Candiacervus
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| Candiacervus Fossil range: Pleistocene |
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 Athene cretensis and Candiacervus ropalophorus
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Prehistoric
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Candiacervus was a genus of deer native to Pleistocene Crete. Their most notable feature, besides their peculiar, spatula-shaped antlers, was their small stature: the smallest species, C. ropalophorus, stood about 40 cm at the shoulders when fully grown. As such, the genus is considered to be a textbook example of insular dwarfism.
Ironically, they were closely related to the giant Irish Elk, with some experts regarding Candiacervus as a subgenus of Megaloceros (Capasso Barbato 1992b).
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[edit] Large variation
The Cretan deer is represented by no less than eight different morphotypes, ranging from dwarf size with withers height of about 40 cm to very large with withers height of about 165 cm. This is explained as a sympatric speciation to occupy all possible empty niches ranging from dense forest to prickly rocks. The coexistence of various environments has been confirmed by studies on the rich fossil avifauna. The most typical Cretan deer are the two smallest sizes, which have not only relatively and absolutely short limbs, but also long and simplified antlers; these species occupied a niche close to that of the wild goat of Crete today: barren rocks with thorny bushes, as shown by features of their osteology and goat-like body proportions. It deviated so much from mainland deer that it is impossible to indicate with certainty its ancestor. Suggested ancestors are Cervus peloponnesiacus and Megaloceros verticornis.
[edit] Taxonomical problems
The Cretan deer is a typical example of taxonomical problems involving endemic insular mammals, due to the much larger variety than on the mainland, and the strong endemism, which obscures taxonomy. De Vos (1979, 1984, 1996) includes the eight morphotypes into one genus (Candiacervus), whereas Capasso Barbato (1992) does not follow this opinion of monophyly, and included the larger species, rethymnensis, major and dorothensis, in Cervus (subgenus Leptocervus) and the smaller species ropalophorus and cretensis in Megaloceros (subgenus Candiacervus), which implies two different ancestors. She also does not recognize sp. II with its three morphotypes, and synonymises them with ropalophorus. At present these two views cannot be properly proven or discarded, but what stays is that the number of deer species is higher than on the mainland, and that they all occupy a different ecological niche.
[edit] Extinction of Candiacervus
![Ill limb bone of Candiacervus from the cave Mavro Muri. See Dermitzakis M., Van der Geer AAE, Lyras G. 2006. Palaeopathological observations on a population of fossil deer from the Late Pleistocene of Crete. In: Kalofourtis, A., Papadopoulos, N., Spiliopoulou, C., Marabellas, K., Chatzioannou, A.. Volume in Honor of Prof. A.S. Koutselinis, pp. 43-51. [in Greek with English summary].](../../../../images/local/thumb/1/1c/Candiacervus-pathology.gif/180px-Candiacervus-pathology.gif)
From the late Middle Pleisocene till the arrival of humans in the Holocene, Crete was inhabited by small elephants, eight types of Cretan deer and a normal sized mouse. The cause of the dramatic faunal turnover, which led to the extinction of the endemic deer and elephants, may simply have been the arrival of paleolithic humans. They could have exterminated the deer either actively by hunting, or passively by destroying its habitat. Another option is a gradual depletion of the eco-system, as indicated by the finding of a complete herd consisting of individuals suffering a bone disease of an osteosclerotic nature. The impact of paleolithic humans is at present still unproven, partly because of the scarcity on published fauna lists from archaeological sites (except for Knossos), partly because of the insecurely dated materials
[edit] Fauna with Candiacervus
The fauna of which Candiacervus is an element, is called Biozone II, or the Mus Zone (after the common mouse). This fauna inhabited Crete between the late Middle and Late Pleistocene, which means between 0.3 and 0.01 million years ago.
The typical fauna elements of this biozone are the common mouse (Mus bateae, M. minotaurus), the dwarf elephant (Elephas antiquus creutzburgi), the Cretan deer (Candiacervus, with the eight species ropalophorus, sp. IIa, b and c, cretensis, rethymnensis, dorothensis and major), the Cretan otter (Lutrogale (Isolalutra) cretensis), and the Cretan shrew (Crocidura zimmermanni) . The common mouse is represented by two species, of which the earlier is M. bateae, which is slightly smaller than the later M. minotaurus, and slightly larger than the common house mouse (Mus musculus). The two species belong to a single lineage with a long term trend for increasing size is attested. The Cretan shrew is one of the rare exceptions to the general rule that mammals change radically on islands. On most islands, the microfauna show the tendency to become large, as is the case of the Cretan mouse of the previous biozone. It is as yet not clear why this did not apply to the Cretan shrew. The Cretan shrew was obviously successful, and was not wiped out during the next faunal turnover, and managed to survive on Crete till the present day . The dwarf elephant may be large compared to the mammoth of the previous period, but it is still about 30% smaller than its mainland ancestor E. antiquus, which has a shoulder height of 3.7 m. The dwarf elephant has strongly curved tusks. It is still a matter of debate why this elephant did not reach a true pygmy size. The Cretan otter is the only carnivore known from the Pleistocene of Crete. Its remains are known from only one locality (Liko Cave), and only in the upper layer thereof (Late Pleistocene). The Cretan otter was less aquatic in life style than the common otter (Lutra lutra) and the smooth otter Lutrogale perspicillata; this is considered a secondary development due to the special conditions on Crete. At first view it may seem strange that the Cretan otter did not develop towards gigantism as in rodents or towards dwarfism as in herbivores, though it seems that it is slightly larger than its mainland ancestor. However, this gigantism and dwarfism is caused by common factors as the absence of predators and limited food resources; for otters these factors are not relevant.
[edit] Karpathos
On the nearby island of Karpathos, Kuss (see references) found deer which were, in his view, similar to the Cretan deer. Therefore, he grouped his species pygadiensis and cerigensis under the genus Candiacervus, but this needs further confirmation. As long as no direct link with Crete is attested, the deer genus of Karpathos is questioned, and better referred to as Cervus.
[edit] References
- Capasso Barbato L. 1992b. Observations on the biostratigraphy of Cretan Pleistocene vertebrates. Il Quaternario 5 (1), 67-76.
- De Vos, J. 1979. The endemic Pleistocene deer of Crete. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, Series B 82 (1), 59-90.
- De Vos J. 1984. The endemic Pleistocene deer of Crete. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, afd. Natuur¬kunde, eerste reeks 31. North Holland Publishing Compa¬ny. Amsterdam, Oxford, New York. 100 pp.
- De Vos, J. 1996. Taxonomy, Ancestry and Speciation of the Endemic Pleistocene Deer of Crete Compared with the Taxonomy, Ancestry and Speciation of Darwin's Finches. In: Reese, 1996, 111-24.
- Dermitzakis, M.D. & J. De Vos 1987. Faunal Succession and the Evolution of Mammals in Crete during the Pleistocene. Neues Jahrbuch Geologischer und Paläontologischer Abhandlungen 173 (3), 377-408.
- Dermitzakis, M.D. & P.Y. Sondaar 1978. The importance of fossil mammals in reconstruction paleogeography with special reference to the Pleistocene Aegean Archipelago. Annales Géologiques des Pays Helléniques 46, 808-40.
- Dermitzakis, M.D. 1977. The quaternary fossil mammals in Caves and Karstic Holes of Crete Island and their Significance. Bulletin de la Société Spéléologique de Grèce 14, 152-90.
- Kuss S.E., 1975. Die pleistozänen Hirsche der ostmediterranen Inseln Kreta, Kasos, Karpatos und Rhodos (Griechenland). Berichte der Naturforschenden Gesellschaft zu Freiburg im Breisgau, 65 (1 2), 25-79
- Mayhew, D.F. 1977. The endemic Pleistocene murids of Crete I-II. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen B, 80 (3), 182-214.
- Reese, D.S., ed. 1996. Pleistocene and Holocene Fauna of Crete and its First Settlers. Monographs in World Archeology 28. Madison, Prehistoric Press.
- Reumer, J.W.F. 1986. Notes on the Soricidae (Insectivora, Mammalia) from Crete. II. The shrew remains from Minoan and Classical Kommos. Bonner zoologische Beiträge 37 (3), 173-82.
- Van der Geer, A.A.E., De Vos, J., Lyras, G., Dermitzakis, M. (2006). New data on the Pleistocene Cretan deer Candiacervus sp. II (Mammalia, Cervinae). In: Kahlke, R.-D., Maul, L. C. & Mazza, P. (Eds.): Late Neogene and Quaternary biodiversity and evolution: Regional developments and interregional correlations. Proceedings of the 18th International Senckenberg Conference (VI International Palaeontological Colloquium in Weimar) vol. II. Courier Forschungsinstitut Senckenberg 256, 131-137. Frankfurt am Main.
- Van der Geer, A.A.E., Dermitzakis, M., De Vos, J. 2006. Crete before the Cretans: the reign of dwarfs. Pharos 13, 121-132. Athens: Netherlands Institute.
- Van der Geer, A.A.E., Dermitzakis, M. 2006. Relative growth of the metapodals in a juvenile island deer: Candiacervus (Mammalia, Cervidae) from the Pleistocene of Crete. Hellenic Journal of Geosciences (formerly Annales Géologiques de Pays Helleniques) 41 (1), 119-125.
- Van der Geer, A.A.E., De Vos, J., Lyras, G., Dermitzakis, M. 2005. The
mounting of a skeleton of the fossil species Candiacervus sp. II from Liko Cave, Crete, Greece. Monografies de la Societat d'Història Natural de les Balears 12, 337-346.
