TABLE 1. Localities used in this study with their ages, species analysed and NAP %.
Locality | Country | Age | Species analysed in this study | Locality used in community-level analyses | Minimum NAP % | Maximum NAP % | Mean NAP % | Reference for age | Reference for pollen record |
Star Carr | UK | MIS 1 | B. primigenius, C. elaphus, C. capreolus, A. alces | yes | 15.0 | 42.0 | 25.3 | Innes et al., 2011; Penkman et al., 2011 | Clark, 1954 |
Late-glacial localities (pollen zone III) | Ireland | MIS 2, Allerød-interstadial | M. giganteus | yes | 45.0 | 93.0 | 74.8 | Watts, 1997; Barnosky, 1986 | Watts, 1977 |
Gough's Cave | UK | MIS 2, Bølling interstadial (GI-1e) | E. ferus, C. elaphus | yes | 89.0 | 94.0 | 91.7 | Currant and Jacobi, 2001; Jacobi and Higham 2009 | Leroi-Gourhan, 1986 |
Whitemoor Haye | UK | MIS 3 | C. antiquitatis | no | 82.3 | 96.7 | 89.5 | Schreve et al., 2013 | Schreve et al., 2013 |
Kent's Cavern (cave earth) | UK | MIS 3 | E. ferus, B. priscus, C. elaphus, R. tarandus, M. giganteus, C. antiquitatis | no | Bocherens and Fogel, 1995; Currant and Jacobi, 2001 | ||||
Isleworth | UK | MIS 5a-d | B. priscus, R. tarandus | no | 86.9 | 94.0 | 90.4 | Penkman et al., 2011; Bates et al., 2014 | Kerney et al., 1982 |
Wretton (Devensian strata) | UK | MIS 5a-d | B. priscus, R. tarandus | no | 80.0 | 98.0 | 89.0 | Lewin and Gibbard, 2010 | West et al., 1974 |
Villa Seckendorff | Germany | MIS 5a-d | E. ferus, E. hydruntinus, B. priscus, B. primigenius, C. elaphus, M. giganteus, C. antiquitatis | no | Ziegler, 1996 | ||||
Aufhausener Höhle | Germany | Last glacial (Würmian) | E. ferus, C. antiquitatis | no | Kley, 1966 | ||||
Upper Rhine valley localities: Brühl (Koller), Otterstadt, Edingen, Ketsch, Lampertheim in der Tanne | Germany | Late Pleistocene | B. primigenius, C. elaphus, C. capreolus, D. dama, A. alces, S. kirchbergensis, C. antiquitatis | no | Koenigswald and Beug, 1988; Dietrich and Rathgeber, 2012 | ||||
Reilingen | Germany | MIS 5e? | S. hemitoechus | no | Ziegler and Dean, 1998 | ||||
Taubach | Germany | MIS 5e | E. ferus, B. priscus, C. elaphus, . capreolus, S. kirchbergensis | no | Brunnacker et al., 1983; van Kolfschoten, 2000 | ||||
Barrington | UK | MIS 5e | B. priscus, B. primigenius, C. elaphus, D. dama, S. hemitoechus | yes | 89.0 | 94.0 | 91.5 | Ashton et al., 2011 | Gibbard and Stuart, 1982 |
Joint Mitnor Cave | UK | MIS 5e | B. priscus, C. elaphus, D. dama, S. hemitoechus | no | Ashton et al., 2011 | ||||
Kirkdale Cave | UK | MIS 5e | B. primigenius | no | Ashton et al., 2011 | ||||
Hoe Grange quarry | UK | MIS 5e | B. priscus, B. primigenius, D. dama | no | Ashton et al., 2011 | ||||
Brundon | UK | MIS 7 | E. ferus, B. priscus, B. primigenius | no | Ashton et al., 2011 | ||||
Ilford | UK | MIS 7 | E. ferus, B. primigenius, C. elaphus, C. capreolus, S. kirchbergensis, S. hemitoechus | yes | 18.0 | 72.0 | 44.6 | Ashton et al., 2011; Penkman et al., 2011 | West et al., 1964 |
Crayford | UK | MIS 7 | E. ferus, B. primigenius, C. elaphus, S. kirchbergensis, C. antiquitatis | no | Ashton et al., 2011; Penkman et al., 2011 | ||||
Aveley (zone II B) | UK | MIS 7 | B. primigenius | no | 9.2 | 57.9 | 28.2 | Ashton et al., 2011; Penkman et al., 2011 | West, 1969 |
Grays Thurrock | UK | MIS 9 | E. ferus, B. primigenius, C. elaphus, D. dama, M. giganteus, S. kirchbergensis | yes | 12.0 | 26.0 | 19.2 | Ashton et al., 2011; Penkman et al., 2011 | Gibbard, 1994 |
Steinheim a.d. Murr, Grube Sammet | Germany | MIS 10 | E. ferus, B. priscus, B. primigenius, C. elaphus, M. giganteus | yes | Schreve and Bridgland, 2002 | ||||
Steinheim a.d. Murr, Grube Sigrist | Germany | MIS 11 | E. ferus, C. elaphus | no | Schreve and Bridgland, 2002 | ||||
Clacton | UK | MIS 11 | E. ferus, B. primigenius, C. elaphus, D. dama, S. hemitoechus | yes | 5.0 | 67.0 | 27.3 | Schreve, 2001; Penkman et al., 2011 | Bridgland et al., 1999 |
Swanscombe (lower loam) | UK | MIS 11 | E. ferus, B. priscus, B. primigenius, D. dama, M. giganteus, S. hemitoechus | yes | 11.0 | 66.0 | 31.6 | Schreve, 2001; Penkman et al., 2011 | Conway, 1996 |
Hoxne | UK | MIS 11 | E. ferus, C. elaphus | no | 12.1 | 37.3 | 23.5 | Schreve, 2000; Penkman et al., 2011 | Mullenders, 1993 |
Frankenbacher Sande | Germany | MIS 11 | E. mosbachensis, B. schoetensacki | no | Van Asperen, 2010 | ||||
Boxgrove (horizons 5 and 4 c) | UK | MIS 13 | E. ferus, B. schoetensacki, C. elaphus, D. roberti, Megacerini sp., S. hundsheimensis, S. cf. megarhinus | yes | 5.0 | 12.0 | 8.5 | Roberts and Parfitt, 1999 | Roberts, 1986 |
Pakefield (pollen zone Cr II) | UK | MIS 15 or MIS 17 | B. schoetensacki, S. hundsheimensis | no | 20.0 | 57.0 | 33.3 | Penkman et al., 2011 | West, 1980 |
Mauer | Germany | MIS 15 | E. mosbachensis, B. schoetensacki, C. elaphus, C. latifrons, S. hundsheimensis | yes | 2.7 | 55.0 | 25.7 | Wagner et al., 2011 | Urban, 1992 |
Süssenborn | Germany | MIS 16 | E. sussenbornensis, E. altidens, B. schoetensacki, C. elaphus, C. sussenbornensis, C. latifrons, Megacerini sp., S. hundsheimensis | no | Kahlke et al., 2010; Kahlke and Kaiser, 2011 | ||||
Voigtstedt | Germany | MIS 17 | C. elaphus, C. sussenbornensis, Megacerini sp., S. hundsheimensis | yes | 1.0 | 22.0 | 11.5 | Maul et al., 2007; Kahlke and Kaiser, 2011 | Erd, 1970 |
West Runton | UK | MIS 17 | Equus sp., B. schoetensacki, C. elaphus, D. cf. roberti, Capreolus sp., Megacerini sp., S. hundsheimensis | yes | 5.0 | 55.0 | 44.6 | Stuart and Lister, 2010; Maul and Parfitt, 2010 | Field and Peglar, 2010 |
TABLE 2. Mean mesowear and body mass values with sample sizes (n) and standard deviations (SD), and minimum, maximum and mean environmental NAP % of the most abundant ungulate species across all localities in which they occur. The species are arranged according to the mean NAP % of their environments from lowest (top) to highest (bottom).
Genus |
sp. |
n (mesowear) |
Mean mesowear value |
SD (mesowear) |
Mesowear value range (of locality means) |
n (body mass) |
Mean body mass (kg) |
Body mass range (kg) |
Minimum NAP% |
Maximum NAP% |
Mean NAP% |
Alces | alces | 8 | 1.00 | 0 | 1.00 | 35 | 433 | 202 - 642 | 12 | 42 | 22 |
Capreolus | capreolus | 22 | 1.07 | 0.08 | 1.00 - 1.17 | 53 | 35 | 22 - 51 | 12 | 42 | 23 |
Stephanorhinus | hundsheimensis | 51 | 1.13 | 0.22 | 1.00 - 1.22 | 63 | 1348 | 999 - 1691 | 1 | 57 | 25 |
Cervalces | latifrons | 13 | 1.11 | 0.16 | 1.00 - 1.23 | 19 | 914 | 593 - 1479 | 3 | 55 | 26 |
Bison | schoetensacki | 22 | 1.46 | 0.04 | 1.42 - 1.50 | 78 | 835 | 314 - 1313 | 3 | 57 | 28 |
Stephanorhinus | kirchbergensis | 52 | 1.03 | 0.07 | 1.00 - 1.17 | 25 | 1844 | 1381 - 2538 | 3 | 72 | 30 |
Cervus | elaphus | 122 | 1.16 | 0.27 | 1.03 - 1.38 | 253 | 211 | 77 - 475 | 1 | 94 | 36 |
Dama | dama | 42 | 1.12 | 0.08 | 1.04 - 1.17 | 124 | 87 | 39 - 145 | 5 | 94 | 39 |
Equus | ferus | 174 | 2.35 | 0.29 | 2.00 - 2.45 | 462 | 499 | 301 - 883 | 5 | 94 | 40 |
Bos | primigenius | 79 | 1.44 | 0.05 | 1.35 - 1.50 | 209 | 1121 | 389 - 2010 | 5 | 94 | 0 |
Stephanorhinus | hemitoechus | 53 | 1.29 | 0.27 | 1.19 - 1.44 | 25 | 1522 | 1181 - 2384 | 5 | 94 | 49 |
Megaloceros | giganteus | 48 | 1.38 | 0.22 | 1.10 - 1.69 | 91 | 687 | 329 - 1228 | 11 | 94 | 54 |
Bison | priscus | 73 | 1.45 | 0.21 | 1.38 - 1.83 | 264 | 1011 | 363 - 1930 | 11 | 94 | 76 |
Coelodonta | antiquitatis | 35 | 2.21 | 0.65 | 1.33 - 2.53 | 28 | 1905 | 1038 - 2958 | 82 | 97 | 90 |
Rangifer | tarandus | 17 | 1.09 | 0.19 | 1.09 | 68 | 129 | 43 - 255 | 80 | 98 | 90 |
TABLE 3. Correlations of mean mesowear value of different ungulate groupings with NAP % in the localities. + = significant positive correlation, no = no correlation. Values indicating significant correlations are emboldened.
Nr. of localities | Correlation | R2 | p | ||
All ungulates | Min. NAP | 11 | + | 0.47 | 0.02 |
Max. NAP | 11 | + | 0.67 | 0.002 | |
Mean NAP | 11 | + | 0.6 | 0.005 | |
Equidae | Min. NAP | 8 | no | 0.09 | 0.46 |
Max. NAP | 8 | no | 0.16 | 0.33 | |
Mean NAP | 9 | no | 0.09 | 0.47 | |
Bovidae ( Bos and Bison) | Min. NAP | 7 | no | 0.11 | 0.46 |
Max. NAP | 7 | + | 0.62 | 0.03 | |
Mean NAP | 8 | no | 0.24 | 0.25 | |
Cervidae | Min. NAP | 8 | (+) | 0.44 | 0.05 |
Max. NAP | 8 | + | 0.54 | 0.02 | |
Mean NAP | 9 | + | 0.58 | 0.02 | |
Rhinocerotidae | Min. NAP | 8 | + | 0.59 | 0.02 |
Max. NAP | 8 | + | 0.93 | <0.0001 | |
Mean NAP | 8 | + | 0.82 | 0.0008 |
TABLE 4. Means comparison of mesowear values of species in the presence/absence of other key ungulate species by paired Wilcoxon tests. M = mean mesowear value. Test statistics (Z and p-values) of the means differences are given for each presence/absence pair for each species (statistically significant values are emboldened). The species presence/absence data per locality were obtained from: Arnold-Bemrose and Newton (1905), Adam (1954), Lister (1984), Ziegler (1996), Schreve (1997), van Kolfschoten (2000) and Currant and Jacobi (2001).
Cervus elaphus | Equus ferus | Bos primigenius | Bison priscus | Dama dama | Stephanorhinus hemitoechus | |||||||
M | M | M | M | M | M | |||||||
A. alces present | 1.26 | Z = 2.14; p = 0.03 | 2.36 | Z = 0;
p = 1 |
1.35 | Z = -2.24; p = 0.02 | 1.34 | Z = -1.51, p = 0.13 | 1.04 | Z = -1.67; p = 0.10 | ||
A. alces absent | 1.10 | 2.35 | 1.48 | 1.45 | 1.17 | |||||||
C. capreolus present | 1.26 | Z = 1.46; p = 0.14 | 2.23 | Z = -1.98; p = 0.05 | 1.44 | Z = -1.10; p = 0.27 | 1.38 | Z = -0.97; p = 0.33 | 1.12 | Z = -0.52; p = 0.60 | 1.25 | Z = -1.00; p = 0.32 |
C. capreolus absent | 1.14 | 2.40 | 1.50 | 1.46 | 1.17 | 1.33 | ||||||
S. kirchbergensis present | 1.18 | Z = 0.55; p = 0.58 | 2.27 | Z = -1.24; p = 0.22 | 1.45 | Z = -0.56; p = 0.57 | 1.38 | Z = -1.07; p = 0.28 | 1.13 | Z = -0.22; p = 0.83 | 1.25 | Z = -1.00; p = 0.32 |
S. kirchbergensis absent | 1.14 | 2.38 | 1.48 | 1.46 | 1.14 | 1.33 | ||||||
Dama dama present | 1.14 | Z = -1.62; p = 0.10 | 2.23 | Z = -1.99; p = 0.05 | 1.41 | Z = -2.05; p = 0.04 | 1.43 | Z = -0.64; p = 0.52 | 1.31 | Z = 1.17; p = 0.24 | ||
Dama dama absent | 1.28 | 2.40 | 1.50 | 1.45 | 1.19 | |||||||
Cervus elaphus present | 2.36 | Z = 1.02; p = 0.31 | 1.43 | Z = -0.19; p = 0.85 | ||||||||
Cervus elaphus absent | 2.28 | 1.44 | ||||||||||
Bos primigenius present | 1.24 | Z = 1.19; p = 0.24 | 2.33 | Z = -0.81; p = 0.42 | 1.43 | Z = -0.28; p = 0.78 | 1.12 | Z = -0.52; p = 0.60 | 1.30 | Z = 0.49; p = 0.62 | ||
Bos primigenius absent | 1.14 | 2.42 | 1.43 | 1.17 | 1.25 | |||||||
S. hemitoechus present | 1.11 | Z = -0.72; p = 0.47 | 2.24 | Z = -1.89; p = 0.06 | 1.45 | Z = -0.59; p = 0.55 | 1.43 | Z = -0.64; p = 0.52 | ||||
S. hemitoechus absent | 1.19 | 2.41 | 1.50 | 1.45 | ||||||||
M. giganteus present | 1.15 | Z = -1.57; p = 0.12 | 2.37 | Z = 1.32; p = 0.19 | 1.46 | Z = 0.24; p = 0.81 | 1.43 | Z = -0.19; p = 0.85 | 1.30 | Z = 0.78; p = 0.44 | ||
M. giganteus absent | 1.32 | 2.28 | 1.44 | 1.44 | 1.20 | |||||||
R. tarandus present | 1.19 | Z = 0.29; p = 0.67 | 2.41 | Z = 0.88; p = 0.38 | 1.50 | Z = 0.58; p = 0.57 | 1.44 | Z = 0.30; p = 0.77 | ||||
R. tarandus absent | 1.18 | 2.32 | 1.45 | 1.43 | ||||||||
B. priscus present | 1.15 | Z = -1.32; p = 0.19 | 2.40 | Z = 2.93; p = 0.003 | 1.48 | Z = 1.00; p = 0.32 | 1.19 | Z = 1.79; p = 0.07 | ||||
B. priscus absent | 1.27 | 2.19 | 1.44 | 1.06 | ||||||||
E. ferus present | 1.19 | Z = 0.35; p = 0.72 | 1.46 | Z = 0.24; p = 0.81 | 1.41 | Z = -0.46; p = 0.64 | 1.13 | Z = -0.22; p = 0.83 | 1.25 | Z = -1.00; p = 0.32 | ||
E. ferus absent | 1.20 | 1.44 | 1.46 | 1.14 | 1.33 | |||||||
C. antiquitatis present | 1.2 | Z = 1.02; p = 0.31 | 2.41 | Z = 2.18; p = 0.03 | 1.50 | Z = -0.56; p = 0.57 | 1.45 | Z = 0.64; p = 0.52 | ||||
C. antiquitatis absent | 1.12 | 2.23 | 1.44 | 1.43 |
TABLE 5. Pairwise correlations (correlation coefficients from pairwise comparisons) of mean body mass between species, and (bottom three rows) correlation coefficients of species’ mean body mass with minimum, maximum and mean NAP percentages in localities. Correlations based on comparisons of three or more pairs are shown. The pairwise correlations and their p-values are given in Appendix 5. Statistically significant (p <0.05) correlations are emboldened.
B. priscus | B. scho. | B. prim. | E. ferus | A. alces | C. latifrons | C. elaphus | D. dama | M. giganteus | C. capreolus | R. tarandus | S. kirch. | S. hem. | S. hund. | C. antiq. | |
Bison priscus | 0.65 | -0.67 | 0.3 | -0.71 | 0.52 | -0.99 | 0.99 | ||||||||
Bison schoet. | -0.78 | -0.35 | 0.37 | ||||||||||||
Bos primigenius | 0.65 | 0.62 | -0.08 | 0.18 | -0.4 | 0.89 | |||||||||
Equus ferus | -0.67 | 0.62 | -0.64 | 0.22 | -0.21 | ||||||||||
Alces alces | |||||||||||||||
Cervalces latifrons | -0.78 | 0.76 | |||||||||||||
Cervus elaphus | -0.35 | -0.08 | -0.64 | 0.76 | -0.07 | -0.91 | -0.98 | -0.75 | -0.41 | ||||||
Dama dama | -0.71 | 0.22 | -0.07 | -0.93 | |||||||||||
Meg. giganteus | 0.52 | -0.21 | -0.91 | ||||||||||||
Capreolus capreolus | -0.4 | -0.98 | |||||||||||||
Rangifer tarandus | -0.99 | ||||||||||||||
Steph. kirch. | |||||||||||||||
Steph. hemit. | 0.99 | 0.89 | -0.75 | -0.93 | |||||||||||
Steph. hund. | 0.37 | -0.41 | |||||||||||||
Coelodonta antiquitatis | |||||||||||||||
Minimum NAP % | -0.06 | -0.64 | 0.1 | -0.89 | 0.55 | -0.91 | 1 | 0.56 | -0.88 | ||||||
Maximum NAP % | -0.22 | -0.58 | 0.17 | -0.52 | 0.46 | -0.63 | 0.81 | 0.66 | -0.06 | ||||||
Mean NAP % | -0.11 | -0.82 | 0.18 | -0.78 | 0.61 | -0.82 | 0.92 | 0.66 | 0.07 |