A Contribution to an Inventory of Lichens from South Sister, Northeastern Tasmania Introduction Methods
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Icmadophila Aversa and Piccolia Conspersa, Two Lichen Species New to Bolivia
Polish Botanical Journal 55(1): 217–221, 2010 ICMADOPHILA AVERSA AND PICCOLIA CONSPERSA, TWO LICHEN SPECIES NEW TO BOLIVIA KARINA WILK Abstract. The species Icmadophila aversa and Piccolia conspersa are reported as new to the lichen biota of Bolivia. The studied material was collected in Madidi National Park (NW Bolivia). The species are briefl y characterized and their ecology and distribution are discussed. Key words: lichenized fungi, new records, Madidi region, Andes, South America Karina Wilk, Laboratory of Lichenology, W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland; e-mail: [email protected] INTRODUCTION Bolivia is still one of the countries least studied While studying the material collected in the biologically, but the data already available indi- Madidi region I identifi ed two interesting lichen cate a potentially high level of biodiversity (Ibisch species – Icmadophila aversa and Piccolia con- & Mérida 2004). Knowledge of the cryptogams, spersa. The species are reported here as new to Bo- including lichens, is particularly defi cient (Feuerer livia. Brief descriptions and notes on their ecology et al. 1998). In the last decade, however, licheno- and worldwide distribution are provided. logical studies have progressed in Bolivia. The most recent works have provided many new dis- MATERIAL AND METHODS coveries: records new to the country, continent or Southern Hemisphere, and species new to The study is based on material collected in 2006–2007 in science (e.g., Ferraro 2002; Feuerer & Sipman Madidi National Park. The collection sites are located in 2005; Flakus & Wilk 2006; Flakus & Kukwa 2007; the Cordillera Apolobamba (Fig. -
H. Thorsten Lumbsch VP, Science & Education the Field Museum 1400
H. Thorsten Lumbsch VP, Science & Education The Field Museum 1400 S. Lake Shore Drive Chicago, Illinois 60605 USA Tel: 1-312-665-7881 E-mail: [email protected] Research interests Evolution and Systematics of Fungi Biogeography and Diversification Rates of Fungi Species delimitation Diversity of lichen-forming fungi Professional Experience Since 2017 Vice President, Science & Education, The Field Museum, Chicago. USA 2014-2017 Director, Integrative Research Center, Science & Education, The Field Museum, Chicago, USA. Since 2014 Curator, Integrative Research Center, Science & Education, The Field Museum, Chicago, USA. 2013-2014 Associate Director, Integrative Research Center, Science & Education, The Field Museum, Chicago, USA. 2009-2013 Chair, Dept. of Botany, The Field Museum, Chicago, USA. Since 2011 MacArthur Associate Curator, Dept. of Botany, The Field Museum, Chicago, USA. 2006-2014 Associate Curator, Dept. of Botany, The Field Museum, Chicago, USA. 2005-2009 Head of Cryptogams, Dept. of Botany, The Field Museum, Chicago, USA. Since 2004 Member, Committee on Evolutionary Biology, University of Chicago. Courses: BIOS 430 Evolution (UIC), BIOS 23410 Complex Interactions: Coevolution, Parasites, Mutualists, and Cheaters (U of C) Reading group: Phylogenetic methods. 2003-2006 Assistant Curator, Dept. of Botany, The Field Museum, Chicago, USA. 1998-2003 Privatdozent (Assistant Professor), Botanical Institute, University – GHS - Essen. Lectures: General Botany, Evolution of lower plants, Photosynthesis, Courses: Cryptogams, Biology -
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. -
(1828-1880): a Catalogue of Lichens Collected in Scandinavia in 1863 Housed in the Herbarium, National Botanic Gardens, Dublin
Glasra 4: 63–84 (2000) Isaac Carroll (1828-1880): a catalogue of lichens collected in Scandinavia in 1863 housed in the Herbarium, National Botanic Gardens, Dublin HOWARD F. FOX National Botanic Gardens, Glasnevin, Dublin 9, Ireland. MARY. J. P. SCANNELL Raglan Road, Ballsbridge, Dublin 4, Ireland. ABSTRACT: A lichen collection, containing 181 numbered packets from Scandinavia, made by the Irish botanist Isaac Carroll during 1863 is documented. Brief biographical notes on Isaac Carroll (1828-1880) are presented. The name Pyrenocollema bryospilum (Nyl.) Coppins is validated. Verrucaria beloniella Nyl. is lectotypified and synonymised with Lecidella stigmatea (Ach.) Hertel & Leuckert. Stigmidium congestum and Lecidea hypopta are added to the floras of Norway and Ireland respectively. Key Words – Herbarium, Lichens, Isaac Carroll. BIOGRAPHICAL NOTES ON ISAAC CARROLL Isaac Carroll, a nineteenth century Irish botanist, was primarily a lichenologist. His contribution to botany has been written up by several researchers in recent decades (Cullinane 1973, Seaward et al. 1982, Parnell 1992, Mitchell 1996). Isaac Carroll was the only son of James Carroll1 and wife Susannah, née Abbott2. Born in Cork City on 28 April 1828, he died in Aghada, Co. Cork on 7 September 1880, aged 52. He was a Quaker and is interred at the Society of Friends' Burial Ground, Summerhill South, Cork (Mitchell, 1995, 1996). He corresponded with botanists from 19 Summer Hill, on the North side of Cork City, in 1858-60, 1862, 1864, 1867-8 and 1870 (Seaward et al., 1982; Mitchell, 1996) and from 79 South Mall in 1858 and 18723. The portrait of Isaac Carroll which now hangs in the herbarium in Glasnevin, was painted by Dublin born artist, Josephine Webb, R.H.A. -
Arthonia Pragensis Spec. Nov. (Ascomycetes, Arthoniales), a New Lichenicolous Fungus from the Czech Republic
C z e c h m y c o l . 47 (2), 1994 Arthonia pragensis spec. nov. (Ascomycetes, Arthoniales), a new lichenicolous fungus from the Czech Republic J a n a H o r á k o v á National Museum, Václavské nám. 68, 115 79 Praha 1, Czech Republic Horáková J. (1994): Arthonia pragensis spec. nov. (Ascomycetes, Arthoniales), a new lichenicolous fungus from the Czech Republic. Czech Mycol. 47: 139-143 Arthonia pragensis Horáková, a new lichenicolous fungus (Ascomycetes, Arthoniales) is de scribed from xerothermic area growing on diabase rocks in Prague in the Czech Republic. This is also the first published report of a lichenicolous fungus from the territory of Prague. Key words: Arthonia, lichenicolous fungi, Czech Republic, taxonomy Horáková J. (1994): Arthonia pragensis spec. nov. (Ascomycetes, Arthoniales), nová lichenikolní houba z České republiky. Czech Mycol. 47: 139-143 Arthonia pragensis Horáková je nová lichenikolní houba (Ascomycetes, Arthoniales) popsaná z xerothermních diabasových skal v Praze v České republice. Jedná se o první publikovaný údaj lichenikolní houby na území Prahy. I ntroduction At present the genus Arthonia (Ascomycetes) includes 44 lichenicolous or para sitic species (incl. Conida and Celidium). This very rich group needs a new taxonomic revision, which should include the nature of the ascus structure. Only the recent research underline amyloidity of ascoapical structures in some lichenized and lichenicolous species of Arthonia (Coppins 1983, 1989; Triebel 1989; Alstrup et Hawksworth 1990) and it points out two different ascus types in this genus (Alstrup et Hawksworth 1990). The species described below, collected once on sunny diabase rocks from xe rothermic area in Prague at the elevation cca 310 m above sea level, belongs to those species of Arthonia having this aspect of amyloidity. -
<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 -
Studies in Lichens and Lichenicolous Fungi: 7
ISSN (print) 0093-4666 © 2011. Mycotaxon, Ltd. ISSN (online) 2154-8889 MYCOTAXON Volume 115, pp. 45–52 January–March 2011 doi: 10.5248/115.45 Studies in lichens and lichenicolous fungi: 7. More notes on taxa from North America James C. Lendemer*1 & Kerry Knudsen2 1Cryptogamic Herbarium, Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY 10458-5126, USA 2The Herbarium, Dept. of Botany & Plant Sciences, University of California, Riverside, CA 92521-0124, USA Correspondence to *: [email protected] & [email protected] Abstract— Acarospora complanata, Fellhaneropsis myrtillicola, and Lecanora stramineoalbida are reported new for North America north of Mexico. Acarospora superfusa is confirmed as occurring in North America. Biatorella rappii is placed in synonymy with Ramonia microspora. Key words— Appalachian Mountains, Magnusson, Sonoran Desert, SE coastal plain. 1. Acarospora complanata H. Magn., Svensk. Bot. Tidskr. 18: 332. 1924. Type: France. Provence-Alpes-Côte D’azur: Var Dist., Massif volcanique de la Courtine, pres Ollisules, 1923, de Crozals (hb. B. de Lesd.[n.v.-presumed destroyed], holotype; UPS! isotype). Acarospora complanata was described from France (Magnusson 1924) and Magnusson recognized it as occurring in Africa and Mexico (Magnusson 1929, 1956). The species forms a brown areolate orbicular thallus with inconspicuous immersed apothecia and an effigurate margin with narrow lobes, a hymenium 80–90 μm high, paraphyses at mid-height mostly 2–2.5 μm in diameter, and a cortex that contains gyrophoric and lecanoric acids (KC+ pink). For a fuller description see Magnusson (1929). Morphologically, the species does not appear related to A. molybdina (Wahlenb.) Trevis, A. macrocyclos Vain., or A. -
A Molecular Phylogeny of the Lichen Genus Lecidella Focusing on Species from Mainland China
RESEARCH ARTICLE A Molecular Phylogeny of the Lichen Genus Lecidella Focusing on Species from Mainland China Xin Zhao1, Lu Lu Zhang1, Zun Tian Zhao1, Wei Cheng Wang1, Steven D. Leavitt2,3, Helge Thorsten Lumbsch2* 1 College of Life Sciences, Shandong Normal University, Jinan, 250014, P. R. China, 2 Science & Education, The Field Museum, Chicago, Illinois, United States of America, 3 Committee on Evolutionary Biology, University of Chicago, Chicago, Illinois, United States of America * [email protected] Abstract The phylogeny of Lecidella species is studied, based on a 7-locus data set using ML and OPEN ACCESS Bayesian analyses. Phylogenetic relationships among 43 individuals representing 11 Leci- della species, mainly from mainland China, were included in the analyses and phenotypical Citation: Zhao X, Zhang LL, Zhao ZT, Wang WC, characters studied and mapped onto the phylogeny. The Lecidella species fall into three Leavitt SD, Lumbsch HT (2015) A Molecular Phylogeny of the Lichen Genus Lecidella Focusing major clades, which are proposed here as three informal groups–Lecidella stigmatea group, on Species from Mainland China. PLoS ONE 10(9): L. elaeochroma group and L. enteroleucella group, each of them strongly supported. Our e0139405. doi:10.1371/journal.pone.0139405 phylogenetic analyses support traditional species delimitation based on morphological and Editor: Nico Cellinese, University of Florida, UNITED chemical traits in most but not all cases. Individuals considered as belonging to the same STATES species based on phenotypic characters were found to be paraphyletic, indicating that cryp- Received: July 20, 2015 tic species might be hidden under these names (e.g. L. carpathica and L. -
Piedmont Lichen Inventory
PIEDMONT LICHEN INVENTORY: BUILDING A LICHEN BIODIVERSITY BASELINE FOR THE PIEDMONT ECOREGION OF NORTH CAROLINA, USA By Gary B. Perlmutter B.S. Zoology, Humboldt State University, Arcata, CA 1991 A Thesis Submitted to the Staff of The North Carolina Botanical Garden University of North Carolina at Chapel Hill Advisor: Dr. Johnny Randall As Partial Fulfilment of the Requirements For the Certificate in Native Plant Studies 15 May 2009 Perlmutter – Piedmont Lichen Inventory Page 2 This Final Project, whose results are reported herein with sections also published in the scientific literature, is dedicated to Daniel G. Perlmutter, who urged that I return to academia. And to Theresa, Nichole and Dakota, for putting up with my passion in lichenology, which brought them from southern California to the Traingle of North Carolina. TABLE OF CONTENTS Introduction……………………………………………………………………………………….4 Chapter I: The North Carolina Lichen Checklist…………………………………………………7 Chapter II: Herbarium Surveys and Initiation of a New Lichen Collection in the University of North Carolina Herbarium (NCU)………………………………………………………..9 Chapter III: Preparatory Field Surveys I: Battle Park and Rock Cliff Farm……………………13 Chapter IV: Preparatory Field Surveys II: State Park Forays…………………………………..17 Chapter V: Lichen Biota of Mason Farm Biological Reserve………………………………….19 Chapter VI: Additional Piedmont Lichen Surveys: Uwharrie Mountains…………………...…22 Chapter VII: A Revised Lichen Inventory of North Carolina Piedmont …..…………………...23 Acknowledgements……………………………………………………………………………..72 Appendices………………………………………………………………………………….…..73 Perlmutter – Piedmont Lichen Inventory Page 4 INTRODUCTION Lichens are composite organisms, consisting of a fungus (the mycobiont) and a photosynthesising alga and/or cyanobacterium (the photobiont), which together make a life form that is distinct from either partner in isolation (Brodo et al. -
Opuscula Philolichenum, 11: 120-XXXX
Opuscula Philolichenum, 13: 102-121. 2014. *pdf effectively published online 15September2014 via (http://sweetgum.nybg.org/philolichenum/) Lichens and lichenicolous fungi of Grasslands National Park (Saskatchewan, Canada) 1 COLIN E. FREEBURY ABSTRACT. – A total of 194 lichens and 23 lichenicolous fungi are reported. New for North America: Rinodina venostana and Tremella christiansenii. New for Canada and Saskatchewan: Acarospora rosulata, Caloplaca decipiens, C. lignicola, C. pratensis, Candelariella aggregata, C. antennaria, Cercidospora lobothalliae, Endocarpon loscosii, Endococcus oreinae, Fulgensia subbracteata, Heteroplacidium zamenhofianum, Lichenoconium lichenicola, Placidium californicum, Polysporina pusilla, Rhizocarpon renneri, Rinodina juniperina, R. lobulata, R. luridata, R. parasitica, R. straussii, Stigmidium squamariae, Verrucaria bernaicensis, V. fusca, V. inficiens, V. othmarii, V. sphaerospora and Xanthoparmelia camtschadalis. New for Saskatchewan alone: Acarospora stapfiana, Arthonia glebosa, A. epiphyscia, A. molendoi, Blennothallia crispa, Caloplaca arenaria, C. chrysophthalma, C. citrina, C. grimmiae, C. microphyllina, Candelariella efflorescens, C. rosulans, Diplotomma venustum, Heteroplacidium compactum, Intralichen christiansenii, Lecanora valesiaca, Lecidea atrobrunnea, Lecidella wulfenii, Lichenodiplis lecanorae, Lichenostigma cosmopolites, Lobothallia praeradiosa, Micarea incrassata, M. misella, Physcia alnophila, P. dimidiata, Physciella chloantha, Polycoccum clauzadei, Polysporina subfuscescens, P. urceolata, -
Large Differences in Carbohydrate Degradation and Transport Potential
bioRxiv preprint doi: https://doi.org/10.1101/2021.08.01.454614; this version posted August 2, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC 4.0 International license. 1 Large differences in carbohydrate degradation and 2 transport potential in the genomes of lichen fungal 3 symbionts 4 5 Philipp Resl1,2, Adina R. Bujold3, Gulnara Tagirdzhanova3, Peter Meidl2, Sandra Freire Rallo4, 6 Mieko Kono5, Samantha Fernández-Brime5, Hörður Guðmundsson7, Ólafur Sigmar 7 Andrésson7, Lucia Muggia8, Helmut Mayrhofer1, John P. McCutcheon9, Mats Wedin5, Silke 8 Werth2, Lisa M. Willis3, Toby Spribille3* 9 10 1University of Graz, Institute of Biology, Universitätsplatz 2, 8010 Graz, Austria 11 2Ludwig-Maximilians-University Munich, Faculty of Biology Department 1, Diversity and 12 Evolution of Plants, Menzingerstraße 67, 80638 Munich, Germany 13 3University of Alberta, Biological Sciences CW405, Edmonton, AB T6G 2R3 Canada 14 4Rey Juan Carlos University, Departamento de Biología y Geología, Física y Química 15 Inorgánica, Móstoles, Spain. 16 5SWedish Museum of Natural History, Botany Department, PO Box 50007, SE10405 17 Stockholm, SWeden 18 7Faculty of Life and Environmental Sciences, University of Iceland, Sturlugata 7, 102 19 Reykjavík, Iceland 20 8University of Trieste, Department of Life Sciences, via L. Giorgieri 10, 34127 Trieste, Italy 1 bioRxiv preprint doi: https://doi.org/10.1101/2021.08.01.454614; this version posted August 2, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity.