Revision of Corallinaceae (Corallinales,Rhodophyta): recognizing Dawsoniolithon gen. nov., Parvicellularium gen. nov. and Chamberlainoideae subfam. nov. containing Chamberlainium gen. nov. and Pneophyllum

A multi‐gene (SSU, LSU, psbA, and COI) molecular phylogeny of the family Corallinaceae (excluding the subfamilies Lithophylloideae and Corallinoideae) showed a paraphyletic grouping of six monophyletic clades. Pneophyllum and Spongites were reassessed and recircumscribed using DNA sequence data integrated with morpho‐anatomical comparisons of type material and recently collected specimens. We propose Chamberlainoideae subfam. nov., including the type genus Chamberlainium gen. nov., with C. tumidum comb. nov. as the generitype, and Pneophyllum. Chamberlainium is established to include several taxa previously ascribed to Spongites, the generitype of which currently resides in Neogoniolithoideae. Additionally we propose two new genera, Dawsoniolithon gen. nov. (Metagoniolithoideae), with D. conicum comb. nov. as the generitype and Parvicellularium gen. nov. (subfamily incertae sedis), with P. leonardi sp. nov. as the generitype. Chamberlainoideae has no diagnostic morpho‐anatomical features that enable one to assign specimens to it without DNA sequence data, and it is the first subfamily to possess both Type 1 (Chamberlainium) and Type 2 (Pneophyllum) tetra/bisporangial conceptacle roof development. Two characters distinguish Chamberlainium from Spongites: tetra/biasporangial conceptacle chamber diameter (<300 μm in Chamberlainium vs. >300 μm in Spongites) and tetra/bisporangial conceptacle roof thickness (<8 cells in Chamberlainium vs. >8 cells in Spongites). Two characters also distinguish Pneophyllum from Dawsoniolithon: tetra/bisporangial conceptacle roof thickness (<8 cells in Pneophyllum vs. >8 cells in Dawsoniolithon) and thallus construction (dimerous in Pneophyllum vs. monomerous in Dawsoniolithon).     

One‐year experiment on the physiological response of the Mediterranean crustose coralline alga,Lithophyllum cabiochae,to elevated pCO2 and temperature

The response of respiration, photosynthesis, and calcification to elevated pCO₂ and temperature was investigated in isolation and in combination in the Mediterranean crustose coralline alga Lithophyllum cabiochae. Algae were maintained in aquaria during 1 year at near‐ambient conditions of irradiance, at ambient or elevated temperature (+3°C), and at ambient (ca. 400 μatm) or elevated pCO₂ (ca. 700 μatm). Respiration, photosynthesis, and net calcification showed a strong seasonal pattern following the seasonal variations of temperature and irradiance, with higher rates in summer than in winter. Respiration was unaffected by pCO₂ but showed a general trend of increase at elevated temperature at all seasons, except in summer under elevated pCO₂. Conversely, photosynthesis was strongly affected by pCO₂ with a decline under elevated pCO₂ in summer, autumn, and winter. In particular, photosynthetic efficiency was reduced under elevated pCO₂. Net calcification showed different responses depending on the season. In summer, net calcification increased with rising temperature under ambient pCO₂ but decreased with rising temperature under elevated pCO₂. Surprisingly, the highest rates in summer were found under elevated pCO₂ and ambient temperature. In autumn, winter, and spring, net calcification exhibited a positive or no response at elevated temperature but was unaffected by pCO₂. The rate of calcification of L. cabiochae was thus maintained or even enhanced under increased pCO₂. However, there is likely a trade‐off with other physiological processes. For example, photosynthesis declines in response to increased pCO₂ under ambient irradiance. The present study reports only on the physiological response of healthy specimens to ocean warming and acidification, however, these environmental changes may affect the vulnerability of coralline algae to other stresses such as pathogens and necroses that can cause major dissolution, which would have critical consequence for the sustainability of coralligenous habitats and the budgets of carbon and calcium carbonate in coastal Mediterranean ecosystems.     

The Role of Encrusting Coralline Algae in the Diets of Selected Intertidal Herbivores

Kalk Bay, South Africa, has a typical south coast zonation pattern with a band of seaweed dominating the mid-eulittoral and between two molluscan-herbivore dominated upper and lower eulittoral zones. Encrusting coralline algae were very obvious features of these zones. The most abundant herbivores in the upper eulittoral were the limpet, Cymbula oculus (10.4 ± 1.6 individuals m⁻²; 201.65 ± 32.68 g.m⁻²) and the false limpet, Siphonaria capensis (97.07± 19.92 individuals m⁻²; 77.93 16.02 g.m⁻²). The territorial gardening limpet, Scutellastra cochlear, dominated the lower eulittoral zone, achieving very high densities (545.27 ± 84.35 m⁻²) and biomass (4630.17 ± 556.13 g.m⁻²), and excluded all other herbivores and most seaweeds, except for its garden alga and the encrusting coralline alga, Spongites yendoi (35.93 ± 2.26% cover). In the upper eulittoral zone, encrusting coralline algae were only present in the guts of the chiton Acanthochiton garnoti (30.5 ± 1.33%) and the limpet C. oculus (2.9 ± 0.34%). The lower eulittoral zone limpet, Scutellastra cochlear also had a large percentage of encrusting coralline algae in its gut with limpets lacking gardens having higher (45.1 ± 1.68%) proportions of coralline algae in their guts than those with gardens (25.6 ± 0.8%). Encrusting coralline algae had high organic contents, similar to those of other encrusting and turf-forming algae, but higher organic contents than foliose algae. Radula structure, grazing frequencies as a percentage of the area grazed (upper eulittoral 73.25 ± 3.60% d⁻¹; lower eulittoral 46.0 ± 3.29% d⁻¹), and algal organic content provided evidence to support the dietary habits of the above herbivores. The data show that many intertidal molluscs are actively consuming encrusting coralline algae and that these seaweeds should be seen as an important food source.     

CLATHROMORPHUM NEREOSTRATUM (CORALLINALES,RHODOPHYTA): THE OLDEST ALGA?1

The longevity of organisms is intrinsically interesting and can provide useful information on their population structure and dynamics and the dynamics of associated communities. With the exception of perennial Laminariales that have rings in the stipe, the life spans of most perennial macroalgae are unknown or based on anecdotal observations. Using morphological analyses combined with the location and time of the rise in ¹⁴C from atmospheric nuclear testing within the thallus, we determined that the growth rate of a specimen of Clathromorphum nereostratum Lebednik from Adak Island was 0.30 mm·yr^?1, the 3⁰ bands within the thallus were annual, and the specimen sampled was 61–75 years old. Living crusts of this species from the same geographic region are reported to be up to 20 cm thick. Assuming our growth rate is typical, C. nereostratum can be approximately 700 years old, the oldest known living alga. This longevity and consistent banding within the thallus suggest that smaller scale sampling and additional chemical analyses of this alga could provide a detailed long‐term record of environmental variation at high latitudes in the North Pacific.     

Microbial to reef scale interactions between the reef-building coral Montastraea annularis and benthic algae

Competition between reef-building corals and benthic algae is of key importance for reef dynamics. These interactions occur on many spatial scales, ranging from chemical to regional. Using microprobes, 16S rDNA pyrosequencing and underwater surveys, we examined the interactions between the reef-building coral Montastraea annularis and four types of benthic algae. The macroalgae Dictyota bartayresiana and Halimeda opuntia, as well as a mixed consortium of turf algae, caused hypoxia on the adjacent coral tissue. Turf algae were also associated with major shifts in the bacterial communities at the interaction zones, including more pathogens and virulence genes. In contrast to turf algae, interactions with crustose coralline algae (CCA) and M. annularis did not appear to be antagonistic at any scale. These zones were not hypoxic, the microbes were not pathogen-like and the abundance of coral-CCA interactions was positively correlated with per cent coral cover. We propose a model in which fleshy algae (i.e. some species of turf and fleshy macroalgae) alter benthic competition dynamics by stimulating bacterial respiration and promoting invasion of virulent bacteria on corals. This gives fleshy algae a competitive advantage over corals when human activities, such as overfishing and eutrophication, remove controls on algal abundance. Together, these results demonstrate the intricate connections and mechanisms that structure coral reefs.     

Does colonization contribute to spatial patterns of common invertebrates in coralline algal turf?

Abstract The potential of colonization to contribute to the spatial patterns of six common invertebrates in coralline algal turf was investigated on a rocky shore near Sydney, Australia. The species, which included two amphipods (Elasmopus warra, Hyale spp.), a small bivalve (Lasaea australis), a fly larva (Limonia marina), and two microgastropods (Amphithalamus incidata and Eatoniella atropurpurea), had a range of dispersal modes (larval dispersal, crawling, swimming, rafting, and passive transport). Field sampling between May 1997 and November 1999 demonstrated that the amphipods were more abundant in low‐shore areas, the fly larvae and bivalves were more abundant in mid‐shore areas, and the abundances of gastropods did not vary with tidal height. Furthermore, abundances of all species varied among patches separated by tens of metres at one time or another. To test whether rates of colonization could contribute to established patterns of abundance, habitat mimics were deployed for 2‐week periods. The supply of new individuals matched long‐term patterns of abundance at different tidal heights for E. warra and L. marina. Colonization rates also differed among patches separated by tens of metres for three of the six species. Overall, there was little evidence to suggest that common species in coralline turf are limited by colonization on local scales, regardless of their major mode of dispersal. However, the potential for colonization to determine patterns of abundance varied from species to species.     

Responses of crustose corallines to epiphyte and canopy cover

Although a dense cover of epiphytes is generally considered to be harmful for some coralline algae (Corallinaceae, Rhodophyta), crustose corallines in the littoral zone seem to be preserved from bleaching when covered by canopy plants and epiphytes during summer. This study aimed to test the responses of coralline crusts to epiphytes and canopy algae and their interaction with grazing limpets. Growth rates and color changes were followed in two crust species in areas with or without canopy algae in the Isle of Man, British Isles. Limpets were excluded, to allow epiphytes to grow upon crusts. Responses were measured both on pieces of crusts upon acrylic plates and on crusts growing naturally on the shore. Fucus canopy and epiphytic Enteromorpha significantly influenced the crusts' growth, depending on season. Epiphytes reduced the light levels beneath by up to 78%, more than the canopy algae (62%). Crusts exposed outside the canopy bleached in summer, but gradually restored their color once they were covered by epiphytes. The fast-growing Phymatolithon lenormandii (Aresch.) Adey recovered its coloration more quickly than the slow-growing P. purpureum (P. et H. Crouan) Woelkerling et Irvine. However, neither crust species could restore its color when epiphytes were reduced by grazing limpets, Patella vulgata L. Bleaching did not kill the crusts, but seemed to interfere with crusts' growth. Restoration of pigmentation was quantified for the first time on bleached coralline crusts. Epiphyte and canopy algae were experimentally shown to be beneficial, probably by providing shade and also protecting crusts from desiccation.     

Cell wall organic matrix composition and biomineralization across reef-building coralline algae under global change

Crustose coralline algae (CCA) are one of the most important benthic substrate consolidators on coral reefs through their ability to deposit calcium carbonate on an organic matrix in their cell walls. Discrete polysaccharides have been recognized for their role in biomineralization, yet little is known about the carbohydrate composition of organic matrices across CCA taxa and whether they have the capacity to modulate their organic matrix constituents amidst environmental change, particularly the threats of ocean acidification (OA) and warming. We simulated elevated pCO₂ and temperature (IPCC RCP 8.5) and subjected four mid-shelf Great Barrier Reef species of CCA to 2 months of experimentation. To assess the variability in surficial monosaccharide composition and biomineralization across species and treatments, we determined the monosaccharide composition of the polysaccharides present in the cell walls of surficial algal tissue and quantified calcification. Our results revealed dissimilarity among species' monosaccharide constituents, which suggests that organic matrices are composed of different polysaccharides across CCA taxa. We also observed that species differentially modulate composition in response to ocean acidification and warming. Our findings suggest that both variability in composition and ability to modulate monosaccharide abundance may play a crucial role in surficial biomineralization dynamics under the stress of OA and global warming.