European reference levels and correlation tables

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13
EUROPEAN REFERENCE LEVELS AND CORRELATION TABLES
Norberl SCHMIDT-KITTLER (ed.)

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Miinchner Geowiss. Abh. (A) 10

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Comments of the editor
Oneof the majoraimsof thesymposiom wasto establish
a biochronological subdivision of the European Paleogene
based on the evolution of mammals. This subdivision
should be applicable to all parts of Europe despite regional
differences in faunal composition. lt is also meant to
provide a stable system of reference irrespective of in-
creasing knowledge on details. Due to differences in
quality of the faunal data from different pads of Europe and
for different periods of the Paleogene, the degree of
accuracy which can be attained by regional subdivisions
varies largely. Because of this, a workable subdivision for
whole Europe should be just as precise as to allow
correlation of biochronological units between all parts of
this bioprovince.
A characteristic of the European bioprovince in the
Paleogene are the marked faunal endemisms during long
periods of the Upper Eocene and Oligocene. They were
partly caused by zoogeographical barriers (e.9. the Upper
Rhine graben, at times connected to the Molasse sea) and
partly due to physicaldifferences, such as lowlands with
fairly high ground-water levels (e.9. continental Molasse
basins, Hampshire basin, Paris basin) and dry karst
highlands (Quercy, Suebian and Franconian Alb). Climatic
gradients may have played an important role as well. The
differences in composition of the regional mammalian
faunas caused by these factors become even more pro-
nounced through faunal docu mentation due to particular
geological settings in the European subprovinces. Of
course, regional concepts of subdivisions have to take into
account the particular kinds of faunal documentation and
geological data. However, for the purpose of establishing
a subdivision applicable on the European scale a more
general, conte)d-independent, concept is needed.
Any kind of biochronology is based on the process of evo-
lution during which features of organisms become trans-
formed and it should be stressed here that using evolution-
ary changes as biochronological markers is independent
from evolutionary theories. Both, gradual character trans-
formation and intermittent rapid change of features of
species are suitable biochronologicalcriteria. As a matter
of fact, in Tertiary mammals gradual change of characters
within lineages is f requent and in many cases welldocu-
mented. ln order to extract biochronologicalcriteria from
such morphoclines, particular evolutionary stages have to
be defined as chronospecies (or -subspecies) by designat-
ing type specimens. These stages can directly be used as
points of reference within a biochronological concept
based on reference levels (niveaux repdres). Contrary to
that, the zone concept in those cases can only be applied
when - independently from erecting a chronospecies by
designating a type specimen - a particular part of a
morphocline is defined as the range of this species. This
once more has to be done using discrete, morphologically
recognizable, evolutionary stages. This is the reason why
reference levels have been favoured by the majority of the
Norberl SCHMIDT-KITTLER (ed.)
participants of the symposium (36:4) as means of bio-
chronological subdivision.
Of course, events of immigration and extinction of faunal
elements are used for subdivision as well. These, how-
ever, can only be taken as reliable in as much as theyare
controlled by the information drawn from evolutionary
lineages.
Another conceptual discussion related to this was cen-
tered on the question of wheter it would be possible to
subdivide the European continental Paleogene into bio-
stratigraphic stages by defining type sections. Localities
within a geological section already provide geochronologic
information by their lithostratigraphic superposition. How-
ever, it is clear that such stratified localities are relevant to
biostratigraphy only in so far as it would be possible to
rangethem chronologically in the same orderas they are
placed by superposition, even in case they would be found
in stratigraphically unrelated beds. it is not suflicient that
one or two levels within a geological sequence provide
good faunas, if at the same time there is few documenta-
tion in other parts of the section. This, however is the most
typical case with continental sections in the European
Paleogene.
The degree of precision which can be attained by a
biochronology depends mainly on two factors:The number
of localities in which biochronologically relevant lineages
are documented, and the size of the samples (e.9. number
of rodent teeth) on which the recognition of successive
evolutionary stages of the lineages is based. lt particulary
means that the degree of precision does not depend on the
geological context. This can best be demonstrated by the
example of karst fissure fillings which play an important
role in the biostratigraphy of the European Paleogene: Due
to the richness of their faunal content they can be arranged
accurately along the time axis though they do not comprise
geologic information on their age (except the age of the
rockwherein they are found).
The particular advantage of a system of reference levels
is that for its definition the best mammal localities available
can be chosen regardless of whether they were found in
sections or not, or whether they are fissure fillings. ln this
way the evolutionary process can be translated into a bio-
chronological scale in most detail. The disadvantage of
type sections for a biochronology based on mammals is
that their biologic information content is (with few excep-
tions)too poor and too irregularly distributed along the time
axis. Because of this they normally cannot reach the
standard of precision which can be attained by a system
of reference levels. This disadvantage is not so strong in
case a good potential of appropriate geological sections is
at hand (e.9. in invertebrate paleontology;e.9. continental
mammaliferous sections in North America) so that a good
choice can be made. lt is, however, certainly relevant to the

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continental Paleogene of Europe because there are only
very few regions where mammal bearing sections exist
and these, with only few exceptions, suffer from poorness
in fossil content. This is another reason why reference
levels have been favoured as means of subdivision of the
European continental Paleogene.
At the same time it was felt that detailed description of as
many mammal bearing sections as possible is very impor-
tant in the future. This will improve the possibilities for con-
necting the phenomena of mammalian evolution and fau-
nal development with data from invertebrate paleontology,
paleobotany and palynology as well as to sedimentologi-
cal data and paleomagnetism. ln order to chose appropri-
ate reference localities different criteria were applied.
These are richness in faunal content, good documentation
on both, macromammals and micromammals, and the po-
sition of the sites within Europe. ln case several good
faunas defining the same levelwere available, the one with
most central position was favoured in order to eliminate
inaccurracy due to biogeographical effects. Because of
the latter pragmatic reason the reference levels mostly
were defined on the basis of French localities.
The faunas of the reference localities form points on the
time axis, not intervalls. Thus no boundaries between the
reference levels are defined and the faunas to be corre-
lated have to be assigned to a particular levelon the basis
of their affinities as expressed by evolutionary stages and
first or last evidence of species. The fact that the intervals
between the reference levels are not defined does not
diminish the exactness of the system of subdivision be-
cause this depends exclusively on the quality of the
reference faunas. lt has already been mentioned above
that the European standard subdivision should be less
detailed as compared to the biochronological resolution
power which can be attained in regional suMivisions due
to (partly also endemic)faunal elements. Thus the horizon-
tal lines which separate the reference levels and correlated
localities in tab.2 are mainly to be taken as facilitations to
reading.
As to the duration of the time intervals lying between the
reference levels no reliable data are at hand since rates of
evolution change and there is no method of measuring
them. The resulution power of absolute dating is too weak
given the fact that much too few calibrations are available
for the European Paleogene and their inaccurracy is of the
same order of magnitude (t 0,5 m.y.) as the intervals to be
measured (average of about 1,2 m.y. between the refer-
ence levelsforthe upperpart of the Paleogene). Judging
from the very detailed documentation of lineages, fossil
information in the European Paleogene is particulary rich
forthe periods MP 16-18 and MP 25-30. On the other hand,
due to the geological conditions in Europe, documentation
of mammalian faunas olderthan Middle Eocene is much
more scattered than in the following period of the
Paleogene and it is evident that the Paleocene corre-
sponds to a big gap in the mammalian report. This is why
the numbers 0-5 were reserved for sites which we hope will
be discovered in the future. ln tab.2 the levels MP 8 and MP
9 are provisionally not separeted (see explanation of M.
Godinoton p.21).
The definition of the reference levels mainly in the French
subprovince leads to the consequence that faunas from
other subprovinces, in so far as they are different, have to
be classified as endemic in respect of the French faunal
compositions. Because of this their faunal elements are
only partly represented in tab.3-7 as "taxa occuring in well
correlated localities". This is for instance the case forthe
Upper Eocene and Lower Oligocene faunas of the Hamp-
shire basin and Southern Germany. lt is an inevitable
effect of the convention adopted. Of course, endemic ele-
ments are not less important but even of particular interest
as keys to the understanding of biogeographical barrieres
and paleoecological differences between paleofaunas.
Munchner Geowiss. Abh. (A) 10

17
Gorrelation tables
Tables2-Twerecompiledbytheactivecooperationof all Correlation of the established reference levels to the
participants during the sessions of the symposium. This marine stages is shown in tab.1 . ln tables 3-7 the reference
was realised by forming two seperate groups of specialists levels are grouped together following the boundaries of
for MP 1-20 and MP 21-30. The coordinators completed marine stages.
and adjusted the data during the time after the symposium.
Table 1 : Subdivision of the European continental Paleogene based on mammals as proposed bythe Symposium in Mainz, February 1987
30 CODERET
29 RICKENBACH
28 PECH DU FRAYSSE
27 BONINGEN
26 MAS oe pnurrtE
25 GAROUILLAS
24 HEIMERSHEIM
23 ITARDIES
22 VILLEBRAMAR
21 SOUMAILLE
Chattian
Rupelian
OLIOGOCENE
EOCENE
Stampian
20 ST. CAPRAISE D. E.
19 ESCAMPS
rs LR oEenuce
17 FONS4
16 ROBTAC
15 LA LIVINIERE
14 EGERKINGEN cr + F
13 GEISELTAL OMK(OBEREMITTELKOHLE)
12 GEISELTAL UMK(UNTEREMITTELKOHLE)
11 GEISELTAL UK(UNTERKOHLE)
10 GRAUVES
8-9 AVENAY
7 DORMML
6 CERNAY
1-5 HAININ
Priabonian
(Ludian) Latdorfian
Bartonian s.l.
Bartonian
Lutetian
Ypresian
PALEOCENE
Norbert SCHMIDT-KITTLER (ed.)

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STANDARD LEVELS SPAIN FRANCE SWITZERLAND W. GERMANY
6ssn ENGLAND
BELGIUM (B)
MP3O CODERET Fraga7,11
Bergasa La Colombidre, Coderet,
Venelles, Th6zel,
Dieupentale
Brochene-Fluh 53,
KLittigen Rottenbuch 2, 6,
Fl6rsheim,
Ehrenstein 4
MP29 RICKENBACH Villanueva del
Rebollar
Comberatidre, La Milloque,
La Mine des Roys,
S'Victor la Coste, Verneuil,
Puyde Mondoury
Rickenbach,
Rances
MP28 PECH DU FRAYSSE Viveldel Rio Portal,
Pech du Fraysse, Cournon,
Pech Desse Fornant 6, 7Gaimersheim 1
MP27 BONINGEN Fraga4,
TorradelCompte,
Minadel Pilar Boujac, Sardle Aarwangen 1
Boningen,
Wynau 1
Ehrenstein 7,
Burgmagerbein 1
Gaimersheim 2
MP26 MAS DE PAUFF]E Gandesa N (drilling) La Devdze, S'Andr6 (N.C.),
Mas de Pauffi6, "StHenri",
Espeyrasse
Miimliswyl-Hardberg,
Oensingen
MP25 GAROUILLAS Carrascosa, Campins
Garouillas, Rigal-Jouet,
Vialenc, Les Matelles,
Aubenas les Alpes,
S' Vincent de Barbeyrargues,
Belgarric
Bumbach Burgmagerbein 2,
Murnau 1,
Habach 4,5
MP24 HEIMERSHEIM L6bratidres 14, Etampes,
La Fert6-Alais, ltteville,
Maintenon Grenchen 1Heimersheim
MP23 ITARDIES Montalban,
Tarrega,
Peraltilla
SrMartin de Castillon,
Les Chapelins, Mounayne,
Itardies, Pech Crabit 1,
Roqueprune 2
Bernloch,
Schelklingen 1
MP22 VILLEBRAMAR Ollala 4, Calaf,
Fonollosa Mas de Got A et B,
La Plante 2,
Villebramar, Lovagny Balm Ronheim 1,
Herrlingen 1,
Mdhren 13, Weinheim
MP21 SOUMAILLES Santpedor,
Espinosade Henares Ronzon, Hoogbutsel, Ravet,
Ruch, Lagny-Thorigny,
Aubrelong 1, Soumailles
Mdhren 1 9, 20,
Ehrenstein 18,
Detan (dssR)
rel (B),
ctiff
MP2O ST, CAPRAISE Huermecesdel Cerro S'Capraise,
Tabarly, Baby 2,
Villeneuve la Comptal2
Frohnstetten,
Neustadt,
Nordshausen
Bay
MP19 ESCAMPS San Cugat
Coanac 1, Audincourt, Sb Croi;
de Beaumont 2, Rosidres 1 -3,
Montmartre, Mas S"" Puelles,
Escamps, Lascours, S' Martin
de Viller6al 2, Pontd'Assou
Mormont-Entreroches,
Obergosgen Neuhausen,
Mdhren 6,
WeiBenburg IHeadon Hill
(HH 6, HH 7)
MP18 LADEBRUGE
Gousnat, Sainte-N6boule,
StMartin de Viller6al 1 ,
S'Croix de Beaumont 1,
Civrac, La D6bruge Gdsgen-Kanal
Plaffenweiler,
Ehrenstein 1A,
Ehrenstein 2,3, 6,
Herrlingen 3
Lacey's Farm,
Quarry,
Headon Hill
(HH 3, HH 4)
Mtinchner Geowiss. Abh. (A) 10

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STANDARD LEVELS PORTUGAL (P)
SPAIN FRANCE SWITZERLAND W. GERMANY
E. GERMANY (GDR) ENGLAND
BELGIUM (B)
MP17 FONS4 Roc de Santa,
Sosis
Aubrelong 2, La Bouffie,
Baby 1, Les Clapids, Fons 1 -7,
Saldme, Les Pradigues,
Les Sorcidres, Penidre,
Malp6ri6, Rosidres 5, Euzel
La Cantine 2,
Lebratidres 1
Hordle,
Headon Hill
(HH1, HH2)
MP16 ROBIAC
Grisolles, Berville,
Lavergne, Le Bretou, Robiac,
Blaye, Le Castrais,
Paris (Monceau),
Paris (Gare du Nord)
Ecl6pens-Gare Heidenheim,
Herrlingen 4Creechbanow,
Hengistbury,
Barton D/E
MP15 LA LIVINIERE 2Pontils La Livinidre 2,
Cesseras
MP14 EGERKINGEN cr + PCapella Le Gu6pelle, Lissieu,
Arcis le Ponsart,
Fontliasmes, lssel Egerkingen a + pGeiseltal-Oberkohle
(GDR)
GEISELTAL-
MP13 OBERE
MITTELKOHLE
Bouxwiller, Aumelas,
La Defense, Dampleux,
ChAteau-Thierry
Geiseltal-Obere
Mittelkohle (GDR)
GEISELTAL-
MP12 UNTERE
MITTELKOHLE
Geiseltal-Untere
Mittelkohle (GDR)
GEISELTAL-
MP11 UNTERKOHLE Geiseltal-Unterkohle
(GDR),
Messel
MP1O GRAUVES Los Saleres
P16montr6, Cuis, Chavot,
Mancy, Monthelon, Venteuil,
Grauves, StAgnan,
Mas de Gimel, Azillanet East-Wittering
MP8+9 AVENAY S6zanne-Broyes, Brasles,
Cond6-en-Brie, Mutigny,
Avenay
Herne Bay,
Harwich
(London Clay),
AbbeyWood
MP7 DORMAAL Silveirinha (P) Pourcy, Meudon, Rians,
Erquelinnes, Palette
Suffolk Pebble
Beds (Kyson,
Ferry Cliff, Bram-
ford, Harwich),
Dormaal(B)
MP6 CERNAY Benu, Cernay,
Marnes de Rilly
MP1-5 HAININ ? Menat ? Walbeck (GDR) Hainin (B)
Table 2: Conelation chart of the most important European mammal localities as erected by the participants ol the Symposium in Mainz,
February 1987.
Norbert SCHMIDT-KITTLEB (ed.)

Miinchner Geowiss. Abh. (A) 10

21
Mammalian Reference Levels MP 1-10
coordinated by
M. GODINOT
The accompanying table of the reference faunas of the
Paleocene and early Eocene reflects the partial knowledge
that we presently have of these epochs in Europe. Few of
these faunas are sufficiently rich to provide a good repre-
sentation of the whole continent; several of them include
only micromammals. We have therelore added supple-
menting information from very well correlated faunas.
Faunas which are rich but not sufficiently well correlated
with the reference faunas have not been used to elaborate
these lists.
The lists which are given for these reference faunas are not
complete. We have limited ourselves to the species which
were adequately studied and to those which have particu-
lar biochronological significance. This explains the omis-
sion of species like Platychoerops daubrei and Corypho-
don eocaenus, the ranges of which in the Eocene are too
long. For some of these localities, more complete faunal
lists have been published previously (RUSSELL, 1964,
1980; GODINOT et al., 1978). Composite faunal lists and
comments on the geological background of the European
fossiliferous regions are to be found in RUSSELL et al.,
1982. The geological context and particularly the difficult
problems of limits in these epochs are discussed in
CAVELIER and POMEROL, 1986. The evolution of Euro-
pean mammalian faunas around the Paleocene-Eocene
transition was synthesized by RUSSELL (1975), and some
of the mammalian dispersal events were discussed by
GODINOT (1982). Recent studies on the faunas of Cer-
nay, Hainin and Silveirinha have modified our knowledge
of these epochs: although more mammalian genera are
known to have persisted from the Paleocene into the
Eocene, the boundary nevertheless represents the point
of the mostdramaticturnover in mammalian history.
ln the Paleocene-early Eocene table, an open numbering
system of 1 to 5 has been proposed to insure stability in the
future if new faunas are discovered. Hainin was chosen as
the reference-fauna because of its position within the
Montian strata in Belgium. lt is not possible at the moment
to place Walbeck and Menat relatively to Hainin, because
Menat has only four mammals and Walbeck has no
multituberculates and in neither locality is stratigraphic
control possible. The study of the Walbeck fauna showed
by the evolutionary grade of several lineages that this
fauna is older than that of Cernay (RUSSELL, 1964).
For the middle of the early Eocene in the table two numbers
have been provided for Avenay. ln the present state of
research, it is difficult to choose, among several succes-
Norbert SCHMIDT-KITTLER (ed.)
sive faunas, the most significant one for a European
reference-level. The fauna of Cond6-en-Brie is rich but
little studied, and is also not very different from that of
Avenay. The Mutigny fauna is older than that of Avenay,
but until now not distinguished from it by a large number of
taxa. The most important differences come from the
revision of the marsupials, which showed within two line-
ages different species in Mutigny and Avenay: lhe Pera-
therium @nstans - P. matronense lineage and the Amphi-
peratherium brabantense -A. sp. 1 lineage (CROCHET,
1980); from this results the artificial overlap of these
species in the table. The fauna of Pourcy could also be a
reference-fauna; it is placed here in the reference-levelof
Dormaal, but it shows the first appearance of a series of
species otherwise known in Avenay and correlated
faunas: Apatemys sigogneaui (RUSSELL et al., 1979),
Microparamys russelli and M. chandoni (HARTEN-
BERGER, 1971), Diacodexis varleti and Bunophorus
cappeftai (SUDRE et al., 1983). These faunas and others
f rom the same period are still under study, and it appears
better to leave for the future the choice of the most
significant reference fauna if more are added to this
interval.
ln the table, the name Sables i Unios et T6r6dines has
been used because we were not able to work with a faunal
list concerning only Grauves. This will be done in the future.
Nevertheless, this locality and the others from the Sables
d Unios are very wellconelated as they are allwithin a river
channelcontained in the Cuisian Sands. The reference-
levelconstituted by all these localities is then an excellent
one. Otherwell-correlated faunas have been used in the
table when they added important information. This is the
case of the Suffolk Pebble Bed localities, complementing
the fauna of Dormaal (which lacks multituberculates,
perissodactyls, Adapidae ...). Other English localities allow
the placement of species ol Hyracotherium , which are
important and recently revised (HOOKER, 1980, 1984).
ln sum, this table shows a selection of the information that
is considered the most significant lor the establishment of
a biochronological framework of European mammalian
faunas. Regionalscales may be more refined, as in the
Paris Basin where at least eight su@essive faunas can be
distinguished between Meudon or Pourcy, and Pr6montr6.
The faunal lists are not complete. They are intended to be
working tools for broad geochronological purposes. ln no
case can they be considered a synthesis of mammalian
evolution in Europe atthattime.

22
References
CAVELIER, C. & POMEROL, C. (1986): Stratigraphy of the Paleo-
gene. - Bull. Soc. G6ol. France, (8), 2: 255-265, Paris.
CROCHET, J.-Y. (1980): Les Marsupiaux du Tertiaire d'Europe. -
279 pp.,241 figs., 40 tab., 2 pls., Paris (Fondation Singer-
Polignac).
GODINOT, M. (1982): Aspects nouveaux des 6changes entre les
faunes mammaliennes d'Europe etd'Am6rique du Nord A la
base de I'Eocdne. - Geobios, m6m. sp6c., 6:403-412,2
figs., Lyon.
GODINOT, M., de BROIN, F., BUFFETAUT, E., RAGE, J.-C. &
RUSSELL, D. (1978): Dormaal: une des plus anciennes
faunes eocdnes d'Europe. - C.R. Acad. Sci. Paris, (D), 287:
1273-1276, Paris.
HARTENBERGER, J.-1. (1971 ): Contribution i l'6tude des genres
Gliravus el Microparamys (Rodentia) de I'Eocdne d'Euro-
pe. - Palaeovertebrata, 4:97-135, 18 figs., 2 tab., 5 pls.,
Montpellier.
HOOKER, J.J. (1980): The succession of Hyracotherium (Perisso-
dactyla, Mammalia) in the English early Eocene. - Bull. Br.
Mus. nat. Hist. (Geol.), 33: 101-1 14, 6 figs., London.
_ (198a): A primitive ceratomorph (Perissodactyla, Mam-
malia) from the early Tertiary of Europe. - Zool. J. Lin. Soc.,
82: 229-244, 22 tigs., London.
RUSSELL, D.E. (1964): Les Mammifdres paleocdnes d'Europe. -
M6m. Mus. nat. Hist. Nat., (c), 13:1-324,74 figs., 16 pls.,
Paris.
_ (1975): Paleoecology of the Paleocene-Eocene Transition
in Europe. - ln: F.S. SZALAY (ed.), Approaches to Primate
Paleobiology. -Contr.to Primatology,5:28-61, 7figs., Basel
(Karger).
_ (1980): Sur les condylarthres cernaysiens Tricuspiodon el
Landenodon (Paleocene Sup6rieurde France). - Palaeo-
vertebrata, M6m. Jubil. R. Lavocat: 127-166,2 figs., 4 pls.,
Montpellier.
RUSSELL, D.E., GODINOT, M., LOUIS, P. & SAVAGE, D.E.
(1979): Apatotheria (Mammalia) de l'Eocdne inf6rieur de
France et de Belgique. - Bull. Mus. nat. Hist. Nat., (4, c), 1:
203-243, 4 figs., 2 tab., 5 pls., Paris.
RUSSELL, 0.E., HARTENBERGER, J.-1., POMEROL, C., SEN,
s., scHM|DT-K|TTLER, N. & V|ANEy-LrAUD, M. (1982):
Mammals and Stratigraphy: The Paleogene of Europe. -
Palaeovertebrata, M6m. extraord. : 1 -77, 23 figs., Montpel-
lier.
suDRE, J., RUSSELL, D.E., LOU|S, P. & SAVAGE, D.E. (1983):
Les Artiodactyles de l'Eocdne inf6rieur d'Europe. - Bull.
Mus. nat. Hist. Nat., (4, c), 5: 281-333, 339-365, 20 ligs, 3
tab., Paris.
Table 3: Occurrence of mammdian taxa in the reference faunas of MP 1-10 and well correlated localities. Chronospecies or -sub-
species forming evolutionary stages of lineages are marked by an asterix. Coordinated by M. Godinot.
Munchner Geowiss. Abh. (A) 10

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MP
units Reference
locality
Taxa occurring in the reference fauna
Taxa restricted to the I rirst appearance I r-ast occurrence
reference level I I
Taxa occurring in well correlated
I
Taxa restricted to the First appearance
reterence level
localities
Last occurrence
MP 10 GRAUVES
Ailuravus michauxi
Plesiarctomys savagei
Hyrachyus stehlini
Propachynolophus gaudryi
Prcpachyrclophus maldani
Aumelasia renieli
Cuisithetium lydekkeil
Palawhiopteryx cl. tupaiodon
Lophiodon rerensis Peradectes louisi
Amphiperatheium mdimum
Arcius lapparcnl
Donrussellia gtallia
Meldimys louisi
Protoadapis rectiilspidens
Protadapis curyicuspidens
Oxyaena menui
Frun@lherium lindgteni
Peratheium matronense
Amphiperatheium sp.'l
Canlius savagei
Lophiaspis cl. maurelti
MP 8+9 AVENAY
Pendetes russelli
Petadectes mutgnyensis
Apatemys mutiniacus
Apatemys sigogneaui
N@matrorella lrciannae
laronycteis ? menui
Archaeonycteis brailloni
Donrussellia louisi
Donrussellia russelli
Patamys ageiensis
Paramys woodi
' Mbroparamys russelli
' Microparamys chandoni
Placentidens lotus
Bunophorus caweltai
Diacodexis varlet
Peradectes louisi
Perutherium matronense
Amphiperathedum sp.1
Amphiryralheilum muimum
Arcius lapparanti
Donrussellia gallia
Meldimys louisi
Paramys w@di
Pseudopanmys teilhardi Arcius fuscus
Cantius erysi
Hyruntheilum lepotinum
Hyracotheilum vulpiceps
Cantius savagei
Lophiaspis cl. maurclti Neoplagiaulax all. sylvani
Peratheium constans
Amphipeatheium brubantense
Beruvius cf. lasseroni
Hyopsodus wardi
MP7 DORMAAL
' Landenodon wouterei
Eochircmys landenensis
Apatemys teilhardi
Teilhatdina belgica
Prolimnocyon n.sp.
Paschathedum dolloi
Microhyus musulus
Palaeonictis gigantea
" Micropatamys nanus
Paramys cl. woodi
Peratherium constans
Amphiperathedum brubantense
Pseudoparumys leilhardi
Plesiadapis cl. ttbuspidens
Plesiadapis all. remensis
Cymbalophus cuniculus
Cantius cl. ewi
Hyopsodus wardi
MP6 CERNAY
N@plagiaulax eoaanus
N@plagiaulil @pei
Neplagiaulax nicolai
Liotomus marehi
Landenodon phelizoni
Landenodon laveati
Cemapia manueli
Adapisorex gaudryi
Chiromyoides campanius
Plesiadapis tricuspidens
Disscus europails
Louisina mkabilis
Arct@yon pilmaevus
Arctocyonides trcuessatli
Tricuspiodon ruelimeyed
Pleuraspidotherium aumoniei
Otlhaspidolhailum e&rardsi
Hainina godhiauxi N@plagiaulax sylvani
Berruvius lassroni
MP 1-5 HAININ
Hainina belgia
Bottius splendidus
Peradectes maandati
Monshyus praevius
Prclatidens waudruae
Hainina godlilauxi
ru
q)

24
Mammalian Reference Levels MP 11-13
coordinated by
J. L. FRANZEN
The mammalian biostratigraphy of the European Middle
Eocene (= Geiseltalian sensu FRANZEN & HAUBOLD
1986) is mainly based on a series of reference levels all
coming from the geologic section of the Geiseltal locality
(German Democratic Republic). Thus there can be no
doubt concerning the gradient of time and the sequence of
reference levels.
Problems still exist with respect to
1) the lack of MP-Unit 1 0 at the Geiseltal locality,
2) the lack of a geologic section comprising mammalian
faunas of MP-Units 10 and 1 1 anywhere,
3) thepoordocumentation of MP-Unit 1 4byfossil mam-
mals coming from the Geiseltal "Oberko6ls" (OK),
4) the uncertainty concerning the biostratigraphic age of
the various fissure fillings of the Egerkingen locality,
and
5) the fact that some major revisions, e.g. of Primates,
Creodonts, Artiodactyls, and Perissodactyls, are not
accomplished as yet, while others, e.g. of the Egerkin-
gen fauna as a whole, would be highly desirable.
The present def inition of the upper limit of the European
Middle Eocene of mammalian biostratigraphy is mainly
based on a correlation with the classical localities of the
Paris basin, while the definition of the lower limit is due to
evolutionary lineages, and a direct correlation with the
palynologic time scale atthe Geiseltal locality. The refer-
ence level of Egerkingen is restricted to the fauna of fissure
fillings a and p.
Only recently some uncertainty arose concerning the
correlation of the mammalfaunaof lssel(Southern Fran-
ce). Based on the Equoidea this locality should be classed
with MP 12 (presence ol Propalaeotherium isselanum
instead of P. helveticum and P. hassiacum respectively,
lack of any Palaeothere sensu stricto). Recent investiga-
tions particularly of the Lophiodonts however, are told to
suggest a post-Lutetian age (MARANDAT, this volume).
Table4: Occurrence of mammalian taxa in the reference faunas of MP 11-13 and well correlated localities. Chronospecies or -sub-
species forming evolutionary stages of lineages are marked by an asterix. Coordinated by J. L. Franzen.
Mrinchner Goowiss. Abh. (A) 10

z
o
o
f
6
I
=
o
I
x
I
+
-
m
u
o
eMP
units Relerence
locality
Taxa occurring in the reference fauna
Taxa restricted to the I first appearance I Ust occurrence
reference level I
Taxa occurring in well correlated localities
Taxa restricted to the I first appearance J Ust occurrence
reterence level I I
MP 13 GEISELTAL
OBERE MITTELKOHLE
Amphilemur eoAenicus
Matth*ia germanica
Pugiodens mius
Europolemur klafti
Nannopithex raabi
Messelobunodon ceciliensis
Anthru@bunodon weigelti
Amphiperathedum giselense
Quercygale helvetica
Lophiotheium pygmaeum
Plagiolophus caniei
Lophiodon uviei
Hapbbunodon cl. muelleil
Rhagatherium kowalevskyi
Prcvivetra gtacilis
Hallensia matthesi
Propalaeotherium voigti
. Prcpalaeothedum isselanum
Paralophiodon buxovillanum
Heterchyus armalus
Alienetherium buwilleri
' Buxolestes hammeli
Praecodens acutus
Alsatbopitheus leemanni
Ped@nodon huetzeleri
Europolemur dunaifi
Paraplagiolophus codiciensis
Buxobune daufueei
Cebehoerus jaegeri
Ceb@hoerus dawsoni
Hyperdichobure hammeli
Tapirulus majoi
' Aurelasia gabireaudi
Dichodon ruatimsyeil
Protadelomys alsaticus
Eogliravus hammeli
Amphipeatheium basbeeense
Archilophus d. depereti
Pachynolophus duvali
Palaeotherium eMenum
Lophiodon tapiroides
Lophiodon leptorhynchum
Menis@don europaeum
Dacrytherium cl. elegans
' Ailuravus picteti
Plesiarctomys spectabilis
Amphiperatheilum goethi
MP 12 GEISELTAL
UNTERE MITTELKOHLE
Europolemur weigelti Cynohyaenodon trux
' Propalaeotherium isselanum
Parulophiodon buxovillanum
Ptodissopsalis ginsburgi
Lophiodon tilholi Provivetra typica
Chasmotheilum Gftied
MP 11 GEISELTAL
UNTERKOHLE
Oxyaenoides bicuspidens
Vulpavoides germanicus
Esthonyx tardus
Prcpalaeotheium argentonicum
Prcpalaeotherium ha,siacum
Lophiothedum n.sp.
Provivera gracilis
Hallensia matthesi
Propalaeotheium paNulum
Propalaeotherium voigti
' Hyrachyus minimus
Leptictidium tobieni
Leptictidium nasutum
Leptictidium auderiense
Buxolestes pisator
Macrccranion lenerum
Pholidocercus hassiacus
Eomanis waldi
Eurotamandua joresi
Palaeochiropteryx spiegeli
Archaeonycteds ttigonodon
Hassianycteris magna
Ha$ianycteris messelensis
Europolemu koenigswaldi
Prcvivefta edingeri
Parcodectes feisti
Miacis ? kessleil
Kopidodon macrognathus
Massilabune maftini
Meselobunodon schaefed
Massilamys beegeri
Massilamys krugi
Micrcpatamys paNus
Ailuravus macrurus
Ptodissopsalis ffiaenius
Albpterodon theriodis
' Aumelasia cl. gabineaudi
Palae@hhopteryx tupaiodon
]\)
(tl

26
Mammalian Reference Levels MP 14-16
coordinated by
J. J. HOOKER
ln deciding on reference localities for the Bartonian,
Robiac was the obvious choice for the most recent faunas.
It occurs in a stratif ied sequence and has a diverse fauna
that is widespread in Europe. lt is easif differentiated from
later faunas by its last occurrences of many taxa, espe-
cially lophiodonts and certain species of palaeothere.
Other faunas attributed to the Bartonian were less easy to
characterise. La Livinidre 2 is demonstrably older than
Robiac according to the evolutionary stage of ils Pseudol-
tinomys and last appearance ol Lophiodon leptorhyn-
chum. lls fauna is rather small, however, and not very
widespread, although it is a stratified locality.
Characterising the lower limit of the Bartonian also posed
problems as stratified faunas of this age arc tare and of low
diversity, so the rich karst localities of Egerkingen a + p
were chosen to typify this interval. lt was felt that the faunas
of the other Egerkingen fissures - yand the Huppersand,
Grey Marl Facies and Aberrant Facies - had too much in
common with Geiseltal oMK, Bouxwiller and the Upper
Calcaire Grossier localities for their Bartonian age to be
undoubted. A further problem is that as there are no
lineages in common between Egerkingen cr + p and La
Livinidre 2, it cannot be excluded that they may be
contemporaneous. The faunaldifferences are likely to be
mainlythe result of ecologicaland collecting bias.
Certain correlations can be made with the marine province
via some of the referred stratified localities. All probably fit
into the type Bartonian of Mayer Eymar (1856), but the
base of the Bartonian as currently used by marine micro-
fossilworkers (base of P13 = upper part of NP16; see
HARDENBOL & BERGGREN 1978)is probably between
La Livinidre 2 and Robiac (see ANADON et al. 1983;
HOOKER 1986:415-429).
References
ANADON, P., FEIST, M., HARTENBERGER, J.-1., tvtUt-len, C. a
VILLALTA-COMELLA, J. de (1983): Un exemple de corr6-
lation biostratigraphique entre 6chelles marines et conti-
nentales dans I'Eocdne: la coupe de Pontils (bassin de
l'Ebre, Espagne). - Bull. Soc. 96ol. France, (7),25, (5):747-
755.
HARDENBOL, J. & BERGGREN, W.A. (1978): A new Paleogene
numerical time scale. ln: COHEE, G.V., GLAESSNER,
M. F. & HEDBERG, H.D., Contributions to the geologic time
scale. - Stud. Geol. Am. Assoc. Pet. Geol.,6: 213-234.
HARTENBERGER, J.-1., SrGE, B. & SUDRE, J. (1968):Nouveaux
gisements de Vert6br6s dans l'Eocdne continental du
Minervois. - C.R. somm. S6anc. Soc. 96ol. France, 1968
(1):22-23.
HOOKER, J.J. (1968): Mammals from the Bartonian (middle/late
Eocene) of the Hampshire Basin, southern England. - Bull.
Br. Mus. nat. Hist., (Geol.) 39 (4):191-478.
MAYER EYMAR, K. (1857): Versuch einer neuen Klassifikation der
Tertidr-Gebilde Europa's. - Verh. schweiz. naturf . Ges., 42:
1 65-1 99.
Table5: Occurrence of mammalian taxa in the reference faunas of MP 14-16 and well correlated localities. Chronospecies or -sub-
species forming evolutionary stages of lineages are marked by an asterix. Coordinated by J. J. Hooker.
Munchner Geowiss. Abh. (A) 10

z
o
q
o
f
@
I
o
i
x
=
-..1
-
m
o
!o
MP
units Reference
locality
Taxa occurring in the reference fauna
Taxa restricted to the First appearanc" I Last occurrence
reference level I
Taxa occurring in well correlated localities
Taxa restricted to the I rir.t appearance I urt occurrence
reference level I
MP 16 ROBIAC
Nerolemu cl. antiquus
Adapis sudtei
Plesiarctomys hutzelei
Ailuravus stehlinschaubi
Sciuroid$ siderclilhius
(= Suevosciurus romani)
Simamphicyon helveticus
Lophiothedum siderolithbun
(= L. robiacense)
Leptolophus s/€hlini
Palaeotheilum pomli
P. castrense robiacense
Lophiodon lauti@nse
A@therulum campichii
Cefuchoerus robia@nsis
Choeropotamus lautricensis
Tapirulus schlossei
Dactytheium elegans
Catodonthedum rcAacense
Robiatheium @urnovense
Rodeina minuta
Xiphodon astrene
Haplomeryx pictei
Amphiperathedum lontense
Pseudoloris paruulus
Helerohyus sudrei
Helerchyus nanus
Ptololomus ? cl. minor
Paracynohyaenodon schlossei
Parcxyaena galliae
Cynohyaenodon lautri@nsis
Paramiacis exilis
Quercygale angustidens
Remys minimus
Archilophus gaudini
Anchilophus dumasii
Plagiolophus ann%tens
Palaetherium siderolithicum
Pseudamphimeryx rcneviei
Pseudamphimeryx pavloviae
Peratherium sudrci
Saturninia mamertensis
' Satuninia hanenbergei
Gliravus robiacensis
' Effomys tobieni
Palaeotherium ruetimeyei
Anchilophus desmarasti
Chasmotherium caftiei
Peralherium bretouense
Satuminia gtandis
Saturninia gilsollensis
Gesnercpilhex ligularis
Nannopithex quaylei
Microchoerus wardi
Mbrochoerus crcechbanowensi1
Pseudolotis crusalonli
Europolemur collinsonae
Suevosciurus authodon
Sciuroides ilssonei
Sciuroides russelli
Theridomys varleti
Heterchyus modnionensis
Vulpavoides @operi
Haplobunodon venatorum
Antfuacobunodon louisi
Plagiolophus cuttisi
P. @tlailhaci
Palaeotheilum lautricense
Amphiperathedum minutum
Peruthedum lavergnense
' Amphiperalheilum bourdellense
Satuminia beata
Plesiarctomys curranti
' Amphiperatheium giselense
Amphipetathedum alt. g@thei
Saturninia intermedia
MP 15 LA LIVINIERE 2
' Pseudoltinomys @setanus
Pachyrclophus livinierensis 1)
Pachynolophus @ssetasicus 1)
Petatheilum sudrei
' Saluminia harTenbergeri
Saturninia intermedia
Gliravus tobiacensis
Parudelomys crusalonti
. Ellonys tobieni
Lophiodon leplorhynchum
MP 14 EGERKINGEN c + p
Necrolemu cl. zifteli
Leptadapis ruetmeyeri
Prcpalaeothedum helvetiilm
Mouillacithedum canieil
Ceb@hoerus ruetimeyeri
Tapiulus depereti
Dichodon catlieri
Palaeotheium ruetimeyei
Palaeotheium castrense
Mixtothedum inlans
Mixtothedum gresslyi (el all.)
Plesiarctomys speclabilis
Ailuruvus picteti
Protadelomys caftieri
Provivefta typica
Alloptercdon thedodis
Prodissopsalis eocaenicus
Anchilophus deperel
Lophiotheium pygmaeum
Plagiolophus caftiei
Palaeotherium e@aenum
Lophiodon rhinocerodes
Lophiodon Nviei
Hyperdichobune nobilis
Meniscodon europaeum
Haplobunodon soloduren*
Haplobunodon muell€ri
Rhagatheium kowalevskyi
Catodontherium fallax
Haploreryx ege*ingensis
Pseudamphimeryx schlossen
Europolemur alf. kJalti
Pivetonia isabenae
' Proadelomys lugdunensis
Hyperdichobune langi
Pachynolophus all. duvalii Amphiperuthedum bastbetgense
Cebochoerus suillus

28
Mammalian Reference Levels MP 17-20
coordinated by
S. LEGENDRE
The Ludian comprises four reference levels (MP 17 to MP
20). The chronological sequence in the Ludian is mainly
based on Palaeotherium taxa (FRANZEN 1968).
The first level, Fons 4, can easily be separated from the
earlier Bartonian level of Robiac (see comments on the
Bartonian tables). The levelof Perridrethough proposed
on the basis of rodent evolution (HARTENBERGER 1973)
and apparently well defined in the Hampshire Basin
(HOOKER this volume), had not been retained because
the arguments were considered to be too weak. Therefore,
the localities of Perridre and others are correlated to the
level of Fons 4 in the tables. The fauna of Fons 4 was
preferred to that of Euzet because the faunal spectrum is
broader, particularly regarding small-sized species.
The next higher level, La D6bruge, iswelldefined, princi-
pally on the basis ot the Palaeotherium group. lt is
characterized by a great number of first appearances.
The locality of Escamps was preferred to that of Montmar-
tre as reference level because itcontains a large number
of small species in addition to larger ones, even though the
Palaeotherium species are not well documented. The
presence of Palaeotherium medium medium indicates an
age for Escamps close to that of Montmartre.
The last Ludian level, Saint-Capraise, is recognized on the
basis of Palaeotherium species. Being the latest level
before the "Grande Coupure", it is important for the
understanding of this faunal phenomenon. So it is chosen
though small mammals, especially rodents, are scarce
and there are no positive arguments other than Palaeo-
therium. Saint-Capraise was preferred to Frohnstetten
because it is stratified.
References
FRANZEN, J. L. (1968): Revision der Gattung Palaeotherium
CUVIER 1804 (Palaeotheriidae, Perissodactyla, Mam-
malia). - lnaug. Diss. Albert Ludwigs-Universitdt, 181 p.,
Freiburg.
HARTENBERGER, J.-L. (1973): Etude syst6matique des Therid-
omyoidea (Rodentia) de l'Eocdne supdrieur. - Mdm. Soc.
g6ol. France, n.S., 1'17: 1-76, Paris.
HOOKER, J. J. (this volume): Mammalian faunal events in the
English Hampshire Basin (late Eocene - early Oligocene)
and their application to European biostratigraphy.
Tabfe6: Occurrence of mammalian taxa in the reference faunas of MP 17-20 and well conelated localities. Chronospecies or -sub-
species lorming evolutionary stages ol lineages are marked by an asterix. Coordinated by S. Legendre.
Munchner Geowiss. Abh. (A) t0

z
o
q
o
f
U'
o
I
=
o
--1
x
-..1
-
m
t
6'
I
MP
units ReJerence
locality
Taxa occurring in the reference fauna
Taxa restricted to the ] rirst appearance I ust occurrence
reference level I
Taxa occurring in well correlated localities
Taxa restricted to the I first appearance Last occurrence
reference level I
MP 20 SAINT CAPRAISE
' Palaeotherium cuftum frchn.
stettense Paragelocus suevicus
. Palaeotherium medium suevi-
cum
Pterodon dasyuroid*
Patriotheidomys altus
Choeropotamus parisiensis
Anoplotherium @mmune
Diplobune s&undaria
Xiphodon gracile
Palaeotheilum magnum
Theridomys bondueili
Amphirhagatherium lrchnstet
tense
Cryptopithecus sidercolithicus Peratheium . tuviei
Amphidozotherium ayluxi
Saturninia tobieni
Adapis betillei
Paradelomys spelaeus
Oltinonys platyceps
Blainvillimys rotundidens
Oxacron couttoisi
Haplomeryx zilteli
Dichodon lrohnsteftense
Amphimeryx murinus
Palaeothedum muehhergi
MP 19 ESCAMPS
"Cynodictis. comprcssidens
' Pseudoltinomys cuvieil
Paroxacrcn sp.
' Palaeotheium medium medium
Amphiperatheium ambiguum
Amphiperutheium exile
Saturninia tobieni
Amphidozothedum cayluxi
Adapis betiilei
Oltinomys plalyceps
Plagiolophus fraasi
Palaeotherium magnum mag-
num
Pseudohymcyon ayluxi
Cynodyctis lacusttis
Suevosciurus minimus
'Acotherulum satuminum
Cuvierimops padsiensis
Leptadapis assolicus [=stintoni]
Adapis paisiensis
Microchoerus ornatus
Theddomys golpei
Moiachoerus simpsoni
Haplomeryx (?) obliquus
Palaeotheium Ndum curtum
Palaeotherium rcnevied
Palaeotheilum crassum crassum
Hyaenodon geryaisi
Parapterodon lostangensis Heterohyus nanus
Satuninia beala
Pseudolotis paruulus
Pseudolotis rcgnanti
' Prototomus ?minot
Plesiarctomys geNaisi
Anoplotheium latipes
Anoplotherium laudllatdi
Plagiolophus annectens
Archilophus ndegondensis
Palaeotherium siderclithicum
'Palaeotherium duvali
MP 18 LA DEBRUGE
Perathedum cayluxi
Thetidomys pseudosiderclithicus
Theidomys petrealensis
Elomeryx crispus
Dichodon stehlini
Palaeotheilum magnum gircndL
cum
Palaeolheium crassum ro-
bustum
Pala@therium medium prrea-
lense
Plerodon dasyuroides
Cynodiclis lacusttis
Plesiarctomys geruaisi
Dichobune leporina
Choeropotamus parisiensis
Dactythedum saturninii
Taphulus hyracinus
Anoplotheium commune _
Anoplotherium latipes
Anoplothedum laurillardi
Oxacron coutloisi
Xiphodon gacile
Dichodon frohnslettense
Amphimetyx murinus
Diplobune sflndaia
Acotherulum satuminum
Plagiolophus minor
Anchilophus radegondensis
Palaeotheium duvali duvali
Palaeotheium m. muehlbergi
' Hyaenodon raquieni
' Palaeotherium cuftum villerea'
lens6
Cryptadapis teftius
' Pseudoltinomys gousnalensis
Mouillacitheium schlossei
Paroxacron valdense
' Palaeotherium muehlbergi tha-
leri
Hyaercdon hebefti
Paradelomys spelaeus
' Elainvillimys totundidens
Patriotheridomys altus
. Haplomeryx zitteli
Hyaenodon minor
Quercygale angustidens
A@therulum pumilum
Dacrytherium ovinum
Leptotheildium lugeoni
Tapirulus peftierensis
MP 17
Eslellomys cansouni
Elfomys paNulus
Pseudoltinomys mamertensis
Remys gaimondi
Theridomys euzetensis
Cebochoerus lontensis
Choeropotamus sudrci
Xiphodon intemedium
Patiliphodon teulonense
Pachynolophus gailmondi
Palaeothedum medium euze-
tense
Lophiotheium eruulum
Hyaenodon minot
Hyaercdon requieni
Quercytherium lenebrosum
Suevosciurus minimus
Treposciurus mutabilis
Dacrytherium ovinum
Haplomeryx euzetensis
Amphiperutheilum bourdellense
Amphiperatherium fontense
Peratheium lavergnense
Gliavus all. tobiacensis
Paradelomys crusalonti
Treposciurus [= Sciuroides] in-
termedius
Mouillacithedum elegans
Pseudamphimeryx reneviei
Anchilophus dumasi
Archilophus gaudini
Peratheilum peniercnse
Necrcmantis adichaster
Lepladapis magnus
Pseudollinomys phosphoicus
Cebochoerus lacusttis
Mixtotherium cuspidatum
Robiacina qrercyi
Choeropotamus depereti
Diplopus aymadi
Pseudamphimeryx hantonensis
Dichodon ilspidalum
Palaeothedum magnum stehlini
Palaeotheium duvali pisum
Palaeothedum muehlbergi prae-
cursum
Palaeotheilum franzeni
Palaeotherium crusatonti
Peralherium cuviei
Pseudorhyncocyon cayluxi
Hyaenodn brachyrhynchus
Prototomus ? buhosus
Ptototomus ? minor
Sciutoides ehrensteinensis
Tapirulus penierensis
A@therulum pumilum
Palaeotherium cuftum villerea'
lense
Heterchyus sudrei
Paroxyaena galliae
Paracynohyaenodon schlossed
Miacis exilis
Anchomomys quercyi
Remys minimus
Pseudamphimeryx pavloviae
Haplobunodon lydeliJei
Propalaeotheium sp.
lU
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MP
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locality
Taxa occurring in the reference fauna
Taxa restricted to the I first appearance I Ust occurrence
reference level I I
Taxa occurring in well correlated localities
Taxa restrided fo the I first appearance I Urt occurrence
reterence level I I
MP 30 CODEBET
Hipposideros (Brachiwosideros)
bransatensis
lssiodoromys bransatensis
Archaeomys laudllardi
Eucri@todon longidens
Plesiosminthus schaubi
' Rhodanomys transiens Gliravus
Archaeomys
Adelomyarion virati
Rhizospalax
Drcmolherium guthi
Columbomys lavocati Amphiperatheium exile
Pseud@ricetodon thaleri
Plesispermophilus
Microbunodon
Bedenomeryx millquensis
Drcmotherium quercyi
MP 29 RICKENBACH
lssiodoromys pseudanaema
Archaeomys helvetius
Eucricelodon pra&usor
Eomys huezeleri
Ro%otherium romani
Rhizospalax
Adelomyarion vireti Anthacotherium cl. magnum Hiwosidercs (Brachipposidercs)
Pseudrcd@todon thalei
Microbunodon
Benedenomeryx milloquereis
Palawhoerus gergovianus
MP 28 PECH DU FRAYSSE
Amphictis ambiguus
lssiodorcmys limognensis
Eomys quercyi
Eomys gigas
Archaeomys intermedius
Prcdrcmotherium elongatum
Drcmotherium quercyi Plesiosminthus promyailon
Eomys zitteli
Eucri@todon dubius
Doli@hoerus quercyi
Bachitherium lavocati
Anthra@lheilum valdense Paratalpa micheli
Dremotherium guthi Pseudrcil@todon philiryi
Lophiomeryx chalaniali
Caenomeryx procommunis
Plesiomeryx adurcensis
Ronzotheilum

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MP 27 BONINGEN ' lssiodorcmys quercyi
'Archaeomys hueEeleil Peratherium antiquum
Eucilcetodon dubius Pseudocilcetodon inrertus Anlhracothedum hippoi(hum Palaeochoerus gergovianus Bachitherium cl. culum
MP 26 MAS DE PAUFFIE
Blainviilimys gominatus
lssiodoromys paultiensis
Archaeomys major
Plesiosminthus prcmyadon
Eomys zitteli Amphiperathedum minutum
Peratherium elegans
Neurcgymnurus cayluxi
Blainvillimys
Pseudoaicetodon moguntiacus
Eucicetodon hubei
Bachitheium insone
Metriotherium mhabile
Elomeryx bofuoniils
Anthracolheium bumbachense
Anthracotherium cuvieri
MP 25 GAROUILLAS
Blainvillimys blainvillei
lssiodorcmys minot
Archa@mys grucilis
Archaeomys geryaisi
Sciuromys quercyi
Cadurcotherium cayluxi
Schizotheium modium
Archaeomys
Pseudocrbetodon ircertus
Bachitherium insune
Anthracotheium magnum
Sciuromys
Amphiperathedum ambiguum Anthracotherium cuvieil
Anthracotheium bumbachense Gliruvus Enuis
Blainvillimys heime6heimensis
MP 24 HEIMERSHEIM ' Taeniodus hexalophodus Euctic€todon huberi
Pseudocicetodon moguntiacus
Blainvillimys heimereheimensis
Gliravus tenuis
Neurogymnurus cayluxi
Ebmeryx boboniils
Metiotheium mirabile
Tetacus nanus
MP 23 ITARDIES
Blainvillimys helmei
PsudMiEtodm montalba-
nensis
lbercmeryx minus
Bachitherium vireti
Bachitheium cunum Amphiryrethedum lamandini
Datbonetus aubrelongensis
Ellomys medius
Lophiomeryx mouchelini
. Taeniodus ilruistiatus
'Theidomys major
Ellomys nanus
Pseud@ricetodon philiwi
Doliochoerus quetcyi Eusmilus bidentatus
Pseudosciurus suevicus
' Sciuromys cayluxi
Eudi@todon atavus
Eomys antiquus
Anthra@theium alsalitum
Bunobrachyodus
Diplobune minot
Parcrecron bergei
Pseudogelous scolti
MP 22 VILLEBRAMAR
Pseudoltircmys major
Blainvillimys grcgailus
Entelodon deguilhemi
Gelocus vilbbramarcnsis
DichobJne jehennei
Ellomys medius
Lophioretyx mouchelini Stenogale gracilis
Stenopbsictis minor
Entelodon
Plagiolophus trasi
' Ibercmeryx matsoui
Elomeryx cluai Elomeryx porcinus Cryptopithws siderclithicus
Plagiolophus minot
MP 21 SOUMAILLE
Blainvillimp langeae
Theidomys aquatilis
Pseudoltinomys gaillardi
Entelodon antiquum
Gelocus communis
Acotherulum quercyi
Eggysodon sp.
Dafuonetus aubrclo|ansis
Eusmilus bidenatus
Plagiolophus minot
Entelodon
'Palaeothedum medium Shamolagus
Tarnomys quercyi
Entelodon magnum
Dichobune leporina
Bothriodon leptorhynchus
Bothtiodon aymatdi
Bothriodon valaunus
Anthru@therium dalmatinum
Anlhncothedum stehlini
Elomeryx wodi
P€eudogelocus suevicus
Pseudopalaeotheium longi-
rostratum
Petatheilum el€gans
Tetruils nanus
Butselia bived
Amphicynodon
Steneoliber
' Sciuromys cayluxi
Eucietodon atavus
Eomys antiquus
Plesispermophilus
Palaeosciutus
Caenomeryx ptocommunb
Plesioreryx adurcensis
Diplobune mircr
Bunobrcchyodus
Elomeryx porcinus
Anthracotheium alsatium
Ronzothailum
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