Oldest fossil ciliates from the Cryogenian glacial interlude reinterpreted as possible red algal spores

The Cryogenian Period experienced two long lived global glaciations known as Snowball Earths. While these events were dramatic, eukaryotic life persisted through them, and fossil evidence shows that eukaryotes thrived during the c. 30-million-year interlude between the glaciations. Carbonate successions have become an important taphonomic window for this interval. One of the most notable examples is the c. 662–635 Ma Taishir Formation (Tsagaan Olom Group, Zavkhan Terrane, Mongolia) which has yielded a number of eukaryotic fossil taxa. Here, we examine more closely the morphology and taxonomic affinity of some of these Taishir fossils previously interpreted as remains of ciliate tintinnid loricae (purportedly the oldest fossil ciliates). New morphological and ultrastructural analyses indicate that these fossils are not ciliate tintinnids. Instead, we propose a new interpretation: that they are algal reproductive structures related to coeval macroscopic organic warty sheets described as putative red algae. We report the first occurrence of these fossils in the earliest Ediacaran Ol Formation, indicating that this taxon persisted through the Marinoan Snowball Earth. A new interpretation of these fossils as putative red algal spores has broad implications for our understanding of biodiversity in the Neoproterozoic Era, specifically during the Cryogenian Period, and for the antiquity of ciliates.     

Digging deeper: why we need more Proterozoic algal fossils and how to get them

Known Proterozoic algal fossils raise compelling questions about the origin and diversification of cyanobacteria and eukaryotic algae, and their ecological influence in deep time. This Perspectives article describes particular examples of persistent evolutionary and biogeochemical issues whose resolution would be aided by additional algal fossil evidence from Proterozoic deposits, which have been the subjects of recent intensive study. New Proterozoic geosciences literature relevant to the early diversification of algae is surveyed. Previously underappreciated algal traits that might improve taxonomic attributions of fossil remains are highlighted. Processes that phycologists could use to improve detection of algal fossils are recommended. Potential geological sources of new Proterozoic fossils are suggested.     

Production of volatile sulfur compounds during the decomposition of algal mats.

Blue-green algal mats incubated anaerobically rapidly produce large amounts of volatile sulfur compounds, including hydrogen sulfide, methyl mercaptan, and dimethyl sulfide. The major organic sulfur compound is methyl mercaptan, in contrast to previous results with marine eucaryotic algae. Light inhibited production of volatile sulfur compounds, apparently because the algae then produced O2, rendering the system aerobic.     

Production of volatile sulfur compounds during the decomposition of algal mats.

Blue-green algal mats incubated anaerobically rapidly produce large amounts of volatile sulfur compounds, including hydrogen sulfide, methyl mercaptan, and dimethyl sulfide. The major organic sulfur compound is methyl mercaptan, in contrast to previous results with marine eucaryotic algae. Light inhibited production of volatile sulfur compounds, apparently because the algae then produced O2, rendering the system aerobic.     

Suitability of a cytotoxicity assay for detection of potentially harmful compounds produced by freshwater bloom-forming algae

Detecting harmful bioactive compounds produced by bloom-forming pelagic algae is important to assess potential risks to public health. We investigated the application of a cell-based bioassay: the rainbow trout gill-w1 cytotoxicity assay (RCA) that detects changes in cell metabolism. The RCA was used to evaluate the cytotoxic effects of (1) six natural freshwater lake samples from cyanobacteria-rich lakes in central Ontario, Canada; (2) analytical standards of toxins and noxious compounds likely to be produced by the algal communities in these lakes; and (3) complex mixtures of compounds produced by cyanobacterial and chrysophyte cultures. RCA provided a measure of lake water toxicity that could not be reproduced using toxin or noxious compound standards. RCA was not sensitive to toxins and only sensitive to noxious compounds at concentrations higher than reported environmental averages (EC₅₀ ≥ 10³ nM). Cultured algae produced bioactive compounds that had recognizable dose dependent and toxic effects as indicated by RCA. Toxicity of these bioactive compounds depended on taxa (cyanobacteria, not chrysophytes), growth stage (stationary phase more toxic than exponential phase), location (intracellular more toxic than extracellular) and iron status (cells in high-iron treatment more toxic than cells in low-iron treatment). The RCA provides a new avenue of exploration and potential for the detection of natural lake algal toxic and noxious compounds.     

Limitations of using extracellular alkaline phosphatase activities as a general indicator for describing P deficiency of phytoplankton in Chinese shallow lakes

Extracellular phosphatase can be produced by phytoplankton to utilize organic phosphorus under phosphorus (P) deficiency. However, there is a controversy about its use as an indicator of P deficiency in natural phytoplankton community inferred by such an “induction–repression” mechanism. Size-fractionation of alkaline phosphatase activity (APA), soluble reactive phosphorus (SRP) concentration, algal density, and composition were determined in six Chinese shallow lakes ranking in gradient of trophic status, where a positive relationship between SRP concentration and algal density was observed. Enzyme-labeled fluorescence (ELF) method was used to localize phosphatase on cell membrane of algae. The so-called algal APA that associated with coarser particle (>3.0 μm) accounted for the largest part of total APA. Within a lake with lower SRP concentration, the “induction–repression” mechanism held true. Contrastingly, both algal and total APA were positively related to SRP concentration based on the data across all study lakes with statistical significance, which may be explained firstly by algal composition. The lakes with higher SRP concentration were dominated by diatoms and green algae, while they easily produced extracellular phosphatases as evidenced by ELFA labeling. In parallel, the lakes with lower SRP concentration were dominated by cyanobacteria, while it was never ELFA-positive; secondly, ELFA-positive dots or structures suggested that, in lakes with higher trophic status, attached bacteria or heterotrophic microorganisms could substantially contribute to extracellular phosphatases for hydrolyzing organophosphoric compounds but probably utilizing the organic moiety as an organic carbon source. This process simultaneously produces inorganic P, leading to the co-occurrence of high phosphate concentration and APA. So, the contributor of APA are complex, which may produce extracellular phosphatase species-specific or not exclusively for P nutrient and consequently make it difficult to normalize APA with the exact biomass estimators. Therefore, it is not reasonable to use APA, normalized or not, as a general indicator for describing P deficiency of phytoplankton in shallow lakes especially eutrophic ones.     

Combining geometric morphometrics,molecular phylogeny,and micropaleontology to assess evolutionary patterns in Mallomonas (Synurophyceae: Heterokontophyta)

Synurophytes, also known as scaled chrysophytes, are ecologically important algae that produce an array of siliceous structures upon which their taxonomy is based. Despite occupying a key position within the photosynthetic heterokonts, the evolutionary history of synurophytes remains poorly constrained. Here, modern and Middle Eocene siliceous scales of the morphotaxon Mallomonas insignis are used as a model to investigate synurophyte evolutionary patterns. Structural details of scale morphology were examined comparatively with scanning electron microscopy and scored for geometric morphometric analyses to assess the stability of shape characters. Although consistent size differences exist (modern scales are larger than Eocene counterparts), the populations cannot be differentiated on the basis of shape or microstructural detail, implying considerable evolutionary stasis in scale morphology. A time‐calibrated relaxed molecular clock analysis using a three‐gene concatenated data set (27 strains) suggests that the M. insignis lineage predates the available fossil record, having diverged from closest congeneric taxa in the Cretaceous (≥94 Ma). However, the molecular analysis also implies that considerable genetic variability is present within several morphotaxa of Mallomonas, implying that substantial genetic variability has arisen despite the retention of uniform scale morphologies, and resulting in the widespread occurrence of cryptic taxa. Results from the synurophyte lineage are consistent with the notion of protracted ghost ranges (>10 Ma) implied by the molecular phylogenies of other algal groups, together pointing to the paucity of the fossil record of these organisms on these timescales.     

CYANOPHYTA-BACTERIA SYSTEMS: EFFECTS OF ADDED CARBON COMPOUNDS OR PHOSPHATE ON ALGAL GROWTH AT LOW NUTRIENT CONCENTRATIONS1

Planktonic blue-green algae are known to be always associated with bacteria. Earlier work has shown that the addition of a bacteria-assimilable carbon source to a normal Zehnder-Gorham culture medium (No. 11) will produce enhanced growth of these algae when atmospheric CO₂ has become the limiting factor. In new work, phosphate-rich culture media were diluted so that they simulated nutrient levels found in the Great Lakes, e.g., Lake Erie. At these low concentrations and when atmospheric CO₂ was not available in a sufficient supply, the addition of sucrose to either a 1/100 or a 1/1000 diluted Z-G, medium (10 mg or 2 mg of sucrose, respectively) also produced enhanced, growth of the tested blue-green algae. The stimulation of algal growth was apparently due to an increased bacterial production of CO₂ and possibly other carbon compounds approaching the composition of the CO₂ molecule. The literature suggests that during vigorous algal growth in lakes, atmospheric CO₂ will be severely limiting. Productive lakes always contain nonliving organic matter. The presence of bacteria-assimilable matter is probably one of the important factors leading to algal bloom.     

Territorial behavior of threespot damselfish (Eupomacentrus planifrons) increases reef algal biomass and productivity

Synopsis Algal growth and damselfish (Eupomacentrus planifrons) territories were studied in two reef habitats at Discovery Bay, Jamaica. Damselfish territories were contiguous in the reef flat (0 to 2.5 m), where the algal composition and biomass varied from territory to territory. In contrast, on the lower reef terrace (22 m), damselfish territories were often spatially segregated. While the algal composition of the territories was more uniform on the reef terrace, the total algal biomass was lower than in the territories on the reef flat. Damselfish are largely herbivorous, and they defend their territories against most intruding fish, including a number of herbivorous species. Areas of the reef terrace outside of damselfish territories were heavily grazed by herbivorous fishes and contained only small quantities of non-crustose algae.The reef terrace territories were characterized by a multispecific turf of algae (greens, blue-greens, and reds) covering the Acropora cervicornis framework and by the leafy, brown alga, Lobophora variegata. A rapid reduction in the biomass of brown algae and filamentous algae was noted when damselfish were permanently removed from their territories. Only calcified, encrusting algae — plants apparently somewhat undesirable as fish food sources — would be common on the terrace zone of this reef if damselfish territories were absent. Damselfish territoriality may significantly influence the dynamics of some reefs by increasing the biomass of the algal turf thereby increasing; reef productivity. Since blue-green algae, potential nitrogen fixers, occur in these algal turfs, the fish may also be indirectly affecting reef nutrition.     

A metabarcoding comparison of windward and leeward airborne algal diversity across the Ko‘olau mountain range on the island of O'ahu,Hawai‘i1

Airborne algae from sites on the windward (n = 3) and leeward (n = 3) sides of the Ko‘olau Mountain range of O‘ahu, Hawai‘i, were sampled for a 16 d period during January and February 2015 using passive collection devices and were characterized using Illumina MiSeq sequencing of the universal plastid amplicon marker. Amplicons were assigned to 3,023 operational taxonomic units (OTUs), which included 1,189 cyanobacteria, 1,009 heterotrophic bacteria, and 304 Eukaryota (of which 284 were algae and land plants). Analyses demonstrated substantially more OTUs at windward than leeward O‘ahu sites during the sampling period. Removal of nonalgal OTUs revealed a greater number of algal reads recovered from windward (839,853) than leeward sites (355,387), with the majority of these being cyanobacteria. The 1,234 total algal OTUs included cyanobacteria, diatoms, cryptophytes, brown algae, chlorophyte green algae, and charophyte green algae. A total of 208 algal OTUs were identified from leeward side samplers (including OTUs in common among samplers) and 1,995 algal OTUs were identified from windward samplers. Barcoding analyses of the most abundant algal OTUs indicated that very few were shared between the windward and leeward sides of the Ko‘olau Mountains, highlighting the localized scale at which these airborne algae communities differ. Back trajectories of air masses arriving on O‘ahu during the sampling period were calculated using the NOAA HY‐SPLIT model and suggested that the sampling period was composed of three large‐scale meteorological events, indicating a diversity of potential sources of airborne algae outside of the Hawaiian Islands.     

Trophodynamics of the sclerosponge Ceratoporella nicholsoni along a shallow to mesophotic depth gradient

The sclerosponge Ceratoporella nicholsoni is a hyper-calcifying high microbial abundance sponge. This sponge has been observed at high densities throughout the Caribbean in the mesophotic zone (30–150 m), as well as cryptic environments in shallow (< 30 m) depths. Given the densities of this sponge, it could play an important role in the cycling of inorganic and organic sources of carbon and nitrogen at mesophotic depths. Additionally, there is broad interest in this sponge as a tool for paleobiology, paleoclimatology and paleoceanography. As a result, it is increasingly important to understand the ecology of these unique sponges in the underexplored Caribbean mesophotic zone. Here we show that this sponge increases in abundance from shallow depths into the mesophotic zone of Grand Cayman Island. We observed no significant differences in the stable isotope signatures of δ¹⁵N and δ¹³C of sponge tissue between depths. A predictive model of sponge diet with increasing depth shows that these sponges consume dissolved organic matter of algal and coral origin, as well as the consumption of particulate organic matter consistent with the interpretation of the stable isotope data. The taxonomic composition of the sclerosponge microbiome was invariant across the shallow to mesophotic depth range but did contain the Phylum Chloroflexi, known to degrade a variety of dissolved organic carbon sources. These data suggest that the depth distribution of this sponge may not be driven by changes in trophic strategy and is potentially regulated by other biotic or abiotic factors.

     

Harmful algal blooms of allelopathic microalgal species: The role of eutrophication

The ability of certain harmful algal species to produce and release chemicals that inhibit the growth of co-occurring phytoplankton species, here considered as allelopathy, is closely associated with competition for limiting nutrient resources. Many phytoplankton cells are known to release elevated amounts of organic compounds under nutrient limitation. Eutrophication alters the nitrogen-to-phosphorus balance and, when nutrient availability is unbalanced, nutrient limitation may result. Algal species that can compete successfully for available growth-limiting nutrient(s) have the potential to become dominant and form blooms. The stress conditions imposed by the shifted nutrient supply ratios can, in some algae, stimulate production of allelochemicals that inhibit potential competitors. Thus, under cultural eutrophication, altered nutrient (N, P) ratios and limiting nutrient supplies can stimulate increased production of allelochemicals, including toxins, by some algal species and accentuate the adverse effects of these substances on other algae. Future investigation on the characterization of the chemical compounds involved in the allelopathic process are needed to advance the study of the mode of action of phytoplankton allelochemicals.     

Tropical freshwater teleosts from Miocene beds of eastern Patagonia, southern Argentina

Loricariid and pimelodid-like siluriforms—along with undetermined percomorph—are described for the first time from Patagonia. Vertebrate fossils, including fish, mammals and birds, were found in beds attributed to the top of the Puerto Madryn Formation. These levels supposedly corresponded to the “Rionegrense marino” of former authors. Mammals occurring in the site suggest a Huayquerian age for the fossiliferous beds. The base of the Huayquerian was dated at about 9 Ma and the top is younger than 6.5 Ma. Radioisotopic dating in the marine shell beds of the Puerto Madryn Formation ranges from 11 to 9 Ma. Consequently, the section described here appears to be younger than the typical Puerto Madryn Formation from which it is separated by an unconformity. The section is correlated with the type Río Negro Formation from northern Patagonia, which also includes Huayquerian fossils. Freshwater fishes were previously unknown in beds younger than the middle Miocene in southern South America. This is also the southernmost record of loricariid fishes. The association of aquatic continental and terrestrial vertebrates indicates for the first time in Península Valdés beds of freshwater origin. The evidence apported by fossils is also in agreement with global climate trends. The local extinction of loricariids in Patagonia possibly occurred much later than the time of deposition.     

Algae as promising organisms for environment and health

Algae, like other plants, produce a variety of remarkable compounds collectively referred to as secondary metabolites. They are synthesized by these organisms at the end of the growth phase and/or due to metabolic alterations induced by environmental stress conditions. Carotenoids, phenolic compounds, phycobiliprotein pigments, polysaccharides and unsaturated fatty acids are same of the algal natural products, which were reported to have variable biological activities, including antioxidant activity, anticancer activity, antimicroabial activity against bacteria-virus-algae-fungi, organic fertilizer and bioremediation potentials.Key words: algae, biological activities, active ingredients     

Vicariance and dispersal in southern hemisphere freshwater fish clades: a palaeontological perspective

Widespread fish clades that occur mainly or exclusively in fresh water represent a key target of biogeographical investigation due to limited potential for crossing marine barriers. Timescales for the origin and diversification of these groups are crucial tests of vicariant scenarios in which continental break‐ups shaped modern geographic distributions. Evolutionary chronologies are commonly estimated through node‐based palaeontological calibration of molecular phylogenies, but this approach ignores most of the temporal information encoded in the known fossil record of a given taxon. Here, we review the fossil record of freshwater fish clades with a distribution encompassing disjunct landmasses in the southern hemisphere. Palaeontologically derived temporal and geographic data were used to infer the plausible biogeographic processes that shaped the distribution of these clades. For seven extant clades with a relatively well‐known fossil record, we used the stratigraphic distribution of their fossils to estimate confidence intervals on their times of origin. To do this, we employed a Bayesian framework that considers non‐uniform preservation potential of freshwater fish fossils through time, as well as uncertainty in the absolute age of fossil horizons. We provide the following estimates for the origin times of these clades: Lepidosireniformes [125–95 million years ago (Ma)]; total‐group Osteoglossomorpha (207–167 Ma); Characiformes (120–95 Ma; a younger estimate of 97–75 Ma when controversial Cenomanian fossils are excluded); Galaxiidae (235–21 Ma); Cyprinodontiformes (80–67 Ma); Channidae (79–43 Ma); Percichthyidae (127–69 Ma). These dates are mostly congruent with published molecular timetree estimates, despite the use of semi‐independent data. Our reassessment of the biogeographic history of southern hemisphere freshwater fishes shows that long‐distance dispersals and regional extinctions can confound and erode pre‐existing vicariance‐driven patterns. It is probable that disjunct distributions in many extant groups result from complex biogeographic processes that took place during the Late Cretaceous and Cenozoic. Although long‐distance dispersals likely shaped the distributions of several freshwater fish clades, their exact mechanisms and their impact on broader macroevolutionary and ecological dynamics are still unclear and require further investigation.     

Nutritional value of detritus and algae in blenny territories on the Great Barrier Reef

The salariin blenny, Salarias patzneri, primarily ingests detrital aggregates and inorganic sediment, with small quantities of filamentous algae. Samples of particulate matter <125 μm and filamentous algae were collected from the territories of S. patzneri and the nutritional value of these resources assessed using organic content and protein/energy ratios calculated from protein, carbohydrate and lipid content. Samples were collected throughout the day, during both summer and winter, to enable temporal comparisons of nutritional value. The particulates <125 μm, which were predominantly amorphic detritus, accounted for 41±2% and 44±3% of the organic matter in the epilithic algal matrix (EAM) during the summer and winter, respectively, whilst filamentous algae accounted for 37±3% and 41±3% of the organic matter in the summer and winter, respectively. Carbohydrate and lipid concentration in the algal samples was significantly greater than in particulate matter <125 μm, although no significant difference was detected in protein levels of algal samples and particulates <125 μm. Consequently, the protein/energy ratio of particulates <125 μm in the summer (11±1 mg kJ⁻¹) and winter (10±1 mg kJ⁻¹) was similar to that of filamentous algae in the summer (9±1 mg kJ⁻¹) and winter (8±1 mg kJ⁻¹). These results suggest detrital aggregates are at least comparable to filamentous algae as a nutritional resource. The large contribution of high quality detritus to organic matter in the epilithic algal matrix collected from S. patzneri territories throughout the day and between seasons provides strong evidence that detritus is a valuable component of small territorial fish diets and is an integral part of coral reef food webs.     

Quantitative and qualitative analyses of dissolved organic matter released from Ecklonia cava Kjellman, in Oura Bay, Shimoda, Izu Peninsula, Japan

Release of dissolved organic matter (DOM) was studied on macroalga, Ecklonia cava Kjellman, by in situ field bag experiments, which were designed to keep the algal body under natural field condition, in Oura Bay, Shimoda, Izu Peninsula, Japan, from August 2003 to May 2005. The experiments were conducted 6 times in different seasons. The concentrations of dissolved organic carbon (DOC) in the experimental bag generally increased during the first 1-2 days, showing the release of DOC from algae. The daily release rates of DOC varied between 0.12 and 5.8 mgC per g (dry wt) of the algal blades per day. The seasonal variability of the DOC release rate was similar to those of the growth and the photosynthetic rates reported by the previous studies on E. cava in the same location. The fractions of the DOC release of the net primary production accounted for 18-62%. These contributions are comparable to other macroalgae in the previous studies (27-43%) using other macroalgae. Analysis of organic composition of DOM using gas chromatography (neutral aldose composition) and spectrometry (UV-visible and fluorescence) indicates that E. cava usually release extracellular products containing mucilaginous polysaccharides containing fucose as a main constituent and colored dissolved organic matter (CDOM). The release of these organic compounds is likely performed as the ordinary metabolism of brown algae. An exceptional release of a considerable amount of protein and carbohydrates mainly composed of mannose was found in June 2004, probably reflecting leakage of the intracellular material by an autolysis of the algal cells. The high DOM release rate in the present study suggests that macroalgae would be one of the important DOM producers in coastal regions.     

Discovery of the mineral brucite (magnesium hydroxide) in the tropical calcifying alga Polystrata dura (Peyssonneliales,Rhodophyta)

Red algae of the family Peyssonneliaceae typically form thin crusts impregnated with aragonite. Here, we report the first discovery of brucite in a thick red algal crust (~1 cm) formed by the peyssonnelioid species Polystrata dura from Papua New Guinea. Cells of P. dura were found to be infilled by the magnesium‐rich mineral brucite [Mg(OH)₂]; minor amounts of magnesite and calcite were also detected. We propose that cell infill may be associated with the development of thick (> ~5 mm) calcified red algal crusts, integral components of tropical biotic reefs. If brucite infill within the P. dura crust enhances resistance to dissolution similarly to crustose coralline algae that infill with dolomite, then these crusts would be more resilient to future ocean acidification than crusts without infill.     

Giving the early fossil record of sponges a squeeze

Twenty candidate fossils with claim to be the oldest representative of the Phylum Porifera have been re‐analysed. Three criteria are used to assess each candidate: (i) the diagnostic criteria needed to categorize sponges in the fossil record; (ii) the presence, or absence, of such diagnostic features in the putative poriferan fossils; and (iii) the age constraints for the candidate fossils. All three criteria are critical to the correct interpretation of any fossil and its placement within an evolutionary context. Our analysis shows that no Precambrian fossil candidate yet satisfies all three of these criteria to be a reliable sponge fossil. The oldest widely accepted candidate, Mongolian silica hexacts from c. 545 million years ago (Ma), are here shown to be cruciform arsenopyrite crystals. The oldest reliable sponge remains are siliceous spicules from the basal Cambrian (Protohertzina anabarica Zone) Soltanieh Formation, Iran, which are described and analysed here in detail for the first time. Extensive archaeocyathan sponge reefs emerge and radiate as late as the middle of the Fortunian Stage of the Cambrian and demonstrate a gradual assembly of their skeletal structure through this time coincident with the evolution of other metazoan groups. Since the Porifera are basal in the Metazoa, their presence within the late Proterozoic has been widely anticipated. Molecular clock calibration for the earliest Porifera and Metazoa should now be based on the Iranian hexactinellid material dated to c. 535 Ma. The earliest convincing fossil sponge remains appeared at around the time of the Precambrian‐Cambrian boundary, associated with the great radiation events of that interval.