Lichen determination keys - neotropical Parmotrema - |
MASON HALE's key to Parmotrema, revised edition:
key to wide-lobed parmelioid species occurring in Tropical America
(genera Canomaculina, Parmotrema, Rimelia, Rimeliella)
by H. Sipman; last update 28 October 2005
This is an update for an unpublished key which Mason HALE distributed to several lichenologists in the late seventies. I have used it until it got dog-ears and became irreadible. Therefore I have made a new, updated version. Following the intention of Mason, it is placed on the Internet for general use. Adriano Spielman provided useful suggestions.
1a Thallus with isidia or dactyles (which may become pustulate or be mixed with soredia) 2 1b Thallus with soredia; soralia linear or round to diffuse, occasionally produced on pustules; no isidial initials present 51 1c Thallus without isidia or soredia 121 2a Cilia completely lacking around lobe tips and in lobe axils 3 2b Cilia well developed around lobe tips to quite sparse and present only in lobe axils 14 3a Medulla yellow or pale yellow orange 4 3b Medulla white (sometimes with patches of orange-red skyrin or other anthraquinones near lower cortex; in decaying plants salazinic acid may cause red staining) 6 4a Isidia slender, dense, not pustular, not hollow; medulla K-, C+ red (gyrophoric acid and entothein); large, leathery plants, similar to P. tinctorum; widespread P. endosulphureum (Hillm.) Hale (Hale 1965: 251) 4b Isidia coarse, pustular, hollow; medulla K+ yellow turning red or C+ red (salazinic or gyrophoric acid) 5 5a Medulla K+ yellow turning red, C- (salazinic acid); Venezuelan Andes P. enteroxanthum Hale (Mycotaxon 5: 434) 5b Medulla K+ yellow, C+ red (gyrophoric acid); southern Brazil (synonymized with P. flavomedullosum) Parmelia elabens Kurok. 6a Isidia coarse, dactyliform or pustular or becoming fatiscent -coralloid 7 6b Isidia thin, cylindrical, simple to sparsely branched, laminal 10 7a Isidia dactyliform 8 7b Isidia pustular or becoming fatiscent-coralloid 9 8a Unidentified fatty acid present; conidia unknown; Argentina P. alidactylatum Estrabou & Adler 8b Caperatic and unidentified aliphatic acids present; conidia sublageniform or weakly bifusiform, 5-7.5 mu long; SE Brazil (Rio Grande do Sul) P. dactylosum Fleig 8c Norlobaridon and loxodic acid present; conidia sublageniform, 6 mu long; SE Brazil (Rio Grande do Sul) P. dissimile Fleig 9a Isidia becoming fatiscent-coralloid, mostly marginal; protocetraric acid present; laciniae developing into large, coralloid (Stereocaulon-like) clusters P. fasciculatum (Vain.) Hale (Hale 1965: 252) 9b Isidia pustular; protocetraric acid present; orange yellow pigment near lower surface (skyrin); Parana P. gibberosum Kurok. 10a Medulla C+ red (lecanoric acid); widespread and common P. tinctorum (Nyl.) Hale (Hale 1965: 264) 10b Medulla C- 11 11a Medulla K+ yellow turning red (norstictic acid) (when strongly applanate and with rhizoid initials near the lobe tips, cf. Parmotremopsis antillensis (Nyl.) Elix & Hale) P. guyanum Hale 11b Medulla K- or K+ slowly yellowish to brownish (protocetraric acid or fatty acids) 12 12a Thallus yellowish green (usnic acid present in cortex; proto- cetraric acid) P. conformatum (Vain.) Hale (Hale 1965: 270) 12b Thallus mineral greenish grey to white (atranorin present in cortex) 13 13a Protocetraric acid present; corticolous at higher elevations in Caribbean region P. peralbidum (Hale) Hale (Hale 1965: 257) 13b Aliphatic acids as in P. praesorediosum present; Brazil (Paraná), saxicolous P. praeisidiosum Fleig 14a (2) Medulla yellow to orange throughout 15 14b Medulla white (sometimes with patches of orange-red skyrin or other anthraquinones near lower cortex; in decaying plants salazinic acid may cause red staining) 18 15a Medulla deep yellow (vulpinic acid); isidia fine, cylindrical; cortex often cracked, revealing the pigment; widespread in the lowlands P. sulphuratum (Nees) Hale (Hale 1965: 312) 15b Medulla pale yellow to orange (no vulpinic acid) 16 16a Medulla pale yellow orange, K+ yellow turning red (salazinic acid); isidia very coarse, tall; Venezuelan Andes, 2100 m, on rock P. enteroxanthum Hale 16b Medulla orange to ochraceous, K+ dark purple (pigment!) or K-; corticolous 17 17a Thallus isidiate; isidia fragile, sometimes ciliate, coralloid- branched and brown-tipped; margins with flaking cortex; medulla completely pigmented, C-, KC- (pigment only); Amazonia, Guianas P. aurantiacoparvum Sipman (Mycotaxon 44: 4) (Acc. to Kurokawa & Moon (1998) identical with next species) 17b Thallus coarsely sorediate or isidiate-sorediate, with laminal and submarginal soralia; unpigmented medulla C-, KC+ rose (alectoronic acid); southern Brazil P. hypomiltoides (Vain.) Kurok. (Hale 1965: 293) 18a (14) Thallus yellowish green (usnic acid dominant in cortex) 19 18b Thallus mineral greenish grey to white (atranorin dominant in cortex) 23 19a Medulla P-, C+ rose (gyrophoric acid) or C- (fatty acid); Caribbean, Mexico, USA, southern Brazil P. xanthinum (Müll. Arg.) Hale (Hale 1965: 275) (incl. P. madagascariaceum (Hue) Hale (Hale 1965: 275); Parmelia aberrans (Vain.) Des Abb. (Hale 1965: 269) 19b Medulla P+ orange or red, C- 20 20a Medulla K- or K+ slowly yellowish to brownish (protocetraric acid) 21 20b Medulla K+ rapidly yellow turning red (salazinic acid) 22 21a Isidia fine; widespread P. conformatum (Vain.) Hale (Hale 1965: 270) 21b Isidia coarse and breaking open; above 2500 m in northern Andes P. fractum (Hale) Hale (Hale 1965: 292) 22a Thallus brown below (darkening with age) and finely short rhizinate to the margin; USA - Mexico, Caribbean Canomaculina neotropica (Kurok.) Elix (P. neotropicum Kurok.) (Mycotaxon 5: 437) 22b Thallus black below with a bare brown marginal zone; widespread P. flavescens (Kremp.) Hale (Hale 1965: 272) 23a (18) Lower surface brown (darkening at the center with age), finely short-rhizinate with scattered much longer rhizines to the margin; upper surface distinctly white-maculate, shiny 24 23b Lower side jet black, sparsely rhizinate with a broad, usually bare, black or brown marginal zone below the lobe tips; upper surface dull 27 24a Medulla K- or K+ slowly yellowish to brownish 25 24b Medulla K+ yellow turning red (salazinic acid) 26 25a Medulla P- (norlobaridone and loxodin); Caribbean & Mexico Canomaculina subtinctoria (Zahlbr.) Elix (Rimeliella subtinctoria (Zahlbr.) Kurok., P. subtinctorium (Zahlbr.) Hale, Canomaculina haitiensis (Hale) Elix = P. haitiense (Hale) Hale) (Hale 1965: 317) 25b Medulla P+ red (fumarprotocetraric and traces of protocetraric acid); southern Brazil Canomaculina fumarprotocetrarica (Marcelli & Hale) Elix (Rimeliella fumarprotocetrarica (Marcelli & Hale) Kurok., P. fumarprotocetraricum Marcelli & Hale) (Mycotaxon 25: 88) 26a Trace of usnic acid often present in cortex; norlobaridone and loxodin lacking; USA - Mexico, Caribbean Canomaculina neotropica (Kurok.) Elix (Rimeliella neotropica (Kurok.) Kurok., P. neotropicum Kurok.) 26b Usnic acid lacking; norlobaridone and loxodin present Canomaculina subtinctoria (Zahlbr.) Elix (Rimeliella subtinctoria (Zahlbr.) Kurok., P. subtinctorium (Zahlbr.) Hale) (Hale 1965: 317) 27a (23) Upper cortex finely reticulately cracked to lobe tips; rhizines rather numerous, usually present till near the margin of the lobe tips, of very unequal length 28 27b Upper cortex continuous, cracking irregularly with age in the center of the thallus; rhizines scarce, absent from a broad marginal zone in the lobe tips 29 28a Medulla K+ yellow turning red, UV- (salazinic acid); widespread Rimelia subisidiosa (Müll. Arg.) Hale & Fletcher (P. subisidiosum (Müll. Arg.) Hale) 28b Medulla K-, UV+ yellow (norlobaridone and lichexanthone); Venezuela Rimelia bonplandii Nata 29a Isidia cylindrical, uniform, mostly unbranched 30 29b Isidia becoming subsorediate or fatiscent-coralloid and tall or pustulate 35 30a Medulla K- 31 30b Medulla K+ yellow or yellow turning red 32 31a Medulla UV+ white (alectoronic acid); orange-red medullary pigment usually present in lower part of medulla; saxicolous in pine-oak forest of Mexico-Honduras P. erasmium (Hale) Hale (Hale 1965: 290) (When medulla unpigmented, try also P. mellissii, couplet 37) 31b Medulla UV- (olivetoric acid); no medullary pigment; Brazil (Rio Grande do Sul), on rock P. horridum Fleig 32a Salazinic acid present; medulla C+ or UV+ 33 32b Stictic acid present; medulla C-, UV- 34 33a Medulla C-, UV+ yellow (lichexanthone); North & Central America P. ultralucens (Krog) Hale 33b Medulla C+ rose, UV- (gyrophoric acid) P. concurrens Hale (Mycotaxon 5: 432) 34a Isidia dense, (partly) ciliate; bare zone below narrow; wide- spread P. crinitum (Ach.) Choisy (Hale 1965: 284) 34b Isidia dense, not ciliate; bare zone below narrow; widespread? P. internexum (Nyl.) Hale ex DePriest & B. Hale 34c Fine-lobulate (no true isidia); mangrove southern Brazil P. lobulatum Marcelli & Hale (Mycotaxon 25: 88) 35a Medulla K+ yellow turning red (salazinic acid) P. coralliforme (Hale) Hale (Hale 1965: 324) 35b Medulla K- or K+ persistently yellow 36 36a Isidia tall, fatiscent-coralloid; medulla K+ yellow (atranorin); Guyana, Amazonia P. flavotinctum (Hale) Hale (Hale 1965: 291) 36b Isidia pustular, not becoming sorediate; medulla K-, P+ orange (protocetraric acid); Parana P. gibberosum Kurok. 36c Isidia short, not fatiscent but becoming sorediate or mixed with soredia; medulla K- 37 37a Medulla strongly UV+ white (alectoronic acid); widespread P. mellissii (Dodge) Hale (Hale 1965: 297) 37b Medulla weakly UV+ pale white or negative (norlobaridone); southern Brazil P. paulense (Zahlbr.) Hale (Hale 1965: 334) 51a (1) Cilia completely lacking at lobe tips and in the lobe axils 52 51b Cilia well developed around lobe tips to quite sparse and present only in lobe axils 67 52a Medulla pale yellow, at least in lower half, to yellow orange throughout 53 52b Medulla white (sometimes with patches of orange-red skyrin or other anthraquinones near lower cortex; in decayed plants salazinic acid may cause red staining) 57 53a Soralia broad and diffuse, mostly submarginal; medulla with gyrophoric acid and entothein; Venezuela - Argentina P. flavomedullosum Hale (Mycotaxon 1: 110) 53b Soralia narrow, linear, marginal for the most part 54 54a Medulla P+ red (protocetraric and echinocarpic acids); South America; resembles P. dilatatum, but with pigmented medulla P. affluens (Hale) Hale 54b Medulla P- 55 55a Medulla C+ rose (entothein, gyrophoric acid and lichexanthone in medulla); soralia strictly marginal; Paraguay - Argentina P. conjunctum Hale (Mycotaxon 1: 108) 55b Medulla C+ yellowish or C- 56 56a Medulla C- or C+ dull yellowish (fatty acid, entothein); cilia may be very sparse; Sao Paulo (Brazil) P. araucariarum (Zahlbr.) Hale (Hale 1965: 238) 56b Medulla C- (diffractaic and barbatic acids), pigmented only in lower half; soredia subgranular; Chiapas P. matudae (Kurok.) Hale ex DePriest & B. Hale 57a (52) Medulla K+ yellow turning red 58 57b Medulla K- or slowly yellowish 59 58a Thallus large and leathery, without pustules; medulla with salazinic acid P. cristiferum (Tayl.) Hale (Hale 1965: 241) 58b Thallus smaller, membraneous, with soredia produced on pustules; medulla with stictic acid; Andean P. bangii (Vain.) Hale (Hale 1965: 281) 58c Thallus smaller, membraneous, without pustules; medulla with norstictic acid; Mexico-Nicaragua, Minas Gerais P. rubifaciens (Hale) Hale (Hale 1965: 261) 59a Medulla P+ red (protocetraric acid); cortex often yellowish, with atranorin and usnic acid, or usnic acid alone 60 59b Medulla P-; cortex grey, with atranorin alone 64 60a Cortex and/or soralia with usnic acid 61 60b Usnic acid completely absent 63 61a Thallus yellowish (no atranorin), membranous; medulla with additionally echinocarpic acid; Amazonia, Guianas P. aptrootii Aubel (Mycotaxon 44: 3) 61b Thallus greyish (with atranorin); medulla without echinocarpic acid 62 62a Soralia yellowish, with usnic acid; lowlands of Caribbean, Central America and Mexico P. dominicanum (Vain.) Hale (Hale 1965: 248) 62b Thallus membranous; soralia white, on marginal laciniae; highlands, widespread P. robustum (Degel.) Hale 63a Thallus coriaceous, without orange pigment near lower surface; widespread P. gardneri (Dodge) Sérus. (Swinscow & Krog 1988: 176) 63b Thallus membranous; with orange anthraquinone pigment scattered near lower cortex; Para (Brazil) - Guianas, lowlands P. subochraceum Hale (Bibl. Lich. 38: 117) 64a Medulla K+ yellowish; always saxicolous P. mordenii (Hale) Hale 64b Medulla K-; usually corticolous; widespread 65 65a Medula C+ red (lecanoric acid) P. austrosinense (Zahlbr.) Hale (Hale 1965: 238) 65b Medulla C- 66 66a Lobes broad (to 10 mm wide); thallus leathery; medulla KC+ rose, UV+ white (alectoronic acid); Uruguay P. exquisitum (Kurok.) DePriest & B. Hale 66b Lobes narrower (4-5 mm wide); membranous; medulla KC-, UV- (unknown fatty acids); soralia on top of dactyls; Argentina P. soredioaliphaticum Estrabou & Adler 66c Lobes narrower (4-5 mm wide); membranous; medulla KC-, UV- (caperatic acid?); soredia not on top of dactyls; widespread P. praesorediosum (Nyl.) Hale (Hale 1965: 258) 67a (51) Medulla orange or pale yellow, especially in lower half 68 67b Medulla white (sometimes with patches of orange-red skyrin or other anthraquinones near lower cortex; in decayed plants salazinic acid may cause red staining) 69 68a Pigment pale yellow, mainly in lower half of the medulla, K-; unpigmented medulla C+ rose (gyrophoric acid); West Indies, South America (if pigment K+ purple, see P. sancti-angelii) P. permutatum (Stirt.) Hale (Hale 1965: 302) 68b Pigment orange; unpigmented medulla C-, KC+ rose (alectoronic acid); coarsely sorediate or isidiate-sorediate, laminal and submarginal soralia; southern Brazil P. hypomiltoides (Vain.) Hale ex DePriest & B.Hale (Hale 1965: 293) 69a Thallus distinctly yellow to greenish yellow (usnic acid with or without atranorin in the cortex) 70 69b Thallus mineral or greenish grey to white (usnic acid lacking in cortex) 75 70a Soralia coarse, pustular, laminal; cortex breaking apart; occurring on soil and humus in paramo P. fractum (Hale) Hale (Hale 1965: 292) 70b Soralia round or linear, mostly marginal; cortex entire; usually corticolous in forests, or on rocks 71 71a Medulla C+ red (gyrophoric acid); Brazil P. nylanderi (Lynge) Hale 71b Medulla C-; known from various countries 72 72a Medulla K+ yellow turning red at once (salazinic acid); soralia on more or less involute lobes; pine-oak forest zone in Mexico P. mirandum (Hale) Hale (Hale 1965: 273) 72b Medulla K- or K+ dull yellowish slowly (protocetraric acid); ciliate mainly in axils; mostly with additionally atranorin in the cortex 73 73a Thallus weakly yellowish green, atranorin dominant in the cortex; soredia often on marginal lobules; medulla with echinocarpic acids; lowlands, widespread P. dilatatum (Vain.) Hale 73b Id., medulla without echinocarpic acid; highlands, widespread P. robustum (Degel.) Hale 73c Thallus mostly strongly yellow green, without or with traces of atranorin in cortex; medulla without echinocarpic acid 74 74a Medulla with malonprotocetraric acid; atranorin absent; Mexico and Caribbean P. viridiflavum (Hale) Hale (Hale 1965: 274) 74b Medulla with protocetraric acid alone; atranorin in the cortex; Ecuador-Venezuela, northern Andes at 2400-3500 m. P. virescens Hale (Mycotaxon 25: 90) 75a (69) Lower side rhizinate to the margin with fine, short rhizines mixed with long ones; lower side pale brown to black 76 75b Lower side with a wide, bare marginal zone or with scattered rhizines of very variable length near the margins; lower side dark brown to black, sometimes pale at the margins 82 76a Lower surface pale brown (sometimes darkening at the center) 77 76b Lower surface brown to black, at least at the center 80 77a Medulla UV+ yellow (lichexanthone) 78 77b Medulla UV- 79 78a Soredia mixed with short cilia; norlobaridone absent; southern Brazil Canomaculina spinibarbis (Kurok.) Elix (P. spinibarbe (Kurok.) Hale) 78b Soredia without cilia; norlobaridone present; Venezuela Canomaculina larensis (López Fig.) Elix (P. larense López Fig.) (Phytologia 44: 89) 79a Medulla with salazinic acid alone; cortex often with additional usnic acid; widespread Canomaculina subsumpta (Nyl.) Elix (Rimeliella subsumpta (Nyl.) Kurok., P. subsumptum (Nyl.) Hale (Hale 1965: 315) 79b Medulla with norlobaridone and loxodin, sometimes with salazinic acid; no usnic acid in cortex; widespread; perhaps better included in C. subsumpta? Canomaculina conferenda (Hale) Elix (Rimeliella conferenda (Hale) Kurok., P. conferendum Hale) (Mycotaxon 5: 433). Salazinic acid-containing specimens are sometimes separated as Canomaculina reitzii (Hale) Elix = P. reitzii Hale (Mycotaxon 5: 439) 80a Lower side brown; medulla with salazinic acid; soralia linear on margins; Mexico - Costa Rica, Caribbean, southern Brazil - Argentina Canomaculina leucosemotheta (Hue) Elix (P. leucosemothetum (Hue) Hale) (Hale 1965: 330) 80b Lower side black; medulla without salazinic acid 81 81a Medulla with stictic acid; soralia laminal, round; Mexico, Andes, Argentina Canomaculina muelleri (Vainio) Elix & Hale 81b Medulla without lichen substance; Andes, Uruguay Canomaculina pilosa (Stizenb.) Elix & Hale 82a (75) Upper cortex usually distinctly and finely reticulately cracked or reticulately white-maculate (low power magnification); lower side rhizinate till margin with rhizines of very unequal length 83 82b Upper cortex continuous, dull; if maculate, the maculae not reticulately arranged but scattered; lower side with wide, bare marginal, brown or white to mottled zone 87 83a Medulla K+ yellow turning red 84 83b Medulla K- or K+ slowly dingy yellow 85 84a Medulla with salazinic acid alone; common and widespread Rimelia reticulata (Tayl.) Hale & Fletcher (P. reticulatum (Tayl.) Choisy) 84b Medulla with norlobaridone and variable amounts of salazinic acid; uncommon Rimelia commensurata (Hale) Hale & Fletcher (P. commensuratum (Hale) Hale) 84c Medulla with salazinic acid and lichexanthone; Parana Rimelia pontagrossensis Eliasaro & Adler 85a Medulla P-, KC- (caperatic acid alone) P. simulans (Hale) Hale 85b Medula P+ orange or KC+ rose (depsidones present) 86 86a Medulla P-, KC+ rose (diffractaic acid and lichexanthone); Brazil Rimelia diffractaica (Essl.) Hale & Fletcher (P. diffractaicum (Essl.) Hale) 86b Medulla P-, KC+ rose (norlobaridone); Mexico - Central America only? Rimelia commensurata (Hale) Hale & Fletcher (P. commensuratum (Hale) Hale) 86c Medulla P+ orange (succinprotocetraric and fumarprotocetraric acids); Brazil - Paraná Rimelia succinreticulata Eliasaro & Adler 87a (82) Lower surface with a broad bare white zone; lobes rather suberect 88 87b Lower surface light brown to black, rarely with mottled white areas at the margin 91 88a Medulla K- 89 88b Medulla K+ yellow turning red 90 89a Medulla UV+ white, KC+ rose (alectoronic acid) P. louisianae (Hale) Hale 89b Medulla UV- or weak, KC+ rose or KC- (norlobaridone and/or protolichesterinic acid); Mexico - Costa Rica P. hababianum (Gyeln.) Hale (Hale 1965: 325) Norlobaridone-containing specimens are sometimes separated as P. yodae (Kurok.) Hale; (see also C. conferenda) 90a Stictic acid present in addition to norstictic acid; Mexico P. hypoleucinum (Stein.) Hale 90b Stictic acid absent, only norstictic present; Mexico P. hypotropum (Nyl.) Hale (Hale 1965: 328) 91a (87) Medulla K+ yellow or yellow turning red at once (salazinic or stictic acids) 92 91b Medulla K- or K+ slowly yellowish 97 92a Medulla K+ persistent yellow (stictic acid) 93 92b Medulla K+ yellow turning red (salazinic acid) 94 93a Soralia orbicular on small lobules or submarginal in revolute lobes; hymenium 70-80 mu high; spores 22-30 x 13-16 mu; Caribbean, Mexico-Panama (temperate regions) P. chinense (Osbeck) Hale & Ahti (syn. P. perlatum (Huds.) Hale) (Hale 1965: 300) 93b Soralia diffuse, coarse and pustular ("cortex disintegrating in large areas near the margins and developing into sorediate pustules"; hymenium 70-80 mu high; spores 26-36 x 11-16 mu; Andean P. bangii (Vain.) Hale (Hale 1965: 281) 94a Cortex finely reticulately cracked or reticulately white- maculate Rimelia reticulata (Tayl.) Hale & Fletcher (P. reticulatum (Tayl.) Choisy) 94b Cortex continuous; cracked only with age at center 95 95a Thallus completely divided into 2-7(-12) mm wide, sublinear laciniae; sometimes with traces of gyrophoric and/or protocetraric acid; Mexico - Chile P. paramoreliense W.L. Culb. & C.F. Culb. 95b Thallus wide-lobes, irregular 96 96a Soralia marginal, often linear; cilia usually very sparse and tiny; widespread P. cristiferum (Tayl.) Hale (Hale 1965: 241) 96b Soralia submarginal on lobules, orbicular; USA P. margaritatum (Hue) Hale (Hale 1965: 296) 96c Soralia on top of laciniae; mountains of Mexico - Costa Rica (temperate) P. stuppeum (Tayl.) Hale (Hale 1965: 308) 97a (91) Medulla P+ red (protocetraric acid) 98 97b Medulla P- 102 98a Soralia pustular, laminal in broad zone; low elevations near Sao Paulo P. madilynae Fletcher (Mycotaxon 25: 88) 98b Soralia coarse, (sub)marginal, subfatiscent, mostly on lobules; saxicolous; protocetraric + gyrophoric acids; Parana (Brazil) P. schindleri Hale (Mycotaxon 25: 89) 98c Soralia powdery in linear or orbicular marginal groups 99 99a Protocetraric acid and echinocarpic acid together P. dilatatum (Vain.) Hale 99b Protocetraric acid alone present 100 100a Cilia conspicuous, 2-3 mm long, distributed around lobe tips; soralia fine; Mexico, southern Brazil P. subarnoldii (Abb.) Hale (Hale 1965: 309) 100b Cilia shorter, often sparse 101 101a Cilia often sparsely developed, mostly in lobe axils; soredia coarse, on marginal laciniae; widespread in the mountains P. robustum (Degel.) Hale 101b Thallus coriaceous; cilia sparse; widespread P. gardneri (Dodge) Sérus. (Swinscow & Krog 1988: 176) 102a (97) Medulla C+ rose or red (gyrophoric acid alone); orange- yellow pigment (rhodophyscin) sometimes present near the lower cortex, K+ purple (if pigment K-, see P. permutatum); conidia weakly sublageniform (6-8 x 1 mu); widespread P. sancti-angelii (Lynge) Hale (Hale 1965: 306) 102b Medulla C+ rose (gyrophoric acid and norlobaridone); no pigment in medulla; conidia filiform; India, SE Brazil P. indicum Hale (Hale 1977: 436) 102c Medulla C- 103 103a Medulla KC-, UV- (no alectoronic acid) 104 103b Medulla KC+ red or rose, UV+ white (alectoronic and/or a- collatolic acid) 105 104a Saxicolous; with protolichesterinic acid; resembling P. praesorediosum but ciliate; widespread P. grayanum (Hue) Hale (Hale 1965: 292) 104b Corticolous; with constipatic acid; Para (Brazil) P. ciliiferum Hale (Bibl. Lich. 38: 112) 105a Lobes rather narrow and laciniate; soralia mostly orbicular on the laciniae, some marginal; marginal zone below narrow, mostly black; Caribbean, Mexico-Guatemala (+ temperate areas) P. arnoldii (DR) Hale (Hale 1965: 279) 105b Lobes broader, not laciniate; soralia linear and marginal: bare zone below broad, brown 106 106a Soredia intermixed with isidia and cilia; widespread P. mellissii (Dodge) Hale (Hale 1965: 297) 106b Soredia without isidial initials or cilia intermingled 107 107a Soralia not pustular, linear; medulla often with skyrin; on trees; cilia 3-6 mm long; widespread P. rampoddense (Nyl.) Hale (Hale 1965: 304) 107b Pustulate-sorediate along margins; saxicolous; cilia 2-4 mm long; Venezuelan Andes P. sorediiferum Hale (Mycotaxon 25: 89) 121a (1) Cilia completely lacking around lobe tips and in lobe axils 122 121b Cilia well developed around lobe tips to quite sparse and present only in lobe axils 136 122a Medulla yellow or yellow orange throughout or for the most part, and especially in the apothecial amphithecium 123 122b Medulla white (sometimes with patches of orange-red skyrin or other anthraquinones near lower cortex; in decayed plants salazinic acid may cause red staining) 125 123a Medulla pale orange to salmon red, P+ red (protocetraric acid); apothecia imperforate; hymenium 75-80 mu high; spores 22-24 x 11-12 mu; Venezuelan Andes P. betaniae Hale (Mycotaxon 25: 87) 123b Medulla P- 124 124a Medulla yellow-orange, C+ red (gyrophoric acid); spores unknown; widespread P. crocoides (Hale) Hale (Hale 1965: 244) 124b Medulla pale yellow, C+ yellow-orange (barbatic and obtusatic acids, entothein); spores 21-26 x 9-13 mu; Mexico (Chiapas 1000 m.) P. myelochroum (Hale) Hale (Hale 1965: 256) 125a Medulla C+ blood red (lecanoric acid); apothecia if present perforate P. andinum (Müll. Arg.) Hale (Hale 1965: 236) 125b Medulla C-; apothecia if present imperforate 126 126a Medulla P- 127 126b Medulla P+ red or orange 128 127a Caperatic acid present; hymenium 60-90 mu high; spores to 15-21 x 7-11 mu long; Mexico-Guatemala, Bolivia-Uruguay-Mato Grosso P. mesotropum (Müll. Arg.) Hale (Hale 1965: 255) 127b Diffractaic and barbatic acid present; hymenium 100-130 mu high; spores 21-30 x 10-15 mu; Mexico, Caribbean; medulla sometimes orange near lower cortex P. mesogenes (Nyl.) Hale (Hale 1965: 254) 128a Medulla K+ yellow turning red (salazinic and/or norstictic acid) 129 128b Medulla K- or faint yellow (protocetraric acid) 131 129a Salazinic acid alone present; hymenium 100-120 mu high; spores 28-32 x 14-16 mu; widespread P. latissimum (Fée) Hale (Hale 1965: 253) 129b As 129a, but thallus lobes deeply laciniate (apothecia unknown); Mexico P. acutatum Kurok. (Kurokawa 2001: 3) 129c Additional medullary substances present (TLC); hymenium under 80 mu high; spores under 25 mu long; South America only 130 130a Norstictic and salazinic acid present; hymenium 80-90 mu high; spores 20-25 x 10-12 mu; Guyana - southern Brazil P. crassescens (Stirt.) Hale (Hale 1965: 240) 130b Norstictic and echinocarpic acid present; spores 20-22 x 14 mu; Mato Grosso, Brazil P. wrightii Ferraro & Elix (Mycotaxon 49: 405) 131a Thallus on rock only; spores up to 15 mu long 132 131b Thallus on bark, occasionally on rock; spores over 15 mu long 133 132a Thallus rather closely adnate on rocks; hymenium 70-80 mu high; spores 11-14 x 5-7 mu; Central America P. eborinum (Hale) Hale (Hale 1965: 249) 132b Thallus laciniate, less closely adnate, on rock; hymenium 50-60 mu high; spores 12-15 x 5-6 mu; Brazil, Colombia P. blanchetianum (Müll. Arg.) Hale (Hale 1965: 281) 133a Lobes with long marginal laciniae; hymenium 90-100 mu high; spores 16-21 x 8-10 mu; Mexico (+ Africa) P. disparile (Nyl.) Hale (Hale 1965: 247) 133b Like 133a, but thallus surface not maculate or finally cracked, and apothecia perforate instead of imperforate; Mexico P. cornigerum Kurok. (Kurokawa 2001: 4) 133c Lobes without conspicuous laciniae 134 134a Protocetraric acid alone present, no pigment near lower cortex; hymenium 70-80 mu high; spores 18-22 x 7-10 mu; widespread (divided by Elix (1998) into P. zollingeri s.str., with fumarprotocetraric and succinprotocetraric acid, P. overeemii (Zahlbr.) Elix, with protocetraric acid and slightly larger spores, and P. platyphyllinum (Vain.) Elix, with protocetraric acid and butlerin derivatives; it is unknown which occur in the Neotropics) P. zollingeri (Hepp) Hale (Hale 1965: 267) 134b With additionally usnic acid in the cortex, but no pigment near lower cortex; spores 16-18 x 10 mu; Andes of Peru P. machupicchuense Kurok. 134c With additionally echinocarpic acid or pigment near lower cortex; South America only 135 135a Orange anthraquinone pigment scattered near lower cortex; hymenium 60-65 mu high; spores 30-33 x 15-18 mu; Para (Brazil) P. cachimboense Hale (Bibl. Lich. 38: 110) 135b Echinocarpic acid present with protocetraric acid; hymenium 100 mu high; spores 20-22 x 10 mu; Amazonia (related to P. dilatatum) P. progenes Hale (Mycotaxon 5: 438) 136a (121) Medulla yellow or yellow orange throughout or for the most part, and especially in the apothecial amphithecium 137 136b Medulla white (sometimes with patches of orange-red skyrin or other anthraquinones near lower cortex; in decayed plants salazinic acid may cause red staining) 142 137a Thallus distinctly yellowish green (usnic acid + atranorin in cortex, protocetraric acid and entothein in medulla); cilia scarce; apothecia eciliate; hymenium 70-80 mu high; spores 25- 27 x 12-13 mu; Venezuelan Andes P. lopezii Hale (Mycotaxon 5: 437) 137b Thallus mineral, greenish or whitish grey (usnic acid lacking) 138 138a Medulla P+ red (protocetraric acid, entothein); laciniate; apothecia ciliate; Amazonia, Venezuela P. cristatum (Nyl.) Hale 138b Medulla P- 139 139a Cortex fragile, breaking apart; medulla pale yellow; fatty acid in medulla; apothecia imperforate; hymenium 80-95 mu high; spores 22-27 x 10-12 mu; southern Brazil P. cryptoxanthoides (Kurok.) Hale 139b Cortex entire; medulla more deeply pigmented 140 140a Medulla bright lemon yellow (vulpinic acid alone); southern Brazil P. cornutum (Lynge) Hale (Hale 1965: 283) 140b Medulla lighter yellow 141 141a Medulla lighter yellow, reddish below (entothein and rhodophyscin); southern Brazil P. lyngeanum (Zahlbr.) Hale 141b Medulla lighter yellow above, orange-red below (entothein and barbatic acid); apothecia ciliate; not yet known from the Neotropics P. appendiculatum (Fée) Hale 142a (136) Thallus yellowish green (usnic acid in cortex) 143 142b Thallus mineral to whitish grey (usnic acid lacking) 144 143a Thallus often white-pruinose; medulla with salazinic (and gyrophoric?) acid; saxicolous; southern Brazil - Uruguay P. delicatulum (Vain.) Hale (Hale 1965: 271) 143b Thallus not pruinose; medulla with salazinic acid and unknown substance Q; corticolous; Corrientes (Argentina) P. masonii Ferraro (1979: 193) 144a Lower surface to the margin covered with fine rhizines, mixed with scattered, much longer rhizines, usually pale to brown, often darkening at the center with age, sometimes completely black; upper side maculate 145 144b Lower surface bare or with sparse, long rhizines (dense in Rimelia), mostly with a jet black center, the marginal zone whitish or light brown to black; upper side maculate or not maculate 148 145a Medulla K+ yellow turning red (salazinic acid) 146 145b Medulla K-; cortex with atranorin only (salazinic acid absent); southern Brazil - Argentina Canomaculina recipienda (Nyl.) Elix (Rimeliella recipenda (Nyl.) Kurok., P. recipiendum (Nyl.) Hale) (Hale 1965: 314) 146a Short rhizines sparsely branched and somewhat intricate; apothecia ciliate; thallus not laciniate; lower surface mostly pale brown, darkening only in the center; norlobaridone absent; Uruguay-Argentina Canomaculina uruguensis (Kremp.) Elix (Rimeliella uruguensis (Kremp.) Kurok., P. uruguense (Kremp.) Hale) (Hale 1965: 342) 146b Short rhizines sparsely branched and somewhat intricate; apothecia not ciliate; thallus laciniate with convex laciniae; lower side black; norlobaridone absent; NW Argentina Canomaculina laciniella Ferraro & Elix 146c Short rhizines simple; apothecia not ciliate; thallus laciniate or not; lower side mostly pale brown; norlobaridone sometimes present 147 147a Thallus not laciniate; southern Brazil Canomaculina subcaperata (Krempelh.) Elix (Rimeliella subcaperata (Krempelh.) Kurok., P. subcaperatum (Kremp.) Hale) (Hale 1965: 314) 147b Thallus laciniate, lacinia flat; Argentina - Bolivia Canomaculina cristobalii (L. I. Ferraro & Elix) Elix (Rimeliella cristobalii L. I. Ferraro & Elix) (Mycotaxon 49: 406) 148a (144) Lower surface with a distinct, broad, bare, white or ivory to brown rim; lobes often suberect 149 148b Lower surface with a brown or black, bare or rhizinate rim (or at most somewhat white-mottled); lobes often more adnate, not suberect 160 149a Medulla K+ yellow turning orange or red (norstictic, stictic or salazinic acid) 150 149b Medulla K- 152 150a Medulla K+ yellow (stictic acid); apothecia if present perforate; rim below often mottled white; hymenium 120-150 mu high; spores 23-30 x 12-18 mu; Mexico-Guatemala, Caribbean, southern Brazil (if thallus applanate, marginally rhizinate and with norsticticacid, cf. Parmotremopsis phlyctina (Hale) Elix & Hale) P. eciliatum (Nyl.) Hale (Hale 1965: 289) 150b Medulla K-; apothecia if present imperforate 151 151a Norstictic acid alone present; Jamaica, USA P. perforatum (Jacq.) Hale (Hale 1965: 335) 151b Norstictic acid present with alectoronic acid; southeastern USA P. subrigidum Egan (Egan et al. 2005) 151c Stictic acid present with norstictic acid P. preperforatum (Culb.) Hale 151d Salazinic acid present; thallus often laciniate; Mexico 600-900 m. P. arteagrum Egan 152a Apothecia if present perforate; spores less than 20 mu long (except P. glaucocarpoides) 153 152b Apothecia if present imperforate; spores usually more than 20 mu long 157 153a Medulla UV+ white (alectoronic acid present) 154 153b Medulla UV- (alectoronic absent) 155 154a Thallus maculate; apothecium rim rarely ciliate; southern Brazil - Argentina P. rigidum (Lynge) Hale (Hale 1965: 338) similar to P. subrigidum, but ascospores 18-33 x 9-19 mu instead of 9-15 x 5-9 mu, pycnoconidia usually <12 mu instead of 11-20 mu, absence of norstictic acid and regular presence of a-collatolic acid (Egan et al. 2005); norstictic acid-deficient P. subrigidum may key out here and can be distinguished by ascospores and pycnoconidia. 154b Thallus not maculate; apothecium rim ciliate; Mexico P. wirthii Hale (Bibl. Lich. 38: 118) 155a Medulla C+ red (gyrophoric, fatty acids); maculate; Brazil (+ Africa) P. hanningtonianum (Müll. Arg.) Hale 155b Medulla C- (gyrophoric acid absent) 156 156a Norlobaridone present (UV negative or weak), and/or protolichesterinic acid; spores 15-20 x 8-10 mu; Mexico?, Africa P. abessinicum (Kremp.) Hale (Hale 1965: 320) 156b Fatty acid present only (UV-); spores 25-30 x 12-15 mu; Africa, neotropical? P. glaucocarpoides (Zahlbr.) Hale 157a (152) Medulla C+ red (gyrophoric acid); hymenium 130-150 mu high; spores 24-28 x 14-16 mu; southern Brazil P. catarinae Hale (Mycotaxon 25: 87) 157b Medulla C- (gyrophoric acid absent) 158 158a Medulla KC-, UV. (protolichesterinic acid); apothecia ciliate, with maculate amphithecium; hymenium 80-100 mu high; spores 20- 26 x 10-16 mu; southern Brazil P. melanothrix (Mont.) Hale (Hale 1965: 332) 158b Medulla KC+ rose, UV+ white (alectoronic acid); rim below uniformly white; exciple often short-dentate and ciliate 159 159a Hymenium 100-140 mu high; spores 26-34 x 12-18 mu; medulla with orange skyrin pigment scattered near lower cortex; surface maculate; Mexico - Guatemala, Caribbean, southern Brazil - Argentina P. subrugatum (Kremp.) Hale (Hale 1965: 341) 159b Hymenium 50-70 mu high; spores 11-22 x 6-12 mu; widespread P. argentinum (Kremp.) Hale (Hale 1965: 322) 160a (148) Lower side rhizinate below to the margin, with rhizines of very unequal length; cortex finely reticulately cracked or white-maculate 161 160b Lower side rhizinate below to margin, with short rhizines; upper side heavily maculate; thallus applanate; no substances in medulla; spores 14-19 x 8-12 mu; southern Brazil to Argentina Canomaculina consors (Nyl.) Elix & Hale 160c Lower side sparsely rhizinate, with wide, bare marginal zone in lobe tips; cortex mostly continuous or irregularly cracked with age toward thallus center 163 161a Medulla K+ yellow turning red, UV- (salazinic acid); lobes often laciniate (cf. P. ruptum (Lynge) Hale ex DePriest & B. Hale, see below); widespread Rimelia cetrata (Ach.) Hale & Fletcher (P. cetratum (Ach.) Hale) 161b Medulla K+ yellow turning red, UV+ yellow (salazinic acid and lichexanthone); upper surface faintly maculate; apothecia perforate; spores 14.5-16 x 10-11 mu; Minas Gerais P. lichexanthonicum Eliasaro & Adler 161c Medulla K- 162 162a Caperatic acid present; SE Brazil only? Rimelia macrocarpa (Pers.) Hale & Fletcher (P. macrocarpum (Pers.) Hale) 162b Norlobaridone and loxodin present; SE Brazil only? Rimelia homotoma (Nyl.) Hale & Fletcher (P. homotomum (Nyl.) Hale) 163a (160) Medulla K+ yellow turning orange or red (stictic or salazinic acid) 164 163b Medulla K- 170 164a Apothecia perforate, at least partly 165 164b Apothecia imperforate 166 164c Apothecia unknown 169 165a Lobes rounded, 8-10 mm wide, membranaceous, in the center often laciniate with many pycnidia; cilia 1.5-5 m long; spores 14-16 x 7-9 mu; oak-pine zone of Mexico, USA, Brazil P. eurysacum (Hue) Hale (Hale 1965: 290) 165b As 165a, but lobes coriaceous, in the center crenate; cilia 1-3 mm long, rare; spores 10-14 x 6-10 mu; SE USA to Mexico P. despectum Kurok. (Kurokawa 2001: 5) 165c As 165b, but upper surface strongly maculate, reticulately cracked in older lobes, and lower side sometimes slightly papillate P. permaculatum Kurok. (Kurokawa 2001: 8) 165d Lobes linear-elongate, 2-5(-10) mm wide, completely laciniate; spores 16-20 x 8-10 mu; Mexico 2250-3300 m. P. moreliense (B. de Lesd.) W. Culb. & C. Culb. 165e If lichexanthone present in medulla (UV+ yellow), see P. lichexanthonicum above 166a Upper cortex reticulately white maculate to cracked; not laciniate, with narrow bare marginal zone below; salazinic acid; apothecia eciliate; hymenium 60-70 mu high; spores 16-18 x 9 mu; Costa Rica - Panama, southern Brazil-Paraguay (intermediate with Rimelia; syn. P. expansum Hale, Mycotaxon 5: 435) P. ruptum (Lynge) Hale ex DePriest & B. Hale 166b Upper cortex continuous or cracked with age toward the thallus center; spores more than 20 mu long 167 167a Cilia sparse, only in lobe axils; salazinic acid present P. latissimum (Fée) Hale (Hale 1965: 253) 167b Cilia usually conspicuous around lobe tips; stictic acid present 168 168a Corticolous; cilia fine; stictic acid; hymenium 120-150 mu high; spores 23-30 x 12-18 mu; Mexico-Guatemala, Caribbean, southern Brazil P. eciliatum (Nyl.) Hale (Hale 1965: 289) 168b Lobulate, stictic acid; mangrove of southern Brazil P. lobulatum Marcelli & Hale 169a Thallus with scattered ciliate warts; Guyana Highland P. verrucisetosum Sipman (Mycotaxon 44: 8) 169b Lobes very wide, 10-20 mm; southern Brazil P. mantiqueirensis Hale (Bibl. Lich. 38: 113) 170a (163) Medulla C+ red; hymenium over (80-)90 mu high; spores 24- 35 mu long 171 170b Medulla C-; hymenium and spores various 173 171a Apothecia perforate; gyrophoric acid present; thallus coriaceous; hymenium 60-80 mu high; spores 18-26 x 7-12 mu; Caribbean, elsewhere? P. eunetum (Stirt.) Hale (Hale 1965: 325) 171b Apothecia imperforate; hymenium over (80-)90 mu high; spores 24-35 mu long 172 172a Olivetoric acid present; southern Brazil P. abnuens (Nyl.) Hale (Hale 1965: 277) 172b Gyrophoric and lecanoric acids present; Guianas P. gradsteinii Aubel (Mycotaxon 44: 6) 173a Medulla UV+ white (alectoronic acid present); apothecia perforate or imperforate 174 173b Medulla UV-; apothecia imperforate 179 174a Apothecia perforate 175 174b Apothecia imperforate 178 175a Orange pigment (skyrin) scattered in medulla near lower surface; thallus papery 176 175b Orange pigment completely absent; thallus coriaceous 177 176a Thallus papery, maculate; apothecia pedicellate, with eciliate rim; hymenium 70-80 mu high; spores 17-22 x 9-12 mu; Mexico - Guatemala P. chiapense (Hale) Hale (Hale 1965: 323) 176b Apothecia perforate (to imperforate); lobes becoming short laciniate-lobulate; hymenium ca. 60 mu high; spores 14-15 x 6 mu; southern Brazil P. brasiliense Hale (Bibl. Lich. 38: 109) 177a Pycnoconidia sublageniform, 6-7 mu long; thallus coriaceous; apothecia substipitate; hymenium 50-70 mu high; spores 12-17 x 7-10 mu; cilia sparse; Brazil, elsewhere? P. maclayanum (Müll. Arg.) Hale (Swinscow & Krog 1988: 183) [incl. P. breviciliatum (Hale) Hale (Hale 1965: 282)] 177b Pycnoconidia filiform, 8-11 mu long; spores 13-18 x 10-14 mu; otherwise as a; Argentina P. pseudobreviciliatum Adler, Elix & Hale 178a Thallus rather fragile, marginal cilia 2-5 mm long; medulla sometimes with skyrin; apothecia ciliate; hymenium 75-90 mu high; spores 17-22 x 6-10 mu; southern Brazil P. wainii (A. Sm.) Hale (Hale 1965: 313) 178b Thallus not fragile, marginal cilia 0.5-1.5 mm; medulla with scattered orange skyrin pigment near lower cortex; apothecia ciliate; hymenium 85-90 mu high; spores 28-30 x 10-15 mu; southern Brazil P. maraense Hale (Bibl. Lich. 38: 114) 179a Medulla P- (protocetraric acid absent, fatty acids present); apothecia estipitate, rarely perforate, not ciliate; hymenium 60-70 mu high; spores 22-25 x 10-12 mu; Para (Brazil) P. paraense Hale (Bibl. Lich. 38: 116) 179b Medulla P+ red (protocetraric acid) 180 180a Lobes broad and rotund, without conspicuous laciniae; cilia very sparse, in axils only, absent on tips; widespread; see note under 134a P. zollingeri (Hepp) Hale 180b Lobes more or less marginally dissected and laciniate; cilia present in axils and on lobe tips, sometimes sparse 181 181a Anthraquinone pigment scattered in medulla near lower cortex; apothecia not ciliate; hymenium 60-65 mu high 182 181b Anthraquinone pigment absent; apothecia sometimes ciliate; hymenium usually over 90 mu high 183 182a Thallus not laciniate, lobes 2-4 mm wide; apothecia with inrolled rim; hymenium 60-65 mu high; spores 24-27 x 15-18 mu; Brazil (Mato Grosso) P. confusum Hale (Bibl. Lich. 38: 113) 182b Thallus laciniate, lobes 5-9 mm wide; hymenium 60-65 mu high; spores 28-30 x 10-12 mu; Brazil (Para) P. pigmentosum Hale (Bibl. Lich. 38: 116) 183a Echinocarpic acid present with protocetraric acid; apothecia not ciliate; hymenium 100 mu high; spores 20-22 x 10 mu; Amazonia (related to P. dilatatum) P. progenes Hale (Mycotaxon 5: 438) 183b Echinocarpic acid absent; lobes marginally dissected and laciniate 184 184a Spores less than 20 mu; North America P. submarginale (Michx.) DePriest & B.Hale (syn. P. michauxianum (Zahlbr.) Hale) 184b Spores 26-34 x 12-16 mu; hymenium 90-110 mu; apothecium exciple dentate-laciniate, ciliate; Bolivia, elsewhere? P. merrillii (Vain.) Hale (Hale 1965: 298)
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