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.     

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.     

Recruiting juvenile damselfish: the process of recruiting into adult colonies in the damselfish Stegastes nigricans

Juveniles of Stegastes nigricans occur in adult colonies, solitarily, and occasionally in juvenile colonies. We concentrated on solitary juveniles and those in adult colonies. We examined the costs and benefits of different settlement strategies, quantified the territory requirements of adults, and investigated the process of how juveniles make the transition to adult territorial fish. An adequate adult territory lies next to those of other adults, is proportional in area to the size of the adult, and contains a refuge tunnel whose entrance is sufficiently large. Compared with solitary juveniles, those <4 cm total length inhabiting adult colonies experienced reduced heterospecific competition for algal food and consequently benefited from a greater density of algae. A cost of recruiting into an adult colony, however, was increased attacks by adults. Juveniles that settled in adult colonies avoided attacks by retreating into small holes inaccessible to adults. As juveniles in adult colonies grew, they were chased less often by adults, whereas they themselves chased adults and heterospecific fish more often. Because territory size correlated with fish size in adult colonies, its area had to expand as the young fish grew, and that expansion was done at the expense of neighbors. Obtaining the space needed by an adult may be possible only when the juvenile settles directly into an adult colony. Juveniles that first settle down solitarily, or in juvenile colonies, may later attempt to enter adult colonies. However, because they do so as larger juveniles, they would have difficulty insinuating themselves into small refuges, which is essential for retreat from the adults. Received in revised form: 4 January 2001 Electronic Publication     

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.     

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.     

Persistence and space preemption explain species‐specific founder effects on the organization of marine sessile communities

Community assembly may not follow predictable successional stages, with a large fraction of the species pool constituted by potential pioneering species and successful founders defined through lottery. In such systems, priority effects may be relevant in the determination of trajectories of developing communities and hence diversity and assemblage structure at later advanced states. In order to assess how different founder species may trigger variable community trajectories and structures, we conducted an experimental study using subtidal sessile assemblages as model. We manipulated the identity of functionally different founders and initial colony size (a proxy of the time lag before the arrival of later species), and followed trajectories. We did not observe any effects of colony size on response variables, suggesting that priority effects take place even when the time lag between the establishment of pioneering species and late colonizers is very short. Late community structure at experimental panels that started either with the colonial ascidian Botrylloides nigrum, or the arborescent bryozoan Bugula neritina, was similar to control panels allowed natural assembling. In spite of high potential for fast space domination, and hence negative priority effects, B. nigrum suffered high mortality and did not persist throughout succession. Bugula neritina provided complex physical microhabitats through conspecific clustering that have enhanced larval settlement of late species arrivals, but no apparent facilitation was observed. Differently, panels founded by the encrusting bryozoan Schizoporella errata led to different and less diverse communities compared to naturally assembled panels, evidencing strong negative priority effects through higher persistence and space preemption. Schizoporella errata founder colonies inhibited further conspecific settlement, which may greatly relax intraspecific competition, allowing resource allocation to colony growth and space domination, thus reducing the chances for the establishment of other species.     

Defense of Food Supply by Eusocial Colonies

Overdispersion of colonies exists in many eusocial insects.Overdispersion can be generated by direct attack on coloniesor founders, by defense of space, by defense of food resourcesbeing harvested, or by exploitative competition. When directcompetitive interactions lead to colony overdispersion, territorialityis said to occur. Whereas solitary territory holders typicallydefend space, most eusocial colonies defend resource patchesrather than space per se. Also unlike solitary territory holders,colonies with forager communication can simultaneously defendseveral spatially separated food patches. A model explores optimalnumbers of scouts (discoverers of patches) and recruits (followers)needed to maximize net rate of energy intake by the colony.Territorial costs are added to the model by requiring a higherinvestment of foragers per unit resource collected. Accordingto the model, optimal colony size and percentage scouts aremore sensitive to changes in patch size than in patch density.If patch defense is required for resource control, a declineoccurs in optimal percentage of scouts; the decline is greatestfor small colonies. Colonies that must defend patches in orderto harvest from them suffer a loss in net energy intake; theloss is greatest for small colonies. It is predicted that amongeusocial insects, those with territorial defense of resourcesshould preferentially visit large patches and have comparativelylarge colony sizes and relatively few scouts. Ways of testingthese predictions are discussed.     

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.     

Effects of coral colony morphologies on mass transfer and susceptibility to thermal stress

This study tested the hypothesis that corals of the same species, but of varying size and shape, may respond differently to thermal stress because of different mass transfer capacities. High mass transfer rates are an advantage under thermal stress, and mass transfer rates are assumed to scale with size. Yet large, corymbose Acropora colonies are more vulnerable to thermal stress than small corymbose Acropora colonies. We took a two-tiered approach to examine the differences in the susceptibility of different coral morphologies to thermal stress. Firstly, the response of several coral species of different sizes and shapes were measured in the field through a thermal stress event. Secondly, diffusion experiments were conducted using gypsum-coral models of different morphologies to estimate mass transfer rates, to test whether dissolution rates differed in accordance with colony morphology and colony size. Coral colonies with a high height to diameter ratio were subjected to more partial mortality than flat colonies. These results agree with mass transfer theory. The diffusion experiments showed that in a low-flow environment, small encrusting colonies had higher rates of dissolution than large flat or small branched colonies. These results, however, disagree with mass transfer theory. We show that the volume of space between colony branches predicts the response to thermal stress in the field. Small encrusting colonies were most likely to maintain mass transfer and were therefore more likely to survive thermal stress than large branched colonies. We predict that an increase in the frequency and intensity of thermal stresses may see a shift from large branched coral colonies to both small colonies, and flat-massive colonies with low aspect ratios.     

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.     

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.     

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.     

Growth-related changes in predation behavior in incipient colonies of the ponerine antEctatomma tuberculatum (Olivier)

Summary We traced the development in the laboratory of 18 young colonies of the arboricolous ponerine antEctatomma tuberculatum. Colony foundation is of the partially-claustral type. During the early stages, when the colony is entirely dependent on the queen's behavior, the growth of the colony in terms of number of workers produced over time was relatively predictable. Afterwards, divergence in colony growth in function of the time increases as fast as the number of workers influences the efficiency of colony provisioning.Comparative analysis indicated clear changes in the predation behavior of foundresses and workers as colonies developed. For any stage of colony growth, all individuals provisioned the nest with dead prey or sugar-rich substances in the same way. However, prey hunting involves two different strategies. Foundresses and nanitic workers (originating from colonies with 9–15 workers) foraged actively, catching prey as the result of random encounters. Post-nanitic foragers (originating from colonies with 20–30 workers) and those from nature colonies developed an ambush strategy. Workers in these colonies gained experience at catching and handling prey during a period when they acted as nest guards, and so tended to be more efficient hunters than poorly experienced foundresses or nanitic foragers. The change in strategy was also positively correlated with an increase in the size of workers as the colony matured. A stable maximum in workers size is apparently reached only after the appearance of efficiently hunting foragers, presumably in numbers sufficient to provide adequate quantity and quality of larval food. Such a correlation between worker size and colony growth, assumed general for all ants, has not been demonstrated for Ponerinae before this work.     

Local interactions drive size dependent space competition between coral and crustose coralline algae

For many species, the outcome of competition for space in homogeneous habitats depends upon relative rates of growth and overgrowth. Size dependence in competition occurs when this balance shifts due to the growth of one or both species. For example, the ability of coral to compete with certain species of crustose coralline algae (CCA) may depend on whether coral colonies are large enough to avoid being overgrown. Spatially implicit models suggest size refuges from competition can improve the persistence of species with a vulnerable life stage. We use spatially explicit simulation models to explore size dependence in competition between coral and competitively dominant CCA in well lit habitat. We determine what conditions allow coral to use size refuges and whether refuges improve the recovery of coral after disturbance. Local interactions in explicit space prevent the maturation of coral into size refuges unless coral grows more rapidly than CCA or coral colonies are allowed to fuse, and mortality mechanisms can limit long‐term persistence even if the refuge is achieved. We contrast results with analogous differential equation models, with and without an explicit maturation delay, to demonstrate how the predicted outcome of competition is frequently reversed when local interactions and individual‐based dynamics are included in models of size‐dependent competition.     

Ecology of cryptic coral reef communities. IV. Community development and life histories of encrusting cheilostome bryozoa

Life histories of encrusting cheilostome species from the cryptic reef community at Rio Bueno, Jamaica, were studied on fouling panels over 3 yr. Recruitment and growth were generally slow compared with those reported for temperate cheilostomes. Most species that became abundant and persisted throughout the study did so through relatively rapid growth to a large size by a few successful colonies, rather than by accumulating great numbers of small colonies. This pattern, which reflects a striking increase in maximum survival with increasing colony size, is the basis for the extremely patchy distributions of bryozoans under corals. Reproduction in these species is delayed, and only a few long-lived, large colonies ever reproduce. Only one species, Drepanophora tuberculatum (Osburn), approached the characteristic opportunistic pattern of high recruitment, small colony size, and early reproduction.Grazing and nesting activities of one yellowtail damselfish greatly affected distributions of major taxa and cheilostome species on one set of panels. Species more abundant on grazed panels are more heavily calcified and showed other protective features, compared with species more abundant elsewhere. Despite intensive grazing by the fish, overgrowth interactions occurred frequently on both sets of panels. The fish affected what organisms were present, but did not obviously reduce the amount of overgrowth.     

Biodiversity and reduced extinction risks in spatially isolated rodent populations

Ecologists have rarely explored the potential influence of local (alpha) biodiversity on the stability and local extinction of spatially isolated populations. Twenty years of annual counts of a small, grazing rodent (Utah prairie dogs, Cynomys parvidens ) from 20 different isolated local populations (colonies) in southern Utah, U.S.A. were analysed. These prairie dogs exhibited large fluctuations and repeated extinctions at individual colonies during the census period. Frequency of extinction at a colony declined dramatically as the number of locally occurring plant species increased. This pattern was not explained by differences among colonies in plant productivity, plant species composition, colony size, or variability in annual counts. Thus, lower extinction risk of consumer populations may be associated with greater resource diversity, and maintaining high local plant diversity may help sustain spatially isolated herbivore populations in fragmented habitats.     

Colony-Site Selection Drives Management Priorities for Yelkouan Shearwater Populations

Abstract: We tested whether colony-site availability could allow for an increase in the unusually small breeding populations of yelkouan shearwater (Puffinus yelkouan) on the islands of the Port-Cros National Park (France) if feral cat eradication were undertaken. Comparisons between colony and noncolony sites indicated yelkouan shearwaters preferred deep-soiled and low-outcrop-covered coastal sites. A substrate cover, light avoidance, and sea proximity model suggested that 17.5% of unoccupied sites are suitable for colony establishment. The low proportion of suitable sites currently used by yelkouan shearwaters suggests that these colonies could be refuges and that feral cat eradication will probably lead to a breeding population increase.     

Experimental evidence that refuges delay insect adaptation to Bacillus thuringiensis

Theoretical projections suggest that refuges from exposure can delay insect adaptation to environmentally benign insecticides derived from Bacillus thuringiensis, but experimental tests of this approach have been limited. We tested the refuge tactic by selecting two sets of two colonies of diamondback moth (Plutella xylostella) for resistance to B. thuringiensis subsp. aizawai in the laboratory. In each set, one colony was selected with no refuge and the other with a 10 per cent refuge from exposure to B. thuringiensis subsp. aizawai. Bioassays conducted after nine selections were completed show that mortality caused by B. thuringiensis subsp. aizawai was significantly greater in the refuge colonies than in the no-refuge colonies. These results demonstrate that the refuges delayed the evolution of resistance. Relative to a susceptible colony, final resistance ratios were 19 and 8 for the two no-refuge colonies compared to 6 and 5 for the refuge colonies. The mean realized heritability of resistance to B. thuringiensis subsp. aizawai was 0.046 for colonies without refuges, and -0.002 for colonies with refuges. Selection with B. thuringiensis subsp. aizawai decreased susceptibility to B. thuringiensis toxin Cry1Ab, but not to Cry1C or B. thuringiensis subsp. kurstaki. Although the ultimate test of refuges will occur in the field, the experimental evidence reported here confirms modelling results indicating that refuges can slow the evolution of insect resistance to B. thuringiensis.     

The role of colony morphology and substratum inclination in the success of Millepora alcicornis on shallow coral reefs

 Millepora species are conspicuous members of shallow coral reefs where they occupy a variety of substrata and produce morphologically complex skeletons. This study focuses on the roles of growth on vertical and horizontal surfaces and the production of encrusting bases and branches (a “sheet-tree” morphology) for the success of the Millepora alcicornis on coral reefs. The effects of inclination were investigated by comparing the size and growth rates of M. alcicornis on vertical and horizontal surfaces at 3–5 m depth, in St. John, US Virgin Islands. The consequences of morphological complexity were investigated by comparing polyp density, chlorophyll content and biomass between encrusting bases and branches; the role of branches in asexual reproduction was also quantified. Colonies on vertical surfaces had larger encrusting bases, longer perimeters and lower densities of branches compared to those on horizontal surfaces. Growth rates also varied significantly between surfaces, largely because colonies on horizontal surfaces shrank in area while those on vertical surfaces increased in area, albeit slowly. Branches were not specialized in comparison to encrusting bases in terms of the density of dactylozooids and gastrozooids, chlorophyll content and biomass, but they were effective asexual propagules. During one storm, 79% of the branches were removed from colonies of M. alcicornis, and 4% attached to the substratum to produce new colonies at a density of ≈0.5 colonies.m^-2. Anecdotal observations suggest that such storms rarely damaged encrusting bases on vertical surfaces, but often destroyed those on horizontal surfaces. Thus, the encrusting bases on vertical surfaces are likely to be large because of greater age rather than faster growth, while those on horizontal surfaces are likely to be small because they are relatively young and short lived. These findings suggest that the success of M. alcicornis is a result, in part, of the beneficial consequences of their “sheet-tree” morphology, that supports: (a) slow growth and resistance to wave damage of encrusting bases on vertical surfaces, and (b) the use of branches as asexual propagules. Accepted: 24 November 1998     

Seasonality of epiphytic development of the hydroid Obelia geniculata on cultivated Saccharina japonica (Laminariaceae, Phaeophyta) in Korea

The occurrence of encrusting colonies of the hydroid Obelia geniculata on farmed Saccharina japonica was examined between December 2007 and July 2008 at a Saccharina farm at Wando on the southwestern coast of Korea. The growth stages of S. japonica can be divided into two phases: an active growth phase from February to the end of May and a decay phase from June to July. There was a significant increase in the level of incrustation by colonies on fronds (measured as the percentage of fronds with encrusting colonies) between February and July (p?<?0.05). The encrusting colonies occurred first on the upper part of the frond in February and progressed to the basal part in July. The abundance of encrusting colonies in relation to the growth phase on farms over time was limited by the harvest of the seaweed crop at the end of the cultivation period in July. The stipes and holdfasts of fronds showed no signs of infestation at any time during the cultivation period. The extent of the infestation appeared to be related to a combination of factors. These could be reduced physiological activity and subsequent tissue aging that occurred simultaneously in the sporophytic life phase of Saccharina frond, and a rapid increase in reproduction and growth of O. geniculata coinciding with rising seawater temperature.