Revision of the Sauropterygian Reptile Genus Cymatosaurus V. Fritsch, 1894, and the Relationships of Germanosaurus Nopcsa, 1928, from the Middle Triassic of Europe

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1 GtOL06Y LIBRARY ,s. ______^^ 0.36FIELDIANA

Geology NEW SERIES, NO. 36

Revision of the Sauropterygian Reptile Genus Cymatosaurus v. Fritsch, 1894, and the Relationships of Germanosaurus Nopcsa, 1928, from the Middle Triassic of Europe

Olivier Rieppel

March 31, 1997 Publication 1484

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iJNOIS LIBRA* URBANA.CHAMPA FIELDIANA

Geology NEW SERIES, NO. 36

Revision of the Sauropterygian Reptile Genus Cymatosaurus v. Fritsch, 1894, and the Relationships of Germanosaurus Nopcsa, 1928, from the Middle Triassic of Europe

Olivier Rieppel

Department of Geology Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 U.S.A.

Accepted June 28, 1996 Published March 31, 1997 Publication 1484

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1997 Field Museum of Natural History ISSN 0096-2651 PRINTED IN THE UNITED STATES OF AMERICA GE0L06Y LIBRARY o^ Table of Contents 3. Fragmentary skull referred to Cymato- saurus fridericianus 8 4. Snout fragment referred to Cymatosau-

Abstract 1 rus cf. C fridericianus 8 zusammenfassung 1 5. The neotype of Cymatosaurus latifrons Introduction 1 (Gurich, 1884) 10 Systematic Paleontology 3 6. The neotype of Cymatosaurus latifrons Cymatosauridae Huene, 1948 3 (Gurich, 1884) 11 Cymatosaurus v. Fritsch, 1894 3 7. The holotype of Cymatosaurus ery- Cymatosaurus fridericianus v. Fritsch, threus E.v. Huene, 1894 12 4 1894 8. The holotype of Cymatosaurus ery- Morphological Description 4 threus E.v. Huene, 1894 13 6 Cymatosaurus latifrons (Gurich, 1884) 9. Isolated parietal referred to Cymatosau- 10 Morphological Description rus erythreus E.v. Huene, 1894 15 multidentatus Huene, Cymatosaurus (Fv. 10. The holotype of Germanosaurus schqf- 14 1958) feri (Arthaber, 1924) 17 Cymatosaurus sp. ("C. erythreus" E.v. 1 1 . The holotype of Germanosaurus schqf- Huene, 1944) 14 feri (Arthaber, 1924) 18 Nothosauridae Baur, 1889 15 12. Isolated parietal referred to Germano- Germanosaurus Nopcsa, 1928 15 saurus schqfferi (Arthaber, 1924) 18 Germanosaurus sp. ("G. latissimus" 13. Phylogenetic interrelationships of the [Gurich, 1884]) 16 Sauropterygia 19 Germanosaurus schqfferi (Arthaber, 1924) 14. Phylogenetic interrelationships of the 18 Reptilia 19 Morphological Description 18 15. Clade rank determination in Cladistic Analysis 22 Sauropter- 20 Phylogenetic Pattern and Stratigraphic ygia 16. Clade versus rank in .. 20 Superposition 29 age Sauropterygia Acknowledgments 33 Literature Cited 33 Appendix 35 List of Tables

List of Illustrations Skull proportions in Cymatosaurus, Ger- manosaurus, and Nothosaurus 9

. Data matrix for the cladistic of 1 The holotype of Cymatosaurus frideri- analysis cianus v. Fritsch, 1894 all Cymatosaurus species 18 2. The holotype of Cymatosaurus frideri- Data matrix for the cladistic analysis of cianus v. Fritsch, 1894 sauropterygian interrelationships 21

UBRARYU. OF!. UHBANA-CHAMPAIGN

Revision of the Sauropterygian Reptile Genus Cymatosaurus v. Fritsch, 1894, and the Relationships of Germanosaurus Nopcsa, 1928, from the Middle Triassic of Europe

Olivier Rieppel

Abstract

Three species are currently recognized within the genus Cymatosaurus from the late Scythian and early Anisian of Europe, viz. Cymatosaurus fridericianus v. Fritsch, 1894, Cymatosaurus latifrons Giirich, 1884, and Cymatosaurus multidentatus (F.v. Huene, 1958). All other previously described species of Cymatosaurus are considered either junior synonyms of Cymato- saurus latifrons (C. gracilis Schrammen, 1899; C. silesiacus Schrammen, 1899) or a nomen dubium (C. erythreus E.v. Huene, 1944). Germanosaurus schqfferi Arthaber, 1924, is recognized as a separate genus and species within the Nothosauridae, sister-group of the Notho- saurinae (including Nothosaurus and the Silvestrosaurus-Ceresiosaurus-Lariosaurus clade). Germanosaurus (JEurysaurus) latissimus (Giirich, 1891) is treated as a nomen dubium. A cladistic analysis based on the critical revision of the genera Cymatosaurus and Germanosaurus improves resolution among Triassic stem-group Eosauropterygia. The resulting cladogram is used as the basis for a comparison of phylogenetic pattern and stratigraphic distribution of the Sauropterygia.

Zusammenfassung

Innerhalb der Gattung Cymatosaurus aus dem obersten Skyth und unteren Anis Europas werden gegenwartig drei Arten als giiltig anerkannt: Cymatosaurus fridericianus v. Fritsch, 1894, Cymatosaurus latifrons Giirich, 1884, und Cymatosaurus multidentatus (F.v. Huene, 1958). Alle anderen beschriebenen Arten von Cymatosaurus sind entweder jungere Synonyme von Cymatosaurus latifrons (C. gracilis Schrammen, 1899; C. silesiacus Schrammen, 1899) oder ein nomen dubium (C. erythreus E.v. Huene, 1944). Germanosaurus schqfferi Arthaber, 1924, wird als selbtandige Gattung und Art innerhalb der Nothosauridae anerkannt, die die Schwestergruppe der Nothosaurinae (Nothosaurus, "Silvestrosaurus," "Ceresiosaurus," und Lariosaurus) reprasentiert. Germanosaurus (Eurysaurus) latissimus (Giirich, 1891) ist ein nomen dubium. Eine kladistische Analyse, die auf der kritischen Revision der Gattungen Cy- matosaurus and Germanosaurus fusst, resultiert in einem besseren Verstandnis der Verwandt- schaftsverhaltnisse unter triassischen Eosauropterygiern. Auf der Grundlage dieser Analyse wird das Muster der Verwandtschaftsbeziehungen mit dem Muster der stratigraphischen Ver- breitung der Sauropterygia verglichen und diskutiert.

Introduction tened skull (and an isolated snout fragment) collected in the lowermost Muschelkalk (mu,, lower The genus Cymatosaurus was erected by Anisian) exposed in the quarries of the "Portland- Fritsch (1894) for an isolated, dorsoventrally flat- Cement-Fabrik" at Halle/Saale, Germany. Fritsch

FIELDIANA: GEOLOGY, N.S., NO. 36, MARCH 31, 1997, PP. 1-38 1 (1894) recognized many similarities that the spec- can no longer be located today) figured a skull imen shares with Nothosaurus, but it differs from from the lower Muschelkalk of Muhlhausen (Thu- the latter genus in having strongly reduced nasals, ringia), which he referred to "Nothosaurus (Cy- a trait that Cymatosaurus shares with Pistosaurus matosaurus) cf. fridericianus." This skull is clear- from the upper Muschelkalk (mo,) of Bayreuth ly that of Nothosaurus marchicus (Rieppel & (Bavaria). Wild, 1996), as is indicated by its proportions (rel- Sauropterygia of equivalent or slightly older atively short rostrum, relatively small upper tem- age (mu,, lowermost Muschelkalk) had previously poral fossae, maxillary tooth row not extending been collected in the Gogolin beds of Upper Si- up to the midpoint of the upper temporal fossa), lesia, and isolated skulls from those deposits had the well-developed nasals, the fused frontals, and been described as Nothosaurus latifrons Gurich, the relatively forward position of the pineal fo-

1884, ox Nothosaurus latissimus Gurich, 1891, re- ramen (see also Schroder, 1914, p. 73). spectively. Fritsch (1894) did not refer to Gurich's Huene (1958) described a new species of the (1884, 1891) material, although in his review of pachypleurosaur genus Anarosaurus Dames, the genus Nothosaurus Koken (1893) had sug- 1890, from the lower Anisian of the Lechtaler gested that Gurich's (1891) species might be re- Alps (Austria), Anarosaurus multidentatus. Re- ferred to a different genus. Schrammen (1899) re- description of the holotype resulted in its identi- viewed the nothosaurs from the lower Muschel- fication as Cymatosaurus (Rieppel, 1995c), indi- kalk of Upper Silesia, referred both of Gurich's cating that the genus had expanded from the Mu- species to the genus Cymatosaurus, and described schelkalk Basin and reached the Alpine Triassic two new species, viz. Cymatosaurus gracilis and during early Anisian times (Rieppel & Hagdorn, C silesiacus. Morphological comparison led him 1996). Throughout the Germanic and Alpine Tri- to conclude that Cymatosaurus must be close to assic, the genus Cymatosaurus remains restricted the common ancestor of Nothosaurus and Pisto- to the lower Muschelkalk and lower Anisian re- saurus. spectively. Productive deposits {orbicularis beds, Freeh (1903) erected the new subgenus Eury- basal middle Muschelkalk) of Esperstadt, Jena, saurus to include Cymatosaurus latissimus, which Querfurt, and Rudersdorf have yielded abundant he found to be intermediate between Cymatosau- material of Nothosaurus (Rieppel & Wild, 1996), rus and Nothosaurus: the frontals are paired and but not a single diagnostic skull fragment of Cy- the nasals reduced in Eurysaurus latissimus as in matosaurus. Cymatosaurus, but the reduced nasals still reach Analysis of the invertebrate fauna from lower the posterior margin of the external naris as in Muschelkalk deposits in the eastern part of the Nothosaurus. Nopcsa (1928a, p. 21, 1928b, p. Germanic Basin shows strong Asiatic affinities 173) replaced the genus name Eurysaurus by Ger- and indicates that these taxa reached the Muschel- manosaurus because the first name was preoccu- kalk Basin from the Paleotethys through the East pied. This had escaped Arthaber (1924), who in Carpathian gate (Kozur, 1974; Hagdorn, 1985; his review of nothosaurs retained Eurysaurus as Urlichs & Mundlos, 1985). In view of this paleo- a subgenus of Cymatosaurus and described a new biogeographical context, it is interesting to note species, Eurysaurus schafferi, from the lowermost that the only possible Cymatosaurus described Muschelkalk (mu,, Gogolin beds) of Gogolin, Up- from outside Europe is Micronothosaurus sten- per Silesia. sioei Haas, 1963, from the upper Muschelkalk of The genus Cymatosaurus was comprehensively Wadi Ramon, Israel. The specimen differs from reviewed by E.v. Huene (1944), who described Nothosaurus in a number of characteristics, such another species, Cymatosaurus erythreus, from as the relatively forward position of the pineal the Buntsandstein of distinct the upper (so 2 , Rot) Rudersdorf foramen, the posterolateral lappets of near Berlin, the earliest representative of its genus frontal approaching the pineal foramen, a narrow in the Germanic Triassic. E.v. Huene (1944) re- postorbital bridge, relatively small upper temporal tained Germanosaurus as a subgenus, to which fenestrae compared to the size of the orbits, no she referred the species gracilis Schrammen, la- evidence for a posterior extension of the tooth row tissimus Gurich, schafferi Arthaber, and silesiacus beyond the posterior margin of the orbit, and a Schrammen; Cymatosaurus ss. would include the poorly ossified occiput with large posterior open- species erythreus E.v. Huene, fridericianus ings for the cranioquadrate passage. Unfortunate-

Fritsch, and latifrons Gurich. ly, some morphological details remain obscure in Jaekel (1911, p. 148, Fig. 161; this specimen Haas's (1963) description, such as the paired or

FIELDIANA: GEOLOGY unpaired condition of the frontal and the relations ciput deeply concave; supraoccipital vertically of the jugal and postorbital along the posterior oriented and in loose connection with the der- margin of the orbit. In view of the many shared matocranium; distinctly reduced nasals that may similarities, Schultze (1970) identified Microno- or may not enter the external naris; frontals thosaurus stensioei as a cymatosauroid, possibly paired; posterolateral processes of frontals closely even belonging to the genus Cymatosaurus. If this approach upper temporal fossa and may enter its identification is valid, and if Haas's (1963) indi- anteromedial margin; parietals incompletely or cation of its stratigraphic occurrence in the upper completely fused; jugal enters posterior margin of Muschelkalk is correct, the specimen would sig- the orbit and remains excluded from upper tem- nificantly expand the geological occurrence of the poral arch; quadratojugal absent. genus Cymatosaurus beyond the central and west- Distribution—Uppermost Buntsandstein and ern European occurrences into the uppermost An- lower Muschelkalk, lower Anisian, Middle Tri- isian or lower Ladinian. assic, Europe—and ?Israel. In fact, Haas (1963, p. 161) specifies that the Comments The cladistic analysis discussed specimen came from "Brotzen's layer D2 (Cera- below indicates a sister-group relationship of Cy- tites beds)," but "Nothosaurus or related genera" matosaurus and Pistosaurus, with Corosaurus are cited by Brotzen (1957, p. 202) as coming representing the sister-taxon of the former two. from the Ceratites zone Dl. The overlying Cer- An argument could therefore be made that the lat- atites zone D2 was correlated by Brotzen (1957) ter two genera be included in the Cymatosauridae. with the Trinodosus zone, and both zones (Dl and This conclusion is not formalized here, because D2) were considered equivalent to the Alpine up- the addition of plesiosaurs and pliosaurs to the per Anisian (Brotzen, 1957, p. 206). More recent analysis may show Pistosaurus to be the sister- analyses equate the Trinodosus zone of the Teth- taxon of plesio- and pliosaurs (see Sues, 1987; yan province with the lower Illyr, of lower upper Storrs, 1991, 1993b). The results reported here Anisian age, which corresponds to the upper low- support the concept of the Pistosauria proposed er Muschelkalk (mu 2 ) (Rieber, 1973; Bucher, by Edinger (1935; see also Sanz, 1983; Alafont & 1988; Budurov et al., 1993; Brack & Rieber, Sanz, 1996), to include Cymatosaurus, Pistosau- 1993). Although geologically younger than the rus, and, by extension, plesio- and pliosaurs. The occurrence of Cymatosaurus in the Gogolin beds Pistosauria may have to be extended to include of Upper Silesia, the Israel cymatosauroid still is Corosaurus, or a new higher taxon may have to in deposits equivalent to the lower Muschelkalk. be named to include Corosaurus and the Pisto- Institutional abbreviations are as follows: bgr, sauria. Bundesanstalt fur Geowissenschaften und Roh-

stoffe, Berlin; Ha, Institut fur Geowissenschaften, Martin-Luther-Universitat, Halle/Saale; Mbg, Cymatosaurus v. Fritsch, 1894 Fachbereich Geowissenschaften, Philipps Univer- sitat, Marburg/Lahn; nhmw, Naturhistorisches Museum, Wien; smns, Staatliches Museum fur 1884 Nothosaurus, Gurich, p. 132, PI. II, 2-4 Naturkunde, Stuttgart. Figs. 1891 Nothosaurus, Gurich, p. 967, Fig. on p. 968.

1894 Cymatosaurus, v. Fritsch, p. 10 Systematic Paleontology 1903 Eurysaurus, Freeh, p. 15. 1928a Cymatosaurus, Nopcsa, p. 44 Sauropterygia Owen, 1860 1928b Cymatosaurus, Nopcsa, p. 173. Eosauropterygia Rieppel, 1994a 1944 Cymatosaurus {Germanosaurus) (par- tim), E.v. Huene, p. 208. Cymatosauridae Huene, 1948 Type Species—Cymatosaurus fridericianus v. Definition—A monophyletic taxon including Fritsch, 1894, from the lower Muschelkalk (lower the genus Cymatosaurus. Middle Triassic), Halle/Saale, Germany. Diagnosis—Small to large eosauropterygians Definition—A monophyletic taxon including with a moderately depressed skull; snout con- the species fridericianus, latifrons, and multiden- stricted; postorbital skull distinctly elongated; oc- tatus.

RIEPPEL: REVISION OF CYMATOSAURUS Diagnosis—Same as for family, of which this sents the posterior part of the dermatocranium, is the only genus. from which most of the braincase has dropped out Distribution—Same as for family, of which (Fig. 3). This indicates incomplete ossification this is the only genus. and hence possibly a juvenile status of the spec- Comments—Lamprosauroides goepperti (Mey- imen (but see further discussion below). The fron- er, 1860; Lamprosauroides replaces Lamprosau- tal enters the anteromedial margin of the upper rus: K. P. Schmidt, 1927, p. 58) was considered a temporal fossa, a derived feature shared with the cymatosauroid by Schrammen (1899, p. 408). In holotype of Cymatosaurus fridericianus. The view of its fragmentary nature, Lamprosauroides specimen is therefore interpreted as a juvenile goepperti (Meyer, 1 860) remains a nomen dubium representative of the latter species. (Rieppel, 1995a). The snout fragment from the lower Muschel- kalk of Halle referred to Cymatosaurus sp. by Fritsch (1894, p. 300, PL 16, Fig. 2) is not diag- Cymatosaurus fridericianus nostic at the species level but differs from Cy- v. Fritsch, 1894 matosaurus fridericianus only by its somewhat smaller size. The specimen can no longer be lo- 1 894 Cymatosaurus fridericianus, v. Fritsch, cated today. A similar snout fragment (smns 7209; p. 281 #, PL 16, Fig. 1; PL 17, PL 18, Fig. 4) is known from the lower Muschelkalk Figs. 1-12. (mu,) of Freudenstadt. 1899 Cymatosaurus fridericianus, Schram- Morphological Description—C. fridericianus men, p. 389 j/, PL 24, Figs. 5a-c. is characterized by a relatively long and slender 1914 Cymatosaurus fridericianus, Schroder, skull with a pronounced rostrum (Fig. 2). The oc- p. ISjf., Figs. 15, 21. cipital condyle is not preserved. Skull length from 1924 Cymatosaurus fridericianus, Arthaber, the tip of the snout to the posterior end of the p. 475, Figs. lOa-b. parietal skull table is 195 mm, the length from the 1928 Cymatosaurus fridericianus, Schmidt, tip of the snout to the mandibular condyle of the p. 393, Fig. 1105. quadrate approximately 238 mm. The occiput is 1934 Cymatosaurus fridericianus, Kuhn, p. deeply excavated, the mandibular articulations lo- 42. cated well behind the assumed level of the occip- 1944 Cymatosaurus fridericianus, E.v. Hu- ital condyle. The external nares are almost twice ene, pp. 198/ 207/ as long as they are broad, and the upper temporal 1964 Cymatosaurus fridericianus, Kuhn, p. 12 fossa is relatively somewhat smaller than in No- 1983 Cymatosaurus fridericianus, Sanz, Fig. thosaurus. Dividing the distance from the tip of lb. the snout to the anterior margin of the external 1995c Cymatosaurus fridericianus, Rieppel, p. naris by the width of the skull at the rostral con- 295, Figs. 8A-B. striction yields a value of 1.64 (1.2-2.5 in Notho- saurus). Dividing the distance from the tip of the Holotype—Skull (Ha, uncatalogued; Figs. 1, snout to the anterior margin of the orbit by the 2). distance from the tip of the snout to the anterior Locus Typicus—Lower Muschelkalk (mu,), margin of the external naris results in an index of Halle/Saale, Germany. 1.87 (1.5-2.0 in Nothosaurus). Dividing the dis- Diagnosis—A species of Cymatosaurus of tance from the tip of the snout to the anterior mar- large size (tip of snout to back end of parietal gin of the upper temporal fenestra by the distance skull table up to 200 mm); three maxillary teeth from the tip of the snout to the anterior margin of preceding paired maxillary fangs; nasals reduced, the external nares yields a ratio of 2.68 (2.3-3.4 excluded from external naris; prefrontal and post- in Nothosaurus). Dividing the longitudinal diam- frontal in contact at dorsal margin of orbit; frontal eter of the external naris by its transverse diameter enters anterior margin of upper temporal fossa. yields a ratio of 1 .94 (the corresponding ratio var- Distribution—Lower Muschelkalk (lower An- ies from 1.0 to 2.2 in Nothosaurus, depending on isian, lower Middle Triassic), central Europe. the species). Dividing the longitudinal diameter of Referred Specimens—bgr S 44/3: a small and the upper temporal fossa by the longitudinal di- incomplete skull from the lower Muschelkalk ameter of the orbit yields values of 1 .76 (left side) (Gogolin Beds, mu,) of Gogolin, Upper Silesia and 1.96 (right side), respectively (the corre- Rieppel, 1994a, Fig. 39). The fragment repre- sponding ratio varies from 2.1 to 3.9 in Notho-

FIELDIANA: GEOLOGY saurus, with N. marchicus at the lower end and compression of the skull. The premaxillary ros- N. mirabilis at the upper end of that range). trum carries five tooth positions on each side; re- The relatively long and slender rostrum, formed placement pits are distinctly elongated and drop- by the paired premaxillae, is distinctly set off shaped, located posteromedial to the functional from the maxillaries a rostral constric- bulging by tooth positions. Three small maxillary teeth pre- tion. suture is located The premaxillary-maxillary cede the paired maxillary fangs. The total count at the anterolateral corner of the external naris. of maxillary teeth cannot be established. The vo- of the Broad posterior (nasal) processes premax- mers are paired elements that meet the maxillae illa the external nares from one another separate at the anterior margin of the internal nares, thus and meet the frontals in an paired interdigitating excluding the premaxillae from the latter. The suture at the level of the anterior of the margin posterior margin of the internal nares is defined orbit. Between the external nares and the orbits, by the palatines. The internal nares are relatively the maxilla is broadened to accommodate the broad and short: division of their longitudinal di- roots of the The nasals are paired maxillary fangs. ameter by their transverse diameter yields a ratio small, bones whose reach- splint-like posterior tip of 2.1; corresponding values vary from 1.28 to es the level of the anterior of the but margin orbit, 3.86 in Nothosaurus, with N marchicus at the which remain excluded from the posterior margin lower end and N mirabilis at the upper end of of the external naris. The lacrimal is lacking. The that range. As compared to Nothosaurus, the in- prefrontal is relatively larger than in Nothosaurus, ternal nares are positioned somewhat farther back located at the anterodorsal corner of the orbit, and relative to the external nares (Table 1). it meets the postfrontal along the dorsal margin The vomers meet the anterior tips of the pter- of the orbit. The postfrontal is a large element ygoids in a transversely oriented interdigitating defining the posterodorsal margin of the orbit as suture behind the level of the posterior margins well as the anterior margin of the upper temporal of the internal naris. The ectopterygoid is a rela- fossa. The postorbital bridge is relatively narrower tively short and broad element located at the an- than in Nothosaurus. Dividing the width of the terior margin of the subtemporal fossa, and it postorbital bridge by the width of the maxilla be- forms a well-developed (ecto-)pterygoid flange tween the external naris and the anterior margin for the origin of the superficial pterygoideus mus- of the orbit yields a value of 0.36 for C. frideri- cle. The quadrate ramus of the pterygoid is well cianus; the corresponding ratio varies from 0.8 to preserved on the right side of the skull and (as in 1 .8 in Nothosaurus (depending on the species). Nothosaurus) shows well-developed flanges at The frontals are large, paired elements that both its medial and lateral edges for the origin of meet the fused parietal at a level somewhat behind the deep pterygoideus muscle. the anterior margin of the upper temporal fossa. Discussion—Among all described species of Unlike any specimens of Nothosaurus (and some Cymatosaurus, the frontal approaches the antero- skulls of Cymatosaurus [see below]), the frontal medial margin of the upper temporal fossa to a enters the anteromedial margin of the upper tem- variable degree. Only in Cymatosaurus friderici- poral fossa. The parietal is a narrow element with anus does the frontal enter the upper temporal fos- the pineal foramen located slightly anterior to its sa and participate in the formation of the ventrally midpoint. The suture between squamosal and pa- descending flange from which originate deep fi- rietal at the posterior margin of the upper tem- bers of the adductor muscles. Unlike in Cy- poral fossa cannot be identified unequivocally. jaw matosaurus three (rather than one) small The squamosal meets the postorbital in an over- latifrons, teeth the —the lapping suture within the upper temporal arch; the maxillary precede maxillary fangs condition compared to the out- anterior tip of the squamosal lies behind the level plesiomorphic The de- of the anterior margin of the upper temporal fossa. group (Nothosauridae: Rieppel, 1994b). of reduction of the nasals external The precise shape and relations of the jugal bone, gree (entering nares or excluded as well as the located at the posteroventral margin of the orbit, therefrom), pres- of a contact between cannot be identified. It did not, however, extend ence or absence prefrontal of the or- into the upper temporal arch, approaching the an- and postfrontal along the dorsal margin is variable all the of terior tip of the squamosal. bit, highly among specimens The ventral view of the skull discloses a large Cymatosaurus ever described, and these charac- fontanelle between the premaxillae, enlarged by ters cannot be used to differentiate separate spe- separation of the premaxillae due to dorsoventral cies within the genus.

RIEPPEL: REVISION OF CYMATOSAURUS Fig. 1. The skull of the holotype of Cymatosaurus fridericianus v. Fritsch, 1894 (Ha, uncatalogued), from the = lower Muschelkalk of Halle/Saale. A, Dorsal view; B, ventral view. Scale bar 20 mm.

Cymatosaurus latifrons 1899 Cymatosaurus gracilis, Schrammen, p. (Gurich, 1884) 402, PI. 23, Figs. 2-3; PL 25, Figs. 6a- b, 7.

1884 Nothosaurus latifrons, Gurich, p. 132, 1899 Cymatosaurus silesiacus, Schrammen, PI. 2, Figs. 3-4. p. 402, PL 21, PL 22, Figs. 1-2; PL 25.

1893 Nothosaurus latifrons, Koken, p. 366 ff., Figs. 3a-e; PL 26, Figs. 1-3. Figs. 4-5. 1903 Cymatosaurus gracilis, Freeh, p. 15. 1899 Cymatosaurus (Nothosaurus) latifrons, 1903 Cymatosaurus latifrons, Freeh, Figs. Schrammen, p. 388/ 2a-c.

FIELDIANA: GEOLOGY Fig. 2. The skull of the holotype of Cymatosaurus fridericianus v. Fritsch, 1894 (Ha, uncatalogued), from the = lower Muschelkalk of Halle/Saale. A, Dorsal view; B, ventral view. Scale bar 20 mm. Abbreviations: ec, ectopterygoid; f, frontal; m, maxilla; n, nasal; p, parietal; pi, palatine; pm, premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; pt, pterygoid; sq, squamosal; v, vomer.

1903 Cymatosaurus silesiacus, Freeh, p. 15. 1928 Cymatosaurus silesiacus, Schmidt, p. 1914 Cymatosaurus gracilis, Schroder, p. 78 394, Figs. 1106a-b.

ff., Fig. 16. 1934 Cymatosaurus gracilis, Kuhn. p. 43. 1914 Cymatosaurus latifrons, Schroder, p. 86 1934 "Nothosaurus" latifrons, Kuhn, ff p. 34. 1914 Cymatosaurus silesiacus, Schroder, p. 1934 Cymatosaurus silesiacus, Kuhn, p. 42. 82 ff 1935 Cymatosaurus silesiacus, Edinger, Fig. 1924 Nothosaurus latifrous, Arthaber, p. 476. 9c. 1924 Eurysaurus latifrous, Arthaber, p. 477. 1944 Cymatosaurus gracilis, E.v. Huene, pp. 1924 Cymatosaurus silesiacus, Arthaber, p. 198/, 208. 473. 1944 Cymatosaurus latifrons, E.v. Huene, pp. 1928 Cymatosaurus (?) gracilis, Schmidt, p. 198/, 207/ 394, Fig. 1107. 1944 Cymatosaurus silesiacus, E.v. Huene, 1928 Cymatosaurus latifrons, Schmidt, p. pp. 198/, 208. 394, Fig. 1108. 1964 Cymatosaurus gracilis, Kuhn, p. 12.

RIEPPEL: REVISION OF CYMATOSAURUS Fig. 4. A snout fragment of Cymatosaurus cf. C. fridericianus (smns 7209) from the lower Muschelkalk of Freudstadt. Scale bar = 20 mm.

Locus Typicus—Lower Muschelkalk (Gogolin beds, mu,), Gogolin, Upper Silesia (Poland); for the neotype: lower Muschelkalk (Gogolin beds, mu,), Krappitz (Krapkowice), Upper Silesia (Po- land). Diagnosis—A species of Cymatosaurus of in-

1 termediate size (tip of snout to back end of partial skull table up to 150 mm); single maxillary tooth preceding paired maxillary fangs; pre- and postfrontal may or may not be in contact at dorsal margin of orbit; frontal may closely approach but does not enter anteromedial margin of upper temporal fossa. Distribution—Lower Muschelkalk (lower An- isian, lower Middle Triassic), central Europe. Referred Specimens—bgr S 44/3 (complete but Fig. 3. A fragmentary skull (bgr S 44/3) from the poorly preserved skull, lower Muschelkalk lower Muschelkalk of Gogolin, Upper Silesia, referred [lower Gogolin beds], Sacrau near Gogolin; smns to Cymatosaurus fridericianus v. Fritsch, 1 894. A, Dor- 10977 Buntsand- = (incomplete skull, uppermost sal view; B, occipital view. Scale bar 20 mm. Abbre- stein [Coelestinschichten, uppermost Rot], Jenzig viations: bo, basioccipital; f, frontal; p, parietal; po, post- near Jena 1994b, 11]). orbital; pof, postfrontal; pt, pterygoid; q, quadrate; sq, [Rieppel, Fig. squamosal. Comments—The original (type) material of Cy- matosaurus latifrons (Gurich, 1 884) can no longer be located today. Freeh (1903) figured the speci- 1964 Nothosaurus {7 Cymatosaurus) latifrons, men after additional preparation, but unfortunate- Kuhn, p. 8. ly published the photograph with retouched suture

1964 Cymatosaurus silesiacus, Kuhn, p. 13. lines. Following Freeh's (1903) interpretation, the nasals remain excluded from the posterior margin Neotype—The skull described by Gurich of the external nares, as in Cymatosaurus frider- (1884) and figured by Freeh (1903, Figs. 2a-c) can no be in 1 longer located public repositories. A small and as yet undescribed species of Cymato- Schrammen's (1899) second specimen of Cyma- saurus from the lower Muschelkalk of Thuringia differs from other of its genus by relatively long and tosaurus "gracilis" (smns 10109; Figs. 5-6) is species narrow upper temporal fenestrae and the presence of a the only complete skull of Cymatosaurus latifrons sagittal crest. The species will be described elsewhere in available and is here the neo- today designated collaboration with R. Werneburg, Natural History Mu- type. seum Schleusingen.

FIELDIANA: GEOLOGY Table 1. Skull proportions in Cymatosaurus, Ger- manosaurus, and Nothosaurus. Fig. 5. The skull of the neotype of Cymatosaurus latifrons (Giirich, 1884) (smns 10109), from the lower Muschel- = kalk of Krapkowice, Upper Silesia. A, Dorsal view; B, ventral view. Scale bar 20 mm.

this to be a derived character, diagnostic of a sep- of the orbit, the restriction of the maxillary tooth arate species for which Cymatosaurus latifrons row to a level in front of the anterior margin of Giirich, 1884, takes priority. the upper temporal fossa, and the location of the Morphological Description—The description pineal foramen at the center of the parietal skull of the skull will primarily be based on the neotype table. (smns 10109; Figs. 5-6); reference will be made Total length of the skull of smns 10109 (as pre- to the referred specimen smns 10977 (Rieppel, served) is 104 mm; the distance from the tip of 1994b, Fig. 11) where necessary. In contrast to the snout to the posterior end of the parietal skull smns 10977, the neotype is strongly compressed table measures 98 mm. The rostrum is narrow and dorsoventrally, and its small size indicates a ju- elongated. Its relative length does not differ in C venile status. Reference of both skulls to Cyma- latifrons and in C fridericianus: dividing the dis- tosaurus is corroborated by the paired frontals and tance from the tip of the snout to the anterior mar- the incompletely fused parietals, the contact of gin of the orbit by the distance from the tip of the prefrontal and postfrontal along the dorsal margin snout to the anterior margin of the external naris

10 FIELDIANA: GEOLOGY Fig. 6. The skull of the neotype of Cymatosaurus latifrons (Gurich, 1884) (smns 10109) from the lower Muschel- = kalk of Krapkowice, Upper Silesia. A, Dorsal view; B, ventral view. Scale bar 20 mm. Abbreviations: ec, ectopterygoid; f, frontal; m, maxilla; n, nasal; p, parietal; pi, palatine; pm, premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; pt, pterygoid; q, quadrate; v, vomer.

yields values of 1.87 for the holotype of C. fri- venile specimen. Schrammen's (1899) first speci- dericianus and 2.0 for the neotype of C latifrons. men of C "gracilis" would show a correspond- Dividing the distance from the tip of the snout to ing ratio of 1.5, whereas in the larger C. "silesi- the anterior margin of the upper temporal fossa acus" (Schrammen, 1899; first specimen) the ra- by the distance from the tip of the snout to the tio is 2.9. The original holotype of Cymatosaurus anterior margin of the external naris yields values latifrons Gurich, 1884, again a relatively small of 2.68 for the holotype of C. fridericianus and skull (tip of snout to posterior end of parietal skull 3.0 and for the neotype of C. latifrons. The some- table approximately 100 mm), shows a corre- what larger value in the latter species may be due sponding value of 1.7. The relatively large size of to relatively larger orbits in this juvenile speci- the orbits is also borne out by the quotient which men. Dividing the distance from the tip of the results from division of the distance between the snout to the anterior margin of the external naris external naris and the orbit by the distance be- by the maximum width of the premaxillary ros- tween the orbit and the upper temporal arch. The trum yields values of 1.4 for the holotype of C. quotient is 2.8 for the holotype of C fridericianus fridericianus and 1.2 for the neotype of C. lati- and 2.3 for the neotype of C latifrons. Schram- frons. Dividing the dorsal bridge between the or- men's (1899) first specimen of C. "gracilis" bits by the dorsal bridge between the external nar- would show a corresponding ratio of 1 .7, whereas " es yields values of 2.4 for the holotype of C fri- in the larger C. silesiacus" (Schrammen, 1899; dericianus and 1 .6 for the neotype of C. latifrons. first specimen) the ratio is 2.6. This indicates a relatively narrower dorsal bridge The paired premaxillae meet the maxillae at the between the orbits in the latter species, again a anterolateral corner of the external naris, and they consequence of relatively large orbits in this ju- form a broad and interdigitating contact with the

RIEPPEL: REVISION OF CYMATOSAURUS 11 Fig. 7. The holotype of Cymatosaurus erythreus E.v. Huene, 1894 (bgr 612), from the upper Buntsandstein of Riidersdorf near Berlin, A, Dorsal view; B, ventral view. Scale bar = 20 mm.

paired frontals between the nasals, at a level be- process of the frontal approaches the upper tem- tween the external nares and the orbits. The ex- poral fossa less closely in smns 10977). The post- ternal nares are relatively broad and rounded. The frontal is a relatively broad element that defines left nasal cannot be identified in smns 10109 be- the posterodorsal margin of the orbit as well as cause of excessive original preparation of the in- the anteromedial margin of the upper temporal ternal naris, but the reduced right nasal can be fossa. It meets the postorbital in a deeply inter- seen to be excluded from the posterior margin of digitating suture within the postorbital arch. The the external naris, unlike in Schrammen's (1899, lateral (ventral) part of the postfrontal, and its re- PI. 23, Fig. 2) "first specimen" of C. "gracilis"; lation to the jugal and to the upper temporal arch, in that respect, the latter specimen resembles the are not preserved on either side of the skull, smns holotype of C. silesiacus (Schrammen, 1899, PI. 10977 shows the postorbital to meet the squa- 21). Laterally, the nasal remains separated from mosal in a broadly overlapping suture within the the prefrontal by a broad contact between an an- upper temporal arch. The jugal is a curved ele- terolateral process of the frontal and the maxilla. ment without posterior process that defines the Exclusion of the reduced nasals from the external posteroventral corner of the orbit. naris and separation of the nasal from the pre- The interdigitating frontoparietal suture is lo- frontal by a frontal-maxillary contact are also ob- cated between the anterior parts of the upper tem- served in smns 10977. The prefrontal appears as poral fenestrae. To judge from the shape of the a rather slender element in dorsal view, lining the lateral margin of the parietal, the anterior, medial, anterodorsal margin of the orbit and establishing and posterior margins of the relatively broad up- a contact with the postfrontal that excludes the per temporal fossa were more or less evenly frontal from the orbit (unlike in Schrammen's curved. The pineal foramen is located at the cen- [1899, PI. 23, Fig. 2] first specimen, but as in the ter of the parietal skull table. In front of the pineal holotype of C. silesiacus, and as in smns 10977). foramen, a distinct suture separates the paired pa- The frontal forms a distinct posterolateral process rietals (the parietals are fully fused in smns that remains narrowly excluded from the antero- 10977). Behind the pineal foramen, the parietal is medial margin of the upper temporal fossa (as in distinctly constricted but not developed into a sag- Schrammen's "first specimen," the posterolateral ittal crest. No trace of a medial longitudinal suture

12 FIELDIANA: GEOLOGY Fig. 8. The holotype of Cymatosaurus erythreus E.v. Huene, 1894 (bgr 612), from the upper Buntsandstein of = Riidersdorf near Berlin. A, Dorsal view; B, ventral view. Scale bar 20 mm. Abbreviations; f, frontal; m, maxilla; pi, palatine; pm, premaxilla; prf, prefrontal; v, vomer.

can be identified in the constricted posterior part crushing of the skull. In the three-dimensionally of the parietal skull table. preserved specimen smns 10977, the suspenso- Preservation of the occiput is incomplete in rium is more vertically oriented, and the mandib- smns 10109. As preserved, the skull contours in- ular articulation is positioned below the posterior dicate a deeply concave occiput and a limited oc- end of the upper temporal fossa, as is typical for cipital exposure of the parietal. Poor preservation the genus in general. makes it impossible to trace the suture between The ventral view of the skull shows five pre- the parietal and squamosal. All posterior braincase maxillary tooth positions and a single maxillary elements (supraoccipital, exoccipitals, opisthotic, tooth preceding the paired maxillary fangs (three and basioccipital [occipital condyle]) are missing. maxillary teeth precede the paired maxillary fangs This indicates a loose connection between the in C. fridericianus). Behind the maxillary fangs, braincase and the dermatocranium, which might 10-11 tooth positions can be counted, bringing be attributed to the juvenile status of the speci- the total number of maxillary teeth up to 13-14. men. However, all posterior braincase elements The same tooth count is obtained in smns 10977. are also missing in the larger specimen smns Posteriorly, the maxillary tooth row does not ex- 10977, as indeed in every other specimen of Cy- tend beyond the level of the posterior margin of matosaurus available today. It is for this reason the orbit. The internal nares are relatively shorter that a tight association of the braincase with the than in C. fridericianus (division of the longitu- dermatocranium in the formation of a closed, dinal diameter by the transverse diameter yields a plate-like occiput (as seen in Simosaurus and No- ratio of 1.6 for smns 10109), but as in the latter thosaurus) is believed to be absent in Cymatosau- species, and unlike Nothosaurus, the internal naris rus (characters 3 1 and 32 of the cladistic analysis, is positioned somewhat further back relative to the discussed below). The suspensorium appears to be external naris (Table 1). distinctly flaring in smns 10109, carrying the A narrow fontanelle ("foramen incisivum") mandibular articulation to a position well lateral separates the premaxillae in ventral view. The of the posterior corner of the upper temporal fos- paired vomers meet the maxillae at the anterior sa. This is, however, an artifact of dorsoventral margin of the internal nares, thus excluding the

RIEPPEL: REVISION OF CYMATOSAURUS 13 premaxillae from the latter. Posteriorly, the vo- Referred Specimen—An isolated parietal (Fig. mers meet the pterygoids in an interdigitating su- 9; bgr 613; original of E.v. Huene, 1994, p. 200, ture. The posterior margin of the internal naris is Fig. 2 in her paper) from the same locality and defined by the palatine. The exact contours of the horizon as the holotype. be but the ectopterygoid cannot identified, pres- Comments—This species was described by E.v. ence of a well-developed (ecto-)pterygoid flange Huene (1944) on the basis of an incomplete skull is distinct on the left side of the skull. Posteriorly, that represents the earliest occurrence of the genus the shows lateral and pterygoid well-developed Cymatosaurus in the Germanic Triassic. The frag- medial ventral flanges for the origin of the pter- ment (Figs. 7, 8) comprises most of the preorbital ygoideus musculature. skull and has a total length of 58.4 mm (as pre-

served). The distance from the tip of the snout to the anterior margin of the external naris is 29 (28) Cymatosaurus multidentatus mm, to the anterior margin of the orbit 54 mm. (F.v. Huene, 1958) The longitudinal diameter of the external naris is 12.2 mm, and the bridge between the external nar- 1958 Anarosaurus multidentatus, F.v. Huene, is and the orbit is 13.5 mm wide. The width of pp. 382-384. the snout at the level of the anterior margins of 1995c Cymatosaurus multidentatus, Rieppel, the external nares is 23.5 mm; the total width of p. 288. the premaxillary rostrum (at the fourth tooth po- is 25.7 mm. The diameter of Holotype—Incomplete lower jaw (Mbg 4791). sition) longitudinal the internal naris is 9.5 mm. Locus Typicus—Lowermost lower Anisian (6 The rostrum is of similar relative as in m above the base), Krabachmasse, Trittwangkopf, length the NNW Stuttgarter Hiitte, Lechtaler Alpen (Arl- genotypical species, Cymatosaurus fridericianus Fritsch as in other berg), Austria. (1894), or, indeed, species of the distance from the Diagnosis—A very incompletely known spe- Cymatosaurus. Dividing cies of Cymatosaurus of small size, characterized tip of the snout to the anterior margin of the orbit by a distinctly heterodont dentition. by the distance from the tip of the snout to the Referred Specimens—Mbg 4792 (isolated anterior margin of the external naris results in a tooth [Huene, 1958, Fig. 2], fragmentary verte- ratio of 1.89 for Cymatosaurus erythreus, com- bra); Mbg 4793 (isolated tooth [Huene, 1958, Fig. pared to 1.87 for Cymatosaurus fridericianus. 3]. ?metatarsal [Huene, 1958, Fig. 6]); Mbg 4794 The rostrum is formed by the premaxillae. (vertebral centrum [Huene, 1958, Fig. 24); Mbg Broad posterior (nasal) processes of the premax- 4795 vertebral centrum (fragmentary [Huene, illae separate the external nares from one another 1958, Fig. 5]).— and meet the frontals in a deeply interdigitating Comments This species was redescribed by suture at a level between the external nares and Rieppel (1995c). the orbits. The preserved anterior ends of the frontals show these elements to be paired, as is char- acteristic of the genus Cymatosaurus. Cymatosaurus sp. ("C. E.v. Huene, erythreus" Damage to the bone surface has rendered the 1944) accurate delineation of the nasals impossible. However, the nasal extends anteriorly along the Synonymy for this material: medial margin of the external naris in those stem- group sauropterygians that do not show reduction 1994 Cymatosaurus erythreus, E.v. Huene, of these elements. No participation of the nasals pp. \93ff., Figs. la-c. can be identified in the medial margins of the ex- 1964 Cymatosaurus erythreus, Kuhn, p. 13. ternal nares in Cymatosaurus erythreus, indicating 1995c Cymatosaurus erythreus, Rieppel, p. reduction of these bones. Likewise, the left exter- 295, Figs. 7, 8C. nal naris indicates the exclusion of the nasal from

its another feature for Holotype—Snout fragment (brg 612; Figs. 7, posterior margin, diagnostic 8). (some) Cymatosaurus. Typicus— The is at the anterodorsal Locus Upper Buntsandstein (so2 , prefrontal exposed Rot), Rudersdorf near Berlin. margin of the left orbit. A lacrimal is lacking

14 FIELDIANA: GEOLOGY shaped than in Nothosaurus, and most of them are confluent with the alveoli of the functional teeth. The skull fragment described by E.v. Huene (1944) as Cymatosaurus erythreus is unquestion- ably a representative of that genus, as is indicated by a number of characters, most notably the exclusion of the (reduced) nasals from the external naris and the paired condition of the frontals. The specimen is too incomplete, however, to be diagnostic for a separate species. Within the genus, it shares with Cymatosaurus fridericianus three small maxillary teeth preceding the maxillary fangs, but this is a plesiomorphic trait in comparison to the outgroup (Nothosauridae). Therefore, Cymatosaurus erythreus E.v. Huene, 1944, is here considered a nomen dubium.

Nothosauridae Baur, 1889

Definition—A monophyletic taxon including the genera Germanosaurus Nopcsa, 1928a, No- thosaurus Miinster, 1834, and Lariosaurus Cu- rioni, 1847 (Ceresiosaurus Peyer, 1931, and Sit- vestrosaurus Kuhn-Schnyder, 1990, are here con- Fig. 9. An isolated parietal referred to Cymatosaurus sidered junior synonyms of Lariosaurus: Rieppel, erythreus E.v. Huene, 1894 (bgr 613), from the upper 1993, and in prep.). Buntsandstein of Rudersdorf near Berlin. Scale bar - 20 Diagnosis—Small to large eosauropterygians with a strongly depressed skull; snout constricted; dorsal exposure of prefrontal reduced; jugal (usu- ally) excluded from posterior margin of orbit; pre- E.v. the lacri- (contra Huene, 1944, p. 195), and maxillary and anterior dentary fangs present. mal duct is enclosed entirely within the maxilla, Distribution—Lower to Upper Triassic, Eu- as is also observed in Nothosaurus. rope and Israel. The ventral aspect reveals relatively short and Comments—Within the Nothosauridae, Ger- broad internal nares, a feature shared with other manosaurus is the sister-taxon of a monophyletic species of Cymatosaurus but a contrast to the rel- clade (Nothosaurinae: Nopcsa, 1923, 1928a,b) in- nares ob- atively narrow and elongated internal cluding the genera Nothosaurus and Lariosaurus. served in Nothosaurus (except N. marchicus). The Diagnostic characters of the Nothosaurinae are of the internal naris is defined posterior margin by frontals fused, parietal skull table strongly con- the palatine and its medial margin by the (paired) stricted, maxillary fangs present, occipital crest vomer. Vomer and maxilla meet in a broad contact present, supraoccipital horizontally oriented and at the anterior margin of the internal naris, thus fused with parietal, occipital crest present, man- excluding the premaxilla from the latter. Between dibular symphysis strongly elongated and fortified the premaxillae and in front of the vomers, a fon- (not known for Germanosaurus), unconstricted tanelle persists, a feature shared with Cymatosau- vertebral centra (not known for Germanosaurus), rus, Pistosaurus (Fritsch, 1 894), and Nothosaurus and no distal expansion of sacral ribs (not known (Rieppel & Wild, 1996). The right premaxilla car- for Germanosaurus). ries five tooth positions; the size of the alveoli indicates that the third and fourth teeth were the largest. Three small maxillary teeth precede the Germanosaurus Nopcsa, 1928 paired maxillary fangs. The replacement pits, located posteromedial to the functional tooth posi- 1903 Eurysaurus, Freeh, p. 15. tions, are more distinctly elongated and drop- 1924 Eurysaurus, Arthaber, p. 476.

RIEPPEL: REVISION OF CYMATOSAURUS 15 1928a Germanosaurus, Nopcsa, pp. 21, 44. Holotype—The skull described by Giirich 1928b Germanosaurus, Nopcsa, p. 173. (1891) and figured by Koken (1893, Fig. 10) and 1944 Cymatosaurus {Germanosaurus) (par- Freeh (1903, Fig. 1) can no longer be located in tim), E.v. Huene, p. 208. public repositories. Locus Typicus—Lower Muschelkalk (Gogolin Type Species—Germanosaurus latissimus Giir- beds, mu,), Sacrau near Gogolin, Upper Silesia ich, 1891, from the lower Muschelkalk, lower (Poland). Middle Triassic, Sacrau, Upper Silesia. Comments—The taxonomic status of "Cyma- Definition—A monophyletic taxon including tosaurus" latissimus (Giirich, 1891) can no longer the species schafferi. be critically evaluated because the original (type) Diagnosis—A large eosauropterygian with a material has been lost. The situation is aggravated constricted snout and a relatively short and broad by the fact that Giirich (1891, figure on p. 968) rostrum. Posterolateral process of paired frontals included only a very schematic figure in his orig- closely approaches upper temporal fossa; parietals inal description of the species. The skull was pre- paired. served in two parts and estimated by Giirich Comments—Of the two species described in (1891) to be approximately 270 mm long. With the genus, Germanosaurus latissimus (Giirich, this size, the specimen is distinctly larger than any 1891) and Germanosaurus schafferi (Arthaber, specimen of Cymatosaurus but closely approach- 1924), only the latter is represented by original es the holotype of Germanosaurus schafferi (Ar- material available today. thaber, 1924). Koken (1893, Fig. 11) indicates four maxillary teeth preceding the paired maxillary fangs, a character that separates "Cymato- Germanosaurus sp. ("G. latissimus" [Giirich, saurus'''' latissimus from the known species of Cy- 1891]) matosaurus. In other respects, however, "Cyma- tosaurus" latissimus resembles Germanosaurus Synonymy for this material: schafferi, with which it shares the reduced nasal bones that enter the posterior margin of the external the the restricted dor- 1891 Nothosaurus latissimus, Giirich, p. 968, nares, paired frontals, Figs. a-b. sal exposure of the prefrontals (broadly exposed in the 1893 Nothosaurus latissimus, Koken, p. 368 Cymatosaurus), which do not contact post- ff, Figs. 6-11. frontal along the dorsal margin of the orbit, and 1899 Cymatosaurus (Nothosaurus) latissi- a jugal bone that extends alongside the postorbital into the does not mus, Schrammen, p. 388/ upper temporal arch (the jugal 1903 Nothosaurus (Eurysaurus) latissimus, extend into the upper temporal arch in Cymato- Freeh, p. 15, Fig. 1. saurus as shown by the specimen smns 10977). 1914 Nothosaurus (Eurysaurus) latissimus, Whether the jugal does, indeed, broadly enter the in Schroder, p. 78. ventral margin of the orbit "Cymatosaurus" latissimus or whether is 1914 Eurysaurus latissimus, Schroder, p. 82 (Koken, 1893, Fig. 10), ff, Fig. 22. was excluded therefrom, can no longer be as- sessed. In of its size the characters 1924 Eurysaurus latissimus, Arthaber, p. 476/ view large and 1928 Eurysaurus latissimus, Schmidt, p. 395, outlined above, "Cymatosaurus" latissimus is Fig. 1110. here referred to the genus Germanosaurus. Ger-

1934 Germanosaurus latissimus, Kuhn, p. 41. manosaurus latissimus (Giirich, 1891) is here

1944 Cymatosaurus latissimus, E.v. Huene, p. treated as a nomen dubium rather than as a senior 199. synonym of Germanosaurus schafferi (Arthaber, 1944 Cymatosaurus (germanosaurus) latissi- 1924), because loss of the holotype (and only mus, E.v. Huene, p. 208. known specimen) prevents critical comparison

1964 Cymatosaurus latissimus, Kuhn, p. 13. with the latter species.

Fig. 10. The skull of the holotype of Germanosaurus schafferi (Arthaber, 1924) (nhmw, uncatalogued), from the = lower Muschelkalk of Gogolin, Upper Silesia, in dorsal view. Scale bar 50 mm. (Courtesy of Dr. G. Hock, Natural History Museum, Vienna.)

16 FIELDIANA: GEOLOGY RIEPPEL: REVISION OF CYMATOSAURUS 17 Fig. 12. An isolated parietal referred to Germano- saurus schafferi (Arthaber, 1924) (bgr, uncatalogued), from the lower Muschelkalk of Gogolin, Upper Silesia. = Scale bar 50 mm. Abbreviations: f, frontal; p, parietal.

Diagnosis—Same as for genus, of which this is the only species represented by original material today. Distribution—Lower Muschelkalk (lower An- isian, lower Middle Triassic), central Europe. Referred Specimens—Isolated parietal (Fig. 12) from the lower Muschelkalk (Gogolin beds, mu,), Gogolin, Upper Silesia (bgr, uncatalogued; Rieppel, 1994a, Fig. 40). Morphological Description—Germanosaurus schafferi is known from a single skull prepared in dorsal view (Fig. 11). The distance from the tip of the snout to the back end of the parietal skull table measures 244 mm, which is distinctly longer than in the skulls referred to the genus Cymato- saurus but approaches the size of "Cymatosau- rus" latissimus (see above). The rostrum is relatively broader than in skulls referred to the genus Fig. 11. The skull of the holotype of Germanosau- with the that the ros- rus schqfferi (Arthaber, 1924) (nhmw, uncatalogued), Cymatosaurus, consequence from the lower Muschelkalk of Gogolin, Upper Silesia, tral constriction is more pronounced in Germa- in dorsal view. Scale bar = 50 mm. Abbreviations: f, nosaurus. Dividing the distance from the tip of frontal; ju, jugal; m, maxilla; n, nasal; p, parietal; pm, the snout to the anterior margin of the external premaxilla; po, postorbital; pof, postfrontal; prf, prefrontal; sq, squamosal.

Table 2. Data matrix for the cladistic analysis of all Cymatosaurus species ever described. Character defini- Germanosaurus schafferi (Arthaber, 1924) tions are given in the Appendix.

1924 Eurysaurus schafferi, Arthaber, p. 1476 ff, Figs, lla-b.

1928 Eurysaurus schafferi, Schmidt, p. 395, Fig. 1111.

1934 Germanosaurus schafferi, Kuhn, p. 42. 1944 Cymatosaurus {Germanosaurus) schafferi, E.v. Huene, p. 208/ 1964 Cymatosaurus schafferi, Kuhn, p. 12.

Holotype—Skull (nhmw, uncatalogued; Figs. 10, 11). Locus Typicus—Lower Muschelkalk (Gogolin beds, rau,), Gogolin, Upper Silesia (Poland). Captorhinidae Ancestor Captorhinidae Testudines Araeoscelidia

Araeoscelidia Claudiosaurus Younginiformes Younginiformes Archosauriformes Rhynchosauria Claudiosaurus Trilophosaurus Placodus Prolacertiformes Choristodera Corosaurus Kuehneosauridae Rhynchocephalia Cymatosaurus Squamata Pistosaurus Testudines Placodus Dactylosaurus Corosaurus Cymatosaurus Serpiano-Neustico Pistosaurus Simosaurus Dactylosaurus Serpiano-Neustico Germanosaurus Simosaurus

Nothosaurus Germanosaurus Nothosaurus Lariosaurus Lariosaurus

Fig. 13. Strict consensus tree (2 MPTs, TL = 283, Fig. 14. Strict consensus tree (2 MPTs, TL = 452, = = = = CI 0.675, RI 0.702) for a monophyletic Sauropter- CI 0.648, RI 0.71 1) for 23 ingroup taxa rooted on ygia rooted on Captorhinidae, Araeoscelidia, Youngini- an all-0 ancestor. For further discussion see text. formes, Claudiosaurus, and Testudines. For further discussion see text.

feri, which falls into the upper range of variability of Cymatosaurus (1.3-2.0) and in between the naris by the maximum width of the premaxillary range of variability between relatively plesiom- rostrum yields values of 0.94 (left side) and 1 .03 orphic {Nothosaurus marchicus) and apomorphic (right side) for the holotype of Germanosaurus {Nothosaurus mirabilis) representatives of the ge- schafferi, 1 .4 for the holotype of C. fridericianus, nus Nothosaurus. Indeed, the skull of Germano- and 1.2 for the neotype of C. latifrons. Dividing saurus schafferi shows an intriguing combination the distance from the tip of the snout to the an- of characters otherwise typical of Cymatosaurus terior margin of the orbit by the distance from the or Nothosaurus. tip of the snout to the anterior margin of the ex- The adult stage of Germanosaurus is indicated ternal naris yields a value of approximately 1.8 by the fusion of the premaxillae in the anterior for the holotype of Germanosaurus schafferi, 1 .9 part of the rostrum. The premaxilla meets the for the holotype of C. fridericianus, and 2.0 for maxilla at the anterolateral edge of the external the neotype of C. latifrons. Dividing the distance naris, in a suture that trends anterolaterally. In Cy- from the tip of the snout to the anterior margin of matosaurus and Nothosaurus, the maxilla shows the upper temporal fossa by the distance from the a distinct lateral bulging in the area between the tip of the snout to the anterior margin of the ex- external nares and orbits, which accommodates ternal naris yields a value of approximately 2.86 the roots of the paired maxillary fangs. In Ger- for the holotype of Germanosaurus schafferi, 2.7 manosaurus, the lateral contours of the skull show for the holotype of C. fridericianus, and 3.0 for a lateral bulging at the level of the external nares, the neotype of C latifrons. Dividing the longitu- but whereas the four, perhaps five, premaxillary dinal diameter of the upper temporal fossa by the teeth (fangs) are distinctly larger than the maxil- longitudinal diameter of the orbit yields a value lary teeth, no maxillary fangs are observed either of 1 .9 for the holotype of Germanosaurus schaf- at the level of the lateral bulging of the maxilla

RIEPPEL: REVISION OF CYMATOSAURUS 19 Sauropterygia

2 £ 3 w•2 Pachypleurosauroidea Nothosauridae

Fig. 15. Clade rank determination in Sauropterygia. Pachypleurosaur interrelationships are based on an in- dependent study (Rieppel & Lin Kebang, 1995). For further discussion see text and Norell and Novacek (1992).

or behind it at the level between the external naris and the orbit. Because a small maxillary tooth is preserved (in the left maxilla) in the position where other Eosauropterygia carry paired maxillary fangs (between the external naris and the orbit), it is concluded that such fangs are absent in Germanosaurus (alternatively, one might assume that both fangs have been shed, and a replacement tooth is observed in situ, in which case the maxillary fangs would be a synapomorphy of the No- thosauridae rather than of the Nothosaurinae). Germanosaurus differs from Nothosaurus, but resembles some specimens of Cymatosaurus, in that the nasals are reduced but still reach the posterior margin of the external nares. As in Cyma- tosaurus, but unlike Nothosaurus, there is no slender anterior process of the nasal lining the medial margin of the external naris that is defined by the premaxilla. Posteriorly, the nasals taper to a blunt tip that is embraced between an anterolateral and an anteromedial process of the frontal. At the level of the anterior margin of the orbit, the anteromedial process of the frontal meets the posterior (nasal) process of the premaxilla, which separates the nasals from one another. The anterolateral pro- Table 3. Data matrix for the cladistic analysis of sauropterygian interrelationships. Character definitions are given in the Appendix. See Rieppel (1994a) and text for further discussion. Table 3. Continued. Table 3. Continued. Table 3. Continued. Table 3. Continued. Table 3. Continued. Table 3. Continued. Table 3. Continued. Table 3. Continued. Table 3. Continued. Table 3. Continued. Table 3. Continued. ,, "Silvestrosaurus (Anisian-Ladinian boundary), chelkalk. Schontaler Symposium 1991. Goldschneck Verlag, Korb. "Ceresiosaurus" (early Ladinian), and Lariosau- Curioni, G. 1847. Cenni un nuovo saurio rus (middle to late Ladinian) at the alpha taxo- sopra fossile dei monti di Perledo sul Lario e sul terreno che lo nomic level must precede the analysis of phylo- racchiude. Giornale del' J.R. Instituto Lombardo di its taxa. genetic interrelationships among terminal Scienze, Lettre ed Arti, 16: 159-170.

Dames, W. 1890. Anarosaurus pumilio nov. gen. nov. sp. Zeitschrift der Deutschen Geologischen Gesell- schaft, 42: 74-85. Acknowledgments Edinger, T 1935. Pistosaurus. Neues Jahrbuch fur Mi- neralogie, Geologie und Palaontologie, Abhandlun- gen, B, 74: 321-359. A number of colleagues granted generous ac- Evans, S. E. 1988. The early history and relationships cess to the collection in their care, making the of the Diapsida. In Benton, M. J., ed.. The Phylogeny of this completion study possible. These are H. U. and Classification of the Tetrapods, 1: 221-260. Clar- Schliiter, Bundesanstalt fur Geowissenschaften endon Press, Oxford. und Rohstoffe, Berlin; H. Haubold, Institut fur Frech, E 1903. Lethaea Geognostica. Handbuch der II. Teil. Das Mesozoicum. 1. Geowissenschaften, Martin-Luther-Universitat, Erdgeschichte. Band. Trias. E. Schweizerbart'sche Verlagsbuchhandlung (E. Halle/Saale; G. Kaufmann, Fachbereich Geowis- Nagele), Stuttgart. senschaften, Philipps Universitat, Marburg/Lahn; Fritsch, K.v. 1894. Beitrag zur Kenntnis der Saurier R. Werneburg, Naturhistorisches Museum Schloss des Halle'schen unteren Muschelkalkes. Abhandlun- Bertholdsburg, Schleusingen; G. Hock, Naturhis- gen der Naturforschenden Gesellschaft zu Halle, 20: torisches Museum, Wien; and R. Wild, Staatliches 273-302. J. A. G. and Rowe. 1988. Museum fur Naturkunde, Stuttgart. Dr. G. Hock Gauthier, A., Kluge, T Am- niote phylogeny and the of fossils. Cla- kindly provided the photograph of Germanosau- importance distics, 4: 104-209. rus shown in Figure 10. H.-D. Sues and H. Hag- Gurich, G. J. E. 1884. Uber einige Saurier des Ober- dorn read an earlier version of this critically paper, schlesischen Muschelkalkes. Zeitschrift der Deutschen offering much helpful advice and criticism. This Geologischen Gesellschaft, 36: 125-144. was study supported by NSF grants DEB- . 1891. Uber einen neuen Nothosaurus von Go- 9220540 and DEB-94 19675. golin, Oberschlesien. Zeitschrift der Deutschen Geo- logischen Gesellschaft, 43: 967-970. Haas, G. 1963. Micronothosaurus stensidi, ein neuer Nothosauride aus dem Oberen Muschelkalk des Wadi Ramon, Israel. Palaontologische Zeitschrift, 37: 161- Literature Cited 178.

Alafont, L. S., and J. L. Sanz. 1996. Un nuevo Sau- Hagdorn, H. 1985. Immigration of crinoids into the ropterigio (Reptilia) en el Triasico de la Sierra de Pra- German Muschelkalk basin, pp. 237-254. In Bayer, des (Tarragona). Cuadernos de Geologia Iberica, 20: U., and A. Seilacher, eds., Sedimentary and Evolu- 313-329. tionary Cycles. Springer, Heidelberg.

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-. 1928b. The genera of reptiles. Palaeobiologica, Schmidt, K. P. 1927. New reptilian generic names. Co- 1: 163-U peia, 1927: 58-59. Norell, M., and M. Novacek. 1992. Congruence be- Schmidt, M. 1928. Die Lebewelt unserer Trias. F. Rau, tween superpositional and phylogenetic pattern: Com- Ohringen. cladistic with fossil records. paring patterns Cladistics, Schrammen, A. 1899. 3. Beitrag zur Kenntnis der No- 8: 319-337. thosauriden des unteren Muschelkalkes in Oberschle- Gesell- Owen, R. 1860. Palaeontology; or, a Systematic Sum- sien. Zeitschrift der Deutschen Geologischen mary of Extinct Animals and Their Geologic Re- schaft, 51: 388-408. mains. Adam and Charles Black, Edinburgh. Schroder, H. 1914. Wirbeltiere der Rudersdorfer Trias. der Preussischen Geolo- Peyer, B. 1931. Die Triasfauna der Tessiner Kalkalpen. Abhandlungen Koniglich Neue 65: 1-98. IV. Ceresiosaurus calcagnii nov. gen. nov. spec. Ab- gischen Landesanstalt, Folge, handlungen der Schweizerischen Palaontologischen Schultze, H.-R 1970. Uber Nothosaurus. Neubeschrei- Gesellschaft, 51: 1-68. bung eines Schadels aus dem Keuper. Senckenbergi- ana 51: 211-237. Rieber, H. 1973. Ergebnisse palaontologisch-stratigra- Lethaea, phischer Untersuchungen in der Grenzbitumenzone Storrs, G. W 1991. Anatomy and relationships of Co- (Mittlere Trias) des Monte San Giorgio (Kanton Tes- rosaurus alcovensis (Diapsida: Sauropterygia) and the sin, Schweiz). Eclogae Geologicae Helvetiae, 66: Triassic Alcova Limestone of Wyoming. Bulletin of 667-685. the Peabody Museum of Natural History, 44: 1-151.

Rieppel, O. 1989. A new pachypleurosaur (Reptilia: . 1993a. The quality of the Triassic sauroptery- Sauropterygia) from the Middle Triassic of Monte San gian fossil record. Revue de Paleobiologie, Volume Giorgio, Switzerland. Philosophical Transactions of Special, 7: 217-228. the of London, B, 323: 1-73. Royal Society 1993b. The systematic position of Silvestro-

. 1993. Middle Triassic reptiles from Monte San saurus and a classification of Triassic sauropterygians Giorgio: Recent results and future potential of analy- (Neodiapsida). Palaontologische Zeitschrift, 67: 177- sis, pp. 131-144. In Mazin, J.-M., and G. Pinna, eds., 191.

34 FIELDIANA: GEOLOGY Sues, H.-D. 1987. Postcranial skeleton of Pistosaurus 4. temporal region of skull relatively high (0) and interrelationships of the Sauropterygia (Diapsida). or strongly depressed (1). Zoological Journal of the Linnean Society, 90: 109- 131. 5. Nasals shorter (0) or longer (1) than fron- Swofford, D. L. 1990. PAUP—Phylogenetic Analysis tal(s). Using Parsimony, Version 3.0. Illinois Natural History 6. Nasals not reduced (0), somewhat reduced Survey, Champaign, 111. (1), or strongly reduced or absent (2). Swofford, D. L., and D. P. Begle. 1993. PAUP—Phy- logenetic Analysis Using Parsimony, Version 3.1 Lab- 7. Nasals do (0) or do not (1) enter external oratory of Molecular Systematics, Smithsonian Insti- naris. tution, Washington, D.C. 8. Nasals meet in dorsomedial suture (0) or are Tschanz, K. 1989. Lariosaurus buzzii n. sp. from the from one another nasal of Middle Triassic of Monte San Giorgio (Switzerland), separated by processes with comments on the classification of nothosaurs. Pa- the premaxillae extending back to the frontal laeontographica, A, 208: 153-179. bone(s) (1). Urlichs, M., and R. Mundlos. 1985. Immigration of 9. The lacrimal is present and enters the exter- cephalopods into the Germanic Muschelkalk Basin nal naris (0) or remains excluded from the exter- and its influence on their suture line, pp. 221-236. In nal naris a contact of maxilla 1 Bayer, U., and A. Seilacher, eds., Sedimentary and by and nasal ( ), or Evolutionary Cycles. Springer-Verlag, Heidelberg. the lacrimal is absent (2). Volz, W. 1902. Proneusticosaurus, eine neue Saurop- 10. The prefrontal and postfrontal are separated terygiergattung aus dem untersten Muschelkalk Ob- by the frontal along the dorsal margin of the orbit erschlesiens. Palaeontographica, 49: 121-164. or a contact of and ex- Young, C.-C. 1959. On a new Nothosauria from the (0), prefrontal postfrontal cludes the frontal from the dorsal of the Lower Triassic Beds of Kwangsi. Vertebrata Pal- margin Asiatica, 3: 73-78. orbit (1). 1960. New localities of in sauropterygians 11. Dorsal exposure of prefrontal large (0) or China. Vertebrata PalAsiatica, 3: 73-78. reduced (1). . 1965. On the new nothosaurs from Hupeh and 12. skull: Kweichou, China. Vertebrata PalAsiatica, 9: 337-356. Preorbital and postorbital region of of -. 1978. A nothosaur from Lu-hsi County, Yun- subequal length (0), preorbital region distinctly nan Province, Vertebrata PalAsiatica, 16: 222-224. longer than postorbital region (1), postorbital region distinctly longer (2).

13. Upper temporal fossa absent (0), present and subequal in size or slightly larger than the orbit Appendix (1), present and distinctly larger than orbit (2), or present and distinctly smaller than orbit (3). The characters listed below are based on the 14. Frontal(s) paired (0) or fused (1) in the data used previously in the analysis of the phy- adult. logenetic interrelationships of Simosaurus (Riep- 15. Frontal(s) without (0) or with (1) distinct pel, 1994a). The data matrix relies heavily on the posterolateral processes. work of Gauthier et al. (1988), further augmented 16. Frontal widely separated from the upper by the addition of characters taken from Evans temporal fossa (0), narrowly approaches the upper (1988) and Storrs (1991, 1993b). Additional ref- temporal fossa (1), or enters the anteromedial erences pertaining to the coding of non-saurop- margin of the upper temporal fossa (2). terygian taxa can be found in Rieppel (1994a). fused in their Coding for the postcranium of Cymatosaurus is 17. Parietal(s) paired (0), posterior or fused in adult. based on the assumption that Proneusticosaurus part only (1), fully (2) Volz, 1902, is a subjective junior synonym of Cy- 18. Pineal foramen close to the middle of the matosaurus Fritsch, 1894 (Rieppel & Hagdorn, skull table (0), is displaced posteriorly ( 1 ), is dis- 1996). placed anteriorly (2), or is absent (3).

19. Parietal skull table broad (0), weakly con- 1. Premaxillae small (0) or large (1), forming stricted constricted (at least posteri- most of snout in front of external nares. (1), strongly orly) (2), or forming a sagittal crest (3). 2. Premaxilla without (0) or with (1) postnarial 20. Postparietals present (0) or absent (1). process, excluding maxilla from posterior margin of external naris. 21. Tabular present (0) or absent (1). absent 3. Snout unconstricted (0) or constricted (1). 22. Supratemporals present (0) or (1).

RIEPPEL: REVISION OF CYMATOSAURUS 35 23. The jugal extends anteriorly along the ven- 39. Dorsal wing of epipterygoid broad (0) or tral margin of the orbit (0), is restricted to a po- narrow (1). sition behind the orbit but enters the latter's pos- 40. Lateral conch on quadrate absent (0) or pres- terior or is restricted to a be- margin (1), position ent (1). hind the orbit without reaching the latter's poste- 41. Palate kinetic (0) or akinetic (1). rior margin (2). 42. Basioccipital tubera free (0) or in complex 24. The extends backward no farther than jugal relation to the pterygoid, as they extend ventrally to the middle of the cheek region (0), or nearly to (1) or laterally (2). the posterior end of the skull (1). 43. Suborbital fenestra absent (0) or present (1). 25. The jugal remains excluded from (0) or en- 44. Pterygoid flanges well developed (0) or ters (1) the upper temporal arch. strongly reduced (1). 26. Postfrontal and (0), with dis- large plate-like 45. Premaxillae enter internal naris (0) or are tinct lateral the dorsal tip of process overlapping excluded (1). the postorbital (1), or postfrontal with reduced lat- 46. Ectopterygoid present (0) or absent (1). eral process and hence more of an elongate shape 47. Internal carotid passage enters basicranium (2). (0) or quadrate ramus of pterygoid (1). 27. Lower temporal fossa absent (0), present 48. Retroarticular process of lower jaw absent and closed ventrally (1), or present but open ven- (0) or present (1). trally (2). 49. Distinct coronoid process of lower jaw ab- 28. Squamosal descends to (0) or remains sent (0) or present (1). broadly separated from (1) ventral margin of 50. without or with skull. Surangular (0) (1) strongly projecting lateral ridge defining the insertion area 29. Quadratojugal present (0) or absent (1). for superficial adductor muscle fibers on the lat- 30. with or without an- Quandratojugal (0) (1) eral surface of the lower jaw. terior process. 51. Mandibular symphysis short (0), somewhat 31. Occiput with paroccipital process forming enforced (1), or elongated and "scoop" -like (2). the lower of the fossa and margin posttemporal 52. Splenial bone enters the mandibular sym- (0), extending laterally paroccipital processes physis (0) or remains excluded therefrom (1). trending posteriorly (1), or occiput plate-like with 53. Teeth set in shallow or deep sockets (0) or no distinct paroccipital process and with strongly superficially attached to bone (1). reduced posttemporal fossae (2). 54. Anterior (premaxillary and dentary) teeth 32. Squamosal without (0) or with (1) distinct upright (0) or strongly procumbent (1). notch to receive distal tip of paroccipital process. 55. Premaxillary and anterior dentary fangs ab- 33. Mandibular articulations at approximately sent (0) or present (1). level with occipital condyle (0), displaced to a 56. One or two caniniform teeth present (0) or level distinctly behind occipital condyle (1), or absent (1) on maxilla. positioned anterior to the occipital condyle (2). 57. The maxillary tooth row is restricted to a 34. Exoccipitals do (0) or do not (1) meet dorsal level in front of the posterior margin of the orbit to the basioccipital condyle. (0), or it extends backward to a level below the 35. Supraoccipital exposed more or less verti- posterior corner of the orbit and/or the anterior on cally occiput (0) or exposed more or less hor- corner of the upper temporal fossa (1), or it ex- izontally at posterior end of parietal skull table tends backward to a level below the anterior one (1). third to one half of the upper temporal fossa (2).

36. Occipital crest absent (0) or present (1). 58. Teeth on pterygoid flange present (0) or absent (1). 37. Quadrate with straight posterior margin (0) or quadrate shaft deeply excavated (concave) pos- 59. Vertebrae notochordal (0) or non-notochord- teriorly (1). al (1). 60. Vertebrae 38. Quadrate covered by squamosal and quadra- amphicoelous (0), platycoelous (1), or other tojugal in lateral view (0), or quadrate exposed in (2). lateral view (1). 61. Dorsal intercentra present (0) or absent (1).

36 FIELDIANA: GEOLOGY 62. Cervical intercentra present (0) or absent elongate (0), short (1), or rudimentary or absent (1). (2).

63. Cervical centra rounded (0) or keeled (1) 84. Scapula represented by a broad blade of ventrally. bone (0) or with a constriction separating a ventral 64. Zygosphene-zygantrum articulation absent glenoidal portion from a posteriorly directed dorsal (0) or present (1). wing (1). 85. The dorsal or of 65. Sutural facets receiving the pedicels of the wing process the eosaurop- neural arch on the dorsal surface of the centrum terygian scapula tapers to a blunt tip (0) or is ven- at its end in the dorsal region are narrow (0) or expanded trally expanded posterior (1). into a cruciform or "butterfly-shaped" platform 86. Supraglenoid buttress present (0) or absent

(1). (1).

66. Transverse processes of neural arches of the 87. One (0) or two (1) coracoid ossifications. dorsal short (0) or region relatively distinctly 88. Coracoid of rounded contours (0), slightly elongated (1). waisted (1), strongly waisted (2), or with expand- 67. Vertebral centrum distinctly constricted in ed medial symphysis (3). ventral view or with lateral (0) parallel edges (1). 89. Coracoid foramen enclosed by coracoid os- 68. Distal end of transverse processes of dorsal sification (0) or between coracoid and scapula (1). vertebrae not in diameter or dis- increasing (0) 90. Pectoral fenestration absent (0) or present thickened tinctly (1). (1). 69. absent or Zygapophyseal pachyostosis (0) 91. Limbs short and stout (0) or long and slen- present (1). der (1). 70. and do Prezygapophyses postzygapophyses 92. Humerus rather straight (0) or "curved" (1). not (0) or do (1) show an anteroposterior trend of 93. Deltopectoral crest well developed (0) or re- increasing inclination within the dorsal and sacral duced (1). region. 94. Insertional crest for latissimus dorsi muscle 71. Cervical ribs without (0) or with (1) a dis- prominent (0) or reduced (1). tinct free anterior process. 95. Humerus with prominent (0) or reduced (1) 72. Pachyostosis of dorsal ribs absent (0) or epicondyles. present (1). 96. The ectepicondylar groove is open and 73. The number of sacral ribs is two (0), three notched anteriorly (0), open without anterior (1), or four or more (2). notch (1), or closed (2) (i.e., ectepicondylar fo- 74. Sacral ribs with or without distinct (0) (1) ramen present). expansion of distal head. 97. Entepicondylar foramen present (0) or ab- 75. Sacral ribs or transverse (and caudal) pro- sent (1). cesses sutured (0) or fused (1) to their respective 98. Radius shorter than ulna (0), longer than centrum. ulna (1), or approximately of the same length (2). 76. Cleithrum present (0) or absent (1). 99. Iliac blade well developed (0), reduced but 77. Clavicles broad or narrow (0) (1) medially. projecting beyond level of posterior margin of ac-

1 lon- 78. Clavicles positioned dorsally (0) or anter- etabular portion of ilium ( ), reduced and no oventrally (1) to the interclavicle. ger projecting beyond posterior margin of acetab- reduced 79. Clavicles do not meet in front of the inter- ular portion of ilium (2), or absent (i.e., to dorsal stub) (3). clavicle (0) or meet in an interdigitating antero- simple medial suture (1). 100. Pubis with convex (0) or with concave (1) ventral 80. Clavicles without (0) or with (1) anterolat- (medial) margin. erally expanded corners. 101. Obturator foramen closed (0) or open (1) in adult. 81. Clavicle applied to the anterior (lateral) (0) or to the medial (1) surface of scapula. 102. Thyroid fenestra absent (0) or present (1).

82. Interclavicle rhomboidal (0) or T-shaped 103. Acetabulum oval (0) or circular (1). (1). 104. Femoral shaft stout and straight (0) or slen- 83. Posterior process on (T-shaped) interclavicle der and sigmoidally curved (1).

RIEPPEL: REVISION OF CYMATOSAURUS 37 105. Internal trochanter well developed (0) or re- 111. Calcaneal tuber absent (0) or present (1). duced (1). 1 12. Foot short and broad (0) or long and slender 106. Intertrochanteric fossa deep (0), distinct but (1). reduced (1), or rudimentary or absent (2). 113. Distal tarsal 1 present (0) or absent (1).

107. Distal femoral condyles prominent (0) or not 114. Distal tarsal 5 present (0) or absent (1). projecting markedly beyond shaft (1). 115. Total number of tarsal ossifications four or more three or 108. Anterior femoral condyle relative to poste- (0), (1), two (2). rior condyle larger and extending farther distally 1 16. Metatarsal 5 long and slender (0) or distinct- (0) or smaller/equisized and of subequal extent ly shorter than the other metatarsals and with a distally (1). broad base (1).

109. The perforating artery passes between as- 117. Metatarsal 5 straight (0) or "hooked" (1). or the distal tragalus and calcaneum (0) between 118. Mineralized sternum absent (0) or present heads of tibia and fibula proximal to the astragalus (1).

(1). 119. The medial gastral rib element always only 110. Astragalus without (0) or with (1) a proxi- has a single lateral process (0) or may have a two- mal concavity. pronged lateral process (1).

38 FIELDIANA: GEOLOGY

A Selected Listing of Other Fieldiana: Geology Titles Available

Status of the Pachypleurosauroid Psilotrachelosaurus toeplitschi Nopcsa (Reptilia, Sauropterygia), from the Middle Triassic of Austria. By Olivier Rieppel. Fieldiana: Geology, n.s., no. 27, 1993. 17 pages, 9 illus. Publication 1448, $10.00

Osteology of Simosaurus gaillardoti and the Relationships of Stem-Group Sauropterygia. By Olivier Rieppel. Fieldiana: Geology, n.s., no. 28, 1994. 85 pages, 71 illus. Publication 1462, $18.00

Revised Phylogeny and Functional Interpretation of the Edrioasteroidea Based on New Taxa from the Early and Middle Ordovician of Western Utah. By Thomas E. Guensburg and James Sprinkle. Field- iana: Geology, n.s., no. 29, 1994. 43 pages, 37 illus. Publication 1463, $12.00

Giant Short-Faced Bear {Arctodus simus yukonensis) Remains from Fulton County, Northern Indiana. By Ronald L. Richards and William D. Turnbull. Fieldiana: Geology, n.s., no. 30, 1995. 34 pages, 20 illus. Publication 1465, $10.00

The Genus Placodus: Systematics, Morphology, Paleobiogeography, and Paleobiology. By Olivier Riep- pel. Fieldiana: Geology, n.s., no. 31, 1995. 44 pages, 47 illus. Publication 1472, $12.00

Pachypleurosaurs (Reptilia: Sauropterygia) from the Lower Muschelkalk, and a Review of the Pachy- pleurosauroidea. By Olivier Rieppel and Lin Kebang. Fieldiana: Geology, n.s., no. 32, 1995. 44 pages, 28 illus. Publication 1473, $12.00

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