Remarks on some Eurasian pliocervines: Characteristics, evolution, and relationships with the tribe...

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Geobios 40 (2007) 113–130

Article original

Remarks on some Eurasian pliocervines: Characteristics, evolution,

and relationships with the tribe Cervini

Remarques sur quelques pliocervines asiatiques et europeennes :

caracteristiques, evolution et rapports avec les cervines

Carmelo Petronio a,*, Tatiana Krakhmalnaya b, Luca Bellucci c, Giuseppe Di Stefano a

a Dipartimento di Scienze della Terra, Universita degli studi di Roma ‘‘La Sapienza’’, P.le A. Moro, 5, 00185 Roma, Italyb National Museum of Natural History of National Academy of Sciences of Ukraine, Kiev, Ukraine

c Dottorato di Paleontologia, Universita degli studi di Roma ‘‘La Sapienza’’, P.le A. Moro, 5, 00185 Roma, Italy

Received 11 July 2005; accepted 16 January 2006

Available online 17 January 2007

Abstract

In order to define the ancestors of the Eurasian genera Axis and Rusa, the representatives of the tribe Pliocervini have been reviewed, with the

exclusion of the genus Pliocervus. The fossil remains referred to Pliocervini stored in the Palaeontological Museum of Mechnikov Odessa National

University and studied by Khomenko [Khomenko, I., 1913. Meotian fauna of v. Tarakliya of Bendery district. Annuaire geologique et

mineralogique de la Russie. 1. The ancestors of modern and fossil Cervinae. 2. Giraffidae and Cavicornia 15, 107–132 (in Russian)] are

reappraised: these remains are attributed to Cervavitus novorossiae (Khomenko, 1913). The fossil deer collected from the locality of

Novoelizavetovka (Odessa region, Ukraine) and observed in the same Museum of the Odessa National University, is referred to Cervavitus

variabilis (Alexeev, 1915).The fossil material from Shanxi and Honan provinces (China), described by Zdansky [Zdansky, O., 1925. Fossil Hirsche

Chinas. Palaeontologia Sinica C 2, 1–94] and Teilhard de Chardin and Trassaert [Teilhard de Chardin, P., Trassaert, M., 1937. The Pliocene

Camelidae, Giraffidae and Cervidae of southeastern Shansi.] is referred to Cervavitus shanxius Dong and Hu, 1994. The phylogenetic relationships

between Pliocervini from China and the European ones are investigated on the basis of the morphological, biochronological and paleogeographical

data. These elements and the suggestions of the more recent genetical and biometrical data may indicate that the origin of the two genera Axis and

Rusa could be recognized into the Chinese pliocervines.

# 2006 Elsevier Masson SAS. All rights reserved.

Resume

Dans cette etude sont revises les restes fossiles des pliocervines observes pour la premiere fois par Khomenko [Khomenko, I., 1913. Meotian

fauna of v. Tarakliya of Bendery district. Annuaire geologique et mineralogique de la Russie. 1. The ancestors of modern and fossil Cervinae. 2.

Giraffidae and Cavicornia 15, 107–132 (en russe)] et places au Museum de l’universite Mechnikov d’Odessa; ces restes sont attribues a Cervavitus

novorossiae; le materiel fossile appele Procervus (= Cervavitus) variabilis par (Alexeev, 1915) provenant de la localite de Novoelizavetovka

(Odessa Region, Ukraine) et depose dans le meme Museum de l’universite d’Odessa est appele Cervavitus variabilis. Le materiel fossile provenant

des provinces de Shanxi et Honan (Chine), decrit par Zdansky [Zdansky, O., 1925. Fossil Hirsche Chinas. Palaeontologia Sinica C 2, 1–94.] et

Teilhard de Chardin et Trassaert [Teilhard de Chardin, P., Trassaert, M., 1937. The Pliocene Camelidae, Giraffidae and Cervidae of southeastern

Shansi.], et observe dans le Museum de Paleontologie de Uppsala (Suede) est rapporte aux sous-especes Cervavitus shanxius shanxius et

C. shanxius minor. Les rapports phylogenetiques entre les pliocervines de la Chine et de l’Europe sont etudies en fonction des elements

morphologiques, biochronologiques et paleogeographiques. Les memes elements ainsi que les apports des nouvelles etudes biometriques et

genetiques, peuvent indiquer que les genres Axis et Rusa ont leur origine au sein des pliocervines de Chine.

# 2006 Elsevier Masson SAS. All rights reserved.

Keywords: Mammalia; Cervavitus; Pliocervines; Miocene; Eurasia

Mots cles : Mammifere ; Cervavitus ; Pliocervines ; Miocene ; Eurasie

* Corresponding author.

E-mail address: [email protected] (C. Petronio).

0016-6995/$ – see front matter # 2006 Elsevier Masson SAS. All rights reserved.

doi:10.1016/j.geobios.2006.01.002

mailto:[email protected]

http://dx.doi.org/10.1016/j.geobios.2006.01.002

C. Petronio et al. / Geobios 40 (2007) 113–130114

1. Introduction

From the Middle Pliocene to the Early Pleistocene,

according to Di Stefano and Petronio (2003) some medium-

sized deer belonging to different genera, spread into continental

and mediterranean Europe: more particularly the genus Rusa is

found throughout the habitats of the northern areas while the

genus Axis is prevalent in the southern areas and in the Italian

peninsula. These cervids occur in large areas and within a

relatively short time span and they are very important for the

biochronological studies of the Plio-Pleistocene mammal

assemblages.

This paper focuses on the study of the Asian and European

Mio-Pliocene pliocervines and also attempts to draw a

phylogenetic hypothesis which links these deer with the

Plio-Pleistocene European genera and species. Tribe Pliocer-

vini in fact represents an important evolutionary passage

between the Middle Miocene Dicrocerini and the Plio-

Pleistocene Cervini (Bubenik, 1990).

The members of this group of extinct deer are particularly

important for any phylogenetic and systematic purposes,

because they have many morphological features that become

subsequently widespread in the more advanced cervids. The

genera and species belonging to the pliocervines show a wide

geographical distribution. They are spread out over the area

covering most of Eurasia, excluding only the artic zones and

the Indian subcontinent. From a biochronological point of view,

the representatives of pliocervines were diffused from the Late

Miocene to the Early Pleistocene.The main purposes of the

following analysis are:

� to
give a systematic reappraisal of the fossil material which
allowed the first description of the subfamily Pliocervinae by

Khomenko (1913) (a tribe since 1968);

� to
compare the forms described by Khomenko (1913) with
the deer remains referred to Pliocervini from other European

faunal assemblages;

� to
review the Chinese fossil material referred to pliocervines
by Zdansky (1925) and by Teilhard de Chardin and Trassaert

(1937);

� to
show the morphological and phylogenetic relationships
between the deer from China, identified on the base of above-

mentioned material and the European ones;

� to
draw an hypothesis on the possible ancestors of the more
evolved Plio-Pleistocene European deer among pliocervines.

The morphological analyses, discussions, and conclusions of

the present paper follow the observation of the species referred to

the tribe Pliocervini with the exclusion of those referred to the

genus Pliocervus; these last species in fact, even if considered to

be part of Pliocervini, do not seem to have any phylogenetic

relationship with the representatives of the tribe Cervini.

2. Historical review of the systematics of Pliocervini

The number of species that belong to Pliocervini and their

characteristics are still not fully known, these uncertainties

dating back to the author who instituted this tribe (see later)

(Vislobokova, 1983; Korotkevich, 1988; McKenna and Bell,

1997). The aim of the following chapter is to draw the history of

the taxonomy and the different opinions on these deer on the

basis of the most important and available sources in literature,

even if not the most complete. We would also like to show how

the opinions on the systematic position of pliocervines into the

family Cervidae changed, and to observe the institution of new

genera and species and their placement (sometimes their

exclusion) inside the tribe.

Khomenko (1913) instituted the subfamily Pliocervinae on

the basis of the Late Miocene fossil deer remains (mainly

antlers) from the village of Tarakliya (Moldova). According to

the author, pliocervines are ‘‘formes de grandeur moyenne

confondant les caracteres des sous-familles Cervulinae et

Cervinae’’. Such peculiarities of their dental and antler

morphology, as well the developed ‘‘Palaeomeryx’’-fold and

the length of pedicles, are similar to the representatives of the

subfamily Cervulinae. Moreover, the presence of the rose, the

shape of the antlers and the position of the tines are shared with

the true cervines.

Khomenko described three new genera with one species for

each genus into his new subfamily: Cervavitus tarakliensis,

Cervocerus novorossiae, and Damacerus bessarabiae. In

addition to these species, the author also included Dremotherium

pentelici Gaud. from Pikermi (Gaudry, 1865), Cervus mather-

onis Gerv. from Mont Leberon (Gervais, 1852) and Cervavus

speciosus Schlos. from China (Schlosser, 1903) to Pliocervinae.

In his paper Khomenko (1913) also refers to a note by Alexeev

(1913) related to a new genus and new species, with the title

‘‘About new form of deer from Petroverovka neighbourhood’’,

but he considers that this genus belongs to the subfamily

Pliocervinae and then he observes ‘‘numerous misunderstand-

ings and vagueness’’ in the note. The main ones are as follows:

the absence of diagnosis, ‘‘new form of deer’’ (in plural for latter

word), presentation of new genus that contains the features of

Cervidae, but which also shows some peculiarities close to more

archaic deer like Dicrocerus, the attribution of the new genus to

Cervinae considering it at the basis of the Cervidae phylogeny.

Khomenko (1913) did not use the names of new genus and

species, considering them as nomen nudum, because, in his

opinion, Alexeev has several forms taking into account the

pictures and description of fossil material.

In 1915, Alexeev published the monograph ‘‘Fauna of

vertebrates of v. Novo-Elizavetovka’’ where he gave diagnosis

and a detailed description of Procervus variabilis Alex. based

on cranial and postcranial remains of this deer. The author does

not reply to Khomenko, but he writes that the deer from Novo-

Elizavetovka (hereafter we shall use recent name Novoeliza-

vetovka) is the form which links the subfamily Cervulinae with

the subfamily Cervinae. However, Alexeev did not point out

that Procervus belongs to Pliocervinae and, on the basis of

some indications in his monograph, we may conclude that he

did not accept this subfamily.

Hilzheimer (1922) instituted the new genus Pliocervus for

the deer previously described by Gervais (1852) as Cervus

matheroni from Mont Leberon.

C. Petronio et al. / Geobios 40 (2007) 113–130 115

Schlosser (1924) investigating fossil deer from Mongolia,

compared them with the remains from China described by him in

1903 as Cervavus speciosus. He replaced the genus ‘‘Cervavus’’

with Cervus (Axis) considering the latter as a more precise

generic name and indicating its position among the true deer.

Zdansky (1925) considered Damacerus bessarabiae as

synonymous with Cervocerus novorossiae even if the descrip-

tion of the latter species seems to be unsatisfactory. However, he

considers Cervavitus tarakliensis to be a valid genus and species.

Teilhard de Chardin and Trassaert (1937), following

Khomenko (1913), observed that Pliocervinae are somewhat

connected with both Cervulinae and Cervinae. These authors

indicated that the most important morphological features of the

Pliocervinae are: pedicles prolonged by the strong frontal

ridges, three-pointed antler (in the most typical forms), the

presence of lateral metacarpals along the entire length, and that

the ‘‘Palaeomeryx’’-fold is sometimes present. As regards the

fossil deer remains from China (Yushe Basin, Shanxi), Teilhard

de Chardin and Trassaert (1937) referred them to Cervocerus

novorossiae. In their diagnosis they observed that the remains

showed the morphological features of Pliocervinae and some

peculiarities, above all they mentioned the lack of the

‘‘Palaeomeryx’’-fold. Teilhard de Chardin and Trassaert

(1937) in addition to Cervocerus novorossiae also described

a new form in the Chinese fossil material: Cervavitus demissus.

Lacking palmated antlers, no remains belonging to Damacerus

bessarabiae are pointed out by Teilhard de Chardin and

Trassaert (1937) but they do not agree with Zdansky (1925) in

considering Damacerus to be an invalid genus.

Simpson (1945) in his classification of mammals included

Khomenko’s Pliocervinae into the subfamily Cervinae. Among

seven species belonging to Cervinae, Simpson considers

Cervocerus, Cervavitus, Procerus (= Procervus, authors

remark) and Pliocervus. Finally, he believes the genus

Damacerus as synonymous with Cervocerus.

Azzaroli (1947) mentioned Cervocerus novorossiae and

indicated that C. variabilis was closely related to this species.

Later, Azzaroli (1952) concluded that in Tarakliya and in

Novoelizavetovka there are two different genera: Cervocerus in

the first locality and Damacerus in the second one. The author

also excludes the Pavlodar deer (Kazakhstan) from the genus

Cervavitus. The systematic position of this form was in fact

problematic and has been investigated by numerous authors

(see later). Finally, he proposed to separate the Chinese remains

referred to this genus as a new species and probably as a new

genus.

Kurten (1952) followed the opinion of Schlosser (1903)

concerning the existence of two main different faunal provinces

in the Chinese Hipparion-fauna, characterized by the presence

of forest and the steppe faunas. Kurten also named the faunas

containing the elements of both previous ones as ‘‘mixed’’

fauna. The forest province is situated in southeastern part of

China and usually includes deer, especially Cervoceros

novorossiae (Kurten, 1952).

Flerov (1950) attributed the deer remains from Tarakliya,

described by Khomenko as three genera, only to the genus

Cervavitus. After investigating the remains of a small deer from

Gusinyy Perelet (Pavlodar, Kazakhstan), the Pavlodar deer,

Flerov (1950) attributed them to the new species Cervavitus

orlovi on the basis of their antler morphology, similar to the

Cervavitus antlers from Moldova (Khomenko, 1913) and from

southern Ukraine (Alexeev, 1915). Flerov in his paper gave

only a short diagnosis based on the features he considered

peculiar, but he did not make a full description of the species.

In 1952, Flerov employs the system of the cerviforms where

only Cervavitus and Pliocervus are given. Later he reconfirms

the presence of the genus Cervavitus (Cervocerus, Damacerus,

and Procervus were included into Cervavitus), however he

wrote about three species in this genus but he did not name them

(Flerov, 1962).

Czyzewska (1960) described a new species of deer from the

Polish locality of Weze as Cervocerus wenzensis. The author

recognized similar features between the deer from Weze and

Cervavitus s.s., above all in the teeth and the skull

characteristics: both deer in fact show a wide frontal bone,

the same position of the pedicles, large-sized frontal fossa,

presence of the upper canines, molars with low crown and

‘‘Palaeomeryx’’-fold. The author also marked the ‘‘inadapta-

tive’’ type of the antlers of the new species among the main

differences with Cervocerus novorossiae. Finally, she included

the genus Cervocerus in the subfamily Cervinae.

Viret (1961) accepted the subfamily Pliocervinae and gave

its generic composition with Cervocerus and Procervus; he

represented the diagnosis of these genera as well.

Korotkevich (1965) established the new species Pliocervus

kutchurganicus from Kuchurgan (Ukraine) studying the fossil

deer material, above all antlers, found in this locality.

Romer (1966) considered Cervocerus to be a good genus in

his classification of vertebrates with three synonyms: Cerva-

vitus, Damacerus, and Procervus.

In 1968, Czyzewska referred Cervocerus wenzensis to the

genus Procapreolus after she studied the deer remains from the

Odessa region and from Moldova. However, she recognizes

some common characteristics between P. wenzensis and

Cervavitus that could in her opinion prove their evolution from

the Cervulinae forms. According to the author, the subfamily

Cervinae consists of five tribes including the tribe Pliocervini.

She writes that pliocervines have close relationships with most

Cervinae and that their attribution to a subfamily proposed by

Simpson (1945) is correct, but taking into account their

primitiveness and some peculiarities, Czyzewska suggests the

distinction of the tribe Pliocervini nov. (= Pliocervinae

Khomenko, 1913) with the genera Cervavitus and Pliocervus

(as incertae sedis). She believes Cervocerus and Procervus as

synonyms of Cervavitus also showing that Procervus Alexeev,1915 ‘‘cannot be maintained because it is a homonym of

Procervus Hodgston in 1847’’. According to the International

Code of Zoological Nomenclature (art. 36.1), although

Czyzewska (1968) was the first to consider Pliocervini as a

new tribe, we have to maintain the authorship of Khomenko

(1913); this also excludes the authorship of Symeonidis (1974),

as often referred by numerous authors.

Czyzewska considers the genera Cervavitus, Cervocerus,

and Damacerus as different stages of the ontogenetic


development of a deer belonging to one genus and one species,

supporting the opinion of several authors. She also supports

Azzaroli (1952) proposal that the deer from China, described by

Zdansky (1925) and by Teilhard de Chardin and Trassaert

(1937) as Cervocerus novorossiae requires further studies and

perhaps belongs to a new genus and species. Czyzewska also

refers the same opinion on the deer from Pavlodar and she does

not assign it to Cervavitus either.

Korotkevich (1970) instituted the new species Cervavitus

sarmaticus from the Late Miocene (Late Sarmatian) of Krivoj

Rog (Ukraine) on the basis of the lower part of a right antler. In

her monograph this author describes material from different

localities which belong to several subfamilies of Cervidae,

including Cervinae. She places the genera Cervavitus and

Pliocervus into this last subfamily without mentioning the tribe

Pliocervini. Korotkevich also describes Cervavitus variabilis

from Novoelizavetovka and Cervocerus novorossiae from

Tarakliya. With regards to the genus Pliocervus, this author

institutes the new subspecies P. kutchurganicus vesti from

Kosyakinskij quarry (Stavropol, Russia).

Korotkevich (1970) was the first to point out the

resemblance between the antler’s shape and structure to the

Pavlodar deer and those of young reindeer observing that both

lack the grooves. She also writes that Cervavitus orlovi strongly

differs in its antlers from the typical Cervavitus and, due to such

morphology, it is excluded by some authors from this genus.

Auberkerova (1974) studied a vertebrate collection from

southeastern Kazakhstan (Yesekartkan) and described the new

species Cervavitus flerowi. This species, according to the

author, is different from all the other representatives of the

genus because of the converging roses, the short pedicles and

the lack of the ‘‘Palaeomeryx’’-fold. Moreover, the perpendi-

cular trez tine and the size of the rose indicate a closer

relationship with the Asian species than the European ones.

Symeonidis (1974) gave a short historical analysis of

Pliocervus pentelici from Pikermi. In this analysis the author

concluded that Cervavitus and Pliocervus had a different shape

of antlers and the systematic position of these two species could

be clarified after the revision not only of the antler remains, but

also of the cranial and postcranial ones. He also describes

some features of the genus Pliocervus and emphasizes its

primitiveness.

Abdrakhmanova (1973, 1974) described the new species

Pliocervus karabastuzicus from Early Pliocene of Kazakhstan

(Karabastuz). The antler is S-curved at the base, his surface is

covered by small furrows which become smoother towards the

top. The section of the antler near the basis is rounded and the

shape of such section of the beam is oval. The brow tine is high

situated, but not higher than the half of the total length of the

antler.

Qiu (1979) instituted a new species from the Huade district

of Inner Mongolia (North China), which he named Cervocerus

huadeensis. The author compares this species with Cervocerus

novorossiae and Cervavitus variabilis and concludes that

Cervocerus huadeensis, Damacerus bessarabiae, Cervocerus

novorossiae, and Dama sericus can be united into one genus or

subgenus.

Vislobokova (1980), after comparing Cervavitus orlovi with

the remains referred to Cervavitus and taking into account the

skull characteristics, instituted the new genus Pavlodaria, that,

according to the author, may be considered as the first example

of the tribe Neocervini, diffused at present time in North

America. Vislobokova (1980) after a detailed examination

confirmed that Pavlodar deer is considerably different from all

Cervinae representatives, also including Cervavitus, and she

showed its real systematic position among Cervidae.

Korotkevich (1988) observes shortly the systematic position

of Pliocervini and their history. She writes that Flerov (1950) at

first referred all deer from Tarakliya to the genus Cervavitus,

and then he (Flerov, 1952) assigned all species from eastern

Europe only to the species Cervavitus variabilis. Korotkevich

considers Cervavitus novorossiae and Cervavitus variabilis to

be two independent species. She also looks closely at the

systematic position of Cervavitus sarmaticus, instituted by her,

which she replaces to the genus Euprox. Korotkevich gives the

following diagnosis of E. sarmaticus: two-tined and large

antlers, short and lacking of a developed upper bifurcation. The

beam is straight and not flattened. The brow tine is rather poor

developed and situated relatively high from the rose.

Korotkevich remarks the separation from the beam to the

brow tine as an important feature of this species which makes

different it from E. furcatus and E. dicranocerus. Korotkevich

(1988) attributes the genera Cervavitus and Pliocervus to the

tribe Pliocervini of the subfamily Cervinae and gives

description (or additions to descriptions) of Cervavitus

variabilis, Cervavitus novorossiae, and Pliocervus kutchurga-

nicus. Korotkevich (1988) shared the opinion of Flerov (1952)

and mentioned that the three genera from Tarakliya had been

described on the basis of their antler morphology, ‘‘whose

variability is in the range of individual variability of recent

deer’’. This author pointed out that representative of Cervavitus

lived on the territory of eastern Europe mainly during the

Meotian Age (Late Miocene) and that their findings were not

confirmed in more earlier or later deposits.

Vislobokova (1990) includes into the subfamily Cervinae

three tribes: Pliocervini, Cervini and Megacerini. The first one

consists of the genera Cervavitus and Pliocervus. Cervocerus,

Damacerus, and Procervus are considered as synonyms of the

genus Cervavitus. She represents, besides generic composition,

diagnosis and distribution of Pliocervini, and also its

comparison with another two tribes referred to Cervinae.

Gentry (1994) observed several publications related to

pliocervines without marking their position in the family

Cervidae. He mentioned Procapreolus loczyi, Pliocervus

pentelici, Cervavitus, and Cervocerus as Late Miocene deer,

confirming the similarity of the two last genera with Muntiacus

and also discussing a little bit about their relationship with

Odocoileinae and Cervinae.

Dong and Hu (1994) studying Cervidae from Hounao

locality (Yushe basin, Shanxi province) gave the description of

two new subspecies of Cervavitus novorossiae: C. novorossiae

shanxius, and C. novorossiae minor. These authors placed

Cervavitus genus into subfamily Pliocervinae. Taking into

account the similarity of Chinese Cervavitus novorossiae with

Fig. 1. Palaeontological Museum of Mechnikov Odessa National University:

antlers of Cervavitus novorossiae N 2384-494 on the left and N 2384-579 on the

right.


the same species described by Zdansky (1925) and Teilhard de

Chardin and Trassaert (1937), they recognized however some

endemic characters in the remains from Hounao. Such features,

like the absence of the ‘‘Palaeomeryx’’-fold and more robust

antlers differentiate the Chinese forms from typical Cervavitus

novorossiae from Tarakliya. Dong and Hu propose that ‘‘the

group found in Shanxi’’ should be a subspecies of Cervavitus

novorossiae and ‘‘the one found at Tarakliya as Cervavitus

novorossiae novorossiae’’. About the second new subspecies C.

novorossiae minor the authors write that it has the same

morphological features as C. novorossiae shanxius, but has

considerable differences in its much smaller dimensions.

Dong (1996) refers the presence of Cervavitus cauvieri in

the Early Pliocene deposits of Languedoc and Roussillon

(France). This species, previously attributed to the genus

Cervus and testified only by few remains, could represent the

latest occurrence of the genus Cervavitus in Europe and also

corresponds to its westernmost presence.

McKenna and Bell (1997) considered Pliocervini as a good

tribe but refer to Symeonidis (1974) as its author. As we showed

above, Czyzewska (1968) was the first to suggest this rank. Inside

the Tribe Pliocervini these scientists classify Cervavitus and

Pliocervus as Czyzewska (1968), Korotkevich (1988), Vislo-

bokova (1990) did earlier. The authors of this classification also

mentioned that Procervus Alexeev, 1913 was nomen nudum.

Dong and Jie (1997) studied the morphological variation of

antlers and teeth of Cervavitus novorossiae. These authors

analyzed annual, environmental, pathological and, random

variations of this species comparing them with Procapreolus

latifrons and Platycemas infans (Teilhard de Chardin and

Trassaert, 1937). In their opinion, the last species is a

pathological variation form of Cervavitus novorossiae while

P. latifrons is a random variation of this species.

Gentry et al. (1999) dealing with the Miocene land mammals

of Europe gave as valid the genera Cervavitus and Pliocervus

among Cervoidea (without marking family, subfamily, and

tribe status).

Azanza (1995, 2000) also assigns Pliocervus to Cervinae;

she also includes to this genus three known species P.

matheroni, P. graecus and P. kutchurganicus and a new one,

instituted by her, P. turolensis.

Di Stefano and Petronio (2003) mention that they consider

the subfamily Pliocervinae at the rank of tribe, regarding

Symeonidis (1974) as its author. They also write that Flerov

(1952) instituted the species Cervavitus sibiricus on the basis of

the Early Pliocene remains collected in western Siberia and

which are ‘‘characterized by a more complicated antler

structure’’. According to these authors the remains attributed

to the species Cervavitus novorossiae from numerous localities

of China could be referred to different forms.

3. European pliocervine morphology

3.1. Tarakliya: Cervavitus novorossiae

After the historical synthesis on the systematic of

Pliocervini, it is important to deal particularly with the

morphology of the species referred to the genus Cervavitus, on

the basis of the fossil material from the eastern Europe localities

stored in the Palaeontological Museum of Mechnikov Odessa

National University.

The deer fossil remains observed by the writers are above all

cast antlers or frontal bone fragments with the basal portion of

the antler. The antlers are very important for the systematics of

Cervinae, not only because these remains are rather frequent in

the fossil assemblages, but also because of their variation that

may indicate the phylogenetic position of a cervid group.

The remains classified as Cervavitus tarakliensis (N 2384-

494 and N 2384-579) consist of two fragmentary antlers

(Fig. 1). They can be probably referred to the same young

individual. The antlers are in fact small-sized; both the pedicle

and the b span (i.e. the part of the beam from the rose to the first

tine, according to Pocock, 1933) are rather long. The rose is

poorly developed. The pedicle and the b span (the portion

between the rose and the first bifurcation of the antler) show

rather circular sections, even if a little flattened. The pedicles

were probably subparallel. Both specimens show a brow tine

curved inward and forming a 458 angle with the beam.

The complete left antler with pedicle (N 257), attributed by

Khomenko to Cervocerus novorossiae, is referable to a young

individual (Fig. 2). Both the pedicle and the b span are rather

long and they are separated by a well-developed rose; the b span

is laterally flattened; the first part of the beam tends to curve


left antler with pedicle of Cervavitus novorossiae (N 257).

Fig. 3. Palaeontological Museum of Mechnikov Odessa National Univers


slightly outwards. The brow tine forms an angle of about 558with the beam, slightly larger than that shown by the antlers

referred to Cervavitus tarakliensis.

A skull fragment (N 366) with pedicles allows to obtain a

general point of view of the frontal appendages (Fig. 3). The

pedicles are rather long with a subcircular section and they are

slightly divergent. From the proximal portion of these

appendages a little crest starts which joins them to the orbit.

The rose is not large and the b span is long. The beam is

laterally flattened in its lower part. The left antler separates in

two tines of the same length in its upper part. The right beam is

different from the previous one because it does not divide itself

and terminates with only one tine. In the general point of view

the morphology of this antler recalls the modern deer of the

genus Axis.

An incomplete right antler can be referred to an adult

individual (N 131). The pedicle is rather long, it is connected

with a fragment of the frontal bone and shows a subcircular

section. The rose is rather large. A remarkable shortening of

the beam between the rose and the brow tine (b span) can be

observed. The beam is also laterally flattened in this part. The

brow tine forms an angle with the beam which is larger than the

ity: skull fragment (N 366) with pedicles of Cervavitus novorossiae.


antlers attributed to Cervavitus tarakliensis. The beam is

broken just beyond the brow tine.

The specimens referred by Khomenko to Damacerus

bessarabiae are a complete left antler with the pedicle and a

fragmentary frontal bone belonging to an adult individual (N

2389-235) (Fig. 4). The rose is smaller if compared with the

previous remains. Moreover, a crest proceeding to the orbit

originates from the pedicle. The b span is as long as in

Cervocerus novorossiae. The brow tine is lacking in the

terminal part; it forms an angle of about 708 with the beam and

is curved in its first part. The beam beyond the brow tine has a

constant section until two-third of its length, where it becomes

strongly flattened forming an evident palm. The upper part of

the beam ends with three tines, those lateral being similar in

length and the central being the shortest one. It is probable that

the complete antler could show a ‘‘lyra’’ morphology.

An incomplete antler (N 504) has been attributed to the same

form: it shows a pedicle similar in size to the previously

described specimen and a long b span.

Finally, an almost complete left antler lacking the upper part

has been described as Damacerus bessarabiae. This antler

belongs to an adult individual; the rose is not well-preserved;

the b span length is comparable to all the other adult specimens

investigated by Khomenko. The brow tine is curved slightly

inwards. Both the beam and the brow tine have a laterally

flattened section; moreover, the beam broadens beyond the

bifurcation and becomes increasingly flattened. It can be


complete left antler with the pedicle (N 235) of Cervavitus novorossiae.

hypothesized that this antler terminates with a palm of two or

three tines.

3.2. Systematic conclusions on the deer from Tarakliya

The morphological data observed and their analysis

indicates that the deer fossil material from Tarakliya cannot

be referred to three different genera as suggested by Khomenko

(1913). The species Cervavitus tarakliensis does not seem to be

valid because it was described on the basis of two young antlers

belonging to the same individual. The remains of young

individuals do not in fact allow satisfactory analysis or

verifications of the diagnostic features and therefore it is

necessary to compare these specimens with adult ones.

The antlers belonging to Cervocerus novorossiae and

Damacerus bessarabiae by Khomenko (1913) also following

other authors (Azzaroli, 1952; Czyzewska, 1968; Korotkevich,

1988), can represent different ontogenetical stages of one

species that can be named Cervavitus novorossiae; to this

species can be attributed also the specimens N 2384-579 and N

2384-494 (young individual) described as Cervavitus tarak-

liensis. In fact, the morphological differences in the antlers of

the Pliocervini from Tarakliya fit into the intraspecific range.

We believe in the opportunity to maintain the genus Cervavitus

for nomenclature stability also taking into account the use of

this name by scientists from Russia and Ukraine (Flerov, 1962;

Korotkevich, 1970,1988; Vislobokova, 1990).

On the basis of the antler morphology observed in the

Paleontological Museum of Mechnikov Odessa National

University and from literature data, we propose the following

diagnosis for the species Cervavitus novorossiae.

It is a medium-small sized deer, a little smaller than the

living sika deer (Cervus nippon), characterized by a peculiar

morphology of the antlers. In the adult individuals the antler

usually constitutes three tines, even if the development of

supernumerary tines is rather frequent. The pedicles are long

and rather separated from each other; the rose is not very large;

the b span is long. Finally, the beam and the brow tine are only

slightly flattened (Korotkevich, 1988).

During the evolution which brings to the advanced cervids, a

progressive decrease of the pedicle (followed by the increase of

its diameter) and b span length can be observed. Taking into

account this evolutionary trend, it seems correct to consider

genus Cervavitus in the Pliocervini which represents the most

ancient tribe in the subfamily Cervinae. The beam of the

pliocervines from Tarakliya tends to curve to the outside in the

first part and to the inside in its the second part. Therefore, from

the anterior point of view, the antlers present a shape that some

authors (Teilhard de Chardin and Trassaert, 1937) describe as

‘‘lyra’’ shaped. In a general point of view this shape is similar to

those showed by the genus Axis.

Dong and Jie (1997) analyze four types of morphological

variations in the Chinese forms referred to Cervavitus

novorossiae: annual, environmental, pathological, and casual

variations. The presence of more palmated forms among the

deer from Tarakliya than in those from northern China, as

suggested by Dong and Jie (1997), could be referred to the


range of the environmental variations due to both geographical

differences and the occurrence of some nutrients. In some

deposits, where the fossil remains are not numerous, some

features belonging to the intraspecific range can be easily

confused with those belonging to the interspecific range.

The antler morphology can represent the synthesis of the

features that indicate an archaic pattern (making this species,

and generally the tribe of Pliocervini, as descendant of some

Middle Miocene species of the genus Dicrocerus) and of the

features that will develop subsequently into the other tribes of

the subfamily Cervinae.

The antler remains of Cervavitus novorossiae have been

previously considered as belonging to individuals of different

age: that allows to hypothesize the ontogenetical development

of its antlers. The young individuals, as observed by

Korotkevich (1970), recall the antler morphology of the

representatives of the subfamily Muntiacinae. It is useful to

remember that pliocervines during their ontogenetical devel-

opment show the following modifications of the antlers:

� in
crease in size;
� in
creasing in the number of tines with three or more tines in
the adult individuals;

� d
evelopment of the pedicles which are subparallel in the
young animals but they separate widely in the adult ones;

� th
e beam is in a right line with the pedicle in the young
individuals while it is curved in the adult ones;

� th
e brow tine forms an angle with the beam which increases
as the age increases until reaching an angle of about 908;
� th e beam is often palmated in the upper part;
� b
oth the pedicle and the b span, decrease in size during
lifetime (this has to be put into relation both with the

increasing of the size of the antlers and with the increasing in

the number of the tines that create static problems to the

animal);

� th


antler of Cervavitus variabilis (N 2265).

e pedicle is laterally flattened in the young individuals but it

becomes more rounded in the adult ones.

As regards the teeth morphology of Cervavitus novorossiae,

it is not possible to follow the same observations as in the antler,

because of the lack of teeth in the material from Tarakliya. We

could only remember that Khomenko (1913) mentioned that the

general features of the upper teeth of the Pliocervinae

representatives were similar to those of Dicrocerus furcatus

(= Euprox furcatus).

3.3. The pliocervines from Novoelizavetovka: Cervavitus

variabilis

The fossil material described as Procervus (= Cervavitus)

variabilis by Alexeev (1915) has been collected from the

locality of Novoelizavetovka (Odessa region, Ukraine). The

faunal assemblage where Cervavitus variabilis has been found

is referable to the Late Miocene, Meotian Age (Korotkevich,

1988; Krakhmalnaya, 1996a, 1996b). The fossil material

observed is stored in the Paleontological Museum of

Mechnikov Odessa National University. Even if there are

seven many deer skeleton remains from this locality, we will

focus the analysis only on the basis of the antlers and teeth

morphology, that is on the same elements found in Tarakliya

where pliocervines were instituted.

The antlers are three-pointed, but usually supernumerary

tines develop in the adult individuals: for example, the

specimen N 2265 (Fig. 5) shows six tines at the top of the

beam but all the adult individuals never have less than three

terminal tines. These tines are rather variable in size and they

can be both subparallel and well-separated from each other; if

separated, these tines tend to curve towards the sagittal axis of

the body. Moreover, the supernumerary tines can be present not

only at the top of the beam but also just beyond the brow tine

(N 2289). Finally, we can observe the difference in shape

between the left and the right antlers of the same individual, like

in the specimens N 2313 and N 2314.

The brow tine is situated very low on the beam, near the rose.

This tine is large-sized, it is dagger-like and shows a peculiar

shape: in fact it is perpendicular to the beam at its basis but

bends towards the top in the terminal part (in this part the tine is

subparallel to the beam). The supernumerary tine of the

specimen N 2289, described before, forms a bisecting line

between the beam and the brow tine; finally this supernumerary

tine has a subelliptical section and sometimes separates in two

branches in the terminal part (as in the specimen N 2283).

The beam is palmated and increasingly flattened laterally

with an increasing of the number of tines.

The antlers are strongly signed by the traces of the blood

vessels which form deep grooves.

The upper canines of Cervavitus variabilis are similar to

those belonging to the genus Muntiacus: they are medium-sized


and curved behind in the terminal part. The teeth show a low

hypsodonty level; the ‘‘Palaeomeryx’’-fold is present on the

lower molars. Alexeev (1915) wrote that the ‘‘Palaeomeryx’’-

fold had been well-recognizable and usually present, but

sometimes in reduced form. As regards the inner wall of the

lower last premolar crown (P4), it is formed by the metaconid,

which goes in the anterio-posterior direction and is connected

with the protoconulid, at the same time its posterior part almost

reaches the entoconid, thus forming a more complete inner wall

in comparison with the same structure of C. novorossiae. In this

case the P4 of C. variabilis, according to the scheme of

Vislobokova (1990), demonstrates the 4th stage of molarization

of premolars, when metaconid is expanded in the anterior

direction closing anterior valley (= second valley under

Vislobokova terminology) and continues to broaden in poster-

ior direction approaching the entoconid.

3.4. Discussion

The morphological analysis of antlers and teeth shows a

close relationship between the pliocervines from Novoeliza-

vetovka and those from Tarakliya. The common features

between the two species are the following:

� s
ize;
� p
edicles morphology (shape, section);
� p
resence of the crest going from the low part of the pedicle to
the orbit;

� p
resence of the ‘‘Palaeomeryx’’-fold.
These species show however some differences, above all in

the antler morphology. The remains referred to Cervavitus

variabilis has a more complicated architecture of the antlers

than the pliocervines from Tarakliya. Both deer are character-

ized by the same three-pointed antler structure which is typical

of the second stage according to Geist (1971). The most

important difference in the antlers is the number of the

supernumerary tines. As already mentioned, the specimen N

2265 from Novoelizavetovka counts six tines at the top of the

beam. Among the antlers from Tarakliya only the specimen N

2389-23 can be compared with those from Novoelizavetovka

for the number of tines and the palm at the top of the beam. The

pedicles are longer in the Tarakliya pliocervines than in those

from the Novoelizavetovka even if they are similar in the shape

and in their section. The brow tine of C. variabilis is more close

to the rose in comparison with C. novorossiae and its

morphology is also different: in fact it is even larger and

demonstrates the peculiar shape already described. The

presence of deep grooves in the antlers of C. variabilis shows

how the development of the antler could be hard for the living

animal.

The presence of antler remains of C. variabilis referable to

individuals of different age allows the analysis of their

ontogenetic development.

The individuals of one year show one-pointed antlers with

subcircular section. The rose is not evident even if an increase at

the basis of the antler can be observed. The pedicle is long with

a circular section. The general shape of the pedicles of adult

individuals can also be observed in those young: in fact, both

the pedicle and the beam are externally curved but the beam

bends inwards at about one half of its length.

The individuals of two years have two-pointed antlers. The

anterior tine is shorter than the posterior one, these tines form

an angle of about 458. The pedicle is stouter and shorter than the

individuals of one year show and the rose is well-developed.

The antlers of this ontogenetic stage resemble those of the

representatives of Muntiacinae (Korotkevich, 1970).

The individuals of three years are characterized by three

pointed antlers: the beam in fact divides in two tines in its

terminal part. In the antlers belonging to the adult individuals

the following trend of development can be observed:

� in
crease in size;
� in
creasing of the number of the points with supernumerary
tines;

� th
e beam becomes increasingly flattened laterally;
� th
e angle between the brow tine and the beam increases
reaching about 908;
� b oth the pedicle and the b span become shorter;
� th
e pedicle has a subcircular section in the young individuals
but it becomes elliptical in adult deer.

The teeth morphology of C. variabilis shows features similar

to those of C. novorossiae. The male individuals of both species

show the upper canine. The morphology of these teeth is similar

to the living Muntiacinae.

The presence of the ‘‘Palaeomeryx’’-fold, which culminates

its evolution in the Late Miocene and Early Pliocene deer, is a

strong diagnostic feature even if it is poorly developed. The

most ancient species showing this feature is Eumeryx culminis

(Late Oligocene) (Vislobokova, 1990). The Early Miocene

genera Amphitragulus and Dremotherium show a more

developed ‘‘Palaeomeryx’’-fold than in the Late Oligocene

forms. The ‘‘Palaeomeryx’’-fold decreases its importance with

the development of secondary structures (styles): in fact,

generally it cannot be observed in forms far into the Middle

Pliocene.

The morphological analysis of the remains of Cervavitus

variabilis lead us to recognize this form at a specific level even

if it shows some convergences with Cervavitus from Tarakliya.

Moreover, as a result of the previously described analysis, the

differences between the pliocervines from Tarakliya and those

from Novoelizavetovka lead us to believe that Cervavitus

variabilis could be a more highly-derived descendent of

Cervavitus novorossiae. The first species in fact shows more

complex antlers with many supernumerary tines, a palmated

beam and the reduction of the pedicle and of the b span.

Another hypothesis could be that the two pliocervines lived in

the same period but each species was adapted to different

ecological niches.

In the previous analysis we do not take into account the

remains of pliocervines from other European localities referred

to other species, above all for the uncertainties on these

remains. The only exception is the Early Pliocene Cervavitus

Fig. 6. Paleontological Museum of Uppsala (Sweden): Pliocervine antler of the

first morphotype (see text).


cauvieri from France. As already mentioned in the historical

chapter, Dong (1996) attributed the remains previously referred

to the species Cervus cauvieri to the genus Cervavitus. This

attribution is based above all on the antler and teeth features of

the French remains that are extremely similar to the

representatives of the genus Cervavitus from eastern Europe

and China. More particularly, the antlers are three-pointed and

show a flattened beam above the terminal tines; the teeth are

brachiodont and characterized by an evident ‘‘Palaeomeryx’’-

fold. The remains are referred to the Ruscinian Mammal Age

(MN 15) corresponding to the Early Pliocene; therefore

C. cauvieri corresponds to the westernmost occurrence of the

genus Cervavitus worldwide and it is also the youngest

occurrence of the same genus in Europe.

As already mentioned in the historical chapter, Auberkerova

(1974) described the new species Cervavitus flerowi from

Yesekartkan (southeastern Kazakhstan). This species shows

antlers rather elongated and slightly bent as a general structure

which makes them more similar to the Chinese remains than the

European ones. The pedicles seem rather short and with

subcircular section; the rose is well-developed; the b span is

rather long and the angle between the brow tine and the beam is

very acute. Auberkerova (1974) pointed out also the lacking of

the ‘‘Palaeomeryx’’-fold which confirms the more closeness of

Cervavitus flerowi with the Chinese Pliocervini. This deer,

according to Auberkerova (1974), occurs in a faunal

assemblage referable to ‘‘the middle of the Late Pliocene’’.

4. Chinese pliocervines morphology

The deer remains from Tarakliya have been compared with

part of the fossil material from Shanxi and Honan provinces

(China) described by Zdansky (1925) and Teilhard de Chardin

and Trassaert (1937). Part of the Chinese material has been

observed in the Paleontological Museum of Uppsala (Sweden),

where a skull with right antler (lacking of tines which are cut at

their base) (loc. 73: Prov. Shansi, Wu-Hsiang-Hsien), four right

antlers (loc. 13,15: Prov. Honan, Hsin-An-Hsien; loc. 70: Prov.

Shansi, Wu-Hsiang-Hsien), one right metacarpus and one right

metatarsus (loc. 73: Wu-Hsiang-Hsien) are stored: all these

remains are referred to Cervocerus novorossiae.

The fossil material observed is just a very small part of the

numerous deer remains from China kept in museums all over

the world and therefore a particular description of the

specimens observed could be poorly significant. However,

direct observations and the analyses by Zdansky (1925) and

Teilhard de Chardin and Trassaert (1937) allow us to conclude

that the Chinese pliocervines show a shorter, stouter and

straighter beam than species of Pliocervini from eastern

Europe; moreover, the first forms generally has shorter pedicles

and large and evident rose than the second ones.

Chinese representatives have usually three tines in the

antlers, even if palmated forms can be found, when there is a

supernumerary tine inside the terminal fork. The unpalmated

antlers show large-sized terminal tines, which separate with an

angle of about 458, while the brow tine forms an angle of about

45–508 with the beam. This last feature makes the Chinese

forms different from the European ones because the last show a

more open angle that reaches 908. The antlers observed are

flattened near the terminal fork; they are very short and robust,

with straight or slightly curved, but very robust, terminal tines.

Notwithstanding the rather small number of antler remains

directly observed, we can recognize two different morpholo-

gical types in the Chinese pliocervines: the more frequent antler

morphology shows small-sized antlers with three small

flattened tines which usually form a palm (Fig. 6); the second

morphology is characterized by a small-sized beam with large-

sized and well-separated tines (both the brow and terminal

tines) (Fig. 7). This general observation does not allow a

hypothesis to be made on the taxonomy and/or on the

relationship between the two morphotypes for the scant and the

doubtful origin of the remains. We must only confirm that these

two morphotypes seem well-defined. The observation of

different antler morphologies was already noted by Teilhard

de Chardin and Trassaert (1937) in the fossil material from the

same locality (Lingtao). The authors in fact distinguished three

groups of antlers on the basis of their general morphologies but,

if we exclude the last group (‘‘Capreolus-like’’ antlers), the

Fig. 7. Paleontological Museum of Uppsala (Sweden): Pliocervine antler of the

second morphotype (see text).


other groups can be separated by their length and by their

general shape: the elongated antlers with a slightly curved beam

and the short antler with a straight beam. The first

morphological features (Di Stefano and Petronio, 2003) could

be linked with the representatives of the genus Axis and the

second ones with the representatives of the genus Rusa. It must

be also remembered that, according to Dong and Hu (1994) two

subspecies of Cervavitus occur in the Early Pliocene deposits of

China, distinguished above all by their size, (Cervavitus

novorossiae shanxius and Cervavitus novorossiae minor).

The cranial remains observed by us are fragmented and show

a rather elongated profile, even if it is not possible to estimate

with certainty the real morphology of the skull. The

neurocranium is rather short and flat with well-separated

pedicles; we cannot observe the crest between these

appendages and the frontal bones; the occipital profile does

not seem very wide. The teeth show rather archaic morpho-

logical features. The upper premolars are wide and, at least P2

and P3, are bilobated; the presence of posterior cingula can also

be observed; the premolar row is rather long if compared with

the molar row; the upper molars lack almost completely

interlobar styles but show evident cingula. We did not see lower

teeth but, on the basis of the figures published by different

authors (Teilhard de Chardin and Trassaert, 1937; Dong and

Hu, 1994), we can confirm that the ‘‘Palaeomeryx’’-fold is not

recognizable.

The morphology of the metapodials seems similar to Axis

nestii and A. eurygonos from the Villafranchian faunas of Italy:

however, the Chinese bones are shorter and slimmer than the

Italian ones and show a more gradual connection between

diaphysis and distal epiphysis. The metapodial remains

observed by us in Uppsala are also longer than the living

Axis even if they maintain the same proportions. We cannot

confirm the holometacarpalian conditions of Chinese Cerva-

vitus as suggested and figured by Zdansky (1925), but the

specimen observed testified the presence of lateral metacarpals

in the proximal end (i.e. plesiometacarpal condition).

As already mentioned in the historical chapter, some species

have been instituted on the basis of the remains yielded in the

various Chinese localities.

Teilhard de Chardin and Trassaert (1937) instituted the

species Cervavitus demissus on the basis of the antler remains

collected from some localities in the Yushe Basin (Shanxi).

These antlers are small-sized and characterized by long

pedicles but not very long b span; the brow tine is well-

developed and forms an angle of about 45–508 with the beam;

the beam is straight and thrown backward at the top as in

Muntiacus and does not show any terminal tine. C. demissus

also occurred in younger deposits (Early Pliocene) than the type

locality (latest Miocene) (Tang and Zong, 1987). As already

suggested by Teilhard de Chardin and Trassaert (1937), the

morphological features of the antlers referred to C. demissus are

rather archaic, with the exclusion of the short b span. The

possible relationships of this species with the other Chinese

representatives of the genus Cervavitus are not clear on the

basis of our knowledge. We can only hypothesize that the

morphological features of C. demissus could correspond to the

most archaic stage of the evolution of Pliocervini preserved in

conservative environmental conditions; in fact, the faunal

associations where the remains of this cervid has been collected

testify tropical or subtropical climates with warm and dense

forest areas interrupted by more opened areas (Tang and Zong,

1987).

The same considerations on a species occurring in a

conservative environment could be done for Cervocerus (=

Cervavitus) ultimus. This species occurs in a rich faunal

assemblage described by Granger (1932) and Colbert (1940)

and testifies the presence of remnant Cenozoic species together

with more evolved forms (Pan and Zong, 1993). Qiu (1979)

instituted the new species Cervocerus huadeensis from the

Huade district of Inner Mongolia (North China). The author

pointed out this species in association with remains referred to

‘‘Cervocerus’’ novorossiae. The cervid from Huade referred to

C. huadeensis shows the following features which make it

different from Cervavitus novorossiae: large four-pointed


antlers with ‘‘dagger-like’’ brow tine and flattened beam

showing a slightly lyre shape. The previous description and

the figures published by Qiu (1979) lead to observe that the

antlers of C. huadeensis are more similar to those referred to the

most archaic representatives of the genus Cervus. More

particularly the remains referred to C. huadeensis could be

attributed to the species Cervus magnus which is a rather

common faunal element in many Chinese deposits of the Early

Pliocene (Zdansky, 1925; Di Stefano and Petronio, 2003).

As already suggested by Di Stefano and Petronio (2003), a

critical reappraisal of all the Chinese fossil material is

necessary to establish more certain philogenetical relationships

between the European Cervavitus novorossiae and the Chinese

species or subspecies, which also seem to be more archaic in the

morphological point of view. This study must be also

accompanied with the revision of the stratigraphical contexts

of the Chinese localities yielding Pliocervine remains.

On the basis of present-day knowledge, there are no

pliocervines in Asia (and in Europe) in the Middle Miocene,

while the genus Cervavitus has been pointed out with numerous

European Cervavitus Chinese Cervavitus

Three or more pointed antler Generally three-pointed antler

Curved development of the beam Straight development of the beam

The upper part of the beam is always laterally flattened and

often palmated in the adult individuals

The upper part of the beam is not very flat and the palmation is unusual

The terminal tines are small-sized and curved inwardly The terminal tines are large and form between them an angle of 458;therefore the antler shows an Y shape

The angle between the brow tine and the beam is about 908 The angle between the brow tine and the beam is about 508The brow tine shows a subelliptical section The brow tine shows a subcircular section

B span generally long also in the adult individuals B span long in the young individuals but shorter in the adult ones

Not very robust rose Well-developed and robust rose

Long pedicles with a subelliptical section Usually not very long pedicles with a subcircular section

Strong development of crests between the pedicles and the orbits Light development of crests between the pedicles and the orbits

Presence of a well-developed ‘‘Palaeomeryx’’-fold in the lower teeth

remains in many Chinese deposits referred to the Late Miocene.

From the biochronological point of view, the first occurrence of

the genus Cervavitus is pointed out in China in the Bahean

Mammal Age (Sigou fauna, Linxia Basin, Tibet), which

correspond to the Vallesian Mammal Age in Europe (about 9–

12.5 m.y.) (Fang et al., 2003), but it became more frequent in

the deposits referred to the Baodean Mammal Age, which

corresponds to the European Turolian Mammal Age (Gu et al.,

1992; Qiu and Qiu, 1995). The genus Cervavitus is a rather

common element in the Chinese faunal assemblages from the

latest Miocene to the most part of the Early Pliocene, above all

in the northern-central Chinese provinces (Shanxi, Honan)

where both historical remains and those yielded from more

recent collection have been collected (Zhang et al., 1978;

Zhaoyuan et al., 1985; Dong and Hu, 1994 inter alios).

Only considering the specific attribution, all the Chinese

Pliocervini remains of this age are referred to the following

species: Cervavitus novorossiae and Cervavitus demissus. The

Pliocervine findings become scarce in China from the Middle

Pliocene, because they are replaced by the representatives of

the tribe Cervini (above all by the genera Axis, Rusa and

Cervus). However, in the Late Pliocene and in the Early

Pleistocene the species ‘‘Cervocerus’’ (= Cervavitus) ultimus

still occurs in China which is probably the last representative of

the tribe Pliocervini (Pan and Zong, 1993).

5. Discussion and comparison

Both the Chinese and European forms belong to the same

evolutionary stage according to the model of the ‘‘Old World

deer’’ (Geist, 1971). This model refers to the Late Miocene –

Early Pliocene ‘‘three-pointed’’ forms of the second stage.

Pliocervines from China and eastern Europe referred to the

genera Cervavitus or Cervocerus show some similar and

diagnostic features, such as the number of tines, the general

antler morphology and the presence of the small-sized upper

canines.

However, the morphological analyses of the adult indivi-

duals allow us to draw the following general differences

between the remains referred to the genus Cervavitus (or

Cervocerus) from Europe and those from China.

On the basis of the compared data of the European and

Chinese populations, we suggest that the division of the species

Cervavitus novorossiae into three subspecies (Cervavitus n.

novorossiae, Cervavitus n. shanxius and Cervavitus n. minor)

proposed by Dong and Hu (1994) could be reconsidered.

The morphological features of the two groups show some

analogies but also strong differences which may indicate their

belonging to the same genus but to different species. In fact, the

characteristics observed in the antler and teeth remains are

hardly linked at a subspecific level between two forms rather

coeval but strongly separated from a palaeobiogeographical

point of view, as shown further in the text. Therefore it seems

more reliable to separate the European remains referred to

Cervavitus novorossiae from the Chinese ones referred to

the same species. From the taxonomical point of view the

European remains are referred, as already mentioned, to

the species C. novorossiae, while the Chinese remains may

be referred to the species C. shanxius, raising to the specific rank

the first subspecies described by Dong and Hu (1994). As

regards the last mentioned authors, on the basis of their

observations and those made by us, the presence of two


subspecies, C. shanxius shanxius and C. shanxius minor may be

confirmed.

According to the Chinese authors the last two subspecies can

be considered as typical Chinese forms with endemical

characteristics. Further studies on more numerous deer material

from China could extend the variability range of the observed

differences and perhaps could allow the relationships among all

the Chinese Pliocervine species and the European ones to be

considered.

As already mentioned the separation at a specific level

between the European and the Chinese forms is also sustained

by the palaeobiogeographical data of the areas where these

cervids lived and, more generally, of Eurasia.

The first thing we have to remember is the presence of a

large basin during the Cenozoic, the Paratethys Basin, in the

area which at the present time spans from the Black Sea to the

sea of Azov, which strongly limited the faunal exchanges from

different part of Eurasia (Fig. 8).

To correlate the stratigraphical scale of the Mediterranean

Basin with the stages of the Eastern Paratethys it is useful to

note that the Mediterranean stratigraphy is connected to

continued marine deposits and it can be referred to the standard

scale, while the Paratethys Basins were isolated for several

periods and therefore they are connected to different regional

stages set on the Central and Eastern Paratethys.

The Paratethysian bioprovince was an open sea between

Eocene and Oligocene. During the Early Oligocene the

isolation increased along all the areas involved (Protopar-

atethys); this phase was followed during the Late Oligocene

and the Early Miocene by the increase of connections with the

open sea (Eoparatethys). During the late Early Miocene the

central and the eastern zones developed into two different

directions (Mesoparatethys), showing subsequently strong

endemism in the late Middle Miocene (Neoparatethys). A

modified scheme from Rogl (1998) is proposed to allow the

correlations between the stages of the Eastern and Central

Paratethys and the same stages of the Mediterranean area

(Fig. 9).

Fig. 8. Reconstruction of sea boundaries of Europe during the Middle Miocene

The presence and the evolution of the Paratethys Basin

during the Oligo-Miocene are determined by the tectonic

movements that, above all from the Miocene with the growth of

the Alpine-Himalayan range, influenced the environment and

the possibility of migratory flows of the continental faunas

(Heintz et al., 1990).

Finally, another element must be added to the palaeobio-

geographical sketch of the late Cenozoic that is the progressive

aridity of wide zones of Eurasia and the following development

of open environments (Fortelius et al., 2002). The raising of

more arid environments strongly influenced the occurrence and

the evolution of the mammals, particularly of the cervids, which

in fact show the tendency to increased regionalism during the

Late Miocene and the Early Pliocene and the possibility to

spread only through defined pathways or ‘‘corridors’’

characterized by rather conservative environments (Heintz

et al., 1990; Di Stefano and Petronio, 2003).

The raising of more opened environments with different

food available also causes the modification of many

characteristics, above all in the antlers and in the teeth, which

will always be more peculiar in the evolution of cervids.

The presence of the upper canines developed as Muntiacus is

an archaic characteristic (Teilhard de Chardin and Trassaert,

1937). Owen (1846) was the first to observe the inversal

relationship between the size of canines in the Cervoidea and

degree of complexity in the antler. The species showing large

canines do not have antlers and those with highly developed

antlers have small or no canines. The large-sized canines are

weapons connected to the territorial defence of resources

(Geist, 1971). The canines and chewing teeth (premolars and

molars) can be put also into relationship. Sabre upper canines

have been found with low crowned molars; more evolved

premolars and molars (selenodont, subhypsodont, hypsodont)

are associated with small canines or often with no canines. The

evolution of the chewing apparate in the late Miocene Cervids

could also be linked with the passage from a ‘‘browsing’’ habit

to a much more ‘‘grazing’’ habit. In the Cervoidea the antlers

substituted the canines as ostentatious characteristic and the

(grey = marine basins, lines = marine platforms, points = continental areas).

Fig. 9. . Correlation table between European and Chinese Mammal Ages and the stages of the Eastern Paratethys.


lack of canines could determine a change in the combat rituals

among the ‘‘grazers’’ in opened or grassland environments.

The presence of supernumerary tines causes an evident

intraspecific variability. These tines could be related to a

different food availability and with hormonal variations (Dong

and Hu, 1994). It could be also hypothesized that the

supernumerary tines helped the living animal for the static

equilibrium, above all for those individuals with large antlers;

the development of supernumerary tines therefore is strongly

variable for the mentioned reasons because every individual

could be different due to availability of food and differing

hormonal levels (Geist, 1971; Bubenik and Bubenik, 1990; Di

Stefano and Petronio, 1997).

Changes in the teeth are associated with the cranial ones. In

fact, the evolution of the size and shape of the teeth, caused

changes in the proportions of the facial region. The structure of

brain region changed in association with the transformation of

the chewing muscle and the development of different move-

ments of the mandible. Vislobokova (1985) observed the

reinforcement of the muscles controlling the flexing-extending

movements in the atlas-occipital articulation. This reinforce-

ment can be also observed in all the Miocene and Early

Pliocene forms with large antlers. The external crest of the

occipital is strongly developed above all in the funicular part of

the nape ligament. This part extended for one third of the scale

of the occipital bone in the genus Cervavitus. In the

evolutionary pattern of Cervinae, this crest becomes always

more reduced: this reduction corresponds to the increased of

importance of the articulation which plays a more important

role; the condyles size increases and they evolve in a different

shape and finally the role of the flexing muscles decreases.

6. Conclusions

The direct observation of the fossil material allowed the

morphological analysis and the comparisons which gave the

opportunity to better define the problems on the Late Miocene

and Early Pliocene Pliocervini, with particular reference to the

genus Cervavitus.

On the basis of the previous analyses and discussions we can

trace the following conclusions:

� th
e deer material from Tarakliya belongs only to one genus
and one species, represented by specimens of different


ontogenetic stages; we accept the name Cervavitus novor-

ossiae for this species;

� th
e species Cervavitus variabilis is confirmed as a valid
species for its homogeneous characteristics; moreover, the

large number of remains observed from the Novoelizave-

tovka (A. Alexeev material) allow analysis of the ontogenetic

development of the antler of this species; we also

hypothesize that C. novorossiae could be the ancestor of

C. variabilis: this last hypothesis can be reliable taking into

account the antler morphology because the characteristics

of C. variabilis seem derived from those of C. novorossiae.

On the basis of the biochronological data this hypothesis is

not confirmed because Tarakliya locality seems younger than

Novoelizavetovka (Korotkevich, 1988; Krakhmalnaya,

1996a, 1996b);

� th
e Chinese pliocervines may be referred to the same genus
Cervavitus as the European ones but they can be separated at

the specific level. Therefore the Chinese remains attributed to

C. novorossiae are referred to the species Cervavitus shanxius

which can be recognized also in two subspecies: Cervavitus

shanxius shanxius and Cervavitus shanxius minor; these

subspecies belong to the same evolutionary stage (sensu

Geist, 1971);

� w
e can hypothesize a first migratory wave from China to
Europe during the late Middle Miocene (this migration could

determinate the evolution of the European Late Miocene

Cervavitus novorossiae and Cervavitus variabilis) and a

second migratory wave taking place in the Early Pliocene

again from China that could determinate the presence of the

Middle and Late Pliocene European true deer (genera Axis

and Rusa).

The genus Cervavitus widely occurs in southeastern Europe,

above all in southern Ukraine and Moldova. It is also pointed

out in Hungary (Hatvan) (Gaal, 1943) but the western boundary

of the range of Cervavitus could be signed by the occurrence of

C. cauvieri in France. The genus Cervavitus never occurred

beyond the 488 north latitude.

In Asia the genus Cervavitus is very common in localities of

many Chinese provinces, above all in the Central ones (Qiu and

Qiu, 1995).

Following the analyses and the conclusions of the previous

chapters, according to the writers, the species referable to the

genus Cervavitus are those in the Fig. 10.

This genus probably is associated with an ecotonal

environment (between woody and more opened areas with

Fig. 10. Table of the species referable to the genus Cervavitus.

temperate climate) on the basis of the taphonomic and faunal

data from the deposits where Cervavitus has been collected

(Alexeev, 1915; Korotkevich, 1970; Fang et al., 2003 inter

alios). The number of remains collected in some deposits may

indicate that this deer lived in groups with many individuals

(Flerov, 1952).

One of the goals proposed in the introduction of this work

was the attempt to relate the genera Axis and Rusa, which,

according to Di Stefano and Petronio (2003), occur in the

Middle and Late Pliocene faunas of Europe, with the Late

Miocene and Early Pliocene pliocervines.

On the basis of the antler morphology we can hypothesize

that the origin of the two genera Axis and Rusa, which

characterize the mammal associations from the Middle

Pliocene to the Middle Pleistocene, could be recognized into

the Chinese pliocervines. In fact, the European pliocervines

represent both in the species C. novorossiae and, above all, in

the species C. variabilis two specialized forms probably

evolved from a deer close to Cervavitus shanxius. This last

species, which occurred in several provinces of China,

probably was the ancestor for two forms of deer specialized

in different niches also considering the changing climate

and the environmental conditions during the Pliocene. In

the late Early Pliocene the species Rusa elegans is in fact

pointed out in China; this species could migrate from Asia to

Europe during the Middle Pliocene with more conservative

populations and could be considered linked to the genus Rusa

from the Late Pliocene and Early Pleistocene deposits of

continental Europe (Di Stefano and Petronio, 2003). Both the

Asian and European forms of this genus seem to be adapted to

temperate climates, they are medium–large-sized and show a

very similar antler morphology with robust, short and straight

beams.

At the same time, during the late Early Pliocene, the species

Axis shansius occurs in China; it seems to be more adapted to

the warm climates of the Mediterranean Europe and it has a

smaller size than Rusa elegans and shows longer, slimmer, and

more bent antlers than this last species.

As regards the evolution and the chronology of the

mentioned event, we must take into account the more recent

genetical and zoological conclusions. According to the

‘‘molecular clock’’ determined form the mithocondrial DNA

and according to the multivariate statistical analysis on skulls,

during the Late Miocene an important evolutionary split which

separates Muntiacini from Cervini occurs (Randi et al., 1998;

Ludt et al., 2004; Meijaard and Groves, 2004). This event could

correspond to the origin of Pliocervini. Subsequently, during

the latest Miocene–Early Pliocene, a further separation is

evident with two well-defined groups: the first group which

comprises the genera Axis and Rucervus and a second group

which comprises the other representatives of the tribe Cervini,

like the genera Cervus s.s., Rusa and Dama (Meijaard and

Groves, 2004; Pitra et al., 2004). These conclusions generally

confirm the paleontological data which, as already mentioned,

indicate the first representatives of the genera Axis and Rusa to

the Early Pliocene of China (Di Stefano and Petronio, 2003)

(Fig. 11).

Fig. 11. Philogenetical sketch of the genus Cervavitus in Asia and Europe.



Acknowledgements

We wish to thank above all the people who allowed us the

observations and the study of the fossil remains: S. Steunes of

the Paleontological Museum of Uppsala, V.A. Topachevsky of

the Department of Vertebrate Palaeozoology, and Palaeonto-

logical Museum of the National Museum (Natural History of

NAS of Ukraine) and N. Podoplelova of the Palaeontological

Museum of Mechnikov Odessa National University. A special

thank to T. Kotsakis (Third University of Rome) and J. Pignatti

(‘‘La Sapienza’’ University of Rome) for the useful sugges-

tions. Finally, we wish to thank J. Gray for the revision of the

English text and A. De Marco for drawing some figures. This

paper has been realized with the contribution of Istituto di

Geologia Ambientale e Geo-Ingegneria IGAG grants.

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Remarks on some Eurasian pliocervines: Characteristics, evolution, and relationships with the tribe...

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Transcript of Remarks on some Eurasian pliocervines: Characteristics, evolution, and relationships with the tribe...