Professional Documents
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OPERA .
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Copenhagen 1996
Opera Botanica
Stefan Ekrnan
This is a taxonomic revision of the corticolous and lignicolous taxa of the genera
Bacldiu and Bacidina (Lecauoraceae, Lecanorales, lichenized Ascomycotina) in the
continental United States and Canada. Twenty-seven species of Bacidia (one of
which is divided into two subspecies) and twelve species of Bacidina are recognized.
Tlie tnorphology, secondary chemistry, ecology, and phytogeography of these spccies
are discussed, and a key is provided. Apotheciu~npigmentation, thallus composition,
presence of crystals in the proper exciple, hymenium height, thickness of paraphyses,
spore shape and size, in Bacidina sometimes also conidial type and tholus structure,
have proved to be the most important characters to distinguish betwcen species.
Circumscriptions of Bacidia and Bacidinn are attempted. Tholus structure, cell struc-ture
of the proper exciple, presence of goniocysts, presence of crystals in the thallus
cortex, and size of conidiogenous cells are the most valuabie characters in the
delimitation of these genera. Nine new taxa arc described: Bacidia d ~ f l i c t a ,B.
helicospom, B. salmonea, Bacldinina aenea, Bn. californica, Bn. crystallifera, Bn.
mmea, Bn, squamellosa, and Bn. varia. Twelve new combinations are introduced:
Bacidia campalea, Bacidia laurocerasi subsp. idahoensis, Bncidina assulata, Bn.
egenvloidea, Byssolomn meadii, Frllhanera floriduna, Hel-!elbna alasketrsis,
Lecania stigmatellu, L, subfuscula, Puchyphiale gj~alizella, Ropulospora phaeo-
pluca, and Ophioparma rubricosa. In addition to tht navIy described species, I2
species of Bacidiu and four species of Bacidinn are correctly reported from the study
area for the first time. The genera Psorella and Tuniniupsis are reduced into syn-onymy
with Bacidia.
Contents Surface
Cortex
Jntroduction Photobiont layer
Objectives Medulla
Material and methods Attachment
T h e North American Bacidia and Bacidina Vegetative dispersal
- an outline Apothecia
Morphology and anatomy Morphology and ontogeny
ThalIus Proper exciple
Growth form Hypothecium
Prothallus Subhymenium
-
/
(~opalospora) A
f f genera in the \
r Bacidia
sensu Zahlbl.
Fig. I. Transfers of species from Bacidia sensu Zahlbruckner to other genera made after the publication o f Catalum lichenurn
universalis (Zahlbruckncr 1921-1940), including those suggested in this study. - The circumscription of the Ectolechiaceae by
M z d a (1986) is probably artificial, being based mainly on the presence of campylidia. The presence of this structure has been
shown to appear in widely different families and thus to be polyphyletic in origin (Sirusiaux 1986, Aptroot & Sipman 1993,
Liicking 1995, Strusiaux 1995). Lucking et al. (1994) doubt the distinctness of the family Ectolechiaceae to Pilocarpaceae.
done here or treating all North American species of Ba- Slightly more than 2000 specimens were examined,
cidiu (but not Bacidina) irrespective of substratum including material from the following herbaria (abbrevi-
choice. Treating all North American Bacidia and Ba- ated according to Holmgren et al. 1990): BG, CANL,
cidiaa was out of the question, due to the time limits set COLO, FH, FLAS, H, LD, MICH, M W , MSC, MU,
on the work. My choice fell on the present delimitation, NY, OSC, S, SFSU, TSB, UBC, UPS, US, WIS, WTU,
for the simple reason that it would be impossible to de- WWB, and the private herbaria of Charis Bratt, WiIla
limit Bacidia properly without also studying members Noble, and Shirley Tucker. My own material, compris-
of Bacidina. '
ing about 3 0 0 specimens, is deposited in LD. In the
course of finding type material for names that might be
applicable to North American species (well over 200
Material and methods names), I have seen material from numerous other her-
baria.
Study area
The study area includes the continental United States Field work
and Canada. This area is henceforth referred to as
"North America", although 1 am aware that other ways Field work was carried out in California, Oregon,
of delimiting this continent have been suggested. Washington, and British Columbia in March-May
1989, in Minnesota, Michigan, Ontario, Quebec, New
Brunswick, Nova Scotia, Maine, Delaware, and Mary-
Material land in September-October 1989, in Louisiana, Missis-
All North American species of Bacidia and Bacidina sippi, and FIorida in October-November 1993, and in
found on tree-bark or lignurn have been treated in their California and Arizona in March-April 1995. From the
entirety, i.e., specimens from all substrata have been in- Iast journey, only some ecological results have been in-
cluded if at least one specimen of the species is corti- cluded in this work, whereas the majority of the collec-
colous or lignicolous. . . . , tions still await revision. Twenty-three out of 28 taxa of
Bacidia and 8 out of 12 taxa of Bacidina have been below) is c. 20 or higher in common species. In uncom-
studied in the field in North America. Two additional mon species, I have used as many collections as possi-
species occurring in North America have been studied in ble, provided they have been reasonably large. It was at-
Sweden but not in North America. tempted to use specimens from various parts of the
distribution range. I have refrained from using more
than one specimen from a single locality. Otherwise,
Morphology and anatomy there is a risk of a single genotype becoming over-repre-
sented in the measurements.
Characteristics of the thallus and apothecia were exam-
ined from hand-cut. sections or squash preparations Measurement accounts
mounted in deionized water, except paraphyses, which
were measured in a 10 % aqueous solution of KOH. Measurements are generally given as "(minimum value
Characters were studied under an ordinary light inicro- observed -) lowest specimen arithmetic mean observed -
scope, a polarized light microscope, and/or an interfer- arithmetic inean of all observations - highest specimen
ence contrast light microscope, depending on what was arithmetic mean observed (- maximum value observed)
most suitable. Lactophenol-cottonblue (LCB) was uscd (S, N, n)", where s is the standard deviation of all ob-
to enhance the contrast when studying excipular or corti- servations, N the number of specimens studied, and n
cal structures. When studying excipular structures, a 0.2 the nwnber of observations per specimen. The totaI num-
U/o aqueous solution of sodium hypochlorite (a comner- ber of observations is thus N X n. When N= 1 or n= 1,
cial solution diluted about 20 times) was sometimes the lowest and highest specimen arithmetic means are
used to bleach pigments and improve the optical clarity not meaningful, and have consequently been left out.
of the gelatinous cell walls. Higher concentrations of hy- The unit of the standard deviation is always the saine as
pochlorite shouId be avoided, since the cell walls may the character itself, but has generally been left out of the
swell and rupture, and thus alter the appearance of the description in order to make them easier to read. All
cell structure. characters were shown to be about normally distributed
The following standard reagents were used to study before using the arithmetic mean and standard deviation
the colour reactions of pigments and solubility of crys- as measures of distribution center and dispersal.
tals: a 10 % aqueous solution of KOH (abbreviated K), a In some cases, where detailed measurements were
commercial 4.4 % aqueous solution of sodium hy- difficult to obtain andlor of little use, the standard form
pochlorite (abbreviated C), and a 50 % soIution of "minimum value observed - maximum value observed"
HNO, (abbreviated N). was employed.
Rout~neinvestigations of the structure of the ascus There are a number of reasons for using this particu-
apex were carried out on gently squashed hand-cut sec- lar method of presenting measurements. Basically, I feel
tions stained with a 0.3 % aqueous solution of iodine, that making a large number of measurements and then
following pretreaanent with K. I decided to use this preparing descriptions using only the minimum and
method after having performed a small pilot study on a maximuin values observed is a huge waste of useful in-
small number specimens of a few randoinly chose11 spe- formation. Fortunately, it has become increasingly com-
cies. Asci of these specimens were stained according to mon to see ineasureinents (particularly of spores) that
the method mentioned, with the same iodine solution are presented as the aritliinetic mean k 1 or 1.5 standard
but without pretreatment with K, as well as with a 0.3 O/o deviations. This method is commendable, but I think
solution of iodine in which lactic acid was substituted such figures are slightly difficult to use in practical de-
for water, wirh and without pretreatment with K. All termination work, particularly if similar species have
four methods were shown to give the same result, but overlapping measures. The advantage of the method
the method finally chosen was the one which gave the used here is that different information can be extracted
optically clearest preparatioiis combined with a low depending on the needs of the user. The disadvantage is
number of ruptured asci. The "lactic-iodine", which has that the descriptions become long and difficult to read. I
lately been used by several authors, Including myself, in think, however, that the advantages outweigh the disad-
this case resulted in a high proportion of ruptured asci, vantages. If one wishes to compare species with each
probably since the solution is very viscous and more other or compare the variation between different distri-
force is needed when squashing the preparations. Partic- bution areas of a single species, one can use the arith-
ular care was always taken not to study asci with rup- metic mean and the standard deviation. The remaining
tured walls, since such asci often show "freak" struc- figures can be used for ordinary determinations. If the
tures. aim is to see whether a single measurements of, i.e.,
The number of speclrnens from which measurements spore length lies within the variation of a particular spe-
of quantitative characters have been extracted (N; see cies, one is confined to the minimuin and maximum val-
line along the edge of the thalIus, often where the speci- shaped, coarse, irregular projections. I have used a ratli-
men meets other lichens. Remarkably, the latter kind of er arbitrary, five-graded scale to describe the surface, ar-
prothallus is present in five out of eight species that are ranged here by increasing height of the projections: (I)
mainly distributed south of the study area, whereas it is Smooth, with no projections, (2) wrinkled, (3) waned,
present in only one out of 18 species endemic to North (4) tuberculate, and (5) subsquamulose. It should be
America or mainly distributed in temperate areas of the notcd that I use the term "warted" for a surface with
northern hemisphere (difference in proportion tested by rather high, often very irregular projections that are
Yates' corrected chi-square; p= 0.006). This is particu- more or less unequal in size and direction. This means
larly interesting, since there is no reason to believe that that I do iiot refer to a regularly papillose surface, nor
the southern species are more closely related to each should it be translated to the anglisized latin word "ver-
other than to other species of Braeidia. Thus, it appears rucose", which is often used to describe the shape of an
that the black, episubstratal prothallus is polyphyletic in entire areole.
origin and that its evolution has somehow been fa- Most species of Bacidiu and Bacidinu have a wrink-
voured by a tropical or subtropical climate. led or wafled thallus surface. A smooth thallus inay oc-
casionally be present in a number of species, but no spe-
Surface cies has an invariably smooth thallus. Likewise, a tuber-
culate to subsquamulose thallus may be present in some
The surface of many species is furnished with variously individuals of a few species, although rarely and never
except in very narrow lumina. They are usually 8-25 p wider, often globose or widely ellipsoid. Often, only the
long, 1-3 p wide, and 5-1 1 times as long as wide. The terminal cells are enlarged, but in some species the out-
walls are gelatinized and thick, and the boundaries be- ermost 2-4(-11) cells are more or less edarged. These
tween adjacent hyphae cannot be discerned. The dis- wide, outer cell lumina can either form a distinctly de-
tance between two adjacent cell lumina in h e interior of limited layer along the rim of the proper exciple, or the
the proper exciple practically always exceeds the width lumina may enlarge gradually towards the rim. Only
of a cell lumen, but more commonly this distance is 2-5 four species regularly possess a distinct layer of en-
times the width of a lumen. larged cells along the rim of the proper exciple thicker
In many species of Bacidia, the outermost cells of the than two cell layers, namely B. labsistens, B. capnpaba,
excipular hyphae, along the rim of the proper exciple, B. mseolu, and B. su@a (Fig. 3C).
differ in shape and size from the inner ones in being Crystals can be found in 14 out of the 27 species of
Fig. 4. Sections of apothecia in Bacidina, unstained but treated with a diluted hypochlorite solution, viewed in interference
contrast.- A. Bacidina chloroticuh (Wong 2709A, CANL), excipular hyphae that are indistinctly radiating and possess ellipsoid
cell lumina. Bar = 20 pm. - B. Bacidina varia (Ekman L1 135 LD), fairly distinctly radiating excipular hyphae with broadly
cylindrical to ellipsoid lumina that become gradually larger towards rim of proper exciple. Bar = 20 pm.
Bacidia (Fig. 3A), and they constitute a good character na and longer and narrower ones. Commonly, they are
for separating some species. They are often minute, up spatially separated. In Bn, ar~loldiana,Bn. egenula, Bn.
to I pm wide, but can in some species be much larger, egenuloideo,and Bn. sq~amellosa,the cell lumina are
up to 10 pm. Only in a few species the crystals are more distinctly wider in the lower, older part of the proper ex-
or less evenly dispersed in part of the proper exciple. ciple than in the upper, younger part. In Bn. varia (Fig.
More commonly, the crystals are aggregated in clusters. 4B), the reverse is true, i.e., the upper lumina are often
Characteristically, the clusters are elongated and radiat- wider than the lower ones. In this species, there is also a
ing, i.e., arranged in the direction of the excipular hy- clear tendency for the hyphae to expand gradually to-
phae. Oil droplets are never present in any of the species wards the rim. The proper exciple of Bn. aenea, Bn.
of Bacidia or Bacidina. cystallifera, and Bn. sp. #l also contain hyphae with
The proper exciple of Bacidina is different from that cell lumina that gradually expand towards the rim, but
in Bacidia (Fig. 4A-B). In some species, the excipular in these species there is no difference between the upper
hyphae are distinctly radiating, in others not. A distinc- and the lower part of the proper exciple. Finally, the
tion between two layers, Iike in some Bocidia, is never proper exciple of Bn. assulata is composed of an appar-
found in Bacidina. The hyphae of the proper exciple are ently fairly unorganized mixture of hyphae with wide
sparingly to abundantly furcate, and usually abundantly and narrow Lumina, although there appears to be a ten-
anastornosed (sparingly to moderately sg in Bn. aenea dency for the upper hyphae to be slightly narrower than
and BB, chioroticulu). The cell lumina in the interior of the lower ones.
the proper exciple are very variable, but all species con- The cell lumina of the excipular hyphae found in the
tain a fair amount of short and wide cell lumina (almost species of Bacidinla are, like in Bacidia, usually con-
globose, f ellipsoid, rectangular, or irregular) with a stricted at the septa, very narrow hyphae excepted. The
width exceeding 2.5 pm and a lengthlwidth ratio less walls are gelatinized, but unlike Bacidia, they are thin.
than 4.0. In Bn. califorfiica, Bn. chloroticula (Fig. 4A), The distance between two adjacent cell lumina is usual-
and Bn. ramsa, the cell lumina are homogeneously glo- ly less than the width of a ceIl lumen. Furthermore, the
bose to ellipsoid throughout the proper exciple, but the species of Bacidina lack enlarged terminal cells of the
remainder of the species contain a mixture of such lurni- excipular hyphae. CrystaIs are found in only one spe-
ly homogeneous. It contains a conical, high or IOW,ocu- faintly amyloid c-layer is, like in Bacidia, narrow in
lar chamber and a more or less conical, blunt or pointed most species. A few species deviate, however, in pos-
axial body h a t never reaches ail through the d-layer sessing a distinctly widened c-layer, namely BR. cvstal-
(Fig. 5A-D). In some species, the axial body is sur- Iifera, Bn. egenuloidea, Bn. squamellosa, and Bn, sp.
rounded by a distinct or fuzzy zone that is very strongly #I. Particularly in Bn. squapnellosa, the c-layer is ex-
amyloid, more so than the remainder of the d-layer. The tremely well developed relative to the d-layer, resulting
width of the axial body never exceeds one third of the in an ascus apex with an appearance unlike any other li-
entire width of the d-layer. The c-Iayer is constantly nar- chen that I know of.
row. All the ascus appearances are variations on one and
Within Bacidina, on the other hand, there is more ex- the same theme: an ocular chamber that may or may not
tensive variation in the composition of the ascus (Fig. be present, and a distinct axial body that is basically
5E-R). A distinct, high or low, more or less conical, conical but varies in height and width. There is a strik-
blunt or pointed ocular chamber is oRen found in Bn. ing difference in variation pattern between the two gen-
aenea, Bn. arnoldiana, Bn, ussu!ata, Bn. caI$ornica, era, however. Whereas Bacidia exhibits rather little var-
Bn. egenula, Bn. egenuloidea, sometimes also in Bn. iation in ascus apex structure, Bacidina, despite the
varia and in Bn, sp. #l. In Bn. chloroticula, Bn,crystal- smaller number of species, is much more variable. An
l$era, Bn, mmeu, and Bn. squamellosa, on the other attempt to explain this difference in variation pattern
hand, an ocular chamber is lacking. An axial body is must necessarily be highly speculative. There are two
present in all species. It is usually more or less conical principal explanations:
or cylindrical, and constitutes one third or more of the l. Contraxy to Bacidia, the ascus apex of several spe-
entire width of the d-layer. In Bn. mmea, the axial body cies of Bacidina has lost its function in the dispersal of
is exkernc, taking a cushion-like, very wide and low the spores. Hence, it is not subjected to any selective
shape. h Bacidina cal@rnicu and Bn. crystallifera, the pressure. Through drift, various deviating ascus apices
axial body vertically extends through the entire d-layer have evolved.
in some, but not all, asci. This is variable even within 2. The ascus apex of several species of Bacidina is
the same specimen. A distinct, strongly amyloid zone highly specialized with respect to the mechanism of
surrounding the axial body can be found in Bn. assula- spore dispersal, which may in its turn be an adaptation
ta, Bn. wlifornicu, and (sometimes) Bn. varia. In the to, e.g., certain climatic or chemical conditions (the rea-
last species, however, most asci Iack this zone. The son for the specialization is actually unimportant to the
Fig. 5. Amyloid reaction of asci. The immature spore content has been omitted from the figures. - A. Bacidin arcevtina. - B. B.
heterochroa. - C . B. medialis. - D. B. rubella. - E. Bacidina aenea. - F,Bn. arnolcliana. - G . Bn. m u l a t a . - H-I. Bn. calflornica.
- J. Bn. chbmticula. - K-L. Bn. ciystall$em. - M. Bn. egenula. - N. Bn. egenuloidea. - 0.Bn. rurne~~. - P. Bn, squamellosa. -
Q . Bn. vuria. - R. Bn. sp. # 1. - Drawings by Ulf Arup.
l?" -
r
-
;"
;g&+E:e;;$
yN"=.PU
Tdb. 2 . The presence of v;irious sporc sllapcs in Rrrririio and Bucidinrt: and the length (p). width (p), Icngthlwidth ratio, and
number of septa per sporc. Thc valucs prescntcd are not the endpoints uf the variatiun. but the interval within which an average of
10 (or in somc cascs 20, denolcd hcrc with an "*") measurements per individual varics. A single value indicates that spores, due to
availabilitv o f ~natcrial.have bccn mcaured in onc specimcn onlv3 and that hence only one average value is obtainable.
17.5-24.5 6.8-14.3
17.1-22.1 4.7-7.5
25.3-36.4 5.0-10.7
3.6-4.4 5.7-6.4
19.5' 10. l *
15.3-17.1 6.6-8.0
oircun~sper.lu 6.0- 15.8 3.0-6.8
difl;rlc/a I 12.4-19.3 4.1-7.8
ji.iesil~/ta -1- 10.8" 7.2"
he1rrorpor.u + 18.0-22.9 7.8-14.8
/ieturnchr.oa + 11.7-18.9 4.5-13.6
hos~hel~r>irle,v 5,4* 3.I*
ig/?irtrii 5,9* 3.0*
i/z,s~t!ori.s 1 6.4-14.1 6.6-1 0.3
lrtrrrcjcc,rrt.si subsp. !(I. + 18.3-3 1.5 1 1.9-22.2
1rtrrrurc.rrrsi subsp. iil, + 23.9-30.4 17.5-23.0
rncdirliis 6.7- 10.7 3.0-4.2
mr(tuhilt.~ T 12.7- 14.4* 5.9-6.3 *
pol~uhrou 11.4-21.6 3.1-1 1.9
r'uugens 15.8-22.8 3.5-12.5
- 5.6-8.7
ru).r elk1 20.8-23.5
ruhriltr - 15.7-24.4 3.2-8.7
17.8-20.1 6.6-7.3
30.2-36.5 6.8-9.4
15.5-30.6 4.3-13.6
4.6- 19.2 3.0-9.7
14.4-2 1.9 6.3-12.6
6.3-1 1.2 3.0-4.3
Tab. 3. The total number of spores with odd and even number of septa, the proportion of odd- to even-septate spores, and the
significance level of the chi-square values obtained from the goodness-of-fit test for species of Bacidia and Bacidina. The null
hypothesis tested was that the number of odd- and even-septate spores are randomly distributed, i. e, that both the number of odd-
and even-septate spores (Odd+ Even)R. Only species in which at least 30 spores have been studied nnd in which the majority of
the mature spores possess more than three septa have been included in the test. The significance levels are denoted * for
0.01q50.05, ** for 0.001<p~0.01,*** for p50.001, and ns (not significant) for p>0.05.
Sign. level
Odd Even Oddeven off
Bacidia
absistens
aggregatu!~
arceulina
auerswddii
campaka
dlzructa
helicospora
heterochroa
insubris
luurocernsi subsp. idahoensis
luumcerasi subsp. Iaumcerasi
poiychrua
reagens
rosella
rubella
salmonea
schweinihii
subincompta
sufisa
Bacidina
caliJornica
crystallifera
ramea
No. of conidial
types la lb lc Id 2 3 4
Bacidia
abxistens
aggregatulu
arceertinu
auerswaldii
bintorina
campalea
circumspecta
d~flacta
friesiana
helicospora
heterochroa
hostheleoides
igniarii
insulan's
laurocerasi
medialis
mutabifis
reugens
roselIu
rubella
msseoln
slalmonea
schweiniizii
subincompta
suflusa
verm$era
Bacidinu
aenea m
nrnoldiana S
nssulata S
cali~ornica S
chlowticula s
ctystall~~era
egenula s (m,Ge.) .
egenlrloidea S
rarneu S
sguamellosa
vuria
sp. # l
The number of spores with an odd number of septa ia and Bacidifia is not random. The most reasonable ex-
are oRen significantly higher than the number of spores planation to this pattern is that, aRer the formation of
with an even number of septa in several species (Tab. the first, median septum, the subsequent septa arise
3 ) . This shows that the septation of the spores in Bacid- more or Iess synchronously in pairs. The synchroniza-
Conidiogenous cells
Bacidina
aenea
a~noldiana
nlifornica .
,hloroficula
crystallifera
egenula
egenuloidea
ramea
sqerarnellosa
varia
S/?. # I
a) More or less curved, non-septate (rarely l-septate), d) Straight to signoid, 0-l-septate, strongly congluti-
not conglutinated, 5-43 X c. 0.5-O.8(-1.0) pm (Fig. SA). nated and hence forming thick bundles, 35-49 X 0.8-1.2
This is by far the commonest conidial type in both Ba- pm (Fig. 8D).
cidia and Bacidina. 2. Oblong, ellipsoid, fusiform, or bacilliform, 0-l(-
b) Straight, non-septate, not conglutinated, 6-17 X 2)-septate, 3-12 X 1.2-2.0 pm (Fig. 8E).
0.5-1.0 pm (Fig, 8B). 3. Falcate (i.e., curved, thickest at the miaale and ta-
c) More or less curved, sometimes straight (variable pering towards the ends), 0-l-septate, 6-25 x 1.2-1.6
within the same species), 3-1 l-septate, not conglutinat- pm (Fig. 8F).
ed, 14-58 X 0.6-1.6 pm (Fig. 8C). 4. Acicular (distinctly thicker at the proximal end
Reaction with
Pigment Colour in pure water K C N
--
uni 2 uni 4 air + atr atr atr atr
0 - ~ ( ~ ~ ~ ~ ~ ~ ~ ~ 4 -
4-0-mcr
uni 4
+-zeo
--m m uni 3
% zco zeo
nor
nor
r e f a b c d e f g r e f a b c d e f g
--
atr i atr atr atr atr found in the species investigated: a in Bucidiu u g g ~ g a m l aB.
,
~ ~ ~ o o ~ * g & m polychroa,
" ' and B. schweinitzii, b in B. rubellu, B. russeola, B.
s u f i s a , and B, heterochvoa from the southeastern U. S., c in
uni 3
B. drflroctu, d in B. cumpulm and B. mutubiiis, e in two
+ zeo zeo spccirncns of Bacidia obsistrns from California and British
Columbia (Ca., Ekman L656, LD, and BC., 1908, ?Macoun,
CANL), f in one specimen of Bacidia absistens from
Washington (Foster 2888, MIN), and g (for comnparison) in
Norwegian and Swedish specimens of the same species.
Secondary substanccs are absent from all other species
investigated. Note that uni I and uni 2 can be separated only
in solvent system A. I am uncertain whether thc rclative order
between uni 3 and uni 4 in solvent system B is really correct.
My preliminary conclusion is based on the difference in
intensity of the spots. Unfortunately, there are no colour
differences. Uni 3 and uni 4, like all other orcinol rneta-
r e f a b c d e f g depsides involved, havc the same, deep purple hue.
idahoensis, B. polychrola, B. subincompta, and Bacidina and once on lignum, Bn. egenula once on bark and once
arnoldiana are predominantly corticolous, but have on rock, and finally Bn. egenuloidea once on lignum
been colIected on lignum a few times. B. suflusa, being and rock. Ba, egenuloidea, which is so far known only
predominantly corticolous as well, has occasionally from the type locality, is the only species treated here
been coIlected on rock. Among the remaining species, that has never been collected on bark. Some of the spe-
Bacidina chlo~oticulahas been collected once on bark cies can be expected on additional subtrates. For exam-
Bacidia:
absistens 4
aggregatula 4
arceutina: 16
auerswaldii 3
biaiorina 1
campalea 4
circumspecta 16
dlfracta 8
frieshnn 1
helicospora 2
hetemchma 40
hosthekoides I
igniarii 1
insularis 2
laurocerusi subsp. ia. 15
burocewi subsp. id. 12
mediulis 10
mutubilis
polychroa
reagens
2
21
3
.
roseliu 2
rubella 14
russeola 1
salmopaea 2
schweinitzii
subincompta
snff;lsa
54
10
16
. m
vermifera 5
Bacidina
aenea
arnoldiana
assulata
cul~ornica
chlomticula
ctystaIIifea
egenulu
egenuloidea
mmea
squarnellosa
varia
sp. # l.
American corticolous and lignicolous species of Bncidia and Bacidina. As "important" have been counted a phorophyte genus
with unknown phorophytes is large in many species. All species except Bacidina egenuloidea are known to inhabit kee-bark.
ple, Bacidia arceutina, B, polychroa, B. rubella, Bacidi- ties that combine high air humidity with a fair amount
na arnoldiana, Bn, chloroticula, and Bn. egenula have of sunlight, such as light forests or forest edges in
been collected on rock in western Europe. Bn. chlorot- swamps, river and creek valleys, and hill and mountain
icda is known to inhabit various anthropogenic substra- slopes close to the sea or near lakes or swamps. Only a
ta as well. few species thrive in deep shade, viz. Buddia heli-
The corticolous finds have been made on a variety of cosporu, B, schweinitzii, and Bacidina squa~ellosu,al-
trees, shrubs, and vines. Bacidia has been collected on though at least the two last are also found under lighter
altogether at least 128 and Bacidina on at least 27 pho- conditions. Similarly, only a few species have been
rophyte species. This is an underestimation, however, found on solitary trees, namely Bacidia rubella and B.
since about 30 % of a e collections lack any information subincompta. Bacidia insularis deviates markedIy from
on the substratum. An additional 20 % of the collections a11 other species in not preferring woodland, but chapar-
have information only on the genus to which the phoro- ral or scrub close to the sea. Contrary to the European
phyte belongs. These have been included in the figures situation, no species is regularly found on dust-enriched
mentioned above only when there is no other phoro- tree trunks (in avenues, in churchyards, on way-side
phyte determined to species that belongs to the same ge- trees e~c.).1n Europe, B, frjeslana, B. laurocerusi, B.
nus (and thus must constitute a phorophyte species dif- roseila, and, particularly, B. rubella are often found un-
ferent from all other although not determined to der such conditions. On the whole, the lichen flora of
species). dust-enriched tree-trunks appears to be much poorer in
The number of phorophytes known to be inhabited by species and individuals than the corresponding Europe-
Bacidia and Bacidina is, of course, to some degree de- an flora.
pendent on the number of collections available. Exam- Many woodland species favour oldgrowths, and are
ining the "common" species of Bacidia and Bacidina, particularly abundant on the rough bark of old trees. A
i.e., those known from more than ten localities, shows number of species, however, have been found on young
that they have been collected on a minimum number of substrata with little competition from other lichens and
phorophytes ranging from 9 to 54. Most species, how- thus appear to be fast colonizers, viz. Bucidia absistens,
ever, inhabit a smaller number of preferred phorophytes, B. arceutina, B. heterochroa, Bacidina chloroticula,
on which the bulk of the collections have been made Bn. cys~ullifera,Bn. egenula, BR. egenuloidea, Bn. ra-
(Tab. 7). mea, and Bn. varia. Apparently, the proportion of fast
The ecological demands of several species of Bacidiu colonizers is distinctly higher in Bacidina than in Ba-
and Bacidina overlap, with the result that some phoro- cidia. The behaviour of Bacidia absistens in North
phytes are more species-rich than others. The phoro- America is remarkable. It is encountered on young
phytes richest in Bucidia are Populus trernuloides (1 0 trunks or branches, sometimes even the thinnest branch-
species), A cer rnacrophyllum (a), Thuja occidenaalis es in the uppermost part of the crown of the trees. In
(g), Ulmus americana (S), Acer saccharurn (7), Frraxi- western Europe, on the other hand, it appears to be re-
plus nigra (7), Populus trichocarpa (7), Fagus grandifu- stricted to large tree-trunks in old woodlands. Possibly,
lia (3,and Liquidambar sfyrac~fia (5). The phoro- the North American populations have had to adapt to a
phytes so far known to be the richest in Bacidina are colonizing way of life, since the trunks of its favourite
F m i n u s caroliniana (3 species), Vitis rotzkndifolia (3), phorophytes are often completely covered with bryo-
Alnus mbm (21, Cephalunthm occiden~alis(Z), F a p phytes.
grandifolia (Z), Sumbucus canademis (2), and UrnbeC It is not uncommon to find one, two, or even k e e
luluria cal$ornica (2). Considering that the species of species of Bacidia and Bacidina in a single locality. A
Bacidiaa are extremely overlooked, this list will proba- higher number is unusual, however. The richest locality
bly soon have to be updated. It should also be noted that known to me is the Burden Research Plantation, East
regionally, other phorophytes than the ones mentioned Baton Rouge Parish in Louisiana, which hosts three
may be the most important. However, the most species- species of Bacidia and four species of Bacidina.
rich phorophytes mentioned altogether house 20 out of
27 species of Bracidila and 9 out of 12 species of Bacidi-
nu. Phytogeography
Habitat Many of the species of Bucidia and Bacidina exhibit
distinctive distribution patterns on the North American
The habitats of Bacidia and Bacidina cover a range of continent. Most of these are previously well-known
woodland from sea level to an altitude of about 3000 fiom other lichens, bryophytes, and vascular plants. The
meters (Bacidia subincompta in the Rocky Mountains causaI relations behind these patterns have been dis-
of Colorado and Arizona). Many species favour locali- cussed elsewhere and are generally not repeated here
Boreal
Temperate
Pan-Noi-tll Ame~-ictun
Easl ~eniperale- Appalachian
Occanic
North Pacific coast
Appalachian + No1-111Pacific coast
Coastal Phi11
T~.opical
Suutl~Pacific const + Coastal Phin
Coaslal Plain
Soulh Pacific consl
Total 28, 12
(BiAaun1955, Irnshaug & Brndo 1966. Schotield 1969, (sub)element has not been previously suggested. This
Gowan 1983). concerns the Californian elerncnt and the southern Pa-
It is important to notc that no species treated here has cific coastCoastal Plain subelement of the tropical ele-
a distribution limited by the availability of suitable pho- ment.
rophytes. Instead, the dishbutions are li~nitedby vari-
ous other ecological, climatic and historic factors. The The boreal elerncnt
classification of the distribution patterns used here has
beell adapted from Brodo (1968). Brodo & Gowan Bacidin ,st~hittcornptuand probably aLo B. vc.rm&?ru
(1983), Brudo & Hawksworth (I977), Gowan (1983), bclong to the boreal element. They constitute 7 % of the
Gowan & Brodo (1988), and Hale (196 1). Representa- taxa uf Bacidia. Both species occur in Europc as well.
tives of the arctic and maritime elements do not occur As pointcd out by Brodo & Hawksworth (19771, the bo-
among the Noilh An~cricanBacidia and Bocidinu. The real element is difficult to interpret on a Noid1 Amcri-
species-richness and the e n d e ~ n i s ~liavc
n been summa- can scale due to the effects of glaciations and subse-
rized in Tab. E. quent recolonizations. Both species, however, arc
Fairly reliable I~ypotheses about distributional pat- coniferous forcst species, mainly confined to Popltlus
terns can be made ror 24 of thc 27 species of Bucidiu /renzuluides. Urodo & Gowan (1983) distinguished be-
and 7 of the 12 species o r Bocidina. The rc~nainingX tween a ilortlem and a southern boreal distribution
spccics are too poorly known to permit crediblc inter- type. B. szihi~zca~nptaexhibits a more or less typlcal
pretations, although educated guesses can bc inadc in southern boreal distiibutioil type. With hesitation, I
most cases: Bacidia hialorinu (north Pacific coast sube- have placed Rircicliu verpnifi~ain thc boreal element al-
lement?), B, ji-iesiana (ditto'?), B. ipiurii (boreal?), Bn- though it is not present in the true borcal zone. Instead,
ciditlu ussnrl(i~a(Ozarkian or, as in Europe, continen- it occupics simikar habitats in the Rocky Mountains and
tal'?), Bn. chlurutiuulu (?l, Bn, crys~ullije~a(Coastal in the Cascades, and can be said to have a "westcm
Plain subelenlent o l the teinperatc cle~nent'?j, Bn. montane" distributiun type. Possibly, it was displaced
egenula (eastern temperale subclcincnt?), and Bn. southwards during thc glaciatons, and has not (yet) rein-
egenuloidea ('!). vaded the northern territories.
Exan~plesother than specles of Ruuidiu and Bacidirla
have been included in the accounl only when the
The temperate element The oceanic subelement
The temperate element is the largest one, constituting Two species, Bacidia arceutina and Bacidina arnol-
50 O/n of the Bacidia aiid 17 % of the Bacidina. The en- diana, can be referred to the oceanic subelement. Bo-
demism is 36 % in Bacidia and 50 % in Bucidina. The cidia arceutina is bicoastal with very few finds on the
latter figure, however, is probably strongly distorted by east coast (none of which have been confirmed in xe-
the fact that that several species of Bucldina have been cent time), whereas Bacidina arnoldiaaa is known
excluded from the analysis. The temperate element can froin humid areas of the Pacific Northwest and the
be divided into a large number of subelements, seven or southern Appalachian Mountains. Neither of these
eight of which are represented among ilie Bacidia and species is endemic.
Bacidina.
tropical species B. canapalea, B. jksconigrescaas (Nyl.) localities are situated in the Great Lakes-Appalachian
ZahIbr, and B. millegruna (Taylor) Zahlbr, than to any regions, but there are a few outliers between the Coast
temperate (Asian, North American, or European) spe- Range and the Cascades in the northern Pacific region.
cies that I know of. Bacidia rubella is known from Europe as well.
MPAC SPAC MONT BORE GLNA OZAR SAPP COPL FLPE BRIS SSWE NSWE
absistens I 1
aggregatula 0. 0
arceutina 1 I
nuerswuldii I 0
biatorinla 1 0
caesiovirens 0 0
carnpalea 0 0
circumspecla 1 1
dlfirncta 0 0
fraxinea 0 0
friesianu 1 0
helicospom 0 0
heterochroa 1 1
hostheleoides 0 0
iglniarii 0 0
incornpm 0 0
insularis 0 1
Inurocerasi 1 0
medialis 0 0
mula bilis 0 0
polychrola 0 0
reagens 0 1
rosella 0 0
rubella I 0
russeoh 0 0
salmonea 1 0
schwrinitzii 0 0
subcircurnspecta 0 0
subincompaplu I 0
suffisa 0 0
vermifem 0 0
(1) NPAC: Northern Pacific coast region, including south to the level of Washington D. C., plus the area
the coast and the Coast Range from central California south of the Great Lakes, south to, but not including,
(north of Marin Co.) to southern British Columbia, plus the Appalachian Mountains and the Ozark Plateau.
low elevations (up to c. 1000 m) on the western slopes (6) OZAR: The Ozark Plateau.
of the Cascades and the Rocky Mountains. (7) SAPP: The southern Appalachian mountains, i.e.,
(2) SPAC: Southern Pacific region, including the the part of the Appalachian mountain range not includ-
coast and the Coast Range in southern CaIifomia north ed in 5.
to (and including) Marin Co. (8) COPL: The Coastal Plain froin Texas to New
(3) MONT: High elevations in the Rocky Mountains, York (except southern peninsular Florida), plus the val-
the Cascades and the Sierra Nevada of western United leys of Mississippi and Tennessee Rivers north 'to south-
States. em Illinois.
(4) BORE: All of Canada and Alaska except the areas (9) FLPE: The Florida Peninsula south of a west-east-
belonging to l or 5. erly line across Tarnpa.
(5) GLNA: The Great Lakes and the northern Appala- (10) BRIS: The British Isles.
chian region, including the Great Lakes, the area imme- (1 1) SSWE: Southern Sweden south of the northern
diately north of the lakes, eastwards to the coast of New distribution limit of Quercus (limes norrlandicus).
Brunswick and Nova Scotia, southwards along the coast (12) NSWE: Northem Sweden north of limes norr-
to New York and along the Appalachian Mountains lundicus.
(15) BRIS
(15) SSWE
( l I ) NPAC
( 6 ) SPAC
(3) MONT
(5) NSWE
(6) BORE
( l 0) GLNA
(6) SAPP
( 5 ) OZAR
(10) COPL
(7) FLPE
Fig. 10. Singlc linkagc clukter analysis o r nine North Arncrican and threc European regions regardii~gspecies composition of the
Baridiu flora. Thc simple lnatchiilg dicholomy (54) coefficicnt w a ~used to calculate similarities between each pair of regions,
using tl~chinary matrix in Tab. 9. Distances bctwccn clusters are E~lclidcan.The abbreviations and delimitations o f the rcgions
(WPAC etc.) AI-eexplained in the tcxt. Thc n~umherof species vf Baci(liu presently known from each 1.cgio11is prcselitcd withiii
brackets.
The choice of the three extra-North American regions efticient measures similarity as thc proportioi~of com-
was bascd on knowledge. The Bacidia flora of these re- mon presences plus coinnloii absences, i.e., (no,,+ n, ,)i
gions is fairly well-known, the British Islcs through the (no,+ nu,+ n,,+ n,,), where n,, is the number of species
work of Coppins (in Purvis et al. 1992) and Sweden absent from both regions, n,, the number of species
through my own studies. Four species occur in thc Euro- prcscnt in both regions, and n,, and n!, the number of
pean regions but not in North Amcrica: Baciclin caesio- spccics prcscnt in only one the two regions. Finally, the
vir-ens S. Fkman & Holieil (Ekn~an& Holien 1995)- R. resulting similarity matrix was used in a single linkage
,fi-uxinea Lonnr. (Ekman & Nordin 1993), Bcicidiu in- cluster analysis.
cu117ptu(Rorrer ex Hook.) Anzi, and B. subcircupns~~eo/a
Coppins (regarding thc two latter. see Coppins et al.
1492 and Coppins in Purvis et al. 1992). Note that Ba-
cidia igniarii has recently been reported for the British
Isles (Purvis et al. 1993). The 27 North American spc- The cluster diagram is shown in Fig. 10. MONT,
cies and 4 additional European species were scored as BORE, and NSWE are the regions that are most similar
present ( I ) or absent (0) in each of thc twelve regions to each other. SAPP and OZAR arc also fairly similar,
(Tab. 9). The similarity in specics composition between as are BRIS and SSWE. FLPE is very different Crom all
each pair of the regions was then calculated using the other regio~ls.NPAC is inore similar to BRTS and SS-
simple inatclling dichotoiny (S4) coefficient. The S4 co- WE than to any other North Anlcrican region. Apart
Bacidia Bacidina
Thallus structure Never dissolved into goniocysts Often dissolved into goniocysts
Proper exciple inside rim Composed of thick-walled hyphae Composed of rather thin-walled
with narrowly cylindrical cell hyphae, most of which have cell
lumina lumina that are broadly ellipsoid,
almost globose, or irregular
Axial body I 1 M of the entire width of the >l13 of the entire width of the
d-layer, vertically not reaching d-layer, in some species vertically
through this reaching all through this
mina varying from narrowly cy1indrical to almost glo- to each other, about as closely as Bacidia and Tondniu.
bose, but at least partly short and wide (even inside the Bacidia, Bacidina, Toaiaia, and Arthrospomrn proba-
rim), f ellipsoid, globose, or irregular (width > 2.5 p, bly together form a rnonophyletic group. Unfortunate-
length > 4 times the width, often distinctly constricted ly, the differences between Bacidia and Bacidina have
near septa. Walls gelatinized, the distance between two often been oversimplified and exaggerated. For exam-
adjacent lumina usually less than the width of a lumen. ple, Bacidina has repeatedly been stated to have a
(5) Hypothecium not chondroid, of distinctly different "paraplectenchymatic" proper exciple contrary to a
texture than the proper exciple and distinctly delimited "prosoplectenchymatic" one in Bacidia. Some species
from this. of Bacidina have an entirely "paraplecteilchymatic"
(6) Paraphyses unbranched or more or less branched in proper exciple, but several species have a mixture of
upper part, not anastomosed, without a thick, gelati- narrow and wide cell lumina. On the other hand, some
nized wall in upper part. species of Bacidia have enlarged cell lumina along the
(7) Hymenium and subhymenium arnyloid, sometimes rim of the proper exciple. Thus, the real difference be-
pigmented in'a f diffuse layer in upper part tween Bacidia and Bacidinu appears to be that Bacidi-
(8) Tholus amyloid, with a narrow or wide c-layer, with n a has at least some wide cell lumina in the inner of
a high or low, blunt axial body (sometimes penetrating the proper exciple, whereas Bacidia does not. Further-
through the entire d-layer), the width of which is more more, it has been stated that Bacidia has "Bacidia
than a third of the entire d-layer; ocular chamber absent type" asci and Bacidina "Lecanora type" asci. ,As I
or present, low or high, blunt or pointed. have indicated earlier, I object to such a statement,
(9) Spores acicular, when mature with three or more m s - since there are no generally accepted definitions of
venal, distinct or indistinct septa. They are colourless, these "ascus types", but also because such a terminolo-
rather thick-walled, and lack perispore or ornamentation. gy can make even minor differences sound fundamen-
(10) Pycnidia with cylindrical conidiogenous cells (4-8 tal. As far as I can understand, the tholus structure in
p long) that produce single conidia terminally. Conid- Bacidia and Bacidina is essentiaIly the same, but the
ia usually filiform and curved. width of the axial body relative to the d-layer is larger
(l l) Contains no lichen substances. in the latter genus. The characters that can be used to
separate Bacidia from Bacidina (including the ones
discussed here) are summarized in Tab. 10.
The relationship between Bacidia and
Bacidina
Cliostornum Fr.
Ecfiidnocymbium Brusse
E. M. Fries 1825: l l 6
Brusse 1987a: 173.
Type: C. corrugurzlm (Ach.) Fr. (leclotype selected by
Timdal 1991). Type: E, spc.cio.~umBrusse (holotype, not seen).
Presently, seven species are recognized in Cliustumutn, Eclridnocjnahium, comprising only the type species, dif-
C, corrugntuni (Ach.: Fr.) Fr.. C. grijlithii (Sm.) Cop- fers from Bacidia and Bacidiraa in having Iecauori~~e
pins, C. lc~;lrc,sum(Rasanen) HoIien & Tensberg (Tans- apothccia, inore abundai~llybranched paraphyses, non-
berg 1992), U, vi!ellinzlm Gowan (Gowau 1990): C. ,flu- septate ellipsoid spores, black and large pyciiidia that
viduhint Hafellner & Kalb {bib & I-Iafellncr 1992). C. are superficial on the thallus, needle-like conidia, and in
sper.mogoniaturn {Zahlbi-.)Kantvilas & Elix, and C. vez- containing stictic acid. The genus was, in my opinion,
due Kantvilas & Elix (Kantvilas & Elix 1995). C. Irlle- referred to the "Biatoraceae" on very dubious grounds.
olnrm Gowan was considered a synonym of C. Iepro.sum
by Tansberg (1992). The boundaries of Clinstnmurn are
still poorly studied, and the genus will almost certainly Eschatogonia Trevis.
"grow" in the future. For example, "Bacidia" pcdIlens
probably belongs here (see E , ~ c / ~ d e01.
d not e.~amined Trevisan 1853: 5.
species).
Uliostomtrm can be separated froin Bucidiu and Ba- Type: E. prolifirra (Mont.) R. Sant., treated as E. mon-
ciditta on account o f the widc and ii-rcgularly shaped zugneL by Trevisan, a superfluous nornei~clatural syno-
cell lumina in the prupcr cxciplc, the often crystal-in- nym (holotypc).
spersed epitheciurn and proper exciple, the l -3-septate
bacilliCorm spores, the branched conidiophores bearing According to Brako (1989) and Kalb & Elix (1 9951, this
subgIobose to ellipsoid to short-bacilliforrn conidia. genus complises a few tropical species that dlffer from
Two of the species contain usnic acid in the thallus and1 Bucidiu and Bacidino in being squamulose, in having a
or the apothecia. Black and widely gaping pycnidia arc lower cortex with a distinct outer layer of unifonnly ar-
characteristic of some species. Clinstornur~zis closdy rc- ranged cells, and i n containing didymic acid and related
lated to Biatom, but the latter genus differs in having substances.
sparsely branched, ampulliform (not cylindrical) conid-
iogenous cells and in having cylindrical, almost parallel
and distinctly radiating txcipular hyphae. Berteliana P. James
Squama~qina,comprising almost 20 species (Poelt 1958, According to Hafellner & Vgzda (1992),this monotyp-
Poelt 1975, V k s k a 1985, Haugan & Timdal 1992), is ic genus differs from Bacidia and Bacidinra in forming
distinguished from Bacidiu and Bacidina on the basis of large, byssoid thalli comprised of thick hyphae. Fur-
the squamulose or almost foliose thallus, lecanorine thermore, the terminal part of the excipular hyphae are
apothecia, and non-septate spores. According to Hertel ungelatinized, the hymenium is unpigmented, the para-
& Rambold 1988: 301), the tholus of S, gypsacea con- physes lack terminal swelling, and the spores are ellip-
tains a tubular structure similar to that found in Porpid- soid and l-septate.
iaceae or Psoraceae sensu Hafellner (1984). These find-
ings were confirmed for sect. Sqzkamarina by Haugan &
Timdai (1992), whereas sect. Petropiucu was found to Toninia A. Massal.
have a tholus resembling the one seen in Lecanora mu-
ralis. Squcdmarina was listed in Bacidiaceae with a que- Massalongo 1852: 107.
ry by Eriksson & Hawksworth (1993). Its placement
near Bacidiu appears very doubtful. Type: T. cinereovirem (Schaer.) A. Massal. (lectotype,
selected by Fink 19 1.0).
- I..
-
. R
. I
0
0 cP
Oo 0
0 f (APWID, APHGT, EXCIP, HYM, PARWID, SPLEN,
SPWID, SEPTA)= 0.0155xAPHGT + 0.00434xHYM +
0.00538xEXCIP + 0.0180wSEPTA - 4.05xAPWID -
0
2.00xPARWID - 0.127xSPLEN - 0.343xSPBRD + 7.90,
- 00
0
characters studied separate the morphs entirely discon- Apart from thallus structure, it is clear that the continu-
tinuously. ous, Bacidia polychroa, and the granular morph, B. dif
The canonical variates plot is presented in Fig. 11, fracta, are separated by a discontinuity in the correlation
and the canonical loadings in Tab. 12. The continuous among a number of characters. 1 conclude that they
morph forms a distinct swarm, whereas the granular should be recognized as distinct species. The position of
morph consists of a fairly distinct swarm with a few the Florida population is slightly more uncertain. Since
outliers. The two Floridan specimens from the same lo- there is rather conspicuous variation within the Florida
cality are rather different from each other, but are only population, and since it is only slightly more extreme
slightly more extreme than the two outliers of the granu- than the extrernest of clear B. dgffracta, I draw the tenta-
lar group (from Minnesota and Massachusetts, respec- tive conclusion that it is best included in B. d~@acta.
tively). The discrimination along the first canonical var- More material is needed, however, since a recognition
iate is perfectly discontinuous, which shows that there is of the Florida population at an infraspecific taxonomic
level may prove to be the best solution. Unfortunately,
my own attempts to find this morph in the field have
Tab. 12. Loadings for the eight characters on the first and been unsuccessful.
second canonical variates (X 100) in a CVA performed to Since the discrimination along the first axis is per-
investigate the distinctness of Bucidia polychma, B. dzflacta, fect, the value of the function defining this axis can be
and a deviant population from Sanford, Florida. The sign
shows whether the character makes a positive or negative used for identification purposes. This may be useful in
contribution to the canonical variate. case there are difficulties classifying the thallus of an
unidentified specimen as granular or continuous. For B.
Canonical varinte polychroa P 0, and for B. dzfructa B 0.
Character I II
APWID
APHGT Bacidia laurocerasi
EXCIP
HYM
PARW ID Background
SPEEN
SPWID B~cicdialaurocerasi S. lat. has a disjunct distribution in
SEPTA North America. It occurs in the coastal lowland and in
the montane rcgioi~of Oregon, Washington, and British Pairwise comparisons of the three distribution groups
Columbia, but also in the Great Lakes and northern Ap- were made. For the qualitative variables THALL and
palachiai~ region of ilortheastern United States and MARG, 1 used Yates' corrected chi-square, but 1 inadc
southeastern Canada. Wlien studying material, I soon the test Inorc coi~scwativeby dividing the ordinary sig-
became aware of slight differenccs in thallus structure, nificancc cut-off values by three (i.e., tlie number of
apothecium colour, and sporc scptatioi~between collec- compa~.isonsmade) in order to compensate for the mul-
tions from the two distributio~~ areas. Therefore, I decid- tiple comparisons. Tlie remainder of the characters are
ed to examine this casc closer. I also included a smalI quantitative and were studied with one-way ANOVAs
numbcr of European specimens in the study in order to using the Scheffk's S proccdurc for post-hoc multiple
see which one of the two potential North American taxa mean comparisons.
is inorphologically closest to the European population. All characters except EXClP werc uscd i l l a PCA per-
formed to investigate thc distiilctncss of the lhree distri-
bution groups.
Methods
and European groups is larger than the overlap between beleive that it is justified to recognize the former taxo-
the western and any of the two other groups. nomically. Since all characters are overlapping, I find
the subspecies level to be the most appropriate one.
There is a name already available for this taxon at the
Conclusions species level, Bacidiu idahoensis H. Magn. This name
is combined here to the subspecies level under B, luu-
Since there are obvious differences between the western rocerasi.
North American group on the one hand, and the eastern The relationship between the European and the east-
North American and the European group on the other, I ern North American groups is more difficult to evaluate.
Despite the low number of specimens studied of the Eu-
ropean group, they differ significantly in two characters,
and there is a clear separation in the PCA. It should be
Tab. 14. Distribution of two qualitative characters (given as noted, however, that all characters are widely overIap-
number of specimens) in Bacidia laumcerasi from western ping, that the significant differences are few, that the
and eastern North America, and from Europe (represented by overlap in the PCA is large, and that European,B. iuu-
specimens from Sweden, Germany, Switzerland, and Great rocerasi is still insufficiently studied. This leads me to
Britain). The abbreviations of the characters are explained in
the text. the conclusion that presently, the best arrangement is to
include both the eastern North American arid the Euro-
pean representatives in the concept of subsp. laurocera-
Western Eastern si. This conclusion must be considered as tentative,
North America North America Europe however.
W= 8) (N=21) (N= 8) The reason why 1 have included so few specimen of
THALL European B. laurocemsi is that it has been considered
cracks few or lacking 1 20 8 peripheral to my work. Naturally, further studies of the
densely cracked 8 1 0 European group and its relationships with other parts of
the world are necessary. Furthermore, when European
MARC B. ~uurocerusihas been studied in more detail, it may
brown to black 2 21 8 prove to contain a significant amount of geographicaIly
brown-red 7 0 0
correlated variation.
Fig. 13. Principal cornpol~e~~ts
plots o r 38 specimens oCBucirIi~ilaumwrasi S. lat Irom western North Ainerica (squares), eastern
North Amcrica (circlcs), and furope (triangles). The first three components (PC 1-111) account for 77 % of the total variance.
One-way ANOVAs were performed on each of these 0.05) in apothecium height, spore width, and spore sep-
characters. APHGT, HYM, SEPTA, SPLEN, and SP- tation. None of the characters studied separate the
WID were used in a PCA in order to study the distinct- groups discontinuously.
ness of the hvo population groups. The PCA plots, in which the specimens have been
plotted against the first three principal components, are
shown in Fig. 14. The component loadings and the
Results amount of variance explained is given in Tab. 17. In the
PCA plots, the western and the eastern popuIations are
Descriptive statistics and the significance of F in the separated, although not: di~continuously.They are main-
ANOVAs are presented in Tab. 16. The western and ly separated by the first principal component, to which
eastern population groups differ significantly (i.e., p< all of the included characters contribute markedly.
Tab. 16. Descriptive statistics and the significance of the F-values obtained from the one-way ANOVAs comparing western and
eastern populations of Bucidia hetemchroa. The abbreviations of the characters are explained in the text. The standard deviation
is abbreviated "S. d." Note that the input values used for APWID, HYM,and PAR are means of five measurements per specimen,
for SPLEN, SPWID, and SEPTA means of 10 measurements per specimen, and for APHGT and EXCIP a single measurement per
specimen. Thus, the values of all but the two latter characters correspond to the minimum and maximum mean values in the
species descriptions. Significance levels of F are denoted * for 0.011 p20.05 and ** for 0.001< p50.01, and ns (not significant) fnr
p> 0.05.
APWIDlmm
APHGTJpn
EXCIPJp
HY M / p
PARWID/p
SPLEMlp
SPWIDlpm
SEPTA
cidia! Some of the taxa included are treated in Excluded 10. Asci with 1-16 spores (usually S), without pronounced
or not examined species. swelling at the apex. A+b-layer of tholus not thickened
Apart from my own observations, I have used the fol- terminally ................................................................... 11
lowing sources of information to construct this key:
Coppins (1983), Timdal (1984), Egea & Torrente 11. Apothecia with poorly developed proper exciple, im-
(1993), Liicking et al. (1994), Printzen (19951, Roux et marginate. Paraphyses and excipular hyphae similar,
al. (1995), and Staiger & Kalb (1995). abundantly branched and anastornosed. Photobiont
usually "micareoid", i.e., fungal hyphae penetrating al-
1. Photobiont T~ntepohliu................................................ 2 gal celIs by intracellular haustoria .................... Micarea
1. Photobiont a green member of the Chlorococcales ......5 11. Apothecia with f well-developed proper exciple, visi-
ble as a distinct margin at least in young apothecia.
2. Apothecia pale yellowish to dark brown; disc concave Paraphyses and excipular hyphae often not similar.
with a distinctly raised margin .....................................3 Photobiont not "micareoid ........................................ 12
2. Apothecia black (although sometimes pruinose); disc
plane to convex ............................................................. 4 12. Asci l-4-spored. Spores with walls c. 2 pm thick .........
................................................................... Meg~lospora
3. Most or all asci with more than 8 spores ... Pachypkiale 12. Asci with 8-16 spores. Spore wall thinner .................13
3. Asci with 8 spores ............................................. Gyalecta
13. Thallus PD+ yellow to orange, K+ vividly yellow (with
4. Asci difficult to separate from ascogenous hyphae when thamnolic acid) ............................................. Loxospora
squashed in K. Spores not fragmenting ......................... 13. Tnallus often PD- and K- to K+ faintly yellow (without
...................... . . . .....................................Lecanactis thamnolic acid) .......................................................... 14
4. Asci easily separated from ascogenous hyphae whem
squashed in K. Spores fragmenting in some species ..... 14. Tholus without axial body but with a strongly amyloid,
................................................................... Bactrospom towards the tip slightly widened tubular structure.
Proper exciple paraplectenchymatic or byssoid (but
5. Thalius squamulose ....................................................... 6 sometimes encrusted with crystals) ............................ 15
5. Thallus crustose, or if squarnulose with pale pink apoth- 14. Tholus with + conical, cylindrical, or lens-shaped axial
ecia .............................................................................
9 body that is sometimes surrounded by a strongly arnyloid
zone. Proper exciple proso- or paraplectenchyrnatic ..... 16
6. Squamules very small, almost granular, usually not ex-
ceeding 0.2 mm in width. Apothecia orange-brow. 15. Proper exciple f byssoid, at least in outer part ..............
Not sorediate ................................... "PsoreHa" pertexta ...................................................................... Byssoloma
6 . At least some squarnuIes exceeding 0.5 mm in width. 15. Proper exciple 'hormal", i.e., composed of hyphae
Apothecia brown to black. Some species sorediate ..... 7 with -t- gelatinized walls ................................ Fellhanera
20. Spores 3-septate, bean-shaped. Upper part of hy- Key to corticolous and lignicolous species of
menium green to blue-green ........................................... Eacidia and Bacidina
............................................... Arthmsporum populorerm
20. Without this combination of characters ..................... 2 1 l . Upper part of hymeniurn green, blue-green, or green-
grey, N+ red to purple (sometimes with a precipitate of
21. Rim of proper exciple covered by a rather thick gelati- blue crystals) ................................................................. 2
nous layer that swells markedly (and finally almost dis- I. Upper part of hymenium colourless to yellow to orange
solves) in K or a dilute hypochlorite solution. Spores to brown to dirty purplish, reaction with N variable .....
with warted perispore in some species ........................... ..................................................................................... I1
................................. The 'IBacidia" sabuletorum group
21. Rim without such gelatinous layer. Spores without 2. Upper part of hyrnenium grey to grey-green, C+ violet.
wartad perispore .................... . . . ............................. 22 ................................. .... ......................... 19. B. reagens
2. Upper part of hymenium green to blue-green, C- ...:..... 3
LL. Excipular hyphae of mature apothecia moderately
branched, running closely parallel to each other; cell +
3. Proper exciple laterally with evenly dispersed, minute
lumina narrowly cylindrical, evenly thick, not marked- crystals (mostly less than 1 pm) ............. 1. B, absistens
ly widened near rim of proper exciple ....................... 23 3. Proper exciple without crystals or with radiating clus-
22. Proper exciple different ........................... ....... 24............ ters off coarse crystals (mostly 1-7 pm) .................. A
23. Apothecia when very young almost globose, with a dis- 4. Hypothecium coIourless or very pale yellowish/brown-
tinct core of young, immature asci and ascogenous hy- ish .................................................................................. 5
phae surrounded by a distinctly delimited proper exci- 4. Hypothecium orange to brown ................................... 7
ple .......................................................... Mycobilimbia
23. Apothecia when very young moderately convex. Imma- 5. Spores acicular. Apothecial margin finally excluded .....
ture asci and ascogenous hyphae dispersed in the ster- ................................................................ 9. B. friesiana
ile tissue, not forming a distinct core distinctly delimit- 5. Spores baciiliform or clavate. Apothecial inargin usual-
ed from the proper exciple ................................. Biatora ly persistent .................................................................. 6
6. Young apotliecia low and flat. Proper exciple mainly 13. Thallus often dissolved into goniocysts. Proper exciple
consisting of regularly radiating hyphae ........................ at least in outer pc>~-tiunwith wide, ellipsoid to globose
.........................................................7. B, uircumspectu cell lumina ............................................ 38. Bn. variu
A. Young apothecia barrcl-sliapcd. Proper exciple below 13. Thallus never dissolved into goniocysts. Proper exciple
hypothecium forlning a thick cushion of irrcgulnrly in- of k cylindrical cell lumina ....................................... 14
tertwined and licl~lybranched hypluac .... 13. B. igntarii
14. Apothecia usually pink. Cell lumina widcr in lower
7. Hypotheci~rmbrown, downwards gi-adually mclging in- (oId) part of proper exciple llian ill upper (young) part
to the dark brow-red to red-black proper exciple ........ ................................................................
16. B. )11edidi,s
...........................................................
24. B. schwuinitzii 14. Aputhecia usually brown-orange to k pi~rple-brown.
7 . Hypotliccium brown in a L distinct zone, in the lower Cell lumina in luwcr part of proper exciple narrowel-
part irsually changing to colourless or yellowish (not than or as wide as Iirmina in uppcr part .........................
gradually merging into a dark proper cxciple) .............8 ......................................................
12. B. hosfheleoidrs
8. Spores acicular, narrower illan 2 ,WII. At lcslst lower part 15. Upper part of hymeniuiii dirty purplish, K+ pure green,
of proper exciple with f ellipsoid cell lumina. Hy- N+ pure purple (oflen with a precipitate of bluc crys-
pothecium K- or K+ greenish brown ... 34. Bn. egen~rla tals). Proper exciple inspersed will1 minute ciystals (<
8. Spores bacilliform, fusiform, clavate. or aciculal-, width 1 W) ............................................
1 . Brrcidio ubsi.siuns
always cxcceding 1.5 pm and ~rsually also 2.0 pii. 15. Upper part of hymenium colourless to yellow to orangc
Cell lumina in proper exciple (outermost cell layers to bruwn; rcactions with K and N variable but not K+
excepted) cylindrical. Hypothecium K- or K+ pm- pure p e e n and N+ pure purple. Proper exciple with or
plisli ...............................................................................
9 without lllir~ulecrystals ...........................................
16
9. Proper exciple, at least rim, dominated by a green to 16. Hypothecium andtor proper exciple immcdiately below
blue-green pigment (K-) ........................14. R. insuluris hypothecium dark (brown-orange, orange-brown, red-
9. Propcr cxciple usually dominated by a ]3rowii, K-I- pur- brown, dark brown, or red-black) ..............................17
plish pigment. Some L pigment deficient speciincns 16. Hypothecium and proper excipIe below hypothecium
dominated by a grccn to blue-green pigment, bui then pale (colourless. palc yellowish, pale orange, or pale
in low concentration and only in upper part of proper brown)
, .....................................
. . . 2l
exciple .........................................................................
10
17. Hypothecium K+ pccn-brown. Thallus with c. 20-50
10. Hypotliecium coluurless or palc orange to brown-or- pn wide goniocysts .........................29. Bn. arnolrbiunu
ange, K-. Spores (average of 10) 1.7-3.2 pn wide, nev- 17. Hypothecium K-, K i intensifying, or K+ purple-red.
er with more than 7 septa. Apothccial margin usually Thallus smooth or granular, granules then mostly ex-
persistent .......................................... 7. B. circ.cun?specla ceeding 50 pm ......................................................... I S
10. Hypothecium pale to dark (often reddish) bmwn, K+
purpIish. Sporcs (average of 10) 2.1-5.1 pm wide, with 1 X. At least young apothecia with abundant, +evenly dis-
3- 13 scpta. Apothecial margin finafly excltrded ............ perscd minute crystals in the epitliecium and rim of
.................................................... 25. B. srihinuornl~fr~ proper exciple (usc poIarizing filters!) ...........................
...........................................................
2 . B, oggregiztuln
1 1 . Spores bacilliform 01. fusiform, straight or only slightly
curved, not coiled in the ascus, lengthlwidth ratio (av- 18. Apotliecia without crystals or with radiating clusters of
erage of 10 or more spores) ncvcr cxceeding 11 (often crystals in the proper exciplc ......................................
19
much less) ............................. .......................
12
11. Spores usually acicular, straight to curred to sigmoid, 19. Brown pigment in apothecia K+ intensifying ................
often coiled in the ascus when inlmaturc, lcngthlwidth ......................................................
24. B. sohweiniizii
ratio (average of 10 or more spores) exceeding 10. IJ' 19. Brown pigmenl in apothecia K+ pu~plc-rcd.............. 20
lengthlwidth ratio not exceeding c. 10 and spores bacil-
lifolm to clavatc, thcn spores coiled in ascus, distinctly 20. Thallus smooth, wrinkled, or waned .. 1 X. B pcjlj~chrou
curved or sigmoid ................................................
14 20. Thallus finely granular .............................
8. B. difpacta
12. Spores firsiforln with blunt cnds, width exceeding 5.5 21. Apothecia (when frcsh) pink, pale yellow, pale grey,
p. Western ........................................
4 B, utlerswulrl'ii or almost while (N.B.! As herbarium specimens age,
12. Spores bacilliform, width never exceeding 5.0 W. pale apothecia often darken to ncar mcdium-orange) ..
Southeastern .....................
. . ............................. 13 .....................................................................................
22
2 1. Apothecia at least partly brown-yellow or k orange to 31. Most young apothecia with a thin thalline margin (visi-
red-brown to black ..................................................... 33 ble at least in section), which is soon suppressed by the
proper exciple. Axial body lens-shaped .. 36. Bn. mmea
22. Epitheciurn and/or proper exciple with crystals ........ 23 3 1. Apothecia without thalline margin. Axial body f coni-
22. Epithecium and proper exciple without crystals ........ 27 cal ................................................................................ 32
23. Proper exciple with coarse crystals (58 pm diarn.) in 32. Hymenium (average of 5) 57-73 pm high. Paraphyses
lower part and (usually) minute crystals (21.5 pm di- fairly abundant in reIation to number of asci. Conidia
m.)in upper part; rim usually with stronger pigmenta- bacillifonn or filiform, the latter non-saptate. Califor-
tion in lower part than in upper. Margin distinctly paler nia .................................................. 3 1. Bn. californica
than disc, usually pinkisll or pale brownish. Extreme 32. Hymenium (average of 5) 42-61 pm high. Paraphyses
South ..................................................... 6. B. campalea rather few in relation to number of asci. Conidia fili-
23. Proper exciple either with clusters of coarse crystals or form, usually 3- I I-septate. Southeastern .......................
with t. evenly dispersed minute crystals; rim with .....................
. . . ................................ 3 8 . Bn. varia
stronger pigmentation in upper part than in lower. At
least upper part of margin (closest to hymenium) usual- 33. Disk of at least some young (not seldom all) apothecia
ly concolorous with disc or only slightly darker or pal- brown-yellow or yellow-brown, without tinge of red.
er .................................................................................. 24 Rim of proper exciple K-, in upper part brown-yellow,
yellow-brown, or almost pure brown, downwards usu-
24. Proper exciple along the rim with a distinct, 4-6 cell ally gradually changing to pale yellow or colourless ....
Iayers thick zone of enlarged cells .......... 26. B. sufusa ................................................................................. 34
24. Proper exciple along the rim without or with a 1-2 cell 33. Apothecial disk at least partly orange to reddish brown
layers thick zone of enlarged cells ............................. 25 to black. Rim of proper exciple at least in part f orange
to pure brown to red-brown to purple-brown (when k
25. At least margin of young apothecia with a thick white brown K+ green-brown or K+ purplish) .................... 35
pruina. Margin persistent ................... 17. B, mutabilis ..
25. Apothecia without or with thin pruina. Margin finally 34. Thallus not dissolving into goniocysts. Proper exciple
excluded ................................................................... 26 (the outermost cell layer excepted) of cells with f nar-
rowly cylindrical lumina. Bicoastal ...... 3. B. arceutina
26. Width of apothecia seldom exceeding 0.3 mm. Spores 34. Thallus partIy dissolving into finely granular gonio-
(average of 10) up to 1.7 pm wide. Ocular chamber cysts. Proper exciple partly of cells with ?I ellipsoid lu-
lacking. Southeastern .....................3 3. Bn. ctystallifera mina. Florida ............................................ 28. Bn. aenea
26. Width of apothecia usually exceeding 0.5 mm. Spores
(average of 10) at least 2.0 p wide. Ocular chamber 35. Thallus entirely consisting of rather large granules (av-
narrow and high. Great Lakes and northwards .............. erage of 20 granules exceeding 70 pm) ..................... 36
............................................................. 2 0 . B. msella 35. Thallus either not at all granular, or partly or entirely
dissolving into finely granular goniocysts (average of
27. Spores (average of 10 or more) 1 .O-1.4 pn wide ...... 28 20 granules less than 60 p) ...................................... 37
27. Spores (average of I0 or more) 1.6-2.8 pm wide ...... 30
36. Apothecia in section f yellowish to pale orange
28. Apothecia with pale yelIow or pale grey disc. Margin throughout, K+ intensifying but never K+ purplish.
paler than disc, persistent. ............. 3 2 . Bn. chloroticuIa Upper cortex with crystals ...................... 21. B, ribella
28. Apothecia with 4 pink disc. Margin concolorous with 36. Apothecia in section with at least upper part of proper
disc, finally excluded. Southeastern ........................... 29 exciple and part of the hymenium +
red-brown, K+
purplish. Cortex without crystals ........... 5 . B. biatorina
29. Thallus partly dissolving into finely granular gonio-
cysts (granules c. 20 pm or less) ............. 39. Bn. sp. #I 37. Outermost 4-8 cell layers of proper exciple with enlarged
29. Thallus of small, deeply incised squarnules, the lobes cell lumina that are distinct h m the inner,narrower ones,
of which are 20-30 p n wide; not dissolving into gonio- or cell lumina gradually but distinctly enlarging towards
cysts .............................................. 37. Bn. squamellosa the rim (if in doubt due to obscuring pigments, use a di-
luted hypochlorite solution to bleach them) ..................38
30. Hyrnenium (average of 5 measurements in one apoth- 37. Outermost 1-2 cell layers of proper exciple with en-
ecium) 75-88 pn high ......................... 23. B. salmonea larged cell lumina that are distinct from the inner, nar-
30. Hymenium (average of 5 measurements in one apoth- rower ones, or entirely without enlarged cell lumina
ecium) 42-73 pm high ................................................ 3 l .. ............................
along rim ............................... 3 9
O w m But 1 2 i . IYY(7
53-57(-66) pm long (S= 8, N= 4, n= 10), (2.1-)2.5-2.6- more or less fusiform spores, the length of which rarely
2.6(-3.3) pm wide (S= 0.2, N= 4, n= 10), (1 1 .l-)17.1- exceeds 10 times the width. B. combinans, although
20.5-22.1(-26.5) times as long as wide (S= 3.7, N= 4, n= very similar to the other species in pigmentation, is easi-
101, with (3-14.7-6.4-7.5(-9) septa (S= 1.7, N= 4, n= 10). ly separated by the short (up to c. 20 pm long), bacilli-
Pycnidia f immersed in thallus, uppermost part of form, usually Ccelled spores and the lack of Bacidia
wall dark brown (even when apothecia are pale), 75- 125 Green in the apothecia.
pm diam. Conidia filiform, curved, non-septate, 15-24
X c. 0.5 pm. Collections examined. U . S. A. Florida. Collier Co.,
Chemistry: Trace amounts of atranorin or no lichen 1993, Ekmltn L1025 (LD). Dade Co., 1993, Ekman
substances at all. ' L1 186, 1 187, 1 188 (LD), 1982, Tucker 25229 (herb.
Pigments: Dark parts of proper exciple and lower Tucker). Sarasota Co., 1981, Thor 21 89 (S).
part of hypothecium K-, N-. Upper part of hymenium
and f orange or brown parts of proper exciple K+ in-
tensifying, N+ intensifying. Greenish parts of proper 3. Bacidia arceutina (Ach.) Arnold
exciple K-, N+ purple. Crystals in proper exciple and
epithecium partly soluble in K but insoluble in N. - Verh. zool.-bot. Ges. Wien 19: 624 (1869). - Lecidea
Schweinitzii Red in proper exciple, hypothecium and luteolu var. arceutina Ach., Method. Lich. 61 (1803). -
pycnidial wall. Rubella Orange in proper exciple, hy- Lecidea arceutina (Ach.) Gray, Nat. Arrang. Brit. Plants
pothecium and hyrnenium. Bacidia Green sometimes in 1: 472 (1821). - Type: Sweden (H-ACH 347E, lecto-
small amounts in proper exciple. type selected here).
Ecology. At West Lake, Dade Co., B. aggregahxla grew Illustrations. Figs 5A, 18 (map), 40C.
abundantly together with B. helerochroa and B. nautabi-
Iis on Conocarpus emc&s, Lugunculari~rucernosa, and Characterization. At least young apothecia f yellowish,
Rhizophora maagle in a mangrove swamp. In Cork- without orange or red hues. Proper exciple distinctly
screw Swamp, Collier Co., B. aggregatula occurred brown-yellow to yellow-brown in the rim, otherwise
sparingly on Acer rubmm in a swamp dominated by old pale. Hypothecium f yellowish. Upper part of hy-
Taxodium a'istickum. At two additional localities, where menium brown-yellow to yellow-brown. Spores acicu-
I did not observe B. aggregatuda myself, it was found lar, 35-71 X 1.2-2.8 pm, with 3-15 septa.
on an unidentified tree in a "swamp forest" and on culti-
vated hardwoods in a botanical garden. Thdlus indeterminate or determinate, thin, continuous,
smooth or wrinkled, rareIy warted or almost granular, f
Distribution. Known from only four localities in Col- cracked or areolate, rarely discontinuous and of discrete
lier, Dade, and Sarasota Counties in southern Florida. - or contiguous, thin areoles; usually paIe grey but some-
Previously known only from the Brazilian type collec- times chalky white, pale yellow-grey or pale green-grey.
tion. Prothallus sometimes present between discrete areoles,
endosubstratal, thin, Iight grey.
Discussion. Bacidia aggregatula is obviously closely re- Apothecia (0.3-)0.4-0.5-0.8(-1.1) mm diam. (S= 0.1,
lated to B. schweinitzii and the neotropical species B. N= 16, n= 5), f plane when young, later becoming con-
horirrta (Nyl.) Zahlbr, and B. combinans (Nyl.) Zahlbr. vex, epruinose. Disc pale brown, yellow-brown, grey-
All four species have a darkIy pigrnented proper exciple brown, or brown to brown-black, rarely pale beige, red-
dominated by Schweinitzii Red, and Bacidia Green is of- brown, or purplish brown to purplish black, at least
ten or always present in the hyrnenium or proper exciple when young with a yellowish hue. Margin in upper part
of all species except B. combinans. B. schweinitzii is sep- concolorous with disc or slightly darker, in lower part
arated from B. aggregutul~by the thicker paraphyses, by usually paler than disc and upper part of margin, pale
the absence of minute crystals in the epithecium and beige to brown, distinct, raised above disc in young
proper exciple, and by the pigmentation: B. schweinitzii apothecia, soon level with the disc, and finally excluded
is usually dominated by either Schweinitzii Red in the in old and convex apothecia.
proper exciple and Bacidia Green in the upper part of the Proper exciple laterally (29-)50(-72) pm wide (S= 12,
hymeniurn, or by Rubella Orange in both the proper exci- N= 16, n= l), without crystals or rarely with diffuse
ple and the upper part of the hymenium. In B. aggregaht- clusters of minute crystals (less than 1 pm in diam.).
In, on the other hand; neither RubelIa Orange nor Bacidia Rim pigmented in a distinct zone, brown-yellow to dark
Green are ever dominant pigments. B. horista differs yellow-brown to almost pure brown in upper part,
from B. aggregutulu in lacking crystals in the proper ex- downwards usually gradually changing to pale yellow or
ciple and apothecium, and in having shorter and wider, colourless, along the edge with a single row of enlarged
cells with lumina that are up to 9 X 8 pm. Lateral interi- barked branches of trees and shrubs. It favours ace^
or paler than rim, colourless, except upper part close to macrophyllum, Alnus mbm, and Populus trichocarpa,
hymenium, which is often pale yellow to brown-yellow. but has occasionally been found on Amelanchier alnifo-
Medullary part usually paler than hypothecium and con- Iia, ArbuCIkS inevtziesii, Cogvlus cordata, Fraxinus latifb-
colorous with lateral interior, colourless to pale yellow. lia, Ilex opacu, Juniperus virginiana, Lonicera sp.,
Hypothecium usually f yellowish, only rarely entirely Oemleria cerasifornais, Prunus ilicgolia, Quercus veiu-
colourless. Hyinenium (56-)63-78-93(-99) pm thick (S= tinu, Ribes bructeosurn, Salix sp., Thuja occidentalis,
8, N= 16, n= 5), lower part colourless, uppermost part and Unabeflu/aria caIfornica.
brown-yellow to dark yellow-brown to almost pure
brown in a thin distinct layer. Paraphyses 1.0-1.6 p n Dirrtribuaion. Coastal Maine, Massachusetts, and New
wide in mid-hyrnenium; apices f clavate or not at all York, and along the West Coast from central California
l .8-2.2-2.7(-3.2) pm wide (S= 0.5, N=
swollen, ( l .Q-) to southern British Columbia. - Europe.
16, n= 5), without internal pigment. Spores acicular,
usually f sigmoid, but sometimes straight or curved, Discussion. Bacidia arceuaina is a characteristic species
sometimes coiled in ascus, (35-)47-54-64(-71) pm long that is only likely to be confused with B. salmonea (see
(S= 7, N= 16, n= 10), (1.2-)l .5-1.8-2.2(-2.8) pm wide that species) and B. Iraurocerasi. B. laurocerusi differs
(S= 0.3, N= 16, n= 10), (18.3-)25.3-30.2-36.4(-46.0) from B. arceutina in having a brown, K+ purplish, N+
times as long as wide (S= 5.5, N= 16, n= 10), with (3-) orange pigment in the proper exciple and the upper part
5.0-7.0-10.7(-15) septa (S= 2.1, N= 16, n= 10). of the hymenium, and in having shorter and narrower
Pycnidia immersed in thallus (only ostiole visible), spores with fewer septa.
colourless, 75-100 p diam. Conidia filifom, curved,
non-septate, 9-1 7 X 0.6-0.8 pm. Collectio~ls examined. CANADA. Brilish Columbia.
Chemistry: No lichen substances detected. 1912, Macoun (CANL). Hope, 1969, Brodo 15641,
Pigments: Apothecia K-, N-. - Arceutina Yellow in Krog & James (CANL). Vancouver Island, Comox,
proper excipIe, hypothecium, and hymenium. 1893, Macoun (CANL, NY, US, WIS). Vancouver Is-
land, Sidney, 1912, 1913, Macoun (FH), 1916, Macoun
Ecology. In hardwood or mixed forests or forest fringes, 308, 326 (UBC). Vancouver Island, Qualicum Beach,
where it grows directly on the bark of hardwoods and 1989, Ekman L756, L757, L758, L759 (LD). Vancouver
shrubs, rarely over bryophytes. It can be found on Island, Victoria, 1893, Macoun (NY), 1989, Ekman
trunks, but more often it is found on thin, srnooth- L77 1, L774 (LD). Vancouver Island, Jordan River,
Disczcssion. B, cumpalea is closely related to B. sufisa, IIIustrations. Figs 6C-D, 8E-F, 2 1 (map), 400.
B, fusconigrescens (Nyl.) Zahlbr., and B. millegrana
(TayIor) Zahlbr. B. s u m a can be separated from B. Chamcterization. Apothecia usually black, flat. Proper
caapulea by the lack of a bluish tinge of the thallus, the exciple with green and brown (sometimes almost black)
lack of minute crystals in the proper exciple (although pigmentation, mainly consisting of distinctly radiating
clusters of large crystals may be present), the narrower hyphae. Hypothecium colourless to brown-orange. Up-
zone of enlarged cells along the edge of the proper exci- per part of hymenium greenish, sometimes also with
ple, and by the thicker paraphysis apices. B. jiisconi- some purplish brown pigmentation. Spores bacilliform
grescens, which is presently known only from tropical to clavate, 11-45 X 1.6-3.7 pm, with 3-7 septa.
America and Asia, differs in having a proper exciple
with abundant minute crystals but no clusters of large Thallus indeterminate, thin to rather thick, either dis-
crystals, in having a thin but distinct, pigmented zone continuous, of discrete or partly contiguous, convex,
along the edge of the proper exciple, in having a K+ sometimes subsquamulose areoles, or continuous, wart-
ed or sometimes smooth, f rimose or without cracks; terior usually paler than rim, or sometimes concolor-
light grey to grey to yellow-grey to green-grey to grey- ous with or (predominantly in lower part of the exci-
green to green-brown to grey-brown. ProthaIlus lacking ple) darker than rim. Medullary part usually paler than
or present between discrete areoles, white, thin, endo- hypothecium and lateraI interior, colourless or rarely
substratai. pale brown or brown-orange. Hypothecium colouriess
Apothecia (0.2-)0.3-0.4-0.7(-0.8) mm diam. (S= or sometimes pale yellow to brown-orange, rareIy pale
0.1, N= 16, n= 5), f plane, remaining so or becoming to medium brown. Hymenium (43-)47-61-73(-78) pm
moderately convex or rarely markedly convex, epru- thick (S= 9, N= 16, n= 5), lower part colourless except
inose, low and flat, 2.1-4.0 times as wide as high. for occasional vertical streaks with green to blue-green
Disc usually pure black, seldom partially or entirely pigment, upper part dirty green to blue-green or partly
blue-black, blue-grey or dirty yellow. Margin con- purplish brown, indistinctly or distinctly delimited.
coIorous with, or rarely slightly paler or darker than Paraphyses 0.8- l. .G ym wide in mid-hymenium; apices
disc, often glossy, distinct, raised above disc in young f clavate or a few only slightly or not at all swollen,
apothecia, later level with the disc, persistent or only (1.2-) 1.7-3.1.-4.1(-6.5) pm wide (S= 1.0, N= 16, n= 51,
rarely excluded. without internal pigment but with an externaI cap of
Proper exciple laterally (19-)46(-69) p wide (S= green or brown pigment. Spores bacilliform to clavate,
13, N= 16, n= l), without crystals, almost entirely straight or slightly curved, not coiled in ascus, (1 1-)12-
formed by regularly radiating hyphae with 0.8-3.5 ,urn 24-37(-45) pm long (S= 7, N= 16, n= 101, (1.6-jl.7-
wide lumina (except for the innermost part of the ex- 2.4-3.2(-3.7) pm wide (S= 0.4, N= 16, n= 10), (5.0-)
ciple, flanking the hypothecium, which consists of a 6.0-10.2-15.8(-20.0) times as long as wide (S= 3.4, N=
few, 0.8-3.0 pm wide, sparsely branched hyphae that 16, n= 101, with (3-)3.0-3.9-6.8(-7)septa (S= 1.4, N=
are not clearly directed towards the rim). Rim grey- 16, n= 10).
+
green to blue-green to brown to black, rarely col- Pycnidia common but not present in all specimens,
ourless or pale green, in upper part, downwards usu- sessile or half-immersed in thallus, uppermost part of
ally gradually changing to colourless or f brown, wall concoIorous with apothecia, 75-300 ym diam.
along the edge with a single cell layer of enlarged Conidia of three types (never more than one type in a
cells with lumina that are up to 8 X 7 p.Lateral in- thallus): ( l ) oblong to fusiform to bacilliform, 0-1-
Discussion. Bucidia circurnspec6a is similar to B. igni- Collections examined. CANADA. British Columbia.
arii, B. reagens, B. schweinitzii, and B. subincornpta. Gabriola Island, 1974, Noble 3008 (herb. Noble). Van-
The distinction of B. igniarii from B. circumpecta is couver Island, Williarn Head, 1975, Noble 4594B (herb.
discussed under the former species. B. subincompta catl Noble), 1989, Ekrnan L766 (LD).Vancouver Island,
be separated by the longer spores, the K+ purplish hy- Mount Douglas, 1991, M. Ryan (herb. M. Ryan). Wells
pothecium, the margin that becomes excluded in old Gray Prov. Park, 1994, McCune 21862 OSC). New
apothecia, and (often) on the distinctly granular thallus. B m s w i c k . Kent Co., 1978, Egger 683b (CANL). Nova
In B, schweinitzii, the hypothecium is brown, and the Scotia. Victoria C o . , 1989, Ekrnan L1069 (LD). Hank
spores acicular. B, mugens has a K+ purple to violet and Co., 1989, Ekrnan L1037 (LD). Onaario. Renfrew Co.,
C+ violet upper hymeniurn. 1985, Wong 3809 (CANL). Quebec. Gatineau Co., 1989,
The main part of North American B. circurnspecta Ekrnan L 1025 (LD) . Saskatchewan. Prince Albert Nat .
agrees completely with European representatives. The Park, 1969, Jesberger 1150 WIS). - U. S. A. Arizona.
range of variation in hymenium height, spore size, spore Coconino Co., 1974, Brodo 20306a (CANL). Arkan-
shape, apothecium pigmentation, and conidial appearance, sas. Izard Co., 1988, Harris 21649a, 21653a (NY). Cal-
however, is much larger in North America than in Europe. ifornia. Santa Monica Range, 1894, 1903, Hasse (US).
In European material, I have not observed forms with a Mendocino Co., 1989, Ekman L790 (LD).Monterey
brown or brown-orange hypothecium, clavate spores, or Co., 1989, Ekman L1219 (LD). San Luis Obispo Co.,
conidial types other than the oblong to f u s i f m to bacilli- 1900, Barker (NY). Santa Barbara Co., 1989, Ekrnan
form. Since the different conidial types never seem to W- L633, L634 (LD). Massachusetts. New Bedford ( F H ) .
cur on the same thallus, and not even at the same locality, I Michigan. Cheboygan Co., 1974, Harris (NY).Delta
have considered the possibility that North Amaican B. cir- Co., 1976, Harris 1 1946a (CANL, MICH), 1969, Harris
cum-pecta in the present sense shouId be split into two or 4034 (Mm, MSC). Menominee Co., 1965, Barris 752b
more species. The variation in conidial type, however, is (MSC). Roscommon Co., 1974, Wang 635 (MSC). Min-
apparently not correlated with any other character, and thus nesota. Anoka Co., 1980, Trana 10096 (MIN). Itasca
I interpret B. circumpecta as one single, variable species. Co., 1977, Trana 2973 (MIN). Lake Co., 1989, Ekrnan
Part of the North American variation within B, cir- L944 (LD). Otter TaiI Co., 1900, Fink 3 (MM). New
York. Brom Co., 1988, Buck (NY). North Dakota. cum ea apothecii minori), apicibus paraphysom an-
Billings Co., 1982, Wetmore 45454a (Mm). McKenzie gustioribus.
Co., 1982, Wetmore 44658 ( M W . Oklahoma. Chero-
kee Co., 1988, Hamis 21362 (NY). Oregon. Linn Co., Eiymology. DlfSractus, broken in pieces.
1970, Pike 1368 (OSC). South Carolina. Chester, Green
(NY).South Dakota. + Lawrence Co., 1960, Anderson fllzcstrations. Figs 22 (map), 40H,43F.
(filed with B. verrnifera, COLO). Washington. Repub-
lic, 1913, Foster (FH). Wisconsin. Dane Co., 1976, Characterization. ThalIus granular; granules 36-97 p
Thomson 18823 (CANL, MIN). diam. Apothecia brown-orange to red-brown to dark
purplish brown, at least some pruinose. Proper exciple
f brown-orange to orange-brown. Hypothecium darker
8. Bacidia diffracts S. Ekman, sp. nov. than proper exciple. Upper part of hymenium colour-
less to orange-brown. Most pigmented parts K+ pur-
Type: Canada, Nova Scotia, Queens Co., Kejimkujik ple-red. Spores acicular, 32-69 X 1.9-4.1 pm, with 3-
Nat. Park, E side of Kejimkujik Lake, c. 700 m SE of l l septa.
the N end of Peter Point, on Fagus grrandiJoolia, alt c.
100 m, 1.989, Ekman L1050 (LD, holotype; CANL, iso- Thallus indeterminate, thin to rather thick, continuous
type). or + discontinuous, finely granuIar, pale grey, pale
greenish grey, yellowish grey, greenish grey, greyish
A simili Bacidia polychroa differt thallo granuloso, green, or grey. Granules (36-)56-67-82(-97) pm d i m .
apotheciis altioribus broportione altitudinis hymenii (S= 16, N= 13, n=5), f globose or irregular in shape.
wide (S= 2.8, N= 23, s 10), with (3-)4.5-9.6-13.6(- been found in planted tree-stands. Known phorophytes
15) septa (S= 3.1, N= 23, n= 10). include (in the Southeast:) Carya illinoensis, Salix spp.,
Pycnidia partly or entirely immersed in thallus, up- Avicennia germinans, Cephalanthus occidentalis, Cor-
permost part of wall concolorous with apothecia, 40- nus foemina, Datum sp., Cornus foemina, Diospyros
100 pm diam. Conidia filiform, curved, non-septate, 5- virgi~ianu,Liquidam bar s pruclflua, Mytsine floridam,
1s X 0.8-1.0 p. Morus rubra, Lysiionaa la~isiliquum, Sambucus ca-
Chemistry: Western specimens contain no lichen sub- nadensis, Piscidiu piscipuiu, Pluneru aquatics, @er-
stances, whereas eastern ones contain atranorin (a1- c m spp., Rhizophora mangle, Allhaeu sp., Celtis laevi-
though sometimes only in trace amounts). gata, Sapiurn sebijerum, Azaleu sp., Forestiera acu-
Pigments: Upper part ofhymenium and rim of proper mina fa, Lage~qstroemiaindica, Nyrrsa sp., PopuIecs del-
exciple K+ purplish, NI- orange-red to red. Hy- toides, Wisteria sinensis, Yitis rotundifolia, Junipems
pothecium and inner parts of proper exciple K+ slightly ashei, J: rrilicicola, (in the West:) Acer macrophyllum,
intensifying, N-. - Laurocerasi Brown in hymenium (as Ceanothus divaricatus, Cupressus macrocarpa, Anus
caps on paraphysis apices), rim of proper exciple, and ponderosa, Pmnus ilicgolia, Quercus agr folia, Q. 10-
pycnidial wall. Rubella Orange in hypothecium and in- batu, Rkus diversiloba, Salix spp., Sambucus coerulea,
ner part of proper exciple. and UmBeflularia culdfornica.
Ecology. This species has a very wide ecological ampli- Distribution. In the west known from California and Or-
tude. It has been found on trunks and branches of a egon, and in the southeast from Texas, Louisiana, Ala-
wide variety of trees and shrubs in forests ranging from bama, Florida, Georgia, South Carolina, and North
shady and humid to sun-lit and dry. Occasionally, it has Carolina. - Probably one of the most widespread spe-
Discrdssion. Judging from neotropical material, Bacidia Ilhstraaions. Figs 19 (map). 41D.
hostheleoides is a very variable species in tha1lus forma-
tion and apothecium pigmentation. The thallus ranges Clauructerizalion. Apothecia black, high and barrel-
from almost smooth to granular, and the apothecia vary shaped to inversely cone-shaped. Proper exciple with
from pink to dark purple-brown. In these respects, the green and brown pigmentation, strongly developed be-
variation much paralIels that of B. mediulis, which is low hypothecium, to the greater part consisting of intri-
obviously a very close relative. There is, however, a dif- cately intertwined hyphae. Hypothecium pale. Upper
ference in that apothecia of B, mediulds are usuaIly f part of hymenium blue-green. Spores bacillifom to cla-
pinkish, whereas they are usually brown-orange to f vate, 12-19 X 2.5-2.9 p~,
with 3 septa.
purple-brown in B, hostheleoides. There is also a differ-
ence in tl~aalluscolour: B. medialis is commonly f deep Thallus indeterminate, rather thin, discontinuous, of dis-
grey-green, whereas B. hos~heleoidesusually is pale crete or contiguous, convex, sometimes subsquamulose
grey to pale green-grey. The entire variation from pink areoles, light brown-grey. Hypothallus not visible.
to purple-brown apothecia and pale grey to dark grey- Apothecia (0.2-)0.3(-0.4) mm diam. (S= 0.05, N= 1 ,
green thallus, however, occurs in both species. n e only n= 1O), f plane, remaining so or becoming slightly con-
constant difference between the two species appears to vex, high and barrel-shaped when young, Iater becom-
be the anatomy of the proper exciple. In B. hosthele- ing inversely cone-shaped, 1.7 times a wide a high. Disc
oides, the lumina in the lower part of the proper exciple pure black. Margin black in upper part, downwards
are as narrow or usually narrower than the ones in the gradually changing to light yeliow-brown, distinct,
upper part of the proper exciple. In B. medialis, on the raised above disc in young apothecia, Iater level with
other hand, the lower part of the proper exciple to a the disc, persistent.
large extent consists of hyphae with lumina that are f Proper exciple Iaterally 35 pm wide (N= 1, n= l),
without crystals, strongly developed below hy- number of spore septa, the only diagnostic character
pothecium; outermost part formed by radiating hyphae positively separating the two species is the anatomy of
(the lumina of which are f cylindrical, 0.8-2.5 pm the proper exciple. In B, circumspectu, the apothecia are
wide), inner (major) part of narrow, intricately inter- low and flat. The main part of the proper exciple is
twined and richly branched hyphae (lumina 0.5-1 pm formed by regularly radiating hyphae. Only a small por-
wide). Rim dark brown-green in uppermost part, down- tion of the innermost part of the excipIe, flanking the
wards gradually changing to dirty brown, pale brown hypothecium, consists of hyphae that are not clearly di-
and then colourless, along the edge without or with a rected towards the rim. In B. igniarii on the other hand,
single cell layer thick zone of cells with enlarged lumi- the young apothecia are high and barrel-shaped with al-
na. Lateral interior usually paler than rim, palebrown in most vertical sides. As they age, the apothecia remain
upper part, downwards gradually changing to colour- high, but become more or less inversely cone-shaped.
less. Medullary part colourIess. Hypothecium very pale This appearance is caused by a strongly developed exci-
brown. Hymenium (42-)M(-47) pm thick (S= 2, N= I , ple below the hypothecium. The outermost part of the
n= 5 ) , lower part coIourless, upper part f blue-green. proper exciple is formed by regularly radiating hyphae,
Paraphyses 1.2-2.0 pm wide in mid-hymenium; apices f but the inner dominating part consists of intricately in-
clavate, (2.0-)2.8(-4.0) pm wide (S= 0.5, N= l , n= 20), tertwined and richly branched hyphae.
many with an external hood of green pigment. Spores Pycnidia have not been observed in the North Ameri-
bacilliform to clavate, slightly curved (banana-shaped) can specimen. In Swedish specimens, l~owever,they are
or (a few) straight, (12-)15(-19) pn Iong (S= 2, N= 1, often present. They are black and sessile, 75-100 pm in
s 20), (2.5-)2.6(-2.9)pm wide (S= 0.2, N= 1, n= 20), diameter. The conidia are bacillifom, straight, 0-1-sep-
(4.3-)5.9(-7.0) times as Iong as wide (S= 0.7, N= 1, n= tate, and 5-12 X 1.2- 1.5 pm, i.e., similar to type I of B.
20), with 3 septa. circ-cupnspecfu(see that species).
Pycnidia not seen in North American specimens. Like in the North American specimen of B. igniarii,
Chemistry: Not examined due to sparsity of material. 3-septate spores are dominating also in North European
Pigments: Upper part of hymenium and uppermost specimens. In some Swedish specimens, however, I
part of proper exciple K- or partly K+ purplish, and N+ have observed 5-septate spores mixed with the 3-septate
purple with a precipitate of blue crystals. Proper exciple ones.
and hypothecium K+ purplish, N+ orange-red. - Bacid-
ia Green in hymenium and uppermost part of exciple. Collections examined. CANADA. Saskutchewan. C y -
Laurocerasi Brown in proper exciple, hymenium (small press Hills, 1964, Jonescu (WIS).
amounts), and hypothecium (small amounts).
Ecology. Bacidia igniarii was collected on a trunk of 14. Bacidia insularis Zahlbr.
Populus ~emuloides1.3 m above the ground.
Annls mycol. 29: 82 (1931). - Type: Ecuador, "Insulae
Distribution. Known from only one single North Ameri- Galapagos: Charles Island, Postoffice Bay, ad cortices
can locality in southwestemmost Saskatchewan. - Eu- laeves", Herre, ZahIbruckner in Kneiff & Hartmann:
rope. Krypt. exs. Vindobon. 3 155 (W, lectotype selected here;
LE), isotype)
Discussion. All previous reports of B. igniarii from
North America have turned out to be erroneous, be- Iliustrutions. Figs 20 (map), 4 1E.
ing based on short-spored forms of B. circumspect^.
B. ignicarii is similar to B. circumspecta, B. schwein- Characte~ization. Apothecia black, often pruinose.
itzii, B. reagens, and B. subincompta. The three last Proper exciple laterally blue-green with only some
species all have longer spores than B. igniarii. Fur- brown pigmentation, otherwise colourless to blue-green.
thermore, B. schweinitzli has an orange-brown to red- Hypothecium f brownish and sometimes also partly
black proper exciple that merges into the hy- blue-green. Upper part of hymenium blue-green. Para-
pothecium, B. reagens a C+ violet upper hymenium, physes 0.8-1.2 pm wide in mid-hymenium. Spores fusi-
and B. subincompta a brown K+ purplish hypo- form to acicular, 26-77 X 3.1-5.3 pm, with 3-15 septa.
thecium. B. ci~.cumspectais the species that is the
most similar to B. igniarii. Although B. igniarii in Thallus indeterminate or determinate, thin to rather
most floras is keyed out as having shorter spores than thick, usually continuous, warted to subsquamulose or
.B. circumspecta, this is not the critical character for sometimes almost granular, f cracked, rarely discontin-
separating these species. Although there are probably uous, of f discrete, thin to rather thick, convex and
differences in average spore length and average warted areoles, light grey, green-grey, brown-grey, or
vertens have nothing to do with B. luurocerasi subsp. Thallus usually thin, almost white to grey, or yellowish
ida hoensis. grey, very rarely grey-green, without cracks or f
Some authors distinguish between Bacidia lau- cracked, rarely areolate. Apothecial margin in upper
~.ocerasiand B. subacerina Vain. They are then mainly part f concolorous with disc, only very rarely with a
separated by apothecium shape. The former is stated to reddish hue. Hymenium (7 1-181-9 1- 106(-118) pm thick
have broadly adnate apothecia that fairly soon become ( S = 9, N= 2 1, n= S ) . Spores (50-157-70-87(-99) p long
convex, and the latter cup-shaped apothecia that re- ( S = 10, N= 21, n= 10), (1.9-)2.5-2.7-3.5(-3.7) p wide
main plane for a longer time. Additional characters ( S = 0.4, N= 21, n= 101, with (7-)I 1.9-15.9-22.2(-28)
have also been used, usually very vague and widely septa (S= 3.6, N= 21, n= 10).
overIapping. In my opinion, the variation observed in
North American as well as European material does not Ecology. Known phorophytes include Abies balsamea,
justify a taxonomic recognition of B. subacerina at the Acer mbmm, Acer sacchamm, Fraxinu~spp.,,Junipe-
species level. The variation is continuous, and both the rus virginiuna, Picea spp., Populus spp., Quercus ru-
supposedly discriminating character states can some- Bra, and Thuja occidentalis.
times be found in a single specimen. Should future in-
vestigations of European B. laurocernsi show that fur- Distribution. The Great Lakes and northern Appalachi-
ther subspecies ought to be recognized (see discussior an regions. - East Asia (?),Europe.
in the chapter Numerical and statistical treatment),
:hen the epithet subacerina may reappear at this taxo- ~ollectionsexamined. CANADA. New ~rulaswick.Al-
iornic level. bert Co., 1980, Gowan 2049, 3188, 4009 (CANL).
Charlotte Co., 1989, Ekman L1085, L1086, L1088,
L1092 (LD). Rent Co., 1978, Egger 147a (CANL).
15a. sut,,. .,,rocerasi King's Co., 1970, Ireland 13476 (CANL). Nova Scotia.
Victoria Co., 1989, E h L1070, L1071 (LD). Hants
Co., 1989, Ekman L1039 (LD). Kings Co., 1989, Ek-
Illustrations. Figs 3B, 6A, 26 (map), 41F. man L1041 (LD). Lunenburg Co., 1972, Brodo 18954
(pigmentation distinctly or indistinctly delimited). Hy- pothecium, proper exciple, and pycnidial wall.
menium (56-)63-77-94(-102) pn thick (F 10, N= 22, n=
5), constituting (19-)29(-39) % of the entire height of the Ecology. Mainly on the coarse bark of old trees in shady
apothecium (S=.6, N= 22, n= l), in lower part colourless or or semi-open, usually humid habitats. It prefers Fruxi-
very pale brown-orange to pale brown, upper part con- nus spp. and Thuja occidentulis, but has also been
colorous or (rarely) with fairly distinct brown-orange pig- found on Ace?.saccharurn, Alnus sp., Ccarya spp., Celtis
mentation. Paraphyses 1.O-1.6 pm wide in rnid-hymenium; occidentalis, Fagus grand$olia, Juglans cinerea, Liri-
apices f clavate or not at all swollen, (1-2-11-7-2.3-3.0(- odendron tulipifera, Ostiya virginiana, Platanus ~ c c i -
3.7) p n wide (S= 0.5, N= 22, n= 5), without internal pig- dentalis, Populus spp., Quercus sp., Salix sp, Taxodium
ment. Spores acicular, straight, curved, or sigmoid, some- distichum, Tilia americana, and Ulmus americana.
times coiled in young asci, (3 1-)38-48-57(-74) pm long (S=
7.4, N= 22, s lO), (1.9-12.3-2.74.0(-5.0) pm wide (S= Distribution. Eastern temperate region with a preference
0.5, N= 22, n= 10), (7.3-11 1-4-18.2-21.6(-30.0) times as for the Great Lakes, Appalachians, and the Ozark Pla-
long as wide (S= 3.8, N= 22, n= lO), with (2-)3.1-6.3- teau. - Europe.
11.9(-15) septa (S= 2.4, N= 22, n= 10).
Pycnidia f immersed in thallus, pale brown or pale Discetirsion. Bacidia polychroa is similar to B. d@racta
orange-brown above, 100-170 pm d i m . Conidia fili- and B. rubella. B. rubella can be separated by the granu-
form, curved, non-septate, 10-17 X 0.6-0.8 pm. lar thallus and the lack of a K+ purple-red pigment (Ba-
Chemistry: Atranorin (usually only trace amounts) or cidia Brown) in the apothecia. For detaiIs on the separa-
no Iichen substances at all. tion of B. polychroa and B. dzflacta, see the latter
Pigments: Most parts of apothecia K+ purple-red, N-. species and the chapter Numerical and statistical ckap-
- Bacidia Brown and Rubella Orange in hymenium, hy- ter.
Ecology. On the bark of hardwoods in shady and humid Discussion. Bacidia reagens is superficially similar to
localities. Known phorophytes include Acer macrophyl- Bucidia absistens, B. subincomptu, B. circu~specta,
lum. Nvssa svlvatica. and Ouercus ugrfoliu. and B. igniarii. Unlike these species, B. mugens con-
tains Thalloidima Green in the apothecia, which reacts
Distribution. Known from a few localities in coastal C+ violet.
southern California and from one locality each in south- I know of only one additional species of Bacidia S.
ern Louisiana and southernmost Georgia. - Bcacidia rea- sir. that contains Thalloidima Green in the apothecia,
gem has previously been reported only from the type col- namely B. violascem K. Ralb & VCzda, described from
lection and one additional collection, both from Rio Hawaii (Kalb & Vgzda 1980). It is very similar to B. re-
Grande do Sul in southern Brazil ( M h e 1935). It is an agens in pigmentation but differs in the shorter and
inconspicuous species likely to escape attention. Thus, it wider, fusiform spores that are usually only 3-4 times as
is uncertain whether it is widely dishbuted in the neo- long as wide.
tropics with some outliers in the subtropical areas of the
U. S, and southern South America, or restricted to sub- Collections examined. U. S . A. Caiijorniu. Monterey
tropical areas north and south of the neokopics. Co., 1989, Ekman L627, L628, L801 (LD). Santa Bar-
bara Co., 1989, Ekman L619, L620 (LD). Georgia.
cidia than to North American B. rosella. A case could t o m verraaIis will take precedence over Bacidia ru-
be made for recognizing North American B. rosellu bella.
taxonomically. Given the overall anatomical similari- The types of Lichen lutereus and Lichen corlicalis
ties and the very small number of known North have not been studied. Lichen lutemus is. next to Lichen
American specimens, I refrain from doing so at the vernalis L., the oldest name for Bacidia rubella, provid-
moment. ed that the type of Lichen lulereus I. F . Gmel. turns out
to fall within-the present concept of the species. In the
Collections examined. CANADA. Ontario. Thunder protologue of Lichen Ititereus, Gmelin (1792) referred
Bay District, 1977, Wetmore 29024 (MIN). + Cochrane to Schrader. It is unlikely that Gmelin ever saw any ma-
District, 1969, Brodo, 1 4 7 1 7 ~(fiIed with B. subin- terial of this species; he probably relied on information
compta, CANL) - U. S. A. Michigan. Keweenaw Co., provided by Schrader. Later, Schrader (1794) made Li-
1980, Wetrnore 413 19B (MIN), 1983, Wetmore 49341 chen luteolus, which is superfluous since he included
(MW- (by direct reference) Lichen lutereus J. F . Gmel. Unfor-
tunately, Schrader erroneously cited Lichen htereus as
"Lichen, luteolus", but from the phrase description ("ci-
21. Bacidia rubella (Hoffm.) A. Massal., nom. cons. nereus, tuberculis, sparsis, lutescentibus") and the refer-
Prop- ence to a page and the number of the species ("Gmel.
Syst. Nat. I1 p. 1359 n. 50'7, it is clear that he referred
Ricerch. Auton. Lich. 1 18 ( 1 852). - Lichen rubellus to Lichen lutereus. This has caused some confusion,
Ehrh., Plant. Crypt. Linn. 196 (1791), nom. nudum. - since Gmelin's treatment included also a "genuine" Li-
Vermcaria rubellu Hoffm., Deutschl. Flora 2: 174 chen luteolus J . F . Ginel. (1792: 1368, no. 161). Thus,
(1796). - Bacidia rubella (Hoffm.)A. Massal., Ricerch. Lichen Iuteodus Schmd. is not only a superfluous syno-
Auton. Lich. 118 (1852). - Type: Germany, Hannover, nym of Lichen lutereus J. F . Gmel., but a younger hom-
Ehrhart: Plant. Crypt. Linn. 196 (GOET, lectotype se- onym of Lichen luteolus J . F . Gmel. All subsequent
lected here). - Proposed conserved type: Sweden, combinations based on Lichen hteolus Schrad. (includ-
"Gotland Prov., Arlingbo par., Lilla Atlings forest ing the much used Bacidia Iuteola), by direct or indirect
meadow, c. 1.6 km ESE Atlingbo church. 57'28'N reference, includes the type of Lichen lutereus and are
18O22'E. On Uimus m i m r " , 1990, Nordin, Sundin & thus nomenclaturally superfluous. If any original mate-
Thor, Moberg: Lichenes selecti exsiccati upsaliensis rial of Lichen lutereus Schrad. can be traced and it turns
107 (UPS,isoiypes in several herbaria, see Moberg out to belong to Bacidiu rubella in the present sense,
1991). then Bacidia luterea would be the correct name for this
?Lichen lutereus J . F . Gmel., Systeina Naturae, ed. species. This name has never been in use, however. To
13, tom 2, pars 2: 1359 (1792). - Lichen luteolus minimize the number of disadvantageous nomenclatural
Schrad., Spicil. Flor. German. 85 (17941, nom. illeg., changes, I suggest that Lichen lulerem J. F. Gmel, be
non J. F. Gmel. 1792 (p. 1368). (superfluous younger rejected, provided that future investigations prove the
homonym). - Lecideu Iuteolu (Schrad.) Ach., Meth. type of this name to agree with B. rubella auct. It should
Lich. 60 (1803), nom, illeg. (superfluous; including the be noted that Lichen Iuterem or a combinatioon based
types of both Lichen Iulereus Gmelin and Verrucaria on this name has never been in use for Bacidila rubella
rubella Hoffm.). - Biraaorca ludeola (Schrad.) fr., K . auct.
svenska VetenskAkad. Handl. 1822: 272 (1822), nom. According to ZahIbruckner (19261, also Lichen corti-
illeg. (superfluous). - Bacidia luteola (Schrad.) Mudd, calk Rutstr, is likely to belong to Bacidia rubella. If Li-
Manual Brit. Lich. 183 (1861), nom. illeg. (superflu- chen Iuterew J . F . Grnel, turns out to be taxonomically
ous). - Type: Not seen. different from Bacidia rubella, Lichen corticalis may be
?Lichen corticalis Rutstr., Spicil. Plant. Cryptog. the oldest name. Rutstrom's herbarium is probably lost,
Suec. 7 (1 794). - Type: Not seen, probably lost. however, and the description (Rutstrom 1794: 7) is very
vague ("leprosus, crusta farinosa, viridicinerea, tubercu-
Nomenclature. The oldest name, the present type of lis subsessilibus, depressiusculis, fermgineis"). Thus,
which is conspecific with B, rubella is Lichen verna- the identity of Lichen corticalis will most likely remain
lis L., better known as Biaaora vernalis ( L . ) Fr. (Jsr- an unsolved problem.
gensen et al. 1994a). Fortunately, this name has been In the protologue of Vewucaria rubella, Hoffmann
proposed to be conserved with a conserved type (1796) included Ehrhart's Plant. Crypt. Linn. 196 and,
agreeing with its present usage (Jargensen et al. indicated by the star to the lower left of the figure pre-
1994b). Lichen vernalis L. is the type species of Bia- ceeding the species name, some unspecified material
torn Fr.: Fr., and unless conserved, the generic name from the area of Gottingen (from the preface: "Asterisci
Bilatora will take precedence over Bacidia, and Bia- signwn, ubi antepositurn est speciebus, eas denotat,
quae circa Gottingen, atque in montibus Hercyniis in- Spores acicular, 3 1-104 X 2.1-4.3 pn,with 3-13 septa.
veniuntur."). I have, however, been unable to trace any
original material other than Ehrhart's exsiccate. The Thallus indeterminate, thin to thick, continuous or &S-
specimen in GOET, which is likely to have been availa- continuous, coarsely grandar, pale grey, greenish or
ble to Hoffmann, contains a very well-developed and yellowish grey, or greyish green, turning yeIlow or
abundantly fertile thaIlus of what is presently known as brownish in the herbarium. Granules (73-)92-139-218(-
Bacidia frminea Lijnnr. (Lonnroth 1858; see also Ek- 290) pm diarn. ( S 47, N= 19, n= 51, globose or usually
man & Nordin 1993). This specimen is selected here as f flattened, almost squamulose and slightly incised.
the Iectotype of B. rubella. Unless conserved with a Prothallus Iacking or present, endosubs~atal,white to
new type, however, the name Bucidin rubella will take pale grey.
precedence over B. fraxinea, a species which is much Apothecia (0.6-10.7-0.9-1.2(-2.0) mm diarn. (S= 0.2,
rarer and has'a much more restricted distribution than B. N= 19, n= 5), 4 plane when young, later becoming con-
rubella auct. In addition, the name B. rubella for B. m- vex, without or with thin to rather thick white pruina on
bella auct. will have to be replaced. AIthough earlier the edge andor disc of young and medium-aged apothecia.
name B. luteoda has been used for this taxon, B. rubella Disc usually brown-orange to orange-brown, sometimes
has been widely used for d least the last few decades. I pale orange, pink-orange, orange, pale orange-brown, or
suggest that Verrucaria P-ubellaBoffm. be conserved brown-red. Margin concolorous with disc or slightly
with a conserved type in order to preserve its present us- paler, level with or raised above disc in young apothe-
age, which also has the desirable effect of retaining the cia, later level with the disc, and finally excluded.
name Bacidia fpuxinea Lonnr. As a conserved type, I Proper exciple laterally (59-)79(- 103) pm wide (S=
suggest a collection distributed by R. Moberg in the ex- 12, N= 19, n= l), without crystals or sometimes with ra-
siccate Lichenes selecti exsiccati upsalknsis. diating clusters of minute crystals (up to 1 pm). Rim al-
most coIourIess or pale yellowish to orange. Lateral in-
Illwtra~iom.Figs 2A, 3D, 5D, 30 (map), 42A. terior usually f concolorous with rim or sometimes
slightly paler, along the edge without or with a single
Characterization. Thallus granular; granuIes 73-290 p cell layer thick zone of cells with enlarged lumina that
d i m . Apothecia usually brown-orange to orange- are up to 6 X 6 p.Medullaty part concoIorous with lat-
brown. Proper exciple, hypothecium, and hyrnenium eral interior part or slightly paler. Hypothecium almost
colour1ess to pale yellowish to orange, K+ intensifying. colourless or pale yellowish to orange, concolorous
Discurrsion. Neotropical B. russeola displays some vari- Characterizution. Apothecia f pinkish or pale orange.
ation in apothecium pigmentation. Most specimens have Proper exciple colourless, pale yellowish or pale brown-
Laurocerasi Brown in the hymenium and react K+ pur- orange; without crystals, along the edge with a l(-2) cell
plish, whereas some are almost or entirely deficient of Iayers thick zone of enlarged cell lumina. Hypothecium
this pigment, and thus do not react with K. Such speci- yellowish or pale brown-orange. Upper part of hy-
mens were named Bacidia subtestacea by Malme menium colourless or very pale brown-orange, without
(1935). The specimens from Texas, however, are very crystals. Spores acicular, 41-82 X 1.2-2.I p,with 3-1 5
similar to mainstream B. ~usseolafrom Central. and septa. Conidia only slightly curved.
South America. They are as well-developed as such
specimens, and the pigment Laurocerasi Brown is Thallus indeterminate, thin, either discontinuous, of dis-
present. crete or contiguous, convex, sometimes almost granular
Among the North American species, Bacidia mseo- areoles, or continuous, f cracked, wrinkled to warted,
/U is likely to be confused only with B. campalea (see pale green-grey to green-grey. Prothallus ofien present
that species) and brown-fruited morphs of B. schwein- between discrete areoIes and sometimes also bordering
itzii. Such specimens of B. schweiniaii differ from B. the thallus, very thin, endophloeodal, pale grey.
russeola in having a f brown hypothecium and proper Apothecia (0.3-j0.4-0.5-0.5(-0.7) mm diam. (S=
exciple. Furthermore, B. schweinibzii often has a granu- 0.1, N= 4, n= 1O), at first plane, later becoming mark-
lar thallus. edly convex, epruinose. Disc f pink, orange-pink,
The types of B. phaeoloma (Miill. Arg.) Zahlbr., B. brown-pink, pale orange, or sometimes pink-brown.
suturalis (Eckfeldt) Zahlbr., B. subaceri~loidesZahlbr., Margin concolorous with disc, raised above disc in
and B. papantlensis Vain. all are typical specimens of B. young apothecia, soon level with the disc, finally ex-
ncsseoia with Laurocerasi Brown in the hymenium. B. cluded.
pawula Malme, as lectotypified here, is a poorly devel- Proper exciple laterally (55-)69(-91) ,um wide ( S =
oped specimen of B. m s e o l a with mostly young apoth- 16, N= 4, n= l), without crystals. Rim colourless,
ecia. , along the edge with a l(-2) cell layers thick zone of
Bacidia chapadensis Malme, described from Brazil enlarged cells that are up to l l X 6 pm. Lateral interi-
by Malrne (1935), is very similar to B. russeola in or colourless or very pale yellowish to pale brown-or-
overall habit, but differs in having slightly longer and ange, rarely brown-orange. Medullary part colourless
wider, 9-17-septate spores and a distinct, 2-3 cell lay- to pale brown-orange. Hypothecium very pale ,yellow-
ers thick zone of enlarged cells along the edge of the ish to pale brown-orange, sometimes with heavier
proper exciple. This species is known only from the pigmented, brown-orange spots. Hyrnenium (72-175-
type collection. 81-88(-93) pm thick (S= 6, N= 4, n= 51, lower part
colourless except for occasionai vertical streaks of
Collections examined. U. S. A. Texas. Higginson (US). very pale brown-orange pigment, uppermost part col-
Brownsville, 1888, Pringle 59 (COLO), 1944, Runyon ourless or very pale brown-orange in a diffuse layer.
3883b (US). Paraphyses 1.0-1.2 pm wide in mid-hymenium; api-
ces f clavate or only weakly swollen, (l.&) 1.9-2.1-
2.3(-4.0) pm wide (S= 0.5, N= 4, n= 10), without in-
23. Bacidia salmonea S. Ekman, sp. nov. ternal pigment. Spores acicular, usually sigmoid,
sometimes straight or curved, coiled in ascus, (41-)
Type: Canada, British Columbia, Vancouver Island, 48-56-61(-82) pm long (S= 9, N= 3, n= IO), (1.2-)1.6-
Port Alberni, Roger Creek City Park, just E of the Ger- 1.7-1.8(-2.1) pm wide (S= 0.3, N= 3, n= 101, (24.1-)
trude St.-Pemberton St. intersection, open parkland, on 30.2-33.6-36.5(-66.0) times as long as wide (S= 8.5,
N= 3, n= l O), with (3-)6.8-7.8-9.4(-15) scpta (S= 2.5, (1926). - Type: Japan, "Fusijama, 6000 ped.", 1879,
N= 3, n= 10). Almquisr (H-NYL 17580, syntype).
Pycnidia half-iinn~crscdin thallus, uppennosl part of Lecideu ubdtrctuns Nyl., Lich. Japon. 68 (1890). -
wall concolorous with apothecia, c. 100 p 1 diam. Co- Bucidiu ubd~izrutuns(Nyl.) Zahlbr., Cat. Lich. Univ. 4:
nidia filiform or sligl~tlythicker at the middle, sIightly Cal kins 33 (H-
166 ( 1 926). - Type: Florida, "on IC)LS.PU",
curvcd, nun-septate, 1 1- 14 X 0.5-0.8 PI. NYL 1758l , lectotype selected here).
Cheinishy: No lichen substances detected. Bucidiu mt.lunocardia Zahlbr., Hedwigia 74: 201
Pigments: Piglnented parts of apothecia K+ intensify- (1934). - Type: China, "Muli: Mountain of Djago",
ing and N-. - Usually small amounts of Rubclla Orangc Rock (W, lectotypc selected hcrc).
in hymenium. hypotheciun, proper exciplc, and pycnid- Bacidia nigrn Zahlbr., Hcdwigia 74: 200 (1934). -
ial wall. Type: China, "Lidjiang snowrange, in the forests of
Ndaza KO, 11 300- 1 2000t", 193 1, Rock ( W , lectotype
Ecology. Preferably on young trunks, branches and selected here).
twigs, rarely on old trurtks In humid, shady to fairly
open forests and rorest edges. All finds are on Acer hromenclu~ure.Bacidia grmulosa Darb., Wiss. Ergebn.
maorophyilum and Popi~lintric-icltocurpa. Schwedisch. Siidpolar-Expedit. 4 ( 1 I): h ( 1 9 12): which
is not conspecific with Bacidia schweinitzii, blocks the
Di.~lribltlion.Along the coast from southcrn Orcgoi~to usc of Patellaria gr-anulosa Michx, in Bacidia. I have
southern British Columbia. - Probably endemic. not sccn thc type of the latter name, but according to
Miiller (1878) it is synonymous with Bacidia schwein-
D i s c ~ . ~ . ~ i Bucidio
on. B. arceutina,
salnior~eais similar to itzii.
B. rosella and scveral species of Brrciclinu with pale The events associated with the description of Biutol-a
apothecia. B. urcezrtinu can be separated by the presence schweinitzii are a complicated matter. In the protologue,
of a brown-yellow pigment in the apothecia, the unpig- it is mentioned as "B. Schweinitzii. Fr. MS." No indica-
rnented and entirely immersed pycnidia, and the strong- tion is given as to why it is named after Lewis David
ly curved conidia. B, rnsclla differs in having a crystal von Schweinitz or why the name is ascribed to Elias
layer in the epithcciu~n,in usually being less pig~nented Fries. The second time the name appeared was in Tuck-
than B. salmonm: and in having scptatc conidia. All spc- erman's exsiccate Lichencs Aincricae Septentrionalis
cies of Baciditza can bc scparatcd from B, salmonea by no. 136 (published in 1854). Here, it is citcd as "Biatora
the structure of thc proper exciple. Schweinitzii, Fr. hb., Tuckenn. in Darlingt. F1. Cestr.
edit, tert. p. 447." Tuckerrnan (1 872) treated il as Biatu-
Additional collt~utiuns auininerl. CANADA. British ra rubt'lla var. sohweinitzii but "Biut. suhweinitzii Fr.
Columbia. Vancouver Island, Port Alberni, 1977, Noble herb. Tuck. a s n. 136" is cited. Later on, Tuckerrnan
61 61 (herb. Noble). - U. S. A. Oregon. Curry Co., (1888) changed his mind and again decided to treat the
1989, Ek~nanL673 (LD). Lincoln Co., 1989, Ekman L taxon as a species and (like in the exsiccate) ascribe the
679, L680, L681 (LD). Tillamook Co., 1989, Ekman protologue in Darlington's Flom Cestrica to Fries and
L7 10, L7 11, L7 14 (LD). Washington. Jefferson Co., himself: "Fr. herb Tuckcnn, in Darlingt. F1. Cestr., edit
1989, Ekman L726, L727 (ED). Whatcorn Co., 1989, 3, p. 447, & in Lich. exs. n. 136". In FH-TUCK, there is
Eknlan, L743, L744, L746, L747, L748 (LD). a specirnen collected by Sc11weinit.z at Salern, North
Carolina, and marked "dedit mihi Fries" (= sent to me
by Fries) by Tuckerman. What appeals to havc. taken
24. Bacidia schweinitzii (Fr. cx E. Michener) A. place is that Schweinitz, during his time as a resident of
Schneid. Salem, North Carolina ( 1 821-35), sent a sample of a li-
chen to Fries, who provisionally named it Biatura sch-
Guide Study Lich. 110 (1898). - Biaturn schweinitzii Fr. weinitzii. Frics thcn passed the material on to Tucker-
ex E. Michener in Darlington, Flora Cestrica ed. 3: 447 man, who saved it for his own herbarium. There is no
(1853). - Biatum ~whellu var, schweinitzii (Fr. ex E. duplicate material preserved in UPS. Ezra Michener,
Michener) Tuck., Genera Lich. Ihh (1872). - Type: U. having discovered thc sainc species in Chester Co.,
S. A., Pennsylvania, "Chcstcr Co. 011 trunks.", Michen- Pennsylvania, was probably informed by Tuckeman
er's Lichens, book 5, pg. 7, specimen marked "126" about the existence of the name Biuturu suhweinihii,
(BPl, lectotype selectcd hcrc). and used it in his account of the lichens in Darlington's
?PafeIEariagranltlosn Michx., Flora Bor.-Americ. ed. Flora Ce.~trica(Darlington 1853). Probably mostly by
2: 320 (1803). - Type: Not seen. coincidence, Michener was the first to provide a valid
Lecidea abd;lcens Nyl., Lich. Japon. 68 (1 890). - Bu- description of Biatnra scltlveinitzii (the protdogue was
(Nyl.) Zalllbr., Cat. Lich. Univ. 4: 166
ciriiu r~bdilucer~s almost certainly written by Michener's hand). The mate-
Fig. 3 1. Known North
American distribution
o f Bacidia schweinitzii.
rial distributed by Tuckerman in his exsiccate soon unable to trace any of Almquist's material of this spe-
thereafter was collected "ad truncos (praesertim Betula cies in S.
excelsae, Fagique) in montibus Novae Angliae", and
has nothing to do with the material sent to him by Fries. IIlus~ations.Figs 8A, 3 1 (map), 42D-E.
Although the collection sent to Fries by Schweinitz
was the first one to bear the name Biabora schweinitzii, ChaructePization. Thallus granular or f rimose. Apothe-
it was almost certainly not seen by Michener. hstead, cia usually orange-brown to dark purple-brown to black.
Michener's diagnosis was based on his own material, Proper exciple at least in inner part orange-brown to
which consists of four collections now preserved in red-black, merging into the pigmentation of the hy-
BPI. One of these, a well-developed and typical speci- pothecium, without minute, evenly dispersed crystals
men, is here chosen as the lectotype. (although sometimes with a few clusters). Hypothecium
Barris (1977) treated Biatora rubella var. schweinih- pale orange-brown to dark brown. Upper part of hy-
ii in Tuckerman (1872) as a different n a p with a dif- +
rnenium either greenish or pale orange-brown. Spores
ferent type than Biatop-a schweinitzii Fr. ex Michener in acicular, 32-88 X 1.9-4.3 p,with 3-15 septa.
Darl. The fact that Tuckerman (1 872) cited his exsiccate
when he made Biatom rubella var. schweinitzii, and Thallus indeterminate, thin to thick, either discontinu-
that Darlington's Flora Cestrica in its turn is cited in ous, of discrete granules or low, convex areoles, or con-
the exsiccate, makes it fairly certain that Tuckerman tinuous, granular or not, then without cracks or f rimose
merely made a new combinations based on the same ep- to areolate, smooth to wrinkled to warted to tuberculate
ithet. or sometimes subsquamulose; grey to green-grey to
I refrain from lectotypifying Lscidea abducens Nyl., grey-green to brown-green. GranuIes f globose, cylin-
since @e only syntype studied is very poor. I have been drical, or flattened, (48-)53-94-121(-145) pn (S= 27,
25. Bacidia subincornpta (Nyl.) AI-nold,nom. cons. Illlrstrations. Figs 32 (map), 42F
prop.
Churucterization. Thallus granular or k rin~ose.
Flora, Jena 53: 472 (1870). - Leoideu subincornj>tcd Nyl., Apothecia purple-brown to black. Proper exciple at
Flora, Jena 48: 147 (1865). - Type: Austria, "An Stam- least IateraIIy with a mixture of greenish and brown-
1nchc11 v011 Sorb-blis uhun~ae~~re.rpilus
unterhalb der scr- isli hues. Hypotheciuln brown, always distinctly dark-
luswai~deober der Waldrast, 5400'. Matrci in Tirol", er than the proper cxciplc bclow, K+ purplish. Upper
1872, Arnold, A r ~ ~ o l dLicli.
: exs. 505 (H-NYL, typ. part of hymenium f green. Paraphyscs 1.2-2.4 pm
cons, prop.; LD, isotype). wide in mid-hymeniu~n.Sporcs fusiform, bacillifor~n,
Lecidea seppurahilis Nyl., Flora, Jcna 48: 147 clavate, or acicular, 19-64 X 1.9-6.2 pn, with 3-13
(1 865). - Bucidim separuhilis (Nyl.) Amold, Flora, Je- scpta.
tla 53: 472 (1870). - Type: Finland, Tavastia australls,
"Hollola", 1863, Norrlin (H-NYL 17424, lectotype se- Thallus indeterminate, rather thin to thick, either dis-
lected here). continuous, of discrete granules or low convex areoles,
Lecideu hegetschweilcri Hepp, Hepp: Systematische or continuous, without cracks nr f rirnose, wrinkled,
Sainn~lungen212 (1852), nom. nudu~n.- Biator-a afro- warted, tuberculate, or distinctly granular. Granulcs f
snnguineu var, heg~'t.vch~~cilet.i Hepp, Hcpp: Flcchten globose or flattened to subsquan~ulosc, (24-)51-74-
Europas no. 23 ( 1 853). - Bucidia hegetschweileri 106(-145) pm (S= 2 5 , N= I l , n= 5), light grey to grey
(Hepp) Vain., Acta Soc. Fauna Flora renn. 53 (I): 21 5 to ycllowish grey to green-grey to grey-green to
(1922), non auct. - Type: 1,Iepp: Flechten Europas no. brown-green to brown. Prothallus lacking or present
23 (BM, lectotypc sclccted here). between discrete areoles/granules, thin, endosubstratal,
white.
Nnmettclat~re.Lecideu subinca~n]~lu Nyl. was original- Apothecia (0.3-)0.4-0.6-1.0(-1.2) mm diam. (S= 0.2.
ly made as a noinen novum for Biufora a!msanguinea N= 18,n= 5), f plane when young, later becoming con-
(Schacr.) I-Icpp (Nylander 1865), since a co~nbination vex, epruinose. Disc purple-brown to brown-black to
based on the latter name was blocked by the already ex- blue-black to pure black, rarely pink-brown 01. f grey-
isting Lecideu utrusang~linca(Hoffm.) Nyl. Unfortunate- brown (f pigment deficient). Margin concoloroi~swith,
ly, Nylander based his interpretation of Biaforrr atrosun- or paler or darker than disc, grcy-brown to dark brown
guinea (Schacr.) Hcpp (the basio~lytll of which is to blue-black to pure black, rarely pink-brown or or-
Lecidea anon;alu rar. afru.sangiritreu Schaer.) on the ange-brown, distinct, level with or raised above disc in
specimens of Hepp's Flechten Europas 286 in H-NYL, young apothecia, later level w'ith the disc, and finally
which coinprises a mixture of Bacidia slrbincornpta in excluded in old and convex apothecia.
the present sense and B. circumsj~ecta.Lecideu arlomnla Proper exciple laterally (40-)56(-74) pm wide (S=
var, atrosunguineu Schaer. was originally bascd on 10, N= 18, n= l), without crystals. Rim colourIess to
Schaerer's Licheiles Helvetici exs. 2 12, which contaiils pale yellow, or orange-brown to red-brown to black-
nice speciil~elisof Bacidiu bzcnr~zpta(Bomcr cx Hook.) brown, often with a green or blue-green tinge at least
Anzi (syntypes in G and LD studied). Unlcss conserved, in upper part (giving rise to different hues uf brown-
the much used namc Bctciclia subiracot?apapan will disap- green or grccn-brown), along the edge with a single
pear into synonymy with Bacidia incoinpfa. The naine cell layer of of enlarged cells with lumina that arc up
Bacidia sllhincnmpta would then be replaced by Buuid- to 8 X 5 p n . Lateral intcrior concolorous with or dark-
ia separabilis (Nyl.) Alliold, a name which has hardly er than riin (never ei~tirelycolourless). Medullary part
ever been used. Ironically, the hasionym of this name, aIways paler than hypothecium and lateral interior,
Lecideci sepurr~bilisNyl., was described on thc samc colourless to pale yellow to pale brown to pale red-
page in the same work as Lccidecr suhincornpta Nyl.! brown. Hypothecium pale to dark (often reddish)
Magnusson (1936) was the first to list B. sepavabilis brown, sometimes with a green-brown tinge in upper
(Nyl.) Arnold as a synui~yinof B, subincumptu (Nyl.) part. Hymeniuin (45-j54-82- 104(- 112) pn thick (S=
Arnold. Arnold's Lich. exs. 505 in 13-NYL is here sug- 17, N= 17, n= 5 ) : lower part colourless except for oc-
gested as the coilservcd type of Bauidiu sirbinco~npta, casional vertical streaks of grccn to bIuc-grccn pig-
Fig. 32. Known North American distribution of Bacidia subincompta.
ment, upper part dirty green to blue-green, indistinctly n= 10), with (3-)3.0-5.8-9.7(-13) septa ( S = 2.3, N= 17,
or distinctly delimited. Paraphyses 1.2-2.4 pm wide in n= 10).
mid-hymenium; apices f clavate or not at all swollen, Pycnidia f immersed in thallus, uppermost part of
sometimes f moniliform, (1.6-)2.2-3.6-5.6(-6.5)pm wall concolorous with apohecia, c. 50 pm dian. Conid-
wide (S= 1.2, N= 19, n= 51, without internal pigment or ia filifom, curved, non-septate, 10-20 X c. 0.8 p.
with brownish grey internal pigmentation forming a Chemistry: No lichen substances detected.
very diffuse cap. Spores hsiforrn, bacilliform, clavate, Pigments: Upper part of hyrnenium R-, N+ purple
or acicular, straight to curved to sigrnoid, (19-)23-37- (with or without a precipitate of blue crystals). Apices
50(-64) pm long (S= 9, N= 17, n= 10), (1.9-)2.1-3.3 of paraphyses when with brownish pigment K+ pur-
-5.1(-6.2) pn wide (S= 1.0, N= 17, n= 10), (3.2-)4.6- plish, N+ orange-red. Hypothecium and proper exciple
12.1-19.2(-22.7)times as long as wide (S= 4.5, N= 17, K+ purplish, N+ orange-red. - Bacidia Green in hy-
tant origiual material appears to be a small specimen in brown or f purplish to black, sometimes mottled, at
PH marked "Lecid spadicea ns." This specimen, which least some pruinose. Proper exciple laterally pale yellow
is typical B, sums, was probably collected in Pennsyl- to black-brown (along the edge with a 4-6 cell layers
vania, where Muhlenberg was active as a clergyman. thick, distinct zone of enlarged cell lumina), without
Lecidea spadicea Ach. antedates Biatora suffusu Fr. by minute and evenly dispersed crystals (but o?en with
l l years..Unless an exception is given to the rules of clusters o f f large crystals). Hypothecium -t yellowish,
priority, the widely and correctly used name Bacidia K+ intensifying. Upper part of hymenium pale yellow to
sums will be replaced by Bacidia spadicea (Ach.) black-brown. Spores 38-91 X 2.5-4.3 pm, with 3-17
Zahlbr. Apparently, after the original publication, 30- septa.
cidiu spadicea has rarely been used, and never in the
correct sense. Instead, B. spadicea has been used for Thallus indeterminate, usually f thick but sometimes
various temperate, subtropical, and tropical taxa, none thin, continuous, without cracks, f rimose to areolate or
of which are conspecific with the type. Thus, to inhibit without cracks, smooth to wrinkled to warted to tuber-
further confusion, I suggest rejection of the name Leci- culate, or sometimes subsquamulose, f yellow-grey to
deu spadicea Ach. (Art. 56.1). green-grey or pale grey to grey. Prothallus lacking.
Apothecia (0.5-)Oh-1 -0-1.2(-1.5) mm d i m . (S= 0.2,
JIlus~ations.Figs 3C, 33 (map), 42G. +
N= 21, n= 5), plane when young, later becoming con-
vex, with thin to thick white pruina on edge andor disc
Characterization. Apothecia f yellow-brown, f orange- of at Ieast some young and medium-aged apothecia.
Woessia and Lichiugoidia, both described by Hawks- Lichenized species. Thallus crustose, episubstratal,
worth & Poelt (1986), antedate Bacidina by almost continuous, without cracks or rimose to areolate to
five years. Both genera were based on a single collec- granular, smooth to wrinkled to warted to subsquamu-
tion each in the anamorphic state referred to the type lose, sometimes discontinuous, of discrete to contigu-
species W. fusarioides and L. gyalectiformis. Both spe- ous areoles. Prothallus absent. Soredia and isidia ab-
cies were thought to be lichenized "conidial fungi", sent. Goniocysts common, sometimes covering most of
green to dark grey-green areoles, some of which dis- (1.0-)2.0(-3.2) pm wide (S= 0.5, N= 1, n= 20. Tholus
solve into granular goniocysts. Goniocysts pale green, with a narrow c-layer. Ocular chamber lacking or rather
loosely aggregated, (16-)24(-35) p diam. (S= 6, N= 1, wide and bIunt; axial body wide and high, conical, blunt
n= 20). or pointed, not penetrating through the entire d-layer.
Apothecia (0.3-)0.4(-0.7) mm d i m . (S= 0.08, N= 1, Spores acicular, clavate, or almost bacillifom, straight
n= 201, at first plane, later becoming convex, epruinose. to curved to sigrnoid, (30-)35(-43)p long ( S 3, N= 1,
Disk brown-yellow to yellow-brown to dark grey- n= ZO), (1.9-)2.2(-2.5) pm wide (S= 0.3, N= I, n= 20),
brown. Margin concolorous with disc, distinct, slightly (13.0-) 16.3(-20.0) times as long as wide (S= 2, N= 1, n=
raised above disc in young apothecia, soon level with 20), with (3-)5.1(-7) sepia (S= 1.6, N= l , n= 20).
the disc, fmally excluded. Pycnidia not seen.
Proper exciple laterdly 43 pm wide (N= l , n= l), Pigments: Apothecia K-, N-. - Arceutina Yellow in
without crystals. Hyphae with fairly thin walls (2.5- proper exciple, hypothecium, and hyrnenium.
5 pm between adjacent cell lumina), rather distinctly ra-
diating towards edge, with cell lumina of widely differ- Ecology. Bacidina aenea was found on the upper side
ing shapes, gradually expanding towards edge (in lateral of the leaning branches of a Fmaxinris caroliniana, at
interior 1-4 pm wide and up to 15 pm long, 3.5-10 the borderline between a small, open marsh and a sea-
times as long as wide). Rim brown-yellow, downwards sonally flooded hammock dominated by Sabal palmer-
gradually changing to colourless. Lateral interior to, Quercus virginiana, and Fruxinus curoiiniana.
brown-yellow, downwards gradually changing to col-
ourless. Medullary part colourless. Hypothecium pale Distribution. Known only from one locality in Sarasota
brown-yellow. Hymenium (7 1-173(-76) pm thick (S= 2, County in southwestern Florida.
N= 1, n= 51, colourless except the uppermost part,
which is brown-yellow. Paxaphyses rather abundant in Discussion. Bucidina aeaea can be separated from all
relation to the number of asci, 1.2 pm wide in mid-hy- other species of Bacidina by the presence of fairly Iarge
menium; apices narrowly clavate or not at all thickened, amounts of a brown-yellow pigment in the apothecia
Cl~aracterization. Thallus entirely dissolving into + 13 pn long, 1.5-7.5 times as long as wide. Rim colour-
green gouiocysts. Apothecia with k brown disc; margin less to palc brown. Latcral intcrior colourIcss. Mcdul-
often paler. Proper exciple colourless or very pale lary part colourless or vcry pale brown. Hypothccium
brown. Hypothecium brownish. Upper part of hy- orange-brown to dark red-brown. Hymenium (42-143-
lneniurn colourless or pale brown. Brown pigment K t 54-70(-74) pn thick (S= 9, N= 5 , n= 51, colourless or
green-brown. Spores acicuIar 35-48 X 0.9-1.9 pm, with pale brown it1 upper part. Paraphyses usually abundant
0-9 septa. Conidia filiform, curvcd, 0-l-scptate. in relation to the number of asci, 1 .O-1.6 pm wide in
mid-hymenium; apices f clavate or only slightly thick-
Thallus indeterminate, thin, starting as small, convex, ened, (1.6-)2.2-2.9-3.4(-4.0) pm wide (S= 0.7, N= 5, n=
discrctc or contiguous areoles (hat very soon dissolve 10). Tholus with a narrow c-layer. Ocular chamber usu-
into k coherent go~liocyststhat may or may not form ally narrow, low, and blunt; axial body widc and high,
coralline or isidia-like structures, grey-green, green- s~bc~liildrical, blunt, ilcvcr pcnctrating through the en-
grey, or pale green. Goniocysts ( 19-)28-32-40(-52) pm tire d-layer. Spores acicular, straight to curvcd to sig-
diam. (S= 9: N= 4: n= 10). inoid, (35-)39-40-42(-48) ,un~long (S= 3: N= 4, n= 10),
Apothecia (0.3-)0.4-0.5-().h(-0.7) mm diam. (S= 0. l , (0.9-)l .2-1.3-1.6(-1.9) pm wide (S= 0.2: N= 4:n= 10):
N= 4, n= 10), at first planc, latcr bccorning f convex: (22.3-)28.0-30.6-33.8(-45.7) times as long as wide (S=
epruinose. Disk grcy-brown to brow11 to dark rcd- 5, N= 4, n= 10), with (0-)1.5-3.2-6.4(-9) septa (S= 2.6,
brown. Margin often palcr than disc: sometimes con- N= 4, n= 1 O), straight or coiled in young asci.
colorous with or darkcr than disc, distinct, raised above Pycnidia immersed in granular thallus, colour~ess,75-
disc in young apothecia: soon level wilh the disc, usual- 125 pm diam. Conidia filiform, curved, 0-l-septate, 21-
ly persistent but sometimes finally excluded. 30xc. 1 . 0 ~ .
Proper exciple laterally (36-)j7(-77) p wide (S= 18, Pigments: Brown parts of apothccia K+ grcen-brown.
N= 5, n= I), without crystals. Hyphae with rather thin - Arnoldiaila Brown in propcr cxciplc, hypothecium,
walls (2-5 pm between adjacent cell lumina), rather dis- and hymenium.
tinctly radiating towards cdgc, with cell lumina oftcn of
different shapes, usually ilarrowcr in uppcr part than in Ecology. Mainly on bark and overgrowing corticolous
iowcr part, in latcral intcrior 1.5-7 PI widc and up to bryophytes on trunks and branches of trees and shrubs.
Once coIlected on the wood of a log. It favours shady aggregated granules (goniocysts). Goniocysts (16-)22(-
to semi-open, always humid habitats. Known phoro- 29) p diam. (S= 5,N= l, n= 10).
phytes in the western part of the distribution are A l m s Apothecia (0.3-)0.4(-0.6)m diam. (S= 0.09, N= 1,
mbra and Sambucus callicarpa. The phorophyte in n= 10), plane, epruinose. Disk pink to brown-orange to
North Carolina is unidentified, but is, according to the purple-brown, often mottled with different colours.
label, a "deciduous tree". In Europe, Ea. arnoldiana is Margin concolorous with disc, distinct, level with the
frequently found on shaded, more or less basic rocks. disc.
Proper exciple laterally 60 p m wide (N= 1, n= l),
Distribution. Known from the Appalachian mountains without crystals. Hyphae with rather thin walls (2-3 pm
of western North Carolina and the coast from Oregon between adjacent cell lumina), rather distinctly radiating
to British Columbia. - Europe. towards edge, of widely differing shapes (almost glo-
bose-ellipsoid-rectangular-elongate), in laterd interior
Discussion. Only likely to be confused with Bn, egenu- 1.5-6 pm wide and up to 15 pm long, mostIy 2-5.5
/a,which also has a distinctly brown hypothecium. Un- times as long as wide. Rim f orange-brown. Lateral in-
like Bn, arnoldiana, Bn. egenuiu has a green pigment terior f orange-brown. Medullary part colourless. Hy-
(Bacidia Green) in the epithecium and a brown, R+ pothecium colourless or very pale yellowish. Hymenium
purplish pigment (Laurocerasi Brown) in the proper ex- (62-)70(-74) pthick (S= 5, N= l, n= S ) , colourless in
ciple. Bn. cal~yornica (q.v.) is the only other North lower part, 1- diffusely and unevenly orange-brown in
American species producing Arnoldiana Brown. upper part. Paraphyses rather abundant in relation to the
number of asci, 1.2 pm wide in mid-hymenium; apices
Collections examined. CANADA. British Columbia. f clavate, (1.6-)2.5(-3.6)p.wide (S= 0.7, N= 1, n=
Vancouver Island, Ucluelet, 1981, Noble 7299 (UBC). 201, without internal pigment. Tholus with a narrow c-
- U. S. A. firth Carolina. Haywood/Transylvania Co., layer. Ocular chamber wide, rather high, blunt; axial
1981, Brodo 23794 & Gowan (CANL). Oregon. Tilla- body wide and high, conical or subcylindrical, blunt,
mook CO., 1989, Ekman L713 (LD). IYmhington. Sk- never penetrating through the entire d-layer, surrounded
agit Co., 1989, Ekman L737 (LD). Whatcom Co., 1980, by a narrow layer that is darker amyloid than the re-
Ryan (WWB). mainder of the d-layer. Spores acicular, straight or
curved, (36-)39(-43) pm long (S= 3, N= 1, n= 10), (1.9-)
2.3(-2.5) pm wide (S= 0.3, N= 1, n= 10), (14.5-)17.2(-
30. Bacidina assulata (Korb.) S. Ekman, comb. nov. 22.7) times as long as wide (S= 3, N= 1, n= 101, with
(5-)7(-9) septa (S= 0.9, N= I , n= 10), straight or coiled
Bacidia rubella var. assubata Korb., Parerga Lich., in young asci.
fasc. 2: 131 (1860). - Bacidia assudata (KM.) Vgzda, Pycnidra immersed in thallus, colourless, c. 150 pm
Lich. sel. exs., fasc. 23, no. 566 (1967). - Type: Po- diam. Conidia filiform, curved, non-septate, c. 10 X 0.5
land, Silesia, "Schreibershau", 1835, Flotow (L, lacto- W.
type selected here). Pigments: Yellowish to orange-brown parts of proper
Secoliga arceutina f. inter-mediaHepp ex Stizenb., exciple, hypothecium, and hymenium K+ intensifying,
Nova Acta Acad. Caesar. Leop. Carol. 30 (3): 42 N-. - Rubella Orange in proper exciple, hypothecium,
(1863). - Bacidia intermedia (Hepp ex Stizenb.) Ar- and hymenium.
nold, Flora, Jena 54: 54 (187 l), nom. illeg., non Hampe
in A. Massal. 1861. - Type: Not designated; isosyntype Ecologv. Collected on the bark of a Quercus sp. in a
iii LD studied. floodplain forest dominated by Qakercus and Calya.
Illzkstmtions. Figs 5G, 35 (map). Distribution. Known from a single locality in northeast-
ern Oklahoma. - Central and eastern Europe. Reports
Churacterizution. Thallus dissolving into, goniocysts in from Scandinavia, the British Isles, and the Mediterra-
small spots. Apothecia brown-orange to purple-brown, nean are based on misidentifications.
often mottled. Proper exciple laterally f orange-brown.
Hypothecium colourIess or yellowish. Hymeniurn high, Discussion. Bacidina assulata is an often misused and
f orange-brown in upper part. Pigment K-t- intensifying. misunderstood name. It has, above all, been used for
Spores acicular. 36-43 X 1.9-2.5 pm, with 5-9 septa. specimens of Bn. phacodes, Bn, urnoidiana, pigment
Conidia filiform, curved, non-septate. deficient forms of Bacidia subincompta, and an un-
described Mediterranean species of Bacidio S. str.
Thallus indeterminate, f continuous, thin to rather Apart from the North American specimen, I have
thick, f warted, in small spots dissolved into loosely seen Bn. assulata only from Central and East Europe.
Ecology. 011 trcc trunks, mainly on rough bark in shady Ill~lstrations.Figs 4A, 5J, 35 (map).
or open, coastaI forests wit11 high air humidity. Known
phorophytes include Ae.5-clrlus califnrniclls, L'~tpr.~sslls IVomenclufirre. The lectntypification made by Diederich
nzacrocurpu, and Uwbellulnria culjfofnmica. et al. (1991) is not valid, since the collection chosen (H-
NYL 17871) is not original material. It was collected in
Distrihzrtic~n. Known from a few localitics in Marin, 1878, the year after the publication of Lecidea clalorn!-
San Mateo, Montcrey, and Santa Barbara Counlies in iutlla Nyl.
the south ;~ndccntral parts of coastal California. - En-
dcmic? Characterizatiora, That lus k dissolvii~ginto goniocysts.
Apothccia vcry small. will1 pale yellow to pale grey
Disc~xsian.From San Luis Obispo Co., there is a fur- disc; margii~ paler, persistent. Proper exciple, Iiy-
ther specimen (Ekman L621, LD), which 1 have not at potliecium, and upper part of hymenium (almost) col-
the moment included in BIT.califirnica. It diffcrs froin ourless. Spores acicular, 17-3 1 X 0.9- 1.2 p,with 0-3
other specimens of B. californi~amainly i l l having nar- septa. Conidia filiform, curved, 0- l -septate.
rower spores (1.2-1.9 p n ) and in having difrerent CO-
nidia (filiform, straight or slightly curved, non-septate, Thallus indeterminate, f thin, continuous, rimose, wriu-
27-73 X 0.8-1.0 pn). It was collected on the upper side kled or warted, grey-green or green-grcy, nut dissolving
of a Icaning branch of Qlterr.cnu ~rgr.(fdiain a rather dry or partly diffusely dissolvi~~g into palc grccn goniocysts.
coastal oldgrowth. Furlher collections are needed to es- Goniocysts loosely aggrcgatcd, (28-)35(-48) pn diain.
tablisl~whether it should be included in Bn. cali{orniccr, (S= 5, N= l , n= 10).
or i f it represents a distinct species. Apothecia (0.15-10.19-0.20-0.2 l(-0.36) mm diam.
Rucidikina cal/fornica is siinilar to Bn. usslllrta (scc (S= 0.05, N= 2, n= 10). at first plane, later sometimes
this species) and Bn. rumea. The lattcr spccics is sepa- becomlng moderately convex, epruinose. Disk pale yel-
rated from Bn, calfornica by the K+ inteiisirying or K+ low or pale grey. Margin paler than disc, milky white or
purplish {not K+ green-brown) pigments in the apotl~e- very pale yellow: distinct, slightly raiscd above disc in
cia and the tci~dcncyto produce a tha1line margin in the young apothecia, soon level wit11 the disc, pcrsistwt.
young apothecia. Specimens of Bn. uulif~wniculacking Proper exciplc laterally (24-)29(-33) p n wide (S= 6,
goniocysts and pigment in the apothecia are very similar N= 2, n= l), without ciystals. Hyphac with veiy thin
to the European Bn. yhacuclcs, the type species of the walls (0.5-1 p n between adjacent cell lumina), not dis-
genus. Rn. phucoclE.r is then separated mainly by the dif- tiilctly radiating towards edge, with cell lumina homo-
ferent conidia (see discussion under Bn. assula!cr) and geileously shaped, in lateral interior 2-8 pn wide and up
the thinner Ilymenium (usually less than 50 pm thick). to l5 pm long, 1.5-4 times as long as wide. Rim colour-
less or very pale brown in uppermost part. Lateral interi-
Additional col/ectioras ~xunzined.U . S. A. Califurnia. or colourless. Medullay part colourless. Hypothecium
Marin Co., 1925, Parks 2883 (COLO), 1989, Ekman colourless. Hyrneniurn (37-)38-43-48(-5 1) pm thick (S=
L639 (LD), 1989, Ekman L645 (LD). Moilterey Co., h, N= 2, n= 5 ) , colourless or vcry palc brown in upper
1984, Bratt 4271, 4277 (herb. Bratt), 1989, Ekman palt. Paraphyses few in relation to thc nuinbcr of asci,
L632 (LD). San Mateo Co., 1967, Jordan 822 (SFSU). + 1.6-2.0 pn~wide in mid-hymeniuin; apices f clavate or
Santa Barbara Co., 1994, Nimis & Tretiach (filcd with only slightly thickened, (2.0-)2.4-3.2-4.1(-6.5) pn wide
B. inszrluris, TSB). (S= 1.2, N= 2, u= 10). Tholus with a narrow c-layer. Oc-
ular cha~nbcrlacking; axial body wide and high, subcy- iilcniuin tenuius sporaeque breviores ct latiores sunt;
lindrical, blunt, never penetrating through the entire d- ambarurn apothecia crystallis careni.
layer. Sporcs acicular, straight or slightly curved to sig-
moid, (17-)24-26-28(-31) p 1 Iong (S= 4, N= 2, n= 10). Etyt~ology.C t y s ~ a l l ~ e v zcarrying
~s, crysials.
(0.9-) 1 .O- 1.1 - 1.2(- 1.2) p n widc (S= 0.2, N= 2, n= l D),
(1 6.0-)23.3-23.9-24.5(-30.0) timcs as long as wide ( S = I/lu,strutions. Figs 5K-L, 3 h (map), 44C.
4. N= 2: 11= 1O), with (0-)0.6-1.2- 1.8(-3) septa (S- l. l ,
N= 2, n= 10). Clruructrriz(ition, Apothccia f pink. Proper exciplc col-
Pycnidia immersed in thallus, colourless, 75- 100 pm ourless, at least in the upper part with minute, evenly dis-
diam. Conidia filiform, curved, 0-l-septate, 30-43 X persed ciystals along the rim. I-Iypothecium (almost) col-
0.8-1.0 pn. ourlcss. Upper part of hynleniuin colourLess but with a
Pigments: All parts of apothecia K-. - Sometiines laycr of minute crystals between the apices of the paraphy-
very small arnouilts of an unidentified brown pigment in ses. Spores acicular, 32-72 X 1.2-1.9 p n , wit11 3- 13 septa.
the proper exciple and hymenium.
Thallus indeterminate, thin, either continuous, smooth
Ecology. Collected once on the smooth bark of Fagzls or warted, slightly rimose or without cracks, or discon-
grundi#hlia and once on rotten wood (probably a log or tinuous, of disci-ete or contiguous, convex, sometiincs
a stump). almost granular, areoles.
Apothecia (0.2-)0.2-0.3-0.3(-0.5) mm diam. ( S = 0.06:
Distl-ibzltion.Collected twice in soutl1e1-n Ontario. --.Eu- N= 3, n= 1 D), at first planc, latcr becoming convex, at
rope. lcast when young with thin pruina. Disk pale pink or
pale orange-pink. Margin concnlol-ous with disc, dis-
Di.~cussion.Buciclinu chlorutio~tlais a rathcr distinctive tinct, level with or raised above disc in young apothccia,
species with its pale grey or pale yellow apothecia with soon level with the disc, finally excluded.
persistent margin. 1t is only likely to be confuscd with Proper exciple latcrally (37-)44(-5 l ) p1 wide (S= 5 ,
Bn. egenulcr and Bn. egenuloid~m.The former is scparat- N= h, n= l), without pigment, a t least in the upper pail
ed from Bn. chlouoticzllu by the presence of a green pig- with minute and evenly disperscd ciystals along the rim
ment (Bacidia Grccn) in the upper part of the hymenium (up to l pm). Hyphae with rather thin walls (2-3.5 pm
and by a brow11 hypothecium. Bn. egenuluiclea is above bctween adjacent cell lumina), not distinctly radiating
all distinguished by thc prcsence of a brown, K+ pur- towards edge, with cell lumina that gradually expand to-
plish pigment (Lauroccrasi Brown) in the proper exci- wards edgc (in lateral interior l .5-3 pn wide and up to
ple, and by the presence of diffcrcnt conidia. 12 pm long, usually 2-4 times as long as wide). Hy-
The type of Bucidiu subchlorodica (Nyl.) Flagey is a polllecium colourless or very pale yellowish. Hymenium
typical specin-ten 01'Ruoidinu chloroticula. (56-159-66-70(-74) p n thick (S= 5, N= 5, n= 5), colour-
less, at least partially with minute crystals in the upper
Collections examined. CANADA. Utttari-iu. Norwich, part (usually as a distinct, contii~uouslayer). Paraphyses
1978, Wong 2709A, 2712 {CANL). Ottawa, 1897, Ma- abuildailt in relation to the number of asci, 1 .O- 1.4 ym
coun 280 1 (CANL). widc in mid-hymenium; apices upwards gradually thick-
ened. (1.6-)2.0-2.1-2.2(-3.2) p n wide (S= 0.4, N= 5, n=
10). Tholus with a rather wide c-layer and a d-layer that
33. Eacidina crystallifera S. Ekrnan, sp. nov. is characteristically tmncate at the apex. Ocular cham-
ber lacking; axial body very wide and high, conical,
Type: U. S. A., Louisiana, "Tunica Hills norih of St. pointed with straight or slightly bulging sidcs, often
FrancisvilIe: wooded ravii~eson Murrill ButIer property, penetrating through the entire d-layer, the sidcs of the
on gravel road l. mi. S of La, route 66; turnoff .25 mi. axial body the11 collapsillg and becoming concavc.
W of junction o r State 66 and U. S. route h l , 30°49'N Spores acicular and straight, curved, or sigmoid, (32-)
9 1"24'W, West Feliciana Parish, covering vine of mus- 40-46-52(-72) pn long (S= 7, N= 4, n= 10)- (1.2-)1.4-
198 1 , Tucker 2 1520 (herb.
cadine (Vifis r.nttr~td~/i)lia)", 1.6-1.7(-1.9) p n wide (S= 0.3, N= 4, n= 10), (20.0-)
Shirley Tuckcr, holotype; LD, isotype) 26.5-29.9-33.4(-44.6) times as long as wide (S= 5, N= 4,
n= 1O), with (3-)4.h-5.7-6.7(- 13) septa (S= 2.4, N= 4, n=
Thallus nunquam in goniocystas solutus. Apothecia ro- 10).
sea, pignierlto destituta vcl quasi, strato crystallifero in Pycilidia not scw.
cpithecio et in margine excipuIi proprii pracdita. Facile Pigments: A11 parts of apothecia K-. - Sornctimes
cum Bacidina aeneu confusa, cuius apothccia pigmen- small amounts of an unidentified pigment (Rubella Or-
turn fulvurn habent et culn Bacidina vavia, cuius hy- ange?) in the hypothecium.
Ecology. On young tree-trunks and vines in shady and epruinose. Disk pale brown to purple-brown to black or
humid forests. Known phorophytes include Fagus gran- occasionally. pale
- -pink, often mottled with different hues
dfolia, Liquidambar s@ructflzaa, Vilis rotundifoiia, and (at least some apothecia f black). Margin concoiorous
Wisteriafiutescens. or darker than disc, in upper part pale brown-purple to
black, in lower part pale pink to pale pink-brown, occa-
Distribution. Known from a few IocaIities in Florida, sionally entirely pale pink, distinct, slightly raised above
coastal Georgia, and southern Louisiana. - Endemic? disc in young apothecia, soon level with the disc, finally
excluded.
Discussion. Bacidina cqvstallfera is an easily recog- Proper exciple laterally (2 1-)28(-34) pm wide (S= 9,
nized species due to its crystal layer in the upper part of N= 2, n= l), without crystals. Hyphae with thin walls
the hymenium. However, in some apothecia the crystals (1-3 pm between adjacent cell lumina), rather distinct-
are rather sparse, and if overlooked, it may be taken for ly radiating towards edge, with cell lumina that are nar-
non-goniocystate morphs of Bn. varia. Apart from hav- rower in upper part than in lower part, in lateral interi-
ing crystals in the hymenum, Bn. cqvstallifera is separat- or 1-5 p- wide and up to 10 pm long, 3-7 times as
ed from Bn, varia in having a higher hymenium, and long as wide in upper part, 1.2-1.7 times as long as
longer and thinner spores. Due to the almost complete wide in lower part. Rim grey-brown to black-brown in
lack of pigmentation and the presence of crystals in the upper part, downwards changing to orange-brown to
apothecia, Bn. ctystallifefe~.a
has some resemblance with colourless (entirely colourless in pale pink apothecia).
Bacidia rosella. Bn. crystallifem, however, has smaller Lateral interior grey-brown to black-brown in upper
apothecia, narrower spores and lacks an ocular chamber. part, downwards changing to colourless or very pale
orange (entirely colourless in pale pink apothecia).
Additional collections examined. U . S . A. Florida. Ala- Medullary part colourless. Hypothecium f orange-
chua Co., 1.993, Ekman E1 154 (LD). Georgia. Chatham brown. Hymenium (40-)40-44-47(-50) pm thick (S= 4,
Co., 1974, Tucker 12224b (herb. Tucker). Louisiana. N= 2, n= 5), colourless in lower part, upwards usually
East Baton Rouge Par., 1982, Tucker 25593 (herb. gradually darkening to pale green, grey-green, or
Tucker), 1993, Ekman L1 120 (LD). East Feliciana Par., green-brown. Paraphyses rather abundant in relation to
1970, Tucker 8478b (herb. Tucker). West Feliciana Par., the number of asci, 1.6 pm wide in mid-hymenium; ap-
1981, Tucker 2 1520 (herb. Tucker). ices f clavate, (2.4-)2.8-3.1-3.4 (-4.9) p wide (S=
0.7, N= 2, n= 10). Tholus with a narrow c-layer. Ocular
chamber rather wide, high, blunt or pointed; axial body
34. Bacidina egenula (Nyl.) VEzda wide and high, conical or subcylindrical, blunt, never
penetrating through the entire d-layer. Spores acicular,
-
Folia Geobot. Phytotax. Praha 25: 432 (1991). Lecidea
egenula Nyl., Flora, Jena 48: 147 (1865). - Bacidia
straight or curved, (16-)21-23-25(-3 1) pn long (S= 5,
N= 2, n= 10), (1.2-11.4-1.5-1.6(-1.9) pm wide (S= 0.2,
egenula (Nyl.) Arnold, Flora, Jena 53: 472 (1870). - N= 2, n= 10), (10.0-)15.1-15.3-15.5(-20.0) times as
Type: France, "Fontainbleau" (H-NYL 17967, lectotype long as wide (S= 3, N= 2, n= lO), with (0-1.1-2.0-2.8(-
selected here). 7) septa (S= 2.0, N= 2, n= 10).
Pycnidia immersed in granular thallus, colourless, 60-
Illustra~ions.,Figs 5M, 36 (map), 43D. 75 pm diarn. Conidia filiform, curved to sigmoid, 0-1-
septate, 9-24 X c. l .O p.
Thallus entirely dissolved into gonio-
Characte~~ization. Pigments: Greenish parts of hymenium K-, N+ purple
cysts. Apothecia pale brown to black (at least some black). without a precipitate of blue crystals. Dark brown parts
+
Proper excipIe laterally brown (K+ purplish), below hy- of proper exciple and hymenium K+ purplish, N+ or-
an&-red, orange or orange-brown parts of proper exci-
pothecium colourless. Hypothecium f orange-brown. Up-
per part of hymeniwn greenish. Spores acicular, 16-31 X ple and hypothecium K+ intensifying and N-, or K+
1.2-1.9 pm, with 0-7 septa. Conidia filiform, curved to sig- slightly green-brown and N-. - Laurocerasi Brown in
moid, 0-1 -septate, not smngly conglutinated. proper exciple and hymenium. Rubella Orange in proper
exciple and hypothecium. Arnoldiana Brown in the hy-
Tnallus indeterminate, thin to rather thick, entirely dis- pothecium of some apothecia. Bacidia Green in hy-
solved into granules (goniocysts), continuous or of dis- menium.
crete to contiguous granules, grey-green or green-grey.
Goniocysts coherent, (24-)45-49-52(-77) pm diam. (S= Ecologv. At the locality in Louisiana, it was collected on
12, N= 2, n= 10). a hematite rock in a pasture, whereas in Ontario, it was
Apothecia (0.2-)0.2-0.3-0.4(-0.5) mm diam. (S= 0.1, found at the base of an UImw aamericaaa. In Europe, it
N= 2, n= 10), at first plane, later becoming f convex, is usually found on shady, siliceous rocks.
hues (K+ intensifying), below hypothecium colourless. whereas type 2 and 3 occur mixed in the same pycnidia.
Hypothecium (almost) colourless. Upper part of hy- Pigments: Brown parts of proper exciple and hy-
menium pale orange or pale brown. Spores acicular, 20- meniurn R+ purplish, N+ orange-red, pale orange to or-
42 X 0.9-2.1 pm, with 0-5 septa. With three types of co- ange-brown parts of proper exciple, hypothecium, and
nidia, the coinrnonest one filiform, straight or slightly hyrnenium R+ intensifying, N-. - Laurocerasi Brown in
curved to sigmoid, 0-l-septate, strongly conglutinated. proper exciple and hymenium. Rubella Orange in proper
exciple, hypothecium, and hymenium.
Thallus indeterminate, thin, entirely dissoIved into gran-
ules (goniocysts) that are f coherent and form isidia-like Ecology. At the only known locality, it was collected
or coralline structures, grey-green. Goniocysts (2 1-)35- both on a granite rock and on "old wood" in an "open
46-57(-77) pm diarn. (S= 15, N= 1, n= 10). wood".
Apothecia (0.2-)0.3-0.3-0.4(-0.4) mm diarn. (S= 0.06,
N= 2, n= 10), plane, epruinose. Disk pale orange to Rislribubion. Known only from the type locality in
brown-orange to dark purple-brown. Margin concolor- Ohio. - Endemic?
Ecology. The Gainesville locality is a more or less Illustrations. Figs 2B, 4B, 5Q, XC, 39 (map), 44F.
swampy forest and adjoining, open and grazed areas
with a few solitary trees, both dominated by Pianera Chumcterizution. Thallus dissolving into goniocysts or
aquotica. Here, Bn. sguumellosa is particularly abun- not; goniocysts coarse, 17-46 pm d i m . Apothecia pink
dant on trunks of Planera aquatica and Cephalunthus to purple-brown. Proper exciple laterally colourless to
occide~ltalisthat are subject to periodical flooding. It is pale red-brown, below hypotheciuin colourless, without
not likely that Bn. sqriainellosa can survive a flooding large inflated cell lumina in Iower part. Hypothecium
that may last for several years, however. Instead, it is colourless to pale red-brown. Bymenium colourless to
probably a fast colonizer once the water has subsided. pale brown-orange. Paraphysis apices thin, 1.6-2.7 pm
The Myalcka River locality is similar in being a swamp (average of 10). Spores acicular, clavate, or almost ba-
forest, but here, Bn. squarnellosa was found on shaded cilliform, 19-50 X 1.2-2.5 p~, with 3-7 septa. Conidia
tree trunks of Fruxinzcs caroiiniana well above the high- filifom, f curved or sigrnoid, usually indistinctly 3-1 1-
water level. septate.
DisMbution. Known from two localities in southern and Thallus indeterminate, thin, usually discontinuous, of
central Florida. - Endemic? discrete or contiguous, convex areoles, sometimes con-
tinuous, wrinkled or warted, usually f rimose but some-
Discmsion . Bacidina squarnellosa is an inconspicuous times areolate or entirely without cracks, pare grey to
species, but easily identified on account of its finely squa- grey-green, often (but not always) partly or entirely dis-
mulose thallus. However, the thallus of depauperate solving into granular goniocysts, then becoming rather
morphs may be confused with the coralloid goniocysts thick. Goniocysts pale green to green, loosely aggregat-
Proper exciple Not prominent, outer pal-t byssoid, Pruminenl, raiscd abovc thallus,
spreading over the lllallus surthcc, modera~elybyasoid: not sprcading
gradi~allythinncr towards edge ovcr thc thallus
pigment (the latter patchily distributed) in the epi- i bears nu close relationship with Bacidia tm-
author, L
thecium o f darkly pign~entedapothccia. In B. sahule- chonu.
toruin, on the other hand, the spores often posscss a
warted pcrisporc and more than three septa, the para-
physes often cxcecd 2 p n in width, and the epi- Bacidia niveocincta G . Merr. in Zahlbr. & Redinger,
thecium of dark apothccia is dominated by a green nom. nud.
pigment.
Lich. Rar. exs. 3 1 1 ( 1933). - Biutora niveocinctu C;. Merr.,
Merrill: Lich. exs. 270 (1913), nom. nud.
Lecania naegeIii (Hepp) Diederich & P. Boom
Evcil though this species has been distributed in two ex-
Bull. Soc. Nat. luxcmb. 95: 154 (1994). - Biutum siccates, its name has not been validated. It may, how-
nuegelii Hepp, Ilepp: Flecht. Eur. 19 (1853). - Bncidia ever, plwvc to have been validly publisl~tdundcr a dif-
nnrgelii (Hepp) Zalilbr.; Ost. bot. L. 59: 439 (1909). - ferent name. Bacidiu niveoci~lclai s a c~r~icolous spe-
Type: Not seen. cies oCFeIIhanem, apparently closely related to thc foli-
icolous F. stnnhopeae (Miill. Arg) Liicking, also known
This coiticolous species has traditionally been named as Badinlicd stanho~?eae(Miill. Arg.) Vgzcla (see Liick-
Bacidia naegelii, but belongs in Lecaaia; see the sec- in:: et al. 1994: 400).
Lion Related genem atad spccie.~groups in the chapter The material distributed in the two exsiccates is prob-
Taxononlic delirizitutions above species 1~.1>el.North ably part of the same collection made by Rapp in 1907.
American speciincns arc generally correctly deter- Bacidia niveocittcto was listed by Lainb (1963).
mined.
Zahlbruckner (June 1926) madc Bcrcidia pcrllidula as a Bacidia rubidofusca (Willey) Zahlbr.
ilolllei~novum: sincc Bacidicr pnllida (Miill. Arg.) Darb.
1912 was blocking the use of a combination based on Cat. Lich. Univ. 4: 139 (1926). - Biuturu rubicliijiiscrr
Biatora ynllida Arnold ex EckIeldt Unforlunatcly, Zahl- Willey, Enum. Lich. New Bedford 23 (1892). - Type:
bruck~~ei- forgot that the name Baoidia pnllidula was al- U. S. A., Massachusetts, New Bedford, Willey (US, hol-
ready occupied for a quite different species made by otypc, not seen by me).
himself a few ~nontlisearlier (Zalilbruckner Feb. 1926).
Bacidia pal/ididlda Zahlbr. (June 1926) is a younger taxo- I have not seen ally material of this spccics, but accord-
nomic synonym of Scnlicios/)nr~~~r trmhrinurn (Ach.) ing to Printze11(1995: 215) it probably bcloilgs in Gval-
Arnold. idea. rt is apparently a teiricolous lichen ("on naked
earth", Willey 1892).
This corticolous species appears to be rather widcsprcad This species was recently treated by Kalb & Staiger
in boreal North America. For a discussion on its cxclu- (1995) as Ophioparma herrci. 1 agree with their conclu-
sion fro111 Bucidiia, see the section Relnfed genera arad sion that this spccics should be included in Ophiopur-
ma. It is similar to the type species of the genus, 0. ven- cies, but it is distantly related to Bacidia and Bacidi-
tosa (L.)Norman, and differs mainly in having a thinner na.
thallus with a more or less granular surface and a differ-
ent ecology. 0,rubrdcosa is characteristic of dry, hard
lignwn and acid bark. It is found from southern Califor- Bacidia scopulicola (Nyl.) A. L. Sm.
nia to southern British Columbia (Noble 19821, and in
humid areas of the western slopes of the Rocky Moun- Monogr. Brit. Lich. 2: 156 (191 1). - Lecidea scopulico-
tains. la Nyl., Flora, Jena 57: 312 (1874). - Type: Not seen.
This saxicolous species is excluded from Bacidia This species, long known as Bracidia subfuscula, be-
and Bacidina on account of the very wide and richly longs in Lecrania; see the section Related genera and
branched paraphyses, the occurrence of oil droplets species groups in the chapter Taxonomic delimitations
in the hymenium, and a very peculiar tholus. It is al- above species level. It is usually terricolous or saxi-
most entirely non-amyloid, but contains a faint, cen- colous, but the type of Biatora siberiensis was collected
tral structure comprised of a small cone just above on bone.
the spore mass with a long, pointed extension at its
apex. I do not understand the affinities of this spe-
Mycologia 4: 131 (1912). - Lecidea microphyllina var. Forteckn. Skandinaviens vhter, 4. Lavax 38 (1936). -
subgranlrlosa Tuck., Proc. Am. Acad. Arts Sci 6278 Bilimbia vereczknduiu 1
' 11. Fr., Lich. Scand. l: 387
(1864). - Type: Not seen. (1874). - Type: Norway, " W. Finm.: Alten, Bosekop",
1864, Th. M. Fries (UPS, lectotype selected here).
A lectotype in FH-TUCK was selected by Lois Brako
in 1987, and, according to her label, synonymized with Bacidia verec~ndulais a member of the Bacidia beck-
Psorella perrexta (Nyl.) Miill. Arg. To my knowledge, hausii group; see the section Related genera and spe-
the lectotypification has not yet been validly pub- cies groups in the chapter Taxonomic delimitations
lished. It was reported as Biatora subgranulosa from a above species level. It is similar to Bacidia heinipolia
single locality in Georgia by Tuckermai~(1888), but 1 (Th. Fr.) Malme but differs in having shorter and wider
have not examined this specimen. Lecidea micro- spores. It is known only from the corticolous type col-
phylliaa var. szkbgrunulosu Tuck. should not be con- lection from n o r - e mNorway. It has been reported from
fused with Lecidea parvifolia f. subgranulosa Tuck., Northwest Territories, Canada, but this record refers to
described in the same work. The latter is a synonym an undescribed member of the Bacidia sabuletorum
of Phyllopsora canounthrina (Vain.) Brako (Brako group.
1989).
LD). - D. B. auerswaldii, X 78 (Ekman L788, LD). - E. B. biatorina, young apothecium, X 115 (Ekman L661, LD). F, B. -
campalea, crystals brilliant white, X 108 (Ekman L1 117, LD). - G. B. circumspecta, X 98 (Ekman L634, LD). - H. B. d~fracta,
x 70 (Ekman L1050, LD, holotype).
141
Fig. 42. Apothecial sections of North American species of Bacidia. Specimens collected in North America were used, unless
otherwise stated. -A. B. rubella, note clusters of crystals in proper exciple, X 83 (Shchepanek 203a, CANL). - B. B. russeola, X
93 (Brazil, Malme: Lich. austroamer. Regnell no. 243, LD). - C. B. salmonea, X 88 (Ekman L710, LD). D. B. schweinitzii,
-
morph with green epithecium and almost black hypothecium and proper exciple, X 60 (Wetmore 35510, MIN). - E. B.
schweinitzii, morph with brown epithecium, hypothecium, and proper exciple, X 75 (1963, Weber, COLO-L36608). F. B. -
subincompta, X 88 (Wetmore 47929, MIN). - G. B. suflusa, X 60 (Wetmore 31371, MIN). - H. B. vermifera, X 103 (Anderson
2825, COLO).
41
Opera B o t 127, 1 O n L
Riatora 5.39-40,32.57:74,70,82,86,8Y:94.99, 1U7. 127-134
Index B intoraccne 40
Biatorina 39
Only names of lichci~shave been included Unless other- biatoi-ina* 8 , 10, 12, l9,2 1,23,28-29,31,34,59,67-68,96
wise statcd. epithets of species and taxa of lower rank biformigera,Tylothallia 45
are treatcd in Bcluidiu or Rnciditto (but combinations in Bilirnbia hh. 12Y-132, 134-135
othcr genera may also occur). Names that have been lec- breviuscula. Phyllopsora 42
totyplfrcd, ncotypirted. or suggested for conservation brialmontii. Tha~nnolecai~ia
brouardii
with a conserved type in this work h a r c bccn mal+kcd Hysroloma 56
with an asterisc. Refercncc in bold is to taxo~lo~nic
treat- bysso~noi~~ha, Aui-iculura 38
ment. caesiovi~~is 10.34-35
californica 9, 14. 16-17,.
19-23.25.28-29,33,59-60,113, 116, 117-118, 123, 127,133
californica, Waynea 45,46
abbrcvians 80 califumicui~~, Haerna~omma 133
abduccns 99- 100, 1 02 caligans . 125
abd~~crans* 99, 102 cnloosensis* 79-80
absistens* X. 12, 13, 19-21.23.25-30,32,34,57-5XXhl-63.74.92 Calnpodia 4
accedens (Amold) Lcaau 46:127, 134 Caloplaca 38
acceder~sscnsu Harris 46.127,134 campalea* 8. 12,
acclit~is 38 13. 15- 19-21-23?27-29. 32-34. 59,63,68-69,82,98. 109
Adelulecia 38 Candelariella 38,42
adscendeiis, Waylea 35 canoumbrina, Phyllnpsora 135
aenen 14-17, 19,21,23,78-29,33,59, 113-115, 125, 126 carneoalbidn. 42.128
aggrcgatula * X. 12.15. 19-21.23.27-29,33-34-58-63-64, 101 camcola, Pachyphiale 4
ahlesii. Lccidea 39 Catillaria 4,41:XO:87: 129
akompsa* 127 Catillariaccac 46
alaskensis, Hertclia~~a* 40, 127 Catinaria 39-40
albcsccns 127, 133 chapadensis 98
albidula, Myrioiiora 42 chlornntha, Ropalospora 56, 128
albohyalina, Lecidea 130 cl~lot.ococcurn,Scolicinspor~un 42-43
alpinn, Schadonia 42 chloroticula 8, 1 1 -
alutacca 76 12. 14-17, 19-21-23.25-31,59. 112. 114. 118-1 19. 121-122
alvearialis. Schistopltlca 42 cinel-cuvii-cns.Toninia 43
arnericana 46 circunispecta* 12: 1 8-24:
apiabica 8, 127 26, 28-29, 32,34,44.58,69-72, X l-82,92: 101, 105, 1 12
a~eulirla* 16-1 7, 19-2 1,23.28->0,32.34,59,64-hh,99. 1 15 clcn~cntis 128
archboldianurn. Con~psocladium 40 Cliostonium 40,57,133
amoldiana* 8, coccocnrpioitles, Ro[fidium 42
10, 1 1 , 14, 16-17, 19-21.23.27-30.32,5RR 112- 114, 11.5-116 coclcstit~a.Toninia 45
aromatica, Toninia 44 cognata, Psorella 43
Aithl-orhaphis 5 coillbii~ans 33.64
Aithoniales 5 compaginala, Phyllnpson 42
Arthmsporun~ 5 : 37-38, 57: 133 Co~npsocladiun~ 38
assulata8 8,l4,1&17,19,21,23,28-29:3I,hO,114,116-118 copladcs 130
atlat~tica* 76-78 corrug~ti~m. Cliosto~nuiii 40
atra,Tcphron~cla 44 corticalis, Lichen ' 94
atrugrisea 82.83. 127 crystallifersr 14-2 1.23.25.28-3 1,59,93, 1 15.1 19- 120. 125-126
atropurpurea, Catinaria 39-40 cuprca, 1-ccania 41,129
atrosanguinca. Biatora, Lccidea aiiomala var. 105,126 curvatuni. Scoliciospomiii 42
auerswaldii* 8, 1O, 12. 15, 18-2 1.23,28-29, 3 2 . 3 4 , 8 , 6 6 - 6 7 cuyabci~sis 33
augustioii* 56. 127 cyrtt.lla, Lccania 22.41
Auriculora 38 cprtellina. Lecnnia 22,41
Bacidiaceae 38.41,41,44.46 dcclir~is* 129
Bacidioniyccs 60 dittracta 8. 10.
Bacidiopsoia 5,38--39 12: 19-21.23.27-29,32,34.47-48.58,72-74,N.b 109
Bactrospon 56. 128 dirnerelloides, Tibellia 44
Badinia 4 Ect~idt~ocyrnbium 40
baglicttoana 8,128 eckfcldtii 78
Bapalmuia 38-39 Ectolechiaccac 4.5
bcckllilusii* 45,57, 128, 135 efilsa*
berengeriana. Lecidea 39 eKi~sella*
egenula* 8, 11, ioessa 131
14, 16-17, 19-21,23,24,27-31,58, 115-1 16, 119, 120-122 jacobi* 57,131
egenuloidea* 8,11,12,14,16-24,27-31,60,119,ltl-122 janeirensis, Squamacidia 43
elaeochroma, Lecidella I2 Japewia 38,41
endoleuca 82, 127 kingmanii I31
endoleucula 82-83 kolaensis, Adelolecia 38
epixanthoides 42, 129 laciniosa 42
erysibe, Lecania 129 laeticolor I29
Eschatogonia ,38,40 laurocerasi 13, 15,
euploca Speerschneidera 43 18-2 I , 23,28-30,32-34,48-5 1,6O, 6 5 , 7 5 - 7 6 , 7 82-85, 127
exilis,Teloschistes , 33 Lecanactis 56,127
fagicola, Pachyphiale 130 Lecmia 5,38,41, 57, 132, 134
Felllunera 4,56, 129, 132 Lecanora 33,38,43,68
finkii* 129 Lecanoraceae 3 8,42,46
flavens* 129 Lecidea 4,6 1,64,66,68,69,79-80,82,
flavicaus, Teloschistes 33. 86,88,94,99,105, 107,110,118, 120,127,129, 130,134-135
flavidulu~n,Cliostomutn 40 Lecideaceae 4,44
floridana, Fellhanera* 129 leprosum, Cliostomum 40
fraxinea 34-35,95 leucophlebarurn, Byssoloma 131-132
fiiesiana* 8, 19,21,23,28-31,34,57,62,67,74 Lichen 93-94,134
fusarioides, Woessia 10,112,115 Lichingoldia 4,112
fusca 135 Iisowskii, Fellhanera I29
fuscella, Lecania 41 Iobulata, Toninia 46
fusconigrescens 32,69,109 hflammia 4
fuscorubella* 88,127 Lopadium 4
fuscula 46 Loxospora 56
gallurae, Scoliciosporum 42 lutereus, Lichen 94
globulosa, Catillaria 129 Iuteola Iuteolus 94-95
granosa* 130,135 luteolum, Cliostomum 40
granulifera 87 lutescens 45-46
granulosa, Patellaria 99 rnajuscula* 79-80
gririthii, Cliostomum 40 meadii, Byssoloma* 131-132
grossa, Catinaria, Megalaria 39,41 medialis* 15-17,19-23,2&29,33-34,58,80,#-X7
Gyalecta 56 Megalaria 38,39,41
Gyalidea 133 Megalospora 56
gyalectiformis, Lichingoldia 112 megapotamica 78
gyalizella, Pachyphiale* 130 meiocarpa, Lecidea 130
gypsacea, Squamarina 44 melanocardia* 99,102
Haernatomma 68,133 Micarea S, 46,56
hegetschweileri* 105, 110-l11 microcmpa, Mycobilimbitl 46,131
helicospora 8, 12, 15, 19-23,28-30,33-34,60,74-76,78,83 microphyllina, Psorella 43
hemipolia 45,135 millegrana 32-33, 69, 109
herbarurn 8,130 mirabilis 11
herrei, Ophioparma 133 molybditiss 86-87
Herteliana 5,40-41, 127 montagnei, Eschatogonia 40
heteroclma* 8, 10, 12, 15- montana, Catinaria, Lecidea vernalis var. 39
21,23,26- 30,33-34,51-53,60,64,69,76-79,83, 117, 127 muralis, Lecanora 44
holomelaenum, Scoliciosporurn 43 mutabilis* 8,
horista 33,64 12, 15, 19,21,23,25,27-29,33-34,59-60,64,79,57-88,93
hostheleoides' 8,15,19-23,28-29,33-34,58,79-80,87 Mycobilimbia 5 41-47 46,57,128-129,134135
hutchinsiae, Lecania 41 Mycoblastus 41
hypno~um,Lecidea 39 Myrionora 38,42
Hypocenonlyce 56 naegelii, Lecania 41, 57, 87, 132, 134
idahoensis 19-20,27-29,32,48-51,60, 84,85-86 neglecta 1I2
igniarii* nigra* 99,102
19-24,28-29,31,34-35,44,58,66,71,80-81,9 101, 112 nigrocIavata, Catillaria 129
illudens 8,44, 130 nivalis 132
impudens, Cecanora 46 niveocincta 132
incompta 34-35, 105, 130 obscura, Toniniopsis* 60-61, 130
insularis* 8,12,19-21,23,28-30,33-34,58,62,67,81-82,127 obscurata, Mycobilimbia 41-42,46, 131, 135
intermedia 116 OpeiYapha 11
inundata 8,18, 112, 130 Ophiopama 5, 133-134
invertens* 82-83 Pachyphiale 5,56, 130