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28 April 2022 The Tübingen collection of ichthyosaurs from the Lower Jurassic (Lower Toarcian) Posidonienschiefer Formation of Württemberg: a historical and curatorial perspective
Henrik Stöhr, Ingmar Werneburg
Author Affiliations +

The Paleontological Collection of Tübingen University houses one of the largest collections of ichthyosaur specimens from the Lower Jurassic (Lower Toarcian) Posidonienschiefer Formation fossillagerstätte in the world. It forms an important basis for numerous past and ongoing studies on the taxonomy, evolutionary morphology, ecology, and other aspects of ichthyosaur biology. The collection includes particularly significant material, such as several type specimens, representatives of rare species, and different ontogenetic stages, which show varying degrees of preservation. Founding fathers of paleontology at Tübingen University, including Friedrich August Quenstedt (1809–1889) and Friedrich v. Huene (1875–1969), assembled the majority of these specimens and conducted extensive research using Tübingen ichthyosaurs. Many more recent publications also use Tübingen material as a reference. Unfortunately, in many cases the identity and provenience of old museum specimens are not adequately known. This has led to confusion, inconsistencies, and errors in the literature. Here we present a detailed assessment of the history and identity of the ichthyosaurs from the Posidonienschiefer Formation in the Tübingen collection by conducting both a comprehensive literature survey and a re-investigation of the entire collection. We consulted document archives and critically compared them to hand-written specimen labels. In total, we were able to clearly identify and illustrate 78 articulated specimens that are now fully accessible for international researchers with clear documentation of their stratigraphical allocation. With respect to old natural history collections in general, we provide a broader discussion on how to deal with historical specimens (which sometimes represent composites of several individuals) and identify a series of challenges when dealing with confusing documentation. Our study attempts to provide means to solve these issues to facilitate and provide a more reliable database for future research.

1. Introduction

Ichthyosaur research has a long history. These animals are among the first Mesozoic reptiles that became known to science. Discoveries of ichthyosaur vertebrae were documented early in the history of paleontological research (Lhuyd 1699). The first articulated skeleton which was recognized as a new type of fossil marine reptile was discovered around 1812 by Mary Anning (1799–1849) resulting in the first scientific descriptions (Home 1819). Those initial publications inspired many scientists (DE La Beche & Conybeare 1821; Conybeare 1822) and contributed to a significant rise of public interest in paleontology in early 19th century Europe. Today, numerous questions regarding the life history of ichthyosaurs remain debated (Buchholtz 2001; Riess 1986). How did the animals breathe, how fast did they swim, and how deep could they dive? How did they orient themselves, did they perform annual migrations, did they move to particular nursery areas for birth? In addition, specific anatomical characteristics of particular taxa remain unexplained such as the extreme elongation of the premaxilla with its full dentition compared to a relatively short lower jaw in Eurhinosaurus longirostris (Quenstedt 1885; Fraas 1891: 64; v. Huene 1922a, 1922b). Despite the availability of a large number of well-prepared specimens and detailed documentation of the stratigraphic distribution (v. Huene 1931), there is still discussion on the taxonomy and the diagnostic features of juveniles (Mcgowan 1979; Mazin 1982; Mcgowan 1986; Godefroit 1994; Motani 1999; Maisch & Matzke 2000; Sander 2000; Maisch 2010; Maxwell 2012; JI et al. 2015; Moon 2019).

Important anatomical details were discovered in the last decades, contributing to a more complete picture of ichthyosaurs in general (Lingham-Soliar 2001; Vinther 2015; Lindgren et al. 2018; Pardo-PÉrez et al. 2018a, 2018b), although Fraas (1888: 280) argued that, based on the enormous quantity of material already available at his time, not much important progress in understanding these animals could be expected. Noteworthy, the phylogenetic origin of ichthyosaurs is less well understood when compared to many other groups of amniotes (Baur 1887; Massare & Callaway 1990; Maisch 2010; Fischer et al. 2013; FrÖbisch et al. 2013; Liu 2015). The extinction of the whole group does not correlate with the mass extinction of other major groups of Mesozoic reptiles (dinosaurs, mosasaurs, plesiosaurs, pterosaurs, and others) at the Cretaceous-Paleogene boundary (66 my ago), but happened already in the Cenomanian (93.9–100.5 my ago) (Massare 1988; Fischer et al. 2016), raising additional scientific questions that have yet to be dealt with using modern scientific and methodological approaches.

One of the richest fossil lagerstätten for ichthyosaurs are those of the Posidonienschiefer Formation (Posidonia Shale, Lias ε) which crops out in the foreland of the Swabian Alb, Baden-Württemberg, Southwestern Germany. These sediments are Lower Jurassic (Early–Middle Toarcian, c. 183–185.6 my) in age. Most ichthyosaur specimens from the Posidonia Shale are stored in the Staatliches Museum für Naturkunde in Stuttgart (acronym: SMNS) and the nearby Paläontologische Sammlung der Universität Tübingen (acronym: GPIT; from the former “Geologisch-paläontologisches Institut Tübingen”), Southern Germany. The paleontological collection of Tübingen is one of the worldwide oldest natural history collections, dating back to the end of the 15th century, when the University of Tübingen was founded (HÖlder 1977; Seidl et al. 2021; Werneburg 2021). With more than one million specimens, it is still one of the worldwide largest university collections of its kind (Werneburg 2016). Founding fathers and pioneers of paleontology and its different sub-disciplines worked in Tübingen, including Friedrich August Quenstedt (1809–1889), Friedrich v. Huene (1875–1969), OTTO Schindewolf (1896–1971) and Adolf Seilacher (1925–2014), a history well represented also by a written document archive (Hinz & Werneburg 2019). Since 2009, the collection is curated by the national German Senckenberg Society, which has the ultimate goal to bring the collection infrastructure to the most modern standards in the field (Werneburg & BÖHme 2018). These include traceable collection management and specimen cataloging. In reality, however, dealing with historical museum material is far more complex than just establishing a logical inventory of specimens. More than 500 years of collection history “accumulated” also the history of science in general, which manifests not only in purely scientific terms, such as different approaches to taxonomy, but also in collection organization and management (Falk et al. 2018).

When studying museum specimens in paleontology, it is important to keep in mind that historical preparations used very different sediments and other materials to “supplement” the specimens on display for a coherently aesthetic impression. In this context, various techniques were applied to vitiate the original impression of the specimen (discussed by Fraas 1891: 51; Hungerbühler 1994; Maisch & Matzke 2000; Maisch 2008). Moreover, in the Posidonia Shale, extreme deformation took place during fossilization in most specimens (Hofmann 1958). Knowing the individual history of each specimen is of vital importance to arrive at sound scientific conclusions. For Posidonia Shale fossils in general and the ichthyosaurs in particular, Bernhard Hauff sen. (1866–1950) is one of the most important figures in paleontology. He unearthed numerous skeletons from his private quarry in Holzmaden, Southern Germany (Fig. 1D), and assembled many specimens from other quarries in the area, which he also documented stratigraphically as exactly as possible at the time. Based on a specifically developed preparation technique, several complete skeletons, some of them with soft tissue preservation, were discovered (Fraas 1894; Hauff 1921). The preserved skins of these specimens show a clear contour of the deceased organism (Fraas 1894; Heller 1965; Keller 1992). In addition, well-preserved stomach contents (Fraas 1891; Keller 1976; BÖttcher 1989) and even embryos were discovered and described from Posidonia Shale ichthyosaurs (Jaeger 1824; Branca 1908; BÖttcher 1990). In 1921, B. Hauff received an honorary doctorate of natural sciences from Tübingen University for his important contributions (v. Huene 1944).

One of the core complexes of the paleontological collection of Tübingen University is the fauna of the Posidonia Shale Fossillagerstätte of Baden-Württemberg, including numerous ichthyosaurs (Figs. 1, 2). Ichthyosaur specimens were continuously acquired for the collection at least since 1843 (e.g., the specimen illustrated in Fig. 3). The specimens housed in Tübingen are very diverse both taxonomically and preservationally. Many were transferred in wooden frames highlighting their aesthetic impression (Bierende 2021). A considerable number of scientific publications deals with the Tübingen ichthyosaur collection, but some of these studies also introduced a number of confusing misinformation in the literature regarding the identity and stratigraphic allocation of particular specimens. Reasons for that might have been: (1) the fluctuating arrangement of the exhibition over decades, which made it difficult to trace superficially described specimens from the literature; (2) missing or changing catalog numbers; as well as, (3) ever-changing opinions about the taxonomic diversity within these ichthyosaurs and the associated emphasis on single taxa (e.g., Godefroit 1994; Sander 1989, 2000; Maisch & Matzke 2000; Maisch 2010) or on selected ontogenetic stages (Hungerbühler 1994).

The last point, taxonomy, is out of our expertise, and resolving this issue should be left to specialists. As a curatorial team in Tübingen, we are able to help in resolving the first two issues, namely by tracing the history of particular specimens. This might sound like a local problem that should be restricted to within-house documentation. In our opinion, this is not the case for several important reasons. Firstly, numerous active researchers from all over the world visit Tübingen regularly to study the diversity, taxonomy, anatomy and taphonomy of ichthyosaurs or rely on published data on the Tübingen collection for comparative purposes in their own research. The Tübingen collection of Lower Jurassic ichthyosaurs is among the three most important ones in the world, alongside those of the Natural History Museum, London, and the Staatliches Museum für Naturkunde, Stuttgart. International researchers require a traceable introduction to the peculiar problems of the historical Tübingen collection to facilitate their studies and to avoid future errors in the literature. Secondly, and more important for a wider audience, we wish to use the Tübingen ichthyosaur collection as an example to highlight and discuss the general issues associated with old scientific collections that have a long research history and illustrate a consistent, specimen-related approach that demonstrates possibilities to solve the collection management issues reported above. Thirdly, we also want to further promote the exceptional collection of Tübingen ichthyosaurs to the scientific community to inspire future research on evolutionary, paleontological and geological topics.

2. Material and methods

2.1. Choice of material

In our study, we concentrated exclusively on articulated ichthyosaur and assemblages of associated material from the Lower Toarcian Posidonienschiefer Formation of Baden-Württemberg (Fig. 1). In addition, there are numerous isolated ichthyosaur elements in the collection; mostly the uniform amphicoelous vertebral centra that are in most cases almost impossible to determine beyond the family or at most genus level. These isolated elements are not treated here, as they are of minor scientific interest. Ichthyosaur material from stratigraphical layers other than the Posdionienschiefer Formation, as well as from other localities housed in the Tübingen collection was not considered in our study. These include, e.g., many important Middle Triassic ichthyosaurs from Monte San Giorgio, Switzerland, and a substantial collection of Middle Jurassic ophthalmosaurids from Peterborough, United Kingdom (Lerche 2021).

2.2. Literature analysis

We started our literature review with the extensive study of v. Huene (1931), who listed most specimens from the Tübingen collection (Fig. 4) and also illustrated several important specimens. We then traced the history of each specimen back to the publications of the founder of the paleontological collection sensu stricto, Friedrich August Quenstedt (Werneburg 2017) (note that the ‘natural history collection’ of Tübingen University is much older, but the paleontological collection was inadequately documented before Quenstedt), and other publications of the 19th century (e.g., Fraas 1891). In the next step, with a clearer understanding of specimen identity, we traced the history of each specimen until today (e.g., Maisch 2008; Pardo-PÉrez et al. 2019; Maxwell & CortÉS 2020) (Fig. 5). In the course of this study, we covered only the most important publications using Tübingen ichthyosaurs. Further, minor important publications exist that only illustrate or briefly discuss Tübingen ichthyosaurs (e.g., HÖlder & Steinhorst 1964); however, these were mainly left out of further consideration because based on the major publications cited here, it was already possible to clearly trace and identify every single specimen.

We had access to an unpublished specimen catalog of Jane Ann Robinson from November 1974, now stored at the University Archive Tübingen (UAT 578/591, here referred to as “Robinson 1974”). Old photographs of the collection distributed among different UAT-resources were also helpful in reconstructing to a certain degree the fate of the specimens' display in the exhibition (Fig. 6). Specimen identification was also possible based on old labels near or attached to the specimens in the collection itself (Fig. 5). We discovered a black wooden box containing old photographs of specimens from the famous Tübingen ichthyosaur researcher Friedrich v. Huene – material now stored at the University Archive Tübingen (UAT 678/592). This box and these photographs had been available to Michael Maisch and were used in his study of Stenopterygius (Maisch 2008: 230). Colleagues cited in the text orally communicated the more recent history of some specimens. Finally, we had access to the Geologenarchiv (GA) housed at the university library of Freiburg, Germany, in which letters from Bernhard Hauff to Friedrich v. Huene are stored, and to the Kreisarchiv Esslingen, Germany (OST). The letters of the archives were used, among others, to get a better idea of previously undefined stratigraphical allocations of particular specimens.

2.3. Specimen numberation

Translations of German quotes are by us. Different acronyms of the Tübingen collection exist: “PV” is the abbreviation for “Petrefactenverzeichnis”, an inventory catalog with limited specimen information used from 1841 until 1988. In the second part of the 20th century, the acronym “GPIT” (Geologisch-paläontologisches Institut Tübingen) was introduced, but only applied to published specimens; e.g., “GPIT/1234-1”, with “-1” referring to the first specimen of material used in a published study numbered “1234”. Since the end of the 20th century, with the development of computer-aided catalogizations, the acronym GPIT was incrementally applied to all specimens in the collection with a supplemented reference to the sub-collection to which the specimen belongs: e.g., “GPIT/ RE/4567”, with “RE” meaning “reptile”-sub-collection. In addition to those numbers, very old oval labels with numbers are associated with some specimens in the collection. These likely correspond to the private numeration system of Gustav SchÜBler (1787–1834) (Hölder 1977), a professor who was associated with the natural history collections (‘Naturalienkabinett’) in Tübingen from 1817 until his death (Werneburg 2021; Loose 2022).

Fig. 1.

Localities and stratigraphy data for the ichthyosaurs from the Posidonienschiefer Formation housed in the Paläontologische Sammlung Tübingen. A – Historical emblem of the collection with the collection's acronym. B – Portrait of Friedrich v. Huene as a young researcher. C – Map of Germany, with the localities around Holzmaden. D – Detail from C with a geological map and locations mentioned in the article. E – Drawing of an ichthyosaur by v. Huene (1922a), GPIT-PV-30017 (see Pl. 3A). F – Historical phylogenetic tree of Posidonia Shale ichthyosaurs by v. Huene (1931), with correspondence to G. G – Section of the Holzmaden region, as drawn by Hauff (1921). H – Tentative correlation with the well-described Posidonia Shale biostratigraphy of Dotternhausen (SChmidröHl et al. 2002).


There was a certain inconsequence in the use of all these numeration systems by scientists introducing several (understandable) confusions. Moreover, some researchers (e.g., Hofmann 1958) introduced their own “numbering system” for these ichthyosaur specimens, which added to the already existing problems. We compiled the available information – including changing taxonomy – and chronologically list, describe and illustrate (Pls. 1–19) every single specimen separately. To avoid confusion in the future, we recently introduced a new cataloging system for the Tübingen collection uniting previous acronyms as ‘GPIT-PV-…’ and starting with 30000 (Werneburg et al. 2021), which is a number never reached by any previous numeration system before. Ichthyosaurs described herein range from GPIT-PV-30009 to GPIT-PV-30083, plus GPIT-PV-30796, GPIT-PV-60576, and GPIT-PV-60577.

2.4. Length measurements

In many cases, it is quite difficult to conduct good measurements directly on the objects. Measurements on photographs are much better but still smaller errors occur by perspective distortion. Historically, due to restorations or rearrangements of objects in the exhibitions, access to specimens in different conditions to measure the specimens existed through time. In many publications, the length of the object is provided but without a clear indication of how the lengths were measured. Moreover, enormous differences emerge when lengths are reconstructed: How important is diagenetic distortion? How exactly can total length be measured when single vertebrae are shifted? What is the influence of the changing number of documented vertebrae (particularly in the tail) in total length estimation? How to traceably provide the angle size at the tail kink? Hence, large differences between measurements can be expected among researchers over time and it is hard to interpret or reconstruct what the respective authors could have meant with the measurements they provided. In our study, we provide a traceable and simple measure. Basically, it is the total length of the actual fossil material in its taphonomic condition. We use our total measurements just to compare with the literature, and provide the scale bar on the photos to prevent future confusion. We provide some examples to illustrate the confusion:

Specimen GPIT-PV-30017 shows an almost perfect skin preservation and good general preservation. Accurate measurement is possible. Hence, in the literature very similar measurements are provided (Hölder 1964: 116 cm, Hungerbühler 1989: 115 cm, Kröner 1999: 117 cm). One exception is v. Huene (1922: pl. 21, fig. 4). The provided length of 112 cm is hard to trace and might represent a reconstructed length.

v. Huene (1931: 361) provided a total length of 320 cm for GPIT-PV-30020, but the total length of the exhibit only measures about 302 cm. Maybe this was due to a typo.

For GPIT-PV-30021, Quenstedt (1858: 220) provided a length of eleven “Fuß”. In most cases, “Pariser Fuß” (Parisian foot) was meant (Hungerbühler 1994: 248). However, given it could mean “Württemberger Fuß”, it would mean a total of 315 cm. When measuring along the vertebral column (Mcgowan 1973: 5, fig. 2B), one receives a length of 290 cm. Exactly that length was used for categorization by Quenstedt (1858: 219: “9½ Fuß”). When adding the distal-most tip of the tail, which was clearly artificially added to the fossil (Pl. 3D), one would receive 310 cm at best. Hence, it is likely that Quenstedt (1858: pl. 26, fig. 12) provided a very generous measure (see also >Maisch 2008: 232: “No. 63, 314 cm”). In addition, it is possible that at the time of measurement the tail tip was still fully covered, but we cannot proof this today.

Finally, v. Huene (1951: 284) provided a length of 520 cm for the wooden frame of GPIT-PV-30024. We measured a plate length of 514 cm and, including the frame, it is 530 cm. Either v. Huene measured without this frame or a new frame was added in the meantime.

3. Results

3.1. Overview of specimens

In total, we identified and illustrated 78 articulated or anatomically associated ichthyosaur specimens from the Posidonienschiefer Formation in the Tübingen collection (Chapter 3.3, Pls. 1–21), including 73 postnatal specimens (juveniles and adults). Three females are associated with four embryos in total (sidenote: actually, these are well-advanced fetuses with – by definition – basically all organs developed, but we keep the terminology “embryo” as it was used only as such in the literature); one additional, isolated embryo is preserved in our collection (Fraas 1891: pl. 6, fig. 3). Two specimens illustrated before cannot currently be located (Quenstedt 1856–1857, pl. 26, fig. 7; Hofmann 1958, pl. 8, fig. 6) and are not illustrated herein. Few further specimens reported in the literature might still exist in our enormous and still largely uncatalogued palaeontological collection, but could not be (re-)discovered based on the limited information available at this time.

Fig. 2.

List of articulated Tübingen ichthyosaur material with known origin and stratigraphical allocation discussed in this study, ordered by the stratigraphy used by Hauff (1921); compare to Fig. 1G, H.


By referring to >Maisch (2008), Maxwell (2012) listed three type specimens, GPIT-PV-30028 being the lectotype of Stenopterygius quadriscissus (Quenstedt, 1856), GPIT-PV-30040, the holotype of S. triscissus (Quenstedt, 1856), and GPIT-PV-30032, the neotype of S. uniter (v. Huene, 1931). These are, in fact, the type specimens of all currently accepted species of Stenopterygius, the most common genus of ichthyosaur from the Posidonienschiefer Formation (>Maisch 2008; Maxwell 2012). Based on the skeleton described as Stenopterygius hauffianus forma typica by v. Huene (1931), Maisch (2008) erected a new genus, Hauffiopteryx, with GPIT-PV-30041 being the lectotype of Hauffiopteryx typicus v. Huene, 1931 (Maxwell & COrtÉs 2020). In addition, there are several type specimens of taxa that are currently not accepted as valid species: GPIT-PV-30065 is the lectotype of Stenopterygius hauffianus v. Huene, 1922 (v. Huene 1922a; Mcgowan 1979), GPIT-PV-30019 is the holotype of Ichthyosaurus longipes V. Wurstemberger, 1876 (Mcgowan 1979), and GPIT-PV-30061 is a syntype of Stenopterygius promegacephalus v. Huene, 1949 (Hungerbühler 1994). Several specimens were described by Quenstedt (1851–1852, 1856–1857, 1867, 1885) and belong to the type series of S. quadriscissus. They were later listed as paralectotypes of S. quadriscissus by Hungerbühler (1994) and include GPIT-PV-30021, GPIT-PV-30054, GPIT-PV-30033, GPIT-PV-30037, GPIT-PV-30060 and GPIT-PV-30016.

A number of specimens are exceptionally well-preserved and may permit detailed anatomical observation beyond previous accounts (v. Huene 1922a, 1926, 1930, 1931; Keller 1976; Caldwell 1997; Maxwell & CortÉS 2020). These specimens include, among others, GPIT-PV-30017, -30020, -30027, -30042, -30048, -30049, -30050, -30063, -30066, -30070, -30075 and -30081.

v. Huene (1922a, 1931, 1949a) recognized different obvious “variations” that might deserve further observations: GPIT-PV-30045 was called Leptopterygius integer Mutatio dissidens and is now known as Suevoleviathan integer (Maisch 1998b; Maisch 2010), GPIT-PV-30051 was called Stenopterygius crassicostatus Mutatio antecedens, and GPIT-PV-30014 was called Stenopterygius megalorhinus Mutatio postera. In several specimens of the Tübingen collection, obvious pathologies were detected, including GPIT-PV-30047 (Hennig 1923; v. Huene 1931), GPIT-PV-30024 (v. Huene 1951; Hofmann 1958), as well as GPIT-PV-44062 (syn. GPIT/RE/07096-431; this isolated element is not further discussed in this paper) (Pardo-PÉrez et al. 2018a), GPIT-PV-30031 and GPIT-PV-30035 (Pardo-PÉrez et al. 2018b), and GPIT-PV-30018 (Pardo-PÉrez et al. 2019).

To our knowledge, the following specimens were not published before and are (except for GPIT-PV-30058) currently not on display. They include four specimens of Stenopterygius quadriscissus: GPIT-PV-30027 (dorsoventrally embedded), GPIT-PV-30030 (without tail), GPIT-PV-30058 (without snout), and GPIT-PV-30070 (with fenestrated coracoids). Three specimens are labelled as Stenopterygius sp.: GPIT-PV-30071 (dorsoventrally embedded, not prepared), GPIT-PV-30073 (disarticulated skull, not prepared), and GPIT-PV-30079 (disarticulated skull, with incision “HAUFF 1904”). Furthermore, one Temnodontosaurus sp. (GPIT-PV-30076, forelimb?, labeled as “multiscissus”) and one “Ichthyosaurus sp.” (GPIT-PV-30080, badly preserved skull, SCHÜBLER-no.: 13550; the genus Ichthyosaurus does, of course, not occur in the Posidonienschiefer-Formation, but the original designation is used here because the badly preserved specimen cannot be more clearly determined at present) were not published before.

A series of five specimens need to be mentioned, because they illustrate how important it is to accurately check the (generally extraordinary) preparations of BERNHARD HAUFF to identify which elements of the specimens actually belong to the specimen in question or not [GPIT-PV-30063 and GPIT-PV-30022: clearly prepared by HAUFF; GPIT-PV-30054 and GPIT-PV-30019: unclear – see Koken (1904: 679): “the old specimens of the Holzmaden Shale were newly prepared with large costs”]. The specimen bought from Mr. FLECK, GPIT-PV-30034, was restored by HAUFF in 1901 (v. Huene 1922a): several elements can be clearly assigned to the main specimen; however, the assignment of single elements is doubtful regarding taxonomic and even anatomical identity. In some cases, the fossil material was damaged during the preparation process or was completely removed. A typical method applied at the time was an intarsia-based process. Synthetic resin interstices (i.e., “Kitt aus Magnesit und Wasserglas, künstlich schwarz gefärbt”; letter from 20.12.1927; GA 80/272, letter 42) closely align the fossil material and veneer the insertion of sediment plates that surround the original specimen. In crucial cases, even parts of bones do not belong to the original specimen and colorations of the sediments whitewash the differences. This situation is particularly problematic when type material is described without detailed preparatory documentation (i.e., GPIT-PV-30019).

3.2. Historical studies on Tübingen Posidonia Shale ichthyosaurs: an overview

In the following, we provide a survey of the major publications dealing with Tübingen Posidonia Shale ichthyosaurs to exemplarily illustrate the diverse challenges posed by these studies when attempting to identify museum specimens properly. In this regard, we also analyze the unique research agenda of each author, which doubtlessly influenced the design of the respective studies and thus the approach to specimen documentation.

3.2.1. Quenstedt's and Fraas' initial studies

FRiedrich August Quenstedt never published a separate study on ichthyosaurs but he described and illustrated several specimens in his monographs on the palaeodiversity of the Swabian Alb. The first comprehensive treatment of “ichthyosaurs of South German Triassic and Jurassic sediments” was published by Fraas (1891).

3.2.2. v. Huene's broad-scale analyses

The study of v. Huene (1922a) on ‘The ichthyosaurs of the Lias and their relationships’ is certainly the most important one to understand the basics for the identification of genera and species erected by this author. Some specimens were documented by photographs, although many more specimens were available in Tübingen at the time. Moreover, a great number of isolated vertebrae and teeth were noticed in the text, but rarely reproduced by drawings. Some descriptions in the text of v. Huene (1922a) were not traceable for us, because it was not clear whether the specimens were actually present in Tübingen or whether they were just added “out of memory” to enable a consistent text flow. Two smaller skulls of Leptopterygius acutirostris were recorded. Although we could locate only one of them clearly (GPIT-PV-30038; Fraas 1891: pl. 12, fig. 2), it is quite possible that GPIT-PV-30026 was cited as the second (see Koken 1905a: 66).

Fig. 3.

Historical drawing of the Tübingen ichthyosaur “Ichthyosaurus tenuirostris” (= Stenopterygius quadriscissus), published by Quenstedt (1852). Combined from the author's pl. 9, fig. 2a + 2b. As in the originally published drawing, the specimen is mirrored (compare to Pls. 2D and 20B; GPIT-PV-30016).


Although seven specimens of Eurhinosaurus longirostris were certainly present at the time (Hauff 1921; v. Huene 1931; incl. GPIT-PV-30012), only five specimens were listed by v. Huene (1922a). We also could only locate four of these with certainty today (GPIT-PV-30010, -30034, -30039, -30062). The putative embryo, namely GPIT-PV-30059, was cited and depicted separately by v. Huene (1922a: pl. 4, fig. 6). One of the E. longirostris specimens (Fraas 1891: pl. 12, fig. 5) was the focus of a morphofunctional discussion, although the specimen is a clear composite of two individuals (Hauff 1921: 34). Two complete skeletons of Stenopterygius zetlandicus were reported by v. Huene (1922a), but only one of them (Seeley 1880: pl. 1, fig. 3) could be located. In a letter to v. Huene (31.05.1922a; GA 80/272, 12; p. 269), Hauff guaranteed that he intended to still sell ‘the’ S. zetlandicus specimen to him. Likely, it was GPIT-PV-30048. This is one of the examples, in which a specimen cannot be traced to its historical origin by the comments in the original publication alone and only archive documents can help to choose the most plausible allocation.

The species S. megacephalus was erected by v. Huene (1922a). Only two specimens were cited (one of them in Amsterdam, The Netherlands, and one in the fossil company Krantz'sche Mineralienhandlung in Bonn, Germany) as “typical old [adult] specimens” of this species. Several younger specimens were said to exist, namely in the Tübingen collection. v. Huene (1922a) clearly listed only one young specimen from Lias ε II, 10. However, neither a clear allocation to the list of v. Huene (1922a) nor to historical photographs of the collection is possible. Apparently, a very large and old specimen (Quenstedt 1856: pl. 26, fig. 12) was later referred to S. megacephalus by v. Huene (1931). For certain ontogenetic considerations, different measures and proportions were noted (V. Huene 1922a: 58) which we cannot trace. As such, the identification of very small specimens in that study remains unclear.

In 1931, Friedrich V. Huene provided the most comprehensive specimen list (Fig. 4) of regional Posidonia Shale specimens (V. Huene 1931; the list was later supplemented in 1974 by the unpublished specimen catalog of Jane Ann Robinson, UAT 578/591, see below). No photographs or drawings were provided, but only their location in the building was noted. Due to a comprehensive, little-documented reorganization of the collection and the whole building in the last century (Werneburg 2021), a clear allocation of the specimens is rarely possible. v. Huene (1922a) already listed many specimens in his comprehensive study before, and in this earlier study, he provided line drawing reconstructions of 20 ichthyosaurs, with five of them clearly belonging to the Tübingen collection: E. longirostris, L. integer, L. acutirostris, S. zetlandicus, and Ophthalmosaurus icenicus (the latter is not from the Posidonia Shale, but from the Lower Oxford Clay of Peterborough, England). Nevertheless, in his ‘images of the Palaeontological Collection Tübingen’, v. Huene (1919) provided a rough visual impression of the exhibition design, but without describing what was visible in the pictures in detail (see also Westphal 1971: 30). In total, v. Huene (1922a) recorded that he had studied 70 specimens of S. quadriscissus. It is not clear, how many of them came from Tübingen, but the author was at least able to reconstruct a cross-section of the pelvic region based on his comprehensive observations (V. Huene 1922b: 281; see also Hennig 1923: 74). In v. Huene (1931), the allocation of specimens is mainly based on the detailed documentation of the stratigraphic layers, in which the specimens were found. It is worth mentioning that also very badly preserved and unprepared material was listed (i.e., GPIT-PV-30012, -30036, -30074), apparently because the stratigraphical allocation of much material for him enabled a better understanding of evolutionary processes.

Fig. 4.

List of specimens in v. Huene (1931: 352) with manual red pencil notations for Tübingen specimens (scan from the collection's library in Tübingen). A) GPIT-PV-30017 (Pl. 3A), B) GPIT-PV-30074 (Pl. 17A), C) GPIT-PV-30036 (Pl. 8A). See Discussion on specimen GPIT-PV-30075.


In their correspondence (GA 80/272), it becomes apparent, how close v. Huene had cooperated with the preparator, amateur paleontologist, and fossil dealer Bernhard Hauff sen. in Holzmaden. The stratigraphical layers of purchased material in Tübingen were often only later (!) allocated to the specimens by Hauff, and also photos of material observed by v. Huene in the Holzmaden collection were only made and sent later to v. Huene. Nevertheless, some aspects that the author listed are difficult to understand. Either some typos exist or some layer information was not correctly updated. The allocation of specimens to the Tübingen collection and the citation of figures appear to be arbitrary and incomplete to some degree.

3.2.3. Hofmann's taphonomy study

Hofmann (1958) discussed taphonomic aspects of fossilization, such as the orientation of the head (abbreviated as KL, “Kopflage”) and the tail (SL, “Schwanzlage”), skin preservation (I, “Hautexemplar”), and disarticulation (V, “stark zerfallenes Exemplar”). One specimen of S. quadriscissus (his “KL III-6”, Lias ε II, 3) was listed as the only skeleton with the special preservation “belly upward” (“Skelett in Bauchlage”). A clear identification was not possible, because the putative specimen in question (GPIT-PV-30023) stems from a different layer (Lias ε I, 2). We could not find a second specimen of S. quadriscissus that this author had illustrated but did not describe in his text (his “BL IV-1” on pl. 8, fig. 6, from Lias ε II, 3). Hence, it must be declared as missing.

Two of the three pregnant females in our collection were drawn by Hofmann (1958, pl. 3: fig. 6, and pl. 4: fig. 5). In that way, the identity of “KL IV-5” (GPIT-PV-30051) and of “SL IV-4” (GPIT-PV-30054) could be clarified. Specimen “SL IV-1” from Lias ε II, 3/4, with two embryos, was not illustrated but might be the lectotype of S. quadriscissus (GPIT-PV-30028), although only one embryo is preserved.

3.2.4. Robinson's inventory list

The unpublished inventory list from 1974 (UAT 578/591) made by JANE ANN Robinson (she was working on plesiosaur and ichthyosaur locomotion at the institute for some time; pers. comm. FRANK Westphal 2020; see also Krahl et al. 2022), summarized all ichthyosaur material, which was on display in the exhibition in the early 1970s. The notations of the cabinets used at this time are not traceable for us. In most cases, specimens and cabinets changed their position in the building. As there are no illustrations in this inventory list, we had to rely on the brief descriptions to identify the specimens (e.g., “a perfectly preserved, well prepared, complete ichthyosaur skeleton with skin preservation”; GPIT-PV-30017). For some aspects, Robinson cited older literature with published illustrations. Unfortunately, several errors appeared: some clear citations were allocated to different objects. Some well-known previously published material was insufficiently described. Based on those discrepancies, it was difficult to locate five specimens listed as “Stenopterygius megacephalus” (“No. 92, 93, 94, 95, 96”), even though the taxonomic attribution was not up-to-date. Most likely “No. 92” was (within the framework of v. Huene's taxonomy) correctly determined as S. megacephalus and could represent specimen GPIT-PV-30016. “No. 93-96” was reported to be from the Holzmaden locality, whereas on all historical photographs available to us (UAT 678/592) Ohmden is always provided as the locality for S. megacephalus specimens. “No. 94 and No. 95” were assigned to Lias ε II, 6. Described as a particularly large object, “No. 94” can be associated with GPIT-PV-30021, whereas the ‘disarticulated tail tip’ of “No. 95” lets us assign the specimen to GPIT-PV-30015. Based on the method of elimination, it is most plausible that two specimen labels were mixed, namely those of GPIT-PV-30018 (“No. 96”) and GPIT-PV-30014 (“No. 99”). Whether the allocation “Lias ε II, 3, from Ohmden” can be assigned to GPIT-PV-30018, is impossible to trace any further. It is most likely that some confusion has happened, because it was not the actual sediment of the specimen itself that was observed, but unfortunately only the surrounding sediment which was secondarily added to the display for aesthetic reasons and to fill up missing parts. The particularly decorative and resistant Fleins (Lias ε II, 3) from the upper Tenuicostatum Zone was traditionally and is still used for such purposes. The correspondence between v. Huene and Hauff already hints at such a compilation (22.10.1929; GA 80/272, 46, p. 340).

3.2.5. Johnson's and McGowan's taxonomies

Johnson (1979) presented a list of ichthyosaurs referred to Stenopterygius of the Tübingen collection. The taxonomic assignment is likely based on the inventory list of Robinson (1974: UAT 578/591). Several uncertainties appeared when we tried to identify the specimens. The collection number “GPIT 7800” cannot be confirmed. However, a well-preserved specimen from the Stuttgart collection, SMNS 7800, exists (Erin Maxwell, pers. comm. 2019). The specimen GPIT-PV-30054 is now referred to S. zetlandicus and was listed by Johnson (1979) as GPIT 1491/6 before. However, exactly this specimen was illustrated as “GPIT Z04” by the same author a few years earlier (Johnson 1977). The inventory of Robinson (1974: UAT 578/591) listed two skeletons as S. megalorhinus, but Johnson (1979) listed three specimens of this species (“GPIT 1491/7”, “GPIT 1491/10”, “GPIT 1491/11”). There is no doubt that specimen GPIT-PV-30063 (“No. 98”, Robinson 1974: UAT 578/591) is identical to “GPIT 1491/7” (Johnson 1979). The illustration of the forelimb (Johnson 1979: fig. 20, p. 77) apparently belongs to GPIT-PV-30032 (“No. 97” of Robinson 1974). The locality is not exactly known, but stratigraphically the specimen belongs to Lias ε II, 10. However, Johnson (1979) allocated it incorrectly to Lias ε II, 11, and named it as “GPIT 1491/11” and not with the correct number “GPIT 1491/10” (Mcgowan 1979). The mirrored image of another forelimb (Johnson 1979, fig. 17a, p. 72) is listed as “GPIT 1491/9”, but would better fit with her “GPIT B5”, because in Robinson (1974, No. 95), GPIT-PV-30015 was clearly described as “GPIT 1491/9”. For the continuous numeration of the old publication acronym, namely “GPIT 1491/…” numbers “GPIT 1491/5” and “GPIT 1491/13” were omitted for unknown reasons. “GPIT 1491/13” belongs to a completely different taxon, Temnodontosaurus trigonodon (GPIT-PV-30035). The specimen “GPIT 1491/5” (i.e., GPIT-PV-30019), which was identified by Mcgowan (1979) as Ichthyosaurus longipes (= S. crassicostatus on the old label of the time), was listed by Johnson (1979), however.

Fig. 5.

Literature, archive, and collection research conducted in this study, exemplified for GPIT-PV-30010 (Pl. 1B). Old labels, photographs, inventory lists, and publications were analyzed and compared.


In the study of Mcgowan (1979), two inventory numbers were provided that we cannot allocate clearly: “GPIT 1491/11” and “GPIT 1491/14”. “GPIT 1491/11” is called S. megalorhinus, Lias ε II, 6, from Holzmaden. Johnson (1979) allocated it to Lias ε II, 10, and Hofmann (1958) (likely as “BL V-1”) to Lias ε II, 3. Perhaps all authors refer to GPIT-PV-30018. Because the original label is lost, we cannot provide a final identification.

“GPIT 1491/14” was described by Mcgowan (1979) as a “partial skeleton with axial skeleton complete just beyond the pelvic region” and he referred the specimen to S. megacephalus, without mentioning the stratigraphic provenance. Johnson (1979) called it a S. quadriscissus from the Holzmaden locality. Consequently, the allocation of the specimen (“No. 110”) of Robinson (1974) appears correct. Our own research in the collection did not result in the identification of any specimen that would fit with the description of Mcgowan (1979). As the taxonomic determination differs between the cited publications, incomplete or bad preservation of the skull is likely. The only possible identification could be GPIT-PV-30025, because this very large object was basically unprepared until 2009 and was placed in front of the institute's old lecture hall on the lower floor (“Erdgeschoß”).

3.2.6. Hungerbühler's and Maisch's Stenopterygius taxonomies

Hungerbühler (1994) discussed the idiosyncratic taxonomic nomenclature introduced by Friedrich August Quenstedt in detail. Several descriptions and illustrations were presented for historical specimens. Moreover, all taxonomic synonyms were listed for the specimens and references to older literature are fairly complete. However, Hungerbühler (1994) limited his study to the historical type material of Stenopterygius. We were not able to identify one specimen (Quenstedt 1856–1857, pl. 26, fig. 7), which has to be declared as missing. Some specimens listed by v. Huene (1922a, 1931) were not considered by Hungerbühler (1994).

A complete treatment of Stenopterygius was presented by Maisch (2008), who discussed more recent publications (Mcgowan 1979; Godefroit 1994; Hungerbühler 1994) and also considered old collection numbers. Maisch (2008: 254) concluded: “It is unsatisfying that, currently, juvenile specimens cannot be determined with confidence”. This quote highlights the situation that the juvenile specimens of Tübingen collection were basically not considered in most publications with the exception that they were taxonomically still relevant for older authors.

The only minor mistake of Maisch (2008) is a double numeration of “GPIT 1491/10”. Both specimens in question (“No. 14”, “No. 44”) were allocated to S. megalorhinus v. Huene 1922a (Nr. 14: v. Huene 1931: pl. 2, fig. 4; Nr. 44: Mcgowan 1979: fig. 6). The stratigraphical allocation permits identification of “No. 14” to GPIT-PV-30032. The identity of “No. 44” remains unclear, although an allocation to GPIT-PV-30037 is most likely. The reason for this confusion is a reference in the text, in which S. triscissus (Quenstedt 1856) is called “No. 44”. After consulting the author (Michael Maisch, pers. comm. 2019), it could be GPIT-PV-30042 (specimen with skin preservation); however, the length provided (189.5 cm) does not fit any of these reported specimens.

3.3. Identification of articulated Posidonia Shale ichthyosaur specimens in the Tübingen Paleontology Collection

In the following, all specimens are discussed in detail and sorted by their collection number. In squared brackets, we refer (1) to the illustration in our paper (Pls. 1–19), (2) we mention the stratigraphical layer and (3) the location; then we provide (4) a tentative, in most cases the most current species determination and (5) refer to earlier mentions of the specimens, notable details, and species allocations in the literature; finally, we (6) list synonymous collection numbers. For each specimen, a discussion follows as separate paragraph(s).

Fig. 6.

Historical view of one hall of the Paleontological collection in Tübingen. All specimens are located at different places in the exhibition now, making it difficult to trace historical references to the place where the specimens were on display in the past. During the present study, all ichthyosaur specimens (in the background) of this picture could be identified, and the descriptions of Koken (1905a) fit very well with this figure. Photo from the collection archive (see also Westphal 1971).


GPIT-PV-30009 [Pl. 1A; Lias ε II, 3; Holzmaden; Stenopterygius quadriscissus; Kröner 1999, figs. 48, 49, length: 56 cm; V. Huene 1922a: 56, Nr. 4; Koken (1905a) cited a very small specimen; syn.: PV 17956, GPIT/RE/00208].

This specimen was possibly reported by v. Huene (1931) as ‘an almost embryonical skeleton from Tübingen’ [“fast embryonales Skelett in Tübingen”] (letter of Bernhard Hauff to Friedrich v. Huene on September 14th, 1926; GA 80/272, letter 35).

GPIT-PV-30010 [Pl. 1B; Lias ε II, 6; Ohmden; Eurhinosaurus huenei; Godefroit 1994, Eurhinosaurus longirostris; Mcgowan 1979: 116, GPIT 439, Eurhinosaurus huenei, Lias ε II, 6, Ohmden; V. Huene 1952: 55, figs. 12, 13, Lias ε II, 4 + Lias ε III, Holzmaden; V. HUENe 1931: 374, see Discussion; V. Huene 1922a, pl. 5, fig. 3-5, pl. 22, fig. 17, Eurhinosaurus longirostris; Koken 1905a: 66, skull and hind limb of a big animal (“Schädel und Hinterflosse eines großen Tieres”); syn.: GPIT/RE/01605, GPIT-PV-30010-1 = forelimb (syn. PV 18350, GPIT/RE/01605), GPIT-PV-30010-2 = hind limb (syn. PV 18351, GPIT/RE/01609), GPIT-PV-30010-3 = humerus (syn. PV 18350a, 1910), GPIT-PV-30010-4 = skull (syn. PV 18349, GPIT 439)].

v. Huene (1931) cited a skeletal fragment of Eurhinosaurus longirostris, from Holzmaden, layer Lias ε II, 6, which is clearly recognizable in v. Huene (1922a, pl. 4, figs. 3–5). The author drew reconstructions several times (1922a: pl. 22, fig. 17; 1926: fig. 6) and reported limbs and a pelvis from Tübingen (V. Huene, 1952, i.e. only mentioned in figure captions). Historically, these body parts were stored separately (see Koken 1905a: 66) and received individual collection numbers.

GPIT-PV-30011 [Pl. 1C; Ohmden, Temnodontosaurus trigonodon; Hölder 1964: 112, fig. 127, Leptopterygius sp., forelimb; v. Huene 1931: 370, Lias ε II, 6; V. Huene 1922a, pl. 22, fig. 19 (?), see Discussion; Quenstedt 1885, pl. 15, fig. 11, p. 206, Ichthyosaurus multiscissus, Lias ε, Ohmden; syn.: GPIT/ RE/01606].

Quenstedt (1885) illustrated a forelimb with malformation from Ohmden (pl. 15, fig. 11) and assigned it to Ichthyosaurus multiscissus. v. Huene (1931) reported a forelimb ‘above the cupboard of pterosaurs’ [“über dem Flugsaurierschrank”] from the Holzmaden locality, layer Lias ε II, 6, and assigned it to Leptopterygius acutirostris. A published reconstruction likely refers to this specimen (V. Huene, 1922a: pl. 22, fig. 19), although in the text, GPIT-PV-30035 is cited. In the current exhibition, the specimen is labelled as Eurhinosaurus longirostris.

GPIT-PV-30012 [Pl. 2A; Lias ε II, 6; Ohmden; Eurhinosaurus longirostris; v. Huene 1931 ?, p. 374, see Discussion; syn.: GPIT/RE/03232].

v. Huene (1931) listed 45 specimens that we could locate in Tübingen. Among those, there is one specimen that we could not identify, namely ‘a badly prepared skull in Tübingen (old) [“Schlecht präparierter Schädel in Tübingen (alt)”], Eurhinosaurus longirostris, Lias ε II, 6’. Most likely, however, a fragment of that specimen, from Ohmden, could be on display.

GPIT-PV-30013 [Pl. 1F; Lias ε II, 6; Holzmaden; Eurhinosaurus longirostris; v. Huene 1931: 374, see Discussion chapeer; syn.: GPIT/RE/03382].

This is a relatively small specimen with one isolated vertebra and one isolated ischium. As written on the original label, this specimen was purchased from Hauff and came from Holzmaden locality. v. Huene (1931) listed an ‘isolated pelvis’ [“isoliertes Becken”] from Lias ε II, 6.

GPIT-PV-30014 [Pl. 2B; Lias ε II, 11; Holzmaden; Stenopterygius triscissus; Maisch 2008, pl. 4, fig. 5, Stenopterygius triscissus, Lias ε II, 1 (wrong); Maisch 1998a: 411, Lias ε II, 2 (wrong), declared as missing specimen; Mcgowan 1979, pl. 2, fig. 4, Stenopterygius megalorhinus, Lias ε II, 2 (wrong); v. Huene 1949, fig. 4, Stenopterygius megalorhinus Mutation postera, Lias ε II, 11; syn.: PV 24306, PV 24334 (?), GPIT/ RE/07297].

This specimen was purchased in 1945 and received the number PV 24306, as written on the label. Apparently, it was prepared for two years and entered the museum in 1947 and received a second number: PV 24334. Based on the related note in the Petrefactenverzeichnis, the object originates from Lias ε II, 11, of Holzmaden. v. Huene (1949c) drew a part of a skull that can be easily associated with this specimen and the author labeled it as Stenopterygius megalorhinus Mut. postera. Mcgowan (1979) apparently confused Arabic and Roman digits and associated the specimen to Lias ε II, 2. Maisch (2008) attempted to correct this mistake, however, in the printed version of the article, it is written as Lias ε II, 1. Furthermore, on the current specimen label, it says: Lias ε II, 3.

Mcgowan (1979) showed this specimen in a mirrored view. In that way, confusion with another object, which looks very similar, is possible in superficial view, namely with: v. Huene (1922a), pl. 9, fig. 1, Stenopterygius megalorhinus, Lias ε II, 4, housed in Kremsmünster, Austria, BMK 813b. When mirroring back to the original view, it is possible to read on the historical labeling: Stenopterygius megacephalus, Lias ε II, 3 from Ohmden, which, however, very likely belongs to GPIT-PV-30016. Maisch (1998) wrote that GPIT-PV-30014 was lost. After the renovation of the exhibition in 2007, it reappeared [M. Maisch, pers. comm. 2019], and it was correctly named in Maisch (2008). Likely, the confusion of labels happened during the restoration before 1979. The current taxonomic determination is S. triscissus (Maisch 2008).

GPIT-PV-30015 [Pl. 2C; Lias ε II, 6; Ohmden; Stenopterygius quadriscissus; Mcgowan 1979: 107, Stenopterygius megacephalus, could not locate this specimen; Hofmann 1958, BL II-2, 170 cm; v. Huene 1931: 359, see Discussion; syn.: GPIT/ RE/07298].

This is one of three specimens, which were named Stenopterygius megacephalus by v. Huene (1931). He described it as a ‘small skeleton in Tübingen at the wall of the reptile hall’ [“Kleines Skelett in Tübingen, an der Wand im Reptilsaal”], which is proven by an old photograph (UAT 678/592), where it is correctly labelled as Lias ε II, 6, from the Ohmden locality. By citing v. Huene (1922a) and by listing three type specimens, Mcgowan (1979) wrote: “two mature (figured) and one immature skeletons, all from Holzmaden. Huene noted that the immature specimen was in the Tübingen collection, but I did not find it.” In this context, it is worth considering that there might be an issue of incorrect translation. When v. Huene (1922a) wrote “klein” (German for ‘small’), he not necessarily meant immature. Only thanks to the fact that the original label is still present on the specimen, we were able to identify it. The specimen, which has a length of about 160 cm, requires fundamental restoration and is currently stored in the collection archive.

GPIT-PV-30016 [Pl. 2D, fig. 2; Lias ε II, 3; Ohmden; Stenopterygius quadriscissus; Godefroit 1994, Stenopterygius quadriscissus (syntype); Hungerbühler 1994, syntype, Stenopterygius quadriscissus; Johnson 1979, fig. 17a, wrongly cited as GPIT 1491/9; Keller 1976, Nr. 12, Stenopterygius megacephalus; v. Huene 1931, p. 359, Lias ε II, 3; Quenstedt 1885, pl. 15, fig. 1, p. 202, fig. 65, Lias ε, Ohmden; Quenstedt 1872: 57; Quenstedt 1852, pl. 9, fig. 2a, 2b, Ichthyosaurus tenuirostris; syn.: GPIT 29/9/2; GPIT/RE/07299].

This large juvenile specimen with 157 cm in total length was already depicted by Quenstedt (1852) as Ichthyosaurus tenuirostris. Johnson (1979) concentrated only on an isolated forelimb, shown in mirrored orientation, and mislabelled it as GPIT 1491/9 (new syn. GPIT-PV-30021). v. Huene (1931) named a ‘small skeleton on the wall of the reptile hall’ [“kleines Skelett an der Wand im Reptilsaal”] as S. quadriscissus. It is worth noting that the tip of the tail was most likely replaced (Robinson 1974). In combination with a slightly different position of the forelimb and the tail, it might likely have been restored by Hauff (PV 17875, May 1903). Hungerbühler (1994) noticed that a new restoration took place in 1985 [he incorrectly wrote “1885”; the earliest documented restoration took place around 1900; see Koken 1904: 679]. The label, which was attached to the specimen later on, Stenopterygius megacephalus (sensu v. Huene), is also not correct. The collection number which can be found at this specimen (“GPIT/RE/2799”) is not correct either and represents a transposed number.

GPIT-PV-30017 [Pl. 3A; Lias ε II, 4; Holzmaden; Stenopterygius quadriscissus; Kröner 1999, figs. 35–43, GPIT 1491/7, 117 cm; Hungerbühler 1989, fig. 4, Stenopterygius quadriscissus, 115 cm; Heller 1965, fig. 2, 15 cm above Schieferfleins, Holzmaden; Hölder 1964: 108, figs. 123, 126, Stenopterygius sp., 116 cm length; v. Huene 1931: 352, Stenopterygius zetlandicus, wrong “1922a” citation; Hennig 1923: 46, pl. 3, I. quadriscissus; v. Huene 1922: 56, Nr. 8, pl. 6, fig. 1, pl. 21, fig. 4, Stenopterygius quadriscissus, Hautexemplar (112 cm, wrong length); Hauff 1921: 35, Lias ε II, 4 (close above 3); Koken 1905a: 64: “[…] das schönste Stück […] mit erhaltener Haut [side note: with carving: 1899]”, lower figure on the plate right after p. 64; syn.: GPIT 328/6/1, GPIT 1491/7, GPIT/RE/07300, GPIT/ RE/09419].

This specimen, currently assigned to Stenopterygius quadriscissus, shows the best soft tissue preservation of any ichthyosaur in the Tübingen collection (Hungerbühler 1989, fig. 4). Its total length is 117.5 cm (Kröner 1999, fig. 35, 117 cm). It was listed by Koken (1905a: 64) as the ‘by far most beautiful specimen […] with preserved skin’ (“Bei weitem das schönste Stück […] mit erhaltener Haut”). It is a juvenile and stems from Lias ε, Holzmaden. Heller (1965) published a good photograph and went into stratigraphical detail: “15 cm above [über] Schieferfleins”, which approximately corresponds to the lower part of Lias ε II, 4 (Figs. 1, 2). Bernhard Hauff sen. reported exactly this detail (Hauff 1921: 35) and signed the specimen as prepared in 1899 in his typical manner with a carving (Koken 1905a: 12: “[…] ist von B. Hauff präpari[e]rt und wurde 1899 angekauft“). v. Huene (1922a) only showed a small part of the skeleton and measured a length of 112 cm, but noticed a length of 118 cm in the text. In addition, a reconstruction in this study refers to this specimen (his pl. 21, fig. 4). v. Huene (1931) incorrectly cited himself referring to the object as S. zetlandicus from the Lias ε II, 11, and with figure number 11 [pl. 6, V. Huene 1922a) instead of figure number 1. He measured a length of 115 cm, Hölder (1964: 108, fig. 123) measured a length of 116 cm, and Robinson (1974; No. 101) listed a length of only ‘less than 1 m’.

Fig. 7.

Display of several juvenile ichthyosaurs (“Jungtierwand”), which was erected at the occasion of the 90th birthday of Friedrich v. Huene in 1965. A – Institute's director ADOLF SEILACHER (1925–2014) arranges the specimens. BFriedrich V. Huene (left) and the former institute's director EDWIN Hennig (1882–1977) (right) discuss in front of the ichthyosaur wall. Photos from the institute's archive.


GPIT-PV-30018 [Pl. 3B; Lias ε II, 3 ?; Holzmaden ?; Stenopterygius quadriscissus; Pardo-PÉrez 2019, fig. 1b, Stenopterygius quadriscissus; Mcgowan 1979, GPIT 1491/11 (?); Hofmann 1958, fig. 19d, Stenopterygius quadriscissus (“Unterkiefer geöffnet”); syn.: GPIT/RE/07301, GPIT 1491/11 (?)].

The label attached to this specimen definitely belongs to specimen GPIT-PV-30014. Therefore, it is no longer possible to trace the fate of the correct label. The GPIT-number was written on this old label in recent times. Maisch (pers. comm. 2019) clearly precluded confusion with Stenopterygius zetlandicus by v. Huene. Hofmann (1958) listed a specimen as “BL V-1, Lias ε II, 3” (which means a combination of embedding type and stratigraphic layer) as Stenopterygius quadriscissus “with opened lower jaw” [“mit geöffnetem Unterkiefer”], his fig. 19d. Actually, there is a specimen in the collection, which perfectly fits with this drawing. However, it is not clear whether the stratigraphic notation refers to the specimen or to the sediment plates, in which the specimen was arranged for presentation. As no other publications seem to exist on this specimen (total length of 195 cm), also matching to the Petrefactenverzeichnis was not successful as almost only short notes, such as “ichthyosaur from ‘xyz’ locality”, are present in this inventory list. Most likely, the specimen is referred to as GPIT 1491/11 by Mcgowan (1979), because it is embedded in a historical frame and was certainly not restored in recent times. Maisch (1998a) reported two different specimens (GPIT 1491/10 from Lias ε II, 2 and GPIT 1491/11 from Lias ε II, 10). To distinguish between these two, he referred to exactly the same individuum, which is for sure GPIT-PV-30032 (Maisch 1998a: 411). Based on the writing style and the preservation, the label was likely attached to the specimen in the 1970s or earlier. A plausible allocation to the inventory list of Robinson (1974) is Stenopterygius megacephalus, Lias ε II, 3, from Holzmaden.

In a letter from Bernhard Hauff sen. to v. Huene dated July 14th, 1930 (GA 80/272, letter 65), a Stenopterygius megacephalus specimen with a length of 175 cm was cited as ‘number 54’. However, no further reference is given to this number. As such, it is not clear whether this specimen was actually purchased by the Tübingen collection. Moreover, the provided length cannot be attributed to a single specimen of the Tübingen collection with certainty.

GPIT-PV-30019 [Pl. 4A; Lias ε II, 11; Holzmaden; Stenopterygius uniter; Maisch 2008, Stenopterygius uniter + Stenopterygius quadriscissus (Komposit); Kröner 1999, fig. 1, “Extremitäten ergänzt” (extremities added); Maisch 1998a: 412, 417, Stenopterygius quadriscissus (skull, thorax) + Stenopterygius megalorhinus (forefins), GPIT 1491/5; Mcgowan 1979: 111, Stenopterygius longipes (holotype); Hofmann 1958, KL IV-2; v. Huene 1931: 364, Stenopterygius crassicostatus; Koken 1905a: 66, Ichthyosaurus crassicostatus; Fraas 1891, pl. 14, fig. 8, Ichthyosaurus crassicostatus; V. Wurstemberger 1876, Ichthyosaurus longipes; syn.: GPIT 1491/5; GPIT no. 83; GPIT/ RE/07323].

This is the uppermost of the three skeletons mounted above each other in the building's stairways towards Sigwartstraße today. The specimen arrangement was erected in 1966 and was established with the help of the local mining company Fischer to create a uniting framing within the Posidonia Shale (‘Seegrasschiefer’). The upper specimen was named Stenopterygius crassicostatus by v. Huene (1931) and has a skull protruding from the plate, Lias ε II, 11. [“Skelett mit aus der Platte ragendem Schädel”]. Mcgowan (1979) correctly cited this specimen as the holotype of S. longipes (V. Wurstemberger, 1876). Maisch (1998a, 2008) clarified that it is actually a composite of S. uniter and S. quadriscissus. In addition, two right ulnae were reported (Maisch 1998a: 415), which could not be identified during a restoration in 2013 (Kröner 1999, fig. 1; in this specimen, the limbs were largely supplemented). Bernhard Hauff sen. wrote in 1921: “The skeletons are embedded horizontally, […] Only very rarely the snout is drilled to the depth; in most cases, this only happens in Ichthyosaurus crassicostatus.” [“Die Skelette lie-gen alle horizontal eingebettet, […] Nur außerordentlich selten kommt es vor, daß die Schnauze mehrere Zentimeter in die Tiefe sich einbohrte, meist ist das nur bei Ichthyosaurus crassicostatus der Fall”]. More detailed information to identify the specimen is not available; however, reference is given to Fraas (1891), who illustrated fore- and hind limbs. Allocating those to the specimen is very difficult though. It must have been on display for a long time based on the dissertation of V. Wurstemberger (1876), who reported “a very nice specimen in Tübingen with strange extremities” [“einem sehr schönen Exemplar in Tübingen mit eigenthümlichen Extremitäten”], and he created a new species name: I. longipes. Probably it is reported to be restored in May 1903 by B. Hauff (“PV 17876”).

GPIT-PV-30020 [Pl. 3C; Lias ε II, 10; Holzmaden; Stenopterygius quadriscissus; Maisch 2008, Stenopterygius quadriscissus; Godefroit 1994, Stenopterygius hauffianus; Hungerbühler 1989, fig. 2, Stenopterygius hauffianus; Mcgowan 1979: 105, Lias ε II, 10, Holzmaden; Keller 1976, fig. 10e, Stenopterygius hauffianus, Nr. 5; Hölder 1964: 109, fig. 124, Stenopterygius hauffianus; Hofmann 1958, pl. 5, fig. 2, BL III-2, 317 cm; v. Huene 1931: 361, Stenopterygius hauffianus, 320 cm; V. Huene 1926, pl. 4, fig. 2, Stenopterygius hauffianus; syn.: GPIT 1491/2; GPIT/RE/07324].

This is the middle specimen of three skeletons in the stairways of the institute (see Discussion on GPIT-PV-30019), determined as Stenopterygius quadriscissus by Maisch (2008). v. Huene (1931) cited himself by naming it: “Stenopterygius hauffianus, v. Huene (1926, pl. 4, fig. 2), 320 cm in length”. Most specimens in the collection are much less than three meters in length. This specimen has an actual length of 290 cm. Hofmann (1958) and Keller (1976) provided illustrations, leaving no doubt about the identity of this specimen, which originates from Lias ε II, 10, Holzmaden.

GPIT-PV-30021 [Pl. 3D; Lias ε II, 6; Ohmden; Stenopterygius quadriscissus; Maisch 2008, Stenopterygius quadriscissus, (paralectotype); Godefroit 1994, Stenopterygius quadriscissus (syntype); Hungerbühler 1994, fig. 3, Stenopterygius quadriscissus (syntype); Mcgowan 1979, GPIT 1491/9; Johnson 1977, pl. 4, C, GPIT C06; Keller 1976, Stenopterygius megacephalus, Nr. 13; v. Huene 1931: 359, S. megacephalus, im Schwäbischen Saal; Fraas 1891, table p. 52, column 10; Quenstedt 1856, pl. 26, fig. 12, Ichthyosaurus tenuirostris, 11 Fuß length; syn.: PV 4750, GPIT C 06, GPIT 1491/9, GPIT/RE/07326].

This is, historically, a relatively old specimen (“PV 4750”), which was described by Quenstedt (1856) as Ichthyosaurus tenuirostris (quadriscissus), but the author depicted only the tail tip of the specimen. v. Huene (1931) listed three specimens of Stenopterygius megacephalus, all from Lias ε II, 6. By the method of elimination, this specimen can only be the one specimen displayed in the ‘Swabian hall’ (“Skelett in Tübingen im schwäbischen Saal”, v. Huene 1931), particularly because the label is identical to a historical photo. Quenstedt roughly measured the specimen as 11 “feet” [German: Fuß] in length, which would be 357.3 cm [Pariser Fuß] or 315.2 cm [Württemberger Fuß], but the correct measurement is 300 cm. Johnson (1977) only depicted a single humerus, which can be associated to this specimen with certainty. Other notations that the author provided are unclear and only depictions would help a clear identification.

GPIT-PV-30022 [Pl. 4B; Lias ε II, 6; Holzmaden; Suevoleviathan integer; Maisch 2020: 155, fig. 1B, Suevoleviathan integer; Maisch 1998b, Suevoleviathan integer, Holzmaden; Maisch 1997: 2, GPIT 328/4/5, “Ichthyosaurusinteger; Hölder 1964: 110, fig. 125, sclerotic ring; Hofmann 1958, SL V-4, 270 cm; v. Huene 1931: 369, Lias ε II, 6, see Discussion; V. Huene 1922a, pl. 4, fig. 5, Leptopterygius integer; syn.: PV 18542, GPIT 328/4/5, GPIT/RE/07328].

Labelled as “PV 18542”, this specimen was included in the collection as Ichthyosaurus integer. v. Huene (1922a, pl. 4, fig. 5) wrote that this specimen has been included in the collection “shortly before the war” [“kurz vor dem Kriege”; i.e., in spring 1914 – compare also to GPIT-PV-30045, purchased from B. Hauff in 1908, and GPIT-PV-30065, purchased in 1912] as Leptopterygius integer, a skeleton with broken snout (“Skelett mit zerbrochener Schnauze”), Lias ε II, 6, Holzmaden locality. Maisch (1998b) referred to it as Suevoleviathan integer, but did not illustrate it. Maisch (2020) illustrated it and noticed that the broken tail was likely added to the specimen. Some teeth and bones were scraped down to the sediment, most likely for aesthetic reasons. Hauff (1921) was only aware of two ‘integer‘-specimens (compare also to GPIT-PV-30045), one of them located in Tübingen. The stratigraphic horizon was clearly identified to lie between “Unterer Stein” and “Steinplatte”.

GPIT-PV-30023 [Pl. 4C; Lias ε I, 2; Holzmaden; Stenopterygius quadriscissus; v. Huene 1931: 349, Lias ε I, 2, Zell, see Discussion; syn.: GPIT/RE/07334].

This specimen, labelled as Stenopterygius quadriscissus, is embedded in dorsal view and was restored in 2009. Limbs and tail are not articulated and are dispersed. Due to its dorsoventral orientation and its small body size, one can easily confuse it with specimen GPIT-PV-30027. During the restoration in 2009, the sediments were covered with paint, which would confuse it to belong to younger stratigraphic layers, namely to Lias ε II, 4 to 10. Unfortunately, the restoration of the specimen is irreversible, making a clear association with the original sediment complicated. Also, potential marks that could indicate the PV-number are no longer present or visible. However, it is likely that v. Huene (1931) associated this specimen with the Tafelfleins (Lias ε I, 2) from Zell locality near Holzmaden (which means most likely “Zell unter Aichelberg”). As such, this specimen is likely the stratigraphically oldest ichthyosaur from the Posidonia Shale housed in the Tübingen collection (at least with reference to V. Huene 1952).

Fig. 8.

Eurhinosaurus longirostris specimens from Eislingen in their original finding situation. Drawings based on field notes. A – The large specimen GPIT-PV-60576; B – the small specimen GPIT-PV-60577. Color legend: Black = shadow of fossil wood, red = identifiable vertebrae, blue = skull material, yellow = appendicular skeleton, brown = ribs, gray = shale sediment.


GPIT-PV-30024 [Pl. 4D,  Supplementary file (Supplement 1.ply); Lias ε II, 6; Ohmden; Eurhinosaurus huenei; Maisch 2022, Eurhinosaurus huenei; Pardo-PÉrez 2017, table 1, Eurhinosaurus longirostris; Hungerbühler 1989, fig. 9, Eurhinosaurus longirostris, 507 cm; Mcgowan 1979: 116, Eurhinosaurus huenei, Lias ε II, 6, GPIT PJ 24361; Hofmann 1958, pl. 6, fig. 3, SL IV-8; V. Huene 1951, pl. 18, fig. 1, Eurhinosaurus longirostris, Holzmaden; syn.: PV 24361, GPIT 1025/18/1, GPIT/RE/07343, GPIT/RE/09412].

This is a complete skeleton from Ohmden, extensively described by v. Huene (1951), illustrated in his pl. 27, fig. 1, and was referred to Eurhinosaurus longirostris, from Lias ε II, 6. Hofmann (1958) reported a rib fracture with callus formation, as v. Huene (1951: 280) did before. Actually, a pseudarthrosis seems to be present in six of the ribs (compare also to Pardo-PÉrez et. al. 2017: 29, table 1).

GPIT-PV-30025 [Pl. 5A; Lias ε II, 3; Holzmaden; Stenopterygius quadriscissus; Mcgowan 1979, GPIT 1491/14 (?); v. Huene 1931: 349, Lias ε II, 3, see Discussion; syn.: PV 17952, GPIT/RE/07344].

This specimen is referred to as Stenopterygius quadriscissus from Holzmaden locality. It was restored for a permanent loan to the Natural History Museum in Luxembourg in 2014, and since then it is on display there. Due to the intense new preparation, the surrounding matrix likely belongs to the “Schieferfleins” (Lias ε II, 3). The historical preparation of this specimen is completely different from that of specimens prepared with the method of Hauff (1921). Only based on that observation, it becomes obvious that v. Huene (1931) named this specimen a ‘three-meter-long skeleton (Hildebrandt) from Lias ε II, 3’. “Hildebrandt” apparently is the person who sold the specimen and certainly refers to Jacob Hildenbrand, who was a younger friend of Friedrich August Quenstedt and who collected several local fossils for the Tübingen collection (UAT 236/59; see also BÖttcher 1990; Fraas 1891; Hölder 1977; Werneburg 2021).

GPIT-PV-30026 [Pl. 5B; Lias ε; unknown locality; Temnodontosaurus trigonodon; Maisch 1998a: 422, GPIT (PV) 8370; V. Huene 1922a: 25; Koken 1905a: 66, “entblößte Unterseite” (exposed ventral side); syn.: GPIT (PV) 8370, GPIT/RE/09269].

v. Huene (1922a: 25) reported four specimens of Temnodontosaurus: first, a large skeleton from Schlierbach (GPIT-PV-30035) and, second, a large skull from Frittlingen (GPIT-PV-30031). We could probably identify a third specimen, which is a smaller skull from Frittlingen (GPIT-PV-30038) and was already reported by Quenstedt (1858) and later illustrated by Fraas (1891).

The fourth specimen, GPIT-PV-30026, was already reported by Koken (1905a: 66), who wrote ‘Another [Ichthyosaurus ingens] with exposed ventral side [hangs] at the southern wall of the same room’ (“Ein anderer [Ichthyosaurus ingens] mit entblößter Unterseite [hängt] an der Südwand desselben Raumes”]. An old photo of the exhibition (Westphal 1971, fig. 1), which likely stems from that time, actually shows the specimen on the southern wall, directly above GPIT-PV-30038.

In the “Petrefactenverzeichnis” (“PV 18800”), another specimen of I. trigonodon is listed. The institute purchased it from the Krantz Company (Bonn, Germany) on May 12th, 1919. Most likely, it stems from another locality and could not fit into the list of v. Huene's four Temnodontosaurus-specimens.

Maisch (1998a) used the number “PV 8370” which is surely not correctly cited from the PV-number given by Quenstedt, but the description fits very well with GPIT-PV-30026 (embedded from ventral side).

Both, the locality Ohmden and the layer Lias ε II, 6, given on the label, cannot be confirmed. Probably confusion exists with GPIT-PV-30038.

GPIT-PV-30027 [Pl. 5C; Lias ε II, 6; Holzmaden; Stenopterygius quadriscissus; v. Huene 1931: 359, see Discussion; syn.: PV 6518, GPIT/RE/09282].

During restoration in 2014, a relatively old PV-number was discovered (“PV 6518”). The specimen must have entered the collection way before 1850 and originated from Holzmaden. v. Huene (1931) just noticed it as a small skeleton from the Lias ε II, 6, laying on its back, but he could not clearly define its taxonomic identity. However, then he named it Stenopterygius megacephalus without providing a clear allocation to the Tübingen collection. Robinson (1974) listed it as “Stenopterygius quadriscissus (exposed belly upward), No. 113”.

GPIT-PV-30028[Pl.5D;LiasεII,3/4;Holzmaden;Stenopterygius quadriscissus; Maisch 2008, pl. 1, fig. 1, Stenopterygius quadriscissus, lectotype; Godefroit 1994, Stenopterygius quadriscissus (syntype); Hungerbühler 1994, fig. 2, syntypes, GPIT 43/0219-1; BÖttcher 1990, pl. 2, fig. 3; Mcgowan 1979, GPIT ‘Q13’, not pregnant; Hofmann 1958, SL IV-1, incorrect as SL IV-4; v. Huene 1931 ?, see Discussion; Koken 1905a: 66, Weibchen (female); Fraas 1891, table p. 52, column 2; Seeley 1880, pl. 1, fig. 2; Quenstedt 1867: 157; Quenstedt 1858: 219; syn.: PV 7532 (partim), No. 102, GPIT 43/0219-1, GPIT/ RE/09283-A].

The pregnant specimen is the lectotype of Stenopterygius quadriscissus, the type species of the genus. The PV-number (“PV 7532”) of this specimen refers to an ichthyosaur with an embryo in its body. Quenstedt (1858: 219) wrote that this cannot be an embryo, but more likely the prey in the pharynx. Mcgowan (1979) shared the opinion of Quenstedt. One drawing by Seeley (1880, pl. 1, fig. 2) nicely proves the identity of the specimen. Hungerbühler (1994) studied the notes of Quenstedt in detail to clearly identify this specimen including one embryo and called it “Qu 1858: 219”. It is worth noting that v. Huene might have largely ignored this specimen. Only the vague and not even unequivocal notation ‘large skeleton’ (“großes Skelett”) in v. Huene (1931) probably refers to this specimen.

There are three mothers with preserved embryos in Tübingen. BÖttcher (1990: 8) wrote: “Eventually, there is a hint to a second skeleton with two embryos in Tübingen, although – based on the information from Mr. Prof. [Dr. Frank] Westphal – only one such skeleton is present there (GPIT 1491/1; v. Huene 1931: 365; Hofmann 1958: pp. 22, 38, 48). The first indication can be found in v. Huene (1931: 351), the second one in Hofmann (1958: 26, 38, 50). The description of Hofmann (1958) fits relatively well to the Tübingen skeleton No. “Qu. 1858: 21”, except for the fact that the skeleton only contains one embryo; curiously, v. Huene made the same mistake. Like Hofmann (1958), he identified the ichthyosaur as Stenopterygius quadricissus from layer εII3 (Hofmann: εII3/4). It was not possible to identify the reason for this mistake.” Nevertheless, Keller (1976, fig. 9) referred to Hofmann (1958, pl. 4b, fig. 5), and one cannot exclude the possibility that, for the second specimen (with two embryos), v. Huene (1931) referred to “GPIT 1491/6” (syn.: GPIT-PV-30054) and not to “Qu.1858: 219” (syn.: GPIT-PV-30028). The figure by Seeley (1880: pl. 1, fig. 2) clearly shows that at this time no fossil material was visible that could hint at a second embryo. In the contrast, the preservation of GPIT-PV-30054 might lead to confusion about negative imprints of the gastralia with a purported second embryo. Moreover, the formulation by BÖttcher (1990) on the chronology is a bit unclear. It is more likely that v. Huene initially made a “mistake” with the second embryo, which might be based on the mass of material that he had previously studied. Of personal respect for v. Huene, who was still active at this time, Hofmann (1958) might have just taken v. Huene's allocation for granted without further checking. This is also obvious as the author correctly described the mother (GPIT-PV-30054) and correctly described and depicted only one embryo inside (Hofmann 1958: pl. 4, fig. 5). Moreover, we cannot clarify what the exact mistake of v. Huene was, either the allocation of the horizon and the taxonomic identification, “just” the incorrect counting of embryos, or the complete confusion of two specimens plus a typo in the writing process. Our investigation revealed the following conclusions:

When compared to the historical drawing of Seeley (1880), no significant material loss can be identified in the fossil (only some vertebrae in the tip of the tail and a few fragments of some rips).

The old number “PV 7532” can be dated to a time significantly earlier than 1855 because “PV 9533” was published in that year (Quenstedt 1855). This is important to identify other objects (e.g., “PV 9775” = GPIT-PV-30054), because the PV-numbers usually provide no dates. “PV 7532” is obviously listed as the only pregnant female in the collection in 1853 (KLÜPFEL 1853).

The stratigraphic layer can be identified as Lias ε II, 3/4. But the only indication for confirming the location and stratigraphy is No. 102 from Robinson (1974).

In a letter of B. Hauff to v. Huene (GA 80/272, letter 46; 22. Okt. 1929) it is written: “It is absolutely clear that specimen No. 112 […] stems from II, 3” [“Es ist ganz sicher, daß St. Nr. 112 mit den kleinen Embryonen aus II, 3 stammt”]. We never found a published reference to specimen number 112; also, it is possible that this specimen [no. 112] does not belong to the collection of Tübingen, because Quenstedt (1852) already knew about this pregnant specimen [“PV 7532”] in 1852. Moreover, whereas other numbers of the inventory of B. Hauff (Hauff'sches Album) (e.g., Nr. 6, Nr. 54) are cited by v. Huene (1931), the author did not report a No. 112. The only allocation left is a S. quadriscissus from the border of Lias ε II, 3/4, with two embryos [“großes Skelett im stratigraphischen Saal”] ( v. Huene 1931: 351). Most likely, it was confused with GPIT-PV-30028, because only a few further specimens with embryos were listed.

Restoration of the pregnant specimens by B. Hauff (see Koken 1904) can be excluded, because Seeley (1880) already drew the specimens in the same condition as they are visible today.

GPIT-PV-30029 [Pl. 19A; Lias ε II, 3/4; Holzmaden; Stenopterygius quadriscissus; Lomax & Massare 2012: 271; Fraas 1891, table on p. 52; syn.: PV 7532 (partim), GPIT/RE/09283-B]. For discussion see GPIT-PV-30028.

GPIT-PV-30030 [Pl. 6A; unclear stratigraphy and location; Stenopterygius quadriscissus; syn.: No. 111; GPIT/RE/09314].

GPIT-PV-30031 [Pl. 6B; Lias ε; Frittlingen; Temnodontosaurus trigonodon; Pardo-PÉrez 2018, Temnodontosaurus trigonodon; Maisch 1998a: 419, Temnodontosaurus trigonodon, GPIT PV 5759; v. Huene 1931: 372, Lias ε II, 11, Frittlingen; V. Huene 1922a: 30, table column 14, skull from Frittlingen; Koken 1905b: 11, Ichthyosaurus ingens, 170 cm in length; Fraas 1891, pl. 11, fig. 11, Ichthyosaurus ingens; Quenstedt 1889, Oberer Lias von Boll; Quenstedt 1885, pl. 14, fig. 21, p. 206, Ichthyosaurus trigonodon, Ohmden; syn.: PV 5759, GPIT 80/14/21; GPIT/RE/09395].

This is a very large ichthyosaur skull from Frittlingen (listed as PV 5759). It is not clear whether this specimen was purchased from a dealer called Christoph Fleck, who offered ‘a large ichthyosaur head’ in a letter from April 28th, 1872 (UAT 236/73). v. Huene (1931) allocated it to Leptopterygius trigonodon, Lias ε II, 11, and measured it as 220 cm (V. Huene 1922a: 25). Fraas (1891, pl. 11, fig. 11) named it Ichthyosaurus ingens from Holzmaden and figured only two teeth with reference to Quenstedt (1885, pl. 14, fig. 21). The PV-number that Fraas provided (“PV 5750”) definitely does not belong to any ichthyosaur material, but the length of 1.75 m fits very well the actual size. The length that Quenstedt provided, namely 3¾ feet (“Fuß”), which is about 114 cm, is more or less fitting with GPIT-PV-30038. The non-broken lower jaw fragment (GPIT-PV-30031) has a length of 130 cm, and a total skull length of about 180 cm can be estimated. As such, this specimen is by far the largest ichthyosaur skull in the collection. The estimate of v. Huene (1931), who reported more than two meters, represents an overestimated value.

In his latest publication, short before his death in the age of 80, Quenstedt (1889) reported a length of seven feet [c. 213 cm] for a Liassic skull from Boll with ‘27 teeth as thick as a thumb’. Actually, 28 teeth are visible on the lower jaw and five more teeth likely belong to the upper jaw. The confusion is likely caused by the advanced age of the author. Actually, there was no other comparable specimen in the collection and the author did not refer to his own previous publications (Quenstedt 1858, 1885).

GPIT-PV-30032 [Pl. 7A; Lias ε II, 10; Holzmaden; Stenopterygius uniter; Maisch 2008, pl. 6, fig. 3, fig. 1B, Stenopterygius uniter, neotype; Maisch 1998a: 411, Stenopterygius megalorhinus, GPIT 1491/11 (wrong); Caldwell 1997, fig. 2C (wrong); fig. 2B = GPIT 1491/11 (wrong); Johnson 1979, fig. 20, GPIT 1491/11 (wrong); Mcgowan 1979, text-fig. 6, Stenopterygius megalorhinus; Keller 1976, fig. 2, Stenopterygius uniter, Nr. 10; Hofmann 1958, BL IV-5, 236 cm; v. Huene 1931, pl. 2, fig. 4, Stenopterygius megalorhinus, Nr. 6, 236 cm; syn.: GPIT 1491/10; GPIT/RE/09406].

This is one of three specimens in the collection that were referred to Stenopterygius megalorhinus by v. Huene (1931). Based on the author's illustration it is clear, that the specimen was no longer located in the museum of Holzmaden, but came to Tübingen sometime after the publication. An intensive conversation existed between Hauff and v. Huene (e.g., July 14th, 1930; GA 80/272, letter 65), in which both discuss the prices of specimens. In particular, they reported a S. megalorhinus specimen, No. 6, 236 cm in length, from Lias ε II, 6. This is the exact information published later ( v. Huene 1931: 357) except for the horizon, which is published as Lias ε II, 10. There is no notation in the “Petrefactenverzeichnis” which would fit to any of that information.

Keller (1976) allocated the specimen to S. uniter, Mcgowan (1979) allocated it again to S. megalorhinus, and Maisch (2008) made it the neotype of S. uniter. In the cited literature, an identification of the specimen (syn. “GPIT 1491/10”) and the association to Lias ε II, 10, is not problematic, due to some good illustrations. Anyhow, Johnson (1979), Caldwell (1997), and Maisch (1998a) cited the specimen with the wrong number “GPIT 1491/11”. Maisch (2008: 242) clearly argued that confusion about the horizons is not really possible based on his experiences.

GPIT-PV-30033 [Pl. 7B; Lias ε II, 3; Holzmaden; Stenopterygius quadriscissus; Maisch 2008, Stenopterygius quadriscissus; Godefroit 1994, Stenopterygius quadriscissus (syntype); Hungerbühler 1994, fig. 5, syntype; Mcgowan 1979, Stenopterygius quadriscissus, Lias ε II, 3; Keller 1976, Stenopterygius eos; Hofmann 1958, BL V-3, “very old discovery“; syn.: GPIT 1491/8, GPIT/RE/09407].

Hungerbühler (1994) identified the following specimens of Stenopterygius quadriscissus as parts of the syntype series of Quenstedt (1856): GPIT-PV-30028, -30054, -30021, as well as this fourth specimen, -30033. Whether this allocation is based on a quotation from McGowan (1979) or on Hungerbühler's (1994) own observation cannot be reconstructed, although the association of specimens is reasonable. Hofmann (1958) reported a very old specimen (“ein sehr alter Fund, BL V-3”) and Keller (1976) referred it to Stenopterygius eos (as seen on a label on a historical photo). v. Huene (1931) listed two specimens of S. quadriscissus as large skeletons and located one of them in the institute's stairway. Both specimens certainly belong to the above-mentioned type-series. Based on reorganizations within the building in the past, further details cannot be tracked.

GPIT-PV-30034 [Pl. 7C; Lias ε II, 6; Holzmaden; Eurhinosaurus huenei; Maisch 2022, Eurhinosaurus huenei; Godefroit 1994, Eurhinosaurus longirostris; v. Huene 1931: 374, Eurhinosaurus longirostris; Hennig 1923: 88, pl. 8; V. Huene 1922a, pl. 9, fig. 3, Lias ε II, 6; Hauff, B. (sen.) 1921: 34, composite; V. Huene 1919, pl. 7, not cited, but visible in the background of a picture; Koken 1905a, second plate after p. 64; Fraas 1891, pl. 12, fig. 5, Ichthyosaurus longirostris; syn.: PV 16589, GPIT/RE/09408].

Fraas (1891, pl. 12, fig. 5) illustrated this large skeleton and allocated it to Ichthyosaurus longirostris. Although he cited Fraas (1891), Mcgowan (1979) did not mention this specimen explicitly. v. Huene (1922a, pl. 9, fig. 3) figured the specimen as Eurhinosaurus longirostris, Lias ε II, 6, Holzmaden locality, and it is visible in the background of a photo that he published in v. Huene (1919). v. Huene did not cite his own referral in 1931. He only noted a large skeleton from “Fleck” [“großes Skelett von Holzmaden in Tübingen (Fleck)”]. It is obvious, that this notation refers to the fossil collector Christoph Friedrich Fleck (Kreisarchiv Esslingen; Signatur: OST 7/1, family register, vol. 1, 1797–1863, p. 351) and not to a locality. The posterior part of the tail and likely also the hind limb belong to another specimen (Hauff 1921). On a historical photo (UAT 678/592), the article of Fraas(1891) is cited and obviously was attached to the specimen, but this label disappeared after a restoration in 2009. In the “Petrefactenverzeichnis”, “Ichthyosaurus longirostris” is listed as “PV 16589” and was purchased for 250 Gulden from Holzmaden – it is not clear whether the Hauff Company placed in Holzmaden or the fossil locality is meant. C. Fleck was also excavating in Holzmaden and selling specimens privately as can be deduced from several notes in the “Petrefactenverzeichnis”. Probably it is reported to be restored in May 1903 (“PV 17874”).

GPIT-PV-30035 [Pl. 7D; Lias ε II, 11; Schlierbach; Temnodontosaurus trigonodon; Pardo-Pérez et al. 2018, fig. 8, Temnodontosaurus trigonodon; Maisch 1998a: 422, Temnodontosaurus trigonodon, GPIT 1491/13; Mcgowan 1979, Leptopterygius burgundiae, Göppingen; v. Huene 1931: 371, Leptopterygius trigonodon, Lias ε II, 11; 7.5 m length; Hennig 1923: 46, Ichthyosaurus acutirostris (= ingens); V. Huene 1922a, pl. 22, fig. 19, p. 30, table column 10, Leptopterygius acutirostris, Lias ε II, 11, Schlierbach; Koken 1905a: 66, Ichthyosaurus ingens; Fraas 1891, table on p. 69, Ichthyosaurus acutirostris, Schlierbach; 7.45 m length; Quenstedt 1852, pl. 9, fig. 6; Quenstedt 1885, pl. 14, fig. 16; Ichthyosaurus platyodon (multiscissus), 23 Fuß length; syn.: PV 4308, GPIT/RE/09409 – Pardo-Pérez et al. (2018) combined different numberation systems and reported a pathology as GPIT/RE/1491/13].

This is the largest complete ichthyosaur skeleton in the Tübingen collection, but it is imperfectly preserved, particularly because of the crude and inadequate original preparation. The right forelimb is incomplete. Quenstedt (1852, pl. 9, fig. 6; see notation in Wagner 1851 [sic: publication year before Quenstedt's final publication]) illustrated the right hind limb, and determined the specimen as Ichthyosaurus platyodon (multiscissus), Schlierbach, pictured in a mirrored way. He noted a huge skeleton (23' Par[iser] Fuß) in the text (Quenstedt 1852: 128).

In the current exhibition, the visible limb on display is described as left hind limb. Fraas (1891: 66) referred the specimen to Ichthyosaurus acutirostris from Schlierbach locality with a length of 7.5 m, and v. Huene (1931) referred it to Leptopterygius trigonodon, Lias ε II, 11, but named it still Leptopterygius acutirostris in 1922 (V. Huene 1922a, pl. 22, fig. 19). Mcgowan (1979) cited it as GPIT 1491/13 from Göppingen (the town to which the Schlierbach locality belongs) and referred it to Leptopterygius burgundiae. The current label in the exhibition says Temnodontosaurus trigonodon from Schlierbach near Göppingen. The oldest note in the Petrefactenverzeichnis that we are clearly able to identify refers to an ichthyosaur skeleton “PV 4308”, and lists a length of 23 feet [c. 747 cm].

In the biography of his father, Ernst Quenstedt (UAT 236/59) reported a large ichthyosaur from Pliensbach (near Göppingen) with a length of 22 Württemberg feet [c. 630 cm]. Klüpfel (1853: 40) called it'a beautiful ichthyosaur specimen of 24 P[arisian] feet, from Ohmden' (“ein schönes Exemplar eines Ichthyosaurus von 24 P[ariser] Fuß [c. 780 cm] Länge aus Ohmden”). Finally, Quenstedt (1889) listed it as Ichthyosaurus multiscissus of 23 Parisian feet from the upper Lias of Boll. The latter locality, however, refers to GPIT-PV-30031 and is not well cited. The notations of all these authors, plus Fraas (1891) and v. Huene (1922a, 1931), clearly show that only one such big specimen was present in Tübingen (see also Koken 1905a, 1905b; Hennig 1923). It is not clear to us why v. Huene personally communicated to the curator Helmut Hölder about an ichthyosaur from Pliensbach with a length of 22 feet, which did not “survive” the transport to the new museum building in the summer semester of 1902 (Hölder 1977: 134). There are two possibilities: either Ernst Quenstedt referred to the ‘23 feet specimen from Schlierbach’ and a ‘22 feet specimen from Pliensbach’ is lost, or there might have been a problem when transporting the large Pliensbach specimen, for example by confusing labels or by breakage of the specimen or by confusion with another large specimen such as GPIT-PV-30031 or with the large vertebra (not listed in the present paper) that Quenstedt (1856: 220) listed to estimate the maximal possible size of “the giant German specimen” (“der deutschen Riesenform”).

GPIT-PV-30036 [Pl. 8A; Lias ε II, 5; Holzmaden; Stenopterygius sp.; V. Huene 1931: 349, see Discussion; syn.: GPIT/ RE/09411].

This broken and unprepared specimen is located below the skeletal reconstruction of an Ophthalmosaurus icenicus (Oxford-Clay of Peterborough, UK, GPIT-PV-31364) in the current exhibition. It illustrates – for education purposes – an imaginary find in a Posidonia Shale quarry. Most likely, it belongs to Stenopterygius quadriscissus, Lias ε II, 5 (Andreas Matzke, pers. comm. 2019), and was listed by v. Huene (1931) as a skele- ton in a geode (“Skelett in einem Laibstein, Lias ε II, 5”). It represents the only specimen from this horizon in our collection.

GPIT-PV-30037 [Pl. 8B; Lias ε; Frittlingen; Stenopterygius triscissus; Godefroit 1994, Stenopterygius longifrons; Hungerbühler 1994, Stenopterygius quadriscissus, syntype, Frittlingen; V. Huene 1952, fig. 6, Stenopterygius longifrons, Lias ε II, 6, Holzmaden: same figure as Hauff 1953, fig. 9; v. Huene 1931: 360, see Discussion, Lias ε II, 11; V. Huene 1922a, pl. 8, fig. 2, Stenopterygius zetlandicus, Lias ε II, 6, Holzmaden; Quenstedt 1858, pl. 26, fig. 11, text fig. on p. 204 [partim], Frittlingen; Quenstedt 1856: 125; Quenstedt 1852, pl. 9, fig. 7, Ichthyosaurus tenuirostris, Lias ε III, Frittlingen; syn.: GPIT 29/9/7, GPIT/ RE/09413].

Hungerbühler (1994) identified this partial juvenile skeleton as a paralectotype of Stenopterygius quadriscissus. Godefroit (1994: 43) correctly cited “GPIT 29/9/7” as one of the syntype series of S. quadriscissus, but the published pictures of this specimen are listed as a synonym of Stenopterygius longifrons (Godefroit 1994: 23). Quenstedt (1851) illustrated the skull in a typical idealized manner and identified the specimen as Ichthyosaurus tenuirostris, Lias ε III, from Frittlingen. Michael Maisch (pers. comm. 2019) agrees with the locality due to the exceptional preservation; however, he is skeptical regarding the stratigraphic layer. Also, Andreas Matzke (pers. comm. 2020) reported that findings from Lias ε III (upper epsilon, see GPIT-PV-30031) are very rare. It is possible that Quenstedt did not get exact information from the collector (the “Dusslinger”, a collector from Dußlingen near Tübingen) and only provides a tentative stratigraphical allocation. More likely is the allocation of v. Huene (1922a, 1949) as Lias ε II, 6. However, in v. Huene (1922a), the author referred to the specimen as Stenopterygius zetlandicus, and called it S. longifrons in v. Huene (1949, textfig. 6). The confusing aspect is that Holzmaden is named as the locality. Of similar inaccuracy is the description of v. Huene (1931): incomplete skeleton, rostral skull part is fallen into pieces [“unvollständiges Skelett, rostrale Schädelpartie zerfallen”], Lias ε II, 10 or 11. Hungerbühler (1994) referred to this notation, which becomes obvious after applying the method of exclusion. Most likely, the notation “rostral” is a confusion with “caudal” with reference to the whole skeleton and not to the skull alone. Currently, the specimen is labelled as Stenopterygius triscissus in the exhibition. Confusion with GPIT-PV-30018 is not possible, because the latter specimen – except for the disarticulated snout – is very well articulated and fairly complete.

GPIT-PV-30038 [Pl. 8C; Lias ε II, 6; Frittlingen; Temnodontosaurus trigonodon; Maisch 1998a: 422, Temnodontosaurus trigonodon, GPIT 171/21/2; Hungerbühler 1989, fig. 10, Leptopterygius burgundiae, GPIT 171/12/2; Mcgowan 1979, Leptopterygius burgundiae, GPIT'B04‘, Frittlingen; v. Huene 1931: 370, see Discussion; V. Huene 1922a: 30, table column 7, (pl. 22, fig. 19), Leptopterygius acutirostris; Koken 1905a: 66, Ichthyosaurus ingens; Fraas 1891, pl. 12, fig. 2, pl. 11, fig. 12, Ichthyosaurus ingens, Lias ε II, 6, Frittlingen; Quenstedt 1858, pl. 26, fig. 13, p. 220, Ichthyosaurus trigonodon, Frittlingen; syn.: PV 5760, GPIT 171/12/2, GPIT/RE/09414].

v. Huene (1931) reported a skull above the plesiosaur cupboard [“Schädel über dem Plesiosaurierschrank”] and referred it to Leptopterygius acutirostris. He probably referred to it already earlier in a graphical reconstruction (V. Huene 1922a, pl. 22, fig. 19). An old photograph proves the stratigraphy as Lias ε II, 6, and refers to the study of Fraas (1891, pl. 12, fig. 2). An error is very likely, because a PV-notation exists from 1919, which says that an Ichthyosaurus trigonodon cranium of unknown origin was purchased from the Krantz Company (“PV 18800”). The PV-number listed by Fraas (1891) (“PV 5750”) is certainly wrong and represents confusion with “PV 5759”, or at least “PV 5760”. In addition, Fraas (1891, pl. 11, fig. 12) pictured one of three teeth referred to by Quenstedt (1858, pl. 26, fig. 13: “[…] drei [kegelförmige Zähne] mit Kieferrest von einem 3¾ ‘langen Schädel aus Oberepsilon von Frittlingen […]”. It is conceivable that the allocation to “upper epsilon” belongs to the second specimen from Frittlingen (GPIT-PV-30031), because it came to the collection at the same time (“PV 5759” + “PV 5760”). Mcgowan (1979) referred to it as Leptopterygius burgundiae and listed the collection number “GPIT B04”.

GPIT-PV-30039 [Pl. 9A; Lias ε II, 6; Holzmaden; Eurhinosaurus longirostris; Godefroit 1994, Eurhinosaurus longirostris; Mcgowan 1979, Eurhinosaurus huenei, Lias ε II, 6; v. Huene 1931: 374, see Discussion, Holzmaden; V. Huene 1922a, pl. 9, fig. 4, p. 35, table, Eurhinosaurus longirostris; Koken 1905a, fourth plate after p. 64; syn.: GPIT 440, GPIT/RE/09415].

This Eurhinosaurus longirostris specimen is represented by a skull and parts of the postcranium. It was prepared by B. Hauff in 1901, and it originated from Holzmaden based on the information scratched into the sediment. The old inventory number, “GPIT 440”, gives its stratigraphic provenance as the Fleins (Lias ε II, 3). More likely, however, is the notation of v. Huene (1922a, pl. 9, fig. 4) as Lias ε II, 6, because of the signature in the sediment mentioned above. v. Huene (1931) described the specimen as a skull having an oblique lower jaw pointing upwards [“Schädel mit schräg aufwärts stehendem Unterkiefer”] and makes correct reference to his older publication.

GPIT-PV-30040 [Pl. 8D; Lias ε; Ohmden; Stenopterygius triscissus; Maxwell 2012, holotype, Stenopterygius triscissus, GPIT 12/0224-2; Maisch 2008, pl. 1, fig. 2, Stenopterygius triscissus, Lias ε II, 6; Hungerbühler 1994, fig. 1, holotype (Quenstedt 1843), GPIT 12/0224-3; Mcgowan 1979, did not identify; Johnson 1979, GPIT C 06; v. Huene 1931: 359, Stenopterygius megacephalus, Lias ε II, 6; Koken 1905a: 66, Stenopterygius triscissus, Original; Fraas 1891, table on p. 52, column 5, p. 51: “bis jetzt noch ein Unikum“; Quenstedt 1885, pl. 15, fig. 10, p. 203, Ichthyosaurus triscissus, Lias ε, Boll; Quenstedt 1858: 219; Quenstedt 1856: 223; syn.: GPIT 12/0224-2, GPIT/RE/09416].

This is the holotype of Stenopterygius triscissus (see Koken 1905a: 66: ‘the original of Quenstedt’: “Ichthyosaurus triscissus QU. Das Original“), Lias ε II, 6, from Ohmden, Boll quarry. Hungerbühler (1994) identified this specimen after comprehensive measurements of body proportions. He concluded that Quenstedt must have known this specimen already in 1843. Unfortunately, there is only one documentation in the Petrefactenverzeichnis (“PV 16763”), leaving it unclear whether a different specimen or a new restoration could have been cited. Nevertheless, it is the oldest almost complete ichthyosaur skeleton in the Tübingen collection. Quenstedt (1885) only illustrated the forelimb, which can clearly be associated with this specimen. v. Huene (1931) explicitly reported three notches in the forelimb [“größeres Skelett (an der Wand im Reptilsaal) mit drei Scissen in der Vorderflosse”], but he determined it as Stenopterygius megacephalus. Mcgowan (1979: 132) not clearly could identify this specimen (Quenstedt 1843: 223). Hungerbühler (1994) cited it in the text with the number “GPIT 12/0224-3”, which, however, became “GPIT 12/0224-2” in the figure caption and was cited as such by Maisch (2008: 232, no. 59) and by Maxwell (2012: 111).

GPIT-PV-30041 [Pl. 9B; Lias ε II, 4/5; Holzmaden; Hauffiopteryx typicus; Maxwell & CortÉs 2020, fig. 1A, fig. 2A, Hauffiopteryx typicus, lectotype; Maisch 2015: 423, 817a, GPIT; Maisch 2008, pl. 7, fig. 3, fig. 1A, Hauffiopteryx typicus, lectotype; Godefroit 1994, Stenopterygius hauffianus; Mcgowan 1979: 105, Stenopterygius hauffianus, Lias ε II, 4–5; Keller 1976, fig. 10c, Stenopterygius hauffianus, Nr. 7; Heller 1965, fig. 1; Hofmann 1958, pl. 8, fig. 1, BL I-2, 190 cm; Hauff 1953, pl. 7b; v. Huene 1931, pl. 2, fig. 1, Stenopterygius hauffianus, Nr. 5 Hautexemplar; syn.: GPIT 1491/4, GPIT/RE/09417].

This is the lectotype of Hauffiopteryx typicus (Maisch 2008), Lias ε II, 4–5, from Holzmaden, and has soft tissue preservation. v. Huene (1931, pl. 2, fig.1: “Nr. 5, neue Nummer, in Tübingen, Hautexemplar”, GA 80/272, letter 30; [likely: BERNHARD HAUFF album:] “No. 774”, St. hauffianus: sent on November 13th, 1925) allocated it to Stenopterygius hauffianus forma typica. Mcgowan (1979) adopted this determination. Thanks to the illustrations in Hofmann (1958, pl. 8a, fig. 1), Heller (1965), Keller (1976), Maisch (2015), and Maxwell (2020), the identification of this specimen is not problematic.

GPIT-PV-30042 [Pl. 9C; Lias ε; Holzmaden; Stenopterygius triscissus; Maisch 2015: 423, 817b, Stenopterygius triscissus, GPIT; (?) Maisch 2008, Stenopterygius triscissus, GPIT 1491/10 (wrong); Kröner 1999, fig. 33, fig. 34, length 120 cm; Caldwell 1997: 12, fig. 5H, Lias ε II, 4, GPIT 1297/11 (wrong); Mcgowan 1979, probably Stenopterygius hauffianus, GPIT 1491/3; Keller 1976, fig. 10d, Stenopterygius quadriscissus, Nr. 8, Lias ε II, 6; Heller 1965, fig. 3, see Discussion; Hofmann 1958, SL II-1, 115 cm; syn.: Re 1297/1, GPIT/RE/09418].

This is another juvenile skeleton with soft tissue preservation labelled as Stenopterygius quadriscissus from Holzmaden in the exhibition. With reference to GPIT-PV-30017, the confusion is enormous, because v. Huene (1931) allocated a specimen with soft tissue (length 115 cm) to the Lias ε II, 4. Heller (1965) and Keller (1976) confirmed this stratigraphic provenance. v. Huene (1922a: pl. 21, fig. 4), however, reported in the text a specimen with skin preservation with a length of 118 cm, although a skin-specimen with a length of 112 cm (GPIT-PV-30017) serves as the reference for his reconstruction. Moreover, directly on the object, a typical signature of Bernhard Hauff and the preparation date “1935” is visible. In a letter of Hauff to v. Huene, dated December 16th, 1929 (GA 80/272, letter 53), clear reference is made to the Tübingen specimen with skin preservation (GPIT-PV-30017). In another letter, dated July 14th, 1930 (GA 80/272, letter 65), a new, 114 cm long [Stenopterygius] quadriscissus with skin preservation is listed as number 225. It is unknown whether this specimen No. 225 was offered for sale or whether only a photograph was sent to v. Huene so that he could conduct his statistical evaluation. A specimen also reported in this letter, no. 54, is clearly published ( v. Huene 1931: 359), whereas no. 225 is not cited in this study. Another specimen in the photo album of Hauff, No. 234, can be clearly identified as GPIT-PV-30042 as the length of the plate (122 cm), the horizon (L ε II, 6), and the year of selling (1935) are clearly indicated.

From the earliest time to this day, it is common practice in the Hauff workshop that the specimens are only roughly prepared before a fine preparation is conducted, just before it is sold or shown on display (Rolf Bernhard Hauff, pers. comm. 2012). As such, we cannot trace whether v. Huene published the specimen GPIT-PV-30042 already in 1931 before the signature of Hauff was added to the object (Klaus Nilkens, pers. comm. 2019). Based on that, any notation in the literature concerning a specimen housed in Tübingen before 1930, the skin-preserved specimen “with a bit more than one meter in length” [“wenig mehr als ein Meter Länge”] can only refer to one specimen (GPIT-PV-30017). The label is hard to read; however, one can identify “RE 1297/1”. Hungerbühler (1994) did not report the two specimens with soft tissue preservation (GPIT-PV-30017 and GPIT-PV-30042). Maisch (2008: 240) referred to the fossil probably as S. triscissus, but called it “GPIT 1491/10”, which was used for a different specimen (compare with specimen GPIT-PV-30032) in the same publication (Maisch 2008: 231, no. 14). This issue was solved by Maisch (2015), where the author figured the specimen by only naming it as “GPIT” (without a collection number). The correctness of his determination has been approved by Erin Maxwell (pers. comm. 2019). However, the (more or less) correct length of this specimen is 119 cm.

GPIT-PV-30043 [Pl. 1D; Lias ε II, 6; Holzmaden; Temnodontosaurus trigonodon; Maisch 1998a: 422, GPIT ohne Nr.: isolierte Hinterextremität; v. Huene 1931: 370, length: 92 cm; V. Huene 1922a, pl. 4, fig. 4, Leptopterygius acutirostris, Lias ε II, 6; syn.: GPIT/RE/01608].

This specimen is an isolated hind limb. It shows a remarkable double notch [“Scisse”] in the astragalus. It was figured by v. Huene (1922a, pl. 4, fig. 4), who ( v. Huene 1931) also reported it as ‘large hind limb’ [“große Hinterflosse”] and referred it to Leptopterygius acutirostris, Lias ε II, 6.

GPIT-PV-30044 [Pl. 1E; Lias ε II, 6; Holzmaden; Temnodontosaurus trigonodon; Maisch 1998a: 422, GPIT PV 18352: isolierte Hinterextremität + Becken; V. Huene 1922a, pl. 4, fig. 3, Leptopterygius acutirostris, Lias ε II, 6, length: 70 cm; syn.: PV 18352, GPIT/RE/01610].

Only represented by a hind limb and one-half of the pelvis, which is presently covered by the wooden frame and, therefore, not visible. It is a bit smaller than GPIT-PV-30043. It was figured as Leptopterygius acutirostris from Lias ε II, 6, by v. Huene (1922a, pl. 4, fig. 3), but seems not to be cited by v. Huene (1931). The PV-number is dated 1911 as the time it was bought together with GPIT-PV-30010.

GPIT-PV-30045 [Pl. 6C; Lias ε II, 10; Holzmaden; Suevoleviathan cf. integer; Maisch 2020: 154, excluding Suevoleviathan, PV 18136 (wrong); Maxwell 2018, table 1, Suevoleviathan integer; Maisch 1998a: 66, fig. 8, cf. Temnodontosaurus sp., Holzmaden, PV 18136 (wrong); v. Huene 1931, pl. 2, fig. 6, Leptopterygius integer; V. Huene 1922a, table on p. 23, Leptopterygius integer Mutation dissidens, Lias ε II, 10; syn.: PV 19136, GPIT/RE/09402].

This incomplete small skeleton was listed by v. Huene (1922a: table on p. 23; 1931, plate 2, fig. 6) as Leptopterygius integer, Lias ε II, 10, Holzmaden. In the figure captions, however, he wrote “Mutation dissidens”, a notation he skipped in the text and did not refer to in this figure, either. Maisch (1998a, fig. 8) allocated the specimen to cf. Temnodontosaurus sp. and insisted on this determination in 2020. In the “Petrefactenverzeichnis”, No. 18136, as provided by Maisch (1998a), lists five specimens of unknown taxonomic status. They were collected by ERNST Koken in 1908. The correct PV-number must be PV 19136, because it is cited as “Leptopterygius disinliga” (should be correctly spelled “L. disinteger”) sold by B. Hauff together with the well-known GPIT-PV-30061 and GPIT-PV-30051 in February 1930.

In 1929, Bernhard Hauff reported a L. integer from Lias ε II, 10 (GA 80/272, letter 53, dated December 16th, 1929), together with three other specimens to be sold (i.e., St. quadriscissus, St. megalorhinus, St. crassicostatus mut. antecedens). It is most unlikely that this could be a misidentification of this rare species. Also, it can be excluded that there are more than two specimens of L. integer in Tübingen (i.e., GPIT-PV- 30022 and GPIT-PV-30045). However, it is likely that St. quadriscis- sus could be GPIT-PV-30061 and St. crassicostatus mut. antecedens GPIT-PV-30051. The cited St. megalorhinus in this letter (GA 80/272) could be GPIT-PV-30032, although there is no indication in the Petrefactenverzeichnis when exactly this specimen was sold. In this case it is not solved which specimen v. Huene (1922) could have meant or where exactly the specimen GPIT-PV-30045 was housed at that time.

GPIT-PV-30046 [Pl. 10A; Lias ε II, 11; Ohmden; Stenopterygius quadriscissus; v. Huene 1931: 360, see Discussion, Stenopterygius zetlandicus; syn.: PV 5556, GPIT/RE/09445].

This very old specimen shows an unusual arrangement of the disarticulated elements. v. Huene (1931) described it as a skeleton without a tail and “with three snout tips [i.e., jaw halves] next to each other” [“Skelett ohne Schwanz mit drei Schnauzenspitzen nebeneinander”] and referred it to Stenopterygius zetlandicus, Lias ε II, 11, from Ohmden. Different specimens are listed as “PV 5554 to PV 5564” [ex. private collection of the medical doctor ERNST Gustav Friedrich Hartmann sen. (1767–1851) from Göppingen, received in 1847], but only the labelling “5556, Ohmden” can be found on this specimen and is still preserved. No other specimen in the collection (and discussed in this manuscript here) shows one of these PV-numbers. However, it is most likely that at least some of the syntype series of Quenstedt can be found among them, because the notations are clearly from some time before 1850 (Quenstedt started his position as curator and institute director in Tübingen in 1837).

GPIT-PV-30047 [Pl. 10B; Lias ε; Bad Boll; Stenopterygius quadriscissus; Hungerbühler 1994, Lias ε II, 4, Boll; Mcgowan 1979, not assignable to a species; Hofmann 1958, SL III-1; v. Huene 1931: 349, Stenopterygius quadriscissus, Lias ε II, 3/4; Hennig 1923: 88, mit Pathologie; Koken 1905a: 66, Ichthyosaurus aduncus; Quenstedt 1885, pl. 16, fig. 27, p. 205, Ichthyosaurus tenuirostris aduncus, Lias ε, Boll; syn.: PV 17909, GPIT 80/16/27, GPIT/RE/09544].

This specimen was described as Ichthyosaurus tenuirostris aduncus by Quenstedt (1885, pl. 16, fig. 27), Lias ε, Bad Boll. The label on the specimen: “Quenstedt 1858” is obviously wrong, due to transposed numbers (1858 vs. 1885). v. Huene (1931) allocated the specimen to Stenopterygius quadriscissus and refers to the description of Quenstedt (1885). Hofmann (1958, SL III-1) did the same. Using this specimen, Hungerbühler (1994) demonstrated how Quenstedt performed his methodology of naming species: Quenstedt considered ontogenetic stages, sub-species, and varieties, pathological forms (compare to Hennig 1923: 88: a ‘healing’ [“Verheilung”] is reported), or related stratigraphic levels to coin a name. This sounds like a complicated taxonomic strategy, but allows a particularly clear specimen identification. v. Huene (1931) recorded this specimen as ‘a small skeleton with upturned snout tip’ [”kleines Skelett mit aufgekrümmter Schnauzenspitze“] from the Lias ε II, 3/4.

The given PV-number named a restoration in June 1904. Obviously, the provenance has been determined at that time.

GPIT-PV-30048 [Pl. 10C; Lias ε; Holzmaden; Stenopterygius uniter; Dick et al. 2016: 426, fig. 3A, SMNS 81961 (wrong); Maisch 2008, pl. 5, fig. 2, Stenopterygius cf. uniter, Lias ε II, 6; Hungerbühler 1989, fig. 5, Stenopterygius cuneiceps, GPIT 1491/12; Mcgowan 1979, Stenopterygius cuneiceps; Keller 1976, figs. 5, 7, Stenopterygius crassicostatus, Nr. 4; Hölder & Steinhorst 1964: 113, fig. 128, Schwanzwirbelsäule (Ichthyosaurier); Hofmann 1958, SL IV-3, Stenopterygius crassicostatus V Huene 1931 ?, see Discussion; V. Huene 1922a ?, pl. 22, fig. 9, S. zetlandicus, 324 cm, reconstructed; syn.: PV 10992, GPIT 1491/12, GPIT/RE/09640].

This is the lower one of the three skeletons mounted above each other in the building's stairways towards Sigwartstraße, with a length of 308 cm. The Petrefactenverzeichnis lists an ichthyosaur vertebra from 1956 and a roughly prepared specimen from 1964 (“PV 24671”). Afterward, in 1965, different plates from Posidonia Shale are correctly listed (“PV 24691”). It is very unlikely that the specimen came to Tübingen after v. Huene died in 1969. Even more so, Hofmann (1958) probably listed this specimen as “SL IV-3” and allocated it to Stenopterygius crassicostatus. Also, correspondence between B. Hauff and v. Huene indicates that the specimens reached the collection in the early 1920s (May 31th, 1922; GA 80/272, letter 12).

The most likely notation is given with “PV 10992”, because a mega-onychit is reported as gut content [“Ichthyosaurus und Loliginiten im Magen, Holzmaden”].

It is a very large, well-preserved skeleton with stomach content. Keller (1976) referred it to S. crassicostatus and to Lias ε II, 6. v. Huene (1931) only reported one specimen that we could not find, namely: S. zetlandicus, skeleton in Tübingen (reptile hall, large) [“Skelett in Tübingen (Reptilsaal, groß)”], Lias ε II, 10. Michael Maisch (pers. comm. 2019) thinks that such a misinterpretation is unlikely; however, it would be the only reason why v. Huene (1931) might have ignored this specimen, given that it was fully prepared in 1931 (see Hennig 1935: 15: ‘In the stairways several complete ichthyosaurs from the same layer [Posidonia Shale of Holzmaden] are on display’ – “Im Treppenhause finden sich aus gleicher Fundlage [Schwarzer Jura von Holzmaden] mehrere vollständige Fischsaurier […]”. v. Huene (1922a) showed several reconstructions based on the Tübingen material. Among those (his pl. 22, fig. 9) is S. zetlandicus with a length of 324 cm (maybe, however, GPIT-PV-30054 is cited here, because it is highlighted that the specimen is incomplete and that the vertebral column was rearranged; see Hungerbühler 1994). Mcgowan (1979) referred it to S. cuneiceps, although he considered that it might not be completely comparable with the holotype. Maisch (2008) referred it to S. cf. uniter.

The well-prepared and arranged specimen could also be cited with “PV 17914” as newly prepared by B. Hauff [“Ichthyos. quadriscissus, neu praepariert, 1904”].

GPIT-PV-30049 [Pl. 11B; Lias ε; unknown locality; Stenopterygius sp.; Maisch & HungerbÜhler 1997, Stenopterygius longifrons, GPIT 1576; v. Huene 1931? syn.: GPIT 1576, GPIT/ RE/09736, GPIT/RE/12812].

This is a relatively well-preserved skull (but half of the snout is missing). v. Huene used the term Stenopterygius zetlandicus in his earlier works, but later he synonymized it with S. longifrons. v. Huene (1931) listed five specimens of S. zetlandicus from Tübingen, which we clearly could locate in the collection. As such, we cannot trace the origin of specimen GPIT-PV-30049.

GPIT-PV-30050 [Pl. 11E; Lias ε; unknown locality; Hauffiopteryx typicus; Maxwell & CortÉs 2020, fig. 3A, B, Hauffiopteryx typicus; syn.: GPIT/RE/12905].

We were not able to trace the origin of this specimen.

GPIT-PV-30051 [Pl. 11A; Lias ε II, 6; Holzmaden; Stenopterygius quadriscissus; Maisch 2008, Stenopterygius quadriscissus; Caldwell 1997, table 2, Stenopterygius hauffianus, GPIT 1491/1; Godefroit 1994, Stenopterygius hauffianus; BÖttcher 1990, pl. 5, fig. 1, GPIT 1491/1; Hungerbühler 1989, fig. 6, S. hauffianus, (with 2 embryos); Mcgowan 1979: 105, S. hauffianus (provisionally), Lias ε II, 6a; Keller 1976, Stenopterygius crassicostatus, Nr. 3; Hofmann 1958, pl. 3, fig. 6, KL IV-5; B. Hauff (jun.) 1953, fig. 30, pl. 14b, Stenopterygius crassicostatus mut. antecedens; Hennig 1935: 15, two embryos; v. Huene 1931, pl. 2, fig. 2, two embryos, length 303 cm; syn.: PV 19138, GPIT 1491/1, GPIT/RE/12910-A].

A pregnant specimen with two embryos. v. Huene (1931) described this specimen as Stenopterygius crassicostatus mut. antecedens. It is an unusual ichthyosaur with two small embryos, depicted in detail by Hauff (1953) and Hölder & Steinhorst (1964). A clear note in the “Petrefactenverzeichnis” (No. “19138”) shows that this specimen was purchased from Holzmaden in 1930 (see also “PV 19137” = GPIT-PV-30061). In the correspondence between v. Huene and Hauff, also a mutation “antecedens” with two embryos is noticed (GA 80/272, letter 50; GA 80/272, letter 53) and Hauff agreed to sell it to Tübingen on December 17th, 1929. Hofmann (1958, pl. 3, fig. 6) depicted an unmistakable delineation referred as well to S. crassicostatus mut. antecedens. Maisch (2008) allocated it to S. quadriscissus, Lias ε II, 6a [sic]. The specimen is unique in its shape and as such, it clearly can be identified in the Tübingen collection.

GPIT-PV-30052 [Pl. 19B: left; Lias ε II, 6; Holzmaden; Stenopterygius quadriscissus; B. HAUFF (Jr.) 1953, fig. 32 (left), Stenopterygius crassicostatus mut. antecedens; HÖLder & Steinhorst 1964: 113, fig. 129, Stenopterygius crassicostatus; syn.: GPIT/RE/12910-B].

Left embryo inside GPIT-PV-30051, positioned a little more caudal inside the mother.

GPIT-PV-30053 [Pl. 19B: right; Lias ε II, 6; Holzmaden; Stenopterygius quadriscissus; B. HAUFF (Jr.) 1953, fig. 32 (right), Stenopterygius crassicostatus mut. antecedens; HÖLder & Steinhorst 1964: 113, fig. 129, Stenopterygius crassicostatus; syn.: GPIT/RE/12910-C].

Right embryo inside GPIT-PV-30051, positioned a little more cranial. Both embryos were probably reported by Caldwell (1997), provided with the wrong number “GPIT 1491/6”.

GPIT-PV-30054 [Pl. 9D; Lias ε II, 10; Holzmaden; Stenopterygius triscissus; Werneburg 2019, fig. 8a, Stenopterygius triscissus; Maisch 2008, pl. 5, fig. 1, Stenopterygius triscissus; Caldwell 1997, table 2, Stenopterygius sp., GPIT 1491/6; Godefroit 1994, Stenopterygius quadriscissus (syntype); Hungerbühler 1994, fig. 4, Stenopterygius quadriscissus, syntype; BÖttcher 1990, pl. 2, fig. 4, GPIT 1491/6; Mcgowan 1979, Stenopterygius quadriscissus, paralectotype, Lias ε II, 10; Johnson 1977, pl. 2, fig. B, GPIT Z04; Keller 1976, fig. 9, Stenopterygius longifrons, No. 2; Hofmann 1958, pl. 4, fig. 5, Stenopterygius longifrons, SL IV-4; v. Huene 1931: 351, second embryo visible = GPIT/RE/09283-B; Hennig 1923: 47, only one embryo; V. Huene 1922a: 47, table, Sp. 10, Stenopterygius zetlandicus; Koken 1905a: 66, Weibchen (female); Seeley 1880, pl. 1, fig. 3; Quenstedt 1867: 157, bent embryo: “gekrümmt liegender Embryo”; syn.: PV 9775 (partim: mother), GPIT 1491/6, GPIT/RE/09287, GPIT/RE/12911-A].

Pregnant female: This second pregnant specimen, with only one embryo, illustrated by Seeley (1880, pl. 1, fig. 3), apparently came to the collection before 1860 (“PV 9775”). The inventory numbers and labels on the specimen provide confusion rather than help in identification. v. Huene (1922a: 46) incorrectly reported two embryos, which was cited (probably followed) by Hofmann (1958, SL IV-1). Hofmann (1958, KL IV-5) depicted the specimen correctly (his pl. 4, fig. 5). The analysis of Hungerbühler (1994), however, is as accurate as the figure of Keller (1976), in which no second embryo is visible on that place where it theoretically could have been present. However, only the structures of interest were depicted: Seeley (1880) illustrated a structure below the skull. It could represent the rostrum of a belemnite, which, however is missing in the specimen today. Hungerbühler (1994) suggested a basisphenoid instead of a belemnite. The stomach contents, which were of interest to Keller (1976), were apparently misinterpreted and not illustrated by Seeley (1880). A new restoration (probably “PV 17910”, inventorized June 20th, 1904) noted by Hungerbühler (1994) obviously refers only to the tail, not to the body. Some areas, which were not considered to belong to the specimen by Keller (1976) are proofed that they actually belong to it. Johnson (1976) illustrated an isolated forelimb. The shadow is visualized in such an inadequate way that one can easily imagine a manipulation of this area in the fossil as can be derived from the record of further preparation (Koken 1904: 689; Hungerbühler 1994).

This specimen is referred to as Stenopterygius triscissus (Maisch 2008), Lias ε II, 10. The previous taxonomic assignments as well as the collection numbers can only be traced based on published figures; i.e., the identification of the object mainly becomes clear by the missing area shaded by Keller (1976) and the drawings of Seeley (1880) and BÖttcher (1990, table 1). The opinion of Hungerbühler (1994) that this could be one of the type specimens of S. quadriscissus of Quenstedt (1856), cannot be refuted by any notation in the “Petrefactenverzeichnis”. The first reference of two pregnant females with one embryo each in the Tübingen collection is by Quenstedt (1867: 157). In this publication, the orientation (stretched out for GPIT-PV-30029; bent for GPIT-PV-30055) and the lengths of the embryos are correctly listed (i.e., 2½' = c. 81 cm).

GPIT-PV-30055 [Pl. 19C; Lias ε II, 10; Holzmaden; Stenopterygius triscissus; Werneburg 2019, fig. 8b, one embryo; syn.: PV 9775 (partim: embryo), GPIT/RE/12911-B].

This is the embryo inside GPIT-PV-30054.

GPIT-PV-30056 to -30061, and GPIT-PV-30047 [Caldwell 1997, table 2, juveniles GPIT unnumbered, Stenopterygius sp.; for further details see further below for each specimen].

Wall with juvenile specimens (“Jungtierwand”): This arrangement of seven juvenile specimens was a present for the 90th birthday of Friedrich V. Huene on March 22nd, 1965 (Fig. 7) A wooden wall has Plexiglas windows with the specimen's silhouettes. This wall was renewed at least once (in 2009) making access for researchers even more complicated today. In consequence, the specimens have not been well-studied. There is an unpublished list by Robinson (1974), which permits allocation of the specimens to their locations and stratigraphic horizons: “Young Ichthyosaurs Display, No. 103–107” [note: only five specimens listed]. Apparently, the original idea of the instititute's director Adolf Seilacher and the curator Frank Westphal was to sort the juveniles in a stratigraphical manner following Lias ε I, 2, up to Lias ε II, 4. This original ordering is not reflected in the catalog numbers. In addition, for specimen GPIT-PV-30061, bed Lias ε II, 2, is listed, instead of the correct Lias ε I, 2 (Hungerbühler 1994). The locations/origins are plausible.

Three of the seven specimens were clearly depicted by Quenstedt (1885, pl. 16, fig. 27, p. 205: GPIT-PV-30047), v. Huene (1931: GPIT-PV-30059), and Hungerbühler (1994, fig. 7: GPIT-PV-30061). The other four specimens cannot be identified in the works published by v. Huene with certainty, but it is plausible that all of them were reported ( v. Huene 1931).

There are two additional very small skeletons in the Tübingen collection (GPIT-PV-30009 and GPIT-PV-30082). As such, there are, in total, nine specimens fitting to the category ‘very small skeleton’ [“sehr kleines Skelett”] erected by v. Huene (1931). Seven of them cannot be identified based on the list of v. Huene (1931) alone. Three of these seven specimens, however, all from the Lias ε II, 3, could be tentatively identified, namely:

No. 310, a very young skeleton with large head [“Nr. 310 sehr junges Skelett mit großem Kopf”]: We cannot associate the number 310 to any inventory system, but based on the information “large head”, we can identify it as GPIT-PV-30058.

A neonatal skeleton [“ebenfalls fast embryonales Skelett”]: Specimen GPIT-PV-30009 is particularly small and well-preserved. The existing inventory number clearly refers to the specimen cited in v. Huene (1931).

A very small skeleton in Tübingen at the wall of the reptile hall [“sehr kleines Skelett in Tübingen an der Wand im Reptilsaal”]: Specimen GPIT-PV-30060 is the largest and best-preserved skeleton on the wall and was already depicted by Quenstedt (1856). For this reason, it might have been presented in the old exhibition to which v. Huene (1931) referred.

GPIT-PV-30056 [Pl. 12A; Lias ε II, 3; Holzmaden; cf. Stenopterygius sp.; V. Huene 1922a: 56, Nr. 6, see Discussion; syn.: PV 17955, GPIT/RE/15001].

Robinson (1974) listed this specimen as “skeleton 4, No. 106, Young Ichthyosaurs Display, Lias ε II, 3”. It is a complete, articulated, well-preserved specimen with good preparation. A very badly preserved label shows the year 1904 and, hence, this skeleton must have been known by v. Huene. Therefore, it could be listed as a young skeleton, 74 cm in length [“74 cm langes, junges Skelett, Tübingen”, v. Huene (1922a: 56)]. “PV 17955”, inventorized December 31st, 1904, gives a perfect match of the size of the embedding plate: 24 cm x 88 cm.

GPIT-PV-30057 [Pl. 12B; Lias ε II, 3; Holzmaden; cf. Stenopterygius sp.; v. Huene 1931 (?): 351, see Discussion; syn.: GPIT/RE/15002].

This specimen is slightly smaller than GPIT-PV-30056. The skeleton is articulated up to the mid-length of the tail. Robinson (1974) listed it as “skeleton 3, No. 105, Young Ichthyosaurs Display, Lias ε II, 3”. It is in generally good condition. A notation shows the year 1926. It could be possible in this case that v. Huene (1931: 351) listed this specimen as “very small skeleton in Tübingen on the wall of the reptile-hall” [“sehr kleines Skelett in Tübingen an der Wand im Reptilsaal”], but there is no additional PV-number to verify this possibility.

GPIT-PV-30058 [Pl. 12C; Lias ε; unknown locality; cf. Stenopterygius sp.; v. Huene 1931 (?): 351, see Discussion; syn.: PV 1851, GPIT/RE/15003].

This specimen was listed by Robinson (1974) as “skeleton 5, No. 107, Young Ichthyosaurs Display, probably Lias ε”. There is no sign visible that could indicate any horizon or locality. The snout is reconstructed in plaster and parts of the tail, most of the hind limb, and distal parts of the forelimb are missing. It is possible that this specimen represents the historically earliest ichthyosaur skeleton in the collection, cited as ‘carcass with thick preserved head’ (“Gerippe mit dickem erhaltenem Kopf”, PV 1851), purchased around 1842 from “Dr. SChmidt” of Metzingen for 25 Gulden. If this is true, v. Huene (1931: 351) most likely referred to it as “No. 310, very young skeleton with a large head in Tübingen” (“Nr. 310, sehr junges Skelett mit großem Kopf in Tübingen”). In case “No. 310” refers to the internal list (“album”) of HAUFF, it cannot be the oldest specimen. Nevertheless, the bad preservation indicates that it could be a very old specimen. That it was nonetheless arranged on the “Jungtierwand” further supports its importance as the supposedly oldest specimen in the collection.

GPIT-PV-30059 [Pl. 12D; Lias ε II, 4; Holzmaden; cf. Stenopterygius sp.; Mcgowan 1979, see Discussion; V. Huene 1951: 277, embryo, Eurhinosaurus longirostris; v. Huene 1931: 374, embryo, Lias ε I, 4 [sic]; V. Huene 1928: 484, embryo, Lias ε II, 4; V. Huene 1922a, pl. 4, fig. 6, Eurhinosaurus longirostris; syn.: PV 17925, GPIT/RE/15005].

This is an isolated embryo allocated to Eurhinosaurus longirostris by v. Huene (1931, pl. 4, fig. 6), Holzmaden locality. It is not clear whether the stratigraphic allocations Lias ε I, 4 (“Aschgraue Mergel”) ( v. Huene 1931: 374) or Lias ε II, 4 (“Unterer Schiefer”) (V. Huene 1922a: 38) are correct. Lias ε I, 4, could be a typo or a later correction by Hauff (see “GA 80/272”). PV-number “17925” seems to be correct and, as such, it might have entered the collection together with “PV 17914” (Ichthyos. quadriscissus) and “PV 17922” (Ichthyosaurus) in spring 1904 (these old numbers could not be identified; see GPIT-PV-30056). Robinson (1974: ‘skeleton 2, No. 104’) cited the wrong PV-number (as “PV 17924”) and Mcgowan (1979) already doubted her determination.

GPIT-PV-30060 [Pl. 13A; Lias ε II, 3; Holzmaden; cf. Stenopterygius quadriscissus; Godefroit 1994, Stenopterygius quadriscissus (syntype); Hungerbühler 1994, fig. 6, Stenopterygius quadriscissus (syntype); v. Huene 1931: 351, Kleines Skelett, Lias ε II, 3; Quenstedt 1856, pl. 26, figs. 3–5, 8–10, Ichthyosaurus tenuirostris; syn.: GPIT 43/26/3, GPIT/RE/15006].

This juvenile specimen was allocated as a syntype of Stenopterygius quadriscissus by Hungerbühler (1994, fig. 6), Lias ε II, 3 from Holzmaden. Quenstedt (1856) illustrated several details of this skeleton (i.e., scapula, coracoid, and humerus), nevertheless identification is slightly difficult. This specimen was listed by Robinson (1974) as ‘skeleton 7, No. 109, Young Ichthyosaurs Display, Lias ε II, 3’; and it was probably listed by v. Huene (1931: 351) as ‘small skeleton in Tübingen on the wall of the reptile hall’ [“Kleines Skelett in Tübingen an der Wand im Reptilsaal”].

GPIT-PV-30061 [Pl. 13B; Lias ε I, 2; Zell; cf. Stenopterygius quadriscissus; Hungerbühler 1994, fig. 7, Stenopterygius promegacephalus; Heller 1965, fig. 4, Stenopterygius quadriscissus; v. Huene 1949: 83, “Hautexemplar” (specimen with skin); Lias ε I, 2; v. Huene 1931: 349, “Kleines Skelett mit typischer Incisur am Coracoid” (small skeleton with typical insicure at the coracoid), PV 19137; syn.: PV 19137, Nr. 223a + Nr. 223b, RE 1297/2, GPIT/RE/15007 (two parts, a + b)].

This specimen (“GPIT 1297/2”) is one of two syntypes of Stenopterygius promegacephalus listed by Hungerbühler (1994). The given PV-number (“PV 19137”) is identical to the original scarified label, which also says 1930 as the date of receipt in the collection (together with the stratigraphical allocation “I, 2”, which was later corrected with black color as “II, 2”). The specimen consists of the actual specimen in the exhibition and an imprint of it in the collection's archive. The smaller part, which clearly shows soft tissue preservation that was sampled for research on melanin (perhaps by Vinther 2015), shows the number 223b of the catalog of the Hauff Company. The image of Heller (1965: fig. 4) shows much more detail than the one of Hungerbühler (1994). But this is only caused by a different presentation of the outline. v. Huene (1931) listed a small skeleton with a broad coracoid with a typical recess from Lias ε I, 2, Zell, and allocated it to S. quadriscissus. This sign is scarified to the sediment as a signature of B[Ernhard] Hauff. In this case, the notation of Robinson (1974) ‘Lias ε II, 2 from Zell’ obviously is wrong. However, this confusion already appeared in the correspondence between Hauff and v. Huene [GA 80/272, letter 53, dated December 16th, 1929; “St. quadriscissus mit recht gut erhaltener Haut an der Rückenflosse aus II, 2” (St. quadriscissus with a quite well-preserved skin at the dorsal fin from II, 2)]. In many letters, “the small St. quadriscissus from II, 2, from the Zell locality” was noticed, before (apparently the same specimen): “der St. quadriscissus aus I, 2 [wurde] photografiert”.

GPIT-PV-30062 [Pl. 14A; Lias ε II, 6; Bad Boll; Eurhinosaurus longirostris; Godefroit 1994, Eurhinosaurus longirostris; V. Huene 1931: 374; V. Huene 1922a: 35, table, Eurhinosaurus longirostris, Lias ε II, 6; Quenstedt 1885, pl. 16, fig. 26, p. 205, Ichthyosaurus longirostris, Lias ε Boll; syn.: PV 10505, GPIT/RE/15008].

This is a large, partly dissociated skull with some postcranial elements from Bad Boll, Lias ε II, 6. Quenstedt (1885, pl. 16, fig. 26) illustrated this skull in an idealized manner as Ichthyosaurus longirostris. v. Huene (1931) cited this publication and listed the specimen as Eurhinosaurus longirostris, “big skull” [“großer Schädel”]. The old PV-entry “10505” allocated it as 5' length [5 Fuß = 146 cm] from Holzmaden, but synonymized it with “PV 9883” from Ohmden.

GPIT-PV-30063 [Pl. 14B; Lias ε II, 4; Holzmaden; Stenopterygius triscissus; Maisch 2008, pl. 4, fig. 1, Stenopterygius triscissus; Mcgowan 1979, Stenopterygius quadriscissus; Johnson 1979, fig. 18c; Keller 1976, fig. 10f, Stenopterygius megalorhinus; Hofmann 1958, KL II-1, 290 cm; v. Huene 1931: 357, Nr. 292, 290 cm long; syn.: PV 19021, GPIT 1491/7, GPIT/ RE/15009].

This is a relatively large specimen with soft tissue preservation. The left forelimb likely does not belong to this specimen. v. Huene (1931) referred to this specimen as Stenopterygius megalorhinus, Mcgowan (1979) and Hofmann (1958, KL II-1) as S. quadriscissus, Maisch (2008) as S. triscissus, Lias ε II, 4. Thanks to the length (290 cm) documented by v. Huene [“Nr. 292, ca. 2,9 m lang”] and the figures by Keller (1976) and Maisch (2008), the specimen can be easily identified. This identification is verified by a letter from Hauff to v. Huene (GA 80/272, letter 22, dated November 4th, 1923), in which a “typical megalorhinus-skeleton with a length of 292 cm” “with a strong skin part at its hip” (“typisches megalorhinus-Skelett von 292 cm Länge” “mit sehr kräftiger Hautpartie am Becken”) is reported.

GPIT-PV-30064 [Pl. 14C; Lias ε II, 10; Holzmaden; cf. Stenopterygius quadriscissus; v. Huene 1931 (?): 360, see Discussion; syn.: GPIT/RE/15010].

There are some skulls in the collection, which are embedded dorsoventrally and are flattened. v. Huene (1931) certainly listed one of them as Stenopterygius zetlandicus: ‘skull in dorsal view, Lias ε II, 10’ [“Schädel in Dorsalansicht, Lias ε II, 10”]. Which skull he meant exactly cannot be evaluated. A very well-preserved and complete specimen of Ichthyosaurus quadriscissus came to the collection in 1908, however, having an incorrect label as ‘Lias ε II, 6’. The correct allocation is scratched into the shale plate as’Lias ε II, 10’.

GPIT-PV-30065 [Pl. 13C; Lias ε II, 7/8; Ohmden; cf. Stenopterygius quadriscissus; Maisch 2008, pl. 7, fig. 1, Stenopterygius hauffianus posterus; Maisch 1998a: 408, Stenopterygius hauffianus (lectotype), GPIT 18387; Godefroit 1994: 45, Stenopterygius hauffianus (syntype), GPIT 18387; Mcgowan 1979, pl. 2, fig. 2, Stenopterygius hauffianus (lectotype); v. Huene 1931: 361, Stenopterygius hauffianus; V. Huene 1922a: 54, “Schädel mit 4 Wirbeln” (skull with four vertebrae), No. 10387; syn.: PV 18387, GPIT/RE/15011].

Acquired in 1912, this small specimen received PV-number “18387” and was listed as Ichthyosaurus macrophthalmus. Hauff (1921) did not know any complete specimens of this very rare species; however, he wrote that it only appears between Lias ε II, 9, and Lias ε II, 11. v. Huene (1931) described it as ‘skull without snout tip and four vertebrae’ [“Schädel ohne Schnauzenspitze und 4 Wirbel”] and referred it to Stenopterygius hauffianus, Lias ε II, 7–8. Unfortunately, Mcgowan (1979) selected it as the lectotype of S. hauffianus and listed it as “GPIT 18387” (p. 105), but in the list of valid species (p. 128) as “GPIT 10387”. Maisch (2008) cited the specimen incorrectly as well as “GPIT 10387” from Ohmden in the text, but as “GPIT 18387” in the illustration (his pl. 7, fig. 1) and as Stenopterygius cf. quadriscissus. v. Huene (1922a) also described a skull, which is still connected to the first four vertebrae, as number “PV 10387” (S. hauffianus). In his published table (V. Huene 1922a: 55), the specimen is correctly listed as number “18387”, but from Holzmaden.

GPIT-PV-30066 [Pl. 15A; Lias ε II, 6/7; Holzmaden; unclear taxonomic determination; Mcgowan 1979: 105, probably Stenopterygius hauffianus, GPIT 1491/3; v. Huene 1931: 361, see Discussion; syn.: GPIT/RE/15012, GPIT 1491/3].

Mcgowan (1979) possibly used the number “GPIT 1491/3” for this specimen. Instead of a figure, he only provided the length of the jaw as 34.6 cm. There is a slightly disarticulated skull in the collection, which is embedded together with postcranial elements. The measurement perfectly fits. There is a notation on it saying that the specimen likely came from B. Hauff to Tübingen in 1923. v. Huene (1931) listed it as Stenopterygius hauffianus, Lias ε II, 6–7, “skull with anterior half of the skeleton, from the Holzmaden locality” [“Schädel und vordere Skeletthälfte, aus Holzmaden”].

GPIT-PV-30067 [Pl. 15B; Lias ε II, 4 (?); Bad Boll; cf. Eurhinosaurus huenei; Maisch 2022, Eurhinosaurus huenei; v. Huene 1931: 364, Stenopterygius crassicostatus; Fraas 1891, pl. 14, fig. 10, Ichthyosaurus longirostris, right forelimb; Quenstedt 1885, pl. 15, fig. 9, p. 203, Ichthyosaurus biscissus, Boll; syn.: GPIT/RE/15013].

This isolated (perhaps left) forelimb was illustrated as Ichthyosaurus biscissus by Quenstedt (1885, plate 15, fig. 9). Fraas (1891, pl. 14, fig. 10) allocated it to Ichthyosaurus longirostris and illustrated the specimen as a left forelimb (in a mirrored manner) together with a coracoid and a scapula (both cannot be located in the collection today, one fragmented coracoid adheres to the limb) from Holzmaden. Another hind limb specimen, called “Wurstemberger's long hind limb” (“Wurstembergers lange Hinterflosse”) was referred to the same specimen by Fraas (Ichthyosaurus longipes; GPIT-PV-30078). v. Huene (1931: 374) listed it as a different specimen when compared to the “hind limb specimen in Tübingen (drawer)” (“Hinterflosse in Tübingen (Schublade)”) and referred both to Eurhinosaurus longirostris. The stratigraphical layer provided (Lias ε I, 4) is doubtful; more likely is Lias ε II, 4 (see GPIT-PV-30059).

GPIT-PV-30068 [Pl. 15C; Lias ε; Bad Boll; Stenopterygius quadriscissus; V. Huene 1952, fig. 10; Fraas 1891: 50; Quenstedt 1885, pl. 14, fig. 18, p. 199, Ichthyosaurus tenuirostris; Quenstedt 1852, pl. 9, fig. 5, Lias ε von Boll; syn.: GPIT 80/15/9, GPIT/RE/15014].

The shoulder girdle and forelimbs from Bad Boll are largely idealized in the drawing of Quenstedt (1852, pl. 9, fig. 5) and were referred to Ichthyosaurus tenuirostris. v. Huene (1952, fig. 10) also drew an idealized picture of this specimen (two coracoids, two scapulae, two claviculae, and one interclavicle), but – compared to Quenstedt (1852) – without the forelimbs. As such, it is impossible to state with certainty which specimen is meant. GPIT-PV-30027 and GPIT-PV-30070 also show this body region in very good preservation. Fraas (1891: 50) referred to the figure of Quenstedt (1885, pl. 14, fig. 18) as “a very nice specimen of the shoulder girdle” (“ein sehr schönes Präparat des Brustgürtels”).

GPIT-PV-30069 [Pl. 14D; Lias ε; unclear locality; cf. Stenopterygius quadriscissus; Quenstedt 1885: 203, Ichthyosaurus communis; syn.: GPIT 80/203-1, GPIT/RE/15015].

The label on this skull has an oval shape as used by Gustav Schübler in the early 19th century (HÖLder 1977). As such, this must be a very old specimen (without any reference to the Petrefactenverzeichnis). Quenstedt reported a skull from Holzmaden with a length of 1¾ ‘, which is almost 51 cm.

GPIT-PV-30070 [Pl. 16A; Lias ε; Bad Boll; Stenopterygius triscissus; Koken 1905a: 64; syn.: PV 16763, GPIT/RE/15016].

Koken (1905a: 64) reported four specimens that ‘one could still call embryos’ [“Vier kann man noch Embryonen nennen”]. He also clearly described this larger specimen as being partly prepared on both sides and as having a complete shoulder girdle [“Das andere von den beiden etwas größeren Exemplaren ist teilweise auf beiden Seiten präpariert und besitzt einen intakt erhaltenen Brustgürtel”]. It is not clear if the old PV-number “16763” (Ichthyosaurus triscissus from Lias ε from Boll) refers to this specimen, but this is most likely because the holotype (GPIT-PV-30040) has been the only one of this taxon in the collection until 1890 (Fraas 1891: 2, 51; V. Wurstemberger 1876).

GPIT-PV-30071 [Pl. 16B; Lias ε; unknown locality; cf. Stenopterygius quadriscissus; syn.: GPIT/RE/15017].

This specimen was only cited by Robinson (1974) as “No. 137”.

GPIT-PV-30072 [Pl. 18C; Lias ε II, 8; Gomaringen; cf. Stenopterygius quadriscissus; syn.: GPIT/RE/15018].

This posterior part of a skeleton came to the Tübingen collection in 1958 as a present from the owner of the Neth quarry near Gomaringen. Although many elements (e.g., hind limbs, skull) were not present or were lost (negative imprints of about 20 thoracal vertebrae are visible), it became part of the collection (Robinson 1974: No. 124), particularly as articulated vertebrate skeletons are very rare from that locality.

GPIT-PV-30073 [Pl. 16C; Lias ε; unknown locality; cf. Stenopterygius quadriscissus; syn.: GPIT/RE/15019].

This specimen was cited by Robinson (1974) as “No. 138”.

GPIT-PV-30074 [Pl. 17A; Lias ε II, 4/5; Holzmaden; cf. Stenopterygius quadriscissus; v. Huene 1931: 349, see Discussion; syn.: GPIT/RE/15020].

This is a small, unprepared skeleton, which was explicitly reported by v. Huene (1931). It came from Holzmaden, Lias ε II, 4/5. Although v. Huene (1931) listed two other ichthyosaurs from this layer (GPIT-PV-30041 and GPIT-PV-30075), there is no other specimen from this horizon in the Tübingen collection, and therefore it was noticed by him despite its bad preservation.

GPIT-PV-30075 [Pl. 17B; Lias ε II, 4/5; Ohmden; cf. Stenopterygius quadriscissus; v. Huene 1931: 361; syn.: PV 9532, GPIT/RE/15021].

This small specimen was named Ichthyosaurus quadriscissus Quenstedt, Lias ε (Ohmden). A clear mark shows the number “PV 9532”; therefore, it must have entered the collection before 1855 (because “PV 9533” was published by Quenstedt 1855). The referral by v. Huene (1931: 361) is Stenopterygius hauffianus, Lias ε II, 4/5, “skull and anterior body part in Tübingen, directly below limestone layer” [“…, Schädel und vorderer Rumpfteil in Tübingen; direkt unter der Kalkbank”]. “Kalkbank” is most likely synonymous with “Unterer Stein” = Lias ε II, 5, in this case. It is noteworthy that Robinson (1974) apparently did not list this specimen. Also important is the finding that in our collection's sample of the printed paper of v. Huene (1931) all specimens of the Tübingen collection are marked with red pencil [as: / or °] – with the exception of this (and few other) specimen(s) (see our Fig. 4).

GPIT-PV-30076 [Pl. 16D; Lias ε; unknown locality; Temnodontosaurus trigonodon; Maisch 1998a: 422; GPIT no number; v. Huene 1931 (?), see Discussion; syn.: GPIT/RE/15022].

This is likely the forelimb of Temnodontosaurus trigonodon (determination by H.S. based on the comparison to the skeleton that he reconstructed for the mayor's house in Eislingen, Germany, in 2006). Quenstedt would have certainly referred it to “Ichthyosaurusmultiscissus. In the past, this specimen was in the exhibition (see Westphal 1971: 30). Whether it appears in the list of v. Huene (1931) is not clear [see GPIT-PV-30067, v. Huene 1931: “Hinterflosse (Schublade)”]. This limb is visible in our Fig. 5. Maisch (1998a) listed some specimens without identification numbers, especially one isolated hind limb (“GPIT ohne Nr.: Isolierte Hinterextremität, Lias epsilon, Holzmaden”).

GPIT-PV-30077 (Lias ε II, 3; unknown locality; Hofmann 1958, pl. 8, fig. 6, BL-IV-1; syn.: GPIT/RE/15023].

This published specimen could not be traced in the collection. It has to be declared as missing.

GPIT-PV-30078[Pl.17C;Liasε;unclearlocality;Eurhinosaurus quenstedti; Maisch 2022, table 1, figs. 3A, B, 4B, Eurhinosaurus quenstedti, isolated forelimb; v. Huene 1931: 374, “Wurstembergers Original” (“longipes”); V. Huene 1922a, pl. 12, fig. 1, Eurhinosaurus longirostris, Lias ε II, 4; Fraas 1891, pl. 14, fig. 10, Ichthyosaurus longirostris, Holzmaden; Quenstedt 1885, pl. 15, fig. 12, p. 203, Ichthyosaurus longipes, Lias ε Ohmden; V. Wurstemberger 1876, Ichthyosaurus longipes; syn.: GPIT/RE/15024].

This is the long limb (“Flosse”) published by V. Wurstemberger (1876), known as the “Wurstembergers Flosse” in v. Huene (1922a, pl. 12, fig. 1). Fraas (1891, pl. 14, fig. 10; see also GPIT-PV-30067), as well as v. Huene (1931: 374), also named it as such, leaving no doubt about the identity of this specimen, although it is now referred to Eurhinosaurus longirostris. Quenstedt (1885, pl. 15, fig. 12) referred it to Ichthyosaurus longipes, further supporting the fact that it represents the specimen of V. Wurstemberger (1876).

GPIT-PV-30079 [Pl. 17D; Lias ε; unclear locality; cf. Stenopterygius quadriscissus; syn.: GPIT/RE/15025].

This specimen was referred to Stenopterygius hauffianus by Robinson (1974; “No. 90”). Why she reported this skull – but not GPIT-PV-30075 – is unclear. Based on the label (B. Hauff 1904), v. Huene should have been aware of this specimen. To our knowledge, there is no publication related to this specimen.

GPIT-PV-30080 [Pl. 11D; Lias ε; Ohmden; cf. Stenopterygius quadriscissus; syn.: Schübler-no.: 13550, GPIT/RE/15026].

This badly preserved, isolated skull has an oval label with a handwriting typical of Gustav Schübler (HÖLder 1977), and hence must be a very old specimen (Schübler died in 1834; see also GPIT-PV-30069). Although a certain number (13550) is given, it is impossible to trace the origin.

GPIT-PV-30081 [Pl. 11C; Lias ε; unknown locality; cf. Stenopterygius quadriscissus; v. Huene 1931, Stenopterygius crassicostatus; syn.: PV 18261, GPIT/RE/15027].

v. Huene (1931: 364) reported a fully prepared skull from the Lias ε II, 10 or 11, which is likely identical with this specimen.

GPIT-PV-30082 [Pl. 18A; Lias ε; Heiningen; cf. Stenopterygius quadriscissus; Koken 1905a: 64; Fraas 1891, pl. 6, fig. 3; syn.: PV 11167; GPIT/RE/16031].

This very small ichthyosaur skeleton was cited as an embryo by Fraas (1891, pl. 6, fig. 3). The “Petrefactenverzeichnis” lists it as “PV 11167” from Heiningen. Robinson (1974; No. 118), as well as BÖttcher (1990) described it correctly (but without illustration). It is not cited in any other publication available to us.

GPIT-PV-30083 [Pl. 18B; Lias ε; Ohmden; cf. Stenopterygius quadriscissus; Quenstedt 1885, pl. 15, fig. 1c, p. 199, Ichthyosaurus tenuirostris, Lias ε, Ohmden; Fraas 1891, pl. 6, fig. 3; syn.: PV 6372, GPIT/RE/16032].

This is a partly articulated tip of a tail, which Quenstedt (1885, pl. 15, fig. 1c) illustrated to highlight the shape of these smallest posterior caudal vertebral centra. He allocated the specimen to Ichthyosaurus tenuirostris from Ohmden. Obviously, the notation on the label, “Holzmaden”, is incorrect.

GPIT-PV-30796 [Pl. 18B; Lias ε; Ohmden; unclear determination; Godefroit 1994, Stenopterygius longifrons; Quenstedt 1856-57, pl. 26, fig. 7, Ichthyosaurus tenuirostris].

This published specimen could not be traced in the collection. It has to be declared as missing.

GPIT-PV-60576 [Fig. 8A; Lias ε III; Eislingen; cf. Eurhinosaurus longirostris; Pardo-PÉrez et. al. 2017: 26, fig. 3; Maisch 2003: 652, fig. 4; syn.: GPIT (NC/20/F/12); GPIT/RE/07096; GPIT/RE/07096-29 to -617].

This specimen was found during an excavation campaign near Eislingen an der Fils in 2002, conducted in cooperation with the “Kreisarchäologie Göppingen” (Anonymous 2002, 2004; Havlik et al. 2002, 2003, 2004). It is well documented that it is one out of at least nine almost complete skeletons. This specimen was found to belong to only one individuum because of the careful drawings, which were done during the excavation. This extremely disarticulated specimen has been excavated from the uppermost layer of Lias ε III, the so-called “Belemnitenschlachtfeld”. During the preparation, it was provisionally determined as Temnodontosaurus sp. (Maisch 2003), but when compared to GPIT-PV-60577, it was found to belong to Eurhinosaurus longirostris.

GPIT-PV-60577 (Fig. 8B; Lias ε III; Eislingen; Eurhinosaurus longirostris; Maisch 2003: 652, fig. 5; syn.: GPIT/ RE/07097; GPIT/RE/07097-1 to 362).

This specimen was found during an excavation-campaign near Eislingen an der Fils in 2003, done in cooperation with the “Kreisarchäologie Göppingen”. It is well documented that it is one out of at least nine almost complete skeletons. This specimen is very similar to GPIT-PV-60576, but slightly better articulated and more complete (especially concerning elements belonging to the skull). It was found in the so-called “Belemnitenschlachtfeld”, the uppermost layer of Lias ε III (see also GPIT-PV-60576).

4. Discussion

4.1. Research history

The present contribution resolves much long-lasting confusion about the identity of historical Posidonia Shale ichthyosaur specimens housed in the Paleontological Collection Tübingen. These impressive, large, and generally well-preserved specimens are of great interest not only to taxonomists, but for any researchers interested in the evolution of Mesozoic marine reptiles (V. Huene 1922a, 1922b, 1922c; Moon & Stubbs 2020), the origin of life history traits (Werneburg 2019), shape and distribution of paleopathologies (Pardo-PÉrez et al. 2019), ontogenetic questions (Miedema & Maxwell 2019), and shape disparities (Maisch 2008), among many other scientific aspects. In addition to the Stuttgart and Holzmaden collections in Southern Germany, Tübingen houses one of the worldwide largest Posidonia Shale ichthyosaur collections. In this respect, our detailed discussion on specimen identity will provide future researchers with an easy tool to perform unambiguous material identification. In addition to a clarification of specimen allocations, we identified different sources of confusion that might constantly occur in historical collections such as in Tübingen.

The first complex of difficulties is caused by the scientists themselves: 1) There was enormous taxonomic heterogeneity through time and the criteria to determine a specimen differs among researchers. 2) Often, an inaccurate, incomplete, and non-verified citation of the literature is apparent. Considering the particular research questions of cited authors, it is crucial to better understand why specimens were described and documented in different ways. 3) Incomplete and schematized illustrations and faulty and incomplete written documentation (e.g., collection number, measurements) of the material cause enormous confusion. 4) Often, difficulties occur when identifying old handwriting on the specimen labels. 5) Difficulties in translating foreign literature occur – most of the older Posidonia Shale ichthyosaur literature is available only in German.

The second complex of difficulties is inherent to the collection itself: 6) Was the specimen well-documented when entering the collection from the field? 7) An incomplete or non-transparent preparation of the specimen adds to the confusion. 8) The arrangements of collections, the place, and the display of specimens in the exhibition changed over time. 9) Labels associated with the specimens deteriorate or are mixed among specimens. 10) The within-house documentation of the collection is not updated or erroneous in itself.

All these issues apply to the Tübingen collection, as documented above. This should not be taken as a harsh criticism of the researcher authorities cited herein nor the highly respected previous curatorial teams. Tübingen is both blessed and burdened by a century-old collection with a very long and complicated history. In each phase of its history, different scientific standards, individual research agendas, and curatorial criteria were adopted – and even our approach may be reconsidered and possibly be found inadequate in the future. However, being aware of all the difficulties, exemplified herein by the historical ichthyosaur collection, is an imperative prerequisite to performing collection-based research for both researchers and curatorial teams, and it requires mutual cooperation. Complete and traceable specimen documentation with photographs, clear measurements, double-checked labels, and so on should be mandatory (although we acknowledge that this is a somewhat idealistic demand).

Coming back to ichthyosaurs, Jaeger (1824) already recognized a close relationship between British and Southern German ichthyosaurs, but also introduced the basis for nomenclatural confusions of synonyms (Fraas 1891). Friedrich August Quenstedt interpreted the standards of taxonomic nomenclature in a very individual and free sense (Hungerbühler 1994). It was one of the big burdens of his life that Quenstedt was unable to make drawings by himself and thus he could not do the illustrations of his many works personally (HÖLder 1977: 123). Although he recruited graphic artists for many of his impressive published plates (Fig. 3); sometimes, these artists did not illustrate the specimens adequately, as Quenstedt often critically noticed (reported by his son Ernst Quenstedt: UAT 236/59). For some publications, Quenstedt also provided several own drawings, but only of small details of the whole specimens. These illustrations were partly idealized, reproduced in a mirrored fashion, and reproduced several times.

Friedrich V. Huene performed a quantitative analysis of the ichthyosaur material ( v. Huene 1931) to study the varieties and species of different genera in the Lias ε sediments (V. Huene 1944: 36). For that, he observed hundreds of Posidonia Shale specimens in different collections. In his scientific biography, v. Huene (1944: 36) wrote: “[…] several studies were published on interesting new ichthyosaur fossils from the Holzmaden locality [V. Huene 1926, 1928, 1930, 1931], which is particularly important, because, in a very fine time-scaled stratigraphy [Hauff 1921: 33], the formation of different varieties and species of several genera can be studied accurately using hundreds of specimens – this is usually not possible in vertebrate paleontology elsewhere, because nowhere else such a detailed horizontation is processed”. Fraas (1891) as well as Hauff (1921) highlighted the detailed stratigraphy available in the profile of Lias ε at Holzmaden locality. Consequently, v. Huene (1931) referred to it and tried to catalog the ichthyosaur specimens in detail. However, as there are so many and also very similar findings, a clear identification is necessary to provide traceable scientific results. As such, a detailed stratigraphy is most helpful for identification. Unfortunately, this aspect is not always adequately covered by v. Huene (1922a, 1931) making clear allocations not possible in many cases. This omission might be mainly explained by the (logistic) inconsistencies of clear documentation in the field (GA 80/272, correspondence with Bernhard Hauff). Moreover, v. Huene often relied on old specimens which were prepared or acquired before Hauff had started to conduct his detailed stratigraphic studies, and some of his specimens were housed in private collections before entering the Tübingen collection (and are now lost).

Unfortunately, not all specimens were illustrated in most publications. Although v. Huene followed the maxim of his early mentor LUDWIG RUETIMEYER (1825–1895) “you have not seen what you did not draw” (cited after HÖLder 1977: 229), only a few specimens were illustrated in his studies (V. Huene 1922a, 1931). One has also to consider that the economic situation in Germany when his monographs were published (1920s and early 1930s) was certainly not favorable for the publication of lavishly illustrated paleontological works. Of course, illustrating hundreds of specimens was out of the scope of his studies as well. Unpublished drawings and photographs (documented by Hinz & Werneburg 2019) are often insufficient, because the species identification is not documented or not informative enough (e.g., GPIT-PV-30021) to correlate the notation of v. Huene or at least the date of origin to the collection of Tübingen.

It is likely that v. Huene (1931) was simply overworked when preparing his list of specimens (which is central to our identification). At the same time when he worked on this study, several skeletal mounts from Brazil and Southern Africa were done by him and his preparators GEORG and Wilhelm Wetzel for the Tübingen collection. He visited several museums (e.g., London) for the comparison of material. He performed a long expedition to South America (May 1928 to March 1929) and organized the shipping of eight tons of material for later examination, and “finally, also the preparation of the material that I collected in the South African Karoo Formation was finished and could be published [by me – already] in 1931” (V. Huene 1944: 36). After publishing many details on Posidonia Shale ichthyosaurs in v. Huene (1922a), v. Huene (1931) apparently concentrated mainly on the erection of a new evolutionary tree of ichthyosaurs than on particular morphological descriptions. In that way, several citation and measurement errors might have appeared.

Visionary Friedrich . Huene always tried to provide a comprehensive evolutionary tree of vertebrates (V. Huene 1936; Werneburg & Betz 2018), but his studies on ichthyosaurs are somewhat confusing, likely due to the general anatomical uniformity of that group, which makes it difficult to identify clear phylogenetic patterns. The excellent work of Hungerbühler (1994) provided much clarity at least for the genus Stenopterygius when the author identified the type material of Quenstedt. In his revision of the same taxon, Maisch (2008) provided photographs and synonymy lists. The juvenile material, however, was not considered in his study, because it would not provide enough characteristics to determine the species – but this approach excluded fossils that were considered valid species before (e.g., Stenopterygius zetlandicus, S. crassicostatus, Ichthyosaurus macrophthalmus). Maxwell (2012: fig. 1) demonstrated the complexity of nomenclatural changes using a comprehensive illustration. Finally, Motani (1999), Sander (2000), Maisch & Matzke (2000), Motani et al. (2015), and others discussed the phylogeny of ichthyosaurs in general, but not referring to individual specimens or discussing specimen identity.

In many publications (Hofmann 1958; Johnson 1977; Johnson 1979; Hungerbühler et al. 1989; Maisch 2015; Pardo-PÉrez et al. 2018a, 2018b, 2019) identification errors occurred that cannot be used for further citations. Either the numbers were incorrectly cited or they form just a continuing number within the respective publication without reference to the actual collection number. Several specimen lists were historically relevant, but today their codification is unavailable or difficult to access for most researchers: “Petrefactenverzeichnis” of Quenstedt (written from 1841 to 1887), Hauff's “Photographien-Album” ( v. Huene 1931, 1949; Hungerbühler 1994), the “Catalogue of Ichthyosaurs” by Robinson (1974) or the old “inventory list of the collection by [the former curators Frank] Westphal and [Alexander] Liebau” (McGowan 1979). Labels on the specimens are or were not always present or correct, could have been mixed in the past in a few cases, and/or typos appear(ed). Research in museums always requires the most careful consideration of all information available. Curatorial teams are requested to clearly document all details of the specimens available and to enable a consistent cataloging, numeration, and traceable location of specimens under their care.

4.2. On the completeness of objects

The fact that most objects were supplemented for conservational reasons can be seen from the glued joints and plate boundaries in the sediment. Sometimes these additions are obvious, such as e.g. at GPIT-PV-30063 (Pls. 14B, 21G). Here, there is a clear contrast between the subsequently added sedimentary plates and the original sediment, which is in direct contact with the bones. In addition, a shadow of skin is preserved on the right front fin, which is either missing or removed in a taxidermy on the left front fin. Furthermore, it is extremely unlikely that with this basically very good preservation only the left humerus should have disappeared naturally without changing the anatomically correct connection of the fin. If this left front fin can actually be assigned to the thorax, then there should in principle be preparation documentation that confirms this fact. Since this can no longer be understood in principle with the historical exhibits, legitimate doubts must be appropriate.

Another object, GPIT-PV-30016 (Fig. 3, Pls. 2D, 20B), was fully depicted by Quenstedt's drafters (Quenstedt 1852: pl. 9, fig. 2), albeit with a slightly different position of the front fin. It is also noticeable that the drawn pelvis can no longer be found in today's presentation. Looking at the surrounding sediment, no plate boundaries are visible within the area. Instead, a thin line of glue surrounds the entire body. It can therefore be assumed that this is a so-called “intarsia work” by Hauff's workshop, especially since this technique was carried out flawlessly and professionally. With the exception of the last 6 cm of the tip of the tail, only a single individual was most likely used. The exact date when this work was carried out cannot be determined with certainty, as there is no corresponding entry in the “Petrefactenverzeichnis”. However, it is known that in 1904 several other specimens (including GPIT-PV-30047; Pl. 10B) were taken to Holzmaden for restoration.

In a study on other Tübingen fossils, v. Huene (1907: 138) mentioned Jacob Hildenbrand, who casted a find in cement and sold it to Quenstedt in 1881 (PV 16718). It is very likely that Quenstedt's assistant from Ohmenhausen was responsible for the fact that the same pouring technique was used in an ichthyosaur (GPIT-PV-30025; Pl. 20H) (v. Huene 1931: 351). This specimen was carefully restored by one of us (H.S.) in 2013 and placed on permanent loan in the Natural History Museum of Luxembourg. Due to the very extensive restoration documentation, the completeness has been proven without any doubt. However, due to excessive pyrite decay, small areas on the bend in the tail and on the rostrum had to be replaced by duplicates.

It is noticeable that some ichthyosaurs have a glue line just on the spine behind the tail bend. Special attention should be paid to this with GPIT-PV-30032 (Pls. 7A, 21C). It cannot be ruled out without doubts that the position was changed here. The situation is different with GPIT-PV-30018 (Pls. 3B, 20C). A clearly reddish appearing sediment coloration behind the tail bend could indicate that it is a different individual. This is definitely not the case, however. These colorings are natural reduction horizons that often occur along fractures.

Almost all of the exhibits listed separately here have glued joints on the caudal spine. Care must be taken to ensure that the size, preservation, staining and embedding position of the bone material appear plausible. It is usually extremely difficult to make a taxonomic statement in this area in order to prove the affiliation. Therefore, justified doubts are also appropriate for GPIT-PV-30022 (Pls. 4B, 20G).

In the case of GPIT-PV-30034 (Pls. 7C, 21D), there is published confirmation that there are two individuals (Hauff 1921: 34). Comparing today's presentation with the historical illustration (Fraas 1891: pl. 12, fig. 5) reveals a displacement of both the hind fin and the tail. Likewise, the preservation, embedding position, and proportion of the hind fin are not exactly consistent with the two fore fins. Furthermore, minor manipulations in the area of the jaw can be noticed, which may be shown extended on display. Due to the extremely inaccessible suspension of the piece, it is almost impossible to examine the exact course of the gluing. A major benefit, however, was that in 2010 the opaque white paint over the sedimentary slabs was removed by former preparator Hans Luginsland.

GPIT-PV-30019 (Pls. 4A, 20E, F), on the other hand, reveals a great need for discussion. This specimen was described in detail in a dissertation (V. Wurstemberger 1876: 41–43). However, there are have no drawings available. It is striking that the lack of a kinked tail is expressively noticed, although one is clearly shown in today's presentation (Mcgowan 1979: 130). All dimensions and descriptions are also perfectly comprehensible. This tail bend was obviously constructed in a later restoration. This restoration may also account for some rib fragments, vertebral centers, and plate boundaries being retouched and added. The statement that this specimen was a composite (Maisch 2008: 243) cannot be confirmed. In a more recent dissertation (Kröner 1999: fig. 1), the extremities are described as “largely supplemented”. The position of the left front fin may well be changed. The contrast to the right front fin is also noticeable when it comes to the state of preservation and embedding. Overall, this specimen is remarkable precise because of the special embedding. The “degree” of the embedding differences between the right and left front fin can be understood surprisingly well in a similar way on the right and left rear fin (Jelle Heijne, oral comm. 2022). The embedding of the left front fin alone is not “natural” in the anatomical sense. It can be called a “rarity”. It is therefore unlikely that this should be a second individual, especially since the anatomy of both front fins is the same. It is quite possible that the number of vertebrae is shown reduced up to the point where the tail bends. Likewise, the position of the tail bend is questionable. Overall, however, the specimen as a whole appears very plausible due to its special embedding position.

A good drawing exists for GPIT-PV-30054 (Seeley 1880: pl. 1, fig. 3; see Pls. 9D, 21F). All details on the thorax (including the embryo) can be found in today's presentation. Because it is a charcoal hatch, the surrounding sediment is shown uniformly [contrary to the drawing of GPIT-PV-30028 on the same panel in Seeley (1880), where individual plate boundaries are sketched; see below]. Therefore, it can only be concluded from the drawn tip of the tail (compared to today's preservation) that the distal spine (behind the bend in the tail) does not belong to the same individual (Hungerbühler 1994: 254). The position of the piece of tail between the kink and the thorax could have been changed before the drawing was made. However, the proportions and the embedding position of the vertebral centers agree that a belonging to the same individual cannot be ruled out.

GPIT-PV-30028 (Pls. 5D, 21B) was carefully restored by H.S. in 2014. There is no doubt that all of the present bone elements belong to one individual (plus the embryo). The work was extensively documented. Indications of an original collection number or a time limit for the creation of the frame construction could not be determined. This specimen was first reported by Quenstedt (1858: 219).

GPIT-PV-30048 (Pls. 10C, 21E) was most likely already purchased in the 19th century (PV 10992 “[...] with loliginites in the stomach”). The condition of the piece is extraordinarily good; the preparation takes this fact into account through also very good work. There are thin glue lines at the outer contour line (especially on the skull and tail), which can, however, be examined properly. Therefore, it is very likely that the right hind fin was originally embedded in a different position. However, it will belong to the same individual. If not, it is very questionable why the same work did not add a corresponding front fin. The state of preservation of the entire caudal spine is so good that no additions or adhesions could be detected. Only a few rib fragments, two front vertebrae with neural arches and small parts of the fins were added. All of this strongly speaks for the fact that the surrounding slabs of slate were restored purely for conservation purposes, with possible subsequent post-preparation of the bones.

There are a few entries in the “Petrefactenverzeichnis” that are associated with a restoration by B. Hauff (sen.). Only “Ichthyosaurus longirostris” (PV 17874, May 1903 = GPIT-PV-30034) and “Ichthyosaurus aduncus” (PV 17909, June 20, 1904 = GPIT-PV-30047) are reported by name, which can be assigned without a doubt. Other specimens of “Ichthyosaurus quadriscissus” are cited, but cannot be assigned individually.

GPIT-PV-30024 (Pls. 4D, 21A) was purchased in 1949 (PV 24361: “from Holzmaden near Kirchheim, Teck”) and described in detail in 1951 [V. Huene 1951: 277: “of Lias e from (Ohmden) Holzmaden”]. From a preparatory point of view, this exhibit represents a fundamentally different dimension of restoration. The following facts can be proven on this specimen:

  • With the exception of a larger area on the left hind fin, all sediment plates are not only supplemented, but also very finely coordinated in terms of colour and structure.

  • It is noticeable that more than half of the thorax is represented by only one side of the ribs. The fact that the other side must also have been preserved can be seen from a few rib ends that are shown in the rear thorax. In addition, carefully supplemented rock plates can be seen between the individual costal arches, particularly in the area that shows the pathologically altered ribs.

  • Elaborate carvings can be found in the sediment on several neural arches of the thorax and most of the gastral ribs, which extremely carefully conceal the transition from various plate boundaries and adhesive joints.

  • The caudal spine from the bend in the tail consists essentially of three larger pieces that fit together very well due to the proportions, the preservation, and the embedding position. However, one can also show glued joints, which suggest that they could be individuals that do not belong to one another. At a minimum, both the position and angle of the kink of the tail must be questioned.

  • The right hind fin consists (especially in the distal area) of several inlays. Extreme care has been taken to ensure that there are no differences in direct comparison with the left rear fin. The assumption even arises that individual phalanges were cut in order to be used again as a mirror-image correspondence.

  • There are synthetic resin supplements at various vertebral centers, but also on the entire distal articular surface of the left humerus. These are color matched to bone preservation. As a result, the exact course of the breaking edge/plate boundary can no longer be followed quickly and easily, as is still possible along the front edge of the front fin.

  • In 2009, restoration work was carried out in the area of the left scapula. Due to pyrite decay, some bone elements have become detached from the subsoil. During the necessary repairs by the former preparator Hans Luginsland, it was found that the surrounding sediment layer was only a few millimeters thick. Given the size of the overall object, it is understandable that when assembling the exhibit, as much material as possible should be stripped down to save weight.

Due to the above facts, the following overall impression derives:

  • The completeness of the original object was either not given or was reduced accordingly to demonstrate a more aesthetic embedding state (missing right front fin, missing half thorax, missing neural arches).

  • The supplementation of the surrounding sedimentary plates was carried out with great care; possibly to create the image of a perfect finding situation, or to make it more difficult to check the original find layer (at least this difficulty was accepted with approval).

  • The suspicion arises that the pathologically altered ribs could have been decisive in making the overall object even more attractive (possibly in order to increase the purchase price; possibly in order to achieve a basic interest in buying, because an obvious justification for the publication was created). In any case, almost all staining that could indicate possible soft tissue preservation in this area of the thorax has been removed, although these clearly exist in both the anterior and posterior thorax.

  • The manipulations carried out on the object itself and in the sediment have been carried out in such a way that it is no longer possible to unequivocally verify that it belongs to a single individual.

4.3. Remarks on the original finding horizons

The by v. Huene's subsequently attempted determinations of the finding horizon contain two major sources of errors. Firstly, the correct attribution of the sample to the original fossil: If the Hauff workshop was commissioned to restore an entire exhibit, it can be assumed that this error has been minimized. Secondly, the correct attribution of the determination in the publication of v. Huene (1931): Several typos appear in this publication (e.g., GPIT-PV-30059), but also obvious misstatements when naming the locality (e.g., GPIT-PV-30037), so that a correct designation of the horizon should be doubted in individual cases. Furthermore, incorrect labelling (e.g., for GPIT-PV-30014 or GPIT-PV-30018) caused significant citation errors. Older photographic documents may help in clarifying (GPIT-PV-30019, GPIT-PV-30040) or supporting the view that it is a single complete and unadulterated rock plate, which is demonstrably not the case (GPIT-PV-30041, GPIT-PV-30045, GPIT-PV-30051).

Basically, it should be noted that all older objects (before Quenstedt 1889) cannot be named exactly horizontally. Quenstedt only distinguished between “upper”, “middle”, and “lower” Lias e, often not even those units. The subsequent fine stratigraphic investigation by v. Huene and B. Hauff sen. cannot be traced beyond doubt today. In addition, it cannot be ruled out that some of the custodian's assistants (Robinson) used the term “Holzmaden” as a synonym for “Posidonia Shale”. As such, the actual sites of Bad Boll or Ohmden, both belonging to the outcrop area of the Posidonia Shale, can no longer be localized with certainty. On the other hand, many exhibition labels are not always correctly researched, so that the old photographs provide a good indication for the identification of the specimens, but there are by no means reliable sources for the exact horizon.


For discussions, we thank Michael Maisch, Andreas Matzke, Hans Luginsland (preparator 1963–2013), Frank Westphal (curator 1963–1980, †), Edgar Bierende and Adrian Tröscher (Tübingen), Jelle Heijne (Bonn), Erin Maxwell (Stuttgart), and preparator Klaus Nilkens and director Rolf Bernhard Hauff (Holzmaden). Madelaine Böhme (Tübingen) is thanked for enabling this research in the collection. Michael Maisch and Erin Maxwell provided very comprehensive reviews that greatly helped to improve our manuscript, but we are of course responsible for all remaining mistakes. We thank Günter Schweigert (Stuttgart) for kindly handling our manuscript. For access to and for help in the University Archive Tübingen we thank Regina Keyler and Susanne Riess-Stumm. Photos of the specimens were taken by Agnes Fatz [Pls. 1A, 1B (-3), 1B (-4), 1F, 2A, 11B, 11C, 11D, 11E, 13B (b), 13C, 14C, 14D, 15A, 15B, 15C, 16D, 17B, 17D, 18A, 18B), by VERENA PIETZSCH [Pls. 2B, 2C, 2D, 3A, 3B, 3D, 4B, 5C, 5D, 6A, 6B, 6C, 7A, 8B, 8C, 8D, 9A, 9B, 9C, 9D, 10A, 10B, 11A, 12A, 12B, 12C, 12D, 13A, 13B (a), 14A, 14B, 16A, 16B, 16C, 17A, 17C), and by H.S. (1B (-1), 1B (-2), 1C, 1D, 1E, 5A, 5B, 7C, 7D, 18C, 19A, 19B, 19C), and were postprocessed by H.S. Valentin Marquardt took photos for Pls. 3C, 4A, 4C, 8A, 10C. Other photos were taken by the former institute's photographers Werner Wetzel (Pl. 4D) and Wolfgang Gerber (Pl. 7B). Christina Kyriakouli (imaging lab Tübingen) is thanked for conducting a 3D-surface scan ( Supplementary file (Supplement 1.ply)). Via chairperson Iris v. Huene, the “Familienverband der Freiherren v. Hoyningen (Hoiningen) gen. Huene e.V.” provided a donation to conduct part of the photo documentation. I.W. was funded by WE 5440/6-1.

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Electronic  Supplementary file (Supplement 1.ply). 3D-surface-model of specimen GPIT-PV-30024 made by Christina Kyriakouli (Tübingen) using an Artec-Eva-scanner and Artec 14 software (size: 48 MB). Available at can be opened in software such as Blender or Meshlab. Surface coloration is part of the file and can be viewed in selected software.

Plate 1. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen. Compare to text. Photographers listed in the acknowledgements of the article. Continued in Pls. 2–20. A – GPIT-PV-30009, B – GPIT-PV-30010, C – GPIT-PV-30011, D – GPIT-PV-30043, E – GPIT-PV-30044, F – GPIT-PV-30013.

Plate 2. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30012, B – GPIT-PV-30014, C – GPIT-PV-30015, D – GPIT-PV-3001643.

Plate 3. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30017, B – GPIT-PV-30018, C – GPIT-PV-30020, D – GPIT-PV-30021.

Plate 4. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30019, B – GPIT-PV-30022, C – GPIT-PV-30023, D – GPIT-PV-30024.

Plate 5. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30025, B – GPIT-PV-30026, C – GPIT-PV-30027, D – GPIT-PV-30028 (mother) with embryo GPIT-PV-30029 (see also Plate 19A).

Plate 6. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30030, B – GPIT-PV-30031, C – GPIT-PV-30045.

Plate 7. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30032, B – GPIT-PV-30033, C – GPIT-PV-30034, D – GPIT-PV-30035.

Plate 8. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30036, B – GPIT-PV-30037, C – GPIT-PV-30038, D – GPIT-PV-30040.

Plate 9. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30039, B – GPIT-PV-30041, C – GPIT-PV-30042, D – GPIT-PV-30054 (mother) with embryo GPIT-PV30055 (see also Pl. 19C).

Plate 10. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30046, B – GPIT-PV-30047, C – GPIT-PV-30048.

Plate11.PhotographsoftheichthyosaurspecimenshousedinthePaläontologischeSammlungTübingen,continued.A–GPIT-PV-30051 (mother) with embryos GPIT-PV-30052 and GPIT-PV-30053 (see also Plate 19B), B – GPIT-PV-30049, C – GPIT-PV-30081, D – GPIT-PV-30080, E – GPIT-PV-30050.

Plate 12. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30056, B – GPIT-PV-30057, C – GPIT-PV-30058, D – GPIT-PV-30059.

Plate 13. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30060, B – GPIT-PV-30061 (with parts a and b), C – GPIT-PV-30065.

Plate 14. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30062, B – GPIT-PV-30063, C – GPIT-PV-30064, D – GPIT-PV-30069.

Plate 15. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30066, B – GPIT-PV-30067, C – GPIT-PV-30068.

Plate 16. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30070, B – GPIT-PV-30071, C – GPIT-PV-30073, D – GPIT-PV-30076.

Plate 17. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30074, B – GPIT-PV-30075, C – GPIT-PV-30078, D – GPIT-PV-30079.

Plate 18. Photographs of the ichthyosaur specimens housed in the Paläontologische Sammlung Tübingen, continued. A – GPIT-PV-30082, B – GPIT-PV-30083, C – GPIT-PV-30072.

Plate19.PhotographsoftheichthyosaurspecimenshousedinthePaläontologischeSammlungTübingen,continued.A–GPIT-PV-30029 (= embryo inside GPIT-PV-30028), B – GPIT-PV-30052 and GPIT-PV-30053 (embryos inside GPIT-PV-30051; see also Plate 11A), C – GPIT-PV-30055 (embryo inside GPIT-PV-30055; see also Pl. 9D).

Plate 20. Hypotheses on the origin of rock plates in selected important specimens. Gray: Original specimen. Green: The green marked rock plates do not belong to the actual fossil and were added to receive a complete picture for the display in the exhibition. It is important to proof whether the added plates belong to the same stratigraphic layer as the fossil. In most cases, however, the Fleins was used to supplement the exhibit, because it is more stable when processing and contains less pyrite. Through time, by decomposition, pyrite results in large problems when storing the exhibit and may irreparably destroy the fossil. Red: The red-marked parts are likely intarsia from other specimens. In some cases (marked with “?”), they could still belong to the original specimen (gray) based on other arguments, but then these body parts were rearranged to fit the final shape of the exhibit. It is not always easy to correctly interpret the glue joints between the plates. Comprehensive restaurations would be necessary to provide a doubtless analysis. However, a clear answer cannot be expected in all cases because an accurate documentation of previous preparations is difficult. Blue: The blue-marked parts are clearly supplemented to the fossil to create the complete exhibit. Inside the actual fossil artificial resin or plaster can be expected. However, it is unclear whether the material represents a modelling or the duplication of a real bone. A clear allocation to the original fossil is extremely rare and hard to document without any doubts. Inbetween the rock plates broad joint gaps can develop through time. Hence, also in these cases, it is not always clear whether these rock areas belong together. Arrows indicate to important areas. Not to scale; compare to Pls .1–19. Continued in Pl. 21.

Plate 21. Continuation of Pl. 20. E shows a reconstruction of GPIT-PV-30019 based on the description of Maisch (2008). It is contrasted to our own reconstruction in F.





















Henrik Stöhr and Ingmar Werneburg "The Tübingen collection of ichthyosaurs from the Lower Jurassic (Lower Toarcian) Posidonienschiefer Formation of Württemberg: a historical and curatorial perspective," Palaeodiversity 16(1), 39-97, (28 April 2022).
Received: 22 December 2020; Accepted: 9 March 2023; Published: 28 April 2022
Bernhard Hauff sen
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