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