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Michael Morlo & Yuriy Semenov

New dental remains of Machairodus KAUP 1833

(Felidae, Carnivora, Mammalia) from the Turolian of Ukraine:
significance for the evolution of the genus
Authors adress: Michael Morlo, Forschungsinstitut Senckenberg, Abteilung Messelforschung,
Senckenberganlage , D.  Frankfurt am Main, Germany,
Yuriy Semenov, Palaeontological Museum, National Museum of Natural History of the National Academy of
Sciences of Ukraine,  Bogdan Khmelnitski Street, Kiev , Ukraine,

New dental remains of Machairodus KAUP,  from Turolian localities of Ukraine are described. While
specimens from Novoukrainka  (late MN ) are very derived and resemble later records from Belka
(late MN ) and Cherevichnoe (MN ), the remains from Novaja Emetovka  (early MN ) are more
plesiomorphic. Previous taxonomy would place these latter specimens in M. aphanistus (KAUP, ), a species
formerly considered to be present from MN  – MN  only, while all other specimens would be assigned to
Amphimachairodus giganteus (WAGNER, ), formerly considered to be present in MN  – MN  only.
As confirmed by the present material, however, both taxa are not as stratigraphically distinct as previously
thought. Moreover, the Ukrainian record provides new evidence for a gradual and mosaic morphological
evolution from M. aphanistus to A.giganteus which is not strictly correlated to cheek tooth size. Consequently,
we consider Amphimachairodus to be a junior subjective synonym of Machairodus. For reasons of taxonomic
stability we retain both species names even though they belong to a single evolutionary lineage.

Neue Reste der Bezahnung von Machairodus KAUP,  aus turolischen Fundstellen der Ukraine werden
beschrieben. Während die Stücke aus Novoukrainka  (spätes MN ) eine sehr fortschrittliche Morphologie

123 kaupia | Darmstädter Beiträge zur Naturgeschichte | Heft 13 | 123–138 | Darmstadt, 28. Dezember 2004 |
aufweisen und darin späteren Resten von Belka (spätes MN ) und Cherevichnoe (MN ) gleichen, sind
die Funde von Novaja Emetovka  (frühes MN ) deutlich plesiomorpher. Die bisher benutzte Taxonomie
würde diese plesiomorphen Stücke zu M.aphanistus (KAUP, ) stellen, eine Art, der bisher nur eine
stratigraphische Verbreitung von MN  bis MN  zugeschrieben wurde. Alle anderen Funde würden
Amphimachairodus giganteus (WAGNER, ) zugeordnet, der bisher nur von MN  bis MN  verbreitet
gewesen sein sollte. Das vorliegende Material belegt allerdings, dass, anders als bisher angenommen, die beiden
Arten stratigraphisch nicht streng zu trennen sind. Darüber hinaus liefert es neue Belege für eine allmähliche
Evolution von M.aphanistus zu A. giganteus, die nicht streng mit der Größe der postcaninen Bezahnung
korreliert. Deshalb interpretieren wir Amphimachairodus als ein jüngeres subjektives Synonym von
Machairodus. Aus Gründen der taxonomischen Stabilität erhalten wir beide Artnamen, auch wenn die Arten
zu einer einzelnen evolutiven Linie gehören.
Introduction The subgenus contains M.(A.) giganteus as the only
The first recognized sabretooth felid genus from species, the stratigraphic distribution of which
Europe, Machairodus, was described by KAUP () was declared to last from MN  to MN  (BEAUMONT
based on an upper canine from MN  of Eppelsheim. , ). The subgenus Machairodus, with
He considered it to belong to the same taxon as a M.(M.)aphanistus as the type species, was said to
canine from the Villafranchium of Val d'Arno, occur only in MN  and MN .
described by CUVIER () as Ursus etruscus sive
cultridens, but a different taxon than Felis aphanista GINSBURG et al. () raised the subgenus
KAUP, , a species from Eppelsheim founded on a Amphimachairodus to genus level, subsequently
mandibular fragment. Later, KAUP () placed both followed by MORALES (), PONS-MOYÁ (,
canines in Machairodus cultridens (CUVIER, ).  |), ALCALÀ (), MARTÌNEZ (), MEIN
He also decided that Felis aphanista and Machairodus (), and ANTÓN et al.(). However, other
giganteus ( WAGNER, ) belong to this taxon and authors still use Amphimachairodus as a subgenus
recognized them as junior subjective synonyms of (e.g., SOTNIKOVA ) or not at all (e.g., MONTOYA
M. cultridens. This taxonomic interpretation lasted ; MORLO ; TURNER & ANTÓN ; GINS-
until FABRINI () who placed the Eppelsheim BURG ; SPASSOV & KOUFOS ; ROUSSIAKIS &
canine into M. aphanistus ( including M. giganteus ) THEODOROU ), that is, implicitly as a junior
but separate from Machairodus cultridens. In so synonym of Machairodus.
doing, M. aphanistus implicitly became the type spe-
cies of Machairodus. Later, BOULE () interpreted In this paper, we focus on new dental remains of
Megantereon megantereon CROIZET & JOBERT,  Turolian Machairodus from Ukraine and thus on the
to be a junior synonym of Machairodus cultridens, species M.aphanistus and M.giganteus (Table ).
but still placed it into the same genus as the separate The numerous postcranial remains of this record will
species M.aphanistus (including M.giganteus). be published elsewhere. It should be noted, however,
MATTHEW () confirmed Machairodus aphanistus that Machairodus is also known from Africa (e.g.,
( including M. giganteus ) as the type species of STROMER ; ARAMBOURG ; KURTÉN ;
Machairodus. He also separated it from Machairodus PETTER & HOWELL ), Asia (e.g., ZDANSKY ;
cultridens on genus level by placing the latter species SOTNIKOVA ; QI ; ZONG & JIANG ), and
into Megantereon. As the oldest available name, North America (see MARTIN  for a summary).
Megantereon cultridens thus became the type species As the most eastern European members yet known,
of the genus Megantereon (even if MATTHEW consi- the specimens described here shed new light on the
dered the species name cultridens not to be valid and evolution and paleobiogeography of the two species.
used M.megantereon instead). The hitter to known Turolian record of Ukrainian
Machairodus consists of only three specimens.
Two other specimens from Eppelsheim, originally
described in the same publication as Felis aphanista One of the oldest, an undescribed (but availably to us
KAUP, , were subsequently placed in by a cast only), complete lower dentition from Krivoj
M. aphanistus. KAUP () himself assigned a lower Rog (early MN ), is morphologically identical to the
canine (Amphicyon sp. KAUP ) to the species type material of M.aphanistus. The other specimens
while BEAUMONT (, ) subsumed Felis prisca come from the localities Issaevo (PAVLOW ;
KAUP, , based solely on an isolated P. SOTNIKOVA in prep.) and Grebeniki (PAVLOW ;
Other Machairodus species had been described from SOTNIKOVA ). While Grebeniki belongs to
Europe by, e.g., ROTH & WAGNER () and MN-zone , the age of Issaevo is unclear and might
RIABININ (), and from Asia (e.g., ZDANSKY , be MN  or MN  (Table ). ›Pseudaelurus
ORLOV  and SENYÜREK ), before BEAUMONT intrepidus‹ (PAVLOW, : pl. , fig. ) from Issaevo is
(, ) subsumed most to two species, the ›small represented by a fragmentary mandible with m.
and earlier‹ M.aphanistus (KAUP, ) and the ›large It was placed in Miomachairodus pseudailuroides
and later‹ M.giganteus (WAGNER, ). Differentia- SCHMIDT-KITTLER,  by SCHMIDT-KITTLER, in
tion of these two species is based on differences Machairodus robinsoni KURTÉN,  by BEAUMONT
between the type material of M.aphanistus from MN  () or in Machairodus cf. aphanistus by
of Eppelsheim (KAUP : pl., figs.  – ; : pl.  b, BEAUMONT (). Furthermore, M.alberdiae
figs. –; BEAUMONT : fig. ) and of M. giganteus GINSBURG et al.,  (= plesiomorphic M.aphanistus
from MN  of Pikermi (WAGNER : pl. , fig. ; sensu MORLO ) from MN  of Los Valles de
BEAUMONT : fig. a). Fuentidueña, was declared to be close to the
Ukrainian specimen by MARTÍNEZ ().
BEAUMONT () used the supposed morphologic
and stratigraphic differences between the two species The presence of a relatively large talonid separates
to place M.giganteus in its own subgenus ›Pseudaelurus intrepidus‹ as well as M.alberdiae from
Amphimachairodus KRETZOI, . This taxon was the holotype of M.aphanistus, but that does not
originally based on M.palanderi ZDANSKY,  which necessarily exclude them from the species, especially
had been included in M.giganteus by BEAUMONT if regarding the relatively late age of the Issaevo
(), thereby suggesting a highly variable and maybe mandible (see MORLO  for the last detailed dis-
sexual dimorphic (TURNER & ANTÓN : pl. , fig. cussion). In contrast to the specimen of Issaevo,
.) species (see also ZDANSKY  and BEAUMONT ›Pogonodon copei‹ (PAVLOW, ) from Grebeniki is
: fig. ). a much more advanced form with additional accesso-
ry cusps in P and an anterior cingulum cusp in P Machairodus aphanistus (KAUP, )
(SOTNIKOVA ). These apomorphic character Holotype: HLMD-Din , fragment of right lower
states contrast to a large, plesiomorphic protocone in mandible with p-m (KAUP : pl.  , figs. – 5; KAUP
P. Due to the peculiar combination of apomorphic : pl. 6 b, figs. a–b; BEAUMONT : fig. ), MN 
and plesiomorphic character states SOTNIKOVA () of Eppelsheim.
interpreted the Grebeniki material as a separate spe- Synonyms: Felis aphanista KAUP, ; Felis prisca
cies, Machairodus copei, while BEAUMONT () had KAUP, ; pars Machairodus cultridens KAUP ;
included it in the highly variable M.giganteus. Epimachairodus romeri SENYÜREK, ; Machairodus
taracliensis in FRANZEN & STORCH ();
Machairodus romeri in SCHMIDT-KITTLER ();
Material and Methods pars Miomachairodus pseudailuriensis (see BEAUMONT
All material was unearthed during field-work headed ; MORLO ); Machairodus laskarevi LUNGU,
by the late DR.E.L.KOROTKEVICH and by Y. SEMENOV, ; Machairodus alberdiae GINSBURG et al., .
and is now housed in the Palaeontological Museum A detailed synonymy list of the species is given in
(National Museum of Natural History of the National MORLO ().
Academy of Sciences of Ukraine) in Kiev. The material New material from the Turolian of Ukraine:
comes from Novoukrainka  (late MN ), Novaja From Novaja Emetovka  (early MN ):
Emetovka  (early MN ), Belka (late MN ), and . NNPM Ca-, right I  (.:.), Fig. .
Cherevichnoe (MN ). Stratigraphy (Table ) and . NNPM Ca-, fragment of right maxilla with
lists of the carnivoran faunas of these localities are (P) and P3 (.:.), Fig. 5.
discussed in SEMENOV (). Assignments to MN- . NNPM Ca-, isolated right i, Fig. .
zones for other localities was mainly done after MEIN . NNPM Ca-, isolated fragment of right p.
(). As is discussed by SEMENOV (), two faunas . NNPM Ca-, isolated fragment of left P (-:.),
exist from Novoukrainka, one from late MN , the Fig. .
other from the latest MN . This is noteworthy, . NNPM Ca-, isolated left p (.:.), Fig. .
because in MEIN () Novoukrainka is placed solely . NNPM Ca-, isolated right m (.:.), Fig. .
at the border of MN  | (= Novoukrainka ). All . NNPM Ca-, isolated fragment of right upper
material discussed herein comes from Novoukrainka  canine (.:.), Fig. .
and thus from the late MN . All measurements are All specimens from Novaja Emetovka  were found
made with calipers to the nearest .mm. Tooth in  in an area of about  m2. They, thus, very
measurements given are max. length to max. breadth. probably belong to a single individual.
Tooth morphology nomenclature follows VAN VALEN
(). Upper teeth are given in capitals (like ›P‹), Machairodus giganteus (WAGNER, )
lowers as ›p‹. Tooth positions verified by alveoli only, Holotype: left mandible with i-, (p), p-m
are indicated by brackets. (WAGNER , : pl. , fig. ; BEAUMONT :
fig. a) from MN  of Pikermi.
Institutional abbreviations Synonyms: M. leoninus ROTH & WAGNER ;
HLMD Hessisches Landesmuseum Darmstadt (D). Pogonodon copei PAVLOW,  (= Machairodus copei
NNPM National Museum of Natural History, PAVLOW,  in SOTNIKOVA ); M. palanderi
Kiev (UA). ZDANSKY, ; M. tingii ZDANSKY, ;
SMF Forschungsinstitut Senckenberg, Frankfurt M. aphanistus var. taracliensis RIABININ, ;
am Main (D). M. irtyschensis ORLOV, . The last taxonomic
overview was provided by MARTÍNEZ ().
New material from the Turolian of Ukraine:

125 Michael Morlo & Yuriy Semenov | Turolian Machairodus from Ukraine |
Systematic Paleontology From Novoukrainka  (late MN ):
Carnivora BOWDICH,  . NNPM -, fragment of left mandible with
Feliformia KRETZOI,  (p), fragment of p, m (.: .), Fig. .
Felidae FISCHER von WALDHEIM,  . NNPM  -, fragment of right juvenile
Machairodus KAUP,  mandible with fragment of dp (. :-), dp
(.:.) and erupting m, base of p still in the
Type species: M.aphanistus (KAUP, ). mandible (KOROTKEVICH : fig. d), Fig. .
Included species: M.giganteus WAGNER, ; . NNPM -, isolated fragment of upper right
M.africanus ARAMBOURG, ; M.robinsoni canine ( . : .), Fig. .
KURTÉN, ; M. kurteni SOTNIKOVA, ; . NNPM -, isolated fragment of p (-.),Fig..
M. coloradensis COOK, ; M. tanneri MARTIN & From Belka (late MN ):
SCHULTZ, . Questionable: M.ischimicus ORLOV, . NNPM -, isolated right p (.:.), Fig. .
 (only known from a right metacarpal ). From Cherevichnoe (MN ):
Synonyms: Pogonodon in PAVLOW (); . NNPM -, lower canine with crenulations on
Amphimachairodus KRETZOI,  except both edges, Fig. 
Amphimachairodus pliocaenicus PONS-MOYÁ,  . NNPM -, juvenile maxilla with dC, erupting
(= Homotherium); pars Epimachairodus KRETZOI, C, dP–, Fig. .
 (see BEAUMONT ); pars Miomachairodus . NNPM -, juvenile mandible with dp
SCHMIDT-KITTLER,  (see BEAUMONT ; (.:.), dp (.:.), and erupting m 
MORLO ). (ca. .:.), Fig. .
. NNPM -, isolated fragment of left P
(.:-), Fig. .
Descriptions and comparisons: Deciduous teeth are preserved in NNPM -,
Because we focus on the similarities and differences a right maxilla from Cherevichnoe (Fig. ). The
of the new specimens, belonging to both species, all maxillar foramen is placed above the anterior margin
are described together. of (d)P (see below). The dC is small, narrow, and
has a tip, which is slightly angled backwards. Its root
Upper dentition: does not extend to the maxillar foramen. Antero-lin-
The isolated right I from Novaja Emetovka  (NNPM gual to it, the uncrenulated tip of the erupting per-
Ca-, Fig. ) has strong internal and external cingu- manent canine is visible. A similar situation can also
lae and an abrasion, which runs from the tip, external be observed in a specimen assigned to M. palanderi
to the base. This abrasion is caused by wear with the (= M.giganteus) by ZDANSKY (: pl. , figs.  – ).
lower canine. ZDANSKY interpreted the preserved teeth as perma-
In the left upper canine from Novaja Emetovka  nent C starting to erupt, vestigial P, and P- erup-
(NNPM Ca-, Fig. ) the posterior crest is strongly ted, suggesting that the permanent C erupted much
crenulated, while the erupting canine in NNPM later than the premolars. This interpretation contrasts
- (Fig. ) shows that this crenulation does not clearly with NNPM -. Moreover, the P of
reaches the tip. A right upper canine from ZDANSKY's specimen resembles a dP in having the
Novoukrainka  (NNPM -, Fig. ) was broken protocone being posteriorly shifted. If this tooth
during lifetime directly above the root. Strong indeed is a dP the specimen differs even more from
abrasion marks at the anterior part of the breakage NNPM - in preserving P and a functional dP.
indicate that the individual continued to live after It thus may belong to a different taxon as was already
the canine was broken, as is also documented in mentioned by ZDANSKY himself. Posterior to the dC
Smilodon (VAN VALKENBURGH & HERTEL ) and of NNPM -, a vestigial, single-rooted tooth is
Nimravides (TURNER & ANTÓN : fig. .). separated from the other teeth by diastemata. Some
adult specimens of Machairodus have a vestigial P,
For P (or dP, see below), only the alveolus is preser- e.g., specimen  in ZDANSKY (: pl. , figs.  –),
ved. The tooth was single-rooted and stood close to M. kurteni (see SOTNIKOVA : fig. ), Machairodus
the upper canine. The right maxillar fragment from MN  of Baltavar (see BEAUMONT ), and
(NNPM Ca-, Fig. ) to which it belonged, comes NNPM Ca- from Novaja Emetovka , described
from Novaja Emetovka . The fragment still contains above. No specimen is known to us documenting a
the P, consisting of an anteriorly enlarged anterior replacement of dP. It could prove whether a vestigial
cingulum, anterior and posterior basal cusps, a tooth as in NNPM - is a dP or an early
dominant paracone, and a posterior cingulum cusp. erupting P. Similarly, the vestigial teeth in the adult
The tooth lacks crenulations. Lingual to the postpara- specimens could represent P or a persisting dP. In
crista, a small but clear enlargement is present, the any case, vestigial teeth vary intraspecifically among
remnant of the protocone. Carnivora and should not be used as taxonomical
characters (see DAYAN et al. ). The dP resembles
The isolated P from Cherevichnoe (NNPM -, P in having an anterior cingulum cusp, an anterior
Fig. ) preserves only the crown. It is a very advanced accessory cusp, and a metastyle. The paracone is low,
tooth with all cristae anterior to the paracone being but still the highest cusp. A well developed protocone
crenulated. A large parastyle cusp and a preparastyle lies directly lingual to the paracone and sits on its
cusp are developed, all separated by deep notches. own root. The tooth can be identified as being deci-
Antero-labial to the preparastyle cusp the anterior duous due to the widely diverging three roots. This is
cingulum builds an additional small cusp. A small also the case in the triple-rooted dP which resembles
protocone exists which is placed lingual to the ante- M  in being much broader than long. It consists of
rior half of the preparacrista. Its most lingual part is a labially pointing parastyle, a low paracone (but
broken which is why the breadth of the tooth cannot nevertheless the highest cusp), a postero-labially
be measured. The postprotocristid is separated from pointing and very short metastyle, and a protocone of
the metastyle by a very deep notch. The long but low nearly equal height to the paracone.
metastyle curves labially in its posterior third. The
posterior end of the tooth is obscured by breakage, Lower dentition:
but a posterior cingulum cusp may have been pre- The i (NNPM Ca-, Fig. ) from Novaja
sent. The P-fragment from Novaja Emetovka  Emetovka  has an external cingulid only and is
(NNPM Ca-, Fig. ) consists of the part anterior highly asymmetrical. Accessory cusps are lacking.
to the tip of the paracone. It differs from NNPM The lower canine from Cherevichnoe (NNPM -
- in lacking any crenulations, having the , Fig. ) shows two crenulated crests with the
parastyle cusp lower and not separated by a notch, crenulations running over the tip. In the p4 NNPM
the notch between preparacrista and preparastyle - (Fig. ) from Novoukrainka  the part ante-
cusp shallower, and the part of the tooth anterior to rior to the preprotocristid is missing. The tooth
this notch being shorter relative to paracone height. shows the typical morphology of Machairodus with
A fairly large protocone is present. high protoconid, distinct posterior basal cusp and
posterior cingulid cusp, all separated from each other
by deep notches. Pre- and postprotocristids are
crenulated and a very small enlargement exists
lingual to the posterior end of the postprotocristid,
resulting in the tooth having its maximal breadth
here. The p-fragment of NNPM - (Fig. ),
from Novoukrainka  additionally shows an anterior Deciduous teeth: Two juvenile mandibles are known,
basal cusp and an anterior cingulid, which is slightly one from Novoukrainka  (NNPM -, Fig. ),
enlarged anteriorly. In contrast to the specimens from and the other from Cherevichnoe (NNPM -,
MN , the left p from Novaja Emetovka  (NNPM Fig. ). Both are nearly in the same ontogenetic
Ca-, Fig. ) is smaller and lacks an anterior stage with dp- present and m1 erupting. However,
cingulid cusp as well as crenulations on the postpro- dp of NNPM - is much more worn, indica-
tocristid. Additionally, the small protoconid is narro- ting a slightly older individual. The dp is broken in
wer than in all other Ukrainian specimens except a NNPM -, leaving only the roots and the ante-
right p-fragment from Novaja Emetovka  (NNPM rior accessory cusp visible. In height, this cusp is
Ca-), which lacks its anterior part in front of the similar to that of NNPM -, but in the latter
preprotocristid. Here, also, the protoconid is relative- specimen the anterior cingulid is slightly prolonged
ly small and narrow and crenulations are lacking on to anteriorly, producing a lower angled anterior cris-
pre- and postprotocristid. Finally, NNPM -, tid of the accessory cusp. Besides the anterior acces-
an isolated right p from late MN  of Belka (Fig. ) sory cusp, a posterior accessory cusp and two basal
is more similar to NNPM - from Novoukrainka posterior cuspids are developed in dp. The crown is
 than to the specimens from Novaja Emetovka . It dominated by the high and lanceolate protoconid.
differs from the latter in being slightly larger, having Both dp's are very similar in being molariform,
the protoconid broader, and the preprotocristid having a high, lanceolate protoconid, a metaconid
crenulated. placed directly posteriorly to the protoconid, and
a very reduced talonid which consists of nothing
The m is represented by adult specimens from more than a cingulid swelling. This last character is
Novoukrainka  (NNPM -, Fig. ) and Novaja obscured in NNPM - due to abrasion.
Emetovka  (NNPM Ca-, Fig. ), as well as in an A carnassial blade is developed in both specimens.
erupting state in the juveniles from Novoukrainka  Due to the strong abrasion of its protoconid,
(NNPM -, Fig. ), and from Cherevichnoe the paraconid of NNPM - seems to be higher
(NNPM -, Fig. ). The nearly completely than in NNPM -. Abrasions occur also on the
preserved specimen from Novaja Emetovka  shows labial side of the postprotocristid which are produ-
plesiomorphic character states in being relatively ced by a functional protocone of dP. The p is still
broad and retaining a talonid. Its carnassial blade is deep in its crypt, but its postero-lingual corner with
highly abraded, so possible crenulations are no longer a pronounced cingulid is visible in NNPM -.
visible. The paraconid is antero-lingually enlarged,
but whether a lingual preparacristid was present is no
longer discernable due to abrasion. However, even Discussion
if it was developed, it must have been very small. In Three parameters have been used to differ between
NNPM - (Fig. ) and - (Fig. ) only the the species M.aphanistus and M.giganteus: stratigra-
unworn most occlusal part is visible because these phic distribution, size, and dental morphology. The
were still erupting teeth. In both specimens, a lingual record of Machairodus from the Turolian of Ukraine
preparacristid is developed at the paraconid. While in provides new information for all three of these.
m  from Cherevichnoe all occlusal crests are crenula- BEAUMONT () believed M.aphanistus to be
ted, only the postprotocristid is crenulated in NNPM Vallesian, and thus occurring in MN  and MN ,
-. The carnassial blade dominates the tooth. and M.giganteus to be Turolian, occurring from
NNPM - lacks a metaconid and has the MN  to MN . Some authors follow the original
talonid reduced to a cingulid swelling. This part is concept of BEAUMONT, e.g., SOTNIKOVA (),
not visible in NNPM -, but it is in the slightly MORALES (), MEIN (), and ROUSSIAKIS &

127 Michael Morlo & Yuriy Semenov | Turolian Machairodus from Ukraine |
damaged specimen NNPM - (Fig. ). This m THEODOROU (). According to GINSBURG
is damaged at the carnassial notch, the blade is (), Machairodus is not present in MN  at all.
abraded, and the most anterior part of the paraconid However, other described specimens from MN  are
is broken. However, if a lingual preparacristid had believed to belong to M.aphanistus (e.g., BONIS 
been present, it would be visible and it is not. The from Kemiklitepe; MONTOYA & MOMPARLER 
postprotocristid is strongly crenulated and the and MARTÍNEZ  from Crevillente-, contra
metaconid and talonid are completely reduced. MORALES  who assigned this latter material to
NNPM -, thus, shows a mixture of very derived M.giganteus) or to M. cf. aphanistus (MORLO 
(strong crenulation of the postprotocristid, lack of from Dorn-Dürkheim ). This led to the idea, that
metaconid and talonid) and more plesiomorphic M.aphanistus was present from MN  to MN , while
(lack of a lingual preparacristid) character states. M.giganteus occurred in MN  and MN  (MARTÍNEZ
). Consequently, questionable specimens were
reassigned to fit the proposed stratigraphic ranges.
M. giganteus from MN  of Terrassa, described by
PONS-MOYÁ (|) as Amphimachairodus
giganteus, was placed in M.aphanistus by MARTÍNEZ
() because the specimen, a fragment of an upper
canine, would not allow for a certain specific assign-
ment. However, its size (PONS-MOYÁ  |) fits
well with M.giganteus (see below). The about
contemporary Ukrainian specimens from Issaevo BEAUMONT : fig. c, : pl.  from MN  of
(plesiomorphic M.aphanistus) and Grebeniki Montrédon; MORLO  from MN  of Dorn-
(plesiomorphic M.giganteus) additionally implied a Dürkheim), increased length of parastyle and prepa-
separation of the species by stratigraphical occurren- rastyle, and enlarged cusps (e.g., BEAUMONT &
ce to be problematic. The material described herein CRUSAFONT PAIRÓ : , pl. , fig.  from MN 
corroborates this and renders a strict stratigraphic of Santiga). The P of apomorphic M.giganteus
separation between the species obsolete: the occur- shows a large metastyle, low paracone, high parasty-
rence of apomorphic specimens in Novoukrainka  le, high preparastyle, reduced protocone (ZDANKSY
show for the first time definitely that M. giganteus : pl. , figs. , , pl. , fig. , pl. , fig. ,
was present in MN , while the much more plesio- BEAUMONT : figs.  c – d), and crenulations on all
morphic specimens of Novaja Emetovka  verify the cristae. The P-fragment from Novaja Emetovka 
presence of M.aphanistus until at least early MN . is similar to that of advanced M.aphanistus from
MN , as known from Eppelsheim (in difference to
Size is the second parameter used to distinguish plesiomorphic specimens as »M.alberdiae« or
between M.aphanistus and M.giganteus. After »M.romeri«). NNPM - from Cherevichnoe
BEAUMONT (), M.aphanistus is the »small and (MN ), instead, reflects the character states of
earlier« species while M.giganteus is »large and M.giganteus closely. Moreover, it resembles the
later«. However, in the same publication BEAUMONT Grebeniki P (SOTNIKOVA ) in having a small
demonstrated P to show a size variability of about cingulum cusp antero-labial to the preparastyle, cor-
% among the »large« M.giganteus (BEAUMONT roborating a close relationship of ›Pogonodon copei‹
). As was verified by SOTNIKOVA (: fig. ) for to M.giganteus (sensu BEAUMONT ).
P and by SPASSOV & KOUFOS (: fig.) for m of
both species, a separation based only on size is also The p of plesiomorphic M.aphanistus is known
not possible with these teeth (see also ROUSSIAKIS & from the holotype HLMD Din  (KAUP : pl.,
THEODOROU  for a recent overview). This is figs.  – , : pl.  b, figs.  a –b). It lacks anterior and
corroborated by additional evidence, given herein, posterior cingulid cusps, has a weak anterior acces-
for the sizes of p (Fig.): Ukrainian M.giganteus is sory cusp, and lacks any crenulations (see also
clearly smaller than several Central and Western BEAUMONT : fig.  from MN  of Charmoille).
European specimens of M.aphanistus. PONS-MOYÁ The complete p (NNPM Ca-) from early MN 
(|: fig.) presented data separating both spe- of Novaja Emetovka  (Fig. ) is very similar to the
cies by the respective sizes of the upper canines, with holotype of M.aphanistus, while SMF DD- from
the upper canine of M.giganteus about as broad, MN  of Dorn-Dürkheim (, see
but antero-posteriorly longer than in M.aphanistus, MORLO ) demonstrates the gradual changes
leading to a relatively more narrow sabre-tooth. towards M.giganteus in having the accessory cusp
Partly based on this evidence, the author placed the higher. The German specimen, however, still lacks
specimen from MN  of Terrassa into M.giganteus. crenulations and an anterior cingulid cusp seen in p
The upper canine of M.aphanistus from Novaja of apomorphic M.giganteus from MN  of Pikermi
Emetovka  (Fig. ) is clearly shorter than in (WAGNER : pl. , fig. ). Still more close to
M.giganteus from Novoukrainka  (Fig. ). Moreover, M.giganteus are two specimens from Crevillente-
both fit well into the respective size variability of the (MONTOYA & MOMPARLER ) which lack an
species given in PONS-MOYÁ (|: fig. ). Howe- anterior cingulid cusp, but have the cingulid enlarged
ver, ROUSSIAKIS & THEODOROU () recently to anteriorly. The holotype of M.giganteus itself
presented data which does not fit into this picture. additionally has a strong anterior accessory cusp, a
Due to the small number of yet measured specimens strong posterior cingulid cusp, and is relatively more
either interpretation has to be confirmed with addi- slender. As was shown by BEAUMONT (: fig. ),
tional material, thus. Other reasons for a metric varia- however, the anterior cingulid cusp might be absent
bility of the upper canine, as, e.g., sexual dimorphism, in M.giganteus (see, e.g., ROTH & WAGNER ;
could not be analysed yet due to the poor specimens RIABININ ; MORALES & SORIA ; ALCALÁ ).
record, but might be involved as well.
The Ukrainian specimens from Novoukrainka  and
Because a separation of M.aphanistus from Belka are very similar to the holotype of M.giganteus,
M.giganteus by sizes of premolars and molars is not especially in being crenulated on all cristids and
possible, several morphological characters were having an anterior cingulid cusp. However, both p
suggested to do so (e.g., BEAUMONT , ; are smaller than all other M.giganteus and also smaller
SOTNIKOVA ; MARTÍNEZ ; SPASSOV & than most p of M.aphanistus, which are morpholo-
KOUFOS ). Here, we focus on P, p, and m gically more plesiomorphic (Fig. ). There is thus no
as the functionally most important cheek teeth. simple relationship between morphology and size as
The P of plesiomorphic M.aphanistus is known was implied by the statement of BEAUMONT ()
from MN  of Eppelsheim (HLMD Din , holotype when declaring M.aphanistus to be the smaller spe-
of »Felis prisca« KAUP, : pl.  b, fig. ; HLMD Din cies relative to M.giganteus. The complete p and the
, BEAUMONT : figs. c–d). It has a short fragmental specimen from Novaja Emetovka  are
metastyle, high paracone, low parastyle cusp, large the smallest p yet known from M.aphanistus.
protocone and lacks a preparastyle. Crenulations are SPASSOV & KOUFOS () recently gave an overview
lacking, at least on the preparacrista. More advanced
specimens show an enlarged metastyle (e.g.,
on the morphological variability of m among Conclusions
Machairodus aphanistus and M.giganteus. They Analysis of the dental remains of Turolian
recognized the existence of two morphotypes with Machairodus from Ukraine supports former
a relatively broad trigonid and the presence of an suggestions based on other European specimens
»expressed metaconid-talonid complex« being (BEAUMONT , , ; MORLO ) that
significant for M.aphanistus, while M.giganteus have a gradual but mosaic evolution occurred from
a narrow trigonid and a reduced talonid with the M.aphanistus to M.giganteus. Between the species,
metaconid no longer recognizable. In Machairodus, large overlaps existed in size and stratigraphic
the m is broadest at the protoconid. Consequently, distribution. Concerning the morphological changes
relative breadth should differ between M.aphanistus in the dentition, P, p, and m show a gradual
and M.giganteus if its trigonid constantly is relatively development of single character states, like the
broader. However, m  sizes, given in MARTÍNEZ increasing length of the metastyle, parastyle, and pre-
(: fig. ), SPASSOV & KOUFOS (: fig. ), parastyle in P, increasing height of the accessory
and ROUSSIAKIS & THEODOROU (: figs.  –) cusp in p, or the degree of reduction in metaconid
do not show such a difference in relative breadth. and talonid of m. However, in single specimens,
The second character, the »metaconid-talonid different characters may show states which were
complex«, is not simply reduced as a whole as was previously assigned to either species, such as the
indicated by SPASSOV & KOUFOS (). This is absence of a lingual preparacristid in NNPM -
shown by an m from MN  of Soblay (VIRET & while metaconid and talonid are highly reduced or
MAZENOT ) which has the metaconid reduced, the absence of a crenulation at the preprotocristid in
while the m from Zillingsdorf (Late Miocene but NNPM - while the lingual preparacristid is
unclear MN-zone, pers. comm. D. Nagel) has the present.
talonid reduced (BEAUMONT : pl. , fig. ). More-
over, the metaconid-talonid complex as a whole is The overlaps in size and stratigraphic distribution
highly, but to a different degree, reduced in as well as the mosaic morphological variability
specimens which were assigned to M.aphanistus of the dentition indicate that the two species are
(BEAUMONT , , MN  from Montrédon; representatives of a single evolutionary lineage.
LUNGU , MN  of Kishinev; MORLO , MN  Consequently, previous generic separations of
of Dorn-Dürkheim). Finally, m  of M.giganteus M.giganteus which did not base on phylogeny but
from MN  of Pikermi (WAGNER : pl. , fig. ) simply on morphological or functional differences
shows a very small metaconid but has the talonid of this species to plesiomorphic M.aphanistus (as
missing. Relative trigonid breadth as well as relative has been done first by GINSBURG et al.  using
size of metaconid and talonid do distinguish plesio- dental morphology and last by ANTÓN et al. 
morphic M.aphanistus (as the holotype) from apo- using functional morphology of the cranium) have
morphic M.giganteus, but a couple of morphologi- to be rejected. If arguing strictly phylogenetically,
cally intermediate specimens exist. Moreover, among even a separation into two species could be questio-
M.giganteus itself, the metaconid-talonid complex ned. In other cases, as the late Miocene mustelid
also is reduced to a very different degree (see Eomellivora wimani (see WOLSAN & SEMENOV )
BEAUMONT : fig. ; SPASSOV & KOUFOS: fig. ), or the Pliocene ursid Ursus minimus (see MORLO &
indicating a graded difference between the species KUNDRÁT ), chrono-subspecies were defined to
rather than the existence of clearly separate morpho- taxonomically reflect morphological changes within
types. Beside a relatively broad trigonid and a relati- a single evolutionary lineage. Such a taxonomic
vely large »metaconid-talonid complex«, the m of procedure is not possible for M.aphanistus and
the holotype of M.aphanistus (KAUP : pl. , figs. M.giganteus, because of their largely overlapping

129 Michael Morlo & Yuriy Semenov | Turolian Machairodus from Ukraine |
 – ; : pl. b, figs.  a – b) lacks the crenulations, as stratigraphic distributions which may reflect a
well as a lingual preparacristid. In the m of the dynamic migration of the genus in all of Europe
M.giganteus holotype (WAGNER, : pl. , fig. ), (and probably Asia), irrespective of the migrant's
the complete carnassial blade (postparacristid and morphological specialization. Based on this new
preprotocristid) is crenulated and a lingual prepara- evidence, J. J. KAUP had a very modern, but too
cristid is present at the paraconid. In the Ukrainian strictly phylogenetic view, when in  he assigned
specimens, m from Novaja Emetovka  (Fig. ) is the specimens of M.aphanistus from Eppelsheim
very similar to the holotype of M.aphanistus while and the holotype of M.giganteus from Pikermi to a
the erupting m from Cherevichnoe (Fig. ) is very single species.
close to advanced M.giganteus. The specimens from
Novoukrainka  also resemble M.giganteus, but show
plesiomorphic characters: NNPM - (Fig. ) Acknowledgements
lacks a lingual preparacristid while the erupting m We thank Mrs. Anika Hebs (SMF) for creating the
of NNPM - (Fig. ) has the lingual prepara- plates and Gregg Gunnell (Univ. of Michigan) for
cristid present, but only the postparacristid crenula- correcting the English. Doris Nagel (Univ. of Vienna)
ted and not the preprotocristid. Such differences as and Gregg Gunnell improved a former version of the
the presence/absence of a lingual preparacristid in manuscript by comments and discussions. Finally,
individuals from a single locality, corroborate the Elmar P. J. Heizmann (Staatliches Museum für Natur-
independent variability of different character states kunde Stuttgart) increased the manuscript's quality
among m  of Machairodus. by his detailed review comments.
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Fig. 1–4: Upper dental
elements of Machairodus
KAUP, 1833 from the Turolian
of Ukraine.

Fig. 1: NNPM Ca-1269, isolated

right I3 of Machairodus
aphanistus (KAUP, 1832) from
Novaja Emetovka 2 (early MN
12) in a: posterior, b: external
view. Notice strong external
wear facet caused by the lower
canine, confirming this to be
an upper I3.

Fig. 2: NNPM Ca-1276, isolated

fragment of right upper
canine of Machairodus
aphanistus (KAUP, 1832) from
Novaja Emetovka 2 (early
MN 12) in a: lingual and
b: labial view. Note crenulated
posterior crest.
1a. 1b.
Fig. 3: NNPM 45-3788, right
maxilla of juvenile
1cm Machairodus giganteus
(WAGNER, 1848) from
Cherevichnoe (MN 13) with dC,
erupting C, dP2-4 in a: lingual
and b: labial view.

Fig. 4: NNPM 38-1760, isolated

2a. 2 b. fragment of right upper
1cm canine of Machairodus
giganteus (WAGNER, 1848)
from Novoukrainka 1 (MN 11)
in a: lingual and b: enlarged
frontal view. Note wear facets
indicating that the tooth was
functional after being broken.


133 Michael Morlo & Yuriy Semenov | Turolian Machairodus from Ukraine |
4 a.

3b. 4b.
1cm 1cm
Fig. 5–7: Upper dental
elements of Machairodus
KAUP, 1833 from the Turolian
of Ukraine.

Fig. 5: NNPM Ca-1270, right

maxillary fragment of
Machairodus aphanistus (KAUP,
1832) from Novaja Emetovka 2
(early MN 12) with (P2) and P3.
P3 in a: labial, b: occlusal, and
c: lingual view. Note single
alveolus of P2 and lingual 5a. 5c.
enlargement of the posterior

Fig. 6: NNPM 45-26319,

isolated fragment of left P4
of Machairodus giganteus
(WAGNER, 1848) from
Cherevichnoe (MN 13) in a: 5b.
labial, b: occlusal, and
c: lingual view. Note large
parastyle, large preparastyle,
and small protocone.

Fig. 7: NNPM Ca-1273, frag-

ment of left P4 of Machairodus
aphanistus (KAUP, 1832) from
Novaja Emetovka 2 (early MN
12 a) in a: occlusal, b: lingual
view. Note low parastyle, low
preparastyle, and large

6a. 7a.




Fig. 8 –11: Lower dental
elements of Machairodus
KAUP, 1833 from the Turolian
of Ukraine.

Fig. 8: NNPM Ca-1271, isolated

right i 2 of Machairodus
aphanistus (KAUP, 1832) from
Novaja Emetovka 2 (early MN
12) in a: posterior, b: internal

Fig. 9: NNPM 45-3380, lower

canine of Machairodus
giganteus (WAGNER, 1848)
from Cherevichnoe (MN 13) in
a: external and b: internal

9a. Fig. 10: NNPM 38-1763, isolated

fragment of left p4 of
Machairodus giganteus
(WAGNER, 1848) from
Novoukrainka 1 (MN 11)
in a: occlusal and b: lingual
view. Note crenulated
8a. 8b.
1cm Fig. 11: NNPM 38-795, left
9b. mandible fragment with (p3),
1cm fragment of p4, and m1 of
Machairodus giganteus
(WAGNER, 1848) from
Novoukrainka 1 (MN 11) in a:
lingual, b: occlusal, and
c: labial view. Note in p 4
presence of an anterior basal
cusp, in m1 reduced lingual
preparacristid, completely
lacking metaconid, and
completely reduced talonid.





135 Michael Morlo & Yuriy Semenov | Turolian Machairodus from Ukraine |

2 cm
Fig.12 –14: Lower dental
elements of Machairodus
KAUP, 1833 from the Turolian
of Ukraine.
Fig. 12: NNPM Ca-1274, isola-
ted left p4 of Machairodus
aphanistus (KAUP, 1832) from
Novaja Emetovka 2 (early MN
12) in a: lingual, b: occlusal,
and c: labial view. Note the
relatively narrow protoconid
and that an anterior basal
cusp and crenulation of the
postprotocristid are lacking.

Fig. 13: NNPM 48-7074, isola-

ted right p4 of Machairodus
giganteus (WAGNER, 1848)
from Belka (late MN 12) in a:
occlusal, b: labial, c: lingual
view. Note low posterior basal 12a.
cuspid and lacking crenula-
tions on the postprotocristid.
Fig. 14: NNPM Ca-1275, isola-
ted right m1 of Machairodus
aphanistus (KAUP, 1832) from
Novaja Emetovka 2 (early
MN 12) in a: labial, b: occlusal,
and c: lingual view. Note the
presence of a small metaconid 14a.
and absence of a lingual pre-

13a. 14b.



Fig. 15–16: Juvenile mandibles
of Machairodus giganteus
(WAGNER, 1848) from the
Turolian of Ukraine.

Fig. 15: NNPM 38-1134, right

juvenile mandible from
Novoukrainka 1 in a: lingual,
b: occlusal, and c: labial view.
d: erupting m1 in occlusal

Fig. 16: NNPM 45-26318, right

15a. juvenile mandible from
Cherevichnoe (MN 13) in
a: lingual, b: occlusal, and
c: labial view. d: erupting m1
in occlusal view.





137 Michael Morlo & Yuriy Semenov | Turolian Machairodus from Ukraine |


Fig. 17: Sizes of p4 of
Machairodus aphanistus (KAUP,
1832) ( ) and Machairodus
giganteus (WAGNER, 1848) ( ).
Ukrainian specimens are given 15,0
in bold. Other data are from
(1975, 1988), MARTÍNEZ (1996), 13,0
MORLO (1997), and
(2004). To allow inclusion of
damaged specimens, unknown
breadth is given as 6.1 mm, 10,0
unknown length as 20.1 mm.
Breadth [mm]
Table 1: Stratigraphic position
of Turolian localities with
remains of Machairodus in 7,0

20,0 22,0 24,0 26,0 28,0 30,0

Length [mm]

Land Mammal Age


M. aphanistus

M. giganteus



Upper 13 Cherevichnoe X

Middle 12b Belka X


Lower 12a Novaja Emetovka 2 X


Upper 11b Grebeniki, Novoukrainka 1 X


11a Krivoj Rog, Issaevo (MN10?) X

139 Michael Morlo & Yuriy Semenov | Turolian Machairodus from Ukraine |

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