Panthera Fossilis
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''Panthera fossilis'' (also known as ''Panthera leo fossilis'' or ''Panthera spelaea fossilis'') is an extinct species of
cat The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the ...
belonging to the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''
Panthera ''Panthera'' is a genus within the family (biology), family Felidae, and one of two extant genera in the subfamily Pantherinae. It contains the largest living members of the cat family. There are five living species: the jaguar, leopard, lion, ...
,'' known from remains found in
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
spanning the
Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...
and possibly into the
Early Pleistocene The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
. Although often historically considered a subspecies of the living
lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'', native to Sub-Saharan Africa and India. It has a muscular, broad-chested body (biology), body; a short, rounded head; round ears; and a dark, hairy tuft at the ...
(''Panthera leo''), ''Panthera fossilis'' is currently considered either a distinct species to be ancestral to or a chronosubspecies of ''
Panthera spelaea ''Panthera spelaea'', commonly known as the cave lion (or less commonly as the steppe lion), is an extinct ''Panthera'' species that was native to Eurasia and northwest North America during the Pleistocene epoch. Genetic analysis of ancient DNA ...
'' (commonly known as the cave lion or steppe lion). In comparison to Late Pleistocene ''Panthera spelaea'' specimens, ''Panthera fossilis'' tends to be considerably larger, up to , considerably exceeding modern lions in size, and making them among the largest cats to have ever lived, along with the South American sabertooth '' Smilodon populator''.


Taxonomic history

The species was first described by Wilhelm von Reichenau in 1906 from remains excavated near Mauer in
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
. ''P. fossilis'' was historically considered a modern
lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'', native to Sub-Saharan Africa and India. It has a muscular, broad-chested body (biology), body; a short, rounded head; round ears; and a dark, hairy tuft at the ...
(''P. leo'')
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
as ''Panthera leo fossilis''. Some authors considered it a subspecies of ''
Panthera spelaea ''Panthera spelaea'', commonly known as the cave lion (or less commonly as the steppe lion), is an extinct ''Panthera'' species that was native to Eurasia and northwest North America during the Pleistocene epoch. Genetic analysis of ancient DNA ...
'' (''Panthera spelaea fossilis'') or treat it as a distinct species. Some employ a subgenus of ''Panthera'', "''Leo''", to contain several lion-like members of ''Panthera'', including ''P. leo'', ''P. spelaea'', '' P. atrox'' and ''P. fossilis''. A 2022 study concluded that ''P. fossilis'' and ''P. spelaea'' represented a
chronospecies A chronospecies is a species derived from a sequential development pattern that involves continual and uniform changes from an extinct ancestral form on an evolutionary scale. The sequence of alterations eventually produces a population that is p ...
lineage, with most differences between the two species explainable by size differences. Results of
mitochondrial genome Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the DNA contained in ...
sequences derived from two
Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 70th parallel north, 72° north latitude in the Chukchi Sea; and on the south ...
n specimens of ''Panthera spelaea'' indicate that it and ''Panthera fossilis'' were distinct enough from the modern lion to be considered separate
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
.


Evolution

The lion lineage is thought to have originated in Africa, with fossils of the lineage on the continent extending back to the
Late Pliocene Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...
, with the ancestors of ''Panthera fossilis'' migrating out of Africa during the Early-Middle Pleistocene transition around 1-0.8 million years ago. The earliest confirmed records of ''Panthera fossilis'' in Europe are from Kozi Grzbiet in Poland, which dates to around 750-700,000 years ago, with records of equivalent age also known from
Pakefield Pakefield is a suburb of the town of Lowestoft, in the East Suffolk district of Suffolk, England. It is located around south of the centre of the town. It 2011 the ward had a population of 6,563. Pakefield has boundaries with Carlton Colvill ...
in England. The earliest confirmed records in Southern Europe are from the Notarchirico site in Italy, dating to ~660–612,000 years ago. Possible earlier European records are known from the Vallparadís locality in Spain, suggested to date to around 1 million years ago.including supplemental material Remains of ''Panthera fossilis'' in
Western Siberia Western Siberia or West Siberia ( rus, Западная Сибирь, p=ˈzapədnəjə sʲɪˈbʲirʲ; , ) is a region in North Asia. It is part of the wider region of Siberia that is mostly located in the Russia, Russian Federation, with a Sout ...
are suggested to date to the late Early Pleistocene, around 1 million years ago. Recent nuclear genomic evidence suggest that interbreeding between modern lions and all Eurasian fossil lions took place up until 500,000 years ago, but by 470,000 years ago, no subsequent interbreeding between the two lineages occurred. The arrival of ''Panthera (spelaea) fossilis'' in Europe was part of a
faunal turnover The turnover-pulse hypothesis, formulated by paleontologist Elisabeth Vrba, suggests that major changes to the climate or ecosystem often result in a period of rapid extinction and high turnover of new species (a "pulse") across multiple different ...
event around the Early-Middle Pleistocene transition in which many of the species that characterised the preceding late
Villafranchian Villafranchian age ( ) is a period of geologic time (3.5–1.0 Ma) spanning the Late Pliocene and Early Pleistocene used more specifically with European Land Mammal Ages. Named by Italian geologist Lorenzo Pareto for a sequence of terrestrial se ...
became extinct. In the carnivore guild, this notably included the giant hyena ''
Pachycrocuta ''Pachycrocuta'' is an extinct genus of hyena. The largest and most well-researched species is ''Pachycrocuta brevirostris'', colloquially known as the giant short-faced hyena as it stood about at the shoulder and it is estimated to have average ...
'' and the sabertooth cat ''
Megantereon ''Megantereon'' is an extinct genus of prehistoric machairodontine saber-toothed cat that lived in Eurasia, Africa and possibly North America from the late Pliocene to the Middle Pleistocene. It is a member of the tribe Smilodontini, and close ...
''. Following the arrival of ''Panthera (spelaea) fossilis'', the lion-sized sabertooth cat ''
Homotherium ''Homotherium'' is an extinct genus of Homotherini, scimitar-toothed cat belonging to the extinct subfamily Machairodontinae that inhabited North America, Eurasia, and Africa, as well as possibly South America during the Pliocene and Pleistocene ...
'' and the "European jaguar" ''
Panthera gombaszoegensis ''Panthera gombaszoegensis'', also known as the European jaguar, is a ''Panthera'' species that lived from about 2.0 to 0.3 million years ago in Europe, as well as likely elsewhere in Eurasia. The first fossils were Excavation (archaeology), exca ...
'' became much rarer, ultimately becoming extinct in the late Middle Pleistocene, with competition with lions suggested to be a likely important factor. The arrival of ''Panthera fossilis'' into Europe was also accompanied by that of the cave hyena (close relatives of the modern
spotted hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus ''Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the IUC ...
), as well as
leopards The leopard (''Panthera pardus'') is one of the five extant cat species in the genus ''Panthera''. It has a pale yellowish to dark golden fur with dark spots grouped in rosettes. Its body is slender and muscular reaching a length of with a ...
. Between 300,000 and 100,000 years ago, ''Panthera fossilis'' evolved into the cave lion ''(Panthera spelaea'')'','' marked by a reduction in body size and a number of changes to its skeletal anatomy (with intermediates between the two dubbed ''Panthera spelaea intermedia'').


Description

Remains of ''P. fossilis'' indicate that it was larger than the modern lion and was among the largest known cats ever, with the largest specimens suggested to have a body length of , shoulder height of and body mass of . Skeletal remains of ''P. fossilis'' populations in Siberia measure larger than those in
Central Europe Central Europe is a geographical region of Europe between Eastern Europe, Eastern, Southern Europe, Southern, Western Europe, Western and Northern Europe, Northern Europe. Central Europe is known for its cultural diversity; however, countries in ...
. Compared to its descendant ''Panthera spelaea''. ''P. fossilis'' had a slightly wider muzzle and nasal region of the skull, though the postorbital and mastoid are narrower, the
orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
s (eye sockets) are smaller, the bullae are less inflated, the canine teeth are more narrow and less flattened, the incisors are smaller, the upper second premolar and upper fourth premolar are narrower, the upper and lower third premolar and lower fourth premolars have smaller cusps. Another notable difference is that the front part of the upper skull surface (the frontal-nasal region) is typically concave, while this is less frequent in ''Panthera spelaea.'' The differences between the skulls of ''P. fossilis'' and ''P. spelaea'' have been described as relatively subtle.'''' In comparison to the modern lion, in addition to being larger, the upper second premolar of ''P. fossilis'' is larger, and the cusp morphology of the upper fourth premolar is different (having a shorter
metastyle Many different terms have been proposed for features of the tooth crown in mammals. The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn i ...
).


Ecology

During the Middle Pleistocene, ''Panthera fossilis'' was the dominant
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
in European ecosystems, likely able to displace every other contemporaneous predator species from kills/carcasses.
Herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s that coexisted with the lion included the
hippopotamus The hippopotamus (''Hippopotamus amphibius;'' ; : hippopotamuses), often shortened to hippo (: hippos), further qualified as the common hippopotamus, Nile hippopotamus and river hippopotamus, is a large semiaquatic mammal native to sub-Sahar ...
, rhinoceroses of the genus ''
Stephanorhinus ''Stephanorhinus'' is an extinct genus of two-horned rhinoceros native to Eurasia and North Africa that lived during the Late Pliocene to Late Pleistocene. Species of ''Stephanorhinus'' were the predominant and often only species of rhinoceros in ...
'' (such as
Merck's rhinoceros ''Stephanorhinus kirchbergensis'', also known as Merck's rhinoceros (or the less commonly, the forest rhinoceros) is an extinct species of rhinoceros belonging to the genus ''Stephanorhinus'' that lived from the end of the Early Pleistocene (arou ...
and the
narrow-nosed rhinoceros The narrow-nosed rhinoceros (''Stephanorhinus hemitoechus''), also known as the steppe rhinoceros is an extinct species of rhinoceros belonging to the genus '' Stephanorhinus'' that lived in western Eurasia, including Europe, and West Asia, as ...
),
straight-tusked elephant The straight-tusked elephant (''Palaeoloxodon antiquus'') is an extinct species of elephant that inhabited Europe and Western Asia during the Middle Pleistocene, Middle and Late Pleistocene. One of the largest known elephant species, mature full ...
,
moose The moose (: 'moose'; used in North America) or elk (: 'elk' or 'elks'; used in Eurasia) (''Alces alces'') is the world's tallest, largest and heaviest extant species of deer and the only species in the genus ''Alces''. It is also the tal ...
,
steppe bison The steppe bison (''Bison'' ''priscus'', also less commonly known as the steppe wisent and the primeval bison) is an extinct species of bison which lived from the Middle Pleistocene to the Holocene. During the Late Pleistocene, it was widely dist ...
,
red deer The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or Hart (deer), hart, and a female is called a doe or hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Ir ...
, roe deer and
fallow deer Fallow deer is the common name for species of deer in the genus ''Dama'' of subfamily Cervinae. There are two living species, the European fallow deer (''Dama dama''), native to Europe and Anatolia, and the Persian fallow deer (''Dama mesopotamic ...
.
Sympatric In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...
predators Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
included
brown bear The brown bear (''Ursus arctos'') is a large bear native to Eurasia and North America. Of the land carnivorans, it is rivaled in size only by its closest relative, the polar bear, which is much less variable in size and slightly bigger on av ...
s,
wolves The wolf (''Canis lupus''; : wolves), also known as the grey wolf or gray wolf, is a canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, including the dog and dingo, though gr ...
, cave hyenas, the large sabertooth cat ''
Homotherium ''Homotherium'' is an extinct genus of Homotherini, scimitar-toothed cat belonging to the extinct subfamily Machairodontinae that inhabited North America, Eurasia, and Africa, as well as possibly South America during the Pliocene and Pleistocene ...
,'' European leopards, and the "European jaguar" ''
Panthera gombaszoegensis ''Panthera gombaszoegensis'', also known as the European jaguar, is a ''Panthera'' species that lived from about 2.0 to 0.3 million years ago in Europe, as well as likely elsewhere in Eurasia. The first fossils were Excavation (archaeology), exca ...
''.


Relationship with humans

The only evidence of human interaction with ''Panthera fossilis'' is from Gran Dolina, Spain, dating to
Marine Isotope Stage Marine isotope stages (MIS), marine oxygen-isotope stages, or oxygen isotope stages (OIS), are alternating warm and cool periods in the Earth's paleoclimate, deduced from Oxygen isotope ratio cycle, oxygen isotope data derived from deep sea core ...
9 (~300,000 years ago), where a specimen of ''Panthera fossilis'' displays cut marks thought to be produced by
archaic humans ''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively calle ...
, who are suggested to have butchered the animal for its flesh.


See also

* History of lions in Europe * '' Panthera leo sinhaleyus'' * '' Panthera pardus spelaea''


References

{{Taxonbar, from=Q511072 fossilis fossilis Pleistocene mammals of Europe Pleistocene carnivorans Pleistocene species extinctions Fossil taxa described in 1906 History of lions in Europe