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Morphology and anatomy of chasmolithic versus epilithic growth: A

taxonomic revision of inconspicuous saxicolous Buellia species from the
Sonoran Desert Region generally ascribed t...

Article  in  Canadian Journal of Botany · April 2004

DOI: 10.1139/b04-028

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 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 pene-
tration. 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 jus-
qu'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 es-
peces 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 Amandi-
nea. 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 Amandi-
nea. 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 has therefore often focused on species with well-developed

epilithic thalli, whereas species that show some degree of
The taxonomy of crustose lichen species with thin and in- endolithic growth have been neglected.
conspicuous thalli is generally poorly resolved. These spe-
The present study focuses on a group of species that in
cies are often overlooked, rarely collected, and their thallus
characteristics have been disregarded as highly variable and North America have generally been treated as Buellia punc-
poorly differentiated. The attention of taxonomic treatments
tata (Hoffm.) A. Massa!. or more recently as Amandinea
punctata (Hoffm.) Coppins & Scheid. In particular, the saxi-
colous material commonly identified as this species is
Received 3 September 2003. Published on the NRC Research poorly understood (Mayrhofer and Moberg 2002). Thallus
Press Web site at http://canjbot.nrc.ca on 25 May 2004. variation has largely been ignored. For example, in his treat-
ment of the North American species of Buellia, Imshaug
F. Bungartz,l T.n. Nash ITI, and B.D. Ryan. School of (1951) states" ... Buellia pullata is, as Tuckerman (1888) in-
Life Sciences, Arizona State University, P.O. Box 87 4601, dicated, a saxicolous form of B. punctata with a well devel-
Tempe, AZ 85 287-4601, USA.
oped thallus. Buellia saxicola de Lesd., on the other hand, is
lCorresponding author (e-mail: frank.bungartz@asu.edu). a saxicolous form of B. punctata with a scant to obsolete

Can. J. Bot. 82: 540-562 (2004) doi: 10.1l39/B04-028 © 2004 NRC Canada
Bungartz et al. 541

thallus ...". The material examined in our study indeed infiltration, this protocol was modified according to
shows some variation in thallus morphology. Several distinct Bungartz and Nash (2004a).
species can nevertheless be distinguished based on thallus Representative specimens were cut with a low-speed dia-
morphology as well as spore ontogeny, apothecial pigments, mond saw and the sections treated according to a protocol
conidial length, and secondary chemistry. Buellia pullata, outlined by Bungartz et al. (2004). To visualize hyphae pen-
for example, is not, as Imshaug (1951) suggested, a mere etrating dark stone substrates, the protocol was modified and
form of B. punctata, but a distinct species. Likewise, it can specimens were stained with lactophenol cotton blue (LCB)
be demonstrated that Beullia saxicola is a synonym of Beul- instead of periodic acid - Schiff's reagent (PAS). Sections
fia sequax (Ny1.) Zahlbr., a taxon not previously docu- were studied with both light microscopy and SEM as dis-
mented from North America, but recognized as a distinct cussed in Bungartz et al. (2004). Some of the sections were
species by Scheidegger (1993). critically point dried and gold coated. All sections were
Six species previously treated in Imshaug's "B. punctata" studied in a JEOL JSM-840A SEM at accelerating voltages
group are distinguished among the Sonoran material. Thallus between 7 and 12 kV in secondary and backscattered emis-
variation in this group is clearly much less pronounced than sion mode.
previously assumed. None of the species presented belongs Specimens were spot tested and routinely examined with
to B. punctata s. str., which apparently does not occur on standardized thin-layer chromatography (TLC) (Culberson
rock substrates in the Sonoran Desert Region. Contrary to and Kristinsson 1970; Culberson and Johnson 1982; White
recent practice, the genus Amandinea is not accepted. The and James 1985; Orange et al. 2001). To differentiate
taxonomy of that genus is currently not clearly resolved, and various xanthones, selected specimens were additionally ex-
all species are therefore treated as members of Buellia s. 1. amined with standardized high-performance liquid chroma-
tography (HPLC; Feige et al. 1993). TLC plates were
Materials and methods interpreted with the computer program WINTABOLITES
(Mietzsch et a1. 1994) and scanned for permanent record
All specimens were examined with light microscopy using (Egan 2001). Spores measurements are given according to
hand- and cryo-sections. Both conventional bright field mi- Nordin (2000). Pigment names follow Meyer and Printzen
croscopy (BF) and differential interference contrast (DIC) (2000).
were used. Selected specimens were also studied with scan- Figure plates were assembled and processed in Adobe
ning electron microscopy (SEM) and transmission electron Photoshop 7.0, and selective contrast adjustments were made
microscopy (TEM) according to a protocol described in de- to emphasize details such as the apothecia in SEM micro-
tail by Bungartz et al. (2002). To improve dehydration and graphs.

Results

Key to the species

lao Thallus chasmolithic, discontinuous, usually of dispersed granules, growing among minerals of the substrate. . . . . . 2
lb. Thallus epilithic, forming a thin, more or less continuous crust on the substrate surface. . . . . . . . . . . . . . . 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 ~m x
6-9 ~m, conidia 4-7 ~m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 ~m x 4-8 ~m, conidia 2-5 ~m . . . . B. ryanii
Sa. Premature spores broad, almost globose, evenly thick walled, septum not thickened; exciple thick, outer carbonized zone
usually >20 ~m 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 ~m 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 persis-
tent 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 mi-
crograph 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, frac-
tured perispore, (2) intermediate layer, (3) proper wall, and (4) endospore (Nash 11426).

Species descriptions Asci 8-spored, clavate, Bacidia-type. Ascospores (Figs. 5,

6, 70) broadly ellipsoid to almost globose, never con-
Buellia christophii Bungartz sp. nov. stricted, 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),
DIAGNOSIS: Thallus saxicolus, crustaceus, rimosus vel rimoso- one-septate; proper septum narrow, not thickening during
areolatus, tenuis vel crassus, fulvus, sine hypothallo. Apo- spore ontogeny (Fig. 70), with septal pore canal, simple pore
thecia sessilia, lecideina, marginibus propriis crassis. and undifferentiated pore plug; no ornamentation visible (in
Excipulum crassum, fulvum, sine pigmento aeruginoso, DIC); spore wall (Fig. 6) differentiated into smooth, thin
carbonaceum. Asci 8-spori. Sporae unisaeptae, late ellip- perispore (0.10-0.20 Jlm), narrow intermediate layer
soideae vel globosae, 9-15 Jlm x 6-10 Jlm. Pycnidia (<10 Jlm), thick proper spore wall (0.50-1.50 Jlm) and mod-
globosa. Conidia bacilliformes, 2-4 Jlm x 1-1.5 Jlm. Thallus erately thickened endospore (0.20-0.40 Jlm).
acida norstictica et connorstictica continens vel has sub- Pycnidia (Fig. 7) rare to common, globose, unilocular, at
stantias deficiens. maturity almost entirely occupied by densely branched co-
nidiophores; conidiogenous cells mostly terminal, rarely also
TYPE: MEXICO. Baja California: lighthouse area near Lagu- intercalary (cf. conidiophore-type V sensu Vobis 1980); pyc-
na Manuela; 28°15'OO*N, 114°07'OO"W; altitude: 100 m; on nidial ontogeny similar to the Umbilicaria-type (sensu Vobis
basalt; 21 February 1993; Nash 33979 (ASU - holotype 1980, Vobis and Hawksworth 1981); conidia (Fig. 7) simple,
designated here). bacilliform, 2.0-4.5 Jlm x 1.0-1.5 Jlm (n = 69) (Fig. 72).
TAXONOMIC NOTE: This species is named in honour of CHEMISTRY: No substances found or with norstictic and con-
Dr. Christoph Scheidegger, Birmensdorf, Switzerland. norstictic acid (low concentrations only detected by TLC or
Figs. 1-7 and 55~3. HPLC). All spot tests negative (very rarely forming orange
needle-shaped crystals with K if observed with the com-
Thallus (Figs. 1, 2, 55~3) crustose, usually rimose to pound scope). UV-. Thallus without amyloid reaction, only
rimose-areolate, thin and more or less continuous but dis- the apothecia amyloid in Lugol's.
tinctly epilithic, never chasmolithic, rarely becoming thick-
ened and distinctly rimose-areolate to subsquamulose, not SUBSTRATE AND ECOLOGY: All specimens examined are from
delimited by a hypothallus; surface matt to shiny, usually low elevations, growing on siliceous mineral-poor coastal
deep brown, rarely pale brown, usually smooth, but fre- rock. The holotype is erroneously labeled as growing "on
quently roughened in areas where the surface appears basalt". The substrate is, however, a very fine grained ande-
eroded, epruinose, phenocorticate. site, with intermediate to silicic acidity, poor in bases.
Apothecia soon sessile, lecideine; proper margin conspicu- DISTRIBUTION: The species is currently known only from
ously thickened, usually persistent, rarely excluded with age; coastal southern California, USA, Baja California, and Baja
disk black, epruinose, plane, rarely becoming slightly convex California Sur, Mexico (Fig. 71).
with age; exciple of aethalea-type (Figs. 3, 4) sensu Schei-
degger (1993), that is, inner excipular hyphae narrow, hyaline, NOTES: The species is superficially similar to B. pullata
prosoplectenchymatous (textura oblita), similar in structure and (Figs. 8, 9, 55~3), and differences have been discussed in
orientation to the paraphyses, often more or less reduced and the note on that species. Table 1 gives an overview of the di-
transient with the dull reddish brown hypothecium agnostic characters available to distinguish all species dis-
(leptoclinoides-brown, textura inricata); outer excipular hyphae cussed here.
parallel (textura oblita), cells moderately swollen and usually
strongly carbonized with various amounts of brown pigments MEXICO. Baja California: Ryan 21513,
MATERIAL EXAMINED:
(cf. elachista-brown), pigmentation continuous with the epihy- Marsh 6198, Nash 34569 (ASU). Baja California Sur: Nash
menium; hymenium hyaline, not inspersed; paraphyses simple 29800 (ASU). USA. California. Los Angeles Co.: Wetmore
to moderately branched, apically swollen, with a brown pig- 73435 (MIN); Hasse Exs. 205 (CAS); Weber L-42660, L-
ment cap (cf. elachista-brown). 42178, L-42662, L-42163 (COLO); Nash 32093, 32100,

© 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 micro-
graph 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.: Britton & Kemp (79), determined by B. Fink] has been an-
Tucker 35170 (SBBG). Monterey Co.: Weber 8242, 8237 notated by Fink as the type of "Buellia substigmata Fink".
(COLO). San Luis Obispo Co.: Tucker 36478 (SBBG); Nash This name has never been published by Fink and the speci-
36982 (ASU). Santa Barbara Co.: Printzen 49, 76 (hb. men is not the type of Buellia substigmatea Mull. Arg. [Pro-
Printzen); Weber L-80178 (COLO); Nash 32386, 41174, ceed. Roy. Soc. Edinburgh 11: 465 (1882)]. The MICH
41181, Ryan 31359. Crayton s. n. (ASU). specimen has correctly been annotated by Imshaug as
Buellia prospersa (Nyl.) Riddle.
Buellia prospersa is the valid name for Rinodina
Buellia prospersa (Nyl.) Riddle (= Amandinea) lecideina, which is, however, not identical
Brook!. Bot Gardens Memoirs 1: 114 (1918) with Buellia lecidina Stein. ex Cohn [Kryptog. Flora von
Lecidea prospersa Nyl., Flora 63: 127 (1880). Type: Vir- Schlesien 2(2) (1879)]. Even though the spelling of the spe-
gin Islands: St. Thomas [original label data. S. Thomae cies epithets "lecideina" and "lecidina" is almost identical,
Antillarum], 1878, Dr. Forel. (H-Nyl 9312! - lectotype se- Buellia lecidina Stein. ex Cohn is a synonym of Rinodina
lected here). occulta Korb.
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
pasture walls [original label data: Eire/Ireland: Co. Ciarrai / Thallus (Figs. 15, 16) crustose, rimose, moderately thin
Kerry, Corea Dhuibhne / Dingle-peninsula, Umgebung des and more or less continuous but distinctly developed and not
Weilers Ballyoughteragh N BailIe an Fheirtearaigh / Bally- chasmolithic, not delimited by a distinct hypothallus or with
ferriter, an Weidenmauem], August 1978, Poelt s. n. (GZU! - a black outline around the thallus; surface matt to more or
holotype). less shiny, usually pale ivory and smooth, rarely gray and
Amandinea lecideina (H. Mayrhofer & Poelt) Scheid. &
slightly roughened (probably as a result of surface damage),
H. Mayrhofer, in Scheidegger, Lichenologist 25: 342 (1993) epruinose, phenocorticate.
Buellia punctata f. crassior (Erichsen) Zahlbr., Cat. Lich. Apothecia soon sessile, lecideine; proper margin excluded
Univ. 8: 591 (1932). - Buellia punctata f. crassior with age; disk black, epruinose, plane, becoming convex
Erichsen, Das Hnke Untertraveufer: 151 (1932). Type: GER- with age; exciple of aethalea-type (Fig. 17) sensu Schei-
MANY, Schleswig-Holstein: Lubeck, Coast of Dummers- degger (1993), that is, inner excipular hyphae narrow,
dorf, boulders along the beach below the village Stulperbank hyaline, prosoplectenchymatous (textura oblita), similar in
[original label data: Deutschland, Schleswig-Holstein, Lu- structure and orientation to the paraphyses, often more or
beck, Dummersdorfer Ufer, Strandblocke unterhalb Stulper- • less reduced and transient with the dull reddish brown hypo-
bank], April 1928, Erichsen (HBG - holotype). thecium (leptoc1inoides-brown, textura inricata); outer
Buellia punctata f. litoralis (Erichsen) Zahlbr., Cat. Lich. excipular hyphae parallel (textura oblita), cells moderately
Univ. 7: 397 (1931). - Buellia myriocarpa var. litoralis swollen and usually strongly carbonized with various
Erichsen, Verh. Bot. Ver. Provinz Brandenburg 72: 48 amounts of a brown pigment (cf. elachista-brown); pigmen-
(1030). Type: GERMANY, Schleswig-Holstein: Eastcoast of tation continuous with the epihymenium; hymenium hyaline,
the Island Alsen, near Kettingholz, on boulders along the not inspersed, paraphyses simple to moderately branched,
beach in the supralittoral zone [original label data: Deutsch- apically swollen, with a brown pigment cap (cf. elachista-
land, Schleswig-Holstein, Insel Alsen, Ostktiste bei Ketting- brown).
holz, an StrandblOcken der supralitoralen Zone], July 1932, Asci 8-spored, clavate, Bacidia-type. Ascospores
Erichsen (HBG - holotype). (Figs. 18, 70) oblong to more or less ellipsoid, not con-
stricted, with obtuse ends, not curved, (9.0-)12.0-[13.5]-
TAXONOMIC NOTES: A specimen from MICH [rocks on a hill, 15.0(-16.0) Ilm x (5.0-)6.2-[7.3]-8.3(-9.5) Ilm (n = 91),
Christiansted, St. Croix, March 17-25, 1923, collected by one-septate; proper septum soon with a more or less persis-

© 2004 NRC Canada

Bungartz et al. 545

tent thickening, becoming reduced only in old spores (i.e., tirely occupied by the long conidia and lined with short,
with a distinct Physconia-type or Orcularia-type ontogeny); scarcely branched conidiophores; conidiogenous cells termi-
ornamentation microrugulate (conspicuous in DIe at an nal (conidiophore-type III sensu Vobis 1980); pycnidial on-
early stage of the ontogeny). togeny similar to the Roccella-type (sensu Vobis 1980,
Pycnidia rare, globose, unilocular; at maturity almost en- Vobis and Hawksworth 1981); conidia (Fig. 19) simple,

© 2004 NRC Canada

 ~
(J)

Table 1. Diagnostic differences of Buellia christophii, Buellia prospersa, Buellia pullata, Buellia ryanii, Buellia sequax, and Buellia tergua.

Thallus growth and UV reaction and

color Thallus morphology Conidia Exciple Spores chemistry
BueUia christophii
Epilithic; pale to Rimose to rimose-areolate, rarely Bacilliform (short) Thick. persistent, brown Broad oblong (almost UV negative, no
deep brown subsquamulose; without globose), equal wall xanthones, ± norstictic
hypothallus thickening, early acid
microrugulate
BueUia prospersa
Epilithic, rarely Rimose to rimose-areolate; Filiform (short to very long) Thin, persistent, brown Oblong-ellipsoid, persis- UV+ yellow to orange,
vaguely usually without a hypothallus, tent septum thickening, usually with
chasmolithic; rarely thallus surrounded by early microrugulate xanthones, ± norstictic
pale ivory black outline acid, ± atranorin
BueUia pullata
Epilithic; deep Rimose to rimose-areolate, (rarely Filiform (short to very long) Thin, persistent, brown Oblong-elIipsoid, brief UV negative, no sub-
brown to gray subsquamulose); without septum thickening, early stances detected
hypothallus microrugulate
BueUia ryanii
Epilithic; olive gray Rimose to rimcfse-areolate (rarely Bacilliform (short) Thin, excluded, Oblong-ellipsoid, no wall UV negative, no sub-
to brownish olive subsquamulose); with distinct aeruginose (N+ violet) thickening, late stances detected
arachnoid black hypothallus microrugulate
BueUia sequox
Chasmolithic, Granular to rimose; without Bacilliform (short) Thin, persistent to Narrow oblong, no wall UV negative, no sub-
inconspicious; hypothallus excluded, brown thickening, late stances or ±Dorstictic
pale brown to microrugulate acid, rarely with
grayish arthothelin
BueUia terguo
Epilithic; "Ieather- Rimose to rimose-areolate; with- Bacilliform (short) Thin to moderately thick. Oblong-ellipsoid, no wall UV negative, no sub-
brown" out hypothallus persistent, aeruginose thickening, late stances detected Q
(N+ violet) microrugulate ?
c...
@
IV
~
~
?=
~ <Xl
.!"
.~, I\:l

§
go
Bungartz et al. 547

long bacilliform to filiform, 7.0-32.0 ~m x <1.0 ~m (n = Buellia pullata Thck.

106) (Fig. 72). Lich. Californ. p. 26 (1866)

CHEMISTRY: No substances found, or with various amount TYPE: USA. California: Rocks on the coast. Bolander 131,
of the following secondary metabolites: norstictic and (FH-3280! - lectotype selected here!), Bolander 181, 150
connorstictic acid, atranorin, or several xanthones (HPLC: (FH-3280! - syntypes).
4,5-dichloro-3-0-methylnorlichexanthone, thuringione, 4,5-
dichlorolichexanthone, 2,4,5-trichlorolichexanthone). Spot TAXONOMIC NOTES: This species is not synonymous with
tests usually negative, rarely K+ yellow to orange, C+ orange. B. punctata, as suggested by Imshaug (1951). Several speci-
UV+ bright yellow to orange. Thallus without amyloid reac- mens from FH are mounted together on one large herbarium
tion, only the apothecia are amyloid in Lugol's. sheet with the FH accession number 3280. A type specimen
is not designated in the protologue. From the available mate-
SUBSTRATE AND ECOLOGY: Growing on siliceous mineral-poor rial we therefore select the specimen labeled Bolander (131)
coastal rock (generally HCI-). as the lectotype. Even though the other specimens are
mounted on the same sheet and have the same FH accession
DISTRIBUTION: Probably cosmopolitan but restricted to coastal number the specimens Bolander 181 and 150 cannot be re-
areas. In the Sonoran Region presently known only from garded as isolectotypes, because they have different collec-
southern California, USA., and Baja California, Baja Cali- tion numbers. Bolander 179, also mounted on the same
fornia Sur, Sonora and Sinaloa, Mexico (Fig. 71). sheet, has bacilliform conidia and it belongs to Buellia
christophii.
NOTE: In general, B. prospersa can be easily recognized by a
pale, distinctly epilithic thallus (Figs. 15, 16) and ascospores Figs. 8-14 and 64-69.
with a persistent median septum thickening (Figs. 18, 70).
Buellia pullata is microscopically similar, but has a deep Thallus (Figs. 8, 9, 64-69) crustose, usually rimose to
brown to dark gray thallus (Figs. 8, 9, 66), which does not rimose-areolate, thin and more or less continuous but dis-
react with UV light. No xanthones were detected in Buellia tinctly epilithic, never chasmolithic, rarely becoming thick-
pullata. Although not previously reported, thalli of B. pros- ened and distinctly rimose-areolate to subsquamulose, not
persa always react UV+ yellow to orange, and xanthones are delimited by a hypothallus; surface matt to more or less
characteristic for this species. Poorly developed thalli of shiny, usually deep brown, rarely pale brown, smooth,
B. prospersa sometimes appear more or less chasmolithic, epruinose, phenocorticate.
and these specimens are then difficult to separate from forms Apothecia soon sessile, lecideine; proper margin excluded
of B. sequax with a well-developed thallus (Fig. 42). If fili- with age; disk black, epruinose, initially plane, soon becom-
form conidia (Fig. 19) can be found, the material is easily ing convex; exciple of aethalea-type (Figs. 11, 12) sensu
recognized, because B. sequax has bacilliform conidia Scheidegger (1993), that is, inner excipular hyphae narrow,
(Fig. 24). In both species pycnidia are, however, quite rare. hyaline, prosoplectenchymatous (textura oblita), similar in
Young ascospores of B. sequax are typically distinctly nar- structure and orientation to the paraphyses, often more or
rowly oblong and not ornamented (Figs. 22, 70). Only less reduced and transient with the dull reddish brown
overmature and often disintegrating ascospores show a weak hypothecium (leptoclinoides-brown, textura inricata); outer
ornamentation, whereas a microrugulate ornamentation usu- excipular hyphae parallel (textura oblita), cells moderately
ally develops in young ascospores of B. prospersa (Fig. 70). swollen and usually strongly carbonized with various
Young ascospores of B. prospersa typically have a distinctly amounts of a brown pigment (cf. elachista-brown); pigmen-
thickened median septum (Figs. 18, 70), which can become tation continuous with the epihymenium; hymenium hyaline,
very pronounced in some specimens. However, in some not inspersed; paraphyses simple to moderately branched,
specimens it is difficult to find spores at this stage of the on- apically swollen, with a brown pigment cap (cf. elachista-
togeny. Median thickening of the spore septum is absent brown).
from all stages of ascospore development in B. sequax Asci 8-spored, clavate, Bacidia-type. Ascospores
(Fig. 70). In the Sonoran Desert Region, B. prospersa is (Figs. 14, 70) oblong to more or less ellipsoid, not con-
restricted to coastal California, USA, and Baja California, stricted, with obtuse ends, not curved, (9.0-)11.1-[12.5]-
Mexico (Fig. 71). Buellia sequax is widely distributed 13.8(-15.0) ~m x (5.0-)6.2-[7.1]-8.0(-10.0) ~m (n = 91),
throughout the Sonoran Desert (Fig. 71). one-septate; proper septum soon thickening during spore on-
togeny, becoming narrow with age (Le., with an indistinct
MATERIAL EXAMINED: MEXICO. Baja California: Schei- Physconia-type ontogeny), with septal pore canal, simple
degger s. n. (hb. Scheidegger); Nash 33982 (ASU). Baja pore and undifferentiated pore plug; ornamentation micro-
California Sur: Nash 12653, 12755 (ASU). Jalisco: Nash rugulate (conspicuous in DIC at an early stage of the ontog-
20765 (ASU). Sinaloa: Moberg 10267 (UPS); Wetmore eny); spore wall (Fig. 14) differentiated into smooth to
71896 (MIN); Nash 33680, 10088, 12248 (ASU); Sonora: cracked, thin perispore «0.20 ~m), narrow intermediate
Nash 25625, 12518, 12521, 10964 (ASU). ITALY. Sardinia: layer «0.10 ~m), moderately thick proper spore wall (0.3-
Scheidegger Inv. Nr. 11097 (hb. Scheidegger). NEW ZEA- 0.5 ~m), and moderately thickened endospore (0.20-
LAND. South Island. Canterbury: Nash 19098 (ASU); Blaha 0.40 ~m).
200 (GZU). Southland: Blaha 152 (GZU). USA. California. Pycnidia (Fig. 13) common, globose, unilocular; at matu-
Santa Barbara Co.: Nash 41312 (ASU). Ventura Co.: Nash rity almost entirely occupied by densely branched conidio-
38657 (ASU). phores; 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 con-
sists of swollen cells, which are carbonized by the dull brown pigment cf. elachista-brown and the aeruginose pigment cinereorufa-
green (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 micro-
graph 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 on- BuelUa ryanii Bungartz sp. nov.
togeny similar to the Roccella-type (sensu Vobis 1980, Vobis
and Hawksworth 1981); conidia (Fig. 13) simple, predomi- DIAGNOSIS: Thallus saxicolus, crustaceus, areolatus vel sub-
nantly filiform, 6.0-43.0 I.lm x <1.0 I.lm (n = 156). squamulosus, tenuis vel crassus, olivaceus. Hypothallus
atratus arachnoideus. Apothecia erumpentia vel sessilia, leci-
CHEMISTRY: No substances found. All spot tests negative. deina, marginibus propriis tenuibus. Excipulum tenue, fulvo-
Thallus without amyloid reaction, only the apothecia are am- caeruleum, pigmentum aeruginosum continens, carbon-
yloid in Lugol's. aceum. Asci 8-spori. Sporae unisaeptae, ellipsoideae vel
oblongae, 9-13 I.lm x 4-8 I.lm. Pycnidia globosa. Conidia
SUBSTRATE AND ECOLOGY: Growing on siliceous mineral-poor bacilliformia, 2-5 I.lm x 1-1.5 I.lm. Materiae chimicae
rock (generally HCl-). nullae.
DISTRffiUTION: Fairly common along the coast of southern TYPE: USA. California. Santa Barbara: Santa Cruz Island,
California, USA, Baja California, and Baja California Sur, 4.5 km E of radar station, ridge crest down N slope;
Mexico. One specimen has also been found further inland in 34°0'15"N, 119°37'30"W; altitude: 287 m; on schist in oak-
Sonora, Mexico (Fig. 71). pine woodland, 9 January 1994; Nash 32448b (ASU -
holotype designated here).
NOTES: The thallus morphology (Figs. 8, 9, 64-69) of this
species is very similar to that of B. christophii (Figs. 1, 2, TAXONOMIC NOTE: The species is named in memory of Dr.
55-62). Pycnidia are relatively common in both species and Bruce D. Ryan, the third author of this publication, who died
if filiform conidia are found (Fig. 13) the material is, thus, from cancer before this article was printed.
reliably distinguished from B. christophii, which has bacil-
liform conidia (Fig. 7). The two species can, however, also Figs. 29-34.
be clearly distinguished by other characters. Buellia chris-
tophii has lecideine apothecia with a thick, rather persistent Thallus (Figs. 29, 30) crustose, areolate to subsqua-
margin (Figs. 1-3). Buellia pullata has a thin margin, which mulose, usually delimited by a distinct black arachnoid
relatively soon becomes excluded (Figs. 8, 9, 11). Although hypothallus; surface matt to more or less shiny, deep olive
the structure of the proper exciple is virtually identical gray to brownish olive, smooth, epruinose, phenocorticate.
(compare Fig. 4 with Fig. 12), the layer of carbonized swol- Apothecia initially immersed appearing aspicillioid, soon
len outer cells is at least twice as thick in B. christophii bursting through the thallus surface and becoming adnate to
(Fig. 3). Young ascospores of B. pullata are oblong to ellip- sessile, lecideine, rarely with remains of necrotic thalline
soid (Fig. 70); in B. christophii, young ascospores are rather material attached to the margin (thalline veil, Fig. 30);
broad, and often appear almost globose (Figs. 5, 70). Spores proper margin more or less persistent, rarely excluded with
of B. christophii have no septum thickening (Fig. 70), age; disk black, epruinose, plane, rarely becoming convex
whereas a median thickening of the spore septum can usu- with age; exciple of aethalea-type (Fig. 31) sensu Schei-
ally be observed at least in some of the younger spores of degger (1993), that is, inner excipular hyphae narrow,
B. pullata (Fig. 70). hyaline, prosoplectenchymatous (textura oblita), similar in
structure and orientation to the paraphyses, often more or
MATERIAL EXAMINED:MEXICO. Baja California: Scheidegger s. less reduced and transient with the dull reddish brown hypo-
n. (hb. Scheidegger); Nash 38267, 38458, 38513b (ASU); thecium (leptoclinoides-brown, textura inricata); outer
Wetmore 75732 (MIN). Sonora: Wetmore 71644 (MIN). excipular hyphae parallel (textura oblita), cells moderately
USA. California. Los Angeles Co.: Weber L-42114 (COLO); swollen and usually strongly carbonized with various
Nash 32174 (ASU); Mendocino Co.: Nash 25476, 11426, amounts of brown and aeruginose (HN0 3+ violet) pigments
11424b (ASU). Monterey Co.: Nash 18896 (ASU). San Diego (cf. elachista-brown and cinereorufa-green); pigmentation
Co.: Bratt 8658 (SBBG). San Luis Obispo Co.: Thcker continuous with the epihymenium; hymenium hyaline, not
28829B (SBBG). Santa Barbara Co.: Bratt 6394, Thcker inspersed; paraphyses simple to moderately branched, api-
34719, Tucker 35711 (SBBG); Nash 32672, 41099, 41227, cally swollen, with a brown pigment cap (cf. elachista-
41249,41311,33012 (ASU), Wetmore 73713 (MIN). Ventura brown) and a diffuse aeruginose pigment (HN0 3+ violet,
Co.: Nash 37059, 37015 (ASU); D.E. Baltzo 7215 (UC). cinereorufa-green).
© 2004 NRC Canada
Bungartz at al. 549

Asci 8-spored, clavate, Bacidia-type. Ascospores premature spores, microrugulate in mature spores (best seen
(Figs. 32, 70) broadly oblong to ellipsoid, with obtuse ends, in DIe).
not curved, mature spores slightly constricted, (9.0-)10.5- Pycnidia (Fig. 33) rare, globose, unilocular; at maturity al-
[11.3]-12.2(-13.0) J.lm x (4.0-)5.6-[6.2]-6.8(-8.0) J.lD1 (n = most entirely occupied by densely branched conidiophores;
100), one-septate; proper septum narrow, not thickening dur- conidiogenous cells mostly terminal, but some also interca-
ing spore ontogeny; ornamentation absent in immature and lary (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, al-
most 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 sec-
tion): 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 crys-
tals, 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 (criti-
cally 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 frac-
tured 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, exciple and epihymenium of B. ryanii is dull olive-brown
Vobis and Hawksworth 1981); conidia (Fig. 34) simple, and the presence of cinereorufa-green may thus be over·
bacilliform to ellipsoid, 2.0-5.0 Jlm x 1.0-1.5 Jlm (n = 73) looked if not tested with HN03 • The spores of the two spe-
(Fig. 72). cies have a similar ontogeny and become constricted with
maturity (Fig. 70). However, spores of B. tergua are dis-
CHEMISTRY: No substances found and all spot tests negative. tinctly larger (10-15 Jlm x 6-9 Jlm) than those of B. ryanii
Thallus without amyloid reaction, only the apothecia are am- (9-13 Jlm x 4-8 Jlm). The ornamentation of Buellia tergua
yloid in Lugol's. spores develops early; it becomes visible at the premature
stage and is distinct at maturity (Fig. 70). In B. ryanii spore
SUBSTRATE AND ECOLOGY: All specimens examined grow on ornamentation is barely discernible in mature spores but
mineral-poor rock substrates such as quarzite and schist in well developed in overmature spores (Fig. 70). The conidia
open oak-conifer forest. of B. ryanii are slightly smaller (2-5 Jlm) than those of
B. tergua (4-7 Jlm) (Fig. 72).
DISTRIBUTION: The species is currently known only from
Santa Cruz Island, along the coast of southern California, MATERIAL EXAMINED: USA. California. Santa Barbara Co.:
USA, and from Isla de Guadalupe, off the coast of Baja Cal- Tucker 35781A, Bratt 3488b (SBBG); C.M. Wetmore 74077
ifornia, Mexico. It has not been found close to the sea shore (MIN); Nash 32445, Ryan 31621 (ASU).
but in open, dry oak-conifer woodland.
NOTES: Scheidegger (1993) mentions the presence of pigment Buellia sequax (Ny!.) Zahlbr.
A (= cinereorufa-green) as characteristic of the aethalea- Cat. Lich. Univ. 7: 410 (1931).
type exciple. All species discussed here, have an exciple Lecidea sequax Ny!., Flora, Jena 58: 302 (1875)
with the structure of the aethalea-type, but the pigment
cinereorufa-green is only present in the exciple and epihy- TYPE: FRANCE, Hautes Alpes: on quartzite along the banks
menium of B. ryanii and B. tergua. The two species are of the river Vienne near Moulin de l' Aiguille [Quartz sur les
quite similar and currently only known from a few localities coteaux de la Vienne pres du Moulin de l'Aiguille], 11 Janu-
off the coast of Baja California, Mexico and southern Cali- ary 1872, Lamy 1083 [H-NYL 9538 - lectotype selected
fornia, USA. Buellia ryanii has an areolate to sub- by Scheidegger (1993)], H-NYL 9539 - isolectotype, M-
squamulose, deep olive gray to brownish olive thallus with a 0023804! - isolectotype].
distinct arachnoid hypothallus (Figs. 29, 30). Buellia tergua Buellia abstracta (Ny!.) H. Olivier, Bull. Acad. Intren.
has a leather brown rimose-areolate thallus with no hypo- Geogr. Bot. 12: 176 (1903). - Lecidea abstracta Ny!., Flora,
thallus (Fig. 35). The apothecial ontogeny is very similar. Jena 66: 102 (1883). Type: FRANCE, Pyrennees: Cauterets,
Apothecia are initially immersed and the margin is covered E. Lamy (H-NYL 9740 - holotype, M-0023906! - isotype).
by the surrounding thallus. Thus, these apothecia look B. punctatula Malme, Arkiv fOr Botanik 21a(14):9 & 14
aspicillioid (Fig. 30). Soon apothecia erupt from the sur- (1927). Type: PARAGUAY. Asun~i6n: "Zapitapunta", on
rounding thallus and become adnate. Rarely, thallus material sunny sandstone [original label data: "Zapitapunta", ad rupes
remains attached to the lecideine margin as a thalline veil. arenarias sat apricas], 21 July 1893, Malme 1424 (S! -
The pigment cinereorufa-green is strongly concentrated in lectotype selected here).
the exciple and epihymenium of B. tergua and in cross sec- Buellia saxicola de Lesd., Ann. Crypt. Exot. 5: 127
tion clearly visible as deeply aeruginose. In contrast, the (1932). Type: USA. New Mexico. San Miguel: around Las
© 2004 NRC Canada
Bungartz et al. 551

Vegas, Agua Zarca, on siliceous rocks [original label data: identical with B. punctata as suggested by Imshaug (1951).
environs de Las Vegas, Agua Zarca, sur roches siliceusesl, The lectotypification of the material designated by Bouly de
Arsene Brouard 1930 (W acquisition 1934 no. 726! -lecto- Lesdain is necessary, because it can generally be assumed
type selected here; UPS!, S! - isolectotypes). that his entire holotype collection, originally located in
Dunkerque (France), has been destroyed during the Second
TAXONOMIC NOTES: The type material of B. saxicola is not World War. The lectotype from Vienna was selected be-
© 2004 NRC Canada
552 Can. J. Bot. Vol. 82, 2004

cause this is the only material that still has a few intact SUBSTRAtE AND ECOLOGY: Growing on a large variety of sili-
pycnidia. Mayrhofer and Moberg (2002) list Buellia myrio- ceous rocks (generally HCl-).
carpa (DC.) De Not. as a synonym of Buellia punctata
(= Amandinea punctata). In 1903, Hasse determined his DISTRIBUTION: The species is widely distributed throughout
exsiccati specimen no. 204 as Buellia myriocarpa (Lamb & the North American Southwest. In the Sonoran Desert Re-
DC.) Mudd. The exsiccati specimen at ASU is identical with gion, specimens have been found from coastal up to subal-
B. sequax. Type material of B. myriocarpa was, however, pine localities (Fig. 71).
not examined. NOlES: Specimens of B. sequax with an unusually well-
Figs. 20--28 and 41-54. developed thallus are sometimes superficially similar to
B. prospersa and diagnostic differences have been discussed
Thallus (Figs. 20, 41-54) crustose, thin and discontinu- under that species.
ous, more or less chasmolithic, that is, forming inconspicu-
SPECIMENS EXAMINED: AUSTRIA. Tirol: Arnold s. n. (M-
ous, poorly delimited granules hidden among the mineral
0061309). MEXICO. Baja California: Scheidegger s. n. (hb.
grains of the substrate; thicker thalli rarely becoming con- Scheidegger), Nash 26137, 38513a, 40151, 26336, 34608
tinuous in parts, rimose to very rarely rimose-areolate, not
(ASU). Baja California Sur: Scheidegger s. n. (hb.
delimited by a hypothallus; surface matt, pale brown to
Scheidegger); van den Boom 25063 (hb. van den Boom);
greyish, smooth to slightly roughened, epruinose, pheno-
Nash 33718, 33858, 40071, 40101, 26144 (ASU) Sonora:
corticate.
Ryan 21689 (ASU). ITALY. Elba: Albertshofer 6723 (M-
Apothecia soon becoming adnate to sessile; lecideine; 0061306), Triebel & Rambold 6209 (M-0061304),
proper margin soon excluded with age; disk black, eprui- Albertshofer s. n. (M-0061303). Sicily: Doppelbaur (M-
nose, plane, becoming convex with age; exciple of aethalea- 0061307). SPAIN. Murcia: Doppelbaur (M-0061305). USA.
type (Figs. 21, 23) sensu Scheidegger (1993), that is, inner Arizona. Apache Co.: Ryan 19184 (ASU) Cochise Co.:
excipular hyphae narrow, hyaline, prosoplectenchymatous Weber S-8754 (COLO); Darrow 1970 (ASU) Coconino Co.:
(textura oblita), similar in structure and orientation to the Nash 35171, Boykin 2725 (ASU) Gila Co.: Nash 28487,
paraphyses, often more or less reduced and transient with Schramm 225 (ASU) Maricopa Co.: Scheidegger s. n. (hb.
the dull reddish brown hypothecium (leptoclinoides-brown, Scheidegger); Nash 9605a (ASU) Pinal Co.: Scheidegger s.
textura inricata); outer excipular hyphae parallel (textura n. (hb. Scheidegger). California. Los Angeles Co.: Wetmore
oblita), cells moderately swollen and usually strongly car- 73249 (MIN); Hasse L-78819 (COLO); Weber L-42778, L-
bonized with various amounts of brown pigment (cf. 42776 (COLO); Bratt 10196 (SBBG); Nash 32187, Hasse
elachista-brown); pigmentation continuous with the epihy- Exs. 204, B.D. Ryan 30929b (ASU); Hasse Exs. 53 (CAS)
menium; hymenium hyaline, not inspersed, paraphyses sim- Merced Co.: Tucker 28820 (SBBG), Tucker 28815 (ASU).
ple to moderately branched, apically swollen, with a brown Monterey Co.: Nash 8035 (ASU). Orange Co.: Weber L-
pigment cap (cf. elachista-brown). 42052 (COLO). San Diego Co.: van den Boom 25192 (hb.
Asci 8-spored, clavate, Bacidia-type (Fig. 25). Ascospores van den Boom); Bratt 3592 (ASU) San Luis Obispo Co.:
(Figs. 22, 26-28, 70) distinctly narrowly oblong (Fig. 22), Tucker 28829A, 36471 (SBBG); Tavares 1409 (UC). Santa
becoming ellipsoid with age, not or rarely constricted with Barbara Co.: Tucker 35689A, 35845, 35781, 36463, 35781,
age, with obtuse ends, not curved, (10.0--)10.7-[11.7]- Bratt 11388, 5709, 3641, 3643, 10316, 3488a (SBBG);
12.7(-14.0) Jlm x (3.0--)3.8-[4.6]-5.4(-6.0) Jlm (n = 50), Printzen 74, 76 (hb. Printzen), Nash 32352, 32656, 32747,
one-septate, proper septum narrow, not thickened during spore 32837, 41392, 41423, 33016, 33018, 32890; Bratt 3488,
ontogeny, with septal pore canal, simple pore and undifferen- 7681, 8879A, Ryan 31216 (ASU). Ventura Co.: Bratt 10392,
tiated pore plug (Fig. 28); ornamentation absent from the Hamber 7 (SBBG) New Mexico. San Juan Co.: Nash 16225,
early spore ontogeny (Fig. 26), becoming visible in mature 16396 (ASU). Valencia Co.: Nash 16004 (ASU). Utah.
spores (Fig. 27); spore wall differentiated into smooth to fis- Washington Co.: Nash 15316 (ASU). Wyoming. Platte Co.:
sured, thin perispore (0.1-0.2 Jlm), narrow intermediate layer Ryan 14306 (ASU).
«0.1 Jlm), thick proper spore wall (0.3-0.5 Jlm) and moder-
ately thickened endospore (0.2-0.3 Jlm). Buellia tergua Bungartz sp. nov.
Pycnidia rare, globose, unilocular; at maturity almost en- DIAGNOSIS: Thallus saxicolus, crustaceus, rimosus vel rimoso-
tirely occupied by densely branched conidiophores; conidio- areolatus, tenuis vel crassus, alutaceus, sine hypothallo.
genous cells intercalary and terminal (conidiophore-type V Apothecia erumpentia vel sessilia, lecideina, marginibus pro-
sensu Vobis 1980); pycnidial ontogeny similar to the priis tenuibus. Excipulum tenue, aeruginosum, pigmentum
Umbilicaria-type (sensu Vobis 1980, Vobis and Hawksworth aeruginosum continens, carbonaceum. Asci 8-spori. Sporae
1981); conidia (Fig. 24) simple, bacilliform, 2.0--4.0 Jlm x unisaeptae, ellipsoideae vel oblongae, 10--15 Jlm x 6-9 Jlm.
1.0--1.5 Jlm (n = 50) (Fig. 72). Pycnidia globosa. Conidia bacilliformia, 4-7 Jlm x 1-
1.5 Jlm. Materiae chimicae nullae.
CHEMISTRY: No substances found or with norstictic acid or
arthothelin (low concentrations only detected by TLC or TYPE: MEXICO. Baja California, Isla de Guadalupe: SE of
HPLC). All spot tests negative (very rarely forming orange southern peak; 28°57'30"N, 118°15'00"W; altitude 600 m;
needle-shaped crystals with K if observed with the com- on basalt, shaded exposure within canyon with rocky out-
pound scope). UV-. Thallus without amyloid reaction, only crops and some native perennials; 3 January 1996, Nash
the apothecia amyloid in Lugol's. 38368 (ASU - holotype designated here).

© 2004 NRC Canada

Bungartz et al. 553

TAXONOMIC NOTE: The species name is derived from the Latin species with cinereorufa-green in the apothecia and diagnos-
word for leather, tergus. tic differences are presented in the notes on that species.
Figs. 35-40. MATERAL EXAMINED: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. Discussion
Apothecia initially immersed appearing aspicillioid, soon
Thallus variation: chasmolithic versus epilithic growth
bursting through the thallus surface and becoming adnate to
All species discussed here are often described as having a
sessile, lecideine, rarely with remains of necrotic thalline
scant or obsolete thallus and were previously interpreted as
material attached to the margin (thalline veil); proper margin
poorly developed forms of B. punctata rather than being
more or less persistent, rarely excluded with age; disk black,
assigned an independent species rank. However, in the So-
epruinose, plane, rarely becoming convex with age; exciple
noran Desert the saxicolous species of the "Buellia punc-
of aethalea-type (Figs. 36, 37) sensu Scheidegger (1993),
tata" group show considerable diversity. The thallus
that is, inner excipular hyphae narrow, hyaline, prosop~ec­
morphology of the species is quite diagnostic and far less
tenchymatous (textura oblita), similar in structure and onen-
variable than commonly implied.
tation to the paraphyses, often more or less reduced and
The extent to which a thallus is developed on the surface
transient with the dull reddish brown hypothecium
or partially hidden within crevices, pores, or fissures is nec-
(leptoclinoides-brown, textura inricata); outer excipular
essarily related to substrate composition and structure. Both
hyphae parallel (textura oblita), cells moderately swollen
epilithic and chasmolithic thalli clearly show some degree of
and usually strongly carbonized with various amounts of
substrate penetration, provided that enough space is avail-
brown and aeruginose (HN0 3+ violet) pigments (cf.
able (Figs. 41-69). The species examined here all grow on
elachista-brown and cinereorufa-green); pigmentation con-
mineral-poor weathering rinds of various siliceous rocks. Al-
tinuous with the epihymenium; hymenium hyaline, not in-
though the composition of these substrates varies consider-
spersed; paraphyses simple to moderately branched, apically
ably, they are all poor in nutrients such as iron, magnesium,
swollen, with a brown pigment cap (cf. elachista-brown) and
or calcium, and no calcium carbonates were found. The fun-
a diffuse aeruginose pigment (HN03+ violet, cinereorufa-
gal hyphae are apparently not able to dissolve their substrate
green).
and instead, hyphal penetration is confined to the. .P~­
Asci 8-spored, clavate, Biatora-type. Ascospores existing crevices (e.g., Figs. 53, 54, 60, 61). Both epilithic
(Figs. 38, 70) broadly oblong to ellipsoid, with obtuse ends, and chasmolithic thalli can thus be distinguished from truly
not curved, mature spores slightly constricted, (10.0-)11.1- endolithic lichens, which develop entirely within their sub-
[12.2]-13.4(-15.0) Jlm x (6.0-)6.0-[6.6]-7.3(-9.0) Jlm (n = strate and are at least partially able to dissolve some of the
68), one-septate; proper septum narrow, not thickening dur- minerals (Bungartz et al. 2004).
ing spore ontogeny (Le., Buellia-type); ornamentation absent Generally, chasmolithic growth (Figs. 41-54) and epilithic
in immature and premature spores, microrugulate in mature growth can be distinguished (Figs. 55-69): epilithic thalli
spores (best seen in DIC). are well developed on the substrate surface, internally dis-
Pycnidia (Fig. 39) rare, globose, unilocular; at maturity al- tinctly stratified (Figs. 10, 63), and have a more or less con:
most entirely occupied by densely branched conidiophores; tinuous surface (Fig. 58, 59, 64, 65). Although these thallI
conidiogenous cells intercalary and terminal (conidiophore- may be thin and inapparent, they are nevertheless largely
type V sensu Vobis 1980); pycnidial ontogeny similar to the covering the substrate (Figs. 1, 2, 8, 9, 55, 66). In contrast,
Umbilicaria-type sensu Vobis 1980, Vobis and Hawksworth chasmolithic thalli grow poorly differentiated among the
1981); conidia (Fig. 40) simple, bacilliform, 4.0-7.0 Jlm x mineral grains (Figs. 41, 44, 47, 50), these thalli are less
1.0-1.5 Jlm (n = 50). stratified and sometimes only consist of loosely associated
hyphae (Figs. 50-52). Even well-developed chasmolithic
CHEMISTRY: No substances found and all spot tests negative.
thalli (Figs. 42, 43) do not establish a continuous surface
Thallus without amyloid reaction, only the apothecia are am- crust. Among the species examined here, only B. sequax
yloid in Lugol's. may consistently be referred to as chasmolithic, that is, the
SUBSTRATE AND ECOLOGY: On siliceous volcanic coastal rock. major part of the thallus remains hidden amo~g the m,ineral
grains, even though some areas may develop Into a dIscon-
DISTRIBUTION: Presently known only from Isla Guadalupe, off tinuous crust (Fig. 47). Specimens of B. sequax with well-
the coast of Baja California, and two localities on the Pacific developed thallus areas can be distinguished from the other
coast of mainland Baja California, Mexico (Fig. 71). epilithic species by the pale thallus colour. Only poorly de-
veloped or damaged thalli of B. prospersa may be confused
NOTES: The thallus of B. tergua is very similar to B. pullata with these well-developed specimens of B. sequax. Com-
(Figs. 8, 9) and B. christophii (Figs. 1, 2), even though .these paring the types of B. abstracta and B. sequax, Scheidegger
two species are generally darker brown. Apothecia of (1993) argued that B. abstracta s. str., with narrow as~o­
B. tergua, erupt from the thallus and young apothecia thus spores and a chasmolitic thallus, cannot reasonably be d~s­
appear aspicillioid. In B. pullata and B. christophii, apothe- tinguished from epilithic and often areolate mat:nal
cia emerge more gradually and young apothecia appear im- assigned to B. sequax s. str. He therefore synonymIzed
mersed but not aspicillioid. Buellia ryanii is the only other 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 as-
pect on rhyolithe (Nash 321(0). Fig. 56. Light micrograph of a thallus on rhyolite (substrate cross section): the thallus is distinctly de-
veloped 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 sur-
face. 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 resin-
embedded, 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 sy- All species treated here have very similar ascospores, and
nonymized. Buellia sequax is, however, far less variable measurements overlap considerably. Nevertheless, diagnostic
than this statement may imply. Chasmolithic growth is not differences can be found in all species by carefully compar-
merely the result of substrate variation, because B. sequax ing their spore ontogeny (Fig. 70). For convenience, four
consistently forms chasmolithic thalli on a variety of differ- stages may be distinguished and some characters are only
ent substrates. We have examined type material of the two present during a brief stage: immature spores are hyaline,
taxa as well as material from M annotated by Scheidegger. premature spores olive, mature spores brown, and over-
Buellia sequax s. str. indeed shows more or less larger and mature spores deep brown. Immature and premature spores
better developed thallus areas, but even well-developed generally have no visible ornamentation, although in some
thalli remain poorly delimited patches and never spread ex- species ornamentation may begin to develop with the onset
tensively into a continuous crust. of pigmentation. Other species barely show any ornamenta-
Among the species discussed here, only B. ryanii forms a tion during their ontogeny, but overmature spores often
distinctly areolate thallus (Figs. 29, 30). The areoles form appear ornamented. This may sometimes be an artefact
independently on a black, arachnoid hypothallus, and indi- ascribed to degeneration of the spores. Wall or septum thick-
vidual areoles regularly become subsquamulose along the enings are usually best developed in premature and mature
margin. All other species have a rimose to rimose-areolate spores and may disappear again in aging spores.
thallus, that is, the surface is initially fissured and only sec- In Rinodina, Giralt and Mayrhofer (1994, 1995) distin-
ondarily breaks into areoles. In exceptionally thick thalli of guish sporesthat form spore septa before apical wall thicken-
B. christophii and B. pullata these secondary areoles can ings (A-ontogeny) from spores with late septum formation,
even become subsquamulose. Generally, thalli of B. christo- where apical thickenings are present before the development
phii, B. pullata, and B. tergua are very similar and easily of a septum (B-ontogeny). Giralt (2001) admits that some
confused (Figs. 1, 2, 8, 9, 35). Buellia tergua is lighter species have an intermediate ontogeny. In spores typically
brown than the deep brown thalli of the other two species. found in Buellia, apical thickenings generally do not develop
However, only a few specimens of B. tergua are currently (Fig. 70). Early septum formation may better be distin-
known, and it is therefore not possible to assess whether the guished from late septum formation based on spore wall
leather-brown colour represents a consistent trait. The three pigmentation. In spores with A-ontogeny, pigmentation de-
species may therefore not be reliably distinguished by their velops after a septum can be distinguished in the immature
thalli alone. spore. Late septum formation of the B-ontogeny develops in
premature spores after the walls have become distinctly pig-
Spore ontogeny and ultrastructure mented. Buellia tergua and B. sequax can tentatively be
In crustose Physciaceae, spore structure has attracted con- assigned to the A-ontogeny, whereas spores of the other spe-
siderable attention (Mayrhofer 1982), and especially in the cies appear to follow the B-ontogeny. This distinction is not
genus Rinodina, several diagnostic types have been recog- easy to assess. Even spores with pigmented walls do not al-
nized because of conspicuous wall and septum thickenings ways have septa, and sometimes these spores can also be ob-
(Mayrhofer and Poelt 1979; Mayrhofer 1984; Giralt 2001). served along hyaline spores with septa. A- and B-ontogeny
In Buellia, Scheidegger (1993) first discovered thickened thus does not appear to be distinctly different in Buellia s. 1.
spore septa in a few species, and Kalb (1986) segregated the Among the specimens examined here, spores of B. pros-
genus Hafellia for species with lateral wall thickenings. The persa are most easily recognized (Fig. 70). They are the
wall thickenings of spores in the Physciaceae are, however, largest spores and have a conspicuous septum thickening
by no means static but the result of a dynamic spore ontog- most distinctly observed in premature spores. This septum is
eny, which is still not completely understood. persistent in mature spores, but eventually reduced in over-

© 2004 NRC Canada

Bungartz at al. 555

mature spores. Because of the thickening, these spores have never present, and the spores could thus also be referred to
been referred to the Physconia-type (Scheidegger 1993). In as the Orcularia-type. Mayrhofer et al. (1999) described
some spores, the septum remains conspicuously thickened a similarly persistent septum thickening in Amandinea
during almost the entire ontogeny. Apical thickenings are insperata (Nyl.) H. Mayrhofer & Ropin. Differences be-

© 2004 NRC Canada

556 Can. J. Bot. Vol. 82, 2004

Fig. 70. Ascospore ontogeny of Buellia species from the tween the two types seem not very distinct. especially if api-
Sonoran Desert with inconspicuous, saxicolous thalli. Immature cal wall thickenings cannot be observed in the Physconia-
ascospores are hyaline, mostly nonseptate (B-ontogeny), but type.
some early septate (A-ontogeny). An olive pigmentation begins A faint and inconspicuous thickening can also be seen in
to develop at the premature stage, some spores also showing the spore ontogeny of Buellia pullata. where it may easily
Bue/lia christophii be overlooked. These spores could thus be assigned alterna-

r··",\\\ tively to the Buellia (= Beltraminea)- as well as the

/""~"":':'.

ft:~' (I
Physconia-type (Fig. 70). A similar situation has been dis-
cussed for B. dispersa. where the septum thickening is also
t:..\ .n restricted to a brief phase of the ontogeny (Bungartz et al.
''- II
\....:..=j
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

e8 8
Bue//ia ryanii stage when they are very broad to almost globose and have a

n
~

l \)
conspicuously, evenly thickened spore wall (Fig. 5). The
broad spore wall of these premature spores is also distinct in
~j 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 previ-
ously 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
Bue/lia pu/lata have examined type material of both taxa (see the discussion
,nII

8
j! on thallus variation). The spores are not very variable but
;f t\ basically follow the same ontogeny, with a premature stage
\\ /j 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 orna-

,
mentation of B. sequax develops early, but it is inconspicu-
ous and can usually only be distinguished in overmature
Bue/lia sequax spores. Nordin (1997, 2000) emphasized spore ultrastructure
of Buellia species with pluriseptate ascospores. In the spe-

{
888
f
~=<i

'J
i\
"""ij
cies studied here, TEM yields little additional information. It
is noteworthy. however. that in B. sequax, spores with a frac-
tured as well as a smooth perispore have been observed in
the TEM (Figs. 26. 27). This emphasizes the dynamic devel-
opment 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 con-
trast. spores of B. tergua have thicker walls, which early de-
velop an ornamentation. The ornamentation is best seen in
mature and overmature spores.
hyaline olive brown deep brown
immature premature mature overmature
Taxonomy
Some of the species described here have bacilliform coni-
dia (Figs. 7, 24, 34, 40, 72) and others filiform conidia

© 2004 NRC Canada

Bungartz et al. 557

Fig. 71. Distribution of Buellia species with inconspicuous saxicolous thalli'in southwestern North America. In the Sonoran Desert Re-
gion, 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

o
Buellia sequax
500
Buel/ia
'---_ _tergua
----"~ ~~
__"'__'_'>!..:W.~"""'_'_'__._i:l
lO9"l9'W

. . Arizona Upland

Lower Colorado Region

==
Subdivisions ofthe Sonoran Desert Floristic Province (Shreve & Wiggins 1964):
Magdalena Region

11IIII r~~~~:F~~iliils ofSonora)

Iml Central GulfCoast
I!mmmmi Vizcaino

© 2004 NRC Canada

558 Can. J. Bot. Vol. 82, 2004

(Figs. 13, 19,72). Otherwise, all species are very similar. In- Fig. 72. Range of conidial length in various saxicolous species
deed, this similarity is the main reason why the species were of Buellia (modified from Scheidegger 1993). The bars indicate
previously generally identified as B. punctata s. 1. We can the range of measurements for each of the species. Buellia
currently not confirm that any saxicolous specimens from pullata overall has not only the longest conidia but also the larg-
the Sonoran Desert Region belong to B. punctata s. str. Ac- est variation in measurements. For Beullia prospersa, the bold
cording to Mayrhofer and Moberg (2002, p. 9), "saxicolous bar indicates the range of measurements given by Scheidegger
and muscicolous material usually placed under this name is (1993), and the narrow bar indicates the variation observed here.
much in need of taxonomic revision". Scheidegger (1993, Measurements overlap considerably, and the asymptotic graph
p. 343) mentions that "Saxicolous material, usually placed shows that the distinction of bacilliform versus filiform is arbi-
under this name, is possibly not homogeneous and is not yet trary and does not represent a discrete character.
completely understood by the author...... According to B.paDllIa
Scheidegger's (1993) description, A. punctata s. str. has B. prospena
B.coniops
spores without a septum thickening, no spore ornamentation, B.punctata
and short fUiform conidia. Sheard and May (1997) noticed B. subdlsciformis
B. lesserata
that the North American material, which they assigned to B.jugorum
A. puncata, has ornamented spores, and a septum thickening B. longispora
can be observed at least during some short period of spore
development. They also report that they have only seen a
B. sardinienstJ
B. saxorum
B.dispena
B. almeriemis
-
•
asymptotic graph
of the measurements
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
B. aethalea
B. spuria
B. concinna
.-
B. ocel/ata
from the Southwest, and it is possible that the specimens B. bama
identified as A. punctata do not strictly belong to that taxon B. leptocline
B. terglUl
(l.W. Sheard, personal communication). B. stella/oto
From the taxa treated here, the newly described species B. vilis
B. ry"m;
are all distinct from B. punctata s. str. because of their short, B. elrrlstoplrll
bacilliform conidia (Figs. 7, 34, 40, 72). In addition, B.s~fllax

B. ryanii and B. tergua are also characterized by having the 10 15 2'0 25 30 35 40 45

apothecial pigment cinereorufa-green, which is absent from conidial length (11m)
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 species from North America into Amandinea. These North
specimens. American species all have fUiform conidia. Most recently,
The genus Buellia s. 1. currently represents an amalgam of Mayrhofer and Sheard (2002) transferred Rinodina
not necessarily closely related species, and various attempts cacuminum (Th. Fr.) Malme into Amandinea because of its
have been made to subdivide this large and heterogeneous fUiform conidia, also citing "preliminary molecular results"
group into smaller, more strictly defined genera. In the pres- (p. 440), which do not support the inclusion of that species
ent publication, we have examined a group of very similar in Rinodina.
species, some of which could have been treated within the Marbach (2000) argued that Sheard and May (1997) pro-
genus Amandinea, solely justified by the filiform conidia. vided an emended genus concept for Amandinea, allowing
Amandinea, however, does not currently constitute a more the inclusion of species where no conidia could be observed.
precisely defmed taxonomic group than Buellia. When Bungartz and Nash (2004b) questioned this approach. They
Scheidegger (1993) described Amandinea, he provided the demonstrated that Buellia turgescens Nyl. ex Tuck., a spe-
first valid description for a genus previously suggested, but cies that Marbach (2000) included in Amandinea without
not validly published, by Choisy (1950). Choisy (1950) dis- having studied the pycnidia, is synonymous with B. badia
tinguished Amandinea from Buellia because of the "pycno- (Fr.) A. Massal. Bungartz and Nash (2004b) showed that
conidies aciculaires", that is, the acicular conidia. Buellia badia is a species with bacilliform conidia, which
Scheidegger (1993) emphasized the differences in conidial therefore must not be included in Amandinea.
length and conidiophore structure as the only diagnostic cha- There is currently little agreement about the characters
racters of the new genus Amandinea. Choisy (1950) trans- characterizing the genus Amandinea. Amandinea may also
ferred Buellia coniops (Wahlenb.) Ach., B. myriocarpa (DC. be characterized by the scarcity of lichen secondary metabo-
ex I. M. Lamb. & DC.) De Not., and B. stigmatea Korb. into lites in the thallus, as suggested by Christoph Scheidegger
the new genus. Scheidegger (1993) selected Amandinea (personal communication). Scheidegger (1993) did not de-
coniops as the type of Amandinea, and also included tect any secondary metabolites in the species he included in
Rinodina lecideina and Buellia punctata. Subsequently, the the genus. Also, Sheard and May (1997) did not document
genus became more widely accepted. Matzer et al. (1994) any secondary metabolites in the North American species.
transferred Rinodina petermannii (Hue) Darb. into Matzer et al. (1994), however, report norstictic acid and er-
Amandinea, emphasizing its filiform conidia. The species in- gosterol peroxide from A. petermannii. Even though
vestigated by Scheidegger (1993), as well as Aman- Scheidegger (1993) did not detect secondary metabolites in
dinea petermannii, are all saxicolous. Sheard and May A. lecideina (= B. prospersa), that species is clearly charac-
(1997) transfered several corticolous Buellia and Rinodina terized by a pale UV+ yellow to orange thallus, a reaction

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Bungartz et al. 559

caused by various xanthones. Several species, which Roccella-type pycnidium (sensu Vobis 1980) therefore rep-
Marbach (2000) included in Amandinea, also have charac- resent the only reliable diagnostic character consistently
teristic secondary metabolites. In addition, Marbach (2000) used to segregate Amandinea from Buellia. Within the
mentioned that Amandinea species often have a pale Physciaceae, conidial length has also been used to distin-
hypothecium, a character more commonly associated with guish Physcia from Phaeophyscia (Moberg 1977), Physcia
Rinodina rather than with Buellia. The species treated by from Hyperphyscia (Choisy 1950, Hafellner et al. 1979), and
Scheidegger (1993) (including the type), and most of the to characterize the genus Mobergia (Mayrhofer et al. 1992).
species treated by Sheard and May (1997), however, have a None of these genera, however, are distinguished solely by
dark hypothecium. All species treated here also have a dark the length of their conidia, and the foliose genera are also
brown hypothecium. well circumscribed by a range of other characters.
In his treatment of corticolous Buellia s. I., Marbach The shape of conidia (bacilliform vs. filiform) is a result
(2000) emphasized branching patterns of the paraphyses and of conidial length, and it does not represent a general differ-
the pigmentation of their apical cells as taxonomically im- ence in conidial structure. Conidial length may vary consid-
portant. According to Marbach (2000, p. 52) large, globose erably, even within the same pycnidium of a single
apical cells are characteristic for Amandinea. "Tafel 1/2: specimen. In fact, B. pullata has predominantly 20-25 Jlm
Paraphysenenden" (pp. 36 and 37, Marbach 2000), however, long filiform conidia, but overall, the conidia measured from
documents only very subtle, if any, differences in the para- a single pycnidium of that species vary from 6 to 43 Jlm in
physes of Buellia s. I. No distinct differences that would jus- length (Fig. 72). The shortest conidia are thus bacilliform
tify a clear segregation of the genus Amandinea are rather than filiform. In B. prospersa (= A. lecideina), the
displayed in this chart. Marbach (2000) did not discuss the range of our measurements (7-32 Jlm, Fig. 72) considerably
different apothecial pigment types in any detail, even though exceeds the variation observed by Scheidegger (1993), who
these differences are clearly taxonomically important (e. g., measured conidia from 15-30 Jlm long. Scheidegger (1993,
in B. ryanii and B. tergua; see also Bungartz and Nash Fig. 5) plotted conidial length of the species in his treatment
2004a). In addition to the paraphysal characters, Marbach to demonstrate the differences between Buellia and Aman-
(2000, p. 52) emphasized that spores without wall thicken- dinea. The shortest conidia were found in Buellia vilis (2.5-
ings are typical for Amandinea. Nevertheless, he included 4 Jlrn). Buellia subdisciformis was the species with the
several species within the genus that have distinctly thick- longest conidia (9-14 Jlm) still treated within Buellia, but
ened septa. As previously discussed, wall thickenings vary conidia of A. punctata were only slightly longer (up to
considerably during spore ontogeny, and the spores of some 15 Jlm). Both A. lecideina and A. coniops had the longest
species (like B. pullata) are difficult to assign to a particular conidia (15-30 Jlm). Plotting our measurements of B. pullata
type. and B. prospersa (= A. lecideina) to this chart results in con-
Conspicuously thickened spore septa are characteristic for siderable overlap, and conidial length can therefore no lon-
several species transferred from Rinodina into Amandinea. ger be regarded as a discrete character. An asymptotic graph
When Scheidegger (1993) validated Choisy's genus descrip- more accurately describes the situation (Fig. 72).
tion of Amandinea, he included two species with distinctly In addition to the conidial length, differences in branching
thickened septa, including the type A. coniops. By transfer- patterns of the hyphae bearing conidia can also be observed.
ring R. insperata into Amandinea, Mayrhofer et al. (1999) Conidiophore structure is, however, closely related to the
also explicitly included a species with distinctly thickened length of the conidia and cannot be treated as an independ-
spore septa. However, taxa transferred from Buellia (e.g., ent character. Long, filiform conidia are generally borne on
Buellia punctata) often lack a distinct septum thickening. shorter and less extensively branched conidiophores. This is
In addition to the filiform conidia, Giralt et al. (2000, a necessary consequence of ejecting conidia from the pyc-
p. 521) provide a list of the following characters typical for nidium. Long filiform conidia would become entangled in
Amandinea: "apothecia lecanorine or lecideine, ascospores densely branched conidiophores. Densely branched conidio-
brown, one-septate, Buellia- or Physconia-type, often with phores can, however, produce larger quantities of short,
rugulate ornamentation; and thallus with norstictic acid or bacilliform conidia and can thus be found in species that
more frequently without secondary lichen compounds....". have only short conidia.
All these characters are regularly found in Buellia and are by In lichen taxonomy, conidia alone have not generally been
no means exclusive to Amandinea. A lecanorine apothecium used to distinguish taxa at the generic level. In some groups,
is generally more common in Rinodina, but some species in conidia of very different lengths can even be found on a sin-
Buellia also show the tendency to form a thalline exciple. In gle thallus. The genus Micarea regularly has both micro-
addition, immersed lecideine apothecia often emerge from conidia and macroconidia formed in different pycnidia on
the thallus, with some thallus material remaining attached to the same thallus. In other groups, conidia of different length
the exciple (see B. ryanii and B. tergua). The ascus of occur in closely related species of the same genus. For ex-
Amandinea is generally described as the Bacidia-type ample, Lecanora himalayae Poelt (from Nepal) and two very
(Marbach 2000, Giralt 2001), and it is thus not different similar species, Lecanora chondroderma Zahlbr. (from
from Buellia s. str. (Rambold et al. 1994). Sl1lChting et al. China) and Lecanora beamanii B. D. Ryan (from high eleva-
(2004) recently transferred several arctic species from Buel- tions in southern Mexico), have straight, 6-8 Jlm long
lia into Amandinea, arguing that in the absence of molecular conidia. In contrast, Lecanora maxima Lynge has curved
data this transfer is only justified because of the filiform conidia that are 11-17(-20) Jlm long. Nevertheless, all taxa
conidia of those species. belong to the section Dactylon Poelt that presumably forms
The filiform conidia borne on type III conidiophores in a a natural group isolated from other species of Lecanora

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560 Can. J. Bot. Vol. 82, 2004

(Ryan 1989). Other genera, for example, Haematomma clade with Bacidia-(or Biatora)-type ascus, usually a dark
(Rogers and Hafellner 1988) or Aspicilia (Clauzade and apothecium, and spores that less frequently have wall thick-
Roux 1984), also include some species with bacilliform and enings (e.g., BueLlia, Dimelaena, Dermatiscum, etc.). The
others with filiform conidia. "Caliciaceae" apparently also belong to this second clade
In the Physciaceae, the validity of using filiform versus (Wedin et al. 2002). The resolution within the clades is cur-
bacilliform conidia as the sole character to justify Aman- rently not sufficient to justify further segregation of the
dinea has recently been questioned. Because of their miform crustose genera Rinodina s. 1. and Buellia s. 1. Until a more
conidia, some terricolous species of the BueLlia epigaea resolved phylogeny is achieved, using both classical and
group would have to be formally transferred into Amandina, molecular data, we advocate including the species currently
even though they do not appear to be closely related to other treated as Amandinea within the genus Buellia s. 1.
species with miform conidia (H. Mayrhofer and U. Grube, Excluding species with filiform conidia from Buellia does
personal communication; Grube and Amp 2001; Trinkaus et not resolve the nomenclatural problems in this rather large
al. 2001). Preliminary molecular studies also do not confirm and heterogeneous genus. According to the code of Botani-
Amandinea as a monophylletic group (H. Mayrhofer per- cal Nomenclature (Greuter et al. 2000), B. disciformis (Fr.)
sonal communication; Grube and Amp 2001; Wedin et al. Mudd. is currently the listed type of the conserved genus
2002). Buellia. Unfortunately, the genus HaJellia Kalb., H. Mayr-
In some lichen species, pycnidia are very rare and are thus hofer & Scheid. is also based on the same type. To avoid this
rarely found in all of the specimens examined. If pycnidia conflict, Moberg et al. (1999) suggested conserving Buellia
are rare, it may sometimes be difficult to assess if these with Buellia aethalea (Ach.) Th. Fr. as a new type. If this
conidia belong to a particular lichen specimen or instead to a proposal is rejected, most species now treated within
lichenicolous fungus growing inside this specimen. A char- Buellia s.l. would then have to be accommodated in a new
acter that is frequently absent and difficult to assess does not genus. This may include some of the species currently
seem to be particularly suitable as the sole diagnostic feature treated within Amandinea. The name Buellia s. str. would
to circumscribe a genus. then refer to species generally treated as HaJellia, a genus
A monophylletic genus Amandinea, with A. coniops as the that would become obsolete. None of the species in Aman-
type, will most likely not include a large portion of species dinea is closely related to that group. If, however, the pro-
currently included by various authors. In the Sonoran Desert, posal by Moberg et al. (1999) is accepted, some of the
the B. mamillana group (F. Bungartz, unpublished data) in- species currently transferred into Amandinea may eventually
cludes species with xanthones that all have long bacilliform tum out to be more closely related to the B. aethalea group
to fusiform conidia. Within Buellia s. 1., that species group is and may still be better accommodated within BueLlia.
isolated, but not closely related to species commonly treated
within Amandinea. Because of its yellowish thallus contain-
ing xanthones, Buellia prospersa (= A. lecideina) shows Acknowledgements
some affinities with the B. mamillana group. Buellia pullata, We are obliged to Dr. Laurence A. Garvie, Department of
on the other hand, is fairly similar to the type species A. co- Geology, Arizona State University (ASU), for access to a
niops, even though the spore septa are hardly thickened dur- slow-speed diamond stone saw. Dr. Donald M. Burt, Depart-
ing their ontogeny. ment of Geology (ASU), identified the rock substrates of the
In summary, the taxonomy of Amandinea is not well re- specimens sectioned. The chemistry of all specimens was
solved. Segregating the species treated here into separate analyzed with TLC at ASU. Specimens from the private her-
genera only on the basis of conidial length would result in a barium of Dr. Christoph Scheidegger were also examined
constrained and arbitrary taxonomy. Nimis (1998) discussed with TLC by Martin Frei Swiss Federal Institute for Forest,
three main criteria that justify the recognition of a genus in Snow and Landscape Research (Birmensdort). Additionally,
lichens: (1) monophylly, (2) phylogenetic analysis of its selected specimens were analyzed with HPLC by Dr. Jack
taxa, and (3) several independent characters that circum- A. Elix from the Australian National University in Canberra.
scribe the new genus. The genus concept of Amandinea cur- Dr. Ulrik S~hting, University of Copenhagen, Denmark,
rently matches none of these criteria. "If only one character and Julia Blaha and Ulrike Grube, University of Graz, Aus-
is involved, this is indeed the weakest possible evidence. It tria (GZU), contributed valuable information to the discus-
could easily be in conflict with the next character to be dis- sion of the taxonomy of Amandinea. Scott Bates (ASU) and
covered..." (Nimis 1998, p. 432). This is exactly the situa- Dr. John Sheard, University of Saskatchewan, Canada,
tion in the genus Amandinea. Including species solely on the kindly reviewed first drafts of the manuscript. Dr. Helmut
basis of conidial length does not help to resolve the taxon- Mayrhofer (GZU) and an anonymous reviewer provided
omy of Rinodina or Buellia, both of which remain large, ar- valuable input to the final version of this publication.
tificial, and heterogeneous genera. Dr. Christian Printzen, Senckenberg Institut, Germany, was
Several recent treatments (Grube and Arup 2001; Wedin very helpful correcting the Latin diagnoses. Dr. Donald
et a1. 2002; Helms et al. 2003) have confirmed two general Pinkava, ASU, provided much support on taxonomic ques-
clades in the Physciaceae: (1) the Physcia (or Rinodina) tions regarding the application of the ICBN. We would like
clade, characterized by a Lecanora-type ascus, a usually to thank all herbaria, which have made specimens available
hyaline hypothecium, and spores that regularly have wall for study, especially Julia Blaha (GZU), who, on short no-
thickenings (e.g., Physcia, Heterodermia, Anaptychia, tice, supplied several specimens of B. prospersa, including
Mobergia, Tornabea, Rinodina, etc.) and (2) the Buellia the type of A. lecideina. On very short notice, Dr. Orvo

© 2004 NRC Canada

Bungartz at al. 561

Vitikainen, Helsinki (H) provided the Nylander specimen of Giralt, M., Etayo, J., and GOmez-Bolea, A. 2000. Amandinea
B. prospersa used for lectotypefication. This study was sup- crassiuscula, a new corticolous species from the Iberian Penin-
ported by a Sigma Xi Grant-in-Aid of Research, sponsored sula. Lichenologist, 32: 521-529.
by the local Sigma Xi chapter at ASU, and two National Greuter, W., McNeill, J., Barrie, ER., Burdet, H.-M., Demoulin,
Science Foundation grants (DEB-0103738, DEB-9701111). V., Filgueiras, T.S., Nicolson, D.H., Silva, P.C., Skog, J.E.,
Trehane, P., Turland, N.J., and Hawksworth, D.L. (Editors).
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