Revision of the genera Parapodemus, Apodemus, Rhagamys and Rhagapodemus (Rodentia, Mammalia)

l

REVISION OF THE GENERA PARAPODEMUS, APODEMUS, RHAGAMYS AND RHAGAPODEMUS (RODENTIA, MAMMALIA)

ELVIRA MARTIN SUAREZ & PIERRE M~EIN MARTIN SUAREZ E. & MEIN R 1998. Revision of the genera Parapodemus, Apodemus, Rhagarnys and Rhagapodemus (Rodentia, Mammalia). [Rdvision des genres Parapodemus, Apodemus, Rhagamys et Rhagapodemus (Rodentia, Mammalia)]. GEOBIOS, 31, 1: 87-97. Villeurbanne, le 28.02.1998. Manuscrit dgpos~ le 23.05.1996; acceptg d~finitivement le 01.10.1996. ABSTRACT - This paper presents a revision of the taxonomy of the genera Apodemus KAuP, 1826, Parapodemus SCHAUB,1938, Rhagamys FORSYTHMAJOR, 1905 and Rhagapodemus KRETZOI,1956. A new classification is proposed, that makes Mus gaudryi DAMES, 1883 from the Late Miocene the only species of the monospecific genus Parapodemus. Other species formerly included in this genus are transferred to the genus Apodemus. Rhagapodemus prirnaevus (HUGUENEY& MEIN, 1965) from the locality of Lissieu is redefined. KEY WORDS: RODENTIA, MURIDAE, TAXONOMY, PHYLOGENY. Rt~SUMt~ - Ce travail contient une r~vision de la taxonomie des genres Apodernus KAuP, 1826, Parapodemus SCHAUB, 1938, Rhagamys FORSYTHMAJOR, 1905 et Rhagapodemus K~ETZOI,1956. Une nouvelle classification est propos~e, qui fait Mus gaudryi DAMSS, 1883 du Miocene final la seule espgce du genre monosp4cifique Parapodemus. D'autres esp~ces, jusqu'h maintenant incluses dans ce genre sont transf~rges au genre Apodemus. Rhagapodemus prirnaevus (HuGUENEY& MEIN, 1965) du gisement de Lissieu est red~fini. MOTS-CL]~S: RODENTIA, MURIDAE, TAXONOMIE, PHYLOGI~NIE.

INTRODUCTION The oldest Muridae known (with the exception of

t h a t in all known populations the s t e p h a n o d o n t y is complete.

T here exist clear differences b e t w e e n the oldest Vallesian, and in t h a t time the representatives of populations t h a t were assigned to t he genus various lineages m a y be distinguished (Mein et al. Parapodemus, like P. lugdunensis (ScHAUB, 1938), 1993): Progonomys, Mus, "Parapodemus", and the and the later populations of Apodemus, e.g. the bett er development of the t m a in the M1 in n e a r l y all Muridae indet, from Can Llobateras. the species, the disposition in chevrons of the cusp The first r e p r e s e n t a t i v e of"Parapodemus" is found pairs of the lower molars, the b e t t e r development in t h e E a r l y Vallesian locality B u z h o r 1 in of the t7 in the M 1 and M 2 of the younger populaMoldavia (Lungu 1981, and new, unpubl i shed tions. This means, t h a t the generic differences (bedata). The mat e r i al is scarce: one M 1 of small size, tween Parapodemus and Apodemus) were based 1.78 x 1.04 mm, with complete stephanodonty, or in upon the degree of development of different charother words, the cusps t4, t5, t6, t9 and t8 united in acter states, which m a y p r e s e n t a problem in intera continuous loop; t7 is not developed, but instead mediate populations: the individuals of one popuof it t h e r e is a continuous high crest (see Mein et lation (per definition one single species) might al. 1993, Fig. 2f). In W. Europe the oldest records of belong to two different genera, according to the "Parapodemus" are from the localities Amb6rieu 1 distribution of character states in the morphoand Soblay (Farjanel & Mein 1984), where it is types. And such a situation is, in our opinion, not associated w i t h Progonomys cathalai SeHAUB, recommendable. 1938. In the Iberian Peninsula it appears later, in t h e L a t e s t Vallesian locality Cascante-Cubla. Therefore, and taking into account t h a t the holotyDuring the E a r l y Turolian "Parapodemus" is a pe of the type-species of the genus Parapodemus is usual component of the fauna. It is interesting, lost, we t hi nk t h a t the best solution for the stabil-

Antemus JACOBS, 1977) are d a t e d as E a r l y

88 ity of the nomenclature is to include in the genus

Apodemus those species that, in our opinion, form one evolutionary lineage with several ramifications of related species. It is not our intention to revive an old polemic on this subject. Therefore, if our new proposal for a classification of Apodemus and Parapodemus causes serious objections, we t h i n k it will be convenient to submit the case to the International Commission on Zoological Nomenclature. Family MURIDAEGray, 1821 Subfamily MURINAE Gray, 1821

Parapodemus SCHAUB,1938

Genus

Type-species - Mus gaudryi DAMES,1883. We propose

to

maintain

the

generic name

Parapodemus SCHAUB,1938 with only one species, P. gaudryi (DAMES, 1883), from the locality of Pikermi (Attica, Greece). Its holotype was figured in Papp (1947), and has subsequently been lost. No other known population may be attributed to this species. Discussion

-

Dames (1883) creates the species

Mus gaudryi on the basis of limited material (several lower molars) from the locality of Pikermi. The specimens were deposited in the Geological Institute Museum of the University of Athens and have been lost. Schaub (1938) creates the genus Parapodemus for fossil Muridae with a dentition similar to t h a t of Apodemus, stating t h a t "Hiigel 7 ist bei ihnen nicht entwickelt" (t7 is not developed). He designates Mus gaudryi DAMEs, 1883 as type-species of the new genus (Schaub 1938, p. 14), and considers the material from Polg~rdi and Pikermi to belong to the same species. In fact the definition of the genus is based upon the upper dentition t h a t has not been found in Pikermi. Papp (1947) figures the original material of Dames and notes t h a t the form from Polg~rdi is much larger and cannot belong to the same species as the mandible from Pikermi. Therefore he creates Parapodemus schaubi PAPP, 1947 (p. 373) for the Polgfirdi population, and then he proposes it as type-species of the genus Parapodemus. This designation is clearly incorrect after article 61 of the ICZN: once a type-species has been designated it can only be changed by the International Commission on Zoological Nomenclature. On the other hand the name P. schaubi is a valid name according to Art. 11 of the ICZN. De Bruijn (1976) studies the Parapodemus fossils from a locality near Pikermi (Chomateri), which is more t h a n a kilometre away from Dames' locality,

and determines them as P. gaudryi. In the cited paper he proposes an emended diagnosis of the genus Parapodemus and designates a neotype for P. gaudryi amongst the material from Chomateri. He suggests t h a t the Parapodemus collection from Chomateri belongs to the same species as the specimens from Polg~rdi (de Bruijn 1976), and therefore considers P. schaubi PAPP, 1947 to be a junior synonym of Mus gaudryi DAMES, 1883 (see also van de Weerd, 1976). Unfortunately the only dental elements t h a t permit a comparison with the material studied by Dames (1883), and figured by Papp (1947), are the M2 and M3, which are not figured by de Bruijn (1976). Mein (1978) analyses the paper by de Bruijn (1976) and treats two different aspects. On the one h a n d he proposes to m a i n t a i n the designation by Papp ofP. schaubi as type-species of the genus, which is an error as demonstrated by van de Weerd & de Bruijn (1978). On the other h a n d Mein (1978) considers the neotype designation by de Bruijn (1976) to be invalid because there are doubts about the correlation of Chomateri with the classical locality of Pikermi, and because the neotype may belong to another species (and even genus) t h a n the lost holotype. The neotype designation should be rejected on the basis of three paragraphs of Art. 75 d of the ICZN: 1.- De Bruijn (1976) designates a neotype without any justification. Art. 75 c of the ICZN states, t h a t a neotype designated as a m a t t e r of curatorial routine has no standing in nomenclature. Since De Bruijn's text makes no reference to the neotype, except in the proper designation, it must be considered invalid. 2.- De Bruijn (1976) says t h a t he chose the richest one of four fossiliferous levels at Chomateri. Any one of these four (or none of them) could be equivalent to the original level of Pikermi. So, the author fails to give evidence t h a t Chomateri and the classical locality of Pikermi represent the same geological horizon, as required by article 75 d(5) of the ICZN for the valid designation of a neotype. In fact these two localities are more t h a n a kilometre apart, and t h a t circumstance, in continental sediments, makes it almost impossible to prove t h a t they represent the same horizon. 3.- In the locality of Chomateri three murids of similar size are found, and one cannot ascertain to which one, if any, of these three belongs the material studied by Dames. The specimens from Pikermi figured by Papp (1947) remind one more of Occitanomys ? neutrum DE BRUIJN, 1976 from Chomateri t h a n of P. schaubi by the absence of accessory labial cusps. De Bruijn (1976) says t h a t P. gaudryi from Chomateri is identical to P. schaubi, whereas

89 Papp (1947) states t h a t P. schaubi is larger t h a n P. gaudryi. For these reasons the neotype designation does not fulfill the requirement of Art. 75 d(4), t h a t the neotype should be consistent with what is known of the former type. Species

excluded

from

the

genus

Parapo-

demus:

- Mus hipparionum SCHLOSSER, 1924 was transferred by Schaub (1938) to the genus Parapodemus, then by Storch (1987) to the genus Karnimata JACOBS, 1978. Finally Mein et al. (1993) consider Karnimata to be a junior synonym of Progonomys, but Mus hipparionum does not belong to the genus Progonomys, for which reason this species is classified as Murinae incertae sedis. Parapodemus lugdunensis SCHAUB, 1938; we transfer this species to the genus Apodemus. - Parapodemus vireti SCHAUB, 1938 was transferred by Michaux (1969) to the genus Valerymys. Later it was transferred to the genus Huerzelerimys MEIN et al., 1993. - Parapodemus coronensis SCHAUB, 1938 is known by very limited populations. It is probable t h a t under this name populations of two different species have been described. On the one hand, the Pleistocene Parapodemus coronensis (Schaub 1938) seems to be related to Apodemus microps KRATOCHVIL• ROSICKY, 1952; this latter species is an oriental form, probably the vicariant species in W. Europe is Apodemus maastrichtiensis VAN KOLFSCHOTEN, 1985. On the other h a n d Kowalski (1956) cites P. coronensis from the Polish locality Podlesice (Pliocene); in its upper molars the t12 is present. The mentioned author does not describe the roots. It is possible t h a t these populations are related to Micromys DEHNE, 1841. - Progonomys orientalis SCHAUB, 1938 was included in the genus Parapodemus by Thaler (1966), and finally it was called Apodemus orientalis by Storch (1987). - Parapodemus similis ARGYROPULO& PIDOPLICHKA, 1939 was transferred to the genus Orientalomys by de Bruijn & van der Meulen (1975). - Mastomys colberti LEWIS, 1939 was included in the genus Parapodemus by Misonne (1969). Black (1972) transfers it to the genus Rattus FRISCO, 1775 and Jacobs (1978) includes it in Karnimata. In his fig. 23 Misonne (1969) figures the only fragm e n t of a maxillary known for this species. It shows t h a t there is no connection whatsoever between t4 and t8, neither in the M ~nor in the M s, though the teeth are very much worn. The M s has four roots, an u n k n o w n feature in "Parapodemus". Consequently we classify this species as Murinae incertae sedis. - Parapodemus schaubi PAPP, 1947 from Polg~rdi is transferred to the genus Apodemus. De Bruijn

(1976) states that the population from Chomateri "appears to be the same species as P. schaubi from Polg~rdi". Therefore the same specific name should be applied to both populations, namely Apodemus schaubi (PEP, 1947) and not Parapodemus gaudryi. - Parapodemus albae K~ETZOI, 1951. This is a murid from the locality of Cs~tkv~r, where it is found together w i t h Neocricetodon schaubi KRETZOI, 1930. This association, as well as its small size (MI: 1.8 x 1.0 mm) make us think, t h a t we are dealing with Apodemus lugdunensis. It was never figured. - Parapodemus jordanicus HAAS, 1966 is based on digested specimens of Apodemus. - Parapodemus adroveri THALER, 1966 was later called Occitanomys adroveri by Michaux (1969). - Parapodemus gaudryi barbarae V~N DE WEERD, 1976 is considered a species, and not a subspecies, by numerous authors. We now call it Apodemus barbarae. - Parapodemus meini MARTIN SUAREZ & FREUDENTHAL, 1993 is transferred to the genus Apodemus. Genus A p o d e m u s KAuP, 1826 Type-species - Mus agrarius P~LA& 1778

- Mnridae with a wide size range. Upper molars with a connecting crest between t4 and t8, which in the course of time becomes inflated and forms a t7. The t6 and t9 are united. The t12 is present in the fossil forms, with a tendency to reduction in the majority of the more recent forms. M 1with t l in an anterior position and three or four roots. M 2without tlbis. Lower molars rarely with longitudinal connections between cusps; the cusp pairs tend to form chevrons. M~ with t m a nearly always present. Diagnosis

N.B. - Niethammer (1978), being a zoologist, bases the diagnosis of Apodemus on dental characters, body size, and soft tissue (op. cit., p. 305). Our diagnosis uses dental characters only and is applicable to both fossil and recent material.

Fossil species included in this genus: Apodemus agustii MARTIN-SUAREZ,1988 Apodemus albae (KRETZOI,1951) =A. lugdunensis SCHAUB,1938 Apodemus alsomioides SCHAUB,1938 Apodemus atavus I-IELLER,1934 Apodemus barbarae (VANDE WEERD, 1976) Apodemus caesareanus BATE, 1942 Apodemus coronensis (SCHAUB,1938) Apodernus debruijni KOTLIA,1992 Apodemus dominans KRETZOI,1959 Apodemus etruscus ENGESSER,1989 Apodernus flavicollis (MELCHIOR,1834) Apodemus gorafensis Rvlz BusTos et al., 1984 Apodemus gudrunae VAN DE WEERD, 1976 Apodemusjeanteti MICHAUX,1969

90 Ieptodus KRETZOI, 1956 levantinus BATE, 1942 lewisi NEWTON, 1899 lugdunensis (SCHAUB, 1938) maastrichtiensis VAN KOLFSCHOTEN, 1985 manu THALER, 1974 Apodemus maxirnus THALER, 1972 Apodemus meini (MARTINSUAREZ• FREUDENTHAL,1993) Apodemus microps KRATOCHVIL& ROSICKY,1952 Apodemus mirabilis (SHEVCHENKO,1965) Apodemus mystacinus (DANDFORD& ALSTON,1877) Apodemus orientalis (ScHAUB, 1938) Apodemus peninsulae THOMAS, 1906 Apodemus qiui Wu WENYU & FLYNN, 1991 Apodemus schaubi (PAPP, 1947) Apodemus sylvaticus (LINNE, 1758) Apodemus whitei HINTON, 1915 Apodemus zhangwagouensis Wu WENYU & FLYNN, 1991 Apodemus Apodemus Apodemus Apodemus Apodemus Apodemus

N.B. - We have tried to cover all the fossil species of Apodemus found in the litterature. It is, however, possible, t h a t we missed some of them, but this does not mean t h a t we do not include them in this genus.

P h y l o g e n y - As we have said in the introduction, the oldest representative of Apodemus is found in the Early Vallesian locality Buzhor (Lungu 1981). We have had the opportunity to study the collection from this locality. The tooth of Apodemus is of small size, with a derived morphology, in which the connecting crest between t4 and t8 is high and well-developed. We think, t h a t the f a u n a from Farafra (Heissig 1982) is not homogeneous. We also think the faunas from Castelnou i (Aguilar et al. 1995), Castelnou 1B (Aguilar et al. 1991a) and Castelnou 3 (Aguilar et al. 1991b) are heterogeneous. These fossiliferous points have yielded mixtures of middle Miocene, late Miocene (Vallesian and Turolian) and Pliocene faunas. In our opinion the origin of Apodemus is unknown. Apodemus lugdunensis is certainly not a descendent ofProgonomys cathalai (as has been proposed by various authors) for several reasons: 1) The oldest european species ofApodemus (Apo~ demus sp. from Buzhor) is older t h a n the oldest record of Progonomys cathalai. Only Progonomys sp. from Siwaliks (Jacobs & Downs 1994) m a y be older t h a n Apodemus. 2) The oldest Apodemus lugdunensis are smaller t h a n Progonomys cathalai, and have a clearly more derived morphology. 3) The M1 of P. cathalai has a third root t h a t is maintained in its descendents (Mein et al. 1993): in P. woelferi BACHMAYER• WILSON, 1970 and in the species of the lineages Huerzelerimys and Anthracomys SCHAUB, 1938. On the other h a n d this third root is not found in Apodemus.

Several localities of the beginning of the Late Vallesian (MN 10) in W. Europe contain Progonomys cathalai, but there are no representatives of the lineage Apodemus. This is the case in Ravin de la Pluie (Bonis & Melentis 1975), Montredon, Masia de] Barbo, Peralejos A and Peralejos 4. The association of P. cathalai and A. lugdunensis is slightly younger and is found in the Vallesian localities of Soblay and Amb~rieu I (Farjanel & Mein 1984). Later, in the Latest Vallesian, P cathalai is substituded by its descendents, P. woelferi or Huerzelerimys minor, t h a t are generally associated with Apodemus lugdunensis. This is the case in the localities Kohfidisch (Bachmayer & Wilson 1980), Amb6rieu 2C, Cascante-Cubla, Cucaldn and Dionay. Figure I represents the chronological distribution and phylogenetic relationships of (the populations and) species of Apodemus. In the localities of Soblay (Farjanel & Mein 1984) and Crevillente 2 (Martin Sufirez & F r e u d e n t h a l 1993) Apodemus lugdunensis is abundant, together with some rare teeth of an Apodemus sp., of very small size and a strongly derived morphology, t h a t cannot be determined, due to the scarcity of the material. From the Turolian onwards, in nearly all the known localities some species of Apodemus is found. During the Early Turolian ( M N l l ) the species A. lugdunensis is found, e.g. in the localities Puente Minero (Alcalfi et al. 1991), Crevillente 2 and 4B (Martin Sufirez & F r e u d e n t h a l 1993), Aguanaces, Vivero de Pinos (Adrover 1986), Eichkoge], Mollon, Lobrieu, Tortajada A, etc... In the "Middle" Turolian (MN12) the representative is A. barbarae, e.g. in the localities Masada del Valle 2, Crevillente 5A and 15, Casa del Acero, Concud, Los Mansuetos, Aljezar B, etc... This is the period when, in our opinion, a first diversification of Apodemus takes place: in Aljezar B we find two species, A. barbarae and Apodemus sp. (Adrover 1986); Crevillente 8 also shows the association of two species: A. barbarae and A. meini. This latter species might be the ancestor of Apodemus sp. from Lissieu. In the Middle and Upper Turolian (MN12 and MN13) we know several localities in which no species of Apodemus is found, but in some of them the genus Castromys is represented. From a biostratigraphic point of view these localities are situated between the disappearance of Apodemus meini and the first appearance of Apodemus gudrunae. The sequence of Crevillente permits this conclusion, since it contains numerous localities in lithostratigraphic superposition (Martin Su~rez & Freudenthal 1994): In CR7 and CR8 we find A. meini; CR17 and CR22 are slightly younger and

91 FIGURE 1 - Chronological distribution and phylogenetic relationships of (the populations and) the european species of Apodemus. Distribution chronologique et relations phyloggn4tiques des (populations et des) esp~ces

A. agrarius O I-

__..:

,/

,i

D.I ~"

d'Apodemus. I I I

Arrondelli , Schemfeld Alquerla

17

!

Orrios Balaruc-2 Pla de la V)lte Csamota-2 S@te C.Castatlo

16 uJ

.~ E

,,.J

=-i

1Lt

Seynes I Pla de la Ville C. Mahora

P~

Mont Hdl*~ne Perpignan

~-

E "~ Hautimagne ~- Cellsneuve

~) Gloria-4

Peralejos-A • La Tour.

,La T o u r - -

?

Apodemus s p .

Lu oieu

:~] CR-7 ~ •~ "

Alcoy

/

~3:ravaca

~

Mont H$l~ne

,,. ,, Vendargues

~ "

/

Aljezar-B Mansuetos

I OR-8

12

o

~i

~

3

~" '~ .~,

Vendargues Gorafe 1, A

~ "~

"Masada del VaUe 7 Valdecebro 3, 6 " Lissieu , ,,,,'

"

13

E

S~te Arquillo-3 J C.Casta5o V.Alta . , - A. agusti~ Mont I--~l~ne Perpignan

Pla de la Ville Belrnez-1 S~te

Apodemussp. 2

Concud Casa del Acero

" ~ CR-15

Pikermi

Parapodemus I ga udryi

"~ CR-5A Masada del Valle 2

ILl Z I,LI O

"~ortaja~a-A

_o .~

1] CR-2 I

i0

~,

~

Soblay

Mollon, Lobrieu Eichkogel, Amberieu-3 Vivero de Pinos Aguanaces, CR-4B CR-2 Puente Minero

Kohfidisch Cucaloa, Dionay ~. Cascante-Cubla '< Amberieu-1, 2C Soblay Buzhor

9

lack

Apodemus, but contain Castromys littoralis

MARTIN SUAREZ •

FREUDENTHAL, 1994. T h e l a s t

localities without Apodemus in the sequence of Crevillente are CR14 and CR31, in which we find

Castromys inflatus

(MEIN, MOISSENET & ADROVER,

1990). In a younger level, CR6, we find Apodemus gudrunae (Apodemus cf. primaevus in de Bruijn et al. 1975). The localities of La Gloria 6 and Velillia 2 in Teruel (Mein et al. 1990) are the only ones wh er e Apodemus gudrunae and Castromys inflatus are associated.

Parapodemus gaudryi from the Middle Turolian locality Pikermi is a descendent of some species of

Apodemus. Apodemus barbarae from the Middle Turo]ian is the most probable ancestor of A. gudrunae from the Late Miocene and the l at t er one probably gave rise to the E a r l y Pliocene A. gorafensis (it is possible t h a t A. gorafensis is al ready p r e s e n t in some Latest Miocene levels). In this lineage t h e r e is a size increase in the course of time and the mor-

92 phological c h a n g e is expressed by t h e r e d u c t i o n of t h e l o n g i t u d i n a l connections b e t w e e n the cusps. The oldest localities in w h i c h A. gorafensis h a s b e e n found, are C a r a v a c a a n d Alcoy (see Fig. 1), a n d this species is r e p r e s e n t e d in n u m e r o u s y o u n g e r localities: Gorafe 1, Gorafe A, Vendargues, P e r p i g n a n , P l a de la Ville, etc. I n t h e Guadix- B a z a B a s i n A. gorafensis gives rise to A. agustii, w h i c h is only k n o w n from its type-locality, C a f i a d a del Castafio 1 (MN15) ( M a r t i n Su~rez, 1988), w h e r e it is associated w i t h A. dominans. D u r i n g t h e E a r l y Pliocene, a n d also from A. gorafensis, a second species arises, A. jeanteti, of similar size or larger, w i t h u p p e r m o l a r s t h a t are practically i n d i s t i n g u i s h a b l e from t h o s e ofA. gorafensis, b u t w i t h M1 t h a t h a v e lost the t m a or the anter o c e n t r a l a n t e r o c o n i d . A. jeanteti h a s b e e n e n c o u n t e r e d in fissure fillings only, e.g. Mont H~l~ne, Belmez, Seynes. T h e s e species of Apodemus of large size h a v e d i s a p p e a r e d in t h e L a t e Pliocene. N e v e r t h e l e s s , it is possible, t h a t the Q u a t e r n a r y Apodemus mystacinus (with a welldeveloped p o s t e r o s i n u s in the u p p e r molars, a n d w i t h a n isolated labial a n t e r o c o n i d in the MI) is a d e s c e n d e n t ofA. gorafensis. D u r i n g t h e entire Pliocene (and possibly in some L a t e s t Miocene locality) in W. E u r o p e a small species, Apodemus dominans, is v e r y a b u n d a n t in m a n y localities: Peralejos A, L a Tour, La Gloria 4, M o n t H~l~ne, B a l a r u c 2, etc. We think, t h a t the E a r l y Pleistocene A. atavus is a d e s c e n d e n t of A. dominans. The Pleistocene species A. sylvaticus a n d A. flavicollis (from t h e e n t i r e P a l e a r c t i c region) m a y well be d e s c e n d e n t s ofA. atavus. So, d u r i n g t h e E a r l y a n d Middle Pliocene (zones MN14, 15 a n d 16A) of W. E u r o p e two lineages of Apodemus are p r e s e n t , one of small size, a n d a second l a r g e r one. A p a r t from these, in some localities as L a Tour, Celleneuve, H a u t i m a g n e , C u e s t a de M a h o r a , etc., we find a n Apodemus of i n t e r m e diate size b e t w e e n A. dominans a n d A. gorafensis. I n the W e s t e r n E u r o p e a n Pleistocene we find A. agrarius, which, in our opinion, is an Asiatic inmig r a n t , t h a t is c o m p l e t e l y u n k o w n from earlier deposits in t h e area. We also find A. maastrichtiensis, w h i c h is a poorly k n o w n Apodemus of small size. We t h i n k it m a y be r e l a t e d to A. coronensis (sensu S c h a u b 1938). A. microps m a y be a d e s c e n d e n t ofA. maastrichtiensis. W i t h t h e n e w definition t h a t we propose, repres e n t a t i v e s of t h e g e n u s Apodemus are k n o w n f r o m t h e E a r l y Vallesian (locality B u z h o r in Moldavia) u n t i l r e c e n t time. This m e a n s t h a t Apodemus a n d Mus are t h e M u r i d a e w i t h the longest t i m e r a n g e k n o w n .

G e n u s R h a g a m y s FORSYTH MAJOR, 1905

Type-species - Mus orthodon HENSEL,1856 W h e n F o r s y t h Major (1905) c r e a t e d the g e n u s it to be closer to t h e we n o w call Apodemus g r o u p Ratten ( w h a t we n u m b e r of reasons, b u t he did n o t give a diagnosis of t h e genus.

Rhagamys he c o n s i d e r e d group Miiuse ( t h a t is w h a t and Micromys) t h a n to t h e call Mus a n d Rattus) for a

Diagnosis.- M u r i d a e of l a r g e size, w i t h a relatively h i g h crown. I n t h e lower m o l a r s the cusp pairs form v e r y p r o n o u n c e d chevrons, even in t h e Ms. I n the M 1 the t2 a n d t3 are united, M 2 w i t h o u t t3 a n d t12. W e a r surfaces flat. Remarks - The populations that have been attributed to

Rhagamys orthodon are insular, from Corsica and Sardinia. They are large-sized in comparison with their continental ancestors. The morphology is strongly derived, e.g. the crowns are very high and the arrangement of the cusps is somewhat different (see the discussion by Schaub, 1938). For the moment we consider it to be a monospecific genus, although the great time range of its populations advocates the possibility that there existed more than one species. Rhagamys orthodon may be a descendent of some species of Rhagapodernus or even ofApodemus. The highly derived motphology that is observed in R. orthodon is due to island conditions. Brandy (1978) considers Rhagarnys orthodon to be a descendent ofRhagapodernus mino5 and the latter one might be a descendent of some other species of Rhagapodemus. In this case, in a systematic classification, all these species should belong to one single genus. But with the available data we cannot resolve the origin ofRhagamys orthodon.

Genus

Rhagapodemus I~ETZOI,

1959.

Type-species - Rhagapodemus frequens KRETZOI,1959. O r i g i n a l d i a g n o s i s - A r r a n g e m e n t of t h e cusps b a s i c a l l y Apodemus-like b u t t h e m o l a r s are h i g h e r crowned a n d t l on M ~ isolated. O t h e r species

Rhagapodemus athensis DE BRUIJN & VAN DER MEULEN, 1975

Rhagapodemus ballesioi MEIN & MICHAUX, 1970 Rhagapodemus hautimagnensis MEIN & MICgAUX, 1970

Rhagapodemus minor (BRANDY, 1978) Rhagapodemus primaevus (HuGUENEY & MEIN, 1965)

Rhagapodemus vanderweerdi DE BRuIJN & VAN DER MEULEN, 1975

Rhagapodemus sp. from E r t e m t e (Storch 1987) Remarks - The populations that have been determined as one of the species ofRhagapodemus have a dental pattern that is similar to the one ofApodemus, but the crown is higher and the cusps are more vertical. The labial cingulum of the lower molars bears various cusps that, in the younger populations, attain the same height as the main cusps. In the upper molars the tl is isolated, and longitudinal connections between the cusps are very rare. The genus Rhagapodernus includes various species. For some of them an ancestor-descendent relationship may be assumed

93 FIGURE 2 - Chronological distribution and phylogenetic relationships of (the populations and) the species of Rhagapodemus. Distri-

Capo Figari] Ft. minor J

UJ

,.J 0.

bution chronologique et relations phylogdndtiques des (populations et des) esp~ces de Rhagapodemus.

Tourkoubounia] R. athensis J

17

Segdes Le grand Serre Lo Fournas-4 Seynes

16

Rebielice IJJ

Pla de la Ville

5

8 -2

Csarnota-2 Layna Poblado lberico Sete Belrnez-1 Orrios-3

R.frequens

15

Mont Helene Perpignan R.hautirnagnensis

14

0•

13

R.primaevus

(Fig. 2). This is the case for R. primaevus fi'om the Late Miocene of Lissieu, t h a t may be considered to be the ancestor ofR. hautimagnensis from the Early Pliocene. This latter one may be considered to be the ancestor of R. frequens from the Middle Pliocene. In this lineage, R. primaevus - R. hautimagnensis - R. frequens, we see a size increase in the course of time, as well as modifications in the morphology: the percentage of M 1 with isolated t l increases with time; the labial cingulums of the lower molars increase in height, and attain the same level as the main cusps in the latest populations; the longitudinal connections between cusps are more abundant in the older populations. In W. Europe R. frequens is the youngest species known of the genus Rhagapodemus and it disappears from the fossil record in the Middle Pliocene. In the Early Pliocene locality of Hautimagne (Mein & Michaux 1970) R. hautimagnensis is associated with another species of the same genus, R. ballesioi, ofsmaller size, whose origin is for the moment unknown. R. ballesioi is also found in the locality of Mont-H~l~ne (Aguilar et al. 1986). In the Pleistocene only two species of this genus are known: Rhagapodemus minor from Capo Figari in Sardinia and R. athensis from Tourkobounia-1 in Greece. Rhagapodemus minor was originally described as Rhagamys minor by Brandy (1978), but in our opinion, the fact t h a t it has been found on an island does not necessarily mean that it be a strictly insular Muridae. Some characters of its morphology lead to consider this population as a species of the genus Rhagapodemus, specially in the M 2 because the t3 is present and the t12 is well developed. Aguilar et al. (1995) created the species Rhagapodemus primitivus based on material from Castelnou 1. The holotype, CTN n°29, is an M s of large size (2.48 x 1.54 ram) which, in our opinion, presents unusual characters for a species of the genus Rhagapodemus: the cusps are oblique and not vertical, the t12 is very much reduced, the connection t9-t8 lies very much backwards (type Apodemus), the crown is low. This morphology

Hautimagne-1 Vendargues Celleneuve Ptolemais-1 La Gloria 4 Peralejos La Tour

R.ballesioi RMaritza • vandeweord_~.

Lissieu agrees with that ofApodemus jeanteti MIC~A~X, 1967, specially with that of the populations from Mont H~l~ne and from Font Estramar. We therefore consider R. primitivus to be a junior synonym of Apodemus jeanteti (which is only known from the Pliocene). The other M ~ (CTN n°26) figured in Aguilar et al. (1995) might have belonged to stone intermediate population between the type-populations ofR. primaevus and R. hautimagnensis. This is one of the reasons that make us think that Castelnou 1, as well as other localities in the same area, contain heterogeneous faunal assemblages. In E. Europe Rhagapodemus is represented by two species: R. vandeweerdi from the Early Pliocene of Maritsa and R. athensis from the Old Pleistocene of Tourkobounia (de Bruijn & van der Meulen 1975). These populations are separated by a large time interval and it is difficult to establish any kind of relation between them.

Discussion on Rhagapodemus

the genera Rhagamys and - We think, t h a t for the benefit

of stability of the nomenclature it is desirable to m a i n t a i n Rhagamys and Rhagapodemus as two separate genera. The only species of the genus Rhagamys, R. orthodon is exclusively insular. The dental patterns of the populations of species attributed to these two genera show differences t h a t support their independency: - The lower molars of the species of Rhagapodemus have a very well-developed labial cingulum, with accessory cusps that, in the youngest populations, attain the same height as the main cuspids. In Rhagamys there are no labial cingulures, since the accessory cusps are completely united to the main cuspids.

94

L

M~ M2 M3 MI M2 M3

W

n

min.

mean

max.

V'

27 25 36 29 39 34

1.56 1.17 0.95 1.77 1.15 0.85

1.806 1.322 1.136 1.997 1.344 1.013

2.00 1.42 1.33 2.26 1.55 1.20

24.72 19.31 33.33 24.32 29.63 34.15

a 0.111 0.059 0.099 0.132 0.088 0.087

L/W

n

rain.

mean

max.

V'

27 25 36 29 39 34

0.93 1.07 0.88 I.i0 i.I0 0.80

1.068 1.159 1.029 1.232 1.232 0.982

1.22 1.30 1.17 1.36 1.44 1.16

26.98 19.41 28.29 21.14 26.77 36.73

0.072 0.059 0.076 0.065 0.067 0.095

1.69 1.14 i.i0 1.62 1.09 1.03

TABLE 1 - M e a s u r e m e n t s of the m o l a r s of Rhagapodemus primaevus from Lissieu (type-locality). Mesures des molaires de R h a g a p o d e m u s p r i m a e v u s de Lissieu (localitg-type).

FIGURE 3 - Rhagapodemus primaevus (HUGUENEY• MEIN, 1965) from its type-locality Lissieu. 1.- M 1 sin., 2.- M ~ dext., 3.- M 2 sin., 4.- M ~ dext., 5.- M 3 sin., 6.- M 3 detx., 7.- M~ sin., 8.- M~ dext., 9.- M~ sin., 10.- M2 dext., 11.- M3 sin., ]2.- M~ dext. Scale r e p r e s e n t s 1 mm.

- T h e a r r a n g e m e n t in c h e v r o n of t h e cusp p a i r s is m u c h m o r e p r o n o u n c e d in Rhagamys orthodon. I t e v e n affects t h e h y p o c o n i d - e n t o c o n i d complex of t h e M3. - I n t h e M1 of t h e species of Rhagapodemus t h e a n t e r o c o n i d is u n i t e d to t h e second p a i r of cusps. I n R. orthodon it is f r e q u e n t l y s e p a r a t e d a n d t h e second p a i r of c u s p s f o r m s a l a m i n a . - I n t h e M1 a n d M2 of t h e species of Rhagapodemus t h e t e r m i n a l h e e l is h i g h w i t h a r o u n d or oval outline, c l e a r l y d i f f e r e n t i a t e d f r o m t h e hypoconid. I n t h e M1 of R. orthodon t h e t e r m i n a l h e e l is complerely c o n n e c t e d to t h e hypoconid, in t h e M2 it h a s t h e s a m e s h a p e as in Rhagapodemus. - I n t h e M 1 of t h e species of Rhagapodemus t h e t l is r o u n d , t h e t2 a n d t3 a r e a t t a c h e d b u t recog-

n i z a b l e as different cusps. I n R. orthodon t h e t l is not round, t2 a n d t3 a r e t o t a l l y fused, t4 is v e r y m u c h s e p a r a t e d f r o m t7, a n d t12 is v e r y v o l u m i nous. - T h e M 2 a n d M 3 of Rhagapodemus h a v e a t3, a n d this cusp lacks c o m p l e t e l y in t h e s a m e m o l a r s of

R. orthodon. - I n all t h e t e e t h of R. orthodon t h e c u s p s t e n d to f o r m t r a n s v e r s a l l a m i n a e , in Rhagapodemus t h e cusps are u n i t e d b u t d o n ' t f o r m s u c h l a m i n a e . - T h e w e a r s u r f a c e of t h e m o l a r s of R. orthodon is flat, in Rhagapodemus t h i s s u r f a c e is a n g u l a r , t h o u g h less m a r k e d l y t h a n in o t h e r m u r i d s w i t h a m o r e b r a c h y d o n t crown. Remarks o n Rhagapodemus primaevus f r o m t h e l o c a l i t y L i s s i e u - T h e locality of L i s s i e u is a

95 k a r s t fissure filling, several metres thick, t h a t has yielded a f a u n a in which dominate the rodents, 90% of t h e m being murids. It is i m p o r t a n t to note t h a t th e f a u n a of Lissieu, does not r e p r e s e n t a level of immigration, since all the species found in this locality were already p r e s e n t in the area beforehand. It is a peculiar faunal association because the Muridae are absolutely dominant, and in all of t h e m the crowns are relatively higher t h a n in contemporaneous forms from other areas. T h er e are t h r e e forms of Muridae with t7, or with connecting crest between t4 and t8: - A small brachyodont Apodemus. Probably this is an A. gudrunae of smaller size t h a n the typepopulation from VDC3 (van de Weerd 1976), of the s a m e size as t h e p o p u l a t i o n from V il l ast ar (Adrover et al. 1993). - A large Apodemus with a r a t h e r simplified morphology, which might be a descendent ofA. meini from Crevillente-7 (Martin Su~rez & F r e u d e n t h a l 1993). - An "Apodemus primaevus" HuaUENEY & MEIN, 1965 of small size and relatively high crown. In our opinion this is t he oldest known population, t h a t m a y be a t t r i b u t e d to the genus Rhaga-

podemus. In th e f r a m e w o r k of this revision we t hi n k it is useful to redefine the species Rhagapodemus primaevus on the basis of t he type-population from Lissieu. An up-to-date characterization of this species m a y contribute to solve the discussion t h a t we have cited in previous p a r a g r a p h s and t h a t p a r t l y still continues (Engesser 1989).

Rhagapodemus primaevus (HuGUENEY & MEIN, 1965) Fig. 3.1-12 Original reference -

Apodernus primaevus HUGUENEY &

MEIN, 1965, p. 112. T y p e - l o c a l i t y - Lissieu (RhSne, France). H o l o t y p e - M 1 sin. (1,95 x 1,22), n ° 65054 deposited in the D~partement des Sciences de la Terre, Universitg ClaudeBernard, Lyon (Hugueney & Mein 1965). M e a s u r e m e n t s : see Table 1.

E m e n d e d d i a g n o s i s - Rhagapodemus of small size and with a relatively low crown; t l completely isolated in only one third of the specimens; the labial cusps of the M1 and M~ are lower t h a n the m ain cuspids. D i f f e r e n t i a l d i a g n o s i s - Rhagapodemus primaevus differs from R. frequens and R. minor by its lesser size and by the lesser relative height of the crown and of the accessory labial cusps of the lower molars. It differs from R. athensis and from R. vanderweerdi by its smaller size and because it lacks a posterior spur in the t2 of the M 1. It differs

from R. hautimagnensis by its smaller size and by having more inclined cusps. R. hautimagnensis has very vertical cuspids. R. ballesioi and R. primaevus are very similar in size, and differ morphologically: the cusps of the molars ofR. ballesioi are more vertical, whilst in R. primaevus t h e y are more inclined. In all the M 1 ofR. ballesioi t he t l is isolated and the cusps t2 and t3 are fused. In th e M 1 of R. primaevus the t l is only isolated in one third of the specimens and the cusps t2 and t3 are very close together and convergent, but t h e r e tips r e m a i n separated (there is enamel between the two cusps). In the M 2 of R. baUesioi the connections of t l and t3 to t5 are t r a n s v e r s e and high and form the anterior wall of the tooth. In R. primaevus these connections are limited to the bases of the cusps. In the lower molars t he cusp pairs form chevrons, which are much more pronounced in R. ballesioi. R e m a r k s - Rhagapodernus prirnaevus is probably the oldest species within the genus. With the available data it is very difficult to decide whether the various species of Rhagapodemus are derived from R. primaevus, (or all derived from an elder, unknown form), or w h e t h e r they stem from different species of Apodemus and the observed similarity between t h e m is due to morphological convergences.

CONCLUSIONS In this paper we present a revision of the taxonom y of various g e n e r a of Muri dae r e l a t e d to Apodemus. Their species are characterized by the presence of a t m a in M1, and of a t7 (or a connecting crest between t4 and t8) in M 1 and M 2.

Parapodemus gaudryi (DAMES, 1883) is reduced to nothing more t h a n a name, as the original collection, including the holotype, has been lost, and t here is no possibility to locate the locality of Pikermi where the type-population was collected. The r e m a i n d e r of the species t h a t had been included in the genus Parapodemus, are transferred to the genus Apodemus or left as Muridae

incertae sedis. The genus Apodemus includes, in our opinion, a monophyletic group of species, of u n k n o w n origin. Its first representatives have been found in deposits of Vallesian age, and have a derived morphology, t h a t distinguishes t h e m from other murids of the same age. T hey cannot be descendents of Progonomys cathalai. By the new definition (a diagnosis is proposed) r e p r e s e n t a t i v e s of th e genus Apodemus are known from t he Vallesian until present time. The taxonomy of the genera Rhagamys and Rhagapodemus is also revised, and the species Rhagapodemus primaevus (HuauENEY & MEIN, 1965) from the locality of Lissieu is redefined.

96 Acknowledgements - We wish to express our gratitude to the Drs Lungu and Sen who put (even unpublished) material at our disposal for study and comparison; to Dr H. de Bruijn who allowed us to study the material from Chomateri deposited in Utrecht; to Dr Freudenthal for his help in the interpretation of the ICZN, for the translation of the manuscript, and because his critical comments are always of great interest for our work. We wish to thank Dr Jacobs and Dr Sen (referees) for their valuable comments. The photographs were made on the Zeiss 950 DSM of the University of Granada. This study was carried out within the frame-work of the Project "cambios clim~ticos en el Sur de Espafia durante el NeSgeno" of the R. Areces Foundation nd PB94-1265 of the DGICYT.

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E. M A R T I N S U A R E Z

Departamento de Estratigraffa y PaleontoIogla Universidad de Granada E-18071 Granada email: [email protected] P. M E I N

UFR des Sciences de la Terre Universit~ Claude-Bernard, Lyon 1 et UMR 5565 du CNRS 43 Bd. du 11 Novembre F-69622 Villeurbanne cedex