Age or paleoenvironment?
What is the cause of the differences between
two Early Miocene small-mammal
assemblages from Kaplangı (Turkey)?
Anne Dangremond
3118584
June 29, 2012
Supervisor: dr W. Wessels
Content
Content .................................................................................................................................................... 2
Abstract ................................................................................................................................................... 4
Introduction............................................................................................................................................. 5
Geological history ................................................................................................................................ 6
Materials and methods ........................................................................................................................... 8
Anomalomys aliveriensis ....................................................................................................................... 10
Description Kaplangı 1 ....................................................................................................................... 10
Description Kaplangı 2 ....................................................................................................................... 13
Discussion .......................................................................................................................................... 15
Evolutionary context of Anomalomys ........................................................................................... 15
Mirrabella cf. crenulata ......................................................................................................................... 16
Description Kaplangı 1 ....................................................................................................................... 16
Discussion .......................................................................................................................................... 17
Comparing with M. crenulata from Keseköy................................................................................. 18
Comparing with other species....................................................................................................... 18
Evolutionary context of Mirrabella ............................................................................................... 18
Eumyarion carbonicus ........................................................................................................................... 19
Description Kaplangı 1 ....................................................................................................................... 21
Discussion .......................................................................................................................................... 22
Comparing E. carbonicus to other E. carbonicus from Harami 1 .................................................. 22
Comparing E. carbonicus to other Eumyarion species .................................................................. 23
Evolutionary context of Eumyarion ............................................................................................... 23
Cricetodon aliveriensis ........................................................................................................................... 25
Description Kaplangı 2 ....................................................................................................................... 27
Discussion .......................................................................................................................................... 29
Comparing the C. Aliveriensis from Kaplangı 2 to C. aliveriensis from Aliveri .............................. 29
Comparing the C. Aliveriensis from Kaplangı 2 to C. tobieni from Horlak 1a ............................... 30
Evolutionary context of Cricetodon .............................................................................................. 31
Democricetodon gracilis ........................................................................................................................ 32
Description Kaplangı 1 ....................................................................................................................... 34
Description Kaplangı 2 ....................................................................................................................... 37
Discussion .......................................................................................................................................... 39
Comparing D. gracilis from Kaplangı to D. gracilis from Sandelzhausen ....................................... 39
Comparison of D. gracilis of Kaplangı to those of Aliveri .............................................................. 40
Comparison of D. gracilis of Kaplangı to those of Karydiá 1 ......................................................... 41
Comparison of D. gracilis of Kaplangı to those of Karydiá 2 ......................................................... 41
Comparing D.gracilis to D. doukasi (Keseköy) ............................................................................... 42
Comparing D. gracilis to D. franconicus (Aliveri) ........................................................................... 42
Evolutionary context of Democricetodon ..................................................................................... 43
Gliridae .................................................................................................................................................. 44
Description ........................................................................................................................................ 46
Discussion .......................................................................................................................................... 46
Discussion and conclusions ................................................................................................................... 47
Age ..................................................................................................................................................... 47
Palaeoenvironment ........................................................................................................................... 48
Migration patterns ............................................................................................................................ 48
Anomolomys aliveriensis ............................................................................................................... 49
Democricetodon gracilis ................................................................................................................ 49
Eumyarion carbonicus ................................................................................................................... 49
Cricetodon aliveriensis .................................................................................................................. 49
Mirrabella cf. crenulata ................................................................................................................. 50
Conclusions........................................................................................................................................ 51
References ............................................................................................................................................. 52
Appendix 1 – Fauna list ......................................................................................................................... 56
Appendix 2: Geological history .............................................................................................................. 57
Abstract
Two assemblages of small mammals near Kaplangı (Anatolia) are described to determine if their
respective differences are due to a difference in palaeoenvironment, age, or both. The main species
composing the assemblages are Anomalomys aliveriensis, Democricetodon gracilis, Eumyarion
carbonicus, Cricetodon aliveriensis, Mirrabella cf. crenulata, Vasseuromys sp. and Glirudinus cf.
euryodon. Their respective ages are determined to be Late MN3 to Early MN4 for both assemblages.
Differences between the populations are determined to be due to a slightly wetter environment in
Kaplangı 2. The Kaplangı assemblages are useful in confirming current theories about faunal
migration from Anatolia to Europe during the Miocene.
Introduction
In the 1960’s the systematic search for fossil mammals started in Anatolia. The lignite exploration
required a good insight in the tectonic and sedimentary history of the complex continental basins
(Ünay et al; 2003). At this day, there is still a lot unknown about many of the Anatolian sedimentary
basins and biostratigraphical research is still necessary. Another reason to do research on fossil
mammals is because they serve as a tool for paleogeographical and paleoclimatological
reconstructions. Furthermore many of the migration patterns of mammals between Europe and Asia
are still not understood and a good biostratigraphical record could help to solve many of these items.
In 1975 Mein defined the Mammal Neogene biochonological framework (MN) as an attempt to
systematize the changes of the Neogene fossil mammal paleofauna’s in the Eastern Mediterranean
and Western Europe. The Neogene is divided into 17 different biozones, however the time
boundaries of these zones are not well defined for the whole of Europe (Agustí et al; 2001). This is
because they are different for each region, therefore it is not an integral system. Even though this
classification is often used for the Eastern Mediterranean, it is mainly suitable for Western Europe.
Therefore Ünay et al (2003) proposed a new biochronological zonation for Anatolia. This biozonation
system divides the Neogene in 16 zones (A to P) based on the stage-evolution of the Muroidea and
Zapodidae. Ünay et al (2003) also correlated these zones to the MN-zones. Kaplangı was preliminarily
assigned to zone E. This will be briefly evaluated with our new results in the discussion.
In this article the MN-zonation will be used because it is the most suitable for correlating the results
from Kaplangı to the available information from Greece and other European localities.
Figure 1 - Map of the North-Eastern Mediterranean region, with the most important locations used in this article,
adapted from Fortelius (2012).
The aim of this article is to determine whether the difference between two Early Miocene smallmammal assemblages found around Kaplangı (Turkey) is due to a difference in age or
paleoenvironment (or both). The geographical distance between the Kaplangı 1 and Kaplangı 2
localities is 150 meters. However, because of the complex geological history of the area,
stratigraphical control is missing. The Kaplangı 1 specimens were found in clayey sediments and the
Kaplangı 2 specimens in clayey sediments with some coal seams.
For this research the molars of the small mammal specimens found in the assemblages were
measured and morphologically compared to other molars to determine the species that are in the
different assemblages. The species composition will be used as an indicator for both the biozone of
the assemblages as well as for their respective palaeoenvirionment
Geological history
Turkey can geologically be divided into three main tectonic units with different characteristics: The
Arabian platform, the Pontides and the Anatolides-Taurides (see figure 1) (Okay, 2008). These units
used to be separated by the Thethys ocean. This ocean existed as a large embayment between
Gondwana and Laurasia at least since the Carboniferous (Stampfli and Borel; 2002), and can be
subdivided into the Neo-Tethys and the Paleo-Tethys (Şengör ; 1984 and 1987, Stampfli; 2000,
Robertson et al; 2004). The sutures that remain after the Tethys oceans has closed indicate the
boundaries between the different geological units. The Izmir-Ankara-Erzincan suture marks the
boundary between the Pontides and the Anatolides-Taurides, and the Assyrian suture connects the
Arabian platform to the Anatolides-Taurides (Okay; 2008).
The Kaplangı area is located in the western part of Anatolia, south of the Afyon zone on the
Anatolide-Tauride Block (fig 2). The coordinates for this location are 38.768004,29.903412.
The Anatolide-Tauride terrane contains a variety of metamorphic, structural and stratigraphic
features that is used to divide it in different zones. However, these zones share some common
stratigraphic elements that distinguish the Anatolide-Tauride terrane as a single paleogeographic
entity. These elements contain a late Precambrian basement, a Paleozoic mixed clastic-carbonate
succession and a thick Upper Triassic to Upper Cretaceous carbonate sequence. This carbonate
sequence was the result of an extensive shallow marine carbonate platform that caused the
deposition of several thousand meters of carbonate during the Mesozoic (Okay, 2008).
During the Late Cretaceous, several basins started to form around the Central Anatolian Crystalline
Complex. These basins might have been interconnected, and were filled with clastic sediments of
Late Cretaceous – Eocene age. By the end of the Oligocene, most of the Anatolian terrains were
merged into a single landmass. The only exception was a narrow seaway between the Arabian
Platform and the Anatolides-Taurides in Southeast Anatolia. This seaway was closed during the
Miocene when the Arabian and Anatolian plates collided. During the Neogene, the tectonic regime
was characterized by extension and strike-slip faulting, creating space for the Miocene basins which
form the present day tectonic frame of Anatolia (Akgün et al, 2007; Okay, 2008).
So it is clear that the Arabian plate and the Anatolian plate collided during the Miocene, creating a
passage for the exchange of different animals from one plate to another. However, it is still unclear
when the connection between Anatolia and Europe developed. This connection probably started to
form, during the Early or Middle Miocene, but this is not definitive yet (Popov et al; 2004).
During the Miocene, most of Central and Western Anatolia was covered with lakes surrounded by
swamps. These subsiding lakes were filled with a sequence of different sediments that were supplied
by volcanoes and uplifted ranges of older metamorphic and sedimentary rocks (Okay, 2008).
Figure 2 - Tectonic map of the North-Eastern Mediterranean region. The red dot indicates the Kaplangı area (After Okay
et al 2000)
Materials and methods
Samples were taken from two different locations (Kaplangı 1 and Kaplangı 2) by Hans de Bruijn and
Gerçek Saraç. The amount of molars per genus and location are given in appendix 1. The amount of
elements per genus are specified in their specific descriptions. A molar may be indicated as
incomplete in cases where it is broken or so worn that some features cannot be identified.
For this article the molars of Eumyarion, Mirrabella, Democricetodon, Cricetodon, Anomalomys and
Gliridae are described. The molars of the other families are mentioned in the fauna list, but are not
described in further detail and therefore not determined on species level.. In total, the Kaplangı 1
assemblage contains 115 molars and the Kaplangı 2 contains 83.
The molars were measured with a Leitz Ortholux microscope with a mechanical stage and measuring
clocks. The measurements are given in millimeter units. The specialized statistics program SPSS was
used for the statistical analyses and graphs.
The molars are described based on their main characteristics. The molars of the upper jaw are
indicated as M1, M2 and M3, those in the lower jaw as M1, M2 and M3. The nomenclature used for the
description of the Cricetidae is after Freudenthal et al (1994), with the following alterations:
-
In the M1 an anteromesoloph is added. This is defined as a ridge that is protruding from
the anterolophule towards the labial edge of the molar. This ridge is also present in the
M1 and is then called anteromesolophid.
-
Because the anterocone spur and the paracone spur are almost always connected, they
are combined to form one ridge, the anterolophule.
-
In the M1 the anteroloph is divided in a labial and a lingual part.
-
Freudenthal(1994) only indicated one protolophule, while in this article an anterior and a
posterior protolophule are indicated.
-
Some molars in the upper jaw have a ridge between the paracone and metacone or
between the protocone and the hypocone.
-
In the lower jaw there can be a ridge between the protoconid and hypoconid and
between the metaconid and the entoconid.
-
In some teeth an anterior spur of the hypocone or hypoconid is present
-
In the M3 a protosinus is present
-
Some of the M3 have a posterior paracone spur or a ridge between the paracone and
metacone
-
Old entoloph = longitudinal crest
The orientation of a protolophule or metalophule is defined as the direction in which the labial part
of the ridge is bending. The orientation of the metalophulid and hypolophid is the direction in which
the lingual part is bending.
The mesoloph or mesolophid can be short medium or long. When the length of the mesoloph(id) is
up to 1/3 of the distance between the entoloph/ectolophid and the edge of the molar it is indicated
as short. When it is 1/3 to 2/3 of this distance it is indicated as being of medium length. The
mesolophid is long when it is longer than 2/3 of the distance between the entoloph/ectolophid and
the edge of the molar. The mesoloph(id) does not need to reach the edge of the molar to be
indicated as long. This is also the case with the ectomesoloph/ectomesolophid, the lingual and labial
anteroloph/anterolophid and posteroloph/posterolophid.
Figure 3 - Terminology of the Cricetidae molars (Theocharopoulos (2000) after Freudenthal et al., 1994)
Anomalomys aliveriensis
Order: Rodentia - Bowdich, 1821
Family: Muridae - Illiger, 1811
Genus: Anomalomys - Gaillard, 1900
Species: Anomalomys aliveriensis - Klein Hofmeijer and de Bruijn, 1985
Type locality: Aliveri (Greece)
Type horizon: Early Miocene (MN4)
Geographical distribution and biochronological range: Aliveri(MN4), Dededag(MN4), Karydia(MN4).
(Fortelius, 2012)
Original diagnosis (Klein Hofmeijer and de Bruijn, 1985): Anomalomys with a relatively narrow M1
and a long M3. The protoloph and metaloph of the M1 and M2 are well developed and transverse. The
posterior arm of the protoconid and the posterior arm of hypoconid are strong. The mesolophids of
the M1 and M2 are always long.
Description Kaplangı 1
Table 1 - measurements of A. aliveriensis from Kaplangı 1 (in mm).
Length
N
M1/
M2/
M3/
M/1
M/2
M/3
3
2
1
3
3
0
Width
Range
Mean Range
Mean
1,39 - 1,47 1,44 0,98 - 1,04 1,01
1,11 - 1,31 1,21 0,95 - 1,01 0,98
0,8 0,80
0,87 0,87
1,33 - 1,46 1,38 0,81 - 0,88 0,86
1,18 - 1,32 1,23 0,96 - 1,02 0,99
-
Figure 4 - Measurements of A. aliveriensis from Kaplangı 1 and 2 compared to other Anomalomys species (in mm)
Although often strongly worn, all molars are high crowned.
M1 – The anterocone is undivided and ridgelike. This complex is located at the labial side of the
molar. The labial anteroloph is long and connected to the anteromesoloph. This anteromesoloph is
long in 2(3) and of medium length in 1(3) molar. There lingual anteroloph is absent. The
anterolophule is oblique. The protosinus is absent. The single protolophule is transverse and
connects lingually with the longitudinal ridge at the point where the sinus is deepest. The mesoloph
is long and reaches the labial border, in 1(3) molar it is connected to the anterior spur of the
metacone. The mesosinus is open in 1(2) and partially closed the other molar. The longitudinal ridge
is straight. The sinus is short but strongly bended towards the anterior side, it is narrow in 2(3) and
broad in 1(3) molar. The metalophule is absent in 1(3), bending towards the anterior side in 1(3) and
transversal in 1(3) molar. In 1(2) molar, a strongly developed anterior spur of the metacone is
present on the lingual edge of the molar and it is connected to the mesoloph. In the other molar, this
anterior spur is much shorter and not connected to the mesoloph. The posterosinus is always narrow
and is closed in 2(3) and open in 1(3) molar. Labial posteroloph is long in 2(3) and of medium length
in 1(3) molar. The lingual posteroloph is absent. The paracone and metacone are much larger than
the protocone and metacone.
M2 – The labial anteroloph is long, it is connected to the paracone in 1(2) molar. The lingual
anteroloph is absent. The anterolophule is complete and oblique. A posterior spur of the paracone is
medium developed. The protolophule is transversal in 1(2) and bending towards the posterior side in
1(2) molar, it connects to the longitudinal ridge at the point where the sinus is the deepest. The
lingual ridge is continuous in 1(2), and interrupted in 1(2) molar. The mesoloph is long, it is
connected to the paracone spur in 1(2) molar. The open sinus is very short and narrow, it bends
strongly towards the anterior side. In 1(2) molar the longitudinal ridge is straight, in the other 1(2) it
is absent. The metalophule is transversal. The metacone has an anterior spur on the longitudinal
edge of the molar which is medium developed in 1(2) and strongly developed in 1(2) molar, it is
connected to the mesoloph in 1(2) molar. The posterosinus is narrow, it is open in 1(2), and closed in
the other molar. The labial posteroloph is long, it is connected to the metacone in 1(2) molar.
M3 - The only M3 present is too worn to determine its features.
M1 – The anteroconid and metaconid are merged and are forming one irregular shaped complex.
The anteroconid part is located at the labial side of the molar and is rounded at the anterior side in
1(2) molar. There is no anterolophid or anterolophulid. The anteromesolophid is short in 2(3) and of
medium length in 1(3) molar. The mesolophid is long, and the ectomesolophid is very short. The
longitudinal ridge is straight in 2(3) and bended in 1(3) molar. The sinusid is broad and short, and is
bending strongly towards the posterior side of the molar. The hypolophulid is transverse and
connects to the longitudinal ridge just in front of the hypoconid in 2(3) molars. In the other molar it is
absent. The entoconid has a medium developed posterior spur. The hypoconid branch is long in 1(2),
and of medium length in 1(2) molar, in the other molar it is not clearly visible, but the lingual
posteroloph is very broad, so it is probably incorporated in this ridge. The posterosinus is closed and
it is broad in 2(3) and narrow in 1(3). The lingual posteroloph is long and connected to the entoconid
spur.
M2 – The metaconid and thick anterlophulid are merged. In 1(3) molar, a lingual anterolophule of
medium length is present. The anterolophulid is absent. A transverse metalophulid is present
between the anteroconid and the metaconid. The anteromesolophid is of medium length. The
posterior spur of the metaconid is strongly developed. The mesolophid is long. The longitudinal ridge
is bended. The sinusid is short and strongly bended towards the posterior side. The hypolophulid is
transverse and it is connected to the longitudinal ridge at the point where the sinusid is deepest.
Both the lingual posterolopnghid and the hypoconid branch are long and the posterolophid is
connected to the entoconid. Therefore the broad posterosinusid is closed.
Description Kaplangı 2
Table 2 - measurements of A. aliveriensis from Kaplangı 2(in mm).
Length
N
M1/
M2/
M3/
M/1
M/2
M/3
Range
2
0
1
3
2
1
1.47
0,97
1,38 - 1,49
1,31 - 1,47
1,02
Width
Mean Range
Mean
1.47
1,00-1.08 1,04
0.97
0.89 0.89
1,44 0,83 - 0,90 0,87
1,39 1,00 - 1,06 1,03
1,02
0,9
0,9
All molars from Kaplangı 2 have higher crowns than those from Kaplangı 1. They are usually less worn
than those from Kaplangı 1.
M1 – The anterocone is ridgelike and fused with the labial anteroloph, and is located at the labial side
of the molar. The lingual anteroloph is absent. The anterolophule is complete and longitudinal, it
merges fluently into the anterocone/anteroloph. The anteromesoloph is long in 1(2) and of medium
length in the other. In 1(2) a strongly developed posterior spur of the paracone is present. The
paracone has an irregular shape. The protolophule is complete and transversal, and it connects
lingually with the longitudinal ridge at the point where the sinus is deepest. The mesoloph is long and
always reaches the labial border of the molar. The longitudinal ridge is slightly bended. The short and
narrow sinus is strongly bended towards the anterior side of the molar. The metalophule is
transverse in 1(2) and absent in the other molar. The metacone has a medium developed anterior
spur on the labial edge of the molar. The labial posteroloph is long and connected to the metacone.
Therefore the narrow posterosinus is closed.
M3 – The only M3 present is too worn to determine its features.
M1 – The anteroconid, metaconid and a short anteromesoloph are forming an irregular shaped
construction which is rounded at the anterior side. It is located at the labial side in 2(3) and in the
middle of the molar in 1(3) specimen. The anterolophulid is absent. In 1(3) molar, a short anterior
arm of the protoconid is present. The mesolophid is long, and the mesosinusid is always open. The
ectomesolophid is very short. The longitudinal ridge is straight in 2(3) and bended in 1(3) molar. The
sinusid is strongly bended towards the posterior side and is broad in 2(3) and narrow in 1(3) molar.
The hypolophulid is transverse and is connected to the longitudinal ridge at the point where the sinus
is deepest. The hypoconid branch is long in 2(3) in the other molar it is not clearly visible, but the
lingual posteroloph is broad, so it is probably fused when worn. The entoconid has a posterior spur
which is connected to the long lingual posterolophid. Therefore the posterosinusid is narrow and
closed.
M2 – The anterolophid and metaconid are strongly connected into an irregular shaped construction.
A small anterosinusid is present. The transverse metalophulid is connected to the labial edge of the
molar at the point where the mesolophid ends. An open transverse sinus is present between the
protoconid and metaconid. The anterolophulid is absent. The posterior spur of the metaconid is
strongly developed, but it is not connected to the long mesolophid. The longitudinal ridge is bended.
The short and narrow sinusid is strongly bended towards the posterior side. The transverse
hypolophulid is connected to the longitudinal ridge just before the hypoconid, at the point where the
sinus is deepest. Both the lingual posterolophid and hypoconid branch are long, but not connected to
the entoconid. Therefore the posterosinusid is open, it is broad in 1(2) and narrow in the other
molar.
M3 – The anterolophid, protoconid and metaconid are merged into a complex structure, both cones
are connected by a lingual anteroloph which is of medium length, and by a transversal metalophulid.
The mesolophid is of medium length and nearly reaches the ridge between the metaconid and the
entoconid. The longitudinal ridge is not complete. The hypolophulid is transverse and ends against
the longitudinal ridge at the point where it is interrupted. This is also the part where the sinusid is
deepest, this sinusid is narrow and is bending strongly towards the posterior side. The hypoconid
spur is of medium length and ends against the long lingual posterolophid.
Discussion
The Anomalomys specimens from Kaplangı 1 and 2 were compared to A. aliveriensis from Aliveri, and
A. minor from Germany which are similar in size to Kaplangı. Based on these comparisons, it was
concluded that the specimens from Kaplangı 1 and 2 belong to the species A. aliveriensis.
When comparing A. aliveriensis from Kaplangı to A. minor it shows that they are similar in size, but its
M1 is narrower, and the M3 is longer. The anterior and posterior arm of the protoconid
(anteromesolophid and mesolophid) are always strong in the M2. The anterior arm of the protoconid
(anteromesoloph) and the anterior arm of the hypoconid (mesoloph) are well developed in all M1. In
the M2, the protolophule and metalophule are always complete and approximately transverse. The
posterolophid and entolophid of the M2 are always connected via the posterior portion of the
longitudinal ridge. The posterior arm of the hypoconid is strong in most molars.
Comparing Anomalomys aliveriensis from Aliveri and Anomalomys aliveriensis from Kaplangı 1 and 2,
shows that major differences between the different populations are absent.
Evolutionary context of Anomalomys
Due to a limited fossil record, there is little knowledge about the origin of Anomalomys. This genus is
mainly present in Eastern and Central Europe, but some older species (A. minor and A. anomalomys)
are known from Greece and Western Turkey (all MN4) (Bolliger, 1999; Fortelius, 2012). Fejfar (1972)
proposed that Anomalomys originated from an unknown Oligocene Asian group of cricetids. The
presence of Anomalomys aliveriensis, the oldest representative of the genus Anomalomys (Klein
Hofmeijer and de Bruijn, 1985), in Kaplangı is in agreement with this idea. Kaplangı is the most
eastern locality were A. aliveriensis has been found, so if Anlomalomys indeed originates from Asia
this is probably one of the oldest specimens found yet. Considering the specimens from Greece and
Turkey are from MN4, the Kaplangı specimens are likely to be of the same age or possibly older.
Mirrabella cf. crenulata
Order: Rodentia - Bowdich, 1821
Family: Cricetidae - Fischer de Waldheim, 1817
Subfamily: Eumyarioninae - Ünay, 1989
Genus: Mirrabella - De Bruijn et al.,1987
Soort: Mirrabella crenulata - de Bruijn and Saraç, 1992
Type locality: Keseköy (a 15cm thick paleosol with gastropods exposed at about 25m above the
lignite seam that is exploited in the Keseköy mine).
Type horizon: Early Miocene (MN 3)
Geographical distribution and biochronological range: Keseköy (Turkey) (de Bruijn and Saraç, 1992)
Original diagnosis (de Bruijn and Saraç, 1992): A Mirrabella with low-crowned cheek teeth. The labial
sinusoid of the M1 is asymmetric because the anterior part of the true longitudinal ridge is absent.
The hypolophid is usually absent in the M1 and incomplete in the M2. The sinus of the M1 is
moderately wide. The entomesoloph is weak or absent. The metalophule of the M1 has a double
lingual connection. The posterolabial corner of the M1 is angular.
Description Kaplangı 1
Table 3 - measurements of M. crenulata from Kaplangı 1(in mm).
Length
N
M1/
M2/
M3/
M/1
M/2
M/3
Range
0
2
0
0
0
0
2,16 - 2,32
-
Width
Mean Range
Mean
2,24 1,68 - 1,79 1,74
-
Figure 5 - Measurements of M. crenulata from Kaplangı 1 compared to other Mirrabella species (in mm).
M2 – Both molars are relatively high crowned. The labial anterolophule is long and connected to the
paracone. The lingual anteroloph is only just indicated. The anteroloph is very short and oblique. The
protolophule is single and located at the posterior side of the paracone. It bends towards the
anterior side. The mesoloph is long and is slightly bending towards the posterior side, it is strongly
connected to the hypocone. It is connected to the strongly developed posterior spur of the paracone
in 1(2). The entomesoloph is absent. The sinus is very deep, narrow and transverse. The metalophule
is single in 1(2) and double in the other 1(2), in this molar the anterior metalophule is incomplete and
very short. The complete metalophules are located at the posterior side of the metaconid and are
bending towards the anterior side. Because the short posteroloph is merged to the metacone, the
posterosinus is very narrow or absent. The posterolabial corner of the molar is absent.
Discussion
Because there are only two molars present of the genus Mirrabella, the species determination will be
less accurate. However, based on the shape of the protocone and hypocone and on the crenulated
enamel, the specimen from Kaplangı 1 are determined to be cf. crenulata
Comparing with M. crenulata from Keseköy
When comparing the Mirrabella cf. crenulata from Kaplangı 1 to those from Keseköy, there are a few
differences. In Keseköy there is a double protolophule and metalophule (although sometimes
incomplete). In Kaplangı 1 the protolophule is always single, and the metalophule is single in 1(2) and
incomplete double in 1(2). In Keseköy there is an extra connection between the protolophule 1 and
the anteroloph, but because the protolophule 1 is absent in Kaplangı 1, there is no such connection.
Comparing with other species
According to the Bruijn and Saraç (1992), M. anatolica can be distinguished from M. crenulata and M.
tuberosa by a better developed metalophule in the M2 . In Kaplangı 1 this metalophule is not strongly
developed.
The main difference between the M. crenulata and M. tuberosa in the M2 can be seen in the shape of
the protocone and hypocone. They are more crescent-shaped in M. tuberosa. In Kaplangı 1, they are
plumper and round which indicates that the specimen from Kaplangı 1 are indeed M. crenulata.
However, because there are only 2 molars available from Kaplangı 1, it is not possible to eliminate all
uncertainty.
Evolutionary context of Mirrabella
Mirabella is not found in many localities and when it is present it is rare. There is only one other
location where M. crenulata was found which is Keseköy (Turkey). The age of these molars is
determined to be MN3. According to de Bruijn and Saraç (1992) the protolophule and metalophule
are becoming more suppressed in younger species in the Mirabella lineage. In the M. crenulata from
Keseköy, the protolophule and metalophule are double while in those from Kaplangı are (mainly)
single. This could indicate that the Kaplangı specimen are slightly younger than those from Keseköy,
however, the limited amount of specimens does not allow formulating a definitive statement on this
subject.
Eumyarion carbonicus
Order: Rodentia - Bowdich, 1821
Familie: Cricetidae - Fischer de Waldheim, 1817
Subfamily: Eumyarioninae - Ünay, 1989
Genus: Eumyarion - Thaler, 1966
Species: E. carbonicus - de Bruijn and Saraç, 1991
Type locality: Harami 1, (top of the main lignite bed exploited in the Harami mines southeast of
Akşehir, Anatolia, Turkey.
Type horizon: Early Miocene, MN 1 or 2
Geographical distribution and biochronological range: Harami (Turkey) MN 1-2
Original diagnosis (de Bruijn and Saraç, 1991): Cusps of cheek teeth subordinate to lophs. Anteroloph
of the M1 usually straight, blade like, with a shallow depression near the middle of its anterior slope
in most specimens. The true anterior arm of the protocone is usually short, but may be long if
connected with the postero-labial spur of the anterocone. The usually well developed posterior spur
of the paracone is situated lingually of the labial border in all M1 and M2 and in most M3. The
protolophule and metalophule of the M2 are parallel and directed forwards. The posterior arm of the
hypoconid is strong in almost all M1 and M2 and preserved as a spur of the posterolophid in most M3
Figure 6 - Measurements of E. carbonicus from Kaplangı 1 compared to other Eumyarion species (in mm).
Description Kaplangı 1
Table 4 - measurements of E. carbonicus from Kaplangı 1(in mm).
Length
N
M1/
M2/
M3/
M/1
M/2
M/3
6
7
3
5
6
0
Width
Range
Mean Range
Mean
1,67 - 1,96 1,83 1,07 - 1,39 1,30
1,30 - 1,51 1,39 1,27 - 1,34 1,29
1,04 - 1,12 1,09 1,10 - 1,18 1,13
1,72 - 1,89 1,79 1,09 - 1,21 1,15
1,43 - 1,49 1,46 1,16 - 1,28 1,20
1,36 - 1,40 1,38 1,06 - 1,15 1,11
M1 – The anterocone is a ridge with a straight transverse edge, it is connected to the protocone by an
anterolophule which is complete in 5(6) molars. From the labial part of the anterocone, a posteriorly
directed spur is present. The anterolophule is longitudinally oriented and ends against the lingual
side of the anterocone. The labial anteroloph is usually very short, and does not reach the paracone.
The lingual anteroloph is usually short, and never ends against the protocone. The anteromesoloph is
of medium length in 2(5) and long in 3(5) molars. Above the anteromesoloph there is a small ovalshaped plateau. The burgee-shaped posterior paracone spur is medium developed in 3(5) and
strongly developed in 2(5) molars. The protolophule is single and it is bending towards the posterior
side. The mesoloph is of medium length, it is connected to the hypocone in 3(5), and there is a small
oval shaped plateau at the labial side of the mesoloph. The metalophule bends towards the anterior
side in 4(6) and is transverse in 2(6) molars. The labial posteroloph is of medium length, and it is
connected to the metacone in 1(4) specimen.
M2 – The anteroloph is long. The protolophule is transverse in 5(7), and in 2(7) specimen it bends
towards the posterior side. The metalophule is transverse in 5(7) and it bends towards the anterior
side in 2(7) molars. In the molars with the transverse protolophule and metalophule, these ridges are
parallel, in the other molars, the ridges are bending towards each other. The mesoloph is of medium
length in 5(6) and short in 1(6) specimen. The mesosinus is very narrow. The posterior spur of the
paracone is well developed in 5(6) and medium developed in 1(6) molars. The labial posteroloph is
long in all 7 specimen, but in only 2 specimen it is connected to the metacone.
M3 – The lingual branch of the anteroloph is absent, the labial branch is long. The protocone and the
hypocone are connected by a ridge along the lingual edge of the molar. The longitudinal crest is
longitudinal. The metalophule is long. The mesoloph is long.
M1 – The lingual arm of the anterolophid is short, the labial branch connects to the base of the
protocone. All molars but one have an anterior metalophid which is directed anteriorly. One molar
also has a partially developed posterior metalophid which is leaning towards the anterior side. The
anterolophulid is incomplete in 3(4) specimen. A double mesolophid is present in all molars, both
the anterior and posterior mesolophid are of medium length in 3(5) and long in 2(5) molars. The
ectomesolophid is of medium length in 4(5) and long in 1(5) molars. In 4(5) molars, the mesolophid is
of equal length as the ectomesolophid , The mesolophid is always located more anteriorly than the
ectomesolophid. A posterior spur of the metaconid is always present. The hypoconid branch is of
medium length in 4(5) and long in 1(5) molar. The posterolophid is long.
M2 – All molars have a double mesolophid. The anterior mesolophid is long in 5(6) and of medium
length in 1(6) specimen. The posterion mesolophid is short in 5(6) molar, and in 1(6) molar it is
absent. In 3(6) molars a short ectomesolophid is present, in the other 3(6) it is absent. A posterior
spur of the metaconid is present in 4(6) specimen. In 1(6) molar, the ectomesolophid is of the same
length as the mesolophid. The hypoconid branch is of medium length in 4(6), short in 1(6) and absent
in 1(6) molar. The labial posteroloph is long and connected to the entoconid.
M3 – The labial anterolophid is of medium length in 2(3) and short in 1(3) molar. It is not connected
to the protoconid. The lingual anterolophid is of medium length in 2(3) and long in 1(3) molar, it is
connected to the metaconid. A longitudinal ridge is present at the labial edge of the molar between
the protoconid and hypoconid. The mesolophid is long in 2(3) and of medium length in 1(3) molar, in
2 specimen the mesolophid is not a straight ridge, but a bit curvy.
Discussion
The scatter plots from figure 6 show that there is no major distinction in size between the different
species and there is a lot of overlap. Therefore the measurements are less important than the
morphological features as a way to determine the species within the genus Eumyarion. Based on
these morphological features it is determined that the specimens from Kaplangı 1 are strongly
comparable to E. carbonicus from Harami 1. This is also agrees with the fact that its location only lies
about 226 kilometers away from the type locality Harami 1. Assuming the geographical distance did
not contain important barriers for migration, it seems plausible that the assemblage of Kaplangı 1
originated during, or close to, the same biozone as that of Harami 1.
Comparing E. carbonicus to other E. carbonicus from Harami 1
Even though the specimens are strongly comparable to E. carbonicus from Harami 1, there are a few
differences:
M1 – In the specimen from Harami 1, the anterocone is often somewhat bifid, but in Kaplangı 1 it is
not. The posterior paracone spur is well developed in all specimen from Harami 1, in Kaplangı 1 it is
well developed in 2(5) and medium developed in 3(5) molars.
M2 – The mesoloph in the carbonicus from Harami 1 is highly variable in length and width, but in
Kaplangı 1 it is mainly of medium length.
M3 – The lingual anteroloph is almost always present in Harami 1, while in Kaplangı 1 it is absent. In
Harami 1 the metalophule is short, but in Kaplangı it is always long.
M1 – In Harami 1, there are usually two metalophulids present, while in Kaplangı 1, the metalophulid
is single.
Comparing E. carbonicus to other Eumyarion species
When comparing E. carbonicus from Harami 1 to E. montanus from Keseköy, the paracone and
metacone are less inflated in the M1 and M2 of E. carbonicus. The posterior spur of the paracone is
better developed in the molars of the upper jaw of E. carbonicus. The lingual part of the protolophule
in the M2 is slightly more anteriorly situated in the E. carbonicus. The posterior arm of the hypoconid
is stronger in E. carbonicus.
E. carbonicus is different from E. latior from Aliveri because of the posterior paracone spur. This spur
is better developed in the upper molars of E. carbonicus. In the M1 of E. latior. The anterocone is
more bifid than in E. carbonicus. In the M2 of E. carbonicus, the protolophule is posterior directed
while in E. latior it is mainly anterior. The posterior arm of the hypoconid is stronger in the molars of
the lower jaw in E. carbonicus.
The most important difference between E. carbonicus and E. bifidus is the anterocone of the M1. In E.
bifidus it is strongly bifid, while in E. carbonicus it is undivided. In E. bifidus, there is a double
metalophule in the M2, while in E. carbonicus it is single. The posterior arm of the hypoconid is better
developed in E. carbonicus.
Evolutionary context of Eumyarion
Eumyarion is recorded in Anatolia for the first time in MN2 (Fortelius, 2012). This is E. carbonicus
from Harami 1 (de Bruijn and Saraç, 1991). In Europe the first species of Eumyarion were found in
MN4, but from MN6 on they were absent (Fortelius, 2012). It was therefore proposed that they
migrated from Asia to Europe (de Bruijn and Saraç, 1991). According to de Bruijn and Saraç (1991)
the medium sized E. carbonicus has one of the most basic dental morphologies ever seen in
Eumyarion molars, and is therefore one of the most primitive species which has been found up to
now. They proposed a dental morpholopy for the hypothetical primitive Eumyarion.
According to de Bruijn and Saraç (1991) this primitive Eumyarion must have had an anteromesoloph
in the M1 which is of medium length. This is the case in 2(5) of the Kaplangı 1 specimen. In the other
3 molars, the anteromesoloph is long. However, in the specimen from Harami 1 the variation in
length is even stronger. The posterior spur of the paracone should be long and burgee-shaped in the
M1 and M2. This is indeed the case in the specimen from Harami 1, but in Kaplangı 1 they are less well
developed. This could indicate that the specimens from Kaplangı 1 are slightly less primitive than
those from Harami. Another difference can be found in the mesoloph of the M1 and M2. In the
primitive Eumyarion the mesoloph should be short and ending freely in the central basin. This is
indeed the case in Harami 1, but in Kaplangı 1 it is mainly of medium length and often connected to
the hypocone. This could also be an indication that these specimens are younger and less primitive
than those of Harami 1.
The M1 is thought to have had a complete metalophulid which is also the case in both Kaplangı 1 and
Harami 1. Another feature, which is thought to have been present in all lower molars, is the double
mesolophid. The anterior mesolophid was probably long and free ending. In Kaplangı 1 this is the
case, but in Harami 1, it is mainly short or of medium length.
When comparing the E. carbonicus from Kaplang 1 to the hypothetical primitive Eumyarion proposed
by de Bruijn and Saraç (1991), and E. carbonicus from Harami 1, it can be concluded that there are
certainly major similarities. However, the specimens from Kaplangı 1 seem to be slightly younger and
less primitive.
Cricetodon aliveriensis
Order: Rodentia - Bowdich, 1821
Familie: Cricetidae - Fischer de Waldheim, 1817
Subfamily: Cricetodontinae - Stehlin and Schaub, 1951
Genus: Cricetodon - Lartet, 1985
Species: C. aliveriensis - Klein Hofmeijer and de Bruijn, 1988
Type locality: Aliveri (Greece), South Quarry.
Type horizon: Early Aragonian, MN4
Geographical distribution and biochronological range: Aliveri (Greece), Karydiá (Greece), Kaplangı 2
(Turkey)
Original diagnosis: C. aliveriensis is a small Cricetodon. The anterocone of the M1 consists of two
cusps of about the same size. The anterior protolophule of the M1 is incomplete, but the posterior
one is complete. The posterior part of the ectoloph of M1 and M2 are either short and low or absent.
A mesolophid is present in all lower molars. The M1 has a double metalophulid
Cricetodon is present in both Kaplangı 1 and 2. However, the Kaplangı 1 specimen are not described
in further detail because there are only 3 molars present and their condition is too poor.
Figure 7 - Measurements of C. aliveriensis from Kaplangı 2 compared to other Cricetodon species (in mm).
Description Kaplangı 2
Table 5 - measurements of C. aliveriensis from Kaplangı 2(in mm).
Length
N
M1/
M2/
M3/
M/1
M/2
M/3
2
3
2
6
3
2
Width
Range
Mean Range
Mean
2,27 - 2,33 2,30 1,51 - 1,52 1,52
1,91 - 2,05 1,98 1,56 - 1,65 1,61
1,74 - 1,77 1,76 1,46 - 1,57 1,52
1,77 -2,35 2,08 1,29 - 1,43 1,35
1,80 - 2,02 1,91 1,50 - 1,55 1,52
1,96 - 2,09 2,03 1,54 - 1,59 1,57
M1 – The M1 has a very prominent anterocone which is strongly bifid. In one specimen it is located at
the far most labial side of the molar, and in the other it is located in the middle. The labial anteroloph
is long in 1(2) and of medium length in 1(2) molar, it is connected to the paracone. The lingual
anteroloph is long and connected to the protocone in 1(2), and of medium length and not connected
in the other. The anterolophule is only visible in 1(2), it is a longitudinal ridge and ends against the
middle part of the anterocone. Both teeth have an anteromesoloph, one long and one of medium
length. The protosinus is broad and deep in 1(2), and narrow and shallow in the other 1(2). A double
protolophule is present. The anterior protolophule is bending towards the posterior side, while the
posterior protolophule is bending towards the anterior side. The posterior protolophule is less
pronounced than the anterior one. The mesoloph is single and short. The mesosinus is open. The
sinus is open and narrow, and it is slightly bended towards the anterior side. The entoloph is bended.
The metalophule is absent, but the metacone is connected to the mesoloph and the posteroloph.
The labial posteroloph is medium or short, and it is always connected to the metacone. Because the
labial posteroloph is almost merged to the metacone, there is no posterosinus present.
M2 – The labial anteroloph is long and connected to the paracone. The lingual anteroloph is long in
1(3), medium in 1(3) and short in 1(3), only the long anteroloph is connected to the protocone. The
labial anteroloph is always stronger and more pronounced than the lingual one. The anterolophule is
oblique in 2(3) molars, and longitudinal in 1(3). In 1(3) molar, an anteromesoloph is present. The
protolophule is single and it is located at the posterior side of the protocone and is bending towards
the anterior side of the molar. The mesoloph is short, and it is connected to the metacone in 1(3)
molar. In 1(3) specimen a mesostyl is present, and there is a ridge between the paracone and
metacone in 1(3) molar. The mesosinus is open in 2(3) specimen. In 1(3) molar, a short
entomesoloph is present. The sinus is narrow in 2(3) molars and it is bending strongly towards the
anterior side in 3(3). The metalophule is bended towards the anterior side. The posterosinus is
narrow. The labial posteroloph is short in 2(3) molars and of medium length in 1(3), in 2(3) molars it
is connected to the metacone. The lingual posteroloph is absent.
M3 – The labial anteroloph is long and connected to the paracone. The lingual anteroloph is of
medium length. The molars have four cones with a strongly separated metacone and hypocone. At
the anterior side of the molar there is a cone-like structure because of the connection between the
anterolophule and the protolophule. The protosinus is open . The protolophule is bending towards
the anterior side in 1(2) molar, and it is transverse in the other 1(2). In both specimen, there is a ridge
at the labial edge connecting the paracone and the metacone, because of this, the mesosinus is
closed. In 1(2) specimen, there is an incomplete longitudinal crest, in the other 1(2) it is absent. There
is a posteriorly directed metalophule, 1(2) is of medium length and the other 1(2) long. Both molars
have a long mesoloph.
M1 - The first molars of the lower jaw are more elongated than those of the upper jaw. The
anteroconid is located in the middle of the anterior edge, and has a rounded pointy anterior edge,
which gives it a triangular appearance. The labial anterolophid is long and connected to the
protoconid. The lingual anterolophid is of medium length, and connected to the metaconid. The
anterolophulid is longitudinal in 3(4) molars and oblique in 1(4). The protosinusoid is broad and
closed. The anterosinusid is broad in 3(4) specimen and it is closed in all 4(4). The metalophulid is
double, the anterior metalophulid is ending at the anterior side of the metaconid and it is bending
towards the posterior side. The posterior metalophulid is bending towards the anterior side in 4(5),
and in 1(5) it is transversal. In 2(5) specimen there is a weak posterior spur of the metaconid. In 4(5)
specimen a short mesolophid is present. There is no ridge at the lingual edge between the
metaconid and the entoconid, so the mesosinusid is open. The entomesolophid is long in 1(5),
medium in 2(5), short in 1(5) and absent in 1(5) molar. The entolophid is bended in 3(5) and is
transversal in 2(5) molars. There is a ridge at the labial edge between the protoconid and the
hypoconid. The broad sinusoid is slightly bended towards the anterior side. The hypolophulid is
ending at the anterior side of the etoconid and is bending towards the posterior side. In 2(5) molars
there is a short hypoconid branch, in the other 3(5) it is absent. The posterosinusid is narrow in 3(5)
and broad in 2(5) specimen. The labial posterolophid is absent. The lingual posterolophid is long and
connected to the entoconid in all specimen.
M2 – The labial anterolophid is long and connected to the protoconid. In 1(3) molar a short lingual
anterolophid is present, in the other 2(3) it is absent. The anterophulid is oblique. The protosinusid is
broad and closed. The anterosinusid is absent. The metalophulid is complete in 2(3) molars, and it is
always bended towards the anterior side. There is no mesolophid, and the mesosinusid is open. The
ectomesolophid is short in 2(3) and of medium size in 1(3) molar. The ectolophid is bended. The
broad sinusoid is not bended. The hypolophulid is ending at the anterior side of the entoconid, and is
bending towards the posterior side of the molar. 1(3) molar has a small hypoconid branch. The
posterosinusid is broad. In all molars the lingual posterolophid is long, and it is connected to the
entoconid in 2(3) molars.
M3 - Only 1(2) molar was suitable for determination. Its labial anteroloph is of medium size and the
lingual anteroloph is absent. The molar has 4 cones with a clearly separated entaconid and
hypoconid. At the anterior side of the molar there is a cone-like structure because of the connection
between the anterolophulid and the metalophulid. The protosinusoid is open. The metalophulid is
bending towards the posterior side and the labial side is ending against the anteroconid. The
mesolophid is of medium length and slightly bended towards the anterior side of the molar. The
mesosinusoid is open. There is no longitudinal crest. The hypolophulid is short. The hypoconid is
large and merged with the broad posterolophid. The posterolophid is not connected to the
entoconid.
Discussion
When comparing the specimen from Kaplangı 2 with other Cricetodon species, they resembled C.
aliveriensis from Aliveri (Greece) and C. tobieni (de Bruijn et al, 1993) from Horlak 1a (Turkey) the
most. When comparing these two species with the molars from Kaplangı 2 in greater detail it is
concluded that they bear the most resemblance to C. aliveriensis. Although the molars from Kaplangı
1 where not suitable for closer determination, they do bear a strong resemblance to those from
Kaplangı 2 and are therefore classified as C. cf. aliveriensis.
Comparing the C. Aliveriensis from Kaplangı 2 to C. aliveriensis from Aliveri
Even though C. aliveriensis from Aliveri and C. aliveriensis from Kaplangı 2 are much alike, there are
still some small differences between both groups.
M1 – The anteromesoloph in the Aliveri specimen is short. In the molars from Kaplangı 2 the
anteromesoloph is either long or of medium length, and it is connected to a posterior spur of the
anterocone which is not present in the Aliveri specimen. The anterocone is oriented slightly more
lingually in the Kaplangı 2 specimen than it is in the molars from Aliveri. In the Kaplangı 2 specimen a
double protolophule is present, the molars from Aliveri only have a single posteriorly protolophule.
In the molars from Aliveri, the labial anteroloph is more pronounced than in the Kaplangı 2 specimen.
It is also connected to the paracone in the Aliveri specimen, but this is not the case in the molars
from Kaplangı 2.
M2 – In the molars from Aliveri, the labial anteroloph is not connected to the paracone, in Kaplangı 2,
they are connected in 2(3) molars.
M2 – The mesolophid is better developed in the molars from Kaplangı 2 than in the molars from
Aliveri. In Kaplangı 2 the posterolophid is like an extension of the hypoconid, while in the molars
from Aliveri it is more like a separate ridge.
Comparing the C. Aliveriensis from Kaplangı 2 to C. tobieni from Horlak 1a
When comparing the C. Aliveriensis from Kaplangı 2 to C. tobieni from Horlak 1a there are also some
differences.
M1 – The molars of C. aliveriensis are more narrow than those of C. tobieni. In the molars of C.
aliveriensis the anteromesoloph is either long or of medium length, while in the molars of C. tobieni
the anteromesoloph is absent. In C. tobieni, the lingual anteroloph is less well developed than it is in
C. aliveriensis. In both species the mesoloph is of medium length, but in C. tobieni it is transverse and
in C. aliveriensis it is bending towards the posterior and connecting to the metacone. In C. tobieni the
posteroloph is better developed. In the molars of C. aliveriensis there is a ridge between the
protocone and the entocone which is absent in C. tobieni.
M2 – The molars of C. tobieni have a better developed lingual anteroloph than those of C. aliveriensis.
In C. aliveriensis there is a ridge between the paracone and the metacone. This ridge is absent in C.
tobieni.
M3 – The molars of C. tobieni are round while the molars of C. aliveriensis have an elongated shape.
In the molars of C. tobieni a double protolophule is present. In C. aliveriensis there is a single
posterior bending protolophule.
M1 – In C. tobieni there is a single posteriorly oriented metalophulid, while in C. aliveriensis there is a
double metalophulid. The anterior metalophulid is connected to the anteroconid. The lingual
anterolophid is less well developed in C. tobieni than it is in C. aliveriensis. In C. aliveriensis the
ectomesolophid is better developed than it is in C. tobieni. The molars from C. aliveriensis have a
ridge between the protoconid and the hypoconid, this ridge is absent in C. tobieni. The lingual
posteroloph is better developed and broader in C. aliveriensis.
M2 - In the molars of C. aliveriensis, the ectomesolophid lies in the extension of the hypolophulid. In
C. tobieni, the ectomesolophid is not always present and if it is, it is located more anteriorly. In C.
aliveriensis, the mesolophid is better developed than in C. tobieni which only has a short mesolophid.
At the anterior edge of the molars of C. tobieni, there is a cone-like structure from where the
anterolophid and the metalophulid are protruding. This is not present in C. aliveriensis.
M3 – The lingual anterolophid is better developed in C. tobieni than it is in C. aliveriensis. In C. tobieni,
the posteriorly oriented mesolophid is short or of medium length. In C. aliveriensis the mesolophid is
anteriorly oriented and better developed. In C. tobieni, some of the molars have an ectomesolophid,
in C. aliveriensis this ectomesolophid is absent.
Evolutionary context of Cricetodon
Cricetodon aliveriensis from Aliveri (MN4)is one the oldest Cricetodon species found thus far(Klein
Hofmeijer and de Bruijn, 1988). It is thought to be the earliest immigrant from Anatolia, but does not
seem to be an ancestor of the younger Cricetodon species of Central or Western Europe. The
ancestor of the Central and Western European species also came from Anatolia and shares a
common ancestor with C. aliveriensis. However these lineages had already split during the Middle or
Late Oligocene (Rummel, 1999).
The presence of C. aliveriensis in Kaplangı supports the theory that it lived in Anatolia, and it is
plausible that it had migrated from there towards Greece. Its close resemblance to C. tobieni from
Horlak 1a indicates that they could be closely related and that C. tobieni might have been an ancestor
of C. aliveriensis (de Bruijn et al, 1993). This would also support the theory of the Cricetodon
migration from east to west.
These results could indicate that these specimens are older or of the same age as those from Aliveri
and Karydiá (MN4), an of the same age or younger than those of Horlak 1a (MN3-4).
Democricetodon gracilis
Order: Rodentia - Bowdich, 1821
Familie: Cricetidae - Fischer de Waldheim, 1817
Genus: Democricetodon - Fahlbusch, 1964
Species: D. gracilis - Fahlbusch, 1964
Holotype: M1 dextral (BSPG 1959 II 247), BSPG Munich, Germany
Type locality: Sandelzhausen, Southern Germany, Bavaria.
Type horizon: Early/Middle Miocene boundary, MN5
Geographical distribution and biochronological range: Greece (MN4), Germany (MN4-5), Switzerland
(MN4-5), Turkey (MN4-5), Austria (MN4-5), Czech Republic (MN5) and Poland (MN6).
Original diagnosis: This Democricetodon is related to D. minor from Sansan, but it is smaller and has a
more graceful appearance. Lower molars: The sides of the crowns are somewhat concave near the
mesosinusid and sinusoid in most molars. The anteroconid is mainly very short and forms a flat
triangle. The mesolophid is very short to half long. The metaconid is curved towards the anterior side
in most molars. Upper molars: The labial side of the crown is mostly undulatory. The anterocone is
relatively narrow and undivided. The mesoloph is long and often reaches the labial edge of the
molar. (translated from German, Fahlbusch 1964)
Figure 8 - Measurements of D. gracilis from Kaplangı 1 and 2 compared to other Democricetodon species (in mm).
Description Kaplangı 1
Table 6 - measurements of D. gracilis from Kaplangı 1(in mm).
Length
N
M1/
M2/
M3/
M/1
M/2
M/3
8
4
2
9
9
6
Width
Range
Mean Range
Mean
1,48 - 1,78 1,61 0,99 - 1,09 1,05
1,15 - 1,26 1,20 0,98 - 1,07 1,03
0,81 - 0,82 0,82
0,86 0,86
1,24 - 1,38 1,31 0,82 - 0,97 0,92
1,16 - 1,30 1,22 0,97 - 1,03 1,00
1,00 - 1,04 1,02 0,80 - 0,91 0,85
M1 – The anterocone is a crest-like cone which is usually more voluptuous at the labial side (6/8
molars). It is usually located at the far labial side of the tooth (5/8 molars), at the middle to labial side
(2/8) and in the middle (1/8). There are always both a labial and a lingual anteroloph present. The
labial anteroloph is usually of medium length in 6, long in 1(8) and short in 1 (8), and is only
connected to the paracone in 2 (8). The lingual anteroloph is usually short (6/8) and in 2 of medium
length, and does not connect to the protocone. The anterolophule is complete in all M1 and is
longitudinal in 4 (8) and oblique in the other 4. The anterolophule ends against the lingual side of the
anteroloph in 6, and in 2 to the middle of the anterocone. The protosinus is mainly narrow (5/8), but
it is always open. 2 out of 7 molars developed a posterior paracone spur, and 4 out of 8 have an
anterior paracone bulge. In all molars, a posterior protolophule is present which always bends
towards the posterior side of the molar. In 4(8) molars, an anterior protolophule is also present. In 1
molar it is incomplete; these anterior protolophules are always bended towards the anterior side of
the molar. The mesoloph is of medium length 5(8) or long 3(8), and is connected to the posterior
paracone spur in 1 molar. In 2(7) molars, a mesostyl is present. In 1 out of 7 a longitudinal ridge along
the labial ridge of the molar connects the paracone and the metacone, while in 3(8) a longitudinal
ridge along the lingual edge of the molar connects the protocone and the hypocone. In 5 out of 6,
the mesosinus is open, while the sinus is open in 6 out of 8 molars. The sinus is narrow in all molars
and usually not bended (5/8), but in 3 it bends lightly towards the posterior side of the molar. In 4(8)
molars the entoloph is U-shaped, while it is straight in the other 4. The metalophule is present in 6(7)
molars, and is always bended towards the posterior edge. All molars have an anterior hypocone spur
at the lingual side of the tooth. The posterosinus is very narrow. In 5(6), a medium sized labial
posteroloph is present, which is connected to the metacone at the postero-labial edge in 3. In 1 out
of 6, the posteroloph is absent.
M2 – The labial anteroloph is present in all M2, but in only 1(4) it is connected to the paracone. In 3(4)
molars, a lingual anteroloph is present (2 long and 1 short), and is connected to the protocone in 1
specimen. The lingual anteroloph is weaker than the labial one. The anterolophule is complete in all
specimen, and is longitudinal in 3(4), and oblique in 1(4). Only 1 out 4 molars has a (weak) posterior
paracone spur. In all molars, an anterior protolophule is present, which bends towards the anterior
edge of the molar. In 1(4) molar, an anterior longitudinal spur is present at the anterior side of the
protolophule. A complete posterior protolophule is present in 3(4) molars, while in 1 specimen it is
incomplete. These protolophules are always bending towards the posterior edge. The mesoloph is
long, and in 2 out of 4 molars connected to the posterior side of the paracone. In 2(4) specimens a
mesostyl is present, and in 1(4) a ridge is present between the paracone and metacone. The
mesosinus is open in 3(4) specimens. An anterior spur of the hypocone is present(at the far lingual
edge), and in 1(4) molar there is a ridge between the protocone and the hypocone at the lingual edge
of the molar. The sinus is straight and broad in 3(4) molars. The metalophule bends towards the
anterior edge in 2(4), and is straight in 2(4). The posterosinus is narrow in 2(4) and broad in 2(4)
molars. The labial posteroloph is long and connected to the metacone at the postero-labial edge.
M3 – The labial anteroloph is long and connected to the paracone. The lingual anteroloph is short.
The molars have four regular cones and a very small cone at the anterior edge of the molar. The
metacone and hypocone are very close and are almost merged. The protosinus is open. The
protolophule ends against the proterocone. One protolophule is bended towards the anterior side,
the other one is straight. The posterior paracone spur is medium developed. A ridge at the labial side
of the molar between the paracone and metacone causes the mesosinus to be closed. The
metalophule is long in 1(2), and incomplete and of medium length in the other. The hypocone is to
the posterior arm of the protocone. A posterosinus is present in 1 out of 2 molars.
M1 - The anteroconid is a single crest-like cone which is rounded at the anterior side. In 4(8)
specimens the lingual part of the anteroconid is slightly more swollen. In 6 molars, the anteroconid is
located at the lingual-middle part of the tooth. In the other 2 it is located at the middle part. The
lingual anterolophid is long in 4(8), medium in 3(8) or short in 1(8), and is connected to the
protoconid in 2(8). The labial anterolophid is of medium length in 2(8), short in 5(8) and absent 1(8).
It is connected to the metaconid in 1(8) specimen. The anterolophulid is complete, and is longitudinal
in 4(8) or oblique in 4(8). The protosinusid is broad in 6(8), and narrow in 2(8) teeth. It is open at the
labial side in 5(8) and closed in 3(8) molars. The anterosinusid is narrow and open. The metalophulid
is located at the anterior side of the metaconid and it is bending towards the posterior side. Only 1(8)
molar has a weakly developed posterior spur of the metaconid, and 1(8) molar has a medium
developed anterior spur of the metaconid. The mesolophid is long in 3(8), medium in 4(8) and short
in 1(8) molar. In 1 specimen, a small plateau is present at the lingual side of the mesolophid. In 3(8)
molars, a longitudinal ridge is present at the lingual edge between the metaconid and the entoconid.
The mesosinusid is open in 6(8), while the sinusid is open in 3(8) molars. An ectostylid is present in
3(8) specimen. The entolophid is U-shaped in 5(8) and straight transversal 3(8) specimen. In 5(8)
molars a longitudinal ridge at the labial edge between the protoconid and the hypoconid is present.
The sinusid is slightly bended towards the anterior side in 7(8), or not bended at all in 1(8) molar. It is
broad in 5(8), and narrow in 3(8) molars. The hypolophulid bends towards the posterior side. In one
specimen, the hypoconid has an anterior spur at the labial edge of the molar. The posterosinusid is
broad in 4(8) and narrow in the other 4(8) molars. The labial posterolophid is short in 4(8) and absent
in the other 4(8) specimen. The lingual posterolophid is long in 7(8) and of medium length in 1(8)
molar. In 6(8)molars it is connected to the entoconid.
M2 – The labial anterolophid is long in 6(9), of medium length in 2(9) and absent in 1(9) molar. It is
connected to the protoconid in 1(9) molar. The lingual anterolophid is short in 6(7) and of medium
length in 1(7) molar, and it is connected to the metaconid. The anterolophulid is complete and it is
longitudinal in 5(7) and oblique in 2(7) specimens. The protosinusid is narrow in 7(9), and open in
7(8) molars. The anterosinusid is narrow and open. The metalophulid bends towards the anterior
edge of the molar. In 2(9)molars, a posterior spur of the metaconid is present, 1 medium and 1
weakly developed. The mesolophid is long in 5(8), medium in 2(8) and short in 1(8) specimen, it is not
connected to the metaconid. The mesosinusid is open. In 2(9) molars an ectostylid is present. The
ectolophid is U-shaped in 8(9), and straight and longitudinally oriented in 1(9) molar. The sinusid is
slightly bended towards the anterior side in 7(9), or not bended at all in 2(9) specimen, it is narrow in
6(9). The hypolophulid is located at the anterior side of the entoconid, and it bends towards the
anterior side. The posterosinusid is broad in 6(8) molars. The lingual posterolophid is long, and it is
connected to the entoconid in 4(7) molars. A labial posterolophid is absent.
M3 – The third molars of the lower jaw are, compared to those of the upper jaw, very elongated. The
labial anterolophid is long in 4(6), medium in 1(6) and short in 1(6) molars, and it is connected to the
metaconid in 2(6) specimen. The lingual anterolophid is of medium length in 3(4), and long in 1(4)
molar, and it is connected to the protoconid in 3(4) specimens. The molars have four main cones and
one cone-like feature at the anterior edge of the molar, this is mainly because the anterolophule and
metalophulid are coming together and form a small bulge. The entoconid and hypoconid are fused
together. The protosinusid is open in 4(6) molars. The metalophulid is complete in 5(5), it bends
towards the anterior side in 4(5), and in 1(5) molar it is straight. The metalophulid ends against the
anterior conid in 3(5) molars. There is a ridge at the lingual side between the metaconid and the
entoconid. A long anterior mesolophid is present in 1(6) molar, while in the other 5 it is absent. The
posterior mesolophid is long and connected to the lingual ridge between the entoconid and the
metaconid. The mesosinusid is closed. The hypolophulid is absent. The entaconid is fused to the
posterolophid. There is no posterosinusid.
Description Kaplangı 2
Table 7 - measurements of D. gracilis from Kaplangı 2(in mm).
Length
N
M1/
M2/
M3/
M/1
M/2
M/3
2
5
4
6
3
0
Width
Range
Mean Range
Mean
0,90 - 0,93 0,92 0,91 - 0,92 0,92
1,07 - 1,23 1,17 1,01 - 1,08 1,06
1,38 - 1,68 1,55 0,96 - 1,06 1,02
1,23 - 1,57 1,35 0,83 - 0,94 0,89
1,14 - 1,24 1,21 1,00 - 1,06 1,03
-
M1 – The anterocone is located at the labial-middle part of the anterior edge of the molar. It is crestlike in 2(3) molars. The labial anteroloph is of medium length in 2(3) and short in 1(3) molar. The
lingual anteroloph is of medium length in 2(3) and long in 1(3) molar. The anterolophule is
longitudinal in 2(4) and oblique in 2(4) molars. The anterolophule is ending against the middle part of
the anterocone in 3(4) and against the lingual part of the anterocone in 1(4) molar. The protosinus is
broad in 1(4) and narrow in 3(4) specimen. It is shallow in 3(4) and deep in 1(4) molar. The posterior
or anterior spur is absent. In 1(4) molar, a double protolophule is present. In the other 3(4) molars a
single protolophule is present which is bending towards the posterior side. The mesoloph is long in
2(4) and of medium length in the other 2(4) molars. The mesosinus is open. The entoloph has an
upside-down U-shape in 3(4) and is longitudinal in 1(4) molar. A longitudinal ridge at the lingual edge
between the protocone and the hypocone is present in 1(4), in the other 3 molars the sinus is open.
The sinus is slightly bended towards the posterior side in 2(4), and not bended at all in the other 2(4).
It is broad in 2(4) and narrow in the other 2(4). In 2(3) molars a metalophule is present, which is
bended towards the posterior side. In 1(3) molar an anterior spur of the hypocone is present. The
posterosinus is narrow. The labial posteroloph is of medium length and the lingual posteroloph is
absent.
M2 – The labial anteroloph is long and connected to the paracone. The lingual anteroloph is long in
3(5), medium in 1(5) and short in 1(5) molar. It is connected to the protocone in 3(5) molars. The
lingual anteroloph is always shorter than the labial anteroloph. The anterolophule is longitudinal. In
3(5) molars, a weakly developed posterior spur of the paracone is present. The protolophule is
always double. These protolophules are bending towards each other and are ending against the
paracone. The mesoloph is long, and it is connected to the paracone spur in 1(5) molar. A mesostyl is
present in 2(5) molars. In 4(5) molars a longitudinal ridge is present at the labial edge between the
paracone and the metacone. Therefore the mesosinus is closed in 4(5) molars. In 3(5) molars a
longitudinal ridge is present at the lingual edge between the protocone and hypocone. Therefore the
sinus is closed in 3(5). In all molars the sinus is broad and straight. The metalophule bends towards
the posterior side of the molar in 2(5) specimen and in the other 3(5) it is straight. The posterosinus
is narrow in 3(5) and broad in 2(5) molars. The labial posteroloph is long in 4(5) and of medium
length in 1(5) molar, in 1 specimen the labial posteroloph is connected to the metacone.
M3 – The labial anteroloph is long, and it is connected to the paracone in 1 molar. The lingual
anteroloph is of medium length in 1, and short in the other, and it is not connected to the protocone.
The protosinus is open. The protolophule is complete and. Both molars have a medium developed
paracone spur. A longitudinal ridge is present at the labial edge between the paracone and
metacone, therefore the mesosinus is closed. In 1(2) molar a longitudinal crest is present. The
metalophule is complete and short in 1(2), and of medium length but incomplete in the other 1(2).
The hypocone is fused to the posterior arm of the protocone. The metacone is fused to the
posteroloph. The mesoloph is of medium length.
M1 - The anteroconid is undivided and rounded at the anterior side. It is located at the middle part of
the anterior edge in 3(4) and in the lingual-middle part in 1(4) molar. The labial anterolophid is of
medium length. The lingual anterolophid is short in 2(3) and of medium length in 1(3) molar. The
Anterolophulid is horizontal. The protosinusid and anterosinusid are broad and open. The
metalophulid is bending towards the posterior side. The posterior spur of the metaconid is medium
developed in 2(3) and weakly developed in 1(3) molar. The mesolophid is of medium length. In 1(3)
molar a mesostylid is present. The mesosinusid is open. In 1(3) molar, an ectostylid is present. The
ectolophid has an upside-down U-shape in 3(4) and it is straight in 1(4) molar. The sinusoid is open. It
is bended towards the anterior side and broad in 3 and narrow in 1 molar. The hypolophulid is
located at the anterior side of the hypoconid and entoconid and is bending towards the posterior
side. The posterosinusid is narrow in 3(4) and broad in 1(4) molar. The lingual posterolophid is long
and connected to the entoconid. The labial posterolophid is absent.
M2 - The labial anterolophid is of medium length and not connected to the protoconid. The lingual
anterolophid is short and connected to the metaconid. The anterolophulid is oblique in 2(3) and
longitudinal in 1(3) molar. The protosinusid is open and broad. The anterosinusid closed and narrow.
The metalophulid is located at the anterior side of the protoconid and metaconid, and is bending
towards the posterior side. In 2(3) molars, a slightly developed posterior spur of the metaconid is
present. The mesolophid is of medium length in 2(3) and long in 1(3) molar. The mesosinusid is open.
The ectolophid has an upside-down U-shape. The sinusoid is slightly bended towards the anterior
side in 2(3) and straight in 1(3) molar. It is broad in 2(3), and narrow in 1(3) molar. The hypolophulid
bends towards the posterior side. The posterosinusid is broad. The lingual posteroloph is long in 2(3)
and of medium length in 1(3) molar. The labial posterolophid is absent.
Discussion
The size of the molars of the different Democricetodon species shown in figure 8 is very similar.
Therefore, when determining the species of Democricetodon present in Kaplangı 1 and 2, the
emphasis lies on dental morphology. Because there are also many similarities in the dental
morphologies of the different Democricetodon species, the main focus lies on 5 important features in
the M1. This is because the M1 shows most variation between the different species. The studied
features are: the position and the shape of the anterocone, the distance between the anterocone
and the protocone and paracone, the length of the mesoloph, and whether the anterocone lies on a
distinct area or if it is incorporated into the rest of the molar. Based on these features, the decision
was made that de Democricetodon molars from both localities belong to the species D. gracilis.
Comparing D. gracilis from Kaplangı to D. gracilis from Sandelzhausen
When comparing the D. gracilis from Kaplangı 1 and 2 to the type specimen from Sandelzhausen
(Germany) (Freudenthal, 1964), there are a few differences:
M1 – In the molars from Sandelzhausen, the anterosinus is closed, but in Kaplangı 1 this is only the
case in 2(8) specimen. In Kaplangı 2 it is always open. The mesoloph is mainly long in the specimen
from Sandelzhausen, but in Kaplangı 1 it is long in 3(8), medium in 4(8) and short in 1(8) molar. In
Kaplangı 2 the mesoloph is long in 2(4) and of medium length in 2(4) molars. The sinus is closed by a
ridge at the lingual side of the molar in the German specimen, in Kaplangı 1 this is only the case in
3(8) specimen. In Kaplangı 2 a ridge is present in 1(4) molar.
M2 – Both Kaplangı 1 and 2 have a double protolophule while the German specimen there is a single
transverse protolophule. In the molars from Sandelzhausen, the sinus is always closed by a lingual
ridge between the protocone and the hypocone, in Kaplangı 1 this is the case in 1(4) molar and in
Kaplangı 2 in 3(5) molars.
M3 – In the specimen from Sandelzhausen, the sinus is absent, in Kaplangı 1 it is present in 1(2) molar
and in Kaplangı 2 in 2(2) molars.
M1 – The lingual anterolophid is long and connected to the metaconid in the molars from
Sandelzhausen. In Kaplangı 1, the lingual anterolophid is of medium length in 2(8), short in 5(8) and
absent in 1(8) molar. It is connected to the metaconid in 1(8) molar. In Kaplangı 2, the lingual
anterolophid is of medium length in 1(3) and short in 2(3) molars. It is never connected to the
metaconid. In Sandelzhausen, the labial anterolophid is long. In Kaplangı 1 it is long in 4(8), of
medium length in 3(8) and short in 1(8) molar. In Kaplangı 2, it is of medium length. In the German
molars, the metalophid and hypolophid are parallel and bending towards the anterior side. In the
specimens from Kaplangı 1 and 2 they are also parallel, but bending towards the posterior side. In
the molars of Kaplangı 1 and 2 there is always a complete metalophulid, while in Sandelzhausen it is
sometimes absent. The ectolophid is always bended in the molars from Sandelzhausen. In Kaplangı 1
this is only the case in 5(8) molars, and in Kaplangı 2 in 1(4) molar. In the specimen from
Sandelzhausen the sinusoid is mainly transversal and sometimes slightly bended. In both Kaplangı 1
and 2, it is mainly bended towards the anterior side of the molar.
M2 – In the molars from Sandelzhausen, the labial anterolophid is long and connected to the
protoconid. In Kaplangı 1 and 2 it is of medium length but it only reaches the protoconid in 1(8)
molar from Kaplangı 1, in Kaplangı 2 it never reaches the protoconid. In the specimen from Germany
the mesolophid is short and connects with the posterior side of the metaconid. In Kaplangı 1, the
mesolophid is long in 5(8), of medium length in 2(8) and short in 1(8). It is not connected to the
metaconid. In Kaplangı 2 the mesolophid is of medium length in 2(3) and long in 1(3) molar, it is also
not connected to the metaconid. The posterosinusid is closed in the specimen from Sandelzhausen,
in Kaplangı 1 it is closed in 4(7) molars, and in Kaplangı 2 in 2(3) molars.
M3 - The lingual anterolophid is long in the German specimen, in Kaplangı 1 it is of medium length in
3(4) and long in 1(4) molar.
D. gracilis from Kaplangı 1 and 2 differs quite a lot from the type specimen of D. gracilis from
Sandelzhausen, but they are more similar to the D. gracilis specimen found nearer to Kaplangı, like
Karydiá 1 and 2 and Aliveri. However, there are still some differences.
Comparison of D. gracilis of Kaplangı to those of Aliveri
M1 – In D. gracilis from Aliveri, the anterolophule is ending at the lingual side of the anterocone in
only 3(42) specimen. In the molars from Kaplangı 1 this is the case in 6(8) molars, in Kaplangı 2 in 1(4)
molars.
M3 – in the molars from Aliveri, the posteroloph is fused to the metacone, while in the molars from
Kaplangı 1 they are not.
M2 - In the Aliveri specimen the mesolophid is of medium length in 24(41), short in 9(41) long in 5(41)
and absent in 4(41) molars. In Kaplangı 1 it is long in 5(8), of medium length in 2(8) and short in 1(8)
molar. In Kaplangı 2 it is of medium length in 2(3) and long in 1(3) molar.
M3 – The molars from Aliveri only have a (short) mesolophid in 2(13) molars, in the Kaplangı 1 and 2
molars, all molars have a long mesolophid.
Comparison of D. gracilis of Kaplangı to those of Karydiá 1
M3 – In Karydiá 1, there is a double protolophule in 3(4) molars, while in Kaplangı 1 and 2 the
protolophule is always single.
M1 – In the specimens from Karydiá 1 the anterosinusid is closed in 3(4), while in the Kaplangı 1
specimen this is the case in 1(8) molar. In Kaplangı 2 it is always open. In Karydiá 1 the mesolophid is
short in 4(6) of medium length in 1(6) and long in 1(6) molar, while in Kaplangı 1 it is of medium
length in 4(8), long in 3(8) and short in 1(8) molar. In Kaplangı 2 the mesolophid is always of medium
length.
M2 - In the specimen from Karydiá 1 the mesolophid is short in 3(7), long in 2(7), of medium length in
1(7) and absent in 1(7) molar. In Kaplangı 1 it is long in 5(8), of medium length in 2(8) and short in
1(8) molar. In Kaplangı 2 the mesolophid is of medium length in 2(3) and long in 1(3) molar.
M3 – The M3 from Karydiá 1 has a very long labial anterolophid and the lingual anterolophid is
absent. However, this is only based on one molar. In Kaplangı 1 the labial anterolophid is long in 4(6),
of medium length in 1(6) and short in 1(6) molar. The lingual anterolophid is of medium length in
3(4) and long in 1(4) molar.
Comparison of D. gracilis of Kaplangı to those of Karydiá 2
M3 – The molars from Karydiá 2 often have a double protolophule while in Kaplangı 1 and 2 it is
always single. In Karydiá 2 the metacone is incorporated into the posteroloph, this is also the case in
Kaplangı 2, but in Kaplangı 1 it is not.
M1 - The anterosinus is closed in 7(11) molars from Karydiá 2, but Kaplangı 1 and 2 it is always open.
In the molars from Karydiá 2, the mesolophid is short in 6(12), long in 3(12), absent in 2(12) and of
medium length in 1(12) molar, while in Kaplangı 1 it is of medium length in 4(8), long in 3(8) and
short in 1(8) molar. In Kaplangı 2 the mesolophid is always of medium length
M2 - In Karydiá 2, the labial anteroloph always ends against the protoconid, while in Kaplangı 1 this
only happens in 1(8) molar. The mesolophid of the molars of Karydiá 2 is short in 5(6) and absent in
1(6) molar. In Kaplangı 1 it is long in 5(8), of medium length in 2(8) and short in 1(8) molar. In
Kaplangı 2 the mesolophid is of medium length in 2(3) and long in 1(3) molar.
M3 – The molars from Karydiá 2 have a long labial anterolophid which ends against the protoconid
and the lingual anterolophid is absent or short. In Kaplangı 1 the labial anterolophid is long in 4(6),
medium in 1(6) and short in 1(6). The lingual anteroloph is long in 1(4) and of medium length in 3(4)
molars.
Comparing D.gracilis to D. doukasi (Keseköy)
When comparing D. gracilis from Kaplangı 1 with D. doukasi from Keseköy (Theocharopoulos, 2000),
D. gracilis is on average larger than D. doukasi and has a more elegant shape. They can mainly be
distinguished because of the posterior paracone-spur which is most of the time present in the D.
Doukasi and only sometimes in the D. gracilis. Another difference is the way the anterocone is
positioned: in D. doukasi, the anterocone is located on a distinct part of the molar, while in D. gracilis
it is more incorporated with the rest of the molar.
Based on the differences between the species Theocharopoulos (2000) suggested that D. doukasi is a
more primitive species than D. gracilis.
Comparing D. gracilis to D. franconicus (Aliveri)
In the M1 of D. franconicus the labial part of the anteroloph always reaches the paracone, but in D.
gracilis it only reaches the paracone in 2 out of 8 molars. The anterocone is often bifid in D.
franconicus while in D. gracilis it is a continuous crestlike cone. The distance between the anterocone
and the protocone/paracone is also smaller in D. franconicus. In D. franconicus, the anterocone is a
distinct part of the molar, while in D. gracilis it is more incorporated with the rest of the molar. The
lingual anteroloph is strong in D. franconicus, but weak in D. gracilis. The mesoloph of the M1 is
mainly long in D. franconicus, but of medium length in D. gracilis.
In the M2 and M3 of D. fahlbush, the lingual anterolophid is well developed. This is not the case in
D.gracilis.
In the M1 of D. franconicus the anteroconid is located close to the protoconid and metaconid, while
in D. gracilis the distance between these cones is twice as big. The anterosinusid in the M1 is
sometimes lingually closed in D. franconicus, but it is always open in D. gracilis. In the M1 of D.
franconicus the mesolophid is long to very long while in D. gracilis it is medium to long. In D.
franconicus 23% of the M1 have an ectomesolophid, while in D. gracilis it is 50%.
The lingual anterolophid of the M2 of D. franconicus is very variable in length while it is mainly short
in D. gracilis.
In the M3 of the D. franconicus only 28% of the molars have a mesolophid, while it is always present
in the molars of D. gracilis.
Evolutionary context of Democricetodon
The Cricetidae seem to have originated in Central Asia during the Eocene (Wang and Dawson, 1994),
and have undergone a major radiation in Western Asia during the Early Oligocene (Ünay, 1989).
During the Oligocene, they migrated towards Anatolia. The oldest records of Democricetodontinae
are from Inkonak and Kargi, both from Central Anatolia. From there they migrated over large parts of
the Northern hemisphere during the Miocene (Theocharopoulos, 2000).
According to Ziegler and Fahlbusch (1986), the double protolophules and posteriorly directed
metalophules in the M2 are increasing through time. When comparing the Kaplangı 1 and 2
specimens to those from Aliveri, Karydiá 1 and Karydiá 2, it turns out that the majority of the molars
have a double protolophule and an anteriorly directed or transversal metalophule, so there are no
major differences when looking at these elements.
Another feature of D. gracilis which changes through time is the mesolophid in the M1. This ridge is
becoming shorter in the younger specimen. When the mesolophids in the molars from Kaplangı are
compared to those from Aliveri, they are all mainly of medium length. The molars from Karydiá on
the other hand, have mainly short mesolophids, which would indicate that they are probably younger
than the Kaplangı specimen.
So based on these features it can be concluded that the specimen from Kaplangı 1 and 2 are of the
same age or older than the specimens from Aliveri and Karydiá (MN 4). This would also agree with
the theory that D. gracilis migrated from east to west, because Kaplangı is the most eastern location
in Anatolia where D. gracilis was found yet.
Gliridae
Order: Rodentia - Bowdich, 1821
Family: Gliridae - Muirhead, 1819
The Gliridae are not identified on the species level due to the small amount of available specimens.
In Kaplangı 1, two different genera are present, the Vasseuromys (Baudelot and de Bonis, 1966) and
Glirudinus (de Bruijn, 1966), while in Kaplangı 2 only Glirudinus is present.
Kaplangı 1
Table 8 - measurements of Vasseuromys sp. and Glirudinus cf. eurydion from Kaplangı 1(in mm).
P4
M1
M2
Vasseuromys M3
sp.
P4
M1
M2
0
0
0
0
3
1
2
M3
0
Glirudinus
cf.
euryodon
Kaplangı 2
Table 9 - of Vasseuromys sp. from Kaplangı 2(in mm).
P4
M1
M2
Vaseuromys M3
sp.
P4
M1
M2
1
2
3
1
1
2
2
M3
0
P4
M1
M2
M3
P4
M1
M2
0
5
1
1
0
2
4
M3
0
Figure 9 - Measurements of Vasseuromys sp. and Glirudinus cf. eurydion from Kaplangı 1 and 2(in mm
Description
The molars of Glirudinus are relatively small, and have a slight concave occlusional surface. The
crowns are very low. These molars have numerous straight thin ridges that form an angle of
approximately 45° with the longitudinal axis of the molar (Daams, 1999). The upper molars have a
very well defined elongated platform along the lingual edge of the molar, which can also be seen in
Glirudinus cf. Euryodon (van der Meulen and de Bruijn, 1982). However, there are not enough molars
available to make a confident statement.
The molars of Vasseuromys are slightly larger than those of the Glirudinus. The occlusion surface is
concave. The longitudinal ridge is continuous and the centrolophid is reaching the labial border
(Daams, 1999).
Discussion
According to Daams (1999), G. euryodon only lived for a very short period of time (MN4). The genus
Vaseuromys on the other hand, mainly lived from MN1 to MN4 (with an exception in MN11), so the
tentative conclusion is that the specimens from Kaplangı lived on the border of MN3 and MN4.
However, the lack of sufficient specimens limits closer determination of the species and therefore
the precision of the age estimates.
Discussion and conclusions
When comparing the faunas from Kaplangı 1 to those of Kaplangı 2 (Appendix 1), it shows that
although the Kaplangı 1 fauna contains more molars, the Kaplangı 2 fauna contains more different
families. In Kaplangı 1 there are 115 molars present in total, divided over 5 families and 11 different
genera. In Kaplangı 2 there are 83 molars in 6 different families and 8 genera. The main differences
are within the families of the Cricetidae, the Sciuridae and the Insectivora. In Kaplangı 1, the
Cricetidae family makes up 61% of the total assemblage, consisting of E. carbonicus, D. gracilis, M. cf.
crenulata and C. cf. aliveriensis. In Kaplangı 2, the Cricetidae make up 45% of the total assemblage,
consisting of D. gracilis and C. aliveriensis. These variations are possibly due to a difference in
palaeoenvironment. The family of the Sciuridae also illustrates some variations between the different
faunas. In Kaplangı 1 there are 3 different genera present (Tamias, Aliveria and Spermophilinus) and
in Kaplangı 2 there is only the genus Tamias. However, this is based on such a limited amount of
specimens that we refrain from drawing any conclusions. The Insectivora are not present in the
larger assemblage of Kaplangı 1, but they do make up 19% of the Kaplangı 2 assemblage. This is also
likely due to a difference in palaeoenvironment, because insectivores require wetter conditions than
the other families present in the assemblages.
Age
For the age determination of the different faunas, we have looked at the individual age estimates
that were discussed in the results-section (table 10).
Table 10 - age estimates of the determined species
Kaplangı 1
Anomalomys aliveriensis
Mirrabella cf. crenulata
Eurmyarion carbonicus
Cricetodon cf. aliveriensis
Democricetodon gracilis
Glirudinus cf. euryodon
Vasseuromys sp.
Estimated age
MN4 or older
MN3 or younger
MN2 or younger
MN 4 or older
MN4 or older
MN4
MN1-4
Kaplangı 2
Anomalomys aliveriensis
Cricetodon aliveriensis
Democricetodon gracilis
Vasseuromys sp.
Estimated age
MN4 or older
MN4 or older
MN4 or older
MN1-4
When averaging the age estimates of the different species it seems that the total fauna of Kaplangı 1
has an estimated age of late MN3 to Early MN4. For Kaplangı 2 this gives an age of MN4 or older, but
because the D. gracilis and A. aliveriensis are very similar in both populations, we argue that Kaplangı
2 can be estimated at the same age as Kaplangı 1: Late MN3 to Early MN4. Thus we conclude that
there are no significant differences in age between the two assemblages.
In the biochronological zonation of Anatolia (Ünay et al, 2003), Kaplangı was assigned to zone E. This
zone correlates to MN4 in the Mammal Neogene biochonological framework defined by Mein (1975).
Based on our results, we conclude that Kaplangı fits in the upper part of zone D (Late MN3) or in the
lower part of zone E (Early MN4).
Palaeoenvironment
To determine the palaeoenvironment of the two Kaplangı assemblages, we have looked at the
individual preferences of the different species. First we will discuss some species that are present in
both localities. The most abundant species in both assemblages is Democricetodon gracilis, according
to Weerd and Daams (1978). This species is adapted to a wet and forested habitat. This is also the
case for Cricetodon (Rummel, 1999), Vasseuromys (de Bruijn, personal communication) and
Glirudinus (Daams et al, 1988). Eumyarion is also thought to have lived in a wet environment (Daams
et al, 1988) with reed-lands or forests (de Bruijn and Saraç, 1999).
Mirrabella, which is only present in Kaplangı 1, is also always found in areas with lacustrine
sequences which indicate forested biotopes (de Bruijn and Saraç, 1992). However it is rare in all the
horizons in which it occurs and it is absent in others that fall within its geographical and
stratigraphical range, probably indicating quite specific habitat requirements which are unknown (de
Bruijn and Saraç, 1992).
The Insectivora, which are only present in Kaplangı 2, also prefer wet and forested biotopes.
However, they prefer denser vegetation and wetter conditions than the other species discussed here
(de Bruijn, personal communication).
We can therefore conclude that the palaeoenvironment in both Kaplangı 1 and 2 is wet and forested.
This is in agreement with the clayey sediment in which the molars were found. There are some small
differences. Kaplangı 1 presumably met some special unknown habitat requirements for Mirrabella
that Kaplangı 2 did not. Keep in mind, though, that this assumption is based on only two molars.
Kaplangı 2 was most likely wetter and contained denser vegetation than Kaplangı 1. This would also
agree with the fact that coal seams were found near the assemblage of Kaplangı 2 (de Bruijn,
personal communication).
Migration patterns
The palaeogeographic maps from Popov et al (2004)(Figure x and x) show that during the Early
Miocene there was no connection between Anatolia and the present day Greece yet. During the
Early-Middle Miocene, these landmasses had already merged and migration of faunas could take
place without major barriers.
The results from Kaplangı can fill some gaps in the knowledge of the migration patterns of different
species.
Anomolomys aliveriensis
When looking at Anomalomys aliveriensis, the specimens from Kaplangı represent the most eastern
finding of this species up to date. The fossil record of Anomalomys is very limited and the results
Kaplangı can therefore add some extra evidence to the theory that Anomalomys originated from
Asia, and migrated towards the west during the Miocene (Feyfar, 1972; Klein Hofmeijer and de
Bruijn, 1985; Bolliger, 1999). The presence of Anomalomys in Greece in MN4 as well as in Kaplangı
during Late MN3 or Early MN4 can indicate that there must have been a connection between Greece
and Turkey in this period.
Democricetodon gracilis
According to Wang and Dawson (1994), the Cricetidae originated in Central Asia during the Eocene,
and migrated towards Anatolia during the Early Oligocene (Theocharopoulos, 2000). The oldest
records of Democricetodontinae are from Inkonak and Kargi, both from Central Anatolia. From there
they migrated over large parts of the Northern hemisphere during the Miocene (Theocharopoulos,
2000). Most specimens of D. gracilis can be found in Central Europe, but the oldest specimens come
from Greece and Anatolia, supporting the theory that they migrated from east to west. The Kaplangı
specimens are farther east, and older than the European findings, which is very much in line with this
theory.
Eumyarion carbonicus
Eumyarion is also thought to have originated from Asia Minor and migrated from there towards the
West in MN4 (de Bruijn and Saraç, 1991). The results from Kaplangı 1 are in agreement with this
theory. The oldest specimen of E. carbonicus can be found east from Kaplangı, in Harami (MN2) and
towards the west, they are becoming younger. Eumyarion is first recorded in Europe during MN4.
This also indicates that the connection between Anatolia and Greece must have existed at the Early
MN4 or Late MN3.
Cricetodon aliveriensis
According to Rummel (1999), Cricetodon also originated from Anatolia and migrated from there
towards Europe. The presence of C. aliveriensis in Kaplangı supports the theory that it lived in
Anatolia, and it is plausible that it had migrated from there towards Greece. Its close resemblance to
C. tobieni from Horlak 1a is an indication that they are closely related and that C. tobieni might have
been an ancestor of C. aliveriensis (de Bruijn et al, 1993). This would also support Rummel’s theory of
the Cricetodon migration from Anatolia to Europe.
Mirrabella cf. crenulata
Because of the lack of specimens we cannot draw firm conclusions on the migration of Mirrabella
species. The only other location where M. crenulata was found is Keseköy, and these specimens were
probably slightly older than those of Kaplangı. But to say that Mirrabella migrated from East to West
is too bold a statement at the current level of knowledge, although it would be in accordance with
the migration patterns that are shown by other Cricetidae genera and species.
Figure 10 - Palaeogeographical map of the Paratethys during MN3 (20.5-19Ma) (Popov, 2004)
Figure 11 - Palaeogeographical map of the Paratethys during MN5 (16-15Ma) (Popov, 2004)
Conclusions
The main conclusions that can be drawn from this research are:
-
Both Kaplangı 1 and 2 are thought to be from the late MN3 or the Early MN4, there are
no major differences in age.
-
The palaeoenvironment in Kaplangı 1 and 2 is both thought to be wet and forested.
However Kaplangı 2 probably had a wetter environment.
-
So the differences between the faunas are mainly caused by the palaeoenvironment
rather than an age difference.
-
The Anomalomidae and most of the Cricetidae are thought to have originated from
Anatolia and migrated from there towards Europe during the Late MN3 or Early MN4
indicating a connection between Anatolia and Greece during this Period.
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Appendix 1 – Fauna list
Kaplangı 1
Kaplangı 2
N
N
Anomalomidae
Anomalomys aliveriensis
12
Cricetidae
Eumyarion carbonicus
Democricetodon gracilis
Mirrabella cf crenulata
Cricetodon cf. aliveriensis
27
38
2
3
Sciuridae
Tamias sp.
Aliveria sp.
Spermophilinus sp.
Spalacidae
Debruijnia sp.
Gliridae
Vasseuromys sp.
Glirudinus cf. euryodon
4
2
Anomalomidae
Anomalomys aliveriensis
9
Cricetidae
Democricetodon gracilis
Cricetodon Aliveriensis
20
18
Sciuridae
Tamias sp.
1
Spalacidae
Debruijnia sp.
2
2
Gliridae
Vasseuromys sp.
12
Insectivora
Talpidae indet.
Soricidae indet.
9
7
2
5
13
Appendix 2: Geological history
Turkey can geologicaly be divided into three main tectonic
units with different characteristics: The Arabian platform,
the Pontides and the Anatolides-Taurides (see figure 1)
(Ketin; 1966). These units used to be separated by the
Thethys ocean. This ocean existed as a large embayment
between Gondwana and Laurussian at least since the
Carboniferous (Stampfli and Borel; 2002), and can be
subdivided into the Neo-Tethys and the Paleo-Tethys
(Şengör ; 1984 and 1987, Stampfli; 2000, Robertson et al;
2004). When the Tethys oceans closed, several sutures
developed that indicate the boundaries between the
Figure 12 - Tectonic setting of Turkey, P = Pontides,
different geological units. The Izmir-Ankara-Erzincan
A-T = Anatolides-Taurides (Okay; 2008)
suture marks the boundary between the Pontides and the
Anatolides-Taurides, and the Assyrian suture connects the Arabian platform to the AnatolidesTaurides (Okay; 2008). In Anatolia both conjugate and exotic margins occur. The boundary between
the Arabian Platform and the Anatolides-Taurides for example, is a conjugate margin. They were
rifted apart during the Triassic, and reassembled during the Miocene. In contrast, the Pontides and
the Anatolides-Taurides were never contiguous. Altough both units were rifted off from Gondwana
(the Pontides during the Ordovician and the Anatolides-Taurides during the Triassic), they used to be
located at widely separated margins (Okay; 2008).
The Anatolian peninsula is surrounded by three different seas: the Black Sea (which is a Cretaceous
back-arc basin, formed by the subduction of the Neo-Tethys (Görür; 1988)), the Aegean (formed by
Oligocene-Miocene north-south extension above the retreating Hellenic subduction zone (Jolivet;
2001)) and the Mediterranean (which is a relic of the southern branch of the Neo-Tethys (Garfunkel;
2004)).
The Kaplangı area is located in the western part of Anatolia, south of the Afyon zone on the
Anatolide-Tauride Block.
The Anatolides-Taurides
The Anatolides-Taurides show Gondwana affinities, but they were separated from the main part of
Gondwana by the southern branch of the Neo-Tethys, it shows a paleozoic stratigraphy which is
similar to the Arabian platform. During the Late Creataceous-Paleocene obduction, subduction and
continental collision episodes, the Anatolide-Tauride terrane underwent much stronger Alpine
deformation and metamorphism than the Pontic zones because of its footwall position. The The
northern margin of the Anatolide-Tauride terrane underwent HP/LT metamorphism at depths of
over 70km under this oceanic thrust sheet. The erosional remnants still occur throughout the
Anatolides-Taurides. When the continental collision started in the Paleocene, the Anatolide-Tauride
was internally sliced and therefore formed a south to south-east vergent thrust pile. In the western
part of Turkey, the contraction continued until the Early- Middle Miocene. Because of the different
ages and types of the Alpine, the Anatolide-Taurides are divided into several zones with different
metamorphic features. These are: the Tavşanlı zone (Cretaceous blueschist belt) in the north, the
Afyon zone (lower grade HP metamorphic belt) in the centre, and the Menderes Massif (Barroviantype Eocene metamorphic belt) in the south. During the Mesozoic, there was an extensive carbonate
platform, where thousands of meters of shallow-water carbonates were being deposited.
The different metamorphic areas also have some common features (that’s why they are combined to
one zone, the Anatolides-Taurides). These are: a late Precambrian crystalline basement, a mixed
clastic-carbonate Paleozoic succession and a thick Upper-Triassic to Upper-Cretaceous carbonate
sequence