Ordovician Paracrinoids from the Baltic: Key Problems of Comparative Morphology of Pelmatozoan Echinoderms

The erect feeding appendages of paracrinoids, brachioles of typical blastozoans and arms of crinoids are morphologically similar in their terminal growth, biserial cover plates, and pinnulation. This is attributed to the inducing effect of the radial ambulacral canal on their growth mode. The uniserial brachioles of Laurentian paracrinoids are homologous to the biserial brachioles of the Baltic Achradocystites and Heckerites, and those of other blastozoans. Based on this assumption, the two Baltic genera, which have a brachiole system plesiomorphic for paracrinoids, and a similar morphology of the theca, are assigned to this class. Brachiolars in brachioles are a new development, homologous to the flooring plates of the food groove and, where present, are the continuations of these plates beyond the theca. The uniserial brachioles of Laurentian paracrinoids evolved from the biserial brachioles as a result of a gradual shift of brachiolars in the neighboring rows and their subsequent fusion in pairs. Brachials in crinoidal arms are a new development that evolved as distal serial growth of radial plates under the induced influence of the incipient radial canals emerging from the closed vestibular cavity, which was an ontogenetic innovation in crinoids. The transformation of a nonorganized small-plated theca into a large-plated, and completely or partly symmetrized theca, or vice versa is possible and results from accelerated or retarded growth of some plate generation in relation to the growth rate of the theca.     

A re‐interpretation of the ambulacral system of Eumorphocystis (Blastozoa,Echinodermata) and its bearing on the evolution of early crinoids

Recent debates over the evolutionary relationships of early echinoderms have relied heavily on morphological evidence from the feeding ambulacral system. Eumorphocystis, a Late Ordovician diploporitan, has been a focus in these debates because it bears ambulacral features that show strong morphological similarity to early crinoid arms. Undescribed and well‐preserved specimens of Eumorphocystis from the Bromide Formation (Oklahoma, USA) provide new data illustrating that composite arms supported by a radial plate that bear a triserial arrangement of axial and extraxial components encasing a coelomic extension can also be found in blastozoans. Previous reports have considered these arm structures to be restricted to crinoids; these combined features have not been previously observed in blastozoan echinoderms. Phylogenetic analyses suggest that Eumorphocystis and crinoids are sister taxa and that shared derived features of these taxa are homologous. The evidence from the arms of Eumorphocystis suggests that crinoid arms were derived from a specialized blastozoan ambulacral system that lost feeding brachioles and strongly suggests that crinoids are nested within blastozoans.     

Cambrian stalked echinoderms show unexpected plasticity of arm construction

Feeding arms carrying coelomic extensions of the theca are thought to be unique to crinoids among stemmed echinoderms. However, a new two-armed echinoderm from the earliest Middle Cambrian of Spain displays a highly unexpected morphology. X-ray microtomographic analysis of its arms shows they are polyplated in their proximal part with a dorsal series of uniserial elements enclosing a large coelomic lumen. Distally, the arm transforms into the more standard biserial structure of a blastozoan brachiole. Phylogenetic analysis demonstrates that this taxon lies basal to rhombiferans as sister-group to pleurocystitid and glyptocystitid blastozoans, drawing those clades deep into the Cambrian. We demonstrate that Cambrian echinoderms show surprising variability in the way their appendages are constructed, and that the appendages of at least some blastozoans arose as direct outgrowths of the body in much the same way as the arms of crinoids.     

A unique edrioasteroid from the upper Middle Cambrian of Iran,its phylogenetic implications and paleoecology

One of the earliest isorophid edrioasteroids from the upper Middle Cambrian-lower Upper Cambrian (upper part of Series 3-lower part of the Furongian Series) of northern Iran is described. It has unusual branched ambulacra, which extend beyond the theca almost to the marginal rim. These unusual features reflect the latent possibility of appearance of separated from the theca and even branching food-gathering appendages, such as arms in crinoids and brachials in blastozoans, in common ancestor of all radially symmetrical echinoderms.     

THE EARLY RADIATION AND PHYLOGENY OF ECHINODERMS

1. Living echinoderms are characterized by an extensive water vascular system developed from the larval left hydrocoel, a complex, multi-plated endoskeleton with stereom structure, and pentamery. Fossil evidence shows that stereom evolved before pentamery, but both were acquired during the Lower Cambrian. 2. Cladistic analysis of Lower Cambrian genera reveals very few characters in common between carpoids and true echinoderms, and that the split between them was the first fundamental evolutionary dichotomy within the Dexiothetica. 3. Helicoplacoids are stem group echinoderms with spiral plating and three ambulacra arranged radially around a lateral mouth. They are the most primitive echinoderms and the first to show a radial arrangement of the water vascular and ambulacral systems. Unlike later echinoderms, their skeleton shows no dorsal/ventral (aboral/oral) differentiation. They were probably sedentary suspension feeders. 4. Camptostroma is the most primitive known pentaradiate echinoderm and, in our view, possibly a common ancestor of all living groups. It had a short conical dorsal (aboral) surface with imbricate plating, a ridged lateral wall and a slightly domed ventral (oral) surface with five curved ambulacra in a 2-1-2 arrangement inherited from the triradiate pattern of the helicoplacoids. Interambulacral areas bore epispires and the CD interambulacrum contained the anus, hydropore and/or gonopore. All parts of the theca had plates in at least two layers. 5. All other echinoderms belong to one of two monophyletic subphyla, the Pelmatozoa and the Eleutherozoa. 6. Stromatocystites is the earliest known eleutherozoan and differs from Camptostroma in having a test with only one layer of plates and having lost the dorsal elongation. In Stromatocystites the dorsal surface is flat and the plating tesselate. Stromatocystites was an unattached, low-level suspension feeder. 7. The lepidocystoids are the earliest known pelmatozoans. They differ from Camptostroma in having an attached dorsal stalk which retained the primitive imbricate plating, and by developing erect feeding structures along the ambulacra. In Kinzercystis, the ambulacra are confined to the thecal surface and erect, biserial brachioles arise alternately on either side. Lepidocystis has a similar arrangement except that, the distal part of each ambulacrum extends beyond the edge of the theca as a free arm. 8. Pelmatozoans diverged more or less immediately into crinoids, with multiple free arms composed of uniserial plates, and cystoids sensu lato, which retained brachioles. Gogia (Lower to Middle Cambrian) is the most primitive known cystoid and differs from Kinzercystis principally in having all plating tesselate, while Echmatocrinus (Middle Cambrian) is the most primitive known crinoid and differs from Lepidocystis in lacking brachioles and in having more than five free arms with uniserial plates. 9. Post Lower Cambrian differentiation of pelmatozoan groups proceeded rapidly, exploiting the primitive suspension-feeding mode of life. Maximum morphological diversity was reached in the Ordovician, but thereafter crinoids progressively displaced cystoid groups and reached their peak diversity during the Carboniferous. The eleutherozoans were slower to diversify, but by the Arenig the earliest ‘sea-stars’ (in reality, advanced members of the eleutherozoan stem group) had reversed their living orientation and had begun to exploit a deposit-feeding mode of life. These in turn led to the ophiuroids, echinoids and holothuroids. 10. The basic echinoderm ambulacrum was already present in the helicoplacoids. It had biserial, alternate flooring plates and complexly plated sheets of cover plates on either side. The radial water vessel lay in the floor of the ambulacrum, external to the body cavity, and gave rise ventrally to short, lateral branches (fore-runners of tube feet) that were used to open the cover plate sheets, and dorsally was connected to internal compensation sacs which acted as fluid reservoirs (and were preadapted for a role in gaseous exchange). Plating on the cover plate sheets was organized and reflected the positions of the lateral branches from the radial water vessel. In Camptostroma, the cover plate sheets had biserially aligned rows of cover plates associated with the lateral branches. 11. Brachioles arose by extension of the lateral branches of the radial water vessel and associated serially aligned cover plates found in Camptostroma. They bear a single alternate series of cover plates. In Lepidocystis the ambulacra extended beyond the edge of the oral surface as true arms. Brachial plates of arms are homologues of primary ambulacral flooring plates, and arms bear multiple series of cover plates. Uniserial ambulacral plating is a derived condition and evolved independently in crinoids, paracrinoids and isorophid edrioasteroids. Pinnules in crinoids arose independently in inadunates and camerates by a progressively more unequal branching of the arms. Thus all parts of the subvective system in crinoids are internally homologous, whereas in cystoids, brachioles and arms (or ambulacra) are not homologous structures. 12. The position of the hydropore is the best reference point in orientating echinoderms. Carpenter's system of identifying ambulacra by letters, arranged clock-wise in oral view with the A ambulacrum opposite the hydropore, is consistent in all echinoderm classes. In all Lower Cambrian pentaradiate echinoderms the anus, gonopore and hydropore lie in the CD interambulacrum and this is accepted as the primitive arrangement. In helicoplacoids we tentatively suggest that the A ambulacrum spiralled down from the mouth while the two ambulacra that spiralled up represent the B + C and D + E ambulacra combined. 13. The pelmatozoan stem arose from a polyplated stalk, via a meric stem to a true column with holomeric (single piece) columnals. This happened independently in the crinoids and the cystoids. 14. Our analysis of echinoderm phylogeny leads us to recommend the following changes to the higher level classification of echinoderms: The phylum Echinodermata includes only those groups with radial symmetry superimposed upon a fundamental larval asymmetry. It has a stem group that contains the triradiate helicoplacoids and a crown group to which all other (pentaradiate) echinoderms belong. The crown group contains two monophyletic subphyla, the Pelmatozoa and the Eleutherozoa, and the Pelmatozoa contains two superclasses, the Crinoidea which are extant and the Cystoidea, which are extinct.     

Evolution Within a Bizarre Phylum: Homologies of the First Echinoderms

SYNOPSIS. The Extraxial/Axial Theory (EAT) of echinoderm skeletalhomologies 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 earlylarval development. Axial skeleton originates in the rudiment,and develops in association with the pentaradially arrangedhydrocoel according to specific ontogenetic principles. Perforateand imperforate extraxial regions are associated with the leftand right somatocoels respectively, are not governed by ontogeneticprinciples of plate addition, and are products of the non-rudimentpart of the larval body. The morphology of even the most bizarreof the earliest echinoderms can be explored using the EAT. Amongthese, edrioasteroid-like taxa best fit the idea that formsexpressing archimery in the sequential arrangement of axial,perforate extraxial, and imperforate extraxial regions are thefirst echinoderms. Metamorphosis is especially marked in cladesthat have a high axial to extraxial skeleton ratio because structuresdeveloping from the non-rudiment part are suppressed in favorof the developing axial elements during this process. However,inearly echinoderms, extraxial skeleton makes up a far largerproportion of the body wall than axial, implying that metamorphosiswas not as significant a part of the developmental trajectoryas it is in more recently evolved taxa. Echinoderm radiationconsists of a succession of apomorphies that reduced the expressionof extraxial components but increased the influence of axialones, with a concomitant increase in the prominence of metamorphosis.     

Arms versus brachioles: Morphogenetic basis of similarity and differences in food-gathering appendages of pelmatozoan echinoderms

The similarity in the skeleton model of the brachiolar food-gathering system of Blastozoa and the arm system of Crinozoa, including the apical growth with enantomorphous displacement of skeletal ele-ments, is explained by the primary organizing role of the radial ambulacral canals, which have the same branching model for ambulacral tentacles. The difference in the positions of brachioles and arms relative to the theca (exothecal and endothecal) is associated with the formation of the primary ambulacral tentacles directly on the body surface of the majority of Blastozoa, particularly, the closed vestibular cavity of crinoids. The supporting skeleton of brachioles arose as a branch of the plates covering the floor of the ambulacrum, if they were present, or formed similarly as a new formation outside the theca. The supporting skeleton of arms, brachials, developed as a result of the serial growth of plates positioned radially at the boundary of the aboral skeleton and tegmen formed due to the appearance of the vestibulum. The hypothesis of the inductive role of hydrocoel and its radial ambulacral appendages, which organize the arrangement of skeletal elements in the morphogenesis of echinoderms, enables the refinement of the principle of skeleton division into the axial and extraxial parts. The axial skeleton has a developmental model formed under the control of the radial ambu-lacral canals. Remaining skeleton is extraxial, subdivided into the symmetrized part arranged under direct or indirect organizing effect of the hydrocoel and unregulated, nonsymmetrized part, which is not connected initially with the influence of the hydrocoel.     

Pelmatozoan arms from the Middle Cambrian of Australia: bridging the gap between brachioles and brachials?

The early Middle Cambrian Monastery Creek Phosphorite (Beetle Creek Formation, Queensland, Australia) contains an assemblage of disarticulated echinoderm ossicles that are exquisitely preserved. Amongst this material we recognize pelmatozoan brachials, radials, basals and holomeric columnals. Although we cannot reconstruct the complete animal with precision, these elements represent the oldest known pelmatozoan with crinoid-like appendages. Key elements include isotomously to heterotomously branched uniserial appendage plates with a tripartite adoral food groove, a longitudinal central canal interpreted as housing entoneural nerve, and differentiated articulation facets. There are also epispire-bearing radials bearing one to four arm insertion-facets, each one pierced by a central neural canal. These canals run internal towards the oral area beneath the external food groove. Co-occuring material includes single truncated cone-shaped basals and holomeric columnals, both with a similar articulation pattern, and irregular, epispire-bearing thecal plates. This mosaic of crinozoan (uniserial isotomous to heterotomous arms with neural canal), blastozoan (epispire-bearing thecal plates, appendage leading to oral thecal food groove without direct connection with body cavity) and apomorphic characters (circumoral instead of basal entoneural plexus) is unexpected and demonstrates that crinoid-like pelmatozoans with uniserial, branched arms appeared significantly earlier than the Tremadocian, when the first articulated crinoid skeletons are found. It also raises questions about the polyphyletic appearance of feeding appendages among pelmatozoan echinoderms.     

Arrays in rays: terminal addition in echinoderms and its correlation with gene expression

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 every echinoderm, from the occasionally bizarre fossils of the Cambrian to the most familiar extant taxa. Using the OPR and other criteria of recognition, echinoderm body wall can be divided into two main regions: extraxial components are associated with the somatocoels, axial components (formed in accordance with the OPR) with the hydrocoel. We compare patterns of development in axial regions of echinoderms with those found in the anterior-posterior axes of the earliest echinoderms as well as other invertebrates. Although axial and extraxial skeletons appear to be composed of the same biomineral matrix, the genes involved in patterning these two skeletal components are likely distinct. During development of the axial skeleton, for instance, the genes engrailed and orthodenticle are expressed in spatial and temporal patterns consistent with the OPR. Other genes such as distal-less seem to demarcate early ontogenetic boundaries between the axial rudiment and the extraxial larval body. There is a complex and pervasive reorganization of gene expression domains to produce the highly divergent morphologies seen in the Echinodermata. We integrate morphological and genetic information, particularly with respect to the origins of radial symmetry in the rudiment, and the concomitant development of the rays.     

Disarticulation patterns in Ordovician crinoids: Implications for the evolutionary history of connective tissue in the Crinoidea

Taphonomic information is examined to evaluate the early history of connective tissues in the Crinoidea. The pattern of stalk segmentation of Middle and Late Ordovician crinoids is consistent with the two-ligament (intercolumnal and through-going ligaments) pattern present in living isocrinid crinoids and interpreted for fossil isocrinids, holocrinids, and Lower Mississippian crinoids. A single rhombiferan was also examined; its taphonomic pattern is also indicative of this style of tissue organization. Furthermore, the taphonomy of all Middle and Late Ordovician crinoids may reflect that they lacked discretely organized muscles between arm brachials, which is consistent with the hypothesis that muscles evolved as a connective tissue between plates only once within the Crinoidea, during the Early Devonian. These data indicate that the two-ligament organization of the stalk is a primitive feature among the Crinoidea and perhaps even among stalked echinoderms. Therefore, the autotomy function of this column-tissue organization among living crinoids is an exaptation. On the other hand, discretely organized muscles as connective tissue in crinoid arms is a derived trait that first appeared during the middle Paleozoic; this adaptation proved very successful for the advanced cladid crinoids.     

Morphology and biomechanical implications of isolated discocystinid plates (Edrioasteroidea, Echinodermata) from the Carboniferous of North America

Detailed examination of isolated thecal plates belonging to three discocystinid edrioasteroids, Spiraclavus nacoensis Sumrall, Hypsiclavus kinsleyi Sumrall, and Giganticlavus bennisoni Sumrall and Bowsher, reveals striking similarity in morphology among these species. Stereom observed in the ambulacral floor plates indicates that ligamentous connective tissue opened the ambulacral cover plates and muscle tissue closed them. The ambulacral floor plates are interpreted as rigid supports for the oral surface with the interambulacral areas acting as flexible integuments of plates. The aboral surface is interpreted as flexible and highly contractile. All discocystinid thecal openings are consistent in morphology with adaptations for thecal pressurization. Extension and contraction of the theca was accomplished by pumping water in and out of an inflatable sac associated with the periproctial opening. The pedunculate zone is interpreted as passively expanding and contracting by relaxing of mutable collagenous tissue and stiffening when the theca was in the desired position. All of these features illustrate that discocystinid edrioasteroids have highly–evolved morphology and function.     

Homology of posterior interray plates in crinoids: a review and new perspectives from phylogenetics,the fossil record and development

Despite their importance for understanding phylogeny, character evolution and classification, well-constrained homology relationships for posterior plating in crinoids have only recently been attempted. Here, we re-evaluate posterior plate homologies in all major crinoid lineages using development, fossil ontogenies and phylogenetic evidence. Based on these lines of evidence, we change terminology for some posterior plates to correct misnomers and make recommendations for updated terminology of others to better reflect homology. Among pentacrinoids (disparids, hybocrinids, eucladids, flexibles and articulates) the relative position of posterior interray plates, not their topology, reflects homology. From proximal to distal, pentacrinoid posterior plates are the radianal, anal X and right sac plate, regardless of the total number of plates in the adult calyx. Camerate posterior plating contrasts with pentacrinoids, but insufficient data are available to resolve homology relationships between these two clades. More examples of early post-larval ontogeny are needed in camerates and other Palaeozoic crinoids.     

Phylogenetic implications of the Protocrinoida: Blastozoans are not ancestral to crinoids

A traditional, widely cited hypothesis for over a century posits the origin of the crinoids from blastozoans. The blastozoan hypothesis is contradicted by the discovery of a new crinoid order, the Protocrinoida. Protocrinoids exhibit many traits that are consistent with a basal crinoid phylogenetic position, but inconsistent with a blastozoan ancestry. Protocrinoids are among the oldest crinoids and are therefore stratigraphically correctly placed. The blastozoan hypothesis in contrast, relies on putative homologies between blastozoans and crinoids taken from taxonomically and stratigraphically disparate representatives of both groups; these disparities indicate homoplasy rather than propinquity of descent. Data supporting these ideas are reviewed here. These findings reinforce insightful observations made by Georges Ubaghs decades ago with less data.     

The anteroposterior axis in echinoderms and displacement of the mouth in their phylogeny and ontogeny

The difference in the position of the anteroposterior axis in larval and adult echinoderms is related to the displacement of the mouth from the anterior end of the body to the posterior end in the phylogeny of echinoderms, which occurred in the course of the reorganization of their body plan from bilateral asymmetrical to radiosymmetrical. Traces of this phylogenetic process have been especially fully preserved in the ontogeny of crinoids. Other recent echinoderms have largely lost such traces. Dislocation of Hox-genes in sea urchins, resulting from the translocation of these genes to the 5′ end of the chromosome and inversion of the anterior Hox-genes, is explained by the necessity to preserve the spatial and temporal colinearity in the course of the convergence of the starting and final stages of the mouth displacement process, similar to the elevation process in crinoids, and inclusion in the basic body plan of the structure of a rudiment now regulated directly by the anterior Hox-genes.     

What a New Model of Skeletal Homologies Tells Us About Asteroid Evolution

The Extraxial-Axial Theory (EAT) is applied to the body wallhomologies of asteroids. Attempts to characterize major platesystems of asteroids as axial or extraxial, particularly thosethat are highly organized into series, can be problematic. However,the Optical Plate Rule (OPR) is instrumental in establishingthat ambulacrals and terminals are axial. It is equally clearthat the region aboral to the marginal frame is a part of theperforate extraxial body wall (with the possible exception ofthe centrodorsal, which is likely imperforate extraxial). Previouslyestablished EAT criteria, particularly those strongly rootedin the embryologically expressed boundary between axial andextraxial body wall in larvae, suggest that marginals, and perhapsadambulacrals, are extraxial in origin. We also explore theextraxial nature and phylogenetic significance of the odontophore.Our data from both juveniles and adults show that plate andtube foot addition sequences occur according to the OPR, andshed light on poorly known homologies of the asteroid mouthframe. These data indicate that the mouth angle ossicle mustat least contain the first ambulacral, although we cannot ruleout the possibility that the first adambulacral also contributesto the construction of this ossicle. The interpretations providedby the EAT for all ossicles suggest a synapomorphy scheme forsomasteroids, ophiuroids, and asteroids.     

Evolutionary history of regeneration in crinoids (Echinodermata)

The fossil record indicates that crinoids have exhibited remarkable regenerative abilities since their origin in the Ordovician, abilities that they likely inherited from stem-group echinoderms. Regeneration in extant and fossil crinoids is recognized by abrupt differences in the size of abutting plates, aberrant branching patterns, and discontinuities in carbon isotopes. While recovery is common, not all lost body parts can be regenerated; filling plates and overgrowths are evidence of non-regenerative healing. Considering them as a whole, Paleozoic crinoids exhibit the same range of regenerative and non-regenerative healing as Recent crinoids. For example, Paleozoic and extant crinoids show evidence of crown regeneration and stalk regrowth, which can occur only if the entoneural nerve center (chambered organ) remains intact. One group of Paleozoic crinoids, the camerates, may be an exception in that they probably could not regenerate their complex calyx-plating arrangements, including arm facets, but their calyxes could be healed with reparative plates. With that exception, and despite evidence for increases in predation pressure, there is no compelling evidence that crinoids have changed though time in their ability to recover from wounds. Finally, although crinoid appendages may be lost as a consequence of severe abiotic stress and through ontogenetic development, spatiotemporal changes in the intensity and frequency of biotic interactions, especially direct attacks, are the most likely explanation for observed patterns of regeneration and autotomy in crinoids.     

The complete mitochondrial genomes of the sea lily Gymnocrinus richeri and the feather star Phanogenia gracilis: signature nucleotide bias and unique nad4L gene rearrangement within crinoids

Complete DNA sequences have been determined for the mitochondrial genomes of the crinoids Phanogenia gracilis (15892 bp) and Gymnocrinus richeri (15966 bp). The mitochondrial genetic map of the stalkless feather star P. gracilis is identical to that of the comatulid feather star Florometra serratissima (Scouras, A., Smith, M.J., 2001. Mol. Biol. Evol. 18, 61-73). The mitochondrial gene order of the stalked crinoid G. richeri differs from that of F. serratissima and P. gracilis by the transposition of the nad4L protein gene. The G. richeri nad4L mitochondrial map position is unique among metazoa and is likely a derived feature in this stalked crinoid. Nucleotide compositional analyses of protein genes encoded on the major sense strand confirm earlier conclusions regarding a crinoid-distinctive T over C bias. All three crinoids exhibit high T levels in third codon positions, whereas other echinoderm classes favor A or C in the third codon position. The nucleotide bias is reflected in the relative synonymous codon usage patterns of crinoids versus other echinoderms. We suggest that the nucleotide bias of crinoids, in comparison to other echinoderms, indicates that a physical inversion of the origin of replication has occurred in the crinoid lineage. Evolutionary rate tests support the use of the cytochrome b (cob) gene in molecular phylogenetic analyses of echinoderms. A consensus echinoderm tree was generated based on cytochrome b nucleotide alignments that placed the asteroids as a sister group to a clade containing the ophiuroids and the (echinoids+holothuroids) with the crinoids basal to the rest of the echinoderm classes: [Crinoid,(Asteroid,(Ophiuroid,(Echinoid,Holothuroid)))].     

Modularity and heterochronies in the evolution of metazoa: Paleontological aspect

All organisms are formed of more or less independent elements, modules. Paleontology deals with morphological modules preserved in the fossil state and allows their evolution within taxa of different levels to be reconstructed. Modularity provides organisms with the ability to evolve, since changes in one module does not influence others, nor disturb the integrity of organism. Each organism may have unique modules represented by a single copy and serial modules developing according to a certain symmetry type. Serial terminal growth is characteristic of ambulacra of echinoderms, in which it is combined with alternating appearance of structures on the right and left of the symmetry plane. The morphology of the solute Maennilia estonica, which has been investigated in detail, shows that the growth model for the brachiola is similar to the model for ambulacra of sea urchins, but without an ocular plate. Probably, the hydrocoel initially induced the appearance of a skeleton necessary for its activity and organized its development according to its own model of terminal growth. Subsequently, the axial skeleton appearing following this pattern could have organized the growth of adjacent parts of the extraxial skeleton following the same model to form a united module. The fusion of modules could have resulted from heterochronies. Extant and extinct material connected with the change in the anteroposterior axis in evolutionary and ontogenetic development of echinoderms provides a prominent example of heterochronies. Heterochronies were the mechanism connecting characters into an integrated ensemble of the body plan. Archaic diversity reflects an attempt to create a new body plan. Various manifestations of archaic diversity show that the emergence of a new higher taxon is connected with the combination of a number of characters united in an integrated complex forming the body plan which is stable from the moment of appearance due to strict recursive relationships between its modules rather than with the acquisition of an individual character, even if it is very important.     

Molecular phylogeny and character evolution of the chthamaloid barnacles (Cirripedia: Thoracica)

The Chthamaloidea (Balanomorpha) present the most plesiomorphic characters in shell plates and cirri, mouthparts, and oral cone within the acorn barnacles (Thoracica: Sessilia). Due to their importance in understanding both the origin and diversification of the Balanomorpha, the evolution of the Chthamaloidea has been debated since Darwin's seminal monographs. Theories of morphological and ontogenetic evolution suggest that the group could have evolved multiple times from pedunculated relatives and that shell plate number diminished gradually (8→6→4) from an ancestral state with eight wall plates surrounded by whorls of small imbricating plates; but this hypothesis has never been subjected to a rigorous phylogenetic test. Here we used multilocus sequence data and extensive taxon sampling to build a comprehensive phylogeny of the Chthamaloidea as a basis for understanding their morphological evolution. Our maximum likelihood and Bayesian analyses separate the Catophragmidae (eight shell plates and imbricating plates) from the Chthamalidae (8-4 shell plates and no imbricating plates), but do no support a gradual reduction in shell plates (8→6→4). This suggests that evolution at the base of the Balanomorpha involved a considerable amount of homoplasy.     

Early rodent incisor enamel evolution: Phylogenetic implications

Incisor enamel microstructure proved to be a very effective tool for assessment of phylogenetic relationships among the Rodentia. Pauciserial and multiserial Schmelzmuster are clearly distinct by structural characters such as orientation of interprismatic matrix, presence or absence of transition zones between Hunter-Schreger bands (HSB), inclination of HSB, enamel thickness, and others. Pauciserial HSB are structurally very close to the earliest known mammalian HSB found in Paleocene arctocyonids. Biomechanical arguments and outgroup comparison with mixodontians indicate that the pauciserial Schmelzmuster is a symplesiomorphy of the Rodentia. Transitional stages from pauciserial to multiserial Schmelzmuster were observed in middle Eocene ctenodactyloids and from pauciserial to uniserial in middle to late Eocene anomalurids. The multiserial Schmelzmuster is considered a synapomorphy of the Hystricognathi, ctenodactylids, and pedetids. Schmelzmuster evolution reflects the early dichotomy of the Rodentia: In the Asian ctenodactyloid lineage a multiserial Schmelzmuster evolved once and in the North American/European ischyromyoid lineage a uniserial Schmelzmuster developed several times convergently. The pauci- to uniserial Schmelzmuster of the anomalurids excludes a close relationship to the phiomyids, because the ctenodactyloid-phiomyid lineage is characterized by the development of a multiserial Schmelzmuster.