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Zooids and extrazooidal skeleton in the order Trepostomata (Bryozoa)
Abstract
Reconsideration of the nature of zooids in trepostomate Bryozoa defines them as physically connected and asexually replicated colony members that housed systems of organs necessary to perform vital functions for the colonies. Zooids known to contain organs in trepostomes are limited to autozooids, the requisite feeding and sexual units, and polymorphs, including macrozooids and two rare zooids of unknown function. Other colony structures are extrazooidal and remain outside zooidal boundaries throughout colony
life. They include the commonly occurring mesopores, exilapores, and styles. This two-part morphologic division of colonies reveals two correlated functions. The essential autozooids dominated the growth patterns and physiology of trepostome colonies; the extrazooidal
parts grew concurrently and passively to connect autozooids and to support and strengthen colonies.
... The skeleton in Palaeostomates is usually laminated, normally in the exozone, or rather hyaline, amorphic in the endozone or in the cores of styles (Tavener-Smith 1969a,b, Armstrong 1970, Blake 1973. In their appearance, the laminated internal walls may be merged, without visible zooecial boundary, or serrated, with distinct boundary between adjacent zooecia (Boardman & Buttler 2005). Fenestrate and cryptostome bryozoans had extensive sheets of external laminated skeletal material, whereas the majority of cystoporates developed a special vesicular skeleton, which filled the space between autozooecia (Utgaard 1983). ...
... Fenestrate and cryptostome bryozoans had extensive sheets of external laminated skeletal material, whereas the majority of cystoporates developed a special vesicular skeleton, which filled the space between autozooecia (Utgaard 1983). Such a vesicular skeleton is in less extent also present in other groups such as in trepostomes (Boardman & Buttler 2005), in ptilodictyines (Karklins 1983, Gorjunova & Lavrentjeva 1993, and in fenestrates (Morozova 2001). A vesicular skeleton is not known from rhabdomesines (Blake 1983b, Gorjunova 1985. ...
... Different types of styles are known in various groups of Palaeostomata. Their function is mainly regarded as protective or structural (Tavener-Smith 1969a, Armstrong 1970, Blake 1973, 1983b, Boardman & Cheetham 1973, Boardman & Buttler 2005. It is supposed that the function of acanthostyles might be to raise exterior membranous walls above zooecial apertures and skeletal surfaces in order to improve communication between zooids (Boardman 1983). ...
... This suggests that the cystid exerts control over polypide formation in the Gymnolaemata (the zooecium size hypothesis; Harvell, 1994), while the polypide (and its possible absence) exerts control on cystid formation in other bryozoan taxa. The zooecium size hypothesis also operates in the other direction: abnormally large polypides (conjoined or double polypides) have been found only in enlarged cystids (Jebram, 1978;Harvell, 1994), and 'macrozooids' in Trepostomata may have contained larger polypides (Boardman & Buttler, 2005). ...
... Modified cystid + polypide Cystid and polypide are modified, and zooids may have protrusible but non-feeding polypides Extrazooidal structure Structures external to zooidal boundaries at all stages of development (Boardman & Buttler, 2005). Can be solid skeleton or space-enclosing skeletal structures (McKinney & Jackson, 1989). ...
... some Disporella species (Boardman, 1998)]. Extrazooidal spine-like 'stylets' may also be present in stenolaemates, which unlike spines, possess a solid core enclosed by laminating sheaths (Blake, 1983;Boardman & Buttler, 2005). A few rare forms arise from zooidal boundaries [zooecial styles, exozonal styles (Boardman & Buttler, 2005)], and should be considered autozooidal appendages instead of extrazooidal structures. ...
Modularity is a fundamental concept in biology. Most taxa within the colonial invertebrate phylum Bryozoa have achieved division of labour through the development of specialized modules (polymorphs), and this group is perhaps the most outstanding exemplar of the phenomenon. We provide a comprehensive description of the diversity, morphology and function of these polymorphs and the significance of modularity to the evolutionary success of the phylum, which has >21000 described fossil and living species. Modular diversity likely arose from heterogeneous microenvironmental conditions, and cormidia (repeated clusters of associated modules) are an emergent property of the cue thresholds governing zooid plasticity. Polymorphs in a colony have, during phylogeny, transitioned into associated non‐zooidal structures (appendages), increasing colonial integration. While the level of module compartmentalization is important for the evolution of bryozoan polymorphism, it may be less influential for other colonial invertebrates.
... Bryozoans described in this paper reveal various types of structures that can be summarized as "stylets" (Blake 1983a: 537). These structures have a generally rod-like or spine-like appearance and have been discussed in many earlier publications (e.g., Tavener-Smith 1969a; Tavener-Smith 1969b; Armstrong 1970; Blake 1873a; Blake 1873b; Boardman & Buttler 2005). The following types of stylets are observed in the Pennsylvanian bryozoans from the Sandia Mountains: acanthostyles, aktinotostyles, microstyles, nodes, and mural spines. ...
... Bryozoans described in this paper reveal various types of structures that can be summarized as "stylets" (Blake 1983a: 537). These structures have a generally rod-like or spine-like appearance and have been discussed in many earlier publications (e.g., Tavener-Smith 1969a; Tavener-Smith 1969b; Armstrong 1970; Blake 1873a; Blake 1873b; Boardman & Buttler 2005). The following types of stylets are observed in the Pennsylvanian bryozoans from the Sandia Mountains: acanthostyles, aktinotostyles, microstyles, nodes, and mural spines. ...
The stenolaemate bryozoan fauna is described from the Pennsylvanian (Atokan-Virgilian) of the Sandia Mountains, New Mexico, USA. The fauna includes 27 species: eight cystoporates, four trepostomes, three cryptostomes, and 12 fenestrates. One species is new: the trepostome Mi shulgella vachardi sp. nov. Eight species are described in open nomenclature: three cystoporates: Fistulipora sp., Fistulamina sp., Cystodictya sp., one trepostome: Trepostomata sp. indet., and four fenestrates: Fabifenestella aff. praevirgosa (Schulga-Nesterenko, 1951), Spinofenestella sp., Laxi fenestella sp., and Penniretepora aff. bellula (Ulrich, 1890). Of the studied units, the Sandia Formation (Atokan-Desmoinesian) contains the most abundant bryozoan association, including 15 species. The Gray Mesa Formation (Desmoinesian) contains 14 species, whereas the Story Member of the Atrasado Formation (Virgilian) contains 12 bryozoan species. The studied samples from the Tinajas Member of the Atrasado Formation (Missourian) contain only two species. Palaeobiogeographic relationships of the Pennsylvanian bryozoan fauna from the Sandia Mountains are mostly restricted to the American continent, with a few connections to the Pennsylvanian of Europe. The bryozoan fauna indicates low-energy to high-energy depositional environments.
... It may be considered that the BSI would be more accurate if it was based on three-dimensional characters such as the volume of space occupied by autozooecial, mesozooecial or exilazooecial chambers, as well as the volume of exozonal and endozonal walls, the portions of acanthostyles that extend beyond the surficial margins of autozooecial walls, and any intrazooecial features such as widely spaced monilae in the exozone, skeletal diaphragms, hemiphragms, and cystiphragms (Boardman 2001;Boardman and Buttler 2005). The effect of these features on the BSI values could be computed by adding those additional characters composed of solid skeleton such as acanthostyles to the left-hand side of the equation alongside EW and ZWT and those of the open spaced features (exilazooecia and mesozooecia) to the right-hand side in combination with MZD. ...
... In several Paleozoic bryozoans such as the encrusting trepostomate Batostoma (Ross 1963;Boardman and Buttler 2005) spaces between zooids were taken up either by expansive mesopores that are horizontally strengthened by diaphragms (Fig. 9B) or by extrazooidal vesicular skeleton between autozooecia, as seen in cystoporate bryozoans (Fig. 6C, 6E). These structures, however, increase the distance between autozooecial apertures and cause an increase in lophophore size. ...
Bryozoans, stromatoporoid sponges, and tabulate corals, all colonial metazoans with lamellar, encrusting growth forms, developed and simultaneously diversified during the Great Ordovician Biodiversification Event (GOBE). After revisiting some classic Lower, Middle, and Upper Ordovician reef localities in Laurentia (Franklin Mountains, west Texas, Mingan Islands in eastern Canada, and Champlain Valley in northeastern United States) and Baltica (northern Estonia) and reviewing the literature, we demonstrate that during the Ordovician a newly emerging consortium of sheet-like bryozoans, stromatoporoid sponges, and tabulate corals locally bound together by microbes, automicrite, and cement and solidly rooted in sediment became the dominant reef-builders globally. The diversification of these sheet-like metazoans (SLM), however, clearly lagged behind the first appearance of their respective skeletal ancestors. Their habitat expansion can be exemplified as a case of simultaneous ecological fitting and ecosystem engineering when the independently evolved shared traits were simultaneously co-opted and became advantageous under globally different environmental conditions. This interaction led to the evolutionary diversification of colonial metazoans during the GOBE and to the expansion of novel reef habitats in previously soft-surface settings; a transformation that forever changed marine reefal ecosystems.
... Distinctly convex lamellar wall microstructure, zooecial boundaries amalgamated. Heterozooecia (mesopores in Boardman and Buttler 2005) usually abundant, originating in the inner exozone, containing rare to abundant basal diaphragms. Distinct difference in diameter between autozooecia and smaller heterozooecia. ...
The type series of Arcticopora christiei Fritz, 1961 – type species of Arcticopora – has been restudied and compared with recently collected material from time-equivalent beds of Ellesmere Island, Canadian Arctic. Taxonomically important characters previously not discussed include diaphragms in both endozone and exozone, irregularly shaped zooecial apertures and growth of acanthostyles. The bryozoan beds containing A. christiei are confirmed as Late Dienerian – Early Smithian (Early Triassic) based on new conodont data. Most Triassic species of Paralioclema and Pseudobatostomella are transferred to Arcticopora. A new family Arc-ticoporidae fam. nov. is erected, including Arcticopora Fritz, 1961; Zozariella Schäfer and Fois, 1987; Vysokella Zágoršek, 1993; and Dyscritellopsis Schäfer and Grant-Mackie, 1994.
... In S. lineata and Leptotrypa minima, macrozooecia are associated with monticules where they are located close to the crests; these zooecia have a larger diameter than autozooecia and have been shown by Boardman and Buttler (2005) to have contained polypides. They contend that they may have been the locus of incurrents on monticules as suggested by Anstey (1981), but these findings are not backed up by studies of monticule dynamics elsewhere. ...
Spatiopora Ulrich, 1882 is a trepostome bryozoan that is found encrusting living orthoconic nautiloids in the Upper Ordovician (Katian) of North America, as do several other bryozoans. These epizoozoan bryozoans are characterized by possessing thin unilaminate zoaria with rows of elongate maculae, which may be monticulate and aligned coaxially to the host growth axis. These develop a distinctive linear shape in response to growing on a conical host, rather than as a response to channelized water flow along the host. Monticules increase in size and spacing adorally until a maximum inter-macular area is reached that results in a decline in surface water flow efficiency, and a new monticular line is inserted. Orthocones normally swam forward at lower velocities that enabled lophophore eversion and feeding, which would have been impossible at the higher speeds reached when the host jetted backwards during escape. Monticules reduced drag and turbulence acting on the orthocones which allowed for more efficient venting of bryozoan macular excurrents. Characteristic elliptical monticule growth continued even after death of the motile host. A Trypanites-bryozoan-orthoconic nautiloid association shows a complex biological and taphonomic relationship between these organisms.
... 159e). In keeping with the terminology introduced by Boardman and Buttler (2005), the term exilapores is used here instead of exilazooecia as previously done by Key et al. (2001 Key et al. ( , 2002). The intermacular autozooecial chamber cross-sectional areas in this colony are on average ten times larger than the macular exilapore chamber crosssectional areas (Key et al. 2001). ...
Colony-wide feeding currents are a common feature of many bryozoan colonies. These feeding currents are centered on excurrent macular chimneys that expel previously filtered water away from the colony surface. In some bryozoans these macular chimneys consist of a branching channel network that converges at a point in the center of the chimney. The bifurcating channels of the maculae are analogous to a stream channel network in a closed basin with centripetal drainage. The classical methods of stream channel network analysis from geomorphology are here used to quantitatively analyze the number and length of macular channels in bryozoans. This approach is applied to a giant branch of the trepostome bryozoan Tabulipora from the Early Permian Kim Fjelde Formation in North Greenland. Its large size allowed 18 serial tangential peels to be made through the 8-mm-thick exozone. The peels intersected two stellate maculae as defined by contiguous exilapores. The lengths of 1460 channels radiating from the maculae were measured and their Horton-Strahler stream order and Shreve magnitude scored. We hypothesize that if fossil bryozoan maculae function as excurrent water chimneys, then they should conform to Hortons laws of stream networks and behave like closed basins with centripetal drainage. Results indicate that the stellate maculae in this bryozoan behaved liked stream channel networks exhibiting landscape maturation and stream capture. They conformed to the Law of Stream Number. They have a Bifurcation Ratio that falls within the range of natural stream channel networks. They showed a pattern opposite that expected by the Law of Stream Lengths in response to behavior characteristic of a centripetal drainage pattern in a closed basin. Thus, the stellate maculae in this bryozoan probably functioned as excurrent water chimneys with the radiating channels serving to efficiently collect the previously filtered water, conducting it to the central chimney for expulsion away from the colony surface.
We used micro-CT and SEM to trace zooidal budding in Hornera (Cyclostomatida: Cancellata) from the ancestrula onwards. Results show that hornerid branches are constructed by dual zooidal budding modes occurring synchronously at two separate budding sites at the growing tips. Frontal autozooids bud from a multizooidal budding lamina. Lateral autozooids bud from discrete abfrontal budding loci by 'exomural budding', a previously undescribed form of budding centred on hypostegal pores in interzooidal grooves on the colonial body wall. These two budding modes are integrated during primary branch morphogenesis, forming composite, developmentally bilaminate, branches.
Trepostome bryozoan Dianulites borealis Astrova, 1965, the earliest known member of
this genus, has been identified from the Early Ordovician of Severnaya Zemlya, Arctic Russia. This
species developed hemispherical colonies which indicate that it lived on a relatively soft substrate with
moderately low rates of sedimentation and erosion. The new record from Severnaya Zemlya expands
the palaeogeographical distribution of Dianulites, known before from the Early Ordovician of Novaya
Zemlya, Arctic Russia.
This monograph deals with the morphology of bryozoans of the orders Cystoporida and Trepostomida from the Latorp and Volkhov horizons (Lower–Middle Ordovician, Floian–Darriwilian stages) of the Leningrad Region. Diagnoses of some families belonging to these orders are revised. Based on morphological features of the bryozoans studied and on the succession of their taxa, four assemblages and three stages in their development are established. In addition, the stratigraphic and paleogeographic distributional patterns of bryozoans in the Lower and Middle Ordovician of the world are analyzed.
A new family, Bimuroporidae, is proposed for a clade of Ordovician trepostome bryozoans. The family is united by several characteristics, including a zooidal ontogenetic progression from mesozooid to autozooid and an integrate wall structure. Discriminant and cladistic analyses of colonies from the Ordovician Simpson Group outcropping in the Arbuckle Mountains and Criner Hills of south-central Oklahoma permit the recognition of eight species belonging to this family. Four species assigned to the new genus Bimuropora are described: B. dubia (Loeblich), B. pollaphragmata n. sp., B. conferta (Coryell), and B. winchelli (Ulrich), as well as four species assigned to the genus Champlainopora Ross: C. chazyensis (Ross), C. ramusculus n. sp., C. pachymura (Loeblich), and C. arbucklensis n. sp.
Delicate symmetrical structures in living chambers of autozooecia of some lower Paleozoic species of the order Trepostomata have been interpreted as indications of polypides with little further elaboration. Applications of the anatomy, mode of skeletal growth, and functional requirements basic to living species permit recognition of a few organs of these partially preserved polypides, their functions, and their relationships to enclosing skeletons. Only one basic polypide anatomy is indicated in lower Paleozoic trepostome species, in contrast to the wide diversity of polypide anatomy and their lack of correlation with skeletal characteristics in living stenolaemates. In general, the longer the evolutionary history, the greater the diversity of skeletal morphology in stenolaemates. Apparently, polypide diversity and lack of correlation with skeletal morphology also have increased over time.
The complex skeletal septa in autozooecial apertures of the lower Paleozoic trepostome Hallopora have been interpreted as reflecting an organ of full-sized feeding polypides. Applications of the basic biological concepts of living species suggest that the septa provided reduced skeletal apertures to fit smaller polypides of polymorphs that secondarily occupied the living chambers of full-sized degenerated feeding polypides.
In many species of lower Paleozoic trepostomes (Bryozoa; class Stenolaemata) transverse partitions called skeletal diaphragms differentiated feeding from non-feeding regions of colonies. It has been thought that each diaphragm floored the living chamber of a feeding polypide. However, analysis of skeletal growth patterns has shown that many diaphragms were too close to colony surfaces or too closely spaced in ontogenetic sequences to have accommodated feeding polypides at any given life horizon. Apparently colonies were capable of maintenance and even robust growth with reduced numbers of active polypides, an interpretation supported by comparison with living stenolaemates. A synthesis of the inferred functions of colonies of the extinct trepostomes with post-Triassic fossil and living stenolaemates suggests that walls of trepostome autozooids grew continuously outward so that living chambers starting from their basal diaphragms ranged from shallow to full-sized on colony surfaces. Under-sized polypides apparently grew with their under-sized living chambers and fed as they regenerated to full size, as in living stenolaemates. Actively feeding colony surfaces included autozooids either having polypides at similar or different stages of polypide regeneration, or fully regenerated. Nonfeeding colony surfaces included autozooids either having degenerated polypides, autozooids with diaphragms too closely spaced to skeletal apertures to have housed polypides, or possibly, autozooids that stopped skeletal growth in proximal regions of some large colonies.
Leioclema asperum (Hall), a typical trepostomatous bryozoan, shows well developed acanthopores which are modified parts of orthodox zooecial walls and not separate skeletal entities. They resulted from locally accelerated forward growth of the wall, hence their distinctive cone-in-cone structure. The comparative absence of growth lines in the axial region of each acanthopore suggests that deposition of calcite in that situation was virtually continuous. There is no reason to believe that the acanthopores were originally hollow and they cannot have housed kenozooids, as is commonly supposed.
Observations on living colonies of 56 species of marine bryozoans from Florida and Panama have shown that these organisms possess a variety of morphological and behavioral adaptations related to feeding activities. In the species studied mean lophophore diameter varies from 187 to 1,012 μm, mouth size from 15 to 91 μm, and tentacle number from 8 to 31. Polypide morphology, particularly introvert length and lophophore symmetry, also varies from species to species, and this variation is linked to behavioral strategy. With respect to individual polypide behavior species can range from passive filterers (e.g. Crisia elongata), to tentacle feeders (e.g. Pasythea tulipifera), to cage-captors (e.g. Bugula neritina), to particle jugglers (e.g. Sundanella sibogae). Colony-wide patterns of morphology and behavior also reflect various methods of dealing with water currents. These range from weakly integrated patterns characterized by separation and a high degree of functional independence of individual polypides to highly integrated patterns in which both skeletal morphology and polypide orientation serve to enhance and/or channel feeding currents. Similar strategies have evolved in all three orders of marine bryozoans, apparently in response to the common problem of changing the characteristics of water flow in the immediate vicinity of the colony so that food particles may be extracted.
Polyzoary (?) forming a simple, flattened, unbranehed, two-edged frond, with sub-parallel sides, consisting of two series of cells, the bases of which rest upon opposite sides of a thin longitudinally-striated central membrane or laminar axis, from which they pass obliquely outwards in opposite directions. The cells open in longitudinal rows on the two flat or slightly convex surfaces of the frond, and have the form of more or less cylindrical tubes, which are septate or are divided transversely by a series of well-developed tabulæ. In the only species known the cells of a few of the median rows of the frond are straight, but those of the lateral rows are oblique. Cell-mouths unknown.
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