The origin of crustaceans: new evidence from the Early Cambrian of China.

One of the smallest arthropods recently discovered in the Early Cambrian Maotianshan Shale Lagerstätte is described. Ercaia gen. nov. has an untagmatized trunk bearing serially repeated biramous appendages (long and segmented endopods and flap-like exopods), a head with an acron bearing stalked lateral eyes and a sclerite and two pairs of antennae. The position of this 520 million-year-old tiny arthropod within the Crustacea is supported by several anatomical features: (i) a head with five pairs of appendages including two pairs of antennae, (ii) highly specialized antennae (large setose fans with a possible function in feeding), and (iii) specialized last trunk appendages (segmented pediform structures fringed with setae). The segmentation pattern of Ercaia (5 head and 13 trunk) is close to that of Maxillopoda but lacks the trunk tagmosis of modern representatives of the group. Ercaia is interpreted as a possible derivative of the stem group Crustacea. Ercaia is likely to have occupied an ecological niche similar to those of some Recent meiobenthic organisms (e.g. copepods living in association with sediment). This new fossil evidence supports the remote ancestry of crustaceans well before the Late Cambrian and shows, along with other fossil data (mainly Early Cambrian in China), that a variety of body plans already coexisted among the primitive crustacean stock.     

A comparison of swimming structures and kinematics in three species of crustacean larvae

Crustacean larvae swim with paired rowing appendages that rotate around the body of the animal. The number of paired rowing appendages varies from one species of larvae to another. In addition, the size of the crustacean larvae is different between species and increases as they grow. The nature of the fluid forces changes as size increases, so the morphology and mechanics of swimming in these animals will change during increases in size. This article demonstrates the changing kinematics of locomotion between three species of crustacean larvae, which swim with one (Artemia franciscana), two (Carcinus maenas) or five (Homarus americanus) pairs of propulsive limbs. The relative change in the surface area and volume ratios of the locomotor structures are also demonstrated.     

介形类系统分类与起源演化的形态、分子和化石证据

介形类(Ostracoda)因其丰富的化石记录和广布的海陆现生代表类群,而被认为是进化生物学中研究生物多样性产生机制和演变历程的颇具潜力的重要模式生物。介形类在甲壳亚门中的谱系发生位置、起源及其内部各类群间的系统关系还存在诸多争议。基于其体制构造的形态学特征,介形类被归入甲壳亚门下的颚足纲(Maxillopoda),但来自18S rDNA序列数据分析却显示Maxillopoda不是单系群。基于化石记录和壳体形态特征,高肌虫(Bradoriida)长期以来被认为是介形类的一个祖先类群,但保存有软躯体的早寒武世化石的研究表明,Bradoriida不是介形类甚至可能也不属于甲壳类。不同的研究者所强调的壳体和肢体形态特征各异,导致介形类最大的现生类群速足目(Podocopida)的四个超科之间的关系也存在诸多推测。壳体和肢体特征在系统演化意义上的不兼容,需要分子生物学等证据的介入。分子、形态和化石证据的积累及各种信息整合是系统演化研究的必然趋势。     

Moulting in the lobopodian Onychodictyon from the lower Cambrian of Greenland

A number of lobopodian taxa from the Cambrian display pairs of sclerotized plates symmetrically positioned along the dorsum of the animal, predominantly above the walking appendages. Most genera were described from complete body fossils exquisitely preserved in the famous Cambrian Lagerstätten, but lobopodian phosphatized plates are found worldwide as typical components of Cambrian small shelly fossil assemblages (SSF). Details regarding intraspecific and ontogenetic variation in lobopod plates are elusive, and the lack of details of ornamentation in Lagerstätte specimens does not minimize the problem. We document here an assemblage of well‐preserved isolated plates of Onychodictyon sp. from the Lower Cambrian (Cambrian Series 2, Stage 4) of North Greenland. Two specimens exhibit perfectly conjoined plates from successive moults. Details of ornamentation and the outline and profile of the fixed plates are identical, but width and length of the underlying plate are 24% larger. These specimens boost the body of evidence that lobopodians moulted but also show that plate outline and ornamentation did not vary during ontogeny.     

Origin of the Ostracoda and their maxillopodan and hexapodan affinities

There are Cambrian fossils attributed to the Ostracoda but the extant subclasses Podocopa and Myodocopa do not appear until the Ordovician. At this time the morphologically similar, free-living ancestors of the now sedentary Thecostraca (Ascothoracida, Acrothoracica and Cirripedia) may have still been extant, and from an ecological point of view it seems likely that, by and large, ostracods replaced them. However, living ostracods have an abbreviated, direct development, and some key aspects of their morphology, such as the nature of the maxillary segment and abdomen, are conjectural. Thus the affinities between these and related taxa remain uncertain; e.g., while some contemporary carcinologists place Ostracoda as a taxon coordinate with the Branchiopoda, Remipedia, Cephalocarida, Maxillopoda, Malacostraca, others tentatively or unequivocally ally them with the Maxillopoda (generally Mystacocarida, Copepoda, Tantulocarida and Thecostraca, and sometimes Branchiura and Pentastomida). Others, largely involved with fossils, have stretched the definition of the Maxillopoda even further, to the point where it seems even less likely a monophyletic taxon. Until recently cladistic analyses utilizing genetic (largely 18S rDNA) as well traditional morphological characteristics have given confusing results regarding the affinities between these taxa, and an important one suggested the Ostracoda might even be diphyletic. Furthermore, a very recent genetic study utilizing protein encoding genes places a podocopine ostracod among the most primitive of the extant crustaceans (Branchiopoda, Cephalocarida Remipedia and Mystacocarida), and then generally at the base of a lineage leading to the Malacostraca, a lineage giving rise to copepods and cirripeds along the way. This indicates these so-called maxillopodan taxa evolved independently from a malacostracan-like ancestor, and if so they are convergent. And finally, from genetic studies it is not only becoming well documented the Crustacea rather than Myriapoda gave rise to the Hexapoda, but it appears the Hexapoda stem from among the lower rather than the higher crustaceans, possibly even from the Ostracoda. Whether there were terrestrial ostracods at the time hexapods appeared in the Lower Ordovician is unknown, but the modest diversity of terrestrial ostracods today are podocopines which also first appeared in the Lower Ordovician. Thus, if current interpretations of living ostracodan and fossil hexapodan body plans are largely correct, it can be hypothesized the Ostracoda are close to the ancestor of the Hexapoda.     

A New Bivalved Arthropod from the Early Cambrian Sirius Passet Fauna, North Greenland

A new bivalved arthropod is described from the Lower Cambrian (?Upper Atdabanian) Buen Formation of North Greenland. Pauloterminus spinodorsalis gen. et sp. nov. possesses a bivalved carapace that covers the head, which has a single pair of antennae, and anteriormost thorax. No mouthparts are visible. The five‐segmented abdomen was limbless and terminated in a telson plus a pair of large, lobate uropods. A suite of at least six biramous thoracic limbs are present: the short endopods are made up of small, serial podomeres, while the exopods are lobate and may have functioned as gills as well as in swimming. Partially infilled guts are occasionally visible, suggesting that this animal may have been a sediment feeder. It is compared to other Cambrian bivalved arthropods, especially the waptiids Chuandianella ovata from the Lower Cambrian Chengjiang fauna (China) and Waptia fieldensis from the Middle Cambrian Burgess Shale (British Columbia). Of these three animals, the Greenland and Chinese taxa appear to be the most closely related. P. spinodorsalis possesses many typical arthropod features, but it also demonstrates more primitive characters that are more reminiscent of the lobopodians.     

Crustacean disparity through the Phanerozoic: comparing morphological and stratigraphic data

Crustaceans have been an important component of marine diversity and biomass since the earliest Phanerozoic. With a relatively well-documented fossil record, they provide an excellent subject for a continuous study of disparity (? bodyplan variety) from the Cambrian to the Recent. A data base of 135 morphological characters forms the basis for cladistic and morphospace studies at the ordinal and sub-ordinal level. Gross cladistic topology is: (Eumalacostraca + Hoplocarida vs Maxillopoda) vs Phyllopoda (paraphyletic). Each of these groups is of approximately equal disparity, and occupies a distinct region of the morphospace plot. A few problematical fossils (e.g. Waptia and Odaraia) fall close to the base of the tree. Comparison of the cladogram with stratigraphic range data indicates the location of probable ghost lineages, and randomization procedures provide a statistical test of the goodness of fit of a given set of stratigraphic ranges to a given tree topology. Disparity indices are calculated at series and stage intervals. Observed range data indicate that Cambrian disparity was approximately one third its present level. The Earliest Ordovician saw a marked decrease, with an increase and subsequent plateau through rest of the period. Increases through the Silurian and Devonian corresponded to the radiation of branchiopods, cephalocarids, and latterly the Eumalacostraca and Hoplocarida. By the end of the Carboniferous, observed disparity had reached over four fifths of Recent levels, and the remaining history of the group saw a gradual but slightly irregular increase up until the end of the Tertiary. Indices of disparity incorporating ghost lineages exhibit less marked peaks and troughs, with fewer perturbations overall. Cladistically-implied disparity in the Lower Cambrian is estimated at three quarters of that in the Recent. Rarefaction is used to compare actual levels of disparity at each time interval with the mean for a similar number of taxa selected randomly from the list of all realized bodyplans. Most intervals preserved a range of forms more disparate than the mean of random samples drawn from the pool of all the taxa considered. From the Triassic to the Recent this difference was intermittently significant. Once occupied, extremes of morphospace tend not to fall vacant again.     

Reappraising the early evidence of durophagy and drilling predation in the fossil record: implications for escalation and the Cambrian Explosion

The Cambrian Explosion is arguably the most extreme example of a biological radiation preserved in the fossil record, and studies of Cambrian Lagerstätten have facilitated the exploration of many facets of this key evolutionary event. As predation was a major ecological driver behind the Explosion – particularly the radiation of biomineralising metazoans – the evidence for shell crushing (durophagy), drilling and puncturing predation in the Cambrian (and possibly the Ediacaran) is considered. Examples of durophagous predation on biomineralised taxa other than trilobites are apparently rare, reflecting predator preference, taphonomic and sampling biases, or simply lack of documentation. The oldest known example of durophagy is shell damage on the problematic taxon Mobergella holsti from the early Cambrian (possibly Terreneuvian) of Sweden. Using functional morphology to identify (or perhaps misidentify) durophagous predators is discussed, with emphasis on the toolkit used by Cambrian arthropods, specifically the radiodontan oral cone and the frontal and gnathobasic appendages of various taxa. Records of drill holes and possible puncture holes in Cambrian shells are mostly on brachiopods, but the lack of prey diversity may represent either a true biological signal or a result of various biases. The oldest drilled Cambrian shells occur in a variety of Terreneuvian‐aged taxa, but specimens of the ubiquitous Ediacaran shelly fossil Cloudina also show putative drilling traces. Knowledge on Cambrian shell drillers is sorely lacking and there is little evidence or consensus concerning the taxonomic groups that made the holes, which often leads to the suggestion of an unknown ‘soft bodied driller’. Useful methodologies for deciphering the identities and capabilities of shell drillers are outlined. Evidence for puncture holes in Cambrian shelly taxa is rare. Such holes are more jagged than drill holes and possibly made by a Cambrian ‘puncher’. The Cambrian arthropod Yohoia may have used its frontal appendages in a jack‐knifing manner, similar to Recent stomatopod crustaceans, to strike and puncture shells rapidly. Finally, Cambrian durophagous and shell‐drilling predation is considered in the context of escalation – an evolutionary process that, amongst other scenarios, involves predators (and other ‘enemies’) as the predominant agents of natural selection. The rapid increase in diversity and abundance of biomineralised shells during the early Cambrian is often attributed to escalation: enemies placed selective pressure on prey, forcing phenotypic responses in prey and, by extension, in predator groups over time. Unfortunately, few case studies illustrate long‐term patterns in shelly fossil morphologies that may reflect the influence of predation throughout the Cambrian. More studies on phenotypic change in hard‐shelled lineages are needed to convincingly illustrate escalation and the responses of prey during the Cambrian.     

The Collins’ monster,a spinous suspension-feeding lobopodian from the Cambrian Burgess Shale of British Columbia

Lobopodians, a paraphyletic group of rare but morphologically diverse Palaeozoic vermiform animals bearing metameric appendages, are key to the origin of extant panarthropods. First discovered in 1983 on Mount Stephen (Yoho National Park, British Columbia), the Cambrian (Wuliuan) Burgess Shale lobopodian nicknamed ‘Collins’ monster’ is formally described as Collinsovermis monstruosus gen. et sp. nov. A formal systematic treatment of the comparable and poorly known lobopodian Acinocricus stichus from Utah is also provided. The body of Collinsovermis is plump and compact but shows the diagnostic suspension-feeding characters of luolishaniid lobopodians. It possesses 14 contiguous pairs of lobopods, lacking space between them. The 6 anterior pairs are elongate, adorned with about 20 pairs of long and slightly curved ventral spinules arranged in a chevron-like pattern. These appendages terminate in a pair of thin claws and their dorsal surfaces are covered in minute spines or setae. The 8 posterior lobopod pairs, which attach to a truncated body termination, are stout and smooth, each terminated by a single strong recurved claw. Each somite bears a pair of dorsal spines; somites 4 and posteriad bear an additional median spine. The spines on somites 1–3 are much shorter than the spines on the remaining somites. The head is short, bears a terminal mouth and a pair of antenniform outgrowths, and is covered by an oblong sclerite. Collinsovermis, plus Collinsium and Acinocricus, are found to comprise a sub-group of stout luolishaniid lobopodians with remarkably long spinules on the front lobopods, interpreted here as a clade (Teratopodidae fam. nov.) This clade is distinct from both the comparatively slenderer Luolishania and a sub-group composed of Facivermis and Ovatiovermis lacking body sclerites. Luolishaniids were mostly sessile forerunners of arthropods that had coupled efficient suspension-feeding devices and, as in Collinsovermis, strong defensive or deterrent features.     

山东寒武系苗岭统三叶虫消化系统的化石证据

三叶虫是寒武纪演化动物群中最引人注目的成员之一,其内部解剖结构一直以来受到广泛关注.与其他非生物矿化软体结构相比,三叶虫的消化系统更容易留下化石记录,为探索其内部结构提供了难得的机会.本文描述了来自山东省潍坊市寒武系馒头组的Proasaphiscus,Lioparia,Deiradonyx和Iranoleesia,以及...     

The Tetraconata concept: hexapod-crustacean relationships and the phylogeny of Crustacea

A growing body of evidence indicates that Crustacea and Hexapoda are sister groups, rather than Hexapoda and Myriapoda. Some recent molecular data even suggest that Mandibulata is not monophyletic, with Myriapoda and Chelicerata instead being sister groups. Here, arguments for homology of the mandible throughout mandibulate arthropods and for a monophyletic Mandibulata will be presented, as well as arguments supporting the taxon Tetraconata (i.e. Crustacea + Hexapoda). The latter include molecular data (nuclear and mitochondrial ribosomal RNAs and protein coding genes), and morphological characters such as ommatidial structure, the presence of neuroblasts and a very similar axonogenesis of pioneer neurons. However, crustaceans are insufficiently sampled for the molecular data, and studies of neurogenesis are lacking for many crustacean taxa. Remipedia, Cephalocarida and Maxillopoda are particularly problematic. This is important for the entire problem, because monophyly of the Crustacea has not yet been proven beyond doubt and several molecular analyses suggest a paraphyletic Crustacea. Here, arguments for the monophyly of the Crustacea are reviewed and two alternatives for the relationships between the five higher taxa Remipedia, Cephalocarida, Maxillopoda, Branchiopoda and Malacostraca are discussed: the Entomostraca concept sensu Walossek with Malacostraca as sister group to Cephalocarida, Maxillopoda and Branchiopoda, and the Thoracopoda concept sensu Hessler with Cephalocarida, Branchiopoda and Malacostraca forming a monophylum.     

Early Cambrian Pentamerous Cubozoan Embryos from South China

Background

Extant cubozoans are voracious predators characterized by their square shape, four evenly spaced outstretched tentacles and well-developed eyes. A few cubozoan fossils are known from the Middle Cambrian Marjum Formation of Utah and the well-known Carboniferous Mazon Creek Formation of Illinois. Undisputed cubozoan fossils were previously unknown from the early Cambrian; by that time probably all representatives of the living marine phyla, especially those of basal animals, should have evolved.

Methods

Microscopic fossils were recovered from a phosphatic limestone in the Lower Cambrian Kuanchuanpu Formation of South China using traditional acetic-acid maceration. Seven of the pre-hatched pentamerous cubozoan embryos, each of which bears five pairs of subumbrellar tentacle buds, were analyzed in detail through computed microtomography (Micro-CT) and scanning electron microscopy (SEM) without coating.

Results

The figured microscopic fossils are unequivocal pre-hatching embryos based on their spherical fertilization envelope and the enclosed soft-tissue that has preserved key anatomical features arranged in perfect pentaradial symmetry, allowing detailed comparison with modern cnidarians, especially medusozoans. A combination of features, such as the claustrum, gonad-lamella, suspensorium and velarium suspended by the frenula, occur exclusively in the gastrovascular system of extant cubozoans, indicating a cubozoan affinity for these fossils. Additionally, the interior anatomy of these embryonic cubozoan fossils unprecedentedly exhibits the development of many new septum-derived lamellae and well-partitioned gastric pockets unknown in living cubozoans, implying that ancestral cubozoans had already evolved highly specialized structures displaying unexpected complexity at the dawn of the Cambrian. The well-developed endodermic lamellae and gastric pockets developed in the late embryonic stages of these cubozoan fossils are comparable with extant pelagic juvenile cubomedusae rather than sessile cubopolyps, whcih indicates a direct development in these fossil taxa, lacking characteristic stages of a typical cnidarian metagenesis such as planktonic planula and sessile polyps.     

Reflections on the Phylogenetic Position of the Cephalocarida

This essay re-evaluates the phylogenetic position of the Cephalocarida in the light of the recently discovered Orsten (Upper Cambrian) crustaceans, the living Remipedia, and new interpretation of the Paleozoic ‘trilobitomorphs’. The Orsten crustaceans reinforce the belief that cephalocarid external morphology is primitive, except to show that the early crustacean trunk limb was almost surely biramous. The epipod appears to be a synapomorphy of the Cephalocarida, Branchiopoda and Malacostraca, thus justifying the existence of the Thoracopoda, which is coequal to the Maxillopoda and Remipedia. External morphology of the Remipedia is too specialized to support the hypothesis that it approximates that of the urcrustacean. Crustaceans and chelicerates are still best regarded as subgroups of the Schizoramia.     

A Parvancorina-like arthropod from the Cambrian of South China

Constraining the origin of animal groups is allowed, to some extent, by discoveries of Cambrian Lagerstätten that preserve both mineralizing and nonmineralizing organisms. A new species is reported here of the Cambrian arthropod Skania, which bears an exoskeleton that shares homologies with the Neoproterozoic (Ediacaran) organism Parvancorina and firmly establishes a Precambrian root for arthropods. A new monophyletic group, Parvancorinomorpha, is proposed as the first clade within the arthropod crown group demonstrably ranging across the Neoproterozoic–Paleozoic transition. The Parvancorinomorpha is interpreted to be the sister group of the Arachnomorpha. Incipient cephalization in Skania and related genera represents a step in the progression toward division of a cephalon from a large posterior trunk as shown in Cambrian arachnomorphs such as naraoiids and the addition of a pygidium and thoracic tergites as shown in the arachnomorph clade basal to trilobites. This evidence can serve as a new calibration point for estimating the divergence time for the last common ancestor of arthropods and priapulids based on molecular clock methods.     

The brachiopod <Emphasis Type="Italic">Eoobolus</Emphasis> from the Early Cambrian Mural Formation (Canadian Rocky Mountains)

The Early Cambrian brachiopod, Eoobolus, is one of the first representatives of the superfamily, Linguloidea, the defining characteristics of which include the classical morphology of oval shells and a pedicle that emerges from between the two valves. The material described here from the Mural Formation (Jasper National Park, Canadian Rocky Mountains) provides well-preserved muscle scars and larval shells that allow a discussion of the muscle system and the larval morphology of Eoobolus. The dorsal larval shell exhibits a morphology similar to other Cambrian linguloids, but also to paterinids, Mickwitzia muralensis, and some rhynchonelliforms. This suggests that there was a lesser degree of disparity among brachiopod larvae in the Cambrian than there is today. The muscle system of Eoobolus is similar to other linguloids, but differs from that of Recent lingulids and discinids by having one or two more pairs of oblique muscles. New data on the distribution of features characteristic of the family Eoobolidae question the validity of this family.     

Dietary partitioning by five sympatric species of stingray (Dasyatidae) on coral reefs

Dietary characteristics and the degree of dietary partitioning by five species of sympatric stingray were assessed using stomach content and sediment analyses within a coral reef lagoon at Ningaloo Reef, Western Australia (the cowtail Pastinachus atrus, blue‐spotted fantail Taeniura lymma, blue‐spotted mask Neotrygon kuhlii, porcupine Urogymnus asperrimus rays and the reticulate whipray Himantura uarnak). A total of 2804 items were recovered from the stomachs of 170 rays and 3215 individual taxa from the environment, which were used in selectivity analyses. Twenty‐four prey taxa were identified from stomach contents and pooled into 10 taxonomic categories for analysis, of which annelids, prawns, brachyurans and bivalves were the most abundant, together accounting for 96% of the diet. Himantura uarnak had the greatest interspecific dissimilarity in diet, consuming a larger proportion of crustaceans, notably penaeids (41% of total diet) than the other four species of rays, all of which had diets dominated by annelids (71–82% of total diet). Crustacean specialization by H. uarnak may exist to maximize resources and reduce competition among sympatric species. The remaining species may partition resources on the basis of space, rather than diet.