Crown Group Research Articles

A molecular phylogeny of the Sematophyllaceae s.l. (Hypnales) based on plastid, mitochondrial and nuclear markers, and its taxonomic implications

AbstractThe Sematophyllaceae s.l. (Sematophyllaceae+Pylaisiadelphaceae) is a family of pleurocarpous mosses that is widely distributed throughout the globe, with centers of diversity in tropical forests. The circumscriptions of the family and its genera have been unstable, due to reductions in morphological complexity and alternative weightings of discrete morphological traits. Based on a sample spanning much of the generic diversity of the family, we inferred the phylogenetic relationships within the Sematophyllaceae s.l. from the variation in eight molecular markers from all three genomes (nuclear, mitochondrial, chloroplast). The Sematophyllaceae s.l. was resolved as monophyletic, as was the Sematophyllaceae s.str.; whereas the Pylaisiadelphaceae was found to be paraphyletic, although its monophyly could not be rejected. The morphological definition of the Pylaisiadelphaceae remains dubious, in the absence of unambiguous synapomorphies. The relationships of the clades of Pylaisiadelphaceae and Sematophyllaceae are discussed with respect to the circumscription of morphogenera, with a focus on the Sematophyllaceae crown clade (Aptychopsis, Chionostomum, Colobodontium, Donnellia, Macrohymenium, Paranapiacabaea, Pterogoniopsis, Rhaphidorrhynchium, Schroeterella, Sematophyllum, Warburgiella). Most genera of Sematophyllaceae were resolved as polyphyletic (e.g., Acroporium, Donnellia, Schroeterella, Sematophyllum, Trichosteleum) indicative of severe homoplasy in their putative diagnostic traits. We propose 4 new genera (Brittonodoxa, Microcalpe, Pocsia, Vitalia) and 19 new combinations (Aptychopsis cylindrothecia, A. estrellae, A. tequendamensis, Brittonodoxa allinckxiorum, B. cataractae, B. lithophila, B. squarrosa, B. steyermarkii, B. subpinnata, Microcalpe subsimplex, Pocsia matutina, Pterogoniopsis paulista, Schroeterella exigua, Trichosteleum amnigenum, T. lonchophyllum, Vitalia caespitosa, V. cuspidifera, V. esmeraldica, V. galipensis)

Early Late Miocene Murine Rodents from the Upper Part of the Nagri Formation, Siwalik Group, Pakistan, with a new fossil calibration point for the Tribe Apodemurini (Apodemus/Tokudaia)

Abstract The early late Miocene is an interval of increased diversification for murine rodents. Whereas the genus Progonomys became widespread throughout Eurasia by 10 Ma, it appears from the known paleontological record that southern Asia is the arena of evolution and diversification at the base of the Murinae. The Siwalik fossil record of the Potwar Plateau in northern Pakistan preserves fossil evidence relevant to unraveling this history. Murine rodents are recorded there throughout the middle Miocene, and diversification began in Siwalik assemblages before 11 Ma. The well-established Progonomys and Karnimata lineages were already present between 11 and 10 Ma, and these represent extant murine crown groups. Here we document diversity in Siwalik murines dating to 10.5 to 10.1 Ma, and clarify their recognition by naming a new species of Karnimata and referring specimens of Progonomys from this interval to P. hussaini. In addition, we define at least two other uncommon murine species that coexist with them. One of these is an early record of Parapodemus, a fossil genus of Tribe Apodemurini, which constitutes a calibration point for the Apodemus/Tokudaia split. Together, these fossil taxa provide further evidence bearing on the major split among murines leading to the clades Murini and Arvicanthini.

New insights in Russula subsect. Rubrinae: phylogeny and the quest for synapomorphic characters

Russula is one of the most speciose genera of mushroom-forming fungi, but phylogenetic relationships among species and subgeneric groupings are poorly understood. Our multi-locus phylogenetic reconstruction places R. firmula, R. rubra, R. rutila and R. veternosa in a well-supported Rubrinae clade, belonging to the Integrae clade of the Crown clade of the genus Russula. Traditional morphology-based classifications placed these four species in two different subsections based on the presence or absence of incrustations on pileocystidia. The Integrae clade also contains R. integra and related species that are traditionally placed in other groups based on their mild taste. Ancestral state reconstruction suggests that the common ancestor of the Crown clade and the Integrae clade probably did not have any incrustations in the pileipellis, had a mild taste, yellow spore print and were associated with angiosperms. All four species of the Rubrinae clade are defined by a darker yellow or ochre spore print, acrid taste and incrustations on pileocystidia. This last character contradicts the former splitting of the group because incrustations were apparently overlooked in R. firmula and R. veternosa. Incrustation type is now highlighted as being important for the delimitation of species and groups within the Crown clade. Pink or red staining of the incrustations in sulphovanillin is present in all species of the Rubrinae clade and a majority of the analysed species of the Integrae clade. The delimitation of the Rubrinae clade and its species circumscriptions are summarised here in a new diagnostic key.

Molecular systematics of the anchovy genus Encrasicholina in the Northwest Pacific.

The anchovy genus Encrasicholina is an important coastal marine resource of the tropical Indo-West Pacific (IWP) region for which insufficient comparative data are available to evaluate the effects of current exploitation levels on the sustainability of its species and populations. Encrasicholina currently comprises nine valid species that are morphologically very similar. Only three, Encrasicholina punctifer, E. heteroloba, and E. pseudoheteroloba, occur in the Northwest Pacific subregion of the northeastern part of the IWP region. These species are otherwise broadly distributed and abundant in the IWP region, making them the most important anchovy species for local fisheries. In this study, we reconstructed the phylogeny of these three species of Encrasicholina within the Engraulidae. We sequenced 10 complete mitochondrial genomes (using high-throughput and Sanger DNA sequencing technologies) and compared those sequences to 21 previously published mitochondrial genomes from various engraulid taxa. The phylogenetic results showed that the genus Encrasicholina is monophyletic, and it is the sister group to the more-diverse "New World anchovy" clade. The mitogenome-based dating results indicated that the crown group Encrasicholina originated about 33.7 million years ago (nearby the limit Eocene/Oligocene), and each species of Encrasicholina has been reproductively isolated from the others for more than 20 million years, despite their morphological similarities. In contrast, preliminary population genetic analyses across the Northwest Pacific region using four mitogenomic sequences revealed very low levels of genetic differentiation within Encrasicholina punctifer. These molecular results combined with recent taxonomic revisions are important for designing further studies on the population structure and phylogeography of these anchovies.

A Burmese amber fossil of <i>Radula</i> (Porellales, Jungermanniopsida) provides insights into the Cretaceous evolution of epiphytic lineages of leafy liverworts

Abstract. DNA-based divergence time estimates suggested major changes in the composition of epiphyte lineages of liverworts during the Cretaceous; however, evidence from the fossil record is scarce. We present the first Cretaceous fossil of the predominantly epiphytic leafy liverwort genus Radula in ca. 100 Myr old Burmese amber. The fossil's exquisite preservation allows first insights into the morphology of early crown group representatives of Radula occurring in gymnosperm-dominated forests. Ancestral character state reconstruction aligns the fossil with the crown group of Radula subg. Odontoradula; however, corresponding divergence time estimates using the software BEAST lead to unrealistically old age estimates. Alternatively, assignment of the fossil to the stem of subg. Odontoradula results in a stem age estimate of Radula of 227.8 Ma (95 % highest posterior density (HPD): 165.7–306.7) and a crown group estimate of 176.3 Ma (135.1–227.4), in agreement with analyses employing standard substitution rates (stem age 235.6 Ma (142.9–368.5), crown group age 183.8 Ma (109.9–289.1)). The fossil likely belongs to the stem lineage of Radula subg. Odontoradula. The fossil's modern morphology suggests that switches from gymnosperm to angiosperm phorophytes occurred without changes in plant body plans in epiphytic liverworts. The fossil provides evidence for striking morphological homoplasy in time. Even conservative node assignments of the fossil support older rather than younger age estimates of the Radula crown group, involving origins for most extant subgenera by the end of the Cretaceous and diversification of their crown groups in the Cenozoic.

At the dawn of the great rise: †Ponomarenkia belmonthensis (Insecta: Coleoptera), a remarkable new Late Permian beetle from the Southern Hemisphere

The first complete beetle body fossil from the Australian early Late Permian fossil site Belmont is described here, †Ponomarenkia belmonthensis sp. nov., attributed to a new extinct family †Ponomarenkiidae. Shortly before a dramatic biodiversity crisis at the end of the Palaeozoic, it documents profound transitions in the evolution of Coleoptera, today by far the most species-rich group of insects and the largest order of organisms. †Ponomarenkia displays transitional states of several important characters, excluding it from the ancestral earliest stem-group coleopterans (e.g. †Tshecardocoleidae), but also from the four ‘modern’ extant suborders. In contrast to †Tshecardocoleidae, †Permocupedidae and †Rhomboleidae, it lacks the ancestral very broad and apically truncated prosternal process and a broad prothoracic postcoxal bridge, features suggesting a position in Coleoptera sensu stricto, i.e. the crown group of beetles. However, it does not share apomorphic features with extant Archostemata (e.g. narrowed neck region), Polyphaga (e.g. internalized propleura), Adephaga (e.g. elongated metacoxae with strongly developed metacoxal plates) or Myxophaga (broadly separated mesocoxae). †Ponomarenkia likely belongs in the stem group of one of the extant suborders or in the stem group of a clade comprising more than one of them.http://zoobank.org/urn:lsid:zoobank.org:pub:BEC74A0C-C902-4326-8549-7F5D5E99E0C3

Genomic Signature of an Avian Lilliput Effect across the K-Pg Extinction

Survivorship following major mass extinctions may be associated with a decrease in body size-a phenomenon called the Lilliput Effect. Body size is a strong predictor of many life history traits (LHTs), and is known to influence demography and intrinsic biological processes. Pronounced changes in organismal size throughout Earth history are therefore likely to be associated with concomitant genome-wide changes in evolutionary rates. Here, we report pronounced heterogeneity in rates of molecular evolution (varying up to $\sim$20-fold) across a large-scale avian phylogenomic data set and show that nucleotide substitution rates are strongly correlated with body size and metabolic rate. We also identify potential body size reductions associated with the Cretaceous-Paleogene (K-Pg) transition, consistent with a Lilliput Effect in the wake of that mass extinction event. We posit that selection for reduced body size across the K-Pg extinction horizon may have resulted in transient increases in substitution rate along the deepest branches of the extant avian tree of life. This "hidden" rate acceleration may result in both strict and relaxed molecular clocks over-estimating the age of the avian crown group through the relationship between life history and demographic parameters that scale with molecular substitution rate. If reductions in body size (and/or selection for related demographic parameters like short generation times) are a common property of lineages surviving mass extinctions, this phenomenon may help resolve persistent divergence time debates across the tree of life. Furthermore, our results suggest that selection for certain LHTs may be associated with deterministic molecular evolutionary outcomes.

Uncovering missing pieces: duplication and deletion history of arrestins in deuterostomes

BackgroundThe cytosolic arrestin proteins mediate desensitization of activated G protein-coupled receptors (GPCRs) via competition with G proteins for the active phosphorylated receptors. Arrestins in active, including receptor-bound, conformation are also transducers of signaling. Therefore, this protein family is an attractive therapeutic target. The signaling outcome is believed to be a result of structural and sequence-dependent interactions of arrestins with GPCRs and other protein partners. Here we elucidated the detailed evolution of arrestins in deuterostomes.ResultsIdentity and number of arrestin paralogs were determined searching deuterostome genomes and gene expression data. In contrast to standard gene prediction methods, our strategy first detects exons situated on different scaffolds and then solves the problem of assigning them to the correct gene. This increases both the completeness and the accuracy of the annotation in comparison to conventional database search strategies applied by the community. The employed strategy enabled us to map in detail the duplication- and deletion history of arrestin paralogs including tandem duplications, pseudogenizations and the formation of retrogenes. The two rounds of whole genome duplications in the vertebrate stem lineage gave rise to four arrestin paralogs. Surprisingly, visual arrestin ARR3 was lost in the mammalian clades Afrotheria and Xenarthra. Duplications in specific clades, on the other hand, must have given rise to new paralogs that show signatures of diversification in functional elements important for receptor binding and phosphate sensing.ConclusionThe current study traces the functional evolution of deuterostome arrestins in unprecedented detail. Based on a precise re-annotation of the exon-intron structure at nucleotide resolution, we infer the gain and loss of paralogs and patterns of conservation, co-variation and selection.

Metamorphosis Is Ancestral for Crown Euarthropods, and Evolved in the Cambrian or Earlier.

Macroevolutionary developmental biology employs fossilized ontogenetic data and phylogenetic comparative methods to probe the evolution of development at ancient nodes. Despite the prevalence of ecologically differentiated larval forms in marine invertebrates, it has been frequently presumed that the ancestors of arthropods were direct developers, and that metamorphosis may not have evolved until the Ordovician or later. Using fossils and new dated phylogenies, I infer that metamorphosis was likely ancestral for crown arthropods, contradicting this assumption. Based on a published morphological dataset encompassing 217 exceptionally preserved fossil and 96 extant taxa, fossils were directly incorporated into both the topology and age estimates, as in "tip dating" analyses. Using data from post-embryonic fossils representing 25 species throughout stem and crown arthropod lineages (as well as most of the 96 extant taxa), characters for metamorphosis were assigned based on inferred ecological changes in development (e.g., changes in habitat and adaptive landscape). Under all phylogenetic hypotheses, metamorphosis was supported as most likely ancestral to both ecdysozoans and euarthropods. Care must be taken to account for potential drastic post-embryonic morphological changes in evolutionary analyses. Many stem group euarthrpods may have had ecologically differentiated larval stages that did not preserve in the fossil record. Moreover, a complex life cycle and planktonic ecology may have evolved in the Ediacaran or earlier, and may have typified the pre-Cambrian explosion "wormworld" prior to the origin of crown group euarthropods.

FRACTURE RESISTANCE AND FAILURE MODE OF TWO RESTORATION DESIGNS MADE OF MONOLITHIC HYBRID AND GLASS MACHINABLE CERAMICS ; IN VITRO STUDY

Statement of the problem: Monolithic hybrid and glass ceramics are indicated for intracoronal and extracoronal ceramic restorations. Available data regarding selection of the appropriate ceramic type for each preparation design and its influence on fracture resistance of tooth restoration complex is still scarce.Purpose: The purpose of this study was to evaluate in vitro the fracture resistance and investigate failure mode of two restoration designs (crowns and inlays) made from lithium disilicate glass ceramic and hybrid ceramic constructed by the CAD /CAM technique.Materials and Methods: A total of twenty ceramic restorations restoring lower molars were designed and fabricated in this study. The restorations were divided into two groups; Group 1: Ten full coverage crowns and Group 2: Ten inlays. Each of the previous groups was further subdivided into two equal subgroups; Subgroup 1: Five restorations constructed from lithium disilicate glass ceramics (e.max CAD) and Subgroup 2: Five restorations constructed from hybrid ceramic (VITA ENAMIC) by the CAD/CAM technology. Two ivorine teeth were prepared following the guidelines for full coverage and inlay ceramic restorations, they were duplicated into stone dies followed by scanning, designing and milling of restorations. The stone dies were duplicated into epoxy dies where the restorations were adhesively cemented on them. All cemented samples were exposed to 2000 thermal cycles (5°C and 55°C in water) with a dwell time of 25 seconds in a computer controlled thermocycler. All specimens were loaded in a universal testing machine where fracture test was done by compressive mode of load applied occlusally using a metallic rod with round tip at cross-head speed of 1mm/min until fracture occurred. Fracture resistance values were recorded in Newtons then they were statistically evaluated and failure modes were analyzed. Results: For group 1 representing crown restorations, a statistically significant difference was found between (e.max CAD) and (VITA ENAMIC) subgroups at (p=0.05) where the highest mean value of maximum load was found in (VITA ENAMIC) (951.77 N ± 81.05) and the lowest mean value of maximum load was found in (e.max CAD) (709.35 N± 49.51). While for the second group representing inlay restorations, no statistically significant difference was found between (e.max CAD) and (VITA ENAMIC) subgroups at (p=0.307) where the highest mean value of maximum load was found in (e.max CAD) (627.71 N ± 80.01) and the lowest mean value of maximum load was found in (VITA ENAMIC) (581.26 N± 51.74). For the e.max CAD subgroup, a statistically significant difference was found between (Crown) and (Inlay) groups at (p=0.005) where the highest mean value of maximum load was found in the crown group (790.35 N± 49.51) and the lowest mean value of maximum load was found in the inlay group (627.71N ± 80.01).Regarding the VITA ENAMIC subgroup, a statistically significant difference was found between (Crown) and (Inlay) groups at (p<0.001) where the highest mean value of maximum load was found in the crown group (951.77 N ± 81.05) and the lowest mean value of maximum load was found in the inlay group (581.26N±51.74).Conclusions: 1. All inlays and crowns constructed from hybrid and glass ceramics in this study revealed fracture load values falling in the range of physiologic masticatory forces 2. When considering restoration of molars with full coverage crowns, VITA ENAMIC is more preferred than e.max CAD ceramic in terms of fracture resistance as it showed a higher statistical significant mean values.3. When planning to restore molars with minimally invasive intracoronal restorations, both ceramics can be used as there was insignificant statistical difference between them , however e.max CAD can be recommended due to its more favorable mode of fracture compared to VITA ENAMIC.4. Irrespective of preparation designs, both ceramics showed insignificant difference regarding fracture resistance mean values.

Oligocene and Miocene albatross fossils from Washington State (USA) and the evolutionary history of North Pacific Diomedeidae

ABSTRACT Albatross fossils have been collected from the late Oligocene Lincoln Creek Formation and the early/middle Miocene Astoria Formation near the townsite of Knappton, Pacific County, Washington (USA). The albatross from the Lincoln Creek Formation, Diomedavus knapptonensis, n. gen. et sp., is smaller than all extant albatrosses and represents the oldest published fossil albatross from the North Pacific Basin. Diomedavus knapptonensis is clearly distinguished from extant albatrosses in several osteological features; some are likely plesiomorphic, supporting a phylogenetic position outside the crown group. The unusual shape of the deltopectoral crest of the humerus suggests that D. knapptonensis also differed from extant albatrosses in its flight performance. A partial skeleton from the Astoria Formation likely represents another new albatross species; however, it is not named because there is no overlap with the skeletal elements of other fossil Diomedeidae. This species, here informally termed the “...

Triassic turtle tracks and the origin of turtles

Turtle (Testudines) tracks, Chelonipus torquatus, reported from the early Middle Triassic (Anisian) of Germany, and Chelonipus isp. from the late Early Triassic (Spathian) of Wyoming and Utah, are the oldest fossil evidence of turtles, but have been omitted in recent discussions of turtle origins. These tracks provide significant clues as to how early the turtle Bauplan originated. Turtle trackways are quite distinctive: the manus and pes form tracks nearly parallel to the midline and indicate an unusually wide gait in which the trackway width is nearly equal to the stride length. These tracks do not fit what would be expected to be made by Triassic Pappochelys or Odontochelys, a supposed prototurtle and an early turtle, respectively. In contrast, these tracks are consistent with what would be expected from the Triassic turtles Proganochelys and Palaeochersis. The features inferred to be present in Triassic turtle tracks support the notion that Odontochelys is a derived aquatic branch of the turtle stem lineage rather than the ancestral state of all turtles. Chelonipus also resembles the Permian track Pachypes dolomiticus, generally assigned to a pareiasaur trackmaker. These revelations highlight the need to consider all available evidence regarding turtle origins, rather than just the body fossils.

Variation in the pelvic and pectoral girdles of Australian Oligo-Miocene mekosuchine crocodiles with implications for locomotion and habitus.

Australian Oligo–Miocene mekosuchines (Crocodylia; Crocodyloidea) display wide diversity in cranial shape and inferred hunting strategies. Terrestrial habitus has been inferred for these distinctive predators. A direct morphological signal for locomotion can be expected in the postcrania, particularly the pelvic and pectoral girdles. Here we describe fossil materials of the girdles, which chart their morphological variation in the subfamily from Eocene through to Middle Miocene. Over this period, both girdles undergo significant morphological changes. Notably, an enclosed, ventrally orientated acetabulum in the ilium is developed in one lineage. This recapitulates the erect parasagittal configuration of the pelvic limb seen in many Mesozoic crocodylomorph lineages, suggesting consistent use of erect high-walking in these mekosuchines. Other pelves from the same Oligo–Miocene deposits display morphology closer to modern crocodilians, suggesting a partitioning of locomotory strategy among sympatric mekosuchines. Plesiomorphic and derived pelvic girdles are distinguishable by parsimony analysis, and the earliest examples of the mekosuchine pelvis more closely resemble gavialids and alligatorids while latter forms converge on crown group crocodylids in the morphology of the iliac crest. This suggests that a revaluation of the base relationship of Mekosuchinae within Eusuchia is necessary.

Phylogeny, taxonomy and biogeography of Neotropical Quiinoideae (Ochnaceae s.l.)

AbstractAmazonia arguably harbours the highest plant species diversity of all wet tropical regions on earth. There is increasing evidence that this enormous diversity is the product of a rather long and complex process and not of a single cause. Here, we investigated divergence times and diversification dynamics in the Neotropical Quiinoideae (Malpighiales: Ochnaceae s.l.) to explore different evolutionary hypotheses for the accumulation of high species diversity. Additionally, we reappraised the phylogenetic relationships and species circumscriptions using sequences of six plastid and nuclear DNA regions and the most comprehensive taxon sampling of the subfamily to date. Molecular dating indicated that Quiinoideae originated during the late Cretaceous, whereas crown group taxa started to diverge at the Eocene‐Oligocene boundary. The lack of rate shifts in this subfamily supports the “museum” model of diversification. The rather constant species accumulation also suggests that the Pleistocene glacial cycles did not play an important role for the diversification of Quiinoideae. While ancestral area reconstruction was inconclusive, the post‐Miocene divergence times of the west Amazonian species provided some evidence that these areas might not have been available for colonization until the demise of the Lake Pebas and Acre wetlands. New infrageneric groups were inferred, and some species were found to be non‐monophyletic, requiring taxonomic reassessment and demonstrating the need for including multiple individuals per species in species‐level phylogenies.

Fracture resistance of CAD-CAM monolithic ceramic and veneered zirconia molar crowns after aging in a mastication simulator

Statement of problemThe demand for ceramic restorations has increased over the past years, and now various machinable materials can be used for chairside computer-aided design and computer aided manufacturing (CAD-CAM) systems. Limited studies of these new materials make it difficult to evaluate their mechanical performance, advantages, and limitations. PurposeThe purpose of this in vitro study was to investigate the fracture resistance of CAD-CAM monolithic ceramic and veneered zirconia molar crowns after thermomechanical aging. Material and methodsA mandibular first molar tooth was prepared, and 12 different experimental groups were generated (n=10). An aging procedure was performed by subjecting the specimens to 12×105 mechanical cycles and 5000 thermocycles. Survival analysis was performed according to the thermomechanical aging. The fracture resistance (load at fracture) of all specimens was evaluated with 1-way analysis of variance after the means had been compared using the Tukey honest significant difference test and Weibull distributions of the experimental groups (α=.05). ResultsAccording to the results, the differences between the fracture resistance of the groups were found to be significant. Restorations in the dual network ceramic crown group fractured catastrophically during thermomechanical aging. In all groups, the highest value was found for monolithic zirconia, followed by monolithic lithium disilicate crowns (P<.05). Both of the monolithic crowns made of lithium disilicate derivates and zirconia crowns veneered with lithium disilicate ceramic showed no differences from each other (P>.05). The Weibull modulus (m) ranged from 4.22 to 8.58, and conventionally veneered and overpressed zirconia showed the lowest Weibull modulus values among the tested groups, indicating greater variation of the data. ConclusionsThe highest fracture resistance was observed for yttria-stabilized zirconia crowns, followed by lithium disilicate derivates. Veneered zirconia restorations generally showed lower resistance than these ceramics. The dual network ceramic crowns fractured during thermomechanical aging.

Hidden morphological diversity among early tetrapods.

Phylogenetic analysis of early tetrapod evolution has resulted in a consensus across diverse data sets in which the tetrapod stem group is a relatively homogenous collection of medium- to large-sized animals showing a progressive loss of 'fish' characters as they become increasingly terrestrial, whereas the crown group demonstrates marked morphological diversity and disparity. The oldest fossil attributed to the tetrapod crown group is the highly specialized aïstopod Lethiscus stocki, which shows a small size, extreme axial elongation, loss of limbs, spool-shaped vertebral centra, and a skull with reduced centres of ossification, in common with an otherwise disparate group of small animals known as lepospondyls. Here we use micro-computed tomography of the only known specimen of Lethiscus to provide new information that strongly challenges this consensus. Digital dissection reveals extremely primitive cranial morphology, including a spiracular notch, a large remnant of the notochord within the braincase, an open ventral cranial fissure, an anteriorly restricted parasphenoid element, and Meckelian ossifications. The braincase is elongate and lies atop a dorsally projecting septum of the parasphenoid bone, similar to stem tetrapods such as embolomeres. This morphology is consistent in a second aïstopod, Coloraderpeton, although the details differ. Phylogenetic analysis, including critical new braincase data, places aïstopods deep on the tetrapod stem, whereas another major lepospondyl lineage is displaced into the amniotes. These results show that stem group tetrapods were much more diverse in their body plans than previously thought. Our study requires a change in commonly used calibration dates for molecular analyses, and emphasizes the importance of character sampling for early tetrapod evolutionary relationships.

The alleged early Miocene Auk Petralca austriaca is a Loon (Aves, Gaviiformes): restudy of a controversial fossil bird

We restudy the holotype specimen of the alleged fossil auk Petralca austriaca from early Miocene marine deposits of the Austrian locality Pucking, which was considered the earliest European representative of Alcidae (auks). The specimen is a partial skeleton consisting mainly of wing bones on two slabs. A recent re-preparation yielded new data on the skeletal morphology of Petralca, which allow more detailed comparisons with auks and loons. Our study shows that the taxon is clearly distinguished from auks in various skeletal features and can be confidently identified as a loon (Gaviiformes). Petralca resembles Colymboides in overall morphology of the wing bones, and clearly is a stem group representative of Gaviiformes. However, some features indicate that it is more closely related to the crown group taxon Gavia than to the stem group taxa Colymboides and Colymbiculus. Unusually thick bone walls of the limb bones indicate well-developed diving capabilities for Petralca austriaca.

A bothremydid from the middle Cenomanian of Portugal identified as one of the oldest pleurodiran turtles in Laurasia

The relatively complete and well preserved shell of a turtle, from the middle Cenomanian of Nazaré (Portugal), is studied here. It is recognized as a member of the crown group Pleurodira and, more specifically, of Bothremydidae. Pleurodira are one of the two lineages of modern turtles, their origin being in Gondwana. Pleurodira are very abundant in the uppermost Cretaceous record of Europe. However, this new finding is one of the few occurrences in the lower Upper Cretaceous of Laurasia. A single member of Bothremydidae had so far been identified in Portugal: the late Campanian-early Maastrichtian Rosasia soutoi. It was the only Cretaceous turtle identified in that country both at specific and at generic levels, being exclusive of Portugal. The taxon from Nazaré is identified as Algorachelus peregrinus, this form is also present in the contemporaneous beds in Spain, and is the oldest member of Bothremydidae in Laurasia. Algorachelus peregrinus is confirmed here to be a coastal form, which facilitated its spread. The two oldest known bothremydids from Laurasia, the European A. peregrinus and the North American Paiutemys tibert, are compared for the first time. They are recognized as closely related taxa. This study provides new data allowing a more precisely characterization of the oldest so far known dispersal event of Pleurodira in Laurasia, which was performed by an African lineage of Bothremydidae that reached the east coast of the Atlantic Ocean at least in the middle Cenomanian, and the west region of that Ocean at least in the late Cenomanian.

Frullania pinnata spec. nov. (Frullaniaceae, Porellales), a new leafy liverwort in mid-Cretaceous Burmese amber from Myanmar

An inclusion in Cretaceous amber from Myanmar is described as a new fossil species in the extant liverwort genus Frullania. The name Frullania pinnata is proposed for the taxon that is characterized by entire underleaves paired with a conspicuous pinnate branching pattern, two distinctive and stable morphological features. Entire underleaves are known in several extant Frullania species but had not previously been documented in Frullania in amber. The combination of morphological characters in this new fossil species is unknown in any crown group lineage of Frullania, and may very well represent a stem lineage element of the genus. This discovery is important because it expands our understanding of the diversity of Frullaniaceae in the Burmese amber forest as well as important ramifications for the phylogenetic reconstruction of extant Frullania lineages.

Long bone histology of Ophiacodon reveals the geologically earliest occurrence of fibrolamellar bone in the mammalian stem lineage

Shared histological characteristics have been observed in the bone matrix and vascularity between Ophiacodontidae and the later therapsids (Synapsida). Historically, this coincidence has been explained as simply a reflection of the presumed aquatic lifestyle of Ophiacodon or even a sign of immaturity. Here we show, by histologically sampling an ontogenetic series of Ophiacodon humeri, as well as additional material, the existence of fibrolamellar bone (FLB) in the postcranial bones of a pelycosaur. Our findings have reaffirmed what previous studies first described as fast growing tissue, and by proxy, have disproven that the highly vascularized cortex is simply a reflection of young age. This tissue demonstrates the classic histological characteristics of true FLB. The cortex consists of primary osteons in a woven bone matrix and remains highly vascularized throughout ontogeny, providing evidence for fast skeletal growth. Overall, the FLB tissue we have described in Ophiacodon is more advanced or “mammal-like” in terms of the osteonal development, bone matrix, and skeletal growth than what has been described thus far for any other pelycosaur taxon. With regards to the histological record, our results remain inconclusive as to the preferred ecology of Ophiacodon due to a similar cortical vascularity pattern exhibited by other carnivorous pelycosaurs. Our findings have set the evolutionary origins of FLB and high skeletal growth rates back approximately 20 million years to the Early Permian, and by phylogenetic extension perhaps the Late Carboniferous.