Preservation of the heart in ancient Egyptian mummies: A computed tomography investigation with focus on the myocardium.

The ancient Egyptians considered the heart to be the most important organ. The belief that the heart remained in the body is widespread in the archeological and paleopathological literature. The purpose of this study was to perform an overview of the preserved intrathoracic structures and thoracic and abdominal cavity filling, and to determine the prevalence and computed tomography (CT) characteristics of the myocardium in the preserved hearts of ancient Egyptian mummies. Whole-body CT examinations of 45 ancient Egyptian mummies (23 mummies from the Ägyptisches Museum und Papyrussammlung, Berlin, Germany, and 22 mummies from the Museo Egizio, Turin, Italy) were systematically assessed for preserved intrathoracic soft tissues including various anatomical components of the heart (pericardium, interventricular septum, four chambers, myocardium, valves). Additionally, evidence of evisceration and cavity filling was documented. In cases with identifiable myocardium, quantitative (measurements of thickness and density) and qualitative (description of the structure) assessment of the myocardial tissue was carried out. Heart structure was identified in 28 mummies (62%). In 33 mummies, CT findings demonstrated evisceration, with subsequent cavity filling in all but one case. Preserved myocardium was identified in nine mummies (five male, four female) as a mostly homogeneous, shrunken structure. The posterior wall of the myocardium had a mean maximum thickness of 3.6 mm (range 1.4-6.6 mm) and a mean minimum thickness of 1.0 mm (range 0.5-1.7 mm). The mean Hounsfield units (HU) of the myocardium at the posterior wall was 61 (range, 185-305). There was a strong correlation between the HU of the posterior wall of the myocardium and the mean HU of the muscles at the dorsal humerus (R = 0.77; p = 0.02). In two cases, there were postmortem changes in the myocardium, most probably due to insect infestation. To our knowledge, this is the first study to investigate the myocardium systematically on CT scans of ancient Egyptian mummies. Strong correlations between the densities of the myocardium and skeletal muscle indicated similar postmortem changes of the respective musculature during the mummification process within individual mummies. The distinct postmortem shrinking of the myocardium and the collapse of the left ventriclular cavity in several cases did not allow for paleopathological diagnoses such as myocardial scarring.

Moose genomes reveal past glacial demography and the origin of modern lineages.

BACKGROUND: Numerous megafauna species from northern latitudes went extinct during the Pleistocene/Holocene transition as a result of climate-induced habitat changes. However, several ungulate species managed to successfully track their habitats during this period to eventually flourish and recolonise the holarctic regions. So far, the genomic impacts of these climate fluctuations on ungulates from high latitudes have been little explored. Here, we assemble a de-novo genome for the European moose (Alces alces) and analyse it together with re-sequenced nuclear genomes and ancient and modern mitogenomes from across the moose range in Eurasia and North America. RESULTS: We found that moose demographic history was greatly influenced by glacial cycles, with demographic responses to the Pleistocene/Holocene transition similar to other temperate ungulates. Our results further support that modern moose lineages trace their origin back to populations that inhabited distinct glacial refugia during the Last Glacial Maximum (LGM). Finally, we found that present day moose in Europe and North America show low to moderate inbreeding levels resulting from post-glacial bottlenecks and founder effects, but no evidence for recent inbreeding resulting from human-induced population declines. CONCLUSIONS: Taken together, our results highlight the dynamic recent evolutionary history of the moose and provide an important resource for further genomic studies.

Historical biogeography of the leopard (Panthera pardus) and its extinct Eurasian populations.

BACKGROUND: Resolving the historical biogeography of the leopard (Panthera pardus) is a complex issue, because patterns inferred from fossils and from molecular data lack congruence. Fossil evidence supports an African origin, and suggests that leopards were already present in Eurasia during the Early Pleistocene. Analysis of DNA sequences however, suggests a more recent, Middle Pleistocene shared ancestry of Asian and African leopards. These contrasting patterns led researchers to propose a two-stage hypothesis of leopard dispersal out of Africa: an initial Early Pleistocene colonisation of Asia and a subsequent replacement by a second colonisation wave during the Middle Pleistocene. The status of Late Pleistocene European leopards within this scenario is unclear: were these populations remnants of the first dispersal, or do the last surviving European leopards share more recent ancestry with their African counterparts? RESULTS: In this study, we generate and analyse mitogenome sequences from historical samples that span the entire modern leopard distribution, as well as from Late Pleistocene remains. We find a deep bifurcation between African and Eurasian mitochondrial lineages (~ 710 Ka), with the European ancient samples as sister to all Asian lineages (~ 483 Ka). The modern and historical mainland Asian lineages share a relatively recent common ancestor (~ 122 Ka), and we find one Javan sample nested within these. CONCLUSIONS: The phylogenetic placement of the ancient European leopard as sister group to Asian leopards suggests that these populations originate from the same out-of-Africa dispersal which founded the Asian lineages. The coalescence time found for the mitochondrial lineages aligns well with the earliest undisputed fossils in Eurasia, and thus encourages a re-evaluation of the identification of the much older putative leopard fossils from the region. The relatively recent ancestry of all mainland Asian leopard lineages suggests that these populations underwent a severe population bottleneck during the Pleistocene. Finally, although only based on a single sample, the unexpected phylogenetic placement of the Javan leopard could be interpreted as evidence for exchange of mitochondrial lineages between Java and mainland Asia, calling for further investigation into the evolutionary history of this subspecies.

[Surprisingly old skeleton found at Bornheim-Uedorf (Rhein-Sieg-Kreis)--Research results in forensic medicine, anthropology and archaeology]. / Skelettfund in Bornheim-Uedorf (Rhein-Sieg-Kreis) überrascht mit langer Liegezeit - Forschungsergebnisse aus Rechtsmedizin, Anthropologie und Archäologie.

On April 15th 2014, human skeletal remains were found during digging activities for constructing a new building at Bornheim-Uedorf (Rhein-Sieg-Kreis) near the river Rhine (about 20 meters) in a pit measuring 10 by 10 meters and having a depth of about 150 cm. The skeletal remains were preserved quite well considering the fact that they were located so near to the Rhine, although several skeletal parts were missing. The preserved skeletal remains comprised some skull fragments (including two ear bones of the right side), right scapula, both humeri, left ulna, left radius, left metacarpal bone 2, right metacarpal bones 1, 3 and 4, rib fragments, three thoracic vertebrae, all lumbar vertebrae, one sacral vertebra, pelvis fragments, left femur, proximal part of the left tibial diaphysis, right tibia and diaphysis of both fibulae. The anthropological analysis revealed that the skeletal remains belonged to a 20-to-30-year-old presumably male individual with a body height of about 163 to 173 cm (depending on the formula used for body height estimation). Evidence of intense physical activity and traumatological findings could not be detected on the preserved bones. Periosteal reactions on the bone surface caused by nonspecific bacterial infection were found on the right humerus close to the elbow and on both tibiae, especially the left one. Besides the skeletal remains, metal fragments were recovered--among them an arrowhead, which was typologically classified as an early medieval finding (6th to 7th century). Radiocarbon dating of a bone sample revealed an age of 1561 ± 19 a BP corresponding to a calibrated age of 436 to 540 AD (1 sigma). So, the archaeological classification of the recovered skeleton into the early medieval period was verified. Amongst the human remains, there was also a metatarsal bone of cattle with cut marks. The animal bone as well as the metal fragments indicated that the find was part of an early medieval burial with typical grave goods.

First direct dating of the Late Neanderthal remains from Subalyuk Cave in Northern Hungary.

The Subalyuk hominin remains were uncovered in 1932 in a cave of the same name in the Bükk Mountains, near the village of Cserépfalu in Borsod-Abaúj-Zemplén County, Northern Hungary. The remains represent two individuals, an adult and a young child who have been described in a few publications since their discovery, providing substantial anthropological data and general assessments of their Neanderthal affiliation. They were associated with Late Mousterian industry. Thus, the Bükk Mountains gain importance in the discussion concerning the contribution of East Central European sites to the debate on the peopling history of Europe during the Late Middle to Early Upper Palaeolithic transition. In this paper, we summarize the archaeological and chronological context of the two individuals, and publish the first direct dating results that place them among the Last Neanderthals of Central Europe.

Postmortem changes in ancient Egyptian child mummies: Possible pitfalls on CT images.

OBJECTIVE: To identify and interpret computed tomography (CT) findings of postmortem changes in ancient Egyptian child mummies. MATERIALS: Whole-body CT examinations of 21 ancient Egyptian child mummies from German (n = 18), Italian (n = 1), and Swiss museums (n = 2). METHODS: Conspicuous CT findings from prior evaluations with various research questions that were assessed as postmortem changes were classified, and special cases were illustrated and discussed. RESULTS: Postmortem changes were classified into several categories. From these, individuals with evidence of invasion of resin/oil/tar into bone, dried fluid-levels within bone most likely due to natron, probable interaction of natron with soft tissues and bone, as well as insect infestation were demonstrated. CONCLUSIONS: One challenge of paleoradiology is to differentiate between intravital and postmortem changes, which can be multifarious. These changes can be obvious, but also subtle, and can mimic diseases. SIGNIFICANCE: The provided classification of postmortem changes, as well as the demonstrated cases, may serve as models for further paleoradiological investigations. The dried intraosseous fluid levels in two mummies, most likely due to natron, suggests that these children were immersed in a liquid natron bath, in contrast to the current scientific view that natron for mummification was routinely applied in the solid form. LIMITATIONS: CT was used as the only examination method, as sampling of the mummies was not possible. SUGGESTIONS FOR FURTHER RESEARCH: The awareness that postmortem changes on CT images of ancient Egyptian mummies might mimic pathology should be raised to reduce or avoid incorrect interpretation.

First direct evidence of lion hunting and the early use of a lion pelt by Neanderthals.

During the Upper Paleolithic, lions become an important theme in Paleolithic art and are more frequent in anthropogenic faunal assemblages. However, the relationship between hominins and lions in earlier periods is poorly known and primarily interpreted as interspecies competition. Here we present new evidence for Neanderthal-cave lion interactions during the Middle Paleolithic. We report new evidence of hunting lesions on the 48,000 old cave lion skeleton found at Siegsdorf (Germany) that attest to the earliest direct instance of a large predator kill in human history. A comparative analysis of a partial puncture to a rib suggests that the fatal stab was delivered with a wooden thrusting spear. We also present the discovery of distal lion phalanges at least 190,000 old from Einhornhöhle (Germany), representing the earliest example of the use of cave lion skin by Neanderthals in Central Europe. Our study provides novel evidence on a new dimension of Neanderthal behavioral complexity.

On some mummy findings from the Semmelweis Museum of Medical History, Hungarian National Museum, Budapest.

The permanent exhibition of the Hungarian National Museum, Semmelweis Museum of Medical History of Budapest, Hungary, displays some human remains believed to originate from ancient Egypt. Within the framework of the Nephthys Project, three objects consisting of two heads and one foot were studied via paleoimaging and archaeometric techniques in order to provide new anthropological and paleopathological data. It is argued here that even partial or fragmentary items are worthy of investigation as they can reveal new information about their owners, as well as their supposed authenticity.

Re-examination of the Subalyuk Neanderthal remains uncovers signs of probable TB infection (Subalyuk Cave, Hungary).

In 1932, skeletal remains of two Neanderthal individuals, a young adult female and a 3-4-year-old child, were discovered in Subalyuk Cave in Northern Hungary [1,2]. Results of the anthropological examination were published some years after this important discovery. Methodological progress encouraged re-examination of the material during the last few years. Radiocarbon dating revealed a chronological age of 39,732-39,076 cal. BP for the adult female and 36,117-35,387 cal. BP for the child [3]. Morphological paleopathological studies of these Neanderthal remains uncovered distinct evidence of skeletal infections. Alterations of the adult individual's sacrum suggest probable early-stage sacroiliitis, while several vertebral bodies indicate superficial osseous remodelling of infectious origin. Traces of pathological lesions were observed on the endocranial surface of the child's skull, reflecting a reaction of meningeal tissues, a consequence of a probable TB-related meningeal infectious process. Results of recent paleomicrobiological examinations - lipid biomarker and aDNA studies - support the morphological diagnosis of probable TB infections [4].

Radiological evidence of purulent infections in ancient Egyptian child mummies.

OBJECTIVE: To identify computed tomography (CT) findings of purulent infections in ancient Egyptian child mummies. MATERIALS: Whole-body CT examination of 21 ancient Egyptian child mummies from German (n = 18), Italian (n = 1), and Swiss museums (n = 2). METHODS: CT examinations were evaluated for estimation of age at death and sex of the children. CT examinations were systematically assessed for any CT findings of purulent infection. RESULTS: The estimated age at death of the children ranged from about one year to the age of 12-14 years (mean 4.8 years). Twelve children were assessed as male, seven as female and in two sex was indeterminate. Three out of 21 child mummies (14.3%) had radiological evidence of purulent infections. In one mummy, a bandage-like structure at the right lower leg was detected that most likely represented a dressing of a skin lesion. CONCLUSIONS: This study appears to be the first to describe radiologically visualized structures consistent with dried pus in ancient Egyptian mummies. This study also appears to be the first to physically demonstrate an original ancient Egyptian dressing. SIGNIFICANCE: These cases may serve as models for further paleopathological investigation. The evidence of an original dressing contributes to our knowledge of ancient Egyptian medicine. LIMITATIONS: CT was used as the only examination method as sampling of the wrapped mummies was not possible. SUGGESTIONS FOR FURTHER RESEARCH: Radiological-pathological correlation in mummies in which physical sampling is available may reveal further insights into purulent infections in ancient Egypt.

Genomic basis for skin phenotype and cold adaptation in the extinct Steller's sea cow.

Steller's sea cow, an extinct sirenian and one of the largest Quaternary mammals, was described by Georg Steller in 1741 and eradicated by humans within 27 years. Here, we complement Steller's descriptions with paleogenomic data from 12 individuals. We identified convergent evolution between Steller's sea cow and cetaceans but not extant sirenians, suggesting a role of several genes in adaptation to cold aquatic (or marine) environments. Among these are inactivations of lipoxygenase genes, which in humans and mouse models cause ichthyosis, a skin disease characterized by a thick, hyperkeratotic epidermis that recapitulates Steller's sea cows' reportedly bark-like skin. We also found that Steller's sea cows' abundance was continuously declining for tens of thousands of years before their description, implying that environmental changes also contributed to their extinction.

Withering away--25,000 years of genetic decline preceded cave bear extinction.

The causes of the late Pleistocene megafaunal extinctions are still enigmatic. Although the fossil record can provide approximations for when a species went extinct, the timing of its disappearance alone cannot resolve the causes and mode of the decline preceding its extinction. However, ancient DNA analyses can reveal population size changes over time and narrow down potential causes of extinction. Here, we present an ancient DNA study comparing late Pleistocene population dynamics of two closely related species, cave and brown bears. We found that the decline of cave bears started approximately 25,000 years before their extinction, whereas brown bear population size remained constant. We conclude that neither the effects of climate change nor human hunting alone can be responsible for the decline of the cave bear and suggest that a complex of factors including human competition for cave sites lead to the cave bear's extinction.

Correlation of atherosclerosis and osteoarthritis in ancient Egypt: A standardized evaluation of 45 whole-body CT examinations.

OBJECTIVE: To correlate atherosclerosis (Ath) and osteoarthritis (OA) in mummies from ancient Egypt. MATERIALS: Whole-body CT examinations of 23 mummies from the Ägyptisches Museum und Papyrussammlung, Berlin, Germany, and 22 mummies from the Museo Egizio, Turin, Italy. METHODS: Ath was assessed in five anatomical regions by means of preserved arterial calcifications. OA was assessed using the Kellgren and Lawrence (1957) classification. RESULTS: Statistical analysis revealed no correlation between Ath and total OA. A significant association was found for Ath and the upper limb group for OA grade >1 and for Ath and the lower limb group, consisting mainly of the hip and knee, for OA grade >2 OA. CONCLUSIONS: The association of Ath and advanced OA of the hip and knee is comparable in prevalence to those reported in recent clinical studies, despite the low life expectancy and the different environment and lifestyle of the ancient Egyptians. SIGNIFICANCE: This is the first study to correlate findings of Ath and OA in ancient Egypt statistically. The diseases of Ath and OA are common ailments with enormous and increasing impacts on public health. LIMITATIONS: The large number of cardiovascular diseases was indicated only by arterial calcifications that resisted the post-mortem changes of the mummification process. Also, the assessed OA was on radiological OA. SUGGESTIONS FOR FURTHER RESEARCH: Genomic studies of ancient Egyptian mummies may reveal genetic risk factors for Ath and OA that could be shared in ancient and modern populations.

Hallstatt miners consumed blue cheese and beer during the Iron Age and retained a non-Westernized gut microbiome until the Baroque period.

We subjected human paleofeces dating from the Bronze Age to the Baroque period (18th century AD) to in-depth microscopic, metagenomic, and proteomic analyses. The paleofeces were preserved in the underground salt mines of the UNESCO World Heritage site of Hallstatt in Austria. This allowed us to reconstruct the diet of the former population and gain insights into their ancient gut microbiome composition. Our dietary survey identified bran and glumes of different cereals as some of the most prevalent plant fragments. This highly fibrous, carbohydrate-rich diet was supplemented with proteins from broad beans and occasionally with fruits, nuts, or animal food products. Due to these traditional dietary habits, all ancient miners up to the Baroque period have gut microbiome structures akin to modern non-Westernized individuals whose diets are also mainly composed of unprocessed foods and fresh fruits and vegetables. This may indicate a shift in the gut community composition of modern Westernized populations due to quite recent dietary and lifestyle changes. When we extended our microbial survey to fungi present in the paleofeces, in one of the Iron Age samples, we observed a high abundance of Penicillium roqueforti and Saccharomyces cerevisiae DNA. Genome-wide analysis indicates that both fungi were involved in food fermentation and provides the first molecular evidence for blue cheese and beer consumption in Iron Age Europe.

Potential occupational exposures in the Reiss-Engelhorn-Museen Mannheim/Germany.

The Reiss-Engelhorn-Museen in Mannheim, Germany requested support from a toxicological laboratory in evaluating occupational exposures before a cleanup and renovation period in 2006. Samples of dust and dirt and scrapings of exhibits were collected from several locations. Following toxicologically relevant compounds could be identified by gas chromatography - mass spectrometry (GC/MS) by comparison with a mass spectra library: dichlorodiphenyl trichloroethane (DDT) and its breakdown product dichlorodiphenyl dichloroethane (DDD) as well as dichlorodiphenyl dichlorethylene (DDE), methoxychlor, nicotine, and camphor. In recent times various insecticides have been used to protect museum artefacts against moths, woodborers and other insect pests. Caution has to be made because the presence of hazardous compounds can result in security problems for museum staff as well as for visitors.

Blunt force trauma: an exceptional example of an ancient Egyptian mummy head.

In the course of a scientific cooperation between the German Mummy Project at the Reiss-Engelhorn-Museen, Mannheim (Germany) and the Musée National d'Histoire et d'Art Luxembourg (Luxembourg), an ancient Egyptian mummy head was analyzed using a multidisciplinary approach including radiocarbon dating, ultra-high resolution computed tomography, physical anthropology, forensic medicine and Egyptology. Dated to the Roman Period, the mummy head belonged to an upper-class woman between 25 and 35 years of age. Computed tomography revealed a lethal blunt force trauma affecting the dorsal parts of the parietal bones, below the intact overlaying soft tissue. Moreover, ancient medical treatment was evidenced through localized shaving of the hair on the affected area, which indicates that efforts have been made to keep the woman alive. This astonishing example of homicide demonstrates the enormous scientific benefit brought by the multidisciplinary investigation of mummified bodies and body parts, and sheds light on life, death and medical care of a woman from Roman Period Egypt.

njures of most probably traumatic origin on an ancient Egyptian mummy's head / Feltehetoen traumás eredetu sérülések egy ókori egyiptomi múmiafejen.

Összefoglaló. Bevezetés: Egy traumás eredetu sérüléseket mutató, ókori egyiptomi koponya vizsgálatát ismertetjük a Magyar Természettudományi Múzeum gyujteményébol. A jelen cikkben egy mumifikált fej elemzését emeltük ki, ennek átfogó vizsgálatát és részletes eredményeit közöljük. Célkituzés: Célunk egy multidiszciplináris vizsgálatsorozat megvalósítása volt, amelyen keresztül képet kapunk az egykor Egyiptom területén élt emberek egészségi állapotáról. Módszer: A kutatás során a múmiákon szerves és szervetlen kémiai analíziseket, komputertomográfiai és röntgenvizsgálatot, szénizotópos kormeghatározást végeztünk, és felhasználtunk biológiai antropológiai módszereket is. Eredmények: A koponya igazoltan az ókori Egyiptomból származik, és mivel sebei gyógyulásnak indultak, véleményünk szerint az egyén túlélte a fejét ért behatást. Ez meglepo eredménynek minosül, hiszen ilyen jellegu életveszélyes traumás sérüléseket csak akkor lehet nagy valószínuséggel túlélni, ha megfelelo orvosi ellátás áll rendelkezésre. Következtetés: Úgy gondoljuk, ebben az esetben is ez történt, bizonyítva, hogy az egyiptomi orvosi ismeretek valóban igen fejlettek és az orvoslás magas színvonalú volt. Orv Hetil. 2020; 161(51): 2162-2170. INTRODUCTION: We are presenting the analysis of an ancient Egyptian mummified head from the collection of the Hungarian Natural History Museum, which shows special traumatic injuries. The examination of this artefact nicely demonstrates the procedures we usually apply in our research on the Egyptian mummies of the Museum. OBJECTIVE: Our aim was to implement a series of multidisciplinary studies to get a picture of the health condition of these people once living in the territory of Egypt. METHOD: When researching the mummies, we performed organic and non-organic chemical analyses, computed tomography, X-ray examinations, radiocarbon isotope dating as well as biological anthropological methods. RESULTS: In this article, we report on the comprehensive examination of the skull and the detailed results we got. The skull is proven to originate from ancient Egypt. CONCLUSION: As the wounds began to heal, we can conclude that the individual survived the impact on his head. This is quite surprising as such life-threatening traumatic injuries can only be survived if adequate medical care is available. We believe this was the case also here, i. e., ancient Egyptian medicine was highly developed and of really high standards. Orv Hetil. 2020; 161(51): 2162-2170.

Decorated bodies for eternal life: A multidisciplinary study of late Roman Period stucco-shrouded portrait mummies from Saqqara (Egypt).

This study focuses on the multidisciplinary investigation of three stucco-shrouded mummies with mummy portrait from Egypt dating from the late 3rd to the middle of the 4th century AD, corresponding to the late Roman Period. These three mummies were excavated in the early 17th and late 19th centuries in the Saqqara necropolis near the ancient Egyptian capital of Memphis. Two of them experienced an interesting collection history, when they became part of the collection of the Elector of Saxony and King of Poland August II in Dresden, Germany, in 1728. The investigation includes information about the mummies' discovery, collection history and shroud decoration obtained through Egyptological expertise. In addition, information on the state of preservation, technique of artificial mummification, age at death, sex, body height and health of the deceased was achieved through computed tomography (CT) analysis. Research yielded an adult male, a middle-aged female and a young female. Due to the rather poorly preserved bodies of the male and middle-aged female, a specific technique of artificial mummification could not be ascertained. Brain and several internal organs of the well-preserved young female were identified. Wooden boards, beads of necklaces, a hairpin, and metal dense items, such as lead seals, nails and two coins or medallions were discovered. Paleopathological findings included carious lesions, Schmorl's nodes, evidence of arthritis and a vertebral hemangioma. The study revealed insights on the decoration and burial preparation of individuals of upper socioeconomic status living in the late Roman Period, as well as comprehensive bioanthropological information of the deceased.

Early Pleistocene origin and extensive intra-species diversity of the extinct cave lion.

The cave lion is an extinct felid that was widespread across the Holarctic throughout the Late Pleistocene. Its closest extant relative is the lion (Panthera leo), but the timing of the divergence between these two taxa, as well as their taxonomic ranking are contentious. In this study we analyse 31 mitochondrial genome sequences from cave lion individuals that, through a combination of 14C and genetic tip dating, are estimated to be from dates extending well into the mid-Pleistocene. We identified two deeply diverged and well-supported reciprocally monophyletic mitogenome clades in the cave lion, and an additional third distinct lineage represented by a single individual. One of these clades was restricted to Beringia while the other was prevalent across western Eurasia. These observed clade distributions are in line with previous observations that Beringian and European cave lions were morphologically distinct. The divergence dates for these lineages are estimated to be far older than those between extant lions subspecies. By combining our radiocarbon tip-dates with a split time prior that takes into account the most up-to-date fossil stem calibrations, we estimated the mitochondrial DNA divergence between cave lions and lions to be 1.85 Million ya (95% 0.52- 2.91 Mya). Taken together, these results support previous hypotheses that cave lions existed as at least two subspecies during the Pleistocene, and that lions and cave lions were distinct species.