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. 2024 Mar;8(3):564-577.
doi: 10.1038/s41559-023-02303-6. Epub 2024 Jan 31.

The ecology, subsistence and diet of ~45,000-year-old Homo sapiens at Ilsenhöhle in Ranis, Germany

Geoff M Smith  1   2 , Karen Ruebens  3   4 , Elena Irene Zavala  5   6 , Virginie Sinet-Mathiot  3   7 , Helen Fewlass  3   8 , Sarah Pederzani  3   9 , Klervia Jaouen  3   10 , Dorothea Mylopotamitaki  3   4 , Kate Britton  11 , Hélène Rougier  12 , Mareike Stahlschmidt  3   13   14 , Matthias Meyer  6 , Harald Meller  15 , Holger Dietl  15 , Jörg Orschiedt  15 , Johannes Krause  16 , Tim Schüler  17 , Shannon P McPherron  18 , Marcel Weiss  3   19 , Jean-Jacques Hublin  3   4 , Frido Welker  20
Affiliations

The ecology, subsistence and diet of ~45,000-year-old Homo sapiens at Ilsenhöhle in Ranis, Germany

Geoff M Smith et al. Nat Ecol Evol. 2024 Mar.

Abstract

Recent excavations at Ranis (Germany) identified an early dispersal of Homo sapiens into the higher latitudes of Europe by 45,000 years ago. Here we integrate results from zooarchaeology, palaeoproteomics, sediment DNA and stable isotopes to characterize the ecology, subsistence and diet of these early H. sapiens. We assessed all bone remains (n = 1,754) from the 2016-2022 excavations through morphology (n = 1,218) or palaeoproteomics (zooarchaeology by mass spectrometry (n = 536) and species by proteome investigation (n = 212)). Dominant taxa include reindeer, cave bear, woolly rhinoceros and horse, indicating cold climatic conditions. Numerous carnivore modifications, alongside sparse cut-marked and burnt bones, illustrate a predominant use of the site by hibernating cave bears and denning hyaenas, coupled with a fluctuating human presence. Faunal diversity and high carnivore input were further supported by ancient mammalian DNA recovered from 26 sediment samples. Bulk collagen carbon and nitrogen stable isotope data from 52 animal and 10 human remains confirm a cold steppe/tundra setting and indicate a homogenous human diet based on large terrestrial mammals. This lower-density archaeological signature matches other Lincombian-Ranisian-Jerzmanowician sites and is best explained by expedient visits of short duration by small, mobile groups of pioneer H. sapiens.

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Conflict of interest statement

The authors declare no competing interests.

Figures

Fig. 1
Fig. 1. Geographic location, stratigraphy and excavation plan for the 2016-2022 excavations at Ranis.
a, Geographic location of Ranis and the main LRJ sites, b, plan of the 2016–2022 excavations and c, stratigraphic sequence of the cave Ilsenhöhle at Ranis. Orange dots in b and c mark the layers and squares that were sampled for sedaDNA. R denotes rockfall events. See Mylopotamitaki et al. for the description of the sedimentary and chronological framework. In a, the location of main LRJ sites (1–7 and 9–15, adapted from Hussain et al; 8, Aldhouse-Green; 16–17, Demidenko and Škrdla). Triangles mark sites with well-contextualized fauna. 1, Ranis; 2, Schmähingen-Kirchberghöhle; 3, Bench Quarry; 4, Kent’s Cavern; 5, Soldier’s Hole; 6, Hyena Den; 7, Badger Hole; 8, Paviland Cave; 9, Robin Hood’s Cave; 10, Grange Farm; 11, Beedings; 12, Spy; 13, Goyet; 14, Nietoperzowa Cave; 15, Koziarnia Green Cave; 16, Líšeň Podolí I; 17, Želešice III. The map was created in QGIS based on Shuttle Radar Topography Mission data V4 (http://srtm.csi.cgiar.org). In b, Each numbered square is 1 m2. The basal sequence including the LRJ layers was excavated in the red area of squares 1003/999, 1003/1000, 1004/999 and 1004/1000. Panels a and b were created with Affinity Designer version 2.3.0.2165.
Fig. 2
Fig. 2. Overview of the bone fragments and ancient mammalian DNA identified across layers 12–7 at Ilsenhöhle in Ranis.
The bone fragment line includes identifications both based on morphology and through ZooMS (for a breakdown by method, see Extended Data Table 1). a, Carnivores, including Canidae, Hyenaidae, Felinae, red fox (V. vulpes) and wolverine (G. gulo). b, Ursidae (Ursus spelaeus, Ursus arctos, Ursus sp.). c, Megafauna, including mammoth (M. primigenius) and rhinoceros (C. antiquitatis). d, Equidae (Equus ferus and Equus sp.). e, Cervidae, including reindeer (R. tarandus) and Cervus sp. f, Bovidae (Bos primigenius, Bison priscus, Bos/Bison). The proportion of aDNA was calculated based on the number of ancient mtDNA fragments assigned to each taxon per layer (Supplementary Tables 3–6). The % on the y axis includes %NISP for bone fragments and percentage of identified sedaDNA (%sedaDNA). Animal silhouettes downloaded from https://www.phylopic.org/.
Fig. 3
Fig. 3. Ecological diversity indices for faunal assemblages from layers 12–7 at Ranis.
a, Shannon–Wiener index, which measures faunal diversity (H), and b, Simpson’s assemblage evenness index (D) show respective increased diversity and assemblage evenness per layer; values for H are normally between 1.5 and 3.5, with larger values indicating greater taxonomic diversity; values for D are between 0 (no taxonomic evenness) and 1 (complete taxonomic evenness); note that different scales are used on the y axis in a and b; see Supplementary Table 7 for a breakdown of NISP, NTAXA and ecological indices. Point size is scaled to the NISP.
Fig. 4
Fig. 4. Bone preservation at Ranis in LRJ layers 8 and 9.
a, Bone surface readability by layer; low readability (0–50% bone surface remaining), high readability (51–100% bone surface remaining), see Methods for further details. b, Bone weathering by layer; low weathering (stages 0 and 1); medium weathering (stages 2 and 3) based on Behrensmeyer. c, COL1ɑ1 508–519 deamidation by layer plotted with bone readability. d, COL1ɑ1 508–519 deamidation by layer plotted with bone weathering stages. Sample sizes in c: layer 8: low readability (n = 22), high readability (n = 168); layer 9: low readability (n = 19), high readability (n = 50). Sample sizes in d: layer 8: low weathering (n = 186), high weathering (n = 4); layer 9: low weathering (n = 64), high weathering (n = 5). Box plots in c and d, box extends from first quartile (Q1 on the left) to third quartile (Q3 on the right) with bold line in the middle representing the median. Lines extending from both ends of the box indicate variability outside Q1 and Q3; minimum/maximum whisker values are calculated as Q1/Q3 ± 1.5 × IQR. Everything outside is represented as an outlier. IQR, interquartile range.
Fig. 5
Fig. 5. Bone surface modifications from Ranis.
a, Carnivore modifications: 1, digested piece (layer 10, 16/116-159503); 2, digested piece (layer 7, 16/116-151389); 3, carnivore gnawing (layer 8, 16/116-151583); 4, U. spelaeus: carnivore tooth pit (layer 11, 16/116-186374); 5, graph illustrates different carnivore modifications as percentage of total number of carnivore modifications (n = 527) across all layers (%modNSP). b, Human modifications: 1, unknown mammal: burnt fragment (layer 11, 16/116-186401); 2, cut-marked fragment (layer 9, 16/116-159345); 3, C. elaphus: marrow fracture (layer 8, 16/116-159070); 4, graph illustrates different human modifications as percentage of total number of human modifications (n = 35) across all layers (%modNSP); for data underlying both graphs, see Supplementary Table 21. All photo scales are in centimetres.
Fig. 6
Fig. 6. Bulk collagen δ13C and δ15N values for mammal remains from layers 12–7 (2016–2022 excavations) and layers XI–VIII from the 1932–1938 excavation at Ranis.
Mammalian isotope data: n = 52. Hominin isotope data: n = 10.
Extended Data Fig. 1
Extended Data Fig. 1
Mammalian ancient DNA recovered from sediment samples from the 2016-2022 excavations at Ilsenhöhle in Ranis. 26 sediment samples were analysed across Layers 7 (n samples = 5), 8 (n samples = 6), 9 (n samples = 3), 10 (n samples = 2), 11 (n samples = 9) and 12 (n samples = 1). Assignments to Cricetidae are not included in this figure. A detailed breakdown of the aDNA data can be found in SI Table 5.
Extended Data Fig. 2
Extended Data Fig. 2
Overview of the length of all piece-plotted bone fragments recovered across Layers 12-7 at Ilsenhöhle in Ranis; B: Overview of length for major taxa from Layers 12-7 at Ilsenhöhle in Ranis. Figure 2a sample sizes are: Layer 12 (n = 18); Layer 11 (n = 565); Layer 10 (n = 92); Layer 9 (n = 115); Layer 8 (n = 244); Layer 7 (n = 722). Figure 2b sample sizes are: Ursidae (Layer 12 n = 1; Layer 11 n = 47; Layer 10 n = 7; Layer 9 n = 23; Layer 8 n = 62; Layer 7 n = 18); Rhinocerotidae sp. (Layer 12 n = 3; Layer 11 n = 13; Layer 10 n = 8; Layer 9 n = 5; Layer 8 n = 28; Layer 7 n = 3); Equidae sp. (Layer 12 n = 1; Layer 11 n = 11; Layer 10 n = 3; Layer 9 n = 7; Layer 8 n = 18; Layer 7 n = 18); Rangifer tarandus (Layer 12 n = 6; Layer 11 n = 32; Layer 10 n = 32; Layer 9 n = 13; Layer 8 n = 67; Layer 7 n = 27); Bos/Bison sp. (Layer 12 n = 1; Layer 11 n = 12; Layer 10 n = 5; Layer 9 n = 9; Layer 8 n = 18; Layer 7 n = 7). Box plot in Extended Data Figs. 2a and 2b: box extends from first quartile (Q1 on left) to third quartile (Q3 on right) with bold line in middle representing (median); Lines extending from both ends of the box indicate variability outside Q1 and Q3; minimum/maximum whisker values are calculated as Q1/Q3 -/ + 1.5 * IQR. Everything outside is represented as an outlier.
Extended Data Fig. 3
Extended Data Fig. 3
Glutamine deamidation values for α1 508 of the bone fragments analysed through ZooMS. These are seen as an indicator for the biomolecular preservation of the bone. Sample sizes are Layer 7 (n = 44), Layer 8 (n = 190), Layer 9 (n = 69), Layer 10 (n = 61), Layer 11 (n = 133) and Layer 12 (n = 16); see SI Tables 14 and SI Table 15. Box plot in Extended Data Fig. 3: box extends from first quartile (Q1 on left) to third quartile (Q3 on right) with bold line in middle representing (median); Lines extending from both ends of the box indicate variability outside Q1 and Q3; minimum/maximum whisker values are calculated as Q1/Q3 -/ + 1.5 * IQR. Everything outside is represented as an outlier.
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