The water vascular system and functional morphology of Paleozoic asteroids

Asteroids of all geologic ages share a single basic body form, surficial skeletal arrangement, and aspects of water vascular construction. In almost all described Paleozoic species, however, either podial pores to the interior of the arm were lacking, or they are directed laterally, above the adambulacrals. They are internal and above the ambulacrals in known post-Paleozoic species and the Pennsylvanian Calliasterella. Certain features of the ambulacral skeletal arrangement also differ. Calliasterella is the closest known Paleozoic relative of post-Paleozoic asteroids. Classifications of asteroids that stress only overall form and surticial skeletal arrangement erroneously include Paleozoic and Holocene species in common ordinal or even lower level groupings. Taxonomic revision is premature: however, most known Paleozoic asteroids represent primitive lineages. Transitional forms allow reconstruction of events leading to the modern arrangement. Ampullar and skeletal arrangements of post-Paleozoic asteroids appear to offer some functional advantages over those of their precursors, but as early as the Ordovician, diverse feeding habits had evolved and ecological roles paralleled those of Holocene species.     

<Emphasis Type="Italic">Compsaster formosus</Emphasis><Emphasis Type="SmallCaps">Worthen &; Miller</Emphasis> (Asteroidea; Echinodermata): A Carboniferous homeomorph of the post-Paleozoic Asteriidae

All adequately known post-Paleozoic asteroids are either assignable to surviving families or closely related families whereas no Paleozoic species assignable to a surviving order has been recognized. The Mississippian speciesCompsaster formosus is similar enough to various Recent taxa in overall form as well as in the form and arrangement of body wall ossicles to raise the issue of affinities: IsC. formosus nested within a phylogenetic branch hitherto known only from post-Paleozoic strata or is it only homeomorphic?The nature of the ambulacral system is critical to interpretation of echinoderms, and post-Paleozoic asteroids share three fundamental ambulacral characters or character suites: dorsal podial pores, staggered positioning of ambulacrals and adambulacrals, and complex articular structures between these two ossicular types.Calliasterella americana, a Carboniferous asteroid, shares the three ambulacral features, although it is distinctive from post-Paleozoic asteroids in other ways.Compsaster formosus exhibits at least two of the three ambulacral characters, although presence of staggering has not been finally established. LikeC. americana, C. formosus differs from post-Paleozoic species in details of ambulacral anatomy as well as aspects of ventral body surface ossicular arrangement. Although approaching crown-group organization,C. formosus nevertheless represents a branch basal to the crown group.Because asteroids are generalists, understanding of life habits ofCompsaster is sketchy in spite of morphological similarities between it and younger genera.Compsaster probably was epifaunal and its overall form is strongly reminiscent of that of Recent predatory asteriids but it is also similar to small-particle feeding echinasterids.     

On the Occurrence and Structure of Choanocyte-like Cells in Some Echinoderms

Choanocyte-like cells with a collar of regularly arranged cylindrical microvilli around the base of the flagellum were observed in the ciliary bands of the Brachiolaria larva of Asterias rubens. The ambulacral ampullae and coelomic epithelia of adult Asterias and coelomic epithelia of Mesothuria contain similar cells with radial lamellae instead of cylindrical microvilli. Other similar but more modified types of cells, in which the inner edges of the radiating lamellae could be recognized as longitudinal ridges in the wall of a cylindrical flagellar pit, were found in ambulacral ampullae of Porania and in coelomic epithelia of Stichopus. Distinct indications of phagocytosis were seen in most of these cells. The present report together with previously published data lend support to the idea, that the choanocyte is a fundamental cell type in metazoans, probably derived phylogenetically from some flagellate ancestor.     

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.     

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.     

What makes an ophiuroid? A morphological study of the problematic Ordovician stelleroid Stenaster and the palaeobiology of the earliest asteroids and ophiuroids

Extant asteroids and ophiuroids [EchinodermataJ are distinguished by differences in arm support, water vascular system structures and in details of arm and jaw structure. However, some lower Palaeozoic taxa show combinations of both asteroid-like and ophiuroid-like characters and their morphology and functional biology is poorly understood. This paper redescribes one such taxon, the middle-upper Ordovician stellate echinoderm Stenaster and clarifies its phylogenetic status. Characters in common with extant and Ordovician ophiuroids, include arm support due primarily to ambulacral ossicles, presence of extensive longitudinal arm musculature, a mobile jaw and an internalised radial water vessel with internalised podial pores. In addition, Stenaster lacks several characters which are conventionally considered to be asteroid-like, for example an axillary, madreporitc, marginal ossicles and a true ambulacral groove. However, in overall shape Stenaster is remarkably asteroid-like, showing short, broad-based arms shared podial basins and a small disc. A cladistic analysis of early asteroids, ophiuroids and somasteroid taxa consistently places Stenaster within the ophiuroids and suggests secondary convergence to asteroids. In functional terms, Stenaster is interpreted as an ophiuroid which has secondarily adopted a semi-infaunal, deposit-feeding mode of life, analogous to that of some extant paxillosid asteroids.     

Types of locomotion in ophiurans

Motion pictures were taken of the locomotion of two species of ophiurans living in the Sea of Japan:Ophiura sarsi vadicoa Djakonov andAmphipholis kochii Lutken.Ophiura sarsi was found to move with the aid of paddling movements of two pairs of arms: The fifth arm (passive) pointed backward. Ophiurans of this species do not use their ambulacral feet. Three main types of locomotor movements were distinguished inAmphipholis kochii, First, locomotion by the "breast stroke" method, in which one arm (the leading) points forward, two point to the side, and two backward. The two side arms are periodically carried forward, after which the disk and the remaining arms are moved with their aid. In this method waves of flexion and extension of the segments spread along the side arms. Second, displacement by pulling with the leading arm pushing with the hind arms, and third, movement by stepping movements of the ambulacral feet. These three methods of locomotion can be used either independently or in various combinations with each other. The ambulacral feet also provide the link between the active arms and the supporting surface by means of which the ophiurans can move forward.Institute of Oceanology, Academy of Sciencs of the USSR, Moscow. Institute of Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Moscow State University. Translated from Neirofiziologiya, Vol. 8, No. 5, pp. 521–528, September–October, 1976.     

A NEW SPECIES OF BRITTLESTAR (OPHIUROIDEA, ECHINODERMATA) FROM THE HUNSRÜCK SLATE (LOWER EMSIAN, LOWER DEVONIAN) OF GERMANY

Abstract:  Lapworthura lehmanni , a new species of ophiuroid, is described from four specimens from the Lower Devonian Hunsrück Slate of Germany. It is the only known ophiuroid in the Hunsrück Slate with paired but unfused ambulacrals, and it exhibits unique rows of spine-bearing dorsal arm ossicles. The genus Lapworthura Gregory was previously known only from the Ordovician of Scotland and the Silurian of England and Australia.     

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.     

New observations on some poorly known protasterid ophiuroids from the Lower Devonian Hunsrück Slate of Germany

Four genera and species of protasterid ophiuroids have been reported from the Lower Devonian Hunsrück Slate:Bundenbachia beneckei, Palaeophiomyxa grandis, Mastigophiura grandis, andPalaeophiura simplex. In addition, the speciesMiospondylus rhenanus has traditionally been recognized as a member of the Lapworthuridae, a group of ophiuroids with paired but unfused ambulacrals. This ambulacral arrangement would make it unique among Hunsrück Slate ophiuroids.Palaeophiura simplex andM. rhenanus were originally based on Single, poorly preserved specimens, and their morphology has remained poorly understood. A restudy of the type material offP. simplex andM. rhenanus and a detailed comparison to other Hunsrück ophiuroids warrants reassignment of these taxa toBundenbachia beneckei.Palaeophiura simplex is a juvenile example of this species andM. rhenanus is a peculiarly preserved adult. A newly prepared, well-preserved specimen ofBundenbachia beneckei expands our knowledge of the morphology of its mouthframe. In addition,Mastigophiura grandis is revised and redescribed based on newly recognized fossil material.     

The Asteroidea (Echinodermata) of the Muschelkalk (Middle Triassic of Germany)

Trichasteropsis Eck, from the Muschelkalk of Germany is the only Triassic asteroid known from more than fragmentary material. Most spécimens representT. weissmanni (Münster) whereasT. senfti ECK,T. bielertorum n. sp., andBerckhemeraster charistikos n. gen. et n. sp., are each known from few individuals.Parsimony analysis hère treats the füll Ordovician to Récent history of the Asteroidea using a somasteroid (a pre-asteroid stelleroid) outgroup. Ambulacral évolution is critical in echinoderm history; the ambulacral arrangement of crown-group asteroids first appears in Paleozoic sister groups, and the subclass Ambuloasteroidea n. subcl. is proposed for Paleozoic and younger taxa with critical ambulacral apomorphies. Muschelkalk asteroids are assigned to the family Trichasteropsiidae n. fam., superorder Forcipulatacea. The recently described Triassic genusNoriaster belongs to the extant family Poraniidae, superorder Valvatacea.Trichasteropsis andNoriaster represent separate major phylogenetic branches of the post-Paleozoic infraclass Neoasteroidea, and together they indicate that diversification of modern-type asteroids was under-way during the Triassic, although the Mesozoic marine révolution largely was a Jurassic and later event. Post-Paleozoic asteroids appear to hâve returned to Paleozoic life modes in spite of new morphological expressions. Trichasteropsis is skeletally robust, suggesting protection from wave impact or predators. It is found in sédiments associated with shell banks but not from within the banks.Trichasteropsis senfti commonly occurs with brachiopods whereasT. weissmanni does not, although brachiopods are found in associated strata. Aspects of morphology of both species are similar to those of récent predatory Asteriidae suggesting similar behavior, but feeding habits ofTrichasteropsis are unverified.     

Colony specificity and chemotaxis in the compound ascidian Botryllus schlosseri

We re-investigated the behavior of hemocytes during the non-fusion (rejection) reaction between genetically incompatible colonies of the ascidian Botryllus schlosseri. In the course of the reaction, hemocytes - mainly morula cells - crowd inside the blind ends of marginal vascular vessels (known as ampullae) of the colonial leading edge (LE) facing the foreign colony which suggests the occurrence of chemotactic attraction of circulating hemocytes towards the ampullar lumen. Then, cells migrate, through the ampullar tips, into the partially fused tunics and contribute to the formation of the necrotic spots along the contact borders which characterize the reaction. Studies on histological sections clearly indicate that, although morula cell concentration is always higher in ampullae of the LE than in those of the lateral (L) part of the colony, their frequency significantly increases in LE ampullae of rejecting colonies with respect to LE ampullae of both fusing and isolated colonies. In addition, in vitro chemotaxis experiments demonstrated that blood plasma from incompatible colonies can stimulate morula cell migration through polycarbonate filters and this passage is inhibited by antibodies raised against mammalian pro-inflammatory cytokines. The possible nature and role of molecules recognized by anti-cytokine antibodies in hemocyte migration are discussed.     

古生代有孔虫实体化石研究方法

古生代有孔虫大部分是通过切片进行研究的,但是普通的切片法很难得到非常好的定向标本,这就产生了很多同物异名,限制了有孔虫在古生代生物地层学、古生态学和古海洋学中的应用。另外由于口孔的内部和外部特征在分类学中的地位越来越重要,所以通过实体化石来研究古生代有孔虫理应受到更多的重视。本文论证了在古生代地层中获得实体化石的可能性,对化石提取的实验步骤进行了系统总结,细述了对实体化石进行扫描电镜和切片研究的全过程,并根据笔者的经验,对一些技术细节及注意事项进行了详细介绍。本文最后建议,古生代的有孔虫研究应该尽可能地利用实体化石来进行。     

Mechanism of perception of changes in the magnetic field by ampullae of lorenzini of elasmobranchs

The mechanisms of reception of changes in the magnetic field by electroreceptor formations were investigated in experiments on Black Sea rays in which spike activity was recorded from single nerve fibers connected with the ampullae of Lorenzini. The responses of the ampullae of Lorenzini to magnetic stimulation were shown to be due to induced electric currents creating potential gradients in the body tissues of the fish and the sea water. On the basis of differences in responses of different ampullae to magnetic stimulation, it is possible to distinguish between magnetic stimuli and other stimuli acting on the electroreceptor system. Potentiation of the receptor response to magnetic stimulation was found as the fish came closer to the "shore." The mechanisms and biological importance of reception of changes in the magnetic field by the ampullae of Lorenzini are discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 10, No. 1, 75–83, January–February, 1978.     

Allogeneic inhibition in a compound ascidian, Batryllus primigenus Oka. I. Processes and features of "Nonfusion" reaction

The processes and features of the “Nonfusion” reaction (NFR), a measure positive allogeneic inhibition, in a compound ascidian, Botryllus primigenus, has been studied.The process and the features of NFR have been observed when two incompatible colonies were placed as to make a contact with (1) ampullae to ampullae, (2) ampullae to margin without ampullae, and (3) cut surface to cut surface of the colonies. NFR appeared immediately after fusion of the test matrices of two incompatible colonies. The ampullae have the ability to loosen or dissolve the outer membrane of the test matrix and to penetrate into the other colony. The first response in NFR was the destruction of test cells around the contact area leading to an appearance of filaments around disintegrated test cells. The second reaction was contraction of the ampullae. Distal parts of the ampullae were thus cut off by constriction and necrosis. The contact area between two incompatible colonies was then separated from the healthy parts of both colonies, and NFR was completed.     

The magnetic diver differential microgasometer

The magnetic differential microgasometer described here is basically an equalarm torsion balance for measuring the reduced weight of submerged objects. The changes of buoyancy (ΔV_gx;ΔRW) are compensated for by the magnetic force acting on the same arm where the change has occurred (substitution method of weighing). Two ampullae, one with the sample and the other with the blank, are placed into the loops mounted at the ends of the arms. The gas volume and the geometry of the ampullae are practically equal. Therefore, the instrument is stable and fairly insensitive to changes of barometric pressure and temperature, and it can be used for gasometric measurements in an open flotation vessel. The applicability of the method for experiments where high blanks cannot be avoided was demonstrated by respiration measurements of multiplying cells and by measurements of cholinesterase activity of muscle end-plate samples.     

Membrane Dynamics of the Contractile Vacuole Complex of Paramecium1

ABSTRACT. Membrane dynamics of the contractile vacuole complex of Paramecium were investigated using conventional electron microscopy of cells so that the vacuoles were serial-sectioned longitudinally and transversely. During systole, vacuolar membrane collapses first into flattened cisternae which undergo further modification into a mass of interconnected small membrane tubules. These tubules retain their connections with the radiating microtubular ribbons; consequently they are found only in the poleward hemisphere. Permanent connections between ampullae and the collapsed vacuole membrane could not be verified nor was a sphincter-like mechanism for closing such a junction observed. Membranes of the ampullae and the collecting canals also collapse to varying extents into arrays of tubules that remain bound to microtubular ribbons during diastole. Thus vacuole, ampullae, and collecting canal membranes all assume tubular forms when internal volume is at a minimum. Having failed to observe a microfilamentous encasement of the vacuole, we suggest that an alternative mechanism for the “contractile” function should be sought. One such is based on fluid volume increase and fluid flow within transiently interconnected tubular membrane systems that cycle between a tubular and a planar membrane form as internal volume is periodically increased and reduced. The driving force for this mechanism might best be sought in the molecular structure of the membranes of the contractile vacuole complex.     

A new brittle star (Ophiuroidea: Ophiodermatina) from the Early Triassic Paris Biota (Bear Lake County,Idaho, USA)

Knowledge on the morphology of Early Triassic ophiuroids is very limited, in spite of the relatively high number of described species. This hampers attempts to use morphology-based phylogenetic analyses in order to explore the early diversification of the ophiuroid modern clades between the latest Paleozoic and the early Mesozoic. Here, a new ophiuroid from the Early Triassic of southern Idaho (USA) is described and assigned to the new taxon Shoshonura brayardi Thuy nov. gen., nov. sp. It is part of an exceptional Early Triassic faunal assemblage recently described as the Paris Biota. On account of the arm morphology and the shape of the arm spines and arm spine articulations, Shoshonura brayardi nov. gen., nov. sp. is considered a basal member of the suborder Ophiodermatina.     

Re-evaluation of the Devonian family Helianthasteridae Gregory, 1899 (Asteroidea: Echinodermata)

Because of their taxonomic and morphologic diversity, the asteroids of the Lower Devonian Hunsrück Slate of Germany are important to both an understanding of the history of the class Asteroidea and to the interpretation of community evolution during the Paleozoic. Helianthaster Roemer, 1863, a large multiarmed Hunsrück asteroid, is redescribed. The Helianthasteridae Gregory is restricted to Helianthaster and Arkonaster Kesling, 1982 (Middle Devonian, Canada); Lepidasterella Schuchert is similar to the other two genera but known specimens are of poor quality and as a result the status of the genus is uncertain. Helianthaster is noteworthy in part because its size, multiarmed state, and presence of pedicellariae suggest certain crown-group genera, yet aspects of the arrangement of the ambulacral column are characteristic of the Paleozoic asteroid evolutionary grade. Neither Helianthaster nor other Hunsrück asteroids appear closely linked to the crown group; instead, functional patterns apparently re-emerged through time, although identification of specific behavior of ancient asteroids is difficult to impossible.      

Die Temperaturempfindlichkeit isolierter Lorenzinischer Ampullen

The ampullae ofLorenzini of elasmobranchs have a high sensitivity to temperature, as has been found bySand (1938) andHensel (1955) in single fibre preparations. Rapid cooling results in a steep, although temporary increase in impulse frequency; rapid warming is followed by a short and temporary inhibition of discharges. This behaviour very closely resembles that of the “cold” receptors of warm blooded animals. During experiments with isolated ampullae ofLorenzini of the infra- and supraorbital group ofScyllium canicula, we found in two animals some ampullae with an opposite response to temperature changes: rapid cooling inhibited the discharges of the ampullae, while rapid changes to warmth initiated a temporary increase in impulse rate. In some of the ampullae a kind of “adaptation” after fast temperature changes was observed which was also different from what has been described by other authors. The high dynamic response after rapid cooling was followed by a strong diminution in discharge rate. Only after several minutes the impulse frequency again rose to a new and constant level. With rapid warming the behaviour of the spike rate was the mirror image: initially, the frequency dropped to low values but then rose again to a level which was higher than before warming. After this maximum the discharge role fell to a constant frequency. From the results it may be concluded that the localization of the ampullae might be of importance for the “specific” function of the receptor as well as for the differentiation of the peripheral stimulus patterns.