Biomechanical evolution of solid bones in large animals: a microanatomical investigation

Graviportal taxa show an allometric increase of the cross‐sectional area of supportive bones and are assumed to display microanatomical changes associated with an increase in bone mass. This evokes osteosclerosis (i.e. an increase in bone compactness observed in some aquatic amniotes). The present study investigates the changes in bones' microanatomical organization associated with graviportality and how comparable they are with aquatically acquired osteosclerosis aiming to better understand the adaptation of bone to the different associated functional requirements. Bones of graviportal taxa show microanatomical changes that are not solely attributable to allometry. They display a thicker cortex and a proportionally smaller medullary cavity, with a wider transition zone between these domains. This inner cancellous structure may enable to better enhance energy absorption and marrow support. Moreover, the cross‐sectional geometric parameters indicate increased resistance to stresses engendered by bending and torsion, as well as compression. Adaptation to a graviportal posture should be taken into consideration when analyzing possibly amphibious taxa with a terrestrial‐like morphology. This is particularly important for palaeoecological inferences about large extinct tetrapods that might have been amphibious and, more generally, for the study of early stages of adaptation to an aquatic life in amniotes.     

Mid-Cretaceous/Albian (Cretaceous) ostracod assemblages from NW Hungary reflecting deep marine,nearshore and non-marine environments

The Albian sediments of the Transdanubian Range in Hungary were deposited in non-marine to fully-marine environments. The studied sections revealed diverse benthic ostracod faunas with moderately-preserved specimens. Forty-six taxa are identified and Cypridea zalanyii n. sp. is newly described. The detailed palaeoecological study of these ostracod faunas provided an opportunity to distinguish eight assemblages. The assemblages include limnic, brackish lagoon, shallow marine, reef and low oxygenated semi-enclosed basin environments in the Northern Tethys region we investigated. This work contributes to the knowledge about Albian non-marine faunas from Europe and explores ecological requirements of brackish faunas.     

New highlights about the enigmatic marine snake Palaeophis maghrebianus (Palaeophiidae; Palaeophiinae) from the Ypresian (Lower Eocene) phosphates of Morocco

Abstract: Palaeophis maghrebianus belongs to the Palaeophiinae (Palaeophiidae). This snake subfamily is relatively poorly known, and it is mainly represented by disarticulated vertebrae and ribs and by a few vertebral segments. Its intracolumnar variability remains also poorly understood. The discovery of new isolated vertebrae and vertebral segments of Palaeophis maghrebianus in the Ypresian (Lower Eocene) Phosphates of Morocco enables us to provide a more detailed diagnosis of this species and to describe its intracolumnar variability. Moreover, the new material reveals that this species could reach gigantic size being, with Palaeophis colossaeus, one of the two longer palaeophiids. The microanatomical and histological analysis of some vertebrae illustrating diverse positions along the vertebral column reveals the presence of osteosclerosis, especially in the anterior and mid‐precloacal regions. The occurrence of this osseous specialization implies a role in buoyancy and body trim control in this taxon, which is considered a shallow marine dweller based on its anatomical features and geological data. Palaeophis maghrebianus also displays a dense vascular network suggesting a growth speed, and thus a metabolic rate, much higher than in the biggest extant snakes.     

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.     

Rib and vertebral micro‐anatomical characteristics of hydropelvic mosasauroids

Houssaye, A. & Bardet, N. 2011: Rib and vertebral micro‐anatomical characteristics of hydropelvic mosasauroids. Lethaia, Vol. 45, pp. 200–209. Mosasauroids are squamates secondarily adapted to an aquatic life that dominated the sea during the Late Cretaceous. Mosasauroids display distinct types of morphologies illustrating steps in the adaptation of this lineage to increasing obligatory habits. Hydropelvic mosasauroids (sensu Caldwell & Palci) were the most highly adapted to an open‐sea environment. Contrary to plesiopelvic forms, they are considered to have relied on a hydrodynamic, rather than hydrostatic buoyancy and body trim control strategies. This led previous authors to consider that these taxa would favour bone lightening (osteoporotic‐like condition) rather than bone mass increase. Although an osteoporotic‐like state was indeed described in Clidastes and Tylosaurus, bone mass increase was reported in Platecarpus. As a matter of fact, the new analysis of vertebral thin sections of various taxa combined with the reanalysis of the rib sections available in Sheldon’s PhD thesis in a micro‐anatomical perspective reveals the absence of both bone mass increase and bone lightening in these organisms. These taxa in fact display a vertebral micro‐anatomy much peculiar within squamates. It characteristically corresponds to a true network of thin trabeculae whose tightness varies between taxa, probably as a result of both species and individual size differences, particularly the latter. In addition, analysis of the pattern of vascularization as observed in hydropelvic mosasauroids, which is unique amongst squamates, suggests that large size in hydropelvic mosasauroids would mainly rely on protracted rather than faster growth rates. □Histology, hydropelvic mosasauroids, micro‐anatomy, ribs, vascularization, vertebrae.     

Squamate reptiles from the middle Eocene of Lissieu (France). A landmark in the middle Eocene of Europe

In Europe, faunas of squamates (lizards and snakes) from the middle Eocene are very poorly known, with the exception of those from the level MP 16 (latest middle Eocene). From the MP 11-MP 15 interval, squamates were previously reported only from Messel (MP 11, earliest middle Eocene) and from the untere and obere Mittelkohle of Geiseltal (MP 12 and MP 13 respectively) in Germany. The present report describes the middle Eocene assemblage of squamates from Lissieu (France), the first fauna reported from the level MP 14. Whereas fossils from Messel and Geiseltal are mostly articulated skeletons, fossils from Lissieu are represented by disarticulated bones; such fossils may be more easily compared to those from other Cenozoic localities, in which bones are almost always disarticulated. The fauna from Lissieu is more diverse than those from the Geiseltal sites and approximately as diverse as that from Messel as they are presently known; it is comprised of 17 distinct taxa. These taxa cannot be all identified to the species or genus level. They belong to iguanids, gekkonids, lacertids, anguids, thecoglossan platynotans, ophidians incertaesedis, boids, ?tropidophiines, “tropidophiids” incertaesedis, booids incertaesedis, and perhaps russellophiids. The fauna includes several new taxa but only a presumed tropidophiine snake may be named on the basis of the available material. The fauna from Lissieu is a mixture of taxa restricted to the middle Eocene and taxa known from older or younger levels. Taxa shared by Lissieu and the few other localities from the middle Eocene of Europe are rare. This fauna from Lissieu represents a stratigraphical landmark for the middle Eocene.     

Molecular diversity of fungal phylotypes co-amplified alongside nematodes from coastal and deep-sea marine environments

Nematodes and fungi are both ubiquitous in marine environments, yet few studies have investigated relationships between these two groups. Microbial species share many well-documented interactions with both free-living and parasitic nematode species, and limited data from previous studies have suggested ecological associations between fungi and nematodes in benthic marine habitats. This study aimed to further document the taxonomy and distribution of fungal taxa often co-amplified from nematode specimens. A total of 15 fungal 18S rRNA phylotypes were isolated from nematode specimens representing both deep-sea and shallow water habitats; all fungal isolates displayed high pairwise sequence identities with published data in Genbank (99-100%) and unpublished high-throughput 454 environmental datasets (>95%). BLAST matches indicate marine fungal sequences amplified in this study broadly represent taxa within the phyla Ascomycota and Basidiomycota, and several phylotypes showed robust groupings with known taxa in phylogenetic topologies. In addition, some fungal phylotypes appeared to be present in disparate geographic habitats, suggesting cosmopolitan distributions or closely related species complexes in at least some marine fungi. The present study was only able to isolate fungal DNA from a restricted set of nematode taxa; further work is needed to fully investigate the taxonomic scope and function of nematode-fungal interactions.     

The evolution of ecological specialization across the range of a broadly distributed marine species

Ecological specialization is an important engine of evolutionary change and adaptive radiation, but empirical evidence of local adaptation in marine environments is rare, a pattern that has been attributed to the high dispersal ability of marine taxa and limited geographic barriers to gene flow. The broad-nosed pipefish, Syngnathus typhle, is one of the most broadly distributed syngnathid species and shows pronounced variation in cranial morphology across its range, a factor that may contribute to its success in colonizing new environments. We quantified variation in cranial morphology across the species range using geometric morphometrics, and tested for evidence of trophic specialization by comparing individual-level dietary composition with the community of prey available at each site. Although the diets of juvenile pipefish from each site were qualitatively similar, ontogenetic shifts in dietary composition resulted in adult populations with distinctive diets consistent with their divergent cranial morphology. Morphological differences found in nature are maintained under common garden conditions, indicating that trophic specialization in S. typhle is a heritable trait subject to selection. Our data highlight the potential for ecological specialization in response to spatially variable selection pressures in broadly distributed marine species.     

A LONG-BODIED LIZARD FROM THE LOWER CRETACEOUS OF JAPAN

Abstract:  Platynotan lizards underwent a dramatic Late Cretaceous radiation into marine habitats. Beginning with small-bodied forms, the lineage culminated with the mosasaurs, large predatory lizards with a world-wide distribution in the Santonian–Campanian. Moreover, the marine squamate radiations of the Cenomanian–Turonian are remarkable in having produced a range of long-bodied, reduced-limbed swimmers (dolichosaurs, adriosaurs, coniasaurs and limbed snakes) that seem to have thrived in the shallow coastal environments of the Western Tethys region. Until now, none of these long-bodied aquatic squamates has been recorded prior to the Cenomanian, none has been recovered from a non-marine locality and none is known from Asia. Here we describe a small, gracile, long-bodied mosasauroid lizard from a swampy continental deposit in the Lower Cretaceous of Japan.     

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

Background

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

Methodology/Principal Findings

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

Conclusions/Significance

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

Speciation in marine environments: Diving under the surface

Marine environments are inhabited by a broad representation of the tree of life, yet our understanding of speciation in marine ecosystems is extremely limited compared with terrestrial and freshwater environments. Developing a more comprehensive picture of speciation in marine environments requires that we 'dive under the surface' by studying a wider range of taxa and ecosystems is necessary for a more comprehensive picture of speciation. Although studying marine evolutionary processes is often challenging, recent technological advances in different fields, from maritime engineering to genomics, are making it increasingly possible to study speciation of marine life forms across diverse ecosystems and taxa. Motivated by recent research in the field, including the 14 contributions in this issue, we highlight and discuss six axes of research that we think will deepen our understanding of speciation in the marine realm: (a) study a broader range of marine environments and organisms; (b) identify the reproductive barriers driving speciation between marine taxa; (c) understand the role of different genomic architectures underlying reproductive isolation; (d) infer the evolutionary history of divergence using model‐based approaches; (e) study patterns of hybridization and introgression between marine taxa; and (f) implement highly interdisciplinary, collaborative research programmes. In outlining these goals, we hope to inspire researchers to continue filling this critical knowledge gap surrounding the origins of marine biodiversity.     

Identifying the plant‐associated microbiome across aquatic and terrestrial environments: the effects of amplification method on taxa discovery

Plants in terrestrial and aquatic environments contain a diverse microbiome. Yet, the chloroplast and mitochondria organelles of the plant eukaryotic cell originate from free‐living cyanobacteria and Rickettsiales. This represents a challenge for sequencing the plant microbiome with universal primers, as ~99% of 16S rRNA sequences may consist of chloroplast and mitochondrial sequences. Peptide nucleic acid clamps offer a potential solution by blocking amplification of host‐associated sequences. We assessed the efficacy of chloroplast and mitochondria‐blocking clamps against a range of microbial taxa from soil, freshwater and marine environments. While we found that the mitochondrial blocking clamps appear to be a robust method for assessing animal‐associated microbiota, Proteobacterial 16S rRNA binds to the chloroplast‐blocking clamp, resulting in a strong sequencing bias against this group. We attribute this bias to a conserved 14‐bp sequence in the Proteobacteria that matches the 17‐bp chloroplast‐blocking clamp sequence. By scanning the Greengenes database, we provide a reference list of nearly 1500 taxa that contain this 14‐bp sequence, including 48 families such as the Rhodobacteraceae, Phyllobacteriaceae, Rhizobiaceae, Kiloniellaceae and Caulobacteraceae. To determine where these taxa are found in nature, we mapped this taxa reference list against the Earth Microbiome Project database. These taxa are abundant in a variety of environments, particularly aquatic and semiaquatic freshwater and marine habitats. To facilitate informed decisions on effective use of organelle‐blocking clamps, we provide a searchable database of microbial taxa in the Greengenes and Silva databases matching various n‐mer oligonucleotides of each PNA sequence.     

A taphonomic approach to the genetic interpretation of clypeasteroid accumulations from the Miocene of Tarragona,NE Spain

Belaústegui, Z., Nebelsick, J.H., Gibert, J.M. de, Domènech, R. & Martinell, J. 2012: A taphonomic approach to the genetic interpretation of clypeasteroid accumulations from the Miocene of Tarragona, NE Spain. Lethaia, Vol. 45, pp. 548–565. Clypeasteroid accumulations are common in Cenozoic shallow marine sediments, particularly in the Neogene, as they are also in analogue modern environments. In this article, four clypeasteroid accumulations from Miocene (Serravallian) shallow marine carbonates of the El Camp de Tarragona Basin (NE Spain) are studied. Two of them are dominated by flat‐shaped Parascutella, whereas the other two consist exclusively of bell‐shaped Clypeaster. The combination of the taphonomic analysis of the tests, the palaeoecological and taphonomic information provided by the associated fossils, and the stratigraphical and sedimentological context of each one of these beds allows interpreting them as autochthonous/parautochthonous accumulations resulting from in situ reworking in moderate energy settings, except for one that constitutes a true sand dollar coquina deposited by a storm event. This contribution explores the potential of thorough taphonomic analysis of marine invertebrate skeletal concentrations and substantiates the importance of clypeasteroid echinoids as producers of shell beds in the Neogene. □ Clypeasteroids, Echinoids, Miocene, Taphonomy, Tarragona.     

The effect of habitat on modern shark diversification

Sharks occupy marine habitats ranging from shallow, inshore environments to pelagic, and deepwaters, and thus provide a model system for testing how gross habitat differences have shaped vertebrate macroevolution. Palaeontological studies have shown that onshore lineages diversify more quickly than offshore taxa. Among onshore habitats, coral reef‐association has been shown to increase speciation rates in several groups of fishes and invertebrates. In this study, we investigated whether speciation rates are habitat dependent by generating the first comprehensive molecular timescale for shark divergence. Using phylogenetic comparative methods, we rejected the hypothesis that shelf (i.e. onshore) lineages have higher speciation rates compared to those occupying deepwater and oceanic (i.e. offshore) habitats. Our results, however, support the hypothesis of increased speciation rates in coral reef‐associated lineages within the Carcharhinidae. Our new timetree suggests that the two major shark lineages leading to the extant shark diversity began diversifying mostly after the end‐Permian mass extinction: the squalimorphs into deepwater and the galeomorphs into shelf habitats. We suggest that the breakdown of the onshore–offshore speciation rate pattern in sharks is mediated by success in deepwater environments through ecological partitioning, and in some cases, the evolution of morphological novelty.     

Palaeoecology of the mid‐Cretaceous siphonate bivalve genus Goshoraia (Mollusca,Veneridae) from Japan

Abstract: The mid‐Cretaceous bivalve Goshoraia Tamura, 1977, endemic to Japan, is an early example of shallow‐marine siphonate bivalves of the family Veneridae Rafinesque, 1815. Three species, including one new, are here described: Goshoraia minor Tashiro and Kozai, 1989 (Aptian), G. crenulata (Matsumoto, 1938; Albian–lower Cenomanian) and G. maedai sp. nov. (middle to ?upper Cenomanian). The habitats of Goshoraia have been extensively compared with those of common Cretaceous, nonsiphonate burrowers, such as trigoniids, which range from tidal flat and shoreface to shelf environments. Depth of burial, which can be estimated from the extent of the pallial sinus, increases from the ancestral G. minor to its descendants G. crenulata and G. maedai sp. nov., documenting that the ability to burrow within this genus improved in time. These morphological and palaeoecological changes may be related to the Mesozoic marine revolution during the mid‐Cretaceous.     

Les ostracodes continentaux au Néogène en Aquitaine (Sud-Ouest France). Répartition spatio-temporelle : le point sur la question

A systematic research of the non-marine ostracodes from the Aquitaine Basin Neogene shows that these animals were locally well represented and able to indicate palaeoecological and palaeoclimatic data, comparable in quality with such data as from Recent continental environments. We provide a list of the taxa, and the stratigraphical distribution during the Lower and the Middle Miocene is indicated, as well as the geographical location in comparison with the estimated shorelines. The recent stratigraphical revisions of some Miocene deposits, mainly marine, from the central, southern and northern parts of the Aquitaine Basin are used. The studied microfauna contains about 20 genera and slightly more than 30 species. Most of the found genera correspond to fresh to brackish ostracodes (particularly Cytheridae and Candonidae). Several genera can indicate environmental characteristics: Candonopsis and Sclerocypris (Upper Aquitanian in age) are intertropical dwellers. Vecticypris from the Lower Miocene could indicate (in analogy with Metacypris) relatively cooler waters existing in these environments.     

A new pachyostotic squamate reptile from the Cenomanian of France

Cenomanian beds from western France have yielded a new pythonomorph squamate (Reptilia) that is described here as Carentonosaurus mineaui gen. et sp. nov. This fossil is referred to non-mosasaurid mosasauroid lizards, i.e. to 'aigialosaurs'. It has been found in sediments of marine origin and its anatomy confirms that it was an aquatic lizard. It is characterized by a combination of characters that has not been reported, thus far, for squamates. Moreover, its vertebral column includes non-pachyostotic cervical vertebrae and highly pachyostotic dorsal vertebrae as in several other 'aigialosaurs'. This new taxon is perhaps restricted to the upper Cenomanian. It lived in shallow and rather warm water of the inner shelf. It is worth mentioning that nearly all pachyostotic squamates are concentrated in the Cenomanian and/or lower Turonian deposits of the present European-North African-Middle East portion of the Tethys. A parallel is drawn between the high percentage of pachyostotic squamates and the fact that this period corresponds to both the largest transgression of the Phanerozoic and the warmest period in the whole of the Mesozoic and Caenozoic.