Middle Palaeozoic Research Articles

A Devonian crinoid with a diamond microlattice.

Owing to their remarkable physical properties, cellular structures, such as triply periodic minimal surfaces (TPMS), have multidisciplinary and multifunctional applications. Although these structures are observed in nature, examples of TPMS with large length scales in living organisms are exceedingly rare. Recently, microstructure reminiscent of the diamond-type TPMS was documented in the skeleton of the modern knobby starfish Protoreaster nodosus. Here we report a similar microlattice in a 385 Myr old crinoid Haplocrinites, which pushes back the origins of this highly ordered microstructure in echinoderms into the Devonian. Despite the low Mg2+/Ca2+ ratio of the 'calcite' Devonian sea, the skeleton of these crinoids has high-Mg content, which indicates strong biological control over biomineralogy. We suggest that such an optimization of trabecular arrangement additionally enriched in magnesium, which enhances the mechanical properties, might have evolved in these crinoids in response to increased predation pressure during the Middle Palaeozoic Marine Revolution. This discovery illustrates the remarkable ability of echinoderms, through the process of evolutionary optimization, to form a lightweight, stiff and damage-tolerant skeleton, which serves as an inspiration for biomimetic materials.

Early-Middle Devonian adakitic magmatism generated by slab retreat in southern West Junggar, NW China: implications for tectonic correlation with central and East Kazakhstan

ABSTRACT The West Junggar terrane (WJT) is known to tectonically correlate with the Central and East Kazakhstan terrane; however, the Middle Palaeozoic magmatism has rarely been reported in the southern WJT, leading to significant uncertainties on the tectonic evolution and correlation of the southwestern Central Asian Orogenic Belt (CAOB). Here, we present the adakitic quartz diorites with zircon U-Pb ages of 390–394 Ma in the Alashankou region to demonstrate the presence of Early-Middle Devonian magmatism in the southern WJT for the first time. They are characterized by low Yb (1.00–1.41 ppm) and Y (8.9–14.4 ppm) but high Sr (301.5–475.9 ppm), Sr/Y (23–41), and (La/Yb)N (8.1–11) values, typical of modern adakites. As suggested by the relatively high SiO2 (64.96–67.71 wt.%), moderate K2O (1.53–3.19 wt.%), but low MgO (1.56–2.31 wt.%), TiO2 (0.28–0.42 wt.%), Cr (≤13.7 ppm), Co (2.7–9.9 ppm), Ni (8.0–18.7 ppm), and Th (3.7–4.8 ppm) concentrations as well as the high positive ɛHf(t) values (+5.6 to +10.3) and low Nb/Ta ratios, the adakitic quartz diorites were probably generated by the partial melting of a slightly thickened lower crust, which consisted mainly of the Ediacaran-Middle Palaeozoic intra-oceanic arc rocks in the southern WJT. Combined with regional data, our new results support that the southern WJT probably evolved as the eastern extension of the Kazakhstan Early-Middle Devonian outer volcanic belt during the Middle Palaeozoic. The Early-Middle Devonian tectonic extension and resultant lower crust melting may thus be related to the slab retreat of the Junggar-Balkhash Ocean.

In the shadow of a giant reef: Palaeoecology of mesophotic coral communities from the Givetian of Anti-Atlas (Morocco)

One of the keys to the success of modern and ancient reef ecosystems is the phenomenon of photosymbiosis. Photosymbiotic corals can thrive down to the limits of the photic zone, and the coral communities living in the lower part of the photic zone are known as mesophotic coral reef ecosystems (MCE). Recent studies have shown that MCEs were already present in the middle Palaeozoic. The reefs of that time reached the peak of their distribution during the Givetian, when they were present even as far as 50°S. One of the southernmost Givetian reefs is the Aferdou el Mrakib reef, exposed in the eastern Anti-Atlas (Morocco). At the base of this reef, numerous small carbonate bioherms are exposed. These structures formed low-relief banks that developed probably at moderate water depths of several tens of meters. In the present contribution, we analyse the paleoecology of these peculiar communities. The assemblage is dominated by platy and encrusting coenitid (Roseoporella) and alveolitid (Alveolites) tabulate corals, as well as abundant platy stromatoporoids and solitary cystyphillid rugose corals. The platy morphology of corals is widely considered as an adaptation related to photosymbiosis, and is especially common in MCEs. In combination with palaeobathymetric and sedimentological data, this characteristic suggests that the studied banks developed at the bottom of the euphotic zone. The MCEs of the Aferdou el Mrakib reef share major taxonomic and ecological similarities with the previously described MCEs from the southern shelf of Laurussia and a reef community from southern China, emphasizing a high degree of cosmopolitanism of the Middle Devonian mesophotic ecosystems. Communities studied here are also the most poleward MCEs described to date.

Stranger than a scorpion: a reassessment of Parioscorpio venator, a problematic arthropod from the Llandoverian Waukesha Lagerstätte

AbstractA relatively uncommon arthropod of the Waukesha lagerstätte, Parioscorpio venator, is redescribed as an arthropod bearing a combination of characters that defy ready classification. Diagnostic features include sub‐chelate ‘great appendages’, a lack of antennae, multiramous anterior trunk appendages, filamentous fan‐like rear trunk appendages, and apparently thin and poorly preserved pleural fields. Phylogenetic analysis resolves this organism as basal to crown‐group Mandibulata and Chelicerata, but its exact placement is inconclusive. Thus, we compare its morphology to several stem groups of arthropods in a discussion of its plausible taxonomic affinities. The examined specimens are probably carcasses and preserve a variety of soft‐tissue details, including muscle blocks in the head, eyes and eye facets, likely ventral nerve cords, a central gut tract and trunk legs with multiple filamentous elements organized into stiff bundles. The preservation habits of P. venator are characterized and compared to previous assessments of Waukesha lagerstätte taxa. Four preservation habits are observed: a phosphatized habit showing flattened to partly three‐dimensional mineralization in francolite; a mouldic habit largely left behind by removed francolite that shows no carbon enrichment despite a darkened colour; sheet‐like or speckled carbonaceous compressions; and scattered pyrite crystals. This redescription highlights both the palaeobiological value of ‘small’ lagerstätten typical of the middle Palaeozoic and the caution that must be taken when interpreting their more enigmatic constituents.

Petrology and geochronology of the San Martín de los Andes batholith: Insights into the Devonian magmatism of the North Patagonian Andes

Evidence is growing that a belt of Devonian plutonism is present in the Northern Patagonian segment of the present-day Andes. This plutonism formed along the paleo-Pacific margin of the Gondwana supercontinent. New whole rock chemistry and SHRIMP U–Pb zircon data for plutonic rocks of the San Martín de los Andes batholith (southwestern Argentina) provide insights into the magmatic and tectonic evolution of the southwestern Gondwana margin during the middle Palaeozoic. The San Martín de los Andes batholith was formed between 390 and 374 Ma and intruded into a Lower Paleozoic metamorphic basement that had been previously affected by Late Ordovician-middle Silurian deformation. Most of the batholith is composed of metaluminous, calc-alkaline tonalites and granodiorites, with subordinate volume of peraluminous garnet-bearing granites. Discrete and disperse gabbroic and dioritic rocks hosted in tonalites and granodiorites occur as either microgranular enclaves or syn-magmatic dykes. Tonalitic, granodioritic and granitic rocks develop transitional contacts among them revealing coetaneous building up throughout the plutonic suite, but when interpreting their chemical variability, they are not strictly comagmatic rocks. Collectively, field, petrographic, and geochemical data of the San Martín de los Andes batholith are interpreted to reflect concurrence of crystal fractionation, crustal assimilation and magma mixing in driving petrological diversification. Yet, gabbroic and dioritic rocks in the San Martín de los Andes batholith display geochemical characteristics of calc-alkaline primitive basalts that concur with what is being hypothesized to be a magmatic arc zone for the Devonian period. This study discusses how to interpret the evolution of the San Martín de Los Andes batholith together with other geodynamic models such as the Chaitenia magmatic arc.

Dictyodora-Nereites dominated ichnofauna in the Silurian−Devonian turbidite-like deposits of central-western Gondwana (San Rafael Block, Argentina): New stratigraphic and bathymetric constraints

The Río Seco de los Castaños Formation (San Rafael Block) holds one of the few middle Palaeozoic (Silurian−Devonian) records worldwide of a Nereites-Dictyodora dominated ichnofauna with sedimentological features such as oscillatory and combined-flow deposits. This record is associated with microbial mats in fine-grained intervals between turbidite-like deposits documented above the storm wave base. In this work, a detailed sedimentological study permitted to divide the unit into two informal sections separated by an erosional unconformity attributed to a sea level fall. Prograding turbidite-like sand-grained deposits document gravitational turbulent flows. The tectonostratigraphic framework allowed to disregard a submarine fan, whereas the unconformity does not support an autocyclic deltaic evolution. Palaeocurrents and palaeoslope indicators to the SW suggest a high source area to the NE, that contrasts with the well-established E-W trend of most of the units of similar age in the Cuyania terrane. Deposition was probably deflected by subaqueous highs with the development of mini-basins most likely structurally controlled.The ichnological analysis allowed to recognise ichnotaxa and intergradation cases between ichnospecies previously unknown for the unit, adding to the most diverse record of Dictyodora and Nereites in central-western Gondwana. Chondrites intricatus, Chondrites targionii, Chondrites?, Dictyodora zimmermanni, and specimens resembling Dictyodora liebeana are recorded for the first time. Nereites missouriensis is documented in intergradation with Nereites jacksoni, whereas Dictyodora scotica is recognised in intergradation with Dictyodora atuelica and Dictyodora tenuis. A decrease in ichnodiversity throughout the section could be related to variation in environmental parameters between the lower and upper sections. The ichnofacies assignment is discussed in terms of ethological categories and ichnodiversity. Some of the defined ichnoassemblages are assignable to a non-standard record of the Nereites Ichnofacies, and one in particular to the Nereites Ichnosubfacies, while another shows some characteristics coincident with the Zoophycos Ichnofacies. A transitional case between ichnofacies for the ichnological content of the unit as a whole, as previously reported in other units, cannot be disregarded.

New Carboniferous ophiuroid from central coastal New South Wales

The ophiuroid Bronthaster retus gen. et sp. nov. is described from a single specimen preserved as external dorsal and ventral moulds, and is assigned to the Protasteridae. It occurs in the Namurian (Levipustula levis Zone) Yagon Siltstone of the Myall Lakes district of central coastal New South Wales. This rock unit forms part of the structurally designated Myall Block at the southeastern coastal termination of the Tamworth Belt. This new genus is the first stelleroid to be described from the Carboniferous in Australia. Most distinctive of the new genus is the arrangement of many small, subrhomboidal to subpentagonal, overlapping ossicles on the dorsal surface of the arms. The clearly exhibited tight relationship between successive ambulacrals in one column in ventral view suggests that some revision of previously inferred relationships within ambulacral columns in protasterids is required. The position of prominent toehooks on each ambulacral ossicle shows that they functioned to prevent lateral movement of the ambulacral column, operating in conjunction with the strong longitudinal musculature to allow arm flexure in the horizontal plane. This feature could potentially identify, within the Protasteridae, an evolutionary lineage that proceeded through the middle Palaeozoic and maintained the linear arrangement of the ambulacral columns using the toehooks, rather than fusion of the ambulacral ossicles across the arm axis. Peter A. Jell [amjell@bigpond.com] and Alex G. Cook, School of Earth Sciences, The University of Queensland, St Lucia 4072, Australia.

Continuity of the Western Qinling and Qaidam‐Qilian blocks: Evidence from Precambrian and Permian–Triassic strata around the Gonghe Basin area, NW China

Tectonic affinity between the Qaidam‐Qilian Block and the Western Qinling Block is an outstanding issue that has not been fully understood. Detrital zircon U–Pb dating and Hf isotope data for the Precambrian and Permian–Triassic strata around the Gonghe Basin area in both the Western Qinling and Qaidam‐Qilian blocks were studied. Detrital zircons from Precambrian strata range in age from Archean to Late Neoproterozoic, with age peaks at ~2.7–2.5, ~1.8–1.6, and ~0.95–0.75 Ga. Except for Precambrian age groups, Early Palaeozoic (550–500 Ma and ~450 Ma) and Late Palaeozoic–Triassic (~250 Ma) age peaks were also detected in the Triassic strata. The juvenile accreted crustal materials were detected at 1.85–1.5 Ga and over ~2.5 Ga based on zircon Hf isotope studies. Different Precambrian and Triassic strata in both the Western Qinling Block and the Qaidam‐Qilian Block show parallel age distribution patterns and Hf isotopes. A Qaidam‐Qilian source contributed to the majority of detritus in Precambrian strata. Besides sources from the Qaidam‐Qilian Block, Late Permian intrusions around the Gonghe Basin area also provided inputs for the Triassic strata. Detrital zircon studies show a strong connection of the Qaidam‐Qilian Block and the majority of detritus in the Gonghe Basin area of the Western Qinling Block. Tectonomagmatic and provenances comparison of the Western Qinling and Qaidam‐Qilian blocks are supportive of their tectonic continuity at least in three stages, that is, in the Precambrian, Middle Palaeozoic, and Permian to Early Triassic.

Llandovery (lower Silurian) graptolites from the Sepon Mine, Truong Son Terrane, central Laos and their palaeogeographical significance

Graptolites are described from the Llandovery of the Sepon mine area, central Laos, part of the Truong Son Terrane. The palaeobiogeographical affinities of the Rhuddanian graptolites are with peri-Gondwanan Europe and Arabia rather than equatorial regions such as South China and Laurentia (Arctic Canada). A review of previous palaeontological research on the middle Palaeozoic of the Indochina terranes reveals an often contradictory picture with vertebrates suggesting close proximity of South China to the Truong Son Terrane in the Devonian and invertebrates and radiolarians providing evidence for a palaeogeographical barrier between the two. Graptolites from the Mojiang area, Yunnan are typical of low latitude Silurian faunas and suggest that this area (Simao or Loei Terrane) was separate from the Truong Son Terrane.

New polybranchiaspiform fishes (Agnatha: Galeaspida) from the Middle Palaeozoic of China and their ecomorphological implications.

The 438–370-million-year-old galeaspids, diversified armoured jawless vertebrates (‘ostracoderms’) from China and northern Vietnam, were assumed to have a benthic feeding habit in a coastal, marine environment. Here, we describe two new genera of galeaspid fishes, Platylomaspis gen. nov. and Nanningaspis gen. nov. from the Middle Palaeozoic of China. The two new forms are characterized by a rostral process and strikingly broad ventral rim, and clustered with Gumuaspis to form a new family, Gumuaspidae, which represents the most primitive clade of Polybranchiaspiformes. They extend the earliest occurrence of Polybranchiaspiformes backward about 19 million years, and expand its geographical distribution from southern China and northern Vietnam to the Tarim Basin, northwestern China. The new taxa exhibit many morphological convergences with modern rays, and might specify a new kind of lifestyle of galeaspids, the half burrowing habit. Probably benefiting from the new lifestyle, the Gumuaspidae has become the longest lasting galeaspid family. The new findings demonstrated that the demersal galeaspids had developed three different kinds of lifestyles: semi-infaunal benthic (half buried), epibenthic, and suprabenthic (nektonic) habits to accommodate to differentiated ecological niches, and reached the peak of their diversity by the Pragian of the Early Devonian.

Tectonic and environmental controls on Palaeozoic fluvial environments: reassessing the impacts of early land plants on sedimentation

The apparent increase in occurrence of meandering fluvial channel systems in the Middle Palaeozoic has long been related to the effects of land-plant colonization. However, evidence for meandering channels in non-vegetated settings is shown by pre-vegetation successions on Earth, from the prevalence of meandering channels on Mars, from physical modelling of meandering channels, and from non-vegetated channels in modern desert basins. In addition, early land plants had small dimensions, were limited in their occurrence, and were dependent on environmental factors. Here, we question the capacity of early land plants to impose the major impacts suggested by current models. We propose that the sudden widespread occurrence on Earth of fluvial deposits indicative of the accumulation of meandering river systems in the Middle Palaeozoic was primarily an effect of environmental and tectonic conditions that prevailed during this period. These conditions induced a worldwide increase in the proportion of meandering rivers, which in turn helped favour the appropriate environment for land-plant colonization of the continents. We propose that land plants opportunistically took advantage of an appropriate global environment, which enabled them to thrive in continental environments. Fluvial environments characterized by single-channel systems and stable floodplains facilitated the greening of the land.

Assessing the fidelity of marine vertebrate microfossil δ18O signatures and their potential for palaeo-ecological and -climatic reconstructions

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.

Seven 365-Million-Year-Old Trilobites Moulting within a Nautiloid Conch.

A nautiloid conch containing many disarticulated exoskeletons of Omegops cornelius (Phacopidae, Trilobita) was found in the Upper Devonian Hongguleleng Formation of the northwestern margin of the Junggar Basin, NW China. The similar number of cephala, thoraces and pygidia, unbroken thoraces, explicit exuviae, and lack of other macrofossils in the conch, indicate that at least seven individual trilobites had moulted within the nautiloid living chamber, using the vacant chamber of a dead nautiloid as a communal place for ecdysis. This exuvial strategy manifests cryptic behaviour of trilobites, which may have resulted from the adaptive evolution induced by powerful predation pressure, unstable marine environments, and competition pressure of organisms occupying the same ecological niche in the Devonian period. The unusual presence of several trilobites moulting within a nautiloid conch is possibly associated with social behaviours in face of a serious crisis. New materials in this study open a window for understanding the survival strategy of marine benthic organisms, especially predator-prey interactions and the behavioural ecology of trilobites in the middle Palaeozoic.

Mitogenomics of Vetigastropoda: insights into the evolution of pallial symmetry

The nucleotide sequences of the complete or nearly complete mitochondrial (mt) genomes of seven vetigastropods were determined:Angaria neglecta(Angarioidea),Phasianella solida(Phasianelloidea),Granata lyrata(Seguenzioidea),Tegula lividomaculataandBolma rugosa(Trochoidea),Diodora graeca(Fissurelloidea) andLepetodrilus schrolli(Lepetodriloidea). While the mt genomes of the superfamilies Angarioidea, Phasianelloidea, Seguenzioidea and Trochoidea conform generally to the ancestral gene order of Vetigastropoda and Gastropoda, those of the superfamilies Fissurelloidea and Lepetodriloidea have suffered important rearrangements. The gene order of the mtDNAofChrysomallon squamiferum, a representative of Neomphalina, was also analysed since it has been proposed to be closely related to Vetigastropoda, and showed a distinct arrangement. The reconstructed phylogenies recovered Neomphalina as a distinct gastropod lineage that is the sister group (only with moderate bootstrap support) of a clade including Vetigastropoda and Neritimorpha + Caeno‐gastropoda while the relative position of Heterobranchia and Patellogastropoda in the gastropod tree could not be determined definitively due to their long branches. Within the monophyletic Vetigastropoda, the superfamily Fissurelloidea was recovered as the sister group of two lineages, one including Lepetodriloidea as the sister group of Seguenzioidea + Halitoidea, the other including Phasianelloidea, Angarioidea and Trochoidea without resolved relationships. The long branches of Fissurelloidea were found to introduce significant tree instability in phylogenetic reconstruction. The new phylogeny supports that the loss of the right pallial gill occurred multiple times in vetigastropod evolution as previously suggested and that Phasianelloidea, Angarioidea and Trochoidea radiated from a common asymmetric (single‐gilled) ancestor that lived in the middle Palaeozoic.

An exceptionally preserved Late Devonian actinopterygian provides a new model for primitive cranial anatomy in ray-finned fishes.

Actinopterygians (ray-finned fishes) are the most diverse living osteichthyan (bony vertebrate) group, with a rich fossil record. However, details of their earliest history during the middle Palaeozoic (Devonian) ‘Age of Fishes' remains sketchy. This stems from an uneven understanding of anatomy in early actinopterygians, with a few well-known species dominating perceptions of primitive conditions. Here we present an exceptionally preserved ray-finned fish from the Late Devonian (Middle Frasnian, ca 373 Ma) of Pas-de-Calais, northern France. This new genus is represented by a single, three-dimensionally preserved skull. CT scanning reveals the presence of an almost complete braincase along with near-fully articulated mandibular, hyoid and gill arches. The neurocranium differs from the coeval Mimipiscis in displaying a short aortic canal with a distinct posterior notch, long grooves for the lateral dorsal aortae, large vestibular fontanelles and a broad postorbital process. Identification of similar but previously unrecognized features in other Devonian actinopterygians suggests that aspects of braincase anatomy in Mimipiscis are apomorphic, questioning its ubiquity as stand-in for generalized actinopterygian conditions. However, the gill skeleton of the new form broadly corresponds to that of Mimipiscis, and adds to an emerging picture of primitive branchial architecture in crown gnathostomes. The new genus is recovered in a polytomy with Mimiidae and a subset of Devonian and stratigraphically younger actinopterygians, with no support found for a monophyletic grouping of Moythomasia with Mimiidae.

Detrital zircon geochronology of the Cretaceous succession from the Iberian Atlantic Margin: palaeogeographic implications

Detrital zircon U–Pb data performed on eight Cretaceous sandstone samples (819 age isotopic results) from the Lusitanian basin (west Portugal) constrain the history of uplift and palaeodrainage of western Iberia following break-up of Pangaea and opening of the North Atlantic Ocean. We examined the links between shifts in provenance and known basinwide unconformities dated to the late Berriasian, Barremian, late Aptian and Cenomanian–Turonian. The detrital zircon record of sedimentary rocks with wider supplying areas is relatively homogenous, being characterized by a clear predominance of late Palaeozoic ages (c. 375–275 Ma) together with variable proportions of ages in the range c. 800–460 Ma. These two groups of ages are diagnostic of sources within the Variscan Iberian Massif. A few samples also reveal significant amounts of middle Palaeozoic (c. 420–385 Ma) and late Mesoproterozoic to early Neoproterozoic (c. 1.2–0.9 Ga) zircon, which are almost absent in the basement to the east of the Lusitanian basin, but are common in terranes with a Laurussia affinity found in NW Iberia and the conjugate margin (Newfoundland). The Barremian unconformity marks a sudden rise in the proportion of c. 375–275 Ma zircon ages accompanied by a decrease in the abundance of the c. 420–385 Ma and c. 1.2–0.9 Ga ages. This shift in the zircon signature, which is contemporaneous with the separation of the Galicia Bank from Flemish Cap, reflects increased denudation of Variscan crystalline rocks and a reduction in source material from NW Iberia and adjoining areas. The late Aptian unconformity, which represents the largest hiatus in the sedimentary record, is reflected by a shift in late Palaeozoic peak ages from c. 330–310 Ma (widespread in Iberia) to c. 310–290 Ma (more frequent in N Iberia). It is considered that this shift in the age spectra resulted from a westward migration of catchment areas following major uplift in northern Iberia and some transport southward from the Bay of Biscay under the influence of a well-established Atlantic circulation.

Juxtaposition of Barrovian and migmatite domains in the Chinese Altai: a result of crustal thickening followed by doming of partially molten lower crust

AbstractOrdovician metasedimentary rocks are the oldest and most extensive sedimentary sequence in the Chinese Altai. They experienced two major episodes of deformation (D1 and D2) resulting in the formation of juxtaposed Barrovian‐type and migmatite domains. D1 is characterized by a penetrative sub‐horizontal fabric (S1), and D2 is marked by upright folds (F2) withNW–SE‐trending axial planes in shallow crustal levels and by sub‐vertical transposition foliations (S2) in the high‐grade cores of large‐scale F2 antiforms. In the Barrovian‐type domain, successive growth of biotite, garnet and staurolite is observed in the S1 fabric. Kyanite included in garnet and plagioclase in the migmatite domain is interpreted to have formed also in S1. In the biotite and garnet zones, the spaced S2 cleavage is marked by biotite and muscovite, and in the staurolite and kyanite zones, the penetrative S2 fabric is characterized by sillimanite, locally with late cordierite. Phase equilibria modelling indicates that the S1 fabric was associated with an increase in pressure and temperature under Barrovian‐type conditions in both domains. The S2 fabric was related to decompression, in which rocks in the biotite and garnet zones well preserve the peak assemblage, and the higher grade rocks in the staurolite and kyanite zones re‐equilibrated to different degrees under high‐temperature/low‐pressure (HT/LP) conditions. The D1 metamorphic history is attributed to the progressive burial related to Early–Middle Palaeozoic crustal thickening and the metamorphism associated with D2 is interpreted to result from exhumation by vertical extrusion. The extrusion of hot rocks was contemporaneous with the formation of gneiss domes accompanied by the intrusion of juvenile magmas at middle crustal levels during the Middle Palaeozoic. Consequently, there is a genetic link between the Barrovian‐type and migmatite domains related to continuous transition of the Barrovian‐type fabric into theHT/LPone during development of domal structures in the southern Altai orogenic belt. This study has a broad impact on the understanding of the thermo‐mechanical behaviour of accretionary orogenic systems worldwide. The lower crustal flow and doming of hot crust, so far reported only in continental collisional settings, seems to be also an integral mechanism responsible for both horizontal and vertical redistribution of accreted material prior to continental collision.

Early vertebrate evolution

AbstractDebate over the origin and evolution of vertebrates has occupied biologists and palaeontologists alike for centuries. This debate has been refined by molecular phylogenetics, which has resolved the place of vertebrates among their invertebrate chordate relatives, and that of chordates among their deuterostome relatives. The origin of vertebrates is characterized by wide‐ranging genomic, embryologic and phenotypic evolutionary change. Analyses based on living lineages suggest dramatic shifts in the tempo of evolutionary change at the origin of vertebrates and gnathostomes, coincident with whole‐genome duplication events. However, the enriched perspective provided by the fossil record demonstrates that these apparent bursts of anatomical evolution and taxic richness are an artefact of the extinction of phylogenetic intermediates whose fossil remains evidence the gradual assembly of crown gnathostome characters in particular. A more refined understanding of the timing, tempo and mode of early vertebrate evolution rests with: (1) better genome assemblies for living cyclostomes; (2) a better understanding of the anatomical characteristics of key fossil groups, especially the anaspids, thelodonts, galeaspids and pituriaspids; (3) tests of the monophyly of traditional groups; and (4) the application of divergence time methods that integrate not just molecular data from living species, but also morphological data and extinct species. The resulting framework will provide for rigorous tests of rates of character evolution and diversification, and of hypotheses of long‐term trends in ecological evolution that themselves suffer for lack of quantitative functional tests. The fossil record has been silent on the nature of the transition from jawless vertebrates to the jawed vertebrates that have dominated communities since the middlePalaeozoic. Elucidation of this most formative of episodes likely rests with the overhaul of early vertebrate systematics that we propose, but perhaps more fundamentally with fossil grades that await discovery.

Mississippian crinoid biodiversity, biogeography and macroevolution

Abstract: The biodiversity and biogeography of 217 genera of Mississippian crinoids from North America and the British Isles shed light on the macroevolutionary turnover between the Middle Palaeozoic and Late Palaeozoic Crinoid Evolutionary Faunas. This turnover resulted from steady differential extinction among clades during the middle Mississippian after crinoids reached their Phanerozoic peak of generic richness during the early Mississippian. This peak richness was primarily a function of Mississippian originations rather than Devonian–holdover taxa. North America had 100 per cent higher generic richness than the British Isles, but rarefaction analysis adjusts the difference to only 37 per cent higher. Rarefaction demonstrated that North America had increased biodiversity, compared to the British Isles, almost entirely among monobathrid camerates, disparids and primitive cladids. In contrast, diplobathrid camerates, advanced cladids and flexibles had the same generic biodiversity between regions, when compared using rarefaction. The early Mississippian radiation resulted from two primary causes: (1) the expansion of Tournaisian carbonate ramps following the Frasnian mass extinction of reef faunas and (2) the predatory release in the Tournaisian following the end‐Famennian Hangenberg extinction of durophagous fishes. A majority of crinoid genera from the British Isles are cosmopolitan. When combined with rarefaction analysis and evidence for more first occurrences in North America, this suggests higher origination rates in North America, especially when carbonate ramps were widespread. With the gradual reduction in the area of carbonate ramps from the early to late Mississippian, in conjunction with the radiation of new durophagous fishes, camerate crinoids in particular experienced continuous background extinction, without replacement, beginning during the earliest Viséan (late Osagean). By middle Viséan time (late Meramecian) advanced cladids were dominant in all settings. This resulted in the transition from the Middle Palaeozoic to the Late Palaeozoic Crinoid Macroevolutionary Fauna.

A brief review of the Middle Palaeozoic vertebrates from Southeast Asia

Here we review the Silurian and Devonian vertebrates from Southeast Asia and their biogeographic affinities. These vertebrate remains are distributed in the Shan-Thai and Indochina terranes as well as the South China block, including the fossil fishes from western Yunnan (a part of the Shan-Thai terrane). The Devonian vertebrates from the Si Ka, Bac Bun and Mia Le formations of northern Vietnam resemble those from the Xitun Formation of Yunnan, South China, and thus belong to the Xitun fauna. The Late Silurian vertebrates from the Dia Giang Formation of My Duc (central Vietnam) are suggestive of the Xiaoxiang fauna of South China. Fossil vertebrate evidence shows the proximity between the South China block and the Indochina terrane in the Middle Palaeozoic and the close relationship between Shan-Thai and east Gondwana in the Middle Devonian.