Since the 1960s, huge progress has been made in reconstructing the multielement skeletons of conodont species and developing a biologically defensible taxonomy. Nevertheless, a widespread prejudice remains that certain parts of the conodont skeleton, particularly the P1 elements, are more informative than others with regard to taxonomy and evolutionary relationships. Here, we test these views. A new partial multielement reconstruction of the skeleton of the biostratigraphically significant conodont originally described as Gnathodus commutatus homopunctatus allows us to conduct a cladistic test of the alternative hypotheses of phylogenetic placement of this species. Our analysis also provides the first test of the hypothesis that Lochriea– species of which are markers for global correlation – is monophyletic and tests hypotheses concerning the origins of the genus. Our results demonstrate that homopunctatus is a species of Lochriea and that the genus is monophyletic. The widely held view that Lochriea arose from a species of Bispathodus is not supported. Our results show that it is difficult to predict a priori which parts of the conodont skeleton carry phylogenetic signal, and provide strong support for the hypothesis that similarity in the morphology of conodont P1 elements alone is not a reliable guide to relationships and taxonomic groupings of conodont species. This is because P1 elements with similar morphologies are convergently acquired in multiple conodont clades, because reliance on the characters of only one of the six or seven morphologically distinct elements of the conodont skeleton ignores phylogenetically significant data and because P1 elements can lack characters that might seem to be diagnostic of a genus. Conodonts are no different to other organisms: ignoring data that have the potential to be phylogenetically informative is unlikely to produce the most reliable hypotheses of evolutionary relationships. We suggest that other biostratigraphically significant hypotheses of relationship between conodont taxa that are based on P1 elements alone should be subject to cladistic testing.

The Aladag Unit is one of the main tectonic units in the Tauride Belt, located in southern Turkey. It includes a continuous Paleozoic carbonate sequence encompassing the mid-Carboniferous boundary, with outcrops being especially well exposed in the Hadim region. The boundary succession lithology is mainly composed of carbonates with intercalated quartz arenitic sandstone layers. Based on foraminifers, four biostratigraphic zones have been defined in the interval from the Upper Serpukhovian to the Lower Bashkirian. These zones are, in ascending order: the Eostaffella ex gr. ikensis-E. postmosquensis Zone (Zapaltyubinsky Horizon, Upper Serpukhovian); the Plectostaffella jakhensis-P. bogdanovkensis Zone, and the Millerella marblensis Zone (Bogdanovsky Horizon, lower Bashkirian); and the Semistaffella sp. Zone (Syuransky Horizon, lower Bashkirian). The mid-Carboniferous boundary occurs between the Eostaffella ex gr. ikensis-E. postmosquensis Zone and the Plectostaffella jakhensis-P. bogdanovkensis Zone. Boundary beds are characterized by eight, repeatedly occurring microfacies types, namely: (1) coated crinoidal packstone; (2) coated bioclastic grainstone; (3) oolitic grainstone; (4) oolitic packstone-grainstone; (5) intraclastic grainstone; (6) mudstone-wackestone; (7) quartz-peloidal packstone; and 8) quartz arenitic sandstone. Based on microfacies stacking patterns, various types of shallowing-upward cycles have been recognized. Depositional sequences and sequence boundaries are correlatable with those described from North America and Russia and Carboniferous global sea-level curves. The duration of cycles has been estimated as 100 ky, suggesting that cycle periodicities correspond to the Milankovitch eccentricity band.

Conodont elements, microfossil remains of extinct primitive vertebrates, are commonly exploited as mineral archives of ocean chemistry, yielding fundamental insights into the palaeotemperature and chemical composition of past oceans. Geochemical assays have been traditionally focused on the so-called lamellar and white matter crown tissues; however, the porosity and crystallographic nature of the white matter and its inferred permeability are disputed, raising concerns over its suitability as a geochemical archive. Here, we constrain the characteristics of this tissue and address conflicting interpretations using ptychographic X-ray-computed tomography (PXCT), pore network analysis, synchrotron radiation X-ray tomographic microscopy (srXTM) and electron back-scatter diffraction (EBSD). PXCT and pore network analyses based on these data reveal that while white matter is extremely porous, the pores are unconnected, rendering this tissue closed to postmortem fluid percolation. EBSD analyses demonstrate that white matter is crystalline and comprised of a single crystal typically tens of micrometres in dimensions. Combined with evidence that conodont elements grow episodically, these data suggest that white matter, which comprises the denticles of conodont elements, grows syntactically, indicating that individual crystals are time heterogeneous. Together these data provide support for the interpretation of conodont white matter as a closed geochemical system and, therefore, its utility of the conodont fossil record as a historical archive of Palaeozoic and Early Mesozoic ocean chemistry.