Eucladoceros

Eucladoceros (Greek for "well-branched antler") is an extinct genus of large deer whose fossils have been discovered across Eurasia, from Europe to China, spanning from the Late Pliocene-Early Pleistocene. It is noted for its unusual comb-like or branching antlers.

Description
Species of Eucladoceros were large-sized deer. European species lie E. dicranios and E. ctenoides are suggested to have reached a body mass of 250-300 kg, while East Asian E. boulei is suggested to have reached body masses of 350 kg. Body size of European species increased over time. E. giulii has been estimated to have had a shoulder height of 1.55 m while E. senezensis has an estimated shoulder height of 1.7 m. Species of Eucladoceros are noted for their branching antlers, with a large number of tines projecting from the front part of the main antler beam. In many species like Eucladoceros ctenoides the antlers have a comb-like branching pattern, while those of the type species Eucladoceros dicranios has a more complex dichotomous branching pattern. The antlers of E. dicranios are proporotionally large relative to body size, and are among the largest antlers known among deer. The teeth of Eucladoceros species are similar in some aspects to those of the genus Cervus, but lack certain derived characters typical of the teeth of that genus.

Distribution
The first find (E. dicranios) was in 1841 by Florentine naturalist Filippo Nesti, director of the "Museo di Storia Naturale di Firenze" ("Museum of Natural History of Florence"). The earliest species of Eucladoceros was described from the Early Pliocene of China. The most abundant fossil remains of Eucladoceros have come from the Early Pleistocene of Europe and China.

The systematics of European forms is confused and up to twelve poorly defined species are reported. The majority of those species names are synonymous, and at present only two or three good species are recognized: E. dicranios from England, Italy and Azov Sea Area in South Russia; E. ctenoides from Greece, Italy, France, Spain, the Netherlands, and England; and E. teguliensis (a senior synonym of E. senezensis) from France, the Netherlands, and Britain. The latter species is regarded by some authors as a subspecies of E. ctenoides, since there are some finds (for instance, from Ceyssaguet, France) that show a transitional character between E. ctenoides and E. teguliensis.

Some poor remains of Eucladoceros are found also in Tajikistan, Pakistan, and India.

Ecology
Analysis of the limbs of Eucladoceros suggest that they are most similar to living deer that occupy open habitats. Dental microwear analysis of Eucladoceros ctenoides suggests that its diet was largely plastic and widely varied according to local conditions.

Taxonomy
The genus was formally described by Hugh Falconer in 1868. Eucladoceros is widely agreed to a be member of the tribe Cervini, though its placement within this group has been debated. A 2017 study analysing the bony labyrinth suggested that Eucladoceros was most closely related to the living genera Rusa (sambar) and Cervus (red deer, elk, sika deer). Some authors have proposed a close relationship to the extinct "giant deer" genus Praemegaceros.

Species

 * Eucladoceros boulei Marcellin Boule (1928), Age: Latest Pliocene - Early Pleistocene; Nihowan, China
 * Eucladoceros ctenoides (former name E. teguliensis) F. Nesti (1841), Age: Early Pleistocene, Late Villafranchian; Locus typicus: Upper Valdarno, Tuscany, Italy
 * Eucladoceros dichotomus (Original citation: Cervus (Elaphurus) dichotomus Teilhard de Chardin & Piveteau; Early Pleistocene of Nohowan; most probably is not a Eucladoceros species)
 * Eucladoceros dicranios Filippo Nesti (1841), Age: Early Pleistocene, Late Villafranchian; Upper Valdarno, Tuscany, Italy. Note: the type species of the genus.
 * Eucladoceros proboulei (Dong Wey), Age: Early Pliocene; China
 * Eucladoceros senezensis Charles Depéret, 1910, Senèze (Haute-Loire), near Brioude, France. Note: some authors regard it as a subspecies of E. ctenoides.
 * Eucladoceros tetraceros Sir Wm. Boyd Dawkins (1878), Age: Early Pleistocene; Peyrolles, Haute-Loire, France. Note: a possible synonym of E. ctenoides.

Literature

 * Azzaroli, A. 1954. "Critical observations upon Siwalik deer". The Proceedings of the Linnean Society of London, 165: 75-83, London.
 * Azzaroli, A. & Mazza, P. 1992. "The cervid genus Eucladoceros in the early Pleistocene of Tuscany". Palaeontographia Italica, 79: 43-100; Pisa.
 * Croitor R. & Bonifay M.-F. 2001. "Étude préliminaire des cerfs du gisement Pleistocène inférieur de Ceyssaguet (Haute-Loire)". Paleo, 13: 129-144.
 * Dong W. & Ye J. 1996. "Two new cervid species from the late Neogene of Yushe Basin, Shanxi Province, China". Vertebrata PalAsiatica, 34 (2): 135-144.
 * Heintz E. 1970. "Les Cervides Villafranchiens de France et d’Espagne". Memoires du Museum national d’histoire naturelle. Ser.C, Sc. De la Terre, 22 (1-2): 1-302, Paris.
 * Vos, J. De, Mol D. & Reumer J. W. F.1995. "Early pleistocene Cervidae (Mammalia, Artyodactyla) from the Oosterschelde (the Netherlands), with a revision of the cervid genus Eucladoceros Falconer", 1868. Deinsea, 2: 95-121.