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OPERA .
..- .

Number 127 - 1996

Stefan Ekman

The corticolous and lignicolous species of

? .
Bacidia and Bacidina in North America
The corticolous and lignicolous species of Bacidia
and Bacidina in North America
The corticolous and lignicolous species of Bacida
and Bacidina in North America
Stefan Ekrnan

Opera Botanica 127

Copenhagen 1996
Opera Botanica

The corticolous and lignicolous species of Bacidia

and Bacidina in North America

Stefan Ekrnan

Ekman, S. 1996. The corticolous and lignicolous species of Bacidia m d Bacidina in

North America. - Opera Bot. 127: 1-148. Copenhagen. ISBN 87-88702-2 1-9

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.

Stefan Ekman, Department of Systematic Botany, Universiv of Lund, 0. Vallgntan

18-20, S-223 6 I Lund, Sweden.

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

Accepted 25-2- I996

Q OPERA
BOTANICA

Opera Bot. 127, 1996

Acknowledgments drawings and the habitus photographs, and Louise and
Patrik Frodtn spent more time than they had testing and
First of all, 1 would like to thank Iilgvar Kiknefelt, who trouble-shooting my key to the species of Becidiu and
has been my supervisor and given me continuous sup- Bacidina. I am very gratehl to Per Magnus Jmgensen,
port throughout my time as a student at the Department who has assisted me in solving a number of complicated
of Systematic Botany in Lund. I would also like to ex- nornenclatural problems. 1 think Klaus Kalb for direct-
press my siilcerest gratitude to Irwin Brodo, Brian Cop- ing my attention to the name Lecaaora campalea, on
pins, RoIand Moberg, and Rolf Santesson for help and which we are together basing a new combination. Ann-
support in many more ways than there is room to relate Margret Sandberg, Henrik Dagnegird, Ingrid Magnus-
here. Twin, Brian, and Eeif Tibell have scrutinized the son, Ingrid Larsson, and Kurt Lindberg are thanked for
manuscript and come with many suggestions for im- technical assistance. Furthermore, I want to express my
provements. For hospitality during my stays in the Unit- gratitude to Charis Bratt, Shirley Tucker, Klaus KaIb,
ed States and Canada, and for help in many other ways, Willa Noble, and the curators of B, BERN, BG, BM,
I want to thank Margie Holmes, Charis Bran, Harry CANL, COLO, E, F, FH, FLAS, G , GLAM, GOET,
Thiers, Bruce McCune, Wilfred Schofield, Willa Nob- GZU, H;HBG, IMI, L, LAM, M, MICR, MW, MSC,
le, Trevor Goward, Clifford Wetmore, Joe Ammirati, MU, NY,OSC, PC, PH, S, SFSU, STR, TSB, TUR,
Katherine Glew, Irwin Brodo, Stephen Clayden, Rich- W C , UPS, US, USAS, W, WELT, WTU, WU, and
ard Hartis, William Buck, Shirley Tucker, Dana Griffin, WWB for arranging loans (or otherwise helping me to
Danyl Grund, the rangers at Lignumvitae Key State Bo- trace important collections) of the herbarium specimens
tanical Site, and the late Mason Hale. I also want to ex- on which this study is based. The Florida Department of
press my sincere gratitude to all my present and former Natural Resources is acknowledged for allowing me to
colleagues at the Department of Systematic Botany. In collect lichens at the Lignumvitae Key State Botanical
particular, my thanks goes to Ulf Amp, Louise Lindb- Site. Financial support for this work was received from
Iom, San-Eric Mattsson, and Lars Froberg for interesting The Royal Swedish Academy of Sciences (Karin och
and endless discussions and constructive criticism, Per Axe1 Binnings fond, Hierta-Retzius' stipendiefond,
Eassen for providing me with the Latin diagnoses, for Hierta-Retzius' fond for vetenskaplig forskning), The
invaluable help with the nomenclature, for helping me Royal Physiographic Society of Lund, The Lund Botan-
to translate numerous Latin texts, and for requesting a ical Society, Anna och Gunnar Vidfelts fond for biolo-
large number of loans on my behalf, Stefan Andersson gisk forskning, and from Lund University (Battramska
for help with statistical problems, and finally the late resestipendiet, Knut och Alice WaIlenbergs stiftelse, C.
Ove Almborn for support and for putting his impressive F. 0. Nordstedts fond, Axe1 Hallstroms donation, Mur-
lichenological library at my disposal. Ulf made all the becks minnesfond).
Introduction sense is unknown, but 1 would roughly &mate it to 60-
90 species worldwide. A vast majority of them has
When De Notaris described Bacidia in 1846, he includ- probabIy already been described. Most are probably cor-
ed only B. rosella (Pers.) De Not. and the species pres- ticolous, but some inhabit various other substrata.
ently known as Pachypkiale carneola (Ach.) Arnold Bacidina is one of the segregate genera from Bacidia
(De Notaris 1846). In the following years, the genus sensu Zahlbruckner, but contrary to many other such
was variously treated. Numerous species were described genera, Bacidina is closely related to Bacidia S. str. The
as new or combined into Bacidia by Arnold, Lettau, and history of this genus is a short one. It was first recog-
Vainio. Other authors, for example Nylander, did not nized as the "Bacidia phacodes-Gruppe" by V k d a
accept the genus at ,all. Bacidia was typified with B. (1983) and was later formally described as Bacidina by
rusella by Fink (1910). It was with Zahlbruckner's Cat- VCzda (1991). Bacidina is, however, antedated by
a l o lichenum
~ universalis that Bacidia received its Woessia and Lichingoldiu, both described in the same
first consistent and comprehensive treatment (Zahl- publication (Hawksworth & Poelt 1986). Until now, 17
bruckner 1921-1940). In Zahlbruckner's taxonomy, Ba- species (including one which I consider to be a syno-
cidia, with a some exceptions, comprised all crustose li- nym) have been formally referred to Bacidina (VCzda
chens with a chlorococcoid photobiont, biatorine or 1991, Lumbsch & VEzda 1992, Farkas & VSzda 1993,
lecideine apothecia, and spores with three or more VEzda 1994, Wirth 1994). Six new species and two
transverse but no longitudinal septa. Altogether, he re- combinations in this genus are added here. Woessia and
ferred 647 accepted species and a large number of addi- Lichingoldia, on the other hand, have (with one excep-
tional synonyms to Bacidia. Similarly, he referred spe- tion) not at all been used in taxonomic or floristic litera-
cies with non-septate spores to Lecidea, species with ture following their publication. The reason why these
one-septate species to ati ill aria, and species with muri- names did not receive any attention is that the authors
form spores to Lopadium. These genera included 1486, obviously were not aware of describing the Bacidia
304, and 110 accepted species, respectively. Lecidea, phacodes group, and nor was it their intention to do so.
Catillaria, Bacidia, together with Toninia, Rhizocar- An attempt to introduce the use of Woessia was recently
pon, and a number of small genera, constituted the fam- published by Serusiaux (1995). In my opinion, a rees-
ily Lecideaceae in the sense of Zahlbruckner. tablishrnent of Woessia would be a clear violation of the
Zahlbruckner's Catalogus has been the most influen- spirit of the present International Code of Botanical No-
tial work of 20th century lichenology, despite the fact menclature (Greuter et al. 1994). Thus, it is suggested
that it was soon realized that the system presented was that Bacidina be conserved against Woessia and Lichin-
highly artificial. For example, Santesson (1952) re- goldia. Pending the final solution of this matter, the use
marked that Zahlbnlckner's system "has become domi- of Bacidina is retained here.
nating in a way strongly detrimental to the development Contrary to most species of Bacidia, those of Bacidi-
of a more natural lichen classification". Regarding Ba- nu are often very inconspicuous. They are small,
cidda, he noted that the delimitation was probably un- "scurfy", often sparingly or not at all fertile, and hence
natural. they are likely to escape attention. On a worldwide ba-
Even in our enlightened time, the ghost of Zahlbruck- sis, aImost half of the presently known species are folii-
ner haunts lichenoIogy. Practically every contemporary coIous. Several species are exclusively corticolous, but
lichen flora, to some extent, still uses a more or less some can be found on practically every substratum
Zahlbrucknerian concept of Lecideu, CaHllariu, and Ba- where competition from other lichens is low, including
cidia. This is, of course, not done in the belief that this such odd ones as brick, rusty iron, and leather: The 24
system is natural, but for practical reasons and since de- presently known species in Bacidina probably constitute
tailed knowledge of many species is still wanting. only a small part of the true number. Many species re-
Since the publication of Catalogus lichenum univer- main to be discovered.
salis a large number of species have been transferred to
other genera, which are not seldom distantly related to Objectives
Bacldia s. str. as typified by B. roseUa (Fig. 1). Particu-
lar attention has been paid to the foliicolous members of The objectives of this study are to provide (1) a taxo-
Bacidia sensu Zahlbruckner, many of which have been nomic revision of thc North American corticolous and
referred to genera in the families Pilocarpaceae and Ec- lignicolous taxa belonging in the genera Bucidia (in the
tolechiaceae, e.g., Fellhanera, Badirnia, Calopadia, Lo- strict sense) and Bacidina, and (2) a preliminary hypoth-
flarnmia, Sporopodium, and Tupellarju (Hafellner 1984, esis on the circumscription of these genera, based main-
Vgzda 1986, Liicking 1992, Malcolm & VGzda 1994; ly on the North American taxa studied.
see also compilation by Farkas & Sipman 1993). The At a critical point in my work during early 1994, 1
number of species referrable to Bacidia in the strict was choosing between delimiting my work the way

Opcra Bot. 127, 1996

 in the) (genera in the\
-: jEctolechiaceaej
yilocar pace-at

-
/
(~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-

Optra Bot 127, 1996

ues (the values within brackets). If, instead, one uses an (Wilkinson 1992). Details of the methods used as well
average of n measurements (ofien 5 or 10, rarely 20), as definitions of the twelve phytogeographic regions are
the same comparison should be made using the lowest found in the chapter Phytogeography.
and highest specimen arithmetic mean (the values flank-
ing the mean). The range of variation of the latter meas- Taxonomy
ure is invariably smaller than the range from the mini-
mum to the maximuin value. Thus, it is often worth the The distinctness of Bacidia polychroa vis-a-vis B. dif
effort to use an average of n measurement instead of fractu, B. lanirocerusi subsp. laurocerasi vis-a-vis sub-
single measurements. It definitely results in more relia- sp, iduhoensis, and western vis-g-vis eastern popula-
ble determinations. tions of B. heterochroa were investigated using two
Measurements, both in the dissecting microscope and statistical methods, one-way analysis of variance (one-
in the light microscope, were usually at first made with way ANOVA) and chi-square, and two numerical meth-
one scale division as the unit. Later on, when preparing ods, canonical variate analysis (CVA) and principal
the description, the unit "scale divisions" was multi- components analysis (PCA), the latter based on a Pear-
plied with the appropriate factor to obtain the measure- son correlation matrix among the original characters. In
ments in p or mm. This has caused some of the nmeas- the PCAs, the combination of characters giving the best
urements in the descriptions to appear rather odd. For separation was chosen. All calculatioiis were performed
example, the reader will notice that the figure 1.6 pm with the SYSTAT 5.2.1 statistical package (Wilkinson
appears rather ofien in the descriptions (ofien for the 1992), except the ANOVAs of Bacidia luurocerasi S .
width of the paraphyses). This is explained by the fact lat., which were performed with SuperANOVA (Abacus
that 1.6 p exactly corresponds to two scale divisions Concepts 1989).
when using the 100x lens in my light microscope! The CVA, also known as canonical discriminant
analysis or simply discriminant analysis, and the PCA
Chemistry are similar in producing a number of orthogonal, linear
axes (called canoilical variates or principal components)
Secondaty metabolites were investigated using high per- composed of the original characters, i.e., a kind of com-
formance thin layer chromatography (HPTLC) accord- posite characters (Dunn & Everitt 1982). A PCA arrang-
ing to the method described by Amp et al. (1993). Ba- es the axes in order of decreasing amount of total vari-
cidiu ,friesiana, B, igniarii, and Bacidina assulata were ance explained, each maximizing the amount of
excepted due to sparsity of material, otherwise a11 spe- variance. In a CVA, on the other hand. each axis maxi-
cies were studied. Between one and l l specimens per mizes the separation between a nurnbkr of pre-defined
species were studied, in a 1 110 specimens. groups along each of the axes.
The measurements used in the data analyses are basi-
Distribution cally the same as the ones used to prepare the descrip-
tions of the species, the only exceptions being that I
In the discussion on phytogeography and in the distribu- have included eight specimens of European B. laurocer-
tion maps, I have consistently ignored literature records, asi in one of the analyses and that I have excIuded spec-
since they are not reliable. imens from which I have been unable to extract a full
set of measurements of all the characters used (hence
the slight discrepancies between the species descrip-
Numerical and statistical methods tions and the tables in the chapter Ntimericnl and statis-
tical treatment). When more than one measurement of a
Phytogeography character was made on a specimen (i.e., when n> l), the
arithmetic mean of the n measurements was used in the
The similarity in composition of Bucidiu species be- subsequent analyses. Prior to the analyses, the distribu-
tween twelve phytogeographic regions (nine North tion of the measurements of each character was studied.
American and three European) was studied. The 27 No distribution was found to be skew enough to violate
North American and four additional European species the assumptions of the methods used. Thus, no transfor-
of Bacidia were scored as present ( l ) or absent (0) in mations were found to be necessary, nor did I have to
each of the twelve regions. The similarity in species exclude any characters for this reason. Methodological
composition between each pair of the regions were then details of the individual analyses are presented in the
calculated using the simple matching dichotomy (S4) chapter Numerical orad statisticul treatment.
coefficient. The resulting similarity matrix was used in a
single Iinkage cluster analysis. The calculations were
performed with the SYSTAT 5.2.1, statistical package

Opera Bot. 127. 1996

 Nomenclature for North America. Bn. upiahica is a foliicolous species
found in the Gulf States. It has been reported to be corti-
The abbreviation of authors follows Brummitt & Powell coious on twigs on occasions (Santesson 1952), bivt I
(1 992). Names that have been lectotypified, neotypified, know of no such finds from this area. North American
or suggested for conservation with a conserved type in coIlections of Bn. inundata are exclusively saxicolous,
this work have been inarked with a11 asterisc in the in- but in Europe it is soinetimcs found on the bark of roots
dex (see p. 145). and tree bases ncar water. The saxicolous, terricolous,
and muscicolous species of Bacidia and Bacidina are still
Taxonomic concepts rather poorly known, and further species should be ex-
pected.
A taxonomic species coilcept has been used (Cronquist
1988: 71). In my opinion, species boundaries are
marked by correlated discontinuities in the variation of Morphology and anatomy
two or more independent characters, i.e., by the absence
Thallus
(or at least extreme rareness) of certain combinations of
character states. A genus is a monophyletic assemblage
Growth form
of species. It is an abstraction without characters of its
own; it is defined by the species included in it. Since The thalli of all treated species of Bacidia and Bacidina
genera have no independent existence in nature, their are lichenized, and crustose or (in Bacidina squamello-
boundaries (and hence their sizc) are somewhat arbi- sa) finely squamulose. In many species, the thallus is
trary. In my opinion, genera should be operational to more or less continuous, often more or less rimose. On-
other than the specialist(s) on the genuslgenera con- ly seldom it is areolate or entirely without cracks. A
cerned. number of species possess a discontinuous thallus, con-
sisting of discrete to contiguous areoles. In several spe-
cies with a normally continuous thallus, young parts of
The North American Bacidia and the same thaIlus or less well-developed specimens may
Bacidina - an outline be more or less discontinuous.
In seven species of Bacidia, B. auerswaldiii, B. biato-
To my present knowledge, 27 species of Bacidia (one of r i m , B. diffruuta, B. hostheleoides, B. mbellra, B. sch-
which is divided into two subspecies) and 12 species of weinitzii, and B. subinoon~pta,the tliallus is commonly
Bacidina occur on bark and lignum in the continental or exclusively granular. The granules are homologous to
United States and Canada. Apart from the nine species areoles, but contrary to normal areoles the granules are
described here as new, 16 are reported (correctly) for not necessarily attached to the substratum by their lower
the first time from this area: Bacidia absistens, B. ag- surface. Instead, they may form obliquely upwards from
gregahtla, B. atkep.swaldii, B. bialorina, B. campuleu, B. old areoles (which may or may not be attached to the
Ji-iesiana (previous reports erroneous), B. heterochma, substratum), thus causing the thallus to become several
B. igniarii (previous reports erroneous), B. insularis, B. granule Iayers thick.
mzktabilis, B. reagens, B. russeola, Bacidina arnoldiana Bucidina squamellosa is the only species in the TWO
(previous reports erroneous), Bn. assulata (previous genera that has a squamulose thallus. These squamules
reports erroneous), Bn. uhloroticula, and Bn. egenula are small, deepIy incised, and vay koin prostrate to as-
(previous reports erroneous). cending. When prostrate, the squamules are virtually in-
Among the corticolous and lignicolous species treat- separable from effigurate areoIes, which indicates that
ed here only three have so far been collected in North the squamlllose habit is easily derived from a crustose
America on soil or rock as well, viz. Bauidiu s u f i s a , one (or vice versa).
Bacidina egenula, and Bn. egenlr/uidea. A number of
exclusively terricolous, saxicolous, or foliicolous spe- Prothallus
cies have been reported for North America, however. In
Bacidia, one marine saxicolous species, B. scopulicolo All species of Bacidina treated here lack a prothallus,
(Nyl.) A. L. Sm., and three terricolous/muscicolous spe- whereas it is present in many species of Bacidia. rt is

h cies, B, bugliettoana (A. Massal & De Not.) Jatta, B.

herbumm (Stizenb.) Arnold, and B. illudens (Nyl.)
Lynge, have been reported. My preliminary findings
show that these deterrninations are correct. Two addi-
usually endosubstratal, very thin, pale grey to white, and
present as a diffuse zone along the edge of the thaIlus
and sometimes also between discrete areoles. rn six spe-
cies (B. aggregatula, B. canzpalea, B. helicospora, B.
tional species of Bacidina, Bn. apiahica (Miill. Arg.) heterochroa, B. mutabilis, and B. russeola), on the other
Zahlbr. and Bn; inundutu (Fr.) VEzda, have been reported hand, the prothallus forms a narrow, episubstratal, black
Fig. 2. Thallus, transverse sections stained with LCB, viewed in interference contrast. - A . Bacidiu rubella (Ekman L960, LD),
transverse section of thallus granule showing brilliant white crystals in cortex of upper side. Bar = 30 p.- B. Bacirdina variu
(Elanan L1 191, LD), section through thallus producing goniocysts in chains that become almost coralloid or isidia-like. Bar = 100
pm. - C. Bacidina califurnica ( E h a n L632, LD), transverse section khowing algal layer (dark), stainable layer (grey), and
epinecral layer @ale). Bar = 10 pm.

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

Opera Bol, t27. 199h

invariably within a species. Subsquamulose individuals, can species, and also the European B. caesiovirens S.
Iike the term implies, have projections high and large Ekrnan & Holien (not treated here). In these species, the
enough as to give the thallus an almost squamulose ap- entire surface of the thallus granules (i.e., also the upper
pearance. one) is identical in anatomy to the lower, ecorticate sur-
In some species, the surface structure is more or less face of B. dz@racta and B, rllbena (Fig. 2A). Possibly,
strongly affected by the presence of goniocysts; see they are not only identical in anatomy, but even truely
Vegetative dispersal below. homologous. B. allerswaldii, B. biaaorina, B. sllbincoomp-
PseudocyphelIae are not present in any species of Ba- tu, and B. caesiovirens prefer humid environments. Per-
cidia or Bacidina, nor is a crystal layer on the surface of haps the lack of a crystal inspersed epinecral layer has
the thallus. Crystals.may be present in the cortex, how- been an advantage in such environments, possibly since
ever (see below). gaseous exchange is facilitated and the risk of pro-
longed desiccation is minimal.
In Bacidino squamellosa, the only known species in
Cortex the genus with a squamulose thallus, the upper as well
as lower surface of the squamules is very similar in
An upper cortex is present in all species of Bacidia and anatomy to the species discussed above, i.e., it is ecorti-
most species of Bucidina. Basically, it consists of two cate and instead comprised of a layer of periclinally ar-
layers (Fig. 2C) similar to the upper cortex of Toninia as ranged, living hyphae stainable in LCB.
described by Timdal (1991). Accordingly, I am retain- Whereas an epinecral layer is lacking in some spe-
ing his terminology: the lower Iayer is comprised of liv- cies, the stainable layer is lacking in some forms of a
ing hyphae that are anticlinally, irregularly, or rarely (in few other species, namely B. schweinitzii, B. sufisa,
some granular forms of Bacidia subincompta and in Ba- and some non-granular specimens of B, subincompta.
cidina ramea) more or less periclinally arranged, the Iu- The epinecral Iayer is well-developed in these species,
mina of which are easily discernable and deeply staina- however.
ble with lactic cotton blue (LCB). This is termed the The distinct stratification present in many species is
stainable layer by Timdal. The upper layer, termed the disrupted by the presence of goniocysts in several spe-
eplnecml layer, consists main1y of periclinall y ar- cies of Bacidina. Goniocysts are discussed below under
mnged, mostly dead hyphae with lumina that are diffi- Vegetative dispersal.
cult to discern and not stainable in ECB (apart from a
few living cell lumina). In all species of Bacidia, the
epinecral layer contains minute crystals (visible at least Photobiont layer
between two polarizing filters) that are soluble in KOH
but insoluble in HNO, and H,SO, (Fig, ZA), These crys- The photobiont of all species is green, unicellular, up to
tals may be absent from some poorly developed individ- c. 15 pm in diameter, and almost certainly belongs in
uals of some species, e.g., B, vermifera. Conversely, the the ChIorococcales. In Bacidina arnoddiana, the photo-
crystals may be very abundant not only in the epinecral biont has been shown to belong to Chlorella saccha-
layer but also in the stainable layer in a few species, rophila (Kriiger) Mig. var. ellipsoideca (Gerneck) Fott &
e.g., B. sufusa. In Bacidina, on the other hand, crystals Novikovi (Tschemak-Woess 1988a, as Woessiafusari-
are never present in the upper cortex (Fig. 2B, C). Dead oides). Tschermak-Woess (1988b) lists no additional
photobiont cells can be found in both of the cortex lay- determination (to genus or species) of the photobiont of
ers in all species. any species presently referred to Bacidia S. str or Ba-
A lower surface which is not attached to the substra- cidina. It has been outside the scope of this'study to
tum and different in anatomy from the upper, corticate identify the photobiont.
one is found in two species of Bucidicd with granular The photobiont layer is commonly continuous and
thallus, B. rubellcl and B. d~ffractu.The lower surface generally does not exceed 50 pm in thickness. In some
consists of a few layers of periclinally arranged, living specimens of Bacidia heterochroa and in several spe-
hyphae that are deeply stainable in LCB. In both the cies of Bacidina with goiliocysts, I have observed a dis-
species mentioned, an upper cortex of normal composi- continuous photobiont layer with colonies of algae sep-
ton is present. arated by more or less wide strands of loosely
Although most species of Bacidia and Bacidina con- interwoven fungal hyphae.
form to the pattern described above, there are several in-
teresting variations and deviations. The most important
ones concern a group of species of Bacidia with granu- Medulla
lar thdlus, namely B. auerswaldii, B. biatorina, and
some forms of B. subincompta among the North Ameri- A medulla is not developed in most species of Buciclia.

Opcm Bol. 127. 1996

Only in species with a thick thallus a medulla is some- finely granular, pale (sometimes yellowish) green mass-
times present, e.g., in Bucidia schweiaitzii and B. suJu- es on the surface of the thallus. Specialized gonio-
sa. A medulla in the strict sense, i.e., a fungal layer be- cystangia, as described by Skrusiaux (1985) for Bauidi-
tween the algal layer and the surface of the substratum, na mirabilis (VEzda) VEzda and four species of
is lacking in all species of Bacidina (but see below un- foliicolous Opegrapha,are not present in any Bacidina
der Vegetative dispersal). treated here. Goniocystate parts of a thallus generally
acquire a completely disrupted algal layer (Fig. 2B). The
algae may become more or less "clumped", i.e., form
Attachment colonies within the fungal tissue, or they may form a
very intempted layer. In any case, there is a marked
The thallus is attached to the substratum by sparse hy- concentration of algae near the surface of the thallus,
phae extending from the lower surface. The hyphae pen- and a considerable amount of the thallus forms a kind of
etrate into the surface of the phorophyte, usually only to medulla. The mitotic activity of both the alga and the
a depth of about 10-100 pm. In one extreme case, I have fungus is obviously very high near the thallus surface.
observed hyphae of Bacidia rubella penetrating c. 400 Goniocysts are formed by the enwrapment of the algal
p into the substrate. cells by the fungus. In Bn. chloroticula, I have observed
that the goniocysts are originally formed from one sin-
gle algal cell, but I have not established whether or not
Vegetative dispersal this holds true for the other eight species with gonio-
cysts. Once formed, the goniocysts may bud off more or
Isidia are absent from all studied species of Bucidia and less immediately, or they may remain to form more or
Bacidina, and soredia are absent from Bacidia. The thal- less coherent aggregations of isidia-like or coraIliile
lus structure of several species of Bacidina is complicat- structure.
ed. In nine out of twelve Noflh American species, the There is no modern, satisfactory definition of a goni-
thallus is sometimes at least in part composed of loosely ocyst. The use of the term goniocyst, the anatomy of
to firmly aggregated granules (Fig. 2B). When firmly various structures named goniocysts, and the possible
aggregated, the granules inay form isidia-like or coral- distinction between a goniocyst and a soredium was dis-
line structures. The thallus granules found in Bacidina cussed in detail by Serusiaux (1985). Although not ex-
have been referred to as goniocysts by Vizda (1980, plicit, it is clear from the context of his paper that SCru-
1990), Skrusiaux (1985), and Lucking (1992). See Tab. siaux favours the definition of a goniocyst used by
1 for an account of the occurrence of goniocysts in the VCzda (1980) with some changes and amendments: the
species of Bacidina treated here. formation of a goniocyst is initiated by the capture of
The function of the so-called goniocysts appears to one single algal cell by the fungal hyphae. The hyphae
vary between and eve11 within the species. In Bn. arnol- quickly enclose the algal cell, all future divisions of
diana, Bn. egenula, and Bn. egenuloidea, the goniocysts which must take pIace within the structure. In compar-
constantly form the main part of the thallus. Even so, ing the goniocystate Bucidiaa mirabilis (VCzda) Vtzda
one cannot rule out the possibility that the goniocysts with available data on sorediate species of Parmeliace-
may function as vegetative dispersal units as well. In ae, he remarks that "the algal divisions are responsible
other species, the thallus is not entirely composed of go- for the formation of soredia", whereas the formation of
niocysts. Instead, the goniocysts are more or less loosely the goniocyst is an initiative of the fungus. However, as
aggregated in distinct or diffuse spots on the thallus. In also pointed out by Sirusiaux, the question of how to
these cases, it is obvious that the goniocysts function as distinguish between a soredium and a goniocyst anatom-
vegetative dispersal units. Bn. varia (Fig. 2B) and Bn. sp ically awaits a final solution.
#l often form large, goniocystate thalli without apothe- In my opiilion, the anatomical/ontogenetical defini-
cia, which implies that in such cases the goniocysts are tion of a goniogyst has some shortcomings. Firstly, one
the main method of dispersal. Furthermore, as in many has to decide whether a goniocyst and a soredium
species with both soralia and apothecia, the number of should be defined by function or by anatomylontogeny.
apothecia in a specimen is more or less inversely pro- I believe it would be a logical flaw assuming that there
portional to the quantity of goniocysts. This indicates must necessarily be a difference in anatomy between a
that the lichen, due to environmental influence andlor soredium and a goniocyst. There are no data showing
genetic predisposal, can use either ascospores or gonio- that soredia consistently arise from more than one algal
cystslsoredia as the principal mean of dispersal. cell, which would be the logical assumption if gonio-
The presence of goniocysts strongIy affects the gener- cysts are to be distinguishable on the basis of this char-
al appearance as well as the anatomy of the thallus. In acter. And why draw the line between the formation
most species of Bacidina, the goniocysts usuaIly form from one and two algal cells? I fail to see the principal
difference; such a line appears to me as arbitrary. Sore- gin is distinct, level with the disc or, more commonly,
dia and goniocysts obviously do not differ in being en- slightly raised. The disc becomes increasingly convex
closed by fungal hyphae. When it comes to the mycobi- with age in many species, and simultaneously, the mar-
ont or the photobiont taking the initiative in the gin becomes increasingly suppressed, finally to be en-
formation of the structure, I doubt that it is possible to tirely excluded. In a number of species, however, the
draw such a clear line, considering that we are dealing disc often or always remains (almost) flat and the mar-
with symbiotic organisms, in which many processes gin distinct: Bacidia absistens (often), B. auerswaldii,
must be coordinated. One should also bear in mind that B, biatorina, B. campalea, B. circumspecta (often), B.
soredia and, most likely, also goniocysts have evolved helicospora, B. heterochroa, B. igniarii, B. insularis, B.
several times. Consequently, there is little reason to be- rnnitabilis, B. rmgens (often), B. russeola, B. vermif.em,
lieve that soredia or goniocysts of widely differing Bacidina chloroticula, and Bn. egenuloidea.
groups are similar in all traits, or not even horno~ogous. A white pruina occurs occasionaIly or invariably in
The functional view of a goniocyst advocated by eight species of Bacidia, namely B. aggregutula (some-
Hawksworth et al. (1 983) and Coppins (1 983) appears to times), B. d1J5racta (always), B, insularis (sometimes),
me as more attractive. In this definition, both goniocysts B, mutubilis (always?), B. polychroa (sometimes), B.
and soredia consist of clusters of algal cells (the number rubella (sometimes), B. schweinitzii (rarely), and B. s y p
of which they originate from being irrelevant) surround- fusa (always). The pruina may vary from thin to thick,
ed by fungal hyphae. The difference lies in the function: and it is variable within the same species whether the
Goniocysts are the main component of the thallus, and pruina occurs on the edge of the apothecium, the disc,
do not normally serve in tlie propagation of the Iichen. or both. It is not uncommon that only young apothecia
Soredia, on the other hand, primarily function as vegeta- are pruinose. In Bacidina, none of the species is pru-
tive dispersal units. A consequence of this definition is inose. I define a pruina as a layer of crystals external to
that the two concepts become overlapping, which in my the tissues. It should not be conhsed with an internal
opinion is logical. The hnction of a "thallus granule" crystal layer, which is present between the paraphyses
can, in some species, be to serve in the propagation as or in the proper exciple of several species (see below).
well as to be the component of which the thallus is built. I have not made detailed ontogenetic studies of the
If the latter definition is used, then several of the North apothecia. The essential features of the apothecial on-
American species of Bacidina may be termed sorediate. togeny of Bacidia and Bacidina, however, appear to
Nevertheless, as a provisional measure, I have cho- agree with the ones described for Lecidella elaeochro-
sen to retain the term goniocyst for the "thallus gran- ma (Ach.) M. Choisy by Letrouit-Galinou (1967, 1968).
ules" formed by the Bacidina, as long as there is no This type of apothecial ontogeny was later referred to as
general agreement on the definition of the terms in- a variant of the Xunthoria-type (Letrouit-Galinou &
volved, Bellemere 1989),
In the granular species of Bacidia discussed previous-
ly, the granules are homologous with nonnal areoles, Proper exciple
and should not be confused with the goniocysts of Ba-
cidina. Anyway, it is likely that the granules of the Ba- The proper exciple of Bacidia and Bacidina is well-de-
cidia species can function as vegetative propagules. veloped, annular, and composed of distinctly or indis-
tinctly radiating hyphae (Fig. 3B-C, 4A-B). The anato-
my of the proper exciple provides some of the best
Apothecia characters to separate these two genera, and alsp to sep-
arate both of them from other genera.
Morphology and ontogcny The excipular hyphae of Bacidia are, at least in the
outer part, distinctly radiating. In some species, the in-
The apothecia are biatorine or lecideine in all North nermost part of the proper exciple forms a more or less
American species of Bacidia and Bacidina except Bn. distinct layer with an intricate texture. The presence or
ramea. In Bn. ramea, the apothecia are zeorine, possess- absence of such a layer varies between closely allied
ing a proper exciple that is as well-developed as in other taxa. For example, such a layer is present in Bacidira ig-
species of Bacidina, as well as a thin thalline margin that niarii, but absent from the closely related B. circum-
soon becomes excluded (Fig. 43E). The thalline margin, specta.
however, is usually easily observed in sections, even in The hyphae of the radiating part of the proper exciple
old apothecia. are abundantly furcate but sparingly or not at all anasto-
The apothecia of Bacidia and Bacidina are usually mosed (Fig. 3B-C).The cell lumina in the interior of the
sessile on the thdlus or slightly constricted below. When proper exciple are usually more or less narrowly cylin-
young, the disc is plane or slightly concave, and the mar- drical and distinctly constricted near the septa (Fig. 3D),
Fig. 3. Sections of apothecia in Bacidia, unstained (A) or stained with LCB (B-D), viewed between two polarizing filters (A) or in
interference contrast (B-D). - A. Bncidia campalea (Ekman L1 177, LD), crystals in proper exciple. Bar = 100 pm, - B. Bacidia
Iaurocerasi subsp. laerrocerasi ( E h a n L1070, LD), very narrow excipular cell lumina. Bar = 20 pm. - C. Bacidia su&a
(Wetmore 60307, MW), layer of enlarged cell lumina along rim of proper exciple. Bar = 30 pmun. - D. Bacidin rubella (Ekman
L950, ED),detail of lateral part of proper exciple showing constrictions o f cell lumina at septa. Bar = 10 p.

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-

Opera Bot. 127. 1996

cies, Bn. u q ~ s t u l l ~ f ~the
i ~ uproper
, exciple of which al- species. 1t is sometimes a good character for separatiilg
ways contains evenly dispersed crystals up to l pm species. 1n many species, the majority of the apices are
wide. more or less swollen, often taking a more or less clavate
The proper exciple of most Bcrcidiu and several Ru- shape. In Bacidia car?tpalra, B. helicuspora, B. hos-
cidina is more or less pigmented; see Pigntentarion.. iheleoides, B. medialis, B, muiahilis, B. russeola, B. suJl
,jii.~cr,and Bacidina aenea, on the other hand, the para-
physes are consistently uniformly thick throughout, or
W ypothecium only sligl~tlythickened or even slightly tapering at the
apices.
A distinct hypothecium of diffcrcnt tcxture than the The upper part of the paraphyses oftcn sccrctcs pig-
proper exciple is prescnt In all stud~cdspcclcs of Bacid- ment into the surroundiilg gelatinous matrix. The pig-
ia and Bacidina {Fig. 3A, C ) . The hyphae arc irregularly ment is either dissolved in the gelatinous mallix, form-
oriented and tl~elrcell lurnilla are veiy varlable in s u e ing a distinct external hood on the apices. or, when in
and shape. The hypotheciu~n is of'len pig~nented (see high concentration, it may form fairly large, irregular
Pignlentutiun). Crystals and or1 droplets are never clu~npsthat are not dissolved in the matrix. In a few
present in the hppolheciuin. spccics, namely Bacidia auer.qwaldii, B heterochron, R.
ve~~inijera, aid occasional forms of B. lcduroceru,~iand
B. subinconzpta, the pigmcnt rcmains within the wall of
the apices of the paraphyscs. Often, the pigmented part
of the wall is distinctly delimited from the unpigmented
A very thin subhymenium is always present betwell the part below, thus foin~ingdistinct intcrnal "caps" of pig-
hymenium and the hypothecium. It is always amyloid ment.
(contrary to the hypothcciuin) and often intermediate in
pigmentation bctwccn thc hypothccium and thc hy- Asci
mcniuin.
The asci are clavate and firmly attached to the ascoge-
nuus hyphae, from which they are delimited by an ob-
Iiquc scptum. The upper part of the asci IS surrounded
by a hzzy, more or less amyloid gelatinous enveIope.
The hynlciliunl is an~yloid,40-130 p n high ~ in Bacidia Likc the ascus itself, the gelatinous envelope is unpig-
and 40-80 p n high in Bacidina. In all species of Buci- incntcd in all species except Bacidia hclicospom. rn this
dia and most of Buridinu, the paraphyses coilstitutc thc species, the ascus wall secretes a brown pigment (Lau-
major part of the tissues, but in lwo species of Bacidia, roccrasi Brown; see beloiv) into the gelatinous enve-
namely Bn, chlomticul(i and Bn. vuriu, the asci are do111- lope. This process starts early, already when the asci are
inating. The liylneniuli~is offen pig~i~ented (see Pign~m- immature and much smaller than thc fully grown ones.
ration), either in a distinct layer in the upper part nf the As in all lichens, the wall of the ascus in Bacidia and
lymeniu~nor in a diffuse zoi~ewith upwards and gradu- Bacidina consists of four layers, the a-, b-, c-, and d-lay-
ally increasing colour illtcnsity. In thc dcscriptions, I ers (I3ellemkre & Letrouit-Galinou 1988: 167: see also
have avoidcd thc term "cpithcciuii~" for the pigmcnted Fig. 15B). Thc a- and b-layers together form the ex-
zone. By definition: this term refers to the layer above oascus. Thcy arc impossible to separate in an ordinary
the asci (cf. Hawksworth et al. 1983), but in n~ailyspc- light inicroscopc, and they are henceforth collec!ively
cies of Bacidici and Baciclin~ithe pig~nentationis not re- tcnncd the a+b-layer. Inwards, they are succeeded by
stricted to this layer. the c-layer, which forms the external part of the en-
A layer of ~ninutecrystals between the apices o r the doascus, and, finally, the d-layer, which constitutes the
paraphyses is present in some species, namely Rucidiu internal part of the enduascus. Thc d-layer is distinctly
uggrcgutulu, B. rusellu, and Ruoiciinrr CF-ystcrllifi,ra.Oil thickened in the apical rcgion, where it forms the major
droplets, however, are never present in any species. part of the apical apparatus. Thc a+b-layer is distinctly
amyloid in all species of Bacidia and Bacidina, al-
though to a varying degree. Commonly, it is more
Paraphyses strongly amyloid in the upper part of the ascus than in
thc lower. The c-layer is only faintly amyloid. The d-
Thc paraphyses are unbranched or branched in the up- layer is basically strongly amyloid, but in Bacidiu and
per part. l11 Bacidia, thcy arc 0.5-2.3 pn wide and in Buuidinu, like in inany other lichens, it contains a less
Bacidina 0.8-2.0 p n wide in mid-hylnenium. Thc width amyloid axial body.
of the apiccs varics considerably bctwccn and within Within Briuidia, thc apical region of the ascus is fair-
Open Bot. 127, 1996
Tab. 1. Goniocysts in North American corticolous and lignicolous Bacidina. Statements refer to North American material only
(but see notes). Notes: - 1. This statement should be interpreted with caution, since only 1-2 North American specimens have been
investigated. - 2. Far from a11 European specimens have goniocysts. - 3. In Europe, many (but far from all) specimens are entirely
dissolved into goniocysts.
Goniocysts All specimens Distribution of Aggregation of Average diarn. o f
present with goniocysts goniocysts goniocysts goniocysts (p)

Bn,uenea yes' small spots loose

Bn. a~nolcliana Ye6 almost entire thallus firm
Bn. assuluta yes1.2 rather small spots loose
Bn, cal fornica no small to rather large spots loose
Bn. chloroticula no rather large spots3 loose to rather firm
Bn. c~stallifera
Bn. egenlrla yes1 entire thaIlus firm
Bn. egenuloidea yes1 entire thallus firm
Bn. rurnea
Bn. squamellosa
Ba. varh no small spots to entire thallus rather loose
Bn. so # l yes1 rather small spots loose

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.

Opera Bot. 127. l996

 ponent in taxonomic revisions of lichens is still fairly
new, covering only about a decade. Many genera are
stiIl insufficiently studied in this respect. It was with the
work of Hafellner (1984) that the importance of this
characeter set in lichen taxonomy was first clearly
pointed out. In my opinion, this was the greatest
achievement of this work, whereas the materiaI studied
was far too small to support many of the rather far-
reaching taxonomical concIusions.
As often is the case when a new character is intro-
duced in taxonomic work, it is given an a priori weight
to the taxonomy, which is hence strongly influenced by
this single character. This is exactly what has happened
in lichen taxonomy over the last decade. Furthermore,
the study of the ascus apex has not seldom been restrict-
ed to generic type species or "representative species",
methods based on the, in my opinion, dangerous as-
Fig. 6. Spore types. Bar = 25 pm. - A. Sigmoid, acicular spore sumption that the ascus apex is invariable within genera
in Bacidia laurocerusi subsp. laumcewi. - B. Straight, acic- or at least species groups within genera. It is commonly
ular spore in Bacidia heteruchroa. - C. BaciIliform spore in seen in the literature, that asci are referred to "ascus
Bacidia circumspectu. - D. Clavate (intermediate between ba- types" (e.g., Bacidia-type, Biatora-type, or Lecanom-
cilliform and acicular) spore in Bacidla cipcurnspecta. - E. type). There is an apparent risk that important taxonom-
Fusiform spore with blunt ends in Blacidia ouerswaldii. -
Drawings by UIf Amp. ic information is lost when an observed ascus is
squeezed into any of a Iimited number of "ascus types"
that, moreover, have never been properly defined.
line of argument). Tnus, it is (or has been) exposed to a In my opinion, the ascus apex characters should be
strong selective pressure. treated like any other character. Its variation within spe-
The most likely reason for the ascus apex to loose its cies, genera, and families should be studied together
importance in the dispersal would be that vegetative dis- with all other characters used. It is then the pattern of
persal has taken over as the mean method of propaga- covariation in the characters studied that, a posteriori,
tion. At first, this may seem as an attractive hypothesis determines the value of the ascus apex, or any other
for what has taken place in Bucidina, since several spe- character for that matter. This reasoning may seem self-
cies, contrary to Bacrdia, produce goniocysts. However, evident, but there are, unfortunately, many examples of
if one makes the plausible assumption that an ascus how ascus apex characters have not been treated this
apex similar to that in Bacidia is the original, plesio- way. In Bacidia and Bacidina, the ascus apex, even
morphic character state in Bacidina, then four specles though variable within the latter genus, has proven to be
stand out as particularly derived from this state, namely of utmost importance to circumscribe the genera, but
Bn cqJstallfera, Bn. egenuluidea, Bn. rarnea, and Bn. aIso to characterize some of the species of Bacidina.
squarnellosa. This group embraces all three North
American species of Bacrdzna that never produce gonio-
cysts (Tab. l), i.e., the specles that are entrrely depend- Spores
ent on spore dispersal for their propagation! It should be
noted, that the probability of randomly picking four out The spores of all Bacidia and Bacidina treated here are
of twelve species that includes all three species depend- long and narrow, colourless, rather thick-walled, and
ent on spore dispersal is 1.8 %. Consequently, the sec- lack perispore and ornamentation. In Bacidia, they are
ond hypothesis appears less contradictory. The three 11-108 pm long, 1.2-7.8 pm wide, and 3-66 times as
North American species are not the only ones not pro- long as wide, whereas they are 16-60 pm long, 0.9-3.3
ducing vegetative propagules. Among the European pm wide, and 9-56 times as long as wide in Bacidina.
species, at least Baczdrna phacodes (the type species of Although often highly variable within a species or even
the genus) and Bn. inund~ranever reproduce vegeta- a single specimen, the spore size is an important charac-
nvely. The reason why Bn. egenuluzdea possesses a ter to distinguish between some species (Fig. 6, Tab. 2).
highly derived ascus apex even though it reproduces The spores are usualIy eight per ascus, or sometimes
vegetatively may be that the ascus apex originally reduced to seven or six. They may or may not be more
volved in a non-goniocystate ancestor of this species. or less coiled in the ascus. This is, however, variable
The use of the ascus apex structure as a regular corn- within the same species, not rarely even within the same
' ?

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.

Aciculnr Clavatc Bacilliform Pusiform Lcngth Ratio Septa

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

specimen. (referred to here as "clavate''). B, verrgens, B, subin-

Thc sporcs arc acicular in nlaily species, i.e., they conzp[c/, and Bn. aenea oftcn producc acicular spores,
posscss o ~ l enarrow, tapering, more or less acute end, bu( occasionally also long-bacillifoin~oncs. I11 B, a~rer-
one (hick and blunt end, and a length/width ratio ex- swaldiii, the spores are invariably fusifom with blunt
ceeding about 10. In a few species, nameIy, Bacidiu civ- ends, in B. hustht~leoides,B, n~edialis,B. ins~rlaris,and
cntrilspcrtu, B. hu.sthcleoides, B. i g ~ ~ i u rB.
l i , t n ~ d i u l i s ,B. B. snrRincumpfu occasiollally so. The namow, tapering
verntiferu, and Bacidina vnr.ia, the spores arc oftcn ci- end of an acicular or clavate spore IS always directed to-
ther bacilliforrn (i.e., evenly wide with blunt ends) or in- wards the proximal end of the ascus, and the wider,
termediate in shape between acicular and bacilliform blunt end towards the distal end. The various spore
 maximurn number of septa is commonly, depending on
Fig. 7. Bacidia helicmpora (Chester 4413, US), squash the species, between seven and fifteen. In Bacidia lau-
preparation in KOH. Released package of eight helically
arranged and strongly congIutinated spores. rocerasi subsp. idahoensis, however, I have encoun-
tered spores with up to 28 septa. In some species, on the
other hand, a majority of the spores never acquire more
shapes present in the species treated here are presented than thee septa (Bacidia cirmmspecta, B. hosthele-
in Fig. 6 and Tab. 2. oida, B. igniurii, B, medialis, B. vermifeu, and Bacidi-
The spores are straight, curved, or more or less "sig- nu varia). In a few species of Bucidi~a(Bn.arnoldiana,
moid". The latter term has commonIy been used to de- Ba. chbroticula, Bn. egenula, and Bn. egenuloidea),
scribe the spores in Bacidiu, but it is actuaIly a rnisno- mature spores with fewer than three septa are often
mm. It implies that the spores are S-shaped in one found.

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

Opera Bot. 127. 1996

Tab. 4. Conidial types in the North American corticolous and lignicolous species of Bacidia and Bacidinu. These are defined in
the text, and are only briefly characterized here: 1 = filiform (a = curved; b = straight; c = multiseptate contrary to a, b, and d; d =
strongly conglutinated, bundle-forming contrary to a-c); 2 = oblong, ellipsoid, fusiform, or bacilliform; 3 = falcate; 4 = acicular.
An "S" denotes a conidial type that occurs singly in a pycnidium, whereas an "m" denotes a conidial type that occur mixed with a
second conidial type in the snme pycnidium. Information within brackets refer to information based on European collections (SW.
= Sweden, Ge. = Germany). The number of conidial types in a species and the number of species with a particular conidial type
refers to North American material, but within brackets the total number (including the information based on European collections)
is given.

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

No. of species 23 (24) 2 4 1 4 (5) 1

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-

Opera Bot. 127, 1996

 or, rarely, in the prothallus. Only in Bacidia circum-
specta and B. igniarii are they sometimes sessiIe. They
are generally 40-175 pm wide in Bacidiu and 60-200
p n in Bacidina, but in Bucidia circumspecta they are
sometimes up to 300 pm wide. In most cases, they are
very similar in pigmentation to the apothecia, except in
some species of Bacidina, which have unpigmented
pycnidia even though the apothecia are pigmented. The
pycnidia are unilocular and globose, ampullifom, or
slightly flattened in shape.

Conidiogenous cells

The inside of the pycnidial cavity is lined by

conidiogenous cells that produce conidia terminally and
lack distinct conidiophores (or the conidiophores can be
said to form the pycnidial wall). The conidiogenous
cells are 7-15 X 1.0-2.5 pm, cylindrical or slightly
tapering in Bacidia, 4-8 X 1 .O- 1.S pm and cylindrical in
Bacidina.
Fig. 8. Conidial types. Bar = 10 pm. - A. Type l a in BaczldiQ
sch~veinitzii.- B.Type I b in Bacidia medialis. - C . Type Ic
in Bacidina varia. - D. Type I d in Bacidina egenuloidea. - E.
Type 2 in Bacidia circumspecta. - F, Type 3 in Bacidia Conidia
ci~cuwpecta.- G.Type 4 in Bacidina egenuloideu. - Draw-
ings by Ulf A m p . Within the North American Bacidia and Bacidina, I
have encountered altogether seven types of conidia, as
tion cannot be entirely perfect, however, otherwise classified by size, shape, curvature, septation, and
even-septate spores would be virtually non-existent. conglutination (Fig. 8). Among the 29 species found
All spore measurements in Bacidia and Bacidim should with pycnidia in North American material, 24 possess
be performed on mature spores that have been released one single conidial type. Three species, Bauidia
from the asci. In practice, the absolute length of the hostheleoides, B. vertnfem, and Bacidina californica
spores is iinpossible to measure, and c~nsequently,I possess two different conidial types. In B. hosthele-
have chosen here to define the length of a spore as the oides, both conidial types occur mixed in the same pyc-
distance between the two ends, irrespective of the cur- nidium. Two species, Bacidia circumspecta and Bacidi-
vature. This means that the absolute length is slightly nu egenuloidea, possess as many as three different
underestimated in most species, since the spores are of- conidial types. In the latter species, two of the three CO-
ten slightly curved or sigmoid. Only in Bacidiu heli- nidial types occur mixed in the same pycnidia. Few oth-
cospom and B. vermifera, both with strongly sigmoid er lichens are known to produce pycnidia containing a
spores, is the underestimation severe. Thc width of the mixture of two conidial types. To my knowledge, they
spores should be measured at the widest part of the have so far been reported from Lecania cyrtellina (NyI.)
spores, i.e., in acicular spores, near the wide, blunt end. Sandst. and L. cyp-tells (Ach.) Th. Fr. (Ekman 1994, but
by then already known from the former species by Brian
Pycnidia Coppins), and from Lecanom schistina (Nyl.) Amold
(Etayo & Diederich 1993) and Opegraphu vuiguta
Pycnidia have been found in North American specimens (Ach.) Ach. (Coppins pers. comrn.)
of 19 out of the 27 species of Bacidia and 10 out of the The folIowing conidial types have been found in the
12 species of Bacidina. In most species, they are small North American Bacidia and Bacidina. The presence of
and inconspicuous, and only present in some specimens the various types in each of the species is presented in
and in in a low number. Thus, they are often difficult to Tab. 4. All conidial types are borne singly on the conid-
find, and they may have been overlooked in some of the iogenous cells, except type 2, which is formed repeated-
10 species in which they have not yet been found. In ly.
some species of Bacidina, on the other hand, they are 1. Filiform, more or less curved or not, 0-1 l-septate,
fairly abundant and thus easy to find. conglutinated or not. The filifom conidia divides into
The pycnidia are more or less immersed in the thallus four types:

Opcra B o t 127. 1996

Tab. 5. Pigments present in the apothecia of the North American corticolous and lignico1ous species of Bacidia and Bacidina. The
pigments arc described in the text. A "+" refers to a pigment that is always or nearly always present, whereas a "+" denotes a
pigment that may or may not be present (including rarely occurring pigments). Unidentified pigments are pigments that occur in so
small amounts as to be impossible to identify with any of the named pigments. It is likely that the unidentified pigments are
synonymous with Rubella Orange.

Bacidia Laurocerasi Rubella Schweinitzii Arceutina Bacidia Thalloidima Arnoldiana unid.

Green Brown Orange Red Yellow Brown Green Brown pigments
Bacidia
absistens
aggrega ruln
arceulina
auerswaldii
bidorina
campaleu
circumspecla
dzficta
friesiana
helicospora
heterochroa
hostheleoides
igniarii
insularis
laurocemsi
medialis
mutabilis
polychroa
reagens
roselIa
rubelIa
russeola
solmonea
schweinitzii
subincompta
stiflusa
vermvera

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

Opera Bot. 127. 19%

Tab. 6. A summary of the pigments found in North American Bncidia and Bacidina: their colours and reactions with K, C, and N.
A minus sign denotes the absence of a reaction.

Reaction with
Pigment Colour in pure water K C N

Bacidia Green I Green to blue-green Purple

Bacidia Green I1 Purplish (dirty) Green Purple
Laurocerasi Brown Reddish brown (grey, grey-brown) Purplish Orange to red
Rubella Orange Orange to orange-brown (yellow) f intensifying f intensifying
Schweinitzii Red Dark red to red-black - (intensifying?) - (intensifying?)
Arceutina Yellow Yellow to yellow brown
Bacidia Brown Brown to red-brown Purple-red
lialloidima Green Green-grey to dirty grey Purple to violet Violet Red
Arnoldiana Brown Brown

than the distal end), 0-7-septate, 16-45 x 1.0-1.6 pm Pigmentation

(Fig. 8G).
The length of the conidia varies considerabIy within a The acetone-insoluble pigments found in the apothecia
pycnidium, between pycnidia on the same thallus, and provide the single most important character set to distin-
between specimens. In some species, the length has guish between species within both Bacidiu and Bacidi-
been found to be rather invariable within a pycnidium aa. They should be studied in transmitted light on mi-
but highly variable between specimens (difference in crotome sections or thin handcut sections mounted in
length up to about 100 %). Finding pycnidia is often a water. The colour caused by these pigment, as seen in
difficult and time-consuming task, and consequently, transmitted light, is determined by their concentration,
only a few pycnidia have been studied in most species. distribution, and by the texture of the tissue in which
The length values of the conidia accounted for in the they are located. Furthermore, mixtures of two or three
descriptions should thus be interpreted conservatively. pigments in the same tissue are fairly common. In re-
The width of the conidia, on the other hand, appears to flected light, as for example when studying the surface
be much less variable. of an apothecium in a dissecting microscope, the colour
The conidia provide important characters to separate of the pigments is often perceived in a different way.
Bacidia and Bacidina from other, related genera. All For examgle, both brown and green pigments may give
species so far found with pycnidia (with the exception a completely black colour to the apothecia in reflected
of Baddia igniarii, which is anyway obviously closely light, and yellow, orange, and brown pigments may give
related to B, circumspecta) at least sometimes produce very similar brown hues. Thus, it is important to study
filifonn conidia (type l). The distribution of the conidi- the pigments in transmitted light and not in reflected
a1 types within the two genera indicates that closely re- light.
lated species are likely to produce similar conidia. For There may be much variation within a species in the
example, Bacidina egenula and Bn. egenuloidea both concentration of various pigments and to some extent
produce the peculiar type 4 conidia. Bn, varia and Bn. also their distribution within the apothecium. The types
sp. #l both produce type l c conidia, and Bucidiu cir- of pigments present, however, are relatively invariable.
cumspect~,B. igniarii, and B. vermdfera all produce Most species contain one or two pigments, but' as many
type 2 conidia. A few cases, however, indicate that the as three may be present, even in the same apothecium
conidial type produced is perhaps not always such a (but then in different parts of the apothecium). In sever-
conservative character as one might believe. h Blacidia al species, occasional pigment deficient specimens oc-
roseliu, the North American specimens differ from the cur that may be difficult to determine. Other patterns
European ones in the type of conidia produced. There appear to be a normal part of the variation within a spe-
are no additional distinct differences between them, oth- cies. h Bacidia schweinihii, for example, there is al-
er than that the North American specimens appear more most a dimorphism. Most of the variation within the
poorly developed. Furthermore, the spectacular .varia- species can be classified into one morph dominated by a
tion in the conidia exhibited by Bacidiu circumspecta green pigment (with bluish or black apothecia) and one
appears to Iack any correlation with other characters or morph with a brown-orange pigment (with orange to
with geographic origin. brown apothecia). There are intermediates, however,
and a few specimens possess apothecia of both colour
morphs. The species of Bacidina are, on average, less

Opera Bot. 127, 1W6

pigmented than Bacidia. Among the species treated varies even between sections from the same apothecium
here, Bn, cal$ornica, Bn. chloroticzdla, Bn. cystulliferu, studied in one and the same slide. Blue crystals appear
BR. rumea, Bn. sguameUosa, Bn. varia, and Bn, sp. #l to be more abundantly formed in sections that are thick
are commonly pigment deficient. Some of these species andlor have a high concentration of pigment. On the
(Bn,cal~ornica,Bn. ramea, and Bn, vuria), however, other hand, I have encountered specimens with very
display an extreme variation in the amount of pigment high amounts of green pigment in the hymenium that do
present, and they include forms with even dark brown not produce a single blue crystal in nitric acid. This im-
apothecia! It should be noted that entirely pigment defi- plies that there is some other factor than amount and
cient apothecia are not pure white, but usually pale pink concentration of pigment that controls the precipitation
or sometimes pale yellow or pale greyish. Obviously, of blue crystals. Perhaps, Bacidia Green consists of a
the texture of the tissues in themselves contribute slight- mixture of similar substances occurring in different pro-
ly to the colour of the apothecia. portions in different specimens. It may also be that the
An attempt is made here to classify the pigments ability to precipitate blue crystals is dependent on the
found in the species studied. 1 am aware that such a texture of the tissues or the presence of quite different
classification is a rather blunt instrument. It is possible, substances. that can function as condensation nuclei for
in some cases even likely, that what is classified here as the crystals.
one single pigment may in reality be comprised of two In most species, Bacidia Green takes the normal,
or more chemica1 substances. If chemical analyses of green hue without any reaction with K (Bacidia Green 1
the pigments found in the apothecia of Bacidia and Ba- in Tab. 5). In B, absistens, on the other hand, many Eu-
cidina would be possible, the conditions for taxonomic ropean and a few North American specimens have a
work in these genera would be greatly enhanced. For the purplish upper part of hymenium, which reacts K+
sake of making the descriptions and discussions sim- green (Bacidia Green I1 in Tab. 5). The reactions of Ba-
pler, the pigments discussed below have been named. cidia Green are pH-dependent and reversible, and the
Some of them already have more or less established purplish colour of B. absistens shifts towards green al-
names originally provided by Bachman (1890). I have ready in very diluted KOH solutions. It is possible that
retained these names (although in an anglicized ver- the colour of the hymenium in B, absistens is affected
sion), and given new, provisional names to others. The by the pH of the substratum and the water trickling over
presence of the pigments in the apothecia in each of the the specimen. There are, however, a few Swedish and
species treated is accounted for in Tab. 5. The colours Norwegian specimens that possess a green hymenium
a i d reactions with K, C, and N have been summarized even though they are from acid habitats, which indicates
in Tab. 6. It should be noted that most, if not all, pig- that the lichen, to some extent, has the ability to regulate
ments occur in other lichen genera than Bacidia and Bu- the pH in its tissues.
cidina, "Intensifying" reactions with K and N men- (2) Lanrocerasi Brown. Usually more or less brown
tioned below refer to reactions where the shade of (often reddish), in low concentration grey-brown or
colour remains unchanged but the colour intensity in- even almost grey. Reacts K+ purplish, C-, and W-l- or-
creases. A case could be made for regarding such reac- ange to red. Usually found in the proper exciple and in
tions as negative. the upper part of the hymenium, sometimes in the hy-
( l ) Bacidia Green (including both the "Bacidiagriin" pothecium. Occurs in 16 species of Bacidia and four
and the "Lecideagriin" of Bachmann). Usually green species of Bucidina.
(sometimes dirty) to blue-green, in more or less acidic (3) Rubella Orange. Yellow in low concentrations,
conditions purplish. Reacts K- (or actually a purification orange to orange-brown in higher concentrations. Re-
of the green) or when purplish K+ green, and C-, N+ acts K+ intensifying (in low amounts sometimes K-), C-,
purple, with or without a precipitate of small bIue crys- and N+ intensifying (in low amounts N-). Found in all
tals. Usually found in the proper exciple and in the up- apothecial tissues. Occurs in 20 (21?) species of Bacid-
per part of the hymenium, rarely in other parts of the ia and five species of Bacidina. - This is the common-
apothecia. Occurs in eight species of Bacidia and one est pigment in Bacidia and Bacidina. In Bacidia muta-
species of Bacidina. - I have chosen to treat the "Bacid- bilis, which normally contains Rubella Orange as the
iagriin" and the "Lecideagriin" of Bachmann (1 890) col- single pigment, I have once observed entirely colourless
lectively. According to Bachrnann, "Bacidiagriin" is apothecial sections of a shade form react distinctly K+
supposed to precipitate small blue granules in nitric ac- yellow. This implies that Rubella Orange is actually a
id, whereas "Lecideagriin" is not. I have found much group of substances, at least one of which is colourless
variation between specimens of the same species in before treatment with K.
their tendency to precipitate blue crystals, without see- 4. Schweinitzii Red. Red, but often in such high con-
ing any other differences in colour or reactions with K, centrations as to appear red-black. Reacts K- (or K+ in-
C, or N. Not seldom, the amount of blue crystals formed tensifying?), C-, and N- (or N+ intensifying?). Occurs in

Opcra Bot. 127. 1996

the hypothecium and proper exciple of two species of Chemistry
Bacidia.
5. Arceutino Yellow. Yellow or brown-yellow, in Eleven of the 25 investigated species of Bacidia and
high concentration brown with only a slight yellowish none of the eleven investigated species of Bacidina
hue. Reacts K-, C-, and N-. Occurs in the proper exci- were found to produce secondary metabolites in con-
pIe, hypothecium, and upper part of the hymenium in centrations detectable with BPTLC. Bacidia fiiesiana,
one species of Bucidio and one species of Bacidina. B. igniarii, and Bacidina assulata were not investigated
6 . Bacidia Brown ("Bacidiabraun"). Pure brown to due to sparsity of material. Among the eleven species of
red-brown. Reacts K+ purple-red, C-, and N-. Occurs in Bacidiu found to contain secondary metabolites, seven
the proper exciple, hypothecium, and upper part of the produced atranorin (occasionally or invariably) as the
hymenium in two species of Bacidia. only substance. Zeorin appears to be a regularly occur-
7. Thalloidima Green ("Thalloidimagriin"). Green- ring substance in two species, and a rare substance in
grey to dirty grey. Reacts K+ purple to violet, C+ violet, one additional species. The chemical patterns found in
and N+ red. Occurs in the proper exciple and upper part the species investigated is shown in Fig. 9.
of the hymenium in one species of Bacidia. Bacidia hetemchroa is the only species treated that
8. Amoldiana Brown. Brown. Reacts K+ green- exhibits a geographically correlated chemical pattern
brown, C-, N-. Possibly synonymous with the "Sphaer- within the study area. In this species, all specimens from
omphdebraun" of Bachmann. Found in the proper exci- California and Oregon were void of secondary metabo-
ple, hypothecium, and upper part of the hymenium. lites, whereas the ones from the Southeast were found
Occurs in three species of Bucidina. to contain atranorin. The chemical difference between
When two or three pigments are present in the same these population groups is correlated with a number of
apothecium, their distribution within the apothecium morphological and anatomical characters. This case is
sometimes overlap. Such mixtures make the individual treated in the chapter Numerical and statistical treat-
pigments difficult to identify. Fortunately, the pigments ment and under the species in the chapter Taxonomy.
do not usually overlap completely. Furthermore, the dis- Only one species, Bacidia absistens, was found to
tribution and relative concentration of the pigments of- have a complex secondary chemistry. Tensbeg et al.
ten vary between different apothecia, which also facili- (1995) reported 4-0-methylcryptochlorophaeicacid, a
tates identification. If nothing else helps, the pigments rare mesa-depside in the orcinol series, as the only ma-
may be slowly bleached using a dilute hypochlorite so- jor secondary product in a number of specimens from
lution (c. 0.2 %). The pigments are variously resistant to Norway and Sweden. I investigated three specimens
bleaching, and during the few minutes it takes for all of from California, Washington, and British Columbia.
the pigments to become completely bleached, their pres- They were found to produce four orcinol meta-depsides
ence and distribution can be studied. For example, when in two different combinations. None of these substances
the sometimes almost black proper exciple of Bacidia is identical to 4-0-rnethylcryptochlorophaeicacid, nor
circumspecta is bleached, the brown pigment (Laurocer- is any of them identical to merochlorophaeic acid,
asi Brown) quickly fades, revealing the presence of a sekikaic acid, or homosekikaic acid. The identity of the
green pigment (Bacidia Green), which is much more re- four substances remains unknown, but there is no doubt
sistant to bleaching. that they are orcinol metu-depsides. This group of sub-
As mentioned earlier, the apothecia of pigment defi- stances produces characteristic, narrow spots that inten-
cient specimens are pale pink, pale yellowish, or pale sify to a deep purple hue during the first few days after
grey. After a few years in a herbarium, however, pig- having been exposed to sulphuric acid and heat. Further
ment deficient apothecia start to darken, and old herbar- investigations of the chemistry of Bacidia abshtens are
ium specimens may become nearly medium-orange. On required. More, large collections of Bacidia absistens
the other hand, the few old collections with originally are needed, since very few of the presently available
pale grey apothecia (concerns Bacidina chlorotiezdia) North American ones are large enough to allow chemi-
seen by me appear not to have been affected by aging. cal investigations.
The darkening of pigment deficient apothecia due to ag-
ing is probably caused by an alteration of the tissue Ecology
structure, not by the formation of a pigment.
The pycnidial wall, when pigmented, always possess- Substratum
es the same pigments as the apothecia. Species with
three pigments in the apothecia, however, usually only Since the ecology has been used to circumscribe the
have two of these pigments in the pycnidia, at least as content of this work, all species are, by definition, corti-
far as can be detected. colous or lignicolous. Most spmies are exclusively cor-
ticolous. Bacidia circumspecta, B. laurocerlasi subsp.

Opera B o t 127. 1996

 uni 1

--
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

Fig. 9. Chromatograms in the solvent systems A, B, and C

obtained by high performance thin Iayer chromatography
' (HPTLC). The figures 1-8 on the left side of the
' chromatograms represent Rf classes according to Culberson

& Kristinsson (1970). The distance betwecn thc dottcd line

(where the acetone extract is applied) and the upper edge
(where the solvent front stops) is 45 mm. Abbreviations of
substances: nor = norstictic acid, atr = akanorin, zeo = zeorin,
uni 1-4 = four unidentified metu-depsides in the orcinol series:
and 4-0-mcr = 4-0-methylcryptochlorophaeicacid. A ''km
means that the substance is only occasionally present. The
standard reference with norstictic acid and atranonn is
abbreviated "ref'. The following chemical patterns have been
+

--
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-

Opera BoI 127. 1996

Tab. 7 . The minimum number of phorophytes, lignicolous and saxicolous occurences, and important phorophytes for the North
covering at least 10 % of the total number of collections with known subskate of that particular species. The number o f collections

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

Opera not. 127. 1986

Tab. 8. The number ol' taxa, number of cndemic taxa, and geographic connections in lhe phytogeugraphic elcmcnts represented
among the corticolnus and lignicolous species uf Brrrirlirr (28 taxa, 27 spccics) and Baciclinu (12 species) ill North Amcrica.
-
Cnntincnts on wl~ichthe Nor111 American spccies are known have been markcd, bid this list is almost certaiiily incurnplete. Only
collections seen by mc or rcasotlahly rcliablc reports have been included. Contine~itsare abbrcviatcd Eu Europe, As = Asia, SA
= South America, A f = Africa, and Au = Australia.

Elcmcnt No, of taxa No of endenlics Spccies known also frotn

Subclcmcnt (Bucidiu, Bacldina) (Brrcidiu, Rucidi~~u) Et1 As SA Af Ail

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.

The northern Pacific coast subelement

The Pan-North American subelement
T h e northern Pacific coast subelement consists of
Only one species, Bacidia circzi~nspecta,can be said to five species with similar ecological demands as the oce-
belong to the Pan-North American subelement, al- anic subelement, viz. Bacidia absisfens, B. arierswaldii,
though it appears that this species avoids much of the B. laurocerasi subsp. idahoensis, B. salrnonea, and Ba-
southeastern coastal plain, the Great Plains, and high cidina ramea. They are mainly restricted to the coastline
altitudes in most of the Rocky Mountains. Bacidia cir- and low elevations of tlie western slopes of the Coast
curnspecta is widespread in Europe. The endemism Range and the Cascades from central California to Brit-
among lichens with this distribution type is low or non- ish Columbia. Bacidia uuerswald~iand B. laurocemsi
existent (Brodo & Hawksworth 1977). subsp. iduhoensis also occur in areas with fairly high
precipitation on the western slope of the mountain rang-
es closest to the east of the Cascades in Idaho. Montana,
The eastern temperate and Appalachian and British Columbia. Bacidina rnmea occurs along the
subelements coast to southern California, but has been referred to
this distribution type, since it appears rather poorly de-
One species, Bacidia schweinitzii, clearly belongs to veloped in the southern IocaIities, and since the north-
the eastern temperate subelement, whereas Bucidia lau- em localities are very rich in well-developed individu-
rocerasi subsp. laurocsrusl belongs to the Appalachian als. In other words, the localities in southern California
subeleinent with a Great Lakes-Appalachian distribu- appear to be a southerly extension of a mainly northern
tion type. Probably, Bacidia rosella also belongs to the distribution.
latter subelement. Three species, viz. Bacidia d ~ ~ a c t a , Bacidia absistens and B. ouerswuldii are known from
B, polychrora, and B. sufisa, have distribution types in- Europe, wllereas tlle rernaindcr of the taxa are endzr~lic.
termediate betweeen these subelements. All three occur Thus, the endemism (60 %) in this subelement is the
over much of the eastern temperate region, but they highest of all.
have a clear preference for the Great Lakes-Appalachi- This subelement corresponds to the "lowland to west-
an-Ozark Plateau regions. Possibly, the intermediate ern montane (humid forests)" plus parts of the "coastal
distribution type, and perhaps also the eastern temper- lowland" distribution type of Brodo & Hawksworth
ate type have been derived fiom an original Appalachi- (1977). The connections of this subelement has been
an distribution type following the glaciations. Among discussed by J~rgensen(l 983).
the species discussed here, Bacidia schweinitzii and B.
Irmurocemsd are known from eastern Asia, B. Iatirocem-
si subsp. laurocerasi and B. polychrola from Europe, The Appalachian-north Pacific coast subelement
whereas B. diffiracta and B. sufsusa are endemic. The
former of the endemics is closely related to B. poly- A single species, Bacidia rubellu, belongs to the Appa-
chma, whereas the latter is inuch closer related to the lachian-north Pacific coast subelement. The bulk of the m

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.

Opera Bot. 127. 1986

Tab. 9. The binary matrix on which the cluster analysis of nine North American and three European regions was based. The
species of Bacidia are coded 1 for regions where it is present and 0 for regions from which it is absent. A species has bee11 coded
a s present even if it is now considered extinct from the region (concerns BRIS and SSWE). The abbreviations of the regions
WPAC etc.) are explained in the text.

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.

Opera Bot. 127. 1996

 Similarities
1.O 0.9 0.8 0.7 0.6
l I I I

(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

Opcm But. 1 2 i . 1996

from FLPE and NPAC, SPAC is the most deviant North (l) Lichcnizcd with grccn, chlorococcoid algae.
American region. The eastern North Amcricaii regions (2) Thallus cmstose, never dissolving into goniocysts.
GLNA, SAPP, OZAR, and COPL are inore similar to Uppcr cortcx two-laycrcd, the uppcr layer an ~pinecral
one another than to other rcgions. layer with crystals.
(3) Biatorine or lecideine apothecia.
(4) Proper exciple well-developed, without thick geIati-
Discussion nous layer along the rim. Hyphae of proper exciple dis-
tinctly radiating, more or less abundantly furcate but
Generalizations based on the results obtained here can- sparingly or not at all anastomosed. Cell lumina long
not be made. Similar studies of tloristic similarity based and narrow (but lumina near rim of proper exciple often
on other genera are required, and ideally the entire li- enlarged), often distinctly constricted at the scpta (not in
chen flora of the regions involved should be used. Never- spccies with veiy narrow cell lumina). Cell walls heavi-
theless, the result must be applicable to the species on ly gelatinized, the distance between two adjacent cell
which it was based, and it calls for an explanation. The lumina (1-)2-5 times the width of a lumen. Some spe-
northern Pacific region is more similar to nemoral parts cies deviate from (his patlem in one or rarely two of
of Europe than to other areas of North America concern- these features.
ing the species composition of Bacidia. A hypothesis to (5) Hypothecium not chondroid, of distinctly different
explain this obscrvation is lhal the West Coast, com- texture than the proper exciple and distinctly delimited
pared to other regions, has orfered better possibilities of from this.
survival during periods of glaciation for species that (6) Paraphyses unbranched or sparsely branched in up-
were once more widespread over the continent. The rea- per part, sparingly anastomosed, not with a thick, gelati-
son why northern Sweden, northern North America, and nized wall in upper part.
the mountainous parts of the United States appear so ( 7 ) Hymeniurn and subhynleniurn amyloid, often dis-
simiiar is probably that thcse regions house part of thc tinctly pigmented in upper part.
same, widespread circumborcal element. It is not sur- (8) Tholus amyloid, with a narrow c-layer, with a l
prising that the Florida Peninsula is so different from high, blunt or pointed axial body, the width of which is
other regions, considering that a majority of the species a third or less of the entire d-layer, and a low or high,
are of tropical origin. Had a neotropical region been in- blunt or pointed ocular chamber.
cluded in this analysis, it would almost certainly had (9) Spores acicular, clavate, bacilliform, or fusifonn,
come out next to the Florida Peninsula in Fig. 10. when mature with three or more transversal, distinct
septa. They are colourless, rather thick-walled, and lack
Taxonomic delimitations above the species perispore or ornamentation.
level ( l 0) Pycilidia with cylindrical conidiogenous cells (7- 15
pm long) that produce conidia terminally. Conidia usu-
I have attempted to characterize Bacidia S. str and Bu- ally filiform and curved, either formed singly or repeat-
cidina, to outlinc thc differences between these genera, edly from the conidiogenous cells.
and to delimit them from other genera. This work is pre- (1 1) Sometimes producing atranorin, rarely zeorin or 4-
scnted hcrc as a preliiniilary hypothesis based mainly on 0-methylcryptochlorophaeic acid or unidentified orci-
North Amelicail species. I have, however, tested it on a no1 rneta-depsidcs.
number of species From other parts of the world, mainIy
Europe and South America, and so far it has proved suc- Bacidina is circumscribed here as to include, species
cessful. with the features mcntioncd bclow. The set of charac-
ters has bccn choscn on thc sarnc grounds as for Bacid-
ia.
Characterization of Bacidia and Bacidina
(1) Lichenized with green, chlorococcoid algae.
Depending on what other genus Bucidiu is compared (2) Thallus crustose, often partly or entirely dissolving
with, a particular set of characters can be used to sepa- into goniocysts. Upper cortex two-layered, the outer lay-
rate them. In the following characterization of Bacidia, I er an epinccral layer without crystals.
have included the minimum amount of characters neces- (3) Biatorine, Iccideine, or exceptionally zeorine (with a
sary to convincingly separate this genus from Bacidina thin thalline margin) apothecia.
and from the gencra trcatcd in the section Related gen- (4) Proper exciplc wcll-developed, without thick gelat-
em and species groups: inous layer along the rim. Hyphae of proper exciple in-
distinctly or distinctly radiating, more or Iess abundantly
furcate but sparingly or not at all anastomosed. Cell lu-
Tab. 10. Observed differences between the genera Bacidia and Bacidinn, as understood in this work.

Bacidia Bacidina

Thallus cortex With minute crystals Without crystals

Thallus structure Never dissolved into goniocysts Often dissolved into goniocysts

Apothecia Usually strongly pigmented, Often weakly and diffusely

not seldom in distinct layers pigmented

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

Terminal cell of conidiophores 7-1 5 pm long 4-8 pm long

Secondary chemistry Often atranorin, rarely other None

substances

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

In my opinion, Bacidia and Bacidina are closely related

Open Bot. 127, 1996

Related genera and species groups walled hyphae with lumina (laterally, just inside the
rim) that are niore 01. less rectangular (up to 12 p n long,
This section concerns a number of genera and spccics gcncrally 1-2 times as long as wide) and not constricted
groups that are related to (11- have recently been thought at thc septa. Furthemiore, the geilus differs fro111Bocidi-
to bc rclatcd to Bucidiu and Bacidina. I have included nu i l l having a nanower axial body, and in having morc
all genera accepted in the "Bacidiaceae" by Eriksso~~ & rcgulnrly rectangular cell lumina witl~outconstrictions
Hawksworth (1993) and some additona1 genera believed at tlic septa inside the rim.
Lo bc rclcvant to thc discussion (Auric~rlorn,Bc~!,alt~iuiu. Although distinct from Buciriiu and Briciclinn. Ar~hr-o-
E.schutugunin, ,Wrgalcir.in, Mvriotzor-U, Physcidiu, Sco- .spol-trri~appears to be vcry closc to Tonittia. As also
licims~~nr~int,Sporuue.srru, Tunitliu, and T~~lothallia). pointed out by Timdal (1 99 11, ~ll-throsporuni differs
Furthermore, tliree species groups, lnai~llyconsisting of froin Tonirtia only in the co~ticoloushabit and in the
species that 1 exclude from Bncicliu and Buciciina, arc polysporous asci. There is a distillcl possibility that AIL
treated. They are all Inore or less closely related to Bu- fhrosporlmi should he included in Toflinicr, considering
cidia a i d Oaoiclirza, and Inay constitute r~ndescribed that tteology hardly is a reason to separate gencrrr and
genera or parts of already described genera. The list OS h a t thcrc are several other examples of octosporous
rclatcd spccics groups could easily have been made spccies that arc closely related to species with polyspo-
longer, but l h a w rcstrictcd the discussion to groups rous asci (in: c.g., Cur?a'elor-iellrr,Crilupl~~ru, Leoania,
thai are well-characterized and rclativcly rich in species. and Lucunora).
It should be noted lhai in a bricf discussion by Hertel &
Rambold (1995), Cur~ip.surl~~diu~ta, Esrhurugunicr, Pltys- Auriculora Kalb
cidiu, and Jupewiu were thoi~ghtnot to bc n~ci~lbers of
the Lecanoraceae (incl. Bacidiaceae). I admit to havc Kalb 1988: 2.
treated the gcncra bclow very inconsistently. The
amount of i n f o i l ~ ~ a t iincluded
o~~ is more or less correlat- Type: A. h)).aoti~orphrr (Nyl.) Kalb (holotype).
ed with my knowledgc and intcrcst in the particular ge-
nus. Tonir~iupsisFrey and Psorellc~Miill. Arg. are con- Thc genus Alrriclrloru was established by Kalb (1988)
sidcred here to be synollylns of Bucicliu and arc trcated Tor thc single species, .4. I?l;ssut~~uryha. It was statcd to
as such under this genus 111 the T~~xonomj: chapter. have "asci silniies typo Lecanoraceae" and was pIaccd
in thc Lecanoraceae by Eriksson & Hawkswwth ( 1989:
199 I b) for this reason. Later studies by Heiissell & Titze
Adelolecia Hertel & Hafellner (1990) led Eriksson & Hawksworth ( 1 991a, 1993) to
put the genus ainong the Lecanorales incertae sedis.
Hafellner 1984: 260. Serusiaux (19931, however, considered the asci ro be of
"Bucidicr-Lype" and refcrrcd .411riclllorn to "Bacidiace-
Type: A. piluti (Hepp) HerteI & Hafelb~er(holotypc). ae" with a query
Aur~iculorcr1~ys.rornul-l;lha has ear-like portions of old
Adelolecia collsists of two known species, A. piluti and decaying hyn~cniaattached to tlie edge oT Lhc younger
A, lcolaensis (Nyl.) Hcitel & Ra~nbold(Hertel & Ram- hymcnia. According to Hensse & Titze (1 990), this pc-
bold 1995). The genus differs fi-urn Bcrcidia and Bucidi- culiar appearancc is caused by the formation of new hy-
na mainly in having more richly branchcd paraphyses menia below thc old oncs, which disintegrate. My own
and 0- I-septate, ellipsoid spores. observaiions of thc tholus show that it is not at all simi-
lar to the ones foui~din Bacidiu. Instead, it possesses a
wide axial body rc-aching all through the d-layer, and is
Arthrosporum A. Massal. thus rather similar Lo thc thulus appearance found in
~ ~ O ~ U I(Ach.)
S L ' O ~ I C ~ O Snin~bri~~l,nr II Arnold (see Hafellner
19x4: 340). Furthennore: Illrriculora differs fro111 Ru-
cidia and Rncirlir~uin having a non-arnyloid hymenial
Type: A. popziirn~.un~ A. Massal (holotype) gcl.

The gcnus Artlar-ospo~umwas re-established by Harell-

ller (1984) to accomodatc tlie single known species, A. Bacidiopsora Kalb
yu~~ntlor-11mA. Massal., previously known as Brrr,iclia
poplllurirr~~(A. Massal.) Trcvis. or Bacilliu ucclinis Kalb 1988: 4.
(Flot.) Zalilbr. ArtJ~r.r~spur~m~
diffcrs froin Baciu'icr by
possessing a proper exciple built up of fairly thin- Type: B. squu~~zz~losula
(Nyl.) Kal b (holotype).
spores with a finely warted perisporc. It diffcrs from L. differs markedly from both Bacidia and Bacidinu by
hypnnru.urn mainly in having consistently two-celled possessing a fruticulose, richly branched, homoioiner-
spores, a much thicker proper cxciplc (latcrally exceed- ous thallus (forming small peltate hemispherical tufts)
ing 100 pm), and a highcr hy~nenium(c. 100 pm). Thus, containing a mixttlre of cyanohacteria and chlorucoc-
Catinal-iu is the oldest generic name for Lecidea hypno- coid algae, an apothecial margin o f thalline origin (al-
rum and its relatives, a group worthy of recognition at though not containing algae), richly branched and anas-
genus level. Carinaria has never been used in this sense, tomosed paraphyses, and in having the thalIus inspcrscd
and this a furlher strong reason why Catinar.ia should with yellowish granules that dissolvc with C ii~loa rose-
be conserved with C. afvopurpurea as the conservcd rcd solutiou (substance not identified). The asci and the
type species spores of Con~psoclurlium,however, are strikingly simi-
lar to Baoiclia 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

Hawksworth et al. 1980: 106,

Compsocladium I. M. Lamb
Type: H. tuylorii (Salwcy) P . Jamcs (holotypc).
Lamb 1956: 157
I-lerteliana compriscs two species, the type species, H.
Type: C. archbolclinnzrm I. M . Lamb (holotype. not seen). faylorii (Salwcy) P. Jaincs, and H. alriskrnsis (NyI.) S.
Ekinan (see E.rulwdec1 ur nut e.'iunlined species). rt dif-
According to Lamb (1456), this monotypic genus, de- fers from Baciu'ia and Bacidinu In having a root-like hy-
scribed from Irian Jaya (formerly Dutch New Guinea), pothecium extending into the thalline medulla, non- or
3-septate spores, and a "paraplectenchymatic" proper cies with the following features: (l) lichenized, crustose
excipIe which is not distinctly delimited from the hy- thallus (or fruticulose, if Thumnolecania is included,
pothecium and composed of densely packed hyphae see below), with or without goniocysts (often termed
with broadly rectangular to ellipsoid cell lumina. blastidia in Lecania terminology, but I fail to see the
difference), (2) lecanorine to biatorine apothecia with a
proper exciple consisting of few to many hyphae, (3)
Japewia Tansberg fairly abundantly furcate but non-anastomosed excipu-
Iar hyphae, the cell lumina of which are narrowly cylin-
drical (1-1.5 pm wide) in the interior of the proper exci-
ple but gradually become thicker towards the rim (up to
5 pm wide), thus giving each hypha a characteristically
Type: J. tornoensis (Nyl.) Tsnsberg (holotype). club-shaped appearance, (4) yellowish, brown, and
blue-green pigments in the apothecia that are often
Three species were referred to Japewiu by T~nsberg markedly unevenly distributed in the upper part of the
(1990). It was placed in the Bacidiaceae by Eriksson & hymeniurn;thus giving the apothecid disc a characteris-
Hawksworth (l99la) "on the basis of ascus structure". tically "dotted" appearance in a wet condition (not seen
Its placement near Bacidia is questionable, however. in pigment deficient specimens), (5) paraphyses that are
Japewia is probably closely related to Mycoblastus Nor- easily separated in K and gradually thicken in the upper
man. Jupewia can be separated from Bacidia and Ba- part, the terminal cell being up to 5 pm, (6) 0-3-, rarely
cidino on account of the more abundantly anastomosed up to 5-septate, ellipsoid, fusiform or bacilliforrn, rather
paraphyses with apices surrounded by a gel coat, very thick-walled spores that are often slightIy curved but
wide asci, and large, thick-walled, non-septate, some- lack perispore and ornamentation, (7) a tholus with a
times abortive spores that often possess a gelatinous fairly low and blunt, more or less conical ocular charn-
perispore. ber, and a fairly high, conical axial body that is often
but not always (often lacking in L. cuprea) surrounded
by a heavily amyloid zone that is darker than the re-
Lecania A. Massal. mainder of the d-layer, (8) strongly curved (or some-
times sigrnoid in L. fuscella), 0.5-0.8 pm wide, O-l-sep-
Massalongo 1853b: 12. tate microconidia and more or less curved, 1.0-2.0 p
wide, 0-l-septate (5-7-septate in L. naegelii) macroco-
Type: L,fuscella (Schaer.) A. Massal. (holotype) nidia.

Traditionally, Lecania has been restricted to species

with a lecanorine apothecial margin. Such a delimita- Megalaria Hafellner
tion of this genus is artificial, however. When several
characters are taken into consideration, it becomes clear Bafellner 1984: 302.
that Lecaaiu includes a variation from typically lecanor-
ine species like L. fuscel/a and L. cyrtella, to intermedi- Type: M. gmssu (Nyl.) Hafellner (holotype).
ate ones with a poorly developed thalline margin (L.
cyrtellina, L. hutchimiae), and finally to a number of Megularia is presently comprised of six accepted spe-
species with a biatorine margin. Four species with a bia- cies (Ekman & Tsnsberg 1996), but the number .will
torine apothecial margin should be referred to Lecania probably increase in the future. It is distinguished from
(all formerly placed in Baeidiu), viz. L. naegelii (Hepp) Bacidia and Bacidina mainly on account of the one-sep-
Diederich & P. Boom, L, cuprea (A. Massal.) P. Boom tate, thick-walled, ellipsoid to subglobose spores, the in-
& Coppins, L. sub@cula (Nyl.) S . Ekman, and L. stig- distinct ocular chamber, and (when applicable) the
naate!la (Tuck.) S. Ekrnan. The first two species were short, ellipsoid, oblong, or ampulliform conidia.
transferred from Bacidia to Lecania by van den Boom
(1992) and van den Boom et al. (1994), unfortunately
without any discussion on the grounds for this. Future Mycobilimbia Rehm
investigations of Lecidea S . lat, and Carillaria S. lat.
may show that several species should be included in Le- Rehm 1889 (1887-1895): 327.
cania. The European saxicolous species of Lecania in
the traditional sense have been treated by Mayrhofer Type: M. obscuurata (Sornmerf.) Rehm (Iectotype select-
(1988). ed by Clements & Shear 1931: 3 15).
In its revised circumscription, Lecania includes spe-
Mycobilimbia is closely related to and very similar to thaleriza and excluded this species and P. s ~ l o p h o m
Biatora, but was separated from this genus by Printzen from Phyllopsora to two undescribed genera in Bacid-
(1995) on the basis of a different apothecial ontogeny. iaceae and Lecanoraceae, respectively.
Four species, M. retramem (syn. M. obscumta auct.),
Catillaria sphaeroides, Bacidia epixanthoides, and Ba-
cidia carneoalbida, were tentatively included. This de- Physcidia Tuck.
limitation is quite different from the one suggested by
Hafellner (1989). The typification of M. obscurata is Tuckerman 1862: 399.
complicated; see note by TimdaI (1991 : 26).
Type: P. wrightii (Tuck.) Tuck. (lectotype selected by
Clernents & Shear 193 1: 322, not seen).
Myrionora R. C. Harris
Kalb & Elix (1995) include seven species in the genus
Harris et al. 1988: 27 Physcidia, which can be separated from Bacidia and
Bacidina on the basis of tlie squamulose or foliose thal-
Type: M. albidula (Willey) R. C. Harris (holotype, not lus that often produces isidia, tl~edistinct upper cortex
seen). lacking an epinecral layer, the thin-waIled, l-3-septate
spores, and the different secondary chemistry.

This monotypic genus appears doubtfully distinct from

Scoliciospomrn, differing from Suoliciospomm cuwa- Rolfidium Moberg
turn Serus, mainly in having straight spores, the number
of which exceeds 30 per ascus (Harris et al. 1988, Seru-
siaux 1993). Polyspory is known in S. curvaturn Skrus.
and straight spores are found in S. chlorococcum Type: R. peltaturn Moberg (holotype, not seen).
(Graewe ex Stenh.) VBzda and S. gallurae VSzda & Po-
elt. A similar variation in the number of spores per as- Two species have been referred to Rolfidiurn, namely
cus is found in Candelarie.iellaand Ropalospora. For the the type species R. peltatlim Moberg and R. coccocarpi-
distinction from Bacidia and Bacidina, see Scoliciospo- oides (Nyl.) Timdal (Timdal 1991). According to
rum. Mobeg (1986), this genus differs from Bacidia and Ba-
cidina in forming small, indistinctly lobate, foliose thal-
li with rhizines, in having ellipsoid, l-septate spores,
Phyllopsora MiiI1. Arg. and large pycnidia containing sparsely branched conidi-
ophores and filiform, more or less straight conidia.

Type: P. breviuscula (Nyl.) Miill. Arg. (lectotype select- Schadonia K6rb,

ed by Clements & Shear 193 l : 3 19, not seen).
Korber 1859: 93.
Phyllopsora is a medium-sized genus with a world-wide
tropical and subtropical distribution. According to Brako Type: S. alpina Korb. (holotype).
(1989, 1991) and Swinscow & E h g (198 l), Phyllopsora
differs from Bacidia and Bacidina mainly in having a usu- Schadonia, comprising only two species (Poelt & Vczda
ally squamulose, sometimes isidiate tlmllus with a well-de- 1981), differs from Bacidia and Bacidina in having
veloped cortex lacking an epinecral layer, a web- or mat- more abundantly branched and anastornosed paraphy-
forming prothallus, a hypothecium and a proper exciple of ses, and asci with 2-8, large and muriform spores.
the same textwe (the texture of the hypothecium being
very similar to the texture of the cortex), 0-l-septate
spores, and in lacking a distinct subhymenium. Swinscow Schistoplace Brusse
and Krog (1981) discussed four species, Phyllopsom
thaleriza (SW.) Gotth. Schneid., P. cornpaginata (Mial. Brusse 1987b: 245.
Arg.) Gotth. Schneid., P. stylophora (Malme) Gotth. Sch-
neid., and Bacidia lacifiiosa Swinscow & Krog, that they Type: S. alvearda/is Bmsse (holotype).
understood as intermediate between Bacidia and PhyIIop
som. Brako (1991) synonymized P. compaginta with P. Two species were referred to Schistoplaca by Brusse

Open Bot. 127. I996

(1987h). The genus was reduced into synonymy with nally and laterally, and shorl-cylindrical conidia
Lecarlan; by Lull~bscll& Feige ( 1994). (Hafellner & Egan 1 98 1 ).

Scoliciosporum A. Massal. Sporacestra A. Massal.

Massalongo 1852: 104. Massalongo 1860: 264.
Type: S. laolontelaenlrr1l (Fldrke) A , Massal,, a younger T ~Bifl~oyn
~ p,-asfita
~ : ~ ~ c &k Mont,,
. llolll, illcg,.
synonym of S. irrr~bririirm(Ach.1 Amold (lectotype se- ~~~,i~~~1856 (hulotype),
lected by VGzda 1978: 414).
Sl~ur-uuerfruis thc oldest generic name for Psurellcz per-
LSculiciosportrnl.to which altogelher nine spccies Iiavc !c-ctrl(Nyl.) Miill. Arg., P. naicr.uphylliraa (Tuck ex Nyl.)
been referred (VGzda 1978, Vixda & Poelt in Niinis 8c Zahlbr, and P. cogn(ifu (Nyl.) Z;~hlbr.,a group which
Poelt 1987, SCnlsiaux 1993), is distinguished rrom Ba- was tentat.ively proposed as a gcnus different from Ba-
cidia and Bacidinrr on account of the poorly developcd cidiu and Phy1lupsor.u by Brako { 1989). In fact, the only
proper exciple consisting of paraphysis-like hyphae, the species incIuded in Sparucestrci by Massalongo, Bialo-
abundantly branched and anastornosed paraphyses, the f a pr-usirzu Tuck. & Mont. (a nonlcll illcgitimum), is
widc axial body reaching through the entire d-layer of synonymous with Psor.ellu p ~ ~ r t e x t ua, Fact which has
thc tholus, and (Coppins 1983: 97) the presence of'some bccn overlooked up until now. A legitinlatc i ~ o i n e no-
i~
dcviatiilg paraphyscs ternlinating in swollen apices that vunl, Biaiora prcrsinalu Tuck., was 111ade later (Tuckcr-
are surrounded by a darkly pigmented cap. It should be man 1888), on which Coppins (l 9x3) based the con~bi-
noted that the tholus of S. chlurur.occum (Graewc cx nation Bacidiu j~r-usinutrr(Tuck.) Coppins.
Stenh.) Vezda, contrary to S. I I ~ ? ~ I . (depicted
~ I ~ U I ~byI SporficrLv!rcrdiffers froill Bcrcidia and Bacidina in
Hafellner 19X4: 340) has a rather distinct, heavily amy- having a finely squamulose thallus, cxcipular hyphae
loid zone surrounding the axial body. that often protrude from the lower side of thc apoth-
eciunl (anchoragc hyphae'!), paraphyses with an apical
swelliiig duc tu a thickened gelatinous wall without a
Solenopsora A. Massal. colresponding widening of the cell luinen (similar to
Massalongo 1855: 20 the paraphyscs of Squamocidic~,see Brako 1989: 7, fig.
3D3, and thin-wallcd spores with indistinct sepia. Ac-
cording to Brako (1989), this species group is separated
Type: S. r-equieraii A. Massal. (holotypc: not seen).
from Phyllopsora on account of the different tissuc
Tlie delilnitatioll of this genus i s still puurly invcstigat- types iri the hypothccium and the proper exciple, the
ed. The core of Solenr)/;r.wruappears to be distinguished
less gelatiriized paraphyses, and tlic longer and septate
froin Bacidia and Bucidrna on account of the placodioid sporcs. Duc to an observed variation in lhc ontogeny of
ur squamulosc thallus, tI~cleca~iorineapothecia, the 1-
Lhc tholus and a suspicio~ithat several specics prcscntly
scptate spores, the lack of both an ocular chaniber and
included in Uucidiu should be included in thc P , ~ o r ~ l l a
pel-textri goup, Urako (1989) at the time refrained from
an axiaI body in thc ascus, the branclicd conidiophores.
treating this group as a scparate genus. With the possi-
and the short-bacillifoin~conidia (Poclt 1969, Hertel 8c
Rambold 1988: Purvis & lamcs in Pun;is et al. 1992). ble exceptio~l or Bcrcidia nlrgu.rlinii (Tuck.) Zal~lbr.,
none of lhe North American species belong here.'Fur-
thermore, I havc sccn the types of nearly all South and
Speerschncidera Trevis. Central American Bocidia S. lat. with long and septate
spores, hut no species is cluse to P,rorella per-lextu.
Thus, judging iiom (he neotropical species, it seems un-
likely that Sparucestrci, if accepted as a genus, is much
Type: S. eiipbcu (Tuck.) Trevis. (hololype, not sccn). richer in species than origii~allythought by Brako.

The monotypic genus Spcerschneidera differs from Bu-

cidia and Bacidina in being foliose, and in having a Squamacidia Brake
thick cortcx of pcriclinally arranged hyphae, paraphyses
with a distincl cap of pignleill in thc disk11 part nf the Bmko 1989: 6.
swollen terminal cell, generally l-septatc sporcs, un-
branched coiiidiophorcs producing conidia both fermi- Type: S. janeircnsis (Miill. Arg.) Brako (holutype).
According to Brako (19891, the monotypic genus Squa- Thamnolecania (Vain.) Gyeln,
macidia differs from Bacidira and Bacidina by possess-
ing a squamulose and isidiate thallus with a cortex lack- Gyelnik 1933: 8
ing an epinecral layer, l-3-septate spores with thin
walls, rod-shaped conidia sometimes produced in Type: T. brialrnontii (Vain.) Gyeln. (lectotype selected
groups of 2-3 terminally on the conidiogenous cells, and by Dodge 1948: 181, not seen).
in having a different secondary chemistry. Furthermore,
my own observations show that Squa~laoeidiuis separat- Gyelnik (1933) recognized two species, and Dodge
ed from Bacidia and Bacidina also in having a quite dif- (1948) described two more species in Tharnnolecania. .
ferent proper exciple, which is composed of hyphae All are fruticulose and more or less Antarctic in distri-
with narrow cell lumina with gelatinous walls, the bor- bution. Hafellner (1984) questioned the distinctness of
ders of which remain distinct and can be easily seen in a Thamnolecunia, since it apparently differs from Leca-
light microscope. Furthermore, the terminaI cells along niu only in the growth form of the thallus.
the rim of the proper exciple are not the least swollen.

TibelIia VGzda & Hafellner

Squamarina Poelt
Hafellner & VGzda 1992: 186.
Poelt 1958: 524.
Type: I: dirnep.elloides VGzda & Hafellner (holotype,
Type: S. gypsacea (Sm.) Poelt (holotype). not seen).

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).

Tephromela M. Cboisy Toniaiu was given a detailed monographic treatment

and a new circumscription by Timda1 (1991). For a long
Choisy 1929: 522. time, the delimitation adopted by Fries (1874) has been
used. In his sense, Toninia included all squamulose spe-
Type: T, atru (Huds.) Hafellner (holotype). cies of the Lecideaceae having septate spores. The new
and improved delimitation provided by Timdal (199 1) is
About 20 species have so far been referred to the genus mostly concerned with the anatomy of the paraphyses
Tephromela (Bertel & Rambold 1985, Poelt & Grube and asci. Timdd recognized 48 species in Toninia.
1993, Rambold 1993, Haugan & Timdal 1994). Teph- Toninia is obviously very closely related to Bacidiu.
romela differs from Bacidiu and Bacidina in having They share features in the paraphyses, asci, spores, and
thick-walled paraphyses, non-septate ellipsoid spores, pigmentation. In particular, group 3 of Timdal (1991:
branched conidiophores producing conidia laterally, ba- 28), containing T, uromatica and its relatives, is very
cilliform conidia, and a different chemistry. n e dark similar to Bacidia cip.curnspecta,B. igniarii, B. illudens,
purplish hymenium and the lecanorine apothecia gener- B. subincompta, and B. schweini~ii.The members of
ally associated with the most well-known species, T, ut- these groups share a similar apotheciurn pigmentation,
m (Huds.) Hafellner, are typical of only a portion of the and bacillifonn or acicular, 3- or more-septate spores
genus. are present within both groups. However, Toninia ap-
pears distinct from Bacidia, and I have so far found no Three species have so far been referred to Way~rea,
corticolous species traditionally treated in Bacidia to be- viz. W. cal$ornica Moberg (Moberg 1990, 1991), K
long in Toninia. stoechadiana (Abbassi Maaf & C. Roux) C. Roux & P.
Toninia can be separated from Bacidila on account of Clerc (Roux & Clerc 1991, Roux & Giralt 19911, and
the anatomy of the proper exciple. Contrary to Bacidia, W. aadscendens V. J. Rico (Rico 1991, Bticaud & Roux
the proper exciple of Toainia is made up of hyphae that 1993). The taxonomy has been summarized by Roux et
are not furcate, but more or less abundantly anastomo- al. (1995). Waynea differs from Bacidia and Bacidina
sed. The lumina are shorter and wider relative to those of by possessing a squamulose thalIus with lip-, cup, or
Bacidia, usually 5-15 (exceptionally 20) pm long and helmet-shaped soralia, a distinct proso- or paraplecten-
1.5-5 pm wide (lumina near rim of proper exciple ex- chymatic upper cortex, rather short, bacillifom to fusi-
cepted), commonly 2-5 times as long as wide (excep- form, 3-septate spores, and cyIindrical excipular cell
tionally up to 8 or down to 0.75 times as long as wide), lumina. Wuyneu shows an interesting western North
hardly ever constricted near septa (T. coelestina except- American-Mediterranean disjunction.
ed). The walls are gelatinized but, compared to Bacidia,
thinner, the distance between two adjacent cell lumina
being about the same as the width of a cell lumen (some-
times shorter, sometimes up to twice the width of a lumi- The Bacidia beckhausii group
num).
A' number of additional differences between Toainia Three species belong to this group: Bacidia beckhausii
and Bacidia were accounted for in the comparison by Korb., B. hemipolia (Th. Fr.) Malrne, and B. verecundu-
Tirndal (1991), in which he used the type species of Ba- la (Th. Fr.) H . Magn. The last is a poorly known species
cidia, B. rosella, as a "model species". Once more spe- that has been collected only once, in northern Norway.
cies of Bacidiu are taken into consideration, it becomes The report from Northwest Territories, Canada, is erro-
clear that the listed differences cannot be used to sepa- neous and refers to an undescribed species in the Baci-
rate the genera. It is clear, however, that Timdal was dia sabuletomm group (see below).
well aware of the dangers with using type species as The Bacidia beckhausii group differs from Bacidia
models. and Bacidivta in having a more poorly developed proper
Toninia differs from Bacidina mainly in having a nar-
exciple, a tholus with a blunt and indistinct ocular
rower axial body, and in having more regularly rectangu- chamber or no ocular chamber at all, an axial body that
lar cell lumina without constrictions at the septa inside reaches all through the d-layer and has sides that are
the rim. Furthermore, compared to Toninia, the species parallel or upwards slightly convergent, a rather distinct
of Bacidina usually contain lower amounts of pigment zone surrounding the axial body that is more heavily
in the apothecia, sometimes no pigments at all. amyloid than the remainder of the tholus, and spores
that are baciIlifom to fusiform and usually three-sep-
tate .
Tylothallfa P.James & H.Kilias
The B. beckhawii group shares the poorly developed
Kilias 1981 : 409. proper exciple and the general appearance of the tholus
with ScoIiciosporuin, and together with the Bacidia lutes-
Type: T. biformigera (Leight.) P . James & H. Kilias cem group it appears to form a Iink between BacidialBa-
(holotype). cidina and ScoIiciosporurn. Contrary to Scoficiospomna,
however, the excipu1a.r hyphae of the B. beckhausii gmup
Two species, i? biformigera (Leight.) P . James & H. do not resemble paraphyses, and the axial body is dis-
Kilias and T. pahiensis (Zahlbr.) Hertel & H. Kilias, tinctly narrower. The B, bech%ausii group is very closely
have been referred to this genus (Kilias 1981, Bertel related to the B. lutacens group, and it seems probable
1983). They differ from Bacidia and Bacidina in having that they together should form a genus that is yet to be
abundantly branched and anastomosed paraphyses, con- described. These groups differ mainly in the amount of
sistently I-septate spores, a different chemistry, and pigment in the apothecia and the presence of oil droplets
(Timdal 1991) smaller conidia. in the hypothecium and proper exciple. B, hemipolia,
however, exhibits a continuous variation in the pigmenta-
tion of the apothecia (not seldom on the same thallus)
Waynea Moberg from darkly pigmented to entirely pigment deficient. De-
spite the close relationship between the B. beckhausii and
Moberg 1990: 249. the B. lutescens groups, I have chosen to treat them sepa-
rately in order not to loose any information. B. beck-
Type: W. californica Moberg (holotype). hausii itself was treated as a Micarea by Poelt & Vizda

Opera Bot 127, 19%

(1977) but it was cxcluded from that genus by Coppins The Bacidiu sabult.tnnlm group almost certainly rep-
( I 983). resents a separate genus. It was included in Mycubilim-
bia by Hafellner ( 1984, 1989), but 1 am coilvillced that
it is not congeneric with the taxoil usually treated as the
The Eacidia lutcscens group type of wvcohilintbia, M obscilrufu auct. (= M ~ e t m m -
era).
The Bacidia /utescens group is comprised of Bacidiu
lutescens Malme, B, an~eriuunu(Fke) Zahlbr., B. ,fusczl- Family
la (Nyl.) Zahlbr., Scoliuiospurum pruinosum (P. Jarnes)
VGzda (also known as Bacidia pruinosn P. James), and Hafellner ( 1984) referred Bacidia to Bacidiaceae. At
one or two apparently undescribed species from south- that time, Bacidina was not described, and none of the
eastern United States. Once thc tropical members of Ba- species later referred to this genus was investigated by
cidiu S . lat. have been studied in more detail, this group Hafellner. Eriksson & Hawksworth (1993) treated Ba-
may prove to be much larger. cidia in the Bacidiaceae, but their concept of this family
The B. 1utescen.r group is separated from Bacidia, Ba- was much wider than Hafellner's. Bacidina was rc-
cidina, and Scoliciosporum in much the same way as ferrcd to Lecanoraceae by these authors, an action
the B. heclthausii group. In addition, all membcrs of the which was based on the statement "Asci tholis gcneri
B. lufescens group have minute KOH-soluble ciystals in Lecanor-a instructi" in the diagnosis of this gcnus by
the propcr exciple and oil droplcts in the hypothecium Vkzda (1991).
and sometimes also in the proper exciplc. The oil drop- In the light of the variation observed within Bacidicz
lets arc usually abundant but may sometimes be sparse. and Bacidina, I endorse the thcory uf HerteI & Ratn-
Furthermore, the apoth~ciaare lnostly weakly and dif- bold ( 1 995), that the Bacidiaccae and the Lecanoraceae,
fusely pigmentcd in the upper part of the hymenium and morc or lcss in the sense of Eriksson & Hawkswor-
along the rim of the proper exciple. Like in the B. beck- (1 993) should be united into one family. Lecanoraceac.
hausii group, the axial body is usually surrounded by a Bacidia, Bacidinn, as well as Toninirr (treated in thc
thin zone that is more heavily pigmcntcd than the re- Catiilariaceae by Eriksson & Hawksworth) should be
mainder of the tholus. This zone is lacking ollly in Sco- referred to this family. This should be vicwcd upon as a
liciosyorilnl pruiaosurn. For a discussion on the rela- tentative hypothesis, however. Contemporary liclienolo-
tionship with the B. becklaazlsii group, sec above. gy should be careful not to flatter itself by thiilkil~gthat
a final ascomycete system, which rcflects true phyloge-
netic relationships, has been achicvcd. There are times
The Bacidia sabuletorum group when I wonder if futurc taxonomisis will not look at
our present-day ascornycctc system in much the same
This group consists of Bacidia sabuleforunz (Schreb.) way as we are now lookii~gupun the spsteln by Zahl-
Leitau (including B. uccedens (Amold) Lettau), R. uc- bruckner (1921-1940)!
cedens sensu R. C. H m i s (19771, non (Arnold) Lettau,
B, tnicrocarpa (Th. Fr.) Lettau, Toainia lobu/ata (Som- Numerical and statistical treatment
rnerf.) Lynge, and an undescribed parasymbiontic or
parasitic species on Lecanoru cf. impuclcns Degel. from During my studies of Bucidicr and Bacidir~a,I have co-
Northwest Territories, Canada. me across some taxonomically con~plicatedcases in need
TRc Bacidia sabuletonrm group is closely rclated to of scrutiny. In these cases, 1 have choscn to use i~arious
Biatom and M?;cobilimbia and differs from thcsc genera numerical and statistical methods to throw some light on
mainly in having a more heavily pigmerltcd hy- the problems. Such methods are powcrful tools to sum-
pothccium, stouter paraphyses, more thick-walled asci, marize the numerical information wailable horn meas-
an oily yellowish content in thc mature spores, a green urements, and can thus provide i~nportantarguments in
pign~ei~t in the upper part of thc hymenium, and in hav- favour of or against cerkaiil taxonoinic solutions. Nahlral-
ing a proper exciple, the riin of which is covered by a ly, they do nut supply any direct evidence. The result of a
rather thick gelatinous layer that swells markedly (and statistical or numerical analysis is always open to inter-
finally almost dissolves) in KOH or dilute hypochloritc pretation, as are the results of any othcr mcthod used by
solutions. Furthermore, B, sabuletorum and T. lobulutu taxonomists.
have a warted perispore (although sometiines over- Three cases with a sufficient number of ineasure-
looked, since it is easily detached frorn thc spores). ments are treated here. Among thc North Ainerican Ru-
Apart from the pigmentation of the hypothecium, all cidiu and Bacidina, thcrc are ce~fainlyfurther cases that
these characters also separate the Bacidiu subuletorii~i would benefit from a nuincrical and statistical analysis.
group from Bucidia and Bacidina. For example, the variation within the present concept of
Tab. 11. Descriptive statistics and the significance of the F-values obtained from the one-way ANOVAs comparing B. difffracia
(granular thallus) and B, polychma (continuous thallus). Extreme morphs from Sanford, Florida, have been excluded. The
abbreviations of the characters are explained in the text. The standard deviation i s abbreviated "S. d." The input values used for
APWID, HYM, and PARWID 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 0.001<p~0.01,*** for p50.001, and ns {not significant) for p>0.05.

Bacidfa polychroa (N= 22) Bacidia dzflacla (N= 13)

Character mean S. d. min-max mean S. d. min-max Significance of F
APWID/mm 0.7 0.1 0.6-0.8 0.8 0. I 0.5-0.9 ns
APHGT/w 273 51 176-359 465 95 344-688 ***
EXClPlpm 63 13 45-95 64 14 52-103 ns
HYMIp 77 10 63-95 X2 6 70-90 ns
PARWIDlp 2.4 0.3 1.7-3.0 2.0 0.3 1.6-2.6 **
SPLENlpm 48 5 38-57 45 5 37-54 ns
SPWlDlw 2.7 0.4 2.3-4.0 2.9 0.2 2.6-3.3 ns
SEPTA 6.3 2.0 3.1-11.9 6.2 1.O 4.1-7.8 ns

Bacidia circumspecta (potential correlations between Methods

conidial types and other characters), the relationship be-
tween h i s species and B, igaiarii, and the relationship Twenty-two specimens of the continuous morph (Bracid-
between B. hostheleoides and B. medialis would have ia polychroa) and 13 specimens of the granular morph
gained from a closer investigation. In these cases, how- (B. d ~ ~ a c twere
a ) used (the same ones used to prepare
ever, the measurements available from North American the descriptions of these species). In addition, two spec-
maierial are too few in number. With the exception of imens of the Florida morph from the same locality were
European Bacidia luurocerasi, the study of extra-North studied. Eight characters were measured on these speci-
American material has not been wihin the scope of the mens:
present study.
1. Apothecium width (APWID).
2. Apothecium height (APHGT).
3. Width of proper exciple lateral to the hymenium
Bacidia poIychroa and B. diffracta (EXCIP).
4. Hymenium thickness (HYM).
Background
5 . Width of paraphysis apices (PARWID).
6. Spore length (SPLEN).
In the eastern United States and Canada there are two
7. Spore width (SPWID).
very similar taxa, both with f brown apothecia dominat- 8. Number of spore septa (SEPTA).
ed by a K+ purple-red pigment (Bacidia Brown). There
is an obvious difference between them in thallus struc-
ture. One, which corresponds to B. polychroa, has a The difference between the continuous (B, polychroa)
continuous, smooth or warted, f rimosc thallus. The and the granular morph (B. d ~ f i a c t a ) ,excluding the
other one, named here as B. diffracta, has a finely gran- Florida specimens mentioned above, was investigated
ular thallus. In addition, there is an apparent, but not with one-way ANOVAs performed on each of these
completely discontinuous, difference in apothecium characters. The distinctness of the granular, the continu-
height. The pattern is further complicated by the exist- ous, and the Florida morphs was investigated with a
ence of a very extreme morph collected a few times in a CVA.
single locality in the Sanford area of Florida. It has very
wide and very high apothecia, and, above all, a thallus
that is not readily classified as granuIar or continuous. Results
In fact, it is partly smooth with almost no cracks at all,
and partIy granular. The question is, are B. polychroa Descriptive statistics and the significance of F in the
and B, dlf~aautadistinct species or just extreme appear- ANOVAs are presented in Tab. l l. The continuous and
ances in a continuously varying species? the granular morph differ significantly in apothecium
height and width of the paraphysis apices. None of the
 l I I l a discontinuity in the correlation between the origii~al
n characters. The first canonical variate is composed
largely by APHGT, but all other characters except
- - SEPTA contribute markedly. The contribution to the
second variate is dominated by EXCIP, but all charac-
0
ters except PARWID contribute clearly.
- - The linear function defining the axes can be extracted
a using the canonical correlation coefficients. The first
axis, CV I, is defined by the function
..
9.

- 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

where APWID is in mm, SEPTA is unitless, and the re-

maining variables are in p Note that the input values
I t
-4 I I
for APWID, HYM, and PARWID are means of five
-4 -2 0 2 4 6 measurements per specimen, for SPLEN, SPWID, and
CV I SEPTA means of ten measurements per specimen, and
Fig. 11. Canonical varintes plot of 22 specimens of Bncidia
for APHGT and EXCIP a single measurement per speci-
polychma (squares), 13 specimens of B, dzficta (circles), men.
and two specimens of a deviant population from Sanford,
Florida (triangles).
Conclusions

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

Opcra Bot. 127. 1996

Tab. 13. Descriplive statistics of quantitativc characters in Baridilin Iuerr~oremsirrom western and eastcrn North Anwrica, and from
Europe (represented by spccirncns from Sweden, Germany. Swit~erland,and Great Britain). Thc abbreviations of the characters
are explained in the tcxt. The standtird deviation is abbreviated "S. d." The input valucs uscd for APWID: HYM, and PAR are
means of five measurements per specimen, for SFLEN, SPWID, and SEPTA meails uf 10 measurements per specimen, and for
EXClP a singIe measurement per specimen. Thus, thc valucs given for all but llie latter character correspond to thc tninirnu~nand
nraxiinum specimen means in thc subspecies descriptions.
p tastern North Amcrica (N= 21) Europe (S=X)
Chal.actcl. mean S. d. min-max mcan S. d, min-max rncan S. d, min-max

APWIVimm 0.8 0.1 0.7-0.9

ExClPlp~ X6 13 69-1U3
H Y Mlpm 108 10 94-125
PARWlDIpnl 1.9 0.4 1.1-2.4
SPLENivm 81 9 69-96
SPWIDipn 3.0 0.1 2.8-3.2
SBPTA 20 2 17-23

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

Eight specilne~lsfrom Europe (Sweden, Germany, Swit- Results

zcrland, and Great Britain) deposited in LD and UPS,
nine specimens from western North Ainei-ica, and 21 Descriptive statistics of the quantitative variables and
specimens from castcn~North America were used. The the distribution of the qualitative variables are presented
North American specii~lcilswere also used to prepare in Tab. 13 and 14, respectively. The result of the pair-
the description of the species. Nine characters were wise comparisoiis of the distribution groups is shown in
scored or measured: Fig. 12. There are significant differences between all
three groups, but obviously all characters are overlap-
l . ThalIus structure (THALL) was scored 0 if the thallus ping. The western populations differ significantly' from
had few cracks or no cracks at all, I if the thallus was both of the other groups in having a diffcrcnt thallus
distinctly cracked. structure, apothecial margin colour, sporc lwgth, and
2. Colour of apothecial margin in upper part (MARG) sporc scptation. The eastenl group differs significanlly
was scored 0 i f it was k brown to black in most apothe- from thc other groups in havii~ga different hymeniurn
cia, 1 if it was f brown-red. height and spore width. The European group does not
3 . Apothecium width (APWID). diffcr significaiitly rrom both of the other groups in any
4. Width of propcr cxciple lateral to the hymenium (EX- character.
CIP). The PCA plot, in which the specimens havc bccn
5. Hyincniunl thickness (HYM). plotted against the first three principal components, is
6. Width of paraphysis apices (PARWID). shown in Fig. 13. The component loadings and thc
7 . Spore length (SPLEN). amount of variance explained is givcil in Tab. 15. Al-
8. Spore width (SPWID). though occupying diffcrcnt spaces in the PCA plot, all
9. Number of spore septa (SEPTA). three swarms overlap. The ovcrlap bctwcen thc easlern
 Fig. 12. A comparison of Bacidin
laupocerasi from western North
America, eastern North America,
and Europe. The abbreviations of
THALL * ** THALL ** the characters are explained in the
MARG *** MARG * text. The qualitative characters
APWID ns APm ns THALL and MARG were com-
parcd with Yates' corrected chi-
EXCIP ns
m *** HYM ns square. The ordinary significance
levels were divided by three to
PARWID ns PARWID ns compensate for the multiple com-
SEPTA ** ' SEPTA *** pmisons. Thus, * denotes 0.0033<
SPLEN ** ~50.017, ** O.O0033<p50.0033,
*** p50.00033, and 11s p> 0.01 7.
MARG is invariable (abbreviated
"inv" in the figure) within both the
European and the eastern group,
and hence the difference could not
be tested. The remaining charac-
ters are quantitative and were ex-
amined with one-way AWOVAs,
using the Scheffk's S procedure
for post-hoc inultiple mean com-
MARG inv SEPTA ns parisons. The significance levels
APWID ns SPLEN ns are dcnoted * for 0 . 0 1 ~p 50.05,
EXCP ns SPWID ** ** for 0.001< p 50.01, *** for
HYM ** p20.001, and ns (not significant)
for p> 0.05.

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.

1 . Apothecium width (APWTD).

2. Apothecium height (APHGT).
Bacidia heterochroa 3. Width of proper exciple lateral to the hymenium (EX-
CIF).
4. Hynlcnium thickness (HYM).
Background 5 . Width of paraphysis apices (PARWID).
6. Spore lengtl~(SPLEN).
In North America, Racidiu heterochruu occurs within 7 . Spore width (SPWID).
two different areas: along the coast in California and 8. Number of spore septa (SEPTA).
Oregon, and in the Gulf States (with a few outliers
along the coast north to North Carolina). It would be in-
correct to assume that this is a disjunct distribution,
Tab. 15. Loadings for the five characters on the first three
however. Bacidia hetemchroa is mainly a tropical spe- principal components ( X 100) in a PCA performed lo
cies, and it is apparently fairly cornmoll in the neotrop- investigate the differences bctwccn population groups of
ics. Thus, there is a distinct possibility that the North Bacidia laurocerasi with different geographic origins
American occurrences are northerIy extensions of one, (western and eastern North America, and Ellrope). The sign
continuous distribution. This, however, is yet to be pro- dcnotcs whcther the character makes a positiue or negative
ven. contribution to thc principal component. The variance
explained (in X)by cach component is given.
The eastern populations of Bacidia heterockroa were
found to consistently produce atranorin, whereas the Principal component
western ones contain no detectable amounts of any sec- Character I I[ IIT
ondary substance. Thus, a search for additional differ-
ences in morphology was carried out. THALL
MARG
APWID
HYM
Methods PARW ID
SPLEN
Fifteen spccimcns rcprescnting the castci-n populatioil SP!YID
group and eight specimens representing the western SEPTA
population group were used in this study (the same ones
Variance
later used to prepare the description of the species). explained
Eight characters were measured:

Opcr.? Bor 127, 199h

Fig. 14. Principal components plots of 23 specimens of Bacidin hetemchroa from western (squares) and southeastern United
States (circles). The first three components account for 90 % 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.

Western North America m= 8) Southeastern North America W=15)

Character mean s, d. min-max mean S. d. min-max Significance of F

APWIDlmm
APHGTJpn
EXCIPJp
HY M / p
PARWID/p
SPLEMlp
SPWIDlpm
SEPTA

Opera Bot. 127, 1998

Conclusions Tab. 17. Loadings for the five characters on the first three
principal components (X 100) in a PCA performed to
investigate the difference between the western and the eastern
It is obvious that the western and eastern groups, in ad- populations groups of Bncidia heterochron. The sign denotes
dition to chemistry, differ slightly in a few morphologi- whether the character makes a positive or negative
cal traits, and that these differences are correlated. Al- contribution to the principal component. The variance
though there are differences, the morphological explained (in %) by each component is given.
variation is completely continuous. Normally, I would
Principal component
argue that a number of morphological differences in ad-
Character I I1 I11
dition to a discontinuous chemical pattern would suffice
for taxonomic recognition at some level. In this case, APHGT 62 -3 2 -65
however, one should remember that the species is main- RYM 64 55 -40
ly distributed outside the study area, and that the differ- SPLEN 88 12 30
ences are fairly small. As long as the chemistry and SPWID 44 -83 I0
morphology of tropical representatives have not been SEPTA 84 13 41
studied, no well-founded taxonomic conclusion can be Variance
drawn. In particular, Central American B. heterochroa explained SO 23 17
should be studied carefully. It is important to establish
whether or not the western North American distribution
area is really continuous with the remainder of the spe-
cies.
Taxonomy variable than single measurements, they have also been
used in the key, although in a simplified manner.
A guide to the use of the key and the "Spores (average of 10 or more) 1.0-1.4 pm wide" is
descriptions such an example (step 27). In the three species that key
out under this alternative, a single measurement of
The aiin of this section is to gather most information spore width may vary from 0.9 to 1.9 pm, whereas, as
needed to correctly understand the structure and the can be seen, an average of ten measurements is much
content of the keys and the descriptions. Some of the less variable.
content below is repeated from the chapters Material In a few cases, particularly when N is low, the mini-
and Meah0d.s and Mbrphology and anatomy, although mum/maximum value observed may be equal to the
in an abridged version. IowesUhighest specimen arithmetic mean. This happens
when all the n observations are equal to the maximud
Arorneaclature. The synonyiny has been kept short. I minimum value. This is an unlikely event but neverthe-
have not necessarily treated all combinations based on a less it occurs. In other cases, the value provided for the
certain epithet, usually only those thought to be impor- arithmetic mean of all observations is the same as the
tant to the nomenclature. Generally, only the basionym lowest or highest specimen mean observed, even when
and the name in BacidiulBacidina have been listed, but N> 1. These cases are always due to rounding; the arith-
in case the basionyrn is below species level, the earliest metic mean of all observations is never exactly equal to
combination at the species level has been added. A few the highest or lowest specimen arithmetic mean when
exceptions from these rules have been made in noinen- N> l .
claturally problematic cases. Unless otherwise explicitly Preparations. Observations on apothecial anatomy
stated, types have been seen by me. should be made on hand-cut sections mounted in pure
The source of the descriptions. The descriptions are water. One or a few median sections should be kept in a
based exclusively on material froin the continental Unit- separate preparation. This preparation is used to study
ed States and Canada. In some cases, however, variation all apothecial characters except pigment reactions, crys-
in European or South American material is discussed, tal solubility, and paraphysis width. The reamaining
but such observations have never been included in the sections should be distributed among at least two addi-
descriptions. tional preparations, which are primarily used to study
Chamcterizations. The characterizations contain the pigment reactions with K, N, and sometimes also C (in
minimum amount of information needed to identify the which case a fourth preparation is needed). Never at-
species. The information included is not necessarily tempt to use more than one reagent on a single prepara-
consistent, since species are variously difficult to identi- tion! Note that reactions with K are stable, whereas re-
fy and to distinguish from other, confusingly similar actions with N and C will soon fade. Once the reaction
species. with K has been observed, this preparation is squashed
Measurement accoun&. Measurements are generally and the paraphyses subsequently measured. The squash-
given as "(minimum value observed -) lowest specimen ing is necessary to properly separate the paraphyses
mean observed - arithmetic mean of all observations - from each other. All measures of paraphyses in the keys
highest specimen mean observed (- maximum value ob- and descriptions refer to squash preparations in K!
served)". The standard deviation of all observations is de- Since the paraphyses may swell slightly in K, it is advis-
noted "S". "W' is the number of specimens studied, and able not to measure them in pure water.
"n" is the number of observations per specimen, the total Apotheciu. Only apothecia old enough to expose a
number of observations thus being N X n. In some char- distinct hymenium should be measured. Thus, it is pos-
acters, only one measurement per specimen has been sible to obtain a lower limit to the variation of the
made (n= l), and hence, specilneil means are identical to apothecium diameter. In apothecia that are not perfectly
the maximum and minimum values observed. In some circular, the "diameter" should be interpreted as the
species, only one specimen has been available for meas- maximum length.
urements (N= l), with the result that specimen means are Proper exciple. The width of the proper exciple is un-
identical to the mean of all observations. In cases like derstood here as the lateral thickness of the proper exci-
these, measurements are accounted for in basically the ple adjacent to the hyrnenium (Fig. 15A). lt should be
same way, but the specimen means have been omitted: measured at about half the height of the hymenium, in
"(minimum vaIue observed -) mean of all observations (- the direction (tangent) of the surface of the hymenium
maximum value observed)". Paraphysis width in mid-hy- near the proper exciple.
meniurn and pycnidial and conidial sizes are accounted An artificial subdivision of the proper exciple has
for in the form "minimum value observed - maximum been used in order to describe the pigmentation, pres-
value observed". Since specimen means are much less ence of crystals etc. (Fig. 15A): (1) Rim. The outermost

Open Bm 127. 1996

few cell laycrs. (2) Luterul interior. Thc part inside the
rim, but still lateral to the hylneniuin and the hy-
pothecium. (3) Mecbrllury part. Thc rcnlainder of the
propcr cxciple inside the rim, i.e., thc oldest parts that
arc coilstricted below the hypothccium.
C~~srt~rls.Crystals in the propcr cxcipie and in the up-
per part of the hymenium can usually easily be seen at
high magnification in an ordinary light microscope.
Should there bc sonle hesitation, use two polarizing fii-
ters, one abovc and one below the preparation. Angle
them as tu minitnize the amount of light that passes
through the preparation, and the crystals will appear as Fig. 15. Terminology used for thc divisions of apothccia and
brilliant white against a dark background. asci. - A. Schc~naticmcdiatl scction through a n apothecium
Hy~nenium.The thickness of the hylneniutl~is meas- showing the hymenium (ttbbreviated "hym"), hypothecium
ured as the distance from the basal septuin of the ascus ("hyp"), and tlie artificial subdivision of the proper exciple
to the upper surface of the apothecium. used to describe the presence o f pigments and crystals: rim
("r"), lateral interior ("l"), and tnedi~llarypart ("m"). The
Tholus. Routine investigations of the structure of thc
double arrow indicatcs how the width of the proper cxciplc
ascus apex arc bcst carried out on geiitly squashed hand- should be measured. - B. Ascus apcx showing Ihe layers of
cut sections staincd with a 0.3 % aqueous solution of io- t l ~ ct h o l ~ ~accordii~g
s tu Bclleinirc & Letrouit-Galinou 1988
dine, following pretreatment with K. Be sure nut to (a l.b, c , d).
study the tholus of asci with a ruptured wall. The de-
nomination of the layers of the tholus follows Belle-
mere & Letrouit-tialinou (1988). The a+b-layer forms lcnt divisiol~(parts of Canada), followed by year of col-
the exoascus, the c-layer thc cxternal part of the en- lcction, collector, collection number, and herbarium. On
doascus, and the d-layer the internal part of the en- inany labels, county/parish has not been specified. In
doascus (Fig. 15B). In Ruci~liaand Bacidina, thc a+b- that case, the name of a village, town, city, lake, moun-
layer is strongIy a~nyloidand the c-layer only slightly tain, or equivalent used on the label has bccn citcd. The
amyloid. Thc d-layer is basically strongly amyloid, but only exception from this n ~ l eare the Minnesota locali-
in most cases it contains a much less amyluid axial ties of Fink, which are generally not given with coun-
body. ties. On the labels of the Fink collcctioi~sin MIN, how-
Spo~-c.s.Only spores that havc bccn released from tlie ever, county infonnation has been added by Clifford
ascus should be studied. Spore length should be meas- Wetmore, and I lravc chosen to use this iilfonnation
ured as the distance between thc cnds o r the spore, Erre- here. Fink's Minnesota collecting sites have been dis-
spective of curvaturc. Thc width should be measured at cussed by Wctillorc (1978).
the widest part of the spore. Mixed collections are fairly common. All admix-
Conidia. Length and width of conidia should bc tures determined and registered by Inc havc bccn list-
nlcasurcd the same way as spores. ed. T admit, however, to having been inconsistent.
Pigmentntion. How the colour of a pig~ncntsis pcr- There are many collections with s~nallad~nixturcsof
ceived is to some degree dependcnt on the thickness of poorly developed specirncils of Bacidia and Bauidinu
the section used. The colours as dcsciibed here refer to that 1 have not bothered to study. In the collectio~l
fairly thin, hand-cut sections (c. 30-60 p n thick). Sever- lists, specimem that occur as admixtures in collections
a! pigments react with K, C, or N. The strength of the of other species arc prcceded by a "+". The name of
reaction is proportionatc to the conceulration or the pig- the species under which the collection is filed is pre-
ment. Thus, it is an advantage to select the darkest ceded by "filed with".
apothecium possible for study. Pigmentation should he Distribnrtiun niups. A single symbol on a distribution
studied in transinitted light i l l a Iight micrnscope with inap may iAepresenlone or more nearby localities.
an (almost) nlaximally opened aperture. Closing the ap- Culuur- yhatu.~.Colour photos of hand-cut apothecial
erlure will increase the contrast bctwccn tissues with sections of all species with distinct internal pigmenta-
different properties, which results in a blurring of the tion are provided (Figs 40A-43E). as are habitus photos
pigmentation. Preferrably, thc source of light shouId be of the nine ilewly described species (Fig. 43F-44F).
of near day-light quality.
Spccinren citution. All specinlens used in this study
are cited, although in a veiy abridged form. The follow-
ing information has been included, to the extent availa-
ble from the labcls: county (comte), parish, nr equiva-
Keys 7.Thallus of grey, placoid squamules that are adpressed to
a black, episubstratal, felty pmthalIus. Not sorediate....
....................................................... "Bacidia" razigustinii
Key to taxa that can be confused with Bacidia
7 . Prothallus indistinct or lacking. Often sorediate .......... 8
and Bacidina
This key contains species, species groups, and genera 8. Thallus with a para- or prosoplectenchymatic cortex,
that are easily confused with Bacidia s, str. and Bacidi- without epinecral layer. Apothecia greyish, bluish, or
nu. It has been designed to work for North American brownish to almost black, with algal cells below hy-
conditions, and includes corticolous or lignicolous taxa pothecium ..................................... Waynea cal~or~ica
that possess a crustose or squarnulose thallus, disc- 8. ThaIlus with a stainable layer and an epinecral layer.
shaped, biatorine (or appearing so), lecideine, or zeorine Apothecia either brown or black, without algal cells
(then with a well-developed proper exciple) apothecia, below hypothecium ................................ Hypocenornyce
and spores with three or more transverse septa but no
longitudinal ones. This key is presented as a service to 9. Apothecia bright red, K-+blue ... Ophioparma ~ubricosa
the reader, since it is often as difficult to determine 9 . Apothecia differently pigmented, not K+ blue ........... 10
whether or not a specimen belongs in BucidialBacidina,
as it is to determine the species. I have chosen to in- 10. Asci polysporous, clavate with a 4 pronounced swell-
clude several genera that are very distantly related to ing at the apex. A+b-layer of tholus thickened tenni-
Bacidia and Bacidina. For example, species of Bactro- nally ...........................
.. ...............................................
spora and Lmospora are commoniy mistaken for Ba- . Ropalospora chlomntha (and A. virirlis, when fertile)

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

Open Bot. 127, 1996

16. Tholus with a -t cylindrical axial body that reaches all 24. Apothecia pale yellow to yolk-yellow, soon becoming
through the d-layer (see, e.g., Hafellner 1984: 340) ... convex. Spores I-3-septate. Epithecium inspersed with
.................................................................................... 17 crystals ....................................................................... 25
16. Thoius with a k conical (or rarely lens-shaped) axial 24. Without this combination of characters .....................26
body that usually does not reach all through the d-lay-
er, at least not in all asci (see, e.g., Hafellner 1984: 25. Apothecia pale yellow. Spores consistently 3-septate ...
26 1-268) ................................................................. 20 ......................................................... "Bacidia" pallens
25. Apothecia yolk-yellow. Spores predominantly I-sep-
.I
17. Paraphyses abundantly branched, some terminating take, but occasionally up to 3-septate .............................
in swollen apices that'are provided with darkly pig- ....................... . . .....................Cliostomum vitellinum
mented caps. Apothecia often shiny ...........................
............................................................. Scoliciosporum 26. Spores often with more than 5 septa. Apothecia not
17. Paraphyses moderately or not at all branched; apices with "dotted" pigmentation in a wet condition. Excipu-
sometimes pignented but not with distinct caps. lar hyphae usually not club-shaped, with narrowly cy-
Apothecia matt ............................................................ 18 lindrical to almost globose cell lumina ..........................
.......................... Bacidia and Bacidina, continue below
18. Hypothecjum brown. Spores with both ends pointed .... 26. Spores with up to 5 septa. Epithecial pigmentation une-
............................................................ "Bacidia" jacobi venly distributed, apothecial disc thus appearing "dot-
18. Hypothecium colourless or yellowish. Spores with at ted" in a wet condition. Excipular hyphae club-shaped,
least one blunt end ................................ . . .............. 19 i.e., narrowly cylindrical in interior of proper exciple
but gradually becoming thicker towards rim ............. 27
19. Apothecia in transection, if at all, faintly and diffusely
pigmented. At least some oil droplets present in hy- 27. Length of spores not exceeding 6 times the width ........
pothecium ...................... The "Bacidia" lutescens group ......................................................... Lecania naegelii
19. Apothecia in transection usually at least partly distinct- 27. Length of spores exceeding 7 times the width ...............
ly pigmented (& greenish). Without oil droplets ............ ........................................................ Lecania stigmaiellu
................................... The "Bacidia" beckhamii p u p

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

Opera Bm. 127. 19%

38, Apothecia variously coloured (but usually not brown- 45. At least some young apothecia with thick white pmina
orange to red-brown). At least some apothecia with on the edge. Spores > 3 p wide .........17. B. mutabilis
white pruina, particularly on the edge. Proper exciple 45. Apothecia without pruina. Spores < 3 pm wide ........46
with a distinct, 4-6 cell layers thick zone of cells with
enlarged lumina along the rim. Eastern ......................... 46. Hyrnenium (average of 5) at least 75 pm high. Most
.................................................................. 26. B, sufisa spores at least 30 times Ionger than wide. Proper exci-
38. Apothecia brown-orange to red-brown, without pruina. ple (outermost 1-2 cell layers excepted) with narrowIy
Proper exciple with cell lumina gradually enlarging to- cylindrical cell lumina only. ................ 23. B. salmoneu
wards edge, or with a f distinct, 6-8 cell layers thick 46. Hymenium (average of S) 70 pm or lower. Most spores
zone of cells with enlarged lumina along the rim. Ex- less than 25 times as long as wide. Proper exciple even
treme South ........................................... 22. B. rwseola in inner part with at least some broadly cylindrical, ir-
regular, or almost globose lumina ..............................
39. Orange to brown pigment of the apothecia K+ green-
brown ............................................... 3 1 . Bn. cal$omica 47. Thallus thin, entireIy dissolved into granular gonio-
39. Apothecium pigments K-, K+ intensifying, or K+ pur- cysts 35-57 pm wide (average of 10) .............................
plish ............................................................................. 40 ........................................................ 3 5. Bn. egenuloidea
47. ThalIus usually not at all or only partly dissolved into
40. Upper part of hymenium and at least upper part of granular goniocysts. If entirely dissolved, then gonio-
proper exciple with a f distinct, dark brown, K+ pur- cysts less than 30 pm wide (average of 10) ............... 48
plish, continuous layer of pigment ............................. 41
40. Upper part of hymenium diffusely and usually weakly 48. Young apothecia often with a thin thalline margin (visi-
pigmented, pale orange to f brown, never with a dis- ble at least in section). Axial body lens-shaped. West-
tinct and continuous layer of a brown, K+ purplish pig- ern ............................................................ 36. Bn. ratnea
ment ...........................................................................
45 48. Apothecia without thalline margin. Axial body conical
to subcylindrical. Central and southeastern U. S. ...... 49
41. Spores (average of 10) 1 6 2 7 pm long, usually strong-
ly curved or sigmoid ............................ 27. B. vermifera 49. Hymenium c. 70 pm high. Pigmented park of apothe-
41. Spores (average of 10) exceeding 35 pm, straight or f cia K+ intensifying. Conidia curved, non-septate .........
cuwed to s i p o i d ....................................................... 42 ..............................................................30. Bn. assulata
49. Hymenium rarely exceeding 60 pm. Pigmented parts of
42. Epithecial pigment forming f distinct grey-brown apothecia K+ purplish or K+ intensifying. Conidia
"caps" over the apices of the paraphyses (best seen by curved or sigmoid, usually 3-1 I-septate .. 38. Bn. voria
squashing strongly pigmcntcd part of apothecium in K)
.......................................................... l I . B. heterochroa
42. Most of the epithecial pigment dissolved in the gelati- Bacidia De Not.
nous matrix or as irregular granules between the apices
of the paraphyses ........................................................ 43 G. Bot. ital. 2: 189 (1846). - Type: B. rosella (Pers.) De
Not. {lectotype selected by Fink 1910: 87).
43. Spores strongly conglutinated, f helically twisted, of- Psorella Miill. Arg., Bull. Herb. Boissier 2, appendix
ten released in packages of 8. Hymenial pigment con- l : 11 (1 894). - Type: P. pannaroidea (C. Knight) Miill.
centrated to the gelatinous matrix surrounding the as- Arg. (lectotype selected by Clements & Shear 1931:
cus apices. Southeastern ................... 10. B. helicospora 319).
43. Spores not conglutinated, although sometimes coiled Toniniopsis Frey, Ber. schweiz. bot. Ges. 35: 73
prior to the release. Hymenial pigment evenly distribut- (1926). - Type: T. obseura Frey (holotype).
ed in upper part. Western and Great Lakes-northern Bacidionayces Cif. & Tomas., Atti 1st. bot. Univ. Lab.
Appalachian ................................................................
44 crittogam. Pavia, ser. 5, 10: 39 & 65 (1953). - Type: B.
rubella (Hoffm.) Cif. & Tomas.
44. Thallus usually rather thick, grey-green, finely areolate.
Margin of at least young apothecia paler than disc, usu- Lichenized species. Thallus crustose, episubstratal, of-
ally f carmine red ........................................................... ten continuous, without cracks or rimose to areolate to
.......................... 15a. B. laurocerasi subsp, idahoensis granular, smooth to wrinkled to warted to subsquamu-
44. Thallus usually thin, t. grey, rarely areolate. Margin of lose, sometimes discontinuous and of discrete to contig-
apothecia f concolorous with disc, f brown to black. uous areoles. Prothallus lacking or present, then either
Great Lakes-northern Appalachian ................................ thin, f endosubstratal, white to grey, between discrete
.......................... 15b. B, laumcerasi subsp. laurocemsi areoles or bordering the thallus, or distinct, dark brown

Opcra Bot. 127, 1996

60
to black, bordering the thallus. Soredia and isidia ab- 66 times as Iong as wide, usually easily separated from
sent. Goniocysts absent. Upper cortex of one inner layer each other at maturity.
of anticlinal or irregular hyphae stainable in LCB and an Pycnidia usually f immersed in thallus or prothal1us
outer epinecral layer (one or the other may be missing), (rarely sessile), usually concolorous with apothecia,
both usually with minute crystals soluble in K but insol- unilocular. Conidiogenous cells lining the pycnidial
uble in N and H,SO,. Photobiont layer 4 continuous, cavity, cylindrical or slightly tapering, 7-15 X 1.O-2.5
photobiont chlorococcoid. Medulla absent or (in thick pm, producing conidia apicalIy, lacking distinct conidi-
areoles) present, rather thin, composed of loosely inter- ophores. Conidia of five different types (one to three
woven hyphae, without crystals. Lower cortex absent. types in one species; usually only one type, rarely two
Apothecia f sessile, biatorine or lecideine, when ma- types mixed in the same pycnidium): (l) filifom, f
ture 0.1-2.0 mm wide, paIe pink or pale yellow to or- curved, non-septate, 5-32 X 0.5-0.8(-1.0) pm, (2) fal-
ange to brown to brown-violet to red-brown to purple- cate, O-l-septate, 6-25 X l L 1 . 6 pm, (3) oblong to ellip-
brown to reddish or bluish black or pure black; pruina soid to fusiform to bacillifom, O-l(-2)-septate, 3-12 X
absent or present, white. Disc plane or slightly concave 1.2-2.0 pm, (4) filifom, straight, non-septate, 6-17 X
when young, often soon becoming + convex. Margin 0.5-1.0 p,and (5) filifom, slightly curved, distinctly
usually slightly raised above disc in young apothecia, 5-9-septate, 27-35 X 1.2-1.6 pm.
later level with the disc, often becoming excluded in old Chemistry: No lichen substances or sometimes atra-
and convex apothecia. norin, zeorin, and rarely meta-depsides in the orcinol se-
Proper exciple without thick gelatinous layer along ries in the thallus. ThaIlus K- or K+ yellow, C-, KC- or
rim, without oil droplets, sometimes with crystals (c. KC+ yelow, PD-, UV-.
0.5-10 p)that are either evenly dispersed or form radi-
ating clusters; composed of radiating, more or less Discussion. The generic description above is based on
abundantly furcate but sparingly or not at all anastomo- my observations on the North American species and
sed hyphae. Cell lumina usually narrowly cylindrical, must be viewed upon as tentative as Iong as Bacidiu has
usually 8-25 pm long and 1-3 pm wide (lumina near rim not been studied worldwide.
of proper exciple excepted), usually 5- 1 1 times as long The type of Psore-ella ppannaroideu (syntype in
as wide, often distinctly constricted near septa (not in WELT!),the type species of Psomlia, fits well within the
species with very narrow lumina). Walls thick and gelat- concept of Bacidia S, sir, used here. Already Brako
inized, the distance between two adjacent cell lumina at (1989) remarked that Psorelia should probably be syno-
least the same as the width of a lumen, often twice the nyinized with Bacidia. Toniniopsis obscura (type speci-
distance, and sometimes even up to five times the width mens in BEW!) is a younger taxonomic synonym of Ba-
of a lumen. Cell lumina along the rim of the proper ex- cidia illudem (Nyl.) Lynge, a species of Bacidia S. str.
ciple often f enlarged in relation to the inner lumina,
forming a distinct zone, although sometimes gradually
enlarging towards the rim of the proper exciple. Hy- 1. Bacidia absistens (Nyl.) Arnold
pothecium colourless to yellowish to brown-orange to
dark brown, without crystals or oil droplets, not chon- Flora, Jena 53: 472 (1870). - Lecidea absistens Nyl.,
droid. Hymenium 40-130 pm high, amyloid, usually Flora, Jena 52: 295 (1869). - Type: France, "Brest.",
colourless in Iower part and pigmented in a distinct or 1868, Crouan (H-NYL 17737b, lectotype selected here).
diffuse layer in the upper part, sometimes with a layer of
crystals in the upper part, without oil droplets. Paraphy- Ilh.strations. Figs 16 (map), 40A.
ses unbranched or f branched in upper part, often also f
anastomosed; apices not at all swollen to -1- clavate, of- Ch~racterization. Apothecia blue-black to black, al-
ten surrounded by pigment or sometimes containing pig- though soinetimes f pigment deficient. Proper exciple
ment in the wall, never with a thick and gelatinized laterally with brown and green pigmentation, with even-
walI. Asci clavate, surrounded by a thin, amyloid gelati- ly dispersed, minute crystals. Hypothecium colourless to
nous sheet; tholus amyloid, with a narrow c-layer, with brown. Upper part of hymeniwn greenish, rarely dirty
a 1 high, blunt or pointed axial body, the width of which purplish. Spores acicular, 40-90 X 1.9-5.0 pm, with 3-
is a third or less of the entire d-layer, and a low or high, 15 septa.
blunt or pointed ocular chamber. Spores eight per ascus,
acicular, clavate, bacilliform, or hsiform, with 3-28 Thallus indeterminate, thin to rather thick, usually dis-
septa, straight, curved, or sigmoid, straight or coiled in continuous, of discrete or contiguous, convex, often
ascus (often variable even within the same apothecium), slightly effigurate areoles that are sometimes warted or
colourless, rather thick-walled, without perispore or or- subsquamulose; sometimes continuous, f cracked, wart-
namentation, 1 1-108 pm long and 1.2-7.8 pm wide, 3- ed, pale grey, yellow-grey, 1 green-grey, or grey-green.

Opera Bot. 127, 1996 61

 a f distinct, 2-1 1 cell layers thick zone of enlarged cells
with lu~ninathat are up to 20 X h m. Lateral interior f
brown, downwards fading to colourless. Medullary part
colourless. Hypothccium colourless, pale yellow, or k
brown. Hymenium (68-j74-86- 112(-116) pn thick (S=
13, N= 8, n= 5). lower part colourless or palc purplish
brown, uppennost part k grccn, blue-green, or green-
blue, rarely dirty purplish or yellow-brown, usually f
distinctly delimited. Paraphyses 1.2-1.6 p wide in
mid-hymenium; apices usually f clavate, solmetimes
wcakly or not a t all swollen, (1.2-)l.X-2.4-3.3(-4.9) pm
wide (S= 0.8, N= 7, n= 10), without internal pigment.
Spores acicular, straight, curvcd, or sigmoid, not coiled
in ascus, (40-)45-58-74(-90) p11Iong (S= 13, N= 7, n=
IO), (1.9-)2.3-3.0-4.1(-5.0) pm widc (S= 0.6, N= 7, ]I=
10), (14.3-)17.5-20.0-24.5(-34.0) tinles as long as wide
(S= 3.9, N= 7 , i1=10), with (3-)6.8-1 1.4-14.3(-15) septa
(S= 3.4, N= 7 , n= 10).
Pycnidia almost entirely immersed in thallus, upper-
most part of wall concolorous with apotl~ecia,50-100
pm diam. Conidia filiform, curved, non-septaie, 7-32 X
c. 0.5 pm.
Chemistry: DiTferei~icombinatioi~sof at least three
unidentified metu-depsides in the orcinol-serics. Appar-
ently, none of these substai~cesis 4-O-incthylcrypto-
chlorophaeic acid, which was reported horn Scandina-
Fig. 16. Known North American distribution of Hncidiu vian specimens of this species (Tsnsberg et al. 1995).
ahsistens.
Pigments: Green parts of hyineniurn and proper exci-
ple K-, N+ purple, with or without a precipitate of sinall
Areoles when discrete (6 1-)88-137-174(-242) p n largc blue crystals. Proper exciple and sometimes hy-
(S= 45, N= 6, n= 10). Prothallus lacking or sometimes pothccium (brow11 parts) K+ purplish, N + brown-orange
present between discrete areoles, then thin, endophloeo- tu orange-red. - Bacidia Green in hylneniuln and proper
dal, pale grey. exciplc. Lauroccrasi Brown in proper exciple, pycnidial
Apothecia (0.3-)0.3-0.7-0.9(- 1 -4) mm diarn. (S= 0.2, wall, and sometimes hypothecium. Rubella Orange
N= h, n= 1 U), plane or moderately convex, remaining so sometitnes in hypothecium, rarely in propcr cxciplc and
or soinctirnes bccoming markedly convcx, cpruinose. hymenium.
Disc usually blue-black to pure black, somctimes grey-
brown, purple-brown, red-brown, or browilish pink, Ecology. On trunks and branches of Acer mucl-uyhyl-
sometimes mottled with different colours. Margin con- lum, Alnus ruhra, Popultrs triohucorpcr, and Urnbellu-
colorous with or paler than disc, k grey, f blue-grey, laria callfornica in shady and humid habitats.
brownish pink, grey-brown, red-brown, purple-brown,
dark brown: or black, sometimes mottled with different Distribution. Along the coast from central ~aliforniato
colours and when f darkly pigmented often paler in the southern British Columbia.- Europe.
lower part (to pale yellow-brown), raised above disc in
young apothecia, soon level with the disc, persistent or Discussiun. Bacidia absistens is similar to B. fr~esiana,
later excluded. B. inswlar-is,and B. s~lbin~onzpiu. All three species dif-
Proper excipIe laterally (43-)62(-74) pm wide (S= l l , fer froin B. absistens in lacking small, evenly dispersed
N= 8, n= l), at least in the inner pait with crystals that crystals in the interior of the proper cxciplc. 111addition,
are usually less than l pm large: sometimes up to 2 pn, B. fricsiana has shorter spores, a lower liymeniuin, and
usually abundant and evenly dispersed but sometimes a pigment deficient inner part of the proper exciple. B.
few and inconspicuous (visible at least in polarized insulari~has a proper exciple dominated by Bacidia
light). Rim f blue-green, green-brown, or f brown in Grcen even in the lower parts, spores that are generally
uppermost part, downwards f brown (seldom blue- fusifornl, thinner paraphyscs, and paraphysis apices that
green), then fading to coIourless, with cell lumina grad- often have distii~ctcaps of Uacidia Green. B. subin-
ually enlarging towards the edge, or along the edge with conzpta has a propcr cxciple which is heavily pigmented
cvcn in the lower parts (downwards changing to colour-
less in B. ahsis!ens).
Onc collection of Bmcidir~ ab.ristens (Fostcr 2888:
MIN) contains orange-brown apothecia in addition to
the ilormally pig~ncntcd,blue-black ones. In the orange-
brown apothccia: thc pigments Bacidia Green and Lau-
rocerasi Browil have bcen replaced by Rubella Orangc,
which snlnetimes occurs in small amounts in the hy-
pothecimn of ilormally pignleilted apothccia. In additlon
to Rubella Orange, small ainounls of Laurocerasi
Brown are produced in the proper exciple. 1 havc ncver
encountcrcd any specimen with orange-brown apothecia
only. In case such foi-ms are found, they can easily be
mistaken for several othcr species with normally f
brown pigmeilled apothccia. However, the presence of Fig. 17. Known North Arncrican distribulion uf Bacidiu
minute. eve~llydispersed crystals in the proper exciple, oggregut~rlu(clots) and B. corupaieu (triangles).
and enlarged cells along lhc cdgc of the proper exciple
is unique to B. uhs-i.stm.s.
vex. epruii~oseor with a very thin pale pruiila on disc
CANADA. Varacouver Is/and.
Colleciions r~-tut~~i~zed. and margin. Disc vciy variable in colour (even within
Colquitz River, 1908. probably collected by Macoun the same specimen), pink, pale grey, pale brown to grcy-
(CANL). Jordan Rivcr, 1966, Brodo 7X97B (CANL). - brown to grey-black (the latter colour in young apbt1li-
U. S. A. Califuunia. I-lumboldt Co., 1989, Ekman L656 cia only), or pale brown-yellow to yellow-brown to or-
(LD). Marin Co., 1978, Hcwlett 1096 (SFSU). Oregun. ange-brown to dark red-brown: allhough very variable
Breitenbush, 1930, A~ldtrsuil 5 19 (NY). Wa.vhington. rarely inottlcd with different colours. Margin in upper
Clallam Co., Foster 2764 (MIN), 1989, Ek~nanL730 part concolorous with or slightly paler or darker than
(LD). Jefferson Co., 1989, Ekman L725 (LD). Pacific disc, pink, pale grey to grey, pale brown to brown, palc
Co., 1989, Ekman L71 7 (LD). Port Angelcs, 1914, Fos- brown-grey to dark grey-brown, pale brown-orangc to
tcr 2888 (MIN). orange-brown to rcd-brown, or pale brown-yellow to
yellow-brown; lower part coilcolorous with upper part
2. Bacidia aggregatula Malme or (particularly in pale apothccia) darker, dark brown or
dark grey-brown; raised above disc in young apothecia,
Ark. Bot. 27A (5): 34 (1935). - Type: Brazil, "Matto so011lcvel with the disc, later bccoming excluded.
Grosso, Guia pr. Cuyabi, in silva cIara ripae fluminis Proper exciple laterally (50-)63(-69) pm wide (S= 9,
Coxipb GuassV, 1894, Maln~c268% (S, lectotype se- N= 4, n=l), along the rim with a layer of minute crystals
lccted here; UPS isolectotype). (at least in the uppcr part) that are up to c. I pm large,
usually abundant and cvcnly dispersed but soineti~nes
Illlrstr-ations. Figs 17 (map), 40B. few and incoi~spicuous (visible at least in polarized
light). Rim coIourless to brow11-orange, along the cdge
Chriructeriza~ion. Apotheciu~n colour very variablc without or with a single laycr of enlarged cells with lu-
even within thc same specimen, pink to 'cd-brown. mina that are up to 11 x 6 pm. Lateral interior brown-
Proper exciplc in the interior brown to red-black, in thc orange to red-black, somctin~eswith dirty green spots
riin wilh a layer of cvcnly dispersed, minute crystals. near the rim. Medullary part palc brown-orange to rcd-
Hypotheciurn colourlcss to red-black, pigmentation black, oftcn darker in the lower part than in the upper
merging into the proper exciplc. Upper part of hy- part. Hypotheciu~n colourless, pale brown, or pale
menium pale orange, with a layer of minute crystals. brown-orangc to dark brown-red, dow~lwardsusually
Spores acicular, 37-66 X 2.1-3.3 pm, with 3-9 septa. darkening and merging into the colour of the exciple.
Hymenium (74-)XO-84-8Y(-93) pm thick (S= j, N= 4, n=
Tl~allusdeterminate, very thin to rather thick, usually 5), lower part colourlcss, uppermost part 1pale orange,
contitluous, without cracks or k cracked, wrinkled, pale with a thin Iayer of minute crystals. Paraphyses 0.5-1.2
grey, pale yellow-grey, f green-grey, or grey-green. Pro- pm wide in mid-hymenium; apices usually k clavatc,
thallus bordering thc individual, narrow, black where it sometimes weakly or not at all swollen, (0.8-)1.7-2.2-
meets other lichens, othciwise pale grey or lacking. 2.4(-3.2) p n wide (S= 0.6, N= 4, n= 10), without inter-
Apothecia (0.5-)0.7-0.8-1.1(- 1.7) mm diam. ( S = 0.3: na1 pigrncnt. Spores acicular, straight, or slightly curved
N= 4, n= IO), plane whcn young, later becoming con- or signloid, sbaight or only slighlly coiled in ascus, (37-)44-

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

Opera B o t 127. l996

Fig. 18. Known North American distribution of Bacidia arceutha.

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,

Open Bat. 127, 19% 65

 (1855). - Lecidea effza (Auersw, ex Rabenh.) Stizenb.,
Nova Acta Acad. Caesar. Leop. Carol. 34 (2): 23
(1 867), nom. illeg., non (Pers. ex Hoffm.) Mudd 1861. -
Bucidia efisa (Auersw. ex Rabenh.) Lettau, Hedwigia
52: 132 (1912), nom. illeg., non (Sm.) Trevisan 1856. -
Lecidea uuerswaldii Hepp ex Stizenb., Ber. Tat. St Gall.
natunv. Ges. 1880/81: 416 (1882). - Bacidiu e-usellu
Zahlbr., Cat. Lich. Univ. 4: 110 (1926), nom. illeg. (su-
perfluous). - Type: Germany, "Leipzig, an einer a l t ~ n
UIme irn Rosenthale, die Rinde m e h e EIlen weit uber-
ziehend", 1855, Auerswald, Rabenhorst: Lich. Eur. exs.
32 (W, lectotype selected here).

Figs 6E, 19 (map), 40D.

J~~ustratiom.

Chamcterization. Thallus finely granular; granules 36-

Fig. 19. Known North American distribution of Baciriin 145 pm diam. Apothecia normally purple-brown to
auerswaldii (dots) and B. igniarii (triangle). black, but often f pigment deficient. Proper excipIe Iat-
erally orange to black-brown. Hypothecium (almost)
colourless. Upper part of hymenium orange to black-
1966, Brodo 7897a (CANL, COLO, WIS), + Vancouver brown. Spores hsiform with blunt ends, 19-33 X 5.6-
Island, Mount Douglas Park, 1989, Eknlan L779 (filed 7.8 pm, with 3-9 septa.
with Bn. ramea, LD).Mayne Island, 1980, Noble 7277
(herb. Noble). Salt Spring Island, 1989, Ekman L782,
L784, L787, L789 (LD). - U. S. A. California. Rum-
Thallus indeterminate, thin, of discrete or contiguous,+
globose or irregular, often slightIy flattened granules,
boldt Co., 1989, Ekman L659 (LD). Mendocino Co., pale grey, pale yellowish grey, pale green to green, or f
1989, Ekrnan L651 (LD), + 1982, Thiers 45648 (filed grey-green to brown-green. Granules (36-)63-72-82(-
with B. heterochroa, SFSU). San Mateo Co., 1967, Jor- 145) pm diarn. (S= 22, N= 3, n= 10), without crystals in
dan 818 (SFSU). Santa Cruz MountaiAs, 1885, probably the cortex. Prothallus sometimes present between dis-
collected by Herre (FH). Maine. Rockport, 1921, Mer- crete granules, very thin, eridophloeodal, pale grey.
rill (FH). South Thornaston, 19 10, Merrill (FH). Massra- Apothecia (0.3-)0.4-0.5-0.6(- 1.1) mm diam. (S= 0.2,
checsetts. Dartmouth, 1888, Willey (US). Fairhaven, N= 2, n= 10), at first plane, remaining so or becoming
1887, Willey (US), New Bedford, Wiiley (FH, US). moderately convex, epminose. Disc brown-orange, or-
New York. Suffolk Co., 1961, Brodo 3209 (MSC). Ore- ange-brown to red-brown to purple-brown to black, in
gon. Benton Co., 1972, Pike 2777, 3117 (OSC), 1989, some apothecia mottled with different colours. Margin
Ekman L638, L682, L683, L684, L686, L689, L696, in upper part concolorous with or slightly darker than
L697, L702, L703 (LD), + 1989, Ekman L688, L691, disc, in lower part commonly distinctly paler, pale pink
L692, L701, L704 (filed with Bpl. mmea, LD). Cuny to orange-brown, distinct, raised above disc in young
Co., 1989, Ekrnan L670, L674, L675 (LD), + 1989, Ek- apothecia, later level with the disc, persistent.
man L673 (filed with B. sa~monea,LD). Lincoln Co., Proper exciple laterally (43-)59(-74) pm wide (S= 15,
1972, Denison (OSC), 1952, Weber S1140 (COLO). N= 3, n= l), without crystals. Rim pale orange to
Washington. Clallam Co., 1989, Ekman L731 (LD), brown-orange to red-brown to black-brown in upper
1969, Brodo 15335 (CANL). Gray's Harbor Co., 1989, part, downwards gradually changing to colouxless,
Ekrnan L718, L719 (LD). Pacific Co., 1989, Ekman along the edge with a single layer of enlarged cells with
L71 5, L71 6 {LD). Skagit Co., 1989, Ekman L734, L738 lumina that are up to 9 X 5 pm. Lateral interior +_ con-
(LD). Whatcom Co., 1914, Herre (FH), 1980, 1982, Ry- colorous with or slightly paler than rim. Medullary part
an (WWB), 1982, Khoades (WWB), 1989, Ekmm colourless. Hypothecium colourless or very pale yellow-,
L740, L741, L742, L745, L749, L 750 (LD). + Jefferson ish. Hyrnenium (74-)80-95-110(-114) pm thick (S= 13,
Co., 1989, Ekman L727 (filed with B. salmonea, LD). N= 3, n= 5), colourless in lower part, pale orange to or-
ange-brown to red-brown to black-brown in a distinct or
diffuse layer in the upper part. Paraphyses 1.0-1.6 pm
4. Eacidia auerswaldii (Hepp ex Stizenb.) Mig. wide in mid-hymenium; apices f clavate, (1. .6-)2.6-3.0-
3.2(-5.7) pm wide (S= 0.8, N= 3, n= 10), without inter-
Kryptogamen-Flora 4 (2): 267 (1931). - Biliinbia $fusu nal pigment or wall with f brown or grey-brown pig-
Auersw. ex Rabenh., Rabenhorst: Lich. Eur. exs. 32 ment. Spores fusiform with bIunt ends, straight, not
coiled in ascus, (19-)23-28-31(-33) pm long (S= 4, N=
3 , n= 10), (5.6-)6.5-6.8-7.0(-7.8) pm wide (S= 0.6, N=
3, n= IO), (2.9-)3.6-4.1-4.4(-5.1) tiines as long as wide
(S= 0.6. N= 3, n= 10). with (3-j5.7-6.1-h.4(-9) septa (S=
1.6, N= 3, n= 10).
Pycnidia not seen.
Chemistry: No lichen substances detected.
Pigments: Uppcr part of hymcniu~nand propcr cxci-
ple pxily K+ intensifying and N+ intensifying, partly
K1- purplish and N+ orange-red, or entirely K+ purplish
and N+ orange-red. Hypothecium K+ intensifying, N+
intensifying. - Laurocerasi Brown and Rubella Orange
in hymenium and proper exciple. Rubella Orange in hy-
pothecium (small quantities).

Ecology. There is no uniform pich~reof the ecology of

B. alter-swaldiclii, sillcc the habitats of the thrcc finds arc
vcry diffcrcnt among themselves. One find was iiladc on
a large Acer muor-ophylhma on a roadside, where it oc-
c urred together with An isurt~eridum nyssuegenlrm, Ba-
cidiu circumspeotu, and LStrungospuri~ ochruphom. An-
other locality is nn a Popullls E,-ichocarpa in a humid
forest don~inatedby Picea engelmanraii, Thuju plicata,
and Betula papvrfer-a. No othcr lichcns wcre prcsent in Fig. 20. Known North Ainericaii distril-rution of Bucidiu
the inlincdiate vicinity, but "Baciclicr" carneoalbida oc- (triangle), B. friesianu (square), and B. instllaris
bi~l~orina
curred abundantly lower down on the same trunk. In the (dots).
third locality: B. auerzl.wuidii was, according to the Ia-
bel, found on a Thijju ylicutu in a gorge with a creek at
the bottom. In this collection, B. uuerswuldii is accoin- 5. Bacidia biatorina (Korb.) Vain.
panied by "Bucirlia" Oeckhuusii and Biutur-idium del-
ite.rcens. Acta Soc. Fauna Flora fenn. 53 (1): 178 (1922). -
Rltuphiospom utrosnngwineu var. biatorinn Korb., Par-
Di.~trihution.Known from one locality in northwestern erga Lich., fasc. 3: 238 (1861). - Type: Gennany, "Wiir-
Montana and two in British Columbia (one on Salt- temberg", Kemmler (L-9 10205-1 689, lectotype selected
spring Island in thc Strait of Gcorgia and o ~ l cin Wclls here).
Gray Provincial. Park). - Europe.
fllzrstrutiuns. Figs 20 (map), 40E.
Discuxsiun. Bucidiu uuctawaliliii is easily recognized by
the very wide, htsiform and blunt-ended spores. Under Charucterizution. Thallus granular, without crystals in
the microscope, it can hardly be confused with any oth- the cortex; granules 48-121 p diam. Apothecia or-
er species. In its overall habit, it resembles B, biutorina ange-brown to dark purplish brown. Proper exciple lat-
and B. r.zlheliu. erally orange-brown to dark red-brown. Hypothecium
The affinities of B. aller.~waldiiare clearly with Ba- (almost) colourless. Upper part of hymenium brown-or-
cidia in thc strictest sensc, dcspitc the shape of the ange to red-brown, the latter pigment K+ purplish.
spores. Ii is probably closely rclatcd to B. hiaforina, Spores acicular, 42-57 X 2.1-2.9 p,with 3- 15 septa.
with which it sharcs a similar apothccium pigmentation
and thc grai~ularthallus. Thallus indeterminate, thin to rather thick, of discrete or
contiguous, globose or irregular, oftcn slightly flattened
Colleelions exautined. C A N A D A . Brilish Columbia. granules, green-grey. GiAanules(48-)85(-121) pm diam.
Saltspring Island, 1989, Ek~nanL788 (LD). Wells Gray (S= 16, N= I, n= 20), without crystals in t l ~ ecortex. Pro-
Provincial Park, 1989, Ekman L796 (LD). - U. S. A. thallus lacking.
Montana. Flathead Co., 1991, McCune 19222 (LD, Apothecia (0.4-)0.6(-0.9) mm diam. (S= 0. I , N= 1, n=
OSC). 10), remaining plane, epruinose. Disc orange-brown to
dark purplish brown, in many apothecia mottled with
diffcrcnt hucs. Margin concolorous with or sligl~tly
darker than disc, distinct, raised above disc in young Collections examined. U. S . A. California. Trinity Co.,
apothecia, later level with the disc, persistent. 1989, Ekman L661 (LD).
Proper exciple laterally 52 pm wide v= 1, n= l),
without crystals. Rim usually orange-brown to dark red-
brown in upper part (rarely colourless), downwards 6. Bacidia campalea (Tuck.) S. Ekman & Kalb,
gradually changing to colourless, along the edge with a comb. nov.
1-2 cell layers thick zone of enlarged cells with lumina
that are up to 12 X 6 p Lateral interior f concolorous Lecanora carnpalea Tuck., Amer. Joum. Sci. Arts, ser.
with rim. Medullary part co1ourIess. Hypothecium col- 2, 28: 205 (1859). - Haematornma campaleurn (Tuck.)
ourless or very pale yellowish. Hymenium (83-)84(-87) Zahlbr., Cat. Lich Univ. 5: 758 (1928). - Type: Cuba,
pm thick (s=2, N=l, n=5), colourless in lower part, 1857, Wrist (FH-TUCK, lectotype selected here).
brown-orange to red-brown in a distinct or diffuse layer Lecidea subluteolu Nyl., Flora, Jena 52: 122 (1869). -
in the upper part. Paraphyses 0.8-1.6 pm wide in mid- Bacidia subluteolu (Nyl.) Zahlbr., Sber. Akad. Wiss.
hymenium; apices f clavate, (1.6-)2.6(-4.0) p wide Wien 111 (1): 395 (1902). - Type: Brazil, 1867,
(S= 0.8, N= 1, n= 20), without internal pigment. Spores GIaziou (H-NYL 17052b, lectotype selected here).
acicular, straight, curved, or sigmoid, sometimes coiled Bacidila v ~ r i e g d aZahlbr., Denksch. Akad. Wiss.,
in ascus, (42-)49(-57) pm long (S= 4, N= 1, n= 20), Wien 83: 129 (1909). - Type: Brazil, "Prov. Sao Paulo.
(2.1-)2.5(-2.9) pm wide (S= 0.2, N= l , n= 201, (16.5-) Prope Rio Grande ad Sao Pauio Railway, 800 m, S. m.",
19.5(-23.0) times as long as wide (S= 1.8, N= l , n= 20), 1901, Schiffner (W, lectotype selected here).
with (3-)10.1(- 15) septa (S= 3.6, N= 1, n= 20). Bucidia vergarensis Rasanen, Revta sudam. Bot. 7:
Pycnidia half-immersed in thallus, pale orange, 50- 13 (1942). - Type: Uruguay, "Tr. y Tres, Vergara, corti-
130 p diam. Conidia filiform, curved, non-septate, 9- cola", 1932, Herter (H, lectotype selected here).
16 X c. 0.5 pm.
Chemistry: No lichen substances detected. IIIustrations. Figs 3A, 17 (map), 40F.
Pigments: Upper part of hymenium and proper exci-
ple K+ intensifying and NI-intensifying, or K+ purplish Characterizution. Apothecia with pink to brown to al-
and N+ orange-red to red. Hypothecium and inner parts most black disc; margin paler. Proper exciple colourIess
of proper exciple K+ intensifying, N+ intensifying. - to brown-orange to dark brown, in lower part with large
Laurocerasi Brown in hyrnenium, proper exciple, and clusters of crystals and in upper part with evenly dis-
pycnidial wall. Rubella Orange in hypothecium (very persed, minute crystals; with cell lumina gradually en-
small quantities), hymenium, proper exciple, and pyc- larging towards the edge, or along the edge with a 6-8
nidial wall. cell Iayers thick zone of enlarged celI lumina. Hy-
pathecium brown-orange to brown. Upper part of hy-
Ecology. h bark crevices of an old Lithocarpm densi- menium orangc-brown to red-brown. Spores acicular,
f i n i s in a fairly open forest (dominated by Quercus 41-71 X 2.5-3.7 pm, with 3-1 1 septa.
garvyana and Pseudotsuga menziesii) in a valley.
Thallus determinate, f thin, continuous, k cracked,
Distribution. Presently known from only one locality in wrinkled, or sometimes partly discontinuous and of dis-
the intermontane region of northern California. - Eu- crete or contiguous, convex areoles; pale bluish grey.
rope. Prothallus sometimes bordering the thallus, 'narrow and
black; sometimes present also between discrete .areoIes,
Discussion. B. rubeilu is very similar to B. biatorieu in thin, endophloeodal or epiphloeodal, pale grey.
habit, but differs in having larger thallus granules with Apothecia (0.5-)0.6-0.7-0.7(-0.9) mm d i m . (S= 0.1,
crystals in the cortex, and in always lacking the brownish, N= 3, n= 10), plane or moderately convex, when young
K+ purplish and N+ orange-red to red pigment, which is or medium-aged usually slightly pruinose on disc and
present in most or all of the apothecia of B. biatorina. margin. Disc pink, pale orange, dirty brown, dark
European specimens of B. biatorina usually have a brown, purple-brown, dark brown-violet, purple-black,
yellowish or brownish green thallus and darker, purple- or brown-black. Margin paler than disc, f pink, brown-
brown to black-brown apothecia with evenly coloured pink, pale pink-brown, f grey-violet, or pale to dark
disks. However, the green-grey thallus and the partly brown, lower part concolorous with or darker than up-
brown-orange apothecia present in the Califomian speci- per part, raised above disc in young apothecia, soon lcv-
men are sometimes found also in European specimens. el with the disc, persistent.
Furthermore, h e Califomian specimen appears to deviate Proper exciple laterally (52-)60(-68) pm wide (S= 8,
from European B. biatorina in having a thinner apothe- N= 3, n= l), in lower part with large clusters of crystals
c i d margin. (single crystals up to 8 p in diarn.) and at least in up-
per part usually with abundant and evenly dispersed purplish pigment (Laurocerasi Brown) dominating the
minute crystals that are up to 1.5 pm in diam. K m col- upper part of the hymenium, and in having wider para-
ourless or f brown-orange to dark brown, often heavier physis apices (min-max 1.6-2.8 pm). B. millegrana,
pigmented in lower part than in upper part, with cell lu- which probably is a pantropical species, differs in hav-
mina gradually enlarging towards the edge, or along the ing a distinctly yellowish thallus with bullate areoles, a
edge with a f distinct, 6-8 cell layers thick zone of en- very distinct, two cell layers thick zone of enlarged cells
larged cells with Iumina that are up to 12 X 6 p n . Later- along the edge of the proper exciple, and in lacking
al interior colourless, or brown-ormge close to the hy- crystals in the proper excipie. B. russeolu is rather simi-
menium. Medullary part colourless to pale yellow. lar to B. carnpalea, and both species share the very wide
Hypothecium brown-orange to brown. Hymenium (78-1 zone of enlarged cells along the edge of the proper exci-
84-98- 105(-112) pm thick (S= l l , N= 3, n= S), lower pie. B. msseola is, however, easily separated by the dif-
part colourless, uppermost part diffusely and unevenly ferently coloured apothecia, the lack of minute crystals
orange-brown to red-brown, with a thin layer of crystals in the proper exciple, and (usually) by the presence of a
above the apices of the paraphyses. Paraphyses 0.5-1.0 K+ purplish pigment in the proper exciple and hy-
pm wide in mid-hymenium; apices not at all swollen or menium. Furthermore, B. keterochroa may occasionally
narrowly clavate, (0.8-) 1.3-1.3-1.4(-2.0) pm wide (S= be mistaken for B. campalea. The former can be sepa-
0.3, N= 3, n= 10), without internal pigment. Spores ac- rated by the distinct, brown cap of pigment in the wall
icular, straight or slightly curved, straight or slightly of the paraphysis apices and by the complete lack of a
coiled in ascus, (41-)52-53-55(-71) pm long (S= 6, N= very wide zone of enlarged cells along the edge of the
3, n= 10), (2.5-)3.2-3.3-3.5(-3.7) pm wide (S= 0.3, N= proper exciple.
3, n= lO), (12.3-)15.3-16.0-17.1(-22.8) times as long as The types of Bacidia subluteola (Nyl.) Zahlbr., B.
wide (S= 2.3, N= 3, n= 10), with (3-)6.6-7.5-8.0(-11) varieguta Zahlbr., and B. vergaremis Eisiinen are typi-
septa (S= 1.5, N= 3, n= 10). cal, well-developed specimens of B. camptalea, and they
Pycnidia not seen. are hence reduced here into synonymy.
Chemistry: Atranorin and zeorin.
Pigments: Pigmented parts of apothecia mainly K+ Collections examined. U . S. A. Florida. Collier CO.,
intensifying, N-, but in parts K+ purplish, N+ orange- 1985, Thor 4645 (S), 1993, Ekman L1176, L1177,
red. - Rubella Orange in proper exciple, hypothecium, L1 178, L1 1.79, L1 180, L1202 (LD).
and hyrnenium. Laurocerasi Brown in hypothecium and
hymenium (small quantities).
7. Bacidia circumspecta (Nyl. ex Vain.) Malme
Ecology. Both localities are swamp fringes along roads,
where B. campalea is growing on the illuininated trees Bot. Notiser 1895: 140 (1895). - Lecidm bacii!ijera f.
closest to the road. It has been found on Acer rubrum, circumspecta Nyl., nom. nudum, Norrlin 8r Nyl: Herb.
Aanona glubm, Piscidia piscipula, and Sulix curolini- Lich. Fenn. 185 (1875). - Lecidea bacillfem var. cir-
ana. cumspectu Nyl. ex Vain., Meddn Soc. Fauna Flora fenn.
10: 22 (1883). - Lecidea circumspecta (Nyl. ex Vain.)
Distribution. Presently known from only two localities Hedl., Bih. K. svenska VetenskAkad. Handl. 18, 3 (3):
in Collier County in southernmost Florida. - Probably 71 (1892). - Type: Finland, "Tavastia australis, Lammi,
widespread in the neotropics, and extending into sub- Evo, supra corticem salicis", 1874, Norrlin, Norrlin &
tropical areas on both sides of the tropical belt. Nyl: Herb. Lich. Fenn. 185 (H, lectotype selected h,ere).

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-

Opcm 001. 127, 1896

Fig. 21. Known North American distribution of Bacidia circumpecta.

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-

Open Bot 127. 1996

septate, 7-12 X 1.2-1.9 p,(2) filiform, curved, non- cumspecta approaches B. subcii-cumspecta Coppins, de-
septate, 8-19 X 0.5-O.S(-1.0) p,and (3) falcate, 0-1- scribed froin Scotland (Coppins et aI. 1992). B. subcir-
septate, 6-25 X 1.2-1.6 p. cumspecta was separated from B. circumspecta by the
Chemistty: No lichen substances detected. filiform, curved conidia (instead of oblong conidia), the
Pigments: Upper part of hymenium K- and N+ purple dominance of a purplish pigment (Laurocerasi Brown?)
(with or without a precipitate of blue crystals), or partly in the l~ymenium(instead of Bacidia Green), the lower
K+ purplish, N+ orange-red. Proper exciple K+ pur- hymenium (30-35 pm instead of 45-55 W), the differ-
plish, N+ purplish. Hypothecium K+ intensiming or ent habitat requirement (hard lignum instead of tree-
very rarely K+ purplish. - Bacidia Green in hymeniurn, trunks). Some North American specimens of B. circurn-
proper exciple, and pycriidial wall. Laurocerasi Brown specta combine curved conidia with having an
in proper exciple, paraphysis apices, pycnidial wall, and abundance of purplish brown paraphysis apices (mixed
very rarely hypothecium. Rubella Orange occasionally with green ones), although the exact combination of
in hypothecium. characters in B. subcircumspecta is not found. Further-
more, some North American specimens are lignicolous.
Ecology. Bacidia circunaspecta is found in a wide varie- The distin'ction between B. szkbcircumspecra and the
ty of habitats, from exposed and dry to to shady and hu- most extreme of North American B, circzkmspecta is
mid ones. It has been collected on Acer grandideatutu, fairly subtle, however, and had B. subcircunaspecta oc-
A. rnacrophyllum, Betula olleghaniensis, Fruxinus spp., curred in North America, 1 am sure it would have been
Malacothamnus fascicukatus, Picea gluuca, Plutanus perceived as the extreme end of a continuous variation
occidentalis, Popzklus grandidentata, P. trernuloides, within B. circum~pectu.Nevertheless, 1 think it is cor-
@ercus agr$oiiu, Q, gurryana, Q. macromrpa, Q. ru- rect to treat B. subcircumspecaa as a separate species. It
bra, Thuja occiden~alis,T. plicaaa, Ulrnus americana, is so far known only from Scotland, where it is sympat-
and rarely on lignum (stumps, fenceposts). ric with B, circumspectu from which it remains morpho-
logically and ecologically distinct. Possibly, B. subcir-
Distribuaion. Probably more or less Panamerican, but cumspecta has evolved through a founder effect
apparently avoiding northern Canada, the extreme following the long-distance dispersal of an ancestor of
Southeast, and high altitudes in the Rocky Mountains B. circumpec~awith an extreme character combination
and the Appalachians. - Europe. from eastern North America to the British Isles.

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

Opera sot. 127. 1496

 Fig. 22. Known world
distribution of Bacidia
d~fracta.

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.

72 Opem BM 127, 19%

sour-i. + Orcgon Co., 1988, Harris 2 17 13a (filed with B. 20), (8.3-) 10.8(- 13.5) times as long as wide (S= 1.3,
poiychroa, NY). Nnr4ih Ccrrolina. + Swain Co., 1976, N= l , n= 20), with (7-)7.2(-9) septa (S= 0.6: N= l , n=
Harris 1 1009 (filcd with B. polychvnrr, NY). Pennsdva- 20).
nia. Bushkill Falls, 1937 ( N Y ) Pike Co., 1940, Thorn- Pycnidia not scen.
soil 2150 {UPS). Tennessee. Cartcr Co., 1985, Harrls Chemistry: Not investigated due to sparsity of materi-
18334 (NY). Vermont. Monkton, 1879, Farlow (NY). al.
Pigments: Upper part of hymeniunl and proper exci-
ple mainly K- and N+ purple (without a precipitate of
9. ~ a c i d i afriesiana (Hcpp) KGrb. small blue crystals). Proper cxciplc partly K+ purplisll.
N+ orange-red.- Bacidia Green in hyme~~ium, r i ~ nof
Parerga Lich., fasc 2: 133 (1 860). - Bintorcr ,#i.iesiana propcr cxciple, and sometimes also in sn~allamounts in
Hepp, Flechten Europas 288 (1 857). - Type: Switzer- I h e hypotheciu~nand the medullary part of thc propcr
land, probably near Ziirich, Hepp (BM-22207b, lecto- exciple. Laurocerasi Brown in the rim of the proper cx-
typc sclected here). ciple, and soinerii~lcsin sinall amoi~ntsin the upper part
of the hymenium.
Il1ustration.r. Figs 20 (map), 4 1 A.
Eccllogv. 011 brai~cl~es of a Su1i.x sp, which fonned a
Chamcferizutiori. Apothccia green-blue to almost scrub in an otherwise open, boggy area along a river,
black, or partly or entirely yellowish. Proper exciple f near its outlet in the sea.
grey-green in the rim, othcrwisc pale, without crystals.
Hypothecium (almost) coluurless. Uppcr part of hy- Dis!ribuiion. Known from only one locality on the coast
nlenium colourless to blue-green. Spores acicutar, 3 1-3 8 of southwestern Oregon. Previous records frorn N o ~ t h
X 2.5-3.7 p n , with 7-9 septa. America are crruncous. They are based on an unde-
scribed species of Scoliciosponrm from the coast of
Thallus indctcnninatc, very thin, discontinuous, or' dis- British Cnlmnbia. - Europe.
crete or contiguous, alnlust granular areoles, pale grey
to pale brown-grey. Areoies whcn k discrete (24-)53(- Discussion. Bacidiu ubxlstens is similar to B. ,jr~e.i~siana,
85) large (S= 1 7 , N= l , n= 20). Prothallus oftcn present but differs in having crystals in the inner of the proper
between dlscrete areoles, very ihin, cndophloeodal, pale cxciple, longer spores, a thicker hyinenium, and in hav-
grey. ing a brown pigment (Laurocerasi Brown) in the lateral
Apothecia (0.1-)0.2(-0.2) mm diam. ( S = 0.04, N= 1, interiur of thc proper excipIe.
n= 20), at first plane, later becoming convex, cpruinose.
Disc green-blue, grey-bIue, blue-black, or palc dirty Collections ~xumiraed. U. S. A. 01.eg.gun. Curry Co.,
brown-yellow, often mottled with different colours. 1989, E h a n L1218 (LD).
Margin concolorous with or paler than disc, blue-grey,
brown-grey, green-bluc, grcy-blue, blue-black, or pale
dirty brown-yellow, Iike the disc oftcn mottled with-dif- 10. Bacidia hclicospora S. Ekman, sp. nov.
ferent colours, raised abovc disc in young apothecia,
soon level with the disc, finally excludcd. Type: U. S. A., "Tcnnessee, Henderson Co., on hard-
Proper exciple laterally 5 3 ,urn wide (N= 1, n= l ) , woods i n mixed pine-hardwood forest (Cornus, Liqui-
without crystals. Rim grey-green, downwards fading dambur, Pinus eohinatn) along road to Greener Ceme-
to pale grey-green, along the edge without or with a tary, I mi. S of US 40, 8 ini. E of intersection wit1 State
single cell layer of enlarged cells with lumina that 32, near Natcllez Trace State Park", 1973. Tucker
are up to 4 X 4 pm. Lateral interlor colourless. Mcd- 11 128b (herb. S. Tucker, holotype).
ullary part colourless to pale blue-green. Hy-
potheciu~ncolourlcss to pale blue-green. H y m e i ~ i u n ~ A simili Bclcidiu Inzu.ucerasi differt sporis firme con-
(38-)47(-55) pm thiclc (S= h, N= l , n= 7), lower part glutinatis et inter se contoriis, pigmciltoque epithecii in
colourless to pale blue-grcen, uppermost part col- matriccin gelatinosam circa ascos contracto. Etiam B.
ourless to blue-green. Paraphyscs c. 1.5 p n wide in heterochr-oae si~nilisquae pigmenturn cinereo-fuscun~
mid-hymenium; apices usually f cIavatc, sometimes in apicibus paraphyson~m et sporas non conglutinatas
weakly or not at all swollen, (1.6-)2.3(-3.2) pm habet.
wide (S= 0.5, N= 1 , n= 20), wiihout intcrnal pig-
ment. Sporcs acicular, straight, curved, or sigirioid, E ~ n t o l o g )Helicu-,
~. spirally twistcd, and sl~ovlls,spore.
not coilcd in ascus, (3 1 -)34(-38) pm Iong (S= 2, N=
l , n= 20), (2.5-)3.1(-3.7) pm wide ( S = 0.3, N= l , n: Ill.ust,a!ions. Figs 7 , 23 (map), 4 1B, 4 3 G .
Chuructrr.izulion. Apothecia purple-brown to black.
Proper exciple red-brown to black-brown in thc rim,
otherwise colourless to orange-brown. Hypothcciuin f
pale yellow to pale orangc-brown. Upper par( of hy-
~neniumk brown, pigmciit accurnulaled around the up-
per part of thc asci. Spores acicular, helical and lightly
coiled, 33-81 X 2.1-3.7 pn, wit11 7-19 septa.

Thallus indeterminate or determinate, thin, usually con-

tinuous. smooth, wrinkled, or rarely warted, without
cracks or f cracked, rarelv discontinuous. o f ? discrctc,
thin areoles, light grey to greenish or yclluwish grcy.
Prothallus so~nctimcspresent as a thin, black line along
the edge of thc thallus.
Apothccia (0.4-)0.4-0.6-0.7(-1 .O) mm diam. (S= 0.1,
N= 7:n= 5), planc or slightly convex, epruinose. Disc
purple-brown to purplish black or pure black. some-
tiines rnottled with orange-brown or beige (particularly
in young apothecia). Margin concolorous wit11 disc or
darker, .purple-brown
. to black, in the lower part some-
times paler, f brownish, distinct, raised above disc in
young apothccia, soon levcl with the disc: persistent.
Prupcr cxciplc lalerally (43-)59(-70) p n wide ( S = 10, Fig. 23 Known world distribution of Hocidiu heiicr>.spura.
N= 7, n= l), without crystals. Rill1 red-brown to black-
brown in a thin distinct zone, only rarely downwards
gradually changing to pale brown, along the edge with a intensifying, N-. - Laurocerasi Brow11in hymenium and
1-2 cell layers thick zone of enlarged cclls with lumina rim of proper cxciplc. Rubclla Orangc in hypotheciu~n
up to X X 5 pm. Lateral interior paler than rim, colour- and iililer part of proper exciple.
less to yellowish to brown-orange to orange-brown or
partly inottIed with red-brown. Medulla~ypait concoior- Ecology. On trcc trunks, preferrably in shady and hunid
ous with or paler than hypotheciun~and lateral interior, habitats. Unfortunately, the labcls of thc majority of thc
colorless to pale orange-brown. liypothcciuin vcry palc speci~nenslack phorophytc data. Know11 phorophytes,
yellow to palc orangc-brown, Hynlcniu~n (74-)76-90- however, include Carpinus enroliniana and Liquidurn-
103(-105) pn thick (S= 9, N= 7 , n= j),lower part col- bar- siymuiJlzru.
ourless except for the brown-pigmented gelatinous ma-
trix surrounding the young asci: uppermost part brown Disfr-ibtltiurl. Known from Maryland, South Carolina,
to black-brown, unevenly pigmented due to the accumu- Tennessee, the Florida Panhandle, Louisiana, southeast-
lation of pigment to the upper part of thc asci. Paraphys- c m Oklaho~na,and southern Illinois. AI1 localities ex-
es 0.8-1.6 pm wide in mid-hymci~ium;apiccs usually ccpt oilc arc situatcd at low elevatioils (all below 180,
not at aIi or only slightly swollc-11, but someti~nesuar- most of them even below 100 m ) in the Coastal Plain or
rowly clavate, (1.0-j1.3-1.6-1.8(-2.8) pi1 widc ( S = 0.4, in the vallcys of Mississippi and Teilnessee Rivers. A
N= 8, n= 5), diffuscly surrounded by brow11 pigmcnt, single find was made at about 400 m abovc sca lcvkl in
~nainly near ascus apiccs. Sporcs acicuiar, hclical. thc valley of Littlc Tcilncsscc River near the south bor-
strongly coi~glutinatedand coiled in packages of X, of- der of Great Sinoky Mountain NationaI Park. - Endcm-
ten very difficult to separate eve11 when squashed in K, ic.
(33-)50-56-64(-81) pm long (S= 9, N= 20, n= S), (2.1-
)2.6-2.8-3.1(-3.7) ynl wide (S= 0.3, N= 5, n= 10): ( 1 1.7- D i ~ c ~ ~ s s iBuciJiu
on. I1~1i~o~pu1.u
is very similar in habit
)18.O-20.6-22.9(-28.3) times as long as wide (S= 3.7, and pigmentation to B. larrrucel-usi, and the spores of
N= 5, n= 10), with (7-)7.X-12.8-14.8(-19) scpta (S= 3.0, the latter may even be helically arranged in young asci.
N= 5, n= 10). I-Iowcvcr, thc sporcs of B. lauro-oce~asiare easily sepa-
Pycnidia not seen. rated at maturity, and they are ncvcr rclcascd in packag-
Chcrnistry: No lichcn substailces detected. es like in B, helicospom. Thc uppcr pait of the hy-
Pigments: Upper part: of hymenium and rim o f proper rneniu~llof B. I(iu,-ucerasi is evenly pigmented, whereas
exciple K+ purplisl~, N+ orange-red to red. Hy- the eplthecial pigmentation of B. helicuspur-a is concen-
pothecium and inner palls of proper exciple K+ slightly trated to the gelatinous matrix surrounding thc asci and
the paraphyses adjacent to the asci. Very young apothe- brown to black but often mottled with various pink, or-
cia of B. helicosporu yet lacking asci are sometimes al- ange, brown, or violet hues. Proper exciple brown or
most pigment deficient in the hymenium. The apothecia red-brown in the rim, otherwise colourless to orange-
of B. helicosporu are usually flatter than in B. laurocer- brown. Hypothecium colourless or pale yellow. Upper
asi. part of hymenium f brown; pigment forming distinct
Bacidia helicospom is also likely to be confused with caps over the apices of the paraphyses. Spores acicular,
B. heterockrou. The latter however, has distinctly 32-73 X 2.5-4.3 p,with 3-15 septa.
brown-pigmented apices of the paraphyses, spores that
are not conglutinated (although sometimes coiled in ThaIlus indeterminate or determinate, very thin to rather
young asci), and wider paraphysis apices. thick, usually continuous, usually wrinkled but some-
times smooth or warted, without cracks, f cracked, or
Additional collections examined U . S. A. Florida. Wash- areolate, rarely discontinuous, of f discrete, thin are-
ington Co., 1993, Ekrnan L1 140 (LD). Illinois. Union oles, light grey to grey to yellow-grey to green-grey to
Co., 1878, Earle (NY). Kentucky. + CaIloway Co., 1975, blue-grey, rarely grey-green or greyish yellow. Prothal-
Buck B34 (filed with B. schweinitzii, Mm).Louisiuna. lus often present as a thin, bIack line bordering the thal-
East Feliciana Par., 1969, Tucker 7833B (herb. Tucker). lus.
Grant Par., 1973, Tucker 11315 & Jones (herb. Tucker). Apothecia (0.5-)0.5-0.7-0.8(- 1.1) mm diam. (S= 0. l ,
St. Helena Par., 1972, Tucker 10149 (herb. Tucker). Web- N= 23, n= 5 ) , plane or moderately convex, remaining so
ster Par., 1973, Tucker 11440B (herb. Tucker). Mayland. or rarely becoming markedly convex, epruinose or rare-
Montgomery Co., 1933, Leonard 2202 (US). Oklahoma. ly thinly pruinose on the edge of a few young apothecia.
Beavers Bend State Park, 1946, Chester 44 13 (US). South Disc purple-brown to purplish black to pure black, often
Carolina. Ravenel 143 (US). Tennessee. Blount Co., mottled with pale pink, pale orange, pale brown-violet,
1956, Sierk & Sharp 32 (US). brownish pink, pink-brown, brown-purple, or greyish
violet. Margin in upper part often concolorous with
disc, sometimes paler than disc (i.e., Iike the Iower part
11. Bacidia heterochroa (MiilI. Arg.) Zahlbr. of the margin), or rarely darker than disc, lower part
concoIorous with or paler than disc, commonly f brown
Cat. Lich. Univ. 4: 204 (1926). - Patellariu heterochroa or f pink; distinct, raised above disc in young apothe-
Miill. Arg., Flora, Jena 63: 280 (1880). -Type: Argenti- cia, soon level with the disc, persistent or only rarely ex-
na, "Buenos Ayres, auf Excaecariu biglandulosa", cluded.
1880, Schnyder (G, lectotype selected here). Proper exciple laterally (46-)67(-120) pm wide (S=
Parellaria subpellzkcida Mull. Arg., Flora, Jena 64: 16, N= 23, n= l), without crystals or rarely with radiat-
522 (1881). - Bacidi~subpellucida (Mull. Arg.) Darb., ing clusters of crystals that are up to 4 pm in d i m .
Wiss. Ergebn. schwedish. Siidpolarexpedit. 4 (11): 49 Rim pigmented in a distinct zone, f brown or red-
(191 2). - Type: Argentina, "Insula de Becreo in Argen- brown in upper part, downwards usually gradually
tinium", 1881, Schnyder (G, lectotype selected here). changing to pale yeI1ow or colourless, along the edge
Patelluri~ atlantica Miill. Arg., Bot. Jb. 5: 137 with a single layer of enlarged celIs with lumina that
(1884). - Bacidira aalantica (Miill. Arg.) Zahlbr., Cat. are up to 9 X 6 pm. Lateral interior paler than rim, col-
Lich. Univ. 4: 178 (1926). - Type: Atlantic Islands, "As- ourless, pale yellow, brown-orange, or orange-brown.
cension, in ramulis", 1883, Naumann 42 (G, lectotype Medullary part usually concolorous with or paler than
selected here). hypothecium and lateral interior, colourless, to paIe
Bacidia alutacea sensu Malme, Ark. Bot. 27A (5): 17 yelIow. Hypothecium colourless or pale yellow. By-
(1935), non (Krempelh.) Zahlbr. menium (74-)76-9 1- 110(-115) p m thick (S= 9, N= 23.
n= 5), lower part colourless except for occasional ver-
Nomenclature. The name Bacidila alutacea was used for tical streaks with brown pigment, uppermost part f
this species by Malme (1935), and his use has been fol- brown, red-brown, grey-brown, or black-brown, usual-
lowed by some authors. The type of that name, however, ly f distinctly delimited. Paraphyses 0.8- 1.6 pm wide
is a quite different species of Baddiu S, str. Bacidia het- in mid-hymenium; apices f clavate or not at all swol-
erochroa is the only species of Bacidiu I know of, which len, (1.2-)l .9-2.6-3.0(-4.0) pm wide (S= 0.6, N= 23,
has been collected in all continents of the world. Thus, n= 5), with grey-brown pigment in the wall forming a
there is a risk that an older name will be found. distinct "cap". Spores acicular, straight, slightly
curved, or sometimes f sigmoid, sometimes coiled in
Illustrations. Figs 5B,6B, 24 (map), 41C. ascus, (32-)36-50-67(-73) p long (S= 8, N= 23, n=
10), (2.5-)3.1-3.4-4.0(-4.3) pm wide (S= 0.4, N= 23,
Characterizatioa. Apothecia usually mainly purpIe- n= 10), (9.3-)11.7-15.1-18.9(-23.9) times as long as

Opera B o t 127, 1996

Fig. 24. Known North American distribution of Bacidia hetemchma.

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-

Opera Bot. 127. l996

77
 cies of Bacidia. I have seen specimens from tropical and MIN, SCSC, UBC, US, WTU). Sail Luis Obispo Co.,
subtropical areas around the world. 1989: Ekman L624 (Ell). San Mateo Co., 1967, Jordan
636: 665, 818, 826 (SFSU), 1974, Brodo 20413, Thiers,
Discztssion. Buci~lin heterochrou is likely to be con- Sigal & Toi-en (CANL), 1978: Thiers 39236 (SFSU): +
fused with B. helicospora (see that species), B. lau- 1967, Jordan 8 1 8 (filed with B. arct'zr!itro, SFSU). Santa
ruuerusi, and the neotropical species B. megupo~ui~zica Barbara Co., 1989, Ekman L637 (LD). San Gabriel
Malme. B. luurocerrasi differs in having none or few Mountains, 1902: Hasse (NY). Santa Cruz, 1888: possi-
pigment caps in the wall of the paraphyscs apices, bly collected by Anderson (NY). Santa Monica, 1897,
more-septate, more distinctly sigmoid spores, and in on- Hasse 536 (NY). Santa Monica Mountains, 1906, Hassc
ly rarely having the apothecium disks mottled with dif- (COLO, US), + 1909, Hasse 2 122 (filed with Bn. ram-
ferent colours. Like B. heterochrou, B. nrega~3ntutnica ecr, MIN). Santa Monica Range, 1902: I.lasse (NY),
forms distinct pigment caps in the wall of the parapllys- Hassc 536 (NY). Florida. 1878, Austiil ( U S ) . Calkills
es apices. These caps, however, consist of a diffcrcnt 76 (NY), Calkins 89 (FH):Calkins 9 1 (COLO). Alachua
greyish brown pigment not rcacting with K. Further- Co., 1982, Griff?n 96, 137 (herb. Tucker), 1982, Griffin
more, according to Malme (1935), B, megapotaniira 137a (FLAS, herb. Tucker, MIN, NY), 1993. Ekman
differs in having shorter sporcs with fewer septa. L1 141, L1 142, L1 143 (LD). Bay Co., 1993, Ekman
Baciclicd eckjeldtii (Miill. Arg.) Zahlbr., described L1217 (LD). Collier Co., 1993, Ekman L1 174, L1203,
from thc vicinity of Monterey in northeastern Mexico, is L1204 (LD). Dade Co., 1982, Tucker 25228 (herb.
very siinilar to B. heteruchruu. It differs in having the Tucker), 1993, Ekman L 1 182, L 1 183, L 1184 (LD).
hymcnial pigment evenIy distributed (not forming dis- Highlands Co., 1965, Wctmorc 13905 (MIN). Wakulla
tinct caps), and in having distinctly sigmoid spores with Co., 1993, Ekinan L1209 (LD). + Caloosa Rivcr, 1878:
more septa (1 1 - 17). In these respects, B. eckfeldtii ap- Austin (in type specimen of Biafor-Uca1oosens.i~Tuck.=
proaches B. lallrocerasi, but the pigmeiitatioil of the Bucidia hostheleoide.~,FH). Fort George, Calkins (CO-
apothecia is more siinilar to B. heteroohruu. The only LO, FH, MIN, NY). Jacksonville, 1 886, Eckfeldt (US).
known specimcn of B. eckfeldtii was collected on lig- Calkins (COLO, NY, US), 1889, Calkins (MINI. San-
num. ford, Rapp (FH), 1909, 1912, 1913, Rapp (FH), 1914,
The western and eastern populations of B. hetcro- Rapp (FH, MIN, US), + 1914, Rapp 22 (fiIcd with Bn.
chroa differ slightly but consistently in the secondary varia and Bn. sp. # l , FH), 1929, Rapp (UPS). Sanibel
chemistry. Furthernlore, the numerical and statistical Island, Hume (NY). Upsala, 1909, Rapp 32 (FH). GEOI--
analysis revealed additonaI differences in morphologi- gia. 1877, Ravenel (US). Glynn Co., 1969, Brodo
cal characters. These differences, however, are slight IIi4j5 (CANL). Ware Co., 1989, Wetmure 64692
and overlapping. I have not been able to detcct any oth- (MIN). Lozrisinnn. 1895, Langlois ( F H ) . Allen Pal-.:
er discontinuous difference between these groups of 1986, Tucker 27294 (herb. Tucker). Calcasieu Par.,
populations other than the chemistry. Further detailed 199 1, Lieveils 5030a (herb. Tucker). Cameron Pal-.,
analyses of the variation pattcms, also includii~gmateri- 1 967, Tucker 6794 (herb, Tucker), 199 1, Lievens 4876c
al from Central and South America, is needed before a (herb. Tucker). East Baton Rouge Par., 1968, Tuckcr
final coi~clusioncan be drawn as to the taxonomic status 729hb (herb. Tucker), 1969, Tucker 7943 (COLO, herb.
of the eastern and western population groups. Tucker, US), 1969, Tucker 7943b (hcrb. Tuckcr), 1970,
The types of Bacidia subpelhcida (Miill. Arg.) Darb. Tucker 8988 (herb. Tucker), 197 1, Tuckcr 9044b (hcrb.
and B, utlanticu (Miill. Arg.) Zahlbr. are typical speci- Tucker), 197 1 , Tucker 9969 (herb. Tucker, MIN), 1974,
inens of B. keterochroa and are thus reduced hcrc into Tuckcr 11698b (CANL), 1977, Tucker 16833, (MIN).
synonymy Evangcline Par., 1969, Tuckcr 8384 (herb. Tucker). Ibe-
ria Par., 1984: Tuckcr 26637 {hcrb. Tuckcr). -+ lbcrrillc
Culleclions examined. U. S. A. Alabama. Baldwin Co., Par., 1974, Tucker 13391 (filed with B. schweinitzii,
1925, Evans 435, 441, 458 (NY, US). California. Con- herb. Tucker). Jefferson Par., 1970, Johnson 45 (herb.
tra Costa Co., 1930, Herre (US). Marin Co., 1925, Parks Tucker). + Livingston Par.: 1968, Tucker 7351 (filed
2883 (COLO), + 1925, Parks 2883 (filed with Bn. cali- with B. schweinitzii, herb. Tucker). Plaquemine Par.,
fornica, COLO), + 1939, Koch 565a (filed with B. insll- 1884, Langlois 589, 593, S . n. (US), 1884, Langlois 599
Iaris, COLO), 1969, Wennekens 77 (SFSU), 1977, Cal- (US), 1886, Langlois 598 (US), 1885, Langlois 186
houn 125 (SFSU), 19X9, Ekmail L640, LG42, L644 (US), 1884-85, Langlois 184 (US). + Sabii~cPar., 1967,
(LD), + 1989, Ekman L645 (filed with Bn. californica, Wetinore 17563, 17587 (filed with B, schn~eir~ibii:
, LD). Mendocino Co., 1973, Thicrs 30885 (SFSU), MIN). + S t . Hclena Par., 1972, Tucker 10149 (filcd wit11
1982, Thiers 45648 (SFSU), 1989, Ekman L650 (LD). + B. helicuspura, herb. Tucker). St. Martin Par., 1889:
Monterey Co., 1966, Weber & Santesson (filed with Bn. Langlois 590 (US). St. Tamrnany Par., 1982, Tucker
californica in Weber: Lich. Exs. Boulder 182, COLO, 24442a (herb. Tucker). Tangipahoa Par., 1975, Tucker
15149 (NY), 1976, Tucker 15285 (herb. Tuckcr). Vcnnil-
ion Par., 199 1, Lievens 4869b (herb. Tucker). + West Fc-
iiciana Par., 1978, Tucker 18 127 (filed with B. schwein-
ihii, herb. Tucker). Banker's. 1894, Langlois (US).
Bayou Silan, 1898, LangIois (US). Bayou Tortue, 1893,
Langlois (US). Bois Charmant, 1893, Langlois (US).
Bois Lobbk, 1891, Langlois (US), + 1893, Langlois X68
(filed with B. schweinitzii, US). Breaux Bridge, 1893,
Langlois 867 (US). Charcnton, 1889, Langlois 59 1 (US).
Grand Coteau, 1894, Langlois (filed with B. schweinifiii,
U S ) . John Durand's, 1894, Langlois (US). Opelousas,
1889, Langlols h17 (US). Petite Prairie, 1884, Lai~glois
5-97 (US). Pointe-h-la-Hache, 1884, Langlois 183, 588,
S , n. (US). St. Leo, 1893, Langlois (US). St. MartinsvilIe,
1893, Langlois 866 (US), 1895, Langlois (CANL, NY, Fig. 25. Known North American distnbutlon of Bacidia
US), 1895, Langlois 475 (NY). North Carolina. Dare (dot) and 8. mlrtubiiis (Irianglc).
hosfheienide.~
Co., 1959, SchalIert L5 10 (WTU). Oregon. + Curry Co.,
1989, Ekman L673 (filed with B. sahtzonea, LD). + Lin-
coln Co., 1989, Ekman L680, L68 1 (filcd with B. salnlo- Wien 83: 128 (1 909). - Type: Brazil, "Sao Paulo. Prope
nun, LD). Tillamook Co., 1989, Ekmai~L709 (LD), + Fazenda BelIa Vista in districtu urbis S. Cruz ad flumen
1989, Ekman L71 1 (filed B. salinonen, LD). So~itlaCaro- Rio Pardo, ca 500 m s m, corticola", 1901, Wettstein &
lina. + Ravenel (tiled with B. .schweinihii, US). Charles- Schiffner (W, lectotypc selected here).
ton Co., 1988, Wet~noreh l 5 5 3 (MIN). Tsar. BurIeson BuciJiu rissoensis Malme, Ark. Bot. 27A (5): 32
Co., 1976, Tucker I5714 (herb. Tucker). Cameron Par.. (1935). - Type: Paraguay, "Colonia Risso", l F Q 7
1990, Licvens 4573b (hcrb. Tucker). Fort Bend Co., MaIme l81 3 (LD, syntype).
1991, Lievens 5344 (herb. Tuckcr). + Hardin Co., 1975, Bncidia nzedia1i.s f, ini#is.czrla Malme, Ark. Bot. 27A
Tucker 14027 (filed with B. sohwuinitzii, herb. Tucker). (5): 32 (1 935). - Type: Brazil, "Coxipo Mirim pr. Cuy-
Robertson Co., 1976, Tucker 15797 (herb. Tucker). abi", 1894, Malme 2346 (S, lectotype selected here).
Travis Co., 1989, Lievens 3594c, Lynch, Kramcr &
Kramer (herb. Tucker). Illustrcrfions. Pig. 25 (map).

Cl~aracterization.Thallus greyish. Apothecia brown-or-

12. Bacidia hostheleoides (Nyl.) Zahlbr. atlge. Proper exciplc vcry pale orange; lumina in lower
part narrower than or as wide as lumina in upper part.
Cat. Lich. Univ. 4: 205 (1926). - Lecideu hosfheleoides Hypothecium very pale orangc. Upper part of hy-
Nyl., Acta Soc. Sci. fenn. 7: 458 (1 863). - Type: Vene- menium very pale orange. Spores bacilliform, fusiform,
zuela, "Nova Granata, Villeta, aIt. I200 m", 1860, or almost clavate, (almost) straight, 16-25 . 2.9-5.0 pm,
Liildig 2668 (PC, lectotype selected here; H, isolecto- with 3-5 septa. - Neotropical speciniclis usually have
type). brown-orange to f purple-brown, seldom k pink apothe-
Putelluria r~!fhscensMiill. Arg., Flora, Jena 67: 467 cia, a colourless to f orange or partly f red-brown prop-
(1884). - Baoidia rufiscens (MiiI1. Arg.) Zahlbr., Cat. cr cxciple, and a colourless to & red-brown upper p,art of
Lich. Univ. 4: 239 (1926). - Typc: Mexico, "Orizaba", the hymenium
1853, Miiller (G, lecioiype selected by Awasthi &
Mathur 1987 as "holotype"). Thallus indeterminate, thin to thick, mainly continuous,
Biaforcl culoosensi.~Tuck., Synopsis North. Am. f cracked and wrinkled, or granular to subsquamulose,
Lich. 2: 4 1 ( I 888). - Bacidiu culoo.sensis (Tuck.) Zahl- partly discontitluous, of discretc to contiguous granules,
br., Cat. Lich. Univ. 4: 184 (1926). - Type: U. S. A., pale grey to pale grecn-grey. Granules (3 1 -)57(-80) pm
Florida, "Caloosa River", 1x78,Austin (FH-TUCK, lec- wide (S= 1 5, N= l , n= 20). Prothallus lacking.
totype selectcd hcrc, excluding specimen d'; US, isolec- Apothecia (0.5-)0.6(-0.8) mm diam. (S= 0.08, N= 1,
totype) n= 20), at first plane, later becoming convex, epruinosc.
Lecidea subpur Nyl., Labuan et Singapore 39 ( 189 1 ). Disc brown-orange. Margin concolorous with disc,
- Bacidiu swbpur (Nyl.) Zahlbr., Cat. Lich. Univ. 4: 243 raiscd above disc in young apothecia, later level with
(1926). - Type: Cuba, Tuckerman: Wright Lich. Cub. the disc, fillally excluded.
1 14 (H-NYL 17370, lectotype selected here). Proper exciple laterally 40 pm wide (N= l , n= l),
Bacidia ve-~ans Zahl br., Denkschr. Akad. Wiss., without crystals; lumina inside rim in upper part 1.2-2.8

O])era Hot. 127. lYYh

pm wide, in lower part 1.O-1.5 pm wide. Rim very pale ellipsoid and distinctly wider than the lumina of the up-
orange, without distinct zone of enlarged cells along the per part of the proper exciple. Possibly, there is also a
edge (lumina gradually expanding towards edge or not difference between the species in the length-width ratio
at all expanded). Lateral interior concolorous with rim. of the spores, but this requires confirmation.
Medullary part colourless. Hypothecium very pale or- Apart from B, mediulis, B. hostheleoides can be con-
ange. Hymeniuin (57-)60(-62) p thick (S= 2, N= 1, n= fused with Bacidina varia and an unidentified (possib1y
5), colourless in lower part, diffusely and very pale or- undescribed) species of Cutillaria sensu Zahlbr. B.
ange in upper part. Paraphyses 1.O-1.6 pm wide in mid- hostheleoides is distinguished from these species in
hymenium; apices only slightly or not at all swollen, much the same way as B. medialis (see this species).
(1.2-)l .8(-2.4) pm wide (S= 0.4, N= 1, n= 20), without Bacidia hmtheleoides is a poorly understood species,
internal pigment. Spores bacilliform, fusiform, or al- which has had the unfortunate destiny of being de-
most clavate, straight or slightly curved, not coiled in scribed several times. T h e types of Bacidiu mfescens
ascus, (16-)l 9(-25) pm long (S= 3, N= 1, n= 20), (2.9-1 (MulI. Arg.) Zahlbr., B. caloosensis ((Tuck.) Zahlbr., B.
3.6(-5.0) pm wide (S= 0.4, N= l , n= 20), (3.8-)5.4(-8.7) subpar (Nyl.) Zahlbr., B. vexans Zahlbr., B. is so ens is
times as long as wide (S= 1.2, N= l , n= 20), with (3-1 Malme; and B. mediulis f. majuscula Malme all repre-
3.1(-5) septa (S= 0.4, N= 1, n= 20). sent typical specimens of B. hostheleoides. In North
Pycnidia half-immersed in thallus, pale orange, 50- American literature, this species has mostly been known
100 pm diam., containing a mixture of two types of co- under the name B. caloosensis.
nidia: (1) filiform, curved, non-septate, 10-14 x 0.5 w,
and (2) short-fusiform to eIlipsoid to obIong, non-sep- Collections examined. U . S. A. Florida. Caloosa River,
tate, 6-9 X 1.6-2.0 pm. 1878, Austin (FH, US).
Chemistry: No lichen substances detected.
Apothecial and pycnidial pigments: All parts of
apothecia and pycnidia K-. - Small amounts of Rubella 13. Bacidia igniarii (Nyl.) Oksner
Orange in proper exciple, hypothecium, hyrnenium, and
pycnidiaI wall. Flor. Lish. Ukraini 2: 166 (1968). - Lecidea ipiarii
Nyl., Flora, Jena 50: 328 (1867). - Type: FinIand, "Ad
Ecology. On the bark of an unidentified hardwood. Polyp, igniarium in Tavastia", 1863, Norrlin (H-NYL
17232, Iectotype selected here).
Distribution. Collected once in southwestern Florida Bacidiia abbbmviuns Th Fr., Lich. Scmd. 2: 362
(the type specimen of Biatoru culoosensis Tuck.). - (1874). - Type: Not designated; isosyntype in LD stud-
Widely distributed in the neotropics. ied.

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

Opera Bot. 127. 1996

grey-green. Prothallus sometimes present between dis- Discussion. Bacidia insularis is similar to B. circuna-
crete areoles, thin, Iight grey. specta and B. sribincompta. Both these species differ
Apothecia (0.4-)0.5-0.7-1.0(-1.7) mm diam. (S= 0.2, from B. insularis in usualIy having the proper exciple
N= 5, n= 10), plane or slightly convex, epruinose or dominated by a brown pigment (Laurocerasi Brown).
with white pruina on edge of young to medium-aged Furthermore, B. circumspecta has narrower spores with
apothecia. Disc black. Margin black, distinct, raised fewer septa, and usually a colourless hypothecium that
above disc in young apothecia, later level with the disc, never contains Bacidia Green. B. subincompta also dif-
persistent. fers in having wider paraphyses. Although similar to the
h o p e r exciple laterally (57-)67(-83) pm wide (S= 11, species mentioned, there is a possibility that B, insuluris
N= 5, n= l), without crystals or with radiating clusters is more closely related to other species with a zone of
of crystals that are up to 3 jm wide. Rim dark blue- enlarged cells along the edge of the proper exciple, i.e.,
green to almost black (often in a distinct zone) in upper B. campalea, B. russeola, and B. suJiusa.
part, downwards changing to pale blue-green, along the The type collection differs from the Californian col-
edge with a f distinct, 1-3 cell layers thick zone of en- lections in a having more abundant white pruina on the
larged cells that are up to 10 X 5 p.Lateral interior margins of the apothecia, a wider zone of enlarged cells
paler than rim, colourless to blue-green, sometimes along the rim of the proper exciple (3-5 cell layers), and
mixed with some brown. Medullary part colourless. Hy- larger conidia (13-22 X c. 0.8 p,non-septate or I-sep-
pothecium f pale brown, dirty brown, green-brown, or tate). h my opinion, these differences do not justify a
sometimes blue-green in upper part and pure brown in separation of the Californian populations as a separate
lower part. Hymenium (74-)82-90-96(-99) pn thick (S= species. B. insularis is a poorly known species, and as
7, W= 5, n= 5), lower part colourless, uppermost part more collections become available, it may turn out that
dark blue-green to almost black in a distinct layer. Para- the variation is continuous.
physes 0.8-1.2 p m wide in mid-hymenium; apices f cla-
vate, (1.6-)2.6-2.9-3.5(-4.9) pm wide (S= 0.8, N= 5, n= Coliectiom examined. U . S . A. California. Catalina Is-
10), unpigrnented or with an external cap of blue-green land, 1911, Hasse (FH), 1912, Hasse (m), 1915,
pigment. Spores fusiforrn or sometimes acicular, (26-) Hasse (Plitt: Lich. exs. ex herb. Hasse rel. 52, COLO).
30-40-55(-77) pm long (S= 12, N= 5, n= 10), (3.1-)3.5- Santa Barbara Co., 1994, Nimis & Tretiach (TSB, 3
4.2-4.7(-5.3) pm wide (S= 0.6, N= 5, n= 10), (5.3-)6.4- coll.). Marin Co., 1939, Koch 565a (COLO).
10.0-14.1(-20.7) times as long as wide (S= 4.1, N= 5, n=
10), with (3-)6.6-7.8-10.3(-15) septa ( S = 2.2, N= 5, n=
10). 15. Bacidia laurocerasi (Delise ex Duby) Zahlbr.
Pycnidia f immersed in thallus, uppermost part of
wall concolorous with apothecia, 100-150 p diam. Cat. LicR. Univ. 4: 2 13 (1926). - Putel!uria luurocerusi
Conidia filifom, curved, non-septate, 7-14 X c. 0.5 W. Delise ex Duby, Botanicon Gallicum 2: 653 (1830). -
Chemistry: No lichen substances detected. Type: Not seen.
Pigments: Upper part of hyrnenium and proper exci- Biatom atrogrisea Delise ex Hepp, Flechten Europas
ple K-, : N purple with a precipitate of blue crystals. 26 (1853). - Bacidia atrogrisea (Delise ex Hepp) Korb.,
Hypothecium K+ slightly intensifying or K+ purplish, Parerg. Lich. 133 (1861). - Type: Switzerland, "Ziirich,
and N+ intensifying, N+ orange-red, or N+ purpIish to an der Rinde junger Tannen und Nussbaume", Hepp,
purple with a precipitate of blue crystals. - Bacidia Hepp: Flecht. Eur. 26 (LD, syntype).
Green in hymenium, hypothecium (small amounts) and Lecidea endoleuculu Nyl., Lich. Japon. 68 (1890). -
proper exciple. Laurocerasi Brown and Rubella Orange Bacidia endoleucula (Nyl.) Zahlbr., Cat. Lich. Univ. 4:
in hypothecium. 194 (1926). - Type: Japan, "Itjigome", 1879, AImquist
(H-NYL 17734, syntype).
Ecology. On branches and twigs of shrubs in f exposed Lecidea invertens Nyl., Acta Soc. Sci. fenn. 26 (10):
or slightly shady habitats, e.g., in Artemisia chaparral. 33 (1900).- Bacidia inveraenrr (Nyl.) Zahlbr., Cat. Lich.
Known phorophytes include Artemisia californica and Univ. 4: 252 (1926). - Type: Japan, "Nagasaki", 1879,
Rhus lauriaa. Alrnquist (H-NYL 1728 1, lectotype selected here).
Bacidia rrubacerina Vain., Acta Soc. Fauna Flora
Distribution. Known from the southern and central parts fenn. 53 (1): 174 (1922). - Type: Not designated; iso-
of coastal California. - Bacidia insularis was described syntypes in ED studied.
on a single collection made by Herre in the Galapagos Bacidia endoleuca auct.
Islands, until now the only known 1ocaIity. Future inves-
tigations may well show that this species is rather wide- Nomenc~uture.I have not been able to trace any original
spread in coastal areas of Central and South America. material of Patellaria laurocermi Delise ex Duby. Fur-

Opera B o t 127. 1996

Fig. 26. Known North American disiibution of Bncidia laurocernsi. - Western (Washington, British Columbia, Idaho, and
Montana): known world distribution of subsp. idahoensis. - Eastern: known North American distribution of subsp, laumcerrari.

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

Opera Bot. 127, 1996

34
& Argus (CANL). Queens C o . , 1989, Ekman L1058 Perce National Forest, 3100 R, on Arnebnchier bark",
(LD). Ontario. Algoma District, Nagamisis Prov. 1949, Cooke & Cooke 25 174 (UPS,holotype).
Park, 1968, Brodo 13883 (CANL). Renfrew Co.,
1986, Wong 4035 (CANL). Thunder Bay District, Nomenclature. The date on the label of the holotype is
Rainbow Falls Prov. Park, 1965, Brodo 6636 given as "14 May 1949", but in the protologue it is erro-
(CANL, COLO). Lake Superior Prov. Park, 1989, neously cited as "May 15, 1940" There are two facts
Ekman L1000 (LD). Slate Islands, 1976, Wetrnore pointing towards the label bearing the correct date and
25256, 25637 (MIN). Prince Edward Island. Brack- the protologue an incorrect date. Firstly, Stevenson
ly Point, 1888, Macoun 2824 (CANL). Quebec. (1971) mentions that most of the material colIected
ComtC Abitibi-Ouest, 1,970, Brodo 16938, Gaudreau, jointly by Dr. Cooke and Mrs Cooke was collected after
Cloutier & Miron (CANL). Compton CO., 1963, Bro- 1942. Secondly, in the same work, two labels are de-
do 498 (CANL). Comtk Gasp&-Est, 1972, Savage 16 picted, one dated June 16, 1939 with the collection
(CANL). Comtb Gatineau, 1977, Brodo 2 1225 number 13257, and one dated November 5, 1949 with
(CANL). - U. S. A. Maine. 1922, Plitt (US). East- the collection number 26271. Provided that the Cookes
port, 1877, Farlow (FB). Mount Desert Island, 1891, numbered.their collections sequentially, the type speci-
White (NY).Rockport, 191 1, MerrilI (FH). Salisbury men of Bacidia idahoensis (numbered 25174) must be
Cove, 1922, Plitt (US). South Thornaston, 1912, collected much closer to November 1949 than to June
Merrill (FH), 1912, Merrill, Merrill: Lich. exs. 260 1939. That the specimen in question really is the holo-
(COLO, FH). South West Harbor, 1907, Sevivey type, and not some other specimen irrelevant to the typi-
(FH). M~ssaclau~setts.Wellesley, 1903, Cumrnings fication of the name, is further supported by the fact that
(FH). Michigan. Alger Co., 1987, Wetmore 59566 in the protologue, the holotype specimen of Bacidio
(MIN). Delta Co., 1976, Harris 11785b (CANL, idahoensis is stated to occur as an "admixture in Per-
MIN), 1976, Harris 12010 (UBC). Emmet Co., 1974, hsaria sp. (conf. multipu~lcta,without spores)". On the
Buck (NY).Keweenaw Co., 1947, Thomson, 2914 label, "Pertusaria cfr. multipuncta" is mentioned, and
(Mm),1983, Wetmore 4927 1 (MIN). Mianesota. this Iichen dominates the collection.
Becker Co., 1974, Wetmore 23768a (MIN). Clearwa-
ter Co., 1963, Hale 23463, 23482 (US), 1972, Wet- Illustrations. Fig. 26 (map).
more 20400, 20513 (MIN), 1972, Wetmore 20820
(CANL, MIN), 1976, Wetmore 26163 (MIN). Hub- Thallus rather thick, grey-green, finely areolalate. Apothe-
bard Co., 1974, Wetmore 23 14 1 (CANL, MIN, US). cial margin paler than disc (particularly conspicuous in
Koochiching Co., Hale 22212, 22215 (US), 1978, young apothecia), with an (orange to) dark carmine red
Wetmore 33463 (MIN), 1979, Wetrnore 37301, to red-brown hue. Hymeniwn (90-194-1 08-125(-13 1)
37881 (MIN), 1978, Wetmore 33428 (CANL, MIN). p n thick ( S l l , N= 9, n= S). Spores (52-169-81-96(-
Lake Co., 1977, Wetmore 26602, 2665 1b (MIN), 108) pm long (S= 11, N= 9, n= IO), (2.5-)2.8-3.1-3.2(-
1989, Ekman L943 (LD). Roseau Co., 1901, Fink 3.7) pm wide (S= 0.3, N= 9, n= 10), with (9-)17.5-19.8-
303 (MIN), 1901, Fink 324 (US). St. Louis Co., 23.0(-28) septa (S= 3.5, N= 9, n= 10).
1973, Wetrnore 2 1697 (MIN), 1978, Wetmore 3 1294,
3 1412, 31716, 32364, 32787, 33695, 33930, 34062, Ecology. Known phorophytes include Abies spp., Acer
34141, 34287, 34539 (MIN), 1978, Wetmore 35414 globrum, A. ~nacrophyIIum,Alnus mbra, Amelanckier
(COLO, MIN), 1979, Wetmore 38856, 38964, 40082 sp., Bewla papyrifem, Cornus nu~tallii,Pseudotsuga
(MIN), 1979, Wetmore 39845 (MIN, NY). New York. memiesii, Quercus gurvuna, Sulk sp., Taws brevifo-
Suffolk Co., 1910 Latham (FH), 1920, Latharn 950 lia, and Thuja plicda.
(US), Latharn (FB). Warren Co., 1982, Harris 16310
(NY). Ohio. Georgesville, 1894, Bogue, Merrill: Distribution. Western coastal lowland and western
Lich. exs. 226 (COLO, US). Wisconsin. Bayfield montane region of Oregon, Washington, British Colurn-
Co., 1987, Wetrnore 61007 (MIN). bia, Idaho, and Montana. The single known locality in
Oregon is unspecified and has not been marked on the
map. - Endemic.
15b. subsp. idahoensis (H.
Magn.) S. Ekman,
comb. et stat. nova Addiaional collectiom examined. CANADA. British
Columbia. Vancouver Island, Old BaIdy Mountain,
Bacidia idahoensis H . Magn., Acta Horti gothoburg. 1975, Noble 4399 (herb. Noble). Vancouver Island, NW
19: 45 (1952). - Type: U. S. A., "Idaho, Idaho Co., one of Nanoose, 1989, Ekman L755 (LD). Vancouver Is-
half mile up Rackliff Ridge Trail from Rackliff Creek land, Thetis Lake, 1989, E h a n L773 (LD). Vancouver
Public Camp Ground, along the Selway River, Nez IsIand, Victoria, 1977, Noble 6459c (herb. Noble). Van-

Opera Bar. 127, 1998 85

 brown-orange. Spores bacillifom, fusiform, or almost
clavate, (almost) straight, 16-40 - 1.9-3.7 p,with 3-7
septa.

ThalIus indeterminate, thin to rather thick, usually con-

tinuous, f cracked to areolate, wrinkled to warted,
sometimes subsquamulose or partly fxberculate to gran-
ular, rarely discontinuous, of discrete to contiguous,
thin, convex areoles; usually grey-green to green-brown,
rarely f grey. Frothallus lacking or present between dis-
crete areoles and along the edge of the thallus, thin, en-
Pig. 27. Known North American distribution of Bacidia dophloeodal, pale grey.
medialis. Apothecia (0.3-)0.3-0.5-0.7(-0.9) mm diarn. (S= 0.6,
N= 9, n= 10), at first plane, later becoming convex,
epruinose, Disc usually pale pink, pale yellow-pink, or
couver Island, Mount Douglas Park, 1991, Ryan (herb. pale yellow, sometimes pale orange to brown-orange to
M. Ryan, LD). Prevost IsIand, 1974, Noble 122Ib orange-brown to red-brown. Margin concolorous with
(herb. Noble). Purden Lake Prov. Park, 1976, Crane & disc, level with or raised above disc in young apothecia,
Noble 5953 (CANL, herb. Noble). Wells Gray Provin- later level with the disc, rather persistent but usually fi-
cial Park, 1989, Ekrnan L800 (LD). - U. S. A. Idaho. nally excluded.
Priest Lake, 1966, Thiers 17346 (SFSU). Montana. Flat- Proper exciple laterally (40-)48(-63) p wide (S= 7,
head Co., 1983, DeBoIt 372 (OSC).Lake Co., 1977, N= 9, n= l), without crystals, lumina inside rim in up-
McCune 8928 (OSC), 1989, McCune 17596, 17624, per part 1.2-3.2 pm wide, in lower part 1.5-5.0 pm
17627, 17630, 17640, 17642 (OSC). Ravalli Co., 1981, wide. R h colourless to paIe brown-orange, without dis-
McCune 11435 (OSC). Oregon. Cusick (US). Wading- tinct zone of enlarged cells along the edge (lumina grad-
ton. Clallam Co., 1989, Elunan L729, L732 (LD). ually expanding towards edge or not at all expanded).
Klickitat Co., 1882, Suksdorf (WTU), 1884, Suksdorf Lateral interior usually concolorous with rim (some-
297 (WTU), 1885, Suksdorf 310 (COLO). Lewis Co., times darker or slightly paler), colourless to pale brown-
1989, Thor 8376 (S), Pierce Co., 1994, Tmsberg 20212 orange, rarely brown-orange. Medullary part colourless
@G). to pale brown-orange. Hypotheci~uncolourless to pale
brown-orange, rarely brown-orange. Hymenium (45)
47-60-68(-74) pm thick (S= 7, N= 9, n= 5), colourless
16. Bacidia medialis (Tuck. ex Nyl.) de Lesd. or partly and diffusely pale yellowish to pale brown-or-
ange. Paraphyses 1.0-1.6 p wide in mid-hymenium;
Bryologist 24: 68 (1921). - Leeidea medialis Tuck, ex apices only slightly or not at a11 swollen, some occa-
Nyl., Annals Sci. Iiat., 4, 19: 346 (1863). - Type: Nica- sionally f clavate, (1 .O-)1.5-1.7-?.l(-3.2) p wide (S=
ragua, Wright (H-NYL 17372 specimen c, lectotype se- 0.4, N= 9, n= 10), without internal pigment. Spores ba-
lected here). cillifom, hsiform, or almost elavate, straight or slightly
Biatora molybdi-ditisTuck., Synops. North Amer. Lich. curved, not coiled in ascus, (16-)20-23-26(-40) pm long
2: 34 (1888). - Bacidia molybcbitis (Tuck.) Zahlbr., Cat. (S= 3,N= 9, n= 101, (1.9-)2.0-2.8-3.4(-3.7) p wide (S=
Lich. Univ. 4: 126 (1926). - Type: U. S. A., "Florida", 0.5, N= 9, B= 101, (4.8-)6.7-8.2-10.7(-16) times as long
1877, Smith 10 (FH-TUCK, lectotype selected here). as wide (S= 1.9, N= 9, n= IQ), with (3-)3.0-3.2-4,2(-7)
Patelluria rosellina Mull. Arg., Revue mycol. 10: 4 septa (S= 0.8, N= 9, n= 10).
(1888). - Bacidia roseflinn (Miill. Arg.) Zahlbr,, Cat. Pycnidia almost entirely immersed in thallus, colour-
Lich. Univ. 4: 238 (1926). - Type: Uruguay, "Montevi- less, c. 75 pm diarn. Conidia filifom, straight or slight-
deo", 1887, Arechavaleta (G, lectotype selected here). ly curved, non-septate, 13-17 X 0.8-1.0 pm.
Chemistry: No lichen substances detected.
Illustrations. Figs 5C, XB,27 (map). Pigments: Pigmented parts of apothecia K+ intensify-
ing, N-. - Without pigments or with RubelIa Orange in
Characterizmtio~r.Thallus usually greenish or brownish. proper exciple, hypothecium, and hymeniurn.
Apothecia usually pale pinkish or yellowish, seldom
pale orange to red-brown. Proper exciple coIourIess to Ecololy. On the bark of hardwoods and shrubs in hu-
pale brown-orange; lumina wider in lower part than in mid or dry, open or more or less shady forests and forest
upper part. Bypothecium colourless to pale brown-or- edges. A few finds are in pIanted tree-stands. K n o m
ange. Upper part of hymeniurn cohourIess to pale phorophytes include Cedrela odonta, Eugenia spp., Ifex
sp., Krugiodenhor~j i r r e ~ t n ~!Was~ichoderzdron
: ,foetidis- ourless. Upper part of hymeniun~colourlcss. Spores ac-
.sim~ttn,Rhizophora nzangle, Sulix ccrru)liniana, S a i n 6 ~ - icular: 33-57 X 2.9-3.7 ,m, with 3-9 scpta. -, Neolropi-
cus canudensis, Illmus u~rlericana,and Xit~zenia amur-i- cal specilllens usually have palc orange to purplisli
concr. black apothecia, a palc ycllow: bruwn-orange, or partly
red-brown rim of the propcr cxciple. which is concolor-
Di~triblttion.Florida, southernmost Louisiana, and one ous with the uppcr p a ~ of
t the hyn~enium.
unspecified locality in Texas (not on thc map). - Proba-
bly pantropical. Thallus dcterminate. thin, continuous, k cracked, wrin-
klcd, pale green-grey. ProthalIus lacking or prcscnt, bor-
Di,scu.rsion. Bacidia inediulis is closely related to, and dering the thallus, narrow, black.
easily confuscd with B. hostheieoides (see that species). Apothecia (0.4-)0.5-0.5-0.6(-m) mln diam. (S= 0.06,
B. rnedialis is superficially similar to Baciclina vauia, N= 2, n= IO), plane or nloderatcly convex, with thick
which call be separated on the basis of the longer and white pn~ina, particularly on the margin. Disc pink.
narrower, usually more-septatc spores that are never dis- Margin pale pink, raiscd above disc in young apothecia,
tinctly bacillifom, and on the anatoiny of the proper ex- soon level ~ ~ i tlic
t h disc, persistei~t.
ciplc. From Florida and Louisiana, l have seen several Proper excip1e latcrally (77-)82(-86) pm wide (S= h,
specimens of a species of Catilluria sensu Zahlbr., N= 2, n= I), with abundant large clusters of crystals
which superficially is vcry similar to RucirIil~niea'iali~. (singlc crystals up to 9 PI dlam.). Rim colourlcss,
It is separated from B. rrzedialis by tlie shorter, cnnstalit- along the edge wit11 a single layer of enlarged cells that
ly 2-celled spores and the lack of an ocular chamber in arc up Lo 5 X 3 p.Lateral interior colourless. Mcdul-
the tholus. I havc not come across any published ~ ~ a n l e lary pall colourless or vely pale yellow. Hyputhccium
for this species. It does not belong in Catiilariu S. str., colourless or very palc ycllow. Hy~neuium(56-)65-68-
but n ~ a ybe related to Lecania naeg~liiand Lecuniu slig- 7I(-74) pn thick (s=6: N=2, 11=5), colourless, upper-
nlnfellu. most p a ~ twith thick layer of crystals. Paraphyses 0.8-
In North American literature, Bucidiu nzedioiis has 1.6 pn widc in mid-hymenium; apices not at all
mostly been known as Bauidiu nlu~)hdi/i.v(Tuck.) Zahl- swollen, slightly tapering, or narrowly clavate, (1.0-)
br. The typcs of the latter ilarne and Bacidia rosellina 1.4-1.6-1.7(-2.8) p wide (S= 0.5, N= 2, n= 20), with-
(Miill. Arg.) Zahlbr, agree colnpletely with B. ~~zedialis, out internal pigment. Spores acicular, straight, curved,
however. Bacidia gmn~lifkru(Miill. Arg.) Zahlbr., de- +
or sigrnoid, straight in ascus, (33-)42-44-47(-57) pn
scribed from Costa Rica: is apparently very closc to (and loi~g(S= 6, N= 2, n= 20), (2.9-)3.3-3.3-3A(-3.7) pn
pcrhaps conspecific with) B. ~~rcclialis. It diffcrs from B. wide (S= 0.3, N= 2, n= 20), (9.3-)12.7-13.6-14.4(-18.4)
medialis mainly in Ilavlng consistently 7-scptate spores times as long as wide (S= 2.3, N= 2, n= 20), wilh (3-)
and a slightly higher hymenium (c. 80 pm). 5.9-6.2-6.3(-9) septa (S= 1.4, N= 2, n=20).
Pycnidia not seen.
Collectiuns ~'~rarnined. U . S . A. Fluridu. 1877, Smith 10 Clicmistry: Atranorin and zeorin.
(FH). Collicr Co.: 1993, Ektnan L1 175 (LD). Dade Co., Pigments: Hypnthecium and propel. exciplc K+ pale
1985, Thor 4641 (S). Monroe Co., 1993, Ekmai~L1 193: yellow, N-. - So~iletimes Rubella Orange in hypo-
Li194: L1195, L1196, L11Y7, L1198, L1201 (LD). Lhecium and medullary part of propcr cxciplc. Un-
Polk Co., 1993, Ekman L 1 167 (LD). Miami, 1924, ex known. K+ yellow substailce in propcr exciple. This
herb. Plitt (US), 1914, Nuttall (US): 1982, Tucker substance is probably not atranorin, since the secondary
25230 (herb. Tucker). Sanford, 19 14, Kapp 5 1 (MW). ~l~etabolites in Bacidia are genera1ly restricted to. the
thallus.

17. Bacidia mutabilis Malme was collected on the branch-

Ecology Bacidia ~~zlltclbilis
es of Xhizophom mungle and Conucc~~p-plis erec!li.s in a
Ark. Bot. 27A (5): 24 (1935). - Typc: Brazil, "Mat10 densc ~nangrovcstand.
Grosso, CuyabQ", 1893, MaIme (S, lectotypc selected
here). Disrrihzitiun. Presently known from only two specirncils
collected at West Lake in Evcrgladcs National Park,
Illustr-utions. Fig. 25 {map). southernmost Florida. - Previously known from a
number of localities in Brazil and Paraguay.
Chamcterizarion. Apothecia pink, with thick whitc
pruina. Proper exciple (almost) colourless, with large Discussion. Althougl~a rather nondescript species, B.
clusters of crystals, along the edge with a single cell lay- mlltabilis call be recognized by a combination of char-
er of enlarged cell lumina. Hypothcciuin (almost) col- acters: dominance or Rubella Orange in tlie apothecia, a
thin or absent zone of enlarged cell lumina along the 2 months older Lerideu subspu~lilict~u (Mull. Arg.) SIi-
edge of the proper exciple, presence of crystals in the zenb.). - Bncicliu subspau'i~~ellu
Zahlbr., Cat. Lich. Univ.
proper exciple, and presence of a thick white pruina in 4: 244 (1 926). - Type: U. S. A., "Jacksonville, Florida",
at least some young apothecia. The Florida specimens 1888, Eckfeldt (H-NYL 17 1 19, lectotype selected here).
of B. rntltahilL.~,however, deviate from South Ameri-
can specimens in being very poorly pigmented and in Nonrenc1utur.e. V~rrucarlrafilscortlhella was not accept-
having more crystals in the proper exciple. These dif- ed by the author in the original publication (Hoffmann
ferences may be entirely due to habitat modification. 1796): and thc namc is consequently invalid (Art. 34. l).
Thc locality in Florida is vcry shady and humid, In the prclacc: EIoffillailll statcs that taxa within brack-
whereas Maline (1935) reported it froin light forests. ets or lacking a number (as: e.g., Verr-I-E~caria,jirscor-I-E~bel-
In typicaI South American B. nzufubilis. the colour of' 10) are doubtful ("Varietates autern, subspecies, atque
the apothecia is very variable, even within the same etiam tales species, quales ut veras statuere adhuic dubi-
thallus. They vary from pale orange to brown-orange to tavi, aut uncis ( ) inclusas in adnotationibus, aut numero
purple-brown to purplish black (disc and margin f con- carentes speciei proxime confini adpositas reperies").
colorous except lower part of margin, which Inay be The type collection of Lecideu ~ ~ h s p a d i c eNyl.
a con-
paler). The upper part of tI~chymeniuln and the edge of tains a mixture of typical Bacidia poivchroa (dominating)
the proper exciple is palc yclluw to brown-urangc (Ru- and B. suffusa. It is clcar, howevcr, that Nylandcr rcfcrrcd
bella Orange), sometimes partIy red-brown (Laurocerasi to B. po/yoh~oa:since thc K-rcaction uf thc hyputl~eciuin
Brown). The remainder of the apothecial (issues is col- and the proper exciple is mentioned in the diagnosis.
ourless to very pale yellow. Along the rim of the proper
exciple, there is a 1-2 cell layers thick zone of cells with I/lu.stmfions.Figs 28 (map), 4 1 G.
enlarged lumina. Crystals are abundant in the proper ex-
ciple, but they do not obscurc thc tissucs as in the Flori- Chal-actel-izdiun. Thallus cracked to areolate. Apothe-
da specimens. cia brown-orangc to red-brown to dark purplish brown,
Bcrcidia ~~zufnbiliscall be confused with, abuvc all. B. soinctin~espruinusc. Proper exciple usually f brown-or-
I-osella and B. suffuusu. R. i.osella can be separated by ange to orange-brown. I-lypothecium darker than proper
the thicker apices of the paraphyses and by the ab- exciple. Upper part of hylneniurn colourless to orange-
sence of a pruina in the strict sense (although i t has a brown. Most pig~lle~lled parts K+ purple-red. Spores ac-
layer of crystals between the paraphyses and in the icular, 31-74 X l. .9-5.0 pm, with 2-15 septa.
proper exciple). B. suffuusa has a wider zone of en-
larged cell lumina along the edge of the proper exci- Thallus indeterminate, thin to thick, either continuous,
ple, a thicker thallus lacking prothallus, and generally finely wrinkled to warted, f cracked or sometimes areo-
larger apothecia. late, or discontinuous, of thin, discrete to contiguous,
convex areoles, almost white to grey or yeIlowish grey.
U . S. A. Florida. Dadc Co.,
Collecfions e~-uri~ined. Apothccia (0.4-)0.6-0.7-0.8(- 1.2) mm dialn. (S= 0.1,
1993, Ekman L1 193, L1 194 (LD). N= 22, n= 5 ) , k plane when young, latcr bccoinii~gcon-
vex, epruinose or with thin to thick white pruina 011
edge and disc of young to medium-aged apothccia. Disc
18. Bacidia polychroa (Th. Fr.) Korb. brown-orange to medium brown to red-brown to pur-
plish brown, rarely orange or dark purple-brown. often
Parerga Lich., fasc. 2: 13 I ( I 860). - Biatovn polyohmn darkening as the apothecia age. Margin concolorous
Th. Fr., Ofvers. K. VetenskAkad Forh. 12 (l): 17 with disc or slightly darker, thick and raised above disc
(1855). - Type: Ukraine, "Ucnnia in Acere campestri", in young apothecia, later level with the disc. and finally
probably coIlected near Charkow, Czernia,jev (UPS:lec- cxcluded in old and convex apothecia.
totype selected here). Proper exciple laterally (45-)63(-95) pm wide (S= 13,
Verrucarlu juscoru heUa Ho ff~n.,Deutsc hl. Flora 2: 175 N= 22, n= l), without crystals or with radiating clusters
(1796), nom. inval. - Lecideu l~i~enlr var. ,Jirscol-zrbellu of rninutc crystals (up to l p). Rim f brown-orange or f
Ach.: Mcth. Lich. G1 (1803). - Secoliga fuscotubella orange-brown, sometiines pale yellowish, pale yellowish
(Ach.) Stizenb., Nova Acta Acad. Caesar. Leop. Carol. 30 brown, or pale to dark brown, along the edge without or
(3): 53 (1863). - Bucidiu ,firscurube/lu (Ach.) Bausch, with a single cell layer thick zone of cells with enIarged
Verh. naturw. Ver. Karlsruhe 4: 107 (1869). - Type: Ger- lumina that are up to 9 X 5 p.Lateral interior usually f
many, "Gennania" (H-ACH 345A, lectotype selected concolorous with rim (sometimes paler or darker). Med-
here). ullary pait concolorous with lateral interior part or slight-
Lecidea .subspudicea Nyl., Sert. Lich. Trop. Labuan ly paler. Hypothecium f brown-orange, f orange-brown,
et Singapore 39 (I X9 I), nom. illeg. (homonym of the 1 - red-brown, or dark brown, darker than proper exciple
Fig. 28. Known North Amer-
ican distribution o f Bacidia
polychroa.

(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.

Opera Bot. 127, 1996

Co., 1976, Wetmore 26441 (CANL, MM). Otter Tail Thallus indeterminate, f thin, either discontinuous, of
Co., 1900, Fink 72, 172 (MIN). Peimington Co., 1900, discrete or contiguous, convex, often almost granular or
Fink 868 (Mm). Pine Co., 1983, Schuster 450, 552, slightly effigurate areoles that are sometimes warted or
859, 1364, 1534, 1946 (MW).Roseau Co., 1901, Fink subsquamulose, or continuous, f cracked to areolate,
221, 274, 301 (Mm). St. Louis Co., 1901, Fiilk 1516, wrinkled or warted, grey-green or grey-brown. Prothal-
1770 (MIN), 1902, Fink 5463 (MTN), 1973, Wetlnore lus sometimes present between discrete areoles, thin,
21685 (MIN), 1977, Wetmore 27534, 27967, 30791, endophloeodal, pale grey.
30860, 30867 (MIN), 1978, Wetmore 3 133Xa, 3 1348, Apothecia (0.3-)0.4-0.4-0.5(-0.7) mm diam. (S=
31372, 31449, 31782, 31783, 31792, 32472, 32693, 0.09, N= 4, n= 10), plane or moderately convex, re-
33047, 33140, 33144, 33269, 33925, 34214, 34431, maining so or sometimes becoming markedly convex,
34448,34668,34768,34850,34913 (MIN), 1978, Wet- epruinose, Disc usually grey-brown to dark purplish
more 34610 (CANL, MW), 1978, Wetmore 34715 brown to black-brown, seldom pink, pale orange,
(NY), 1978, Wetmore 32480, 33378 (COLO, MIN), blue-grey, or yeIlow-brown, sometimes mottIed with
1979, Wetmore 38263, 38605, 39156, 39737, 39830, different colours. Margin concolorous with disc or
40265,40269 (MINI, 1986, Trana 13331, 13493 (MW), slightly dxker, sometimes paler in lower part (to pale
1986, Wetrnore 57599 (MIN, NY). Collegeville, 1910, grcy) than in upper part, level with or raised above
Hansen 1 (MW). Minneapolis, 1896, Fink 101 (US). disc in young apothecia, soon level with the disc, per-
Red Lake, 1900, Fink 892 (CANL, MIN). Snowbank sistent.
Lake, 1897, Fink 861 (US). St. Cloud, 1897, Morgan Proper exciple laterally (26-)M(-60) pm wide (S= 17,
(US). Missouri. Oregon Co., 1988, Hanis 2 17 13a (NY). N= 4, n= l), without crystals. Rim grey-brown to green-
New York, + Buffalo, Wilson (filed with B. s~~fiusa, grey to dirty green in upper part, downwards often
US). East Galway, 1893, Burt (FH), + 1893, Burt (filed changing to colourless to pale orange, with cell lumina
with B. sums, FH). Newcomb, 1936, Lowe 6007 (NY). gradually slightly enlarging towards the edge (wider in
Richfield Springs, Willcy (US). Shushan, Plitt 28 (US). lower part than in upper part). Lateral interior colourless
Wmensburg, 1933, Lowe 3649a (NY). North Carolina. to pale orange to grey-brown; if grey-brown then down-
Swain Co., 1976, Harris 11009 (NY), 1987, Harris wards usually changing to colourless or pale orange.
20975 (NY). Ohio. 1897, Bogue (NY).Bogue 874 Medullary part colourless to brown-orange. Hy-
(NY). Orwell, 1891, Bogue, Memill: Lich. exs. 171 pothecium very pale yellowish to brown-orange, rarely
(COLO, US). Sugar Grove, 1893, Bogue 419 (NY). colourless. Rymenium (53-157-65-82(-84) pm thick (S=
South Carolina. Lamoille Co., 1985, Harris 18256 1 I , N= 4, n= 5), lower part colourless, upper part green-
(WY). Chester, 1882, Green (US), 1883, EckFeldt 48, grey or dirty green, sometimes with grey-brown spots.
102 (CANL). Vermont. Brandon, 1920, Dutton 989 Paraphyses l .O-1.2 pm wide in mid-hymenium; apices
(COLO), 192 1, Merrill 1 189 (WIS). Jamaica, 1906, Do- clavate or only slightIy swolIen, sometimes not thick-
bbin (NY).Leicester, 19 15, Merrill 864 (US). West fir- ened or even slightly tapering, (0.8") 1.3-2.3-2.9(-4.4)
ginia. Mercer Co., 1963, Brodo B4596, 4610 & Brodo p wide (S= 0.8, N= 4, n= 10), without internal pig-
(CANL). ment but sometimes with a distinct external hood of
greenish or brownish pigment. Spores acicular or occa-
sionally long-bacillifom, straight or slightly curved, oc-
19. Bacidia reagens WIalrne casionally sigrnoid, not coiled in ascus, (22-)29-37-59(-
69) pm long ( S 14, N= 4, n= 10),
Ark. Bot. 27A (5): 20 (1935). - Type: Brazil, "Rio (1.6-)1.8-2.0-2.6(-2.9) pm wide (S= 0.4, N= 4, n= ,10),
Grande do SuI: Santo Angelo pr. Cachoeira", 1893, (1 1.2-1 15.8-17.8-22.8(-28.0) times as long as wide (S=
Malme 970 (S, lectotype selected here). 3.8, N= 4, n= 10), with (3-)3.5-6.8-12.5(-15) septa (S=
4.0, N= 4, n= 10).
Illwtraaions. Figs 29 (map), 41H. Pycnidia not seen.
Chemistry: No lichen substances detected.
Characaerizdion. Apothecia usually grey-brown to Pigments: Edge, inner upper part of proper exciple
black-brown. Proper exciple with some green-grey to (sometimes), and upper part of hymenium K+ purple to
dirty green and some orange pigmentation. Hy- violet, C+ violet, N+ red. Hypothecium and most of the
pothecium very pale yellowish to brown-orange. Upper inner part of proper exciple K+ intensifying, C-, N-. -
part of hymeniurn dominated by a green-grey to dirty Thalloidima Green in proper exciple and upper part of
green pigment (same as in proper exciple), which reacts hymenium. Laurocerasi Brown in upper part of hy-
C+ violet. Spores acicular (rarely Iong-bacilliform), 22- menium. Rubella Orange in proper exciple and hy-
69 X 1.6-2.9 pm,with 3-15 septa. potheciurn.

Opera Bot. 127, 1996

Fig. 29. Known North American distribution of Bacidia reagens (squares), B, rosella (triangles), and B. russeola (star). Known
world distribution of Bacidin salmonea (dots).

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,

Opera Bot. 117, 1996

Fig. 30.Known North American distribution of Bncidia rubellu.

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

Opera Bot. 127, 19%

(LD). Linn Co., 1972, Rossman 610 W Y ) . Vermont. brown disc; margin paler. Proper exciple very pale yel-
Chittenden Co., 1890, Pringle (COLO). + Brandon, low to orange-brown, rim sometimes slightly darker (to
1921, Merrill 1189 (filed with B. polychroa, WIS). red-brown), never with evenly dispersed, minute crys-
Brattlcboro, Frost (MIN). Virginia. Highland Co., 1975, tals (but sometimes with large clusters), with cell lumi-
Harris 9941 (MIN). + Madison Co., 1966, Wetmore na gradually enlarging towards the edge, or along the
15391 (filed with B, schweinitzii, MIN). Washington. edge with a f distinct, 6-8 cell layers thick zone of en-
Skagit Co., 1989, Ekman L733, L735, L736 (LD). Sno- larged cell lumina. Hypothecium very pale yellow to
hoinish Co., 1994, Tmsberg 21678, 21688 (BG). Wis- pale orange. Upper part of hymenium pale brown-or-
consin. Ashland Co., 1987, Wetmore 6058 1 (MIN). ange to 1 red-brown. Parts of proper exciple and hy-
menium usually K+ purplish. Spores acicular, 41-67 .
2.5-3.7 pm, with 5-9 septa.
22. Bacidia russeola (Kremp.) Zahlbr,
Thallus determinate, rather thin to rather thick, continu-
Cat. Lich. Univ. 4: 239 (1926). - Lecidea m s e o l a ous, f cracked, wrinkled to warted, pale grey or pale
Kremp., Flora, Jena 6 1: 5 18 ( 1878). - Type: Argentina, green-grey. Prothallus lacking or present, bordering the
1872-74, Lorentz & Hieronymus (M, lectotype selected thallus, narrow, black or brown.
here). Apothecia (0.5-)OS-0.6-0.6(-0.9) mm diam. (S= 0.1,
Patellaria phaeolornta Miill. Arg., Flora, Jena 67: 690 N= 2, n= 10), plane or moderately convex, epruinose.
(1884). - Bacidia phaeoloma (Mull. Arg.) Zahlbr., Cat. Disc brown-orange to brown-red to red-brown, or occa-
Lich. Univ. 4: 232 (1926). - Type: Cuba, 1878, Gray, sionally purple-brown to purplish black. Margin paler
Tuckerman: Wright Lich. Cub. 220 (G, lectotype select- than disc, pale brown-orange, pale orange-brown, pale
ed here). to dark red-brown, f brown-red, or purple-brown, lower
Biatora sutur.alis Eckfeldt, Bull. Toney bot. Club 21 : part paler than upper part; raised above disc in young
394 (1894). - Bucidia sutumlis (Eckfeldt) Zahlbr., Cat. apothecia, soon level with the disc, persistent.
Eich. Univ. 4: 245 (1926). - Type: Mexico, "San Luis Proper exciple laterally (43-)48(-52) pm wide ( S 6,
Potosi", 1888, Pringle 183 (PH, lectotype selected N= 2;n= l), without crystals or with radiating clusters
here). of crystals that are up to 7 p.Rim 1 brown-orange or
Patellaria subacerina Miill. Arg., Flora, Jena 69: 308 red-brown in upper part, downwards gradually changing
(1886). - Bacidia subucerinoldes Zahlbr., Cat. Lich. to colourless, with cell lumina gradually enlarging to-
Univ. 4: 242 (1926) (N.B.!The existence of Bacidia su- wards the edge, or along the edge with a f distinct, 6-8
bacerina Vain. 1922 necessitated a nomen novum). - cell layers thick zone of enlarged cells with lumina that
Type: Jamaica, "Gordon-Town. Comm. Joshua 1885" are up to 14 X 5 pm. Lateral interior f concolorous with
(G, lectotype selected here). rim,f orange-brown in upper part, downwards gradual-
Bacidia papontiensis Vain., Dansk Botan. Arkiv 4 ly changing to colourless. Medullary part very pale yel-
(1 1): 20 (1926). - Type: Mexico, "Papantla", 1841, low/orange. Hypothecium very pale yellow to pale or-
Liebrnann 7379 (TUR-V, syntype). ange. Hymenium (69-)75-79-83(-87) pm thick (S= 6,
Bacidia ~ubtesaaceaMalme, Ark. Bot. 27A (5): 23 N= 2, n= 5 ) , lower part colourless, uppermost part pale
-
(1935). Type: Paraguay, "Asuncion, Villa Morra", brown-orange to orange-brown, sometimes with spots
1893, Malme 156 1 (S, lectotype selected here). of red-brown. Paraphyses 0.8-1.6 pm wide in mid-hy-
Bacidia pawuku Malme, Ark. Bot. 27A (5): 33 menium; apices weakly or not at all swollen, sometimes
(1935). - Type: Paraguay, "Colonia Riso. In cortice Pip- even slightly tapering, (0.8-) l .6-1.6-I.7(-2.8) pm wide
eris, in silva clara", 1893, MaIme, bark pieces a and d (S= 0.5, N= 2, n= 1O), without internal pigment. Spores
(S, lectotype selected here). acicular, straight, curved, or sigmoid, straight or slightly
coiled in ascus, (4 1-)53-56-58(-67) pm long (S= 6, N=
Nomencluture. The original material of Bacidia pawula 2, n= 10), (2.5-)2.9-3.0-3.1(-3.7) pm wide (S= 0.4, N=
Malrne, a single collection in S, is heterogenous. Two of 2, n= 10), (14.3-) 17.8-18.9-20.1(-27.0) times as long as
the four bark pieces (marked "a" and "P) agree well wide (S= 3.6, N= 2, n= 10), with (5-)6.6-7.0-7.3(-9)
with the diagnosis, whereas the other two (marked "b" septa (S= 0.9, N= 2, n= 10).
and "c") represent a different species of Bacidia S. str. Pycnidia not seen.
with pale greenish epithecium and short, 3-septate, ba- Chemistry: Trace amounts of atranorin.
cilliform spores. Pigments: Edge of proper exciple and upper part of
hymenium K-l- purplish, N+ orange, other parts K+ in-
Illwtrations. Figs 29 (map), 42B. tensifying, N- or N+ intensifying. - Laurocerasi Brown
in proper exciple and hyrnenium. Rubella Orange in hy-
Churucteriza~ion.Apothecia with brown-orange to red- pothecium, proper exciple, and upper part of hymenium.

Opera Bot. 127. I996

Ecology. Very little is known about the ecology of this Acer macrophyIIum, alt. 10 m, 1989, Ekman L753 (LT),
species, except that one of the three specimens was col- holotype).
lected on Celtis laevigata. In Brazil and Paraguay, this
species occurs in open forests (Malme 1935). A simili Bacidia rosella differt apotheciis obscuriori-
bus, conidiis non septatis; strato crystallifera epithecii
Dis~ibution.Collected twice near Brownsville, Texas, caret. Etiarn B. arceufinae similis a qua differt conidiis
with a 56-year interval, and once in an unspecified lo- subrectis et apotheciis pigmento fulvo carentibus.
cality in Texas (not on the map). - Widely distributed in
the neotropics. There are several finds of this species EQmology. Salmoneus, salmon-pink.
from Cuba, and it may well occur in southern Florida as
well. Illustrations. Figs 29 (map), 42C, 43H.

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,

Opera Bot 127, 19%

N= 27, n= S). Prothallus lacking or sometimes present al wall. Rubella Orange in hypothecium, proper exciple,
between discrete areoles/granules, thin, endosubstratal, pycnidial wall, and sometimes in hymenium. Scllwein-
white to pale grey. itzii Red often in hypothecium, proper exciple, and pyc-
Apothecia (0.5-)0.6-0.9- 1.4(- 1.8) mm diam. (S= 0.3, nidial wall.
N= 27, n= 5), f plane when young, later becoming
convex, usually epruinose, rarely with thin white prui- Ecologv. On the trunks of trees, sometimes branches of
na on margin of young to medium-aged apothccia. trees, shrubs, and vines, mainly in shady and humid local-
Disc in most cases either orange-brown or dark purple- ities, but sometimcs also in more or less open habitats
brown to purple-black to blue-black to pure black, such as forest edges and solitary trccs. The cormnonest
rarely pink to greyish pink, bluish or purplish grey, phorophytes are Acer spp., Fagus grcrnd$olin, Liqui-
palc brown, grcy-brown, or grcyish blue. Margin usu- dambar s!yrac~~ltia, Li~iOu't'?ldrOnhilipifra, Quercus
ally concoluruus with or paler than disc, rarely slightly spp., and Tltuja occidentc~lis,but it has also been found
darker than disc, rather thick: level with or raised on Aesculus sp., Be!da spp., Carpinll,~caroliniuna, Car-
above disc in young apothecia, later level with the ya spp., Celti.7 laevigata, Cephalanlhu.~occideatalis,
disc, and finally excluded. Chionanthws virginicus, Cornus spp., Cra!uegus sp.,
Proper exciple laterally (52-)87(-138) pm wide (S= Euonymus uinericatn~s,Frminu.7 spp., Humumelb vir-
25, N= 27, n= l), sometimes with radiating clusters of giniana, Ilex sp., Juglcrras spp., Juniperus virginiana,
up to 7 pm wide crystals. Rim almost colourless or pale Mngnoliu spp., !tlums rlrbrn, Nyssa spp., Osatya virgira-
yellow to orange-brown to dark brown to dark red- ianu, Piceu sp., Plut~musocciclentalis, Populus tremll-
brown, often bluish or greenish near the hymenium, ceIl loides, Salix spp., Suyium seb$eeuum, Sns- safras aalbi-
lumina gradually slightly enlarging towards the edge, dum, Taxodiurn Jistichutn, Tecoina stans, Tilin
near the edge up to 11 X 4 pn. Lateral interior usually americaraa, Ulmus anrer.icana, and V i ~ i rottind$jIiu.
s
darker than rim, pale yellow/brown to orange-brown to
dark red-brown to brown-black to red-black. Medullary Distribwfion. The castern temperate region of eastern
part darker than or sometimes concolorous with lateral North America, wherc it is thc by far commonest spe-
interior part, orange-brown to dark brown-red to red- cies o f Bucicliu. - Previously thought to bc a North
black. Hypothecium pale orange-brown to orange- American endemic, but it has turned out that it occuls in
brown to dark brown, downwards gradually ~nergingin- eastern Asia (at least China and Japan) as well, from
to the colouration of the propcr exciple. Hymenium where it has been described under at Ieasl thrcc different
(66-)69-92-110(- 118) pm thick (S= 11, N= 27, n= 5), names (see syyonymy).
lower part coIourless except for occasional vertical
streaks with blue-green pigment, upper part either bluc- Disc~.wion.Bacidia schweinilzii is similar to B. uggre-
green (rarely pale blue-green or dark green) or pale or- g a ~ l a B.
, circumspecta, B, igniarii, B. russeolu, and R.
ange-brown, indistinctly or distinctly delimited. Para- subincon~pta.For distinguishing characters, see these
physes 1.2-2.0 pm wide in mid-hymenium; apices f species.
clavate or not at all swollen, {l .2-) 1.6- 1.9-2.7(-4.0) pm Bacidia schweinitzii is the most variable spccics of
wide (S= 0.5, N= 27, n= 5), without internal pigment. Bacidiu in North America. There is a large variation in
Spores acicuIar, straight to curved to sigmoid, (32-)44- thallus structure, from granular to almost smooth,
56-73(-88) pm long (S= 9, N= 27, ii= 10), (1.9-)2.5-2.8- which much parallels the variation seen in B, subin-
3.2(-4.3) pm wide (S= 0.4, N= 27: n= 10), (10.3-)15.5- compta. There is also extensive variation in the pig-
20.2-30.6(-46.7) tiines as Iong as wide (S= 4.6, N= 27, mentation of the apothecia. Most apothecia are elther
n= lO), with (3-14.3-7.9-13.6(-15) septa (S= 2.8, N= 27, black with a blue-green upper hymeniuln and more or
n= 10). Iess red-black hypothecium and proper exciple, or
Pycnidia f immersed in thallus or prothallus, upper- brown with brown upper hymenium and more or less
most part of wall concolorous with apothecia, 75-175 brown hypotheciunl and proper exciple. lntermediate
pm diam. Conidia filiform, curved, non-septate, 9-20 X apotl~eciaare sornetimcs found, and in some cases I
c. 0.7-0.8 pm. have observed brown as well as black apothecia mixed
Chemistry: Usually no lichcn substances, but some- on the same thallus. In most thalli, however, all apoth-
times with atranorin. ecia are either brown or black. Although all combina-
Pigments: Yellowish to orange-brown parts K+ intcn- tions of apothecium pigmentation and thallus structure
sifying, N- or N+ intensifying. Dark reddish brown to occur within all parts of the distribution range, there is
red-black parts K-, N-. Green parts K-, N+ purple with a a clear tendency that individuals with brown apothecia
precipitate of dark blue crystals (in apothecia with k andlor nun-granular thallus are more common in the
blue-grew hymenium). - Bacidia Green often in hy- South than in thc North. An explanation for this may
menium, uppermost part of proper exciple, and pycnidi- be that only a part of the variation within the species
has been involved in the recolonization of the North af- Kantmd (CANL). Hairy Lake, 1902, unknown collector
ter the latest glaciation(s). (CANL). Chelsea, 1907, Macoun 51 (CANL). - U. S.
The Japanese Bacidia abducens (Nyl.) Zahlbr., and A. Alabama. Lee Co. 1893, 1897, Earle & Baker (MW,
the Chinese B. melanocardia Zahlbr. and B. nigra ZahI- NY, US). Pike Co., 1964, Johnsoii (NY).Randolph Co.,
br. should be included in B. schweinitzii. All three have 1987, Bowers & Raynes 12561b-1 (herb. Tucker). Tal-
+
a continuous, non-granular thallus with black apothe- ladega CO, 1900, PolIard & Maxon 239 (US). Tuscaloo-
cia containing Bacidia Green in the hymeniurn. Morphs sa Co., 1987, Bowers & White 13 103 (herb. Tucker).
with granular thallus or lacking Bacidia Green in the hy- Brooklyn, 1860, Beaumont 72309 (FH). Dadeville,
menium are so far not known outside North America. 1900, Pollard and Maxon I30 (US). Arkansas. Newton
The type of Bucidia dbductans (Nyl.) Zahlbr. from Flor- C o . , 1979, Ryan (WWB), 1979, Brodo 23491 (CANL),
ida is a typical specimen of B. schweinihii with black 1988, Harris 21511 (NY). Stone Co., 1988, Harris
apothecia containing Bacidia Green in the hymenium 21618 (NY), + 1988, H m i s 21592 (filed with B, cd+
and a partly smooth, partly granular thallus. frcscla, NY), + 1993, Tmsberg 19258 (filed with B. suf-
fuse, BG). Van Buren Co., 1993, Tansberg 19227,
Additional collections examined. CANADA. Lac Trum- 19236 (BG). Mahern, 1899, Russell 23 (US). Connecli-
blant, 1922, Plitt (US). New Brunswick. Albert Co., cut. Fairfield Co., 1926, Evans 852 (FH, US). Litchfield
1980, Gowan 3051 (CANL), 1981, Gowan 4362, 4989, Co., 1949, Hale 140 (US). New Haven Co., 1925,
5066 & Brodo (CANL), 1989, Ekman L1079 (LD). Musch 342 (FH, US). New London Co., 1925, Nichols
Charlotte Co., 1989, Ekman L1089, L1090, L1094 & Musch 576 (FH, US). D. C. Lehnert (US). Delaware.
(LD). Nova Scotiu. Kings Co., 1972, Brodo 18908 Sussex Co., 1966, Brodo 9540 & Brassard (CANL, CO-
(CANL), 1989, Ekman L1042 (LD). Queens Co., 1989, LO), 1966, Brodo 9563 & Brassard (CANL), 1989, Ek-
Ekman L1045, L1046, L1047, L1055 (LD). Ontario. inan L1047, L1 101, L1 102, L1 104 (LD). Wilmington,
Brant Co., 1944, Cain (CANL). Dufferin Co., 1984, 1890, Commons 42 m). Florida. Calkins (FH, US).
Wong 3595, Krug & Warren (CAWL), Lake Superior 1886-87, Calkins (MW). Alachua Co., 1915, Nelson
Prov. Park, 1989, Ekman L996, L997, L999, L1007, (FH, US), 1993, Ekman L1 145, L1 146, L1 147 (LD).
L1010, L1011, L1012, L1013, L1015, L1016, L1017 Holmes Co., 1993, Ekrnan L1 138 (LD). Leon Co.,
(LD). Frontenac Co., 1941, Cain (CANL, COLO, NY, 1990, Tucker 29736, 29737, 29739B, 29771, 30169
US). Hastings Co., 1989, Buck 17530, 17539 (NY). (herb. Tucker), 1991, Tucker 303 18 (herb. Tucker). Lib-
Lanark Co., 1986, Wong 3975 (CANL). Muskoka Dis- erty Co., 1993, Ekman L1214, L12 16 (LD). Nassau Co.,
trict, Univ. of Toronto Forest, 1964, Denison 487 1987, Harris 21 153 (NY). Orange Co., 1993, Ekman
(CANL). Peel Co., 1946, Cain (CANL, OSC, SFSU, L1 165 (LD).Wakulla Co., 1993, Ekrnan L121 1, L1212
UBC). Renfrew Co., 1973, Brodo 20571b (CANL), (LD). Jacksonville, 1888, 1889, 1890, 1892, Calkins
1975, Wong 1832, Ireland, White & Dugal (CANL), (CANL, FH, NY, US). Sanford, 1911, 1912, 1914,
1985, Brodo 25255 (CANL), 1985, Wong 3805, 3810 1915, Rapp (FH, US). Six Mile Creek, Calkins (COLO,
(CANL). Simcoe Co., 1964, Denison 489 (CANL), NY). Georgia. Chatman Co., 1974, Tucker 12224a
1967, Krug 223, 224a (CANL). York Co., 1941, Cain (herb. Tucker). Rabun Co., 1981, Brodo 23888
(CANL, MY), 1946, Cain (COLO), 1953, Cain (CANE). (CANL). Towns Co., 1976, Harris 1 1210 (NY). White
Algonquin Prov. Park, 1900, Macoun (CANL), 1972, Co., 1976, Harris 11148, 11168 (NY). Savannah, 1917-
Dickson 91 (CANL). Belleville, 1868 (CANL), 1892, 18, Rervey 78, 89 (FH, US). Illinois. Union Co., 1968,
probably collected by Macoun (CANL). Belleville and Skorepa 4122 (US). Canton (NY). Idiana. Owen Co.,
Ottawa, 1881, Macoun 63 (CANL). Carleton Place, 1988, Hanis 21824 (NY). Putnarn Co., 1893, Under-
1900, Macoun (CANL). Carp, 1907 (CANL). Emo, wood 13 (NY), 1970, Brodo 17084 (CANL). Iowa. Fay-
1901, Fink 842 (MM).Kingston, 1864, Drurnmond etfe Co., 1894, Fink (MW). Ke~eatucb.Calloway Co.,
(FH). Rritannia, 1904, Macoun 173 (CANL), 1904, 1975, Buck B34 (MW). Madison Co., 1941, Allen 44,
probably collected by Macoun (CANL). Ottawa, I89 1, 208, 5 15 (US). Rockcastel Co., 1927, Fink (CANL).
Macoun, 83, 484, 512 (CANL), 1892, Macoun 2815, Louisiana. Acadia Par., 1894, Langlois 1020 (NY).Al-
2821 (CANL), 1892, Macoun 342 (US), 1895, Macoun len Par., 1986, Tucker 27296b (herb. Tucker). + Ascen-
(US), 1902, Macoun 192 (CANL, COLO, US). Quebec. sion Par., 1981, Tucker 21473 (filed with B. SUJL-~,
Argenteuil Co., 1984, Brodo 250 17a (CANL). Brome herb. Tucker). Assumption Par., 1981, Tucker 21638c
Co., 1965, Denison 29 (CANL). Gatineau Co., 1968, (herb. Tucker). Caldwell Par., 1984, Tucker 26355
Brodo 13340 (CANL), 1969, Brodo 16331 (CANL), (herb. Tucker). East Baton Rouge Par., 1971, Tucker
1971, Hanes (CANL), 1973, Brodo 19134 (CANL), 9087 (herb. Tucker), 1974, Tucker 1 1835 (CANL, CO-
1989, Ekman L1024, L1026, L1027, L1030 (LD). Pon- LO, MIN, UBC), 1974, Tucker 1 1835a, 1 1836b, 1 1987
tiac Co., 1969, Brodo 14849 (CANL). ComtC Portneuf, (herb. Tucker), 1974, Tucker 12571 (herb. Tucker,
1975, Brodo 20678b (CANL). Sherbrooke Co., 1972, CANL, MIN), 1975, Tucker 13991 (herb. Tucker),

Opcm Boi. 127, 1996

morc 60236, 60245, 60828, 60863 (MIN). Bayficld Co., because Nylander himself annotated this specimen
1965, Wctinore 13378 (COLO, MIN, US), 1987, Wet- "Leciclea subincurplptct". it is a well-developed and typi-
more 61 006, 6 1183 (MTN). Douglas Co., 1946: Thorn- cal specimen, and there are isotypes in several other her-
son, Thornson: Llch, wisconsincnscs 36 (CANL, CO- baria.
EO: MW: US). Regarding the nomenclaturc of thc nalnc Baciclia he-
get.~chw~tleri,see B. ~~ernaferu.

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-

Oprrn Bot. 127, 1996

meniuin, uppermost part of cxciplc, and so~nctimesalso Wells, 1 978, Bird 8r Thoillsoil (W IS). Oniurio. Algoma
upper part o r hypolhecium. Lauroccrasi Brown in lly- District, Nagaga~nis Prov. Park, 1968, Brodo 13462
pothecium, proper exciple, pycnidial wall, and apices of (CANL). Cochranc District, Moose Factory. 1969, Bro-
the parapliyses. do 14682, 1 4 7 1 7 ~(CANL). Lcnnox & Addington Co.,
1972, Wong S j j c , Shchepanek k Lcy (CANL). Thun-
E c n l o ~ v .Mainly on rough bark of tree-trunks, prcfcra- der Bay District. 16 km SW u f Suomi, 1972, Garton
hly in shady and humid forests, but sometimes a l w in 15327, 15337 (CANL). Slatc Islands, 1977, Wetmore
more or less open places like thin forests, forest edgcs, 28593 (MIN), 1977, Wellnore 28596 (CANL, MIN).
a i d solitary trces. Ft~a.xinu.7 spp., Poptrlus spp, and Thzr- YUICOFI.Tom Creek Tower Area, 1981, Rosic 71 1
ju urciclentnlis are the most c o ~ n m o ~ phorophytes,
i but it (CANL). - U. S. A. Alaska. C. 3U0 km NNW o r Fair-
has also been ,collected on Abies sp., Acer saccltnrtmt, banks,] 989. Thor 9081 (S). Chisik Lland, 1987, Talbot
Alnus sp., Qlret.czi.s g(irt;vunu, Snlix sp., U ~ I I Zanlet-ica-
~IS X70 14-2a ( W IS). Wrangcl-St. Blias Nat. Presenle, c.
nu, and lignum of Pinrir sp. 390 km E of Anchorage, 1989, Thor 879 1 (S). A r ~ ~ o a a .
ApacIic Co., 1975, Nasli 11739 (COLO). Colorado.
Disfriblition. IH the Great Lakes region, in tlie tnoiltane or Boulder Co., 196 1, Anderson 15 13 (CANL, MIN).
arctic areas from southe1.n Arizona to Alaska, and one i s o n1 955, Weber (COLO). lows. Fayette,
C u ~ ~ ~ ~ Co..
outlicl. in thc Willamette Valley oforegnn. - Europe. 1894, Fink (NY). ~ M i c h i ~ rGogebic
n. C O . , 1989, Ekman
L97 1, L976, L978, L979 (LD), + 1989, Ekrnan L972
Discussion. Bcluidia sl~bincortzp!cris superficially very (filed wilh B, pokchroa, LD). Keeweenaw Co., 1983,
siinilar to B. sch~veit~itzii. However, in B. schweitritzii, Weirnore 4732 I , 47929 (MIN). Ontonagoil Co., 1989,
the hypotl~eciu~n and at least a largc part of the proper Ekman L987 (LD). Minne.~o!o.BeItrami Co., 1900, Fink
exciple is continuousIy and evenly darkly pigincnted, 596 (MTN). Clearwater Co., 1972, Wetmore 20343
whereas in B. szlhincomplu, the hypotheciu~nis k brown (MIN). Cook Co., 1986, Welnlorc 57107 (MIN, NY).
in a distinct zonc and much paler below. Lakc Co., 1989, Ekman L952, L9.59, L961 (LD). + Lake
Bacidia subir~compta is a well-defincd species, al- of thc Woods Co., 1901, Fink 400 (filed with B, sufuuscu,
though with an amazing amount of variation in spore MIN). Ottcr Tail Co., 1900, Fink 376 (MIN}. Roscau
shape and thallus appearance. The variatioil in thallus Co., 1901, Fink 234, 323 (MIN). St. Louis Co., 1901,
appearance is similar to that exhibittd by tlic close rela- Fink 1468 (!MIN), 1986, Wetmore 57637, 57642 (MW).
tive B. .rchweinitzii. The variation in spurc slyape, how- iMuntuna. Ravalli Co., 1980, McCune 10824 (LD). Ore-
ever, is unsurpassed by any other species of Bacidiu that gon. Benton Co., 1989, E h a n Lh95 (LD).
I know of. Withln B. szlbin~111upt~z. 4-celled and fusi-
form as well as multicelled and acicular spores can bc
found. The variation in spore shape is always modcratc 26. Bacidia suffusa (Fr.) A. Schneid.
within the same apotlieciu~n,but in some specimens, ba-
cillifonn sporcs arc found in young apothecia, whereas Guide Study Lich. 1 10 ( 1 898). - B i a ~ o r usl!j/usa Fr.,
acicular spores are found in uld apothecia. Consequent- Syst. Orb. Veget. 285 (1825). - - Typc: U. S. A., Minne-
ly, it seems likely thal a t least part of thc variation is due sota, Mahnomen Co., "Easl of Washington Lake (18
to the age of the apotheciun~in which the spores are miles NE of Mahnornen). 111 swamp arca with balsam
produced. The remainder of the variation must bc due to fir, sprucc, ta~narack", 1976, Welmorc 26404 (US, neo-
cither geneticaI variation, or environmental influcnce. type sclcctcd here; CANL, MlN, isotypes).
Since specimens with very difTeret~tspore types can bc Leoidea sprrdiicecr Ach., Sy~iops. Lich. 34 (1 8 14),
found in similar habitats, and since specimens with the nom, rcjic. prop. - Bacidiu spu~liucw (Ach.) ~ a h l b r . ,
same spore type can he found in very diKerent habitats, Denkschr. math.-naturw. KI. Akad. Wiss. Wicn 83:
it seems probablc tllat a large portion of the variation is I28 (1909). - Typc: North America, specimen marked
actually genctically bascd. ''Le~~itf.
.spu~li~euns.", ex hcrb. Mdllenberg (PH, lecto-
Pa~tiaIly pigmcnt dcficie~~tspecimens wltl~ pink- type selected here).
brown apothccia have been collected a few times in
North America. Entirely pig~ncnt deficient specimens ~Vontcnclu!~rr.c.I have bee11 unable to trace any original
with pale pink apothecia have so far ooly bceil observed nlatcrial of Baciu'io ~ujfu.su.Therefore, a ncotype is se-
in Europe. lected, a wcll-developed collection kept in three of the
major hcrbaria in North America.
Collecfions exornined. C A N A D A . Brirish ColllmRiu. Lecidea spadicea Ach. was, according Lo thc intro-
Wells Gray Provincial Park, 1989, Ekman L794 (LD). duction o r .Yjvlupsis Methndiccr Liclrenzim. based on ma-
Nurthwest Terr-ifories. Mackenzie District, Little Gull terial collected by C. H. E. Muhlenherg. There is no
River, 1966, Scottcr 8274c (H). C. 5 ~ n SE
i of Nornlai~ original material in l-I-ACH, BM, or UPS. The only ex-
 Fig. 33. Known world
distribution of Bacidia
suffusu.

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.

Opera Bol. 127, 19%

108
Disc pale yellow-brown, yellow-brown, brown, deep or- cidentalis, Tilia amei-icana, Ulmus americana, and an-
ange-brown, purple-brown, brownish purple, dark pur- thoxylupn clava-herculis.
ple, brown-black, purple-black, ox pure black, some-
times mottled with different colours. Margin Distribution. In the eastern temperate region, but com-
concolorous with disc or sometimes slightly darker, f monest in the Great Lakes area. In addition, there are
thin, level with or slightly raised above disc in young two old and probably mislabelled specimens marked
apothecia, persistent, excluded only in some old and "Idaho" and "British Columbia", respectively. - Endem-
convex apothecia. ic.
Proper exciple laterally (52-)67(-86) pm wide ( S =
12, N= 21, n= 1j, usually with radiating clusters of up Discussion. The abundance of crystals in the proper ex-
to 10 pm wide crystals. Rim pale yellow to yellow- ciple, the pigmentation, and the zone of enlarged cells
brown to orange-brown to dark-brown to black-brown, along the edge of the proper exciple indicate that B. suf-
often darker in upper part than in lower part, along the fuse is more cIosely related to the tropical species B.
edge with a 4-6 cell layers thick, distinct zone of cells cumpaleu, B. fisconigrescens (Nyl.) Zahlbr, and B. mil-
with enlarged lumina that are up to 12 p long and up legraaa (TayIor) Zahlbr. than to any of the species oc-
to 6 p wide. Lateral interior paler than or f concolor- curring within the distribution range of B. s u f f i a . For
ous with rim (rarely darker than rim). Medullary part differences between B. cumpalea and B. sufisa, see the
paler than lateral interior part, almost colourless to former species. B. millegrana can be separated from B.
pale yellowish. Hypothecium f yellowish, concoIorous sufisa by the presence of a narrower, two cell layers
with or slightly darker than Iateral interior of proper thick zone of enlarged lumina along the edge of the
exciple. Hymenium (7 1-)74-94-115(-127) pm thick proper exciple, and by the absence of crystals and pruina
(S= 12, N= 21, n= 5), lower part colourless except for in the apothecia. B. fisconigrescens is very similar to B.
occasional vertical streaks with f orange-brown pig- sufisa, also regarding the zone of enlarged cells along
ment, upper part pale yellow to pale brown-orange to the edge of the proper exciple, but is mainly separated
orange-brown to red-brown to black-brown, pigmenta- by the presence of k evenly distributed, minute crystals
tion indistinctly or sometimes distinctly delimited. Par- in the proper exciple and by the darker pigmentation
aphyses 1.2-2.0 pm wide in mid-hymenium; apices (apothecia f black).
narrowly clavate or not at all swollen, (1.2-)l .5-1.R- Although perhaps not closely related, B, sufusa may
2.0(-2.8) pm wide (S= 0.3, N= 21, n= 5), without inter- be conhsed also with B. rosella (see that species), B.
nal pigment. Spores acicular, straight, curved, or sig- dzgi-acta, and B. polychroa. The latter two species can
moid, sometimes coiIed in ascus, (38-348-57-69(-9 1) be separated by the brown, K+ purple-red hypothecium.
p long (S= 8.0,N= 19,n= lO), (2.5-)2.7-3.2-3.6(-4.3)
pm wide (S= 0.4, N= 19, n= IO), (10.3-114.4-17.9- Additional collections examined. CANADA. British Co-
21.9(-29.5) times as long as wide (S= 3.4, N= 19, n= lumbia. Vancouver Island, 1875, Macoun 2808 (CANL,
101, with (3-)6.3-7.9-12.6(-17) septa (S= 2.5, N= 19, probably mislabelled). Ontario. Renfrew Co., 1983,
n= 10). Brodo 25122, Brodo, Thomson & Thornson (CANL).
Pycnidia f immersed in thallus or prothallus, upper-
, Algonquin Park, 1900 (CANL). Belleville and Ottawa,
most part of wall concolorous with apothecia, 100-125 1881, Macoun (US). Belleville, 1882, Macoun 2805
pm diam. Conidia Bliform, curved, non-septate, 10-27 (CANL), 1889, Macoun 3049 (CANL), + 1882, proba-
X c. 0.8 pm. bly collected by Macoun (filed with B. poiychrou,
Chemistry: Atranorin (sometimes only as a trace). CANLj. Carp, 1907 (CANL). Edmonton, 1893, White
Pigments: Yellow to orange parts of apothecia K+ in- 320 (CANL). Navan, 1902 (CANL). Ottawa, 1884, Ma-
tensifying, N+ intensifying, 1 dark brown parts K+ pur- coun 2804 (CANL), 1891, Macoun 273-74 (CANL),
plish, N+ orange-red. - Laurocerasi Brown in proper 1892, Macoun 2807 (CANL), 1896, Macoun (US),
exciple, hyrnenium, and pycnidial wall. Rubella Orange 1900, Macoun 22 1 (CANL, COLO, US), 1902, Macoun
in proper exciple, hypothecium, hymenium, and pycnid- 192 (CANL), 1905, Macoun 19 1 (CANL, US), + 1891,
ial wall. Macoun 102 (filed with B. poiychroa, US), + 1891, Ma-
coun 275-276 (filed with B, polychroa, CANL) + 1901,
Ecology. Mainly on the rough bark of tree trunks in Macoun 192 (filed with B. schweinitzii, CANL). Pres-
shady and humid forests (often swamp forests). There col?, BiIlings (CANL). Quebec. Comtk Argenteuil,
are a few finds on rock. Many corticolous finds are on 1984, Brodo 25032 (CANL). Gatineau Co., Brodo
Acer spp. and Frminus spp., but it has also been col- 16373 (CANL), 1973, Brodo 19122 & Hawksworth
lected on Carpinus caroliniana, Fugus grundijiolia, Lir- (CANL). Buckinghamn, 1896, Macoun 391 (CANL). -
iodendron tulipifem, Liquidcambar siyraciflua, Populus U. S. A. Arhrarus. Stone Co., 1993, Tansberg 19258
sp., Quercus alba, Q. macrocappa, Sulk sp., Thuja oc- @G). Florida. + Jacksonville, 1888, Eckfeldt (filed

Opera Bot. 127. 19%

with type specimen of Lecidea subspadicea Nyl.= B. 3 1371, 34234, 34255, 34932, 34935 (CANL, MIN),
polychroa, H-NYL), Idaho. 1889, Leiberg (US, proba- 1979, Wetmore 38899,40219 (MW), + 1978, Wetmore
bly mislabelled). Illinois. Calkins 56 my). + Calkins 3 1334 (filed with B. rubella, Mm). Winona Co., 1980,
114 (filed with B. polychroa, NY). Cook Co., 1897, Brako 2520 (MIN). Yellow Medicine Co., 1899, Fink
Calkins (NY).Canton, unknown collector (NY), Wolf 529, 536, 540 (MIN). Cobden, 1878, Earle (NY).Red
(US), + unknown collector (filed with B. polyckroa, Lake, 1900, Fink 918, 936, 1015, 1041 (MM). St.
NY), + Wolf 17 (filed with B. polychroa, US), + Wolf Cloud, 1897, Morgan (US). Missixsippi. Madison Co.,
(filed with B. polychroa, US). Deer Park, Calkins 113 1971, Tucker 9856 (herb. Tucker). Sharkey Co., 1978,
(NY). Iowa. Bremer Co., 1894, Pink (MW, NY, US), Johnson 7038, 7039, 7044, 7045, 7046, 7066, 7083,
1898, Fink (MM).Fayette Co., 1894, Fink (CANL, 7084, 7089, 7116, 7121 (NI'). New Jersey. Andover,
MW, NY, US), 1894, Fink, Cummings: Decades N. unknown collector (NY). Closter, 1876, Austin 723
Am. Lich. 171 (FH, NY, US), + 1894, Fink (filed with (NY), Austin (NY). Springdale, 1937 (NY). New York.
B. polychroa, US), + 1894, Fink (filed with B. schwein- Belleville (NY). Buffalo, Wilson (NY, US). East Gal-
itzii, MW), 1895, Fink (CANL, NY, US). Henry Co., way, 1886, 1893, Burt (FH). Fishkill, 1850, Russell
1898, Savage (Mm). Iowa City, Shimek (MIN). Fay- (FH). Jamesville, Willey (US). Oneida, Merrill (FH).
ette, 1894, Fink, Curnmings et al.: Lich. Boreali-Am. Shushan, 1907, Fink (NY). North Carolina. Swain Co.,
102 (COLO), 1894, Fink (MIN, NY). Near Fayette, 1987, Harris 20979,21000 (NY).Ohio. + 1897, Bogue
1894, Fink (CANL, MW). Kansas. Manhattan, 1893, 874, S. n. (filed with B. polychroa, NY). Willson 841
Waters & Reed 36 (NY). Louisiana. Ascension Par., (NY). Columbus, 1893, Bogue 215 (NY). + Orwell,
1981, Tucker 21468, 21473 (herb. Tucker). East Baton 1891, Bogue, Merrill: Lich. exs. 17 1 (filed with B. poIy-
Rouge Par., 1980, Tucker 21335 (herb. Tucker). Iber- chroa, COLO). Pennsylvania. Willey (US).South Caro-
ville Par., 1974, Tucker 13317b (herb. Tucker). Sabine lina. Curtis (NY) . Chester, Green (NI'). Wisconsin.
Par., 1970, Tucker 8883 (CANL, COLO), + 1970, Ashland Co., 1987, Wetmore 60307, 60902 (MIN),
Tucker 8883 (filed with B. dtgrocta, herb. Tucker). St. 1965, Denison 862 (CA_NIJ,Sl.LCr.q EaJ&,J&Ki',&gk:
Belena Par., 1972, Tucker 10072 (herb. Tucker). West er 138 (NY). dd m d - h a m m b g d m o
Feliciana Par., 1981, Tucker 21525 (herb. Tucker). +
Chenal, 1894, Langlois (filed with B. schweinitzii, US).
Duchamps, 1894, Langlois 860, S. n. (US). Maryland. 27. Bacidia verrnifera (Nyl,) Th. Fr.
Montgomery Co., 1902, Maxon 622 (US), 1907, Fink
87 (US). Massachuseats. New Bedford, Clarke (FH), Lichenographia Scand. l: 363 (1874). - Lecidea vermif-
Willey (PH, US). Michigan. Alger Co., 1987, Wetmore , era NyL, Bot. Notiser 1853: 98 (1853). - Type: Swe-
58764, 59587 (MIN). Antrim Co., 1965, Swanson den, "Holmiae (Marieberg) ad quercus", Nylander (H-
(MICB). Delta Co., 1976, Harris 11890 (CANL). Dick- NYL 17507, lectotype selected here).
inson Co., 1971, Harris 7606 (MIN). Gogebic Co., Bacidia hegetschweileri auct., non (Hepp) Vain.
1975, Harris 9998a (MM), 1975, Buck B230 (NY),
1989, Ekman L966, L967, L974, L984 (LD).Onton- Nornencla~re.The name Bucidja hegetschweileri has
agon Co.,, 1989, Ekrnan L994, L995 (LD). Minnesota. often been used for this species. This name first ap-
Blue Earth Co., 1899, Fink 29, 37, 130, 138, 141 peared as Lecidea hegetschweileri Hepp (a nornen nu-
(MIN), 1899, Gardner 77, 79, 97 (NY). Brown C o . , dum) in Hepp's Systematische Sammlungen no. 212 in
1899, Fink 274 (MIN). Cass Co., 1976, Wetmore 1852. In the following year, the name was validly pub-
25951, 25969 (MIN). Clearwater Co., 1963, Hale lished as Biatora atrosanguinea var. hegetschweileri in
22819 (US), 1972, Wetmore 20327, 20354, 20359 another of Hepp's exsiccates, Flechten Europas no. 23.
(MIN), 1974, Wetmore 22826 (MIN), 1976, Wetmore In the protologue, the collection in Systematische
25820 (MIN), 1978, Wetrnore 36328, 36336 (MIN). Sammlungen, a collection in herb. Schaerer (Flotow:
Hennepin-Ramsey Co., 1896, Fink 194 (MW).Bub- Lich. exs. no. 91c), and an unspecified number of col-
bard Co., 1957, Anderson (COLO). Itasca Co., 1977, lections from the vicinity of Ziirich in herb. Hepp. are
Trana 3 130 (MIN). Koochiching Co., 1979, Wetmore cited. All these specimens are syntypes in addition to
3741 6 (MIN, NY). Lake of the Woods Co., 1901, Fink Flechten Europas no. 23. The protologue also includes
400 (MIN). Olmsted Co., 1975, Wetmore 24163 line drawings of spores that are 3-7-septate and strnight
(MIN), 1977, Trana 2898 (MINI. Otter Tail C o . , 1900, or only slightly curved.
Fink 345 (MIN). Pine Co., 1983, Schuster 258 (CANL, I have not been abIe to trace any specimen of System-
MIN). Rice Co., 1975, Wetmore 24224 (MIN). St. atische Sammlungen no. 212, nor have I been able to
Louis Co., 1902, Fink, Hibbard Photo 5443 (MIN), find any schedae. A stray specimen of this "exsiccate"
1973, Wetmore 21696 (MM),1978, Wetmore 32669, in UPS (no. 126) shows a handwritten label. Clearly,
32694, 35494, 35502, 35533 (MIN), 1978, Wetmore Systematische Samrnlungen was not a true exsiccate.

Opera Bot. 127, 1996

110
Lynge (1915) saw the series in B, but according to H.
Sipman (in Iitt.) it is no longer present (probably de-
stroyed). All syntypes seen by me (BM, G,LD, M, STR)
belong to Bacidia subinconap~a,except one (one of the
four in G) which belongs to B. hegetschweibri auct.,
i.e., B, vermifera. Biatora atrosanguinea var. hegetsch-
weileri cannot be typified with a specimen of B, heget-
schweileri auct., however. The line drawings of spores
show that Hepp's intention was to describe the lichen
that belongs to B. subiplcomptu. A lectotype belonging
to B. hegetschweileri auct. would thus be in serious con-
flict with the protologue due to the very different spores
(Art. 9.13b).
The use of Bacidia hegetschweileri was established
by Vainio (1922), who was also the first to combine the
name to the species level. Bacidia vei-inifera, on the oth-
er hand, was validly published as Lecidea vermifera in
1853, and has since been used in a sense consistent with
the lectotype selected here. Indeed, the name Bacidia
vermfera has been in use for this species, parallel to
Bacidia hegetschweileri, as late as in the 1970s. Conse-
quently, B, vermifera is the oldest name at the species
level and has never been used in a sense conflicting
with its type.
Since the name Bacidiu hegetschweileri is known by Fig. 34. Known North American distribution o f Bacidia
few, and since the name B, vermifem has been used for vermifera.
the same taxon also in relatively recent times, I believe
the best way to minimize the confusion is to discontinue
the use of Bacidia heebchweileri and reduce it into matt, distinct, raised above disc in young apothecia,
synonymy with B. subincornp~a (see synonymy under soon level with the disc, persistent or only rarely ex-
this species). Accordingly, the use of B. vermifera cluded.
shouId be resumed. Proper exciple laterally (26-)35(-45) pm wide (S= 6,
N= 8, n= l), without crystals. Rim black-brown or dark
IIlustratiow. Figs 34 (map), 42H. red-brown, downwards sometimes gradually changing
to k red-brown or colourless, along the edge without a
Characterization. Apothecia (almost) black. Proper ex- layer of enlarged cells. Lateral interior slightly paler
ciple laterally red-brown to black-brown (rim darkest). than or sometimes concolorous with rim. ~ e d u l l G part
Rypothecium pale. Upper part of hymenium dark red- colourless or very pale brownish. Hypothecium colour-
brown to black-brown; much of the pigment confined to less or very pale brownish. Hyrnenium (45-)48-56-64(-
the wall of the parahysis apices. Spores bacilliform to 68) pm thick (S= 6, N= 8, n= 5 ) , lower part colourless
clavate, strongly curved or sigmoid, 14-31 X 2.1-3.1 except for occasional vertical streaks with brown pig-
m. ment, upper part dark red-brown to black-brown, indis-
tinctly delimited. Paraphyses 1.2-2.4 pm wide in mid-
Thallus indeterminate, f thin, either continuous, warted, hymenium; apices f clavate (or sometimes a few
f rimose to areolate, ox discontinuous, of discrete or slightIy or not at all swollen), (1.2-)3.1-3.5-4.1(-6.1) pm
contiguous, convex, warted, sometimes subsquamuIose, wide (S= 1.1, W= 8, n= 10), many with brown pigment
oRen poorly delimited areoIes, white to pale grey to in the waIl. Spores bacillifom to clavate, usually
grey to brown-grey. ProthalIus lacking or present be- strongly curved or sigmoid but sometimes only slightly
tween discrete areoIes, thin, white to pale grey, endo- curved or almost straight, coiled in ascus, (14-)16-20-
substratal. 27(-3 1j long (S= 4, N= 7,U= 10j, (2.1 -12.3-2.6-2.9(-
Apothecia (0.2-)0.3-0.3-0.4(-0.5) mm d i m . (S= 0.06, 3.1) pm wide (S= 0.3, N= 7, II= 101, (5.2-)6.3-7.8-11.2(-
N= 8, n= 5 ) , f plane, remaining so or becoming moder- 13.5) times as long as wide (S= 2.2, N= 7, n= 10), with
ately convex or rarely markedly convex, epruinose. Disc (3-)3.0-3.4-4.3(-7) septa (S= 0.9, N= 7, n= 10).
usually pure black, seldom partialIy or entirely dark pur- Pycnidia half-immersed in thallus, uppermost part of
ple-brown. Margin concolorous with disc, glossy or wall concolorous with apothecia, 40-75 ym d i m . CO-

Opem Bot. 127, 1996

nidia of two types: ( I ) ellipsoid or short-bacilliform, i.e., fungi not producing ascomata and entirely de-
non-septate, 2.8-3.6 X 1.2-1.6 pm, and (2) bacilliform, pendent on conidia for their dispersal. It is clear from
straight or slightly curved, non-septate, 6-9 X 0.5-0.8 the context that the authors did not associate W. fisari-
m. oides or L. gyulectiformis with the Bacidia phacodes
Chemistry: No lichen substances detected. group.
Pigments: All brown parts of apothecium and pycnid- The type specimen of Woessia j%sarioides belongs to
ial wall K+ purplish and N+ orange-red. - Laurocerasi Bn. arnoldiana (according to Serusiaux 1995 and iso-
Brown in proper exciple, hypothecium (small amounts), type in priv. herb, of Tschermak-Woess!), whereas the
hymeniurn, and pycnidial wall. type specimen of Lichingoldia gyalectifornais belongs
to Bacidina inundata (Fr.) VGzda. The describing au-
Ecology. Preferably in shady and humid sites such as thors interpreted the type collection of Lichingoldia gy-
canyons, but sometimes also in semiopen habitats. It is akctiformis as a mixture of this species and Bucidinu
often found near streams and lakes. Known phorophytes inundata. The possibility that these names were conspe-
include Populus angustfolia, P. tremuloides, P, tricho- cific was actually discussed , but it was ruled out due to
carpa, Quercus Pnacrocarpa, and SaIix sp. ''the different appearance of the thalli" and since "the
conidiomata are not found intermixed with the ascoma-
Distribution. Known from a few scattered localities in ta". In the type material (isotype in UPS!), I find the
Alberta, Saskatchewan, South Dakota, Montana, Idaho, apothecia and pycnidia to be mixed on the same thalIus,
Colorado, and California, all at altitudes exceeding which is not dimorphic. Furthermore, the measurements
1000 m. - Europe. of conidia provided by the authors refer, according to a
slide madeby one of the authors and filed with the type,
Discussion. B. verrngera is similar to B, circumspecta to filaments of an alga overgrown by Bacidina inuada-
and B. igniarii but can be separated from both these spe- fa! True conidia are, however, present in the same prep-
cies by the absence of a green pigment (Bacidia Green) aration, and they measure 21-36 pm in length (which is
in the hymeniurn and the proper exciple. Furthermore, normal to Bn. inundata), not 70-85 pm as suggested by
the spores of B, circumspecta are not as strongly curved the authors.
or sigrnoid as in B. vermfera. Twenty-five species are presently recognized in Ba-
c i d i m (26 if Bacidina neglecaa VEzda is considered as
Collections examined. CANADA. Alberta. Waterton distinct from Bn. chlomticulu), including the species
Lakes Area, 1964, Jonescu (WIS). Saskatchewaa. Cy- described in or combined into Bacidina in this work.
press Bills, 1964, Jonescu (WIS). - U. S. A. California. Although still a "young" genus, Bacidina has gained ac-
Tulare Co., 1985, Blakeman 360 (MW. Colorado. ceptance also in important reference works (e.g., Farkas
Boulder Co., 1973, Carmer 757 (CANL, COLO). Lar- & Sipmm 1993, Esslinger 1995, Wirth 1995). Woessiu
imer Co., 1962, Anderson 2825 (COLO). Idaho. Shos- and Lichingoldia, on the other hand, have not received
hone Co., 1987, McCune 16508 (OSC). Souah Dakota. any attention at all, with the recent unfortunate excep-
Lawrence Co., 1960, Anderson (COLO). Pennington tion of SCrusiaux (1995), who attempted to replace the
Co., 1960,. Anderson (COLO, MSC). name Baciddna with Woessia for exactly the same ge-
nus. The reason for doing so was purely nomenclatural,
Bacidina Vgzda, nom. cons. prop. i. e., there were no underlying taxonomical considera-
tions. The present Code strongly discourages this kind
Folia Geobot. Phytotax. Praha. 25: 43 1 (1991). - Type: of action: "Name changes made for purely nomenclatu-
Bn. phacodes (holotype). ral reasons ... ... are to be avoided" (Greuter et al. 1994:
Lichingoldia D. Hawksw. & Poelt, P1. Syst. Evol. xv). The tools for avoiding an undesirable name change
154: 203 (1986). - Type: L. gvalecfiformis D. Hawksw. from Bacidina to Woessia are provided by the Code
& Poelt (holotype). (Art. 14). I propose that Bacidina be conserved against
Woessiu D. Hawksw. & Poelt, P1. Syst. Evol. 154: both Woessia and Lichingoldiu (Ekman 1996). Pending
207 (1986). - Type: jbarioides D. Hawksw., Poelt the final settlement, 1 urge that Bacidina should be re-
& Tscherm.-Woess (holotype). tained.

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

Opera Bot. 127, 1996

thallus, usually f pale green. Upper cortex (not perispore or ornamentation, 16-60 pm long and 0.9-3.3
present in goniocysts) of one inner layer of anticlinal pm wide, 9-56 times as long as wide, easily separated
or irregular hyphae stainable in LCB and an outer from each other at maturity.
epinecral layer (one or the other may be missing), +
Pycnidia immersed in thallus, usually concolorous
without crystals. Photobiont layer continuous or dis- with or paler than apothecia, unilocular. Conidiogenous
continuous, photobiont ch1orococcoid. Medulla and cells lining the pycnidial cavity, cylindrical, 4-8 X 1.0-
lower cortex absent. 1.5 pm, producing conidia apically, lacking distinct CO-
Apothecia sessile, biatorine, rarely with an addi- nidiophores. Conidia of five different types (one to three
tional, thin thalline margin, when mature 0.1-2.0 mm types in one species; usually only one type, rarely two
wide, often weakly pigmented (pale pink or pale yel- types mixed in the same pycnidiurn): (I) filiform, f
low to pale orange), sometimes darker (yellow-brown curved, non-septate (rarely l-septate), 7-30 X 0.5-1.0
to orange to brown to red-brown to purple-brown to p,(2) bacillifom, 0(-2)-septate, 7-9 X 1.6 p,(3) fili-
black); pruina absent. Disk plane or slightly concave +
form, straight or curved to slightly sigmoid, distinctly
when young, often soon becoming f convex. Margin 3- l l -septate, 14-57 X 0.6-1.0 pm, (4) filifom, straight
usually slightly raised above disc in young apothecia, or slightly curved to sigmoid, strongly conglutinated, 0-
later level with the disc, often becoming excluded in l-septate, 9-14 X 0.6-0.8 p,and (5) acicular, non-sep-
old and convex apothecia. tate, 16-20 X l .O pm.
Proper exciple without thick gelatinous layer along Chemistry: No lichen substances. Negative to K, C,
rim, without oil droplets, rarely with crystals (c. l p) KC, PD, and UV.
that are evenly dispersed along the rim of the proper
exciple; composed of indistinctly or distinctly radiat- Discussion. As with Bacidia, the generic description of
ing, sparingly or abundantly furcate, usually moderate- Bacidina presented above is based on my observations
ly to abundantly anastomosed hyphae. Cell lumina on the North American species and must be regarded as
very variable in shape, from narrowly cylindrical to al- provisional.
most globose, 3-1 5 pm long and 1.5-8 pm wide (lumi-
na near rim of proper exciple excepted), usually 1-8
times as Iong as wide, but at least partly short and 28. Bacidina aenea S. Ekman, sp. nov.
wide (width exceeding 2.5 p,length less than 4 times
the width), widely cylindrical to almost globose or ir- Type: U. S. A., Florida, Sarasota Co., Myakka River
regular; often distinctly constricted near septa (not in State Park, c. 3 km S of the W end of Upper Myakka
very narrow lumina). Walls f thin, the distance be- lake, seasonally flooded hammock, on Fraxinus carolin-
tween two adjacent cell lumina usually less than the iana, 1993, Ekman L1207 (LD, holotype).
width of a lumen. Cell lumina along the rim of the
proper exciple rarely f enlarged in relation to the inner Thallus per maculas in goniocystas solutus. Apothecia
lumina, never forming a distinct zone. Hypothecium h l v a ad cervina ad fusca, pigmenturn hlvum (idem
usually colourless, sometimes yellowish to brown, quod in Baddia urceutina) continentes. Facile curn Ba-
without crystals or oil droplets, not chondroid. Hy- cidina clystullfera confusa, quae stratum crystallifer-
menium 40-80 pm high, amyloid, usually colourless in urn in epithecio et in margine excipuli proprii, item curn
lower part and k pigmented in a diffuse layer in the Bacidina variu quae hymenium tenuius, item cum Ba-
upper part, rarely with a crystal layer in the upper part, cidina sp. #l quae hymeniurn tenuius; apices paraphyso-
without oil droplets. Paraphyses unbranched or f rum latiores, et goniocystas tenuiores habent.
branched in upper part, sometimes very sparingly anas-
tomosed; apices not at all swollen to f clavate, some- Etymology. Aeneus, (here) brazen.
times surrounded by pigment, never containing pig-
ment in the wall, never with thick and gelatinized wall. IIlustrations. Figs 5E, 36 (map), 43A, 44A.
Asci clavate, surrounded by a thin, amyloid gelatinous
sheet; tholus amyloid, with a narrow or wide c-layer, Characterization. Thallus dissolved into goniocysts in
with a high or low, blunt axial body (sometimes pene- spots. Apothecia brown-yellow to yellow-brown to dark
trating through the entire d-layer), the width of which bey-brown. Proper exciple laterally brown-yellow. Hy-
is more than a third of the entire d-layer; ocular cham- pothecium and upper part of hymenium brown-yellow.
ber absent or present, low or high, blunt or pointed. Pigment K-. Spores acicular, clavate, or almost bacilli-
Spores eight per ascus, acicular to almost bacillifom, form, 30-43 X 1.9-2.5 pm, with 3-7 septa.
with 0-13 septa, straight, curved, or sigmoid, straight
or coiled in ascus (often variable even within the same Thdlus indeterminate, thin, discontinuous, of discrete
apothecium), colourless, rather thick-walled, without or contiguous, convex, sometimes almost granular, dark

Opera 001. 127. 1996

 8-1 yihmu ,~kmqs?ar%stgisxs twq I n m31 uan gn
hra a l l d . tnr-I 4- +.ud nA..n 4- 6-1 -I W-:*
Fig. 35. Known North American distribution of Bacidina arno!dmna (dots), Bn. assulata (star), and Bn. chloroticula (squares).
Known world distribution of Bacidina californica (triangles)

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

Opera Bot 127. 19%

(Arceutina Yellow). Furthermore, Bn. aenea is scparat-
ed froin Bn. crl;stullifera by the lack of crystals in the
proper exciple and hy~nenium,and by thc widcr spores,
from Bn. sp. # l by the higher hymcniuin, i~airowcrpara-
physis apices. widcr sporcs, and coarser gotliocysts, and
from Bn. var~.iaby a highcr hymcniun~,and possibly also
by slightly larger apothccia and finer go~~iocysts. The
apothecia of Bn. aenea arc superficially very similar to
the apothecia of Bncicliu arcezrti~~a due to the presence
of the same pigment.

29. Bacidina arnoldiana (Korb.) V. Wirth & Vi.zda

Stuttg. Reitr. Naturk. A. 5 17: h2 ( 1994) - Bacidiu arnol-

dianu Korb.. Parerga Lich., fasc. 2: 134 (1 860). - Type:
Germany, "ad saxa umbrosa calcarca in sylvaticis vallis
Rosenthal prope Eichstaedt Havariac", Amold, Ainold:
Lich. sel. Gcrm. 131 (L, lcctotype selected here).
Woc.~sicrfuL~arioides0 , Hawksw., PoeIt & Tsc11erm.-
Woess, PI. Syst. Evol. 1 54: 207 (1 986). - Type: Austria,
"Burgenland, Be7. Oberwart, Quercus-Sturnpf im Misch-
wald SW iiber Bad Tatzmannsdorf, nahe der Strasse
nach Oberwart, f 350 m", 1980, Poelt (herb. Tscher-
mak-Woess, isotype). Fig. 36. Known NOI-th Alncrican distribution of Bircidina
acnea (square), Bn. o~stullijkra (dots), Bn, egenrfla (tri-
angles), and Bn, egtltnrloiclru (star).

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.

Opera Bot. 127, 1996

116
Correctly determined and representative specimens Iinam et pigmenturn hydrato kalico intensius coloraturn
have been distributed by Vgzda in Lichenes selecli ex- habent.
siccati no. 566.
Bacidina assulatu is closely related and very similar Etymology. Calforuicus, Californian.
to the type species of Bacidinra, Bn. phacodes (Korb.)
VEzda, a species which is known with certainty only Illus~rations.Figs 2C, SH-I,35 (map), 43C,44B.
from Europe. Bn, phacodes differs from Bn. assu~atain
almost completely lacking pigments in the apothecia, Characterization. Thallus dissolved into goniocysts or
and (Swedish and French specimens) in possessing ba- not. Apothecia usually pale pink to orange-brown. Prop-
cilliforrn to filiform, straight or only slightly curved, 1- er exciple usually colourless to pale orange-brown. Hy-
5-septate conidia that measure 16-32 X 1 -0-1.2 p.Ac- pothecium colourless to pale brown-orange. Upper part
cording to Coppins (in Purvis et al. 1992), British of hymenium colourless to f brownish. Pigment K+
specimens of Bn. phacodes have conidia up to 60 p m green-brown. Spores acicular, 3 1-53 X 1.9-3.3 p,with
long, 1.5 p wide, and up to 7-septate. h Bn. assulata, 0-9 septa. Conidia filifom, curved, non-septate, or ba-
on the other hand, the conidia are strongly curved and cillifom, straight, 0-2-septate.
non-septate. In specimens from Poland (the type speci-
men) and the Czech Republic, they measure 10-43 X Thallus indeterminate, thin to thick, either continuous,
0.8-1.0 pm (i.e., generally slightly larger than in the wrinkled or warted, f rimose, or discontinuous, of dis-
American specimen). Furthermore, the thallus of Bn. crete to contiguous, convex, sometimes almost granular
phacodes is never dissolved into goniocysts, whereas in areoles, only seldom dissolved into loosely aggregated
most specimens of Bn. assulata small parts of the thal- granules (goniocysts) in small or large spots; grey,
lus are dissolved. green-grey, grey-green, or yellow-grey, except granular
Bacidina assulata is also very similar to the North spots, which are pale green. Goniocysts (21-)30(-52)
American species Bn. calfornica and Bn. varia. Forms pm diam. (S= 7 , N= l , n= 20).
of Bn. californica with pigmented apothecia can be sep- Apothecia (0.2-10.4-0.5-0.7(-0.9) mm diam. (S= 0.2,
arated by the brown, K+ green-brown pigment (Amol- N= 6, n= 10), f plane when young, later becoming con-
diana Brown). Furthermore, Bn, californica differs in vex, epruinose. Disk pale pink to pale yellow to pale or-
the presence of a distinct ocular chamber and a narrow, ange to orange-brown, rarely darker (red-brown to pur-
heavily amyloid zone along the outer edge of the axial ple-brown). Margin concolorous with disc or slightly
body. Bn, vuria can be distinguished from Bn,assulutu paler or darker, distinct, level with or slightly raised
by the lower hymenium (rarely exceeding 60 pm), the above disc in young apothecia, later level with the disc.
presence of a K+ purplish pigment in the apothecia finally excluded.
(Laurocerasi Brown), and the curved to sigmoid, septate Proper exciple Iaterally (29-)48(-69) wide (S= 14,
conidia. N= 5, n= l), without crystals. Hyphae with rather thin to
rather thick walk (1.5-2.5 pm between adjacent cell lu-
Collectiom examined. U . S . A. OkIuhoma. Cherokee mina), rather distinctly radiating towards edge; cell lu-
Co., 1988, Harris 21341 (W). mina f homogeneously shaped, in lateral interior 1.5-2
pm wide and up to 12 pm long, 2.5-6.5 times as long as
wide. Rim usually colourless to pale orange-brown,
31. Bacidina californica S. Ekman, sp. nov. sometimes orange-brown to red-brown, then darkest in
upper part and downwards gradually changing to col-
Type: U. S. A., "California. Monterey Co.: on Umbellu- ourless. Lateral interior colourless or pale orange-
lariu californica, floor of canyon at Big Sur", 1966, brown. Medullaly part colourless. Hypothecium colour-
Weber & Santesson, Weber: Lich. exs. 182 (COLO, less, very pale yellowish, or pale brown-orange.
holotype; LD, MIN, SFSU, UBC, US, WTU, isotypes). Hymenium (52-)57-64-73(-74) p m thick (S= 8, N= 6, n=
NB! In addition to Bn, culfornica, Bacidiu heterochroa 5), in lower part colourless except where vertical streaks
is present in most type specimens. of pigment reaches deep down in the hyrnenium; in up-
per part colourless, pale yellow-brown, pale orange-
Thallus aiiquando per maculas in goniocystas solutus, brown, rarely orange-brown to red-brown. Paraphyses
plenunque autem non solutus. Apothecia plerumque rather abundant in relation to the number of asci, 1.2-
pallide rosea ad aurantiaco-brunnea, pigrnentum brun- 1.6 p wide in mid-hymeniurn; apices f clavate or
neum hydrato kalico viridescens continentia. Pycnidia some not at all or only slightly thickened, (1.6-)2.2-2.5-
biformia, conidiis aut filifomibus curvatis non septatis, 2.8(-3.7) pm wide (S= 0.6, N= 6, n= 10). Tholus with a
aut bacillifomibus rectis 0-2-septatis. Facile cum Ba- narrow c-layer. Ocular chamber wide, low or high,
cidina ramea confusa, cuius apothecia rnarginem thal- blunt; axial body wide and high, conical, blunt, some-

Opera Bol. 127. 1996

117
times penetrating through the eiltirc d-lnycr. Spores ac- 32. Bacidina chloroticula (Nyl.) VGzda & Poclt in
icular, straight to curved to sig~noid,(3 1 -)35-4 1-46(-53) VGzda
p long (S= 5, N= h, n= 101, (1.9-)1.9-2.4-2.8(-3.3) Fun
wide (S= 0.4, N= 6, n= l@), (12.2-)16.7-17.5-19.7(- Folia Geobot. Phytotax. Praha 25: 432 (1991). - Leciclen
22.0) times as luilg as widc (S= 2, N= 6, n= 101, with chlor-oficula Nyl., Flora, Jcna 60: 564 (1877). - Bucidi(i
(0-)3 .h-5.7-7.5(-9) septa (S= 2. l , N= 6, n= 10). chlor.nliczlla (Nyl.) A. L. Sm., Munugr. Brit. Lich. 2:
Pycnidia f i~nrnerscdin thallus, coloul-less or pale or- 155 (1 9 1 1). - Type: Ireland, "in rivulo propc Kylein-
angc to pale brown, of two types: ( l ) 70-100 in diam; ore", 1877, Larbalestier (11-NYL 17648, lectotype se-
conidia iiliform, curved, non-septate, 7-12 X 0.6-0.8 lected here).
pm, (2) 75-225 pn in diam; conidia bacilliforn~: Leciilrn suhchioruticm Nyl.. Flora, Jena h 1: 45 1
straight, non-septate or s o ~ n cI -2-septatc, 7-9 X 1.6 pm. ( I 878). - Brrcidia sr~hchlorotica (Nyl.) Flagey, Catal.
Pigments: Pig~nentedpaits of apothecia K+ green- Lich. AlgCrie h4 (1896). - Type: France: "Corsica,
brown, N-. - Sometilnes Ar~loldia~~a Brown in proper Ajaccio, supcr corticcm Cupulariae viscosae", 1878,
exciplc, hypotheciuin, and hytnenium. Norrlii~(1.1-NYL 17954, lectolype selected here).

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.

Open B o t 127, 1996

Distribution. Known from one station in southern On- ous or darker than disc, in upper part pale orange to red-
tario and one in northernmost Louisiana. Probably much brown to purplish black, in lower part pale orange to
overlooked and possibly with a wide (eastern temper- brown-orange to dark purple-brown, distinct, slightly
ate?) distribution. - Europe. raised above disc in young apothecia, later level with
the disc.
Discussion. Bacidina egenula is very similar to Bn. Proper exciple laterally (46-)5 l(-56) p wide (S= 7,
egenuloidea. The latter species differs in lacking Bacid- N= 2, n= l ) , without crystals. Hyphae with rather thin
ia Green in the hymenium and in having very long and walls (1-3 p between adjacent cell lumina), rather dis-
almost straight conidia that are strongly conglutinated. tinctly radiating towards edge, with cell Iumina that are
Bn, egenula also resembles Bn. chloroticuku (see this narrower in upper part than in lower part, in lateral inte-
species). rior 1-4 pm wide and up to 14 p long, 4-8 times as
In a European collection of Bn. egenula (Oberfrank- long as wide in upper part, 1.5-4 times as long as wide
en, Engelhardsberger Brunnen, 1855 Arnold, L), anoth- in lower part. Rim orange-brown, grey-brown, or brown
er conidial type has been observed in addition to the one in upper part, downwards often gradually changing to
mentioned above and reported by Coppins (in Purvis et co1ourless:Lateral interior either colourless, or pale or-
al. 1992). It is found mixed in the same pycnidia as the ange to pale brown in upper part and downwards chang-
common type. These conidia are straight or curved, ing to colourless. Medullary part colourIess. Hy-
evenly thick or slightly thicker towards the proximal pothecium colourless or very pale orange. Hymenium
end, distinctly 3-7-septate, 32-45 X 1.O-1.6 pm. A very (43-)45-49-54(-56) p thick (S= 5, N= 2, n= 5), colour-
similar type of pycnidium with two conidial stages is less in lower part, diffusely pale orange or pale brown
found in B egenuloidea (see this species). in upper part. Parapkyses rather abundant in reIation to
the number of asci, 1.6 wide in mid-hymenium; api-
Collections examined. CANADA. Ontario. Ottawa, ces f clavate or some not at all or only slightly thick-
1976, Robitaille 149.4 (CANL). - U. S. A. Louisiana. ened, (1.6-)3.0-3.1-3.1(-4.9) p wide (S= 1.O, N= 2, n=
Union Par., 1970, Kay (herb. Tucker). 10). Tholus with a wide c-layer. Ocular chamber wide,
rather low, pointed; axial body wide and high, conical
or subcylindrical, blunt, never penetrating through the
35. Bacidina egenuloidea (Fink) S. Ekman, entire d-layer. Spores acicular, straight or slightly
comb. nov. curved to sigmoid, (20-125-29-34(-42) pm Iong (S= 6,
N= 2, n= 101, (0.9-)1.1-1.5-1.9(-2.1) pm wide (S= 0.5,
Bacidia egenuloidea Fink, Bull. Ohio S. Univ. 25: 346 N= 2, n= lO), (14.7-)l8.3-20.5-22.6(-29.0) times as
(1921). -Type: U. S. A., Ohio, Preble Co., "West Alex- long as wide ( S 5, N= 2, n= 1O), with (0-10.5-1.5-2.4(-
andria, damp field, granite", 1914, Fink 320 (MU-L608, 5) septa (S= 1.g, N= 2, n= 10).
lectotype selected here). Pycnidia immersed in thallus, colourless, 75-200 pm
diam. Conidia of three types: (1, the commonest type)
Il~ustrations.Figs 5N, 8D, G, 36 (map). filifom, straight or slightly curved to sigmoid, strongly
conglutinated, 0-I-septate, 35-49 X 0.8-1.2, pm, (2) fili-
Characterization. Thallus entirely dissolved into gonio- form, curved, not conglutinated, non-septate, 9-14 X
cysts. Apothecia with pale orange to dark purple-brown 0.6-0.8 pm, and (3) acicular (thicker at the proximal
disc; margin concolorous or darker. Proper exciple lat- end), straight or curved, not conglutinated, non-septate,
erally with various brown (K+ purplish) and orange 16-20 1.0 pm. Type 1 occur in separate pycnidia,
4

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?

Opera But. 127, 1996

 Characterization. Thallus not dissolved into goniocysts.
Apothecia with pale yellow to dark red-brown, often
mottled disc; proper margin usually paler; rnargo thalli-
nus present at least in young apothecia, thin, pale grey-
ish. Proper exciple (almost) colourless, except colour-
less to dark brown rim. Hypothecium (almost)
colourless. Upper part of hymenium colourless to red-
brown. Spores acicular, 25-57 X 1.2-2.5 pm, with 3-7
septa. Conidia filifonn, curved, non-septate.

Thallus indeterminate, thin, usually continuous, wrin-

kled or warted, f rimose to areolate, sometimes partly or
entirely discontinuous, of discrete or contiguous, con-
vex areoles, pale grey, pale yellow-grey, or pale green-
grey, never dissolving into granular goniocysts.
Apothecia (0.3-)0.4-0.5-Oh(-0.7) mm diam. (S= 0.1,
N= 9, n= IO), at first plane, later becoming convex,
epruinose. Disk pale yellow, pale pink, pale orange,
pale to medium brown-orange, pale to medium orange-
brown, or pale to dark red-brown, often mottled with
different hues. Proper margin usuaIly paler but sorne-
times concolorous with disc, milky white, pale grey,
pale pink, pale yellow, pale brown, seldom orange-
brown to dark red-brown, distinct, level with or raised
above disc in young apothecia, soon level with the disc,
finally excluded, sometimes appearing tomentose due to
protruding hyphae. Margo thallinus rather thin, con-
Fig. 37.Known world distribution of Bacidina ramea. colorous with the hallus, in at least some of the young
apothecia covering 50-100 % of the proper exciple,
soon excluded (but easily visible in section even in old
Discussion. Bacidina egenuloidea is similar to Bn. chlo- apothecia).
roticula and Bn. egenula (see these species). Proper exciple laterally (26-)45(-86) pm wide (S= l l ,
N= 9, n= l), without crystals. Hyphae with rather thin
Additional calkctions examined. U . S . A. Ohio. West walls (1.5-4 pm between adjacent cell lumina), not dis-
Alexandria, 1914, Fink 3 13 (MICH). tinctly radiating towards edge, with cell lumina f of the
same shape throughout the entire exciple, in lateral inte-
rior 2-3.5 pm wide and up to 9 pm long, 1-3.5 times as
36. ~acidinaramea S. Ekrnan, sp. nov. long as wide. Rim colourless, pale yellowish, or pale to
dark brown. Lateral interior colourless or very pale yel-
Type: Canada, Vancouver Island, NW of Victoria, lowish. Medullary part colourless or very pale yellow-
Thetis Lake Park, just S of the SE-most part of Thetis ish. Hypothecium colourless or very pale yellowish. Hy-
Lake, on Acer macrophylluna, alt. 80 m, 1989, Ekman menium (55-)57-62-65(-68) pm thick (S= 4, N= 9, n=
L767 (LD, holotype) . 5), colourless or diffusely pale yellow to pale brown to
red-brown, usually darker in upper part than in lower
Thallus nunquam in goniocystas solutus. Apothecia flava part. Paraphyses abundant in relation to the number of
vel pallide rosea ad fermginea, saepe variegata (pigmen- asci, 1.2-2.0 (occasionally up to 3.2) pm wide in mid-
turn hydrato kalico intensius coloratum), juniora margine hymenium; apices usually f clavate sometimes weakly
thallina praedita. Pycnidia conidiis filifomibus curvatis or not at all thickened, (1.6-)2.0-2.8-3.5(-5.3) pm wide
non septatis. Facile cum Bacidina califonim confusa, (S= 0.7, N= 9, n= 10).ThoIus with a narrow c-layer. Oc-
cuius apothecia margine thallina carent, pigrnentumque ular chamber lacking; axial body very wide, low, blunt,
brunneum hydrato kalico viridescens continent. cushion-like, never penetrating through the entire d-lay-
er. Spores acicular, straight or slightly curved/sigmoid,
Eay~no/ogy.Ra~neus,belonging to a branch. (25-)3 1-41-49(-57) pm long (S= 7, N= 9, n= 1 O), (1.2-)
1.6-1.9-2.2(-2.5) p wide (S= 0.3, N= 9, n= l 0), ( 1 2 5 )
Illustrations. Figs 5 0 , 37 (map),43E, 44D. 18.5-22.2-28.5(-37.0) times as long as wide (S= 5, N=

Opern Bot 127, 1996

 of B. varia. B. squameIlosu can then be separated fiom
B. varia by the wider paraphysis apices, the longer and
narrower spores, and the large, inflated cell lumina in
the proper exciple.

Additional collections marnined. U . S. A. Flo~.ida.Ala-

chua Co.,1993, Ekman L1149, E1150, L1151 (LD).
Sarasota Co., 1993, Ekman L1206 (LD).

38. Bacidina varia S. Ekman, sp. nov.

Type: U. S. A., Florida, Monroe Co., Lignumvitae Key

Fig. 39. Known North American distribution of Bacidina (N of Lower Matacumbe Key), tropical virgin forest, on
varia.
D~ypetesdiversfolia, 1993, Ekman L1 191 (LD, holo-
type).
1.6 pm wide in mid-hymenium; apices f clavate,
(2.4-)3.1-3.5-4.0(-5.7) p wide (S= 1.0, N= 3, n= Thallus saepe partim vel totaliter in goniocystas laxe
10). Tholus with an extremely wide c-layer. Ocular aggregatas solutus. Apothecia pallide rosea ad por-
chamber lacking; axial body wide and high, conicaI phyrea, pigmentum hydrato kalico immutatum vel in-
or semiglobose, blunt or pointed, filling up most of tensius coloraturn, aliquando etiam pigmentum hy-
the d-layer but never penetrating at its apex. Spores drato kalico purpurascens continentes. Pycnidia
acicular and straight, curved or sigmoid, (37-)45-46- conidiis filiformibus curvatis ad sigmoideis 0-1l -sep-
48(-60) 'pmlong (S= 5, N= 2, n= 10), (0.9-11.3-1.3- tatis. Facile cum Bacidina ueneu confusa quae hy-
1.4(-1.6) pm wide ( S = 0.2, N= 2, n= 10), (26.9-)33.9- menium altius et pigmenturn fulvum in apotheciis,
36.2-38.5(-55.7) times as long as wide (S= 8, N= 2, item cum Bacidina'sp. #l quae goniocystas tenuiores;
n= 10), with (3-)3.2-3.8-4.4(-7) septa (S= 1.4, N= 2, apices paraphysorum latiores, sporasque angustiores,
n= 10). item cum Bacidina crystallifem quae hymenium al-
Pycnidia immersed in thallus to sessile, colourless, tius, sporasque longiores et angustiores stratumque
75-100 pm diam. Conidia filifom, slightly curved, in- crystalliferum in epithecio et in margine excipuli pro-
distinctly 3-7-septate, 41-58 X 1.O-1.2 p. prii habent.
Pigments: All parts of apothecia K-, N-. - Pigments
absent. Etymology. Varius, variable.

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-

Opera Bot. 127. l996

124
ed in coralloid structures, (17-)25-28-29(-46) p diam. clude CephaIanthus occidentalis, Drypetes divers$olia,
(S= 7, N= 4, n= 10). F m i n u s caroliniuna, Ilex sp., Lngerstroemiu iadica,
Apothecia (0.2-)0.2-0.3-0.4(-0.6) mm diarn. (S= 0.09, Magnolia grandijlbm, Su bul palmerto, Sambakcus ca-
N= 8, n= 10), at first plane, later becoming convex, nudensis, and Yitis rrotunddfodia. One find was made on
epruinose. Disk pale pink, pale grey-pink, pale orange, a polypore on the trunk of a still living Quercus virgifl-
pale grey-brown, or f brown-orange to purple-brown, iaaa.
sometimes mottled with different hues. Margin con-
colorous with or slightly paler than disc, distinct, leveI Distribution. Known from southern Louisiana and most
with ox raised above disc in young apothecia, soon level parts of Florida. - Endemic?
with the'disc, finally excluded.
Proper exciple laterally (25-)40(-52) pm wide (S= l l , Discussion. Bacidina varia is remarkably variable in
N= 9, n= l), without crystals. Hyphae with rather thin to thallus structure and apothecium pigmentation. The
rather thick walls (1-3.5 pm between adjacent cell lumi- thaIlus varies from being not at all dissolved to entirely
na), not distinctly radiating towards edge, with cell lu- dissolved into goniocysts. The apothecia are usually
mina of widely differing shapes, often shorter and wider pale pink. but can sometimes become much darker.
in upper part than in lower part, gradually expanding to- Some morphs of Bn. varia resembles the European "Ba-
wards edge (in lateral interior 1-3.5 p wide and up to cidia "caligans (Nyl.) A. L. Sm. (a species of Bacidina,
15 pm long, 1-1 1 times as long as wide). Rim colour- but the combination has not yet been made). In the latter
less, pale yellowish, or pale orange-brown. LateraI inte- species, however, the apothecial margin is generally
rior colourless, pale yellowish, pale brown-orange, or darker than the disk due to a concentration of Laurocer-
pale red-brown. Medullary part colourless. Hy- asi Brown to the uppermost part of the proper exciple.
pothecium colourless, pale yellowish, pale brown-or- Bn. varia is also similar to B. aenea, Bn. cyslullifera,
ange, or pale red-brown. Hymenium (40-142-52-61(-62) and Bn. sp. # I (see these species).
pm thick (S= 7, N= 9, n= S ) , colourless, pale yelIowish,
or pale brown-orange. Pmphyses usually rather few in Additional codlections examined. U . S . A. FIorida. Ala-
relation to the number of a c i , 0.8-1.6 p wide in mid- chua Co., 1993, Ekman 1152 (LD). Polk Co., 1993, Ek-
hymenium; apices f clavate or not at all thickened, man L1 169, L1 170, L1 171 (ED). Sarasota Co., 1993,
(0.8 -)I -6-2.0-2.7(-4.4) pm wide (S= 0.6, N= 9, n= 10). Ekman L1208 (LD). Sanford, 1914, Rapp (FH, US).
Tholus with a narrow c-layer. Ocular chamber lacking Louisiana. East Baton Rouge Par., 1993, Ekman L1 116,
or wide and blunt, sometimes surrounded by a narrow L1 122, L1 123, L1 124 (LD), 1983, Tucker 25618 (herb.
zone that is more heavily amyloid than the remainder of Tucker), 1992, Tucker 32 181 (herb. Tucker). St. Helena
the d-layer; axial body wide and high, conical, blunt, Par., 1986, Tucker 27953 (herb. Tucker). St. Tarnmany
never penetrating h u g h the emire d-layer. Spores ac- Par., 1993, Ekrnan L1 135 (LD); 1993, Tucker 33490
icular to clavate or sometimes almost bacillifom, (herb. Tucker).
straight to curved to sigrnoid, (19-)23-29-41(-50) pm
long (S= 7, N= 8, n= 101, (1.2-)1.7-2.0-2.4(-2.5) pm
wide (S= 0.3, N= 8, n= 10), (8.5-)10.0-14.8-23.5(-33.0) 39. Bacfdina sp. # I
times as long as wide (S= 6, N= 8, n= 10), with (3-13.0-
4.4-6.6(-7) septa (S= 1.7, N= 8, n= 10). Illustrations. Figs 5R, 38 (map).
Pycnidia immersed in thalIus that is not dissolved in-
to goniocysts, uppermost part of wall concoloraus with Characterization. Thdlus partly dissolving into gonio-
apothecia, 75-175 ym diarn. Conidia filiform, f curved cysts; goniocysts h e , 9-2 1 ym diarn. Apothecia f pink.
or sigmoid, indistinctly 3-1 l-septate or sometimes non- Proper exciple, hypothecium, and hyrnenium (almost)
septate, 14-55 X c. 0.6- 1.0 p. colourless. Paxaphysis apices wide, 3.2-3.3 pm (average
Pigments: All parts of apothecia R- to + intensifying, of 10). Conidia filifom, straight or f curved, indistinct-
N-, except pale red-brown parts of proper exciple, hy- ly 5-1 l-septate
pothecium, and hymenium, which react K+ purplish,
N+ orange. - Sometimes small amounts of Rubella Or- Thallus indeterminate, thin, either discontinuous, of dis-
ange, sometimes also Laurocerasi Brown, in proper ex- crete or contiguous, convex areoles, or continuous,
ciple, hypothecium and hyrneniurn. wrinkled or warted, without cracks, green to grey-green,
partly dissolving into granular goniocysts. Goniocysts
Ecology. Usually on trunks or branches of trees and loosely aggregated, paIe green, (9-112-14-16(-21) pm
shrubs in shady and humid as well as open and rather d i m . {S= 3, N= 2, n= 20).
dry forests. Sometimes also in more or less open stands Apothecia (0.3-10.3-0.3-0.3(-0.4) mm diam. (S= 0.03,
of shrubs in man-made habitats. Known phorophytes in- N= 2, n= 10), plane, later becoming convex, epruinose.

Opera Bm 127, 19%

Disk pale pink to pink. Margin concolorous disc, dis- physis apices, the narrower spores, and the wider cell
tinct, raised above disc in young apothecia, soon level lumina in the proper exciple, from B. aenea by the low-
with the disc, finally excluded. er hymenium and the lack of a brown-yellow pigment
Proper exciple laterally 43 pm wide (N= 1, n= l), (Arceutina Yellow) in the apothecia, and from Bn. ciys-
without crystals and without internal pigment. Hyphae talliyera by the lower hymenium, wider paraphysis api-
with rather thin walls (2.5-4 pm between adjacent cell ces, and the lack of crystaIs in the hymenium and proper
lumina), f distinctly radiating towards edge, cell lumina exciple.
gradually expanding towards edge (in lateral interior 2-5
pm wide and up to l l pn long, 1.5-3.5 times as Iong as Collections examined. U . S . A. Florida. Sanford, 1914,
wide). Hypothecium colourless or very pale yellowish. Rapp (FH). Louisiana. East Baton Rouge Par., 1993,
Hymenium (51-)53(-57) pm thick (S= 2, N= 1, n= 5), Ekrnan L1 121 (LD).
colourless. Paraphyses abundant in relation to the
number of asci, 1.2-2.0p n wide in mid-hymenium; api- Exsiccates
ces f clavate, (1.6-)3.2-3 .2-3.3(-5.7) pm wide (S= 1.0,
N= 2, n= 10), without pigment. Tholus with a rather This i s an account of all the exsiccate specimens I have
wide c-layer. Ocular chamber narrow or wide, f blunt; seen containing North American Bucidia and Bacidina.
axial body very wide and high, conical, blunt, filling up The exsiccates listed are those that were accepted by
most of the d-layer but never penetrating at its apex. Lynge (1915) or Sayre (1969), with the exception of J.
Spores acicuIar, curved or sigmoid, (30-)37(-43) pm Macoun's "Canadian Lichens", which I do not consider
long (S= 4, N= 1, n= 10), (1.2-)1.4(-1.9)ym wide (S= to be an exsiccate. Also included here, but not listed by
0.2, N= 1, s 10), (22.7-)27.9(-33.0) times as long as Lynge or Sayre, is Plitt's "Lichenes Exsiccati ex Her-
wide (S= 4, N= 1, n= 10), with (3-)3.4(-S) septa ( S = 0.7, bario Dr. H. E. Hasse, relicti". The exsiccate numbers
N= l , n= 10). below are followed by the name of the taxon under
Pycnidia immersed in thallus that is not dissolved into which it was issued, the herbaria in which I have seen
goniocysts, uppermost part of wall concolorous with this collection, and finally my determination.
apothecia, 50-125 pm diam. Conidia filifom, straight or
f curved, indistinctly 5-1 l -septate, 41 -57 X c. I .O p. Cummings: Decades of N. Am. Lich.
Pigments: All parts of apothecia K-, N-. - Sometimes 17 1. Biatorasums (FH, N Y , U S ) ,Bacidia suflusa
small amounts of an unidentified yellowish pigment 208. Biatora rubella (NY,U S ) , Bacidia polychroa
(Rubella Orange?) in the hypothecium. 228. Biatora rubella (NY,US),Bacidia polychroa
247. Biatom schweiaitzii (NY),Bacidia schweinitzii
Ecology. The specimen from Sanford was collected on 258. Biatom rubella (FH, NY, U S ) ,Bacidia polychruu
Sarnbucus canadensis, where it grew together with Ba- 267. Biatora schweinitzii (COLO, NY, U S ) , Bacidia
cidia heterochrora and Bacidina varia. At the locality in schweinitzii
Baton Rouge, I observed Bn. sp. # 1 growing on old
stems of Vitis rotundifoiiu in a dense, shady and rather Cummings et al.: Lich. Boreali-Am.
humid forest, forming rather large sterile patches with 102. Biaaora sufusa (COLO), Bacidia suffusa
few apothecia. Few other lichens were present near Bn. 169. Biatora sckweinihii (COLO),Bacidia schweinitzii
s p . # 1. 195. Biatora rubella (COLO), Bacidia polychroa
200. Biatom schweinikii (COLO, UBC),Bacidiu schwein-
Distribution. 1 know of only two sparsely fertiIe collec- itzii
tions, one from Sanford, Florida, and one from Baton
Rouge, Louisiana. - Endemic?. Farlow Herb.: Reliquiae Tuckermanianae
16. Biutoru sclaweinitzii (CANL, COLO), Bacia'ia schweirz-
Discussion. I am fairly certain that the taxon I refer to itzii
here as Bn. sp # 1 is a distinct species, although obvi-
ously closely related to Bn. varia. The reason for not de- Merrill: Lich. Exs.
scribing it is that the material presently availabIe is very 30. Bacidia rosella (COLO, M I N , NY, U S , WIS),Ba-
sparse and includes only a few apothecia. None of these cidia rubella
colIections is suitable as a holotype. Additional speci- 119. Bacidia atrosanguinea (COLO, m),Bacidia
mens with abundant apothecia are needed before this li- schweinitzii
chen can be formally described. 171. Biatora rubelda (COLO, US ), Bacidia polychma,
Bacidina sp. # l is similar to Bn. aenea, Bn. crystal- in COLO also with an admixture of B, sufisa
Igera, and, above all, Bn. vap.ia. Bn. sp # l is separated 214. Biatura schweinitzii (COLO, F H , U S ) , Baciddu
from Bn. varia b y the finer goniocysts, the wider para- schweinitzii

Opera Bot. 127, 1996

prothaIlus consisting of a web of hyphae that are pur- Bacidia carneoalbida (Miill. Arg) Coppins
ple-brown in the microscope. Apothecia slightly con-
cave to flat, with orange-brown disc and a thick, red- Lichenologist 2 1: 99 ( 1 989). - Pateliaaria carneoalbida
brown to black margin. Proper exciple orange-brown Miill. Arg., Flora, jena 51: 50 (1868). - Biatora car-
to red-brown along rim, internally colourless to pale neoalhida (Miill. Arg.) Coppins in Coppins, P. James &
yellow, without crystals or oil droplets, consisting of D. Hawksw., Lichenologist 24: 367 (1992). - Type:
hyphae with lumina that are -1- ellipsoid (3.0-3.5 pm Switzerland, Bern, Thun, "Stockhorn", 1867, Fischer
wide), often protruding from the lower part (anchorage (G, lectotype selected by Gilbert et al. 1988 as "holo-
hyphae?). Hypotheciurn pale yellow. Hymenium col- type"; not seen by me).
ourless to pale yellow, c. 50 pm high. Paraphyses Bacidia sphaeroides auct., non (Dicks.) Zahlbr.
sparsely branched in upper part, apical cell sometimes
swollen to 3 pm, the wall of which is not thick and ge- This muscicolous and corticolous lichen, often referred
latinized. Spores filiform, distinctly l-3-septate, 25-30 to as Bacidia sphaeroides, belongs in Mycobilipnbia
x c . 1.2 p. (Printzen 1995); see the section Related genera and
The affinities of this corticolous taxon are unclear, species groups in the chapter Tmonomic delimitations
but it does not belong in Bacidia or Bacidina. It should a hove species level.
possibly be referred to Sporacestra.

Ropalospora chlorantha (Tuck.) S. Ekman

Bacidia bagliettoana (A. Massal. & De Not. in A.
Massal.) Jatta Bryologist 96: 586 (1993) - Bicatora chloraatha Tuck.,
Proc. Am. Acad. Arts Sci. 1: 252 (1848). - B a c j i a
Sylloge Eich. Ital. 42 1 (1900). - Scoliciospomm bagliet- chlomntha (Tuck.) Fink, Contr. U.S. natn. Herb. 14 (1):
toanurn A. Massal. & de Not. in A. Massal., Memor. Li- 91 (1910). - Type: U. S. A., New Hampshire, "White
chenogr. 126 (l 853). - Type: Not seen. Mts", 1847, Tuckerman (FH-TUCK, lectotype selected
by Ekman 1993)
This species is a member of Bacidia S. str. It grows di-
rectly on soil or, more often, on bryophytes overgrow- This corticolous species, which has traditionally been
ing soil. North American deterrninations are generally treated as Bacidia chlomnthra, was transferred to Ropal-
correct. ospom by Ekman (1993). Since the publication of that
paper, one additional species traditionally treated in Bu-
cidia has turned out to belong in Ropalosporu, namely
Bacidia beckhausii Kiirb. the Japanese corticolous species Ropa1ospor.a phaeo-
placa (Zahlbr.) S. Ekman comb. nova (Bacidia phaeo-
Parerga Lich., fasc. 2: 134 (1860). - Type: Gennany, placa Zahlbr., Botan. Magaz. Tokyo 41: 333, 1927;
"Westphalen", Beckhaus (L-9 10.137 1363, lectotype se- type: Japan, "Insula Nippon, Takayu", 1904, Faurie
lected here.). 5882, W, lectotype selected here). It is obviously closely
related to R. chlorantha, but differs in having the proper
This corticolous species belongs in the Bacidia beck- exciple inspersed with minute K-soluble crystals and in
hausii group; see the section Related genera and spe- usually having eight (in some asci up to 12 or 16?)
cies groups in the chapter Taxonomic delimitations spores per ascus.
above species level.

Bacidia clementis Hasse

Bacidia brouardii (de Lesd.) ZahIbr.
Bryologist 13: 61 (1910). -Type: U. S. A., California, "on
Cat. Lich. Univ. 10: 359 (1939). - Bilirnbia brouardii de Heteromeles, San Clemente Island", 1903, Trask (FH,
Lesd., Annls Cryptog. exot. 5: 105 (1932). - Type: Not syntype).
seen.
This is a younger synonym of Buctrospom patellario-
I have not been able to trace any original material of this ides var. patellarioides.
species. It was described from New Mexico, where it
was found on siliceous rocks (Bouly de Lesdain 1932).
Unfortunately, the diagnosis gives no clue as to the ge-
neric affinities.

Opera Bot. 127, 1996

Lecania cuprea (A. Massal.) P. Boom & Coppins in This saxicolous taxon belongs in Lecania. I am not sure
P. Boom whether it is just a depauperate form of Lecania e~ysibe
(Ach.) Mudd or a separate species.
Nova Hedwigia 54: 234 (1992). - Bilimbia cuprea A.
Massal., Lotos B: 77 (1 856). - Bacidia cuprea (A. Mas-
sal.) Lettau, Hedwigia 52: 132 (1912). - Lecidea cupre- Fellhanera floridana (Tuck.) S. Ekman, comb. nova
orosema Nyl ex Stizenb., Nova Acta Acad. Caesar.
Leop. Carol. 34 (2): 9 (18671, made as a nomen novum Bia~omJoridana Tuck., Synops. N . Am. Eich. 2: 39
due to the existence of Lecidea cuprea Sommerf. - Bia- (1888) - Bacidia floridana (Tuck.) Zahlbr. in Engler &
tora cupreorosella (Nyl. ex Stizenb.) Tuck., Genera Prantl, Natiirl. Pflanzenfam. l (1): 135 (1905). - Type:
Lich. 164 (1872). - Bacidia cupreorosella ( N y l . ex Sti- U. S. A., Florida, "Caloosa River", 1878, Austin (FH-
zenb.) A. Schneid., Guide Study Lich. 109 (1898) - TUCK, lectotype selected here).
Type: Not seen. Bilimbiajhkii Vain., Mycologia 2 1: 35 (1 929). - Ba-
cidia Pnkii (Vain.) Zahlbr., Cat. Lich. Univ. 8: 401
This saxicolous species has traditionaIly been treated as (1932). -.Type: Puerto Rico, "Rio Piedras", 1915, Fink
a species of Bacidia, but it belongs in Lecania; see the 534 (TUR-V, lectotype selected here).
section Related genera and species groups in the chap- Bacidiu kaeticolor Malme, Ark. Bot. 27A (5): 8
ter Tmonomic delimitarions above species level. I have (1 935). - Type: Paraguay, "inter Lambart et Asuncion",
not checked the determinations of the North American 1893, Malme 1647 (LD, paratype).
specimens.
Fellhanem jloridana is transferred here to the genus
Fellhanera on account of the distinctly paraplectenchy-
Bacidia declinis (Tuck.) Zahlbr. matic proper exciple, tholus structure (dark tubular
structure that fades towards the tip of the ascus), abun-
Cat. Lich. Univ. 4: 109 (1926). - Lecidea declinis Tuck., dantly branched paraphyses, and short, 3-septate spores.
Genera Lich. 182 (1 872). - Type: U. S. A., Massachu- It is exclusively corticolous and IignicoIous, but appears
setts, "NB" (= New Bedford), 1867, Willey (FA-TUCK, to be closely related to the foliicolous FeUhaneru
lectotype selected here). lisowskii (Vzzda) VEzda (described as Bncidia lisowskii
by VGzda 1980). F. Joridana differs sIightIy from the
This name is a younger synonym of Cutillaria nigrocla- latter in the pigmentation of the thallus and the apothe-
m t a (Nyl.) Schuler. cia, and in the non-halonate spores.
Short description of the type specimens studied:
Thallus thin, f granular, yellowish. Apothecia up to 0.6
Bacidia epixanthoides (Nyl.) Lettau mm wide, orange-brown to red-brown, margin con-
colorous with disc or slightly paler. Proper exciple of
Aedwigia 52: 133 (1912). - Lecidea epixanthoides Nyl., hyphae with almost sphaerical lumina ("paraplectenchy-
Flora, Jena 48: 5 (1865). - Biatora epixanahoides (Nyl.) matic"), yellow to orange-brown along rim, pale yellow
Diederich, Trav scient. Mus, nat. Hist. nat. Euxemb. 14: inside. Hypothecium pale yellow. Hyrnenium 40-50 pm
57 (1989). - Type: Not seen. high, with yellow to orange-brown epithecium (this and
rim of proper exc. K+ citrine yellow), pigment when in
This mainly corticolous species has for a long time high concentration appearing crystalline. Paraphyses
been treated as a Bacidia, but belongs in Myco- abundantly branched. Spores 3-septate, slightIy con-
bilirnbia (Printzen 1 9 9 5 ) ; see the section Related stricted at septa, 12-14 . 3-5 pm.
genera and species groups in the chapter Taxo-
nomic delimitations above species level. I have not
checked the determinations of the North American Bacidia globulosa (Florke) Hafellner & V. Wirth in
specimens. Wirth

Flecht. Baden-Wiirttemb. 5 11 (1987). - Lecidea globu-

Bacidia flavens (WiIley in Tuck.) Zahlbr. loss Florke, Deutsche Lich. 10: 1 (1821). - Catillaria
globulosa (Florke) Th. Fr., Lichenogr. scand. 2: 575
Cat. Lich. Univ. 4: 1 12 (1926). - Biatoraflavens Willey (1 874). - Type: Not seen.
in Tuck., Synops. N. Am. Lich. 2: 34 (1888). - Type: U.
S. A., Massachusetts, "New Bedford", Willey (US, lec- This lichen has traditionally been called Catillaria
totype selected here). globulosa, but the name Bacidia globtilosu has been used

Opera Bot. 127, 1996

however, it is occasionally found on the lower parts of Byssoloma meadii (Tuck.) S. Ekrnan, comb. nova
tree-trunks or on cxposed tree-roots etc. m similar habitats.
Biatova meadii Tuck., Synops. N . Anl. Lich. 2: 129
(1888) - Bacidia nzeaclii (Tuck.) Zahlbr.. Cat. Licl~.
Bacidia ioessa Herre Univ. 4: 122 (1926). - Type: U. S. A., Florida, Mead
(FA-TUCK, lectotype seIected here).
Proc. Wash. Acad. Sci. 12: 98 (1910). -Type: Not seen.
This poorly known species is transferred here to
I haw bccn unable to trace any original material of this B~~ssulotna on account of the tholus (a dark tubular
saxicolous species. Judging from the diagnosis, it prob- structure that fades towards the tip of the ascus),
ably does not belong neither in Bacidia nor Bacidina. byssoid proper exciplc, short 3-scptatc spores, and
drop-shaped to short-bacillifurm conidia. It is very
closely related to the mucl1 inore common and wide-
Bacidia jacobi (Tuck.) Hasse spread B y s s u l o ~ ~leucuphlt.bur-urn
~u (Nyl.) Vainio. I
have been in doubt whether B. meadii should be rec-
Contr. U.S. natn. Herb. 17: S 1 (1913). - Biatora jucobi ognized as a distinct species. I have, however, see11
Tuck., Synops. N. Am. Lich. 2: 48 (1888). - Typc: U. S. both species several times in Florida and Louisiana,
A., California, "San Dicgo", 1875, Palmer (FI-I-TUCK, where they are often found in the same localities, and
lectotype selected here). it has not been a problem to keep them apart. This
has convinced me that they are in fact different spe-
Short description of the Icctotypc: Tliallus rather thick, cies. Byscolomra meadii is found in southeastern Unit-
pale, wrinkled to warted. Apothecia black, c. 0.5 mm in ed States, and appears to be exclusively corticolous,
diain., flat, with margin level with the disc. Proper exci- contrary to B. leucophlebarupn. I11 localitics whcrc B.
ple mostly colourless, but the interior gradually merging mendii is found on bark, I have searched for it in vain
into the pigmentation of the hypothccium, with crystals on leaves, even if the foliicalous lichen flora as a
in the rim and as radiating clustcrs in thc interior. Hy- whole has been rich. B. inuudii does not seem to
pothccium brown, K-. Hyrneilium c. 75 pn high, col- agree with any of the species treated by Kalb & V&-
ourless exccpt for brown spots in the epithecium, which da (1990) and Liicking ( 1 992). The differences be-
is also inspersed with cryslals. Paraphyses unbranched tween B. nreadii and B. leucophlehrarum are summa-
or sparsely branched in upper part, with apical swelling rized in Tab. 18.
to c. 4 pm, some apical cells with a distinctly delimited,
brown (K-) hood. Ascus with a pointed, conical ocular
chamber and a rather narrow axial body that reaches all Bacidia microcarpa (Th. Fr.) Lettau
through the d-layer. Spores acicular with both ends
pointed, 5-septate, c. 30 X 2.5 pn. Hedwigia 52: 132 (19 12). - Bilimhia ~?hscut-a!uvar.
This is a corticolous spccics, the affinities of which I do naicrocarpa Th. Fr., Nova Acta R. Socf Ycient. upsal.,
not understand. I excludc it from Bucidia and Bucidznu on scr. 3, 3: 283 (1861). -- Typc: Norway, "M. Finmark,
account of thc tholus: the pure brown and K- pigment in Varanger, Mortensnaes", 1857, Th. M. Frics (UPS, svn-
the hypothcciuin, and the spores having both ends pointed. pe).
The isolectotype in US is a different taxon, which be-
longs in the Arthoniales. This specimen was probably This is an arctic species overgrowing terricolous bryo-
never seen by Tuckerman. phytes, and belongs in the Bucidia snb~tletorwm
group; see the section Aelnted genera und .species
groups in the chapter Tcrxonornic drlimltatio~uabove
Bacidia kingmanii Hasse level. I have not examined the North Ameri-
.~pecIe.~.
can specimens. The name Bacidia microcarpa has
Bryologist 14: 101 ( 1 91 l), as "kingman?'.- Type: U. S. A., been widely misused for forms of B. sahuleloru~~z
California, "Mt. WiIson Trail Road, San Gabriel Mts", with predominantly 3-septate spores. The specimen
19 10, Kingman (FH, 3 syntypes). depicted as "Mycobilimbia obscurafa" by Hafcllncr
(1984: 309, note the warted perispore) is probably
This is a saxicolous species with unclear affinities. It is such a form. B. nzicrocar-pa can be distinguished
excluded from Bacidia and Bacidina on account of the horn B. subwleturu~non accounl of thc rcgularly 3-
densely packcd and thin-walled excipular hyphae, the septate spores lacking a perispore, slightly thinner
richly branched paraphyses, and the tholus, which, al- paraphyses (only a few exceeding 2 ,m1 in width),
though similar to Bucidia, lacks a discernabIe c-layer. and the dominance of a brown pigment over a green
Tab. 18. A co~nparisonhettvcen Bl,ssolotnu /eirr.ophlehclrrtrn (Nyl.) Vainio and B, n~eridii(Tuck.) S. Eknlan.

Apothccia 0.3-U.5(-0.6) iilln wide. grcyish n r 0.2-0.3-0.5)mm, palc dirty

brownish, rarely yellowish (pigment ycllow
dcficicnt ~norphs)

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

Pale 10 dark brown, rarely cnlourlcss Colourless or palc yellowish

(pigment deficient specimens)
Pycnidia Cuininon, 100- 150 pm wide Rare, 50-75 pm wide

Ecology Mostly roliicolous or corticolous, ' Cofliculous. preferring twigs and

rarely saxicolous bra~lches

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.

Bacidia pallens (Kullh.) ZahIbr

Bacidia nivalis Follmann
Cat. Lich. Univ. 4: 134 (1926). - Bilinahicd pallens
Philippia 4: 30 (1979). - Type: U. S. A:. Wasl~ingtoi~, Kullh., Not. Sallsk. Fauna FI. Fcnn. Forh. 1 1 : 274
"....iin Bercich dcr Sominerschneegrenze am Nor- ( l 8 7 1). - Type: Finland, Tavastia australis, "Mustiala,
dosthang des Mount Baker iibcr dcm Austinpass, What- ad corticein tiliae", 1869, Ktllll~em(H: lcctotype se-
corn County", 1969, Follmann, FolImailfi: Lichenes exs- lected by Printzen 1995: 203 as "holotype"; UPS. iso-
iccati selecti a museo historiae naturalis Casseleilsi cditi type).
262 (LD, isotype).
The lichen traditionally callcd Bacidia prrllens, a corti-
The pigmentation, thc structure of the proper exciple, culous species, i s excluded here from Bacidia and Ba-
and Ihe lholus appearailce shows that this saxicolous li- cidina on account of the "paraplectenchymatic" proper
chen is related to Porpidia. Contrary to the claiins by its exciple composed of hyphae with ellipsoid, globose, or
irregular cell lumina, the usnic acid containing apothe- delimi~utions
species gr-ozlps in the chapter Tc~~tonontic
cia, the crystal-inspersed proper exciple and epithecium, above species level.
and the bacillifos~n to narrowly elliptical to narrowly
drop-shaped conidia. The affinities of Bacidicr pallens
are almost surely with Cliuston~~tm. Bacidia ravenelii (Tuck.) Zahlbr.

Cat. Lich. Univ. 4: 138 (1926). - Riaioru ruvenelii

Bacidia pallidula Zahlbr. Tuck., Synops. N . Am. Lich. 2: 34 (1888). - Type: U. S.
A., Florida, "Caincsvillc", 1877, Ravenel 65 (FH-
Cal. Lich. Univ. 4: 231 (June 1926), nom. il1eg.. non TUCK, two syntypes).
(Kremp.) Zahlbr. (Feb. 1926). - Biaturu yallidu Arnold
ex Eckfeldt. Bull. Torrey bot. Club 22: 254 (1895). - This is a saxicolous species with unclear arfinilies. It is
Type: Canada, Newfoundland, 1894, Waghorn 13 (PH, distantly related to Bcrcicl'ia and Bncidinu.
lectotype selected here).

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).

Bacidina phacodes (Korb.) VGzda

Ophioparma rubricosa (Miill. Arg.) S. Ekman,
Folia Geobot. Phytotax. Praha 25: 432 (1991). - Bacidia comb. nova
yhucades Korb., Parerga Lich., fasc. 2: 130 (1860). -
Type: Gem~any, (1) "Miinchen", 1853, Amold, (2) Pafellaria ruhricnsa Miill. Arg., Hedwigia 34: 142
Eichsddt, 1858, Anlold (L, syntypes). (1895). - Bacidia ruhrico.w (Mull. Arg.) Zahlbr., Cat.
?Seoulig(~at.cezrtinu var rrlhesccns Stizenb., Nova Ac- Lich. Univ. 4: 139 (1926). - Type: U. S. A., Idaho, "Lake
ta Acad. Caesar. Leop. Carol. 30 (3): 43 (1x63). - Bm- Fend d'OreilIen, 1880, Eckfcldt 28 (G, holotype)
cidia alhe.~cens (Stizenb.) Uausch, Verhandl. natulw. Bucidia herrei Zahlbr., Aiu~lsinycol. 6: 130 (1908). -
Ver. Karlsruhc 4: 106 (1 869). - Type: Not seen. Ophiopurnru hewei (Zahlbr.) Kalb 8c Staiger, Biblthca
lichenol. 58: 193 (1995). - Typc: U. S. h., California,
The few North American collections labelled Bacidia Santa C n ~ zMountains, Devil's Calion, on dead Acieno-
phucodes or its probable synonym B. r~lbescensgeneral- storna fascictllcdhlnr, 2300 ft., 1906, Herre 943 (W, holo-
ly contain other species of Baciiiinu, such as Bn, cali- type, not seen by me; FH, isotype, seen).
jowtica, Bn. vuriu, or Bn. sp. # 1 . B ~ ~ c i dyhucvdtls
i~u is a Haemaiomrna paciflcum Hasse, Bryologist 13: 62
corticolous species that does not occur in North Ameri- (1910): as '>acijica". - Type: U. S. A., California, Santa
ca. C ~ u zMountains, ncar Grizzly Peak, on bark of Psetldo-
tsugu tuxifvliu, 2700 ft., 1906, Herre 1033 (F, lectotype
selected by Kalb 8c Staiger 1995: 194, not seen by me).
Arthrosporum populorum A. Massal. Hrzematum~na calfornic~rm Sigal & D. R. Toren, '

Bryologist 78: 71 (1975). - Type: U . S. A . , California,

Memor. Lichcnogr. 128 (1 853). - Bacidia populor.zlm Lake County, north shore of Lake Pillsbury, at Sunsel
(A.Missal.) Trevis., Trevisan: Lichenotheca Veneta no. Campground, 55 1 m, 1973, Thiers 30899 (SFSU, holo-
163 (1869). - Typc: Not sccn. typc, not seen by me).

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 species is a member of Bacidia s. str. It is charac-

Bacidia sabdetorum (Schreb.) Lettau teristic of marine rocks, where it occurs in the Vermcar-
ica zone, prefmably on shady surfaces. In North Ameri-
Hedwigia 52: 132 (1912). - Lichen sabuletomm ca, it is found along both the west and the east coast.
Schreb., Spicil. Flor. Lipsiensis 134 (1771). - Myco-
bilimbia sabuletomm (Schreb.) Hafellner, Beih. Nova
Hedwigia 79: 3 10 (1984). - Type: Not seen. Lecania stigmatella (Tuck.) S. Ekman, comb. nova
?Bilimbia accedens Arnold, Flora, Jena 45: 391
(1862). - Bacidia accedenr; (Arnold) Lettau, Hedwigia Biatora stigmatella Tuck., Genera Lich. 167 (1872). -
52: 131 (1912). - Type: Not seen. Bacidia stigmatella (Tuck.)Zahlbr., Cat. Lich. Univ. 4:
Secoliga fuscorubella var. propinqua Stizenb., Nova 241 (1926). - Type: U. S. A., Louisiana, 1853, Hale
Acta Acad. Caesar. Leop. Carol. 30 (3): 58 (1863). - Ba- (FH-TUCK, lectotype seected here).
cidia propinquu (Stizenb.) Amold, Flora 49: 531
(1866). - Type: Switzerland, "Ziirich, an der Rinde von This is a corticolous species found in the southeastern
Pop. tremulu", Hegetschweiler, Hepp: Flecht. Eur. 519 United States. It is closely related to Lecania naege/ii,
(LD, syntype). and their apothecia are almost identically pigmented. L.
stigmatella, traditionally treated as a Bacidia, differs
This muscicoIous and corticolous species belongs in the from L. naegelii in having narrower spores, filiform,
Bacidia sabuletontm group; see the section Related curved, O- l -septate conidia, and a thallus that often dis-
genera and species groups in the chapter T m n o m i c integrates into goniocysts. For a discussion on its inclu-
delimitations above species level. sion in Lecania, see the section Related genera and
I agree with Coppins (in Punis et al. 1992) that the specia groups in the chapter Taxonomic delimitu~iow
European Bacidia accedens is just an intensely pigmen- above species level.
ted morph of B. sabuletomm. The variation from B.
sabuletom to "B. accedens" is occasionally seen in a
single specimen. I have, however, not seen the types in- Lecania subfuscula (Nyl) S. Ekman, comb. nova
volved, and that is why I have listed B. accedens with a
query above. The North American B, accedens sensu Lecidea subfuscula Nyl., Flora, Jena 48: 604 (1865). -
Harris (1977) is a distinct species awaiting formal de- Bacidia subfuscula (Nyl.) Th. Fr., K. svenska Veten-
scription. skPJrad. Forh. 7 (2): 35 (1867). - Type: Iceland, "Reik-
javik", 1860, Lindsay (H-NYL 17363, lectotype select-
ed by Printzen 1995: 223 as "holotype").
Bacidia saxicola Looman Biatora siberiemis, Willey in Rothr., Proc. U.S. natn.
Mus. 7 (1): 6 (1884). - Bacidia siberiemis (Willey in
Bryologist 65: 297 (1963). - Type: Canada, Saskatch- Rothr.) Zahlbr., Cat. Lich. Univ. 4: 146 (1926). - Type:
ewan, "Larnond C. P., 1700 feet, on granite rock", U. S. A., Alaska (as "Siberia"), 1880, Bean (US, lecto-
1960, Looman 600 128 (USAS, holotype). type selected here).

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-

Opera Bet. 127, 1g%

Bacidia subgranulosa (Tuck.) Riddle Bacidia verecundula (Th. Fr.) B.Magn.

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).

Bacidia tetramera @e Not.) Coppins in Gilbert et al.

Lichenologist 20 237 (1988). - Bilimbia tetrainera De

Not., Giorn. Bot. ital. 2: 191 (1846). - Type: Not seen.
Lecidea sphaeroides var o bscurata Somrnerf., Suppl .
F1. Lapp 165 (1826). - Mycobilirnhiu obscuratu (Som-
merf.) Rehm, Rabenh. Krypt.-FI. 2, 1 (3): 328 (1 890). -
Bacidiu ohscumta (Sommerf.) Zahlbr, in Engler, Nat.
Pflanzenfam. 1 (1*): 135 (1905). - Type: Not seen.
Bacidia fusca auct.

This species belongs in Mycobilimbia (Printzen 1995).

It usually overgrows bryophytes and plant debris on the
ground in boreal or arctic areas. I have not examined
any North American material.

Bacidia trachona (Ach.) Lettau

Hedwigia 52: 133 (1912). - Vewucariu trachona Ach.,

Meth. Lich., suppl. 16 (1 803). - Type: Sweden (H-ACH
823, lectotype selected here).

All North American collections named Bacidia rracho-

na are saxicolous. I have studied only a few of them,
and they have turned out to belong to B. granosa. It
seems likely that Bacidicr trachona shouId be excluded
from Bacidia S. str. on account of the pycnidia, which
are are large, multi-chambered, and contain narrow, 0.8-
1.0 pm wide, conidiogenous cells producing small, el-
lipsoid conidia with two guttulae. I have not seen any
fertile material of this species.

upem Bot. 127. 19%

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Opera Bot. 127, 19%

Fig. 40. Apothecial sections of North American species of Bacidia. Specimens collected in North America were used. A . B.
absistens, X103 (Ekman L656, LD). B. B. aggregattlla, X 85 (Ekman L1 188, LD). C. B. arceutina, X 108 (Ekman L696,
- -

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).

Jpera Bot. 127, 19"'

 .
-
ial sections of North American species of Bacidia. Specimens collected in North America were used, unless
-A. B. friesiana, unusually weakly pigmented specimen, but similar to the only North American one, X 140
1942, Almborn, LD). - B. B. helicospora, X 108 (Tucker 11315 & Jones, herb. Tucker). - C. B. heterochroa,
42a, herb. Tucker). -D. B. igniarii, X 103 (Sweden, 1866, Blomberg, LD). -E. B. insularis, X 85 (1994, Nimis
-18385). - F. Bacidia laurocersi subsp. laurocerasi, X 78 (Gowan 3188, CANL). - G. B. polychroa, X 83
4668, MIN). -H. B. reagens, . 130 (Wetmore 65677, MIN).

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).

Opera Bot. 127, 1996

 America were u s e d : A: Bn. aenea, X 130 (Ekman L1207, LD, holotype). - B. Bn. arnoldiana, 98 (Brodo 23794,
. - C. Bn. californica, unusually darkly pigmented specimen (this species is often pigment deficient), X 103 (Bratt 4277,
ratt). - D. Bn. egenula, X 150 (Robitaille 149.4, CANL). - E. Bn. mmea, young apothecium with well-developed
margin, X 123 (Ekman L643, LD). - F. B. diffracts, X 12 (Ekman L1050, LD, holotype). - G. B. helicospora, X 12
11315 & Jones, herb. Tucker). -H. B. salmonea, X 19 (Ekman L753, holotype). i I
' /
:/
Fig. 44. Habitus of North Arnencan species of Baczdzna. -A. Bn aenea, X 15 (Ekman L1207, holotype). - B. Bn callfornzca, X
16 (Ekman L632, LD). - C Bn ciystalllfera, X 20 (Ekman L1120, LD, holotype). - D. Bn ramea, X 14 (Ekman L767, LD,
holotype). - E. Bn squamellosa, X 20 ). -F. Bn varza, X 13 (Ekman L1 191, LD, holotype)

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

opem Bor 127. 1946

pacificum, Haematomma 133 Scoliciosporum 5,38,42-43,45,57,74, 128
pahiensis, Tylothallia 45 swpuIicola 8,134
pdlen~ 57, 132-133 SecoIiga 116,130,133,134
pallida Arnold ex Eckfeldt 133 separabilis* 105
paIlida (Mull. Arg.) Darb. 133 siberiensis* 134
pallidula* 133 Solenopsora 43
palrnularis, Bapalmuia 39 sp. # 1 11,14,
parmnellii* 130 16-17, 19,21,23-25,28-29,33,59,115, 123,125-126, 133
pannaroidea, Psorella 60-6 1 spadicea* 107-108
papantlensis 97-98 speciosum, Echidnocylnbium 40
parvuIa* 97-98 Speerschneidera 43
Patellaria 76,79,82,86,99, 127-128, 133 spermogoniatum,Cliostomwn 40
pateIlarioides, Bactrospora I28 sphaeroides nuct. 128
peltaturn, Rolfidium 42 sphaeroides, Catillaria 42
pertexta, PsorelIa 43,56,135 Sporacestra 5,38,43, 128
Pertusaria 33 Sporopodiurn 4
phacodes 4,19, 112,116-1 18, 133 Squamarina 44
phaeoloma* 97-98 Squarnacidia 42-43
phaeoplaca, Bacidia, Ropalospora* 128 squamellosa 8,10,14,16-19,21,23,25,28-30,33,59,123-124
Phyllopsorn 42 squamulosula, Bacidiopsora 38-39
Physcidia 38,42 stanhopene, Fellhanera, Badimia 132
pilati, Adelolecia 38 stigrnatella,Lecania* 41,57,87, 134
Pilocarpaceae 4, 5 , 3 9 stoechadiana, Waynea 45
p~ly~hroa* 12, 19- stylophora, PhyIlopsora 42
21,23,27-30,32, 34,47-48, 58,73,88-9I,83,96, 109, 126 subacerina 82,84
populorum, Arthrosporum 38,57,133 subacerina, Patellaria* 97
Porpidia 39,132 subacerinoides* 97-98
Porpidiaceae 44 subchlorotica* 118-119
prasina, Biatora 43 subcircumspecta 34-35,71
prasinata, Biatora 43 subhscula, Lecania 41, 134
prolifera, Eschatogonia 40 subgranulosa 135
propinqua 134 subincompta* X, 10, 15, 19-21,23,
pminosum, Scoliciosporum 46 27-31,34,44,58,62, 71, 81-82,92, 101,105-107, 111, 116
Psoraceae 44 subluteola* 68-69
Psorella 38,56,60-61 subpar* 79-80
ramea 10, 12, 14, 16-2 1,23,25,28-30,32,59-60,122-123 subpellucida* 76,78
ravenelii 133 subspadicea* 88
reagens* 8, 12, 19-21,23,28-29,33-34,57,71,81,91-93 subspadicella* 88
requienii, Solenopsora 43 subtestacea* 97-98
Rhaphiospora 67 sufi~a* 8, 10-13, 15,
Rhizocarpon 4 19-21,23,27-29,32,34,59-60,69, 82, 8 8 , 4 3 , 1 0 7 0 126
rissoensis 79-80 suturalis* 97-98
Rolfidium , 42 Tapellaria 4
Ropalospora >,42,56, 128 taylorii, Herteliana 40,127
rosella* 4, 15, 19-21, Teloschistes 33
23,24,28-30,32,34,45,59,88,92,93-94,99,109,120, 126 Tephromela 44
msellina* 86-87 testtlcea 33
rubella* 8-13, 16- tetramera 42, 135
17,19-21,23,27-30,32,34,59-60,67-68,89,93,94-97, 126 thaleriza, Phyllopsora 42
rubidofusca 133 Thamnolecania 41,44
rubricosa, Ophiopma 56, 133-134 TibeIlia 44
mfescens 79-80 Tonitria 4,5, 10,37-38,44-45, 130
W seola* 8, Toniniopsis ' 38,60-61, 130
12,13,15,19,21,23,27-29,33-34,60,69,82,92,97-98, 101 tornoensis, lapewia 41
sabuletorum 45-46,57, 131-132,134-135 trachona* 135
salmonea 19-21,23,28-29,32,34,59-60,65,92-93,98-99 Tylothallia 38,45
sanguineoaba, Lecidea 39 urnbrinum, Scoliciosporum 38,43,133
saxicola 134 varia 9, 11, 14-17, 19-
Schadonia 42 25,28-30,33,58-60,80,87, 115, 1 17, 120,124-126, I33
schistina, Lecanora 22 variegata* 68-69
Schistoplaca 42-43 ventosa, Ophiopama 134
schweinikii* 8, 10-12, 19-24, verecundula* 45,135
27-30,32,34,44,58,64,71,81,98,99-105, 107, 126, 130 vergarensis* 68-69

Opera BoL 127. 1996

verrnifera* 10, 12, 15, 19-24,28-29,3 1,34,60, 105,110-112 vlridis, Ropalospora
vemnlis, Biatora 39 vltellinum, Cliostomum
Verrucana 88,94-95, 135 vulgata, Opegrapha k' T:
. X < , ,>.
.
vexans* 79-80 Waynea
vezdae, Cliostomum 40 Woessia
violascens 92 wrightli, Physcidia

Open Bet. 127. 1596