Bryozoans of the order Melicerititida: Morphological features and position of the order in the taxonomic structure of the class Stenolaemata

Morphological features of a distinctive group of Post-Paleozoic bryozoans belonging to the order Melicerititida (Stenolaemata) are discussed. They include funnel-shaped zooecia, facettes, semicircular zooecial apertures with a straight proximal edge, calcified opercula, and various types of vicarious eleozooecia, which resemble avicularia of cheilostome bryozoans. These morphological structures are unique within the class Stenolaemata. They not only differentiate these bryozoans both morphologically and evolutionarily from Cyclostomata, in which some authors place them, but also from all the other orders of this class.     

Morphogenesis in the individual and historical development of marine post-Paleozoic Tubuliporida (bryozoa,Stenolaemata)

The principal features of the morphogenesis in the individual and historical development of marine post-Paleozoic bryozoans of the order Tubuliporida (=Cyclostomata, part.) are discussed. Throughout their history (Ordovician–Recent), Tubuliporida retained the morphological type of the tubu-lar zooid with a terminal aperture. This may suggest similarities not only between the zooidal anatomy of fossil and living Tubuliporida but also between the ontogenetic processes in their ancestrulae and zooids and similarities in astogeny in general. After the two types of reproduction, sexual and vegetative, which lead to the formation of new colonies that could only grow by budding, had been developed, the develop-ment of different types of colonial organization was of special importance in the evolution of the post-Pale-ozoic Tubuliporida.     

Evolution of the colonial growth habit in the ordovician bryozoans of the class stenolaemata: Feeding adaptations (Leningrad Region,Russia)

Based on the study of the growth habits and the relief of the colony surface in bryozoans of the class Stenolaemata from the Lower (Latorp horizon) and Middle (Volchov and Kunda horizons) Ordovician of the Leningrad Region, these bryozoans are shown to develop from the simple, unilaminate colonies (B_I^β) to the massive colonies with a nodular surface and smooth columnar colonies (B_II^α), which subsequently evolved into the columnar-spiral (B_II^β) and more complex erect branching and fenestrate constructions (B_II^γ), and subsequently into the branching, articulate colonies (B_III^α). The apertures of autozooecia and the character of their arrangement on the colony surface changed correlatively from the circular (B_I^β) to polygonal and roundedpolygonal, randomly arranged apertures, and subsequently to the oval apertures (B_III^α) arranged in strictly regular longitudinal or longitudinal-diagonal rows or in a quincuncial pattern. Thus, the development of growth habits in the bryozoans under consideration has a progressive character. It is expressed in the progressive increase in the complexity of growth habits of colonies and in the more regular arrangement of apertures and other structures on the colony surface. The directionality of morphological changes in the growth habits of colonies of Ordovician bryozoans was apparently closely associated with the development of more complex environmental interactions of these bryozoans, especially with water currents supplying food particles. It is suggested that the high competitive ability of bryozoans of the class Stenolaemata at early stages of its development in the basin of Baltoscandia was apparently due to the better use of food resources.     

Rare bryozoans (Stenolaemata) with bilateral colonies from the Jurassic and Cretaceous of the East European Platform

New bryozoans characterized by a bilateral vertical colonies colonies—Cardioecia refuga sp. nov. (Tubuliporida), Elea lyapini sp. nov., E. troshkovensis sp. nov., and E. taylori sp. nov. (Melicerititida) from Middle Jurassic (Middle Callovian) of the Moscow region (Russia), and Biforicula legitima sp. nov. (Melicerititida) from the Upper Cretaceous (Lower Campanian) of the Southern Donets Basin (Ukraine)—are described. All species described in this paper belong to genera that have been recorded for the first time in the East European Platform. Some morphological structures in the colonies of these bryozoans and some distinctive features of the order Melicerititida that emphasize the difference of this order from bryozoans of other orders of the class Stenolaemata are examined.     

Modern Data on the Innervation of the Lophophore in Lingula anatina (Brachiopoda) Support the Monophyly of the Lophophorates

Evolutionary relationships among members of the Lophophorata remain unclear. Traditionally, the Lophophorata included three phyla: Brachiopoda, Bryozoa or Ectoprocta, and Phoronida. All species in these phyla have a lophophore, which is regarded as a homologous structure of the lophophorates. Because the organization of the nervous system has been traditionally used to establish relationships among groups of animals, information on the organization of the nervous system in the lophophore of phoronids, brachiopods, and bryozoans may help clarify relationships among the lophophorates. In the current study, the innervation of the lophophore of the inarticulate brachiopod Lingula anatina is investigated by modern methods. The lophophore of L. anatina contains three brachial nerves: the main, accessory, and lower brachial nerves. The main brachial nerve is located at the base of the dorsal side of the brachial fold and gives rise to the cross neurite bundles, which pass through the connective tissue and connect the main and accessory brachial nerves. Nerves emanating from the accessory brachial nerve account for most of the tentacle innervation and comprise the frontal, latero-frontal, and latero-abfrontal neurite bundles. The lower brachial nerve gives rise to the abfrontal neurite bundles of the outer tentacles. Comparative analysis revealed the presence of many similar features in the organization of the lophophore nervous system in phoronids, brachiopods, and bryozoans. The main brachial nerve of L. anatina is similar to the dorsal ganglion of phoronids and the cerebral ganglion of bryozoans. The accessory brachial nerve of L. anatina is similar to the minor nerve ring of phoronids and the circumoral nerve ring of bryozoans. All lophophorates have intertentacular neurite bundles, which innervate adjacent tentacles. The presence of similar nerve elements in the lophophore of phoronids, brachiopods, and bryozoans supports the homology of the lophophore and the monophyly of the lophophorates.     

Tentacle structure in freshwater bryozoans

Tentacles are the main food‐gathering organs of bryozoans. The most common design is a hollow tube of extracellular matrix (ECM), covered with ten columns of epithelial cells on the outside, and a coelothelium on the inside. Nerves follow the ECM, going between the bases of some epidermal cells. The tentacle musculature includes two bundles formed by myoepithelial cells of the coelothelium. The tentacles of freshwater (phylactolaemate) bryozoans, however, differ somewhat in structure from those of marine bryozoans. Here, we describe the tentacles of three species of phylactolaemates, comparing them to gymnolaemates and stenolaemates. Phylactolaemate tentacles tend to be longer, and with more voluminous coeloms. The composition of the frontal cell row and the number of frontal nerves is variable in freshwater bryozoans, but constant in marine groups. Abfrontal cells form a continuous row in Phylactolaemata, but occur intermittently in other two classes. Phylactolaemata lack the microvillar cuticle reported in Gymnolaemata. Abfrontal sensory tufts are always composed of pairs of mono‐ and/or biciliated cells. This arrangement differs from individual abfrontal ciliary cells of other bryozoans: monociliated in Stenolaemata and monociliated and multiciliated ones in Gymnolaemata. In all three groups, however, ciliated abfrontal cells probably serve as mechanoreceptors. We confirm previously described phylactolemate traits: an unusual arrangement of two‐layered coelothelium lining the lateral sides of the tentacle and oral slits in the intertentacular membrane. As previously reported, tentacle movements involved in feeding differ between bryozoan groups, with phylactolaemates tending to have slower movements than both gymnolaemates and stenolaemates, and a narrower behavioral repertoire than gymnolaemates. The morphological and ultrastructural differences between the freshwater species we studied and marine bryozoans may be related to these functional differences. Muscle organization, tentacle and coelom size, and degree of confluence between tentacle and lophophore coeloms probably account for much of the observed behavioral variability.     

New data on the colonial morphology of the Jurassic bryozoans of the class Stenolaemata

The radiation of the Jurassic bryozoans of the class Stenolaemata, which started in seas of the Bajocian and Bathonian of western Europe, is shown to be continued in basins of eastern Europe during the extensive Middle Callovian transgression. The taxonomic composition of stenolaemate bryozoans from the Jurassic of central European Russia and main features of their colonial morphology are discussed.     

Zooid orientation structures and water flow patterns in Paleozoic bryozoan colonies

Anstey, Robert L. 1981 12 15: Zooid orientation structures and water flow patterns in Paleozoic bryotoan colonies. Lethaia . vol. 14, pp. 287–302. Oslo. ISSN 0024–1164.
By means of direct physical evidence provided by zooecial orientation structures, active water flow systems in Paleozoic bryozoans are inferred to be variously centripetal, centrifugal, or basipetal. Monticules, previously assessed as excurrent water outlets, fall into three additional functional types: incurrent, bypassed, and funnel. In one species circular zoarial fenestrations served as excurrent water outlets. Water flow patterns are strongly correlated with zoarial growth form, which vanes in a general way with inferred habitat conditions in ancient environments. Monticular astogeny and phylogeny include a graded series of sizes, types, and functions. Analogy with zooidal polarities in extant stenolaemates suggests that colony bases and centripetal monticules in the Paleozoic orders were anally budded, but that erect branches and centrifugal monticules were orally budded, a character shared only by the freshwater Phylactolaemata. * Bryozoa, Stenolaemata, functional morphology, monticule function, hydrodynamics, feeding currents, Palaeozoic .     

奥陶纪苔藓动物的多样性演变——兼论苔藓动物的起源

苔藓动物是一类多为海生、滤食性的群体生物。奥陶纪是苔藓动物发生、演化辐射和灭绝的重要时期,也是苔虫礁形成的最早时期。已知最老的化石苔藓动物发现于中国特马豆克晚期。构成苔藓动物基本分类框架的狭唇纲(包括变口目、隐口目、泡孔目和管孔目)和宽唇纲(包括窗孔目和栉口目)也都是在奥陶纪时期逐步形成的,其中,变口目出现于特马豆克期Tr2时间段,在弗洛期和大坪期,多样性较低,但从达瑞威尔期开始,经桑比期至凯迪期,多样性不断增高,并出现辐射。隐口目(特别是"双叶类隐口目苔虫")也经历了与变口目相类似的发展过程,但它首次出现的时间要相对略迟于变口目。这两个目在整个奥陶纪苔藓动物群中一直占据主导地位。泡孔目、管孔目和窗孔目,先后首次出现在弗洛期Fl2时间段、大坪期Dp1和Dp2时间段,但它们在整个奥陶纪期间一直处于低多样性态势。至于栉口目,它首次出现的时间可能更迟,在凯迪期Ka4时间段,犹如昙花一现。苔藓动物的演化在接近奥陶纪末时呈两幕式灭绝,一次发生在凯迪期Ka2时间段(可能相当于塔凯和安斯蒂的"拉夫塞伊灭绝"),另一次发生在赫南特期Hi2时间段(可能相当于塔凯和安斯蒂的"赫南特灭绝")。分子生物学和形态学证据表明,苔藓动物属原口动物,而不是以前长期认为的后口动物,或介于原口动物和后口动物之间的过渡类型;而且,苔藓动物与腕足动物、帚形动物之间没有直接的亲缘关系。苔藓动物可能起源于一种叫原内肛动物的生物,它们的目一级分类单元之间的系统发育关系目前尚未形成共识,本文绘制的谱系图还有待于化石记录的不断补充和分子生物学研究的逐步介入以使其日趋完善。     

Particle capture in ciliary filter‐feeding gymnolaemate and phylactolaemate bryozoans – a comparative study

Riisgård, H.U., Okamura, B. and Funch, P. 2009. Particle capture in ciliary filter‐feeding gymnolaemate and phylactolaemate bryozoans – a comparative study. —Acta Zoologica (Stockholm) 91 : 416–425. We studied particle capture using video‐microscopy in two gymnolaemates, the marine cheilostome Electra pilosa and the freshwater ctenostome Paludicella articulata, and three phylactolaemates, Fredericella sultana with a circular funnel‐shaped lophophore, and Cristatella mucedo and Lophophus crystallinus, both with a horseshoe‐shaped lophophore. The video‐microscope observations along with studies of lophophore morphology and ultrastructure indicated that phylactolaemate and gymnolaemate bryozoans with a diversity of lophophore shapes rely on the same basic structures and mechanisms for particle capture. Our study also demonstrates that essential features of the particle capture process resemble one another in bryozoans, brachiopods and phoronids.     

New Stenolaematous bryozoans from the Jurassic of central European Russia (Moscow city and the Moscow and Kostroma Regions)

New bryozoans from the Middle Callovian (Middle Jurassic) of Moscow city (Reptomulticava pileola sp. nov.) and the Moscow Region (Spirodella radiolobata gen. et sp. nov.) and from the Middle Oxfordian (Upper Jurassic) of the Kostroma Region (Hyporosopora mittai sp. nov.) are described. All three bryozoans belong to the class Stenolaemata. Some features of their colonial organization and environmental conditions are discussed.     

Bryozoan (Stenolaemata) records from the upper Callovian (Middle Jurassic) of the Moscow region

Encrusting bryozoans (Stenolaemata, Tubuliporida) discovered from upper Callovian deposits (Middle Jurassic) near the town of Kolomna in the Moscow region are described. They belong to two new species: Microeciella kolomnensis sp. nov. and Diplosolen akatjevense sp. nov. These bryozoans encrust a fragment of the large shell of cephalopod mollusk (ammonite), which is an unusual substrate not only for bryozoans but also for the other encrusting organisms.     

Key novelties in the evolution of the aquatic colonial phylum Bryozoa: evidence from soft body morphology

Molecular techniques are currently the leading tools for reconstructing phylogenetic relationships, but our understanding of ancestral, plesiomorphic and apomorphic characters requires the study of the morphology of extant forms for testing these phylogenies and for reconstructing character evolution. This review highlights the potential of soft body morphology for inferring the evolution and phylogeny of the lophotrochozoan phylum Bryozoa. This colonial taxon comprises aquatic coelomate filter‐feeders that dominate many benthic communities, both marine and freshwater. Despite having a similar bauplan, bryozoans are morphologically highly diverse and are represented by three major taxa: Phylactolaemata, Stenolaemata and Gymnolaemata. Recent molecular studies resulted in a comprehensive phylogenetic tree with the Phylactolaemata sister to the remaining two taxa, and Stenolaemata (Cyclostomata) sister to Gymnolaemata. We plotted data of soft tissue morphology onto this phylogeny in order to gain further insights into the origin of morphological novelties and character evolution in the phylum. All three larger clades have morphological apomorphies assignable to the latest molecular phylogeny. Stenolaemata (Cyclostomata) and Gymnolaemata were united as monophyletic Myolaemata because of the apomorphic myoepithelial and triradiate pharynx. One of the main evolutionary changes in bryozoans is a change from a body wall with two well‐developed muscular layers and numerous retractor muscles in Phylactolaemata to a body wall with few specialized muscles and few retractors in the remaining bryozoans. Such a shift probably pre‐dated a body wall calcification that evolved independently at least twice in Bryozoa and resulted in the evolution of various hydrostatic mechanisms for polypide protrusion. In Cyclostomata, body wall calcification was accompanied by a unique detachment of the peritoneum from the epidermis to form the hydrostatic membraneous sac. The digestive tract of the Myolaemata differs from the phylactolaemate condition by a distinct ciliated pylorus not present in phylactolaemates. All bryozoans have a mesodermal funiculus, which is duplicated in Gymnolaemata. A colonial system of integration (CSI) of additional, sometimes branching, funicular cords connecting neighbouring zooids via pores with pore‐cell complexes evolved at least twice in Gymnolaemata. The nervous system in all bryozoans is subepithelial and concentrated at the lophophoral base and the tentacles. Tentacular nerves emerge intertentacularly in Phylactolaemata whereas they partially emanate directly from the cerebral ganglion or the circum‐oral nerve ring in myolaemates. Overall, morphological evidence shows that ancestral forms were small, colonial coelomates with a muscular body wall and a U‐shaped gut with ciliary tentacle crown, and were capable of asexual budding. Coloniality resulted in many novelties including the origin of zooidal polymorphism, an apomorphic landmark trait of the Myolaemata.     

Bryozoans are returning home: recolonization of freshwater ecosystems inferred from phylogenetic relationships

Bryozoans are aquatic invertebrates that inhabit all types of aquatic ecosystems. They are small animals that form large colonies by asexual budding. Colonies can reach the size of several tens of centimeters, while individual units within a colony are the size of a few millimeters. Each individual within a colony works as a separate zooid and is genetically identical to each other individual within the same colony. Most freshwater species of bryozoans belong to the Phylactolaemata class, while several species that tolerate brackish water belong to the Gymnolaemata class. Tissue samples for this study were collected in the rivers of Adriatic and Danube basin and in the wetland areas in the continental part of Croatia (Europe). Freshwater and brackish taxons of bryozoans were genetically analyzed for the purpose of creating phylogenetic relationships between freshwater and brackish taxons of the Phylactolaemata and Gymnolaemata classes and determining the role of brackish species in colonizing freshwater and marine ecosystems. Phylogenetic relationships inferred on the genes for 18S rRNA, 28S rRNA, COI, and ITS2 region confirmed Phylactolaemata bryozoans as radix bryozoan group. Phylogenetic analysis proved Phylactolaemata bryozoan's close relations with taxons from Phoronida phylum as well as the separation of the Lophopodidae family from other families within the Plumatellida genus. Comparative analysis of existing knowledge about the phylogeny of bryozoans and the expansion of known evolutionary hypotheses is proposed with the model of settlement of marine and freshwater ecosystems by the bryozoans group during their evolutionary past. In this case study, brackish bryozoan taxons represent a link for this ecological phylogenetic hypothesis. Comparison of brackish bryozoan species Lophopus crystallinus and Conopeum seurati confirmed a dual colonization of freshwater ecosystems throughout evolution of this group of animals.     

New bryozoans (Stenolaemata) from the Middle Jurassic of Moscow City and the Moscow Region

Five new species of stenolaematous bryozoans are described from the Middle Callovian of Moscow City (Krasnopresnenskii Region) and the Moscow Region (Gzhel’ Railroad Station and the quarry between the small towns of Rechitsy and Troshkovo): Entalophora alexeevi sp. nov., Cellulipora retshitsiensis sp. nov., Diplosolen pravus sp. nov., Siphodictyum primarium sp. nov., and Ceata kamushkiensis sp. nov. The last four species belong to the genera that have never been recorded from deposits older than the Lower Cretaceous. The presence of different types of heterozooecia in these bryozoans shows that the functional polymorphism of zooids developed considerably in the class Stenolaemata as early as the Middle Jurassic.     

Organogenesis during budding and lophophoral morphology of Hislopia malayensis Annandale, 1916 (Bryozoa, Ctenostomata)

Background  

Bryozoans represent a large lophotrochozoan phylum with controversially discussed phylogenetic position and in group relationships. Developmental processes during the budding of bryozoans are in need for revision. Just recently a study on a phylactolaemate bryozoan gave a comprehensive basis for further comparisons among bryozoans. The aim of this study is to gain more insight into developmental patterns during polypide formation in the budding process of bryozoans. Particular focus is laid upon the lophophore, also its condition in adults. For this purpose we studied organogenesis during budding and lophophoral morphology of the ctenostome bryozoan Hislopia malayensis.     

A model of particle capture by bryozoans in turbulent flow: significance of colony form

A hydrodynamic model was developed to examine particle capture by lophophores of encrusting bryozoans. Particle capture rate is predicted to increase with increasing speed of the feeding current. There should be a large feeding advantage when lophophores are tightly packed and excurrents are vented through chimneys. This prediction contradicts conclusions of an earlier model study and suggests that selection for colony integration has a basis in the acquisition of food. If lophophores are not tightly packed, particle-capture patterns depend on two key ratios: the advection ratio (feeding current velocity to shear velocity) and the separation ratio (lophophore spacing to lophophore diameter). At high separation ratios, particle capture rates should be fairly uniform among zooids. At high advection ratios, lophophores located near the upstream colony edge should experience higher rates of particle capture. Rates of particle capture in turbulent flows should greatly exceed those in laminar flows (of identical speed) when excurrent waters are locally remixed into the flow above lophophores. However, when lophophores are tightly packed and excurrents are vented through chimneys, feeding rates should be identical in turbulent and laminar flows. Thus, colonies that vent excurrents through chimneys may be uniquely able to exploit weak laminar flows.     

Body cavities in bryozoans: Functional and phylogenetic implications

Based on morphological evidence, Bryozoa together with Phoronida and Brachiopoda are traditionally combined in the group Lophophorata, although this view has been recently challenged by molecular studies. The core of the concept lies in the presence of the lophophore as well as the nature and arrangement of the body cavities. Bryozoa are the least known in this respect. Here, we focused on the fine structure of the body cavity in 12 bryozoan species: 6 gymnolaemates, 3 stenolaemates and 3 phylactolaemates. In gymnolaemates, the complete epithelial lining of the body cavity is restricted to the lophophore, gut walls, and tentacle sheath. By contrast, the cystid walls are composed only of the ectocyst-producing epidermis without a coelothelium, or an underlying extracellular matrix; only the storage cells and cells of the funicular system contact the epidermis. The nature of the main body cavity in gymnolaemates is unique and may be considered as a secondarily modified coelom. In cyclostomes, both the lophophoral and endosaccal cavities are completely lined with coelothelium, while the exosaccal cavity only has the epidermis along the cystid wall. In gymnolaemates, the lophophore and trunk cavities are divided by an incomplete septum and communicate through two pores. In cyclostomes, the septum has a similar location, but no openings. In Phylactolaemata, the body cavity is undivided: the lophophore and trunk coeloms merge at the bases of the lophophore arms, the epistome cavity joins the trunk, and the forked canal opens into the arm coelom. The coelomic lining of the body is complete except for the epistome, lophophoral arms, and the basal portions of the tentacles, where the cells do not interlock perfectly (this design probably facilitates the ammonia excretion). The observed partitioning of the body cavity in bryozoans differs from that in phoronids and brachiopods, and contradicts the Lophophorata concept.     

Ciliary feeding structures and particle capture mechanism in the freshwater bryozoan Plumatella repens (Phylactolaemata)

Abstract. In contrast to marine bryozoans, the lophophore structure and the ciliary filter‐feeding mechanism in freshwater bryozoans have so far been only poorly described. Specimens of the phylactolaemate bryozoan Plumatella repens were studied to clarify the tentacular ciliary structures and the particle capture mechanism. Scanning electron microscopy revealed that the tentacles of the lophophore have a frontal band of densely packed cilia, and on each side a zigzag row of laterofrontal cilia and a band of lateral cilia. Phalloidin‐linked fluorescent dye showed no sign of muscular tissue within the tentacles. Video microscopy was used to describe basic characteristics of particle capture. Suspended particles in the incoming water flow, set up by the lateral ‘pump’ cilia on the tentacles, approach the tentacles with a velocity of 1–2 mm s^‐1. Near the tentacles, the particles are stopped by the stiff sensory laterofrontal cilia acting as a mechanical sieve, as previously seen in marine bryozoans. The particle capture mechanism suggested is based on the assumed ability of the sensory stiff laterofrontal cilia to be triggered by the deflection caused by the drag force of the through‐flowing water on a captured food particle. Thus, when a particle is stopped by the laterofrontal cilia, the otherwise stiff cilia are presumably triggered to make an inward flick which brings the restrained particle back into the downward directed main current, possibly to be captured again further down in the lophophore before being carried to the mouth via the food groove. No tentacle flicks and no transport of captured particles on the frontal side of the tentacles were observed. The velocity of the metachronal wave of the water‐pumping lateral cilia was measured to be ～0.2 mm s^‐1, the wavelength was ～7 μm, and hence the ciliary beat frequency estimated to be ～30 Hz (～20 °C). The filter feeding process in P. repens reported here resembles the ciliary sieving process described for marine bryozoans in recent years, although no tentacle flicks were observed in P. repens. The phylogenetic position of the phylactolaemates is discussed in the light of these findings.     

Intracolonial variability in new bryozoans from the Middle Jurassic of the Bryansk Region (Russia)

Two new species of encrusting bryozoans (Stenolaemata, Tubuliporida) from the Middle Callovian (Middle Jurassic) deposits of the Bryansk Region, Oncousoecia guzhovi sp. nov. and Microeciella fokinensis sp. nov., are described. Their intracolonial variability, which is due to differences in astogenetic stages and to external factors, is discussed.