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Early Cretaceous ichthyosaur diversity has markedly increased in recent years with the discovery of new taxa in the field and in museum collections. This has led to new characters, and a better und...

The postcranial axial skeleton of actinopterygian fishes is typically divided into three regions: (1) an anterior abdominal region, (2) a posterior caudal region and (3) those vertebrae supporting the caudal fin. However, in some actinopterygians, the axial skeleton is more finely subdivided, with up to six morphologically distinctsub-regions recognized. Phylogenetic continuity and homology of structures across these sub-regions have not been investigated in detail, either between or among groups. We examine variation in axial regionalization in saurichthyid fishes, a clade of extinct non-teleostean actinopterygians with highly variable axial skeletal morphology but an otherwise conservative body plan, and compare these findings to other non-teleostean actinopterygians to assess conservation of a regionalized axial skeleton within bony fishes. We document up to eight distinct regions in the vertebral column of Triassic Saurichthys: (1) a postoccipital region, (2) an anterior and (3) a posterior abdominal region, (4) a transitional region spanning the abdominal– caudal boundary, (5) an anterior and (6) a posterior caudal region and (7) preural and (8) ural regions. Based on taphonomical and morphological evidence, the transitional region appears to function in axial stiffening in the area of the median fins, whereas the abdominal region is highly flexible. The degree to which these axial regions are osteologically differentiated is highly variable across Saurichthyidae, implying iterative evolution of differentiation and de-differentiation over relatively short geological timescales. Such variably expressed regionalization was also identified in the outgroup non-teleostean actinopterygians Birgeria and Australosomus. Despite variation in morphological disparity, the regions identified in saurichthyids correlate well with those documented in some teleosts and Paleozoic actinopterygians, suggesting potential deep patterning homology but independent evolution of specific regionalized axial morphologies in response to changing functional demands.

Bite traces on fossil bones are key to deciphering feeding ecology and trophic interactions of vertebrate past ecosystems. However, similarities between traces produced by different carnivorous taxa with similar dentitions, and misidentifications due to equifinality, hinder confident identifications of the bite makers. Here, we correlate bite traces with macroscopic wear and microanatomy of the teeth of the pseudosuchian archosaur Batrachotomus kupferzellensis from the Triassic Lower Keuper fossil lagerstätten (southern Germany), untangling its feeding habits and shedding light on the bite traces generated by ziphodont teeth (teeth with serrated carinae). Individually, bite traces reflect tooth morphology, whereas composite bite traces and their frequency are related to feeding behaviour and explain tooth macroscopic wear and microanatomy. Therefore the identification of the bite maker is possible by analysing composite bite traces, their location on bones, and their relative abundance. In addition, tooth macroscopic wear and microanatomy are proven as independent lines of evidence of feeding ecology. Comparing bite traces on fossil and present-day bone assemblages, we observe that bone modifications by the crocodylomorph lineage (from Triassic pseudosuchian archosaurs to extinct and extant crocodylians) are strikingly similar, including taxa with and without ziphodont teeth. Such a set of features differs from bone modification assemblages produced by taxa with similar ziphodont teeth outside the pseudosuchian lineage, such as theropod dinosaurs and the Komodo monitor, suggesting phylogeny is a better predictor of feeding ecology among saurian reptiles than tooth morphology.

Fil: Fernandez, Marta S.. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Division Paleontologia Vertebrados; Argentina

Viviparity has evolved independently at least 12 times in ray-finned fishes. However, the fossil record of actinopterygian viviparity is poor, with only two documented occurrences. Both of these are from the non-teleost actinopterygian Saurichthys, and include S. curionii and S. macrocephalus from the Middle Triassic Meride Limestone (Monte San Giorgio, Switzerland). Here, we present new data on the reproductive biology of these species, giving unprecedented insights into their life-history. Based on positional and preservational criteria, six specimens were identified as unambiguously gravid. Embryos were positioned dorsal to the gastrointestinal tract, parallel to the axial skeleton and to each other, in the posterior two-thirds of the abdominal region. A minimum of 16 embryos are preserved in the most fecund females and, based on the largest preserved embryos and smallest preserved neonates, birth must have occurred at 7–12% of maternal fork length. Embryonic crania and teeth are relatively well-ossified, however ossification of the parietal region is delayed. In the postcranium, the median scale rows and lepidotrichia are ossified, but not the lateral scale rows. Ossified squamation and gradual allometric growth suggests that neonates did not undergo metamorphosis and were relatively precocial. When considered in a phylogenetic context, neither live birth nor internal fertilization appears to represent the primitive state for saurichthyid fishes. Key words: Actinopterygii, Saurichthys, Triassic, Monte San Giorgio, viviparity, allometry

Saurichthyidae are a family of elongate, piscivorous actinopterygian fishes with a stratigraphic range extending from the late Permian to the Middle Jurassic. There are four recognized Early Jurassic species, all from Europe and all referred to the genus Saurorhynchus. Only a single non-European Jurassic occurrence has been reported, a partial Saurorhynchus skull from the Fernie Formation of western Alberta, Canada. Here, we evaluate the Saurorhynchus material from Canada, including a new skull from Ya Ha Tinda, Alberta, and a fragmentary mandible from Canyon Creek, Alberta. All diagnostic Canadian Saurorhynchus specimens are consistent with the European species Saurorhynchus acutus, on the basis of external narial morphology and reduced dermal ornamentation. Saurorhynchus acutus was present in the Toarcian of western North America from the tenuicostatum Zone to the late serpentinum – early bifrons Zone, similar to the stratigraphic range documented in Europe. The absence of divergence between eastern Panthalassan and western Tethyan saurichthyids is surprising, given that divergence has been well-documented in coeval invertebrates. Saurorhynchus may have had slower rates of morphological evolution than contemporaneous marine invertebrates; alternatively greater mobility, broader environmental tolerance, or both in these fish may have allowed ongoing gene flow between the two oceanic basins in the early Toarcian.

Changing predator-prey interactions during the Mesozoic Marine Revolution (MMR) profoundly altered the trajectory of marine tetrapod evolution. Here, we assess potential signatures of this landmark transition through the fossil record of skeletal pathologies in ichthyosaurs — iconic marine reptiles that developed increasingly ‘fish-like’ body plans over time. We surveyed a stratigraphically constrained sample of 200 Middle Triassic ichthyosaur specimens and compared the type, distribution and prevalence of pathologies with an approximately equivalent assemblage of Early Jurassic age. Overall, skeletal pathologies were equally prevalent in these groups, and most often manifested in species >4 m long. However, pathological bones were found to be concentrated in the hind limbs and tail of Triassic ichthyosaurs, whereas the jaws, forelimbs, and ribcage were preferentially affected in Jurassic taxa. We posit that the occurrence of ankylosed zygapophyses in the caudal peak of Triassic ichthyosaurs could represent a functional by-product of their primitive ‘eel-like’ swimming. Conversely, increased instances of broken ribs in Jurassic ichthyosaurs may infer ramming or tail strike behaviours that characterise morphologically ‘fish-like’ marine tetrapods, such as modern toothed whales. Different categories of skeletal pathologies thus evidently reflect structural modifications in the ichthyosaur body plan, and indirectly coincide with ecological turnover during the MMR.

The Oligocene was a period of profound climatic and biotic changes, coinciding with a shift from a mostly ice-free warmhouse world at the Eocene-Oligocene boundary to a globally cooler, more seasonal climate. The Rauenberg locality (Baden-Wurttemberg, Germany) is one of the most significant early Oligocene fossil assemblages in Europe, containing both marine and terrestrial elements of fauna and flora. Preservation is often superb and comprises complete and articulated skeletons with soft tissue preservation. The diverse assemblage provides critical insights into the paleoenvironment of the Upper Rhine Valley. We reassess diversity at the locality, resulting in a list of 302 taxa found at the site, 207 of which are marine (52% of taxa represented by macrofossils). The rest are interpreted as originating from the coastal environment. Molluscan, echinoderm, and plant macrofossil diversity are assessed here for the first time. Based on these new results, we interpret Rauenberg as representing a fully marine assemblage deposited in a moderately shallow, low-energy tropical-subtropical environment. Productivity was high, and seafloor anoxia was intermittently developed. There is no evidence for long-term brackish influence or mangrove swamps, and no direct evidence for the development of seagrass meadows. On land, warm, frost-free conditions permitted the development of prevailingly evergreen sclerophyllous broad-leaved forests along with pine and palm-rich coastal forests on sandy soils. The marine invertebrate fauna shows more northerly affinities, whereas the vertebrate fauna is distinctly Paratethyan.

The first ophthalmosaurid ichthyosaur from the Upper Jurassic deposits of the Central–Northern Apennines (Marche, Italy) is here described for the first time. The specimen is relatively complete and is referred toGengasaurus nicosiaigen. et sp. nov. based on a unique combination of characters, including a peculiar condition of the preaxial accessory facet on the humerus. The faunal association of the ichthyosaur-bearing level indicates a late Kimmeridgian – earliest Tithonian age, and its finding contributes significantly to our knowledge of the diversity of Late Jurassic ichthyosaurs from the Western Tethys. Two shark teeth assigned to the order Hexanchiformes were also recovered in association with the ichthyosaur specimen, suggesting that scavenging of the carcass might have occurred.Gengasauruscan be referred to Ophthalmosauridae based on the reduced extracondylar area of the basioccipital, and the presence of a preaxial digit. It differs fromOphthalmosaurusspp. in several respects, including the shape of the posterior basisphenoid, the shape of the supraoccipital, the anteriorly deflected preaxial facet of the humerus, and a proximodistally shortened ulna. The new taxon actually shares diagnostic characters with both members of the two main lineages recovered in previous phylogenetic analyses, more nested within Ophthalmosauridae. The affinities ofGengasaurusto genera from both the northern and southern hemispheres also suggest that connectivity between pelagic habitats was high during the early Late Jurassic, allowing dispersal of some forms, followed by local, endemic divergence.

Parental care is widespread among vertebrates and the observed patterns of parental care and investment are extremely diverse. Among amphibians, caecilians (Gymnophiona) exhibit considerable variation in reproductive modes, including both oviparity and viviparity, combined with highly unusual investment strategies (e.g. skin-feeding and intrauterine feeding). In the present study, current knowledge on the reproductive modes is integrated into an analysis of the evolutionary scenario of parental investment of caecilians. Phylogenetically basal caecilians possessing a biphasic life cycle that includes an aquatic larval stage invest in macrolecithal eggs directly corresponding to size at hatching. Some phylogenetically derived caecilians (i.e. the Teresomata) have a smaller clutch size and show a reduction to either medium-yolked (mesolecithal) or small-yolked (microlecithal) eggs. Via alternative pathways of parental investment, such as intrauterine feeding in viviparous taxa and maternal dermatotrophy in oviparous taxa, teresomatan caecilians increase both offspring size and quality. However, more data regarding reproductive biology are needed to obtain a fully resolved understanding of the evolution of reproduction in caecilian amphibians.

Reconstructing ecological niche shifts during ontogeny in extinct animals with no living analogues is difficult without exceptional fossil collections. Here we demonstrate how a previously identified ontogenetic shift in the size and shape of the dentition in the early Toarcian ichthyosaur Stenopterygius quadriscissus accurately predicts a particular dietary shift. The smallest S. quadriscissus fed on small, burst-swimming fishes, with a steady shift towards faster moving fish and cephalopods with increasing body size. Larger adult specimens appear to have been completely reliant on cephalopods, with fish completely absent from gut contents shortly after onset of sexual maturity. This is consistent with a previously proposed ontogenetic niche shift based on tooth shape and body size, corroborating the idea that dental ontogeny may be a useful predictor of dietary shifts in marine reptiles. Applying the theoretical framework used here to other extinct species will improve the resolution of palaeoecological reconstructions, where appropriate sample sizes exist.

Among the numerous ichnofossils found in the Upper Jurassic Solnhofen-type Nusplingen Fossillagerstatte of southwestern Germany, there is a unique find of a fish associated with a trail, which must be interpreted as a mortichnium. Like similar cases of fatally injured crustaceans from the same locality, this fish mortichnium resulted from an unsuccessful attack on the fish by a predator and does not provide evidence for a hostile environment.

Saurichthyidae, a speciose group of Early Mesozoic actinopterygian fishes, was previously documented as ranging from the Late Permian to the end of the Early Jurassic. Saurichthyids were most diverse in the Triassic: only two species persisted into the Early Jurassic, restricted to marine sediments in Europe and North America. Here, I report previously undescribed specimens from the Middle Jurassic of the District of Goppingen, Baden-Wurttemberg, Germany. The new material consists of a partial skull, and a second skull with associated lower jaw, both from lower Aalenian sediments of the marine Opalinuston Formation. Although poor preservation prevents referral to species, these specimens are consistent with the Early Jurassic taxon Saurorhynchus based on the elongate jaws anterior to the orbit, organization of the dentition, and structure and ornamentation of the lateral mandible. These finds extend the stratigraphic range of Saurichthyidae into the Middle Jurassic, eliminating any connection between the extinction of this family and the early Toarcian Oceanic Anoxic Event and supporting minimal faunal change across the Toarcian–Aalenian boundary.

The Early Jurassic Toarcian Oceanic Anoxic Event is considered one of the most dramatic environmental perturbations of the Mesozoic. An elevated extinction rate among marine invertebrates accompanied rapid environmental changes, but effects on large vertebrates are less understood. We examined changes in ichthyosaur body size in the Posidonia Shale of the Southwest German Basin spanning the extinction interval to assess how environmental changes and biotic crisis among prey species affected large reptiles. We report no species-level extinction among the ichthyosaurs coinciding with peak invertebrate extinction. Large ichthyosaurs were absent from the fauna during the extinction interval, but became more abundant in the immediate aftermath.Stenopterygius quadriscissus, the most abundant species during the extinction interval, increased in body size after the biotic event. Rapid invasion by large taxa occurred immediately following the extinction event at the end of the first ammonite zone of the early Toarcian. Greater mobility permitting exploitation of ephemeral resources and opportunistic feeding behavior may minimize the impacts of environmental change on large vertebrates.

Cretaceous ichthyosaurs were relatively diverse in temperate latitudes, but few species have been described from the palaeotropics. Here, we describe a new ophthalmosaurid ichthyosaur, Muiscasaurus catheti gen. et sp. nov., from the Barremian–Aptian aged Paja Formation of Colombia. This species is known only from a partial skull and differs from all other ichthyosaurs in the unusual configuration of the external narial opening, slender rostrum, narrow postorbital region, and gracile dentition. It is the second ichthyosaur described from the Paja Formation, suggesting moderate taxonomic and ecological ichthyosaur diversity in the region during the Early Cretaceous.

Saurichthys, characterized by a long slender body and an elongated rostrum, is one of the most iconic genera of Late Paleozoic–Early Mesozoic fishes. The genus was particularly speciose in the Triassic, with a global distribution in both marine and freshwater habitats. Here, we describe two new species from the Middle Triassic Besano Formation of Monte San Giorgio, Switzerland, Saurichthys breviabdominalis sp. nov. and Saurichthys rieppeli sp. nov. S. breviabdominalis is characterized by a proportionately long operculum, short abdominal region and rib-like mid-lateral scales, whereas S. rieppeli is divergent from other Middle Triassic saurichthyids in the block-like haemal arches, fringing fulcra on the pelvic and unpaired fins, and reduction of the squamation to a single row in the abdominal region. Phylogenetic analysis places S. rieppeli in a basal position relative to congeners from the Alpine Triassic, and supports previous hypotheses regarding the convergent evolution of reduced squamation within saurichthyids. S. breviabdominalis forms a monophyletic group with species from the same locality, suggesting divergence in sympatry. This finding has implications for our understanding of disparity and character evolution in saurichthyid fishes, as well as ecomorphological divergence and resource partitioning between closely related fishes in Triassic marine ecosystems. © 2015 The Linnean Society of London

Paleopathologies document skeletal damage in extinct organisms and can be used to infer the causes of injury, as well as aspects of related biology, ecology and behavior. To date, few studies have been undertaken on Jurassic marine reptiles, while ichthyosaur pathologies in particular have never been systematically evaluated. Here we survey 41 specimens of the apex predator ichthyosaur Temnodontosaurus from the Early Jurassic of southern Germany in order to document the range and absolute frequency of pathologies observed in this taxon as a function of the number of specimens examined. According to our analysis, most observed pathologies in Temnodontosaurus are force-induced traumas with signs of healing, possibly inflicted during aggressive interactions with conspecifics. When the material is preserved, broken ribs are correlated in most of the cases with traumas elsewhere in the skeleton such as cranial injuries. The range of cranial pathologies in Temnodontosaurus is similar to those reported for extinct cetaceans and mosasaurs, which were interpreted as traces of aggressive encounters. Nevertheless, Temnodontosaurus differs from these other marine amniotes in the absence of pathologies in the vertebral column, consistent with the pattern previously documented in ichthyosaurs. We did not detect any instances of avascular necrosis in Temnodontosaurus from southern Germany, which may reflect a shallow diving life style. This study is intended to provide baseline data for the various types of observed pathologies in large ichthyosaurs occupying the 'apex predator' niche, and potentially clarifies aspects of species-specific behavior relative to other ichthyosaurs and marine amniotes.

Ichthyosaurs, a lineage of extinct Mesozoic marine reptiles, have garnered attention in both the palaeontological and developmental literature for the unique limb morphology seen in derived genera. These morphologies include an increase in the number of phalanges per digit (hyperphalangy) and in the number of digits (hyperdactyly), but most interestingly also a shift in element identity. Elements distal to the stylopodium acquire characteristics of mesopodial elements, such as a rounded, nodular shape and a loss of perichondral bone on the anterior and posterior surfaces. Here, we examine numerous aspects of the loss of proximodistal identity in ichthyosaur limbs including phylogenetic progression of the loss of perichondral bone, histology and microstructure of the elements retaining perichondral bone in derived taxa, and correlates of intraspecific variation in degree of perichondral bone reduction in a derived ichthyosaur,Stenopterygius quadriscissus. Results show that loss of limb element identity occurred progressively over ichthyosaurian evolution, and the notches seen on the anterior surface of limb elements in derived ichthyosaurs are homologous to the long bone shafts in terrestrial tetrapods. Variation in the number of notches inS. quadriscissuscan best be explained through delayed ossification of the anterior perichondrium, indicating a heterochronic component to the loss of identity. We propose a developmental mechanism – gradual expansion of the polyalanine region of HoxD13 over ichthyosaurian evolution – to explain the progressive loss of limb regionalization as well as the heterochronic delay in perichondral ossification.

Ophthalmosauridae is a clade of derived thunniform ichthyosaurs that are best known from Callovian (late Middle Jurassic) to Cenomanian-aged (Late Cretaceous) deposits in both the Northern and Southern Hemispheres. Ophthalmosaurids arose prior to the Early–Middle Jurassic boundary, however, very little is known about their diversity and distribution in the earliest phase of their evolutionary history during the Aalenian–Bathonian (Middle Jurassic) interval. Here we describe new diagnostic ophthalmosaurid material from the Early Bajocian (Middle Jurassic) of Alaska. The specimen, UAMES 3411, is a partial disarticulated skull that was discovered in the Middle Jurassic Tuxedni Formation, which was deposited in shallow marine settings outboard of the then-accreting Wrangellia composite terrane. The new material is significant in that it is the first Jurassic ichthyosaur described from Alaska, one of the oldest ophthalmosaurids known and the only Middle Jurassic ophthalmosaurid described from the Northern Hemisphere. The new material adds to a rapidly growing data set on ophthalmosaurid diversity and suggests that the clade was geographically widespread by the Early Bajocian, very early in its evolutionary history.

An expedition to Melville Island in Nunavut, Canada, recovered the fragmentary fossils of several plesiosaurs from non-marine deposits of the Hauterivian–Aptian Isachsen Formation. These plesiosaur fossils are some of the oldest Early Cretaceous records of the group in North America, and they likely predate the formation of a continuous Western Interior Seaway. The plesiosaurs from Melville Island appear to be primarily juveniles, and would have been living in a region that experienced at least seasonally cool temperatures. The presence of these fossils in a fluvial deposit support previous suggestions that juvenile plesiosaurs may have preferentially inhabited shallower waters rather than open marine environments. These fossils also show that polycotylid plesiosaurs were able to rapidly disperse and colonize high latitude coastal regions, as their occurrence in Arctic Canada only slightly postdates the first confirmed appearance of the group in Australia.

Teleosauroid extinction at the Jurassic/Cretaceous boundary has been hypothesized to be an artifact of sampling only temperate paleolatitudes. here, we describe the definitive youngest record of Teleosauroidea, from the upper Barremian of the paleotropics. The preserved material consists of dorsal vertebrae, dorsal ribs, dorsal and ventral osteoderms and epipodial remains. We refer the specimen to Teleosauroidea based on the large hourglass-shaped amphiplatyan morphology of the dorsal centra and rectangular dorsal osteoderms, which are much wider than long and have a straight anterior edge. The South American specimen is one of the largest known teleosauroids, with an estimated body length of 9.6 m. This is the first evidence of a marine crocodylomorph recorded from the Paja Formation of Colombia. The survival of Teleosauroidea in the paleotropics of northern Gondwana following the group’s extinction in europe supports the hypothesis that water temperature played a role in controlling the diversity and distribution of these large marine predators. doi: 10.5710/AMGH.26.09.2019.3269

The fossil record of the ichthyosaur genus Stenopterygius comprises a large number of specimens of all ontogenetic stages. This makes the taxon a good model for ontogenetic studies in a Mesozoic ma...

International audience; The Posidonienschiefer Formation (Toarcian) of Holzmaden, Baden-Wurttemberg in southwestern Germany has yielded several excellently preserved plesiosaurian specimens and received considerable research attention. The plesiosaurians found within these deposits are always significantly outnumbered by ichthyosaurs, and close examination of these rare specimens is crucial to a better understanding of the diversity and palaeoecology of Plesiosauria in this very peculiar ecosystem. The plesiosaurian specimen SMNS 51945 found in this area is a juvenile individual consisting of a partial, crushed skull and an exquisitely preserved post-cranial skeleton. Its anatomical characters seem to differ from the long-necked plesiosauroids Microcleidus brachypterygius and Seeleyosaurus guilelmiimperatoris that are the most abundant taxa within the plesiosaurian assemblage. The post-cranial skeleton preserves very likely soft tissues composed of buff-coloured and dark-coloured structures around the vertebral column and hindlimb of the animal. A network of buff-coloured fibres located posterior to the hindlimb most likely represents phosphatised collagen fibres as already found in some ichthyosaur specimens, confirming that wing area in plesiosaurians was much larger than that suggested by skeletal remains alone. The specimen also contains gastroliths (sand-sized grains mainly composed of quartz) in the stomach cavity suggesting the animal spent at least some of its time in shallow coastal waters, tens or hundreds of kilometres from the final place of burial.

An incomplete rostrum of a platypterygiine ichthyosaur (IGVR 94574) has recently been found near Tregnago (Verona, Italy), in the Lessini Mts. This is the first Cretaceous ichthyosaurian fossil found in Italy outside the northern Apennines, where fragmentary remains have been recovered since the 19th century. IGVR 94574 is a mid-distal rostral fragment infilled by a white, fine-grained limestone matrix. It consists of the premaxilla, nasal, vomer, dentary, and splenial, together with 20 teeth. Even if the exact horizon of provenance is unknown, analysis of the microfossil content of the matrix (planktonic foraminifera and calcareous nannofossils) allowed us to assign it to the lower part of the upper Albian. This is also the first Cretaceous ichthyosaur in Italy with a precise age: all previously reported fossils from the Apennines could not be dated due to the complete absence of microfossils in the surrounding matrix. We assign IGVR 94572 to Platypterygiinae; generic specification is not possible pending the taxonomic revision of the genus Platypterygius. Finally, the Tregnago rostrum is strikingly similar to the specimen IPUM 30139 from Gombola (Modena, Italy), stored in the collections of the Paleontological Museum of Modena and Reggio Emilia University and also attributable to a platypterygiine ichthyosaur. The comparison between these two fossils reveals the teeth are very similar both in shape and size.

Understanding morphological integration is one of the central goals of evolutionary developmental biology. Despite its applicability to questions of paleontological interest, there are few studies on integration in fossil vertebrates. In this study, we examine limb integration in the Lower Jurassic ichthyosaurStenopterygius quadriscissus, with the aim of examining the effect of ontogeny and anagenetic changes over short geological time spans on metrics of limb integration. Both ontogenetic and stratigraphic effects had a significant influence on measured values of integration, the identity of strongly integrated elements, and some common ratio values such as the relative integration of the forelimb to the hind limb, or within-limb to between-limb integration. Ontogenetic effects were relatively greater, although this could be linked to sample size. Although adults showed the lowest levels of overall integration, they possessed high levels of integration between serially homologous elements, something that was unexpected due to strong divergence in limb size and perhaps functional differences in derived ichthyosaurs. Ontogenetic differences in the relative integration of the forelimb to the hind limb are probably related to early locomotor demands on the forelimb. We conclude that if samples are pooled, the resulting pattern of integration may not reflect any one subsample but will be a composite created through the superposition of several variables. Pooling data in paleontological studies of integration has a non-trivial effect on the results obtained.