Table 1.
Dataset used for the statistic analyses.
| Species | Order | Sub-order | Average reconstructed cranial orientation (in °) | Average neutral head posture (in °) | Body mass (in kg) | Diet: browser/grazer/mixed/other | Frontal head to head butting: yes/no | Habitat: open/closed/mixed/rocky/semi-aquatic |
|---|---|---|---|---|---|---|---|---|
| Elephas maximus | Afrotheria | Paenungulata | ? | 39 (n = 105) | 2720 | Mixed | Yes | Mixed |
| Heterohyrax brucei | Afrotheria | Paenungulata | ? | 32 (n = 95) | 2 | Browser | No | Rocky |
| Loxodonta africana | Afrotheria | Paenungulata | 28 (n = 3) | 48 (n = 103) | 4000 | Mixed | Yes | Open |
| Procavia capensis | Afrotheria | Paenungulata | 21 (n = 4) | 19 (n = 121) | 4 | Grazer | No | Rocky |
| Orycteropus afer | Afrotheria | Tubulidentata | 41 (n = 2) | 42 (n = 37) | 70 | Other | No | Open |
| Hexaprotodon liberiensis | Artiodactyla | Hippopotamidae | 27 (n = 2) | 50 (n = 27) | 262 | Browser | No | Semi-aquatic |
| Hippopotamus amphibius | Artiodactyla | Hippopotamidae | 20 (n = 3) | 59 (n = 74) | 2300 | Grazer | No | Semi-aquatic |
| Addax nasomaculatus | Artiodactyla | Ruminantia | 45 (n = 1) | 48 (n = 75) | 93 | Grazer | Yes | Open |
| Aepyceros melampus | Artiodactyla | Ruminantia | 53 (n = 2) | 24 (n = 41) | 58 | Mixed | Yes | Open |
| Alcelaphus buselaphus | Artiodactyla | Ruminantia | 68 (n = 8) | 55 (n = 67) | 185 | Grazer | Yes | Open |
| Alces alces | Artiodactyla | Ruminantia | 44 (n = 2) | 33 (n = 28) | 543 | Browser | Yes | Closed |
| Ammotragus lervia | Artiodactyla | Ruminantia | 51 (n = 1) | 43 (n = 71) | 103 | Mixed | Yes | Rocky |
| Antidorcas marsupialis | Artiodactyla | Ruminantia | 42 (n = 3) | 38 (n = 61) | 43 | Mixed | Yes | Open |
| Antilocapra americana | Artiodactyla | Ruminantia | 42 (n = 1) | 36 (n = 140) | 55 | Browser | Yes | Open |
| Antilope cervicapra | Artiodactyla | Ruminantia | 35 (n = 3) | 23 (n = 121) | 37 | Grazer | Yes | Open |
| Axis axis | Artiodactyla | Ruminantia | 46 (n = 1) | 27 (n = 122) | 78 | Grazer | Yes | Open |
| Axis kuhlii | Artiodactyla | Ruminantia | ? | 24 (n = 8) | 36 | Grazer | ? | Open |
| Axis porcinus | Artiodactyla | Ruminantia | 35 (n = 1) | 28 (n = 117) | 43 | Grazer | ? | Open |
| Babyrousa babyrussa | Artiodactyla | Ruminantia | ? | 38 (n = 13) | 90 | Other | No | Closed |
| Babyrousa celebensis | Artiodactyla | Ruminantia | ? | 29 (n = 18) | 90 | Other | No | Closed |
| Bison bison | Artiodactyla | Ruminantia | 70 (n = 3) | 51 (n = 119) | 613 | Grazer | Yes | Mixed |
| Bison bonasus | Artiodactyla | Ruminantia | ? | 49 (n = 44) | 610 | Mixed | Yes | Mixed |
| Bos gaurus | Artiodactyla | Ruminantia | ? | 33 (n = 37) | 825 | Mixed | No | Closed |
| Bos grunniens | Artiodactyla | Ruminantia | ? | 49 (n = 13) | 775 | Grazer | Yes | Open |
| Bos javanicus | Artiodactyla | Ruminantia | 32 (n = 1) | 29 (n = 71) | 650 | Grazer | ? | Closed |
| Bos taurus | Artiodactyla | Ruminantia | 36 (n = 2) | 39 (n = 89) | 650 | Grazer | Yes | Open |
| Boselaphus tragocamelus | Artiodactyla | Ruminantia | 36 (n = 4) | 28 (n = 127) | 205 | Mixed | No | Open |
| Bubalus bubalis | Artiodactyla | Ruminantia | ? | 31 (n = 112) | 771 | Mixed | Yes | Closed |
| Bubalus depressicornis | Artiodactyla | Ruminantia | 51 (n = 3) | 30 (n = 81) | 250 | Mixed | ? | Closed |
| Bubalus mindorensis | Artiodactyla | Ruminantia | 49 (n = 1) | ? | 240 | Grazer | No | Closed |
| Budorcas taxicolor | Artiodactyla | Ruminantia | 54 (n = 2) | 46 (n = 121) | 300 | Mixed | Yes | Closed |
| Capra caucasica | Artiodactyla | Ruminantia | ? | 36 (n = 60) | 96 | Mixed | Yes | Rocky |
| Capra falconeri | Artiodactyla | Ruminantia | 52 (n = 1) | 30 (n = 48) | 71 | Mixed | Yes | Closed |
| Capra hircus | Artiodactyla | Ruminantia | 45 (n = 4) | 32 (n = 164) | 80 | Mixed | Yes | Open |
| Capra ibex | Artiodactyla | Ruminantia | 56 (n = 3) | 25 (n = 72) | 99 | Grazer | Yes | Rocky |
| Capreolus capreolus | Artiodactyla | Ruminantia | 43 (n = 1) | 28 (n = 92) | 28 | Mixed | No | Mixed |
| Cephalophus dorsalis | Artiodactyla | Ruminantia | 40 (n = 3) | ? | 20 | Browser | No | Closed |
| Cephalophus leucogaster | Artiodactyla | Ruminantia | 38 (n = 4) | ? | 18 | Browser | No | Closed |
| Cephalophus natalensis | Artiodactyla | Ruminantia | 30 (n = 3) | 25 (n = 68) | 13 | Browser | No | Closed |
| Cephalophus niger | Artiodactyla | Ruminantia | 36 (n = 3) | 15 (n = 100) | 20 | Browser | No | Closed |
| Cephalophus nigrifrons | Artiodactyla | Ruminantia | 31 (n = 5) | ? | 17 | Browser | No | Closed |
| Cephalophus sylvicultor | Artiodactyla | Ruminantia | 43 (n = 3) | 31 (n = 148) | 63 | Browser | No | Closed |
| Cervus albirostris | Artiodactyla | Ruminantia | ? | 30 (n = 61) | 165 | Grazer | Yes | Mixed |
| Cervus elaphus | Artiodactyla | Ruminantia | 41 (n = 5) | 22 (n = 186) | 128 | Mixed | Yes | Mixed |
| Cervus nippon | Artiodactyla | Ruminantia | 35 (n = 1) | 19 (n = 20) | 95 | Mixed | Yes | Closed |
| Connochaetes gnou | Artiodactyla | Ruminantia | ? | 62 (n = 14) | 134 | Mixed | Yes | Open |
| Connochaetes taurinus | Artiodactyla | Ruminantia | 68 (n = 4) | 60 (n = 74) | 213 | Grazer | Yes | Open |
| Dama dama | Artiodactyla | Ruminantia | 48 (n = 3) | 25 (n = 69) | 35 | Grazer | Yes | Mixed |
| Damaliscus lunatus | Artiodactyla | Ruminantia | 55 (n = 6) | ? | 155 | Grazer | Yes | Open |
| Damaliscus pygargus | Artiodactyla | Ruminantia | 54 (n = 2) | 50 (n = 81) | 71 | Grazer | Yes | Open |
| Elaphodus cephalophus | Artiodactyla | Ruminantia | 27 (n = 1) | 33 (n = 15) | 22 | Mixed | No | Closed |
| Elaphurus davidianus | Artiodactyla | Ruminantia | 46 (n = 2) | 38 (n = 73) | 183 | Grazer | No | Open |
| Gazella dorcas | Artiodactyla | Ruminantia | 30 (n = 3) | ? | 15 | Browser | No | Open |
| Gazella spekei | Artiodactyla | Ruminantia | ? | 28 (n = 154) | 20 | Mixed | ? | Open |
| Gazella subgutturosa | Artiodactyla | Ruminantia | 42 (n = 2) | ? | 28 | Mixed | No | Open |
| Gazella thomsonii | Artiodactyla | Ruminantia | 46 (n = 3) | ? | 21 | Grazer | Yes | Open |
| Giraffa camelopardalis | Artiodactyla | Ruminantia | 32 (n = 5) | 24 (n = 187) | 1317 | Browser | No | Open |
| Hemitragus jemlahicus | Artiodactyla | Ruminantia | 59 (n = 2) | 40 (n = 79) | 75 | Mixed | Yes | Rocky |
| Hippocamelus sp. | Artiodactyla | Ruminantia | 56 (n = 1) | ? | 55 | Browser | ? | Mixed |
| Hippotragus equinus | Artiodactyla | Ruminantia | 46 (n = 1) | 36 (n = 78) | 262 | Grazer | Yes | Open |
| Hippotragus niger | Artiodactyla | Ruminantia | 44 (n = 2) | 40 (n = 140) | 228 | Grazer | Yes | Open |
| Hydropotes inermis | Artiodactyla | Ruminantia | ? | 28 (n = 5) | 12 | Mixed | No | Open |
| Kobus ellipsiprymnus | Artiodactyla | Ruminantia | 42 (n = 4) | 31 (n = 76) | 211 | Grazer | Yes | Mixed |
| Kobus leche | Artiodactyla | Ruminantia | 46 (n = 2) | 27 (n = 59) | 91 | Grazer | Yes | Open |
| Kobus megaceros | Artiodactyla | Ruminantia | ? | 17 (n = 68) | 90 | Grazer | Yes | Open |
| Kobus vardoni | Artiodactyla | Ruminantia | 39 (n = 2) | ? | 66 | Grazer | Yes | Open |
| Litocranius walleri | Artiodactyla | Ruminantia | 33 (n = 3) | 14 (n = 181) | 41 | Browser | No | Open |
| Madoqua sp. | Artiodactyla | Ruminantia | 25 (n = 3) | 24 (n = 97) | 6 | Browser | No | Open |
| Mazama americana | Artiodactyla | Ruminantia | 35 (n = 1) | ? | 17 | Mixed | No | Closed |
| Moschus moschiferus | Artiodactyla | Ruminantia | 25 (n = 1) | 32 (n = 16) | 12 | Mixed | No | Closed |
| Muntiacus reevesi | Artiodactyla | Ruminantia | 43 (n = 2) | 30 (n = 78) | 21 | Mixed | No | Closed |
| Naemorhedus crispus | Artiodactyla | Ruminantia | 50 (n = 1) | ? | 38 | Mixed | No | Rocky |
| Naemorhedus goral | Artiodactyla | Ruminantia | 41 (n = 2) | 38 (n = 33) | 29 | Mixed | No | Rocky |
| Naemorhedus griseus | Artiodactyla | Ruminantia | ? | 38 (n = 10) | 29 | Mixed | ? | Rocky |
| Naemorhedus sumatraensis | Artiodactyla | Ruminantia | 47 (n = 3) | ? | 29 | Mixed | ? | Closed |
| Nanger dama | Artiodactyla | Ruminantia | ? | 23 (n = 41) | 46 | Mixed | Yes | Open |
| Nanger granti | Artiodactyla | Ruminantia | 39 (n = 4) | ? | 56 | Mixed | Yes | Open |
| Nanger soemmeringii | Artiodactyla | Ruminantia | ? | 25 (n = 108) | 42 | Mixed | Yes | Open |
| Neotragus batesi | Artiodactyla | Ruminantia | 28 (n = 3) | ? | 4 | Browser | No | Closed |
| Nesotragus moschatus | Artiodactyla | Ruminantia | 26 (n = 2) | ? | 4 | Browser | No | Closed |
| Nilgiritragus hylocrius | Artiodactyla | Ruminantia | 61 (n = 2) | ? | 90 | Grazer | Yes | Rocky |
| Odocoileus virginianus | Artiodactyla | Ruminantia | ? | 19 (n = 22) | 79 | Mixed | Yes | Closed |
| Okapia johnstoni | Artiodactyla | Ruminantia | 40 (n = 2) | 34 (n = 193) | 283 | Browser | No | Closed |
| Oreamnos americanus | Artiodactyla | Ruminantia | 50 (n = 3) | 52 (n = 52) | 91 | Grazer | No | Rocky |
| Oreotragus oreotragus | Artiodactyla | Ruminantia | 34 (n = 3) | 15 (n = 262) | 14 | Browser | Yes | Rocky |
| Oryx beisa | Artiodactyla | Ruminantia | ? | 48 (n = 48) | 163 | Mixed | Yes | Open |
| Oryx dammah | Artiodactyla | Ruminantia | 46 (n = 1) | 44 (n = 87) | 137 | Grazer | Yes | Open |
| Oryx gazella | Artiodactyla | Ruminantia | 49 (n = 2) | 45 (n = 60) | 163 | Grazer | Yes | Open |
| Oryx leucoryx | Artiodactyla | Ruminantia | ? | 43 (n = 60) | 75 | Grazer | Yes | Open |
| Ourebia ourebi | Artiodactyla | Ruminantia | 31 (n = 3) | ? | 12 | Grazer | No | Open |
| Ovibos moschatus | Artiodactyla | Ruminantia | 42 (n = 2) | 46 (n = 21) | 475 | Mixed | Yes | Open |
| Ovis ammon | Artiodactyla | Ruminantia | 36 (n = 1) | 23 (n = 53) | 163 | Mixed | Yes | Rocky |
| Ovis aries | Artiodactyla | Ruminantia | 37 (n = 2) | 22 (n = 104) | 110 | Mixed | Yes | Open |
| Ovis canadensis | Artiodactyla | Ruminantia | 55 (n = 2) | ? | 98 | Browser | Yes | Rocky |
| Ovis orientalis | Artiodactyla | Ruminantia | 31 (n = 1) | 28 (n = 11) | 48 | Grazer | Yes | Rocky |
| Panolia eldii | Artiodactyla | Ruminantia | ? | 20 (n = 60) | 93 | Grazer | ? | Open |
| Pantholops hodgsonii | Artiodactyla | Ruminantia | 46 (n = 3) | ? | 33 | Grazer | Yes | Open |
| Pelea capreolus | Artiodactyla | Ruminantia | 45 (n = 3) | ? | 25 | Mixed | No | Open |
| Philantomba monticola | Artiodactyla | Ruminantia | 16 (n = 1) | 26 (n = 23) | 7 | Browser | No | Closed |
| Procapra gutturosa | Artiodactyla | Ruminantia | 38 (n = 3) | ? | 30 | Grazer | No | Open |
| Pseudois nayaur | Artiodactyla | Ruminantia | 65 (n = 1) | 28 (n = 73) | 55 | Grazer | Yes | Rocky |
| Pudu puda | Artiodactyla | Ruminantia | 22 (n = 1) | 27 (n = 30) | 10 | Browser | No | Closed |
| Rangifer tarandus | Artiodactyla | Ruminantia | 45 (n = 2) | 35 (n = 189) | 99 | Grazer | Yes | Mixed |
| Raphicerus campestris | Artiodactyla | Ruminantia | 28 (n = 5) | 28 (n = 9) | 11 | Mixed | No | Open |
| Redunca arundinum | Artiodactyla | Ruminantia | 34 (n = 2) | 23 (n = 128) | 73 | Grazer | Yes | Open |
| Redunca fulvorufula | Artiodactyla | Ruminantia | 39 (n = 3) | 23 (n = 33) | 29 | Grazer | Yes | Open |
| Rucervus duvauceli | Artiodactyla | Ruminantia | ? | 23 (n = 17) | 180 | Grazer | ? | Open |
| Rupicapra rupicapra | Artiodactyla | Ruminantia | 42 (n = 2) | 36 (n = 84) | 37 | Grazer | Yes | Rocky |
| Rusa alfredi | Artiodactyla | Ruminantia | ? | 24 (n = 10) | 53 | Mixed | ? | Closed |
| Rusa timorensis | Artiodactyla | Ruminantia | 47 (n = 1) | 29 (n = 32) | 117 | Grazer | ? | Open |
| Rusa unicolor | Artiodactyla | Ruminantia | 38 (n = 1) | 21 (n = 17) | 273 | Mixed | Yes | Mixed |
| Saiga tatarica | Artiodactyla | Ruminantia | 43 (n = 2) | 38 (n = 2) | 48 | Grazer | Yes | Open |
| Sylvicapra grimmia | Artiodactyla | Ruminantia | 30 (n = 4) | ? | 19 | Browser | No | Mixed |
| Syncerus caffer caffer | Artiodactyla | Ruminantia | 51 (n = 2) | 27 (n = 70) | 648 | Grazer | Yes | Open |
| Syncerus caffer nanus | Artiodactyla | Ruminantia | ? | 24 (n = 72) | 320 | Mixed | No | Closed |
| Taurotragus derbianus | Artiodactyla | Ruminantia | ? | 27 (n = 55) | 700 | Browser | Yes | Closed |
| Taurotragus oryx | Artiodactyla | Ruminantia | 49 (n = 4) | 30 (n = 53) | 470 | Browser | Yes | Mixed |
| Tetracerus quadricornis | Artiodactyla | Ruminantia | 31 (n = 3) | ? | 17 | Browser | No | Mixed |
| Tragelaphus angasii | Artiodactyla | Ruminantia | 29 (n = 3) | 22 (n = 76) | 78 | Mixed | Yes | Closed |
| Tragelaphus euryceros | Artiodactyla | Ruminantia | 25 (n = 1) | 26 (n = 197) | 330 | Browser | No | Closed |
| Tragelaphus imberbis | Artiodactyla | Ruminantia | 46 (n = 1) | 21 (n = 252) | 82 | Browser | Yes | Closed |
| Tragelaphus scriptus | Artiodactyla | Ruminantia | 26 (n = 3) | 25 (n = 59) | 53 | Browser | Yes | Closed |
| Tragelaphus spekei | Artiodactyla | Ruminantia | 35 (n = 1) | 20 (n = 125) | 98 | Mixed | Yes | Closed |
| Tragelaphus strepsiceros | Artiodactyla | Ruminantia | 41 (n = 4) | 23 (n = 78) | 214 | Browser | Yes | Mixed |
| Tragulus javanicus | Artiodactyla | Ruminantia | 33 (n = 2) | 15 (n = 144) | 4 | Browser | No | Closed |
| Tragulus nigricans | Artiodactyla | Ruminantia | ? | 12 (n = 50) | 4 | Browser | No | Closed |
| Catagonus wagneri | Artiodactyla | Suoidea | ? | 46 (n = 47) | 35 | Browser | No | Closed |
| Phacochoerus africanus | Artiodactyla | Suoidea | 55 (n = 12) | 46 (n = 75) | 73 | Grazer | Yes | Open |
| Potamochoerus porcus | Artiodactyla | Suoidea | 45 (n = 1) | 53 (n = 30) | 85 | Other | Yes | Closed |
| Sus barbatus | Artiodactyla | Suoidea | ? | 55 (n = 7) | 97 | Other | ? | Closed |
| Sus cebifrons | Artiodactyla | Suoidea | ? | 51 (n = 14) | 28 | Browser | ? | Closed |
| Sus scrofa | Artiodactyla | Suoidea | 37 | 47 (n = 79) | 200 | Other | No | Closed |
| Tayassu pecari | Artiodactyla | Suoidea | ? | 43 (n = 21) | 33 | Other | No | Closed |
| Camelus bactrianus | Artiodactyla | Tylopoda | 15 (n = 2) | 2 (n = 136) | 570 | Mixed | No | Open |
| Camelus dromedarius | Artiodactyla | Tylopoda | 13 (n = 3) | -1 (n = 127) | 450 | Browser | No | Open |
| Lama guanicoe | Artiodactyla | Tylopoda | ? | 15 (n = 89) | 120 | Grazer | No | Open |
| Lama lama | Artiodactyla | Tylopoda | ? | 9 (n = 48) | 94 | Grazer | No | Rocky |
| Vicugna pacos | Artiodactyla | Tylopoda | 23 (n = 1) | 17 (n = 55) | 66 | Grazer | No | Open |
| Vicugna vicugna | Artiodactyla | Tylopoda | 17 (n = 1) | 20 (n = 66) | 50 | Grazer | No | Rocky |
| Equus asinus | Perissodactyla | Equidae | 53 (n = 2) | 58 (n = 88) | 275 | Mixed | No | Open |
| Equus burchelli | Perissodactyla | Equidae | 47 (n = 2) | 59 (n = 109) | 280 | Grazer | No | Open |
| Equus caballus | Perissodactyla | Equidae | 44 (n = 4) | 59 (n = 135) | 250 | Grazer | No | Open |
| Equus grevyi | Perissodactyla | Equidae | ? | 66 (n = 182) | 401 | Grazer | No | Open |
| Equus hemionus | Perissodactyla | Equidae | ? | 58 (n = 117) | 325 | Grazer | No | Open |
| Equus zebra | Perissodactyla | Equidae | 50 (n = 1) | 61 (n = 156) | 274 | Grazer | No | Open |
| Ceratotherium simum | Perissodactyla | Rhinocerotidae | 38 (n = 9) | 57 (n = 140) | 1930 | Grazer | No | Open |
| Dicerorhinus sumatrensis | Perissodactyla | Rhinocerotidae | 25 (n = 1) | ? | 1400 | Browser | ? | Closed |
| Diceros bicornis | Perissodactyla | Rhinocerotidae | 36 (n = 4) | 35 (n = 80) | 1129 | Browser | No | Open |
| Rhinoceros sondaicus | Perissodactyla | Rhinocerotidae | 28 (n = 2) | ? | 1890 | Browser | ? | Semi-aquatic |
| Rhinoceros unicornis | Perissodactyla | Rhinocerotidae | 34 (n = 2) | 17 (n = 83) | 1416 | Mixed | No | Open |
| Tapirus bairdii | Perissodactyla | Tapiridae | ? | 21 (n = 34) | 200 | Browser | No | Semi-aquatic |
| Tapirus indicus | Perissodactyla | Tapiridae | 14 (n = 1) | 28 (n = 117) | 263 | Browser | No | Semi-aquatic |
| Tapirus terrestris | Perissodactyla | Tapiridae | 25 (n = 3) | 23 (n = 149) | 200 | Browser | No | Semi-aquatic |
Data on body mass, diet, head-butting behaviour, and habitat are from the literature (see Supplementary Table S1). n represents the number of measurements.