Regular and flexible modes of division and hystero-ontogenetic growth in the Silurian rugose coral Stauria favosa

New modules arise in colonial corals as the result of asexual reproduction. The Silurian rugosan Stauria favosa ordinarily exhibits cerioid coralla with a characteristic cross-shaped axial structure and a typical pattern of parricidal increase. Quadripartite increase at the sites of the four protosepta is most common, whereas cases of tripartite increase are rare. Parental protosepta are transformed into dividing walls, where the four protosepta first appear with a definite polarity in offset corallites. Daughter corallites inherit metasepta as metasepta, and catasepta as catasepta, within the same quadrants as those of the parent. Metasepta are inserted serially, following Kunth's rule, as is characteristic of rugosan protocorallites. As each daughter corallite derived immediately from the same parent is arranged with identical polarity, it grows equally and evenly both individually and as a group. Daughters thus form protosepta and metasepta under strict phylogenetic and developmental constraints. However, individual corallites grow and reproduce autonomously, by using all available skeleton and space of the parent. Although each module cannot modify essential modes of division, flexibility of the system was via changes in the density and arrangement of corallites, and regulating modes of growth, in tandem with adjacent corallites within the corallum. It is probable that regularity, due to constraints of several origins, as well as flexibility are typical of other rugosan colonies and played an important role in growth dynamics between corallites and corallum.     

Genetic control of septal numbers and the species problem in a fossil solitary scleractinian coral

Intraspecific morphological variations of a Pleistocene solitary scleractinian coral, Cylindrophyllia orientatis (Yabe & Eguchi), have been examined based on 792 specimens. The specimens are discoidal to short cylindrical in shape, with no significant change in their diameter during skeletal growth. Septal arrangements of the coralla are observed on upper and basal surfaces. Septal numbers do not change through the ontogeny of each corallum, even when the last cycle of septa is incomplete. Septal arrangements and numbers are controlled by intrinsic genetic factors. Heights of the coralla are controlled by environmental factors where they lived If growth rates are presumed to be constant, heights can be regarded as indicating age of specimens. Assuming that this is the case, the survivorship curve shows that this fossil population had a constant death rate. Two varieties exist in this population: one has 20–28 septa, the other 30–48 septa, showing a dimorphic feature. Scleractinian coral, intraspecific variation, population, septa, species problem ,     

Boring sponges,an increasing threat for coral reefs affected by bleaching events

Coral bleaching is a stress response of corals induced by a variety of factors, but these events have become more frequent and intense in response to recent climate‐change‐related temperature anomalies. We tested the hypothesis that coral reefs affected by bleaching events are currently heavily infested by boring sponges, which are playing a significant role in the destruction of their physical structure. Seventeen reefs that cover the entire distributional range of corals along the Mexican Pacific coast were studied between 2005/2006, and later between 2009/2010. Most of these coral reefs were previously impacted by bleaching events, which resulted in coral mortalities. Sponge abundance and species richness was used as an indicator of bioerosion, and coral cover was used to describe the present condition of coral reefs. Coral reefs are currently highly invaded (46% of the samples examined) by a very high diversity of boring sponges (20 species); being the coral reef framework the substrate most invaded (56%) followed by the rubbles (45%), and the living colonies (36%). The results also indicated that boring sponges are promoting the dislodgment of live colonies and large fragments from the framework. In summary, the eastern coral reefs affected by bleaching phenomena, mainly provoked by El Niño, present a high diversity and abundance of boring sponges, which are weakening the union of the colony with the reef framework and promoting their dislodgment. These phenomena will probably become even more intense and severe, as temperatures are projected to continue to rise under the scenarios for future climate change, which could place many eastern coral reefs beyond their survival threshold.     

Genetic variation and population structuring in two brooding coral species (<Emphasis Type="Italic">Siderastrea stellata</Emphasis> and <Emphasis Type="Italic">Siderastrea radians</Emphasis>) from Brazil

Siderastrea stellata and S. radians are scleractinian coral species that present a remarkable overlap of diagnostic characteristics and sympatric distribution. Moreover, both are viviparous with similar reproductive strategies and with a gregarious larval behavior. Samples of both species from the Brazilian coast were analyzed using 18 isozymic loci to quantify their genetic variability and populational structure. Results confirmed species identity, high intrapopulational variability and revealed moderate genetic structuring among all samples (S. stellata: F (ST) = 0.070; S. radians: F (ST) = 0.092). Based on genotypic diversity analysis, there was evidence that local recruitment may have a minor role in the populations (mean, G ( o ):G ( e )= 1.00 +/- 0.0003 SD for S. stellata and 0.99 +/- 0.0023 SD for S. radians). Deviations towards heterozygote deficiencies found in both Siderastrea species could be explained by the Wahlund effect, since there was evidence that populations might be composed of colonies of different ages. In S. radians it is also likely that there is some inbreeding occurring in the studied populations. Despite the brooding pattern and the gregarious larval behavior, our data suggest the occurrence of gene flow along the Brazilian coast. This is the first study on population genetics of Brazilian reef corals.     

Long distance dispersal and connectivity in amphi-Atlantic corals at regional and basin scales

Among Atlantic scleractinian corals, species diversity is highest in the Caribbean, but low diversity and high endemism are observed in various peripheral populations in central and eastern Atlantic islands and along the coasts of Brazil and West Africa. The degree of connectivity between these distantly separated populations is of interest because it provides insight into processes at both evolutionary and ecological time scales, such as speciation, recruitment dynamics and the persistence of coral populations. To assess connectivity in broadly distributed coral species of the Atlantic, DNA sequence data from two nuclear markers were obtained for six coral species spanning their distributional ranges. At basin-wide scales, significant differentiation was generally observed among populations in the Caribbean, Brazil and West Africa. Concordance of patterns in connectivity among co-distributed taxa indicates that extrinsic barriers, such as the Amazon freshwater plume or long stretches of open ocean, restrict dispersal of coral larvae from region to region. Within regions, dispersal ability appears to be influenced by aspects of reproduction and life history. Two broadcasting species, Siderastrea siderea and Montastraea cavernosa, were able to maintain gene flow among populations separated by as much as 1,200 km along the coast of Brazil. In contrast, brooding species, such as Favia gravida and Siderastrea radians, had more restricted gene flow along the Brazilian coast.     

Preservation of in situ reef framework in regions of low hurricane frequency: Pleistocene of Curaçao and Bonaire, southern Caribbean

Luxuriant fringing reefs along the southwestern shores of the Caribbean islands of Curaçao and Bonaire (12°N), located outside the most frequent hurricane tracks, are rarely affected by major storms. Consequently, reef growth and long-term preservation are potentially optimal and distinct from reefs experiencing greater hurricane frequency. Hurricane Lenny (November 1999) took an unusual west-to-east track, bisecting the Caribbean Basin north of these islands, but generated heavy waves (3-6  m) that severely damaged reefs along the normally leeward shores. Massive coral colonies >100 years old were toppled, but even at the most severely damaged sites, 82–85% of colonies remained in growth position. Late Pleistocene (125  ka) elevated reefs in the Lower Terrace of Curaçao record even higher proportions of corals in growth position (93%), possibly reflecting a low hurricane frequency during the Pleistocene highstand. In comparison, coeval Pleistocene reefs in regions that today experience a high hurricane frequency (Great Inagua Island and San Salvador, Bahamas) have lower proportions of corals preserved in growth position (79% and 38%, respectively). These results are consistent with the hypothesis that reefs in regions experiencing very low hurricane frequency, like the southern Caribbean, are more likely to be preserved with corals in primary growth position in comparison to regions with higher hurricane frequency.     

Short-term progressive early diagenesis in density bands of recent corals:Porites Colonies,Mauritius Island,Indian Ocean

Summary The growth history of some recentPorites colonies of Mauritius Island (Indian Ocean) was dated by sclerochronological methods. Couples of high-density and low-density bands represent the annual growth rate of the corals and allow the growth pattern of every year in the corallum to be counted. The growth and structure of the skeletons ofPorites solida andPorites lutea were investigated. Older parts of the aragonitic skeleton in these 10 to 20 year old corals show various secondary microstructures resulting from alterations and thickenings of the elements of the skeleton. The primary needle-like aragonite crystals are absent in older parts of the corallum and blocky aragonitic cements can occur. Pores and primary skeletal elements are overgrown by new microstructures. These microstructures are caused by secondary cementation and exhibit frontal zones (Stirnzonen), zigzag-like and pseudolamellar-structures. The lamellar structures can be compared with similar structures in the exoskeleton of some Rugosa. A very short early diagenesis within the recent corals is responsible for the thickening and alteration of skeletal elements. It occurs only 4 to 5 years after the formation of the skeleton and tends to increase in importance in older parts of the corallum. Nevertheless, there is no proof for any alteration of aragonite to calcite.     

Life-history strategies of a species of Catenipora (Tabulata; Upper Ordovician; southern Manitoba, Canada)

Detailed study of coralla by transverse serial sections permits the determination and evaluation of life-history strategies (survival and growth characteristics) in response to different physical environments, for Catenipora foerstei Nelson, 1963 from the Selkirk Member, Red River Formation, in Manitoba. We recognize various modes of corallite increase: one type of axial increase, four types of lateral increase, and agglutinated patches of corallites in association with normal, undamaged corallites; and one type of axial increase, one type of lateral increase, and temporary agglutinated patches from the recovery processes of corallites damaged by sediment or bioclast influx. In addition, the formation of new ranks by lateral increase is the most effective method for rapid growth of a corallum or for reconstructing part of a corallum damaged by physical disturbances. Fluctuations in the tabularial area of corallites occur in cycles over vertical intervals ranging from 3.20-7.90 mm. We consider each cycle to represent annual growth. Average annual growth of the three coralla ranges from 4.20-6.27 mm. According to correlations between annual growth cycles and other growth characteristics, a high frequency of offsetting is associated with rapid vertical growth. Specifically, annual growth is relatively high in association with episodes of sediment or bioclast influx, probably generated by storms. In some coralla, however, annual growth is highest in the cycle characterized by few new corallites or by extraordinarily high rates of offsetting by normal, undamaged corallites as well as damaged corallites. This suggests that vertical growth could also be affected by factors other than storm-related disturbance.     

An illustrated key to the genera and subgenera of the Recent azooxanthellate Scleractinia (Cnidaria,Anthozoa), with an attached glossary

The 120 presently recognized genera and seven subgenera of the azooxanthellate Scleractinia are keyed using gross morphological characters of the corallum. All genera are illustrated with calicular and side views of coralla. All termes used in the key are defined in an illustrated glossary. A table of all species-level keys, both comprehensive and faunistic, is provided covering the last 40 years.     

Distribution and Abundance of Fauna on Living Tissues of Two Brazilian Hermatypic Corals (Mussismilia hispida (Verril 1902) and Siderastrea stellata Verril, 1868)

This study describes the distribution and abundance patterns of the associate fauna on the living surface of the corals Siderastrea stellata Verril, 1868 and Mussismilia hispida (Verril 1902) using a non-destructive method, on the northern coast of Rio de Janeiro State. For each coral species, infestation density and proportions of infested colonies, colonies attached and unattached to the substrate were estimated. A total of 474 colonies of S. stellata and 452 colonies of M. hispida were examined. The barnacle Ceratoconcha floridana (Pilsbry, 1931) was the dominant coral associate found, followed by gall-crabs of the family Cryptochiridae Paulson, 1875 and the bivalve Lithophaga bisulcata (d’Orbigny, 1842). Both coral species presented similar patterns of infestation dominance. S. stellata colonies were more commonly infested and showed a greater mean infestation density of 0.62 ind/cm² at Armação dos Búzios, whereas M. hispida colonies had infestation densities of only 0.20 ind/cm². Infestation density does not appear to impact negatively on corals of Armação dos Búzios. A clear negative relationship between the number of associates in the coral colony and coral size was found. Evidently abundance and frequency of occurrence of associated fauna is highly related to coral community structure and composition and the results highlight the importance of local scale studies.     

Does Dark-Spot Syndrome Experimentally Transmit among Caribbean Corals?

Over the last half-century, coral diseases have contributed to the rapid decline of coral populations throughout the Caribbean region. Some coral diseases appear to be potentially infectious, yet little is known about their modes of transmission. This study experimentally tested whether dark-spot syndrome on Siderastrea siderea was directly or indirectly transmissible to neighboring coral colonies. We also tested whether open wounds were necessary to facilitate disease transmission. At the completion of the experiments, we sampled bacterial communities on diseased, exposed, and healthy coral colonies to determine whether bacterial pathogens had transmitted to the susceptible colonies. We saw no evidence of either direct or waterborne transmission of dark-spot syndrome, and corals that received lesions by direct contact with diseased tissue, healed and showed no signs of infection. There were no significant differences among bacterial communities on healthy, exposed, and diseased colonies, although nine individual ribotypes were significantly higher in diseased corals compared with healthy and exposed corals, indicating a lack of transmission. Although our experiments do not fully refute the possibility that dark-spot syndrome is infectious and transmissible, our results suggest that in situ macroscopic signs of dark-spot syndrome are not always contagious.     

The Association of Two Gall Crabs (Brachyura: Cryptochiridae) with the Reef-building Coral Siderastrea stellata Verrill, 1868

The family Cryptochiridae includes a small group of cryptic crabs that inhabit cavities in scleractinian corals. They are represented in Brazil by two species. Troglocarcinus corallicola, found from Maranhão to Bahia states and that occurs in many species of the coral families Mussidae, Faviidae, Caryophillidae and Siderastreidae. Opecarcinus hypostegus, reported from the state of Pernambuco, is mainly associated with Agaricia fragilis. The range of O. hypostegus is extended to Rio Grande do Norte and Alagoas states. Males are usually smaller than females, and couples may be eventually found sharing cavities. Ovigerous females and males of O. hypostegus are recorded for the first time inhabiting adjoining cavities on colonies of Siderastrea stellata, an endemic reef coral widespread along the Brazilian littoral.     

Regular mode of increase, and constrained but variable growth, in the Silurian rugose coral Stauria favosa

Stauria favosa exhibits a typical pattern of axial division (tripartite or quadripartite). The four protosepta first appear with a definite polarity in offset corallites, and metasepta are inserted serially, following Kunth's rule. There are no variations in essential modes of asexual reproduction. However, at the corallum periphery, up to four offset corallites appear to have been derived by discrete offsetting within the parent calices. Detailed observations of vertical morphological changes, however, indicate that this apparently unique mode of increase is derived from axial division at the sites of the four protosepta, as usual in this species. Just after the initiation of division, the contraction of polyps occurred in response to somewhat accidental, deteriorating habitat conditions. The combination of asexually divided corallites and rejuvenescence resulted in phaceloid daughter corallites, which subsequently underwent partial mortality at the corallum periphery. Stauria favosa always exhibits not only regular modes of asexual reproduction and septal insertion under the strict control of phylogenetic and developmental constraints, but also variable growth forms of corallites and coralla. This variability, though within limited options, is palaeoecologically significant for successfully coping with unpredictable habitats.     

Severity of the 1998 and 2005 bleaching events in Venezuela, southern Caribbean

This study describes the severity of the 2005 bleaching event at 15 reef sites across Venezuela and compares the 1998 and 2005 bleaching events at one of them. During August and September 2005, bleached corals were first observed on oceanic reefs rather than coastal reefs, affecting 1 to 4% of coral colonies in the community (3 reef sites, n = 736 colonies). At that time, however, no bleached corals were recorded along the eastern coast of Venezuela, an area of seasonal upwelling (3 reefs, n = 181 colonies). On coastal reefs, bleaching started in October but highest levels were reached in November 2005 and January 2006, when 16% of corals were affected among a wide range of taxa (e.g. scleractinians, octocorals, Millepora and zoanthids). In the Acropora habitats of Los Roques (an oceanic reef),no bleached was recorded in 2005 (four sites,n = 643 colonies). At Cayo Sombrero, a coastal reef site, bleaching was less severe in 1998 than in 2005 (9% of the coral colonies involving 2 species vs. 26% involving 23 species, respectively). Our results indicate that bleaching was more severe in 2005 than in 1998 on Venezuelan reefs; however, no mass mortality was observed in either of these two events.     

Short-term ecological consequences of a major oil spill on Panamanian subtidal reef corals

A major oil spill (8,000,000 liters; 50,000 barrels) occurred in Bahía Las Minas on the Caribbean coast of Panama in April 1986, and oil slicks from the refinery landfill and mangroves were still common there after 21/2 years. We studied short-term effects of the spill on common shallow subtidal reef corals, at the individual, population, and community levels. Numbers of corals, total coral cover, and species diversity based on cover decreased significantly with increased amounts of oiling. Cover of the large branching coral Acropora palmata decreased most. Frequency and size of recent injuries on massive corals increased with level of oiling, particularly for Siderastrea siderea. Growth of three massive species (Porites astreoides, Diploria strigosa, and Montastrea annularis, but not S. siderea) was less at oiled reefs in the year of the spill than during the 9 previous years. Subtidal coral reefs, particularly those along protected coasts, may suffer extensive damage from chronic exposure after major oil spills. Mailing Address from the USA: Smithsonian Tropical Research Institute, APO Miami 34002-0011, USA     

Coral mortality and interaction with algae in relation to sedimentation

The impact of sedimentation on coral–algal interactions was studied by monitoring tissue mortality and radial growth in two coral species, Colpophyllia natans and Siderastrea siderea, over a continuum of sediment input intensities. This study sets out to investigate (1) whether sedimentation can facilitate algal overgrowth of corals and (2) whether this was a significant cause of coral mortality. Over a 15-month period, 198 coral colonies were tagged and photographed at six sites along two replicate gradients of sediment input, spanning high inputs near river mouths to low inputs at exposed headlands. Photographs were taken so that they covered the interface between colonies and algae. Radial growth was measured along colony edges in contact with algae and unaffected by tissue loss from causes other than competition with algae. To establish whether algal overgrowth was a significant cause of coral mortality, tissue mortality on the colony surface area visible in the photographs was related to different causes, including sediment smothering, disease, and algal overgrowth. Radial growth became negative with increasing proximity to river mouths in C. natans and remained negative or close to zero throughout the gradients in S. siderea, overall suggesting that sedimentation can facilitate algal overgrowth on corals. However, the analysis of tissue mortality revealed that algal overgrowth was a relatively minor cause of tissue loss. In contrast, the most important cause of coral mortality in relation to sedimentation was from sediment smothering, probably during intense episodes of deposition associated with heavy rainfall. We conclude that sedimentation may lead to reef degradation by causing coral mortality through sediment smothering and burial, and then by suppressing the regrowth of surviving adult colonies through increased competition with algae.     

Highly stable symbioses among western Atlantic brooding corals

The reproductive mode of corals largely determines how zooxanthellae (Symbiodinium spp.) are acquired. Typically, broadcast spawning corals obtain symbionts from the surrounding environment, whereas most brooders transfer symbionts from maternal parent to offspring. Brooding corals are therefore predicted to harbor stable communities of Symbiodinium. This study documents the associations between Symbiodinium spp. and brooding corals in response to seasonal environmental fluctuations. Between March 2002 and December 2005, endosymbiont identity was determined seasonally from replicate colonies (n = 6) of three brooding species, Agaricia agaricites, Porites astreoides and Siderastrea radians, from shallow environments (1–4 m) of the Florida Keys and Bahamas. Symbionts were identified via denaturing gradient gel electrophoresis (DGGE) of the internal transcribed spacer 2 (ITS2) region. No change was detected in the Symbiodinium communities harbored within these brooding colonies. Additionally, no change in symbiosis was observed through a moderate bleaching event, thereby demonstrating that some bleached corals recover without changing symbionts.     

Holocene coral reef accretion in Hawaii: a function of wave exposure and sea level history

 In the high Hawaiian Islands, significant accretion due to coral reef growth is limited by wave exposure and sea level. Holocene coral growth and reef accretion was measured at four stations off Oahu, Hawaii, chosen along a gradient in wave energy from minimum to maximum exposures. The results show that coral growth of living colonies (linear extension) at optimal depths is comparable at all stations (7.7–10.1 mm/y), but significant reef accretion occurs only at wave sheltered stations. At wave sheltered stations in Hanauma Bay and Kaneohe Bay, rates of long term reef accretion are about 2.0 mm/y. At wave exposed stations, off Mamala Bay and Sunset Beach, reef accretion rates are virtually zero in both shallow (1 m) and deeper (optimal) depths (12 m). At wave sheltered stations, such as Kaneohe Bay and Hanauma Bay, Holocene reef accretion is on the order of 10–15 m thick. At wave exposed stations, Holocene accretion is represented by only a thin veneer of living corals resting on antecedent Pleistocene limestone foundations. Modern coral communities in wave exposed environments undergo constant turnover associated with mortality and recruitment or re-growth of fragmented colonies and are rarely thicker than a single living colony. Breakage, scour, and abrasion of living corals during high wave events appears to be the major source of mortality and ultimately limits accretion to wave sheltered environments. Depth is particularly important as a modulator of wave energy. The lack of coral reef accretion along shallow open ocean coastlines may explain the absence of mature barrier reefs in the high Hawaiian Islands. Accepted: 14 May 1998     

Community patterns on a submerged barrier reef at Barbados,West Indies

A quantitative survey of a submerged barrier reef was undertaken in Barbados, West Indies, over a two year period (1971–73). Photo-line transects were employed to obtain coverage data on corals and other benthic organisms. Light, sedimentation, currents, oxygen, temperature and salinity were also monitored. Results indicate corals cover about 30 per cent of the bottom with living colonies; another 7 per cent is contributed by other zoobenthos. The most abundant coral species are Montastrea annularis, M. cavernosa and Siderastrea siderea, each contributing between 4 and 5 per cent of bottom cover. Light is the only physical factor monitored that correlated significantly with biomass; sedimentation may have a secondary effect. Most of the barrier reef is composed of mixed coral associations forming a biologically accommodated community. Comparisons are made between the barrier reef in Barbados and deeper reefs in Jamaica and Curacao. Reefs are, in the main, similar but coral species and community structure differences do occur.     

Pedogenesis of late Quaternary deposits, northern Kyonggi Bay, Korea: Implications for relative sea-level change and regional stratigraphic correlation

Late Pleistocene and early Holocene paleosols are described from construction pits and boreholes in the northern Kyonggi Bay, west coast of Korea. Thin-section petrography, chemistry, clay mineralogy and geotechnical properties of the paleosols were examined to infer relative sea-level fluctuations during the late Quaternary. Relict laminae and gradational contacts with underlying tidal rhythmites confirm that the late Pleistocene paleosol is a pedogenically altered tidal deposit. Illuvial clay coatings, rootlets and enrichment of kaolinite and chemically stable oxides indicate that pedogenesis occurred on a well-drained, stable surface during the last glacial period. Redoximorphic features, such as drab-colored root traces and sphaerosiderite, however, suggest that waterlogged conditions developed temporarily due to a rise of the water table at the end of the last glacial period. As sea level rose, an early Holocene paleosol formed in freshwater bog deposits, burying the late Pleistocene paleosol. The early Holocene paleosol has abundant sphaerosiderites and organic material. The presence of weakly developed illuvial clay coatings, oxidized siderites and pellets, however, indicates oxidizing condition associated with water-table fall occurred during the early Holocene. Despite the similarity in the degree of chemical and clay mineralogical weathering, the late Pleistocene paleosol is distinguished from its early Holocene counterpart by thicker and more abundant illuvial clay coatings, more Fe₂O₃ and a greater degree of consolidation. Duration of pedogenesis might be a primary cause for the difference of the pedofeatures with climate playing a secondary role. The widespread presence of similar late Pleistocene paleosols highlights its significance as a key stratigraphic marker for regional correlations along the west coast of Korea.