A multiscale analysis of coralline algae Lithophylloideae (Corallinophycidae, Rhodophyta) shedding new light on understanding cryptic diversity.

Cryptic diversity abounds in many biological species, posing challenges to our understanding of biological diversity, conservation and management. Taking the common coralline algae, the subfamily Lithophylloideae as an illustration, this study delved into the implications of cryptic diversity through global-level phylogenetic and geographical analysis based upon Lithophylloideae molecular data worldwide, as well as a multi-locus time-calibrated phylogeny to elucidate their possible evolutionary process. The multiscale analysis revealed the polyphyly in current concept of the genus Lithophyllum. Geographic isolation resulting from the Tethys terminal event (TTE) has led to two distinct distribution regions for this so-called cosmopolitan genus: one regionally distributed along European coasts/Mediterranean that should include the taxonomical Lithophyllum; others widely distributed, particularly among pan-tropic waters, suggesting at least five groups to be rediscovered within the subfamily Lithophylloideae. Meanwhile, the cryptic genus Titanoderma, lacking morphological identification features with Lithophyllum, exhibited differences in distribution and evolutionary patterns consistent with their ecological habits, thus supporting their separation. This study provided useful hints for cryptic diversity, which advocated an integrative thinking to investigating global cryptic diversity and exploring the broad linkages between phylogenetic relationships and evolutionary origin, biogeography, morphological and ecological traits to achieve a more comprehensive understanding of biodiversity.

It's time to broaden what we consider a 'blue carbon ecosystem'.

Photoautotrophic marine ecosystems can lock up organic carbon in their biomass and the associated organic sediments they trap over millennia and are thus regarded as blue carbon ecosystems. Because of the ability of marine ecosystems to lock up organic carbon for millennia, blue carbon is receiving much attention within the United Nations' 2030 Agenda for Sustainable Development as a nature-based solution (NBS) to climate change, but classically still focuses on seagrass meadows, mangrove forests, and tidal marshes. However, other coastal ecosystems could also be important for blue carbon storage, but remain largely neglected in both carbon cycling budgets and NBS strategic planning. Using a meta-analysis of 253 research publications, we identify other coastal ecosystems-including mud flats, fjords, coralline algal (rhodolith) beds, and some components or coral reef systems-with a strong capacity to act as blue carbon sinks in certain situations. Features that promote blue carbon burial within these 'non-classical' blue carbon ecosystems included: (1) balancing of carbon release by calcification via carbon uptake at the individual and ecosystem levels; (2) high rates of allochthonous organic carbon supply because of high particle trapping capacity; (3) high rates of carbon preservation and low remineralization rates; and (4) location in depositional environments. Some of these features are context-dependent, meaning that these ecosystems were blue carbon sinks in some locations, but not others. Therefore, we provide a universal framework that can evaluate the likelihood of a given ecosystem to behave as a blue carbon sink for a given context. Overall, this paper seeks to encourage consideration of non-classical blue carbon ecosystems within NBS strategies, allowing more complete blue carbon accounting.

In situ decrease in rhodolith growth associated with Arctic climate change.

Rhodoliths built by crustose coralline algae (CCA) are ecosystem engineers of global importance. In the Arctic photic zone, their three-dimensional growth emulates the habitat complexity of coral reefs but with a far slower growth rate, growing at micrometers per year rather than millimeters. While climate change is known to exert various impacts on the CCA's calcite skeleton, including geochemical and structural alterations, field observations of net growth over decade-long timescales are lacking. Here, we use a temporally explicit model to show that rising ocean temperatures over nearly 100 years were associated with reduced rhodolith growth at different depths in the Arctic. Over the past 90 years, the median growth rate was 85 µm year-1 but each °C increase in summer seawater temperature decreased growth by a mean of 8.9 µm (95% confidence intervals = 1.32-16.60 µm °C-1, p < .05). The decrease was expressed for rhodolith occurrences in 11 and 27 m water depth but not at 46 m, also having the shortest time series (1991-2015). Although increasing temperatures can spur plant growth, we suggest anthropogenic climate change has either exceeded the population thermal optimum for these CCA, or synergistic effects of warming, ocean acidification, and/or increasing turbidity impair rhodolith growth. Rhodoliths built by calcitic CCA are important habitat providers worldwide, so decreased growth would lead to yet another facet of anthropogenic habitat loss.

Changes in maerl-associated macroalgal community dynamics as evidence of anthropogenic pressure.

BACKGROUND AND AIMS: Maerl-associated communities have received considerable attention due to their uniqueness, biodiversity and functional importance. Although the impacts of human activities are well documented for maerl-associated macrofauna, the spatio-temporal variations of macroalgae have comparatively been neglected, and the drivers that influence their dynamics are poorly known. We investigate the links between maerl-associated macroalgal communities, anthropogenic pressures and environmental conditions, and hypothesize that sites under human pressure would exhibit different dynamics when compared to reference sites. METHODS: To better understand community variation through space and time, four subtidal maerl beds under different pressures were consistently monitored over one year in the bay of Brest, Brittany, France. Both macroalgae community monitoring and environmental data were acquired through field sampling and available models. KEY RESULTS: Higher macroalgal biomass was observed within eutrophic sites, especially in summer (more than ten times higher than in the Unimpacted site), caused by free-living forms of opportunistic red macroalgae. The Dredged site also exhibited distinct macroalgal communities during summer from the Unimpacted site. Nutrient concentrations and seasonality proved to be key factors affecting the macroalgal community composition, although dredging and its effects on granulometry also had a strong influence. Over the long term, fewer than half of the species identified during historical surveys were found, indicating major temporal changes. CONCLUSIONS: Human pressures have strong impacts on maerl-associated macroalgal communities. Nutrient concentrations and dredging pressure appear as the main anthropogenic factors shaping maerl-associated macroalgal communities. Additionally, our results suggest historical changes in maerl-associated macroalgal communities over 25 years in response to changes in local human pressure management. This study suggests that maerl-associated macroalgal communities could be used as indicators of anthropogenically driven changes in this habitat.

Investigating seed bank potential of crustose coralline algae using DNA metabarcoding.

To examine the potential for the autogenic ecosystem engineers, crustose coralline algae (CCA), to serve as seed banks or refugia for life stages of other species, it is critical to develop sampling protocols that reflect the diversity of life present. In this pilot study on two shallow water species of CCA collected from Raoul Island (Kermadec Islands; Rangitahua) New Zealand, we investigated two preservation methods (ethanol vs. silica gel), sampled inner and outer regions of the crusts, and used DNA metabarcoding and seven genes/gene regions (16S rRNA, 18S rRNA, 23S rRNA, cox1, rbcL, and tufA genes and the ITS rRNA region) to develop a protocol for taxa identification. The results revealed immense diversity, with typically more taxa identified within the inner layers than the outer layers. As highlighted in other metabarcoding studies and in earlier work on rhodoliths (nodose coralline algae), reference databases are incomplete, and to some extent, the use of multiple markers mitigates this issue. Specifically, the 23S rRNA and rbcL genes are currently more suitable for identifying algae, while the cox1 gene fares better at capturing the diversity present inclusive of algae. Further investigation of these autogenic ecosystem engineers that likely act as marine seed banks is needed.

Coral larval settlement induction using tissue-associated and exuded coralline algae metabolites and the identification of putative chemical cues.

Reef-building crustose coralline algae (CCA) are known to facilitate the settlement and metamorphosis of scleractinian coral larvae. In recent decades, CCA coverage has fallen globally and degrading environmental conditions continue to reduce coral survivorship, spurring new restoration interventions to rebuild coral reef health. In this study, naturally produced chemical compounds (metabolites) were collected from two pantropical CCA genera to isolate and classify those that induce coral settlement. In experiments using four ecologically important Caribbean coral species, we demonstrate the applicability of extracted, CCA-derived metabolites to improve larval settlement success in coral breeding and restoration efforts. Tissue-associated CCA metabolites induced settlement of one coral species, Orbicella faveolata, while metabolites exuded by CCA (exometabolites) induced settlement of three species: Acropora palmata, Colpophyllia natans and Orbicella faveolata. In a follow-up experiment, CCA exometabolites fractionated and preserved using two different extraction resins induced the same level of larval settlement as the unfractionated positive control exometabolites. The fractionated CCA exometabolite pools were characterized using liquid chromatography tandem mass spectrometry, yielding 145 distinct molecular subnetworks that were statistically defined as CCA-derived and could be classified into 10 broad chemical classes. Identifying these compounds can reveal their natural prevalence in coral reef habitats and facilitate the development of new applications to enhance larval settlement and the survival of coral juveniles.

Settlement cue selectivity by larvae of the destructive crown-of-thorns starfish.

Population irruptions of crown-of-thorns starfish (COTS) cause extensive degradation of coral reefs, threatening the structure and function of these important ecosystems. For population irruptions to initiate and spread, large numbers of planktonic larvae have to successfully transition into their benthic life-history stage (i.e. settlement), whereby larval behaviour and the presence of settlement cues may shape spatial patterns of recruitment and adult densities. Our results demonstrate that a wide range of coralline algae species induce COTS larvae to settle; however, the capacity to promote settlement success varied manyfold among algal species, ranging from greater than 90% in Melyvonnea cf. madagascariensis to less than 2% in Lithophyllum cf. kotschyanum and two Porolithon species at 24 h. Because many coralline algae species that promote high settlement success are prevalent in shallow reef habitats, our findings challenge the hypothesis that COTS larvae predominantly settle in deep water. Considering both larval behaviour and algal ecology, this study highlights the ecological significance of coralline algae communities in driving recruitment patterns of COTS. More specifically, the local abundance of highly inductive coralline algae (especially, Melyvonnea cf. madagascariensis) may explain some of the marked spatial heterogeneity of COTS populations and the incidence of population irruptions.

Seasonal upwelling conditions promote growth and calcification in reef-building coralline algae.

Crustose coralline algae (CCA) are important components of reef ecology contributing to reef framework construction. However, little is known about how seasonal upwelling systems influence growth and calcification of tropical CCA. We assessed marginal and vertical growth and net calcification rates of two dominant but morphologically different reef-building CCA, Porolithon antillarum and Lithophyllum cf. kaiseri, in a shallow coral reef of the Colombian Caribbean during upwelling and non-upwelling seasons. Growth and calcification rates varied seasonally with higher values during the upwelling compared to the non-upwelling (rainy) season. Annual vertical growth showed rates of 4.48 ± 1.58 and 4.31 ± 2.17 mm · y-1 , net calcification using crust growth estimates of 0.75 ± 0.30 g and 0.68 ± 0.60 g CaCO3 · cm-2 · y-1 and net calcification using the buoyant weight method of 1.49 ± 0.57 and 0.52 ± 0.11 g CaCO3 · cm-2 · y-1 in P. antillarum and L. kaiseri, respectively. Seawater temperature was inversely related with growth and calcification; however, complex oceanographic interactions between temperature and resource availability (e.g., light, nutrients, and CO2 ) are proposed to modulate CCA vital rates. Although CCA calcification rates are comparable to hard corals, CCA vertical accretion is much lower, suggesting that the main contribution of CCA to reef construction is via cementation processes. These results provide baseline data on CCA in the region and generate useful information for monitoring the impacts of environmental changes on tropical upwelling environments.

New branched Porolithon species (Corallinales, Rhodophyta) from the Great Barrier Reef, Coral Sea, and Lord Howe Island.

Porolithon is one of the most ecologically important genera of tropical and subtropical crustose (non-geniculate) coralline algae growing abundantly along the shallow margins of coral reefs and functioning to cement reef frameworks. Thalli of branched, fruticose Porolithon specimens from the Indo-Pacific Ocean traditionally have been called P. gardineri, while massive, columnar forms have been called P. craspedium. Sequence comparisons of the rbcL gene both from type specimens of P. gardineri and P. craspedium and from field-collected specimens demonstrate that neither species is present in east Australia and instead resolve into four unique genetic lineages. Porolithon howensis sp. nov. forms columnar protuberances and loosely attached margins and occurs predominantly at Lord Howe Island; P. lobulatum sp. nov. has fruticose to clavate forms and free margins that are lobed and occurs in the Coral Sea and on the Great Barrier Reef (GBR); P. parvulum sp. nov. has short (<2 cm), unbranched protuberances and attached margins and is restricted to the central and southern GBR; and P. pinnaculum sp. nov. has a mountain-like, columnar morphology and occurs on oceanic Coral Sea reefs. A rbcL gene sequence of the isotype of P. castellum demonstrates it is a different species from other columnar species. In addition to the diagnostic rbcL and psbA marker sequences, the four new species may be distinguished by a combination of features including thallus growth form, margin shape (attached or unattached), and medullary system (coaxial or plumose). Porolithon species, because of their ecological importance and sensitivity to ocean acidification, need urgent documentation of their taxonomic diversity.

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.

Red algae acclimate to low light by modifying phycobilisome composition to maintain efficient light harvesting.

BACKGROUND: Despite a global prevalence of photosynthetic organisms in the ocean's mesophotic zone (30-200+ m depth), the mechanisms that enable photosynthesis to proceed in this low light environment are poorly defined. Red coralline algae are the deepest known marine benthic macroalgae - here we investigated the light harvesting mechanism and mesophotic acclimatory response of the red coralline alga Lithothamnion glaciale. RESULTS: Following initial absorption by phycourobilin and phycoerythrobilin in phycoerythrin, energy was transferred from the phycobilisome to photosystems I and II within 120 ps. This enabled delivery of 94% of excitations to reaction centres. Low light intensity, and to a lesser extent a mesophotic spectrum, caused significant acclimatory change in chromophores and biliproteins, including a 10% increase in phycoerythrin light harvesting capacity and a 20% reduction in chlorophyll-a concentration and photon requirements for photosystems I and II. The rate of energy transfer remained consistent across experimental treatments, indicating an acclimatory response that maintains energy transfer. CONCLUSIONS: Our results demonstrate that responsive light harvesting by phycobilisomes and photosystem functional acclimation are key to red algal success in the mesophotic zone.

Transcriptomic stability or lability explains sensitivity to climate stressors in coralline algae.

BACKGROUND: Crustose coralline algae (CCA) are calcifying red macroalgae that play important ecological roles including stabilisation of reef frameworks and provision of settlement cues for a range of marine invertebrates. Previous research into the responses of CCA to ocean warming (OW) and ocean acidification (OA) have found magnitude of effect to be species-specific. Response to OW and OA could be linked to divergent underlying molecular processes across species. RESULTS: Here we show Sporolithon durum, a species that exhibits low sensitivity to climate stressors, had little change in metabolic performance and did not significantly alter the expression of any genes when exposed to temperature and pH perturbations. In contrast, Porolithon onkodes, a major coral reef builder, reduced photosynthetic rates and had a labile transcriptomic response with over 400 significantly differentially expressed genes, with differential regulation of genes relating to physiological processes such as carbon acquisition and metabolism. The differential gene expression detected in P. onkodes implicates possible key metabolic pathways, including the pentose phosphate pathway, in the stress response of this species. CONCLUSIONS: We suggest S. durum is more resistant to OW and OA than P. onkodes, which demonstrated a high sensitivity to climate stressors and may have limited ability for acclimatisation. Understanding changes in gene expression in relation to physiological processes of CCA could help us understand and predict how different species will respond to, and persist in, future ocean conditions predicted for 2100.

Geographic distance, sedimentation, and substrate shape cryptic crustose coralline algal assemblages in the world's largest subtropical intertidal algal reef.

Algal reefs, concreted by crustose coralline algae (CCA), are the main biotic reefs in temperate waters but rare in the subtropics and tropics. The world's largest known intertidal algal reef in the subtropics is the Taoyuan Algal Reef (TAR) located in the northwestern coast of Taiwan. The biodiversity and ecology of the TAR are scarcely explored, and now the reef is imperiled by industrialization. Here, we document cryptic species of CCA in Taiwan, particularly the TAR, by sequencing the psbA genes of over 1800 specimens collected across Taiwan. We also examine the ecological background of the TAR by surveying its benthic composition and measuring its environmental parameters. Our data reveal that the TAR harbours a high diversity of cryptic CCA species (27 molecular operational taxonomic units, or mOTUs), many of which are potentially new to science (18 mOTUs) and/or endemic to the TAR (9 mOTUs). Comparing the CCA species inventory of the TAR with the rest of Taiwan shows that the TAR represents a unique hotspot of CCA taxa in the waters of Taiwan. Our analyses show that variation in the CCA assemblages in the TAR is associated with geographic distance, sedimentation, and substrate type (for example, reef vs. hermit crab shell), suggesting that dispersal limitation and contemporary environmental selection shape the CCA assemblages in the TAR. The data from this study can inform the monitoring of human impacts on the health of the TAR and contribute to our understanding of the ecological processes underlying algal reef development.

Coralline photosynthetic physiology across a steep light gradient.

Coralline algae (CA) are globally distributed and fulfil many important roles within coastal ecosystems. In this study, photosynthetically active radiation (PAR) measured for 616 days at 2 and 10 m in a temperate subtidal kelp forest in southern New Zealand provided context to photosynthesis vs. irradiance relationships for, and pigment concentrations of, an articulated coralline alga, Arthrocardia sp. and a crustose coralline species assemblage within the Hapalidiales order. The maximum photosynthetic rate Pmax of the Arthrocardia sp. (20.38 ± 2.38 µmol O2. gDW-1 h-1) was significantly higher than the Pmax of crustose coralline spp. (3.72 ± 0.74 µmol O2. gDW-1 h-1) at the same 2 m stratum. Pigment concentration of Arthrocardia sp. was significantly higher than that of crustose coralline spp. at the same depth, while pigment concentration of crustose coralline spp. at 2 and 10 m were not significantly affected by depth. The photosynthetic characteristics of these coralline algae represent a shade acclimated organism with low saturation irradiance (all Ek < 100 µmol photons m-2 s-1). Despite sevenfold difference in average daily dose between 2 and 10 m there was no significant effect of depth on the photosynthetic performance of crustose coralline algae measured. The lack of evidence for acclimation to low light could be because periods of clear water provide enough light to maintain photosynthesis, lower energetic requirements of species found at depth or constraints on the synthesis of photosynthetic pigments at greater depth.

Understanding coralline algal responses to ocean acidification: Meta-analysis and synthesis.

Ocean acidification (OA) is a major threat to the persistence of biogenic reefs throughout the world's ocean. Coralline algae are comprised of high magnesium calcite and have long been considered one of the most susceptible taxa to the negative impacts of OA. We summarize these impacts and explore the causes of variability in coralline algal responses using a review/qualitative assessment of all relevant literature, meta-analysis, quantitative assessment of critical responses, and a discussion of physiological mechanisms and directions for future research. We find that most coralline algae experienced reduced abundance, calcification rates, recruitment rates, and declines in pH within the site of calcification in laboratory experiments simulating OA or at naturally elevated CO2 sites. There were no other consistent physiological responses of coralline algae to simulated OA (e.g., photo-physiology, mineralogy, and survival). Calcification/growth was the most frequently measured parameters in coralline algal OA research, and our meta-analyses revealed greater declines in seawater pH were associated with significant decreases in calcification in adults and similar but nonsignificant trends for juveniles. Adults from the family Mesophyllumaceae also tended to be more robust to OA, though there was insufficient data to test similar trends for juveniles. OA was the dominant driver in the majority of laboratory experiments where other local or global drivers were assessed. The interaction between OA and any other single driver was often additive, though factors that changed pH at the surface of coralline algae (light, water motion, epiphytes) acted antagonistically or synergistically with OA more than any other drivers. With advances in experimental design and methodological techniques, we now understand that the physiology of coralline algal calcification largely dictates their responses to OA. However, significant challenges still remain, including improving the geographic and life-history spread of research effort and a need for holistic assessments of physiology.

Nitrogen Fixation Influenced by Phosphorus and Nitrogen Availability in the Benthic Bloom-forming Cyanobacterium Hydrocoleum sp. Identified in a Temperate Marine Lagoon.

The nitrogen-fixing, non-heterocystous cyanobacterium Hydrocoleum sp. (Oscillatoriales) is a common epiphytic and benthic bloom-former in tropical and subtropical shallow water systems but shares high phylogenetic similarity with the planktonic, globally important diazotroph Trichodesmium. Multiphasic observations in this study resulted in unexpected identification of Hydrocoleum sp. in mass accumulations in a coastal lagoon in the Western temperate North Atlantic Ocean. Hydrocoleum physiology was examined in situ through measurements of N2 and CO2 fixation rates and expression of genes involved with N2 fixation, CO2 fixation, and phosphorus (P) stress. Bulk N2 fixation rates and Hydrocoleum nifH expression peaked at night and were strongly suppressed by dissolved inorganic nitrogen (DIN). The expression of high affinity phosphate transporter (pstS) and alkaline phosphatase (phoA) genes of Hydrocoleum was elevated during the night and negatively responded to phosphate amendments, as evidence that these mechanisms contribute to P acquisition during diazotrophic growth of Hydrocoleum in situ. This discovery at the edge of the previously known Hydrocoleum habitat range in the warming oceans raises intriguing questions about diazotrophic cyanobacterial adaptations and transitions on the benthic-pelagic continuum.

Global distribution and diversity of marine euendolithic cyanobacteria.

Euendolithic, or true-boring, cyanobacteria actively erode carbonate-containing substrata in a wide range of environments and pose significant risks to calcareous marine fauna. Their boring activities cause structural damage and increase susceptibility to disease and are projected to only intensify with global climate change. Most research has, however, focused on tropical coral systems, and limited information exists on the global distribution, diversity, and substratum specificity of euendoliths. This metastudy aimed to collate existing 16S rRNA gene surveys along with novel data from the south coast of South Africa to investigate the global distribution and genetic diversity of endoliths to identify a "core endolithic cyanobacterial microbiome" and assess global diversification of euendolithic cyanobacteria. The cyanobacterial families Phormidesmiaceae, Nodosilineaceae, Nostocaceae, and Xenococcaceae were the most prevalent, found in >92% of categories surveyed. All four known euendolith clusters were detected in both intertidal and subtidal habitats, in the North Atlantic, Mediterranean, and South Pacific oceans, across temperate latitudes, and within rock, travertine tiles, coral, shell, and coralline algae substrata. Analysis of the genetic variation within clusters revealed many organisms to be unique to substratum type and location, suggesting high diversity and niche specificity. Euendoliths are known to have important effects on their hosts. This is particularly important when hosts are globally significant ecological engineers or habitat-forming species. The findings of this study indicate high ubiquity and diversity of euendolithic cyanobacteria, suggesting high adaptability, which may lead to increased community and ecosystem-level effects with changing climatic conditions favoring the biochemical mechanisms of cyanobacterial bioerosion.

Phymatolithopsis gen. nov. (Hapalidiales, Corallinophycidae, Rhodophyta) based on molecular and morpho-anatomical evidence.

A multigene (psbA, rbcL, 18S rDNA) molecular phylogeny of the genus Phymatolithon showed a polyphyletic grouping of two monophyletic clades within the Hapalidiales. DNA sequence data integrated with morpho-anatomical comparisons of type material and of recently collected specimens were used to establish Phymatolithopsis gen. nov. with three species, P. prolixa comb. nov., the generitype, P. repanda comb. nov. and P. donghaensis sp. nov. Phymatolithopsis is sister to Mesophyllum and occurs in a clade distinct from Phymatolithon and boreal species currently assigned to Lithothamnion. Morpho-anatomically, Phymatolithopsis is comprised of species that are non-geniculate and encrusting, bear epithallial cells with rounded walls (not flared), subepithallial initials that are usually as short as or shorter than their immediate inward derivatives, conceptacle primordia from all stages forming superficially directly from subepithallial initials, mature carposporangial conceptacles with a discontinuous fusion cell, gonimoblast filaments that develop at the margins of the fusion cell around the periphery of the carposporangial conceptacle chambers, and multiporate tetra/bisporangial conceptacles. Phymatolithopsis can be distinguished from Phymatolithon by the origin of its conceptacle primordia, which are initiated superficially, directly from the layer of subepithallial initials below the epithallial cells and the distribution of gonimoblast filaments in carposporangial conceptacles, that are at the margins of the fusion cells.

Consequences of Warming and Acidification for the Temperate Articulated Coralline Alga, Calliarthron Tuberculosum (Florideophyceae, Rhodophyta).

Global climate changes, such as warming and ocean acidification (OA), are likely to negatively impact calcifying marine taxa. Abundant and ecologically important coralline algae may be particularly susceptible to OA; however, multi-stressor studies and those on articulated morphotypes are lacking. Here, we use field observations and laboratory experiments to elucidate the impacts of warming and acidification on growth, calcification, mineralogy, and photophysiology of the temperate articulated coralline alga, Calliarthron tuberculosum. We conducted a 4-week fully factorial mesocosm experiment exposing individuals from a southern CA kelp forest to current and future temperature and pH/pCO2 conditions (+2°C, -0.5 pH units). Calcification was reduced under warming (70%) and further reduced by high pCO2 or high pCO2 x warming (~150%). Growth (change in linear extension and surface area) was reduced by warming (40% and 50%, respectively), high pCO2 (20% and 40%, respectively), and high pCO2 x warming (50% and 75%, respectively). The maximum photosynthetic rate (Pmax ) increased by 100% under high pCO2 conditions, but we did not detect an effect of pCO2 or warming on photosynthetic efficiency (α). We also did not detect the effect of warming or pCO2 on mineralogy. However, variation in Mg incorporation in cell walls of different cell types (i.e., higher mol % Mg in cortical vs. medullary) was documented for the first time in this species. These results support findings from a growing body of literature suggesting that coralline algae are often more negatively impacted by warming than OA, with the potential for antagonistic effects when factors are combined.

Growth Resilience of Subarctic Rhodoliths (Lithothamnion glaciale, Rhodophyta) to Chronic Low Sea Temperature and irradiance.

Rhodolith beds are pervasive marine biological systems in the subarctic North Atlantic. Limited knowledge about effects of temperature and irradiance on rhodolith growth limits the ability to anticipate the response of rhodolith beds to this ocean's chronic low, yet changing sea temperature and irradiance regimes. We carried out a 149-d laboratory experiment with Newfoundland Lithothamnion glaciale rhodoliths to test the predictions that growth (i) is inhibited at temperatures of ~0.5°C and (ii) resumes as temperature increases above 0.5°C, albeit at a higher rate under high than low irradiances. Rhodoliths were grown in experimental tanks at near-zero (~0.7°C) seawater temperatures during the first 85 d and at temperatures increasing naturally to ~6°C for the remaining 64 d. Rhodoliths in those tanks were exposed to either low (0.02 mol photons·m-2 ·d-1 ) or high (0.78 mol photons·m-2 ·d-1 ) irradiances during the entire experiment. Rhodoliths grew at a linear rate of ~281 µm·year-1 (0.77 µm·d-1 ) throughout the experiment under both irradiance treatments despite daily seawater temperature variation of up to 3°C. Near-zero temperatures of ~0.5 to 1.0°C did not inhibit rhodolith growth. Model selection showed that PAR-day (a cumulative irradiance index) was a better predictor of growth variation than Degree-day (a cumulative thermal index). Our findings extend to ~0.5°C the lower limit of the known temperature range (~1 to at least 16°C) over which growth in L. glaciale rhodoliths remains unaffected, while suggesting that the growth-irradiance relationship in low-light environments at temperatures below 6°C is less irradiance-driven than recently proposed.