0000000000118167
AUTHOR
Rich Mooi
Phylogeny and origin of Jurassic irregular echinoids (Echinodermata: Echinoidea).
27 pages; International audience; A phylogenetic analysis of Jurassic irregular echinoids is realized to explore the origin and early evolution of this important subset of echinoids. The phylogeny is based on 39 characters and considers data from apical system architecture, the corona including tuberculation and spines, Aristotle's lantern, and general test shape. Results corroborate the monophyly of Irregularia, and clarify the phylogenetic interrelationships existing between the main groups of irregular echinoids. Specializations of the Aristotle's lantern, spines, tubercles and phyllodes constitute the apomorphies for different taxa, as for the whole of Irregularia. The phylogenetic sign…
Images are not and should not ever be type specimens: a rebuttal to GarraffoniFreitas.
Note. This original form of this rebuttal was submitted to Science on 3 March 2017 (limited to 300 words as per Science editorial policy) but rejected on 13 March 2017. Herein, we elaborate on our original Science submission in order to more fully address the issue without the length limitations. This rebuttal is followed by the list of the signatories who supported our original submission.
Athenacrinus n. gen. and other early echinoderm taxa inform crinoid origin and arm evolution
AbstractIntermediate morphologies of a new fossil crinoid shed light on the pathway by which crinoids acquired their distinctive arms. Apomorphies originating deep in echinoderm history among early nonblastozoan pentaradiate echinoderms distinguish Tremadocian (earliest Ordovician) crinoid arms from later taxa. The brachial series is separated from the ambulacra, part of the axial skeleton, by lateral plate fields. Cover plates are arrayed in two tiers, and floor plates expressed podial basins and pores. Later during the Early Ordovician, floor plates contacted and nestled into brachials, then were unexpressed as stereom elements entirely and cover plates were reduced to a single tier. Inco…
Echinoderm evolution since 1972 and since the cambrian: tales from a dozen IECS.
5 pages; International audience; Progress made since the first IEC - which took place in Washington, DC in 1972 has drastically altered some of our views on echinoderm evolution. In addition to publications in scientific journals, there are thousands of pages of peer-reviewed material in the proceedings of past IECs, and in Echinoderm Studies. These pages bear witness to some of the most outstanding milestones in this progress. This paper aims to highlight one third of a century of maturing ideas dealing with several key aspects of echinoderm evolution as witnessed by 12 IECs. Conceptual changes have taken place in a scientific milieu characterized by rapid adoption of new theoretical and p…
How Hox genes can shed light on the place of echinoderms among the deuterostomes.
19 pages; International audience; BACKGROUND: The Hox gene cluster ranks among the greatest of biological discoveries of the past 30 years. Morphogenetic patterning genes are remarkable for the systems they regulate during major ontogenetic events, and for their expressions of molecular, temporal, and spatial colinearity. Recent descriptions of exceptions to these colinearities are suggesting deep phylogenetic signal that can be used to explore origins of entire deuterostome phyla. Among the most enigmatic of these deuterostomes in terms of unique body patterning are the echinoderms. However, there remains no overall synthesis of the correlation between this signal and the variations observ…
Reappraisal of ambulacral branching patterns in blastozoans
6 pages; International audience; The type of ambulacral bifurcation is an important diagnostic character in primitive echinoderms. However this character is rarely used in blastozoan phylogenetic studies, whereas the corresponding character is well used in crinoid analyses. The main reason for this gap is the absence of definition of patterns for all blastozoans, resulting in terminological confusion. Although crinoid arms and blastozoan ambulacra are not homologous, the well-defined crinoid terminology has been used to describe the different topologies of ambulacral bifurcation. Unilateral and bilateral patterns have been recognized in blastozoans, and associated with the mineralization of…
Evolution to the extreme: origins of the highly modified apical system in pourtalesiid echinoids
The apical system of the genus Pourtalesia displays a plate architecture that falls so far outside that typical of other echinoids that plate homologies remain problematic. A new approach using the Extraxial–Axial Theory (EAT) that develops homologies for the Echinodermata is proposed. The exploration of apical plate patterns throughout ontogenetic sequences shows that the typical holasteroid pattern found in the youngest specimens undergoes a series of disturbances that result in a multiple disjunction accompanied by isolation or disappearance of certain genital plates. We propose a new interpretation of the apical architecture of the genus that agrees with: (1) the plate addition processe…
Are homalozoans echinoderms? An answer from the extraxial-axial theory
Homalozoans include four classes of non-pentamerous Paleozoic echinoderms: Homostelea (cinctans), Ctenocystoidea (ctenoid-bearing homalozoans), Homoiostelea (solutes), and Stylophora (cornutes and mitrates). Their atypical morphologies have historically made it difficult to relate them to other classes. Therefore, their systematic positions have been represented by two hypotheses (H): as stem taxa to echinoderms (H1) or as stem taxa to chordates (H2). These conclusions rest on previous inability to recognize synapomorphies with more crownward echinoderms, resulting in a forcing of the homalozoans down the phylogenetic tree that is more artifactual than evolutionary. The Extraxial-Axial Theo…
Exceptionally preserved soft parts in fossils from the Lower Ordovician of Morocco clarify stylophoran affinities within basal deuterostomes.
10 pages; International audience; The extinct echinoderm clade Stylophora consists of some of the strangest known deuterostomes. Stylophorans are known from complete, fully articulated skeletal remains from the middle Cambrian to the Pennsylvanian, but remain difficult to interpret. Their bizarre morphology, with a single appendage extending from a main body, has spawned vigorous debate over the phylogenetic significance of stylophorans, which were long considered modified but bona fide echinoderms with a feeding appendage. More recent interpretation of this appendage as a posterior “tail-like” structure has literally turned the animal back to front, leading to consideration of stylophorans…
Radial Symmetry, the Anterior/Posterior Axis, and Echinoderm Hox Genes
20 pages; International audience; The strangeness of echinoderm pentaradiality results from superposition of radial symmetry onto ancestral deuterostome bilaterality. The Extraxial- Axial Theory shows that echinoderms also have an anterior/posterior (A/P) axis developed independently and ontogenetically before radiality. The A/P axis is first established via coelomic stacking in the extraxial region, with ensuing development of the pentamerous hydrocoel in the axial region. This is strongly correlated with a variety of gene expression patterns. The echinoid Hox cluster is disordered into two different sets of genes. During embryogenesis, members of the posterior class demonstrate temporal, …
The morphology, ontogeny, and inferred behaviour of the deep-sea echinoid Calymne relicta (Holasteroida).
The deep-sea holasteroid Calymne relicta was first described from a few fragments discovered by the HMS ‘Challenger’ in the Bermuda abyssal plain more than a century ago. In addition to re-examining the type material, we describe herein new specimens from unpublished material collected between 3720 and 4860 m during three scientific expeditions that took place on both sides of the North Atlantic between 1966 and 1991. The new material includes juvenile and adult specimens in sufficiently good preservational state to allow a full redescription, including all types of appendages, some of which have never been described. These new observations confirm the atypical characteristics of C. relicta…
What a New Model of Skeletal Homologies Tells Us About Asteroid Evolution1
The Extraxial-Axial Theory (EAT) is applied to the body wall homologies of asteroids. Attempts to characterize major plate systems of asteroids as axial or extraxial, particularly those that are highly organized into series, can be problematic. However, the Optical Plate Rule (OPR) is instrumental in establishing that ambulacrals and terminals are axial. It is equally clear that the region aboral to the marginal frame is a part of the perforate extraxial body wall (with the possible exception of the centrodorsal, which is likely imperforate extraxial). Previously established EAT criteria, particularly those strongly rooted in the embryologically expressed boundary between axial and extraxia…
Arrays in rays: terminal addition in echinoderms and its correlation with gene expression
Summary The echinoderms are deuterostomes that superimpose radial symmetry upon bilateral larval morphology. Consequently, they are not the first animals that come to mind when the concepts of segmentation and terminal addition are being discussed. However, it has long been recognized that echinoderms have serial elements along their radii formed in accordance with the ocular plate rule (OPR). The OPR is a special case of terminal growth, forming elements of the ambulacra that define the rays in echinoderms. New elements are added at the terminus of the ray, which may or may not be marked by a calcified element called the terminal plate (the “ocular” of sea urchins). The OPR operates in eve…
Evolution Within a Bizarre Phylum: Homologies of the First Echinoderms
SYNOPSIS. The Extraxial/Axial Theory (EAT) of echinoderm skeletal homologies describes two major body wall types: axial and extraxial. The latter is subdivided into perforate and imperforate regions. Each of the regions has a distinctly different source in early larval development. Axial skeleton originates in the rudiment, and develops in association with the pentaradially arranged hydrocoel according to specific ontogenetic principles. Perforate and imperforate extraxial regions are associated with the left and right somatocoels respectively, are not governed by ontogenetic principles of plate addition, and are products of the non-rudiment part of the larval body. The morphology of even t…
Biodiversity of Antarctic echinoids: a comprehensive and interactive database
Eighty-one echinoid species are present south of the Antarctic Convergence, and they represent an important component of the benthic fauna. “Antarctic echinoids” is an interactive database synthesising the results of more than 100 years of Antarctic expeditions, and comprising information about all echinoid species. It includes illustrated keys for determination of the species, and information about their morphology and ecology (text, illustrations and glossary) and their distribution (maps and histograms of bathymetrical distribution); the sources of the information (bibliography, collections and expeditions) are also provided. All these data (taxonomic, morphologic, geographic, bathymetri…
La radiation des échinodermes au Paléozoïque inférieur, l'exemple des blastozoaires.
10 pages; Le sous-phylum Blastozoa est un des groupes d'échinodermes les plus diversifiés (dix classes) au début du Paléozoïque. Après révision critique de leur squelette, leurs morphologies en apparence très variées sont en fait homogènes. Leur diversité montre deux pics (Drumien, Sandbien) liés par un événement de fortes apparitions génériques au Cambrien supérieur-Ordovicien inférieur. Les blastozoaires montrent un fort endémisme au Cambrien et un important provincialisme à l'Ordovicien inférieur et moyen. Ils deviennent cosmopolites à l'Ordovicien supérieur, par plusieurs événements migratoires. Ils sont restreints à la Laurentia et à Baltica au Silurien inférieur.