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The macrostructural anatomy and functional morphology of dendrochirotid sea cucumber’s (Echinodermata) calcareous rings
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In the field of correlative microscopy, light and electron microscopy form a powerful combination for morphological analyses in zoology. Due to sample thickness limitations, these imaging techniques often require sectioning to investigate small animals and thereby suffer from various artefacts. A recently introduced nanoscopic X-ray computed tomography (NanoCT) setup has been used to image several biological objects, none that were, however, embedded into resin, which is prerequisite for a multitude of correlative applications. In this study, we assess the value of this NanoCT for correlative microscopy. For this purpose, we imaged a resin-embedded, meiofaunal sea cucumber with an approximate length of 1 mm, where microCT would yield only little information about the internal anatomy. The resulting NanoCT data exhibits isotropic 3D resolution, offers deeper insights into the 3D microstructure, and thereby allows for a complete morphological characterization. For comparative purposes, the specimen was sectioned subsequently to evaluate the NanoCT data versus serial sectioning light microscopy (ss-LM). To correct for mechanical instabilities and drift artefacts, we applied an alternative alignment procedure for CT reconstruction. We thereby achieve a level of detail on the subcellular scale comparable to ss-LM images in the sectioning plane.
Recent studies have shown that micro-computed tomography (µCT) must be considered one of the most suitable techniques for the non-invasive, three-dimensional (3D) visualization of metazoan hard parts. In addition, µCT can also be used to visualize soft part anatomy non-destructively and in 3D. In order to achieve soft tissue contrast using µCT based on X-ray attenuation, fixed specimens must be immersed in staining solutions that include heavy metals such as silver (Ag), molybdenum (Mo), osmium (Os), lead (Pb), or tungsten (W). However, while contrast-enhancement has been successfully applied to specimens pertaining to various higher metazoan taxa, echinoderms have thus far not been analyzed using this approach. In order to demonstrate that this group of marine invertebrates is suitable for contrast-enhanced µCT as well, the present study provides results from an application of this technique to representative species from all five extant higher echinoderm taxa. To achieve soft part contrast, freshly fixed and museum specimens were immersed in an ethanol solution containing phosphotungstic acid and then scanned using a high-resolution desktop µCT system. The acquired datasets show that the combined visualization of echinoderm soft and hard parts can be readily accomplished using contrast-enhanced µCT in all extant echinoderm taxa. The results are compared with µCT data obtained using unstained specimens, with conventional histological sections, and with data previously acquired using magnetic resonance imaging, a technique known to provide excellent soft tissue contrast despite certain limitations. The suitability for 3D visualization and modeling of datasets gathered using contrast-enhanced µCT is illustrated and applications of this novel approach in echinoderm research are discussed.
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The first morphological observation of a euryalid brittle star, Asteronyx loveni, using non-destructive X-ray micro-computed tomography (µCT) was performed. The body of euryalids is covered by thick skin, and it is very difficult to observe the ossicles without dissolving the skin. Computed tomography with micrometer resolution (approximately 4.5–15.4 µm) was used to construct 3D images of skeletal ossicles and soft tissues in the ophiuroid’s body. Shape and positional arrangement of taxonomically important ossicles were clearly observed without any damage to the body. Detailed pathways inside the vertebral ossicles, lateral arm plates, and arm spines for passage of nerves and water vascular structures were observed. Inter-vertebral muscles were also observed. Forms and 3D arrangements of many important taxonomical characters of the euryali xonomy of the class Ophiu ds were scrutinized by µCT in high enough resolution for taxonomic description of ophiuroids.
Ophiuroid systematics is currently in a state of upheaval, with recent molecular estimates fundamentally clashing with traditional, morphology-based classifications. Here, we attempt a long overdue recast of a morphological phylogeny estimate of the Ophiuroidea taking into account latest insights on microstructural features of the arm skeleton. Our final estimate is based on a total of 45 ingroup taxa, including 41 recent species covering the full range of extant ophiuroid higher taxon diversity and 4 fossil species known from exceptionally preserved material, and the Lower Carboniferous Aganaster gregarius as the outgroup. A total of 130 characters were scored directly on specimens. The tree resulting from the Bayesian inference analysis of the full data matrix is reasonably well resolved and well supported, and refutes all previous classifications, with most traditional families discredited as poly- or paraphyletic. In contrast, our tree agrees remarkably well with the latest molecular estimate, thus paving the way towards an integrated new classification of the Ophiuroidea. Among the characters which were qualitatively found to accord best with our tree topology, we selected a list of potential synapomorphies for future formal clade definitions. Furthermore, an analysis with 13 of the ingroup taxa reduced to the lateral arm plate characters produced a tree which was essentially similar to the full dataset tree. This suggests that dissociated lateral arm plates can be analysed in combination with fully known taxa and thus effectively unlocks the extensive record of fossil lateral arm plates for phylogenetic estimates. Finally, the age and position within our tree implies that the ophiuroid crown-group had started to diversify by the Early Triassic.
Several derived sand dollar (Echinoidea: Clypeasteroida) families are characterized by the presence of Gregory’s diverticulum, an accessory organ of the digestive tract. This soft tissue structure is composed of a central tubular cecum that gives off multiple lobes into the periphery of the test. Most notable are the organ’s capacity to selectively store sand grains that the animal has taken up from the surrounding sediment as well as the gradual reduction of Gregory’s diverticulum during ontogeny. Several aspects of the biology of this structure have remained unexplored, including the organ’s precise morphology and structural diversity. In order to provide a concise basis for future histological, physiological, and functional analyses, a comprehensive comparative morphological and phylogenetic study across numerous taxa was undertaken. Taxon sampling comprised over 100 clypeasteroid species, including various fossil taxa. This extensive dataset permits establishing a concise terminology that incorporates all of the organ’s substructures. In addition, three-dimensional models of Gregory’s diverticulum are presented that provide an improved spatial understanding of the organ’s morphology in situ. The combined data from dissection, X-ray imaging, microcomputed tomography, and magnetic resonance imaging reveal a previously unknown variability of the structure, which also yields several phylogenetically informative morphological characters. Among those sand dollar families that possess Gregory’s diverticulum, the organ is present in two distinct shapes, which can be distinguished by the number, shape, and location of substructures. In addition, the data provide unequivocal evidence that Gregory’s diverticulum is absent in the extant taxa Rotulidae and Astriclypeidae, but also in the enigmatic Marginoproctus.
Available data on the formation of echinoderm skeleton are reviewed based on the literature. The development and structural features of the skeleton and morphological, histological, and molecular data on the skeleton formation in different echinoderm groups are described. Recent data on the study of gene regulatory networks and their effect on biomineraization and skeleton formation are also included. The skeletogenic mechanisms are in general highly conservative within echinoderms at the levels of genes, gene regulatory networks, and involved cell populations. At the same time, the formation mechanism of the echinoderm skeleton is unique and has not been recorded in other taxonomic groups.
A new genus and a new species of dendrochirotid, Thandarum hernandezi gen. et sp. nov., is described from Buenos Aires coast in the Southwestern Atlantic. For this species is quite significant the body U shape, up to 14.08 mm (along the trivium), with double row of podia restricted to the ambulacra. Other features are ossicles from body wall with buttons and 4 pillar tables, tube feet with rods and end plate star-shaped; the introvert with rosettes and tentacles with rods. These characteristics require the recognition of a new genus, and the new species and represents the first sclerodactylid reported in Argentinean waters.
O'Loughlin, P. M., and VandenSpiegel, D. 2007. New apodid species from southern Australia (Echinodermata: Holothuroidea: Apodida). Memoirs of Museum Victoria 64: 53-70 A new chiridotid genus is erected: Archedota O'Loughlin gen. nov. In addition, seven apodid species, new to science, are described (O'Loughlin as author) for the rocky shallows, continental shelf and continental slope of southern Australia: chiridotids Archedota lapidea, Taeniogyrus papillis, Taeniogyrus tantulus, Trochodota epiphyka; myriotrochids Prototrochus burni, Prototrochus staplesi, Prototrochus taniae. Taeniogyrus heterosigmus Heding, Taeniogyrus roebucki (Joshua), Trochodota allani (Joshua) and Trochodota shepherdi Rowe are discussed. Keys are provided for southern Australian species of Taeniogyrus Semper and Trochodota Ludwig. A table is provided distinguishing Tasman Sea myriotrochid species.
Various interpretations of the holothurian system and phylogeny are critically reviewed and the main characters that form the basis of the existing systematics of this group are analyzed. A system of holothurians based on thorough analysis of their morphology and anatomy is proposed. Four subclasses are recognized in the class Holothuroidea: Arthrochirotacea, Synaptacea, Elpidiacea, and Holothuriacea. The subclass Arthrochirotacea includes the extinct Paleozoic order Arthrochirotida. The subclass Synaptacea includes the order Synaptida with two suborders and three families. The subclass Elpidiacea includes the order Elasipodida with four families. The subclass Holothuriacea includes four orders: Aspidochirotida with five families; Dendrochirotida with 15 families (14 extant and one extinct); Molpadiida with three families; Gephyrothuriida with one family and two genera Gephyrothuria and Hadalothuria. The order Gephyrothuriida is re-established. The order Dactylochirotida Pawson et Fell, 1965 is synonymized under the order Dendrochirotida. A new suborder Cucumariina and new family Mesothuriidae are described. The family Vaneyellidae is synonymized under the family Cucumariidae. Four subfamilies are classified as families: Cladolabidae, Sclerothyonidae, Monilipsolidae, and Thyonidiidae.
Continuous improvements in the resolution of three-dimensional imaging have led to an increased application of these techniques in conventional taxonomic research in recent years. Coupled with an ever increasing research effort in cybertaxonomy, three-dimensional imaging could give a boost to the development of virtual specimen collections, allowing rapid and simultaneous access to accurate virtual representations of type material. This paper explores the potential of micro-computed tomography (X-ray micro-tomography), a non-destructive three-dimensional imaging technique based on mapping X-ray attenuation in the scanned object, for supporting research in systematics and taxonomy. The subsequent use of these data as virtual type material, so-called "cybertypes", and the creation of virtual collections lie at the core of this potential. Sample preparation, image acquisition, data processing and presentation of results are demonstrated using polychaetes (bristle worms), a representative taxon of macro-invertebrates, as a study object. Effects of the technique on the morphological, anatomical and molecular identity of the specimens are investigated. The paper evaluates the results and discusses the potential and the limitations of the technique for creating cybertypes. It also discusses the challenges that the community might face to establish virtual collections. Potential future applications of three-dimensional information in taxonomic research are outlined, including an outlook to new ways of producing, disseminating and publishing taxonomic information.
Tomographic imaging techniques such as micro-computed tomography (μCT) and magnetic resonance imaging (MRI) permit the gathering of digital anatomical data from whole animal specimens non-invasively. The resulting datasets can be used for direct observation of the two-dimensional tomographic image data as well as for manual and semi-automated three-dimensional modelling. Freshly fixed specimens as well as preserved museum material can be successfully analyzed using this approach, giving the zoomorphologist a powerful tool for large-scale comparative studies. In order to demonstrate the principle suitability of non-invasive imaging in echinoderm research, μCT scans of 199 and MRI scans of 92 sea urchin (Echinodermata: Echinoidea) species were acquired, resulting in a total of 203 analyzed echinoid species. The taxa selected represent 50 of the currently recognized 60 extant sea urchin families. The present article lists all species that have been analyzed so far and provides information about the scanning parameters employed for each dataset. Furthermore, the workflow established to generate three-dimensional models of sea urchins is outlined. Using a number of examples from μCT as well as MRI scans performed on echinoids, the potential of the systematic approach described here is highlighted. Finally, the suitability of non-invasive imaging techniques for the study of other echinoderm taxa is assessed based on multimodal datasets of representative species.
The relationships of post-Palaeozoic echinoids at family level are established through phylogenetic analysis of 169 taxa and 306 skeletal characters (excluding pedicellariae). Previous phylogenetic analyses of echinoids have either examined specific subgroups in detail or have looked at a relatively small number of taxa selected from across the class, with sparse sampling potentially affecting the reliability of results adversely. Our new analyses represent a compromise between encompassing the diversity of form that exists, while keeping the number of taxa to a level that does not make rigorous analysis impossibly time-consuming. In constructing the taxon-character data matrix we have encountered a surprising lack of primary data on plating pattern, lantern, and girdle structure for many supposedly "well-known" taxa. A well-resolved phylogenetic hypothesis was obtained and is used as the basis for a formal classification. Characters generally have a high retention index (>0.7) but low consistency index (<0.25) suggesting that, although characters are largely retained after they first evolve, most also undergo occasional reversal or convergence. Although parts of the resulting trees are only weakly supported (e.g. the precise sister group of the Irregularia), other parts are unambiguously resolved. Not unexpectedly, deep nodes are often not supported by unique apomorphies and higher taxa acquire their characteristic set of features over time. Diagnoses based on crown group taxa thus often fail to encompass fossil stem-group members adequately. Establishing the relationships of taxa at the root of large groups is hampered by limited character resolution. The influence of fossil taxa on the topology was explored by comparing the tree topologies obtained with and without their inclusion. We show that removal of fossils from stem groups makes no difference where their crown group is morphologically conservative, but has a major influence where extant sister groups are separated by large morphological gaps. Completeness of the echinoid record and its match to the stratigraphical record of first occurrences is tested using various metrics and found to be highly congruent, with irregular echinoids showing a higher congruence than regular ones.
Holothuroids, or sea cucumbers, are an abundant and diverse group of echinoderms with over 1400 species occurring from the intertidal to the deepest oceanic trenches. In this study, we report the first phylogeny of this class, based on a cladistic analysis of 47 morphological characters. We introduce several previously unconsidered synapomorphic characters, examine the relationships between representatives from all extant families and assess the assumptions of monophyly for each order and subclass. Maximum-parsimony analyses using three rooting methods recovered well-supported and identical topologies when two small and apparently derived families, Eupyrgidae and Ge-phyrothuriidae, were removed. The results suggest that the higher-level arrangement of Holothuroidea warrants a considerable revision. Apodida was sister to the other holothuroids. The monophyly of Dendrochirotida was not supported and the group may be paraphyletic. A randomization test using Wills' gap excess ratio found significant congruence between the phylogeny and the stratigraphic record of fossil members, suggesting that the fossil record of holothuroids is not as incomplete as is often stated. The fossil-calibrated tree indicated that several groups of holothuroids survived the end-Permian mass extinction and that the clade composed of Dendrochirotida, Dactylochirotida, Aspidochirotida and Molpadiida rapidly radiated during the Triassic.
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.
Despite descending from heavily calcified ancestors, the holothuroid skeleton is fully internal and composed of microscopic ossicles and a ring of plates bound by connective tissue, the calcareous ring. The calcareous ring exhibits a complex and poorly understood morphology; as a result, establishing unambiguous homology statements about its macrostructure has been challenging and phylogenetic studies have had to simplify this important structure. Here, we provide the first broad comparative study of Dendrochirotida calcareous rings using micro-computed tomography (μCT). A detailed description of the three-dimensional macrostructure of the calcareous ring of 10 sea cucumber species, including rare and type specimens, is presented. The structures observed were highly variable at the subfamily level, especially at the point of tissue attachment. The relationship between the calcareous ring and its associated organs, and their functional morphology are discussed. To aid future phylogenetic studies, we listed 22 characters and performed a preliminary cladistic analysis. The topology obtained supports the idea that the simple, cucumariid ring is ancestral to the mosaic-like phyllophorid ring; however, it did not support the monophyly of the cucumariids. It also did not support the family Sclerodactylidae, which was described based on the ring morphology. Differently from the dermal ossicles, which are highly homoplastic, the general homoplasy index of the calcareous ring characters was relatively low. This result highlights the importance of this structure for phy-logenetic inference. Unfortunately, time since collection, rough collection methods and fixation can damage the skeleton, and the calcareous ring is often overlooked in taxonomic descriptions. The data presented here will improve our understanding of holothuroid relationships and facilitate studies on holothuroid functional morphology and biomechanics. K E Y W O R D S comparative morphology, Holothuroidea, internal anatomy, phylogeny, skeleton, three-dimensional imaging, μCT 1 | INTRODUC TI ON Echinoderms are characterized by their complex and lightweight en-doskeleton, formed by an intricate mesh of magnesium-rich calcium carbonate-the stereom-which most likely evolved after the split between the echinoderms and the other deuterostomes (David & Mooi, 1998; Nebelsick et al., 2015; Smith, 1990). Because it is more easily preserved and identified than other characteristic echinoderm
Here, we review the taxonomy of the Apodida species from the Brazilian coast and oceanic
islands. Eleven species are reported, four Chiridotidae (including Chantalia conandae gen. et
sp. nov.) and seven Synaptidae (including Yemoja gen. nov.). Chantalia conandae gen. et sp.
nov. is a shallow-water species diagnosed by having 12 tentacles, 4–6 Polian vesicles and
vase-shaped ciliated funnels along the mesenteries, and by lacking a calcareous ring, sensory
cups and ossicles. Based on the morphology of the calcareous ring (dumbbell-shaped plates)
and of the tentacles (12, simple), we propose the new genus Yemoja gen. nov. to
accommodate Leptosynapta brasiliensis (Yemoja brasiliensis comb. nov.). To date, there are no
records of Myriotrochidae in Brazil. In addition to species descriptions and photographs, we
provide novel data on the calcareous ring stereom and a discussion about the calcareous
ring morphology in Apodida, an identification key for the Brazilian species, a synoptic table
with morphological characteristics of the Brazilian genera, and a synoptic table comparing
the ossicleless Apodida genera. Finally, we provide the geographic and bathymetric
distribution of the Apodida species reported for the South Atlantic and adjacent Southern
Ocean. Based on our compendium, the Synaptidae is most diverse in warmer waters, while
the Myriotrochidae is most diverse in colder waters. Also, the Atlantic coasts of South
America and Africa have a surprisingly low with their adjacent Antarctic waters. We expect
that dedicated holothuroid research will greatly increase the diversity of Apodida throughout
the South Atlantic Ocean, especially off the continental shelf.
Recent efforts to reconstruct the phylogeny of brittle stars (ophiuroids) have shown the need for more objective and reproducible data collection methods than the traditional visual examination and verbal description of morphological characters. Complex skeletal structures may be better understood in three dimensions than in two dimensions obtained from techniques like scanning electron microscopy (SEM). We test this hypothesis using three types of three-dimensional tomographic imaging methods—lab-based micro-CT, X-ray microscopy and synchrotron-based tomography—to examine the morphology of ophiuroid arms, and compare them with two-dimensional data obtained from SEM. We describe the advantages and disadvantages of each instrument and set of parameters in terms of the ease and efficiency of data collection for morphometric analyses. We present new morphological observations obtained by digital sectioning of three-dimensional images that could not be achieved with SEM. Overall, our findings suggest that three-dimensional imaging has a high potential to address the gaps in knowledge of the internal ophiuroid skeleton, which will be pivotal to providing morphological characters that will aid in phylogenetic reconstructions.
Two new myriotrochid genera and a new species are described on the basis of material collected during the Russian-German expeditions KuramBio and KuramBio II in the Kuril-Kamchatka Trench area. Within the family Myriotrochidae eight types of wheel-like deposits are distinguished. Psilotrochus gen. n. and Sonnetrochus gen. n. possess 10 tentacles and wheels of unique types. Currenly Myriotrochidae consists of 52 recent species belonging to 10 genera. Number of species decreases from 26 at depths 0–2000 m to 10 at 6001–8000 m, and remains equal to 10 at 8001–10,000 m. The most speciose genera Myriotrochus (17 species) and Prototrochus (16 species) are the most eurybathic (2– 9174 m and 192–10730 m respectively) and the most widespread being known in all five oceans. The myriotrochids are widely distributed in the hadal (ultra-abyssal) zone. They were found in 16 trenches. The wheels of the recent types appeared for the first time in the Middle Triassic. Some wheels from Upper Permian resemble the wheels of the recent species.
Inclusion of fossils can be crucial to address evolutionary questions, because their unique morphology, often drastically modified in recent species, can improve phylogenetic resolution. We performed a cladistic analysis of 45 cassidulids with 98 characters, which resulted in 24 most parsimonious trees. The strict consensus recovers three major cassiduloid clades, and the monophyly of the family Cassidulidae is not supported. Ancillary analyses to determine the sensitivity of the phylogeny to missing data do not result in significantly different topologies. The taxonomic implications of these results, including the description of a new cassiduloid family and the evolution of some morphological features, are discussed. Cassiduloids (as defined here) most probably originated in the Early Cretaceous, and their evolutionary history has been dominated by high levels of homoplasy and a dearth of unique, novel traits. Despite their high diversity during the Palaeogene, there are only seven extant cassiduloid species, and three of these are relicts of lineages dating back to the Eocene. Future studies of the biology of these poorly known species, some of which brood their young, will yield further insights into the evolutionary history of this group.
Starfish (order: Asteroidea) possess a complex endoskeleton composed of thousands of calcareous ossicles. These ossicles are embedded in a body wall mostly consisting of a complex collagen fiber array. The combination of soft and hard tissue provides a challenge for detailed morphological and histological studies. As a consequence, very little is known about the general biomechanics of echinoderm endoskeletons and the possible role of ossicle shape in enabling or limiting skeletal movements. In this study, we used high‐resolution X‐ray microscopy to investigate individual ossicle shape in unprecedented detail. Our results show the variation of ossicle shape within ossicles of marginal, reticular and carinal type. Based on these results we propose an additional classification to categorize ossicles not only by shape but also by function into ‘connecting’ and ‘node’ ossicles. We also used soft tissue staining with phosphotungstic acid successfully and were able to visualize the ossicle ultrastructure at 2‐μm resolution. We also identified two new joint types in the aboral skeleton (groove‐on‐groove joint) and between adambulacral ossicles (ball‐and‐socket joint). To demonstrate the possibilities of micro‐computed tomographic methods in analyzing the biomechanics of echinoderm skeletons we exemplarily quantified changes in ossicle orientation for a bent ray for ambulacral ossicles. This study provides a first step for future biomechanical studies focusing on the interaction of ossicles and soft tissues during ray movements.
Thyone and Havelockia are worldwide-distributed genera represented in the Western Atlantic by eight and three species, respectively. In this paper, we describe two new shallow-water species from the Brazilian coast: Thyone waltinhoi sp. nov. and Havelockia oraneae sp. nov. These genera are very similar morphologically but the presence of subdivisions in their calcareous rings separates them into different families, Thyone in Phyllophoridae and Havelockia in Sclerodactylidae. To observe the presence of subdivisions, we submerged the calcareous rings in an iodine solution prior to morphological analyses. Thyone waltinhoi sp. nov. differs from its Western Atlantic congeners in its dermal ossicles and Havelockia oraneae sp. nov. differs from its South-western Atlantic congeners in the composition of its introvert ossicles. In addition to species descriptions, illustrations and morphological diagnoses of both genera are provided, along with a synoptic table of their Western Atlantic species, and a discussion of the taxonomic significance of the dermal ossicles and the calcareous ring.
Here we use synchrotron radiation-based micro-computed tomography (SRµCT) images of type specimens to confidently place Cassidulus malayanus in a new genus (Kassandrina gen. nov.) that would not have been discovered with traditional techniques, and to describe a new species of Cassidulus (Cassidulus briareus sp. nov.) from Australia and designate a neotype for Cassidulus caribaearum. We also provide remarks describing the taxonomic history of each taxon and a diagnostic table of all living cassidulid species, and extend the known geographic and bathymetric range of C. caribaearum and C. malayanus. Besides rendering novel morphological data, the SRµCT images provide significant insights in the evolution of bourrelets of these cassiduloid echinoids.
The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data. In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.
Sea cucumbers (Holothuroidea) are a morphologically diverse, ecologically important, and economically valued clade of echinoderms; however, the understanding of the overall systematics of the group remains controversial. Here, we present a phylogeny of extant Holothuroidea assessed with maximum parsimony, maximum likelihood, and Bayesian approaches using approximately 4.3 kb of mt- (COI, 16S, 12S) and nDNA (H3, 18S, 28S) sequences from 82 holothuroid terminals representing 23 of the 27 widely-accepted family-ranked taxa. Currently five holothuroid taxa of ordinal rank are accepted. We find that three of the five orders are non-monophyletic, and we revise the taxonomy of the groups accordingly. Apodida is sister to the rest of Holothuroidea, here considered Actinopoda. Within Actinopoda, Elasipodida in part is sister to the remaining Actinopoda. This latter clade, comprising holothuroids with respiratory trees, is now called Pneumonophora. The traditional Aspidochirotida is paraphyletic, with representatives from three orders (Molpadida, Dendrochirotida, and Elasipodida in part) nested within. Therefore, we discontinue the use of Aspidochirotida and instead erect Holothuriida as the sister group to the remaining Pneumonophora, here termed Neoholothuriida. We found four well-supported major clades in Neoholothuriida: Dendrochirotida, Synallactida, Molpadida and a new taxon, Persiculida. The mapping of traditionally-used morphological characters in holothuroid systematics onto the phylogeny revealed marked homoplasy in most characters demonstrating that further taxonomic revision of Holothuroidea is required. Two time-tree analyses, one based on calibrations for uncontroversial crown group dates for Eleutherozoa, Echinozoa and Holothuroidea and another using these calibrations plus four more from within Holothuroidea, showed major discrepancies, suggesting that fossils of Holothuroidea may need reassessment in terms of placing these forms with existing crown clades.
The degree of isolation of oceanic islands makes these environments an excellent model for evolutionary studies. Proper knowledge of the species composition of oceanic islands, however, is required to better understand evolutionary processes (e.g. speciation events). A 3-year survey in the shallow waters (up to 30 m) of the Trindade and Martin Vaz oceanic insular complex, and a literature review on the data published for these islands and for Fernando de Noronha and São Pedro and São Paulo oceanic archipelagos have been conducted to document the biodiversity of echinoids and holothuroids from these isolated Brazilian oceanic islands. Sixteen species were collected and characterized morphologically, including two first records for the South Atlantic and one for Brazil. Comparison with conspecific specimens from the Brazilian coast and congeners was also done. Species richness increased from six to 18; the richness in Trindade Island being the highest among the South Atlantic oceanic islands. However, these islands remain undersampled beyond 30-m depth. Endemism was very low, suggesting the potential role of oceanic currents and seamounts as stepping-stones in transoceanic dispersal of species to remote islands. The Brazilian oceanic islands are impoverished oceanic outposts of the Brazilian Province; nevertheless, endemic species and intraspecific morphological variations compared with the mainland suggest they may also be regions of speciation. Documenting their biodiversity is critical for effective management and conservation of their marine ecosystems.
Echinoderm skeletons are described within a hierarchical framework ranging from complete organisms to the ultrastructural level. They consist of numerous elements which can be isolated, connected by soft tissue or locked together in rigid structures. The top level considers skeletons as a whole with all associated elements and the basic symmetry of the echinoderms. The next level deals with the structural analysis and modeling of echinoids with respect to the growth parameters and stress resistance of the corona. The flexibility and movement has also been studied for the stalks and arms of recent and fossil crinoids. The next level deals with the elaborate morphology and types of symmetry found in single skeletal elements. The numerous types of stereom architectures found within the elements of all echinoderms are highly correlated to specific functions. A high number of recent studies concern the last hierarchical level on ultrastructure and biomineralization. Lightweight aspects of the skeleton are especially present at the level of conjoined plates, single elements and the stereom.
In a detailed study of the Blue Lias (Hettangian-Sinemurian) at four localities in southern Britain holothurian sclerites were found in 76% of samples, sometimes in abundance. There are eleven sclerite morphospecies, four of them new. Four types of calcareous ring are described and, where possible, their associated sclerite morphospecies identified. The fauna is of limited palaeoenvironmental and biostratigraphical use, although changes in size frequency and relative abundance are potentially informative for the latter. Assemblages are difficult to recognize possibly due to taphonomic effects. The fauna is broadly similar across northwestern Europe but with regional variations both in sclerite composition and abundance. -from Author
In this paper, we propose the reassignment of three western Atlantic species of the order Dendrochirotida to the family Sclerodactylidae (Euthyonidiella occidentalis comb. nov., Euthyonidiella arenicola comb. nov., and Thandarum manoelina comb. nov.) with a discussion on the classification of the dendrochirotids based on the morphology of the calcareous ring. We also propose the synonymization of Euthyonidiella dentata with Euthyonidiella occidentalis comb. nov., desig- nate a neotype for this species and for Thandarum manoelina, and describe a new species of the genus Cucumaria from Brazil
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
Echinoderms are a major group of invertebrate deuterostomes that have been an important component of marine ecosystems throughout the Phanerozoic. Their fossil record extends back to the Cambrian, when several disparate groups appear in different palaeocontinents at about the same time. Many of these early forms exhibit character combinations that differ radically from extant taxa, and thus their anatomy and phylogeny have long been controversial. Deciphering the earliest evolution of echinoderms therefore requires a detailed understanding of the morphology of Cambrian fossils, as well as the selection of an appropriate root and the identification of homologies for use in phylogenetic analysis. Based on the sister-group relationships and ontogeny of modern species and new fossil discoveries, we now know that the first echinoderms were bilaterally symmetrical, represented in the fossil record by Ctenoimbricata and some early ctenocystoids. The next branch in echinoderm phylogeny is represented by the asymmetrical cinctans and solutes, with an echinoderm-type ambulacral system originating in the more crownward of these groups (solutes). The first radial echinoderms are the helicoplacoids, which possess a triradial body plan with three ambulacra radiating from a lateral mouth. Helicocystoids represent the first pentaradial echinoderms and have the mouth facing upwards with five radiating recumbent ambulacra. Pentaradial echinoderms diversified rapidly from the beginning of their history, and the most significant differences between groups are recorded in the construction of the oral area and ambulacra, as well as the nature of their feeding appendages. Taken together, this provides a clear narrative of the early evolution of the echinoderm body plan.
In this paper, we diagnose a new genus of Sclerodactylidae, Coronatum gen. et sp. nov., from shallow waters of the southwestern Atlantic Ocean. Coronatum baiensis sp. nov. has a compact calcareous ring with short posterior processes and a unique set of body wall ossicles, composed of two-pillared tables, which do not have any morphological affinities with Euthyonidiella dentata and Pseudothyone belli, the other two known Brazilian sclerodactylid species. Identification keys and photographs of living specimens and ossicles are provided. This paper increases to twelve the number of known Sclerodactylinae genera and to two the number of species of this subfamily in this region.
The fossil echinoderm Palaeocucumaria, from the early Devonian Hunsrück Slate of southwestern Germany, has been studied using both traditional techniques and X-ray microtomography, and its anatomy clarified. Phylogenetic analysis shows that it is a stem-group holothurian with a combination of characters that help understand how the modern (crown-group) holothurian body plan developed. Echinoids and holothurians have evolved along different paths, by differential growth of the larval- and rudment-derived body regions. Palaeocucumaria shows that late stem-group holothurians had a water vascular organization with a single external madreporite and calcified stone canal leading to the aboral end of the peripharyngeal coelom, and five primary radial water vessels that gave rise to tentacle-like tube-feet. This fossil data, in combination with a molecular phylogeny based on 18 s-like rRNA gene sequence data, is used to order evolutionary steps in the making of the crown-group holothurian body plan.
Recent sampling in the Rockall Trough, Porcupine Seabight and Porcupine Abyssal Plain, in the NE Atlantic, has yielded 200 specimens of apodous holothurians belonging to seven species of the family Myriotrochidae Théel from depths between 1000 and 4310 m. These include the type species of a new genus and two new species of existing genera. Parvotrochus belyaevi gen. et sp. nov. is described from some minute specimens from the Rockall Trough that possess both exceptionally small wheel-like deposits and numerous, large curved rods in the body wall and tentacles. Myriotrochus clarki sp. Nov. has affinities to M. vitreus (M. Sars) but shows differences in the shape of the plates of the calcareous ring and in having larger, frequently abnormally formed, wheels. Siniotrochus myriodontus sp. Nov. is similar to the type species of this genus, S. phoxus, Pawson, but differs in the arrangement of the teeth on the wheels. A single small specimen of Prototrochus Belyaev et Mironov from the Whittard Canyon, northern Bay of Biscay, is similar to P. minutus (Östergren), a species known only from the Sea of Japan, but differs in some characteristics of the wheels and tentacles. A new subspecies, P. zenkevitchi rockallensis subsp. nov., is proposed for a form of P. zenkevitchi (Belyaev) that was the most common myriotrochid encountered. P. zenkevitchi was known previously only from a few localities in the Pacific and S Atlantic deep-sea trenches. Similarly, two species previously known only from the NE Pacific, Myriotrochus bathybius H. L. Clark and M. giganteus H. L. Clark, are also recorded from the N Atlantic for the first time. The synonymy of M. giganteus and M. sp. ex gr. macquariensis-giganteus Belyaev et Mironov is proposed.
The greatest number both of specimens and species came from the areas most intensively sampled. The present records show that several species have world-wide distributions. The wide geographic separation of many records is the result of both poor sampling effort in the deep sea, particularly with fine-meshed gear, and the difficulties in sampling infaunal animals in the deep sea. Box core samples in the Rockall Trough suggest that the myriotrochids are more common in this area than would be supposed from epibenthic sledge data. Hence, myriotrochids may be more prevalent in the deep sea than previously thought.
(1) Catch connective tissue is defined as the collagenous connective tissue whose mechanical properties can be changed rapidly (in seconds or minutes) under nervous control.
(2) Catch connective tissues are found in all five classes of Echinodermata. They function in tone control of the tissues and in autotomy.
(3) The change in mechanical properties occurs in viscosity.
(4) Muscle cells are not responsible for the viscosity change.
(5) The viscosity change is controlled by nervous activities. Neurosecretory-like cells with large electron-dense granules are found in all the catch connective tissues so far studied.
(6) The viscosity change is quite likely caused by the change in the ionic environment in the connective tissues, which alters the weak (non-covalent) interactions between extracellular macromolecules in the tissue.
Logical equivalence between the notions of homology and synapomorphy is reviewed and supported. So‐called transformational homology embodies two distinct logical components, one related to comparisons among different organisms and the other restricted to comparisons within the same organism. The former is essentially hierarchical in nature, thus being in fact a less obvious form of taxic homology. The latter is logically equivalent to so‐called serial homology in a broad sense (including homonomy, mass homology or iterative homology). Of three tests of homology proposed to date (similarity, conjunction and congruence) only congruence serves as a test in the strict sense. Similarity stands at a basic level in homology propositions, being the source of the homology conjecture in the first place. Conjunction is unquestionably an indicator of non‐homology, but it is not specific about the pairwise comparison where non‐homology is present, and depends on a specific scheme of relationship in order to refute a hypothesis of homology. The congruence test has been previously seen as an application of compatibility analysis. However, congruence is more appropriately seen as an expression of strict parsimony analysis. A general theoretical solution is proposed to determine evolution of characters with ambiguous distributions, based on the notion of maximization of homology propositions. According to that notion, ambiguous character‐state distributions should be resolved by an optimization that maximizes reversals relative to parallelisms. Notions of homology in morphology and molecular biology are essentially the same. The present tendency to adopt different terminologies for the two sources of data should be avoided, in order not to obscure the fundamental uniformity of the concept of homology in comparative biology.
“A similar hierarchy is found both in ‘structures’ and in ‘functions’. In the last resort, structure (i.e. order of parts) and function (order of processes) may be the very same thing […].”
L. von Bertalanlfy
“[…] it is the fact that certain criteria enable us to match parts of things consistently which suggests that mechanisms of certain kinds must have been involved in their origin.”
N. Jardine and C. Jardine