Paleobiology

The Devonian-Carboniferous boundary is allegedly marked by one of the most catastrophic global extinctions associated with sedimentation of the Hangenberg black shale. A dense sampling of the Kowala section in the Holy Cross Mountains, Poland, challenges this view, showing that the faunal dynamics across the Hangenberg black shalewas not more dramatic than that across the preceding Kowala black shale. Quantitative analysis and biologically meaningful conodont apparatus study of the Kowala material offer probably the most complete record of faunal change in the latest Famennian and earliest Tournaisian among those sampled bed-by-bed for ammonoids and conodonts. It appears that the faunal dynamics of both cephalopods and conodonts was controlled by environmental changes that resulted in numerous immigrations and disappearances of particular lineages. Only a small fraction of lineages persisted long enough at the place, and transformed their morphology fast enough, to leave a record of their evolution. Most of the evolution apparently took place elsewhere. Locations of remote refugia where these lineages evolved in the time span bracketed by the Kowala and Hangenberg black shale events remain to be identified. Conodont apparatus study on geographically distant Vietnamese locality Cat Ba provides evidence that at least in the latest Famennian some conodont species unknown from Poland were present in Vietnam. Moreover, the contribution of species known from both localities to Polish and Vietnamese fossil assemblages was dramatically different.

<p> Conodonts were the first vertebrates to develop mineralized dental tools, known as elements. Recent research suggests that conodonts were macrophagous predators and/or scavengers but we do not know how this feeding habit emerged in the earliest coniform conodonts, since most studies focus on the derived, ‘complex’ conodonts. Previous modelling of element position and mechanical properties indicate they were capable of food processing. A direct test would be provided through evidence of <italic>in vivo</italic> element crown tissue damage or through <italic>in vivo</italic> incorporated chemical proxies for a shift in their trophic position during ontogeny. Here we focus on coniform elements from two conodont taxa, the phylogenetically primitive <italic>Proconodontus muelleri</italic> Miller, 1969 from the late Cambrian and the more derived <italic>Panderodus equicostatus</italic> Rhodes, 1954 from the Silurian. Proposing that this extremely small sample is, however, representative for these taxa, we aim to describe in detail the growth of an element from each of these taxa in order to the test the following hypotheses: (1) <italic>Panderodus</italic> and <italic>Proconodontus</italic> processed hard food, which led to damage of their elements consistent with prey capture function; and (2) both genera shifted towards higher trophic levels during ontogeny. We employed backscatter electron (BSE) imaging, energy-dispersive X-ray spectroscopy (EDX) and synchrotron radiation X-ray tomographic microscopy (SRXTM) to identify growth increments, wear and damage surfaces, and the Sr/Ca ratio in bioapatite as a proxy for the trophic position. Using these data, we can identify whether they exhibit determinate or indeterminate growth and whether both species followed linear or allometric growth dynamics. Growth increments (27 in <italic>Pa. equicostatus</italic> and 58 in <italic>Pr. muelleri</italic> ) were formed in bundles of 4–7 increments in <italic>Pa. equicostatus</italic> and 7–9 in <italic>Pr. muelleri</italic> . We interpret the bundles as analogous to Retzius periodicity in vertebrate teeth. Based on applied optimal resource allocation models, internal periodicity might explain indeterminate growth in both species. They also allow us to interpret the almost linear growth of both individuals as an indicator that there was no size-dependent increase in mortality in the ecosystems where they lived <italic>e.g</italic> ., as would be the case in the presence of larger predators. Our findings show that periodic growth was present in early conodonts and preceded tissue repair in response to wear and damage. We found no microwear and the Sr/Ca ratio, and therefore the trophic position, did not change substantially during the lifetimes of either individual. Trophic ecology of coniform conodonts differed from the predatory and/or scavenger lifestyle documented for “complex” conodonts. We propose that conodonts adapted their life histories to top-down controlled ecosystems during the Nekton Revolution. </p>

A fossil larva lacking segmentation of the calcified carapace, closely resembling the trilobite protaspis, has been found associated with other skeletal elements of an angarocaridid Girardevia species in the mid Darriwilian of central Siberia. The presence of protaspis larvae in the angarocaridids, generally believed to represent a branch of the Aglaspidida, supports their proximity to trilobites and proves a low position on the arthropod phylogenetic tree but does not necessarily contradict the chelicerate affinity. The cephalic appendages of angarocaridids bore massive gnathobases with detachable spines, closely similar to those known in extant xiphosurans and in their probable Cambrian relatives. The stratigraphic succession of the angarocaridids, their phosphatized cuticle pieces being abundant in the Ordovician strata of Siberia, shows a gradual improvement of mechanical resistance of their carapaces, eventually resulting in a honeycomb structure. The associated benthic mollusc assemblage is dominated with the bellerophontids showing high mortality at metamorphosis and only the limpet-like Pterotheca, infaunal bivalves, and scaphopods being able to survive this in a substantial number. This suggests a strong selective pressure from predators equipped with well-skeletonised oral apparatuses able to crush mineralized body covers of their prey. Possibly, these were some of the associated conodonts of appropriate size and co-evolving towards their ability to crush more and more resistant cuticle. Less likely candidates for durophagy are endoceratid or orthoceratid cephalopods. Also the angarocaridids themselves, equipped with robust gnathobases of cephalic appendages, apparently predated on benthic shelly animals.

ABSTRACTBicuspid, tricuspid and tetracuspid postcanine teeth of a new non-mammaliaform eucynodont, Polonodon woznikiensis gen. et sp. nov. from the mid-late Carnian (early Late Triassic) of Woźniki clay-pit, Silesia (southern Poland) show incipient root division. They are similar to teeth of Dromatheriidae from the Carnian (early Late Triassic) to the Rhaetian (late Late Triassic) of Europe, India, and USA and the dentition of brasilodontids from the early Norian (mid Late Triassic) of Brazil. The P. woznikiensis teeth differ from those of the latter group mostly in the absence of cingulum. Some of the new fossils from Silesia provide the oldest Laurasian record of eucynodont teeth with the main cusp (a) anterior edge very long as the mesial cusp b is placed much lower than cusp c (distal). The contemporaneous Alemoatherium huebneri, from Gondwana, had similar postcanines. The findings from Poland indicate that this postcanine morphology was present in non-mammaliaform cynodonts from both hemispheres as early as the mid-late Carnian. The distal end of the humerus from the same locality is also described.

Abstract It is widely accepted that ornithodirans (bird lineage) and some pseudosuchians (crocodilian lineage) achieved fully erect limb posture in different ways. Ornithodirans have buttress-erected hindlimbs, while some advanced pseudosuchians have pillar-erected hindlimbs. Analysis of the musculoskeletal apparatus of the early dinosauriform Silesaurus opolensis challenges this view. This ornithodiran had pillar-erected hindlimbs like some pseudosuchians. This condition could be autapomorphic or represents a transitional state between adductor-controlled limb posture of early dinosauromorphs and the buttress-erected hindlimbs of dinosaurs. This sequence of changes is supported by Triassic tracks left by animals of the dinosaurian lineage. It was associated with the strong development of knee flexors and extensors. Furthermore, the forelimbs of Silesaurus were fully erect, analogously to those of early sauropods. Members of both lineages reduced the muscles related to the protraction, retraction and bending of the limb. They used forelimbs more as a body support and less for propulsion. A similar scapula and humerus construction can be found in the Lagerpetidae and Lewisuchus, suggesting that long, slender, fully erected forelimbs are primitive for all Dinosauromorpha, not just Silesauridae. Early dinosaurs redeveloped several muscle attachments on the forelimb, probably in relation to bipedality.

Bagaceratops rozhdestvenskyi is a ceratopsian dinosaur from the Late Cretaceous Baruungoyot Formation of the Gobi Desert, closely related to Protoceratops spp. Several Bag. rozhdestvenskyi skulls demonstrate a wide range of variation in their morphology and size. Here I argue that the observed variability is most likely of intraspecific nature. Specimens classified in a few allegedly distinct species from the same or near-contemporary sediments, namely Gobiceratops minutus, Lamaceratops tereschenkoi and Platyceratops tatarinovi from Baruungoyot Formation, and Magnirostris dodsoni from Bayan Mandahu, are younger subjective synonyms of Bag. rozhdestvenskyi. They plausibly represent an ontogenetic series within the latter. Breviceratops kozlowskiiis a distinct taxon. The evolutionary relationships within Protoceratopsidae are complicated by the mosaic distribution of plesiomorphic and derived features in distinct species. I suggest that taxa distribution and observed changes in morphology are an evidence for the ancestral position of Protoceratops andrewsi among protoceratopsids. It implies possible temporary separation between the geological formations of the Gobi Desert yielding distinct protoceratopsid species. The novel evolutionary scenario suggests number of convergences that occurred in Protoceratopsidae and Ceratopsoidea (reduction of the premaxillary dentition, fusion of nasals, development of the accessory antorbital fenestra). Present study reveals the significance of the intraspecific and ontogenetic variation in the study of the neoceratopsian taxonomy.