UPDATING OF MN ZONES

P.Mein

Departement des Sciences de la Terre

27-43 Bd. du 11 Novembre
696ZZ Villeurbanne Cedex, France
The recent numerous advances in our knowledge about fossil mammals have
required a review and revision of the MN zonation (Mein, 1975, 1976).
In some areas (especially in Spain) the succession of biological events among
Cenozoic mammals can be identified in a very sharp manner, especially with the
evolutionary grade method of some rodent lineages.

Unfortunately, small mammal species generally have a limited geographical

distribution that does not allow long distance correlations. So in this new chart,
biological events are mentioned only at the generic level with special references to
faunal records of west and central Europe.
The revised chart of MN zones, common taxa, generic first appearances in
Europe, and generic last occurrences in Europe are shown in table 1. Table 1 follows
the format of previous charts of MN zones (e.g., 1975 and 1979) with numerous additions and slight revision. MN zones are grouped in table 1 according to mammal ages
of Fahlbusch (1976). Another notable change is that the designated reference fauna
for each subdivision is printed in capitals. The most important biological event is the
first appearance of a new genus, but we also note the last occurrences.' For each
mammalian age, the list of the main genera are also given.
To allow greater ease of correlation we have tried to simplify the subdivision
and have retained formal subdivision (a and b) only for MN z.
For the same reason, we have not split the last Neogene unit MN 17 (corresponding to MN 17 + 18 of Guerin). With the new recognition of the NeogenePleistocene boundary at the top of the Olduvai event, correlated with the beginning of
the Eburonian cooling, the majority of paleontologists now place within the Neogene
some faunas, like Seneze, which were formerly considered Pleistocene.
In this revision, we have tried to keep the change of localities position to a
minimum and have added several new and well studied faunas.

For the Agenian age, there are no major modifications, only the addition of new
Spanish localities.
For the Orleanian age, the French locality Artenay is placed again in MN 4; that
determination is based on the new record of Democricetodon (Bulot, oral communication). So, as previously argued by Ginsburg, the Proboscidean datum is placed at the
beginning of MN 4 (for western and central Europe). During this interval the biological events are so numerous that it was impossible to note all the changes.
European Neogene Mammal Chronology
Edited by E.H. Lindsay et a/.
Plenum Press, New York, 1990

73

....

-..I

2a

2b

loiN

PAULHAIC

W. Asia
N. Africa

Eucrlcotodon
Paeudocrtcetodon
lolelloolodon
Stonooflbor

Amphiporathorium
Plozoduo
Amphllaguo
Tltanomyo

Amphltraguluo

Wlochberg

Boudry 2

PyrlmontChallomyo

Dromothorlum

Tomordlngon

Calnothorlum

Proto cerathorium

Dlacorathorlum

rornant 11

Agyopo
------- ------------ ---------

Haplocyon
Cophalogolo
Yoongrlnla
Ploolctlo
Amphlctlo
Polaoogalo

lollomoryx
Lophlomoryx
Prodremothorium

Paracerathorium
Aprotodon

Protalactaga

~~~~::;:.

T~!1:l:.rr-

Warculnomya

Hyotherlum

Proatlurua

Amphlcyon
Cyneloo
Palaoogalo

Apoomyo

Tltanomyo

Dlmylechlnuo

Phylotlllon
Orlomoryx

Pooudodryomyo
Horpootldoo
Potamothorlum

Armantomya

RIHonorla

Aurollachooruo

Brolllana
Ploologalo
Parataplruo
Xonohyuo

Prologuo

-----------

lllrobolla
Deperetom yo
Eumyarlon

Other

rlnt Appearance
W+C Europe

Palaeo:scaptor Lagopsls

Agyopolaguo

Other

Common Taxa

W+C Europe

Hyotherlum

RoHonburg
3,5,8

Saulcot

S.E. Europe

Haraml 1
Cynoloe
--- ------ ------------- Amphlcyon

E. Europe

AGENIAN

European Mammal Neogene (MN) Zones.

Wolooonburg 6

Welaenau

La Chaux

hauaen

Schaff-

Heaaler

~~~r.~h

C. Europe

lololooac

IIONTAIGU
Alllao
Gano

loloncalvlllo

Colina

Lambert
Chavrochoo

Marcotn

Cocumant
Valquomado

Ballzac

Solloo-our-Chor

LAUGNAC

Navarrete

Bouzlguoo

lorane a

W. Europe

Table 1.

Elomeryx

Bedenomeryx

Eotaplruo

Protaplruo

Palaeochoerua

Proolluruo

Rhodanomyo

Dromothorlum

Pleuroceroa

Horpostldos
Proallurua

Tltanomys
Plozoduo

------------

-----------

Other

Last Occurrence

W+C Europe

-.1
C1l

loiN

Leo ben
Puttenhauoen

Qta Pombolro
Pontlgno
Chela
Savlgno
t.lunebrega

Rou1cherod

Strakonlco

Elblowald

Folnln
LA ROt.IIEU

NOUYIIIO

Estropouy

Hona du J~:r."n~Y
Unlv. Colo. Llsboa
Leo Bolllaux

Valhondo

Cootablanca

Serre de Verge
Agroda

Chlllours
t.loll Calopa

Moratllla

Arlenay
Qta Narlgao

Bunol
Valtorea

Co reo lea

~~y;~~urg

Chomatov
Echzoll
Bluing on

burg 5

Welaaen-

Schnaltholm
WINTERSHOFWEST

Tuchorlco

Dol nice
Potorobuch

Forothart
Qla Farlnholra
Rombach
Balgnoaux
Erketohofon
t.lorallneo
Aorotroln Orechov

t.lonlroal

Boaugency
Tarazona
lavers
Vloux Collongos

Toffalba Thonay Oggenhof

PONT LEVOY
Qta Pod relzas
franzenbad
Vermeo 1

moosen

Langen-

Maaaendorf

C. Europe

W. Europe

Kln!vtoh

Blshlubo
AIIYIM

Bania Luka

t.lala t.llllva

Luzovlk

Komollnl

Slbnlca

S.E. Europe

Gemorok

t.lurefte

Hoskoy

Chloo

W. Asia

Rotom

AI Sarrar
AI Dabllyeh

Yeroham

H. Africa

Haraml :S

-- .LWo_t.ltcu._

Horlak 1

Wadi
t.loghara

---

Jabal t.lldra

------ ----- ------- ---------Gobel Zollon

Koylbulax

E. Europe

Archaoobolodon

Zygolophodon

Gompholhorlum

Prodolnothorlum

Albanohyuo

~:?:!:~'!~x

~~T~~~~~:~um

Anchlthorlum

Ursovua

~~";~,:~:~a

P11udaeluruo
Horpootldos

Spermophlllnus

Prolagus
Lagopolo

t.llrabello
Eumyarlon
Deperotomyo

Other

Common Taxa

W+C Europe

ORLEAN IAN

Stephanocemaa
Procervulua

Acteocemaa

Lagomeryx

Anchlthorlum
Amphlmoschuo
Xonohyua
Brachyoduo

Ureavua
Pseudarctoa

Llgerlmys

Mtcromeryx
Oorcatherlum
Stephanocemoa

Proaantorhlnua
Larlotothorlum
PROBOSCIDEA
Bunollotrlodon
Eotrogus

~:~T!~C:~:therlum

Megacrlcetodon
Anomalomya
Neocometee
Hyoenallouroa

Ptychoprolaguo
Oemocrlcetodon

Heteroprox

Conohyuo
Sanlthorlum
Dlcroceroa

Eomuacordlnua

Keramldomys
Anchlthorlomys
Crlcetodon
Chollcotherlum
Chllotherlum
Galndatherlum

Plloplthecus

Sanlthorlum

Aurellachoeruo

Bunollslrlodon

Hlspanotherlum

~::~::Y~!~Ium

Anchllherlomyo

Palaeoaclurua

Amphlporalherlum
Llgerlmyo

-------sr::::rlce

Bunollotrio don
Canthumeryx
Doreathorium

Shamallna
Araboamlnthus
Crlcetodon

Antomuo

Orlomeryx

Paralaplruo
Xonohyuo
Andegameryx

~~:~~r.:~~~~hllua

Eucrlcotodon
Paeudocrlcetodon

Amphllaguo

Amphltraguluo

Brachyoduo
Procervulua

Protacerothertum
Olacerotherlum

Pooudolherldomyo
Cynoloo

(continued)

----------

---------

Other

Last Occurrence

W+C Europe

----------- lofolloolodon
Alberlona

Turkomyo

Other

First Appearance
W+C Europe

en

.....

loiN

Slmorro

SAN SAN

Uot

Arroyo dol Val

Wanchonea

Paracuolloo 3,5

Torll

La Grena11ere

Corro dol Otoro

La Grlvo lol

Palencia

La Clotornlga

Solera

Excobooa
St. Gaudono

San Qulrzo
La Grlvo L3

W. Europo

Plackla

Sarlcay

Prbreza

Paoolar

Candlr

Plloplthocuo

Benl lolollal

AI Jadldoh

Dryopllhocua

Crlcelodon

Prolaguo
Lagopala

~~:~~f::~r:~~"

Kubano-

Paradlcerow

Parcrocuto

lollohyaona

turolagua

Slvapllhocuo
Crouzella

Euprox

Protlcllthorlum

roraythla

Zygolophodon

Plalybolodon
Gomphothorlu m Choerolophodon

Capralragoldeo _

Mesaceratherlum

Plltoayon
Plealaceratherlum

Hyaonallouroo
Homlcyon
Olnocyon

Goblcyon
Tungurlctlo
Percrooulo

"''~~i~~

Capratragoldoo
Hypoodonluo

Archcooboladon

Taucanamo

Colnotherlum

Pooudocyon

Gllrudlnuo
Taxodon

Doamanodo Ptychoprolagua
Schizo-

-----------

~TI~~~~r~=

Dakkamya

Prodolnothorlum

----------

Palaeomeryx

Ko:1;nochoerua

Neocomete1

Tolralophodon
Dolnolhorlum

Pro ragocerue

Hlopanomyo

DromacaraBollalovlna
choerue
thorium
Eotrav~
Kubanotragu Aceratherlum
Prodolnothorlum lnlanathorlum Llotrlodon_ Llotrlodon
Palaeomeryx

Hyothorlum
Conohyua
Albonohyuo
Dorcathorlum
Logomoryx
lollcromeryx
Olcrocerue

Larlelothorlum
Ph;ylotlllon
Challcothorlum

Anchllhorlum

Sansano1mllu1

~:~t!~:r~~~=

lochyrlctla

Proia lactaga

lolelaaaylmya

Alloptox

thorium

Coucaso-

Turkomyo

-----------

Hlapanotherlum

----------

Plalybalodon

Zramyo

t.le;apodotoa

Othor

Last Ocourronce
W+C Europo

Pleslosorex
Crlcetulodon Crlcolodon
Doperetamyo

Othor

First Appearance
W+C [uropo

Galorlx
Ooomanolla
Lanthanothorlurr
Cruoafonllna
Konyalagomyt Wuscardlnus
Plotlodlmylua

Othor

Common Taxa

W+C Europe

Afrlcanomya
lufJJ.U'______ lolegacrlcetodon
Domocrlcotodon lologaflodotoa

Palanlak 6

N. Africa

---------- ------------ --------- -----------

Koroth.l
lollkulov

Oppolo

----- ---------- ---------

Kloln
Hadorodorf
Stolnborg
Gclldborg
Schononborg
BloloGerlach
Noudorf
metschav
Sandbo'l,
Houdarf p.

Szontondrol

hausen

Sandolz-

Hasno1

mund

Goorgono-

Ohnlngen

STEIHHEIIol

1~;ro

-au
Lodo

Sofca

Yen I
Eoklhlaa

W. Aala

~~~ 1 Etephan

S.E. Europe

Boyroklepo 3

E. Europe

Klolnoloonbaoh
Pneworno
Vormeo 2
Glv;enhouoon II

C. Europe

ASTARACIAN

Table 1 (continued)

-.J
-.J

10

liN

Amborlou 1

Hammer-

Rudabanya

Charmolllo

AzambuJolra 1
Nombrovllla
JuJurloux
Can llata 1

Eppoloholm

Howonogg

Comanutl 2

Buzhor

Kalfa

Broil a

Atavaoka

Lapuohna

Sobaotopal
lalovena
Hovorany
Vamltoa

Galsolborg
Aumel1ter
Grooolappon

--

Balclk

Eldar

Vooondorf

Gotzendorf

Borlolav

Groooulova

Kalnary

Kohfldloch

Suchoma1ty

E. Europe

C. Europe

achmlodo
Carrllanga
Douo Ia Fantalno llarktl

Lao Valloo do
Fuontlduona

Can Ponslc
St. Joan do B.
Podreyguoros 2C

CAN UOBATERES

VIlla do Caval

Lyon X Rouooo

llontrodan
IIASIA DEL BARBO

La Tarumba
Sob lay

Torraooa

Cucalon

W. Europe

Amama 1

N. Africa

Kalakoy
Corak
Yorlor
Karaln
A Kin
Bayrak-

Karaozu
Koudlct
ot Tluo

Kobaahta

Agha Jarl Ouod Zro

Dendll

W. Aola

Yaoolorom
Blod Douarah

Proochotona

Parapodomuo
Eplmorlonoo
Eozapu1

Palaoomyo

Tragonthorlum

loflapotaurlota

~r:~:::r~~::.n

lollotragocoruo
Totralophadan

~~ra'::traguo

Korynochoeruo
Dorcathorlum
loflcromoryx

Oryctoropuo

Choorolophadon

Chllathorlum

Plio hyrax
lndarctoo
Mochalroduo
Dlcorothlnuo
Hlpparlon

loflcrootonyx

Pllopeteo

Progonomyo

Micro ocr ce ue

Tragoportax

Adcrocuta

Sanoanoomlluo
Poeudaeluruo

Uroavuo

Eurolaguo
llogacrlcotodon
llyogll

htm-

~1no1orex

lofotacordylodon

Amphlcyon
Trocharlon
Llotrlodon
Lagomoryx
Dtcrocero

lofacholroduo :r:!~o\;~".ru
Dlcoroo
Anchlthorlum
Hlpparlon
Gomphothorlum

Samothorium

Plloplthocus
Cryaplthecu1
Zramyo
Sonouoolmyo
Buzhoromyo Domocrlcotodon
Eomuacardlnu
Eumyarlon
SchizoAgnotherlum
galerbc

Progonomyo

~~rca:.~;

brlacoruo
ProotroptiS:.!1!9.!__

(continued)

IProttctllhorlum
Mlomachalroduo
Dlcoryphochoeru

Dakkamyo

----------

Afrlcanomyo

Other

Laot Occurronco
W+C Europe

Schlzochooruo
Bohlin Ia
lollatragaceruo
Honanothorium
Samotraguo Euprox

Adcrocuta

Ouzoceroe

Pooudomerion

Sarmatamyo

Alllopuo
Lophocrlcotu1

Other

Fltot Appearance
W+C Europe

Byzantlnla
=~~~~1::r:ta
Hlopanomyo
Alloopalax
Zramyo
Crlcotulodon
llyocrlcotadon
lluocardlnuo
Spormophlllnu
Protalactaga
Albanonola

Pralaguo

Galorlx

Cruoafontlna

Other

Common Taxa
W+C Europe

lofachalroduo
Ravin do Ia
Thalaoalcllo
Plulc
Lycyaona
_!!p_o.J!_
Kaotollloo-~----lndarctoo
Jobol Somono
Pllavlvorrapo
lofahmutUroavuo
Bou
kay
Hanlfla5
Challcothorlum
Taplruo
Elmo
Bou
Acorathorlum
Akcakoy Hanlfla 1

Ravin doo
Zouavoo(RZ1)
Bladrak

Lofkon

S.E. Europe

VALLES IAN

-..!

(X)

11

12

13

MH

CREVILLENTE 3
Dlonoy

PI ora

Lobrlou
Amberlou 3
llollon
La Cantero
St. Bouzlle

Casa del Acero

(Cerro do Ia Garita

Con cud

Wr. Leberon

Cucuron Slade

LOS MANSUETOS

Loa Al)ozareo

EL ARQUILLO
La Por1ora
LIIIIOU
lotollna do Sequra

Vonla dol lotoro

Crevlllenlo 6

Ubrllla

Salobrona

Arenaa del Roy

W. Europe

Mamay

Angustovka

E. Europe

Prochoma
Kayacllbl
Vathylakkos
Ravin dos
Zouavoa(RZ5

Cekrace

Garkln

I.Moravha

Grobenlkl

Amama 2

Sldl Solem

------------Ifoli"iir.lF --------Samoa 1,4

gozl

Klnlk

Bacclnollo VC Poraholy
Raopopeny

Coakvar

Wenacer

Sahabl
Wadi Hatrun
Khondok
ol Oualda

H. Africa

Anouroaorex

Amblycoptua
Blarlnolla

Galorlx
Schlzogalerlx
Diboll a
Doamanolla

Mooopllhocuo

Delnothorlum
Zygolophodon

lllcrostonyx
Gazella
Tragoportax

Dlcerorhlnus

Hlpparlon
Challcolhorlum

Adorocuta

~:r..~~~=~ropo

~f:;:,~g=~~~

Atlantoxerua

Ellomya

Eozapua

"Valorymyo"

Occltanomyl

Paropodemuo

Kowalakla

Ruaclnomya

Prolagua

Orycleropue
Stogototrabolodon
Choorolophodan

Dlceros

Paramocholrodu

Pooudomerton

Other

Common Taxa

W+C Europe

------- ----- Hlopanomyo

Taaklnpaoa

W. Asia

Sumeg
DornDurkhelm

Elloavotouk

Novo

Taraklla

Clmlslla

Cobrucl

Salonlkl

Choma tori

Samoa 5
Plkorml

Dlllko
Monaolorl
Ano Wotochl

S.E. Europe

Corak
Yelr
Kucuc

Elchkogol

Tlhana

Bucclnollo VI

Tardoabanya

Tudorovo

Bacclnollo V F'ranzovka
Hatvon
Polgardl

Caolno
Ballavar

Brlolgholla

C. Europe

TUROLIAN

Table 1 {continued)

Wacaca

Holladothorlum

Protoryx

Toplrlscuo
Cervovltua
Psoudotraguo

Proapola~

"Valorymyo"
Dlpoldto

~~~r.:':amy

Anacue

Gazella

Hlopanodorcoa

Stephanomyo

Ruactnomya

Amblycoptuo

Paracamelua

Dlnofolla
Agrlolherlum

:C~aamapor1hotoa
. .~:~.r::r:.

Hyolrlx
Apodemua
Paraolhomya
Crlootua

~:.::~:::::.~~~

Crlothorlum
Ploaladdax
Palaeorea1
Hlsldorcaa

-----

pf.~~~~UI

Selene-

Protatoro

lhroudla

~r:~'~y.

Rodunca
Paraoamolu

lachymomya

acopt11

lollcrolodon
Analolomya
Mlcroto-

Other

rtrst Appearance
W+C Europe

Ooconnalhorlum

Anomalomyo
Alloopalax

Lanthanothorlum

Mlcromoryx

Percrocuta

Hlapanomyo

Dlnoaorex
Ploalodlmyluo

Pllocorvuo
Hollodothor1um
Dolnolhorlum
Tetralophodon

Cervavltua

Acoralhorlum

Adcrocuta
Thala01lctls

--

Hlsldorcaa

Samothorlum
Plosladdox

Slvapllhocua

CrlcetuIoden

Zromyo
Byzantlnla

Mlcroatonyx
Palaooryx
Palaeoreae
Choorolphodon

Wyocrlootodan

Other

Lao! Occurrence
W+C Europe

(I)

....

14

15

liN

Solo

Torrato

Hautlmaen

Wantpolllor

Collonouvo
PoraloJoiJ'
Hautorlvo

Caravaca
Alcoy

Vondargu..
La Gloria 4

Colada I

Orrloo 1

Bacclnollo
VI

Ootramoo
1,11

Kaoolaklno

PODLESICE

Dorkovo

Frunzovka

Cugonouka

Antlpavka

Kamonokoo

La Calera

Ootramoo 1

Plolomolo I

Spllla 2,1

S.E. Europe

Calla

W. Asia

lchkoul

Aln Brlmba

N. Africa

~h=~:::,o:,~mus

Stophanomyo
Caotlllomp
Paraothomyo
Occltanomp

Ruoalnomp

Oryctolagus
Hypolaguo

~~':.\\~:Iague

Blarlnokloo
Diboll a

Macaca

plthoCUI

~=~:~~~OCUI

Plolomalo 1

Kardla

Warltoa

Dinar
Aklaky

lgdoll

AIn
Cuottura

Ouod
Athmonla

Anaj'ICUI

Zygalophadon

Para boo
Gazella

Crolz:etoceroa

Suo

Hlpparlon
Dlcororhlnuo
Taplruo

Nycloroutoo
Canto
Chaomaporthole
Pochycrocuta
Ursuo

Hyotrlx

Slvathorlum

Arctomolos
Baranogalo

lhrondla
Eulmuo
Plloopalax

merfone1

Poeudo-

Mooocrlcotuo
EplmotJonoo

~::rz~~~~~~.

Condylura

Other

Common Taxa

W+C Europe

- - - Cao
------- ---- Argoub
Prror,otaurlota
or

Odoooa
(Catacombo)
Borootl,
Maluotonl
Cluporcon

Kagul

Wozo

E. Europe

Kutchurgan

lvanovoe

Coamota 2

Wolforoholm

C. Europe

VIllalba Rio
Go dolo
Wont Helene

La Gloria 2

Porplgnan
(Serra! don
Vaoquor)
La Juliana

C.'J1:~~a A~ta

Orrlon 1,7

Gorafo 2

Layna

W. Europe

RUSCINIAN

Paracervulue

Vormola
Moloo

~~r,,o,!,;~uo
Pachycrocuta

Suo

Poracenulua
Orycloropuo

Ploologulo
Pllovlvorropo

Spermaphlllnuo
Promlmomyo
Coladonola

Paranou.:-- "Protatera"
roeorex

Wlcroapalax
Pllaoolovlnla

Aratomyo

Lemmue

Kowalokla
Occltanomyo
Vlllanyla
Paraothomyo
Borsodla
Coorla
Synaptamyo Dlnofollo

Weaocrlcetua
Eolramomy
Proopalax

Eploarlculuo
Promlmomyo
Trllophomyo

af:::~ne.:l~:.

Dallcho-

Dolomyo
Crlcotuluo
Scluruo
Paralluruo

Mlmomyo

Trlochlzolaguo

Ochotonoldoo

Boremondla
Oryctolaguo
Pllopentalaguo

(continued)

Stegotetrabolodon

Lophocrlcetue
Microtoocoplo

Pr"OiiCiiiiiOnaAmblycapluo

Other

Last Occu.,..nco
W+C Europe
Mooopllhocuo
Galorlx

Other

nrst Appearance
W+C Europe

()')

16

17

MN

TRIVERSA

ARONDELU

Eocorlhuela

Arclllo

Uryv 1

Hajnacka

Balaruc 2

Caponi

Kvabobl

Roblellce

Tuluceotl

W. Asia
N. Africa

Limn I

Tourkobounla
Gulyazl

Amama 3

Bulla Rogla

Orouaae

Dhobol

------------ 1------- -------

S.E. Europe

------ ----------Cornateotl

Clupercenl

Slallna 2

Llventoovka

Khapry

-------Beremend 5

Oolramoo7

Montopoll

Dragoneoll

Oolramoo 3

Chorlootll

Kadzielna

--~-----

Botfla XIII

E. Europe

Tegeler

Brielle

C. Europe

VIa Ieite

r--------------

Woreda

El Rincon
Seyne1

Etoualrea

Valdeganga 1.11

Pardlnoo
Vlllaroyo
Roccaneyra

ST. VALLIER

Puobla do Valverde

Seneze
Chlllac
Lo Coupe!

t.tontou 5

W. Europe

Ana neue

Suo
Gazellooplra
Pllotraguo
Crolzetoceroa

Homotherlum
Dlcerorhlnuo
Taplruo

Meganthereon

Aclnonyx

Nycteroutoo
Canlo

Pachycrocuta

Caotlllomyo
~r;,o.,:~therlum
Hyotrlx

Pllomyo

~::;:~r:

Prolaguo
Oryctolagus
Hypolaguo

Beremendla

Blarlnoldos

De1mana

acaca

Paradollchopllhocuo
Dollchor/clthocuo

Paracamelu1

Other

Common Taxa
W+C Europe

"VI LLAr\YIAN"

Table 1 (continued)

Kr.lll:::~~

Leptoboo
Pllolraguo
Gazellosplra

Aclnonyx

Homolherlum

Meganthereon

Coorla

Paradollcho-

Mammuthu

Equuo

Vlrotalluruo

Heaperldocerua

Eucladocoroo

Vulpeo
Lynx

Lemmua

Lagurodon
Clothrlonomyo

Llbralcos
lotegalovlo

Hippopotamus

Allophalomys

-----------

---------

Other

First Appearance
W+C Europe

Procamptocerua

Agrlotherlum

Dlnofollo

Trllophomyo
Bloncomyo

Zygolophodon

Gazella

Hlpparlon

Chaomopor!hetoo

Parallurua

Stophanomyo

Anancua

Pllolraguo
Gallogoral

-----------

----------

Other

Last Occurrence

W+C Europe

Table 2..

Principal reference localities and fauna.

MNl
F- 47 Paulhiac
Marsupialia

Amphiperatherium frequens

lnsectivora

Amphechinus arvernensis; Paratalpa micheli; Clapasorex

bonisi

Chiroptera

Nyctinomus stehlini; Myotis sp.

Lagomorpha

Piezodus tomerdingensis; Titanomys visenoviensis

Rodentia

Eucricetodon collatus; Peridyromys murinus; Bransatoglis

concavidens; Bransatoglis fugax; Glirudinus glirulus;
Heteroxerus paulhiacensis; Heteroxerus lavocati; Rhodanomys
schlosseri; Pseudotheridomys parvulus; Plesiosminthus cf.
myarion

Carnivora

Cephalogale ginesticus; Cephalogale gracile ursinus;

Haplocyon elegans; Haplocyon crucians; Haplocyonopsis
crassidens; Amphicyon cf. astrei; Ysengrinia tolosana;
Amphicyonine sp. 1; Amphicyonine sp. 2.; Plesictis solidus;
Plesictis palustris; Plesictis' cultellatus; Plesictis sp. 1;
Plesictis sp. 2.; Proailurus lemanensis

Perissodactyla

Paleotapirus cf. d.ouvillei; Brachypotherium lemanense;

Aceratherium paulhiacensis; Diceratherium pleuroceros;
Dicerorhinus cf. minutus

Artiodactyla

Hyotherium major; Cainotherium geoffroyi; Dremotherium

feignouxi; Amphitragulus major; Amphitragulus cf. lemanensis
MNZa

F - 03 Montaigu
Marsupialia

Amphiperatherium frequens

Chiroptera

Hydromops stehlini; Palaeonycteris robustus; Rhinolophus

lemanensis

Insectivora

Amphechinus edwardsi; Dimylechinus bernouillii; Dimylus

paradoxus; Oligosorex antiqrms; Soricines 1, 2.

Lagomorpha

Titanomys visenoviensis; Piezodus cf. branssatensis;

Amphilagus cf. ulmensis

Rodentia

Eucricetodon gerandianus; Rhodanomys schlosseri;

Pseudotheridomys parvulus; Plesiosminthus myarion;
Pseudodryomys aljaphi; Heteroxerus paulhiacensis;
Palaeosciurus feignouxi; Steneofiber eseri; Peridyromys
murinus; Vasseuromys priscus; Vasseuromys rugosus;
Microdyromys sp.

Perissodactyla

Paleotapirus douvillei; Diaceratherium pleuroceros;

Protaceratherium tagicum; Mesaceratherium paulhiacensis;
Diceratherium aginense; Moropus sp. or Phylotillon sp.

Carnivora

Pseudocyon crassidens; Cynelos rugosidens; Cynelos

lemanensis; Haplocyon elegans; Haplocyon crucians;
Pseudocyonopsis gerandianus; Potamotherium valetoni;
Cephalogale depereti; Cephalogale gracile; Stenogale julieni;
Stenogale brevidens; Proailurus lemanensis; Plesictis
lemanensis; Plesictis? croizeti; Plesictis julieni; Plesictis
stenoplesictoides; Plesictis robustus?; Plesiogale angustifrons;
Palaeogale minuta; Paragale hurzeleri; Amphictis lemanensis;
Amphictis antiquus; Herpestides antiquus
(cont.)
81

Table Z (continued)
Artiodactyla

Hyotherium major; Palaeochoerus tyPus; Dremotherium

feignouxi; Amphitragulus elegans; Amphitragulus lemanensis;
Oriomeryx major; Amphitragulus boulangeri; Amphitragulus
gracilis; Cainotherium commune

Pholidota

Necromanis sp.
MNZb

F - 4 7 Laugnac
Marsupialia

Amphiperatherium freguens

Insectivora

Proscapanus primaevus; Carposorex sylviae

Chiroptera

Myotis insignis

Lagomorpha

Prolagus cf. vasconiensis

Rodentia

Eucricetodon aquitanicus; Melissiodon sp.; Peridyromys

occitanus; Peridyromys brailloni; Vasseuromys rugosus;
Glirudinus cf. modestus; Steneofiber eseri; Pseudotheridomys
parvulus

Carnivora

Haplocyon elegans; Haplocyonoides mordax; Cynelos

rugosidens; Cynelos lemanensis; Amphicyon cf. astrei;
Ysengrinia sp.; Plesictis aff. solidus; Plesictis laugnacensis;
Palaeogale minuta; Plesiogale angustifrons; Amphictis
aginensis; Herpestides collectus; Proailurus lemanensis

Perissodactyla

Palaeotapirus cf. douvillei; Diaceratherium aginense;

Plesiaceratherium platyodon; Diceratherium cf. pleuroceros;
Diceratherium sp.; Protaceratherium minutum

Artiodactyla

Xenohyus venitor; Hyotherium major; Cainotherium geoffroyi;

Cainotherium commune; Dremotherium feignouxi;
Amphitragulus gracilis; Oriomeryx cf. major; Andegameryx
andegaviensis
MN3

D- Wintershof-West

82

Marsupialia

Amphiperatherium freguens wintershofense

Insectivora

Heterosorex neumayrianus subseguens; "Sorex" pusilliformis;

"Sorex" stehlini; Soricella discrepans; Paenelimnoecus
micromorphus; Cordylodon intercedens; Plesiodimylus
hUrzeleri; Talpa sp.

Chiroptera

Rhinolophus aff. lemanensis; Rhinolophus sp.; Megaderma sp.

Lagomorpha

Prolagus vasconiensis; Amphilagus ulmensis

Rodentia

Ameniscomys selenoides; Plesispermophilus decedens;

Paracitellus eminens; Heteroxerus wintershofensis;
Palaeosciurus fissurae; "Ratufa" obtusidens; Miopetaurista
dehmi; Pseudocricetodon sp. ?; Melissiodon dominans;
Pseudodryomys ibericus; Pseudodryomys gregarius; Glirudinus
modestus; Glirudinus gracilis; Heteromyoxus schlosseri;
Bransatoglis spectabilis; Miodyromys biradiculatus;
Peridyromys murinus?; Ligerimys lophidens; Ligerimys
antiquus; Apeomys turkheimae

Carnivora

Cynelos rugosidens schlosseri; Cynelos lemanensis helbingi;

Amphicyon steinheimensis bohemicus; Megamphicyon
giganteus; Pseudocyon sansaniensis aff. intermedius;

Pseudarctos bavaricus dehmi; Phoberocyon huerzeleri;

Hemicyon sp.; Hemicyon dehmi; Ursavus elmensis; Plesictus
aff. pygmaeus; Plesictis aff. sicaulensis; Plesictis mayri;
Plesictis vireti; Plesictis humilidens; Amphictis aff. antiquus;
Martes laevidens; Laphyctis vorax; Laphyctis comitans;
Palaeogale minuta; Palaeogale hyaenoides; Plesiogale
postfelina; Broiliana nobilis; Stromeriella franconica;
Miomephitis pilgrimi; Semigenetta elegans; Progenetta
praecurrens; Pseudaelurus transitorius
Perissodactyla

Anchitherium aurelianense; Tapirus aff. intermedius;

Rhinocerotidae indet.; Mesaceratherium aff. simorrense;
Protaceratherium minutum; Diaceratherium aurelianense

Artiodactyla

Aureliachoerus aurelianensis; Cainotherium bavaricum;

Procervulus dichotomus praelucinus; Amphitragulus cf.
boulangeri; Palaeomeryx aff. kaupij Andegameryx serum;
Lagomeryx parvulus

Pholidota

N ecromanis franconica; N ecromanis parva

MN4

F - 33 La Romieu
Insectivora

Galerix cf. exilis; Talpa sp.; Proscapanus sp.; Heterosoricinae;

Miosorex grivensis; Soricidae indet.

Chiroptera

Hipposideros collongensis; Myotis sp.

Lagomorpha

Prolagus aff. oeningensis; Lagopsis penai

Rodentia

Megacricetodon cf. collongensis; Megacricetodon bezianensis;

Democricetodon mutilus; Democricetodon aff. romieviensis;
Democricetodon sp. 1; Democricetodon sp. 2.; Eumyarion cf.
weinfurteri; Melissiodon cf. dominans; Ligerimys antiquus;
Ligerimys florancei; Peridyromys aquatilis; Peridyromys
occitanus; Peridyromys gregarius; Miodyromys biradiculatus;
Pseudodryomys ibericus; Glirudinus modestus; Microdyromys
koenigswaldij Microdyromys sp.; Bransatoglis cadeoti;
Miopetaurista sp.; Heteroxerus rubricati; Spermophilinus
bessana; Palaeosciurus cf. fissurae; Steneofiber depereti
carnutense

Fissipeda

Megamphicyon giganteus; Pseudocyon sansaniensis;

Semigenetta repelini; Pseudaelurus quadridentatus

Perissodactyla

Anchitherium aurelianense; Brachypotherium brachypus;

Plesiaceratherium lumiarense; Lartetotherium sansaniensis;
Prosantorhinus germanicus

Artiodactyla

Aureliachoerus aurelianensis; Bunolistriodon lockharti;

Hyotherium soemmeringi; Cainotherium miocaenicum;
Dorcatherium cf. naui; Procervulus dichotomus; Lagomeryx
d. rutimeyeri; Palaeomeryx kaupi

Proboscidea

Gomphotherium angustidens; Deinotherium cuvieri

MNS

F- 36 Faluns Pont Levoy- Thenay

Primates

Pliopithecus piveteaui

Insectivora

Plesiodimylus sp.; Lanthanotherium sp.; Soricidae

Lagomorpha

Prolagus oeningensis; Amphilagus ulmensis

(continued)

83

Table Z (continued)
Rodentia

Spermophilinus besana; Miopetaurista cf. lappi; Miodyromys

cf. aegercii; Steneofiber depereti; Cricetodon aureus;
Megacricetodon collongensis; Megacricetodon lappi;
Democricetodon mutilus; Eumyarion weinfurteri

Perissodactyla

Chalicotherium grande

Artiodactyla

Procervulus dichotomus; Dicroceros elegans parviceros

MN6

F- 33 Sansan
Primates

Pliopithecus antiquus, Crouzelia auscitanensis

Insectivora

Dinosorex sansaniensis; Paenelimnoecus crouzeli; Miosorex

desnoyersianus; Oligosorex prevostianus; Blarinella robustus;
Talpa minuta; Alloscapanus sansaniensis; Proscapanus sp.;
Mygalea antiqua; Lanthanotherium sansaniensis;
Lanthanotherium tobieni; Pseudogalerix exilis; Galerix sudrae;
Amphechinus ginsburgi; Plesiodimylus chantrei

Chiroptera

Myotis murincftdes; Myotis elegans; Eptesicus noctuloiiies;

Eptesicus campanensis; ?Miomegaderma gaillardi

Lagomorpha

Prolagus oeningensis

Rodentia

Miopetaurista sansaniensis; Blackia parvula; Spermophilinus

bredai; Heteroxerus cf. grivensis; "Sciurusn sp.; Cricetodon
sansaniensis; Democricetodon gaillardi; Democricetodon
crassus; Megacricetodon minor; Megacricetodon gersi;
Eumyarion medium; Eumyarion sp.; Eomuscardinus
sansaniensis; Myoglis meini; Paraglis astaracensis;
Microdyromys miocaenicus; Miodyromys aegercii;
Keramidomys octaviae; Steneofiber sansaniensis;
Trogontherium minutum

Carnivora

Amphicyon major; Pseudocyon sansaniensis; Pseudarctos

bavaricus; Alopecocyon goeriachensis; Hemicyon sansaniensis;
Plithocyon armagnacensis; Schlossericyon viverroldes; Martes
sansaniensis; Martes muncki; Ischyrictis zibethciides;
Proputorius sansaniensis; Taxodon sansaniensis; Mionictis
dubia; Semigenetta sansaniensis; Viverra medica;
Stenoplesictis aurelianensis; Pseudaelurus quadridentatus;
Pseudaelurus turnauensis; Sansanosmilus palmidens

Perissodactyla

Anchitherium aurelianense; Chalicotherium grande;

Aceratherium tetradactylum; Brachypotherium brachypus;
Lartetotherium sansaniensis

Proboscidea

Gomphotherium angustidens; Deinotherium laevius

Artiodactyla

Listriodon splendens; Taucanamo sansaniense; Dorcatherium

crassum; Dicroceros elegans; Heteroprox larteti; Micromeryx
flourensianus; ?Strogulognathus sansaniensis; Palaeomeryx
magnus; Eotragus sansaniensis
MN7

D - Steinheim

84

Insectivora

Mioechinus intermedius; Galerix socialis; Heterosoricinae;

Soricinae; ?Proscapanus sansaniensis

Chiroptera

Vespertilionidae; Eptesicus campanensis; Chiroptera indet.

Lagomorpha

Prolagus oeningensis; Lagopsis verus; Eurolagus fontannesi

Rodentia

Spermophilinus bredai; Microdyromys complicatus

miocaenicus; Miodyromys aegercii; Myoglis meini;
Megacricetodon minor; Megacricetodon germanicus;
Democricetodon mutilus; Democricetodon gaillardi;
Eumyarion cf. latior; Anomalomys gaudryi

Carnivora

Paralutra jaegeri; lschyrictis mustelinus; Martes cf. filholi;

Proputorius sp.; Trochotherium cyamoides; Trocharion
albanense; Amphicyon steinheimensis; Amphicyon sp.;
Amphicyonopsis? serus; Hemicyon gtiriachensis; Ursavus cf.
intermedius; Pseudaelurus quadridentatus; Pseudaelurus
lorteti; Sansanosmilus jourdani; Semigenetta sansaniensis

Perissodactyla

Anchitherium aurelianense; Aceratherium simorrense;

Lartetotherium sansaniense; Lartetotherium steinheimense;
Phyllotillon fraasi

Proboscidea

Gomphotherium angustidens

Artiodactyla

Listriodon splendens; Conohyus simorensis; Albanohyus

pygmaeus; Micromeryx flourensianus; Euprox furcatus;
Heteroprox larteti; Palaeomeryx eminens; Dorcatherium
crassum
MN8

CH- Anwil
lnsectivora

Galerix socialis; Lanthanotherium sansaniense; Plesiosorex

schaffneri; Dinosorex pachygnathus; Plesiodimylus chantrei;
Metacordylodon schlosseri; Alloscapanus sansaniensis;
Scaptonyx edwardsi; Talpa minuta; Desmanella stehlini

Chiroptera

Scotophilus sp.; Tadarida sp.; Nyctinomus helveticus;

Miomegaderma gaillardi

Lagomorpha

Prolagus oeningensis; Lagopsis verus; Eurolagus fontannesi

Rodentia

Spermophillinus bredai; Albanensia albanensis; Miopetaurista

gaillardi; Forsythia gaudryi; Blackia miocaenica; Sciuropterus
sp.; Deperetomys hagni; Cricetodon sp.; Megacricetodon
schaubi; Megacricetodon similis; Megacricetodon germanicus;
Democricetodon brevis; Democricetodon freisingensis;
Eumyarion latior; Anomalomys gaudryi; Neocometes brunonis;
Microdyromys koenigswaldi; Microdyromys complicatus;
Paraglirulus werenfelsi; Paraglirulus conjunctus; Glirudinus
undosus; Eomuscardinus aff. sansaniensis; Muscardinus sp.;
Miodyromys aegercii; Miodyromys hamadryas; Paraglis sp.;
Myoglis meini; Keramidomys mohleri; Keramidomys
anwilensis; Eomyops catalaunicus; Trogontherium minutum

Carnivora

Trochotherium cyamoides; Leptoplesictis filholi; Semigenetta

sansaniensis; Ursavus brevirhinus; Pseudarctos bavaricus;
Carnivora indet.

Perissodactyla

Anchitherium aurelianense; Rhinocerotidae indet.

Artiodactyla

Listriodon splendens; Conohyus simorrensis; Suidae indet.;

Albanohyus pygmaeus; Dorcatherium sp.; Micromeryx
flourensianus; Euprox furcatus; Cervidae indet.; Palaeomeryx
sp.; Eotragus sp.
(continued)

85

Table Z (continued)
MN9

E - Can Llobateres
Primates

Rahonapithecus sabadellensis; Dryopithecus brancoi;

Sivapithecus sp.?

lnsectivora

Plesiodimylus chantrei; Metacordylodon sp.; Postpalerinaceus

vireti; Parasorex socialis; Lanthanotherium sanmigueli; Talpa
minuta; Talpa vallesensis; Dinosorex aff. sansaniensis;
Miosorex grivensis; Crusafontina excultus

Chiroptera

Chiroptera sp. 1; sp. Z

Lagomorpha

Lagopsis cf. verus; Amphilagus fontannesi; Prolagus crusafonti

Rodentia

Progonomys cathalai; Hispanomys thaleri; Megacricetodon cf.

minor; Eumyarion leemanni; Cricetulodon sabadellensis;
Anomalomys cf. gaillardi; Eomyops catalaunicus;
Keramidomys pertesunatoi; Heteroxerus grivensis;
Heteroxerus cf. rubricati; Spermophilinus bredai; Albanensia
grimmi; Miopetaurista crusafonti; Eomuscardinus vallesiensis;
Muscardinus crusafonti; Paraglirulus werenfelsi; Paraglirulus
conjunctus; Glirudinus undosus; Myoglis meini; Steneofiber
jaegeri; Steneofiber depereti; Trogontherium minutum

Carnivora

Indarctos vireti; Ursavus primaevus; Ursavus brevirhinus;

Protursus simpsoni; Amphicyon major; Simamphicyon
batalleri; Canidae indet.; Martes mellibulla; Martes sp.;
Ischyrictis petteri; Sabadellictis crusafonti; Circamustela
dechaseauxi; Marcetia santigae; Mesomephitis medius;
Promephitis pristinidens; Trocharion albanense; Paralutra sp.;
Trochictis narcisoi; Semigenetta ripolli; Protictitherium
crassum; Protictitherium gaillardi; Hyaenidae indet.;
Machairodus aphanistus

Perissodactyla

Hipparion catalaunicum; Chalicotherium grande; Tapirus

priscus; Dicerorhinus sansaniensis; Dicerorhinus
schleiermacheri; Aceratherium incisivum; Alicornops
simorrense

Proboscidea

Prodeinotherium laevius; Deinotherium giganteum;

Tetralophodon longirostris

Hyracoides

Pliohyrax sp.

Artiodactyla

Albanohyus pygmaeus; Parachleuastochoerus crusafonti;

Listriodon splendens; Hyotherium soemmeringi; Micromeryx
flourensianus; Euprox dicranocerus; Dorcatherium sp.;
Capreolus sp.; Miotragocerus aff. panoniae
MN 10

E - Masia del Barbo (Teruel)

86

Insectivora

Galerix sp.; Postpalerinaceus vireti; Crusafontina excultus

Lagomorpha

Prolagus crusafonti

Rodentia

Hispanomys peralensis; Hispanomys sp.; Democricetodon aff.

sulcatus; Kowalskia aff. fahlbuschi; Progonomys hispanicus;
Progonomys calathai; Muscardinus crusafonti; Tempestia
hartenbergeri; Eliomys sp.; Spermophilinus aff. bredai;
Heteroxerus; Atlantoxerus adroveri; Palaeomys castoro1des;
Dipoides problematicus

Carnivora

Adcrocuta eximia

Perissodactyla

Hipparion cf. primigeniurn; Aceratherium incisivum

Artiodactyla

Decennatherium pachecoi
MN 11

E - Crevillente Z
Insectivora

Galerix sp.; Dibolia sp.

Lagomorpha

Prolagus crusafonti

Rodentia

Kowalskia fahlbuschi; Parapodemus lugdunensis; Occitanomys

sondaari; Valerymys vireti; Eliomys truci; Muscardinus sp.;
Heteroxerus aff. grivensis

Carnivora

Indarctos atticus; Mustelidae indet.; Plioviverrops guerini;

Amphimachairodus giganteus; Paramachairodus ~

Perissodactyla

Hipparion sp.; Dicerorhinus schleiermacheri

Proboscidea

Tetralophodon longirostris

Artiodactyla

Microstonyx major; Dorcatherium naui; Eostylocerus aff.

pierensis; Cervidae indet.; Birgerbohlinia schaubi; Giraffinae
indet.; Tragoportax gaudryi

MNIZ
Los Mansuetos (E - Teruel)
Jnsectivora

Galerix sp.; Dibolia major; Desmanella crusafonti; Talpa

minuta; Blarinella dubia; Paenelimnoecus repenningi

Chiroptera

Megaderma vireti

Lagomorpha

Prolagus crusafonti; Alilepus turolensis

Rodentia

Ruscinomys schaubi; Parapodemus barbarae; Occitanomys

adroveri; Valerymys turoliensis; Spermophilinus turolensis;
Atlantoxerus adroveri; Eliomys cf. truci; Dipo1Cles
problematicus

Carnivora

Canis cipio; Enhydriodon lluecai; Baranogale adroveri;

Adcrocuta eximia; Thalassictis adroveri; Plioviverrops guerini;
Lycyaena; Paramachairodus giganteus; Metailurus parvulus
Hipparion concudense aguirrei; Aceratherium incisivum

Perissodactyla
Artiodactyla

Euprox dicranocerus; Gazella deperdita; Tragocerus

amalthaeus; Hispanodorcas

Proboscidea

Tetralophodon longirostris
UN 13

El Arquilla 1 (E- Teruel)

Insectivora

Galerix sp.; Desmanella crusafonti; Blarinella dubia

Lagomorpha

Prolagus crusafonti; Alilepus turolensis

Rodentia

Ruscinomys schaubi; Cricetus cf. kormosi; Valerymys

turoliensis; Stephanomys ramblensis; Occitanomys adroveri;
Apodemus gudrunae; ?Myomimus sp.

Carnivora

Sivaonyx lehmani; Adcrocuta eximia; Thalassictis adroveri;

Lycyaena sp.; Machairodus giganteus; Metailurus parvulus;
Fortunictis acerensis

87

Table Z (continued)
Perissodactyla

Hipparion periafricanum; Hipparion primigenium truyolsi;

Hipparion gromovae gromovae; Dicerorhimus schleiermacheri

Proboscidea

Zygolophodon turicensis

Artiodactyla

Hexaprotodon crusafonti; Euprox sp.; Gazella deperdita;

Tragoportax sp.
MN 14

P - Podlesice
Insectivora

Talpa minor; Desmana nehringi; Sorex alpinoides; Sorex

dehneli; Sorex runtonensis; Sorex minutus; Petenyia hungarica;
Paranourosorex ~ Blarinoides mariae; Petenyiella gracilis;
Soriculus kubinyii

Chiroptera

Rhinolophus delphinensis; Rhinolophus grivensis; Miniopterus

schreibersi; Plecotus crassidens; Myotis podlesicensis; Myotis
danutae; Myotis dasycneme subtilis; Myotis cf. aemulus;
Myotis cf. exilis

Carnivora

Baranogale helbingi; Vormela cf. petenyi; Mustela sp.

Rodentia

Sciurus cf. warthae; Tamias cf. orlovi; Pliopetaurista cf.

dehneli; Blackia polonica; Pliopetes hungaricus; Pliopetaurista
sp.; cf. Sciurotamias; Leptodontomys aff. catalaunicus;
Keramidomys mohleri; Estramomys sp.; Protozapus
intermedius; Anomalomys sp.; Parapodemus coronensis;
Kowalskia magna; Kowalsia polonica; Cricetus sp. 1; Cricetus
sp. Z; Epimeriones progressus; Baranomys kowalskii; Prosomys
insuliferus; "Trilophomys" canterranensis; Glirulus pusillus;
Muscardinus pliocaenicus; Muscadinus cf. dacicus; Glis minor

Lagomorpha

Ochotonid indet.; Leporid indet.

MN 15

F- 66 Perpignan (Serrat d'en Vacquer)

88

Primates

Mesopithecus monspessulanus; Dolichopithecus ruscinensis

Insectivora

Galerix depereti, Talpa sp.; Episoriculus gibberodon

Lagomorpha

Prolagus depereti; Oryctolagus laynensis

Rodentia

Mimomys davakosi-occitanus; Ruscinomys europaeus;

Trilophomys pyrenaicus; Trilophomys schaubi; Blancomys
neglectus; Cricetus angustidens; Stephanomys donnezani;
Castillomys crusafonti; Occitanomys montheleni; Paraethomys
meini; Paraethomys jaegeri; Rhagapodemus hautimagnensis;
Apodemus dominans; Apodemus jeanteti; Glis minor;
Muscardinus pliocaenicus; Eliomys intermedius; Pliopetaurista
pliocaenica; Castor praefiber; Hystrix primigenia

Carnivora

Dinofelis diastemata; Felis issiodorensis; Felis aff.

maniculata; Viverra pepraxi; Pachycrocuta pyrenaica;
Nyctereutes donnezani; Canis michauxi; Canis adoxus; Ursus
ruscinensis

Perissodactyla

Hipparion crassum; Dicerorhinus megarhinus; Dicerorhinus

miguelcrusafonti; Tapirus arvernensis minor

Proboscidea

Anancus arvernensis; Zygolophodon borsoni

Artiodactyla

Korynochoerus provincialis; Sus arvernensis minor;

Croizetoceros pyrenaicus; Paracervulus australis; Gazella aff.
borbonica; Alephis ~

Tubulidentata

Orycteropus deperetl

Sirenia

Felsinotherium

MN 16
I- Villafranca d'Asti (Triversa)
Primates

Macaca florentina

Rodentia

Castor sp.

Carnivora

Viverra cf. pepratxi; Acinonyx pardinensis; Chasmaporthetes

lunensis

Perissodactyla

Dicerorhinus jeanvireti; Tapirus arvernensis

Proboscidea

Anancus arvernensis; Zygolophodon borsoni

Artiodactyla

Sus arvernensis minor; Cervus cf. cusanus; Leptobos

stenometopon

MN 16
I- Villafranca d'Asti (Arondelli)
Insectivora

Blarinotdes mariae; Episoriculus gibberodon; Sorex sp.;

Petenyia hungarica; Beremendia fissidens; Talpa cf. minor

Lagomorpha

Prolagus savagei; Hypolagus sp.

Rodentia

Mimomys polonicus; Mimomys (Cseria) gracilis; Apodemus cf.

alsomyoides; Muscardinus; Glirulus pusillus

Carnivora

Parailurus cf. hungaricus; Baranogale helbingi

MN17
F- 26 Saint Vallier
Primates

Macaca cf. florentina

Lagomorpha

Ochotonid indet.; Oryctolagus lacosti

Rodentia

Mimomys pliocaenicus; Hystrix refossa; Castor plicidens

Carnivora

Nyctereutes megamastoides; Vulpes alopecoides; Felis (Lynx)

issiodorensis; Panthera schaubi; Acinonys pardinensis;
Megantereon megantereon; Homotherium crenatidens;
Pachycrocuta perrieri; Chasmaporthetes lunensis; Enhydrictis
ardea; Baranogale helbingi; Aonyx bravardij Meles thorali;
Ursus etruscus

Perissodatyla

Equus stenonis vireti; Dicerorhinus etruscus

Proboscidea

Anancus arvernensis; Mammuthus meridionalis

Artiodactyla

Croizetoceros ramosus medius; Cervus philisi valliensis;

Euctenoceros senezensis; Leptobos stenometopon; Gallogoral
meneghini; Gazella borbonica; Gazellospira torticornis

89

The Astaracian is an age with less generic changes, There are difficulties to
delimit the MN 7 unit with generic events; these difficulties occur partially with the
classical placement of the Manchones fauna in MN 6. In fact, this local fauna is more
young than the Sansan local fauna. The Caproid~s migration (visible in the Manchones
fauna) is an important generic event as shown by Koehler (1988).
Some events, formerly associated with the early Turolian, such as the occurrence of several Hipparion or the first appearance of Parapodemus, seem now to occur
in late Vallesian. The Austrian locality Kohfidisch fits better in late Vallesian than
early Turolian.
The subdivision of Turolian into three units is relatively easy in western Europe
but more difficult in eastern Europe and west Asia. For this reason the position of
eastern localities are only suggested here without absolute certainty. The first
appearance of Hystrix seems to occur earlier in North Africa and eastern Europe than
it does in western Europe.
The Ruscinian has few new records other than the discovery of Mimomys in
Perpignan (Serrat-d'en-Vacquer), which is a good indicator for MN 15.
Persons studying "Villafranchian" faunas generally work with precise biological
events. It seems that the arrival of many important Villafranchian genera occur not
exactly at the same time and in western Europe will allow the further subdivision of
MN zones.
During the Miinchen Symposium in 1975 about mammalian stratigraphy
(Fahlbusch, 1976), some localities were selected to illustrate each MNzone. It.seemed
useful to give here (table Z) the complete mammalian faunal list of these localities.
REFERENCES

Fahlbusch, V., 1976. Report on the Internatioanl Symposium on Mammalian Stratigraphy of the European Tertiary. Newsletter Stratigraphy, v. 5 (Z/3), p. 160-167.
Kohler, M., 1988. Boviden des tUrkischen Miozans (Kanozoikum und Braukohlen der
Tiirkei. 28). Paleont. i Evol. t. 21, 1987, p. 133-246.
Mein, P., 1975. Resultats du groupe de travail des vertebres: Biozonation du Neogene
mediterraneen a partir des mammiferes, in Senes, J. (ed.), "Report on Activity
of the RCMNS Working Groups (1971-1975f," Bratislava, p. 78-81.
Mein, P., 1976. Biozonation du Neogene mediterraneen partir des mammiferes, in
Proceedings, VIth Congress, RCMNS, Bratislava, September 4-7, v. z.

90