The systematic position ofSilvestrosaurus and a classification of Triassic sauropterygians (Neodiapsida)

The Triassic marine reptiles at the base of the extinct Order Sauropterygia, traditionally known as nothosaurs, represent an unnatural assemblage of superficially similar forms. The traditional grouping of these animals has generally relied on the flawed basis of “primitive” characteristics whereas two distinct nothosaurian lineages can now be identified through the use of cladistic methodology. The Pachypleurosauria are the most plesiomorphic, but not the earliest, sauropterygians known. The Nothosauriformes include not only ‘nothosaurs’ but also the monophyletic group Plesiosauria and seemingly also the Placodontia. Rigorous coding of characters for the recently described nothosaurian genusSilvestrosaurus allows expansion of the phylogenetic data base for the Sauropterygia. The revised analysis reaffirms recent studies of relationship for the sauropterygians and demonstrates the affinity ofSilvestrosaurus with the Nothosauriformes. The explicit hierarchical classification scheme presented incorporates the hypothesized phylogenetic relationships of these long-misunderstood taxa. The traditional Suborder Nothosauria, now seen to be paraphyletic, must be abandoned.     

三叠纪胎生主龙型海洋爬行动物

现代爬行动物的生殖方式包括卵生、卵胎生和“胎生”三种类型.总体而言,卵生即母体在陆地上产出羊膜卵,并依靠外界环境加以孵化;卵胎生系指羊膜卵在母体内孵化,随即产出幼子,其间胚胎的营养来源和代谢废物均限于羊膜卵内;所谓“胎生”,是指某些类群的卵在母体内孵化后没有立刻产出,胚胎与母体建立了某种程度的营养乃至气体交换关系,此后再以幼体的形式娩出.卵胎生和“胎生”见于有鳞类.在化石中,由于无法识别胚胎与母体的营养联系等信息,通常只以“生产方式”加以定义,即除卵生以外,所有“含胚胎的化石”都统称为“胎生”,目前“胎生”已有的准确记录包括鱼龙类、鳍龙类、沧龙类和离龙类.近期报道的“怀孕恐头龙”显示原龙类可能也有此种生殖方式,但标本的保存状态显示不能排除其“同类相食”之可能.本文记述了产自云南罗平地区中三叠系内一独立保存的、完整的圆胚状化石,标本的大小、形态、姿态和埋藏环境显示这是一个处于孵化后期、因某种原因而流产的胚胎.头后骨骼显示该标本代表了一种全新类型的海生原龙类,但是由于在个体发育中很多性状特征会有较大变化,因此胚胎乃至幼体标本不宜作为正型标本而建立新属种.这也是又一个主龙型爬行动物胎生的确切证据;同时这也进一步反映了盘县—罗平动物群中原龙类的高度多样性.     

A new phylogenetic hypothesis of turtles with implications for the timing and number of evolutionary transitions to marine lifestyles in the group

Evolutionary transitions to marine habitats occurred frequently among Mesozoic reptiles. Only one such clade survives to the present: sea turtles (Chelonioidea). Other marine turtles originated during the Mesozoic, but uncertain affinities of key fossils have obscured the number of transitions to marine life, and the timing of the origin of marine adaptation in chelonioids. Phylogenetic studies support either a highly‐inclusive chelonioid total‐group including fossil marine clades from the Jurassic and Cretaceous (e.g. protostegids, thalassochelydians, sandownids) or a less inclusive chelonioid total‐group excluding those clades. Under this paradigm, these clades belong outside Cryptodira, and represent at least one additional evolutionary transition to marine life in turtles. We present a new phylogenetic hypothesis informed by high resolution computed tomographic data of living and fossil taxa. Besides a well‐supported Chelonioidea, which includes protostegids, we recover a previously unknown clade of stem‐group turtles, Angolachelonia, which includes the Late Jurassic thalassochelydians, and the Cretaceous–Palaeogene sandownids. Accounting for the Triassic Odontochelys, our results indicate three independent evolutionary transitions to marine life in non‐pleurodiran turtles (plus an additional two‐three in pleurodires). Among all independent origins of marine habits, a pelagic ecology only evolved once, among chelonioids. All turtle groups that independently invaded marine habitats in the Jurassic–Cretaceous (chelonioids, angolachelonians, bothremydid pleurodires) survived the Cretaceous–Palaeogene mass extinction event. This highlights extensive survival of marine turtles compared to other marine reptiles. Furthermore, deeply‐nested clades such as chelonioids are found by the middle Early Cretaceous, suggesting a rapid diversification of crown‐group turtles during the Early Cretaceous.     

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.     

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.     

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.     

Sea snakes (Laticauda spp.) require fresh drinking water: implication for the distribution and persistence of populations

Dehydration and procurement of water are key problems for vertebrates that have secondarily invaded marine environments. Sea snakes and other marine reptiles are thought to remain in water balance without consuming freshwater, owing to the ability of extrarenal salt glands to excrete excess salts obtained either from prey or from drinking seawater directly. Contrary to this long-standing dogma, we report that three species of sea snake actually dehydrate in marine environments. We investigated dehydration and drinking behaviors in three species of amphibious sea kraits (Laticauda spp.) representing a range of habits from semiterrestrial to very highly marine. Snakes that we dehydrated either in air or in seawater refused to drink seawater but drank freshwater or very dilute brackish water (10%-30% seawater) to remain in water balance. We further show that Laticauda spp. can dehydrate severely in the wild and are far more abundant at sites where there are sources of freshwater. A more global examination of all sea snakes demonstrates that species richness correlates positively with mean annual precipitation within the Indo-West Pacific tropical region. The dependence of Laticauda spp. on freshwater might explain the characteristically patchy distributions of these reptiles and is relevant to understanding patterns of extinctions and possible future responses to changes in precipitation related to global warming. In particular, metapopulation dynamics of the Laticauda group of sea snakes are expected to change in relation to projected reductions of tropical dry-season precipitation.     

Revised paleoecology of placodonts – with a comment on ‘The shallow marine placodont Cyamodus of the central European Germanic Basin: its evolution,paleobiogeography and paleoecology’ by C.G. Diedrich (Historical Biology,iFirst article, 2011, 1–19, doi: 10.1080/08912963.2011.575938)

A recent article published by Diedrich (2011a, Hist Biol. iFirst online, 1–19, doi: 10.1080/08912963.2011.575938) aspired to provide a complete revision of the known material of the placodont genus Cyamodus Meyer, 1863 from the Germanic Basin of central Europe. It is the latest in a series of similar articles by the same author (see Diedrich 2010, Palaeogeogr Palaeoclimatol Palaeoecol. 285(3–4):287–306; 2011b, Nat Sci. 3(1):9–27 for overview) focussing on the European members of the Placodontia (Reptilia: Sauropterygia), a diverse group of enigmatic marine reptiles known from Triassic shallow marine deposits. In a similar fashion to some previous works by Diedrich (see Tintori 2011, Palaeogeogr Palaeoclimatol Palaeoecol. 300(1–4):205–207 for similar points of criticism), this newest article demonstrates a narrow scope of presenting and discussing data, including omitted articles relevant to the topic, and over-interpretation of results, all with the aim of embedding the idea of placodonts being herbivorous Triassic ‘sea cows’ feeding on macroalgae (Diedrich 2010, 2011b). The present contribution is intended to clarify mistakes and misinterpretations made by Diedrich (2011a), to incorporate vital citations previously omitted which allow alternative interpretations, and to put the paper into perspective by including a more general evolutionary and paleoecological overview of the remaining placodonts.     

The Enigmatic Marine Reptile Nanchangosaurus from the Lower Triassic of Hubei,China and the Phylogenetic Affinities of Hupehsuchia

The study of the holotype and of a new specimen of Nanchangosaurus suni (Reptilia; Diapsida; Hupehsuchia) revealed a suite of hitherto unrecognized characters. For example, Nanchangosaurus has bipartite neural spines and its vertebral count is nearly identical to that of Hupehsuchus. It differs from the latter in having poorly developed forelimbs despite the advanced ossification in the rest of the skeleton. Other differences all pertain to hupehsuchian plesiomorphies retained in Nanchangosaurus, such as low neural spines. The relationship of Hupehsuchia within Diapsida was analyzed based on a data matrix containing 41 taxa coded for 213 characters, of which 18 were identified as aquatic adaptations from functional inferences. These aquatic adaptations may be vulnerable to the argumentation of character homology because expectation for homoplasy is high. There is an apparent incongruence between phylogenetic signals from aquatic adaptations and the rest of the data, with aquatic adaptations favoring all marine reptiles but Helveticosaurus to form a super-clade. However, this super-clade does not obtain when aquatic adaptations were deleted, whereas individual marine reptile clades are all derived without them. We examined all possible combinations of the 18 aquatic adaptations (n = 262143) and found that four lineages of marine reptiles are recognized almost regardless of which of these features were included in the analysis: Hupehsuchia-Ichthyopterygia clade, Sauropterygia-Saurosphargidae clade, Thalattosauria, and Helveticosaurus. The interrelationships among these four depended on the combination of aquatic adaptations to be included, i.e., assumed to be homologous a priori by bypassing character argumentation. Hupehsuchia always appeared as the sister taxon of Ichthyopterygia.     

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.     

Diversity dynamics of post‐Palaeozoic crinoids – in quest of the factors affecting crinoid macroevolution

Following the end‐Permian biotic crisis which led to the near extinction of crinoids, this echinoderm class rebounded rapidly during the Mesozoic, resulting in forms with important morphological and behavioural novelties. However, quantitative patterns of crinoid diversity during the Mesozoic remain largely unexplored. Here, we report results of analyses of the evolutionary dynamics of post‐Palaeozoic crinoid genera spanning a time interval between 250 and 70 Myr. We show that crinoids reached their Mesozoic peak of genus‐level richness during the Late Jurassic. We also document a major reorganization of different ecological crinoid groups in the Mesozoic. More specifically, the diversity of sessile forms generally increased towards the mid‐Mesozoic but decreased significantly starting in the Cretaceous, whereas the number of motile crinoid genera increased linearly during the Mesozoic. The possible role of biotic and abiotic factors in crinoid evolution is discussed.     

Salt glands in a Tithonian metriorhynchid crocodyliform and their physiological significance

Our knowledge of Mesozoic tetrapods is based mainly on osteological evidence. The discussion of the evolution of any homeostatic system is highly speculative because direct non-osteological evidence is uncommon. Here we report an extraordinarily well-preserved cast of a pair of lobulated protuberances in the skull of the marine metriorhynchid crocodiliform Geosaurus from the Tithonian (Jurassic) of Patagonia (Argentina). These protuberances are interpreted as representing salt glands. Based on their topology, these glands are identified as the nasals. Optimization of this character on a phylogenetic tree permits us to infer the ancestral condition for archosaurs. The relationship between salt gland and diet is also analysed. The presence of hypertrophied salt glands in the skull of Geosaurus suggests that as early as 140 million years ago, some Mesozoic marine reptiles had evolved an extra-renal osmoregulatory system. This achievement was an important clue in the successful colonization of marine environments. Salt glands preclude the risk of lethal dehydration and allow marine reptiles to include an important amount of invertebrates in their diet.     

中国三叠纪海生爬行动物化石研究的回顾与进展

中国三叠纪海生爬行动物化石研究始于20世纪50年代,近10年来取得了重要的进展。此类化石在华南分布广泛,已见于多个省区的十余处地点,涉及自下三叠统奥伦尼克阶至上三叠统诺利阶的至少7个层位,并显示出由东向西产出层位逐渐升高的趋势。我国的三叠纪海生爬行动物化石门类齐全,属种丰富,已知类群包括鱼龙类、海龙类、檐齿龙类、始鳍龙类、原龙类、初龙类和湖北鳄类,显示出典型的西特提斯动物群特征,同时也体现了与东太平洋动物群的某些联系,以及一定的地方色彩。这些化石为研究三叠纪海生爬行动物各个类群的起源、演化和绝灭以及海洋环境的变迁提供了新的材料。     

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.     

A bizarre new toothed mysticete (Cetacea) from Australia and the early evolution of baleen whales

Extant baleen whales (Cetacea, Mysticeti) are all large filter-feeding marine mammals that lack teeth as adults, instead possessing baleen, and feed on small marine animals in bulk. The early evolution of these superlative mammals, and their unique feeding method, has hitherto remained enigmatic. Here, I report a new toothed mysticete from the Late Oligocene of Australia that is more archaic than any previously described. Unlike all other mysticetes, this new whale was small, had enormous eyes and lacked derived adaptations for bulk filter-feeding. Several morphological features suggest that this mysticete was a macrophagous predator, being convergent on some Mesozoic marine reptiles and the extant leopard seal (Hydrurga leptonyx). It thus refutes the notions that all stem mysticetes were filter-feeders, and that the origins and initial radiation of mysticetes was linked to the evolution of filter-feeding. Mysticetes evidently radiated into a variety of disparate forms and feeding ecologies before the evolution of baleen or filter-feeding. The phylogenetic context of the new whale indicates that basal mysticetes were macrophagous predators that did not employ filter-feeding or echolocation, and that the evolution of characters associated with bulk filter-feeding was gradual.     

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.     

Ecophysiological steps of marine adaptation in extant and extinct non-avian tetrapods

Marine reptiles and mammals are phylogenetically so distant from each other that their marine adaptations are rarely compared directly. We reviewed ecophysiological features in extant non-avian marine tetrapods representing 31 marine colonizations to test whether there is a common pattern across higher taxonomic groups, such as mammals and reptiles. Marine adaptations in tetrapods can be roughly divided into aquatic and haline adaptations, each of which seems to follow a sequence of three steps. In combination, these six categories exhibit five steps of marine adaptation that apply across all clades except snakes: Step M1, incipient use of marine resources; Step M2, direct feeding in the saline sea; Step M3, water balance maintenance without terrestrial fresh water; Step M4, minimized terrestrial travel and loss of terrestrial feeding; and Step M5, loss of terrestrial thermoregulation and fur/plumage. Acquisition of viviparity is not included because there is no known case where viviparity evolved after a tetrapod lineage colonized the sea. A similar sequence is found in snakes but with the haline adaptation step (Step M3) lagging behind aquatic adaptation (haline adaptation is Step S5 in snakes), most likely because their unique method of water balance maintenance requires a supply of fresh water. The same constraint may limit the maximum body size of fully marine snakes. Steps M4 and M5 in all taxa except snakes are associated with skeletal adaptations that are mechanistically linked to relevant ecophysiological features, allowing assessment of marine adaptation steps in some fossil marine tetrapods. We identified four fossil clades containing members that reached Step M5 outside of stem whales, pinnipeds, sea cows and sea turtles, namely Eosauropterygia, Ichthyosauromorpha, Mosasauroidea, and Thalattosuchia, while five other clades reached Step M4: Saurosphargidae, Placodontia, Dinocephalosaurus, Desmostylia, and Odontochelys. Clades reaching Steps M4 and M5, both extant and extinct, appear to have higher species diversity than those only reaching Steps M1 to M3, while the total number of clades is higher for the earlier steps. This suggests that marine colonizers only diversified greatly after they minimized their use of terrestrial resources, with many lineages not reaching these advanced steps. Historical patterns suggest that a clade does not advance to Steps M4 and M5 unless these steps are reached early in the evolution of the clade. Intermediate forms before a clade reached Steps M4 and M5 tend to become extinct without leaving extant descendants or fossil evidence. This makes it difficult to reconstruct the evolutionary history of marine adaptation in many clades. Clades that reached Steps M4 and M5 tend to last longer than other marine tetrapod clades, sometimes for more than 100 million years.     

The effect of body size on skeletal articulation and completeness in Jurassic Ichthyopterygia (Reptilia)

A taphonomic investigation of Jurassic ichthyopterygians from the Posidonienschiefer Formation initially supports the common assumption of better preservation with increasing body size. However, there is also an obvious increase in the range of preservational states with decreasing body size, suggesting greater susceptibility of smaller carcasses to the effects of biostratinomic processes. An investigation of articulation and completeness separately against body size indicates a stronger link of completeness to preservation; bigger elements were more difficult to move. Size ranges, which crucially relate to ontogenetic stages, suggest the extent of disarticulation and loss of completeness is greater in juvenile and embryonic skeletons (less than 200 cm ±) compared to those of adults (over 200 cm). The study has important implications for the preservation of different body size ranges, in particular, those representing ontogenetic stages in ichthyopterygians and other marine reptile groups, but, more generally, for vertebrates entering the fossil record.     

Effect of sea water concentration on hyphal growth and antimicrobial metabolite production in marine fungi

We studied the effect of sea water concentration in a culture medium on fungal growth and the production of antimicrobial metabolites. Most of the marine fungal isolates were identified as members of the same genera as terrestrial isolates, such asAspergillus andTrichoderma. Many of the marine fungi isolated grew more abundantly as the sea water concentration increased. The production of antimicrobial materials was improved as the sea water concentration increased. Even though the marine fungi were considered to be similar to fungi from terrestrial environments, from a mycological perspective, the two types have different physiological characteristics. The fungi from marine samples are useful microbial resources in the search for new bioactive compounds.