A New Species of Lecanora S. Lat., Growing on Lasallia Pustulata

Home , Carbonea, Carbonea vitellinaria, Japewia, Japewiella, Lasallia pustulata, Lecanora argopholis

The Lichenologist 40(2): 111–118 (2008) 2008 British Lichen Society doi:10.1017/S0024282908007469 Printed in the United Kingdom

A new species of Lecanora s. lat., growing on Lasallia pustulata

Sergio PEuREZ-ORTEGA and Javier ETAYO

Abstract: The new species Lecanora lasalliae Pe´rez-Ortega & Etayo is described from Spain. It is included provisionally in Lecanora s. lat as characters such as Lecanora-type ascus, exciple composed of thick radiating hyphae and the usual presence of algal cells in the excipulum or its lichenicolous habitus on Lasallia pustulata, do not ﬁt well within any known genus of lichenicolous or lichenized fungi. Its taxonomic afﬁnities with several taxa are discussed, including the parasitic Lecanora gyrophorina. Key words: Carbonea, Lecidea, lichenicolous fungi, Nesolechia, Phacopsis, Protoparmelia, Ramboldia, Scoliciosporum, Spain

Introduction and compare it to other genera with licheni- The umbilicate genus Lasallia Me´rat does colous species with Lecanora-type ascus with not host many species of fungi; so we were which the species could be related or surprised to find several healthy thalli of confused. Lasallia pustulata (L.) Me´rat. with small patches of apothecia growing on the thallus Material and Methods margins, mainly mixed with clusters of isidia. Because of the frequent presence of The material was examined using standard micro- scopical techniques. Photographs were taken with a dispersed algae in the exciple, thick excipu- Leica Mz75 stereomicroscope and a Zeiss Axioskop2 lar hyphae, the nature of the pigments in Plus microscope equipped with differential contrast. paraphyses and excipulum, and the Amyloid reactions were tested with Lugol’s reagent, Lecanora-type ascus, we hesitated to include either without or with a pre-treatment with KOH (I and K/I respectively). Microscopic measurements were it in any known genus. It might be related to made on material mounted in water. Excipular pig- other parasitic Lecanora species growing on ments have been studied in water, K (KOH) and N Umbilicaria, for example Lecanora gyropho- (nitric acid). rina Poelt & Zhurb. We are reluctant to Specimens selected for comparison. Ramboldia stuartii describe a new genus, as long as no similar (Hampe) Kantvilas & Elix. Australia: Tasmania: summit of Mt Charles, 42(09#N 146(15#E, 1040 m, 4 ii species are known and there is an absence of 2006, Kantvilas 112/06 (FR). molecular data by which to define more Ramboldia petraeoides (Nyl. ex C. Bab. & Mitt.) accurately the boundaries among the differ- Kantvilas & Elix. Australia: Tasmania: summit of ent natural groups within the Lecanoraceae. Mt Pearse, 41(30#S 145(37#E, 1000 m, 5 ii 2006, Kantvilas 97/06 (FR). In this paper we will describe the new species Ramboldia insidiosa (Th. Fr.) Hafellner. Spain: Ciudad Real. Horcajo de los Montes, Sierra del Chorito, forest of Pinus pinaster,39(20#21.1$N, S. Pérez-Ortega: Forschungsinstitut Senckenberg, Abt. 4(15#42.2$W, 636 m, 21 ix 2005, Pérez-Ortega 903 Botanik und Molekulare Evolutionsforschung, (hb. Pe´rez-Ortega). D-60325 Frankfurt am Main, Germany. Protoparmelia badia (Hoffm.) Haffelner. Spain: Departamento de Biología Vegetal II, Facultad de Huesca. Biescas, Valle de Tena, ladera norte de Penã Farmacia, Universidad Complutense, Plaza de Ramón Telera. Ibon de Piedrafita. 30TYN1361, 1600-1700 m, y Cajal s/n, Ciudad Universitaria, E-28040 Madrid, 10 ix 1994, L.G. Sancho 4281 (MAF). Spain. Email: [email protected] Carbonea vitellinaria (Nyl.) Hertel. Austria: Tirol: J. Etayo: Navarro Villoslada 16, 3( dcha., E-31003, O} tztaler Alpen, Obergurgl, entrance of ‘‘Rotmoostal’’ Pamplona, Spain. c. 250 m NNE of ‘‘Scho¨nwieshu¨tte’’, 46(51#00.9$N,

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11(01#08.7$E, talus slope along dirt road from blue. Hymenium hyaline, not inspersed, Obergurgl to Scho¨nwieshu¨tte, on Candelariella vitellina, c. 40 m high, the upper part brown, K 2250 m, 25 vii 2005, Printzen & Pérez-Ortega 901 (hb. Pe´rez-Ortega). olive-brown, N , KI+ blue. Paraphyses Carbonea supersparsa (Nyl.) Hertel. Austria: Tirol: numerous, simple or branched especially O} tztaler Alpen, Obergurgl, entrance of ‘‘Rotmoostal’’ near the base or in the apex (Fig. 1E), not c. 250 m NNE of ‘‘Scho¨nwieshu¨tte’’, 46(51#00.9$N, anastomosing, 2–2·5 m wide, capitate with 11(01#08.7$E, talus slope along dirt road from Obergurgl to Scho¨nwieshu¨tte, on Lecanora polytropa, slightly enlarged tips of 3–4·5 m wide, api- ca. 2250 m, 25 vii 2005, Printzen & Pérez-Ortega 902 cal cells with a extra-cellular brown pigment, (hb. Pe´rez-Ortega). K+ pale brown to olive-brown, and glued by a thin gel (Fig. 1D). Asci clavate, 8-spored, 30–369–13 m, ascus wall apically The Species strongly thickened and forming a tholus Lecanora lasalliae Pe´rez-Ortega & KI+ blue, with a non-reactive thin axial Etayo sp. nov. body, outermost layer of the ascal wall and tholus KI+ blue, close to the Lecanora-type Lichen lichenicola in thallo Lasalliae pustulatae crescens. Ascomata apothecioidea, aggregata, fusca vel nigra, (Fig. 1F, G). Ascospores ellipsoid to ovoid, superficialia, 0·15–0·4(–0·5) mm in diametro. Excipu- straight, simple, hyaline, (6·5–)7–9(–10·5) lum subhyalinum vel fuscum, 20–50 m grossum. 3·5–5·5 m (Fig. 1H–K). Hypothecium hyalinum, c. 100 m altum. Hymenium Pycnidia not seen. hyalinum, c.40m altum. Epihymenium fuscum, K+ pallide fuscum vel fusco olivaceum. Asci clavati, octosporati, 30–369–13 m. Ascosporae ellipsoideae Chemistry. No lichen secondary products vel ovoideae, hyalinae, simplices (6·5–)7–9 (–10·5) detected by TLC. 3·5–5·5 m. Typus: Spain, La Rioja, Ezcaray, Valle del rió Oja, Ecology and distribution. Lichenicolous, 30TVM9675, 943 m, on Lasallia pustulata on siliceous outcrops, 7 September 2004, V. Atienza (VAL-Lich— growing on Lasallia pustulata thalli, mainly holotypus). in the periphery, usually amongst clusters of isidia, but very rarely growing on them. (Fig. 1) Although we cannot conclude that Lecanora lasalliae is strictly lichenicolous, it is very Thallus absent or restricted to very small likely to be so since two of the localities granules amongst the ascomata; granules not where it was found were examined carefully corticated, brownish. and it was not found growing on other Apothecia dark brown to black, sessile, substrata. Although usually no damage was frequently grouped and regularly constricted observed on the host, in some cases when at the base, 0·15–0·4(–0·5) mm; disc flat, apothecia grew in marginal areas of the thalli finally slightly convex (Fig. 1A). Margin free of isidia, a degradation of the thallus was thin, sometimes excluded when mature, detected, and brown, apparently necrotic then apothecia virtually emarginate. Excipu- areas were evident. The apothecia are aggre- lum in longitudinal section pale but darken- gated in small patches and are very different ing to brown at the outer edge, pale brown in size and shape to the apothecia of the to hyaline in its inner part (Fig. 1B); 20– host. So far, the species is known only from 50 m thick, composed of radiating hyphae three mountainous localities in the north of with wide lumina, c. 5–7 m, in the outer Spain. part (Fig. 1C), and narrower in the inner Additional specimens examined. Spain: Asturias: part, c. 2–4 m; with gelatinized walls; Parque Natural de las Fuentes del Narcea, Deganã e external cells with extra-cellular brown pig- Ibias, road from Moal to Mt. Connio, 43(02#N, ment; not cupular. Photobiont green, not 6(41#W, 1026 m, on Lasallia pustulata on siliceous always present, not forming a defined layer, outcrops, 16 v 2005, S. Pérez-Ortega & A. Álvarez- Lafuente (hb. Pe´rez-Ortega 904, hb. Etayo); Parque cells dispersed in the exciple, 6–14 m Natural de las Fuentes del Narcea, La Viliella, Erica diam., sometimes shared in autospores. australis subsp. aragonensis shrub formation close to Hypothecium c. 100 m high, hyaline, KI+ the path from La Viliella to Vega de La Rondiella.

F.1. Lecanora lasalliae. A, habitus, apothecia; B, longitudinal section of an apothecium; C, detail of excipulum with radiating thick-walled hyphae; D, paraphyses, apex with brown pigment; E, branched paraphyses; F, ascus, detail of apex ; G, ascus with spores; H, I, J & K, spores, variability in age and shape. B, in water; C & D, lactophenol blue;F&G:Lugol after pre-treatment with K; E, H–K, in K. Scales: A=0·3 mm; B=40 m; C, D–F, G=5 m; H–K=2·5 m.

Monte La Viliella, 29TPH9162, 1020 m, 11 ix 2002, Lecanora Ach. 1810 Pérez-Ortega & J. M. Formentí (VAL-Lich). Type species: Lecanora allophana Nyl. Discussion of generic assignment This genus is characterized by asci of the Lecanora-type, simple hyaline ascospores, There are several lecideoid or lecanoroid and crustose thalli, with an apothecial genera including lichenicolous species with margin usually containing algal cells (Ryan Lecanora-type asci; viz. Carbonea (Hertel) et al. 2004). Lecanora s. str. is defined by Hertel, Lecanora Ach., Nesolechia A. Massal., species with atranorin and presence of Phacopsis Tul., Protoparmelia M. Choisy, oxalate crystals in their amphithecium. Ramboldia Kantvilas & Elix and Scolicio- However, within Lecanora, many species sporum A. Massal. We have not found any without these characteristics are found, and genus into which the new species could fit although many efforts have been made in the flawlessly. Below, we discuss the candidate past to identify more natural groups, most of genera to house the new species and the the relationships within the group remain reasons why we have decided to include it in unresolved. Meanwhile, the genus in a less Lecanora s. lat. restricted sense has been used to place several taxa with a combination of Lecanora- Carbonea (Hertel) Hertel 1983 type ascus and hyaline, simple ascospores. We have followed this practice with Type species: Carbonea atronivea (Arnold) Hertel L. lasalliae. Its ascus type, the hyaline, simple ascospores and especially the presence The genus Carbonea is superficially com- of algal cells in the excipulum are the parable and hosts several lichenicolous characters that push us to take this decision. species. Its excipulum composed of thick There are some examples of lichenicolous radiating hyphae with rather wide lumina is species with uncertain affinities that have very similar to that observed in L. lasalliae been placed within the genus Lecanora, (Fig. 1C). However, it differs in the exist- such as L. usneicola Etayo, on Usnea from ence of an intensely pigmented (carbonized) Antarctica (Etayo & Sancho 2006). An exciple lacking algal cells, and a green earlier example is Lecanora gyrophorina Poelt (K; N+ purple) pigment in the upper & Zhurbenko, a lichenicolous lichen grow- hymenium; in L. lasalliae the excipulum is ing on Umbilicaria hyperborea in Siberia K olive-brown and N, and regularly (Poelt et al. 1996), the systematic position of shows dispersed algal cells (although some- which also remains uncertain. It differs from times excluded when very old!). Further- L. lasalliae in its larger ascomata 0·4–0·8 mm more, the paraphyses are simple in Carbonea diam., with a typically lighter and rela- but usually branched in L. lasalliae. Other tively thick thalline margin with algae in the characters such as the size and shape of amphithecium, an unknown yellow pig- ascospores or the ascus-type and shape may ment (K) in the exciple and presence of fit well with our species. Recently Knoph norstictic acid. The presence of similar char- et al. (2004) enlarged the concept of acters in both species and the systematic Carbonea to include species without such a affinities of their hosts might indicate a close dark exciple. However we prefer to follow relationship. Hafellner (2004), keeping the genus homo- geneous. A similar choice was followed by Lecidea Ach. 1803 Etayo & Sancho (2006), who included a new lichenicolous species growing on Usnea Type species: Lecidea fuscoatra (L.) Ach. (=Lecanora usneicola Etayo) at least provisionally in Lecanora instead of Carbonea Lecidea is a large and artificial genus where mainly because of the pigmentation of the many species of microlichens have been exciple. placed, based on the occurrence of hyaline,

non-septate ascospores within lecideine growing on Paraparmelia and Xantho- apothecia (Hertel & Printzen 2004). During parmelia, P. falcispora Triebel & Rambold the past few decades numerous taxonomic growing on Usnea spp. and P. thallincola works have split more natural groups from (A. Massal.) Triebel & Rambold living on Lecidea into new genera, for example, Fusci- several parmelioid genera (Triebel & dea Wirth & Veˇzda (Wirth & Veˇzda 1972), Rambold 1995; Aproot & Triebel 2002). Carbonea Hertel (Hertel 1983), Farnoldia The last two species are easily differentiated Hertel (Hertel 1983), Immersaria Rambold from L. lasalliae by their pale to dark brown & Pietschmann (Rambold 1989), Japewia hypothecium, absence of algal cells in their Tønsberg (Tønsberg 1990), Japewiella exciple and their different hosts. Phacopsis Printzen (Printzen 1999), Ramboldia australis closely resembles L. lasalliae, but it Kantvilas & Elix (Kantvilas & Elix 1994), also lacks algae in its excipulum, has larger or more recently Hertelidea Printzen & ascospores and its hypothecium, although Kantvilas (Printzen & Kantvilas 2004). hyaline, is KI. Furthermore, it grows Despite removal of these new segregated on different hosts, Paraparmelia spp. and genera, Lecidea remains a heterogeneous Xanthoparmelia spp. genus, in which many new species with uncertain taxonomic position are placed. Protoparmelia M. Choisy 1929 We considered the possibility of including Type species: Protoparmelia badia (Hoffm.) Hafellner the new species in Lecidea. However ‘true’ Lecidea species always lack algae in their By the presence of algae in the exciple, exciple and have a Lecidea-type ascus, which together with its ascus type, the new species is moderately thick-walled and apically is reminiscent of the genus Protoparmelia. thickened, with a I+ blue outermost gelati- This genus is separated from Lecanora by nous layer (Hertel & Printzen 2004). its brown pigmentation, generally smaller ascospores, straight conidia borne both Phacopsis Tul. 1852 acrogenously and pleurogeneously, lack of atranorin and a somewhat different ascus Type species: Phacopsis vulpina Tul. type (Ryan, Nash & Hafellner 2004). Within this genus, species related to Protoparmelia & Nesolechia A. Massal. 1855 atriseda (Fries) R. Sant. & V. Wirth, differing Type species: Nesolechia oxyspora (Tul.) A. Massal. in some respects from the P. badia group, were included in the section Phaenora (Poelt Phacopsis and Nesolechia are two closely & Leuckert 1991). Species in this section are related genera of obligately lichenicolous mostly lichenicolous on crustose, silicicolous fungi included in the family Parmeliaceae lichens. All species so far known develop (Triebel et al. 1995; Persˇoh & Rambold their own thallus and following Poelt & 2002). Owing to their similarities they were Leuckert (1991), are further characterized synonymized by Triebel & Rambold (1988) by their close parasitic connection to lichens, although not all authors concur (e.g. Dore´ their ﬁliform, often curved conidia and et al. 2006), and recent molecular studies the frequent presence of norstictic acid (at (Persˇoh & Rambold 2002) have sug- least in the excipulum). Lecanora lasalliae gested that Phacopsis s. lat. is polyphyletic. may seem to be an extreme species adapted Although the habitus of L. lasalliae is really to a lichenicolous life-style on Lasallia, with different, both genera have some characters a much reduced thallus and scarce algae in common with it, but most species in them conﬁned to the exciple. The absence of induce distinct cecidia on the host thallus norstictic acid is also shared with the species and the apothecia are frequently immersed P. loricata Poelt & Veˇzda, a parasite on and lack an apothecial margin. Three Lecanora umbrosa Degel. However, the Phacopsis species have raised apothecia as in excipulum in L. lasalliae differs from that of L. lasalliae, viz. P. australis Aproot & Triebel Protoparmelia, which is cupular (Miyawaki

1991). The remaining species in the section developed, prosoplechtenchymatous proper differ from L. lasalliae in the presence of exciple, composed of radiating, branched norstictic acid, the presence of thallospores, and anastomosed hyphae with thick, conglu- a more or less well-developed thallus (some- tinated walls that are similar to the paraphy- times with effigurate margin), and different ses, abundantly branched and anastomosed crustaceous hosts: P. leproloma (R. Sant.) paraphyses, broadly clavate asci surrounded Rambold & Hertel on Lecidea paupercula by a gelatinous, amyloid sheet, with a well- Th. Fr.; P. placentiformis (Steiner) Poelt on developed, amyloid tholus containing a wide Bellemerea cupreoatra (Nyl.) Clauzade & cylindrical axial body surrounded by a Cl. Roux; P. cupreobadia (Nyl.) Poelt and P. deeper amyloid zone and a poorly devel- atriseda (Fries) R. Sant. on yellow Rhizo- oped, bluntly conical ocular chamber, more carpon, P. phaeonesos Poelt on Aspicilia or less of the Lecanora-type (Ekman & myrinii (Fr.) Hafellner and P. gesamia Poelt Tønsberg 2004). The ascospores are typi- & Grube on Lecanora argopholis (Ach.) Ach. cally bent spirally or sickle-shaped with attenuated ends. The new species share Ramboldia Kantvilas & Elix 1994 some features with Scoliciosporum especially Type species: Ramboldia stuartii (Hampe) Kantvilas & sensu Hafellner (2004). This author pro- Elix posed the combination S. intrusum (Th. Fr.) Hafellner for Lecidea intrusa Th. Fr. Ramboldia is a recently described genus (=Carbonea intrusa (Th. Fr.) Rambold & that shares many characters with L. lasalliae. Triebel), a lichenicolous lichen growing on It also includes one lichenicolous species, saxicolous lichens with ellipsoid 0–1(–3)- R. insidiosa (Th. Fr.) Hafellner, which grows septate ascospores. In our opinion, the on Lecanora varia (Hoffm.) Ach. (Kantvilas new taxon does not fit well in the homo- & Elix 2007). Ramboldia is circumscribed by geneous genus Scoliciosporum. The principal its lecideine apothecia, an internally unpig- differences between Scoliciosporum and L. mented excipulum composed of radiating, lasalliae, apart from spore shape, concern the branched and anastomosing hyphae, asci paraphyses and excipular hyphae, which are more or less of the Lecanora-type, with an much more richly branched and anastomo- amyloid tholus and broadly diverging axial sed in the former. Furthermore, the lumina body, sparsely branched and anastomosing of excipular hyphae in Scoliciosporum are paraphyses and simple, hyaline, non- much narrower and algae are always lacking halonate ascospores. Although most of the in the excipulum. characters that define the genus are present in L. lasalliae, obvious differences arise from a different excipulum, with much wider Other groups hyphae, which are not anastomosed and The new species should also be compared not or almost not branched, a somewhat with some taxa with a similar external different ascus type, since in L. lasalliae the appearance or that occur in similar hosts. axial body does not diverge as in Ramboldia. This is the case of the Umbilicaria specialized Furthermore, the brown excipular and epi- lichenicolous Gyrophthorus crustulosae hymenial pigment of L. lasalliae is not found (Creveld) Hafellner & Sancho. This species in Ramboldia. was initially described as belonging to the genus Phacopsis,asP. crustulosae Creveld Scoliciosporum A. Massal. 1852 (Creveld 1981), but Hafellner & Sancho (1990) demonstrated that this fungus has Type species: Scoliciosporum holomelaenum (Flo¨rke) A. Massal.=S. umbrinum (Ach.) Arnold perithecia, so it is not possible to confuse it with L. lasalliae. Polycoccum umbilicariae Scoliciosporum is characterized by its (Linds.) D. Hawksw. is another perithecial biatorine apothecia with an annular, poorly fungus that can be found on Lasallia thalli,

but its morphology isdistinct. Dark apoth- Ekman, S & Tønsberg, T. (2004) Scoliciosporum.In ecia similar to those in L. lasalliae are present Lichen Flora of the Greater Sonoran Desert Region, Vol. 2 (T. H. Nash III, B. D. Ryan, P. Diederich, in some genera of the order Leotiales (e.g. C. Gries & F. Bungartz, eds): 504–505. Tempe: Rhymbocarpus Zopf or Llimoniella Hafellner Lichens Unlimited. & Nav.-Ros.). A thorough revision of these Etayo, J. & Sancho, L. G. (2006) Two lichenicolous lichenicolous fungi was carried out by ascomycetes on austral species of Usnea (Lecano- Diederich & Etayo (2000). A very different rales, Ascomycota). Nova Hedwigia 83: 483–488. Hafellner, J. (2004) Notes on Scoliciosporum intrusum. ascus-type readily separates them from L. Fritschiana 49: 29–41. lasalliae. No species of those leotialean gen- Hafellner, J. & Sancho, L. G. (1990) U} ber einige era is known from umbilicarioid lichens. lichenicole Pilze und Flechten aus den Gebirgen Zentralspaniens und den Ostalpen. Herzogia 8: 363–382. Conclusions Hertel, H. (1983) U} ber einige aus Lecidea und Mela- nolecia (Ascomycetes lichenisati) auszuschliessende The lichenicolous species Lecanora lasalliae, Arten. Mitteilungen der Botanischen Staatssammlung growing on the margins of Lasallia pustulata, München 19: 441–447. Hertel, H. & Printzen, C. (2004) Lecidea.InLichen is described from the north of Spain. We Flora of the Greater Sonoran Desert Region, Vol. 2 tentatively include it in the genus Lecanora (T. H. Nash III, B. D. Ryan, P. Diederich, since the new taxon does not perfectly fit any C. Gries & F. Bungartz, eds): 287–309. Tempe: other described genus. We have been Lichens Unlimited. Kantvilas, G. & Elix, J. A. (1994) Ramboldia,anew tempted to include it in either Ramboldia or genus in the family Lecanoraceae. Bryologist 97: Protoparmelia, both sharing many characters 301–329. with the new taxon, but without a perfect fit. Kantvilas, G. & Elix, J. A. (2007) The genus Ramboldia Also, we are reluctant to create a new genus (Lecanorales): a new species, key and notes. for this single species and prefer to wait until Lichenologist 38: 135–141. Knoph, J., Rambold, G., Triebel, D. & Kainz, C. new similar species are found or new insights (2004) Carbonea.InLichen Flora of the Greater from other sources are added. We have Sonoran Desert Region, Vol. 2 (T. H. Nash III, B. D. made several unsuccessful attempts to Ryan, P. Diederich, C. Gries & F. Bungartz, eds): obtain ITS and SSU sequences from L. 54–55. Tempe: Lichens Unlimited. Miyawaki, H. (1991) Protoparmelia badia in Japan. lasalliae. Perhaps in the future when new Hikobia 11: 29–32. fresh collections are available and DNA Persˇoh, D. & Rambold, G. (2002) Phacopsis – a licheni- sequences can be obtained, the systematic colous genus of the family Parmeliacae. Mycological position of this new taxon will be clarified. Progress 1: 43–55. Poelt, J. & Leuckert, C. (1991) Der Formenkreis von The authors thank P. Diederich and an anonymous Protoparmelia atriseda (Lichenes, Lecanoraceae) in reviewer for their valuable comments. SPO is indebted Europa. Nova Hedwigia 52: 39–64. to the Ministerio de Educacioń y Ciencia (MEC, Spain) Poelt, J., Zhurbenko, M. & Matzer, M. (1996) Lecanora for a postdoctoral grant. gyrophorina sp. nov., eine bemerkenswerte, auf Umbilicaria hyperborea parasitierende Flechte aus Sibirien. Herzogia 12: 1–6. R Printzen, C. (1999) Japewiella gen. nov., a new lichen Aptroot, A. & Triebel, D. (2002) A new Phacopsis genus and a new species from Mexico. Bryologist species on Paraparmelia and Xanthoparmelia in 102: 714–719. southern Africa. Nova Hedwigia 74: 405–409. Printzen, C. & Kantvilas, G. (2004) Hertelidea, Creveld, M. (1981) Epilithic lichen communities in genus novum Stereocaulacearum (Ascomycetes the alpine zone of southern Norway. Bibliotheca lichenisati). Bibliotheca Lichenologica 88: 539–553. Lichenologica. 17: 1–287. Rambold, G. (1989) A monograph of the saxicolous Diederich, P. & Etayo, J. (2000) A synopsis of the lecideoid lichens of Australia (excl. Tasmania). genera Skyttea, Llimoniella and Rhymbocarpus Bibliotheca Lichenologica 34: 1–345. (lichenicolous Ascomycota, Leotiales). Lichenolo- Ryan, B. D., Nash III, T. H. & Hafellner, J. (2004) gist 32: 423–485. Protoparmelia.InLichen Flora of the Greater Sonoran Dore´, C. J., Cole, M. S. & Hawksworth, D. L. (2006) Desert Region, Vol. 2 (T. H. Nash III, B. D. Ryan, Preliminary statistical studies of the infraspecific P. Diederich, C. Gries & F. Bungartz, eds): variation in the ascospores of Nesolechia oxyspora 425-430. Tempe: Lichens Unlimited. growing on different genera of parmelioid lichens. Ryan, B. D., Lumbsch, H. T., Mesuti, M. I., Printzen, Lichenologist 38: 425–434. C., Suliwa, L. & Nash III, T. H. (2004) Lecanora.In

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Accepted for publication 7 March 2008