''Panthera fossilis'' (also known as ''Panthera leo fossilis'' or ''Panthera spelaea fossilis''), is an extinct species of

cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...

belonging to the

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

Panthera ''Panthera'' is a genus within the family (biology), family Felidae that was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group. Reginald Innes Pocock revised the classification of this genus in 1916 as co ...

,'' known from remains found in Eurasia spanning the

Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. The ...

and possibly into the

Early Pleistocene The Early Pleistocene is an unofficial sub-epoch in the international geologic timescale in chronostratigraphy, being the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently estimated to span the time ...

. ''P. fossilis'' has sometimes been referred to by the

common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contrast ...

s steppe lion or cave lion, though these names are conventionally restricted to the later related species '' P. spelaea,'' to which ''P. fossilis'' is probably ancestral.

Discoveries

It was first described from remains excavated near

Mauer Mauer is the German word for ''wall''. It may also refer to: Places *Mauer, Vienna, a former village of Lower Austria that has been part of Vienna since 1938 * Mauer bei Amstetten, a village in the municipality of Amstetten, in Lower Austria * Ma ...

Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...

. Bone fragments of ''P. fossilis'' were also excavated near

Pakefield Pakefield is a suburb of the town of Lowestoft in the north of the English county of Suffolk. It is located around south of the centre of the town. In 1931 the parish had a population of 1774. Pakefield has boundaries with Carlton Colville and ...

in the

United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...

, which are estimated at 680,000 years old. Bone fragments excavated near

Isernia Isernia () or, in Pliny and later writers, ''Eserninus'', or in the Antonine Itinerary, ''Serni''. is a town and ''comune'' in the southern Italian region of Molise, and the capital of province of Isernia. Geography Situated on a rocky crest ri ...

Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...

are estimated at between 600,000 and 620,000 years old. The first Asian record of a ''fossilis'' lion was found in the

Kuznetsk Basin The Kuznetsk Basin (russian: Кузнецкий угольный бассейн, Кузбасс; often abbreviated as Kuzbass or Kuzbas) in southwestern Siberia, Russia, is one of the largest coal mining areas in Russia, covering an area of aroun ...

in western

Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part of ...

and dates to the late

Evolution

''P. fossilis'' is estimated to have evolved in

Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...

about 600,000 years ago from a large pantherine cat that originated in the

Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...

Olduvai Gorge The Olduvai Gorge or Oldupai Gorge in Tanzania is one of the most important paleoanthropology, paleoanthropological localities in the world; the many sites exposed by the gorge have proven invaluable in furthering understanding of early human ev ...

about 1.2–1.7 million years ago. This cat entered Eurasia about 780,000–700,000 years ago and gave rise to several lion-like forms. The first fossils that can be definitively classified as ''P. fossilis'' date to 610,000 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.

Characteristics

Bone fragments of ''P. fossilis'' indicate that it was larger than the modern lion and was among the largest cats. Skeletal remains of ''P. fossilis'' populations in Siberia measure larger than those in

Central Europe Central Europe is an area of Europe between Western Europe and Eastern Europe, based on a common historical, social and cultural identity. The Thirty Years' War (1618–1648) between Catholicism and Protestantism significantly shaped the area' ...

. Compared to a modern lion, ''P. fossilis'' had a slightly wider skull and

nasal cavity The nasal cavity is a large, air-filled space above and behind the nose in the middle of the face. The nasal septum divides the cavity into two cavities, also known as fossae. Each cavity is the continuation of one of the two nostrils. The nasal c ...

, smaller

orbit In celestial mechanics, an orbit 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 object or position in space such as a p ...

s, less inflated bullae, less specialized lower teeth, reduced lower

premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...

s and smaller

incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...

Taxonomic history

''P. fossilis'' was historically considered an early

lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'' native to Africa and India. It has a muscular, broad-chested body; short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphi ...

(''P. leo'')

subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...

as ''Panthera leo fossilis''. Some authors considered it a subspecies of ''

Panthera spelaea ''Panthera spelaea'', also known as the Eurasian cave lion, European cave lion or steppe lion, is an extinct ''Panthera'' species that most likely evolved in Europe after the third Cromerian interglacial stage, less than 600,000 years ago. Phylo ...

'' (''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 anagenesis, 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 populatio ...

lineage, with most differences between the two species explainable by size differences. Results of

mitochondrial genome Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...

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 72 degrees north latitude in the Chukchi Sea; and on the south by the tip ...

n specimens of ''Panthera spelaea'' indicate that it and ''Panthera fossilis'' were distinct enough from the modern lion to be considered separate

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

Palaeobiology

This lion coexisted with

early humans ''Homo'' () is the genus that emerged in the (otherwise extinct) genus ''Australopithecus'' that encompasses the extant species ''Homo sapiens'' (modern humans), plus several extinct species classified as either ancestral to or closely related ...

and

prehistoric Prehistory, also known as pre-literary history, is the period of human history between the use of the first stone tools by hominins 3.3 million years ago and the beginning of recorded history with the invention of writing systems. The use of ...

fauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. Zoo ...

. A mandible from the early hominid ''

Homo heidelbergensis ''Homo heidelbergensis'' (also ''H. sapiens heidelbergensis''), sometimes called Heidelbergs, is an extinct species or subspecies of archaic human which existed during the Middle Pleistocene. It was subsumed as a subspecies of ''H. erectus'' in ...

'' was excavated in 1907 at Mauer, Germany.Schoetensack, O. (1908). Der Unterkiefer des ''Homo heidelbergensis'' aus den Sanden von Mauer bei Heidelberg. Ein Beitrag zur Paläontologie des Menschen. Leipzig: Engelmann.

Herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

s that coexisted with the lion included the

hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...

narrow-nosed rhinoceros The narrow-nosed rhinoceros (''Stephanorhinus hemitoechus'') is an extinct species of rhinoceros that lived in western Eurasia and North Africa during the Pleistocene. It first appeared in Europe some 600,000 years ago, and survived there until ...

straight-tusked elephant The straight-tusked elephant (''Palaeoloxodon antiquus'') is an extinct species of elephant that inhabited Europe and Western Asia during the Middle and Late Pleistocene (781,000–30,000 years before present). Recovered individuals have reac ...

southern mammoth ''Mammuthus meridionalis'', or the southern mammoth, is an extinct species of mammoth native to Europe and Central Asia from the Gelasian stage of the Early Pleistocene, living from 2.5–0.8 mya. Taxonomy The taxonomy of extinct elephant ...

moose The moose (in North America) or elk (in Eurasia) (''Alces alces'') is a member of the New World deer subfamily and is the only species in the genus ''Alces''. It is the largest and heaviest extant species in the deer family. Most adult mal ...

steppe bison The steppe bisonSeveral literatures address the species as ''primeval bison''. or steppe wisent (''Bison'' ''priscus'')
– Y ...

and

fallow deer ''Dama'' is a genus of deer in the subfamily Cervinae, commonly referred to as fallow deer. Name The name fallow is derived from the deer's pale brown colour. The Latin word ''dāma'' or ''damma'', used for roe deer, gazelles, and antelopes, ...

Sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...

predators Predation is a biological interaction where 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 the ...

included

bear Bears are carnivoran mammals of the family Ursidae. They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats throughout the Nor ...

wolves The wolf (''Canis lupus''; plural, : wolves), also known as the gray wolf or grey wolf, is a large Canis, canine native to Eurasia and North America. More than thirty subspecies of Canis lupus, subspecies of ''Canis lupus'' have been reco ...

hyena Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the clas ...

s and

saber-toothed cats Sabretooth or sabertooth may refer to: Animals * Saber-toothed cat, several prehistoric felines ** ''Smilodon'', a prehistoric genus of felidae * Sabertooth fish, a deep-sea fish found in the tropics ** Sabre-toothed blenny, ''Aspidontus taenia ...

References