The Evolution of Marine Reptiles

Reptiles have repeatedly invaded marine environments despite their physiological constraints as air breathers. Marine reptiles were especially successful in the Mesozoic as major predators in the sea. There were more than a dozen groups of marine reptiles in the Mesozoic, of which four had more than 30 genera, namely sauropterygians (including plesiosaurs), ichthyopterygians, mosasaurs, and sea turtles. Medium-sized groups, such as Thalattosauria and Thalattosuchia, had about ten genera, whereas small groups, such as Hupehsuchia and Pleurosauridae, consisted of only two genera or less. Sauropterygia and Ichthyopterygia were the two longest surviving lineages, with 185 and 160 million years of stratigraphic spans, respectively. Mesozoic marine reptiles explored many different swimming styles and diets. Their diet included fish, cephalopods, other vertebrates, and hard-shelled invertebrates, whereas no herbivore is known at this point. Sauropterygians and ichthyopterygians gave rise to cruising forms that probably invaded outer seas. Intermediate forms that led to the cruising species are known in Ichthyopterygia but not as much in Sauropterygia. Discovery of new fossils should eventually reduce the gap in the fossil record.     

The locomotor ecomorphology of Mesozoic marine reptiles

The aftermath of the end-Permian mass extinction provided ecological opportunities for many groups of reptiles, marking the beginning of reptile dominance of the Mesozoic oceans. Clades such as ichthyosaurs, thalattosuchians, sauropterygians, mosasaurs and turtles evolved a remarkable diversity of ecological niches and became important components of aquatic ecosystems. Locomotion is a key aspect of ecology, crucial for many biological functions such as foraging and migration. However, the evolution of locomotory adaptations across all Mesozoic marine reptiles remains poorly understood. Here we present multivariate and disparity analyses based on body proportions, body size and post-cranial proxies for locomotion in 125 species of Mesozoic marine reptiles. Our analysis highlights key anatomical transformations in the evolution of swimming modes, characterizing two divergent evolutionary paths in the transition from drag-based to lift-based propulsion in both the axial and appendicular spectrum. Analyses against geological time do not show evidence for an explosive radiation after the end-Permian extinction, pointing instead to a gradual increase in locomotory disparity during the whole Mesozoic, which reached the highest levels in the Cretaceous. Our analysis also provides insight into the evolution of locomotion in particular clades. Some notable findings are the high aquatic specialization in the earliest ichthyosauromorphs and the morphospace overlap between mosasauroids and ichthyosauromorphs.     

A Giant Chelonioid Turtle from the Late Cretaceous of Morocco with a Suction Feeding Apparatus Unique among Tetrapods

Background

Secondary adaptation to aquatic life occurred independently in several amniote lineages, including reptiles during the Mesozoic and mammals during the Cenozoic. These evolutionary shifts to aquatic environments imply major morphological modifications, especially of the feeding apparatus. Mesozoic (250–65 Myr) marine reptiles, such as ichthyosaurs, plesiosaurs, mosasaurid squamates, crocodiles, and turtles, exhibit a wide range of adaptations to aquatic feeding and a broad overlap of their tooth morphospaces with those of Cenozoic marine mammals. However, despite these multiple feeding behavior convergences, suction feeding, though being a common feeding strategy in aquatic vertebrates and in marine mammals in particular, has been extremely rarely reported for Mesozoic marine reptiles.

Principal Findings

A relative of fossil protostegid and dermochelyoid sea turtles, Ocepechelon bouyai gen. et sp. nov. is a new giant chelonioid from the Late Maastrichtian (67 Myr) of Morocco exhibiting remarkable adaptations to marine life (among others, very dorsally and posteriorly located nostrils). The 70-cm-long skull of Ocepechelon not only makes it one of the largest marine turtles ever described, but also deviates significantly from typical turtle cranial morphology. It shares unique convergences with both syngnathid fishes (unique long tubular bony snout ending in a rounded and anteriorly directed mouth) and beaked whales (large size and elongated edentulous jaws). This striking anatomy suggests extreme adaptation for suction feeding unmatched among known turtles.

Conclusion/Significance

The feeding apparatus of Ocepechelon, a bony pipette-like snout, is unique among tetrapods. This new taxon exemplifies the successful systematic and ecological diversification of chelonioid turtles during the Late Cretaceous. This new evidence for a unique trophic specialization in turtles, along with the abundant marine vertebrate faunas associated to Ocepechelon in the Late Maastrichtian phosphatic beds of Morocco, further supports the hypothesis that marine life was, at least locally, very diversified just prior to the Cretaceous/Palaeogene (K/Pg) biotic crisis.     

Pterosaur dietary hypotheses: a review of ideas and approaches

Pterosaurs are an extinct group of Mesozoic flying reptiles, whose fossil record extends from approximately 210 to 66 million years ago. They were integral components of continental and marginal marine ecosystems, yet their diets remain poorly constrained. Numerous dietary hypotheses have been proposed for different pterosaur groups, including insectivory, piscivory, carnivory, durophagy, herbivory/frugivory, filter‐feeding and generalism. These hypotheses, and subsequent interpretations of pterosaur diet, are supported by qualitative (content fossils, associations, ichnology, comparative anatomy) and/or quantitative (functional morphology, stable isotope analysis) evidence. Pterosaur dietary interpretations are scattered throughout the literature with little attention paid to the supporting evidence. Reaching a robustly supported consensus on pterosaur diets is important for understanding their dietary evolution, and their roles in Mesozoic ecosystems. A comprehensive examination of the pterosaur literature identified 314 dietary interpretations (dietary statement plus supporting evidence) from 126 published studies. Multiple alternative diets have been hypothesised for most principal taxonomic pterosaur groups. Some groups exhibit a high degree of consensus, supported by multiple lines of evidence, while others exhibit less consensus. Qualitative evidence supports 87.3% of dietary interpretations, with comparative anatomy most common (62.1% of total). More speciose groups of pterosaur tend to have a greater range of hypothesised diets. Consideration of dietary interpretations within alternative phylogenetic contexts reveals high levels of consensus between equivalent monofenestratan groups, and lower levels of consensus between equivalent non‐monofenestratan groups. Evaluating the possible non‐biological controls on apparent patterns of dietary diversity reveals that numbers of dietary interpretations through time exhibit no correlation with patterns of publication (number of peer‐reviewed publications through time). 73.8% of dietary interpretations were published in the 21st century. Overall, consensus interpretations of pterosaur diets are better accounted for by non‐biological signals, such as the impact of the respective quality of the fossil record of different pterosaur groups on research levels. That many interpretations are based on qualitative, often untestable lines of evidence adds significant noise to the data. More experiment‐led pterosaur dietary research, with greater consideration of pterosaurs as organisms with independent evolutionary histories, will lead to more robust conclusions drawn from repeatable results. This will allow greater understanding of pterosaur dietary diversity, disparity and evolution and facilitate reconstructions of Mesozoic ecosystems.     

Absence of Suction Feeding Ichthyosaurs and Its Implications for Triassic Mesopelagic Paleoecology

Mesozoic marine reptiles and modern marine mammals are often considered ecological analogs, but the extent of their similarity is largely unknown. Particularly important is the presence/absence of deep-diving suction feeders among Mesozoic marine reptiles because this would indicate the establishment of mesopelagic cephalopod and fish communities in the Mesozoic. A recent study suggested that diverse suction feeders, resembling the extant beaked whales, evolved among ichthyosaurs in the Triassic. However, this hypothesis has not been tested quantitatively. We examined four osteological features of jawed vertebrates that are closely linked to the mechanism of suction feeding, namely hyoid corpus ossification/calcification, hyobranchial apparatus robustness, mandibular bluntness, and mandibular pressure concentration index. Measurements were taken from 18 species of Triassic and Early Jurassic ichthyosaurs, including the presumed suction feeders. Statistical comparisons with extant sharks and marine mammals of known diets suggest that ichthyosaurian hyobranchial bones are significantly more slender than in suction-feeding sharks or cetaceans but similar to those of ram-feeding sharks. Most importantly, an ossified hyoid corpus to which hyoid retractor muscles attach is unknown in all but one ichthyosaur, whereas a strong integration of the ossified corpus and cornua of the hyobranchial apparatus has been identified in the literature as an important feature of suction feeders. Also, ichthyosaurian mandibles do not narrow rapidly to allow high suction pressure concentration within the oral cavity, unlike in beaked whales or sperm whales. In conclusion, it is most likely that Triassic and Early Jurassic ichthyosaurs were ‘ram-feeders’, without any beaked-whale-like suction feeder among them. When combined with the inferred inability for dim-light vision in relevant Triassic ichthyosaurs, the fossil record of ichthyosaurs does not suggest the establishment of modern-style mesopelagic animal communities in the Triassic. This new interpretation matches the fossil record of coleoids, which indicates the absence of soft-bodied deepwater species in the Triassic.     

Der Frühe und Mittlere Jura. Lebensraum Jurameer

Central Europe preserves sedimentary rocks of Jurassic age, deposited in a subtropical sea, which permit insights into Mesozoic marine ecosystems by their extensive fossil record. Calcareous plankton contributed to the formation of marine sediments for the first time and became an important factor in the global Carbon cycle ever since. Ammonites, Belemnites, fish and marine reptiles shifted through the open water, while a varied fauna dwelled upon and borrowed within the sea floor. Faunal associations as well as depositional environments changed, however, and as these changes are recorded in the rocks, they help to reconstruct the vicissitudes of a mesozoic sea, its changing sea level and current patterns, and what influence they had for the evolution of marine faunas.     

Palaeopathology and injury in the extinct mosasaurs (Lepidosauromorpha, Squamata) and implications for modern reptiles

Three fossilized dentaries provide an insight into the healing of fractures in a major group of extinct marine predators, mosasaurs. The data has implications for modern day reptiles in which such information is scanty. All three dentaries have callus formation. Both dentaries of Mosasaurus hoffmanni show fracture non-union, possibly resulting from intervening tissue. They also show evidence of osteomyelitis. Bone remodeling is complete in an earlier fracture in one of the M. hoffmanni dentaries. Given that the most recent fracture (non-union) occurred at a new and much deeper part of the dentary and not in the region of the earlier fracture, it can be assumed that remodeled bone in mosasaurs (probably in reptiles generally) could withstand powerful stresses such as those encountered during predation and fighting. The splenial bone, attached in life to the dentary only by connective tissue, may have acted as a natural splint to immobilize the fracture to the dentary and maintain its alignment during healing. Possible explanations for the injuries are feeding on hard-shelled prey, such as turtles, and fighting.     

Trophic convergence drives morphological convergence in marine tetrapods

Marine tetrapod clades (e.g. seals, whales) independently adapted to marine life through the Mesozoic and Caenozoic, and provide iconic examples of convergent evolution. Apparent morphological convergence is often explained as the result of adaptation to similar ecological niches. However, quantitative tests of this hypothesis are uncommon. We use dietary data to classify the feeding ecology of extant marine tetrapods and identify patterns in skull and tooth morphology that discriminate trophic groups across clades. Mapping these patterns onto phylogeny reveals coordinated evolutionary shifts in diet and morphology in different marine tetrapod lineages. Similarities in morphology between species with similar diets—even across large phylogenetic distances—are consistent with previous hypotheses that shared functional constraints drive convergent evolution in marine tetrapods.     

Grazing bioerosion in Jurassic seas: a neglected factor in the Mesozoic marine revolution?

Grazing bioerosion, notably by chitons, gastropods and regular echinoids, is a powerful destructive force in many recent shallow-marine environments and impacts significantly on sessile epibionts through grazing predation and/or unselective dislodgement. Grazing bioerosion was an important component of a major phase of biotic escalation; the Mesozoic marine revolution. Recent investigations of hard substrates in southern British Jurassic marine formations have identified widespread ichnofossils attributable to grazing activity by gastropods and/or chitons, and regular echinoids. The co-occurring benthic macrofaunas include groups that would have been vulnerable to grazing disturbance and dislodgement; notably articulate brachiopods. The emerging ichnological evidence strengthens the argument for grazing bioerosion as a significant contributor to the Mesozoic–Cenozoic decline of the articulate brachiopods, and their retreat to deep-water and/or cryptic refugia.     

The evolution of the modern avian digestive system: insights from paravian fossils from the Yanliao and Jehol biotas

The avian digestive system, like other aspects of avian biology, is highly modified relative to other reptiles. Together these modifications have imparted the great success of Neornithes, the most diverse clade of amniotes alive today. It is important to understand when and how aspects of the modern avian digestive system evolved among neornithine ancestors in order to elucidate the evolutionary success of this important clade and to understand the biology of stem birds and their closest dinosaurian relatives: Mesozoic Paraves. Although direct preservation of the soft tissue of the digestive system has not yet been reported, ingested remains and their anatomical location preserved in articulated fossils hint at the structure of the digestive system and its abilities. Almost all data concerning direct evidence of diet in Paraves comes from either the Upper Jurassic Yanliao Biota or the Lower Cretaceous Jehol Biota, both of which are known from deposits in north-eastern China. Here, the sum of the data gleaned from the thousands of exceptionally well-preserved fossils of paravians is interpreted with regards to the structure and evolution of the highly modified avian digestive system and feeding apparatus. This information suggests intrinsic differences between closely related stem lineages implying either strong homoplasy or that diet in each lineage of non-ornithuromorph birds was highly specialized. Regardless, modern digestive capabilities appear to be limited to the Ornithuromorpha, although the complete set of derived feeding related characters is restricted to the Neornithes.     

Morphological and biochemical evidence for the evolution of salt glands in snakes

Vertebrate salt glands have evolved independently multiple times, yet there are few hypotheses about the processes underlying the convergent evolution of salt glands across taxa. Here, we compare the morphology and molecular biology of specialized salt-secreting glands from a marine snake (Laticauda semifasciata) with the cephalic glands from semi-marine (Nerodia clarkii clarkii) and freshwater (N. fasciata) watersnakes to look for evidence of a salt gland in the former and to develop hypotheses about the evolution of snake salt glands. Like the salt gland of L. semifasciata, the nasal and anterior/posterior sublingual glands in both species of Nerodia exhibit a compound tubular shape, and express basolateral Na(+)/K(+)-ATPase (NKA) and Na(+)/K(+)/2Cl(-)cotransporter (NKCC); however, the abundance of NKA and NKCC in N. fasciata appears lower than in N. c. clarkii. Aquaporin 3 (AQP3) is also basolateral in the sublingual glands of both species of Nerodia, as is abundant neutral mucin; both AQP3 and mucin are absent from the salt gland in L. semifasciata. Thus, we propose that the evolution of the snake salt gland by co-option of an existing gland involved at least two steps: (i) an increase in the abundance of NKA and NKCC in the basolateral membranes of the secretory epithelia, and (ii) loss of AQP3/mucus secretion from these epithelia.     

An enigmatic marine reptile,Hispaniasaurus cranioelongatus (gen. et sp. nov.) with nothosauroid affinities from the Ladinian of the Iberian Range (Spain)

An incomplete skull of a marine reptile with an atypical elongation of the postorbital region is described. The find comes from the Muschelkalk facies (Cañete Formation) of the Villora section (Iberian Range, Cuenca Province, Spain), characterised by a shallow marine (intertidal) environment and dated as Ladinian in age. The small skull has a rectangular shape, lacking, as preserved, upper temporal openings and a parietal foramen. The upper temporal openings might be secondarily closed. However, the absence of a parietal foramen and squamosals in the preserved part and the incompleteness of the pterygoids make a posteriorly postponed location of the upper temporal openings also conceivable. Teeth are all broken off but alveolar spaces indicate large and massive maxillary dentition. Micro-CT-data revealed a highly vascularised inner structure of the dorsal skull elements, which might indicate special feeding adaptations. Adding the new find to an existing phylogenetic analysis of Triassic marine reptiles reveals eosauropterygian, especially nothosauroid, affinities. However, morphological differences to nothosauroids justify the erection of a new genus and species for this enigmatic marine reptile. Its atypical morphology, without any extinct or modern analogue, fits well with the continuously increasing diversity of Triassic marine reptiles, exhibiting various specialised feeding strategies urn:lsid:zoobank.org:act:6D75AEC7-A5C5-4844-B71A-8215AB099134     

Early Mesozoic evolution of alivincular bivalve ligaments and its implications for the timing of the ''Mesozoic marine revolution''

The early Mesozoic radiation of the Pteriomorphia was accompanied and furthered by the development of several new types of alivincular ligaments. These new types evolved as modifications of the primitive alivincular-areate (new term) ligament, which is characterized by an ontogenetic shift of both the central resilium and the straight lateral ligament in the direction of main shell growth. Arching of the attachment surface of the ligament led to the alivincular-arcuate (new term) ligament type, which has been realized by the Ostreidae only. By contrast, a replacement of the lateral ligament by hinge teeth, limiting the (primary) ligament to a central groove (alivincular-fossate, new term), has evolved independently in three families (Dimyidae, Plicatulidae and Spondylidae). Functionally, both kinds of modification effectively impede shearing of the valves and are interpreted as an antipredatory adaptation advantageous in the cemented habit of these families. The alivincular-alate (new term) ligament of the Entoliidae and Pectinidae differs from the other types of alivincular ligaments by different growth directions of resilium and lateral ligament, which result in an internal position of the resilium suitable for fast and powerful opening of the valves. This arrangement is an important prerequisite for effective swimming, which, in its turn, is a behaviour chiefly used to escape from predator attacks. The simultaneous early Mesozoic appearance of different antipredatory adaptations within independent clades hints at increased predator pressure as a stimulant and may therefore point to a contemporaneous proliferation of durophagous predators. Hence, an important aspect of the 'Mesozoic marine revolution' might have started earlier than previously thought.     

Ultrastructural features of the salt gland of Tamarix aphylla L.

Summary The salt gland in Tamarix is a complex of eight cells composed of two inner, vacuolate, collecting cells and six outer, densely cytoplasmic, secretory cells. The secretory cells are completely enclosed by a cuticular layer except along part of the walls between the collecting cells and the inner secretory cell. This non-cuticularized wall region is termed the transfusion are (Ruhland, 1915) and numerous plasmodesmata connect the inner secretory cells with the collecting cells in this area. Plasmodesmata also connect the collecting cells with the adjacent mesophyll cells.There are numerous mitochondria in the secretory cells and in different glands they show wide variation in form. In some glands wall protuberances extend into the secretory cells forming a labyrinth-like structure; however, in other glands the protuberances are not extensively developed. Numerous small vacuoles are found in some glands and these generally are distributed around the periphery of the secretory cells in association with the wall protuberances. Further, an unusual structure or interfacial apparatus is located along the anticlinal walls of the inner secretory cells. The general structure of the gland including the cuticular encasement, connecting plasmodesmata, interfacial apparatus, and variations in mitochondria, vacuoles, and wall structures are discussed in relation to general glandular function.     

Ion secretion by salt glands of desert iguanas (Dipsosaurus dorsalis)

Unlike the NaCl-secreting salt glands of many birds and reptiles, the nasal salt glands of lizards can secrete potassium as well as sodium, with either chloride or bicarbonate as the accompanying anion. The factors responsible for initiating secretion by the gland and the rates of cation and anion secretion were studied in the desert iguana, Dipsosaurus dorsalis. Lizards were given combinations of ions for several days, and secreted salt was collected daily and analyzed for sodium, potassium, chloride, and bicarbonate. Maximum total cation secretion rate was 4.4+/-0.38 micromol/g/d. Cation secretion ranged from 24% to 100% potassium; even high NaCl loads did not abolish potassium secretion. Maximum bicarbonate secretion was about 0.5 micromol/g/d; chloride was the predominant anion. Secretion rate increased only in response to those treatments that included potassium and/or chloride; sodium ions and other osmotic loads (e.g., sucrose) did not increase secretion. This is in contrast to birds and some other reptiles with salt glands, which initiate NaCl secretion in response to any osmotic load. The specificity of the response of the salt gland of Dipsosaurus may be related to the ecological importance of dietary potassium and chloride for herbivorous desert lizards.     

Coping with excess salt: adaptive functions of extrarenal osmoregulatory organs in vertebrates

In all organisms, changing environmental conditions require appropriate regulatory measures to physiologically adjust to the altered situation. Uptake of excess salt in non-mammalian vertebrates having limited or no access to freshwater is balanced by extrarenal salt excretion through specialized structures called ‘salt glands’. Nasal salt glands of marine birds are usually fully developed in very early stages of their lives since individuals of these species are exposed to salt soon after hatching. In individuals of other bird species, salt uptake may occur infrequently. In these animals, glands are usually quiescent and glandular cells are kept in a fairly undifferentiated state. This is the situation in ‘naive’ ducklings, Anas platyrhynchos, which have never been exposed to excess salt. When these animals become initially osmotically stressed, the nasal glands start to secrete a moderately hypertonic sodium chloride solution but secretory performance is meager. Within 48 h after the initial stimulus, however, the number of cells per gland is elevated by a factor of 2–3, the secretory cells differentiate and acquire full secretory capacity. During this differentiation process, extensive surface specializations are formed. The number of mitochondria is increased and metabolic enzymes and transporters are upregulated. These adaptive growth and differentiation processes result in a much higher efficiency of salt excretion in acclimated ducklings compared with naive animals. Receptors and signal transduction pathways in salt gland cells controling the adaptive processes seem to be the same as those controling salt secretion, namely muscarinic acetylcholine receptors and receptors for vasoactive intestinal peptide. This review focusses on signal transduction pathways activated by muscarinic receptors which seem to fine-tune salt secretion in salt-adapted ducklings and may control adaptive growth and differentiation processes in the nasal gland of naive animals.     

Life‐history strategies indicate live‐bearing in Nothosaurus (Sauropterygia)

In Sauropterygia, a diverse group of Mesozoic marine reptiles, fossil evidence of viviparity (live‐bearing) only exists for Pachypleurosauria and Plesiosauria, and was assumed to also be the case for nothosaurs. Previous studies have successfully applied an extant squamate model to sauropterygian life‐history traits. In extant squamates, oviparity and viviparity are associated with differences in life‐history trait combinations. We establish growth curves for Nothosaurus specimens based on their humeral histology. We then analyse life‐history traits derived from these curves and compare inferred traits to those of modern squamates and pachypleurosaurs to assess their reproduction mode. We show that birth to adult size ratios (i.e. birth size divided by the mother's size) provide good estimates of clutch sizes in extant squamates and in viviparous extinct marine reptiles, but these ratios cannot discriminate viviparous and oviparous squamates. Thus, large ratios do not indicate viviparity in fossil taxa to which the extant squamate model is applicable. Applying differences in birth size, age at maturation, and maximum longevity that are observed between extant viviparous and oviparous squamates to our Nothosaurus sample, we identified 7 out of 24 specimens as being potentially viviparous. Conversely, they suggested oviparity for many nothosaurs but also for many pachypleurosaur samples. Under the assumption that the entire clade Pachypleurosauria was viviparous, the majority of nothosaurs would also have been viviparous as they comprised trait combinations similar to those seen in pachypleurosaurs. Overall, this suggests that within nothosaurs and pachypleurosaurs both reproduction modes existed in different taxa.     

Salt marsh foraminifera from the Great Marshes, Massachusetts: environmental controls

By using sites in the Great Marshes at Barnstable (Massachusetts, USA) this study examines the effects of a set of environmental parameters on the foraminiferal distribution. The studied parameters are: elevation above mean high water; salinity of the porewater; various sediment characteristics; vegetation; and food source. Relations between the environmental parameters and foraminiferal properties (frequencies, densities and diversities) are quantified with correlation coefficients. For the first time Siphotrochammina lobata and Balticammina pseudomacrescens are documented in the New England region.

The following species show a significant correlation with one or more of the studied parameters and are designated as key-species: Haplophragmoides manilaensis, Jadammina macrescens, Balticammina pseudomacrescens, Miliammina fusca and Tiphotrocha comprimata. Based on cluster analysis and the presence, absence or dominance of the key-species characteristic associations are distinguished. The distribution of three associations is indicative of specific marsh environments: the marsh fringe, the middle marsh and the marsh edge. These three marsh units are separated by their own salinity regime, flooding and sediment characteristics.

The marsh fringe is typified by the H. manilaensis Association and experiences freshwater input (seepage, surface runoff and rainwater) and only slight marine influence, resulting in low salinity values (2.5–20‰). The width of the marsh fringe is variable, dependent on the amount of seepage which in turn is controlled by the permeability of the basement and the peat. The J. macrescens Association characterizes the middle marsh where salinities are controlled by infiltration of sea- and rainwater and by evaporation. Salinity values are higher than 20‰, while temporarily salinity can reach extreme high values during periods without flooding and high evaporation rates (e.g., 44‰). The fully marine M. fusca Association occupies the daily flooded marsh edge where the salinities have the same values as Cape Cod bay water (ca. 28‰).

Unlike many other salt marshes the distribution of foraminiferal assemblages in the Great Marshes does not show a vertical zonation with respect to mean high water. This shows that a worldwide applicable model for paleoenvironmental studies in salt marshes based on foraminifera is not feasible. Each salt marsh has its own characteristics. Regional factors such as climate play an important role in the salinity regime, while the local upland characteristics determine if seepage takes place. Thus each marsh has its own foraminiferal fingerprint showing the opportunistic behaviour of the salt marsh agglutinants. A surface study is an indispensable first step in assessing the value of foraminifera as paleo-ecological indicators.     

The mixosaurid ichthyosaur Contectopalatus from the Middle Triassic of the German Basin

The skull of the mixosaurid species Contectopalatus atavus (Quenstedt, 1851-52) is the most bizarre of any known ichthyosaur. It possesses a very high sagittal crest formed by the nasal, frontal and parietal bones which grows higher during ontogeny. This skull structure - found to a lesser extent in the other mixosaurid genera Mixosaurus and Phalarodon - is a synapomorphy of the family Mixosauridae. It is here interpreted as correlated with a unique arrangement of the jaw adductor musculature among tetrapods, with the internal jaw adductors extending over most of the skull roof up to the external narial opening. This reconstruction would increase the biting force considerably and the hypothesis is supported by peculiarities of the dentition and jaws of Contectopalatus. Contectopalatus probably reached a length of about 5 meters. It is therefore the largest known mixosaurid and one of the largest Triassic ichthyosaurs. The general text-book picture of mixosaurs as small, rather unspecialized, primitive ichthyosaurs is incorrect. Mixosaurs were a highly specialized, uniquely adapted and very diverse ichthyosaur family, some members of which rank among the marine top predators of their time.     

MARINE REPTILES FROM THE LOWER CRETACEOUS OF SOUTH AUSTRALIA: ELEMENTS OF A HIGH-LATITUDE COLD-WATER ASSEMBLAGE

Abstract: The Lower Cretaceous rocks of South Australia have yielded a diverse marine reptile assemblage of up to five families of plesiosaur (including a new cryptoclidid or cimoliasaurid, indeterminate elasmosaurids, a possible polycotylid, rhomaleosaurids, and pliosaurid) and one family of ichthyosaur (ophthalmosaurid). Other common associated vertebrates include chimaerids and osteichthyans. Sharks, dipnoans and dinosaurs are uncommon and marine turtles are notably absent. The main fossil‐producing strata belong to the Lower Aptian–Lower Albian Bulldog Shale although the Upper Albian Oodnadatta Formation has produced isolated elements. Both these units comprise finely laminated shaly mudstones and claystones deposited in a transgressive shallow coastal, epicontinental marine environment. Estimates of palaeolatitude place South Australia between 60° and 70°S, in the late Early Cretaceous. Sedimentary structures (including lonestone boulders and glendonites), fossils, isotope data and climatic modelling also indicate that seasonally cool–cold conditions (possibly with winter freezing) prevailed during deposition of the Bulldog Shale. This contrasts markedly with climate regimes typically tolerated by modern aquatic reptiles but suggests that some of the South Australian Mesozoic taxa may have possessed adaptations (including elevated metabolic levels and/or annual migration) to cope with low temperatures. A high proportion of juvenile plesiosaur remains in the Bulldog Shale might also indicate that nutrient‐rich cold‐water coastal habitats functioned as both ‘safe calving grounds’ and refuges for young animals prior to their entering the open sea as adults. The occurrence of plesiosaurs and ichthyosaurs in the high‐latitude Lower Cretaceous of southern Australia, along with plesiosaurs and mosasaurs in the Upper Cretaceous of South America, Antarctica, New Zealand and the Chatham Islands, demonstrates that Mesozoic marine reptiles utilized southern high‐latitude environments over a considerable period of time, and that these records do not represent casual occupation by isolated taxa.