Climate change in Europe between 90 and 50 kyr BP and Neanderthal territorial habitability

of the treatment of the climate data and the method of biome reconstruction (see also S2 File). Although the climate model simulations we use are from an earlier generation of climate modelling, there is not a significant body of more recent work covering the time period we are studying (for instance the widely used protocol Paleo Model Intercomparison Project (PMIP) [15] do not include periods of glacial inception). In fact, a recent effort to make more widely available paleoclimate reconstructions [16], which relies on the dataset described by [17] for this period is itself derived from one of the same sets of climate simulations [13] as we consider here and uses a similar method of ecological reconstruction to ours. However, since our con- clusions regarding the suitability of the environment for Neanderthal occupation are indepen- dent of which of our two climate reconstructions we use, for simplicity the ecological niche analysis presented in this paper uses reconstructions based on only one of the two climate models, the single, continuous transient simulation of [14]. Friction and carrying capacity are derived from the biome distribution reconstructed for the climate in each period. Friction is defined as the difficulty of moving in a given environ- ment, with the density of vegetation affecting human movement. Values of friction have been considered, one for each global biome type (table 3.2 by [18]): ranging from 0.1 for environ- ments where mobility is very easy, to 1 for environments across which it is practically impossi-

ble to spread, like ice sheets (see also S3 File). Carrying capacity is an indicator for the available quantity of flora and fauna, corresponding to the maximum number of individuals who can be Fig 1. Location of the 123 Neanderthal fossil sites (1 Goyet Caves; 2 Scladina; 3 La Chaise Bourgois-Delaunay; 4 La Ferrassie; 5 Roc de Marsal; 6 Le Portel; 7 La Rouquette; 8 Grotte de Saint Marcel; 9 Hohlenstein-Stadel; 10 Hunas; 11 Buhlen; 12 Sarstedt; 13 Kefalonia; 14 Grotte de Kalamakia; 15 Tata; 16 Tabun; 17 Grotta Maggiore di S. Bernardino; 18 Grotta de’ Santi; 19 Riparo Mochi; 20 Grotta Madonna dell’Arma; 21 Arma delle Manie; 22 Altai Mountain; 23 Abric Romani; 24 Grotte de Lezetxiki; 25 Valdegoba; 26 El Castillo; 27 Alle-Noir Bois; 28 Korolevo; 29 Staroselye; 30 Kabazi; 31 Prolom II; 32 Teshik-Tash; 33 Kulna; 34 Marillac; 35 Sandougne et abri Brouillard; 36 Grotte de L’Hyene; 37 Regourdou; 38 Grotte d’Unikoté; 39 Grottes d’Isturitz et d’Oxocelhaya; 40 Pech l’Azé1; 41 Abri Laborde; 42 La Brèche de Genay; 43 Grotte de la Verrerie; 44 Loton; 45 Préletang; 46 Abri Pié Lombard; 47 Salzgitter-Lebenstedt; 48 Warendorf; 49 Kokkinopilos; 50 Riparo Zampieri; 51 Caverna delle Fate; 52 Tor Faraj; 53 Conceicao; 54 Mezmaiskaya Cave; 55 Chagyrskaya; 56 Divje Babe; 57 Pinilla del Valle; 58 Grotte de Cotencher; 59 Dederiyeh; 60 Molodova; 61 Kiik-Koba; 62 Obi-Rakhamat Grotto; 63 La Quina; 64 Subalyuk; 65 Ein Qashish; 66 Amud; 67 Grotta della Ghiacciaia; 68 Guattari; 69 Grotta di Cotariova; 70 Vilas ruivas; 71

Cueva Anton; 72 Banyoles; 73 Salzofen Cave; 74 Temnata Cave; 75 Fonseigner; 76 Grottes du Coupe-Gorge; 77 Sites du Cotentin; 78 Mauran; 79 Le Moustier; 80 Fieux; 81 La Chapelle aux Saints; 82 Hortus; 83 Grotte Boccard; 84 Königsaue; 85 Shanidar; 86 Kebara; 87 Riparo Tagliente; 88 Grotta Taddeo; 89 Grotta Tina; 90 Bisceglie; 91 Grotta di Uluzzo; 92 Tor Sabiha; 93 Ochoz; 94 El Salt; 95 Gibraltar 2; 96 Cova del Gegant; 97 Cueva del Boquette; 98 Gibraltar 1; 99 Région de la Löwenbourg; 100 Lynford Quarry; 101 Pin Hole Cave; 102 Kryegjata B; 103 Fonds de forêt; 104 Zobiste; 105 Kadar; 106 Roche-Cotard Cave; 107 Grotte de Gatzarria; 108 La Roquette; 109 Combe Grenal; 110 Grottes du Tuteil et de Caougno; 111 Moula Guercy; 112 Auzières 2; 113 Grotte Vaufrey; 114 Calascio; 115 Molare; 116 Northsea shore; 117 Stajnia; 118 San Antao do Tojal; 119 Furninha; 120 Gabasa; 121 El Sidron; 122 Grotte des Plaints; 123 Quneitra). Topographic data ‘NOAA National Centers for Environmental Information. 2022: ETOPO 2022 15 Arc-Second Global Relief Model. NOAA National Centers for Environmental Information. https://doi.org/10.25921/fd45-gt74. Accessed 2024". https://doi.org/10.1371/journal.pone.0308690.g001 PLOS ONE Neanderthal habitability PLOS ONE | https://doi.org/10.1371/journal.pone.0308690 February 26, 2025 3 / 17