Fossil Old World monkeys (Primates, Cercopithecidae) from the Pliocene of Dorkovo, Bulgaria

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Abstract
Two colobine species occur at the early Pliocene (MN 14) locality of Dorkovo: Dolichopithecus ruscinensis Depéret, 1889 and Mesopithecus mons-pessulanus (Gervais, 1849). Such an association is known from at least three other Ruscinian or earliest Villafranchian sites, and at two other localities a cercopithecine taxon joins the two colobines. The large colobine from the Kuchurgan localities of Ukraine has previously been considered either a dis-tinct genus or a separate species of Dolichopithecus, but it is here syno-mymized with D . ruscinensis; an isolated Kuchurgan molar previously published as a macaque is here tentatively re-identified as a colobine, perhaps M . monspessulanus. No temporal patterning has been observed either for asso-ciations among these Pliocene genera or of size change within the two Dorkovo taxa.
159
GEODIVERSITAS • 2005 • 27 (1) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com
Fossil Old World monkeys
(Primates, Cercopithecidae)
from the Pliocene of Dorkovo, Bulgaria
Eric DELSON
Department of Anthropology, Lehman College and the Graduate School,
City University of New York, NYCEP (The New York Consortium in Evolutionary Primatology)
and
Department of Vertebrate Paleontology, American Museum of Natural History,
Central Park West at 77th street, New York, NY 10024 (USA)
delson@amnh.org
Herbert THOMAS
UMR-CNRS 5143 Paléodiversité, département Histoire de la Terre,
Muséum national d’Histoire naturelle et Collège de France, case postale 38,
8 rue Buffon, F-75231 Paris cedex 05 (France)
hthomas@mnhn.fr
Nikolaï SPASSOV
National Museum of Natural History, Bulgarian Academy of Sciences,
1 Tzar Osvoboditel, 1000 Sofia (Bulgaria)
nspassov@nmnh.bas.bg
Delson E., Thomas H. & Spassov N. 2005. Fossil Old World monkeys (Primates,
Cercopithecidae) from the Pliocene of Dorkovo, Bulgaria. Geodiversitas 27 (1) : 159-166.
ABSTRACT
Two colobine species occur at the early Pliocene (MN 14) locality of
Dorkovo: Dolichopithecus ruscinensis Depéret, 1889 and Mesopithecus mons-
pessulanus (Gervais, 1849). Such an association is known from at least three
other Ruscinian or earliest Villafranchian sites, and at two other localities a
cercopithecine taxon joins the two colobines. The large colobine from the
Kuchurgan localities of Ukraine has previously been considered either a dis-
tinct genus or a separate species of Dolichopithecus, but it is here syno-
mymized with D . ruscinensis; an isolated Kuchurgan molar previously
published as a macaque is here tentatively re-identified as a colobine, perhaps
M . monspessulanus. No temporal patterning has been observed either for asso-
ciations among these Pliocene genera or of size change within the two
Dorkovo taxa.
KEY WORDS
Mammalia,
Primates,
Colobinae,
Dolichopithecus ruscinensis,
Mesopithecus monspessulanus,
Pliocene,
Ruscinian,
Bulgaria.
INTRODUCTION
Discovered in 1982 during geological prospect-
ing, the Pliocene locality of Dorkovo (S-W
Bulgaria) is situated in the western Rhodope
Mountains in the Neogene deposits of the
Chepino depression, about 15 km north of the
town of Rakitovo and about 120 km E-SE of
Sofia. The site of Dorkovo, 500 m WNW of the
village of the same name, is exceptionally rich in
fossil mammals, but 90% of the remains belong to
a single proboscidean species, Anancus arvernensis
(Croizet & Jobert, 1828). The fossil level is an im-
mense concentration of skeletons in a layer more
than 1 m thick covering an area of several hundred
square meters. Only about 10% of the fossilifer-
ous zone was excavated by a Franco-Bulgarian
team in a systematic way using a 1-m grid, during
three field seasons (1985-1987). Extraction of the
bones was difficult because they were tightly inter-
locked and due to a high concentration of radon
gas. Preliminary results of the first field season
have been published (Thomas et al. 1986).
The vertebrate fauna comprises some 30 species,
but until now, only the birds (Boev 1998),
Anancus (Metz-Muller 1995, 2000) and hippari-
ons (Alberdi & Alcala 1999) have been described.
In addition to the cercopithecid monkeys dis-
cussed in the present paper, the large mammals
noted or described from the locality are as fol-
lows: Ursus ex gr. ruscinensis Depéret, 1890;
Mustelidae Fischer de Waldheim, 1817 indet.;
Hyaenidae Gray, 1821 indet. (size of Chasma-
porthetes Hay, 1921); Anancus arvernensis
(Croizet & Jobert, 1828); Mammut borsoni
(Hays, 1834); Cervidae Goldfuss, 1820 indet.;
Suidae Gray, 1821 indet.; Bovidae Gray, 1821
indet.; Stephanorhinus megarhinus (de Christol,
1834); and Hipparion crassum Depéret, 1890
(Thomas et al. 1986; Metz-Muller 1995, 2000;
Alberdi & Alcala 1999; Spassov 2003).
This faunal association is typical of the
Ruscinian, especially of the MN 14 unit. The
abundance of Prosomys insuliferus (Kowalski,
1958; Arvicolidae Gray, 1821) confirms this age,
which is further supported by magnetostrati-
graphic data indicating reverse polarity correlated
with the Gilbert chron (Thomas et al. 1986).
Characteristic features of the Dorkovo fauna
include: the relative scarcity of hipparions and
Mammut borsoni; the dominance of cervids
among the artiodactyls; the extreme abundance
of Anancus arvernensis representing at least
100 individuals; and the presence of the
Dolichopithecus-Mesopithecus association. This
association indicates a prevalence of open wood-
land habitats in the Dorkovo region c. 4.5 Ma
(Thomas et al. 1986; Spassov in press).
Delson E. et al.
160
GEODIVERSITAS • 2005 • 27 (1)
RÉSUMÉ
Les cynomorphes fossiles (Primates, Cercopithecidae) du Pliocène de Dorkovo,
Bulgarie.
Deux colobinés sont présents dans le gisement pliocène (MN 14) de
Dorkovo : Dolichopithecus ruscinensis Depéret, 1889 et Mesopithecus mons-
pessulanus (Gervais, 1849). La coexistence de ces deux primates caractérise au
moins trois autres gisements d’âge ruscinien ou villafranchien basal. Dans
deux autres gisements, ces colobinés sont associés à un cercopithéciné. Le
grand colobiné des localités de Kuchurgan en Ukraine considéré auparavant
comme un genre distinct ou comme une espèce différente de Dolichopithecus
est interprété ici comme un synonyme de D. ruscinensis. Une molaire isolée de
Kuchurgan identifiée précédemment à celle d’un macaque est attribuée à un
probable colobiné, peut-être M. monspessulanus. Aucune répartition chrono-
logique particulière n’a pu être observée quant à l’association de ces genres
pliocènes ni quant aux changements de taille chez les deux taxons de
Dorkovo.
MOTS CLÉS
Mammalia,
Primates,
Colobinae,
Dolichopithecus ruscinensis,
Mesopithecus monspessulanus,
Pliocène,
Ruscinien,
Bulgarie.
A BBREVIATIONS
DKVDorkovo collection (in NMNH);
MNHN Muséum national d’Histoire naturelle,
Paris;
NMNH National Museum of Natural History,
Sofia.
MATERIALS AND METHODS
The specimens which form the focus of this paper
comprise 10 primate teeth from Dorkovo: NMNH
DKV 78, 79, 82 and 479-485. Two smaller teeth
(DKV 480 and 481) are identified as Mesopithecus
monspessulanus, while the eight larger specimens
are identified as Dolichopithecus ruscinensis.
Comparative metrical data are provided for the
equivalent teeth only from the major samples
known for each species (see Delson 1973 for a dis-
cussion of most material). The large sample of
Dolichopithecus ruscinensis from Perpignan is
housed mainly in Paris and Perpignan. The sam-
ples of Mesopithecus monspessulanus from Mont-
pellier and Fornace RDB (Villafranca d’Asti) are
housed in Paris, Lyon, Montpellier and Basel.
Specimens were measured by the senior author
according to the definitions in Delson (1973),
using Helios dial or Fowler digital calipers fitted
with needle points; measurements were recorded
in tenths of mm.
SYSTEMATICS
Order PRIMATES Linnaeus, 1758
Hyporder ANTHROPOIDEA Mivart, 1864
Infraorder CATARRHINI
E. Geoffroy Saint-Hilaire, 1812
Parvorder EUCATARRHINI Delson, 1977
Superfamily CERCOPITHECOIDEA Gray, 1821
Family CERCOPITHECIDAE Gray, 1821
Subfamily COLOBINAE Blyth, 1875
Genus Dolichopithecus Depéret, 1889
Dolichopithecus ruscinensis Depéret, 1889
M ATERIAL EXAMINED. NMNH DKV 78, right m3
in fragment of mandible (Fig. 1E); DKV 79, left male
c1 (Fig. 1B); DKV 82, partial left m1? (Fig. 1D); DKV
479, heavily worn female right c1 or possibly left C1
(or even right i2); DKV 482, left male C1 crown
(Fig. 1A); DKV 483, right M3? (Fig. 1C); DKV 484,
left male p3, nearly unworn (Fig. 1F); DKV 485,
heavily worn right female c1 or possibly left i2.
D ESCRIPTION
These specimens are essentially identical in mor-
phology and size to fossils of this species from the
type locality, Perpignan (France; MN 15a), which
has yielded the largest collection to date. D . rusci-
nensis is known from 16 localities, of which half
have produced only one specimen. In addition to
Perpignan and Dorkovo, only Baraolt-Capeni and
Malusteni (Romania; see Delson 1973 and Szalay
& Delson 1979 unless otherwise noted) and the
Kuchurgan sites (Ukraine; Maschenko 1991, see
below) have produced more than three or four iso-
lated teeth, while a well preserved mandible is
known from Megalo Emvolon (Greece; Koufos et
al. 1991). Other nearby localities include Beresti
(Romania), Ciuperceni-2 (Romania; Terzea
1979) and Ptolemais (Greece; Doukas & De
Bruijn 2002), each yielding only one or two teeth.
Delson (1973) reported no significant differences
in size or morphology of this species across locali-
ties. The biochronologic range of this taxon
extends from the Early Pliocene (MN 14) of
Montpellier (France) and Beresti (age following
Radulescu & Samson 2001) to the earlier Late
Pliocene of Balaruc-2 (France; MN 16? if the age
assignment in Fejfar et al. 1997 is accepted;
Radulescu et al. 2003 suggested that Baraolt-
Capeni is better placed in MN 15b). The report-
ed Late Miocene occurrence of D . ruscinensis at
Pestlörinc (near Budapest, Hungary; see Delson
1974) was based on an incorrect age estimate for
that site, which is actually of MN 15 age (Delson
1994).
The dentition of colobines has been described in
detail by Szalay & Delson (1979: 321-323, 383,
384), and these specimens do not provide any
novel features. The m3 (DKV 78) presents clear
colobine features, including the high cristodont
lophids, deep median lingual (talonid) notch
between the lophids, and short trigonid basin
(mesial fovea). The hypoconulid is long,
Fossil monkeys from Dorkovo (Bulgaria)
161
GEODIVERSITAS • 2005 • 27 (1)
displaced buccally and situated nearly in line with
the buccal cusps; there is no tuberculum sextum.
The fragmentary lower molar (DKV 82) is bro-
ken distal to the hypolophid, and it is not possi-
ble to determine definitively to which molar it
belongs; its size suggests an m1, and it may be
noted that its DW is smaller than any of 18
known from Perpignan, while its MW is equal
to the smallest thereof. The upper molar (DKV
483) is identified as an M3 due to its reduced
distal loph and lack of a distal contact facet. The
male p3 (DKV 484) has the typical strongly
sloping mesial honing flange of cercopithecids
corresponding to a large and projecting C1 (as
in DKV 482), which in turn complements the
tall and slender c1 (DKV 79). It is not possible
to distinguish canine teeth by subfamily, but the
lack of any cercopithecine molars suggests that
they are best identified as D . ruscinensis . Two
other worn anterior teeth, as noted above, can-
not be definitively identified. DKV 479 presents
a worn crown and a complete root; so little
crown morphology remains that it might repre-
sent either a female canine (upper or lower) or
possibly a lower i2. DKV 485 is a more incisi-
form tooth: if it is a lower right canine, the
external face is smooth with a slight cingular
base, while internal face rises in inverted “V”
below the cingulum; the distolingual face pres-
ents a complex wear surface, and the root is
gracile. Its morphology, however, appears differ-
ent from that of the equivalent tooth in female
mandible MNHN PER 004. The colobine mor-
phology of these teeth excludes their allocation
to known European Pliocene cercopithecines
(see below), and their size (Table 1) separate
them from Mesopithecus. Neither Delson (1973)
nor any other author has found any temporal
pattern of size change in Dolichopithecus across
its time range.
Delson E. et al.
162
GEODIVERSITAS • 2005 • 27 (1)
A
B
C
D E1 E 2 F
G
F IG. 1. A - F , lower and upper teeth of Dolichopithecus ruscinensis Depéret, 1889; A , left male C1 (DKV 482), lingual view; B , left
male c1 (DKV 79), lingual view; C , right M3? (DKV 483), occlusal view; D , left m1? (DKV 82), occlusal view; E , right m3 in fragment of
corpus (DKV 78), E1 occlusal view, E2 lingual view; F , left male p3 (DKV 484), buccal view; G , upper molar of Mesopithecus
monspessulanus (Gervais, 1849), left M3? (DKV 480), occlusal view. Scale bars: A, B, 1 cm; C-G, 2 cm.
D ISCUSSION
Gremyatski (1958) described several colobine
mandible fragments from the Kuchurgan sites
(Voinichevo river, including Novopetrovka and
other find-spots; probably MN 15 after Fejfar et
al. 1997), which he named Adelopithecus hypsilo-
phus Gremyatski, 1958. Szalay & Delson (1979)
tentatively synonymized the two genera, but they
had not seen the full publication, and the speci-
mens had not yet been located in Moscow.
Alexejeva (1964) had also described a subadult
mandible from Novopetrovka, and in 1982
Delson examined several isolated teeth and casts
from there in the Kiev collection. Maschenko
(1991) assessed the systematic position of Adelo-
pithecus and synonymized it with Dolichopithecus,
but he continued to recognize the Kuchurgan
material as the separate species D . hipsulophus (an
incorrect spelling-lapsus calami). Maschenko
(1991) reported (also in a letter to Delson) that
the mandible which Gremyatski had used as
holotype was lost but that two others remained,
which he compared with casts of Perpignan spec-
imens. He did not mention Alexejeva’s specimen.
Maschenko argued that several features (especially
the tall, narrow and mesiodistally short symphysis
Fossil monkeys from Dorkovo (Bulgaria)
163
GEODIVERSITAS • 2005 • 27 (1)
T ABLE 1. — Measurements (in mm) of lower and upper teeth of Dolichopithecus ruscinensis Depéret, 1889 from Dorkovo compared
to those from the sample from Perpignan. Abbreviations: N , number of specimens; Min., minimum value; Max., maximum value;
for canines and lower third premolars: , male individuals only; , female individuals only; for canines: L , mesiodistal length (note
that length is smaller than width for lowers as thus defined); W , labiolingual width (taken perpendicular to length); H , mesiobuccal
height of crown; RH, height of root below cervix for lowers; for lower premolars: W , maximum buccolingual width; FL, length
of mesiobuccal flange (taken from cusp apex to mesiobuccal end); for molars: MW, maximum mesial buccolingual width, taken
across paraloph or protolophid cusps at cervix; DW, maximum distal buccolingual width, taken across metaloph or hypolophid
cusps at cervix; L ,maximum mesiodistal length, taken near cervix; RW, relative molar width = 100 × DW/MW; RL, relative molar
length = 100 × MW/L.
Dorkovo (DKV No.) Perpignan
479 485 79 484 82 78 NMean Min. Max.
 / / / /
L 5.3 4.5 7.3 12/11 4.6/7.4 3.5/6.3 5.3/8.4
c1
W 7.2 6.4 11.5 12/11 6.9/10.8 6.2/9.1 7.8/12.1
H 7.3 9.8 19.3 11/9 8.5/19.2 6.7/13.3 10.9/24.1
RH 14.5 11.3 20+ 6/5 12.9/23.8 11.9/22.5 14.7/24.7
p3
W 5.5 -/5 -/7.1 -/6.2 -/9.0
FL 18.0 -/5 -/15.2 -/14.5 -/16.0
MW 7.0 18 6.8 6.0 7.7
m1? DW 6.8 18 7.1 6.2 8.0
RW 102.9 18 96.6 89.6 103.1
MW 8.7 15 8.3 7.6 9.3
DW 8.4 18 8.0 7.2 8.8
m3 L 13.5 18 13.1 11.8 14.7
RW 103.0 15 104.9 90.5 112.7
RL 64.4 15 63.9 58.5 72.4
482 483
L 9.3 8/6 6.0/9.1 5.6/8.1 6.5/10.7
C1 W 14.0 8/6 7.9/12.6 7.0/11.4 8.7/13.9
H 30.0 7/5 9.4/22.9 8.2/17.3 10.2/29.0
MW 9.1 10 9.1 8.6 9.7
DW 7.3 10 8.1 6.9 9.3
M3? L 9.5 10 9.7 8.4 10.6
RW 119.7 10 113.0 104.3 124.6
RL 95.8 10 93.9 89.3 102.4
and narrow p3 talonid) of the Kuchurgan speci-
mens significantly distinguished them from those
of Perpignan and other sites, but in fact these
appear to fall within the expected range of varia-
tion. D . hypsilophus Gremyatski, 1958 is here for-
mally synonymized with D . ruscinensis Depéret,
1889.
Genus Mesopithecus Wagner, 1839
Mesopithecus monspessulanus (Gervais, 1849)
M ATERIAL EXAMINED. DKV 480, unworn left M3?
(Fig. 1G); DKV 481, left p4.
D ESCRIPTION
The p4 is comparable to several from Montpellier
and Fornace RDB (Villafranca d’Asti; again, see
Delson 1973 and Szalay & Delson 1979 for
details). The upper dentition of this taxon is rep-
resented by only five other teeth: three upper
molars, one incisor and one canine. The
Dorkovo upper molar lacks a distal contact facet
and is metrically most similar to the presumed
M3 from the Red Crag (see Tables 2 and 3).
D ISCUSSION
Research in progress is evaluating the distinction
between M . monspessulanus and M . pentelicus
Wagner, 1839, and the species allocation of sev-
eral late Miocene samples. Tentatively, it appears
that specimens from MN 13 sites at Baltavar,
Polgardi and Varpalota (Hungary), Monticino
Quarry and Casino (Italy) and Maramena
(Greece) are best included in M . pentelicus, while
long-lost material from Gravitelli cannot be accu-
rately determined (see also Rook 1999). The
Dytiko sites in Greece, especially Dytiko-2
(Bonis et al. 1990, 1997; Koufos et al. 2004;
Delson pers. obs.) appear to yield remains of
both species; if that interpretation is correct,
these would be the oldest known representatives
of M . monspessulanus. The co-occurrence of the
two species requires additional documentation.
Up to now, they have been considered as
chronospecies, but if they are indeed found
together it suggests that the two are biospecies:
M . pentelicus may have given rise to M. monspes-
sulanus and briefly coexisted with its daughter
species before becoming extinct. The latter taxon
continues through MN 14 and 15, with the
youngest individuals perhaps derived from MN
16 localities such as Hajnacka (Slovak Republic),
and the Red Crag (England).
CONCLUSIONS
In their original report on the Dorkovo fauna,
Thomas et al. (1986) mentioned the co-occur-
Delson E. et al.
164
GEODIVERSITAS • 2005 • 27 (1)
T ABLE 2. Measurements (in mm) of the p4 of Mesopithecus
monspessulanus (Gervais, 1849) from Dorkovo (DKV 481)
compared to those for the combined sample from Montpellier
and Fornace RDB (Villafranca d’Asti). Abbreviations: N , number
of specimens; Min., minimum value; Max., maximum value;
W ,maximum buccolingual width; L , length; FL, length of mesio-
buccal flange (taken from cusp apex to mesiobuccal end);
RW,relative width = 100 × W/L.
Dorkovo Montpellier and
DKV 481 Fornace/Villafranca d’Asti
p4 NMean Min. Max.
W 4.9 64.3 4.0 4.4
L 6.1 65.4 4.8 6.1
FL 5.4 35.0 4.8 5.3
RW 80.3 680.3 70.5 89.6
T ABLE 3. — Measurements (in mm) of the M3(?) of Mesopithecus
monspessulanus (Gervais, 1849) from Dorkovo (DKV 480)
compared to those for all other known upper molars attributed
to the species. Abbreviations: MW, maximum mesial buccolin-
gual width, taken across paraloph cusps at cervix; DW, maxi-
mum distal buccolingual width, taken across metaloph cusps at
cervix; L , maximum mesiodistal length, taken near cervix;
RW,relative molar width = 100 × DW/MW; RL, relative molar
length = 100 × MW/L.
DKV 480 Wölfersheim Ivanovce Red
Crag
M3? M1? M2? M3?
MW 7.3 5.2 6.0 7.0
DW 6.1 5.1 5.8 6.5
L 7.9 5.6 6.4 7.6
RW 119.7 102.0 103.0 108.0
RL 92.4 93.0 94.0 92.0
rence of Dolichopithecus and Mesopithecus as a
rare phenomenon in the Ruscinian, and Spassov
(2003) noted that this association is characteristic
for the Ruscinian of the southern half of Europe.
In fact, four species of cercopithecid are known
in MN 14-16, and their joint presence is not
uncommon. In addition to the two colobines
mentioned, macaques (broadly referable to
Macaca cf. sylvanus subspp.) appeared in Europe
in the latest Miocene (Köhler et al. 2000) and
persisted into the Late Pleistocene, while
Paradolichopithecus arvernensis is now known
from the Late Pliocene of Senèze (France),
Graunceanu (Romania), and Vatera-F (Lesvos,
Greece), with fragmentary remains of smaller-
sized individuals occurring at earlier Pliocene
localities. The most common association is that
at Dorkovo: Mesopithecus and Dolichopithecus co-
occur also at Perpignan, Baraolt-Capeni, and
Wölfersheim (Germany); they are joined by
Macaca at Montpellier and by cf. Paradoli-
chopithecus at Malusteni. Macaca and Dolicho-
pithecus co-occur at Cova Bonica (Spain) and
Balaruc-2, while Macaca and Mesopithecus are
found at Fornace RDB (Italy). Tesakov &
Maschenko (1992) described an isolated tooth
from Grebeniki-2 (a Kuchurgan locality) which
they identified as a macaque, but which appears
colobine (perhaps M. monspessulanus) from their
drawing; in either case, a second taxon accompa-
nies Dolichopithecus in the Kuchurgan region.
Among younger localities, the only co-occurrence
is of Paradolichopithecus and cf. Macaca at
Senèze. There does not appear to be any temporal
pattern in these associations.
Acknowledgements
We thank the curators of numerous paleontolog-
ical collections who have permitted us to examine
specimens in their care. We thank the two
anonymous reviewers for their useful comments
on a previous version of this manuscript. We
(especially NS and HT) express our gratitude to
the Municipality of Dorkovo and to the ex-
mayor Yanko Duzov for their support during the
excavations (1985-1987), which were partially
supported by the Bulgarian Academy of Sciences.
ED’s research and travel to study European fossil
Cercopithecidae has been supported by a series of
grants from the PSC-CUNY Faculty Research
Awards Program, most recently 64542-33 and
65509-34, for which he is most grateful.
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    • "This is however only true for the permanent dentition, because not all the deciduous dentition is known from Pikermi (i.e., the anterior deciduous teeth are missing; see Zapfe, 1991) and sample sizes for the available deciduous teeth are much smaller. Unfortunately, the attribution at the species level of the more complete sample of deciduous teeth from Maramena is open to different interpretations , having been previously attributed to either M. pentelicus (Kullmer and Doukas, 1995; Delson et al., 2005) or Mesopithecus sp. (Koufos, 2009a). "
    [Show abstract] [Hide abstract] ABSTRACT: We report dental remains of the extinct colobine monkey Mesopithecus from the Turolian (MN13, Late Miocene, ca. 6.23 Ma) locality of Venta del Moro (Valencia, Spain). They include most of the deciduous dentition and the unerupted germs of the first molars of a single infantile individual, as well as two lower left lateral incisors from two additional individuals. On the basis of morphometric comparisons, mainly based on the M1s, these remains are attributed to the Late Miocene species Mesopithecus pentelicus. They represent a significant addition to the knowledge of the deciduous dentition of this taxon, much less well-known than the permanent dentition. Although this genus was widely distributed from the Late Miocene through the Pliocene across Europe, southwestern Asia, Pakistan, and China, until now its occurence in the Late Miocene of the Iberian Peninsula had not been documented conclusively. Hence, the reported remains considerably enlarge southwestwards the known geographic distribution of Mesopithecus. The presence of this genus at Venta del Moro must be understood within the framework of the significant faunal turnover that took place in European faunas during the latest Turolian (the second Messinian mammalian dispersal), which is further documented at this locality by the occurrence of other eastern immigrants. At the same time, the presence of M. pentelicus at this site agrees well with previous paleoenvironmental and sedimentological evidence, indicating a lacustrine depositional environment with strong hydrologic seasonality.
    Full-text · Article · Nov 2015
    • "In the transitional Turolian/Ruscinian faunas of the studied area, Mesopithecus is known from MAR (Kullmer and Doukas 1995), but, as the available material consists of isolated teeth, it is difficult to say if it is represented by one or two forms. The typical Ruscinian cercopithecoids Dolichopithecus ruscinensis and M. monspessulanus co-exist in the early Ruscinian locality of Dorkovo (Delson et al. 2005). The main late Miocene carnivore genera (Adcrocuta, Amphimachairodus, Paramachaerodus, Metailurus) became extinct, whereas other ones entered in the early Pliocene with new species (Promeles macedonicus, Martes lefkonensis) and some made their first occurrence (Lutra affinis). "
    [Show abstract] [Hide abstract] ABSTRACT: Even though there are numerous late Miocene mammal localities in the southern Balkans, those of late Turolian and early Ruscinian age are uncommon. Using the available data, mainly from Greece and Bulgaria, we compiled information about the faunal and palaeoenvironmental changes at the Miocene/Pliocene boundary. The analysis of the faunal elements indicates that several Miocene taxa disappeared, whereas new taxa appeared. The faunal composition of the mammal zones suggests a decrease of the bovids and giraffids and an increase of cervids and suids from MN 12 to MN 14. The available faunal assemblages from zones MN 12–MN 14 are compared with a set of modern and fossil assemblages from known environments to determine their palaeoenvironment. The results indicate an increase of the closed character of the environment from MN 12 to MN 14, suggesting a gradual increase of the humidity and the development of more forestial conditions. The rodent faunas from northern Greece ranging from MN 13 to MN 14 show a characteristic aridification of the biotopes and the introduction of African elements in the last part of the MN 13 (during the Messinian Salinity Crisis), with a subsequent return of humid-and/or-forested biotopes towards the MN 13/14 boundary. The faunal and sedimentological data for the Greek localities correlated with the Turolian/Ruscinian boundary suggests a general open landscape with several water spots of variable sizes surrounded by dense forests. © 2015, Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg.
    Full-text · Article · Jul 2015
    • "On the other hand, it is interesting that Myanmarcolobus shows similarity in lower molar morphology to Dolichopithecus in having relatively obliquely running transverse ridges, a mesiodistally long asymmetrical median lingual notch, a largely concave median buccal cleft, and a distinct mesiobuccal notch on the lower molars (Figs. 5, 8 and 9). Myanmarcolobus is most similar to Dolichopithecus, especially to the specimens from the Early Pliocene of southeastern Europe, such as Dolichopithecus hypsilophus (Fig. 5b) and Dolichopithecus balcanicus, among fossil colobines discovered in Eurasia (Koufos et al., 1991; Maschenko, 1991 Maschenko, , 2005 Delson et al., 2005; Spassov and Geraads, 2007). However, large size differences in the lower molars between Myanmarcolobus and Dolichopithecus and the cohesiveness of lower molar size in Dolichopithecus suggest heterogeneity between them (Fig. 7). "
    [Show abstract] [Hide abstract] ABSTRACT: Here we report two kinds of colobine fossils discovered from the latest Miocene/Early Pliocene Irrawaddy sediments of the Chaingzauk area, central Myanmar. A left mandibular corpus fragment preserving M1-3 is named as a new genus and species, Myanmarcolobus yawensis. Isolated upper (M1?) and lower (M2) molars are tentatively identified as Colobinae gen. et sp. indet. Although both forms are medium-sized colobines, they are quite different from each other in M2 morphology. The isolated teeth of the latter show typical colobine-type features, so it is difficult to identify their taxonomic position, whereas lower molars of Myanmarcolobus have unique features, such as a trapezoid-shaped long median lingual notch, a deeply concave median buccal cleft, a strongly developed mesiobuccal notch, and rather obliquely running transverse lophids. Compared with fossil and living Eurasian colobine genera, Myanmarcolobus is most similar in lower molar morphology to the Pliocene Dolichopithecus of Europe rather than to any Asian forms. In Dolichopithecus, however, the tooth size is much larger and the median lingual notch is mesiodistally much shorter than that of Myanmarcolobus. The discovery of Myanmarcolobus in central Myanmar is the oldest fossil record in Southeast Asia not only of colobine but also of cercopithecid monkeys and raises many questions regarding the evolutionary history of Asian colobine monkeys.
    Full-text · Article · Jun 2015
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