Large dwellers of the Silurian Halysites biostrome: rhizosessile life strategies of cystiphyllid rugose corals from the Llandovery of Gotland

An exceptionally well‐preserved, unusual biostrome composed of the framebuilding cateniform tabulate coral Halysites catenularius (Linnaeus, 1767) bears an assemblage of the relatively large solitary cystiphyllid rugosan Cystiphyllum visbyense Wedekind, 1927. The corallites of solitary cystiphyllids are embedded within the ranks of the halysitid colonies, which developed on a soft, muddy substrate and in relatively turbid water. The cystiphyllid larvae successively settled mostly on the ranks of halysitid colonies and on colonies of the tiny phaceloid rugose coral Nanophyllum ramosum Johannessen, 1995, whereas calice‐in‐calice recruitment was not successful for these cystiphyllid corals. Further growth of C. visbyense was supported by rhizoid structures, which were most frequently developed on the cardinal (convex) side of the corallite. The process of formation of the rhizoid structures is here discussed and explained in detail, showing that they were formed by the extension of the basal ectodermal tissue of the polyp. The cystiphyllids, which settled on the walls of living corallites of halysitid colonies, used sweeper tentacles to kill the smaller polyps of the colony to maintain the space around them and expand. Hence, they ultimately used the halysitid colonies only as a hard substrate to stabilize their position on the soft muddy sediment.     

Modes of regeneration and adaptation to soft‐bottom substrates of the free‐living solitary scleractinian Deltocyathoides orientalis

Scleractinian corals adapt to various substrate conditions with a variety of growth morphologies and modes of life. The azooxanthellate solitary scleractinian Deltocyathoides orientalis exhibits slightly flattened, bowl‐shaped corallites. This study describes in detail the modes of skeletal regeneration after fragmentation in association with exquisitely adaptive strategies of the corals for life on soft substrates. Larger fragments of individuals retaining almost two‐thirds to five‐sixths of the original skeletal area inherit the densely dilated, lower central skeleton, so as to keep a stable life position on soft substrates and regenerate the lost parts promptly. Even highly fragmented individuals preserving less than 10% of the original skeleton still regenerate and repair. Fragmented individuals with almost one‐sixth to one‐third original skeleton actively maintain a posture with the oral disc upward using movements of remaining tentacles. Damaged and missing soft tissues are then efficiently regenerated to form a mouth and gastrovascular cavity near the new centre of the corallum. Every regenerated individual reuses skeleton and soft tissues, and is capable of burrowing before the completion of growth morphology. The mode of regeneration characteristic of D. orientalis is thus effective and adaptive for maintenance of a stable life position on soft substrates for this solitary scleractinian. As fragmentation in deeper‐water, soft‐bottom settings is likely due to predation rather than turbulence, the rapid corallum regeneration and burrowing strategy may both represent adaptive strategies for life on soft substrates and exploitation of new niches, such as an infaunal mode of life, in a predator‐rich environment.     

Halysitid tabulates: sponges in corals' clothing

Abundant pyritic pseudomorphs of monaxonic siliceous spicules (ophirhabds and ?heloclones) have been found entrapped in the calcareous skeleton of the halysitid tabulate Quepora ?agglomeratiformis (Whitfield) from late Ordovician limestones of Frobisher Bay, Baffin Island, Canada. The finding indicates a poriferan (choristid or sublithistid) affinity of halysitids, early Palaeozoic marine fossils related so far to corals. They probably derived from a monaxonic group of early demosponges that adapted during the Ordovician to Ca²⁺ stress conditions in epicontinental seas by excreting the excessive Ca²⁺ influx to their tissues as variously designed chains of basally secreted calcareous tubes.     

Species-specific growth responses of favositid corals to soft-bottom substrates

Gibson, Michael A. & Broadhead, Thomas W. 1989 07 15: Species-specific growth responses of favositid corals to soft-bottom substrates. Lethaia , Vol. 22, pp. 287–299. Oslo. ISSN 0024–1164.
Species of favositid corals from the Upper Silurian and Lower Devonian of Tennessee, USA, exhibit structural modifications related to corallum geometry, interfacial skeletal material, and biotic associations that enabled them to survive in terrigenous mud rich environments. Favosites conicus Hall (Lower Devonian) had a flat, holotheca-covered base and a radial pattern of colony growth, but apparently had a short life span and may not have survived beyond the first reproductive cycle (monocarpous). It was adapted for living between major episodes of terrigenous mud influx. F. foerstei (Lower Devonian) had a convex, pseudoholotheca-covered base and a modified axial pattern of colony growth. Its large size, in comparison to that of F. conicus , suggests a longer lived colony (polycarpous), in which continued upward and outward growth enabled it to survive episodic sediment influx. F. forbesi (Upper Silurian) exhibited radial growth to form either (1) a globose corallum that was symbiotic with the stalks of living crinoids permitting the colony to live entirely above the substrate, or (2) a Gorallum with a steeply convex, holotheca-covered base that represents a bottom-dwelling colony in which the rate of growth probably only slightly exceeded the rate of sediment accumulation. * Functional morphology, astogeny, paleoecology, Tabulata .     

Effect of colony size and surrounding substrate on corals experiencing a mild bleaching event on Heron Island reef flat (southern Great Barrier Reef,Australia)

In January–May 2006, Heron Island in the Great Barrier Reef experienced a mild bleaching event. The effect of colony size, morphology and surrounding substrate on the extent of bleaching was explored. In contrast with previous studies, colony size did not influence bleaching sensitivity, suggesting that there may be a threshold of light and temperature stress beyond which size plays a role. Also contrasting with previous studies, massive corals were more affected by bleaching than branching corals. Massive corals surrounded by sand were more affected than the ones surrounded by rubble or dead coral. It is hypothesized that light reflectance from sand increases stress levels experienced by the colonies. This effect is maximized in massive corals as opposed to branching corals that form dense thickets on Heron Island. These results emphasize the importance of the ecological dynamics of coral communities experiencing low, moderate and high levels of bleaching for the understanding of how coral communities may change under the stress of climate change.     

Allometry of resource capture in colonial cnidarians and constraints on modular growth

1. Indeterminacy in growth of colonial organisms, such as corals, is commonly attributed to their modular construction which frees the colony from the allometric constraints that limit the size of single modules. However, as a colony grows, there may be a decrease in resource availability to interior modules because of active depletion and/or passive deflection by modules on the exterior. The effects of 'self-shading' on resource capture in modular animals are modelled using a simple allometric growth function.
2. The model assumes that resource capture by a module scales as an exponent ( γ ) of colony size (i.e. number of modules). Data taken from the literature indicate that model values of γ for light and prey capture range from – 0·80 to – 1·16 for branching and encrusting corals. Module-specific rates of resource use (i.e. metabolism) are less affected by colony size. Therefore, as a colony grows, net resource state eventually reaches zero, making further growth unsustainable or determinate.
3. The model also predicts an inverse relationship between module size and colony size such as that observed in Caribbean corals. This negative correlation results from the additive effects of module size and colony size on the degree of self-shading.
4. Resource capture is affected by growth form and flow regime, and the interaction between them can account for some of the morphological variation in corals and other colonial suspension feeders.     

Self‐organization and emergent individuality of favositid corals adapted to live on soft substrates

Seilacher, A. & Thomas, R.D.K. 2011: Self‐organization and emergent individuality of favositid corals adapted to live on soft substrates. Lethaia, Vol. 45, pp. 2–13. Secondary soft‐bottom dwellers share the problem that their ancestors, attached to hard substrates, had lost their mobility. On soft substrates, only a limited number of alternative tricks are available to maintain sessile organisms in life position or to right them following disturbance. Consequently, convergent adaptations have emerged in unrelated members of this ecological guild. Those of favositid corals are of particular interest because, on account of their small polyps, they were always colonial. A comparative analysis of forms adopted by favositids on soft substrates shows that key elements of their adaptive paradigms could have been achieved by self‐organization. This arose without centralized control by means of inherited reaction norms of individual polyps to local environmental conditions. The unique spiral growth habit of Favosites turbinatus Billings and its emergent individuality at the colony‐level of organization are explained in these terms. ‘Suicidal’ lids entombed altruistic marginal polyps, forming a secondary epitheca as the growing colony settled into the sediment. These lids also record the size and spacing of soft tentacles in this species. □Coloniality, Devonian, Favositida, Favosites turbinatus, morphogenesis, self‐organization, Tabulata.     

ORDOVICIAN HALYSITID CORALS FROM NEW SOUTH WALES

Three halysitid coral species are described from the Ordovician of New South Wales. Quepora calamus sp.nov. and Quepora sp. are recorded from the Cargo Creek Limestone, and Halysites praecedens sp.nov. from the Bowan Park and Canomodine Limestones. The species of Halysites is the first to be described from the Ordovician. It occurs in beds correlated with the LTpper Eastonian (= Middle-Upper Caradoc or Barneveld–Eden). A review of the early history and distribution of halysitids is presented.
Sections of Ordovician and Silurian species of Halysites have shed new light on the nature of interstitial and peripheral budding in the genus.     

The effect of flow and morphology on boundary layers in the scleractinians Dichocoenia stokesii (Milne-Edwards and Haime) and Stephanocoenia michilini (Milne-Edwards and Haime)

Mass transfer characteristics of scleractinian corals are affected by their skeletal morphology and the concentration gradients that develop as a consequence of the interactions of their morphology and biomass with the overlying seawater. These interactions can have a profound effect on coral metabolism. In this study, boundary layer characteristics were compared between different size colonies of the corals Dichocoenia stokesii and Stephanocoenia michilini to determine the relative roles of colony size and corallite structures (i.e. surface roughness) in mass transfer. Colonies of both species were rounded in shape, but differed in small-scale roughness as measured by the elevation of corallites. Additionally, D. stokesii had a greater aspect ratio than S. michilini, and their colonies were slightly taller for a given diameter. Boundary layers were characterized by placing dead coral skeletons in a flow tank and estimating shear velocities (u(*)) at different flow speeds. The effects of flow speed, size, and roughness on shear velocities were estimated for two juvenile size classes (10-20 and 30-40 mm diameter) of each species that were exposed to unidirectional flow regimes (4 and 17 cm s(-1)). Shear velocities were significantly greater in high, compared to low flow, and there was a significant interaction between colony size and surface roughness; the interaction was caused by a difference in magnitude, rather than direction, of the effect of roughness and size on u(*). Thus, there was a greater degree of turbulence at high flow compared to low flow, regardless of roughness or size, and the greatest turbulence occurred over large colonies of D. stokesii at high flow. Together, these results suggest that boundary layers around small corals are heavily influenced by upstream roughness elements, and more strongly affected by flow regimes than skeletal features. The relationship between colony morphology (i.e. aspect ratio and, possibly, surface roughness) and boundary layer characteristics may be non-linear in small corals.     

Patterns of distribution of calcite crystals in soft corals sclerites

The gross morphology of soft coral surface sclerites has been studied for taxonomic purposes for over a century. In contrast, sclerites located deep in the core of colonies have not received attention. Some soft coral groups develop massive colonies, in these organisms tissue depth can limit light penetration and circulation of internal fluids affecting the physiology of coral tissues and their symbiotic algae; such conditions have the potential to create contrasting calcifying conditions. To test this idea, we analyzed the crystal structure of sclerites extracted from different colony regions in selected specimens of zooxanthellate and azooxanthellate soft corals with different colony morphologies, these were: Sarcophyton mililatensis, Sinularia capillosa, Sinularia flexibilis, Dendronephthya sp. and Ceeceenus levis. We found that the crystals that constitute polyp sclerites differ from those forming stalk sclerites. We also observed different crystals in sclerites located at various depths in the stalk including signs of sclerite breakdown in the stalk core region. These results indicate different modes of calcification within each colonial organism analyzed and illustrate the complexity of organisms usually regarded as repetitive morphological and functional units. Our study indicates that soft corals are ideal material to study natural gradients of calcification conditions. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

High-Throughput Method for Automated Colony and Cell Counting by Digital Image Analysis Based on Edge Detection

Counting cells and colonies is an integral part of high-throughput screens and quantitative cellular assays. Due to its subjective and time-intensive nature, manual counting has hindered the adoption of cellular assays such as tumor spheroid formation in high-throughput screens. The objective of this study was to develop an automated method for quick and reliable counting of cells and colonies from digital images. For this purpose, I developed an ImageJ macro Cell Colony Edge and a CellProfiler Pipeline Cell Colony Counting, and compared them to other open-source digital methods and manual counts. The ImageJ macro Cell Colony Edge is valuable in counting cells and colonies, and measuring their area, volume, morphology, and intensity. In this study, I demonstrate that Cell Colony Edge is superior to other open-source methods, in speed, accuracy and applicability to diverse cellular assays. It can fulfill the need to automate colony/cell counting in high-throughput screens, colony forming assays, and cellular assays.     

Improvements on colony morphology identification towards bacterial profiling

Colony morphology may be an indicator of phenotypic variation, this being an important adaptive process adopted by bacteria to overcome environmental stressors. Furthermore, alterations in colony traits may reflect increased virulence and antimicrobial resistance. Despite the potential relevance of using colony morphological traits, the influence of experimental conditions on colony morphogenesis has been scarcely studied in detail. This study aims to clearly and systematically demonstrate the impact of some variables, such as colony growth time, plate colony density, culture medium, planktonic or biofilm mode of growth and strain genetic background, on bacterial colony morphology features using two Pseudomonas aeruginosa strains. Results, based on 5-replicate experiments, demonstrated that all variables influenced colony morphogenesis and 18 different morphotypes were identified, showing different sizes, forms, colours, textures and margins. Colony growth time and composition of the medium were the variables that caused the highest impact on colony differentiation both derived from planktonic and biofilm cultures. Colony morphology characterization before 45 h of incubation was considered inadequate and TSA, a non-selective medium, provided more colony diversity in contrast to P. aeruginosa selective media. In conclusion, data obtained emphasized the need to perform comparisons between colony morphologies in equivalent experimental conditions to avoid misinterpretation of microbial diagnostics and biomedical studies. Since colony morphotyping showed to be a reliable method to evaluate phenotypic switching and also to infer about bacterial diversity in biofilms, these unambiguous comparisons between morphotypes may offer a quite valuable input to clinical diagnosis, aiding the decision-making towards the selection of the most suitable antibiotic and supportive treatments.     

Estimating surface area of sponges and gorgonians as indicators of habitat availability on Caribbean coral reefs

A rapid method to estimate the three-dimensional (3D) surface area (SA) of marine gorgonians and sponges from field measurements of colony height, diameter, and morphology was developed as an indicator of habitat availability for fish and invertebrates. Colony characteristics for sponges and gorgonians were compiled from field measurements, expert judgment, and taxonomic literature, and employed to generate 3D images using computer-aided design software. The images were used to test various statistical models and geometric surrogates that best estimated SA using only height and diameter measurements. A morphological classification system was devised using shapes and relative proportions of sponges and gorgonians which are commonly found in shallow waters (<25 m depth) of the Central Western Atlantic Ocean. Regression models (linear, quadratic, or cubic) were found to be more robust than geometric surrogates, exhibiting greater accuracy at range extremes. Statistical models explained over 90% of the variation in SA and forecast errors of less than 20%. The best models for estimating SA are presented for eight sponge and nine gorgonian morphologies. Application of these methods with existing estimators for stony corals SA can be used as an indicator of structural habitat availability, which is an important ecosystem service of coral reefs.     

Influence of different substrates on the evolution of morphology and life‐history traits of azooxanthellate solitary corals (Scleractinia: Flabellidae)

Sessile organisms are influenced considerably by their substrate conditions, and their adaptive strategies are key to understanding their morphologic evolution and traits of life history. The family Flabellidae (Cnidaria: Scleractinia) is composed of the representative azooxanthellate solitary corals that live on both soft and hard substrates using various adaptive strategies. We reconstructed the phylogenetic tree and ancestral character states of this family from the mitochondrial 16S and nuclear 28S ribosomal DNA sequences of ten flabellids aiming to infer the evolution of their adaptive strategies. The Javania lineage branched off first and adapted to hard substrates by using a tectura‐reinforced base. The extant free‐living flabellids, including Flabellum and Truncatoflabellum, invaded soft substrates and acquired the flabellate corallum morphology of their common ancestor, followed by a remarkable radiation with the exploitation of adaptive strategies, such as external soft tissue [e.g. Flabellum (Ulocyathus)], thecal edge spine, and transverse division (e.g. Placotrochus and Truncatoflabellum). Subsequently, the free‐living ancestors of two genera (Rhizotrochus and Monomyces) invaded hard substrates independently by exploiting distinct attachment apparatuses such as tube‐like and massive rootlets, respectively. In conclusion, flabellids developed various morphology and life‐history traits according to the differences in substrate conditions during the course of their evolution. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 184–192.     

Light-enhanced calcification and dark decalcification in isolates of the soft coral Cladiella sp. during tissue recovery

Light-enhanced calcification is a general characteristic of zooxanthellate corals, suggesting a link between calcification by the coral and photosynthesis by the zooxanthellae, but the relationship between zooxanthellae and coral hosts during this process has not been elucidated. We hypothesized that the effects of tissue injury on the coral fragments used in experiments studying calcification might obscure that link. To detect the effects of tissue injury on light-enhanced calcification, we measured calcification rates (sclerite formation) in the soft coral Cladiella sp. by the alkalinity anomaly method during a 36-day experiment following injury associated with coral fragmentation. In the 2 weeks after colony fragmentation, the calcification response did not show a relation with light intensity. The typical light-enhanced calcification pattern was not noticed until day 15 of tissue recovery. The calcification rate of this soft coral increased with light intensity and time of tissue recovery and was comparable to that of hard corals exposed to similar experimental conditions. However, Cladiella sp. decalcified in the dark. The diurnal calcification-decalcification cycles probably control sclerite size and shape.     

Coral bleaching: the winners and the losers

Sea surface temperatures were warmer throughout 1998 at Sesoko Island, Japan, than in the 10 preceding years. Temperatures peaked at 2.8 °C above average, resulting in extensive coral bleaching and subsequent coral mortality. Using random quadrat surveys, we quantitatively documented the coral community structure one year before and one year after the bleaching event. The 1998 bleaching event reduced coral species richness by 61% and reduced coral cover by 85%. Colony morphology affected bleaching vulnerability and subsequent coral mortality. Finely branched corals were most susceptible, while massive and encrusting colonies survived. Most heavily impacted were the branched Acropora and pocilloporid corals, some of which showed local extinction. We suggest two hypotheses whose synergistic effect may partially explain observed mortality patterns (i.e. preferential survival of thick-tissued species, and shape-dependent differences in colony mass-transfer efficiency). A community-structural shift occurred on Okinawan reefs, resulting in an increase in the relative abundance of massive and encrusting coral species.     

Influence of substrate type on periphyton biomass and nutrient state at contrasting high nutrient levels in a subtropical shallow lake

How substrate affects periphyton biomass and nutrient state at different, but high, nutrient levels was tested in three large enclosures in a hypereutrophic subtropical shallow lake. We compared periphytic characteristics (1) on three hard substrates (stone, bamboo, and wood) incubated for 2 weeks and 1 year, respectively, to investigate the existence of the influences of substrate type at hypereutrophic levels, and (2) on artificial plants with contrasting (parvopotamid-like and myriophyllid-like) soft substrate morphology. In general, periphytic biomass and nutrient state were sensitive to variations in nutrient level, incubation time, hard substrate type (except 2-week incubated) and substrate morphology, but to a varying extent. The periphyton nutrient content increased with increasing nutrient levels on most substrates. Long-time incubated substrates supported more periphytic biomass, had a higher nutrient content and autotrophic proportion, while the effect of nutrient level on nutrient content in the periphyton was independent of incubation time. The effects of hard substrate type on periphyton characteristics were much weaker than those of nutrient level. By contrast, the effects of soft substrate morphology on periphyton biomass and carbon: nutrient ratios surpassed those of nutrient level. Chlorophyll a, dry mass, and ash free dry mass were much higher on parvopotamid than on myriophyllid substrates. Our results show that periphyton biomass and nutrient state are influenced by both substrate and nutrient level even in hypereutrophic lakes, which might have cascading effects on the benthic food web.     

Consequences of fission in the coral <Emphasis Type="Italic">Siderastrea siderea</Emphasis>: growth rates of small colonies and clonal input to population structure

Colony age and size can be poorly related in scleractinian corals if colonies undergo fission to form smaller independent patches of living tissue (i.e., ramets), but the implications of this life-history characteristic are unclear. This study explored the ecological consequences of the potential discrepancy between size and age for a massive scleractinian, first by testing the effect of colony origin on the growth of small colonies, and second by quantifying the contribution of ramets to population structure. Using Siderastrea siderea in St. John (US Virgin Islands) as an experimental system, the analyses demonstrated that the growth of small colonies derived from sexual reproduction was 2.5-fold greater than that of small ramets which were estimated to be ≈100 years older based on the age of the parent colonies from which they split. Although fission can generate discrete colonies, which in the case of the study reef accounted for 42% of all colonies, it may depress colony success and reef accretion through lowered colony growth rates.     

Reproduction and life strategies in the Paleozoic tabulate coral Paleofavosites capax (Billings)

Asexual and sexual modes of colony formation in a tabulate coral Paleofavosites capax are recognized from the early Silurian Gun River Formation of Anticosti Island. Québee. Colonies produced by asexual fragmentation comprise monospecific 'clumped populations'. They are characterized by circular and concave bases, and lack a protocorallite origin of colony growth. Sexually produced colonies, where in situ , are always dispersed and characterized by conical bases with a definite protocorallite point origin of colony growth. Asexual colony formation by fragmentation in P. capax appears to have been an adaptation to a habitat of muddy substrates. Sexual reproduction in this species probably played a minor role but was necessary for the maintenance of gene diversity and long-distance dispersal. A comparison of corallite size distributions between populations demonstrates that intrapopulation variation in the 'dispersed population' and the conical colonies in 'transported populations' of P. capax is significantly larger than the variation in the 'clumped populations'. It is suggested that this difference reflects the two modes of reproduction. The above observations are significant to systematic studies because they show that estimates of species morphologic parameters can he seriously biased even when based on a relatively large sample size from a well-defined population if that population is largely a result of asexual colony formation.     

The relationship between corallite morphology and colony shape in some massive reef-corals

Multivariate statistical tests are performed on three coral species to determine if a relationship exists between colony shape and corallite morphology. Hemispherical to branched colony shapes are emphasized. The material consists of specimens of Montastraea annularis and M. cavernosa from modern reef habitats in Jamaica. Colonies of the more branched M. limbata from the Neogene of the Dominican Republic are analyzed for comparison. The characters studied consist largely of linear measurements of colony dimensions and corallite structures in transverse thin-sections. The results show that different complexes of corallite characters vary within each species. With the possible exception of a complex describing theca thickness in M. annularis, none of these corallite complexes is related to variation in colony shape within species. Colony shape variation within species is related to upward colony growth rate. These results indicate that corallite morphology and colony shape can be represented by two different vectors of variation within species. Comparisons of intraspecific with interspecific patterns suggest that a strong genetic component explains colony shape variation in corals.