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5 November 2025

New Records of Panthera gombaszoegensis (Kretzoi, 1938) from Europe

and
1
Department of Paleozoology, University of Wrocław, Sienkiewicza 21, 50-335 Wrocław, Poland
2
School of Geography, Queen Mary University of London, Mile End Road, London, E1 4NS, UK
*
Author to whom correspondence should be addressed.
Quaternary2025, 8(4), 65;https://doi.org/10.3390/quat8040065
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Abstract

New postcranial material of Panthera gombaszoegensis, a large pantherine felid, is analyzed from the English site of Corton (early Middle Pleistocene, possibly 0.7–0.6 mya) and the Polish site of Rogóżka Cave (0.45–0.35 mya). Both records are attributable to Panthera gombaszoegensis gombaszoegensis. This robust chronosubspecies is characteristic of the late Early and Middle Pleistocene, ca. last 1.5 mya. Both findings contribute valuable data on the knowledge of the species. The most likely factors that contributed to the extinction of P. gombaszoegensis were intraspecific competition with African newcomers, such as P. s. fossilis and C. crocuta, combined with climatic fluctuations and shifts in prey availability.

1. Introduction

Panthera gombaszoegensis was a characteristic faunal component of the Eurasian palaeocommunities by the last 2 mya [,,]. Until ca. 0.9 mya, when Panthera spelaea fossilis (von Reichenau, 1906) and Panthera pardus (Linnaeus, 1758) arrived, it was the only large pantherine felid present in Europe []. Its evolutionary success is evidenced by the enormous spatial distribution and the finding of its fossils in a wide range of habitats [,,,,,].
Remains of this felid are relatively abundant in the fossil record; however, many aspects of its evolutionary origin and taxonomic placement remain controversial. The earliest record of P. gombaszoegensis is controversial, despite general agreement of its African origin [,]. The oldest African fossil material is that from Kromdraai (1.9 mya) []; however, this remains controversial []. Considerably older African findings suggest that at least a jaguar-like felid existed already in Africa during the Late Pliocene. A fragment of maxilla from the South African site of Laetoli (3.5 mya) was attributed to P. aff. gombaszoegensis [] or Panthera leo (Linnaeus, 1758) []. However, later this specimen was described as a jaguar-like common stem species of pantherines [,]. It has also been noted that African pantherine specimens between 3.5 and 2.0 mya are not diagnostic at a species level. Specimens dated between 2.0 and 1.0 mya have historically been attributed to P. leo and Panthera pardus (Linnaeus, 1758) [], rather than to a jaguar-like felid. According to this parsimonious hypothesis, there is no direct link between the African specimens and the European P. gombaszoegensis [,].
Panthera gombaszoegensis is not known to have appeared in Eurasia before 2.0 mya, and most fossils have been known from southern and western Europe and S-W Asia []. The split of the jaguar lineage is estimated to have occurred between 2.6 and 2.0 mya, and the fossil records well align with the molecular data []. The species later dispersed into North America through Beringia 0.9–0.8 mya [,]. Three Pleistocene forms are commonly recognized and have been classified as independent chronosubspecies [,,,,,,], distinct species [,,,,,], or extinct subspecies of the extant Panthera onca (Linnaeus, 1758) [,,,]. Morphologically, P. gombaszoegensis is distinct from P. onca, and in this paper, these two taxa are regarded as separate species []. Additionally, analysis showed that the cranial features of P. gombaszoegensis are closer to those of the extant Panthera tigris (Linnaeus, 1758), while the mandibular morphology aligns more closely with P. onca []. A broader comparative overview showed that P. gombaszoegensis stays closer to the ancestral Asian pantherine lineage, now represented by P. tigris and Panthera uncia (Schreber, 1775), rather than to the tiger specifically [].
The oldest Panthera gombaszoegensis toscana (Schaub, 1949) is relatively small and gracile, with narrow P3 and P4, a broad, high, and short p3, a wide p4, and a narrow m1 without the median lingual bulge [,]. The occurrence of this chronosubspecies is temporarily restricted to the middle Early Pleistocene (2.0–1.6 mya) [,]. The large and robust Panthera gombaszoegensis gombaszoegensis (Kretzoi, 1938) possesses a wide P3–P4, an elongated, low, and narrow p3, a broad p4, and a wide m1, with a well-developed lingual median bulge [,,,]. Contemporarily to P. g. toscana, Asian chronosubspecies Panthera gombaszoegensis georgica (Hemmer et al., 2010) is a small jaguar with a narrow and high P3, a narrow P4 without a broadened heel-shelf and a prominent lingual cingulum edge, and a narrow m1 lacking lingual median bulge []. These features exhibit a high degree of intraspecific variability, making it difficult to determine precise chronological ranges for the various forms, especially those from Europe. Many European records, dated to 1.6–0.3 mya, are generally determined as P. g. gombaszoegensis or simply as P. gombaszoegensis [,,,,].
A review of the spatial and temporal distribution of P. gombaszoegensis documents its presence at more than 100 African and Eurasian localities, spanning a time range between 2.5 and 0.3 mya []. From Poland, its presence has been recorded from seven sites: Żabia Cave (1.7–1.5 mya), Kozi Grzbiet (0.8–0.7 mya), Południowa Cave (0.6–0.5 mya), Tunel Wielki Cave (0.6–0.5 mya), Draby 3 and 8 (0.45–0.35 mya), Biśnik Cave, layer 19ad (0.4–0.35 mya), and Komarowa Cave (0.4–0.3 mya). Three British records date to the Middle Pleistocene: Pakefield (0.75–0.7 mya), West Runton (0.7–0.6 mya), and Swanscombe (0.45–0.35 mya). Here, we describe additional postcranial material from England and Poland in an evolutionary context. In addition, palaeobiological aspects such as size trends and potential causes of extinction are briefly discussed.

2. Geographical and Geological Setting

Rogóżka Cave (50°14′39″ N 16°51′10″ E; 643 m a. s. l.; German name: Wolmsdorfer Tropfsteinhöhle) was located within a recently abandoned marble quarry (Figure 1). The 350 m long cave was discovered on 4.11.1885 during marble extraction and was filled with yellow-brown sandy loams. Excavations were first conducted in 1885 and 1894. Subsequently, in the 1920s, excavations were led by Arndt, who documented the discovery of U. spelaeus bones. Survey excavations were conducted by F. Pax in 1937, and by L. Zotz in 1935–1937, though no significant discoveries were recorded. After 1933, the destruction of the cave intensified due to quarry activity. Although the cave no longer exists, quarry workers claimed that a significant portion of the cave was not destroyed but rather lies behind a mound that obscures the entrance (Figure 2). Workers at the Heinrich und Lipst quarry regularly reported the dense accumulation of U. spelaeus bones from the cave as it was gradually destroyed. Due to the frequency of these reports, the State Office for the Protection of Prehistoric Monuments in Wrocław reported the discovery of bones in 1936, 1937, 1938, and 1942. Some of these specimens were deposited in Wrocław museums, where extensive documentation related to these finds is preserved. The faunal assemblage from Rogóżka Cave (MIS 10/9–1) consists of 43 species [,,,,].
Figure 1. Location of sites which yielded the analyzed Panthera gombaszoegensis remains: 1—Corton, 2—Rogóżka Cave. Scale bar 500 km.
Figure 2. Rogóżka Cave: (A)—the cave entrance to the cave and entrance shaft for tourists (1930s), (B)—cave plan, (C)—cave entrance, (D)—current state of preservation [,,].
Most of the fossil material was dated to the Late Pleistocene (last 130 kya). However, ancient forms such as Plecotus cf. rabederi Wołoszyn, 1987, Sorex runtonensis Hinton, 1911, and Macaca sp. from the deepest layers suggest an older age. Cave bear remains from the same layer as the jaguar material showed far less advanced morphology than those from the uppermost layers. Metrically and morphologically, these bones are related to Ursus deningeri hercynicus Rode, 1935, rather than U. spelaeus, suggesting a late Middle Pleistocene age (MIS 11–9).
Corton (52°31′42.5″ N 1°44′31.6″ E) is a coastal site in north Suffolk, England, which has early Middle Pleistocene exposures of the Cromer Forest-bed Formation [CF–bF]. The site is geologically significant as it represents the stratotype locality of the Anglian Glacial Stage []. The CF-bF sediments at Corton overlie sands of the Crag Group and underlie glaciofluvial sands and gravels associated with the Anglian Glaciation []. The two specimens of P. gombaszoegensis from the site were found within the silty clay deposits of the lower Rootlet Bed (Figure 3), which represents the basal level of the CF–bF at the site. The lower Rootlet Bed has been interpreted as a low-energy floodplain with standing water pools, surrounded by areas of temperate woodland, as indicated by pollen records []. These lower layers of the Rootlet Bed are often covered by beach sand and only exposed at low tide during scouring conditions. The two specimens were recovered approximately 100 m apart, and from slightly different sedimentary contexts within the lower Rootlet Bed. They also exhibit markedly different surface preservation, with MS 14 being significantly more pristine than MS 22, which shows substantial carnivore gnawing and gravel rolling. These differences in spatial distribution, sedimentary setting, and preservation suggest that the specimens likely do not belong to the same individual. The CF–bF deposits at Corton are dated to 0.7–0.6 mya; however, more precise dating is required to further correlate the site with nearby similarly aged localities such as Pakefield, West Runton, and Norton Subcourse. Pollen from Quercus, Tilia, and Ulmus was recorded from the Rootlet Bed at Corton. It was suggested that this was washed into the fluvial system from the temperate woodland that surrounded the floodplain [,]. This deciduous woodland likely made a suitable habitat and hunting ground for P. gombaszoegensis.
Figure 3. Exposed sediments of the Cromer Forest-bed Formation at Corton, 2023. The gray deposits exposed on the foreshore represent the lower Rootlet Bed. Remains of Panthera gombaszoegensis were found approximately 100 m apart within this layer. These deposits represent a low-energy freshwater environment, most likely a river floodplain.

3. Materials and Methods

The material from P. g. gombaszoegensis from Corton is stored in the Mark Spencer Collection (private collection) in Suffolk, England. The first bone is the right radius (MS 22) of an immature individual, with both proximal and distal epiphyses unfused. The second bone is a right metacarpal 5 (MS 14) with a slightly damaged distal epiphysis. The postcranial material from Rogóżka Cave is housed in the Department of Paleozoology, University of Wrocław, and consists of three bones: a right metatarsal 2 lacking the distal epiphysis (Rog. 21.1), a right metatarsal 4 also lacking the distal epiphysis (Rog. 21.2) and the third medial phalanx of the pes (Rog. 21.3) with a slightly damaged distal end. Measurements were taken point to point using an electronic caliper to the nearest 0.01 mm. Measuring schemes were adapted and modified from the literature (Figure 4, [,,]). Anatomical nomenclature is in accordance with the current International Committee on Veterinary Gross Anatomical Nomenclature (I.C.V.G.A.N.) []. Throughout the text, upper teeth are denoted using capital letters (e.g., P4), while lower teeth are indicated with lowercase letters (e.g., p4). The Marine Isotope Stage (MIS) boundaries follow the framework established in Lisiecki & Raymo [].
Figure 4. Scheme of measurements of Panthera gombaszoegensis bones: L—total length, pL—proximal epiphysis length, pB—proximal epiphysis breadth, mL—middle shaft length, mB—middle shaft breadth, dL—distal epiphysis length, dB—distal epiphysis breadth.

4. Results

Radius.
Description. The robust radius is cranio-caudally flattened and has a slightly concave caudal surface. The articulation surface for the scapholunate is broad and short (Figure 5).
Figure 5. Right radius (MS 22) of Panthera gombaszoegensis from Corton: (a)—lateral view, (b)—cranial view, (c)—medial view, (d)—caudal view, (e)—proximal view, (f)—distal view. Scale bar 40 mm.
Comparison. In comparison, the radius of Homotherium is slimmer in lateral view, with a straighter distal half, and a less pronounced median diaphysis enlargement. The radius of P. s. fossilis is significantly larger but less robust; its diaphysis is straighter and narrower, and both epiphyses are proportionally smaller in lateral view. P. pardus displays a smaller and narrower radius overall, with a similarly straight and narrow diaphysis, and proportionally smaller epiphyses. Additionally, all grooves and muscle attachments are less well developed in P. pardus.
Metacarpal 5.
Description. Metacarpal 5 from Corton is a large and massive bone with a strongly flattened and thickened proximal epiphysis (Figure 6). Laterally, the proximal epiphysis forms a semilunar convex surface. The intermetacarpal articulation is flattened and set at right angles to that of the hamatum. Both proximal articulation facets run gently from mesial to distal and protract medially. The medial articulation surface for the mc 4 shows two rounded areas: the larger, mesial area, extends distally into a triangular region. Externally, the head presents a large tuberosity, which provides attachment for strong ligaments that bind the bone to the hamatum, cuneiform, and pisiform. In addition, there is a large tuberosity on the palmar surface. The shaft is robust and short, triangular in cross-section, and arches more markedly medially than any other metacarpal bone. The distal epiphysis is strongly convex externally and concave internally. The lateral tuberosity is larger and lower than the medial one.
Figure 6. Postcranial material of Panthera gombaszoegensis: (1)—right metacarpal 5 from Corton (MS 14), (2)—right metatarsal 4 from Rogóżka Cave (Rog. 21.2), (3)—right metatarsal 2 from Rogóżka Cave (Rog. 21.1), (4)—right phalanx II-3-p from Rogóżka Cave (Rog. 21.3). Scale bar 20 mm, (a)—cranial view, (b)—lateral view, (c)—caudal view, (d)—medial view, (e)—proximal view.
Metatarsal 2.
Description. The metatarsal 2 from Rogóżka Cave (Rog. 21.1) has a massive and curved diaphysis, which is front-distally flattened and rectangular (Figure 6). Both (lateral and medial) articular surfaces of the proximal epiphysis are broad. The articular surface at the proximal end of the shaft is formed by a medial concavity that bulges anteroposteriorly. Posteriorly, the bone tapers significantly and exhibits a triangular shape. The distal articulation is irregularly shaped, large, and rounded. The medial epicondyle is more strongly developed than the lateral one.
Metatarsal 4.
Description. Metatarsal 4 from Rogóżka Cave (Rog. 21.2) has a prominent and rectangularly shaped outline of the proximal articulation facet of mt 5 (Figure 6). In the frontal view, the elongated and stout diaphysis widens strongly towards the proximal epiphysis. It exhibits a medial crista and a medial articulation facet that slopes medially. The articular surface for the cuboid bone is convex and frontally inclined. The lateral surface of the proximal epiphysis is rectangular, with a deep convexity located posterior to it. The articular surface is convex, inclined, and rectangular, and ends in the front in the form of a large, curved edge. On the cranial side, the proximal articular surface ends in a horizontal pattern. In the lateral view, the shaft is nearly straight and robust (Figure 6).
Comparison. The dimensions of all three described metapodials and single phalanx from Corton and Rogóżka Cave fit well, both metrically and morphologically, within the range of variability of P. gombaszoegensis (Table 1 and Table 2). Specimens from both sites belonged to relatively large and robust individuals. They are larger and noticeably more robust than those of P. pardus, but distinctly smaller and with proportionally larger epiphyses than those of P. s. fossilis. No substantial differences were observed between the metapodials from Corton and Rogóżka Cave and those from other P. gombaszoegensis populations (Table 1 and Table 2).
Table 1. Measurements of the postcranial material of Panthera gombaszoegensis gombaszoegensis (L-total length, pL-proximal epiphysis length, pB-proximal epiphysis breadth, mL-middle shaft length, mB-middle shaft breadth, dL-distal epiphysis length, dB-distal epiphysis breadth).
Table 2. Comparison of the sizes of metacarpal 5, metatarsals 2 and 4, and the third medial phalanx of the cats of the genus Panthera: Panthera gombaszoegensis from Corton and Rogóżka Cave, Panthera spelaea fossilis, extant Panthera leo, Panthera gombaszoegensis, and Panthera pardus (fossil and extant). For the measurements, the mean (bold) and ranges (range of variation as minimum and maximum value) are reported. Abbreviations: L—total length, pB—proximal epiphysis breadth, mB—middle shaft breadth, dB—distal epiphysis breadth. Data from own measurements and [,,,].
The differences in the examined material are significant enough to clearly exclude other forms of the genus Panthera. This is not to mention other large felids, such as Megantereon cultridens, Cuvier, 1824, Acinonyx pardinensis (Croizet & Jobert, 1828), or Homotherium latidens, Owen, 1846, sensu lato, whose metapodia are characterized by completely different morphology. Despite the significant degree of morphological unification of the metapodials and phalanges of cats from the genus Panthera, the described material from both sites, represented by bones of relatively large size and very massive structure, clearly allows them to be classified as belonging to P. gombaszoegensis. Metric differences often allow for precise identification, even in the case of isolated bone finds whose state of preservation is far from satisfactory. One such example is metatarsal 4 of P. s. fossilis from Notarchirico []. The surface of the bone is incompletely preserved, and the true size would have been greater than the obtained measurements. It was well documented in the bivariate comparison of the shaft diameters, in which the metatarsal 4 from Notarchirico plots among the smallest individuals. Contrary to that, the relatively better preserved distal end is larger and well above the range of extant P. leo [].

5. Discussion

Following a presence spanning more than 1.8 mya, the widespread and eurytopic P. gombaszoegensis had become extinct in Eurasia by around 0.3 mya. The disappearance of large carnivores, especially eurytopic species such as P. gombaszoegensis, would not have occurred rapidly, but would have been a long-spanning process because of multiple factors [,,,,]. Previously, P. gombaszoegensis was regarded as a forest dweller, closely associated with water []. This idea is strongly connected to the biology of the extant P. onca, which is also more flexible ecologically to varying habitats than previously thought. Before the decline under human pressure, P. onca also occupied open, dry pampas grasslands, as far as southern Patagonia []. Its Pleistocene relative, P. gombaszoegensis, was an even more eurytopic species, closer to P. pardus [,,,,,]. If P. gombaszoegensis was a typical forest dweller, the large-scale reduction in tree cover during the glacial may have contributed to its extinction, like Megantereon cultridens, Cuvier, 1824. However, P. gombaszoegensis records are more numerous from the Middle Pleistocene than from the Early Pleistocene, indicating that it not only survived but also increased in numbers [,,,,,].
It is highly probable that environmental changes did not play a primary role in the disappearance of P. gombaszoegensis, and there were likely other decisive factors. A broad prey spectrum was one of the key factors that contributed to its evolutionary success, enabling it to persist for a long time across dynamically changing and diverse habitats []. This dietary flexibility also contributed to the prolonged survival of P. gombaszoegensis despite pressure from African newcomers such as P. s. fossilis and C. crocuta ssp. Other highly specialized felids with narrower prey spectrums, such as A. pardinensis or M. cultridens, became extinct much earlier []. In this context, the availability of potential prey appears to have played a decisive role in the population dynamics of P. gombaszoegensis []. Despite its eventual reduction in range, this felid remained extremely successful compared to other large extinct forms such as saber-toothed cats []. The degree of dietary specificity was a key factor influencing the species’ long-term survival. P. gombaszoegensis was a generalist hypercarnivore, able to consume a wide prey spectrum [,,]. This versatility would have been advantageous to P. gombaszoegensis during periods when the availability of a large prey declined [,,].
Panthera gombaszoegensis was part of a stable carnivore palaeocommunity which remained unchanged at the species level for nearly 1 mya, between 2.0 and 1.0 mya [,,]. It held a high, though not dominant, position in the Early Pleistocene palaeohierarchy, and coexisted with multiple competitors, including saber-toothed cats, large canids, and hyenas [,,]. This stability was disrupted by the arrival of three newcomers of African origin, which occurred between 0.9 and 0.85 mya []. Particularly significant was the arrival of two powerful and social carnivores. The first, C. crocuta, is a highly intelligent, active hunter and aggressive scavenger, operating in large clans. At roughly the same time, P. s. fossilis appeared in Europe, a giant felid reaching a size of a big bear, with a body mass of up to 500 kg [,,]. Although the precise ecological impact of the lion on P. gombaszoegensis remains unclear, it was likely significant. The arrival of both species had a significant impact on the previous stability of the carnivore palaeoguild.
The relatively small body size of early leopards may have been an adaptive response to reduce competition with other large carnivores, particularly P. gombaszoegensis, where these two felids coexisted [,,]. Leopards are well adapted to living alongside other large sympatric carnivores, such as lions, tigers, wolves, and hyenas. Their shy behavior, like avoiding areas frequented by larger carnivores, targeting smaller prey, hunting at different times of day or night, and taking their carcasses high up trees, may reduce direct competition []. Being metrically intermediate between Panthera leo (Linnaeus, 1758) and P. pardus, P. gombaszoegensis occupied a distinct ecological niche, which reduced interspecific pressure. After the extinction of P. gombaszoegensis, P. pardus became widespread in Europe, increasing in size and appearing to ecologically replace the extinct jaguar []. The arrival of P. s. fossilis particularly disrupted the previously stable European felid paleoguild. A strong decline of P. gombaszoegensis is inferred in regions where lions found favorable habitat conditions [].
Human pressure may have also been a factor, resulting in the decline of P. gombaszoegensis. Hominins, likely including Homo heidelbergensis and possibly early Homo neanderthalensis, had larger body sizes and greater encephalization compared to previous members of Homo, which would have resulted in higher energetic demands. The consequence was an increase in the consumption of high-protein meat from large-bodied carcasses, such as ungulates, obtained from regular hunting and scavenging. This shift resulted in a new ecological dynamic in which competition between hominins and carnivores, such as the jaguar, was increased, which may have played a considerable role in the eventual extinction of P. gombaszoegensis [,,].

6. Conclusions

Panthera gombaszoegensis was a long-lived and successful felid, and one of the dominant carnivores during the Early Pleistocene. Its broad ecological flexibility and generalist feeding strategy gave it an advantage over more specialized carnivores. Since its arrival in Europe ~2 mya, P. gombaszoegensis coexisted with a relatively stable carnivore paleoguild, which remained unchanged at the species level for almost 1 mya. Currently, the spatial and temporal occurrence of P. gombaszoegensis includes more than 100 African and Eurasian localities, ranging between 2.5 and 0.3 mya. Among them are three British sites, including Pakefield (0.75–0.68 mya), West Runton (0.7–0.6 mya), and Swanscombe (0.42–0.37 mya), and seven Polish localities, including Żabia Cave (1.7–1.5 mya), Kozi Grzbiet (0.8–0.7 mya), Południowa Cave (0.6–0.5 mya), Tunel Wielki Cave (0.6–0.5 mya), Draby 3 and 8 (0.45–0.35 mya), Biśnik Cave, layer 19ad (0.4–0.35 mya), and Komarowa Cave (0.4–0.3 mya). In this paper, we describe two new sites, Corton, England (0.7–0.6 mya), and Rogóżka Cave, Poland (0.45–0.4 mya), broadening our knowledge of the species’ distribution. Intraspecific competition with African newcomers such as P. s. fossilis and C. Crocuta, combined with climatic fluctuations and shifts in prey availability, were likely factors that contributed to the extinction of P. gombaszoegensis.

Author Contributions

Conceptualization, methodology, software, and funding acquisition, A.M.; investigation, A.M. and A.B.; writing—original draft preparation, A.M. and A.B.; visualization, A.M. and A.B.; writing—review and editing, A.M. and A.B. All authors have read and agreed to the published version of the manuscript.

Funding

The research was financed by a grant from the University of Wrocław entitled “The Middle Pleistocene Revolution—how the modern theriofauna of Eurasia was developed”, grant no. BPIDUB.4610.6.2021.KP.A.

Data Availability Statement

All information where readers can obtain the research data required to reproduce the work is provided within the manuscript. All material, if currently present, is available to study in museums or private collections.

Acknowledgments

The authors are grateful to Mark Spencer for providing access to the Corton material [MS 14 and MS 22] from his private collection. A.B. is grateful to Simon Parfitt and Adrian Lister for helpful initial discussions on the Corton material. We are very grateful to A. Iannucci and two anonymous reviewers for their insightful comments and remarks which significantly improved the manuscript.

Conflicts of Interest

The authors declare no conflicts of interest.

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Climate During the Last Glacial Maximum in the Southern Sangre de Cristo Mountains, Colorado, U.S.A.
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