A test of whether rates of speciation were unusually high during the Cambrian radiation

The Cambrian radiation represents an interval when nearly 20 animal phyla appear in the fossil record in a short geological time span; however, whether this radiation also represents a period of extremely rapid speciation remains unclear. Here, a stochastic framework is used to test the null hypothesis that diversity changes in one of the dominant Early Cambrian groups, the olenelloid trilobites, could be produced by tempos of speciation known to have operated during later time periods. Two continuous-time models, the Yule model and the birth and death process model, and one discrete-time model, the Bienaymé-Galton-Watson branching process model, were used. No statistical evidence for uniquely high rates of speciation during the radiation in these trilobites was found when the continuous-time models were used with low or moderate extinction rates, the rates typically associated with the Cambrian radiation, although the p values are fairly low or, in one case, significant when high extinction rates were used. However, rates of speciation were higher than the average Phanerozoic rates of speciation. The discrete-time model produced equivocal results: either rates were unusually high or the model is inapplicable during the Cambrian radiation. This suggests that there was nothing unique about evolutionary processes relating to the tempo of speciation during the Cambrian radiation.     

The Cambrian radiation of bilaterians: Evolutionary origins and palaeontological emergence; earth history change and biotic factors

Evidence from a variety of research areas, including phylogenetic palaeobiogeographic studies of trilobites, indicates that there may be a fuse to the Cambrian radiation, with a duration on the order of 20–70 myr. Evolution in trilobites appears to have been powerfully influenced by the tectonic changes occurring at the end of the Neoproterozoic: especially the breakup of Pannotia. This continental fragmentation may have also elevated opportunities for vicariance and speciation in trilobites, and other metazoans, given that speciation rates at this time period were high, though not phenomenally so. This provides clear evidence that abiotic factors played an important role in motivating evolution during this key episode in the history of life; biotic factors probably also played a role. The evidence for the role of biotic factors is considered in light of information from some problematic Cambrian taxa. These may show affinities with modern problematic pseudocoelomate phyla, although Cambrian and modern exponents differ dramatically in body size.     

中国寒武纪三叶虫生物地理趋议

寒武纪三叶虫除北疆和藏南尚无可靠报道外,在中国各地均有广泛分布。三叶虫证据表明,除兴安区的额尔古纳一兴安地体和早寒武世中天山一北山地体显示了与西伯利亚和劳伦陆块在动物群方面的密切相关外,中国寒武纪的所有板块和大部分地体（包括中及晚寒武世的中天山～北山地体）均系当时东泛冈瓦纳的重要组成部分,在生物地理上彼此密切关联。对这些属于东泛冈瓦纳的中国区块的生物地理格局的进一步探讨,主要侧重于它们台地相浅水三叶虫动物群之间的对比,特别是立足于来自动物群纪录比较完整的华南、塔里木和华北板块的证据。综合分析研究表明这些板块和有关的地体在中及晚寒武世应归属同一生物地理区,而在早寒武世则可划分为两个生物地理亚区：一个由华南、塔里木板块和安南或印支地体组成,另一仅以华北板块为代表。     

A tiny eye indicating a planktonic trilobite

Abstract: Cambrian trilobites mainly lived on the sea floor, and up till now few, if any, unequivocally planktonic trilobites have been reported from earlier than the Ordovician. The late Cambrian (Furongian) to late Ordovician olenids are a distinctive group of benthic (sea‐floor dwelling) or nekto‐benthic trilobites. Here we show, however, that one recently described, miniaturized and very spiny olenid species, Ctenopyge ceciliae must have been planktonic (passively drifting or feebly swimming in the upper waters of the sea). This interpretation is based not only upon body form but also on the analysis of its visual system and may be one of the earliest records of the planktonic realm being invaded by trilobites.     

Contrasting patterns of disparity suggest differing constraints on the evolution of trilobite cephalic structures during the Cambrian ‘explosion’

Trilobites are an abundant group of Palaeozoic marine euarthropods that appear abruptly in the fossil record c. 521 million years ago. Quantifying the development of morphological variation (or ‘disparity’) through time in fossil groups like trilobites is critical in understanding evolutionary radiations such as the Cambrian ‘explosion’. Here, I use geometric morphometrics to quantify ‘cumulative disparity’ in functionally-important structures within the trilobite cephalon across their initial radiation during Cambrian Series 2. Overall cephalic disparity increased rapidly and attained a maximum within several million years. This pattern is dominated by the cephalic outline (in particular the genal spines), reflecting rapid, convergent expansion to the extremes of morphospace in a few early families. In contrast, removing the outline and focusing on structures such as the glabella and eye ridges (associated with feeding and vision, respectively) showed a more gradual increase in disparity, closer in line with taxonomic diversity and supporting the hypothesis of a relatively accurate trilobite fossil record. These contrasting patterns suggest that disparity in different structures was constrained in different ways, with extrinsic (ecological) factors probably having the major impact on overall disparity. It also implies that patterns of disparity in isolated substructures cannot necessarily be taken individually as representative of overall morphologies.     

Origin of planktotrophy--evidence from early molluscs

The size of early ontogenetic shells (protoconchs) of ancient benthic molluscs suggests that feeding larvae occurred at about 490 myr (approximately, transition from Cambrian to Ordovician). Most studied Ordovician protoconchs were smaller than Cambrian ones, indicating smaller Ordovician eggs and hatchlings. This suggests substitution of nutritious reserve matter such as yolk by plankton as an energy source for larvae. The observed size change represents the first direct empiric evidence for a late Cambrian to Ordovician switch to planktotrophy in invertebrate larvae. It corroborates previous hypotheses about a possible polyphyly of planktotrophy. These hypotheses were primarily based on molecular clock data of extant clades with different types of larva, change in the overall body size, as well as increasing predation pressure on Early Paleozoic sea floors. The Early Ordovician is characterized by an explosive radiation of benthic suspension feeders and it was suggested that planktotrophy would prolongate escape from benthic predation on hatchlings. This biological escalation hypothesis does not fully explain why planktotrophy and suspension feeding became important at the same time, during a major biodiversification. An additional factor that probably included availability of nutrients must have played a role. We speculate that an increasing nutrient supply and availability of photoautotrophic plankton in world oceans have facilitated both planktotrophy and suspension feeding, which does not exclude a contemporaneous predation-driven escalation. It is very likely that the evolution of planktotrophy as well as increasing predation contributed to the Ordovician radiation.     

The Phylogeny and Systematics of Blind Cambrian Ptychoparioid Trilobites

The paraphyletic trilobite suborder Ptychopariina includes a large proportion of Cambrian trilobite diversity and is probably ancestral to most groups of post-Cambrian trilobites. Resolution of the phylogenetic relationships within the group is therefore crucial to a better understanding of the initial radiation of trilobites. The recognition of approaches that can successfully resolve the relationships of ptychoparioid taxa is an important first step towards this aim. Cladistic analysis was used to determine relationships within the Cambrian ptychoparioid trilobite family Conocoryphidae, and to test claims that the family is polyphyletic. Ninety-seven characters were coded for 40 conocoryphid species and nine non-conocoryphids. The results indicate that the family consists of four distantly related clades. Three are recognized here as distinct families, including an extensively revised Conocoryphidae, and the families Holocephalidae and Atopidae. The fourth clade is referred to the subfamily Acontheinae (Corynexochida) as the new Tribe Hartshillini. Analysis of the disparity of these four clades shows that they are significantly less morphologically variable than the original polyphyletic taxon, demonstrating the possible effects of taxonomic error on macroevolutionary studies of morphological disparity.     

Burlingiid trilobites from Norway, with a discussion of their affinities and relationships

Four Middle and Upper Cambrian burlingiid trilobites from the Oslo Region, Norway, are described including Burlingia angusta sp. nov. from the Ptychagnostus punctuosus Zone and Schmalenseeia athrotryphe sp. nov. from the lower part of the Lejopyge laevigata Zone. New complete material previously attributed to Schmalenseeia jagoi Whittington is assigned to Burlingia . Cladistic analysis supports the genera Burlingia and Schmalenseeia as currently understood, including the placement of the controversial middle Middle Cambrian Schmalenseeia acutangula Westergård in Schmalenseeia , even though it lacks typical characters of the genus such as the median ridge on the preglabellar field. The analysis also supports burlingiid monophyly, and suggests that Schmalenseeia was derived from a broadly Burlingia –like ancestor, with S. acutangula displaying how the transition may have occurred. The broader relationships of Burlingia remain obscure, although similarities between burlingiids and the arthropod Kleptothule from the Early Cambrian Sirius Passet fauna are discussed: these include overall form, lack of functional hinges in the thorax, and details of the cephalic region. It is unclear whether these similarities represent general progenetic features, are functional convergences or, less likely, represent a genuine relationship.     

Head segmentation of trilobites

Although trilobites have provided research subjects for more than two centuries, their head segmentation has remained unresolved. Four glabellar furrows (SO and S1–S3) marking the segmental boundaries are generally present in the cephalic axis, but there are trilobites with one more pair of furrows, the so‐called S4, in the cephalic axis, causing confusion in understanding trilobite head segmentation. Recent advances in developmental biology and palaeontology have shed light on the arthropod head problem, and thus, trilobite head segmentation can be reviewed in the light of this knowledge. Based on the information from the anatomy of exceptionally preserved trilobites and artiopodans closely related to trilobites, it is inferred that trilobite head contains five segments: the anteriormost ocular segment potentially associated with the hypostome, the antennal segment and the following three segments with walking legs. When present, the S4 furrows are situated where the eye ridges meet the cephalic axis of trilobites, indicating that the furrows are incised ‘within’ the anteriormost segment in trilobites with an anteriorly enlarged frontal lobe. Trilobites of the Order Redlichiida, the most primitive stock, show variable conditions in the frontal glabellar conditions, while in other more derived groups, the condition is rather constant. The frontal glabellar condition, therefore, could provide a clue to elucidate the unresolved Cambrian trilobite phylogeny and the Cambrian roots of the post‐Cambrian trilobites.     

The Cambrian evolutionary 'explosion': decoupling cladogenesis from morphological disparity

The origin and differentiation of major clades is often assumed to have occurred in tandem with the 'explosion' of fossil evidence of diverse morphologies ('disparity') at the base of the Cambrian. Evidence is presented that this was not the case. Biogeographical and morphological differentiation among the earliest trilobites reveals incompleteness in the known early Cambrian record; similar evidence can be accrued for other major groups. Phylogenetic analysis proves the likelihood of 'ghost' lineages extending into the Precambrian. The important events in the generation of clades were earlier than the Cambrian 'explosion', at which time the groups become manifest in the fossil record. It is likely that the important phylogenetic changes happened in animals of small size; sister taxa of major groups are shown to be small animals. Decoupling cladogenesis from the Cambrian 'explosion' removes the necessity of invoking unknown evolutionary mechanisms at the base of the Phanerozoic. Genes controlling development may have played a role in generating new morphologies, through heterochrony for example, in the early differentiation of metazoan body plans.     

Abnormalities in early Paleozoic trilobites from central and eastern China

Malformations are common in trilobites, but the majority of described specimens are from Europe and North America. Only a few abnormal trilobites have been reported from China. Ten abnormal trilobites from Cambrian, Ordovician and Silurian strata in central and eastern China are documented. The abnormal Ordovician trilobites are found for the first time in China. All malformations occur in the thoraxes and pygidia, and were caused by a sub-lethal predatory attack, genetic or embryological malfunction, or injury sustained during molting. It is difficult to identify the predators of the six injured trilobites, but potential predators include Cambrian non-trilobite arthropods, Ordovician cephalopods, and Silurian eurypterids, chondrichthyes or cephalopods, even cannibalistic trilobites. Abnormal specimens caused by sub-lethal predatory attacks mainly occur in Cambrian strata in China and other areas in the world, and are rare in post-Cambrian strata. This pattern may reflect the rise of predators and increased predation in the post-Cambrian, which led to an increased trilobite fatality rate, thus reducing the probability that injured specimens would become fossilized.     

New bradoriids from the Lower Cambrian Yanwangbian formation of southern Shaanxi Province,Central China

Located in the northwest margin of the South China Block during the Cambrian, the Lower Cambrian Yanwangbian Formation of southern Shaanxi Province, Central China, has yielded a diverse fossil assemblage that includes abundant bradoriid arthropods, inarticulate brachiopods, hyolithids, trilobites, priapulids, lightly sclerotized arthropods, and ichnofossils. This assemblage represents the biodiversity of Early Cambrian marine communities in South China. Here we describe two new bradoriids from this stratigraphic unit, Sanlangella xixiangensis n. gen. et n. sp. and Kunmingella pentagona n. sp. On the basis of its carapace morphology, S. xixiangensis is assigned to the family Comptalutidae Öpik. The specimens of this species are preserved either in clay minerals or in calcium phosphate. Scanning electron microscopy analysis of phosphatized specimens reveals the microscopic reticulate surface ornaments that form a polygonal pattern on the anterodorsal node, around the base of mid-dorsal spine and along the lateroadmarginal rim. K. pentagona is described as the third species of this preexisting genus, based on the eight specimens with a distinctive pentagonal outline, a prominent anterodorsal curve and an anterodorsal sulcus.     

Asymmetry of Early Paleozoic trilobites

Asymmetry in fossils can arise through a variety of biological and geological mechanisms. If geological sources of asymmetry can be minimized or factored out, it might be possible to assess biological sources of asymmetry. Fluctuating asymmetry (FA), a general measure of developmental precision, is documented for nine species of lower Paleozoic trilobites. Taphonomic analyses suggest that the populations studied for each taxon span relatively short time intervals that are approximately equal in duration. Tectonic deformation may have affected the specimens studied, since deviations from normal distributions are common. Several measures of FA were applied to 3–5 homologous measures in each taxon. Measurement error was assessed by the analysis of variance (ANOVA) for repeated measurements of individual specimens and by analysis of the statistical moments of the distributions of asymmetry measures. Measurement error was significantly smaller than the difference between measures taken on each side of a specimen. However, the distribution of differences between sides often deviated from a mean of zero, or was skewed or kurtosic. Regression of levels of FA against geologic age revealed no statistically significant changes in levels of asymmetry through time. Geological and taphonomic effects make it difficult to identify asymmetry due to biological factors. Although fluctuating asymmetry is a function of both intrinsic and extrinsic factors, the results suggest that early Cambrian trilobites possessed genetic or developmental mechanisms used to maintain developmental stability comparable to those of younger trilobites. Although the measures are biased by time averaging and deviations from the normal distribution, these data do not lend strong support to 'genomic' hypotheses that have been suggested to control the tempo of the Cambrian radiation.     

Cambrian agnostid communities in Tasmania

Jago, J. B.: Cambrian agnostid communities in Tasmania.
Two or possibly three different agnostid trilobite assemblages can be distinguished in the late Middle and early Upper Cambrian sequences of northern and western Tasmania. This is significant because in recent years agnostid trilobites have been widely used in local and international correlations of Middle and Upper Cambrian rocks. The three assemblages recognized are (1) an agnostid assemblage in which polymerid trilobites are abscnt, rare or present as thanatocoenotic fossils, (2) a ptychagnostid-non-nepeid assemblage, and (3) a nepeid-clavagnostid-peronopsid assemblage in which non-agnostid trilobites are abundant but ptychag-nostids are absent. It is proposed that assemblage (1) represents an open sea fauna, with assemblages (2) and (3) occurring in progressively shallower water.     

Two middle Cambrian diceratocephalid trilobites, Cyclolorenzella convexa and Diceratocephalus cornutus, from Korea: development and functional morphology

Park, T.-Y. & Choi, D.K. 2010: Two middle Cambrian diceratocephalid trilobites, Cyclolorenzella convexa and Diceratocephalus cornutus , from Korea: development and functional morphology. Lethaia , Vol. 43, pp. 73–87.
Silicified sclerites of the latest middle Cambrian trilobites, Cyclolorenzella convexa and Diceratocephalus cornutus , have been recovered from the Sesong Formation, Korea. Their morphological similarity and stratigraphic occurrences suggest that D. cornutus is a descendant of C. convexa . The ontogenies of both trilobites demonstrate that a pair of long frontal horns in the cephalon of D. cornutus is an evolutionarily novel structure. It is inferred that redeployment of some pre-existing regulatory gene played a significant role in constructing the frontal horns of D. cornutus . The frontal horns may have been a defensive structure to deter predators. The facial suture of D. cornutus , which extends onto the frontal horns and splits them into the dorsal and ventral halves, was a solution to enable easier forward egression during ecdysis. □ Functional morphology, Korea, Middle Cambrian, ontogeny, trilobites .     

安徽淮南早寒武世蠕虫化石的发现

Ｐａｌａｅｏｓｃｏｌｅｘｈｕａｉｎａｎｅｎｓｉｓｓｐ．ｎｏｖ．是我国华北地区寒武纪地层中首次发现的蠕虫化石，也是我国寒武纪地层中所发现的蠕虫化石的最高层位。     

THE AGE AND RELATIONSHIPS OF THE MAJOR ANIMAL PHYLA

Given the uncertainties in the fossil record and the paucity of informative morphological characters, there is still considerable uncertainty as to the phylogenetic affinities and times of origins of essentially all of the phyla of animals. A multilocus analysis of amino-acid sequence data for mitochondrial genes suggests that the major triploblast phyla began diverging approximately 630 million years ago. These results support the hypothesis that the so-called Cambrian radiation of animals actually initiated about 100 million years prior to the Cambrian, as the fossil evidence suggests. In addition, phylogenetic analysis supports the monophyly of animals, an early (~900 million years ago) branching off of the cnidarian lineage, the monophyly of deuterostomes and protostomes, and the inclusion of nematodes in the protostome lineage. The results of this study suggest that, with appropriate levels of taxon sampling and a focus on conserved regions of protein-coding sequence, complete mitochondrial genome analysis may be sufficiently powerful to elucidate the genealogical relationships of many of the animal phyla.     

Gut content fossilization and evidence for detritus feeding habits in an enrolled trilobite from the Cambrian of China

Enrolment was a major defensive strategy for trilobites that significantly contributed to their evolutionary success. The ability to enrol also helped to constrain the morphological evolution of trilobites, which in part was driven by the need to improve this capability to encapsulate the soft parts of the body within the mineralized dorsal exoskeleton. Here, we describe a unique example of gut content fossilization in an enrolled trilobite from the Cambrian of China, and we propose a taphonomic scenario that considers the possible implication of enrolment in this exceptional preservation. A micro‐facies analysis indicates that the specimen was entombed during an obrution event and remained intact due to limited infaunal activity. The encapsulation of the body did not prevent the decay of soft tissues, but it permitted the delicate gut content to be protected during diagenesis. In isolating the decaying soft tissues from the external environment, enrolment might also have favoured the establishment of microenvironmental conditions conducive to the precipitation of pyrite framboids. Within the gut, the formation of such crystals may have led to the consolidation of the ingested material. These results suggest that fossilized gut contents might be quite common in enrolled trilobites. Textural and compositional analyses reveal that the gut content is similar to the sediment surrounding the fossil except for the presence of pyrite framboids that indicate higher initial organic matter content. The complete enrolment of the body argues against an accidental ingestion of this material or a diagenetic origin for it. Accordingly, detritus feeding habits are inferred for this ptychopariid trilobite.     

The stability of thoracic segmentation in trilobites: a case study in developmental and ecological constraints

The decline in origination rate of new metazoan body plans following the Cambrian radiation has been suggested to reflect developmental canalization in derived taxa, limiting their ability to evolve forms with radically different morphotypes. Segmentation is a fundamental aspect of arthropod body plan, and here we show that a derived trilobite that secondarily converged on a morphotype characteristic of basal members of the clade also reverted to a pattern of segmental variability common among basal trilobites. Hence a secular trend in loss of variability of the trilobite thorax was not due to the evolution of an inviolable developmental constraint. This result challenges the notion of developmental canalization in phylogenetically derived taxa. Rather, early variability in trilobites may be the result of ecological factors that promoted segment-rich thoracic morphotypes during Cambrian time.     

New Lower Cambrian trilobites from Pleistocene erratic boulders of northern Germany and Denmark and their bearing on the intercontinental correlation

Pleistocene erratic boulders in northern Germany and Denmark yield upper Lower Cambrian ellipsocephaloid trilobites described asEpichalnipsus anartanus n. gen., n. sp. andBerabichia erratica n. sp. They are supposedly derived from an unidentified late Early Cambrian unit from central Sweden and most probably come from an otherwise poorly fossiliferous stratigraphie interval of theHolmia kjerulfi Zone. Although both species are new, their systematic affinities permit a refined intercontinental correlation between Baltica and western Gondwana. An erratum to this article is available at .