Morphology and anatomy ofchasmolithic versus epilithic growth: a taxonomic revision of inconspicuous saxicolous Buellia species from the Sonoran Desert Region generally ascribed to the "Buellia puncfafa" group F. Bungartz, T.H. Nash III, and B.D. Ryan Received 3 September 2003. Published on the NRC Research Press Web site at http://canjbot.nrc.ca on 25 May 2004. F. Bungartz,l T.IL Nash m, and B.D. Ryan. School of Life Sciences, Arizona State University, P.O. Box 874601, Tempe, AZ 85287·4601, USA. ·Corresponding author (e-mail: frank.bungartz@asu.edu). Can. J. Bot. 82: 540-562 (2004) doi: lO.1l39/B04-028 © 2004 NRC Canada 540 Morphology and anatomy ofchasmolithic versus epilithic growth: a taxonomic revision of inconspicuous saxicolous Suellia species from the Sonoran Desert Region generally ascribed to the "Suellia punctata" group F. Bungartz, T.H. Nash III, and B.D. Ryan Abstract: Six saxicolous species of Buellia, which were previously generally identified as Buellia punctata (Hoffm.) A. Massal., were examined from the Sonoran Desert. None of the species belongs to Beullia punctata s. str. Though inconspicuous, it can be demonstrated that the thallus morphology of these species is quite distinct and far less variable than previously assumed. Most species are epilithic, even though their thalli also show some degree of substrate penetration. Buellia sequax (Nyl.) Zahlbr., not previously reported from North America, is exclusively chasmolithic. Three new species with epilithic thalli, Beullia christophii Bungartz sp. nov., Beullia ryanii Bungartz sp. nov., and Beullia tergua Bungartz sp. nov., are described. Two species with filiform conidia are not treated in the genus Amandinea. We discuss why the current delimitation of this genus, based solely on conidial shape, is rejected: Buellia pullata Tuck., with filiform conidia, is consequently not transferred into Amandinea. Amandinea lecideina (H. Mayrhofer & Poelt) Scheid. & H. Mayrhofer is synonymized with Beullia prospersa (Nyl.) Riddle. Key words: North America, taxonomy, Physciaceae, Amandinea, conidia, thallus morphology, new species. Resume: Les auteurs ont examine six especes de Buellia venant dans Ie desert de Sonoran; ces eSpCces ont ete jusqu'ici identifiees comme Buellia punctata (Hoffm.) A. Massal. Aucune de ces entites appartient au Beullia punctata s. str. Bien que ce soit peu evident, on peut demontrer que la morphologie du thalle de ces especes est assez differente, et beaucoup moins variable qu'on l'a consideree auparavant. La plupart des especes sont epilithiques, meme si leurs thalles montrent egalement un certain degre de penetration dans Ie substrat. Le Buellia sequax (Nyl.) Zahlbr., jamais rapporte auparavant en Amerique du Nord, est exclusivement chasmolithique. Les auteurs decrivent trois nouvelles especes munies de thalles epilithiques : Beullia christophii Bungartz sp. nov., Beullia ryanii Bungartz sp. nov. et Beullia tergua Bungartz sp. nov. En depit de leurs conidies filiformes, deux especes ne sont pas attribuees au genre Amandinea. Les auteurs discutent pourquoi la presente delimitation de ce genre, basee uniquement sur la forme des conidies, est rejetee : consequemment on ne transtere pas Ie Buellia pullata Tuck., muni de conidies filiformes, avec les Amandinea. On place l'Amandinea lecideina (H. Mayrhofer & Poell) Scheid. & H. Mayrhofer en synonymie avec Ie Beullia prospersa (Nyl.) Riddle. Mots eles : Amerique du Nord, taxonomie, Physciaceae, Amandinea, conidies, morphologie du thalle, nouvelles especes. [Traduit par la Redaction] Introduction The taxonomy of crustose lichen species with thin and inconspicuous thalli is generally poorly resolved. These species are often overlooked, rarely collected, and their thallus characteristics have been disregarded as highly variable and poorly differentiated. The attention of taxonomic treatments Received 3 September 2003. Published on the NRC Research Press Web site at http://canjbot.nrc.ca on 25 May 2004. F. Bungartz,l T.n. Nash ITI, and B.D. Ryan. School of Life Sciences, Arizona State University, P.O. Box 87 4601, Tempe, AZ 85 287-4601, USA. lCorresponding author (e-mail: frank.bungartz@asu.edu). Can. J. Bot. 82: 540-562 (2004) has therefore often focused on species with well-developed epilithic thalli, whereas species that show some degree of endolithic growth have been neglected. The present study focuses on a group of species that in North America have generally been treated as Buellia punctata (Hoffm.) A. Massa!. or more recently as Amandinea punctata (Hoffm.) Coppins & Scheid. In particular, the saxicolous material commonly identified as this species is poorly understood (Mayrhofer and Moberg 2002). Thallus variation has largely been ignored. For example, in his treatment of the North American species of Buellia, Imshaug (1951) states" ... Buellia pullata is, as Tuckerman (1888) indicated, a saxicolous form of B. punctata with a well developed thallus. Buellia saxicola de Lesd., on the other hand, is a saxicolous form of B. punctata with a scant to obsolete doi: 10.1l39/B04-028 © 2004 NRC Canada Bungartz et al. thallus ...". The material examined in our study indeed shows some variation in thallus morphology. Several distinct species can nevertheless be distinguished based on thallus morphology as well as spore ontogeny, apothecial pigments, conidial length, and secondary chemistry. Buellia pullata, for example, is not, as Imshaug (1951) suggested, a mere form of B. punctata, but a distinct species. Likewise, it can be demonstrated that Beullia saxicola is a synonym of Beulfia sequax (Ny1.) Zahlbr., a taxon not previously documented from North America, but recognized as a distinct species by Scheidegger (1993). Six species previously treated in Imshaug's "B. punctata" group are distinguished among the Sonoran material. Thallus variation in this group is clearly much less pronounced than previously assumed. None of the species presented belongs to B. punctata s. str., which apparently does not occur on rock substrates in the Sonoran Desert Region. Contrary to recent practice, the genus Amandinea is not accepted. The taxonomy of that genus is currently not clearly resolved, and all species are therefore treated as members of Buellia s. 1. Materials and methods All specimens were examined with light microscopy using hand- and cryo-sections. Both conventional bright field microscopy (BF) and differential interference contrast (DIC) were used. Selected specimens were also studied with scanning electron microscopy (SEM) and transmission electron microscopy (TEM) according to a protocol described in detail by Bungartz et al. (2002). To improve dehydration and 541 infiltration, this protocol was modified according to Bungartz and Nash (2004a). Representative specimens were cut with a low-speed diamond saw and the sections treated according to a protocol outlined by Bungartz et al. (2004). To visualize hyphae penetrating dark stone substrates, the protocol was modified and specimens were stained with lactophenol cotton blue (LCB) instead of periodic acid - Schiff's reagent (PAS). Sections were studied with both light microscopy and SEM as discussed in Bungartz et al. (2004). Some of the sections were critically point dried and gold coated. All sections were studied in a JEOL JSM-840A SEM at accelerating voltages between 7 and 12 kV in secondary and backscattered emission mode. Specimens were spot tested and routinely examined with standardized thin-layer chromatography (TLC) (Culberson and Kristinsson 1970; Culberson and Johnson 1982; White and James 1985; Orange et al. 2001). To differentiate various xanthones, selected specimens were additionally examined with standardized high-performance liquid chromatography (HPLC; Feige et al. 1993). TLC plates were interpreted with the computer program WINTABOLITES (Mietzsch et a1. 1994) and scanned for permanent record (Egan 2001). Spores measurements are given according to Nordin (2000). Pigment names follow Meyer and Printzen (2000). Figure plates were assembled and processed in Adobe Photoshop 7.0, and selective contrast adjustments were made to emphasize details such as the apothecia in SEM micrographs. Results Key to the species lao Thallus chasmolithic, discontinuous, usually of dispersed granules, growing among minerals of the substrate. . . . . . lb. Thallus epilithic, forming a thin, more or less continuous crust on the substrate surface. . . . . . . . . . . . . . . 2 3 2a. Thallus with a pale yellowish tinge, UV+ yellow to orange, with xanthones; premature ascospores ellipsoid, with distinct septum thickening; conidia filiform. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. prospersa 2b. Thallus pale brown to gray, UV-, without xanthones; premature ascospores narrowly oblong, without septum thickening; conidia bacilliform. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sequax 3a. Young apothecia erumpent, outer exciple in cross section carbonized by an aeruginose, HN03+ violet pigment, conidia bacilliform. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3b. Young apothecia emergent, outer exciple in cross section carbonized by a dull brown pigment, HN0 3-, conidia bacilliform to filiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4a. Thallus "leather" brown, rimose, not delimited by a distinct hypothallus, exciple deeply aeruginose, ascospores 10-15 ｾｭ x 6-9 ｾｭＬ conidia 4-7 ｾ ｭ Ｎ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. tergua 4b. Thallus olive gray to brownish olive, distinctly areolate in the center, with an intermediate undifferentiated zone delimited by a distinct black hypothallus; exciple dull fuscous brown, ascospores 9-13 ｾ ｭ x 4-8 ｾｭＬ conidia 2-5 ｾｭ . . . . B. ryanii Sa. Premature spores broad, almost globose, evenly thick walled, septum not thickened; exciple thick, outer carbonized zone usually >20 ｾｭ in cross section, conidia bacilliform . . . . . . . . . . . . . . . . . . . . . . . . B. christophii 5b. Premature spores oblong to ellipsoid, not conspicuously broadened, with a thin spore wall; septum thin or thickened, exciple thin, outer carbonized zone usually <20 ｾｭ in cross section, conidia filiform . . . . . . . . . . . . . . . . . . . 6 00. Thallus pale, usually with a yellow tinge, UV+ yellow to orange, with xanthones; spores with a distinct, more or less persistent septum thickening . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. prospersa 6b. Thallus deep brown, rarely dull gray, UV-, without xanthones; spores with a brief stage of inconspicuous septum thickening . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. pullata © 2004 NRC Canada 542 Can. J. Bot. Vol. 82, 2004 Figs. 1-7. Buellia christophii. Fig. 1. Overview of the epilithic rimose thallus (Nash 33979 - holotype). Fig. 2. Close-up of an epilithic rimose-areolate thallus (Nash 32100). Fig. 3. Light micrograph of the athalea-type exciple (cross section in water): the thick outer part consists of swollen cells, which are carbonized with the dull brown pigment cf. elachista-brown (Nash 33979 - holotype). Fig. 4. Scanning electron microscope (SEM) micrograph of an exciple cross section (Nash 32100). Fig. 5. Light micrograph of broad premature ascospores with a conspicuous thickened wall (Nash 33979 - holotype). Fig. 6. TEM micrograph ofa premature ascospore: (1) thin, smooth perispore, (2) intermediate layer, (3) thick proper wall, and (4) endospore (Nash 33979 - holotype). Fig. 7. Light micrograph of a pycnidium with bacilliform conidia (Nash 33979 - holotype). Figs. 8-14. Buellia pullata. Fig. 8. Overview of the epilithic rimose-areolate thallus (Bolander 131 - lectotype). Fig. 9. Close-up of the epilithic rimose-areolate thallus (Nash 41227). Fig. 10. Light micrograph of a thallus cross section: (e) epinecral layer, (c) cortical layer, (a) algal layer (Nash 41227). Fig. 11. Light micrograph of an athalea-type exciple (cross section in water): the thin outer part consists of swollen cells, which are carbonized with the dull brown pigment cf. elachista-brown (Nash 41227). Fig. 12. SEM micrograph of an exciple cross section (Nash 11426). Fig. 13. Light micrograph of a pycnidium with filiform conidia (Nash 41227). Fig. 14. TEM micrograph of a mature ascospore: (1) thick, fractured perispore, (2) intermediate layer, (3) proper wall, and (4) endospore (Nash 11426). Species descriptions Buellia christophii Bungartz sp. nov. Thallus saxicolus, crustaceus, rimosus vel rimosoareolatus, tenuis vel crassus, fulvus, sine hypothallo. Apothecia sessilia, lecideina, marginibus propriis crassis. Excipulum crassum, fulvum, sine pigmento aeruginoso, carbonaceum. Asci 8-spori. Sporae unisaeptae, late ellipsoideae vel globosae, 9-15 Jlm x 6-10 Jlm. Pycnidia globosa. Conidia bacilliformes, 2-4 Jlm x 1-1.5 Jlm. Thallus acida norstictica et connorstictica continens vel has substantias deficiens. DIAGNOSIS: MEXICO. Baja California: lighthouse area near Laguna Manuela; 28°15'OO*N, 114°07'OO"W; altitude: 100 m; on basalt; 21 February 1993; Nash 33979 (ASU - holotype designated here). TYPE: This species is named in honour of Dr. Christoph Scheidegger, Birmensdorf, Switzerland. TAXONOMIC NOTE: Figs. 1-7 and ＮＳｾＵ Thallus (Figs. 1, 2, ＩＳｾＵ crustose, usually rimose to rimose-areolate, thin and more or less continuous but distinctly epilithic, never chasmolithic, rarely becoming thickened and distinctly rimose-areolate to subsquamulose, not delimited by a hypothallus; surface matt to shiny, usually deep brown, rarely pale brown, usually smooth, but frequently roughened in areas where the surface appears eroded, epruinose, phenocorticate. Apothecia soon sessile, lecideine; proper margin conspicuously thickened, usually persistent, rarely excluded with age; disk black, epruinose, plane, rarely becoming slightly convex with age; exciple of aethalea-type (Figs. 3, 4) sensu Scheidegger (1993), that is, inner excipular hyphae narrow, hyaline, prosoplectenchymatous (textura oblita), similar in structure and orientation to the paraphyses, often more or less reduced and transient with the dull reddish brown hypothecium (leptoclinoides-brown, textura inricata); outer excipular hyphae parallel (textura oblita), cells moderately swollen and usually strongly carbonized with various amounts of brown pigments (cf. elachista-brown), pigmentation continuous with the epihymenium; hymenium hyaline, not inspersed; paraphyses simple to moderately branched, apically swollen, with a brown pigment cap (cf. elachista-brown). Asci 8-spored, clavate, Bacidia-type. Ascospores (Figs. 5, 6, 70) broadly ellipsoid to almost globose, never constricted, with obtuse ends, not curved, (9.0-)10.2-[11.7)13.2(-15.0) Jlm x (6.0-) 6.8-[7.7]-8.6(-10.0) Jlm (n = 81), one-septate; proper septum narrow, not thickening during spore ontogeny (Fig. 70), with septal pore canal, simple pore and undifferentiated pore plug; no ornamentation visible (in DIC); spore wall (Fig. 6) differentiated into smooth, thin perispore (0.10-0.20 Jlm), narrow intermediate layer (<10 Jlm), thick proper spore wall (0.50-1.50 Jlm) and moderately thickened endospore (0.20-0.40 Jlm). Pycnidia (Fig. 7) rare to common, globose, unilocular, at maturity almost entirely occupied by densely branched conidiophores; conidiogenous cells mostly terminal, rarely also intercalary (cf. conidiophore-type V sensu Vobis 1980); pycnidial ontogeny similar to the Umbilicaria-type (sensu Vobis 1980, Vobis and Hawksworth 1981); conidia (Fig. 7) simple, bacilliform, 2.0-4.5 Jlm x 1.0-1.5 Jlm (n = 69) (Fig. 72). No substances found or with norstictic and connorstictic acid (low concentrations only detected by TLC or HPLC). All spot tests negative (very rarely forming orange needle-shaped crystals with K if observed with the compound scope). UV-. Thallus without amyloid reaction, only the apothecia amyloid in Lugol's. CHEMISTRY: All specimens examined are from low elevations, growing on siliceous mineral-poor coastal rock. The holotype is erroneously labeled as growing "on basalt". The substrate is, however, a very fine grained andesite, with intermediate to silicic acidity, poor in bases. SUBSTRATE AND ECOLOGY: DISTRIBUTION: The species is currently known only from coastal southern California, USA, Baja California, and Baja California Sur, Mexico (Fig. 71). The species is superficially similar to B. pullata (Figs. 8, 9, ＬＩＳｾＵ and differences have been discussed in the note on that species. Table 1 gives an overview of the diagnostic characters available to distinguish all species discussed here. NOTES: MEXICO. Baja California: Ryan 21513, Marsh 6198, Nash 34569 (ASU). Baja California Sur: Nash 29800 (ASU). USA. California. Los Angeles Co.: Wetmore 73435 (MIN); Hasse Exs. 205 (CAS); Weber L-42660, L42178, L-42662, L-42163 (COLO); Nash 32093, 32100, MATERIAL EXAMINED: © 2004 NRC Canada Bungartz at al. 543 © 2004 NRC Canada 544 Can. J. Bot. Vol. 82, 2004 Figs. 15-19. Buellia prospersa (all specimens: Wetmore 71896). Fig. 15. Overview of the epilithic rimose-areolate thallus. Fig. 16. Close-up of Fig. 15. Fig. 17. Light micrograph of the athalea-type exciple (cross section in water): the thin outer part consists of swollen cells, which are carbonized with the dull brown pigment cf. elachista-brown. Fig. 18. Light micrograph of mature ascospores. Fig. 19. Light micrograph of filiform conidia. Figs. 20-28. Buellia sequax. Fig. 20. Chasmolithic granular thallus (Nash 33018). Fig. 21. Light micrograph of the athalea-type exciple (cross section in water): the thin outer part consists of swollen cells, which are carbonized with the dull brown pigment cf. elachista-brown (Nash 32187). Fig. 22. Light micrograph of a characteristically narrow, mature ascospore (Nash 32187). Fig. 23. Scanning electron microscope (SEM) micrograph of an exciple cross section (Nash 33018). Fig. 24. Light micrograph of conidiophores with bacilliform conidia (Nash 32187). Fig. 25. Transmission electron microscope (TEM) micrograph of a Bacidia-type ascus tip (for designation of the different layers see Bellemere 1994): a- and b-Iayer are missing because of fixation artifacts; (c) outer electron opaque c-layer; (dl) dl-Iayer, that is, the outer tholus, which has a distinctly laminated, fibrillar structure (in light microscopy this outer part stains deep blue with Lugol's iodine); (d2) d2-layer, that is, the inner tholus, which is not layered and more or less homogeneous (not staining in Lugol's iodine); and (oc) ocular chamber (Nash 32718). Fig. 26. TEM micrograph of a premature ascospore: (1) thin, smooth perispore, (2) intermediate layer, (3) proper wall, and (4) endospore (Nash 32718). Fig. 27. TEM micrograph of mature ascospore: (I) thick, fractured perispore, (2) intermediate layer, (3) proper wall, and (4) endospore. (Ryan 21689). Fig. 28. TEM micrograph of the spore septum (s) with a simple pore plug (p) (Nash 32718). 32174a, Nash 32208, Nash 32141 (ASU). Mendocino Co.: Tucker 35170 (SBBG). Monterey Co.: Weber 8242, 8237 (COLO). San Luis Obispo Co.: Tucker 36478 (SBBG); Nash 36982 (ASU). Santa Barbara Co.: Printzen 49, 76 (hb. Printzen); Weber L-80178 (COLO); Nash 32386, 41174, 41181, Ryan 31359. Crayton s. n. (ASU). Britton & Kemp (79), determined by B. Fink] has been annotated by Fink as the type of "Buellia substigmata Fink". This name has never been published by Fink and the specimen is not the type of Buellia substigmatea Mull. Arg. [Proceed. Roy. Soc. Edinburgh 11: 465 (1882)]. The MICH specimen has correctly been annotated by Imshaug as Buellia prospersa (Nyl.) Riddle. Buellia prospersa is the valid name for Rinodina (= Amandinea) lecideina, which is, however, not identical with Buellia lecidina Stein. ex Cohn [Kryptog. Flora von Schlesien 2(2) (1879)]. Even though the spelling of the species epithets "lecideina" and "lecidina" is almost identical, Buellia lecidina Stein. ex Cohn is a synonym of Rinodina occulta Korb. Buellia prospersa (Nyl.) Riddle Brook!. Bot Gardens Memoirs 1: 114 (1918) Lecidea prospersa Nyl., Flora 63: 127 (1880). Type: Virgin Islands: St. Thomas [original label data. S. Thomae Antillarum], 1878, Dr. Forel. (H-Nyl 9312! - lectotype selected here). Rinodina lecideina H. Mayrhofer & Poelt, Bibl. Lich. 12: Figs. 15-19. 112 (1979). Type: IRELAND. Kerry: Dingle Peninsula. Near the village Ballyoughteragh, N BailIe near Ballyferriter: on Thallus (Figs. 15, 16) crustose, rimose, moderately thin pasture walls [original label data: Eire/Ireland: Co. Ciarrai / and more or less continuous but distinctly developed and not Kerry, Corea Dhuibhne / Dingle-peninsula, Umgebung des chasmolithic, not delimited by a distinct hypothallus or with Weilers Ballyoughteragh N BailIe an Fheirtearaigh / Ballya black outline around the thallus; surface matt to more or ferriter, an Weidenmauem], August 1978, Poelt s. n. (GZU! less shiny, usually pale ivory and smooth, rarely gray and holotype). slightly roughened (probably as a result of surface damage), Amandinea lecideina (H. Mayrhofer & Poelt) Scheid. & epruinose, phenocorticate. H. Mayrhofer, in Scheidegger, Lichenologist 25: 342 (1993) Apothecia soon sessile, lecideine; proper margin excluded Buellia punctata f. crassior (Erichsen) Zahlbr., Cat. Lich. with age; disk black, epruinose, plane, becoming convex Univ. 8: 591 (1932). - Buellia punctata f. crassior with age; exciple of aethalea-type (Fig. 17) sensu ScheiErichsen, Das Hnke Untertraveufer: 151 (1932). Type: GERdegger (1993), that is, inner excipular hyphae narrow, MANY, Schleswig-Holstein: Lubeck, Coast of Dummershyaline, prosoplectenchymatous (textura oblita), similar in dorf, boulders along the beach below the village Stulperbank structure and orientation to the paraphyses, often more or [original label data: Deutschland, Schleswig-Holstein, Lubeck, Dummersdorfer Ufer, Strandblocke unterhalb Stulper- • less reduced and transient with the dull reddish brown hypothecium (leptoc1inoides-brown, textura inricata); outer bank], April 1928, Erichsen (HBG - holotype). excipular hyphae parallel (textura oblita), cells moderately Buellia punctata f. litoralis (Erichsen) Zahlbr., Cat. Lich. swollen and usually strongly carbonized with various Univ. 7: 397 (1931). - Buellia myriocarpa var. litoralis amounts of a brown pigment (cf. elachista-brown); pigmenErichsen, Verh. Bot. Ver. Provinz Brandenburg 72: 48 tation continuous with the epihymenium; hymenium hyaline, (1030). Type: GERMANY, Schleswig-Holstein: Eastcoast of not inspersed, paraphyses simple to moderately branched, the Island Alsen, near Kettingholz, on boulders along the apically swollen, with a brown pigment cap (cf. elachistabeach in the supralittoral zone [original label data: Deutschbrown). land, Schleswig-Holstein, Insel Alsen, Ostktiste bei KettingAsci 8-spored, clavate, Bacidia-type. Ascospores holz, an StrandblOcken der supralitoralen Zone], July 1932, (Figs. 18, 70) oblong to more or less ellipsoid, not conErichsen (HBG - holotype). stricted, with obtuse ends, not curved, (9.0-)12.0-[13.5]15.0(-16.0) Ilm x (5.0-)6.2-[7.3]-8.3(-9.5) Ilm (n = 91), TAXONOMIC NOTES: A specimen from MICH [rocks on a hill, one-septate; proper septum soon with a more or less persisChristiansted, St. Croix, March 17-25, 1923, collected by © 2004 NRC Canada Bungartz et al. tent thickening, becoming reduced only in old spores (i.e., with a distinct Physconia-type or Orcularia-type ontogeny); ornamentation microrugulate (conspicuous in DIe at an early stage of the ontogeny). Pycnidia rare, globose, unilocular; at maturity almost en- 545 tirely occupied by the long conidia and lined with short, scarcely branched conidiophores; conidiogenous cells terminal (conidiophore-type III sensu Vobis 1980); pycnidial ontogeny similar to the Roccella-type (sensu Vobis 1980, Vobis and Hawksworth 1981); conidia (Fig. 19) simple, © 2004 NRC Canada ｾ (J) Table 1. Diagnostic differences of Buellia christophii, Buellia prospersa, Buellia pullata, Buellia ryanii, Buellia sequax, and Buellia tergua. Thallus growth and color BueUia christophii Epilithic; pale to deep brown BueUia prospersa Epilithic, rarely vaguely chasmolithic; pale ivory BueUia pullata Epilithic; deep brown to gray BueUia ryanii Epilithic; olive gray to brownish olive BueUia sequox Chasmolithic, inconspicious; pale brown to grayish BueUia terguo Epilithic; "Ieatherbrown" @ ｾ IV ｾ UV reaction and chemistry Thallus morphology Conidia Exciple Spores Rimose to rimose-areolate, rarely subsquamulose; without hypothallus Bacilliform (short) Thick. persistent, brown Broad oblong (almost globose), equal wall thickening, early microrugulate UV negative, no xanthones, ± norstictic acid Rimose to rimose-areolate; usually without a hypothallus, rarely thallus surrounded by black outline Filiform (short to very long) Thin, persistent, brown Oblong-ellipsoid, persistent septum thickening, early microrugulate UV+ yellow to orange, usually with xanthones, ± norstictic acid, ± atranorin Rimose to rimose-areolate, (rarely subsquamulose); without hypothallus Filiform (short to very long) Thin, persistent, brown Oblong-elIipsoid, brief septum thickening, early microrugulate UV negative, no substances detected Rimose to rimcfse-areolate (rarely subsquamulose); with distinct arachnoid black hypothallus Bacilliform (short) Thin, excluded, aeruginose (N+ violet) Oblong-ellipsoid, no wall thickening, late microrugulate UV negative, no substances detected Granular to rimose; without hypothallus Bacilliform (short) Thin, persistent to excluded, brown Narrow oblong, no wall thickening, late microrugulate UV negative, no substances or ±Dorstictic acid, rarely with arthothelin Rimose to rimose-areolate; without hypothallus Bacilliform (short) Thin to moderately thick. persistent, aeruginose (N+ violet) Oblong-ellipsoid, no wall thickening, late microrugulate UV negative, no substances detected Q ? c... ｾ ?= <Xl .ｾ , go .!" I\:l § Bungartz et al. long bacilliform to filiform, 7.0-32.0 106) (Fig. 72). 547 ｾｭ x <1.0 ｭｾ (n = CHEMISTRY: No substances found, or with various amount of the following secondary metabolites: norstictic and connorstictic acid, atranorin, or several xanthones (HPLC: 4,5-dichloro-3-0-methylnorlichexanthone, thuringione, 4,5dichlorolichexanthone, 2,4,5-trichlorolichexanthone). Spot tests usually negative, rarely K+ yellow to orange, C+ orange. UV+ bright yellow to orange. Thallus without amyloid reaction, only the apothecia are amyloid in Lugol's. SUBSTRATE AND ECOLOGY: Growing on siliceous mineral-poor coastal rock (generally HCI-). DISTRIBUTION: Probably cosmopolitan but restricted to coastal areas. In the Sonoran Region presently known only from southern California, USA., and Baja California, Baja California Sur, Sonora and Sinaloa, Mexico (Fig. 71). NOTE: In general, B. prospersa can be easily recognized by a pale, distinctly epilithic thallus (Figs. 15, 16) and ascospores with a persistent median septum thickening (Figs. 18, 70). Buellia pullata is microscopically similar, but has a deep brown to dark gray thallus (Figs. 8, 9, 66), which does not react with UV light. No xanthones were detected in Buellia pullata. Although not previously reported, thalli of B. prospersa always react UV+ yellow to orange, and xanthones are characteristic for this species. Poorly developed thalli of B. prospersa sometimes appear more or less chasmolithic, and these specimens are then difficult to separate from forms of B. sequax with a well-developed thallus (Fig. 42). If filiform conidia (Fig. 19) can be found, the material is easily recognized, because B. sequax has bacilliform conidia (Fig. 24). In both species pycnidia are, however, quite rare. Young ascospores of B. sequax are typically distinctly narrowly oblong and not ornamented (Figs. 22, 70). Only overmature and often disintegrating ascospores show a weak ornamentation, whereas a microrugulate ornamentation usually develops in young ascospores of B. prospersa (Fig. 70). Young ascospores of B. prospersa typically have a distinctly thickened median septum (Figs. 18, 70), which can become very pronounced in some specimens. However, in some specimens it is difficult to find spores at this stage of the ontogeny. Median thickening of the spore septum is absent from all stages of ascospore development in B. sequax (Fig. 70). In the Sonoran Desert Region, B. prospersa is restricted to coastal California, USA, and Baja California, Mexico (Fig. 71). Buellia sequax is widely distributed throughout the Sonoran Desert (Fig. 71). EXAMINED: MEXICO. Baja California: Scheidegger s. n. (hb. Scheidegger); Nash 33982 (ASU). Baja California Sur: Nash 12653, 12755 (ASU). Jalisco: Nash 20765 (ASU). Sinaloa: Moberg 10267 (UPS); Wetmore 71896 (MIN); Nash 33680, 10088, 12248 (ASU); Sonora: Nash 25625, 12518, 12521, 10964 (ASU). ITALY. Sardinia: Scheidegger Inv. Nr. 11097 (hb. Scheidegger). NEW ZEALAND. South Island. Canterbury: Nash 19098 (ASU); Blaha 200 (GZU). Southland: Blaha 152 (GZU). USA. California. Santa Barbara Co.: Nash 41312 (ASU). Ventura Co.: Nash 38657 (ASU). MATERIAL Buellia pullata Thck. Lich. Californ. p. 26 (1866) TYPE: USA. California: Rocks on the coast. Bolander 131, (FH-3280! - lectotype selected here!), Bolander 181, 150 (FH-3280! - syntypes). TAXONOMIC NOTES: This species is not synonymous with B. punctata, as suggested by Imshaug (1951). Several speci- mens from FH are mounted together on one large herbarium sheet with the FH accession number 3280. A type specimen is not designated in the protologue. From the available material we therefore select the specimen labeled Bolander (131) as the lectotype. Even though the other specimens are mounted on the same sheet and have the same FH accession number the specimens Bolander 181 and 150 cannot be regarded as isolectotypes, because they have different collection numbers. Bolander 179, also mounted on the same sheet, has bacilliform conidia and it belongs to Buellia christophii. Figs. 8-14 and 64-69. Thallus (Figs. 8, 9, 64-69) crustose, usually rimose to rimose-areolate, thin and more or less continuous but distinctly epilithic, never chasmolithic, rarely becoming thickened and distinctly rimose-areolate to subsquamulose, not delimited by a hypothallus; surface matt to more or less shiny, usually deep brown, rarely pale brown, smooth, epruinose, phenocorticate. Apothecia soon sessile, lecideine; proper margin excluded with age; disk black, epruinose, initially plane, soon becoming convex; exciple of aethalea-type (Figs. 11, 12) sensu Scheidegger (1993), that is, inner excipular hyphae narrow, hyaline, prosoplectenchymatous (textura oblita), similar in structure and orientation to the paraphyses, often more or less reduced and transient with the dull reddish brown hypothecium (leptoclinoides-brown, textura inricata); outer excipular hyphae parallel (textura oblita), cells moderately swollen and usually strongly carbonized with various amounts of a brown pigment (cf. elachista-brown); pigmentation continuous with the epihymenium; hymenium hyaline, not inspersed; paraphyses simple to moderately branched, apically swollen, with a brown pigment cap (cf. elachistabrown). Asci 8-spored, clavate, Bacidia-type. Ascospores (Figs. 14, 70) oblong to more or less ellipsoid, not constricted, with obtuse ends, not curved, (9.0-)11.1-[12.5]x (5.0-)6.2-[7.1]-8.0(-10.0) ｾｭ (n = 91), 13.8(-15.0) ｭｾ one-septate; proper septum soon thickening during spore ontogeny, becoming narrow with age (Le., with an indistinct Physconia-type ontogeny), with septal pore canal, simple pore and undifferentiated pore plug; ornamentation microrugulate (conspicuous in DIC at an early stage of the ontogeny); spore wall (Fig. 14) differentiated into smooth to narrow intermediate cracked, thin perispore «0.20 ｾｭＩＬ moderately thick proper spore wall (0.3layer «0.10 ｾｭＩＬ and moderately thickened endospore (0.200.5 ＬＩｭｾ 0.40 ＮＩｭｾ Pycnidia (Fig. 13) common, globose, unilocular; at maturity almost entirely occupied by densely branched conidiophores; conidiogenous cells mostly terminal, some intercalary © 2004 NRC Canada 548 Can. J. Bot. Vol. 82, 2004 Figs. 29-34. Buellia ryanii. Pig. 29. Overview of the epilithic areolate thallus delimited by a distinct, black arachnoid hypothallus (Nash 38368 - holotype). Pig. 30. Close-up of Fig. 29: young apothecia aspicillioid, emerging from the thallus partially covered by a thalline veil (arrows). Pig. 31. Light micrograph of the athalea-type exciple (cross section in water): the thin, dull olive outer part consists of swollen cells, which are carbonized by the dull brown pigment cf. elachista-brown and the aeruginose pigment cinereorufagreen (Nash 38368 - holotype). Pig. 32. Light micrograph of mature ascospores (Nash 38368 - holotype). Fig. 33. Light micrograph of a pycnidium with bacilliform conidia (Nash 32448b). Fig. 34. Light micrograph of conidiophores with bacilliform conidia (Nash 32448b). Figs. 35-40. Buellia tergua (all specimens: Nash 38368 - holotype). Fig. 35. Rimose-areolate thallus. Fig. 36. Light micrograph of the athalea-type exciple (cross section in water): the moderately thickened, strongly aeruginose outer part consists of swollen cells, which are carbonized by the dull brown pigment cf. elachista-brown and the aeruginose pigment cinereorufa-green. Pig. 37. Light micrograph of an exciple (cross section in HN03): the aeruginose pigment reacts violet and is partially diluted, thus the structure of the hyphal cells becomes more distinct Fig. 38. Premature (p) and mature (m) ascospore. Fig. 39. Light micrograph of a pycnidium with bacilliform conidia. Fig. 40. Light micrograph of conidiophores with bacilliform conidia. (cf. conidiophore-type III sensu Vobis 1980); pycnidial ontogeny similar to the Roccella-type (sensu Vobis 1980, Vobis and Hawksworth 1981); conidia (Fig. 13) simple, predominantly filiform, 6.0-43.0 I.lm x <1.0 I.lm (n = 156). No substances found. All spot tests negative. Thallus without amyloid reaction, only the apothecia are amyloid in Lugol's. CHEMISTRY: SUBSTRATE AND ECOLOGY: Growing on siliceous mineral-poor rock (generally HCl-). DISTRffiUTION: Fairly common along the coast of southern California, USA, Baja California, and Baja California Sur, Mexico. One specimen has also been found further inland in Sonora, Mexico (Fig. 71). The thallus morphology (Figs. 8, 9, 64-69) of this species is very similar to that of B. christophii (Figs. 1, 2, 55-62). Pycnidia are relatively common in both species and if filiform conidia are found (Fig. 13) the material is, thus, reliably distinguished from B. christophii, which has bacilliform conidia (Fig. 7). The two species can, however, also be clearly distinguished by other characters. Buellia christophii has lecideine apothecia with a thick, rather persistent margin (Figs. 1-3). Buellia pullata has a thin margin, which relatively soon becomes excluded (Figs. 8, 9, 11). Although the structure of the proper exciple is virtually identical (compare Fig. 4 with Fig. 12), the layer of carbonized swollen outer cells is at least twice as thick in B. christophii (Fig. 3). Young ascospores of B. pullata are oblong to ellipsoid (Fig. 70); in B. christophii, young ascospores are rather broad, and often appear almost globose (Figs. 5, 70). Spores of B. christophii have no septum thickening (Fig. 70), whereas a median thickening of the spore septum can usually be observed at least in some of the younger spores of B. pullata (Fig. 70). BuelUa ryanii Bungartz sp. nov. DIAGNOSIS: Thallus saxicolus, crustaceus, areolatus vel subsquamulosus, tenuis vel crassus, olivaceus. Hypothallus atratus arachnoideus. Apothecia erumpentia vel sessilia, lecideina, marginibus propriis tenuibus. Excipulum tenue, fulvocaeruleum, pigmentum aeruginosum continens, carbonaceum. Asci 8-spori. Sporae unisaeptae, ellipsoideae vel oblongae, 9-13 I.lm x 4-8 I.lm. Pycnidia globosa. Conidia bacilliformia, 2-5 I.lm x 1-1.5 I.lm. Materiae chimicae nullae. USA. California. Santa Barbara: Santa Cruz Island, 4.5 km E of radar station, ridge crest down N slope; 34°0'15"N, 119°37'30"W; altitude: 287 m; on schist in oakpine woodland, 9 January 1994; Nash 32448b (ASU holotype designated here). TYPE: NOTES: MEXICO. Baja California: Scheidegger s. n. (hb. Scheidegger); Nash 38267, 38458, 38513b (ASU); Wetmore 75732 (MIN). Sonora: Wetmore 71644 (MIN). USA. California. Los Angeles Co.: Weber L-42114 (COLO); Nash 32174 (ASU); Mendocino Co.: Nash 25476, 11426, 11424b (ASU). Monterey Co.: Nash 18896 (ASU). San Diego Co.: Bratt 8658 (SBBG). San Luis Obispo Co.: Thcker 28829B (SBBG). Santa Barbara Co.: Bratt 6394, Thcker 34719, Tucker 35711 (SBBG); Nash 32672, 41099, 41227, 41249,41311,33012 (ASU), Wetmore 73713 (MIN). Ventura Co.: Nash 37059, 37015 (ASU); D.E. Baltzo 7215 (UC). MATERIAL EXAMINED: The species is named in memory of Dr. Bruce D. Ryan, the third author of this publication, who died from cancer before this article was printed. TAXONOMIC NOTE: Figs. 29-34. Thallus (Figs. 29, 30) crustose, areolate to subsquamulose, usually delimited by a distinct black arachnoid hypothallus; surface matt to more or less shiny, deep olive gray to brownish olive, smooth, epruinose, phenocorticate. Apothecia initially immersed appearing aspicillioid, soon bursting through the thallus surface and becoming adnate to sessile, lecideine, rarely with remains of necrotic thalline material attached to the margin (thalline veil, Fig. 30); proper margin more or less persistent, rarely excluded with age; disk black, epruinose, plane, rarely becoming convex with age; exciple of aethalea-type (Fig. 31) sensu Scheidegger (1993), that is, inner excipular hyphae narrow, hyaline, prosoplectenchymatous (textura oblita), similar in structure and orientation to the paraphyses, often more or less reduced and transient with the dull reddish brown hypothecium (leptoclinoides-brown, textura inricata); outer excipular hyphae parallel (textura oblita), cells moderately swollen and usually strongly carbonized with various amounts of brown and aeruginose (HN0 3+ violet) pigments (cf. elachista-brown and cinereorufa-green); pigmentation continuous with the epihymenium; hymenium hyaline, not inspersed; paraphyses simple to moderately branched, apically swollen, with a brown pigment cap (cf. elachistabrown) and a diffuse aeruginose pigment (HN0 3+ violet, cinereorufa-green). © 2004 NRC Canada Bungartz at al. Asci 8-spored, clavate, Bacidia-type. Ascospores (Figs. 32, 70) broadly oblong to ellipsoid, with obtuse ends, not curved, mature spores slightly constricted, (9.0-)10.5[11.3]-12.2(-13.0) J.lm x (4.0-)5.6-[6.2]-6.8(-8.0) J.lD1 (n = 100), one-septate; proper septum narrow, not thickening during spore ontogeny; ornamentation absent in immature and 549 premature spores, microrugulate in mature spores (best seen in DIe). Pycnidia (Fig. 33) rare, globose, unilocular; at maturity almost entirely occupied by densely branched conidiophores; conidiogenous cells mostly terminal, but some also intercalary (cf. conidiophore-type V sensu Vobis 1980); pycnidial © 2004 NRC Canada 550 Can. J. Bot. Vol. 82, 2004 Figs. 41-54. Chasmolithic growth in Buellia sequax on various substrates. Fig. 41. Light micrograph of a thallus on a sandstone, almost entirely composed of quartz (substrate cross section): the poorly delimited thallus is growing between the quartz crystals (Nash 33018). Fig. 42. Light micrograph showing the thallus aspect of well-developed cortical granules (arrows) between feldspar crystals (Nash 32718). Fig. 43. Light micrograph of a thallus with corticate granules growing between feldspar crystals (substrate cross section): arrows point at the algal layer (Nash 32718). Fig. 44. Scanning electron microscope (SEM) micrograph of two apothecia and poorly delimited hyphae growing between quartz crystals of a sandstone (specimen critically point dried, gold coated; Nash 33018). Fig. 45. Light micrograph of a chasmolithic thallus almost entirely hidden in the fissures of a foliated metamorphic, low-grade phyllite (Weber L-42776). Fig. 46. Light micrograph of a thallus on phyllite (substrate cross section): the phyllite contains some quartz crystals, and it is rich in mica and thus more or less foliated; hyphae have been stained with lactophenol cotton blue, which results in the darkening of the upper inhabited layers of the substrate (Weber L-42776). Fig. 47. Natural break of a sandstone where the fractures between the quartz crystals are inhabited by the thallus (Nash 40151). Fig. 48. SEM micrograph of the chasmolithic thallus on phyllite (critically point dried, gold coated): the minerals appear somewhat coated but no distinct hyphae are visible (Weber L-42776). Fig. 49. Cross section of the specimen growing on phyllite (critically point dried, gold coated): the thallus hyphae (t) are closely conglutinated between the mineral crystals (c) of the substrate (Weber L-42776). Fig. 50. SEM micrograph of a thallus growing on sandstone (critically point dried, gold coated): the thallus consists of hyphae and algal cells growing irregularly between the quartz crystals (Ryan 21689). Fig. 51. Detail of Fig. 50: thallus hyphae (t) growing between quartz crystals (c). Fig. 52. Detail of Fig. 51: the haustorial hyphae (h) are closely wrapped around algal cells (a); these thallus areas are developed between the quartz crystals (c) of the substrate. Fig. 53. SEM of a chasmolithic thallus on sandstone (cross section resin embedded, polished, and carbon coated): (a) apothecium, (t) thallus, (c) quartz crystal (Ryan 21689). Fig. 54. SEM of a chasmolithic thallus on a more or less metamorpic volcanic rock with fractured and sheared mineral crystals (cross section resin embedded, polished, and carbon coated): (s) stipe of the apothecium, (e) exciple, (t) thallus, (c) sheared mineral crystals (Nash 32187). ontogeny similar to the Umbilicaria-type (sensu Vobis 1980, Vobis and Hawksworth 1981); conidia (Fig. 34) simple, bacilliform to ellipsoid, 2.0-5.0 Jlm x 1.0-1.5 Jlm (n = 73) (Fig. 72). No substances found and all spot tests negative. Thallus without amyloid reaction, only the apothecia are amyloid in Lugol's. CHEMISTRY: SUBSTRATE AND ECOLOGY: All specimens examined grow on mineral-poor rock substrates such as quarzite and schist in open oak-conifer forest. The species is currently known only from Santa Cruz Island, along the coast of southern California, USA, and from Isla de Guadalupe, off the coast of Baja California, Mexico. It has not been found close to the sea shore but in open, dry oak-conifer woodland. DISTRIBUTION: Scheidegger (1993) mentions the presence of pigment A (= cinereorufa-green) as characteristic of the aethaleatype exciple. All species discussed here, have an exciple with the structure of the aethalea-type, but the pigment cinereorufa-green is only present in the exciple and epihymenium of B. ryanii and B. tergua. The two species are quite similar and currently only known from a few localities off the coast of Baja California, Mexico and southern California, USA. Buellia ryanii has an areolate to subsquamulose, deep olive gray to brownish olive thallus with a distinct arachnoid hypothallus (Figs. 29, 30). Buellia tergua has a leather brown rimose-areolate thallus with no hypothallus (Fig. 35). The apothecial ontogeny is very similar. Apothecia are initially immersed and the margin is covered by the surrounding thallus. Thus, these apothecia look aspicillioid (Fig. 30). Soon apothecia erupt from the surrounding thallus and become adnate. Rarely, thallus material remains attached to the lecideine margin as a thalline veil. The pigment cinereorufa-green is strongly concentrated in the exciple and epihymenium of B. tergua and in cross section clearly visible as deeply aeruginose. In contrast, the NOTES: exciple and epihymenium of B. ryanii is dull olive-brown and the presence of cinereorufa-green may thus be over· looked if not tested with HN03 • The spores of the two species have a similar ontogeny and become constricted with maturity (Fig. 70). However, spores of B. tergua are distinctly larger (10-15 Jlm x 6-9 Jlm) than those of B. ryanii (9-13 Jlm x 4-8 Jlm). The ornamentation of Buellia tergua spores develops early; it becomes visible at the premature stage and is distinct at maturity (Fig. 70). In B. ryanii spore ornamentation is barely discernible in mature spores but well developed in overmature spores (Fig. 70). The conidia of B. ryanii are slightly smaller (2-5 Jlm) than those of B. tergua (4-7 Jlm) (Fig. 72). USA. California. Santa Barbara Co.: Tucker 35781A, Bratt 3488b (SBBG); C.M. Wetmore 74077 (MIN); Nash 32445, Ryan 31621 (ASU). MATERIAL EXAMINED: Buellia sequax (Ny!.) Zahlbr. Cat. Lich. Univ. 7: 410 (1931). Lecidea sequax Ny!., Flora, Jena 58: 302 (1875) TYPE: FRANCE, Hautes Alpes: on quartzite along the banks of the river Vienne near Moulin de l' Aiguille [Quartz sur les coteaux de la Vienne pres du Moulin de l'Aiguille], 11 January 1872, Lamy 1083 [H-NYL 9538 - lectotype selected by Scheidegger (1993)], H-NYL 9539 - isolectotype, M0023804! - isolectotype]. Buellia abstracta (Ny!.) H. Olivier, Bull. Acad. Intren. Geogr. Bot. 12: 176 (1903). - Lecidea abstracta Ny!., Flora, Jena 66: 102 (1883). Type: FRANCE, Pyrennees: Cauterets, E. Lamy (H-NYL 9740 - holotype, M-0023906! - isotype). B. punctatula Malme, Arkiv fOr Botanik 21a(14):9 & 14 (1927). Type: PARAGUAY. ＺｮＶｩｾ ｵｳａ "Zapitapunta", on sunny sandstone [original label data: "Zapitapunta", ad rupes arenarias sat apricas], 21 July 1893, Malme 1424 (S! lectotype selected here). Buellia saxicola de Lesd., Ann. Crypt. Exot. 5: 127 (1932). Type: USA. New Mexico. San Miguel: around Las © 2004 NRC Canada 551 Bungartz et al. Vegas, Agua Zarca, on siliceous rocks [original label data: environs de Las Vegas, Agua Zarca, sur roches siliceusesl, Arsene Brouard 1930 (W acquisition 1934 no. 726! -lectotype selected here; UPS!, S! - isolectotypes). TAXONOMIC NOTES: The type material of B. saxicola is not identical with B. punctata as suggested by Imshaug (1951). The lectotypification of the material designated by Bouly de Lesdain is necessary, because it can generally be assumed that his entire holotype collection, originally located in Dunkerque (France), has been destroyed during the Second World War. The lectotype from Vienna was selected be© 2004 NRC Canada 552 cause this is the only material that still has a few intact pycnidia. Mayrhofer and Moberg (2002) list Buellia myriocarpa (DC.) De Not. as a synonym of Buellia punctata (= Amandinea punctata). In 1903, Hasse determined his exsiccati specimen no. 204 as Buellia myriocarpa (Lamb & DC.) Mudd. The exsiccati specimen at ASU is identical with B. sequax. Type material of B. myriocarpa was, however, not examined. Figs. 20--28 and 41-54. Thallus (Figs. 20, 41-54) crustose, thin and discontinuous, more or less chasmolithic, that is, forming inconspicuous, poorly delimited granules hidden among the mineral grains of the substrate; thicker thalli rarely becoming continuous in parts, rimose to very rarely rimose-areolate, not delimited by a hypothallus; surface matt, pale brown to greyish, smooth to slightly roughened, epruinose, phenocorticate. Apothecia soon becoming adnate to sessile; lecideine; proper margin soon excluded with age; disk black, epruinose, plane, becoming convex with age; exciple of aethaleatype (Figs. 21, 23) sensu Scheidegger (1993), that is, inner excipular hyphae narrow, hyaline, prosoplectenchymatous (textura oblita), similar in structure and orientation to the paraphyses, often more or less reduced and transient with the dull reddish brown hypothecium (leptoclinoides-brown, textura inricata); outer excipular hyphae parallel (textura oblita), cells moderately swollen and usually strongly carbonized with various amounts of brown pigment (cf. elachista-brown); pigmentation continuous with the epihymenium; hymenium hyaline, not inspersed, paraphyses simple to moderately branched, apically swollen, with a brown pigment cap (cf. elachista-brown). Asci 8-spored, clavate, Bacidia-type (Fig. 25). Ascospores (Figs. 22, 26-28, 70) distinctly narrowly oblong (Fig. 22), becoming ellipsoid with age, not or rarely constricted with age, with obtuse ends, not curved, (10.0--)10.7-[11.7]12.7(-14.0) Jlm x (3.0--)3.8-[4.6]-5.4(-6.0) Jlm (n = 50), one-septate, proper septum narrow, not thickened during spore ontogeny, with septal pore canal, simple pore and undifferentiated pore plug (Fig. 28); ornamentation absent from the early spore ontogeny (Fig. 26), becoming visible in mature spores (Fig. 27); spore wall differentiated into smooth to fissured, thin perispore (0.1-0.2 Jlm), narrow intermediate layer «0.1 Jlm), thick proper spore wall (0.3-0.5 Jlm) and moderately thickened endospore (0.2-0.3 Jlm). Pycnidia rare, globose, unilocular; at maturity almost entirely occupied by densely branched conidiophores; conidiogenous cells intercalary and terminal (conidiophore-type V sensu Vobis 1980); pycnidial ontogeny similar to the Umbilicaria-type (sensu Vobis 1980, Vobis and Hawksworth 1981); conidia (Fig. 24) simple, bacilliform, 2.0--4.0 Jlm x 1.0--1.5 Jlm (n = 50) (Fig. 72). No substances found or with norstictic acid or arthothelin (low concentrations only detected by TLC or HPLC). All spot tests negative (very rarely forming orange needle-shaped crystals with K if observed with the compound scope). UV-. Thallus without amyloid reaction, only the apothecia amyloid in Lugol's. Can. J. Bot. Vol. 82, 2004 SUBSTRAtE AND ECOLOGY: Growing on a large variety of siliceous rocks (generally HCl-). The species is widely distributed throughout the North American Southwest. In the Sonoran Desert Region, specimens have been found from coastal up to subalpine localities (Fig. 71). DISTRIBUTION: Specimens of B. sequax with an unusually welldeveloped thallus are sometimes superficially similar to B. prospersa and diagnostic differences have been discussed under that species. NOlES: AUSTRIA. Tirol: Arnold s. n. (M0061309). MEXICO. Baja California: Scheidegger s. n. (hb. Scheidegger), Nash 26137, 38513a, 40151, 26336, 34608 (ASU). Baja California Sur: Scheidegger s. n. (hb. Scheidegger); van den Boom 25063 (hb. van den Boom); Nash 33718, 33858, 40071, 40101, 26144 (ASU) Sonora: Ryan 21689 (ASU). ITALY. Elba: Albertshofer 6723 (M0061306), Triebel & Rambold 6209 (M-0061304), Albertshofer s. n. (M-0061303). Sicily: Doppelbaur (M0061307). SPAIN. Murcia: Doppelbaur (M-0061305). USA. Arizona. Apache Co.: Ryan 19184 (ASU) Cochise Co.: Weber S-8754 (COLO); Darrow 1970 (ASU) Coconino Co.: Nash 35171, Boykin 2725 (ASU) Gila Co.: Nash 28487, Schramm 225 (ASU) Maricopa Co.: Scheidegger s. n. (hb. Scheidegger); Nash 9605a (ASU) Pinal Co.: Scheidegger s. n. (hb. Scheidegger). California. Los Angeles Co.: Wetmore 73249 (MIN); Hasse L-78819 (COLO); Weber L-42778, L42776 (COLO); Bratt 10196 (SBBG); Nash 32187, Hasse Exs. 204, B.D. Ryan 30929b (ASU); Hasse Exs. 53 (CAS) Merced Co.: Tucker 28820 (SBBG), Tucker 28815 (ASU). Monterey Co.: Nash 8035 (ASU). Orange Co.: Weber L42052 (COLO). San Diego Co.: van den Boom 25192 (hb. van den Boom); Bratt 3592 (ASU) San Luis Obispo Co.: Tucker 28829A, 36471 (SBBG); Tavares 1409 (UC). Santa Barbara Co.: Tucker 35689A, 35845, 35781, 36463, 35781, Bratt 11388, 5709, 3641, 3643, 10316, 3488a (SBBG); Printzen 74, 76 (hb. Printzen), Nash 32352, 32656, 32747, 32837, 41392, 41423, 33016, 33018, 32890; Bratt 3488, 7681, 8879A, Ryan 31216 (ASU). Ventura Co.: Bratt 10392, Hamber 7 (SBBG) New Mexico. San Juan Co.: Nash 16225, 16396 (ASU). Valencia Co.: Nash 16004 (ASU). Utah. Washington Co.: Nash 15316 (ASU). Wyoming. Platte Co.: Ryan 14306 (ASU). SPECIMENS EXAMINED: Buellia tergua Bungartz sp. nov. DIAGNOSIS: Thallus saxicolus, crustaceus, rimosus vel rimosoareolatus, tenuis vel crassus, alutaceus, sine hypothallo. Apothecia erumpentia vel sessilia, lecideina, marginibus propriis tenuibus. Excipulum tenue, aeruginosum, pigmentum aeruginosum continens, carbonaceum. Asci 8-spori. Sporae unisaeptae, ellipsoideae vel oblongae, 10--15 Jlm x 6-9 Jlm. Pycnidia globosa. Conidia bacilliformia, 4-7 Jlm x 11.5 Jlm. Materiae chimicae nullae. CHEMISTRY: MEXICO. Baja California, Isla de Guadalupe: SE of southern peak; 28°57'30"N, 118°15'00"W; altitude 600 m; on basalt, shaded exposure within canyon with rocky outcrops and some native perennials; 3 January 1996, Nash 38368 (ASU - holotype designated here). TYPE: © 2004 NRC Canada 553 Bungartz et al. The species name is derived from the Latin word for leather, tergus. species with cinereorufa-green in the apothecia and diagnostic differences are presented in the notes on that species. Figs. 35-40. MATERAL EXAMINED: TAXONOMIC NOTE: MEXICO. Baja California: Moberg 8588 (UPS); Nash 38366a, Nash 8412, 8736, 38442 (ASU). Thallus (Fig. 35) crustose, rimose to rimose-areolate, not delimited by a distinct hypothallus; surface matt, usually deep brown and smooth, epruinose; phenocorticate. Apothecia initially immersed appearing aspicillioid, soon bursting through the thallus surface and becoming adnate to sessile, lecideine, rarely with remains of necrotic thalline material attached to the margin (thalline veil); proper margin more or less persistent, rarely excluded with age; disk black, epruinose, plane, rarely becoming convex with age; exciple of aethalea-type (Figs. 36, 37) sensu Scheidegger (1993), that is, inner excipular hyphae narrow, hyaline, ﾭ｣･ｾｰｯｳｲ tenchymatous (textura oblita), similar in structure and onentation to the paraphyses, often more or less reduced and transient with the dull reddish brown hypothecium (leptoclinoides-brown, textura inricata); outer excipular hyphae parallel (textura oblita), cells moderately swollen and usually strongly carbonized with various amounts of brown and aeruginose (HN0 3+ violet) pigments (cf. elachista-brown and cinereorufa-green); pigmentation continuous with the epihymenium; hymenium hyaline, not inspersed; paraphyses simple to moderately branched, apically swollen, with a brown pigment cap (cf. elachista-brown) and a diffuse aeruginose pigment (HN03+ violet, cinereorufagreen). Asci 8-spored, clavate, Biatora-type. Ascospores (Figs. 38, 70) broadly oblong to ellipsoid, with obtuse ends, not curved, mature spores slightly constricted, (10.0-)11.1[12.2]-13.4(-15.0) Jlm x (6.0-)6.0-[6.6]-7.3(-9.0) Jlm (n = 68), one-septate; proper septum narrow, not thickening during spore ontogeny (Le., Buellia-type); ornamentation absent in immature and premature spores, microrugulate in mature spores (best seen in DIC). Pycnidia (Fig. 39) rare, globose, unilocular; at maturity almost entirely occupied by densely branched conidiophores; conidiogenous cells intercalary and terminal (conidiophoretype V sensu Vobis 1980); pycnidial ontogeny similar to the Umbilicaria-type sensu Vobis 1980, Vobis and Hawksworth 1981); conidia (Fig. 40) simple, bacilliform, 4.0-7.0 Jlm x 1.0-1.5 Jlm (n = 50). CHEMISTRY: No substances found and all spot tests negative. Thallus without amyloid reaction, only the apothecia are amyloid in Lugol's. SUBSTRATE AND ECOLOGY: On siliceous volcanic coastal rock. Presently known only from Isla Guadalupe, off the coast of Baja California, and two localities on the Pacific coast of mainland Baja California, Mexico (Fig. 71). DISTRIBUTION: The thallus of B. tergua is very similar to B. pullata (Figs. 8, 9) and B. christophii (Figs. 1, 2), even though .these two species are generally darker brown. Apothecia of B. tergua, erupt from the thallus and young apothecia thus appear aspicillioid. In B. pullata and B. christophii, apothecia emerge more gradually and young apothecia appear immersed but not aspicillioid. Buellia ryanii is the only other NOTES: Discussion Thallus variation: chasmolithic versus epilithic growth All species discussed here are often described as having a scant or obsolete thallus and were previously interpreted as poorly developed forms of B. punctata rather than being assigned an independent species rank. However, in the Sonoran Desert the saxicolous species of the "Buellia punctata" group show considerable diversity. The thallus morphology of the species is quite diagnostic and far less variable than commonly implied. The extent to which a thallus is developed on the surface or partially hidden within crevices, pores, or fissures is necessarily related to substrate composition and structure. Both epilithic and chasmolithic thalli clearly show some degree of substrate penetration, provided that enough space is available (Figs. 41-69). The species examined here all grow on mineral-poor weathering rinds of various siliceous rocks. Although the composition of these substrates varies considerably, they are all poor in nutrients such as iron, magnesium, or calcium, and no calcium carbonates were found. The fungal hyphae are apparently not able to dissolve their substrate and instead, hyphal penetration is confined to the. ﾭｾｐＮ existing crevices (e.g., Figs. 53, 54, 60, 61). Both epilithic and chasmolithic thalli can thus be distinguished from truly endolithic lichens, which develop entirely within their substrate and are at least partially able to dissolve some of the minerals (Bungartz et al. 2004). Generally, chasmolithic growth (Figs. 41-54) and epilithic growth can be distinguished (Figs. 55-69): epilithic thalli are well developed on the substrate surface, internally distinctly stratified (Figs. 10, 63), and have a more or less con: tinuous surface (Fig. 58, 59, 64, 65). Although these thallI may be thin and inapparent, they are nevertheless largely covering the substrate (Figs. 1, 2, 8, 9, 55, 66). In contrast, chasmolithic thalli grow poorly differentiated among the mineral grains (Figs. 41, 44, 47, 50), these thalli are less stratified and sometimes only consist of loosely associated hyphae (Figs. 50-52). Even well-developed chasmolithic thalli (Figs. 42, 43) do not establish a continuous surface crust. Among the species examined here, only B. sequax may consistently be referred to as chasmolithic, that is, the major part of the thallus remains hidden ｡ｭｯｾｧ the m,ineral grains, even though some areas may develop Into a dIscontinuous crust (Fig. 47). Specimens of B. sequax with welldeveloped thallus areas can be distinguished from the other epilithic species by the pale thallus colour. Only poorly developed or damaged thalli of B. prospersa may be confused with these well-developed specimens of B. sequax. Comparing the types of B. abstracta and B. sequax, Scheidegger (1993) argued that B. abstracta s. str., with narrow ｡ｳｾｯﾭ spores and a chasmolitic thallus, cannot reasonably be ､ｾｳﾭ tinguished from epilithic and often areolate mat:nal assigned to B. sequax s. str. He therefore synonymIzed B. abstracta with B. sequax, which he regarded as a highly © 2004 NRC Canada 554 Can. J. Bot. Vol. 82, 2004 Figs. 55-69. Variation of epilithic growth in Buellia christophii (Figs. 55-ti3; Nash 32100, on silica-rich rhyolithe, Nash 33979 :... holotype, on a vesicular, fme-grained volcanic andesite) and B. pullata (Figs. 64-ti9; Nash 32672, on a fme-grained sandstone with more or less metamorphic, laminated, and flattened grains). Fig. 55. Light micrograph showing the rimose-areolate epilithic thallus aspect on rhyolithe (Nash 321(0). Fig. 56. Light micrograph of a thallus on rhyolite (substrate cross section): the thallus is distinctly developed on the surface (Nash 321(0). Fig. 57. Detail of Fig. 56: a single areole with an apothecium. Fig. 58. Scanning electron microscope (SEM) micrograph of a thallus on andesite (substrate cross section critically point dried, gold coated): the thin thallus is distinctly developed on the surface (Nash 33979 - holotype). Fig. 59. Detail of Fig. 58: the hyphae are restricted to the substrate surface. 60. Backscattered SEM micrograph (resin-embedded, polished, carbon coated): (a) apothecium, (t) epilithic thallus, and (h) hyphae penetrating natural crevices (Nash 32100). Fig. 61. Detail of Fig. 60: (t) thallus, (c) substrate crevice penetrated by a bundle of hyphae; the arrow points to a narrow fissure. Fig. 62. SEM micrograph of a thallus on andesite (substrate cross section resinembedded, polished, carbon coated): (a) apothecium, (t) thallus, (s) substrate (Nash 33979 - holotype). Fig. 63. Detail of Fig. 62: (c) thallus cortex, (a) alga, (q) quartz crystal. Fig. 64. SEM micrograph of the rimose-areolate thallus on metamorphic sandstone (specimen critically point dried, gold coated). Fig. 65. Detail of Fig. 64. Fig. 66. Light micrograph of the rimose-areolate thallus. Fig. 67. Light micrograph of a thallus on sandstone (substrate cross section): the thallus areoles are distinctly developed on the surface. Fig. 68. SEM micrograph of a thallus on sandstone (substrate cross section critically point dried, gold coated); the thallus areoles are distinctly developed on the substrate surface: (a) apothecium, (t) thallus, (c) natural cavity penetrated by lichen hyphae. Fig. 69. SEM micrograph of a thallus on sandstone (substrate cross section resin-embedded, polished, carbon coated): (t) thallus, (s) substrate. polymorphic species. We agree that both taxa can be synonymized. Buellia sequax is, however, far less variable than this statement may imply. Chasmolithic growth is not merely the result of substrate variation, because B. sequax consistently forms chasmolithic thalli on a variety of different substrates. We have examined type material of the two taxa as well as material from M annotated by Scheidegger. Buellia sequax s. str. indeed shows more or less larger and better developed thallus areas, but even well-developed thalli remain poorly delimited patches and never spread extensively into a continuous crust. Among the species discussed here, only B. ryanii forms a distinctly areolate thallus (Figs. 29, 30). The areoles form independently on a black, arachnoid hypothallus, and individual areoles regularly become subsquamulose along the margin. All other species have a rimose to rimose-areolate thallus, that is, the surface is initially fissured and only secondarily breaks into areoles. In exceptionally thick thalli of B. christophii and B. pullata these secondary areoles can even become subsquamulose. Generally, thalli of B. christophii, B. pullata, and B. tergua are very similar and easily confused (Figs. 1, 2, 8, 9, 35). Buellia tergua is lighter brown than the deep brown thalli of the other two species. However, only a few specimens of B. tergua are currently known, and it is therefore not possible to assess whether the leather-brown colour represents a consistent trait. The three species may therefore not be reliably distinguished by their thalli alone. Spore ontogeny and ultrastructure In crustose Physciaceae, spore structure has attracted considerable attention (Mayrhofer 1982), and especially in the genus Rinodina, several diagnostic types have been recognized because of conspicuous wall and septum thickenings (Mayrhofer and Poelt 1979; Mayrhofer 1984; Giralt 2001). In Buellia, Scheidegger (1993) first discovered thickened spore septa in a few species, and Kalb (1986) segregated the genus Hafellia for species with lateral wall thickenings. The wall thickenings of spores in the Physciaceae are, however, by no means static but the result of a dynamic spore ontogeny, which is still not completely understood. All species treated here have very similar ascospores, and measurements overlap considerably. Nevertheless, diagnostic differences can be found in all species by carefully comparing their spore ontogeny (Fig. 70). For convenience, four stages may be distinguished and some characters are only present during a brief stage: immature spores are hyaline, premature spores olive, mature spores brown, and overmature spores deep brown. Immature and premature spores generally have no visible ornamentation, although in some species ornamentation may begin to develop with the onset of pigmentation. Other species barely show any ornamentation during their ontogeny, but overmature spores often appear ornamented. This may sometimes be an artefact ascribed to degeneration of the spores. Wall or septum thickenings are usually best developed in premature and mature spores and may disappear again in aging spores. In Rinodina, Giralt and Mayrhofer (1994, 1995) distinguish sporesthat form spore septa before apical wall thickenings (A-ontogeny) from spores with late septum formation, where apical thickenings are present before the development of a septum (B-ontogeny). Giralt (2001) admits that some species have an intermediate ontogeny. In spores typically found in Buellia, apical thickenings generally do not develop (Fig. 70). Early septum formation may better be distinguished from late septum formation based on spore wall pigmentation. In spores with A-ontogeny, pigmentation develops after a septum can be distinguished in the immature spore. Late septum formation of the B-ontogeny develops in premature spores after the walls have become distinctly pigmented. Buellia tergua and B. sequax can tentatively be assigned to the A-ontogeny, whereas spores of the other species appear to follow the B-ontogeny. This distinction is not easy to assess. Even spores with pigmented walls do not always have septa, and sometimes these spores can also be observed along hyaline spores with septa. A- and B-ontogeny thus does not appear to be distinctly different in Buellia s. 1. Among the specimens examined here, spores of B. prospersa are most easily recognized (Fig. 70). They are the largest spores and have a conspicuous septum thickening most distinctly observed in premature spores. This septum is persistent in mature spores, but eventually reduced in over© 2004 NRC Canada Bungartz at al. mature spores. Because of the thickening, these spores have been referred to the Physconia-type (Scheidegger 1993). In some spores, the septum remains conspicuously thickened during almost the entire ontogeny. Apical thickenings are 555 never present, and the spores could thus also be referred to as the Orcularia-type. Mayrhofer et al. (1999) described a similarly persistent septum thickening in Amandinea insperata (Nyl.) H. Mayrhofer & Ropin. Differences be© 2004 NRC Canada Can. J. Bot. Vol. 82, 2004 556 Fig. 70. Ascospore ontogeny of Buellia species from the Sonoran Desert with inconspicuous, saxicolous thalli. Immature ascospores are hyaline, mostly nonseptate (B-ontogeny), but some early septate (A-ontogeny). An olive pigmentation begins to develop at the premature stage, some spores also showing Bue/lia christophii r··",\\\ ＯＢ ｾ ＺＧ Ｎ fｴＺｾ ' (I .n t:..\ ''II \....:..=j Bue//ia ryanii ｮl \) ｾ ｪｾ e8 8 Bue/lia pu/lata ,nII 8 , 888 j! ;f t\ \\ /j ＿ｾ］ＧＮ｜ Bue/lia sequax f ｩ＼］ｾ { i\ """ij 'J hyaline olive brown immature premature mature deep brown overmature tween the two types seem not very distinct. especially if apical wall thickenings cannot be observed in the Physconiatype. A faint and inconspicuous thickening can also be seen in the spore ontogeny of Buellia pullata. where it may easily be overlooked. These spores could thus be assigned alternatively to the Buellia (= Beltraminea)- as well as the Physconia-type (Fig. 70). A similar situation has been discussed for B. dispersa. where the septum thickening is also restricted to a brief phase of the ontogeny (Bungartz et al. 2002). In spores of both B. prospersa and B. pullata a microrugulate ornamentation forms early. It is barely visible in premature spores. but becomes distinct in mature spores. Spores of B. christophii are very distinct in the premature stage when they are very broad to almost globose and have a conspicuously, evenly thickened spore wall (Fig. 5). The broad spore wall of these premature spores is also distinct in the TEM (Fig. 6). Mature spores of the species become less globose and are more ellipsoid to oblong (Fig. 70). With spore maturity the conspicuously thickened wall disappears, and spore ornamentation cannot be distinguished in any of the stages. Spores of B. sequax are often difficult to distinguish from B. pullata. B. tergua, and B. ryan;;. Premature and mature spores of B. sequax are distinctly narrowly oblong. Their shape is very characteristic but transient. Scheidegger (1993, p. 356) suggests that .....The type of B. abstracta combines a chasmolithic thallus and narrow spores and was thus previously thought to be a good species." He continues to say that intermediate forms with B. sequax can be found and that B. sequax therefore constitutes a polymorphic species. We have examined type material of both taxa (see the discussion on thallus variation). The spores are not very variable but basically follow the same ontogeny, with a premature stage of characteristically narrow spores. This stage. however, is easily overlooked if the apothecium contains a majority of ellipsoid spores at a later stage of the ontogeny. Spore ornamentation of B. sequax develops early, but it is inconspicuous and can usually only be distinguished in overmature spores. Nordin (1997, 2000) emphasized spore ultrastructure of Buellia species with pluriseptate ascospores. In the species studied here, TEM yields little additional information. It is noteworthy. however. that in B. sequax, spores with a fractured as well as a smooth perispore have been observed in the TEM (Figs. 26. 27). This emphasizes the dynamic development of the spore. Immature and premature spores have a smooth perispore (Figs. 26. 70). which becomes fissured and eventually fractured with maturity (Figs. 27. 70). Spores of B. ryanii and B. tergua usually become more or less constricted along the septum (Fig. 70). This constriction is not obvious in young spores. but it is usually pronounced in overmature spores. Buellia ryan;; has the smallest spores. with no distinct ornamentation and thin spore walls. In contrast. spores of B. tergua have thicker walls, which early develop an ornamentation. The ornamentation is best seen in mature and overmature spores. Taxonomy Some of the species described here have bacilliform conidia (Figs. 7, 24, 34, 40, 72) and others filiform conidia © 2004 NRC Canada 557 Bungartz et al. Fig. 71. Distribution of Buellia species with inconspicuous saxicolous thalli'in southwestern North America. In the Sonoran Desert Region, Buellia christophii, Beullia ryanii, Beullia prospersa, and Beullia tergua are currently only known from coastal localities. Buellia sequax is widely distributed throughout the region. 1120 06' W ____ J • I iLBuellia christophii - Buellia prospersa • • , I i Buellia pullata Buellia ryanii • Buel/ia '---_ _tergua Ｍ Ｂｾ Buellia sequax o ｟ ＢＧ ｟ ＾ＡＮ Ｚｗ ｾＢ Ｇ｟ Ｎ｟ｩＺｬ ｾ lO9"l9'W 500 == Subdivisions ofthe Sonoran Desert Floristic Province (Shreve & Wiggins 1964): . . Arizona Upland Lower Colorado Region Iml Central GulfCoast Magdalena Region 11IIII ｳｬｩ ｾｆＺ ｲ ofSonora) I!mmmmi Vizcaino © 2004 NRC Canada Can. J. Bot. Vol. 82, 2004 558 (Figs. 13, 19,72). Otherwise, all species are very similar. Indeed, this similarity is the main reason why the species were previously generally identified as B. punctata s. 1. We can currently not confirm that any saxicolous specimens from the Sonoran Desert Region belong to B. punctata s. str. According to Mayrhofer and Moberg (2002, p. 9), "saxicolous and muscicolous material usually placed under this name is much in need of taxonomic revision". Scheidegger (1993, p. 343) mentions that "Saxicolous material, usually placed under this name, is possibly not homogeneous and is not yet completely understood by the author...... According to Scheidegger's (1993) description, A. punctata s. str. has spores without a septum thickening, no spore ornamentation, and short fUiform conidia. Sheard and May (1997) noticed that the North American material, which they assigned to A. puncata, has ornamented spores, and a septum thickening can be observed at least during some short period of spore development. They also report that they have only seen a single specimen with short filiform conidia (averaging <15 mm) and all other specimens have longer conidia. Sheard and May (1997) did not examine saxicolous material from the Southwest, and it is possible that the specimens identified as A. punctata do not strictly belong to that taxon (l.W. Sheard, personal communication). From the taxa treated here, the newly described species are all distinct from B. punctata s. str. because of their short, bacilliform conidia (Figs. 7, 34, 40, 72). In addition, B. ryanii and B. tergua are also characterized by having the apothecial pigment cinereorufa-green, which is absent from B. punctata s. str. The presence of broad, almost globose spores and a thick exciple also separates B. christophii from B. punctata s. str., even if conidia are not found in all of the specimens. The genus Buellia s. 1. currently represents an amalgam of not necessarily closely related species, and various attempts have been made to subdivide this large and heterogeneous group into smaller, more strictly defined genera. In the present publication, we have examined a group of very similar species, some of which could have been treated within the genus Amandinea, solely justified by the filiform conidia. Amandinea, however, does not currently constitute a more precisely defmed taxonomic group than Buellia. When Scheidegger (1993) described Amandinea, he provided the first valid description for a genus previously suggested, but not validly published, by Choisy (1950). Choisy (1950) distinguished Amandinea from Buellia because of the "pycnoconidies aciculaires", that is, the acicular conidia. Scheidegger (1993) emphasized the differences in conidial length and conidiophore structure as the only diagnostic characters of the new genus Amandinea. Choisy (1950) transferred Buellia coniops (Wahlenb.) Ach., B. myriocarpa (DC. ex I. M. Lamb. & DC.) De Not., and B. stigmatea Korb. into the new genus. Scheidegger (1993) selected Amandinea coniops as the type of Amandinea, and also included Rinodina lecideina and Buellia punctata. Subsequently, the genus became more widely accepted. Matzer et al. (1994) transferred Rinodina petermannii (Hue) Darb. into Amandinea, emphasizing its filiform conidia. The species investigated by Scheidegger (1993), as well as Amandinea petermannii, are all saxicolous. Sheard and May (1997) transfered several corticolous Buellia and Rinodina Fig. 72. Range of conidial length in various saxicolous species of Buellia (modified from Scheidegger 1993). The bars indicate the range of measurements for each of the species. Buellia pullata overall has not only the longest conidia but also the largest variation in measurements. For Beullia prospersa, the bold bar indicates the range of measurements given by Scheidegger (1993), and the narrow bar indicates the variation observed here. Measurements overlap considerably, and the asymptotic graph shows that the distinction of bacilliform versus filiform is arbitrary and does not represent a discrete character. B.paDllIa B. prospena B.coniops B.punctata B. subdlsciformis B. lesserata B.jugorum B. longispora B. sardinienstJ B. saxorum B.dispena B. almeriemis B. aethalea B. spuria B. concinna B. ocel/ata B. bama - asymptotic graph of the measurements • .- B. leptocline B. terglUl B. stella/oto B. vilis B. ry"m; B. elrrlstoplrll ｸ｡ｬ ｦｾｳＮｂ 10 15 2'0 25 30 35 40 45 conidial length (11m) species from North America into Amandinea. These North American species all have fUiform conidia. Most recently, Mayrhofer and Sheard (2002) transferred Rinodina cacuminum (Th. Fr.) Malme into Amandinea because of its fUiform conidia, also citing "preliminary molecular results" (p. 440), which do not support the inclusion of that species in Rinodina. Marbach (2000) argued that Sheard and May (1997) provided an emended genus concept for Amandinea, allowing the inclusion of species where no conidia could be observed. Bungartz and Nash (2004b) questioned this approach. They demonstrated that Buellia turgescens Nyl. ex Tuck., a species that Marbach (2000) included in Amandinea without having studied the pycnidia, is synonymous with B. badia (Fr.) A. Massal. Bungartz and Nash (2004b) showed that Buellia badia is a species with bacilliform conidia, which therefore must not be included in Amandinea. There is currently little agreement about the characters characterizing the genus Amandinea. Amandinea may also be characterized by the scarcity of lichen secondary metabolites in the thallus, as suggested by Christoph Scheidegger (personal communication). Scheidegger (1993) did not detect any secondary metabolites in the species he included in the genus. Also, Sheard and May (1997) did not document any secondary metabolites in the North American species. Matzer et al. (1994), however, report norstictic acid and ergosterol peroxide from A. petermannii. Even though Scheidegger (1993) did not detect secondary metabolites in A. lecideina (= B. prospersa), that species is clearly characterized by a pale UV+ yellow to orange thallus, a reaction © 2004 NRC Canada Bungartz et al. caused by various xanthones. Several species, which Marbach (2000) included in Amandinea, also have characteristic secondary metabolites. In addition, Marbach (2000) mentioned that Amandinea species often have a pale hypothecium, a character more commonly associated with Rinodina rather than with Buellia. The species treated by Scheidegger (1993) (including the type), and most of the species treated by Sheard and May (1997), however, have a dark hypothecium. All species treated here also have a dark brown hypothecium. In his treatment of corticolous Buellia s. I., Marbach (2000) emphasized branching patterns of the paraphyses and the pigmentation of their apical cells as taxonomically important. According to Marbach (2000, p. 52) large, globose apical cells are characteristic for Amandinea. "Tafel 1/2: Paraphysenenden" (pp. 36 and 37, Marbach 2000), however, documents only very subtle, if any, differences in the paraphyses of Buellia s. I. No distinct differences that would justify a clear segregation of the genus Amandinea are displayed in this chart. Marbach (2000) did not discuss the different apothecial pigment types in any detail, even though these differences are clearly taxonomically important (e. g., in B. ryanii and B. tergua; see also Bungartz and Nash 2004a). In addition to the paraphysal characters, Marbach (2000, p. 52) emphasized that spores without wall thickenings are typical for Amandinea. Nevertheless, he included several species within the genus that have distinctly thickened septa. As previously discussed, wall thickenings vary considerably during spore ontogeny, and the spores of some species (like B. pullata) are difficult to assign to a particular type. Conspicuously thickened spore septa are characteristic for several species transferred from Rinodina into Amandinea. When Scheidegger (1993) validated Choisy's genus description of Amandinea, he included two species with distinctly thickened septa, including the type A. coniops. By transferring R. insperata into Amandinea, Mayrhofer et al. (1999) also explicitly included a species with distinctly thickened spore septa. However, taxa transferred from Buellia (e.g., Buellia punctata) often lack a distinct septum thickening. In addition to the filiform conidia, Giralt et al. (2000, p. 521) provide a list of the following characters typical for Amandinea: "apothecia lecanorine or lecideine, ascospores brown, one-septate, Buellia- or Physconia-type, often with rugulate ornamentation; and thallus with norstictic acid or more frequently without secondary lichen compounds....". All these characters are regularly found in Buellia and are by no means exclusive to Amandinea. A lecanorine apothecium is generally more common in Rinodina, but some species in Buellia also show the tendency to form a thalline exciple. In addition, immersed lecideine apothecia often emerge from the thallus, with some thallus material remaining attached to the exciple (see B. ryanii and B. tergua). The ascus of Amandinea is generally described as the Bacidia-type (Marbach 2000, Giralt 2001), and it is thus not different from Buellia s. str. (Rambold et al. 1994). Sl1lChting et al. (2004) recently transferred several arctic species from Buellia into Amandinea, arguing that in the absence of molecular data this transfer is only justified because of the filiform conidia of those species. The filiform conidia borne on type III conidiophores in a 559 Roccella-type pycnidium (sensu Vobis 1980) therefore represent the only reliable diagnostic character consistently used to segregate Amandinea from Buellia. Within the Physciaceae, conidial length has also been used to distinguish Physcia from Phaeophyscia (Moberg 1977), Physcia from Hyperphyscia (Choisy 1950, Hafellner et al. 1979), and to characterize the genus Mobergia (Mayrhofer et al. 1992). None of these genera, however, are distinguished solely by the length of their conidia, and the foliose genera are also well circumscribed by a range of other characters. The shape of conidia (bacilliform vs. filiform) is a result of conidial length, and it does not represent a general difference in conidial structure. Conidial length may vary considerably, even within the same pycnidium of a single specimen. In fact, B. pullata has predominantly 20-25 Jlm long filiform conidia, but overall, the conidia measured from a single pycnidium of that species vary from 6 to 43 Jlm in length (Fig. 72). The shortest conidia are thus bacilliform rather than filiform. In B. prospersa (= A. lecideina), the range of our measurements (7-32 Jlm, Fig. 72) considerably exceeds the variation observed by Scheidegger (1993), who measured conidia from 15-30 Jlm long. Scheidegger (1993, Fig. 5) plotted conidial length of the species in his treatment to demonstrate the differences between Buellia and Amandinea. The shortest conidia were found in Buellia vilis (2.54 Jlrn). Buellia subdisciformis was the species with the longest conidia (9-14 Jlm) still treated within Buellia, but conidia of A. punctata were only slightly longer (up to 15 Jlm). Both A. lecideina and A. coniops had the longest conidia (15-30 Jlm). Plotting our measurements of B. pullata and B. prospersa (= A. lecideina) to this chart results in considerable overlap, and conidial length can therefore no longer be regarded as a discrete character. An asymptotic graph more accurately describes the situation (Fig. 72). In addition to the conidial length, differences in branching patterns of the hyphae bearing conidia can also be observed. Conidiophore structure is, however, closely related to the length of the conidia and cannot be treated as an independent character. Long, filiform conidia are generally borne on shorter and less extensively branched conidiophores. This is a necessary consequence of ejecting conidia from the pycnidium. Long filiform conidia would become entangled in densely branched conidiophores. Densely branched conidiophores can, however, produce larger quantities of short, bacilliform conidia and can thus be found in species that have only short conidia. In lichen taxonomy, conidia alone have not generally been used to distinguish taxa at the generic level. In some groups, conidia of very different lengths can even be found on a single thallus. The genus Micarea regularly has both microconidia and macroconidia formed in different pycnidia on the same thallus. In other groups, conidia of different length occur in closely related species of the same genus. For example, Lecanora himalayae Poelt (from Nepal) and two very similar species, Lecanora chondroderma Zahlbr. (from China) and Lecanora beamanii B. D. Ryan (from high elevations in southern Mexico), have straight, 6-8 Jlm long conidia. In contrast, Lecanora maxima Lynge has curved conidia that are 11-17(-20) Jlm long. Nevertheless, all taxa belong to the section Dactylon Poelt that presumably forms a natural group isolated from other species of Lecanora © 2004 NRC Canada 560 (Ryan 1989). Other genera, for example, Haematomma (Rogers and Hafellner 1988) or Aspicilia (Clauzade and Roux 1984), also include some species with bacilliform and others with filiform conidia. In the Physciaceae, the validity of using filiform versus bacilliform conidia as the sole character to justify Amandinea has recently been questioned. Because of their miform conidia, some terricolous species of the BueLlia epigaea group would have to be formally transferred into Amandina, even though they do not appear to be closely related to other species with miform conidia (H. Mayrhofer and U. Grube, personal communication; Grube and Amp 2001; Trinkaus et al. 2001). Preliminary molecular studies also do not confirm Amandinea as a monophylletic group (H. Mayrhofer personal communication; Grube and Amp 2001; Wedin et al. 2002). In some lichen species, pycnidia are very rare and are thus rarely found in all of the specimens examined. If pycnidia are rare, it may sometimes be difficult to assess if these conidia belong to a particular lichen specimen or instead to a lichenicolous fungus growing inside this specimen. A character that is frequently absent and difficult to assess does not seem to be particularly suitable as the sole diagnostic feature to circumscribe a genus. A monophylletic genus Amandinea, with A. coniops as the type, will most likely not include a large portion of species currently included by various authors. In the Sonoran Desert, the B. mamillana group (F. Bungartz, unpublished data) includes species with xanthones that all have long bacilliform to fusiform conidia. Within Buellia s. 1., that species group is isolated, but not closely related to species commonly treated within Amandinea. Because of its yellowish thallus containing xanthones, Buellia prospersa (= A. lecideina) shows some affinities with the B. mamillana group. Buellia pullata, on the other hand, is fairly similar to the type species A. coniops, even though the spore septa are hardly thickened during their ontogeny. In summary, the taxonomy of Amandinea is not well resolved. Segregating the species treated here into separate genera only on the basis of conidial length would result in a constrained and arbitrary taxonomy. Nimis (1998) discussed three main criteria that justify the recognition of a genus in lichens: (1) monophylly, (2) phylogenetic analysis of its taxa, and (3) several independent characters that circumscribe the new genus. The genus concept of Amandinea currently matches none of these criteria. "If only one character is involved, this is indeed the weakest possible evidence. It could easily be in conflict with the next character to be discovered..." (Nimis 1998, p. 432). This is exactly the situation in the genus Amandinea. Including species solely on the basis of conidial length does not help to resolve the taxonomy of Rinodina or Buellia, both of which remain large, artificial, and heterogeneous genera. Several recent treatments (Grube and Arup 2001; Wedin et a1. 2002; Helms et al. 2003) have confirmed two general clades in the Physciaceae: (1) the Physcia (or Rinodina) clade, characterized by a Lecanora-type ascus, a usually hyaline hypothecium, and spores that regularly have wall thickenings (e.g., Physcia, Heterodermia, Anaptychia, Mobergia, Tornabea, Rinodina, etc.) and (2) the Buellia Can. J. Bot. Vol. 82, 2004 clade with Bacidia-(or Biatora)-type ascus, usually a dark apothecium, and spores that less frequently have wall thickenings (e.g., BueLlia, Dimelaena, Dermatiscum, etc.). The "Caliciaceae" apparently also belong to this second clade (Wedin et al. 2002). The resolution within the clades is currently not sufficient to justify further segregation of the crustose genera Rinodina s. 1. and Buellia s. 1. Until a more resolved phylogeny is achieved, using both classical and molecular data, we advocate including the species currently treated as Amandinea within the genus Buellia s. 1. Excluding species with filiform conidia from Buellia does not resolve the nomenclatural problems in this rather large and heterogeneous genus. According to the code of Botanical Nomenclature (Greuter et al. 2000), B. disciformis (Fr.) Mudd. is currently the listed type of the conserved genus Buellia. Unfortunately, the genus HaJellia Kalb., H. Mayrhofer & Scheid. is also based on the same type. To avoid this conflict, Moberg et al. (1999) suggested conserving Buellia with Buellia aethalea (Ach.) Th. Fr. as a new type. If this proposal is rejected, most species now treated within Buellia s.l. would then have to be accommodated in a new genus. This may include some of the species currently treated within Amandinea. The name Buellia s. str. would then refer to species generally treated as HaJellia, a genus that would become obsolete. None of the species in Amandinea is closely related to that group. If, however, the proposal by Moberg et al. (1999) is accepted, some of the species currently transferred into Amandinea may eventually tum out to be more closely related to the B. aethalea group and may still be better accommodated within BueLlia. Acknowledgements We are obliged to Dr. Laurence A. Garvie, Department of Geology, Arizona State University (ASU), for access to a slow-speed diamond stone saw. Dr. Donald M. Burt, Department of Geology (ASU), identified the rock substrates of the specimens sectioned. The chemistry of all specimens was analyzed with TLC at ASU. Specimens from the private herbarium of Dr. Christoph Scheidegger were also examined with TLC by Martin Frei Swiss Federal Institute for Forest, Snow and Landscape Research (Birmensdort). Additionally, selected specimens were analyzed with HPLC by Dr. Jack A. Elix from the Australian National University in Canberra. Dr. Ulrik Ｌｧｮｩｴｨｾｓ University of Copenhagen, Denmark, and Julia Blaha and Ulrike Grube, University of Graz, Austria (GZU), contributed valuable information to the discussion of the taxonomy of Amandinea. Scott Bates (ASU) and Dr. John Sheard, University of Saskatchewan, Canada, kindly reviewed first drafts of the manuscript. Dr. Helmut Mayrhofer (GZU) and an anonymous reviewer provided valuable input to the final version of this publication. Dr. Christian Printzen, Senckenberg Institut, Germany, was very helpful correcting the Latin diagnoses. Dr. Donald Pinkava, ASU, provided much support on taxonomic questions regarding the application of the ICBN. We would like to thank all herbaria, which have made specimens available for study, especially Julia Blaha (GZU), who, on short notice, supplied several specimens of B. prospersa, including the type of A. lecideina. On very short notice, Dr. Orvo © 2004 NRC Canada Bungartz at al. Vitikainen, Helsinki (H) provided the Nylander specimen of B. prospersa used for lectotypefication. This study was supported by a Sigma Xi Grant-in-Aid of Research, sponsored by the local Sigma Xi chapter at ASU, and two National Science Foundation grants (DEB-0103738, DEB-9701111). References Bellemere, A. 1994. Asci and ascospores in ascomycete systematics. In Ascomycete systematics: problems and perspectives in the nineties. NATO ASI Series, Series A: Life Sci. Vol. 269. Edited by D.L. Hawksworth. International Mycological Institute, Surrey, UK. pp. 111-126. Bungartz, E, and Nash, T. H., ill. 2004a. Buellia subalbula (Nyl.) MUll. 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