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Palaeloxodon cypriotes, the dwarf elephant of Cyprus: Size and scaling comparisons with P. falconeri (Sicily-Malta) and mainland P. antiquus

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We provide the first detailed biometric study of Palaeoloxodon cypriotes, the dwarf elephant of Cyprus, based on the Bate collection from Imbohary. Molar morphology indicates derivation from P. antiquus, while molar proportions are unchanged from that species, paradoxically implying an allometric shift. To maintain function, enamel is relatively thick and plates have been lost, producing a molar lamellar frequency analogous to P. antiquus milk molars of the same size. Body size was similar to or slightly larger than P. fal- coneri, but the teeth were even smaller. Perhaps as a result, plate loss was more extreme than in that species. Sparse material of slightly larger dwarf elephants from Achna are of uncertain status.
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... Fossil-bearing localities on Cyprus have usually yielded specimens referred to an extremely small-sized endemic elephant, Palaeoloxodon cypriotes (Bate, 1903). This endemic species together with the Sicilian Palaeoloxodon falconeri (Busk, 1867) are the smallest elephants ever evolved, having an adult height of about 1 m (Ambrosetti, 1968;Davies and Lister, 2001). However, there is also sporadic evidence for the presence of a larger elephant form on Cyprus (Vaufrey, 1929;Boekschoten and Sondaar, 1972;Reese, 1995;Iliopoulos et al., 2011). ...
... There is general consensus among authors that this insular form descended from a P. antiquus ancestral population that migrated to Cyprus sometime during the MiddleeLate Pleistocene (Caloi et al., 1996;Palombo, 2004;Herridge, 2010;van der Geer et al., 2010). P. cypriotes is known mainly from dental material, the size of which is slightly smaller than that of the Siculo-Maltese species P. falconeri (Table 2; Davies and Lister, 2001;Herridge, 2010). The richest P. cypriotes-bearing sites are the type locality, P ano Díkomo-Imbohary, on the southern slopes of Pentad aktylos Range, and Akrotíri-Aet okremnos at the southernmost promontory of the island (Bate, 1903c(Bate, , 1904a1904b, 1905Reese, 1995Reese, , 2001. ...
... Some additional but scanty remains of larger-bodied elephants, first found on Cyprus (Achna locality) in 1924, have raised doubts about their attribution to Bate's species (Vaufrey, 1929;Boekschoten and Sondaar, 1972;Reese, 1995;Davies and Lister, 2001;Herridge, 2010); however, they were inadequate for taxonomic evaluation, as their number and preservation state are poor. Over the years more fragmentary dental and postcranial material kept appearing on the SE side of the Mesaoría Basin (the lowland between the Tr oodos and Pentad aktylos mountains), mainly as a result of private collectors' activities. ...
Article
Cyprus, the largest Eastern Mediterranean island, hosted a highly impoverished endemic mammalian fauna during the Pleistocene to early Holocene times. This was a result of its extreme biogeographic isolation since its formation, which prevented the immigration of most terrestrial mammals, except for those with apparent sea channel crossing abilities. The main faunal elements are the extremely dwarfed hippo Phanourios minor, commonly found in many sites across the island, and the dwarf elephant Palaeoloxodon cypriotes. The latter is a very small-sized elephant species, comparable in size with the Siculo-Maltese Palaeoloxodon falconeri. Larger dental specimens found sporadically during the last century, raised the possibility that a second endemic elephant, larger than P. cypriotes, may have also existed in Cyprus. Here we describe a skull recently excavated in the coastal area of Xylophágou, SE Cyprus, which provides evidence that, indeed, two elephant species have existed on the island. The larger species, Palaeoloxodon xylophagou n. sp., is still strongly dwarfed and characterised by elongated, low and wide skull, diverging tusk alveoli and comparatively large molars. Dimensionally the dentition is distinctly larger than P. cypriotes and close to Palaeoloxodon tiliensis, though the skull size is intermediate between P. tiliensis and P. falconeri. Both Cypriot elephant species exhibit morphological affinities with Palaeoloxodon antiquus, which is their probable ancestor. Stratigraphic data suggest that P. xylophagou is older (late Middle Pleistocene), while P. cypriotes is more recent (latest Pleistocene to early Holocene) and may have descended from the former or – less probably – evolved as a result of a separate, more recent colonisation event.
... The enamel thickness of the teeth of the island form is relatively more and the number of plates is reduced: the average plate number at the third molars is 11 instead of 18, as in the ancestral form. Hence, the loss of the tooth's plates in the BOREAS teeth of the Cypriot dwarf elephant is more significant than that in the other island dwarf, P. falconeri (Davies & Lister 2001). ...
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A pioneer comprehensive study of several diminutive last‐generation woolly mammoth teeth (M3) found on the coast of the East Siberian Sea between the mouths of the Alazeya and Malaya Kuropatoch'ya rivers was conducted. Two teeth belonged to one individual. These teeth have a similar lamellar frequency and enamel thickness as teeth of Mammuthus primigenius Blumenbach. The molar crowns from the lower Alazeya region are similar in size to those of the small Late Pleistocene–Holocene mammoths from Wrangel Island. However, the number of plates (17–19, excluding talons) is much lower than that in the teeth of typical Late Pleistocene M. primigenius (23–25). The age data of the examined teeth are beyond the limits of the 14C dating method (>45 000 years BP). Nevertheless, palaeobotanical data allow correlation of the enclosing sediments with the warm Kazantsevo Interglacial (Eemian, MIS 5e) and reconstruction of the average annual temperature, which was warmer than present‐day temperatures. These conditions are confirmed by the δ18O isotopes from the structurally bound carbonate in tooth enamel. The ancient landscape was wetter and more forested than modern landscapes. The diminution of M3 size and loss of posterior plates were a result of the overall decrease in body size, likely in response to landscape change and narrowing of resource space. Mammoths from the lower Alazeya region demonstrate a stage of significant size reduction, although the dwarfing was not finalized. Their teeth are the oldest amongst the small teeth found in west Beringia.
... Changes in body size occurred in other time periods as well, so it has been shown that towards the end of the Pleistocene correlating with warming, mammoths (Vartanyan et al. 1993, Agenbroad et al. 1999, Guthrie 2004, horses (Forsten 1991, Guthrie 2003 and red deer (Lister 1989) decreased in size. There are other examples of size changes on islands (the island rule, Foster 1964): size reduction or dwarfing of large mammals on islands, for example the dwarfed elephants of the Mediterranean Islands (Palombo 2001, Davies & Lister 2001, Raia et al. 2003, or size increase to gigantism of smaller mammals (mainly rodents and leporids) like e.g. Nuralagus rex (Quinata et al. 2011). ...
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The question whether the European wildcat, adapted to cooler climate than other small to medium sized felids, shows changes in body mass or size in response to climate change as indicated for other animals is addressed. The literature yielded body mass data of individual specimens covering the time span from about 1860 to 1960 Also the records of collections were accessed to record weight and body length. These mainly cover the time after 1950 Additionally, three cranial measures, gsl, cbl and zw were measured as indicators of size in the collections representing Germany and Slovakia. Museum records of weight from the last 60 years alone do not show a statistically significant change over time or mean annual temperature. But they do so for body length. The combined data from literature and collections from both regions show a statistically significant decrease in weight over time. As the samples from the literature and museum records represent different time periods, prior to and after 1950, it is difficult to decide if the literature data might be unrealistically high or if there was a real decrease in weight. The German and Slovakian samples differ statistically in the studied parameters, which complicates the picture. Overall the indications of changes in size of wildcats with time or mean annual temperature are not consistent in the studied regions and therefore difficult to assess. Even though there is ample material and substantial literature the collected specimens in the collections do mainly represent relatively short time periods and the available data on weight are also unevenly distributed in time. This supports the necessity to collect large series of specimens over time.
... A revision of the Sicilian and Maltese dwarf elephants is, however, beyond the scope of this research and therefore the current taxonomy is kept consistent with the literature. Boekschoten and Sondaar (1972) and Davies and Lister (2001) mentioned the presence of two pygmy elephants in Cyprus: one is Palaeoloxodon cypriotes and the second is a larger, still unnamed, form. This form, known from the locality Xylophagou (Iliopoulos et al., 2011), might represent a second invasion or an earlier anagenenetic form of P. cypriotes. ...
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During the Late Pleistocene, Naxos and adjacent areas, including Delos and Paros, constituted a mega-island, here referred to as palaeo-Cyclades. The extensive low-lying plains with lakes and rivers provided a suitable habitat for elephants. Due to long-term isolation from the mainland and mainland populations, these elephants evolved miniature size. The species found on Naxos had a body size of about ten percent of that of the mainland ancestor, Palaeoloxodon antiquus. During the glacial periods of the Late Pleistocene, P. antiquus may have migrated eastwards and southwards in search of better conditions and reached the islands. The dwarf species of the various Southern Aegean islands (e.g. Crete, Tilos, Rhodos, palaeo-Cyclades) are each the result of independent colonisation events. The very small size of the Naxos species respective to the dwarf elephants from Crete is explained as due to the lack of competitors. The only other elements of the contemporaneous fauna were a rock mouse (Apodemus cf. mystacinus) and a shrew (Crocidura sp.). Submergence of the area, climate change, volcanism, hunting by humans or a combination of these factors during the terminal Pleistocene may have caused the extinction of this endemic fauna.
... Pohlig (1893) describes and figures a large right tusk, 2 m in length and with a maximum diameter of 137 mm, possessing a clear spiral curvature. A more marked curvature of the tusks, with respect to the condition in P. antiquus, also characterizes P. falconeri and Palaeoloxodon cypriotes (Ambrosetti, 1968;Davies and Lister, 2001). All the other complete tusks from Puntali Cave are smaller than that described by Pohlig, ranging in (axial) length from 1120 to 1380 mm. ...
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... African elephants are highly sexually dimorphic, females weighing around 2800 kg and males around 5000 kg (Kingdon, 1979). At an estimated 7673 kg (Shipman, 1992) the Columbian mammoth, Mammuthus columbi, was significantly larger than an African elephant, and at an estimated 200 kg (Davies and Lister, 2001), Palaeoloxodon cypriotes, the pygmy elephant of Cyprus, was significantly smaller. Age estimates using African elephants are likely to underestimate age in M. columbi and overestimate age in P. cypriotes. ...
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Submitted to: Dept. of Biology. Vol. 2: Plates. Thesis (Ph. D.)--Harvard University, 1971.