Acute vision in the giant Cambrian predator Anomalocaris and the origin of compound eyes

Until recently, intricate details of the optical design of non-biomineralized arthropod eyes remained elusive in Cambrian Burgess-Shale-type deposits, despite exceptional preservation of soft-part anatomy in such Konservat-Lagerst?tten. The structure and development of ommatidia in arthropod compound eyes support a single origin some time before the latest common ancestor of crown-group arthropods, but the appearance of compound eyes in the arthropod stem group has been poorly constrained in the absence of adequate fossils. Here we report 2-3-cm paired eyes from the early Cambrian (approximately 515 million years old) Emu Bay Shale of South Australia, assigned to the Cambrian apex predator Anomalocaris. Their preserved visual surfaces are composed of at least 16,000 hexagonally packed ommatidial lenses (in a single eye), rivalling the most acute compound eyes in modern arthropods. The specimens show two distinct taphonomic modes, preserved as iron oxide (after pyrite) and calcium phosphate, demonstrating that disparate styles of early diagenetic mineralization can replicate the same type of extracellular tissue (that is, cuticle) within a single Burgess-Shale-type deposit. These fossils also provide compelling evidence for the arthropod affinities of anomalocaridids, push the origin of compound eyes deeper down the arthropod stem lineage, and indicate that the compound eye evolved before such features as a hardened exoskeleton. The inferred acuity of the anomalocaridid eye is consistent with other evidence that these animals were highly mobile visual predators in the water column. The existence of large, macrophagous nektonic predators possessing sharp vision--such as Anomalocaris--within the early Cambrian ecosystem probably helped to accelerate the escalatory 'arms race' that began over half a billion years ago.     

华南布尔吉斯页岩型软躯体化石生物群对比研究

目的梳理华南布尔吉斯页岩型软躯体化石生物群的产出层位、生物面貌和沉积环境。方法对10个华南布尔吉斯页岩型软躯体化石生物群进行对比研究。结果除了岩家河生物群之外的9个布尔吉斯页岩型生物群代表了一系列生活在泥砂质基底的、从时空分布上逐渐过渡的具有高度相关性的生物群落。结论布尔吉斯页岩型化石生物群在中晚寒武世应该会有更大规模的分布。     

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.     

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.     

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.     

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.     

Primitive deuterostomes from the Chengjiang Lagerst?tte (Lower Cambrian, China).

Cambrian fossil-Lagerst?tten (sites of exceptional fossil preservation), such as those from Chengjiang (Lower Cambrian) and the Burgess Shale (Middle Cambrian), provide our best window into the Cambrian 'explosion'. Such faunas are known from about 40 localities, and have yielded a widely disparate series of taxa ranging from ctenophores to agnathan fish. Recent excavations of the Chengjiang fossil-Lagerst?tte, known from a series of sites near Kunming in Yunnan, south China, have resulted in the discovery of several new forms. In conjunction with material described earlier, these provide evidence for a new group of metazoans, the vetulicolians. Several features, notably a series of gill slits, suggest that this group can throw light on an early stage of deuterostome diversification.     

New anomalocardid frontal appendages from the Guanshan biota, eastern Yunnan

Anomalocaridids were large predators of the Cambrian seas at the top of the trophic pyramid. Complete anomalocaridid specimens have been rarely discovered and the rigid isolated frontal appendages and mouthparts are more commonly preserved. Here we study new material of the frontal appendages from the Wulongqing Formation, Cambrian Stage 4, Series 2 near Kunming, eastern Yunnan. Two new forms of anomalocaridid frontal appendages are described, namely Anomalocaris kunmingensis sp. nov. and Paranomalocaris multisegmentalis gen. nov., sp. nov. The frontal appendage of A. kunmingensis sp. nov. probably comprises 15 podomeres of which the first one has a weakened skeletoned, the second one is armed with small spines, and the third one is armed with remarkably robust proximal ventral spines with 6 anisomerous auxiliary spines; paired auxiliary spines are associated with podomeres 4–14; podomeres 12–14 are armed with paired dorsal spines, and the last podomere bears 2 distal spines, one spine distinctly larger than the other. The frontal appendage of P. multisegmentalis tapered backwards, consisting of 22 visible podomeres; the most ventral spine is armed with 5 pairs of auxiliary spines, and podomeres 12–21 bear dorsal spines, the last podomere with 2 small distal spines. The new material provides additional evidence for our understanding of the diversity of anomalocaridids in the Cambrian. The morphology of these new finds may indicate the importance of different feeding strategies of anomalocaridids in the Cambrian ecosystem.     

The earliest-known ancestors of Recent Priapulomorpha from the Early Cambrian Chengjiang Lagersttte

It was suggested and eventually has been profferedthat the representatives of all extant phyla, including someminor group, occurred during the Cambrian radia-tions[1—10]. Many priapulid-like fossil worms have beenreported from Cambrian Lagerst?tten, such as the EarlyCambrian Chengjiang Lagerst?tte[11—16], the Middle Cam-brian Burgess Shale[17], and the Middle Cambrian Kailifauna[18]. The palaeoscolecidan worms from these Lager-st?tten have been regarded as either a stem group of thepri…     

凯里动物群与我国早期后生动物演化的初步研究

作者所发现的中寒武世凯里动物群,属于带软躯体化石的布吉斯页岩型的动物群,是继云南澄江动物群之后我国发现的又一个早期后生动物群。它和新近发现的晚震旦世庙河生物群,分别填补了晚震旦世、中寒武世我国带软躯化石生(动)物群的空白,形成了我国多时期带软躯化石的生(动)物群体系。对于研究我国及世界早期后生动物演化并不断揭示其演化规律有重要意义。     

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.     

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.     

The earliest-known ancestors of Recent Priapulomorpha from the Early Cambrian Chengjiang Lagerstaette

The taxonomy of an early ancestor of Recent Priapulidae, Xiaoheiqingella peculiaris (= Yunnanpriapulus halteroformis Huang et al., 2004) from the Early Cambrian Chengjiang fossil Lagerstaette, 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 additional extremely rare fossil priapulid worm, Paratubiluchus bicaudatus gen. nov., sp. nov. is also described herein. Its diagnostic characters are: an introvert bearing 25 longitudinal 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 Priapulidae 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 between the priapulids in mature-form and the priapulids with lorica.     

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.     

中国扬子海盆——世界上罕见寒武纪生命大爆发和辐射进化的化石库

在简要论述我国扬子地块或扬子海盆,包括现今云南、贵州、湖南和湖北等地,寒武纪地层中相继发现的以澄江生物群、凯里生物群、牛蹄塘生物群和清江生物群等为代表的特异埋藏群及其与加拿大布尔吉斯页岩生物群的对比的同时,讨论了最近发现的湖北清江生物群的产出时代和层位、组合特征,并对上述生物群的生态和埋藏环境进行了分析与对比.认为大规模两侧对称动物门类的出现和多门类后生动物骨骼化是构成寒武纪动物大爆发的最主要特征.以上生物群的发现和研究,不仅展示寒武纪大爆发的客观存在,也说明在漫长地质历史时期,生物进化并非都是沿着达尔文渐进论的进化轨迹,从低级向高级发展;只要环境许可,生物在相对短暂时间内,采用一种爆发式和多样性辐射进化的模式,也可以出现大量门一级生物,而且这种进化模式在生物进化的历程中具有划时代的重要意义.前寒武纪罗迪尼亚(Rodinia)超大陆裂解和聚合而引发的、并波及我国南方的距今10亿～8亿年左右的晋宁造山事件,以及雪球事件后因气候变暖,大气和水中有机质和氧及二氧化碳含量的急剧变化,尤其是大气和水中氧气含量的增加,为我国扬子海盆寒武纪大爆发提供了必要的前提,也为动物的繁衍和骨骼化创造了极好的生态条件.但只有那些生活在海水相对较深,表层水柱充氧而底层缺氧、古地理位置处于扬子碳酸盐岩台地陆棚或陆棚边缘或台缘斜坡或凹陷中的岩家河、澄江、牛蹄塘、清江、凯里等生物群,才有可能在缺氧的海底被保存下来,并形成深色和黑色页岩或泥岩特异埋藏群或化石库.     

Palaeoecology: a gigantic fossil arthropod trackway

A unique, complex trackway has been discovered in Scotland: it was made roughly 330 million years ago by a huge, six-legged water scorpion that was about 1.6 m long and a metre wide. To my knowledge, this is not only the largest terrestrial trackway of a walking arthropod to be found so far, but is also the first record of locomotion on land for a species of Hibbertopterus (Eurypterida). This evidence of lumbering movement indicates that these giant arthropods, now extinct, could survive out of water at a time when the earliest tetrapods were making their transition to the land.     

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 soft-bodied mollusc with radula from the Middle Cambrian Burgess Shale

Odontogriphus omalus was originally described as a problematic non-biomineralized lophophorate organism. Here we re-interpret Odontogriphus based on 189 new specimens including numerous exceptionally well preserved individuals from the Burgess Shale collections of the Royal Ontario Museum. This additional material provides compelling evidence that the feeding apparatus in Odontogriphus is a radula of molluscan architecture comprising two primary bipartite tooth rows attached to a radular membrane and showing replacement by posterior addition. Further characters supporting molluscan affinity include a broad foot bordered by numerous ctenidia located in a mantle groove and a stiffened cuticular dorsum. Odontogriphus has a radula similar to Wiwaxia corrugata but lacks a scleritome. We interpret these animals to be members of an early stem-group mollusc lineage that probably originated in the Neoproterozoic Ediacaran Period, providing support for the retention of a biomat-based grazing community from the late Precambrian Period until at least the Middle Cambrian.     

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