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Quaternaire, Hors-série, (4), 2011, p. 229-236
MASTERS OF THE LOST WORLD:
A HYPOTHETICAL LOOK AT THE TEMPORAL
AND SPATIAL DISTRIBUTION OF LION-LIKE FELIDS
n
Martin SABOL 1
ABSTRACT
A hypothetical scenario for the temporal and spatial distribution of extinct and extant lion-like felids is presented on the basis
of the latest data on their morphology, genetics, biogeography, and (palaeo-)ecology with the distinguishing of three to four possible
“phylogenetic lineages” (African, Asian, European, and American? one). The presence of two to three relative allopatric forms of
maneless lion-like felids in the northern Holarctic region during the Late Pleistocene (P. spelaea, P. vereshchagini, and P. atrox?) is
assumed as a result of parallelism within the pantherine cats.
Key-words: Lion-like felids, Pliocene, Pleistocene, Holocene, Distribution.
RÉSUMÉ
LES MAÎTRES D’UN MONDE PERDU : HYPOTHÈSES SUR LA RÉPARTITION SPATIO-TEMPORELLE DES «LION-LIKE
FELIDS» FOSSILES
Une hypothèse sur la répartition spatio-temporelle des «lion-like felids» fossiles et des lions actuels est présentée compte tenu
des données les plus récentes sur leur morphologie, leur génétique, la biogéographie, et la paléoécologie ou l’écologie, avec distinc-
tion entre trois ou quatre lignées phylogénétiques possibles (africaine, asiatique, européenne et américaine ?). La présence de deux
ou trois formes allopatriques de «lion-like felids» sans crinière dans la région nord holarctique au Pléistocène supérieur (P. spelaea,
P. vereshchagini, et P. atrox ?) pourrait résulter d’évolutions parallèles à l’intérieur du groupe phylétique des Panthera.
Mots-clés : «lion-like felids», Pliocène, Pléistocène, Holocène, répartition.
Manuscrit reçu le 23/09/2010, accepté le 29/03/2011
1 Department of Geology and Palaeontology, Faculty of Natural Sciences, Comenius University, Mlynská dolina, SK – 842 15 Bratislava,
Slovak Republic. Courriel: sabol@fns.uniba.sk
1 - INTRODUCTION
Present opinions on taxonomical position of fossil
lion-like felids are not unified. So far, they are most
often regarded either as extinct subspecies of Panthera
leo or they are assigned to some different fossil species,
respectively only to P. spelaea with some subspecies.
The main problem of fossil lion-like felids is the occur-
rence of relatively great regional differences that can
reflect an existence of several phylogenetic lineages. The
problem cannot be resolved only by using the morpho-
metric data and indices since they are often overlapping.
Besides statistic and genetic analyses, a research based
on knowledge of external variability of (pre)historic and
extant lion subspecies in the connection with (palaeo-)
biogeographic and (palaeo-)environmental study could
also yield interesting data. Based on that and the latest
published results of the research, a possible scenario for
the temporal and spatial distribution of lion-like felids
can be proposed with distinguishing of three to four
possible “phylogenetic lineages” (Fig. 1).
2 - TEMPORAL AND SPATIAL DISTRIBUTION
OF LION-LIKE FELIDS
2.1 - PLIO-PLEISTOCENE AFRICAN LION-LIKE
ANCESTORS
The Panthera genus, probably of Asian origin (Mazák,
2010), includes extant and extinct big pantherine felids
that common ancestor is known from the Late Pliocene
4-3 Ma (Turner and Antón, 1997). Probably during this
period, the first divergence of the genus took place and
the first representatives of lion-like felids occurred in
eastern Africa. Their earliest known fossils have been
found at Laetoli site in Tanzania, dated to around 3.5 Ma
(3.47 ± 0.12 Ma) (Barry, 1987). According to Hemmer
1112-052 Mep.indd 229 03/02/12 10:19
230
et al. (2001), they display characters of lion, leopard,
and jaguar, whereas Werdelin and Lewis (2005) attri-
buted these fossils to the Panthera specimen in size of
small lion. A fossil record of lion-like felids, older than
3 Ma, is also known from next East African site of Omo
Usno – Shungura C-F (Howell and Petter, 1976). In the
following period, the lion-like felids lived except eastern
Africa (Laetoli, Member G of the Shungura Sequence,
2.3 - 1.9 Ma; Turner, 1990) also in the south of the conti-
nent (Sterkfontein, Mb 4, 2.8 - 2.4 Ma; Turner, 1986).
About the same age range, a fossil record of lion-sized
Panthera is also known at the North African sites of Ahl
Al Oughlam (2.5 Ma; Geraads, 1997) and Tighenif (the
Earliest Pleistocene; Geraads, 2008), although it did
not has to belong to the P. leo lineage (Geraads, 2008).
All these Plio-Pleistocene large African cats (Panthera
shawi – Panthera aff. leo – Panthera nov. sp.) represent
probably smaller basal forms resembling a leopard by
their probably spotted body – a character that is preserved
in the shape of juvenile patches in modern lions and is
also known from some prehistoric and historic paintings
of adult specimens. They probably lived in the transi-
tional environmental type of peripheral primeval forest
and mosaic savannah (Lupták, 2009). We can consider
them (minimally eastern African ones) to be ancestors of
subsequent lion-like felids (Hemmer, 1974), the evolu-
tionary history of that started in eastern Africa in connec-
tion with the expansion of open habitats of woodland
and savannah type (the expansion of C4 plants – grasses)
during the Early Pleistocene 2 to 1.5 Ma (Yamaguchi et
al., 2004).
2.2 - AFRICAN LINEAGE
The earliest fossil record of lion representatives (within
the scope of the paper named as Panthera “leo”) is
known from the sites of Olduvai (layer I, 1.87 - 1.7 Ma)
in Tanzania (Petter, 1973), Koobi Fora (1.88 - 1.39 Ma)
(Werdelin and Lewis, 2005) and Turkana (Okote Mb.,
1.6 - 1.4 Ma) in Kenya (Leakey, 1976), Kromdraai A
(1.5 Ma) in South Africa (Turner, 1986; Lupták, 2009),
or from layers of Mb. L at Laetoli (Werdelin and Lewis,
2005). These assumed lineal descendants of the above-
mentioned Plio-Pleistocene basal forms were probably as
large as or somewhat larger than modern lions (Hemmer,
1974) and their males were even maneless (Yamaguchi
et al., 2004).
The evolution of Panthera leo and its extant subspe-
cies took place in Africa probably during the late Middle
Pleistocene. Based on the latest analysis of FIV virus
(Antunes et al., 2008), the extant lions are not descen-
dants of one panmictic population and evolved in some
Pleistocene refugiums in eastern (P. leo njarasensis?)
and southern Africa around 170 to 320 ky BP. These lions
are phylogenetically distinct from the Pleistocene lion-
like felids of the northern Holarctic region despite their
geographical proximity (Yamaguchi et al., 2004). They
also differ from earlier forms by the presence of male
mane (Janczewski et al., 1995) that represents a secon-
dary character evolved during the sexual selection, maybe
connected with the evolution of group-living reproducti-
vely active females (West and Packer, 2002). On the other
hand, the mane development was linked to disadvantage
in thermoregulation and restricted male activity (Nagel
et al., 2003).
Approximately 100 ky ago, the maned lions expanded
within the second migration wave to northern Africa
and Asia, where they gradually replaced Late Pleisto-
cene lion-like felids (Yamaguchi et al., 2004). After their
extinction, maned lions probably penetrated Europe about
6 to 8 ky ago (Yamaguchi et al., 2004; Stuart and Lister,
in press), usually assigned to Persian lion (Panthera leo
persica) (Gromov and Baranova, 1981). Based on the
archaeological record from Greece, Bulgaria, Moldova,
Hungary, and Ukraine dated to the period from the Late
Neolith to the Early Iron Age, however, Lupták (2009)
considers the extinct separate Holocene (historical) lion
subspecies (Panthera leo europaea). His assumption is
also based on historical illustration, mainly from both the
Minoan (2 600 - 1 425 BC) and the Mycenaean (1 570
- 1 125 BC) periods, where figured lions are similar to
Egyptian ones corresponding to African and Asian lions
in body size. Their specific characters are represented by
the conspicuous dorsal mane extending up to two thirds
of the back line and the elongated hair on the margin of
front legs or a sporadic hair hem along abdominal sides
(Lupták, 2009). Some figured lions with abdominal mane,
however, could indicate a presence of hybrids from Asia
Minor (Hemmer, 1967). The earliest written reference
on historical European lions comes from Homer, who
mentioned the “yellow lion” in his Iliad from 8th Century
BC (Lupták, 2009). In 480 BC, lions even attacked a
supplying caravan of Persian king Xerxes in Greece, but
according to Aristotle, they were already very rare in 330
BC and disappeared there about 100 BC (Schaller, 1980).
In the area to the east of Caucasus, the Holocene lions
(P. leo persica) survived up to 10th Century (Gromov and
Baranova, 1981). The Persian lions went extinct in Meso-
potamia in 1935, whereas they were even mentioned in
Iran in 1940s (Turner and Antón, 1997).
The recent distribution of lions, caused by both the
human activity and the impact of climatic changes, is
limited only to sub-Saharan Africa and one population in
northwestern India (Gir Forest, Kathiawar) (Yamaguchi
et al., 2004; Lupták, 2009). Within the species Panthera
leo, the historical and extant subspecies are divided to
three groups: persica group – P. leo persica (South Asian
lions), group close relatives to persica group – P. leo leo
(lions from Atlas Mountains), and senegalensis group –
P. leo senegalensis (West African lions), P. leo azandica
(lions from northeastern Congo), P. leo nubica (East
African lions), P. leo bleyenberghi (lions from southwes-
tern Africa), P. leo krugeri (lions from southeastern
Africa), and P. leo melanochaita (Cape lions) (Hemmer,
1974, 1978). The phylogenetic dividing of extant lions
to persica group and senegalensis group took place
probably during the period 74 - 203 ky ago (Burger et al.,
2004). At the Pleistocene-Holocene boundary, the second
lion exodus from southern refugium to eastern Africa
occurred, what caused the population inbreeding (e.g.
1112-052 Mep.indd 230 03/02/12 10:19
231
extant population of Serengeti lions represents a result of
interbreeding of three different recent populations). The
latest molecular research found 11 specific haplotypes
(M1 to M11) within single regional subpopulations of
P. leo, shedding a light on the taxonomy of the species.
The physical barriers (such as Sahara, Great Rift Valley,
or dense rain forests) played also one of major roles for
the haplotype divergence (Lupták, 2009).
2.3 - EUROPEAN LINEAGE
At the end of Early Pleistocene, probably still before
the Cromerian complex, the first migration wave of lion-
like felids from Africa (Panthera “leo”) to western part of
Asia (Near East, Central Asia?) took place. Subsequently,
they migrated from Asia as far as Europe at the beginning
of Middle Pleistocene (Panthera fossilis). Although the
earliest fossil records of these early European lion-like
felids, about one third larger than extant lions, comes
from the Italian site of Isernia la Pineta (0.7 Ma; Sala,
1990) and Pakefield in England (0.75 - 0.68 Ma, OIS
17-19; Lewis et al., 2010), their phylogenetic diver-
gence is marked by their dispersal over Europe in the
early Middle Pleistocene (Burger et al., 2004) during the
period after OIS 16. These evolutionary descendants of
African basal “lion” forms are assumed by some palaeon-
tologists to be only an earlier form of cave lion (Panthera
spelaea fossilis) specialized in open country hunt (e.
g. Argant, 1991; Baryshnikov and Boeskorov, 2001;
Barycka, 2008). During the Middle Pleistocene, they
spread over Europe and their fossil remains are mainly
known from sites in England (Kurtén, 1968; Jánossy,
1969), Spain (García, 2003), France (Bonifay, 1971;
Prat and Thibault, 1976; Argant, 1991, 2000), Germany
(Heller, 1949; Adam, 1959; Hemmer and Schütt, 1969;
Jánossy, 1969; Wolsan, 1993; Ambros et al., 2005),
Poland (Barycka, 2006, 2008), Moravia in Czech Repu-
blic (Musil, 1969; Thenius, 1972; Ábelová, 2005), and
Hungary (Jánossy, 1986; Wolsan, 1993). Lion fossils
from the Greek cave of Petralona, determined previously
as P. fossilis (Kurtén and Poulianos, 1977), are recently
attributed to P. spelaea (Baryshnikov and Tsoukala,
2010). The latter authors, however, mention lion fossils
from “old collection” (collected from the cave floor),
whereas Kurtén and Poulianos (1977) described lion
remains from “new collection”. They have been found
within the sedimentary groups A (top stalagmite) and C
(layer 11, 210 cm below the top stalagmite). The age of
the whole sedimentary filling of the Petralona Cave does
not exceed 800,000 years (Baryshnikov and Tsoukala,
2010) and its upper fossiliferous levels with pantherine
fossils are dated to ≤ 0.4 Ma (Kahlke et al., in press),
corresponding with age younger than OIS 11. Since the
fossil record from the site consists of mixed assemblages
of the Middle to Late Pleistocene, it is not excluded that
some lion fossils (minimally mandible fragment No. 136)
could be attributed to P. fossilis, as assumed by Kurtén
and Poulianos (1977).
The fossil record of lion-like felids from the Middle
Pleistocene sites situated in Eastern Europe and Caucasus
area, traditionally attributed to cave lions (Vereshchagin,
1971), is questionable. In the case of valid stratigraphy,
however, the fossil remains can belong either to P. fossilis
or to another representative of Middle Pleistocene lion-
like felids (see “2.4. Asian lineage”).
In spite of the fact that the relationship of the Middle
Pleistocene P. fossilis and the subsequent Late Pleisto-
cene cave lion is currently not clear because of the insuf-
ficient knowledge of the ancient form (Sotnikova and
Nikolskiy, 2003), it is not excluded that lion-like felids
from European plains and lowlands began to disperse
also in the Alps area at the end of the Middle Pleisto-
cene (or earlier?). Probably about this period (~ OIS 6?),
cave lion (Panthera spelaea) could evolve from one local
montane population. Subsequently, they dispersed in
mountainous areas of the entire Central European region
and penetrated territories of Western and Eastern Europe
during the Late Pleistocene. Findings of lion-like felids
from Austrian site of Repolusthöhle (late Middle Pleisto-
cene – Late Pleistocene; Döppes et al., 2008), mentioned
as Panthera leo cf. fossilis – P. cf. spelaea (Wolsan,
1993), can be an evidence of this hypothesis. These
fossils show rather certain differences from typical cave
lions, but probably they do not represent the remains of
P. fossilis relict population (Schütt and Hemmer, 1978).
On the other hand, the (south-)eastern immigration is
also not excluded since lion-like fossils, determined as P.
cf. spelaea, have been found in Middle Pleistocene depo-
sits of the Bear Cave in Israel (Tchernov and Tsoukala,
1997) and P. spelaea is also mentioned within the late
Middle Pleistocene mammalian assemblage of the Petra-
lona Cave in Greece (Baryshnikov & Tsoukala, 2010),
although exact species determination of the latter one
can be questioned because of the possible presence of
P. fossilis at the site (see above). Based on the record,
however, the hypothesis of the (Central-)European origin
of the species seems more verisimilar. During the Late
Pleistocene, the area of cave lion distribution extended
from Britannia and Pyrenees up to Ural and Caucasus
(Vereshchagin, 1971) with vertical distribution almost
up to 3,000m a.s.l. (Rabeder et al., 2000). The species
reached its optimum number in the second half of the
Last Glacial. After this period, however, the number of
cave lions gradually decreased up to the end of Weichse-
lian, when P. spelaea occurred mainly in south areas with
more temperate climate (Musil, 1986).
The main problem of cave lions (or Late Pleistocene
lion-like felids resp.) is the existence of relatively large
regional differences that theoretically allow dividing
them to several forms (subspecies?), mirroring a possible
existence of some phylogenetic lineages (Musil, 1996).
According to Krofel (2004), the body size of various
cave lion specimens could depend rather on food sources
than on abiotic factors and vegetation like in extant lions.
Schütt and Hemmer (1978) are thinking even about
genetically separate populations isolated by mountainous
ridges, and they distinguish the West European popula-
tion with narrow p4 and the East European one with wide
p4 (Hemmer, 1974). The situation is compared from
stratigraphical viewpoint almost to recent conditions in
1112-052 Mep.indd 231 03/02/12 10:19
232
the south of Africa, where two lion subspecies – P. leo
melanochaita and P. leo krugeri – persisted “side by
side”. Also skulls display two types – there are known
skulls with straight and convex profile. The latter ones
have shorter and wider nasalia, demonstrating a simila-
rity with skulls of lion-like felids from both the European
Middle Pleistocene and the Russian samples (Barycka,
2008). Yamaguchi et al. (2004) also assume the presence
of two morphologically distinguishable forms in Eastern
and Central Europe based on the ratio of breadth of the
postorbital constriction – interorbital breadth (Hemmer,
1974). According to the authors, one form was more
similar to North American and Siberian lion-like felids,
whereas another one resembled more extant North African
and Asian lions. This great variability could be caused by
the invasion of Siberian lion-like felids (Hemmer, 1974)
or by the invasion of extant lions from southwestern Asia
within the second migration wave (Yamaguchi et al.,
2004). A possibility of genetic interchange (Burger et al.,
2004), however, is questionable, relating with the taxono-
mical status of cave lion as a separate species (P. spelaea)
or only as a subspecies (P. leo spelaea) (Barnett et al.,
2009). Based on the phylogenetic analysis, the genetic
sequences of cave lions form a clade that is most closely
related to the extant lions from Africa and Asia (Barnett
et al., 2009). The cave lion sequences, however, represent
lineages that were isolated from lions in Africa and Asia
since their dispersal over Europe about 600 ky BP. These
lineages went extinct without mitochondrial descendants
and their clade is a sister to the extant lions in phyloge-
netic tree, what means that cave lions may be excluded
from or included within the species Panthera leo (Burger
et al., 2004). If cave lion was a separate species (Sotnikova
and Nikolskiy, 2006), however, the genetic interchange
between P. spelaea and other immigrating taxa of lion-
like felids was impossible. Also according to Christiansen
(2008), cave lion is not on the lion lineage, as traditio-
nally assumed, but represents a successive outgroup to
the lion + leopard. Although its position is only tentative,
it is supported by other lines of evidence such as brain
anatomy (Groiss, 1996), validating a common sharing of
plesiomorphic characters with tigers without the sharing
of common synapomorphic ones within the cranial struc-
ture (Sotnikova and Nikolskiy, 2006). This character
mosaic indicates a separate species status of P. spelaea
(Vereshchagin, 1971; Baryshnikov and Boeskorov, 2001;
Sotnikova and Nikolskiy, 2006). Moreover, it is not
excluded that relict populations of steppe lion-like felids
could also persist in lowlands of western, southern, and
eastern Europe (sporadically also in Central Europe,
mainly in the Pannonian area; Musil, 1986) during the
Late Pleistocene.
Before 48 ky BP, some cave lion populations started
to stagnate, undergoing major genetic alterations that
resulted to their extinction at the end of the Pleistocene.
In following period (after 46 - 48 ky BP), a rapid popula-
tion expansion in connection with a dramatic decline in
genetic diversity within cave lion populations is observed,
reflecting the genetic drift or genetic stagnation. The
stagnation could reflect major changes in the availabi-
lity of prey or in the ecology. The environmental changes
represent an alternative explanation since the stagnation
period corresponds to Heinrich event 5a, associated with
a cold stadial between Dansgaard-Oeschger interstadials
14 and 15 (Barnett et al., 2009), all within the OIS 3
– a period of major changes in the genetic structure of
Quaternary megafaunal populations of the Northern
Hemisphere before their end-Pleistocene mass extinc-
tion. Probably, the extinction of cave lion was caused by
a complex of adverse factors including genetic changes
in relation with environmental ones as a result of climatic
changes. One of the terminal data for cave lion in Europe
is at 12 375 ± 50 BP (14 378 cal BP) from Zigeuener-
fels, Sigmaringen in Germany (Stuart and Lister, 2007,
in press) or at 12 248 ± 66 BP (14 141 cal BP) from Le
Closeau in France (Stuart and Lister, in press), although
the species could survive in some European areas up to
the start of Holocene.
2.4 - ASIAN LINEAGE
Early forms of Asian lion-like felids can represent
hypothetical descendants of Early Pleistocene African
immigrants (P. “leo” ssp.), representing a result of
assumed more eastern migration(?). Exact data for the
validation or the disproving of this assumption, however,
are so far missing and it is only used within the proposed
hypothetical look at the temporal and spatial distribution
of lion-like pantherine cats. They dispersed into Indian
subcontinent (including Ceylon?) and Siberia probably
still before OIS 8 and also findings from Middle Pleis-
tocene sites situated eastwards from Ural Mountains
(Vereshchagin, 1971; Sotnikova, 2006) could belong
to specimens of the hypothetical one. According to
Yamaguchi et al. (2004), they probably did not spread
into south-eastern Asian montane rain forests, although
there is mentioned a regional form of large pantherine in
China (Panthera youngi) from the period 0.5 – 0.4 Ma,
considered to be a close relative to P. spelaea (Lupták,
2009).
The Middle Pleistocene Asian/Siberian lion-like felids
are probably ancestors of the Late Pleistocene ones
from Beringia (Panthera vereshchagini), originally
considered to be only a lesser Pleistocene subspecies
of cave lion (P. spelaea vereshchagini) (Baryshnikov
and Boeskorov, 2001). More recently detailed study of
ancient DNA (Barnett et al., 2009), however, does not
find any evidence for the differentiation of Beringian lion
populations from other spelaea ones (Stuart and Lister,
in press) in spite of certain differences from European
Late Pleistocene cave lions (Baryshnikov and Boeskorov,
2001). From this viewpoint, the taxonomical status of
Beringian “lion” remains open until more data on their
morphology and genetics will be available, although their
separate species status could be more probable from the
palaeobiogeographical and palaeoecological viewpoint.
According to Baryshnikov and Boeskorov (2001), they
were evolutionary descendants of P. fossilis, living in
conditions with low winter temperature extremes, a
lengthy duration of snow cover, and an elusive or growth-
1112-052 Mep.indd 232 03/02/12 10:19
233
limiting food base. These conditions caused the evolu-
tion of the economical form expending less energy to
maintain vital functions in connection with a decrease
in body size for the satisfaction of its hunger and the
maximization of its foraging efforts. To the contrary of
this opinion, however, the body size of P. vereshchagini
could be increased by the effect of low temperature like
in recent Siberian tigers, representing the largest subspe-
cies of Panthera tigris. Based on the radiometric dating of
fossils from NE Siberia (Lena and Arga-Yurekh Rivers)
and Alaska (Fairbanks Creek), the Beringian lion-like
felids went extinct probably in the period about 11 925 ±
70 BP (13 920 cal BP) to 12 450 ± 60 BP (14 640 cal BP),
or 12 525 ± 50 BP (14 797 cal BP) respectively (Barnett
et al., 2009; Stuart and Lister, in press), although later
survivals may yet be discovered in some areas (Stuart and
Lister, in press).
2.5 - AMERICAN LINEAGE
It is not excluded that next phylogenetic lineage of
big cats entered North America probably also during
the (early?) Middle Pleistocene or sooner, represen-
ting a possible ancestor of the biggest lion-like felids
(Panthera atrox). They remained isolated from panthe-
rine populations in other parts of the world (Yamaguchi
et al., 2004) probably already by Illinoian continental
glacier (Harington, 1969). New data estimates their
genetic isolation at around 340 ky BP and some analysis
place P. atrox inside the basal diversity of spelaea-form,
suggesting P. atrox was derived from a Beringian popu-
lation (Barnett et al., 2009). Found affinities with jaguars
(Simpson, 1941; Kurtén, 1965; Thenius, 1972), however,
can rule out this derivation (Christiansen and Harris,
2009) and refer to the relationship with ancestral branch,
from which big cats of the Old World evolved (Merriam
and Stock, 1932). The mainland form of North American
large felids was spread to the south of the continental
glacier southern boundary (Baryshnikov and Boeskorov,
2001) perhaps as far south as Peru (Yamaguchi et al.,
2004), except of eastern Canada and northeastern USA
because of dense forest (Lupták, 2009). Their distribution
was also related to climatic cycles and glacial/interglacial
environment (Yamaguchi et al., 2004). Ecological barriers
could cause the isolation of mainland North American
lion-like felids, although they could penetrate also the
non-glaciated territories in the area between Beringia and
central North America during warm oscillations. So far
an unknown intraspecific competition between atrox and
spelaea (vereshchagini) can also represent an alternative
explanation of their isolation since environmental and
ecological barriers did not operate over the entire period
of separation (Barnett et al., 2009). On the other hand,
provided that P. atrox is a different pantherine species
(Sotnikova and Nikolskiy, 2003, 2006; Christiansen,
2008; Christiansen and Harris, 2009), the isolation can
be more likely interpreted as a result of the interspecific
competition and not of the intraspecific one. The last
occurrence of P. atrox is mentioned at 11 355 ± 55 BP
from Edmonton in Canada (Barnett et al., 2009).
2.5 - PARALLELISM OF ALLOPATRIC FORMS
Within the Late Pleistocene lion-like felids from the
Northern Hemisphere area, two to three allopatric forms
can be distinguished – P. spelaea (western Europe to Ural
and Caucasus), P. vereshchagini (Siberia to Alaska and
Yukon), and P. atrox? (to the south of Wisconsinan glacier)
– as Baryshnikov and Boeskorov (2001) mentioned on
the basis of skull measurements and body size. Males of
P. spelaea and P. vereshchagini were probably maneless
since the cooler climate did not allow the development of
manes and a seasonal change of fur is rather assumed in
these animals (Nagel et al., 2003), similar to that in Sibe-
rian tigers. Also males of the North American mainland
pantherine cat (P. atrox) probably did not take pride in
mane because of the systematic position of the species
and its assumed different social system (Wheeler and
Jeferson, 2001; Christiansen and Harris, 2009). Based on
abovementioned, it is not excluded that these lion-like felid
taxa represent a phenomenon of the parallelism within
phylogenetic lineages of Northern Holarctic pantherine
cats and the resolving of their species status will depend
on results from both nuclear and morphological data.
3 - CONCLUSION
Based on the latest morphological, molecular, genetic,
biogeographic, and environmental analyses, a possible
scenario for the temporal and spatial distribution of
extant lions and their extinct close relative lion-like felids
can be as follows:
1) The earliest lion-like representatives of Panthera
evolved in eastern Africa during the Late Pliocene (3.5
- 2.5 Ma). They can be regarded as ancestors of later lion-
like felids, which probably spread up to southern and
(maybe) northern Africa;
2) The earliest fossils of Panthera “leo” have been
found in formations of eastern and southern Africa, dated
to the period 1.8 - 1.4 Ma;
3) At the end of Early Pleistocene, probably still before
the Cromerian complex, the first migration wave of
lion-like felids from Africa took place. They penetrated
western Asia, whence they spread up to Europe (Panthera
fossilis) about 0.7 Ma;
4) Hypothetical early forms of Asian lion-like felids
(Panthera “leo” ssp.) probably spread up to Indian
subcontinent (including also Ceylon?) and Siberia in
the period before OIS 8. About the same time, another
pantherine lineage probably entered the Americas and
gave rise to Panthera atrox in the late Middle Pleistocene
or in the early Late Pleistocene of North America, remai-
ning isolated from big cat populations in other parts of
the world;
5) During the Late Pleistocene, a descendant of the
Middle Pleistocene Siberian lion-like felids persisted in
Beringia (Panthera vereshchagini), but purportedly never
penetrated the American mainland because of presence
of the continental glacier between Alaska and the south
of North America;
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234
6) Lion-like felids from European lowlands began to
disperse in the Alps area probably during the late Middle
Pleistocene (or earlier?). Cave lion (Panthera spelaea)
evolved probably from one of these local populations of
ancient form about OIS 6. Subsequently, they dispersed
in mountainous areas of the entire Central European
region and during the Late Pleistocene they also pene-
trated territories of Western and Eastern Europe. It is also
not excluded that relict populations of steppe lion-like
felids (P. fossilis and/or P. “leo” ssp.?) could persist in
lowlands of western, southern, and eastern Europe during
the Late Pleistocene;
7) About 320 - 190 ky BP, maned felids (Panthera leo)
evolved in Africa, phylogenetically differing from Pleis-
tocene lion-like ones of the northern Holarctic region.
Approximately 100 ky ago, they expanded to northern
Africa and Eurasia, where gradually replaced original
maneless forms;
8) After the end-Pleistocene extinction of maneless
lion-like felids, probably Persian lions (Panthera leo
persica) penetrated Europe about 6 to 8 ky BP, although
there are some evidences for the existence of a separate
Holocene European lion subspecies (Panthera leo euro-
paea);
9) The recent distribution of lions is limited by both the
human activity and the impact of climatic changes (e.g.
in Saharan region) only to area of sub-Saharan Africa and
northwestern India.
It is also possible to distinguish several “phylogenetic
lineages” within the lion-like felids with the Early Pleis-
tocene African Panthera “leo” as a common ancestor
(fig. 1): 1. African lineage Panthera “leo” – Panthera leo
(including extinct and extant subspecies of maned lions);
2. Asian lineage Panthera “leo” – Panthera “leo” ssp. –
Panthera youngi / Panthera vereshchagini; 3. European
lineage Panthera “leo” – Panthera fossilis – Panthera
spelaea; and 4. questionable American lineage Panthera
sp. / Panthera “leo” ssp. – Panthera atrox. The existence
of two to three allopatric forms of relative lion-like felids
in the northern Holarctic region during the Late Pleis-
tocene (P. spelaea, P. vereshchagini, and P. atrox?) is
very interesting and it can be assumed also as a result of
parallelism within these pantherine cats.
ACKNOWLEDGEMENT
This research was supported by the Grant Agency for
Science, Slovakia (projects APVV-0280-07 and Vega
1/0176/10). The author also gratefully acknowledges the
helpful comments and suggestions of R. Ballesio and A.
Testu.
REFERENCES
ÁBELOVÁ M., 2005 - Doprovodná fauna veľkých cicavcov medvedej
jaskyne “Za Hájovnou”, Javoříčsky kras. Přírodovědné studie muzea
Prostějovska, 8, 171-184.
ADAM K.D., 1959 - Mittelpleistozäne Caniden aus dem Heppenloch
bei Gutenberg (Würtenberg). Stuttgarter Beiträge zur Naturkunde,
27, 1-46.
AMBROS D., HILPERT B., KAULICH B., REISCH L., & ROSEN-
DAHL W., 2005 - Steinberg-Höhlenruine bei Hunas (HFA A 236).
In: D. Ambros, Ch. Gropp, B. Hilpert, & B. Kaulich (eds.): Neue
Forschungen zum Höhlenbären in Europa, Abhandlung Band 45,
Naturhistorische Gesellschaft Nürnberg e.V., Nürnberg, 325-342.
ANTUNES A., TROYER, J.L., ROELKE M.E., PECON-SLAT-
TERY J., PACKER C., WINTERBACH CH., WINTERBACH
H., HEMSON G., FRANK L., STANDER PH., SIEFERT L.,
DRICIRU M., FUNSTON P.J., ALEXANDER K.A., PRAGER
K.C., MILLS G., WILDT D., BUSH M., O’BRIEN S.J., &
JOHNSON W.E., 2008. - The evolutionary dynamics of the Lion
Panthera leo revealed by host and viral population genomics. Plos
Genetics, 4 (11), 1-11.
ARGANT A., 1991 - Carnivores quaternaires de Bourgogne. Docu-
ments du Laboratoire de Géologie, 115, 304 p.
ARGANT, A., 2000 - Les sites paléontologiques du Pléistocène moyen
en Mâconnais. Bulletin de la Société Préhistorique Française, 97,
4, 609-623.
BARNETT R., SHAPIRO B., BARNES I., HO S.Y.W., BURGER J.,
YAMAGUCHI N., HIGHAM T.F.G., WHEELER H.T., ROSEN-
DAHL W., SHER A.V., SOTNIKOVA M., KUZNETSOVA T.,
BARYSHNIKOV G.F., MARTIN L.D., HARINGTON C.R.,
BURNS J.A., & COOPER A., 2009 - Phylogeography of lions
(Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene
reduction in genetic diversity. Molecular Ecology, 18, 1668-1677.
BARRY J.C., 1987 - The large carnivores from the Laetoli region of
Tanzania. In: M. D. Leakey & J. M. Harris (eds.): Laetoli, a Pliocene
site in northern Tanzania, Clarendon Press, Oxford, 48-52.
BARYCKA E., 2006 - Geographic and stratigraphic interpretations of
Panthera spelaea material from Polish sites – a new aspects. Scripta
Facultatis Scientarium Naturalium Universitatis Masarykianae
Brunensis, Geology, Vol. 33-34 (2003-2004), 14-15.
BARYCKA E., 2008 - Middle and Late Pleistocene Felidae and Hyae-
nidae of Poland. Fauna Poloniae – Fauna Polski 2ns, Museum and
Istitute of Zoology Polish Academy of Science, Warszawa, 228 p.
BARYSHNIKOV G.F. & BOESKOROV G., 2001 - The Pleistocene
cave lion, Panthera spelaea (Carnivora, Mammalia) from Yakutia,
Russia. Cranium, 18 (1), 7-23.
BARYSHNIKOV G.F., & TSOUKALA E., 2010 - New analysis of
the Pleistocene carnivores from Petralona Cave (Macedonia, Greece)
based on the Collection of the Thessaloniki Aristotle University.
Geobios, 43 (4), 389-402.
BONIFAY M.-F., 1971 - Carnivores Quarternaires du Sud-Est de la
France. Mémoires du Muséum National d´Histoire Naturelle, C,
XXXI (2), 377 p.
BURGER J., ROSENDAHL W., LOREILLE O., HEMMER H.,
ERIKSSON T., GÖTHERSTRÖM A., HILLER J., COLLINS
M.J., WESS T., & ALT K.W., 2004. - Molecular phylogeny of the
extinct cave lion Panthera leo spelaea. Molecular Phylogenetics and
Evolution, 30, 841-849.
CHRISTIANSEN P., 2008 - Phylogeny of the great cats (Felidae:
Pantherinae), and the influence of fossil taxa and missing charac-
ters. Cladistics, 24, 977-992.
CHRISTIANSEN P. & HARRIS J.M., 2009 - Craniomandibular
morphology and phylogenetic affinities of Panthera atrox: implica-
tions for the evolution and paleobiology of the lion lineage. Journal
of Vertebrate Paleontology, 29, 3, 934-945.
DÖPPES D., KEMPE S., & ROSENDAHL W., 2008 - Dated Paleon-
tological cave sites of Central Europe from Late Middle Pleistocene
to Upper Pleistocene (OIS 5 to OIS 8). Quaternary International,
187, 97-104.
GARCÍA N.G., 2003 - Osos y otros carnívoros de la Sierra de
Atapuerca. Fundacion Oso de Asturias, Asturia, 576 p.
GERAADS D., 1997 - Carnivores du Pliocène terminal de Ahl Al
Oughlam (Casablanca, Maroc). Geobios, 30, 1, 127-164.
GERAADS D., 2008 - Plio-Pleistocene Carnivora of northwestern
Africa: A short review. C. R. Palevol, 7, 591-599.
GROISS J.T., 1996 - Der Höhlentiger Panthera tigris spelaea
(GoldFuss). Neues Jahrbuch für Geologie and Paläontologie, Abh.
7, 399-414.
GROMOV I.M., & BARANOVA, G.I., 1981 - Katalog mlekopitajush-
chih SSSR. Pliocen - sovremennost. Nauka, Leningrad, 456 p.
HARINGTON C.R., 1969 - Pleistocene remains of the lion-like cat
(Panthera atrox) from the Yukon Territory and northern Alaska.
Canadian Journal of Earth Sciences, 6, 1277-1288.
HELLER F., 1949 - Beiträge zur Geologie und Paläontologie des
Tertiars und des Dilluviums in der Umgebung von Heidelberg. Ursus
(Plionarctos) stehlini Kretzoi, der kleine Bär aus den altdiluvialen
1112-052 Mep.indd 234 03/02/12 10:19
235
Fig. 1: Hypothetical look at the temporal and spatial distribution of single “phylogenetic lineages” of lion-like felids with maneless (probably spotted) Panthera “leo” as a common ancestor mentioned within the scope
of the paper. Stratigraphy is based on the Global Chronostratigraphical correlation table for the last 2.7 Ma (http://www.quaternary.stratigraphy.org.uk/charts); animal illustrations © Lupták & Csurma, 2009.
Fig. 1 : Hypothèses sur la distribution spatio-temporelle des lignées phylo-génétiques des “lion-like felids” sans crinière (et probablement tachetés) à partir de l’ancêtre commun Panthera “leo”. La chronostratigraphie est basée
sur “Global Chronostratigraphical correlation table” pour les 2,7 Ma précédents (http://www.quaternary.stratigraphy.org.uk/charts) ; illustrations © Lupták & Csurma, 2009.
1112-052 Mep.indd 235 03/02/12 10:19
236
Sanden von Mauer. Bammenthal und Mainz-Wiesbaden. Sitzungbe-
richte der Heidelberger Akademie der Wissenschaften, Abh. II.
HEMMER H., 1967 - Über das Aussehen der klein-bzw. Vorderasiatis-
chen Löwen, Panthera leo ssp. Säugetierkundl. Mitt., 15, 50-53.
HEMMER H., 1974 - Untersuchungen zur Stammesgeschichte der
Pantherkatze (Pantherinae), Teil III: Zur Artgeschichte des Löwen
Panthera (Panthera) leo (linnaeus, 1758). Ver öffentlichungen der
Zoologischen Staatssammlung München, 17, 167-280.
HEMMER H., 1978 - The evolutionary systematics of living Felidae:
present status and current problems. Carnivore, 1, 71-79.
HEMMER H., KAHLKE R.-D., & VEKUA A.K., 2001 - The jaguar –
Panthera onca gombaszoegensis (KreTzoi, 1938) (Carnivora: Felidae)
in the late Lower Pleistocene of Akhalkalaki (South Georgia; Trans-
caucasia) and its evolutionary and ecological significance. Geobios,
34, 475-486.
HEMMER H. & SCHÜTT G., 1969 - Ein Unterkiefer von Panthera
gombaszoegensis (Kretzoi, 1938) aus den Mosbacher Sanden. Mz.
Naturw. Arch., 8, 90-101.
HOWELL F.C., & PETTER G., 1976 - Carnivora from Omo Group
formations, Southern Ethiopia. In Y. Coppens, F. C. Howell, G. L.
Isaac & R. E. F. Leakey (eds.), Earliest Man and Environments in
the Lake Rudolph Basin. University of Chicago Press, Chicago,
314-331.
JANCZEWSKI D.W., MODI W.S., STEPHENS J.C., & O´BRIEN
S.J., 1995 - Molecular evolution of mitochondrial 12S RNA and
cytochrome b sequences in the pantherine lineage of Felidae. Mole-
cular Biology Evolution, 12 (4), 690-707.
JÁNOSSY D., 1969 - Stratigraphische Auswertung der europäischen
mittelpleistozänen Wirbeltierfauna. Teil I-II. Bereiches der Deuts-
chen Geschichtlichen geologichen Wissenschaften, A, Geologie und
Paläontologie., 14, 4-5, 367-438, 573-643.
JÁNOSSY D., 1986 - Pleistocene vertebrate fauna of Hungary. Else-
vier, Amsterdam – Oxford – New York – Tokyo, 208 p.
KAHLKE R.-D., GARCÍA N., KOSTOPOULOS D. S., LACOMBAT
F., LISTER A.M., MAZZA P.P.A., SPASSOV N., & TITOV,
V. , in press - Western palearctic palaeoenvironmental conditions
during the early and early Middle Pleistocene inferred from large
mammal communities, and implications for hominin dispersal in
Europe. Quaternary Science Reviews (2010), doi:10.1016/j.quas-
cirev.2010.07.020.
KROFEL M., 2004 - A find of Panthera leo spelaea Goldfuss (Felidae,
Mammalia) from unknown stratigraphic position of Potočka zijalka
(Slovenia). In: M. Pacher, V. Pohar & G. Rabeder (eds.): Potočka
Zijalka. Palaeontological and Archaeological Results of the
Campaigns 1997-2000. Mitteilungen der Kommission für Quar-
tärforschung der Österreichischen Akademie der Wissenschaften,
Band 13, 82-88.
KURTÉN B., 1965 - The Pleistocene Felidae of Florida. Bulletin of
Florida State Museum, 9 (6), 215-273.
KURTÉN, B., 1968 - Pleistocene Mammals of Europe. London, 317
p.
KURTÉN B., & POULIANOS, A.N., 1977 - New stratigraphic and
faunal material from Petralona Cave, with special reference to the
Carnivora. Anthropos, 4, 47-130.
LEAKEY M.G., 1976 - Carnivora of the east Rudolph succesion. In: Y.
Coppens, F. C. Howell, G. L. Isaac & R. E. F. Leakey (eds.): Earliest
Man and Environments in the Lake Rudolph Basin. University of
Chicago Press, Chicago, 302-313.
LEWIS M., PACHER M., & TURNER A., 2010 - The larger Carni-
vora of the West Runton Freshwater Bed. Quaternary International,
228, 116-135.
LUPTÁK P., 2009 - Externá variabilita a taxonómia súčasných a vyhy-
nutých poddruhov leva (Panthera leo). Gazella, 36, 33-150.
MAZÁK J.H., 2010 - What is Panthera palaeosinensis? Mammal
Revue, 40 (1), 90-102.
MERRIAM J.C., & STOCK CH., 1932 - The Felidae of Rancho La
Brea. Carnegie Institution of Washington Publications, 442, 1-227.
MUSIL R., 1969 - Eine karstspalte mit mittelpleistozänen Funden im
Kalksteinbruch Žernavá. Časopis moravského musea – Acta Musei
Moraviae, Vědy přírodní – Scientiae naturales, LIV, 85-96.
MUSIL R., 1986 - Paleobiography of terrestrial communities in Europe
during the Last Glacial. Sborník Národního muzea v Praze – Acta
Musei Nationalis Pragae, XLI B (1985), 1-2, 1-89.
MUSIL R., 1996 - Čertova pec a její fauna. Slovenský kras, XXXIV,
5-56.
NAGEL, D., HILSBERG S., BENESCH A., & SCHOLZ J., 2003
- Functional morphology and fur patterns in Recent and fossil
Panthera species. Scripta Geologica, 126, 227-241.
PETTER G., 1973 - Carnivores Pleistocènes du Ravin d´Olduvai
(Tanzanie). In: L. S. B. Leakey, R. J. G. Savage & S. C. Coryndon
(eds.): Fossil vertebrates of Africa 3, Academic Press, London,
43-100.
PRAT F. & THIBAULT C., 1976 - Le Gisement de Nauterie à la
Romieu (Gers) fouilles de 1967 a 1973. Nauterie I. Mémoires du
Muséum National d´Histoire Naturelle, Série C, Science de la terre,
t. XXXV, 96 p.
RABEDER G., NAGEL D., & PACHER M., 2000 - Der Höhlenbär.
Jan Thorbecke Verlag, Stuttgart, 111 p.
SALA B., 1990 - Panthera leo fossilis (V. Reich, 1906) (Felidae) de
Isernia La Pineta (Pléistocène Moyen Inférieur d´Italie). Geobios,
23, 2, 189-194.
SCHALLER G.B., 1980 - Zlaté stíny, pádící kopyta. Mladá fronta,
Praha, 272 p.
SCHÜTT G., & HEMMER H., 1978 - Zur Evolution des Löwen
(Panthera leo L.) im europäischen Pleistozän. Neues Jahrbuch für
Geologie und Paläontologie, Mt. 4, 228-255.
SIMPSON G.G., 1941 - Large Pleistocene Felines of North America.
American Museum Natural History Novitates, 1136, 1-27.
SOTNIKOVA M.V., 2006 - Pliocene-Early Pleistocene carnivore
assemblages of Transbaikalian Area, Russia. In: N. V. Alexeeva, M.
A. Erbajeva & A. G. Mironov (eds.): Stratigraphy, paleontology
and paleoenvironment of Pliocene-Pleistocene of Transbaikalia and
interregional correlations. Volume of Abstracts, INQUA, Ulan-Ude,
84-85.
SOTNIKOVA M., & NIKOLSKIY P., 2003 - Systematic position of
the cave lion (Panthera spelaea). 3rd International Mammoth Confe-
rence, Program and Abstracts, Occasional Papers in Earth Sciences,
No. 5, 142-143.
SOTNIKOVA M., & NIKOLSKIY, P., 2006 - Systematic position
of the cave lion Panthera spelaea (Goldfuss) based on cranial and
dental characters. Quaternary International, 142-143, 218-228.
STUART A.J., & LISTER A.M., 2007 - Patterns of Late Quaternary
megafaunal extinctions in Europe and northern Asia. Courier Fors-
chung-Institute Senckenberg, 259, 287-297.
STUART A.J., & LISTER A.M., in press - Extinction chronology of
the cave lion Panthera spelaea. Quaternary Science Reviews (2010).
doi:10.10.16/j.quascirev.2010.04.023.
TCHERNOV E., & TSOUKALA E., 1997 - Middle Pleistocene
(Early Toringian) carnivore remains from Northern Israel. Quater-
nary Research, 48, 122-136.
THENIUS E., 1972 - Die Feliden (Carnivora) aus dem Pleistozän von
Stránská skála. In: R. Musil (ed.): Stránská skála I 1910-1945, Studia
Musei Moraviae – Anthropos, 20 (N. S. 12), 121-135.
TURNER A., 1986 - Miscellaneous carnivore remains from Plio-Pleis-
tocene deposits in the Sterkfontein Valley (Mammalia: Carnivora).
Annals of the Transvaal Museum, 34, 203-226.
TURNER A., 1990 - Late Neogene/Lower Pleistocene Felidae of
Africa: evolution and dispersal. Quartärpaläontologie, 8, 247-256.
TURNER A., & ANTÓN M., 1997 - The Big Cats and their fossil rela-
tives. Columbia University Press, New York, 236 p.
VERESHCHAGIN N.K., 1971 - Peshcherniy lev i ego istoria
v Golarktike i v predelach SSSR. Trudy zoologičeskogo instituta, 49,
123-199.
WERDELIN L., & LEWIS M.E., 2005 - Plio-Pleistocene Carnivora
of eastern Africa: species richness and turnover patterns. Zoological
Journal of the Linnean Society, 144, 121-144.
WEST P.M., & PACKER C., 2002 - Sexual selection, temperature,
and the lion´s mane. Science, 297, 1339-1343.
WHEELER T. H. & JEFFERSON G. T., 2001 - Selection bias in
the predator collection from Rancho La Brea. Journal of Vertebrate
Paleontology, 21 (3, Supplement), 113A.
WOLSAN M., 1993 - Évolution des Carnivores Quaternaires en
Europe Centrale dans leur Contexte Stratigraphique et Paléoclima-
tique. L´Anthropologie, 97, 2/3, 203-222.
YAMAGUCHI N., COOPER A., WERDELIN L., & MACDO-
NALD D.W., 2004 - Evolution of the mane and groupliving in the
lion (Panthera leo): a review. Journal of the Zoological Society of
London, 263, 329-342.
1112-052 Mep.indd 236 03/02/12 10:19
