Nipping the Cambrian "explosion" in the bud?

In recent years, two schools of thought have emerged with regard to the Cambrian "explosion". One argues that it was very quick, with phyla tumbling into existence in a virtual geological instant. The other view has a more relaxed temporal perspective. It looks to slow aeons of cryptic metazoan history, which led to a final breakthrough in the Cambrian, not in evolution but of fossilization potential. Yet both views have serious difficulties. Now, in a recent issue of Biological Reviews, Graham Budd and S?ren Jensen(1) argue for a third way. In an intriguing blend of functional morphology, the fossil record and cladistic thinking, they suggest that the assembly of metazoan bodyplans took place in a surprisingly straightforward manner.     

Integrin and cadherin clusters: A robust way to organize adhesions for cell mechanics

The evolutionary emergence of animals is one of the most significant episodes in the history of life, but its timing remains poorly constrained. Molecular clocks estimate that animals originated and began diversifying over 100 million years before the first definitive metazoan fossil evidence in the Cambrian. However, closer inspection reveals that clock estimates and the fossil record are less divergent than is often claimed. Modern clock analyses do not predict the presence of the crown‐representatives of most animal phyla in the Neoproterozoic. Furthermore, despite challenges provided by incomplete preservation, a paucity of phylogenetically informative characters, and uncertain expectations of the anatomy of early animals, a number of Neoproterozoic fossils can reasonably be interpreted as metazoans. A considerable discrepancy remains, but much of this can be explained by the limited preservation potential of early metazoans and the difficulties associated with their identification in the fossil record. Critical assessment of both records may permit better resolution of the tempo and mode of early animal evolution.     

The origin of animal phyla and the new phylum Procoelomata

BergstrÖm, Jan 1989 07 15: The origin of animal phyla and the new phylum Procoelomata. Lethaia , Vol. 22, pp. 259–269. Oslo. ISSN 0024–1164.
A model of metazoan evolution presented previously (BergstrÖm 1986 in Zoologica Scripta 15 ) explains deuterostomian characters as derived from protostomian ones through loosening of the constraints in the spiralian type of morphogenesis. This fits phylogenies derived from studies of molecular sequences. The model helps explain (1) the well-known mixture of proto- and deuterostomian features in several groups; (2) the difficulties in making a phylogeny based on comparative anatomy, and (3) the fossil explosion in the Cambrian. Since protostomian features such as a ciliated locomotory sole and a pelagic larva with ciliary bands are widely distributed in branches of the phylogenetic tree, they must have been present in the stem of the tree. Most probably the stem forms were pseudosegmented, which helps explain how segmentation, oligomery and non-segmentation could evolve repeatedly in derived groups. Origination of new phyla involved macroevolutionary changes primarily in the mode of feeding and locomotion. The stem phylum, from which most other phyla appear to have been derived directly, is here named the Procoelomata. Machaeridian-type animals are referred to it. The Ediacaran-type Precambrian fossils Cannot be placed in the metazoan evolutionary tree. * Biochemical evolution, Cambrian fossil explosion, Deuterostomia, eukaryote evolution, Machaeridia. macroevolution, Precambrian fossils. Procoelomata, Protostomia .     

Vestiges of a beginning? Paleontological and geochemical constraints on early animal evolution

Inspired by molecular clock estimates, some biologists have proposed that animals in general, and bilaterian metazoans in particular, began to diverge substantially earlier than fossils indicate. Balavoine and Adoutte [Science 280 (1998) 397] specifically hypothesized that the Cambrian explosion documents parallel radiations within three major bilaterian clades that diverged from one another relatively early in the Neoproterozoic Era. The geological record is permissive with respect to such hypotheses, but not encouraging. The earliest evolution of animals certainly took place before the initial appearance of phosphatic animal microfossils or Ediacaran macrofossils, but bilaterian clades need not have been part of this metazoan “pre-history”. Alternatively, the evolution of large size, made possible by late Neoproterozoic oxygen increase, may have provided the selective environment in which stem bilaterians differentiated. Cambrian events per se appear to have begun in the wake of environmental perturbation and accompanying extinction near the Proterozoic–Cambrian boundary.     

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.     

From Vendian to Cambrian: the beginning of morphological disparity of modern metazoan phyla

Continuity of the transition from Precambrian to the Phanerozoic metazoan fauna at the phylum level is analyzed. The discrete traces of feeding on bacterial mats by Dickinsonia and similar organisms are explained by extracorporeal digestion, characteristic of a placozoan level of organization, as in the extant Trichoplax. The absence of a morphologically developed anterior end of the body, of food-gathering appendages of any kind, and of appendages responsible for movement suggest that these animals were at this level of organization. Probably, an expanded placozoan level of organization can be assumed for most Vendian animals. Against this background, new characters emerged in the Cambrian to be included in the body plan of extant animal phyla. The relationships between the morphological archetype (body plan) and morphogenetic archetype are considered. It is shown that major features of a morphogenetic archetype could be retained from the time the taxon was formed.     

THE IMPORTANCE OF PREADAPTED GENOMES IN THE ORIGIN OF THE ANIMAL BODYPLANS AND THE CAMBRIAN EXPLOSION

The genomes of taxa whose stem lineages branched early in metazoan history, and of allied protistan groups, provide a tantalizing outline of the morphological and genomic changes that accompanied the origin and early diversifications of animals. Genome comparisons show that the early clades increasingly contain genes that mediate development of complex features only seen in later metazoan branches. Peak additions of protein‐coding regulatory genes occurred deep in the metazoan tree, evidently within stem groups of metazoans and eumetazoans. However, the bodyplans of these early‐branching clades are relatively simple. The existence of major elements of the bilaterian developmental toolkit in these simpler organisms implies that these components evolved for functions other than the production of complex morphology, preadapting the genome for the morphological differentiation that occurred higher in metazoan phylogeny. Stem lineages of the bilaterian phyla apparently required few additional genes beyond their diploblastic ancestors. As disparate bodyplans appeared and diversified during the Cambrian explosion, increasing complexity was accommodated largely through changes in cis‐regulatory networks, accompanied by some additional gene novelties. Subsequently, protein‐coding genic richness appears to have essentially plateaued. Some genomic evidence suggests that similar stages of genomic evolution may have accompanied the rise of land plants.     

EDIACARAN AND CAMBRIAN INDEX FOSSILS FROM SONORA, MEXICO

Abstract:  The Cambrian index fossil Treptichnus pedum is reported from the Puerto Blanco Formation near Pitiquito, Sonora, Mexico, and new occurrences of the Neoproterozoic index fossil Cloudina are reported from the underlying La Ciénega Formation. Considered together, these fossils constrain the location of the Ediacaran/Cambrian boundary in Mexico. The suite of fossils in this sequence is important because it provides an independent biostratigraphic datum for constraining the position of Laurentia during the onset of metazoan diversification, for testing proposed lithostratigraphic correlations among western North American Neoproterozoic–Cambrian successions, and for testing regional tectonic paradigms such as the Mojave-Sonora Megashear hypothesis.     

Eggs and embryos from the Cambrian

The early evolution of metazoans is a major focus of biological attention, but is the historical record revealed in the Cambrian “explosion” an accurate reflection of original events? The key questions concern the nature of the earliest animals and when they originated. One widely-mooted suggestion is that planktotrophic larvae, typified by the annelidan trochophore and echinoid pluteus, existed long before the metazoan radiations evident in the Cambrian fossil record. This idea could be consistent for recent evidence of divergence times, based on molecular “clocks,” of phyla appearing well before the Cambrian. Now a surprising new discovery of eggs with blastomeres and embryos with well-defined anatomy from the Cambrian (c.530 Myr ago) of China and Siberia promises to extend the arena of debate. In one case a convincing ontogeny can be traced from eggs to adult tube-dwelling cnidarians. In the other example a possible protostome, unhatched and wrapped around the egg, shows segmentation and possibly nascent sclerites. In both, these cases development is direct, i.e., there is no evidence for any planktotrophic larval stage. The implications for our perceptions of both the Cambrian ‘explosion’ and metazoan phylogeny could be considerable. BioEssays 20 :676–682, 1998.© 1998 John Wiley & Sons Inc.     

Morphological and developmental macroevolution: a paleontological perspective

Evidence of the morphological evolution of metazoans has been preserved, in varying degrees of completeness, in the fossil record of the last 600 million years. Although extinction has been incessant at lower taxonomic levels, genomic comparisons among surviving members of higher taxa suggest that much of the developmental systems that pattern their bodyplans has been conserved from early in their history. Comparisons between the origin of morphological disparity in the record and patterns of genomic disparity among living taxa promise to be interesting. For example, Hox cluster composition varies among major taxa, and the fossil record suggests that many of the changes in Hox clusters may have been associated with late Neoproterozoic evolution among minute benthic vermiform clades, from which crown bilaterian phyla arose just before or during the Cambrian explosion. Study of genomic differences among crown classes and orders whosetiming and mode of origin can be inferred from morphological data inthefossil record should throw further light on the timing and mode of origin of genomic disparities.     

Palaeoenvironmental interpretation of microphytoplankton diversity trends in the Cambrian–Ordovician of the northern Sahara Platform

Analysis of vertical fluctuations in assemblage diversity and relative abundances of microphytoplankton morphological classes permits palaeoenvironmental reconstructions of the Cambrian–Ordovician stratal sequences in the Hassi-R'Mel area (north-central Algeria) and northern Rhadames Basin (southern Tunisia). Abundance fluctuations of acritarch morphological classes and assemblage diversity appear to be related to changes in depositional facies. The present palaeoenvironmental interpretation is primarily based on a comparison with microphytoplankton distributional trends in modern depositional environments. Where possible, the observations are tested against palaeoenvironmental information derived independently from previous lithostratigraphical, macropalaeontological, and petrographical studies of the formations investigated. The present data support the following palaeoenvironmental scenario: during Late Cambrian times, relatively shallow water, neritic conditions prevailed throughout the study area, with evidence for a more proximal environment in southern Tunisia. In the early Tremadoc, open marine, offshore conditions characterized the central Algerian region, but a shallower water setting is indicated for southern Tunisia. During the late Arenig–Llanvirn interval, an open marine, offshore setting is envisaged for the southern Tunisian region, with more proximal conditions prevailing in the Algerian Hassi-R'Mel area. During the entire Ashgill, an essentially restricted marine environment was established throughout the study area. Some stratigraphically important acritarch species (e.g. Acanthodiacrodium angustum) appear to be facies-sensitive; the recognition of palaeoecological control on microphytoplankton distribution has important consequences on the biostratigraphic application of these microfossils.     

Molecular clocks, molecular phylogenies and the origin of phyla

Erwin, Douglas H. 1989 07 15: Molecular clocks, molecular phylogenies and the origin of phyla. Lethaia , Vol. 22, pp. 251–257. Oslo. ISSN 0024–1164.
Protein, RNA and DNA sequences have been widely used to construct phylogenies and to calculate divergence times using a molecular clock. Reliance on molecular information is particularly attractive when fossil evidence is missing or equivocal, as in the Cambrian metazoan radiation. I consider the applicability of molecular clocks and phylogenetic analysis of molecular data to the origin of metazoan phyla, and conclude that molecular information is often ambiguous or misleading. Amino acid sequences are of limited use because the redundancy of the genetic code masks patterns of descent, while in a nucleotide sequence only four potential states exist at each site (the four nucleotide bases). In each case, homoplasy may often go undetected. The application of a molecular clock to resolve the timing of the metazoan radiation is unwarranted, while molecular phylogenetic reconstruction should be approached with care. A potentially more useful technique for phylogenetic reconstruction would be the use of patterns of genome structure and organization as characters. * Molecular clock, phylogenetics, metazoan radiation, origin of phyla .     

Receptor tyrosine kinase, an autapomorphic character of metazoa: identification in marine sponges

In the present review we summarize sequence data obtained from cloning of sponge receptor tyrosine kinases [RTK]. The cDNA sequences were mainly obtained from the marine sponge Geodia cydonium. RTKs (i) with immunoglobulin [Ig]-like domains in the extracellular region, (ii) of the type of insulin-like receptors, as well as (iii) RTKs with one extracellular speract domain, have been identified. The analyses revealed that the RTK genes are constructed in blocks [domains], suggesting a blockwise evolution. The phylogenetic relationships of the sequences obtained revealed that all sponge sequences fall into one branch of the evolutionary tree, while related sequences from higher Metazoa, human, mouse and rat, including also invertebrate sequences, together form a second branch. It is concluded that the RTK molecules have evolved in sponges prior to the "Cambrian Explosion" and have contributed to the rapid appearance of the higher metazoan phyla and that sponges are, as a taxon, also monophyletic. Due to the fact that protein tyrosine kinases in general and RTKs in particular have only been identified in Metazoa, they are, as a group qualified, to be considered as an autapomorphic character of all metazoan phyla.     

Avian diversity and behavior in an Eocene coastal plain,Svalbard: the ichnological evidence

Abstract

This study presents an ichnotaxonomical assessment of bird footprints from a coastal plain setting in the Eocene lower to middle Aspelintoppen Formation in Brongniartfjellet and Storvola, Svalbard. These footprints are unique evidence for Paleogene birds from Svalbard and the second evidence of Paleogene avifauna of the Arctic (along with scarce fossil remains from Ellesmere Island). The analyzed footprints are assigned to six ichnotaxa belonging to three ichnofamilies: Aquatilavipes isp., avian footprint morphotype A and B (ichnofamily Avipedidae); Gruipeda cf. G. abeli, Gruipeda cf. G. dominguensis (ichnofamily Gruipedidae); and Gyeongsangornipes isp. (ichnofamily unknown). Associated invertebrate ichnofossils include Helminthoidichnites tenuis, Cochlichnus anguineus, and Helminthopsis isp. The avian footprints reflect small, medium-sized and rare large birds that can be morphologically compared to those of modern relatives including crane, heron, plover, moorhen, gallinule, oystercatcher and curlew. It is also proposed that in situ sets of shorebird footprints showing a preferred orientation, not composing trackways and showing overprinting, can be used as an indicator of the position and orientation of the shoreline.     

陕南早寒武世早期Quadrapyrgites再研究

处于"寒武纪生命大爆发"序幕阶段的梅树村期,生物类群大规模辐射,身体构型快速革新,与前寒武纪生物群面貌明显不同.最近在陕南宁强宽川铺地区梅树村期地层中发现了大量五辐射Punctatus及部分四辐射四方塔形壳属Quadrapyrgites,其中包括1个新种Quadrnpyrgites undulatuscostalis sp.nov..在此基础上对Quadrapyrgites进行了属征补充.双胚层腔肠动物的出现标志着地球生命史的真后生动物演化开端,在生物起源演化历程上占据着关键的位置.本文为研究真后生动物起源演化、生物辐射、体型构建提供了重要实证.     

Colour in Burgess Shale animals and the effect of light on evolution in the Cambrian

Diffraction gratings are reported from external surfaces of the hard, protective parts of Wiwaxia corrugata, Canadia spinosa and Marrella splendens from the Burgess Shale (Middle Cambrian (515 million years), British Columbia). As a consequence, these animals would have displayed iridescence in their natural environment: Cambrian animals have previously been accurately reconstructed in black and white only. A diversity of extant marine animals inhabiting a similar depth to the Burgess Shale fauna possess functional diffraction gratings. The Cambrian is a unique period in the history of animal life where predatory lifestyles and eyes capable of producing visual images were evolving rapidly. The discovery of colour in Cambrian animals prompts a new hypothesis on the initiation of the ''Big Bang'' in animal evolution which occurred during the Cambrian: light was introduced into the behavioural systems of metazoan animals for the first time. This introduction, of what was to become generally the most powerful stimulus in metazoan behavioural systems, would have triggered turbulence in metazoan evolution.     

The cambrian evolutionary ‘explosion’ recalibrated

The sudden appearance in the fossil record of the major animal phyla apparently records a phase of unparalleled, rapid evolution at the base of the Cambrian period, 545 Myr ago. This has become known as the Cambrian evolutionary ‘explosion’, and has fuelled speculation about unique evolutionary processes operating at that time. The acceptance of the palaeontological evidence as a true reflection of the evolutionary narrative has been criticised in two ways: from a reappraisal of the phylogenetic relationships of the early fossils, and from predicitions of molecular divergence times, based on six appropriate metazoan genes. Phylogenetic analysis of the arthropods implies an earlier, Precambrian history for most clades, and hence an extensive period of cladogenesis unrecorded by fossils. A similar argument can be applied to molluscs, lophophorates and deuterostomes. Molecular evidence implies divergence between clades to at least 1000 Myr ago. The apparent paradox between the sudden appearance of recognisable metazoans and their extended evolutionary history might be explained by a sudden Cambrian increase in body size, which was accompanied by skeletisation. A new paradigm suggests that the ‘explosion’ in the record may have been decoupled from the evolutionary innovation.