Candiacervus
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''Candiacervus'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
deer Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...
native to
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
Crete Crete ( el, Κρήτη, translit=, Modern: , Ancient: ) is the largest and most populous of the Greek islands, the 88th largest island in the world and the fifth largest island in the Mediterranean Sea, after Sicily, Sardinia, Cyprus, and ...
. Due to a lack of other herbivores, the genus underwent an
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
, filling niches occupied by other taxa on the mainland. Due to the small size of Crete, the genus underwent
insular dwarfism Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is disti ...
, the smallest species, ''C. ropalophorus'', stood about 40 cm at the shoulders when fully grown, as can be inferred from a mounted skeleton. 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. They were traditionally considered to be related to the giant
Irish elk The Irish elk (''Megaloceros giganteus''), also called the giant deer or Irish deer, is an extinct species of deer in the genus ''Megaloceros'' and is one of the largest deer that ever lived. Its range extended across Eurasia during the Pleisto ...
, with some experts regarding ''Candiacervus'' as a
subgenus In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
of ''
Megaloceros ''Megaloceros'' (from Greek: + , literally "Great Horn"; see also Lister (1987)) is an extinct genus of deer whose members lived throughout Eurasia from the early Pleistocene to the beginning of the Holocene and were important herbivores durin ...
''. However, van der Geer (2018) finds them closer to ''Dama''.


Taxonomy

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 Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
. This obscures taxonomy, because many endemic features of ''Candiacervus'' are not unique but are found in other island deer as well, such as ''
Cervus astylodon ''Cervus astylodon'', the Ryukyu dwarf deer, is a recently extinct species of cervid that was endemic to the Ryukyu islands (Okinawa, Ishigaki Island, Ishigaki, Kume, Tokunoshima). It lived throughout the Pleistocene, going extinct as recently as ...
'' (Ryukyu Islands) and ''Hoplitomeryx'' (Southern Italy). De Vos (1979, 1984, 1996) identified eight morphotypes into one genus (''Candiacervus''), whereas Capasso Barbato (1992) 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''. On the nearby island of
Karpathos Karpathos ( el, Κάρπαθος, ), also Carpathos, is the second largest of the Greek Dodecanese islands, in the southeastern Aegean Sea. Together with the neighboring smaller Saria Island it forms the municipality of Karpathos, which is part o ...
, Kuss 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

The Cretan deer is represented by no less than eight different
morphotype In biology, polymorphism is the occurrence of two or more clearly different morphs or forms, also referred to as alternative ''phenotypes'', in the population of a species. To be classified as such, morphs must occupy the same habitat at the s ...
s, ranging from dwarf size with
withers The withers is the ridge between the shoulder blades of an animal, typically a quadruped. In many species, it is the tallest point of the body. In horses and dogs, it is the standard place to measure the animal's height. In contrast, cattle ar ...
height of about 40 cm to very large with withers height of about 165 cm. This is explained as a
sympatric speciation Sympatric speciation is the evolution of a new species from a surviving ancestral species while both continue to inhabit the same geographic region. In evolutionary biology and biogeography, sympatric and sympatry are terms referring to organi ...
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 Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweight s ...
. 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 '' Pseudodama peloponnesiacus'' and ''
Praemegaceros ''Praemegaceros'' is an extinct genus of deer, known from the Pleistocene and Holocene of Western Eurasia. It contains the subgenera ''Praemegaceros,'' ''Orthogonoceros'' and ''Nesoleipoceros''. It has sometimes been synonymised with ''Megalocer ...
verticornis''.


Ecology

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 In biostratigraphy, biostratigraphic units or biozones are intervals of geological strata that are defined on the basis of their characteristic fossil taxa, as opposed to a lithostratigraphic unit which is defined by the lithological properties of ...
are the common mouse (''Mus bateae'', ''M. minotaurus''), the dwarf hippo (''
Hippopotamus creutzburgi ''Hippopotamus creutzburgi'', the Cretan dwarf hippopotamus, is an extinct species of hippopotamus from the island of Crete. ''Hippopopotamus'' colonized Crete probably 800,000 years ago and lived there during the Middle Pleistocene. Bones of ''H ...
'') the
dwarf elephant Dwarf elephants are prehistoric members of the order Proboscidea which, through the process of allopatric speciation on islands, evolved much smaller body sizes (around ) in comparison with their immediate ancestors. Dwarf elephants are an example ...
(''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 The Cretan otter ''(Lutrogale cretensis)'' is an extinct otter that was endemic to Crete during the Pleistocene. Taxonomy It was a close relative of the smooth-coated otter ''(L. perspicillata)'', whom today lives only in southern Asia but had a ...
(''Lutrogale (Isolalutra) cretensis''), and the Cretan shrew (''Crocidura zimmermanni'').


Extinction

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. 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 The Paleolithic or Palaeolithic (), also called the Old Stone Age (from Greek: παλαιός ''palaios'', "old" and λίθος ''lithos'', "stone"), is a period in human prehistory that is distinguished by the original development of stone too ...
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
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
, as indicated by the finding of a complete herd consisting of individuals suffering a bone disease of an
osteosclerotic Osteosclerosis is a disorder that is characterized by abnormal hardening of bone and an elevation in bone density. It may predominantly affect the medullary portion and/or cortex of bone. Plain radiographs are a valuable tool for detecting and ...
nature (see X-ray photograph).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. PDF
/ref>_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.


__References_


__External_links_


Article_on_the_mounting_of_a_''Candiacervus''_skeleton

Scientific_information_on_''Candiacervus''



New_data_on_the_Pleistocene_Cretan_deer_Candiacervus_sp._II_(Cervinae,_Mammalia)
(2006) {{Taxonbar.html" ;"title="n Greek with English summary
PDF
/ref> 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.


References


External links


Article on the mounting of a ''Candiacervus'' skeleton

Scientific information on ''Candiacervus''



New data on the Pleistocene Cretan deer Candiacervus sp. II (Cervinae, Mammalia)
(2006) {{Taxonbar">from=Q3655192 Prehistoric deer Pleistocene even-toed ungulates Pleistocene mammals of Europe Prehistoric even-toed ungulate genera Prehistoric Crete