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1307 Fungi Representing 1139 Infrageneric Taxa, 317 Genera and 66 Families ⇑ Jolanta Miadlikowska A, , Frank Kauff B,1, Filip Högnabba C, Jeffrey C
Molecular Phylogenetics and Evolution 79 (2014) 132–168 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families ⇑ Jolanta Miadlikowska a, , Frank Kauff b,1, Filip Högnabba c, Jeffrey C. Oliver d,2, Katalin Molnár a,3, Emily Fraker a,4, Ester Gaya a,5, Josef Hafellner e, Valérie Hofstetter a,6, Cécile Gueidan a,7, Mónica A.G. Otálora a,8, Brendan Hodkinson a,9, Martin Kukwa f, Robert Lücking g, Curtis Björk h, Harrie J.M. Sipman i, Ana Rosa Burgaz j, Arne Thell k, Alfredo Passo l, Leena Myllys c, Trevor Goward h, Samantha Fernández-Brime m, Geir Hestmark n, James Lendemer o, H. Thorsten Lumbsch g, Michaela Schmull p, Conrad L. Schoch q, Emmanuël Sérusiaux r, David R. Maddison s, A. Elizabeth Arnold t, François Lutzoni a,10, Soili Stenroos c,10 a Department of Biology, Duke University, Durham, NC 27708-0338, USA b FB Biologie, Molecular Phylogenetics, 13/276, TU Kaiserslautern, Postfach 3049, 67653 Kaiserslautern, Germany c Botanical Museum, Finnish Museum of Natural History, FI-00014 University of Helsinki, Finland d Department of Ecology and Evolutionary Biology, Yale University, 358 ESC, 21 Sachem Street, New Haven, CT 06511, USA e Institut für Botanik, Karl-Franzens-Universität, Holteigasse 6, A-8010 Graz, Austria f Department of Plant Taxonomy and Nature Conservation, University of Gdan´sk, ul. Wita Stwosza 59, 80-308 Gdan´sk, Poland g Science and Education, The Field Museum, 1400 S. -
An Evolving Phylogenetically Based Taxonomy of Lichens and Allied Fungi
Opuscula Philolichenum, 11: 4-10. 2012. *pdf available online 3January2012 via (http://sweetgum.nybg.org/philolichenum/) An evolving phylogenetically based taxonomy of lichens and allied fungi 1 BRENDAN P. HODKINSON ABSTRACT. – A taxonomic scheme for lichens and allied fungi that synthesizes scientific knowledge from a variety of sources is presented. The system put forth here is intended both (1) to provide a skeletal outline of the lichens and allied fungi that can be used as a provisional filing and databasing scheme by lichen herbarium/data managers and (2) to announce the online presence of an official taxonomy that will define the scope of the newly formed International Committee for the Nomenclature of Lichens and Allied Fungi (ICNLAF). The online version of the taxonomy presented here will continue to evolve along with our understanding of the organisms. Additionally, the subfamily Fissurinoideae Rivas Plata, Lücking and Lumbsch is elevated to the rank of family as Fissurinaceae. KEYWORDS. – higher-level taxonomy, lichen-forming fungi, lichenized fungi, phylogeny INTRODUCTION Traditionally, lichen herbaria have been arranged alphabetically, a scheme that stands in stark contrast to the phylogenetic scheme used by nearly all vascular plant herbaria. The justification typically given for this practice is that lichen taxonomy is too unstable to establish a reasonable system of classification. However, recent leaps forward in our understanding of the higher-level classification of fungi, driven primarily by the NSF-funded Assembling the Fungal Tree of Life (AFToL) project (Lutzoni et al. 2004), have caused the taxonomy of lichen-forming and allied fungi to increase significantly in stability. This is especially true within the class Lecanoromycetes, the main group of lichen-forming fungi (Miadlikowska et al. -
Lichen Functional Trait Variation Along an East-West Climatic Gradient in Oregon and Among Habitats in Katmai National Park, Alaska
AN ABSTRACT OF THE THESIS OF Kaleigh Spickerman for the degree of Master of Science in Botany and Plant Pathology presented on June 11, 2015 Title: Lichen Functional Trait Variation Along an East-West Climatic Gradient in Oregon and Among Habitats in Katmai National Park, Alaska Abstract approved: ______________________________________________________ Bruce McCune Functional traits of vascular plants have been an important component of ecological studies for a number of years; however, in more recent times vascular plant ecologists have begun to formalize a set of key traits and universal system of trait measurement. Many recent studies hypothesize global generality of trait patterns, which would allow for comparison among ecosystems and biomes and provide a foundation for general rules and theories, the so-called “Holy Grail” of ecology. However, the majority of these studies focus on functional trait patterns of vascular plants, with a minority examining the patterns of cryptograms such as lichens. Lichens are an important component of many ecosystems due to their contributions to biodiversity and their key ecosystem services, such as contributions to mineral and hydrological cycles and ecosystem food webs. Lichens are also of special interest because of their reliance on atmospheric deposition for nutrients and water, which makes them particularly sensitive to air pollution. Therefore, they are often used as bioindicators of air pollution, climate change, and general ecosystem health. This thesis examines the functional trait patterns of lichens in two contrasting regions with fundamentally different kinds of data. To better understand the patterns of lichen functional traits, we examined reproductive, morphological, and chemical trait variation along precipitation and temperature gradients in Oregon. -
One Hundred New Species of Lichenized Fungi: a Signature of Undiscovered Global Diversity
Phytotaxa 18: 1–127 (2011) ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ Monograph PHYTOTAXA Copyright © 2011 Magnolia Press ISSN 1179-3163 (online edition) PHYTOTAXA 18 One hundred new species of lichenized fungi: a signature of undiscovered global diversity H. THORSTEN LUMBSCH1*, TEUVO AHTI2, SUSANNE ALTERMANN3, GUILLERMO AMO DE PAZ4, ANDRÉ APTROOT5, ULF ARUP6, ALEJANDRINA BÁRCENAS PEÑA7, PAULINA A. BAWINGAN8, MICHEL N. BENATTI9, LUISA BETANCOURT10, CURTIS R. BJÖRK11, KANSRI BOONPRAGOB12, MAARTEN BRAND13, FRANK BUNGARTZ14, MARCELA E. S. CÁCERES15, MEHTMET CANDAN16, JOSÉ LUIS CHAVES17, PHILIPPE CLERC18, RALPH COMMON19, BRIAN J. COPPINS20, ANA CRESPO4, MANUELA DAL-FORNO21, PRADEEP K. DIVAKAR4, MELIZAR V. DUYA22, JOHN A. ELIX23, ARVE ELVEBAKK24, JOHNATHON D. FANKHAUSER25, EDIT FARKAS26, LIDIA ITATÍ FERRARO27, EBERHARD FISCHER28, DAVID J. GALLOWAY29, ESTER GAYA30, MIREIA GIRALT31, TREVOR GOWARD32, MARTIN GRUBE33, JOSEF HAFELLNER33, JESÚS E. HERNÁNDEZ M.34, MARÍA DE LOS ANGELES HERRERA CAMPOS7, KLAUS KALB35, INGVAR KÄRNEFELT6, GINTARAS KANTVILAS36, DOROTHEE KILLMANN28, PAUL KIRIKA37, KERRY KNUDSEN38, HARALD KOMPOSCH39, SERGEY KONDRATYUK40, JAMES D. LAWREY21, ARMIN MANGOLD41, MARCELO P. MARCELLI9, BRUCE MCCUNE42, MARIA INES MESSUTI43, ANDREA MICHLIG27, RICARDO MIRANDA GONZÁLEZ7, BIBIANA MONCADA10, ALIFERETI NAIKATINI44, MATTHEW P. NELSEN1, 45, DAG O. ØVSTEDAL46, ZDENEK PALICE47, KHWANRUAN PAPONG48, SITTIPORN PARNMEN12, SERGIO PÉREZ-ORTEGA4, CHRISTIAN PRINTZEN49, VÍCTOR J. RICO4, EIMY RIVAS PLATA1, 50, JAVIER ROBAYO51, DANIA ROSABAL52, ULRIKE RUPRECHT53, NORIS SALAZAR ALLEN54, LEOPOLDO SANCHO4, LUCIANA SANTOS DE JESUS15, TAMIRES SANTOS VIEIRA15, MATTHIAS SCHULTZ55, MARK R. D. SEAWARD56, EMMANUËL SÉRUSIAUX57, IMKE SCHMITT58, HARRIE J. M. SIPMAN59, MOHAMMAD SOHRABI 2, 60, ULRIK SØCHTING61, MAJBRIT ZEUTHEN SØGAARD61, LAURENS B. SPARRIUS62, ADRIANO SPIELMANN63, TOBY SPRIBILLE33, JUTARAT SUTJARITTURAKAN64, ACHRA THAMMATHAWORN65, ARNE THELL6, GÖRAN THOR66, HOLGER THÜS67, EINAR TIMDAL68, CAMILLE TRUONG18, ROMAN TÜRK69, LOENGRIN UMAÑA TENORIO17, DALIP K. -
Lichens and Associated Fungi from Glacier Bay National Park, Alaska
The Lichenologist (2020), 52,61–181 doi:10.1017/S0024282920000079 Standard Paper Lichens and associated fungi from Glacier Bay National Park, Alaska Toby Spribille1,2,3 , Alan M. Fryday4 , Sergio Pérez-Ortega5 , Måns Svensson6, Tor Tønsberg7, Stefan Ekman6 , Håkon Holien8,9, Philipp Resl10 , Kevin Schneider11, Edith Stabentheiner2, Holger Thüs12,13 , Jan Vondrák14,15 and Lewis Sharman16 1Department of Biological Sciences, CW405, University of Alberta, Edmonton, Alberta T6G 2R3, Canada; 2Department of Plant Sciences, Institute of Biology, University of Graz, NAWI Graz, Holteigasse 6, 8010 Graz, Austria; 3Division of Biological Sciences, University of Montana, 32 Campus Drive, Missoula, Montana 59812, USA; 4Herbarium, Department of Plant Biology, Michigan State University, East Lansing, Michigan 48824, USA; 5Real Jardín Botánico (CSIC), Departamento de Micología, Calle Claudio Moyano 1, E-28014 Madrid, Spain; 6Museum of Evolution, Uppsala University, Norbyvägen 16, SE-75236 Uppsala, Sweden; 7Department of Natural History, University Museum of Bergen Allégt. 41, P.O. Box 7800, N-5020 Bergen, Norway; 8Faculty of Bioscience and Aquaculture, Nord University, Box 2501, NO-7729 Steinkjer, Norway; 9NTNU University Museum, Norwegian University of Science and Technology, NO-7491 Trondheim, Norway; 10Faculty of Biology, Department I, Systematic Botany and Mycology, University of Munich (LMU), Menzinger Straße 67, 80638 München, Germany; 11Institute of Biodiversity, Animal Health and Comparative Medicine, College of Medical, Veterinary and Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK; 12Botany Department, State Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany; 13Natural History Museum, Cromwell Road, London SW7 5BD, UK; 14Institute of Botany of the Czech Academy of Sciences, Zámek 1, 252 43 Průhonice, Czech Republic; 15Department of Botany, Faculty of Science, University of South Bohemia, Branišovská 1760, CZ-370 05 České Budějovice, Czech Republic and 16Glacier Bay National Park & Preserve, P.O. -
On Rhizocarpon Viridiatrum (Wulfen) Körb. in North-Eastern Iran
ON RHIZOCARPON VIRIDIATRUM (WULFEN) KÖRB. IN NORTH-EASTERN IRAN M. Haji Moniri & S. Kamyabi Received 27. 07. 2011. Accepted for publication 08.11.2011. Haji Moniri, M. & Kamyabi, S. 2011 12 31: On Rhizocarpon viridiatrum (Wulfen) Körb. in north-eastern Iran. - Iran. J. Bot. 17 (2): 276-281. Tehran. Morphology, anatomy, chemistry and distribution of the species Rhizocarpon viridiatrum (Rhizocarpaceae, Lichenized Fungi) in NE Iran are presented based on observations from 357 samples collected in Razavi Khorasan province during 2006-2007, an analysis of lichen substances and a literature survey. Mahroo Haji Moniri (correspondence <[email protected]>) & Saleh Kamyabi, Department of Biology, Faculty of Science, Mashhad Branch, Islamic Azad University, Mashhad, Iran, Rahnamaie St., Mashhad. Keywords. Lichenized Fungi, Rhizocarpon viridiatrum, Razavi Khorasan. Rhizocarpon viridiatrum (Wulfen) Körb. . . # $ ! " # + , (Rhizocarpaceae, Lichenized Fungi) Rhizocarpon viridiatrum ! (!' &$ !' % , !& 8 9 : ;$' <$9=' &# , 2007 2006 2 #3 2!4 1 1 2 ( &0 !& 357 . ! > (, Introduction Khorasan province, have been reported (Seaward et al. The lichen genus Rhizocarpon introduced by Ramond 2008; Moniri et al. 2009). Razavi Khorasan province in 2 in Flore Francaise ed. III (Lamarck and Candolle, the northeast of Iran with an area of c. 127,432 km 1805) has been subject to taxonomic debate. Originally biogeographically belongs to Irano-Touranian region its species were included in Lecidea, Buellia and (Takhtajan, 1977). Razavi and Northern Khorasan Diplotomma. Since 1871 new attempts have been made provinces have more mountain ranges than any other to delimit a separate genus Rhizocarpon Lam. ex DC. parts in the country, including Ala Dagh, Binaloud, as a cosmopolitan with ca. -
<I> Lecanoromycetes</I> of Lichenicolous Fungi Associated With
Persoonia 39, 2017: 91–117 ISSN (Online) 1878-9080 www.ingentaconnect.com/content/nhn/pimj RESEARCH ARTICLE https://doi.org/10.3767/persoonia.2017.39.05 Phylogenetic placement within Lecanoromycetes of lichenicolous fungi associated with Cladonia and some other genera R. Pino-Bodas1,2, M.P. Zhurbenko3, S. Stenroos1 Key words Abstract Though most of the lichenicolous fungi belong to the Ascomycetes, their phylogenetic placement based on molecular data is lacking for numerous species. In this study the phylogenetic placement of 19 species of cladoniicolous species lichenicolous fungi was determined using four loci (LSU rDNA, SSU rDNA, ITS rDNA and mtSSU). The phylogenetic Pilocarpaceae analyses revealed that the studied lichenicolous fungi are widespread across the phylogeny of Lecanoromycetes. Protothelenellaceae One species is placed in Acarosporales, Sarcogyne sphaerospora; five species in Dactylosporaceae, Dactylo Scutula cladoniicola spora ahtii, D. deminuta, D. glaucoides, D. parasitica and Dactylospora sp.; four species belong to Lecanorales, Stictidaceae Lichenosticta alcicorniaria, Epicladonia simplex, E. stenospora and Scutula epiblastematica. The genus Epicladonia Stictis cladoniae is polyphyletic and the type E. sandstedei belongs to Leotiomycetes. Phaeopyxis punctum and Bachmanniomyces uncialicola form a well supported clade in the Ostropomycetidae. Epigloea soleiformis is related to Arthrorhaphis and Anzina. Four species are placed in Ostropales, Corticifraga peltigerae, Cryptodiscus epicladonia, C. galaninae and C. cladoniicola -
A Multigene Phylogenetic Synthesis for the Class Lecanoromycetes (Ascomycota): 1307 Fungi Representing 1139 Infrageneric Taxa, 317 Genera and 66 Families
A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families Miadlikowska, J., Kauff, F., Högnabba, F., Oliver, J. C., Molnár, K., Fraker, E., ... & Stenroos, S. (2014). A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families. Molecular Phylogenetics and Evolution, 79, 132-168. doi:10.1016/j.ympev.2014.04.003 10.1016/j.ympev.2014.04.003 Elsevier Version of Record http://cdss.library.oregonstate.edu/sa-termsofuse Molecular Phylogenetics and Evolution 79 (2014) 132–168 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families ⇑ Jolanta Miadlikowska a, , Frank Kauff b,1, Filip Högnabba c, Jeffrey C. Oliver d,2, Katalin Molnár a,3, Emily Fraker a,4, Ester Gaya a,5, Josef Hafellner e, Valérie Hofstetter a,6, Cécile Gueidan a,7, Mónica A.G. Otálora a,8, Brendan Hodkinson a,9, Martin Kukwa f, Robert Lücking g, Curtis Björk h, Harrie J.M. Sipman i, Ana Rosa Burgaz j, Arne Thell k, Alfredo Passo l, Leena Myllys c, Trevor Goward h, Samantha Fernández-Brime m, Geir Hestmark n, James Lendemer o, H. Thorsten Lumbsch g, Michaela Schmull p, Conrad L. Schoch q, Emmanuël Sérusiaux r, David R. Maddison s, A. Elizabeth Arnold t, François Lutzoni a,10, -
Further Investigations on Rhizocarpon of North-Eastern Iran: R
Hindawi Publishing Corporation Journal of Mycology Volume 2014, Article ID 528041, 5 pages http://dx.doi.org/10.1155/2014/528041 Research Article Further Investigations on Rhizocarpon of North-Eastern Iran: R. geographicum Mahroo Haji Moniri Department of Biology, Faculty of Sciences, Islamic Azad University, Mashhad Branch, Mashhad 917 56 89 119, Iran Correspondence should be addressed to Mahroo Haji Moniri; h [email protected] Received 15 January 2014; Accepted 7 April 2014; Published 29 April 2014 Academic Editor: Simona Nardoni Copyright © 2014 Mahroo Haji Moniri. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Morphology, anatomy, secondary chemistry, ecology, and distribution of Rhizocarpon geographicum (Rhizocarpaceae, lichenized Ascomycota) in north-eastern Iran are investigated and discussed. 1. Introduction the elevation of the sites ranges from 1340 to 1880 m. Over 537 Rhizocarpon thalli were investigated of which 311 (58%) Iran has an exceptional variety of biomes which has resulted belong to R. geographicum. As a result much new information in an extensive diversity in many organism groups including became available on the species of Rhizocarpon [11–13]. Mor- lichens. The history of lichenological exploration in north- phological and anatomical characteristics of the thalli and eastern Iran is fairly extensive, dating back to the middle fruiting bodies were examined with a stereomicroscope and a of the twentieth century [1, 2]. Since 2004, however, the light microscope. Preparations were made in distilled water, investigations in the present Khorasan provinces have much 10% KOH and KI. -
British Lichen Society Bulletin No
1 BRITISH LICHEN SOCIETY OFFICERS AND CONTACTS 2010 PRESIDENT S.D. Ward, 14 Green Road, Ballyvaghan, Co. Clare, Ireland, email [email protected]. VICE-PRESIDENT B.P. Hilton, Beauregard, 5 Alscott Gardens, Alverdiscott, Barnstaple, Devon EX31 3QJ; e-mail [email protected] SECRETARY C. Ellis, Royal Botanic Garden, 20A Inverleith Row, Edinburgh EH3 5LR; email [email protected] TREASURER J.F. Skinner, 28 Parkanaur Avenue, Southend-on-Sea, Essex SS1 3HY, email [email protected] ASSISTANT TREASURER AND MEMBERSHIP SECRETARY H. Döring, Mycology Section, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, email [email protected] REGIONAL TREASURER (Americas) J.W. Hinds, 254 Forest Avenue, Orono, Maine 04473-3202, USA; email [email protected]. CHAIR OF THE DATA COMMITTEE D.J. Hill, Yew Tree Cottage, Yew Tree Lane, Compton Martin, Bristol BS40 6JS, email [email protected] MAPPING RECORDER AND ARCHIVIST M.R.D. Seaward, Department of Archaeological, Geographical & Environmental Sciences, University of Bradford, West Yorkshire BD7 1DP, email [email protected] DATA MANAGER J. Simkin, 41 North Road, Ponteland, Newcastle upon Tyne NE20 9UN, email [email protected] SENIOR EDITOR (LICHENOLOGIST) P.D. Crittenden, School of Life Science, The University, Nottingham NG7 2RD, email [email protected] BULLETIN EDITOR P.F. Cannon, CABI and Royal Botanic Gardens Kew; postal address Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, email [email protected] CHAIR OF CONSERVATION COMMITTEE & CONSERVATION OFFICER B.W. Edwards, DERC, Library Headquarters, Colliton Park, Dorchester, Dorset DT1 1XJ, email [email protected] CHAIR OF THE EDUCATION AND PROMOTION COMMITTEE: position currently vacant. -
Bulletin of the California Lichen Society
Bulletin of the California Lichen Society Volume 5 No. 1 Summer 1998 The California Lichen Society seeks to promote the appreciation, conservation, and study of the lichens. The focus of the Society is on California, but its interests include the entire western part of the continent. Dues are $15 per year ( $20 for foreign subscribers) payable to The Cal ifornia Li chen Society, 362 Scenic Avenue, Santa Rosa, CA, 95407. Members receive the Bulletin and notices of meetings, field trips, and workshops. The Bulletin of the California Lichen Society (ISSN 1 093-9148) is edited by Isabelle Tavares, Shirley Tucker, William Sanders, Richard Moe, and Darrell Wright and is produced by Richard Moe. The Bulletin welcomes manuscripts on technical topics in lichenology relating to western North America and on conservation of the lichens, as well as news of lichenologists and their activities. Manuscripts may be submitted to Richard Moe, Bulletin of the California Lichen Society, University Herbarium, 1001 Valley Life Sciences Bldg . #2465, University of California, Berkeley, CA 94 720-2465. The best way to submit manuscripts apart from short articles and announcements is by E-mail or on diskette in Word Perfect or Microsoft Word format; ASCII format is a very good alternative. Manuscripts should be double-spaced. Figures are the usual line drawings and sharp black and white glossy photos, unmounted, and must be sent by surface mail. A review process is followed . Nomenclature follows Esslinger and Egan's Sixth Checklist (The Bryologist 98: 467-549, 1995). and subsequent on-line updateshttp://www.ndsu.nodak.edul instruct/chcklst/chcklst7 .htm. -
Ecological Specialization of Lichen Congeners with a Strong Link to Mediterranean-Type Climate: a Case Study of the Genus Solenopsora in the Apennine Peninsula
Università degli Studi di Trieste ar Ts Archivio della ricerca – postprint Ecological specialization of lichen congeners with a strong link to Mediterranean-type climate: a case study of the genus Solenopsora in the Apennine Peninsula Anna GUTTOVÁ, Zuzana FAČKOVCOVÁ, Stefano MARTELLOS, Luca PAOLI, Silvana MUNZI, Elena PITTAO and Silvia ONGARO Abstract: An ecological biogeographical perspective provides an understanding of the factors that shape the geographical distribution of organisms, their biodiversity and ecological speciation. Focusing on members of the lichen genus Solenopsora, which are strongly linked to a Mediterranean-type climate, we aimed to depict their environmental niches in the Apennine Peninsula. This area represents their ecological optima, as well as the biogeographical centre of distribution. On the basis of occurrences of Solenopsora congeners, we identified the key ecological factors that shape their environmental niches. Applying an ensemble approach, which merges the results of Random Forest, GLM and MaxEnt algorithms, suitability maps were developed. These are mainly influenced by geological substratum, temperature and precipitation. Occurrence of Solenopsora taxa seems to be mainly governed by low variability in diurnal temperature and tolerance to dryness, with precipitation in the range of 0–20 mm in the driest month and a minimum temperature of >5 °C in the coldest month. The sensitivity to diurnal temperature, an important indicator for climate change, suggests that the taxa confined to Mediterranean bioclimatic types (i.e. Solenopsora grisea, S. marina, S. olivacea subsp. olbiensis and S. olivacea subsp. olivacea) might be good indicators of climatic stability. The geological substratum was a strong limiting factor and separated the taxa into three groups: those growing on calcareous, siliceous and ultramafic substrata.