Extremes of Pit Infestation and Growth Deformity in a Crinoid Column,Permian of Timor

Crinoids are diverse and well-known from the Permian of Timor, but the literature has failed to document the numerous specimens of crinoid pluricolumnals from the fauna, many showing unusual morphology or yielding palaeoecological information. A curious and instructive specimen demonstrates the relationship between a living Permian crinoid and coeval invasive, pit-forming, invertebrates in detail. The pit-former is not preserved; most likely it was unmineralized or, if mineralized, then the shell simply dropped out. The infesting organism made pits assigned to the ichnospecies Oichnus paraboloides Bromley. The pit-former was unusually site selective. Either (1) one spatfall attached to just one side of the elevated (either up-current or down-current) or recumbent column and each individual centered their pits on the sutures between adjacent columnals; or (2) a single individual migrated along the column. The living crinoid showed an extreme reaction to this infestation. Excess stereom growth on the side of the pits transformed what was a circular column by addition of a thick, triangular ridge on the pitted side.     

Epi- and endobiontic organisms on Late Jurassic crinoid columns from the Negev Desert, Israel: Implications for co-evolution

Columns of the articulate crinoids Millericrinus and Apiocrinites from the Upper Jurassic (Upper Callovian) Zohar and Matmor formations of the Negev Desert of Israel display abundant encrusting organisms of about ten species, as well as diverse trace fossils produced by endobionts. Pluricolumnals were colonized by epi- and endobiontic organisms both during life and post-mortem. Skeletonized encrusting organisms include abundant ostreid bivalves (which evidently colonized both live and dead crinoid columnals), two types of serpulid worms, encrusting foraminifera, three species of bryozoans, and small encrusting sclerosponges. Several types of borings are present: Trypanites (possibly produced by sipunculids), Gastrochaenolites (crypts of boring lithophagid bivalves), elliptical barnacle? borings, and channel-like annelid? borings. In addition, approximately 16% of the pluricolumnals display circular parabolic embedment pits assignable to the ichnogenus Tremichnus. They are associated with substantial deformation of the containing columnals and were probably the work of host-specific ectoparasitic organisms. Discovery of Tremichnus on Jurassic crinoids extends the range of this trace by almost 100 million years, providing evidence for one of the longest-ranging host-parasite interactions documented thus far (over 200 million years). The relationship of epibionts to the Jurassic crinoids thus ranged from simple utilization of dead hard substrate to probable opportunistic commensalism in forms that colonized the live upright stems, as in some oysters, through host-specific parasitism in the case of Tremichnus.     

Stereom morphogenesis and differentiation during regeneration of adambulacral spines of Asterias rubens (Echinodermata,Asteroida)

Summary The very first mineral deposits appearing in regenerating fractured adambulacral spines of Asterias rubens are minute polyhedrons that cover the surface of fractured trabeculae. Polyhedrons fuse together forming a fold from which a microspine differentiates. Microspines develop into long linear trabeculae which send out lateral processes at regular length intervals. Lateral processes from adjacent trabeculae fuse together, bridging the trabeculae and giving the regenerate the typical meshwork structure of stereom. Most of the regenerate is built up according to this growth pattern which ensures its longitudinal growth. Simultaneously, the initial fascicular stereom of the stub sends out short radial processes which branch into upward and downward directed subprocesses. The latter fuse with their equivalents located above or below, building up longitudinal rows of stereom meshes. These rows then bridge together by additional branched or unbranched lateral processes, so forming a new stereom layer which progressively covers the whole stub. Up to three new layers of stereom are formed in this way at the stub periphery. These become continuous with the stereom layers of the regenerate by fusion of reciprocal subprocesses, so ensuring the continuity between the stub and the regenerate. In both structures the first stage of mineralization results in an open stereom. Stereom thickening occurs in a second stage of mineralization (that is chronologically separated from the formation of the open stereom) and results in the differentiation of the original stereom fabrics (i.e. fascicular stereom). Regeneration of removed spines starts with the formation of a new spine base made of labyrinthic stereom. The development of the latter mostly relies on short branched and unbranched processes which fuse with each other or with predifferentiated meshes. After completion of its base, the regenerating spine lengthens and thickens similarly to the regenerating fractured spines. The diversity of the stereom growth processes observed in the present work may be reduced to the combination of one to three elementary events, viz. the development of long linear processes, of short unbranched processes and of short branched processes. A survey of the literature allows the suggestion that the implementation of these elementary events is sufficient to describe most types of stereom morphogenesis.Senior research assistant NFSR (Belgium)     

²⁶Mg labeling of the sea urchin regenerating spine: Insights into echinoderm biomineralization process

This paper reports the results of the first dynamic labeling experiment with regenerating spines of sea urchins Paracentrotus lividus using the stable isotope ²⁶Mg and NanoSIMS high-resolution isotopic imaging, which provide a direct information about the growth process. Growing spines were labeled twice (for 72 and 24 h, respectively) by increasing the abundance of ²⁶Mg in seawater. The incorporation of ²⁶Mg into the growing spines was subsequently imaged with the NanoSIMS ion microprobe. Stereom trabeculae initially grow as conical micro-spines, which form within less than 1 day. These micro-spines fuse together by lateral outgrowths and form a thin, open meshwork (inner stereom), which is subsequently reinforced by addition of layered thickening deposits (outer stereom). The (longitudinal) growth rate of the inner stereom is ca. 125 μm/day. A single (ca. 1 μm) thickening layer in the stereom trabeculae is deposited during 24 h. The thickening process is contemporaneous with the formation micro-spines and involves both longitudinal trabeculae and transverse bridges to a similar degree. Furthermore, the skeleton-forming cells remain active in the previously formed open stereom for at least 10 days, and do not migrate upwards until the end of the thickening process. The experimental capability presented here provides a new way to obtain detailed information about the skeleton formation of a multitude of marine, calcite producing organisms.     

Fine structure of the dorsal arm plate of Ophiocoma wendti: Evidence for a photoreceptor system (Echinodermata,Ophiuroidea)

Summary Three structures in the dermis of the dorsal arm plate (DAP) of the brittlestar, Ophiocoma wendti, appear to comprise a photoreceptor system. The upper surface of the DAP bears transparent, knob-like, microscopic structures which are expanded peripheral trabeculae (EPT) of the calcite stereom. The EPT are part of the photoreceptor system and can facilitate light transmission through the DAP by decreasing light refraction, reflection and absorption that occur at stereom/stroma interfaces. Bundles of nerve fibres located below the EPT are a second component of the system, and may function as primary photoreceptors. The intensity of light impinging on the putative sensory tissue is regulated by the diurnal activity cycle of chromatophores, the third element of the system. During the day the chromatophores cover the EPT and thereby shade the nerve fibres. At night they retract into inter-trabecular channels, uncovering the EPT and thereby exposing the nerve fibres to transmitted light. Thus, the transparent stereom may play a role in photoreception, in addition to its generally recognized skeletal function. Although ciliated cells that may be sensory are present in the epidermis of Ophiocoma wendti, they do not appear to be photoreceptors. Functional analogues of the brittlestar photoreceptor system in other echinoderms are discussed, emphasizing the relationship between photosensitivity and the transparency of the stereom in several classes of Echinodermata.     

Anatomical features of the euryalid snake star Astrobrachion constrictum (Ophiuroidea: Asteroschematidae)

Abstract. The morphology of Astrobrachion constrictum , a representative of the little known Euryalida, was examined by light and electron microscopy. The tegument is smooth and lacks calcified elements. Many features of A. constrictum were found to conform to the usual pattern of ophiuroid anatomy. The skeletal elements are generally composed of uniform labyrinthic stereom. The radial shields, however, consist of overlapping stereom plates. The 5 oral shields serve as madreporites, each with a U-shaped canal leading into an axial sinus. A 3-lobed axial organ is associated with each madreporite and may play some sensory or secretory role.     

Middle Cambrian cystoid (sensu lato) stem columnals from Bornholm, Denmark

A rich material of echinoderm fragments from two Middle Cambrian stratigraphical levels on Bornholm are preserved due to phosphatization of the original calcitic stereom. Preservation of echinoderms in this way - not previously recorded from the Middle Cambrian - permits detailed analysis of the three-dimensional stereom structure. Identifiable are fragments of stylophorans and eocrinoids. Stem columnals, most likely from eocrinoids, show a wide and advanced morphological variation indicating articulation similar to that of crinoids. The material from the Exsulans Limestone/Kalby marl ( Ptychagnostus gibbus Zone) represents stem-bearing cystoids older than Akadocrinus from Bohemia. The Andrarum Limestone ( Sole-nopleura brachymetopa Zone) contains echinoderm fragments from a higher stratigraphical level, a level correlatable with that from which the oldest North American stem–bearing cystoid, Eustypocystis , has been recorded.     

The attachment of collagenous ligament to stereom in primary spines of the sea-urchin, Eucidaris tribuloides

The similar proximal and distal attachments to the stereom of primary spine ligament in the echinoid Eucidaris tribuloides are described, from thin sections and SEM studies on frozen and fractured spine articulations and ligaments from decalcified material. The orthogonal structure of the general stereom is modified on the attachment zones where bundles of collagen cylinders enter approximately hexagonally arranged channels. Straps of collagen extend in parallel series between adjacent bundles via regularly placed ports and collagen loops rather than non-striated 'tendons' pass over skeletal trabeculae. The regular pattern of collagen straps is most evident on the proximal and distal attachment zones. Mechanical features of the non-adhesive mode of attachment are considered, together with similarities and differences between insertion of muscle cells and mutable collagenous tissue (ligament) in echinoderms.     

Ultrascale and microscale growth dynamics of the cidaroid spine of Phyllacanthus imperialis revealed by 26Mg labeling and NanoSIMS isotopic imaging

Growth dynamics of the primary spine of the cidaroid sea urchin Phyllacanthus imperialis was assessed for the first time using pulsed ²⁶Mg‐labeling and NanoSIMS isotopic imaging. The sea urchin was incubated twice (for 48 h) in artificial seawater with elevated level of ²⁶Mg. After each labeling event, the sea urchin was returned for 72 h to seawater with natural isotopic abundance of ²⁶Mg. NanoSIMS ion microprobe was subsequently used to visualize the labeled regions of the spine with submicrometer lateral resolution. The growth of the new skeleton was restricted to the distalmost and peripheral portions of the spine. Skeletogenesis involved mostly the deposition of continuous thickening layers and lateral growth involving bridges between previously formed trabeculae. The timescale of formation of individual thickening layers (ca. 1 µm in width) on the stereom trabeculae was on the order of 1 day. Longitudinal growth occurred mainly at the periphery in the form of small portions of the thickening deposits or more massive microspines that appeared to branch and fuse with those above and below. These microspines were found to grow at about 10 µm/day. These results reveal that the skeletal growth of a juvenile cidaroid spine is complex and highly heterogeneous, with different extension rates depending on the stage of the stereom development and/or direction of the growth fronts. The growth pattern observed here at the submicrometer scale provides direct evidence supporting the earlier suggestions that the lamellar structure of echinoderm stereom is formed by periodic deposition of continuous mineral layers. J. Morphol. 275:788–796, 2014. © 2014 Wiley Periodicals, Inc.     

Skeleton resorption in echinoderms: Regression of pedicellarial stalks inSphaerechinus granularis (Echinoida)

Summary When a globiferous pedicellaria ofSphaerechinus granularis injects its venom, the head autotomizes whereas the stalk remains on the test and enters a regression process with concomitant resorption of its supporting ossicle (i.e. the rod). Scanning electron microscope investigations of the morphological changes undergone by the stereom of resorbing rods show that: (1) resorption proceeds both axially and laterally, and leads to a reduction of approximately 80% of the original length of the rod, (2) secondary growth of new stereom processes occurs concomitantly with resorption but never ensures even a partial regeneration of the rod, and (3) resorption and secondary growth stop before the rod is totally destroyed leaving a static stump that remains in place up to 190 days. Particular resorption figures result from either the axial or the lateral resorption of the rod shaft. These consist chiefly of terraced conical cupules, dense cylinders and concentric lamellae whose walls or edges are typically made of closely piled and/or aligned subprismatic crystallites. Whatever their location along the rod, these crystallites always organize strictly parallel to the rod axis. Whether the crystallites are mosaic blocks composing larger monocrystalline units or discrete monocrystals themselves is for the moment unclear. A growth model, which accounts for the observed resorption figures, is proposed for the shaft of pedicellarial rods. According to this model, the early growth of the shaft would produce elongated, interconnected trabeculae (initial trabeculae) made of densely piled and perfectly aligned crystallites. Thickening and coalescence of adjoining trabeculae would progressively occur by adjunction around the initial trabeculae of successive and concentric layers of similarly arranged crystallites. Coalescent trabeculae would then be cemented together in a perforate stereom layer by the final deposition of larger crystallite layers surrounding the whole shaft periphery. Growth of secondary stereom processes occurs both in the resorbing rod (here the newly formed processes are resorbed soon after they have been produced) and in rods where resorption has stopped. These are always irregular processes that localize near or on the actual sites of resorption. It is suggested these processes result from an uncontrolled activation of the skeleton-forming cells in areas where the concentration of calcium ions increases as a consequence of calcite resorption.     

Die Feinstruktur der Gallenblase und der Gallengänge

Zusammenfassung Die Feinstruktur des Ductus hepaticus communis, des Ductus cysticus und des Ductus choledochus der Maus sowie der Ductus hepatici und des Ductus choledochus der Ratte werden untersucht.Im Ductus hepaticus communis und im Ductus cysticus der Maus kommen Zellen vor, die den Hauptzellen der Gallenblase vergleichbar sind. Außerdem sind in beiden Ductus und vorwiegend im Ductus choledochus neutrale Mukosubstanzen sezernierende Zellen sowie Wanderzellen, Bürstenzellen, Becherzellen und basalgekörnte Zellen vorhanden. Die letztgenannten drei Zelltypen befinden sich nur in einigen Abschnitten der untersuchten Gänge.Die Feinstrukturen des Epithels der Ductus hepatici der Ratte ähneln jenen des Epithels des Ductus hepaticus der Maus. Der Ductus choledochus enthält dagegen vorwiegend Zellen, die reich an Filamenten und Desmosomen sind. Auch bei der Ratte sind Wanderzellen, Bürstenzellen, Becherzellen und basalgekörnte Zellen festzustellen.Die unterschiedliche Struktur des Ductus choledochus von Maus und Ratte wird diskutiert.

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On the fine structure of gall bladder and biliary ducts

Summary Fine structures of the ductus hepaticus communis, ductus cysticus and ductus choledochus of mice as well as of the ductus hepatici and ductus choledochus of rats have been investigated.In the ductus hepaticus communis and ductus cysticus of the mouse there are cells that are similar to the principal cells of the gall bladder epithelium. In addition both these ducts and especially the ductus choledochus contain cells with mucigen granules. There are three further cell types, irregularly distributed in the epithelium of the three ducts: brush cells, goblet cells and basal granulated cells. Macrophages (wandering cells) are present in epithelia.In the ductus hepatici of the rat the epithelial cells are displaying fine structures similar to those of the ductus hepaticus of the mouse. The ductus choledochus however presents cells with many filaments and desmosomes. Again between these cells there are macrophages, brush cells, goblet cells and basal granulated cells.The differences in the fine structure of the epithelium of the ductus choledochus of mouse and rat are discussed.

Herrn Prof. Dr. H. Ruska danke ich für die wertvolle Diskussion des Manuskriptes.     

Microscopical Anatomy of the Cyrtocrinid Cyathidium meteorensis (sive foresti) (Echinodermata,Crinoidea)

Light microscopy and scanning electron microscopy of three specimens of Cyathidium meteorensis (order Cyrtocrinida) revealed some special morphological features. The brachial articulation is provided with a long tendon at the aboral side; the entire articulation surface, including the areas where ligaments attach is built up by labyrinthic stereom. The calycinal ossicle lacks any internal vestiges of a pentameric composition; vertical planes with changing stereom direction lie irrespectively of radial or interradial planes. Gut, ovary and testis are histologically inconspicuous, and the location of the gonads inside the calyx is quite unusual. Both sexes develop outer gonoducts which probably are functionally adapted madreporic canals. The coelomic system differs from that of other crinoids in that a chambered organ is completely lacking. Simultaneously, the aboral nervous subsystem has no aboral nerve centre and simply terminates aborally in the ring-shaped commissure. A glandular axial organ is absent, as are typical sacculi. The data are in accordance with two proposals made previously for Holopus rangii, viz., that the animals can feed raptorially, and that cyrtocrinids probably have evolved by loss of aboral calycinal ossicles of ancestors. In part, our observations differ from those in Cyathidium foresti, so we have chosen to use the species name meteorensis which has been considered a synonym of foresti.     

Micro‐ to nanostructure and geochemistry of extant crinoidal echinoderm skeletons

This paper reports the results of micro‐ to nanostructural and geochemical analyses of calcitic skeletons from extant deep‐sea stalked crinoids. Fine‐scale (SEM, FESEM, AFM) observations show that the crinoid skeleton is composed of carbonate nanograins, about 20–100 nm in diameter, which are partly separated by what appears to be a few nm thick organic layers. Sub‐micrometre‐scale geochemical mapping of crinoid ossicles using a NanoSIMS ion microprobe, combined with synchrotron high‐spatial‐resolution X‐ray micro‐fluorescence (μ‐XRF) maps and X‐ray absorption near‐edge structure spectroscopy (XANES) show that high Mg concentration in the central region of the stereom bars correlates with the distribution of S‐sulphate, which is often associated with sulphated polysaccharides in biocarbonates. These data are consistent with biomineralization models suggesting a close association between organic components (including sulphated polysaccharides) and Mg ions. Additionally, geochemical analyses (NanoSIMS, energy dispersive spectroscopy) reveal that significant variations in Mg occur at many levels: within a single stereom trabecula, within a single ossicle and within a skeleton of a single animal. Together, these data suggest that physiological factors play an important role in controlling Mg content in crinoid skeletons and that great care should be taken when using their skeletons to reconstruct, for example, palaeotemperatures and Mg/Ca palaeo‐variations of the ocean.     

Properties,morphogenesis, and effect of acidification on spines of the cidaroid sea urchin Phyllacanthus imperialis

Cidaroid sea urchins are the sister clade to all other extant echinoids and have numerous unique features, including unusual primary spines. These lack an epidermis when mature, exposing their high‐magnesium calcite skeleton to seawater and allowing the settlement of numerous epibionts. Cidaroid spines are made of an inner core of classical monocrystalline skeleton and an outer layer of polycrystalline magnesium calcite. Interestingly, cidaroids survived the Permian‐Triassic crisis, which was characterized by severe acidification of the ocean. Currently, numerous members of this group inhabit the deep ocean, below the saturation horizon for their magnesium calcite skeleton. This suggests that members of this taxon may have characteristics that may allow them to resist ongoing ocean acidification linked to global change. We compared the effect of acidified seawater (pH 7.2, 7.6, or 8.2) on mature spines with a fully developed cortex to that on young, growing spines, in which only the stereom core was developed. The cortex of mature spines was much more resistant to etching than the stereom of young spines. We then examined the properties of the cortex that might be responsible for its resistance compared to the underlying stereomic layers, namely morphology, intramineral organic material, magnesium concentration, intrinsic solubility of the mineral, and density. Our results indicate that the acidification resistance of the cortex is probably due to its lower magnesium concentration and higher density, the latter reducing the amount of surface area in contact with acidified seawater. The biofilm and epibionts covering the cortex of mature spines may also reduce its exposure to seawater.     

X-ray microCT study of pyramids of the sea urchin Lytechinus variegatus

This paper reports results of a novel approach, X-ray microCT, for quantifying stereom structures applied to ossicles of the sea urchin Lytechinus variegatus. MicroCT, a high resolution variant of medical CT (computed tomography), allows noninvasive mapping of microstructure in 3-D with spatial resolution approaching that of optical microscopy. An intact pyramid (two demipyramids, tooth epiphyses, and one tooth) was reconstructed with 17 microm isotropic voxels (volume elements); two individual demipyramids and a pair of epiphyses were studied with 9-13 microm isotropic voxels. The cross-sectional maps of a linear attenuation coefficient produced by the reconstruction algorithm showed that the structure of the ossicles was quite heterogeneous on the scale of tens to hundreds of micrometers. Variations in magnesium content and in minor elemental constitutents could not account for the observed heterogeneities. Spatial resolution was insufficient to resolve the individual elements of the stereom, but the observed values of the linear attenuation coefficient (for the 26 keV effective X-ray energy, a maximum of 7.4 cm(-1) and a minimum of approximately 2 cm(-1) away from obvious voids) could be interpreted in terms of fractions of voxels occupied by mineral (high magnesium calcite). The average volume fraction of mineral determined for a transverse slice of the demipyramid near where it joins an epiphysis was 0.46; for a slice 3.3 mm adoral it was 0.70. Local volume fractions of mineral approached 1, and, away from resolvable voids, considerable portions of the demipyramids had volume fractions of calcite at or below approximately 0.33. MicroCT imaging of a demipyramid before and after infiltration with a high absorptivity fluid (sodium polytungstate) confirmed the determination of the volume fractions of minerals.     

Effects of larval antlions Euroleon nostras (Neuroptera, Myrmeleontidae) and their pits on the escape-time of ants

Abstract. Pit-building antlions are predators with a unique predation strategy, namely using pitfall traps constructed in loose sand to catch prey. Here, prey escape-time in the field is measured by introducing ants into one of four different treatment arenas. The first treatment lacks pits and antlions, the second includes 10 antlions that did not build pits, the third comprises eight artificially constructed pits, and the fourth is a treatment of eight antlions in pits and two without pits. Their pits are of a similar size to those used in the third treatment. When antlions are present without constructing pits, they impede the dispersal of prey. The mean escape-time for one half of the released ants is twice as long with antlions present as without them. When pits are present, the time taken for one half of the released ants to escape the predator is more than 10 times as long as when pits are absent. Escape-time from artificial pits is three times that from nonpit building antlions. Pits hinder the escape of ants and therefore increase the amount of time that the prey is available for capture. In the area where the pits are occupied by antlions, escape-time is four times longer than in a treatment with similar sized artificial pits. Thus, it appears that not only a pit, but also the presence of antlions influences the capture success.     

Cleaning Up After a Meal: The Consequences of Prey Disposal for Pit‐Building Antlion Larvae

Predators use a variety of strategies for capturing prey. Trap‐building predators can save on searching and encountering costs by investing in the construction and maintenance of traps such as webs and pits. However, what to do with partially consumed prey poses a potential problem. Antlion larvae (Myrmeleon acer) catch ants in conical pits, and dispose of partially consumed carcasses by flicking them a short distance away. We tested whether this prey‐disposal behaviour affects the effectiveness of antlion pits. We observed ant behaviour around artificially constructed pits and compared falls into pits with clean margins to those with conspecific ant carcasses or control objects around the pit edge. The presence of objects near pits affected the behaviour of live ants, and reduced the effectiveness of pits. Live ants spent the most time examining fresh ant carcasses, but the presence of any object near pits deterred pitfalls. Ants fell into pits significantly more often when pit edges were clean, suggesting that antlions could incur a prey capture cost from flicking carcasses from pits as well as from the accumulation of other debris around pit margins.     

红树族植物次生木质部附物纹孔的电镜观测

应用扫描电子显微镜详细观察了红树族4属、10种、1变种植物次生木质部管状分子附物纹孔的分布和形态, 应用Carnoy 2.0软件和扫描电镜下采集的照片, 测定了管间梯状附物纹孔丰富度指标和管间梯状纹孔数量特征指标。结果显示, 红树族植物次生木质部管状分子侧壁具附物纹孔。所观察的植物附物纹孔的分布和形态变化大。附物纹孔丰富度指标与管间梯状纹孔数量特征指标的逐步回归分析表明, 导管侧壁附物纹孔丰富度随纹孔口面积百分比的增大而增大。据此推测, 红树族植物附物纹孔丰富度与纹孔几何构造及数量特征有关。附物纹孔是红树族植物稳定存在的一个木材解剖性状。综合生态-系统演化的观点, 红树族植物具附物纹孔可能是受系统演化关系控制的生态适应结果。