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The early Cambrian Carinachitidae, a family in the subclass Conulata, are intriguing and important small shelly fossils. Their gently tapering, tube-shaped skeletons consist of convex faces separated from each other by broad, deep corner sulci, and they exhibit triradial, pentaradial, or predominantly tetraradial symmetry. However, the morphology of the aperture and the modes of growth of carinachitid skeletons as well as the anatomy of their soft parts are unknown. Examination of a single new, exceptionally well-preserved specimen of tetramerous Carinachites spinatusQian, 1977, collected from the lower Cambrian Kuanchuanpu Formation in South China, reveals: (1) that its aperture is connected to a small mass of relic soft tissue and (2) that the apertural end of each of the four faces is developed into a subtriangular lappet or oral lobe that is smoothly folded toward the long axis of the tube, partially closing the tube aperture. Similarities between thorn-like spines on the faces and the oral lobes indicate that the transverse ribs were periodically displaced from the perradial portion of the aperture during formation of new ribs. In addition, the tube walls may have undergone secondary thickening during growth. The growth pattern of the tube and the spatial relationships between the tube aperture and soft parts are analogous to those of co-occurring olivooids. These findings further strengthen the previously proposed hypothesis that coeval carinachitids, olivooids, hexangulaconulariids, and Paleozoic conulariids are closely related taxa within the subphylum Medusozoa. Finally, carinachitids most likely represent an evolutionary intermediate between olivooids and hexangulaconulariids.
Scalidophoran worms diversified in the Cambrian Fortunian, as indicated by recent reports from this stage, with two described species and two more unnamed forms exclusively from Orsten-type Lagerstätten yielding three-dimensionally phosphatized fossils. Here, we report new material of scalidophoran worms in Orsten-type preservation from the Cambrian Fortunian Xinli section in northern Sichuan Province, South China. At least five forms of scalidophoran worms were recovered from this location, including Eokinorhynchus rarusZhang et al., 2015 and four unnamed taxa—Forms A, B, C, and D. Co-occurring disassociated spinose small shelly fossils might also be isolated cuticular elements of these early scalidophoran worms. The ontogeny of Eokinorhynchus rarus is revised. Forms A, C, and D are assigned to total-group Scalidophora to indicate their uncertain positions within Scalidophora, while Form B might be a close relative of Eokinorhynchus rarus. The current work highlights the significance of Orsten-type Lagerstätten in uncovering the morphology, ontogeny, and taxonomy of early Scalidophora and Cycloneuralia, made particularly available by the new finds in China.
Malongitubus kuangshanensisHu, 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 racemataWalcott, 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 kutorginates are commonly the most abundant rhynchonelliform brachiopod found in the early Cambrian; they are also some of the oldest known rhynchonelliforms, first appearing in the Unnamed Series 2 (Atdabanian equivalent) and becoming extinct sometime in Cambrian Series 3 (Amgaian equivalent). Moreover, kutorginates are the first known member of the rhynchonelliforms for which we have a detailed knowledge of their soft-part anatomy, including the lophophore, digestive tract, and pedicle-all exceptionally preserved in Kutorgina chengjiangensisZhang et al., 2007 from the early Cambrian Chengjiang Lagerstätte of southern China. The stout and annulated pedicle in the original report was described as protruding between the valves; however, newly collected better—preserved material now clearly shows that the pedicle actually protrudes from the apical perforation of Kutorgina chengjiangensis. This type of apical pedicle has also been described from other early Cambrian rhynchonelliforms, including the problematic chileate Longtancunella chengjiangensis (Zhang et al., 2011a). Exceptionally preserved similar pedicles are also known to emerge apically from the Silurian chileate dictyonellid Eichwaldia subtrigonalisBillings, 1858, as well as from the recently described Silurian chileate Trifissura rigidaHolmer, Popov, and Bassett, 2014. However, it is clear that the only other exceptionally preserved kutorginate—a silicified Nisusia-was provided with an adult pedicle emerging between the valves from a posterior gap; thus, Nisusia has two pedicle openings. However, the apical foramen may represent the earliest attachment of the larvae, which subsequently became nonfunctional through ontogeny. It is suggested that both types of attachment strategies may have appeared early in the stem lineage of the Rhynchonelliformea.
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 AnomalocarisWhiteaves, 1892, PeytoiaWalcott, 1911, and HurdiaWalcott, 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.
The Cambrian Yanwangbian assemblage (Series 2, Stage 4) in South Shaanxi, China, is one of the Burgess Shale-type faunas as it represents the only relatively diverse Cambrian biota from the north margin of the Yangzte Platform, South China. The paleoscolecids (Cycloneuralia) illustrated herein are one of the major components of the fauna, although they appear to be much less abundant than skeletonized fossils, according to available collections. A new taxon, Shaanxiscolex xixiangensis new genus new species, is described based on the scleritome pattern: each annulus has two rows of alternating Hadimopanella-type plates positioned close to the borders, and a mosaic pattern of microplates occurs between the plates and within intersegmental furrows. The occurrence of the new taxon confirms a fairly diversified and widespread distribution of paleoscolecidan worms recognized in the early Cambrian of South China. Moreover, the ecology of paleoscolecids is reappraised based on burial position of the S. xixiangensis and gut contens of Cambrian taxa from South China, hinting that paleoscolecids (at least some taxa) were both deposit feeders and carnivores, as well as active bioturbators in the substrates of the Cambrian sea, which sheds new light on the ‘Cambrian Substrate Revolution.’
Disarticulated net-like plates of the lobopod Microdictyon had a near cosmopolitan distribution from the early to middle Cambrian but are yet to be documented from the North China Platform. Here we report isolated plates of Microdictyon from the lower Cambrian Xinji Formation (Stage 4, Series 2) of the North China Platform, extending the paleogeographic distribution of Microdictyon in the early Cambrian. The plates of Microdictyon from the Xinji Formation are similar to those of other species established on the basis of isolated plates but do bear some new characters, such as mushroom-shaped nodes with a single inclined platform-like apex and an upper surface that displays radial lines. However, the plates documented here are left under open nomenclature due to inadequate knowledge of intraspecific and ontogenetic variation and low specimen numbers. Through comparison of the node shapes of the isolated plates of different Microdictyon species, we consider that low mushroom-shaped nodes could be a primitive and conservative character of Microdictyon while tall mushroom-shaped nodes may be a derived character. Subtle differences in shape and number of node apices may also represent intraspecific or ontogenetic variation.
New morphological features of the mobergellan Discinella micans (Billings, 1871) from the lower Cambrian (Stage 4) of Northeast Greenland and southern Labrador are described. The new features include: (1) the morphology of the larval shell, which is shown to be cap-shaped, subcircular, and with impressions of the internal muscle attachment scars; (2) a range of unusual shell deformations (changes in growth direction resulting in thickened shells, partial detachment of shell laminae and subsequent regrowth, internal projections of shell material increasing the depth of the shell by up to 150%, disturbances and irregular fusion of muscle scars). In addition, we provide new details about the variability in number and shape of the anteriormost internal muscle scars, which often fuse and may vary in number from one to three (resulting in nine to 11 scars in total). Together the new observations provide additional strength for the hypothesis that mobergellan shells represent opercula of an as yet unknown tubular organism.
Rare specimens of eldonioids recovered from the lower Cambrian (Series 2, Stage 4) Emu Bay Shale (EBS) Konservat-Lagerstätte represent the first record of the group for the Cambrian of East Gondwana. The disc-shaped body of the EBS taxon bears fine concentric corrugations on the dorsal surface and, ventrally, a series of internal lobes that have primary and secondary bifurcations, as well as a coiled sac. It appears to be most similar to Rotadiscus and Pararotadiscus of the Cambrian Chengjiang and Kaili biotas of South China, respectively. While the structure of the internal lobes would indicate that this occurrence in the EBS represents a new taxon within the Rotadiscidae, lack of detail regarding the precise number of internal lobes and the condition of the circumoral tentacles warrants a more conservative approach in leaving the genus and species under open nomenclature. The EBS specimens also host trace fossils, including the remains of a burrow, which are generally lacking in the body-fossilbearing layers of the Konservat-Lagerstätte interval. These traces appear to have been made by small organisms and are similar to traces associated with the discs of Pararotadiscus guizhouensis (Zhao and Zhu, 1994) from the Kaili Biota. The available taphonomic, paleoenvironmental, and ichnological evidence indicates that the EBS eldonioids are most likely vagrants that were transported or settled into the ‘preservational trap’ and subsequently exposed on the substrate for a brief period before burial, thereby allowing organisms to exploit their carcasses for nutrients or other purposes.
Some rare microscopic cycloneuralians are present in the Cambrian of South China, represented by Eopriapulites and Eokinorhynchus (both early Cambrian), fossil embryos of Markuelia (middle to late Cambrian), and palaeoscolecids (early to late Cambrian). Among them, palaeoscolecids are relatively diverse and abundant. Here, we describe new material of three-dimensionally phosphatized and microscopic cycloneuralians from the Paibian Stage of Wangcun Lagerstätte, western Hunan, South China. New material includes fossil embryos assignable to Markuelia sp., two other types of fossil embryos, and three species of palaeoscolecids, including Dispinoscolex decorusDuan, Dong, and Donoghue, 2012, Schistoscolex hunanensisDuan, Dong, and Donoghue, 2012, and Austroscolex sinensis new species. The palaeoscolecid fragments differ mainly in size and armor of the trunk annuli. Since Eokinorhynchus and Eopriapulites occurred the earliest among the Cambrian cycloneuralians, it is proposed here that: (1) cycloneuralians originated in the Cambrian Fortunian small shelly faunas rather than in the early Cambrian macrobenthos, (2) ancestral cycloneuralians may have simple trunk armor, and (3) Eopriapulites represents an ancestral cycloneuralian.
Radiodontan body elements, some belonging to Peytoia and Hurdia and some unassigned, have been reported from the Langston Formation (Spence Shale Member), Wheeler Formation, and Marjum Formation of the middle Cambrian (Series 3) of Utah. These identifications are reassessed in light of recent work on the morphology of the radiodontan Hurdia. New specimens of Hurdia are identified from the Spence Shale, representing mouthparts (oral cones), cephalic carapace H-elements, frontal appendages, and a single isolated swimming flap. The shape of the H-elements allows H. victoriaWalcott, 1912 to be identified from the Spence Shale for the first time. The flap is larger and more complete than any reported from the Burgess Shale and allows for a better understanding of the morphology of Hurdia swimming flaps. A 3D model of a Hurdia frontal appendage indicates that there is only one morph of Hurdia frontal appendage found in both species, and apparent morphological differences between disarticulated appendages reflect a preservational continuum caused by varying oblique angles relative to the seafloor. Peytoia should no longer be reported from the Spence Shale, but its presence is confirmed in the Wheeler and Marjum formations. New mouthparts (oral cones) of Hurdia from the Spence Shale and Peytoia from the Marjum Formation with surface textures of submillimeter-diameter raised nodes are described. These new features have not been observed in material from the Burgess Shale and suggest slight differences in preservation.