A New Look at Ichthyosaur Long Bone Microanatomy and Histology: Implications for Their Adaptation to an Aquatic Life

Background

Ichthyosaurs are Mesozoic reptiles considered as active swimmers highly adapted to a fully open-marine life. They display a wide range of morphologies illustrating diverse ecological grades. Data concerning their bone microanatomical and histological features are rather limited and suggest that ichthyosaurs display a spongious, “osteoporotic-like” bone inner structure, like extant cetaceans. However, some taxa exhibit peculiar features, suggesting that the analysis of the microanatomical and histological characteristics of various ichthyosaur long bones should match the anatomical diversity and provide information about their diverse locomotor abilities and physiology.

Methodology/Principal Findings

The material analyzed for this study essentially consists of mid-diaphyseal transverse sections from stylopod bones of various ichthyosaurs and of a few microtomographic (both conventional and synchrotron) data. The present contribution discusses the histological and microanatomical variation observed within ichthyosaurs and the peculiarities of some taxa (Mixosaurus, Pessopteryx). Four microanatomical types are described. If Mixosaurus sections differ from those of the other taxa analyzed, the other microanatomical types, characterized by the relative proportion of compact and loose spongiosa of periosteal and endochondral origin respectively, seem to rather especially illustrate variation along the diaphysis in taxa with similar microanatomical features. Our analysis also reveals that primary bone in all the ichthyosaur taxa sampled (to the possible exception of Mixosaurus) is spongy in origin, that cyclical growth is a common pattern among ichthyosaurs, and confirms the previous assumptions of high growth rates in ichthyosaurs.

Conclusions/Significance

The occurrence of two types of remodelling patterns along the diaphysis, characterized by bone mass decrease and increase respectively is described for the first time. It raises questions about the definition of the osseous microanatomical specializations bone mass increase and osteoporosis, notably based on the processes involved, and reveals the difficulty in determining the true occurrence of these osseous specializations in ichthyosaurs.     

A basal thunnosaurian from Iraq reveals disparate phylogenetic origins for Cretaceous ichthyosaurs

Cretaceous ichthyosaurs have typically been considered a small, homogeneous assemblage sharing a common Late Jurassic ancestor. Their low diversity and disparity have been interpreted as indicative of a decline leading to their Cenomanian extinction. We describe the first post-Triassic ichthyosaur from the Middle East, Malawania anachronus gen. et sp. nov. from the Early Cretaceous of Iraq, and re-evaluate the evolutionary history of parvipelvian ichthyosaurs via phylogenetic and cladogenesis rate analyses. Malawania represents a basal grade in thunnosaurian evolution that arose during a major Late Triassic radiation event and was previously thought to have gone extinct during the Early Jurassic. Its pectoral morphology appears surprisingly archaic, retaining a forefin architecture similar to that of its Early Jurassic relatives. After the initial latest Triassic radiation of early thunnosaurians, two subsequent large radiations produced lineages with Cretaceous representatives, but the radiation events themselves are pre-Cretaceous. Cretaceous ichthyosaurs therefore include distantly related lineages, with contrasting evolutionary histories, and appear more diverse and disparate than previously supposed.     

Ichthyosauria: their diversity, distribution, and phylogeny

The Ichthyosauria is the group of Mesozoic marine reptiles that was most highly adapted to the aquatic environment. The first ichthyosaurs from the upper Lower Triassic (Spathian) already show a suite of unique characters (very large eyes, elongate snout, deeply amphicoelous vertebrae, limb modified to fins) correlated with a fully aquatic existence and probably were unable to leave the water. The key evolutionary innovation was vivipary, giving birth to live young, which is documented by the fossil record since the end of the Anisian. Major evolutionary trends in the locomotor apparatus are the increasing modification of the fin skeleton to a mosaic of bones and the change from anguiliform swimming in the earliest forms to thunniform swimming in the Jurassic and later forms, as evidenced by the shortening of the body and the evolution of a semilunate tail fin. Almost from the beginning, ichthyosaurs had a cosmopolitan distribution which was retained until their extinction in the Cenomanian. Ichthyosaurian diversity is greatest in the Middle Triassic with piscivorous, heterodont, and durophagous forms. Jurassic diversity is greatest in the Liassic, declining to one genus (Platypterygius) in the Cretaceous. Although skull characters indicate that ichthyosaurs were diapsids, their exact position within Diapsida is unclear. A cladistic analysis of the well known genera clarifies relationships within the Ichthyosauria. Most basal areGrippia andUtatsusaurus, followed by the Mixosauridae (Mixosaurus andPhalarodon). The Shastasauridae (Cymbospondylus, Shonisaurus, Besanosaurus) are the most advanced Triassic forms and represent the sistergroup of all post-Triassic ichthyosaurs. These are clearly monophyletic and are termed here the Neoichthyosauria.     

Mikadocephalus gracilirostris n. gen., n. sp., a new ichthyosaur from the Grenzbitumenzone (Anisian-Ladinian) of Monte San Giorgio (Switzerland)

An almost complete skull of a new ichthyosaur from the Middle Triassic Grenzbitumenzone Beds of Monte San Giorgio (Kanton Tessin, Switzerland) represents one of the most complete and best preserved finds of a large Triassic ichthyosaur cranium. Its affinites with other Triassic ichthyosaur taxa are discussed and it is demonstrated to represent a new genus and species,Mikadocephalus gracilirostris, which does not fit into any of the currently recognized families of Triassic ichthyosaurs. Remarkable similarities in cranial structure exit to postTriassic ichthyosaurs, with which a number of important apomorphies are shared.     

Evidence for the longest stratigraphic range of a post-Triassic Ichthyosaur: a Leptonectes tenuirostris from the Pliensbachian (Lower Jurassic) of Switzerland

A three-dimensional well-preserved ichthyosaur skull and parts of the postcranial skeleton are attributed to the species Leptonectes tenuirostris (Conybeare, 1822). It was found vertically embedded in Pliensbachian deposits representing three successive biozones (ibex to margaritatus Zone). The find is dated as early Late Pliensbachian (margaritatus Zone) by a rich ammonite and ostracod fauna. It is the first record of the genus Leptonectes from Switzerland and from the Late Pliensbachian. It is so far the best preserved and most complete ichthyosaur from this time interval worldwide. With diagnostic specimens known from the Rhaetian (Late Keuper) up to the early Late Pliensbachian (Middle Liassic), L. tenuirostris (Conybeare, 1822) has the most extensive stratigraphic range documented for any post-Triassic ichthyosaur so far.     

The skull and endocranium of a Lower Jurassic ichthyosaur based on digital reconstructions

Even after 200 years of study, some details of the cranial anatomy of ichthyosaurs, one of the most successful groups of marine vertebrates in the Mesozoic, are still unclear. New information on the braincase, palate and occiput are provided from three‐dimensional scans of an exceptionally preserved ichthyosaur (‘Hauffiopteryx’ typicus) skull from the Toarcian (183–174 Ma, Lower Jurassic) of Strawberry Bank, England. This ichthyosaur has unusual, hollow, tubular hyoid bars. The occipital and braincase region is fully reconstructed, creating the first digital cranial endocast of an ichthyosaur. Enlarged optic lobes and an enlarged cerebellum suggest neuroanatomical adaptations that allowed it to be a highly mobile, visual predator. The olfactory region also appears to be enlarged, suggesting that olfaction was more important for ichthyosaurs than has been assumed. Phylogenetic analysis suggests this ichthyosaur is closely related to, but distinct from, Hauffiopteryx, and positioned within Thunnosauria, a more derived position than previously recovered. These results further our knowledge of ichthyosaur cranial anatomy in three dimensions and provide a platform in which to study the anatomical adaptations that allowed ichthyosaurs to dominate the marine realm during the Mesozoic.     

High Diversity in Cretaceous Ichthyosaurs from Europe Prior to Their Extinction

Background

Ichthyosaurs are reptiles that inhabited the marine realm during most of the Mesozoic. Their Cretaceous representatives have traditionally been considered as the last survivors of a group declining since the Jurassic. Recently, however, an unexpected diversity has been described in Upper Jurassic–Lower Cretaceous deposits, but is widely spread across time and space, giving small clues on the adaptive potential and ecosystem control of the last ichthyosaurs. The famous but little studied English Gault Formation and ‘greensands’ deposits (the Upper Greensand Formation and the Cambridge Greensand Member of the Lower Chalk Formation) offer an unprecedented opportunity to investigate this topic, containing thousands of ichthyosaur remains spanning the Early–Late Cretaceous boundary.

Methodology/Principal Findings

To assess the diversity of the ichthyosaur assemblage from these sedimentary bodies, we recognized morphotypes within each type of bones. We grouped these morphotypes together, when possible, by using articulated specimens from the same formations and from new localities in the Vocontian Basin (France); a revised taxonomic scheme is proposed. We recognize the following taxa in the ‘greensands’: the platypterygiines ‘Platypterygius’ sp. and Sisteronia seeleyi gen. et sp. nov., indeterminate ophthalmosaurines and the rare incertae sedis Cetarthrosaurus walkeri. The taxonomic diversity of late Albian ichthyosaurs now matches that of older, well-known intervals such as the Toarcian or the Tithonian. Contrasting tooth shapes and wear patterns suggest that these ichthyosaurs colonized three distinct feeding guilds, despite the presence of numerous plesiosaur taxa.

Conclusion/Significance

Western Europe was a diversity hot-spot for ichthyosaurs a few million years prior to their final extinction. By contrast, the low diversity in Australia and U.S.A. suggests strong geographical disparities in the diversity pattern of Albian–early Cenomanian ichthyosaurs. This provides a whole new context to investigate the extinction of these successful marine reptiles, at the end of the Cenomanian.     

First diagnostic marine reptile remains from the aalenian (middle jurassic): a new ichthyosaur from southwestern Germany

Background

The Middle Jurassic was a critical time in the evolutionary history of ichthyosaurs. During this time interval, the diverse, well-studied faunas of the Lower Jurassic were entirely replaced by ophthalmosaurids, a new group that arose sometime prior to the Aalenian-Bajocian boundary and by the latest middle Jurassic comprised the only surviving group of ichthyosaurs. Thus, the Middle Jurassic Aalenian-Bathonian interval (176–165 million years ago) comprises the time frame during which ophthalmosaurids not only originated but also achieved taxonomic dominance. However, diagnostic ichthyosaur remains have been described previously from only a single locality from this interval, from the Bajocian of Argentina.

Methodology/Principal Findings

In this paper, we describe a new species of ichthyosaur based on a partial articulated specimen from the Middle Jurassic of southwestern Germany. This specimen was recovered from the Opalinuston Formation (early Aalenian) and is referable to Stenopterygius aaleniensis sp. nov. reflecting features of the skull and forefin. The genus Stenopterygius is diverse and abundant in the Lower Jurassic of Europe, but its presence has not previously been confirmed in younger (Middle Jurassic) rocks from the northern hemisphere.

Conclusions/Significance

This specimen represents the only diagnostic ichthyosaur remains reported from the Aalenian. It bears numerous similarities in size and in morphology to the Lower Jurassic species of the genus Stenopterygius and provides additional evidence that the major ecological changes hypothesized to have occurred at the end of the Toarcian took place sometime after this point and most likely did not occur suddenly. There is currently no evidence for the presence of ophthalmosaurids in the northern hemisphere during the Aalenian-Bathonian interval.     

New finds ofOmphalosaurus and a review of Triassic ichthyosaur paleobiogeography

Triassic ichthyosaurs are very widespread and diverse but most taxa are poorly known. New discoveries, such as described in this paper, underscore the first statement but only slowy invalidate the second. The first associated skeleton of the unusual durophagous ichthyosaurOmphalosaurus (O. wolfi) is reported from the Middle Triassic (earliest Ladinian) of the northern Alps. The vertebrae are of the ichthyosaurian type, firmly establishing the ichthyosaurian affinities of the genus. A round bone from the Muschelkalk of Franconia, Germany, is identified and described as a left humerus ofOmphalosaurus sp. These finds greatly extend the paleogeographic and temporal range ofOmphalosaurus which previously was only known from the western cratonic margin of North America and from Spitsbergen. The new records ofOmphalosaurus allow a revised differential diagnosis of the genus and a revision of its species. Valid species ofOmphalosaurus areO. nettarhynchus, O. nevadanus, andO. wolfi. In conjunction with such new insights into Triassic ichthyosaur distribution, it is necessary to address the validity of the poorly known genera such as the Middle TriassicPessosaurus. This taxon must be considered invalid because it lacks diagnostic characters. In assessing progress in research on Triassic ichthyosaurs and their paleobiogeography, it becomes apparent that supra-specific diversity appears largely known, at least for the pre-Norian record of North America, Europe, and East Asia.     

Microanatomical and Histological Features in the Long Bones of Mosasaurine Mosasaurs (Reptilia,Squamata) – Implications for Aquatic Adaptation and Growth Rates

Background

During their evolution in the Late Cretaceous, mosasauroids attained a worldwide distribution, accompanied by a marked increase in body size and open ocean adaptations. This transition from land-dwellers to highly marine-adapted forms is readily apparent not only at the gross anatomic level but also in their inner bone architecture, which underwent profound modifications.

Methodology/Principal Findings

The present contribution describes, both qualitatively and quantitatively, the internal organization (microanatomy) and tissue types and characteristics (histology) of propodial and epipodial bones in one lineage of mosasauroids; i.e., the subfamily Mosasaurinae. By using microanatomical and histological data from limb bones in combination with recently acquired knowledge on the inner structure of ribs and vertebrae, and through comparisons with extant squamates and semi-aquatic to fully marine amniotes, we infer possible implications on mosasaurine evolution, aquatic adaptation, growth rates, and basal metabolic rates. Notably, we observe the occurrence of an unusual type of parallel-fibered bone, with large and randomly shaped osteocyte lacunae (otherwise typical of fibrous bone) and particular microanatomical features in Dallasaurus, which displays, rather than a spongious inner organization, bone mass increase in its humeri and a tubular organization in its femora and ribs.

Conclusions/Significance

The dominance of an unusual type of parallel-fibered bone suggests growth rates and, by extension, basal metabolic rates intermediate between that of the extant leatherback turtle, Dermochelys, and those suggested for plesiosaur and ichthyosaur reptiles. Moreover, the microanatomical features of the relatively primitive genus Dallasaurus differ from those of more derived mosasaurines, indicating an intermediate stage of adaptation for a marine existence. The more complete image of the various microanatomical trends observed in mosasaurine skeletal elements supports the evolutionary convergence between this lineage of secondarily aquatically adapted squamates and cetaceans in the ecological transition from a coastal to a pelagic lifestyle.     

Ichthyosauria from the Upper Lias of Strawberry Bank,England

Abstract: Eight remarkably preserved specimens of ichthyosaurs from the lower Toarcian (Lower Jurassic) of Strawberry Bank (Ilminster, Somerset, England) are described fully for the first time. Whereas previously these ichthyosaurs were assigned to one species, Stenopterygius hauffianus, our study shows there are two, Stenopterygius triscissus and Hauffiopteryx typicus. S. triscissus is a small‐ to medium‐sized ichthyosaur up to 3.5 m long, characterised by three apomorphies: long and slender rostrum, large elliptical supratemporal fenestra and bipartite pelvis. H. typicus is a small ichthyosaur up to 2.95 m long, with five apomorphies: short and extremely slender rostrum, very large orbit, small rounded supratemporal fenestra and tripartite pelvis, which is fused distally. Cladistic analysis is equivocal about their relationships, suggesting either that Hauffiopteryx and perhaps also Stenopterygius are members of a clade Eurhinosauria or that this clade does not exist, and both genera are members of a wider clade Thunnosauria. Further, the clade Stenopterygiidae, in which Hauffiopteryx had been located, is not identified. Most striking is that the specimens are all juveniles (five specimens) or infants (three specimens), ranging from one‐tenth to one‐half the normal adult length of the species.     

The skull of a large Lower Triassic ichthyosaur from Spitzbergen and its implications for the origin of the Ichthyosauria

An incomplete ichthyosaur skull from the Lower Triassic (Spathian) Sticky Keep Formation of Spitzbergen is one of the earliest ichthyosaurs known and by far the largest of the most basal ichthyosaurs. The specimen shows an extremely plesiomorphic cheek region of the skull and allows for the first time a confident reconstruction of the grundplan condition of the lower cheek region of ichthyosaurs. It unequivocally demonstrates the presence of a deep emargination of the lower cheek, which is bordered dorsally by the quadratojugal, squamosal and jugal, and therefore a plesiomorphic lack of contact between jugal and quadratojugal. The specimen also allows an evaluation of recent skull reconstructions of several Lower Triassic taxa which probably contain severe misinterpretations. The structure of the temporal and cheek region of the specimen can be most parsimoniously explained by the initial presence of a lower temporal fenestra. This provides important new evidence that the ancestry of ichthyosaurs might have lain within the Diapsida or their stem-group.     

Late Jurassic ichthyosaurs from the Neuquén Basin,Argentina

Late Jurassic ichthyosaurs are well represented in the Tithonian of the Neuquén Basin, in northwestern Patagonia, Argentina. Most of the ichthyosaur material from the Neuquén Basin was originally identified as Ophthalmosaurus. Recently, the new ichthyosaur genus Caypullisaurus was described, based on an almost complete mature specimen from Cerro Lotena. Some material previously referred to Ophlhalmosaurus has been referred to the new genus. However, both genera are present in the Tithonian of the Neuquén Basin. The discovery of an articulated forefin in Cajón de Almanza (near Loncopue, Neuquén) confirms the presence of Ophthalmosaurus in the uppermost Tithonian of the Neuquén Basin.     

On the purported presence of fossilized collagen fibres in an ichthyosaur and a theropod dinosaur

Since the discovery of exceptionally preserved theropod dinosaurs with soft tissues in China in the 1990s, there has been much debate about the nature of filamentous structures observed in some specimens. Sinosauropteryx was the first non‐avian theropod to be described with these structures, and remains one of the most studied examples. Despite a general consensus that the structures represent feathers or feather homologues, a few identify them as degraded collagen fibres derived from the skin. This latter view has been based on observations of low‐quality images of Sinosauropteryx, as well as the suggestion that because superficially similar structures are seen in Jurassic ichthyosaurs they cannot represent feathers. Here, we highlight issues with the evidence put forward in support of this view, showing that integumentary structures have been misinterpreted based on sedimentary features and preparation marks, and that these errors have led to incorrect conclusions being drawn about the existence of collagen in Sinosauropteryx and the ichthyosaur Stenopterygius. We find that there is no evidence to support the idea that the integumentary structures seen in the two taxa are collagen fibres, and confirm that the most parsimonious interpretation of fossilized structures that look like feather homologues in Sinosauropteryx is that they are indeed the remains of feather homologues.     

The Liassic ichthyosaur<Emphasis Type="Italic">Stenopterygius</Emphasis> cf.<Emphasis Type="Italic">quadriscissus</Emphasis> from the lower Toarcian of Dobbertin (northeastern Germany) and some considerations on lower Toarcian marine reptile palaeobiogeography

An incomplete skull of the lower Toarcian ichthyosaurStenopterygius cf.quadriscissus is described from the lower Toarcian of Dobbertin (Mecklenburg, northeastern Germany). It represents both the northeasternmost occurrence of this ichthyosaur genus and the first diagnostic specimen from East Germany. It therefore extends the palaeobiogeographic range ofStenopterygius considerably and demonstrates that this ichthyosaur also inhabited the Germanic Basin east of the Rhenish Massif by Early Jurassic times. A palaeobiogeographical pattern is evident within the genusStenopterygius, with the ubiquitous speciesS. longifrons andS. hauffianus on one hand, andS. megalorhinus andS. quadriscissus on the other hand, which appear not to have ranged northwest of the London-Brabant Massif. It is suggested that, whereas the Rhenish Massif was not an effective barrier for dispersal of ichthyosaurs in Western Europe during early Toarcian times, the London-Brabant Massif played a rather significant role, as is also shown by the fossil record of other marine reptile groups. A provincialism of early Toarcian marine reptiles is suggested for Western Europe, with a northwestern province which contains the British occurrences, an intermediate, Subgermanic province in France and the Benelux countries, and a southeastern province in the Germanic Basin. The British and Germanic provinces are each characterized by a typical assemblage of ichthyosaurs, plesiosaurs, and marine crocodiles, wheres the intermediate Subgermanic region shows an intermingling of faunal elements.      

Osteohistological insight into the early stages of growth in Mussaurus patagonicus (Dinosauria,Sauropodomorpha)

Here, we describe the bone histology of juvenile specimens of the basal sauropodomorph Mussaurus patagonicus and interpret its significance in terms of the early growth dynamics of this taxon. Thin sections from three juvenile specimens (femur length, 111–120 mm) of Mussaurus were analysed. The sampled bones consist of multiple postcranial elements collected from the Late Triassic Laguna Colorada Formation (El Tranquilo Group, Patagonia). The cortical bone is composed of fibrolamellar bone tissue. Vascularisation is commonly laminar or plexiform in the long bones. Growth marks are absent in all the examined samples. The ‘epiphyses’ of long bones are all formed by well-developed hypertrophied calcified cartilage. The predominance of woven-fibred bone matrix in cortical bones indicates a fast growth rate in the individuals examined. Moreover, given the existence of growth marks in adult specimens of Mussaurus, as in other sauropodomorphs, and assuming that the first lines of arrested growth was formed during the first year of life, the absence of growth marks in all the bones suggest that the specimens died before reaching their first year of life. Compared with the African taxon Massospondylus carinatus (another basal sauropodomorph for which the bone histology has been previously studied), it appears that Mussaurus had a higher early growth rate than Massospondylus.     

Long Bone Histology of Sauropterygia from the Lower Muschelkalk of the Germanic Basin Provides Unexpected Implications for Phylogeny

Background

Sauropterygia is an abundant and successful group of Triassic marine reptiles. Phylogenetic relationships of Triassic Sauropterygia have always been unstable and recently questioned. Although specimens occur in high numbers, the main problems are rareness of diagnostic material from the Germanic Basin and uniformity of postcranial morphology of eosauropterygians. In the current paper, morphotypes of humeri along with their corresponding bone histologies for Lower to Middle Muschelkalk sauropterygians are described and interpreted for the first time in a phylogenetic context.

Methodology/Principal Findings

Nothosaurus shows a typical plesiomorphic lamellar-zonal bone type, but varying growth patterns and the occurrence of a new humerus morphotype point to a higher taxonomic diversity than was known. In contrast to the enormous morphological variability of eosauropterygian humeri not assigned to Nothosaurus, their long bone histology is relatively uniform and can be divided into two histotypes. Unexpectedly, both of these histotypes reveal abundant fibrolamellar bone throughout the cortex. This pushes the origin of fibrolamellar bone in Sauropterygia back from the Cretaceous to the early Middle Triassic (early Anisian). Histotype A is assigned to Cymatosaurus, a basal member of the Pistosauroidea, which includes the plesiosaurs as derived members. Histotype B is related to the pachypleurosaur Anarosaurus. Contrary to these new finds, the stratigraphically younger pachypleurosaur Neusticosaurus shows the plesiomorphic lamellar-zonal bone type and an incomplete endochondral ossification, like Nothosaurus.

Conclusions/Significance

Histological results hypothetically discussed in a phylogenetical context have a large impact on the current phylogenetic hypothesis of Sauropterygia, leaving the pachypleurosaurs polyphyletic. On the basis of histological data, Neusticosaurus would be related to Nothosaurus, whereas Anarosaurus would follow the pistosaur clade. Furthermore, the presence of fibrolamellar bone, which is accompanied with increased growth rates and presumably even with increased metabolic rates, already in Anarosaurus and Cymatosaurus can explain the success of the Pistosauroidea, the only sauropterygian group to survive into the Jurassic and give rise to the pelagic plesiosaur radiation.     

Short-snouted toothless ichthyosaur from China suggests Late Triassic diversification of suction feeding ichthyosaurs

Background

Ichthyosaurs were an important group of Mesozoic marine reptiles and existed from the Early Triassic to the early Late Cretaceous. Despite a great diversity in body shapes and feeding adaptations, all share greatly enlarged eyes, an elongated rostrum with numerous conical teeth, and a streamlined body.

Methodology/Principal Findings

Based on new material from China and the restudy of Shastasaurus pacificus, we here reinterpret the classical large-bodied Late Triassic ichthyosaur genus Shastasaurus to differ greatly from the standard ichthyosaurian body plan, indicating much greater morphological diversity and range of feeding adaptations in ichthyosaurs than previously recognized. Phylogenetic analysis indicates a monophyletic clade consisting of the giant Shonisaurus sikanniensis, Guanlingsaurus liangae, and Shastasaurus pacificus to which the genus name Shastasaurus is applied. Shastasaurus liangae comb. nov. is from the Late Triassic (Carnian) Xiaowa Formation of Guizhou Province, southwestern China. The species combines a diminutive head with an entirely toothless and greatly reduced snout. The species also has by far the highest vertebral count among ichthyosaurs (86 presacral vertebrae and >110 caudal vertebrae), a count that is also very high for tetrapods in general. A reduced toothless snout and a diminutive head is also apparently present in the giant S. sikanniensis and presumably in S. pacificus.

Conclusions/Significance

In analogy to many modern odontocetes, Shastasaurus is interpreted as a specialized suction feeder on unshelled cephalopods and fish, suggesting a unique but widespread Late Triassic diversification of toothless, suction-feeding ichthyosaurs. Suction feeding has not been hypothesized for any of the other diverse marine reptiles of the Mesozoic before, but in Shastasaurus may be linked to the Late Triassic minimum in atmospheric oxygen.