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Author: Hungerbuhler, Axel Title: Cranial anatomy and diversity of the Norian phytosaurs of southwestern Germany.

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Your claim will be investigated and, where appropriate, the item in question will be removed from public view as soon as possible. Cranial anatomy and diversity of the Norian phytosaurs of Southwestern Germany

Volume 2 Appendices and Figures

by Axel Hungerbiihler

A thesis submitted to the University of Bristol in accordance with the requirements for the degree of Doctor of Philosophy in the Faculty of Science (Department of Earth Sciences), May 1998. Appendix A Data matrix

Appendix A

Data matrix

361 Table A.l: Data matrix nJa =character not applicable; 1 =missing data; (1) =score questionable because of preservational state; 0-1, 1-2, 2-3 =polymorphic character states.

Abbrev.: A., Angistorhinus; E., Ebrachosuchus; F., Francosuchus; M., Mesorhinosuchus; P., Paleorhinus; "Pa.", "Parasuchus"; Pr., Promystriosuchus. character Eu· Proto· E. F. M. P. P. magna· P. P. scurri· ·Pa. Pro A. A. parkeria champsa neukLlmi broili jraasi bransoni culus sawini ensis hislopi" ehlersi grandis taJainti 1 snout length 0 1 3 '1 '1 2 1 1 '1 2 3 2 1 2 ventral rim of m 0 0 0 0 1 0 0 1 0 0 0 0 1 I 3 pm and internasal septum nJa nJa 1 1 '1 '1 1 '1 0 1 1 1 1 4 lateral rim of m 0 1 1 0 0 0 1 0 0 0 0 0 0 5 nature of dentition 0 0 '1 '1 1 1 1 '1 1 1 '1 1 1 6 number of teeth 0 0 2 1 1 1 1 1 1 1 2 2 1 7 anterior extent of sm nJa nJa 1 0 1 1(1) 0 '1 1 0 ? 0 0 8 posterior extent of sm nJa nJa 1 1 '1 1 1 '1 1 1 '1 0 0 9 elevation of naris 0 0 0 0 0 0 0 0 0 0 0 0 0 10 naris position 0 0 1 1 2 1 1 1 1 1 2 3 3 11 infranasal recess 0 0 1 1 0 0 0 0 0 0 0 0 0 12 preorbitaI depression 0 0 0 0 0 0 0 0 0 0 0 0 0 Vl 13 length of prf 0 nJa 1 1 0 0 0 ? '1 1 '1 1 1 ~ 14 SCUlpture of skull roof 0 1 0 0 0 0 0 0 0 0 0 0 0 15 pre·infratemporal shelf 0 0 0 0 0 0 0 0 0 0 0 0 0 16 size of infratemporal fen. 0 1 2 1 1 1 0 1 1 1 1 2 2 17 anterior process of j 0 1 0 0 0 0 0 0 0 0 0 1 1 18 height of j 0 1 1 1 '1 2 2 1 1 1 2 1 1 19 j-qj Suture 0 0 1 1 0(1) 1 '1 1 1 0 1 1 1 20 length I width of po·sq bar 0 2 0 2 1 1 1 1 1 1 1 0 1 211ength of par 0 1 0 0 0 0 0 0 0 0 0 1 1 22 posterior extent of par 0 0 0 1 0 1 1 1 1 0 1 0 0 23 par·soc complex 0 0 0 0 0 0 0 0 0 0 0 0 0 i....0- 24 overhang of par 0 1 0 0 1 1 '1 1 1 1 1 2 2 >< 25 border supratemporal fen. 0 0 0 0 0 0 0 0 0 0 0 0 0 > tj e Pl 3 ....~ >< Table A.1: Data matrix (continued) nla = character not applicable (0, 1); ? =missing data; (?) =score questionable because of preservational state; 0-1, 1-2,2-3 =polymorphic character states.

Abbrev.: A., Angistorhinus; E., Ebrachosuchus; F., Francosuchus; M., Mesorhinosuchus; P., Paleorhinus; "Pa.", "Parasuchus"; Pr., Promystriosuchus. character Eu- Proto- E. F. M. P. P. magna- P. P. scurri- "Pa. Pro A. A. parkeria chilmpsa neukami broili fraasi bransoni culus sawini ensis his/opi" ehlersi grandis talainti 26 dorsal groove of sq 0 0 1 1 ? I 1 1 I 1 1 0 0 27 lateral ridge of sq 0 1 I 1 I I 1 I I 1 I 0 1 28 medial lamella of sq 0 0 ? ? ? 0 7 0 0 0 0 0 2 29 vertical part of sq 0 0 0 0 0 0 0 0 0 0 0 I I 30 posterior process of sq 1 0 0 1 ? I 1 (7) 0 0 0 0 2 2 31 extent of sq 0 0 0(7) I (7) 7 I 1 0 I 0 I I 1 32 shape of tip of sq 0 1 1 0 ? 0 1 0 0 0 0 1 1 33 depression of par-sq bar 0 0 0 0 0 0 0 0(7) 1 0 0 0 0 34 shape of par-sq bar 0 0 0 0 0 0 0 7 0 0 0 1 1 35 quadratojugal process of sq 0 0 1 1 7 I 0 7 1 1 0 1 1 36 paroccipital process of sq 0 0 0 0 ? I 0 0 1 1 0 1 1 37 soc shelf 0 0 1 1 1 0 1 0 1 0 I 0 0 w 38 soc I posttemporal fenestra 0 nla ? 0 ? ? 7 7 0 0 1(7) 1 0 ~ 39 paroccipital process of opo 0 0 0 0 0 ? 7 7 0 0 0 1 1 40 size of quadrate foramen 0 nla 0 0 ? 0 7 0 1 1 1 (7) 1 1 41 location of quadate foramen 0 nla 1 1 ? 0 0 0 0 1 1 1 1 42 height of posttempora! fen. 0 nla 0 0 1 (7) 0 7 ? 0 0 1 (7) 0 1 43 pm reaching choana 0 7 0 7 1 1 1 7 0 1 I 1 0 44 anterior extent of pal 0 0 1 1 1 3 (7) 3 7 1 3 3 (7) 3 (7) 2 45 medial extension of pal nla nla 0 0 7 1 0 7 1 0 0 0 0 46 pal meet in midline 0 0 0 0 7 0 0 7 0 0 0 0 0 47 suborbital fenestra 0 0 0 0 0 2 7 7 1 0 2 1 2 48 interpterygoid vacuity 0 0 0 0 1 2 7 7 2 2 0 0 2 ie: >< 49 position of nervus XU 0 7 7 0 7 ? 7 7 7 0 7 7 7 > o S g Ei- >< Table A.l: Data matrix (continued)

? =missing data; (?) =score questionable because of preservational state; 0-1, 1-2,2-3 = polymorphic character states.

Abbrev.: Ar., Arribasuchus; B., Brachysuchus; L., Leptosuchus; My., Mystriosuchus; N., Nicrosaurus; Ps., Pseudopalatus; R., Rutiodon; S., Smilosuchus. character B. L L s. R. N. N. Ar. Ps. My. My. megalodon crosbiensis adamanensis gregorii carolinensis kapjJi species B buceros pristinus planirostris species B I snout length I 1 1 1 2 1 2 1-2 2 3 2 2 ventral rim of m 1 0 0 1 0 1 0 1 0 0 1 3 pm and internasal septum 1 1 1 1 ? 1 1 1 1 1 1 4 lateral rim of m 0 0 0 0 0 0 0 0 1 1 1 5 nature of dentition 1 2 2 2 1 2 1 2 1 1 1 6 number of teeth 1 1 1 1 1 1 1 1-2 2 2 2 7 anterior extent of sm 0 1 0 1 ? 1 0 1 1 0 0 8 posterior extent of sm 0 0 0 0 ? 0 0 0 0 0 0 9 elevation of naris 0 0 0 0 0 0 0 1 1 0 0

10 naris position 3 3 3 3 3 3 3 3 3 3 3 I 11 infranasal recess 0 0 0 0 0 1 1 0 0 0 0 12 preorbital depression 0 0 0 0 0 0 0 0 0 1 1 VJ 13 length of prf 1 1 1 1 1 1 1 1 1 1 1 ~ 14 sculpture of skull roof 0 0 0 0 0 0 0 0 0 1 1 15 pre-infratemporal shelf 0 0 0 0 0 0 0 1 1 2 2 16 size of infratemporal fen. 2 2 2 2 1 2 2 2 2 2 2 17 anterior process of j 0 0 0 0 0 0 0 0 0 0 0 18 height of j 1 1 1 2 1 1 0 1 0 0 0 19 j-qj suture 1 1 1 1 1 1 1 1 1 1 1 20 length I width of po-sq bar 0 I 1 1 1 2 2 2 2 2 2 21 length of par 1 0 0 0 0 0 0 0 0 0 0 22 posterior extent of par 0 1 1 1 1 1 1 1 1 1 1 :g o 23 par-soc complex 0 0 0 0 ? 0 0 1 1 1 1 ::l ~ 24 overhang of par 2 1 1 1 1 1 1 2 2 2 2 >= 25 border supratemporal fen. 0 0 0 0 0 0 0 0 0 1 1 :> o a ~

~.>= Table A.1: Data matrix (continued)

? =missing data; (?) =score questionable because of preservational state; 0-1, 1-2,2-3 = polymorphic character states.

Abbrev.: Ar., Arribasuchus; B., Brachysuchus; L., Leptosuchus; My., Mystriosuchus; N., Nicrosaurus; Ps., Pseudopalatus; R., Rutiodon; S., Smilosuchus. character B. L L s. R. N. N. Ar. Ps. My. My. megalodon crosbiensis adamanensis gregorii' carolinensis kapffi species B buceros pristinus planirostris species B 26 dorsal groove of sq 0 0 0 0 0-\ 0 0 0 0 0 0 27 lateral ridge of sq 0 0 0 0 1 0 0 0 0 1 1 28 medial lamella of sq 0 0 1 0 I 2 2 3 3 1 I 29 vertical part of sq 0 I 0 1 0 0 0 0 0 0-1 0 30 posterior process of sq 1 3 2 2 2 2 2 3 3 1 0 31 extent of sq 0 I I 1 I I I 1 I 0 0 32 shape of tip of sq I 0 1 0 1 1 0 0 0 0 0 33 depression of par-sq bar 0 2 2 2 2 2 2 3 3 3 3

34 shape of par-sq bar 1 0 0 0 0 0 0 0 0 0 0 I 35 quadratojugal process of sq I 1 I 0 I 0 1 0 I I 1 36 paroccipital process of sq 1 I I I 1 I I 1 I 1 1

37 soc shelf 0 1 I I I 2 2 2 2 I 1 i VJ 38 soc I posttemporal fenestra ? 0 1 1 ? 0 0 ? 1 0 0 8i 39 paroccipital process of opo 1 I 1 1 ? 1 1 1 1 1 1 40 size of quadrate foramen 1 1 I 1 ? 1 I 1 I 0 0 41 location of quadate foramen 1 1 1 1 ? 1 1 I I I I 42 height of posttemporal fen. 0 0 0 0 ? 0 0 0 0 I I 43 pm reaching choana 0 0 I 0-1 0 I I ? ? 0 I 44 anterior extent of pal 3 3 2 2 3 3 3 3 ? 3 3 45 medial extension of pal 0 I 0 0 0 I I 0 ? 0 0 46 pal meet in midline 0 ? 0 0 0 I I 0 ? 0 0 47 suborbital fenestra 0 2 I 2 I I I 2 2 2 2 48 interpterygoid vacuity 2 I I 2 0 2 2 2 I 2 2 i ~ 49 position of nervus XII 0 ? ? 0 ? I I ? ? ? 0 >< --- -- "------» t::I ~ ~ 3 x·~ Appendix B Data collection

Appendix B Data collection

366 Appendix B Data collection

taxon ratio preorbitall orbital ratio prenariall narial + source + postorbital length postnariallength Ebrachosuchus 3.8 1.59 KUHN (1936: fig. 4) neukami Paleorhinus 3.2 1.09 LEES (1907: fig. 1) bransoni MURRY (1986) Paleorhinus 2.7 1.12 DUTUIT (1977a: fig. lA) magnoculus Paleorhinus 2.6 1.01 LONG & MURRY (1995: fig. 24F) sawini MURRY (1986) "Parasuchus 3.4 1.09 CHATTERJEE (1978: fig. 2) hislopi" Promystriosuchus 3.7 1.64 CASE (1922: pI. II fig. A) ehlersi Angistorhinus 3 1.57 MEHL (1915: fig. 3) grandis Angistorhinus 2.4 1.16 DUTUIT (1977a: fig. 2C) talainti Brachysuchus 2.9 1.28 CASE (1929: fig. 1) megalodon Leptosuchus 2.7 1.15 - 1.22 CASE (1922: fig. 25B) crosbiensis CASE & WHITE (1934: fig. 2) Leptosuchus 2.4 1.4 CAMP (1930: pI. 1) adamanensis Smilosuchus 2.4 - 2.5 1.1 - 1.26 CAMP (1930: pI. 6) gregorii COLBERT (1947: pI. 4 fig. 1) Rutiodon 3 1.5 - 1.59 GREGORY (1962a); carolinensis GREGORY (l962a: fig. 8) Nicrosaurus 2.44 - 2.74 1.15 - 1.34 pers. obs. (SMNS 4379. kapffi SMNS 5726. SMNS 5727) Nicrosaurus 3.41 1.63 pers. obs. (SMNS 12593) species B Pseudopalatus 3 - 3.2 1.6 - 1.64 LONG & MURRY (1995: fig. 40A) pristinus MEHL (1928b: pI. I fig. A) Arribasuchus 2.5 1.14 holotype (Huene 1915: fig. 11) buceros 2.8 - 3.3 1.36 - 1.53 BALLEW (1989: pI. 8 fig. A) MEHL (1922: figs. 1.3) Mystriosuchus 3.9 - 4.1 2.6 MCGREGOR (1906: pI. 6 fig. 1) planirostris pers. obs. (SMNS 13240) Mystriosuchus 3.3 2.2 pers. obs. (GPIT 261/001) species B

Table B.I: Data for character 1: ratios of the snout (preorbital skull) length and the rostral (prenarial skull) length.

367 Appendix B Data collection

taxon range of tooth no. of source positions specimens Ebrachosuchus neukami 56-58 1 KUHN (1936) Francosuchus sp. 43 1 KUHN (1936) Paleorhinus bransoni 36 1 GREGORY (1962a) Paleorhinus magnoculus 43-47 1 DUTIJIT (1977b) Paleorhinus sawini 39 1 GREGORY (1962a) "Parasuchus" hislopi 45 1 CHATIERJEE (1978) Promystriosuchus ehlersi 47 1 CASE (1922) Angistorhinus grandis 42 1 GREGORY (1962a) Angistorhinus talainti 40-41 1 DUTIJIT (1 977a) Brachysuchus megalodon 46 1 GREGORY (1962a) Leptosuchus crosbiensis 39-45 7 GREGORY (1962a) Leptosuchus adamanensis 40-43 1 CAMP (1930) Smilosuchus gregorii 37-41 3 CASE & WHITE (1934); GREGORY (l962a); Rutiodon carolinensis 42 1 GREGORY (1962a) Nicrosaurus kapffi 34-45 5 pers. obs. (Tab. 2.2.3) Nicrosaurus species B 39 2 pers. obs. (SMNS 4059, BMNH42745) Arribasuchus buceros 41-47 3 CAMP (1930); GREGORY (1962a); MEHL (1922) Pseudopalatus pristinus 48-50 1 GREGORY (1962a) Mystriosuchus planirostris 48 1 MCGREGOR (1906) Mystriosuchus species B 50 1 pers. obs. (GPIT 2611001)

Table B.2: Data for character 6: range and variation in the number of tooth positions. The third column indicates the number of specimens on which the data is based.

368 Appendix B Data collection

taxon jugal height I skull height source Francosuchus angustifrons 0.29 KUHN (1936: pI. 10 fig. 5) Paleorhinus bransoni 0.31 LEES (1907: fig. 3) Paleorhinus magnoculus 0.38 DUTUIT (1977a: fig. 2) Paleorhinus sawini 0.29 LONG & MURRY (1995: fig. 24B) "Parasuchus" hislopi 0.21 CHATIERJEE (1978: fig. 3) Promystriosuchus ehlersi 0.37 CASE (1922: pI. 11 fig. B) Angistorhinus grandis 0.25 MEHL (1915: fig. 3) Angistorhinus talainti 0.21 DUTUIT (1 977a: fig. 2C) Brachysuchus megalodon 0.26 CASE (1929: fig. 3) Leptosuchus crosbiensis 0.19 - 0.22 CASE (1922: fig. 25A) CASE & WHITE (1934: pI. 1) Leptosuchus adamanensis 0.25 CAMP (1930: pI. 2 fig. a) Smilosuchus gregorii 0.34 - 0.36 CAMP (1930: pI. 5) COLBERT (1947: pI. 3 fig. 1) Rutiodon carolinensis 0.25 COLBERT (1947: fig. 7) Nicrosaurus kapffi 0.19 - 0.28 pers. obs. (SMNS 4378. 4379. 5726. 5727, BMNH 42743) Nicrosaurus species B 0.18 pers. obs. (SMNS 12593. BMNH 42745) Arribasuchus buceros 0.20 - 0.27 CAMP (1930: fig. 4d) LONG & MURRY (1995: fig. 410) MEHL (1922: fig. 1) Pseudopalatus pristinus 0.17 - 0.20 BALLEW (1989: pI. 6 fig. C) LONG & MURRY (1995: fig. 41C) MEHL (l928b: pI. 1 fig. C) Mystriosuchus planirostris 0.12 MCGREGOR (1906: pI. 7) Mystriosuchus species B 0.14 pers. obs. (GPIT 2611001)

Table B.3: Data for character 18: ratio of the minimum height of the jugal at the jugal quadratojugal bar to the total skull height in this area.

369 Appendix B Data collection

taxon length I width postorbito- source squamosal bar Ebrachosuchus neukami 9.4 KUHN (1936: pI. 8 fig. Ie) Francosuchus angustifrons 3.6 KUHN (1936: pI. 9 fig. 2b) Paleorhinus bransoni 7.2 LEES (1907: fig. 1) 5 LONG & MURRY (1995: fig. 24F) Paleorhinus magnoculus 4.4 DUTUIT (1977a: pI. 1) Paleorhinus sawini 7 LONG & MURRY (1995: fig. 24C) Paleorhinus scurriensis 7.3 LANGSTON (1947: fig. 1) "Parasuchus hislopi" 5.3 CHATTERJEE (1978: figs. 2, 3) Promystriosuchus ehlersi 7.9 CASE (1922: pI. 11) Angistorhinus grandis 10 MEHL (1915: fig. 3) Angistorhinus talainti 5 DUTUIT (1977a: fig. 2C) Brachysuchus megalodon 10 CASE (1929: figs. 1,3) Rutiodon carolinensis 8 GREGORY (1962b: fig. 8) Leptosuchus crosbiensis 7 CASE (1922: fig. 25B) 6.5 CASE & WHITE (1934: fig. 2) Leptosuchus adamanensis 4.4 CAMP (1930: pI. 1) Smilosuchus gregorii 4.2 CAMP (1930: pI. 6) Nicrosaurus kapffi 3.1 SMNS 4379 3.05 SMNS 5726 2.85 SMNS 5727 Nicrosaurus species B 2.35 SMNS 12593 3 BMNH42745 Pseudopalatus pristinus 2.1 MEHL (l928b: pI. 1) Arribasuchus buceros 2.5 HUENE (1915: fig. 11) Mystriosuchus planirostris 3.7 MCGREGOR (1906: pIs. 6, 7) Mystriosuchus species B 3.8 GPIT 2611001

Table B.4: Data for character 20: ratio of the length to width of the postorbito squamosal bar.

370 Appendix B Data collection

taxon depression source in % of skull height Leptosuchus crosbiensis 14-17% BALLEW (1989: fig. 2A) CASE & WHITE (1934: fig. 3) Leptosuchus adamanensis 26% CAMP (1930: pI. 2 fig. e) Smilosuchus gregorii 21-23% BALLEW (1989: fig. 2B) CAMP (1930: pI. 5) COLBERT (1947: pI. 7 fig. 3) Nicrosaurus kapffi 19-27% pers. obs. (SMNS 4378, 4379, 5726, 5727, BMNH 42743) Nicrosaurus species B 21% pers. obs. (SMNS 12593, BMNH 42745) Pseudopalatus pristinus 26-30% BALLEW (1989: fig. 2D) MEHL (l928b: fig. 4) Arribasuchus buceros 26-30% BALLEW (1989: fig. 2C) HUENE (1915: fig. lld) Mystriosuchus planirostris 31-38% MCGREGOR (1906: pI. 8 fig. 4) pers. obs. (SMNS 10460, 13007, 13240, uncat. no. 184, 185) Mystriosuchus species B 26% pers. obs. (GPIT 2611001)

Table B.5: Data for character 33: depression of the parieto-squamosal bar below the skull roof, indicated as a percentage of absolute skull height in occipital view.

371 Appendix B Data collection

taxon length of the quadratojugal source process of the squamosal Ebrachosuchus neukami 80-90% KUHN (1936: fig. 4) Francosuchus 80-100% KUHN (1932: fig. 6, 1936: fig. 1) Paleorhinus bransoni 70-80% LEES (1907: fig. 3) Paleorhinus magnoculus 0% DUTUIT (1977a: fig. 2) Paleorhinus scurriensis >50% LANGSTON (1949: fig. 1) "Parasuchus" hislopi 80-100% CHATTERJEE (1978: figs. 3,5) Promystriosuchus ehlersi <10% CASE (1922: pI. 11 fig. B) Angistorhinus grandis 60% MEHL (1915: fig. 3) Angistorhinus talainti 70% DUTUIT (1977a: fig. 2C) Brachysuchus megalodon 50-60% CASE (1929: fig. 3) Leptosuchus crosbiensis 70-90% CAMP (1930: fig. 9) CASE (1922: fig. 25A) CASE & WHITE (1934: pI. 1) Leptosuchus adamanensis 60-80% CAMP (1930: fig. 4a-c, 5b, pI. 2 fig. a) Smilosuchus gregorii 20-35% CAMP (1930: pI. 5) COLBERT (1947: pI. 3 fig. 1) Rutiodon carolinensis 90% DOYLE & SUES (fig. 1) Nicrosaurus kapffi 30-50% pers.obs. Nicrosaurus species B >50% pers.obs. Arribasuchus buceros 50% MEHL (1922: fig. 1) Pseudopalatus pristinus 75% CAMP (1930: fig. 4d) Mystriosuchus planirostris 90-100% pers.obs. Mystriosuchus species B 90-95% pers.obs.

Table B.6: Data for character 35: length of the quadratojugal process of the squamosal, measured as a percentage of the distance it borders the infratemporal fenestra posteriorly in lateral view.

372 Table B.7: Data for character 46: variation in the suborbital fenestra in phytosaurs.

Indicated are column 1: the total number of the openings present, in the case of taxa scored as (2) followed in brackets by the traits 2a to 2c as defined in the text, column 2: the shape and width of the single, or alternatively the posterior, fenestra, in the case of the taxa with a single opening followed in brackets by the score, column 3: contribution of the maxilla to the single (or alternatively the posterior) fenestra, column 4: the bones surrounding the anterior fenestra when two fenestrae are present.

Abbrev.: ect, ectopterygoid; m, maxilla; pal, palatine; pm, premaxilla; pt, pterygoid; taxon number single elongate single or posterior: single or posterior: anterior foramen: remarks foramen: width foramen: shape foramen: +m configuration Ebrachosuchus neukami I less broad (1) elongate +m - - Francosuchusangustijrons I less broad (1) elongate +m - - Mesorhinosuchus jraasi I very broad (0) elongate +m - pt contributes to choana (7) ! Paleorhinus bransoni I (2a) - oval no - - . Paleorhinus scurriensis I less broad (I) elongate +m - - "Parasuchus" hislopi I less broad (1) elongate no - - Promystriosuchus ehlersi 1 (2a) - foramen +m - pt contributes to choana! I Angistorhinus grandis I less broad elongate +m - pm contributes to choana! Angistorhinus talainti none (2c) - - - - - Brachysuchus megalodon 1 less broad (1) elongate +m - - w Leptosuchus crosbiensis 1 (2a) - oval no - - ....:Jw Leptosuchus adamanensis 1 slit-like (1) elongate no - ROMER (1956: fig. 7lC) illustrates two fenestrae Smilosuchus gregorii 1 (2a) - oval no - - Rutiodon carolinensis 1 slit-like (1) elongate +m - - Nicrosaurus kapjJi 1 slit-like (1) elongate +m - ontogenetically subdivided into 2 or 3 fenestrae ~ Nicrosaurus species B 1 slit-like (1) elongate +m - - ~ Arribasuchus buceros 2 (2b) oval no m/pal/ect - 8- - ~. Pseudopalatus pristinus 1 (2a) - oval no - MEHL (1922: fig. 3) shows 2 ttl fenestrae t:j Mystriosuchus planirostris ~ 1 (2a) - oval no - number not verified s:>:> (') Mystriosuchus species B 2 (2a) - oval no m/pal/ect + small foramen pal/ect ~ gp. Appendix C Localities

Appendix C The phytosaur-bearing localities and their stratigraphic position

The localities in the Stubensandstein from Wtirttemberg that have yielded identifiable phytosaur remains are localised here or, in the cases of insufficient or conflicting data, their most likely topographic position is discussed. The sites are treated in geographical order from north to south. Illustrations are based on the topographic and the corresponding geological maps, identified by their four digit sheet number (Topographische Karte [TK] 1:50 000 and 1:25 000, and Geologische Karte von Baden Wtirttemberg 1:25000, both series published by Landesvermessungsamt Baden Wtirttemberg). The location of a site is determined as accurately as possible, given in Gauss-KrUger coordinates according to the German grid system. The stratigraphic position of each site is discussed and determined as exactly as possible. In several cases, data from the literature were cross-checked in the field, and the sources of data (publications, personal field observations) are indicated. The identified phytosaur material from each site, most of which is referred to in the descriptive part of this study, is listed. Phytosaur remains, which are described or mentioned in the literature but which can not be located in a collection at present, are usually not included. Additionally, indeterminable phytosaur material is included in those cases when there is sufficient evidence that the listed skeletal elements are derived from the same site and approximately the same beds, and thus are likely to represent a contemporaneous association.

1. Affaltrach near Lowenstein

TK 1:25 000: sheet 6922 Lowenstein. Site locations: unknown.

An attempt has been made to sort out the complex history of finds from Lowenstein in northeastern Wiirttemberg, which include the syntypes of Belodon plieningeri and the historically important von HUGEL-collection. Finds in the LOwenstein region have evidently been made several times in the last century and probably at different locations, and were incorporated in, or subsequently passed on to, a number of private and public

374 Appendix C Localities

collections. It is necessary to distinguish among these various finds, which in the past have often been confused and are commonly (but incorrectly) treated as a single association.

1.1 Collections

1.1.1 The syntypes of Belodon plieningeri MEYER, 1844

In the 1840s, the specimens were kept in the collection of the "Centralstelle des landwirtschaftlichen Vereins" (MEYER in MEYER & PLIENINGER 1844: 43). This collection came under the auspices of the "Verein ftir vaterUindische Naturkunde" in 1850, and was finally merged with the collection of the later SMNS in 1864 (ZIEGLER 1986). According to official documents, a small, representative part of the collection was also given to Ttibingen (M. WARTH, SMNS, pers. comm., 1994). However, it remains obscure what happened to the type material of Belodon plieningeri. It appears from the published records that no researcher actually studied, or was able to examine, the original type material after the specimens had been studied by MEYER and PLIENINGER (MEYER'S drawings published later 1847-55 were probably made when he borrowed the material in 1842). The specimens are evidently not present in either the SMNS nor the GPIT, and their present repository is unknown.

1.1.2 The von HUGEL collection

The fossils collected by Freiherr von HUGEL around the year 1850 (KURR 1852) represent the first substantial collection of vertebrates from the Stubensandstein of Wtirttemberg. PLIENINGER (1857: 405) had the whole assemblage at his disposal until at least 1856. However, except for the brief listing in PLIENINGER (1857), the majority of von HUGEL's specimen have never been described. The only two specimens, which have been figured and thus can be identified with certainty, were acquired in 1862 by the SMNS as part of the von KAPFF collection (SMNS archives). KAPFF must have acquired them at least two years earlier (MEYER 1860b). The present whereabouts of the remaining collection is unknown. Since von HUGEL's collection is of great historical importance, I include below a list of the specimens which were once part of it, based on PLIENINGER (1857: 405-415) and my own records.

375 Appendix C Localities

• SMNS 54708, Nicrosaurus kapffi, anterior part of a left premaxilla. • SMNS uncat. no. 8, cf. Nicrosaurus species B or Mystriosuchus pianirostris, fragment of the posterior symphyseal part of a right mandible (PLIENINGER 1857: pI. 8 fig. 2; MEYER 1861: pI. 29 figs. 8-10). • cf. Nicrosaurus kapffi, part and counterpart of a very fragmentary upper jaw (PLIENINGER 1857: pI. 8 figs. 3-4). • series of rib fragments, among them one ?cervical rib (p. 490). The series includes also four gastralia found in parallel orientation, one of which is complete (p. 492). • several vertebrae, all deformed; among them one centrum with transverse processes preserved and one neural arch (p. 481). • 10 osteoderms, with "edge-like transverse ridges" (p. 406, my translation); among them eight osteoderms of either trapezoidal shape with keels or rhomboidal unkeeled scutes (p. 514). The description does not match aetosaur scutes but rather suggests phytosaurian osteoderms, and PLIENINGER noted the similarity to scutes of Mystriosuchus from the ESER collection. • several limb bones, among them one humerus (p. 509) and three femoral fragments (pp. 509-510). • several cranial pieces showing a sculptured surface, which were only partially freed from their hematitic coating; among them possibly one surangular (p. 470-71). • several (at least two) small, bicarinate, and serrated tooth crowns and one small recurved, conical tooth crown (the latter might correspond to a crown labelled "Lowenstein", SMNS uncat.).

1.1.3 Specimens in the GPIT collection

The GPIT also keeps a small collection of vertebrate remains from Lowenstein. Specimens once housed in the collections of the "Landwirtschaftlicher Verein" cannot be identified any more. It is also possible, that some of von HUGEL's specimens ended up in Ttibingen. The most likely option is, however, that the specimens (or some of them) listed below were acquired by QUENSlEDT in the course of surveying the geological map sheet LOwenstein.

• GPIT 2093,001, phytosaur cf. Nicrosaurus species B or Mystriosuchus pianirostris, mandibular fragment • GPIT 2093,002, indeterminable bone fragment • GPIT 2140,000, long splint-like bone fragment • GPIT 2202,000, Paratypothorax, paramedian osteoderm • GPIT 2204,000, Paratypothorax, large paramedian osteoderm

376 Appendix C Localities

• GPIT 2205,000, Paratypothorax, paramedian osteodenn • GPIT 2206,002, aetosaur, two fragments of osteoderms • GPIT 2207,008, aetosaur, eight fragments of osteodenns; from collection 1. C. L. HEHL (1774-1853, 1806 to 1817 mining inspector, 1840 to 1852 curator ofthe mineralogical collection at the SMNS)

1.2 Localities and stratigraphy

Lowenstein, and the surrounding LOwensteiner Berge, has a longstanding history of quarrying of the Stubensandstein. Only a little contemporaneous infonnation about the localities (and consequently their stratigraphic position) has been recorded in print. The type locality of Belodon plieningeri is given as "Lowenstein" (MEYER & PLIENINGER 1844: 43, 103), and so read the specimen labels of all the GPIT specimens. This information is too vague to allow an identification, and thus BENTON & WILD (1993) suggest a number of possible sites. These include sites on the Wohlfahrtsberg close to the town centre of Lowenstein, for example, which were already in operation in 1827 (JAEGER 1827), and the quarries are still well exposed nowadays. It is safe to say that all specimens from the von HUGEL collection came from one single site (PLIENINGER 1857: 450,451). The quarry was not at Lowenstein itself but from the surroundings of the village Affaltrach a few kilometres to the north of LOwenstein (KURR 1852). The localities in BENTON & WILD (1993) are all centred around LOwenstein, and are therefore probably too far south. There are no former quarries indicated on the map in the close vicinity of Affaltrach.

Based on a note in QUENSTEDT (1874), BRENNER (1973) refers all "Lowenstein" fossils to the upper Stubensandstein without specifying the specimens. This is in accord with KURR's (1852) stratigraphical comment on the von HUGEL collection. However, these old stratigraphic determinations have to be treated with caution, since at that time the knowledge of Stubensandstein stratigraphy was poor, and conflicting, non-standardised terminology was widely used (see LANG 1909). Moreover, PLIENINGER (1857: 406), the only person who worked on the collection, was evidently ignorant about even the broad stratigraphic derivation ofthe Affaltrach specimens. BENTON & WILD (1993) list three Nicrosaurus in the SMNS as being derived from the lower Stubensandstein, which according to the context includes not only the two surviving von HUGEL specimens but also a phytosaur clavicle found as early as 1836 also near Affaltrach (SMNS 60, WILD 1991), but not necessarily at the same site. Since the provenance of all the Lowenstein Mfaltrach specimens is unknown, any claims about their stratigraphic derivation are speculative and must be considered unsubstantiated. Finally, ENGEL mentions that "such

i Or" BRISTOL 377 LIBRARY Appendix C Localities

scutes [Phytosaurus, i.e. most likely what is now known to be aetosaur osteoderms] occur in the red mudstones below the Stubensandstein" (1908: 170, my translation), Le. probably the Basisletten of the lowermost Stubensandstein (BRENNER 1973), but this does not necessarily apply to the types of Belodon nor the phytosaur remains from Affaltrach.

2. Stromberg Mountains

TK 1:50000: sheet L6918 Bretten (overview). TK 1:25 000: sheet 6919 Giiglingen.

WeiBer Steinbruch, also named quarry BURRER, Pfaffenhofen, R34976 H54320

Phytosaur remains in the Stromberg Mountains were found at several localities: near Ochsenbach in the quarry IpPICH (c. R349855 H543180; lower Stubensandstein; LINCK 1968), in the so-called "Ceratodus"-quarry (R3497625 H5432000; lower Stubensandstein; LINCK 1968) and an unspecified locality in the 4th Stubensandstein (= upper Stubensandstein, BERCKHEMER 1938). Only the WeiBer Steinbruch to the southeast of Pfaffenhofen yielded identifiable remains. This quarry is more famous for the abundant prosauropod dinosaurs, and some theropod and sphenosuchian remains (HUENE 1907/08, 1932; SERENO & WILD 1991). BRENNER (1973: 157) and BENTON & WILD (1993) also mention a phytosaur skull from Ochsenbach (quarry IpPICH), but no such skull is present in the SMNS collection. The WeiBer Steinbruch covers a huge area of about two hectares, and the rocks worked are middle Stubensandstein (BRENNER 1978a). A brief history of the quarry can be found in SERENO & WILD (1991). The site was abandoned in 1914 and is still potentially accessible; however, intrusive geological field work is currently difficult, since the overgrown quarry cliffs are now under environmental protection. The phytosaurs came exclusively from the once targeted sandstone bed about four metres in thickness ("Werkstein") just at the base ofthe quarry (BERCKHEMER 1938; FRAAS 1913b, 1914; LINCK 1968), which also yielded temnospondyls (FRAAS 1913c). The bed is identified as sandstone scI of the middle Stubensandstein (STOLL 1929). The prosauropods and other cursorial vertebrates were mostly confined to the overlying sequence of mud- and siltstones interbedded with thinner dolomitic limestone and sandstone beds ("Upper Faule" with "Brecciensandstein").

378 Appendix C Localities

List of specimens: • SMNS 12593, Nicrosaurus species B • SMNS 12593, 5 phytosaur teeth • SMNS 12593/2, Nicrosaurus species B, skull • SMNS 1267114, cf. Mystriosuchus planirostris, tip of snout • SMNS uncat., Mystriosuchus planirostris, lower jaw • SMNS 12594, osteoderm • SMNS 12595, osteoderm • SMNS 1267111-3, 3 articulated strings of vertebrae, found close together on the same bedding plane (HUENE 1922) • SMNS 12986, articulated pelvis and femora • SMNS uncat. no. 142, scapulocoracoid

3. Murrhardt

TK 1:25 000: sheet 7023 Murrhardt.

Quarry SCHLIPF, (?) R35414 H54257.

In 1945, the articulated, almost 2 m long skeleton of a phytosaur was recovered in the quarry SCHLIPF from the so-called "Fleins" (BERCKHEMER 1949). The specimen is reposited at the SMNS and tentatively referred to Nicrosaurus species B (R. WILD, pers. comm., 1994), pending its final preparation. BENTON & WILD (1993) give the location as R35414 H54257. The Fleins is the term used by local quarrymen for the usually calcareous cemented, commercially workable beds of the lower Stubensandstein. Thus, the skeleton is likely to be derived from the lower part of the Stubensandstein. It must be noted, however, that the Fleins is often locally subdivided into two distinct beds (Hauptfleins and Upper Fleins, EISENHlIT 1971). The latter stratum already represents the base of the middle Stubensandstein (BRENNER 1973: tab. 1), and for this reason, the stratigraphic referral must be treated with some caution.

379 Appendix C Localities

4. Stuttgart-Heslach and Stuttgart-Kaltental

TK 1:50000: sheet L7320 Stuttgart-Slid (overview). TK 1:25 000: sheet 7220 Stuttgart-Slidwest.

Quarry "Billert": ridge top above EIsenbach Valley, R3509760 H5409760 (WILD 1989) or floor ofthe EIsenbach Valley, R3509800 H5400825 (STROBEL & WURM 1977). Cliff "Heslacher Wand": R35 1058 H540260. Quarry "Kohlhau": R35 105 H54007.

Locating the former sites at Heslach and Kaltental in the vicinity of Stuttgart is of outstanding importance for the stratigraphic assessment of Stubensandstein reptiles. These quarries have not only yielded the phytosaur material collected by KApPF between 1859 and 1876, including almost all specimens of Nicrosaurus kapfji, but also the types of the prosauropod Sellosaurus gracilis, the aetosaurs Aetosaurus ferratus and Paratypothorax andressi, and all remains known of the rauisuchian Teratosaurus suevicus. Unfortunately, contemporaneous sources concerning the exact provenance of these finds are rather vague, and identification must largely rely on circumstantial evidence.

4.1 Localities

It is evident that KAPFF's first collections were made near Heslach in a large quarry (or a chain of smaller quarries) stretching for the distance of 70-80 metres from west to east (KApPF 1859). These finds include probably the majority of material described by MEYER (1861). One of the syntypes of Nicrosaurus kapfji, SMNS 4060, was recovered later in December 1859 in the course of enlarging the same quarry and is therefore not included in KApPF's publication. The provenance of most of the finds after 1861 (MEYER 1863, 1865; SMNS and BMNH, undescribed specimens listed in subsections 2.2.2 and 2.3.2) is still unknown. A single remark in FRAAS (1877) indicates, that new quarries were opened later in the 1870s "to the left and right side of the main road of Kaltental" (0. FRAAs 1877: 2, my translation). This corresponds to the "Billert" site (see below) and a small infilled area opposite to the road leading to Heslach (R3509875 H54OO425). The new sites also produced semionotid and phytosaur remains as the Heslach site, and FRAAs' remark might suggest that in later times the centre of vertebrate finds shifted from Heslach to the Kaltental area.

380 Appendix C Localities

Two different localities have been discussed as the main source of KApPF's finds. The first is a locally well known cliff face, the "Heslacher Wand" at the mouth of the Nesenbach Valley to the east of Heslach (BRENNER 1973; BENTON & WILD 1993: locality 16). The large outcrop today runs for more than 200 metres from northwest to southeast. Alternatively, WILD (1991) and BENTON & WILD (1993: locality 17) suggested the large quarry "Kohlhau" on the Sonnenberg to the northeast of Kaltental, already indicated in the first issue of the local geological map from 1865, which is now infilled and used as a sports ground.

A secure fixed point to start an identification is the only vertebrate-bearing site that can be pinpointed with reasonable accuracy, a former quarry in the field "Billert" on the ridge separating the rivers Nesenbach and EIsenbach (0. FRAAS 1877; STROBEL & WURM 1977; WILD 1989). The quarry location might be indicated by a mudstone lens within the Stubensandstein in recent issues of the geological map, an area long since claimed by the outskirts of Kaltental. Here, KAPPF recovered the famous slab with 22 AetosaurusJerratus during 1874 and 1875 (KAPPF 1875). The quarry also produced phytosaur remains (0. FRAAs 1877), but the alleged occurrence of Nicrosaurus kapffi (STROBEL & WURM 1977) can not be substantiated. According to KApPF (1875: 303), he uncovered the type of Sellosaurus gracilis in 1871 (or in 1872, SMNS archives) at the very spot where the type of Teratosaurus suevicus was found in 1860, "about 112 hour [c. 2000 metres] away" from the Aetosaurus-site. The distance is in accord with the Heslacher Wand, but not with the quarry Kohlhau, which is much closer. Strangely, there is no mention of phytosaurs at the Teratosaurus site in KAPPF's (1875) account, but these are not the subject of his paper. The Heslacher Wand is also the most likely site of the former "Heslacher Sandgrube below the Jagerhaus" (E. FRAAs 1910), although I was unable to find evidence for a nearby forrester's house, and the "large Stubensandstein quarry above Heslach" (0. FRAAs 1877), according to both authors the main fossil site. The size of the Heslacher Wand also matches well the dimensions given by KApPF (1859).

4.2 Stratigraphy

The stratigraphic position of each site is debated. An inspection of the Nesenbach Valley showed, that no section spanning the whole Stubensandstein can be measured. With the exception of the Heslacher Wand, only patchy small outcrops, which are difficult to correlate, are exposed at the moment. According to composite logs in BRENNER (1978a, b), only lower Stubensandstein (26-30 metres) and middle Stubensandstein (c. 30 metres) crop out on the slopes of the Nesenbach Valley.

381 Appendix C Localities

The quarry Billert has been referred either to the lower Stubensandstein (WILD 1989; BENTON & WILD 1993) or to the middle Stubensandstein (BRENNER 1973; BRAUHAUSER & FRANK 1932; STROBEL & WURM 1977). Strangely, STROBEL & WURM's placement of the quarry, which is exactly at the interface of Stubensandstein and underlying Kiselsandsteinschichten, contradicts their stratigraphic interpretation. A final decision depends largely on locating accurately the quarry face in 1874: if a placement exactly at the slope of the EIsenbach Valley is correct (STROBEL & WURM's locality data), lower Stubensandstein is reasonable. Indeed, former quarrying activities can still be traced at several spots on the slope just above the EIsenbach (pers. obs., 1996). If the location was more south towards the ridge top (WILD'S data), the quarry could have well included middle Stubensandstein strata. This view is supported by the outline of infilled areas to the west on the geological map. In summary, while lower Stubensandstein seems to be more likely, the other alternative cannot be completely ruled out at the moment. The Heslacher Wand section was mostly placed in the lower Stubensandstein (BENTON & WILD 1993; MULLER 1955; STROBEL & WURM 1977). However, in all lithostratigraphic studies, the present outcrop was identified as middle Stubensandstein, sandstone units scI and sc2 (STOLL 1929, corrected by BRENNER 1978a). A figure of the quarry face in the late 1870s is given in O. FRAAS (1877: fig. p. 2). BRENNER (1973a: 136) identifies the basal sandstone bed (top of sandstone sel) of his log as source of KAPFF's finds by correlating this bed with a 200 cm thick sandstone described by O. FRAAS (1877) as being at the foot of the quarry face. However, FRAAS' log was actually measured in the quarry Billert, and does not correspond in detail with the Heslacher Wand section as exposed today. Moreover, the former quarry floor at Heslacher Wand is covered by extensive spoil heaps, and it is not known if still deeper beds had been quarried in the 1850s and 1860s.

In summary, the quarry Billert is the only vertebrate-bearing site in the Heslach Kaltental region which can be identified with certainty. Apart from the Aetosaurus ferratus slab, no Stubensandstein specimen kept in a collection can be assigned unquestionably to a particular site. However, it is concluded that the types of Teratosaurus suevicus, Sellosaurus gracilis, and at least the phytosaur finds of KAPFF prior to 1861 are derived from the Heslacher Wand or a site very close nearby. The stratigraphic provenance of all specimens from Heslach and Kaltental (i.e. the majority of phytosaur remains considered in this study) can be restricted to lower or middle Stubensandstein. A more detailed referal is presently not applicable, although BRENNER's suggestion of the Heslach specimens being derived from the middle Stubensandstein seems likely. It is possible that a thorough small-scale remapping and

382 Appendix C Localities

correlation of the sandstone beds throughout the Nesenbach Valley could result in a detailed local lithostratigraphic framework, which could be used to test BRENNER's disputed identification. However, because of the lack of detailed excavation records, the placement of the existing phytosaur specimens in the framework will always meet some doubts.

List of identified phytosaur remains of KAPFF's 1858-1860 campaigns, from Heslach, most likely from Heslacher Wand: • SMNS 4060, Nicrosaurus kapJfi, syntype, snout and partial lower jaw • SMNS 4059, Nicrosaurus species B, partial skull (KAPFF 1859: specimens III, and 1,1) • SMNS uncat. no. 6, postsymphyseal part of mandible (KAPFF 1859: specimen II,I; MEYER 1861: 302) • SMNS uncat. no. 64 [ex 4060], left scapula (KAPFF 1859: specimen 1,3; MEYER 1861: pI. 35 figs. 2-3) • SMNS uncat. no. 80, eight associated osteoderms (KAPFF 1859: specimen 1,10)

The reasonably accurate provenance of these specimens in the quarry is given in KAPFF (1859). Moreover, KApPF lists numerous additional mandibular fragments and postcrania of phytosaurs from the same site. Although detailed measurements are given as well, no additional specimen in the SMNS collection can be identified unequivocally. MEYER (1861) subsequently described and illustrated the finds made until 1859, and more material that has been collected in the two following years. This might suggest that all material in MEYER (1861) is derived from Heslacher Wand.

5. Stuttgart-Gaisburg

TK 1:25 000: sheet 7221 Stuttgart-Stidost.

Quarry "Raichberg", R35 16250 H5404150.

At least two quarries were in operation at the beginning of the century to the southwest of Gaisburg (LANG 1909). The log given for the large quarry on the Raichberg spanned almost the entire middle Stubensandstein (STOLL 1929; LANG 1909: 117, "upper sand quarry"). The specimen label of Nicrosaurus kapJfi SMNS 13078, found in 1912, reads "sand pit AbelsbergstraBe", which corresponds perhaps to LANG's "lower sand pit" (1909: 117, log units 4a-17). BRENNER (1978a) places the recorded logs of

383 Appendix C Localities

both quarries in the middle Stubensandstein. Therefore it is likely that SMNS 13078 (and tibia SMNS 13077) are derived from middle Stubensandstein, substantiating the occurrence of Nicrosaurus kapffi at Magstadt in the same beds. The coordinates presented in BENTON & WILD (1993) are most probably mistaken.

6. Botnang

TK 1:25000: sheet 7220 Stuttgart-Siidwest.

Quarry "EISENBARTH", probably R35 10100 H5404900.

In 1939, the SMNS recovered the partial skull (SMNS 18645), parts ofthe lower jaws (SMNS 18646), and a scute (SMNS 18644) of a specimen of Nicrosaurus kapffi from the quarry EISENBARTH near Stuttgart-Botnang (BERCKHEMER 1939; SMNS archives). The present location of the specimens is unknown; they were probably destroyed in the war. A quarry is indicated at R35 10100 H5404900. According to the position, the stratigraphic level is probably lower Stubensandstein.

7. Magstadt near Sindelfingen

TK 1:25000: sheet 7220 Stuttgart-Siidwest (also on sheet 7219 Weil der Stadt, BENTON & WILD 1993).

Sandwerk Magstadt, R350120 H540000.

BENTON & WILD (1993) give the coordinates of Sandwerk Magstadt near Sindelfingen as R350120 H540000. The quarry, which is now abandoned but still accessible, operated in the middle Stubensandstein. A log of the quarry face has been measured, but it proved impossible to refer sandstone beds exposed to a particular subunit of the middle Stubensandstein (M. URLICHS, pers. comm., 1998). The phytosaurs from Magstadt were collected in 1974 and 1976 by M. URLICHS (SMNS) at the base of the same sandstone bed, along with numerous temnospondyl remains. Most important is SMNS 56989, a longitudinally split snout of Nicrosaurus kapffi (still unprepared). The specimen represents the most well-founded evidence for the occurrence of Nicrosaurus kapffi in the middle Stubensandstein. The phytosaur material from the quarry includes:

384 Appendix C Localities

• SMNS 56989, Nicrosaurus kapffi, snout, largely unprepared • SMNS 50818, sacral vertebra (natural mould) • SMNS 56990, clavicle • SMNS 51440, vertebral centrum and rib • SMNS 56991, paramedian osteoderm

8. Riibgarten near Tiibingen

TK 1:25 000: sheet 7421 Metzingen

Quarry close to castle Wildenau, R35 12975 H5379300.

The ruin of castle Wildenau is indicated on the map as a faint dotted circle just to the east of the deep castle moat. The former quarry "Wildenau" is located about 30 metres to the east of Wilden au. about 10 metres down the hill slope (310 to 320 D.D.). This location is fully in accord with the itinerary given in MEYER & PLIENINGER (1844) and QUENS1EDT (1850). The quarry cliffs are now toppled over and heavily overgrown, and measuring a log would require a lot of clearing. The present height of the quarry face varies between 5 and 10 metres. Fossil wood is still abundant in boulders scattered on the quarry floor. The quarry "Wildenau" is commonly referred to the upper Stubensandstein (BRENNER 1973; BENTON & WILD 1993), probably by analogy to an active quarry situated 500 metres to the north (R35 12850 H5379700). This quarry spans a section in the upper Stubensandstein (T. AIGNER, GPIT, pers. comm., 1994), however, the quarry face is situated at a height from 355 to 370 a.D. The top of the middle Stubensandstein a few kilometres away from Rtibgarten can be placed at 340 a.D. (BRENNER 1978a: fig. 2; 1978b: 227-228). The first Stubensandstein (lower Stubensandstein) pinches out towards the southwest of Wtirttemberg and is already drastically reduced in thickness in the Schonbuch area (BRENNER & VILLINGER 1981; compo KRIMMEL 1980: fig. 32), and for this reason not worth commercial quarrying. The topographical position indicates, that the Wilden au site must be placed within the middle Stubensandstein. The quarry "Wildenau" is the site of the first find of a phytosaur in 1826. Phytosaurus cylindricodon (for the history of the collection see PLIENINGER in MEYER & PLIENINGER 1844). Since the quarry was abandoned already sometime in the 1830s (QUENSTEDT 1850, 1885-87), postcrania in the GPIT collection (HUENE 1922) labelled "Rtibgarten" must come from a different locality.

385 Appendix C Localities

9. Kayh near Tiibingen

TK 1:25000: sheet 7419 Herrenberg.

Quarries "West 1" and "West 2" of JUNGHANS et al. (1997), R 349550 H53S260.

The quarries near Kayh to the northwest of TObingen represent the only phytosaur bearing site, of which the stratigraphy and lithology is excellently documented and a facies analysis was undertaken (STOLL 1929; JUNGHANS et al. 1997). Moreover, unpublished data of the location and the history of the vertebrate finds is preserved (GPIT archives). The site was recently cleared from rubble and classified as a protected natural monument. The locality consists of a string of abandoned quarries at the southern rim of the Schonbuch Forest. Vertebrate remains were found spatially and stratigraphically restricted to a section named "West 1" and "West 2" in JUNGHANS et al. (1997). According to STOLL (1929) and corroborated by lithologic criteria of the adherent matrix (JUNGHANS, pers. comm., 1993), all fossils are derived from the basal beds of sandstone sc2 in the middle Stubensandstein. At Kayh, these beds are interpreted as a repetitive series of crevasse splay deposits (JUNGHANS et al. 1997). Between 1923 and 1927, numerous fishes, mostly fragmented phytosaur postcrania, phytosaur teeth, aetosaur scutes, coprolites, and indeterminable bone fragments (now GPIT Inv.-No. 2070) were collected by quarry men. Finally, STOLL recovered the skull of a Nicrosaurus species B GPIT 2070,001 during fieldwork. Only recently, corresponding beds in one of the other quarries yielded plant remains and numerous semionotid fishes (Semionotus bergeri, GPIT uncat.).

10. Trossingen and Aixheim

TK 1:25 000: sheet 7817 Rottweil. TK 1:25000: sheet 7818 Wehingen. TK 1:25000: sheet 7917 Villingen-Schwenningen. TK 1:25 000: sheet 7918 Spaichingen.

Quarry "Untere Miihle I": R3474925 H5427875. Quarry "Untere MOhie II": R3475280 H5428400. Quarries around Neuhaus: R3477080 H5430400; R3477110 H5430350. Quarries "Wannengrund": R347125 H542905.

386 Appendix C Localities

Towards the southwest, the sandstone beds of the Stubensandstein gradually decrease in thickness and are substituted by mudstones deposited in a playa plain. In the region of Trossingen and Aixheim, the lower and upper Stubensandstein are mostly represented by uniform silt- and claystones, interbedded with dolomitic limestones and occasional very thin sandstone beds. The site of the famous Plateosaurus-bonebeds (SANDER 1992), usually placed in the top unit of the Middle Keuper, the Knollenmergel, is indeed contemporaneous at least in parts with the third Stubensandstein (upper Stubensandstein) of central and northern Wiirttemberg. According to this geological context, all the numerous quarries that once worked in the sandstone beds in the region can be safely referred to the middle Stubensandstein (BRENNER & VILLINGER 1981). STOLL (1929) identified sandstone sc3 as the vertebrate-bearing bed, and this is confirmed by the logs and descriptions for the Neuhaus quarries (LANG 1909) and Untere Miihle (HUENE 1915b; pers. obs.). BRENNER & VILLINGER (1981) presented a number of logs recently measured in the north and northwest of Trossingen. They showed, that although sandstone sc3 is still present with considerable thickness, here sandstone sc2 is the dominant bed. For this reason, it cannot be excluded that in the area west of Trossingen, which includes the quarries "Wannengrund", beds from the sandstone sc2 were once quarried as well.

Isolated postcranial remains of phytosaurs were said to be quite common in the Stubensandstein around Aixheim and Trossingen, whereas skulls and articulated skeletons must have been exceptional rarities (BERCKHEMER 1931). Unfortunately, it was a common practice to label fossils or to refer to specimens in print as being derived generally from "Aixheim" or "Aldingen", and for this reason many finds cannot be referred to a particular site any more.

10.1 Quarries around Neuhaus

The first recorded quarries near Aixheim were established in the Prim Valley (ESER 1907) around Neuhaus (formerly also termed Neuwirtshaus) in at least the 1850s, and a number of small quarries continued to work at the same spot until the 1930s (BERCKHEMER 1931). The coordinates of two of the more recent quarry locations are taken from GAFFNEY (1990). Both these, and all former quarry locations, are now infilled and inaccessible. Only two phytosaur specimens are referred to the Neuhaus quarries in litteris, but it is reasonable to assume that much more material was recovered and acquired by museums over the years (comp. GAFFNEY 1990). ESER's accounts on the topology of the Stubensandstein outcrops in the Aixheim region (ESER 1862; 1907: 645,659)

387 Appendix C Localities

strongly suggests that the types series of Mystriosuchus pianirostris was also collected at Neuhaus.

Identified specimens from the quarries "Neuhaus": • type series of Mystriosuchus pianirostris: MCZ 1018, partial skull; MCZ 1019A, B, 3 fragments of the rostrum; ?MCZ 1022, isolated frontal and parietal • SMNS 13007, Mystriosuchus pianirostris, skull and cervical column (quarry "JEITER"; SMNS archives) • SMNS 13240, Mystriosuchus planirostris, skull (quarry "JEITER"; SMNS archives)

10.2 Quarries "Wannengrund"

Around the turn of the century, quarrying activities intensified because of the need for building stones for the new church of Aixheim. The area northeast of "Wannengrund" was exploited and soon became the place of the most productive quarries in terms of phytosaur finds (SCHMIDT 1914, HUENE 1923). Apparently fossils were not only collected by locals and sent to the museums as usual, but a major excavation by the SMNS under supervision of E. FRAAs took place in 1903, funded by a generous donation of 1000 Reichsmark by the industrialist E. von SIEGLIN (BERCKHEMER 1931). Quarrying must have ceased here only a few years later (HUENE 1923).

Identified specimens from the quarries "Wannengrund": • SMNS 11126(1), Mystriosuchus planirostris, partial skull • SMNS 11126(2), Mystriosuchus planirostris, complete skull, [ABEL 1923: fig. 2; now lost] • SMNS 11126(3), Mystriosuchus pianirostris, skull [SMNS archives, now not to identify or lost] • SMNS 11127, osteoderms • SMNS 11128/1, left femur • SMNS 11128/2, left femur • SMNS 11128/3, left femur • SMNS 1112814, right scapula • SMNS 11128/5, right clavicle • SMNS 11129. pubis • SMNS 11130/1, part of a mandible • SMNS 11130/2, left coracoid

388 Appendix C Localities

10.3 Quarries near Untere Miihle

Of the two quarries near Untere Mtihle, the one situated on the southern slope of the Trosselbach Valley (R3474925 H5427875, here designated Untere Miihle I), close to the famous Plateosaurus-bonebed at Obere Miihle (SANDER 1992, and references therein), is identified as the type locality of Mystriosuchus species B. The quarry is still identifiable in the field, but now largely infilled and overgrown. Untere Mtihle II (R3475280 H5428400) is still accessible, as a small quarry in the same bed, which has allegedly also yielded vertebrates (LANG 1909). A notable exception concerning the referral of specimens to a particular site is the collection from Untere Miihle (= Untere Mtihle I) in the GPIT. The quarry was constantly monitored, and finds collected and shipped to von HUENE, probably for several years. The body of Untere Miihle specimens now represent the largest association of vertebrate remains from the Stubensandstein, that is derived from a single sandstone bed (sandstone unit sc3) at one location. Untere Mtihle I also produced three articulated partial skeletons of the prosauropod dinosaur Sellosaurus gracilis (HUENE 1915b).

Untere Mtihle collection: • GPIT 249/001 Mystriosuchus species B • GPIT 2074.000, Mystriosuchus planirostris, basicranium • GPIT 2149.002, Mystriosuchus planirostris, right posteroventral cheek • GPIT 2150.000, Mystriosuchus planirostris, quadrate and quadratojugal • GPIT 2149.000, Mystriosuchus planirostris, postorbito-squamosal bar • GPIT 2149.001, Mystriosuchus planirostris, right squamosal • GPIT 2149.003, jugal fragment • GPIT 2100.004, (?) caudal rib • GPIT 2111.000, spinal process • GPIT 2116.001, spinal process with prezygapophyses • GPIT 2116.002-003, two postzygapophyses • GPIT 2217.000, spinal process with postzygapophyses • GPIT 2215.000, centrum of caudal vertebra • GPIT 2091.000, right scapulocoracoid • GPIT 2218.000,left scapula • GPIT 2099.001, right scapula • GPIT 2099.002, left scapula • GPIT 2072.000, left scapula • GPIT 2104.00 I, right scapula • GPIT 2104.002, left scapula

389 Appendix C Localities

• GPIT 2104.003, (?) left scapula • GPIT 2104,004, (?) scapula • GPIT 2137.000, left scapula • GPIT 2138.000, right scapula • GPIT 2107.000, coracoid • GPIT 2090.000, (?) right clavicle • GPIT 2100.002, left clavicle • GPIT 322/035, left humerus (HUENE 1922: fig. 35) • GPIT 2087.000, (?) left humerus • GPIT 2038.000, (?) left humerus • GPIT 2079.000, left humerus • GPIT 2094.000, right humerus • GPIT 2100.001, left humerus • GPIT 2122.000, left radius • GPIT 2224.000, radius • GPIT 2119.000, left ulna • GPIT 322/049, right femur (HUENE 1922: fig. 49) • GPIT 2033.001-002, right femur • GPIT 2087.000, (?) left femur • GPIT 2131.000, right femur • GPIT 2100.003, (?) right tibia • GPIT 2123.000, left tibia • GPIT 3221055-056, metatarsals (HUENE 1922: figs. 55-56) • GPIT 2132.000, (?) metatarsal

390 Figure 1.1: Phytosaur localities and palaeogeography of the Keuper succession.

A Map of central Europe showing the localities of important phytosaur finds. B Detail of Figure 1.lA showing Southwest Germany with the phytosaur localities treated in the text and Appendix C. Modified from BEmON (1993). C Palaeogeographical map of central Europe in the Norian superimposed on the modern map Figure 1.IA showing the former highlands and the distribution of the central basin facies (Steinmergel-Keuper) and marginal basin facies (Sandstein-Keuper) in the Germanic Keuper Basin. Based on ZIEGLER (1982).

391 • Schilfsandstein middle Stubensandstein • Stubensandstein, unspecified • highlands • marginal basin deposits Lowenstein f:·::·:·:] Pfaffenhofen/ D central basin deposits Botnang • Murrhardt

Sindelfingen.----"'"• .... Salzgitter* :+'t1 e V Bernburg *Halberstadt N \~ Coburg* * Tubingen Rubgarten Ebrach* ! ! •25 km

B Figure 1.2: Stratigraphy and lithology of the Keuper and the Stubensandstein deposits.

A Idealised lithostratigraphical section of the Keuper succession and chronostratigraphic framework. The stratigraphical positions of important phytosaur genera are plotted on the log. Log based on AIGNER & BACHMANN (1992). B Lithology and lithostratigraphy of the Stubensandstein deposits. Modified from HORNUNG (1998).

392 litho- _.... JJthoIogy Chronostrat. L thostrat graphy detailed IItnItIgrllphy IItnItIgrllphy -- I~ System I Stage KncIonnorvaI ; (1an5) ! I ....II II - t .. t t II Jurassic IHettangian Lias Lias a ~ DI4 I 4. Haw1gendIetten r:: ca .-.- _ I 4. Slubensandstein to) 0 ca ...r:: ~ - ~1%*)*;~;:~:.:"·.·· .• '"' ~ t ." E c;; - _ CD 0 .... "'" Knollenmergel ~ ... '" I cI3 I 3. Hangendlelten c.~ I c. E r:: ~~~~>-___I _ c :l~. Cd C!) Coburgosucluu ..; I ?- 003 3. Slubensandstein til ~ Q) ..- mI2 2. Hangendletlen '""0 CI) Stubensandstein .:.::.•.. .:.:.-:~ If:' . .2.~a ~tu.~~S:."~~~I.~ .• : •• :... .. MyslTiosuchus ...... Z I Q" -C/) mo2.3 Ic:3 E Iftld( til 1"'1 $!~~~ndslelD .111 I ,F 1. Stubensandstein .....: .•:..... NicroSilunu "C .... "0 mZD Obere ZWtschenleUen C EN' ·····1···· ~'" :::I ---E "0 CO ~ Kieselsandstein cap r:: C/) ~g ~21~.iba""""""" ~ Francosuchll$ Ebrachosuchus "0 ca C "0 0 8 Q) E NI ~·ZWtschenIetten- 1 Rote Wand r:: .c i mo2.,i scI .... I :::l ..,K '""Cd ~ SclWf"""."Gipskeuper -~...... •.' ... YI:'1Cljo"r_*-Ac.* \:::::::~~:):~::::.: :.~ ::\}~~/.:~::.:-; 1k!-2 I 1. Hangendlellan U en I-< I .----- [3J...... ' ....:. sandstone "'-2 .....1 ...... Ik!-,I_- CI) c: :t Lettenkeuper ...:".i .. i~~·'!teuPer-HaupisaOdsiein" ...... ::.:: . c=J pelite .. ,-, ..9 • ~ ~ dolomitic -,,",","""A"< ~Grenzbone!'e~~ limestone uB I Rote BasIsIetten Muschel- U. Muschelkalk A L I ~ kalk ~ limestone B\fa..*: I Figure 2.1: Nicrosaurus kapffi (MEYER, 1860), skull material used in this study. The specimens are grouped into high-domed, gracile (top) and low-domed, robust skull morphs (centre). SMNS 54706 (bottom) is not referable to a particular morpho

Hatched areas: restored or damaged surfaces; irregularly dashed area: exposed articular surface; scale bars indicate 100 mm.

393 High-domed, gracile specimens

SMNS4378 ~ ~ (inverted)

BMNH 42743

Low--domed, robust specimens

_.------<'-..r- SMNS4379

SMNS5725 (inverted) ~~ ~~~~ SMNS5726

SMNS 13077 (Inverted)

not referred to a morphotype

SMNS54706 Figure 2.2: Cranial measurements of Nicrosaurus kapfji (Tab. 2.2.1) and Nicrosaurus species B (Tab. 2.3.1).

Shaded areas: restored surfaces.

394 1----9---J r·····'····1-----9----1 r 10

l~.~ ...... If.1 . . 1--21~ i23 i 13--.1 r------8:--~------~ 1----22----J Figure 2.3: Nicrosaurus kapfji (MEYER, 1860), SMNS 4060, lectotype.

Snout fragment SMNS 4060; two additional posterior jaw fragments that possibly pertain to the same animal are not figured. A, B Right lateral view. Diagonal hatching indicates restored areas. e Dorsal view. D Detail of the prenarial crest in a position indicated in Figure 2.2B, showing an arrangement of three rounded holes still infilled with matrix. E Cross-section of the snout at level of the 8th premaxillary alveolus, as indicated in Figure 2.2B, to illustrate the cavernous internal structure of the prenarial crest. ? indicate areas obscured from view.

Mandibular fragment SMNS 4060a, found in occlusion with the snout SMNS 4060. F Dorsal view.

395 -•

B .----100mm

F Figure 2.4: Nicrosaurus kapfji (MEYER, 1860), SMNS 54708, paralectotype, anterior left fragment of a snout.

A Lateral view of the external side in its present state of preservation. Note in particular the steeply downturned tip of the snout. B Lateral view of the external side, redrawn after MEYER (l861: pI. 31 fig. 9).

396 A

B Figure 2.5: Nicrosaurus kapffi (MEYER, 1860), SMNS 4378.

A Dorsal view (copy of MEYER 1863: pI. 38). B Ventral view (copy of MEYER 1863: pI. 39). C Right lateral view (copy of MEYER 1863: pI. 40).

397 , . " '

100 100 mm mm

m100m

~-~ ...... ,"" ~ . W..; ... 1';; ,-;:.I l" ;- - - . 'It :1 ' . .. -

~-., ( ~ ,.

~. ; ~ , ~ :L, "'- -. ~. -:;r- - I~ '~n ) '~.,.

/', ." \ - i '. !

\ , , - i;,: '- /' / / /

" "

8 c A Figure 2.6: Nicrosaurus kapffi (MEYER, 1860), SMNS 4378, right posttemporal fenestra in posterior view.

Note the lateral extent of the squamosal process of the parietal on the parieto-squamosal bar and the medioventral extension of the squamosal onto the posterior face of the paroccipital process.

Abbrev.: bs, basisphenoid; eoc, exoccipital; opo, opisthotic; par, parietal; soc, supraoccipital; sq, squamosal. Hatched areas: restored surface.

398 j

j squamosal process of par j parietal process of sq j

..... a::J eoc 10mm paroccipital j process 01 opo j

j Figure 2.7: Nicrosaurus kapffi (MEYER, 1860), SMNS 4378, posterior part of the braincase and the posttemporal fenestra seen through the right infratemporal fenestra.

The paroccipital lamella, a medioventral extension of the paroccipital process of the squamosal is exposed on the anterior face of the paroccipital process of the opisthotic, and the pterygoid process of the squamosal is visible on top of the pterygo-quadrate plate.

Abbrev.: ep, epiotic; opo, opisthotic; par, parietal; pro, prootic; pt, pterygoid; q, quadrate; sq, squamosal; Roman numerals indicate cranial nerves. Stippled area: matrix; hatched area: restored surface.

399 j j j j j j j j j j j j j parleto-s j barqua mosal j j j POsltemporal ~ tenestra fOmm j j °P/SlhoUc j lamella of sq ---1hC> j j Pterygoid j prOCess of sq j j j j j j j j PI j j j j j j j j j j j j j j j j j j j Figure 2.8: Nicrosaurus kapffi (MEYER, 1860), SMNS 4379, dorsal view.

Abbrev.: f, frontal; j, jugal; 1, lacrimal; m, maxilla; n, nasal; par, parietal; pm, premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; q, quadrate; qj, quadratojugal; sm, septomaxilla; sq, squamosal. Hatched area: restored surface. Photo: R. HARLING (SMNS).

400 I 700 mm Figure 2.9: Nicrosaurus kapffi (MEYER, 1860), SMNS 4379, palate and palatal vault in ventral view.

The outline of the palatine on the palatal vault is well preserved. The areas of pitting and transverse ridges on the palate are indicated by circles.

Abbrev.: bs, basisphenoid; ect, ectopterygoid; j, jugal; m, maxilla; pal, palatine; pm, premaxilla; ps, parasphenoid; pt, ptyerygoid; qj, quadratojugal; v, vomer. Dotted areas: matrix; hatched area: restored surface. Photo: R. HARLING (SMNS).

401 palatal plale

transverse ridges pits

suborbital fenestra

palatal axis ---1---=:::....4------'

pterygoid flange ~ subtemporal t fenestra ~mm ~ Quadrate process of pt Figure 2.10: Nicrosaurus kapffi (MEYER, 1860), SMNS 4379, right lateral view.

Abbrev.: f, frontal; j, jugal; 1, lacrimal; n, nasal; q, quadrate; qj, quadratojugal; m, maxilla, pm, premaxilla; po, postorbital; pof, postfrontal; pt, pterygoid; sq, squamosal. Hatched area: restored surface. Photo: R. HARLING (SMNS).

402 E 0. \

't: a. Figure 2.11: Nicrosaurus kapffi (MEYER, 1860), SMNS 4379, skull in posterolateral view from ventral to illustrate the main anatomical structures of the temporal and occipital regions treated in the text..

Abbrev.: opo, opisthotic; q, quadrate; pt, pterygoid; sq, squamosal. Dotted areas: matrix; hatched area: restored surface. Photo: R. HARLING (SMNS).

403 palatal .. pterygoid flange --l00mm - - ridge

bean-shaped groove pterygo-quadrate plate \ pteroccipital fenestra

pterygo-quadrate bar (overhang) groove on quadrate wing 01 pt

groove on pterygoid wing 01 q

lateral ridge

ventral sulcus

triangular recess

ventral _--I knob jugular groove

anterior shelf of dorsal sulcus paroccipital process medial extension of sq posterior posterior ridge process of sq of paroccipital process parieto- squamosal bar ventral ridge olopo Figure 2.12: Nicrosaurus kapffi (MEYER. 1860). SMNS 5725. dorsal view.

Abbrev.: f, frontal; j,jugal; 1, lacrimal; m, maxilla; n, nasal; pm, premaxilla; po, postorbital; prf, prefrontal; sm, septomaxilla. Photo: R. HARLING (SMNS).

404 -- 100 mm

po Figure 2.13: Nicrosaurus kapffi (MEYER, 1860), SMNS 5725, ventral view.

Abbrev.: ect, ectopterygoid; j, jugal; m, maxilla; pal, palatine; pm, premaxilla; v, vomer. Dotted areas: matrix. Photo: R. HARLING (SMNS).

405 previous alveolar rim

100 mm

interpterygoid hssure Figure 2.14: Nicrosaurus kapffi (MEYER, 1860), SMNS 5725, right lateral view.

Abbrev.: f, frontal; j, jugal; 1, lacrimal; n, nasal; rn, maxilla, pm, premaxilla; prf, prefrontal. Irregularly dashed areas: damaged bone surface. Photo: R. HARLING (SMNS).

406 - E c. Figure 2.15: Nicrosaurus kapffi (MEYER, 1860), SMNS 5726, dorsal view.

407 "",.J. .. _I'.. !:O ...•

100 mm Figure 2.16: Nicrosaurus kapffi (MEYER, 1860), SMNS 5726, ventral view.

Abbrev.: boc, basioccipital; bs, basisphenoid; ect, ectopterygoid; j, jugal; m, maxilla; opo, opisthotic; pal, palatine; pm, premaxilla; pro, prootic; pt, ptyerygoid; q, quadrate; qj, quadratojugal; v, vomer. Hatched areas: restored surface; irregularly dashed areas: damaged bone surface. Photo: R. HARLING (SMNS).

408 dorsal wing of pal

100 mm Figure 2.17: Nicrosaurus kapffi (MEYER, 1860), SMNS 5726, pterygo-quadrate plate and paroccipital process of the opisthotic in posteromedial view from ventral.

Abbrev.: opo, opisthotic; pro, prootic; q, quadrate; qj, quadratojugal; pt, pterygoid; sq, squamosal. Main fractures on the pterygo-quadrate plate and the quadrate are indicated by dashed lines.

409 pterygo-quadratal bar ventral process of cranlo-mandlbular Quadrate wing 01 pi Joint 01 q dorsal process 01 quadrate wing 01 pt recess ot q pterocclpital lenestra

jugular .,,\r.~; groove .. foramen quadrati

posterior ridge 01 paroccipital process

_____ e paroccipital process 01 opo 10mm tympanic vertical posterior fossa face of opo

paroccipital process 01 sq Figure 2.18: Nicrosaurus kapffi (MEYER, 1860), SMNS 5727, dorsal view.

Abbrev.: q, quadrate; qj, quadratojugal; sq, squamosal. Hatched areas: restored parts or surfaces. Photo: R. HARLING (SMNS).

410 100 mm Q

panctat foramen Figure 2.19: Nicrosaurus kapfji (MEYER, 1860), SMNS 5727, ventral view.

Dotted areas: matrix; hatched areas: restored parts or surfaces. Photo: R. HARLING (SMNS).

411 100 mm Figure 2.20: Nicrosaurus kapfji (MEYER, 1860), SMNS 5727, right lateral view.

Hatched areas: restored 1?arts or surfaces. Photo: R. HARLING (SMNS).

412 c Figure: 2.21 Nicrosaurus kapffi (MEYER, 1860), SMNS 54706, isolated right cheek and temporal region

A Anterior part of right jugal and postorbital in lateral view. Note the sutural facets of the jugal to maxilla and lacrimal, the small contribution of the jugal to the orbital rim, and the presence and dorsal extent of a lamina into the infratemporal fenestra, resulting in an external ventral rece s

B Fragment in medial view. The medial spine and the anteroventral recess of the jugal, the anterior lamina of the jugal and the facet for the jugal process of the ectopterygoid are indicated. Note al 0 the presence of an epiphyseal canal within the interparietal suture and the ventral extent of the squamo al process of the parietal onto the occipital shelf. The preservation is similar to a specimen of Rutiodon carolinensis (GREGORY 1962b: fig. 3), especially regarding the "continually sloping" dorsal rim of parieto- quamosal bar as a probable preservational artefact.

Abbrev.: eoc, exoccipital; j, jugal; I, lacrimal; m, maxilla; opo, opi thotic; par, parietal; po, postorbital; pof, postfrontal; pro, prootic; q, quadrate; qj, quadratojugal; sq, squamosa\. Dotted areas: matrix.; hatched areas: restored parts or surfaces; irregularly dashed areas: damaged bone surface or rugose facets.

413 knob descending orbital nm lamina squamosal process of pa r po po dorsal j contacts sulcus orbit interparietal suture posterior process of sq

epiphyseal l;'''~-'-'~;:,(. covered canal ,.. ? medial facet for I spine po L. parieta l process of sq

ventral sulcus

paroccipital process of opo

j-Iac _ suture -"

medial ~ sp in e facet for m anterior .. ':';'~... :.:\ foramen recess of j ..... ? •• ..i quadrati j-ect suture A (dorsal process)

c; 100 mm

B j-eet suture Figure 2.22: Nicrosaurus kapffi (MEYER, ) 860), BMNH 42743, dorsal view.

Abbrev.: f, frontal; j, jugal; I, lacrimal; m, maxilla; n, nasal; par, parietal; pm, premaxilla; po, postorbital; pof, postfronta); prf, prefrontal; q, quadrate; qj, quadratojugal; sm, septomaxilla; sq, squamosal. Dotted areas: matrix; hatched areas: restored parts or surfaces; irregularly dashed area: damaged bone surface. Photo and print: Photographic Unit, The Natural History Museum.

414 Q Figure 2.23: Nicrosaurus kapffi (MEYER, 1860), BMNH 42743, ventral view.

Abbrev.: boc, basioccipital; bs, basisphenoid; eCl, ectopterygoid; j, jugal; m, maxilla; opo, opisthotic; pal, palatine; pm, premaxilla; pt, ptyerygoid; q, quadrate; ? represents unclear configuration. Dotled areas: matrix; irregularly dashed areas: damaged bone surface. PhOlO and print: Photographic Unit, The Natural History Museum.

415 ~ 100 ~ mm ~ ~

_~:..&-_ ridge

.. ,.,,, _l.----Pl /fs..:?'~ Figure 2.24: NicrosaLlrLls kapffi (MEYER, 1860), BMNH 42743, right lateral view.

Abbrev.: ect, ectopterygoid; f, frontal; j, jugal; I, lacrimal; n, nasal; q, quadrate; qj , quadratojugal; m, maxilla, n, nasal; pm, premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; sm, scptomaxill a; sq, squamosal. Dotted area: matrix; hatched areas: restored surface; irregularly dashed areas: damaged bone surfaces. Photo and print: Photographic Unit, The Natural History Museum.

416 E a. Figure 2.25: Nicrosaurlls kapffi (MEYER, 1860), BMNH 42743, right posterior part of the skull in posteromedial view from ventral.

Note the steeply sloping quadrate, the sutures on the pterygo-quadratal plate, the sharp laminae bordering the stapedial and jugular groove the foramina of the basioccipital, the posteroventrally facing tubera, and the shape of the basipterygoid process.

Abbrev.: boc, basioccipital; bs, basisphenoid; eoc, exoccipital; pro, prootic; pt, pterygoid; q, quadrate; soc, supraoccipital. Dotted area: matrix.

417 ecl

eCloPlertgoiO palalal V/i1"9 0\ pI tiog6

::.:",: :

oasiSpMI"OiO process 01 pI ...... :.:.:.:.:.: ...: ..: ..:.: ... :.. :.. ::: •. :..: ... .. :.,' ... ::;'.~:: sulCU S

oasipleryQOiO process 01 os ~ .- . .•... _. ... . ,.., ...... ,;..:...... ::;11 IUoeral su rlace oloS

eral IUO c surlace 01 bO

I"UI(\el"l lorarnil"a

. _~ _,., oCClpilal ·-"'<':1; cono)'l6

eoC o,U30r31e V/ll"g 01 pl. Clorsal ptoCess plerygoiO soC V/,ng 01 0. tOUc;nel"eo ar ea o,uaorale V/Il"g 01 pl. proc ess ~en\tal pro parocCIpital process Figure 2.26A: Nicrosaurus kapffi (MEYER, 1860), mandible.

A: BMNH 42744, mandible in dorsal view.

Teeth are indicated schematically by fIlled circles, empty alveoli by hollow circles.

Abbrev.: art, articular; d, dentary; part, prearticular; sa, surangular; sp, splenial.

418 1: .go~ _____..., ",II) c~ "'g a. Figure 2.26B, C: Nicrosaurus kapffi (MEYER, 1860), mandible.

B: BMNH 42744, right mandibular ramus in lateral view.

c: right mandibular ramus in medial view, composite diagram from SMNS 5730 (symphysis, and anterior section of postsymphyseal ramus) and SMNS uncat. no. 9 (prearticular-articular complex), including information from SMNS 4380 (anterior process of prearticular).

Abbrev.: a, angular; art, articular; d, dentary; part, prearticular; part-art, prearticular articular complex; sa, surangular; sp, splenial.

419 coronoid process

a tOOmm infrafenestral retroarticular external process of d process ridges mandibular fenestra on sa poslglenoidal B process triangular postglenoidal fo-______,mandibular 'I facet symphysIs

~ ~

symphyseal rugosities foramen canal ---l00mm-- lateral process anterior process of part-art c of part ('coronoid") Figure 2.27: Nicrosaurus kapffi (MEYER, 1860), SMNS 5727 and SMNS 13078 in ventral view to indicate the teeth preserved and to illustrate the extent of the dental sets of the upper dentition.

Blank circles represent the size and shape of the alveoli. The position and outline of teeth, or tooth fragments, is indicate in black. Shaded: restored areas. Dashed outlines: reconstructed alveoli.

420 tip-of-snout set I rosette tip-ol-snout set

anterior constriction __* and diastema

premaxilla set premaxilla set

posterior constriction at pm/m-suture

maxilla set maxilla set -

SMNS 13078

SMNS 5727 Figure 2.28: Nicrosaurus kapffi (MEYER, 1860), SMNS 13078, tip-of snout set, right premaxillary teeth (1) and (2).

A Lateral view (labial face of the teeth). B Detail of the flange at mid-length of the posterior carina of pm (2). An undulating, vertical groove distinguishes the denticles from the flange, and the interdenticle grooves extend onto the flange. C Anterior view (mesial face of the teeth), with three cross-sections of the tooth crown of pm (2). Arrows indicate posterior direction. Note the split and partially doubled anterior carina.

421 . '. ,"'. ,~ ...... ~ "

10 ~ mm ~

/ •

mm'0

c Figure 2.29: Nicrosaurus kapffi (MEYER, 1860), SMNS 13078, tip-of-snout set, left premaxillary tooth (3).

A Lateral view (labial face of the tooth). B Medial view, seen slightly from ventrally (exposing the lingual and parts of the mesial face of the tooth), to show the long, fully serrated carinae. C Anterior view (mesial face of the tooth), with a cross-section near the base of the crown. Note the strong bilateral asymmetry of the tooth crown. Arrow indicates posterior direction.

Figure 2.30: Nicrosaurus kapffi (MEYER, 1860), SMNS 13078, tip-of-snout set, left premaxillary tooth (4)

A Lateral view (labial face of the tooth, tooth crown height somewhat foreshortened). Note the tooth axis pointing posteriorly and ventrally. B Ventromedial view (lingual face of the tooth). C Anterior view, seen slightly from laterally, with two oval cross-sections at mid-length and near the apex of the crown. Arrows indicate posterior direction. The tooth is set up high on the lateral face of the anterior constriction, and only the apex of the crown is deflected ventrally. The faint mesial carina is unserrated.

422 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 8 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 10 mm 1 1 1 1 B 1 1 1 1 1 1 1 1 ., 1 c 1 1 Figure 2.31: Nicrosaurus kapffi (MEYER, 1860), SMNS 13078, premaxilla set, right premaxillary teeth (6) and (7). Pm (6) not fully erupted.

A Lateral view (labial face of the teeth), showing the close spacing and the distal carina of pm (7). B Ventromedial view (lingual face of the teeth). C Anterior view, seen slightly from laterally (mesial and parts of the labial face of the teeth), with two cross-sections of pm (7) near the base and the apex of the crown. Note the oval section near the apex. The tooth crowns are strongly arched and display the mesial carinae. Arrows indicate posterior direction.

Figure 2.32: Nicrosaurus kapffi (MEYER, 1860), SMNS 13078, premaxilla set, left premaxillary tooth (14).

A Lateral view (labial face of the tooth). B Medial view (lingual face of the tooth). Narrow mesial and distal flanges are present. C Anterior view (mesial face of the tooth), with two cross-sections at the base and at mid-length of the crown. Note the kink-like alignment of root and crown, the extent of the carina over the entire crown, and the lingual flattening of the apex. Arrows indicate posterior direction.

423 .. eM _W' 10mm A B c

-81 8, .... -., 10mm A B c Figure 2.33: Nicrosaurus kapjfi (MEYER, 1860), SMNS 13078, premaxilla set, left premaxillary tooth (17). Tooth not fully erupted.

A Lateral view (labial face of the tooth). B Medial view (lingual face of the tooth). Note the broadening of the flanges compared to pm (14). C Anterior view (mesial face of the tooth), with two cross-sections at mid-length and near the apex of the crown. The cross-sections are D-shaped. The gutter-like fitting of the premaxilla rim is visible in the succeeding alveolus, including an replacement tooth pm (18). Arrows indicate posterior direction.

Figure 2.34: Nicrosaurus kapjfi (MEYER, 1860), SMNS 5727, premaxilla set, right premaxillary tooth (20).

A Lateral view (labial face of the tooth). Note the posterior inclination of the straight crown. B Medial view (lingual and part of the mesial face of the tooth). Both flanges show maximum expansion in the premaxilla set. The basal part of the distal carina is unserrated. C Posterior view (distal face ofthe tooth), with two cross-sections at the base and mid length of the crown. Note the slender appearance, the comparatively broad flange, the high vault of the labial face, and the more mesial orientation of the anterior carina compared to pm (17). Arrows indicate posterior direction.

424 ~.r -..r A 8 c

~j .1

~ 10 ~mm A 8 c Figure 2.35: Nicrosaurus kapffi (MEYER, 1860), SMNS 5727, maxilla set, left maxillary tooth (1) (top, left-hand side). A Lateral view (labial face of the tooth), indicating the wide-spaced basal striation. B Posterior view (distal face of the tooth), with two cross-sections near mid-length and at the apex of the crown. The singular, small, unserrated distal carina is shown. Arrows indicate posterior direction.

Figure 2.36: Nicrosaurus kapffi (MEYER, 1860), SMNS 13078, maxilla set, right maxillary tooth (3) (top, right-hand side). A Lateral view (labial face of the tooth). B Posterior view (distal face of the tooth), with two cross-sections at the base and the apex of the crown. Note the stout appearance of the crown and the lengthening of the carina. Arrows indicate posterior direction.

Figure 2.37: Nicrosaurus kapffi (MEYER, 1860), SMNS 13078, maxilla set, left maxillary tooth (6) (centre). A Lateral view (labial face of the tooth). The distal carina extends over the whole crown. B Medial view (lingual and part of the distal face of the tooth). A long mesial carina appears for the flrst time, the distal part being still un serrated. C Posterior view (distal face of the tooth), with two cross-sections at mid-length and near the apex of the crown. The flattening of the lingual side and the mesiodistal asymmetry at the base is visible. Arrows indicate posterior direction.

Figure 2.38: Nicrosaurus kapffi (MEYER, 1860), SMNS 5727, maxilla set, left maxillary tooth (17) (bottom). A Lateral view (labial face of the tooth), with two cross-sections at the base and near the apex of the crown. Two large flanges flanking the high central vault on the labial side, the mesial flange being shorter and steeper. Apically, there is a strong ridge followed by a depression. Arrows indicate posterior direction. B Medial view (lingual face of the tooth). Note the constriction at the base of the crown. C Apical view, showing the recurvedness of the apex overhanging the root, and the mesiodistal asymmetry. D Detail of the mesial carina at the apex of the crown, taken from SMNS 5727, left m (19). E Detail of the distal carina at mid-length of the crown, taken from left m (17).

F SMNS 5727, m (14) to m (20) in ventrolateral view to show the imbricate pattern of the tooth arrangement and the continuous cutting edge in the posterior part of the maxilla.

425 A 8

10 i mm~ A B c

Ie -, C 10mm

E Figure 2.39: Nicrosaurus kapffi (MEYER, 1860), reconstruction of the skull in dorsal view.

The skull morphology is based on SMNS 4379, corrected for distortion, with additional information from SMNS 5726 (skull table, jugal and lacrimal).

Abbrev.: f, frontal; j, jugal; l,lacrimal; m, maxilla; n, nasal; par, parietal; pm, premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; q, quadrate; qj, quadratojugal; sm, septomaxilla; soc, supraoccipital; sq, squamosal. Shaded area indicates the outline of the infranasal groove.

426 \ Figure 2.40: Nicrosaurus kapffi (MEYER, 1860), reconstruction of the skull and mandible in right lateral view.

The skull morphology is based on SMNS 4379, the dentition on SMNS 5727, and the mandible on BMNH 42744.

Abbrev.: a, angular; d, dentary; ect, ectopterygoid; f, frontal; j, jugal; 1, lacrimal; m, maxilla; n, nasal; pm, premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sm, septomaxilla; sp, splenial; sq, squamosal. Shaded area indicates the outline of the infranasal groove.

427

Figure 2.41: Nicrosaurus kapffi (MEYER, 1860), reconstruction of the skull in occipital view.

The morphology is based on SMNS 5727, corrected for distortion, with the sutural configuration according to SMNS 5726. Further information is included from SMNS 4378 (squamosal-opisthotic contact), SMNS 4379 (supraoccipital shelf), and data extrapolated from Nicrosaurus species B BMNH 42745 (supraoccipital-squamosal contact).

Abbrev.: boc, basioccipital; bs, basisphenoid; ect, ectopterygoid; eoc, exoccipital; opo, opisthotic; par, parietal; po, postorbital; proc., process; pt, pterygoid; q, quadrate; qj, quadratojugal; soc, supraoccipital; sq, squamosal.

428 I~~~ ____..I-JJ

UlQ. Figure 2.42: Nicrosaurus kapjfi (MEYER, 1860), reconstruction of the life appearance of the animal based on SMNS 4378.

429

Figure 2.43: Nicrosaurus kapJfi (MEYER, 1860), skulls in dorsal view illustrating the variants of four individually variable characters.

A SMNS 5726 B SMNS 5725

The anterior rims the external nasal openings lie at a considerable distance behind the anterior rim of the antorbital fenestra in SMNS 5726 (A), but terminate at level with that opening in SMNS 5725 (B). The external shape of the septomaxilla (dotted) is narrow and tapering in SMNS 5726 (A), but broad and multipronged in SMNS 5725 (B). The jugal (shaded) is excluded from the antorbital fenestra in SMNS 5726 (A), but forms the posteroventral rim of that opening in SMNS 5725 (B). The jugal of SMNS 5726 (A) touches the orbit on the left-hand side, but does not contribute to the orbital rim on the right-hand side, as in SMNS 5725 (B).

Abbrev.: f, frontal; l,lacrimal; m, maxilla; n, nasal; par, parietal; pm, premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; q, quadrate; qj, quadratojugal; sq, squamosal.

430 100 mm

100 mm Figure 2.44: Nicrosaurus kapffi (MEYER, 1860), sexual dimorphism.

Reconstructions of the high-domed morph (top line), based on SMNS 4378 (A, B) and BMNH 42743 (C), and the low-domed morph (bottom line), represented by SMNS 5727 (D-F).

A, D Reconstruction of the skull in occipital aspect. Note the narrower skull with steeper cheeks and the more parallel parieto-squamosal bars in the high-domed specimen. The different shape of the paroccipital process represents individual variation. Modified after MEYER (1863: pI. 41 fig. 4). n, E Skulls in right lateral view. Note the different orientation of the orbits and the outline of the prenarial crest. C, F Hypothetical cross-section through the snout at the premaxilla-maxilla suture, drawn to the same scale as B and E. Note the narrower snout and the less well-rounded crest top in the high-domed morpho

431 LL

: : : : \: /' ~ ..... ~

E E -8

« Figure 2.45: Nicrosaurus species B, SMNS 4059, dorsal view.

Modified after MEYER (1861: pI. 28), with most of the snout omitted, to illustrate the present state of preservation. The configuration of the skull elements is indicated according to personal observation.

The right side, which is separated from the left by a cleft and tucked underneath, is graphically rotated around the median axis of the skull in a perspective corresponding to the dorsolateral aspect. For this reason, the external nasal openings appear to be widely separated. Two osteoderms of unknown affinities are attached to the right side of the snout.

Abbrev.: f, frontal; 1, lacrimal; m, maxilla; n, nasal; par, parietal; pm, premaxilla; po, postorbital; pof, postfrontal; q, quadrate; qj, quadratojugal; sq, squamosal. Dotted area: obscured by matrix; hatched areas: restored parts or surfaces; irregularly dashed areas: damaged bone surfaces.

432 right squamosal process 100 mm Figure 2.46: Nicrosaurus species B, SMNS 4059, ventral view.

Modified after MEYER (1861: pI. 29 fig. 1) to illustrate the present state of preservation. The configuration of the skull elements is indicated according to personal observation.

Abbrev.: ect, ectopterygoid;j,jugal; m, maxilla; opo, opisthotic; pal, palatine; par, parietal; po, postorbital; pm, premaxilla; pt, ptyerygoid; q, quadrate; qj, quadratojugal; v, vomer. Dotted area: obscured by matrix; hatched areas: restored parts or surfaces; irregularly dashed areas: damaged bone surfaces.

433 nutricnt 1J..1~U4--'oramina

qj __--rJ"

skun roof baslpterygold 100 process mm squamosal procosses of par supratemporal fenestra Figure 2.47: Nicrosaurus species B, SMNS 12593, dorsal view.

Abbrev.: f, frontal; j, jugal; 1, lacrimal; m, maxilla; n, nasal; par, parietal; pm, premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; q, quadrate; qj, quadratojugal; soc, supraoccipital; sm, septomaxilla; sq, squamosal. Dotted area: matrix; hatched areas: restored parts or surfaces; irregularly dashed areas: damaged bone surface. Photo: S. POWELL (Bristol).

434 100 mm

internal antortlital lamina

sq Figure 2.48: Nicrosaurus species B, SMNS 12593, details of the temporal region.

A Temporal region in dorsal view.

B Cross section of the postorbito-squamosal bar at the posterior edge of the infratemporal fenestra drawn to the same scale. Note the thin medial lamella of the squamosal.

Abbrev.: opo, opisthotic; par, parietal; q, quadrate; sq, squamosal. Dotted areas: obscured by matrix; hatched area: restored parts or surfaces; irregularly dashed areas: damaged bone surfaces.

435 B

par po supratemporal fenestra

quadratojugal process of sq

foramen magnum supraoccipital sholl posnemporal sqamosal fenestra opisthotic process of par process of sq parietal process of sq A Figure 2.49: Nicrosaurus species B, SMNS 12593, ventral view.

Abbrev.: boc, basioccipital; bs, basisphenoid; ect, ectopterygoid; j, jugal; m, maxilla; opo, opisthotic; pal, palatine; par, parietal; po, postorbital; pm, premaxilla; pt, ptyerygoid; q, quadrate; qj, quadratojugal; sq, squamosal; v, vomer; ? represents probably a fish bone included in overlying matrix.

Dotted areas: matrix; irregularly dashed areas: damaged bone surface. Photo: S. POWELL (Bristol)

,- 436 100 mm

paroccipital process Figure 2.50: Nicrosaurus species B, SMNS 12593, palatine and choanal region in ventral view.

A Prechoanal part of the palate in ventral view.

B Left palatine in medioventral view with four hypothetical transverse cross-sections. The most posterior section is drawn after the section caused by the tenninal fracture. Note the different scale of the drawing and the cross-sections.

Abbrev.: m, maxilla; pal, palatine; pm, premaxilla; v, vomer. Dotted areas: matrix; irregularly dashed area: damaged bone surfaces.

437 pm pm

alveoli (covered)

...... 1 10mm

palatal ridge

pal m

8 Figure 2.51: Nicrosaurus species B, SMNS 12593, left side of the occipital region in ventral view.

Abbrev.: boc, basioccipital; bs, basisphenoid; eoc, exoccipital; opo, opisthotic; pt, pterygoid; q, quadrate. Dotted area: matrix; irregularly dashed area: damaged bone surface.

438 bs

fontanene . ~ <.;;~:.(~.;..... ; ::~':.:':.

'bort/enack' of fracture stapedial groove

median ridge ofboe

stapedial groove

OCCipital pterygo condyle quadrate plate

pterocCipital fenestra boc

- ••• -=:J 101lll1• paroccipital process of cpo Figure 2.52: Nicrosaurus species B, SMNS l2593, lateral view of the right side.

Abbrev.: ect, ectopterygoid; f, frontal; j, jugal; I, lacrimal; m, maxilla; n, nasal; pm, premaxill a; po, postorbital; pof, postfrontal; q, quadrate; qj, quadratojugal; sm, septomaxilla; sq, squamosal. Dotted areas: matrix; hatched areas: restored parts or surfaces; irregularly dashed area: damaged bone surfaces. Photo: S. POW ELL (Bristol)

439 ~ 8- CT Figure 2.53: Nicrosaurus species B, SMNS 12593, left posterior part of the skull in lateral view.

Abbrev.: j, jugal; par, parietal; po, postorbital; pof, postfrontal; q, quadrate; qj, quadratojugal; sq, squamosal.

Hatched areas: restored parts or surfaces; irregularly dashed area: damaged bone surface or cross-section of bone.

440

Figure 2.54: Nicrosaurus species B, SMNS 12593, occipital view from ventral.

Abbrev.: b c, basioccipital; bs, basisphenoid; opo, opisthotic; par, parietal; pt, pterygoid; q, quadrate; cU, quadratojugal; oc, supraoccipital; sq, squamosal. Dotted areas: obscured by matrix; hatched area: restored parts or surfaces; irregularly dashed areas: damaged bone surfaces. Photo: S. POWELL (Bristol)

441 50mm

edge of soc supraOCCIpital shetl

paroccipital process of sq

_ DOst omooral . les:ru

tuberal surface of boc tuberal sur1ace of bs Figure 2.SS: Nicrosaurus species B, BMNH 42745, reconstruction of the skull in dorsal view.

Specimen in the illustration of MEYER (1865b: pI. 25 fig. 1) to the left. The right figure illustrates the present state of preservation with the configuration of the skull elements indicated according to personal observation. Missing areas of the skull are omitted. The original state of preservation is figured by HUENE (1911: pI. 15).

Abbrev.: f, frontal; j , jugal; 1, lacrimal; m, maxilla; opo, opisthotic; pal, palatine; par, parietal; po, postorbital; pof, postfrontal; pm, premaxilla; prf, prefrontal; pt, ptyerygoid; q, quadrate; qj, quadratojugal; sm, septomaxilla; soc, supraoccipital; sq, squamosal; v, vomer; Hatched area: restored parts or surfaces; irregularly dashed areas: damaged bone surfaces.

442 100 mm

paroccipital squamosal process of SQ process of par Figure 2.56: Nicrosaurus species B, BMNH 42745, reconstruction of the skull in ventral view.

Specimen in the illustration of MEYER (l865b: pI. 25 fig. 1) to the left. The right figure illustrates the present state of preservation with the configuration of the skull elements indicated according to personal observation. Missing areas of the skull are omitted. The original state of preservation is figured by HUENE (1911: pI. 15).

Abbrev.: boc, basioccipital; bs, basisphenoid; ect, ectopterygoid; j, jugal; m, maxilla; opo, opisthotic; pal, palatine; par, parietal; po, postorbital; pm, premaxilla; pt, ptyerygoid; q, quadrate; qj, quadratojugal; sq, squamosal; v, vomer. Dotted area: obscured by matrix; hatched areas: restored parts or surfaces; irregularly dashed areas: damaged bone surfaces.

443 If:) 0'4' 0 J ~. ? C ' :. C ') (. 100 mm ;; (; r • ,c. c ,.., (.

"I b .( ") -. , .-r r; r, f "

" -. r 0 ' 1 0" C l... ~, \...." ( , . '" " ty j .' i t"I - ' "" C: ,... ,~ J- •• '",. .. Figure 2.57: Nicrosaurus species B, BMNH 42745, reconstruction of the skull in lateral right view.

Specimen in the illustration of MEYER (1865b: pi. 25 fig. 1) to the left. The right figure illustrates the present state of preservation with the configuration of the skull elements indicated according to personal observation. Missing areas of the skull are omitted. The original state of preservation is figured by HUENE (1911: pi. 15).

Abbrev.: ect, ectopterygoid; f, frontal; j , jugal; 1, lacrimal; m, maxilla; n, nasal; pm, premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; q, quadrate; qj, quadratojugal; sm, septomaxilla; sq, squamosal. Hatched areas: restored parts or surfaces; irregularly dashed area: damaged bone surface.

444 c:- ~c. ';cE_ '., ca :> E.c: '.~ CD . ~ a.

~~ . ~ '· .. t . L . ~ . 0 r. I ' ( 'U , ~ ~ '1 i 10 ~\ qr U 1,1 ., ~ I Q " " 'i ~-.\ I

E E ~ Figure 2.58: Nicrosaurus species B, BMNH 42745, occipital view of otic region from right side and ventral.

The base of the paroccipital process of the opisthotic is highlighted. The foramen jugale is subdivided, and the foramen nervi hypoglossi (XII) is located in the jugular recess.

Abbrev.: opo, opisthotic; pro, prootic. Dotted area: obscured by matrix; irregularly dashed area: damaged bone surface.

445 jugular ----~4-:=!ifl.:8_~;.._;:;~.. recess

foramen jugulare

XII

pro

stapedial groove Figure 2.59: Nicrosaurus species B, BMNH 42745, occipital view of the braincase.

Abbrev.: boc, basioccipital; bs, basisphenoid; eoc, exoccipital; soc, supraoccipital; sq, squamosal. Hatched area: restored surface; irregularly dashed areas: damaged bone surfaces.

446 lamina of par onto supraoccipital shell

postlemporal fenestra ---_·1::'.:;,,;..;.. :~~ parieto-squamosal bar

parietal process of sq edge of supraOCCIpital shell

bee

10 ~ mm~ Figure 2.60: Nicrosaurus species B, GPIT 2070.001, dorsal view.

Abbrev.: f, frontal; j, jugal; m, maxilla; n, na al; pm, premaxilla; po, postorbital. Hatched areas: restored part or surfaces. Photo and print: W . GERBER, GPIT.

447

Figure 2.61: Nicrosaurus species B, GPIT 2070.001, detail of the palate.

Abbrev.: m, maxilla; pal, palatine; pm, premaxilla; v, vomer. Hatched areas: restored parts or surfaces. Photo: W. GERBER, GPIT.

448 10 S" mm e Figure 2.62: Nicrosaurus species B , GPIT 2070.001, lateral view of the right side.

Abbrev.: j, jugal; m, maxilla; po, postorbital. Hatched areas: restored parts or surfaces. Photo and print: W. GERBER, GPIT.

449

Figure 2.63: Nicrosaurus species B, SMNS uncat. no. 11, lateral view.

Abbrev.: m, maxilla; n, nasal; pm, premaxilla; sm, septomaxilla. Dotted area: matrix; hatched areas: restored surfaces; irregularly dashed areas: damaged bone surfaces.

450 E E c 8

E 0. in Q) U "iii "C: c '"e! 0. Figure 2.64: Nicrosaurus species B, SMNS uncat. no. 11, medial view.

The orientation of the bone trabeculae is indicated by dashes. Arrows indicate the extent of the external nasal opening.

Abbrev.: n, nasal; pm, premaxilla; sm, septomaxilla. Dotted areas: matrix.

451

Figure 3.1: Mystriosuchus planirostris (MEYER, 1863), skull in dorsal and lateral view.

Skull morphology is taken from MCGREGOR (1906: pIs. 6, 7). The configuration of the temporal region is modified based on SMNS 13007, the narial region on SMNS uncat. no. 184.

Abbrev.: boc, basioccipital; bs, basisphenoid; j, jugal; m, maxilla; n, nasal; par, parietal; po, postorbital; pof, postfrontal; pm, premaxilla; prf, prefrontal; q, quadrate; qj, quadratojugal; sm, septomaxilla; soc, supraoccipital; sq, squamosal; v, vomer.

452

Figure 3.2: Mystriosuchus pianirostris (MEYER, 1863), SMNS 13007, occipital region in dorsal view.

Abbrev.: eoc, exoccipital; opo, opisthotic; par, parietal; soc, supraoccipital; sq, squamosal. Dotted areas: matrix.

453 supratemporal descending lamina fenestra 01 parieto-squamosal bar

squamosal squamosal process of par process 01 par supraoccipital lamina of par parietal process 01 sq

parietal process of sq

paroccipital II:I:III1 process 01 opo 10mm Figure 3.3: Mystriosuchus planirostris (MEYER, 1863), right squamosal and paroccipital process in posterolateral (A, C) and dorsal (B, D) view to illustrate the variation. Without scale.

A, B: SMNS uncat. no. 184. The extremity of the squamosal is constricted fOrming a blunt knob; the descending part of the squamosal shows a broad furrow; there is a deep and long recess between the squamosal and the opisthotic process leading into the posttemporal fenestra.

C, D SMNS 10260. The tip of squamosal is pointed but without a knob-like extension; the descending part of squamosal narrow and rounded without a furrow; there is a short triangular recess between the squamosal and the opisthotic process.

Irregularly dashed areas: damaged bone surface.

454 nro:l 10mm B

II:I1

c Figure 3.4: Mystriosuchus species B, GPIT 261/001, lateral view of the right side.

Abbrev.: f, frontal; j, jugal; 1, lacrimal; m, maxilla; n, nasal; par, parietal; pm, premaxilla; pof, postfrontal; prf, prefrontal; q, quadrate; qj, quadratojugal; sm, septomaxilla. Small arrows indicate the anterior and posterior rim of the external nasal opening.

455

Figure 3.5: Mystriosuchus species B, GPIT 261/001, occipital view.

Abbrev.: boc, basioccipital; ect, ectopterygoid; eoc, exoccipital; opo, opisthotic; par, parietal; po, postorbital; pt, pterygoid; q, quadrate; qj, quadratojugal; soc, supraoccipital; sq, squamosal; t, supernumerary occipital bone. Stippled areas: matrix; hatched areas: restored surface; irregularly dashed areas: damaged bone surface. Photo: GPIT, HUENE-archive; print: W. GERBER, GPIT.

456 ledge inclined medial 01 par plane of sq

parieto postero-Iateral squamosal bar surface of sq

triangular recess

posttemporal ...... ,~~~'=~!~~=====~~~:-~4- fenestra

--=:::::------i~-_L_ posterior ridge

pterygoid wing 01 q

pt 111M - -100 mm - - Figure 3.6: Mystriosuchus species B, GPIT 261/001, cross-section through the right postorbito-squamosal bar, posterior aspect.

Figure 3.7: Mystriosuchus species B, GPIT 2611001, dorsolateral view of the right temporal region seen through the supratemporal fenestra.

Abbrev.: ep, epiotic; Is, 1aterosphenoid; par, parietal; pro, prootic; sq, squamosal; t, supernumerary occipital bone. Without scale.

457 ovemang --

Inclined lateral medial surlace -----squamosal ridge rimot Infratemporal fenestra

~ second 10mm ridge

parietal ledge

supratemporal fenestra supraoccipilal shell

crecenl-shaped crest

vena capitis dorsahs

parielal process 01 sq Figure 3.8: Mystriosuchus species B, GPIT 2611001, posterolateral view of the right otic region and the pterygo-quadrate plate.

Abbrev.: boc, basioccipital; bs, basisphenoid; eoc, exoccipital; opo, opisthotic; pro, prootic; pt, pterygoid; q, quadrate; Roman numerals indicate cranial nerves. Stippled areas: matrix.

458 paroccipital process 01 opo

pteroccipital fenestra

ventrolateral ridge 01 boc

_ 1 pterygoid .. -100mm-- procesS 01 bs Figure 3.9: Mystriosuchus species B, GPIT 261/001, right lateral view of the braincase.

Abbrev.: boc, basioccipital; bs, basisphenoid; eoc, exoccipital; ep, epiotic; ept, epipterygoid; Is, laterosphenoid; opo, opisthotic; os, orbitosphenoid; par, parietal; pro, prootic; ps, parasphenoid; pt, pterygoid; q, quadrate; t, supernumerary occipital bone; Roman numerals indicate cranial nerves. Stippled areas: matrix.

459 par

parleto squamosal bar posnemporat fenestra ?hypophyseal 10ramen

lamina olsq

eoe

bs 50mm inlemal carotid anery Figure 3.10 (top): Mystriosuchus species B, referred snout fragment GPIT 26111717 in ventral view. The anterior end of the snout is to the right.

Fragmentary remains of a large phytosaur, perhaps Mystriosuchus species B, from quarries in the region around Trossingen and Aixheirn.

Figure 3.11 (left-hand side): GPIT 2145.000, postsymphyseal fragment of a mandibular ramus in dorsal view. The medial side is to the right.

Figure 3.12 (right-hand side): GPIT 2146.000, postsymphyseal fragment of a right mandibular ramus in dor al view. The anterior end is at the top.

460 100mm

50 mm

50 mm Figure 5.1 (top): Cladogram resulting from a cladistic analysis ofPhytosauria, modified from BALLEW (1989: fig. 1).

Paleorhinus comprises Ebrachosuchus, Francosuchus, Mesorhinosuchus, Paleorhinus, "Parasuchus" and Promystriosuchus, and Angistorhinus includes Angistorhinus and Brachysuchus megalodon of this study. The generic composition of Rutiodon and Pseudopalatus in BALLEW (1989) is indicated in the top line.

Figure 5.2 (bottom): Cladogram constructed on the basis of the taxonomic scheme of Phytosauria presented in LONG & MURRY (1995).

Paleorhinus comprises Ebrachosuchus, Francosuchus, Mesorhinosuchus, Paleorhinus , "Parasuchus" and Promystriosuchus, and Angistorhinus includes Angistorhinus and Brachysuchus megalodon of this study. Arribasuchus includes Pseudopalatus buceros and Pseudopalatus mccauleyi of BALLEW (1989). The taxa which were defined by shared derived characters by LONG & MURRY are underlined.

461 Rutiodon Pseudopalatus sensu BAllEW (1989) sensu BAlLEW (1989)

T I I I I I

I BALLEW (1989)

Mystriosuchidae Rutiodonlidae

Paleo- ~ Angisto Rutlodonlioae PseydQpalatjnae rhininae ~ mininae r---1 r---1 ,...---,

T T

I I LONG &MURRY (1995) Figure 5.3 (top): Consensus tree (50% majority-rule) resulting from 5 most parsimonious trees obtained by equal weight parsimony analysis of 49 characters (see Appendix A, Table A.l).

The numerals indicate the percentage of the trees which include the clade.

Figure 5.4 (bottom): Consensus tree (50% majority-rule) resulting from 35 most parsimonious trees obtained by parsimony analysis of 49 characters.

Characters 1,6, 10, 16, 18,20,24,28,30,33,37,44, and 48 of the data matrix (see Appendix A, Table A.l) were treated as ordered (linear). The numerals indicate the percentage of the trees which include that particular clade.

462 1st analY multista tes~~aracters unordered

2nd characters 13 multlstaan~IYStiS: e ordered Figure 5.5: Strict consensus tree calculated from trees 10,25, and 26 out of the 35 most parsimonious trees from Figure 5.4, selected after obtaining a similar tree by successive approximations character weighting optimised to rescaled consistency index (RCI) and consistency index (CI).

The nodes that define the taxa treated in the text are indicated by letters. Note that only previously established suprageneric taxa, or their synonymous clades, are named. The numbers indicate the characters that diagnose a particular node. The state of multi state characters is given in parenthesis. Character states are explained in 5.3.2. Characters or character states that are homoplastic within phytosaurs are marked with an asterisk (*). Reversals are indicated by a minus symbol (-).

463 .~ ~ f ~ ~ rS .~ ~ ~"J ~ ~ ~,.~ ~f W .~ ~ ~ § 9 ~ 0 ~/I.~ I I ~ ~ i t .f ~ h~.~ .{{J~ /f i ~. J1~ ~o"J .f l' :J' ~9F~f9~~f§t!tl~/~f~#~~1 ~ § "Jtri ~~ ~ ! :$ ~ ~ ~ ~ ~~ ~'l '-J6 ~t$ tri ! {§ ~ ~ ~ ~ 9 .~ f or;; ;:,"J ,§l ~ .or-j 9 ~ ~ ~ '§ ·f·~ ~ ~ 9 c1i /~ ~{J ~;s '::-.rti ~ ~ $ ~ .~ .~ & ~ ~ ~ .~ .~ .~ ~ '{! f: ~,§' '§ ~ ~- ~- ,§l ~ ~ ~ f .$0 f f i ~o ~i? i flJf flJf flJf -I .j .j #::5 .~'-j #::5 .o~ ~tri ~tri .~~ l ~~ if ~~~~~~~~~~~~~~~4~~##~1~~

"-IC:11*, K M -24 LJ~Y"- "- "- "- I G, -13, I, Angistorhlnin.. ~ I 0, 1[2",6[21',23, 18[2], -41 21,24*,34, -37 I 24[2]*,33[3],47[2]* N: Pseudopalatinae 20[2]*, 37[2]*

"H: Phytosauridae 8*,10[3],16[2]*,-26, -27, 30[2]*, 39

"-0: 40*, 44[3]*

"B: 13*,19,41*

"-A: Phytosauria Figure 5.6A-C (top): Equally parsimonious alternatives to the configuration of the interrelationships of Rutiodon carolinensis, Leptosuchus crosbiensis, Leptosuchus adamanensis, and Smilosuchus gregorii illustrated in Figure 5.5. All subclades are realised among the 35 most parsimonious trees of Figure 5.4.

The numbers indicate the characters that diagnose a particular node. The state of multi state characters is given in parenthesis. Character states are explained in 5.3.2, and for a discussion of the character distribution, see 5.4.3.1. Characters or character states that are homoplastic within phytosaurs are marked with an asterisk (*). Reversals are indicated by a minus symbol (-).

Figure 5.7 (bottom): Cladogram resulting from a bootstrap analysis (algorithm: heuristic, 1000 replications) on the basis of the data matrix Appendix A, Table A.1.

Characters 1,6, 10, 16, 18,20,24,28,30,33,37,44, and 48 were treated as ordered (linear). The numerals indicate the per cent distribution of a clade among the resulting MPTs of all replications (only values> 50% are included).

464 " 1[2]*,28[1]* "16[2]*, -27* no ~ ~-'T'::"'_...J autapomorphy ~_...... ,.._--J

A B c •

L..

7 1 0

1 7 l2 Y 15 8 Bootstrap analysis: 13 multistate characters ordered