The genus and species Yuyuanozoon magnificissimi gen. et sp. nov., a new fossil vetulicolian, is reported from the Early Cambrian Chengjiang Fauna in Yunnan of China. It has a bipartite body plan and five pairs of cha...The genus and species Yuyuanozoon magnificissimi gen. et sp. nov., a new fossil vetulicolian, is reported from the Early Cambrian Chengjiang Fauna in Yunnan of China. It has a bipartite body plan and five pairs of chain-formed gill sacs, showing the general characteristics of Ventulicolia. However, the exclusive 'atrial cavity' and external gill observed indicate that the new form is different from those previously described as vetulicolians, probably representing a lineage developed within deuterostomes and more primitive than urochordates.展开更多
The paper deals with a new species of megaalgal fossil-Enteromophites intestinalis sp. nov. found in Chenejiang Biota in the early Cambrian. A morphological comparison of the living Enteromorpha and fossil Enteromophi...The paper deals with a new species of megaalgal fossil-Enteromophites intestinalis sp. nov. found in Chenejiang Biota in the early Cambrian. A morphological comparison of the living Enteromorpha and fossil Enteromophites indicates that there probably is a close relationship between these two genera. Based on a study on the surroundings of living Enteromorpha, Enteromophites intestinalis sp. nov. might have lived in the environment of sea or salt water.展开更多
The Lower Cambrian Chengjiang Lagerstatte provides a good window to explorethe origin and radiation of early bilaterians. Here we describe a netted sclerite-bearing wormTabelliscolex hexagonus gen. et sp. nov., and te...The Lower Cambrian Chengjiang Lagerstatte provides a good window to explorethe origin and radiation of early bilaterians. Here we describe a netted sclerite-bearing wormTabelliscolex hexagonus gen. et sp. nov., and tentatively assign it to palaeoscolecidans ofpriapulids. The cuticle of the animal is covered with two kinds of platy sclerites which areconstructed by hexagonally arranged tubercles Similar structures of the sclerites can be seen onsome Cambrian palaeoscolecidans and lobopods, so, this new species is critical for understanding therelationships between lobopods and palaeoscolecidans.展开更多
The bivalved arthropod Branchiocaris? yunnanensis Hou, 1987 is redescribed on the basis of new fossil material from the early Cambrian Chengjiang biota, South China. In total, 81 specimens have been examined. The car...The bivalved arthropod Branchiocaris? yunnanensis Hou, 1987 is redescribed on the basis of new fossil material from the early Cambrian Chengjiang biota, South China. In total, 81 specimens have been examined. The carapace comprises two identical valves, each valve are sub-circular in outline, ranging from 24 mm to 58 mm in length, and from 15 mm to 46 mm in height. The dorsal margin is straight and bears two short cardinal spines. The valve surface is either smooth or ornamented with polygonal reticular structures, which may result from different preservation conditions. Two morphotypes have been recognized based on the presence or absence of a subdorsal swelling of the valve, which is convex dorsolaterally and extends beyond the dorsal margin when it appears. Statistic and ontogenetic analyses show both morphotypes grow isometrically and have the same growth trajectory. Therefore, the two morphotypes are interpreted as dimorphism within the same species. Our results are helpful for investigating the taxonomy and ontogeny of arthropod fossils and suggest that dimorphism might be fairly common in these early arthropods.展开更多
This paper gives a brief report on the new occurrence of the Early CambrianQiongzhusi' an Chengjiang fauna in the Haikou area, Kunming and its assemblage feature andthe stratigraphic section of the horizon yieldin...This paper gives a brief report on the new occurrence of the Early CambrianQiongzhusi' an Chengjiang fauna in the Haikou area, Kunming and its assemblage feature andthe stratigraphic section of the horizon yielding these fossils. Eight genera and 8 species ofTrilobitoidea are described here, embracing Leanchoilia asiatica Luo et Hu (sp. nov.), Yohoiasinensis Luo et Hu (sp. nov.), Zhongxinia speciosa Luo et Hu (gen. et sp. nov.), Xandarellaspectuculum Hou et al., Dianchia mirabilis Luo et Hu (gen. et sp. nov.), Kuamaia lata Hou,Retifacies longispinus Luo et Hu (sp. nov.) and Sinoburius lunaris Hou et al.展开更多
Well known for its abundant and extraordinary soft-tissue fossils,the Chengjiang fauna has witnessed the main phase of the Cambrian explosion and the first great congress of the ancestors of nearly all major phyla of ...Well known for its abundant and extraordinary soft-tissue fossils,the Chengjiang fauna has witnessed the main phase of the Cambrian explosion and the first great congress of the ancestors of nearly all major phyla of animals on Earth.The large-scale survey and exploration of the fauna by Chinese paleontologists,partly in collaboration with international scientists,over the past 30 years can be broadly divided into three stages.In the first decade since 1984(19841994),a large number of invertebrates,including basal animals and the protostomes of the early animal tree were discovered,but the subkingdom Deuterostomia was completely unknown.Then in the second decade(19952005),the great discoveries of various deuterostomes led to the construction of the basic framework of the Deuterostomia and then to the formation of the tripartite phylogenetic trees of early animals(TPTEA,including basal animals,protostomes and deuterostomes),for the first time.In the third decade(after 2005),the academic community began to think about the internal relationships between the formation of TPTEA and the multiepisode Cambrian explosion,leading to the new hypothesis of the three-episode Cambrian explosion.The Chengjiang fauna is important for deciphering the fauna structure,paleoecological environment,and so on.However,its core academic values mainly rest on two aspects.Firstly,The Chengjiang fauna,as the main phase witness of the Cambrian explosion,has created a nearly complete phylogenetic framework of the TPTEA on Earth for the first time.The three-phase radiation hypothesis reveals the essential connotation of the Cambrian explosion:a step-wise divergent evolution of animals,from basal to highly advanced groups,lasting about 40 million years.In the first phase,it gave birth to a bulk of basal animals(including some now extinct“animal”groups)in the latest Ediacaran,probably including some pioneer protostomes.The second phase took place in the first epoch of the Cambrian period(Terreneuvian),giving rise to the main invertebrate protostomes with a persistent prosperity of basal animals.The third phase proceeded in Cambrian Epoch 2(represented by the Chengjiang fauna),which not only maintained the prosperity of basal animals and protostomes,but also,more importantly,gave birth to all the main phyla of the subkingdom Deuterostomia.Thus,the rudimental framework of the whole TPTEA has been shaped,with the termination of the major innovation events of the Cambrian explosion.Here,we discussed the evolutionary properties of Ediacaran biota,small shelly fossils and the Chengjiang fauna in the Cambrian explosion with emphasis on the biological properties of several important animal groups.The order Myllokunmingiida is the only known oldest vertebrate,while Yunnanozoon and Haikouella are neither vertebrates nor stem-group chordates but a special group of basal deuterostomes;Cheungkongella is a credible ancestor of the urochordate and it supports the classical hypothesis on the origin of the urochordates;and the gill slits were first invented in the members of the phylum Vetulicolia to provide key information on the origin of the deuterostomes.The second core value of the Chengjiang fauna is of profound humanistic and philosophical significance:the discoveries of the‘first gill openings’,‘first brain’,‘first vertebrae’and‘first heart’provide the pivotal evidence for solving the unsolved mystery of the origin of the main basic human organs as described in Darwin’s“The Descent of Man”.In addition,the morphological and anatomical information of the Chengjiang fauna can provide important clues for a better understanding of most components of Ediacaran and Cambrian metazoans.展开更多
Over one hundred arthropod fossil species have been described from the famous Chengjiang Lagerst?tte(South China, Cambrian Stage 3, ca. 518 Ma) including a diverse assemblage of radiodonts–a group containing Anomaloc...Over one hundred arthropod fossil species have been described from the famous Chengjiang Lagerst?tte(South China, Cambrian Stage 3, ca. 518 Ma) including a diverse assemblage of radiodonts–a group containing Anomalocaris and its relatives. These iconic stem-group euarthropods include some of the largest animals of the time, and some are known from hundreds of specimens. A longstanding conundrum has been the rarity or absence of hurdiids from Cambrian Series 2 Lagerst?tten like Chengjiang. This is because radiodonts are generally common in such deposits and the oldest radiodont ever discovered is a hurdiid. Furthermore, this family displays the widest geographic and temporal ranges of all radiodont families, and the highest diversity. Here we document the first hurdiid frontal appendages from Chengjiang, which display unique features within the family and may provide insights for understanding the character evolution of hurdiid appendages. The palaeoenvironmental distribution of hurdiids suggests that the rarity of hurdiids in Chengjiang may be due to a preference for deeper water environments,and the later success of this family from the Wuliuan onwards may relate to their ability to tolerate cooler water temperatures than other radiodont families. The palaeogeographical, palaeoenvironmental, and stratigraphical patterns observed in hurdiids maybe caused in part by the limited distributions of Konservat-Lagerst?tten in the Cambrian as well.展开更多
Ancestral lobopodians,which diversified and flourished in the Cambrian seas,have long drawn much attention in that not only their living counterparts,onychophorans and tardigrades,but euarthropods(Chelicerata,Myriapod...Ancestral lobopodians,which diversified and flourished in the Cambrian seas,have long drawn much attention in that not only their living counterparts,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展开更多
Burgess Shale-type deposits provide a wealth of information on the early evolution of animals.Questions that are central to understanding the exceptional preservation of these biotas and the paleoenvironments they inh...Burgess Shale-type deposits provide a wealth of information on the early evolution of animals.Questions that are central to understanding the exceptional preservation of these biotas and the paleoenvironments they inhabited may be obscured by the post-depositional alteration due to metamorphism at depth and weathering near the Earth’s surface.Among over 50 Cambrian BST biotas,the Chengjiang and Qingjiang deposits are well known for their richness of soft-bodied taxa,fidelity of preservation,and Early Cambrian Age.While alteration via weathering has been well-investigated,the thermal maturity of the units bearing the two biotas has not yet been elucidated.Here we investigate peak metamorphic temperatures of the two deposits using two independent methods.Paleogeotemperature gradient analyses demonstrate that the most fossiliferous sections of the Chengjiang were buried at a maximum depth of∼8500 m in the Early Triassic,corresponding to∼300°C,while the type area of the Qingjiang biota was buried at a maximum depth of∼8700 m in the Early Jurassic,corresponding to∼240°C.Raman geothermometer analyses of fossil carbonaceous material demonstrate that peak temperatures varied across localities with different burial depth.The two productive sections of the Chengjiang biota were thermally altered at a peak temperature of approximately 300°C,and the main locality of the Qingjiang biota experienced a peak temperature of 238±22°C.These results from two independent methods are concordant.Among BST deposits for which thermal maturity has been documented,the Qingjiang biota is the least thermally mature,and therefore holds promise for enriching our understanding of BST deposits.展开更多
The Cambrian Explosion is real Fossil evidence from Chengjiang reveals an assemblage of the fauna, which is represented by at least 100 species. They belong to animal phyla that are still extant, including sponges, cn...The Cambrian Explosion is real Fossil evidence from Chengjiang reveals an assemblage of the fauna, which is represented by at least 100 species. They belong to animal phyla that are still extant, including sponges, cnidarians, ctenophorans, priapulids, annelids(?), arthropods,brachiopods, phoronids, echinoderms and chordates. There are even a number of animal body plans at phylum or subphylum level that are extinct, including: medusiform Eldonia, Facivermis, Tardipolyda, chanccelariids, Dinomischus, hyoliths, Vetuliculid,展开更多
Twenty black shale samples, which are free from the influence of weathering, were collected from the Chengjiang Fauna-bearing horizon, central Yunnan Province, yielding a Pb-Pb isochron age of 534±60 Ma. Although...Twenty black shale samples, which are free from the influence of weathering, were collected from the Chengjiang Fauna-bearing horizon, central Yunnan Province, yielding a Pb-Pb isochron age of 534±60 Ma. Although this age is younger than both the Rb-Sr isochron age and 40Ar-39Ar age, it should represent the lower isotopic age limit of the Chengjiang Fauna.展开更多
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 composi...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.展开更多
The Chengjiang Formation is the earliest continental clastic deposit after the Jinning Orogeny in central Yunnan Province,and therefore its accurate depositional age is significant for understanding the formation and ...The Chengjiang Formation is the earliest continental clastic deposit after the Jinning Orogeny in central Yunnan Province,and therefore its accurate depositional age is significant for understanding the formation and evolution of the Neoproterozoic rift basins in southern China.However,hampered by accuracy of the dating technique,the existing age data for the Chengjiang Formation are not very reliable.A large number of magmatic zircons were obtained from the tuff interbed in the lower part of the Chengjiang Formation in Jinyang area and the bottom part of the Chengjiang Formation in Dongchuan area,central Yunnan Province,and high-precision SHRIMP U-Pb dating was carried out on these zircons.The results show that the weighted mean 206 Pb/238 U ages are 797.8±8.2 and 803.1±8.7 Ma respectively.In combination with related geological evidence,it has been demonstrated that the bottom boundary age of the Chengjiang Formation should be 800±5 Ma,whereas the top boundary age could be ca 725 Ma.By synthetically analyzing the latest age data for the relevant strata,it has been confirmed that the Chengjiang Formation should be correlated with the Suxiong Formation and the Kaijianqiao Formation in western Sichuan Province,the Liantuo Formation in the middle and lower Yangtze,the Hongchicun Formation and the Shangshu Formation in northern Zhejiang Province,the Puling Formation in southern Anhui Province,the Luokedong Formation and the Majianqiao Formation in northwestern Jiangxi Province,the Wuqiangxi Formation in northwestern Hunan Province,the Fanzhao Formation and the Qingshuijiang Formation in southeastern Guizhou Province,and the Sanmenjie Formation and the Gongdong Formation in northern Guangxi Province.Sedimentary cycle analysis shows that the sedimentary filling sequence of the Neoproterozoic rift basins in southern China can be divided into four cycles.Among them,Cycle II began at ca.800 Ma,accompanied by intensive tectonic-thermal events.The zircon U-Pb ages from the bottom of the Chengjiang Formation reported in this paper indicate that the Neoproterozoic Kangdian rift subbasin probably started to develop at ca.800 Ma and therefore missed Cycle I of the Neoproterozoic sedimentary filling sequence in southern China.展开更多
The taxonomy of an early ancestor of Recent Priapulidae, Xiaoheiqingella peculiaris (= Yunnanpriapulus halteroformis Huang et al., 2004) from the Early Cambrian Chengjiang fossil Lagerst?tte, is revised. Morphological...The taxonomy of an early ancestor of Recent Priapulidae, Xiaoheiqingella peculiaris (= Yunnanpriapulus halteroformis Huang et al., 2004) from the Early Cambrian Chengjiang fossil Lagerst?tte, is revised. Morphological characters comprise a pair of caudal appendages rather than a single appendage flanking the trunk end and a possible urogenital duct found inside the preanal region. An addi- tional extremely rare fossil priapulid worm, Paratubiluchus bicaudatus gen. nov., sp. nov. is also described herein. Its diagnostic characters are: an introvert bearing 25 longitudi- nal rows of scalids, a distinct neck region, no annulus on the oval trunk, and a pair of caudal appendages. The proportion of body parts is similar in size to that of loricate larvae of Recent priapulids and larva-formed Palaeopriapulitidae. Taking account of the features of Xiaoheiqingella, bicaudal appendages are considered to be a synapomorphy of Pri- apulidae and Tubiluchidae. Paratubiluchus gen. nov. is most likely a candidate for the ancestor of the Tubiluchidae; it probably originated from a larva-formed priapulid with 25 rows of scalids, thus representing an intermediate link be- tween the priapulids in mature-form and the priapulids with lorica.展开更多
The Cambrian explosion was coined to describe the geologically sudden appearance of numerous bilaterian body plans(Phyla)around the Ediacaran-Cambrian transition,around 565-520million years ago.Many explanations and c...The Cambrian explosion was coined to describe the geologically sudden appearance of numerous bilaterian body plans(Phyla)around the Ediacaran-Cambrian transition,around 565-520million years ago.Many explanations and conjectures have been postulated in order to explain the pattern and duration of this explosive radiation of many different phyla of early metazoans.Here,we focus on the evolution of a phylum of marine suspension-feeding animals—the brachiopods,as exemplified by the exceptionally preserved taxa from the celebrated Chengjiang Konservat Lagerst(a|¨)tte(Yunnan,China).The abundant soft-bodied preservation at these fossil quarries gives us the only firm insights into what brachiopods looked like and how they functioned and lived when they first appeared on the Earth.Studies of Chengjiang brachiopods demonstrate that the early animals developed a remarkably varied organization of tissues and organs shortly after the onset of Cambrian explosion.In the marine suspension-feeding brachiopods,most importantly the tentaculate feeding structure of early brachiopods is already differentiated into two shapes of lophophore,anteriorly coiled(spiralled)and posteriorly arching tentacle crowns and the unique latter type was previously not documented from fossil and living brachiopods.Also unlike any known Recent brachiopod,all the known Cambrian brachiopods from Chengjiang have an open digestive tract that was disposed either as a Ushaped gut in linguliform and stem group brachiopods,or straight gut with a posterior anus in some calcareous-shelled stocks.Moreover,in contrast toliving lingulids,all the Cambrian brachiopods have an epibenthic lifestyle either cemented by a ventral valve or attached by variable pedicles to establish complex ecological community encompassing primary tierers and variable secondary tierers.It is therefore assumed that brachiopods were the first benthic metazoan that achieved their success in ecological stratification and tiering complexity by late Atdabanian.The setae are also important for the brachiopod suspension-feeding life style,and in the Chengjiang braehiopods they include two types cilia-like and spine-like setae.The mantle canals of different braehiopod species are also distinctly variable in arrangement,mainly disposed in pinnate,baeulate and peripheral conditions.Of these,the peripheral disposition of mantle canals is for the first time proposed here so as to differentiate from the bifurcate condition in recent lingulids in that the former is devoid of posteriorly extending main trunks of sinus,but possesses a diverging dorsal vascula media in dichotomy.展开更多
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 arti...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.展开更多
The origin of arthropods has long been one ofthe most hotly-debated subjects. Arthropods used to bethought closely related with annelids, but the two groups arenow believed to be separated into two major realms within...The origin of arthropods has long been one ofthe most hotly-debated subjects. Arthropods used to bethought closely related with annelids, but the two groups arenow believed to be separated into two major realms withinProtostomia on the basis of new molecular data. Although itis generally held by paleontologists that arthropods should berooted in the early lobopods-.a kind of worm-like creaturewith non-segmented legs, no intermediate forms have beenfound to bridge them. Here we report an organism with amixture of characters, including features characteristic ofarthropods (e.g., primary cephalization with paired eyes,paired antennae, and preliminary tagmosis) and of lobopods(e.g., worm-like body design, the dorsal spines, andnon-segmented limbs or lobe-like legs). The discovery of therare transitional form may throw new light on the origin ofarthropods and suggests that the most primitive arthropodsbegan with paired uniramous legs and the biramous onesevolved later.展开更多
A new Chengjiang-type fossil assemblage is reported herein from the lower part of the Hongjingshao Formation at Xiazhuang village of Chenggong,Kunming,Yunnan.The fossil assemblage,named as Xiazhuang fossil assemblage,...A new Chengjiang-type fossil assemblage is reported herein from the lower part of the Hongjingshao Formation at Xiazhuang village of Chenggong,Kunming,Yunnan.The fossil assemblage,named as Xiazhuang fossil assemblage,yields predominantly soft-bodied fossils,including arthropods,brachiopods,priapulids,lobopods and some problematic taxa,with arthropods being the most dominant group.Preservation and composition of the fossil assemblage are very similar to the typical Chengjiang biota,which is preserved in the middle Yu’anshan Formation in the large area of eastern Yunnan.The associated trilobites demonstrate that the soft-bodied fossil assemblage belongs to the late Qiongzhusian in age(Stage 3,Cambrian),suggesting that the Hongjingshao Formation is probably a diachronous lithostratigraphic unit ranging from the upper Qiongzhusian to the lower Canglangpuan stages in eastern Yunnan.The fossil assemblage from the Xiazhuang area fills up the missing link between the typical older Chengjiang biota and the younger Malong and Guanshan biotas,making eastern Yunnan a unique area in the world to reveal the early evolutionary history of animals and palaeocommunity dynamics during the‘‘Cambrian explosion’’.展开更多
The efforts of labor-intensive collecting in the Early Cambrian Chengjiang deposits in eastern Yunnan Province, China led to the discovery of many horizons containing exceptionally well preserved soft-bodied fossil co...The efforts of labor-intensive collecting in the Early Cambrian Chengjiang deposits in eastern Yunnan Province, China led to the discovery of many horizons containing exceptionally well preserved soft-bodied fossil concentrations, many of which can be assigned to either monospecific concentrations or paucispecific concentrations. The features of these fossil concentrations support the hypothesis that frequent storm events producing tempestites mainly contributed to the preservation of abundant soft- bodied fossils in the Chengjiang deposits, and indi- cate that the balance of the ecological web in this region was probably frequently destroyed or upset by such storm events during that geological time. Ani- mals in a fossil assemblage in such fossil concentra- tions probably occupied similar ecological biotopes.展开更多
Facivermis yunnanicus (Hou & Chen, 1989), from the Early Cambrian Chengjiang Lager- st?tte, a worm-like fossil with 5 pairs of tentacles and a perceived shrunken end, has been regarded as related to polychaetes, l...Facivermis yunnanicus (Hou & Chen, 1989), from the Early Cambrian Chengjiang Lager- st?tte, a worm-like fossil with 5 pairs of tentacles and a perceived shrunken end, has been regarded as related to polychaetes, later it has been variously interpreted as lobopods, Pentastoma and lopho- phorates. Newly discovered complete specimens by the ELI field team show that the taxon has, in addition to the 5 pairs of appendages, a pear-shaped trunk end bearing two or three circles of hooks. Accordingly, based on these important morphological characters, reconsideration of its affinities is provided and the taxonomy is remedied herein. Because the five pairs of appendages of Facivermis yunnanicus resemble the appendages of the fore-trunk of lobopod Mi- raluolishania (Liu & Shu, 2004), it seems that they are homologous structures. Therefore, the affinities of Facivermis are regarded here as being close to the lobopods, in addition, the lobe-like appendages of Facivermis are very crucial to exploring the origin of the appendages of lobopods and arthropods.展开更多
基金supported by the National Natural Science Foundation of China(Grants 40073007,40173002,40232020)the Natural Scienee Foundation of Yunnan Province(Grant 97D007Z)
文摘The genus and species Yuyuanozoon magnificissimi gen. et sp. nov., a new fossil vetulicolian, is reported from the Early Cambrian Chengjiang Fauna in Yunnan of China. It has a bipartite body plan and five pairs of chain-formed gill sacs, showing the general characteristics of Ventulicolia. However, the exclusive 'atrial cavity' and external gill observed indicate that the new form is different from those previously described as vetulicolians, probably representing a lineage developed within deuterostomes and more primitive than urochordates.
文摘The paper deals with a new species of megaalgal fossil-Enteromophites intestinalis sp. nov. found in Chenejiang Biota in the early Cambrian. A morphological comparison of the living Enteromorpha and fossil Enteromophites indicates that there probably is a close relationship between these two genera. Based on a study on the surroundings of living Enteromorpha, Enteromophites intestinalis sp. nov. might have lived in the environment of sea or salt water.
基金supported by the Natural Science Foundation of China(NSFC)the Ministry of Science and Technology(CMST)of China
文摘The Lower Cambrian Chengjiang Lagerstatte provides a good window to explorethe origin and radiation of early bilaterians. Here we describe a netted sclerite-bearing wormTabelliscolex hexagonus gen. et sp. nov., and tentatively assign it to palaeoscolecidans ofpriapulids. The cuticle of the animal is covered with two kinds of platy sclerites which areconstructed by hexagonally arranged tubercles Similar structures of the sclerites can be seen onsome Cambrian palaeoscolecidans and lobopods, so, this new species is critical for understanding therelationships between lobopods and palaeoscolecidans.
基金financed by the Major Basic Research Project of the Ministry of Science and Technology of China (Grants 2013CB837100, 2013CB806400)the Natural Science Foundation of China (Grants 41202007, 41272019 and 40830208)
文摘The bivalved arthropod Branchiocaris? yunnanensis Hou, 1987 is redescribed on the basis of new fossil material from the early Cambrian Chengjiang biota, South China. In total, 81 specimens have been examined. The carapace comprises two identical valves, each valve are sub-circular in outline, ranging from 24 mm to 58 mm in length, and from 15 mm to 46 mm in height. The dorsal margin is straight and bears two short cardinal spines. The valve surface is either smooth or ornamented with polygonal reticular structures, which may result from different preservation conditions. Two morphotypes have been recognized based on the presence or absence of a subdorsal swelling of the valve, which is convex dorsolaterally and extends beyond the dorsal margin when it appears. Statistic and ontogenetic analyses show both morphotypes grow isometrically and have the same growth trajectory. Therefore, the two morphotypes are interpreted as dimorphism within the same species. Our results are helpful for investigating the taxonomy and ontogeny of arthropod fossils and suggest that dimorphism might be fairly common in these early arthropods.
基金This work is a contribution to Project 92D087 supported by the Yunnan Provincial Fundation for Applied Basic Researches and Project 8502004-1A supported by the fund from the Ministry of Geology and Mineral Resources for important basic researches in the
文摘This paper gives a brief report on the new occurrence of the Early CambrianQiongzhusi' an Chengjiang fauna in the Haikou area, Kunming and its assemblage feature andthe stratigraphic section of the horizon yielding these fossils. Eight genera and 8 species ofTrilobitoidea are described here, embracing Leanchoilia asiatica Luo et Hu (sp. nov.), Yohoiasinensis Luo et Hu (sp. nov.), Zhongxinia speciosa Luo et Hu (gen. et sp. nov.), Xandarellaspectuculum Hou et al., Dianchia mirabilis Luo et Hu (gen. et sp. nov.), Kuamaia lata Hou,Retifacies longispinus Luo et Hu (sp. nov.) and Sinoburius lunaris Hou et al.
基金Natural Science Foundation of China(41672009,41621003,41772010,41720104002)The Strategic Priority Research Program of Chinese Academy of Sciences(XDB18000000,XDB26000000)。
文摘Well known for its abundant and extraordinary soft-tissue fossils,the Chengjiang fauna has witnessed the main phase of the Cambrian explosion and the first great congress of the ancestors of nearly all major phyla of animals on Earth.The large-scale survey and exploration of the fauna by Chinese paleontologists,partly in collaboration with international scientists,over the past 30 years can be broadly divided into three stages.In the first decade since 1984(19841994),a large number of invertebrates,including basal animals and the protostomes of the early animal tree were discovered,but the subkingdom Deuterostomia was completely unknown.Then in the second decade(19952005),the great discoveries of various deuterostomes led to the construction of the basic framework of the Deuterostomia and then to the formation of the tripartite phylogenetic trees of early animals(TPTEA,including basal animals,protostomes and deuterostomes),for the first time.In the third decade(after 2005),the academic community began to think about the internal relationships between the formation of TPTEA and the multiepisode Cambrian explosion,leading to the new hypothesis of the three-episode Cambrian explosion.The Chengjiang fauna is important for deciphering the fauna structure,paleoecological environment,and so on.However,its core academic values mainly rest on two aspects.Firstly,The Chengjiang fauna,as the main phase witness of the Cambrian explosion,has created a nearly complete phylogenetic framework of the TPTEA on Earth for the first time.The three-phase radiation hypothesis reveals the essential connotation of the Cambrian explosion:a step-wise divergent evolution of animals,from basal to highly advanced groups,lasting about 40 million years.In the first phase,it gave birth to a bulk of basal animals(including some now extinct“animal”groups)in the latest Ediacaran,probably including some pioneer protostomes.The second phase took place in the first epoch of the Cambrian period(Terreneuvian),giving rise to the main invertebrate protostomes with a persistent prosperity of basal animals.The third phase proceeded in Cambrian Epoch 2(represented by the Chengjiang fauna),which not only maintained the prosperity of basal animals and protostomes,but also,more importantly,gave birth to all the main phyla of the subkingdom Deuterostomia.Thus,the rudimental framework of the whole TPTEA has been shaped,with the termination of the major innovation events of the Cambrian explosion.Here,we discussed the evolutionary properties of Ediacaran biota,small shelly fossils and the Chengjiang fauna in the Cambrian explosion with emphasis on the biological properties of several important animal groups.The order Myllokunmingiida is the only known oldest vertebrate,while Yunnanozoon and Haikouella are neither vertebrates nor stem-group chordates but a special group of basal deuterostomes;Cheungkongella is a credible ancestor of the urochordate and it supports the classical hypothesis on the origin of the urochordates;and the gill slits were first invented in the members of the phylum Vetulicolia to provide key information on the origin of the deuterostomes.The second core value of the Chengjiang fauna is of profound humanistic and philosophical significance:the discoveries of the‘first gill openings’,‘first brain’,‘first vertebrae’and‘first heart’provide the pivotal evidence for solving the unsolved mystery of the origin of the main basic human organs as described in Darwin’s“The Descent of Man”.In addition,the morphological and anatomical information of the Chengjiang fauna can provide important clues for a better understanding of most components of Ediacaran and Cambrian metazoans.
基金supported by funds from the National Natural Science Foundation of China (Grant Nos. 41930319, 41890844, 41890845 and 41720104002)the Strategic Priority Research Program of the Chinese Academy of Sciences (Grant No. XDB26000000)+3 种基金111 Project (Grant No. D17013)Key Scientific and Technological Innovation Team Project in Shaanxi Provincefunded by the China Postdoctoral Science Foundation (Grant No. 2022M712570)support from a Herchel Smith Postdoctoral Fellowship (University of Cambridge)。
文摘Over one hundred arthropod fossil species have been described from the famous Chengjiang Lagerst?tte(South China, Cambrian Stage 3, ca. 518 Ma) including a diverse assemblage of radiodonts–a group containing Anomalocaris and its relatives. These iconic stem-group euarthropods include some of the largest animals of the time, and some are known from hundreds of specimens. A longstanding conundrum has been the rarity or absence of hurdiids from Cambrian Series 2 Lagerst?tten like Chengjiang. This is because radiodonts are generally common in such deposits and the oldest radiodont ever discovered is a hurdiid. Furthermore, this family displays the widest geographic and temporal ranges of all radiodont families, and the highest diversity. Here we document the first hurdiid frontal appendages from Chengjiang, which display unique features within the family and may provide insights for understanding the character evolution of hurdiid appendages. The palaeoenvironmental distribution of hurdiids suggests that the rarity of hurdiids in Chengjiang may be due to a preference for deeper water environments,and the later success of this family from the Wuliuan onwards may relate to their ability to tolerate cooler water temperatures than other radiodont families. The palaeogeographical, palaeoenvironmental, and stratigraphical patterns observed in hurdiids maybe caused in part by the limited distributions of Konservat-Lagerst?tten in the Cambrian as well.
文摘Ancestral lobopodians,which diversified and flourished in the Cambrian seas,have long drawn much attention in that not only their living counterparts,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
基金supported by the Natural Science Foundation of China(Nos.41930319,41621003,and EAR-1554897)the 111 Project(No.D17013)the Natural Science Basic Research Plan of Shaanxi Province(No.2022JC-DW5-01).
文摘Burgess Shale-type deposits provide a wealth of information on the early evolution of animals.Questions that are central to understanding the exceptional preservation of these biotas and the paleoenvironments they inhabited may be obscured by the post-depositional alteration due to metamorphism at depth and weathering near the Earth’s surface.Among over 50 Cambrian BST biotas,the Chengjiang and Qingjiang deposits are well known for their richness of soft-bodied taxa,fidelity of preservation,and Early Cambrian Age.While alteration via weathering has been well-investigated,the thermal maturity of the units bearing the two biotas has not yet been elucidated.Here we investigate peak metamorphic temperatures of the two deposits using two independent methods.Paleogeotemperature gradient analyses demonstrate that the most fossiliferous sections of the Chengjiang were buried at a maximum depth of∼8500 m in the Early Triassic,corresponding to∼300°C,while the type area of the Qingjiang biota was buried at a maximum depth of∼8700 m in the Early Jurassic,corresponding to∼240°C.Raman geothermometer analyses of fossil carbonaceous material demonstrate that peak temperatures varied across localities with different burial depth.The two productive sections of the Chengjiang biota were thermally altered at a peak temperature of approximately 300°C,and the main locality of the Qingjiang biota experienced a peak temperature of 238±22°C.These results from two independent methods are concordant.Among BST deposits for which thermal maturity has been documented,the Qingjiang biota is the least thermally mature,and therefore holds promise for enriching our understanding of BST deposits.
文摘The Cambrian Explosion is real Fossil evidence from Chengjiang reveals an assemblage of the fauna, which is represented by at least 100 species. They belong to animal phyla that are still extant, including sponges, cnidarians, ctenophorans, priapulids, annelids(?), arthropods,brachiopods, phoronids, echinoderms and chordates. There are even a number of animal body plans at phylum or subphylum level that are extinct, including: medusiform Eldonia, Facivermis, Tardipolyda, chanccelariids, Dinomischus, hyoliths, Vetuliculid,
基金This research project was granted jointly by the National Natural Science Foundation of China(Grant No.40073001)the Key Project(97D0072)of Yunnan Provincial Foundation.
文摘Twenty black shale samples, which are free from the influence of weathering, were collected from the Chengjiang Fauna-bearing horizon, central Yunnan Province, yielding a Pb-Pb isochron age of 534±60 Ma. Although this age is younger than both the Rb-Sr isochron age and 40Ar-39Ar age, it should represent the lower isotopic age limit of the Chengjiang Fauna.
基金supported by National Natural Science Foundation of China (Grant Nos. 40725005, 40930211, J0630967)National Basic Research Program of China (Grant No. 2006CB806401)
文摘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.
基金supported by National Natural Science Foundation of China (Grant Nos. 41030315,41072088)the China Geological Survey(Grant No. 1212011121105)
文摘The Chengjiang Formation is the earliest continental clastic deposit after the Jinning Orogeny in central Yunnan Province,and therefore its accurate depositional age is significant for understanding the formation and evolution of the Neoproterozoic rift basins in southern China.However,hampered by accuracy of the dating technique,the existing age data for the Chengjiang Formation are not very reliable.A large number of magmatic zircons were obtained from the tuff interbed in the lower part of the Chengjiang Formation in Jinyang area and the bottom part of the Chengjiang Formation in Dongchuan area,central Yunnan Province,and high-precision SHRIMP U-Pb dating was carried out on these zircons.The results show that the weighted mean 206 Pb/238 U ages are 797.8±8.2 and 803.1±8.7 Ma respectively.In combination with related geological evidence,it has been demonstrated that the bottom boundary age of the Chengjiang Formation should be 800±5 Ma,whereas the top boundary age could be ca 725 Ma.By synthetically analyzing the latest age data for the relevant strata,it has been confirmed that the Chengjiang Formation should be correlated with the Suxiong Formation and the Kaijianqiao Formation in western Sichuan Province,the Liantuo Formation in the middle and lower Yangtze,the Hongchicun Formation and the Shangshu Formation in northern Zhejiang Province,the Puling Formation in southern Anhui Province,the Luokedong Formation and the Majianqiao Formation in northwestern Jiangxi Province,the Wuqiangxi Formation in northwestern Hunan Province,the Fanzhao Formation and the Qingshuijiang Formation in southeastern Guizhou Province,and the Sanmenjie Formation and the Gongdong Formation in northern Guangxi Province.Sedimentary cycle analysis shows that the sedimentary filling sequence of the Neoproterozoic rift basins in southern China can be divided into four cycles.Among them,Cycle II began at ca.800 Ma,accompanied by intensive tectonic-thermal events.The zircon U-Pb ages from the bottom of the Chengjiang Formation reported in this paper indicate that the Neoproterozoic Kangdian rift subbasin probably started to develop at ca.800 Ma and therefore missed Cycle I of the Neoproterozoic sedimentary filling sequence in southern China.
基金supported by the National Natural Science Foundation of China(Grant NOS.40332016 and 30270207)the National "973" Project(Grant No.G-2000077702)
文摘The taxonomy of an early ancestor of Recent Priapulidae, Xiaoheiqingella peculiaris (= Yunnanpriapulus halteroformis Huang et al., 2004) from the Early Cambrian Chengjiang fossil Lagerst?tte, is revised. Morphological characters comprise a pair of caudal appendages rather than a single appendage flanking the trunk end and a possible urogenital duct found inside the preanal region. An addi- tional extremely rare fossil priapulid worm, Paratubiluchus bicaudatus gen. nov., sp. nov. is also described herein. Its diagnostic characters are: an introvert bearing 25 longitudi- nal rows of scalids, a distinct neck region, no annulus on the oval trunk, and a pair of caudal appendages. The proportion of body parts is similar in size to that of loricate larvae of Recent priapulids and larva-formed Palaeopriapulitidae. Taking account of the features of Xiaoheiqingella, bicaudal appendages are considered to be a synapomorphy of Pri- apulidae and Tubiluchidae. Paratubiluchus gen. nov. is most likely a candidate for the ancestor of the Tubiluchidae; it probably originated from a larva-formed priapulid with 25 rows of scalids, thus representing an intermediate link be- tween the priapulids in mature-form and the priapulids with lorica.
基金supported by the National Natural Science Foundation of China(4070200541072017+8 种基金40830208)the 973 Project(2013CB835002)111 Project of China(P201102007)Swedish Research Council(VR 2009-43952012-1658 to LEH)ZZF acknowledges grants from the Fok Ying Tung Education Foundationthe China Scholarship Counciland the Ministry of Education of China(FANEDD200936NCET-11-1046)
文摘The Cambrian explosion was coined to describe the geologically sudden appearance of numerous bilaterian body plans(Phyla)around the Ediacaran-Cambrian transition,around 565-520million years ago.Many explanations and conjectures have been postulated in order to explain the pattern and duration of this explosive radiation of many different phyla of early metazoans.Here,we focus on the evolution of a phylum of marine suspension-feeding animals—the brachiopods,as exemplified by the exceptionally preserved taxa from the celebrated Chengjiang Konservat Lagerst(a|¨)tte(Yunnan,China).The abundant soft-bodied preservation at these fossil quarries gives us the only firm insights into what brachiopods looked like and how they functioned and lived when they first appeared on the Earth.Studies of Chengjiang brachiopods demonstrate that the early animals developed a remarkably varied organization of tissues and organs shortly after the onset of Cambrian explosion.In the marine suspension-feeding brachiopods,most importantly the tentaculate feeding structure of early brachiopods is already differentiated into two shapes of lophophore,anteriorly coiled(spiralled)and posteriorly arching tentacle crowns and the unique latter type was previously not documented from fossil and living brachiopods.Also unlike any known Recent brachiopod,all the known Cambrian brachiopods from Chengjiang have an open digestive tract that was disposed either as a Ushaped gut in linguliform and stem group brachiopods,or straight gut with a posterior anus in some calcareous-shelled stocks.Moreover,in contrast toliving lingulids,all the Cambrian brachiopods have an epibenthic lifestyle either cemented by a ventral valve or attached by variable pedicles to establish complex ecological community encompassing primary tierers and variable secondary tierers.It is therefore assumed that brachiopods were the first benthic metazoan that achieved their success in ecological stratification and tiering complexity by late Atdabanian.The setae are also important for the brachiopod suspension-feeding life style,and in the Chengjiang braehiopods they include two types cilia-like and spine-like setae.The mantle canals of different braehiopod species are also distinctly variable in arrangement,mainly disposed in pinnate,baeulate and peripheral conditions.Of these,the peripheral disposition of mantle canals is for the first time proposed here so as to differentiate from the bifurcate condition in recent lingulids in that the former is devoid of posteriorly extending main trunks of sinus,but possesses a diverging dorsal vascula media in dichotomy.
基金supported by National Natural Science Foundation of China (Grant No. 40730211)National Basic Research Programme of China (Grant No. 206CB806400)Natural Science Foundation of Department of Science and Technology of Yunnan Province (Grant No. 2005D002Z)
文摘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.
文摘The origin of arthropods has long been one ofthe most hotly-debated subjects. Arthropods used to bethought closely related with annelids, but the two groups arenow believed to be separated into two major realms withinProtostomia on the basis of new molecular data. Although itis generally held by paleontologists that arthropods should berooted in the early lobopods-.a kind of worm-like creaturewith non-segmented legs, no intermediate forms have beenfound to bridge them. Here we report an organism with amixture of characters, including features characteristic ofarthropods (e.g., primary cephalization with paired eyes,paired antennae, and preliminary tagmosis) and of lobopods(e.g., worm-like body design, the dorsal spines, andnon-segmented limbs or lobe-like legs). The discovery of therare transitional form may throw new light on the origin ofarthropods and suggests that the most primitive arthropodsbegan with paired uniramous legs and the biramous onesevolved later.
基金supported by the Program of Chinese Academy of Sciences(KZZD-EW-02-2)the National Basic Research Program of China(2013CB835006)+2 种基金the National NaturalScience Foundation of China(41002002,41372021,J1210006)the Natural Science Foundation of Jiangsu Province(BK2012893)the National Science and Technology Major Project(2011ZX05008)
文摘A new Chengjiang-type fossil assemblage is reported herein from the lower part of the Hongjingshao Formation at Xiazhuang village of Chenggong,Kunming,Yunnan.The fossil assemblage,named as Xiazhuang fossil assemblage,yields predominantly soft-bodied fossils,including arthropods,brachiopods,priapulids,lobopods and some problematic taxa,with arthropods being the most dominant group.Preservation and composition of the fossil assemblage are very similar to the typical Chengjiang biota,which is preserved in the middle Yu’anshan Formation in the large area of eastern Yunnan.The associated trilobites demonstrate that the soft-bodied fossil assemblage belongs to the late Qiongzhusian in age(Stage 3,Cambrian),suggesting that the Hongjingshao Formation is probably a diachronous lithostratigraphic unit ranging from the upper Qiongzhusian to the lower Canglangpuan stages in eastern Yunnan.The fossil assemblage from the Xiazhuang area fills up the missing link between the typical older Chengjiang biota and the younger Malong and Guanshan biotas,making eastern Yunnan a unique area in the world to reveal the early evolutionary history of animals and palaeocommunity dynamics during the‘‘Cambrian explosion’’.
基金the National Natural Science Foundation of China (Grant Nos. 40332016, 32070207) the Program for Changjiang Scholars and Innovative Research Team in University (PCSIRT).
文摘The efforts of labor-intensive collecting in the Early Cambrian Chengjiang deposits in eastern Yunnan Province, China led to the discovery of many horizons containing exceptionally well preserved soft-bodied fossil concentrations, many of which can be assigned to either monospecific concentrations or paucispecific concentrations. The features of these fossil concentrations support the hypothesis that frequent storm events producing tempestites mainly contributed to the preservation of abundant soft- bodied fossils in the Chengjiang deposits, and indi- cate that the balance of the ecological web in this region was probably frequently destroyed or upset by such storm events during that geological time. Ani- mals in a fossil assemblage in such fossil concentra- tions probably occupied similar ecological biotopes.
基金This work was supported by the National Science Foundation of China (Grant Nos. 40332016, 30270207)the Science Foundation of Yunnan Province (2002D0033Q)the Program for Changjiang Scholar and Innovative Research Team in the University and Ministry of Science and Technology of China (Grant No. 2000077700).
文摘Facivermis yunnanicus (Hou & Chen, 1989), from the Early Cambrian Chengjiang Lager- st?tte, a worm-like fossil with 5 pairs of tentacles and a perceived shrunken end, has been regarded as related to polychaetes, later it has been variously interpreted as lobopods, Pentastoma and lopho- phorates. Newly discovered complete specimens by the ELI field team show that the taxon has, in addition to the 5 pairs of appendages, a pear-shaped trunk end bearing two or three circles of hooks. Accordingly, based on these important morphological characters, reconsideration of its affinities is provided and the taxonomy is remedied herein. Because the five pairs of appendages of Facivermis yunnanicus resemble the appendages of the fore-trunk of lobopod Mi- raluolishania (Liu & Shu, 2004), it seems that they are homologous structures. Therefore, the affinities of Facivermis are regarded here as being close to the lobopods, in addition, the lobe-like appendages of Facivermis are very crucial to exploring the origin of the appendages of lobopods and arthropods.