J . van der Made Museo Nacional de Ciencias Naturales, M a d r i d

B i oge og r ap hy and stratigrap hy of the Mio-Pleistocene mammals of Sardinia and the description of some fo s s i l s

Made, J. van der, 1999 - Biogeography and stratigraphy of the Mio-Pleistocene mammals of Sardinia and the description of some fossils - in: Reumer, J.W. F. & De Vos, J. (eds.) - EL E P H A N T S H AV E A S N O R- K E L! PA P E R S I N H O N O U R O F PA U L Y. SO N D A A R - DEINSEA 7: 337-360 [ISSN 0923-9308]. Published 10 December 1999

The Mio-Plio-Pleistocene faunal succession of Sardinia is discussed in the context of paleogeography and stratigraphy. Some material of a particular stratigraphic or taxonomic interest is described. T h e name Sus sondaari is introduced to replace the preoccupied name Sus nanus. Sardinia may have be- come an island during the Late Oligocene. Some of the Early Miocene taxa on Sardinia may have sur- vived there from the time when Sardinia was still connected with the main land, others arrived later to the island. Part of these taxa survived till the Late Miocene, when Sardinia (and Corsica) became con- nected to the island of Tu s c a n y. At this moment faunal exchange occurred between these areas. Some of the taxa known from the Baccinello area may have come from Sardinia. During the Messinian, Sardinia became connected to the mainland again, the endemic fauna went largely or completely extinct and a balanced fauna entered. During the earliest Pliocene Sardinia (and Corsica) became isolated again and an endemic fauna was formed. Possibly only once, at the beginning of the Middle Pleistocene, new elements were introduced. Twice Sardinia changed from mainland to insular conditions. Both times an endemic fauna with reduced diversity evolved from a mainland fauna. This occurred also in other islands. A possible explanation for such a process is that the carrying capacity of an island is insuff i- cient for viable populations of large carnivores. The larger the island, the larger the carnivore that can survive on it. The dog C y n o t h e r i u m could survive on Sardinia. The absence of large carnivores may lead to increased competition between herbivores, resulting in reduction in species abundance. Only when large carnivores are lacking, certain typical island adaptations, such as low gear locomotion, evol- ve.

Correspondence: J. van der Made, Museo Nacional de Ciencias Naturales, c. José Gutiérrez Abascal 2, 28006 Madrid, Spain, e-mail [email protected]

Keywords: Sardinia, Capo Figari, Capo Mannu, Mandriola, Messinian Salinity Crisis, Miocene, Pliocene, Pleistocene

I N T RO D U C T I O N Already during the earlier part of the past ted fossils from Capo Figari. This collection c e n t u r y, fossil mammals were collected from is stored in the Naturhistorisches Museum Sardinia. Collecting continued and, particu- Basel (NMB) and the Istituto di Geologia larly in the last thirty years, a better view of Firenze (IGF). It is not known whether they the Miocene and Pliocene was obtained. A come from a single fissure or from several, locality of particular interest is Capo Figari. nor whether this, or these fissures are the At the end of the 19th century and/or the ones we know now. In the first half of the beginning of the 20th, Forsyth Major collec- 20th century, collections were made at Capo

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Figari by D. Bate (stored in the Natural (NHML), a team from Utrecht (IVAU) (De History Museum, London - NHML) and E.G. Bruijn & Rümke 1973) made a more extensi- Dehaut (stored in Turin) and in 1970 a team ve collection. Both collections are possibly from the University of Rome collected from not from the same locality, but the fossilisa- probably the same fissure as Dehaut (Gliozzi tion suggests that they are from the same & Malatesta 1980). Up to then, all material beds. Upper Miocene remains were only from Capo Figari seems to have been consi- recently described (Cordy & Ginesu 1994). dered as of the same age, though estimates of this age ranged from Pliocene to Middle Although I never excavated with Paul at P l e i s t o c e n e . Corbeddu, I have followed the research on Sardinia from near. Since I studied the little A team of the University of Utrecht, lead by pig from Capo Figari (Van der Made 1988), I Paul Sondaar, collected in 1977 from several have a particular interest in the older part of d i ffe rent localities at Capo Figari. Two of the the history of Sardinia. I planned to write a localities are fissure fillings at the top of the paper on the subject, but this remained a plan hill near the signal station. They are named for many years. Kotsakis discussed the taxo- Capo Figari I and Capo Figari II. There are n o m y, stratigraphy and biogeography of some more localities near the sea. The collec- Sardinian mammals in many papers (Esu & tion from Capo Figari I includes remains of Kotsakis 1983; Kotsakis 1980, 1984a and b; the insectivores Nesiotites corsicanus a n d Kotsakis & Palombo 1979). The description Talpidae indet., the bat Chiroptera indet., the and discussion of some inedited material lagomorph P rolagus figaro?, the rodents from Capo Figari, Capo Mannu and Oschiri, Rhagamys minor and Ty r rhenoglis and some original ideas on the ‘Baccinello f i g a r i e n s i s, and the bovid Nesogoral melonii, faunas’, also found at Sardinia, results in a whereas Capo Figari II yielded Ta l p i d a e new interpretation of the Sardinian stratigra- indet., Chiroptera indet., P rolagus sard u s , phy and biogeography. There are many cita- R h a g a m y s cf. o rt h o d o n , Ty r rhenoglis tions of aberrant species from the localities. f i g a r i e n s i s, Ty r rh e n i c o l a sp. (primitive), the These include recent or Holocene remains latter genus being an arvicolid rodent. that later workers supposed to have been Possibly the first publication of diff e r e n t mixed with the Pleistocene fossils, or remains faunal lists for the different localities are the of fossil mainland species of micro mammals, lists of the glirids by Zammit Maempel & De that are supposed to have been brought to the Bruijn (1982). This and the excavations at island in the stomachs of birds of prey. Such Corbeddu gave rise to the recognition of two citations are not discussed here. faunal units: the Middle and Late Pleistocene (and even Holocene) ‘Ty r rh e n i c o l a f a u n a ’ M e a s u rements and their abb rev i a t i o n s and the earlier ‘N e s o g o r a l f a u n a ’ ( S o n d a a r Measurements after Van der Made (1989, 1986; Sondaar et al. 1984, 1986, 1988). 1 9 9 6 ) : D A P = Antero posterior diameter; Late Pliocene remains from Nuraghe Su DAPd = DAP of the distal part of a bone; Casteddu were first described by Esu & DAPp = DAP of the proximal part of a bone; Kotsakis (1979). Early Pliocene remains were D Ta = Transverse diameter of the anterior lobe of a described by Pecorini et al. (1973) from the t o o t h ; locality of Mandriola at Capo Mannu. A t e a m DTd = Transverse diameter of the distal part of a bone; from the University of Utrecht collected also DTp = Transverse diameter of the posterior lobe of a from Capo Mannu. Though the first few fos- tooth or proximal part of a bone; sils from the lower Miocene of Oschiri were DTpf = Transverse diameter of the proximal facet of a collected at the beginning of the 20th century b o n e ;

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Hla = Height of the crown of a tooth measured at the (1980) believed that the ancestor of N e s i o t i t e s labial side; entered Crosica and Sardinia in the latest Hli = Height of the crown of a tooth measured at the Miocene and suggested that the shrews from lingual side; the Baleares and those from Corsica and Lint = Internal or medial length of the astragalus; Sardinia might belong to different lineages. Lext = External or lateral length of the astragalus; N. corsicanus is cited from Capo Figari I Lm = Minimal or median length of the astragalus. (Zammit Maempel & De Bruijn 1982). T h e latter species is cited from a number of younger THE TA X A localities: Cava Albastro, Cava Grande and The distribution of the taxa is indicated in Santa Lucia in the Is Oreris area (Gliozzi e t Figure 1. Tooth nomenclature used in this a l . 1984), Monte San Giovanni (Bate 1945b), section is after Van der Made (1996). Dragonara (Malatesta 1970) and Corbeddu (Sondaar et al. 1988). I n s e c t i vo r a The available data make it likely that the E r inacei dae i n det. shrews arrived during the Messinian to Pecorini et al. (1973) mentioned an indeter- Corsica, Sardinia and the Baleares. From the minate genus of erinaceid from the locality of Pliocene onwards, the Corso-Sardinian and Mandriola at Capo Mannu that did not seem Balearic populations became isolated from to be known from Europe. The material was the mainland population and from each other. not described. Both insular populations independently beca- me larg e r. The evolution in both lineages was Crocidosorex antiquus (PO M E L , 1853) parallel. The genus N e s i o t i t e s as applied at De Bruijn & Rümke (1973) described present is polyphyletic. I am not in favour of material assigned to this species from the use of more names than necessary (Va n Oschiri. This soricid is known from the main- der Made 1988); if the ancestor of an island land of Europe from the localities Saulcet, species or lineage is known, the classification Chavroches and Montaigu-le-Blin. in a different genus obscures relationships. I propose to place the Corso-Sardinian species A s o r i c u l u s (= E p i s o r i c u l u s) a ff. g i b b e r o d o n in the genus A s o r i c u l u s. ( P E T É N Y I , 1864) Asoriculus corsicanus (BAT E, 1945) Geotrypus oschiriensis RÜ M K E, 1973 Asoriculus similis (HE N S E L , 1855) The species was described on the basis of Hensel based the species S o rex similis o n material from Oschiri. Other species of the material from Monreale de Bonaria near same genus are known from a number of Cagliari. Bate (1945b) named the genus Eocene, Oligocene and Early Miocene N e s i o t i t e s for shrews from the Baleares, European localities. Corsica and Sardinia, indicated N. hidalgo from Majorca as type species and described Nuragha schreuderae RÜ M K E, 1973 Nesiotites corsicanus from the Teppa di Rümke (1973) named this species and genus Lupino cave in Corsica. E p i s o r i c u l u s is of talpid on the basis of material from considered to be the ancestor of N e s i o t i t e s Oschiri. The species could not be related to (Reumer 1980 a, b). Esu & Kotsakis (1979) any of the known forms. described E p i s o r i c u l u s a ff. g i b b e ro d o n f r o m the late Pliocene locality of Nuraghe Su Talpidae indet. / Ta l p a s p . Casteddu (the species name g i b b e ro d o n i s Talpa tyrrhenica BAT E, 1945 currently placed in the genus A s o r i c u l u s Bate (1945a) based the species Talpa tyrrh e - KR E T Z O I , 1959; see Hutterer 1994). Kotsakis n i c a on material from Monte San Giovanni.

339 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

Figure 1 Range chart for the Neogene and Pleistocene endemic mammals from Sardinia.The localities are indicated from old (left) to young (ri g h t ) . Ve r tical lines indicate faunal exchange with mainland or Tu s c a ny.The position relative to the early Middle Pleistocene event is unknown for the possibly mixed collections of Major (FM), Dehaut (De) and Thaler (Th). Question mark s , ‘ c f .’ and ‘ a f f .’ indicate that the genus is present in that locality, but that the specific assignation is insecure. Lineages are indicated as a grey rectangle, the actual presence is indicated in the common way in bl a c k . For Macaca majori, a hypothetical early entry is indicated (grey and question mark s ) .

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The species is also known from Cava A l b a P r i m a t e s (Gliozzi et al. 1984) and Dragonara Oreopithecus c f . b a m b o l l i i (Malatesta 1970). Material from Capo Mannu Cordy & Ginesu (1994) and Kotsakis et al. was assigned to Talpidae indet. (non (1997) reported the primate O reopithecus c f . Desmaninae) (Pecorini et al. 1973), from b a m b o l l i i from Fiume Santo. O. bambollii Capo Figari I & II to Talpidae indet. (pers. was known from the endemic island faunas comm. of P. Y. Sondaar to Van der Made, from Tuscany and has been placed in the 1988) and from Nuraghe Su Casteddu to Cercopithecoidea and in the Hominoidea. Ta l p a sp. (Esu & Kotsakis 1979). It is possi- C u r r e n t l y, the latter position is the more ble that these indeterminate remains belong accepted and the genus is supposed to be a to the T. tyrrh e n i c a lineage, which thus descendant of D ry o p i t h e c u s (Moyà-Solà & entered in Corso-Sardinia during the Köhler 1997). O re o p i t h e c u s is known from M e s s i n i a n . the Baccinello V1 and V2 faunas (Rook 1993). These faunas are correlated to MN 11 C h i ro p t e r a and MN 12or MN 13 respectively (Engesser 1989), though evidence of the suid in the V 2 C h i roptera indet. / Ve s p e r tilionidae indet. fauna suggests that this level might be as old Rhinolophus ferrumequinum (SC H R E B E R , as late MN 11 or 8.0-8.5 My (see discussion 1 7 7 4 ) on H. etru s c u s below); the V1 fauna being Rhinolophus hipposideros (BE C H S T E I N, 1800) still older. The last record of D ry o p i t h e c u s i s Myotis myotis (BO R K H A U S E N, 1979) from the lower part of MN 10 (Andrews e t Myotis capaccinii (BO N A PA RT E , 1837) a l . 1996). It seems likely that O re o p i t h e c u s Nyctalus cf. lasiopterus (SC H R E B E R , 1780) arrived at the islands, during a sea level low Miniopterus schreibersi (KU H L, 1819) stand. There are two sea level low stands, Material from Capo Mannu was assigned to that are of interest here, one at 11.5 My and Vespertilionidae (de type myotodonte) another 8.2 My ago (Haq et al. 1987, Miller (Pecorini et al. 1973), from Capo Figari I & et al. 1996). The one 8.2 My ago is later than II to Chiroptera indet. (pers. comm. of P. Y. the extinction of D ry o p i t h e c u s . This would Sondaar to Van der Made, 1988) and from mean a very early presence of O re o p i t h e c u s Nuraghe Su Casteddu to Chiroptera indet. either in the islands of Tuscany or Corso- (Esu & Kotsakis 1979). Myotis nattere r i w a s Sardinia, or in both. cited from Dragonara (Malatesta 1970), but O re o p i t h e c u s was recently interpreted to be Kotsakis (1987) described M. myotis; it is not terrestrial, bipedal and shuffling with short clear to me whether this is based on the same stride length and low speed (Köhler & Moyà- material, though this seems likely. Solà 1997). It was supposed to have evolved Rhinolophus ferru m e q u i n u m a n d this behaviour because of the absence of car- Vespertilionidae indet. were cited from nivores in the insular environment where it Corbeddu (Sondaar et al. 1988, Kotsakis lived. The large carnivore H y a e n a rc t o s ? 1987). Rhinolophus ferru m e q u i n u m, a n t h r a c i t e s first described from Monte Bamboli Rhinolophus hipposidero s, Myotis capaccinii, has now been cited from Fiume Santo. Its N y c t a l u s cf. l a s i o p t e r u s and M i n i o p t e ru s presence at the first locality has been explai- s c h re i b e r s i were described from a cave ned away by doubting the stratigraphical con- between Punta Padrebellu and Omo Morto text (pers. comm. S. Moyà-Solà). This is not (Kotsakis 1987). These species are living possible anymore, after the find at Fiume mainland species that apparently were not Santo; O re o p i t h e c u s and the large carnivore hindered by narrow sea straits. were sympatric. The interpretation of O re o p i t h e c u s as slow and terrestrial due to the absence of carnivores is not satisfactory.

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H o m o s p . Figari II and thus survived into the Middle Sondaar et al. (1986) assigned human Pleistocene, presence of M a c a c a in the remains from Corbeddu to H o m o sp. and Middle Pleistocene of Sardinia does not hypothesised that man reached Sardinia exclude that the lineage entered during the during the early Middle Pleistocene. Spoor & Messinian. A detailed comparison of M . Sondaar (1986) describe some of the speci- majori with the mainland forms may shed mens in more detail and differences with light on the question. H o m o s a p i e n s were pointed out. Sondaar e t a l . (1991) interpreted the lithic industry of L ago m o r p h a Sardinia to be different from that of the main- land. Prolagus figaro LÓ P E Z, 1975 Prolagus sardus WA G N E R , 1825 Macaca majori AZ Z A R O L I , 1946 P rolagus figaro was first described from Gliozzi & Malatesta (1980) stated that Capo Figari (though the name seems to refer Dehaut based Ophthalmomegas lamarmorae to the Barber of Seville) on the basis of on primate molars and, what appeared later to material collected by Thaler (López Martínez be, a skull cap of an eagle, from Capo Figari, & Thaler 1975). It is a large species, much reason for Azzaroli (1946) to name l a rge r than P. sard u s . Wagner seems to have the species Macacus majori, based on based P. sard u s on material from Bonaria material collected by Major and stored in the near Cagliari (Tobien 1935). Material from IGF (Gliozzi & Malatesta 1980). There is Capo Mannu (López Martínez & T h a l e r also material of the macaque in the Major 1975) and from Capo Figari I (Zammit collection in the NMB. The reason that Maempel & De Bruijn 1982) was assigned to Dehaut included material that in reality P. cf. f i g a ro. P. sard u s is also present in the belongs to two species in the description of a Thaler collection from Capo Figari (suggest- new species does not invalidate that taxon. I ing that material from different levels was do not know the early literature on this mixed) and it was mentioned by Major from macaque, but the possibility exists that Capo Figari, though material in the NMB col- D e h a u t ’s taxa are available and, depending on lection is large and more likely represents P. the type, that the species name is a senior f i g a ro. P. sard u s is cited from Capo Figari II synonym of Macaca majori. (Zammit Maempel & De Bruijn 1982), Bonaria and Monte San Giovanni (Bate The species was also found at Cava A l b a s t r o , 1945a), San Giovanni in Sinis (Kotsakis where also Nesiotites similis, P ro l a g u s 1980), Dragonara (Malatesta 1970), s a rd u s, R h a g a m y s o rt h o d o n and Ty r rh e n i c o l a Corbeddu (Sondaar et al. 1986, 1988), Cava were found (Gliozzi et al. 1984). I interpreted Albastro, Cava Grande, Santa Lucia and from this as an indication that M. majori b e l o n g s a cave between Punta Padrebellu and Omo to the ‘Ty r rh e n i c o l a f a u n a ’ ( Van der Made Morto (Kotsakis 1987) and a P ro l a g u s s p . 1988), implying a Middle to Late Pleistocene from Buggerru, all in the Is Oreris area age. Kotsakis (1980) considered M. flore n t i n a (Gliozzi et al. 1984). The subspecies P. f i g a ro from the Villafranchian as the probable ance- d e p e re t i is present in the locality of stor of M. majori. Sondaar (1987) considered Perpignan (MN 14). P. f i g a ro f i g a ro seems to Macaca majori to be part of the ‘N e s o g o r a l have entered Sardinia during the Messinian. fauna’. M a c a c a first appeared in Europe in P. sard u s is no descendant of P. f i g a ro, but MN 13 (Andrews et al. 1996), that is during seems to have entered Sardinia during the the Messinian Crisis. M a c a c a may have Pleistocene (López Martínez & Thaler 1975). reached Sardinia then. In view of the fact that N e s o g o r a l is now also known from Capo

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R o d e n t i a Brandy assumed that this species gave rise on Corsica and Sardinia to the R. minor-o rt h o - Sardomys dawsonae DE BR U I J N, 1973 d o n lineage, which is characterised by size Sardomys antoniettae DE BR U I J N , 1973 increase. The type of R. minor is from the Pereddamys rayi DE BR U I J N, 1973 collections of Thaler at Capo Figari. The list These ctenodactylid genera and species were of this material (Brandy 1978) includes main- described on the basis of material from ly primitive forms, but also P rolagus sard u s. Oschiri. They were supposed to have come Some mixture seems to have occurred. I from Asia through A f r i c a . could not consult Hensel’s publication and do not know the type locality of R . o rt h o d o n . aff. Huerzlerimys turoliensis (MICHAUX, 1969) The type species of the genus R h a g a m y s Kotsakis et al. (1979) cited ‘aff. Va l e ry m y s MA J O R , 1905 is R . o rt h o d o n . The genus is t u ro l i e n s i s ’ from Fiume Santo. This is a endemic to Corsica and Sardinia. Material mainland species, currently placed in from Capo Figari I and in the Forstyh Major H u e r z e l e r i m y s , that evolved during the collection of Capo Figari was reported to be second half of MN 11 from H. vire t i, which small by Zammit Maempel & De Bruijn in turn evolved from the MN 10 H. minor (1982), who where unaware of Brandy’s ( Van Dam 1997). One of the characteristic p a p e r. The material probably represents R . changes in this lineage is size increase. m i n o r. Material from Capo Figari II was Engesser (1989) used these murids for the assigned to R . cf. o r t h o d o n and from Capo stratigraphy of the V0-2 faunas from which Figari III to R . o rt h o d o n . R. o rt h o d o n is cited he described respectively: Va l e rymys vire t i, V. from Bonaria (Bate 1945b), Monte San o re o p i t h e c i and Anthracomys majori. T h e Giovanni (Bate 1945a), Dragonara (Malatesta whole of this evolution of three successive 1970), Corbeddu (Sondaar et al. 1988), cave forms may well have occurred within MN 11 . between Punta Padrebellu and Omo Morto In that case the radiometric date from the V 2 (Kotsakis 1987) and from Cava A l b a s t r o , level and the stage of evolution of Cava Grande, Santa Lucia and Buggeru in the Hippopotamodon etru s c u s would fit very well Is Oriris area (Gliozzi et al. 1984). (see also discussion on the suid). It is diff i- cult to interpret the Fiume Santo murid. Tyrrhenicola henseli MA J O R , 1905 A primitive Ty r rh e n i c o l a is cited from Capo Apodemus mannu TH A L E R , 1973 Figari (collection Thaler; Brandy 1978) and Thaler (Pecorini et al. 1973) described the Capo Figari II (pers. comm. P. Y. Sondaar to giant form Apodemus mannu from the locali- Van der Made 1988). Ty r rh e n i c o l a h e n s e l i i s ty of Mandriola at Capo Mannu. A p o d e m u s cited from the Major collection from Capo first appeared in Europe in MN 13, which is Figari (Gliozzi & Malatesta 1980), Siniscola during the Messinian Crisis. Mandriola is levels C and E (Mezzabotta et al. 1 9 9 6 ) , believed to be just posterior to this event Bonaria (Bate 1945b), Monte San Giovanni (MN 14). (Bate 1945a), Dragonara (Malatesta 1970), Corbeddu (Sondaar et al. 1988), cave Rhagapodemus hautimagniensis ME I N & between Punta Padrebellu and Omo Morto MI C H A U X , 1970 (Kotsakis 1987), Cava Albastro, Cava Rhagamys minor BR A N D Y, 1978 Grande, Santa Lucia and Buggerru in the Is Rhagamys orthodon (HE N S E L , 1856) Oriris area (Gliozzi et al. 1984).

The murid Rhagapodemus hautimagniensis i s Mezzabotta et al. (1996) considered known from the mainland of Europe and Ty rrh e n i c o l a a subgenus of M i c ro t u s and stu- from Capo Mannu (Pecorini et al. 1 9 7 3 ) . died M. (T.) henseli from a number of

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localities and concluded that those from Capo Gliridae indet. Figari and Dragonara are the most primitive, Kotsakis et al. (1997) cited an indeterminate those from Monte San Giovanni and Bonaria glirid of large size from Fiume Santo. are intermediate and those from Corbeddu are more derived. Eliomys (Tyrrhenoglis) figariensis ZA M M I T MA E M P E L & DE BR U I J N, 1982 Peridyromys a ff . m u r i n u s (PO M E L , 1853) Eliomys (Tyrrhenoglis) majori EN G E S S E R , Microdyromys a ff . k o e n i g s w a l d i DE BR U I J N, 1 9 7 6 1 9 6 5 Engesser (1976) named the glirid species and Glis major DE BR U I J N, 1973 genus Ty r rhenoglis majori; the type material These glirids are present in Oschiri, which is is from the collections from Capo Figari by the type locality of G. major. They have M a j o r. Zammit Maempel & De Bruijn (1982) European affinities. De Bruijn & Rümke included Ty r rh e n o g l i s as a subgenus in (1973) described all material, but Daams E l i o m y s , described E. (T.) figariensis f r o m (1981) placed the material that originally was type locality Capo Figari I as well as from assigned to M y o m i m u s sp. in P e r i d y ro m y s Capo Figari II and assigned the material a ff. m u r i n u s. described by Pecorini et al. (1974) from Capo

Figure 2 C y n o t h e r i u m? sp. from Capo Figari (FM).Ty 5363 proximal fragment of a left humeru s : a anteri o r, b l a t e r al and, c p r oximal views.Ty 5362 distal fragment of a left humeru s : d d i s t a l , e p o s t e ri o r, f m e d i a l , g a n t e r i o r, and h l a t e r al views.The bar represents 2 cm.

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Figure 3 Bivariate plots of the distal (above) and proximal (below) humerus of C y n o t h e r i u m ? s p. from Capo Figari (squares) and C. s a r d o u s from Corbeddu (diamonds).

345 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

Mannu to E. (T. ) a ff. f i g a r i e n s i s . The same of the former species. There are two frag- authors assigned material from Nuraghe Su ments of a humerus (Fig. 2) in the Major col- Casteddu, described by Esu & Kotsakis lection from Capo Figari (NMB), Ty 5363 (1979), as H y p n o m y s to E . ( T. ) a ff. m a j o r i . and Ty 5362. There is no f o r a m e n e n t e p i c o - The stratigraphic distribution suggests that d y l o i d e u m, which occurs in Felidae and there were already two parallel lineages in Hyaenidae. The specimens are considerably the Pliocene. The separation into two lineages l a rge r than seven specimens from Corbeddu might be the result of the separation of the (Fig. 3). The specimens from Capo Figari populations on Corsica and Sardinia during may belong to a canid, even to C y n o t h e r i u m , high sea level stands. Engesser (1976) suppo- but the size difference is such, that it seems sed that T. majori evolved from A n t h r a c o g l i s unlikely they present Cynotherium sard o u s . of the Baccinello V1 or V2 fauna. If this is They might represent an ancestral form. A n the case, the lineage must have survived the alternative to the idea that the corso-sardinian Messinian crisis on Sardinia and/or Corsica. dogs evolved from Canis estru s c u s or a simi- A l t e r n a t i v e l y, there may have been parallel lar form, is that a form close to Canis cipio evolution; it is not common for island endi- or ‘C a n i s’ m o n t i c i n e n s i s entered Corsica and mics to survive in fully continental condi- Sardinia during the Messinian. These dogs t i o n s . were present in Turolian localities like Concud, Venta del Moro and Brisighella C a r n i vo r a (Rook 1992). Still another alternative is that it is related to N y c t e re u t s or X e n o c y o n . T h e ‘ H y a e n a r c t o s ’ a n t r a c i t e s WE I T H O F E R, 1889 first showed similarities in skull proportions Kotsakis et al. (1997) cited ‘ H y a e n a rc t o s ’ and of the second no skulls are known and a n t r a c i t e s from Fiume Santo. The species is Eisenmann (1990) did not further discuss the based on material from Monte Bamboli. two as possible ancestors; see also Kotsakis (1984) discussed the material from Eisenmann & Van der Geer 1999. Monte Bamboli, which has been attributed also to A m p h i c y o n, and concludes that it is Enhydrictis galictoides MA J O R, 1902 d i fficult to decide whether the material Algarolutra majori (MA L AT E S TA, 1978) belongs to I n d a rctos, Hyaenarc t o s o r Sardolutra ichnusae (MA L AT E S TA, 1977) A g r i o t h e r i u m. Two localities that can be cor- Megalenhydris barbaricina WI L L E M S E N & related to the V2 level have this large carni- MA L AT E S TA, 1987 vore, which thus seems to be typical for this Four mustelids are known from Sardinia; one l e v e l . musteline and three lutrines. The endemic musteline Enhydrictis galictoides is supposed C y n o t h e r i u m ? to have descended from Enhydrictis ard e a . Cynotherium sardous ST U D I AT I , 1857 This mainland species is known from the The species Cynotherium sard o u s was based Villafranchian localities of St. Vallier and on material from Bonaria. C. sard o u s is also Olivola (Kotsakis, 1980). Malatesta and the type species of the genus. The species is Willemsen (1986) named the genus also described from Dragonara (Malatesta, A l g a ro l u t r a with the species C y r n a o n y x 1970) and is cited from Cava Grande (Gliozzi m a j o r i as type species. This species is based et al. 1984) and Corbeddu (Sondaar et al. on material from Dragonara. Wi l l e m s e n 1988, Eisenmann 1990, Eisenmann & Va n (1992) named the genus S a rd o l u t r a. The type der Geer 1999). Kotsakis (1980) reviewed species is ‘N e s o l u t r a’ i c h n u s a e and the type literature on the dog and concluded that its of this species is an isolated skeleton found in ancestor is likely to be either Canis etru s c u s the Grotta di Nettuno in beds that are or Cuon majori, which might be a synonym probably of Weichselian age. The lutrine

346 VAN DER MADE: Sardinian Mio-Pleistocene mammals

Megalenhydris barbaricina is based on level V2 (also known under the mistaken material from the Ispiginoli cave near name of ‘M i c rostonyx choero i d e s ’). T h e D o rgali, an isolated find of unknown a ffinities of this form with the large Tu r o l i a n stratigraphic position. The genus is monospe- mainland suids of the genus M i c ro s t o n y x cific. have been clear since a long time (Pilgrim 1926, Hünermann 1969, Van der Made & P ro b o s c i d e a Moyà-Solà 1989, Van der Made 1997a). Pilgrim (1926) introduced the names Mammuthus lamarmorae (MA J O R , 1883) D i c o r y p h o c h o e ru s and M i c ro s t o n y x . Pickford The types of this species are from Fontana (1988) showed that H i p p o p o t a m o d o n has Marimenta near Gonessa. Malatesta (1954) priority over D i c o r y p h o c h o e r u s. It is becom- described a molar from Tramariglio. T h e s e ing increasingly clear that H i p p o p o t a m o d o n remains indicate a dwarf form close to and M i c ro s t o n y x form a closely related group Mammuthus meridionalis. M. lamarmorae i s of suids, that all could conveniently be placed also cited from San Giovanni in Sinis in H i p p o p o t a m o d o n (see classification by Va n (Kotsakis 1980). der Made 1997b) and in reality there is little reason for recognising a monospecific genus Pe r i s s o d a c t y l a ? E u m a i o c h o e ru s.

Some scraps of enamel from Oschiri (IVA U ) H. etru s c u s is known from Monte Bamboli might represent a perissodactyl. If this would and from the Baccinello V2 level, but not be the case, this is probably the only perisso- from the V1 level. It seems to have evolved dactyl known from an endemic island fauna. from an early stage of evolution of the H . Perissodactyls are no good swimmers and e r ymanthius bre v i d e n s (MN 11) - H. e. therefore they do not form part of the typical e r y m a n t h i u s (MN 12) lineage, suggesting that island faunas. It is possible that this animal it entered in MN 11 or early in MN 12 in was on Sardinia when it separated from south- Tuscany (Van der Made 1997a). Considering ern France during the Oligocene and then the ages for MN 11 (Van Dam 1997), this is survived some time on the island. in line with the radiometric date of 8.0-8.5 My for the V2 level (Hürzeler 1982), but A r t i o d a c t y l a older than the MN 12/13 age suggested by Engesser (1989). Haq et al. (1987) reported a Suidae sp. sea level low stand 8.2 My ago. Even though Some remains from Oschiri (IVAU) represent islands remain isolated to some extend, such a suid and indicate shortening of the legs, an moments tend to be the moments when new adaptation common in endemic island elements are introduced in the endemic envi- environment. The remains clearly belong to a ronment. If H. etru s c u s was really present on suid and not to a palaeochoerid. The only Sardinia, it is likely to have entered 8.2 My suid known of this age are H y o t h e r i u m ago simultaneously in Corso-Sardinia and in (MN1-6) and X e n o c h o e rus (MN 2b-3). Tu s c a n y, which then formed one large island.

Hippopotamodon etruscus (MI C H E L O T T I , Sus sondaari new name 1 8 6 1 ) ? Sus a f f. sondaari new name A suid is cited from Fiume Santo (Kotsakis e t Sus nanus was the name I gave to a small pig a l . 1997). This is probably ‘E u m a i o c h o e ru s’ . in the Major collection from Capo Figari Hürzeler (1982) introduced the name ( Van der Made 1988). The name is however, E u m a i o c h o e ru s for the species ‘M i c ro s t o n y x ’ preoccupied by Sus scrofa nanus NE H R I N G, e t ru s c u s from Monte Bamboli and Bacinello 1864, which is a domestic pig. A r e p l a c e m e n t

347 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

name should be given, and what other name ‘Amphitragulus boulangeri’ could be given on this occasion than Sus son - Comaschi Caria (1953) described a mandible daari in honour of Paul Sondaar? Holotype, from Sardara as Amphitragulus boulangeri. diagnosis etc. remain the same as published The specimen is an isolated find, but was by Van der Made (1988). The material from assumed to be of Early Miocene age. T h e Capo Mannu figured by me as S u s a ff. n a n u s description was short and taxonomically becomes of course S u s a ff. s o n d a a r i . important information was omitted. Sardinia being an island then, it seems unlikely that The little pig is a descendant of Sus arvernen - the mainland A. boulangeri was present. T h e s i s and arrived at Sardinia during the Messinian age of the ruminant from Sardara is under (Van der Made 1988). Evolutive tendencies in discussion (Esu & Kotsakis 1983): whereas it this endemic lineage are: size decrease, increase has been originally attributed to the in crown height of the first incisors, P4 and Burdigalian, it might even be of Oligocene molars, increase in enamel thickness, simplifi- age. As long as its taxonomy and age are not cation of molar structure (loss of crests or clear the find is of very limited value in bio- lobes of the cusps), reduction in size of the g e o g r a p h y. anterior premolars, loss of the p1, and shorten- ing of the anterior part of the mandible. A Megaloceros sp. peculiarity for an endemic island pig is that its Megaloceros? cazioti (DEPÉRET, 1897) third metacarpal (the only postcranial element Cervus cazioti DEPÉRET, 1897 is based on known) is not short compared to the tooth row material from Corsica, but also occurs on and is even long compared to its own DAP and Sardinia and Cervus algarensis Comaschi DT values; it is slender. Moreover, the crest on Caria, 1955 is based on remains from Il the distal articulation surface is pronounced Cantaro of Weichselian age. In addition, and the proximal articulation surface flat as in Megaceros cretensis and Orthogonoceros ver - artiodactyls that are more cursorial than the ticornis have been cited from Sardinia (Caloi majority of pigs. The dental adaptations are to & Malatesta 1974, Kotsakis 1980). Some of be expected in an island form, but possible these species have also been placed in cursorial adaptations instead of a change to Nesoleipoceros, but this genus is not common- low-gear locomotion is unexpected. ly accepted. Megaceros, Praemegaceros and Megaceroides are considered to be A n t h r a c o t h e r i i d a e synonyms of Megaloceros (Lister 1993). Most authors coincide in that the remains from Some bones from Oschiri (IVAU) are inter- Sardinia and Corsica belong to a megacerine; preted as an anthracothere. It has been sugge- if this is really the case, the name Megaloceros sted that A n t h r a c o t h e r i u m was present on should be applied. Kotsakis (1980) already Corsica and Sardinia, when these separated noted that there are size differences in the from the mainland; when the Corso-Sardinian material. Fossils in the collection of Major island became connected to the Tu s c a n from Capo Figari are larger than those from island, this form may have given rise to the Corbeddu assigned to M. cazioti. It seems like- A n t h r a c o t h e r i u m known from Tuscany (Va n ly that there was more than one deer species der Made 1999). on Sardinia, though the presence of M. creten - sis, an endemic of Crete, and M. verticornis, of B a c h i t h e r i u m ? the mainland, seems unlikely. The different Some bones from Oschiri (IVAU) might Sardinian species might, or might not form a represent a B a c h i t h e r i u m. This form may lineage. A revision of all material seems neces- have been present on Sardinia, when the sary, as already stressed by Kotsakis (1980). island became separated from the mainland. A C e rv u s sp. is cited from Bonaria (Bate

348 VAN DER MADE: Sardinian Mio-Pleistocene mammals

1945b), N e s o l e i p o c e ros cazioti f r o m mainland. It is also possible that this rumi- Dragonara (Malatesta 1970) and M e g a l o c e ro s nant gave rise to ‘U m b r i o t h e r i u m’ w h e n cazioti is described and figured from Sardinia came into contact with Tu s c a n y. Corbeddu (Klein Hofmeijer 1996). Kotsakis Kotsakis et al. (1997) cited ‘G i r a f f i d a e i n d e t . (1980) mentioned deer from Grotta I (medium size), G i r a f f i d a e indet. II (small dell'Inferno, Portovesme and Carmas and s i z e ) ’ from Fiume Santo. The larger ‘giraff i d ’ Maritza. The robusticity index (L/DTd) of the may well be identical with ‘U m b r i o t h e r i u m’ . metacarpal of M. cazioti from Corbeddu is 5.6 (average measurements from Klein Maremmia lorenzi HÜ R Z E L E R , 1983 Hofmeijer 1996). M. soleilhacus ( ‘ M. vert i - Cordy & Ginesu (1994) and Kotsakis et al. c o r n i s’) has been mentioned as a possible (1997) cited M a re m m i a l o re n z i from Fiume ancestral form to M. cazioti (Kotsakis 1980). Santo. Hürzeler & Engesser (1976) introdu- This form tends to have a robusticity index of ced these specific and generic names for the less than 5 (Van der Made in press). endemic bovid M. haupti from the V1 level E u c l a d o c e ro s , A r v e r n o c e ro s and C e rv u s t e n d in the Baccinello basin, but the names are to have indices between 6.5 and 5, D a m a - l i k e available since the publication of the diagno- deer have even more slender metapodials. sis by Hürzeler (1983), who also introduced Either M .? c a z i o t i descended from M. the name M a re m m i a l o re n z i for bovids from s o l e i l h a c u s and became more cursorial (con- the younger V2 level. The presence of trary to island evolution tendencies), or we M a re m m i a l o re n z i in Fiume Santo suggests should consider the possibility that it evolved that the locality is of about the same age as from one of the other genera. The lack of a the V2 level, that is, MN 11 or about 8.0-8.5 bez-tine seems to eliminate C e rv u s e l a p h u s My (see discussion on M. etru s c u s). as a candidate. E u c l a d o c e ros dicranios and E . t e g u l e n s i s seem to have too complex antlers. Tyrrhenotragus gracillimus (WE I T H O F E R , E u c l a d o c e ros giulii has a brow tine that 1 8 8 8 ) originates high above the burr and, though ‘Bovidae indet. I (small size)’ not well known, the antlers do not seem to be ‘Bovidae indet. II (very small size)’ very complex. The possibility of a descen- Kotsakis et al. (1997) cited the three bovids dance from D a m a-like deer, a form close to from Fiume Santo. The type locality of T. E u c l a d o c e ros tetracero s, or from E. giulii, or g r a c i l l i m u s is Monte Bamboli. Hürzeler & from A rv e r n o c e ro s merits further investiga- Engesser (1976) introduced the name t i o n . Ty rrh e n o t r a g u s, which remained a nomen nudum. Kostakis (1984) considers the name R u min an tia sp. valid because of the publication by T h o m a s ‘ G i r a ffid ae indet. I & II’ (1984). A T. a ff. g r a c i l l i m u s is cited from the A ruminant approximately the size of a fal- V1 level (Hürzeler & Engesser 1976). low deer was found at Oschiri. The animal is, Thomas (1984) interpreted Ty r rh e n o t r a g u s t o h o w e v e r, no deer and resembles in its molar be a neotragine, though he noted that the structure an animal from Casteani that has position of the horn core was posterior, been classified either as a bovid or a giraff i d , approaching the position in the Cepha- and was named Umbriotherium azzaro l i i b y lophini. This is already a derived character, Hürzeler & Engesser (1976). This name though not yet fully developed as in the however remained a nomen nudum (Kotsakis recent cephalophines. The orientation of the 1984a, b). It is well possible that the animal horn core also approaches the state in the from Oschiri is no cervid, bovid or giraff i d , Cephalophini. Work in progress suggests that but a primitive ruminant, present in Corsica the Cephalophini have an origin outside A f r i c a , and Sardinia when they separated from the not later than the Late Miocene.

349 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

Ty r rh e n o t r a g u s may well be one of the first ‘low gear locomotion’ and that is typical for c e p h a l o p h i n e s . dwarfed island species (Leinders & Sondaar 1974). Pecorini et al. (1973) mentioned N e s o g o r a l s p . ‘ C a p r i n a e ? ’ from the locality Mandriola at Nesogoral melonii (DE H A U T, 1911 ) Capo Mannu in the Capo Mannu Formation. Gliozzii & Malatesta (1980) introduced the There are some specimens of a caprine in the name N e s o g o r a l for the bovid ‘A n t i l o p e’ I VAU collection that are assigned here to m e l o n i i, a species that is based on material N e s o g o r a l sp. from Capo Figari (Dehaut collection). T h e y concluded that N e s o g o r a l and an endemic A left p2 (Fig. 4a-c; DAP 6.8, DTa 4.1, DTp bovid from the Baleares, M y o t r a g u s , are 5.4, Hli >7.3, Hla >7.1) has a pointed proto- closely related and that the ancestor of both conid and a protoprecristid that rapidly reached the islands during the Messinian becomes lower away from the tip of the pro- crisis, when the Mediterranean sea level tocone at its end it curves lingually without dropped several kilometres. A striking diff e - forming a cusp that could be called paracon- rence between the two endemic genera is that id. The protoendocristid is directed postero- M y o t r a g u s has extremely shortened metapo- l i n g u a l l y. There is a large triangular facet dials, whereas those of N e s o g o r a l are not covering most of the posterior part of the shortened. Shortened metapodials form part tooth. The size is roughly comparable to that of a set of adaptations that has been named of the homologous tooth in N. melonii a s

Figure 4 N e s o g o ra l s p. from Capo Manu . Left p2: a o c c l u s a l , b lingual and c buccal views. Right m1-2: d occlusal and e l i n- gual views. Distal fragment of right tibia: f distal and g p o s t e r ior views.The bar represents 0.5 cm for figures a-c, and 1 cm for figures d-g.

350 VAN DER MADE: Sardinian Mio-Pleistocene mammals

indicated by Gliozzi & Malatesta (1980). may have been comparable to those of the m1 Several early caprines have recently been of N. melonii as given by Gliozzi & described. Aragoral mudejar from La Roma Malatesta (1980). 2 (Spain; MN 10; Alcalá & Morales 1997, Alcalá 1994) and N o r b e r tia hellenica f r o m The central part of an u p p e r m o l a r has the Maramena (Greece; MN13/14; Köhler et al. crests arranged in a bovid manner. It seems to 1995) have much lower p2. have been rather hypsodont; besides it does not show any important characters and it can- A worn selendont right m1-2 (Figure 4d-e; not be measured. D A P 14.3, DTa 8.6, DTp 9.1) has a fairly flat lingual wall like in bovids and unlike in cer- A distal part of a r i gh t tibi a (Fig. 4f-g; vids. Like the Caprinae, it lacks a buccal sty- DAPd 18.0) has the morphology that is typi- lid. The tooth is too much worn to show a cal for ruminants (deep groves for the proxi- possible goat fold. The size is in the range for mal pulley of the astragalus, oriented in antero- the m2 of N. melonii as given by Gliozzi & posterior direction). In size it goes well with Malatesta (1980). the dental remains.

The anterior part of a left lower m o l a r ( D Ta N e s o g o r a l is a much more hypsodont animal 6.7) is much worn, but of similar morphology than A r a g o r a l and N o r b e rt i a . Unless as the previous tooth. Though the second lobe hypsodonty was acquired in insular environ- may have been wider, the size of this tooth ment, the latter genera cannot be ancestral to

Figure 5 N e s o g o ra l a f f . m e l o n i i from Capo Figari II. Right astra g a l u s : a p o s t e r i o r, b l a t e r a l , c a n t e r i o r, d m e d i a l , e distal and f p r oximal views.The bar represents 2 cm.

351 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

Figure 6 Bivariate plot of the astra g a l u s . Squares = N e s o g o ral melonii from Capo Figari oc; t r iangle pointing upwards = N e s o g o ra l? from Capo Figari II; Diamonds = ‘M eg a l o c e r o s ’ c a z i o t i from Corbeddu.

N e s o g o r a l. The affinities of the Capo Mannu material seem to be much more with The right astragalus (Fig. 5; measurements N e s o g o r a l than with the other early caprines in Table 1) has the typical ruminant morpho- and are assigned here to N e s o g o r a l sp. A p a r t logy with a distal pulley that is in line with from Capo Mannu, N e s o g o r a l has only been the proximal pulley. The specimen is much cited from Capo Figari (Major Collection, smaller than the astragali from Corbeddu Dehaut Collection) and in particular from assigned to M e g a l o c e ros cazioti (Fig. 6) and Capo Figari I (Van der Made, 1988). In it is even smaller than those of N e s o g o r a l October 1988, I visited Capo Figari together m e l o n i i from the old collections from Capo with P. Weesie and R.D. Kahlke. I took a Figari by Major stored in the NMB. The size small sample from Capo Figari I and a larg e r d i ffe rence is such that the possibility exists one from Capo Figari II. This material is tem- that it belongs to a different species. The astraga- porarily stored in the IVAU. The sample from lus, which is much smaller than the astragali Capo Figari II included Ty r rh e n i c o l a, P. from Corbeddu assigned to M e g a l o c e ro s c a z i o t i s a rd u s, possibly ‘A s o r i c u l u s’ and a ruminant (Fig. 6), is assigned here to N . a ff. m e l o n i i. astragalus. The astragalus was in the same sediment as the vole remains.

352 VAN DER MADE: Sardinian Mio-Pleistocene mammals

Ta ble 1 Measurements (in mm) of the astragalus of N e s o g o ral melonii from Capo Figari oc and N e s o g o ra l ? from Capo Figari II.

PALEOBIOGEOGRAPHY OF island, or may have got there during MP29 or SARDINIAN LAND MAMMALS MN 1. C ro c i d o s o rex antiquus and the suid may have got onto the island during MN 1-2 Late Oligocene - Early Miocene (second event), and various possibilities exist During the late Oligocene Corsica and for the rest of the fauna. Sardinia seem to have separated from Southern France (Esu & Kotsakis 1983). De Eustatic sea level was high between 35 and Bruijn & Rümke (1973) interpreted the 30 My ago and then dropped nearly instantly microfauna of Oschiri as being endemic and (geologically speaking) some 75 m and i n s u l a r. G e o t r y p u s last appears in localities remained this low, or even lower between 30 that at present are placed in MN 1 and and 25 My ago, after which it rose again for a C ro c i d o s o rex antiquus is found in localities couple of millions of years. There are several that at present are placed in MN 1 and 2 (De fluctuations in the 30-25 My interval, but sea Bruijn & Rümke 1973; De Bruijn et al. level was always 120-40 m below the level of 1992). The last citation of A n t h r a c o t h e r i u m i s 33-24 My ago (Haq et al. 1987). T h e from MP 29 (Brunet & Vianey-Liaud 1987), Corsican and Sardinian plate became the oldest European suid is from MN 1 (Va n disconnected from the main land during the der Made 1990, 1994). The fauna includes Oligocene (Dercourt et al. 1986, Rögl & ctenodactylids, which are known from A s i a Steininger 1983, Rögl 1998). Whereas this and Africa, but not from Europe, though they was a slow and unidirectional tectonic are known from the Oligocene of Mallorca process, the fluctuating sea level is likely to and the Villafranchian of Sicily (Kotsakis have had a more direct effect on the paleoge- 1984). The ctenodactylids are believed to o g r a p h y. It is thus to be expected that during have reached Sardinia during the Late the early phase of separation, Corsica and Oligocene (De Bruijn & Rümke 1973). T h e Sardinia may have been intermittently con- l a rge mammals may all be of European nected to the mainland, whereas later, when origin, though the affinities of the large they moved more southwards, the island(s) ruminant are not clear. The simplest model were intermittently closer and further away for the population of Sardinia (a single Early from the mainland. The period of 30-25 My Miocene event) becomes more complex with ago corresponds to units MP 24-29 (Legendre the incorporation of the large mammals from & Lévêque 1997). Taking this into account, Oschiri. The A n t h r a c o t h e r i u m was either on A n t r a c o t h e r i u m and G e o t r y p u s may have Sardinia when it separated from the continent been on Corsica and Sardinia when these or got there when it was already an island (at areas became insular. The ctenodactylids the same moment as the ctenodactylids?), but entered Sardinia when it was insular, because not later than MP 29. G e o t r y p u s might also they remained restricted to Sardinia. This is have been on Sardinia when it became an likely to have occurred during one of the sea

353 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

level lowstands during MP 28-29. Probably part of the Middle and/or Late Miocene or still later (MN1/2) C ro c i d o s o rex antiquus a n d may have occurred only during sea level low the suid entered Sardinia. stands. There was a sea level low stand about 11.5 My ago and another one about 8.2 My Late Miocene ago (Haq et al., 1987; Miller et al., 1996). Nothing is known of the Middle Miocene These moments are also the moments when land mammals of Corsica and Sardinia. T h e distance of the islands to the mainland is presence of O re o p i t h e c u s and M a re m m i a i n smallest and when dispersals from the main- the locality of Fiume Santo suggests that land to the islands are more likely. Ta k i n g Sardinia, Corsica and Tuscany constituted this into account, it seems likely that 11.5 My one large island during part of the Late ago O re o p i t h e c u s , Ty r rh e n o t r a g u s a n d Miocene. The locality of Fiume Santo is cor- M a re m m i a coming from the mainland related to the V2 level of the Baccinello reached Tuscany and probably Corsica and basin. Rook (1993) correlated the localities of Sardinia. At the same time, A n t h r a c o t h e r i u m Casteani, Montemassi and Ribolla with the and ‘U m b r i o t h e r i u m’, coming from Sardinia Baccinello V1 level. There is a bunodont and Corsica, may have reached Tu s c a n y. anthracothere from Casteani (Kotsakis 1984). H i p p o p o t a m o d o n reached Tu s c a n y, Corsica This form must have lived for a long time in and Sardinia 8.2 My ago, marking the onset isolation. The last bunodont anthracotheres in of the V2 fauna. Europe went extinct during the latest Oligocene. In Africa no younger bunodont Pliocene - Early Pleistocene anthracothere is known than the lower During the Messinian the Mediterranean Miocene (?) one from Malembe became disconnected from the Atlantic Ocean ( ‘P a l a e o c h o e r u s’ d a r t e v e l l e i of Hooijer and large parts of the Mediterranean became 1963). In the Indian Subcontinent the last land; Corsica, Sardinia and Tuscany became l a rge bunodont anthracothere is from the connected to the mainland (Rögl & Lower Miocene; a small bunodont Steininger 1983). Evaporation in the a n t h r a c o t h e r e possibly lived on till the Late Mediterranean was then, and still is, greater Miocene, but is an unlikely ancestor for the than the influx by precipitation and rivers. It anthracothere from Tu s c a n y. A n t h r a c o t h e r i u m is possibly the sea level low stands 6.3 and is known from the Oligocene of the North of 5.8 My ago (Haq et al. 1987), that caused the I t a l y, but this was no isolated area and isolation of the Mediterranean and triggered A n t h r a c o t h e r i u m could not survive here till what is known as the Messinian Salinity the Late Miocene. It could have been present Crisis. The beginning of the Pliocene is in Tuscany since the Oligocene, but in view marked by the re-establishment of the of the known presence of an anthracothere in connection with the Atlantic Ocean resulting Sardinia, the most likely origin for the tuscan in the flooding of the Mediterranean. T h i s anthracothere is the one from Oschiri (Va n happened about 5 My ago. der Made 1999). The ‘U m b r i o t h e r i u m’ f r o m Casteani might be a descendant of the larg e When, during the Messinan, Tu s c a n y, Corsica ruminant from Oschiri, as suggested above. If and Sardinia became connected to the main- the two artiodactyls from Casteani evolved land, these areas became accessible to main- from the Oschiri forms, there must have been land species, including carnivores. T h i s some connection between Tuscany and meant the extinction of the island faunas. Sardinia in ‘V1 time’ (or earlier). When, in the Pliocene, the Mediterranean flooded again, Tuscany remained part of the Connections between Sardinia, Corsica and mainland, but Corsica and Sardinia became Tuscany may have been permanent during islands again. The fauna living on these

354 VAN DER MADE: Sardinian Mio-Pleistocene mammals

islands became isolated. Geologically comm. D.S. Reese). speaking, species diversity dropped immediately and species acquired island M i ddle and Late Pleistocene adaptations, such as gigantism in small mam- The younger faunas are characterised by the mals and dwarfism in large mammals. T h e presence of P rolagus sard u s , which is inter- Capo Mannu fauna is from shortly after the preted as a new arrival and not a descendant isolation. Pecorini et al. (1973) suggest an of P. f i g a ro (López Martínez & Thaler 1975), age of, in terms of Neogene Mammal Units, A. similis and R. o rt h o d o n , which are inter- MN 14. S u s a ff. s o n d a a r i is already dwarfed preted as advanced evolutive stages of line- and Apodemus mannu became already larg e , ages that where present already, and showing that the fauna was already isolated. Ty rrh e n i c o l a h e n s e l i , ‘M e g a l o c e ro s’ c a z i o t i A number of animals that are not frequently and possibly C y n o t h e r i u m, which are new found in island environment is present in this arrivals. The transition to this new fauna is fauna: Erinaceidae, Talpidae and Bovidae. assumed to have occurred during a regression This is possibly due to the fact that the fauna at the beginning of the Middle Pleistocene, came walking to the islands, instead of swim- some 0.8 My ago, by Sondaar et al. ( 1 9 8 6 ) , ming. M a c a c a is possibly no good swimmer who also assumed H o m o to have arrived at e i t h e r, and may have arrived at Sardinia at this moment in Sardinia. M a m m u t h u s m a y this time. A s o r i c u l u s , E l i o m y s and the ance- have arrived at this moment in Sardinia, or stor of N e s o g o r a l are likely to have reached some 1.6-1.8 My ago, as pointed out above. both Corsica-Sardinia and the Baleares during the Messinian, giving rise to The Middle and Late Pleistocene fauna is N e s i o t i t e s, H y p n o m y s and M y o t r a g u s in the indicated by the name ‘Ty r rh e n i c o l a f a u n a ’ Baleares and to A s o r i c u l u s c o r s i c a n u s - (Sondaar et al. 1984, 1986; Sondaar 1987). s i m i l i s, Ty rrh e n o g l i s and N e s o g o r a l i n ESR dates on M. cazioti tooth enamel by Dr. Sardinia and Corsica. Intermittent connec- Motoji Ikeya (Dpt. of Earth and Space tions between the latter two islands may have Science, Osaka Univ.) are 450,000 ± 20% resulted in two lineages in Ty r rh e n o g l i s. y B P for Santa Lucia I, 366,950 ±20 % yBP Possibly stratigraphically interesting evolu- for Capo Figari II, and 15,375 ± 20 % and tion occurred in this period in the A s o r i c u l u s, 14,490 + 20 % yBP for two samples from R h a g a p o d e m u s-R h a g a m y s , Ty rrh e n o g l i s a n d Corbeddu (pers. comm. D.S. Reese). As S u s lineages. New arrivals from the mainland suggested by the find from Capo Figari II, in Sardinia might be expected during low sea N e s o g o r a l (as well as E . (Ty r rh e n o g l i s) f i g a - level stands. These occurred about 3.8, 2.9 r i e n s i s ) persisted quite some time after the and 1.6 My ago (Haq et al. 1987), allowing new arrivals. M. majori is another large mam- for small differences due to advance in mal that (if it really arrived at Sardinia during dating. There is little evidence for such arri- the Messinian), persisted after this faunal vals, though the arrival of M a m m u t h u s 1.6 or turnover event. A l t e r n a t i v e l y, M a c a c a a r r i v e d 1.8 My ago cannot be ruled out. as part of this event. There is evidence for a faunal phase that could be termed ‘N e s o g o r a l The fauna described above is the one indica- - Ty r rh e n i c o l a - fauna’, though it is of little ted by the name ‘N e s o g o r a l f a u n a ’ after its practical use, in view of the fact that the most emblematic taxon (Sondaar et al. 1 9 8 4 , older elements in this fauna seem to be rare. 1986; Sondaar 1987). The youngest locality of this ‘fauna’ is Capo Figari I. An ESR date on N e s o g o r a l tooth enamel by Dr. Motoji Ikeya (Dpt. of Earth and Space Science, Osaka Univ.) is 1,807,500 ± 20% yBP ( p e r s .

355 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

ENDEMIC ISLAND FAUNAS small populations tend to be genetically less O R I G I N ATING IN SITU F ROM A diverse and are more vulnerable (for instance an BALANCED FAU N A epidemic might wipe out the entire population). Paul Sondaar is the person who most contributed Some examples of carnivore population densi- to the model of island evolution that we are ties are (in numbers per km2): Lion 0.38-0.06, familiar with now (for instance, Sondaar 1986). P. pard u s 0.03-0.05, L. lynx 0.02-0.10, C ro c u t a Though the model is of course much more com- 0.12-0.24, U. arc t o s 0.007-0.67; C a n i s l u p u s plex, the most characteristic features are mentio- 0.002-0.04, N y c t e re u t e s 0.1-2, Cuon alpinus ned here. Proboscideans, cervids and hippos are 0.35-0.90, Vulpes vulpes 1-2, G u l o 0 . 0 0 2 - 0 . 0 0 5 , the most common endemic large mammals on Mustela erminea 10 (Nowak 1991). The higher Mediterranean islands, as well as on many other densities may well be due to artificial circum- islands elsewhere. This is because they are good stances. It can be observed that felids tend to swimmers. Large mammals become dwarfed in have low densities and that within each family, such environments and small animals become population densities increase with decreasing giants. Apart from their size, the animals have body size. other typical adaptations, that evolved parallel on many islands: ‘low gear locomotion” (short Not all islands can support a population of larg e legs, changes in posture), stereoscopic view etc. carnivores. The surface of Malta is 246 km2, of Since carnivores do not get to the islands (they Mallorca some 3600 km2 and of Sardinia some are not such good swimmers), high gear loco- 24000 km2. Malta might support about 10 pan- motion is no advantage, nor is a wide visual thers, Mallorca 144 and Sardinia 960. Sardinia field; instead the economy and stability of low might support some 4320 hyenas, some 25200 gear locomotion and stereoscopic view are raccoon dogs, some 15000 dholes or 5040 wol- advantageous. The absence of carnivores leads ves. Populations of these sizes will not all be to overpopulation, which is the cause for adap- viable. Population sizes frequently fluctuate and tations in the dentition (extreme hypsodonty, one of the reasons for that may be the interac- reduction of premolars etc.). Thus, the typical tion between predator and prey. Mathematical endemic island fauna, as on Cyprus and Crete, models and experiments in limited environ- results largely from the fact that carnivores ments may show increasing amplitude of fluctu- could not reach the island. But in the case of ation, leading to the extinction of the predator Sardinia, two times (in the Late Oligocene and (Hassel 1976). Such fluctuations seem to be in the Early Pliocene) the island started as part more common to small or poor ecosystems. A l l of the mainland, only later to become isolated. this suggests that not all islands are able to sup- It is to be expected that a balanced mainland port populations of large carnivores. The larg e r fauna became isolated on the island. So carnivo- the island, the larger the carnivore that may sur- res and all other kind of animals were present vive on it. Carrying capacity might be the rea- that are not commonly found in endemic island son that on Mallorca no large carnivore survived, faunas. Still impoverished island faunas develo- while on Sardinia the canid C y n o t h e r i u m, but ped. W h y ? no larger carnivores, nor felids could survive.

Herbivore biomass is much bigger than In the absence of carnivores, competition carnivore biomass, because carnivores need between the herbivores becomes more l a rge populations of herbivores to maintain accentuated. This may be the reason why island themselves. Therefore, the carnivore biomass faunas, starting as balanced faunas, become supported by a certain land surface is much impoverished in herbivores as well. Islands with smaller than the herbivore biomass supported. a single carnivore might be relatively rich in Carnivores need huge territories. Moreover, herbivore species. These species would then not populations need a certain size to be viable; change to ‘low gear locomotion’ and lack a

356 VAN DER MADE: Sardinian Mio-Pleistocene mammals

number of other typical island adaptations, such Teruel, Aragón, Spain) - Comptes Rendus de as stereoscopic view etc. There seem thus to l'Académie des Sciences Paris, 324, série IIa: exist two ways in which an endemic island spe- 9 4 7 - 9 5 3 cies can originate: (1) it arrives on an island and Andrews, P., Harrison, T., Delson, E., Bernor, R.L. & becomes isolated (generally by ‘sweepstake Martin, L., 1996 - Distribution and biochronology of route’), (2) a population lives on a piece of land European and southwest Asian Miocene catarrhines - that becomes an island, isolating the population. in: Bernor, R.L., Fahlbusch, V. & Mittmann, H.W. It is of interest to note that proboscideans, deer (eds.) - The Evolution of Western Eurasian Neogene and hippos are common in the first group, whe- Mammal Faunas - New York: Columbia University reas the cases of the Baleares and Sardinia sug- Press: 168-207 gest that bovids, anthracotheres, primates, hed- Azzaroli, A., 1946 - La scimmia fossile della Sardegna - gehogs, moles and possibly pigs and dogs may R i v. Sci. Preist.1: 68-76 be among the second group. Soricids, glirids Bate, D.M.A., 1945a - The Pleistocene mole from and murids seem to be present in any case and Sardinia - Annals and Magazine of Natural History perissodactyla and carnivores never or rarely 11 (12): 448-461 survive in insular environments. Cyprus, Crete Bate, D.M.A., 1945b - Pleistocene shrews from the and many of the smaller Greek islands were l a rger western Mediterranean islands - Annals and colonised by animals that used the sweepstake Magazine of Natural History 11 (11): 738-769 route. The faunal composition of the Baccinello B r a n d y, L.D., 1978 - Données nouvelles sur l'évolution faunas suggests, that the area was originally du rongeur endémique fossile corso-sarde R h a g a m y s connected to the mainland or became connected F. Major (1905) (Mammalia, Rodentia) - Bulletin de to islands with such faunas. la Société géologique de France (7), 20 (6): 831-835 Brunet, M. & Vianey-Liaud, M., 1987 - Mammalian AC K N OW L E D G E M E N T S Reference Levels MP 21-30 - Münchner In the first place I thank Paul Sondaar for intro- Geowissenschaftliche Abhandlungen, Reihe A ducing me to the interesting and bizarre world Geologie und Paläontologie, 10: 30-31 of endemic island mammals. A number of pers- De Bruijn, H., Daams, R., Daxner-Höck, G., Fahlbusch, ons gave me access to material or information V., Ginsburg, L., Mein, P., Morales, J., Heizmann, E., or helped me in any other way: H. de Bruijn, A . M a y h e w, D.F., Van der Meulen, A.J., Schmidt-Kittler, Currant, B. Engesser; J. Hooker, M. Ikeya, T. N. & Telles Antunes, M., 1992 - Report of the Kotsakis, A.J. van der Meulen, J. Morales, D.S. RCMNS working group on fossil mammals, Reese, J. Rodríguez, L. Rook, D. Torre. I am R e i s e n s b u rg 1990. Newsletters on Stratigraphy 26 receiving support from the Ministerio de (2/3): 65-11 8 Educación y Cultura, the Dirección General de De Bruijn, H. & Rümke, C.G., 1973 - On a peculiar Enseñanza Superior (project PB93-0066-C03- mammalian association from the Miocene of Oschiri 03) and the ‘Unidades A s o c i a d a s ’ program of (Sardinia) - Proceedings of the Koninklijke the DGICYT. The study of material from Nederlandse Akademie van Wetenschappen B 77 (1): Oschiri in the NHML was possible through an 4 6 - 7 9 exchange program of the Royal Society and the Comaschi Caria, I., 1953 - L'Amphitragulus boulangeri C S I C . Pomel, primo mammifero terrestre segnalato nel Miocene della Sardegna - Rivista Italiana di R E F E R E N C E S Paleontologia 59: 91-99 Alcalá Martínez, L., 1994 - Macromamíferos neógenos C o r d y, J.M. & Ginesu, S., 1994 - Fiume Santo (Sassari, de la fosa de Alfambra - Teruel - Instituto de Estudios Sardaigne, Italie): un nouveau gisement à Turolenses & Museo Nacional de Ciencias Naturales, Oréopithèque (Oreopithecidae, P r i m a t e s, Mammalia) Teruel & Madrid: 1- 554 - Comptes Rendus de l'Académie des Sciences Paris Alcalá, L. & Morales, J., 1997 - A primitive caprine from 318 série II: 697-704 the Upper Vallesian of La Roma 2 (Alfambra, Daams, R., 1981 - The dental patten of the dormice

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