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Antiarch placoderm fishes were an abundant component of the Middle Paleozoic vertebrate assemblages. Despite a large number of known taxa and specimens, the morphology and function of the skeletal elements of their jaws is inadequately known. Because of this, questions regarding their feeding modes and their roles in the trophic webs remain open. We present a skeletomuscular model of the antiarch jaw apparatus with an attempt to reconstruct its potential biomechanical function. The position of the upper jaw suborbital bones within the plane of the ventral side of the fish armor is suggested to represent the natural “mouth closed” position. During mouth opening, the suborbitals rotated rostrally with simultaneous depression and inward rotation of the infragnathals. The ball-and-socket jaw articulation might ensure this combined movement. Recently described lower jaw elements of Livnolepis zadonica (Obrucheva, 1983) and Bothriolepis sp. from the Upper Devonian (lower Famennian) of Central Russia demonstrating very deep and porous blades of the oral division of the infragnathals queried the structure of these bones in other antiarchs. Observed porosity reflects intense vascularization to supply blood to a connective tissue underlying a supposed keratinous sheath, which protected and strengthened the jaws, as well as made possible scraping tough food objects, such as thallus algae, from the substrate.Having evolved during the Silurian in the Pan-Cathaysian zoogeographical province, antiarchs migrated to Gondwana during the Emsian and later to Euramerica during the Eifelian. Supposedly, antiarchs became the first macrophytophagous vertebrates occupying the trophic level of primary consumers during the late Silurian–Early Devonian. This event diversified the only previously existing predator–prey interrelationships between filter-feeding agnathans and predatory gnathostomes.

Jensen et al . ( 1 ) question evidence presented of a chambered heart within placoderms, citing its small size and apparently ventral atrium. However, they fail to note the belly-up orientation of the placoderm within one nodule, and the variability of heart morphology within extant taxa. Thus, we remain confident in our interpretation of the mineralized organ as the heart.

A middle Famennian shallow-water vertebrate assemblage is described from the shallow-water shelf deposits exposed in the western Mongolia Hushoot Shiveetiin gol section from the Baruunhuurai Terrane of the Central Asian Orogenic Belt (CAOB). The low diversity, yet abundant chondrichthyan fauna represents a typical shallow-water shark biofacies but lacks the expected protacrodont and phoebodont species, instead comprising almost exclusively of ctenacanthiform shark teeth and a new genus and species Junggarensis ambiguus gen. et sp. nov. This new species appears highly derived, likely occupying the shallow-water niche protacrodont and orodont sharks typically inhabit. The identification of tooth variation in Junggarensis ambiguus gen. et sp. nov. has allowed for the synonymy of previous indeterminate Famennian shark teeth and in turn confirms vertebrate faunal connections between the Central Asian Orogenic Belt and areas along the northern margins of Gondwana.

The origin and early diversification of jawed vertebrates involved major changes to skeletal and soft anatomy. Skeletal transformations can be examined directly by studying fossil stem gnathostomes; however, preservation of soft anatomy is rare. We describe the only known example of a three-dimensionally mineralized heart, thick-walled stomach, and bilobed liver from arthrodire placoderms, stem gnathostomes from the Late Devonian Gogo Formation in Western Australia. The application of synchrotron and neutron microtomography to this material shows evidence of a flat S-shaped heart, which is well separated from the liver and other abdominal organs, and the absence of lungs. Arthrodires thus show the earliest phylogenetic evidence for repositioning of the gnathostome heart associated with the evolution of the complex neck region in jawed vertebrates.

The Australian Mesozoic fish fauna is considered to be depauperate in comparison with fish faunas in the Northern Hemisphere. However, due to its geographical location as a potential radiation cent...

Discoveries from the Late Devonian Gogo Formation, in the Canning Basin, Western Australia have provided insights into the origin and evolution of many unique gnathostome features such as the origins of teeth, internal fertilisation, air-breathing, transitional tissues between bone and cartilage, and insights into the fin to limb transition. Although vertebrate studies have dominated evolutionary work, invertebrate studies have added important insights into the palaeoecology of the site and demonstrated close faunal affinities along the margins of northern Gondwana and China. Geochemical analyses have broadened the understanding of the pathways involved in the exceptional preservation of this Devonian Konservat-Lagerstatte. Fossils from the Gogo Formation show extensive soft tissue preservation through phosphatization recording anatomical details not normally obtained from fossil sites.

The pursuit of simple, yet fair, unbiased, and objective measures of researcher performance has occupied bibliometricians and the research community as a whole for decades. However, despite the diversity of available metrics, most are either complex to calculate or not readily applied in the most common assessment exercises (e.g., grant assessment, job applications). The ubiquity of metrics like the h-index (h papers with at least h citations) and its time-corrected variant, the m-quotient (h-index ÷ number of years publishing) therefore reflect the ease of use rather than their capacity to differentiate researchers fairly among disciplines, career stage, or gender. We address this problem here by defining an easily calculated index based on publicly available citation data (Google Scholar) that corrects for most biases and allows assessors to compare researchers at any stage of their career and from any discipline on the same scale. Our ε′-index violates fewer statistical assumptions relative to other metrics when comparing groups of researchers, and can be easily modified to remove inherent gender biases in citation data. We demonstrate the utility of the ε′-index using a sample of 480 researchers with Google Scholar profiles, stratified evenly into eight disciplines (archaeology, chemistry, ecology, evolution and development, geology, microbiology, ophthalmology, palaeontology), three career stages (early, mid-, late-career), and two genders. We advocate the use of the ε′-index whenever assessors must compare research performance among researchers of different backgrounds, but emphasize that no single index should be used exclusively to rank researcher capability.

Chondrichthyes (Elasmobranchii and Holocephali) are distinguished by their largely cartilaginous endoskeleton that comprises an uncalcified core overlain by a mineralised layer; in the Elasmobranchii (sharks, skates, rays) this mineralisation takes the form of calcified polygonal tiles known as tesserae. In recent years, these skeletal tissues have been described in ever increasing detail in sharks and rays but those of Holocephali (chimaeroids) have been less well-described, with conflicting accounts as to whether or not tesserae are present. During embryonic ontogeny in holocephalans, cervical vertebrae fuse to form a structure called the synarcual. The synarcual mineralises early and progressively, anteroposteriorly and dorsoventrally, and therefore presents a good skeletal structure in which to observe mineralised tissues in this group. Here we describe the development and mineralisation of the synarcual in an adult and stage 36 elephant shark embryo (Callorhinchus milii). Small, discrete, but irregular blocks of cortical mineralisation are present in stage 36, similar to what has been described recently in embryos of other chimaeroid taxa such asHydrolagus, while inCallorhinchusadults, the blocks of mineralisation have become more irregular, but remain small. This differs from fossil members of the holocephalan crown group (Edaphodon), as well as from stem group holocephalans (e.g., Symmorida,Helodus, Iniopterygiformes), where tessellated cartilage is present, with tesserae being notably larger than inCallorhinchusand showing similarities to elasmobranch tesserae, for example with respect to polygonal shape.

Burrow, C.J., Turner, S., Trinajstic, K. & Young, G.C., 27 February 2019. Late Silurian vertebrate microfossils from the Carnarvon Basin, Western Australia. Alcheringa 43, 204–219. ISSN 0311-5518.A...

The study of early actinopterygians (ray-finned fishes) from the Devonian has been hampered by imperfect preservation in the majority of taxa. The Late Devonian (early Frasnian) Gogo Formation of north-western Western Australia is notable in producing complete fossil actinopterygians with exceptional three-dimensional preservation of both the dermal and endoskeletal anatomy. Four taxa have been described and have proved invaluable in understanding the anatomy of early representatives of this clade. Here, we present a fifth Gogo taxon, based on a single exceptionally preserved specimen and representing a new genus and species of early ray-finned fish. The neurocranium was CT scanned, permitting a detailed examination of the endocast. The new taxon possesses unusual features including a greatly enlarged spiracular opening and extensive spinose ornamentation on the dermal skull bones, median ridge scutes and lepidotrichia. The endocast displays a mosaic of characters, some of which are similar to Mimipiscis and non-actinopterygian outgroups, while other features are more akin to Raynerius from the Late Devonian of France as well as several stratigraphically younger taxa. A phylogenetic analysis resolves the new form as sister taxon to fishes from the Late Devonian of the Northern Hemisphere and all post-Devonian actinopterygians, confirming that the assemblage of fossil ray fins from the Gogo Formation is part of the wider Devonian actinopterygian radiation .

Significant new material of Late Devonian Gogo Formation fish fossils is still surfacing. Collecting in the past decade has uncovered the first Gogo shark fossils (Gogoselachusplus another new undescribed taxon), the first acanthodian (Halmacanthodes ahlbergi), the first coelacanth, as well as the first placoderm embryos. Recent studies have elucidated the nature of placoderm claspers, pelvic girdles, synarcuals and embryos, the structure of their teeth, a description of well-preserved muscles in placoderms, and how muscles attach to bones. Molecular biomarkers have also been identified in Gogo fossils. There are now five basal ray-fin fishes in the fauna, including one undescribed new taxon. The lungfish fauna from Gogo is the most diverse known for any Devonian site, with 10 genera and 12 species. The dermal skeleton and endocast of the dipteridRhinodipterus kimberleyensishave been described in detail from CT scans; and the ontogenetic stages of neurocranium formation inGriphognathus. New specimens of the tetrapodomorph fishGogonasus andrewsaehave shed further light on its endocranium, pectoral girdle and fin. Through their exceptional preservation of both hard and varied kinds of soft tissues, the Gogo fishes remain crucial for resolving key debates on the diversification, physiology, biomechanics and phylogenetic relationships of early gnathostomes.

Many species of lizard use caudal autotomy, the ability to self-amputate a portion of their tail, regenerated over time, as an effective anti-predation mechanism. The importance of this tactic for survival depends on the degree of predation risk. There are, however, negative trade-offs to losing a tail, such as loss of further autotomy opportunities with the regenerated tail vertebrae being replaced by a continuous cartilaginous rod. The common consensus has been that once a tail has been autotomised and regenerated it can only be autotomised proximal to the last vertebral autotomy point, as the cartilage rod lacks autotomy planes. However, anecdotal evidence suggests that although the regenerated portion of the tail is unable to autotomise, it can re-regenerate following a physical shearing event. We assessed re-regeneration in three populations of the King’s skink (Egernia kingii), a large lizard endemic to south-west Western Australia and surrounding islands. We show that re-regeneration is present at an average of 17.2% across the three populations, and re-regenerated tissue can comprise up to 23.3% of an individual’s total tail length. The ability to re-regenerate may minimise the costs to an individual’s fitness associated with tail loss, efficiently restoring ecological functions of the tail.

Conodont biogenic apatite has become a preferred analytical target for oxygen isotope studies investigating ocean temperature and palaeoclimate changes in the Palaeozoic. Despite the growing application in geochemically-based palaeoenvironmental reconstructions, the paucity or absence of conodont fossils in certain facies necessitates greater flexibility in selection of robust oxygen-bearing compounds for analysis. Vertebrate microfossils (teeth, dermal denticles, spines) offer a potential substitute for conodonts from the middle Palaeozoic. Vertebrate bioapatite is particularly advantageous given a fossil record extending to the present with representatives across freshwater to fully marine environments, thus widening the scope of oxygen isotope studies on bioapatite. However, significant tissue heterogeneity within vertebrates and differential susceptibility of these tissues to diagenetic alteration have been raised as potential problems affecting the reliability of the oxygen isotope ratios as palaeoclimatic proxies. Well-preserved vertebrate microfossils and co-occurring conodont fossils from the Upper Devonian and Lower Carboniferous of the Lennard Shelf, Canning Basin, Western Australia, were analysed using bulk (gas isotope ratio mass spectrometry, GIRMS) and in-situ (secondary ion mass spectrometry, SIMS) methodologies, with the latter technique allowing investigation of specific tissues within vertebrate elements. The δ18Oconodont results may be interpreted in terms of palaeolatitudinally and environmentally sensible palaeo-salinity and -temperature and provide a baseline standard for comparison against vertebrate microfossil δ18O values. Despite an absence of obvious diagenetic modification, GIRMS of vertebrate denticles yielded δ18O values depleted in 18O by 2–4‰ relative to co-occurring conodonts. SIMS analysis of dentine tissues exhibited significant heterogeneity, while hypermineralised tissues in both scales and teeth produced δ18O values comparable with those of associated conodonts. The susceptibility of permeable phosphatic fossil tissues to microbial activity, fluid interaction and introduction of mineral precipitates post-formation is demonstrated in the dentine of vertebrate microfossils, which showed significant heterogeneity and consistent depletion in 18O relative to conodonts. The hypermineralised tissues present in both teeth and scales appear resistant to many diagenetic processes and indicate potential for palaeoclimatic reconstructions and palaeoecological investigations.

The evolution of jaws marked an important transition in the evolution of vertebrates, allowing them to become successful, active predators. Some developmental events associated with this transition can be deduced from the fossil record of jawless vertebrates, with new discoveries suggesting that the separation of the nasal sacs in jawless vertebrates was a key step in the evolution of jaws. Some members of the earliest jawed vertebrates, the placoderms, have recently been shown to have teeth and hypobranchial muscles similar to those in extant chondrichthyans. Associated with the evolution of jaws was the evolution of a moveable neck, which allowed the first jawed vertebrates to raise their heads and increase the size of their mouths, making it more efficient for food and water intake. The evolution of jaws and associated musculature enabled gnathostomes to occupy new niches and to radiate, leading to the great diversity observed today. New fossil discoveries continue to populate the gnathostome stem, adding to our understanding of the early diversification of this successful group of vertebrates.

Chondrichthyans are one of two major clades of living jawed vertebrates, with a rich fossil record potentially extending back to the Late Ordovician (455 million years ago, mya). The main groups of chondrichthyans include the chimaeroids, sharks, and skates and rays. This chapter outlines the major events in chondrichthyan evolution, focusing on features of the cranium, jaw and jaw musculature, and gill arch skeleton. The “spiny sharks” (acanthodians) and other stem chondrichthyans have recently been shown to exhibit a mosaic of chondrichthyan and osteichthyan characters. Taxa such as iniopterygians and chondrenchelyiforms, resolved as stem group chimaeroids, appear in the Carboniferous and display dramatic body forms and unusual fin morphology. Chondrichthyans also show a considerable range of dentitions, both in terms of morphology and development, particularly modified in the chimaeroids. In addition to their differing tooth morphologies, chondrichthyans have several types of jaw suspensions to support a range of feeding and breathing modes. Sharks have well-developed brains that vary according to the environment rather than phylogeny. Their senses are also well-developed and finely tuned to best perform in their particular ecological niche. The long evolutionary history of chondrichthyans and their great diversity as well as the retention of some primitive characters make them good models for evolutionary and developmental studies.

A diverse microvertebrate fauna is described from the Virgin Hills and Napier formations, Bugle Gap Limestone Canning Basin, Western Australia. Measured sections at Horse Spring and Casey Falls (Virgin Hills Formation) and South Oscar Range (Napier Formation) comprise proximal to distal slope carbonates ranging in age from the Late Devonian Frasnian to middle Famennian. A total of 18 chondrichthyan taxa are identified based on teeth, including the first record of Thrinacodus tranquillus, Cladoides wildungensis, Protacrodus serra and Lissodus lusavorichi from the Canning Basin. A new species, Diademodus dominicus sp. nov. is also described and provides the first record of this genus outside of Laurussia. In addition, the upper range of Australolepis seddoni has been extended to Late Devonian conodont Zone 11, making it the youngest known occurrence for this species. The Virgin Hills and Napier formations microvertebrate faunas show close affinities to faunas recovered from other areas of Gondwana, including eastern Australia, Iran, Morocco and South China, which is consistent with known conodont and trilobite faunas of the same age.

a b s t r a c t The Late Devonian was a time of major evolutionary change encompassing the fifth largest mass extinction, the Frasnian-Famennian event. In order to establish a chronological framework for global correlation before, during, and following the Frasnian-Famennian mass extinction, we carried out a coupled magnetostratigraphic and biostratigraphic study of two stratigraphic sections in the Upper Devonian carbonate reef complexes of the Lennard Shelf, in the Canning Basin, Western Australia. Magnetostratigraphy from these rocks provides the first high-resolution definition of the Late Devonian magnetic polarity timescale. A 581-m-reference section and an 82-m overlapping section through the marginal slope facies (Napier Formation) of the Oscar Range as well as a 117-m section at Horse Spring (Virgin Hills Formation) were sampled at decimeter to meter scale for magnetostratigraphy. Conodont biostratigraphy was used to correlate both sections, and link magnetostratigraphic polarity zones to a globally established biostratigraphy. As table, Characteristic Remanent Magnetization (ChRM) with dual polarities (NE, shallowly upward and SW, shallowly downward) is recovered from ∼60% of all samples, with magnetite inferred to be the chief magnetic carrier from thermal demagnetization characteristics. These directions define a geomagnetic pole at 49.5 ◦ S/285.8 ◦ E and α95 = 2. 4( n = 501), placing the Canning Basin at 9.9 ◦ S during the Late Devonian, consistent with carbonate reef development at this time. Ac onservative interpretation of the magnetostratigraphy shows the recovery of multiple reversals from both sections, not including possible cryptochrons and short duration magnetozones. Field tests for primary remanence include positive reversal tests and matching magnetozones from an overlapping section in the Oscar Range. As trong correlation was found between magnetic polarity stratigraphies of the Oscar Range and Horse Spring sections, and we correlate 12 magnetostratigraphic packages. The relative stratigraphic thicknesses of the isochronous sediments from these two sections indicate that carbonate accumulation was ∼4.5× faster in the middle slope deposits at Oscar Range than in the more distal, lower slope Horse Spring deposits for the middle Frasnian through Famennian. The magnetic field during the Late Devonian underwent a relatively high reversal frequency with good potential for regional and global correlation, and should prove useful in deciphering a high-resolution chronostratigraphy across the Lennard Shelf to enable higher confidence examination of reef development across a major biotic crisis.

High-resolution, time-significant correlations are integral to meaningful stratigraphic frameworks in depositional systems, but may be difficult to achieve using traditional sequence stratigraphic or biostratigraphic approaches alone, particularly in geologically complex settings. In steep, reefal carbonate margin-to-slope systems, such correlations are essential to unravel shelf-to-basin transitions, characterize strike variability, and develop predictive sequence stratigraphic models – concepts which are currently poorly understood in these heterogeneous settings. The Canning Basin Chronostratigraphy Project (CBCP) integrates multiple independent datasets (including biostratigraphy, magnetostratigraphy, stable isotope chemostratigraphy, and sequence stratigraphy) extracted from Upper Devonian (Frasnian and Famennian) reefal platform exposures along the Lennard Shelf, Canning Basin, Western Australia. These were used to generate a well-constrained stratigraphic framework and shelf-to-basin composite reconstruction of the carbonate system. The resultant integrated framework allows for unprecedented analysis of carbonate margin-to-slope heterogeneity, depositional architecture, and sequence stratigraphy along the Lennard Shelf. Systems tract architecture, facies partitioning, and stacking patterns of margin to lower-slope environments were assessed for six composite-scale sequences that form part of a transgressive-to-regressive supersequence and span the Frasnian-Famennian (F-F) biotic crisis. Variations are apparent in margin styles, foreslope facies proportions, dominant resedimentation processes, downslope contributing sediment factories, and vertical rock successions, related to hierarchical accommodation signals and ecological changes associated with F-F boundary. We present these results in the form of carbonate margin-to-basin sequence stratigraphic models and associations that link seismic-scale architecture to fine-scale facies heterogeneity. These models provide a predictive foundation for characterization of steep-sided flanks of reefal carbonate platform systems that is useful for both industry and academia. This study emphasizes the utility of an integrated stratigraphic approach and the insights gained from better-constrained facies and stratal architecture analysis; insights that were not achievable with traditional sequence stratigraphic or biostratigraphic techniques alone.

Reproduction in jawed vertebrates (gnathostomes) involves either external or internal fertilization. It is commonly argued that internal fertilization can evolve from external, but not the reverse. Male copulatory claspers are present in certain placoderms, fossil jawed vertebrates retrieved as a paraphyletic segment of the gnathostome stem group in recent studies. This suggests that internal fertilization could be primitive for gnathostomes, but such a conclusion depends on demonstrating that copulation was not just a specialized feature of certain placoderm subgroups. The reproductive biology of antiarchs, consistently identified as the least crownward placoderms and thus of great interest in this context, has until now remained unknown. Here we show that certain antiarchs possessed dermal claspers in the males, while females bore paired dermal plates inferred to have facilitated copulation. These structures are not associated with pelvic fins. The clasper morphology resembles that of ptyctodonts, a more crownward placoderm group, suggesting that all placoderm claspers are homologous and that internal fertilization characterized all placoderms. This implies that external fertilization and spawning, which characterize most extant aquatic gnathostomes, must be derived from internal fertilization, even though this transformation has been thought implausible. Alternatively, the substantial morphological evidence for placoderm paraphyly must be rejected., Travel to Tallinn, London and Washington for J.A.L. to examine specimens was supported by Flinders University. G.C.Y., J.A.L. and K.T. acknowledge support from the Australian Research Council. ARC DP1092870

Newly discovered pelvic and reproductive structures within placoderms, representing some of the most crownward members of the gnathostome stem group and the most basal jawed vertebrates, challenge established ideas on the origin of the pelvic girdle and reproductive complexity. Here we critically review previous descriptions of the pelvic structures in placoderms and reinterpret the morphology of the pelvic region within the arthrodires and ptyctodonts, in particular the position of the pelvic fin and the relationship of the male clasper to the pelvic girdle. Absence of clear articular surfaces on the clasper and girdle in the Arthrodira, along with evidence from the Ptyctodontida, suggest that these are separate structures along the body. We describe similarities between the pectoral and pelvic girdles and claspers, for example, all these have both dermal and perichondral (cartilaginous) components. Claspers in placoderms and chondrichthyans develop in very different ways; in sharks, claspers develop from the pelvic fin while the claspers in placoderms develop separately, suggesting that their independent development involved a posterior extension of the 'competent stripes' for fin development previously limited to the region between the paired pectoral and pelvic fins. Within this expanded zone, we suggest that clasper position relative to the pelvic fins was determined by genes responsible for limb position. Information on early gnathostome reproductive processes is preserved in both the Ptyctodontida and Arthrodira, including the presence of multiple embryos in pregnant females, embryos of differing sizes and of different sexes (e.g. male claspers preserved in some embyros). By comparison with chondrichthyans, these observations suggest more complex reproductive strategies in placoderms than previously appreciated.

The close association of anoxic or dysoxic sedimentary rocks and the major Late Devonian (Frasnian–Famennian) mass extinction has focused considerable attention on anoxia as the major cause or as a major factor in a multicausal scenario. The record of the Late Devonian biotic crisis in the well-known reef complexes of northwestern Australia (Canning Basin), in contrast to many localities elsewhere, does not display sedimentological evidence of anoxia through the Frasnian–Famennian boundary interval. Analysis of continuous drill core through this interval has yielded three positive δ 13 C isotopic excursions, only one of which coincides with total organic carbon (TOC) maxima in our data. Multi-element geochemical proxies suggest that TOC maxima preceding positive shifts in δ 13 C most likely resulted from higher productivity caused by nutrient influx from continental weathering, given the close association between TOC maxima and regional relative sea-level falls. Our interpretation supports the view that anoxia was not a fundamental driver of mass extinction and stresses the importance of integrated data sets and understanding regional controls on environmental changes and/or stresses.

Facies architecture and platform evolution of an early Frasnian reef complex in the northern Canning Basin of north-western Australia were strongly controlled by syn-depositional faulting during a phase of basin extension. The margin-attached Hull platform developed on a fault block of Precambrian basement with accommodation largely generated by movement along the Mount Elma Fault Zone. Recognition of major subaerial exposure and flooding surfaces in the Hull platform (from outcrop and drillcore) has enabled comparison of facies associations within a temporal framework and led to identification of three stages of platform evolution. Stage 1 records initial ramp development on the hangingwall dip slope with predominantly deep subtidal conditions that prevented any cyclic facies arrangements. This stage is characterised by basal siliciclastic deposits and a major deepening-upward facies pattern that is capped by a sequence boundary towards the footwall (north-west) and a major flooding surface towards the hangingwall. Stage 2 reflects the bulk of platform aggradation, significant platform growth towards the hangingwall and the development of reef margins and cyclic facies arrangements. Thickening of this stage towards the hangingwall indicates that accommodation was generated by rotation of the fault block and overlying platform. Stage 3 records a major flooding and backstep of the platform margin. The Hull platform illustrates important elements of margin-attached carbonate platforms in a half-graben setting, including: (i) prominent, but limited, coarse siliciclastic input that does not have a major detrimental effect on carbonate production near the rift margin in arid to semi-arid settings; (ii) wedge-shaped accommodation created by syn-depositional rotation of fault blocks and tilting of the hangingwall dip slope, resulting in shallow-water facies and subaerial exposure up-dip of the rotational axis and deeper water facies down-dip; and (iii) evolution of a ramp to rimmed shelf, coincident with a sequence boundary–flooding surface, that is accelerated by tilting of the hangingwall dip slope during fault-block rotation.

Royal Society Open Science 4(7), 170214 (2017). doi:10.1098/rsos.170214, Published by Royal Soc. Publ., London

Photic zone euxinia (PZE) has proven important for elucidating biogeochemical changes that occur during oceanic anoxic events, including mass extinction and conditions associated with unique fossil preservation. Organic geochemical analyses of a 380 Ma invertebrate fossil, which included well-preserved soft tissues, from the Gogo Formation (Canning Basin, Western Australia) showed biomarkers and stable isotopic values characteristic of PZE and a consortium of sulfate-reducing bacteria, which lead to exceptional fossil and biomarker preservation. The carbonate concretion contained phytoplankton, green sulfur bacteria (Chlorobi), and sulfate-reducing bacteria biomarkers with an increasing concentration toward the nucleus where the fossil is preserved. The spatial distribution of cholestane unequivocally associated with the fossilized tissue and its high relative abundance to the total steranes suggest that the fossil is a crustacean. The presence of an active sulfur cycle in this Devonian system, including sulfate reduction and the resulting PZE, played a pivotal role in the preservation of soft tissue from the fossil and its associated low-maturity biomarker ratios.

A new acanthodiform acanthodian Halimacanthodes ahlbergi n. gen. n. sp., based on a single uncrushed, partial articulated specimen, represents the first acanthodian collected from the Gogo Formation, Canning Basin, Western Australia. The delicate perichondral ossifications of the lower jaws, branchial skeleton and endoskeletal shoulder girdle are preserved in 3D. H. ahlbergi n. gen. n. sp. is characterised by having a tall slender scapular shaft, paired pelvic fin spines, mineralised branchial arches and scales with an anterior median depression and a microornament of tiny smooth tubercles randomly distributed on the crown. In having paired pelvic spines and lacking prepelvic spines, it is most closely related to Howittacanthus kentoni from Frasnian lacustrine deposits of eastern Victoria. Isolated scales, here attributed to the new taxon, are found in the coeval Gneudna Formation, Carnarvon Basin, Western Australia.

The synarcual is a structure incorporating the anterior vertebrae of the axial skeleton and occurs in vertebrate taxa such as the fossil group Placodermi and the Chondrichthyes (Holocephali, Batoidea). Although the synarcual varies morphologically in these groups, it represents the first indication, phylogenetically, of a differentiation of the vertebral column into separate regions. Among the placoderms, the synarcual of Cowralepis mclachlani Ritchie, 2005 (Arthrodira) shows substantial changes during ontogeny to produce an elongate, spool-shaped structure with a well-developed dorsal keel. Because the placoderm synarcual is covered in perichondral bone, the ontogenetic history of this Cowralepis specimen is preserved as it developed anteroposteriorly, dorsally and ventrally. As well, in the placoderm Materpiscis attenboroughi Long et al., 2008 (Ptyctodontida), incomplete fusion at the posterior synarcual margin indicates that both neural and haemal arch vertebral elements are added to the synarcual. A survey of placoderm synarcuals shows that taxa such as Materpiscis and Cowralepis are particularly informative because perichondral ossification occurs prior to synarcual fusion such that individual vertebral elements can be identified. In other placoderm synarcuals (e.g. Nefudina qalibahensis Lelievre et al., 1995; Rhenanida), cartilaginous vertebral elements fuse prior to perichondral ossification so that individual elements are more difficult to recognize. This ontogenetic development in placoderms can be compared to synarcual development in Recent chondrichthyans; the incorporation of neural and haemal elements is more similar to the holocephalans, but differs from the batoid chondrichthyans.

A full description of a complete and articulated, three-dimensionally preserved, placoderm fish, Materpiscis attenboroughi (Ptyctodontida), from the Late Devonian Gogo Formation is presented. The jaw articulation is unique in being preserved in situ, providing new morphological information on the articulation of the quadrate, metapterygoid, and labial skeleton. In addition, separate hyoid and opercular cartilages are identified, thus confirming their presence for the first time within placoderms. Preserved orbital ossifications indicate an anteriomesial orientation of this braincase region. The complete scapulocoracoid provides additional morphological information on the pectoral fin articulation. A new specimen of Austroptyctodus gardineri preserving the anterior portion of the head and jaws allows for revised descriptions of the endocranium and visceral skeleton, including the most complete endocranial ossifications recovered to date. As well, elements of the labial skeleton not known previousl...

The present paper involves a detailed evaluation of specific steroid biomarkers by gas chromatography–mass spectrometry (GC–MS) and GC-metastable reaction monitoring (MRM) analyses of several crude oils and source rocks from the East Sirte Basin. 24-Norcholestanes, dinosteranes, 4α-methyl-24-ethylcholestanes and triaromatic steroids have been identified in both source-rocks and crude oils of the East Sirte Basin. Diatoms, dinoflagellates (including those potentially associated with corals) and/or their direct ancestors are amongst the proposed sources of these biomarkers. These biomarker parameters have been used to establish a Mesozoic oil–source correlation of the East Sirte Basin. Hydropyrolysis of an extant coral extract revealed a similar distribution (although immature) of dinosteranes and 4α-methyl-24-ethylcholestanes also observed in the Sirte oils and source-rocks. This is consistent with the presence of dinoflagellates present during the deposition of the Mesozoic aged East Sirte Basin Formations. A good data correlation for the rock extracts revealed a similar distribution of 3,24-dimethyl triaromatic steroids, 3-methyl-24-ethylcholestanes, 4-methyl-24-ethylcholestanes and 2-methyl-24-ethylcholestanes observed in one of the oil families and associated source-rocks for the East Sirte Basin.

Biomarker ratios, together with stable carbon (δ 13 C) and hydrogen (δD) isotopic compositions of individual hydrocarbons have been determined in a suite of crude oils ( n = 24) from the East Sirte Basin to delineate their sources and respective thermal maturity. The crude oil samples are divided into two main families (A and B) based on differences in source inputs and thermal maturity. Using source specific parameters including pristane/phytane (Pr/Ph), hopane/sterane, dibenzothiophene/phenanthrene (DBT/P), Pr/ n -C 17 and Ph/ n -C l8 ratios and the distributions of tricyclic and tetracyclic terpanes, family B oils are ascribed a marine source rock deposited under sub-oxic conditions, while family A oils have a more terrigenous source affinity. This genetic classification is supported by the stable carbon isotopic compositions (δ 13 C) of the n -alkanes. Using biomarker maturity parameters such as the abundance of Pr and Ph relative to n -alkanes and the distribution of sterane and hopane isomers, family A oils are shown to be more thermally mature than family B oils. The contrasting maturity of the two families is supported by differences between the stable hydrogen isotopic compositions (δD) of Pr and Ph and the n -alkanes, as well as the δ 13 C values of n -alkanes in their respective oils.

The Gogo Formation of Western Australia preserves a unique Late Devonian (Frasnian) reef fauna. The exceptional three-dimensional preservation of macrofossils combined with unprecedented soft-tissue preservation (including muscle bundles, nerve cells, and umbilical structures) has yielded a particularly rich assemblage with almost 50 species of fishes described. The most significant discoveries have contributed to resolving placoderm phylogeny and elucidating their reproductive physiology. Specifically, these discoveries have produced data on the oldest known vertebrate embryos; the anatomy of the primitive actinopterygian neurocranium and phylogeny of the earliest actinopterygians; the histology, radiation, and plasticity of dipnoan (lungfish) dental and cranial structures; the anatomy and functional morphology of the extinct onychodonts; and the anatomy of the primitive tetrapodomorph head and pectoral fin.

An almost complete but predominantly disarticulated ptyctodont fish, Kimbryanodus williamburyensis n. gen., n.sp. from the Late Devonian Gneudna Formation, is described. The fossils occur as three-dimensionally preserved isolated plates, and this has allowed the reconstruction of the fish. A taxonomic revision of the ptyctodonts was undertaken based on recently described Australian taxa and new reconstructions of Australian, American and European specimens. The phylogenetic analysis supports a threefold division of the ptyctodonts, with Rhamphodopsis being the most basal taxon and the other ptyctodonts divided into those possessing a median dorsal spine, spinal plate and simple V-shaped overlap of the anterior lateral and anterior dorsolateral plates and those taxa which do not.

Intraspecies variation, polymorphism and asymmetric traits are observed within two families of Arthrodira, the Incisoscutidae and Camuropiscidae, from the Gogo Formation in northern Western Australia. Individual plates of the head and trunk shield show considerable variation between individuals. Plates that show the greatest degree polymorphic traits are the rostral (R), marginal (M), submarginal (SM), preorbital (PrO), anterior dorsolateral, anterior median ventral (AMV) and posterior ventrolateral (PVL) plates. The paths of the sensory line canals are the most variable feature and the dermal plates of the cheek show the greatest asymmetry. It is apparent that if anatomical data in arthrodires are to be interpreted with greater precision, detailed knowledge of intraspecies variation, polymorphic and asymmetric traits is essential. How these variables are treated in cladistic analysis is also critical. Here multistate characters were coded differently in five discrete analyses, each analysis yielding a different number of trees and relationships. It was concluded that including and coding for multistate characters gave the most robust tree. In addition, further morphological characters from a new specimen of Gogosteus sarahae Long (1994) indicates many of the characters used to separate this genus from Incisoscutum are inconsistent and so it is here considered that the genus Gogosteus is a junior synonym of Incisoscutum.