Hyoliths are extinct enigmatic organisms of early lophotrochozoan affinity known globally from the Palaeozoic Era and were especially diverse and abundant in the Cambrian Period. However, the commonly incomplete preservation of hyolith exoskeletons and our limited knowledge of their soft anatomy makes their ecological and biological aspects unclear. Konservat-Lagerstätte are crucial windows to unlock the mysteries of hyoliths. Here we report a new occurrence of exceptionally preserved hyolithid hyoliths from the middle Cambrian Mantou Formation (Miaolingian, Wuliuan) in Shandong Province, North China. The preserved soft organs of the new species Novakotheca weifangensis sp. nov. include a U-shaped gut and possible pharynx, oesophagus, stomach and digestive gland, which provide significant new information for the reconstruction of the digestive system of hyolithids. Two taphonomic modes of hyoliths described herein are recognized: soft tissue preservation through pyritization and three-dimensional shell preservation through phosphatization. Morphological variations due to different preservational pathways in the same species are revealed, highlighting the taphonomic bias on taxonomy. The ecological association between hyoliths and small brachiopod epibionts is a direct example of species interactions, providing insights into the ecological structures and adaptability of early animals during Cambrian time.

The hurdiid radiodont Cambroraster is characterized by a prominent horseshoe-shaped head carapace, which bears conspicuous posterolateral spinous processes and covers half of the body length, and a pair of frontal appendages possessing rake-like endites. Here we report the discovery of characteristic central and lateral elements as well as a possible appendage endite for Cambroraster from the Cambrian Wuliuan Mantou Formation of North China. This discovery suggests that Cambroraster had dispersed over a wide biogeographic range and expands the geographic range of Hurdiidae. The increasing discoveries of exceptionally preserved fossils in the Upper Shale Member of the Mantou Formation indicate that this horizon is a promising Burgess Shale–type Lagerstätte.

The arthropod Sidneyia Walcott, 1911 is a remarkable animal of the Burgess Shale biota (Cambrian Miaolingian, Wuliuan; British Columbia, Canada), which has not been confidently reported from other Cambrian Konservat-Lagerstätten. Here we report the discovery of Sidneyia cf. inexpectans from the Wuliuan Mantou Formation of North China, which substantially expands the known palaeogeographical distribution of this genus. Our discovery suggests that Sidneyia had much greater dispersal ability than hitherto thought. It also confirms the presence of exceptionally preserved fossils in the Wuliuan Mantou Formation, one of the rare Burgess Shale-type deposits of North China.

Non-spicular sponges constitute >8% of the extant sponge biodiversity at the species level, yet their evolutionary history is poorly known due to a sparse fossil record. The genus Vauxia, previously only known from middle Cambrian (Miaolingian, Wuliuan) Lagerstätten, was regarded as the earliest fossil record of non-spicular demosponges. Here we describe the first vauxiid sponge, Vauxia leioia new species, from the early Cambrian Chengjiang Biota (Series 2, Stage 3). This sponge exhibits a double-layered fibrous skeleton: the mesh and fiber thickness of the endosomal layer are irregular while the dermal layer, which directly connects with the endosomal skeleton without intermediate supporting fibers, is regular in both aspects. Measurements using scanning electron microscope and Raman spectroscopy revealed that the endosomal fibers are composed of carbonaceous material, but are tomographically indiscernible from the host rock, while the dermal fibers are preserved as impressions without obvious accumulation of carbonaceous material. Although the original composition of the dermal skeleton is now hard to establish, we cannot rule out that it was siliceous. The morphological characters of V. leioia n. sp. represent an intermediate state between other Vauxia species and the recently established vauxiid genus Angulosuspongia. However, more data are required to reconstruct the phylogenetic relationship among these taxa.

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Malongitubus kuangshanensis Hu, 2005 from the early Cambrian Chengjiang Lagerstätte of China is redescribed as a pterobranch and provides the best evidence to demonstrate that hemichordates were present as early as Cambrian Stage 3. Interpretation of this taxon as a hemichordate is based on the morphology of the branched colony and the presence of resistant inner threads consistent with the remains of an internal stolon system. The presence of fusellar rings in the colonial tubes cannot be unambiguously proven for Malongitubus, probably due to early decay and later diagenetic replacement of the thin organic material of the tubarium, although weak annulations are still discernible in parts of the tubes. The description of M. kuangshanensis is revised according to new observations of previously reported specimens and recently collected additional new material. Malongitubus appears similar in most features to Dalyia racemata Walcott, 1919 from the Cambrian Stage 5 Burgess Shale, but can be distinguished by the existence of disc-like thickenings at the bases of tubarium branching points in the latter species. Both species occur in rare mass-occurrence layers with preserved fragmentary individuals of different decay stages, with stolon remains preserved as the most durable structures. Benthic pterobranchs may have occurred in some early Cambrian shallow marine communities in dense accumulations and provided firm substrates and shelter for other benthic metazoans as secondary tierers.

The radiodontans, including anomalocaridids and their allies, are enigmatic stem-group euarthropods and are the most ancient apex giant predators known from the fossil record. Most studies on their feeding behaviors have emphasized their diverse and abundant raptorial frontal appendages, while the oral cone surrounding the mouth opening in these animals has attracted less attention. At present, three oral cone morphotypes are known, from Anomalocaris Whiteaves, 1892, Peytoia Walcott, 1911, and Hurdia Walcott, 1912, respectively. In this paper, we report on a novel form of radiodontan oral cone from the Guanshan Lagerstätte (Cambrian Series 2, Stage 4) in the Wulongqing Formation, eastern Yunnan, South China. This oral cone is unique in combining features seen in Peytoia/Hurdia and Anomalocaris. It possesses a Peytoia/Hurdia-type ‘tetraradial’ configuration comprising a 32-plate outer ring that consists of four perpendicularly arranged large plates and 28 small plates, in addition to furrowed folds and scale-like nodes on plate surfaces otherwise seen only in Anomalocaris. As an intermediate morphotype, the Guanshan oral cone improves our understanding of the occurrence and morphological disparity of radiodontan oral cones, illuminates future investigations on potentially variable radiodontan feeding mechanisms, and reveals possible evolutionary transformations of these peculiar feeding structures. The resolution of current radiodontan phylogeny would be potentially improved by new knowledge on other body parts apart from frontal appendages in future studies.

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.

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 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.