The phylogenetic status of Homo heidelbergensis – a cladistic study of Middle Pleistocene hominins

AbstractThe Ceprano human calvarium, dated around 400,000 yr, is a well-known fossil specimen. It represents significant evidence of hominin presence in the Italian peninsula during the Middle Pleistocene and may be considered representative of an archaic variant of the widespread and polymorphic species Homo heidelbergensis. Since its discovery (March 1994), systematic surveys in the Campogrande area near Ceprano, central Italy, identified 12 localities (CG1-12) with archaeological and/or paleontological assemblages. On this basis, fieldwork was carried out at Campogrande between 2001 and 2006, including drilled cores and excavations, allowing a detailed description of the stratigraphic and paleoenvironmental context associated with the human fossil specimen and the archaeological materials. In the present paper we focus on the stratigraphy and sedimentological features of the uppermost deposits, coupled with a detailed appraisal of the available lithic assemblages that mostly belongs to overlying sediments (CG9 and CG10 localities). We conclude that the Ceprano hominin died in a floodplain environment with a low topographic gradient, where a fluvial meandering channel occurred. The archaeological materials describe a network of sites that document common behavioural features of human groups of the mid-to-late Middle Pleistocene, representing evidence of the regionalization observed across Europe after Marine Oxygen Isotope Stage 12.

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New Middle Pleistocene hominin cranium from Gruta da Aroeira (Portugal)

Proceedings of the National Academy of Sciences ◽

10.1073/pnas.1619040114 ◽

2017 ◽

Vol 114 (13) ◽

pp. 3397-3402 ◽

Cited By ~ 38

Author(s):

Joan Daura ◽

Montserrat Sanz ◽

Juan Luis Arsuaga ◽

Dirk L. Hoffmann ◽

Rolf M. Quam ◽

...

Keyword(s):

Homo Erectus ◽

Middle Pleistocene ◽

Homo Heidelbergensis ◽

Incipient Stage ◽

Sima De Los Huesos ◽

Time Period ◽

Nasal Floor ◽

Fossil Hominins ◽

The Right ◽

Important Evidence

The Middle Pleistocene is a crucial time period for studying human evolution in Europe, because it marks the appearance of both fossil hominins ancestral to the later Neandertals and the Acheulean technology. Nevertheless, European sites containing well-dated human remains associated with an Acheulean toolkit remain scarce. The earliest European hominin crania associated with Acheulean handaxes are at the sites of Arago, Atapuerca Sima de los Huesos (SH), and Swanscombe, dating to 400–500 ka (Marine Isotope Stage 11–12). The Atapuerca (SH) fossils and the Swanscombe cranium belong to the Neandertal clade, whereas the Arago hominins have been attributed to an incipient stage of Neandertal evolution, toHomo heidelbergensis, or to a subspecies ofHomo erectus. A recently discovered cranium (Aroeira 3) from the Gruta da Aroeira (Almonda karst system, Portugal) dating to 390–436 ka provides important evidence on the earliest European Acheulean-bearing hominins. This cranium is represented by most of the right half of a calvarium (with the exception of the missing occipital bone) and a fragmentary right maxilla preserving part of the nasal floor and two fragmentary molars. The combination of traits in the Aroeira 3 cranium augments the previously documented diversity in the European Middle Pleistocene fossil record.

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The Sima de los Huesos (Burgos, northern Spain): palaeoenvironment and habitats of Homo heidelbergensis during the Middle Pleistocene

Quaternary Science Reviews ◽

10.1016/j.quascirev.2010.11.008 ◽

2011 ◽

Vol 30 (11-12) ◽

pp. 1413-1419 ◽

Cited By ~ 22

Author(s):

Nuria García ◽

Juan Luis Arsuaga

Keyword(s):

Middle Pleistocene ◽

Northern Spain ◽

Homo Heidelbergensis ◽

Sima De Los Huesos

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Humans of the Middle Pleistocene: The controversial calvarium from Ceprano (Italy) and its significance for the origin and variability of Homo heidelbergensis

Quaternary International ◽

10.1016/j.quaint.2015.12.047 ◽

2016 ◽

Vol 411 ◽

pp. 254-261 ◽

Cited By ~ 26

Author(s):

Giorgio Manzi

Keyword(s):

Middle Pleistocene ◽

Homo Heidelbergensis

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Crown outline analysis of the hominin upper third molar from the Megalopolis Basin, Peloponnese, Greece

Words, Bones, Genes, Tools: DFG Center for Advanced Studies - Ancient Connections in Eurasia ◽

10.51315/9783935751377.001 ◽

2021 ◽

pp. 13-36

Author(s):

Carolin Röding ◽

Julia Zastrow ◽

Heike Scherf ◽

Constantin Doukas ◽

Katerina Harvati

Keyword(s):

Homo Sapiens ◽

Third Molar ◽

Principal Component ◽

Middle Pleistocene ◽

Homo Heidelbergensis ◽

Human Fossil ◽

Geological Surveys ◽

Geological Age ◽

Outline Analysis ◽

Upper Third

The left upper third molar from the Megalopolis Basin is enigmatic due to its problematic preservation and context. The Megalopolis molar is the only possible human fossil known to date from the Megalopolis Basin. It was found on the surface during geological surveys in 1962-63. Based on the faunal assemblage collected during the same survey, it was proposed to be of Middle Pleistocene age and possibly one of the oldest human fossils in Europe. However, its actual geological age is unknown. In the past, dental crown outline analysis has been successfully used to differentiate between hominin species and populations. We applied the method to upper third molars, attempting to shed light on the affinities of the Megalopolis specimen. Principal component analysis (PCA) of the crown outline shape grouped the Megalopolis molar with our Homo sapiens sample; however, the PCA in form space, including shape plus size, as well as Procrustes distances based on overall shape, grouped it with our Neanderthal comparative sample. We conclude that its most likely identification is as a member of the Neanderthal lineage. However, we urge further analyses with an increased fossil comparative sample to include representatives of Homo heidelbergensis, which is underrepresented in our study. The Megalopolis molar contributes to the scarce Pleistocene human fossil record of Greece and highlights the potential of the Megalopolis Basin for yielding further paleoanthropological finds.

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Before the Emergence of Homo sapiens: Overview on the Early-to-Middle Pleistocene Fossil Record (with a Proposal about Homo heidelbergensis at the subspecific level)

International Journal of Evolutionary Biology ◽

10.4061/2011/582678 ◽

2011 ◽

Vol 2011 ◽

pp. 1-11 ◽

Cited By ~ 13

Author(s):

Giorgio Manzi

Keyword(s):

Homo Sapiens ◽

Early Pleistocene ◽

Middle Pleistocene ◽

Last Common Ancestor ◽

Homo Heidelbergensis ◽

Deep Roots ◽

Incipient Species ◽

And Diffusion ◽

Special Case ◽

Subspecific Level

The origin of H. sapiens has deep roots, which include two crucial nodes: (1) the emergence and diffusion of the last common ancestor of later Homo (in the Early Pleistocene) and (2) the tempo and mode of the appearance of distinct evolutionary lineages (in the Middle Pleistocene). The window between 1,000 and 500 thousand years before present appears of crucial importance, including the generation of a new and more encephalised kind of humanity, referred to by many authors as H. heidelbergensis. This species greatly diversified during the Middle Pleistocene up to the formation of new variants (i.e., incipient species) that, eventually, led to the allopatric speciation of H. neanderthalensis and H. sapiens. The special case furnished by the calvarium found near Ceprano (Italy), dated to 430–385 ka, offers the opportunity to investigate this matter from an original perspective. It is proposed to separate the hypodigm of a single, widespread, and polymorphic human taxon of the Middle Pleistocene into distinct subspecies (i.e., incipient species). The ancestral one should be H. heidelbergensis, including specimens such as Ceprano and the mandible from Mauer.

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