Bardet&Fernandez 2000 PDF
Bardet&Fernandez 2000 PDF
Bardet&Fernandez 2000 PDF
503–511
Copyright q 2000, The Paleontological Society
0022-3360/00/0074-0503$03.00
ABSTRACT—Two ichthyosaurian specimens from the Upper Jurassic lithographic limestones of Bavaria, namely an almost complete
skeleton with soft tissue impression and another partial one, are described for the first time. Both belong to the same taxon, which is
mainly characterized by a long and slender snout; numerous small, delicate, packed, and well-anchored teeth; a medium size orbit; a
reduced triangular squamosal in the cheek region; an angular largely exposed laterally reaching as far anteriorly as the surangular; a
humerus with three distal facets for radius, intermedium and ulna; an extrazeugopodial element anterior to the radius; a very reduced
hindlimb; packed polygonal paddle elements; and a bipartite pelvis with a distally greatly expanded puboischiatic complex. This
combination of characters permits differentiation from all other known genera; moreover, it could be compared to the species inquirendae
Ichthyosaurus leptospondylus Wagner, 1853a. A new genus, Aegirosaurus, is created and proposed as a new combination for this
species. Aegirosaurus clearly belongs to the clade Ophthalmosauria because of an angular largely exposed laterally and reaching as far
anteriorly as the surangular, and the occurrence of an extrazeugopodial element anterior to the radius and the associated digit distal to
it. A systematic review of ichthyosaurs from the lithographic limestone of Bavaria (most of them destroyed during World War II)
reveals the occurrence of probably three different taxa, namely Aegirosaurus, an indeterminate form close to Ophthalmosaurus or
Caypullisaurus, and an indeterminate one possibly close to Nannopterygius.
TABLE 1—Skeleton measurements (in cm) of Aegirosaurus leptospondylus. packed polygonal elements becoming more spaced and rounded
distally; bipartite pelvis composed of a puboischiatic complex
Measurements BSPHGM SM without foramen and with a distal end expanded more than twice
L (jaw length) 31.4 56.5 the width of the proximal one.
K (skull length) 29.4 56 Etymology.—From Aegir, God of the oceans and the sea-
J (snout length) 19.5 41.5 shores in the Germanic and Scandinavian mythology and Sau-
D (postorbital segment) 1.9? 4.5
H (orbital length) 8.4 10.4 ros, lizard in Greek.
A (orbit center–anterior skull) 23.8 47
C (orbit center–hindmost skull) 5.6 8 AEGIROSAURUS LEPTOSPONDYLUS (Wagner, 1853a)
I (premaxillar segment) 15.2 31.3 new combination
B (prenarial segment) 16 38
Figures 1–8, Tables 1, 2
TABLE 2—Ratios of Aegirosaurus leptospondylus and of other Ophthalmosauria (sensu Motani, 1999a, 1999b). Ratios as defined by McGowan (1976) and
Kirton (1983): OR 5 Orbital ratio (H/L), SR 5 Snout ratio (J/L), PmxR 5 Premaxillary ratio (I/J sensu Kirton, 1983), PnR 5 Prenarial ratio (B/L), PoR
5 Postorbital ratio (D/L), OPI 5 Orbital position index (C/A). Ratios of Brachypterygius, Ophthalmosaurus, Caypullisaurus and Platypterygius based on
McGowan (1976) and Fernández (1997).
FIGURE 1—Aegirosaurus leptospondylus n. comb., SM (neotype), almost complete skeleton with soft tissue impression, Solnhofen Formation, early
Lower Tithonian, Schrandel quarry district, South of Langenaltheim, Bavaria. Global view. Scale equals 20 cm.
1993) and leading to the final buried position mentioned above. sea floor in a lateral position depends on the depth to which they
The lateral rotation of the body could explain the loss of both sank. Here, the carcass has probably intruded the sediment by
the left paddle and girdle. As suggested by Allison (1988), the about 50 percent (see Martill, 1993, fig. 6). This specimen,
soft tissues are subject to bacterial degradation whether the water called ‘‘Munich 5’’ by McGowan (1976), was erroneously en-
column is oxic or anoxic. Thus, the preservation of soft tissue visaged to be Wagner’s more complete skeleton (Häberlein spec-
impression, as well as the complete lack of any scavenger or imen). As previously noted, the old collections from the Munich
epifauna activity on the specimen suggest that its burial was Museum were completely destroyed and BSPHGM 1954 I 608
rapid. was purchased after the war (P. Wellnhofer, personal commun.).
BSPHGM 1954 I 608 is a partial articulated skeleton, 0.70 m Skull and mandible.—In both specimens the skull and man-
long, exposed on a lithographic slab in left lateral view (Fig. 2). dible are complete and articulated. The SM skull is 56 cm long
The preserved parts of the skeleton are the completety articu- and lacks only the anteriormost part of the premaxillae (Fig. 3).
lated skull and trunk region with disarticulated girdles and pad- The BSPHGM skull is complete and is about 30 cm long (Fig.
dles. The tail region is absent. According to Martill (1993), the 4). Both skulls are mainly characterized by a delicate and long
completeness of the body outline of carcasses that arrive on the snout fitted with numerous teeth, and a medium size orbit. The
FIGURE 2—Aegirosaurus leptospondylus n. comb., BSPHGM 1954 I 608 (referred specimen), partial skeleton, Solnhofen Formation, early Lower
Tithonian, Steinbruch am Geisberg quarry, Apfeltal, South of Solnhofen, Bavaria. Global view. Scale equals 20 cm.
506 JOURNAL OF PALEONTOLOGY, V. 74, NO. 3, 2000
FIGURE 5—Aegirosaurus leptospondylus n. comb., SM (neotype), Sol- trapezoidal and larger than the radius; the radiale is ovoid; the
nhofen Formation, early Lower Tithonian, Schrandel quarry district, intermediium is wedge-shaped and elongated; the ulnare is qua-
South of Langenaltheim, Bavaria. Forelimb with soft tissue impression, drangular; there is a pentagonal extrazeugopodial element an-
1, photograph; 2, interpretative drawing. Abbreviations: e, extrazeu- terior to the radius and a pisiform posterodistal to the ulna, nei-
gopodial element; H, humerus; i, intermedium; p, pisiform; R, radius; ther in contact with the humerus; the metacarpal V contacts the
r, radiale; ti, tissue impression; U, ulna; u, ulnare; 2–4, distal carpals; ulnare and the distal carpal 4; digit I is considered to be lost;
ii-v, metacarpals; grey zones are covered with matrix; dotted line ele- there are three digits anterior to the primary axis (sensu Motani,
ments are preserved as matrix impression. 1999a), which corresponds to digit IV, the longest one with 23
elements. It should be noted that between digits III and IV there
In BSPHGM, only about 34 presacral vertebrae are preserved are two small isolated elements that may or not be remains of
in articulation (see Fig. 2). an additional extra digit. The elements are strongly packed and
In both specimens, the height of the presacral centra is about polygonal in the proximal part of the paddle but become more
twice their length. The neural arches are rather low, rectangular, rounded and spaced in its distal part, as in Brachypterygius and
and slightly posteriorly recurved in the trunk region. They be- Caypullisaurus. The epipodial construction of Aegirosaurus (in-
come very low in the caudal region. In BSPHGM, some dis- termedium contacting the humerus) is comparable to that of Bra-
placed vertebrae show that the neural canal was large. chypterygius (McGowan, 1997) and is quite different from that
Ribs are preserved in articulation with the vertebrae in both of Ophthalmosaurus, Platypterygius and Caypullisaurus in
specimens. The anterior presacral ribs are short, wide and bear
confluent parapophyses and diapophyses located anteriorly. The
other ribs are very long and slender, as usual in ichthyosaurs.
Some delicate gastralia are preserved disarticulated in
BSPHGM.
Limbs.—Right fore- and hindlimbs are completely preserved
in articulation and with soft tissue impression in SM (Figs. 1, 5,
6). In BSPHGM, disarticulated elements of possibly both fore-
paddles and of the right hindlimb are preserved (Figs. 2, 7). In
both specimens, the hindlimb is strongly reduced compared to
the animal size, and is about half the size of the forelimb.
The humerus is a massive, short element, clearly belonging
to the derived ‘morphotype 3’ of Motani (1999a). As in Bra-
chypterygius, Ophthalmosaurus, Caypullisaurus and Platypter-
ygius, there are three distal facets. Those for the radius and ulna
are similar in size whereas the third facet is smaller and herein
interpreted as for the intermedium, like in Brachypterygius. The
facets are angulated and clearly separated from each other, es- FIGURE 7—Aegirosaurus leptospondylus n. comb., BSPHGM 1954 I 608
pecially the radial facet which is antero-distally oriented, also (referred specimen), Solnhofen Formation, early Lower Tithonian,
Steinbruch am Geisberg quarry, Apfeltal, South of Solnhofen, Bavaria.
like in Brachypterygius. The humerus proximal head is almost Limb and girdle interpretative drawings; 1, forepaddle and pectoral
flat and the distal extremity is wider than the proximal one. girdle; 2, hindpaddle and pelvic girdle. Abbreviations: c, centra; cl,
Based on the conservative topological features of the primary clavicle; co, coracoid; F, femur; H, humerus; ic, interclavicle; il, ilium;
axis and digital arch as described by Motani (1999a), the ho- m, mandible; o, orbit; pi, puboischiatic complex; r, ribs; s, skull; sc,
mology of the Aegirosaurus forefin elements is identified as fol- scapula. Grey zones indicate cuts on the lithographical slab. Scale
lows (see Fig. 5.2): the radius is roughly pentagonal; the ulna is equals 5 cm.
508 JOURNAL OF PALEONTOLOGY, V. 74, NO. 3, 2000
at least Ophthalmosaurus, Brachypterygius, Caypullisaurus and BARTHEL, K. W., N. H. M., SWINBURNE, AND S., CONWAY MORRIS.
Platypterygius (Motani, 1999a, 1999b). Within the Ophthalmo- 1990. Solnhofen: A study of Mesozoic palaeontology. Cambridge
sauria, Ophthalmosaurus, Caypullisaurus and Platypterygius University Press, Cambridge, 236 p.
differ from Aegirosaurus by their epipodial arrangement (extra- BAUER, F. 1898. Die Ichthyosaurier des oberen weissen Jura. Palaeon-
tographica, 44:283–328.
zeugopodial element 1 radius 1 ulna). Aegirosaurus shares BLAINVILLE, H. DE. 1835. Système d’Herpétologie. Nouvelles Annales
with Brachypterygius the same forefin construction and espe- du Museum d’Histoire Naturelle, 4:37–295.
cially the humerus-intermedium contact, but differs from it in CONYBEARE, W. D. 1822. Additional notices on the fossil genera Ich-
several cranial characters such as the delicate snout fitted with thyosaurus and Plesiosaurus. Transactions of the Geological Society
numerous small teeth, the number of sclerotic plates and the of London, 1st series, 1(1):103–123.
short jugal. DE LA BECHE, H. T., AND W. D. CONYBEARE. 1821. Notice on the
Pending a more precise description and illustration of several discovery of a new fossil animal, forming a link between the Ichthy-
taxa, notably from Russia (Arkhangelsky, 1997, 1998, 1999; Efi- osaurus and the crocodile, together with general remarks on the os-
mov, 1998, 1999a, 1999b) and a detailed comparative study of teology of the Ichthyosaurus. Transactions of the Geological Society
of London, 1st series, 5(2):559–594.
the taxa included into the Ophthalmosauria, precise phylogenetic EFIMOV, V. M. 1998. An Ichthyosaur, Otschevia pseudoscythica gen. et
relationships of Aegirosaurus among Ophthalmosauria cannot be sp. nov. from the Upper Jurassic strata of the Ulyanovsk Region (Vol-
traced yet. ga region). Paleontological Journal, 32(2): 87–191.
. 1999a. Ichthyosaurs of a New Genus Yasykovia from the Upper
CONCLUSION Jurassic Strata of European Russia. Paleontological Journal, 33(1):
A systematic review of the ichthyosaurs from the Late Juras- 91–98.
sic of Bavaria, based on the bibliographical revision of old col- . 1999b. A New Family of Ichthyosaurs, the Undosauridae fam.
lections (most destroyed during World War II—see Historical nov. from the Volgian Stage of the European Part of Russia. Pale-
ontological Journal, 33(2):174–181.
account and Appendix) and the description of two new speci- FRAAS, E. 1891. Die Ichthyosaurier der süddeutschen Trias- und Jura-
mens, reveals that three different taxa were probably present Ablagerungen. Laupp’schen Buchandlung, Tübingen, 81 p.
during Tithonian times during deposition of the lithographic FERNÁNDEZ, M. S. 1997. A new ichthyosaur from the Tithonian (Late
limestones of the Solnhofen area: Aegirosaurus leptospondylus Jurassic) of the Neuquén Basin, Northwestern Patagonia, Argentina.
(Wagner, 1853a) new combination, an indeterminate form close Journal of Paleontology, 71:479–484.
to Ophthalmosaurus or Caypullisaurus (Häberlein specimen— . 1998. Nuevo material de Caypullisaurus bonapartei Fernández
see Appendix) and an indeterminate one possibly close to Nan- (Reptilia: Ichthyosauridae) del Jurásico superior de la Cuenca Neu-
nopterygius (Solhnofen specimen—see Appendix). This study quina, Argentina. Ameghiniana, 35(1):21–24.
shows that ichthyosaurs from the lithographic limestones of Ba- FRICKHINGER, K. A. 1994. Die Fossilien von Solnhofen: Dokumentation
der aus den Plattenkalken bekannten Tiere und Pflanzen—The Fossils
varia were not as scarce as previously thought and that during of Solnhofen: Documenting the Animals and Plants known from the
Tithonian time, ichthyosaurs were more diversified worldwide Plattenkalks. Soldschneck, Korb, 333 p.
than classically assumed. GODEFROIT, P. 1994a. Les reptiles marins du Jurassique inférieur en
Lorraine belgo-luxembourgeoise. Unpublished Ph.D. thesis, Catholic
ACKNOWLEDGMENTS University of Louvain-la-Neuve, 359 p.
The authors greatly thank M. Schwegler (Schwegler Museum, . 1994b. Les reptiles marins du Toarcien (Jurassique inférieur) bel-
Langenaltheim, Bavaria) for his generosity of making available go-luxembourgeois. Mémoires pour servir à l’Explication des Cartes
the neotype specimen for study and for his hospitality. We Géologiques et Minières de la Belgique, 39:1–98.
HUENE, F. VON. 1922. Die Ichthyosaurier des Lias und ihre Zusammen-
warmly thank P. Wellnhofer (BSPHGM, Munich, Bavaria) for hänge. Sebrüder Borntraeger, Berlin, 114 p.
giving us the opportunity to study the specimens, as well as for HULKE, J. W. 1870. Note on some teeth associated with two fragments
his great assistance and kindness. Many thanks also to S. Chap- of jaw from Kimmeridge Bay. Quartely Journal of the Geological
man (NHM, London) for the access to the ichthyosaurian col- Society of London, 26:172–174.
lections in her care. Special thanks to M-A. Lançon (CNRS, JAECKEL, O. 1904. Eine neue Darstellung von Ichthyosaurus. Zeitschrift
Paris) for her extensive help in the German literature translations der deutschen geologischen Gesellschaft, 56:26–34.
and to P. K. Tubbs (ICZN, NHM, London) for his helpful no- JOHNSON, R. 1977. Size independent criteria for estimating relative age
menclatural advice. We are grateful to D. Serrette (CNRS, and the relationship among growth parameters in a group of fossil
MNHN, Paris) and to G. Bergmeier (BSPHGM, Munich) for the reptiles (Reptilia: Ichthyosauria). Canadian Journal of Earth Science,
photographs. Finally, we thanks the two referees J. Massare 14:1916–1924.
KIPRIJANOFF, W. 1881. Studien über die fossilen Reptilien Russlands.
(New-York) and R. Motani (Toronto) for their helpful and con- Theil 1: Gattung Ichthyosaurus König aus dem severischen Sandstein
structive comments which have permitted us to greatly improve oder Osteolith der Kreide-Gruppe. Mémoires de l’Académie Impéri-
our manuscript. ale des Sciences de St.-Pétersbourg, 28(8), 103 p.
KIRTON, A. M. 1983. A review of British Upper Jurassic ichthyo-
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dans l’argile de Kimmeridge par M. Lennier, au Cap de la Hève près lithographic limestones of Bavaria was mentioned as ‘an exemplar
le Havre. Comptes Rendus Hebdomadaires de l’Académie des Sci- found with polygonal bones in the flippers and draughtboard-shaped
ences, 53:267–273. vertebral centra’ (Quenstedt, 1852, p. 129). It was assigned to I. lep-
. 1861b. D’un nouveau reptile très voisin du genre Ichthyosaurus, tospondylus by Wagner (1861) and later considered by Fraas (1891, p.
trouvé dans l’argile du Kimmeridge de Bléville, au nord du cap la 74) as the new type of this species. This small and rather well preserved
specimen came from Solnhofen and was part of the Häberlein collection
Hève du Havre. Comptes Rendus Hebdomadaires de l’Académie des
(Pappenheim). This specimen was 90 cm long and included the skull
Sciences, 53:999–1001. and mandible, teeth, vertebrae, ribs, pectoral girdle and paddle elements.
WAGNER, A. 1852. Neu aufgefundene Saurier Ueberreste aus den lith- Following the description given by Wagner (1861, p. 120–121), Fraas
ographischen Schiefern und dem obern Jurakalke. Abhandlungen der (1891, p. 74–75) and Bauer (1898, p. 295–299), this individual was
Mathematische-Physische Classe der königlische bayerische Akade- estimated to have been about 1.5 m long, of which the skull occupies
mie der Wissenschaft, 6:661–710. 35 cm. It was mainly characterized by a large orbit (compared to the
. 1853a. Die Characteristic einer neuen Art von Ichthyosaurus aus animal size) filled entirely by the sclerotic ring; numerous, small, slen-
den lithographischen Schiefern und eines Zahnes von Polyptychodon der and almost smooth teeth (height about 1.2 cm) weakly anchored in
aus dem Gründsandsteine von Kelheim. Bulletin der königlische Aka- the jaws (they were all preserved separately); a robust mandible; short
demie der Wissenschaft, Gelehrte Anzeigen, 3:25–35. vertebrae with a diameter more than twice the length (diameter about
. 1853b. Beschreibung einer fossilen Schildkröte und etlicher an- 2.3 cm, length about 0.8 cm); a pectoral girdle composed of slender
derer Reptilien—Ueberreste aus den lithographischen Schiefern und scapula, rounded and unnotched coracoid and pointed interclavicle; and
dem Gründsandsteine von Kelheim. Abhandlungen der Mathematis- a pelvic girdle represented only by an elongated ilium (pubis of Wagner,
che-Physische Classe der königlische bayerische Akademie der Wis- 1861; Bauer, 1898, p. 299). The most diagnostic elements described by
senschaft, 7(1):239–264. Bauer (1898, p. 299) were those of the paddle. The humerus (basi-
. 1861. Neue Beiträge zur Kenntniss der urweltlichen Fauna des phenoid of Wagner, 1861) was described as short and robust (length 5
BARDET AND FERNANDEZ—JURASSIC ICHTHYOSAUR FROM BAVARIA 511
3.2 cm, proximal width 5 2.4 cm, distal width 5 3.2 cm) with three The specimen was 1.45 m long (estimated length about 2–2.5 m) and
distal facets. The two largest for radius and ulna were almost equal in mainly characterized by a long and slender skull about 41 cm long;
size (1.6 cm and 2 cm) and at 131 degrees to each other; whereas the large orbits (diameter about 7 cm) completely filled by the sclerotic
smallest (few mm) for the pisiform was dorsally oriented. The femur ring; elongated nares tapering anteriorly as in the Meyer specimen; pre-
was preserved as an impression in the matrix. It was described as short maxillae long and slender, forming part of the lower narial opening;
(length 5 1.9 cm, proximal width 5 1 cm, distal width 5 1.5 cm) with maxillae excluded from the nares; jugals located above the maxillary
two distal facets (0.9 cm and 0.7 cm). Finally, about 100 paddle ele- and forming the lower orbital margin; triangular nasals taking part in
ments, ranging from 1.5 cm to 0.3 cm were preserved, the largest being the nares and articulating with premaxillae and lacrymals; pterygoids
penta- or hexagonal whereas the smallest being rounded. bearing two branches at an angle of 50–60 degrees; basicranium robust;
According to Wagner (1861), this specimen belongs to the same spe- teeth small (height about 0.7 to 1.5 cm) and very variable in crown
cies as the Oberndorfer one and differs from it only in its relative size ornamentation, being either smooth or striated; mandible about 37.5 cm
and mandible height. On the contrary, for Fraas (1891, p. 74–75), the long with a powerful dentary; fifty short vertebrae (diameter about 2
Häberlein specimen was clearly different from the Oberndorfer one and cm, length about 1.2 cm) with diapophyses confluent with neurapo-
considered as the new type and only specimen referable to I. leptos- physes; short and robust femur (height 5 2 cm) with two articular
pondylus. Finally, Bauer (1898), considered these two specimens as facets; polygonous paddle elements (2 mm to 2 cm); pectoral girdle
belonging to the same species, the Häberlein one being considered as including elongated coracoids (length 5 6.54 cm, width 5 4.4 cm) with
a juvenile. Bauer (1898) also noted that the humerus of the Häberlein an anterior notch, scapulae (length 5 7.5 cm) and clavicle (length 5
specimen looks like that of Ophthalmosaurus cantabrigiensis and that 11 cm); and a tripartite pelvic girdle composed of rodlike ilium (length
its hindlimb was very reduced compared to the forepaddle. 5 4.24 cm), bootlike ischium (length 5 2.93 cm) and sticklike pubis
For Fraas (1891), the short vertebrae and great number of paddle (length 5 1.98 cm).
elements suggest a species close to Ophthalmosaurus or Baptanodon. This specimen was assigned by Bauer (1898) to I. trigonus var. pos-
We confirm here his view, arguing that the Oberndorfer and Häberlein thumus along with all the ichthyosaurs from the Late Jurassic of Ba-
specimens belong to different taxa. The above mentioned characters, varia. Some characters exhibited by the Solnhofen specimen such as a
namely a large orbit, small teeth all preserved out of the jaws (which slender skull, a medium size orbit (ratio estimated to about 0.19), small
seem thus edentulous), the humerus shape and the important reduction teeth, small and short thoracic vertebrae, an elongated and anteriorly
of the hindlimb, indicate that the Häberlein specimen has possible af- notched coracoid, a tripartite pelvis and finally a femur with two distal
finities with Ophthalmosaurus or Caypullisaurus. Nevertheless, the very facets, tend to exclude Brachypterygius and Ophthalmosaurus but are
poor illustration of the material prevents any accurate identification. It comparable to Nannopterygius. However, because of the scarcity of the
is considered as an indeterminate form, possibly close to Ophthalmo- illustrations given by Bauer (1898) it is not possible to assign the Sol-
saurus or Caypullisaurus. nhofen specimen with certainty to this genus and it is thus only con-
sidered as an indeterminate ichthyosaur, possibly close to Nannoptery-
BMNH 42833 (London).— An incomplete skull from the lithograph-
gius.
ical limestones of Eichstätt, part of the Krantz collection (Bonn), re-
Tail (Munich State Museum, destroyed during WWII).—The lunate
ferred to I. leptospondylus (Meyer, 1863, p. 222) and later to I. posthu- tail 90 cm high, with soft tissue impression, was from a large specimen
mus (Fraas, 1891, p. 73). estimated to be about 4 m long, referred to I. trigonus var. posthumus
This portion of skull, 31.2 cm long, preserved the snout back to the (Bauer, 1898; Merriam, 1908). It comes from the lithographic lime-
nares. Its total length was estimated to about 42 cm (Meyer, 1863). The stones of an indeterminate locality in Bavaria. The angle of downturn
snout is long, slender (length about 28 cm) and not markedly demar- (about 80 degrees) was very high. There were about 12 vertebrae pre-
cated from the skull in lateral view. It bears numerous minutely ridged served before the tail-bend and 70 after, the last one being elongated
or smooth, small teeth, well anchored in the dental groove and extend- with bulged borders. This specimen does not exhibit characters permit-
ing almost under orbit. The nares is elongated (2.7 cm) and tapers an- ting a precise identification and may be only assigned to an indeter-
teriorly. On the basis of these combination of characters, this specimen minate ichthyosaur.
is here referred as to cf. Aegirosaurus leptospondylus. Rib and vertebrae (Munich State Museum, destroyed during
One tooth and two vertebrae (Munich State Museum, destroyed dur- WWII).—A slab from Kelheim mentioned by Bauer (1898, p. 295) in-
ing WWII).—An isolated tooth from Eichstätt (Fraas, 1891, pl. 11, fig. cluded about 8 abdominal ribs and 12 vertebrae. As no description was
18) and a vertebra from Solnhofen (Fraas, 1891, pl. 12, fig. 5) were given and as the material is poorly diagnostic, it could only be assigned
attributed to I. posthumus. Another vertebra from Kelheim (Fraas, 1891, to an indeterminate ichthyosaur.
pl. 12, fig. 6) was referred to as I. leptospondylus. These three speci- Paddle elements (Munich State Museum, destroyed during WWII).—
mens do not show any diagnostic character permitting a precise iden- A slab from Solnhofen mentioned by Bauer (1898, p. 300) consisted of
tification and they are assigned to indeterminate ichthyosaurs. articulated paddle elements exhibiting four primary digits (radial, ulnar,
Solnhofen specimen (Munich State Museum, destroyed during intermedium and pisiform). No detailed description was given of this
WWII).—An incomplete skeleton unearthed from the lithographic lime- material. The limb construction and especially the occurrence of an
stones of Solnhofen, including skull elements, vertebrae, ribs, parts of intermedium suggests either Aegirosaurus or Brachypterygius but it is
pectoral and pelvic girdles. The description given by Bauer (1898, p. impossible to determine in this specimen, which is referred as an in-
300–309) was extensive but the illustrations were very poor. determinate ichthyosaur.