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Klebsormidium Flaccidum, Preliminary Experiments in Culture Have Shown a Very Large Range of Morppghological Variation

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Klebsormidium Flaccidum, Preliminary Experiments in Culture Have Shown a Very Large Range of Morppghological Variation Subaerial Algae in Tropical Rainforests, part II Juan M. Lopez‐Bautista The University of Alabama 1 Overview 9 Bidiiodivers ity and systematics of subilbaerial algae 9 Collection and study of subaerial algae •Molecular systematics of subaerial microchlorophytes •Evolution of subaerial microchlorophytes 2 Molecular systematics of subaerial microchlorophytes 3 Klebsormidium Klebsormidium is one of the most common genera of terrestrial Charophyta It consists of few species of green algae occurring on soil, subaerial surfaces and semi- aquatic habitats all around the world Pisa Italy 2005 The systematics of this genus has been entirely based on traditional morphology. The species-level relationships in Klebsormidium have not been examined with molecular tools Klebsormidium Galway Ireland 2004 Klebsormidium - One of the most widespread genera of terrestrial and freshwater green algae - Cosmopolitan distribution - Uniseriate filaments; no differentiated holdfast; one parietal chloroplast covering a half- 2/3 of the cell wall, with a single pyrenoid - 22 species currently described - Species-level systematics based almost entirely on morphology - Reproduction by biflagellate zoospores Principal morphological characters used for species identification in Klebsormidium 1) Width of filaments 2)Length of filaments 3) Cell shape 4) Presence/absence of constrictions between adjacent cells 5) Occasional presence/absence of biseriate parts 6) Presence of H-shaped pieces 7) Shape of the chloroplast Long Short Characters observable in culture 1)Growth habit: filamentous or fragmented 2) Presence/absence of a superficial hydrorepellent layer 3) Inducibility of zoospore release 4) Morphology of the zoosporangial aperture 5) Germination pattern of sporelings Fragmented Filamentous CITIES SAMPLED - Bergen (Norway) - Stockholm (Sweden) - Copenhagen ( Denmark) - Galway (Ireland) - London, Plymouth and Manchester (England) - HbHamburg, RtkRostock and Konstanz (Germany) - Bordeaux and Marseilles (()France) - Porto (Portugal) - Pisa, Siena and Pavia (Italy) - La Valletta (Malta) - Koper (Slovenia) - Prague (Czech Republic) Morphology (field) Very uniform morphology; virtually no differences between strains from different cities • Filaments mostly long, sometimes mixed with short fragments • Filaments unbranched, 6-9 µm wide • Slight constrictions between adjacent cells present in many strains • Cells regularly cylindrical • Chloroplast with smooth margin • Globular enlargements present in one strain (Pisa) • No biseriate parts or pseudo-branches Morphology (culture) • All strains growing well in both media used • Production of superficial hydrorepellent layer observed in the strains from Bergen, Galway , Hamburg and Konstanz • The strains from Marseilles, Pisa, La Valletta, Hamburg and Porto showed a marked tendency to fragmentation; after a few weeks the strains from Pisa and Marseilles consisted entirely of short fragments (2-5 cells) • Release of zoospores easily inducible only in the strain from Galway; release aperture indistinct • Sporelings in the Galway strain germinating with unipolar and bipolar pattern Although the morphology of the field material was virtually identical to Klebsormidium flaccidum, preliminary experiments in culture have shown a very large range of morppghological variation • Most strains produce only 3D filaments that remain submerged • Others populations produce also a superficial layer of parallel filaments that cover completely the surface of the medium, • In other strains, the filaments get fragmented into many short fragments, giving the cultures the appearance of a green “soup” Submerged Submerged Short Fragmented filaments Filaments + filaments Superficial layer Samples originally corresponding to Klebsormidium flaccidum are spread along the rbcL tree Samples originally corresponding to Klebsormidium flaccidum are spread along the rbcLtree On the other hand , samples corresponding to different morphological species are grouped together Samples originally corresponding to Klebsormidium flaccidum are spread along the rbcLtree On the other hand, samples corresponding to different morphological species are grouped together Furthermore, specimens not possible to ascribe to any known taxa, and from distant locations, are found to be genetically similar! Conclusions • The populations of Klebsormidium occurring in European cities belong to at least 4 different evolutionary lineages • In the field material, these show an almost identical morphology and there are no morphological characters useful to distinguish them • More differentiation observed in culture, in which different strains show different growth habits • The taxonomy of Klebsormidium needs critical reconsideration; none of the morphological characters used for identification have phylogenetic relevance • The chtitiharacterization of Kleb sormidi um flacc idum, type species of the genus, is a taxonomic mess that can be solved only sequencing the type specimen; depending on its identity, the whole genus may require a radical reassessment Lessons learned 9 Overall, these experiments and analyses indicated that a great deal of genetic diversity is hidden behind a very similar morphology 9 Morphology and cytology, even in “simple” and “common” forms, can be misleading 9 Unialgal cultures and phylogenetic analyses are needed to re-evaluate these so called cosmopolitan species Spongiochrysis hawaiiensis Dense yellow-orange coating believed to be Trentepohlia by the local scientists Interesting unicellular green alga, reproducing with a budding-like mechanism known only in two subaerial species of the class Trebouxiophyceae: • Marvania geminata Hindák 1976 • Marvania aerophytica Neustupa & Sehjonová 2003/Stichococcus ampulliformis Handa et al. 2003 •However, sequences of the 18S rRNA, showed unequivocally that this alga is a member of the Cladophorales Siphonocladales lineage (Ulvophyceae) •Since this group was so far known to include basically marine species with complex thalli, this was a discovery of exceptional interest, that shed new light on the phylogeny of this algal group •AtAutosporu ltilation hihas arisen separately at least 3 times (twice in Trebouxiophyceae and once in Ulvophyceae) Phylogenetic tree for a 18S rDNA sequence alignment of 1732 characters, with 55 representatives of the Viridiplantae and Spongiochrysis hawaiiensis. In a restricted analysis limited to 41 representat ives o f t his group, * * our samples were included in * the “Aegggagrop ila-clade” with high support 100/1.00 ML tree 18S rDNA (BS/Bayesian) Phylogenetic tree for an 18S rDNA sequence alignment of 1595 characters, with 41 representatives of the Siphonocladales Cladophorales complex, two outgroup taxa, and Spongiochrysis hawaiiensis * * * These data suggest that a marine ancestor gave origin to Spongiochrysis • Simplification of the thallus • Carotenoid production Similar trends during the colonization of terrestrial habitats have been found in the evolution of green algae Phylogenetic tree for an 18S rDNA sequence alignment of 1595 characters, with 41 representatives of the Siphonocladales Cladophorales complex, two outgroup taxa, and Spongiochrysis hawaiiensis Phylogenetic Considerations This alga represents a new subaerial lineage from the class Ulvophyceae It is the first known subaerial member of the Cladophorales Siphonocladales lineage For the un ice llu lar ha bit and f or th e sub aeri al h abi tat thi s speci es i s well differentiated from all other members of this lineage and can be considered the first known successful step of these algae into subaerial habitats New genus, new species: Spongiochrysis hawaiiensis Lessons learned 9 The subaerial habit has developed in an algal group that was formerly believed to be entirely aquatic (the Cladophorales) 9 A new subaerial lineage exists in the class Ulvophyceae 9 It also showed that identical morphologies and identical mechanisms of reproduction have developed independently in separated green algal lineages 9 Most importantly, great evolutionary surprises can be found in forest environmen ts, if ifdtild detailed surveys based on mod ern me thdlthodologi es are carri ided out The Order Trentepohliales The group is unusual (to say the least!) in many respects Unusual features 1. Exclusively terrestrial and orange-red (unlike the majority of green algae, which are aquatic and green) 2. Differentiated sexual and asexual reproductive structures including an absolutely intriguing, unique zoosporangial abscission process 3. Remarkable bilaterally keeled flagella and flagellar apparatus (i.e. basal bodies and associated components), and 4. Evolutionarily puzzling phragmoplast-mediated cytokinesis found no where else in the entire chlorophyte lineage and analogous to but distinct from the phhltragmoplast-meditddiated cyt tkiokinesi s of some green al gae and dthl the land pl ant s i n the charophycean lineage The order Trentepohliales Five genera currently recognized: Phycopeltis Printzina Trentepohlia Cephaleuros Discoid Prostrate Erect Discoid Reduced Epiphytic Epiphytic or Epiphytic or Obligated Endophytic epilithic epilithic epiphyte Isomorphic Isomorphic Isomorphic Heteromorphic Heteromorphic 28 species 8 species 38 species 15 species 4 species Trentepohlia Martius 1817 The genus Trentepohlia is the most diversified of the trentepohliacean algae, consisting of branched heterotrichous filaments, growing epilithic or epiphytic on the bark of trees, or in lichenic associations at exposed habitats forming conspicuous masses, usua
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