The effects of ambient flow velocity,colony size,and upstream colonies on the feeding success of Bryozoa. II. Conopeum reticulum (Linnaeus), an encrusting species

The effects of ambient flow velocity, colony size, and the presence of an actively-feeding colony upstream on the feeding success of the encrusting bryozoan Conopeum reticulum (Linnaeus) were studied. Zooids from both large and small colonies showed a reduction in feeding as flow velocity increased, however, the reduction in feeding was less for zooids from large colonies except at very fast ambient flow velocities. The greater pumping capacity of large colonies may result in a relatively greater per zooid feeding success from moving water. The presence of an actively-feeding colony upstream was found to enhance the feeding of zooids on downstream colonies. Diversion of flowing water by actively-feeding colonies upstream may account for the observed enhancement of feeding by zooids on colonies downstream.The results from this study on an encrusting species are compared with results from a previous study on feeding from flow by an arborescent bryozoan, and the feeding performances of these two colony types are related to their respective flow microhabitats.     

Developmental pattern of colonies in the polystyelid ascidian,Polyandrocarpa misakiensis

Summary

The growth pattern of zooids formed asexually by budding was studied in the colonial ascidian, Polyandrocarpa misakiensis. Each colony started from a blas- tozooid (the first generation) on the glass plate in two series of experiments. To evaluate the growth of colonies, lineage of all the zooids of three successive generations was traced on photographs which were taken once a week. The zooids of the first generation produced many buds from any basal margin of the zooidal body, and those of the second generation produced a small number of buds mainly from anterior parts of the zooidal body. The zooids of the second generation produced by early budding of mother zooids were clearly more prolific than those produced by late budding. Circular colonies which developed around a zooid of the first generation consisted of stratified zones of successive generations. Each zone was composed of two subzones; the outer one mainly containing early-produced zooids, and the inner one mainly containing late-produced zooids. The zooids in the marginal area of colony are early-produced ones from generation to generation. The seawater temperature may influence the growth of zooids and/or the frequency of budding.     

Colony Structure in Flustra foliacea (Linnaeus) (Bryozoa,Chéilostomata)

Encrusting forms presumably represent the original mode of growth in the bryozoans. They usually exhibit a relatively low degree of colony integration. The numerous different erect colony types, certainly independently derived from this stage, show a higher degree of integration. Eventually forms arise with a strict pattern only little influenced by the environment. The Flustra foliacea colony holds an intermediate position in this respect. It forms an initial crust on a foreign substratum such as rock. The crust forms marginal lobes. The lobes collide two and two. The two components of each collision then rise back to back, owing to lack of space, into bilaminar fronds. The two zooid layers composing these erect branches and mutually encrusting the “auto-substratum” of their respective basal walls, grow much more rapidly than the initial crust on the “heterosubstratum” of the rock to finally comprise more than 95 % of the colony volume. The different rate of growth on “auto-” and “heterosubstratum” respectively is interpreted as indicating a preference for the “autosubstratum” and as the main force in the formation of erect branches. The Fl. foliacea case indicates one of the pathways from encrusting to erect colonies in the bryozoans. It is probably valid for at least some of the erect bilaminar colonies appearing in different parts of the bryozoan system.     

COLONY FORM AND THE EXPLOITATION OF SPATIAL REFUGES BY ENCRUSTING BRYOZOA

The sheet-runner continuum model of unilaminar encrusting colony growth is reassessed for cheilostome Bryozoa. It is concluded that the model does not adequately account for the existence of spatially predictable refuges from mortality, which can be selected by the larva at the time of settlement.
A third end-point category of colony form, named the spot colony, is recognized for species settling in small spatially predictable refuges and growing to small, early maturing colonies of determinate or semi-determinate size. Similar colonies are reported from spatially restrictive substrates such as flexible algal fronds and single sediment grains on particulate seabeds.
Runner growth is also reappraised. In some cases, uniserial growth may be regarded as a primary adaptation for growth in linear refuges, or on maze-like or strongly three-dimensional surfaces where multiserial growth is impossible, rather than as a general fugitive strategy adopted by competitively inferior forms.
A revised classificatory model for encrusting growth is proposed. This consists of two continua, sheet-ribbon-runner and sheet-patch-spot. It is suggested that an improved ecological classification of encrusting growth might be framed as a series of coupled settlement/growth strategies.     

Adaptive architectural trends in encrusting ectoprocts

Ectoproct colonies with membraniporiform, celleporiform and Iunulitiform A growth-forms have particular habitat preferences on the continental shelf. A colonial capability model for each growth-form, based on a functional analysis of its structure at different levels, tries to portray the adaptive range of the growth-form and includes inferences from the structure of the individual zooids, their polymorphs, their arrangement within the colony as controlled by budding patterns, and their relation to external environmental stresses. Each encrusting growth-form on the Sahul Shelf, northwest Australia, has distinct structural and functional characters which can be arranged in adaptive architectural trends. Deviations from the fundamental membraniporiform growth-form include the development of vertical growth, the sacrifice of individual zooids, the incorporation of substrate within the colony, and variations in the heterozooids. The celleporiform growth-form is subdivided to recognize the massive, 'nodular' colony as a special category.     

Determination of polarity and bilateral asymmetry in palleal and vascular buds of the ascidian Botryllus schlosseri.

Reversal of the bilateral asymmetry of the zooids was induced in a series of colonies of Botryllus schlosseri. Palleal buds from colonies with normal or reversed bilateral asymmetry were isolated in the early stages from the parental zooids and cultured in the vascularized tunic of the same colony or of another colony with opposite asymmetry. Vascular budding was induced in colonies with either type of asymmetry.The bud polarity was shown to depend on the vascularization; the test vessel entering the isolated palleal bud always causes the entrance point to become the posterior end of the developing zooid. On the contrary, the bilateral asymmetric type is predetermined in the bud primordium; the isolated palleal buds develop the type of asymmetry of their parents, even when grafted in the test of a colony with opposite asymmetry. Since the same was also true of the vascular buds, it is concluded that the information for the kind of bilateral asymmetry to be developed is conveyed by the epidermal envelope of the bud. The epidermis of the parental zooids influences the palleal buds, whereas the wall of the test vessels, epidermal extrusions of the zooids, influences the vascular buds.     

Multizooidal Budding in Parasmittina trispinosa (Johnston) (Bryozoa,Cheilostomata)

Two principally different wall types occur in the bryozoan colony: Exterior walls delimiting the super-individual, the colony, against its surroundings and interior walls dividing the body cavity of the colony thus defined into units which develop into sub-individuals, the zooids. In the gymnolaemate bryozoans generally, whether uniserial or multiserial, the longitudinal zooid walls are exterior, the transverse (proximal and distal) zooid walls interior ones. The radiating zooid rows grow apically to form “tubes” each surrounded by exterior walls but subdivided by interior (transverse) walls. The stenolaemate bryozoans show a contrasting mode of growth in which the colony swells in the distal direction to form one confluent cavity surrounded by an exterior wall but internally subdivided into zooids by interior walls. In the otherwise typical gymnolaemate Parasmittina trispinosa the growing edge is composed of a series of “giant buds” each surrounded by exterior walls on its lateral, frontal, basal and distal sides and forming an undifferentiated chamber usually 2–3 times as broad and 3 or more times as long as the final zooid. Its lumen is subdivided by interior walls into zooids 2–3, occasionally 4, in breadth. This type of zooid formation is therefore similar to the “common bud” or, better-named, “multizooidal budding” characteristic of the stenoleamates but has certainly evolved independently as a special modification of the usual gymnolaemate budding.     

Cell proliferation and growth inZoothamnium niveum (Oligohymenophora, Peritrichida) — Thiotrophic bacteria symbiosis

Chemolithoautotrophic, sulphide-oxidizing (thiotrophic) symbioses represent spectacular adaptations to fluctuating environmental gradients and survival is often accomplished when growth is fuelled by sufficient nourishment through the symbionts leading to fast cell proliferation. Here we show 5′-bromo-2′ deoxyuridine (BrdU) pulse labelling of vegetative growingZoothamnium niveum, a colonial ciliate obligately associated with thiotrophic ectosymbionts, and demonstrate age related growth profiles in three heteromorphic host cell types. At the colony’s apex, a large top terminal zooid performed high proliferation activity, which decreased significantly with increasing colony age but was still present in old colonies indicating that this cell possesses lifelong cell division potential. In contrast, terminal branch zooids proliferated independent of colony age but appeared to be limited by their cell division capacity predetermined by branch size, thus leading to the strict, feather-shaped colony form. Appearance of labelled terminal branch zooids allowed us to distinguish a highly proliferating apical colony region from an almost inactive, senescent basal region. In macrozooids attached to the colony, extensive BrdU labelling suggests that DNA synthesis occurs in preparation for a new generation. As motile swarmers, the macrozooids seem to be arrested in the cell cycle and mitosis and cell division occur when the swarmer settles and transforms into a top terminal zooid buildingup a new colony.     

Larger foraminifera as a substratum for encrusting bryozoans (Late Oligocene,Tethyan Seaway,Iran)

Considering the diversity and abundance of larger foraminifera examined from a wide range of Late Oligocene to Early Miocene palaeoenvironments in the Tethyan Seaway, encrusting bryozoans make extremely little use of their tests as substratum. Significant encrustations by bryozoans were exclusively found on large (ø c. 6 cm), undulating tests of Lepidocyclina spp., on which, however, a remarkable 34 taxa of encrusting bryozoans were recorded. This shallow-water fauna of Chattian age was analyzed in respect of the bryozoan taxa present, colony growth type, and mode of budding, colony size, as well as onset of reproduction. Taxic and morphological similarities between the fossil assemblage and modern faunas encrusting mobile substrata indicate a long history of bryozoans as part of the interstitial habitat, while the tests of certain larger foraminifera may have played a significant role in the evolution of shallow-water bryozoans by providing substrata for encrusting species in otherwise unfavorable environments.     

On the rate of budding and proximal attenuation of the graptoloid rhabdosome

Earlier models of the morphogenesis in graptoloid colonies can be improved by taking into account the rate of growth and budding. It is assumed that both these factors are controlled by a specific function of the morphogen, here called for convenience the blastogen , and both are responsible for the attenuation of the proximal part of the rhabdosome. The permanence of this attenuation may theoretically be explained by the universal advantage of differentiation between the proximal and the distal part of the colony. Some aspects of possible adaptive significance of such colony organization are discussed. □ Graptoloid colonies, morphogenesis, mathematical model.     

Kin or self‐recognition? Colonial fusibility of the bryozoan Celleporella hyalina

We estimated fusion frequency with respect to coancestry in the bryozoan Celleporella hyalina, whose briefly planktonic sexually produced larvae settle on algal substrata and proceed to form encrusting colonies by iterative budding. Frequency of fusion between paired colonies growing on an artificial substratum was positively correlated with coefficient of relatedness, with allorecognition ability increasing during the early stages of colonial growth after larval settlement. Parents repressed the growth of F1 progeny with which they had fused. The results are concordant with the Feldgarden-Yund model of selection for self-recognition, which regards fusion with kin as an inevitable source of error whose cost diminishes with increasing relatedness. Contrary to fusion compatibility, gametic compatibility is negatively correlated with coancestry, indicating a need for further research on the possibility of common or linked genetic control that has opposite effect at somatic and gametic levels.     

Structure of Adnate Colony Portions in Crisiidae (Bryozoa Cyclostomata)

The “rhizoidsrdquo; surrounding the base of the erect “colony” emanating from the ancestrula in the Crisiidae, especially the simple species Crisidia cornuta (L.), are a regular adnate system of autozooids. Each autozooid is composed of a proximal adnate part and a distal peristome (in some species kenozooids are possibly intercalated). The autozooid peristomes support erect branches identical in budding and structure with the branch emanating from the erect peristome of the ancestrula. Thus, the complete crisiid colony consists of an adnate system of ancestrula and autozooids, which form erect branches from their peristomes. The adnate zooid system is comparable in autozooid morphology and budding pattern with simple uniserial stomatoporids. The tentative hypothesis is proposed that the crisiid group has developed from primitive stomatoporids; the adnate zooid system of the stomatoporids apparently evolved peristomial budding to produce the erect colony branches characteristic of crisiids.     

Hemichordate skeletal growth: shared patterns in Rhabdopleura and graptoloids

Rhabdopleura shows several features of skeleton growth that are also seen in graptoloids. The similarities between the growth patterns, in terms of the plan towards which the zooids aimed and of their response to environmental disturbance, are profound. Both demonstrate a high degree of genetic control, not only on the gross morphology of the tube or theca but also on the pattern of increments by which this must be achieved. The execution of this 'blueprint' is facilitated by spatial awareness in the zooids of both groups. The main variable left to the graptoloid zooid was the number of increments used in building a theca. Variations in the number of increments probably reflect differences in the productivity of the environment and hence the amount of spare energy in the colony budget. An important new observation is that mortality is common amongst zooids in both Rhabdopleura and graptoloids, with new animals taking over tube or thecal building from where the previous zooid left off. This is identifiable in the increment patterns of tubes and thecae. Several generations of zooids can inhabit a rhabdopleuran tube and can be demonstrated to have inhabited a graptolite theca. This means that innate senescence was not a major cause of death for graptolite colonies. It also means that all thecae might have been continuously occupied and that the colony could have survived significantly bleak environmental conditions by large-scale zooid mortality followed by regeneration. □ Graptolite, ecology, hemichordate, pterobranch, RHABDOPLEURA, growth.     

Relative size predicts competitive outcome through 2 million years

Competition is an important biotic interaction that influences survival and reproduction. While competition on ecological timescales has received great attention, little is known about competition on evolutionary timescales. Do competitive abilities change over hundreds of thousands to millions of years? Can we predict competitive outcomes using phenotypic traits? How much do traits that confer competitive advantage and competitive outcomes change? Here we show, using communities of encrusting marine bryozoans spanning more than 2 million years, that size is a significant determinant of overgrowth outcomes: colonies with larger zooids tend to overgrow colonies with smaller zooids. We also detected temporally coordinated changes in average zooid sizes, suggesting that different species responded to a common external driver. Although species‐specific average zooid sizes change over evolutionary timescales, species‐specific competitive abilities seem relatively stable, suggesting that traits other than zooid size also control overgrowth outcomes and/or that evolutionary constraints are involved.     

Does polymorphism predict physiological connectedness? A test using two encrusting bryozoans

A colonial lifestyle necessitates communication between colony members to coordinate functions and enable resource sharing through physiological integration. Colonial integration is predicted to increase with both the size of the colony and the level of specialization (polymorphism). In modular colonies, although integration might be reflected in structural characteristics such as module spacing or branching patterns, physiological integration is fundamentally dependent on the level of connectedness between modules. In cheilostome bryozoans, funicular tissue links adjacent zooids through pores within zooid walls and is the most likely means of nutrient transport within colonies. We sought to determine whether the relative numbers of pores (septulae) and pore plates (septal chambers) per zooid differed across colony regions in a monomorphic species, Watersipora subtorquata, and one showing some polymorphism, Mucropetraliella ellerii. Within each species, the morphology of pore plates corresponded to functional predictions based on their position within the zooid, and connection numbers per zooid increased with colony size. Contrary to expectations, however, the more complex species, M. ellerii, had significantly fewer porous connections per zooid than W. subtorquata. Physiological connectedness was therefore not predicted by simple assessment of polymorphism in these species and may not be sufficient to infer colonial integration in related taxa.     

Jurassic corals as emersion indicators

In Recent coral reefs, emersion has resulted in the development of special growth forms of colonial corals which have been named microatolls. These colonies grow almost exclusively in a horizontal direction. In the upper part of the colony, subaerial exposure leads to the decay of the living body. The growth then stops and the skeleton is later colonized by various boring and encrusting organisms. Here we show the first record of a massive Mesozoic coral colony displaying the main features of emersion. The colony has been collected in perireefal Oxfordian (Jurassic) limestones from the Jura Mountains (France). It is situated exactly at the expected place in a shallowing-upward sequence between infralittoral buildups and supralittoral limestones. We suggest that such growth structures could be more common than previously thought in ancient coralliferous sediments and add some new details to discriminate between these colonies.     

Paraboloid colony bases in Paleozoic stenolaemate bryozoans

Dendroid stenolaemate bryozoan colonies with paraboloid bases developed initially in the same manner as stenolaemate colonies with broadly encrusting bases. Their unique shape is related to the narrowly cylindrical shape of the encrusted surface (algal stipe?) and to radially differentiated rates of growth from the point of colony origin. The colony shape is interpreted as an adaptation to unconsolidated substrates in relatively quiet though not necessarily deep water.     

Quantitative observations on asexual reproduction of Aeolosoma viride (Annelida,Aphanoneura)

Agametic reproductive activity (via paratomy) of Aeolosoma viride was analyzed throughout the life cycle in individually reared specimens. Aeolosoma viride is organized in linear chains of 3–4 zooids; the main zooid is anterior, and the secondary zooids are positioned posterior to the main zooid in inverse order with respect to their degree of growth, the most advanced being at the posterior end, and those less advanced nearer the main zooid. On average, worms lived 66±10 d and produced 57±6 offspring. A budding area located in the sub‐terminal part of the main zooid produced chaetigers that formed the origin of the secondary zooids. A growth zone was located in the posterior end of each secondary zooids. Fission occurred between the penultimate and the last zooid of the chain. Just before fission, the growth zone of each secondary zooid became a budding area. Agametic reproduction was via multiple paratomy with linear succession of the secondary zooid and terminal fission. The structure of the chain was therefore modulated by the interaction of the processes of budding, growth, cephalic differentiation, and fission, which occurred continuously and on different timescales. Values of parameters describing paratomic activity (interval between origin of the zooids, time to produce a chaetiger, growth time of the zooids, and interval between the fission of the filial chains) are low early in an individual's life, but increase during senescence. Due to its relatively rapid lifecycle and high reproductive activity, A. viride is a convenient experimental organism for the study of agametic reproduction.     

The genera Cribrilina and Collarina (Bryozoa, Cheilostomatida) in the British Isles and North Sea Basin, Pliocene to present day

Seven species assigned to the genus Cribrilina are illustrated by scanning electron microscopy (SEM) and redescribed. At the present day, three species occur in the area studied: C. punctata (Hassall), C. cryptooecium Norman and C. annulata (Fabricius). The presence of Collarina balzaci (Audouin), which has previously been confused with Cribrilina punctata , is also reported. The following species of Cribrilina are known as Pliocene and Pleistocene fossils: C. punctata, C. cryptooecium, C. watersi Andersson, C. puncturata (Wood) and C. mucronata Canu & Lecointre. In addition, C. annulata and C. spitsbergensis Norman have been reported as fossils, but these occurrences could not be confirmed here by the examination of actual material. Lectotypes are selected for C. spitsbergensis, C. watersi, C. puncturata and C. mucronata. Type material of the American fossil C. marylandica , which has been synonymised with C. punctata , is illustrated.     

Effect of zooid spacing on bryozoan feeding success: is competition or facilitation more important?

Most Recent bryozoan species are encrusting sheets, and many of these colonies have densely packed feeding zooids. In this study, I tested whether tight packing of feeding zooids affects food capture. Colonies of a bryozoan with an encrusting sheet form (Membranipora membranacea) were dissected to produce individuals whose feeding zooids were (1) closely packed, (2) more widely spaced, or (3) isolated. For each type, rates of particle ingestion were measured in still water and in a freestream velocity of 2.7 cm s(-1). Ingestion rate increased when zooids were closest together, probably because of reduced refiltration and increased feeding current strength farther from the lophophores. The mean incurrent velocity within 0.02 cm above the center of the lophophore was 0.28 cm s(-1) regardless of zooid spacing; however, when the incurrent velocity was measured more than 0.1 cm from the lophophores, zooids that were close together or spaced one zooid's width apart had significantly faster incurrent velocities than single zooids. Flow visualization suggests that isolated zooids and those spaced far apart refilter more water than zooids that are close together. These results along with the observed trend of increased zooid integration over evolutionary time suggest that the benefits of increasing coordination outweigh the consequences of intrazooid competition.