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Molecular analyses of the recently described Xanthoria polessica collected from the type locality and of the Australian species X. coomae were used to determine whether these taxa are distinct species or fall within the wide phenotypic variation of the well-known and cosmopolitan X. parietina. Our results clearly indicate that both taxa should be considered as synonyms of X. parietina since the infraspecific morphological range is accommodated by the observed variation in the thallus and lobe size, their color, position of apothecia, the shape of ascospores, and width of ascospore septum.

The lichen family Teloschistaceae from the Galapagos is revised. Most of the species belong to the Caloplacoideae, two to Teloschistoideae and a few to Xanthorioideae, three subfamilies not validly published, which is remedied here. Four different datasets were analyzed using Bayesian inference. For the bulk of the species, a combined dataset of nrITS, nrLSU and mrSSU was analyzed. Additionally, three analyses were performed using nrITS to further investigate phylogenetic relationships within and between species in each subfamily, and in the genera Xanthomendoza and Squamulea. Four new genera are described: Lacrima, Oceanoplaca, Phaeoplaca, Sucioplaca. Twenty-four species are reported, of which ten are new to science: Caloplaca nigra, Lacrima galapagoensis, Oceanoplaca chemoisidiosa, O. sideritoides, Phaeoplaca tortuca, Squamulea chelonia, S. humboldtiana, S. osseophila, S. oceanica, and Xanthomendoza leoncita. Several new combinations are proposed and three species of Xanthomendoza are reduced to synonymy. Several new combinations and species placed into synonymy do not occur in the Galapagos, but are treated as a consequence of our taxonomic revision. Morphology, anatomy, secondary chemistry, distribution and molecular phylogenetic affiliation are presented for each species and a key is provided. Eight different chemical patterns are quantitatively described based on HPLC analyses. The new genus Lacrima includes L. galapagoensis, a species without vegetative propagules, and two densely isidiate species, L. epiphora and L. aphanotripta that are morphologically similar to ‘Caloplaca’ wrightii. The only species of Galapagos Teloschistaceae that contains xanthones is placed into Huneckia. Oceanoplaca includes two species with the new anthraquinone isidiosin, O. isidiosa and O. chemoisidiosa, while a third species, O. sideritoides, does not contain this secondary metabolite. Phaeoplaca camptidia has previously been reported from Galapagos, but our phylogenetic analysis suggests that it is a new species, here named P. tortuca. An isolated position is occupied by ‘Caloplaca’ diplacia, which we place in it its own monotypic genus Sucioplaca. Some Galapagos Teloschistaceae can be considered a ‘residue’ of unresolved Caloplaca s.l., i.e. the corticolous C. floridana is possibly related to the saxicolous C. nigra, while C. cupulifera can currently not be placed. Squamulea remains particularly problematic and includes S. phyllidizans, that is nested among otherwise unresolved Squamulea species. Based on molecular data, S. phyllidizans is close to ‘Huriella’. ‘Huriella’ flakusii, described from Peru, is confirmed to occur in the Galapagos and the genus is reduced to synonymy with Squamulea. The Squamulea squamosa/subsoluta group remains largely unresolved, but the new species S. chelonia, S. humboldtiana, S. oceanica, and S. osseophila are phylogenetically distinct. Foliose Teloschistaceae are represented only by one species, described as Xanthomendoza leoncita, while the only fruticose species, Teloschistes chrysophthalmus and T. flavicans, are cosmopolitan.

A replacement name Obscuroplaca for the illegitimate genus Phaeoplaca is introduced. The three known species are transferred as Obscuroplaca camptidia, O. ochrolechioides, and O. tortuca.

We inventoried lichens in Kenai Fjords National Park in Alaska, USA We assembled the known information on occurrence and ecology of lichens in this park by combining field, herbarium, and literature studies. Our results provide baseline data on lichen occurrence that may be used in resource condition assessments, vulnerability assessments, long-term ecological monitoring, and resource management. We report a total of 616 taxa of lichenized fungi from the Park, plus an additional five subspecies and three varieties, all of which are new additions to the National Park Service database for this park unit. An additional five species of nonlichenized lichenicolous fungi are reported here. Eight non-lichenized fungi that are traditionally treated with lichens are also included, most of these associated with bark of particular host species. Four taxa new to North America are reported here (Arctomia delicatula var. acutior, Aspicilia dudinensis, Myriospora myochroa, and Ochrolechia bahusiensis), along with 44 species new to Alaska. Numerous species have been confirmed using ITS barcoding sequences. Also several records assigned to the genus level are reported, many of those are likely new species.

Blastenia is a widely distributed lichen genus in Teloschistaceae. We reconstructed its phylogeny in order to test species delimitation and to find evolutionary drivers forming recent Blastenia diversity. The origin of Blastenia is dated to the early Tertiary period, but later diversification events are distinctly younger. We recognized 24 species (plus 2 subspecies) within 6 infrageneric groups. Each species strongly prefers a single type of substrate (17 species occur on organic substrates, 7 on siliceous rock), and most infrageneric groups also show a clear substrate preference. All infrageneric groups tend to have the Mediterranean and Macaronesian distribution, but some epiphytic species have much larger geographic ranges and some evolved after a long-distance dispersal outside the region. Chlorinated and nonchlorinated anthraquinone chemosyndromes co-occur in apothecia of most species, but the chemotype has been secondarily reduced in some lineages. One infrageneric group has a marked reduction in apothecial size, associated with a substrate shift to twigs. Only seven species have vegetative diaspores; they also produce apothecia but have smaller ascospores. Genome sizes (22-35 Mb in Blastenia) are significantly higher in epilithic species. Within-species genetic variation is low in widely distributed species but high in some epilithic species with small geographical ranges. New taxa are: B. afroalpina, B. anatolica, B. caucasica, B. gennargentuae, B. herbidella subsp. acidophila, B. lauri, B. monticola, B. palmae, B. psychrophila, B. purpurea, B. relicta, B. remota, B. xerothermica, and B. xerothermica subsp. macaronesica. New combinations are: B. festivella and B. subathallina; both names and B. catalinae are lectotypified. (Less)

The lichen family Teloschistaceae is one of the larger families of lichenized fungi. Currently it includes one very large heterogenous genus, Caloplaca, with some 1000 or more species with a vast variation in morphology, anatomy and chemistry. The rest of the family is split into 1015 smaller genera, each with 20 or fewer species. There is no modern classification of the family based on molecular data. Here we attempt a first phylogenetic evaluation of a large part of the family, including a total of 337 species. Of these, 162 were used in a combined phylogenetic analysis of the ribosomal RNA sequence markers nrITS, nrLSU and mrSSU, using Bayesian inference. We also analysed all species using nrITS data, split into four different analyses. As a result we propose a new classification of the family, where a total of 39 genera are recognized, of which 31 are newly described or resurrected. The new genera are: Athallia, Austroplaca, Bryoplaca, Calogaya, Cerothallia, Flavoplaca, Gondwania, Haloplaca, Orientophila, Pachypeltis, Parvoplaca, Rufoplaca, Shackletonia, Scutaria, Sirenophila, Solitaria, Squamulea, Stellarangia, Teloschistopsis, Usnochroma, Variospora, Villophora and Wetmoreana. Resurrected genera are Blastenia, Dufourea, Follmannia, Gyalolechia, Leproplaca, Polycauliona, Pyrenodesmia and Xanthocarpia. The species Orientophila subscopularis is described as new. A third subfamily, Teloschistoideae, is proposed to accommodate the genus Teloschistes and related genera, parallel to the two previously recognized subfamilies Xanthorioideae and Caloplacoideae. We also show the large plasticity in both morphological and anatomical characters between closely related species within genera, indicating the low value of these as evolutionary markers. The secondary chemistry is a better marker in some parts of the family. We recognize a large number of geographically delimited clades with clear centres of evolution, but often showing large variation in morphology and anatomy. (Less)

23 species of the lichen genus Caloplaca from Svalbard are described and/or discussed. The descriptions are natural language descriptions based on characters for each species coded into LIAS (Global Information System for Lichenized and Non-Lichenized Ascomycetes). A total of 37 Caloplaca species are listed for Svalbard, of which two species: C. elvebakkiana and C. scabrosa are described as new to science. C. alaskensis is recorded for the first time from Svalbard. A key to the Caloplaca species known from Svalbard is provided.

A molecular phylogeny of the Lecanora rupicola group is presented, based on ITS sequence analyses. The study includes saxicolous and corticolous members of the Lecanora rupicola group as well as other Lecanora species with pruinose apothecia. A phylogenetic hypothesis for species in Lecanora s. lat. and various other genera in Lecanoraceae, based on an alignment-free distance estimation technique, shows that the Lecanora rupicola group forms a monophyletic clade within Lecanoraceae. Affinities to the core group of Lecanora are not well supported, likewise the monophyly of Lecanora s. str. with other species groups in Lecanora, such as the lobate taxa (and Rhizoplaca) is not supported. A more detailed analysis involving Lecanora species with pruinose apothecial discs was carried out with model-based Bayesian Markov chain Monte Carlo (B/MCMC) tree sampling. The results suggest the monophyly of the Lecanora species that are characterized by the presence of chromones. Corticolous as well as saxicolous species are included. Lepraria flavescens is closely related to the Lecanora swartzii subgroup, and the new name Lecanora rouxii nom. nov. is introduced for that species. Other Lecanora species with pruinose discs are riot closely related to the Lecanora rupicola group.

Caloplaca gyalectoides S. Kondratyuk & Karnefelt from the Cape Province, South Africa, and Caloplaca chilensis S. Kondratyuk, Karnefelt, Froden & Arup with an associated parasymbiont Arthonia tetraspora S. Kondratyuk & Karnefelt occurring mainly along the coast in central Chile are described as new. Possible associated taxa are briefly discussed. Distribution maps are provided for all three species.

Rhizoplaca Zopf is a genus characterized by an umbilicate thallus with an upper and a lower cortex, as well as a cupulate hypothecium. It has been considered to be related to Lecanora Ach., the type genus of the Lecanoraceae and, in particular, to the lobate species of this genus. The phylogeny of Rhizoplaca, the monotypic Arctopeltis thuleana Poelt, and a number of representatives of different groups of Lecanora is studied, using sequences from the nuclear ribosomal internal transcribed spacer (ITS) regions. The results suggest an origin for Rhizoplaca species within the large genus Lecanora. A well-supported monophyletic assemblage includes the umbilicate type species Rhizoplaca melanophthalma (DC.) Leuck. & Poelt, the lobate Lecanora novomexicana H. Magn., and five vagrant Rhizoplaca species. Rhizoplaca chrysoleuca (Sm.) Zopf and Rhizoplaca subdicrepans (Nyl.) R. Sant. form a separate well-supported group and Rhizoplaca peltata (Ram.) Leuck. & Poelt is more closely related to Lecanora muralis (Schreb.) Rabenh. Together with data on secondary chemistry, the results show that the umbilicate thallus with a lower and an upper cortex, as well as apothecia with a cupulate hypothecium found in Rhizoplaca and A. thuleana, have developed several times in independant lineages in Lecanora. The thallus morphology in lecanoroid lichens is highly variable and does not necessarily reflect phylogenetic relationships.Key words: Rhizoplaca, Lecanora, Lecanorales, phylogeny, ITS.

The phylogeny of the lichen family Candelariaceae was investigated using nucleotide sequences from the ITS region of the nu-rDNA. Twenty-three species of Candelariella, six species of Candelaria, two species of Candelina and two species of Placomaronea were included in the study. Acarospora cervina and Pleopsidium chlorophanum were used as outgroup species. The phylogenetic analyses were performed using MP and Bayesian MCMC inference. The resulting trees were poorly resolved and strong support was only found for terminal clades. However, the results indicate that polyspored asci have evolved a limited number of times within the family and appear within four clades. One of these clades comprises the core group of Candelariella, including the type species C. vitellina. Placomaronea and Candelina both form strongly supported monophyletic clades, but neither genera are distinctly morphologically separated from Candelariella, and their positions in the tree are uncertain. The genus Candelaria is probably polyphyletic and should possibly be restricted to comprise only polyspored species with a lower cortex.

We compiled literature, intensively studied 15 sites as a group, and collected opportunistically in other areas of the Yosemite National Park. We report a total of 562 species of lichenized fungi from the Park, adding 461 species to the total of 101 species reported for the Park by the National Park Service database. An additional 22 lichenicolous fungi are reported here. Two nonlichenized fungi associated with young living twigs of particular host species are also included. An additional 75 species that are known from nearby areas in the Sierra Nevada, but not yet from Yosemite, are listed. Fourteen species are apparently newly reported for the Sierra Nevada, with an additional 17 species new to California, and five species new to North America ( Gyalidea fritzei, Pyrenopsis reducta, Lecanora pseudosarcopidoides, L. sarcopidoides, L. subravida ). Two taxonomic changes are included here: Verrucaria carbonusta Breuss is newly described, and Lecidea fuscoatrina Hertel & Leuckert is synonymized under the earlier but neglected name, L. cascadensis H. Magn.

The recently invalidly published name Teloschistaceae subfam. Teloschistoideae Arup, Sochting & Froden subfam. nov. is validated here by providing a diagnosis for the taxon (ICN Art. 38).

The number of undescribed species of lichenized fungi has been estimated at roughly 10,000. Describing and cataloging these would take the existing number of taxonomists several decades; however, the support for taxonomy is in decline worldwide. In this paper we emphasize the dire need for taxonomic expertise in lichenology. We bring together 103 colleagues from institutions worldwide to describe a total of 100 new species of lichenized fungi, representing a wide taxonomic and geographic range. The newly described species are: Acarospora flavisparsa, A. janae, Aderkomyces thailandicus, Amandinea maritima, Ampliotrema cocosense, Anomomorpha lecanorina, A. tuberculata, Aspicilia mansourii, Bacidina sorediata, Badimia multiseptata, B. vezdana, Biatora epirotica, Buellia sulphurica, Bunodophoron pinnatum, Byssoloma spinulosum, Calopadia cinereopruinosa, C. editae, Caloplaca brownlieae, C. decipioides, C. digitaurea, C. magnussoniana, C. mereschkowskiana, C. yorkensis, Calvitimela uniseptata, Chapsa microspora, C. psoromica, C. rubropulveracea, C. thallotrema, Chiodecton pustuliferum, Cladonia mongkolsukii, Clypeopyrenis porinoides, Coccocarpia delicatula, Coenogonium flammeum, Cresponea ancistrosporelloides, Crocynia microphyllina, Dictyonema hernandezii, D. hirsutum, Diorygma microsporum, D. sticticum, Echinoplaca pernambucensis, E. schizidiifera, Eremithallus marusae, Everniastrum constictovexans, Fellhanera borbonica, Fibrillithecis sprucei, Fissurina astroisidiata, F. nigrolabiata, F. subcomparimuralis, Graphis caribica, G. cerradensis, G. itatiaiensis, G. marusa, Gyalideopsis chicaque, Gyrotrema papillatum, Harpidium gavilaniae, Hypogymnia amplexa, Hypotrachyna guatemalensis, H. indica, H. lueckingii, H. paracitrella, H. paraphyscioides, H. parasinuosa, Icmadophila eucalypti, Krogia microphylla, Lecanora mugambii, L. printzenii, L. xanthoplumosella, Lecidea lygommella, Lecidella greenii, Lempholemma corticola, Lepraria sekikaica, Lobariella sipmanii, Megalospora austropacifica, M. galapagoensis, Menegazzia endocrocea, Myriotrema endoflavescens, Ocellularia albobullata, O. vizcayensis, Ochrolechia insularis, Opegrapha viridipruinosa, Pannaria phyllidiata, Parmelia asiatica, Pertusaria conspersa, Phlyctis psoromica, Placopsis imshaugii, Platismatia wheeleri, Porina huainamdungensis, Ramalina hyrcana, R. stoffersii, Relicina colombiana, Rhizocarpon diploschistidina, Sticta venosa, Sagenidiopsis isidiata, Tapellaria albomarginata, Thelotrema fijiense, Tricharia nigriuncinata, Usnea galapagona, U. pallidocarpa, Verrucaria rhizicola, and Xanthomendoza rosmarieae. In addition, three new combinations are proposed: Fibrillithecis dehiscens, Lobariella botryoides, and Lobariella pallida.

A total of 20 species of Caloplaca have been found to occur, obligately or occasionally, in the littoral and supralittoral zones on seashore rocks in continental North America. Eighteen species are marine to maritime and two are mainly inland species with occasional occurrence on the shore. On the west coast there are 16 species occurring on seashore rocks, while the east coast contains only five species. Europe and North America share 40% of the littoral species. A total of 12 species seem to be endemic to western North America. None of the 18 marine to maritime species is known to occur also in the southern hemisphere. A key, a short description for each of the treated species, and statement of their distribution is given. The species are also depicted in color photos.

Twenty species ofCaloplaca occur on seashore rocks in North America. Twelve of these have been studied taxonomically in previous papers, and the remaining eight are included in this study. Four of the eight species are new to science: C. ignea, C. impolita, C. inconspecta, and C. ludificans. All four species seems to be endemic to western North America. Caloplaca ignea and C. impolita are members of section Gasparrinia, while C. inconspecta and C. ludificans belong to section Caloplaca. The otherfour species--C. alcarum Poelt, C. brattiae W. A. Weber, C. coralloides (Tuck.) Hult., and C. litoricola Brodo-have been poorly known in some respects. Knowledge about C. alcarum in continental North America is still very poorsince only very scanty material is available. In a series of articles I have reported on the species of Caloplaca occurring on seashore rocks in North America (Arup 1992a,b, 1993a,b, 1994). This paper completes that series and deals with a remaining group of eight species. Four of these species are new to science, and the remaining four have been poorly known in some respects, e.g., relation to other spe- cies, ecology, or distribution. Two of the new species, C ignea and C. impolita, have well-developed marginal lobes and belong to section Gasparrinia. The other new species, C in- conspecta and C ludificans, are members of section Caloplaca. The new species could not be matched with any already existing taxa in the literature from Mexico and New Mexico (Bouly de Lesdain 1914, 1929, 1932, 1933, 1942), South America (e.g., Malme 1926), Europe (e.g., Clauzade & Roux 1985), or North America (e.g., Rudolph 1955). In Arup (1995) there is a key to all the species of Caloplaca occurring on seashore rocks in North America.

The species ofthe genus Caloplaca occurring on seashore rocks in eastern North America have been studied with regard to taxonomy, geographical distribution, and ecology. Five species were found to inhabit the coastal rocks. The species also occur in Europe, and three of them in Asia. Three of the species, C. microthallina (Wedd.) Zahlbr., C. scopularis (Nyl.) Lettau, and C. verru- culifera (Vainio) Zahlbr. occur exclusively on seashore rocks, while C. lithophila H. Magn. and C. fraudans (Th. Fr.) H. Olivier occur also in inland localities. One-way analysis of variance of both morphological and anatomical characters was carried out on the American and the European populations of the species, except C. fraudans. No significant differences were found in C. lithophila, but in the other three species significant differences were recorded in two or three characters. However, the differences were mostly very small, the overlaps large, and no discontinuities could be detected. Therefore no infraspecific taxa were recognized. The lichen flora occurring on seashore rocks in eastern North America has previously been studied on several occasions. In the late 19th and the early 20th century contributions to the knowledge of lit- toral lichens were made mainly with scattered col- lections by botanists such as E. Tuckerman, H. Wil- ley, W. G. Farlow, and G. K. Merrill. Degelius (1940) was one of the first lichenologists to investigate the lichens on seashore rocks in North America more thoroughly. He studied the flora of Maine and com- pared it with the European flora, and found great similarities. The work of Degelius inspired I. M. Lamb (1954) to carry out similar studies in Nova Scotia, Canada. The most extensive collections of littoral lichens in eastern North America were made by R. M. Tay- lor. His work mainly concerned the vertical distri- bution patterns, interspecific associations, and the general geographical distribution of the species col- lected, but also comprised taxonomical studies of the genus Verrucaria and a key to the 21 species included in the study (Taylor 1974). One of the latest studies in eastern North America, including littoral lichens, is that of the flora of Fundy National Park, New Brunswick, Canada, by Gowan and Bro- do (1988). They pointed out that the maritime prov- inces are still not very well known and that the need for further studies is high. This study was carried out to investigate the spe- cies of Caloplaca occurring on seashore rocks in eastern North America, and to compare the Euro- pean and the American populations of the species. No extensive studies have previously been done on this group in this region. Three of the species, oc- curring also in northern Europe, have been treated by Nordin (1972), but only for the European range of the species.

Caloplaca luteominia (Tuck.) Zahlbr. and C. bolanderi (Tuck.) Magn. both have a western, marine-maritime distribution in North America. They are similar in most respects but differfrom each other in the color of their apothecia and pycnidia, which in C. luteominia are orange and in C. bolanderi scarlet. Caloplaca luteominia and C. bolanderi are very closely related and can only be separated by the color of the apothecia and pycnidia, which is due to chemical differences. The new combination C. luteominia var. bolanderi (Tuck.) Arup is therefore suggested. Caloplaca laeta Magn. is a synonym of C. luteominia. Lectotypes for Placodium luteominium and Blastenia ferruginea fo. bolanderi are selected.

Two sorediate species ofCaloplaca occur on seashore rocks in North America, Caloplaca flavogranulosa sp. nov., and C. citrina (Hoffm.) Th. Fr. Caloplaca citrina s.l. occurs on many substrates in many different environments, but the treatment of C. citrina in this paper comprises only the form occurring on seashore rocks. This form may represent a distinct taxon, separate from C. citrina s.s., but more extensive studies of the entire complex must be carried out to settle this problem. Caloplaca flavogranulosa is described herefor the first time. It has previously been confused with several species, mostly nonsorediate. Caloplaca flavogranulosa is characterized by a discon- tinuous areolate to lobate or squamulose, yellowish-orange thallus, prominent prothallus, coarse granules, and soredia which are marginal or covering most of the thallus. Caloplaca flavogranulosa differs from C. citrina by having prominent prothallus, convex areoles with a narrow base, granules, and larger soredia.

Caloplaca bolacina (Tuck.) Herre and C. stantonii sp. nov. belong to a group of species in North America with more or less lobate, subsquamulose to squamulose thalli. The two species occur mainly on rocks in the coastal mountains and on maritime rocks in western North America. Caloplaca bolacina is here lectotypified and provided with a thorough description. Caloplaca stan- tonii, sometimes confused with C. bolacina, was in 1989 distributed in Lich. Exs. Colo No. 672 by W. A. Weber but is described for the first time in this paper. The present study was carried out to clarify the taxonomy of C. bolacina and C. stantonii and their relationship to a group of similar species. This group, here referred to as the C. squamosa group, comprises many species, both European andAmerican. A key to the treated taxa is included. The taxonomy within the C. squamosa group is not dealt with. The results show that C. bolacina, C. stantonii, and the C. squamosa group are separated from each other principally in several anatomical characters as well as in gross morphology.

Caloplaca rosei Hasse and C. marina (Wedd.) Zahlbr. both grow on seashore rocks and are widely distributed along the North American west coast. Caloplaca marina occurs in Europe as well, but not on the North American east coast. Due to their similarities to one another and the lack of correctly determined specimens in herbaria, the two species have been very much misunderstood in North America. Thepresent study, based on my own material and on herbarium material, was carried out to clarify their relationship and to establish whether or not the North American populations of C. marina are conspecific with those in Europe. A large set of morphological and anatomical characters was studied and analyzed using a computer-assisted principal components analysis. The results show that C. rosei and C. marina, even though closely related, are well separated by morphological traits such as continuity, areolation, and surface of the thallus, together with the shape, disk, and thalline margin of the apothecia. Within C. marina two subgroups could be recognized, corresponding well with their disjunct distribution on the two continents. The American population is designated C. marina subsp. americana Arup, differing from subsp. marina mainly in ecology but also having narrower spores and spore septum and a thicker hymenium. Caloplaca rosei Hasse and C. marina (Wedd.) Zahlbr. are two closely related lichen species within the large family Teloschistaceae. The two species are characterized by their yellow to orange, crustose, areolate thalli, often distinct hypothallus, and abundant lecanorine apothecia. Both species grow mainly in the supralittoral zone on seashore rocks and are part of the "orange belt" just above the black Verrucaria zone. Sometimes, however, they also occur in adjacent zones. Caloplaca rosei is rather common along most of the American west coast, but does not occur on the American east coast. Caloplaca marina is a more widespread species, occurring not only in western North America but also in Europe, from where it originally was described. It has also been reported several times from the American east coast (e.g., Degelius 1940; Lamb 1954; Taylor 1974) but all these records seem to be erroneous. The taxonomy of C. marina in Europe has been treated by Nordin (1972), but the relationship of C. rosei and C. marina has never been scrutinized. The two species have been misunderstood in North America for several reasons. In addition to being similar to one another in many respects, they have been confused with other species as well. The fact that they often grow together (Fig. 1), and even upon each other, may be part of the reason there are hardly any correctly determined specimens in herbaria. Most reports of one or the other species refer to a mixture of both (e.g., Benton et al. 1977; Noble 1982; Otto 1968; Ryan 1988). The purpose of this study was to clarify the taxonomic relationship between C. marina and C. rosei, and to obtain a good picture of their geographical distribution and ecology in North America. I also wanted to study geographical variation within the species and to find out if C. marina in Europe and North America are conspecific. The study is part of my doctoral thesis project on "The Littoral Species of Caloplaca in North America." MATERIALS AND METHODS The study is based primarily on materials collected on a field trip to North America in 1989. I spent eight weeks on the west coast where I collected from the Los Angeles area up to Vancouver Island in British Columbia, and three weeks on the east coast, from Massachusetts to Quebec. In addition to my own material I have studied herbarium material from the following institutions: ASU, Bratt (priv. herb.), CANL, COLO, FH, LAM, LD, LISU, MIN, S, SFSU, TUR, UBC, UPS, WTU, and WWB. The collections of C. marina from Europe included in the study were selected to give a good representation from throughout the European range of the species. A total of almost 230 specimens of both species are included in the study. My own collections are deposited at the Botanical museum in Lund (LD). The specimens were examined by interference and light microscopy. Sections of the apothecia were made both by hand and with a freezing microtome and embedded in water or in phenol-cottonblue. All measurements of anatomical features were carried out on sections mounted in water. 0007-2745/92/148-160$1.45/0 This content downloaded from 157.55.39.35 on Thu, 01 Sep 2016 05:37:25 UTC All use subject to http://about.jstor.org/terms 1992] ARUP: CALOPLACA MARINA AND C. ROSEI 149 The chemistry was examined by high performance thinlayer chromatography (HPTLC) as described by Arup et al. (1992), a somewhat different procedure than the standardized method developed by Culberson (1972, 1974). The plates used were glass HPTLC-Fertigplatten Merck no. 5629, silica-gel 60 F254, 0.2 mm thick. The silicon particles on these plates are smaller and of more uniform size than on the normal TLC plates, resulting in an ability to detect also low concentrations of the substances applied. The chamber used was a Camag Horizontal Developing Chamber, 10 x 10 cm. This method differs from the normal in that two fronts of the same solvent move toward each other, meeting in the middle on a horizontal plate, and the amount of substance applied on the plate is much smaller, ca. 3 01. The solvents used were: A, toluene: dioxan: acetic acid (18:6:0.8 ml); B, cyclo-hexane: methyl tert.-butyl ether : formic acid (13:10:2 ml); and C, toluene: acetic acid (20:3 ml). A large set of morphological and anatomical characters was studied, and 19 of these were scored on a total of 118 specimens (42 specimens of C. rosei, 39 American and 37 European specimens of C. marina) and analyzed using principal components analysis (PCA), run on a Macintosh SE 30 computer. A PCA simplifies multivariate data by assigning most of the variation to a smaller number of "principal components" (Dunn & Everitt 1982). All calculations were performed with SYSTAT (Wilkinson 1986). Only specimens for which all 19 traits could be scored were used in the analysis. The characters used in the PCA are listed in Tables 1 and 2 and discussed in the following section.