The Bryologist 107(4), pp. 459 479
Copyright q 2004 by the American Bryological and Lichenological Society, Inc.
The Genus Buellia Sensu Lato in the Greater Sonoran Desert Region: Saxicolous
Species with One-septate Ascospores Containing Xanthones
FRANK BUNGARTZ
Botanische Staatssammlung München, Menzinger Straße 67, D-80638 München, Germany; e-mail: bungartz@
bsm.mwn.de
JOHN A. ELIX
Department of Chemistry, The Faculties, Australian National University, Canberra, ACT 0200, Australia; e-mail:
john.elix@anu.edu.au
THOMAS H. NASH III
Arizona State University Lichen Herbarium, School of Life Sciences, P.O. Box 87 4601, Tempe, AZ 85287-4601,
U.S.A. e-mail: tom.nash@asu.edu
Abstract. Six species of saxicolous Buellia s.l. containing xanthones are reported from the
Greater Sonoran Desert Region: Buellia concinna, B. halonia, B. mamillana, B. prospersa, B.
subaethalea, and B. trachyspora. All species have a pale yellow to greenish yellow thallus characterized by the presence of xanthones. Nevertheless, they all show distinct characters and may
consequently only be distantly related. Buellia concinna, a taxon previously reported only from
Europe, is the valid name for the North American B. semitensis. Buellia subaethalea, first described by Bouly de Lesdain, but subsequently largely ignored, is reported for the first time from
the region. Buellia trachyspora, a subtropical species barely reaching the Sonoran Region, is
distinguished from B. mamillana.
Keywords. Buellia, lichen taxonomy, lichenized ascomycetes, North America, saxicolous species, Sonoran Desert, xanthones.
Lichen species with xanthones can usually easily
be recognized by their yellowish thallus color, frequently have a C1 conspicuously orange thallus
reaction and usually fluoresce yellow or orange in
UV light. Xanthones are lichen metabolites derived
from acetyl CoA via the acetyl polymalonyl pathway. Unlike many other lichen substances, they are
difficult to distinguish with routine thin-layer chromatography (TLC). High performance liquid chromatography (HPLC) is therefore the preferred
method of identification. In Europe, Scheidegger
and Ruef (1988) pioneered analyzing xanthones
from saxicolous species of Buellia. Xanthones are
not common in Buellia and it is currently difficult
to evaluate whether species containing xanthones
belong to Buellia s.str. All species from the Sonoran Region are well distinguished from one another by distinct morphological, anatomical, and ultrastructural characters. Even though some of these
species may be closely related, it is unlikely that
all species with xanthones share a common ancestry. Segregating these species from Buellia s.l. does
not resolve the taxonomy of the species, and it is
currently not advisable to erect new genera that are
monotypic or contain only a few species. All spe-
cies are therefore included in Buellia s.l. and detailed descriptions are provided including morphological, anatomical, ultrastructural, and chemical
characters.
MATERIAL
AND
METHODS
All specimens were examined with light microscopy
using hand- and cryosections. Both conventional bright
field microscopy (BF) and differential interference contrast (DIC) were used. Selected specimens were also studied with transmission electron microscopy (TEM) according to a protocol described in detail by Bungartz et al.
(2002). To improve dehydration and infiltration, this protocol has been modified according to Bungartz and Nash
(2004b).
All specimens were spot tested and routinely examined
with standardized thin-layer chromatography (Culberson
& Johnson 1982; Culberson & Kristinsson 1970; Orange
et al. 2001; White & James 1985). If norstictic acid was
present, clusters of orange, needle-shaped crystals were
formed in KOH. These are best observed in the compound
microscope. TLC-plates were interpreted with the computer program WINTABOLITES (Mietzsch et al. 1994),
and scanned for permanent record (Egan 2001). In addition, a subset of specimens were analyzed using standardized high performance liquid chromatography (HPLC,
Elix et al. 2003). All specimens examined here fluoresce
in UV-light, even though some reactions may be pale
(possibly caused by a low concentration of xanthones).
0007-2745/04/$2.25/0
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THE BRYOLOGIST
Thallus medulla, cortex, and transverse sections of the
apothecia were tested with Lugols iodine. A highly concentrated or even saturated stock solution (Common 1991)
was used if reactions were tested directly on the specimen.
When testing the medulla, caution was applied that reactions were not confused with black hyphae from a hypothallus below. If in doubt, thallus transverse sections
were carefully re-examined with the compound microscope. To observe sufficient detail within the ascus apex,
apothecium transverse sections were tested with a 0.3%
IKI solution after a pre-treatment with 5% KOH. Ascus
type terminology follows Rambold et al. (1994).
The thalli frequently contain crystals of various origins.
A simple test with H2SO4 can be indicative for the presence of Ca oxalates (Bungartz & Nash 2004b), but for all
specimens examined here this reaction was negative and
no clusters of needle-shaped crystals were observed.
Terminology referring to spore types is currently in a
state of flux and it appears problematic to adopt a single
concept. Structurally identical types within Buellia s.l. and
Rinodina s.l. have probably evolved independently
(Helms 2003), and it may thus be problematic to refer to
these types with the same name. Spores measurements are
given according to Nordin (2000). Spore ornamentation
follows the terminology of Scheidegger (1993). The term
‘‘psilate’’ has not been used. Using a technical term like
‘‘psilate’’ implies that a particular ornamentation should
be present. Psilate spores are, however, smooth and thus
lack distinct ornamentation. Buellia trachyspora has
spores with coarse ornamentation not previously described
from Buellia. It is described here as coarsely rugulate or
‘areolate’.
Scheidegger (1993) described several distinct exciple
types, but his concepts have not generally been adopted
because the distinction of the different types relies on
cumbersome examination of the excipular plectenchyma.
Only thin microtome sections (,15 mm) allow this differentiation to be observed. To clarify Scheidegger’s
(1993) concept, line drawings of all types are provided
here (Figs. 10–11). Structural variation and differences in
pigmentation are mentioned within the descriptions. References to the fungal plectenchyma follow Korf (1973),
van Brummelen (1967), and Ryan et al. (2001). Pigment
names follow Meyer and Printzen (2000). Their protocol
to identify acetone insoluble pigments according to color
reactions with HNO3, KOH, and HCl was applied. Most
pigments were thus identified. Some color reactions can
be misleading because pigments do not occur as pure colors, but are ‘‘superimposed’’ i.e., mixed in different concentrations. In addition, colorless lichen substances like
atranorin and norstictic acid react with KOH and further
obscure the observations. We are reasonably confident that
the most commonly present brown pigment is elachistabrown, even though the reactions are less obvious than
described by Meyer and Printzen (2000).
The terminology of Vobis (1980) and Vobis and Hawksworth (1981) has been adopted to describe conidiophore
types and pycnidium ontogeny. For a detailed discussion
of the relevance of conidia and pycnidia as taxonomic
characters in Buellia s.l., see Bungartz et al. (2004).
Detailed specimen information about all collections
at ASU is available at: http//seinet.asu.edu/collections/
selection.jsp.
BUELLIA CONCINNA Th. Fr., Nova Acta Reg. Soc.
Scient. Upsal. Ser. 3, 3: 332. 1860.
TYPE: NORWAY. FINMARK. Varanger, Nesseby, 30 August 1857, Fries s.n. (UPS, lectotype designated by Foucard et al. 2002; BM, GZU, M!, O, PC, isolectotypes).
[VOL. 107
Buellia semitensis Tuck., Syn. N. Amer. Lich. 2: 95. 1888.
TYPE: U.S.A. CALIFORNIA. Mariposa Co. Granitic rocks,
Bolander 329 ex Tuckerman sheet 3281 (FH!, lectotype
selected here). Note: The lectotypification is necessary
because Tuckerman (1888, p. 96) did not refer to a
particular specimen in the protologue: ‘‘Granitic rocks,
Yosemite Valley and elsewhere, California, Bolander:’’
For additional synonyms see Scheidegger and Ruef
(1988) and Foucard et al. (2002).
Thallus (Figs. 1A–B) crustose, thin to moderately thickened, closely aggregated or dispersed, epilithic; granular to minutely bullate; prothallus absent; thallus surface smooth and shiny, rarely matt,
usually pale yellow to greenish yellow, rarely becoming pale yellow (older herbarium specimens),
epruinose, phenocorticate; medulla lacking Ca-oxalate (H2SO42). Apothecia lecideine; (0.2–)0.3–
0.6(–0.8) mm in diameter; soon sessile; proper margin prominent, usually persistent, rarely excluded
with age, black; disc black, epruinose or rarely with
faint, yellow pruina, plane, rarely becoming slightly
convex with age; exciple distinct, of leptocline-type
(Figs. 1C, 10D) i.e., exciple thick, distinct, and not
distinctly differentiated into inner and outer part,
hyphae thin-walled, prosoplectenchymatous and
usually 6 densely interwoven (textura intricata),
dull fuscous brown throughout, becoming 6 carbonized by various amounts of brown pigments (cf.
elachista-brown), pigmentation continuous with
epihymenium, hypothecium deep reddish brown
(leptoclinoides-brown, textura intricata); hymenium hyaline, not inspersed; paraphyses simple to
moderately branched, apically swollen, with brown
(cf. elachista-brown) pigment cap and diffuse,
brown pigment (HNO31 violet, cinereorufa-green).
Asci 8-spored, clavate, Bacidia-type. Ascospores
(Figs. 1D–I) soon becoming pigmented, initially olive, brown at maturity, ellipsoid to 6 crescentshaped i.e., often distinctly curved and characteristically with tapered ends, rarely constricted,
(13.0–)15.6–[18.5]–21.3(–24.0) 3 (6.0–)7.9–[8.9]–
9.9(–11.0) mm (n 5 60); one-septate (very rarely
with additional median septa forming at both ends),
proper septum becomes thickened early during
spore ontogeny (Physconia-type; Figs. 1D–E); ornamentation microrugulate, becoming rugulate with
age (best seen in DIC); septum with septal pore
canal, simple pore, and undifferentiated pore plug;
spore wall (Figs. 1G–I) differentiated into cracked,
thin perispore (0.11–0.17 mm), narrow intermediate
layer (,0.03 mm), thick proper spore wall (0.37–
0.50 mm) and moderately thickened endospore
(0.24–0.34 mm). Pycnidia rare, urceolate to globose, unilocular, at maturity almost entirely occupied by densely branched conidiophores; conidiogenous cells mostly terminal, rarely also intercalary (cf. conidiophore-type V); pycnidial ontogeny
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BUNGARTZ ET AL.: SAXICOLOUS BUELLIA
461
FIGURE 1. Buellia concinna (A–F. light micrographs; G–I. TEM micrographs). — A. Granular thallus with sessile
apothecia (Nash 22836). — B. Close-up of granules with apothecia (Nash 22836). — C. Transverse section of the
leptocline-type exciple i.e., largely undifferentiated throughout and composed of intricately interwoven, mesodermatous
hyphae (Nash 22836). — D. Mature ascospores with 6 tapered ends and thickened septum (arrow; Nash 22836). —
E. Premature, ellipsoid pale ascospore with distinct septum thickening and mature, crescent-shaped ascospore with
deeply pigmented wall and moderately thickened septum (Nash 11791). — F. Overmature, 6 citriform ascospore (Nash
22836). — G. Premature ascospore (Nash 14655): (as) ascus wall; (s) mucilaginous sheath; (1) perispore; (2) intermediate layer; (3) proper spore wall; (4) endospore (at this stage of the ontogeny the proper spore wall and the endospore
are not yet fully differentiated). — H. Mature ascospore (Nash 14655). — I. Wall layers in a mature ascospore (Nash
14655): (as) ascus wall; (s) mucilaginous sheath; (1) perispore; (2) intermediate layer; (3) proper spore wall; (4)
endospore (at this stage of the ontogeny the proper spore wall and the endospore are fully differentiated).
462
THE BRYOLOGIST
similar to Umbilicaria-type; conidia simple, bacilliform to ellipsoid, 2.0–4.0 3 1.5–2.0 mm (n 5 20).
Chemistry (Fig. 7). Spot test reactions usually
C1 orange or sometimes 6 pinkish, K1 yellow,
KC1 orange, P1 orange, CK1 orange. UV1 pale
or bright yellow to orange. The thallus medulla reacts weakly amyloid or not amyloid. The hymenium is also amyloid. The thallus is characterized by
the following xanthones: arthothelin, isoarthothelin,
6-O-methylarthothelin, 4,5-dichloronorlichexanthone, 4,5-dichloro-6-O-methylnorlichexanthone,
asemone and thiophanic acid. In addition, gyrophoric, lecanoric and orsellinic acid are often present.
Substrate and ecology. On a variety of hard,
siliceous (HCl2) rock substrates, often on vertical
cliffs. Scheidegger and Ruef (1988) reported that
this species often grows as a juvenile parasite on
crustose and rarely foliose lichens. The North
American specimens are not lichenicolous, but otherwise match the description of B. concinna in
Scheidegger and Ruef (1988), Scheidegger (1993),
and Foucard et al. (2002). Isolectotype material of
B. concinna, examined during a visit to M, as well
as a specimen from Scheidegger’s private herbarium are not lichenicolous and agree well with the
North American material.
Distribution (Fig. 6). Widely distributed
throughout the Northern Hemisphere. Within the
Sonoran Desert Region only moderately common
at higher elevations (montane to subalpine).
Notes. Buellia concinna is easily recognized by
its distinct granular thallus and the characteristic,
crescent-shaped spores. This unusual spore shape is
not present in all spores of an apothecium, but can
usually be found in at least some of the sections
examined from an apothecium. No other species
from the Sonoran Region has similar spores.
Selected specimens examined.—AUSTRALASIA.
NEW ZEALAND. Nash 19094 (ASU).—EUROPE.
FRANCE. PYRÉNÉES-ORIENTALES. Scheidegger 7840 (hb.
Scheidegger). NORWAY. FINMARK. Fries s.n. (MSC69240).—NORTH AMERICA. MEXICO. HIDALGO. Nash
38098 (ASU). SONORA. Nash 12483 (ASU). U.S.A. ARIZONA.
Apache Co. Nash 27088 (ASU), Hertel 40189 (M). Cochise
Co. Nash 14596 (ASU); Nordin 5178 (UPS). Coconino Co.
Nash 7575 (ASU). Santa Cruz Co. Nash 18578 (ASU). CALIFORNIA. El Dorado Co. Ryan 23591a (ASU). Fresno Co.
Thiers 34410 (ASU). Lake Co. Sigal s.n. (ASU). Los Angeles Co. Ryan 26137b (ASU). San Diego Co. Ryan
25821b (ASU). Shasta Co. Nash 22836 (ASU). Tulare Co.
Nash 11287 (ASU). Tuolumne Co. Ryan 24231 (ASU). COLORADO. Clear Creek Co. Anderson 6673 (ASU). El Paso
Co. Anderson 8293 (ASU). SOUTH DAKOTA. Custer Co. Anderson 7879 (ASU). Unknown Co. Anderson s.n. (MSC63646).
BUELLIA
1866.
[VOL. 107
HALONIA
(Ach.) Tuck., Lich. Californ. 26.
Lecidea halonia Ach., Meth. Lich. 47. 1803. TYPE:
SOUTH AFRICA. CAPE PROVINCE. Cape of Good
Hope. On hard maritime rock [original label data: Caput bonae spei, in saxis durissimis ad littora Africae
australioris], Osbeck s.n. (H-Ach–362!, lectotype selected here; several isolectotypes, UPS-ACH, isolectotype); ex herb. Acharius (H-Nyl 9211, possibly a type
fragment).
Baeomyces capensis Taylor in Hook., London Journ. Bot.
6: 186. 1847. TYPE: SOUTH AFRICA. WESTERN CAPE.
Cape of Good Hope. ex hb. Hooker sub. ‘‘9311 Lecidea’’ in Taylor herb. sheet 1460 (FH, lectotype selected
here; BM-000660172, BM-000660173, isolectotypes).
Lecidea disciformis var. halonia (Ach.) Nyl., Mém. Soc.
Imp. Sci. Natur. Cherbourg 5: 126. 1857.
Buellia disciformis var. halonia (Ach.) Boist., Nouv. Flore
Lich. 2: 235. 1903.
Diploicia capensis (Taylor) Dodge, Beih. Nov. Hedwigia
38: 157. 1971.
Possible synonyms:
Buellia flavoareolata (Nyl.) Müll. Arg., Hedwigia 31: 283.
1892.
Lecidea flaveoareolata Nyl., Annal. Scienc. Nat. Bot., ser.
4, 3: 166. 1855. TYPE: CHILE. QUEBRADA. Quilmenco,
318459 S, 718059 W. (H-Nyl, syntype). Note: Specimens
from Chile (Coquimbo Province) distributed by Weber
Lich. Exs. 564a, b (ASU, SBBG), and material collected
by Imshaug on an Expedition to Santa Clara (Juan Fernandez Islands, Chile; MSC-0005485, MSC-0005486,
MSC-0005487, MSC-0005488) are morphologically and
anatomically similar, if not identical to B. halonia.
These specimens were analyzed with TLC by R. C.
Harris, who detected atranorin and an unknown xanthone. Imshaug annotated several other specimens (see
specimens examined) from San Juan Fernandez (Chile)
as B. halonia. These specimens were not examined by
TLC or HPLC, but agree well with B. halonia. The type
of B. flavoareolata has not been studied, but it is very
likely that it is a synonym of B. halonia.
Thallus (Figs. 2A–B) crustose, thick, 6 continuous, epilithic; areolate; prothallus distinct, delimiting thallus as black outline; thallus surface smooth
and matt to 6 shiny, usually yellowish green to pale
yellow, rarely becoming pale yellow (older herbarium specimens), epruinose, phenocorticate; medulla
lacking Ca-oxalate (H2SO42). Apothecia lecideine;
(0.2–)0.3–0.5(–0.7) mm in diameter; initially immersed, young apothecia frequently 6 aspicillioid,
soon bursting through thallus surface and becoming
adnate to sessile; proper margin prominent, usually
persistent, rarely excluded with age, black, frequently with coarse thallus fragments attached; disc
black, epruinose, sometimes with yellowish pruina,
plane, rarely becoming strongly convex with age;
exciple distinct, of leptocline-type (Figs. 2C–D,
10D) i.e., exciple thick, distinct, and not distinctly
differentiated into inner and outer parts, hyphae
thin-walled, prosoplectenchymatous, and usually 6
densely interwoven (textura intricata), dull fuscous
brown to aeruginose throughout, becoming 6 car-
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BUNGARTZ ET AL.: SAXICOLOUS BUELLIA
463
FIGURE 2. Buellia halonia (Nash 41434; A–F. light micrographs; G–L. TEM micrographs). — A. Areolate thallus
with immersed sessile apothecia. — B. Close-up of areoles and apothecia: immature apothecia are immersed and large
chunks of thalline material (arrows) often remain attached during emergence. — C. Transverse section of the leptoclinetype exciple. — D. Detail of outer exciple: mesodermatous hyphae are not considerably swollen at apices and 6
interwoven throughout exciple. — E. Ascus with eight mature ascospores (arrow indicates septum thickening). — F.
Mature ascospore with distinct septum thickening (arrow). — G. Immature ascospore. — H. Wall layers in an immature
ascospore: (as) ascus wall; (1) outer layer (perispore and mucilaginous sheath not distinctly differentiated); (2) inner
layer (proper wall and endospore not distinctly differentiated). — I. Mature ascospore: septum thickening of endospore
is not yet developed. — J. Pycnidium. — K. Conidiophores with bacilliform conidia. — L. Wall layers in mature
ascospore: (as) ascus wall; (s) mucilaginous sheath; (1) perispore; (2) intermediate layer; (3) proper spore wall; (4)
endospore.
464
THE BRYOLOGIST
bonized by various amounts of brown and aeruginose pigments (cf. elachista-brown & cinereorufagreen), pigmentation continuous with epihymenium, but aeruginose pigment not extending into
deep reddish brown hypothecium (leptoclinoidesbrown, textura intricata); hymenium hyaline, not
inspersed; paraphyses simple to moderately
branched, apically swollen, with brown pigment
cap (cf. elachista-brown) and diffuse, aeruginose
pigment (HNO31 violet, cinereorufa-green). Asci
8-spored, clavate, Bacidia-type. Ascospores (Figs.
2E–I, L) soon becoming pigmented, initially olive,
brown at maturity, oblong to ellipsoid, usually not
constricted, with obtuse ends, not curved, (11.5–)
12.4–[14.1]–15.7(–19.0) 3 (6.0–)6.8–[7.4]–8.0(–
9.0) mm (n 5 60); one-septate, proper septum soon
but only briefly thickened during ontogeny (Physconia-type; Figs. 2E–F); ornamentation weakly microrugulate (in DIC); septum with septal pore canal, simple pore, and undifferentiated pore plug;
spore wall (Figs. 2H, L) differentiated into smooth,
thin perispore (0.14–0.20 mm), narrow intermediate
layer (, 0.05 mm), thick proper spore wall (0.35–
0.44 mm), and moderately thickened endospore
(0.18–0.34 mm). Pycnidia (Fig. 2J) rare, urceolate
to globose, unilocular, at maturity almost entirely
occupied by densely branched conidiophores; conidiogeneous cells mostly terminal, rarely also intercalary (cf. conidiophore-type V, Fig. 2K); pycnidial
ontogeny similar to Umbilicaria-type; conidia simple, bacilliform, 5.0–7.0 3 1.0–1.5 mm (n 5 20).
Chemistry (Fig. 7). Thalli typically react K1
yellow, P1 orange, C1 orange, KC1 orange, and
CK1 orange. The thallus fluorescence is typically
UV1 bright yellow to orange. The thallus is not
amyloid, but the hymenium reacts amyloid. The
following xanthones were detected with HPLC: arthothelin, isoarthothelin, 2,4-dichloronorlichexanthone, 2,5-dichloronorlichexanthone, 2,7-dichloronorlichexanthone, 4,5-dichloronorlichexanthone,
5,7-dichloronorlichexanthone, thiophanic, and thiophaninic acid. Two bis-xanthones, eumitrin X and
eumitrin Y, have been detected with HPLC in part
of the specimens. Their chemical structure is currently unresolved. Atranorin is regularly present,
frequently accompanied by norstictic acid.
Substrate and ecology. Growing abundantly on
maritime siliceous mineral-poor coastal rock (generally HCl2).
Distribution (Fig. 6). Currently known from
the Pacific coast of North America, South Africa,
Australia, and Chile (see notes on B. flaveoareolata, which is possibly a synonym of B. halonia).
Reports of B. halonia from Europe belong to B.
concinna according to Scheidegger and Ruef
(1988). In the Sonoran Region, very common along
[VOL. 107
the coast of southern California and Baja California.
Notes. Buellia halonia has frequently been confused with Lecidella asema (Nyl.) Knoph & Hertel,
a crustose lichen with similar color, often growing
in the same habitat. In the field, granulose thalli of
L. asema can usually be distinguished from areolate
thalli of B. halonia. This distinctly different morphology is often sufficient for identification, but the
two species are more reliably distinguished by microscopic examination: Lecidella asema has simple,
hyaline spores. Spores of Buellia halonia are brown
and one-septate.
Selected specimens examined.—AUSTRALASIA.
AUSTRALIA. SOUTH AUSTRALIA. Streimann 54946 (MSC354293).—AFRICA. SOUTH AFRICA. CAPE PROVINCE. Eaton s.n. (BM, H-Nyl 2210); unknown collector (H-Nyl
9212).—NORTH AMERICA. MEXICO. BAJA CALIFORNIA. Nash 4910, 8719, 25186, Ryan 21332 (ASU); Kalb
24668a (hb. Kalb); Weber L-36628 (COLO); Wetmore
63671 (MIN). BAJA CALIFORNIA SUR. Nash 40017 (ASU).
U.S.A. CALIFORNIA. Lake Co. Toren s.n. (SFSU), Hasse s.n.
(ASU). Los Angeles Co. Marsh 6500, Nash 32020, Ryan
30942 (ASU); Weber L-42957 (COLO). San Diego Co. Bratt
5628 (SBBG). San Luis Obispo Co. Nash 36976, Riefner
88-431 (ASU). San Mateo Co. Shushan S14688 (ASU). Santa Barbara Co. Nash 32573, 33102, Ryan 31184, Weber
Lich. Exs. 680 (ASU); Weber L-80190 (COLO). Ventura Co.
Nash 37035 (ASU); Weber S1734 (COLO).—SOUTH
AMERICA. CHILE. JUAN FERNANDEZ. Imshaug 37584
(MSC-113121), 38241 (MSC-113136), 38236 (MSC-113137).
BUELLIA MAMILLANA (Tuck.) W. A. Weber, Mycotaxon 27: 493. 1986.
Rinodina mamillana Tuck., Proc. Amer. Acad. 7: 226.
1866 (1868). TYPE: U.S.A. HAWAII. Honolulu. Oahu.
Volcanic Rocks, Mann s.n. ex sheet no. 2124a (FH, holotype; W!, isotype).
Lecidea glaziouana Kremp., Flora 59: 317. 1876. TYPE:
BRAZIL. RIO DE JANEIRO. On granitic rocks [original
label data: ad saxa granitica], Glaziou 3506 (M0023763!, lectotype selected here; M-0023765!, BM000660175!, C!, G! W!, isolectotypes).
Lecidea recepta Kremp., Flora 59: 318. 1876. TYPE: BRAZIL. RIO DE JANEIRO. On granitic rock. [original label
data: ad saxa granitica], 1869, Glaziou 3295 (M0023811!, lectotype selected here; M-0023810!, isolectotype).
Buellia glaziouana (Kremp.) Müll. Arg., Flora 63: 19.
1880.
Buellia recepta (Kremp.) Müll. Arg., Linnaea 43: 36.
1880.
Rinodina thomae Tuck., Syn. N. Americ. Lich. 1: 209.
1882. TYPE: U.S.A. ALABAMA. Lawrence Co. Sandstone
Rocks at Moulton, 348289 N 878179 W, ca. 190 m, Peters s.n. (5 ex Tuckerman sheet no. 2125) (FH, holotype; MICH, isotype).
Rinodina contiguella Vain., Étude Lich. Brésil 1: 164.
1890. TYPE: BRAZIL. RIO DE JANEIRO. On granitic rock
[original label data: ad saxa granitica], Glaziou 3506
(M!, lectotype selected here).
Lecidea cohibilis Nyl., Sertum ich. Trop. Labuan et Singapore 41. 1891. TYPE: U.S.A. TENNESSEE. At Lookout
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BUNGARTZ ET AL.: SAXICOLOUS BUELLIA
Tower mountain, on sandstone, Calkins s.n. (H-Nyl, holotype; F, FH, MICH, MO, NY, US, isotypes).
Buellia contiguella (Vain.) Malme, Bih. K. Svenska Vet.Akad. Handl. 28: 5. 1902.
Buellia moreliiensis de Lesd., Lich. Mex. 26. 1914. TYPE:
MEXICO. MICHOACÁN. Morelia. On volcanic rocks
(US!, lectotype selected here). Note: The material from
US is selected here as the lectotype because the holotype
material was probably destroyed in World War II. Imshaug (1955) reported that he had seen type material
from Magnusson’s private herbarium.
Buellia poliocheila Vain., Ann. Acad, Sci. Fenn. ser. A
6(7): 85. 1915. TYPE: U.S.A. VIRGIN ISLANDS. St. Thomas. On rocks at Ma Folie, Biese s.n. (C, holotype; TUR,
isotype).
Melanaspicilia contiguella (Vain.) Vain., Ann. Acad. Sci.
Fenn. ser. A 6: 79. 1915.
Buellia sensitiva Zahlbr., Mycologia 22: 78. 1930. TYPE:
U.S.A. PUERTO RICO. Naranjito. In an open field, 25
November 1915, Fink 102 (MICH!, holotype).
Buellia cohibilis (Nyl.) Zahlbr., Cat. lich. univ. 7: 344.
1931.
Buellia antillarum de Lesd., Bull. Museum Hist. Nat. Paris II 15: 470. 1934. TYPE: FRENCH WEST INDIES.
GUADELOUPE ISLAND. Cove south of Bouillante. On volcanic rock at sea level. [original label data: Anse du
Sud de Bouillante] collector not known (PC, holotype).
Buellia thomae (Tuck.) Imshaug comb. ined., in Imshaug
University Microfilms no. 2607. 1951.
Buellia glaziouana var. poliocheila (Vain.) Imshaug, Farlowia 4: 493. 1955.
Buellia glaziouana var. sensitiva (Zahlbr.) Imshaug, Farlowia 4: 494. 1955.
Possible synonym:
Buellia subglaziouana S.R. Singh & D.D. Awasthi, Biological Memoirs 6: 190. 1981. TYPE: INDIA. MADHYA
PRADESH. Hoshangabad district, Pachmarhi, on way to
Chhota Mahadeo, altitude ca 1,050 m, 26 January 1973,
Singh 73.157 (AWAS, holotype). Note: No type material
of B. subglaziouana has been studied. Singh and Awasthi (1981) described B. subglaziouana as different from
B. glaziouana because of a deep iodine-blue medulla
reaction. However, contrary to the protologue, B. glaziouana also has a distinctly I1 blue medulla. Buellia
subglaziouana could therefore be considered a synonym of B. mamillana (5 B. glaziouana).
Thallus (Figs. 3A–B) crustose, thin to moderately thick, 6 continuous, epilithic; rimose, rarely becoming rimose-areolate; prothallus distinct, delimiting thallus as black outline; thallus surface smooth
and matt, rarely 6 shiny, usually pale greenish yellow, rarely gray (older herbarium specimens),
epruinose, phenocorticate; medulla lacking Ca-oxalate (H2SO42). Apothecia of mamillana-type
(Figs. 3C–D, 11A–B) i.e., initially lecanorine, but
soon becoming lecideine; (0.3–)0.6–0.9(–1.1) mm
in diameter; usually adnate to sessile, rarely remaining immersed; proper margin thickening with
maturity, inconspicuous in immature apothecia, but
soon becoming distinct, brownish black; disc dark
brown to blackened, epruinose, plane, usually 6
convex with age; exciple of immature apothecia
thin, indistinct but soon thickening i.e., inner ex-
465
cipular hyphae distinct, hyaline but becoming pigmented with age, prosoplectenchymatous (textura
angularis), distinct, not reduced, similar in structure
and orientation to deep reddish brown hypothecium
(leptoclinoides-brown, textura intricata), outer excipular hyphae short-celled, cells angular, 6 swollen (textura angularis), and usually 6 carbonized
with various amounts of brown pigments (cf. elachista-brown), pigmentation continuous with epihymenium; hymenium hyaline, not inspersed; paraphyses simple to moderately branched, apically swollen, with brown (cf. elachista-brown) pigment cap.
Asci 8-spored, clavate, Bacidia-type. Ascospores
(Figs. 3E–G) soon becoming pigmented, initially
olive, brown at maturity, oblong to ellipsoid, usually not constricted, with obtuse ends, not curved,
(10.5–)12.0–[13.8]–15.7(–18.5) 3 (5.5–)6.3–[7.0]–
7.8(–10.0) mm (n 5 60); one-septate, proper septum
soon but only briefly thickened during spore ontogeny (6 Physconia-type); ornamentation rugulate;
septum with septal pore canal, simple pore, and undifferentiated pore plug; spore wall (Fig. 3G) differentiated into smooth to cracked, thick perispore
(0.32–0.43 mm), indistinct intermediate layer
(,0.01 mm), thin proper spore wall (0.09–0.16 mm)
and moderately thickened endospore (0.17–0.28
mm). Pycnidia rare, urceolate to globose, unilocular, at maturity almost entirely occupied by densely
branched conidiophores; conidiogeneous cells
mostly terminal, rarely also intercalary (cf. conidiophore-type V); pycnidial ontogeny similar to Umbilicaria-type; conidia simple, fusiform, 6.0–14.0
3 1.0–1.5 mm (n 5 20).
Chemistry (Fig. 8). Thalli usually react K1
yellow, P2 or P1 yellow, C2, KC2, CK2, but
these spot tests are sometimes weak. Thalli generally fluoresce UV1 bright or pale yellow to orange.
Both thallus medulla and apothecia react distinctly
amyloid, even though Vainio (1915) and Malme
(1902) reported a negative iodine reaction. Imshaug
(1955) first noticed this reaction and Weber (1986)
confirmed his observation. The following xanthones where detected with HPLC: 4,5-dichloro-3O-methylnorlichexanthone, 4,5-dichlorolichexanthone, 4-chlorolichexanthone. Atranorin is typically
also present, frequently accompanied by haematommic acid and methyl b-orsellinate (both artifacts
formed by hydrolysis of atranorin). 29-O-methylperlatolic, divaricatic, norstictic, hypostictic, stictic,
methylstictic, cryptostictic acid, and one unknown
trace substance have occasionally also been detected by HPLC.
Substrate and ecology. On shaded, steep, 6
vertical, siliceous rock faces (HCl2).
Distribution (Fig. 6). Known from tropical to
subtropical localities throughout the world (Mexi-
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FIGURE 3. Buellia mamillana (A–E. light micrographs, F–G. TEM micrographs). — A. Rimose thallus with immersed to 6 sessile apothecia (Nash 28439). — B. Close-up thallus and apothecia (Nash 25875; arrows indicate thalline
part of margin in some premature apothecia). — C. Transverse section of immature apothecium (Nash 25604) with
distinct thalline outer exciple [ca 60 mm wide; (c) cortex; (a) algae], and a narrow (ca 10 mm wide) inner exciple
(arrows). — D. Transverse section of premature apothecium (Nash 25604): proper exciple (arrows) is expanding (ca
30 mm wide) while the outer, thalline exciple (th) becomes excluded (ca 35 mm wide). — E. Mature ascospores with
rugulate ornamentation (Nash 25604). — F. Mature ascospore (Nash 37836). — G. Wall layers in mature ascospore
(Nash 37836): (as) ascus wall; (s) mucilaginous sheath; (1) perispore; (2) intermediate layer; (3) proper spore wall; (4)
endospore.
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BUNGARTZ ET AL.: SAXICOLOUS BUELLIA
co: Bouly de Lesdain 1914, Caribbean: Imshaug
1955, 1957, Brazil: Malme 1902, India: Singh &
Awasthi 1981, southeastern USA: Tuckerman
1888). In the Sonoran Region, a montane subtropical species, especially common in the Sierra Madre
Occidental and Southern Baja California.
Notes. Based on the development of the apothecia, Imshaug (1955) distinguished three lines of
development in B. mamillana and recognized two
of these lines as varieties: 1) with apothecia remaining immersed, 2) with apothecial initials frequently abortive (var. poliocheila), and 3) with a
depsidone and a K1 red spot test reaction, forming
crystals (var. sensitiva). This range of variation can
be explained if apothecial anatomy is examined
carefully. All apothecia are initially immersed, but
eventually emerge from the thallus with a distinct
thalline margin. Immature apothecia do not appear
abortive because spores are frequently formed even
in young apothecia that are not yet enclosed by a
distinctly colored envelope.
The considerable variation in thallus chemistry,
as confirmed by HPLC, would suggest that Imshaug’s varieties might correlate with distinct
chemotypes and if so, these varieties would deserve
separate taxonomic recognition. Material examined
here could, however, not be segregated into distinct
chemotypes. It was not possible to correlate the
chemical variation with any of the ‘‘developmental
lines’’ described by Imshaug (1955), therefore no
distinct varieties are recognized here.
Selected specimens examined.—CENTRAL AMERICA. BRITISH ANTILLES. ST. LUCIA. Imshaug 30160,
30160 (MSC-347926, MSC-347926). COSTA RICA. Pittier
s.n. (BM-000660174, BM-000671861). CUBA. LA HABANA.
Imshaug 24815 (MSC-75987). DOMINICAN REPUBLIC.
DE LA VEGA. Imshaug 23738 (MSC-76115); Wetmore
23738 (MSC-76115). ECUADOR. GALAPAGOS ISLANDS.
Weber Lich. Exs. 110 (M-0061342); Hill s.n., ex sheet no.
2124a (FH). FRENCH ANTILLES. GUADELOUPE. LeGallo
2707 (MSC-60972). SAINT-BARTHÉLEMY. LeGallo 2665
(MSC-61050), 2655 (MSC-61038). JAMAICA. ST. ANDREW
PARISH. Imshaug 14454, 13808 (MSC-76027, MSC-76062).
SAINT VINCENT & THE GRENADINES. SAINT VINCENT . Imshaug 30568, 30568 ( MSC-347923, MSC347923).—NORTH AMERICA. MEXICO. BAJA CALIFORNIA. Ryan 21461 (ASU). BAJA CALIFORNIA SUR. Nash
29793, 33778, 16953 (ASU). CHIHUAHUA. Nash 31295
(ASU). NAYARIT. Nash 39192 (ASU). SINALOA. Nash 10086,
12214 (ASU). SONORA. Marsh 5918, Nash 25518, 25524,
Ryan 21625, 21636 (ASU); Wetmore 69963 (COLO). U.S.A.
ARIZONA. Cochise Co. Nash 9653b, Ryan 10704 (ASU).
Gila Co. Nash 28439. Pima Co. Nash 7984 (ASU). Santa
Cruz Co. Nash 25875 (ASU). ARKANSAS. Garland Co. Wetmore 86251 (MSC-379125). HAWAIIAN ISLANDS. Kauai.
Nash 20829 (ASU); Hawaii. Bailey 353 (DUKE). LOUISIANA.
Natchitoches Co. Tucker 17523 (M-0061343). TENNESSEE.
Hamilton Co. Calkins 224, 236 (MSC-146694, MSC120013). TEXAS. Nacogdoches Co. Wetmore 17638
(MIN).—SOUTH AMERICA. ARGENTINA. TUCUMAN.
Lamb 5164 (MSC-117950). BRAZIL. RIO DE JANEIRO. Vai-
467
nio Lich. Bras. Exs. 74 (M-0061345, BM-000671862); Glaziou s.n. (MSC-122653); Lamb 5014 (MSC-338305). SÃO
PAULO. Kalb Lich. Neotrop. Exs. 46 (M-0061341); Lamb
5013 (MSC-117955). CHILE. ISLA DE PASCUA (5 EASTER
ISLAND). Skottsberg s.n. (BM-000671861). PARAGUAY.
DEPARTMENTO DE PARAGUARI. Malme 1486B (MSC129597).
BUELLIA PROSPERSA (Nyl.) Riddle, Brookl. Bot.
Gardens Memoirs 1: 114. 1918.
Recent synonyms.—Rinodina lecideina Mayrhofer &
Poelt, Bibl. Lich. 12: 112. 1979. Amandinea lecideina
(H. Mayrhofer & Poelt) Scheid. & H. Mayrhofer, in
Scheidegger, Lichenologist 25: 342. 1993.
For a detailed description see Bungartz et al. (2004).
BUELLIA SUBAETHALEA de Lesd., Lichens du Mexique 27. 1914.
TYPE: MEXICO. MICHOACÁN. Morelia, Santa Maria
mountain ridge [original label data: Loma Santa Maria]
2,000 m, Frère Arsène Brouard 3654 (MSC 48484!, lectotype selected here; US!, S!, isolectotypes). Note: The lectotype is selected here because the original material in
Dunkerque was probably destroyed in World War II.
Thallus (Figs. 4A–B) crustose, thin to moderately thickened, 6 continuous, epilithic; granular-areolate to verrucose; prothallus absent or delimiting
thallus as black outline; thallus surface smooth and
matt to 6 shiny, usually pale yellow, rarely becoming yellowish gray (in the herbarium), epruinose,
phenocorticate; medulla lacking Ca-oxalate
(H2SO42). Apothecia lecideine; (0.1–)0.2–0.4(–0.6)
mm in diameter; immersed, becoming adnate to
sessile; proper margin thin, reduced, inconspicuous,
black; disc black, epruinose, plane, rarely becoming
slightly convex with age; exciple narrow, poorly
differentiated, of aethalea-type (Figs. 4C, 10B) i.e.,
inner excipular hyphae narrow, hyaline, prosoplectenchymatous (textura oblita), often reduced, similar in structure and orientation to paraphyses, transient with deep reddish brown hypothecium (leptoclinoides-brown, textura intricata), outer excipular hyphae parallel, moderately swollen (textura
oblita) and usually strongly carbonized with various amounts of brown pigments (cf. elachistabrown, HNO32), pigmentation continuous with
epihymenium; hymenium hyaline, not inspersed;
paraphyses simple to moderately branched, apically
swollen, with brown (cf. elachista-brown) pigment
cap. Asci 8-spored, clavate, Bacidia-type. Ascospores (Figs. 4D–I) soon becoming pigmented, initially olive, brown at maturity, oblong to ellipsoid,
often constricted, with obtuse ends, not curved,
(12.0–)14.2–[16.4]–18.6(–22.0) 3 (6.0–)7.1–[8.1]–
9.0(–11.0) mm (n 5 60); one-septate, proper septum
developing soon and rarely thickened during spore
ontogeny, with two dark bands across each cell
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FIGURE 4. Buellia subaethalea [A–D. light micrographs (Arsène 3654 — lectotype), E–I. TEM micrographs (Ryan
27029)]. — A. verrucose-areolate thallus. — B. Close-up of verrucose-areolate thallus with 6 immersed to sessile
apothecia. — C. Transverse section of mature apothecium with aethalea-type exciple. — D. Mature, rugulate ascospore
with two distinctly darkened bands (arrows). — E. Ascus with immature ascospores. — F. Immature ascospore: (as)
ascus wall; (1) outer layer (perispore and mucilaginous sheath not distinctly differentiated); (2) inner layer (proper wall
and endospore not distinctly differentiated). — G. Wall in immature ascospore. — H. Wall layers in mature ascospore:
(s) mucilaginous sheath; (1) perispore; (2) intermediate layer; (3) proper spore wall; (4) endospore. — I. Mature
ascospore. Note: the two bands that are distinctly visible in the light microscope (Fig. 4A), do not correspond with
any distinct wall structure in the TEM (Figs 4F–I).
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BUNGARTZ ET AL.: SAXICOLOUS BUELLIA
(Fig. 4D; similar to Physconia- and/or Bicinctatype); ornamentation rugulate; septum with septal
pore canal, simple pore, and undifferentiated pore
plug; spore wall (Figs. 4G–J) differentiated into
fractured, thick perispore (0.31–0.47 mm), narrow
intermediate layer (,0.04 mm), moderately thickened proper spore wall (0.17–0.29 mm) and moderately thickened endospore (0.12–0.22 mm). Pycnidia rare, urceolate to globose, unilocular, at maturity almost entirely occupied by densely branched
conidiophores; conidiogeneous cells mostly terminal, rarely also intercalary (cf. conidiophore-type
V); pycnidial ontogeny similar to Umbilicaria-type;
conidia simple, bacilliform, 7.0–11.5 3 1.0–1.5 mm
(n 5 20).
Chemistry (Fig. 8). The thallus reacts K1 yellow to orange (forming crystals), P1 yellow to orange, C2, KC2, and CK2. Thalli show a dull fluorescence i.e., UV1 deep orange. The thallus is not
amyloid, but the hymenium reacts amyloid. The
only xanthone found in this species is 4,5-dichlorolichexanthone. In addition, norstictic, cryptostictic, methylstictic, and stictic acid were detected.
Substrate and ecology. On a variety of siliceous (HCl2) rock substrates.
Distribution (Fig. 6). A montane, subtropical
species, infrequent in southeastern Arizona and
probably more common towards the Sierra Madre
Occidental. Originally described from Michoacán,
Mexico but little is presently known about the extent of the distribution.
Notes. Buellia subaethalea is superficially similar to B. concinna, but the thallus is verrucoseareolate rather than granular and the oblong to ellipsoid spores are usually distinctly ‘‘banded’’ and
never crescent-shaped (see Discussion–Asci and
Ascospores).
Selected specimens examined.—NORTH AMERICA.
MEXICO. CHIHUAHUA. Nash 31382 (ASU). DURANGO. Nash
31087 (ASU). SINALOA. Nash 10147 (ASU). SONORA. Nash
10871 (ASU). U.S.A. ARIZONA. Apache Co. Nash 26815
(ASU). Cochise Co. Ryan 10748 (ASU); Weber L-36928
(COLO). Gila Co. Nash 39439 (ASU). Pima Co. Darrow
722, Nash 21208 (ASU). Santa Cruz Co. Nash 25353 (ASU).
TEXAS. Brewster Co. Nash 15037b (ASU).
BUELLIA TRACHYSPORA Vain., Annal. Acad. Sc.
Fennic. ser. A 6(7): 84. 1915.
TYPE: U.S.A. VIRGIN ISLANDS. St. John Co. Reef Bay,
on a cliff along a cataract [original label data: Indias occ.
S. Jan, Reef Bay, In rupe ad cataractam], 7 March 1906,
Boergesen 19067-3 (5 Vainio 09521a) (C!, lectotype;
TUR-10017!, isolectotype, both selected by Imshaug 1955).
Note: The former Danish Virgin Island ‘‘St. Jan’’ is now
part of the U.S.A. and more frequently referred to by its
English name ‘‘St. John’’. Imshaug (1955) regarded material in C as the holotype because most of Vainio’s tropical material is deposited there. However, the protologue
469
does not mention a particular herbarium (C or TUR). Imshaug’s annotation thus must be interpreted as a lectotypification (see Art. 10.5 ICBN). The material in TUR is thus
isolectotype material. Only a small fragment of this isolectotype material (Boergesen 9521a) was made available
for study at ASU. This fragment is annotated as ‘‘only part
of the type (# 9521) S. Huhtinen, Curator, Turku University Herbarium (TUR)’’. The lectotype material at C is a
well developed, large specimen and annotated in Vainio’s
handwriting as ‘‘type’’.
Buellia gyrosa Vain., Annal. Acad. Sc. Fennic. ser A.
6(7): 85. 1915. TYPE: U.S.A. VIRGIN ISLANDS. St. John
Co. On rock. 15 March 1906, Boergesen s.n. (Vainio
9583; C!, lectotype; TUR, isolectotype, both selected by
Imshaug 1955). Note: Imshaug (1955) studied type material from C and TUR and argued that B. gyrosa
‘‘. . . differs from B. trachyspora only in the gyrose nature of the apothecia. . . ’’ (p. 505). The older apothecia
of the type specimen have a folded margin. We agree
with Imshaug’s (1955) assessment that this variation
‘‘. . . scarcely warrants species recognition’’ (p. 505).
Again Imshaug’s (1955) annotation is in effect a lectotypification.
Thallus (Figs. 5A–B) crustose, thin, 6 continuous, epilithic; rimose to rimose-areolate; prothallus
distinct, delimiting thallus as black outline; thallus
surface smooth and matt, not shiny, usually pale
greenish yellow, rarely gray (herbarium specimens), epruinose, phenocorticate; medulla lacking
Ca-oxalate (H2SO42). Apothecia lecideine; (0.3–)
0.4–0.9(–1.8) mm in diameter; soon sessile; proper
margin black, conspicuously thickened and frequently fractured, always persistent, not excluded
with age; disc black, epruinose, plane, not becoming convex with age; exciple very thick, distinctly
differentiated into inner and outer parts, of trachyspora-type (Figs. 5C–D, 11C) i.e., inner excipular
hyphae large, leptodermatous, 6 isodiametric cells,
reddish brown, paraplectenchymatous (textura angularis), transient with deep reddish brown hypothecium (leptoclinoides-brown, textura intricata),
outer excipular hyphae short-celled, cells globose,
mesodermatous (textura globularis) and extremely
carbonized with large amounts of brown pigments
(cf. elachista-brown), pigmentation continuous
with epihymenium; hymenium hyaline, not inspersed; paraphyses simple to moderately branched,
apically swollen, with brown pigment cap (cf.
elachista-brown). Asci 8-spored, clavate, Bacidiatype. Ascospores (Figs. 5E–G) soon becoming pigmented, initially olive, brown at maturity, oblong
to ellipsoid, sometimes constricted, with obtuse
ends, not curved, (14.0–)18.8–[22.6]–26.5(–34.0)
3 (8.0–)8.7–[9.6]–10.5(–12.0) mm (n 5 60); oneseptate (occasionally additional septa forming at
both ends of spore cells), proper septum and spore
wall narrow, not thickening during spore ontogeny
[Beltraminea (5 Buellia)-type]; ornamentation
coarsely rugulate to areolate; septum with septal
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FIGURE 5. Buellia trachyspora (Nash 20750; A–E. light micrographs, F–G. TEM micrographs). — A. Rimose
thallus. — B. Close-up of apothecia. — C. Transverse section of mature apothecium with trachyspora-type exciple:
(1) outer, strongly carbonized exciple; (2) inner, paraplectenchymatous exciple; (3) hypothecium extending into inner
exciple. — D. Detail of outer and inner exciple: (1) strongly carbonized cells of outer exciple cannot be distinguished
from one another; (2) leptodermatous, 6 isodiametric cells of inner exciple. — E. Mature ascospores with strongly
rugulate to areolate ornamentation. — F. Mature Ascospore. — G. — Wall layers in mature ascospore: (s) mucilaginous
sheath; (1) perispore; (2) intermediate layer; (3) proper spore wall; (4) endospore.
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BUNGARTZ ET AL.: SAXICOLOUS BUELLIA
pore canal, simple pore, and undifferentiated pore
plug; spore wall (Fig. 4G) differentiated into
coarsely fractured, thick perispore (0.27–0.97 mm),
narrow intermediate layer (,0.05 mm), thick proper
spore wall (0.38–0.53 mm) and thick endospore
(0.27–0.43 mm). Pycnidia rare to common, urceolate to globose, unilocular, at maturity almost entirely occupied by densely branched conidiophores;
conidiogenous cells mostly terminal, rarely also intercalary (cf. conidiophore-type V); pycnidial ontogeny similar to Umbilicaria-type; conidia simple,
bacilliform to fusiform, 10.0–13.0 3 0.5–1.0 mm
(n 5 20).
Chemistry (Fig. 9). All spot tests are negative
(K2, P2, C2, KC2, CK2), but the thallus has a
UV1 bright orange fluorescence. The thallus medulla is not amyloid and apothecia react amyloid.
Only xanthones detected with HPLC: 4,5-dichloro3-O-methylnorlichexanthone, 4,5-dichlorolichexanthone. No other substances were found.
Substrate and ecology. On sheltered siliceous
rock (HCl2), usually close to rivers and streams.
Distribution (Fig. 6). The species was first described from the Virgin Islands and included by
Imshaug (1957) in his Catalogue of the West Indian
Lichens (Cuba, Jamaica, Puerto Rico, Virgin Islands: St. Thomas, St. John). This subtropical to
tropical species barely reaches the southernmost
Greater Sonoran Desert Region in southwestern
Chihuahua (Barranca del Cobre, La Bufa, COLOWeber L-65270). Additional collections at ASU were
collected near Compostella in Nayarit, Mexico
(outside the Greater Sonoran Desert Region).
Notes. Although this species has a unique apothecial anatomy, it has been confused with B. mamillana. Thalli of the two species are indeed similar,
but young apothecia of B. trachyspora emerge early
during the ontogeny and are never remotely lecanorine (not even a thalline collar or a thalline veil
has been observed in the material).
Selected specimens examined.—MEXICO. NAYARIT.
Nash 20751 (ASU). CHIHUAHUA. Weber L-65270 (COLO).
KEY
TO THE
SPECIES
1. Inner exciple distinctly paraplectenchymatous,
with large, leptodermatous isodiametric cells; outer exciple very strongly carbonized (small globular cells barely distinguished even in thin microtome sections); spores with coarsely rugulate to
areolate ornamentation (clearly visible at 4003)
--------------------------------------------------------- Buellia trachyspora
1. Inner and outer exciple not as above; spores
smooth to rugulate but not coarsely areolate (ornamentation, if present, barely visible at 4003)
----------------------------------------------------------------------------------------- 2
2. Thallus granular to bullate or verrucose-are-
3.
3.
5.
5.
471
olate i.e., entirely composed of small granules
or areoles with distinctly granular surface ------ 3
2. Thallus rimose to areolate, not verrucose or
granular ------------------------------------------------------------------ 4
Mature spores oblong to ellipsoid, with obtuse
ends, not curved, with two spore cells covered by
broad, darkened band; thallus granular-areolate to
verrucose --------------------------------------- Buellia subaethalea
Mature spores ellipsoid to 6 citriform, often with
6 tapered ends, frequently curved (crescentshaped), the two spore cells never covered by
darkened band; thallus granular to minutely bullate ------------------------------------------------------ Buellia concinna
4. Apothecial disc deep brown, rarely blackened; young apothecia remaining immersed,
or if emergent, distinctly lecanorine i.e., with
distinct thalline margin; thalline margin darkening and eventually excluded; apothecia becoming lecideine with age; ascospores rugulate --------------------------------------------- Buellia mamillana
4. Apothecial disc black, not brownish black;
young apothecia sometimes emerging with
thalline veil but never distinctly lecanorine;
ascospores smooth to inconspicuously microrugulate ------------------------------------------------------------------ 5
Thallus thick, areolate; usually with distinct black
prothallus along thallus margin; apothecia pruinose or epruinose; exciple thick, of intricately intertwined hyphae, carbonized by diffuse, aeruginose (HNO31 violet) pigment; conidia bacilliform
to fusiform -------------------------------------------- Buellia halonia
Thallus thin, rimose to rimose-areolate; with or
without faint black prothallus; apothecia epruinose; exciple thin, of narrow, parallel hyphae with
swollen end cells, carbonized by brown, non-aeruginose pigment (HNO32); conidia filiform ------------------------------------------------------------------ Buellia prospersa
DISCUSSION
Thallus. Even though all species can be characterized by xanthones, thallus color varies considerably among the species. Some species like B.
prospersa and B. mamillana may have very pale
thalli and the presence of xanthones in these thalli
may thus be overlooked. Thalli of B. subaethalea
and B. concinna become sordid yellow with prolonged herbarium storage. Buellia halonia is usually greenish yellow rather than pale yellow. The
presence of xanthones in the thalli is generally indicated by a yellow to orange fluorescence in longwave (ca 350 nm) ultraviolet light. Intensity of this
fluorescence; however, varies from pale to very
bright even among thalli of the same species. Buellia concinna and B. halonia both have a C1 and
KC1 orange thallus, but the other species usually
have no indicative C (or KC) reaction even though
their thalli contain xanthones.
Thallus morphology for each of the species is
quite distinct and most species can usually be recognized even without anatomical analysis. Buellia
prospersa has thin, usually inconspicuous, rimose
thalli (Bungartz et al. 2004). In comparison, thalli
of B. halonia are much thicker and distinctly are-
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FIGURE 6. Distribution of species treated here in the Greater Sonoran Desert Region (Floristic Provinces according
to Shreve & Wiggins 1964).
olate (Figs. 2A–B). The thallus of Buellia concinna
is composed of small and 6 dispersed granules
(Figs. 1A–B), whereas B. subaethalea has a verrucose surface of an essentially areolate thallus
(Figs. 4A–B). Buellia mamillana and B. trachyspora have very similar smooth, rimose thalli which
are finely fissured (Fig. 3A–B, 5A–B). These two
species are difficult to distinguish by their thalline
characters alone and have thus been confused.
Apothecia. Buellia trachyspora and B. mamillana have apothecia that are unique among species
in the genus Buellia. Apothecia of B. trachyspora
become very large (up to 1.8 mm in diameter). Unlike in B. mamillana these large apothecia very
2004]
BUNGARTZ ET AL.: SAXICOLOUS BUELLIA
FIGURE 7.
473
HPLC chromatograms of Buellia concinna (Nash 38098) and Buellia halonia (Nash 17166).
soon emerge and become sessile. They remain distinctly flattened, with a plane disc and a thick margin (Fig. 5B). Due to the extreme carbonization of
the outer exciple, this margin often becomes
coarsely fractured in dry herbarium specimens. The
carbonized cells of the outer exciple are small and
6 globose, and can barely be distinguished because
of their strong carbonization. The inner exciple is
entirely composed of large, isodiametric cells and
thus distinctly paraplectenchymatous (Figs. 5C–D,
474
THE BRYOLOGIST
FIGURE 8.
[VOL. 107
HPLC chromatograms of Buellia mamillana (Nash 29793) and Buellia subaethalea (Nash 25353).
11C). Because of the angular shape of these cells
the plectenchyma may be referred to as textura angularis. The inner exciple of B. trachyspora is entirely composed of thin-walled cells that are not
carbonized. In contrast, many other Buellia species
have an outer exciple of 6 angular cells, which are
smaller, carbonized, and mesodermatous. This texture has also been referred to as textura angularis
but is quite different from the texture seen in B.
trachyspora (Bungartz & Nash 2004b,c; Bungartz
2004]
BUNGARTZ ET AL.: SAXICOLOUS BUELLIA
FIGURE 9.
475
HPLC chromatogram of Buellia trachyspora (Weber L-65279).
et al. 2004; Scheidegger 1993). The exciple of B.
trachyspora is therefore unique and cannot be assigned to any of the types described by Scheidegger
(1993).
Buellia mamillana is the only species currently
known from the Sonoran Region with apothecia developing a distinct thalline margin (Figs. 3B–D).
Other species currently included in Buellia s.l. may
have some irregular thalline collar (Bungartz &
Nash 2004b) or a thalline veil (Bungartz & Nash
2004a), but only in Buellia mamillana do the young
apothecia develop a distinct lecanorine margin with
a cortex and algal layer embracing the young apothecium (Figs. 3C, 11A). As a result, the species
was originally described as a Rinodina. It is unusual
that this initially well developed thalline margin
subsequently becomes excluded and replaced by
hyphae from the proper exciple (Figs. 3D, 11B).
During this process the deep brown pigmentation
of the disc extends more and more across the initially pale margin. This shift in pigmentation is a
result of internal growth processes. The inner exciple expands and the thalline outer part becomes
excluded. During this ontogeny, apothecia may
even have a lecanorine exciple on one side, and a
lecideine exciple on the other side of a transverse
section. This ontogeny is unusual, and Weber
(1986) who transferred the species into Buellia, did
not describe this unusual ontogeny. He interpreted
the apothecia as ‘‘clearly lecideine’’, possibly because older apothecia indeed become lecideine.
Apothecia of the other species are not as unusual.
Scheidegger (1993) already described the intricately interwoven excipular hyphae of B. halonia (Figs.
2C–D) and assigned it to the leptocline-type. This
exciple type is not significantly differentiated
throughout and it is initially less strongly pigmented inside but soon becomes evenly pigmented.
Buellia concinna has an exciple similar to B. halonia, but lacks the aeruginose pigment. Like in B.
halonia, excipular hyphae of B. concinna are 6
evenly pigmented, little differentiated, and intricately interwoven (Figs. 1C, 10D). Despite this,
Scheidegger (1993) assigned the exciple of B. concinna to the dispersa-type (Fig. 10C), which he described as composed of hyphae radiating from a
deep reddish brown hypothecium, with a 6 hyaline
transition zone and a strongly carbonized outer layer of moderately swollen cells. The exciple of B.
concinna does not show this differentiation and
thus should be assigned to the leptocline-, not the
dispersa-type.
The exciple of B. subaethalea is less prominent
and the inner hyphae are thinner than in B. halonia
or B. concinna. In B. subaethalea only the outer
cells are usually 6 swollen and strongly pigmented
(Fig. 4C). The exciple of B. prospersa is similar
(Bungartz et al. 2004). Both species belong struc-
476
THE BRYOLOGIST
[VOL. 107
FIGURE 10. Line drawings of exciple types in Buellia s.l. — A–B. Variation of the aethalea-type exciple, characterized by thin, more or less parallel hyphae, similar in orientation and structure to paraphyses. — A. Apothecia
remaining immersed and exciplular hyphae strongly reduced (e.g., Buellia aethalea and Buellia eganii, see Bungartz
& Nash 2004a). — B. Apothecia emerging and excipular hyphae becoming pronounced (e.g., Buellia subaethalea, or
B. spuria and B. stellulata, see Bungartz & Nash 2004a). — C. The dispersa-type: inner, mesodermatous hyphae radiate
from hypothecium towards outer exciple. Pigmentation in young apothecia does not extend far into central exciple, but
with age exciple becomes evenly pigmented. Outermost cells are distinctly swollen, shorter, and more isodiametric
(e.g., Buellia dispersa and B. nashii, see Bungartz 2004, Bungartz et al. 2002). — D. The leptocline-type: hyphae are
mesodermatous, little differentiated throughout, and with age more and more intricately interwoven (e.g., Buellia
concinna and B. halonia).
turally to the aethalea-type (Fig. 10B), even though
they lack the aeruginose pigment cinereorufagreen.
Asci and Ascospores. All species have Bacidiatype asci and a dark hypothecium, characters typically associated with the Buellia-clade rather than
the Rinodina-clade (Helms et al. 2003). Ascospores
of all species are not pigmented at very early stages, soon become olive, and are distinctly brown at
maturity. Buellia trachyspora has spores with no
distinct septum thickening and may thus be assigned to the Buellia (5 Beltraminea)-type, even
2004]
BUNGARTZ ET AL.: SAXICOLOUS BUELLIA
477
FIGURE 11. Line drawings of exciple types in Buellia s.l. (continued). A.–B. The mamillana-type (currently known
only from Buellia mamillana). — A. At early stage of ontogeny a distinct thalline exciple is still present. — B. Thalline
exciple becomes excluded by a progressively expanding proper exciple. — C. The trachyspora-type: differentiated into
strongly carbonized outer exciple of small, globular cells and reddish brown, large-celled and distinctly paraplectenchymatous inner exciple (currently known only from B. trachyspora). — D. The vilis-type: hyaline hypothecium of
thin, strongly interwoven hyphae extends into inner exciple and reacts very strongly iodine-blue; outermost exciple is
deeply pigmented with blackish red pigment atra-red (currently known only from Buellia vilis, see Scheidegger 1993
and Bungartz 2004).
though the spores are usually 6 constricted along
the septum and younger spores show some inconspicuous swelling where the septum forms. The ascospores of all other species have a distinct septum
thickening at least during some part of their ontogeny. They can thus be referred to the Physconiatype. In most species this septum thickening is
ephemeral i.e., it is most pronounced in premature
spores but becomes reduced in mature and overmature spores. Only B. prospersa has a septum,
which usually is more persistent and considerably
thicker than in any of the other species. These
spores could, therefore, also be referred to as Orcularia-type. None of these spore types refers to
strictly distinctive categories, but represent an idealized concept of an inherently dynamic and transient development of the spore. For example, even
though both B. concinna and B. subaethalea may
formally be assigned to the same spore type, their
spore ontogenies are distinctly different from all of
the other species. Spores of B. concinna develop a
distinctive shape (Figs. 1D–F). The premature to
mature ascospores are characteristically curved and
have 6 tapered or pointed ends. These spores are
478
THE BRYOLOGIST
best described as a ‘‘half-moon’’ or crescent-shaped
(Fig. 1E). The characteristic shape is not always
observed because some spores do not become
curved. These spores nevertheless develop 6 tapered ends and thus appear 6 citriform. All other
species have spores with obtuse ends.
Old spores of B. subaethalea frequently become
6 constricted, but are more importantly characterized by two, broad, and distinctly darkened bands
across each of the two spore cells (Fig. 4D). In the
light microscope these bands feign lateral wall
thickenings that must not be confused with distinct
structural lateral thickenings in Callispora-type ascospores. Ultrastructural observations currently
suggest that the ‘‘banding’’ does not correspond
with structural layers of the lateral cell walls. However, comparing the ultrastructure of ascospores at
different ontogenetical stages may elucidate whether these patterns correspond to some ultrastructural
differentiation of the spore wall. Similar ‘‘bands’’
are known from ascospores of Rinodina (for example, the Bicincta-type) and may be caused by a
phenomenon similar to the torus.
Spore ornamentation is a result of the perispore
differentiation. Thus, the ultrastructure of this outer
spore wall layer corresponds to the ornamentation
observed in the microscope. In B. trachyspora, the
perispore appears coarsely fractured in the TEM
(Figs. 5F–G) and a rugulate to areolate ornamentation (Fig. 5E) is visible in light microscope even
at 4003 magnification. In other species, spore ornamentation can also be observed, but is much less
distinct. Buellia subaethalea also has rugulate ascospores (Figs. 4D, I), but the ornamentation is
barely visible at 4003 and distinct only at 1,0003
magnification. The rugulate ornamentation of Buellia mamillana is even less distinct (Fig. 3E) and the
perispore is not strongly fractured (Fig. 3G). Finally, B. halonia and B. prospersa both have a microrugulate ornamentation. This fine ornamentation is
quite distinct at 1,0003 in B. prospersa, but barely
visible in B. halonia.
Conidiomata. The conidiomata of all species
are quite similar. They develop in urceolate to globose pycnidia (Fig. 2J) that are almost entirely occupied by conidiophores. These conidiophores are
branched and predominantly form conidia at the
apices (Fig. 2K). Some conidia are also formed intercalary on short bayonet-like protrusions. The conidiophores do, however, not anastomose at their
apices and thus resemble more closely type V instead of type VI (Vobis 1980). Buellia concinna has
the shortest, 6 ellipsoid conidia. Buellia halonia
and B. subaethalea have slightly longer, bacilliform
conidia. Conidia of B. mamillana and B. trachyspora are bacilliform to fusiform and B. prospersa
has filiform conidia. Because of its long conidia, B.
[VOL. 107
prospersa also has been accommodated in the genus Amandinea [as Amandinea lecideina (H. Mayrhofer & Poelt) Scheid. & H. Mayrhofer]. The segregation of an entire genus on this single character
is, however, problematic (Bungartz et al. 2004).
Chemistry (Figs. 7–9). Detailed information on
the thallus chemistry of the species is provided in
the species descriptions. Buellia concinna and B.
halonia are the only species that contain asemone,
thiophanic acid, and arthothelin together with the
closely related isoarthothelin. The other species
contain a variety of different lichexanthones that
are also present in B. concinna and B. halonia.
Buellia subaethalea is the only species that contains
a single xanthone (4,5-dichchlorolichexanthone). In
contrast, B. mamillana has the most complex chemistry, regularly with a variety of xanthones, depsides, and depsidones. This chemical variation does
not appear to correlate with any morphological difference.
Distribution (Fig. 6). Of all the species examined only B. halonia shows a distinctly coastal,
maritime distribution. Buellia concinna and B. subaethalea are montane to subalpine species and
possibly occur in the same habitats. Buellia mamillana and B. trachyspora show subtropical affinities. Buellia mamillana extends far into the Sonoran
Desert Region and typically grows along mountain
streams. Buellia trachyspora grows in similar habitats but barely extends into the southernmost part
of the Sonoran Region.
ACKNOWLEDGMENTS
We are grateful to John Sheard, University of Saskatchewan, Canada and Scott Bates and James Lendemer ASU,
Arizona, for reviewing the original manuscript. Roland
Moberg, Museum of Evolution, Uppsala and an anonymous reviewer provided additional instructive advice.
Hannes Hertel and Dagmar Triebel, Botanische Staatssammlung München, helped resolve questions regarding the
correct typification of several taxa. Ulrik Søchting and
Eric Hansen, University of Copenhagen (Denmark) and
Seppo Huhtinen, University of Turku (Finland) provided
valuable advice on the correct typification of Buellia trachyspora. Several lichenologists responded quickly to a
literature search for rare taxonomic literature posted on
the lichen list server. Loans from the following herbaria
to ASU are greatly appreciated: BM, C, CANL, COLO, FH, H,
hb. Scheidegger, hb. Kalb, hb. Sheard, L, M, MICH, MSC,
SBBG, TUR, UPS, and W. This study was supported by a
Research Grant in Plant Systematics from the International Association of Plant Taxonomists (IAPT) and National
Science Foundation Awards to ASU (DEB-0103738, DEB9701111). A research visit of the first author to MSC was
supported by a National Science Foundation Award (DBI0237401).
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