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.     

Morphological plasticity in scleractinian corals

When describing coral shape and form the term phenotypic plasticity, i.e. environment-induced changes in morphology, is often used synonymously with intraspecific variation. Variation, however, may simply be due to genetic differentiation (polymorphism). Of the 1314 extant scleractinian coral species, less than 20 have been tested for plastic responses. Morphological plasticity has important implications for coral identification, as skeletal features used in coral systematics are directly affected by environment. Furthermore, plastic changes can indicate how corals acclimatise to environmental change. The studies that have examined phenotypic plasticity in corals experimentally can be divided into two groups, i.e. 'non-clonal'—those that have transplanted whole colonies or fragments of colonies (but not treated the fragments as clones) to new environments, and 'clonal'—those that have transplanted colony fragments and used them as clone-mates. The use of clone-mates is preferable as it facilitates the identification of among-genotype variation for plasticity. The heterogeneous nature of the reef environment makes identifying the parameters that affect coral morphology difficult in the field, but there are also many problems conducting suitable aquarium experiments. Nevertheless, evidence to date suggests light and water movement are the most important variables inducing change. As these factors are known to be axiomatic to coral growth, it is possible that associated plastic changes in corals are adaptive; however, this hypothesis is yet to be tested rigorously.     

Coral reef restoration in the Eastern Tropical Pacific: feasibility of the coral nursery approach

Due to the worldwide degradation of coral reefs, the active restoration of these ecosystems has received considerable attention in recent decades. This study investigated (1) the feasibility of using coral nurseries for restoration projects, (2) the minimum size required for a Pocillopora damicornis (Pocilloporidae) coral fragment to survive and grow in a nursery, and (3) the optimal transplant size of a fragment when transplanted to a degraded reef at Gorgona Island (Colombian Pacific). For this investigation, 230 fragments were transplanted directly to El Remanso reef, and another 150 fragments were maintained in in situ nurseries. Every 2 months, the length, weight, and survival of the fragments were recorded. After growing for 134 days in the nurseries, the 52 surviving fragments were transplanted to El Remanso reef, and after 5 months, the same variables were measured. Among the nursery‐reared fragments, the largest (4 to <8 cm) had the highest survival and growth rates, whereas among the directly transplanted fragments, the smallest fragments (<2 cm) had the highest survival and growth rates. However, the nursery‐reared fragments acquired greater structural complexity (arborescent morphology), and they were all alive 156 days after transplantation and presented a maximum linear growth rate of over 2 cm, which was higher than that of the directly transplanted fragments. Apparently, the arborescent morphology acquired during the nursery period provides advantages to the colonies that favor greater success when transplanted. Therefore, nursery‐reared fragments of P. damicornis between 2 and 4 cm are the most appropriate for use in restoration projects.     

Transplantation and cultivation of fragments of coral colonies of various scleractinian species on a reef in Vietnam

Six coral species of the genus Acropora and two species of the genus Porites were studied during experiments on cultivation of reef-building scleractinian corals. The research has established species-specific factors and others affecting regeneration of fragments and growth of new colonies in these coral species. The accretion of donor fragments and new branches averaged from 40 to 160 mm per year, depending on the coral species, colony size, and season of transplantation. An average monthly accretion of medium and larger transplants and growth of new branches were 1.2–1.3 times higher at spring cultivation than at autumn transplanting. When transplanted, coral fragments of medium and larger sizes survived well and showed higher growth rates in all species studied. These transplants developed the highest number of new branches, and their buds and formed the largest colonies. Prolongation of the cultivation time from 1 to 1.5 years caused a 1.2–1.4 fold accretion of transplants.     

Effects of frequent fish predation on corals in Hawaii

The abundance of lesions from fish bites on corals was quantified at nine shallow reefs in the main Hawaiian Islands. There were on average 117 bite scars m⁻² on Pocillopora meandrina tissue from the barred filefish Cantherhines dumerilii, 69 bites m⁻² on Porites compressa tissue, and 4 bites m⁻² on Porites lobata tissue from the spotted puffer Arothron meleagris. Across sites, the frequency of A. meleagris bites on P. compressa per unit area of living coral cover declined exponentially with increasing coral cover. P. compressa nubbins in two size classes (1–2 cm and 4–5 cm) were transplanted onto six study reefs. Nubbins in the small size class were entirely removed by bites from A. meleagris, while nubbins ≥4 cm were only partially consumed, leaving them able to recover. At sites with abundant P. compressa, predation had little effect on transplanted nubbins; at sites where P. compressa comprised less than 5% of living cover, all nubbins were preyed upon. A. meleagris bite lesions on P. compressa were monitored through time and fully recovered in 42 ± 4 days. A model of the risk of over-predation (a second predation event before the first is healed) decreased exponentially with increasing coral cover and increased linearly with increasing lesion healing time. The increased risk of over-predation at low coral cover could indicate an Allee effect limiting the recovery of coral populations if coral cover is substantially reduced by natural or anthropogenic disturbances.     

Contrasting Light Spectra Constrain the Macro and Microstructures of Scleractinian Corals

The morphological plasticity of scleractinian corals can be influenced by numerous factors in their natural environment. However, it is difficult to identify in situ the relative influence of a single biotic or abiotic factor, due to potential interactions between them. Light is considered as a major factor affecting coral skeleton morphology, due to their symbiotic relation with photosynthetic zooxanthellae. Nonetheless, most studies addressing the importance of light on coral morphological plasticity have focused on photosynthetically active radiation (PAR) intensity, with the effect of light spectra remaining largely unknown. The present study evaluated how different light spectra affect the skeleton macro- and microstructures in two coral species (Acropora formosa sensu Veron (2000) and Stylophora pistillata) maintained under controlled laboratory conditions. We tested the effect of three light treatments with the same PAR but with a distinct spectral emission: 1) T5 fluorescent lamps with blue emission; 2) Light Emitting Diodes (LED) with predominantly blue emission; and 3) Light Emitting Plasma (LEP) with full spectra emission. To exclude potential bias generated by genetic variability, the experiment was performed with clonal fragments for both species. After 6 months of experiment, it was possible to detect in coral fragments of both species exposed to different light spectra significant differences in morphometry (e.g., distance among corallites, corallite diameter, and theca thickness), as well as in the organization of their skeleton microstructure. The variability found in the skeleton macro- and microstructures of clonal organisms points to the potential pitfalls associated with the exclusive use of morphometry on coral taxonomy. Moreover, the identification of a single factor influencing the morphology of coral skeletons is relevant for coral aquaculture and can allow the optimization of reef restoration efforts.     

Simultaneous extension of both basic microstructural components in scleractinian coral skeleton during night and daytime,visualized by in situ 86Sr pulse labeling

Using in situ (12 h) pulse-labeling of scleractinian coral aragonitic skeleton with stable ⁸⁶Sr isotope, the diel pattern of skeletal extension was investigated in the massive Porites lobata species, grown at 5 m depth in the Gulf of Eilat. Several microstructural aspects of coral biomineralization were elucidated, among which the most significant is simultaneous extension of the two basic microstructural components Rapid Accretion Deposits (RAD; also called Centers of Calcification) and Thickening Deposits (TD; also called fibers), both at night and during daytime. Increased thickness of the ⁸⁶Sr-labeled growth-front in the RADs compared to the adjacent TDs revealed that in this species RADs extend on average twice as fast as TDs. At the level of the individual corallite, skeletal extension is spatially highly heterogeneous, with sporadic slowing or cessation depending on growth directions and skeletal structure morphology. Daytime photosynthesis by symbiotic dinoflagellates is widely acknowledged to substantially increase calcification rates at the colony and the corallite level in reef-building corals. However, in our study, the average night-time extension rate (visualized in three successive 12 h pulses) was similar to the average daytime extension (visualized in the initial 12 h pulse), in all growth directions and skeletal structures. This research provides a platform for further investigations into the temporal calibration of coral skeletal extension via cyclic growth increment deposition, which is a hallmark of coral biomineralization.     

Skeletal response of <Emphasis Type="Italic">Lophelia pertusa</Emphasis> (Scleractinia) to bioeroding sponge infestation visualised with micro-computed tomography

The skeleton morphology of the azooxanthellate cold-water coral Lophelia pertusa can be strongly influenced by invasive boring sponges that infest corallites in the still living part of the colony. Atypically swollen corallites of live Lophelia pertusa from the Galway Mound (Belgica Carbonate Mound Province, Porcupine Seabight, NE Atlantic), heavily excavated by boring organisms, have been examined with a wide range of non-destructive and destructive methods: micro-computed tomography, macro- and microscopic observations of the outer coral skeleton, longitudinal and transversal thin sections and SEM analyses of coral skeleton casts. As a result, three excavating sponge species have been distinguished within the coral skeleton: Alectona millari, Spiroxya heteroclita and Aka infesta. Furthermore, four main coral/sponge growth stages have been recognised: (1) cylindrical juvenile corallite/no sponge cavities; (2) flared juvenile corallite/linear sponge cavities (if present); (3) slightly swollen adult corallites/chambered oval sponge cavities; (4) very swollen adult corallites/widespread cavities. The inferred correlation between corallite morphology and boring sponge infestation has been detected in micro-computed tomography (micro-CT) images and confirmed in sponge trace casts and peculiar features of coral skeleton microstructure. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorised users.     

Species composition and morphologic variation of Porites in the Gulf of California

Morphometric analysis of corallite calices confirmed that from the late Miocene to the Recent, four species of Porites have inhabited the Gulf of California: the extinct Porites carrizensis, the locally extirpated Porites lobata and the extant Porites sverdrupi and Porites panamensis. Furthermore, large-scale spatial and temporal phenotypic plasticity was observed in the dominant species P. panamensis. Canonical discriminant analysis and ANOVA demonstrated that the calice structures of P. panamensis experienced size reduction between the late Pleistocene and Recent. Similarly, PERMANOVA, regression and correlation analyses demonstrated that across the 800 km north to south in the gulf, P. panamensis populations displayed a similar reduction in calice structures. Based on correlation analysis with environmental data, these large spatial changes are likely related to changes in nutrient concentration and sea surface temperature. As such, the large-scale spatial and temporal phenotypic variation recorded in populations of P. panamensis in the Gulf of California is likely related to optimization of corallite performance (energy acquisition) within various environmental scenarios. These findings may have relevance to modern conservation efforts within this ecological dominant genus.     

Phenotypic plasticity in sex allocation for a simultaneously hermaphroditic coral reef fish

Phenotypic plasticity can facilitate reproductive strategies that maximize mating success in variable environments and lead to differences in sex allocation among populations. For simultaneous hermaphrodites with sperm competition, including Serranus tortugarum a small coral reef fish, proportional male allocation (testis in total gonad) is often greater where local density or mating group size is higher. We tested whether S. tortugarum reduced male allocation when transplanted from a higher density site to a lower density site. After 4 months, transplants mirrored the sex-allocation patterns of the resident population on their new reef. Transplants had significantly lower male allocation than representatives from their source population, largely as a result of reduced testis mass relative to body size.     

Effects of fish predation and seaweed competition on the survival and growth of corals

On Caribbean coral reefs, high rates of grazing by herbivorous fishes are thought to benefit corals because fishes consume competing seaweeds. We conducted field experiments in the Florida Keys, USA, to examine the effects of grazing fishes on coral/seaweed competition. Initially, fragments of Porites divaracata from an inshore habitat were transplanted into full-cage, half-cage, and no-cage treatments on a fore-reef. Within 48 h, 56% of the unprotected corals in half-cage and no-cage treatments (62 of 111) were completely consumed. Stoplight parrotfish (Sparisoma viride) were the major coral predators, with redband parrotfish (S. aurofrenatum) also commonly attacking this coral. Next, we transplanted fragments of P. porites collected from the fore-reef habitat where our caging experiments were being conducted into the three cage treatments, half in the presence of transplanted seaweeds, and half onto initially clean substrates. The corals were allowed to grow in these conditions, with concurrent development of competing seaweeds, for 14 weeks. Although seaweed cover and biomass were both significantly greater in the full-cage treatment, coral growth did not differ significantly between cage treatments even though corals placed with pre-planted seaweeds grew significantly less than corals placed on initially clean substrate. This surprising result occurred because parrotfishes not only grazed algae from accessible treatments, but also fed directly on our coral transplants. Parrotfish feeding scars were significantly more abundant on P. porites from the half and no-cage treatments than on corals in the full cages. On this Florida reef, direct fish predation on some coral species (P. divaracata) can exclude them from fore-reef areas, as has previously been shown for certain seaweeds and sponges. For other corals that live on the fore-reef (P. porites), the benefits of fishes removing seaweeds can be counterbalanced by the detrimental effects of fishes directly consuming corals. Received: 31 May 1997 / Accepted: 2 September 1997     

Coral Transplantation: Regeneration and Growth of Acropora Fragments in a Nursery

Abstract Coral reef degradation has been widely reported for the past 20 years. Because the recovery rate is usually low, various methods of restoration have been explored in different regions of the world. Among the effective and commonly used methods to restore coral communities is the transplantation of coral colonies or fragments. In this investigation fragments of Acropora pulchra were used in a semiprotected nursery in southern Taiwan between 1996 and 1998 to test, in situ, the possible effects of different factors on the generation of new branches and the initial skeletal extension rates of transplants. The variables under study here were the origin and length of the fragments, their new orientation, presence of tissue injury, and position in the fragment. All these factors were found to make a difference in either one or both aspects of coral growth (i.e., branching frequency and skeletal extension rate). These two factors clearly determine the success rate of a small fragment developing into a large colony that has a much higher probability to survive and grow on its own. It is now obvious that the efficiency of coral generation through fragment culture can be enhanced if the variables examined here are taken into consideration. Once coral colonies are formed, they can be fragmented again to generate more corals or can be transplanted to a suitable site.     

In situ nurseries enhance coral transplant growth in sedimented waters

In situ nurseries have been a crucial part of coral reef restoration initiatives for the past two decades. However, the advantages over direct transplantation in sedimented waters has yet to be examined. In the present study, we showed that Pachyseris speciosa and Pocillopora damicornis fragments reared in in situ nurseries (NR) in Singapore’s sedimented waters grew significantly faster (by three to five times) than those which were directly transplanted (DT) onto the substrates. The increased growth rate during the nursery phase augmented the size of NR transplants, and had a flow-on effect on their performance during the post-transplantation phase. Overall, the maximum diameter of the NR transplants was 1.8–2.7 times larger than DT transplants after 11 months. The growth enhancement of the nursery-reared transplants improved the cost-effectiveness of our restoration effort: the estimated cost per centimetre growth of NR transplants was one-fifth of the DT corals despite the additional costs incurred to construct the nurseries. These results highlight that coral nurseries are beneficial to reef restoration in chronically sedimented waters.     

Mediation of growth by conspecific neighbors and the effect of site in transplanted fragments of the coral Porites attenuata Nemenzo in the central Philippines

To investigate facilitative and competitive effects of conspecific neighbors on coral growth, fragments of Porites attenuata were reciprocally transplanted between two reefs. Transplants were interspersed with dead coral fragments or live conspecifics and monitored for growth. Over 13 months, transplant performance differed between sites. Those from degraded Bais Bay grew significantly slower in all treatments compared with those from more pristine Apo Island, and Bais fragments branched less (Bais: 1.5ǂ.16/transplant; Apo: 8.1ǂ.66/transplant). Treatment effects were observed in Apo; fragments with live conspecifics branched less but grew taller, adding significantly more surface area than those with dead neighbors. Clonal fragments differed significantly in their responses, indicating environmentally induced effects. These interactions affected growth and morphology prior to physical contact between fragments. This illustrates a benefit of manipulating the biotic environment of early transplants: dense stocking in ocean nurseries could stimulate early growth, and subsequent spacing would allow lateral branching and reduce competition.     

Size matters when it comes to the survival of transplanted yellow gorgonian fragments

The yellow gorgonian Eunicella cavolini is a Mediterranean endemic coral. Mediterranean populations underwent sharp decreases in the last decades due to climatic anomalies. In the meanwhile, populations in the Sea of Marmara thrive under conditions exempted from significant thermal variations thanks to the peculiar oceanography of the basin. However, they are vulnerable to anthropogenic activities, such as fishing, coastal constructions, dredging and landfilling. Major infrastructure projects at two remote islands, Yassıada and Sivriada, put healthy gorgonian communities at risk. We observed many dead colonies when Yassıada was constructed and before operations commenced on Sivriada, we transplanted gorgonian fragments 13 km away to a site at lower risk from construction. The transplantation site already hosts yellow gorgonian as sparse colonies; but also, it sits within an important fishing area, and entangled nets over rocks are common. Therefore, together with a local NGO, we initiated conservation efforts simultaneously with the transplantation works. 300 gorgonian fragments in the 4.5–15.4 cm size range were transplanted in two years (2017 – 2019). At six months, the survival of the colonies was significantly size dependent. Fragments of 10 – 15.4 cm initial sizes had the highest survival rate (up to 90 %), while small fragments yielded substantially lower survival. Despite severe impacts from fishing nets and mucilaginous entanglement, 50 % of fragments larger than 10 cm survived after 4.5 years of plantation. Besides, several recruits were observed among 4.5 years old transplants. Therefore, when restoration by raw transplantation actions is planned for gorgonians with thin scleraxis, a minimum fragment size of 10 cm is recommended. The study was a notable example of science/NGO collaboration with substantial local conservation success by using extensive media coverage of conservation efforts as leverage to declare the recipient area as a Marine Protected Area.     

Changes in scleractinian coral <Emphasis Type="Italic">Seriatopora hystrix</Emphasis> morphology and its endocellular <Emphasis Type="Italic">Symbiodinium</Emphasis> characteristics along a bathymetric gradient from shallow to mesophotic reef

The algae living endosymbiotically within coral are thought to increase algal pigmentation with increasing depth to capture the diminishing light. Here, we follow distribution of the hermatypic coral Seriatopora hystrix along a 60-m bathymetric gradient in the Gulf of Eilat, Red Sea, to study coral ecophysiology and response to light regimes. Combining work on coral morphology, pigment content and genotyping of the photosymbiont, we found that total chlorophyll concentration per zooxanthellae cell and the dark- and light-acclimated quantum yield of photosystem II did not vary significantly along the 60-m gradient. However, the chlorophyll a/c ratio increased with depth. This suggests that the symbiotic algae in S. hystrix possess a mechanism for acclimatization or adaptation that differs from previously described pathways. The accepted photoacclimatory process involves an increase in chlorophyll content per alga as light intensity decreases. Based on corallite and branch morphology, this research suggests that S. hystrix has two depth-dependent ecophenotypes. Above 10 m depth, S. hystrix exhibits sturdier colony configurations with thick branches, while below 30 m depth, colonies are characterized by thin branches and the presence of a larger polyp area. Between 10 and 30 m depth, both ecophenotypes are present, suggesting that corallite morphology may act as another axis of photoacclimation with depth.     

Reciprocal transplantation of the heterotrophic coral Tubastraea coccinea (Scleractinia: Dendrophylliidae) between distinct habitats did not alter its venom toxin composition

Tubastraea coccinea is an azooxanthellate coral species recorded in the Indian and Atlantic oceans and is presently widespread in the southwestern Atlantic with an alien status for Brazil. T. coccinea outcompete other native coral species by using a varied repertoire of biological traits. For example, T. coccinea has evolved potent venom capable of immobilizing and digesting zooplankton prey. Diversification and modification of venom toxins can provide potential adaptive benefits to individual fitness, yet acquired alteration of venom composition in cnidarians is poorly understood as the adaptive flexibility affecting toxin composition in these ancient lineages has been largely ignored. We used quantitative high‐throughput proteomics to detect changes in toxin expression in clonal fragments of specimens collected and interchanged from two environmentally distinct and geographically separate study sites. Unexpectedly, despite global changes in protein expression, there were no changes in the composition and abundance of toxins from coral fragments recovered from either site, and following clonal transplantation between sites. There were also no apparent changes to the cnidome (cnidae) and gross skeletal or soft tissue morphologies of the specimens. These results suggest that the conserved toxin complexity of T. coccinea co‐evolved with innovation of the venom delivery system, and its morphological development and phenotypic expression are not modulated by habitat pressures over short periods of time. The adaptive response of the venom trait to specific predatory regimes, however, necessitates further consideration.     

Grazing dynamics on a Caribbean reef-building coral

Corals in certain Caribbean coral reef habitats are constantly grazed-on due to the territorial marking behavior of the stoplight parrotfish Sparisoma viride. We studied the grazing dynamics on the Caribbean reef-building coral Montastraea annularis. We transplanted colonies to algae-dominated reefs (Rosario Islands, Cartagena, Colombia), where they encountered higher grazing pressure. We counted grazed polyps every month throughout a year. Over the course of a year, 4,101 different grazed polyps were counted on lobe-like M. annularis transplants ( n =23). Grazing was evaluated on a monthly basis as the probabilities of all the possible transitions among four grazing categories (0%, >0–1%, >1–5%, >5% grazed tissue), uncovering a dynamic process. Higher transition probabilities were always between 0 and 1% (coral tissue grazed) grazing states, indicating that grazing did not usually exceed 1% per coral per month. The probability of remaining uninjured in a month was 0.19, 0.17 of a change from 0–1% grazing state, 0.31 of remaining at 1%, and of full recovery from 1% grazing was 0.16. More than one month was usually required for complete recovery ( P<<1) probably due to both steady grazing pressure and slow regeneration rates. Since the marking behavior of the parrotfish was not as common on other zones of the reef no comparison on the grazing among environments was possible. In spite of this, it is possible to have stable transplanted populations of corals such as M. annularis on algae-dominated Caribbean reef environments due to their tolerance to the natural grazing pressure.Communicated by: K. S. Sealey     

Seasonal growth bands in Famennian rugose coral Scruttonia kunthi and their environmental significance

Large colonies of rugose coral Scruttonia kunthi occurring in the upper Famennian of Sudetes (southern Poland) reveal distinct growth banding in their skeletons. They were investigated for internal structural characteristics and stable isotopic composition. The skeletal tissue consists of alternating light and dark bands which differ in thickness, density and morphology of structural elements, and in occurrence of corallite contraction and rejuvenescense. Darker parts with densely arranged thick skeletal elements are thin in comparison to lighter parts. In addition, they include frequently offsets and contraction of corallites. A couplet of dense and less dense bands is interpreted to represent most probably an annual cycle. The calculated growth rate for Scruttonia kunthi varied from 6 mm/yr to 12 mm/yr. Growth-band formation was influenced environmentally. Oxygen isotopic data provide an evidence that high-density bands were formed in the season of higher environmental stress, with relatively warmer temperatures and higher sedimentation rates. Carbon isotopic signatures are very uniform, and thus enigmatic. They indicate that at least growth rate of the skeleton and seawater temperature had no influence on the coral δ¹³C.