No abstract
The timing and nature of biotic recovery from the devastating end-Permian mass extinction (252 Ma) are much debated. New studies in South China suggest that complex marine ecosystems did not become re-established until the middle–late Anisian (Middle Triassic), much later than had been proposed by some. The recently discovered exceptionally preserved Luoping biota from the Anisian Stage of the Middle Triassic, Yunnan Province and southwest China shows this final stage of community assembly on the continental shelf. The fossil assemblage is a mixture of marine animals, including abundant lightly sclerotized arthropods, associated with fishes, marine reptiles, bivalves, gastropods, belemnoids, ammonoids, echinoderms, brachiopods, conodonts and foraminifers, as well as plants and rare arthropods from nearby land. In some ways, the Luoping biota rebuilt the framework of the pre-extinction latest Permian marine ecosystem, but it differed too in profound ways. New trophic levels were introduced, most notably among top predators in the form of the diverse marine reptiles that had no evident analogues in the Late Permian. The Luoping biota is one of the most diverse Triassic marine fossil Lagerstätten in the world, providing a new and early window on recovery and radiation of Triassic marine ecosystems some 10 Myr after the end-Permian mass extinction.
Although palaeontological evidence from exceptional biota demonstrates the existence of diverse marine communities in the Early Cambrian (approx. 540-520 Myr ago), little is known concerning the functioning of the marine ecosystem, especially its trophic structure and the full range of ecological niches colonized by the fauna. The presence of a diverse zooplankton in Early Cambrian oceans is still an open issue. Here we provide compelling evidence that chaetognaths, an important element of modern zooplankton, were present in the Early Cambrian Chengjiang biota with morphologies almost identical to Recent forms. New information obtained from the lowermost Cambrian of China added to previous studies provide convincing evidence that protoconodont-bearing animals also belonged to chaetognaths. Chaetognaths were probably widespread and diverse in the earliest Cambrian. The obvious raptorial function of their circumoral apparatuses (grasping spines) places them among the earliest active predator metazoans. Morphology, body ratios and distribution suggest that the ancestral chaetognaths were planktonic with possible ecological preferences for hyperbenthic niches close to the sea bottom. Our results point to the early introduction of prey-predator relationships into the pelagic realm, and to the increase of trophic complexity (three-level structure) during the Precambrian-Cambrian transition, thus laying the foundations of present-day marine food chains.
Exceptional fossil specimens with preserved soft parts from the Maotianshan Shale (ca 520 Myr ago) and the Burgess Shale (505 Myr ago) biotas indicate that the worldwide distributed bivalved arthropod Isoxys was probably a non-benthic visual predator. New lines of evidence come from the functional morphology of its powerful prehensile frontal appendages that, combined with large spherical eyes, are thought to have played a key role in the recognition and capture of swimming or epibenthic prey. The swimming and steering of this arthropod was achieved by the beating of multiple setose exopods and a flap-like telson. The appendage morphology of Isoxys indicates possible phylogenetical relationships with the megacheirans, a widespread group of assumed predator arthropods characterized by a pre-oral 'great appendage'. Evidence from functional morphology and taphonomy suggests that Isoxys was able to migrate through the water column and was possibly exploiting hyperbenthic niches for food. Although certainly not unique, the case of Isoxys supports the idea that off-bottom animal interactions such as predation, associated with complex feeding strategies and behaviours (e.g. vertical migration and hunting) were established by the Early Cambrian. It also suggests that a prototype of a pelagic food chain had already started to build-up at least in the lower levels of the water column.
An updated reconstruction of the body plan, functional anatomy and life attitude of the bradoriid arthropod Kunmingella is proposed, based on new fossil specimens with preserved soft parts found in the lower Cambrian of Chengjiang and Haikou (Yunnan, SW China) and on previous evidence. The animal has a single pair of short antennae pointing towards the front (a setal pattern indicates a possible sensory function). The following set of seven appendages (each composed of a 5-segmented endopod and a leaf-like exopod fringed with setae) is poorly differentiated, except the first three pairs (with possible rake-like endopodial outgrowths, smaller exopods) and the last pair of appendages (endopod with longer and more slender podomeres). The endopods are interpreted as walking legs with a possible role in handling food particles (marginal outgrowth with setae). The leaf-like exopods may have had a respiratory function. The trunk end is short, pointed, flanked with furcal-like rami and projects beyond the posterior margin of the carapace. The attachment of the body to the exoskeleton is probably cephalic and apparently lacks any well-developed adductor muscle system. The inferred life attitude of Kunrningella (e.g. crawling on the surface of the sediment) was that of a dorsoventrally flattened arthropod capped by a folded dorsal shield (ventral gape at least 120 O), thus resembling the living ostracode Manawa. The animal was also probably able to close its carapace as a response to environmental stress or to survive unfavourable conditions (e.g. buried in sediment). The anterior lobes of the valves are likely to have accommodated visual organs (possibly lensless receptors perceiving ambient light through the translucent head shield). Preserved eggs or embryos suggest a possible ventral brood care. The presence of Kunmingella in coprolites and its numerical abundance in Chengjiang sediment indicate that bradoriids constituted an important source of food for larger predators. Kunmingella differs markedly from the representatives of the crown group Crustacea (extant and Cambrian taxa) and from the stem group derivatives of Crustacea (exemplified by phosphatocopids and some 'Orsten' taxa) in showing no major sign of limb specialization (e.g. related to feeding strategies). Although it resembles other Chengjiang euarthropods in important aspects of its body plan (eg. uniramous antennae, endopod/exopod configuration), Kunrningella possesses several features (e.g. antennal morphology, post-antennular appendages with 5-segmented endopods) which support the view that bradoriids may be very early derivatives of the stem line Crustacea. OArthropoda,
Detailed sedimentological and quantitative taphonomical analyses of 11,974 fossil specimens from an early Cambrian (Stage 3) Chengjiangtype deposit near Haikou, Yunnan, reveals significant relationships between the original depositional environments and the composition and preservation of their respective fossil assemblages. In general, the Maotianshan Shale is characterized by superimposed couplets of laminated background and thin event mudstone layers representing two distinct taphofacies, A and B, respectively. Fossils in taphofacies A consist predominantly of indeterminate organic elements and fecal or algal strings with few, poorly preserved, soft-bodied animals. Among those, disarticulated arthropods account for 84.3% of specimens (mostly isolated valves of Kunmingella douvillei) and 51.4% of species. Poriferans represent 7.4% of specimens and 22.9% of species. Fossils in this taphofacies have undergone significant pre-or syn-burial decay and represent limited timeaveraged assemblages exhibiting low species richness. By contrast, taphofacies B contains greater numbers of species and specimens and better preserved soft-bodied animals. Taphofacies B represents mostly smothered organisms by distal tempestites. Arthropods are also dominant in taphofacies B, both in terms of species richness (41%) and abundance of specimens (44%). Poriferans, priapulids, lobopods, and brachiopods exhibit similar low species richness (6-8% each), but poriferans and lobopods are numerically rare, at around 1% each, whereas priapulids and brachiopods make up 26% and 24% of specimens, respectively. The arthropod Kunmingella douvillei (19%), the priapulid Cricocosmia jinningensis (19%), and the brachiopod Diandongia pista (18%) are the most abundant species in taphofacies B. Fossil assemblages in taphofacies A and B have similar recurrent and abundant species and similar temporal trends in evenness and richness, but taphofacies A captures only a portion of the species that are preserved in taphofacies B. These results suggest that the fossil assemblages present in both taphofacies represent a single local community subjected to two different taphonomic processes and imply similar recurrent environmental conditions within the section studied.
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