Shixue Hu

The Luoping biota: exceptional preservation, and new evidence on the Triassic recovery from end-Permian mass extinction

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.

QUANTITATIVE ANALYSIS OF TAPHOFACIES AND PALEOCOMMUNITIES IN THE EARLY CAMBRIAN CHENGJIANG LAGERSTÄTTE

Abstract Detailed sedimentological and quantitative taphonomical analyses of 11,974 fossil specimens from an early Cambrian (Stage 3) Chengjiang-type 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 time-averaged 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.

Community structure and composition of the Cambrian Chengjiang biota

Based on previously published species data (228 species in over 18 phyla) and field sampling (114 species and 18406 individuals) in the Chengjiang-Haikou-Anning area, we analyzed quantitatively the paleocommunity composition and structure of the Cambrian Chengjiang biota (Cambrian Series 2, eastern Yunnan, China). Arthropods dominate the community both in species diversity (species: 37%) and in abundance (individuals: 51.8%). Priapulids (individuals: 22.6%) and brachiopods (individuals: 16.3%) follow in abundance rank. The arthropod Kunmingella douvillei (26.2%), the priapulid Cricocosmia jinning-ensis (15.4%), and the brachiopod Diandongia pista (11%) are the three most abundant species. Ecological analyses show that the community was dominated by epifaunal organisms (species: 63%, individuals: 68.4%) followed by infaunal organisms (species: 11.9%, individuals: 25.9%), nektobenthic organisms (species: 11.5%, individuals: 2.6%), and pelagic organisms (species: 5.3%, individuals: 3.1%). The diverse feeding strategies, dominated by suspension feeders (species: 35.6%, individuals: 26.1%) and hunter/scavengers (species: 31.1%, individuals: 40.4%), indicate the former existence of a complex food chain and intense competition. Epifaunal vagrant omnivores (28.2%), infaunal vagrant hunter/scavengers (19.8%), epifaunal sessile suspension feeders (17.7%), and epifaunal vagrant hunter/scavengers (15.3%) were the most abundant ecological groups, represented primarily by arthropods, poriferans, priapulids, and brachiopods. Ecological group analyses reveal that the early Cambrian Chengjiang biota is similar in community patterns and functional relations to modern biotas in shallow marine settings.

Diversity and species abundance patterns of the Early Cambrian (Series 2, Stage 3) Chengjiang Biota from China

Abstract Lagerstätten from the Precambrian–Cambrian transition have traditionally been a relatively untapped resource for understanding the paleoecology of the “Cambrian explosion.” This quantitative paleoecological study is based on 10,238 fossil specimens belonging to 100 animal species, 11 phyla, and 15 ecological categories from the lower Cambrian (Series 2, Stage 3) Chengjiang biota (Mafang locality near Haikou, Yunnan Province, China). Fossils were systematically collected within a 2.5-meter-thick sequence divided into ten stratigraphic intervals. Each interval represents an induced time-averaged assemblage of various event (obrution) beds of unknown duration. Overall, the different fossil assemblages are taxonomically and ecologically similar, suggesting the presence of a single community type recurring throughout the Mafang section. The Mafang community is dominated by epibenthic vagile hunters or scavengers, sessile suspension feeders, and infaunal vagile hunters or scavengers represented primarily by arthropods, brachiopods, and priapulids, respectively. Most species have low abundance and low occurrence frequencies, whereas a few species are numerically abundant and occur frequently. Overall, in structure and ecology the Mafang community is comparable to the Middle Cambrian (Series 3, Stage 5) Burgess Shale biota (Walcott Quarry, Yoho National Park, British Columbia, Canada). This suggests that, despite variations in species identity within taxonomic and ecological groups, the structure and ecology of Cambrian Burgess Shale-type communities remained relatively stable until at least the Middle Cambrian (Series 3, Stage 5) in subtidal to relatively deep-water offshore settings in siliciclastic soft-substrate environments.

A gigantic nothosaur (Reptilia: Sauropterygia) from the Middle Triassic of SW China and its implication for the Triassic biotic recovery

The presence of gigantic apex predators in the eastern Panthalassic and western Tethyan oceans suggests that complex ecosystems in the sea had become re-established in these regions at least by the early Middle Triassic, after the Permian-Triassic mass extinction (PTME). However, it is not clear whether oceanic ecosystem recovery from the PTME was globally synchronous because of the apparent lack of such predators in the eastern Tethyan/western Panthalassic region prior to the Late Triassic. Here we report a gigantic nothosaur from the lower Middle Triassic of Luoping in southwest China (eastern Tethyan ocean), which possesses the largest known lower jaw among Triassic sauropterygians. Phylogenetic analysis suggests parallel evolution of gigantism in Triassic sauropterygians. Discovery of this gigantic apex predator, together with associated diverse marine reptiles and the complex food web, indicates global recovery of shallow marine ecosystems from PTME by the early Middle Triassic.

CARBONATE RETICULATED RIDGE STRUCTURES FROM THE LOWER MIDDLE TRIASSIC OF THE LUOPING AREA, YUNNAN, SOUTHWESTERN CHINA: GEOBIOLOGIC FEATURES AND IMPLICATIONS FOR EXCEPTIONAL PRESERVATION OF THE LUOPING BIOTA

ABSTRACT Exceptionally preserved carbonate reticulated ridge structures are documented from the lower Middle Triassic Guanling Formation of the Luoping area, eastern Yunnan Province, southwestern China. Macrostructures suggest these reticulated ridge structures are morphologically similar to the growth-related structures in ancient and modern microbial mats. Other such features as minute load structures and pyrite crystals beneath the reticulated layers on bedding surfaces are overlain by a black carbonaceous shale layer. Clay minerals in the reticulated layers are oriented parallel to bedding at the top, and they usually form wavy laminae. These characteristics indicate that these carbonate reticulated ridge structures share a similar microstructure to their siliciclastic analogs, and are also comparable with that of wrinkle structures. SEM imaging shows that many tiny objects—coccoid spheres, rod-like structures and filamentous fibrils—are distinct within those ultra-thin, wavy laminae, suggesting similar accretion and growth models to modern microbial mats. Accordingly, the Luoping reticulated ridge structures are of biogenic origin. These microbial mats are associated with the early Middle Triassic Luoping Biota and they are thought to have played a crucial role in the preservation of the fossils through sealing and microbial coats. An SEM-EDS analysis on fossil fragments and their surrounding matrix reveals that microbial mats also played an important role in the phosphatization of Luoping fossils.

Macrurous Decapoda from the Luoping Biota (Middle Triassic) of China

Abstract A large collection of macrurous decapod crustaceans is recorded from the middle–late Anisian (Middle Triassic) Guanling Formation in Yunnan Province, China. A remarkable assemblage of over 20,000 vertebrate, invertebrate, and plant fossils collectively referred to as the Luoping Biota has been collected from quarries in the vicinity of the city of Luoping. Among these, arthropods including the decapods are the most common element although articulated fish and reptiles are also common. The decapods represent new taxa, including Koryncheiros luopingensis n. gen. n. sp. within Clytiopsidae, a newly elevated family within Erymoidea; Tridactylastacus sinensis n. gen. n. sp. within Glypheidae; and Yunnanopalinura schrami n. gen. n. sp. within Palinuridae. A single specimen has been referred to Palinuridae sp. Koryncheiros luopingensis exhibits a unique cheliped architecture and the second through fourth chelipeds are subchelate, an extremely rare configuration. Tridactylastacus sinensis also exhibits subchelate closures of pereiopods 2–4, but it bears a distinctive subchelate first pereiopod with an intercalated spine between the fingers on the distal margin of the propodus. Yunnanopalinura schrami represents the oldest occurrence of Palinuridae and Achelata. Collectively, these expand our knowledge of Chinese decapods significantly in that only six species of fossil decapods have been described previously from the country.

Nothosaur foraging tracks from the Middle Triassic of southwestern China

The seas of the Mesozoic (266-66 Myr ago) were remarkable for predatory marine reptiles, but their modes of locomotion have been debated. One problem has been the absence of tracks, although there is no reason to expect that swimmers would produce tracks. We report here seabed tracks made by Mesozoic marine reptiles, produced by the paddles of nothosaurs (Reptilia, Sauropterygia) in the Middle Triassic of the Luoping localities in Yunnan, southwestern China. These show that the track-making nothosaurs used their forelimbs for propulsion, they generally rowed (both forelimbs operating in unison rather than alternately), and the forelimb entered medially, dug in as the paddle tip gained purchase, and withdrew cleanly. These inferences may provide evidence for swimming modes, or it could be argued that the locomotory modes indicated by the tracks were restricted to such contact propulsion. Such punting behaviour may have been used to flush prey from the bottom muds.

Penaeoid Decapoda (Dendrobranchiata) from the Luoping Biota (Middle Triassic) of China: systematics and taphonomic framework

Abstract Two new genera, Anisaeger and Distaeger, and three new species, Anisaeger brevirostrus, A. spiniferus, and Distaeger prodigiosus, extend the range of the Aegeridae (Dendrobranchiata, Penaeoidea) into the Middle Triassic (Anisian) of China. Seven decapod crustacean species are now known from the Luoping biota of southern China. Morphological features of shrimp that are present but rarely mentioned in the neontological literature are recognized as potentially useful in classifying fossil material, including a diaeresis on the exopod of the uropods and multiarticulate flagellae on pleopods. Unusual taphonomic features of the shrimp include fractured cuticle, preservation in lateral, dorsal, and ventral position, and twisted cephalothoraxes.

A New Shrimp (Decapoda, Dendrobranchiata, Penaeoidea) from the Middle Triassic of Yunnan, Southwest China

Abstract A new penaeoid shrimp collected from the Middle Triassic Member II of the Guanling Formation in the vicinity of the city of Luxi, Yunnan, southwest China, is a new species, Aeger luxii n. sp. The new species possesses prominent spinose third maxillipeds, which is one of the typical characteristics of Aeger. The new species differs from the type species, Aeger tipularius from the Jurassic Solnhofen Plattenkalk, in having a long, smooth rostrum with no subrostral spines. The new taxon increases the diversity of Chinese decapods, and further expands our knowledge of the phylogeny and evolution of the Mesozoic decapods. The find is the first complete specimen of Aeger in the Middle Triassic, and reveals a close biogeographic connection of the marine ecosystem between Eastern and Western Tethys.