Modern optics in exceptionally preserved eyes of Early Cambrian arthropods from Australia

Despite the status of the eye as an "organ of extreme perfection", theory suggests that complex eyes can evolve very rapidly. The fossil record has, until now, been inadequate in providing insight into the early evolution of eyes during the initial radiation of many animal groups known as the Cambrian explosion. This is surprising because Cambrian Burgess-Shale-type deposits are replete with exquisitely preserved animals, especially arthropods, that possess eyes. However, with the exception of biomineralized trilobite eyes, virtually nothing is known about the details of their optical design. Here we report exceptionally preserved fossil eyes from the Early Cambrian (～ 515 million years ago) Emu Bay Shale of South Australia, revealing that some of the earliest arthropods possessed highly advanced compound eyes, each with over 3,000 large ommatidial lenses and a specialized 'bright zone'. These are the oldest non-biomineralized eyes known in such detail, with preservation quality exceeding that found in the Burgess Shale and Chengjiang deposits. Non-biomineralized eyes of similar complexity are otherwise unknown until about 85 million years later. The arrangement and size of the lenses indicate that these eyes belonged to an active predator that was capable of seeing in low light. The eyes are more complex than those known from contemporaneous trilobites and are as advanced as those of many living forms. They provide further evidence that the Cambrian explosion involved rapid innovation in fine-scale anatomy as well as gross morphology, and are consistent with the concept that the development of advanced vision helped to drive this great evolutionary event.     

Phylogeny and life habits of Early Arthropods-Predation in the Early Cambrian Sea

We investigated two new arthropods from the Maotianshan-Shale fauna of southern China in the course of our research on life strategies, particularly predation, in Early Cambrian marine macrofaunal biota. One form clearly belongs to the so-called "great-appendage" arthropods, animals that were, most likely, active predators catching prey with their first pair of large, specialized frontoventral appendages. Based on this, we hypothesize that the new species and many others, if not all, of the "great-appendage" arthropods were derivatives of the chelicerate stem lineage and not forms having branched off at different nodes along the evolutionary lineage of the Arthropoda. Rather, we consider the "great-appendage" arthropods as belonging to a monophyletic clade, which modified autapomorphically their first pair of appendages (antennae in general arthropod terminology) into raptorial organs for food capture. The second new form resembles another Maotianshan-Shale arthropod, Fuxianhuia protensa, in sharing a head made of only two separate segments, a small segment bearing oval eyes laterally, and another bearing a large tergite, which forms a wide shield freely overhanging the subsequent narrow trunk segments. This segment bears a single pair of rather short anteriorly directed uniramous appendages, considered as the "still" limb-shaped antennae. Particularly the evolutionary status of head and limbs of these two forms suggests that both are representatives of the early part of the stem lineage toward the crown-group of Arthropoda, the Euarthropoda. These forms appear rather unspecialized, but may have been but simple predators. This adds to our hypothesis that predation was a common, if not dominant feeding strategy in the Cambrian, at least for arthropods.     

Possible developmental mechanisms underlying the origin of the crown lineages of arthropods

The extraordinarily preserved, diverse arthropod fauna from the Lower Cambrian Maotianshan shale, central Yunnan (southwest China), represents different evolutionary stages stepping from stem lineages towards crown arthropods (also called euarthropods), which makes this fauna extremely significant for discussion of the origin and early diversification of the arthropods. Anatomical analyses of the Maotianshan shale arthropods strongly indicate that     

An epipodite-bearing crown-group crustacean from the Lower Cambrian

Crown-group crustaceans (Eucrustacea) are common in the fossil record of the past 500 million years back to the early Ordovician period, and very rare representatives are also known from the late Middle and Late Cambrian periods. Finds in Lower Cambrian rocks of the Phosphatocopina, the fossil sister group to eucrustaceans, imply that members of the eucrustacean stem lineage co-occurred, but it remained unclear whether crown-group members were also present at that time. 'Orsten'-type fossils are typically tiny embryos and cuticle-bearing animals, of which the cuticle is phosphatized and the material is three-dimensional and complete with soft parts. Such fossils are found predominantly in the Cambrian and Ordovician and provide detailed morphological and phylogenetic information on the early evolution of metazoans. Here we report an Orsten-type Konservat-Lagerst?tte from the Lower Cambrian of China that contains at least three new arthropod species, of which we describe the most abundant form on the basis of exceptionally well preserved material of several growth stages. The limb morphology and other details of this new species are markedly similar to those of living cephalocarids, branchiopods and copepods and it is assigned to the Eucrustacea, thus representing the first undoubted crown-group crustacean from the early Cambrian. Its stratigraphical position provides substantial support to the proposition that the main cladogenic event that gave rise to the Arthropoda was before the Cambrian. Small leaf-shaped structures on the outer limb base of the new species provide evidence on the long-debated issue of the origin of epipodites: they occur in a set of three, derive from setae and are a ground-pattern feature of Eucrustacea.     

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.     

Arthropod trace fossils from the Zhujiaqing Formation (Meishucunian, Yunnan) and their palaeobiological implications

1　Biostratigraphicbackgroundandgeologi calsettingIneasternYunnan ,thesupra tointertidaldolomitesoftheXiaowaitoushanMember (Upper mostDengyingFormation =LatestSinian)areun conformablyoverlainbythesubsequent (earliestCambrian) phosphoritesoftheZhujiaqingFormation(MeishucunianStage) .TheoutcropsnearMeishucuninJinningCounty ,southwestofKunmingrepresentsakeysectionforthePC/CtransitionontheYangtzeplatform[1] .TheZhujiaqingFormationbelongstotheEarlytoMiddleMeishucunian (=Nemakit DaldynSt…     

A giant Ordovician anomalocaridid

Anomalocaridids, giant lightly sclerotized invertebrate predators, occur in a number of exceptionally preserved early and middle Cambrian (542-501?million years ago) biotas and have come to symbolize the unfamiliar morphologies displayed by stem organisms in faunas of the Burgess Shale type. They are characterized by a pair of anterior, segmented appendages, a circlet of plates around the mouth, and an elongate segmented trunk lacking true tergites with a pair of flexible lateral lobes per segment. Disarticulated body parts, such as the anterior appendages and oral circlet, had been assigned to a range of taxonomic groups--but the discovery of complete specimens from the middle Cambrian Burgess Shale showed that these disparate elements all belong to a single kind of animal. Phylogenetic analyses support a position of anomalocaridids in the arthropod stem, as a sister group to the euarthropods. The anomalocaridids were the largest animals in Cambrian communities. The youngest unequivocal examples occur in the middle Cambrian Marjum Formation of Utah but an arthropod retaining some anomalocaridid characteristics is present in the Devonian of Germany. Here we report the post-Cambrian occurrence of anomalocaridids, from the Early Ordovician (488-472?million years ago) Fezouata Biota in southeastern Morocco, including specimens larger than any in Cambrian biotas. These giant animals were an important element of some marine communities for about 30?million years longer than previously realized. The Moroccan specimens confirm the presence of a dorsal array of flexible blades attached to a transverse rachis on the trunk segments; these blades probably functioned as gills.     

Progress and significance in research on the early Middle Cambrian Kaili Biota, Guizhou Province,China

Important progress in research on the Kaili Biota has been made recently. Many interesting components from Chengjiang Biota and Burgess Shale Biota have been discovered, e.g. Microdictyon of lobopodia; Ottoia, Palaeoscolex of worms; Naraoia, Marrella of Trilobitioidea, Mollisonia, anamalocarids and other non-trilobite arthropods; and new sorts of echinoder-mas, macroalage fossils and so on. Recent work on the Kaili Biota has resulted in the following developments: (i) an increase in the number of animal genera, up to more than 100 genera in total, so that the Kaili Biota has become the third most diverse of the Burgess Shale-type Biota after the Burgess Shale and Chengjiang Biotas; and (ii) the most noteworthy fossils in the Kaili Biota are echinoderms, non-trilobite arthropods and soft-bodied medusiform fossils, especially the most diverse echinoderms. The progress provides envidence for the biodiversity of marine organisms presented after the "Cambrian Explosion" and serves as a link between the earlist Cambrian Chengjiang Biota and late early Middle Cambrian Burgess Shale Biota. It is of great significance in the reconstruction of the Cambrian palaeoplate, palaeongeography and in research on taphonomy.     

A Cambrian micro-lobopodian and the evolution of arthropod locomotion and reproduction

The evolutionary success of arthropods, the most abundant and diverse animal group, is mainly based on their segmented body and jointed appendages, features that had evolved most likely already before the Cambrian. The first arthropod-like animals, the lobopodians from the Early Cambrian, were unsclerotized and worm-like, and they had unjointed tubular legs. Here we describe the first three-dimensionally preserved Cambrian lobopodian. The material presented of Orstenotubulus evamuellerae gen. et sp. nov. is the smallest and youngest of a lobopodian known. O. evamuellerae shows strikingly detailed similarities to Recent tardigrades and/or onychophorans in its cellular-structured cuticle and the telescopic spines. It also shows similarities to other, longer known lobopodians, but which are ten times as large as the new form. These similarities include the finely annulated body and legs, which is characteristic also for Recent onychophorans, and paired humps continuing into spines situated dorsally to the leg insertions, a feature lacking in the extant forms. The morphology of O. evamuellerae not only elucidates our knowledge about lobopodians, but also aids in a clearer picture of the early evolution of arthropods. An example is the single ventral gonopore between a limb pair of O. evamuellerae, which indicates that a single gonopore, as developed in onychophorans, tardigrades, pentastomids, myriapods and insects, might represent the plesiomorphic state for Arthropoda, while the paired state in chelicerates and crustaceans was convergently achieved. Concerning life habits, the lateral orientation of the limbs and their anchoring spines of the new lobopodian imply that early arthropods were crawlers rather than walkers.     

Lower Cambrian small shelly faunas from Zhejiang (China)and their biostratigraphical implications

Despite a long history of research on the Early Cambrian in China most available data on small skeletal fossils concern fossil associations of the shallow carbonate platform. Information on skeletal fossils from marginal shelf environments of the Yangtze Platform is scanty, which may reflect the rarity of fossils in deeper sedimentary environments but is also due to limitation of carbonate distribution and outcrops, difficulties in fossil extraction, and a general research focus on the Precambrian-Cambrian boundary beds on the carbonate platform. Here we present a documentation of Meishucunian to Qiongzhusian small skeletal fossils from the lower Hetang Formation and the chert unit at its base from the Jiangshan region, Zhejiang Province, representing a relatively deep shelf environment compared to the inner shelf region. The earliest association (Meishucunian) from the chert unit underlying the Hetang Formation is mainly characterized by the occurrence of Protohertzina anabarica, P. unguliformis, Fengzuella zhejiangensis, and Kaiyangites novilis, which differs somewhat in composition from SSF-associations of typical inner shelf deposits. The enigmatic skeletal fossil Fengzuella zhejiangensis, which exhibits an unusual secretional growth mode previously unrecognized from the Early Cambrian, is described in detail. A younger (Qiongzhusian) fossil association contains numerous arthropod remains, such as disarticulated spines of arthropods (Jiangshanodus- and Kijacus-type), which have previously been considered as conodont-like fossils, and bradoriid valves.     

A palaeontological solution to the arthropod head problem

The composition of the arthropod head has been one of the most controversial topics in zoology, with a large number of theories being proposed to account for it over the last century. Although fossils have been recognized as being of potential importance in resolving the issue, a lack of consensus over their systematics has obscured their contribution. Here, I show that a group of previously problematic Cambrian arthropods from the Burgess Shale and Chengjiang faunas form a clade close to crown-group euarthropods, the group containing myriapods, chelicerates, insects and crustaceans. They are characterized by modified or even absent endopods, and two pre-oral appendages. Comparison with reconstructions of the crown-group euarthropod ground plan and recent investigations into onychophorans demonstrates that these two appendages are the first antenna (of extant crustaceans) and a more anterior appendage associated with an ocular segment. The latter appendage has been reduced in all crown-group euarthropods. Its most likely relic is as a component of the labrum. These fossils thus tie together results from disparate living groups (onychophorans and euarthropods).     

The occurrence of the genus Marrella(Trilobitoidea) in Asia

An important discovery in the Kaili Biota has been made recently, and many interesting components from ChengjiangBiota and Burgess Shale Biota have been discovered. Among them Marrella, only known from mid-Middle Cambrian Burgess Shale Biotaof British Columbia, Canada, is one of the most important arthropods fossils. Because the Kaili Biota is older than the Burgess Shale Biota,the strange head shield of Marrella occurring in the Kaili Biota outside Laurentia ranges from mid-Middle Cambrian to early Middle Cam-brian and is significant for the reconstruction of palaeogeography and evolutionary study of early metazoa. In the present paper Marrellasp. is reported for the first time in Asia.     

An armoured Cambrian lobopodian from China with arthropod-like appendages

Cambrian fossil Lagerst?tten 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 Lagerst?tte (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.     

Doushantuo embryos preserved inside diapause egg cysts

Phosphatized microfossils in the Ediacaran (635-542 Myr ago) Doushantuo Formation, south China, have been interpreted as the embryos of early animals. Despite experimental demonstration that embryos can be preserved, microstructural evidence that the Doushantuo remains are embryonic and an unambiguous record of fossil embryos in Lower Cambrian rocks, questions about the phylogenetic relationships of these fossils remain. Most recently, some researchers have proposed that Doushantuo microfossils may be giant sulphur-oxidizing bacteria comparable to extant Thiomargarita sp. Here we report new observations that provide a test of the bacterial hypothesis. The discovery of embryo-like Doushantuo fossils inside large, highly ornamented organic vesicles (acritarchs) indicates that these organisms were eukaryotic, and most probably early cleavage stage embryos preserved within diapause egg cysts. Large acanthomorphic microfossils of the type observed to contain fossil embryos first appear in rocks just above a 632.5 +/- 0.5-Myr-old ash bed, suggesting that at least stem-group animals inhabited shallow seas in the immediate aftermath of global Neoproterozoic glaciation.     

Sophisticated particle-feeding in a large Early Cambrian crustacean

Most Cambrian arthropods employed simple feeding mechanisms requiring only low degrees of appendage differentiation. In contrast, post-Cambrian crustaceans exhibit a wide diversity of feeding specializations and possess a vast ecological repertoire. Crustaceans are evident in the Cambrian fossil record, but have hitherto been known exclusively from small individuals with limited appendage differentiation. Here we describe a sophisticated feeding apparatus from an Early Cambrian arthropod that had a body length of several centimetres. Details of the mouthparts resolve this taxon as a probable crown-group (pan)crustacean, while its feeding style, which allowed it to generate and handle fine food particles, significantly expands the known ecological capabilities of Cambrian arthropods. This Early Cambrian record predates the major expansions of large-bodied, particle-handling crustaceans by at least one hundred million years, emphasizing the importance of ecological context in driving adaptive radiations.     

A Chengjiang-type fossil assemblage from the Hongjingshao Formation （Cambrian Stage 3） at Chenggong,Kunming, Yunnan

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’’.     

Neuroanatomy of sea spiders implies an appendicular origin of the protocerebral segment

Independent specialization of arthropod body segments has led to more than a century of debate on the homology of morphologically diverse segments, each defined by a lateral appendage and a ganglion of the central nervous system. The plesiomorphic composition of the arthropod head remains enigmatic because variation in segments and corresponding appendages is extreme. Within extant arthropod classes (Chelicerata, Myriapoda, Crustacea and Hexapoda--including the insects), correspondences between the appendage-bearing second (deutocerebral) and third (tritocerebral) cephalic neuromeres have been recently resolved on the basis of immunohistochemistry and Hox gene expression patterns. However, no appendage targets the first ganglion, the protocerebrum, and the corresponding segmental identity of this anterior region remains unclear. Reconstructions of stem-group arthropods indicate that the anteriormost region originally might have borne an ocular apparatus and a frontal appendage innervated by the protocerebrum. However, no study of the central nervous system in extant arthropods has been able to corroborate this idea directly, although recent analyses of cephalic gene expression patterns in insects suggest a segmental status for the protocerebral region. Here we investigate the developmental neuroanatomy of a putative basal arthropod, the pycnogonid sea spider, with immunohistochemical techniques. We show that the first pair of appendages, the chelifores, are innervated at an anterior position on the protocerebrum. This is the first true appendage shown to be innervated by the protocerebrum, and thus pycnogonid chelifores are not positionally homologous to appendages of extant arthropods but might, in fact, be homologous to the 'great appendages' of certain Cambrian stem-group arthropods.     

Comparative study of Cambrian lobopods <Emphasis Type="Italic">Miraluolishania</Emphasis> and <Emphasis Type="Italic">Luolishania</Emphasis>

The rare fossil Miraluolishania described by Liu et ah from the Lower Cambrian Chengjiang Lagerstatte in 2004 is regarded as an arthropod sphinx because it bears mosaic features of both Iobopods and arthropods. The discovery of this rare transitional form offers direct fossil evidence for exploring the relationship between Iobopods and arthropods. However, some scientists consider Miraluolishania to be a junior synonym of Luolishania because the former superficially resembles the latter in general appearance. Considering the significant differences between the two taxa, a thorough comparative study of Miraluolishania and Luolishania leads to the conclusion that there are definitely two different genera. Nevertheless, the ＂Luolishania＂ of the Haikou area is indeed ＂Miraluolishania＂, whereas Luolishania is most likely the typical genus of the Maotianshan area of Chengjiang County.     

Early Cambrian protoconodonts and conodont-like fossils from China: Taxonomic revisions and stratigraphic implications

Tooth likephosphaticprotoconodontsand“con odont likefossils”arecharacteristiccomponentsoftheEarlyCambriansmallshellyfossils (SSF )withaworldwidedistribution .Theyarerepresentedexclu sivelybyisolatedscleritesandareoften presentinaceticacid etchedresiduesfromLowerCambriancar bonates .SincethefirstreportofProtohertzinaMis sarzhevsky 1973fromthebasalCambrianoftheYangtzePlatformin 1977[1] ,andalongwithenthusi asticresearchactivitiesonthesmallshellyfossilsandcandidatesectionsforaglobalPrecamb…     

Skeletal faunas from the Qiongzhusian of southern Shaanxi: biodiversity and lithofacies-biofacies links in the Lower Cambrian carbonate settings

The Xiaoyang section, located in Zhenba County of Shaanxi Province, is important both for documenting the biodiversity of the Qiongzhusian Stage (Lower Cambrian) and for analyzing lithofacies-biofacies links in carbonate depositing environments on the Yangtze Platform.The skeletal fossils from the Xihaoping Member, including linguloid valves, cambroclavid and chancelloriid sclerites, genal and occipital spines of polymerid trilobites, and tubular fossils, are described here as the Eoobolus-Cambroclavus fauna, which mainly occurred in peritidal environments.The fossils from the argillaceous limestones of the lower Shuijingtuo Formation, containing lingulate brachiopods, eodiscoid trilobites, bivalved arthropods, microdictyoniid sclerites and siliceous sponge spicules, are regarded as the Palaeobolus-Hupeidiscus-Kunmingella fauna, which presumably lived in subtidal or deeper water environments.The faunal diversity in the Xihaoping Member is lower than that in the lower Shuijingtuo Formation.The investigations of the skeletal fossil assemblages from carbonate lithofacies in southern Shaanxi will significantly contribute to a detailed documentation of the biodiversity during the Qiongzhusian interval on the Yangtze Platform and add new knowledge on the Cambrian bioradiation.