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FIELDIANA
Geology
NEW SERIES, NO. 40
Sauropterygia from the Middle Triassic
of Makhtesh Ramon, Negev, Israel
Olivier Rieppel
Jean-Michel Mazin
Eitan Tchernov
February 26, 1999
^lblication 1499
§ PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
Si
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Croat, T. B. 1978. Flora ofBarro Colorado Island. Stanford University Press, Stanford, Calif., 943 pp.
Grubb, P. J., J. R. Lloyd, and T D. Pennington. 1963. A comparison ofmontane and lowland rain forest in
Ecuador. I. The forest structure, physiognomy, and floristics. Journal ofEcology, 51: 567-601.
Langdon, E. J. M. 1979. Yage among the Siona: Cultural patterns in visions, pp. 63-80. In Browman, D. L.,
and R. A. Schwarz, eds.. Spirits, Shamans, and Stars. Mouton Publishers, The Hague, Netherlands.
Murra, J. 1946. The historic tribes of Ecuador, pp. 785-821. In Steward, J. H., ed., Handbook of South
American Indians. Vol. 2, The Andean Civilizations. Bulletin 143, Bureau of American Ethnology,
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FIELDIANA
Geology
NEW
SERIES, NO. 40
Sauropterygia from the Middle Triassic
M
of akhtesh Ramon, Negev, Israel
Olivier Rieppel Jean-Michel Mazin
DepartmentofGeology Laboratoire de Geobiologie
FieldMuseum ofNatural History Universite de Poitiers
Roosevelt Roadat Lake Shore Drive Faculte des Sciences Fondamentales etAppliquees
Chicago, Illinois 60615 40, Avenue du RecteurPineau
U.S.A. 86022 Poitiers
France
Kitiin Tchernov
DepartmentofEvolution, Systematics, and Ecology
Hebrew University
Givat-Ram
Jerusalem 91904
Israel
Accepted March 31, 1998
Published February 26, 1999
Publication 1499
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
© 1999 Field Museum of Natural History
ISSN 0096-2651
PRINTED IN THE UNITED STATES OF AMERICA
GEOLOGYLIBRARY
N
Table of Contents 17. Left pubis ofNothosaurus cf. gi-
ganteus 18
18. Left scapula ofNothosaurus giganteus .. 18
Abstract 19. Holotype ofNothosaurus haasi Riep-
1
Introduction pel, Mazin, and Tchernov 19
1
Geological Provenience of the Material 20. Partial skull ofNothosaurus haasi
3 Rieppel, Mazin, and Tchernov 20
Systematic Paleontology 4 21. Holotype ofNothosaurus haasi Riep-
Phylogenetic Interrelationships within pel, Mazin, and Tchernov 21
the Genus Nothosaurus 60 22. Partial skull ofNothosaurus haasi
Stratigraphic and Geographic Distribu- Rieppel, Mazin, and Tchernov 24
tion of the Genus Nothosaurus 69 23. Neural arches ofNothosaurus haasi
Paleobiogeographic History of the Sau- Rieppel, Mazin, and Tchernov 26
ropterygians from makhtesh ramon .. 73 24. Neural arches ofNothosaurus haasi
Summary and Conclusions 78 Rieppel, Mazin, and Tchernov 27
Acknowledgments 78 25. Humeri ofNothosaurus haasi Rieppel,
Literature Cited 79 Mazin, and Tchernov 28
Appendix I: Character Definitions for 26. Humeri ofNothosaurus haasi Rieppel,
Table 5 83 Mazin, and Tchernov 29
27. Femora ofNothosaurus haasi Rieppel,
Mazin, and Tchernov 30
28. Femur ofNothosaurus haasi Rieppel,
List of Illustrations Mazin, and Tchernov 31
29. Holotype ofNothosaurus tchernovi
Haas 32
30. Holotype ofNothosaurus tchernovi
1. Location of the Triassic outcrops at Haas 33
Makhtesh Ramon 2
31. Incomplete skulls ofNothosaurus
2. Stratigraphy of the Triassic at Makh- tchernovi Haas 34
tesh Ramon 4
32. Partial skulls ofNothosaurus tchernovi
3. Holotype of IPsephosaurus mosis Haas 35
Brotzen, 1957 5
33. Partial skull ofNothosaurus tchernovi
4. Incomplete skull of IPsephosaurus Haas 35
mosis Brotzen, 1957 6
5. Lowerjaw fragment of IPsephosaurus 34. Basioccipital ofNothosaurus tchernovi
Haas 37
mosis Brotzen, 1957 6
35. Lowerjaw ofNothosaurus tchernovi
6. Holotype of IPsephosaurus mosis
Haas 38
Brotzen, 1957 7
36. Vertebrae ofNothosaurus tchernovi
7. Incomplete skull of IPsephosaurus
Haas 39
mosis Brotzen, 1957 9
8. Postcranial remains of pachypleuro- 37. Vertebrae ofNothosaurus tchernovi
Haas 40
saurs 10
9. Partial skull of Simosaurus sp 12 38. Vertebrae ofNothosaurus tchernovi
Haas 42
10. Partial skull of Simosaurus sp 13
11. Pectoral girdle ofSimosaurus sp 14 39. Stylopodial elements ofNothosaurus
12. Interclavicula of Simosaurus sp 15 tchernovi Haas 43
13. Sacral rib of Simosaurus sp 15 40. Holotype of Lariosaurus stensioei
14. Left mandible ofNothosaurus cf. N. (Haas) 44
giganteus 16 41. Holotype of Lariosaurus stensioei
15. Dorsal neural arch ofNothosaurus cf. (Haas) 45
N
giganteus 17 42. Interrelationships of species ofLario-
16. Right scapula ofNothosaurus cf. N. gi- saurus 61
ganteus 17 43. Skull fragments ofNothosaurus sp 62
in
44. Premaxillary rostrum ofNothosaurus List of Tables
sp 62
45. Mandibular symphysis ofNothosaurus
sp 63
46. Neural arch ofNothosaurus sp 64 1. Skull proportion in Nothosaurus 22
2. Measurements for humeri ofNothosaurus
47. Scapulae ofNothosaurus sp 64
haasi 28
48. Coracoids ofNothosaurus sp 65 3. Measurements for femora ofNothosaurus
49. Ilia ofNothosaurus sp 67 haasi 30
50. Astragali ofNothosaurus sp 68 4. Measurements for humeri ofNothosaurus
51. Lowerjaw fragment of sauropterygian .. 69 tchernovi 44
52. Lowerjaw fragment of sauropterygian .. 69 5. Data matrix for the analysis of the phylo-
53. Dorsal vertebra of sauropterygian 70 genetic relationships ofLariosaurus .. 48-60
54. Dorsal vertebrae of sauropterygians 70 6. Measurements for neural arches ofNo-
55. Interrelationships of Nothosauridae 72 thosaurus sp 63
56. Clade rank versus age rank for Notho- 7. Measurements for coracoids ofNothosau-
sauridae 72 rus sp 65
57. Cladistic relationships of three species 8. Measurements for ilia ofNothosaurus sp. .. 66
ofNothosaurus 76 9. Data matrix for the analysis of the phylo-
58. Paleobiogeographic scenario 77 genetic relationships ofNothosaurus 71
Sauropterygia from the Middle Triassic of Makhtesh
Ramon, Negev, Israel
Olivier Rieppel Jean-Michel Mazin Eitan Tchernov
Abstract
The Sauropterygia of the Middle Triassic Muschelkalk of Makhtesh Ramon, Negev, are
reviewed, and their evolutionary and paleobiogeographic history is reconstructed on the basis
of cladistic analysis of their interrelationships. The fauna includes cyamodontoid placodonts
(IPsephosaurus) as well as Eosauropterygia such as possible pachypleurosaurs, Simosaurus
N
sp., at least three diagnosable taxa ofNothosaurus (Nothosaurus cf. N. giganteus, haasi n.
sp., and N. tchernovi), and one lariosaur (Lariosaurus stensioei).
The use of cyamodontoids for historical biogeographic analysis remains impeded because
the diagnosis oflower Muschelkalk taxa (IPsephosaurus, Cyamodus) remains incomplete. The
taxic composition ofthe eosauropterygian fauna, as well as the pattern ofcladistic relationships
ofits nothosauriancomponents, indicates paleobiogeographic affinities oftheMakhtesh Ramon
fauna with the Germanic and Alpine realm, supporting the hypothesis of a Burgundian Gate
thatconnectedthe Germanic basin withthe southernbranchofthedeveloping Neotethysduring
upper Anisian and Ladinian times.
Refined analysis ofNothosaurus documents the occurrence of the sister species N. haasi n.
sp. and N. tchernovi at Makhtesh Ramon, which provides evidence fordichotomous speciation
and habitat partitioning within the intraplatform basin habitat characteristic of the northern
Gondwanan shelf(southern margin ofthe developing southern branch ofthe Neotethys) during
Anisian and Ladinian times.
Introduction saurus, along with bones and teeth "ofnumerous
nothosaurian and other reptiles" (Brotzen, 1955,
The first sauropterygian fossil reported from p. 404). In the same year, Peyer (1955) exhibited
Palestine (Israel) was a right humerus of Notho- tetrapods from Makhtesh Ramon at the annual
saurus from the Wady Ayun Musa northeast of meeting ofthe Swiss Paleontological Society, in-
the Dead Sea (Cox, 1924). Cox's (1924) work fo- cluding a skull fragment and ilium ofa large No-
cused mainly on invertebrates, and he found the thosaurus and vertebrae ofthe prolacertilian Tan-
Trans-Jordan Trias to yield a "fauna remarkable ystropheus. The stratigraphy ofthe Makhtesh Ra-
for its unique association of 'Alpine' and 'Ger- mon Muschelkalk was reviewed by Brotzen
man' elements" (Cox, 1932, p. 93). The occur- (1957) in a study that also included the formal
rence of vertebrate fossils in the Middle Triassic description of two cyamodontoid placodonts,
deposits of Makhtesh Ramon in the Negev (Fig. namely Psephosaurus mosis and P. picardi. The
1), southern Israel, was first signaled by Shaw following years witnessed a series of papers by
(1947). A vertebra collected in the area by J. the late Professor G. Haas systematically describ-
Wahrmann in 1949 was described by Swinton ing the vertebrate fossils from the Muschelkalk of
(1952). FurthercollectionsfromMakhteshRamon Makhtesh Ramon (Haas, 1959, 1969, 1975: Pla-
were reported by Brotzen (1955), who identified codontia; Haas, 1963: Micronothosaurus sten-
what he believed to be the earliest occurrence of sioei; Haas, 1980: Nothosaurus tchernovi; Haas,
a cyamodontoid placodont of the genus Psepho- 1981: Simosaurus).
FIELDIANA: GEOLOGY, N.S., NO. 40, FEBRUARY 26, 1999, PP. 1-85
LEBANON
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FIELDIANA: GEOLOGY
During a visit to the Hebrew University of Je- associated the occurrence ofIPsephosaurusmosis
rusalem (huj), one of us (J.-M. M.) surveyed the Brotzen, 1957, Nothosaurus sp., and Tanystro-
extensive collections of vertebrate remains from pheus with that of Beneckeia levantina in his
Makhtesh Ramon that had accumulated since "level 3." The occurrence ofIP. picardi Brotzen,
Haas's work and selected a portion of the mate- 1957, IPsephosaurus spp., Placodus, and Notho-
rial, which was taken out on loan to Paris to be saurus spp. was associated with that of Israelites
prepared using acid. This material forms the core (Hungarites) ramonensis, Protrachyceras wahr-
of the present descriptions, to which was added mani, and Pr. curionii var. ramonensis in "level
other, less well-preserved and/or prepared mate- 5." Translated into his latest revisions (Parnes,
rial that is diagnostic, and a review ofpreviously 1986; see also Parnes et al., 1985), these associ-
described taxa. ations with invertebrate index fossils place the oc-
currence of IP. mosis in the Levantina level of
the Gevanim Formation of late Pelsonian (early
late Anisian) time (Middle Member of the Gev-
Geological Provenience of the anim Formation, upper Bithynian, upper lower
Material Anisian, according to Druckman, 1974). The type
section at Makhtesh Ramon is 68 m thick (Druck-
The marine Middle Triassic (Muschelkalk) of man, 1974). In the following descriptions, these
Makhtesh Ramon in the Negev was deposited in deposits will colloquially be referred to as Be-
a basin that formed by rapid subsidence ofa pre- neckeia beds.
viously continental area (Freund et al., 1975; By contrast, IP. picardi, IPsephosaurus spp.,
Hirsch, 1984). The boundary ofthe subsiding ba- Placodus, and Nothosaurus spp., as indeed the
sin ran across the central and northern Negev bulk of the sauropterygian material here de-
(Freundet al., 1975). Subsidence precededthe on- scribed, with the exception of IP. mosis and pos-
set ofrifting in late Anisian times along what was sibly the Nothosaurus fragment HUJ-Pal. 223, de-
to become the North African coast, and which rive from the middle and upper part of the Fos-
crossed the present Levant coast and Syria to siliferous Limestone Member of the Saharonim
reach the margin of the Neotethys Ocean (Gar- Formation of late Anisian (middle and late Illyr-
funkel & Derin, 1984; May, 1991). The result of ian) and early Ladinian (Fassanian) age. Druck-
these events, initiating the opening ofthe southern man (1974) assigns these deposits to the Lower
branch ofthe Neotethys (Sengoret al., 1984), was Member of the Saharonim Formation, straddling
an embayment ofthe Neotethys that was centered the Anisian-Ladinian boundary; the type section
in Israel and southwestern Syria (May, 1991; Fig. at Makhtesh Ramon is 45 m thick. In the follow-
2). Triassic strata in Israel reach a thickness of ing descriptions, these deposits will colloquially
500 to 1,100 m and are particularly well exposed be referred to as Ceratites beds.
at the Makhtesh Ramon locality in the Negev, lo- The occurrence of IP. mosis, collected by
cated in the center ofthe Triassic-Jurassic "low" Brotzen (1957, p. 199) in situ from the Beneck-
(Freund et al., 1975). eia beds, which on their surface also yielded Hy-
Unfortunately, stratigraphic control on the pro- bodus and reptilian bones preliminarily identified
venience of the sauropterygians within these de- as those of nothosaurs and possibly Tanystro-
posits is rather poor (Brotzen, 1955, 1957; Peyer, pheus, is interesting as it is diachronic with the
1955), most of them having been picked up from later occurrence of abundant cyamodontoid and
the float. However, revision ofthe stratigraphy of nothosaur material. Following the latest strati-
the Middle Triassic deposits at Makhtesh Ramon graphic correlation proposed by Parnes (1986),
by Parnes (1962, 1965, 1975. 1986; Parnes et al., the occurrence of IP. mosis in Israel (upper Pel-
1985; Zak, 1986) provides some indication ofthe sonian) corresponds in time to the occurrence of
age of the horizons from which the material de- Cyamodus tarnowitzensis Giirich, 1884, in the
scribed here is derived (Fig. 2). Karchowice beds of the lower Muschelkalk of
In his initial approach to the biostratigraphy of Upper Silesia (Szulc, 1991; see also Parnes,
the Makhtesh Ramon exposures, Parnes (1962) 1975, Table 3). By contrast, the bulk of the sau-
FlG. 1. Location ofthe Triassic outcrops at Makhtesh Ramon. Negev. Israel (after Parnes, 1986).
RIEPPEL ET AL.: SAUROPTERYGIA FROM MAKHTESH RAMON
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