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Calcium oxalate crystals persist in Antithamnion thalli cultured in artificial seawater medium (ASW) containing reduced calcium concentrations or in darkness. Differential interference contrast microscopy. Fig. 9. Antithamnion antillanum. Thalli grown in ASW containing 2.5 mM CaCl 2 released spores, and the young sporelings began to form calcium oxalate crystals (arrows) in the axial cells. Treated in sodium hypochlorite to remove organic material. Fig. 10. Antithamnion sparsum. Thalli grown for three weeks in ASW containing only 1 mM CaCl 2 still had calcium oxalate crystals in axial cells. Treated in sodium hypochlorite. Fig. 11. Antithamnion sparsum. Starch grains are absent but calcium oxalate crystals are abundant among the chloroplasts in the parietal cytoplasm of thalli maintained in complete darkness for 14 weeks.  

Calcium oxalate crystals persist in Antithamnion thalli cultured in artificial seawater medium (ASW) containing reduced calcium concentrations or in darkness. Differential interference contrast microscopy. Fig. 9. Antithamnion antillanum. Thalli grown in ASW containing 2.5 mM CaCl 2 released spores, and the young sporelings began to form calcium oxalate crystals (arrows) in the axial cells. Treated in sodium hypochlorite to remove organic material. Fig. 10. Antithamnion sparsum. Thalli grown for three weeks in ASW containing only 1 mM CaCl 2 still had calcium oxalate crystals in axial cells. Treated in sodium hypochlorite. Fig. 11. Antithamnion sparsum. Starch grains are absent but calcium oxalate crystals are abundant among the chloroplasts in the parietal cytoplasm of thalli maintained in complete darkness for 14 weeks.  

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A survey of 18 species of the Ceramiales grown in culture revealed calcium oxalate crystals in Antithamnion antillanum Børgesen, A. callocladum Itono, and A. sparsum Tokida. The needle-shaped crystals were present within the cytoplasm of cells of the indeterminate axes but not in cells of the determinate lateral branches. No such crystals were pres...

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... of more mature thalli, the apical cell and two or three expanding cells subtending it did not have crystals. However, by the time five axial cells were produced, calcium oxalate crystals were visible in some of the older of these cells. Even in thalli grown in 2.5 mM CaCl 2 , the new elongate cells of the indeterminate axis formed crystals (Fig. 9). After exposure to ASW containing 1.0 mM CaCl 2 for only five days, thalli of A. nipponicum showed loss of colour in scattered cells of both indeterminate axes and lateral branches. However, cell death was not pervasive; in response to loss of their subtending axial cells, the basal cells of the determinate lateral branches were ...

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... Pennington also grew filaments under different wavelengths of light and reported that complete darkness and nearly all colours of monochromatic light caused fragmentation and dissolution of calcium oxalate crystals. However, observations of S. hatillensis (C.M. Pueschel, unpublished) and several species of the red alga Antithamnion (Pueschel & West 2007c) revealed the presence of calcium oxalate crystals after weeks of complete darkness. ...
... In specimens prepared by freeze substitution, microfilaments were found to partially ensheath the crystals (Babuka & Pueschel 1998). Pueschel & West (2007c) reported that acicular crystals were also present in Antithamnion antillanum, A. callocladum, and A. sparsum. The effects of a wide range of calcium concentrations were tested on these species and A. nipponicum Yamada & Inagaki, which does not produce calcium oxalate crystals. ...
... In a study of Antithamnion in South Africa, Norris (1987) reported needle-like crystals of unknown composition in axial cells of A. diminuatum, A. secundum, and A. antillanum. Given the crystals' acicular morphology, anatomical distribution, and concurrence of these findings on A. antillanum with those of Pueschel & West (2007c), there is little doubt that the structures described were calcium oxalate crystals. ...
Article
The morphology, anatomical and cellular localisation, abundance, and taxonomic distribution of calcium oxalate deposits in algae are described. Comparisons are drawn to the same features of calcium oxalate crystals in embryophytes. A review of the early history of work on these mineral deposits in algae reveals a wider distribution of taxa and greater research activity than has generally been recognised in modern literature. Documentation of the presence of calcium oxalate in Vaucheria (Vaucheriales), the sole genus of Ochrophyta known to deposit this mineral, is provided for the first time. Additional species of Dasycladales are reported to be calcium oxalate mineralisers. Although calcium oxalate crystals are a consistent feature of some species, they appear only sporadically in others. It is proposed that three different mechanisms of calcium oxalate deposition operate amongst the algae: (1) unregulated, spontaneous precipitation of stored oxalic acid; (2) constitutive mineralisation involving an organic matrix that determines crystal form; and (3) regulation of crystal form by soluble agents that affect nucleation or deposition on the mineral surface. Members of Bryopsidales that deposit acicular calcium oxalate crystals do so constitutively and are not accumulators of oxalic acid; the opposite is true of many Dasycladales, which deposit calcium oxalate in the bipyramidal form as a result of spontaneous precipitation of oxalic acid. Several species of Spirogyra (Zygnematales) are the only representatives of the Charophyceae known to deposit calcium oxalate; their crystals are cruciate, have an organic matrix, and appear to be constitutive. Among red algae, some species of Antithamnion (Ceramiales) deposit acicular crystals constitutively; whereas, sequestered oxalic acid in Spyridia (Ceramiales) can precipitate to form bipyramidal crystals. Calcium oxalate–depositing members of Cladophorales are numerous; crystal morphologies and perhaps their mode of deposition vary. Vaucheria deposits morphologically diverse intracellular calcium oxalate but was also found to produce bipyramidal crystals by spontaneous precipitation. Monitoring acid dissolution of calcium oxalate by microscopic observation provides insights into crystal structure. Possible ecological and physiological functions of calcium oxalate crystals and of soluble oxalate production in algae are evaluated.
... It has been proposed that these inclusions might serve as a seasonal nitrogen store (Pueschel 1992), but this idea has not been tested in red algae. Calcium oxalate crystals are common in higher plants and are present in some algal groups, including red algae (Pueschel 1995), but the physiological functions usually assigned to such inclusions in higher plants are unlikely to apply to the algae (Pueschel and West 2007). Progress has been made in the characterization of refractile inclusions that are associated with some kinds of specialized vegetative cells (Paul et al. 2006) and can form distinctive structures, such as the corps en cerise in cortical cells of Laurencia (Reis et al. 2013). ...
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Macroalgae in mesophotic coral ecosystems are generally understudied compared to corals and fishes yet may be more abundant than coral-dominated reefs given their lower depth limits (> 200 m) and ability to grow over soft and hard bottom habitats. These assemblages are abundant and diverse globally, with changing species composition with increasing depth. Ubiquitous macroalgal assemblages include the red algal rhodolith beds and nongeniculate and Peyssonneliales assemblages; green algal Halimeda beds, meadows, and bioherms and Caulerpa spp. beds; and brown algal Lobophora spp. or Distromium spp. beds, Sargassum spp., and kelps. The use of molecular techniques is elucidating macroalgal diversity and rates of endemism, and molecular data and phylogenetic analyses often show strong cryptic diversity or pseudodiversity when compared with morphoanatomical analyses. Mesophotic macroalgae are important as habitat and may serve as seedbanks or refugia for ecosystem resilience following environmental stress. Invasive algal blooms may be deleterious, particularly with the removal of native herbivores or increasing nutrients. Geomorphologically, calcified species such as rhodoliths and Halimeda spp. are significant global producers of calcium carbonate. Abiotic factors influencing the abundance and distribution of mesophotic macroalgae include temperature, water clarity, nutrients, and currents. In general, threats include rhodolith mining, oil spills, sedimentation, ocean acidification, invasive species, bottom trawling, and eutrophication. The impacts of global warming at mesophotic depths are unknown. Future studies should focus on collections for molecular analyses to evaluate population-level dynamics and connectivity between shallow and mesophotic depths and in situ manipulations to determine competitive interactions and ecophysiological processes in these low-light environments.
... It has been proposed that these inclusions might serve as a seasonal nitrogen store (Pueschel 1992), but this idea has not been tested in red algae. Calcium oxalate crystals are common in higher plants and are present in some algal groups, including red algae (Pueschel 1995), but the physiological functions usually assigned to such inclusions in higher plants are unlikely to apply to the algae (Pueschel and West 2007). Progress has been made in the characterization of refractile inclusions that are associated with some kinds of specialized vegetative cells (Paul et al. 2006) and can form distinctive structures, such as the corps en cerise in cortical cells of Laurencia (Reis et al. 2013). ...
Chapter
Rhodophyta, or red algae, comprises a monophyletic lineage within Archaeplastida that includes glaucophyte algae and green algae plus land plants. Rhodophyta has a long fossil history with evidence of Bangia-like species in ca. 1.2 billion-year-old deposits. Red algal morphology varies from unicellular, filamentous, to multicellular thalloid forms, some of which are sources of economically important products such as agar and carrageenan. These species live primarily in marine environments from the intertidal zone to deep waters. Freshwater (e.g., Batrachospermum) and terrestrial lineages also occur. One of the major innovations in the Rhodophyta is a triphasic life cycle that includes one haploid and two diploid phases with the carposporophyte borne on female gametophytes. Red algae are also well known for their contribution to algal evolution with ecologically important chlorophyll-c containing lineages such as diatoms, dinoflagellates, haptophytes, and phaeophytes all containing a red algal-derived plastid of serial endosymbiotic origin. Analysis of red algal nuclear genomes shows that they have relatively small gene inventories of 6,000–10,000 genes when compared to other free-living eukaryotes. This is likely explained by a phase of massive genome reduction that occurred in the red algal ancestor living in a highly specialized environment. Key traits that have been lost in all red algae include flagella and basal body components, light-sensing phytochromes, and the glycosylphosphatidylinositol (GPI)-anchor biosynthesis and macroautophagy pathways. Research into the biology and evolution of red algae is accelerating and will provide exciting insights into the diversification of this unique group of photosynthetic eukaryotes.
... For instance, needle-like crystals have been reported in the vacuoles of the siphonous green algae Penicillus (Friedmann et al., 1972) and Chlorodesmis (Ducker, 1967). Such crystals inclusions were also found in red algae, e.g. in Antithamnion kylinii (Pueschel, 1995) and Spyridia filamentosa (Pueschel and West, 2007). The diversity of crystal types in algae has been shown to be of systematic importance. ...
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Calcium oxalate is the most widely distributed inorganic crystal in plants, occurring in over 200 plant families. This wide distribution suggests that they constitute an important biomineralization process in plants. The variation in CaOx crystal shape and cell types producing them indicates that crystals may have evolved many times independently in different plant lineages and may probably serve multiple functions. Their distribution among lycophytes and ferns is poorly documented and has led to the assumption that they are rare or absent in these lineages. Only a few reports indicate that CaOx crystals are present in several fern families, but their distribution in this group has never been assessed on a broad scale. Our aim was to determine and compare shape, abundance as well as distribution patterns of calcium oxalate crystals across ferns and lycophytes. Such knowledge is crucial to infer hypotheses about the structural-functional evolution of CaOx-formation in land plants. Our observations were plotted on a phylogenetic tree in order to test whether certain patterns appear on different phylogenetic levels. It was also investigated whether distribution patterns or particular crystal types could be related to fern ecology or potential function(s).
... It has been proposed that these inclusions might serve as a seasonal nitrogen store (Pueschel 1992), but this idea has not been tested in red algae. Calcium oxalate crystals are common in higher plants and are present in some algal groups, including red algae (Pueschel 1995), but the physiological functions usually assigned to such inclusions in higher plants are unlikely to apply to the algae (Pueschel and West 2007). Progress has been made in the characterization of refractile inclusions that are associated with some kinds of specialized vegetative cells (Paul et al. 2006) and can form distinctive structures, such as the corps en cerise in cortical cells of Laurencia (Reis et al. 2013). ...
Chapter
Rhodophyta, or red algae, comprises a monophyletic lineage within Archaeplastida that includes glaucophyte algae and green algae plus land plants. Rhodophyta has a long fossil history with evidence of Bangia-like species in ca. 1.2 billion-year-old deposits. Red algal morphology varies from unicellular, filamentous, to multicellular thalloid forms, some of which are sources of economically important products such as agar and carrageenan. These species live primarily in marine environments from the intertidal zone to deep waters. Freshwater (e.g., Batrachospermum) and terrestrial lineages also occur. One of the major innovations in the Rhodophyta is a triphasic life cycle that includes one haploid and two diploid phases with the carposporophyte borne on female gametophytes. Red algae are also well known for their contribution to algal evolution with ecologically important chlorophyll-c containing lineages such as diatoms, dinoflagellates, haptophytes, and phaeophytes all containing a red algal-derived plastid of serial endosymbiotic origin. Analysis of red algal nuclear genomes shows that they have relatively small gene inventories of 6,000–10,000 genes when compared to other free-living eukaryotes. This is likely explained by a phase of massive genome reduction that occurred in the red algal ancestor living in a highly specialized environment. Key traits that have been lost in all red algae include flagella and basal body components, light-sensing phytochromes, and the glycosylphosphatidylinositol (GPI)-anchor biosynthesis and macroautophagy pathways. Research into the biology and evolution of red algae is accelerating and will provide exciting insights into the diversification of this unique group of photosynthetic eukaryotes.
... No alga has been found to have raphides. Where needles (acicular crystals) have been found, they appear singly, not in bundles of needles; they are surrounded by a vacuolar membrane or crystal chamber but not both, are generally very small, or reside individually in the cytoplasm and not the cell vacuole [14][15][16][17][18]. Single needles also occur in the cell vacuole of both lightly calcified parts of otherwise heavily calcified seaweeds [19][20][21]. ...
... Pueschel [16] found small, cruciate crystals in the peripheral cytoplasm but not within the vacuole of the fresh water green alga, Spirogyra. Antithamnion, a marine red alga, had single needles up to 30 µm in length in the peripheral cytoplasm [15,18]. The length of the needles in Antithamnion, though shorter than those in C. minus, suggested that they may adversely affect herbivores [18], but the bipyramidal morphology of crystals (apparently surrounded by an organic matrix = the vacuolar membrane?) in Chaetomorpha, a marine green seaweed, probably negates their role in grazer defense [26]. ...
... Antithamnion, a marine red alga, had single needles up to 30 µm in length in the peripheral cytoplasm [15,18]. The length of the needles in Antithamnion, though shorter than those in C. minus, suggested that they may adversely affect herbivores [18], but the bipyramidal morphology of crystals (apparently surrounded by an organic matrix = the vacuolar membrane?) in Chaetomorpha, a marine green seaweed, probably negates their role in grazer defense [26]. Other papers [14,15,17] also describe needles, in some cases more than 100 µm long. ...
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The vacuole of utricles, the outermost cell layer of the siphonous green seaweed, Codium minus, had numerous single needles and needle bundles. The crystals composing each needle appeared arranged in a twisted configuration, both ends were pointed, and each needle was contained in a matrix or membrane; bundles of needles appeared enclosed by a matrix. Chemical and electron diffraction analysis indicated that the needles consisted of calcium oxalate. This is the first paper on terrestrial plant-like raphides in an alga.
... Calcium oxalate crystals in the bryopsidalean green seaweed Callipsygma wilsonis J. Agardh are triangular plates that are located within the cytoplasmic layer and move with streaming cytoplasm (Pueschel & West 2007b). Among the red algae, needle-like calcium oxalate crystals are known from several species of Antithamnion (Pueschel 1995;Pueschel & West 2007a), and the membrane-bounded crystals are embedded within the cytoplasmic layer. ...
... Embryophytes deposit calcium oxalate within membranous crystal chambers located inside the central vacuole of specialized idioblast cells (Francheschi & Horner 1980). The association of calcium oxalate crystals in C. coliformis with the periphery of the cytoplasm but not within the central vacuole could be viewed as an intermediate condition between the intravacuolar crystal chambers of embryophytes and the membrane-bounded crystals immersed within the cytoplasm of S. hatillensis (Pueschel 2001), Callipsygma wilsonis (Pueschel & West 2007b), and several species of the red algal genus Antithamnion (Pueschel 1995;Pueschel & West 2007a). ...
Article
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Living cells of field-collected specimens of the giant-celled marine green alga Chaetomorpha coliformis (Montagne) Kutzing were found to have birefringent cellular inclusions whose composition was determined to be calcium oxalate on the basis of their reactions to diagnostic chemical solubility tests and the Yasue cytochemical staining procedure. The inclusions consisted of individual bipyramidal crystals up to 50 mu m in greatest dimension and variously sized aggregates of much smaller crystals. Some aggregates consisted of rosettes or spheres of radiating elements that resemble embryophyte druses more than any other calcium oxalate deposit yet reported in the algae. Both single crystals and aggregates occurred in the same cells with no discernible pattern to their distributions. The calcium oxalate crystals were anchored on the vacuolar face of the thin cytoplasmic layer rather than being dispersed within the voluminous central vacuole. Light and transmission electron microscopy demonstrated the presence of a nearly continuous layer of small vacuoles between the organelle-rich parietal cytoplasm and the large central vacuole, and the calcium oxalate crystals were associated with this layer of vacuoles. The occurrence of mineralization in the parietal vacuoles, but not the large central vacuole, indicates that differentiation of vacuoles according to ionic contents may be occurring.
... It turns out, however, that calcium oxalate crystals occur more widely within the Plantae, occurring in a number of coenocytic green ulvophycean algae (Pueschel & West 2007a, and references therein) and florideophyte red algae (Pueschel 1995, Pueschel & West 2007b2007c, as well as in the microscopic filamentous green (charophycean) algal filament Spirogyra hatillensis . Oxalic acid synthesis does not invariably involve calcium oxalate precipitation, since oxalate anions contributes about 20% to the vacuolar anionic charge in the giant-celled marine ulvophycean Acetabularia mediterranea (Saddler 1970) yet this organism has not been reported as precipitating calcium oxalate.. Little functional research has been done on calcium oxalates in algae, but the functions proposed for vascular plants by Franceschi and Nakati (2005) are at least plausible for other members of the Plantae. ...
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Organismic and Evolutionary Biology Skeletal biomineralisation by microbial eukaryotes significantly affects the global biogeochemical cycles of carbon, silicon and calcium. Non-skeletal biomineralisation by eukaryotic cells, with precipitates retained within the cell interior, can duplicate some of the functions of skeletal minerals, e.g., increased cell density, but not the mechanical and antibiophage functions of extracellular biominerals. However, skeletal biomineralisation does not duplicate many of the functions of non-skeletal biominerals. These functions include magnetotaxis (magnetite), gravity sensing (intracellular barite, bassanite, celestite and gypsum), buffering and storage of elements in an osmotically inactive form (calcium as carbonate, oxalate, polyphosphate and sulfate; phosphate as polyphosphate) and acid-base regulation, disposing of excess hydroxyl ions via an osmotically inactive product (calcium carbonate, calcium oxalate). Although polyphosphate has a wide phylogenetic distribution among microbial eukaryotes, other non-skeletal minerals have more restricted distributions, and as yet there seems to be no definitive evidence that the alkaline earth components (Ba and Sr) of barite and celestite are essential for completion of the life cycle in organisms that produce these minerals.
... De acordo com Leliaert & Coppejans (2004), a ocorrência deste tipo particular de cristal em uma espécie indica que esta presença não é dependente do ambiente, sendo seu desenvolvimento geneticamente controlado e, portanto, com potencial para ser utilizado como característica diagnóstica. Pueschel & West (2007a), através de estudos experimentais com fornecimento de luz e cálcio em culturas de talos de Antithamnion Nägeli, demonstraram estabilidade na deposição de oxalato de cálcio em muitas espécies do gênero, sugerindo que estes cristais são específi cos. Por outro lado, Pueschel & West (2007b) demonstraram que em Spyridia fi lamentosa (Wulfen) Harv. a deposição de cristais de oxalato de cálcio é dependente do fornecimento de cálcio no meio, logo, esta característica deve ser usada com parcimônia na taxonomia desta espécie. ...
... raia de Cacha-Pregos, crescendo em poças próximas a árvores de mangue, parcialmente enterrada por sedimento arenoso/lamoso e na praia de Buraquinho, arribada. Foi encontrada associada à Chaetomorpha brachygona, C. aerea, Cladophoropsis membranacea, Dictyopteris delicatula, Acanthophora spicifera, Hypnea musciformis e a cianobactérias fi lamentosas.Price (1967) sugere que C. crassa e C. linum sejam co-específi cas, enquantoTaylor (1960),Nizamunddin & Begum (1973),Schneider & Searles (1991) eLittler & Littler (1997) as consideram como distintas. Segundo estes autores estas espécies não apresentam célula basal e crescem formando emaranhados. Chaetomorpha crassa é caracterizada por apresentar cél ...
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This paper presents a taxonomic study of the genera Chaetomorpha and Rhizoclonium occurring in the littoral of Bahia. The species were defined based on habit, presence or absence of lateral rhizoids, basal cell type, shape and cells dimensions. The presence of crystalline inclusions in cells was also verified. It were identified nine species, seven to the genus Chaetomorpha (C. aerea (Dillwyn) Kütz., C. antennina (Bory) Kütz., C. brachygona Harv., C. clavata Kütz., C. crassa (C. Agardh) Kütz., C. minima Collins & Herv. and C. nodosa Kütz.) and two to the genus Rhizoclonium (R. africanum Kütz. and R. riparium (Roth) Kütz. ex Harv.). Chaetomorpha crassa, C. minima and C. nodosa are new additions to the Northeastern of Brazil and C. aerea, C. clavata and R. africanum are additions to the Bahia. Three crystals types were found: 1. clusters of fine needle-shaped of silica crystals presents in C. aerea, C. antennina, C. brachygona, C. clavata and C. crassa; 2. octahedral of calcium oxalate crystals found in C. antennina and C. clavata and, 3. globular aggregates of cone-shaped calcium carbonate crystals observed in C. clavata. Fertile thallus of C. antennina, C. clavata, C. nodosa, R. africanum and R. riparium are being illustrated for the first time for the Brazilian coast. The vegetative and reproductive structures are described in detail and a comparison with similar species is provided.
... De acordo com Leliaert & Coppejans (2004), a ocorrência deste tipo particular de cristal em uma espécie indica que esta presença não é dependente do ambiente, sendo seu desenvolvimento geneticamente controlado e, portanto, com potencial para ser utilizado como característica diagnóstica. Pueschel & West (2007a), através de estudos experimentais com fornecimento de luz e cálcio em culturas de talos de Antithamnion Nägeli, demonstraram estabilidade na deposição de oxalato de cálcio em muitas espécies do gênero, sugerindo que estes cristais são específi cos. Por outro lado, Pueschel & West (2007b) demonstraram que em Spyridia fi lamentosa (Wulfen) Harv. a deposição de cristais de oxalato de cálcio é dependente do fornecimento de cálcio no meio, logo, esta característica deve ser usada com parcimônia na taxonomia desta espécie. ...
... raia de Cacha-Pregos, crescendo em poças próximas a árvores de mangue, parcialmente enterrada por sedimento arenoso/lamoso e na praia de Buraquinho, arribada. Foi encontrada associada à Chaetomorpha brachygona, C. aerea, Cladophoropsis membranacea, Dictyopteris delicatula, Acanthophora spicifera, Hypnea musciformis e a cianobactérias fi lamentosas.Price (1967) sugere que C. crassa e C. linum sejam co-específi cas, enquantoTaylor (1960),Nizamunddin & Begum (1973),Schneider & Searles (1991) eLittler & Littler (1997) as consideram como distintas. Segundo estes autores estas espécies não apresentam célula basal e crescem formando emaranhados. Chaetomorpha crassa é caracterizada por apresentar cél ...
Article
Full-text available
This paper presents a taxonomic study of the genera Chaetomorpha and Rhizoclonium occurring in the littoral of Bahia. The species were defined based on habit, presence or absence of lateral rhizoids, basal cell type, shape and cells dimensions. The presence of crystalline inclusions in cells was also verified. It were identified nine species, seven to the genus Chaetomorpha (C. aerea (Dillwyn) Kütz., C. antennina (Bory) Kütz., C. brachygona Harv., C. clavata Kütz., C. crassa (C. Agardh) Kütz., C. minima Collins & Herv. and C. nodosa Kütz.) and two to the genus Rhizoclonium (R. africanum Kütz. and R. riparium (Roth) Kütz. ex Harv.). Chaetomorpha crassa, C. minima and C. nodosa are new additions to the Northeastern of Brazil and C. aerea, C. clavata and R. africanum are additions to the Bahia. Three crystals types were found: 1. clusters of fine needle-shaped of silica crystals presents in C. aerea, C. antennina, C. brachygona, C. clavata and C. crassa; 2. octahedral of calcium oxalate crystals found in C. antennina and C. clavata and, 3. globular aggregates of cone-shaped calcium carbonate crystals observed in C. clavata. Fertile thallus of C. antennina, C. clavata, C. nodosa, R. africanum and R. riparium are being illustrated for the first time for the Brazilian coast. The vegetative and reproductive structures are described in detail and a comparison with similar species is provided.