Middle Pleistocene Steppe Lion Remains from Grotte de la Carrière (Têt Valley, Eastern Pyrenees)

Abstract

Late Pleistocene cave lions are one of the most iconic species of Northern Hemisphere Quaternary taphocoenoses. Despite their often-scarce record in cave environments, their ubiquitous distribution across Eurasia and North America assemblages attests to their position as top ice-age predators. Nevertheless, the origins of these former large felids, their distribution during the Middle Pleistocene, and their paleoecology during co-existence with the scimitar-toothed cat Homotherium remain debated. Here we describe for the first time an abundant collection of large-sized and stout felid remains from the recently discovered site of Grotte de la Carrière in Eastern Pyrenees, with an estimated age corresponding to MIS 9. Our results highlight the larger size of Middle Pleistocene lions compared to Late Pleistocene ones as well as a trend of decreasing in size, which has been previously stated by other authors. Grotte de la Carrière steppe lions have similar morphological and biometrical parameters to those of other samples from MIS 11–9, being larger and stouter than younger latest Middle Pleistocene-Late Pleistocene forms and slightly smaller than older MIS 15–12 forms.

Early occurrence of lion (Panthera spelaea) at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy)

“…Note that the Notarchirico specimen is damaged and the true size would have been greater than the measured values. The raw data used in the comparison are from Marciszak et al (2021) and Prat‐Vericat et al (2022); see the supplementary dataset. [Color figure can be viewed at wileyonlinelibrary.com]…”

“…Pleistocene lions (or lion‐like felids) form a group close to the extant Panthera leo , but they had already diverged from it in the Early Pleistocene, being assigned to distinct species ( Panthera fossilis and Panthera spelaea ) or subspecies of the cave lion Panthera spelaea (the latter approach is followed herein), based mainly on craniodental metrics and morphology (Argant and Brugal, 2017; Prat‐Vericat et al, 2022; Sabol et al, 2022; Marciszak et al, 2023). Although the lion dispersal is often considered to be an important bioevent within the Galerian faunal turnover (Palombo et al, 2008), the species is poorly documented during the early Middle Pleistocene (Prat‐Vericat et al, 2022). Until now, the oldest published record from southwestern Europe was an isolated upper carnassial from Isernia La Pineta (Sala, 1990), dated at ~583–561 ka (at the end of the interglacial MIS 15; Peretto et al, 2015).…”

The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea

“…The cave lion species was not monotypic and included three chronologically successive subspecies. The oldest, largest and longest-existed was Panthera spelaea fossilis (von Reichenau, 1906) whose presence is documented in at least 62 Eurasian sites dated between 1200 and 300 kyr BP (David 1980(David , 1999Sala 1990;Lewis et al 2010;Hemmer 2011;Sotnikova and Foronova 2014;Marciszak et al 2019Marciszak et al , 2020aPrat-Vericat et al 2022). The second subspecies, on average smaller and less massive, P. s. intermedia Argant et Brugal, 2017, has been erected based on the material from Igue-des-Rameaux (France, 300-250 kyr BP; Argant and Brugal 2017;Brugal et al 2020;Persico 2021); morphologically similar lions with 'intermediate' features are known from various European sites dated to 300-180 kyr BP (Hemmer 1974(Hemmer , 2011Schütt and Hemmer 1978;Argant et al 2007;Argant 2010;Marciszak and Stefaniak 2010;Marciszak et al 2014Marciszak et al , 2020a.…”

“…The three subspecies of the cave lion differ in size and several morphological features. As compared to P. s. fossilis, P. s. spelaea possesses: larger incisors; more flattened and narrower canines; narrower P3 with shorter and higher paracone, more reduced protocone and metastyle and weaker distal cingulum; narrower P4 with shorter protocone, smaller and lower parastyle, and paracone shorter than metastyle; narrower p3 with shorter and lower protoconid; shorter and narrower p4 with longer and lower protoconid; more elongated and narrower m1, which is longer than p4 and has higher and longer protoconid and weaker or absent median lingual bulge (Kurtén 1960;Dietrich 1968;Hemmer and Schütt 1970;Ballesio 1975;Schütt and Hemmer 1978;Argant 1988Argant , 1991Groiss 1992Groiss , 2002Gužvica 1998;Baryshnikov and Boeskorov 2001;Hemmer 2003Hemmer , 2004Bona 2006;Sotnikova and Nikolskiy 2006;Argant et al 2007;Hankó 2007;Barycka 2008;Baryshnikov and Tsoukala 2010;Bona and Sardella 2012;Sabol 2014;Sotnikova and Foronova 2014;Marciszak et al 2014, Prat-Vericat et al 2022. According to Argant and Brugal (2017), P. s. intermedia is smaller than P. s. fossilis and has: less massive muscle attachments; a shorter and triangular-shaped mandibular ramus with shallower and shorter masseteric fossa; narrower canines; narrower P4 with almost straight buccal margins; narrower m1 with less developed median bulge; and more gracile metapodials and calcaneus.…”