Candiacervus

Candiacervus is an extinct genus of deer native to Pleistocene Crete.[1] Due to a lack of other herbivores, the genus underwent an adaptive radiation, filling niches occupied by other taxa on the mainland. Due to the small size of Crete, the genus underwent insular dwarfism[2], the smallest species, C. ropalophorus, stood about 40 cm at the shoulders when fully grown, as can be inferred from a mounted skeleton.[3] Some species (C. ropalophorus) are noted for their peculiar, spatula-shaped antlers, though other species have normal albeit miniaturized antlers. Other features are the relatively short limbs, the massivity of the bones and the simplified antler.[4]

Candiacervus
Temporal range: Pleistocene
Candiacervus cretensis antler
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Family: Cervidae
Genus: †Candiacervus
Kuss, 1975
Species
  • †Candiacervus ropalophorus de Vos, 1984
  • †Candiacervus cretensis (Simonelli, 1907) (type)
  • †Candiacervus dorothensis (Capasso Barbato, 1990)
  • †Candiacervus rethymensis Kuss, 1975
  • †Candiacervus major (Capasso Barbato and Petronio, 1986)
  • †Candiacervus devosi van der Geer, 2018
  • †Candiacervus listeri van der Geer, 2018
  • †Candiacervus reumeri van der Geer, 2018

They were traditionally considered to be related to the giant Irish elk, with some experts regarding Candiacervus as a subgenus of Megaloceros.[5] However van der Geer (2018) finds them closer to Dama.[6]

Taxonomy
Hippopotamus creutzburgi and C. cretensis

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. This obscures taxonomy, because many endemic features of Candiacervus are not unique but are found in other island deer as well,[7] such as Cervus astylodon (Ryukyu Islands) and Hoplitomeryx (Southern Italy). De Vos (1979, 1984, 1996)[8][9][10] identified eight morphotypes into one genus (Candiacervus), whereas Capasso Barbato (1992)[11] included the larger species, rethymnensis, major and dorothensis, in Cervus (subgenus Leptocervus) and the smaller species ropalophorus and cretensis in Megaloceros (subgenus Candiacervus), implying two different ancestors, and she also did not recognize sp. II with its three morphotypes, instead referring it to ropalophorus. A new paper published in 2018 rejected the conclusion of Capasso Barbato (1992) and formally named the three morphotypes of De Vos' Candiacervus sp. II C. devosi, C. listeri, and C. reumeri.[12]

On the nearby island of Karpathos, Kuss[13] 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.

Description
Athene cretensis and C. ropalophorus

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.[14] 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,[15] 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 Pseudodama peloponnesiacus and Praemegaceros verticornis.

Ecology

The fauna of which Candiacervus is an element is called Biozone II, or the Mus Zone (after the common mouse).[16] This fauna inhabited Crete between the late Middle and Late Pleistocene, which means between 0.3 and 0.01 million years ago.[17]

The typical fauna elements of this biozone are the common mouse (Mus bateae, M. minotaurus), the dwarf hippo (Hippopotamus creutzburgi) 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).

Extinction
Ill limb bone of Candiacervus from the cave Mavro Muri, showing an osteosclerotic pathology.

From the late Middle Pleistocene till the arrival of humans in the Holocene, Crete was inhabited by small elephants, eight types of Cretan deer and a normal sized mouse.[18] 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.[19] They could have exterminated the deer either actively by hunting, or passively by destroying its habitat. Another option is a gradual depletion of the ecosystem, as indicated by the finding of a complete herd consisting of individuals suffering a bone disease of an osteosclerotic nature (see X-ray photograph).[20] 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.

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References
  1. 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.PDF
  2. Van der Geer, A.A.E., De Vos, J., Dermitzakis, M., Lyras, G., 2009. Hoe dieren op eilanden evolueren. Majorca, Ibiza, Kreta, Sardiniie, Sicilie, Japan, Madagaskar, Malta. Utrecht: Veen Magazines; ISBN 978-90-8571-169-8.Ga naar Bruna Archived 2013-09-28 at the Wayback Machine
  3. 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
  4. 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.
  5. Capasso Barbato L. 1992b. Observations on the biostratigraphy of Cretan Pleistocene vertebrates. Il Quaternario 5 (1), 67-76.
  6. van der Geer, Alexandra (2018). "Uniformity in variety: Antler morphology and evolution in a predator-free environment". Palaeontologia Electronica: 1–31. doi:10.26879/834.
  7. Van der Geer, A.A.E. 2005. Island ruminants and the evolution of parallel functional structures. In: Cregut, E. (Ed.): Les ongulés holarctiques du Pliocène et du Pléistocène. Actes Colloque international Avignon, 19-22 septembre. Quaternair, 2005 hors-série 2: 231-240. Paris. PDF
  8. De Vos, J. 1979. The endemic Pleistocene deer of Crete. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, Series B 82 (1), 59-90
  9. 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.
  10. 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.
  11. Capasso Barbato L. 1992b. Observations on the biostratigraphy of Cretan Pleistocene vertebrates. Il Quaternario 5 (1), 67-76.
  12. van der Geer, A.A.E. (2018). "Uniformity in variety: Antler morphology and evolution in a predator-free environment". Palaeontologia Electronica (21.1.9A): 1–31. doi:10.26879/834.
  13. 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.
  14. 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.
  15. 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. link PDF
  16. 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.
  17. 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.
  18. Reese, D.S., ed. 1996. Pleistocene and Holocene Fauna of Crete and its First Settlers. Monographs in World Archeology 28. Madison, Prehistoric Press.
  19. Sondaar, P.Y., Van der Geer, A.A.E. 2005. Evolution and Extinction of Plio-Pleistocene Island Ungulates. In: Cregut, E. (Ed.): Les ongulés holarctiques du Pliocène et du Pléistocène. Actes Colloque international Avignon, 19-22 septembre. Quaternair, 2005 hors-série 2: 241-256. Paris.
  20. 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] PDF
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