Anabaena Fuscovaginata (Nostocales), a New Cyanobacterial Species from Periphyton of the Freshwater Alkaline Marsh of Everglades, South Florida, USA
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Fottea, 10(2): 235–243, 2010 235 Anabaena fuscovaginata (Nostocales), a new cyanobacterial species from periphyton of the freshwater alkaline marsh of Everglades, South Florida, USA. Jan MA R E š University of South Bohemia, Faculty of Science, Branišovská 31, CZ–37005 České Budějovice, Czech Republic, e–mail: [email protected] Abstract: A new species of the nostocalean genus Anabaena was discovered in periphytic cyanobacterial mats of the Everglades marsh, South Florida, USA. In some of the 10 samples where it occurred, the species dominated the cyanobacterial community, however, it was always part of a rich assemblage of microbiota, including many types characteristic for alkaline herbaceous wetlands of Caribbean America or freshwater tropical marshes. On the basis of typical morphological traits and occurrence in a periphytic habitat, I suggest that the species is a member of the genus Anabaena in its original sense, valid after recent separation of planktic taxa. As the most distinctive morphological feature, it has firm, coloured sheaths that have never been reported for this genus before. Key words: alkaline marshes, Anabaena, cyanobacteria, Everglades, morphology, periphyton, taxonomy Introduction to material from either countries with few laboratory facilities, inaccessible sites, or highly The concept of a species in Cyanobacteria and specific habitats. A phycologist has to answer the characters necessary for taxonomic description question: Is it worth describing the morphological of new taxa have been a subject of continual diversity of the material even if I cannot debate among concerned researchers. The modern characterize its genetic identity? In my opinion, polyphasic approach, introduced for cyanobacteria it is always useful to record the biodiversity as by AN A GNOSTIDIS & KOMÁRE K (1985, 1988, 1990), well as possible, thus providing at least basic KOMÁRE K & AN A GNOSTIDIS (1986, 1989) and since data for improvements by further researchers. then further developed and applied by scientists This approach is important in taxonomically all over the world (recently e.g. CA S A M A TT A et al. little explored and ecologically distinct habitats 2006; FIORE et al. 2007; FINSINGER et al. 2008; where there is a great number of unknown (but MCGREGOR & RA SMUSSEN 2008; PA LINS ka & characteristic) types of cyanobacteria. Examples MA RQU A RDT 2008, and many others), has proved to of such environments are wetlands and forests be the best available tool for reliable and objective of the tropics, high mountains, and other centers taxonomical conclusions. This approach requires of biodiversity. In these situations, I suggest that the synthesis of detailed description of classical new cyanobacterial taxa should be described morphological traits, ecology, life history, cell according to the International Code of Botanical ultrastructure and molecular phylogeny into a Nomenclature (MCNEILL et al. 2006), based congruent picture of a species defined by both exclusively on morphological and ecological unique genetic and phenotypic markers. characteristics. It should be emphasised that the Unfortunately, in some cases it is not possible descriptions should be as precise and detailed as to fulfil all these expectations and provide complete possible, and always accompanied by good–quality characterization of a species within a single study. images of the material for further comparisons. For various reasons, some cyanobacterial taxa do One habitat that fits these considerations not grow well in cultures or cannot be returned to is the alkaline (sub)tropical shallow marsh. the laboratory alive for cultivation and molecular Such wetlands are largely found in warm studies. In our experience, this especially applies parts of America, especially the Caribbean 236 MA R E š : Anabaena fuscovaginata, a new species region: Yucatán Peninsula and South Florida to 14 µg.L–1). The locality was mostly ecologically (Everglades), Cuba, Venezuela, Puerto Rico, similar to the original oligotrophic alkaline wetland, and at minor sites elsewhere. This type of marsh dominated by a community of emergent and submersed is characterised by raised pH (e.g. in Belize macrophytes with abundant periphyton encrusted by calcium carbonate. pH = 7.5 – 8.8, after REJMÁN K OVÁ et al. 2004), The material was preserved in 1.5% usually a nutrient–poor environment, fluctuating formaldehyde and observed by optical microscopy water levels, and characteristic communities of with differential interference contrast. Morphology aquatic plants with extensive accumulations of of the species was documented by means of digital calcified cyanobacteria–dominated periphytic photography and drawings based on examination of 10 mats. Thorough observations of the ecology, different samples (mats). hydrology, geology and biota and possible means of protection of this peculiar type of wetland can be found in DA VIS & OGDEN (1994), REJMÁN K OVÁ Results et al. (1996), REJMÁN K OVÁ & KOMÁR K OVÁ (2000) and RI C H A RDSON (2008). The described species of Anabaena dominated The species composition of cyanobacteria the periphytic assemblage at least in some of the at several sites has been studied using modern samples. In material from site C3 it sometimes taxonomic approaches. For instance, more than contributed up to 50% of total biomass according 80 species were found in the wetlands of Cuba to my estimations. It retained its characteristic (14 new taxa to science, cf. KOMÁRE K 1984a,b, features also in samples where it was scarce. The 1989a,b, 1995, 2005) and 65 species in the observed set of populations provided sufficient alkaline marshes of northern Belize (19 new to morphological and ecological data to separate science, cf. KOMÁRE K & KOMÁR K OVÁ –LEGNEROVÁ it from existing taxa as a new species under the 2007; TURI cc HI A et al. 2009). On the other hand, International Code of Botanical Nomenclature. the systematics of the periphytic cyanobacteria of Florida Everglades have been little studied, and Anabaena fuscovaginata MA R E š , sp. nov. usually only a few dominant species have been Diagnosis: Stratum micro– ad macroscopicum, noted (GLE A SON 1974; VYM A Z A L & RI C H A RDSON prostratum, submersum. Fila in fasciculis dense 1995; MCCORMI ck & O´DELL 1996). Application of agglomerata et intricata vel inter cyanobacteria a more comprehensive examination (MA R E š 2006) periphytica dispersa; mucilago sine colore, revealed great diversity of morphotypes, some of diffusus, interdum promptus. Trichoma sinuata, which evidently represented unknown species. So uniseriata, non ramosa, ad dissepimenta far only one of them has been described as a new constricta, 2.5–5 µm lata, paucim ad extremitatis species (Wolskyella floridanaMareš et Kaštovský, attenuata, metamerica, cum (2–) 6–18 (–25) cf. MA R E š et al. 2008). cellulis vegetatibus inter heterocytas. Akinetes In this study, I present a new species of heterocytarum utrinque contiguae, solitariae Anabaena (Nostocales, Cyanobacteria) from the vel raro 2 (–3)–seriatae. Vaginae adsunt vel periphyton of Everglades that in my opinion also gelatinosae, laeves, hyalinae vel luteo–fuscae, ad deserves valid taxonomic description due to its 2 µm latae, praecipue ad heterocytis akinetisque conspicuous phenotype and unique locality of evolutae. Cellulae cylindricae ad doliiformae, occurrence. isodiametricae vel paucim longior quam latae, 2.5–3.5 (–5) × 2.5–7 (–8) µm, aeruginosae, contentu praecipue granuloso; cellula apicalis Materials and Methods paucim elongata, plus minusve obtuse conica. Heterocytae intercalares, solitariae, sphaerice The samples of cyanobacterial mats were collected cylindricae, praecipue duplo longiores quam latae, in November 1999 inside the borders of the Water 3.5–5 (–7) × (5.5–) 7.5–10 (–12) µm. Akinetes Conservation Area 2A of Everglades, South Florida, cylindricae ad cylindricae–ovatae, apicibus USA, at the sampling points C3, C4 and C5 (for maps rotundatis, 7–10 (–13) × (10–) 14–20 (–25) µm, and detailed description of the area see VYM A Z A L et al. 2002). These were the rather less impacted or heterocytas contiguis, episporio laevi, pallide et unimpacted sites of a nutrient enrichment gradient translucente lutescenti in akinetis maturis. caused by long–term agricultural runoff (water Habitatio: Periphytice metaphyticeque inter column total phosphorus concentration less or equal algas cyanobacteriasque, interdum in massis, in Fottea, 10(2): 235–243, 2010 237 paludibus alkalinis, oligotrophicis, subtropicis (former subg. Anabaena) have been shown to cum vegetatio submerso. – Locus classicus: have significant genetic differences from planktic America septentrionalis, Florida – Everglades, in types (former subg. Dolichospermum) that area dicto Water Conservation Area 2A . are phylogenetically closer to Aphanizomenon Collectio conservata deposita in: BRNM HY 1.431 (GUGGER et al. 2002; RA J A NIEMI et al. 2005; depositus (coll. Jan Kaštovský, 23.09.1999); KOMÁRE K & KOMÁR K OVÁ 2006). The taxonomy iconotypus : figura nostra 1. of planktic cyanobacteria formerly included in the genus Anabaena has been further studied, and Filaments densely agglomerated and entangled in recently, new genera Dolichospermum (KOMÁRE K clusters or dispersed among other cyanobacteria & Za p o m ě l o v á 2007, 2008; Wack LIN et al. 2009) in periphyton; trichomes are sometimes and Sphaerospermopsis (Za p o m ě l o v á et al. 2009, enveloped by a fine and diffluent colourless 2010) have been separated from Anabaena on the mucilage. Trichomes wavy,