John A. Talent

Middle Palaeozoic extinction events: Faunal and isotopic data

Abstract At least 9 and possibly as many as 12 extinction events of global or near-global impact can be discriminated in the mid-Palaeozoic (earliest Silurian to Early Carboniferous) on the basis of brachiopod, coral, conodont and ammonoid data, and the history of carbonate build-ups. Isotopic data from whole-rock samples are presented for three of these events, based on Australian carbonate sequences constrained by conodont data. These data represent the initial phase of a more extensive investigation using C, O, S, Sr and Nd isotopic signatures derived from conodonts, articulate and inarticulate brachiopod shell, and fish remains from numerous Australian and European sequences. The aim of the project is to identify isotopic responses to extinction events, and address causes for these changes. In limestone sections analysed so far, variations in carbon isotope compositions on a whole-rock scale are most marked at horizons that can be correlated with times of significant reduction in biomass and diversity. This is despite the fact that the whole rocks are in fact multicomponent systems with respect to carbon, in part arising from diagenetic rearrangements of carbon distribution within the scale of the whole rock. Thus, the carbon isotope data do provide evidence on a whole-rock scale for fundamental changes in the global carbon cycle that are correlative with extinction events. It is, however, unlikely that the magnitude of the isotopic shift will be precisely documented from such whole-rock analyses. The oxygen isotope results are less obviously related to the times of carbon isotope excursion; this contrast in apparent resilience to post-depositional modification of pristine isotope compositions exemplifies the extreme caution needed in evaluating isotopic data from carbonates.

The plant Leclercqia (Lycopsida) in Gondwana: implications for reconstructing Middle Devonian palaeogeography

Abundant and well-preserved material of the ligulate lycopsid genus Leclercqia is reported from a new Middle Devonian locality in northeastern Queensland (Australia). The plants occur in a chert horizon in the Storm Hill Sandstone of the Dosey-Craigie Platform. Lithological data and conodont analyses combined with information from in situ spores provide an age for the plant levels ranging from Eifelian, possibly Middle Eifelian, to Early Givetian. Plant taxonomic identification is based on vegetative and fertile stems that display both external morphology and anatomy. This material represents the best documented occurrence of Leclercqia outside Laurussia and possibly the earliest in Gondwana; it provides evidence that colonization of Gondwana by the species L. complexa was contemporaneous to that of Siberia and Kazakhstan. Analysis of the distribution patterns of L. complexa suggests that it was adapted to a wide range of environments, but within certain limits which we hypothesize to be those of a climatic belt. Such considerations support previous studies using other biological data, such as faunas and palynomorphs, for reconstructing Devonian palaeogeography. They favour a close proximity of Laurussia and Gondwana rather than the occurrence of a wide ocean separating the two palaeocontinents in Middle Devonian times.

Oldest coelacanth, from the Early Devonian of Australia

Coelacanths are well-known sarcopterygian (lobe-finned) fishes, which together with lungfishes are the closest extant relatives of land vertebrates (tetrapods). Coelacanths have both living representatives and a rich fossil record, but lack fossils older than the late Middle Devonian (385–390 Myr ago), conflicting with current phylogenies implying coelacanths diverged from other sarcopterygians in the earliest Devonian (410–415 Myr ago). Here, we report the discovery of a new coelacanth from the Early Devonian of Australia (407–409 Myr ago), which fills in the approximately 20 Myr ‘ghost range’ between previous coelacanth records and the predicted origin of the group. This taxon is based on a single lower jaw bone, the dentary, which is deep and short in form and possesses a dentary sensory pore, otherwise seen in Carboniferous and younger taxa.

LITHOSTRATIGRAPHY, CONODONT BIOSTRATIGRAPHY AND DEPOSITIONAL ENVIRONMENT OF THE MIDDLE DEVONIAN (GIVETIAN) TO EARLY CARBONIFEROUS (TOURNAISIAN) LIPAK FORMATION IN THE PIN VALLEY OF SPITI (NW INDIA)

Bed-by-bed lithostratigraphic sections combined with sequence stratigraphy and conodont biostratigraphy provide new information on the depositional environment and age of the Lipak Formation in the Pin Valley (Spiti). The formation comprises mixed siliciclastic and calcareous sediments at lower levels, richly fossiliferous limestones with two distinct sandstone incursions at higher levels, and dark mudstones followed by a thin siltstone interval. The upper limit of the Lipak Formation is defined by the angular unconformity below the sandstones of the Permian Gechang Formation. Lithologic correlation with sections in upper Lahaul indicates that, in the Pin Valley, the formation has been truncated just below its characteristic gypsum horizon. The lower boundary of the Lipak Formation is gradational from coastal arenites of the Muth Formation; the mappable boundary is drawn at the first appearance of dark carbonaceous, argillaceous siltstone and shale. Sedimentary structures, microfacies and conodont faunas indicate a general shallow marine depositional environment of the Lipak Formation in the Pin Valley; five sequence stratigraphic units have been distinguished. Conodont data demonstrate that the lowest 33 m of the Lipak Formation of the Pin Valley is mid to late Early varcus Subzone with characteristic species of Icriodus and Bipennatus . A previously unrecognised hiatus at c. 33 m above the base, at the boundary of sequence stratigraphic units S1 and S2, represents the interval Middle varcus Subzone to at least the end of the late Famennian Early expansa Zone. Because this hiatus does not correspond to a mappable boundary, no division of the Lipak Formation into named stratigraphic units is suggested, but we refer informally to the sediments represented by cycle S1 as Lipak A, and the sediments represented by cycles S2-S5 as Lipak B. Determination of S1 as Early varcus Subzone provides a maximum age for the gradationally underlying Muth Formation. At 75 m above the base of the composite Lipak Formation section, a 58 cm black to dark grey shale interval within late Famennian fossiliferous limestones conceivably correlates with the Hangenberg Event (end-Middle praesulcata Zone). Younger conodont faunas of the Lipak Formation -dominated by species of Clydagnathus with species of Bispathodus and Pseudopolygnathus also represented- is shown to extend to the mid-Tournaisian Early crenulata Zone.

Low-grade metamorphism in the Palaeozoic sequences of the Townsville hinterland, northeastern Australia*

The transition from diagenesis to low‐grade metamorphism for the Ordovician ‐ Early Carboniferous sequences of the Burdekin and Clarke River Basins and the Broken River ‐ Camel Creek region is characterised by b ‐cell dimension and Kübler Index (KI = illite crystallinity) of K‐white micas, clay‐mineral assemblages and conodont colour alteration index (CAI). Data indicate that: (i) there is a regional metamorphic pattern characterised by anchizonal conditions, with consistently higher grade in the Camel Creek region (or Subprovince); (ii) anomalies are explicable in terms of location relative to areas of differing intensity of deformation (e.g. the Ruxton Formation) and inferred greater depth of burial (e.g. southern versus northern outcrop tracts of Quinton Formation); (iii) the regional pattern in the Broken River ‐ Gray Creek area shows general alignment with the regional grain; (iv) there is a generalised north to south increase in CAI for the Broken River region (‘Top Hut’ to Broken River, 6), with higher values towards the Montgomery Range intrusions; (v) in the Burdekin Basin, there arehigher CAI values north of Mt Podge and ‘Star’, and lower values (

EARLIEST TRIASSIC CONODONTS FROM CHITRAL, NORTHERNMOST PAKISTAN

Extensive tracts of very shallow water carbonates in the valleys of the Yarkhun and Mastuj rivers of Chitral (northernmost Pakistan) previously though to be Permian (or Cretaceous) are shown by conodonts from two horizons in sequences 110 km apart—near Torman Gol (Mastuj valley) and near Sakirmul (upper Yarkhun valley)—to include earliest Triassic (Scythian—Induan) horizons. Both faunas have Isarcicella staeschei Dai & Zhang , Is. lobata Perri , Is. turgida (Kozur et al.) and Hindeodus parvus (Kozur & Pjatakova), whereas Is. Isarcica (Huckriede) has been recognised only in the Torman Gol occurrence. The presence, respectively, of Is. staeschei in the Sakirmul and Is. isarcica in the Torman Gol occurrences, allows discrimination of the staeschei and isarcica zones respectively the third and the fourth conodont biozones of the Early Triassic conodont biozonation of Perri (in Perri & Farabegoli 2003). Such faunas, consisting mainly of isarcicellids and hindeodids but lacking gondolellids, are characteristic of restricted sea environments across the Permian–Triassic boundary and in the earliest Triassic in other Tethyan areas. The conodont faunas from these two occurrences are remarkably similar, nearly contemporaneous, and indicate shallow water biofacies. They are inferred to equate with the Ailak Dolomite, a sequence of Late Permian–?Late Triassic dolostones discriminated farther up the Yarkhun valley and extending eastwards into the upper Hunza region of northernmost Pakistan. The Zait Limestone and Sakirmul carbonate sequence are consistent with extension of the previously inferred Triassic carbonate platform at least 110 km farther to the SW than previously supposed.

Late Ludfordian Correlations and the Lau Event

Changes in whole conodont faunas and δ13C values are combined to achieve high-resolution correlations of Upper Silurian successions in many areas (primarily Gotland, Skane, Lithuania, Bohemia, Austria, Sardinia and Queensland); other areas are correlated with lower precision. Four of the widely recognised subzones average considerably less than 0.1 Ma and a fifth interval less than 0.01 Ma. The main constraints on resolution and precision now achievable are the amount of, and the precision in, new or previously published data from each local section; centimetric scale collecting would be worthwhile in many sections. Some stratal characters are more widespread during certain intervals than might have been expected, for example, the presence of oncolites and algal coatings in the Icriodontid Zone and the lower part of the O. snajdri Zone. Similarly, the Dayia navicula bloom in the Upper P. siluricus Subzone was widespread. So also were muddy-sandy sediments followed by oolite low in the O. snajdri Zone; crinoids flourished widely when the lower part of the Lower Icriodontid Subzone was formed. Closely spaced samples show that, in some intervals, the now well-known δ13C spike was modified considerably by fluctuations; that is, it is not a smooth plateau. The best-documented fluctuations (19 analyses) are 2.5 smooth cycles with up to 2.7‰ in amplitude during ca 12,000 years or less in the base of the O. snajdri Zone. An enigma is the depletion of the spike in some sections, especially in the best Bohemian section.

NEW ONYCHODONTIFORM (OSTEICHTHYES; SARCOPTERYGII) FROM THE LOWER DEVONIAN OF VICTORIA, AUSTRALIA

Abstract The Onychodontiformes is a poorly known sarcopterygian fish group, with four genera currently described, predominantly from the Middle-Late Devonian. A new onychodont, Bukkanodus jesseni n. gen. and sp., from the Fairy Formation, Victoria, Australia, is of Early Devonian (mid-late Pragian) age, representing one of the oldest known occurrences of this group. Other Pragian onychodonts are represented by a single lower jaw from China, while older occurrences (Lochkovian) include a lower jaw also from China and isolated teeth from Nevada. The Australian material, though disarticulated, includes skull, jaw, and palatal and dental specimens. These specimens share characteristics with younger onychodont taxa, including a laterally compressed tooth whorl with main and accessory rows containing an equal number of teeth, an anteriorly arched shape of the premaxilla, a lateral rostral bone participating in the orbital margin, parietals separated in the midline by small bones, the insertion of the basisphenoid region of the braincase into a posterior area on the parasphenoid, and an ethmoidal sensory canal running along the dorsal margin of the premaxilla rather than through the bone. Bukkanodus jesseni differs from other onychodonts in the presence of a distinct vomer, coronoids with fang pairs, a restricted herringbone pattern of ribs on enamel ridges of teeth, and the presence of a cluster of large pores on certain skull and jaw bones. The course of the ethmoid sensory canal and this cluster of pores are plesiomorphic features for the Onychodontiformes, also occurring in primitive sarcopterygian taxa such as Youngolepis, Powichthys, and Kenichthys. The opening of this cluster on the internal surface of the premaxilla resembles the rostral organ of derived coelacanths.

LATE DEVONIAN-TOURNAISIAN CONODONTS FROM THE EASTERN KHYBER REGION, NORTH-WEST PAKISTAN

Conodonts (62 species and subspecies) from acid-leaching of 226 samples from four sections through the Ali Masjid Formation west of Misri Khel in the former South Khyber Agency, north-west Pakistan, are documented by illustrations and distributional data. These indicate that most of this unit, at least in that area, spans the interval Middle crepida Zone (low in the Famennian) to at least Early crenulata Zone (mid-Tournaisian), though a fauna from low in one of the sections produced conodonts indicative of the Late falsiovalis Zone (early Frasnian). Two major hiatuses are inferred: between the Late falsiovalis and Middle crepida zones, and between the Late expansa and Early duplicata zones. Coherence of the conodont biostratigraphy accords with lithologic alignments between the sections.

Devonian trilobites from the Shogrām Formation, Kurāgh, Chitral (NW Pakistan)

Late Devonian trilobites from horizons close to the Frasnian-Famennian boundary in the Shogrām Formation at Kurāgh, Chitral (NW Pakistan) are described. A new species of Asteropyginae, Neocalmonia chitralensis sp. nov., and a new subpecies, Neocalmonia batillifera orientalis subsp. nov., are described; these extend the range of Asteropyginae eastwards from Iran and southern Afghanistan. The Upper Kellwasser Event is located within KUR 19 of Talent et al. (1999).