The effects of ambient flow velocity,colony size,and upstream colonies on the feeding success of bryozoa. I. Bugula stolonifera Ryland,an arborescent species

The effects of ambient flow velocity, colony size, and the presence of upstream colonies on the feeding success of the arborescent bryozoan, Bugula stolonifera (Ryland), were studied. Faster ambient flow velocities were found to reduce feeding of zooids of small colonies but not of large colonies. Zooids from different regions of colonies dominated in feeding at different ambient flow velocities: upstream zooids dominated in feeding from slow ambient flow: zooids from central regions dominated in feeding from fast ambient flow. These results are interpreted with respect to the branching morphology of colonies. Finally, evidence that upstream colonies interfere with the feeding success of zooids of colonies downstream was obtained.     

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.     

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.     

Effects of ambient flow and injury on the morphology of a fluid transport system in a bryozoan

Many organisms use fluid transport systems that are open to the external environment for suspension feeding or gas exchange. How do factors related to the environment, such as injuries and ambient currents, affect remodeling of these systems? In the bryozoan Membranipora membranacea, the lophophores (crowns of ciliated tentacles) form a canopy over the colony. The lophophores pump seawater from above the colony through themselves to capture food particles. The seawater then flows under the canopy to exit the colony at chimneys (openings in the canopy) or at the canopy edge. To test whether either ambient flow speed or injury affects remodeling of this system, I measured changes in chimney size and spacing in colonies grown in flow tanks at different ambient flow speeds, and in colonies in which I killed patches of zooids. There was no effect of either ambient flow speed or injury size on chimney remodeling. Injury did not induce chimney formation. In addition, chimneys formed at the canopy edge, indicating that high pressure under the canopy did not induce chimney formation. These results suggest that ambient flow, injury, and the pressure under the canopy may have little effect on the remodeling of this fluid transport system.     

A new early cyclostome bryozoan from the Lower Ordovician (Volkhov Stage) of Russia

Goryunovia hemiseptata n.gen. n.sp. is an unusual Lower Ordovician cyclostome bryozoan, represented by a single colony from the Middle Volkhov Stage (Arenig) of the Leningrad Oblast. The encrusting uniserial colony consists of large, slender zooids, many with closely-spaced partial diaphragms (hemisepta), and has a distinctive trichotomous branching pattern. Skeletal organization is of the sagenellid grade: calcified interior walls appear to be present, and the exterior frontal walls lack pseudopores. As details of microstructure and early astogeny are unknown, it is difficult to interpret the phylogenetic relationships ofGoryunovia until further material is forthcoming. However, there is some resemblance with the enigmatic hederellids, which appear in the Upper Silurian and possibly range into the Triassic.     

Schischcatella (Fenestrata,Bryozoa) from the Devonian of the Rhenish Massif,Germany

Abstract: The Devonian fenestrate bryozoan, Schischcatella Waschurova, 1964 , possessed colonies in the form of low, erect bifoliate fronds that grew from an encrusting sheet‐like base with autozooecia arranged in biserial, bifurcating rows. This growth habit is unique in fenestrates, which normally had unilaminate arborescent colonies. Originally, Schischcatella was described from the Lower Devonian of Tajikistan. This article describes a new species, S. heinorum sp. nov., from the Middle Devonian of the Eifel (western Rhenish Massif, Germany) with additional material from the Lower Devonian of the Kellerwald (eastern Rhenish Massif, Germany). External and internal morphologies of this bryozoan have been studied using abundant material. The growth habit of Schischcatella suggests a completely different pattern of feeding currents than that in the normal fenestrate colony. The outflow of the filtered water occurred only on edges of colonies between rami. In the absence of chimneys (areas of vertical water expelling), such a functional morphology may have restricted extension of the colony in a distal direction. The evolution of Schischcatella is apparently an example of paedomorphosis, the genus evolved from an unknown semicosciniid species by the early ontogenetic interruption of colony development and further changes in the mode of growth.     

Epibiont foraminifera of Sertella frigida (Waters) (Bryozoa, Cheilostomata) from Terranova Bay, Ross Sea, Antarctica

Several arenaceous and calcareous foraminifera epibionts of Sertella frigida (Waters) (Bryozoa, Cheilostomata), collected in Terranova Bay during the 1989–90 Italian Antarctic expedition, were studied. Arenaceous species, such as Psammosphaera fusca Schulze f. adhearescens Rhumbler, Haplophragmoides canariensis d'Orbigny, Portatrochammina antarctica (Parr), and Trochammina arctica Hedley, Hurdle and Burdett, are a conspicuous component of the epifaunal community. The calcareous foraminifers encrusting S. frigida are mostly represented by Cibicides refulgens Montfort and Rosalina globularis d'Orbigny. Foraminifera were found only in the basal part of the bryozoan colonies. The presence of many juveniles provided evidence that foraminifers were reproducing at the time of sampling. Psammosphaera fusca f. adhaerescens was found exclusively inside the zooids of S. frigida. The large number of individuals associated with the bryozoan suggest that P. fusca f. adhaerescens finds optimum conditions (shelter, food, and grains for shell building) for growth inside the zooids of S. frigida. The other species occurred most commonly adhered to the trabeculae or to the zooid orifices of S. frigida. An elevated position offers a better chance to catch food particles from the overlying water column. Foraminifers could benefit also from bryozoan feeding currents. The availability of resuspended organic material could provide a more consistent source of food in a highly seasonal oligotrophic environment. Received: 8 February 1995/Accepted: 30 June 1996     

Effect of colony size on the competitive outcome of encrusting colonial organisms

The relative importance of colony size ratio of interacting species was studied in Tomioka Bay, Japan. Six encrusting colonial species belonging to the following three different taxonomic groups were tested: Ascidia (three species), Bryozoa (two) and Porifera (one). Colonies of these organisms were grown in the community of sessile organisms developed on plastic panels. Logistic regression analysis was carried out to determine the effect of size ratio on the competitive outcome of interacting colonies. The results between all possible combinations among these six species did not show a significant size effect in competitive outcome (i.e. a larger colony size did not always prove important in the success of a competitive interaction with smaller colonies of other species). On the contrary, competitive success depends on the types of species interacting. Certain species such asDidemnum moseleyi (ascidian) andHaliclona sp. (sponge), in spite of being smaller in colony size, won in competitive interactions with larger colonies of other species such asDiplosoma mitsukurii (ascidian) andWatersipora subovoidea (bryozoan). These results contradict the one reported earlier: that the larger the colony size, the more chance the colony will have to win in competitive interactions.     

The inference of extrazooidal feeding currents in fossil bryozoan colonies

Taylor, Paul D. 1979 01 15: The inference of extrazooidal feeding currents in fossil bryozoan colonies. Lethaia , Vol. 12, pp. 47–56. Oslo. ISSN 0024–1164.
Previous studies on live bryozoan colonies have shown that the feeding autozooids in a colony may cooperate in differing ways to produce an extrazooidal watercurrent system organised on a colony-wide or subcolony-wide basis. The presence of an extrazooidal current system may be inferred in fossil stenolaemate bryozoans which exhibit either a differential spacing of open autozooecial apertures or a systematic variation across the marial surface in the orientation of automoecial distal portions. By inference, aggregations of autozooecial apertures represented loci of inhalant extrazooidal flow whereas zoarial protuberances (e.g. monticules) with outwardly leaning autozooecia acted as loci of exhalant extrazooidal flow. Bryozoans having automoecia opening obliquely into gaps or fenestrules in their zoaria probably drew a unidirectional extrazooidal current of water through the fenestrules. Extrazooidal water currents may function to accelerate colony clearance rate, decrease the chances of recycling filtered water, aid spermatoman and larval dispersal, and clear sediment from the colony surface.     

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.     

Flow and polypide distribution in the cheilostome bryozoan Bugula and their inference in Archimedes

Cilia-generated flow in the absence of ambient current is directed from frontal to reverse sides of branches in Bugula turrita, B. turbinata, B. neritina , and B. stolonifera , whether axes of feeding lophophores are perpendicular to the basal plane of branches or are tilted toward distal ends of branches. Ambient current less than 5 cm per second interacts with cilia-generated flow, but ambient flow of 15 cm per second destroys self-generated colonial flow and severely hampers feeding. Polypides are located in the more distal, younger portions of colonies, in species with and without polypide recycling, whereas zooids in the more proximal, older portions are senesced. Presence of feeding polypides in distal but not in proximal portions of the larger spiralled colonies of B. turrita and B. turbinata results in downward, slightly radially directed flow through the colony. The colonial flow passes directly from one whorl to the next-proximal so that water exits from low around the colony perimeter, and a proximally expanding conical stagnant zone occupies the interior of the colony. A substantial percentage of zooecia in distal whorls of well-preserved Archimedes is filled by sediment and inferred to have been occupied by actively feeding polypides. whereas spar-filled zooecia capped by terminal diaphragms were apparently senesced during the latter part of a colony's existence. The capped zooecia constitute an increasing percentage of the total in more proximal whorls. Generally similar colony form and inferred similarity in distribution of current-generating polypides in spiralled colonies of Bugula and in Archimedes suggest that colony-generated flow in Archimedes was similar to that in Bugula , passing downward and then outward, and only through the distal whorls of the colonies.     

Hoeverella krauseae gen. et sp. nov., an unusual uniserial cheilostome bryozoan from the campanian of hannover

Hoeverellakrauseae gen. et sp. nov. is a peculiar runner-like anascan cheilostome bryozoan known from a single well-preserved specimen encrusting a belemnite guard. The colony comprises about 60 zooids, each of fusiform/pyriform shape and having a small “opesia” within which is situated an enigmatic ring structure. Several interpretations are possible for the ring structure: “regeneration”, closure plate, foreign organism inhabiting the zooecium, or socket of articulation for an erect branch or anAetea- like erect distal tube. Unusually for a runner-like bryozoan only one branch ramification occurs in the entire colony. The systematic position ofHoeverella is uncertain; however, it may be related to the electrid genusHerpetopora.     

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.     

Will variation among genetic individuals influence species responses to global climate change?

Increased anthropogenic CO₂ emissions in the last two centuries have lead to rising sea surface temperature and falling ocean pH, and it is predicted that current global trends will worsen over the next few decades. There is limited understanding of how genetic variation among individuals will influence the responses of populations and species to these changes. A microcosm system was set up to study the effects of predicted temperature and CO₂ levels on the bryozoan Celleporella hyalina. In this marine species, colonies grow by the addition of male, female and feeding modular individuals (zooids) and can be physically subdivided to produce a clone of genetically identical colonies. We studied colony growth rate (the addition of zooids), reproductive investment (the ratio of sexual to feeding zooids) and sex ratio (male to female zooids) in four genetically distinct clonal lines. There was a significant effect of clone on growth rate, reproductive investment and sex ratio, with clones showing contrasting responses to the various temperature and pH combinations. Overall, decreasing pH and increasing temperature caused reduction of growth, and eventual cessation of growth was often observed at the highest temperature, especially during the latter half of the 15‐day trials. Reproductive investment increased with increasing temperature and decreasing pH, varying more widely with temperature at the lowest pH. The increased production of males, a general stress response of the bryozoan, was seen upon exposure to reduced pH, but was not expressed at the highest temperature tested, presumably due to the frequent cessation of growth. Further to the significant effect of pH on the measured whole‐colony parameters, observation by scanning electron microscopy revealed surface pitting of the calcified exoskeleton in colonies that were exposed to increased acidity. Studying ecologically relevant processes of growth and reproduction, we demonstrate the existence of relevant levels of variation among genetic individuals which may enable future adaptation via non‐mutational natural selection to falling pH and rising temperature.     

Organization and Evolution of Animal Colonies

Based on the theory of organization and evolution of colonies in an extinct group of hemichordate graptolites (Urbanek, 1960, 1990) the relationship between the events in the late astogeny of bryozoan colonies and their somatic and reproductive cycle is proposed. The bryozoan colonies with simple organization are compared with graptoloid colonies and their structure is interpreted within the framework of the morphogen gradient theory. A morphogen produced by the founder-zooid (oozooid) diffuses along the long axis of the colony and controls the phenotypic expression of the size and structure of zooids. Evolutionary changes in the graptoloid colonies involve introduction of new characters and their spreading is also accompanied by gradient changes of their manifestation. Evolutionary mechanisms in bryozoan colonies are considered in terms of penetrance and expressivity of genes. In contrast to graptolites, many bryozoan colonies display multiple zones of astogenetic changes and repetitions.     

Some regularities in the development of encrusting colonies of Cribrilina annulata (Fabricius, 1780)

The growth of encrusting colonies was studied with mathematical model. It was shown that encrusting growth takes place under increasing competition for the substrate inside the colony. The model was tested on the example of Cribrilina annulata collected in White Sea on Laminaria saccarina. All colonies were mapped, zooids were measured and genealogical connections between them were established. A number of gradients were revealed by statistical methods. The intensity of budding decreases in astogeny according to theoretical predictions. It was shown that development of Cribrilina annulata colonies is strictly determined by gradients that can be caused by shortage of substrate space. It leads to the suppression of budding and changing in zooid size. Increasing substrate shortage is predictable and caused by the regularities in of zooid budding. The growth of colony stops after exhausting of potentially available substrate.     

Zooid size and growth rate of the bryozoan Cryptosula pallasiana Moll in relation to temperature, in culture and in its natural environment

The size of cheilostome bryozoan zooids has been widely discussed for its potential in inferring palaeotemperatures, based on correlations between zooid size and temperature. Studies in both the natural environment and under experimental laboratory conditions have shown that an increase in temperature significantly decreases zooid size in a range of bryozoan taxa. In order to test the effect of temperature on zooid size, the cheilostome bryozoan Cryptosula pallasiana was for the first time successfully cultured under laboratory conditions. C. pallasiana was grown at 14 °C and 18 °C using Rhodomonas sp. as a food organism. Zooid size, tentacle number and growth rate were measured over a period of 26 days. For comparison, zooids from colonies of C. pallasiana collected from the natural environment were measured in winter and summer months. Results showed that colonies grown in laboratory culture had significantly longer and wider zooids at 14 °C than at 18 °C. The specific growth rate of C. pallasiana doubled from 14 °C to 18 °C. Comparison of tentacle number in culture showed a significantly higher value at lower temperatures. This may be related to differing food availability, longer polypide life spans, or a shift of energy use at colder temperatures. In nature the zooids were significantly longer in colonies sampled in July than in January, a clear difference from laboratory results. The utility of cheilostome Bryozoa as indicators of environmental change and their potential for studies of paleotemperature are highlighted.     

A new encrusting interstitial marine fauna from Brazil

Abstract. This paper reports the second occurrence of a sand‐grain encrusting interstitial epifauna dominated by bryozoans and polychaetes at a site thousands of kilometers from the first described occurrence of such a fauna 20 years ago. Such faunas seem to have gone almost unrecorded in the marine ecological literature, but they are potentially geographically widespread and ecologically significant, deserving recognition and further study by benthic ecologists. Although rooted‐erect and free‐living lunulitiform bryozoans can be abundant in soft‐bottom habitats, the presence of encrusting forms was, until recently, considered to be limited to patches of hard substrata. In 1985 and 1988, a new and seemingly unique habitat for encrusting bryozoans and other organisms on single grains of shell or sand was reported from the coastal waters of Florida, USA. Here we report a second discovery of an interstitial encrusting fauna from the continental shelf off the state of São Paulo, Brazil. In addition to the cupuladriid Discoporella umbellata, several species of bryozoans (9 cheilostomes, 3 ctenostomes, and 1 cyclostome) were found encrusting on or boring into sand grains from the 4 stations examined. Four species were found exclusively on sand to gravel size grains. The most abundant colonies, with ～1300–1500 colonies m^?2, belonged to a new species of Cleidochasma. New species of Trypostega and Reginella, each with up to 200–300 colonies m^?2, were also discovered. The grain‐encrusting bryozoans were characterized by their small size, and by the fact that sexual reproduction was initiated very early in colony growth; brood chambers (for the development of embryos into larvae) occurred in colonies having only a few zooids. Colonies of boring ctenostome and cheilostome bryozoans were even more abundant than those of grain encrusting forms, being present in almost every piece of shell (～5000–5500 colonies m^?2). The fauna also included representatives of other groups of encrusting organisms, especially tubeworms (11,000–13,000 tubes m^?2). Planned work on samples from additional stations on the São Paulo shelf will no doubt yield a larger number of species from various taxa and perhaps show some overlap in sand fauna species between the Brazilian and Floridian sites. In addition to the unique species of single grain encrusters, colonies of bryozoan species characteristic of larger subtidal hard substrata were also found on sand or gravel size grains, indicating that an interstitial refuge may be available to some epifaunal taxa and suggesting that this interstitial refuge, which remains almost completely unknown to benthic ecologists, may play a large role in determining distributions of those taxa.     

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.     

The effect of water flow on growth and reproduction of celleporella hyalina (l.) (bryozoa : cheilostomata)

Colonies of Celleporella hyalina (L.), settled on glass Petri dishes, were grown in the Menai Straits. Water flow over the colonies was either unrestricted or increasingly restricted by short and long funnels fitted over the Petri dishes. Under all conditions, colonies grew exponentially for the first few weeks, but then growth decelerated. Growth rate at standard size was minimal in winter (December–February). Restricted water flow enhanced growth during winter but retarded growth at other times. Colonies began sexual reproduction at a size of ≈57 autozooids, but at different ages depending on growth conditions. When colonies had accumulated 1000–2000 autozooids, the number of reproductive zooids (males plus females) per autozooid became asymptotic in the ratio of 1.0 : 2.0 : 3.6 in conditions of restricted, semi-restricted, and unrestricted water flow, respectively. The proportion of sexual zooids that were reproductively active was greater among colonies experiencing greater water flow and, for all treatments, was greatest in June and least in winter. The experimentally induced trends in growth and reproduction were reversed by translocating colonies from one regime of water flow to another. Colonies had lower budding rates when grown close to neighbouring colonies than when isolated, but the ratio of reproductive zooids to autozooids was similar in both treatments. Ratios of female to male zooids varied among colonies but with no clear relation to the experimental conditions, suggesting that relative investments in male and female functions are largely under genetic control. The life history of C. hyalina is well-suited to the ephemeral frondai tissue of Laminaria saccharina L. (Lamour), commonly used as a substratum in British waters. Early sexual maturity and sexual activity, concurrent with somatic growth, maximize average sexual output in the circumstances of unpredictable longevity.