Or If Leprose Then with a Distinct Well Developed Prothallus (Not Lepraria)

KEY TO KEYS

1. Thallus not entirely leprose, with portions of the thallus distinct not dissolved into soredia/granules; or if leprose then with a distinct well developed prothallus (not Lepraria)...... …2 2. Thallus AND/OR soralia yellow or orange pigmented...... …Key 1 (Page 2) 2. Thallus AND/OR soralia not yellow or orange pigmented, but red pigments can be present...... …3 3. Thallus UV+ bright yellow (lichexanthone present)…...... Key 2 (Page 7) 3. Thallus UV-, UV+ dull orange/pink or orange or blue-white (without lichexanthone)...... …4 4. Photobiont Trentepholia...... …Key 3 (Page 9) 4. Photobiont not Trentepholia…...... 5 5. Thallus OR soredia with norstictic acid, K+ yellow to red producing large crystals in water mount...... …...... Key 4 (Page 12) 5. Thallus OR soredia without norstictic acid, K-, K+ other colors, or K+ yellow to red but NOT producing large crystals in water mount...... …6 6. Thallus squamulose and on bark (Hypocenomyce species)…...... Key 5 (Page 11) 6. Thallus not squamulose and on bark (not Hypocenomyce species)...... …7 7. Thallus instantly K+ intense lemon yellow, P+ intense orange (thamnolic acid present)...... …Key 6 (Page 13) 7. Thallus K-, K+ other colors, or K+ yellow but not usually intense, P+ or P- (without thamnolic acid) ...... …8 8. Thallus AND/OR soralia/medulla C- AND UV+ blue-white (alectoronic acid, confluentic acid, divaricatic acid, perlatolic acid, etc. present)...... …Key 7 (Page 14) 8. Thallus AND/OR soredia/medulla not C- AND UV+ blue-white...... …9 9. Thallus C+ AND/OR KC+ pink-red (erythrin, gyrophoric acid, lecanoric acid present)...... …...... Key 8 (Page 16) 9. Thallus NOT C+ AND/OR KC+ pink-red...... 10 10. Thallus AND/OR soralia/medulla K- AND C+ yellow-orange OR KC+ yellow or KC+ yellow-orange (xanthones or usnic acid present), usually distinctly greenish or yellowish in color…...... Key 9 (Page 19) 10. Thallus AND/OR soralia/medulla K+ or K-, C-, KC- (or KC reaction not different from K reaction)…...... 11 11. Thallus distinctly green, often with a white prothallus; isidiate or lobulate; southeastern North America (Phyllopsora)…...... Key 10 (Page 21) 11. Thallus not with the above combination of characters; from anywhere in North America… …………………………………………………………………..…………………………….12 12. Thallus K- or K+ yellowish AND P+ yellow, P+ orange-red, or P+ red (argopsin, fumarprotocetraric acid, pannarin, protocetraric acid, psoromic, or stictic acid present) …… ………………………………………………………………………...... …Key 11 (Page 23) 12. Thallus K+/-, P-, UV- (atranorin, and species that do not react to spot tests)...... …Key 12 (Page 26) 1. Thallus entirely leprose and lacking a prothallus (Lepraria s.l.)...... …Key 13 (Page 30)

1 KEYS TO YELLOW, YELLOW-GREEN, AND ORANGE STERILE CRUSTOSE LICHENS

Note: This key does not include species with usnic acid which would be classified as “green”. It does include some xanthone containing species that are often very yellow. These species are also keyed out in the xanthone/usnic acid key.

GENERIC KEY

1. Thallus K+ violet/magenta, anthraquinone pigments present in the cortex...... …Caloplaca/Fulgensia 1. Thallus K- or K+ some other color than above, anthraquinone pigments absent from the cortex...... …2 2. Growing on soil in arctic/boreal habitats...... …3 3. Thallus UV+ bright orange (rhizocarpic acid)...... Arthrorhaphis citrinella (Ach.) Poelt 3. Thallus UV- or UV+ dull orange (calycin?)…...... Candelariella “granuliformis” Westberg 2. Not growing on soil in arctic/boreal habitats...... …4 4. Thallus rusty orange; medulla UV+ blue-white (confluentic acid); on rocks in boreal habitats...... …Porpidia melinodes (Körber) Gowan & Ahti (fig. 16D) 4. Thallus distinctly yellow or yellow-green; medulla not UV+ blue-white (confluentic acid absent); ecology various...... …5 5. Corticolous; in the coastal plain of the southeastern USA; soralia punctiform; thallus of thin to indistinct flat green areoles ...Buellia wheeleri R.C. Harris (fig. 20B) 5. Primarily on wood; not in the coastal plain of the southeastern USA; soralia and thallus not as above...... 6 6. Xanthones present; thallus C+/KC+ orange...... …7 7. On driftwood in maritime habitats of the arctic/boreal regions; thallus areolate with discrete soralia...... ……………………………………………………………...…Lecanora orae-frigidae R. Sant. (fig. 20B) 7. On other substrates in other habitats; thallus granulose/leprose...... …8 8. Thallus leprose, composed of ecorticate granules, rare in North America?...... Lecanora expallens Ach. (fig. 20A) 8. Thallus granulose, composed of corticate granules, common in western North America...... …Pyrrhospora quernea (Dicks.) Körber (fig. 19E-F) 6. Xanthones absent; thallus not C+/KC+ orange; distribution various...... …9 9. Thallus entirely leprose...... …see key to Chrysothrix 9. Thallus areolate, at least initially…...... 10 10. On bark or wood...... …11 11. Thallus composed of corticate granules, 50-100 µm in diameter...... …...... Candelariella xanthostigma (Pers.) Lettau (fig. 19A) 11. Thallus composed of soredia arising from the breakdown of areoles…...... 12 12. Asci 8-spored; widespread except in the Great Lakes-Appalachians...... …Candelariella xanthostigmoides (Müll. Arg.) R.W. Rogers (fig. 19B) 12. Asci polysporous; distribution various…...... 13 13. Soredia small, 20-40-(50) µm in diameter...Candelariella efflorescens R.C. Harris & Buck 13. Soredia large, 50-100 µm in diameter...... …Candelariella xanthostigma (fig. 19A) 10. On non-calcareous rock…...... 14

2 14. Thallus P+ orange-red (pannarin present w/ zeorin and rhizocarpic acid)...... …Lecanora subaurea Zahlbr. (fig. 20D) 14. Thallus P- (pannarin absent)…...... 15 15. Soralia excavate and crater-forming...... …Lecanora reagens Norman 15. Soralia not as above…...... Lecanora epanora (Ach.) Ach. (fig. 20C)

KEY TO SOREDIATE CALOPLACA SPECIES

Only species of Caloplaca with yellow/orange thalli are keyed out here, the color of the apothecia does not matter. This includes the species in which the either whole thallus is yellow/orange as well as those in which only the lichenized diaspores are yellow/orange. Specifically this excludes species in which the entire thallus (i.e., including the diaspores) is gray. Note that some species with gray thalli have yellow/orange apothecia. These are keyed out separately in Key 12 because they are frequently sterile. 1. On calcareous rocks or soil/humus derived from calcareous rocks...... …...2 2. Interior deserts or arctic; thallus often densely pruinose; discrete soralia absent; schizidia or “warts” present; ascospores simple or 2-celled and not polarilocular...... …...... …3 3. Thallus without distinct marginal lobes; ascospores simple...... Fulgensia bracteata (Hoffm.) Räsänen 3. Thallus with distinct marginal lobes; ascospores various...... 4 4. Thallus “warted”; ascospores 2-celled...... …Fulgensia deserotrum (Tomin) Poelt 4. Thallus schizidiate; ascospores simple...... ….....…Fulgensia subbracteata (Nyl.) Poelt 2. Distribution various; thallus not typically densely pruinose; discrete soralia present in most species; ascospores polarilocular...... 5 5. Thallus saxicolous (distributions various)...... …...6 6. Thallus lobed at margin, orange, forming rosettes...... …...…7 7. Granulose isidia present, soralia absent …Caloplaca verruculifera (Vain.) Zahlbr. (fig. 21E-F) 7. Soralia present, granulose isidia absent...... …8 8. Soralia terminal on short lobes or marginal along the margins of the areoles; typical of exposed sites...... Caloplaca decipiens (Arn.) Blomb. & Forss. (fig. 21A-B) 8. Soralia laminal; typical of shaded sites...... …9 9. Lobe tips white pruinose; thallus forming small rosettes; soralia excavate, lighter than thallus...... …………………..... Caloplaca cirrochroa (Ach.) Th. Fr. (fig. 21C-D) 9. Lobe tips not white pruinose; thallus a thin areolate +/- continuous crust; soralia +/- excavate, concolorous with thallus…...... Caloplaca obliterans (Nyl.) Blomb. & Forss. 6. Thallus not lobed at margins...... ….10 10. Thallus leprose, pale yellow-greenish; typical of shaded overhangs and humid bluffs...... …………………………...... Caloplaca chrysodeta (Vainio ex Räsänen) Dombr. (fig. 20E) 10. Thallus areolate (at least initially), color otherwise; habitats otherwise...... 11 11. Thallus brownish/greenish-orange, isidiate/isidio-sorediate; northern...... ….Caloplaca xanthostigmoidea (Räsänen) Zahlbr. 11. Thallus yellow to orange, sorediate; distribution various...... …...... 12 12. Thallus orange to red-orange, soralia excavate, laminal; typical of shaded sites……… ……...... …Caloplaca obliterans (Nyl.) Blomb. & Forss. 12. Thallus yellow to yellow-orange, soralia not excavate or laminal; typical of exposed sites...... 13 13. Thallus +/- squamulose-areolate; soralia marginal; ascospores 10-12 µm long...... ….Caloplaca flavocitrina (Nyl.) H. Olivier (fig. 22F) 13. Thallus areolate, often dissolved almost completely into soredia; soralia irregular, spreading to dissolve the areoles; ascospores 11-14 µm long………………………… ...... ….Caloplaca citrina (Hoffm.) Th. Fr. 3 5. Muscicolous/terricolous (all northern boreal)...... 14 14. Thallus isidiate to isidio-sorediate, brown/greenish-orange; common...... Caloplaca xanthostigmoidea 14. Thallus sorediate, yellow-orange; less common...... …...... ….15 15. Thallus areolate/squamulose; soralia marginal; terricolous……………………………………... …………………………………………………...... Caloplaca tominii (Savicz) Ahlner (fig. 20F) 15. Thallus areolate, dissolving almost entirely into soredia; soralia irregular, spreading to dissolve the areoles; muscicolous/lignicolous...... Caloplaca citrina (Hoffm.) Th. Fr. 1. On bark, non-calcareous rocks, or humus/soil derived from non-calcareous rocks...... …...... 16 16. On bark or wood...... 17 17. Thallus well developed and distinct...... ….18 18. Extreme northern Alaska or elsewhere in the high arctic; thallus composed of convex areoles; soralia laminal and expanding to dissolve the areoles...... Caloplaca alaskensis Wetmore 18. Not from the extreme high arctic; thallus various...... …....19 19. Thallus distinctly areolate to subsquamulose; typically on wood...... ….....20 20. Thallus yellow-orange, areolate....group of problematical taxa included in C. citrina s.l. (see Wetmore, Bryologist, 107(4): 510. 2004). 20. Thallus orange-red, areolate to subsquamulose...... Calopalca microphyllina (Tuck.) Hasse (fig. 22A-B) 19. Thallus continuous; typically on bark...... 21 21. Southwestern North America; soredia pale, cream colored; thallus pruinose...... …………………………………………………...... Caloplaca persimilis Wetmore 21. Eastern North America/Great Plains; soredia yellow-orange, not pale; thallus...... epruinose...... Caloplaca chrysophthalma Degel. (fig. 22E) 17. Thallus not well developed, immersed or appearing as a thin continuous film...... 22 22. Boreal/arctic regions; thallus a thin continuous film; soredia yellow, coarse-granular and not arising in distinct soralia...... …..Caloplaca xanthostigmoides 22. Southeastern coastal plain; thallus immersed/indistinct; soredia orange, arising in discrete soralia...... Caloplaca granularis (Mull. Arg.) Zahlbr. (fig. 22C-D) 16. On non-calcareous rocks, or humus/soil derived from non-calcareous rocks...... ….23 23. Thallus with distinct marginal lobes……………………………………………...... 24 24. Known from Mexico and the West Indies, not yet from North America; soralia laminal, excavate, crateriform/punctiform...... [Caloplaca cupulifera (Vainio) Zahlbr.] 24. Known from North America; soralia marginal, not excavate and cupuliform...... 25 25. Thallus large and with distinct well developed marginal lobes, large enough to be confused with a Xanthoria; throughout western North America...... …Caloplaca decipiens 25. Thallus smaller, marginal lobes poorly developed, obviously not a Xanthoria; restricted to coastal western North America...... Caloplaca stellata Wetmore & Karnef. 23. Thallus without distinct marginal lobes...... Caloplaca flavocitrina (fig. 22F)

KEY TO SOREDIATE CANDELARIELLA SPECIES

Keyed out in the general key above. There are two common sorediate species, and a common third granulose species. There is also an undescribed species on organic matter in arctic/boreal habitats. An additional key from Lendemer and Westberg (Opuscula Philolichenum, 8: 75-81. 2010.) is reproduced below as it may be helpful.

1. Thallus foliose, although sometimes mostly dissolved into soredia with only a few intact lobes 4 remaining...... 2 2. Asci polysporous; soredia produced marginally; widespread throughout the continent……………………… …………………………………………………………………...... Candelaria concolor (Dicks.) Stein 2. Asci 8-spored; soredia produced on the lower surface of the thallus; coastal southwest with occasional inland localities...... …. Candelaria pacifica M. Westb. & Arup (in press) 1. Thallus crustose...... 3 3. On mosses and plant remains on the ground in arctic-alpine habitats…………………………………………... ………………………………………………………………….....Candelariella granuliformis M. Westb. ined. 3. On bark or wood in other habitats...... …....4 4. Asci 8-spored; widespread except in the Great Lakes-Appalachian Mountains...... ……………………………...... …...Candelariella xanthostigmoides (fig. 19B) 4. Asci polysporous; distribution various...... 5 5. Soredia small, 20-40-(50) µm in diameter, true soredia arising from the breakdown of thalline areoles; Appalachian-Great Lakes...... ….Candelariella efflorescens 5. Soredia large, 50-100 µm in diameter, “soredia” actually granules that form the primary thalline units; widespread...... Candelarella xanthostigma (fig. 19A)

KEY TO CHRYSOTHRIX IN EASTERN/CENTRAL NORTH AMERICA

This key is taken from Harris and Ladd (Opuscula Philolichenum, 5: 29-42. 2008) with additions and corrections. This key SHOULD NOT be used for western North America. The genus is badly in need of revision in that region.

1. Photobiont chlorococcoid, the cells + isodiametric; thallus chemistry various…………...... 2 2. On bark or wood……………………………………………………………………………………………..…3 3. Granules small, 15-14 µm in diameter; calycin or leprapinic? acid or pinastric acid or rhizocarpic acid as the major substance; eastern North America.……………………………………………...... 4 4. Thallus bright yellow, UV- (leprapinic? acid or pinastic acid or calycin); granules 25-45 µm; on hardwoods or conifers………………………………………………………………………………...5 5. Major substance pinastric or leprapinic? acid (K-, KC-); on trees and wood……………………6 6. Pinastric acid major; widespread……….…Chrysothrix xanthina (Vainio) Kalb (fig. 19C) 6. Leprapinic? acid major; specimens on Quercus and palm, southern Coastal Plain or on Abies in Maine/Michigan………………………………………...Chrysothrix insulizans s.l. 5. Major substance calycin (K+ slowly reddish); typically on Quercus; Alabama, Florida, Georgia ……...... …………………………………………………………………………..Chrysothrix sp. 4. Thallus dull pale yellow to pale greenish-yellow, UV+ dull to bright orange (rhizocarpic acid); granules small, 15-25-(30) µm in diameter, loose and without binding hyphae; only on conifers………………………………………………….Chrysothrix chamaecyparicola Lendemer 3. Granules coarse, 35-80 µm in diameter; calycin and/or pinastric acid; not seen from eastern North America……………………………………………………...Chrysothrix candelaris (L.) J.R. Laundon 2. On rock, or humus/organic matter associated with rock………………………………………………………..7 7. On organic matter/humus associated with rock; thallus C+ pink, gyrophoric acid present, typically dull greenish-yellow to pale-yellow; very rare?...... Chrysothrix susquehannensis Lendemer & Elix 7. On rock; thallus C-, gyrophoric acid absent, not as above………………………………………………...8 8. On acidic siliceous rock, thallus K- to K+ slowly reddish……………………………………………9 9. Thallus thick, attached to the rock by rhizohyphae (these sometimes not evident); usually easily separated from rock……………………………………………………………………………...10

5 10. Thallus UV+ dull orange (leprapinic? acid), of loosely aggregated granules with numerous projecting hyphae, bicolor in section, yellow above, whitish to brown below (figure 8); Ozarks, Pennsylvania, SE USA…..Chrysothrix onokoensis (Wolle) R.C. Harris & Ladd 10. Thallus UV- (calycin, vulpinic acid, zeorin) usually of more tightly compacted granules, mostly without obvious projecting hyphae, + uniform yellow in section; Ontario, Vermont………………………………………Chrysothrix chlorina (Ach.) J.R. Laundon 9. Thallus thin, lacking rhizohyphae, of scattered to contiguous granules or leprose; rarely easily separable from the rock………………………………………………………………………….11 11. Thallus K- (pinastric or rhizocarpic acid), forming a thin, + continuous, granular crust...12 12. Thallus bright yellow to bright greenish yellow, UV- (pinastric acid); normally on bark, rarely on rock……………………………...……………Chrysothrix xanthina 12. Thallus pale yellow to pale greenish yellow, UV+ dull to bright orange (rhizocarpic acid); normally on rock, often in heavily shaded crevices of fieldstone walls, rarely on wood or bark ………………………………Psilolechia lucida (Ach.) M. Choisy 11. Thallus K+ slowly reddish (calycic + leprapinic? acid), forming small + round patches on rock, sometimes forming large continuous, rimose or rimose-areolate patches with rounded soralium-like outliers ……………Chrysothrix insulizans R.C. Harris & Ladd 8. On calcareous rock; thallus K+ instantly magenta-purple (parietin)……………...... …………………………………………………………………Caloplaca chrysodeta (Vainio) Dombr. 1. Photobiont Stichococcus, the cells subrectangular to elongate; thallus containing vulpinic acid...... ………………………………………………………………………….…sterile Chaenotheca furfuracea (L.) Tibell

6 KEY TO LICHEXANTHONE CONTAINING SPECIES

Note: In some species, particularly Pertusaria species, the lichexanthone is distributed unevenly across the thallus. Thus sometimes parts of the thallus will react UV+ while others will be UV-, in these cases often the UV+ regions are concentrated around the soralia. Some of the species keyed out here also produce other UV+ substances (e.g., 2-0-methylperlatolic acid or haemathamnolic acid). The UV+ blue-white reactions of these substances are not always obvious because they can be masked by the reaction of lichexanthone.

1. Saxicolous; SW (AZ/Mexico, disjunct in Channel Islands); soralia K+ dirty yellow-brown (haemathamnolic acid)...... …...... Pertusaria moreliensis de Lesd. 1. Corticolous; distribution various; chemistry various...... …2 2. Photobiont Trentepholia; tropical southern Florida; not sorediate or isidiate, but rather with fluffy mounds of ascomata that can be sterile and are frequently UV+ blue-white (2-0-methylperlatolic acid)...... …………………………………………………...... …Cryptothecia evergladensis Seavey (fig. 1E) 2. Photobiont not Trentepholia; not with the above combination of characters...... …...... 3 3. Medulla or soralia C+ pink/red (lecanoric acid present)...... …4 4. Apothecia always abundant and obvious; soralia sparse and scattered, poorly developed...... …...... unreported sorediate morphotype of Ochrolechia africana Vain. (fig. 2E-F) 4. Apothecia rare or absent; soralia abundant, well developed...... …………….…5 5. Soredia fine, arising from true soralia; soralia convex to +/- excavate; north-temperate...... Ochrolechia arborea (Kreyer) Almb. (fig. 1A-B) 5. Soredia coarse, actually pruina on discoid apothecia; soralia absent; tropical SE with disjunctions in S Appalachians and Canadian Maritimes……………………………...... Pertusaria velata (Turner) Nyl. lichexanthone chemotype (syn. P. pulchella Malme) 3. Medulla and soralia C- (lecanoric acid absent); all in the SE USA…………...... 6 6. Soralia discrete not coarsely pustular; zeorin absent...... …7 7. Very rare?, known only from Apalachicola; soralia K+ yellow (atranorin)...... ……………………………………...... ….Haematomma leprarioides (Vain.) Vain. 7. More common and widespread distributed in the SE; soralia K+ or K-...... 8 8. Soralia not discrete, coarsely pustular; zeorin present………………...... ………………...... Megalospora pachycheila (Tuck.) Sipman (fig. 1C-D) 8. Soralia discrete, not pustular; zeorin absent ...... …9 9. Soralia bitter tasting, P-, K-, KC+ fleeting purple-red in section (picrolichenic acid present); rare, known only from Florida...... ….Pertusaria ventosa Malme (fig. 2B) 9. Soralia not bitter tasting, P+, K+/-, KC+/- (picrolichenic acid absent); more common and widespread in the SE...... 10 10. Soralia K-, P+ yellow in section (baeomycesic acid present)...... …………………………...... …..Pertusaria floridana Dibben (fig. 2C-D) 10. Soralia K+ yellow, dirty-brown, or red-purple (thamnolic, haemothamnolic, hypothamnolic, or stictic acid present)...... …..11 11. Soralia K+ deep red-purple (note, use only a small amount of reagent, and reaction is fast); hypothamnolic acid…………………...... ………………………………...... Pertusaria hypothamnolica Dibben (fig. 1E) 11. Soralia K+ strong yellow or dirty yellow-brown...... …..12

7 12. Soralia K+ strong yellow (thamnolic acid)...... ………...... Pertusaria trachythallina Erichsen lichexanthone chemotype 12. Soralia K+ dirty yellow-brown...... 13 13. Common; haemathamnolic acid present...... Pertusaria commutata Müll. Arg. (syn. P. copiosa Dibben) (fig. 2A) 13. Very rare, known only from Marion Co. FL; stictic acid present...... ………………………………...... P. sp. Buck 16889 of Harris 1995

8 KEY TO SPECIES WITH TRENTEPHOLIA PHOTOBIONT

1. Thallus with sporodochia/synnema not soralia, and conidia not soredia...... …2 2. Conidia muriform; sporodochia stalked (e.g., synnema), though occasional sessile forms occur; lecanoric acid absent; north temperate with occasional occurrences in the SE on the roots of old growth Fagus grandifolia...... … Dictyocatenulata alba Finley & E.F. Morris (fig. 4A) 2. Conidia transversely septate or muriform; sporodochia not stalked; lecanoric acid present (thallus C+ OR C-); tropical SE………………………………………………………………………………………...... 3 3. Sporodochia white; conidia 1-2-(4) celled; tropical SE; thallus C+ red...... …………………………………………………………………...…Tylophoron protrudens Nyl. (fig. 6C-D) 3. Sporodochia brown; conidia simple; extreme SE (Collier Co., FL); thallus C-...... …Tylophoron moderatum Nyl. (fig. 6E-F) 1. Thallus with soredia or isidia, not sporodochia ……………………………………………………………………..4 4. Thallus isidiate……………………………………………………………………………………………….…5 5. Thallus P+ yellow or orange (stictic satellites, salazinic, or psoromic acid present)…...... 6 6. Isidia crusted with red pigment; extreme tropical Florida...... ………………………………………………………Herpothallon rubroechinatum Frisch & G. Thor 6. Isidia not crusted with red pigment; distribution various...... …7 7. Thallus P+ orange (stictic satellites or salazinic acid)...... …8 8. Thallus K+ yellow turning red (salazinic acid)...... …Diorygma antillarum (Vain.) ined. 8. Thallus K+ dirty yellowish (stictic satellites)…...... Heiomasia sipmanii ined. 7. Thallus P+ yellow (psoromic acid)...... …9 9. Thallus fluffy or cottony…….…Herpothallon echinatum Aptroot, Lücking & Will-Wolf 9. Thallus compact, not fluffy and cottony...... …10 10. Isidia abundant, long and branching; ascospores brown, 2-celled; psoromic acid only……………………………………...... Thelotrema eximium R.C. Harris (fig. 5D-E) 10. Isidia sparse, short and globose; ascospores hyaline, submuriform; psoromic acid + unknown accessory……………..Thelotrema halei (Tuck. & Mont.) Zahlbr. (fig. 6A-B) 5. Thallus P- (without the above substances)…...... 11 11. Thallus fluffy or cottony, red pigmented, UV+ blue-white (confluentic acid)...... …...…………………..…Herpothallon rubrocinctum (Ehrh.) Aptroot, Lücking & G. Thor (fig. 5F) 11. Thallus compact, not fluffy, not red pigmented, UV-...... …12 12. Thallus dull, blue-gray; isidia thick and coarse; medulla inspersed with large crystals...... …………………………………………………...…Reimnitzia santense (Tuck.) Kalb (fig. 5B-C) 12. Thallus shiny or dull, brown to orange-brown or yellow-brown; isidia small and fragile; medulla not inspersed …………………………………………………………………………..…13 13. Thallus thick and shiny, yellow-brown, with distinct well developed prothallus; rare except in tropical SE……………………...... Clathroporina isidiifera R.C. Harris (fig. 4E-F) 13. Thallus thin and dull, brown to orange-brown...... …14 14. Common throughout SE; thallus yellowish- or orange-brown, isidia usually short and fragile; perithecia pale, wall orange-brown.....…Porina scabrida R.C. Harris (fig. 4C-D) 14. Rare?, known only from Illinois; thallus brown; isidia long and branched; perithecia dark, wall purple-black……...... …Pseudosagedia isidiata (R.C. Harris) R.C. Harris

9 4. Thallus sorediate…………………………………………...... 15 15. Coastal S/C California; C+ red (erythrin or lecanoric acid)...... ….16 16. Saxicolous; erythrin major………………...... …Dirina catalinariae f. sorediata Tehler 16. Corticolous; lecanoric acid major……....Schismatomma pluriloculare (Zahlbr.) Zahlbr. (fig. 6A) 15. Not coastal S/C California; C+/-…………….…...... 17 17. C+ red/pink (erythrin or gyrophoric acid present)…...... 18 18. Corticolous……………………….…...... 19 19. Thallus blue-white; extreme tropical Florida...... ………………………………………Cryptothecia punctosorediata Sparrius & Saipunkaew 19. Thallus thin, immersed, brown; Appalachian…...... Opegrapha sp. nov. 18. Saxicolous... 20 20. Erythrin major; Appalachian/Ozarks; soralia small and punctiform...... ……………………………………………Dirina massiliensis f. sorediata Müll. Arg. Tehler 20. Gyrophoric acid major; northern temperate; soralia irregular, arising from cracks in the thallus………………………………...... Opegrapha gyrocarpa Flot. (fig. 3B) 17. C- (erythrin or gyrophoric acid absent)…...... 21 21. Thallus P+ yellow or orange-red (psoromic, stictic, or protocetraric acid present)…...... 22 22. Saxicolous; P+ yellow (psoromic acid)…...... Opegrapha moroziana Lendemer (fig. 3C) 22. Corticolous, P+ orange or red (stictic or protocetraric acid)...... …23 23. Common and widespread; soralia punctiform with gray soredia; P+ orange (stictic acid)…………………………………...…Nadvornikia sorediata R.C. Harris (fig. 3E-F) 23. Rare, tropical SE; soralia not punctiform, with white soredia contrasting distinctly with the thallus; P+ red (protocetraric acid). …Myriotrema erodens R.C. Harris (fig. 4B) 21. Thallus P- (those substances absent)…...... 23 24. Saxicolous; UV+ blue-white (confluentic acid)...... …Opegrapha zonata Körber (fig. 3D) 24. Corticolous; UV-, no substances…....Opegrapha corticola Coppins & P. James (fig. 3A)

10 KEY TO HYPOCENOMYCE SPECIES

1. Thallus C+ red (lecanoric acid)...... … Hypocenomyce scalaris (Ach.) M. Choisy 1. Thallus C-……………………………………………………………………………………………………………2 2. Thallus K+ strong yellow (thamnolic acid)...... …Hypocenomyce sierrae Timdal 2. Thallus K- (thamnolic acid absent)...... …3 3. Thallus P+ orange-red (fumarprotocetraric acid)……………………………………………………………... ……………………………………………Hypocenomyce anthracophila (Nyl.) P. James & Gotth. Schneid. 3. Thallus P-...... …4 4. Squamules large; soralia excavate, in hoods…...... ………..Waynea californica Moberg (fig. 29E) 4. Squamules smaller; soralia not excavate and in hoods..Hypocenomyce castaneocinerea (Räsänen) Timdal

11 KEY TO SPECIES WITH NORSTICTIC ACID

1. Thallus with coralloid isidiate; coastal western North America…...... Pertusaria coccodes (Ach.) Nyl. (fig. 8E-F) 1. Thallus sorediate or with small globose isidia that are easily detached; distribution various...... …2 2. Saxicolous………………………………...... 3 3. Thallus entirely dissolving into irregular granulose-isidioid structures or soredia; primarily western....…4 4. Thallus blue-gray, dissolving into soredia; conspicuous white prothallus usually present; in humid microhabitats...…Phlyctis speirea G. Merr. (saxicolous populations that are common in western North America but very rare in the east) (fig. 8A) 4. Thallus brown or lead-gray, dissolving into irregular granulose-isidioid structures; without a white prothallus; not restricted to humid microhabitats; +/- arid western/central North America...... ………………………………………………………….… Aspicilia cf. simoensis Räsänen (fig. 7A-B) 3. Thallus remaining intact, with propagules more-or-less restricted to certain areas; eastern/south- central/northern-boreal………………...... 5 5. Thallus corticate, greenish-brown; uncommon; northern-boreal with disjunctions in the Appalachians…………………...... Baeomyces carneus (Retz.) Flörke (fig. 7C-D) 5. Thallus ecorticate, bluish-white; very common; eastern North America, Ozarks and adjacent central US……………………………...... Phlyctis petraea R.C. Harris ined. (fig. 8A-B) 2. Corticolous………………………………...... …6 6. Thallus without discrete soralia, soredia arising from irregular breakdown of the thallus...... …………………………………………………...………………………Phlyctis speirea G. Merr. (fig. 8A) 6. Thallus with discrete soralia; soredia arising in soralia...... …7 7. Cortex UV+ bright orange (thiopanic acid); soralia hemispherical; arid SW North America (Arizona)………………………...... …Pertusaria azulensis de Lesd. (fig. 8D) 7. Cortex UV- (thiophanic acid absent); soralia various; northern/boreal...... …8 8. Soralia discoid, plane to convex; soredia white to pale-blue; thallus thick and well developed; atranorin absent………………...... Pertusaria waghornei Hulting (fig. 8C) 8. Soralia punctiform, +/- excavate; soredia usually dark; thallus thin not well developed; atranorin present………………...... …Buellia griseovirens (Turner & Borrer ex Sm.) Almb. (fig. 7E-F)

12 KEY TO SPECIES WITH THAMNOLIC ACID

1. Pseudopodetia present; thallus white; on humus or soil; high elevations of western North America...... …………………………………...... …Leprocaulon subalbicans (Nyl.) I.M. Lamb & A.E. Wade 1. Pseudopodetia absent; thallus, distribution, and ecology not as above…...... 2 2. Soralia discrete, circular, discoid…...... Pertusaria trachythallina Erichsen (fig. 9F) 2. Soralia irregular, not discrete and circular…...... 3 3. Schizida present; on hardwoods; temperate eastern North America extending S into the Coastal Plain…... ………………………………………………….…Variolaria pustulata (Brodo & Culb.) ined. (fig. 9C-D) 3. Soredia present; typically on conifers, rarely on hardwoods; distribution various…...... 4

4. Thallus squamulose, at least in part; red pycnidia often present; SE & Ozarks...... ……………………...... Cladonia ravenelii Tuck. (fig. 9A-B) 4. Thallus crustose; pycnidia not red, and not obvious; northern/boreal...... ………………………...... …Loxospora elatina (Ach.) A. Massal. (fig. 9E)

13 KEY TO UV+ BLUE-WHITE SPECIES

1. Thallus isidiate…...... 2 2. Thallus gray-brown; southeastern North America; alectoronic acid...... …………………………………………...... …Protoparmelia isidiata Diederich et al. (fig. 18B) 2. Thallus white; PNW; divaricatic acid (+ gyrophoric acid in the apothecia)...... …………………………………………...... …Loxosporopsis corallifera Brodo et al. (fig. 18A) 1. Thallus sorediate or blastidiate…………………...... 3 3. On rock…………………………………...... …4 4. Medulla K+ bubbles in squash mount (confluentic acid present); soralia +/- excavate…...... 5 5. Thallus orange…………………...... Porpidia melanodes (Korb.) Gowan & Ahti (fig. 16D) 5. Thallus white/gray………………...... …Porpidia tuberculosa (Sm.) Hertel & Knoph (fig. 16C) 4. Medulla K- in squash mount (divaricatic acid present); soralia convex, not excavate...... …6 6. Areoles thin and dispersed; arctic; rare…...... Fuscidea gothoburgensis (H. Magn.) V. Wirth & Vězda 6. Areoles thick, +/- continuous; arctic-boreal and/or Appalachian...... …7 7. Ascospores 0-1 septate, ellipsoid and frequently bent...... …………………………………...... … Fuscidea recensa (Stirton) Hertel et al. (fig. 16B) 7. Ascospores simple, globose-subglobose…...... Fuscidea appalachensis Fryday (fig. 16A) 3. On bark, humus, soil, or wood.…...... 8 8. Southeastern US/Ozarks……………...... …9 9. Thallus dirty-green, +/- granulose; restricted to bark of trees in very high humidity inland sites; apothecia pale brown, often present; lobaric acid…...... Scoliciosporum pensylvanicaum R.C. Harris (fig. 17E) 9. Thallus not dirty green, continuous or areolate; not restricted to the above habitats; apothecia less frequent or unknown; lobaric acid present or absent…...... 10 10. Appalachians and corticolous; lobaric acid present…...... Bacidia lobarica Tønsberg 10. Primarily Coastal Plain or Piedmont; lobaric acid absent...... …11

11. Thallus dark gray-brown; soralia large and capitate; alectoronic acid...... …………………………………...... … Protoparmelia capitata Lendemer (fig. 18B) 11. Thallus usually lighter; soralia smaller and not capitate; alectoronic acid absent...... …12 12. On hardwoods; thallus white; soralia irregular; sphaerophorin present...... ………………………………...... … Haematomma americanum Staiger & Kalb (fig. 18D) 12. On conifers or rotting wood; thallus distinctly light-gray; soralia +/- convex; perlatolic acid...... …………………………...... Hertelidea pseudobotryosa R.C. Harris et al. (fig. 18E-F) 8. Northern/boreal and/or western...... …13 13. Thalli forming +/- regular circular colonies with a distinct prothallus...... …………………………Ropalospora viridis (Tønsberg) Tønsberg & Fuscidea pusilla Tønsberg (fig. 17D) 13. Thalli not as above…………………...... 14 14. Thallus dirty-green, +/- granulose; lobaric acid present...... …15 15. Restricted to inland suites; ascospores not spirally arranged in the ascus; eastern North America...... ………………………………...... … Scoliciosporum pensylvanicaum R.C. Harris (fig. 17E) 15. Restricted to oceanic sites?; ascospores spirally arranged in the ascus; PNW and NE...... ………………………………...... …Scoliciosporum pruinosum (P. James) Vězda (fig. 17F) 14 14. Thallus not dirty-green and entirely granulose; with or without lobaric acid...... …16 16. Thallus blastidiate or not obviously sorediate, typically small; restricted to high humidity habitats, especially Thuja swamps; lobaric acid present…… Protoparmelia hypotremella Herk et al. (fig. 17A-B) 16. Thallus distinctly sorediate, usually larger and not so restricted; lobaric acid absent…...... 17 17. Thallus yellow, usnic acid present in addition to planaic acid and atranorin; arctic boreal...... ……………………… ...... … Mycoblastus alpinus (Fr.) Hellb. (fig. 16E) 17. Thallus white or gray/brown, usnic acid absent...... …18 18. Thallus gray/brown…...... 19 19. Thallus KC+ fleeting purple (lobaric acid); areoles dissolving into brown-green soredia; high elevations of PNW…...... Lecidea porphyrospoda (Anzi) Th. Fr. 19. Thallus KC- (lobaric acid absent); areoles +/- persisting; distribution various...... …20 20. Soredia dark; PWN; sphaerophorin…...... Rinodina disjuncta Sheard & Tønsberg 20. Soredia pale; arctic-boreal; perlatolic acid…...... ………………… ...... Mycoblastus caesius (Coppins & P. James) Tønsberg (fig. 16F) 18. Thallus white…………...... …21 22. On conifer bark; ascospores globose; divaricatic acid…...... ………………………...... Lecidea nylanderi (Anzi) Th. Fr. (fig. 17C) 22. On rotting logs of conifer bark; ascospores ellipsoid; perlatolic acid...... …………………………...... … Hertelidea botryosa (Fr.) Printzen & Kantvilas

15 KEY TO SPECIES WITH C+ AND/OR KC+ PINK-RED THALLUS

Note: In some regions Ochrolechia androgyna (Hoffm.) Arn. can occur on rocks. Although this phenomenon is usually rare, O. androgyna was found to commonly occur on rocks in oceanic Maine.

1. Typically on rock or soil (not humus)………………………………………………...... 2 2. Thallus C+ and KC+ pink/red (erythrin or gyrophoric or lecanoric acid present)…...... 3 3. Thallus with cephalodia; medulla often UV+ blue-white (confluentic acid); arctic...... ……………………………………………………. Amygdalaria panaeola (Ach.) Hertel & Brodo (fig. 13B-C) 3. Thallus lacking cephalodia; medulla without confluentic acid; not arctic…...... 4 4. Thallus thick, ecorticate, byssoid and distinctly green in the field; Channel Islands and coastal Central California; erythrin present……………… .....…Lecanographa insolita Lendemer & K. Knudsen (fig. 13F) 4. Thallus not as above; distribution various; erythrin lacking…………………...... …5 5. Thallus isidiate; eastern North America…………….. ...…Ochrolechia yasudae (Vain.) Oshio (fig. 10A) 5. Thallus sorediate or granulose-isidiate; distribution various………………...... 6 6. Thallus continuous, placodioid, with the soralia arising from cracks in the thallus...... ………………..…..Trapelia placodioides Coppins & P. James (common in E, rare in W) (fig. 11E-F) 6. Thallus not continuous and placodioid, soralia discrete…………………...... …7 7. Thallus squamulose; on soil; arid northwestern NA……………………...... …………………………………………………...…Trapelia steppica McCune & Camacho (fig. 11C) 7. Thallus crustose/areolate; substrate and distribution various…………...... 8 8. Thallus brown, gray or white, bullate-areolate; distribution various...... …9 9. On maritime rocks in PNW; fatty acids present; with a distinct brown prothallus...... ………………………………………….… Ropalospora hibernica (P. James & Poelt) Tønsberg 9. Not on maritime rocks in PNW; fatty acids absent; without a distinct brown prothallus…...... 10 10. On rock; areoles widely dispersed, light brown or white-pruinose...... …………………………………………………………..…Trapelia obtegens (Th. Fr.) Hertel ? 10. Typically on soil; areoles not widely dispersed, not white-pruinose...... …..11 11. Thallus lead-gray, soralia typically darker than the thallus; rarely on soil or rock...... ………………………………. …Trapeliopsis flexuosa (Fr.) Coppins & P. James (fig. 11A) 11. Thallus brown or light gray, soralia pale or dark; frequently on soil...... …12 12. Thallus and soralia with irregular brown-orange pigmented patches, but the pigment K-; soredia pale…………………….. ..…Trapeliopsis granulosa (Hoffm.) Lumbsch (fig. 11B) 12. Thallus and soralia without pigmented patches; soredia dark; western North America…. ……………………………..…...Trapeliopsis bisorediata McCune & Camacho (fig. 11D) 8. Thallus green, immersed/indistinct; eastern North America……...... ……….……………….…Biatora chrysantha (Tuck.) Zahlbr. ? (unusual saxicolous populations) 2. Thallus C-, KC+ red (alectorialic acid present) …………………………………………………………………13 13. High elevations of southwestern North America (Arizona); atranorin and thamnolic acid present; thallus distinctly brown………………………………………………………... Lecanora cavicola Creveld (fig. 13A-B) 13. Appalachians to Texas; alectorialic acid only; thallus gray-white………………...... …Fuscidea praeruptorum (Du Rietz & H. Magn.) V. Wirth & Vězda 1. Typically on bark, humus, bryophytes, or lignum……………………………………...... 14 14. Thallus C-, KC+ red (alectorialic acid present); boreal region and high elevations of the Appalachians; on wood and conifers…………………………………………...... … Pycnora sorophora (Vain.) Hafellner (fig. 15E) 16 14. Thallus C+ and KC+ pink/red (gyrophoric or lecanoric acid present); distribution and substrate various….....15 15. Thallus isidiate; typically on humus or wet rotting logs…………………………...... ……………..………………………….. ...…Placynthiella icmalea (Ach.) Coppins & P. James (fig. 15B & D) 15. Thallus sorediate; on various substrates…………………………………………...... …………..…16 16. Thallus UV+ yellow (lichexanthone), at least patchily………………………...... ………………………………………………….………….Ochrolechia arborea (Kreyer) Almb. (fig. 1A-B) 16. Thallus UV- (lichexanthone lacking)…………………………………………...... …17 17. Thallus creamy white (Ochrolechia species)………………………………...... …18 18. Tropical SE……………………………………………....…Ochrolechia antillarum Brodo (fig. 10D) 18. Not in the tropical SE……………………………………………………...... …19 19. Thallus granular-isidiate; variolaric acid present; arid SW……………...... …………………………………………………………… Ochrolechia subisidiata Brodo (fig. 10B-C) 19. Thallus sorediate; variolaric acid absent; boreal/northern extending into the mountains of the SW.... …………………………………………………… Ochrolechia androgyna (Hoffm.) Arn. (fig. 10E-F) 17. Thallus not creamy white (not Ochrolechia)………………………………...... …20 20. Soralia irregular and thallus more-or-less leprose; PNW…………………...... …21 21. In ruderal habitats; thallus dark green and scurfy………………………...... ………………….………………………….....Scoliciosporum sarothamni (Vain.) Vězda (fig. 14A-B) 21. Not in ruderal habitats; thallus not dark green and scrufy……………..…...... 22

22. 5-0-methylhiascic acid major in addition to gyrophoric acid……....………………………………… …………………………………………………………...… Biatora aureolepra T. Sprib. & Tønsberg 22. 5-0-methylhiascic acid absent/minor………………...… Biatora vacciniicola (Tønsberg) Printzen 20. Not as above; distribution various……………………………………..…...... 23 23. Thallus and soralia distinctly light or dark green NOT lead-gray, gray, or brown; areoles usually minute and +/- immersed or otherwise indistinct…………………………...... …24 24. Thallus P+ red (pannarin present)…………………….…..Biatora printzenii Tønsberg (fig. 14F) 24. Thallus P- (pannarin absent)………………………………………...... 25 25. On humus and rotting logs………………………………………...... 26 26. Thallus usually dark brown contrasting strongly with the soralia...... ……………………………………..…Placynthiella dasaea (Stirton) Tønsberg (fig. 15A & C) 26. Thallus green, contrasting or not with the soralia………………...... 27 27. Soralia small, discrete; areoles dispersed and minute………………………………………. ……………………………………….…Trapelia corticola Coppins & P. James (fig. 12E-F) 27. Soralia not small and discrete; areoles forming a well developed continuous crust...... …………………….…Trapeliopsis viridescens (Schrad.) Coppins & P. James (fig. 12C-D) 25. On bark or over mosses………………………………………………...... 28 28. Soralia very discrete; corticolous; apothecial sections C+ red; southern Appalachians...... …………………………………..… Biatora appalachensis Printzen & Tønsberg (fig. 14C-D) 28. Soralia more irregular; corticolous or over mosses; apothecial sections C+ or C-; Great Lakes/Northern Appalachians or PWN…………………………………………………………29 29. Apothecial sections C-; Great-Lakes/Northern Appalachians...... …………………………………………….….Biatora chrysantha (Tuck.) Zahlbr. (fig. 14E) 29. Apothecial sections C+; PNW…………….Biatora chrysanthoides Printzen & Tønsberg 17 23. Thallus gray or brown; areoles distinct and obvious……………………...... 30 30. Not sorediate; high elevations of AZ; on conifer wood and bark; apothecia required for ID…………………………………………………………Lecidea trapelioides Printzen (fig. 13E) 30. Sorediate; distribution various; substrate various…………………………………...... 31 31. Thallus and soralia with orange-red pigmented patches, the pigment K+ purple; humid PNW ………………………………Trapeliopsis pseudogranulosa Coppins & P. James (fig. 12A-B) 31. Thallus and soralia without K+ pigment patches……………………...... 32 32. Thallus squamulose and large; arid NW……………………...... ………………………………………..Trapeliopsis steppica McCune & Camacho (fig. 11C) 32. Thallus areolate; distribution various………………………...... 33

33. Thallus P+ orange-red (argopsin present)………………………………………………... ………………………………………..Micarea leprosula (Th. Fr.) Coppins & A. Fletcher 33. Thallus P- (argopsin absent)…………………………………...... 34 34. Thallus brown, with punctiform soralia; PNW……………...... ………………………………...... Schaereria corticola Muhr & Tønsberg (fig. 15F) 34. Thallus lead-gray or light-gray, not brown; soralia not punctiform; distribution various………………………………………………………...... …35

35. Thallus lead-gray; soredia usually dark; lacking K- pigment patches...... …………………………...Trapeliopsis flexuosa (Fr.) Coppins & P. James (fig. 11A) 35. Thallus lighter gray; soralia usually pale; often with brown-orange pigment patches that are K-…………………….. Trapeliopsis granulosa (Hoffm.) Lumbsch (fig. 11B)

18 KEY TO SPECIES WITH USNIC OR XANTHONES

1. Thallus isidiate (both with xanthones)...... …2 2. Tropical SE...... Megalaria pannosa (Zahlbr.) Fryday & Lendemer (fig. 23D) 2. PNW...... Pertusaria flavocorallina Coppins & Muhr 1. Thallus sorediate………………...... …3 3. On rock………………………...... 4 4. Thallus C-, KC- (coronotone); zeorin present in addition to coronatone and unknown xanthone; PNW………. …………………………….… ...... Bacidia viridifarinosa Coppins & P. James 4. Thallus C+ and/or KC+ yellow-orange (usnic acid or different xanthones); distribution various...... …5 5. Thallus UV-, C- and KC+ gold (usnic acid without xanthones)...... …6 6. Thallus granulose sorediate, with a distinct white prothallus; arid SW; atranorin, zeorin, usnic acid...... ………………………...... …Lecanora brodoana Lumbsch 6. Thallus with discrete soralia that dissolve the thallus, without a white prothallus; northern/boreal; zeorin and usnic acid…………...... …Lecanora orothsea (Ach.) Ach. 5. Thallus UV+ dull or strong orange, C+/KC+ yellow-orange (other xanthones)...... …7 7. Thallus placodioid; schizidia present; coastal Baja California Mexico...... ……………………...... … Lecanora xanthonica B.D. Ryan & Poelt 7. Thallus areolate/granulose, not placodioid; soredia present; not in coastal Baja California…...... 8 8. Thallus dissolving into soredia; without discrete soralia...... ……………………...... … Lecidella meiococca (Nyl.) Hertel & Knoph 8. Thallus not dissolving into soredia; with discrete soralia...... ……………………...... … Lecidella scabra (Taylor) Hertel & Leuckert (fig. 23A) 3. On bark, wood, or humus...... …9 9. Thallus leprose, with a distinct fibrous prothallus; on hardwoods; Appalachian-Great Lakes...... ………………………………………………………………….………..…Lecanora thysanophora R.C. Harris 9. Thallus not leprose with a distinct fibrous prothallus; substrate and distribution various…...... 10 10. Appalachians/Great Lakes...... …11 11. Southern Appalachians; soralia discrete, convex, often yellowish and appearing pigmented; atranorin + xanthones………………...... …Vainionora americana Kalb et al. (fig. 23E) 11. More widespread; soralia irregular, appearing totally diffuse; atranorin absent...... …………………………...... … Biatora pontica Printzen & Tønsberg (fig. 23B) 10. Other geographic regions…...... 12 12. Thallus P+ orange (stictic acid present); tropical SE...... Pertusaria expolita R.C. Harris (fig. 23C) 12. Thallus P- (stictic acid absent); distribution various…...... 13 13. Thallus composed of corticate granules; C+ orange (xanthones only); SW and both coasts...... …………………...... … Pyrrhospora quernea (Dicks.) Körb. (fig. 19E-F) 13. Thallus areolate, immersed, or otherwise not as above; C+ or C- (xanthones present or absent); distribution various...... …14 14. On wood in boreal coastal/maritime areas; thallus C+ yellow-orange, (atranorin, zeorin, and xanthones); distribution various...... …15 15. Soralia discrete; very common on driftwood…...... Lecanora oraefrigidae R. Sant. (fig. 20B) 15. Soralia irregular and diffuse; not common…...... Lecanora expallens Ach. (fig. 20A) 19 14. On bark in other habitats; thallus C+ or C- (xanthones present or absent); all PNW…...... 16 16. Zeorin present...... …17 17. Coronatone and unknown xanthone...... … Bacidia viridifarinosa Coppins & P. James 17. Other xanthone present...... … Rinodina sheardii Tønsberg 16. Zeorin absent...... …18 18. Thallus UV+ dull to strong orange or red, C+ (xanthones present)...... …19 19. Thallus dispersed areolate…...... Rinodina flavosoralifera Tønsberg (fig. 23F) 19. Thallus +/- continuous…...... Biatora disjuncta Printzen & Tønsberg 18. Thallus UV-, C- (xanthones absent)...... …20 20. Thallus composed of flat areoles; soralia punctiform; soredia yellow; usnic acid, isousnic acid, and atranorin...... … Biatora flavopunctata (Tønsberg) Hinteregger & Printzen 20. Thallus +/- immersed or indistinct; soralia convex and diffuse; soredia green; 2-0- methylsulphurellin, usnic acid, and atranorin…...... Lecanora jamesii J.R. Laundon

20 KEY TO SPECIES OF PHYLLOPSORA

The key presented here is reproduced (in simplified form) from Harris (1995). The genus is badly in need of a modern revision in North America. The number of species increases as one travels south in the Coastal Plain, especially once one enters the Southeastern Plains Ecoregion and the Southern Coastal Plain Ecoregion. There are only two or three common species in the Mid-Atlantic Coastal Plain (i.e. N of the GA/SC border), Piedmont, and low elevations of the Appalachian Mountains: Phyllospora confusa Swinsc. & Krog, P. corallina (Eschw.) Müll. Arg. var. corallina, and P. parvifolia (Pers.) Müll. Arg.) var. parvifolia. The user should consider the above when naming specimens. The primary reason I have chosen to include this group here is that species of Phyllopsora are commonly sterile and the morphological terminology used by Brako/Harris is hard to understand without illustrations. Hopefully users will find this helpful. The only species likely be confused with Phyllospora is Bacidia schweinitzii (Fr. ex E. Michener) A. Schneider, sterile thalli of that species typically produce some black pycnidia.

1. Thallus P-, argopsin or pannarin absent…...... 2 2. Prothallus white, although sometimes poorly developed...... 3 3. Squamules small (if you don’t think your specimen has squamules follow this lead); thalli not forming circular rosettes with what appear to be marginal lobes...... 4 4. Thallus with marginal proliferations/lobules (if you don’t think your specimen has isidia follow this lead)………………...... 5 5. Squamules very small, with short globose isidia…...... Phyllospora kalbii Brako 5. Squamules larger, with elongate structures that look intermediate between lobules and isidia…...... 6 6. Squamules adnate, with narrow proliferations...... 7 7. Proliferations long, abundant, forming overlapping layers...…Phyllospora confusa Swinsc. & Krog 7. Proliferations short, not as abundant, not forming overlapping layers...... ……………...... …Phyllopsora corallina (Eschw.) Müll. Arg. var. corallina 6. Squamules erect, distinctly lobulate; RARE...... …Phyllopsora parvifoliella (Nyl.) Müll. Arg. 4. Thallus with isidia...... 7 7. Isidia short, globose…...... Phylloposora kalbii Brako 7. Isidia elongate, not globose...... 8 8. Isidia long and becoming tangled…...... Phyllospora isidiolyta (Vainio) Brako 8. Isidia obpyriform (fatter at the base and constricted at the point of attachment)...... …Phyllopsora corallina var. rappiana Brako 3. Squamules large; thalli forming circular rosettes with what appear to be marginal lobes...... 9 9. Marginal proliferations abundant, narrow, ecorticate, smaller than the primary squamules; COMMON…. ………………...... …Phyllopsora parvifolia (Pers.) Müll. Arg. var. parvifolia 9. Marginal proliferations sparse, not narrow and ecorticate, resembling the primary squamules; RARE…... …………………...... …Phyllopsora parvifolia var. breviuscula (Nyl.) Brako 2. Prothallus black...... 10 10. Isidia long, slender, and abundant…...... Phyllopsora furfuracea (Pers.) Zahlbr. 10. Isidia short, cylindrical, less abundant...... …Phyllopsora halei (Tuck.) Zahlbr. 1. Thallus P+ orange-red, argopsin or pannarin present…...... 11

21 11. Isidiate; squamules epruinose and small (if you don’t think your specimen has squamules follow this lead); thalli not forming circular rosettes with what appear to be marginal lobes...... 12 12. Argopsin the major substance...... …13 13. Lacking phyllopsorins; common?…...... Phyllopsora corallina var. santensis (Tuck.) Brako 13. Containing phyllopsorins; rare...... …Phyllopsora corallina var. ochroxantha (Nyl.) Brako 12. Vicanicin the major substance...... …Phyllopsora corallina var. glaucella (Vainio) Brako 11. Lobulate; squamules pruinose and large; thalli forming circular rosettes with what appear to be marginal lobes. …………………………...... 14 14. Argopsin present...... …Phyllopsora buettneri var. glauca (de Lesd.) Brako (note this is not reported to have zeorin but I have seen material from GA with zeorin) 14. Pannarin and zeorin present...... …Phyllopsora buettneri var. munda (Malme) Brako

22 KEY TO P+ SPECIES WITH ARGOPSIN, BAEOMYCESIC ACID, FUMAR/PROTOCETRARIC ACID, PANNARIN, PSOROMIC ACID, OR STICTIC ACID AGG.

1. Thallus K-, P+ instantly strong yellow (psoromic or baeomycesic acid present)...... …2 2. On bark and rarely on rock; thallus thick, bluish-green, irregularly sorediate, and often appearing fluffy, UV- (psoromic acid present)…...…Phlyctis boliviensis Nyl. (syn. P. ludoviciensis (Müll. Arg.) Lendemer) (fig. 25A) 2. On soil; thallus thin, white, schizidiate, not fluffy, UV+ dull orange (baeomycesic acid present)...... ………………………………………………….....…sterile forms of Dibaeis baeomyces (L.) Rambold & Hertel 1. Thallus K- or K+ dirty yellow, P+ various (other substances)…...... 3 3. On rock……………………………………………...... …4 4. Thallus isidiate…………………………….…...... 5 5. Thallus brown; isidia short and scruffy; arctic/boreal?, rare.....Rhizocarpon furfurosum H. Magn. & Poelt (s. str. contains stictic acid, but North American material contains norstictic?) 5. Thallus gray; isidia long and cylindrical; southern Appalachians, rare; fumarprotocetraric acid……………. ……………………………………………...... … Pertusaria andersoniae Lendemer 4. Thallus sorediate, blastidiate, or schizidiate...... …6 6. Thallus with discrete soralia…...... Porpidia soredizodes (J. Lahm) J.R. Laundon (fig. 26A-B) (note a rare sorediate chemotype of P. albocaerulescens (Wulfen) Hertel & Knoph will also key out here) 6. Thallus with schizidia, blastidia, or pustules...... …7

7. Boreal/northern; in humid shaded microhabitats; thallus green, schizidiate; stictic acid...... …………………………………………...... …Baeomyces rufus (Huds.) Rabent. 7. Eastern temperate or arid central/western; habitats various; thallus brown or gray, sorediate or blastidiate; argopsin or substictic acid…...... 8 8. Eastern temperate; on sun exposed rock; thallus brown-gray, blastidiate; blastidia black; apothecia almost always present; argopsin…...... Halecania pepegospora (H. Magn.) v.d. Boom 8. Arid central/western North America; on sheltered rocks usually in humid microhabitats; thallus lead- gray or brownish, irregularly pustulose-sorediate; apothecia rare; substictic acid...... …………………………….….Aspicilia cf. simoensis Räsänen (substictic acid chemotype) (fig. 7A-B) 3. On bark, mosses, humus, or wood……………...... …9 9. On humus or old wood, very rarely on bark…...... 10 10. On old wood; soralia distinctly excavate with darkened soredia...... …11 11. Oceanic, bi-coastal; typically on driftwood; stictic acid...... ………………………………………………...... Xylographa vitilago (Ach.) J.R. Laundon (fig. 26E-F) 11. Inland; western North America; not typically on driftwood; fumarprotocetraric acid...... ……………………………………………...... … Xylographa soralifera Holien & Tønsberg 10. On humus or mosses……………………...... 12 12. Thallus isidiate; arctic/boreal; fumarprotocetraric acid…...... Pertusaria dactylina (Ach.) Nyl. 12. Thallus sorediate or schizidiate; distribution various; stictic acid or argopsin with gyrophoric acid…..13 13. Thallus schizidiate, placodioid with marginal radiating lobes; on humus; stictic acid...... ……………………………………...... … Baeomyces placophyllus Ach. 13. Thallus sorediate, areolate; on mosses; argopsin + gyrophoric acid...... ………………………………………...... … Micarea leprosula (Th. Fr.) Coppins & A. Fletcher

23 9. On bark……………………………………...... …14 14. Appalachians, temperate eastern North America or southeastern US...... …15 15. Thallus appearing isidiate, but “isidia” actually stipitate pycnidia; on conifers in humid microhabitats; fumarprotocetraric acid… ...... Micarea neostipitata Coppins & P. May (fig. 24A) 15. Thallus sorediate, schizidiate, pustulose, otherwise not isidiate…...... 16 16. Thallus C+ pinkish (gyrophoric acid in addition to argopsin); temperate...... ………………………………...... …Biatora printzenii Tønsberg (fig. 14F) 16. Thallus C- (gyrophoric acid absent); temperate or not…...... 17 17. Thallus without discrete soralia, pustulose, blastidiate, or with irregular diffuse soralia…...... 18 18. Thallus composed of dispersed minute areoles, coarsely blastidiate; pannarin...... ………………………………………………………………… Rinodina excrescens Vain. (fig. 26F) 18. Thallus +/- continuous, pustulose or with diffuse soralia; chemistry various…...... 19 19. Thallus distinctly blue, pustulose-sorediate; pannarin and zeorin; common...... ……………………………...... Megalospora porphyritis (Tuck.) R.C. Harris 19. Thallus white-gray or green, with irregular diffuse soralia; fumarprotocetraric acid present; rare……………………………...... …20 21. Thallus distinctly white-gray; atranorin and zeorin...... …………………………...... … Megalaria pulverea (Borrer) Hafellner & E. Schreiner 21. Thallus green; usnic acid...... Lecanora conizaeoides Nyl. ex Cromb. (fig. 24C) 17. Thallus with discrete soralia (note the soralia may appear coarse and +/- pustulose)...... 21 21. Thallus UV+ dull orange-red or pink (coronatone present); tropical SE; stictic acid...... ……………………………………...... … Pertusaria expolita R.C. Harris (fig. 23C) 21. Thallus UV- (xanthones absent); temperate/Appalachians; without stictic acid...... …22 22. Soralia white, discoid; protocetraric acid present…...... Pertusaria multipunctoides Dibben 22. Soralia not white and discoid; other substances present...... …23 23. Thallus continuous, often forming circular rosettes; fumarprotocetraric acid present…...... 23 24. Thallus distinctly greenish-brown; atranorin absent...... …………………………...... … Fuscidea arboricola Coppins (fig. 24B) 24. Thallus lead-gray; atranorin present..… Mycoblastus fucatus (Stirton) Zahlbr. (fig. 24D) 23. Thallus +/- dispersed areolate, not forming rosettes; pannarin and zeorin present…...... 25 25. Soralia pigmented yellowish; areoles convex...... …………………………...... …Rinodina efflorescens Malme (fig. 25C-D) 25. Soralia not pigmented yellowish; areoles +/- flat...... ………………………...... … Rinodina willeyi Sheard & Giralt (fig. 25B) 14. Other regions…………………………...... …26 26. Thallus blastidiate; high elevations of AZ; pannarin and zeorin……...... Rinodina perreagens Sheard 26. Thallus sorediate; PNW/boreal-arctic North America; various substances…...... 27 27. Soralia irregular and diffuse; atranorin and fumarprotocetraric acid present...... …28 28. Zeorin present; apothecia black, ascospores hyaline, 2-celled...... …………………………………...... … Megalaria pulverea (Borrer) Hafellner & E. Schreiner 28. Zeorin absent; apothecia red or brownish…...... 29 29. Apothecia red………………...... Ramboldia cinnabarina (Sommerf.) Kalb et al. (fig. 24E) 29. Apothecia brownish/pale…...... Cliostomum flavidulum Hafellner & Kalb 24 27. Soralia discrete……………………...... …30 30. Thallus immersed and imperceptible...... …31 31. Soralia minute, excavate; argopsin…...... Biatora efflorescens (Hedl.) Räsänen 31. Soralia larger; hemispherical; fumarprotocetraric…...... Pertusaria pupillaris (Nyl.) Th. Fr. 30. Thallus not immersed, typically well developed...... …32 32. Thallus +/- dispersed areolate...... 33 33. Soralia pigmented yellowish; pannarin and zeorin. Rinodina efflorescens Malme (fig. 25C-D) 33. Soralia not pigmented yellowish; stictic acid with atranorin and zeorin...... ……………………………...... … Rinodina stictica Sheard & Tønsberg 32. Thallus continuous………………...... 34 34. Thallus creamy white; stictic acid...... ……………………………...... Coccotrema pocillarium (C.E. Cumm.) Brodo (fig. 26C-D) 34. Thallus brown or gray; fumarprotocetraric acid...... …35 35. Thallus lead-gray; atranorin present...... … Mycoblastus fucatus (Stirton) Zahlbr. (fig. 24D) 35. Thallus olive-brown or blue-gray; atranorin absent…...... 36 36. Thallus olive-brown; soralia small and plane...... … Lecidea praetermissa Tønsberg 36. Thallus blue-gray; soralia large and hemispherical...... ………………………...... Pertusaria borealis Erichsen (fig. 24F)

25 K- OR K+ YELLOWISH SPECIES WITHOUT EASILY IDENTIFIED SUBSTANCES, LACKING SUBSTANCES, OR WITH ATRANORIN

1. Thallus cortex K+ purple in section viewed under the compound microscope, Thalloidima Green present...... 2 2. Thallus on bark or wood…………………...... 3 3. On bark………………………………...... …4 4. Typically on hardwoods (Populus); thallus immersed; soredia indistinct and easily overlooked; apothecial margins orange………………………………………………………….…Caloplaca ahtii Søchting (fig. 28F) 4. Typically on conifers; thallus well developed, not immersed; soredia obvious...... ………………………………………...... …Caloplaca pinicola Wetmore 3. On lignum……………………………...... …5 5. Apothecial margins gray; widespread in the arid west...... …Caloplaca pinicola Wetmore 5. Apothecial margins orange; rare, known from vicinity of ND & SD...... ………………………………………...... …Caloplaca lignicola Wetmore (fig. 28C) 2. Thallus on rock…………………………...... …6 6. Thallus with irregular lobules/blastidia…...... Caloplaca chlorina (Flot.) Olivier (fig. 28A-B) 6. Thallus with soredia…………………….…...... 7 7. Apothecial margins gray; soralia laminal and marginal, very irregular...... ………………………………………...... …Caloplaca pratensis Wetmore (fig. 27E) 7. Apothecial margins orange; soralia marginal and discrete...... ……………………………………...... …Caloplaca soralifera Vondrák & Hrouzek (fig. 27D) 1. Thallus cortex K- or K+ yellow, not K+ purple in section, Thalloidima Green absent...... …8 8. Thallus squamulose and with distinct hair-like papillae...... …………………………………………………...… Agonimia opuntiella (Buschardt & Poelt) Vězda (fig. 29D) 8. Thallus squamulose or not, without distinct hair-like papillae...... 9 9. Thallus emerald/jade colored, composed of flat squamules with a raised rim and marginal soralia...... …………………………………………...... …Normandina pulchella (Borrer) Nyl. (fig. 29C) 9. Thallus not as above……………………...... 10 10. On rock……………………………...... …11 11. Thallus with marginal lobes, C+ KC+ yellow where pigmented; coastal southern California; atranorin +diploicin……………….… Diploicia canescens (Dicks.) A. Massal. 11. Thallus without marginal lobes, C- K+/-, unpigmented; not in coastal southern California; chemistry various…………………………….…...... 12 12. Thallus composed of convex bullate areoles; soralia laminal; soredia dark; arid SW; atranorin, gangaleoidin, roccellic acid………...... … Lecanora pannonica Szatala (fig. 31A) 12. Thallus not as above; soralia various; soredia various; distribution various; without gangaleoidin …13 13. Thallus forming small lobed rosettes; soralia laminal and deeply excavate; Appalachians...... ………………………………...... ….Caloplaca reptans Lendemer & Hodkinson ined. (fig. 27F) 13. Thallus not as above; soralia not laminal and deeply excavate; mostly not in Appalachians...... …14 14. Southern Appalachians; thallus granulose sorediate, without discrete soralia; apothecia frequently present, ascospores 2-celled; atranorin and zeorin…………………………………………. ………………………………...... Megalaria beechingii Lendemer (fig. 31B) 14. Not in the Southern Appalachians; thallus not granulose sorediate, frequently with discrete soralia; apothecia absent or ascospores not 2-celled…...... 15

26 15. Humid sheltered microhabitats in the SE and south Central USA; thallus thin, pale; soralia distinctly white; atranorin only, in very high concentration...... ……………………………...... …Caloplaca yuchiorum Lendemer & C.A. Morse 15. Not in humid sheltered microhabitats in the SE, occasional in south Central USA; thallus not as above; atranorin and zeorin present or no substances...... …16 16. Soralia not distinctly marginal; Pacific NW; in oceanic habitats; atranorin, zeorin, porphyrilic acid, +/- bourgeanic acid…...... Haematomma ochroleucum (Neck.) J.R. Laundon 16. Soralia distinctly marginal; not in the PNW; in inland habitats; lichen substances lacking..17 17. Thallus brown, epruinose; soralia not becoming excavate; perithecia occasionally present……………………...... Verrucaria furfuracea (de Lesd.) Breuss (fig. 27C) 17. Thallus tan, pruinose; soralia distinctly excavate, often hollowing out the areoles; apothecia very rare…...... Acarospora moenium (Vain.) Räsänen (fig. 27A-B) 10. On bark, wood, humus, or mosses…...... 18 18. Soralia circular, discoid, KC+ lavender in section, bitter tasting (picrolichenic acid present); thallus with pseudocyphellae………………..…...... Pertusaria amara (Ach.) Nyl. 18. Soralia discoid or not, KC- or KC+ yellow not KC+ lavender in section, not bitter tasting (picrolichenic acid absent); thallus with or without pseudocyphellae…...... 19 19. Thallus lobed, appearing foliose, small or large…...... 20 20. Thallus large, resembling a Physcia or Dirinaria; medulla often pigmented, C+ and KC+ yellow; coastal southern California……………………………………Diploicia canescens (Dicks.) A. Massal. 20. Thallus smaller, resembling a minute Phaeophyscia; medulla not pigmented, C-, KC-; widespread.. …………………………………...... … Hyperphyscia adglutinata (Flörke) H. Mayrh. & Poelt 19. Thallus not lobed, not appearing foliose, size various...... …21 21. Thallus blastidiate, without soralia or isidia; no substances present; on bark...... …22 22. Eastern North America and Ozarks; thallus minute and almost imperceptible...... ………………………………...... … Rinodina papillata H. Magn. (fig. 30A-B) 22. Central California; thallus obvious…...... Rinodina poeltiana Giralt & Obermayer 21. Thallus sorediate, with soralia or isidiate; substances present or absent; on various substrates…...23 23. Thallus isidiate…………………...... …24 24. Tropical south/southeast; thallus with pseudocyphellae; isidia tall, cylindrical...... ……………………………...... …Caloplaca wrightii (Tuck.) Fink (fig. 29A) 24. Arid west/central or boreal/northern; thallus without pseudocyphellae; isidia various...... …24 25. Arctic/boreal; on humus; with large coarse isidia…... Pertusaria panyrga (Ach.) A. Massal. 25. Arid western interior species; on bark and wood; with irregular small granular isidia...... …………………………...... …Caloplaca furfuracea H. Magn. (fig. 28D) 23. Thallus sorediate……………...... 26 26. On soil or rarely VERY rotten organic matter; northern/Appalachian; with diffuse green soralia and thin gelatinous film-like thallus; no substances...... ………………………...... … Trapeliopsis gelatinosa (Schrad.) Coppins & P. James (fig. 29F) 26. On bark or mosses; distribution various; soralia and thallus various; substances present or absent.…………………………………………………………………………………………...... 27 27. Soralia distinctly cupuliform and excavate; not common; western…...... 28 28. Thallus distinctly squamulose; widespread in southern/central California...... ………………………...... … Waynea californica Moberg (fig. 29E) 27 28. Thallus areolate; known only from one locality in Yosemite NP...... …………………………...... … Caloplaca lecanoroides Lendemer 27. Soralia not cupuliform, excavate or not; some species more frequent; distribution various...... ………………………...... …29 29. Soredia consistently dark in color, often darker than the thallus; atranorin present…...... 30 30. Soralia excavate; soredia contrasting distinctly against a pale thallus that is often immersed; atranorin and placodialic acid...... ……………………...... … Buellia arborea Coppins & Tønsberg (fig. 30E) 30. Soralia plane or convex; soredia blending in against the thallus and substrate; atranorin and zeorin…………...... … Rinodina griseosoralifera Coppins (fig. 30D) 29. Soralia concolorous with the thallus, or lighter in color, occasionally darkened but not typically; atranorin present or absent...... 31 31. Soralia large, white, and irregular; southern/central California; allopertusaric acid present………………...... Pertusaria albescens (Huds.) M. Choisy & Werner (fig. 31C) 31. Soralia smaller not large and irregular; distribution various; allopertusaric acid absent.... ………………………...... …32 32. Tropical and subtropical southeast…...... 33 33. Thallus greenish, with large diffuse soralia; fibrous white prothallus often present; atranorin and placodialic acid agg...... ………………...... …. Haematomma guyanense Staiger & Kalb (fig. 30F) 33. Thallus gray or white, with discrete discoid soralia; never with fibrous white prothallus; no substances…...... Pertusaria ophthalmiza (Nyl.) Nyl. (fig. 31D) 32. Other regions...... …34 34. Thallus K+ yellow, atranorin present (observing this reaction in section may be helpful)…………...... 35 35. Eastern North America or boreal/northern...... …36 36. Soralia diffuse and irregular, not discrete; caperatic acid present; often on conifers…...... Cliostomum leprosum (Räsänen) Holien & Tønsberg 36. Soralia discrete and regular in shape; caperatic acid present or absent; on conifers or not…...... 37 37. Thallus distinctly dispersed areolate; zeorin present...... …………...... … Rinodina degeliana Coppins (fig. 30C) 37. Thallus not as above; zeorin absent…...... 38 38. Northern/Great Lakes; two unknown terpenoids that behave like fatty acids but color up after charring present...... ………...... … Lecanora allophana f. sorediata Schrad. 38. Temperate? Coastal?; roccellic acid present. … Lecanora farinara Borrer 35. Western/southwestern………………………………………………………...…39 39. Soralia circular and excavate; zeorin absent; widespread in southwest… ……………………….………………………………...Lecanora impudens Degel. 39. Soralia irregular and eroding; zeorin present; known from one site in the Channel Islands………………………………...… Lecanora carneolutescens Nyl. 34. Thallus K-, atranorin absent…...... 40

28 40. Without fatty acids or pseudoplacodialic acid…...... 41 41. Soredia distinctly greenish or yellowish when fresh…...... 42 42. On bark; soralia small and discrete; Appalachians/Great Lakes...... ……………...... … Lecania croatica (Zahlbr.) Kotlov 42. On mosses at base of trees, rarely on bark; soralia larger and irregular; northern/boreal and Great Lakes...... ……………...... … Mycobilimbia epixanthoides (Nyl.) Vitik. et al. (fig. 31F) 41. Soredia white or gray when fresh…...... 43 43. Thallus continuous and soralia discoid...... …………...... … Pertusaria ophthalmiza (Nyl.) Nyl. (fig. 31D) 43. Thallus areolate or not obviously continuous, soralia discoid or not...... …44 44. Soralia circular, not excavate, distinctly elevated above the thallus; PNW... ………...... … Caloplaca sorocarpa (Vainio) Zahlbr. (fig. 29B) 44. Soralia excavate, not circular and distinctly elevated above the thallus; PNW or Great Plains...... …45 45. Mid-western North America; on large old hardwoods; apothecia orange; epihymenium K+...... Caloplaca ulcerosa Coppins & P. James (fig. 28E) 45. Widespread but rare; on various hardwoods; apothecia brown; epihymenium K-...... Caloplaca obscurella (Lahm ex Körber) Th. Fr. 40. Fatty acids or pseudoplacodialic acid present...... …46 46. Fatty acids present; soralia small and discrete; apothecia red; PNW...... ………...... … Ramboldia subcinnabarina (Tønsberg) Kalb et al. 46. Pseudoplacodialic acid present; soralia irregular and diffuse; apothecia brown- black…...... Lecidea leprarioides Tønsberg (fig. 31E)

29 PRELIMINARY KEYS TO LEPRARIA S.L. IN NORTH AMERICA.

Throughout these keys the term “hypothallus” refers to the layer of fungal tissue that underlies the lichenized granules. The thickness of a given thallus is due to the thickness of the hypothallus. Species with a thin hypothallus are those typically referred to as having a thallus that is “dispersed,” “lacking a medulla,” or “thin and continuous.” Similarly, species with a thick hypothallus are those that are typically referred to as having “a medulla.” Differences in the presence/absence of a well developed pseudocortex on the thalline granules are also used to distinguish taxa in the keys below. Species in which the granules have a well developed pseudocortex do not appear “fluffy” or “cottony” and are almost all are members of the L. caesioalba-neglecta group. The granules of these taxa have a characteristic gestalt because they are corticate and compact instead of ecorticate and composed of loosely intertwined hyphae. This morphology is apparently either an adaptation to, or induced by, the ecology of these taxa, all of which occur in exposed habitats.

KEY USING TLC & MORPHOLOGY

This key is meant for use with specimens that have been analyzed with Thin Layer Chromatography.

1. Pannarin and zeorin present………………………………………………………………………...... 2 2. Pannaric acid 6-methyl ester present; terricolous; coastal southern California and Channel Islands...... ……. ……………...... …………………L. terricola Lendemer 2. Pannaric acid 6-methyl ester absent; not terricolous, typically saxicolous; widely distributed throughout North America………………………………...... ….L. adhaerens K. Knudsen, Elix & Lendemer 1. Pannarin absent, zeorin present or absent………………………...... …………….…………………………….3 3. Argopsin present, zeorin absent………...... …………………………..L. santamonicae K. Knudsen & Elix 3. Argopsin absent, zeorin present or absent…...... ………………………………………………………..…...4 4. Divaricatic acid present ………………...... ………………………………………………………………5 5. Zeorin absent; saxicolous on non-calcareous rocks; Appalachians and Ozarks...... …L. cryophila Lendemer 5. Zeorin present; saxicolous or corticolous; distribution various………………………...... ……………...6 6. Nordivaricatic acid present in high concentration; well developed hypothallus present; typically on calcareous rocks but corticolous in North America?; present in Pacific Northwest?...... ………………………………………………………………………...... L. crassissima (Ach.) Lettau 6. Nordivaricatic acid absent or present only as trace; well developed hypothallus absent; corticolous and rarely saxicolous on non-calcareous rocks in humid habitats; throughout………….....…L. incana (L.) Ach. 4. Divaricatic acid absent……………………………………...... ………………………………………...…7 7. Barbatic acid present; Ozarks………...... ……………………………………………L. barbatica Lendemer 7. Barbatic acid absent…………………………...... ………………………………………………………...…8 8. Xanthones present……………………………………………………………...... ………………………..9 9. Hypothallus well developed; on calcareous rocks; thiophanic acid and arthothelin present…………...... …………………………………………………………………………...... ……….L. disjuncta Lendemer 9. Hypothallus thin and not well developed; on non- or weakly calcareous rocks; 5,7-dichloro-3-O- methylnorlichexanthone, 3-O-methylasemone present……………...... …………L. xanthonica Lendemer 8. Xanthones absent…………………………………………………...... ………………………………….10 10. Obtusatic acid present……………………………………………...... ……….L. moroziana Lendemer 10. Obtusatic acid absent………………………………………………...... ……………………………...11 11. Usnic acid and zeorin present……………………………………………...... …………………..…12 12. Thallus distinctly leprose, granules distinct; widely distributed…....…L. ecorticata J.R. Laundon s.l. 12. Thallus continuous not distinctly leprose; southern California………………………...... ………...... L. texta K. Knudsen, Elix & Lendemer 11. Usnic acid absent, zeorin present or absent………………………………...... ………………….…13 30 13. Stictic acid present………………………………………………………...... ……………………...14 14. Zeorin absent; rangiformic acid present…………...... …...L. caesioalba (Ach.) J.R. Laundon s.l. 14. Zeorin present; roccellic/angardianic acid present or absent……...... …………………………...15 15. Hypothallus thick and well developed; ubiquitous…………...... ……………L. lobificans auct. 15. Hypothallus thin and poorly developed; uncommon…………...... …….…L. elobata Tønsberg 13. Stictic acid absent………………………………………………...... ………………………………16 16. Alectorialic acid present……………………………………...... ………………………………..17 17. Porphyrilic acid present; thallus composed of large flat areoles; arctic…...... ………………….. ………………………………………………………………..…...... L. gelida Tønsberg & Zhurb. 17. Porphyrilic acid absent; thallus composed of distinct granules; not arctic...... ……………..…18 18. Hypothallus thin; granules with a distinct pseudocortex, not appearing fluffy…...... ………... ………………………………………………………...... …………...L. neglecta (Nyl.) Erichsen 18. Hypothallus thick, well developed; granules without a pseudocortex, appearing fluffy…...... ……………………………………………………………...... …….…L. eburnea J.R. Laundon 16. Alectorialic acid absent………...... …………………………………………………………....…19 19. Squamatic acid present……………...... …………………………………….L. squamatica Elix 19. Squamatic acid absent………………………...... …………………………………………..…20 20. Fumarprotocetraric/protocetraric acid present……………...... …………………………….21 21. Fumarprotocetraric acid present (major)…………………………...... ………………...…22 22. Thallus thick, with a well developed hypothallus………………………...... ………..…23 23. Atranorin absent; boreal North America…...... ……L. torii Perez-Ortega & T. Sprib. 23. Atranorin present; widely distributed…………...... …………………………………24 24. Thallus with “crisped” marginal lobes resembling Normandina; darkened rhizohyphae present; roccellic/angardianic acid present...L. normandinoides Lendemer & R.C. Harris 24. Thallus without “crisped” marginal lobes; darkened rhizohyphae absent; roccellic/angardianic acid absent …………...... ………………L. nivalis J.R. Laundon 22. Thallus thin, without a well developed hypothallus…...... ……………………………..25 25. Granules with a distinct pseudocortex; atranorin present; saxicolous or corticolous; not known from the SE coastal plain…………...... …L. caesioalba s. str. 25. Granules without a distinct pseudocortex; atranorin absent; corticolous and typically on conifers; SE coastal plain...... …..L. friabilis Lendemer, Knudsen & Elix 21. Protocetraric acid present (major)……...... ………………………………………………..26 26. Atranorin absent; granules extremely large (>1 mm); high elevations of the Appalachians…………………………...... ………………………………L. lanata Tønsberg 26. Atranorin present; granules smaller; distribution various…...... ……………………..…27 27. Thallus with “crisped” marginal lobes resembling Normandina; darkened rhizohyphae present; roccellic/angardianic acid present……...... ……..L. normandinoides 27. Thallus without “crisped” marginal lobes; darkened rhizohyphae absent; roccellic/angardianic acid absent……………………………………...... ……..…L. nivalis 20. Fumarprotocetraric/protocetraric acid absent……...... ……………………………………..28 28. Psoromic acid present………………...... ……………………...…L. caesioalba chemotype 28. Psoromic acid absent………...... ……………………………………….……………….…29 29. Norstictic acid present……………………...... ……………..…L. caesioalba chemotype 29. Norstictic acid absent………………………………...... ……………………………….30 30. Thamnolic acid present…...... ……………………..L. aurescens Orange & Wolseley 30. Thamnolic acid absent……………...... ……………………………………………...31

31 31. Lecanoric acid present; high elevations of Arizona……...... L. lecanorica Tønsberg 31. Lecanoric acid absent; distribution various……………...... …………………...…32 32. Salazinic acid present; Appalachians……...... ……...... …L. salazinica Tønsberg 32. Salazinic acid absent; distribution various………...... ………………………….33 33. Zeorin present…………………………………………...... ……………....…34 34. Pallidic acid present; temperate………...... …….…L. caesiella R.C. Harris 34. Porphyrilic acid present; Greenland……………...... L. zeorinica Saag et al. 33. Zeorin absent…………………...... …………………………………………..35 35. Granules with a distinct pseoducortex; members of the L. caesioalba-neglecta group……………...... ……………….……………………………….36 36. Rangiformic acid present; porphyrilic acid absent……...... ……………… ...... L. borealis Tønsberg & Lohtander 36. Roccellic/anagardianic acid present; porphyrilic acid present or absent…………...... ………………………………………………………...37 37. Porphyrilic acid present…...... L. alpina (de Lesd.) Tretiach & Baruffo 37. Porphyrilic acid absent…………...... …………L. caesioalba chemotype 35. Granules without a distinct pseudocortex; not members of the above group….38 38. Dibenzofurans present………...... ……………………………………..39 39. Pannaric acid 6-methylester present (major)……...... ……………….40 40. Thallus resembling Cladonia squamules; terricolous; coastal southern California………………………...... ….L. xerophila Tønbserg 40. Thallus not as above; not in coastal southern California……...... …... ……………………………………...... L. vouauxii (Hue) R.C. Harris 39. Pannaric acid 6-methylester absent or trace……...... ……………….…41 41. Pannaric acid present (major); common in boreal North America and rare in Appalachians…...... ………...L. membrancea (Dicks.) Vain. 41. 4-0-oxypannaric acid 2-methylester present (major); widely distributed and very rare……...... …..L. diffusa (J.R. Laundon) Kukwa 38. Dibenzofurans absent……...... ………………………………………….42 42. Roccellic/angardianic acid present; thallus with “crisped” marginal lobes; saxicolous…………...... …………...L. normandinoides chemotype 42. Roccellic/angardianic acid absent; thallus not as above…...... ………43 43. Nephrosterinic acid present…...... L. rigidula (Hue) J.R. Laundon 43. Jackinic/rangiformic acid present……...... ……...L. jackii Tønsberg

KEY USING SPOT TESTS & MORPHOLOGY

Note that this key is intended for use by those without access to TLC, or as a “first attempt” to identify a specimen whose chemistry will be confirmed using the TLC and the key above. For the most part I have structured the key to use spot rests (K, C, KC, P, and UV) that produce consistent results, however concentrations of the various substances can be variable and result in variable spot test results. Spot tests are often more reliable when performed on an acetone extract of the thallus. Several groups of species that differ in the presence of different terpenoids/fatty acids which cannot be detected with spot tests are keyed out collectively.

1. Thallus UV+ blue-white (divaricatic acid, barbatic acid, obtusatic acid, or squamatic acid present)...... 2 2. Thallus PD+ orange-red (protocetraric acid and baeomycesic acid in addition to squamatic acid); inland Florida and southern Georgia barrier islands……...... …………………………………………...L. squamatica 2. Thallus PD- (protocetraric and baeomycesic acids absent); not present in the above region, but widely distributed elsewhere……………………………………...... ……………………………………………….....3 32 3. Thallus with K+ dark purple patches (anthraquinones present in addition to isousnic and obtusatic acids); high elevations of the Appalachians……………………………...... …………………...L. moroziana 3. Thallus K- (anthraquinones absent); mostly not high elevations of the Appalachians (except L. cryophila). ……………………………………………...... …………………………………………………..4 4. Thallus KC+ gold (usnic acid present in addition to barbatic acid); Ozarks……………...... L. barbatica 4. Thallus KC- or KC+ pink/red (usnic acid absent)………………………………………………...... …...5 5. Corticolous (rarely saxicolous); thallus thin, with a poorly developed hypothallus; C-/KC-…....L. incana 5. Saxicolous (rarely corticolous); thallus thick, with a well developed hypothallus; C+/KC+ pink- red……………………...... ………………………………………………………………………………6 6. On non-calcareous rocks; Appalachian…………………...... ………………………….L. cryophila 6. On calcareous rocks (and conifers?); western?...... L. crassissima 1. Thallus UV- or UV+ dull orange……………………………………………………...... ……………………...7 7. Thallus greenish-yellow, acetone extract of thallus K-, C-, KC+ golden yellow (usnic acid present).. ……………………………………………………………...... ……………………………………………....8 8. Thallus obviously leprose, composed of granules…………………...... ………………………………….9 9. Saxicolous on calcareous rocks; acetone extract or thallus positive with Dimroth’s solution (xanthones present); rare………………………………………...... ………………………....L. disjuncta 9. Saxicolous on non-calcareous rocks or corticolous; acetone extract or thallus positive or negative with Dimroth’s solution (xanthones present or absent); more common than the above taxon...... 10 10. Rare; acetone extract or thallus positive with Dimroth’s solution; eastern…...... …..L. xanthonica 10. Common; acetone extract or thallus negative with Dimroth’s solution; widespread…...... L. ecorticata 8. Thallus not distinctly leprose, appearing to be a continuous network of hyphae and algae with delimited margins; saxicolous; southern California……………...... ……………………………...... L. texta 7. Thallus not usually greenish yellow, acetone extract of thallus K+/-, C+/-, KC- or KC+ not gold (usnic acid absent)………………………………………………...... ………………………………………………..11 11. Thallus KC+ pink/red (alectorialic acid present)……………...... ……………………………………...12 12. Thallus composed of large areole-like structures; arctic……...... …………………………...…L. gelida 12. Thallus composed of distinct granules; not arctic……………………...... …………..………………..13 13. Thallus thin, without a well developed hypothallus; granules with a distinct pseudocortex; saxicolous or corticolous…………………………...... …………………………………….....L. neglecta 13. Thallus thick, with a well developed hypothallus; granules without a distinct pseudocortex; muscicolous, lignicolous, or corticolous at the bases of trees……...... ……………………...L. eburnea 11. Thallus KC- (alectorialic acid absent)...... 14 14. Thallus K+ red, P+ yellow or orange (salazinic or norstictic acid present)……...... ……………….....15 15. Thallus P+ yellow, K+ red crystals in water mount; western?...... ………………………………………………...... …………...L. caesioalba chemotype (norstictic acid) 15. Thallus P+ orange, K+ red but not producing crystals in water mounts; Appalachian……...... ……...... ……………………………………………………………………… ...L. salazinica (salazinic acid) 14. Thallus K- or K+ yellow, P- or P+ various...... 16 16. Thallus C+ pink/red (lecanoric acid present); high elevations of Arizona……...... …....L. lecanorica 16. Thallus C-………………………………………………………………………...... ……………....17 17. Thallus P+ yellow, orange, or red (argopsin, fumarprotocetraric, pannarin, protocetraric, psoromic, stictic or thamnolic acid present)………………………………………………...... ……..18 18. Thallus P+ intense yellow (psoromic acid or thamnolic acid present)……...... ……………....19 19. Thallus K+ yellow (thamnolic acid); granules without a distinct pseudocortex; extreme southeastern Coastal Plain (Florida); corticolous...... L. aurescens 19. Thallus K-; granules with a distinct pseudocortex; widely distributed outside of the southeastern Coastal Plain (where it is absent)……………...... ……….L. caesioalba chemotype 33 18. Thallus P+ orange or red (argopsin, fumarprotocetraric, pannarin, protocetraric, or stictic acid present)…………………………………………...... ………………………………………...20 20. Thallus with distinct “crisped” marginal lobes that resemble Normandina; Appalachian- Ozarks…………...... ……………………………………………………….....L. normandinoides 20. Thallus without distinct “crisped” marginal lobes…………...... …………………………...21 21. High elevations of the Appalachians; granules extremely large (>1 mm) and “fluffy”...... …………………………………………………………………………………...... …...L. lanata 21. Not in the above habitat; granules smaller and appearance various……...... ………….....22 22. Terricolous in coastal soil communities in southern California…...... ………..L. terricola 22. Not terricolous in the above habitats…………………………...... ………………….....23 23. Thallus thin, without a well developed hypothallus...... 24 24. Corticolous on conifers; southeastern Coastal Plain………...... ………....L. friabilis 24. Not corticolous on conifers in the southeastern Coastal Plain……...... …………...25 25. Thallus distinctly lead-gray or blue-gray; always in sheltered habitats; saxicolous or rarely muscicolous (occasional corticolous collections are also known)…....L. adhaerens 25. Thallus greenish or blue-white; in various habitats and on various substrates...... 26 26. Thallus greenish-blue, saxicolous in exposed habitats in southern California…...... L. santamonicae 26. Thallus blue-white or white……………...... ………………………………...27 27. Granules with a distinct pseudocortex; typically saxicolous in exposed habitats and rarely corticolous……...... L. caesioalba (various chemotypes) 27. Granules without a distinct pseudocortex; typically saxicolous in sheltered habitats…………………...... …………………………….....L. elobata 23. Thallus thick, with a well developed hypothallus………...... ………………………..28 28. Common everywhere except southern California and Florida; thallus light greenish or bluish…………………………………………...... ……...L. lobificans auct. 28. Rare except in the Pacific Northwest; thallus dark green or white…...... ………....29 29. Thallus K+ yellow (atranorin present)……………...... ………………....L. nivalis 29. Thallus K- (atranorin absent)…………………………………...... ……...... L. torii 17. Thallus P-...……………...... ……………………………………………………………………..30 30. Coastal southern California; thallus white and resembling Cladonia squamules….…...L. xerophila 30. Not in coastal southern California; thallus otherwise………………………………...... ……..31 31. Granules with a distinct pseudocortex…………...... members of the L. caesioalba- neglecta group (L. alpina, L. borealis, L. caesioalba roccellic acid chemotype, L zeorinica) 31. Granules without a distinct pseudocortex; not members of the above complex……...... ………32 32. Thallus with a distinctly yellowish hue (dibenzofurans present)………………...... ……….. ………………………………………………...... …..L. diffusa, L. membranacea, L. vouauxii 32. Thallus blue-white or greenish, not with a yellowish hue (dibenzofurans absent)…...... 33 33. Eastern temperate……………………………………………………...... …....L. caesiella 33. Boreal/western……………………………………………………...... ……………...…34 34. Thallus greenish; granules without long projecting hyphae……...... ………….L. jackii 34. Thallus grayish; granules with long projecting hyphae………...... …………L. rigidula