Endohyalina, the Genus in the Physciaceae to Accommodate the Species of the Rinodina Ericina-Group
Total Page:16
File Type:pdf, Size:1020Kb
Mycol Progress (2010) 9:37–48 DOI 10.1007/s11557-009-0616-2 ORIGINAL ARTICLE Endohyalina, the genus in the Physciaceae to accommodate the species of the Rinodina ericina-group Mireia Giralt & Pieter P. G. van den Boom & John A. Elix Received: 1 June 2009 /Revised: 10 August 2009 /Accepted: 12 August 2009 /Published online: 16 September 2009 # German Mycological Society and Springer 2009 Abstract The genus Endohyalina is characterized by and a simple method for determining the secondary crustose, autonomous, or obligately lichenicolous thalli, chemistry of lichenicolous fungi is provided. lecideine apothecia with a hymenium often more or less inspersed with oil droplets and a brown hypothecium, Keywords Generic recircumscription . Lichenicolous Bacidia-type asci, small Dirinaria-type ascospores devel- species . Lecanoromycetes . Newcombinations . Newspecies oping with type B ontogeny, bacilliform conidia and containing diploicin as the major secondary metabolite. The genus is based on four species previously included in Introduction Rinodina—R. ericina s. lat., R. insularis, R. interjecta and R. kalbii—and on two lichenicolous species from the All the species of the Rinodina ericina-group treated in the Canary Islands described here as new, Endohyalina present paper are considered to be congeneric. They are brandii and E. diederichii. The generic type, Endohyalina characterized as autonomous lichens with a crustose thallus rappii, is reduced to synonymy with E. ericina whereas E. or obligately lichenicolous species with an epikapylic to circumpallida is excluded from the genus and returned to endokapylic thallus, by containing diploicin and related Buellia s. lat. Except for the thalline growth form and the substances (except for R. kalbii Giralt & Matzer), an often common lichenicolous habit, the diagnostic characters of more or less inspersed hymenium, and by having short Endohyalina are akin to those of Diploicia. New chemical bacilliform conidia. Further, all possess small Dirinaria- data on Endohyalina insularis and E. kalbii are reported, type or similar ascospores, with pronounced internal wall thickenings (= rinodinoid ascospores), developing with type B ontogeny (where apical wall thickenings become distinct before the septum develops), Bacidia-type asci, lecideine M. Giralt Departament de Bioquímica i Biotecnologia (Àrea de Botànica), apothecia and a brown hypothecium. Because of this Facultat d”Enologia de Tarragona, Universitat Rovira i Virgili, combination of diagnostic characters, the generic position Marcel·lí Domingo s/n, of these species has oscillated between Buellia s. lat. and 43007 Tarragona, Spain Rinodina (Ach.) Gray. e-mail: [email protected] Hafellner (1979) transferred Buellia saxatilis f. insularis P. P. G. van den Boom (*) Arnold into Rinodina on the base of its rinodinoid ascospores. Arafura 16, Since then, many authors (e.g., Rambold et al. 1994;Giralt 5691 JA Son, The Netherlands 2001; Mayrhofer and Lambauer 2004;Sheard2004; Kaschik e-mail: [email protected] e-mail: [email protected] 2006) have discussed its generic position. The same ascospore character induced Kalb and Hafellner (1992)todescribeR. J. A. Elix madeirensis Kalb & Hafellner as a Rinodina. Giralt and Research School of Chemistry, Australian National University, Matzer (1994) pointed out that the most reliable features to Building 33, Canberra ACT 0200, Australia distinguish Buellia from Rinodina were those of the asco- e-mail: [email protected] spores and retained R. kalbii and R. madeirensis in Rinodina. 38 Mycol Progress (2010) 9:37–48 Subsequently, Giralt (2000) transferred Buellia ericina (Nyl.) distinct radiate-plicate marginal lobes, squamulose, foliose, Jatta to Rinodina for the same reason. umbilicate, and fruticose representatives are known. Apart Giralt (2000, 2001) considered all the above species to from thallus morphology, microscopic characters of the be closely related and noted that they may belong to an asci, ascospores, hymenium, and conidia as well as undescribed genus in the Physciaceae, since none of the chemistry have been used to segregate the genera of hitherto known crustose genera within this family pos- Physciaceae. Even after considering all these characters, sessed the same combination of characters (i.e., rinodinoid the species of the R. ericina-group cannot be clearly ascospores, type B ontogeny, Bacidia-type asci, lecideine assigned to any of the known genera of Physciaceae as apothecia and brown hypothecium), or to the placodioid they are currently defined. genus Diploicia A. Massal. If this was so, Diploicia would The combination of a crustose (autonomous) or an have to be recircumscribed to include microlichen species epikapylic to endokapylic (lichenicolous) thallus, Dirinaria- with independent or lichenicolous habit (with crustose or type ascospores with pronounced wall thickenings (rinodinoid endokapylic thalli). At the time, Giralt (2000) provisionally ascospores), ascospore-ontogeny of type B, Bacidia-type asci, retained the species in Rinodina in the expectation that lecideine apothecia with a poorly developed proper exciple further taxa might well be found with the same diagnostic (aethalea-type), a brown hypothecium, bacilliform conidia, characters. and the presence of diploicin (except for R. kalbii) exclude the Subsequently, three additional species have been found species of the R. ericina-group from the following genera of to belong to the R. ericina-group, namely Buellia interjecta Physciaceae: Müll. Arg., transferred to Rinodina by Mayrhofer et al. (1992) because of its rinodinoid ascospores, and the two 1. The clearly delimited genus Buellia de Not. nom. cons. or lichenicolous species described in this paper. Buellia s. str. (= Hafellia Kalb, H. Mayrhofer & Scheid., In our opinion, the new data described in the present cf. Gams 2004), with the generic type B. disciformis (Fr.) work confirm that the species of R. ericina-group are Mudd and all the species previously accommodated in congeneric and should be accommodated in a genus other Hafellia (and several other Buellia) are characterized by than Buellia s. lat. or Rinodina: either (1) a new genus of the typically large, Callispora-type ascospores with Physciaceae, or (2) the recircumscribed genus Diploicia. variable septation (from uniseptate to muriform) which However, when revising the 15 generic segregates of develop with a particular ontogeny (distinct from types A Buellia s. lat. erected by Marbach (2000), especially those and B) where subapical wall thickenings become distinct containing diploicin, it became apparent that Endohyalina before the septum is inserted and the hymenium is rappii (Imshaug ex R.C. Harris) Marbach could belong to strongly inspersed with oil droplets. In addition, they the R. ericina-group. A detailed study of the type specimen exhibit a very well-developed proper exciple which is not and additional material established that E. rappii is of the aethalea-type. Although some species contain conspecific with Rinodina ericina. As E. rappii is the type diploicin (B. demutans (Stirt.) Zahlbr., B. capitis-regum species of Endohyalina Marbach, the “new genus” in the W. A. Weber, H. desertica Marbach,B.dissa(Stirt.) Physciaceae [see (1) above] to include the species of the R. Zahlbr., Buellia oidalea (Tuck.) Tuck, B. oidaliella A. ericina-group already exists, namely the genus Endohya- Nordin,B.parastata(Nyl.) Zahlbr., B. procellarum A. lina. Alternatives (1) and (2) are further discussed below. Massal., B. pseudotetrapla (Pusswald) Elix, B. reginae Further alternative solutions to the problem include (3) Bungartz, and B. tetrapla (Nyl.) Müll. Arg.), their to maintain the species of the R. ericina-group in the genus morphological characters readily distinguish them from Rinodina, a solution discredited by the fact that molecular the R. ericina-group. data clearly indicate that R. insularis belongs to the Buellia- 2. The genera Rinodina, Rinodinella H. Mayrhofer & group rather than to the Physcia-group (Helms et al. 2003; Poelt and Mobergia H. Mayrhofer & Sheard, with Kaschik 2006; Wedin et al. 2002), or (4) to retain these Lecanora-type asci, lecanorine apothecia, and a color- species in the polyphyletic Buellia s. lat., an option which less hypothecium. does not resolve the taxonomic position of these well- 3. The genera Amandinea M. Choisy ex Scheid. & H. delimited taxa but obscures them in the heterogeneous Mayrhofer and Tetramelas Norman, with ascospores assembly that comprises Buellia s. lat. with slight internal wall thickenings or without thicken- ings (not rinodinoid) developing with type A ontogeny. (1) The case for accommodating the species of the Rinodina 4. The placodioid genera Australiaena Matzer, H. ericina-group in the genus Endohyalina Mayrhofer & Elix and Dimelaena Norman, are sepa- rated from the R. ericina-group, the former by its A wide range of thalline forms is observed within the lecanorine to biatorine apothecia, Lecanora-type asci, Physciaceae, where typically crustose, placodioid with and filiform conidia; and the latter by its cryptoleca- Mycol Progress (2010) 9:37–48 39 norine apothecia and Buellia-type ascospores develop- tive growth forms of the thallus, being placodioid in ing with ontogeny of type A. Another placodioid genus Diploicia, and foliose in the latter two genera. The is the sterile (and for this reason not discussed here) inclusion of the genus Diplotomma within this clade is Coscinocladium Kunze, included in the Physciaceae doubtful (Crespo et al. 2004).