Lichen myco- and photobiont diversity and their relationships at the edge of life (McMurdo Dry Valleys, Antarctica)

FEMS Microbiology Ecology

Volumn 82, Issue 2, 2012, Pages 429-448

  Pérez Ortega, Sergio ^a   Ortiz Álvarez, Rüdiger ^a   Allan Green, T G ^b   de los Ríos, Asunción ^a  

^a MUSEO NACIONAL DE CIENCIAS NATURALES   (Spain)

^b UNIVERSIDAD COMPLUTENSE DE MADRID   (Spain)

Author keywords

Ascomycota; Lichen forming fungi; Photobionts; Selectivity; Spatial structure; Symbiosis

Indexed keywords

ALGAL DNA; FUNGAL DNA; RIBOSOMAL SPACER DNA;

BRYOPHYTE; ENVIRONMENTAL CONDITIONS; EXTREME EVENT; FUNGUS; LICHEN; POPULATION DISTRIBUTION; SPATIAL ANALYSIS; SPECIES DIVERSITY; SYMBIOSIS; VASCULAR PLANT;

ANTARCTICA; ARTICLE; BAYES THEOREM; BIODIVERSITY; BIOLOGICAL MODEL; CLASSIFICATION; ECOSYSTEM; FUNGUS; GENETICS; GREEN ALGA; HAPLOTYPE; LICHEN (ORGANISM); MICROBIOLOGY; MOSS; NUCLEOTIDE SEQUENCE; PHYLOGENY; SEQUENCE ALIGNMENT; SPECIES DIFFERENCE; STATISTICAL MODEL;

ANTARCTIC REGIONS; BASE SEQUENCE; BAYES THEOREM; BIODIVERSITY; BRYOPHYTA; CHLOROPHYTA; DNA, ALGAL; DNA, FUNGAL; DNA, RIBOSOMAL SPACER; ECOSYSTEM; FUNGI; HAPLOTYPES; LICHENS; LIKELIHOOD FUNCTIONS; MODELS, GENETIC; PHYLOGENY; SEQUENCE ALIGNMENT; SPECIES SPECIFICITY;

ANTARCTICA; EAST ANTARCTICA; MCMURDO DRY VALLEYS;

ASCOMYCOTA; BRYOPHYTA; BRYOPHYTES; FUNGI; TRACHEOPHYTA;

EID: 84867717167     PISSN: 01686496     EISSN: 15746941     Source Type: Journal    
DOI: 10.1111/j.1574-6941.2012.01422.x     Document Type: Article

References (114)

1. Akaike H (1974) A new look at the statistical model identification. IEEE Trans Auto Control 19: 716-723.
2. Aoki M, Nakano T, Kanda H & Deguchi H (1998) Photobionts isolated from Antarctic lichens. J Mar Biotechnol 6: 39-43.
3. Bačkor M, Peksa O, Škaloud P & Bačkorová M (2010) Photobiont diversity in lichens from metal-rich substrata based on ITS rDNA sequences. Ecotoxicol Environ Saf 73: 603-612.
4. Beck A (1999) Photobiont inventory of a lichen community growing on heavy-metal-rich rock. Lichenologist 31: 501-510.
5. Beck A, Friedl T & Rambold G (1998) Selectivity of photobiont choice in a defined lichen community: inferences from cultural and molecular studies. New Phytol 139: 709-720.
6. Beck A, Kasalicky T & Rambold G (2002) Myco-photobiontal selection in a Mediterranean cryptogam community with Fulgensia fulgida. New Phytol 153: 317-326.
7. Begerow D, Nilsson H, Unterseher M & Maier W (2010) Current state and perspectives of fungal DNA barcoding and rapid identification procedures. Appl Microbiol Biotechnol 87: 99-108.
8. Blaha J, Baloch E & Grube M (2006) High photobiont diversity associated with the euryoecious lichen-forming ascomycete Lecanora rupicola (Lecanoraceae, Ascomycota). Biol J Linn Soc 99: 283-293.
9. Castello M & Nimis PL (1995a) A critical revision of Antarctic lichens described by C. W. Dodge. Studies in Lichenology with Emphasis on Chemotaxonomy, Geography and Phytochemistry (Knoph JG, Schrüfer K & Sipman HJM, eds), pp. 71-92. Festschrift Christian Leuckert, Bibliotheca Lichenologica, J. Cramer, Berlin, Stuttgart.
10. Castello M & Nimis PL (1995b) The lichen vegetation of Terra Nova Bay (Victoria Land, Continental Antarctica). Scripta Lichenologica - Lichenological Papers Dedicated to Antonín Vezda (Farkas EE, Lücking R & Wirth V, eds), pp. 43-55. Bibliotheca Lichenologica, J. Cramer, Berlin, Stuttgart.
11. Castresana J (2000) Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Mol Biol Evol 17: 540-552.
12. Clement M, Posada D & Crandall K (2000) TCS: a computer program to estimate gene genealogies. Mol Ecol 9: 1657-1660.
13. Convey P & Stevens MI (2007) Antarctic biodiversity. Science 317: 1877-1888.
14. Crawford AJ, Lips KR & Bermingham E (2010) Epidemic disease decimates amphibian abundance, species diversity, and evolutionary history in the highlands of central Panama. P Natl Acad Sci USA 107: 13777-13782.
15. Crespo A & Pérez-Ortega S (2009) Cryptic species and species pairs in lichens: a discussion on the relationship between molecular phylogenies and morphological characters. An Jard Bot Madr 66 (suppl 1): 71-81.
16. Cubero OF, Crespo A, Fatehi J & Bridge PD (1999) DNA extraction and PCR amplification method suitable for fresh, herbarium-stored, lichenized, and other fungi. Plant Syst Evol 216: 243-249.
17. De la Torre JR, Goebel BM, Friedmann EI & Pace NR (2003) Microbial diversity of cryptoendolithic communities from the McMurdo Dry Valleys, Antarctica. Appl Envrion Microbiol 69: 3858-3867.
18. De los Ríos A, Wierzchos J, Sancho LG, Green TGA & Ascaso C (2005a) Ecology of endolithic lichens colonizing granite in continental Antarctica. Lichenologist 37: 383-395.
19. De los Ríos A, Sancho LG, Grube M, Wierzchos J & Ascaso C (2005b) Endolithic growth of two Lecidea lichens in granite from continental Antarctica detected by molecular and microscopy techniques. New Phytol 165: 181-190.
20. De Wever A, Leliaert F, Verleyen E, Vanormelingen P, Van der Gucht K, Hodgson DA, Sabbe K & Vyverman W. (2009) Hidden levels of phylodiversity in Antarctic green algae: further evidence for the existence of glacial refugia. Proc Biol Sci 276: 3591-3599.
21. Del Prado R, Cubas P, Lumbsch HT, Divakar PK, Blanco O, Amo De Paz G, Molina MC & Crespo A (2010) Genetic distances within and among species in monophyletic lineages of Parmeliaceae (Ascomycota) as a tool for taxon delimitation. Mol Phylogenet Evol 56: 125-133.
22. Doering M & Piercey-Normore MD (2009) Genetically divergent algae shape an epiphytic lichen community on Jack Pine in Manitoba. Lichenologist 41: 69-80.
23. Drummond AJ & Rambaut A (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evol Biol 7: 214.
24. Drummond AJ, Ho SYW, Phillips MJ & Rambaut A (2006) Relaxed phylogenetics and dating with confidence. PLoS Biol 4: e88.
25. Edgar RC (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Res 32: 1792-1797.
26. Engelen A, Convey P & Ott S (2010) Life history strategy of Lepraria borealis at an Antarctic inland site, Coal Nunatak. Lichenologist 42: 339-346.
27. Excoffier L, Smouse PE & Quattro JM (1992) Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data. Genetics 131: 479-491.
28. Fell JW, Scorzetti G, Connell L & Craig S (2006) Biodiversity of micro-eukaryotes in Antarctic Dry Valley soils with < 5% soil moisture. Soil Biol Biochem 38: 3107-3119.
29. Felsenstein J (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39: 783-791.
30. Fernández-Mendoza F, Domaschke S, García MA, Jordan P, Martín MP & Printzen C (2011) Population structure of mycobionts and photobionts of the widespread lichen Cetraria aculeata. Mol Ecol 20: 1208-1232.
31. Friedl T (1989) Comparative ultrastructure of pyrenoids in Trebouxia (Microthamniales, Chlorophyta). Plant Syst Evol 164: 145-159.
32. Friedmann EI (1982) Endolithic microorganisms in the Antarctic cold desert. Science 215: 1045-1053.
33. Friedmann EI, Garty J & Kappen L (1980) Fertile stages of cryptoendolithic lichens in the dry valleys of Southern Victoria Land. Antarct J Unit States 15: 166-167.
34. Galun M & Bubrick P (1984) Physiological interactions between partners of the lichen symbiosis. Encyclopedia of Plant Physiology, New Series, Volume 17. Cellular Interactions (Linskens HF & Heslop-Harrison J, eds), pp. 362-401. Springer-Verlag, Berlin, Heidelberg.
35. Gardes M & Bruns TD (1993) ITS primers with enhanced specificity for basidiomycetes - application to the identification of mycorrhizae and rust. Mol Ecol 2: 113-118.
36. Green TGA (2009) Lichens in arctic, Antarctic and alpine ecosystems. Ökologische Rolle der Flechten. Rundgespräche der Kommission für Ökologie 36. Bayerische Akademie der Wissenschaften (Herausgeber), Verlag Dr. Freiderich Pfeil, München, 190 pages, pp. 45-65.
37. Green TGA, Schroeter B & Sancho LG (2007) Plant life in Antarctica. Functional Plant Ecology, 2nd edn (Pugnaire FL & Valladares F, eds), pp. 389-433. CRC Press, Boca Raton, FL.
38. oS, Queen Maud Mountains, suggests preglacial survival of species in the Ross Sea region Antarctica. Polar Biol 34: 1211-1220.
39. Grundt HH, Kjølner S, Borgen L, Rieseberg LH & Brochman C (2006) High biological species diversity in the arctic flora. PNAS 103: 972-975.
40. Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W & Gascueln O (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Syst Biol 59: 307-321.
41. Guzow-Krzemińska B (2006) Photobiont flexibility in the lichen Protoparmeliopsis muralis as revealed by ITS rDNA analysis. Lichenologist 38: 469-476.
42. Hale ME (1987) Epilithic lichens in the Beacon sandstone formation, Victoria Land, Antarctica. Lichenologist 19: 269-287.
43. Hall TA (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis. http://www.mbio.ncsu.edu/BioEdit/bioedit.html.
44. Hart M & Sunday J (2007) Things fall apart: biological species form unconnected parsimony networks. Biol Lett 3: 509-512.
45. Hauck M, Helms G & Friedl T (2007) Photobiont selectivity in the epiphytic lichens Hypogymnia physodes and Lecanora conizaeoides. Lichenologist 39: 195-204.
46. Hawksworth DL (2001) The magnitude of fungal diversity: the 1.5 million species estimate revisited. Mycol Res 105: 1422-1432.
47. Hawksworth DL & Honegger R (1994) The lichen thallus: a symbiotic phenotype of nutritionally specialized fungi and its response to gall producers. Plant Galls. Organisms, Interactions, Populations (Williams MAJ, ed), pp. 77-98. The Systematics Association Special Volume, Clarendon Press, Oxford.
48. Helms G, Friedl T, Rambold G & Mayrhofer H (2001) Identification of photobionts from the lichen family Physciaceae using algal-specific ITS rDNA sequencing. Lichenologist 33: 73-86.
49. Hertel H (1987) Progress and problems in taxonomy of Antarctic saxicolous lecideoid lichens. Progress and Problems in Lichenology in the Eighties (Peveling E, ed), pp. 219-242. Bibliotheca Lichenologica, J. Cramer, Berlin, Stuttgart.
50. Hertel H (1988) Problems in monographing Antarctic crustose lichens. Polarforschung 58: 65-76.
51. Kappen L & Valladares F (2007) Opportunistic growth and desiccation tolerance: the ecological success of poikilohydrous autotrophs. Handbook of Functional Plant Ecology (Pugnaire FI & Valladares F, eds), pp. 9-80. Marcel Dekker, Inc., New York, Basel.
52. Kappen L, Friedmann EI & Garty J (1981) Ecophysiology of lichens in the dry valleys of southern Victoria Land, Antarctica. I. Microclimate of the cryptoendolithic lichen habitat. Flora 171: 216-235.
53. 2 exchange on Umbilicaria aprina under extreme light stress in the cold. Oecologia 113: 325-331.
54. Kelly LJ, Hollingsworth PM, Coppins BJ, Ellis CJ, Harrold P, Tosh J & Yahr R (2011) DNA barcoding of lichenized fungi demonstrates high identification success in a floristic context. New Phytol, 191: 288-300.
55. Kroken S & Taylor JW (2000) Phylogenetic species, reproductive mode, and specificity of the green alga Trebouxia forming lichens with the fungal genus Letharia. Bryologist 103: 645-660.
56. Kroken S & Taylor JW (2001) A gene genealogical approach to recognize phylogentic species boundaries in the lichenized fungus Letharia. Mycologia 93: 38-53.
57. Leavitt SD, Johnson LA & St. Clair LL (2011) Species delimitation and evolution in morphologically and chemically diverse communities of the lichen-forming genus Xanthoparmelia (Parmeliaceae, Ascomycota) in western North America. Am J Bot 98: 175-188.
58. Lefébure T, Douady CJ, Gouy M, Trontelj P, Briolay J & Gibert J (2006) Phylogeography of a subterranean amphipod reveals cryptic diversity and dynamic evolution in extreme environments. Mol Ecol 15: 1797-1806.
59. Librado P & Rozas J (2009) DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinformatics 25: 1451-1452.
60. Lohse K (2009) Can mtDNA barcodes be used to delimit species? A response to Pons et al. (2006). Syst Biol 58: 439-442.
61. Lumbsch HT & Leavitt S (2011) Goodbye morphology? A paradigm shift in the delimitation of species in lichenized fungi. Fung Divers 50: 59-72.
62. Matesanz S, Gimeno TE, de la Cruz M, Escudero A & Valladares F (2011) Competition may explain the fine-scale spatial patterns and genetic structure of two co-occurring plant congeners. J Ecol, 99: 838-848.
63. Monaghan MT, Wild R, Elliot M et al. (2009) Accelerated species inventory on Madagascar using coalescent-based models of species delineation. Syst Biol 58: 298-311.
64. Muggia L, Grube M & Tretiach M (2008) Genetic diversity and photobiont associations in selected taxa of the Tephromela atra group (Lecanorales, lichenised Ascomycota). Mycol Prog 7: 147-160.
65. Muggia L, Zellnig G, Rabensteiner J & Grube M (2010) Morphological and phylogenetic study of algal partners associated with the lichen-forming fungus Tephromela atra from the Mediterranean region. Symbiosis 51: 149-160.
66. Nakayama T, Watanabe S, Mitsui K, Uchida H & Inouye I (1996) The phylogenetic relationship between the Chlamydomonadales and Chlorococcales inferred from 18S rDNA sequence data. Phycol Res 44: 47-55.
67. Nei M (1987) Molecular Evolutionary Genetics. Columbia University Press, New York, NY.
68. Nelsen MP & Gargas A (2008) Dissociation and horizontal transmission of codispersing lichen symbionts in the genus Lepraria (Lecanorales: Stereocaulaceae). New Phytol 177: 264-275.
69. Nevo E (2001) Evolution of genome-phenome diversity under enviromental stress. PNAS 98: 6233-6240.
70. Ohmura Y, Kawachi M, Kasai F & Watanabe M (2006) Genetic combinations of symbionts in a vegetatively reproducing lichen, Parmotrema tinctorum, based on ITS rDNA sequences. Bryologist 109: 43-59.
71. Oksanen J, Blanchet FG, Kindt R, Legendre P, O'Hara RB, Simpson GL, Solymos P, Stevens MHH & Wagner H (2011) Vegan: Community Ecology Package. R package version 1.17-7. http://CRAN.R-project.org/package=vegan.
72. Orange A, James PW & White FJ (2001) Microchemical Methods for the Identification of Lichens. British Lichen Society, London, pp. 101.
73. Otálora MAG, Martínez I, O′Brien H, Molina MC, Aragón G & Lutzoni F (2010) Multiple origins of high reciprocal symbiotic specificity at an intercontinental spatial scale among gelatinous lichens (Collemataceae, Lecanoromycetes). Mol Phylogenet Evol 56: 1089-1095.
74. Øvstedal DO & Lewis-Smith RI (2001) Lichens of Antarctica and South Georgia: A Guide to Their Identification and Ecology. Studies in Polar Research, Cambridge University Press, Cambridge, UK, pp. 411.
75. Papadopoulou A, Monaghan MT, Barraclough TG & Vogler AP (2009) Sampling error does not invalidate the Yule-coalescent model for species delimitation. A response to Lohse (2009). Syst Biol 58: 442-444.
76. Paradis E (2010) Pegas: an R package for population genetics with an integrated-modular approach. Bioinformatics 26: 419-420.
77. Paradis E, Claude J & Strimmer K (2004) APE: analyses of phylogenetics and evolution in R language. Bioinformatics 20: 289-290.
78. Peat HJ, Clarke A & Convey P (2007) Diversity and biogeography of the Antarctic flora. J Biogeogr 34: 132-146.
79. Peksa O & Škaloud P (2011) Do photobionts influence the ecology of lichens? A case study of environmental preferences in symbiotic green alga Asterochloris (Trebouxiophyceae). Mol Ecol 20: 3936-3948.
80. Pérez-Ortega S, de los Ríos A, Crespo A & Sancho LG (2010) Symbiotic lifestyle and phylogenetic relationships of the bionts of Mastodia tessellata (Ascomycota, incertae sedis). Am J Bot 97: 738-752.
81. Piercey-Normore MD (2009) Vegetatively reproducing fungi in three genera of the Parmeliaceae show divergent algal partners. Bryologist 112: 773-785.
82. Pinzón-Navarro S, Barrios H, Múrria C, Lyal CHC & Vogler AP (2010) DNA-based taxonomy of larval stages reveals huge unknown species diversity in neotropical seed weevils (genus Conotrachelus): relevance to evolutionary ecology. Mol Phylogenet Evol 56: 281-293.
83. Pointing SB, Chan Y, Lacap DC, Lau MCY, Jurgens JA & Farrell RL (2009) Highly specialized microbial diversity in hyper-arid polar desert. P Natl Acad Sci USA 106: 19964-19969.
84. Pons J, Barraclough TG, Gómez-Zurita J, Cardoso A, Duran DP, Hazell S, Kamoun S, Sumlin WD & Vogler AP (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Syst Biol 55: 595-609.
85. Posada D (2008) jModelTest: phylogenetic model averaging. Mol Biol Evol 25: 1253-1256.
86. Powell JR (2012) Accounting for uncertainty in species delineation during the analysis of environmental DNA sequence data. Methods Ecol Evol 3: 1-11.
87. Prentice ML, Bockheim JG, Wilson SC, Burckle LH, Hodell DA, Schlüchter C & Kellog DE (1993) Late Neogene Antarctic glacial history: evidence from central Wright Valley. The Antarctic Paleoenvironment: A Perspective on Global Change, Vol. 60 (Knnet J & Warnke D, eds), pp. 207-250. American Geophysical Union, Washington, DC.
88. R Development Core Team (2010) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
89. Rambaut A & Drummond AJ (2007) Tracer v1.4. Available at http://beast.bio.ed.ac.uk/Tracer.
90. Romeike J, Friedl T, Helms G & Ott S (2002) Genetic diversity of algal and fungal partners in four species of Umbilicaria (lichenized ascomycetes) along a transect of the Antarctic Peninsula. Mol Biol Evol 19: 1209-1217.
91. Ronquist F & Huelsenbeck JP (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572-1574.
92. Ruprecht U, Lumbsch HT, Brunauer G, Green TGA & Türk R (2010) Diversity of Lecidea (Lecideaceae, Ascomycota) species revealed by molecular data and morphological characters. Antarct Sci 22: 727-741.
93. Sabbe K, Hodgson DA, Verleyen E, Taton A, Wilmorte A, Vanhoutte K & Vyverman W (2004) Salinity, depth and the structure and composition of microbial mats in continental Antarctic lakes. Freshw Biol 49: 296-319.
94. Sancho LG, Green TGA & Pintado A (2007) Slowest to fastest: extreme range in lichen growth rates supports their use as an indicator of climate change in Antarctica. Flora 202: 667-673.
95. Schloss PD, Westcott SL, Ryabin T et al. (2009) Introducing mothur: open-source, platform-independent, community-supported software for describing and comparing microbial communities. Appl Environ Microbiol 75: 7537-7541.
96. Schmidt HA, Strimmer K, Vingron M & von Haeseler A (2002) TREE-PUZZLE: maximum likelihood phylogenetic analysis using quartets and parallel computing. Bioinformatics 18: 502-504.
97. Seifert KA (2009) Progress towards DNA barcoding of fungi. Mol Ecol Resour, 9: 83-89.
98. Seppelt RD, Green TGA & Schroeter B (1995) Lichens and mosses from the Kar Plateau, Southern Victoria Land, Antarctica. NZ J Bot 33: 203-220.
99. Seppelt RD, Türk R, Green TGA, Moser G, Pannewitz S, Sancho LG & Schroeter B (2010) Lichen and moss communities of Botany Bay, Granite Harbour, Ross Sea, Antarctica. Antarct Sci 22: 691-702.
100. Sipman HJM & Aptroot A (2001) Where are the missing lichens?. Mycol Res 105: 1433-1439.
101. Smith RIL (1988) Classification and ordination of cryptogamic communities in Wilkes Land, continental Antarctica. Vegetatio 76: 155-166.
102. Taton A, Grubisic S, Brambilla E, De Wit R & Wilmorte A (2006) Cyanobacterial diversity in natural and artificial microbial mats of Lake Fryxell (McMurdo Dry Valleys, Antarctica): a morphological and molecular approach. Appl Environ Microbiol 69: 5157-5169.
103. Tavaré S (1986) Some probabilistic and statistical problems in the analysis of DNA sequences. Lect Math Life Sci 17: 57-86.
104. Templeton AR, Crandall KA & Sing CF (1992) A cladistic analysis of phenotypic associations with haplotypes inferred from restriction endonuclease mapping and DNA sequence data. III. Cladogram estimation. Genetics 132: 619-633.
105. Tschermak-Woess E (1978) Myrmecia reticulata as a phycobiont and free-living - free-living Trebouxia - the problem of Stenocybe septata. Lichenologist 10: 69-79.
106. Vondrák J, Říha P, Arup U & Søchting U (2009) The taxonomy of the Caloplaca citrina group (Teloschistaceae) in the Black Sea region; with contributions to the cryptic species concept in lichenology. Lichenologist 41: 571-604.
107. Vrijenhoek RC, Schutz SJ, Gustafson RG & Lutz RA (1994) Cryptic species of deep-sea clams (Mollusca, Bivalvia, Vesicomyidae) in hydrothermal vent and cold-seep environments. Deep-Sea Res II 41: 1171-1189.
108. Werth S & Sork VL (2010) Identity and genetic structure of photobiont of the epiphytic lichen Ramalina menziesii on three oak species in Southern California. Am J Bot 97: 821-830.
109. White TJ, Bruns T, Lee S & Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. PCR Protocols: A Guide to Methods and Applications (Innis MA, Gelfand DH, Sninsky JJ & White TJ, eds), pp. 315-322. Academic Press Inc., San Diego.
110. Wirtz N, Lumbsch HT, Green TGA, Türk R, Pintado A, Sancho LG & Schoeter B (2003) Lichen fungi have low cyanobiont selectivity in maritime Antarctica. New Phytol 160: 177-183.
111. Wirtz N, Printzen C & Lumbsch HT (2008) The delimitation of Antarctic and bipolar species of neuropogonoid Usnea (Ascomycota, Lecanorales): a cohesion approach of species recognition for the Usnea perpusilla complex. Mycol Res 112: 472-484.
112. Wornik S & Grube M (2010) Joint dispersal does not imply maintenance of partnerships in lichen symbioses. Microb Ecol 59: 150-157.
113. Yahr R, Vilgalys R & DePriest PT (2004) Strong fungal specificity and selectivity for algal symbionts in Florida scrub Cladonia lichens. Mol Ecol 13: 3367-3378.
114. Yahr R, Vilgalys R & DePriest PT (2006) Geographic variation in algal partners of Cladonia subtenuis (Cladoniaceae) highlights the dynamic nature of a lichen symbiosis. New Phytol 171: 847-860.