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A Taxonomic Study of the Calcicolous Endolitic Species of the Genus Verrucaria (Ascomycotina, Verrucariales) with the Lid-Like and Radiately Opening Involucrellum

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A Taxonomic Study of the Calcicolous Endolitic Species of the Genus Verrucaria (Ascomycotina, Verrucariales) with the Lid-Like and Radiately Opening Involucrellum A taxonomic study of the calcicolous endolitic species of the genus Verrucaria (Ascomycotina, Verrucariales) with the lid-like and radiately opening involucrellum. Josef Halda Muzeum a galerie Orlických hor, Jiráskova 2, 516 01 Rychnov n. Kn., [email protected] Abstract This study describes anatomy, ontogenesis and history of taxonomy of calcicolous endolitic species of the genus Verrucaria, which have been treated as Bagliettoa and Protobagliettoa (Verrucaria sect. Sphinctrina), and that are characterized by black, flat (lid-like) radiate crackling involucrellum, completely endolitic thallus and presence of macrosferoid cells formed by hy- phae in the lower part of the medullar layer. Most of the revised taxa were described by Servít (more than 100), and show a large taxo- nomic heterogeneity. Consequently, a detailed study had to be done to unify the group again. The characteristic characters have been recognized as the size and shape of the involucrellum and the development of the cortex layer. Following diacritic characters were recognized just morphological variations that originate regularly during ontogenesis. After revision the num- ber of species reducedfrom more than twenty to four: V. baldensis, V. limborioides, V. marmorea and V. parmigerella. The genus Bagliettoa differs from the genus Verrucaria in different forming of involucrellum, other significant characters (ascus morphology and further structures of hymenium), usually used to define lichen genera are identical. Therefore, a new section Bagliettoa has been estab- lished within the genus Verrucaria, which is well defined by the way of opening and shape of involucrellum. The genus Protobagliettoa has been separated from the genus Bagliettoa artifi- cially for the absence of ascospores in 1955 by Servít. The separation of genera Bagliettoa and Protobagliettoa to the family Bagliettoaceae is formal too, because other families in Verrucariales have not been established afterwards on the base of different structure of involucrellum. There- fore, the revision disapproves such unjustified separation and recommends to keep all the studied species of genus Verrucaria (Verrucariaceae). From the practical point of view, the revi- sion can help to determine all of the mentioned species outside. Eleven species (Limboria sphinctrina Duf., Protobagliettoa lactea (Arnold) Servít, V. bosniaca Servít, V. dalmatica Servít, V. grummanni Servít, V. gyelnikii Servít, V. parmigera J. Steiner, V. serbica Servít, V. steineri Kuan, V. subconcentrica (J. Steiner) Servít and V. suzaeana Servít), and also other 75 varieties and forms, were synonymized with the species V. baldensis A. Massal. Two species (V. ceracea J. Steiner and V. quarnerica Zahlbr.) and 4 varieties or forms were restricted to synonyms of the V. limborioides (A. Massal.) Clauzade & Roux. The species V. cazzae Zahlbr., one of its varieties, and one of its forms were restricted to synonyms of the species V. marmorea (Scop.) Arnold. Two species (P. kutakiana Servít and V. sphinctrinella Zschacke) and 20 variet- ies or forms were restricted to the synonyms of the species V. parmigerella Zahlbr. A new neotype and an isoneotype of the species V. marmorea there have been established, 19 taxa have been lectotypified (Protobagliettoa lactea (Arnold) Servít var. ochracea Servít, Verrucaria bagliettoaeformis (Hazsl.) Servít var. caesia (Arnold) Servít, Verrucaria baldensis A. Massal., Verrucaria baldensis A. Massal. var. canici Servít f. bakonyensis Servít, Verrucaria baldensis A. Massal. var. spilomatica A. Massal., Verrucaria calciseda DC. f. alocyza Arnold, Verrucaria calciseda DC. var. lactea Arnold, Verrucaria cazzae Zahlbr., Verrucaria inaequata (Servít) Servít f. helvetica Servít, Verrucaria inaequata (Servít) Servít var. transsilvanica Servít, Verrucaria parmigera J. Steiner, Verrucaria parmigera J. Steiner var. calcivoroides J. Steiner, Verrucaria parmigera J. Steiner var. calcivoroides J. Steiner f. arnoldiana Servít, Verrucaria pinguis J. Steiner f. alocizoides J. Steiner, Verrucaria sphinctrina Ach. var. tiroliensis Servít, Verrucaria sphinctrinella Zschacke var. elevata Servít f. spermogonifera (Arnold) Servít, Verrucaria steineri Kuan f. scrobicularis Servít, Verrucaria subconcentrica (J. Steiner) Servít var. nigroaureolata (Servít) Servít f. limitata Servít and Verrucaria subconcentrica (J. Steiner) Servít var. nigroaureolata (Servít) Servít f. rauca Servít). 2 Acta Mus. Richnov., Sect. natur. 10(1): 1-148, 2003 Content A. General part ....................................................................................... 3 I. Introduction ............................................................................................... 3 II. Servíts concept ......................................................................................... 8 II.-1. Generic concept .................................................................................... 8 II.-2. Interspecific and infrageneric distribution according to different thallus ..................................................................................................... 10 III. Characters of the studied species .......................................................... 11 III.-1. Morphology of thallus and ascoma .................................................. 11 III.-2. Ontogenesis ....................................................................................... 13 III.-3. Ecological claims and geographic distribution ................................ 16 III.-4. Taxonomical characters .................................................................... 17 III.-5. Described taxa ................................................................................... 18 III.-6. Table of characters ............................................................................ 20 B. Taxonomic part ................................................................................ 23 Bagliettoa A. Massal. ..................................................................................... 23 Protobagliettoa Servít..................................................................................... 23 IV. Studied species ...................................................................................... 23 IV.-1. Position of the studied species .......................................................... 23 IV.-2. A key for species determination of the studied species ................... 24 V. Verrucaria baldensis A. Massal................................................................... 24 V.-1. Geographic distribution in Europe .................................................... 29 V.-2. Extraeuropean distribution ................................................................ 38 V.-3. Commented list of synonyms.............................................................. 39 V.-4. Iconographia of type specimens....................................................... 106 VI. Verrucaria limborioides (A. Massal.) Clauzade & Roux ........................... 109 VI.-1. Geographic distribution in Europe ................................................ 111 VI.-2. Commented list of synonyms .......................................................... 111 VI.-3. Iconographia of type specimens ..................................................... 112 VII. Verrucaria marmorea (Scop.) Arnold .................................................... 113 VII.-1. Geographic distribution in Europe ............................................... 115 VII.-2. Commented list of synonyms ......................................................... 116 VII.-3. Iconographia of type specimens .................................................... 117 VIII. Verrucaria parmigerella Zahlbr. ............................................................ 118 VIII.-1. Geographic distribution in Europe.............................................. 121 VIII.-2. Extraeuropean distribution .......................................................... 124 VIII.-3. Commented list of synonyms ....................................................... 124 VIII.-4. Iconographia of type specimens .................................................. 130 C. Discussion ...................................................................................... 131 D. Summary ........................................................................................ 134 E. Literature cited ............................................................................... 136 F. List of synonyms ............................................................................. 140 G. Index ............................................................................................. 144 J. Halda: A taxonomic study of the calcicolous endolitic species of the genus Verrucaria.. 3 A. General part I. Introduction The aim of this study is to bring new knowledge on ascoma morphology and variability of thallus of selected endolitic species of the genus Verrucaria (V. baldensis, V. limborioides, V. marmorea and V. parmigerella) after more than fifty years, that could help to clarify their taxonomy and thus contribute to defining and distribution of the individual species that is so little known about, obtain a summarizing outline and revise interspecific and infraspecific relations of species, their variations and forms that are mainly recognized as morphological variations. The Bagliettoa group contains a large number of described taxa that are hardly recognized in field as well as microscopically. Further Zschackes (1933-1934) and
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