Influence of light on algal symbionts of the deep water coral Leptoseris fragilis

Photoadaptations of zooxanthellae living within the deep water coral Leptoseris fragilis taken from the Gulf of Aqaba (Red Sea) were studied. Specimens-collected in summer 1988 between 110 and 120 m depth —were transplanted to 70 and 160 m. At each depth individuals were exposed in their natural growth position (oral side facing the surface) or in a reverse growth position (oral side facing the bottom). After 1 yr of exposure the corals were collected and the zooxanthellae were isolated. As a function of the availability of light with depth and growth position several algal parameters showed changes which are related to photoadaptations. The relatively low density of zooxanthellae of 0.15x10⁶ cellsxcm^-2 at a natural growth depth of 116 m decreased to 0.0034x10⁶ cellsxcm^-2 (2%) at 160 m in specimens growing with a natural orientation. In corals with a downward-facing oral surface at the same depth (160 m) only degenerated algae could be observed. With respect to depth dependence the volume of the algae decreased from 728 m³ at 116 m to 406 m³ at a depth of 160 m and the content of pigments increased. The augmentation of peridinin per cell was low (two times at 160 m compared to 116 m). Chlorophyll a and in particular chlorophyll c ₂ concentrations per cell were enhanced. Compared to natural amounts at 116 m, chl a was five times and chl c ₂ eight times higher at 160 m. At all depths the chl c ₂ content per cell was higher than for chl a. The formation of chl a/chl c ₂ complexes as light harvestor is discussed. Light harvesting, with chl c ₂ prevailing may be explained as a special type of chromatic adaptation of L. fragilis in a double sense: (1) in the habitat light short wavelengths predominate. This light can be directly absorbed with pigments such as chl a and chl c ₂. (2) Host pigments absorb visible violet light and transform these wavelengths, less suitable for photosynthesis, into longer ones by means of autofluorescence. The emitted longer wavelengths fit the absorption maxima of the algal pigments. Thus the host supports photosynthesis of his symbionts. Corals exposed at 160 m depth with a downward facing oral surface were alive after 1 yr and the host wavelength transforming pigment system was still present, but zooxanthellae were absent or degenerated. The light field at 160 m seems therefore to be critical: the combined photoadaptations of host and symbionts, allowing photosynthesis under barren light conditions, seem to be exhausted. In L. fragilis the photoadaptive strategies of host and symbionts cooperate harmoniously. In addition, the adaptations are interlocked with the particular light situation of the habitat with respect to light quantity and quality. The cooperation of physical and organismic parameters examplifies how evolution and, in particular, coevolution has led to optimal fitness.     

Phylogeny of the order Zoantharia (Anthozoa,Hexacorallia) based on the mitochondrial ribosomal genes

Zoantharia (or Zoanthidea) is the third largest order of Hexacorallia, characterised by two rows of tentacles, one siphonoglyph and a colonial way of life. Current systematics of Zoantharia is based exclusively on morphology and follows the traditional division of the group into the two suborders Brachycnemina and Macrocnemina, each comprising several poorly defined genera and species. To resolve the phylogenetic relationships among Zoantharia, we have analysed the sequences of mitochondrial 16S and 12S rRNA genes obtained from 24 specimens, representing two suborders and eight genera. In view of our data, Brachycnemina appears as a monophyletic group diverging within the paraphyletic Macrocnemina. The macrocnemic genus Epizoanthus branches as the sister group to all other Zoantharia that are sampled. All examined genera are monophyletic, except Parazoanthus, which comprises several independently branching clades and individual sequences. Among Parazoanthus, some groups of species can be defined by particular insertion/deletion patterns in the DNA sequences. All these clades show specificity to a particular type of substrate such as sponges or hydrozoans. Substrate specificity is also observed in zoantharians living on gorgonians or anthipatharians, as in the genus Savalia (Gerardia). If confirmed by further studies, the substrate specificity could be used as reliable character for taxonomic identification of some Macrocnemina.     

Sexual reproduction, larval development and benthic planulae of the solitary coral Monomyces rubrum (Scleractinia: Anthozoa)

The reproductive biology of the solitary ahermatypic coral Monomyces rubrum was studied in northeastern New Zealand between January 1996 and January 1998. The period of oogenesis lasted around 11 months, from late January to December, while spermatogenesis was more rapid, starting in late August and culminating in a spawning period in early December. Reproduction commenced at a polyp size of around 1,000 mm³ (5-6 years old) and the maximum estimated fecundity of the largest corals (7,000 mm³) was no more than 200 eggs. Oocytes were probably fertilized while within the mesentery and were shed into the coelenteron where they developed, via a solid blastula stage, for approximately 1 month. Planulae were relatively large, 3-4 mm in length and 1-2 mm diameter at the time of release, and crawled or swam immediately to the substratum. Peaks of planula shedding were semi-lunar in January 1997, but only one peak was observed in January 1998. The production of a few large rapidly settling larvae by this member of the family Flabellidae is consistent with the trend for solitary short-lived corals from other families to brood larvae rather than spawn gametes.     

Seasonal cycle of sexual reproduction of the Mediterranean soft coral Alcyonium acaule (Anthozoa, Octocorallia)

Alcyonium acaule (Cnidaria, Octocorallia) is a common, hard-bottom soft coral in the northwestern Mediterranean Sea. This study describes sexual reproduction and the gamete development cycle of this soft coral. A population at 15–18 m depth in the Marine Protected Area of the Medes Islands (42º02′55″ N, 3º13′30″ E) was sampled from July 1994–August 1995. A. acaule is gonochoristic and a surface brooder, spawning once a year in early summer. The mean diameter of ripe spermatic sacs was 400 ± 91 (SD) μm, and the mean diameter of mature oocytes was 473 ± 37 (SD) μm. There were 30 spermatic sacs polyp^?1 in males and 14 oocytes polyp^?1 in females. Different phases of gametogenesis in female and male colonies were examined separately with respect to seasonal changes in bottom temperature and solar irradiance. The data suggest that the relatively constant temperatures in January–April are probably not related to oocyte maturation, but that rising temperatures in May could affect sperm maturation. Rapidly increasing solar irradiance in March may be the trigger for vitellogenesis and oocyte maturation, although the mechanism for this in anthozoans is not understood.     

Synergistic effects of temperature,salinity and light on the hermatypic coral Montipora verrucosa

Temperature tolerance in the reef coral Montipora verrucosa (Lamarck) is affected by salinity and light. Low salinity reduces ability of the coral to survive shortterm exposure to elevated temperature. High natural light intensity aggravates damage sustained by corals at high temperature. In long-term growth experiments, high light intensity caused substantial loss of zooxanthellar pigment, higher mortality rates, reduced carbon fixation and lowered growth rate at both upper and lower sublethal temperatures Effects of light at optimal temperature were less dramatic. Interactions between physical environmental factors appear to be most important near the limits of tolerance for a given factor. Acclimation capability was indicated, and was influenced by both thermal history and pigmentation state of stressed corals.Contribution No. 543 of the Hawaii Institute of Marine Biology.     

Distribution,abundance and survival of juvenile hermatypic corals (Scleractinia) and the importance of life history strategies in the parent coral community

The distribution and abundance of juvenile corals were examined at depths from 3 to 37 m on the reefs of Curaçao and Bonaire (Netherlands Antilles). Juveniles of Agaricia agaricites were most abundant (60.6%), followed by Helioseris cucullata (8.3%). The large massive corals such as Montastrea annularis, M. cavernosa and branched species such as Madracis mirabilis and Acropora palmata had few juveniles. This, combined with species characteristics, shows that these species employ very different life history strategies. In some species the abundance of juveniles over the reef paralleled that of larger colonies, but not for example in Agaricia agaricites. The composition of the coral community was apparently no direct function of juvenile abundance. A change in angle of settlement of A. agaricites juveniles with increasing depth, from vertical to horizontal surfaces, seems to reflect the preferred light intensity. We studied the survival of juvenile corals during a half-year period. One-third remained unharmed, one-third died or disappeared, and one-third was limited in growth by factors such as spatial competition. This was the same for all depths, but factors influencing survival varied with depth.     

Seasonality in reproduction of the deep-water pennatulacean coral Anthoptilum grandiflorum

The deep-sea pennatulacean coral Anthoptilum grandiflorum exhibits a cosmopolitan distribution and was recently determined to serve as habitat for other invertebrates and fish larvae in the northwest Atlantic. Colonies collected at bathyal depths between 2006 and 2010 in eastern Canada were analysed to determine their fecundity and characterize spatial and temporal trends in their reproductive cycle. Anthoptilum grandiflorum is a gonochoric broadcast spawner with a sex ratio that does not differ significantly from equality (although one hermaphrodite colony was observed). In male colonies, all the spermatocysts synthesized become mature over the annual cycle, while only ~21 % of the initial production of oocytes reaches maturity in female colonies. Female potential fecundity based on mature oocytes just before spawning was on average 13 oocytes polyp^?1; male potential fecundity was 48 spermatocysts polyp^?1. The spawning period of A. grandiflorum differs between geographic regions, from April (in southern Newfoundland) to July (in Labrador), closely following regional spring phytoplankton blooms after accounting for the deposition of planktic detritus. Release of oocytes by a live colony held in the laboratory was recorded in April 2011, coincident with field data for similar latitudes. Seawater temperatures at the time of spawning were around 3.6–4.8 °C in all regions and in the laboratory. Early stages of gametogenesis were detected in colonies collected shortly after the spawning season, and early and late growth stages occurred successively until December. Mature colonies were observed between April and July (depending on latitude). The diameter of mature oocytes (~1,100 μm maximum diameter) is consistent with lecithotrophic larval development.     

Degradation and proliferation of zooxanthellae in planulae of the hermatypic coral Stylophora pistillata

Zooxanthellae in different stages of two opposite processes, degradation and proliferation, were found in the planulae of hermatypic corals. The formation of new zooxanthellae is balanced by degraded zooxanthellae in newly released planulae. The number of dividing zooxanthellae and degraded zooxanthellae during the day amounted to approximately 2 to 3% of the standing stock. In settled planulae and particularly in motionless planulae of Stylophora pistillata (Esper, 1797), the degraded zooxanthellae outnumbered proliferous zooxanthellae. The proliferation and degradation of zooxanthellae and the extrusion of degraded remnants of zooxanthellae are significantly phased. Swimming planulae are more autotrophic than motionless planulae. The physiological parameters of settled planulae with exoskeleton are similar to those of adult polyps. The significance of zooxanthella degradation in the vital functions of planulae is discussed. We suggest that the degradation of zooxanthellae in planulae occurs by the digestion of symbionts by host cells. Received: 5 March 1997 / Accepted: 6 August 1997     

Effect of temperature and temperature adaptation on calcification rate in the hermatypic coral Pocillopora damicornis

Using ⁴⁵Ca incorporation into the coral skeleton as a measure of calcification rate, the effect of temperature on clacification rate was studied in the hermatypic coral Pocillopora damicornis. Both immediate and long-term (adaptation) effects were investigated. Temperature has a marked effect on rate — an effect that varies depending on the temperature history of the coral (i.e., temperature adaptation occurs). P. damicornis showed both 27° and 31°C temperature optima, one or the other being dominant depending on the natural water temperature to which the coral was adapted. The two optimum temperatures may indicate two isoenzymes or two alternate metabolic pathways involved in the calcification process.     

Host specificity in sponge-encrusting zoanthidea (Anthozoa: Zoantharia) of Barbados,West Indies

Six species of common Caribbean Zoanthidea, Parazoanthus swiftii, P. parasiticus, P. catenularis, P. puertoricense, Epizoanthus cutressi, and Epizoanthus sp., are virtually restricted to living on surfaces of reef-dwelling sponges. Quantitative surveys on Barbados reefs indicate that substrate specificity is relatively high among these zoanthids with three restricted to a single primary host sponge species and three restricted to three closely related sponges. One species, P. swiftii, exhibits a broad range of acceptable secondary substrates, due to its unique ability to execute migrational spread in the adult polyp stage. Variations in substrate specificity have been noted between island populations within the extensive Caribbean range and appear to be due to different species compositions of local sponge communities and slight differences in zoanthid larval settling specificities.     

Effects of pCO2 on the interaction between an excavating sponge,Cliona varians,and a hermatypic coral,Porites furcata

Rising dissolved pCO₂ is a mounting threat to coral reef ecosystems. While the biological and physiological impacts of increased pCO₂ are well documented for many hermatypic corals, the potential effects on bioerosion processes remain largely unknown. Increases in pCO₂ are likely to modify the direct interactions between corals and bioeroders, such as excavating sponges, with broad implications for the balance between biologically mediated deposition and erosion of carbonate in reef communities. This study investigated the effects of three levels of CO₂ (present-day, mid-century and end-of-century projections) on the direct interaction between a bioeroding sponge, Cliona varians, and a Caribbean coral, Porites furcata. Increased pCO₂ concentrations had no effect on the attachment rates of C. varians to the corals, and we observed no significant impact of pCO₂ on the survival of either the coral or sponges. However, exposure to end-of-century levels of CO₂-dosing (~750 μatm) reduced calcification in P. furcata and led to a significant increase in sponge-mediated erosion of P. furcata. These findings demonstrate that pCO₂ can enhance erosional efficiency without impacting survival or competitive vigor in these two species. While few studies have considered the influence of pCO₂ on the competitive outcomes of interactions between corals and other reef organisms, our study suggests that assessing the impacts of changing pCO₂ on species interactions is crucial to adequately predict ecosystem-level responses in the future.     

Molecular evidence for multiple lineages in the gorgonian family Plexauridae (Anthozoa: Octocorallia)

Octocorals are diverse and abundant on many marine hard substrates, and, within this group, members of the family Plexauridae are an important component of tropical reef assemblages, especially in the Caribbean. To understand historical relationships within this large and diverse assemblage, and to test the monophyly of the family and some of its genera, DNA sequences of two mitochondrial loci (msh1 and ND2, ~1,185 bp) were analyzed from 46 species in 21 genera from deep and shallow waters in the tropical western Atlantic and in the tropical western and eastern Pacific (plus 9 taxa in the closely related Gorgoniidae and 1 species of the more distantly related Alcyoniidae). Five strongly supported clades were recovered. Three large clades correspond roughly to the Plexauridae, Paramuriceidae, and Gorgoniidae, and two smaller clades were comprised of taxa previously assigned to several families. Astrogorgia sp. did not group with any of the clades. The mutual relationships among the five clades remain unclear. Several genera previously regarded as unrelated appear to be grouped among the three families; e.g. Hypnogorgia sp. (Paramuriceidae) falls within a clade consisting of both Pacific and Atlantic Muricea spp. (Plexauridae), while Swiftia sp., Scleracis sp., and an Atlantic Thesea sp. (all Paramuriceidae) group with the gorgoniids. In several instances, genera containing Atlantic and Pacific species were recovered as monophyletic (Muricea spp., Bebryce spp.). However, in at least three cases (Echinomuricea spp., Thesea spp., Villogorgia spp.), placement of Atlantic and Pacific species in the same genus may reflect convergence of sclerite morphology. The results indicate a strong need for reexamination of octocoral taxonomy using a combination of molecular, morphological, and chemical evidence.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-005-1592-y.Communicated by J.P. Grassle, New Brunswick     

A systematic study of some Black Corals species (Antipatharia,Hexacorallia) based on rDNA internal transcribed spacers sequences

A phylogenetic analysis based on rDNA internal transcribed spacers (ITS) sequences was performed on 15 species of black corals assigned to the genera Antipathes, Stichopathes, Cirrhipathes, Rhipidipathes, Antipathella, Myriopathes and Cupressopathes recorded from the Messina Strait (Mediterranean Sea) and the Bunaken Marine Park (Celebes Sea, Indonesia). The phylogenetic analysis shows that the examined species are grouped in two main branches corresponding to the families Antipathidae-Aphanipathidae and Myriopathidae. While among the Myriopathidae species there is a very small genetic distance, the Antipathidae-Aphanipathidae clade shows a high degree of divergence between different genera. According to this study, the taxon Antipathes? sp. 3 characterised by a bush-like corallum without a well-defined axis, probably belongs to a new undescribed genus of the family Antipathidae.     

Sexual reproduction in the coral Pocillopora verrucosa (Cnidaria: Scleractinia) in KwaZulu-Natal, South Africa

The mode of reproduction in Pocillopora verrucosa Ellis and Solander, 1786 varies between geographically isolated regions. This scleractinian coral is common along the KwaZulu-Natal coast, and its reproductive mode and period of reproduction were assessed using histological preparations. The study was undertaken from 1992 to 1994 and showed that P. verrucosa is a simultaneous hermaphrodite and broadcast spawner in KwaZulu-Natal. Gametogenesis occurs from October to January, with the gametes maturing simultaneously so that spawning can be synchronised at new moon in January (mid-summer). Zooxanthellae are present in the mature oocytes. Received: 10 September 1997 / Accepted: 6 July 1998     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panama, and Galapagos Islands (Ecuador)

Reproductive ecology of two major reef-building corals in the eastern Pacific [Pocillopora damicornis (Linnaeus) andPocillopora elegans Dana] was investigated between 1984 and 1990 in Costa Rica, Panama (Gulf of Chiriqui and Gulf of Panama), and the Galapagos Islands (Ecuador) following the 1982–83 El Niño disturbance. Mature spermatocytes and oocytes were found in both species and were usually present in the same polyp in nearly equal ratios. Numerous zooxanthellae were observed in mature, unfertilized oocytes. Although spawning in field populations was not observed, it is likely that both pocilloporids are simultaneous hermaphroditic spawners, as evidenced by the disappearance of mature gametes after full moon. This is in contrast with most known pocilloporid corals that brood and release planula larvae. Corals were reproductively most active in the thermally stable environments of Costa Rica and the Gulf of Chiriqui (Panama) where 32 to 90% of all colonies contained gametes. In the moderately varying thermal conditions in the Galapagos Islands, 16 to 40% of colonies contained gametes, and in the pronounced seasonal upwelling environment of the Pearl Islands (Panama) only 6 to 18% of colonies contained gametes. Year-round reproduction occurred in Costa Rica and the Gulf of Chiriqui, whereas reproduction was confined to warm periods in the seasonally varying environments of the Galapagos Islands and the Gulf of Panama. Pocilloporid corals in Costa Rica and the Gulf of Chiriqui demonstrated lunar spawning activity, with mature gametes present a few days before and after full moon. Some limited spawning may have occurred also at new moon. While frequent gamete maturation has been demonstrated in this study, the relatively low rates of larval recruitment occurring on eastern Pacific coral reefs disturbed by the 1982–83 El Niño suggest that the recovery of important frame-building corals could be greatly prolonged.     

Sexual and asexual reproduction of Anthopleura dixoniana (Anthozoa: Actiniaria): periodicity and regulation

Reproduction of the sea anemone Anthopleura dixoniana (Haddon and Shackleton) from the high intertidal zone of southern Taiwan (120°41 E; 22°01N) was studied from April 1987 through March 1989. A. dixoniana spawns once a year, in July, and divides asexually by longitudinal fission throughout the year, with a peak in July. During the spawning season, sea anemones>3 mm pedal dise diameter can be sexed, and display a 1:1 sex ratio. Dividing sea anemones are significantly larger than non-dividing individuals, and increase in body size before fission. Under laboratory conditions, individuals kept at 28 C and fed had larger oocytes and a higher division rate than those kept at 18, 22, 25 or 32°C or starved. The division rate significantly influenced the oocyte diameter. The present study revealed for the first time, that a long photoperiod (14 h hight:10 h dark) significantly enhances the growth of oocytes in A. dixoniana under laboratory conditions.     

Limited gene flow in the brooding coral Favia fragum (Esper, 1797)

Understanding population connectivity in corals is particularly important as these organisms are increasingly threatened by abiotic and biotic factors. This study examined the population genetic structure of the brooding coral Favia fragum across four locations in the Caribbean and Western Atlantic using mitochondrial and nuclear markers. Morphological features were also compared to test whether genetic diversity corresponds with skeletal morphology. When comparing across distantly related Caribbean and Bermudian locations, F _ST values were high and significant, indicating strong genetic structure. At a local scale, significant genetic structure was found among reefs in Panama, while no genetic structure was found among reefs within Barbados, Bermuda or Jamaica. Surprisingly, a single haplotype for each of the three markers examined was found in Bermuda, where samples varied significantly from all other locations in three out of four morphological features analyzed. These data indicate that gene flow of F. fragum may occur locally among reefs but is highly restricted at distant locations. Furthermore, isolated populations, such as that of Bermuda, must be self-seeding to maintain the observed genetic uniformity.     

Gonadal morphology and gametogenesis in the sea pen Pennatula aculeata (Anthozoa: Pennatulacea) from the Gulf of Maine

The ultrastructural features of gametogenesis are described in male and female colonies of the sea pen Pennatula aculeata. Specimens were collected for observation and fixation at 113 to 231 m depth in the Gulf of Maine, USA, in August 1993. The species is gonochoric, and all stages of gametogenesis are observed in both male and female colonies >45 mm in height. Gametogenesis shows several features that differ from sea anemones. The developing oocytes and sperm cysts are completely encompassed by gastrodermally derived follicle cells, and they are released from the mesenteries into the coelenteron before they are fully differentiated. Following maturation in the coelenteron, the eggs and intact sperm cysts are expelled through the mouths of the autozoids during spawning. The expulsion of sperm cysts suggests that they function as primitive spermatophores, perhaps as a way of reducing sperm dilution. Vitellogenesis results in the biosynthesis of lipid droplets which are the sole nutrient reserves in the egg. Heterosynthetic vitellogenesis is characterized by the importation of lipid precursors into the oocyte, and there is some indirect evidence that hypertrophic follicle cells play a role in production, transport, and/or mediation of these precursors. Spermatogenesis is similar to that of other anthozoans. The spermatozoon has a cone-shaped head, a posterior nuclear fossa, a ring of lipid-like bodies in the midpiece, a prominent cytoplasmic collar surrounding the proximal flagellum, and a single mitochondrion, but the posterior region of the sperm also contains previously undescribed concentric rings of cisternae resembling smooth endoplasmic reticulum. Received: 23 March 1998 / Accepted: 31 July 1998     

Relationship of life-history patterns to depth in deep-water caridean shrimps (Crustacea: Natantia)

Deep-water shrimps, distributed on the steep outer reef slopes of tropical Pacific Islands, were obtained by setting baited traps in depths down to 850 m in the vicinity of Laucala Bay, Fiji, over the period 1979 to 1983. Life-history variables were estimated and interspecific comparisons made between Parapandalus serratifrons, Plesionika longirostris, Heterocarpus ensifer, H. gibbosus, H. sibogae, H. laevigatus (Pandalidae) and Saron marmoratus (Hippolytidae) which have different depth distributions. Results suggested that, interspecifically, reproductive lifespan tended to increase with increasing depth of distribution and that mean egg sizes of each species increased with increasing depth. Brood sizes tended to increase inconsistently with depth, although relative brood size (number of eggs per “standar” 15 g individual) appeared unrelated to depth. Although annual reproductive effort varied inconsistently with depth, reproductive effort totalled over the lifespan of the Heterocarpus species tended to increase with increasing depth. Some of the findings are counter to competition-based ecological theory, and it is proposed that adult predation decreases with increasing depth, allowing deeper-water species to have an extended lifespan, an increased degree of iteroparity, and a corresponding increase in lifetime reproductive effort. Further, we suggest that the probabilities of larval survival, which appear to decrease with increasing depth, are offset by the production of larger eggs by deeper-water species.