Els líquens saxícoles i terrícoles del Parc Natural de Cap de Creus, amb un estudi filogenètic aplicat a la sistemàtica dels gèneres Diploschistes i Ingvariella

[cat] Una de les línies de treball del Grup de Liquenologia del Departament de Biologia Vegetal (Unitat Botànica) de la Universitat de Barcelona ha tingut com a objectiu estudiar la diversitat dels líquens i fongs liquenícoles del territori català, així com la descripció de les comunitats liquèniques d’aquesta àrea. Malgrat la contribució d’aquests estudis al coneixement de la flora liquenològica de Catalunya, encara queden moltes àrees del territori que no s’han estudiat amb deteniment. Una d’aquestes àrees és la inclosa dins del Parc Natural de Cap de Creus, on els extensos afloraments de roques àcides estan ocupats, de forma parcial o total, per comunitats liquèniques. Vàrem endegar un estudi florístic que tenia com a objectius principals elaborar un catàleg dels líquens saxícoles i terrícoles del Parc, examinar les diferències morfològiques, anatòmiques o ecològiques dels tàxons estudiats respecte les dades bibliogràfiques per avaluar-ne la variabilitat i caracteritzar les preferències ecològiques de les espècies estudiades. Com a resultat d’aquest estudi, s’ha elaborat un catàleg florístic format per 263 espècies de líquens i 31 de fongs liquenícoles. Del total de tàxons citats, un és primera cita per Europa, vuit per la península Ibèrica i 34 per Catalunya. L’anàlisi fitogeogràfic determina que la flora liquènica està formada per tres grups de distribució principals: espècies mediterrànies, temperades-mediterrànies i d’àmplia distribució. Addicionalment, la flora es veu enriquida amb espècies de distribució atlàntica i de distribució centreuropea o àrtico-boreal. En el plantejament inicial d’aquesta tesi vam decidir incorporar un element addicional de caire més taxonòmic i filogenètic. Fent ús de les dades i espècimens recollits per al treball florístic, vam seleccionar aquells tàxons que resultessin conflictius i amb necessitat de ser revisats. Per aquest motiu, vam plantejar un examen exhaustiu centrat en els gèneres Diploschistes i Ingvariella. El gènere Diploschistes (Graphidaceae) presentava interessants qüestions no resoltes, com la delimitació del gènere, la validesa taxonòmica i sistemàtica dels tres grups d’espècies establerts en base a la morfologia dels apotecis, i la delimitació de diverses espècies. Per resoldre aquestes qüestions vam reconstruir diverses filogènies basades en caràcters emprats tradicionalment en taxonomia (morfològics, anatòmics, químics, ecològics), marcadors moleculars (nuITS, mtSSU i nu LSU) i la combinació d’ambdós tipus de caràcters. Els resultats confirmen que Diploschistes és monofilètic i demostren, per primera vegada, que els tres grups morfològics considerats dins del gènere són monofilètics, i se’n proposa una classificació infragenèrica. Les anàlisis filogenètiques revelen novetats en algunes espècies: l’existència d’un clade dins D. ocellatus, corresponent a espècimens sense apotecis que només tenen picnidis, i la presència d’un clade que agrupa tots els espècimens de la subespècie D. diacapsis subsp. neutrophilus, i que es proposa considerar al nivell de l’espècie. El gènere monotípic Ingvariella fou escindit de Diploschistes a partir de la morfologia de l’exciple, i mantingut dins de la família Thelotremataceae (= Graphidaceae) dins de l’ordre de les Ostropales. L’estudi detallat de la morfologia i anatomia d’Ingvariella ens va suggerir possibles afinitats amb altres famílies de l’ordre. Els objectius d’aquest estudi van ser avaluar la monofília d’Ingvariella i investigar la seva posició sistemàtica dins l’ordre. S’han inferit les relacions filogenètiques amb una matriu molecular (basada en els marcadors nuLSU i mtSSU), on s’ha inclòs una àmplia representació de membres de les Ostropales. Els resultats revelen que Ingvariella és monofilètic i pertany a la família de les Stictidaceae. Amb la inclusió del primer gènere de líquens saxícoles dins aquesta família, s’expandeix considerablement la diversitat ecològica de les Stictidaceae, dins les quals s’han inclòs fongs sapròfits, líquens corticícoles, i espècies que poden ésser facultativament liquenitzades. A més dels resultats esmentats, s’inclouen noves dades sobre les relacions entre altres famílies dins les Ostropales.[eng] This thesis is structured in two different parts. The first part, is an exhaustive floristic study of the saxicolous, epiphytic (only if sporadically growing on rocks), and terricolous lichens, and their lichenicolous fungi from the Cap de Creus Natural Park (NE Iberian Peninsula). The study reports 263 lichen taxa and 31 lichenicolous fungi from the study area, of which one is new for Europe, eight new for the Iberian Peninsula, and 34 for Catalonia. From a phytogeographical point of view, the flora is mainly composed of Mediterranean temperate species and Mediterranean lichens in a broad sense, enriched with a considerable number of Atlantic species in the most humid sites of the Park, and some species of boreal distribution. The large number of different microclimatic conditions in the area, combined with frequent extensions of rock outcrops, makes the Cap de Creus Natural Park an extraordinary region for the diversity of saxicolous and terricolous lichens. During the floristic study, we detected conflictive and unresolved taxonomic and systematic questions regarding the genera Diploschistes and Ingvariella. In the second part of the thesis, we have tried to clarify these questions within a phylogenetic framework. In the revision of Diploschistes, the reconstruction of the phylogenetic relationships of the genera within its family (Graphidaceae) confirmed that the genus is monophyletic. We also reconstructed the phylogenetic relationships within Diploschistes using a combined dataset of morphological, chemical, and molecular data. The results show that the three groups traditionally distinguished within the genus (based on ascomatal morphology) are monophyletic. Therefore, it is proposed a new subgeneric treatment for Diploschistes, and it is confirmed the taxonomical value of fruiting body types. In The monotypic genus Ingvariella was segregated from Diploschistes based on fruiting body anatomy, and placed within the Thelotremataceae (= Graphidaceae), in the order Ostropales. Our morphological observations, however, suggest affinities to other families within the order. We inferred the phylogenetic relationships of a broad array of ostropalean fungi based on mtSSU and nuLSU. The results demonstrate that Ingvariella is monophyletic and belongs to the Stictidaceae, expanding the broad ecological diversity of this family

First report of the pantropical species Diploschistes rampoddensis from Europe

The lichen species Diploschistes rampoddensis, previously known only from a few localities in tropical and subtropical Asia and Oceania, is reported here for the first time in Europe. A detailed description, including macro- and microscopic characters, and comparisons with closely related taxa are also provided. Molecular analyses based on the nrITS were used to confirm this new record for the European lichen biota

Hongos liquenizados y liquenícolas de la Sierra de Albarracín (Teruel, España)

Se presenta un catálogo de 462 hongos liquenizados y liquenícolas de la Sierra de Albarracín (Teruel, Aragón, España), como resultado de la IV Campaña de Recolección organizada por la Sociedad Española de Liquenología (SEL). Diplotomma hedinii es novedad para la Península Ibé- rica y Lepraria leuckertiana constituye una segunda cita peninsular, 69 taxones son novedad para Aragón y 86 lo son para la provincia de Teruel

Lichenized and lichenicolous fungi from the Sierra de Albarracín (Teruel, Spain)

Se presenta un catálogo de 462 hongos liquenizados y liquenícolas de la Sierra de Albarracín (Teruel, Aragón, España), como resultado de la IV Campaña de Recolección organizada por la Sociedad Española de Liquenología (SEL). Diplotomma hedinii es novedad para la Península Ibérica y Lepraria leuckertiana constituye una segunda cita peninsular, 69 taxones son novedad para Aragón y 86 lo son para la provincia de Teruel.A catalogue of 462 lichenized and lichenicolous fungi from Sierra de Albarracín, (Teruel, Spain), is presented as the result of the SEL (Sociedad Española de Liquenología) IV collecting field trip. Diplotomma hedinii is recorded for the first time for the Iberian Peninsula and Lepraria leuckertiana is registered for the second time. The checklist also includes 69 taxa new for Aragón Community and 86 new for the province of Teruel

Hongos liquenizados y liquenícolas de la Sierra de Albarracín (Teruel, España)

[spa] Se presenta un catálogo de 462 hongos liquenizados y liquenícolas de la Sierra de Albarracín (Teruel, Aragón, España), como resultado de la IV Campaña de Recolección organizada por la Sociedad Española de Liquenología (SEL). Diplotomma hedinii es novedad para la Península Ibérica y Lepraria leuckertiana constituye una segunda cita peninsular, 69 taxones son novedad para Aragón y 86 lo son para la provincia de Teruel.[eng] A catalogue of 462 lichenized and lichenicolous fungi from Sierra de Albarracín, (Teruel, Spain), is presented as the result of the SEL (Sociedad Española de Liquenología) IV collecting field trip. Diplotomma hediniiis recorded for the first time for the Iberian Peninsula and Lepraria leuckertiana is registered for the second time. The checklist also includes 69 taxa new for Aragón Community and 86 new for the province of Teruel

A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families

The Lecanoromycetes is the largest class of lichenized Fungi, and one of the most species-rich classes in the kingdom. Here we provide a multigene phylogenetic synthesis (using three ribosomal RNA-coding and two protein-coding genes) of the Lecanoromycetes based on 642 newly generated and 3329 publicly available sequences representing 1139 taxa, 317 genera, 66 families, 17 orders and five subclasses (four currently recognized: Acarosporomycetidae, Lecanoromycetidae, Ostropomycetidae, Umbilicariomycetidae; and one provisionarily recognized, ‘Candelariomycetidae’). Maximum likelihood phylogenetic analyses on four multigene datasets assembled using a cumulative supermatrix approach with a progressively higher number of species and missing data (5-gene, 5 + 4-gene, 5 + 4 + 3-gene and 5 + 4 + 3 + 2-gene datasets) show that the current classification includes non-monophyletic taxa at various ranks, which need to be recircumscribed and require revisionary treatments based on denser taxon sampling and more loci. Two newly circumscribed orders (Arctomiales and Hymeneliales in the Ostropomycetidae) and three families (Ramboldiaceae and Psilolechiaceae in the Lecanorales, and Strangosporaceae in the Lecanoromycetes inc. sed.) are introduced. The potential resurrection of the families Eigleraceae and Lopadiaceae is considered here to alleviate phylogenetic and classification disparities. An overview of the photobionts associated with the main fungal lineages in the Lecanoromycetes based on available published records is provided. A revised schematic classification at the family level in the phylogenetic context of widely accepted and newly revealed relationships across Lecanoromycetes is included. The cumulative addition of taxa with an increasing amount of missing data (i.e., a cumulative supermatrix approach, starting with taxa for which sequences were available for all five targeted genes and ending with the addition of taxa for which only two genes have been sequenced) revealed relatively stable relationships for many families and orders. However, the increasing number of taxa without the addition of more loci also resulted in an expected substantial loss of phylogenetic resolving power and support (especially for deep phylogenetic relationships), potentially including the misplacements of several taxa. Future phylogenetic analyses should include additional single copy protein-coding markers in order to improve the tree of the Lecanoromycetes. As part of this study, a new module (‘‘Hypha’’) of the freely available Mesquite software was developed to compare and display the internodal support values derived from this cumulative supermatrix approach.Keywords: Classification, Multi-gene phylogeny, Lichenized fungi, Systematics, Cumulative supermatrix, Lecanoromycete

First report of the pantropical species Diploschistes rampoddensis from Europe.

The lichen species Diploschistes rampoddensis, previously known only from a few localities in tropical and subtropical Asia and Oceania, is reported here for the first time in Europe. A detailed description, including macro- and microscopic characters, and comparisons with closely related taxa are also provided. Molecular analyses based on the nrITS were used to confirm this new record for the European lichen biota

Schizoxylon as an experimental model for studying interkingdom symbiosis.

Experiments to re-synthesise lichens so far focused on co-cultures of fungal and algal partners. However, recent studies have revealed that bacterial communities colonise lichens in a stable and host-specific manner. We were therefore interested in testing how lichenised fungi and algae interact with selected bacteria in an experimental setup. We selected the symbiotic system of Schizoxylon albescens and the algal genera Coccomyxa and Trebouxia as a suitable model. We isolated bacterial strains from the naturally occurring bacterial fraction of freshly collected specimens and established tripartite associations under mixed culture experiments. The bacteria belong to Actinobacteria, Firmicutes and Proteobacteria and corresponded to groups already found associated with fungi including lichens. In mixed cultures with Coccomyxa, the fungus formed a characteristic filamentous matrix and tightly contacted the algae; the bacteria distributed in small patches between the algal cells and attached to the cell walls. In mixed cultures with Trebouxia, the fungus did not develop the filamentous matrix, but bacterial cells were observed to be tightly adhering to the fungal hyphae. Our experiments show that this tripartite fungal-algal-bacterial model system can be maintained in culture and can offer multiple opportunities for functional studies based on experiments under controlled conditions in the laboratory

Rhagadodidymellopsis endocarponis gen. et sp. nov. and Arthopyrenia symbiotica (Dothideomyceta), two lichenicolous fungi growing on Endocarpon species.

The lichenicolous fungus Rhagadodidymellopsis endocarpi (Dothideomyceta) growing on the thallus of the terricolous lichen Endocarpon pusillum is described from Spain and Australia as new to science. The new genus and species is compared with other taxa from the genera Didymellopsis and Zwackhiomyces (Xanthopyreniaceae, Collemop-sidiales, Dothideomyceta), in particular with D. perigena, a species also having hyaline didymospores and also growing on Endocarpon. Rhagadodidymellopsis endocarpi is char-acterized by its almost completely superficial stromatic ascomata with a coarse and irregular surface, and an ascomatal wall of very irregular thickness, and ascospores smaller than those of D. perigena. We also compare the new species with other Endocarpon parasites, including Arthopyrenia symbiotica. This is a misunderstood species, originally described as Verrucaria symbiotica, which we also discuss in detail in this study