Chengjiang Biota Research Articles

New tubicolous problematic fossil with some “lophophorate” affinities from the Early Cambrian Chengjiang biota in south China

A new tubicolous problematic fossil Cambrocornulitus rarus n. gen. n. sp. (Cornulitida?) from the Early Cambrian Chengjiang biota with well-preserved soft body tissues is described. This tubicolous fossil probably had a biomineralized shell, and shares some affinities with cornulitids and lophophorates. C. rarus has a straight to curved, conical tube with external transversal rings (ridges), longitudinal ribs (striae) and possibly microlamellar wall structure. The preserved soft body of C. rarus exhibits several important morphological characteristics, such as a possible lophophore and nephridial ridges. The new genus had a somewhat similar mode of life to the gregarious type of cornulitids. Some of the studied tubes of C. rarus show malformed development.

Multi-Segmented Arthropods from the Middle Cambrian of British Columbia (Canada)

A new arthropod,Kootenichela deppin. gen. n. sp., is described from the Stanley Glacier exposure of the middle Cambrian (Series 3, Stage 5) Stephen Formation in Kootenay National Park (British Columbia, Canada). This taxon possesses a number of primitive arthropod features such as an elongate, homonomous trunk (consisting of at least 29 segments), poorly sclerotised trunk appendages, and large pedunculate eyes associated with an anterior (ocular) sclerite. The cephalon encompasses a possible antenna-like appendage and enlarged raptorial appendages with a bipartite peduncle and three spinose distal podomeres, indicative of megacheiran (“great-appendage” arthropod) affinities. The relationships of megacheirans are controversial, with them generally considered as either stem-euarthropods or a paraphyletic stem-lineage of chelicerates. An extensive cladistic analysis resolvedKootenichelaas sister-taxon to the enigmaticWorthenella cambriafrom the middle Cambrian (Series 3, Stage 5), Burgess Shale Formation in Yoho National Park (British Columbia), which is herein reinterpreted as a megacheiran arthropod. Based on their sister-group relationship, both taxa were placed in the new family Kootenichelidae, to whichPseudoiuliafrom the Chengjiang biota is also tentatively assigned. All of these taxa possess an elongate, multi-segmented body and subtriangular exopods. This family occupies a basal position within a paraphyletic Megacheira, the immediate outgroup of Euarthropoda (crown-group arthropods). The resultant topology indicates that analyses that have resolved megacheirans as stem-chelicerates have done so because they have rooted on inappropriate taxa, e.g., trilobitomorphs and marrellomorphs.

A sclerite-bearing stem group entoproct from the early Cambrian and its implications

The Lophotrochozoa includes disparate tentacle-bearing sessile protostome animals, which apparently appeared in the Cambrian explosion, but lack an uncontested fossil record. Here we describe abundant well preserved material of Cotyledion tylodes Luo et Hu, 1999, from the Cambrian (Series 2) Chengjiang deposits, reinterpreted here as a stem-group entoproct. The entoproct affinity is supported by the sessile body plan and interior soft anatomy. The body consists of an upper calyx and a lower elongate stalk with a distal holdfast. The soft anatomy includes a U-shaped gut with a mouth and aboral anus ringed by retractable marginal tentacles. Cotyledion differs from extant entoprocts in being larger, and having the calyx and the stalk covered by numerous loosely-spaced external sclerites. The description of entoprocts from the Chengjiang biota traces the ancestry of yet another lophotrochozoan phylum back to the Cambrian radiation, and has important implications for the earliest evolution of lophotrochozoans.

Spatial variation in the diversity and composition of the Lower Cambrian (Series 2, Stage 3) Chengjiang Biota, Southwest China

Strata hosting the Lower Cambrian (Series 2, Stage 3) Chengjiang Biota (Maotianshan Shale Member, Yu'anshan Formation) occur throughout the eastern part of Yunnan Province, Southwest China. In this study, literature-based faunal inventories from 10 areas (representing 34 localities), together with 22,038 new specimens collected at three localities from three of the 10 areas, were analyzed quantitatively to assess large-scale spatial variation in taxonomic diversity and composition. Our analyses show substantial co-variation between local paleoenvironmental settings and species diversity, and suggest the presence of three general taphofacies in the Maotianshan Shale Member, from west to east: low diversity (26 species) characterizes the Wuding area, where fossiliferous strata have undergone extensive bioturbation; high diversity (215 species) characterizes the Chengjiang–Haikou–Anning areas, where fossiliferous strata consist of stacked couplets of thin event and background mudstone layers; moderate diversity (55 species) characterizes the Malong area, where fossiliferous strata consist of indistinctly bedded shale showing fewer single-event layers but abundant background layers. In spite of variations in sampling effort between sites, spatial patterns in species diversity and community composition were controlled primarily by variation in depositional environments along a proximal offshore to lower shoreface gradient, which influenced both ecological and taphonomic factors. The offshore settings (Haikou–Chengjiang–Anning areas) show relatively high species diversity and exceptional preservation of soft-bodied fossils, reflecting rapid burial in distal tempestites. Relatively high degrees of bioturbation, time-averaging and hydrodynamic disturbance caused a substantial decrease in the quality of preservation and number of soft-bodied species in the Wuding and Malong areas. Arthropods are the most diverse group in most localities, followed by priapulids, poriferans, brachiopods, lobopods and hyoliths.This study shows that the Chengjiang Biota lends itself very well to high resolution characterization of spatial variation in taxonomic diversity, faunal composition and fossil preservation. Furthermore, the Chengjiang Biota may provide a unique opportunity to assess the roles of environmental factors, taphonomy and ecological controls on species diversity at local to regional spatial scales.

Evolution from an anoxic to oxic deep ocean during the Ediacaran–Cambrian transition and implications for bioradiation

The Ediacaran–Cambrian transition, one of the most critical intervals in Earth's history, is marked by dramatic biological, oceanic and geochemical turnovers. Here high-resolution carbon and sulfur isotopic data respectively for organic carbon and pyrite, and iron speciation data are presented from the deep-water Liuchapo and Niutitang Formations on the Yangtze block, South China. The carbon isotopic data, together with biostratigraphic and radiometric dating, offer the compelling evidence for the placement of Ediacaran–Cambrian boundary within the Liuchapo Formation (chert succession), and for its correlation with shallow-water equivalents elsewhere. In this context, iron speciation and sulfur isotopic data further suggest a predominant anoxic and ferruginous deep ocean over the transitional time until the middle Early Cambrian (Atdabanian or Stage 3) when the deep ocean was rapidly oxygenated. Coincidently, during this interval, large-body metazoans (i.e., sponges) abruptly appeared in the deep ocean, which was temporally associated with the highly diversified large-body skeletonized animals (i.e., Chengjiang Biota) which colonized in shallow-water niches particularly in southwestern China. This scenario suggests a causal link between deep oceanic oxygenation and the explosive diversification of large-body skeletonized organisms in the Early Cambrian.

New artiopodan arthropods from the early Cambrian Emu Bay Shale Konservat-Lagerstätte of South Australia

The Emu Bay Shale Konservat-Lagerstätte (Cambrian Series 2, Stage 4) on Kangaroo Island, South Australia, is the source of two new non-biomineralized artiopodan arthropods.Squamacula buckorumn. sp. is the first record outside of China of a genus otherwise known only from its type species,S. clypeata, from the Chengjiang biota. The Australian species displays the long cephalic doublure and spiniform exopod setae that are apomorphic for this genus, provides new information on the alimentary tract and midgut glands (the latter preserved as three-dimensional, permineralized structures), and indicates interspecific variability in trunk segment numbers. The distribution ofSquamaculastrengthens the biogeographic connections between early Cambrian “Burgess Shale-type” biotas of Australia and South China.Australimicola spriggin. gen. n. sp. represents a monotypic genus resolved in a cladistic analysis of Cambro-Ordovician artiopodans as most closely related to or within Conciliterga (a clade containingHelmetia,Kuamaia,Kwanyinaspis,Rhombicalvaria,Saperion,Skioldia, andTegopelte). Compared with other members of this clade from Chengjiang and the Burgess Shale, the new genus is diagnosed by an elongate trunk with 23 thoracic tergites having spatulate pleural tips and a small pygidium possessing a single, elongate pair of pleural spines, with specimens also showing a hypostome attached to an anterior (or prehypostomal) sclerite, antennae, short endopods, an annulated alimentary tract, and a series of three-dimensional, permineralized midgut glands. An alternative relationship betweenAustralimicolaand the Early Ordovician–Early Devonian Cheloniellida explains the shared anterior flexure of trunk pleurae but forces dubious homologies in other characters, such as dorsally-articulated furcae versus spines.

Peduncular attached secondary tiering acrotretoid brachiopods from the Chengjiang fauna: Implications for the ecological expansion of brachiopods during the Cambrian explosion

Brachiopods are usually thought to contribute little to the tiering complexity from the Paleozoic to the Recent, mainly due to the fact that their recent representatives live primarily in lower tiers directly above or below the water–sediment interface. Here we present the first and oldest record of varied levels of secondary tiering in minute brachiopods attached by exceptionally preserved thread-like pedicles around the branched fronds of the algae-like Malongitubus kuangshanensis Hu, 2005. The specimens illustrated herein were recovered from the Chengjiang fauna (Series 2, Stage 3) in the Lower Cambrian Heilinpu Formation at the Kuangshan section in Malong County, Yunnan Province, southern China. The micro-morphology and oval outline of the attached brachiopods demonstrate that they can be assigned to acrotretoid brachiopods (Linguliformea, Lingulata, Acrotretoidea), described here as Kuangshanotreta malungensis gen. et sp. nov. This is the first report on the occurrence of acrotretoid brachiopods in the Lower Cambrian muddy deposits from southern China. The posterior margins of the Kuangshanotreta shells are invariably either in direct contact with, or directed towards, and then in situ attached to the algal frond of M. kuangshanensis, indicating a secondary tiering in the ecological structure of Early Cambrian brachiopods. The acrotretoid Kuangshanotreta/algae malongitubus association represents both the first and oldest evidence into the enigmatic paleoecology of the diverse acrotretoid linguliformean stock that comprises an important component of the Cambrian evolutionary fauna, and sheds light on medium-high levels of secondary tiering (+5 to +10cm) Cambrian soft substrate suspension-feeding communities. When compared to other Chengjiang brachiopods, the miniature morphology and concomitant weight reduction of the shell of K. malungensis may be a good adaption to a suspended epifaunal tiering life style. We infer that the diversification of micromorphic and miniaturized acrotretoids that occurred from the Middle Cambrian to Early Ordovician may be the result of the increased availability of habitable surfaces provided by high levels of tiering in new types of ecological spaces.

Soft Anatomy of the Early Cambrian ArthropodIsoxys curvirostratusfrom the Chengjiang Biota of South China with a Discussion on the Origination of Great Appendages

An updated reconstruction of the body plan, functional morphology and lifestyle of the arthropod Isoxys curvirostratus is proposed, based on new fossil specimens with preserved soft anatomy found in several localities of the Lower Cambrian Chengjiang Lagerstatte. The animal was 2–4 cm long and mostly encased in a single carapace which is folded dorsally without an articulated hinge. The attachment of the body to the exoskeleton was probably cephalic and apparently lacked any well-developed adductor muscle system. Large stalked eyes with the eye sphere consisting of two layers (as corneal and rhabdomeric structures) protrude beyond the anterior margin of the carapace. This feature, together with a pair of frontal appendages with five podomeres that each bear a stout spiny outgrowth, suggests it was raptorial. The following 14 pairs of limbs are biramous and uniform in shape. The slim endopod is composed of more than 7 podomeres without terminal claw and the paddle shaped exopod is fringed with at least 17 ...

Skimming the surface with Burgess Shale arthropod locomotion

The first arthropod trackways are described from the Middle Cambrian Burgess Shale Formation of Canada. Trace fossils, including trackways, provide a rich source of biological and ecological information, including direct evidence of behaviour not commonly available from body fossils alone. The discovery of large arthropod trackways is unique for Burgess Shale-type deposits. Trackway dimensions and the requisite number of limbs are matched with the body plan of a tegopeltid arthropod. Tegopelte, one of the rarest Burgess Shale animals, is over twice the size of all other benthic arthropods known from this locality, and only its sister taxon, Saperion, from the Lower Cambrian Chengjiang biota of China, approaches a similar size. Biomechanical trackway analysis demonstrates that tegopeltids were capable of rapidly skimming across the seafloor and, in conjunction with the identification of gut diverticulae in Tegopelte, supports previous hypotheses on the locomotory capabilities and carnivorous mode of life of such arthropods. The trackways occur in the oldest part (Kicking Horse Shale Member) of the Burgess Shale Formation, which is also known for its scarce assemblage of soft-bodied organisms, and indicate at least intermittent oxygenated bottom waters and low sedimentation rates.

Redox changes in Early Cambrian black shales at Xiaotan section, Yunnan Province, South China

The Xiaotan section, situated on the southwestern Yangtze Platform, South China, contains an unusually expanded sedimentary record, spanning the upper Neoproterozoic to the Cambrian and representing more than 40My of marine sedimentation. Three distinct black shale units of the Shiyantou and Yuanshan formations were analysed for redox-sensitive trace elements and iron speciation, in order to constrain the evolution of the marine environment during the Early Cambrian metazoan biodiversification on the southwestern Yangtze Platform. For comparison we also include black shale samples of the Shiyantou Fm. from the Meishucun section, samples from a previous study covering the Shiyantou and Yuanshan Fm. at Dapotuo near Chengjiang, and samples from the strongly condensed Early Cambrian Niutitang Fm. from the Zhongnan section situated on the platform margin. Th/U and FeHR/FeT ratios reveal that bottom waters on the southwestern Yangtze Platform were commonly anoxic and ferruginous into the Cambrian, but became oxic after deposition of the black shales of the lower Yuanshan Formation, which elsewhere contains the Chengjiang Biota. Average Mo/TOC ratios, used as an indicator of the size of the Mo reservoir, increase up-section, and together with generally increasing S/C ratios suggest a trend towards normal (Phanerozoic) marine conditions. A distinctive Ni peak within the upper Shiyantou Fm. at Xiaotan and maximum Mo concentrations at the base of the Yuanshan Fm. can be correlated to the Ni–Mo sulphide layer within the lower Niutitang Fm. and thus indicate two separate pulses of sedimentary-exhalative nickel enrichment and subsequent, redox controlled molybdenum accumulation. A strong decoupling of organic carbon and pyrite sulphur within both sections further suggests upwelling of H2S-rich waters onto the platform. We argue here for an opening of the depositional environment on the northwestern Yangtze Platform which led to the upwelling of sulphidic waters, ventilation of the seafloor and ultimately its colonisation by newly emergent animal groups during the ‘Cambrian Explosion’.

Lobopodian phylogeny reanalysed

Arising from J. Liu et al. , 526–530 (2011)10.1038/nature09704 ; Liu et al. reply Liu et al.1 described an ‘armoured’ lobopodian, Diania cactiformis, from the Chengjiang Lagerstatte (China; Cambrian, stage 3); this fossil bears potentially arthropod-like articulated and possibly sclerotized appendages, but lacks a sclerotized body. A cladistic analysis resolved Diania as sister-taxon to arthropods. From this phylogenetic position the authors tentatively inferred that arthropodization (sclerotization of limbs) may have preceded arthrodization (sclerotization of body elements) in arthropod evolution. Although we concur with the reasoning behind this inference, it rests on a phylogenetic placement that our analysis of the published data set does not reproduce.

New Bivalved Arthropods from Mid‐Cambrian Kaili Biota of Southeastern Guizhou, Southwest China

Abstract:Bivalved arthropods are very important elements in Mid‐Cambrian Kaili Biota. Three genera, Alicaris, Pseudoarctolepis, and Forfexicaris, are herein recorded for the first time, including three new species: Alicaris kailiensis sp. nov., Forfexicaris reticulata sp. nov., and Pseudoarctolepis semicircularis sp. nov., of which Alicaris kailiensis sp. nov. could be assigned to crustaceomorphs; the other two forms are grouped as proschizoramians. These forms provide important information about the distribution and species diversity of a poorly‐known element of Cambrian fauna, and also bridge the biotic evolutionary gap between Early Cambrian Chengjiang Biota in Southwest China and Mid‐Cambrian burgess shale biota in North America.

A rare onychophoran-like lobopodian from the Lower Cambrian Chengjiang Lagerstätte, southwestern China, and its phylogenetic implications

Lobopodians, which diversified and flourished in the Cambrian seas, have long drawn much attention in that not only their extant close relatives, onychophorans and tardigrades, but euarthropods (Chelicerata, Myriapoda, Crustacea, and Hexapoda) may have been deeply rooted in stem-group lobopodians. Antennacanthopodia gracilis new genus and species is described and interpreted here as an “unarmoured” lobopodian from the Chengjiang fossil Lagerstätte (Early Cambrian, —520 Ma), Yunnan, southwestern China. This animal shares with other known Cambrian lobopodians such plesiomorphies (primitive characters) as onychophoran-like overall appearance; a metamerically segmented body covered by slightly sclerotized cuticle, and paired, unjointed lobopodal legs. Antennacanthopodia is also featured by a pair of frontal antennae, potential ocellus-like lateral visual organs, second antennae, a straight, voluminous midgut, diminutive spines arrayed on the leg and the trunk, well-developed leg musculature, highly sclerotized terminal leg pads, and presumptively a pair of posteriormost appendicules. This new taxon, with innovative characters (autapomorphies), furthers our understanding of early lobopodian diversification. Antennacanthopodia is considered closely allied to extant Onychophora based on considerable anatomical similarities. Taken together its “two-segmented” cephalization and appendage-bearing “ocular segment”, this new form may shed some new light on the arthropod groundplan.

A new arthropodJugatacaris agilisn. gen. n. sp. from the Early Cambrian Chengjiang biota, South China

A new arthropodJugatacaris agilisn. gen. n. sp. with excellent soft anatomy is reported from the Early Cambrian Chengjiang biota. Its “bivalved” carapace with a dorsal fin-like fusion is distinct from those of other Cambrian arthropods. A pair of stalked eyes and a prominent median eye protrudes the carapace anteriorly. The cephalon, attaching to the carapace through the lateral adductor muscles at maxillary segment, bears an antennule, antenna and mandible. The trunk is comprised of a large number of segments (varying from 55 to 65), two-thirds of which is covered by the carapace. Each segment, except for the posterior three to five, carries a pair of uniform biramous appendages. The endopodite is composed of 30 podomeres and a terminal claw. The oar-shaped exopododite is fringed with filaments and distally bears a broad setiferous lobe. Trunk terminates with a conical telson and a pair of broad furcal rami serving as steering devices. Internal features like gonad and gut have also been found. The presence of the food groove combined with other morphological features indicates thatJugatacariswas a filter feeder. The appendages design associated with the overall body-plan supports the view thatJugatacarisis a crustaceanomorph. The accurate phylogenic assessment will remain the subject of debate until more information becomes available.

An obolellate brachiopod with soft-part preservation from the Early Cambrian Chengjiang fauna of China

The obolellates represent a poorly understood group of the oldest known cosmopolitan calcareous rhynchonelliform brachiopods. They made their first appearance in the early Atdabanian and became extinct at the end of the Middle Cambrian. Consequently, any information concerning the soft anatomy of this ephemeral lineage of brachiopods has great phylogenetic significance. This is the first report on two specimens of an obolellate with fine preservation of soft parts including the pedicle, marginal setae and possible imprints of a spiral lophophore, recovered from the early Cambrian Chengjiang Konservat Lagerstätte of Kunming, southern China. The setae are thin and densely fringed along the shell margin. The stout pedicle is distinctly composed of densely stacked tabular bodies, lacking a central coelomic lumen; it emerges through a possible foramen, and slightly tapers posteriorly with the distal end attached to exoskeletons of other organisms.

A venomous arthropod in the Early Cambrian Sea

A number of recently evolved animals possess poison glands for feeding and/or defense. However, examples of such animals are rare in the fossil record. We report a fossil arthropod Isoxys curvirostratus from the Early Cambrian Chengjiang biota of China. This species is regarded as the oldest known venomous arthropod based on the presence of venomous glands in its head region. The adult animal is 2–5 cm long and the body is covered entirely with a carapace. The presence of large stalk eyes and a pair of stout grasping appendages with a terminal spine suggest it was raptorial. Interestingly, the two pear-shaped, three-dimensionally preserved objects that are present in the head region and at the base of the grasping appendages closely resemble the venom glands of some living arthropods in size, shape, and position. These features indicate that the presence of venomous predators could date back 520 million years. Furthermore, our observations suggest that the feeding strategies and organs adapted for this purpose had already reached a high level of diversity and anatomical sophistication in the Early Cambrian ecosystems.

An armoured Cambrian lobopodian from China with arthropod-like appendages

Cambrian fossil Lagerstätten preserving soft-bodied organisms have contributed much towards our understanding of metazoan origins. Lobopodians are a particularly interesting group that diversified and flourished in the Cambrian seas. Resembling 'worms with legs', they have long attracted much attention in that they may have given rise to both Onychophora (velvet worms) and Tardigrada (water bears), as well as to arthropods in general. Here we describe Diania cactiformis gen. et sp. nov. as an 'armoured' lobopodian from the Chengjiang fossil Lagerstätte (Cambrian Stage 3), Yunnan, southwestern China. Although sharing features with other typical lobopodians, it is remarkable for possessing robust and probably sclerotized appendages, with what appear to be articulated elements. In terms of limb morphology it is therefore closer to the arthropod condition, to our knowledge, than any lobopodian recorded until now. Phylogenetic analysis recovers it in a derived position, close to Arthropoda; thus, it seems to belong to a grade of organization close to the point of becoming a true arthropod. Further, D. cactiformis could imply that arthropodization (sclerotization of the limbs) preceded arthrodization (sclerotization of the body). Comparing our fossils with other lobopodian appendage morphologies--see Kerygmachela, Jianshanopodia and Megadictyon--reinforces the hypothesis that the group as a whole is paraphyletic, with different taxa expressing different grades of arthropodization.

Relation between anatomy and lifestyles in Recent and Early Cambrian chaetognaths

The Burgess Shale-type Lagerstätten of the Early Cambrian Maotianshan Shale record an apparently sudden conquest of pelagic niches by ten phyla of metazoans. One of these phyla is Chaetognatha, a group of predatory marine worms. Given their role as major predators in modern planktonic ecosystems, the chaetognaths discovered in the Maotianshan Shale (Yunnan Province, South China) suggest that the pelagos at the time was already quite complex. Modern chaetognaths, however, can be divided into benthic and pelagic forms; the pelagic nature ofEognathacantha ercainellashould therefore be validated by strong morphological evidence.Knowing that planktonic animals present morphological adaptations that increase their buoyancy, we studied the drag produced during the active phase of chaetognath locomotion for the modern formsParaspadella gotoi(benthic) andSagitta elegans(pelagic). By using a motion model developed by Jordan in 1992, we could calculate the resistive force produced by the undulatory movement of chaetognaths' bodies.This mechanistic approach evaluates the effect of three motion parameters (relative speed, total length, and tail ratio) on the drag force produced during locomotion. Our results show that the increase of size contributes to higher drag while the shorter tail of the pelagic form balances this effect by reducing the wetted surface subject to friction. For chaetognaths, therefore, a bigger body (both in length and width) and a shorter tail indicate a pelagic lifestyle, a finding that can be applied to the study of the fossilEognathacantha ercainella. A discriminant analysis can confirm that the Early Cambrian chaetognath presents a pelagic morphology with similarities to modern bathypelagic and mesopelagic species.

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.

Phylogeny and evolutionary significance of vermiform animals from the Early Cambrian Chengjiang Lagerstätte

The Early Cambrian Chengjiang Biota of China has preserved many species of vermiform animals in exquisite detail, which provide a unique window into the origin and early evolution of different vermiform phyla. This article reviews and discusses the forty-five Chengjiang vermiform animals discovered so far, and specifically comments on the phylogeny, modes of life, evolutionary significance as well as existing debates concerning the Chengjiang priapulid and lobopodian phyla. This article analyzes existing problems with Chengjiang vermiform morphological research and highlights various factors that affect the final preserved fossil morphology, including taphonomy and shape change caused by vermiform movement, as well as proposing that Anningvermis multispinosa and Corynetis brevis are synonymous. Chengjiang vermiform animals displayed a great diversity of morphologies and ecological roles, which is significant for understanding the origin and evolution of early metazoans; the potential for future research is enormous but needs to be broadened by interdisciplinary work involving taphonomy, geochemistry, cladistics, and genetics.