The effect of insularity on the five-horned deer Hoplitomeryx (Late Miocene, Italy)

Island studies increase our understanding of the effects of habitat fragmentation. The study of the Tertiary paleo-island Gargano is an important contribution, because of the long-term isolation under less fluctuating climatic conditions, free from anthropogenic influences; such a situation does not exist in the Quaternary period nor in the Holocene period. This makes the Gargano a unique case to study the effects of insularity in isolation. Here, a highly endemic, unbalanced vertebrate fauna evolved including the five-horned deer Hoplitomeryx. Its post-cranial material contains four size groups, based on the metapodals. In this study, the humerus and radius are described. The question whether the morphotypes are chronomorphs or ecomorphs is addressed. Sexual dimorphism is ruled out as the underlying principle of size separation in this case, based upon body mass estimations and data from living deer. Chronomorphs is the best explanation for the Megaloceros cazioti lineage (Pleistocene, Sardinia) and the Myotragus balearicus lineage (Pliocene–Holocene, Mallorca). Ecomorphs are a better explanation for the size groups of Candiacervus (Late Pleistocene, Crete) and Cervus astylodon (Late Pleistocene, Ryukyu Islands, Japan). An adaptive radiation into several trophic types took place, promoted by the ecological meltdown of the ancestral niche. The drive behind this speciation is increased interspecific competition. For Hoplitomeryx, although the hypothesis of chronomorphs cannot be discarded, that of ecomorphs seems most likely, based upon the coexistence of two or more size groups per fissure, and upon the presence of a huge morphotype, larger than mainland species, in the younger fissures.

Read more in VAN DER GEER A.A.E. (2008). The effect of insularity on the Eastern Mediterranean early cervoid Hoplitomeryx: the study of the forelimb. Quaternary International 182, 1: 145-159. See http://dx.doi.org/10.1016/j.quaint.2007.09.021 or ask me a pdf (geeraae@geol.uoa.gr).

For more general information of this enigmatic Late Miocene 'deer', see my Wikipedia page at http://en.wikipedia.org/wiki/Hoplitomeryx

The postcranial of the deer Hoplitomeryx (Mio-Pliocene; Italy): another example of adaptive radiation on Eastern Mediterranean Islands.

During the Late Miocene a highly endemic vertebrate fauna evolved on Gargano Island (south-east coast of Italy), comprising others the giant soricid Deinogalerix, the giant barn owl Tyto gigantea, the giant hamster Hattomys, and the 'prongdeer' Hoplitomeryx with five horns and sabrelike ('moschid' type) upper canines. The Hoplitomeryx skeletal material forms a heterogenous group, containing four size groups; within the size groups different morphotypes may be present. All size groups share the same typical Hoplitomeryx features. These are: one central nasal horn and a pair of pronged orbital horns, protruding canines, complete fusion of the navicocuboid with the metatarsal, distally closed metatarsal gully, a non-parallel-sided astragalus, and an elongated patella. The different size groups are equally distributed over the excavated fissures, and are therefore not to be considered chronotypes. The hypothesis of an archipelago consisting of different islands each with its own morphotype cannot be confirmed.
The situation with several co-existing morphotypes on an island is paralleled by Candiacervus (Late Pleistocene, Crete, Greece). Opinions about its taxonomy differ, and at present two models prevail: one genus for eight morphotypes, or alternatively, two genera for five species. The second model is based upon limb proportions only, but these are invalid taxonomic features for island endemics, as they change under influence of environmental factors that differ from the mainland. Also in Hoplitomeryx the morphotypes differ in limb proportions, but here different ancestors are unlikely, because in that case they all ancestors must have shared the typical hoplitomerycid features. The morphosphere of Hoplitomeryx is too coherent to assume two or more different ancestors, and indicates a monophyletic origin of all morphotypes.
The large variation is instead explained as an example of adaptive radiation, starting when the Miocene ancestor colonized the island. The range of empty niches promoted its radiation into several trophic types, yielding a differentiation in Hoplitomeryx. The shared lack of large mammalian predators and the limited amount of food in all niches promoted the development of derived features in all size groups (apomorphies).

For full text, see VAN DER GEER, A.A.E. (2005). The postcranial of the deer Hoplitomeryx (Mio-Pliocene; Italy): another example of adaptive radiation on Eastern Mediterranean Islands.van der Geer. Monografies de la Societat d'Història Natural de les Balears 12: 325-336. Palma de Mallorca. For a free pdf [1,018 kb]: http://users.uoa.gr/~geeraae/publications/2005-IMEDEA-Hoplitomeryx.pdf. See my website http://users.uoa.gr/~geeraae for three more publications on this bizarre and enigmatic insular 'deer' of the Late Miocene.

For more general information of this enigmatic Late Miocene 'deer', see my Wikipedia page at http://en.wikipedia.org/wiki/Hoplitomeryx

New data on the Late Pleistocene Cretan deer Candiacervus sp. II

For our museum, we mounted a skeleton of the endemic Late Pleistocene Cretan deer Candiacervus sp. II (Liko Cave), using bones of different individuals. This composite skeleton contributes to the study of taxonomy of insular ungulates as it reveals some additional features that were not detected in the isolated elements. Candiacervus sp. II differs from all known recent and extinct mainland deer, mainly in its proportions. Although its considerably shortened distal limbs were already noted in the past, Candiacervus sp. II now appears at the same time to have had a more or less normal vertebral column length relative to continental large deer, and moderately upwards curved lumbar section, both features reminding us more of the insular dwarf bovid Myotragus than of the mainland small deer Axis axis. Combined with an increased massivity of all bones and pronounced muscle scars, this change in body proportions appears to indicate that Candiacervus sp. II evolved towards the niche of goat-like bovids in rocky environments. Other additional diagnostic features are the horizontally directed transversal processus of the vertebras, fusion of the lateral metacarpal to the main metacarpal, a tail length of ten vertebras, a more pronounced difference between anterior and posterior hooves, and the presence of lateral toes upto the third phalanx, anterior as well as posterior.

Read more in VAN DER GEER A.A.E., DE VOS J., LYRAS G.A., DERMITZAKIS M.D. (2006). New data on the Pleistocene Cretan deer Candiacervus sp. II (Mammalia, Cervinae). Courier Forschungsinstitut Senckenberg 256: 131-137. For a pdf, send an e-mail to geeraae@geol.uoa.gr. For more Candiacervus publications, visit our websites at http://users.uoa.gr/~geeraae and http://users.uoa.gr/~glyras

For more general info on the extinct Cretan deer, see http://en.wikipedia.org/wiki/Candiacervus