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NEW SYSTEMATICS AND REVISED GENERIC CLASSIFICATION OF PARMELIOID (, ) BASED ON MULTIGENE PHYLOGENETIC ANALYSIS

PARSYS 08’ PROJECT Led by SYSTEMOL, UCM

List of Parmelioid genera

1. Bulborrhizina Kurok. 2. Bulbothrix Hale 3. Elix & Hale 4. Cetrariastrum Sipman 5. Cetrelia W.L. Culb. & C.F. Culb 6. Everniastrum Hale ex Sipman 7. Flavoparmelia Hale 8. Flavopunctelia Hale 9. Hypotrachyna (Vain.) Hale 10. Karoowia Hale 11. Essl. 12. O. Blanco, A. Crespo, Divakar, Essl., D. Hawksw. & Lumbsch 13. Melanhalea O. Blanco, A. Crespo, Divakar, Essl., D. Hawksw. & Lumbsch 14. (Asahina) Elix & Hale 15. Omphalodiella Henssen 16. Omphalodium Meyen & Flot. 17. Omphalora T.H. Nash & Hafellner 18. Parmelaria Awasthi 19. Ach. 20. Hale 21. Elix & Hale 22. Parmelinopsis Elix & Hale 23. Parmeliopsis (Nyl.) Nyl. 24. A. Massal. 25. Parmotremopsis Elix & Hale 26. Placoparmelia Henssen 27. Petr. 28. Pseudoparmelia Lynge 29. Krog 30. Relicina (Hale & Kurok.) Hale 31. Relicinopsis Elix & Verdon 32. Xanthoparmelia (Vain.) Hale

Genera numbers 1, 25, 26 and 28 are not included in our study. Genera numbers 10, 15, 18 and 31 are considered synonym from our study. Genera numbers 16, 17 are considered as non parmelioid from phylogenetic studies

Addenda: The taxa mentioned below are representative of the new genera in press.

33. : Parmelina labrosa, Parmelina conlabrosa, Parmelina endoleuca 34. Emodomelanelia: Parmelia massonii

1 35. Nipponoparmelia: Parmelia laevior, Parmelia pseudolaevior and Parmelia isidioclada 36. Notoparmelia: Parmelia crambiocarpa, Parmelia cunninghamii, Parmelia subtestacea, Parmelia signifera, Parmelia tenuirima 37. Pseudomelanelia: Melanelia disjuncta, Melanelia sorediata, Melanelia tominii. 38. Remototrachyna: Hypotrachyna adducta, Hypotrachyna ciliata, Hypotrachyna costaricensis, Hypotrachyna crenata, Hypotrachyna infirma, Hypotrachyna koyaensis, Hypotrachyna scytophylla

Total number of genera in Parmelioid from this study is 34.

GENERA AND TYPE SPECIES

Genera Type species

1. Bulborrhizina Kurok. Bulborrhizina africana Kurok. 2. Bulbothrix Hale Bulbothrix bicornuta (Lynge) Hale 3. Canoparmelia Elix & Hale Canoparmelia texana (Tayor) Elix & Hale 4. Cetrariastrum Sipman Cetrariastrum ecuadoriense (Sant.) Sipman 5. Cetrelia Culb. & Culb Cetrelia cetrarioides (Del. ex Duby) Culb. & Culb. 6. Everniastrum Hale ex Sipman Everniastrum cirrahatum (Fr.) Hale ex Sipman 7. Flavoparmelia Hale Flavoparmelia caperata (L.) Hale 8. Flavopunctelia Hale Flavopunctelia flaventior (Stirt.) Hale 9. Hypotrachyna (Vain.) Hale Hypotrachyna brasiliana (Nyl.) Hale 10. Karoowia Hale Karoowia adhaerense (Nyl.) Hale 11. Melanelia Essl. Melanelia stygia (L.) Essl. 12. Melanelixia O. Blanco et al. Melanelixia glabra (Schaer) O. Blanco et al. 13. O. Blanco et al. Melanohalea exasperata (De Not.) O. Blanco et al. 14. Myelochroa (Asah.) Elix & Hale Myelochroa aurulenta (Tuck.) Elix & Hale 15. Omphalodiella Henssen Omphalodiella patagonica Henssen 16. Omphalodium Meyen & Flot. Omphalodium pisacomense Meyer & Flotow 17. Omphalora Nash & Hafellner Omphalora arizonica (Tuck. ex Willey) 18. Parmelaria Awasthi Parmelaria thomsonii (Stirt.) Awasthi 19. Parmelia Ach. Parmelia saxatilis (L.) Ach. 20. Parmelina Hale (Hoffm.) Hale 21. Parmelinella Elix & Hale Parmelinella wallichiana (Taylor) Elix & Hale 22. Parmelinopsis Elix & Hale Parmelinopsis horrescens (Taylor) Elix & Hale 23. Parmeliopsis (Nyl.) Nyl. Parmeliopsis ambigua (wulf. in Jacq) Nyl. 24. Parmotrema A. Massal. Parmotrema perforatum (Jacq.) Mass. 25. Parmotremopsis Elix & Hale Parmotremopsis antillensis (Nyl.) Elix & Hale

2 26. Placoparmelia Henssen Placoparmelia patagonica Hensen 27. Pleurosticta Petr. (Neck.) Elix & Lumbsch 28. Pseudoparmelia Lynge Pseudoparmelia cyphellata Lynge 29. Punctelia Krog Punctelia borreri (Sm.) Krog 30. Relicina (Hale & Kurok.) Hale Relicina relicinula (Mull Arg.) Hale 31. Relicinopsis Elix & Verdon Relicinopsis intertexta (Mont. & Bosch) Elix & Verdon 32. Xanthoparmelia (Vain.) Hale Xanthoparmelia conspersa (Ach.) Hale

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

1. Pleurosticta (2) 2 sp. Northern hemisphere (Eurasia and North Africa). Temperate. Antioceanic. Brown cortex HNO3 + violet. Medullary metabolites: β-orcinol depsidones Broad lobes. Adnate thallus. Pored epicortex. Lacking pseudocyphellae. Simple rhizines Isolichenan Large (12-16 x 6-11 µm).

4 SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

2 Melanelia disjuncta group Pseudomelanelia (3) (at least Melanelia disjuncta, Melanelia sorediata, Melanelia tominii) Northern hemisphere (Eurasia and North Africa). Temperate. Antioceanic. Brown cortex HNO3 - Medullary metabolites: Orcinol depsides (perlatolic acid, stenosporic acid, girophoric acid) Lobes sublinear, plane to convex, inflated. Adpresse thallus Not pored epicortex Pseudocyphellae flat efigurate. Simple rhizines Ascospores (8-12x4-7 µm)

5 SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

3. Emodomelamelia massonii (1) (at least Parmelia massoniii) Northern hemisphere (Asia, Himalaya). Temperate. Antioceanic. Brown cortex HNO3 + pale to dark blue green Medulla: several strains: negative/ or K-, C-, Pd+ red-orange Medullary metabolites: β-Orcinol depsidones, aliphatic acids. Lobes moderate to narrow, sublinear, sometimes developing secondary lobes Thallus adnate to pulvinate or panniform. Not pored epicortex Pseudocyphellae (fenestrations) efigurate on lobes surface. Simple rhizines simple, rarely furcated. Ascospores 13-16 x 6-9 µm

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

4. Melanohalea (20) Northern hemisphere (Eurasia, North Africa and North America). Temperate. Antioceanic. Brown cortex HNO3- Medullary metabolites β- Orcinol Depsidones or none Lobes subiregular, plain to concave. Adnate to adpress thallus Not pored epicortex Pseudocyphellae on tuberculae. Simple rhizines Ascospores 6-20x4-12 µm

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

5. Melanelixia (11) North and Southern hemispheres; Southern species are monophyletic and nested in terminal . Temperate. Antioceanic. Brown cortex HNO3 - Medullary metabolites: Orcinol Depsides Lobes subiregular, plain to concave. Adnate thallus Pored or fenestrate epicortex with efigurate maculae. Pseudocyphellae Simple rhizines Ascospores 9-15x5-11 µm

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

6. Cetrelia (15) Northern hemisphere (overall in south East Asia and North America) but also rarely in Southern hemisphere. Temperate and tropical. Oceanic. Gray cortex with Atranorin Medullary orcinol depsides and orcinol depsidones Lobes broad and both sides exposed Not adnate thallus. Not pored epicortex Psudocyphellae punctiform Simple rhizines Apothecia commonly perforate Isolichenan Ascospores 11-22 x 6-12 µm

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7. Xanthoparmelia (700) Southern hemisphere but also represented in the Northern one. Temperate and xero tropical regions. Oceanic (mediterranean), subarid to arid. Foliaceous (laminar or umbilicate), crustaceous or vagrant. Laking cilia. Cortex green-yellowish or white or brown HNO3 + (with or without usnic acid and with or without melanoid pygment). Medullary orcinol depsides and orcinol depsidones; β-orcinol depsides, β-orcinol depsidones, aliphatic acids, antraquinones, amino acid derivatives. When foliaceous thalli are adnate Pored epicortex Pseudocyphellae lacking but some strongly maculate (fenestrations) Simple rhizines Apothecia common Xantholichenan Ascospores vacuolar aparat arachiform

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8. Canoparmelia s. str.(texana group: C. texana, C. caroliniana, C. concrescens, C. nairobiensis) (4) Northern and Southern hemispheres. Pantemperate but Pantropical or subtropical at higher elevation. Oceanic. Foliaceous. Cortex gray to white. Medullary metabolites orcinol depsides and β-orcinol depsides. Upper surface not reticulate but weak maculae (slightly fenestrate) Lobes sublinear. Laking cilia. Thallus adnate Pored epicortex Pseudocyphellae lacking but effigurate maculae (fenestrations) present Simple rhizines Isolichenan Ascospores 7-15x6-8µm

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9. Canoparmelia crozalsiana group. Includes: C.crozalsiana, C. carneopruinata, C. schelpei, C. inhamiensis) (4) Northern and Southern hemispheres. Pantemperate Oceanic. Foliaceous. Cortex gray to white. Medullary metabolites β-orcinol depsides and depsidones,. Upper surface strongly reticulate with maculae (fenestrations) delimiting concave areolae. Lobes subiregular. Laking cilia. Thallus adnate Pored epicortex Simple rhizines Ascospores 8-13x4-8 µm

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

10. Austroparmelina (7) (includes all the species of Parmelina from Australia) Southern hemisphere. Temperate. Moderately oceanic. Foliaceous Cortex gray to white. Medullary orcinol and β-orcinol depsides, aliphatic acids, antraquinones, amino acid derivatives. Lobes sublinear ciliate especially in lobe axils. Thallus adnate Pored or slightly fenestrate epicortex. Pseudocyphellae lacking, more or less maculatae. Simple rhizines Isolichenan Apothecia: lacking dark (black) stain below anphithecium. Ascospores more globose than in Parmelina.

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11. Parmotrema (250) (incl. Rimelia, Rimeliella, Parmotremopsis, some Canoparmelia, Parmelaria) Southern and Northern hemispheres. Temperate. Oceanic (mainly in coastal regions). Foliaceous usually with both surfaces exposed. Cortex gray to white. Medullary orcinol depsides and β-orcinol depsidones, aliphatic acids, antraquinones, amino acid derivatives. Lobes broad, rotund, most commonly ciliate. Thallus not adnate Pored or slightly fenestrate epicortex (efigurate micromaculae in some species). Pseudocyphellae lacking. Simple rhizines X-lichenan Apothecia: perforate or not. Ascospores large (20-35 x 12-18), subglobose and thick- walled.

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

12. Flavoparmelia (40) Southern and Northern hemispheres. Temperate. Moderately oceanic. Foliaceous. Cortex yellow. Medullary orcinol depsides and β-orcinol depsides, and β-orcinol depsidones, aliphatic acids, antraquinones, secalonic acids, amino acid derivatives. Lobes broad, sub- rotund, lacking cilia. Thallus adnate Pored epicortex. Pseudocyphellae lacking. Simple rhizines isolichenan Apothecia: not perforate. Ascospores (10-14 x 7-10).

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13. Flavopunctelia (5) Southern and Northern hemispheres in all continents except Australia. Tropical to temperate. Moderately oceanic. Foliaceous. Cortex yellow. Medullary orcinol depside (Lecanoric acid), aliphatic acids. Lobes broad, sub- rotund, lacking cilia. Thallus adnate Nonpored epicortex. Pseudocyphellae punctiform. Simple rhizines Isolichenan ? Apothecia: not perforate. Ascospores (11-16 x 5-10).

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14. Nessolechia Lichenicolous fungi.

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15. Punctelia (30) Southern and Northern hemispheres more abundant in South America and subtropical Africa. Tropical to temperate. Moderately oceanic. Foliaceous. Cortex gray (atranorin). Medullary orcinol depside (Lecanoric acid), aliphatic acids. Lobes broad, sub- rotund, lacking cilia. Thallus adnate Nonpored epicortex. Pseudocyphellae punctiform. Simple rhizines to fasciculate. Isolichenan Apothecia: not perforate. Ascospores (10-27 x 8-18).

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

16. Bulbotrix Hale pr. s. str. (at least 4 species; from morphological features it could be 15 species. Important problem: typus circumscription is not confirmed) Tropical. Pantropical. Only the 3 species with bicornutae spores are restricted to neotropic (Brasil). From semiarid tropical regions at 2300m to lowlands rainforests. Cortex gray to white Medullary chemistry: Orcinol depsides and orcinol depsidones. Pored epicortex. Pseudocyphellae lacking. Bulbate cilia (this character is not exclusive because is also present in Relicina) Rhizines dichotomic. Ascospores small (5-10x4-6 μm)

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17. Bulbotrix Hale (2)= the includes Parmelinella wallichiana typus of the . The name of this genus will be Parmelinella. (number or species would be about 6) Tropical. Paleotropical (South East Asia, India, East Africa and South Africa). Subtropical to temperate 750m to 3000m. Cortex gray to white Medullary chemistry: β-Orcinol depsidones. Pored epicortex. Pseudocyphellae lacking. Non-bulbate (Parmelinella wallichiana) or bulbate cilia (this character is not exclusive because is also present in Relicina) Rhizines simples. Ascospores medium to large (10-20x6-10 μm)

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

18. Myelochroa (28) Northern hemisphere (East Asia specially Japan). Temperate. Moderately oceanic. Foliaceous Cortex gray to white. Medulla yellow-orange. Medullary metabolites β-orcinol depsidones, triterpenes, secalonic acids. Lobes sublinear ciliate, cilia simple. Thallus adnate Pored epicortex. Pseudocyphellae lacking, emaculatae. Simple to squarroselly branched rhizines. Isolichenan Apothecia: lacking dark (black) stain below anphithecium. Ascospores more globose than in Parmelina.

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

19. Parmelina (8) Northern hemisphere (specially Mediterranean regions of North America and Eurasia and North Africa). Temperate. Moderately oceanic. Foliaceous Cortex gray to white. Medulla white. Medullary metabolites Orcinol depsides, aliphatic acids. Lobes subrotund ciliate, cilia simple mainly in the lobe axils. Thallus adnate Pored epicortex. Pseudocyphellae lacking, emaculate or maculatae. Simple rhizines. Isolichenan Apothecia: with dark (black) stain below the anphithecium. Ascospores more oval than in Austroparmelina.

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20. Remototrachyna (12) Northern hemisphere (specially temperate high altitudes from 2000-3500m in Hymalaya and surrounding regions). One species in Costa Rica. Foliaceous Cortex gray to white. Medulla white. Medullary metabolites Orcinol depsides, β-Orcinol depsidones and aliphatic acids. Lobes subrotund not ciliate. Thallus variate but frecuently adnate Pored epicortex. Pseudocyphellae lacking, emaculate. Dichotomic rhizines. Isolichenan Apothecia: three layered hypothecium. Ascospores: larger and more ellipsoid comparing with Hypotrachyna s. str.

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

21. Parmeliopsis (6) Northern hemisphere. Temperate and boreal. Specially in conifers forests. Foliaceous Cortex gray to yellow greenish. Medulla white. Medullary metabolites Orcinol depsides. Lobes linear, not overlapping, not ciliate. Thallus adnate Pored epicortex. Pseudocyphellae lacking, emaculate. Simple rhizines. Isolichenan Conidiophores Psora type.

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

22. Everniastrum (40) 27 sp. Pantropical but especially in Neotropic (Central and South America) in high mountains. Foliaceous or sub-fruticulous. Cortex gray. Medulla white. Medullary metabolites Orcinol depsides and depsidones, β-Orcinol depsidones, benzyl esters and aliphatic acids. Lobes linear elongate, canaliculated subdichotomically divides, ciliate. Cilia simple. Pored epicortex. Pseudocyphellae lacking, emaculate. Branched rhizines, some dichotomus, some furcatae or bundles of tufted. Isolichenan Ascospores: large (10-20x5-10µm.

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

23. Cetrariastrum (incl. Everniastrum lipidiferum from Peru) (4) 3 sp. Neotropic (South America). in high mountains. Sub-fruticulous. Cortex gray. Medulla white. Medullary metabolites: aliphatic acids. Lobes linear elongate, canaliculated irregularly branched, ciliate. Cilia simple. Pored epicortex. Pseudocyphellae lacking, emaculate. Rhizines simple or dichotomus. Isolichenan Ascospores: Smaller than in Everniastrum (9-12x5-6µm).

26 SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

24. Hypotrachyna s. str.(typus species H. brasiliana). (9 species were analysed) Including following Hypotrachyna: H. osseoalba, H. immaculata, H. booralensis, H. pseudosinuosa, H. polydactyla, H. livida, H. cf. leiophylla Pantropical (subtropical to temperate regions in low altitudes but in tropics up to 2800m). Foliaceous Cortex gray to white. Medulla white. Medullary metabolites Orcinol depsidones, β-Orcinol depsidones. Lobes sublinear truncate not ciliate. Thallus variate but frecuently adnate Pored epicortex. Pseudocyphellae lacking. Dichotomic rhizines. Isolichenan Apothecia: two layered hypothecium. Ascospores: medium size (7-12x4-8 µm).

27 SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

25. Parmelinopsis (typus species P. horrescens). Including following Hypotrachyna species: H.revoluta, H. neodisecta, H. execta, H.britannica) (31) Pantemperate, oceanic. Foliaceous Cortex gray to white. Medulla white. Medullary metabolites Orcinol depsides, β-Orcinol depsidones. Lobes sublinear, truncate, ciliate. Thallus variate but frecuently adnate Pored epicortex. Pseudocyphellae lacking. Rhizine simple to sparsely furcated or dicotomic. Isolichenan Apothecia: two layered hypothecium. Ascospores: medium size (7-20x6-14µm).

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SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

26. Hypotrachyna gr. laevigata (typus species ?). Including following Hypotrachyna species: H.laevigata, H. taylorensis, H. endochlora, H.rockii, H. physcioides, H. imbricatula, H. degelii, H. carraccensis, H. reducens, H. sinuosa, H. fissicarpa). (15) Pantemperate, oceanic. Foliaceous Cortex gray to white and also yellowish green. Medulla white or yellow. Medullary metabolites Orcinol depsides, β-Orcinol depsides, Orcinol depsidones, β- Orcinol depsidones. Lobes sublinear, to linear, truncate. Thallus variate but frecuently adnate Pored epicortex. Pseudocyphellae lacking. Rhizine dicotomic. Isolichenan Apothecia: two layered hypothecium. Ascospores: medium size (8-22x5-14µm).

29 SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

27. Nipponoparmelia (typus species N. laevior). Including following Parmelia subgenus Nipponoparmelia Kurokava species:Parmelia laevior, P. pseudolaevior, P. isidioclada, P. sochintiana sp. Nova (4) Asian species (specially Japanese) Temperate moderately oceanic. Foliaceous Cortex gray to white. Medulla white. Medullary metabolites β-Orcinol depsidones, Orcinol depsides. Lobes subiregular. Thallus variate but frecuently adnate Non-pored epicortex. Pseudocyphellae only at margins of terminal lobes discontinuous. Rhizine simple to furcated. Ascospores: medium size (12-17x6-10µm).

30 SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

28. Notoparmelia (typus species P. tenuirrima). Including following Parmelia from Australasia species:Parmelia tenuirrima, P. crambidiocarpa, P. subtestacea, P. signifera, P.cunninghamii. (13 likely) Australasia.Temperate moderately oceanic. Foliaceous Cortex gray to white. Medulla white. Medullary metabolites β-Orcinol depsidones, Orcinol depsidones. Lobes subiregular. Thallus variate but frecuently adnate Non-pored epicortex. Pseudocyphellae efigurate, laminal. Rhizine furcatae to multibranched . Ascospores: medium size (11-21x6-10µm).

31 SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

29. Parmelia (typus species P. saxatilis). Including following Parmelia species: Parmelia saxatilis, P. ernstiae, P. submontana, P. pinnatifida, P. discordans, P. omphalodes, P. serrana, P. sulcata, P. squarrosa, P. barrenoae, P.adaugescens, P. marmariza.(30) Eurasia, North Africa and North America.Temperate moderately oceanic. Foliaceous Cortex gray to white. Medulla white. Medullary metabolites β-Orcinol depsidones, Orcinol depsidones. Lobes subiregular. Thallus variate but frecuently adnate Non-pored epicortex. Pseudocyphellae efigurate, laminal. Isolichenan. Rhizine simple to furcatae and squarrose. Ascospores: medium size (10-27x6-15µm).

32 SHORT DIAGNOSE OF THE STUDIED GENERA OF PARMELIOID GENERA

30. Relicina (typus species R. relicinula). Including also Relicinopsis intertexta and Relicinopsis malaccensis. (56) Pantropical. Tropical, subtropical to temperate. Foliaceous Cortex yellowish green. Medulla white. Medullary metabolites β-Orcinol depsides and depsidones, Orcinol depsides and depsidones and aliphatic acids. Lobes sublinear. Cilia bulbate or not. Thallus variate but frecuently adpress. Pored epicortex. Pseudocyphellae lacking. Isolichenan. Rhizine simple to agglutinated. Ascospores: small size (5-8x3-5µm).

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SYSTEMOL Members contributors: Ana Crespo, Paloma Cubas, Pradeep K. Divakar, Ruth del Prado, Sergio Pérez Ortega, Zuzana Ferencova, Guillermo Amo de Paz, Oscar Blanco Alcalá, Beatriz Roca Valiente, M Carmen Molina.

COAUTHORS IN PARSYS 08’

*DIVAKAR, PRADEEP K.1, CRESPO, ANA1, KAUFF, FRANK2, DEL PRADO, RUTH1, PEREZ-ORTEGA, SERGIO1, AMO DE PAZ, GUILLERMO1, FERENCOVA, ZUZANA1, BLANCO, OSCAR1, ARGUELLO, ARTURO1, MILLANES, ANA3, MOLINA, M. CARMEN3, NORMORE, M.P.4, WEDIN, MATS5, APTROOT, ANDRE6, BUNGARTZ, FRANK7, CALVELO, SUSANA8, CANDAN, MEHMET9, COLE, MARIETTE10, ELIX, JONH A.11, ERTZ, DAMIEN12, GOFFINET, BERNARD13, KNIGHT, ALLISON14, LENDEMER, JAMES15, LINDBLOM, LOUISE16, LUECKING, ROBERT17, LUTZONI, FRANCOIS18, MATTSSON, JAN-ERIC19, MESSUTI, M. INÉS20, PERLMUTTER, GARY21, RICO, VICTOR J.1, SPRIBILLE, TOBY22, STEFFEN, U. PAULS23, SWEAT, K.24, THELL, ARNE25, THOR, GÖRAN26, URBANAVICHUS, GENADII27, and LUMBSCH, H.THORSTEN17

1Universidad Complutense de Madrid, Departamento de Biología Vegetal II, Plaza de Ramón y Cajal s/n, 28040 Madrid, Spain; 2Molecular Phylogenetics FB Biologie, 13/276 TU Kaiserslautern Postfach 304967653 Kaiserslautern, Germany; 3Universidad Rey Juan Carlos, Área de Biodiversidad y Conservación, c/ Tulipán s/n., 28933 Móstoles (Madrid), Spain; 4Dipartimento di Biologia Universit di Trieste Via Giorgieri 10 Trieste, Italy; 5Cryptogamic Botany Swedish Museum of Natural History P.O. Box 50007 SE-104 05 Stockholm, Sweden; 6ABL Herbarium, G.v.d.Veenstraat 107, NL-3762 XK, Soest, Netherlands; 7Charles Darwin Foundation for the Galapagos Islands, Av. 6 de Diciembre N 36-109 y Pasaje California, Post Box 17-01-3891, Quito, Ecuador; 8Centro Regional Universitario Bariloche, Universidad Nacional del Comahue, 8400 Bariloche, Argentina; 9Department of Biology, Faculty of Science, Anadolu University, Eskişehir, Turkey; 103010 West 112th St., 55431-3815, Bloomington. MN, USA; 11Dept. of Chemistry, Australian National University, P.O. Box 4, 0200, Canberra, Australia; 12National Botanic Garden of Belgium, Department of Bryophytes-Thallophytes, Domaine de Bouchout, B-1860 Meise, Belgium; 13Ecology and Evolutionary Biology, 75 North Eagleville road, University of Connecticut, Storrs CT, 06269-3043 USA; 14Department of Botany, University of Otago, Box 56, Dunedin, New Zealand; 15The Academy of Natural Sciences of Philadelphia, Botany Department, The Academy of Natural Sciences of Philadelphia, 1900 Benjamin Franklin Pky. 19103, Philadelphia, USA; 16Dept of biology (BIO), Univ of Bergen, Allegaten 41, 5007, Bergen, Norway; 17Department of Botany, The Field Museum, 1400 S. Lake Shore Drive Chicago, IL 60605, USA; 18Dept. of Biology, Duke University, Box 90338, 27708, Durham, North Carolina USA; 19School of Life Science, Södertörns högskola, SE-141 89 Huddinge, Sweden; 20Dept. Botanica, Centro regional Universitario Bariloche, Universidad Nacional del Comahue Quintral

34 1250 8400SC de Bariloche, Rio Negro Argentina; 21University of North Carolin Herbarium, CB 3280 Coker may, University of North Carolina, Chapel Hill NC 27599- 3280,USA; 22University of Göttingen, Untere Karsp 2, D-37073 Göttingen, Germany; 23Department of Zoology, The Field Museum, 1400 S. Lake Shore Drive Chicago, IL 60605, USA 24Department of Integrated Natural Sciences, Arizona State University, West Campus AZ, USA, 25Lund University, Dept. of Systematic Botany, Ostra-Vallgatan 18- 20, S-22361Lund, Sweden; 26Department of Ecology, Swedish University of Agricultural Sciences, Box 7002, SE-750 07 Uppsala, Sweden; 27Institute of the Industrial Ecology оf the North, Kola Science Center, Russian Academy of Sciences Fersmana str., 14a 184209, Apatity Murmansk region, Russia.

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