Fangchen Zhao

Sponge grade body fossil with cellular resolution dating 60 Myr before the Cambrian

Significance Phylogenomic extrapolations indicate the last common ancestor of sponges and eumetazoans existed deep in the Cryogenian, perhaps 200 million years (Myr) before the Cambrian (541 Ma). This inference implies a long Precambrian history of animals phylogenetically allied with sponges. However, there is yet little unequivocal paleontological evidence of Precambrian sponges. Here, we present a newly discovered 600-Myr-old fossil preserved at cellular resolution, displaying multiple poriferan features. The animal was covered with a dense layer of flattened cells resembling sponge pinacocytes, displaying a hollow tubular structure with apparent water inflow and outflow orifices. Although requiring additional specimens of similar form for confirmation, this finding is consistent with phylogenomic inference, and implies the presence of eumetazoan ancestors by 60 Myr before the Cambrian. An extraordinarily well preserved, 600-million-year (Myr)-old, three-dimensionally phosphatized fossil displaying multiple independent characters of modern adult sponges has been analyzed by SEM and synchrotron X-ray tomography. The fossilized animal (Eocyathispongia qiania gen. et sp. nov.) is slightly more than 1.2 mm wide and 1.1 mm tall, is composed of hundreds of thousands of cells, and has a gross structure consisting of three adjacent hollow tubes sharing a common base. The main tube is crowned with a large open funnel, and the others end in osculum-like openings to the exterior. The external surface is densely covered with flat tile-like cells closely resembling sponge pinacocytes, and this layer is punctuated with smaller pores. A dense patch of external structures that display the form of a lawn of sponge papillae has also survived. Within the main funnel, an area where features of the inner surface are preserved displays a regular pattern of uniform pits. Many of them are surrounded individually by distinct collars, mounted in a supporting reticulum. The possibility cannot be excluded that these pits are the remains of a field of choanocytes. The character set evinced by this specimen, ranging from general anatomy to cell type, uniquely indicates that this specimen is a fossil of probable poriferan affinity. So far, we have only this single specimen, and although its organized and complex cellular structure precludes any reasonable interpretation that its origin is abiogenic, confirmation that it is indeed a fossilized sponge will clearly require discovery of additional specimens.

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.

Decimetre-scale multicellular eukaryotes from the 1.56-billion-year-old Gaoyuzhuang Formation in North China

Fossils of macroscopic eukaryotes are rarely older than the Ediacaran Period (635–541 million years (Myr)), and their interpretation remains controversial. Here, we report the discovery of macroscopic fossils from the 1,560-Myr-old Gaoyuzhuang Formation, Yanshan area, North China, that exhibit both large size and regular morphology. Preserved as carbonaceous compressions, the Gaoyuzhuang fossils have statistically regular linear to lanceolate shapes up to 30 cm long and nearly 8 cm wide, suggesting that the Gaoyuzhuang fossils record benthic multicellular eukaryotes of unprecedentedly large size. Syngenetic fragments showing closely packed ∼10 μm cells arranged in a thick sheet further reinforce the interpretation. Comparisons with living thalloid organisms suggest that these organisms were photosynthetic, although their phylogenetic placement within the Eukarya remains uncertain. The new fossils provide the strongest evidence yet that multicellular eukaryotes with decimetric dimensions and a regular developmental program populated the marine biosphere at least a billion years before the Cambrian Explosion.

A deep root for the Cambrian explosion: Implications of new bio- and chemostratigraphy from the Siberian Platform

Much uncertainty remains as to the temporal relationship between the Ediacaran and Cambrian biotas, yet this is critical to our understanding of the rise of metazoans. Here we present new high-resolution carbon isotope chemostratigraphy and biostratigraphy for a terminal Ediacaran to Cambrian succession on the eastern Siberian Platform, Russia, which shows the presence of a succession of diverse fossil assemblages before the start of the basal Cambrian negative carbon isotope excursion (BACE). Soft-bodied Ediacaran biota ( Beltanelliformis ) occur before the start of the late Ediacaran positive carbon isotope plateau (EPIP), a mixed Ediacaran and Cambrian skeletal biota ( Cloudina , Anabarites , Cambrotubulus ) appears within the EPIP, and diverse Cambrian-type small shelly fossils including Protohertzina and other protocondonts, halkieriids, chancelloriids, hyoliths, hyolithelminthes, and the burrowing trace fossil Diplocraterion appear at the beginning of the BACE. These integrated data show that taxa attributed to so-called Ediacaran and earliest Cambrian skeletal biotas in fact overlap without notable biotic turnover, and thus refute the presence of a large isotope excursion coincident with mass extinction of all Ediacaran biota. We propose a new biozone, the Cloudina - Namacalathus - Sinotubulites Assemblage Zone, to precede the known small shelly fossil (SSF) zones. These observations raise doubts as to whether there is any true separation between the Ediacaran and Cambrian skeletal biotas, and suggest that there is a deep root for the Cambrian explosion of metazoans.

Meroblastic cleavage identifies some Ediacaran Doushantuo (China) embryo-like fossils as metazoans

Ediacaran Doushantuo (Formation) embryo-like fossils (EDEFs, ca. 600 Ma) from South China display cellular and sub-cellular structures and provide a unique window on the early evolution of multicellular eukaryotes. But there have been widely disparate interpretations of these fossils. Here we report new fossil embryo-like forms from the Doushantuo phosphorite that exhibit a meroblastic cleavage pattern. Our results from high-resolution propagation phase contrast–synchrotron radiation X-ray microtomography (PPC-SRµCT) demonstrate that these fossils preserve features directly comparable to those of modern meroblastic animal embryos that utilize discoidal cleavage. Given that discoidal-type meroblastic cleavage occurs only in metazoans, the phylogenetic positions of these fossils probably fall into the animal branch of the holozoan tree. Meroblastic as well as holoblastic cleavage forms were thus present by ca. 600 Ma, substantiating the conclusion derived from molecular clock estimates that a variety of metazoan lineages had evolved by the mid-Ediacaran after the termination of the Marinoan glaciation, if not earlier.

Appendages of an early Cambrian metadoxidid trilobite from Yunnan, SW China support mandibulate affinities of trilobites and artiopods

Appendage anatomy contributes crucial data for understanding the evolution and ecology of Euarthropoda. The Palaeozoic trilobites show a great diversity of exoskeletons in the fossil record. However, soft parts, especially appendages, have only been discovered from a few trilobite species. Here we report extraordinarily preserved appendages in the trilobite species Hongshiyanaspis yiliangensis Zhang & Lin in Zhang et al . 1980 (Redlichiida, Metadoxididae) from a single mudstone layer of the Xiazhuang fossil assemblage within the Hongjingshao Formation (Cambrian Series 2, Stage 3) near Kunming, Yunnan, SW China. The appendages exhibit the common architecture revealed by other trilobites and artiopods by consisting of a pair of uniramous antennae followed by a series of paired homonomous biramous limbs. The antennae in holaspid individuals comprise up to 27 spinous podomeres and their ontogeny occurs by lengthening of the podomeres. The post-antennal biramous limbs are similar to those in other polymerid trilobites and artiopods by having a single-segmented protopodite and an endopodite comprising seven segments, but possess a unique wide tripartite exopodite with long setae. Sophisticated appendage anatomy, including the body–limb junction, fine setae, putative muscle bundles and duct-type tissues, are also revealed. Appendages of trilobites, artiopods and other upper stem-group euarthropods are compared and summarized. The H. yiliangensis appendages highlight the high morphological disparity of exopodites and the conservativeness of endopodites in trilobites and artiopods. This morphological pattern, together with similar body patterning seen in crustaceans but not in chelicerates, supports the mandibulate affinities of trilobites and at least some artiopods.

First report of Wiwaxia from the Cambrian Chengjiang Lagerstätte

Research was supported by the Chinese Academy of Sciences (KZZD-EW-02-2); the National Basic Research Program of China (2013CB835006); the National Natural Science Foundation of China (41472012); the Natural Science Foundation of Jiangsu Province (BK2012893); and Clare College, Cambridge.

A New Helmetiid Arthropod From The Early Cambrian Chengjiang Lagerstätte, Southwest China

Abstract A new arthropod, Haifengella corona new genus new species is described from the early Cambrian Chengjiang Lagerstätte (Series 2, Stage 3), Yunnan Province, southwest China. It is readily assignable to helmetiida based on gross morphology of the tergum. The new helmetiid is unique in having the marginal spines extending over one-third of the total body width. The weakly sclerotized tergum consists of six thoracic tergites with edge-to-edge tergite articulations. The sub-trapezoidal cephalic shield has a pair of long spines projecting from each posterolateral corner. A prehypostomal sclerite (anterior sclerite) recesses in the anterior margin of the cephalic shield, and a pair of bulges that are close to the prehypostomal sclerite in the cephalic shield are presumed to be the position of ventral eyes. Each of the thoracic tergites exhibits a pair of long spines projecting from the posterolateral corners. The semicircular pygidium carries one terminal spine and two pairs of lateral spines.