ACTA CARSOLOGICA	30/1	8	115-124	LJUBLJANA 2001
COBISS: 1.01
THE SMALL VERTEBRATE FAUNA (RODENTS, INSECTIVORES, AND REPTILES) OF [ANDALJA 1A (ISTRIA, CROATIA)
FAVNA MALIH VRETENČARJEV (GLODALCI, @UŽKOJEDI IN PLAZILCI) ŠANDALJE 1A (ISTRA, HRVATSKA)
JEAN-PIERRE AGUILAR1 & JEAN-YVES CROCHET1 & JACQUES MICHAUX2 & ANDREJ MIHEVC3 & MAJA PAUNOVIČ4
1	J.-P. Aguilar : Universite de Montpellier 2, Institut des Sciences de l'Evolution, Laboratoire de Paleontologie, CC 064, Place Eugene Bataillon, 34095 MONTPELLIER CEDEX 5, FRANCE,
E-mail: aguilar@isem.univ-montp2.fr
2	J.-Y. Crochet: Universite de Montpellier 2, Institut des Sciences de l'Evolution, Laboratoire de Paleontologie, CC 064, Place Eugene Bataillon, 34095 MONTPELLIER CEDEX 5, FRANCE,
E-mail: crochet@isem.univ-montp2.fr
3	J. Michaux: Universite de Montpellier 2, Institut des Sciences de l'Evolution, Laboratoire de Paleontologiede l'EPHE, CC 064, Place Eugene Bataillon, 34095 MONTPELLIER CEDEX 5, FRANCE,
E-mail: michaux @isem.univ-montp2.fr
4	A. Mihevc: Karst Research Institute, ZRC SAZU, Titov trg 2, SI - 6230 POSTOJNA, SLOVENIA, E-mail: mihevc@zrc-sazu.si
5	M. Paunovic: Institute of Quaternary Paleontology and Geology, Ante Kovacica 5, 1000 ZAGREB, CROATIA, E-mail: mpaunovi@public.srce.hr
Prejeto / received: 25. 1. 2001
Izvleček	UDK: 568/569(119.1)(497.5)
551.791:568/569(497.5)
Jean-Pierre Aguilar & Jean-Yves Crochet & Jacques Michaux & Andrej Mihevc & Maja Paunovič: Favna malih vretenčarjev (glodalci, žužkojedi in plazilci) Šandalje 1A (Istra, Hrvaška)
Pod lokacijo Šandalja 1A je opisana spodnje srednje pleistocenska favna malih vretenčarjev, ki vsebuje ostanke glodalcev, žužkojedov in plazilcev. Kostne breče so bile najdene leta 1999 v kamnolomu Šandalja pri Puli, njihova natančna lega v primerjavi znanim kostnim brečam Šandalje 1 pa ni natačno znana. Kljub temu, njena izpričana manjša - Biharijska starost - govori v prid manjši starosti favne velikih sesalcev Šandalje 1 in z njo povezanim prodnjakom.
Ključne besede: paleontologija, rodentia, insectivora, reptilia, pleistocen, Šandalja, Hrvaška.
Abstract	UDC: 568/569(119.1)(497.5)
551.791:568/569(497.5)
Jean-Pierre Aguilar & Jean-Yves Crochet & Jacques Michaux & Andrej Mihevc & Maja Paunovič: The small vertebrate fauna (Rodents, Insectivores, and Reptiles) of Šandalja 1A (Istria, Croatia)
Is described under the name of Šandalja 1A, a Lower to Middle Pleistocene fauna of small vertebrates including rodents, insectivores and reptiles. Extracted from a bone breccia found in 1999 in the Šandalja quarry near Pula, its accurate localization with respect to the previously known bone breccia of Šandalja 1 is not known. Nevertheless this dating - a Biharian age - is congruent with the younger age now advocated for the fauna of large mammals of Šandalja 1 and its associated chopper.
Key words: Palaeontology, Rodentia, Insectivora, Reptilia, Pleistocene, Šandalja, Croatia.
INTRODUCTION
In the quarry of Sandalja, located 4 km from the town of Pula (Istria, Croatia), a bone breccia was discovered in 1961. Characterized by a rich fauna of large mammals, a chopper was also found, and the assemblage received a Middle or Late Villafranchian age (Malez, 1968, 1975; Paunovic 1984). In 1981 Lumley compared the Sandalja 1 with Vallonnet because of the similarity of the Sandalja's 1 chopper and industry from Vallonet, as well as of the faunal assemblages, placing both sites in the Günz and Günz/Mindel period dated from 1 500 000 to 700 000 years B.P. In 1992 Malez et al. described the equid sample from Sandalja 1 which consists of only two distal metapodia, astragalus and calcaneum, one phalanx and a magnum as Equus sp. and compared the remains also with Middle Pleistocene equids from Mosbach and Tiraspol finding no big differences. The faunal assemblage of the Sandalja 1 most probably belongs to the Late Early Pleistocene or perhaps to the Middle Pleistocene ie. Biharian similarly to the other faunal assemblages from the bonne breccias found in Croatia. Today, this fossilliferous locality is compared to the Vallonet in France (Paunovic, 2000) and an age of 1 My (Tautavel Conference, 2000) is thus proposed.
In October 1999, in the same quarry, new blocks of a bone breccia similar in texture to the one described by Malez (1975) were sampled but the relative location of the old and new site remains vague. The previously found fauna and site was named Sandalja 1, and the one discovered in 1999, Sandalja 1A. It has been possible to collect small vertebrates (rodents, insectivores and reptiles) from the new breccia. The rodents of Sandalja 1A indicate a Lower to Middle Pleistocene age. A more intensive prospecting of the quarry as well as a revision of the fauna of Sandalja 1 will make it possible to know if these two assemblages are identical or slightly different in age.
1 - FAUNA OF SANDALJA 1A
A - Rodents
Measurements are expressed in mm Apodemus sylvaticus (Fig. 1a to 1d)
Material: five incomplete mandibles, four maxillaries including two complete ones, and several isolated teeth, SA 1A n°67 to 133
Measurements
	n	L min	L moy	L max	L min	L moy	L max
m1	19	1,75	1,88	2,01	1,05	1,12	1,24
m2	18	1,19	1,28	1,40	1,03	1,10	1,23
m3	1		1,06			0,93	
M1	18	1,75	1,94	2,15	1,19	1,27	1,37
M2	14	1,21	1,32	1,46	1,12	1,17	1,23
M3	2	0,86		0,91	0,84		0,89
These molars have all the characteristics of the species Apodemus sylvaticus. Many Pleistocene populations have been decribed by Pasquier (1974). The average size of these molars is larger than that of the Croatian population from Podumci 1 (Malez & Rabeder, 1984) and French populations of the end of the Middle Pleistocene (Pasquier, 1974). On the other hand, the size is similar to the one of the Slovenian locality of Črni kal 2 and 3 allocated to the Middle Pleistocene (Aguilar et al. 1998).
Fig. 1: Rodents of [andalja IA - Apodemus sylvaticus: 1a - m1 d. SA 1A n°111 ; 1b - M1 s. SA 1A n° 76; 1c - M2 s. SA 1A n° 92; 1d - M2 d. SA 1A n° 100 - Apodemus cf. microps: 1e - m1 s. SA 1A n° 66 - Allocricetus bursae: 1f - M2 d. SA 1A n° 63; 1g - M3 s. SA 1A n° 65 - Glis sp.: 1h - M1 d. SA 1A n°36; 1i - m1 d. SA 1A n° 20 - Clethrionomys cf. glareolus: M3 SA 1A n° 134 - Arvicolid cf. Microtus: m3 SA 1A n° 135
Sl. 1: Glodalci iz [andalje 1A - Rodents of [andalja IA - Apodemus sylvaticus: 1a - m1 d. SA 1A n°111; 1b - M1 s. SA 1A n° 76; 1c - M2 s. SA 1A n° 92; 1d - M2 d. SA 1A n° 100 - Apodemus cf. microps: 1e - m1 s. SA 1A n° 66 - Allocricetus bursae: 1f - M2 d. SA 1A n° 63; 1g - M3 s. SA 1A n° 65 - Glis sp.: 1h - M1 d. SA 1A n°36; 1i - m1 d. SA 1A n° 20 - Clethrionomys cf. glareolus: M3 SA 1A n° 134 - Arvicolid cf. Microtus: m3 SA 1A n° 135
Apodemus cf. microps (Fig. 1e)
Material : 1 m1 (1,66 x 0,96) SA 1A n° 66
This molar differs from those of A. sylvaticus, by its small size and it is characterized by a reduced outer cingular margin. It is referred to A. microps which has been recognized in Kamik (Kowalski, 1960).
Allocricetus bursae (Fig. 1f-g)
Material : 1 mandibule with m1-m3 (2,00 x 1,16; 1,54 x 1,32; 1,56 x 1,18); 2 m2 (1,58 x 1,29; 1,54 x 1,25); 1 m3 (1,58 x 1,13) ; 3 M2 (1,57 x 1,29; 1,54 x 1,35; 1,50 x 1,25); 2 M3 (1,23 x 1,07; 1,29 x 1,11). SA 1A n° 57 to 65
The size of these molars is a little smaller than the size observed in A. ehiki. It enters the size variation known for populations of A. bursae and A. croaticus of Pleistocenes age described from Croatia : Tatinja draga and Razvodje (Paunovi} & Rabeder, 1996). The two M3 seem less reduced in size than those of A. croaticus and a small mesoloph is still present on the M2. These characteristics would bring this small population close to the species Allocricetus bursae. This species was formerly recognized in the locality of [andalja 1 (Malez, 1975).
Clethrionomys cf. glareolus (Fig. 1j)
Material : 1 M3 (1,74 x 0,88) SA 1A n° 134
This rooted tooth shows the typical round outline of the salient lingual angles and backward curved buccal synclines.
Arvicolid cf. Microtus (Fig. 1k)
Material : 1 m3 (1,45 x 0,78) SA 1A n° 135
This unique molar devoid of roots is of small size, and shows the characteristically confluent T1 and T2 triangles.
Glis sp. (Fig. 1h-i)
Material : isolated teeth (see table below) SA 1A n° 20 to 56
	n	L min	L moy	L max	L min	L moy	L max
d4	2	1,21		1,24	1,03		1,05
p4	2	1,32		1,37	1,32		1,35
m1	4	1,88	1,98	2,06	1,71	1,79	1,87
m2	6	2,02	2,07	2,12	1,88	1,93	2,01
m3	4	2,01	2,09	2,20	1,66		1,92
D4	1		1,11			1,33	
P4	1		1,42			1,62	
M1	5	1,84	1,87	1,91	1,89	1,95	2,03
M2	9	1,83	1,93	2,03	2,05	2,13	2,22
M3	3	1,50	1,62	1,74	1,80	1,84	1,89
Morphology and size of these teeth are comparable with those known for the much larger assemblage described from the Slovenian site of Velika Pirešica (in prep).
B - Insectivores:
The remains of insectivores extracted from Šandalja 1A are scarce but their interest for the dating of the fauna is significant. They are attributed to 4 taxa. Teeth measurements are taken according to Rümke (1985, p. 25, Fig. 4) for Talpidae and to Reumer (1984, p. 7, Fig. 4) for Soricidae. Measurements are given in millimeter. Morphological terms are borrowed from these two authors.
Talpidae
Genus Talpa
Talpa fossilis or europaea
The determination of this form in Šandalja 1A is based on the discovery of two isolated teeth: 1 m2 dext. (2,53 X 2,03) and 1 m3 dext. (1,98 X 1,16).
The species T. fossilis Petenyi, 1864 and T. europaea Linne, 1758 are known to be difficult to separate according to tooth morphology. Some authors assert the synonymy of the two taxa whereas others regard them as good chronological index species. In this case, T fossilis is used for the lower Pleistocene time interval (Crochet & Michaux, 1981, p.135).
The main interest of the discovery of a mole is environmental as Talpa indicates rather soft but not too clayey soils, and open space.
Soricidae
Subfamily Crocidurinae
Crocidurina kornfeldi Kormos, 1934
Material : 1 right P4, 1 right M2 (highly worn; with a broken hypocone) and 1 left well preserved m2 (1,55 x 0,92).
The P4 has a broken parastyle, no parastylar point, and a weakly individualized protocone. The lack of hypocone is to be noticed. Cingulums are reduced and the posterior notch is not very deep but the latter character is variable among populations (Reumer o.c., p.21, pl. 1).
Referring to Reumer's recent critical revision of the group (o.c.), the three specimens are attributed to C kornfeldi. According to Reumer C kornfeldi is confined in the Upper Villanyan (Kislang phase) and Lower Biharian (Betfia phase) in spite of its recognition in a few older localities.
Subfamily Soricinae
Drepanosorex Kretzoi, 1941
Drepanosorex sp.
A fragmentary right maxillary (with the two last unicuspid teeth and the P4 which lacks the postero-labial part) can be referred to Drepanosorex Kretzoi 1941 because of the coloured tips of the teeth (exodaenodonty). But the size of the Šandalja 2 specimen (P4: (1,31) X 1,24; A3 0,5 X 0,57; A2 : 0,5 X 0,61) is too weak for an attribution to any species referred to this genus.
Besides some isolated mentions in a few Pliocene localities, the genus is well represented in Villanyan and Biharian faunas.
Soricid indet.
One highly worn right m2 (1,62 X 0,95) cannot be referred to above mentioned taxon. As it is not possible to see if the tip of the cones are coloured because of the wear, the tooth is determined as Soricidae indet.
C - Reptiles:
Among the microvertebrate remains, only 16 pieces of reptiles can be determined (table 1): they belong to lizards (Ophisaurus pannonicus, Archaeolacerta and Lacerta) and snakes (Elaphe).
Inv. Nr.	SPECIES	DESCRIPTION
Sa 1A-1	Archaeolacerta sp.	dermal plate, dorsal
Sa 1A-2	Archaeolacerta sp.	dermal plate, dorsal
Sa 1A-3	Archaeolacerta sp.	dermal plate
Sa 1A-4	Ophisaurus pannonicus	dermal plate
Sa 1A-5	Lacerta sp.	dermal plate
Sa 1A-6	Serpentes indent.	vertebra
Sa 1A-7	Lacerta sp.	dentary
Sa 1A-8	Lacerta sp.	dentary
Sa 1A-9	Lacerta sp.	dermal plate
Sa 1A-10	Lacerta sp.	dermal plate
Sa 1A-11	Elaphe sp.	vertebra
Sa 1A-12	Elaphe sp.	vertebra
Sa 1A-13	Serpentes indent.	vertebra
Sa 1A-14	Serpentes indent.	vertebra
Sa 1A-15	Serpentes indent.	vertebra
Sa 1A-16	Elaphe sp.	vertebra
These remains of reptiles testify to the existence of a karstic environment with mixed vegetation (forest and maquis).
AGE OF THE MICROFAUNA OF ŠANDALJA 1A
The weak representation of Arvicolids does not allow an accurate dating. However the presence the Clethrionomys with an evolved tooth morphology the rootless vole of the genus Microtus indicate the Biharian, a time interval ranging between -1,6 and -0,6 My according to the calibration suggested by Nadachowski (1990). The estimated age on the basis of these rodent is more recent than the classical localities of the "Massif Central" in France which are referred to the mammalian zone MN 16/17 (that is they are Villanyan) (de Bruijn et al. 1992; Torre et al. 1992), their ages are close to -2,5 My.
The remainding rodents are not usefull for a dating. A. bursae is known from the end of the Villanian up to the Biharian (Nadachowski, 1990). The fieldmouse species Apodemus sylvaticus, A. flavicolis and A. microps are known in Poland in Lower Biharian (Nadachowski, 1989, 1990) localities, but also in of Croatia (Malez & Rabeder, 1984; Paunovic & Rabeder, 1996) as well as in Slovenia (Aguilar et al. 1998). The dormouse Glis sp. is only known up to now in Slovenian sites (Aguilar et al. 1998). The insectivores recognized in Šandalja 1A are known in the Villanyan and Biharian.
The composition of the rodent fauna of Šandalja 1A is similar to the ones of the Slovenian localities of Črni kal 2 and 3 which are referred to the Middle Pleistocene (Aguilar et al, 1998). Although the breccia of Šandalja 1 and Šandalja 1A are very similar (great amount of pieces of bones of large mammals) it is difficult to say that they are contemporaneous because they exact origin is no more known. Nevertheless, it is worth to point that the two species Allocricetus bursae and Mimomys sp. recognized in Šandalja 1 do not contradict the age advocated for the fauna of Šandalja 1A. Preparation of newly collected blocks of breccia in Šandalja 1A at the Institute of Quaternary Paleontology and Geology in Zagreb as well as a revision of the fauna of Šandalja 1 will undoubtedly bring new data to solve this question, more fossils of voles are urgently needed. If the Šandalja I fauna is effectively younger than estimated by Malez (1968), the two faunas of Šandalja I and IA may be contemporaneous and thus the chopper would be Biharian in age.
Publication n° 2000-xxx ISEM, UMR 5554
ACKNOWLEDGMENTS
We would thank the Slovenian Ministry for Science and Technology for its financial assistance, the Management of the Valtura penitentiary institution in Pula who allowed us the access to the [andalja locality, Mr. Claude Requirand and Frank Senegas who helped us on the field.
REFERENCES
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FAVNA MALIH VRETENČARJEV (GLODALCI, ŽUŽKOJEDI IN PLAZILCI) ŠANDALJE 1A (ISTRA, HRVAŠKA)
Povzetek
V	jami v kamnolomu Šandalja pri Puli je bila leta 1961 odkrita kostna breča s kostmi velikih sesalcev. Najden je bil tudi prodnjak (Malez, 1968, 1975; Paunovic, 1984). Danes Šandaljo primerjajo z nahajališčem Vallonet v Franciji (Paunovic, 2000), njegova starost pa naj bi bila okrog 1 Ma.
V	oktobru 1999 so bili v istem kamnolomu najdeni bloki kostne breče s podobno teksturo, kot jo je opisal Malez (1975), vendar pa nahajališče blokov ni natančno znano, zato oznaka Šandalja 1A. V breči smo lahko zbrali drobne vretenčarje (glodalce, žužkojede in plazilce). Glodalci iz teh breč kažejo spodnjo do srednje pleistocensko starost breče.
Slaba zastopanost arvicolidov ne omogoča natančnega datiranja, vendar pa Clethrionomys z razvito zobno morfologijo in voluharica vrste Microtus kaže na Biharijsko starost oziroma časovni interval med -1,6 Ma in 0,6 Ma.
Ostanki glodalcev niso primerni za datiranje. A. bursae je znan za Villanian in Biharian (Nadachowski, 1990). Vrste poljskih miši Apodemus sylvaticus, A. flavicolis in A. microps so znane na Poljskem že v spodnje Biharianskih nahajališčih (Nadachowski, 1989, 1990) na Hrvaškem (Malez & Rabeder, 1984; Paunovic & Rabeder, 1996) in v Sloveniji (Aguilar et al. 1998). Polhi, Glis sp. so do sedaj znani le iz slovenskih nahajališč (Aguilar et al. 1998). Sestava favne glodalcev v Šandalja 1A je podobna nahajališčema Črni kal 1, Črni kal 2 in Črni kal 3, ki so srednje pleistocenske starosti (Aguilar et al, 1998).
Čeprav sta si kostni breči Šandalje 1 in Šandalje 1A zelo podobni (velik del kosov kosti velikih sesalcev) je težko reči, ali sta sočasni, saj natančne lokacije breč niso več znane. Kljub temu pa je vredno poudariti, da vrsti, Allocricetus bursae in Mimomys sp. iz Šandalje 1, ne nasprotujeta starosti, ki jo kaže favna iz Šandalje 1A.
Preparacija novih blokov kostne breče iz lokacije Šandalja 1A na Inštitutu za kvartarno paleontologijo in geologijo v Zagrebu kot tudi revizija favne iz Šandalje 1 bo nedvomno pomembno prispevala k natančnejšemu določanju starosti favne, predvsem s pomočjo voluharic. Če je favna Šandalja 1 mlajša kot predpostavlja Malez (1968), sta favni Šandalje 1 in Šandalje 1A lahko sodobni in je prodnjak Biharijski po starosti.