Late Pleistocene Dwarf Elephants from the Aegean Islands of Kassos and Dilos, Greece

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Ann. Zool. Fennici 51: 27–42 ISSN 0003-455X (print), ISSN 1797-2450 (online) Helsinki 7 April 2014 © Finnish Zoological and Botanical Publishing Board 2014

Late Pleistocene dwarf elephants from the Aegean islands of Kassos and Dilos, Greece

Sevket Sen1,*, Eric Barrier2 & Xavier Crété3

1) Laboratoire de Paléontologie du Muséum, CR2P-CNRS-UPMC, 8 rue Buffon, F-75005 Paris, France (*corresponding author’s e-mail: [email protected]) 2) iSTeP, Université Paris 6, CNRS, Case 129, 4 Place Jussieu, F-75252 Paris, France 3) 280 Chemin de la Villa des Chênes, F-30250 Aubais, France

Received 17June 2013, final version received 3 Jan. 2014, accepted 20 Jan. 2014

Sen, S., Barrier, E. & Crété, X. 2014: Late Pleistocene dwarf elephants from the Aegean islands of Kassos and Dilos, Greece. — Ann. Zool. Fennici 51: 27–42.

Dwarf elephants are quite well known from the western Mediterranean islands, but they are still poorly documented in most eastern Mediterranean islands. This paper reports on the discovery of a third lower molar of a dwarf elephant from the island of Kassos, which is situated in the southern Aegean, between Crete and Karpathos. This molar is determined as Palaeoloxodon aff. creutzburgi, a dwarfed species known from the Late Pleistocene caves and deposits in Crete and derived from the mainland straight-tusked elephant P. antiquus. An upper molar (M3) from the island of Dilos, previously referred to Elephas antiquus or E. mnaidriensis, is also redescribed. It is almost identical to the upper molars of a paleoloxodontine elephant from Naxos. Its small size and lack of sufficient comparative material lead us to determine it as Palae- oloxodon sp. The palaeogeographic evolution of the Aegean region during the Pleisto- cene and its impact on the dispersal of elephants in the Aegean islands are discussed.

Introduction trial mammals reached Mediterranean islands through temporary land bridges. Sondaar (1977) Endemic insular mammals are commonly proposed another alternative, assuming that the recorded in Pleistocene deposits and cave infill large mammals may have reached islands by of Mediterranean islands. They display particu- swimming and small mammals by passive trans- lar adaptational features and evolutionary trends, port on natural rafts. Whatever the way terrestrial different from that of their continental relatives. mammals used to reach and to populate Mediter- Elephants, hippos and ruminants (in particular ranean islands, their affinities with mainland deer) for large mammals, and a few murids mammals indicate palaeobiogeographic relation- and insectivores for small mammals are the ships of islands with the surrounding mainland. main mammalian taxa settled in the Mediter- The “island rule” in mammals, i.e. impact of ranean islands during the Quaternary (Masseti island conditions on the trends towards dwarfism 2009, 2012). They apparently reached islands of large mammals and towards gigantism of when particular conditions favored their disper- small mammals, is also an interesting phenom- sal. Azzaroli (1977) hypothesized that terres- enon that is largely discussed in the literature 28 Sen et al. • Ann. ZOOL. Fennici Vol. 51

(see e.g., Foster 1964, Van Valen 1973, Palombo several fossil-elephant experts recognized the 2007, and references therein). These authors and genus status of Palaeoloxodon (e.g. Caloi et others (e.g. Thaler 1973, Case 1978, Lomolino al. 1996, Doukas & Athanassiou 2003, Inuzuka 1985, 2005, Angerbjörn 1986, Damuth 1993, & Takahashi 2005, Stuart 2005, Ferretti 2008, Palombo, 2004, 2007, 2009, Meiri et al. 2006, Herridge & Lister 2012). We decided to follow Raia & Meiri 2006, 2011, Van der Geer et al. them. 2010, Lomolino et al. 2012) provided several The systematic status of island elephants is explanations for these trends. Among them the also the subject of current debate. For instance, surface of the island and its distance to the main- Vaufrey (1929) referred to Elephas falconeri land, its food resources, intra- and interspecific the specimens from Sicily and Malta, but also competition, and absence of predation pressure from Cyprus and Crete. Later studies recognized are considered the main factors that impacted on the distinct species status of elephants from the the evolution of island populations. eastern Mediterranean islands (see Sondaar & The occurrence of Pleistocene endemic Boekschoten 1967, Kuss 1970, 1973, Mol et al. mammals on Mediterranean islands indicates 1996, Poulakakis et al. 2002a and references that they had particular ecosystems quite dif- therein). However, during the last decade, sev- ferent from that of the surrounding mainland, eral authors questioned the occurrence of some due to their isolated conditions. Their mam- dwarfed elephant species on several islands. malian faunas are unbalanced, and the lack of Consequently, elephant populations on each iso- carnivorous predators is particularly noticeable. lated island should have evolved independently Among these endemic mammals, elephants are (Palombo 2004, 2007, Theodorou et al. 2007, particularly remarked because of the spectacular Ferretti 2008, Herridge & Lister 2012). Con- reduction of their size. Dwarf elephants are also cerning the Aegean realm, this question is cru- known from the western Mediterranean islands cial for the systematic attribution of elephant such as Malta, Sicily, Sardinia, and from the remains that are known from several islands. The eastern Mediterranean islands such as Cyprus, only named Pleistocene elephants species are Crete and several Aegean islands. from Crete: the small Mammuthus creticus (Bate This paper reports on the discovery of a small 1907), the average-sized Palaeoloxodon creutz- elephant molar from the Late Pleistocene on burgi (Kuss 1965) and a possible third large- Kassos and reviews the systematic assignment of sized species that is compared to P. antiquus another elephant molar from Dilos. (Falconer & Cautley 1847) (Mol et al. 1996, European Pleistocene elephants are grouped Poulakakis et al. 2002a and references therein). in the genera Mammuthus Brookes, 1828 (M. Elephant fossils that belonged to a species of the meridionalis, M. trogontherii, M. primigenius) size of P. creutzburgi or larger have also been and Elephas Linnaeus, 1758 or Palaeoloxodon found on several south Aegean islands: Naxos, Matsumoto, 1924. Opinions differed during the Dilos, Kythnos, Serifos, Milos, Paros, Kalym- last half-century on the generic attribution of nos, Rhodos and recently Kassos (Sondaar 1971, the species grouped under the last two genera Kuss 1973, Dermitzakis & Sondaar 1979, Caloi names. Maglio (1973) included the species et al. 1996, Palombo 2001, Doukas & Athanas- antiquus in the genus Elephas, while Aguirre siou 2003, Tsoukala et al. 2011, Masseti 2012). Enríquez (1968–1969) preferred the genus The remains from Kythnos, Serifos, Milos and name Palaeoloxodon to distinguish fossil Eur- Paros are just mentioned in the literature without asian strait-tusked elephants from the extant and any description and illustrations (Palombo 2001, subfossil Asian Elephas. Since that time, the 2004) while the specimens from other islands species antiquus waltzed between Elephas and are better known. In the present paper, we also Palaeoloxodon or was nominated as Elephas discuss the systematic status of some findings (Palaeoloxodon) antiquus (Mol et al. 2007). from these islands based on morphological com- This was also the case for the Mediterranean parisons with the Kassos and Dilos molars. The island elephants presumably derived from the tooth terminology follows Maglio (1973). species antiquus. During the last two decades, Ann. Zool. Fennici Vol. 51 • Pleistocene dwarf elephants from the Aegean islands 29

Fig. 1. Simplified geological map of Kassos Island (modified after Barrier 1984). The arrows indicate the localities where the elephant molar and deer fossils were found.

Localities and their geological 63) “the Karpathos/Kasos deer must be a differ- context ent and probably new genus”. According to De Vos (2006: 129) the Kassos deer “is a species The small island of Kassos (surface 49 km2) is on its own and different from Candiacervus situated in the southern part of the Aegean Sea, cerigensis”. The occurrence of an elephant on between the large islands of Crete and Karpa- Kassos is new, and its discovery contributes to thos. It is a mountainous island whose highest a better understanding of Pleistocene palaeobio- peak reaches 601 m a.s.l. It is mainly formed geographic context of this island. of Mesozoic and Palaeogene limestones; in the One of us (Xavier Crété) discovered the lowlands of the island some marine Neogene Kassos elephant molar when he was hiking in deposits unconformably cover the substratum. early August 2011 along the northwestern slopes Quaternary deposits are piedmont screes and of the Kefalos Mountain (465 m) in the western alluvions in riverbeds, especially in the northern part of the island. The geographical coordinates and western parts of the island (Barrier 1979, and the altitude of the site where he found this 1984), in addition to karstic deposits that may molar are 35°23´45.6´´N and 26°24´´24.7´´E, have been formed locally in caves, fissures or and 165–170 m a.s.l., respectively (Figs. 1 and fault scarps. These Quaternary deposits are not 2). The elephant molar was in a loose block known as fossiliferous, except in a locality at probably fallen from a nearby karstic infill. about 600 meters east of the last houses of the Indeed, to the north of the Kefalos Mountain, capital Fry, on the coast, where Kuss (1969) an approximately E–W trending normal fault recorded cranial and postcranial remains of a forms cliffs in the Eocene limestones (Figs. 1 fossil deer in cemented alluvial fan deposits. He and 2). Along these fault cliffs, occasional fis- referred these fossils to Cervus cretensis, and sure fillings and footwall accumulations contain later (Kuss 1975) to Candiacervus cerigensis, brecciated red deposits. They are strongly eroded which he had described from Late Pleistocene because of the young topography and barren of the neighbouring island of Karpathos. Der- landscapes. The age of this fault as well as of mitzakis and Sondaar (1979) suggested that the sediments that are accumulated on its footwall occurrence of the same species on both islands is not known. The sediment in which the ele- indicates the lack of a real barrier between phant molar was preserved is a pinkish-reddish Kassos and Karpathos. For Sondaar et al. (1996: brecciated marl, consolidated by a calcareous 30 Sen et al. • Ann. ZOOL. Fennici Vol. 51

Fig. 2. Northwestern slopes of the Kefalos Mountain on Kassos. The arrow indicates the site where the elephant molar was found. Note the fault cliffs along the slopes of the mountain marked by light-coloured rock out- crops at the level of the arrow.

Fig. 3. A lower left third molar (m3) of Palaeoloxodon aff. creutzburgi from Kassos, Greece. (A) Occlusal view, and (B) lateral view with an indication (white line) of the crown-root limit. Scale bar = 5 cm.

Fig. 4. An upper left third molar (M3) of Palaeoloxodon cement. It was prepared at the Natural History sp. from Dilos, Greece (cast, original stored at Dilos Museum of Paris, using successive baths of 5% Archeology Museum). (A) Occlusal view, and (B) lateral diluted formic acid. view. Scale bar = 5 cm. In addition to the Kassos elephant molar (Fig. 3), we redescribe a third upper molar from Dilos (Fig. 4). The tiny island of Dilos (3.5 km2) monuments in the ancient city of Dilos, to the is part of Cyclades, and is situated some 20 east of the Apollo temple, 4 m a.s.l. (Cayeux miles northwest of the large island of Naxos, and 1908). He referred it to Elephas antiquus Fal- between Rhinia and Mykonos. Cayeux (1908) coner & Cautley, 1847. Later, Vaufrey (1929: fig. reported on the discovery of an elephant molar in 38) illustrated this molar and speculated whether the alluvions of the Inopos River at the northern it belongs to Elephas mnaidriensis Adams, 1874, part of Dilos. This molar was found during a a species previously defined from Malta, or to survey excavation to study the substrate of the a small sized Elephas antiquus. This record Ann. Zool. Fennici Vol. 51 • Pleistocene dwarf elephants from the Aegean islands 31 was often mentioned in the literature (e.g. Der- Material from Kassos Island mitzakis & Sondaar 1979, Symeonidis & The- odorou 1982, Kotsakis 1990, Caloi et al. 1996, An isolated left third lower molar (m3) labelled Poulakakis et al. 2002a, Doukas & Athanas- as KAS-1 (Fig. 3). Its basal length is 165 mm; siou 2003, Tsoukala et al. 2011); most of these preserved crown length 162 mm; maximum authors noted that this molar belonged to a width 54 mm at the level of 3rd plate; maximum somewhat dwarfed elephant without however crown height 72.9 mm at the 7th plate. This suggesting any systematic assignment. Its sim- specimen is stored at the Arhaeological Museum ilarities in size and dental pattern to the Kassos of Kassos, and its casts are available at the Nat- molar led us to restudy the Dilos molar from a ural History Museums of Paris (MNHN) and cast kept at the Paris Museum. London (NHM). The southern Aegean region is formed of several tectonic structures, the most prominent of them being (1) the south Aegean volcanic arc, Description which extends approximately from the Gulf of Corinth in the west to Kos in the east (including In occlusal view, the Kassos molar is moderately Cyclades), (2) the Cretan arc, which extends curved laterally, so its labial face is concave. The from Peloponnese to Rhodos via the largest occlusal surface is concave and progressively Aegean island of Crete, and (3) the Cretan Basin tapers backwards. The stage of wear indicates that separates these two arcs. South and south- that it belonged to an adult individual. It bears west of the Cretan arc, the Hellenic trench is a nine plates plus talon. The advanced stage of subduction zone where the African plate plunges wear of the anterior plates, which are also partly beneath Eurasia. On the other hand, the Anato- damaged, and the lack of the anterior root indi- lian plate is extruded southwestward under the cate that some anterior plates may have been effect of the collision of the African plate with lost. All plates but the last one are progressively the Anatolian plate in southeastern Turkey. The abraded. The lamellar frequency is 6.5. The talon southward extension of the Aegean–West Ana- forms a narrow platelet as high as the last plate. tolian region since the late Miocene onwards The crown height increases from the first pre- is related to a differential advancement of the served plate to the 7th plate, and then it slightly upper plates (Greece and Anatolia) over the decreases towards the back. On the occlusal view, African plate (Piper & Perissoratis 2003, Agos- the plates form narrow and slightly oblique ridges tini et al. 2010). In other words, the southern with almost regular antero-posterior width. In Aegean realm is subsiding since at least the other terms, the width of the plates does not early Pliocene. The effects of the subsidence change significantly in their central, lateral and on the dispersal of Pleistocene elephants in the medial parts, except when there is anterior and/ southern Aegean realm will be discussed in the or small posterior enamel bulges near the center. Discussion. The enamel bands are delicately wrinkled. The enamel thickness varies between 1.8–2.2 mm depending on the wear stage. On the occlusal Systematic study surface, the enamel loops of the plates and the cement areas between the plates occupy similar Palaeoloxodon aff. creutzburgi (Kuss, surfaces. The roots are not preserved. The Kassos 1965) molar is determined as an m3 based on the following characters: Order Proboscidea Illiger, 1811 Family Elephantidae Gray, 1821 1. In occlusal view, its distal part is laterally Genus Palaeoloxodon Matsumoto, 1924 curved from the 6th plate backward. The curve makes an angle of 160°. In that it is Type locality: Kalo Chorafi cave, Rethymnon, Crete. similar to the left m3 from Simonelli Cave Holotype: left m1, Nr 22/1963 (Kuss 1965: pl. 3). in Crete illustrated by Kotsakis (1980) and 32 Sen et al. • Ann. ZOOL. Fennici Vol. 51

referred to P. creutzburgi, as well to the left burgi (Kuss 1965). Some papers mentioned the m3 of P. tiliensis from Tilos (Theodorou presence of a third form a little larger than the 1983: pl. 3, fig. 1), right m3 of “Elephas type specimens of P. creutzburgi from Kalo antiquus” from Kythera Island (Kuss 1967, Chorafi cave and about 20% smaller than the fig. 2), and many other m3 referred to Palae- mainland species Palaeoloxodon antiquus; this oloxodon sp. from the European mainland third form was referred to P. antiquus, P. cf. and Mediterranean islands. The m3s from antiquus or to a new species, Palaeoloxodon Coumbes Cave (Crete) are less curved than chaniensis (Kuss 1973, Dermitzakis & Sondaar the above-mentioned specimens (Symeo- 1979, Symeonidis & Theodorou 1982, Der- nidis & Theodorou 1982). The holotype of mitzakis & de Vos 1987, Symeonidis et al. 2000, P. creutzburgi, and the only lower molar 2001, Poulakakis et al. 2002a). from the type locality Kalo Chorafi (Crete), Mammuthus creticus from the Early-Middle is a worn left m1 (Kuss 1965). Its occlusal Pleistocene site of Cape Meleka near Rethymnon surface is straight. Theodorou (1983) also is a very small sized elephant and it has peculiar illustrated two m2 from Tilos, the one with molar characters (reduced number of lamellae, a similar occlusal wear, and these molars low crown, wide occlusal surface, broad cement are barely curved, nothing to compare with areas, simple enamel bands) clearly different the curvature seen on the Kassos molar. The from that of the Kassos molar. Bate (1907: 243) general curvature trend on the m3 of Palaeo- noted that “except with regard to the immense loxodon sp. is that the less a tooth is abraded difference in size the characteristics of the the more it appears to be laterally curved molars (of Cape Meleka), more especially in the [e.g. curved m3 in Bate (1907: pl. 22, fig. 6) lowness of the crowns, appear to resemble more and Palombo & Ferretti (2005: fig. 12), but closely those of E. meridionalis than any other almost not curved m3 from Coumbes Cave of the larger Elephants of the Mediterranean (Symeonidis & Theodorou 1982: pl. 3–5)]. region“. Lastly, studying the Bate’s collection at 2. The distal part of the Kassos molar tapers the NHM-London and some new material, Her- backward. In a similar degree of attrition, ridge and Lister (2012) confirmed morphological the m1 from Kalo Chorafi as well as the m2 evidence on its ancestral relationships with Early from Tilos Island have the distal edge rather Pleistocene Mammuthus meridionalis or even rounded and lack the talon. with Late Pliocene M. rumanus. These authors 3. The Kassos tooth is worn up to its eight pre- estimated that M. creticus may have a body mass served plate. In such a degree of attrition, the about 310 kg and shoulder height 1.13 m. Mam- distal face of an m1 or an m2 should have a muthus creticus was found associated with the pressure mark of the following tooth. This murid Kritimys kiridus, and it apparently reached is not the case on the Kassos molar. For the Crete during the first migration wave of land identification of a molar from Simonelli Cave mammals in Early Pleistocene (Mayhew 1977, in Crete, Kotsakis (1980: 115) compared 1996), or ca. 0.9 Ma ago (Reese et al. 1996, it with the lower molars of Palaeoloxodon Sondaar & Van der Geer 2005, Van der Geer et antiquus, and observed that on m2s “there al. 2010). However, all these dates need to be is always a flattening however small, on the confirmed or evaluated with new dating methods posterior side due to the contact with the and more accurate analyses, as noted by Stuart growing m3”. Such a contact is not observed and Lister (2012). on the Kassos molar. A second elephant species from Crete is Palaeoloxodon creutzburgi (Kuss 1965). It was recorded in Crete at more than 15 localities, all Comparison of the Kassos m3 supposedly of latest Middle or Late Pleistocene age (Poulakakis et al. 2002a). This is a large Dwarf elephants are best known from the neigh- sized form having palaeoloxodontine affinities bour island Crete with two species: Mammuthus in its dentition. The systematic status of palae- creticus (Bate 1907) and Palaeoloxodon creutz- oloxodontine elephant remains from Crete has Ann. Zool. Fennici Vol. 51 • Pleistocene dwarf elephants from the Aegean islands 33 been debated since 1960s. Lastly Poulakakis (1965), Kotsakis (1980) and Poulakakis et al. et al. (2002a) reviewed the material from the (2002a) referred to P. creutzburgi. Cretan localities and retained two subspecies, From other Aegean islands some elephant Elephas (Palaeoloxodon) antiquus creutzburgi remains of the size of P. creutzburgi were for specimens similar in size to that of the also recovered. From Rhodos, Symeonidis et Kalo Chorafi cave, andElephas (Palaeoloxodon) al. (1974) recorded postcranial remains of a antiquus chaniensis for the larger sized elephant dwarfed elephant that they tentatively referred remains from the caves of Vamos and Coumbes to a possible subspecies of P. antiquus. From in Crete. We estimate that the taxon creutzburgi Naxos, Mitzopoulos (1961) described a palate deserves the species status, and here it will be with both M2 that Symeonidis et al. (2000) retained as P. creutzburgi. Its type material from referred to their new species Elephas chanien- the Kalo Chorafi cave includes two upper-cheek sis. An M3 from Dilos (see below) that Cayeux teeth identified as P2 and P3, and one damaged (1908) determined as Elephas antiquus also rep- and worn m1 (Kuss 1965). This m1 is clearly resents a dwarfed elephant. All these findings wider than the Kassos m3 (75.2 mm contra 54 cannot be compared with the Kassos m3. mm) and also all m3s of “large elephant” known Elephas tiliensis Theodorou et al. (2007) from other sites in Crete. However, it should be from Charkadio Cave on Tilos is well docu- noted that the anterior molars, in particular when mented with many skeletal parts, cranial and they are well worn, might be larger than the m3, postcranial bones, and several cheek teeth. They as already noted by Bate (1907) on the mandible were extensively described and illustrated by with the m2–m3 from Kharoumes Bay in Crete. Theodorou (1983) who referred it in his Ph.D. Concerning the size of molars, it is not possible thesis, following Symeonidis (1972), to Palae- to distinguish any size group among the avail- oloxodon antiquus falconeri. Later on, Theo- able specimens from Crete, as well as including dorou et al. (2007: 23) referred the Tilos ele- the m3s from Kythera and Naxos islands (Mitzo- phant to a new species because (in the diagno- poulos 1961, Kuss 1967). The size and different sis): “even if its dimensions and morphology ratios of the Kassos m3 fall close to the aver- were equal to other endemic Mediterranean ele- age values of palaeoloxodontine elephants from phants, its isolation is more than adequate for the Crete (Table 1). description of a new endemic species”. In other The lower molar morphology is best words, this species is new not because it is mor- observed on the type specimen (m1) from Kalo phologically different from the other elephant Chorafi, on the m2 (not M2 as determined by species, but because it is endemic to Tilos, and Poulakakis et al. 2002a) from Katharo Basin, on thus it is supposed to be genetically isolated from the m2–m3 from Kharoumes Bay (Bate 1907) the other elephants hence deserving a new spe- and on the m3s from Grida Avlaci, Simonelli cies status. According to the measurements given Cave and Coumbes Cave (Simonelli 1908, Kot- by Theodorou (1983), the size of the molars of sakis 1980, Symeonidis & Theodorou 1982). In this species is smaller than or in the lower range all these specimens, as well as on the Kassos of variation of Palaeoloxodon creutzburgi from m3, the plates are narrow, the interplate cement Crete (Table 1). The characters of cheek teeth areas are narrower than the plates, the central such as lamellar frequency, enamel thickness, part of plates is a little larger (mesio-distal) slightly folded enamel, lamellae with parallel than the lateral parts, the plates bear anterior enamel bands, oblique position of the lamellae and posterior enamel bulges in their central in the posterior half of the m3, presence of a part, the enamel is gently folded but much less small central loop on the lamellae and crown than in Palaeoloxodon antiquus molars of sim- height of upper molars are also similar to that ilar wear stage, the plates slightly abraded are of P. creutzburgi. According Poulakakis et al. composed of one wide central loop and two or (2002b, 2006) and Theodorou et al. (2007), Ele- more lateral rounded rings. In all these features, phas tiliensis inhabited Tilos between ca. 50 000 the Kassos m3 cannot be distinguished from and 4000 BP, i.e. at least partly contemporaneous the lower molars of Cretan localities that Kuss with P. creutzburgi. Although smaller, the mor- 34 Sen et al. • Ann. ZOOL. Fennici Vol. 51 et al. Mol et al. 1996 References 1982 Symenoidis & Theodorou Simonelli 1908 Kotsakis 1980 Kuss 1965 Bate 1907, Kuss 1966 This paper Lister 1998 & Tsoukala Lister Kuss 1967 Theodorou 1983 Bate 1907 Ferretti 2008 Markov 2007 Poulakakis 2002 3 2.5 2.5 2.5 (mm) 3–3.5 1–2.5 1.5–2.5 1.8–2.2 2.3–3.3 2.5–3.0 Enamel 1.8–2.88 thickness less than 4 ? 9 6 8 10 LF 5.2 6.5 5.5 6–7 4.5* 4.75 5.5–6 6.6–7 6.5–7.7 2 – 2.1 1.2 1.35 1.55 index – – 133 172 104 95.3 72.9 (mm) 79.4* 39–50 Height Hypsodonty 4.3 3.9 3.2 L / W 2.4* 2.1* 1.6* 3.38 2.87 2.92 3.06* 2.62–2.87 75 60 64 56 67 63 47 54 86 60 83 75.2 (mm) Width 33–41 48–53 ? 216 370 145 242 180* 125* 125* 140* 165* (mm) 121.3* Length 165–181 170–174 110–160* >8 x9p >7p x11p x12p x17p x12p x10p x14p x>9p x>8p x>9p plates preserved

1 1 1 1 1 2 n n umber of 1 1 1 1 3 2 2 1 Kythera I sland, Greece Crete Avlaci, Grida Simonelli Cave, Crete Simonelli Cave II , Crete Katharo Basin, Crete Coumbes Cave, Crete Kalo Chorafi cave, Crete Kharoumes, Crete Kassos, Greece Ryahovo, Bulgaria Tilos I sland, Greece Tilos Cape Maleka, Crete Puntali Cave, Sicily Grevena, Greece

(m1)

sp. creutzburgi aff. aff. Palaeoloxodon Palaeoloxodon creutzburgi Palaeoloxodon creutzburgi Palaeoloxodon creutzburgi P. creutzburgi (m2) P. P. creutzburgi aff. Table 1. Comparative Table measurements and ratios of the third lower molars (m3, except an m1 from Kalo Chorafi and an m2 from ‘>’ = part of the tooth missing. Katharo) n = number of specimens, x = talon, p plate, L / W length/width ratio, LF = lamellar frequency. oflocalities in Greece, Bulgaria and I taly. Elephantidae from several Species Locality Palaeoloxodon creutzburgi * measured from incomplete tooth, true value greater than the given one. Palaeoloxodon creutzburgi Palaeoloxodon antiquus Elephas tiliensis Mammuthus creticus Palaeoloxodon mnaidriensis Palaeoloxodon antiquus P. Ann. Zool. Fennici Vol. 51 • Pleistocene dwarf elephants from the Aegean islands 35 phological pattern of its cheek teeth shows great stored at the Archaeological Museum of Dilos similarities with that of P. creutzburgi. Island. This label also mentions that this tooth Kuss (1967) described and illustrated a “was found in the sandy alluvium of the former right m3 from Kythera as belonging to Elephas torrent Inopos, during the excavations near the antiquus. This m3 is larger than any other m3 sanctuary of Apollo”. from Crete and Kassos (Table 1), it has more plates preserved, although it is in other characters quite similar to both P. antiquus and P. Description creutzburgi. However, the lack of central enamel bulges on the plates resembles more the pattern The Dilos molar should be an M3 because it of P. antiquus. Masseti (2009, 2012) referred to tapers backward though the wear abraded almost P. antiquus a lower jaw fragment from Kalymnos all plates except the last two, and there is a talon with the m3 (?), which has the anterior and poste- typical for a last upper molar. Its occlusal surface rior parts damaged. Masseti did not provide any is almost flat. It preserves nine plates plus talon. measurements for this molar but he illustrated it The advanced wear degree suggests that some with a scale. Using this scale, the estimated max- anterior plates may have been lost. The wear is imum width of this molar should be less than 40 progressive backward, and the two last plates mm, which is smaller than any other lower molar are not yet abraded. The lamellar frequency is referred to P. creutzburgi, and of course infinitely 6.1. The crown height rapidly increases from small as compared with the molars of P. antiquus. the first preserved plate to the 7th plate, and This molar bears seven preserved plates that are then it slightly decreases towards the back. In similar in shape and pattern to that of P. creutz- occlusal view, the plates form narrow lamellae, burgi and P. tiliensis. The size of the Kalym- almost perpendicular to the longitudinal axis. nos molar makes it the smallest palaeoloxodon- The first six preserved plates are composed of tine elephant of the Aegean islands. We have to two unequal enamel loops, which are more or emphasize that Kalymnos is close to the western less confluent depending on the degree of wear; Anatolian coasts and it is on the continental pla- the lingual loop is larger than the labial one. teau. The status of the Kalymnos specimen cannot The remaining three plates display each one big be solved without a detailed comparative study. central and two small lateral rings. The enamel is Despite the great similarities of the Kassos delicately folded and the enamel wear surfaces molar with those from Crete, we preferred to are inclined mesially. The talon is a small cusp determine it as P. aff. creutzburgi, due to the fact at the mid depth of the crown. The roots are not that P. creutzburgi is still poorly documented preserved. in Crete (limited number of lower molars to be compared with Kassos molar), and that only a more abundant material from the type locality Comparison of P. creutzburgi or from similar age localities in Crete can solve its taxonomic status. As noted above, Cayeux (1908) described this tooth from Dilos as a last upper milk molar of Elephas antiquus. According to Vaufrey (1929) Palaeoloxodon sp. who gave an illustration, the Dilos specimen is the last molar (M3) and it can be referred to Ele- Material from Dilos phas mnaidriensis, a species previously known from Malta and Sicily. Since that time the sys- An isolated left third upper molar (M3) (Fig. 4). tematic status of this specimen remained open. Preserved crown length 156 mm; maximum As shown above, this specimen is a left M3 width 59 mm at the level of 3rd plate; maximum belonging to an adult individual. Its narrow crown height 110 mm at the 7th plate. A cast of plates, unequal rings and gently folded enamel this specimen is preserved at MNHN. According bands indicate a palaeoloxodontine elephant. Its to the label of this cast, the original specimen is size is about 60% of the strait-tusked continental 36 Sen et al. • Ann. ZOOL. Fennici Vol. 51

elephant P. antiquus. Upper cheek teeth of large sized palaeoloxodontine elephants from Aegean islands are poorly documented: one P2 and one P3 from the Kalo Chorafi cave in Crete referred to P. creutzburgi (Kuss 1965, 1973), a palate with both M2s from Naxos tentatively referred to Elephas melitensis (Mitzopoulos 1961) and Athanassiou 2001 Markov 2007 References Kuss 1965 Kuss 1965 Mitzopoulos 1961 This paper Theodorou 1983 Palombo & Ferretti 2005 several upper cheek teeth from Tilos referred to a new species Elephas tiliensis (Theodorou

3 1983, Theodorou et al. 2007). The Dilos M3 is 2.0 3.8 2.6 2–3 1.5–3 2–2.9 1.7–3.9 Enamel

thickness a little larger and more hypsodont than the specimens from Naxos (Table 2). On the contrary, the LF 5.4 6.5 7.1 5.9 6.1 5.0 pattern of the occlusal surface is very similar in 7.7–9 6.9–7.2 having equal number of plates, a narrow shape, unequal enamel loops, gently folded enamel, and 2.31 1.86 1.72 1.92 2.58* index three loops on the unworn plates. The M3s from 2.37–2.63* Hypsodonty Tilos are smaller than those of Dilos and Naxos, with however a similar occlusal pattern. The lamellar frequency is quite similar on the Dilos 86 45* 66* 110 170 183* (mm) Height and Naxos molars, but higher on the Tilos molars 101–104 220–250 (Table 2). Based on the size and tooth pattern of 3.2

L / W these molars, it can be supposed that the Dilos 3.11 1.17 1.46 2.77 2.64 2.28 3.70 and Naxos specimens belong to the same species, while those from Tilos probably belonged 71 59 90 63.3 64.8 (mm) Width 45–46 47–53 93–95 to a different species. Concerning the P2 and P3 from the Kalo Chorafi cave in Crete, the type locality of P. creutzburgi, it will be hazardous to 156 280 73.8 94.6 230* (mm) Length

95–124* compare them with the Dilos M3. They appar- 125–127 340–355 ently represent a larger sized form, and the plates are not divided in unequal loops as on the Dilos >5 >5 >9

>9p and Naxos molars. In summary, the poor avail- x14p x14p x19p plates (mm) >7–11 preserved

N umber of able knowledge on the upper dentition of palaeoloxodontine elephants from the Aegean islands does not allow a definite systematic assignment of the Dilos M3, hence its determination as Palaeoloxodon sp.

Discussion

Locality Ryahovo, Bulgaria Kalo Chorafi cave, Crete Kalo Chorafi cave Dilos, Greece Greece Tilos, Greece Penios Valley, Pleistocene dwarf elephants are well known from the western Mediterranean islands of Malta and Sicily. Palaeoloxodontine species from these islands are, from the smaller to the larger, Palae- oloxodon falconeri (Busk, 1867), Palaeoloxo- sp. (M2, 2) n axos, Greece sp. (M3, 1) don melitensis (Falconer 1862, in Busk 1867) and Palaeoloxodon mnaidriensis (Adams, 1874) (Ambrosetti 1968, Caloi et al. 1996, Palombo 2001, Marra 2005, Ferretti 2008, Masseti 2009), P. antiquus (M3, 1) P.

number of specimens) = length/ plate, L / W = talon, p = x = 2. Comparative measurementsand ratios of the upper molars Elephantidae several localities in Greece, Bulgaria and I taly. from Table ‘>’ = part of the tooth missing. width ratio, LF = lamellar frequency. Species (tooth, creutzburgi (P2, 1) P. * measured from incomplete tooth, true value greater than the given one. P. creutzburgi (P3, 1) P. Palaeoloxodon Palaeoloxodon tiliensis (M3, 3) P. antiquus (M3 average, 6) P. antiquus (M3, 2) P. various locs i taly, although the possible synonymy between these Ann. Zool. Fennici Vol. 51 • Pleistocene dwarf elephants from the Aegean islands 37 species is discussed since Pohlig (1893) (see ferent from the elephant of Kassos and Dilos in also Ambrosetti 1968, Caloi et al. 1996, Ferretti being significantly smaller and in having lower 2008). These authors demonstrated the affin- crowned cheek teeth. Boekschoten and Sondaar ities of these species with the “normal” sized (1972) reported the presence of a larger elephant strait tusked elephant of Europe, Palaeoloxodon in Cyprus based on some tooth fragments that are antiquus. Some of these western Mediterranean not reliable for any comparisons. Pleistocene elephants are in some extent simi- How many elephant species did inhabit the lar in size and morphology to the elephants of Aegean islands during the Pleistocene? This cru- the Aegean islands. For instance, Palaeoloxodon cial question is difficult to answer reliably due mnaidriensis is similar in size to Palaeoloxodon to the fact that the elephant remains from many creutzburgi from Crete (Table 1). The length/ islands, except in Tilos, are still quite scarce width ratio, number of lamellae and lamellar and fragmentary. The available material and the frequency, gently folded enamel bands and the state of paleontological studies do not allow progress of the occlusal wear on cheek teeth are for estimating variations in size and morphol- also characters shared by P. mnaidriensis and P. ogy because of sexual dimorphism, intra-specific creutzburgi. variation or degree of dwarfism. However, the similarities of their tooth Another major question currently debated pattern are the characters inherited from their concerns the endemic evolution of island ele- common ancestor Palaeoloxodon antiquus. phants, and consequently the occurrence of inde- These are also reliable differences between these pendent species on each island (Palombo 2004, two species, for instance for the m3, sharp lamel- 2009, Theodorou et al. 2007, Ferretti 2008). This lar edges in P. mnaidriensis but square shaped in view assumes a genetic isolation of elephants P. creutzburgi, lamellae generally curved back- after they reached the islands, and consequently ward in the first, straight in the second, and so they became independent species on each island. on. Despite the above-mentioned similarities, For instance Palombo (2004, 2009) and Ferretti it is assumed here that the dwarfed elephants (2008) questioned the species identity of Sicilian of Malta and Sicily have an independent evo- and Maltese elephants despite their great size lutionary history from their mainland relatives and morphological similarities. It is conceivable as well as from the other island elephants. Con- that elephants may have evolved independently sequently, they represent independent species. on islands that remained isolated from the main- Indeed, the western Mediterranean islands have land (e.g. Cyprus and Malta). We do not know their own palaeobiogeographic context (Marra for certain how and when the Aegean islands 2005). During the Pleistocene they were never became isolated from the mainland, and also connected to the Aegean islands, neither had from each other, in particular when Pleistocene they any mammalian exchange. elephants inhabited these islands. As seen in This is also the case for Cyprus, which Fig. 5, the distances between the Aegean islands became definitely isolated from the mainland that yielded elephant fossils are short. after flooding of the Mediterranean at the end of In addition, we have to take into account the the Messinian Salinity Crisis, ca. 5.3 Ma (Hadjis- fact that the Aegean region is currently under- terkotis et al. 2000). The dwarf elephant Palae- going a widespread regional extension, which oloxodon cypriotes (Bate 1903) inhabited Cyprus can be dated back at least to the Late Miocene. during the Middle-Late Pleistocene (Simmons This is due to the convergence of the Afro-Ara- 1999). It is probably derived from Palaeolox- bian and Eurasian plates and the southwestward odon antiquus although remarkably reduced in motion of the Anatolian block. Using seismic size, i.e. about 1 to 1.2 m at the withers (Davies reflection profiles and available chronostrati- & Lister 2001; G. Iliopoulos unpubl. data). Its graphic data, Piper and Perissoratis (2003) esti- molar pattern displays the primitive characters mated the southward Quaternary extension rate of the paleoloxodontine type with low-crowned of the Aegean realm as 37 mm/year and the cheek teeth, gently folded enamel and low lamel- subsidence rates for the last 0.2 Myr at about lar frequency. Palaeoloxodon cypriotes is dif- 2.5 m/kyr in the Santorini-Anafi Valley region, 38 Sen et al. • Ann. ZOOL. Fennici Vol. 51

Fig. 5. Present day map of the southern Aegean, showing the islands inhabited by Mammuthus creticus (Crete), Palaeoloxodon creutzburgi (Crete), Palaeoloxodon spp. (Kassos, Rhodos, Dilos, Naxos and possibly Kalymnos) and Palaeoloxodon tiliensis (Tilos). The dashed lines around the islands and continent indicate –100 m depth that was exceeded during the last glacial sea level drop. The red arrows indicate the possible dispersal routes of island elephants. and up to 1.6 m/kyr on the North Aegean Shelf. Crete and Palaeoloxodon cypriotes in Cyprus. In According to Agostini et al. (2010), the subduc- addition, the available fossil documentation does tion rates in the southern Aegean may reach 4 not inform us on the possible sexual dimorphism cm/year during the Quaternary. Indeed, a final and/or size variability of the palaeoloxodontine tectonic phase beginning at about 0.8 Myr ago elephants on Aegean islands. markedly changed the fault pattern in the South Poulakakis et al. (2002a) discussed the pos- Aegean arc, and all the Aegean Basin underwent sible origination of the elephant species in Crete. a fault-controlled subsidence (Piper & Perissora- It is hypothesized that the pigmy elephant Mam- tis 2003). Thus, when P. creutzburgi and its rela- muthus creticus is derived from Mammuthus tives inhabited the southern Aegean islands, one meridionalis based on its older age (ESR dates can speculate how large these islands were (pos- ranging from 846 000 to 475 550 years; Reese sibly even larger than today), and how deep were et al. 1996, see above) and the similarities of its the channels in between. This implies that these dental pattern with the Mediterranean mammoth. islands may have intermittently been connected This species may have reached Crete during the to the mainland and/or the area of the Aegean sea level lowering in the late Early Pleistocene. Pleistocene islands was larger than today. They For the entrance in Crete of the less dwarf might have sheltered larger sized individuals elephant Palaeoloxodon creutzburgi, Poulakakis and the elephants might not have been totally et al. (2002a: 177) suggest that “the most proba- isolated from their mainland relatives as it hap- ble scenario for colonization of Crete by Middle pened for species such as Mammuthus creticus in and Late Pleistocene mammals was the ‘Island Ann. Zool. Fennici Vol. 51 • Pleistocene dwarf elephants from the Aegean islands 39

Sweepstakes’ route“. Fossil elephant researchers land species P. antiquus. The size of the Kassos generally accept that P. creutzburgi is probably and Dilos molars indicates individuals smaller derived from the strait-tusked continental spe- than P. creutzburgi from its type locality the cies P. antiquus. This species is well represented Kalo Chorafi cave in Crete. on the continents surrounding the Aegean Sea. Immigration of elephants as well as of other It has been recorded in at least 23 Pleistocene mammals onto the Aegean islands probably localities all over Greece (Tsoukala & Lister occurred in several phases, the one in the Early 1998, Athanassiou 2001, Doukas & Athanassiou Pleistocene with Mammuthus creticus, which is 2003, Tsoukala et al. 2011) as well in the Balkans known only from Crete, and the second during (Markov 2007) and Asia Minor (Becker-Platen & the late Middle or Late Pleistocene with the Sickenberg 1968, Albayrak & Lister 2011). palaeoloxodontine elephants. This latter group Indeed, the sea level dropped several times is much less dwarfed than the first one, and during the Pleistocene. For instance, the sea it inhabited several south Aegean islands, i.e. level was about –120 m during the Marine Iso- Crete, Kassos, Rhodos, Kalymnos, Dilos and topic Stage 6 (MIS) (about 140 kyr ago), –100 m probably Naxos. Its occurrence on these islands during MIS 4 (about 60 kyr ago) and –125 m is probably due to sea level lowering during the during MIS 2 (about 20 kyr ago) (Antonioli et al. Middle-Late Pleistocene. 2006). In such a palaeogeographic context, several Aegean islands, today isolated and far from the continent, became connected to the mainland Acknowledgements (Greece or Turkey) or their distance from the mainland considerably diminished (Fig. 5). In It is a great pleasure for us to contribute to this volume in addition, we have to take into account the current honour of Mikael, with whom one of us (S. Sen) hunted mammalian fossils in Turkey, China and more recently in regional extension and fault-controlled subsid- Iran, and shared some exciting discussions making and ence of the Aegean region since the Late Mio- unmaking the systematics and palaeobiogeography of many cene. All these parameters indicate that during groups of mammals. We would like to thank Anu, Suvi and the Pleistocene, the island surfaces and envi- Jussi for inviting us to join the group of Mikael’s friends. ronmental conditions were probably different Pascal Tassy (MNHN, Paris) and Marco Masseti (Firenze) of their present conditions. These events may provided useful comments and literature on fossil elephants. Vincent Pernègre (MNHN, Paris) hunted all fossil elephant have facilitated the dispersal to these islands of molars in the palaeontology collections of the Natural His- deer, elephants and hippopotamuses that were tory Museum of Paris to be compared with the Kassos and common on the mainland, hence their occur- Dilos elephants. Philippe Loubry (CNRS-MNHN, Paris) rence in Crete and several other Aegean islands. made the photos. Philippe Richir prepared the cast of the When sea level rose, these islands became iso- Kassos elephant molar. We are grateful to the referees Vic- toria Herridge, Juha Saarinen and an anonymous referee for lated and their inhabitants evolved under the new their valuable comments and suggestions that allowed to insular conditions. greatly improve this paper.

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