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A revision of European Plesiosminthus (Rodentia, Dipodidae), and new material from the upper Oligocene of Teruel (Spain)

Matthijs Freudenthal and Elvira Martín-Suárez

Plain Language Abstract

Plesiosminthus is an extinct genus of rodents, belonging to the Dipodidae, a family that is currently found across the Northern Hemisphere. The nearest extant relatives of Plesiosminthus are the birch mice (Sicista). The genus Plesiosminthus originated in Asia in the early Oligocene and dispersed into Europe in the late Oligocene. In the early Miocene it became extinct. In this paper we describe two new species and we discuss the European members of the genus. Our material from Spain represents the oldest European occurrence of the genus.

Resumen en Español

Revisión de los Plesiosminthus europeos (Rodentia, Dipodidae), y material nuevo del Oligoceno final de Teruel (España)

En este trabajo revisamos el registro europeo de las especies del género Plesisosminthus y describimos dos nuevas especies. Siguiendo las indicaciones del Código Internacional de Nomenclatura Zoológica, la especie Plesiosminthus promyarion es un nomen dubium y no se debe utilizar. Por primera vez describimos una colección rica del Oligoceno superior (MP 27) de España. Hasta ahora sólo se conocían Plesiosminthus en yacimientos más recientes (MP 28). En Europa se conocen diez especies de Plesiosminthus: nueve del Oligoceno final y una del Mioceno inicial. Para la mayor parte de estas especies no es posible establecer relaciones de parentesco antecesor-descendiente.

Palabras clave: Mammalia, Dipodidae, Oligoceno, Mioceno, nuevas especies.

Traducción: Authors

Résumé en Français

Une révision des Plesiosminthus (Rodentia, Dipodidae) européens et une description de nouveau matériel de l'Oligocène supérieur de Teruel (Espagne)

Le registre fossile européen de Plesiosminthus, un genre de rongeurs, est révisé et deux nouvelles espèces sont décrites. Plesiosminthus promyarion est considéré comme un nomen dubium d'après le Code International de Nomenclature Zoologique et ce nom ne devrait donc plus être utilisé. Jusqu'à présent, ce genre était quasiment inconnu dans les localités plus anciennes que MP28. Pour la première fois, nous décrivons des échantillons conséquents provenant de l'Oligocène supérieur (MP27) d'Espagne. Dix espèces européennes sont connues pour l'instant, une du Miocène inférieur, et les autres de l'Oligocène supérieur. Pour la plupart d'entre elles, il n'est pas possible d'identifier des relations d'ancêtres à descendants.

Mots-clés : Mammalia ; Dipodidae ; Oligocène ; Miocène ; nouvelle espèce

Translator: Antoine Souron

Deutsche Zusammenfassung

Überarbeitung des europäischen Plesiosminthus (Rodentia, Dipodidae) und neues Material aus dem oberen Oligozän von Teruel (Spanien)

Der europäische Nachweis der Nagetier-Gattung Plesiosminthus wird überarbeitet und zwei neue Arten werden beschrieben. Es wurde festgestellt, dass Plesiosminthus promyarion im Sinne der internationalen Regeln der Zoologischen Nomenklatur ein nomen Dubium ist und dass dieser Name nicht länger benutzt werden sollte. Bis jetzt war die Gattung aus Fundstellen die älter als MP28 sind kaum bekannt. Wir beschreiben zum ersten Mal reichhaltige Proben aus dem oberen Oligozän (MP 27) von Spanien. Bis jetzt sind 10 europäische Arten bekannt, eine aus dem unteren Miozän und die anderen aus dem oberen Oligozän. Bei den meisten können keine Vorfahren-Nachfahren-Beziehungen festgestellt werden.

Schlüsselwörter: Mammalia; Dipodidae; Oligozän; Miozän; neue Art

Translator: Eva Gebauer

Arabic

Translator: Ashraf M.T. Elewa

 

 

FIGURE 1. Terminology of molars. Figures represent left-hand molars.

figure1 

FIGURE 2. Distribution of morphology values of protoconid hind arm in m2 (filled circles) and protolophule in M2 (open circles). Enclosing circles and connecting lines indicate possible relationships.

 figure2

FIGURE 3. Plesiosminthus margaritae n. sp. from Mirambueno 1 (occlusal views). 1, m1 sin., RGM 558099; 2, m2 sin., MIR1 168; 3, m3 sin., RGM 558149; 4, M1 sin., RGM 558200; 5, M2 sin., MIR1 189; 6, M3 sin., MIR1 196; 7, m1 dext., RGM 558105; 8, m2 dext., RGM 558138; 9, m3 dext., MIR1 178; 10, M1 dext., MIR1 187; 11, M2 dext., MIR1 193 (Holotype); and 12, M3 dext., MIR1 197.

figure3 

FIGURE 4. Scatter diagrams of molar length (L) and width (W) of Plesiosminthus margaritae n. sp. from Mirambueno 1 (in mm). 1, m1; 2, m2; 3, m3; 4, M1; 5, M2; and 6, M3.

 figure4

FIGURE 5. Scatter diagrams of molar length (L) and width (W) of Plesiosminthus cf. margaritae n. sp. from Vivel del Río 1 (in mm). 1, m1; 2, m2; 3, m3; 4, M1; 5, M2; and 6, M3.

figure5 

FIGURE 6. Scatter diagrams of molar length (L) and width (W) of Plesiosminthus aff. conjunctus (cross) and Plesiosminthus sp. (circle) from Mirambueno 2A, compared with the distribution area (rectangle) of Plesiosminthus conjunctus molars from HERR8 (in mm). 1, m1; 2, m2; 3, m3; 4, M1; 5, M2; and 6, M3.

 figure6

FIGURE 7. m1 length (1) and width (2) of Plesiosminthus species. The bars represent the minimum and maximum in mm, a tick marks the mean, and the number of specimens is given to the right.

figure7 

FIGURE 8. m2 length (1) and width (2) of Plesiosminthus species. The bars represent the minimum and maximum in mm, a tick marks the mean, and the number of specimens is given to the right.

 figure8

FIGURE 9. m3 length (1) and width (2) of Plesiosminthus species. The bars represent the minimum and maximum in mm, a tick marks the mean, and the number of specimens is given to the right.

figure9 

FIGURE 10. M1 length (1) and width (2) of Plesiosminthus species. The bars represent the minimum and maximum in mm, a tick marks the mean, and the number of specimens is given to the right.

figure10 

FIGURE 11. M2 length (1) and width (2) of Plesiosminthus species. The bars represent the minimum and maximum in mm, a tick marks the mean, and the number of specimens is given to the right.

figure11 

FIGURE 12. M3 length (1) and width (2) of Plesiosminthus species. The bars represent the minimum and maximum in mm, a tick marks the mean, and the number of specimens is given to the right.

 figure12

FIGURE 13. Relation of mean length values (in mm) of m1, m2, m3, M1, M2, M3 of Plesiosminthus species. Number of specimens per sample in Table 7.

figure13 

 
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Measurements of Plesiosminthus samples. V' = coefficient of variation as defined by Freudenthal and Cuenca Bescos (1984): 100R/M, where R is the range, and M is the median of the distribution. σ = standard deviation.

  Length   Width
 
m1 N Min. Mean Max. V' σ   N Min. Mean Max. V' σ
CLUZEL 10 1.13 1.210 1.32 15.5 0.055   10 0.76 0.830 0.93 20.1 0.046
CHAVR 36 1.09 1.190 1.31 18.3 0.013   36 0.75 0.870 0.97 25.6 0.049
LAUT2 19 1.20 1.260 1.39 14.7 0.053   25 0.82 0.890 0.92 11.5 0.042
SAULC 15 1.07 1.260 1.39 26.0 0.059   15 0.79 0.880 0.94 17.3 0.044
FORN11 27 1.08 1.222 1.36 23.0 0.063   27 0.76 0.856 0.92 19.0 0.042
VENINF 19 1.13 1.270 1.39 20.6 0.064   19 0.83 0.910 1.02 20.5 0.058
BF53 4 1.30 1.320 1.34 3.0 0.016   4 0.96 1.035 1.08 11.8 0.054
THZ 142 1.19 1.330 1.47 21.1 0.062   142 0.82 0.920 1.02 21.7 0.045
COD3 65 1.18 1.300 1.44 19.8 0.066   65 0.86 0.960 1.06 20.8 0.046
DIEU 33 1.15 1.28 1.40 19.6     33 0.82 0.93 1.04 23.7  
PARR 15 1.17 1.280 1.44 20.7     16 0.86 0.940 1.12 26.3  
SAY1 7 1.07 1.140 1.22 13.1     8 0.70 0.750 0.79 12.1  
HERR8 8 1.17 1.240 1.33 12.8     8 0.82 0.879 0.96 15.7  
PDF 45 1.06 1.150 1.26 17.2 0.049   45 0.74 0.820 0.92 21.7 0.043
PDES 57 0.98 1.140 1.23 22.6 0.049   58 0.67 0.800 0.88 27.1 0.039
GAIM 34 1.05 1.160 1.25 17.4 0.051   34 0.73 0.840 0.94 25.1 0.051
RDUBEY 19 1.04 1.109 1.20 14.3 0.051   19 0.68 0.792 0.88 25.6 0.057
VIV1 42 1.02 1.109 1.23 18.7 0.062   47 0.68 0.776 0.88 25.6 0.050
MIR2A 17 1.02 1.125 1.22 17.9 0.051   18 0.72 0.808 0.89 21.1 0.035
MIR1 17 1.02 1.119 1.19 15.4 0.056   21 0.72 0.796 0.85 16.6 0.039

 

m2 N Min. Mean Max. V' σ   N Min. Mean Max. V' σ
CLUZEL 8 1.10 1.170 1.26 13.6 0.058   8 0.88 0.930 0.98 10.8 0.033
CHAVR 48 1.02 1.120 1.22 17.9 0.050   48 0.82 0.930 1.00 19.8 0.041
LAUT2 28 1.10 1.160 1.22 10.3 0.034   29 0.86 0.950 1.01 16.0 0.048
SAULC 24 1.07 1.170 1.28 17.9 0.057   24 0.82 0.920 1.02 21.7 0.050
FORN11 24 1.00 1.116 1.20 18.2 0.052   24 0.77 0.898 0.98 24.0 0.054
VENINF 25 1.07 1.190 1.30 19.4 0.049   24 0.88 0.950 1.00 12.8 0.041
BF53 4 1.34 1.373 1.40 4.4 0.032   4 1.04 1.048 1.06 1.9 0.010
THZ 123 1.01 1.190 1.33 27.4 0.065   123 0.83 0.950 1.09 27.1 0.051
COD3 59 1.09 1.210 1.34 20.6 0.055   59 0.86 0.990 1.09 23.6 0.049
DIEU 30 1.05 1.17 1.28 19.7     30 0.82 0.92 1.02 21.7  
PARR 13 1.03 1.190 1.35 26.9     16 0.84 0.960 1.11 27.7  
SAY1 6 0.95 1.050 1.15 19.0     6 0.77 0.840 0.89 14.5  
HERR8 15 1.16 1.197 1.26 8.3     15 0.85 0.923 1.00 16.2  
PDF 28 0.99 1.140 1.25 23.2 0.067   28 0.79 0.880 0.98 21.5 0.047
PDES 54 0.99 1.120 1.23 21.6 0.048   54 0.79 0.850 0.94 17.3 0.038
GAIM 32 1.05 1.140 1.21 14.2 0.041   31 0.79 0.870 0.96 19.4 0.039
RDUBEY 9 1.00 1.106 1.18 16.5 0.066   9 0.80 0.841 0.90 11.8 0.044
VIV1 51 0.96 1.063 1.17 19.7 0.050   52 0.73 0.842 0.92 23.0 0.046
MIR2A 16 1.05 1.116 1.21 14.2 0.048   18 0.83 0.893 0.94 12.4 0.032
MIR1 41 0.99 1.093 1.17 16.7 0.041   42 0.78 0.869 0.98 22.7 0.041

 

m3 N Min. Mean Max. V' σ   N Min. Mean Max. V' σ
CLUZEL 13 0.83 0.920 0.99 17.6 0.044   13 0.75 0.820 0.88 16.0 0.043
CHAVR 39 0.78 0.880 0.99 23.7 0.056   39 0.68 0.790 0.90 27.8 0.047
LAUT2 26 0.88 0.960 1.04 16.7 0.051   26 0.75 0.820 0.90 18.2 0.044
SAULC 18 0.90 0.950 1.05 15.4 0.036   18 0.76 0.810 0.89 15.8 0.040
FORN11 18 0.80 0.928 1.00 22.2 0.042   18 0.68 0.799 0.86 23.4 0.043
VENINF 8 0.82 0.900 0.98 17.8 0.052   8 0.70 0.780 0.84 18.2 0.046
BF53 5 1.16 1.220 1.32 12.9 0.062   5 0.98 1.032 1.10 11.5 0.063
THZ 97 0.73 0.890 1.01 32.2 0.049   97 0.70 0.800 0.97 32.3 0.052
COD3 32 0.74 0.860 0.99 28.9 0.053   32 0.70 0.800 0.88 22.8 0.045
DIEU 20 0.68 0.79 0.90 27.8     20 0.70 0.77 0.84 18.2  
PARR 2 0.84 0.890 0.94 11.2     3 0.81 0.850 0.88 8.3  
SAY1 1   0.710         1   0.650      
HERR8 14 0.92 0.961 1.05 13.2     14 0.76 0.819 0.87 13.5  
PDF 21 0.83 0.950 1.08 26.2 0.057   21 0.71 0.790 0.88 21.4 0.046
PDES 16 0.86 0.920 1.00 15.1 0.041   14 0.68 0.750 0.81 17.4 0.037
GAIM 11 0.81 0.930 0.99 20.0 0.050   11 0.72 0.780 0.80 10.5 0.024
RDUBEY 6 0.80 0.883 0.92 14.0 0.043   6 0.72 0.760 0.82 13.0 0.044
VIV1 39 0.76 0.839 0.94 21.2 0.045   40 0.63 0.724 0.82 26.2 0.044
MIR2A 20 0.79 0.867 0.96 19.4 0.051   20 0.68 0.773 0.89 26.8 0.056
MIR1 45 0.78 0.867 0.99 23.7 0.046   45 0.67 0.773 0.86 24.8 0.045

 

P4 N Min. Mean Max. V' σ   N Min. Mean Max. V' σ
CLUZEL 7 0.47 0.550 0.60 24.3 0.042   7 0.49 0.560 0.65 28.1 0.038
CHAVR 18 0.36 0.470 0.60 50.0 0.051   18 0.43 0.530 0.65 40.7 0.049
SAULC 21 0.40 0.480 0.58 36.7 0.053   21 0.42 0.520 0.64 41.5 0.054
THZ 34 0.47 0.610 0.71 40.7 0.043   34 0.57 0.650 0.73 24.6 0.035
COD3 29 0.48 0.590 0.66 31.6 0.040   29 0.51 0.620 0.73 35.5 0.049
GAIM 1   0.500         1   0.600      
VIV1 7 0.49 0.536 0.59 18.5 0.036   7 0.52 0.559 0.60 14.3 0.027
MIR2A 2 0.56 0.565 0.57 1.8     2 0.47 0.505 0.54 13.9  
MIR1 1   0.540         1   0.590      

 

M1 N Min. Mean Max. V' σ   N Min. Mean Max. V' σ
CLUZEL 17 1.09 1.160 1.30 17.6 0.053   17 0.95 1.040 1.11 15.5 0.043
CHAVR 49 1.06 1.130 1.23 14.8 0.044   49 0.93 1.030 1.14 20.3 0.056
LAUT2 20 1.04 1.150 1.26 19.1 0.054   18 0.95 1.030 1.12 16.4 0.049
SAULC 28 1.06 1.150 1.25 16.5 0.047   28 0.90 1.030 1.15 24.4 0.055
FORN11 30 1.04 1.150 1.26 19.1 0.050   30 0.96 1.036 1.12 15.4 0.053
VENINF 27 1.08 1.200 1.30 18.5 0.049   27 0.98 1.080 1.19 19.4 0.049
BF53 6 1.14 1.217 1.38 19.0 0.089   6 0.97 1.122 1.24 24.4 0.088
THZ 119 1.06 1.230 1.36 24.8 0.058   119 0.99 1.140 1.28 25.6 0.059
COD3 71 1.14 1.250 1.39 19.8 0.051   71 1.06 1.160 1.34 23.3 0.052
DIEU 30 1.14 1.22 1.30 13.1     30 1.02 1.15 1.28 22.6  
PARR 13 1.09 1.220 1.37 22.8     13 0.98 1.120 1.22 21.8  
SAY1 3 1.03 1.090 1.14 10.1     2 0.94 0.970 1.00 6.2  
HERR8 18 1.04 1.139 1.21 15.1     18 1.00 1.111 1.21 19.0  
PDF 30 0.96 1.080 1.22 23.9 0.067   30 0.90 1.010 1.17 26.1 0.065
PDES 35 1.00 1.060 1.15 14.0 0.035   35 0.87 0.960 1.02 15.9 0.035
GAIM 28 1.01 1.090 1.18 15.5 0.044   28 0.93 1.010 1.10 16.7 0.045
RDUBEY 12 0.88 1.023 1.13 24.9 0.088   12 0.84 0.948 1.08 25.0 0.080
VIV1 70 0.93 1.050 1.18 23.7 0.049   69 0.85 0.970 1.10 25.6 0.048
MIR2A 30 0.96 1.075 1.19 21.4 0.050   32 0.83 0.973 1.10 28.0 0.049
MIR1 49 0.93 1.062 1.16 22.0 0.048   51 0.85 0.962 1.11 26.5 0.051

 

M2 N Min. Mean Max. V' σ   N Min. Mean Max. V' σ
CLUZEL 18 0.92 1.020 1.11 18.7 0.052   17 0.88 0.980 1.10 22.2 0.054
CHAVR 33 0.98 1.030 1.12 13.3 0.057   33 0.88 1.000 1.10 22.2 0.057
LAUT2 26 0.98 1.080 1.16 16.8 0.047   28 0.93 1.030 1.09 15.8 0.049
SAULC 21 0.96 1.050 1.16 18.9 0.051   21 0.91 1.010 1.08 17.1 0.048
FORN11 16 0.92 1.016 1.10 17.8 0.055   16 0.88 0.959 1.06 18.6 0.058
VENINF 26 0.96 1.070 1.17 19.7 0.057   24 0.94 1.000 1.06 12.0 0.036
BF53 5 1.10 1.164 1.22 10.3 0.048   5 1.07 1.134 1.18 9.8 0.040
THZ 111 0.96 1.110 1.25 26.2 0.065   111 0.89 1.020 1.13 23.8 0.051
COD3 63 1.01 1.130 1.24 20.4 0.054   63 0.97 1.070 1.21 22.0 0.046
DIEU 26 1.00 1.10 1.20 18.2     26 0.97 1.08 1.20 21.2  
PARR 14 1.01 1.100 1.17 14.7     14 0.91 1.060 1.18 25.8  
SAY1 3 0.95 0.990 1.05 10.0     3 0.80 0.870 0.92 14.0  
HERR8 12 1.02 1.085 1.18 14.5     12 1.01 1.077 1.15 13.0  
PDF 27 0.90 1.010 1.12 21.8 0.067   25 0.88 0.970 1.08 20.4 0.053
PDES 16 0.95 1.040 1.12 16.4 0.050   16 0.88 0.960 1.02 14.7 0.036
GAIM 23 0.99 1.030 1.10 10.5 0.034   22 0.94 0.990 1.05 11.1 0.032
RDUBEY 17 0.88 1.000 1.08 20.4 0.054   17 0.84 0.951 1.05 22.2 0.051
VIV1 74 0.87 0.982 1.15 27.7 0.049   69 0.84 0.935 1.09 25.9 0.047
MIR2A 24 0.91 1.006 1.08 17.1 0.050   20 0.87 0.953 1.07 20.6 0.048
MIR1 43 0.87 1.007 1.08 21.5 0.046   44 0.84 0.967 1.10 26.8 0.054

 

M3 N Min. Mean Max. V' σ   N Min. Mean Max. V' σ
CLUZEL 18 0.68 0.780 0.85 22.2 0.050   18 0.76 0.840 0.90 16.9 0.036
CHAVR 15 0.66 0.740 0.86 26.3 0.055   15 0.71 0.820 0.90 23.6 0.047
LAUT2 20 0.71 0.780 0.90 23.6 0.048   19 0.78 0.850 0.94 18.6 0.049
SAULC 19 0.69 0.770 0.89 25.3 0.056   19 0.77 0.840 0.95 20.9 0.052
FORN11 7 0.72 0.774 0.82 13.0 0.034   7 0.80 0.836 0.90 11.8 0.038
VENINF 1   0.780         1   0.780      
BF53 5 0.92 0.950 1.02 10.3 0.040   5 0.96 1.022 1.12 15.4 0.067
THZ 87 0.67 0.790 0.90 29.3 0.044   87 0.65 0.800 0.89 31.2 0.045
COD3 13 0.63 0.750 0.79 22.5 0.048   13 0.68 0.760 0.86 23.4 0.053
DIEU 6 0.61 0.70 0.78 24.5     6 0.66 0.75 0.84 24.0  
PARR 2 0.79 0.830 0.87 9.6     2 0.83 0.890 0.95 13.5  
SAY1 1   0.670         1   0.730      
HERR8 11 0.74 0.824 0.89 18.4     11 0.82 0.882 0.94 13.6  
PDF 9 0.68 0.750 0.82 18.7 0.046   9 0.71 0.780 0.85 17.9 0.044
PDES 3 0.74 0.760 0.77 4.0 0.015   3 0.74 0.770 0.79 6.5 0.025
GAIM 3 0.63 0.720 0.85 29.7     2 0.82 0.860 0.89 8.2  
RDUBEY 4 0.64 0.738 0.80 22.2 0.068   4 0.72 0.773 0.81 11.8 0.041
VIV1 31 0.63 0.698 0.83 27.4 0.051   29 0.66 0.733 0.81 20.4 0.041
MIR2A 15 0.60 0.702 0.76 23.5 0.047   16 0.71 0.783 0.84 16.8 0.039
MIR1 37 0.63 0.700 0.77 20.0 0.035   37 0.70 0.772 0.86 20.5 0.038
 

freudentMatthijs Freudenthal. Departamento de Estratigrafía y Paleontología, Universidad de Granada, Avda. Fuentenueva s/n, E-18071 Granada, Spain; Naturalis Biodiversity Center, P.O. Box 9517, NL-2300 RA Leiden, The Netherlands. This email address is being protected from spambots. You need JavaScript enabled to view it.

Matthijs Freudenthal is a former curator of vertebrate paleontology of the National Museum of Natural History, Naturalis, Leiden, the Netherlands. He is actually working at the University of Granada, Spain. He is one of the pioneers of fossil micromammal research in Europe. His most recent research covers a variety of subjects like taxonomy of Gliridae, correspondance analysis in vertebrate paleontology, the insular faunas from Gargano (Italy) and body mass estimates in fossil rodents.

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martinElvira Martín-Suárez. Departamento de Estratigrafía y Paleontología, Universidad de Granada, Avda. Fuentenueva s/n, E-18071 Granada, Spain. This email address is being protected from spambots. You need JavaScript enabled to view it.

Elvira Martín Suárez is professor of vertebrate paleontology at the University of Granada, Spain. After completing her study in Biology, she specialized in the field of fossil vertebrates at the University of Lyon (France) and the National Museum of Natural History, Leiden, the Netherlands. Her main scientific interest are the fossil rodents from the Mio-Pliocene transition. 

 

TABLE 1. Species of Plesiosminthus described from the European Oligocene and Miocene.

Species Type-locality
P. schaubi Viret, 1926 (type-species) Coderet, France
P. myarion Schaub, 1930 Chavroches, France
P. promyarion Schaub, 1930 Puy-de-Montdoury, F rance
P. bavaricus Freudenberg, 1941 Gaimersheim, Germany
P. winistoerferi Engesser, 1987 Brochene Fluh 53, Switzerland
P. conjunctus Ziegler, 1994 Herrlingen 8, Germany
P. moralesi Álvarez Sierra, Daams and Lacomba Andueza, 1996 Sayatón 1, Spain
P. admyarion Comte, 2000 Thézels, France
P. meridionalis Comte, 2000 Venelles inf., France
P. margaritae n. sp. (this paper) Mirambueno 1, Spain
P. moniqueae n. sp. (this paper) Pech Desse, France
Plesiosminthus n. sp., Ziegler, 1994 Herrlingen 8, Germany
Plesiosminthus sp., Engesser and Mödden, 1987 Mümliswil-Hardberg, Switzerland
Plesiosminthus sp., Comte, 2000 Venelles inf., France

 

TABLE 2. Locality codes, species contents and most important reference of the localities studied. Zones are European mammal zones (Biochrom’97, 1997).

Code Locality Zone Species Reference
BF53 Brochene Fluh 53 MP30 P. winistoerferi Engesser (1987)
CANAL Canales MP28 Plesiosminthus sp. Alvarez Sierra et al. (1999)
CHAVR Chavroches MN2 P. myarion Hugueney and Vianey-Liaud (1980)
CLUZEL Cluzel MN2 P. myarion Hugueney and Vianey-Liaud (1980)
COD3 Coderet 3 MP30 P. schaubi Hugueney (1969)
DIEU Dieupentale MP30 P. schaubi Baudelot and Olivier (1978)
FORN11 Fornant 11 MN1 P. myarion Engesser (1987)
GAIM Gaimersheim MP28 P. bavaricus Kristkoiz (1992)
HERR8 Herrlingen 8 MP28 P. conjunctus Ziegler (1994)
LAUT2 Lautern 2 MN1 P. myarion Ziegler and Werner (1994)
MIR1 Mirambueno 1 MP27 P. margaritae this paper
MIR2A Mirambueno 2A MP27 P. aff. conjunctus this paper
OBL Oberleichtersbach MP30 P. winistoerferi Engesser and Storch (2008)
PARR Parrales MP29 P. schaubi Alvarez Sierra et al. (1999)
PDES Pech Desse MP28 P. promyarion Hugueney and Vianey-Liaud (1980)
PDES Pech Desse MP28 P. moniqueae this paper
PDF Pech-du-Fraysse MP28 P. promyarion Hugueney and Vianey-Liaud (1980)
RDUBEY Ruisseau du Bey MP28 P. promyarion Engesser (1987)
SAULC Saulcet MN1 P. myarion Hugueney and Vianey-Liaud (1980)
SAY1 Sayatón 1 MP29 P. moralesi Alvarez et al. (1996)
THZ Thézels MP30 P. admyarion Comte (2000)
VENINF Venelles inf. MP30 P. meridionalis Comte (2000)
VIV1 Vivel del Río 1 MP28 P. cf. margaritae this paper

 

TABLE 3. Frequencies of character states of the protolophule of M2, number of observations, and morphology values. Ant. = anterior, trans = transverse, ant.plus = anterior plus an incomplete posterior connection. * protolophule posterior, not double.

Sample ant. or trans ant. plus double n MV
CHAVR 0 3 97 33 0.985
FORN11          
VENINF 4 0 96* 25 0.960
THZ 95 0 5 97 0.050
COD3 97 0 3 63 0.032
HERR8 17 8 75 12 0.790
PDF          
GAIM 39 52 9 23 0.350
PDES 74 13 13 16 0.195
RDUBEY          
VIV1 57 4 39 69 0.410
MIR2A 19 7 74 27 0.775
MIR1 28 37 35 43 0.535

 

TABLE 4. P-values of t-test comparing the samples from THZ and COD3; df = degrees of freedom.

Molar Comte
(2000)
this paper df
m1 3.18 3.092 205
m2 2.02 -2.162 180
m3 3.00 2.828 127
M1 2.39 -2.483 188
M2 2.06 -2.178 172
M3 3.00 2.832 98

 

TABLE 5. Frequencies of character states of the protoconid hind arm of m2, number of observations, and morphology values. Data from Kristkoiz (1992, figure 84) and this paper.

Sample complete incomplete absent n MV
CHAVR 12 50 38 50 0.630
FORN11 26 52 22 27 0.480
VENINF 0 0 100 26 1.000
COD3 0 0 100 59 1.000
THZ 9 9 82 127 0.865
HERR8 82 18 0 17 0.090
PDF 36 53 11 28 0.375
GAIM 50 41 9 31 0.295
PDES 32 45 23 53 0.455
RDUBEY 31 69 0 13 0.345
VIV1 31 16 53 52 0.610
MIR2A 83 0 17 16 0.170
MIR1 60 26 14 41 0.270

 

TABLE 6. Frequencies of character states of the protoconid hind arm of m3, number of observations, and morphology values.

Sample complete incomplete absent n MV
CHAVR 5.4 0.0 94.6 37 0.946
VENINF 0.0 0.0 100.0 8 1.000
COD3 0.0 0.0 100.0 32 1.000
THZ 0.0 0.0 100.0 97 1.000
HERR8 0.0 0.0 100.0 14 1.000
PDF 8.0 32.0 60.0 25 0.760
GAIM 14.9 27.3 63.6 11 0.730
PDES 25.0 43.8 31.2 16 0.531
RDUBEY 0.0 0.0 100.0 6 1.000
VIV1 5.1 0.0 94.9 39 0.949
MIR2A 20.0 15.0 65.0 20 0.725
MIR1 23.2 11.6 65.1 43 0.710
 

TABLE 7. Number of specimens for samples in Figure 13.

Sample m1 m2 m3 M1 M2 M3
BF53 4 4 5 6 5 5
CHAVR 36 48 39 49 33 15
COD3 65 59 32 71 63 13
GAIM 34 32 11 28 23 3
HERR8 8 15 14 18 12 11
MIR1 17 41 45 49 43 37
PDES 57 54 16 35 16 3
SAY1 7 6 1 3 3 1
THZ 142 123 97 119 111 87
VENINF 19 25 8 27 26 1