phru

s la

ppon

icus

G

yll.

E

laph

rus

ripa

rius

(L

.)

* Dia

chei

la a

rcti

ca

(Gyl

l.)

*D. p

olit

a (F

ald.

) 1

Lor

icer

a pi

lico

rnis

(F

.)

Dys

chir

ius

glob

osus

(H

bst.

) P

eril

eptu

s ar

eola

tus

1 (C

reut

z.)

Tre

chus

sec

alis

(P

ayk.

) T.

rub

ens

(F.)

1

T. o

btus

us E

r./

1 4-

stri

atus

(S

chrk

.)

Tre

chus

sp.

B

embi

dion

bi

punc

tatu

m (

L.)

B

. ni

tidu

lum

(M

arsh

.)

*B.

daur

icum

(M

ots.

) 1

1 B

. (T

este

diol

um)

sp.

1 B

. sc

happ

eli

Dej

. B

. ae

neum

Ger

m.

R

unic

olor

Cha

ud.

2

121

1 1

1 1

1 1

1 1

2 1

11

1 1

1 1

1 1

1

1

1

11

1 3

11

1

1 1 11

1

1 1

1

e~

B.

guttu

la (

F.)

Bem

bidi

on s

p.

* Pat

robu

s as

sim

ilis

Cha

ud.

Pat

robu

s sp

. B

rady

cellu

s ru

ficol

lis

(Ste

ph.)

Poe

cilu

s le

pidu

s (L

eske

) P

oeci

lus

sp.

Pte

rost

ichu

s pu

mili

o (D

ej.)

P.

dilig

ens

(Stu

rm)

P.

vern

alis

(Pa

nz.)

P

. ni

grita

(Pa

yk.)

P

. ni

ger

(Sch

all.)

P

. m

elan

ariu

s (I

ll.)

P

. m

adid

us (

F.)

P

tero

stic

hus

sp.

Aba

x sp

. Sy

nuch

us n

ival

is

(Pan

z.)

Cal

athu

s m

elan

ocep

h-

alus

(L

.)

~4go

num

eri

ceti

(Pan

z.)

A.

mue

lleri

(H

bst.)

A

. ful

igin

osum

(P

anz.

) A

gonu

m s

p.

Am

ara

luni

colli

s (S

ehdt

e)

A.

quen

seli

(Sch

~nh.

) A

mar

a sp

. A

. (C

yrto

notu

s) s

p.

HA

LIP

LID

AE

B

rych

ius

elev

atus

(V

anz.)

H

alip

lus

sp.

DY

TIS

CID

AE

H

yphy

drus

ova

tus

(L.)

C

oela

mbu

s im

pres

so-

punc

tatu

s (S

chal

l.)

Hyd

ropo

rus

palu

stri

s (L

.)

1 1

1 1

1 1 1

1 4

1 2 1 1

1

1

1 1 1 113

2 2

¢ 7"

Tab

lel(

cont

inue

d)

0 1

2 3A

3B

4A

4B

5

6 7

8 9

10

11

12

13

14

15

16

17

18

19 2

021

2223

26

2729

30

31

32

33

34

35

36

37

38

39

40

41

2 1

22

43

1 I

11

19

1

34

14

11

1

I1

12

511

Hyd

ropo

rus

sp.

Pot

amon

ecte

s gr

iseo

stri

atus

(G

eer)

P

. as

sim

ilis

(Pay

k.)

Pot

amon

ecte

s sp

. A

gabu

s bi

pust

ulat

us

(L.)

A

. st

urm

i (G

yll.

) *A

. ar

ctic

us (

Payk

.)

Aga

bus

sp.

llybi

us s

p.

Rha

ntus

sp.

C

olym

bete

s fu

scus

(L

.)

*C.

dola

brat

us

(Pay

k.)

Col

ymbe

tes

sp.

Gra

phod

erus

sp,

A

ciliu

s sp

. D

ytis

cus

sp.

1 1

I 1

1

1 1

1 1

GY

RIN

IDA

E

Gyr

inus

min

utus

F.

1 G

. aer

atus

Ste

ph.

?1

1 3

G. a

erat

us

2 1

1 1

Step

h./m

arin

us

Gyl

l.

Gyr

inus

sp.

1

1 1

1 1

1 1

RH

YS

OD

IDA

E

Rhy

sode

s su

lcat

us

(F.)

HY

DR

AE

NID

AE

H

ydra

ena

gr.

ripa

ria

Kug

. H

. gr

acili

s H

ydra

ena

sp.

Och

theb

ius

gr.

fove

olat

us

Ger

m.

Och

theb

ius

sp.

2 1

1 3

34

1

2 i

2

43

t

1

2

22

1

1 1 2

1

1

1

1 1

1 1

I 1

1 I

I

,2 4~

4~

Lim

nebi

us s

p.

Hyd

roch

us s

p.

Hel

opho

rus g

rand

is Il

L

* H.

sibi

ricu

s (M

ots.

) H

. aq

uatic

us (

L.)

*H

. ob

long

us L

eC.

type

11

. gla

cial

is V

illa

H

. br

evip

alpi

s Bed

el

H.

?fla

vipe

s F.

H.

gr. m

inut

us F

. H

elop

horu

s sp

. C

oelo

stom

a or

bicu

lare

(F

.) Sp

haer

idiu

m s

p.

Cer

cyon

mel

anoc

epha

- /u

s (L

.)

Cer

cyon

sp.

M

egas

tern

um

bole

to-

phag

um (

Mar

sh.)

C

rypt

ople

urum

m

inu-

tu

rn (

F.)

H

ydro

bius

fusc

ipes

(L

.)

Ana

caen

a sp

. La

ccob

ius

sp.

Eno

chru

s af

finis

( T

hunb

. )/c

oar c

tatu

s (G

r.)

Eno

chru

s sp

. C

haet

arth

ria

sem

inu-

lu

m (

Hbs

t.)

HIS

TE

RID

AE

P

lega

deru

s vu

lner

atus

(P

anz.

) P

arom

alus

fla

vico

rnis

(H

bst.

) H

iste

r sp

.

12

13

1 16

1

1 1

1 1

22

1 2 1

1

2

1

1

1

2 1

11

22

11

2

1 1

1 1

2 5

47

4

1 1 I

34

14

3

21

1

12

12

2

3 3

1

1 11

11

7 3

53

1

6 2

43

1 1

1

1 12

2

1 7

.¢

E"

xt~

SIL

PH

IDA

E

Than

atop

hilu

s sp

. P

tero

lom

a fo

rsst

roem

i (G

yll.

) N

ecro

phor

us s

p.

1

1 1

2 1

1

Tab

le 1

(co

ntin

ued)

0 l

2 3A

3B

4A

4B

5

6 7

8 9

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

G

CA

TO

PID

AE

C

hole

ra s

p.

1 C

atop

s sp

. 1

1 I

1

LIO

DID

AE

G.

sp.

1 2

SC

YD

MA

EN

IDA

E

Neu

raph

es s

p.

1 1

I 3

2

PT

ILID

AE

A

crot

rich

is s

p.

I 1

1

ST

AP

HY

LIN

IDA

E

Mic

rope

plus

te

sser

ula

Cur

t.

Meg

arth

rus

sp. 1

M

egar

thru

s sp

.2

Pro

tein

us s

pp.

Eus

phal

erum

spp

. A

crul

ia i

nfla

ta (

Gyl

l.)

Pyc

nogl

ypta

lur

ida

(Gyl

l.)

Acr

oloc

ha c

f. s

ult'u

la

(Ste

ph.)

O

mal

ium

cae

sura

G

rav.

*O

loph

rum

?co

nsim

ile

(Gyl

l.)

O. f

useu

m (

Gra

v.)

*0.

bore

ale

(Pay

k.)

Olo

phru

m s

p.

Euc

neco

sum

br

achy

pter

um

(Gra

y.)

Aci

dota

cre

nata

(F

.)

A.

crue

ntat

a (M

ann

h.)

Le

stev

a sp

. G

eodr

omic

us n

igri

ta

(Mal

l.)

G. p

lagi

atus

(F

.)

G.

cf.

kunz

ei H

eer

Ant

hoph

agus

sp.

1 1

1

I 2 1

I 2

2 4

2 8

6

2 2

1 2

I 21

15

10

1

1

3 2

2 1

3 2

1 1

1 1

1

I~1

12

1

32

1

3

I

1 1

1 1

1

1 1

1 1

I 1

8 1

I 1

1 1

1 2 1

2 1

1 2

1 1

1 1

1 1 1

e~

e~

o~

e~

t~

* Bor

eaph

ilus h

enni

ngi-

an

us S

ahib

. *B

. nor

dens

kioe

ldi

Mhk

l.

Apl

oder

us c

aela

tus

(Gra

y.)

Oxy

telu

s in

seca

tus

Gra

v.

O. p

iceu

s (L

.)

O. l

aque

atus

(M

arsh

.)

O. s

culp

tura

tus

Gra

v.

*0.

?pol

itus

Er.

O

. niti

dulu

s G

rav.

1

*0.

gibb

ulus

Epp

. O

xyte

lus

sp.

Pla

tyst

hetu

s co

rnut

us

(Gra

v.)

Ble

dius

sp.

St

enus

sp.

1

1 E

uast

hetu

s b i

punc

tatu

s (L

jung

h)

Scop

aeus

sp.

La

thro

bium

1

term

inat

um G

ray.

X

anth

olin

us s

p. s

l. 1

Stap

hylin

us s

p.

Phi

lont

hus/

Que

dius

1

1 sp

. M

ycet

opor

us s

p.

1 B

olito

bius

sp.

1

Tach

ypor

us

chry

som

elin

us (L

.)

Tach

ypor

us s

p.

Tach

inus

rufip

es

(Cre

er)

T. l

atic

ollis

Gra

v.

T. c

ortic

inus

Gra

v.

T. e

long

atus

Gyl

l.

T. ?

fimet

ariu

s Gra

v.

Tach

inus

sp.

Myl

laen

a sp

. A

leoc

hari

nae

inde

t.

1011

1

PS

EL

AP

HID

AE

B

atri

sode

s sp

. B

ryax

is s

p.

1 1

1 2

1

1 1

1 1

1

1 1

1 1

I1

1 2

12

12

1

1

1

1 6

2 1

1 1

1 1

3 1

1 4

1 2

1 7

3 1

1 2

1 I

1 1

2 2

4

12

12

14

21

1

1

11

111

1 12

1 1

1 1

1 1

1 1

1 1

1 1

1 1

2 2

1 5

1 2

4 5

5 2

8 1

1 2

1 ?1

1

1 1

1 1

2 4

126

134

8 14

109

7 1

6 26

128

4 10

4

1 1

3 3

8 5

3 15

4

19

10

5 1

74

2

¢%

-&

U,

Tab

le 1

(co

ntin

ued)

.~

CA

NT

HA

RID

AE

R

hago

nych

a sp

, M

alth

odes

sp.

ME

LY

RID

AE

H

aplo

cnem

us s

p.

Das

ytes

sp.

EL

AT

ER

IDA

E

Am

pedu

s cf

. ni

gerr

imus

(La

c.)

Ade

loce

ra m

urin

a ( L

. I

Cte

nice

ra

pect

inie

orni

s (L

.)

C.

eupr

ea (

F.)

Cte

nice

ra s

p.

Pro

ster

num

te

ssel

latu

m

(L.)

Se

lato

som

us a

eneu

s (L

.)

Den

tico

llis

line

aris

(L

.)

Fle

utia

uxel

lus

mar

itim

us (

Cur

t.)

Ela

teri

dae

inde

t, p

l. sp

.

EU

CN

EM

IDA

E

?Hyl

ocha

res

dubi

us

(Pil

l. M

itt.

) G

. sp

.

TH

RO

SC

IDA

E

Thro

scus

car

inifr

ons

Bon

v.

BU

PR

ES

TID

AE

A

gril

us s

p.

G.

sp,

HE

LO

DID

AE

in

det.

0 1

2 3A

3B

4A

4B

5

6 7

8 9

1 I

1 1

I t

1 1

2 1

1 I

1 1

1 1

l 1

1 1

2 2

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

1 I

3

1 1

3

2 1

1 1

1 2

1 1

1 5

t 2

1

2 1

1

1

18

42

1

1 1

o~ 7"

DR

YO

PID

AE

D

ryop

s sp

. E

lmis

t.a

enea

(M

Oll

.)

2 E

solu

s pa

ralle

lipip

edus

(M

oll.

) O

ulim

nius

tube

rcul

atus

(M

Oll

.)

O.

trog

lody

tes

(Gyl

l.)

Oul

imni

us s

p.

1 Li

mni

us

1 1

1 vo

lckm

ari

(Pan

z.)

L. o

pacu

s (M

Oll

.)

Lim

nius

sp.

N

orm

andi

a ni

tens

(M

011.

) R

iolu

s sp

. 1

BY

RR

HID

AE

C

ytilu

s sp

. B

yrrh

us s

p.

* Sim

ploc

aria

m

etal

lica

(Stu

rm)

OS

TO

MID

AE

N

emos

oma

elon

gatu

m

(L.)

NIT

IDU

LID

AE

C

ater

etes

sp.

M

elig

ethe

s sp

. E

pura

ea s

p.

Poc

adiu

s fe

rrug

ineu

s (F.)

RH

IZO

PH

AG

IDA

E

Rhi

zoph

agus

de

pres

sus

(F.)

R

hizo

phag

us s

pp.

2 1

2 3

1 1

1 1

1

1 2

7 4

8 5

5 1

?1

6 9

4 6

2 3

1 2

3 4

2 2

2 1

4 1

35

6

2 3

2 2

8 1

3 3

1 5

2 3

5 2

8 4

3 1

1 3

1 1

2 1

3 1

1 1

18

7

3 10

I1

1

12

71

44

8

1 1

01

01

83

21

71

71

3

8 8

7 1

1 6

2 1

1 3

7 2

3 7

15

21

10

4

8 10

7

6 9

6

1 1

1 1

11

1

11

1

45

12

1

1

1 1

1 2

1 2

1 1

1 1

1 1 1

3

2 1

1 2

1 2

1 1

4 1

41

21

48

21

26

2

4 2 1

1 2

1

1 1

1 1

1 1

CU

CU

JID

AE

M

onot

oma

brev

icol

lis

Aub

6 A

irap

hilu

s sp

. Si

lvan

opru

s fa

gi (

Gu6

r.)

Tab

le I

(c

onti

nued

)

0 1

2 3A

3B

4A

4B

5

6 7

8 9

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 4

0 41

1 P

edia

cus

derm

esto

ides

(F

.)

Laem

ophl

oeus

bi

mac

ulat

us (

Pay

k.)

Laem

ophl

oeus

sp.

CR

YP

TO

PH

AG

IDA

E

Cry

ptop

hagu

s sp

. C

(M

icra

mbe

) sp

. 2

3 1

A to

mar

ia s

p.

3

PH

AL

AC

RID

AE

P

hala

crus

car

icis

1

2 S

turm

P

hala

crus

sp.

O

libru

s sp

. 1

1 I

G.

sp.

I

LA

TH

RID

IID

AE

E

nicm

us s

p.

I C

orti

cari

a/C

orti

cari

na

2 2

1 1

5 6

sp.

type

CO

LY

DII

DA

E

Pyc

nom

erus

ter

ebra

ns

(oi.)

D

itom

a cr

enat

a ( F

. )

Col

ydiu

m e

long

atum

(F

.)

C fi

lifo

rme

(F.)

EN

DO

MY

CH

IDA

E

?End

omyc

hus

sp.

1 12

11

4

e~

CO

CC

INE

LL

IDA

E

?Epi

lach

na s

p.

Scym

nus

sp.

Chi

loco

rus

bipu

stul

atus

(L.

) * H

ippo

dam

ia

7-m

acul

ata

( Gee

r)

1

1

H.

arct

ica

(Sch

neid

er)

Har

mon

ia 4

-pun

ctat

a (P

ont.)

CIS

IDA

E

Sulc

acis

?bi

dent

ulus

(R

osh.

) C

/s s

p.

AN

OB

IID

AE

D

ryop

hilu

s pus

illus

(G

yll.)

E

rnob

ius

cf.

nigr

inus

(S

turm

) E

rnob

ius

sp.

Ano

bium

pun

ctat

um

(Gee

r)

A. o

f. fu

lvic

orne

Stu

rm

A. c

f. pe

rtin

ax (

L.)

A

nobi

um s

p.

Dor

cato

ma

sp.

PT

INID

AE

P

tinus

fur

(L

.)

OE

DE

ME

RID

AE

O

edem

era

luri

da

( Mar

sh.)/

vire

scen

s (L

.)

AN

TH

ICID

AE

A

nthi

cus

sp.

MO

RD

EL

LID

AE

A

nasp

is h

umer

alis

(F

.)

Ana

spis

sp.

SE

RR

OP

AL

PID

AE

D

irca

ea 4

-gut

tata

(P

ayk.

)/au

stra

lis

Fair

m.

SC

AR

AB

AE

IDA

E

Geo

trup

es s

p.

Ont

hoph

agus

ve

rtic

icor

nis (

Lai

ch.)

O

ntho

phag

us s

p.

1 2 1

1

1 3 1

E-

.¢

E"

oa

4~

Tab

le 1

(co

ntin

ued)

o

c

Aph

odiu

s t.

foss

or

(L.)

*A.

hold

erer

i R

eitt

er

A.

rufip

es (

L.)

A

. t.

fim

etar

ius

(L.)

A

phod

ius

spp.

Se

rica

bru

nnea

( L

.)

?Tri

odon

ta s

p.

Ano

mal

a sp

. V

algu

s he

mip

teru

s (L

.)

LU

CA

NID

AE

C

eruc

hus

chry

som

elin

us

(Ho

ch.)

CE

RA

MB

YC

IDA

E

Rha

gium

bifa

scia

tum

iF

.)

dlos

tern

a ta

baci

colo

r (G

eer)

CH

RY

SO

ME

LID

AE

D

onac

ia c

lavi

pes

F.

D.

?ver

sieo

[ore

a (B

rah

m)

D,

thal

assi

na G

erm

, D

. ei

nere

a H

bst.

D

onae

ia s

p.

Don

aeia

/Pla

teum

aris

sp

. P

late

umar

is s

p.

Mae

ropl

ea

appe

ndic

ulat

a (P

anz.

) Le

ma

sp.

Cry

ptoc

epha

lus

?mor

aei

Cry

ptoc

epha

lus

sp.

Ado

xus

obsc

urus

( L

.)

0 1

2

2 2

11

2

!

3A

3B

4A

4B

5

1 1

6 5

2 4

67

89

2 1

2 1

1 1

1 1

1 1

1

1 1

1 1

1 l

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

2 1

06

1

1

1 2

1 3

1 1

2 1

1 I

1 1

1 1

1 1

2 7

4 1

5 4

2 1

1 1

1 !

1 1

l l

1 1

I

1

1 1

,2

-m

e,

4t~

Chr

ysom

ela

sp.

C. ?

cere

alis

L.

Chr

ysoc

hloa

sp.

P

haed

on s

p.

Pla

giod

era

vers

icol

ora

(Lai

ch.)

P

hyto

dect

a vi

min

alis

(L

.)

Phy

tode

cta

sp.

Phy

llode

cta

latic

ollis

Su

ffr.

P

hyllo

dect

a sp

. G

aler

uca

tana

ceti

(L.)

Lu

peru

s sp

. A

gela

stic

a al

ni (

L.)

P

hyllo

tret

a sp

. A

phth

ona

sp.

Hal

tica

sp.

Cre

pido

dera

sp.

C

halc

oide

s fu

lvic

orni

s (F

.)

Cha

etoc

nem

a sp

. C

assi

da s

p.

BR

UC

HID

AE

B

ruch

us/B

ruch

idiu

s sp

. B

ruch

idiu

s fas

ciat

us

(Ol.

) B

. de

bilis

(GyU

.)

B.

unic

olor

/deb

ilis

(Gyn

.)

AN

TH

RIB

IDA

E

Bra

chyt

arsu

s ne

bulo

sus

( For

st. )

G

. sp

.

SC

OL

YT

IDA

E

Scol

ytus

cf

. am

ygda

li (G

u&

.)

S. i

ntri

catu

s (R

atz.

) S.

cf.

mal

i (B

echs

t.)

S. c

f. e

arpi

ni (R

atz.

) S.

sco

lytu

s (F

.)

S. r

atze

burg

i Jan

son

6

4

2 8

1

5 4

6 3

2 4

3 5

2 3

6 1

1 1 4

2 1

2 2

2

2

2

1 1 1

2 1

4 1

1

1 I

1

1 1

E"

Tab

le 1

(co

ntin

ued)

0 l

2 3A

3B

4A

4B

5

6 7

8 9

10

11

12

13

14

15

16

17

18

19 2

0 21

22

23

26

27

29

30

31

32

33

34

35

36

37

38

39 4

0 41

S. m

ulti

stri

atus

1

(Mar

sh.)

H

ylas

tes

ater

(P

ayk.

) 1

?1

3 1

2 8

4 H

ylur

gops

pal

liat

us

1 6

3 2

5 4

6 1

1 2

(Gyl

l.)

Pol

ygra

phus

1

1 5

3 6

5 3

39

11 4

2

2 3

,~

poli

grap

hus

(L,)

P~

?su

bopa

cus

Tho

rns.

1

1 2

2 H

yles

inus

ole

iper

da

1 ~

- (F.)

~-

Lepe

risi

nus

vari

us

1 (F.)

Kis

soph

agus

hed

erae

I

""

(Sch

mit

t.)

Kis

soph

agus

/ ~

""

Xyl

echi

nus

sp.

~ P

ityo

phth

orus

2

1 !

! pi

tyog

raph

us (

Rat

z.)

~.

Pit

yoph

thor

us s

p.

I t,,

,,,

Pit

yoge

nes

I I

2 2

I 5

2 8

4 "~

ch

aleo

grap

hus

(L.)

P

. tr

epan

atus

2

I 2

5 2

(N6r

dl.)

~"

P

. bM

enta

tus

(Hbs

t.)

1 P

ityo

phth

orus

sp.

1

Pit

yokt

eine

s sp

inid

ens

5 8

2 ,~

(R

eitt

er)

P.

curv

iden

s (G

erm

.)

4 "~

P

. vo

ront

zow

i 10

12

8 (J

acob

s.)

Ort

hoto

mic

us

1 su

tura

lis

(GyU

.)

O.

lari

cis

(F.)

I

?Ort

hoto

mic

us s

p.

l lp

s se

xden

tatu

s 1

2 1

1 2

(Boe

rner

) X

yleb

orus

sax

esen

i I

1 (R

atz.

) X

. dr

yogr

aphu

s 1

(Rat

z.)

Xyl

oter

us d

omes

ticu

s 3

(L.)

X

ylot

erus

sp.

1

PL

AT

YP

OD

IDA

E

Pla

typu

s ox

yuru

s (D

uf.)

P.

cylin

drus

(F.

)

CU

RC

UL

ION

IDA

E

Pse

laph

orhy

nchi

tes

nanu

s (P

ayk.

) P

. to

men

tosu

s (G

yll.

) R

hync

hite

s ?a

urat

us

(Sco

p.)

Rhy

nchi

tes

sp.

Byc

tiscu

s be

tula

e (L

.)

B. p

opul

i (L

.)

Dep

orau

s tr

istis

(F

.)/se

min

iger

Rtt

. 1)

. bet

ulae

(L

.)

Api

on c

erdo

Ger

st.

Api

on s

pp.

Otio

rhyn

chus

gr.

cla

v-

ipes

(B

onsd

.)

O.

dubi

us (

StrU

m.)

O

. ruf

ifron

s (G

yU.)

O

tiorh

ynch

us s

p.

Phy

llobi

us/

Pol

ydru

sus

sp.

Pol

ydru

sus

sp.

Stro

phos

oma

capi

tatu

m (

Gee

r)

Bar

ynot

us o

bscu

rus

(F.)

Sito

na

of.

flave

scen

s (M

arsh

.)

Sito

na s

p.

Cle

onus

(sl.)

sp.

D

ryop

htho

rus

cort

ical

is (

Payk

.)

Rhy

ncol

us e

long

atus

(G

yU.)

Rhy

ncol

us s

p.

Sten

osce

lis s

ubm

uric

a-

tus

(Sch

finh

.)

Bag

ous

sp.

Not

aris

acr

idul

us (

L.)

N

. ae

thio

ps (

F.)

Ant

hono

mus

sp.

1 5 1

91

2 2

1 1

1

2112

11

1 1

1 1

1 1

11

1

22

23

21

1

1 1

2 1

322

1 1

1 211

5 5

1 1

3 1

1

231

1 1 1 14

1

1 1

7 3

2 1

1 1

1

1 11

11

11

21

1

1

1

11

31

35

31

14

21

1

.¢

7"

Tab

le 1

(co

ntin

ued)

0 I

2 3A

3B

4A

4B

5

6 7

8 9

l0

ll

12

13

14

15

16

17

18

19 2

0 2l

22

23

26 2

7 29

30

31

32

33

34

35

36 3

7 38

39

40

41

Bra

chon

yx p

inet

i 1

1 1

(Pay

k.)

Cur

culio

ven

osus

1

l 1

1 (G

ray.

) C

. gl

andi

um M

arsh

. 2

3 1

.~

C. p

yrrh

ocer

as

1 I

3 2

5 3

2 1

1 1

6 1

Mar

sh.

Pis

sode

s pi

ni (

L.)

1

P.

?har

cyni

ae (

Hbs

t.)

1 ~"

M

agda

lis

niti

da

2 2

3 4

1 1

(Gyl

l.)

Alo

phus

tri

gutt

atus

Lim

noba

ris

T-al

bum

I

l (L

.)/p

ilis

tria

ta

(Ste

ph.)

E

ubry

chiu

s ve

lutu

s I

Phy

tobi

us m

uric

atus

B

ris.

/gra

natu

s G

yll.

-

~"

Mic

relu

s er

icae

t

1 1

2 (G

yll.

) F

Gym

netr

on

sp.

l 1

Mia

rus

sp,

Ano

plus

pla

ntar

is

8 2

1 2

(Nae

zen)

R

hync

haen

us

quer

cus

2 2

2 2

1 2

2 2

-Q

(L.)

R_

ave

llan

ae

1 1

~-~

(Oon

ov.)

R

. ru

sci

(Hbs

t.)

3

R.

foB

orum

(M

fill

.)

2 1

~ 1

1 3

1 1

1 R

. (P

seud

orch

este

s)

1 sp

. R

hync

haen

us

sp.

1 1

1 R

ham

phus

pul

icar

ius

1 1

1 !

1 (H

bst.

) 1

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41 23

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

0 260 0 lO I I I I I I I I I I l l I I I r i l l I I I I I l l l l l t l l " l I I g. .

l l

......... ~m~mmm _ _

m mmm m m m m m m

.-.I

x --e

~o 133

$ $ (--

--i

GP-A7b

GP-A7a

GP-A6

GP-A5

GP-A4

-GP-A3 -

GP-A2b

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

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 oak- dependent 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).

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. Running- water species are markedly predominant (up to 71 individuals in sample 14) in the forest domi- nated 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 standing- water Coleoptera.

Running-water Coleoptera being dependent on

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

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

10 0 60 10 20 0 30 0 110 i-~ i i [ i l ' l , ~'-~ I ' - ~ i 11 ,1"~ i i I I I [ ~ i 7 ,

~.~ I'rl I'rl 7- : 7

I '~ I '~ m 0 3 I'rl m "13 "X3 Z Z

O O O O

m r ~

I II , , ,i I ~ 1 I I 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 pre- vailed 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. Dung- beetles are however more abundant in the

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41 25

41 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

20 10 0 700 40 10 0 80 m r- . I i i i ~ i i i ~1 i i 1 i 1--,i f i i i i ) i i I

-..t

¢o

F i g . 8. C o m p a r i s o n o f t h e o c c u r r e n c e s o f Abies p o l l e n a n d

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-

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

I

m m m m

l,,

"n ¢.rl ~10

~--4 t:~ H E:J

g~g g ~To

Fig. 9. Aquatic Coleoptera, uals (I).

I I I GP-A6 I l l | | GP-A~

, .n.. l ' GP-A4 |

' l | | | • GP-A3

number of taxa (T) and individ-

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 r r 7

41 40 39 38 37 36 35 34 33 32 31 30 28 2 l 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 abun- dance of tree-dependent and running-water Coleoptera. This unit may be divided in two sub- units 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 run- ning-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 pro- nounced 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 assem- blages 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 running- water 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, cold- adapted 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 o f faunal units

Main faunal units - -GP-AI: characterized by the occurrence of

cold-adapted Coleoptera, the scarcity of tree- dependent 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, Palaeoecology 114 (1995) 1-41 27

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 usu- ally lives on the peaty soil of open tundra, some- times 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 circum- polar 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 grass- lands 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 moun- tains, 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 south- ernmost part of its area it is restricted to moun- tains. 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

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 ecol- ogy do not fit with such an hostile environment, like Plagiodera versicolor and Gonioctena viminalis that feed upon willows (but not upon dwarf wil- lows 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 rela- tively 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 corres- ponds to a period of very sudden climatic improve- ment leading to the temperate conditions that prevail in the overlying faunal unit (GP-A2).

During most of the GP-A1 unit, the local envi- ronment 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 (Notio- philus, Bembidion dauricum, Amara quenseli) sug- gests 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 appear- ance of trees and tree-dependent Coleoptera sug- gesting a lag in their response time, possibly

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

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

GP-A2a faunal unit This unit is characterized by the abundance of

deciduous tree-dependent Coleoptera and by the total absence of conifer-dependent and cold- adapted 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, Chal- coides fulvicornis, Byctiscus populi, Rhynchaenus rusci and Rhamphus pulicarius. Bark-beetles are represented by many species, including Scolytus multistriatus and S. scolytus that are mainly elm- dependent. 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 envi- ronment 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 repre- sented only by Stenus, Lathrobium and several species of Donaeia, truly aquatic taxa are very abundant. This group is dominated by running- water 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 running- water beetles and the rarity of species of standing water suggests that the sediment was carried into the area by running water or even directly depos- ited by it.

GP-A2b faunal 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 conifer- dependent species, mostly scolytids and weevils, such as Hylastes ater, Hylurgops palliatus, Ips sexdentatus, Polygraphus polygraphus, Pityo- phthorus 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 tree- dependent 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 29

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 sug- gests 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 rain- fall; 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

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

GP-A3 faunal unit This unit is limited to sample 10 and is charac-

terized 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 elonga- tus. Bearing in mind the decidedly non-forest char- acter of the assemblage as a whole, it is likely that these specimens really belong either to the overly- ing 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 dete- rioration was perhaps too short to allow the north- ern 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 cold- adapted 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 running- water 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 assem- blage fairly similar to that described in GP-A2b unit. This assemblage is characterized by a rich fauna of tree-dependent species indicating the pres- ence 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 scolyt- ids Orthotomicus laricis and O. suturalis which are both parasite of many coniferous trees, Xyloterus domesticus that lives exclusively on various decidu- ous 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 tree- dependent 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 unci- nata. 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 bimacu- latus, 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 tomen- tosus (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 open- ing 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. Phala- crus caricis is a small phytophagous Coleoptera feeding on smutted Carex (Thomson, 1958), it is another typical marsh species. A few taxa indica- tive of standing water are recorded (Agabus, Enochrus, Anacaena, Coelostoma, Helophorus). However, running-water taxa (Elmis, Esolus, Oulimnius, Limnius, Normandia) are again domi- nant 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 pub- lished data. Nevertheless the presence of this insect in GP-A4 unit probably denotes climatic condi- tions 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, Palaeoecology 114 (1995) 1-41 31

taxa do not disappear totally since Dircaea australis/quadriguttatus, Polygraphus polygraphus (on conifers), Deporaus betulae and Anoplus plan- taris (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 discov- ered at La Grande Pile during the coring opera- tions, 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 preda- tor ground-beetle Synuchus nivalis that lives on dry and sandy soils (Koch, 1989) suggests an open environment also.

GP-A6 faunal 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, Paromalus flavicornis, Laemophloeus bima- culatus, Colydium filiforme, Ceruchus chrysomeli- nus, Polygraphus polygraphus, Hylurgops palliatus, Scolytus intricatus, S. ratzeburgi, Strophosoma capitatum, Curculio pyrrhoceras, Dryophthorus cor- ticalis, Brachonyx pineti and Magdalis nitida. These species indicate a return of a truly forested land- scape. However, the occurrence of a single indivi- dual 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 pro- vides a hint that the climate was cooler than in either unit GP-A2 or GP-A4. Running-water aquatic genera such as Normandia, Esolus,

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, sug- gesting 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, Hydro- philidae). 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 ecolog- ical 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 biogeographi- cal 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 consis- tent 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). Diacheila polita, last found in GP-A1, reap- pears 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 peat- bogs, and on mossy lake shores in the north of the British Isles and in Fennoscandia (Balfour- Browne, 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 palaearc- tic 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 wide- spread in the Siberian taiga and tundra. In the southern part of its area this species is apparently restricted to forest although it is common else- where 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 pre- viously 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 henningia- nus (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 sta- phylinid, 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 spo- radic 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 corre- lation of the Grande Pile sedimentary levels with the British sediments where similar fossil assem- blages 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 33

It is convenient to divide GP-A7 unit into two subunits:

--GP-A7a (samples 21-35) is characterized by the almost complete disappearance of tree- dependent species, the first indications of the re-appearance of the most cold-adapted Coleoptera, the persistence of running-water bee- tles 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 holder- eri, 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 sup- ported by the occurrence of the crustacean Notostraca Lepidurus arcticus Pallas which appears in huge numbers (223 mandibles were

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 pre- vailing 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 corre- lated (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 Isotope stages Beaulieu and Reille (1992a) Woillard and Moock (1982)

GP-A7b Wiirmian Pleniglacial 2 GP-A7a Wttrmian Pleniglacial 3 samples missing Lower Wiirmian Pleniglacial 4 GP-A6 Saint Germain II 5a GP-A5 Mrlisey II 5b GP-A4 Saint Germain I 5c GP-A3 Mrlisey I 5d GP-A2b Eemian Interglacial 5e GP-A2a Eemian Interglacial 5e GP-A1 Riss (Linexert) 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 palaeoenviron- ment 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 attrib- uted 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 deteri- orations according to both pollen and insect analy- sis. Though occurring in low numbers, running- water 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 sup- ported by some pollen data. Thus Fig. 1 in Beaulieu and Reille (1992a) shows that shallow and calm water plants such as Isoetes are practi- cally 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 com- bined with the scarcity of lake or pond taxa certainly indicate the presence or even the predomi- nance 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 accumu- lation 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 35

(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 biogeo- graphical, ecological and climatical data can thus be obtained from the rich literature that has been devoted to Coleoptera during the two last centu- ries. Moreover a recent computerized palaeocli- mate 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 quan- titative 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 especi- ally significant from a palaeoclimatic point of view because their distribution is not likely to be linked to the occurrence of particular host-plants but

imposed by climatic factors. Usually, the food chains of such species ultimately depend on micro- organisms (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, Cara- bidae and Scarabaeidae families (e.g. Elaphrus lapponicus, Diacheila arctica, Diacheila polita, Bembidion dauricum, Amara quenseli, Potamonectes griseostriatus, Potamonectes assimilis, Agabus arcti- cus, Colymbetes dolabratus, Helophorus glacialis, Helophorus sibiricus, Helophorus oblongus, Ptero- loma forsstroemi, Pycnoglypta lurida, Eucnecosum brachypterum, Boreaphilus henningianus, Borea- philus 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 ground- beetle 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 repre- sented 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 deterio- ration of Mrlisey I is clearly visible in sample 10,

3 6 P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41

10 10 10 10 0 30 i TM i-~ i-'n F - ~ I I ,

4, !I"i' 'l,ilE" ,o I i l t I i ! G 39 P-A7b 3 8

36 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 a,t

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

~0 19 18

16 15

13 12 11 10

9

?

5 4B

4A 313 3A

2 1 O

~

~ ~ NN

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 reappear- ance 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, Palaeoecology 114 (1995) 1-41 3 7

'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,,-,~, ~P-~. G~AZ. I ~ GP-A4 Q,.-,. GP,.A6 GP,.A7a GP-A7b

c L E E M I SG I M II SG II PW LW

"C

+ 10'

- 1 0 ,

. 2 0 ¸

- 30

iii iif: iiliiii:i - i ii ::liii i - ii:: iii i i:i!l !!i/ii!ili!!!i!!!!

. . . . . . . . . . . . . . . . . . . . . . . . . . . . ,..2122-- ;21212; ,°o12222-2221h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Fig. 12. "Mutual climatic range" climatic reconstruction for the Coleoptera from La Grand Pile (°C). The horizontal axis is approximately equivalent to time B.P.a. Insect samples, b, Entomological biostratigraphic units, c. Pollen chronozones of Beaulieu and Reille (1992a), L= Linexert (or Riss), E= Eemian, M I= M61isey I, M H= M61isey II, SG I= Saint Germain I, SG H= Saint Germain II, PW= Pleniwllrm, LW= Late WOrm.

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 reconstruc- tion 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 occur- rence 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 sedi- mentary gaps were recognised during pollen analy- ses (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 conti- nental Western Europe gives a detailed picture of the drastic changes in the insect fauna that corre- late 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 Riss- Eemian lead to the replacement of a cold-adapted fauna and of a tundra-type environment by a tree- dependent 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 (how- ever 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 estab- lishment of a very cold-adapted insect fauna, con- currently 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 indi- cate 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 pres- ence 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 pro- gram 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.

References

Aaby, B. and Diggerfeldt, G., 1986. Sampling techniques for lakes and bogs. In: B.E. Berglund (Editor), Handbook of Holocene Palaeoecology and Palaeohydrology. Wiley, Chichester, pp. 181-194.

Atkinson, T.C., Briffa, K.R. and Coope, G.R., 1987. Seasonal temperatures in Britain during the past 22,000 years, reconstructed using beetles remains. Nature, 325: 587-592.

Atkinson, T.C., Briffa, K.R., Coope, G.R., Joachim, M.J. and Perry, D.W., 1986. Climatic calibration of coleopteran data. In: B.E. Berglund (Editor), Handbook of Holocene Palaeoecology and Palaeohydrology. Wiley, Chichester, pp. 851-858.

Angus, R,B., 1973. Pleistocene Helophorus [Coleoptera Hydrophilidae] from Borislav and Starunia in the Western Ukraine with a reinterpretation of M. Lomnicki's species. Description of a new Siberian species, and a comparison with British Weichselian faunas. Philos. Trans. R. Soc. London, B 265: 299-326.

Balachowsky, A.S., 1949. Faune de France. 50, Colroptrres Scolytidae. Libr. Fac. Sci., Paris, 320 pp.

Balfour-Browne, F., 1950. British Water Beetles, 2. Ray Soc., London, 394 pp.

Beaulieu, J.-L. de and Reille, M., 1984. A long Upper

P. Ponel/Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1995) 1-41 39

Pleistocene pollen record from Les Echets, near Lyon, France. Boreas, 13:111-132.

Beanlieu, J.-L. de and Reille, M., 1992a. The last climatic cycle at la Grande Pile (Vosges, France): a new pollen profile. Quat. SCI. Rev., 11: 431-438.

Beaulieu, J.-L. de and Reille, M., 1992b. Long Pleistocene pollen sequences from the Velay Plateau (Massif Central, France). I. Ribains maar. Veget. Hist. Archaeobot., 1: 233-242.

Beaulieu, J.-L. de, Monjuvent, G., Nicoud, G., Richard, H. and Seret, G. (with the collaboration of: Campy, M., Clerc, J., Dricot, E., Either, U., Mandier, P., Ponel, P., Reille, M., Rousseau, D.D., Ruffaldi, P. and Wansard, G.), 1992. Long pollen sequences and the last glaciations from the Southern Alps to the Vosges mountains. In: 8th Int. Palynol. Congr., Aix-en-Provence, 13-16th Sept. 1992, Excursion I. Cah. Micropaltontol., 7: 215-257.

Boulton, G.S., 1993. Two cores are better than one. Nature, 366: 507-508.

Buckland, P.C. and Coope, G.R., 1991. A Bibliography and Literature Review of Quaternary Entomology. Collis Publ., Dep. Archaeol. Prehist., Univ. Sheffield, 85 pp.

Callot, H.J., 1990. Hydradephaga. In: Catalogue et atlas des Coltopt~res d'Alsace. Soc. Alsac. Entomol. Mus. Zool. Univ. Strasbourg, 68 pp.

Ch~tenet, G. du, 1986. Guide des Coltoptbres d'Europe. Delachaux et Niestlt, Lausanne, 480 pp.

Coope, G.R., 1961. A Pleistocene coleopterous fauna with arctic affinities from Fladbury, Worcestershire. Q. J. Geol. Soc. London, 118: 103-123.

Coope, G.R., 1962. Coleoptera from a peat interbedded between two Boulder Clays at Burnhead near Airdrie. Trans. Geol. Soc. Glasgow, 24: 279-286.

Coope, G.R., 1968. An insect fauna from Mid Weichselian deposits at Brandon, Warwickshire. Philos. Trans. R. Soc. London, B 254: 425-456.

Coope, G.R., 1969. The response of coleoptera to gross thermal changes during the Mid Weichselian interstadial. Mitt. Int. Ver. Theor. Angew. Limnol., 17: 173-183.

Coope, G.R., 1973. Tibetan species of dung beetle from Late Pleistocene deposits in England. Nature, 245: 335-336.

Coope, G.R., 1975. Mid-Weichselian climatic changes in Western Europe re-interpreted from coleopteran assem- blages. In: R.P. Suggate and M.M. Cresswell (Editors), Quaternary Studies. R. Soc. N. Z., Wellington, pp. 101-108.

Coope, G.R., 1977. Fossil coleopteran assemblages as sensitive indicators of climatic changes during the Devensian (Last) cold stage. Philos. Trans. R. Soc. London, B 280: 313-340.

Coope, G.R., 1986. Coleoptera analysis. In: B.E. Berglund (Editor), Handbook of Hoiocene Palaeoecology and Palaeohydrology. Wiley, Chichester, pp. 703-713.

Coope, G.R., 1987. The response of late Quaternary insect communities to sudden climatic changes. In: J.H.R. Gee and P.S. CAller (Editors), Organization of Communities Past and Present. Blackwell, Oxford, pp. 421-438.

Coope, G.R., 1990. The invasion of Northern Europe during the Pleistocene by Mediterranean species of Coleoptera. In:

F. di Castri, A.J. Hansen and M. Debussche (Editors), Biological Invasions in Europe and the Mediterranean Basin. Kluwer, Dordrecht, pp. 203-215.

Coope, G.R., 1994. The response of insects faunas to glacial- interglacial climatic fluctuations. Philos. Trans. R. Soc. London, B 344: 19-26.

Coope, G.R. and Angus, R.B., 1975. An ecological study of a temperate interlude in the middle of the last glaciation, based on fossil Coleoptera from Isleworth, Middlesex. J. Animal Ecol., 44: 365-391.

Coope, G.R. and Sands, C.H.S., 1966. Insect faunas of the last glaciation from the Tame Valley, Warwickshire. Proc. R. Soc. London, B 165: 389-412.

Coope, G.R., Shotton, F.W. and Strachan, I., 1961. A Late Pleistocene fauna and flora from Upton Warren, Worcestershire. Philos. Trans. R. Soc. London, B 244: 379-421.

Comet, C., 1988. Interprttation paltotcologique des diatomtes /t la Grande Pile (France). MOrt. Soc. R. Bot. Belg., 10: 77-86.

Freude, H., 1971. Rhysodidae. In: Freude-Harde-Lohse, Die Irdifer Mitteleuropas. Goecke and Evers, Krefeld, pp. 93-94.

Goeury, C., 1988. Acquisition, gestion et reprtsentation des donntes de l'analyse pollinique sur micro-ordinateur. In: Actes 10~ne Symp. Assoc. Palynol. Langue Fr. Inst. Fr. Pondichtry, Tray. Sect. Sci. Tech., 25: 405-4-416.

G.R.I.P. members, 1993. Climate instability during the last interglacial period recorded in the GRIP core. Nature, 364: 203-207.

Guignot, F., 1947. Faune de France. 48. Coltopt~res Hydrocanthares. Lechevalier, Paris, 287 pp.

Guiot, J., Beaulieu, J.-L. de, Cheddadi, R., David, F., Ponel, P. and Reilte, M., 1993. The climate in Western Europe during the last Glacial/Interglacial cycle derived from pollen and insect remains. Palaeogeogr. Palaeoclimatol. Palaeoecol., 103: 73-93.

Guiot, J., Pons, A., Beaulieu, J.-L. de and Reille, M., 1989. A 140,000-year continental climate reconstruction from two European pollen records. Nature, 338: 309-313.

Guiot, J., Reille, M., Beaulieu, J.-L. de and Pons, A., 1992. Calibration of the climatic signal in a new pollen sequence from La Grande Pile. Clim. Dyn., 6: 259-264.

Hammond, P., Morgan, A. and Morgan, A.V., 1979. On the gibbulus Group of Anotylus, and fossil oeeurences of Anotylus gibbulus (Staphylinidae). Syst. Entomol., 4: 215-221.

Hansen, M., 1987. Fauna Entomologica Scandinavica. 18. The Hydrophiloidea (Coleoptera) of Fennoscandia and Denmark. Brill, Scan. Sci. Press, Leiden, Copenhagen, 254 pp.

Herv~, P., 1951. Apropos de captures de Ceruchus chrysomeli- nus Hoch. darts les Alpes-Maritimes. Conditions ~tholog, iques, climatiques et foresti~res. Entomologiste, 7: 30-35.

Hoffmann, A., 1945. Faune de France. 44. Coltopt~res Bruchides et Anthribides. Libr. Fac. Sci., Paris, 184 pp.

Hoffmann, A., 1954. Faune de France. 59. Col~opt~res Curculionides (2~me partie). Libr. Fac. Sci., Paris, pp. 487-1208.

Hoffmann, A., 1958. Faune de France. 62. Col~opt~res

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

Curculionides (36me partie). Libr. Fac. Sci., Paris, pp. 1209-1840.

Johnsen, S.J., Clausen, H.B., Dansgaard, W., Fuhrer, K.. Gundestrup, N., Hammer, C.U., Iversen, P., .Iouzel, J., Stauffer, B. and Steffensen, J.P., 1992. Irregular glacial interstadials recorded in a new Greenland ice core. Nature, 359: 311-313.

Koch, K., 1989. Die Kafer Mitteleuropas, Okologie 1. Goecke and Evers, Krefeld, 440 pp.

Koch, K., 1992. Die Kafer Mitteleuropas, Okologie 3. Goecke and Evers, Krefeld, 389 pp.

Lefkovitch, L.P., 1958. A revision of the European Laemophloeinae (Coleoptera: Cucujidae). Trans. R. Entomol. Soc. London, 111:95 -118.

Lemdahl, G., 1990. Water-surfaces as insect traps and some consequences for palaeoentomology. Quat. Newsl., 62: 26 29.

Leseigueur, L., 1972. Col6opt6res Elateridae de la faune de France continentale et de Corse. Bull. Soc. Linn. Lyon, 382 pp. (suppl.).

Lindroth, C.H., 1985. Fauna Entomologica Scandinavica. 15 (part 1). The Carabidae (Coleoptera) of Fennoscandia and Denmark. Brill, Scand. Sci. Press, Leiden, Copenhagen, 226 pp.

Lindroth, C.H., 1986. Fauna Entomologica Scandinavica. 15 (part 2). The Carabidae (Coleoptera) of Fennoscandia and Denmark. Brill, Scand. Sci. Press, Leiden, Copenhagen, pp. 227 598.

Louis, A. and Smeets, J., 1981. Etude d6taill6e des fluctuations algales comme param6tres des fluctuations thermales dans un refroidissement (M61isey II) de l'Interglaciaire Riss- Wtlrrn. Stud. Algol. Lov., 10, 241 pp.

Louis, A., De Cremer, D. et coll., 1981. Les fluctuations algales param6tres des fluctuations thermales au St Germain II et/i l'Eowfirm. Stud. Algol. Lov., 11,370 pp.

Louis, A. and Ermin, L., 1983. Les fluctuations climatiques au Pr~-E6mien et au d6but de l'E6mien d~duites des fluctuations algales. Stud. Algol. Lov., 12, 126 pp.

Louis, A. and Peters, J., 1983. Les fluctuations climatiques de l'E~mien, du M~lisey I e t du St Germain I d6duites des fluctuations algales fossiles. Stud. Algol. Lov., 13, 270 pp.

Louis, A. et al., 1983. Fluctuations climatiques au N6ow~rm et Pr6bor6al d6duites des fluctuations algales. Stud. Algol. Lov., 15, 190pp.

Lucht, W.H., 1987. Die K~fer Mitteleuropas, Katalog. Goecke and Evers, Krefeld, 342 pp.

Mani, M.S., 1968. Ecology and Biogeography of High Altitude Insects. Junk, The Hague, 527 pp.

Paulian, R., 1959. Faune de France. 63. Col~opt6res Scarab6ides. Libr. Fac. Sci., Paris, 298 pp.

Ponel, P., Beaulieu, J.-L. de and Tobolski, K., 1992. Holocene palaeoenvironment at the timberline in the Taillefer Massif, French Alps: a study of pollen, plant macrofossils and fossils insects. Holocene, 2:117-130.

Ponel, P. and Coope, G.R., 1990. Lateglacial and Early Flandrian Coleoptera from La Taphanel, Massif Central,

France: Climatic and Ecological Implications. J. Quat. Sci., 5:235 249.

Ponel, P., Etlicher, B., Beaulieu, J.-L. de, Debard, E., Thinon, M., Vasari, A. and Petiot, R., 1991. La fin de la derni6re glaciation dans le Cantal (France): la tourbi&e de la Taphanel et son environnement. Quaternaire, 2: 147-163.

Pons, A., Campy, M. and Guiot, J., 1989. The last climatic cycle in France: the diversity of records. Quat. Int., 3/4: 49-55.

Pons, A., Guiot, J., Beaulieu, J.-L. de and Reille, M., 1992. Recent contributions to the climatology of the last glacial interglacial cycle based on french pollen sequences. Quat. Sci. Rev., 11: 439-448.

Reille, M. and Beaulieu, J.-L. de, 1990. Pollen analysis of a long younger Pleistocene continental sequence in a Velay maar (Massif Central, France). Palaeogeogr. Palaeoclimatol. Palaeoecol., 80:35 48.

Sainte-Claire Deville, J., 1914. Catalogue critique des Col~opt6res de la Corse. Caen, 573 pp.

Seret, G., 1967. Les syst6mes glaciaires du bassin de la Moselle et leurs enseignements. Soc. R. Beige G6ogr., 77, 577 pp.

Seret, G., Dricot, E. and Wansard, G., 1990. Evidence for an early glacial maximum in the French Vosges during the last glacial cycle. Nature, 346:453 456.

Seret, G., Guiot, J., Wansard, G., Beaulieu, J.-L. de and Reille, M., 1992. Tentative palaeoclimatic reconstruction linking pollen and sedimentology in La Grande Pile (Vosges, France). Quat. Sci. Rev., 11: 425-430.

Shotton, F.W., Keen, D.H., Coope, G.R., Currant, A.P., Gibbard, P.L., Aalto, M., Peglar, S.M. and Robinson, .I.E., 1993. The Middle Pleistocene deposits of Waverley Wood Pit, Warwickshire, England. J. Quat. Sci., 8: 293-325.

Shotton, F.W. and Osborne, P.J., 1965. The fauna of the Hoxnian Interglacial deposits of Nechells, Birmingham. Philos. Trans. R. Soc. London, B 248: 353-378.

Strand, A., 1946. Nord-Norges Coleoptera. Tromso Mus. Arsh. Naturhist. Avd., 34, 67 ( 1 ), 629 pp.

Taylor, B.J. and Coope, G.R., 1985. Arthropods in the Quaternary of East Anglia--Their role as indices of local palaeoenvironments and regional palaeoclimates. Mod. Geol., 9: 159-185.

Tessier, L., Beaulieu, J.-L. de, Cof~teaux, M., Edouard, J.-L., Ponel, P., Rolando, C., Thinon, M., Thomas, A. and Tobolski, K., 1993. Holocene palaeoenvironments at the timberline in the French Alps--a multidisciplinary approach. Boreas, 22: 244-254.

Theobald, N., Thi6baut, J. and Bernatzky, M., 1974. Carte g6ologique /~ 1/50,000, Giromagny XXXV-20. BRGM, Orleans.

Thomson, R.T., 1958. Coleoptera Phalacridae. In: Handbooks for the Identification of British Insects, V, 5 (b). R. Entomol. Soc. London, London, 17 pp.

Tiberghien, G., 1969. Nouvelles observations sur Rhysodes sulcatus F. [Col. Rhysodidae]. Entomologiste, 25: 85-92.

Woillard, G., 1973. Mise en 6vidence de l'E~mien sur le plateau de Haute-Sa6ne. C. R. Acad. Sci., Paris, 276: 939-942.

Woillard, G., 1974a. Recherches sur le P16istoc+ne dans l'Est

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

de la Belgique et dans les Vosges lorraines. Thesis. Univ. Cathol. Louvain, 216 pp.

Woillard, G., 1974b. Expos6 des recherches palynologiques sur le Plgistoc~ne dans l'Est de la Belgique et des Vosges lorraines. Trav. Lab. Palynol. Phytosociol., Univ. Cathol. Louvain, 19 pp.

Woillard, G., 1975. Recherches palynologiques sur le P16istoc.bne dans l'est de la Belgique et dans les Vosges lorraines. Acta Geogr. Lov., 14, 118 pp.

Woillard, G., 1977a. Comparison between the chronology from the beginning of the classical Eemian to the beginning of the classical Wllrm in Grande Pile peat-bog, and other chronolo- gies in the world. In: V. Sibrava (Editor), Quaternary Glaciations in the Northern Hemisphere. Rep., 4, Praha, pp. 72-82.

Woillard, G., 1977b. Grande Pile: a continuous climatic record for the last 140,000 years. In: Abstr. 10th INQUA Congr., Birmingham, 1977, p. 505.

Woillard, G., 1978a. Grande Pile peat-bog: a continuous pollen record for the last 140,000 years. Quat. Res., 9: 1-21.

Woillard, G., 1978b. The last interglacial-glacial cycle at Grande Pile in northeastern France. Trav. Lab. Palynol. Phytosociol., Univ. Louvain, 21 pp.

Woillard, G., 1979. The last interglacial-glacial cycle at Grande Pile in Northeastern France. Bull. Soc. Beige G-6ol., 88: 51-69.

Woillard, G. and Moock, W.G., 1982. Carbon dates at Grande Pile: correlation of land and sea chronologies. Science, 215: 159-161.

Zanetti, A., 1987. Fauna d'Italia. 25. Coleoptera Omaliinae. Calderini, Bologna, 472 pp.