PART 1

PART 1 

INTRODUCTION 

Sadly this introduction is a litany of apologies. This second part appears almost five years to the day 

after the publication of the first, "Some Florida Lichens". Its publication is occasioned by the Third Tuckerman 

Workshop in Gainesville, Florida as the first was by the meeting of The American Bryological and 

Lichenological Society in Wakulla Springs, Florida. Obviously I had hoped to have a second part finished 

much earlier but without an absolute deadline such as the Tuckerman Workshop, procrastination rules. I 

badly underestimated the time necessary to produce the manuscript, deal with accumulated specimens, 

changes and errors. As a result much has been left unfinished. The presentation is only a skeleton intended 

to document Florida's lichen diversity, provide names by which it can be manipulated and encourage further 

investigations. It is definitely not "user friendly", especially for nonprofessionals. Descriptions, bibliography, 

illustrations and maps are not included although a glimpse of the future is included in the form of computer 

generated maps for Calicium. Human limitations have also led me to confine the scope primarily to materials 

held by The New York Botanical Garden. Study of specimens from other herbaria and new collections will 

make this preliminary survey obsolete within a matter of months. (A prime example is the Graphidaceae 

where the species curve shows no sign of leveling off.) This will be exacerbated by the rapidly changing 

generic concepts in crustose lichens resulting from release from the constraints of artificial systems and the 

introduction of new characters and the techniques to study them. 

"More Florida Lichens" consists of two parts. The first is mostly the keys from "Some Florida Lichens" 

with corrections and a few additions plus a few new family treatments. It is assumed that most users of "More 

Florida Lichens" have the first part and therefore, in order to save space, the reprinted keys are in 8pt type with 

additions and corrections in bold face. The second part is an expanded treatment of the pyrenolichens. It is 

given a separate section and called a" tour" since it is not confined to Florida taxa and like a tour covers 

whatever seems relevant, hitting the high points and avoiding the slums. It is essentially a complete treatment 

of the corticolous pyrenolichens for eastern North America. However, for some families that are special 

favorites, the digressions are global. 

As always I wish to thank my companion, Bill Buck, for moral support as well as editing and indexing. 

Dana Griffin, III is thanked for arranging many pleasant field trips and making numerous interesting specimens 

available. I am grateful to the Florida Department of Natural Resources for permission to collect in a number 

of state parks and preserves. 

ARTHONIALES 

1

OPEGRAPHACEAE Stizenb. 

KEY TO GENERA 

1. Exciple or hypothecium dark; exciple not byssoid; hypothecium, if dark, usually 

confluent with exciple ................................................................................................................................ 2 

2. Ascomata not aggregated, not in raised stroma-like structures............................................................3 

3. Ascoma elongated, lirelliform; ascospores with a gelatinous sheath............................................... 4 

4. Ascoma not pruinose; ascus of vulgata-type ............................................................. Opegrapha 

4. Ascoma pruinose; ascus of grumulosa-type ........................................................ [Lecanographa] 

3. Ascoma rounded, apothecioid; ascospores with or without a sheath .............................................. 5 

5. Asci and ascospores not fragile; paraphysoids tightly interconnected, 

not separating easily; ascus abietina- or grumulosa-type ............................................................ 6 

6. Ascus abietina-type, endoascus strongly hemiamyloid with conspicuous 

apical ring; ascospores without sheath ................................................................................... 7 

7. Ascoma not pruinose or disk with bright colored pruina 

[orange, yellow or greenish], KOH+ yellow or purple; exciple 

carbonaceous, hyphae not distinguishable in KOH............................................Cresponea 

7. Ascoma pruinose; pruina the same color as thallus, KOH-; 

exciple dark brown, hyphae in outer part distinguishable in KOH ......................Lecanactis 

6. Ascus grumulosa-type, endoascus weakly hemiamyloid with apical 

ring little or not differentiated; ascospores with a sheath ................................. [Lecanographa] 

5. Asci and ascospores fragile; paraphysoids dichotomously branched, 

easily separating; ascus of Bactrospora-type...........................................................Bactrospora 

2. Ascomata aggregated or in raised stroma-like structures..................................................................... 8 

8. Ascoma perithecioid, in weakly raised stroma-like structures; 

ascospores 4-celled, 23-27 × 2-3 µm; roccellic acid; 

Duval and Putnam counties ......................................................................Chiodecton malmei Thor 

8. Ascomata with exposed disk, not perithecioid; PD+ or C+ 

(protocetraric acid, psoromic acid or erythrin ..................................................................................... 9 

9. Thallus C-, PD+ yellow or red orange .......................................................................................... 10 

10. Thallus PD+ red orange (protocetraric acid); ascomata asteriform, 

in raised "stroma" which is constricted at base; disk brown, 

white pruinose; ascospores 4-celled, 50-70 × 4 µm; 

peninsular Florida............................Syncesia [treated in 1990 as Chiodecton sp. Buck 3254] 

10. Thallus PD+ yeliow (psoromic acid); ascomata round, 

in rows .................................................................................Enterographa quassiaecola Fée 

9. Thallus C+ red (erythrin), PD-; ascomata asteriform in raised "stroma" 

which is constricted at base; disk light to dark brown, white pruinose; 

ascospores 4-celled, tapered, 50-70 × 4-5 µm; Duval County ........................................................ 

2

..............................................................Syncesia [treated in 1990 as Chiodecton perplexum Nyl.] 

1. Exciple not evident or byssoid (when pruina dissolved in KOH); hypothecium colorless 

or light brown and small, confined to base of ascoma ............................................................................ 11 

11. Hypothecium light brown, small, only at base of ascoma.............................................................. 12 

12. Exciple not evident; epithecium not byssoid; ascomata immersed, asteriform; 

disk exposed, orange or red brown; ascospores 4-celled, 15-20 × 5 µm; 

psoromic acid; Duval, Leon and St. Lucie counties .................................................................... 

................................................................................ Sclerophyton inscriptum (Nyl.) Müll. Arg 

12. Ascomata densely pruinose; exciple and epithecium pale, byssoid 

hypothecium brown; ascospores 4-celled, ± clavate, ca. 24-31 × 6 µm; 

Bay County.......................................................................................... Genus? sp. Harris 35756 

11. Hypothecium colorless ................................................................................Enterographa/Mazosia 

BACTROSPORA Massal. 

Since "Some Florida Lichens" was published Egea and Torrente have revised the genus 

(Lichenologist 25: 211-255. 1993). Their study has resulted in the reduction of one of the Florida species to 

synonymy and the addition of one species to the mycota. 

1. Ascospores narrow, 3-4 µm wide.............................................................................................................. 2 

2. Ascospores long, 35-130 µm ................................................................................................................ 3 

3. Ascospores 12-26-celled; exciple or exciple and epithecium containing 

small granules; tissues IKI-, KOH/IKI + bluish................................................................................... 4 

4. Ascomatal margin smooth; ascospores 12-20-septate, 

47-85(-90) × 2-3(-3.5) µm; Collier, Duval, Lee, Levy and 

Polk counties ........................................................................ B. myriadea (Fée) Egea & Torrente 

4. Ascomatal margin with sparse, small teeth; ascospores 

70-130 × 2-2.5(-3) µm; Dade, Levy and Monroe 

counties ........................................................................ B. denticulata (Vainio) Egea & Torrente 

3. Ascospores 10-celled, 35-50 × 3 µm; exciple and epithecium lacking 

granules; parts of exciple and base of hymenium IKI+ bluish, 

KOH/IKI+ deep blue; thallus IKI+ bluish; common, south to 

Sarasota County......................................................................................B. mesospora R. C. Harris 

2. Ascospores short, (4-)6-celled, 22-27 × 3-4 µm; on lignum or bark 

of Taxodium; Franklin, Liberty and Wakulla counties...................................B. brevispora R. C. Harris 

1. Ascospores broad, 18-20-celled, 80-90 × 7-10 µm; Calhoun, 

Duval and Nassau counties........................................................................... B. macrospora R. C. Harris 

NOTES 

Bactrospora denticulata (Vainio) Egea & Torrente 

3

Bactrospora integrispora Seaver, Mycologia 16: 8. 1924. Type. VIRGIN ISLANDS. St. Thomas: on 

decorticated wood, 10 Mar 1923, Seaver 754 (NY, holotype), syn. nov. 

Seaver's name, appearing in a mycological journal, was not picked up by Zahlbruckner and thus not 

included by Egea & Torrente. (Also omitted, Seaver's report of Bactrospora dryina (Ach.) Massal. from North 

Dakota (Mycologia 1: 112. 1909.) is Schizoxylon berkelyeanum (Dur. & Lév.) Fuckel s. lat.) The Aptroot 

specimen from Key Largo was incorrectly assigned to Dade County instead of Monroe County in Egea and 

Torrente (1993). 

Bactrospora macrospora R. C. Harris 

Henssen in Henssen and Thor (1990) transferred this species to Melampylidium Stirton. I am 

uncomfortable with this on several grounds. First, the decision is based on material from North Carolina which 

the authors admit differs from my description of B. macrosporum. Second, I am cited for pointing out the 

different ascospore type. This is true but this was before Egea and Torrente's (1993) treatment of 

Bactrospora and I was simply ignorant of the range of ascospore variation in the genus. I did comment on the 

identity of the ascomatal type within Bactrospora. Third, Egea and Torrente (1995) present evidence for 

synonymizing Melampylidium with Bactrospora. Until more convincing evidence for recognizing 

Melampylidium and B. macrospora as a member of the genus, I prefer to retain the species in Bactrospora. 

Bactrospora nematospora R. C. Harris = B. myriadea 

CHIODECTON Ach. 

Only a single species, C. malmei Thor, is in North America, and it is diagnosed in the key to genera. 

Two other species C. perplexum and sp. Buck 3254 would seem to belong in Syncesia Taylor (Tehler, 1993) 

but there is as yet no treatment of the species. 

CRESPONEA Egea & Torrente 

This genus was recently separated from Lecanactis and revised by Egea and Torrente (Mycotaxon 

48: 301-331. 1993). They reported C. proximata from Florida. 

1. Ascospores 3-4(-5)-septate, 15-22(-24) × (4-)4.5-5.5 µm; margin black, raised, 

cracked; disk chartreuse pruinose; Alachua County........................... C. flava (Vainio) Egea & Torrente 

1. Ascospores 5-7(-8)-septate, 25-38(-40) × 5-7(-7.5) µm; margin black, thick, 

slightly raised, ± smooth; disk chartreuse pruinose; Monroe County (Keys)............................................... 

....................................................................................................... C. proximata (Nyl.) Egea & Torrente 

NOTES 

Cresponea flava (Vainio) Egea & Torrente 

This species is close to C. chloroconia (Tuck.) Egea & Torrente. They annotated a Calkins collection, 

purportedly from Florida, as C. chloroconia but did not cite it in the publication. The Cresponea is associated 

with sterile Chaenotheca furfuracea (L.) Tibell and is surely not from Florida. 

Cresponea flava is new to North America. FLORIDA. Alachua County: Paynes Prairie State Preserve, 

vicinity of Alachua Sink, 29°37'N, 82°18'W, mixed hardwoods over limestone, 1 Dec 1992, Harris 29432 (NY). 

ENTEROGRAPHA Fée & MAZOSIA Massal. 

4

1. Ascomata immersed or emergent; not resembling Byssoloma; margin not white .................................... 2 

2. Growing on bark .................................................................................................................................... 3 

3. Ascomata immersed rounded or lirelline.......................................................................................... 4 

4. Ascospores 8-celled...................................................................................................................... 5 

5. Ascomata rounded, arranged in short rows or irregular groups, 

surrounding area somewhat raised in age; psoromic acid; 

ascospores 34-37 × 5-6 µm, with thick gelatinous sheath; 

Monroe County (Keys)...............................................................................E. quassiaecola Fée 

5. Ascomata solitary, immersed, with slightly raised margin, lirelliform 

to weakly asteriform; disk yellowish; ascospores 26-35 × 4.5 µm; 

TLC not done; Collier County ....................................................Enterographa sp. Buck 22882 

4. Ascospores 4-celled; ascomata scattered; no substances; 

throughout Florida ................................................................ Mazosia ocellata (Nyl.) R. C. Harris 

3. Ascomata emergent, Lecanora-like or weakly lirelline, with distinct thalline margin; 

disk pale yellowish to brown, weakly pruinose; ascospores 7-8-celled, median cells 

occasionally somewhat swollen, 30-45 × 3-4 µm; psoromicacid; Franklin, Gulf and 

Okaloosa counties................................................................................E. lecanoroides R. C. Harris 

2. Growing or limestone; thallus thick, pale buff brown; ascomata immersed; 

disk dark, rounded; ascospores 20-24 × 5-5.5 µm; Monroe County 

(Key West) ........................................................................................... Enterographa sp. Thaxter 415 

1. Ascomata minute, resembling Byssoloma; margin white; disk brown, 

white pruinose; ascospores 6(-7)-celled,18-22 × 3-3.5 µm; Seminole 

County ...........................................................Enterographa [Schismatomma rappii (Zahlbr.) R C. Harris] 

LECANACTIS Fr. 

1. Ascospores 6-8-celled, 30-40 × 3-4 µm; ascomata gray brown, white pruinose, 

strongly constricted at base, round, irregular or slightly elongate; schizopeltic acid; 

Duval County.......................................................................................................L. epileuca (Nyl.) Tehler 

1. Ascospores 4-celled, 20-24 × 4 µm; ascomata as above; schizopeltic acid; 

Duval County................................................................................................... Lecanactis sp. Calkins 52 

NOTES 

Lecanactis subattingens (Nyl.) R. C. Harris 

= L. epileuca (Nyl.) Tehler. See Egea & Torrente (Biblioth. Lichenol. 54: 1-205. 1994). 

OPEGRAPHA Ach. 

Although badly in need of revision, the key to Opegrapha is merely reprinted from 1990. 

1. Ascospores 4-celled.................................................................................................................................. 2 

5

2. Lirellae black or white pruinose, not pigmented .................................................................................... 3 

3. Lirellae sessile, black, shiny, not pruinose .......................................................................................... 4 

4. Ascospores rounded at one or both ends ....................................................................................... 5 

5. Lirellae smaller, short, fusiform or once branched, rarely asteriform ..........................................6 

6. Hymenium not inspersed; ascospores 13-15 × 4-5 µm; 

south to Manatee County .....................................................................................O. atra Pers. 

6. Hymenium inspersed; disk exposed; ascospores 

13-15 × 4-4.5 µm; Duval County ................................................. Opegrapha sp. Calkins 604 

5. Lirellae coarse, elongate, branched; ascospores 13-15 × 4-5 µm; 

Collier, Dade and Duval counties .............................................................. O. candida Müll. Arg. 

4. Ascospores tapering at both ends, 18-19 × 

4-4.5 µm; lirellae small; disk not exposed; 

Okeechobee and Polk counties ...........................................................Opegrapha sp. Britton 819 

3. Lirellae initially immersed in bark of Taxodium; disk a strongly flexuose 

slit becoming quite broadly exposed; margin thick, black or densely white 

pruinose; ascospores 25-28 × 4-5 µm, cells ± equal; Franklin and Liberty 

counties ......................................................................................................... O. cypressi R. C. Harris 

2. Lirellae encrusted with bright orange pigment, short, straight; 

ascospores 17-20 × 4 µm; Collier County ................................................... O. aurantiaca B. de Lesd. 

1. Ascospores 6-14-celled............................................................................................................................. 7 

7. Growing on bark or wood ...................................................................................................................... 8 

8. Disk not covered by dense white layer .............................................................................................. 9 

9. Disk broadly expanded, red brown.............................................................................................. 10 

10. Ascospores 5-8-celled, 16-20 × 4-5 µm; lirellae long, flexuose, 

branched; margin not carbonized below hymenium; south to 

Polk County ....................................................................................... O. longissima Müll. Arg. 

10. Ascospores 9-12-celled, 38-50 × 4 µm; lirellae short, orbicular 

or elongate and strongly flexuose; margin carbonized below 

hymenium; Dade County...................................................................Opegrapha sp. Britton 2 

9. Disk hidden or if exposed, then black ......................................................................................... 11 

11. Ascospores 6-celled; lirellae fusiform or once branched ...................................................... 12 

12. Ascospores narrow, 3-4 µm wide; disk hidden................................................................. 13 

13. Thallus whitish, continuous or rimose, 

sometimes scanty; ascospores 24-30 × 3-4 µm; 

south to Polk County ............................................................................ O. vulgata Ach. 

13. Thallus white, thick, matte, almost byssoid; 

6

ascospores 18-24 × 3 µm; Louisiana, 

not yet found in Florida...........................................................[O. leucoplaca Müll. Arg.] 

12. Ascospores broader, 4.5-6 µm wide; disk becoming exposed, black .............................. 14 

14. Hymenium not inspersed; ascospores18-20 × 5-6 µm; 

Duval County........................................................................................... O. varia Pers. 

14. Hymenium inspersed; ascospores 20-27 × 4.5-5 µm; 

Southwest Florida, Mar 1878, Smith & Austin 82.............................Opegrapha sp. 82 

11. Ascospores 8-14-celled......................................................................................................... 15 

15. Ascospores narrow, 3-4 µm wide.................................................................................... 16 

16. Ascospores 12-14-celled, 30-55 × 3-4 µm; thallus dark; 

disk hidden; lirellae fusiform or once branched; south to 

Polk County ..................................................... O. viridis (Ach.) Behlen & Desberger 

16. Ascospores 8-celled, 32-43 × 3 µm; thallus pale; 

disk hidden; lirellae branched, flaring out at base; 

Dade and Monroe counties ................................................ Opegrapha sp. Small 471 

15. Ascospores broader, 5-7 µm wide .................................................................................. 17 

17. Hymenium not inspersed; disk becoming exposed; 

lirellae small, fusiform to linear, rarely once branched; 

ascospores 7-8-celled, 35-40 × 5-7 µm; Dade County.................................................. 

.........................................................................................Opegrapha sp. Britton 799 

17. Hymenium inspersed; disk hidden; lirellae coarse, 

simple to weakly asteriform; ascospores 7-8-celled, 

40-45 × 5-6.5 µm; Collier County .....................................Opegrapha sp. Britton 652 

8. Disk becoming broadly expanded, covered by a dense layer of white 

material; margin black, usually lacking at base; lirellae long, flexuose, 

branched; ascospores rarely produced?, (4-)6-celled, ca. 24 × 5 µm; 

Duval, Polk and Putnam counties ........................................................................... O. astraea Tuck. 

7. Growing on bamboo culms or rock ...................................................................................................... 18 

18. Growing on bamboo culms (normally a foliicolous species); 

ascospores narrow, 4-6-celled, 18-25 × 3-4 µm; Louisiana, 

not seen from Florida ..............................................................................................[O. filicina Mont.] 

18. Growing on limestone; lirellae sessile, fusiform to linear, 

not branched; ascospores 5-6-celled, 20-22 × 4 µm; Keys................................................................ 

..............................................................................................................Opegrapha sp. Britton 869 

SCHISMATOMMA Massal. 

Tehler (1993) has removed S. rappii, the only Florida species, from Schismatomma s. str. and 

suggested that it be placed provisionally in Enterographa s. lat. 

7

SCLEROPHYTON Eschw. 

Only the single species, S. inscriptum (Nyl.) Müll. Arg., is in Florida. Egea & Torrente (Bryologist 98: 

207-217. 1995) have revised Sclerophyton s. lat. for North America. They provide excellent illustrations. In 

1990 I suggested that Sclerophyton should be restricted to the hyaline spored species with dendroid, 

immersed ascomata and I am still of this opinion. Egea and Torrente retained the old, broad definition, 

recognizing that there are two distinct groups in the genus so constituted, but provided no reasons as to why 

they should be joined in a single genus. Superficially S. inscriptum (North America) and S. elegans Eschw. 

(Brazil) are identical and both contain psoromic acid. My first thought was to synonymize them. Egea and 

Torrente suggested that they are conspecific but did not formally make the synonymy. However, examination 

of an isotype of S. elegans (H-Nyl 4583) revealed that the ascospores are slightly smaller (13-14.5 × 4-5 µm) 

than in S. inscriptum, (15-20 × 5 µm). Pending study of more material, I agree the two should be kept 

separate. 

GRAPHIDALES 

GRAPHIDACEAE Dumortier 

I had hoped to completely revise this family. However, I have had to be content with a few minor additions and corrections. I 

now have collections of at least a dozen species not treated in 1990 and expect that more will turn up, making this one of the most 

speciose families among the lichen-forming fungi in Florida. 

KEY TO GENERA 

1. Ascomata immersed to sessile, mostly elongated with a narrow, often slit-like disk; 

not aggregated in a pseudostroma ......................................................................................................................................................... 2 

2. Spores colorless................................................................................................................................................................................... 3 

3. Spores transversely septate...................................................................................................................................................Graphis 

3. Spores submuriform to densely muriform........................................................................................................................... Graphina 

2. Spores brown ....................................................................................................................................................................................... 4 

4. Spores transversely septate....................................................................................................................................... Phaeographis 

4. Spores submuriform to densely muriform................................................................................................................ Phaeographina 

1. Ascomata with brown disk, cup-like or aggregated in a pseudostroma................................................................................................. 6 

5. Ascomata cuplike with a thick raised margin; disk brown; hymenium not inspersed; 

ascospores colorless to weakly tinted, muriform with 9-11 longitudinal cells, 

33-48 × 13-16 µm; no substances; throughout Florida......................................................Gyrostomum scyphuliferum (Ach.) Nyl. 

5. Ascomata aggregated in a pseudostroma.......................................................................................................................................... 6 

6. Spores muriform, colorless becoming brownish in age, 35-50 × 8-9 µm; 

pseudostroma well developed, usually gray pruinose; hymenium not inspersed; 

no substances; throughout Florida......................................................................................................... Glyphis cicatricosa Ach. 

6. Spores transversely 4-6-celled, brown; pseudostroma poorly developed, 

usually whitish............................................................................................................................................................. Sarcographa 

GLYPHIS Ach. 

Only a single species in North America. Redinger (1934) in his treatment for Brazil synonymized the names which also appear in 

the North American list, choosing Glyphis cicatricosa Ach. from among the competing synonyms. See generic key for diagnosis. 

GRAPHINA Müll. Arg. 

1. Lirellae not carbonized ............................................................................................................................................................................ 2 

8

2. Ascospores 1(-2)/ascus....................................................................................................................................................................... 3 

3. Norstictic acid present; ascospores 80-115 × 24-38 µm; 

Alachua, Duval, Levy and Putnam counties ........................................................................................................Graphina n. sp. 

3. No lichen substances detected;thallus rough, with a lumpy, scurfy look; 

lirellae crowded, short and irregular; disk often exposed; ascospores 

85-105 × 20-28 µm; south to Marion County ................................................................................................ G. cypressi Müll. Arg. 

2. Ascospores 2-8/ascus.......................................................................................................................................................................... 4 

4. Thallus UV-........................................................................................................................................................................................ 5 

5. Ascospores longer than 40 µm and/or wider than 20 µm............................................................................................................. 6 

6. Lichen substances present (usually PD+) ................................................................................................................................. 7 

7. Protocetraric acid; thallus pale gray; lirellae white, emergent; ascospores 

2-4/ascus, 110-140 × 15-17 µm; south to Lake County.......................................................G. abaphoides (Nyl.) Müll. Arg. 

7. Norstictic acid or stictic acid satellites; ascospores at least initially 8/ascus ....................................................................... 8 

8. Stictic acid satellites; lirellae paler than thallus ................................................................................................................. 9 

9. Hypostictic and hyposalazinic acids; thallus dull, pale green gray; lirellae 

white, immersed, elongate and branched; ascospores 40-70 × 13-17 µm; 

north to Lake Okeechobee........................................................................................... G. virginea (Eschw.) Müll. Arg. 

9. Constictic acid; lirellae "mealy" due to multiple crumbling layers of exciple; 

ascospores 35-78 × 18-36 µm [2-4/ascus according to Wirth & Hale 

but U.S. material initially 8/ascus]; Seminole County ....................................................... G. peplophora Wirth & Hale 

8. Norstictic acid; lirellae concolorous with thallus, crowded, irregular; 

ascospores 60-85 × 13-17 µm; Duval and Levy counties.........................................G. subvirginalis (Nyl.) Müll. Arg. 

6. Lichen substances absent; lirellae crowded, irregular with white crumbling exciple; 

ascospores IKI-, 35-78 × 18-36 µm; Seminole County............................................................... G. leuconephela auct. Amer. 

5. Ascospores shorter than 40 µm and/or narrower than 20 µm.................................................................................................... 10 

10. Ascospores 1-2 × 4(-6)-celled, 15-20 × 7-9 µm .................................................................................................................... 11 

11. Lirellae fissurine; disk not exposed; no substances; 

peninsular Florida..............................................................................................................G. incrustans (Fée) Müll. Arg. 

11. Lirellae long, regularly branched; disk exposed, reddish; norstictic acid; 

Collier County ................................................................................................................ G. floridana (Tuck.) R. C. Harris 

10. Ascospores more than 1-2 × 4-celled, over 20 µm long....................................................................................................... 12 

12. Thallus PD- (no subsances)............................................................................................................................................ 13 

13. Exciple well developed, yellow to orange, darker in KOH; 

lirellae raised, elongate, flexuose; ascospores 2-3 × 6-8-celled, 

22-32 × 10-11 µm; Duval County ........................................................................................ G. scolecitis (Tuck.) Fink 

13. Exciple not evident; lirellae short,crowded; disk becoming exposed, white 

pruinose; ascospores 2-4 × 6-8-celled, 22-24 × 7-9 µm; Marion County....................................Graphina sp. 23599 

12. Thallus and lirellae PD+ yellow (psoromic acid); ascospores elliptical, 

IKI+ dark violet, 20-30(-40) × 13-19 µm; often sterile but combination 

of fissurine lirellae and psoromic acid is diagnostic; south to Lake 

County.......................................................................................................................G. columbina (Tuck.) Wirth & Hale 

4. Thallus UV+ yellow (lichexanthone); lirellae imnersed; disk pale red brown; 

margin white, patchily developed; ascospores 35-50 × 12-15 µm, found only 

in Liberty County specimen, sterile from Duval and Wakulla counties but could be 

Graphis "anguinoides"....................................................................................................................... G. anguina (Mont.) Müll. Arg. 

1. Lirellae carbonized, at least patchily so or at apices of exciple............................................................................................................ 14 

9

14. Ascospores 2-8/ascus...................................................................................................................................................................... 15 

15. Exciple carbonized to base of hymenium ................................................................................................................................... 16 

16. Exciple not striate ................................................................................................................................................................... 17 

17. Exciple covered with thin white layer; salazinic acid; ascospores 8/ascus, 

1-2 × 4-5-celled, 12-16 × 5-8 µm; Dade County ................................................................ G. marescens (Fée) Müll. Arg. 

17. Exciple naked, black; lirellae elliptical toweakly elongate with broad 

red brown disk; ascospores 2-4/ascus, 35-50 × 17-25 µm; 

no substances; Lake County..................................................................................................................Graphina sp. 1271 

16. Exciple becoming striate, bleeding purplish in KOH; norstictic acid; 

ascospores 2-4 × 6-8-celled, 20-35 × 10-15 µm; Dade County......................................................Graphina sp. Britton 680 

15. Exciple only carbonized at apices, becoming striate.................................................................................................................. 18 

18. Ascospores 2-4/ascus, 30-50 × 20-27 µm; norstictic acid; 

exciple carbonized almost to base but inner part yellowish; 

Florida, without locality ...............................................................................................G. antillarum (Vainio) Zahlbr. 

18. Ascospores 8/ascus, 30-40 × 15-17 µm; stictic acid; Louisiana, 

not yet known from Florida ...................................................................................... [G. parilis (Krempelh.) Müll. Arg.] 

14. Ascospores 1/ascus ......................................................................................................................................................................... 19 

19. No substances; ascospores 65-90 × 20-30 µm; lirellae semi-immersed; 

disk narrow, not pruinose; margin black, shiny, not striate, open below; 

throughout Florida ................................................................................................................................G. xylophaga R. C. Harris 

19. Protocetraric acid; ascospores 120-165 × 35-40 µm; lirellae immersed, 

disk broad, whitish pruinose; margin pruinose, often only patchily carbonized; 

Collier County ....................................................................................................................................G. platyleuca (Nyl.) Zahlbr. 

NOTES 

Graphina adscribens Nyl.) Müll. Arg. 

I have now examined the type of this species (H-Nyl 6824) and it does not match anything in North America. The Florida 

collection in Nylander's herbarium under this name is Graphina scolecitis. Thus Graphina adscribens can be excluded from the 

North American mycota with confidence. 

Graphina columbina (Tuck.) Wirth & Hale 

The lead for this species was inadvertenly omitted from "Some Florida Lichens". 

Graphina platyleuca (Nyl.) Zahlbr. 

New to North America. FLORIDA. Collier County: Collier-Seminole State Park, just W of boat basin, 25°59'N, 81°35'W, 

mangroves at edge of salt marsh, 7 Dec 1992, Harris 30065 (NY). 

Graphina subvirginalis (Nyl.) Müll. Arg. 

I was badly confused in 1990, mixing two obviously distinct taxa. Graphina subvirginalis has a variable number of 

ascospores, 2-8/ascus (type, H-Nyl 6851) and another undescribed species has only 1/ascus. I have not had time to re-examine 

material but I presume that Graphina intertexta (Müll. Arg.) R. C. Harris will prove to be a synonym of G. subvirginalis. 

GRAPHIS Adanson 

1. Exciple carbonized; spores 4-celled only in G. afzelii............................................................................................................................. 2 

2. Margin not striate; hymenium inspersed or not ................................................................................................................................... 3 

3. Ascomata sessile, covered with thin white layer; exciple well developed below hymenium .......................................................... 4 

4. Lirellae C+ red (lecanoric acid); ascospores 4-celled, 16-23 × 6-9 µm; 

common, throughout the state .................................................................................................................................G. afzelii Ach. 

4. Lirellae and thallus UV+ yellow (lichexanthone), PD+ red (protocetraric acid); 

ascospores ca. 22-23-celled, 60-95 × 7-9 µm; Highlands County .....................................................................G. candidata Nyl 

10

3. Ascomata immersed or sessile, black, not covered by a white layer.............................................................................................. 5 

5. Epihymenium colorless, KOH- ..................................................................................................................................................... 6 

6. Exciple lacking below hymenium ............................................................................................................................................. 7 

7. Lichen substances absent.................................................................................................................................................... 8 

8. Ascospores 8-11-celled, 25-40 µm long .......................................................................................................................... 9 

9. Disk not exposed; exciple carbonized to base of hymenium; 

hymenium not inspersed; ascospores 8-10-celled, 25-40 × 7-9 µm; 

Columbia, Duval and Lake counties ..................................................................................................... G. lineola Ach. 

9. Disk becoming exposed, whitish or bluish pruinose; exciple 

mostly not carbonized to base; hymenium not inspersed; 

ascospores 8-11-celled, 30-40 × 7-9 µm; Duval County ............................................................... G. scripta (L.) Ach. 

8. Ascospores 11-13-celled, 50-70 × 8-11 µm; hymenium inspersed; 

Duval, Franklin and Highlands counties .............................................................................................. G. pavoniana Fée 

7. Norstictic, stictic or protocetraric acids............................................................................................................................... 10 

10. Norstictic acid only....................................................................................................................................................... 11 

11. Lirellae flexuose, branching, sessile to semi-immersed; 

hymenium not inspersed; ascospores 6-8-celled, 18-35 × 5-9 µm; 

peninsular Florida............................................................................................................................ G. librata Knight 

11. Lirellae short, straight, immersed; hymenium inspersed; 

ascospores 8(-9)-celled, 30-35 × 8-9 µm; Dade County..................................................... Graphis sp. Britton 664 

10. Stictic acid (with or without norstictic acid) or protocetraric acid.................................................................................. 12 

12. Lirellae immersed; margin and disk white pruinose; hymenium 

not inspersed; ascospores 8-11-celled, 25-35 × 7-8 µm; stictic acid 

or protocetraric acid; Dade and Palm Beach counties ....................................................... G. caesiella Vainio s. lat 

12. Lirellae semi-immersed, black; margin and disk not pruinose; 

hymenium inspersed; ascospores 10-11-celled 30-40 × 7-8 µm; 

norstictic plus stictic acid; Dade County.............................................................................. G. leptocarpa Fée s. lat. 

6. Exciple entire, carbonized below hymenium.......................................................................................................................... 13 

13. Ascospores 11-15-celled, 50-60 × 8-10 µm..................................................................................................................... 14 

14. Hymenium inspersed; lirellae mostly immersed; norstictic acid (rarely 

acid deficient); throughout the state................................................................................................. Graphis sp. 23509 

14. Hymenium not inspersed; lirellae sessile; no substances; Polk County................................. G. anfractuosa Eschw. 

13. Ascospores 6-8-celled, 18-25 × 6-7 µm; hymenium inspersed; norstictic acid; 

Duval and Polk counties......................................................................................G. desquamescens (Fée) Zahlbr. s. lat 

5. Epihymenium orange, KOH+ purple; margin not striate; disk exposed, orange....................................................................... 15 

15. Exciple closed; hymenium yellowish inspersed; ascospores 8-celled, 

27-34 × 7-8 µm; norstictic acid; Dade County..........................................................................G. chromothecia R. C. Harris 

15. Exciple lacking below; hymenium not inspersed; ascospores 8-celled, 

23-30 × 6-7 µm; no substances; Franklin and Liberty counties..........................................................G. inversa R. C. Harris 

2. Margin striate; hymenium not inspersed............................................................................................................................................ 16 

16. Thallus UV+ yellow (lichexanthone)............................................................................................................................................. 17 

17. Lirellae narrow, flexuose, branched; margin weakly striate; ascospores 

6-8-celled, 20-30 × 7-9 µm; south to Polk County ................................................................................ G. lucifica R. C. Harris 

17. Lirellae rather broad, short, often straight, sparsely branched; margin 

strongly striate; ascospores 11-14-celled, 50-75 × 9-12 µm; 

Franklin and Liberty counties ............................................................................................................... G. haleana R. C. Harris 

11

16. Thallus UV-................................................................................................................................................................................... 18 

18. Ascospores 14-19-celled, 55-80 µm long .............................................................................................................................. 19 

19. Stictic acid; exciple strongly developed below; ascospores ca. 14-celled, 

55 × 8-9 µm; Southwest Florida, Mar 1878, Smith & Austin 91.................................................................. Graphis sp. 91 

19. Norstictic acid; exciple barely closed below; ascospores 16-19-celled, 

70-80 × 10-11 µm; Louisiana, not yet known from Florida ..................................................[G. lumbricina Vainio s. lat.] 

18. Ascospores 8-12-celled, 20-45 µm long ................................................................................................................................ 20 

20. Margin carbonized to base of hymenium.......................................................................................................................... 21 

21. No substances; ascospores 10-12-celled, 35-45 × 7-9 µm; 

Duval and Lake counties....................................................................................................G. striatula (Ach.) Sprengel 

21. Stictic acid; ascospores 8-celled, 23-28 × 6-7 µm; Leon County .................................................... Graphis sp. 25050 

20. Margin carbonized above, orange below; ascospores 9-10-celled, 

25-27 × 6-7 µm often sterile; no substances; Duval and Marion counties ............................................G. subelegans Nyl. 

1. Exciple not carbonized; ascospores 2-4-celled .................................................................................................................................... 22 

. 

22. Ascospores 4-celled; no substances ............................................................................................................................................... 23 

23. Lirellae fissurine; exciple not evident.......................................................................................................................................... 24 

24. Margin not swollen; ascospores to 17 × 7 µm ....................................................................................................................... 25 

25. Thallus white, ecorticate; lirellae not gaping, minute, visible only as 

thin lines; ascospores 15-17 × 6-7 µm; Duval and Polk counties.................................................G. illiterata R. C. Harris 

25. Thallus greenish, with shiny prosoplectenchymatous cortex; 

lirellae gaping; ascospores 11-13 × 5-6 µm; Levy County.................................................... G. subnitidula Nyl. ex Tuck. 

24. Margin swollen; ascospores 17-20 × 8-10 µm; Duval, Lake and 

Marion counties ......................................................................................................... G. insidiosa (Knight & Mitten) Hook. f. 

23. Lirellae long, dichotomously branched; disk hidden to exposed; 

exciple orangish, well developed laterally or at base ................................................................................................................. 26 

26. Disk not exposed; exciple well developed laterally ............................................................................................................... 27 

27. Margin not striate; ascospores 15-17 × 6-8 µm; 

Duval and Liberty counties........................................................................................................................G. subparilis Nyl. 

27. Margin striate; ascospores 22-24 × 10-13 µm; East Florida, 1877, Smith..................................................G. rufula Mont. 

26. Disk exposed; "exciple" well developed below hymenium; 

ascospores 10-13 × 5 µm; Collier, Dade and Osceola counties .................................................................G. grammatis Fée 

22. Ascospores 2-celled or 8-9-celled ................................................................................................................................................... 28 

28. Ascospores 2-celled; norstictic acid; thallus whitish, shiny, UV-; lirellae crowded, initially 

with closed ± pruinose lips, later disk exposed, pale, pruinose; Duval County................................................ G. turbulenta Nyl. 

28. Ascospores 8-9-celled, 21-31 × 7.5-9 µm; thallus whitish, dull, UV+ yellow (lichexanthone); 

lirellae imnersed; disk pale red brown; margin white, patchily developed; Bay, Jefferson and 

Lafayette counties ......................................................................................................................................G. anguinoides ined. 

NOTES 

Graphis "anguinoides" ined. 

This species is superficially indistinguishable from Graphina anguina. I had actually placed a couple specimens in this 

species on the external characters of ascoma and UV+ thallus. When I checked the ascospores of some recent collections, I 

discovered a taxon with transversely septate, not muriform, ascospores. To my mind this is another nail in the coffin of the 

ascospore genera, Graphis vs. Graphina. It is inconceivable to me that genera defined on the single character of ascospore 

septation should have evolved parallel species with identical multicharacter complexes of thallus and ascoma. 

12

GYROSTOMUM Fr. 

Only the single species, G. scyphuliferum (Ach.) Nyl. in North America although there are two additional apparently undescribed 

species in Puerto Rico. See generic key for diagnosis. 

PHAEOGRAPHINA Müll. Arg. 

1. Exciple not carbonized ............................................................................................................................................................................ 2 

2. Disk initially hidden, finally exposed; margin white, separating from thallus; 

margin and disk densely white pruinose; hymenium not inspersed; ascospores 

8/ascus, 2-4 × 8-celled, 27-35 × 10-13 µm; no substances; Highlands and 

Polk counties ........................................................................................................................................................... P. asteroides Fink 

2. Disk exposed from beginning; disk weakly white pruinose; hymenium not 

inspersed; ascospores 8/ascus, 1-2 × 6-celled, 20-30 × 10-11 µm; 

norstictic acid; Highlands County.............................................................................................................Phaeographina sp. 18150B 

1. Exciple carbonized, at least at apices..................................................................................................................................................... 3 

3. Lirellae immersed or if emergent, not white......................................................................................................................................... 4 

4. Disk broad, bluish pruinose or not pruinose .................................................................................................................................... 5 

5. Lirellae immersed; disk not or weakly pruinose; exciple usually carbonized below; 

hymenium inspersed; ascospores 8/ascus, 2-4 × 6-8-celled, 24-30 × 9-12 µm; 

no substances; Duval and Lake counties ..........................................................................................................P. explicans Fink 

5. Lirellae emergent, large; disk usually bluish pruinose; exciple carbonized only 

at apex; hymenium inspersed; ascospores 8/ascus, muriform, 65-75 × 16-23 µm; 

no substances; south to Highlands County ..........................................................................P. caesiopruinosa (Fée) Müll. Arg. 

4. Disk narrow, white pruinose; margin black, carbonized only at edges; hymenium 

inspersed; ascospores 2-6/ascus, very variable, 35-50 × 13-24 µm; norstictic acid; 

Keys ..............................................................................................................................................Phaeographina sp. Small 10114 

3. Lirellae strongly emergent with thin white covering, cup-shaped, oval to weakly elongate 

and once branched; margin and base thick, heavily carbonized; hymenium inspersed; 

ascospores 8/ascus, 100-110 × 24-27 µm; no substances; Dade County............................. P. quassiaecola (Fée) Müll Arg. s. lat. 

PHAEOGRAPHIS Müll. Arg. & SARCOGRAPHA Fée 

1. Lirellae red; exciple not carbonized; ascospores 6-10-celled, 21-35 × 9-11 µm; 

Pinellas and Sarasota counties...............................................................................................................P. haematites (Fée) Müll. Arg. 

1. Lirellae white, brown or blackish ............................................................................................................................................................. 2 

2. Ascospores 4-celled ............................................................................................................................................................................. 3 

3. Ascomata not erumpent ................................................................................................................................................................... 4 

4. Hymenium inspersed..................................................................................................................................................................... 5 

5. Exciple entire, carbonized below .............................................................................................................................................. 6 

6. Lirellae simple to loosely asteriform, not surrounded by whitish "stroma"; 

ascospores 15-16 × 5-6 µm; no substances or rarely norstictic acid; 

south to Polk County .............................................................................................................P. sericea (Eschw.) Müll. Arg. 

6. Lirellae densely branched, surrounded by a weakly raised whitish "stroma"....................................................................... 7 

7. No substances; ascospores17-21 × 7-8.5 µm; Highlands County ..................... Sarcographa tricosa (Ach.) Müll. Arg. 

7. Stictic acid agg.; ascospores 16-21 × 5.5-6 µm; Dade County...................................................................................... 

.....................................................................................................................Sarcographa labyrinthica (Ach.) Müll. Arg. 

5. Exciple carbonized only at apex; hymenium inspersed; ascospores 16-21 × 6-10 µm; 

13

no substances; not yet found in Florida .........................................................................................[P. inusta (Ach.) Müll. Arg.] 

4. Hymenium not inspersed............................................................................................................................................................... 8 

8. Lirellae densely branched or crowded and irregular, surrounded by 

distinct whitish area ................................................................................................................................................................... 9 

9. Norstictic acid; whitish area weakly raised, not mealy; ascospores 15-20 × 5-6 µm; 

Polk and Seminole counties.......................................... Sarcographa intricans sensu Harris 1990 non (Nyl.) Müll. Arg. 

9. No substances; lirellae in mealy decorticate patches, brown, irregular, 

crowded ; ascospores 13-16 × 5-6 µm; Puntnam County .............................................................Phaeographis sp. 23697 

8. Lirellae simple to loosely branched, not surrounded by whitish area; 

ascospores 15-18 × 5-6 µm; norstictic acid or norstictic acid plus 

stictic acid or rarely acid deficient; Duval, Polk and Volusia counties.......................................P. subtigrina (Vainio) Zahlbr. 

3. Ascomata erumpent with more or less erect bark flaps ................................................................................................................ 10 

10. No substances; hymenium inspersed; ascospores 16-20 × 7-8 µm; 

Volusia County...............................................................................................................................P. subfulgurata (Nyl.) Zahlbr. 

10. Norstictic acid; hymenium not inspersed; ascospores 19-21 × 7-8 µm; 

Florida, without locality, Wilson...............................................................................................................Phaeographis sp. 1233 

2. Ascospores 6-10-celled....................................................................................................................................................................... 11 

11. Ascomata round to slightly elongated, not branched .................................................................................................................. 12 

12. Thallus UV+ yellow (lichexanthone); margin not raised; ascospores 6-celled, 

18-24 × 6-7 µm; south to Polk County............................................................................ P. punctiformis (Eschw.) Müll. Arg. 

12. Thallus UV-(no substances); margin often erect, jagged; ascospores 8-10-celled, 

25-40 × 7-10 µm; south to Polk County......................................................................................P. lobata (Eschw.) Müll. Arg. 

11. Ascomata elongate, often branched ............................................................................................................................................ 13 

13. Exciple not carbonized; hymenium not inspersed ................................................................................................................. 14 

14. Margin not striate............................................................................................................................................................... 15 

15. Lirellae simple or sparingly branched; disk weakly white pruinose; 

ascospores (4-)6-celled, 20-25 × 7-9 µm; norstictic acid; 

Highlands and Volusia counties.......................................................................................... Phaeographis sp. 23787B 

15. Lirellae much branched, angular, crowded, surrounded by poorly 

developed whitish area; disks bluish white pruinose; ascospores 

(4-)6-celled, 22-24 × 6-7 µm; stictic acid; Highlands, Lake and 

Volusia counties ....................................................................................... Sarcographa medusulina (Nyl.) Müll. Arg. 

14. Margin striate; lirellae long, flexuose, branched; disk becoming 

exposed, brown; ascospores 6-celled, 25-27 × 9-10 µm; 

Florida, without locality, Wilson......................................................................................................Phaeographis sp. 1259 

13. Exciple carbonized, at least at apex ..................................................................................................................................... 16 

15. Ascomata not erumpent ................................................................................................................................................... 17 

17. Hymenium inspersed with orange pigment, KOH+ dull purple; 

lirellae black, short, little branched; exciple carbonized at apex; 

ascospores 8-celled, 30-40 × 8-11 µm; norstictic acid; 

Monroe County ..................................................................................................................... P. multicolor R. C. Harris 

17. Hymenium colorless, KOH-; exciple entire; ascospores 6-8-celled, 

26-42 × 8-11 µm; norstictic acid; not yet found in Florida..............................................[P. dendritica (Ach.) Müll. Arg.] 

16. Ascomata erumpent with distinct erect bark flaps; exciple entire; 

hymenium inspersed; thallus often with a reddish tinge; ascospores 

6-7-celled, 27-32 × 8-9 µm; no substances; northern Florida ............................................ P. erumpens (Nyl.) Müll. Arg. 

NOTES 

14

Sarcographia labyrinthica (Ach.) Müll. Arg. 

Restored to the North American mycota. FLORIDA. Dade County: Everglades National Park, Mahogany Hammock, on large vine, 

2 Jan 1976, Buck B798 (NY). 

THELOTREMATACEAE (Nyl.) Stizenb. 

KEY TO GENERA 

1. Growing on bark ...................................................................................................................................................................................... 2 

2. Ascoma an apothecium........................................................................................................................................................................ 3 

3. Upper hymenium and ascus sheath I-; paraphyses mostly simple ................................................................................................. 4 

4. Periphysoids absent; columella present or absent....................................................................................................................... 5 

5. Exciple colorless to pale reddish brown; columellar structures, 

if present, noncarbonized..........................................................................................................................................Myriotrema 

5. Exciple carbonized; carbonized columella often present .........................................................................................Ocellularia 

4. Exciple with periphysoids on the inner face; columella always lacking.......................................................................Thelotrema 

3. Upper hymenium and ascus sheath I+ blue green; paraphyses somewhat 

anastomosed below, strongly so above the asci; apothecia immersed; hymenium 

separated from bark by thick layer of colorless crystals; periphysoids absent; 

ascospores colorless, vermiform, 30-40-celled, I-, ca. 85-120 × 7-9 µm; 

Seminole County ............................................................................................."Ocellularia" stictidea (Nyl.) V_zda ex Gilenstam 

2. Ascoma a mazaedium........................................................................................................................................................Nadvornikia 

1. Growing on soil or over Cladonia; spores brown, muriform ..............................................................................................Diploschistes 

DIPLOSCHISTES Norman 

1. Growing on Cladonia, at least initially; ascospores 4/ascus, 18-32 × 6-15 µm; 

lecanoric and diploschistesic acids; Polk County .................................................................................. D. muscorum (Scop.) R. Sant. 

1. Growing on soil, forming a continuous crust; ascospores 6-8/ascus, 20-28 × 8-14 µm; 

gyrophoric acid plus small amount of lecanoric acid; Gadsden County ............................................................ D. hypoleucus Zahlbr. 

MYRIOTREMA Fée 

1. Thallus not sorediate ............................................................................................................................................................................... 2 

2. Ascospores brown, submuriform to densely muriform ........................................................................................................................ 3 

3. Spores 8/ascus, submuriform .......................................................................................................................................................... 4 

4. Thallus PD+ orange or red............................................................................................................................................................ 5 

5. Thallus PD+ orange (stictic acid agg.)...................................................................................................................................... 6 

6. Ascomata rounded, not crowded .......................................................................................................................................... 7 

7. Norstictic acid lacking; ascomata round, slightly raised; disk not pulling 

away from thallus; Bay, Jackson, Liberty, Sarasota and Wakulla 

counties ........................................................................................................................... M. subcompunctum (Nyl.) Hale 

7. Norstictic acid in addition to stictic acid; ascomata smaller, more 

irregular; disk pulling away from thallus, Dade, Lake, Monroe and 

Seminole counties ................................................................................................................ M. compunctum (Ach.) Hale 

6. Ascomata angular, crowded, often in whitish, decorticate areas; 

15

ascospores 1-2 × 4-celled, 13-16 × 7-10 µm; throughout Florida .............................................M. glaucescens (Nyl.) Hale 

5. Thallus PD+ red (protocetraric acid); ascomata large, emergent with 

a simple to divided columella; ascospores 1-3 × 5-6-celled, 16-24 × 12-14 µm; 

Collier, Dade, Lee and Lake counties............................................................................................... M. bahianum (Ach.) Hale 

4. Thallus PD- (no substances), with large polygonal crystals, usually with 

pockets of bright red pigment; ascospores 2 × 4-celled, 14-18 × 10-14 µm; 

throughout Florida ....................................................................................................................................M. wightii (Taylor) Hale 

3. Spores 1-2/ascus, densely muriform, 70-130 × 20-30 µm; thallus PD+ orange 

(stictic acid agg.); common, south to Highalnds County ...............................................................M. reclusum (Krempelh.) Hale 

2. Ascospores colorless; apothecia smaller ........................................................................................................................................... 8 

8. Spores submuriform........................................................................................................................................................................ 9 

9. Apothecia small, 0.3-0.4 mm across, immersed; thallus light gray with 

numerous crystals; psoromic acid; ascospores 1-3 × 3-5-celled, 

9-20 × 4-6 µm; south to Hillsborough County .....................................................................................M. rugiferum (Harm.) Hale 

9. Apothecia larger, to 1.0 man, emergent, with broad rounded pore; 

thallus pale tan with numerous crystals; stictic acid agg. or acid deficient; 

ascospores 1-2 × 6-7-celled, 18-25 × 6-8 µm; throughout Florida ................................................... M. peninsulae R. C. Harris 

8. Spores transversely septate.......................................................................................................................................................... 10 

10. Cortex UV+ yellow, medulla PD- (lichexanthone, hypoprotocetraric acid); 

ascospores 4-celled, 10-12 × 5-6 µm; Dade and Monroe counties ................................................... M. glauculum (Nyl.) Hale 

10. Cortex UV-, medulla PD+ yellow (psoromic acid); ascospores 3-4-celled, 

9-15 × 4-7 µm; Alachua County....................................................................................................... M. terebratulum (Nyl.) Hale 

1. Thallus sorediate; protocetraric acid; ascospores colorless, 6-10-celled, 

42-46 × 9-11 µm; throughout Florida ................................................................................................................ M. erodens R. C. Harris 

NADVORNIKIA Tibell 

1. Thallus sorediate; throughout Florida ..............................................................................................................N. sorediata R. C. Harris 

1. Thallus not sorediate; Highlands County..................................................................................................N. hawaiiensis (Tuck.) Tibell 

OCELLULARIA G. Meyer 

1. Ascospores colorless .............................................................................................................................................................................. 2 

2. Ascospores transversely septate ......................................................................................................................................................... 3 

3. Ascoma smaller; medulla C-; ascospores under 50 µm long........................................................................................................... 4 

4. Thallus PD+ yellow or PD- ............................................................................................................................................................ 5 

5. Thallus PD+ yellow (psoromic acid).......................................................................................................................................... 6 

6. Columella present .................................................................................................................................................................. 7 

7. Ascospores 6-8-celled, 17-24 × 6-7 µm; apothecial margin 

not erect; pore usually small; columella undivided; common, 

throughout Florida ................................................................................................................ O. granulosa (Tuck.) Zahlbr. 

7. Ascospores 8(-9)-celled, 23-33 × 5-6 µm; apothecial margin 

erect; disk ± broad; columella becoming reticulate; rare, 

Collier County .............................................................................................................O. emersa (Krempelh.) Müll. Arg. 

6. Columella absent; ascospores 6-8-celled, 25-32 × 9-11 µm; 

Monroe County (Keys)..................................................................................................................Ocellularia sp. Small 9122 

5. Thallus PD- (no substances); apothecia emergent; rim often eroding 

16

to expose black rim around pore; columella present, black ..................................................................................................... 8 

8. Ascospores 4-6-celled, 13-20 × 5-8 µm; thallus dull, ecorticate; 

Hillsborough County ............................................................................................................ O. concolor Meyen & Flotow 

8. Ascospores 4-celled, 13-17 × 6-7 µm; thallus ± glossy, thinly 

corticate; Seminole County ...................................................................................................Ocellularia sp. Schallert 1664 

4. Thallus PD+ red (cinchonarum unknowns); apothecia emergent with round 

pore; columella present; ascospores 9-11-celled, 27-37 × 7.5-8.5 µm; 

Duval County ...................................................................................................................................... O. cavata (Ach.) Müll. Arg. 

3. Ascoma large, emergent, hemispherical; medulla pink, C+ pink (gyrophoric 

acid); ascospores very large, 19-23-celled, 100-130 × 20-35 µm; south to 

Lake County...........................................................................................................................................................O. americana Hale 

2. Ascospores densely muriform, 1-2/ascus, over 100 µm long ............................................................................................................. 9 

9. Thallus PD- (no substances), dull, usually roughened with small warts; 

medulla not conspicuous; south to Seminole County........................................................................O. sanfordiana (Zahlbr.) Hale 

9. Thallus PD+ orange (praestans unknown), shiny, very irregular; medulla 

conspicuous; Duval, Gulf and Nassau counties ..............................................................................O. praestans (Müll. Arg.) Hale 

1. Ascospores brown................................................................................................................................................................................. 10 

10. Ascospores submuriform ................................................................................................................................................................ 11 

11. Thallus PD- (no substances); ascospores 2 × 4-celled, 16-18 × 8-11 µm, 

with reticulate ornamentation; columella absent; Columbia and 

Wakulla counties ..................................................................................................................................O. retispora R. C. Harris 

11. Thallus PD+ yeiiow psoromic acid); ascospores 2 × 4-celled, 

13-16 × 8-10 µm, with weak striate ornamentation; known only 

from Louisiana ................................................................................................................................................[O. fissa (Nyl.) Hale] 

10. Ascospores transversely 4-celled ................................................................................................................................................... 12 

12. Apothecia immersed with small pore; columella undivided, not carbonized; 

pycnidia conspicuous, orange; hypoprotocetraric acid; ascospores 

15-18 × 8-10 µm, weakly granular or rugose warted; Columbia, 

Franklin, Leon, Liberty and Wakulla counties........................................................................O. leiostoma (Tuck.) R. C. Harris 

12. Apothecia immersed to emergent with broadly exposed disk; columella 

divided, carbonized; pycnidia not conspicuous; psoromic acid; 

ascospores 9-14 × 4-8 µm, smooth; Hillsborough County ......................................................... O. auberiana (Mont.) Hale 

NOTES 

Ocellularia auberiana (Mont.) Hale 

I excluded this species from North America in "Some Florida Lichens" as I could find no supporting specimen. I have since 

collected it. FLORIDA. Hillsborough County: Hillsborough River State Park, 28°09'N, 82°14'W, on oak, 3,4 Dec 1992, Harris 29684, 29722 

(NY). 

Ocellularia concolor Meyen & Flotow 

New to North America. FLORIDA. Hillsborough County: Hillsborough River State Park, 28°09'N, 82°14'W, on Carya and 

Liquidamabar, 3 Dec 1992, Harris 29680, 29686 (NY). 

Ocellularia emersa (Krempelh.) Müll. Arg. 

New to North America. FLORIDA. Collier County: Fakahatchee Strand State Preserve, Royal Palm Hammock, 26°01'N, 81°25'W, 

Buck 22763 (NY). 

THELOTREMA Ach. 

1. Thallus isidiate......................................................................................................................................................................................... 2 

2. Thallus PD+ yellow (psoromic acid), thinner ...................................................................................................................................... 3 

17

3. Ascospores brown, 2-celled, 8-11 × 5-6 µm, reticulately ornamented; 

isidia thick, long, often recumbent, "branching" by regeneration; 

apothecia emergent, doughnut-like; Liberty County.................................................................................. T. eximium R. C. Harris 

3. Ascospores colorless, submuriform, 4 × 4-celled, 15-21 × 9-12 µm; 

isidia sparse, initially globose; apothecia emergent, resembling Porina nucula Ach.; 

unknown substance in addition to psoromic acid; Leon, Okaloosa 

and Santa Rosa counties .............................................................................................................. T. halei (Tuck. & Mont.) Zahlbr. 

2. Thallus PD- (no substances), thick, filled with crystals, often separating 

from the substrate; isidia sparse, coarse; ascospores brown, submuriform, 

16-28 × 7-12 µm; south to Sarasota County...........................................................................................................T. santense Tuck. 

1. Thallus not isidiate................................................................................................................................................................................... 4 

4. Ascospores colorless .......................................................................................................................................................................... 5 

5. Ascospores transversely septate .................................................................................................................................................... 6 

6. Thallus PD- (no substances); ascomata small, immersed ......................................................................................................... 7 

7. Ascospores purple with iodine ................................................................................................................................................ 8 

8. Apothecia 0.4-0.6 mm across; thallus poorly developed, not corticate; 

medulla not obvious; ascospores 9-15-celled, 25-40 × 6-8 µm; 

northern Florida ...........................................................................................................................................T. subtile Tuck. 

8. Apothecia 0.2-0.35 m across; thallus usually well developed, 

weakly corticate; medulla thick; ascospores 8-13-celled, 25-45 × 6-8 µm; 

south to Polk County ............................................................................................................................. T. lathraeum Tuck. 

7. Ascospores colorless with iodine, 11-14-celled, 24-38 × 6-7 µm; 

apothecial disk mostly flush with thallus; Lake County ..........................................................T. alborosellum (Nyl.) Tuck .? 

6. Thallus PD+ yellow or orange; apothecia larger......................................................................................................................... 9 

9. Thallus PD+ orange (stictic acid agg.).................................................................................................................................. 10 

10. Apothecia with thick, recurved and split margin; ascospores 7-10-celled, 

30 × 6-8 µm; Florida Panhandle ......................................................................................... T. dilatatum (Müll. Arg.) Hale 

10. Apothecia emergent with broad round pore; exciple free from margin 

forming a conspicuous inner ring; ascospores to 30-celled, 

120-140 × 15-20 µm; exact locality unknown, known only from 

type of Ocellularia floridensis ......................................................................................T. porinoides Mont. & v.d. Bosch 

9. Thallus PD+ yellow (psoromic acid); apothecia emergent, round with broad 

round pore and smooth margin; exciple separating from margin; ascospores 

4-6-celled, 13-18 × 6-7 µm; Franklin and Liberty counties .............................................................T. floridense R. C. Harris 

5. Spores submuriform to muriform; thallus PD- (no substances) ................................................................................................... 11 

11. Ascospores large, over 50 µm long, 1-4/ascus ...................................................................................................................... 12 

12. Ascospores thick walled, 2-4/ascus, 65-90 × 15-20 µm ................................................................................................... 13 

13. Apothecia immersed when young, becoming urceolate with low, 

thick margin; exciple coarse, pinkish tan, often filling pore and 

covering the disk; Panhandle ...............................................................................................................T. adjectum Nyl. 

13. Apothecia with large, open pore; margin regular, thinner; 

exciple fragile, whitish, free from margin; Duval County ....................................................... T. lepadinum (Ach.) Ach. 

12. Ascospores thin walled, IKI-, 1-2/ascus, 50-80 × 15-20 µm; apothecia 

white pruinose with erect, jagged margin; south to Hillsborough 

County ......................................................................................................................................T. leprocarpum (Nyl.) Tuck. 

11. Ascospores 25-35(-45) × 12-15(-18) µm, initially 8/ascus but several may abort; 

apothecia small, immersed, radially striate and lacerate; Citrus, Columbia, 

Jackson, Leon and Union counties ................................................................................... T. defectum Hale ex R. C. Harris 

18

4. Ascospores light to dark brown......................................................................................................................................................... 14 

14. Ascospores 5-10 transversely septate......................................................................................................................................... 15 

15. Ascospores 6-celled, 17-20 × 6-7 µm; stictic acid agg.; 

apothecia with rim recurved and splitting; disk white 

pruinose; Collier County..................................................................................................................T. platycarpoides Tuck. 

15. Ascospores 7-11-celled, 33-60 × 6-10 µm; no substances; 

apothecia emergent with broad, round pores; exciple pulling 

away from margin; Santa Rosa and Seminole counties....................................................................... T. lacteum Krempelh. 

14. Ascospores densely muriform, 1/ascus, 105-140 × 24-30 µm; 

no substances; apothecia emergent with broad, round pores; 

exciple pulling away from margin; Dixie, Liberty, Marion 

and Santa Rosa counties ............................................................................................................. T. monospermum R. C. Harris 

NOTES 

Thelotrema platycarpoides Tuck. 

Thelotrema platycarpoides was originally described from Cuba. Hale (1981) was in error describing the ascospores as "3-4loculate". 

Tuckerman described them as 6-celled and an examination of an isolectotype (BUF) corroborates this. (The very similar T. 

platycarpum Tuck. does have 4-celled ascospores and also differs in larger apothecia.) 

New to North America. FLORIDA. Collier County, Fakahatchee Strand State Preserve, Royal Palm Hammock, 26°01'N, 81°25'W, 

6 Dec 1992, Harris 29940 (NY). 

"CALICIALES" 

"Calciales" are used here in an informal, highly artificial, sense to include not only the Caliciaceae and 

Mycocaliciaceae but mazaedioid genera belonging to other families or of uncertain position. Mycocaliciaceae 

and Caliciaceae are clearly not at all closely related. The familial disposition of Heterocyphelium Vainio and 

Tylophoron Nyl. is uncertain. The first is monotypic and treated in the generic key. The second is left in the 

Caliciaceae until its relationships are better understood. 

KEY TO GENERA 

1. Apothecia stalked or if almost sessile, then very small ............................................................................ 2 

2. Asci disintegrating, forming a mazaedium ........................................................................................... 3 

3. Ascospores simple ............................................................................................................................ 4 

4. Ascospores small (5 µm or less), more or less globose, pale, 

with weak ornamentation formed by cracking of exospore; 

lichenized, on bark or leaf bases of Sabal................................................................ Chaenotheca 

4. Ascospores larger (to 17µm), lemon-shaped, dark, 

with strong reticulate ribbing; apothecia top-shaped with short, 

thick stalk; parasymbiont on Pertusaria; so far only Gulf Coast 

from Dixie to Collier County.........................................................Sphinctrina tubiformis Massal. 

3. Ascopores 2-celled, brown .................................................................................................. Calicium 

2. Asci persistent, not forming mazaedium (Mycocaliciaceae) ................................................................5 

5. Growing on old wood or bark ............................................................................................................ 6 

6. Stalk dark, usually of more or less parallel hyphae; 

19

ascospores simple with pointed ends; ascus tip not 

pierced by narrow canal ...........................................................................................Mycocalicium 

6. Stalk pale, usually of irregularly arranged hyphae 

(very short in C. pilosa); ascospores simple or 2-celled, 

usually with rounded ends but sometimes as in Mycocalicium; 

ascus tip pierced by a narrow canal .................................................................. Chaenothecopsis 

5. Growing on the polypore, Trichaptum biforme (Fr.) Ryv.; 

ascospores 2-celled, pale, 10-14 × 3-5 µm; 

northern Florida .............................................................Phaeocalicium polyporaeum (Nyl.) Tibell 

1. Apothecia immersed or sessile, large ...................................................................................................... 7 

7. Ascospores 3-4-celled............................................................................................................................ 8 

8. Thallus UV+ yellow (lichexanthone); ascoma formed of a lower 

perithecioid chamber and upper cup-shaped chamber holding loose 

ascopores; ascospores 4-celled........................................................................................................... 

............................................... Pyrgillus javanicus (Mont. & v.d. Bosch) Nyl. (See Pyrenulaceae) 

8. Thallus UV-; ascoma not 2-chambered; ascospores usually 3-celled, 

with middle cell largest; no substances; Seminole County.................................................................. 

................................................................................. Heterocyphelium leucampyx (Tuck.) Vainio 

7. Ascospores 2-celled............................................................................................................................... 9 

9. Apothecial margin and thallus C-, KOH+ yellow or red, PD+ orange 

(norstictic or stictic acid)................................................................................................................... 10 

10. Norstictic acid agg. present; ascospores 13-17 × 8-9 µm (Tibell, 

1976); peninsular Florida south to Highlands County .................................................................... 

....................................................................................... Thelomma carolinianum (Tuck.) Tibell 

10. Stictic acid agg. present; ascospores 7-9 × 5-6 µm (Tibell,1987) .................................................. 

...........................................................................................Nadvornikia (See Thelotremataceae) 

9. Apothecial margin C+ red (lecanoric acid), KOH-, PD-; 

ascospores with very dark middle region ......................................................................... Tylophoron 

CALICIACEAE Chev. 

CALICIUM Pers. 

1. Lowerside of capitulum white pruinose or not pruinose ............................................................................ 2 

2. Apothecia relatively large, over 0.5 mm tall............................................................................................ 3 

3. Thallus KC+ orange (xanthones), areolate and yellowish or more 

often in Florida material not evident; stalk IKI-; ascospores 

10.5-14 × 4.5-7.5 µm (Tibell, 1987); common, peninsular Florida ................... C. hyperelloides Nyl. 

3. Thallus KC- (no substances), granular; stalk IKI+ blue; 

ascospores 9-14 × 4-7.5 µm (Tibe11, 1987); Bradford, 

Leon and Marion counties .................................................................................... C. lenticulare Ach. 

20

2. Apothecia minute, ca. 0.2 mm tall; capitulum with faint yellow pruina?; 

ascospores with pointed ends, strongly spirally ornamented, 10-13 × 4.5-6 µm; 

on twigs; Citrus and Hamilton counties ........................................................ C. adspersum Pers. s. lat. 

1. Lowerside of capitulum brown pruinose; ascospores with spiral 

ornamentation ........................................................................................................................................... 4 

4. Spores larger, mostly over 11 × 6 µm ................................................................................................... 5 

5. Thallus UV+ yellow (lichexanthone), KC+ weak orange, with same 

unknown as Florida C. salicinum, not shiny; ascospores 

9-14 × 5.5-7 µm (Tibell, 1987); peninsular Florida ................................ C. chlorosporum F. Wilson 

5. Thallus UV-, KC+ red-orange (xanthones); ascospores 12.5-15.5 

× 7-8 µm; Gulf Coast from Taylor to Collier County ...................................... C. leucochlorum Tuck. 

4. Spores small, 7-11 × 3.5-5 µm (Tibell, 1987); thallus with unknown 

substance; Bay and Lake counties.................................................................C. salicinum Pers. s. lat. 

NOTES 

Calicium adspersum Pers. s. lat. 

Tibell (1987) considered this a variable species. Whether it is commodious enough to encompass the 

Florida material is open to question. The two collections, although very minute in stature, are placed here 

because of the faint yellow pruina and the rather narrow, mostly pointed ascospores. Its ecological 

preferences may prove to be significant, one collection is from small oak twigs, the other from a stem of the 

shrub Ceratiola. 

Calicium salicinum Pers. s. lat. 

The chemistry of this group is highly problematical. Tibell (1987) reported norstictic acid. In northern 

North American, European, Andean South American and Australian material I find a spot that runs much like 

norstictic acid but seems to differ in color and shape. I get no red needles with KOH on thallus sections. I 

regard norstictic acid as possible but unlikely. The Florida specimens and another from Puerto Rico contain a 

different substance in high concentration. It has some of the characteristics of and is possibly a dibenzofuran. 

This same substance is also found in C. chlorosporum accompanied by lichexanthone. Since the lowland 

neotropical population of C. salicinum is chemically distinct its taxonomic status should perhaps be reviewed. 

21

CHAENOTHECA Th. Fr. 

1. Base of capitulum brown to whitish, not pruinose; photobiont 

Chlorococcalean........................................................................................................................................ 2 

2. Thallus PD+ yellow (unknown); apothecia relatively tall and slender; 

ascospores 2.7-4.3 µm in diameter, more or less smooth (Tibell,1980); 

common, penisular Florida.....................................................................C. brunneola (Ach.) Müll. Arg. 

2. Thallus PD+ red-orange (virensic acid); apothecia relatively short and 

stout; ascospores 4-5 µm in diameter, rather coarsely ornamented; 

Hillsborough, Levy, Putnam and Sarasota counties.........................................C. floridana R. C. Harris 

1. Base of capitulum yellow pruinose; photobiont Trentepohlia; 

ascospores 3-4 µm in diam.; Bay County............................................. C. hispidula (Ach.) Zahlbr. s. lat. 

NOTES 

Chaenotheca brunneola (Ach.) Müll. Arg. s. lat. 

I cannot make out the chemistry reported by Tibell (1980,1987) for the brunneola-hygrophilastemonea 

complex. Most Florida collections of C. brunneola contain an unknown identical to that found in 

22

three collections of C. stemonea tested (New York, Minnesota, Sweden). It runs about the same as 

baeomyesic acid but the spot characteristics are different. Three collections of C. brunneola (Florida, Maine & 

Sweden) contain no detectable substances, four contain baeomycesic and squamatic acids (Florida, New 

York, North Carolina & Sweden). Tibell reported baeomycesic acid in1980 but rejected it in 1987. It seems 

probable that his unknowns B & C are the same as the single unknown spot that I get with my TLC system. 

However, it is clear that more work needs to be done to confirm this. The baeomycesic acid strain is known 

from a single collection at Gold Head Branch State Park (Clay County) where the unknown substance strain 

also occurs. Most specimens checked had asci in chains. Collections from Duval and Hernando counties had 

asci arising singly. 

Chaenotheca floridana R. C. Harris, sp. nov 

Ab aliis acidum virensicum continentia, ascosporis grosse ornamentis, 4-5 µm diametro et apothecis 

pallidis brevibus et crassiusculis differt. 

Type. FLORIDA. Putnam County: Ocala National Forest, old Johnson Field Campground along Oklawaha River 

just W of U.S. Hwy. 19, swamp forest, 7 Dec 1988, Buck 16798 (NY, holotype; B, UPS, isotypes). 

Thallus whitish, granular leprose, PD+ red-orange, virensic acid, unknown (tr.), protocetraric acid (tr.). 

Phycobiont trebouxioid. Apothecia relatively short and stout, to 1 mm tall. Stalk to 0.3 mm across at base 

tapering to ca. 0.1 mm, light orange brown (more or less concolorous with spore mass), appearing white 

pruinose due to wefts of loose hyphae or occasionally covered with thallus particles. Capitulum obconical to 

globose, to 0.5 mm in diameter. Asci not in chains. Ascospores globose or appearing slightly angular due to 

coarse plate-like ornamentation, 4-5 µm in diameter, yellow brown. 

I originally included this species in C. brunneola s. lat. until I studied its chemistry. A subsequent reevaluation 

showed that it also differs in the pale, shorter apothecia and slightly larger, coarsely ornamented 

ascospores. 

Additional specimens. FLORIDA. Hillsborough County: Hillsborough River State Park, Rapids Trail 

along river, 28°09'N, 82°14W, on trunk of Sabal, 3 Dec 1992, Buck 22641; Levy County: Cedar Key Scrub 

State Preserve, along Co. Rd. 347 ca. 1.5 mi N of Fla. Hwy. 24, on Sabal leaf bases, 30 Nov 1992, Buck 

22425, 22434, 22436; Sarasota County: Myakka River State Park, along Fla. Hwy. 72 ca. 0.7 mi SE of park 

entrance, on trunk of Sabal, 5 Dec 1992, Buck 22717, 22731 (all NY). 

Chaenotheca hispidula (Ach.) Zahlbr. s. lat. 

The single specimen varies from the North American norm in having the asci in chains and in smaller 

ascospores reaching only 4 µm which is below the minimum given by Tibell (1980). It is possible that the 

deviation is clinal as the locality is considerably south of the range given by Tibell. 

THELOMMA Massal. 

Thelomma carolinianum (Tuck.) Tibell is endemic to the Coastal Plain. The other species of the 

genus occur in western North America and Europe. See key to genera. 

TYLOPHORON Nyl. ex Stizenb. 

1. Thallus and ascomatal margin C+ red (lecanoric acid); ascomata 

0.6-1.2 mm diam. (Tibell, 1982); ascospores 12-15 × 6.5-8 µm; 

sporodochia white; conidiospores 1-2(-4)-celled, colorless, 

rough-walled; throughout Florida.................................................................................. T. protrudens Nyl. 

1. Thallus C-; ascomatal margin C+ red (lecanoric acid, 2'-O-methylperlatolic 

acid); ascomata 0.4-0.8 mm diam. (Tibell, 1982); ascospores 10-14 × 5-6 µm; sporodochia 

23

own (not known in Florida); conidiospores simple, 

brown; Collier County ...................................................................................................T. moderatum Nyl. 

NOTES 

Tibell (1991) described the sporodochial anamorph of T. moderatum but did not mention one for T. 

protrudens. However, Florida material always has conspicuous sporodochia. In fact, seven of the ten Florida 

collections consist of sporodochia only. The same anamorph was also found on Lindig 1 from Colombia. The 

number of cells in a conidium is apparently determined by how the hyphae break to form the chlamydosporelike 

conidia. In T. moderatum the sporodochia apparently do not occur on the same thallus as the ascomata. 

However, to my eye two collections of T. protrudens have both on the same thallus. There is only a single 

chemotype in Florida as neither lichexanthone nor the red ascomatal pigment was detected. Tylophoron 

protrudens is probably common but not often collected by lichenologists as it somewhat resembles members 

of the Aphyllophorales. It might also be confused with Cryptothecia striata Thor (Chiodecton montagnei auct.) 

that also seems sterile and is C+. A section of a sporodochium with conidiospores readily identifies the 

Tylophoron. 

SPHINCTRINACEAE Choisy 

SPHINCTRINA Fr. 

Only S. tubiformis is known from Florida. See key to genera. 

MYCOCALICIACEAE A. Schmidt 

KEY TO CHAENOTHECOPSIS AND MYCOCALICIUM 

1. Apothecia and thallus lacking lichen substances; ascospores simple, 

elliptical to fusiform.................................................................................................................................... 2 

2. Apothecia distinctly stalked, not hairy.................................................................................................... 3 

3. Exciple and stalk brown or stalk almost colorless; exciple not 

incurved and rugose......................................................................................................................... 4 

4. Stalk brown to dark brown; stalk hyphae more or less parallel .................................................... 5 

5. Cells of lower part of exciple not enlarged, not paraplectenchymatous .................................. 6 

6. Apothecia relatively broad and flat, ± conical, often 

2-3/stalk; stalk relatively short and stout; ascospores 

6-8 × 2.5-3.5 µm; on old wood; common, south to 

Palm Beach County ............................................... Mycocalicium subtile (Pers.) Szatala 

6. Apothecia narrowly obconical or obpyriform, often 2-3/stalk; 

stalk long, slender and flexuous; ascospores pale, 6-7 × 2.5-3 µm; 

rare, Washington County ................................Chaenothecopsis savonica (Räs.) Tibell? 

5. Cells of lower part of exciple enlarged and paraplectenchymatous; 

ascospores 6-8 × 2.5-3.5 µm; on old wood; Citrus and Sumter counties................................. 

24

.................................................................................... Mycocalicium albonigrum (Nyl.) Fink 

4. Stalk almost colorless to pale olive or pale brown; hyphae irregularly arranged.......................... 7 

7. Ascospores arranged end to end ............................................................................................. 8 

8. On Sabal palmetto; ascospores 7-9 × 3-4 um; common, 

south to Collier County ............................... Chaenothecopsis rappii (Nadv.) R. C. Harris 

8. On pine bark; ascospores 6-7 × 2.5-4 µm; rare, 

Franklin County ........................................................................Chaenothecopsis sp. 1393 

7. Ascospores obliquely arranged, 7-8 × 3.5-4 µm; on Quercus; 

rare, Citrus County .................................................................... Chaenothecopsis nana Tibell 

3. Exciple and stalk reddish, redder in KOH; exciple incurved, longitudinally 

rugose; ascospores 8-11 × 4.5-5.5 µm; on old wood; Santee Canal, 

South Carolina, not yet found in Florida ...................... [Mycocalicium ravenelii (Tuck. ex Nyl.) Fink] 

2. Apothecia sessile or with a very short stalk, hairy due to projecting 

excipular hyphae; ascospores simple; growing on thallus of sterile 

Tylophoron protrudens; Wakulla County................................Chaenothecopsis pilosa Tibell & Kalb 

1. Apothecia containing norstictic acid; ascospores 2-celled, ± cylindrical 

with rounded ends, 7-11 × 3-3.5 µm; on old wood; Bradford County and 

Okefenokee Swamp, Georgia............................................... Chaenothecopsis norstictica R. C. Harris 

NOTES 

CHAENOTHECOPSIS Vainio 

Chaenothecopsis norstictica R. C. Harris, sp. nov. 

Unica Mycocalicacearum acidum norsticticum continens. 

Type. GEORGIA. Charlton County: Okefenokee National Wildlife Refuge, Suwanee Canal Recreation Area, 

along "Swamp Trail" on Chesser Island, swamp forest, 12 Dec 1987, Buck 15462 (NY, holotype). 

Thallus not evident, on lignum, no algae seem to be associated with stalk bases. Cladonia ravenelii 

Tuck. is also on the wood but the Chaenothecopsis does not seem to be directly associated with it. Apothecia 

dark brown, slightly shiny, ca. 0.9 mm tall. Stalk dark brown; hyphae seem more or less parallel but hard to 

tell. Capitulum obconical, ca. 0.3-0.4 mm in diameter. Exciple and upper stalk bleeding yellow in KOH with 

subsequent formation of typical crystals of norstictic acid (confirmed by TLC), norstictic acid apparently 

present in substantial concentration. Asci with tip pierced by narrow canal. Ascospores pale brown, more or 

less cylindrical with rounded ends, 2-celled, 7-11 × 3-3.5 µm; septum dark. 

Chaenothecopsis norstictica is apparently unique among the Mycocaliciaceae in producing a 

depsidone. The chemistry of the family otherwise seems confined to various unknown pigments or rarely the 

pulvinic acid group. 

Additional specimens. FLORIDA. Bradford County: Lake Butler Wildlife Management Area, Raiford 

Tract, along Co. Rd. 125, 6.1 mi ENE of U.S. 301 at Lawtey, 30°04'N, 82°09'W, pine scrub around Taxodium 

swamp, 3 Dec 1994, Buck 27333, 27343, 27348 (NY). 

25

Chaenothecopsis pilosa Tibell & Kalb, Nova Hedwigia 60: 211. 1995. 

New to North America. FLORIDA. Wakulla County: Apalachicola National Forest, along Forest Service 

Rd. 309 at Lost Creek just S of Leon Co. line, 5.6 mi W of Fla. Hwy. 267, swamp forest, on Tylophoron 

protrudens on Fraxinus, 4 Dec 1988, Harris 23372 (NY). 

Chaenothecopsis rappii (Nádv.) R. C. Harris, comb. nov. 

Mycocalicium rappii Nádv., Ann. Mycol. 40: 139. 1942. 

This species has simple pointed ascospores as in Mycocalicium but Tibell includes other species with this 

ascospore type in Chaenothecopsis. The stalk is pale with irregularly arranged hyphae and the ascus tip has 

a narrow canal which are considered by Tibell, 1987) as better indicators of generic placement if one wishes to 

recognize Chaenothecopsis as distinct from Mycocalicium. It would be simpler not to. 

MYCOCALICIUM Vainio 

Two species are known from Florida. See key above. Mycocalicium ravenelii is close to M. calicioides 

(Nádv.) Tibell. Both seem to me quite distinct from M. subtile, the generitype, and probably should be 

segregated into Sphinctrinella Nádv. (type, S. calicioides). 

PHAEOCALICIUM A. Schmidt 

Only the single species, P. polyporaeum (Nyl.) Tibell, occurs in Florida and is on the polypore, Tricaptum 

biforme (Fr.) Ryv. (Polyporus pargamenus Fr.). See key to genera. 

LEOTIALES 

BAEOMYCETACEAE Dumortier 

(incl. Icmadophilaceae Triebel) 

I do not think that recognition of Icmadophilaceae is warrented and does not seem to be in line with 

general practice in the leotioid ascomycetes. The generic split between Baeomyces Pers. and Dibaeis 

Clements is well supported. Dibaeis baeomyces (L. fil.) Rambold & Hertel ( syn. Baeomyces roseus Pers.) is 

the only species in Florida. It is obviously at the southern edge of its North American range. It was collected 

on a road bank in Escambia County and disintegrated sandstone at Rock Hill, Washington County. 

LECANORALES 

COLLEMATACEAE Zenker 

KEY TO GENERA 

1. Ascospores septate; thallus without reticulate maculae......................................................................................................................... 2 

2. Thallus with cortex...............................................................................................................................................................Leptogium 

2. Thallus without cortex.............................................................................................................................................................. Collema 

1. Ascospores simple; thallus with reticulate markings (maculae) due to absence of photobiont; 

apothecia large, with very thick wrinkled margin; thallus tomentose below; 

reported by Moore (1970) from Citrus County, not seen by me ......................................................... Physma byrsaeum (Ach.) Tuck. 

26

COLLEMA Wigg. 

[The non-lichenized loculoascomycete, Myriangium duriaei Montagne & Berkeley forms small cushions with apothecium-like ascomata 

and can in the field be confused with a tiny Collema. The brown, cellular tissue, lack of photobiont and globose asci with muriform 

ascospores readily distinguish it in the lab.] 

1. Lobes large and broad ............................................................................................................................................................................ 2 

2. Lobes longitudinally ridged and pustulate, isidiate or not.................................................................................................................... 3 

3. Isidia absent (C. pulcellum Ach. s. lat.)............................................................................................................................................. 4 

4. Ascospores narrower, under 5 µm wide ....................................................................................................................................... 5 

5. Apothecial disks strongly white pruinose; south to Polk County ................................................................................................. 

.........................................................................................................................C. pulcellum var. leucopeplum (Tuck.) Degel. 

5. Apothecial disks not pruinose; Duval County ......................................................................C. pulcellum Ach. var. pulcellum 

4. Ascospores broad, over 5 µm wide; apothecial disk occasionally pruinose; 

north to Flagler County.............................................................................C. pulcellum var. subnigrescens (Müll. Arg.) Degel. 

3. Isidia present, mainly on ridges and pustules; apothecial disk pruinose; proper exciple 

paraplectenchymatous; Citrus, Collier, Hernando and Levy counties (sterile), 

"S.W. Florida" (with apothecia) ..............................................................C. furfurceum (Arn.) Du Rietz var. luzone (Räs.) Degel. 

2. Lobes not ridged or pustulate, with ± globose isidia; a single scrappy 

collection from Madison County tentatively assigned here ........................................................................... C. subflaccidum Degel. 

1. Lobes narrow, often anastomosing......................................................................................................................................................... 6 

6. Ascospores rod-shaped with more than 2 cells or if broader, submuriform ...................................................................................... 7 

7. Apothecia subglobose, mostly immersed in the thallus; ascospores globose to ellipsoid, submuriform ..................................... 8 

8. Thallus foliose; proper exciple paraplectenchymatous; ascospores ellipsoid, 

2-3 × 4(-6)-celled, 20-40 × 11-15 µm (Degelius); Dade County.................................................................... C. pustulatum Ach. 

8. Thallus subfoliose; proper exciple prosoplectenchymatous; ascospores globose 

to subglobose, 2 × 3-4-celled, 8.5-13 µm in diameter or 10.5-17 × 8.5-10.5 µm; 

Jackson County and reported by Degelius (1974) from Caloosa River ......................................................C. callibotrys Tuck. 

7. Apothecia cup-shaped, sessile to stalked; ascospores narrow, ca. 2 µm wide, 

transversely septate with blunt ends; Leon and Seminole counties ................................................................ C. leptaleum Tuck. 

6. Ascospores 2-celled, 8.5-21 × 3-6.5 µm; thallus lobes small and crowded; 

apothecia usually numerous; rare, Citrus and Hillsborough counties......................................................C. conglomeratum Hoffm. 

LEPTOGIUM (Ach.) Gray 

Since the bulk of NY's collections of Leptogium are on loan and the new treatment, when it appears, is likely to encompass 

considerable change, I include Moore's key for completeness and convenience. 

1. Thallus larger, clearly foliose................................................................................................................................................................... 2 

2. Thallus isidiate...................................................................................................................................................................................... 3 

3. Thallus wrinkled................................................................................................................................................................................. 4 

4. Lobes distinct and not anastamosing, the margins not thickened; isidia not granular ................................................................ 5 

5. Thallus longitudinally wrinkled; isidia concentrated along the ridges; 

throughout Florida ......................................................................................................................L. isidiosellum (Riddle) Sierk 

5. Thallus irregularly wrinkled; isidia marginal or scattered on the surface ................................................................................. 6 

6. Isidia mostly marginal and associated with apothecia; on hardwood trees; 

throughout Florida ............................................................................................................. L. marginellum (Swartz) S. Gray 

27

6. Isidia scattered on the surface of the thallus; on trees and rocks; throughout 

the state .................................................................................................................... L. austroamericanum (Malme) Dodge 

4. Lobes irregularly anastamosing, the margins thickened; isidia becoming granular; 

scattered throughout Florida .........................................................................................................................L. milligranum Sierk 

3. Thallus not wrinkled; throughout Florida.................................................................................... L. cyanescens (Rabenh.) Körber 

2. Thallus not isidiate................................................................................................................................................................................ 7 

7. Thallus smooth, not wrinkled............................................................................................................................................................ 8 

8. Thallus not pitted .......................................................................................................................................................................... 9 

9. Thallus blue-gray, lobes 1-5 mm broad; scattered throughout Florida ......................................... L. azureum (Swartz) Mont. 

9. Thallus gray-brown, lobes 2-3 mm broad; Duval County?, reported by Sierk 

on the basis of a Calkins collection..........................................................................................................L. juniperinum Tuck. 

8. Thallus pitted; peninsular Florida ........................................................................................................... L. microstictum Vainio 

7. Thallus wrinkled.............................................................................................................................................................................. 10 

10. Spores acicular or fusiform ...................................................................................................................................................... 11 

11. Spores acicular, 50-85 µm long; thallus gray; Sarasota and Seminole counties.................................... L. adpressum Nyl. 

11. Spores fusiform, 25-40 µm long; thallus usually brownish; peninsular Florida 

north to Levy County...............................................................................................................L. fusisporum (Tuck.) Dodge 

10. Spores ellipsoid or subfusiform................................................................................................................................................ 12 

12. Lobes distinct, not anastamosing........................................................................................................................................ 13 

13. Thallus naked above, shiny, with large pustules; cited by Moore 

from four counties, not found by me nor have I seen her material, 

Florida records possibly equal the next species............................................................. L. corticola (Taylor) Tuck. 

13. Thallus sparsely hairy above; hairs short with cylindrical cells; thallus naked 

below, dull, with small pustules ................................................................................................ Leptogium sp. 31815 

12. Lobes irregularly anastomosing .......................................................................................................................................... 14 

14. Thallus heavily wrinkled longitudinally, lobes narrow and 

elongate or inflated and hollow ................................................................................................................................ 15 

15. Apothecia not at the ends of hollow lobes; cortical cells 

usually more than 5 µm wide.............................................................................................................................. 16 

16. Thallus thickened but not warty, generally appearing gray 

or gray-green ................................................................................................................................................. 17 

17. Thallus gray; thalloid exciple lobulate; scattered throughout 

the state ................................................................................................... L. phyllocarpum (Pers.) Mont. 

17. Thallus green or gray-green; thalloid exciple not lobulate; 

scattered throughout the peninsula................................................................L. chloromelum (Ach.) Nyl. 

16. Thallus warty, usually appearing bluish; north to Lake 

and Seminole counties..................................................................................................... L. floridanum Sierk 

15. Apothecia at the ends of elongate hollow lobes; cortical cell 

usually less than 5 µm wide; peninsular Florida from Levy and 

Marion counties south to Palm Beach County......................................................................L. stipitatum Vainio 

14. Thallus wrinkled but flattened, lobes rounded; Lake, Polk and 

Seminole counties ............................................................................................................................ L. sessile Vainio 

1. Thallus small, appearing almost crustose; reported from Seminole County by Sierk ....................L. tenuissimum (Dickson) Körber 

28

PILOCARPACEAE Zahlbr. 

There are more Florida species of this family in genera other than Byssoloma. I have chosen to treat 

only Byssoloma at this time since the most common North American species has not been previously 

recognized. Further, the generic limits in the family are not clear to me at this time and the other North 

American species seem to fall into groups which may merit generic status. The key owes much to Santesson 

(1952). 

BYSSOLOMA Trevisan 

1. Thallus UV-; hypothecium dark brown; mostly on leaves.......................................................................... 2 

2. Margin whitish, KOH-; disk brownish to black ....................................................................................... 3 

3. Disk yellow brown to dirty brown, often mottled; exciple pale; 

ascospores10-18 × 2.5-4 µm; on leaves and bark; common, 

south to Polk County .....................................................................B. leucoblepharum (Nyl.) Vainio 

3. Disk black; exciple often greenish tinted; ascospores 

10-17 × 3-5 µm; on leaves; Lake, Seminole, Taylor, 

and Wakulla counties ................................................................. B. subdiscordans (Nyl.) P. James 

2. Margin yellow brown, KOH+ yellow; disk brown to brownish black; 

ascospores 9-15 × 3-4 µm; on leaves; Highlands, 

Marion and Seminole counties ........................................................... B. tricholomum (Mont.) Zahlbr. 

1. Thallus UV+ orange (xanthones); hypothecium yellowish; byssoid margin 

pale; disk yellowish becoming yellow brown, often mottled; ascospores 

10-15 × 2.5-4 µm; on bark; common, south to Polk County........................................................................ 

........................................................................................................B. pubescens V_zda ex R. C. Harris 

NOTES 

Byssoloma pubescens V_zda ex R. C. Harris, sp. nov. 

Superficie similis B. leucoblephari sed thallo UV+ aurantiaco et hypothecio pallido. 

Type. FLORIDA. Seminole County: Sanford, on oak, May 1911, Rapp (NY, holotype). 

Thallus corticolous, gray green, dull, smooth, continuous, UV+ orange (xanthones). Apothecia 

sessile, orbicular, flattened, tightly adnate, 0.2-0.7 mm diam.; byssoid margin moderately to slightly developed, 

usually not spreading, whitish, KOH-; disk pale whitish yellow to dirty yellowish, often mottled with darker 

yellow brown, sometimes entirely darkening to dirty brown. Epithecium ± colorless in section. Hymenium 

nearly colorless or yellowish. Hypothecium nearly colorless or yellowish, rarely light brownish. Ascospores 4celled, 

10-15 × 2.5-4 µm. Microconidia lageniform, 3-4 × 1.5 µm. 

This very commonly collected species sometimes grows mixed with the rather similar B. 

leucoblepharum for which it has been mistaken until recently. Byssoloma pubescens is easily distinguished 

under the ultraviolet lamp by the intense orange fluorescence of the thallus. Also under the dissecting 

microscope the absence of a dark hypothecium is readily evident. The taxon was first recognized by V_zda 30 

years ago when he annotated the type specimen with this name. I believe all other species of the genus have 

a dark hypothecium. This raises the question as to generic limits, which in the Pilocarpaceae are, I think, 

rather fuzzy. Byssoloma, apparently can have an essentially non-byssoid margin but I am unaware of the 

29

everse, i.e., anything other than Byssoloma with a byssoid margin. Thus far B. pubescens is always on bark, 

ranging from tiny twigs to large tree trunks, mainly in mesic or swamp forest habitats. In North America it is a 

species of the southern Coastal Plain and is also known from Cuba and Puerto Rico. 

Additional specimens. FLORIDA. Bay County: N of Co. Rd. 388 at Econfina Creek, 7.9 mi W of U.S. 

231 at Youngstown, 30°23'N, 85°34'W, 1 Dec 1994, Buck 27223, N of Co. Rd. 388, 4.5 mi W of Fla. Hwy. 77, 

30°20'N, 85°44'W, 1 Dec 1994, Buck 27275; Calhoun County: W of St. Rd. 71, 6.6 mi N of Gulf Co. line, 

30°16'N, 85°10'W, 11 Dec 1993, Buck 24769; Dixie County: Steinhatchee Wildlife Management Area, 

Pumpkin Swamp, along Co. Rd. 358 ca. 7 mi NW of Cross City, 29°41'N, 83°13'W, on Sabal leaf base, 4 Dec 

1993, Harris 31584, 31591, along Sauls Rd. ca. 0.4 mi N of Co. Rd. 351, ca. 6 mi NE of Cross City, 29°41'N, 

83°04'W, 4 Dec 1993, Buck 24354; Duval County: Big Talbot Island St. Park, ca. 0.7 mi S of northern 

boundary, on Lyonia ferruginea, 17 Dec 1987, Buck 15513, Jacksonville, Calkins; Escambia County: along Co. 

Rd. 99A at Brushy Creek, 4.3 mi W of Fla. 97 at Walnut Hill, 30°53'N, 87°32'W, 8 Dec 1993, on Acer, Buck 

24550, on Ilex, Harris 31933; Franklin County: Apalachicola National Forest, Hickory Landing Rec. Area, ca. 2 

mi S of Sumatra, 23 May 1976, Harris 11304; Gilchrist County: Waccasassa Flats, along Co. Rd. 232, ca. 7.5 

mi N of Trenton, 29°44'N. 82°48'W, 5 Dec 1993, Buck 24415; Hamilton County: Bee Haven Bay, N of Co. Rd. 

6, E of Jasper, 30°30'N, 82°45'W, on branches of Taxodium,15 Dec 1993, Harris 32472, Holton Creek Wildlife 

Management Area, 30°26'N, 83°04'W, on Quercus, 5 Dec 1994, Harris 36099; Jackson County: Black Spring, 

ca. 6 mi S of Marianna, 3 Dec 1988, Harris 23330, 23346; Jefferson County: E of Fla. 59, 8.4 mi S of Wacissa, 

30°16'N, 84°03'W, 12 Dec 1993, Buck 24885; Lafayette County: along Co. Rd. 357, 6.4 mi Se of Fla. Hwy. 51, 

29°51'N, 83°15'W, 2 Dec 1994, Buck 27308; Lake County: Ocala National Forest, ca. 2 mi SE of Alexander 

Springs, on Magnolia virginiana, 6 Dec 1988, Buck 16735; Liberty County: Apalachicola National Forest, 

Ochlockonee River at Forest Serv. Rd. 13 crossing, 3 May 1990, Harris 25026; Madison County: 0.3 mi on dirt 

road W of Co. Rd. 150, NNE of Greenville, 30°36'N, 83°33'W, on Quercus, 14 Dec 1993, Buck 24888, Harris 

32369, 32382; Marion County: Ocala National Forest, ca. 0.75 mi NE of Doe Lake, 11 Dec 1988, Buck 16901, 

just E of Oklawaha River of Fla. Hwy. 316, 10 Aug 1985, Harris 18036; Polk County: Nalcrest, on Ilex, 28 May 

1989, Wheeler s.n.; Santa Rosa County: Blackwater River State Forest, Jackson Red Ground Trail from Red 

rock Rd., ca. 12 mi NE of Milton, 30°47'N, 86°53'W, 9 Dec 1993, Buck 24623, 24624, Harris 31990, 32002; 

Wakulla County: Edward Ball Wakulla Springs State Park, ca. 30°14'N, 84°18'W, on bamboo culm, 26-27 Dec 

1990, Buck 18991-A. GEORGIA. Charlton County: Okefenokee National Wildlife Refuge, N edge of Middle 

Island, 22 Nov 1989, Wetmore 65012; Coffee County: Broxton Rocks Ecological Preserve, on dead azalea, 

16-17 Dec 1993, Harris 32722, Ricketson Tract, on Vitis, 4 Feb 1995, Buck 27481. MISSISSIPPI. Winston 

County: Tombigbee National Forest, Choctaw Wildlife Area, 33°16'N, 89°06'W, on Magnolia, 28 Sep 1992, 

Harris 28661. NORTH CAROLINA. Dare Couny: Kitty Hawk, Nearing s.n., Nags Head Woods Ecological 

Preserve, 35°58'N, 75°06'W, on Liquidambar, 29 Sep 1993, Harris 31031 (all NY). 

CUBA. Prov. Santiago de Cuba: vicinity of Museo de la Francesca, "La Isabelita," ca. 2 km SE of La 

Gran Piedra, ca. 1000 m, 7 Apr 1982, Harris 14599 (HAC, NY). PUERTO RICO. Distr. Mayagüez: Maricao State 

Forest, Area de Recreo "Lookout," along Hwy 120 at Km 13.9, ca. 800 m, 28 May 1989, Buck 17241, 17265 

(all NY). 

CLADONIACEAE Zenker 

CLADINA (Nyl.) Nyl. 

1. Podetia yellowish, usnic acid present; PD+ red or PD-; branching dichotomous.................................................................................. 2 

2. Colonies without dense rounded heads; PD+ red (fumarprotocetraric acid); 

throughout Florida ................................................................................................................... C. subtenuis (des Abb.) Hale & Culb. 

2. Colonies with dense rounded heads; PD- (atranorin and perlatolic acid); 

throughout Florida ........................................................................................................................ C. evansii (des Abb.) Hale & Culb. 

1. Podetia not yellowish, usnic acid absent; PD+ red, fumarprotocetraric acid present............................................................................ 3 

3. Atranorin present; branching dichotomous or whorled........................................................................................................................ 4 

4. Branching dichotomous; pycnidial gel reddish; Everglades and Panhandle 

extending into southern Georgia....................................................................................................... C. sandstedei (des Abb.) Ahti 

4. Branching whorled; pycnidial gel colorless; Duval County (I have come to 

30

doubt the occurrence of this species in Florida. It needs to be confirmed 

with recent collections) ...................................................................................................................................[C. rangiferina (L.) Nyl.] 

3. Atranorin absent; branching dichotomous; Bay, Franklin and Hamilton counties ....................... C. subtenuis f. cinerea (Ahti) Ahti 

CLADONIA Hill ex Browne 

1. Apothecia and/or pycnidia red................................................................................................................................................................. 2 

2. Podetia simple or sparsely branched................................................................................................................................................... 3 

3. Podetia corticate, not sorediate/decorticate .................................................................................................................................... 4 

4. Thallus KOH+, PD+ deep yellow (thamnolic acid)........................................................................................................................ 5 

5. Usnic acid absent; podetia gray................................................................................................................................................ 6 

6. Podetia well developed, often branched; squamules small, often dissolving into 

sorediate masses; didymic acid; throughout Florida ................................................................................. C. ravenelii Tuck. 

6. Podetia poorly developed; squamules large, at most marginally sorediate, 

with broad yellow band below; pigments; throughout Florida ..............................................................C. hypoxantha Tuck. 

5. Usnic acid present; podetia yellowish, short, turbinate; didymic and squamatic acids 

or in one collection from Hillsborough County didymic acid is replaced by grayanic 

acid; often on Taxodium; throughout Florida .......................................................................................... C. abbreviatula Merr. 

4. Thallus KOH-, PD-; usnic acid present ......................................................................................................................................... 7 

7. Barbatic acid; squamules esorediate; northern tier of counties ................................................................. C. cristatella Tuck. 

7. Grayanic and squamatic acids; squamules often marginally sorediate; 

Lake, Liberty and Polk counties ........................................................................................................... C. anitae Culb. & Culb. 

3. Podetia sorediate or becoming mostly decorticate.......................................................................................................................... 8 

8. Barbatic and/or thamnolic acid...................................................................................................................................................... 9 

9. Soredia granular...................................................................................................................................................................... 10 

10. Podetia becoming decorticate; exposed stereome translucent; didymic acid; 

throughout Florida ............................................................................................................................................................. 11 

11. Barbatic acid........................................................................................................ C. didyma (Fée) Vainio var. didyma 

11. Thamnolic acid, often with accessory barbatic acid ......................C. didyma var. vulcanica (Zoll. & Moritzi) Vainio 

10. Podetia mostly corticate with a few sorediate patches....................................................................see C. ravenelii above 

9. Soredia farinose; barbatic acid; exposed stereome opaque; Leon County................................................... C. bacillaris Nyl. 

8. Squamatic, didymic and usnic acids; squamules abundantly sorediate; 

podetia very short; south to Polk County ......................................................................................................C. incrassata Flörke 

2. Podetia abundantly branched, forming lax, tangled colonies on sand; usnic, didymic, 

baeomycesic and squamatic acids; throughout Florida ................................................................................................C. leporina Fr. 

1. Apothecia and/or pycnidia brown or pallid............................................................................................................................................ 12 

12. Squamules smaller, not loose from substrate and curling into balls.............................................................................................. 13 

13. Podetia repeatedly dichotomously branched, "shrubby"; thallus PD- 

or PD+ yellow (baeomycesic acid); usnic acid present .............................................................................................................. 14 

14. Cortex lacking or poorly developed........................................................................................................................................ 15 

15. Podetia greenish with poorly developed cortex; podetial wall thin; cartilaginous 

tissue more continuous; Rock Hill Preserve,Washington County................................................................................... 16 

31

16. Thallus coarse, generally somewhat upright to sprawling ; 

tips abruptly narrowed, generally not curved downward; 

usnic acid only in Florida material .................................................................C. caroliniana Schweinitz ex Tuck. 

16. Thallus thin; tips gradually narrowing and downward pointing; 

usnic acid only in Florida material .................................................................................. C. dimorphoclada Robbins 

15. Podetia gray-white, ecorticate; podetial wall very thick, with scattered 

cartilaginous strands; south to Broward County ........................................................................ C. pachycladodes Vainio 

14. Cortex well developed............................................................................................................................................................ 17 

17. Podetia coarse, erect, hollow, perforate; PD-, squamatic acid; 

Highlands and Okaloosa counties ........................................................................................................ C. perforata Evans 

17. Podetia delicate, in lax, tangled, recumbent colonies; PD+ yellow 

(baeomycesic and squamatic acids); north to Polk County......................................... C. subsetacea Robbins ex Evans 

13. Podetia simple or sparingly branched........................................................................................................................................ 18 

18. Thallus PD-............................................................................................................................................................................ 19 

19. Podetia not forming cups ................................................................................................................................................ 20 

20. Squamules coarse, maculate; podetia proliferating, perforate, 

broadly expanded above; numerous apothecia on ultimate 

proliferations; squamatic acid; on Taxodium; Franklin and 

Liberty counties........................................................................................................................... C. buckii R. C. Harris 

20. Squamules thinner, emaculate; podetia mostly simple, mostly decorticate, 

narrower than the apothecia; barbatic acid; Duval and Seminole counties................................................................... 

................................................................................................................................... C. botryocarpa Merr. ex Sandst. 

19. Podetia forming broad cups, coarsely sorediate/areolate; grayanic acid; 

northern Florida .........................................................................................................................C. grayi Merr. ex Sandst. 

18. Thallus PD+ yellow, orange or orange-red ........................................................................................................................... 21 

21. Thallus PD+ yellow or orange .......................................................................................................................................... 22 

22. Podetia not forming cups ........................................................................................................................................... 23 

23. Squamules linear, large; podetia rarely squamulose, broader at top; 

norstictic acid or norstictic and stictic acids ......................................................................................................... 24 

24. Norstictic and stictic acids; south to Polk County ..............................................................C. polycarpia Merr. 

24. Norstictic acid ± atranorin; northern Florida.................................................................. C. polycarpoides Nyl. 

23. Squamules irregular, small; podetia often squamulose; thamnolic or 

baeomycesic acid ................................................................................................................................................. 25 

25. Thallus KOH+ deep yellow (thamnolic and barbatic acids)........................................................................... 26 

26. Cortex dispersed, coarsely areolate/squamulose; podetia 

short, rarely branched; central Florida ......................................................................... C. santensis Tuck. 

26. Cortex continuous; podetia taller, with few squamules, 

commonly branched, especially above; south to Polk County .................................... C. floridana Vainio 

25. Thallus KOH- ................................................................................................................................................. 27 

27. Squamules and podetia C-, KC-(baeomycesic and 

squamatic acids); podetia densely squamulose; 

south to Charlotte County..................................................................... C. beaumontii (Tuck.) Vainio 

27. Squamules and podetia C+, KC+ blue-green (strepsilin, 

baeomycesic and squamatic acids); podetia usually not 

densely squamulose, often lacking; rare, Holmes and 

Washington counties ........................................................................... C. strepsilis (Ach.) Grognot 

32

22. Podetia forming abruptly flared, centrally proliferating cups; 

psoromic acid; north to Seminole County..........................................................................C. rappii Evans var. rappii 

21. Thallus PD+ orange-red (fumarprotocetraric acid).......................................................................................................... 28 

28. Podetia forming broad cups ....................................................................................................................................... 29 

29. Cups not centrally proliferating, sorediate/areolate.............................................................................................. 30 

30. Fumarprotocetraric and either grayanic or cryptochlorophaeic acids ............................................................ 31 

31. Grayanic acid (this strain may not be in Florida) .................................................C. grayi Merr. ex Sandst. 

31. Cryptochlorophaeic acid; rare, Franklin County ..................................C. cryptochlorophaea Asahina 

30. Fumarprotocetraric acid only; reported from peninsular Florida 

by Moore, NY holds a couple of dubious records................................. C. chlorophaea (Sommerf.) Sprengel 

29. Cups centrally proliferating, abruptly flared; smoothly 

corticate; south to Polk County..........................................................C. rappii Evans var. exilior (Abbayes) Ahti 

28. Podetia not forming cups or forming very narrow, irregular cups ............................................................................ 32 

32. Podetia not sorediate ........................................................................................................................................... 33 

33. Podetia simple, short; apothecia much exceeding stalk or 

podetia gradually swelling to the apothecia.................................................................................................... 34 

34. Squamules small, unlobed, appressed, imbricate; 

apothecia exceeding stalk; south to Polk County................................... C. peziziformis (With.) Laundon 

34. Squamules larger, somewhat lobed, usually recurved; 

apothecia ± equal to stalk; Duval County, a somewhat 

dubious Calkins collection....................................................................................... C. sobolescens Nyl. 

33. Podetia taller, branched, often with poorly developed cups; 

squamules lobed, erect to recurved; south to Polk County............................................... C. simulata Robbins 

31. Podetia sorediate ................................................................................................................................................. 35 

35. Primary squamules at most marginally sorediate........................................................................................... 36 

36. Podetia awl-shaped or irregular; soredia farinose or 

granular; grayanic acid lacking [C. ochrochlora Flörke 

will key here. There is no trustworthy record for 

the state] .................................................................................................................................................... 37 

37. Podetia with farinose to granular soredia above, 

granular soredia mixed with isidioid squamules below; 

cortex dissolved except for basal collar; throughout 

Florida....................................................................................................C. subradiata (Vainio) Sandst. 

37. Podetia often mostly or partly corticate with patches 

of coarse soredia; exposed stereome translucent; 

throughout Florida ..........................................................................C. ramulosa (With.) Laundon s.lat. 

36. Podetia short, stubby with coarse granular soredia; 

containing grayanic acid; rare, Union County...........................................C. cylindrica (Evans) Evans 

35. Primary squamules dissolving into a sorediate crust; 

podetia tiny, pointed; Franklin County.......................................................................................... C. nana Vainio 

12. Squamules very large, curled into balls when dry; usually without podetia; 

on sand; south to Polk County ............................................................................................................................ C. prostrata Evans 

BIATORACEAE Stizenb. 

33

PHYLLOPSORA Müll. Arg. 

Note: This treatment mainly follows Brako (1990). Sterile isidiate crusts which are doubtfully 

squamulose should also be sought in the key to sterile isidiate crusts. 

1. Thallus PD- ............................................................................................................................................... 2 

2. Prothallus pale (in North America rarely darkening in old, abused specimens).................................... 3 

3. Squamules with marginal proliferations ± parallel to the substrate, flattened 

to some degree and not distinctly cylindrical; proliferations initially simple but 

usually repeatedly branched or proliferate ....................................................................................... 4 

4. Squamules large; lobes more than 0.5 mm wide .......................................................................... 5 

5. Marginal proliferations usually abundant, narrow, mostly ecorticate, 

forming a distinct white fringe; cortex less than 50 µm thick; ascospores 

ellipsoid to short fusiform, 8-12.5 × 2-3 µm; south to Highlands 

County ................................................................P. parvifolia (Pers.) Müll. Arg. var. parvifolia 

5. Marginal proliferations scarce, broader, similar to primary squamules; 

cortex over 50 µm thick; ascospores ovoid, 7-10.5 × 2.5-4 µm; Alabama, 

not yet known from Florida........................................ [P. parvifolia var. breviuscula (Nyl.) Brako] 

4. Squamules smaller, 0.1-0.5 mm wide .......................................................................................... 6 

6. Squamules adnate; proliferations narrow in respect to squamule, 

initially ± ovate becoming elongated, with constrictions or branched ......................................7 

7. Cortex type 2; very irregular, ± non-linear proliferations usually 

abundant and forming overlapping layers; ascospores 8.5-12 

× 2.5-3 µm; common, south to Hillsborough County.............................................................. 

........................................................................................... P. confusa Swinscow & Krog 

7. Cortex type 1; proliferations less abundant less commonly elongated; 

ascospores 7.5-11.5 × 2-3 µm; reported by Brako from Alachua 

and Duval counties.................................... P. corallina (Eschw.) Müll. Arg. var. corallina 

6. Squamules erect; proliferations broad in respect to squamule or 

squamules adnate; proliferations coarse and inflated-looking, not 

much elongated ....................................................................................................................... 8 

8. Squamules ± erect with broad, flat lobules; ascospores 

4-6.5 × 2-3 µm; Lake County ............................................. P. parvifoliella (Nyl.) Müll. Arg. 

8. Squamules closely adnate, often contiguou and forming ± 

continuous crust; proliferations coarse, appearing inflated; 

an abberant population from Sharkey County, Mississippi, 

not yet known from Florida; referred in hb. by Brako 

to .............................................................................................. [cf. P. corallina var. corallina] 

3. Squamules with ± globose to cylindrical isidia or squamules lacking isidia 

and marginal proliferations .............................................................................................................. 9 

9. Thallus isidiate ............................................................................................................................ 10 

10. Squamules medium, 0.1-0.5 mm wide ................................................................................. 11 

34

11. Isidia cylindrical .............................................................................................................. 12 

12. Isidia common, often obpyriform, constricted at base 

and broadest just above base; atranorin and parvifoliin; 

ascospores 6.5-10 × 2.5-3.5 µm; south to 

Sarasota County .................................................... P. corallina var. rappiana Brako 

12. Isidia rare, cylindrical; squamules mostly with marginal 

proliferations; no substances; seen by me only from 

South Carolina........................................[P. corallina (Eschw.) Müll. Arg. var. corallina] 

11. Isidia nearly globose or irregular, not much elongated or weakly 

elongated by buddiing; no substances; ascospores 6.5-12.5 

× 2-3.5 µm; Louisiana and Oklahoma, not yet known from 

Florida ....................................................................................................... [P. kalbii Brako] 

10. Squamules/areoles minute, ca. 0.1 mm wide; isidia long, 

cylindrical, tangled; no substances; ascospores 

7.5-12.5 × 2-3.5 µm; Leon County .............................................P. isidiolyta (Vainio) Riddle 

9. Thallus not isidiate or lobulate; see couplet 5 ................................. [P. parvifolia var. breviuscula] 

2. Prothallus dark, abundant ................................................................................................................. 13 

13. Squamules minute, inconspicuous; thallus dominated by long 

slender isidia; furfuracein; ascospores 6-12 × 2-2.5 µm; 

Dade and Seminole Counties ...........................................................P. furfuracea (Pers.) Zahlbr. 

13. Squamules/areoles large, tightly adjoined forming an almost 

continuous crust, separated by white lines formed by fibrils, 

isidiate; isidia sparse, short, cylindrical; atranorin; Louisiana, 

not yet known from Florida ........................................................................[P. halei (Tuck.) Zahlbr.] 

1. Thallus PD+ orange (argopsin agg. or pannarin) .................................................................................... 14 

14. Thallus isidiate, not pruinose (P. corallina (Eschw.) Müll. Arg. s. lat.)............................................... 15 

15. Containing argopsin agg. as major substances .......................................................................... 16 

16. Lacking phyllopsorins; ascospores 7-13 × 2.5-3 µm; 

common, south to Sarasota County........................P. corallina var. santensis (Tuck.) Brako 

16. Containiing phyllopsorins; ascospores 6.5-12 × 2.5-3.5 µm; 

reported by Brako from Florida without locality (prob. Lake 

County) ...............................................................P. corallina var. ochroxantha (Nyl.) Brako 

15. Containing vicanicin as major substance with traces of 

norvicanicin and argopsin; ascospores 7-11 × 2.5-3 µm; 

reported by Brako from Dade and Seminole 

counties ........................................................................P. corallina var. glaucella (Vainio) Brako 

14. Thallus not isidiate, pruinose; squamules large 

(P. buettneri (Müll. Arg.) Zahlbr. s. lat.) ............................................................................................ 17 

35

17. Containing argopsin agg.; ascospores 7-12 × 2.5-3 µm; 

Wakulla County ...................................................P. buettneri var. glauca (B. de Lesd.) Brako 

17. Containing pannarln, phyllopsorin and zeorin; ascospores 

10-12 × 2.5-3 µm; reported by Brako from Louisiana, 

not yet found in Florida.................................................. [P. buettneri var. munda (Malme) Brako] 

NOTES 

Phyllopsora buettneri (Müll. Arg.) Zahlbr. var. glauca (B. de Lesd.) Brako 

This variety of P. buettneri has not been reported from North America previously. The specimen was 

determined by Lois Brako. 

FLORIDA. Wakulla County: Swamp forest, Lost Creek at Forest Service Rd. 309, just S of Leon County 

line, 4 Dec 1988, Harris 23375 (NY). 

PARMELIACEAE Zenker 

KEY TO THE GENERA 

1. Thallus foliose; central strand absent ..................................................................................................................................................... 2 

2. Lobes ciliate on margins or in axils...................................................................................................................................................... 3 

3. Cilia not swollen at base ................................................................................................................................................................... 4 

4. Lobes broad, 4-20 mm wide; cilia usually conspicuous ............................................................................................................... 5 

5. Underside pale at margin, black in center, with only one type of rhizine ................................................................................. 6 

6. Cortex not or only weakly maculate, not or only slightly cracked; 

rhizines lacking in marginal zone; chemistry various......................................................................................... Parmotrema 

6. Cortex maculate and abundantly cracked; rhizines usually reaching margin; 

salazinic acid .............................................................................................................................................................. Rimelia 

5. Underside uniformly pale to medium brown, with numerous short rhizines 

and fewer long ones ; Florida species isidiate; rare (previously 

included in Parmotrema) ......................................................................................................................................... Rimeliella 

4. Lobes narrow, 0.5-6.0 mm wide; cilia inconspicuous................................................................................................................... 7 

7. Medulla yellow, especially below soralia, C+ yellow (pigments, terpenoids); 

soralia pustular, breaking down to form coarse soredia; south to Seminole County.................................................................. 

................................................................................................................................Myelochroa aurulenta (Tuck.) Elix & Hale 

7. Medulla white, C+ pink (gyrophoric or hiasic acid complex) ............................................................................. Parmelinopsis 

3. Cilia swollen at base (bulbate) ......................................................................................................................................................... 8 

8. Cortex gray (atranorin) ................................................................................................................................................. Bulbothrix 

8. Cortex yellow (usnic acid) ................................................................................................................................................ Relicina 

2. Lobes not ciliate on margins or in axils................................................................................................................................................ 9 

9. Thallus not pseudocyphellate......................................................................................................................................................... 10 

10. Cortex or medulla yellow.......................................................................................................................................................... 11 

11. Lobes narrow, 0.5-6.0 mm wide.......................................................................................................................................... 12 

12. Cortex yellow (usnic acid) ............................................................................................................................................... 13 

13. Growing on bark; thallus pustulate; pustules breaking down 

to form coarse soredia; divaricatic acid; south to Lake 

County................................................................................................................ Parmeliopsis subambigua Gyelnik 

13. Growing on sandstone; isidiate; stictic acid agg.; known only from 

Rock Hill, Washington County, new to Florida ......................................Xanthoparmelia conspersa (Ach.) Hale 

36

12. Medulla yellow; thallus not sorediate or isidiate ............................................................................................................. 14 

14. Medulla C+, KC+ orange (pigments); apothecia laminal; common, 

throughout Florida ............................................................................... Pseudoparmelia sphaerospora (Nyl.) Hale 

14. Medulla C-, KC- (vulpinic acid); apothecia marginal; 

Jackson and Walton counties....................................................... Vulpicida viridis (Schweinitz) Mattson & Lai 

11. Lobes broad, 4-20 mm wide............................................................................................................................... Parmotrema 

10. Cortex gray or brownish; medulla white...................................................................................................................................... 15 

15. Lobes narrow, 0.5-6.0 mm wide............................................................................................................................................ 16 

16. Thallus with "normal" rhizines, thick felty rhizine mat lacking.......................................................................................... 17 

17. Cortex KOH+ yellow; pycnidia laminal........................................................................................................................ 18 

18. Underside with inconspicuous pale to brown, simple rhizines............................................................................. 19 

19. Cortex KOH+ pale yellow (atranorin) .......................................................................................... Canoparmelia 

19. Cortex KOH+ deep yellow, PD+ deep yellow (thamnolic acid); 

isidiate; lobes linear; cortex shiny; Duval County.................................Imshaugia aleurites (Ach.) Meyer 

18. Underside with conspicuous dark, dichotomously branched rhizines; 

medulla KC+ purple (lividic acid agg.) .............................................................................................. Hypotrachyna 

17. Cortex KOH-; pycnidia conspicuous, marginal; fatty acids; 

on pine; south to Lake County ......................................................................... Tuckermanopsis fendleri (Nyl.) Hale 

16. Thallus with a thick spongy mat of rhizines below; marginally isidiate; 

divaricatic acid; rare, Escambia County, new to Florida ..........................................Anzia ornata (Zahlbr.) Asahina 

15. Lobes broad, 4-20 mm wide................................................................................................................................. Parmotrema 

9. Thallus white pseudocyphellate, isidiate; medulla C+ red (lecanoric acid); 

south to Seminole County ...............................................................................................................Punctelia rudecta (Ach.) Hale 

1. Thallus fruticose with a central solid cartilaginous strand ............................................................................................................. Usnea 

ANZIA Stizenb. 

Anzia ornata (Zahlbr.) Asahina has previously been reported from Alabama and North Carolina. Thus, its occurrence in 

northernmost Florida is not especially notable. FLORIDA. Escambia County: along Co. Rd. 99A at Brushy Creek, 4.3 mi W of St. 

Rd. 97 at Walnut Hill, 30°53'N, 87°32'W, floodplain woods, on Magnolia, 8 Dec 1993, Harris 31913 (NY). 

BULBOTHRIX Hale 

1. Thallus not isidiate; apothecia present ................................................................................................................................................... 2 

2. Apothecia lacking bulbate rhizines (ecoronate); medulla C+ red (lecanoric acid); 

south to Polk County ............................................................................................................................ B. confoederata (Culb.) Hale 

2. Exciple with bulbate rhizines (coronate); medulla C+ pink (gyrophoric acid); 

Texas, not yet known from Florida ................................................................................................................. [B. coronata (Fée) Hale] 

1. Thallus isidiate; apothecia absent........................................................................................................................................................... 3 

3. Medulla C+ pink to red, KOH-; lobes linear ......................................................................................................................................... 4 

4. Gyophoric acid present; south to Polk County.........................................................................................B. goebelii (Zenker) Hale 

4. Lecanoric acid present; south to Lake County ........................................................................................ B. laevigatula (Nyl.) Hale 

3. Medulla C-, KOH+ red (salazinic acid); lobes short, truncate to rounded; 

isidia often globose, brown; south to Polk County...............................................................................................B. isidiza (Nyl.) Hale 

NOTES 

37

Bulbothrix isidiza (Nyl.) Hale 

In the previous edition I speculated that this species might be recently introduced. However, an examination of Moore's specimens 

of Canoparmelia salacinifera revealed several collections of B. isidiza, putting an end to this notion. 

CANOPARMELIA Elix & Hale 

1. Thallus isidiate........................................................................................................................................................................................ 2 

2. Medulla PD+ orange or red................................................................................................................................................................. 3 

3. Medulla KOH- (protocetraric acid) .................................................................................................................................................. 4 

4. Isidia globose to cylindrical; throughout Florida....................................................................... C. amazonica (Nyl.) Elix & Hale 

4. Isidia inflated, somewhat lobulate; Monroe County (Keys) ................................................... C. martinicana (Nyl.) Elix & Hale 

3. Medulla KOH+ red (saiazinic acid); lobe tips unmarked; 

south to Highlands County ....................................................................................................... C. salacinifera (Hale) Elix & Hale 

2. Medulla PD- (perlatolic acid); lobe tips with reticulate 

white maculae; throughout Florida...................................................................................................C. caroliniana (Nyl.) Elix & Hale 

1. Thallus sorediate ..................................................................................................................................................................................... 5 

5. Soralia capitate on upturned lobules; medulla KC+ purple (cryptochlorophaeic acid); 

throughout Florida .............................................................................................................C. cryptochlorophaea (Hale) Elix & Hale 

5. Soralia pustular, laminal; medulla KC+ pinkish (divaricatic acid); Leon and 

Seminole counties according to Moore, not seen by me..................................................................... C. texana (Tuck.) Elix & Hale 

See Parmeliopsis. 

FORAMINELLA Fricke-Meyer 

HYPOTRACHYNA (Vainio) Hale 

1. Thallus pustular-sorediate....................................................................................................................................................................... 2 

2. Thallus UV+ yellow (lichexanthone); very common, south to 

Polk County ................................................................................................................................. H. osseoalba (Vainio) Park & Hale 

2. Thallus UV- (atranorin); Duval and Wakulla counties ........................................................................ H. pustulifera (Hale) Skorepa 

1. Thallus not pustulate-sorediate, UV-; apothecia common; south to Lake County.............................................. H. livida (Taylor) Hale 

IMSHAUGIA Fricke-Meyer 

Only I. aleurites (Ach.) Meyer known from Florida. Previously treated as Parmeliopsis. 

MYELOCHROA (Asahina) Elix & Hale 

Only the single species M. aurulenta (Tuck.) Elix & Hale is known from Florida. 

PARMELINOPSIS Elix & Hale 

1. Thallus isidiate......................................................................................................................................................................................... 2 

2. Isidia usually tipped with ciiia; hiasic acid complex; south to 

Pinellas County............................................................................................................................ P. horrescens (Taylor) Elix & Hale 

2. Isidia usually eciliate; gyrophoric acid and a UV+ white unknown; 

northern part of the state ................................................................................................................. P. minarum (Vainio) Elix & Hale 

1. Thallus sorediate; gyrophoric acid and a UV+ white unknown .............................................................................................................. 3 

3. Soredia coarse, derived from pustules; Marion, Wakulla and 

Walton counties........................................................................................................................... P. spumosa (Asahina) Elix & Hale 

38

3. Soredia farinose, not derived from pustules; Leon County .....................................................P. cryptochlora (Vainio) Elix & Hale 

PARMELIOPSIS (Stizenb.) Nyl 

Only the single species P. subambigua Gyelnik is known from Florida. It was treated in the previous edition as Foraminella. 

PARMOTREMA Massal. 

1. Lobes lacking marginal cilia .................................................................................................................................................................... 2 

2. Thallus isidiate...................................................................................................................................................................................... 3 

3. Medulla white, C+ red (lecanoric acid); common, throughout Florida .................................................... P. tinctorum (Nyl. ) Hale 

3. Medulla yellow. C+ pink (gyrophoric acid, ± echinocarpic acid); 

throughout Florida ...................................................................................................................... P. endosulphureum (Hillm.) Hale 

2. Thallus sorediate or lacking both soredia and isidia ........................................................................................................................... 4 

4. Thallus sorediate .............................................................................................................................................................................. 5 

5. Medulla KOH-, PD+ red or PD-..................................................................................................................................................... 6 

6. Medulla PD+ red (protocetraric acid)........................................................................................................................................ 7 

7. Soralia white ........................................................................................................................................................................... 8 

8. Protocetraric acid only; common, throughout Florida.................................................... P. gardeneri (Dodge) Sérusiaux 

8. Protocetraric acid with echinocarpic acid, usnic acid and unknowns; 

common, throughout Florida ......................................................................................................P. dilatatum (Vainio) Hale 

7. Soralia yellowish (usnic acid); south to Polk County, 

reported by Moore, not seen by me ....................................................................................... P. dominicanum (Vainio) Hale 

6. Medulla PD- (fatty acids); generally smaller than most Parmotrema; 

common, throughout Florida ....................................................................................................P. praesorediosum (Nyl.) Hale 

5. Medulla KOH+ red, PD+ orange ................................................................................................................................................... 9 

9. Salazinic acid present; common, Peninsular Florida.................................................................... P. cristiferum (Taylor) Hale 

9. Norstictic acid present; Dade and Volusia counties ........................................................................P. rubifaciens (Hale) Hale 

4. Thallus not sorediate; medulla PD+ red (protocetraric acid); conidia sublageniform; 

Dade and Monroe counties.......................................................................................................................P. zollingeri (Hepp) Hale 

1. Lobes with marginal cilia ....................................................................................................................................................................... 10 

10. Thallus isidiate................................................................................................................................................................................. 11 

11. Cortex or medulla yellow........................................................................................................................................................... 12 

12. Cortex yellowish (usnic acid); medulla white; medulla C- (fatty acids) 

or C+ pink (gyrophoric acid, i.e., including P. madagascariaceum (Hue) Hale); 

south to Marion County ........................................................................................................P. xanthinum (Müll. Arg.) Hale 

12. Medulla bright yellow (vulpinic acid); common, Peninsular 

Florida...................................................................................................................... P. sulphuratum (Nees & Flotow) Hale 

11. Cortex gray; medulla white......................................................................................................................................................... 13 

13. Medulla KOH+ yellow or red, UV- or UV+ yellow in lower part ............................................................................................ 14 

14. Medulla KOH+ yellow (stictic acid agg.); throughout Florida ........................................................P. crinitum (Ach.) Hale 

14. Medulla KOH+ red (salazinic acid); lower medulla UV+ yellow 

(lichexanthone); south to Marion County .................................................................................P. ultralucens (Krog) Hale 

39

13. Medulla KOH-, UV+ bright blue white (alectoronic acid); isidia often ciliate; 

south to Lake County........................................................................................................................ P. mellissii (Dodge) Hale 

2. Thallus sorediate or neither sorediate nor isidiate................................................................................................................................ 15 

15. Thallus sorediate ............................................................................................................................................................................. 16 

16. Medulla KOH-, UV+ bright blue white (alectoronic acid); cortex not maculate; 

common throughout Florida ............................................................................................................ P. rampoddense (Nyl.) Hale 

16. Medulla KOH+ yellow (stictic acid agg.); cortex maculate; 

common throughout Florida ......................................................................................................P. hypoleucinum (Steiner) Hale 

15. Thallus not sorediate....................................................................................................................................................................... 17 

17. Cortex maculate; medulla PD+ yellow (norstictic acid) or PD-................................................................................................. 18 

18. Norstictic acid only present (rarely acid deficient); 

south to Highlands County ....................................................................................................P. perforatum (Jacq.) Massal 

18. Alectoronic acid with or without norstictic acid present; 

common, throughout Florida ..........................................................................................................P. rigidum (Lynge) Hale 

17. Cortex not maculate; medulla PD+ red (protocetraric acid); conidia rod-shaped; 

throughout Florida .................................................................................................................... P. michauxianum (Zahlbr.) Hale 

PSEUDOPARMELIA Lynge 

Only the single species P. sphaerospora (Nyl.) Hale is known from Florida. 

PUNCTELIA Krog 

Only the single species P. rudecta (Ach.) Krog is known from Florida. 

RELICINA (Hale & Kurok.) Hale 

1. Thallus isidiate; medulla KOH+ red (norstictic/salazinic acid agg.); 

Baker County, reported by Moore, not seen by me, but recently collected 

at Broxton Rocks, Coffee county, Georgia..................................................................................................... R. abstrusa (Vainio) Hale 

1. Thallus not isidiate or sorediate; medulla KOH-, PD+ red (fumarprotocetraric acid agg.); 

Monroe County (Keys)............................................................................................................................ R. eximbricata (Gyelnik) Hale 

RIMELIA Hale & Fletcher 

1. Thallus sorediate; throughout Florida .........................................................................................R. reticulata (Taylor) Hale & Fletcher 

1. Thallus isidiate; south to Polk County............................................................................... R. subisidiosa (Müll. Arg.) Hale & Fletcher 

RIMELIELLA Kurokawa 

1. Norlobaridone present; Bradford County........................................................................................R. subtinctoria (Zahlbr.) Kurok. 

1. Norlobaridone absent; Bradford and Wakulla counties ......................................................R. neotropica (Kurok. & Hale) Kurok. 

TUCKERMANOPSIS Gyelnik 

Only T. fendleri (Nyl.) Hale occurs in Florida. Tuckermanopsis viridis has been removed to Vulpicida. See key to genera. 

USNEA Dill. ex Adanson 

1. Cortex, medulla or axis red or pink ......................................................................................................................................................... 2 

2. Cortex not red; medulla or axis red or pink.......................................................................................................................................... 3 

3. Main axes hollow; medulla red or pink............................................................................................................................................. 4 

4. Medulla usually red; norstictic acid present; sorediate-isidiate; south to 

40

Lake County U. baileyi (Stirton) Zahlbr. 

4. Medulla usually pink; diffractaic acid; isidiate-sorediate or sorediate; south 

to Lee County ..............................................................................................................................................U. perplectata Motyka 

3. Main axes solid................................................................................................................................................................................. 5 

5. Thallus shrubby to pendant without numerous perpendicular branchlets; 

apothecia rare; medulla pink or axis red; isidiate-sorediate......................................................................................................... 6 

6. Axis reddish; fatty acids; south to Lake Okeechobee................................................................................ U. mutabilis Stirton 

6. Axis unpigmented; medulla pale pink; diffractaic acid; branches 

often coarse; Duval County..............................................................................................................................U. ceratina Ach. 

5. Thallus shrubby with numerous perpendicular branchlets; main branches often 

ending in apothecia; not sorediate or isidiate; medulla red; chemistry various; 

south to Lake Okeechobee ................................................................................................................U. strigosa (Ach.) A. Eaton 

2. Cortex red mottled; medulla and cortex not pigmented; shrubby to subpendant; 

isidiate or isidiate-sorediate; stictic acid agg. (or in one collection from Hamilton 

County galbinic acid agg.); south to Lake County .............................................................................................. U. rubicunda Stirton 

1. Cortex, medulla or axis not red or pink ................................................................................................................................................... 7 

7. Soredia and/or isidia present .............................................................................................................................................................. 8 

8. Thallus shruby or subpendant.......................................................................................................................................................... 9 

9. Medulla loose .............................................................................................................................................................................. 10 

10. Thallus not inflated; salazinic acid ....................................................................................................................................... 11 

11. Isidiate-sorediate; cortex with numerous raised pseudocyphellae; 

Lake and Leon counties ...........................................................................................................................Usnea sp. 23493 

11. Isidiate; cortex smooth and shiny; Dade County ............................................................................ Usnea sp. Small 5853 

10. Thallus inflated and somewhat articulate; isidiate-sorediate; 

protocetraric acid; Lake County ...................................................................................................................Usnea sp. 23508 

9. Medulla compact; isidiate-sorediate; protocetraric acid; south to 

Lake County................................................................................................................................ U. subscabrosa Nyl. ex Motyka 

8. Thallus pendant, often with numerous perpendicular branchlets; 

galbinic acid agg.; south to Polk County......................................................................................U. dimorpha (Müll. Arg.) Motyka 

7. Soredia and isidia absent; thallus long, pendent; axis brownish; diffractaic acid 

and/or constictic acid; south to Polk County............................................................................................................U. trichodea Ach. 

VULPICIDA Mattson & Lai 

Only a single species, V. viridis (Schweinitz) Mattson & Lai in Florida. It was included in Tuckermanopsis in the previous 

edition. See key to genera. 

XANTHOPARMELIA (Vainio) Hale 

Since non-calcareous rock is almost entirely lacking in Florida, it is only with recent access to the Rock Hill Preserve that 

this genus has been added to the lichenota. Only X. conspersa (Ach.) Hale is known from the state. Many thanks to Angus 

Gholson for guiding Bill Buck and myself around the site. 

PHYSCIACEAE Zahlbr. 


1. Thallus foliose.......................................................................................................................................................................................... 2 

41

2. Rhizines present.................................................................................................................................................................................. 3 

3. Upper cortex paraplectenchymatous; lower cortex present .......................................................................................................... 4 

4. Medulla white (bright red orange in Phaeophyscia rubropulchra), PD-; cortex UV-; 

apothecia with thalline margin..................................................................................................................................................... 5 

5. Cortex KOH+ yellow .....................................................................................................................................................Physcia 

5. Cortex KOH- ......................................................................................................................................................Phaeophyscia 

4. Medulla often orangish or yellowish, if white then cortex UV+ yellow 

or medulla PD+ orange; apothecia without thalline margin...............................................................................................Pyxine 

3. Upper cortex prosoplectenchymatous; lower cortex often lacking............................................................................Heterodermia 

2. Rhizines lacking or very sparse and poorly developed ...................................................................................................................... 6 

6. Divaricatic acid or sekikaic acid agg. present; underside black; lobes separable from 

the substrate; hypothecium brown .......................................................................................................................................Dirinaria 

6. No substances present; lobes entirely adnate to the substrate; hypothecium pale.................................................. Hyperphyscia 

1. Thallus crustose ...................................................................................................................................................................................... 7 

7. Ascospores with irregularly thickened walls ....................................................................................................................................... 8 

8. Ascospores small, under 30 µm long; hymenium not inspersed.............................................................................Rinodina s. lat. 

8. Ascospores larger, over 30 µm long; hymenium inspersed ................................................................................................ Hafellia 

7. Ascospores with uniformly thin walls or somewhat thickened only at 

septum (Buellia imshaugiana)............................................................................................................................................................. 9 

9. Conidiospores oblong or rod shaped......................................................................................................................................Buellia 

9. Conidiospores filiform, curved.........................................................................................................................................Amandinea 

BUELLIA De Notaris 

including AMANDINEA Scheid. & Mayrh. & HAFELLIA Kalb et al. 


2. Thallus PD- or PD+ yellow (norstictic or baeomycesic acid).............................................................................................................. 3 

3. Ascospores 8/ascus ........................................................................................................................................................................ 4 

4. Ascospores 15 µm or more long and 5 µm or more wide........................................................................................................... 5 

5. Apothecial disk naked ............................................................................................................................................................. 6 

6. Ascospores 2-celled ............................................................................................................................................................ 7 

7. Norstictic acid present; ascospore wall not or weakly ornamented ............................................................................... 8 

8. Hymenium without oil droplets or rarely with some oil in upper part.......................................................................... 9 

9. Ascospore wall thickened at septum; ascospores 

18-22 × 7-8.5(-10) µm; common, throughout Florida................................................ B. imshaugiana R. C. Harris 

9. Ascospore wall not thickened at septum; ascospores 

16-23 × 6-9 µm; common, south to Polk County..........................................................B. curtisii (Tuck.) Imshaug 

8. Hymenium with numerous oil droplets throughout ................................................................................................... 10 

10. Ascospores 14-22 × 6-8 µm ................................................................................................................................ 11 

11. Hymenium greenish above, KOH+ purplish; common, 

throughout Florida ..................................................................................... Hafellia bahiana (Malme) Sheard 

11. Hymenium brownish above, KOH-; throughout Florida.................................................. B. curatellae Malme 

42

10. Ascospores 22-41 × 11-21 µm............................................................................................................................ 12 

12. Ascospores 22-31 × 11-13 µm, darkly pigmented, wall pitted; 

northern Florida ..............................................................Hafellia callispora (Knight) H. Mayrh. & Sheard 

12. Ascospores 30-41 × 16-21 µm, lightly pigmented; 

wall not pitted; south to Sarasota County ...................................................... Hafellia parastata (Nyl.) Kalb 

7. Norstictic acid lacking; ascospore wall strongly ornamented; 

ascospores 14-20 × 8-11 µm; peninsular Florida .......................................................................B. leucomela Imshaug 

6. Ascospores becoming 4-celled, 17-21 × 6-8 µm; norstictic acid; 

north to Seminole County..................................................................................................B. lauricassiae (Fée) Müll. Arg. 

5. Apothecial disk yellow-green pruinose; thallus yellowish (xanthones); 

ascospores 19-28 × 8-10 µm; reported by Imshaug (1955) but 

not seen ................................................................................................................. B. melanochlora (Krempelh.) Müll. Arg . 

4. Ascospores 15 µm or less long and mostly 5 µm or less wide................................................................................................. 13 

13. Thallus not pigmented......................................................................................................................................................... 14 

14. Hymenium without oil droplets ...................................................................................................................................... 15 

15. Exciple at most paler brown within.......................................................................................................................... 16 

16. Lichen substances present (xanthones, atranorin, baeomycesic, 

barbatic, norstictic acids).................................................................................................................................... 17 

17. Containing other substances; exciple not paler within ................................................................................ 18 

18. Thallus PD-............................................................................................................................................. 19 

19. Thallus UV+ (lichexanthone and barbatic acid); 

disk not pruinose; ascospores 10-12.5 × 4-5.5 µm; 

throughout Florida .................................................................................... B. catasema (Tuck.) Tuck. 

19. Thallus UV-, KC+ orange (xanthones); disk white 

pruinose; ascospores 12-15 × 4.5-6 µm; Franklin 

and Seminole counties..........................................................................B. pachnidisca R. C. Harris 

18. Thallus PD+ yellow (baeomycesic and squamatic acids) 

(see below) ............................................................................................................. B. caloosensis Tuck. 

17. Atranorin and norstictic acid or atranorin only; exciple paler 

within; ascospores 10-15 × 5-7 µm; common, throughout 

Florida.......................................................................................................................... B. stillingiana Steiner 

16. Lichen substances lacking; tips of paraphyses swollen, 

dark-capped; ascospores 9-16 × 6-7 µm; throughout 

Florida ............................................................................... Amandinea punctata (Hoffm.) Coppins & Scheid. 

15. Exciple with a strongly differentiated colorless layer; ascospores 

9-12 × 4-5 µm; south to Lake County........................................................B. amphidexia Imshaug ex R. C. Harris 

14. Hymenium with abundant oil droplets; ascospores 9-13 × 4-5 µm; 

atranorin; Duval, Franklin, Lake and Seminole counties.........................................B. rappii Imshaug ex R. C. Harris 

13. Thallus containing bright pink or red pigment..................................................................................................................... 20 

20. Thallus thick, bullate, shiny, PD-; pigment red (xanthones and 

chiodectonic acid); medulla ascospores 8.5-12 × 4-5.5 µm; Dade 

and Liberty counties .............................................................................................................. B. coccinea (Fée) Aptroot 

20. Thallus thin, weakly areolate, PD+ yellow (baeomycesic and 

squamatic acids); pigment pink; ascospores 9-13 × 4-5.5 µm; 

Duval, Lake, Volusia and Wakulla counties ................................................................................. B. caloosensis Tuck. 

3. Ascospores more than 8/ascus..................................................................................................................................................... 21 

43

21. Hymenium with abundant oil droplets; ascospores 12-16/ascus; 

upper hymenium KOH+ purplish; south to Polk 

County............................................................................Hafellia bahiana (Malme) Sheard var. pleiotropa (Malme) Sheard 

21. Hymenium without oil droplets; ascospores 24-32/ascus; upper 

hymenium KOH-; south to Polk County.....................................................................Amandinea polyspora (Willey) ined. 

2. Thallus PD+ orange-red (fumarprotocetraric acid); ascospores 13.5-16.5 × 6.5-7.5 µm; 

Franklin, Levy, Liberty, Santa Rosa and Wakulla counties .................................................................... B. rubifaciens R. C. Harris 

1. Thallus with yellow soredia (KC+ orange) in punctiform to diffuse, confluent soralia; 

ascospore wall strongly ornamented; ascospores 16-19 × 9-11.5 µm; south to Polk County........................ B. wheeleri R. C. Harris 

DIRINARIA (Tuck.) Clements 

1. Thallus not sorediate, pustulate or isidiate ............................................................................................................................................. 2 

2. Apothecial disk with white pruina or pruina lacking ............................................................................................................................ 3 

3. Sekikaic acid agg. present; throughout Florida ................................................................................................D. confusa Awasthi 

3. Divaricatic acid present; south to Polk County ..................................................................................... D. confluens (Fr.) Awasthi 

2. Apothecial disk with purple pruina; divaricatic acid; throughout Florida 

......................................................................................................................................................... D. purpurascens (Vainio) Moore 

1. Thallus sorediate, pustulate or isidiate ................................................................................................................................................... 4 

4. Medulla white....................................................................................................................................................................................... 5 

5. Sorediate, pustulate or isidiate-pustulate ....................................................................................................................................... 6 

6. Sorediate, soralia mostly orbicular, eroded to capitate; soredia farinose ................................................................................. 7 

7. Lobes plane, not confluent, irregularly branched; apices truncate; thalli mostly 

smaller than next; divaricatic acid; throughout Florida .................................................. D. picta (Swartz) Clements & Shear 

7. Lobes often convex, confluent; apices flabellate; 

divaricatic acid; throughout Florida ................................................................................................D. applanata (Fée) Awasthi 

6. Pustulate or isidiate-pustulate; divaricatic acid; throughout Florida...................................................D. aegialita (Ach.) Moore 

5. Isidiate; isidia subglobose to cylindrical, not forming pustules or soredia 

divaricatic acid; Citrus and Okeechobee counties.......................................................................... D. papillulifera (Nyl.) Awasthi 

4. Medulla red; divaricatic acid; Levy and Seminole counties....................................................................D. leopoldii (Stein) Awasthi 

HAFELLIA Kalb, Mayrhofer & Scheidegger 

The treatment follows Sheard (Bryologist 95: 79-87. 1992) although I am somewhat uncomfortable about including Buellia bahaiana 

in Hafellia. The three species and one variety known from Florida are included in the key to Buellia above. 

HETERODERMIA Trevisan 

1. Thallus lobes shorter, not strap-like, mostly closely adnate to substrate............................................................................................... 2 

2. Lobes sorediate or isidiate; apothecia rare......................................................................................................................................... 3 

3. Sorediate .......................................................................................................................................................................................... 4 

4. Underside corticate ...................................................................................................................................................................... 5 

5. Salazinic acid and unknown PQ4 present; common, north 

of the Everglades.........................................................................................................H. albicans (Pers.) Swinscow & Krog 

5. Salazinic acid and unknown PQ4 absent; common, south 

to Highlands County ................................................................................................................H. speciosa (Wulfen) Trevisan 

4. Underside not corticate ................................................................................................................................................................ 6 

44

6. Underside white or purplish gray in the center, KOH- ............................................................................................................. 7 

7. Thallus coarse with coarse marginal soredia; rhizines dark................................................................................................. 8 

8. Underside white or weakly purplish toward the center, 

not yellowish toward tips; norstictic acid rarely present; 

south to Lake Okeechobee ..................................................................................H. japonica (Sato) Swinscow & Krog 

8. Underside purplish, yellowish toward tips, ultimate parts 

white; norstictic acid present; Marion County ...........................................................H. casarettiana (Massal.) Trevisan 

7. Thallus delicate with fine soredia in terminal, somewhat labrose, soralia; 

rhizines pale; norstictic acid lacking in N. American material; 

Lake and Marion counties ..............................................................................................H. galactophylla (Tuck.) W. Culb. 

6. Underside orange, KOH+ purple; south to Lake Okeechobee ................................................. H. obscurata (Nyl.) Trevisan 

3. Isidiate; underside orange, KOH+ purple; Leon, Marion and Okaloosa counties ....................................... H. crocea R. C. Harris 

2. Lobes neither sorediate nor isidiate; apothecia present...................................................................................................................... 9 

9. Rhizines simple; norstictic acid absent; Panhandle ......................................................................... H. echinata (Taylor) W. Culb. 

9. Rhizines branched; norstictic acid usually present; reported by Kurokawa 

from Dade County, not seen by me .................................................................................................H. barbifera (Nyl.) K. P. Singh 

1. Thallus lobes narrow, long, strap-like, subfruticose, loosely attached to substrate; 

tips of lobes often sorediate below; with or without salazinic acid; Marion County 

and south, reported by Moore, not seen .......................................................................................................... H. leucomelos (L.) Poelt 

HYPERPHYSCIA Müll. Arg. 


2. Lower surface pale, conspicuous black hypothallus lacking; 

soralia marginal; south to Polk County ..................................................................................H. adglutinata (Flörke) Mayrh. & Poelt 

2. Lower surface dark, conspicuous black hypothallus present; 

soralia laminal; Dade County ..........................................................................................................................H. minor (Fée) Awasthi 

1. Thallus esorediate; Duval and Levy counties, reported by Moore, not seen ......................................H. syncolla (Tuck. ex Nyl.) Kalb 

PHAEOPHYSCIA Moberg 

1. Thallus sorediate; apothecia usually lacking .......................................................................................................................................... 2 

2. Medulla white........................................................................................................................................................................................ 3 

3. Tips of lobes naked .......................................................................................................................................................................... 4 

4. Soralia capitate, borne on raised, lateral lobes; underside dark; Panhandle 

(fide Esslinger, not seen by me).....................................................................................................P. pusilloides (Zahlbr.) Essl. 

4. Soralia marginal; underside pale; Baker County (fide Esslinger, not 

seen by me) ..................................................................................................................................... P. chloantha (Ach.) Moberg 

3. Tips of lobes with erect, colorless hairs; St. Johns County ............................................................. P. cernohorskyi (Nádv.) Essl. 

2. Medulla orange-red; Panhandle fide Esslinger, not seen by me ....................................................... P. rubropulchra (Degel.) Essl. 

1. Thallus not sorediate; apothecia present; Panhandle .................................................................................. P. ciliata (Hoffm.) Moberg 

PHYSCIA (Schreber) Michaux 


2. Lobes white spotted, including lobe ends; zeorin present or absent; ascospores Physcia- 

or Pachysporaria-type .......................................................................................................................................................................... 3 

45

3. Spores Physcia-type; zeorin present ................................................................................................................................................ 4 

4. Lobes narrow, often convex, not confluent, branching irregular; apothecial 

disk not or only weakly pruinose; margin thin, smooth, not raised; 

ascospores small, 15-17 × 6-7.5 µm; Calhoun, Dixie, Gadsden 

and Hamilton counties..............................................................................................................................P. pumilior R. C. Harris 

4. Lobes broader, flatter, often confluent, branching more nearly dichotomous; 

apothecial disk normally strongly pruinose; margin thick, often becoming 

crenulate, usually strongly raised; ascospores larger, 18-25 × 8-12 µm; 

mostly north of Mason-Dixon line..........................................................................................[P. aipolia (Ehrh. ex Hampe) Fürnr.] 

3. Spores Pachysporaria-type, 19-20 × 8-9 µm; zeorin absent or trace; 

Dade County and Florida Keys ................................................................................................................... Physcia sp. Small 7569 

2. Lobes not white spotted or if white spotted then only in the older parts, 

not on lobe ends; zeorin absent; apothecia with rhizines on back; ascospores 

Pachysporaria-type, 20-25 × 8-11 µm; throughout Florida............................................................................ P. neogaea R. C. Harris 


5. Soralia mainly marginal....................................................................................................................................................................... 6 

6. Underside dark or pale, not striate; lower cortex paraplectenchymatous ..................................................................................... 7 

7. Underside pale; soralia marginal, not pustular ............................................................................................................................ 8 

8. Medulla KOH-; zeorin absent; thallus not "frosted", thin and fragile; 

Lake County, "SW Florida" and Louisiana...........................................................................................................P. crispa Nyl. 

8. Medulla KOH+ yellow; zeorin present; thallus "frosted", thicker; 

margins often undulate; Collier and Hillsborough counties ..............................................................P. undulata Moberg 

7. Underside dark; soralia marginal, often pustular; soredia coarse; 

zeorin present; throughout Florida ..................................................................................................P. sorediosa (Vainio) Lynge 

6. Underside pale at tips with dark longitudinal lines; lower cortex more or less 

prosoplectenchymatous; lobe margins pruinose especially at tips; common, 

throughout Florida ..........................................................................................................................................P. atrostriata Moberg 

5. Soralia mainly laminal; underside pale ............................................................................................................................................... 9 

9. Lower cortex prosoplectenchymatous; zeorin absent, unknown terpenoid present; 

northern Florida .................................................................................................................................................P. americana Merr. 

9. Lower cortex paraplectenchymatous; zeorin present; Columbia and Dade counties................................ P. poncinsii Hue 

NOTES 

Physcia poncinsii Hue 

New to North America. FLORIDA. Columbia County: W side of Woodland Avenue, off Co. Rd. 138, 0.5 mi W of U.S. Hwy. 27, 

29°51'N, 82°39'W, on Berchemia scandens, 5 Dec 1994, Harris 36037 (NY); Dade County: Coral Gables, Fairchild Tropical Garden, 

4 Nov 1982, Tucker 25266 (LSU). 

Physcia undulata Moberg 

New to North America. FLORIDA. Collier County: Fakahatchee Strand State Preserve, Royal Palm Hammock, on Ilex, 6 Dec 

1992, Harris 29942; Hillsborough County: Hillsborough River State Park, on Carya, 3 Dec 1992, Harris 29691. LOUISIANA. West 

Feliciana Parish: Locust Grove Cemetery, ca. 4 mi from Bains, on grave stone, 13 Sep 1975, Buck B596 (all NY). 

PYXINE Fr. 

1. Thallus UV-; medulla PD+ orange or PD-............................................................................................................................................... 2 

2. Medulla white or tinted yellow or orange, PD+ orange (testacein agg.) ............................................................................................. 3 

3. Lobes with marginal dactyls (isidiate/pustulate) ................................................................................................................................ 4 

4. Dactyls fragile, almost immediately breaking down into granular soredia; 

46

PD+ dirty orangish; ascospores 4-celled; throughout Florida ....................................................... P. eschweileri (Tuck.) Vainio 

4. Dactyls coarse, not becoming sorediate (may be broken in handling); 

PD+ red orange; ascospores 2-celled; Walton County............................................................................... P. coralligera Malme 

3. Lobes with laminal soralia; ascospores 2-celled; Leon, Marion and 

Walton counties......................................................................................................................... P. eschweileri (Tuck.) Vainio s. lat. 

2. Medulla yellow-orange, PD-; soredia marginal; not yet definitely known from Florida ...............................[P. sorediata (Ach.) Mont.] 

1. Thallus UV+ yellow (lichexanthone); medulla PD- ................................................................................................................................. 5 

5. Soredia or dactyls present................................................................................................................................................................... 6 

6. Medulla white or pale yellow, KOH+ yellow or KOH-...................................................................................................................... 7 

7. Medulla white; Peninsular Florida......................................................................................................... P. cocoes (Swartz) Nyl. 

7. Medulla yellow; Alachua, Dixie and Hamilton counties ............................................................................P. subcinerea Stirton 

6. Medulla orange to pale salmon, KOH+ purple ................................................................................................................................ 8 

8. Lobes with laminal soralia; throughout Florida ............................................................P. caesiopruinosa (Nyl.) Imshaug s. lat. 

8. Lobes with marginal dactyls; south to Seminole County............................................ P. caesiopruinosa (Nyl.) Imshaug s. str. 

5. Soredia and dactyls lacking; medulla yellow; Lake Okeechobee southward....................................... P. berteriana (Fée) Imshaug 

NOTES 

Pyxine caesiopruinosa (Nyl ) Imshaug 

As noted in 1991 there are two taxa included under this name which are readily separable by their asexual reproductive structures 

and terpene patterns. I have now had a chance to examine Nylander's type. Although fragmentary it is clearly the taxon with marginal 

dactyls. It still remains to examine the type of 

P. albovirens (G. Meyer) Aptroot. Aptroot's illustration (not type) of P. albovirens is clearly the laminal soraliate taxon which is also in the 

West Indies but I see no point in taking up albovirens until the correct application of the name is determined. 

RINODINA (Ach.) S. Gray 

1. Thallus white, gray or brownish............................................................................................................................................................... 2 

2. Thallus gray or brownish; apothecial margin often poorly developed; no lichen substances ........................................................... 3 

3. Thallus lacking isidioid granules ..................................................................................................................................................... 4 

4. Apothecia sessile, sometimes constricted below; thallus usually somewhat 

areolate, often poorly developed, not shiny................................................................................................................................ 5 

5. Ascospore lumina not constricted in the middle; apothecia with at least 

remnants of thalline margin...................................................................................................................................................... 6 

6. Ascospore lumina globose to slightly longer than broad; 

ascospores 16-18 × 7-8 µm; south to Polk County..........................................................................R. applanata H. Magn. 

6. Ascospore lumina distinctly longer than broad; ascospores 15-18 × 6-7.5 µm; 

Leon and Volusia counties.................................................................................................................. Rinodina sp. 23268 

5. Ascospore lumina elongated, constricted in the middle, bowling pin- 

or dumbell-shaped; ascospores 17-20 × 7.5-9 µm; apothecia without 

thalline margin; Sarasota County......................................................................................... Buellia placodiomorpha Vainio 

4. Apothecia immersed in continuous, smooth, shiny thallus; ascospores 

with subglobose lumina, 15-19 × 9-10 µm; Dade County............................................................... Rinodina sp. Tucker 25286 

3. Thallus tan, with isidioid granules; ascospores with globose lumina, 

16-19 × 9-10 µm; Polk and Dade counties ......................................................................................... Rinodina sp. Tucker 25528 

2. Thallus and apothecial margin white or pale gray, well developed, 

resembling a Lecanora; ascospores Physcia-type, 18-20 × 9-11 µm; 

atranorin, zeorin; Duval, Lake and Seminole counties.................................................................................R. granuligera H. Magn. 

47

1. Thallus and apothecial margin greenish yellow; ascospores Pachysporaria-type, 

20-27 × 11-13 µm; central Florida ...................................................................................................................R. lepida (Nyl.) Müll. Arg. 

PERTUSARIALES 

PERTUSARIACEAE Körber 

OCHROLECHIA Massal. 

1. Thallus not pustulate-isidiate, smooth to rugose, rugae often vermiform; 

apothecial cortex C-, medulla C+ red (gyrophoric acid with accessory 

lichexanthone and methylhiascic acids); throughout the state...................................................................................O. africana Vainio 

1. Thallus pustulate-isidiate; pustules and thalline rugae very irregular, occasionally 

breaking down into a granular sorediose crust; apothecia mostly without 

cortex; apothecial medulla C+ red, (gyrophoric acid and 5-O-methylhiascic acid); 

Liberty and Marion counties ................................................................................................................................... O. antillarum Brodo 

PERTUSARIA DC. 

1. Ascomata discoid, lecanorate, often heavily pruinose or sorediate above or sterile and 

soraliate; ascospores l-2/ascus when present [if photobiont is Trentepohlia or chemistry 

is not one of those below, these will be treated in key to sterile sorediate crusts]................................................................................ 2 

2. Thallus UV+ yellow (lichexanthone)..................................................................................................................................................... 3 

3A. Lecanoric acid; wart C+ red; often fertile; south to Lake County....................................................................P. pulchella Malme 

3B. Hypothamnolic acid; wart KOH+ lavender, KC+ red, PD-; 

sometimes fertile; Duval, Liberty and Walton counties ......................................................................P. hypothamnolica Dibben 

3C. Picrolichenic acid; wart KOH-, C-, KC+ violet, PD-; 

sterile; Putnam and Wakulla counties ...............................................................................................................P. ventosa Malme 

3D. Baeomycesic acid, unknowns, squamatic acid (tr.); 

wart KOH-, C-, KC-, PD+ yellow; often fertile; 

south to Lee County ........................................................................................................................................ P. floridana Dibben 

3E. Haemathamnolic acid; wart KOH+ yellow becoming brownish, 

C-, KC-, PD+ orange; rarely fertile; throughout the state ..............................................................................P. copiosa Erichsen 

3F. Thamnolic acid; wart KOH+ yellow, C-, KC-, PD+ orange; 

south to Lake County.....................................................................................................................................Pertusaria sp. 25229 

3G. Stictic acid; wart KOH+ yellow, C-, KC-, PD+ pink-orange; 

Marion County .....................................................................................................................................Pertusaria sp. Buck 16889 

2. Thallus UV- or UV+ pinkish [if appropriate chemistry is not found, specimen is either a 

lichexanthone deficient form of the above group or not a Pertusaria] ............................................................................................... 4 

4. Thallus UV-, cortical xanthone lacking; usually fertile ..................................................................................................................... 5 

5A. Lecanoric acid; wart C+ red; south to Orange County........................................................................... P. velata (Turner) Nyl. 

5B. Picrolichenic acid; wart KOH-, C-, KC+ violet, PD-; Duval County.......................................................... P. amara (Ach.) Nyl. 

5C. Thamnolic acid; wart KOH+ yellow, C-, KC-, PD+ orange; 

Volusia County..................................................................................................................................P. trachythallina Erichsen 

5D. Fumarprotocetraric acid; wart KOH-, C-, KC-, PD+ 

orange-red; Leon and Wakulla counties........................................................................................ P. multipunctoides Dibben 

4. Thallus UV+ pinkish (coronatone); stictic acid agg.; wart KOH+ yellow, C-, KC-, 

PD+ pink-orange; sterile; Duval, Highlands, Liberty and Nassau counties ............................................... P. expolita R. C. Harris 

1. Apothecia with small pore-like disk......................................................................................................................................................... 6 

6. Ascospores 8/ascus ............................................................................................................................................................................. 7 

7. Cortex UV+ orange, pink or UV- ...................................................................................................................................................... 8 

8. Cortex UV+ orange (thiophaninic acid)......................................................................................................................................... 9 

48

9. Medulla PD+ orange ............................................................................................................................................................... 10 

10. Stictic acid agg. present; ostioles often yellow and raised; 

ascospores 45-75 × 21-33 µm; throughout the state.............................................................................P. texana Müll. Arg. 

10. Virensic and an unknown (PD-) present; ostioles not yellow and raised; 

ascospores 60-70 × 28-35 µm; Highlands and Okaloosa counties.............................................. P. virensica R. C. Harris 

9. Medulla PD-............................................................................................................................................................................. 11 

11. Containing variolaric acid; ascospores basally biseriate to uniseriate, 

65-90 × 30-40 µm; ostiolar area often depressed and yellow; 

Duval, Leon and Liberty counties.................................................................................................P. epixantha R. C. Harris 

11. Containing unknown substance (perlatolic/confluentic acid group); 

ascospores uniseriate, 55-60 × 25-28 µm; ostiolar area not yellow; 

Duval County .......................................................................................................................................Pertusaria sp. 23902 

8. Cortex UV+ pinkish (coronatone) or UV- .................................................................................................................................... 12 

12. Cortex UV-; medulla PD- (unknown substance); ascospores uniseriate, rough, 

70-95 × 30-40 µm; south to Highlands County ........................................................................................ P. iners R. C. Harris 

12. Cortex UV+ pinkish (coronatone); medulla PD+ pink-orange (stictic acid agg.); 

apothecial warts very irregular; ascospores uniseriate, smooth, 

50-60 × 27-30 µm; Lake County.............................................................................................................Pertusaria sp. 23573 

7. Cortex UV+ yellow (lichexanthone); medulla K-, P- (4-O-methylperlatolic acid); 

ascospores uniseriate, 55-100 × 28-40 µm; south to Marion County.................................................P. paratuberculifera Dibben 

6. Ascospores basically 2 or 4/ascus (1-5) ............................................................................................................................................... 13 


14. Cortex UV+ pinkish or UV-.......................................................................................................................................................... 15 

15. Medulla PD+ pink-orange (stictic acid agg.); hymenium not inspersed................................................................................ 16 

16. Ascospores smooth, 50-100 × 23-43 µm; Duval County (reported by Dibben 

from Lee and Seminole counties also) ............................................................................. P. leucostoma (Bernh.) Massal. 

16. Ascospores rough, 75-130 × 27-43 µm; throughout the state...................................................P. tetrathalamia (Fée) Nyl. 

15. Medulla PD- (traces of unknown); hymenium densely inspersed; 

ascospores smooth, 63-100 × 30-45 µm; throughout the state...................................................... P. sinusmexicani Dibben 

14. Cortex UV+ orange (xanthone X-l); medulla PD+ pink-orange (stictic acid agg.); 

ascospores smooth, 90-105 × 30-35 µm; St. Lucie and Pinellas counties ........................................ Pertusaria sp. Britton 829 


17. Medulla PD+ orange .................................................................................................................................................................... 18 

18. Medulla PD+ pink-orange (stictic acid agg.).......................................................................................................................... 19 

19. Ascospores rough, 55-125 × 28-45 µm; ostiole pale; cortical xanthone 

thiophaninic acid; throughout the state ......................................................................................... P. xanthodes Müll. Arg. 

19. Ascospores smooth, 65-135 × 27-40 µm; ostiole dark; cortical xanthone, 

xanthone X-l; reported by Dibben from Duval and Seminole counties .......................................P. pustulata (Ach.) Duby 

18. Medulla PD+ orange-red (fumarprotocetraric acid); cortex UV- 

(unknown xanthone); ascospores somewhat roughened, 

75-135 × 30-45 µm; Lake and Marion counties .....................................................................................P. subpertusa Brodo 

17. Medulla PD- (variolaric acid); cortex UV- (no xanthone); apothecia mostly 

immersed; ascospores somewhat roughened, 100-120 × 30-40 µm; Marion 

and Wakulla counties...................................................................................................................................P. obruta R. C. Harris 

49

PART 2 

THE 10¢ TOUR OF THE PYRENOLICHENS 

A TWENTY-YEAR PERSPECTIVE 

In 1975 I submitted my thesis which, while dealing primarily with Arthopyrenia s. lat., also included a 

primitive attempt to break away from Zahlbruckner's system, then in use. Over the ensuing 20 years my 

mistakes as regards Arthopyrenia and the more general system have become evident, new ideas and 

information have become available, critical specimens have been examined and a large number of new 

records have accumulated. This anniversary seems an appropriate time to try again. The system proposed is 

not intended in any way to be final but only an interim report. As noted in the introduction the title "tour" is 

given to this part so that I am free to pick and choose what is treated. This is not the proverbial 50¢ tour, only 

the 10¢ version, since I badly miscalculated the amount of errors, changes and specimens to be waded 

through. The most interesting part, the higher categories, is left undone. They naturally fall at the end of such 

a project since I have worked from the bottom (species) upwards. I do not like the current tendency to assign 

new genera, etc., to "incertae sedis". This conveys absolutely no information. It is useless. It is an admission 

of failure, of lack of general knowledge, of laziness, which one is surely forced to admit to on occasion but 

should not become a way of life. Given a choice, I have chosen to err on the side of controversy. This, I 

50

hope, leads to more intensive study and ultimately to a better result. Nothing is set in stone. I do not consider 

some of the groups treated here to be lichens. Not only do I feel there is no consistent association with a 

photobiont, they fail the basic definition that requires some visible morphological evidence of interaction 

between fungus and photobiont. I have omitted for now perithecoid forms belonging to the Gyalectales s. lat. 

and Lecanorales. Also the non-lichen Mycoglaena, Micropeltidaceae, is omitted. The following table shows 

the changes over 20 years in the systematic arrangement of the corticolous pyrenolichens. Due to time 

constraints I have not been able to work through the mostly saxicolous Verrucariaceae nor the perithecioid 

genera of parasites/ parasymbionts, some of which probably are confamilial with lichen-forming taxa. 

ZAHLBRUCKNER (1926) 

PYRENULACEAE 

Anthracothecium 

Arthopyreneiella 

Arthopyrenia 

Belonia 

Blastodesmia 

Clathroporina 

Coccotrema 

Leptorhaphis 

Microthelia 

Monoblastia 

Polyblastiopsis 

Porina 

Pseudopyrenula 

Pyrenula 

Stereochlamys 

Thelopsis 

TRYPETHELIACEAE 

Bottaria 

Laurera 

Melanotheca 

Tomasellia 

Trypethelium 

HARRIS (1974) 

PLEOSPORACEAE 

Arthopyrenia 

Blastodesmia 

Leptorhaphis 

Microthelia 

Mycomicrothelia? 

Mycoglaena? 

Mycoporum 

Polyblastiopsis 

Sporoschizon 

Tomasellia 

PYRENULACEAE 


Bottaria 

Eopyrenula 

Lithothelium 

Melanotheca 

Microglaena s. str. 

Parathelium 

Parmentaria 

Plagiocarpa 

Pleurotheliopsis 

Pyrenastrum 

PARATHELIACEAE 

Campylothelium 

Parathelium 

Plagiotrema 

Pleurotrema 

HARRIS (1995) 

LOCULOASCOMYCETES 

PLEOSPORALES 

NAETROCYMBACEAE 

Jarxia 

Leptorhaphis 

Naetrocymbe 

Tomasellia 

MICROPELTIDACEAE 

Mycoglaena 

MYCOPORACEAE 

Mycoporum 

ARTHOPYRENIACEAE 

Arthopyrenia 

Julella 

Mycomicrothelia 

XANTHOPYRENIACEAE 

Pyrenocollema 

Zwackhiomyces 

ZAHLBRUCKNER (1926) 

Pleurothelium 

ASTROTHELIACEAE 

Astrothelium 

Cryptothelium 

51 

Lithothelium 

Parmentaria 

Pyrenastrum 

STRIGULACEAE

Haplopyrenula 

Micropyrenula 

Microtheliopsis 

Phyllobathelium 

Phylloblastia 

Phylloporina 

Raciborskiella 

Strigula 

Trichothelium 

XANTHOPYRENIACEAE 

Xanthopyrenia 

MYCOPORACEAE 

Asteroporum 

Dermatina 

Mycoporellum 

HARRIS (1975) 

PYRENULACEAE 

Pyrenula 

STRIGULACEAE 

Acrocordia 

Anisomeridium 

Monoblastia 

Pleurotrema 

Pyrenocollema 

Strigula 

TRICHOTHELIACEAE 

Clathroporina 

Porina 

Trichothelium 


Astrothelium 

Campylothelium 

Cryptothelium 

Laurera 

Polymeridium 


Trypethelium 

* Genera in square brackets probably doomed to synonymy. 

VERRUCARIALES 

52 

HARRIS (1995) 

DACAMPIACEAE 

Dacampia 

Eopyrenula 

MELANOMMATALES s. lat. 

THELENELLACEAE 

?Celothelium 

[Chromatochlamys]* 

Thelenella 

STRIGULACEAE 

Phyllobathelium 

Strigula 

PYRENULACEAE 

Acrocordiella 


Clypeopyrenis 

Distopyrenis 

Granulopyrenis 

Lachrymospora 

Lithothelium 

Mazaediothecium 

Mycopyrenula 

Parapyrenis 

Polypyrenula 

Pyrenographa 

Pyrenowilmsia 

Pyrenula 

Pyrgillus 


Architrypethelium 

Astrothelium 

Bathelium 

[Campylothelium] 

[Cryptothelium] 

Exiliseptum 

Laurera 

Polymeridium 


Trypethelium 

HYMENOASCOMYCETES 

TRICHOTHELIALES 

TRICHOTHELIACEAE 

Clathroporina 

Porina 

Segestria 

Trichothelium 

XYLARIALES 

PLEUROTREMATACEAE 

Pleurotrema, etc. (see Barr, 1994) 

I have not had time to deal with this order in a thorough fashion, treating only the rather few Florida 

species. However, my current thought is that the Verrucariales probably should be merged with the

Chaetothyriales and a number of parasitic/parasymbiontic genera currently placed either in Verrucariaceae or 

Mycosphaerellaceae should be transferred to the Microtheliopsidaceae that is more closely allied to 

Chaetothyriales than Verrucariales. 

VERRUCARIACEAE Zenker 

1. Thallus crustose, on rock; ascospores simple or 2-celled ........................................................................ 2 

2. Ascospores simple ................................................................................................................ Verrucaria 

2. Ascospores 2-celled............................................................................................................... Thelidium 

1. Thallus squamulose, on bark, mosses, lichens or rock; ascspores simple, 

transversely septate or muriform.............................................................................................................. 3 

3. Thallus of large to small, brown or tan, ± orbicular or irregular squamules, 

without a raised margin, not sorediate; on bark, soil or rock................................................................ 4 

4. Ascospores simple, 8/ascus; hymenial algae absent; on bark or soil........................ Catapyrenium 

4. Ascospores muriform, 2/ascus; hymenial algae present; on limestone 

or cement ...................................................................................................................... Endocarpon 

3. Thallus of tiny, green gray, orbicular or ± irregular squamules, usually with 

a raised margin or marginal soralia; over bark, mosses or other lichens; 

ascomata very rare; Dixie, Marion and Suwannee counties ................................................................... 

................................................................................................... Normandina pulchella (Borrer) Nyl. 

CATAPYRENIUM Flotow 

1. Growing on calcareous soil; thallus usually quite orbicular, 

not imbricate,brown; Gadsden County.................................................... C. squamulosum (Ach.) Breuß 

1. Growing on bark (occasionally on shaded limestone); 

thallus pale, tan or grayish, imbricate when well developed; 

Leon County .........................................................................................C. tuckermanii (Mont.) Thomson 

NOTES 

I thank Othmar Breuß for determining C. squamulosum. Both species known from Florida belong to the 

segregate genus Dermatocarpella Harada (1993). Based on my limited knowledge of this group, Harada's 

genera seem tenable but the European arbiters have not yet spoken. The combination for tuckermanii has not 

been made however Dermatocarpella squamulosa (Ach.) Harada is available. The genus just barely makes it 

into northern Florida. Harada divides Catapyrenium as follows. 

1. Exciple colorless........................................................................................................................................ 2 

2. Pycnidia of Xanthoria-type; lobes round; attached to substrate by 

rhizohyphae over almost the whole lower surface ...........................................Dermatocarpella Harada 

2. Pycnidia of Staurothele-type; lobes linear, canaliculate below, 

basally attached, almost stipitate; lower surface naked ................................. Neocatapyrenium Harada 

1. Exciple brown to almost black................................................................................................................... 3 

3. Pycnidia unknown; exciple brown; upper surface ± scabrose, dull; 

upper cortex paraplectenchymatous; on soil or bark ................................................Catapyrenium Ach. 

3. Pycnidia of the Staurothele-type; exciple almost black; upper 

53

surface smooth, ± glossy; upper cortex pachydermatous; on 

soil or rock ......................................................................................................... Scleropyrenium Harada 

ENDOCARPON Hedwig 

1. On mortar between bricks (HCl+); thallus brown, relatively undivided; 

ascospores 23-31 × 11-16 µm; Dade County.............................................. Endocarpon sp. Calkins 147 

1. On HCl- rock along river; thallus light gray, becoming lobate; 

ascospores 29-41 × 11-16 µm; Hillsborough County .................................. Endocarpon sp. Buck 22648 

NOTES 

Calkins 147 resembles some forms assigned to E. pusillum Hedwig but the upper cortex is composed of 

relatively thin walled hyphae. Buck 22648 is smaller and more divided than E. pusillum and also differs in thin 

walled cortical cells. There are several Cuban species but I have not seen adequate material. Endocarpon is 

in need of a careful revision in the Americas. 

NORMANDINA Nyl. 

Normandina pulchella (Borrer) Nyl. is probably undercollected because of its small size and is more 

common in Florida than the few specimens would indicate. Applying Occam's Razor I prefer not to recognize 

Lauderlindsaya J. David & D. Hawksw. (= ascomata of Normandina). Three similar 

squamulose/subsquamulose species with ascomata seems to me a simpler explanation than three similar 

species each with three similar parasymbionts. There is no "hard" data such as attempts to recombine 

ascospores with isolated algae or comparative molecular studies of ascospores and vegetative hyphae. 

THELIDIUM Massal. 

1. Thallus epilithic, ± continuous to areolate, greenish; ascomata globose, 

sessile, ca. 0.2 mm diam., wall brown-black; ascospores 2-celled, 

20-23 × (8-)9-11(-13.5) µm; on natural limestone and cement rubble; 

Clay, Collier and Taylor counties..................................................................... Thelidium sp. Buck 22736 

1. Thallus mostly endolithic, greenish?; ascomata immersed, subglobose, 

clypeate; exciple colorless; ascospores 2-celled, 17-20 × 7.5-8.5 µm; 

Monroe County (Key West) .............................................................................Thelidium sp. Thaxter 403 

NOTES 

Buck 22736 may be referable to T. minutulum Körber but comparative material was not available. 

VERRUCARIA Ach. 

1. Ascomata deeply immersed in limestone; black clypeus restricted to a disk 

or cap around ostiole; ascomatal wall colorless........................................................................................ 2 

2. Ascomata numerous, crowded; clypeus a small (ca. 0.2 mm) flat disk; 

thallus endolithic, white; all collections sterile with immature asci; 

periphyses very abundant and conspicuous; Collier, Dade, Monroe 

and Taylor counties ..................................................................................... Verrucaria sp. Britton 861 

54

2. Ascomata scattered; clypeus large (0.5-0.7 mm), flat to hemispherical, 

(depending on erosion of surrounding limestone), often with a distinct 

papilla; thallus endolithic, white; ascospores 17-22(-24) × 9-11 µm; 

Monroe County (Key West) ............................................................................. V. riddleana R. C. Harris 

1. Ascomata sessile or with base immersed in thallus or limestone; black 

clypeus hemispherical; ascomatal wall colorless or dark.......................................................................... 3 

3. Ascospores larger, 15-30 × (7-)9-16 µm; thallus endolithic or 

epilithic; ascomata 0.35-0.6 mm diam. ................................................................................................. 4 

4. Thallus white, endolithic or epilithic ................................................................................................... 5 

5. Exciple dark; clypeus not broadly spreading; ascomata 

0.4-0.5 mm diam.; ostiolar area depressed; ascospores 

21-30 × 11-14 µm; Dade and Monroe counties.................................. Verrucaria sp. Britton 870 

5. Exciple pale; clypeus spreading, 0.5-0.6 mm diam.; 

ascospores 2-28 × 10.5-14.5 µm; Dade and 

Monroe counties................................................................................ Verrucaria sp. Thaxter 397 

4. Thallus green gray or olive green to brown green............................................................................. 6 

6. Ascospores 15-23 × (7-)9-11 µm; thallus olive or brown 

green, areolate; ascomata 0.35-0.4 mm diam.; Taylor 

County ................................................................................................ Verrucaria sp. Buck 24842 

6. Ascospores 23-27 × 11-12.5 µm; thallus green gray, rimose 

or areolate; ascomata ca. 0.4 mm diam.; Taylor and 

Wakulla counties ................................................................................ Verrucaria sp. Buck 18979 

3. Ascospores small, 9-16 × 4-6.5 µm; thallus epilithic, tan to brown; 

ascomata 0.25-0.3 mm diam. ............................................................................................................... 7 

7. Ascospores 9-12 × (4-)5-5.5(-6.5) µm; exciple usually dark; 

Dade, Hillsborough and Monroe counties .............................................. Verrucaria sp. Buck 22655 

7. Ascospores 13-16 × 5.5-6.5 µm; exciple pale; 

Dade County............................................................................................ Verrucaria sp. Britton 741 

NOTES 

There is so little known about subtropical/tropical Verrucaria that I have not ventured to describe any new 

taxa. Only three species have Caribbean types and I have seen material of only one of these. One could 

shoehorn some of the Florida taxa into European species, i.e., Britton 870 into V. muralis Ach. or Thaxter 397 

into V. calkinsiana Servít, but in my present state of ignorance it doesn't seem worth it. This has obviously 

been the practice in the West Indies. 

Verrucaria riddleana R. C. Harris, nom. nov. 

Verrucaria mamillaris Riddle in Britton & Millspaugh, Bahama fl. 522. 1920 (non Hepp in Zollinger, 

1854). Type. BAHAMAS. Watling's Island: vicinity of Cockburn Town, 13 Mar 1917, Britton & 

Millspaugh 6130, not seen. 

New to North America. I have not seen the type collection but two "paratypes" agree with the Florida 

material. It is no surprise to find a species desribed from the Bahamas on Key West. I have no recent 

collections. FLORIDA. Monroe County: Key West, Feb 1898, Thaxter 395, 420, s.n. (NY). 

55

PLEOSPORALES 

NAETROCYMBACEAE Höhnel ex R. C. Harris, fam. nov. 

Naetrocymbeen Höhnel, Sitzungsber. Kaiserl. Akad. Wiss. Math.-Naturwiss. Cl., Abt. 1, 118: 1200. 

1909, nom. inval. [not in Latin or form complying with ICBN]. 

Familia Pleosporalium pseudoparaphysibus peculiaribus areis refractivis ad extremos cellulorum, ascis 

obpyriformibus loculo apicali lato, plerumque in sectione subtriangulari, ascosporis hyalinis vel brunneis, 

plerumque granularibus ornamentatis et microconidiis breviter bacillaribus. 

Type. Naetrocymbe Körber. 

Eriksson (1981) had an extensive description of the type species of Naetrocymbe as N. fumago where he 

illustrated the odd physes with the ends of the cells usually ± refractive and differentially staining. It is also 

described under Arthopyrenia rhyponta. The validation of this family is necessitated due to facing up to the 

fact that Arthopyrenia s. lat. contains two major disparate elements. This is more extensively discussed under 

Arthopyreniaceae. In addition to the short-celled pseudoparaphyses, the group is supported by the obpyriform 

asci with a rather distinctive apical region (lacking a nasse often seen in Arthopyreniaceae), ornamented 

ascospores, short rod-shaped microconidia and a mainly temperate/boreal distribution. Currently I accept four 

genera in the Naetrocymbaceae, Jarxia, Naetrocymbe and Tomasellia with reasonable confidence. 

Leptorhaphis is placed here more or less by default. The physes are not quite right, the asci are more oblong 

although they seem to have a similar apical region and macroconidia are known. As far as I have been able 

to ascertain over the years none of the members of this family are lichen-forming. In my opinion it makes 

about as much sense to include Peziza or Sordaria in checklists of lichens as it does to include members of 

the Naetrocymbaceae. 


1. Ascospores colorless (sometimes weakly tinted when postmature)......................................................... 2 

2. Ascomata with a single chamber .......................................................................................................... 3 

3. Ascospores soleiform, 1-septate to submuriform ........................................................ Naetrocymbe 

3. Ascospores narrowly fusiform arcuate, 1-5-septate..................................................... Leptorhaphis 

2. Ascomata compound with several/many centra under a single 

clypeus; ascospores 4-celled, 14-20 × 4-5.5 µm; Duval 

County ............................................................................. Tomasellia americana (Willey) R. C. Harris 

1. Ascospores brown, 1-septate (rarely 3-septate when postmature); 

tropical/subtropical (but poorly known)..............................................................................................Jarxia 

JARXIA D. Hawksw. 

Stud. Mycol. 31: 93. 1989. 

Jarxia is distinguished from Mycomicrothelia in coarser physes with refractive areas at the cell ends, 

obpyriform asci and differently proportioned, thinner walled ascospores. The genus is probably overlooked 

due to its small ascomata and likely to be more common than the few collections indicate. 

1. Ascospores (23-)25-28 × (5.5-)6.5-7(-8.5) µm; on Ilex; 

56

Duval County...................................................................................J. thelenula (Müll. Arg.) D. Hawksw. 

1. Ascospores 17-21 × 5-7 µm; on Ilex; Monroe and 

Duval counties....................................................................................................... J. ilicicola R. C. Harris 

Jarxia ilicicola R. C. Harris, sp. nov. 

NOTES 

Jarxiae thelenulae similis sed ascosporis minoribus, 17-21 × 5-7 µm. 

Type. FLORIDA. [Duval County: Jacksonville?] on Ilex cassine, Calkins, Fla. Lich. 152 (NY, holotype). 

Thallus indicated by grayish blotch on bark, with blackish antagonism zone. Ascomata scattered, 

hemispherical to subglobose, superficial to ca. ½ immersed, 0.15-0.2 mm diam.; wall not melanized at base. 

Asci obpyriform, ca. 60 × 20-30 µm, with irregularly arranged ascospores. Ascospores yellow-brown, 2-celled, 

rarely? 4-celled in age, 17-21 × 5-7 µm, finely granular ornamented. Microconidia not found. 

Additional specimens. FLORIDA. Duval County: Ft. George [Island], on Ilex, Calkins, N. Amer. Lich. 280; 

Monroe County: hammock, S end of Big Pine Key, 11 Dec 1921, Small et al. 10159 (both NY). 

Jarxia thelenula (Müll. Arg.) D. Hawksw. 

New to North America. Reported by Hawksworth from Cuba and the Dominican Republic. 

FLORIDA. Duval County: Jacksonville, [on Ilex], Calkins, N. Amer. Lich. 255 (NY). TEXAS. Webb County: 

University of Wisconsin monkey colony, 30 mi N of Loredo on U.S. Hwy. 83, 2 Dec 1972, Iltis 27420 (MIN). 

LEPTORHAPHIS Körber 

The genus is almost entirely northern with only a single scrappy collection from Florida. It seems close to 

L. amygdali (Massal.) Zwackh and L. parameca (Massal.) Körber (both on Prunus) in having a broad basal 

fringe, IKI+ bluish hymenial gel and similar ascospore shape. However, the ascospores are broader, 37-47 × 

4-5.5 µm, and it does not occur on Prunus. Better material is need before it is formally described. For the 

northern species the excellent revision by Aguirre-Hudson (Bull. Br. Mus. Nat. Hist. (Bot.) 21: 85-192. 1991) 

should be consulted. Otherwise a pyrenocarpous specimen with filiform ascospores from Florida is probably a 

Celothelium. 

FLORIDA. Monroe County: hammock, S end of Big Pine Key, 11 Dec 1921, Small et al. 10154-D (NY). 

NAETROCYMBE Körber 

Santessoniolichen Tomaselli & Cifferi, Arch. Bot. (Forli) 28: 5. 1952. ≡ Santessoniomyces Cifferi & 

Tomaselli, Ist. Bot. Reale Univ. Reale Lab. Crittog. Pavia Atti, ser. 5, 10: 29,57. 1953. Type. S. 

punctiformis ("Fr.") sensu Jatta [sic]. 

?Sporoschizon Riedl, Sydowia 14: 334. 1960. Type. Sporoschizon petrakianum Riedl (= Naetrocymbe 

punctiformis s. lat.). 

My previous intent was to recognize N. rhyponta as a monotypic Arthopyrenia separate from the 

punctiformis-group (which I have occasionally annotated as Santessoniolichen). The "retypification" of 

Arthopyrenia has changed that. Also, I have recently concluded that the differences are minor. The most 

obvious being the dark brown subiculum usually present in N. rhyponta. Foucard (1992) indicated from his 

study of ample material that it probably is of no taxonomic significance. Eriksson (1987) recognized the type 

of Naetrocymbe, N. fumago, as belonging in Arthopyrenia s. lat. and Foucard (1992) further considered it to be 

synonymous with Arthopyrenia rhyponta. Although I have not seen type material of N. fumago, I concur with 

57

their decisions and the resurrection of the genus and family is the result. Sporoschizon is synonymized with 

question as I have not seen the holotype of Sporoschizon petrakianum but an isotype (Krypt. Vind. 4454, S) 

does not show any ascospore fragmentation which was the basis of Riedl's genus and is synonymous with 

Naetrocymbe punctiformis s. lat. I am making only such combinations in Naetrocymbe as affect North 

America (although it is my hope they will not be included in the checklist). I hope this is the last time I have to 

deal with this peripheral non-lichenized taxon. It will be omitted from a final treatment of Florida lichens. 

KEY TO NAETROCYMBE IN NORTH AMERICA 

1. Ascospores 2-4-celled............................................................................................................................... 2 

2. Ascopsores 2-celled (rarely 4-celled when postmature) ....................................................................... 3 

3. Physes persistent .............................................................................................................................. 4 

4. Ascospores smaller, 26 µm or less long ....................................................................................... 5 

5. Ascomata superficial, flattened, often surrounded by dark hyphal ring..................................... 6 

6. Ascospores 19-25 × 5.5-7.5 µm; on Fraxinus; Maine, 

Minnesota and Vermont ...................................................... [N. fraxini (Massal.) R. C. Harris] 

6. Ascospores 15-20 × 4.5-5.5(-6.5) µm; on a variety of 

smooth barks; northern U.S. (southern limit in East is 

Long Island)................................................................... [N. punctiformis (Pers.) R. C. Harris] 

5. Ascomata at least initially immersed, ± globose ....................................................................... 7 

7. Ascospores 22-26 × 6-7.5 µm; known only from Florida, Duval, 

Gilchrist, Levy and Seminole counties ...............................N. cedrina (Zahlbr.) R. C. Harris 

7. Ascospores 16-22 × 4-5 µm; asci slender, 55-70 × 12-16 µm; 

rare, Seminole County and Alabama ........................N. atractospora (Zahlbr.) R. C. Harris 

4. Ascospores large, 25-32 × 6-8 µm; ascomata superficial, flattened; 

Maine, Europe .................................................................... [N. megalospora (Lönnr.) R. C. Harris] 

3. Physes dissolving with only fragments remaining; ascospores 

14.5-17 × 4-5 µm; Monroe County (Keys) ..................................N. quassiaecola (Fée) R. C. Harris 

2. Ascospores 4-celled.............................................................................................................................. 8 

8. Ascomata resting in extensive dark brown subiculum, 

0.2-0.3 mm diam.; ascospores 17-20 × 5.5-7 µm; 

northern U.S.? (no "good" American material seen) ........................ [N. rhyponta (Ach.) R. C. Harris] 

8. Ascomata not surrounded by a subiculum, 0.1-0.2 

mm diam.; ascospores 17-21 × 5-6 µm; subtropical, 

Duval County....................................................................N. atomarioides (Müll. Arg.) R. C. Harris 

1. Ascospores 6-celled, 20-23 × 5-5.5 µm; physes persistent; 

Duval County............................................................................................. Naetrocymbe sp. Calkins 157 

NOTES 

Naetrocymbe atomarioides (Müll. Arg.) R. C. Harris, comb. nov. 

58

Arthopyrenia atomarioides Müll. Arg., Bot. Jahrb. Syst. 6: 406. 1885. Type. CUBA. Wright, Lich. 

Cub. II. 629 (G, holotype). 

Verrucaria subpunctiformis Nyl., Lich. ins. guin. 51. 1889. Arthopyrenia floridana Zahlbr., Cat. lich. 

univ. 1: 332. 1922, nom. nov. Type. FLORIDA. [Duval County:] Ft. George, 1887, Calkins 

(FH, MICH, NY, US, isotypes?). 

Naetrocymbe atractospora (Zahlbr.) R. C. Harris, comb. nov. 

Arthopyrenia atractospora Zahlbr., Ann. Mycol. 33: 33. 1935. Type. FLORIDA. [Seminole County:] 

Sanford, on Quercus, Jul 1928, Rapp137 (W 2451, holotype). 

Additional specimen. ALABAMA. Baldwin County: Yupon Point, 1 Mar 1925, Evans 210 (MICH). 

Naetrocymbe cedrina (Zahlbr.) R. C. Harris, stat. et comb. nov. 

Arthopyrenia sanfordensis var. cedrina Zahlbr., Ann. Mycol. 33: 33. 1935. Type. FLORIDA. 

[Seminole County] Sanford, on Taxodium, Oct 1922, Rapp 135 (W 2459, holotype). 

The type and the Gilchrist County collection are on Taxodium, the Duval County and Alabama collections 

on Myrica. Microscopically rather similar to N. fraxini, this species is distinct in the smaller more globose 

ascomata, substrate and distribution. 

Naetrocymbe fraxini (Massal.) R. C. Harris, comb. nov. 

Arthopyrenia fraxini Massal., Ric. auton. lich. crost.167, f. 333. 1852. Type. "Ad truncos Fraxini 

ubique in regno Veneto. Massal.", Massal. Lich. Ital. 298A (BM, lectotype, Coppins, 1988, not 

seen by me). 

Naetrocymbe megalospora (Lönnr.) R. C. Harris, comb. nov. 

Arthopyrenia megalospora Lönnr., Flora 41: 634. 1858. Type. SWEDEN. Fries, Lich. Suec. 244 

(FH-Tuck 4067, isolectotype, Harris, 1975). 

Foucard (Graphis Scripta 4: 49-60. 1992) has placed this in synonymy with N. fraxini on no better 

grounds than those by which I recognize it, speculation. The ascospores seem significantly larger to me. I 

prefer to keep the question open and therfore make the combination in Naetrocymbe. 

Naetrocymbe massalongiana (Hepp) R. C. Harris, comb. nov. 

Sagedia massalongiana Hepp, Flecht. Eur. 444. 1857. Type. SWITZERLAND. Zurich and Liestal, 

Hepp (NY). 

This species has been previously synonymized with N. saxicola (see below) but the asci and ascospores 

are larger than in the probable type of N. saxicola (Massalongo, Lich. Ital. 348). The asci are 75-85 × 20-25 

µm vs. ca. 50 × 20 µm and are less distinctly pyriform than in N. saxicola. The ascospores are 18-25 × 6-7(-8) 

µm vs. 16-20 × 4-5 µm. A broader study may find these differences trivial and recognition here is mainly 

intended as a stimulus for such a study. There are apparently two colections distributed in Hepp 444 and I 

have no way of making an intellegent lectotype. Hepp's description is also apparently mixed. The illustration 

of the ascus and the measurements of ascus and ascospores agree with the Naetrocymbe. The 4-celled 

ascospores would seem to belong to Trichothelium lineare which is mixed in one of the NY packets of Hepp 

444. Its previous synonymization with N. saxicola would seem to indicate maintaining the name for the 

Naetrocymbe. It is not known from North America. 

Naetrocymbe punctiformis (Pers.) R. C. Harris, comb. nov. 

Verrucaria punctiformis Pers., Ann. Bot. (Usteri) 11: 19. 1794. Arthopyrenia punctiformis auct. 

non Massal. Type. "Verrucaria punctiformis", sine loca in herb. Persoon (L, lectotype, Riedl, 

1977, not seen). 

Arthopyrenia padi Rabenh., Lich. Eur., fasc. 14, 390. 1858. Type. "an Prunus Padus bei 

Hermsdorf in Bilaer Grunde in der sächse Schweiz." (MICH, isotype). 

One of the benefits of the removal of this species to a genus other than Arthopyrenia is that it allows 

Persoon's sacred, although commonly misapplied, epithet to be taken up again legitimately. (Because of the 

existence of Arthopyrenia punctiformis Massal. which excluded Persoon's name, I have been using A. padi 

59

Rabenh.) Foucard (Graphis Scripta 4: 53. 1992) designated a neotype. However, Riedl (Sydowia 29: 115- 

138. 1977) described Persoon's material at great length and provided a photograph of its label. Until there is 

some statement by Prof. Riedl that this Persoon material is lost or destroyed, I consider the neotype to be 

superseded. 

Naetrocymbe quassiaecola (Fée) R. C. Harris, comb. nov. 

Verrucaria epidermidis var. γ. quassiaecola Fée, Essai crypt. écorc. 84. 1825. Arthopyrenia 

quassiaecola (Fée) Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 30(3): 26. 1888. Type. 

AMERICA. On Quassia excelsa (G, lectotype; G, isolectotype). 

The ascospores of Florida material are slightly larger than those of the type. There is a single specimen 

from Puerto Rico. The physes dissolve leaving only fragments embedded in gel as in Arthopyrenia salicis 

Massal. s. lat. I include quassiaecola in Naetrocymbe but suspect that the dissolving hamathecium indicates 

that it really belongs elsewhere but have no suggestions at this time. 

Naetrocymbe rhyponta (Ach.) R. C. Harris, comb. nov. 

Verrucaria rhyponta Ach., Kongl. Vetensk. Akad. Nya Handl. 30: 150. 1809. Arthopyrenia 

rhyponta (Ach.) Massal., Ric. auton. lich. crost. 166. 1852. Type. SWEDEN. On Fraxinus (H-Ach 

772, the three left hand pieces, lectotype, selected here). 

Naetrocymbe saxicola (Massal.) R. C. Harris, comb. nov. 

Arthopyrenia saxicola Massal., Symm. lich. nov. 107. 1855. Pyrenocollema saxicola (Massal.) 

Coppins in Coppins et. al., Lichenologist 24: 368. 1992. Type. ITALY. Massalongo, Lich. Ital. 348 

(NY, isotype?). 

Although not in North America, I treated this species briefly in my thesis, placing it in Pyrenocollema, and 

now wish to correct my misunderstanding of its taxonomic position. I was uneasy about its disposition and left 

it to Coppins to do the dirty work of making the transfer. In restudying some of the problems in Pyrenocollema 

for this volume, I realized that the asci were pyriform and that the hamathecium and microconidia were of the 

Naetrocymbe-type. I do not know what the biology of this species is. It may be a parasite as it grows with 

cyanobacteria on limestone with no obvious thallus formation. (This association with cyanobacteria suggested 

Pyrenocollema.) I have rummaged briefly for an appropriate parasite/parasymbiont genus but none is 

immediately obvious. Careful morphological studies combined with the different? biology of N. saxicola and N. 

massalongiana (segregated from N. saxicola above) may well find that generic recognition is indicated. 

TOMASELLIA Massal. 

The genus is treated in a narrower sense than previously. Most of the species have been transferred to 

Mycoporum (see Mycoporaceae for discussion). There is only a single species in Florida, T. americana 

(Willey) R. C. Harris, based on a single Calkins collection from Duval County which is, thus, unfortunately a 

little suspect. 

MYCOPORACEAE Zahlbr. 

This group has turned out to be one of my major early failures. In my thesis work I was so impressed by 

the compound nature of the ascomata that I did not look closely enough at the characters of the ascus and 

hamathecium. In re-evaluating this group I have come to the conclusion that most of the taxa referred to 

Mycoporellum Müll. Arg. and Mycoporopsis Müll. Arg. are not congeneric with Tomasellia Massal. and, at least 

tentatively, not confamilial. In 1975 I declined to recognize 2-celled versus 4-celled ascospores as a generic 

difference. I have since found taxa with ascospores intermediate between transversely septate and muriform. 

Further, on the basis of ascus and hamathecium I believe that two species with simple, uniloculate ascomata 

should be placed in this group also. Assuming all transversely sepatate species are placed in one genus, 

traditional mycological taxonomy would recognize four genera, Mycoporum (compound ascomata, muriform 

ascospores), Mycoporellum (compound ascomata, transversely septate ascospores), unnamed (simple 

60

ascomata, muriform ascospores) and unnamed (simple ascomata, transversely septate ascospores). 

However, I do not feel that these distinctions are supported by any additional characters and have no 

predictive value. Therefore, I recognize only the single genus Mycoporum Flotow ex Nyl. as have Eriksson 

(1981) and Barr (1987), but with a broader circumscription. 

Both Mycoporum and Tomasellia have compound ascomata with several to many locules, each with its 

own ostiole (except for the ostioles the ascomata superficially resemble those of Arthonia and have often been 

confused). The asci of Mycoporum are obpyriform to obclavate with the wall in the upper third very much 

thickened and with virtually no stipe or with a broad, short stipe. Those of Tomasellia arthonioides (Massal.) 

Massal. (the generitype) are ± cylindrical with relatively even wall thickenings and a short, broad stipe. The 

hamathecium in Mycoporum consists of very irregular, slightly gelatinizing and somewhat disarticulating 

physes which are almost entirely obscured by irregular oil droplets. (If you can make out the physes clearly, it 

probably isn't a Mycoporum). That of Tomasellia consists of narrower, more regular physes which may also 

gelatinize somewhat, having a refractive body at one or both ends of the cells as in Arthopyrenia rhyponta 

(Ach.) Massal. and lacks oil droplets. The ascospores of Mycoporum are thin walled, often collapsing, and 

tend to turn yellow brown with a conspicuous granular ornamentation when old. Those of Tomasellia are 

comparatively thicker walled and do not readily become brownish and ornamented. The microconidia in both 

genera are short, ca. 5 µm, rod-like or oblong. Possible oblong macroconidia are found in M. buckii assigned 

to Mycoporum with some reservation. The biological status of Mycoporum is unclear. I was incorrect in 

suggesting Trentepohlia as a photobiont. There is no morphologically differentiated lichen thallus although in 

section thick walled superficial hyphae are present which become brown and more convoluted near the 

ascomata. Some species seem not to be lichen-forming. Others usually seem to have a cover of "nonstandard" 

algae (i.e., not similar to known photobionts). It may be that Mycoporum is weakly or facultatively 

lichen-forming. If so, it represents another independent development of the lichen lifestyle. Tomasellia seems 

definitely not lichen-forming. 

Purvis et al. (1992) include two species in Mycoporum, M. hippocastani (DC.) Coppins and M. quercus 

(Massal.) Müll. Arg., both of which seem to me to belong elsewhere. They have ± spherical asci, ± 

paraplectenchymatous hamathecial tissue and the ascoma is not of the "clypeal" type but a stroma pushing 

through the upper layers of bark, possibly non-ostiolate. My current thought is that these two species should 

be referred to Cyrtidula Minks. Vainio (1921) emended the genus retaining only one of the original species 

and that can serve as a lectotypification with C. pteleodes (Ach.) Minks (= hippocastani). The combination C. 

quercus (Massal.) Minks exists. The other combination seems not to have been made. Cyrtidula 

hippocastani (DC.) R. C. Harris, comb. nov. (Verrucaria hippocastani DC., Fl. franç. 6: 172. 1815.). I have 

no records of either from North America. 

Of the species referred to Tomasellia in addition to the type, T. arthonioides, I would retain T. americana 

(Minks ex Willey) R. C. Harris, T. diffusa (Leighton) Lahm., T. gelatinosa (Chev.) Zahlbr. and T. macularis 

(Minks ex Willey) R. C. Harris, comb. nov., (Cyrtidula macularis Minks ex Willey, Enum. lich. New Bedford 

33. 1892. Type. MASSACHUSETTS. New Bedford, Willey, (MICH, isosyntype), treated in 1975 but combination 

not validly published). Tomasellia americana and T. macularis are maintained provisionally pending reexamination 

of type material. All four differ somewhat from the type species in more obclavate asci and are 

kept in Tomasellia on the basis of compound ascoma and hamathecial type. The genus Athrismidium Trevisan 

(syn. Beckhausia Hampe ex Körber, nom. inval.), typified with A. gelatinosa, is available for the others if one 

wishes to restrict Tomasellia to its type species. Tomasellia diffusa was included in Melanthecopsis Dodge. 

However, the type of this genus, M. follmanii Dodge, is a Pyrenula, close to or synonymous with P. cubana 

(Müll. Arg.) R. C. Harris. Melanothecopsis sinensis (Krempelh.) Dodge is a coelomycete with brown 2-celled 

conidiospores. The other two species, M. coarctella (Stirton) Dodge and M. subpuncta (Nyl.) Dodge are 

unknown to me. Tomasellia americana is treated in the Naetrocymbaceae, T. arthonioides and T. diffusa do 

not occur in North America, T. gelatinosa is known from Maine and Newfoundland and T. macularis from 

Massachusetts. 

The following are possible generic synonyms of Mycoporum but cannot be placed definitively at this time 

as I have not seen type material, Mycoporopsis Müll. Arg., lectotype (Riedl, 1962), Mycoporum sorenocarpum 

Knight, and Nothostroma Clements, holotype, Mycoporopsis roseola Müll. Arg. 

MYCOPORUM Flotow ex Nyl., nom. cons. 

61

Mém. Soc. Sci. Nat. Cherbourg 3: 186. 1855. Type. M. elabens Flotow ex Nyl. 

Bottaria Massal., Misc. lichenol. 43. 1856. Holotype. B. composita Massal. 

Mycoporellum Müll. Arg., Rev. Mycol. (Toulouse) 6: 14. 1884. Lectotype (Riedl, 1962). Mycoporum 

sparsellum Nyl. 

Syngenosorus Trevisan, Conspect. Verruc. 15. 1860. Lectotype (chosen here). Melanotheca arthoniella Nyl. 

Chlorodothis Clements, Genera of fungi 50. 1909. Holotype. Mycoporellum lahmii Müll. Arg. (= Mycoporum 

lacteum) 

Sciodothis Clements, Genera of Fungi 50. 1909. Holotype. Mycoporopsis leucoplaca Müll. Arg. (= 

Mycoporum lacteum) 

1. Ascoma compound, with several to many chambers, each with a pore, 

each containing many asci ........................................................................................................................ 2 

2. Ascospores transversely septate .......................................................................................................... 3 

3. Ascospores mostly 2-celled, some becoming 4-celled when old...................................................... 4 

4. Ascospores fusiform, relatively slender, 2.5-4 times longer than broad........................................ 5 

5. Ascospores larger, over 15 × 5 µm; ascoma without well developed 

brown basal layer; macroconidia not known ............................................................................. 6 

6. Ascospores 17-20(-22) × 5-6(-7) µm; very common, throughout 

Florida .................................................................... M. eschweileri (Müll. Arg.) R. C. Harris 

6. Ascospores 20-27 × 6.5-8 µm; not uncommon, throughout 

Florida ...................................................................................M. lacteum (Ach.) R. C. Harris 

5. Ascospores small, 10-13 × 3-3.5 µm; ascoma with a distinct brown 

basal layer; macroconidia oblong, ca. 5 × 2 µm; on Taxodium or 

old wood; Collier and Duval counties ...................................................... M. buckii R. C. Harris 

4. Ascospores relatively short and "fat", 1.5-2.5 times longer than broad, 

17-22(-25) × 7.5-10 µm; upper cell markedly shorter than lower; 

common, throughout Florida ............................................................................M. sparsellum Nyl. 

3. Ascospores mostly 4-celled, only immature ascospores 2-celled..................................................... 7 

7. Ascospores 17-22 × 5.5-7 µm; physes obscured by oil; 

rare, Citrus and Escambia counties ..................................M. californicum (Zahlbr.) R. C. Harris 

7. Ascospores 15-17 × 4-5.5 µm; physes not obscured by 

oil; rare, Duval County............................................. Tomasellia americana (Willey) R. C. Harris 

2. Ascospores muriform ............................................................................................................................ 8 

8. Ascospores over 30 × 12 µm ........................................................................................................... 9 

9. Ascospores 30-38(-43) × 12-17(-18) µm, with 8-9 rows of locelli; 

Franklin, Gilchrist and Okaloosa counties.......................M. compositum (Massal.) R. C. Harris 

9. Ascospores (36-)39-48(-55) × (14-)16-21(-24) µm, with (9-)10(-11) 

rows of locelli; Duval County ........................................................ M. pycnocarpoides Müll. Arg. 

8. Ascospores 23-28(-30) × 9-12 µm, with 8 rows of locelli; not uncommon, 

northern Florida ........................................................................................M. acervatum R. C. Harris 

62

1. Ascoma simple, with only a single chamber ........................................................................................... 10 

10. Ascospores mostly 2-celled, 4-celled and brownish in old age, 

28-40 × 8-12 µm; Lake County.......................................................M. antecellens (Nyl.) R. C. Harris 

10. Ascospores muriform, 22-37 × 7-9.5 µm, with 8-9 rows of locelli; 

not seen to become brownish; northern Florida.................................... M. uniloculatum R. C. Harris 

Mycoporum acervatum R. C. Harris, sp. nov. 

NOTES 

Similis M. compositi et M. pycnocarpoidis sed ascosporis minoribus, 24-29 × 9-11 µm. 

Type. FLORIDA. Clay County: Gold Head Branch State Park, vicinity of Loblolly Trail, mixed hardwoods along 

stream, 29 Nov 1992, Harris 29260 (NY, holotype). 

Thallus indicated by whitish blotch on bark, in type patchily covered by small clumps of algae. Ascomata 

orbicular to ± irregular in outline, flattened, 0.5-1.0 mm diam. Asci obpyriform, 60-75 × 30-40 µm, with 

ascospores irregularly arranged. Ascospores oblong-elliptical with rounded ends, slightly constricted at 

median septum, colorless (no brown ascospores observed), muriform, with 8 rows of 2-6 cells, 23-28(-30) × 9- 

12 µm. Pycnidia with microconidia, oblong, 3 × 1.5 µm. 

Mycoporum acervatum is distinguished from its near relatives M. compositum, M. elabens and M. 

pycnocarpoides only on ascospore size. Since it has no really distinctive character, I have given it an epithet 

which could apply to most of the genus, meaning "in small heaps" in reference to the appearance of the 

compound ascoma. I am not really sure how to interpret the ascospore variation in this group. We have no 

theoretical basis. Why should basically similar ascospores vary in size? Is it the result of random mutation 

fixed in isolated populations? Probably not in this case as compositum and pycnocarpoides co-occur widely in 

the Neotropics and Africa and both occur with acervatum in Florida. Is there some microhabitat or dispersal 

advantage related to size? Is it just minor genetic "noise", e.g., big vs. little humans? If the latter, then all four 

should probably be lumped into a single species. I am not even sure how one would go about addressing this 

problem. This question aside, M. acervata does differ in a minor way from compositum and pycnocarpoides in 

that I have never observed brown ascospores which are common at maturity in those species. Mycoporum 

acervatum is closest in ascospore size to the European M. elabens but the latter has larger, hence fewer 

locelli per row. 

Additional specimens (a few from old notes not re-examined 1995). ALABAMA. Conecuh County: Brooklyn, 

1860-61, Beaumont 218 (FH-Tuck 3735). FLORIDA. Bradford County: Montgomery Conference Center, 4 mi 

NW of Keystone Heights, on branches of Quercus margaretta, 11 Feb 1993, Griffin 390 (FLAS); Duval County: 

Jacksonville, Calkins, NAL 78 (FH), NAL 163 (NY); Leon County: Apalachicola National Forest, Leon Sinks 

Geological Area, oak-pine woods, on Quercus, 2 Dec 1988, Harris 23272 (NY); Marion County: Ocala, 31 Feb 

1891, Underwood 1843 (NY); Walton County: Lakewood, Lakewood Park, highest point in Florida, 345 ft, oakhardwood 

forest, on Quercus, 30 Nov 1988, Harris 23077 (NY). SOUTH CAROLINA. without location, Ravenel 

(FH-Tuck 3735). 

Mycoporum antecellens (Nyl.) R. C. Harris, comb. nov. 

Verrucaria antecellens Nyl., Flora 49: 86. 1866. Arthopyrenia antecellens (Nyl.) Müll. Arg., Flora 

53: 485. 1870. Mycoporopsis antecellens (Nyl.) Riedl, Sydowia 15: 268. 1961[1962], nom. 

inval., (basionym not cited). Type. ENGLAND. Sussex: Tilgate Forest, Larblastier 77 (H-Nyl 

767, holotype). 

Pyrenula zwackhii Hepp, Fl. Eur. 954. 1867. Type. GERMANY. Near Heidelberg, Zwackh (FH, FH- 

Tuck 4066, NY, isotypes). 

63

Verrucaria subantecellens Nyl., Flora 59: 364. 1876, nom. nud. Arthopyrenia subantecellens 

Müll. Arg., Bot. Jahrb. Syst. 6: 403. 1885. Type. CUBA. Wright, Verr. Cub. 2 (MSC, holotype 

fragment; H-Nyl 769, isotype). 

Arthopyrenia griseola Müll. Arg., Rev. Mycol. (Toulouse) 10: 182. 1888. Type. PARAGUAY. 

Paraguarí, Balansa (G, holotype). 

Presumably on the basis of the ascospores that soon become 4-celled, brownish and granular 

ornamented, Riedl removed this species from Arthopyrenia to Mycoporopsis even though the ascomata are 

uniloculate. Placement in Mycoporum ( incl. Mycoporopsis sensu Riedl) is further supported by the large 

ascospores, large asci, thick walled in the upper part and the characters of the hamathecium. Purvis et al. 

(1992) spelled the epithet "antecellans". All the specimens in Nylander's herbarium have the "-ens" spelling 

and my small Latin dictionary gives "antecello, -ere" the participle of which should end in "-ens". The British are 

noted scholars of Latin and perhaps there is some linguistic point I have missed. However, until I am 

corrected, I will spell the epithet "antecellens". 

Mycoporum buckii R. C. Harris, sp. nov. 

Similis M. sparselli sed ascomatis planioribus, strato brunneo sub hymeniis, ascosporis minoribus, 9.5-12 

× 2.5-3 µm et macroconidiis oblongiis, ca. 5 × 2 µm. 

Type. FLORIDA. Duval County: Flood plain forest E of San Pablo River (Intracoastal Waterway), W of 17th St. 

N., dead portion of trunk of oak, 19 Jun 1975, Buck B175 (NY, holotype; MICH, isotype? not seen). 

Thallus a whitish blotch on substrate. Ascomata black, superficial, compound, multiostiolate, with many 

chambers, orbicular to elliptical (elongated with the grain of the wood), 0.5-1.0 mm diam, or to ca. 0.5 × 1.0 

mm, with a brown basal layer. Asci obclavate to ± fusiform, 40-55 × 10-14 µm, with ascospores weakly 

biseriately to irregularly arranged. Ascospores fusiform, colorless, 2-celled, 10-13 × 3-3.5 µm. Microconidia 

not found. Pycnidia with macroconidia, oblong, colorless, ca. 5 × 2 µm. 

Mycoporum buckii is anomalous in several ways, the narrow asci which are not basally swollen, the well 

developed brown layer at the base of the ascoma and the presence of what can be interpreted as 

macroconidia. However, I would not feel comfortable creating a new genus and believe it is best 

accommodated in Mycoporum pending more thorough investigation. It is easily recognized by the small 

ascospores. Some species of Strigula have ascospores this small but have small, uniloculate ascomata. In 

addition to the cited collections it has been collected in Belize by Mason Hale. 

Additional specimens. FLORIDA. Collier County: Fakahatchee Strand State Preserve, 1.2 mi from N 

entrance on W. J. James Memorial Scenic Drive, open Taxodium swamp, on trunk of Taxodium, 6 Dec 1992, 

Buck 22817 (NY). DOMINICAN REPUBLIC. Prov. Monte Cristi: SE of El Morro de Monte Cristi, ca. 20 m, low dry 

forest near sea, 9 Jan 1987, Harris 19603 (JBSD, NY). 

Mycoporum californicum (Zahlbr.) R. C. Harris, comb. nov. 

Mycoporellum californicum Zahlbr., Ann. Mycol. 10: 363. 1912. Mycoporopsis californica (Zahlbr.) 

Riedl, Sydowia 16: 216. 1962[1963]. Tomasellia californica (Zahlbr.) R. C. Harris in Tucker & 

Jordan, Wasmann J. Biol. 36: 87. 1979, comb. inval. (33.2). Type. CALIFORNIA. Santa Monica 

Mtns., Hasse 1003 (FH, isotype). 

This species is separated from M. eschweileri, that has similar sized ascospores, by the early 

development of 4-celled ascospores, and from Tomasellia americana by larger ascospores and hamathecium 

obscured by oil droplets. Its distribution is mainly western. The easternmost record previously was from 

Texas. However, a number of ± western species are turning up in northern Florida. 

64

Mycoporum compositum (Massal.) R. C. Harris 

in Egan, Bryologist 90: 164. 1987. Bottaria composita Massal., Misc. lichenol. 43. 1856. Type. ad 

cort. Chinae [comm. Hampe] (VER, holotype). 

Mycoporum pycnocarpum Nyl., Flora 41: 38. 1858 (lapsus pyrenocarpa). Dermatina pyrenocarpa 

(Nyl.) Zahlbr., Cat. lich. univers. 1: 550. 1922. Type. MEXICO. Sierra de Cristóbal, 1855, Fr. 

Müller (H-Nyl 4223, holotype; MICH, isotype?). 

Arthonia tumidula Leighton, J. Linn. Soc., Bot. 9: 200. 1867. Arthothelium tumidulum (Leighton) 

Zahlbr., Cat. lich. univers. 2: 137. 1922. Type. [CANADA.] St. Mary's, Lake Superior, Richardson 

158 (BM, lectotype, selected by Makhija & Patwardhan, 1995; BM, isolectotype (neither seen); 

NY, isolectotype). 

Mycoporum integrum Vainio, Mycologia 21: 40. 1929. Type. PUERTO RICO. On bark near Aibonito, 

3 Jan 1916, Fink 1800 (FH(2), MICH-Fink 8633, NY, "isolectotypes". (I have not examined 

material in Vainio's herbarium but his published ascospore size agrees with M. compositum. 

Other duplicates (MICH-Fink 8953, NY) have the larger ascospores of M. pycnocarpoides. The 

official designation of a lectotype can wait pending examination of material at Turku.) 

Mycoporum pycnocarpum var. ohiense Nyl., Mém. Soc. Sci. Nat. Cherbourg 5:135. 1867, nom. 

nud. Mycoporum ohiense Nyl. ex Fink, Lich. Fl. U.S. 67. 1935. Dermatina ohiensis (Fink) 

Zahlbr., Cat. lich. univ. 10: 108. 1938. Lectotype (Harris, 1973). OHIO. Lea (ex hb. Hooker) (H- 

Nyl 4234). 

Tomasellia cubana Müll. Arg. 

Bot. Jahrb. Syst. 6: 397. 1885. Type. CUBA. Wright, Verr. Cub. 126 (NY, isotype). 

This name is a synonym of Arthopyrenia consanguinea Müll. Arg. 

Mycoporum difforme Minks, Mycoporellum difforme (Minks) Fink 

Described from Massachusetts but I have not been able to locate type. From the description by Riedl 

(1961) it is a synonym of M. lacteum and should be removed from the checklist. 

Mycoporum eschweileri (Müll. Arg.) R. C. Harris, comb. nov. 

Mycoporellum eschweileri Müll. Arg., Flora 71: 526. 1888, Tomasellia eschweileri (Müll. Arg.) R. 

C. Harris in Tucker & Harris, Bryologist 83: 19. 1980. Type. BRAZIL. Bahia (G, holotype). 

Tomasellia esenbeckiana (Fée) Müll. Arg. = Arthopyrenia esenbeckiana 

Tomasellia exumana Riddle 

in Britton & Millspaugh, Bahama Fl. 528. 1920. Type. BAHAMAS. Great Exuma: on bark near 

Georgetown, 22-28 Feb 1905, Britton & Millspaugh 3135 (NY, isotype). 

This is a synonym of Anisomeridium anisolobum (Müll. Arg.) Aptroot. 

Mycoporum lacteum (Ach.) R. C. Harris, comb. nov. 

Verrucaria stigmatella var. lactea Ach., Lichenogr. universalis 227. 1810. Verrucaria lactea (Ach.) 

Eschw. in Martius, Fl. bras. enum. pl. 1: 126. 1833. Mycoporellum lacteum (Ach.) Zahlbr., Cat. 

lich. univers. 1: 554. 1922. Tomasellia lactea (Ach.) R. C. Harris in Tucker & Harris, Bryologist 

83: 19. 1980. Type. Amer. meridion., in Copaifera, [Swartz], (H-Ach 779A, holotype; BM-Ach, 

UPS, isotypes). 

Mycoporopsis leucoplaca Müll. Arg., Flora 69: 316. 1886. Mycoporellum leucoplacum (Müll. Arg.) 

Zahlbr., Nat. Pflanzenfam. 1(1*): 78. 1903. Type. BRAZIL. Minaes Geraes, Glaziou (FH, FH- 

Tayl 118, US, isotypes). 

Mycoporellum lahmii Müll. Arg., Flora 70: 78. 1887. Type. SOUTH AFRICA. Transvaal: near 

Lydenburg, Wilms, comm. Lahm 61 (G, holotype). 

?Mycoporum difforme Minks ex Willey, Enum. lich. New Bedford 32. 1892. Type not seen. 

Mycoporopsis exigua Zahlbr., Sitzungsber. K. Akad. Wiss. Wien. Math.-Naturw. Cl. 111(1): 360, 

pl. 2, f. 24-26. 1902. Type. BRAZIL. Near Theresiopolis, Höhnel, Lich. Bras. 159 (W, holotype). 

Mycoporellum hassei Zahlbr. in Hasse, Bryologist 15: 46. 1912. Type. CALIFORNIA. Catalina 

65

Island: near Avalon, Hasse 1321 (W, holotype; FH, isotype). 

Mycoporellum deserticola Fink in Hedrick, Mycologia 22:248. 1930. Mycoporopsis deserticola 

(Fink) Riedl, Sydowia 16: 218. 1962[1963]. Type. PUERTO RICO. Yauco, 1 Jan 1916, Fink 

1688 (MICH, holotype). 

Tomasellia leucostoma Müll. Arg. 

See Arthopyrenia confluens R. C. Harris 

Mycoporum mycoporoides (Müll. Arg.) R. C. Harris, comb. nov. 

Arthopyrenia mycoporoides Müll. Arg., Rev. Mycol. (Toulouse) 10: 4. 1888. Type. URUGUAY. 

Montevideo, 1887, Arechavaleta (G, holotype). 

This species has uniloculate ascomata and in hamathecial and ascal characters seems very close to M. 

antecellens. In view of the epithet Müller had some inklng of its generic affinities. It differs in having 6-celled 

ascospores. 

Mycoporum pycnocarpoides Müll. Arg. 

Bot. Jahrb. Syst. 20: 290. 1895. Type. TANZANIA? Usambara, im Lutindi-Hochwald, Holst 3373 (G, 

holotype). 

Mycoporopsis rappii (Zahlbr.) Riedl = Melaspilea Nyl. s. lat. (see Arthopyrenia). 

Mycoporum sparsellum Nyl. 

Ann. Sci. Nat. Bot. sér. 5, 7: 343. 1867. Mycoporellum sparsellum (Nyl.) Müll. Arg., Rev. Mycol. 

(Toulouse) 4: 14. 1884. Tomasellia sparsella (Nyl.) R. C. Harris in Tucker & Jordan, Wasmann 

J. Biol. 36: 87. 1979 comb. inval (33.2). Type. COLOMBIA. Monte del Morro, Lindig (H-Nyl 4292, 

lectotype, selected here). 

Mycoporum uniloculatum R. C. Harris, sp. nov. 

Quoad ascos et ascosporas similis M. acervatae sed ascomatis uniloculatis, quoad ascomatas similis M. 

antecellentis sed ascosporis muriformibus. 

Type. FLORIDA. Liberty County: Apalachicola National Forest, pine-oak scrub S of Forest Serv. Rd. 113, 0.8 mi 

W of County Road 12, on small branches of Quercus, 6 May 1990, Harris 25258 (NY, holotype). 

Thallus indicated by pale olive brown blotch apparently due to a supeficial layer of algae in clumps that do 

not seem to be surrounded by hyphae. Ascomata orbicular, flattened, 0.1-0.2 mm diam., with a single 

hymenium, with a weak clypeal ring. Physes with short, irregular cells, mostly obscured by oil droplets. Asci 

obpyriform, ca. 60-75 × 30-35 µm, with irregularly arranged ascospores. Ascospores fusiform, constricted at 

median septum, colorless, muriform, with (6-)8 rows of 1-4 locelli, 22-32 × 7-9.5 µm. Pycnidia with short rodlike 

microconidia, 3-5 × 1 µm. 

As noted in the introduction this species is included in Mycoporum on the basis of features of ascus and 

hamathecium. In ascospore size M. uniloculata is close to M. acervata but has a narrower, more pointed, 

fusiform shape and fewer cells in a row. The general shape and size of the ascospores is close to M. 

antecellens except that they are muriform. All the collections are on oak twigs or small branches from pineoak 

scrub habitat. It is probably common but overlooked. 

Additional specimens. FLORIDA. Clay County: Gold Head Branch State Park, along Florida Trail from 

cabins to clay pit in SW corner of park, pine-oak scrub, 29 Nov 1992, Harris 29241; Marion County: Ocala 

National Forest, Huges Island, slightly moister oak woods surrounded by dry sand pine scrub, 22 Apr 1967, 

Harris 2007-A; Okaloosa County: 1.2 mi W of Fla. Hwy. 85 on Antioch Road (Co. Rd. 4), ca. 4 mi S of 

Crestview, oak-pine scrub, 5 May 1990, Buck 17917 (all NY). 

66

XANTHOPYRENIACEAE Zahlbr. 

It has been suggested that this family should be synonymized with the Arthopyreniaceae. I prefer to keep 

it separate on the basis of physes which are more intricately and densely branched and interconnected, 

differently shaped asci, differently shaped ascospores and elliptical (not rod-shaped) microconidia. The 

differences do not seem major but I think that the relationships of the Xanthopyreniaceae probably lie 

elsewhere in some "non-lichenized" family. Also since I have invested considerable energy in making the 

Arthopyreniaceae homogeneous, I am reluctant to "mess it up". The inclusion of Zwackhiomyces is basically 

arbitrary. There are few useful characters avaiable to place these simple/highly reduced taxa. Essentially 

there are no characters which would exclude Zwackhiomyces from the Xanthopyreniaceae but equally there 

are no unique shared characters to support inclusion. Collemopsidium Nyl. and Didymellopsis (Saccardo) 

Clements & Shear are also homeless and if the positioning of Zwackhiomyces in the Xanthopyreniaceae 

works out, perhaps, they too could be dumped here. 

1. Associated with cyanobacteria; pigment in ascomatal wall amorphous, located 

in the hyphal cell wall; ascospores thin-walled, readily collapsing, not becoming 

brownish and granular-roughened .....................................................................................Pyrenocollema 

1. Parasitic or parasymbiontic on lichens with green algal photobiont; pigment in 

ascomatal wall fine granular, located between hyphae; ascospores with thicker 

wall, not readily collapsing (until overmature), becoming brownish and granular 

ornamented ..................................................................................................................... [Zwackhiomyces] 

PYRENOCOLLEMA Reinke 

In my thesis I pointed out the problems involved in defining species in Pyrenocollema. They have not 

been resolved and even seem to have worsened. The major unresolved questions are: does the 

host/photobiont matter (and therefore does the rock substrate matter, i.e., preference of the cyanobacteria) or 

should the taxonomy be based entirely on fungal characters? 

1. Fresh water aquatics, semi-aquatics or on rock in moist places...............................................................2 

2. Photobiont blue-green in color; on calcareous rock .............................................................................. 3 

3. Ascomata sessile to semi-immersed, ascospores 17-23 × 8-11 µm; 

eastern United States ..................................................[P. prospersellum (Nyl.) R. C. Harris] 

3. Ascomata immersed, with a broad clypeus; ascospores 18-23 × 8-9.5 µm; 

Bermuda............................................................................................ [P. farlowii (Riddle) R. C. Harris 

2. Photobiont yellow brown to brown due to colored sheath (Gleocapsa-like) 

or thallus dark brown ............................................................................................................................ 4 

4. Growing on limestone; ascspores 16-18(-20) × 5-7 µm; Monroe County 

(Key West), Puerto Rico.............................................................P. atlanticum (Vainio) R. C. Harris 

4. Growing on a variety of rock, usually ± aquatic; ascospores 

(18-)20-27 × 8-11 µm; temperate North America, Europe .................................................................. 

.....................................................................................[P. tichothecioides (Arnold) R. C. Harris s. lat.] 

1. Marine, in tidal zone ....................................................................... [P. halodytes (Nyl.) R. C. Harris s. lat.] 

[In my thesis I took a very broad view of this species. It is divided in Purvis et al. (1992) as follows. 

I am not sure at this time how this view applies in North America. 

A. On calcareous rocks or shells; clypeus absent or not spreading; 

ascomata not cracked around ostiole ................................................................................................B 

67

B. Ascomata sessile and flattened, or immersed and globose, 0.2-0.25 mm 

diam.; on calcareous rocks or shells................................................P. halodytes (Nyl.) R. C. Harris 

B. Ascomata immersed or protruding, 1-1.5 mm diam.; on chalk or 

limestone................................................................................. P. sublitorale (Leighton) R. C. Harris 

A. On siliceous rock; clypeus laterally spreading; ascomata cracked 

around ostiole; ascospores 15-20 × 6-9 µm.................................... A. orustense (Erichsen) Fletcher] 

NOTES 

Pyrenocollema atlanticum (Vainio) R. C. Harris, comb. nov. 

Collemopsidium atlanticum Vainio, Mycologia 21: 34. 1929. Type. PUERTO RICO. Yauco, 28 Dec 

1915, Fink 1402 (MICH, isotype). 

The material of Collemopsidium iocarpum (Nyl.) Nyl., the type species of the genus, was still out on loan 

when I was in Helsinki so that I am unable to say if Collemopsidium Nyl. (1881) and Pyrenocollema Reinke 

(1895) are synonymous. If they should prove to be, although Collemopsidium has priority, I assume 

Pyrenocollema would be conserved since it is thoroughly embedded in both North American and European 

literature. The isotype of C. atlanticum consists mostly of a species of Lichenaceae. There are, however, a 

few patches of pale bluish "thallus" supporting the conclusion drawn from microscopic characters to include 

the Florida and recent Puerto Rican specimens in this species. Pyrenocollema atlanticum is quite similar in 

aspect to P. caesium but is distinguished by the smaller ascospores. 

New to North America. FLORIDA. Monroe County: Key West, Feb 1898, Thaxter 412. PUERTO RICO. 

Reserva Forestal Guánica, along Hwy. 333 at Km 4.8, coastal scrub forest, 26 May 1989, Buck 17135 (both 

NY). 

Pyrenocollema caesium (Nyl.) R. C. Harris 

Sagedia nylanderi Hepp, Flecht. Eur. 440. 1857. Thelidium nylanderi (Hepp) Lönnroth, Öfvers. 

Förh. Kongl. Svenska Vetensk.-Akad. 15: 284. 1859. Arthopyrenia nylanderi (Hepp) Riedl, 

Linzer Biol. Beitr. 22: 154. 1990. Type. SWITZERLAND. "An Alpenfindlingen, am Ufar bei Zürich", 

Hepp. (NY, US, isotypes). 

Pyrenocollema ceuthocarpoides (Müll. Arg.) R. C. Harris, comb. nov. 

Arthopyrenia ceuthocarpoides Müll. Arg., Flora 67: 468. 1884. Type. BRAZIL. São Paulo: "Ad saxa 

gneissiaca rivuli Iguape prope Apiahy.", Puiggari (G, holotype). 

Pyrenocollema epigloeum (Nyl.) R. C. Harris, comb. nov. 

Verrucaria epigloea Nyl. Flora 69: 464. 1886. Arthopyrenia epigloea (Nyl.) Zahlbr., Cat. lich. univ. 

1: 279. 1921. ≡ Pyrenocollema tremelloides Reinke, Jahrb. Wiss. Bot. 28: 461. 1886.Type. 

"Supra Nostoc saxa dolomitica incolens inundata fluminis Narenta in Herzegovinia (Lojka)", 

Lojka Lich. univ. 249 (MICH, NY, isotypes). 

This is the type of the genus but I far as I know the oldest epithet has not been combined in 

Pyrenocollema previously. 

Pyrenocollema farlowii (Riddle) R. C. Harris, comb. nov. 

Thelidium farlowii Riddle, Bull. Torrey Bot. Club 43: 147. 1916. Type. BERMUDA. 1881, Farlow 

(MICH, NY, isotypes?). 

Pyrenocollema prospersellum (Nyl.) R. C. Harris 

Sagedia nylanderi β. hudsoniana Hepp, Flecht. Eur. 945. 1867. Thelidium nylanderi var. 

hudsoniana (Hepp) Zahlbr., Cat. lich. univ. 1: 135. 1921. Type. [NEW JERSEY.] "Auf 

Serpentinblöcken am rechten Ufer des Hudson, in der Nähe von Hoboken bei Newyork", 

1861, Uhl (NY, US, isotypes). 

68

Pyrenocollema tichothecioides (Arn.) R. C. Harris s. lat. 

Since my thesis I have examined the type material of this species, the type of Xanthopyrenia Bachm., 

and it is clearly congeneric with Pyrenocollema. This is where most of the obvious problems occur. Michigan 

material cited in my thesis has an ascospore size nearer that of P. strontianense (Swinscow) R. C. Harris but 

are on calcareous rock. Material reported from Louisiana on sandstone is not available but more recent 

collections seem to be better placed in or near Lichenothelia D. Hawksw. A recent collection on sandstone 

from Mississippi is closer to P. strontianense but apparently differs in asomatal characters. Specimens from 

the West seem to be closer to P. tichothecioides. Until a large amount of material is amassed and studied at 

one time, these problems cannot be resolved. 

Pyrenocollema zonatum (Müll. Arg.) R. C. Harris, comb. nov. 

Arthopyrenia zonata Müll. Arg., Flora 71: 549. 1888. Type. BRAZIL. "Saxicola ad flumen Iguape 

prope Iporanga in Brasilia meridionali", Puiggari 3025, 3027 (G, syntypes; BM, M, isosyntypes). 

ZWACKHIOMYCES Grube & Hafellner 

The species described here as Z. arenicola has the ascomatal pigmentation, physis type, ascospore 

type and lifestyle of Zwackhiomyces. Not having seen verified material of a real Zwackhiomyces, I am not 

sure about the ascus type. The apex is not in agreement with Grube's illustrations and the ascus is broadest 

at or below the middle. Also the ascospores have a diffuse gel sheath. I would not like to think another genus 

is needed but the possibility exists. 

1. Ascospores 28-37 × 10-13 µm; on unidentifiable areolate thallus or without 

obvious host; southern Georgia ......................................................................... [Z. arenicola R. C. Harris] 

1. Ascospores 24-27 × 7.5-8.5 µm; without obvious host but green algae present; 

Louisiana ..............................................................................................[Zwackhiomyces sp. Harris 11406] 

Zwackhiomyces arenicola R. C. Harris, sp. nov. 

NOTES 

Ascomatis in areolis lichenis ignoti vel super saxum arenaceum nudum et ascosporis 29-37 × 9-13 µm. 

Type. GEORGIA. Johnson County: 2 mi N of Adrian on Georgia Highway 15, 32°34'N, 82°35'W, ca. 80 m, small 

Altamaha Grit sandstone outcrop on slope, 18 Mar 1995, Harris 36282 (NY, holotype) 

Ascomata initially immersed in and then erupting through small, white, unidentifiable areoles on moist 

sandstone or growing sessile and naked directly on moist sandstone with a layer of green algae, subglobose, 

ca. 0.2-0.35 mm diam.; when growing immersed, ascomata often surrounded by a layer of paler brown 

hyphae, when growing exposed, ascomata black and glossy; ascomatal wall pale below when immersed, 

entirely dark when exposed. Asci elliptical to pyriform, ca. 110-135 × 30-45 µm, with eight irregularly arranged 

spores. Ascospores 28-37 × 10-13 µm, 2-celled, becoming 4-celled when overmature, ovoid, constricted at 

septum, becoming brownish and granular ornamented with age, with a watery gel sheath; upper cell slightly 

shorter and broader than the lower; pycnidia not found. 

All the known collections are from Altamaha Grit which is proving to be a habitat for many interesting 

lichens. The species is so inconspicuous that its absence from similar outcrops elsewhere in the South may 

be a collection (or lack of) phenomenon. A very similar species differing only in smaller ascospores occurs on 

sandstone in Louisiana. Assuming Z. arenicola is correctly assigned to genus, it is distinctive in the larger 

ascospores. Other species of Zwackhiomyces are reported by Grube (Nova Hedwigia 51: 283-360. 1990) 

from Clauzadea, Lecanora, Mycobilimbia and Porpidia. I am unable to determine the host for Z. arenicola, 

partly I believe, since the habitat is sufficiently wet that the lichen does not form a typical thallus. 

69

Additional speciemns. GEORGIA. Coffee County: Broxton Rocks Ecological Preserve, Ricketson Tract, ca. 

10 mi NE of Broxton, ca. 3 mi S of Ocmulgee River, 31°44'N, 82°44'W, ca. 70 m, extensive, dissected 

sandstone outcrop [Altamaha Grit], 4 Feb 1995, Buck 27461, 27495; Johnson County: as type, Buck 27570; 

Washington County: ca. 3 mi N of Harrison along County Road 206 (Peacock Road), near headwaters of 

Rocky Creek, 32°53'N, 82°44'W, ca. 140 m, Altamaha Grit sandstone flats, 18 Mar 1995, Buck 27543 (all NY). 

DACAMPIACEAE Körber 

EOPYRENULA R. C. Harris 

When I described this genus I placed it in the Pyrenulaceae, largely on the basis of IKI+ blue hymenial 

gel, long rod-shaped microconidia and brown ascospores. It was given the generic name with youthful 

naiveté, speculating that the genus might be ancestral to Pyrenula. Aptroot (1991) placed Eopyrenula in the 

Pleomassariaceae. This was accepted by Barr (Mycotaxon 49: 129-142. 1993). Eriksson (Lichenologist 25: 

310. 1993) was dubious about this placement. Personally I find it absurd. The type species of Pleomassaria 

and Splanchnonema have massive clavate asci with thick tips with indistinct, rounded ocular chambers ( with 

an apical nasse? in Pleomassaria siparia (Berk. & Br.) Sacc.), very coarse physes and large ascospores with 

thick gelatinous sheaths. Eopyrenula has smaller cylindrical asci with thin walls evenly thickened in water 

mounts, with a round or elongate ocular chamber in KOH, physes ca. ½ the width of those in 

Pleomassariaceae and medium sized ascospores without a gelatinous sheath. The only character I can see 

linking Eopyrenula to the Pleomassariaceae is the production of brown, septate macroconidia. Casting 

around for a better home for Eopyrenula, I ran across notes made for another purpose on Byssothecium 

Fuckel and Dacampia Massal., both placed in the Dacampiaceae. Eopyrenula has distinctive ascospores with 

the end cells paler than the middle cells. This seems to be due to a two layered spore wall, the inner pale, the 

outer dark. The outer layer apparently forms from the center outwards not covering the end cells which are 

therefore pale. This also seems to be the case in Byssothecium and possibly Dacampia, both of which have 

pale end cells. Both genera lack a gelatinous ascospore sheath as in Eopyrenula. The asci are ± cylindrical 

with ocular chamber similar to Eopyrenula. The pseudoparaphyses are slender as in Eopyrenula. For these 

reasons I suggest placing Eopyrenula in the Dacampiaceae. Barr described the ascomatal wall of 

Dacampiaceae as three layered with the outer pseudoparenchymatous. The ascomatal wall of Eopyrenula is 

of textura intricata. However, to my eye, the wall of Byssothecium circinans Fuckel also consists of textura 

intricata. Some Dacampiaceae have a pore in the ascospore septa which is not found in Eopyrenula. Also 

Eopyrenula-type macroconidia do not seem to be known from the Dacampiaceae. Even with these 

anomalies, Dacampiaceae seem to me a much better fit than Pleomassariaceae or Pyrenulaceae. 

Eopyrenula is well lichenized with Trentepohlia and would seem to represent another independent line of 

lichenization. In the Dacampiaceae Dacampia and Pyrenidium Nyl. are lichen parasites on a wide variety of 

hosts. 

There are three species of Eopyrenula known to me from North America (none of which occur in Florida), 

E. intermedia Coppins ex Aptroot in eastern North America, E. parvispora R. C. Harris & Aptroot in the Pacific 

Northwest and an apparently undescribed species on spruce from Minnesota. The potentially undescribed 

taxon may be the European E. leucoplaca (Wallr.) R. C. Harris as it has 2-celled macroconidia but the 

substrate is unique and I have almost no material of E. leucoplaca for comparison at this time. 

Type. Arthopyrenia Massal. 

ARTHOPYRENIACEAE W. Watson 

70

The family is defined here more narrowly than generally used in the last decade. The mechanics of this 

are discussed under Arthopyrenia and see also Naetrocymbaceae. Jarxia, Leptorhaphis and Naetrocymbe 

(Arthopyrenia rhyponta-punctiformis group) are removed to the Naetrocymbaceae. The restricted family is 

held together by clypeate ascomata, ascomatal wall KOH+ dark green [in the three species (incl. A. lapponina) 

studied by Foucard (1992); I have not assessed this character], pseudoparaphyses (cellular 

pseudoparaphyses), asci in a basal layer, asci fissitunicate, cylindrical to clavate with an apical nasse, 

ascospores colorless or brown, often granular ornamented, septum or initial septum submedian, 2-celled to 

muriform, pycnidia with rod shaped to filiform microconidia, occasionally associated with Trentepohlia, growing 

on bark, less commonly old wood and highest diversity in the tropics. The physes are often long and slender 

(easily distinguished from the sparse, coarse physes of the Naetrocymbaceae) and easily confused with 

paraphysoids (trabecular pseudoparaphyses) suggestive of the Melanommatales but they clearly taper from 

broad anastomosing bases. The cylindrical to clavate, stipitate asci (markedly obovate and nearly estipitate in 

Naetrocymbaceae) to some extent also suggest Melanommatales but the apical nasse would seem be 

anomalous in that order. Placement in the Pleosporales is also supported by the basal (rather than peripheral) 

origin of the asci. It does not seem unreasonable to include Julella in Arthopyrenia but more study is needed 

as there are no really clear intermediates and there are unresolved generic problems within Julella. I do not 

consider the vast majority of the family to be lichenized but a few, e.g., Arthopyrenia lyrata, seem consistently 

associated with Trentepohlia and perhaps continued treatment by lichenists is not inappropriate. 

1. Ascospores colorless ................................................................................................................................ 2 

2. Ascospores 2-4-celled.......................................................................................................Arthopyrenia 

2. Ascospores submuriform to muriform..........................................................................................Julella 

1. Ascospores brown, 2-celled to submuriform, usually granular ornamented ..................Mycomicrothelia 

ARTHOPYRENIA Massal. 

Ric. auton. lic. crost. 165. 1852. Lectotype (Fink, 1910). Arthopyrenia analepta (Ach.) Massal. 

≡ Leiophloea (Ach.) S. Gray, Nat. arr. Brit. pl. 1: 495. 1821. Verrucaria*Leiophloea Ach., Methodus 

Suppl. 24. 1803. Lectotype (Riedl, 1962). L. analepta (Ach.) S. Gray 

≡ Pyrenyllium Clements, Genera fungi 41, 173. 1909. Holotype. P. analeptum (Ach.) Clements 

Cifferiolichen Tomaselli in Tomaselli & Ciferri, Arch. Bot. (Forlì) 28: 4. 1952. ≡ Mycocifferia Tomaselli in 

Ciferri & Tomaselli, Ist. Bot. Reale Univ. Reale Lab. Crittog. Pavia Atti, ser. 5, 10: 28, 56. 1953. Holotype. 

Arthopyrenia lapponina Anzi (= A. analepta). 

In my unpublished thesis I made three major mistakes regarding Arthopyrenia, 1) Not splitting out the 

taxa here referred to Naetrocymbe (I came close, see Harris, 1975: 39), 2) Interpreting Lichen analeptus Ach. 

as illegitimate, 3) Accepting Riedl's lectotypification (If it is maintained, the nomenclatural changes consequent 

to the division of Arthopyrenia are distinctly unpleasant. Cifferiolichen is the next available name for the A. 

lapponina group.). As regards the first, I placed too much emphasis on microconidia. Arthopyrenia fraxini 

Massal. has the longer rod-shaped microconidia typical of A. analepta and I thought it represented a link 

between the two genera. It does not. All the other characters are typical for Naetrocymbe. The microconidia 

are anomalous. (With 20 years of retrospect the ascus and hamathecium characters seem extremely strong 

and necessitate the recognition of two families as is done in this work.) This division of the genus reopens the 

can of worms involved in typifying Arthopyrenia. Lichen analeptus is not illegitimate even though Acharius cited 

Verrucaria olivacea Pers. in synonymy since the name Lichen olivaceus (Melanelia olivacea (L.) Essl.) had 

already been used in the genus Lichen in the Lichenographiae sueciae prodromus. This makes possible the 

acceptance of Fink's typification of the genus. In 1975 I accepted Rield's choice of A. rhyponta (Ach.) Massal. 

as a temporary expedient, thinking that I would change it as soon as possible and believing that no one else 

would seriously endorse such an atypical species as the generitype. I erred grievously on both counts. In 

1994 I was finally able to study the relevant Acharian material in Helsinki. When I discovered that a specimen 

of the taxon currently known as A. lapponina was the obvious lectotype for A. analepta, the way was cleared to 

71

preserve Arthopyrenia in the broadest possible sense. Although Fink's typification may have been 

mechanical, it is not required that it be rejected if it is the most logical, preserving current usage. [If this is 

unacceptable to some, then consider this a proposal to overturn Riedl's lectotype and to reselect A. analepta.] 

The problem remains to decide whether Acharius was describing a new species or creating a new name for 

Verrucaria olivacea Pers. There is no direct indication in his work itself. In the interests of stability, I have 

chosen to consider Lichen analeptus Ach. as a new species. If this is totally unpalatable, then, since 

Arthopyrenia in this sense would require conservation no matter what (see below), it could be lectotypified with 

A. cerasi (Schrader) Massal. which would have the same effect. There is an appropriate Schrader collection 

in H-Ach of A. cerasi in the accepted sense. Either way the bulk of the taxa in temperate, subtropical and 

tropical regions are retained in Arthopyrenia. (In North America seven taxa are transferrred to Naetrocymbe 

but only N. punctiformis is represented by more than a few records so that the real effect is very minimal.) 

Arthopyrenia cinchonae is one of the most commonly collected pansubtropical/pantropical species, likewise A. 

analepta (syn. A. lapponina) in temperate regions and A. plumbaria in the Pacific Northwest, etc. In boreal 

regions Naetrocymbe is perhaps more important and names will have to change in the British and 

Scandinavian mycotas unless it is decided to ignore the differences and retain a broad view of Arthopyrenia. 

(Arthopyrenia seems a possible test case for the meaning of "Names in Common Usage". Will a small area 

with an essentially depauperate (but "well known") mycota sieze precedence over the rest of the world to 

retain Arthopyrenia in the sense of Naetrocymbe?) This typification allows the family name Arthopyreniaceae 

to be retained for three large well known genera, Arthopyrenia, Julella and Mycomicrothelia, that have been 

assigned to this family for some years already. This is also an "economical" solution as it does not require 

many new names, especially at the familial and generic levels (except for the validation of the previously 

proposed Naetrocymbaceae). A side benefit of realizing Lichen analeptus is not illegitimate and obligately 

typified with Verrucaria olivacea Pers. [= Porina s. lat., V. olivacea is not to be typified by the type of the 

misapplied name cited by Persoon (Art. 33 Note 2) as Hawksworth (1985: 133) proposed] is that Leiophloea 

(Ach.) S. Gray (1821) and Pyrenillium Clements (1909) do not have priority over Pseudosagedia (Müll. Arg.) 

M. Choisy (1949), recently resurrected by Hafellner and Kalb. However, since Riedl (1962) lectotypified 

Leiophloea with Lichen analeptus, Arthopyrenia will have to be conserved over Leiophloea or A. cerasi chosen 

as conserved type. 

In my thesis I included a small group of species in Arthopyrenia with some hesitation at the end of the 

genus as the "Arthopyrenia bifera group". The ascospores were distinctive in having ring-like wall thickenings 

forming partial subdivisions of the cells (and in postmature ascospores often ultimately a complete septum). 

Aptroot refers to this type as "biocellate", Vainio as "porospore". Over the succeeding years I have come to 

the conclusion that this is not significant but is in fact a more extreme expression of the general tendency for 

the ascospore cells to be medially constricted in Arthopyrenia. This is most conspicuous in A. 

cinereopruinosa, A. lyrata and A. plumbaria but shows up to some extent even in such "typical" species as A. 

analepta and A. cinchonae. Within the Arthopyreniaceae one might wish to give this ascospore type generic 

recognition but there is no correlation with any other feature and the other characters agree with those of 

Arthopyrenia, so I prefer to maintain a single genus. The same ascospore type occurs also in 

Mycomicrothelia suggesting that it may be a familial tendency. 

KEY TO ARTHOPYRENIA IN NORTH AMERICA 

1. Ascospores 2-celled (rarely 4-celled when postmature) ........................................................................... 2 

2. Ascospore cells not pinched in nor wall with internal ring-like thickenings............................................ 3 

3. Ascospores narrowly ovate or narrowly elliptical, length/width ratio 

2.5:1 or greater................................................................................................................................. 4 

4. Ascospores mostly more than 20 × 7 µm..................................................................................... 5 

5. Ascomata superficial, complanate to hemispherical, with or without 

72

a basal fringe; ascospores 20-30 × 7-9.5(-11) µm; microconidia 

4-5 × 0.5 µm; on a wide variety of smooth barks, very common, 

throughout Florida ................................................................... A. cinchonae (Ach.) Müll. Arg. 

5. Ascomata immersed, with a broad clypeus; ascospores 20-28(-32) × 

6-8(-9.5) µm; microconidia 7-8 × 0.5 µm; on Taxodium; Dade and 

Seminole counties ................................................................................. A. taxodii R. C. Harris 

4. Ascospores mostly less than 23 × 7.5 µm.................................................................................... 6 

6. Ascospores 15-23 × 5-7.5 µm .................................................................................................. 7 

7. Ascomata usually complanate with a broad clypeus, often fusing 

with others; ascospores 17-23 × 5-7 µm; microconidia arcuate, 

20-27 × 0.5 µm; subtropical; Dade, Duval, Monroe and Sarasota 

counties .................................................................................A. planorbis (Ach.) Müll. Arg. 

7. Ascomata hemispherical to subglobose, with weak basal fringe; 

ascospores 15-22 × 5.5-7.5 µm; microconidia 7-10 × 0.5 µm; 

Massachusetts, Newfoundland, Europe.....................................[A. analepta (Ach.) Massal.] 

6. Ascospores 12-17 × 4-5.5 µm; microconidia 4-7 × 0.5 µm; rare, 

Liberty County ........................................................................................A. minor R. C. Harris 

3. Ascospores ovate, length/width ratio 1.5-2:1; 13-15 × 7-8(-10) µm; 

cells markedly unequal; microconidia 8-10 × 0.5 µm; Lake, Levy, 

Seminole and Wakulla counties...............................................................A. oblongens R. C. Harris 

2. Ascospore cell(s) pinched in ± mid cell or wall with internal ring-like thickenings 

as if additional septa were forming, often near apices (ultimately 4-celled in 

postmature stage) ................................................................................................................................. 8 

8. Ascospore wall with granular ornamentation, often with an internal 

ring-like thickening; southern ............................................................................................................ 9 

9. Ascospores 4-8/ascus................................................................................................................. 10 

10. Ascospore wall usually without internal thickening, merely 

constricted; ascospores only very rarely 4-celled when postmature, 

18-30 × 6.5-9.5(-12) µm; microconidia 6-9 × 0.5 µm; always? 

associated with Trentepohlia; throughout Florida...................................... A. lyrata R. C. Harris 

10. Ascospore wall with an internal ring-like ridge in both cells 

which forms a complete septum in postmature spores; 

ascospores 27-37 × 10-12 µm; microconidia 6-8 × 0.5 µm; 

Monroe County...............................................................................A. majuscula (Nyl.) Zahlbr. 

9. Ascospores 2/ascus, 37-48 × 15-16 µm; ostiole occasionally 

eccentric; microconidia 5-10 × 0.5 µm; common, throughout 

Florida ..............................................................................................A. malaccitula (Nyl.) Zahlbr. 

8. Ascospore wall smooth and lacking any internal thickening; northern............................................ 11 

11. Ascospores smaller, 12-15(-18) × 4-5 µm; spore cells weakly 

constricted; microconidia 6-8 × 1 µm; Massachusetts, Tennessee and 

73

West Virginia ...............................................................................................[A. degelii R. C. Harris] 

11. Ascospores (13-)15-26 × 4.5-9 µm; cells usually with stong constrictions................................ 12 

12. Ascospores with both ends rounded, 18-22 × 7-8(-9) µm; upper 

cell usually somewhat larger; asci 60-80 × 17-22 µm; microconidia 

10-12 × 0.5 µm; rare, Vermont, Newfoundland, California and 

Oregon .........................................................................[A. cinereopruinosa (Schaerer) Massal.] 

12. Ascospores with one or both ends pointed, (13-)15-20 × 4.5-6.5(-7.5) µm; 

cells ± equal; asci 75-110 × 12-18 µm; microconidia 10-12(-15) × 0.5 µm; 

Pacific Northwest...............................................................[A. plumbaria (Stizenb.) R. C. Harris] 

1. Ascospores 4-celled................................................................................................................................ 13 

13. Ascospores less than 25 µm long ..................................................................................................... 14 

14. Ascomata immersed, the tip surrounded by a broad clypeus often 

confluent with others; ostiole surrounded by a whitish ring; 

ascospores 17-22 × 6-8 µm; microconidia 6-9 × 0.5 µm; 

Dade and Monroe counties ......................................................................A. confluens R. C. Harris 

14. Ascomata lacking a broad clypeus, not confluent; ostiole not 

surrounded by a whitish ring; ascospores 17-23 × 6-7.5 µm; 

microconidia 10-13 × 0.5 µm; reported from British Columbia 

by Noble et al., 1987............................................................................[A. cerasi (Schrader) Massal.] 

13. Ascospores 28-40(-45) × 9-12(-15) µm, with distinct granular ornamentation 

and thick gelatinous epispore; microconidia 9-12 × 0.5 µm; Collier, Dade 

and Monroe counties ................................................................................. A. exasperata R. C. Harris 

NOTES 

Much of the information for this genus comes from my unpulished thesis (1975). To avoid possible 

confusion I have tried to account for all of the names included in it. The synonymy from the thesis and that 

accumulated in the 20 years since is also given. 

Arthopyrenia analepta (Ach.) Massal. 

Ric. auton. lic. crost. 165. 1852. Lichen analeptus Ach., Lichenogr. suec. prodr. 15. 1798. Type. 

SWEDEN. (H-Ach 796-B, lectotype, selected here). 

Arthopyrenia lapponina Anzi, Comment. Soc. Crittogam. Ital. 2: 25. 1864. Type. ITALY. Bormio, 

Anzi, Lich. Lang. 347 (FH-Tuck 4084, isotype). 

Verrucaria epidermidis var. fallax Nyl, Bot. Notiser 1852: 178. 1852. Verrucaria fallax (Nyl.) Nyl., 

Flora 55: 363. 1872. Arthopyrenia fallax (Nyl.) Arnold, Verhandl. Zool.-Bot. Ges. Wien 23: 

505. 1873. Type. FINLAND. Holmia, Nylander (H-Nyl 955, holotype). 

Pyrenula punctiformis var. fallax f. betulae Hepp, Flecht. Eur. 452. 1857. Type. SWITZERLAND. 

Zürich, Hepp (FH, isotype) 

Arthopyrenia fallax f. crategina Steiner in Zahlbr., Ann. Naturhist. Hofmus. Wien 24: 283. 1911. 

Type. YUGOSLAVIA. "Ad litus meridionalis lacus "Veldser See", Steiner, Krypt. Vind. 1763a 

(FH, US, isotypes). 

The material in the Acharian herbarium under Verrucaria analepta contains three pieces of bark labelled 

"Suecia" (marked A, B, C) and therefore candidates for the lectotype. Pieces A and C lack ascospores but the 

hamathecium is of the "Naetrocymbe-type". Piece C also has microconidia in accord with Naetrocymbe. 

Piece B has hamathecium, asci, ascospores and microconidia identical to those of the species currently 

known as Arthopyrenia lapponina. I have selected Piece B as the lectotype as it contains good material of 

both teleomorph and anamorph and to stabilize the usage of Arthopyrenia in the broadest possible sense (see 

74

above). The selection of piece B is further endorsed by the fact that the only specimen cited by Massalongo 

under A. analepta (Schaerer, Lich. Helv. 287) is also this species. Some probably have not gotten over the 

loss of the epithet "fallax" and may be upset by this further name change but I remind them that the epithet 

"analepta" was widely used in the past in this sense. (Of the three older European collections at NY not 

previously annotated (Czechoslovakia, England and Switzerland), two were this species and one A. 

cinereopruinosa.) 

Arthopyrenia annulata R. C. Harris ined. = A. majuscula 

Arthopyrenia antecellens (Nyl.) Arnold 

Transferred in this publication to Mycoporum. 

Arthopyrenia atomarioides Müll. Arg. = Naetrocymbe 

Arthopyrenia atractospora Zahlbr. = Naetrocymbe 

Arthopyrenia bifera Zahlbr. = A. malaccitula 

Arthopyrenia cedrina (Zahlbr.) ined. = Naetrocymbe 

Arthopyrenia cerasi (Schrader) Massal. 

Ric. auton. lic. crost. 167. 1852. Verrucaria cerasi Schrader, Ann. bot. (Usteri) 22: 86. 1797. Type 

not seen. 

Arthopyrenia cineropruinosa (Schaerer) Massl. 

Symm. lich. nov. 117. 1855. Verrucaria cinereopruinosa Schaerer, Lich.helv. spic. sect. 6, 343. 

1836. Type not seen. 

Pyrenula punctiformis var. cinereopruinosa f. hederae Hepp, Flecht. Eur. 105. 1853. Type. 

SWITZERLAND. Zürich, Hepp (FH, FH-Tuck 4073, isotypes). 

Pyrenula punctiformis var. cinereopruinosa f. pinicola Hepp, Flecht. Eur. 106. 1853. Arthopyrenia 

pinicola (Hepp) Massal., Symm. lich. nov. 118. 1855. Type. SWITZERLAND. Zürich, Hepp 

(FH, FH-Tuck 4073, isotypes). 

Arthopyrenia stigmatella var. elabens Massal., Symm. lich. nov. 120. 1855. Type. ITALY. "Ad 

truncos pinorum in Prov. Patavinia (Rua)", Massal. Lich. Ital. 202 (FH, FH-Tuck 4077, isotypes). 

Pyrenula punctiformis var. cinereopruinosa f. buxicola Hepp ex Rabenh., Lich. Eur. 630. 1862, 

nom. inval. sine descr. 

Arthopyrenia convexella Müll. Arg., Flora 66: 288. 1883. Type. BRAZIL. São Paulo: Apiaí, Dec 

1879, Puiggari 1004 (G, holotype). 

Arthopyrenia transposita Müll. Arg., Bull. Soc. Bot. Belgique 31: 39. 1892. Type. NEW ZEALAND. 

1883, Knight (G, holotype). 

Arthopyrenia fallax var. conspurcata Steiner in Beck & Zahlbr., Ann. Naturhist. Hofmus. Wien 12: 

94. 1898. Type. AUSTRIA. Carinthia, near Klagenfurt, Steiner, Krypt. Vind. 269 (US, isotype). 

Arthopyrenia ligustri Britz., Hedwigia 43 (Beibl.): (37). 1904, hom. illeg. Type. GERMANY. 

Augsburg, westliche Hugelzug, Britzelmayer (US, isotype). 

Arthopyrenia japonica Vainio, Bot. Mag. (Tokyo) 32: 163. 1918. Type. JAPAN. Izumo Prov., 1 

Mar 1914, Yasuda 101 (TUR-Vainio 32112, holotype). 

Arthopyrenia cinchonae (Ach.) Müll. Arg. 

Flora 66: 287. 1883. Verrucaria cinchonae Ach., Syn. meth. lich. 90. 1814. Type. AMERICA. On 

Cinchona officinalis (H-Ach, holotype). 

Verrucaria prostans Mont., Ann. Sci. Nat. Bot. sér. 2, 19: 53. 1843. Type. FRENCH GUIANA. 

Leprieur 215 (PC-Mont, holotype). 

Verrucaria alboatra var. detergens Nyl., Flora 52: 125. 1869. Type. BRAZIL. Prope Rio de Janeiro, 

Glaziou 1915 (M, isotype). 

75

Verrucaria concamerata Stirton, Proc. Phil. Soc. Glasgow 13: 192. 1881. Porina concamerata 

(Stirton) Zahlbr., Cat. lich. univ. 1: 377. 1922. Type. ASSAM. Watt (BM, isotype). 

Arthopyrenia nieteriana Müll. Arg., Flora 66: 288. 1883. Type. CEYLON. Nieter 30, ex hb. Hampe, 

1877 (G, holotype). 

Arthopyrenia planipes Müll. Arg., Flora 73: 345. 1890. Type. AFRICA TROPICO-ORIENTALI. Rev. J. 

Hannington (BM ex K, holotype). 

As indicated by the number of synonyms and their wide distribution A. cinchonae is common, 

conspicuous and widespread. It is one of the most common species of the oak-pine woods of Florida, mainly 

on oak but not at all particular as to substrate. Over the years I have seen hundreds of specimens from drier 

areas of both Neo- and Paleotropics. 

Arthopyrenia confluens R. C. Harris, nom. nov. 

Tomasellia (sect. Oligomeris) leucostoma Müll. Arg., Flora 68: 257. 1885. [non Arthopyrenia 

leucostoma (Ach.) Massal. (1852)] Type. "in Cort. Cascarillae ex hb. Hampe, 1877". (G, 

holotype). 

Arthopyrenia degelii R. C. Harris, sp. nov. 

Ascis sat parvis, 48-60 × 11-13 µm et ascosporis parvis, bilocularibus, uno cellulo vel ambis cellulis in 

medio constrictis, 12-15 × 4-5 µm. 

Type. TENNESSEE. Great Smoky Mountains, Cherokee Orchard, on Hamamelis, 11 Sep 1939, Degelius (US, 

holotype; FH, US, isotypes). 

Forming brown, rarely tan, blotches on smooth bark, often Hamamelis. Ascomata superficial, 

hemispherical to complanate, 0.3-0.5 mm diam., clypeate, not melanized below. Asci narrowly elliptical, 48-60 

× 11-13 µm, with 8 biseriate to subbiseriate spores. Ascospores narrowly elliptical, 2-celled, on or both cells 

constricted near the middle (spore outline sinuose), 12-15(-18) × 4-5 µm, with a thin sheath. Microconidia rodshaped, 

6-8 × 1 µm. 

Arthopyrenia degelii is characterized by the sinuose ascospore outline, small ascospores, small, narrow 

asci and small hemispherical ascomata. Arthopyrenia minor is similar in ascospore size but the ascospores 

do not have median constrictions. Arthopyrenia degelii seems ± Appalachian in distribution. 

Additional specimens. MASSACHUSETTS. Bristol County: New Bedford, Willey (US). NORTH CAROLINA. 

Gaston County: Crowder's Mountain, 11 Aug 1886, Green (NY); Haywood County: Great Smoky Mtns. Natl. 

Park, at Big Creek, 45 mi ESE of Knoxville, 2000-2300 ft, on Liriodendron, 25 Aug 1977, Wetmore 29846 

(MIN, NY); Macon/Jackson County: Whiteside Mountain, ca. 5.5 mi E of Highlands, 4400-4930 ft, on 

Hamamelis, 10 Jun 1981, Harris 13756 (NY). TENNESSEE. Carter County: Doe River Gorge, ca. 1 mi N of U.S. 

Hwy. 19E, ca. 5 mi NW of town of Roan Mountain, ca. 800 m, on Hamamelis, 26 Sep 1993, Harris 30978 

(NY). WEST VIRGINIA. sine loc., on Hamamelis, Calkins, (FH, NY); Fayette County: sine loc., on Hamamelis, 

Nuttall 85 (FH), Nuttall s.n. (MICH, NY); Greenbrier County: Greenbrier Run, 1 Jun 1880, coll. unknown (FH); 

Taylor County: Grafton, on Hamamelis, 17 Dec 1880, coll. unknown (MSC). 

Arthopyrenia esenbeckiana (Fée) R. C. Harris, comb. nov. 

Melanotheca esenbeckiana Fée, Essai crypt. écorc. suppl. 74. 1837. Tomasellia esenbeckiana 

(Fée) Müll. Arg. Type. ANTILLES. on Exostemma, (G, lectotype, selected here). 

I am still of the opinion that this species does not occur in North America as I suggested in Egan (1987). 

It has nothing to do with either Melanotheca or Tomasellia. The characters of microconidia, ascus and 

hamathecium suggest placement in Arthopyrenia as defined in this work. The compound ascoma is 

anomalous although A. planorbis (Ach.) Müll. Arg. has a strong tendency for the clypei to fuse laterally. If one 

saw only ascospores and asci, one would identify it as A. cinchonae (Ach.) Müll. Arg. 

76

Arthopyrenia exasperata R. C. Harris, sp. nov. 

Primo aspectu Polymeridi catapasti similis sed differt ascosporis exasperatis, 36-44 × 

12.5-15 µm, perisporio crasso et microconidiis longioribus, 9-12 × 0.5 µm. 

Type. FLORIDA. Collier County: Big Cypress National Preserve, Oasis Ranger Station along U.S. Hwy. 41, 

along Florida Trail N of hwy., 25°50'N, 81°03'W, Taxodium swamp, 9 Dec 1992, Buck 22976 (NY, holotype). 

Forming whitish patches on bark. Ascomata hemispherical to subglobose, mostly immersed, 0.4-0.6 mm 

diam., with slightly spreading clypeus or not, little or not melanized below. Asci cylindrical, 140-160 × 20-28 

µm, with 8 biseriate to nearly uniseriate spores. Ascospores narrowly elliptical to narrowly ovate, 4-celled, 28- 

40(-45) × 9-12(-15) µm, distinctly granular ornamented, with a thick gelatinous sheath. Microconidia 9-12 × 

0.5 µm. 

In 1975 I placed this species in Polymeridium (Trypetheliaceae) but when I revised that genus (Harris, 

1993) I realized that it did not have Trypetheliaceae type of ascus and hamathecium but that of 

Arthopyreniaceae. In 1975 I designated a different type collection but since I have not re-examined the 

material at MSC, I have chosen a recently collected and studied specimen. There is some possibility that the 

MSC material is mixed and the specimens cited below need to be restudied. 

Additional specimens. FLORIDA. Dade County: Everglades National Park, NE of Pa-hay-okee Overlook, 

Harris 2814-B, 2829-A, 2836-C (MSC); Monroe County: Key West, Jan 1898, Thaxter 122 (MICH). 

Arthopyrenia fraxini Massal. = Naetrocymbe 

Arthopyrenia lapponina Anzi = Arthopyrenia analepta 

Arthopyrenia lyrata R. C. Harris 

in Tucker & Harris, Bryologist 83: 6. 1980. 

Arthopyrenia majuscula (Nyl.) Zahlbr. 

Cat. lich. univ. 1: 332. 1922. Verrucaria majuscula Nyl., Ann. Sci. Nat. Bot. sér. 4, 20: 253. 1864. 

Type. BONIN ISLANDS. Wright (H-Nyl 824, holotype; BM, isotype). 

This was treated in my thesis as Arthopyrenia annulata ined. Any specimens so annotated are to be 

referred here. 

Arthopyrenia malaccitula (Nyl.) Zahlbr. 

Cat. lich. univ. 1: 284. 1921. Verrucaria malaccitula Nyl., J. Linn. Soc., Bot. 20: 61. 1883. Type. 

MALACCA. Water Islands, 1864, Maingay 191 (H-Nyl 1706, holotype; BM, isotype). 

Arthopyrenia bifera Zahlbr., Ann. Mycol. 33: 34. 1935. Type. FLORIDA. [Seminole County:] Sanford, 

27 Apr 1929, Rapp 80 [publ. as 86] (W, holotype). 

Didymella gigantea Räsänen, Arch. Soc. Zool. Bot. Fenn. "Vanamo" 3: 89. 1949. Type. BORNEO. 

Sandakan, Myburgh, 1921, Elmer 20019 (H, lectotype, selected here; H, isolectotype). 

Arthopyrenia megalospora Lönnr. = Naetrocymbe 

Arthopyrenia minor R. C. Harris 

in Tucker & Harris, Bryolgoist 83: 7. 1980. 

Arthopyrenia oblongens R. C. Harris, sp. nov. 

Species distincta sporis ovatis, bilocularibus, cellulis insigniter inaequalibus, 13-15 × 7-8 µm. 

77

Type. FLORIDA. Sanford, Apr 1930, Rapp (MICH, holotype; LD, isotype). 

Forming a whitish blotch on bark of ericaceous shrubs. Ascomata ± subglobose, immersed, 0.2-0.3 mm 

diam., with a broad, elongate clypeus, 0.3-0.5 × 0.7-1.0 mm, little or not melanized below. Asci narrowly 

elliptical to cylindrical, 75-110 × 14-16 µm, with 8 uniseriate to subbiseriate spores. Ascospores ovate, 2celled, 

13-15(-17) × 7-8.5(-10) µm, smooth; cells markedly unequal, the lower narrower and shorter. 

Microconidia rod-shaped, 8-12 × 0.5-1 µm. 

The broad ascospores with strongly unequal cells are anomalous in Arthopyrenia and more suggestive of 

Anisomeridium but hamathecium, ascus and microconidium type place it here. It may be possibly related to A. 

analepta having evolved unusually broad ascospores. As far as I can tell A. oblongens is restricted to 

ericaceous substrates and is known only from Florida. 

Additional specimens. FLORIDA. Lake County: Ocala National Forest, ca. 2 mi W of Alexander Springs, 

dwarf oak scrub, on Lyonia, 6 Dec 1988, Harris 23620 (NY); Levy County: Cedar Key Scrub State Preserve, 

along Co. Rd. 347 ca. 1.5 mi N of Fla. Hwy. 24, 29°12'N, 83°01'W, dry oak-ericad scrub, 30 Nov 1992, Buck 

22421, 22423, 22453 (NY), Harris 29357 (FLAS, NY); Seminole County: Sanford, 28 Jun 1928, Rapp 709 

(MICH); Wakulla County: Apalachicola National Forest, Bradwell Bay Wilderness, along Apalachicola Trail at 

Sopchoppy River, 29 Nov 1988, Harris 22979 (NY). 

Arthopyrenia padi Rabenh. = Naetrocymbe punctiformis 

Arthopyrenia planorbis (Ach.) Müll. Arg. 

Mém. Soc. Phys. Genève 30: 27. 1888. Verrucaria planorbis Ach., Syn. meth. lich. 92. 1814. 

Type. "in America ad Crotonis Cascarillae". (H-Ach, holotype). 

Arthopyrenia planior Müll. Arg., Bot. Jahrb. Syst. 6: 404. 1885. Type. CUBA. Wright, Lich. Cub. 

II: 627a, 627b (G, syntypes), 627b (US, isosyntype). 

Arthopyrenia planorbiculata Müll. Arg., Bot. Jahrb. Syst. 6: 405. 1885. Type. CUBA. Wright, Verr. 

Cub. 64 (G, holotype; US, isotype). 

Arthopyrenia plumbaria (Stizenb.) R. C. Harris 

in Egan, Bryologist 90: 163. 1987. Verrucaria plumbaria Stizenb. in Hasse, Erythrea 3: 44. 1895. 

Porina plumbaria (Stizenb.) Hasse, Contr. U. S. Natl. Herb. 17: 12. 1913. Type not seen. Pyrenula 

herrei Fink in Hedrick, Mycologia 25: 309. 1933. Type. CALIFORNIA. Santa Cruz Peninsula, Herre 

(MICH, holotype). 

Arthopyrenia rappii Zahlbr., Ann. Mycol. 33: 34. 1935. Mycoporopsis rappii (Zahlbr.) Riedl, Sydowia 

16: 272. 1962[1963]. Type. FLORIDA. Seminole County, Fort Reed, on Hicoria, Jan 1929, Rapp 

138 [publ. as 183] (W, holotype). 

This is a species of Melaspilea Nyl. s. lat. 

Arthopyrenia rhyponta (Ach.) Massal. = Naetrocymbe 

Arthopyrenia salicis Massal. 

Position uncertain. A related species is included in Naetrocymbe as a temporary expedient. Like 

Naetrocymbe it is not lichen-forming and the most satisfactory solution as far as I am concerned would be to 

exclude if from the checklist as having no relevance for lichenologists. I wish I had the "silver bullet" to kill off 

the perpetually "undead" irrelevancies in checklists. 

Arthopyrenia taxodii R. C. Harris, sp. nov. 

Sat similis Arthopyreniae cinchonae quoad ascos et ascosporas sed differt ascomatis in corticem Taxodii 

78

immersis clypeo lato et microconidiis 7-8 × 0.5 µm. 

Type. TEXAS. Houston, on Taxodium distichum, 1869, Ravenel 60 (FH-Tuck 4022, holotype; US, isotype; FH, 

MICH, PH, US, isotypes?). 

Forming a whitish blotch on bark of Taxodium. Ascomata subglobose, immersed, 0.2-0.3 mm diam. with 

a broad clypeus; wall poorly melanized. Asci cylindrical, 100-130 × 14-20 µm, with 8 nearly uniseriate to 

subbiseriate spores. Ascospores narrowly ovate to narrowly elliptical, 20-28(-32) × 6-8(-9.5) µm, smooth. 

Microconidia 7-8 × 0.5 µm. 

Arthopyrenia taxodii is distinctive in the immersed ascomata and substrate. Most species of 

Arthopyrenia occur on smooth bark of hardwoods. It seems closer to the "Peltosphaeria" group of Julella than 

to the rest of Arthopyrenia. 

Additional specimens. FLORIDA. Duval County: Jacksonville, [Calkins] (MICH); Seminole County: Sanford, 

Rapp s.n. (FLAS), 11 May 1932, Rapp 29 (FLAS), Upsala, 11 May 1932, Rapp 2014 (FLAS, MICH). 

Arthopyrenia texensis (Cooke) D. Hawksw. 

I have not seen the specimen involved and since Hawksworth's concept of Arthopyrenia was very broad, 

I cannot place the species even as to genus. 

JULELLA Fabre 

Just as I was nearly finished with this volume, Aptroot and v.d. Boom published a paper with notes on 

Julella (Mycotaxon 56:1-8. 1995). We have reviewed essentially the same group of species and come up with 

very different results. Depending on one's view point, Aptroot and v.d. Boom have indulged in an orgy of 

"lumping" or I have indulged in an orgy of "splitting". 

As with other muriform-spored taxa, the possibility exists that Julella should be merged with its 

transversely septate sister taxon, Arthopyrenia. I am reluctant to take this step as Julella is probably 

heterogenous. For instance, I am sorely tempted to reinstate Peltosphaeria Berlese and some of the tropical 

taxa seem to form a relatively distinct clade. Further, I am not happy with the specific level taxonomy 

presented below. Many of the specimens come from dry, harsh habitats and "external" characters of clypeus 

and ascomatal immersion are highly variable. Also as with many muriform-spored taxa, one is never quite 

sure ascospore differences are due to genetics or age. Even worse, the microconidia, which I normally 

consider a highly conservative character, don't seem to provide much help. I suspect that there are more host 

specific microspecies but I do not now have either enough specimens or enough time to make sense of them. 

A few names that have been placed in Julella (or Peltosphaeria) whose types I have not examined are 

included in the key or commented on in the notes despite this basic ignorance. 

1. Asci with 8 ascospores (rarely 1-several aborting).................................................................................... 2 

2. Ascospores remaining colorless ........................................................................................................... 3 

3. Asci clavate to pyriform with biseriately to irregularly arranged spores.............................................4 

4. Asci narrower clavate or short-cylindrical...................................................................................... 5 

5. Ascospores 24 µm long or longer ............................................................................................. 6 

79

6. Microconidia known............................................................................................................... 7 

7. Microconidia filiform, curved, 16-27 µm long; ascospores 

24-34 × 8.5-11 µm, with 8-10 rows of cells; Collier, Polk 

and Washington counties......................................................... J. variiformis R. C. Harris 

7. Microconidia rod-shaped, 6-9 µm long; ascospores 

24-33 × 10-15 µm, with 8(-10) rows of cells; 

Dade County.................................................................................... J. asema R. C. Harris 

6. Microconidia not known; ascospores 18-32 × 10-15 µm, 

5-7 × 2-3-septate; Hawaiian Islands; no material seen ........................................................... 

..................................................................................... [Peltosphaeria sandwichensis Petrak] 

5. Ascospores 25 µm long or shorter ............................................................................................ 7 

7. Microconidia known............................................................................................................... 8 

8. Ascospores 22-25 × 11-14 µm, rather regularly 5-6 × 2-3-septate; 

microconidia 5-8 µm long; oceanic?, Europe .......................[J. sericea (Massal.) Coppins] 

8. Ascospores 17-23 × 7-9 µm, irregularly submuriform to muriform, 

4-6 × 1-2-septate; microconidia 6-10 µm long; a single questionable 

Calkins record from Duval County ...................................J. fallaciosa (Arn.) R. C. Harris 

7. Microconidia not known; ascospores 13-19 × 7-9 µm, 

3-5 × 1-2-septate; California; no material seen .................. [Peltosphaeria californica Petrak] 

4. Asci broad, mostly pyriform but occasionally long-ellipsoidal; 

ascospores 26-38 × 12-16 µm, with 8-10 rows of cells; 

microconidia 10-20 µm long; peninsular Florida ..........................J. sublactea (Nyl.) R. C. Harris 

3. Asci long-cylindrical, 200-300 × 20-25 µm with uniseriate spores..................................................... 9 

9. Ascospores larger, 24 × 9 µm or over ......................................................................................... 10 

10. Ascospores large, 29-46 × (9.5-)11-14 µm ........................................................................ 11 

11. Ascospores 36-46 × 12-13(-15) µm, rather irregularly 

7-11 × 1-2-septate, with large cells; microconidia 6-7 µm 

long; on bark of Taxodium; Collier, Dade, Duval and 

Wakulla counties .........................................................................J. taxodii R. C. Harris 

11. Ascospores 29-37 × 11-14 µm, regularly 7-11 × 2-4-septate, 

with smaller cells; microconidia 10-11 µm long; on decorticate 

stems of shrubs; California ............................... [J. vitrispora (Cooke & Harkness) Barr] 

10. Ascospores smaller, 24-30(-35) × 9-12 µm (NB: 3 choices below).................................... 12 

12. Microconidia 5-6 µm long; ascospores 24-30 × 9-11 µm, 

with 8-9 rows of cells; on Xanthoxylum; Flagler County...................Julella sp. 2485-B 

12. Microconidia 8-9 µm long; ascospores 24-35 × 11-12.5 µm, 

with 6-8 rows of cells; Namibia.................................................................[Harris 18533] 

12. Microconidia 15-17 µm, curved; ascospores 25-30 × 9-12 µm, 

with 7-8 rows of cells; Namibia.................................................................[Harris 18519] 

80

9. Ascospores small, 16-22 × 8-9 µm; ascomata on old wood, raised, 

± constricted at base; clypeus elongated in direction of grain; 

Dominican Republic ........................................................................................[Julella sp. 19512-A] 

2. Ascospores becoming brown, ± biseriate, 28-36 × 12-16 µm; ascomata 

pyriform, immersed; anamorph in culture a Phoma;on intertidal 

mangrove wood; India, Australia .............................................................[?J. avicenniae (Borse) Hyde] 

1. Asci with 2 or 4 ascospores..................................................................................................................... 13 

13. Asci with 2 ascopores........................................................................................................................ 14 

14. Ascospores over 35 × 15 µm ...................................................................................................... 15 

15. Growing on a wide variety of bark in dry habitats; ascoma not 

strongly immersed; clypei often fused; ostioles often surrounded 

by a whitish ring; ascospores 40-51 × 17-21 µm, with 9-11 rows 

of cells; Dade and Seminole counties, also "Caloosa River", 

Austin ...................................................................................J. geminella (Nyl.) R. C. Harris 

15. Growing on Buxus sempervirens in Europe; ascoma immersed; 

clypei may be fused; ascospores (26-)40-50 × (10-)16-20 µm, 

with (4-)16 rows of cells (Barr, 1986) .............................................................. [J. buxi Fabre] 

14. Ascospores 27-35 × 10-12.5 µm, with 8-9 rows of cells; 

microconidia 14-17 × 0.5 µm; Texas ....................................... [J. dispora (Müll. Arg.) R. C. Harris] 

13. Asci with 4 ascospores...................................................................................................................... 16 

16. Ostiole apical, colorless............................................................................................................... 17 

17. Ascospores 30-36 × 12-15 µm, with 7-9 rows of cells; ascomata 

± superficial, with narrow hyphal ring; microconidia 15-18 × 0.5 µm; 

eastern North America and Europe................................................... [J. lactea (Massal.) Barr] 

17. Ascospores 30-36 × 14-17 µm with 6 rows of cells; ascomata 

immersed under a broad hyphal ring (thin clypeus?); microconidia 

7.5-11 × 0.5 µm; Australia ................................................... [J. exiguella (Bailey) R. C. Harris] 

16. Ostiole lateral, red pigmented, olive brown in KOH; ascospores 

40-50(-62) × 15-18(-22) µm, with 8-10 rows of cells; Bahamas 

and Puerto Rico....................................................................... [J. declorans (Riddle) R. C. Harris] 

Julella asema R. C. Harris, sp. nov. 

NOTES 

Julellae sublacteae similis sed ascis angustioribus et J. variiformis sed ascosporis latioribus, 24-33 × 10- 

15 µm, 7(-9) × 2-4-septatis, microconidiis 7-9 µm longis. 

Type. FLORIDA. Dade County: Everglades National Park, just N of Paurotis Pond, Paurotis-buttonwood 

hammocks surrounded by red mangrove, on Rhizophora mangle, 11 May 1967, Harris 2663-A (NY, holotype). 

81

Thallus indicated by whitish discoloration of bark. Ascomata orbicular, initially immersed and flat, 

becoming hemispherical, 0.5 mm diam., with or whithout a dark surrounding clypeoid ring; ostiole sometimes 

surrounded with pale ring. Asci oval to clavate, with 8 ascospores ± biseriately arranged. Ascospores with one 

end narrower, often slightly constricted at median septum, 7(-9) × 2-4-septate, 24-33 × 10-15 µm, often with a 

gel sheath. Microconidia rod-shaped, 6-7 µm. 

This species is perhaps poorly defined and rather characterless which is reflected in the epithet meaning 

"without a mark or sign". In ascospore size it is not very different from J. sublactea and J. variiformis, J. 

sublactea differs in broad pyriform asci, J. variiformis has narrower ascospores, both have longer 

microconidia. Dominican and Venezuelan material seems to agree quite well. 

Additional specimens. FLORIDA. Dade County: Coconut Grove, on mango, 23 Dec 1919, Britton & Britton 

665. DOMINICAN REPUBLIC. Prov. La Romana: Isla Catalina, 0-10 m, 18 Dec 1987, Zanoni et al. 37318[D]. 

VENEZUELA. Falcón: Península de Paraguaná, Cumbre de Cerro Colorado, zona xerófita, 200 m, López & 

Wingfield 22504 (all NY). 

Julella avicenniae (Borse) Hyde 

This species is retained in Julella with considerable hesitation. The brown ascospores and Phoma 

anamorph are anomalous. 

Julella dactylospora Rehm 

Hedwigia 40: 110. 1901. 

= Anthracothecium columellatum (Vainio) Zahlbr. fide Aptroot & v. d. Boom (1995). 

Julella decolorans (Riddle) R. C. Harris, comb. nov. 

Campylothelium decolorans Riddle in Britton & Millspaugh, Bahama Fl. 532. 1920. Type. 

BAHAMAS. Exuma Chain: Cay N of Wide Opening, on Drypetes, 18 Feb 1905, Britton & 

Millspaugh 2795 (FH, holotype; NY, isotype). 

The red pigmented ostiole and ± 4-spored asci make this a very distinctive species. The asci are initially 

8-spored but 4-6 routinely abort with most commonly 4 surviving although other numbers are encountered. It 

seems to be a dry forest species and I would expect it to turn up in Florida. 

Julella dispora (Müll. Arg.) R. C. Harris, comb. nov. 

Polyblastia dispora Müll. Arg., Flora 60: 79. 1877. Polyblastiopsis dispora (Müll. Arg.) Zahlbr., Cat. 

lich. univ. 1: 348. 1922. Type. TEXAS. Near Dallas, Boll (US, isotype). 

?Polyblastia fallaciuscula Müll. Arg., Bot. Jahrb. Syst. 6: 407. 1885. Polyblastiopsis fallaciuscula 

(Müll. Arg.) Zahlbr., Cat. lich. univ. 1: 349. 1922. Type. CUBA. Wright, Verr. Cub. 131. 

The fragment of the holotype of Polyblastia fallaciuscula I have seen lacked ascospores. It is tentatively 

assigned here based on the ascospore size given by Müller, 26-30 × 12-15 µm. Geographically it is more 

likely to be an immature specimen of J. geminella. 

Julella exiguella (Bailey) R. C. Harris, comb. nov. 

Trypethelium exiguellum Stirton ex Bailey, Queensland Agric. J. 5: 39. 1899. Type. AUSTRALIA. 

Queensland: Thursday Island, Jun 1897, Bailey s.n. (BM, holotype?). 

Julella fallaciosa (Arn.) R. C. Harris 

Mycoglaena yasudae Vainio, Bot. Mag. (Tokyo) 35: 75. 1921. Type. JAPAN. Prov. Kozuke, 7 May 

1918, Yasuda 238 (TUR-Vainio 31023), syn. nov. 

Julella geminella (Nyl.) R. C. Harris, comb. nov. 

Verrucaria geminella Nyl., Expos. syn. pyrenocarp. 40. 1858. Polyblastiopsis geminella (Nyl.) 

Zahlbr., Cat. lich. univ. 1: 349. 1922. Type. "Ad cortices in Mexico." (not seen). 

82

?Polyblastiopsis alboatra Zahlbr. in Rechinger, Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr. 

81: 230. 1907. Type. SAMOA. Upolu, auf Clerodendron im Mangrove Sumpfe bei Mulinuu, 

Rechinger 2797 (not seen). 

Polyblastiopsis rappii Zahlbr., Ann. Mycol. 33: 36. 1935. Type. FLORIDA. [Seminole County:] 

Sanford, on Amorpha fruticosa, Apr 1933, Rapp 133 (W, holotype; FLAS F-27978, isotype). 

Although I have not seen the types of Verrucaria geminella or Polyblastiopsis atroalba, the 2-spored asci 

and ascospore size suggest this disposition. Julella geminella is apparently pantropical, with specimens seen 

from Florida, Bermuda, Puerto Rico, Bequia, St. Lucia, Tanzania, India, Hong Kong and Papua New Guinea. 

Julella leopoldina Rehm, Ann. Mycol. 6: 319. 1908. 

From the description I suspect this is an Anthracothecium or Pyrenula. 

Julella luzonensis P. Henning 

Hedwigia 47: 257. 1908. 

= Pyrenula sp. (Harris, 1989) or Anthracothecium prasinum (Eschw.) R. C. Harris (Aptroot & v.d. Boom, 

1995). The type collection may be mixed and the typification needs to be carefully investigated. 

Julella macrospora Döbbler, Mitt. Bot. München 14: 191. 1978. 

Julella phycophila Döbbler, Mitt. Bot. München 14: 193. 1978. 

Both are described as being aparaphysate. They certainly have nothing to do with Julella. However, I 

am not aware that they have been transferred elsewhere. 

Julella manakonensis Kern, Ber. Schweiz Bot. Ges. 69: 282. 1959. 

From the description and illustrations I surmise that this is referable to either Anthracothecium or 

Pyrenula. 

Julella sericea (Massal.) Coppins 

Purvis et al. (1992) and Aptroot and v.d. Boom (1995) suggest that J. sericea might be synonymous with 

J. fallaciosa. From the few specimens of J. sericea (including Massal. 262) I have studied, the ascospores 

seem broader, more ellipsoidal, more divided and more regularly divided. The distribution seems 

Mediterranean/ oceanic. I am inclined to keep the taxa separate. 

Julella sublactea (Nyl.) R. C. Harris 

Verrucaria sublactea Nyl., Flora 69: 464. 1886. Type. CORFU. S. Nicolo[?], an junger Oelbäume, 

May 1886, P. Sydow s.n. (NY, isotype?). 

Polyblastia transwaalensis Müll. Arg., Flora 70: 79. 1887. Type. SOUTH AFRICA. Transvaal: 

Lydenburg, Wilms, comm. Lahm no. 52 (G, holotype). 

Mycoglaena collosporella Vainio, Bol. Soc. Broteriana, sér. 2, 6: 176. 1930. Type. MOZAMBIQUE. 

Palma, 1916, Pires de Lima 774 (TUR-Vainio 34328, holotype). 

Clathroporina exiguella Zahlbr., Ann. Mycol. 33: 38. 1935. Type. FLORIDA. [Seminole County:] 

Sanford, on Garberia fruticosa, Oct 1932, Rapp 119 (FLAS F-30032, isotype). 

Clathroporina amygdalina sensu Fink, Lich. Fl. U.S. 56. 1935. 

This species is distinctive in having a whitish thallus and long, unusually variable microconidia. I had 

originally thought that Polyblastia alba Müll. Arg. from South Africa was synonymous but the type has slightly 

smaller ascospores and microconidia only 8-10 µm long. It antedates Verrucaria sublactea by a few months 

and would have priority if my present stance turns out to be incorrect. McCarthy (1995) suggested that 

Clathroporina exiguella should be referred to Thelenella. One collection from Puerto Rico and one from 

Venezuela seem referable to P. alba. 

Julella taxodii R. C. Harris, sp. nov. 

Ascomatis immersis clypeis grandis, 1-1.5 mm diam., ascis anguste cylindraceis, 220-300 × 20-25 µm, 

ascosporis uniseriatis, 9-10 × 1-2-septatis, 36-46 × 12-15 µm, microconidiis bacilliformibus, 6-7 µm et ad 

Taxodium crescentibus. 

83

Type. FLORIDA. Dade County: Everglades National Park, cypress dome between Mahogany Hammock and 

Pa-hay-okee Overlook, on Taxodium, 16 May 1967, Harris 2947-A (NY, holotype). 

Thallus not evident or a whitish discoloration of the bark. Ascomata immersed to somewhat emergent 

with age, ± spherical, 0.45-0.7 mm diam., usually with a broad clypeus, 0.6-1.8 mm diam. Asci narrowly 

cylindrical, 200-300 × 20-25 µm, often with a long stipe, with 8 uniseriate ascospores. Ascospores fusiform, 

rather irregularly 9-10 × 1-2-septate with large cells, 36-46 × 12-13(-15) µm, often with an obvious gel sheath. 

Microconidia rod-shaped, 5-7 × 0.5 µm. 

Julella taxodii is conspicuous on Taxodium due to the very broad clypeus and is probably locally 

common. It was widely distributed by Calkina under the name Pyrenula subcynerea. Its nearest relative is 

probably J. vitrispora from which it differs in larger ascospores and shorter microconidia. There are two 

Greater Antillean collections which are very close, differing mainly in substrate. They are included here 

tentatively. 

Additional specimens. FLORIDA. Collier County: Big Cypress National Preserve, Oasis Ranger Station 

along U.S. Hwy. 41, along Florida Trail N of hwy., 9 Dec 1992, Buck 22957 (NY); Dade County: NE of Pa-hayokee 

Overlook, 3 May 1967, Harris 2819, 2832, S of Pa-hay-okee Overlook, 12 May 1967, Harris 2779-A 

(NY); Duval County: Jacksonville, Calkins (NY); Wakulla County: Appalachicola National Forest, along Co. Rd. 

368, 3.6 mi from Co. Rd. 319 at Crawfordsville, 29 Nov 1988, Harris 22948 (NY). GEORGIA. Ware County: 

Okefenokee National Wildlife Refuge, N end of Cedar Hammock Canal, 3 mi NW of boat landing, 25 Nov 

1989, Wetmore 65182 (MIN, NY), 1 mi W of Chesser Island along boat trail, 29 Nov 1989, Wetmore 65450 

(MIN) (all on Taxodium). 

Not on Taxodium. DOMINICAN REPUBLIC. Prov. Monte Cristi: El Morro de Monte Cristi, 237 m, limestone 

mesa by sea, 19°54'N, 71°39'W, 8 Jan 1987, Buck 13947 (NY). PUERTO RICO. Bosque Estatal de Guajataca, 

Vereda Juan Pérez near S end of reserve, 18°24'N, 66°58'W, ca. 300 m, humid forest, 17 Jan 1992, Harris 

27876 (NY). 

Julella variiformis R. C. Harris, sp. nov. 

Ascomatis hemisphaericis, ascosporis 25-33 × 9-10.5 µm, 7 × 1-3-septatis et microconidiis filiformibus, 

arcuatis, ca. 22 µm longis. 

Type. PUERTO RICO. Mun. Cabo Rojo: Sierra Bermeja, Cerro Mariquita, from W end of Laguna Cartagena to 

summit, on cactus, ca. 18°00'N, 67°07'W, 35-300 m, dry forest over chert, 15 Jan 1992, Harris 27732 (NY, 

holotype). 

Thallus indicated by whitish or grayish area of disrupted bark. Ascomata extremely variable, from forms 

with immersed ascomata and the clypeus flush with the bark surface and a whitish ring around the ostiole 

(young forms?) to forms ± superficial and hemispherical with a dark hyphal, clypeoid ring well developed to 

absent, on barks which erode occasionally and thus raised on pedestals of bark held together by the ascoma, 

black, centrum ca. 0.5 mm diam. but clypeus to 2 mm diam. and irregular groups of fused clypei to several 

mm; clypeus dense to diffuse hyphae in upper bark layer. Asci fusiform to ± clavate, with 8 mostly biseriate 

ascospores. Ascospores with long-ovoid or short-cylindric, 7 × 1-3-septate, 24-34 × 8.5-11 µm, often with an 

obvious gel sheath. Microconidia filiform, arcuate, (16-)22-27 × 0.5 µm. 

As noted above in this circumscription the ascomata are extremely variable and hence the specific 

epithet. The type is a form with superficial hemispherical ascomata. It is certainly possible that several 

microspecies are included. The characters that hold the putative taxon together are the filiform microconidia 

and the medium sized, rather narrow ascospores. Julella variiformis may be difficult to separate from J. 

asema without microconidia and one has to depend on ascospore width. Julella variiformis seems to be a 

species of exposed, often dry, scrub habitats. 

84

Additional specimens. FLORIDA. Collier County: Big Cypress National Preserve, Oasis Ranger Station on 

U.S. Hwy. 41, along Florida Trail N of hwy., on Bumelia, 9 Dec 1992, Harris 30153; Polk County: Nalcrest, 12 

Mar 1989, Wheeler s.n.; Washington County: Rock Hill Preserve, 4.1 mi SE of Chipley, Griffin 2919[A]. 

BAHAMAS. Cat Isalnd: The Bight and Vicinity, on Coccoloba krugii, 1-6 Mar 1907, Britton & Millspaugh 5922. 

PUERTO RICO. Reserva Forestal Guánica, Bahía de la Ballena at Km 9.5 on Hwy. 333, ± sea level, dry coastal 

scrub over rough limestone, 26 May 1989, Harris 23984, 23985, 24013; Vereda El Ojo de Agua at Km 3.8 on 

Hwy. 334, ca. 600 ft, semi-dry scrub, 1 Jun 1989, Harris 24152; Maricao State Forest, Santana Ridge, 

secondary moist scrub, 27 May 1989, Harris 24044; Bosque Estatal de Susúa, S edge of reserve, ca. 275 m, 

dry scrub forest over exposed serpentine, 13 Jan 1992, Buck 21268; as type, Harris 27712-A (all NY). 

Jullela vitrispora (Cooke & Harkness) Barr 

E. Britton 4367 (NY) from Puerto Rico on Byrsonema has ascospores which fall into the range of J. 

vitrispora but differs in fewer, less regular longitudinal septations and in microconidia only 4-6 µm long. If 

future collections maintain these differences, it should be given taxonomic recognition. 

Julella zenkeriana P. Henning 

Bot. Jahrb. Syst. 38: 127. 1905. 

= Laurera megasperma (Mont.) Riddle fide Aptroot & v.d. Boom (1995). 

MYCOMICROTHELIA Keissler 

?Ornatopyrenis Aptroot, Biblioth. Lichenol. 44: 127. 1991. Type. Microthelia queenslandica Müll. Arg. 

Ornatopyrenis was assigned by Aptroot to the Trypetheliaceae. I have seen no material. The reasons 

for tentatively assigning it to synonymy of Mycomicrothelia are discussed in the introduction to the 

Trypetheliaceae. 

KEY TO FLORIDA SPECIES 

1. Ascospores 13.5-16 × 6-7 µm (Hawksworth, 1985); on a variety of trees, 

perhaps favoring Myrica;common, throughout Florida.................M. subfallens (Müll. Arg.) D. Hawksw. 

1. Ascospores 17-20(-21) × 7-8(-9) µm (Hawksworth, 1985); on Fraxinus; 

Duval, Liberty and Sarasota counties............................................ M. willeyana (Müll. Arg.) D. Hawksw. 

NOTES 

Mycomicrothelia oleosa Aptroot 

This species is unique in having an inspersed hymenium. Its range is extended to Brazil. Brako 5640 

shows some endospore development forming a ring near the ends of the ascospores forming a rounded 

chamber as in some species of the sister genus Arthopyrenia. In Arthopyrenia these endospore rings often go 

on to form septa but that was not observed here. 

BRAZIL. Mato Grosso: Serra do Cachimbo, 763 km N of Cuiabá on Cuiabá-Santarém highway (Br-163), 

ca. 400 m, 09°35'S, 54°55'W, 22 Apr 1983, Brako 5640; Pará: Serra do Cachimbo, Aeroporta Cachimbo, 20 

km N of border with Mato Grosso on Cuiabá-Santarém highway (Br-163), 430-480m, 09°22'S, 54°54'W, 27 

Apr 1983, Brako 6208, near Base Aerea do Cachimbo, along Cuiabá-Santarém highway (Br-163), 430-480 m, 

ca. 09°22'S, 54°54'W, 28 Apr 1983, Brako 6323 (all INPA and NY). 

Mycomicrothelia rubrostoma Aptroot 

Aptroot either did not notice or ignored the fact that this species has well developed endospore (to 4 µm) 

giving the locules a bowling pin shape. Since all the other characters seem to agree with Mycomicrothelia, I 

am satisfied that it is correctly placed. There is a second species with this ascospore type in Brazil with 

shorter, elliptical to subspherical ascospores. It is left undescribed pending more ample material. 

85

MELANOMMATALES s. lat. 

PYRENULACEAE Rabenh. 

Since my treatment of Pyrenulaceae in eastern North America (Mem. New York Bot. Gard. 49: 74-107. 

1989) and the preliminary treatment in "Some Florida Lichens", Aptroot has published a treatment of the 

"peripheral" genera of Pyrenulaceae and Requienellaceae (Biblioth. Lichenol. 44: 1-178. 1991). Unfortunately 

this work is flawed by numerous careless errors and poor observation. An extensive critique of the 

morphological characters and cladistic results was published by Eriksson (Lichenologist 25: 307-311. 1993). 

(From the point of view of a cladistic-know-nothing, Aptroot's errors, especially incorrect generic and familial 

placements, would seem to make many of Eriksson's comments on the cladistic trees rather irrelevant.) In 

spite of these problems much of the skeleton is sound and should not be discarded. Further, as a result of reexamining 

some of the material studied by Aptroot, I have modified some of my ideas regarding the familial 

placement of some of these genera. I had believed that the taxa with short, coarse, essentially unbranched 

physes should be referred to the Requienellaceae. Restudy of hamathecial and ascal characters in Pyrenula 

have led me to think that these are differences of degree and not kind. Requienella has a unique ascus type 

with a "tholus" staining with various dyes (Boise, 1986). The taxa previously thought to belong to the 

Requienellaceae also have rod-shaped microconidia, shorter than the filiform, often curved microconidia of the 

Pyrenulaceae but, again, it seems to be a matter of degree. Requienellaceae sensu Aptroot seem to be nonlichen-forming. 

I surmise this may be an adaptation to xeric habitats similar to Polymeridium in the 

Trypetheliaceae. As a consequence I am restricting the Requienellaceae to Requienella, transferring the 

other genera placed in the family by Aptroot to the Pyrenulaceae along with the resurrected genera, 

Acrocordiella and Mycopyrenula. Another possibility, of course, would be to erect another family to 

accommodate Distopyrenis, Granulopyrenis, Lachrymospora and Pyrenographa and possibly Acrocordiella 

and Parapyrenis. The uniformly rod-shaped microconidia of the first four (not filiform as in Pyrenulaceae s. 

str.) make this a tempting solution (microconidia not known for the last two). Perhaps adding to the confusion, 

there are a wide variety of ascomatal types, pyrenuloid, parathelioid, pyrenastroid and possibly even 

mycoporoid, none of which seems to align with ascospore type. There are so few specimens known it seems 

best to more or less maintain the status quo pending more material and, perhaps, even molecular studies 

which would seem useful in dealing with what seems to be a highly reduced group. 

In 1989 when I abandoned ascomatal orientation and arrangement as generic characters, I did not 

mention the case where many ascomata with apical ostioles are included within a pseudostroma filled with 

pigment and/or oxalate(?) crystals. This type is very common in the Trypetheliaceae but very rare in the 

Pyrenulaceae. To repair the previous omission two such taxa are included below although neither is reported 

from Florida, P. concastroma and P. wrightii. 

KEY TO GENERA OF PYRENULACEAE AND REQUIENELLACEAE 

1. Ascomata perithecioid............................................................................................................................... 2 

2. Ascospores some shade of brown, mostly thick walled........................................................................ 3 

3. Ascospores 4-celled to muriform; physes long and slender 

(except in Acrocordiella and Mycopyrenula which have 4-celled 

ascospores and are non-lichenized) ................................................................................................ 4 

4. Lichenized; thallus usually conspicuous........................................................................................ 5 

86

5. Endospore absent or poorly developed..................................................................................... 6 

6. Ascomata with an outer pseudostromatic shell; ascospores 

dark brown, relatively thin walled (endospore poorly 

developed); lumina nearly cylindrical or cubical; mainly 

subtropical and tropical .............................................................................. Anthracothecium 

6. Ascomata lacking outer pseudostromatic shell; ascospores 

medium brown or yellow brown; thin walled (endospore not 

readily detectable); temperate North America and Europe 

............................................................................................ [Eopyrenula, see Dacampiaceae] 

5. Endospore well developed; various shades of brown; 

lumina rounded; mostly distoseptate............................................................................Pyrenula 

4. Not lichenized; no thallus evident.................................................................................................. 7 

7. Ascospores 4-celled; physes sparse, coarse; Europe .............................................................. 8 

8. Ascomata immersed in twigs of Ribes ............................................................. [Acrocordiella] 

8. Ascomata superficial on young bark of Betulaceae ........................................[Mycopyrenula] 

7. Ascospores 3-7 transversely septate to submuriform............................................................... 9 

9. Occurring in Europe and eastern North America; ascospores 

not distoseptate with a single eccentric euseptum; physes 

relatively long, abundant; ascomata large, immersed in soft 

bark of deciduous trees.................................................................... [Requienella, see below] 

9. Occurring in the West Indies; ascospores with 4 distosepta and 

a single terminal euseptum; known only from the type (not seen 

by me, placed only provisionally in Pyrenulaceae) .......................................... [Polypyrenula] 

3. Ascospores 2-celled or rarely submuriform (no 4-celled ascospores 

yet known in this group); physes coarse, relatively short, rather abruptly 

tapering; non-lichenized except for Clypeopyrenis; microconidia rod-shaped, 

to 15 µm, longer in Clypeopyrenis................................................................................................... 10 

10. Ascospores with well developed endospore, larger; not 

lichenized; physes coarse ........................................................................................................... 11 

11. Ascospores 2-celled.............................................................................................................. 12 

12. Ascospores elliptical, oblong or subspherical; septum median........................................ 13 

13. Ascospores with unlayered endospore without pigment granules ............................. 14 

14. Ascus tip thickened, with a distinct apical chamber (may be 

necessary to use stain + KOH) .......................................................... Distopyrenis 

14. Ascus wall uniform, thin, without obvious apical chamber; 

Indonesia, Australia?...........................................................................[Parapyrenis] 

13. Ascospores with an inner colorless layer surrounding the lumina; 

pigment granules in the endospore......................................................[Granulopyrenis] 

12. Ascospores teardrop-shaped, with septum strongly eccentric; 

87

Madagascar............................................................................................... [Lachrymospora] 

11. Ascospores submuriform ...................................................................................................... 15 

15. Ascoma highly melanized, elongated and somewhat 

graphid-like in aspect; "Pacific", from Galapagos to 

Australia ......................................................................................................[Pyrenographa] 

15. Ascoma with eccentric ostiole, not resembling a graphid .........................[Granulopyrenis] 

10. Ascospores 2-celled, with only the slightest hint of endospore 

at the median septum, ca. 7 × 2.5-3 µm; thallus well developed, 

lichenized; physes slender, Pyrenula-type; West Indies, 

Central America and northern South America ........................................................ [Clypeopyrenis] 

2. Ascospores colorless .......................................................................................................................... 16 

16. Ascospores ovate, 4-celled; ascomata not pigmented, simple or 

compound (astrothelioid or pyrenastroid); ostiole apical or lateral.............................. Lithothelium 

16. Ascospores ± cylindrical, 4-celled, often bent, ca. 15-21 × 3-4.5 µm; 

ascoma not compound, with apical ostiole covered with orange 

pigment, KOH+ purple; South Africa .......................................................................[Pyrenowilmsia] 

1. Ascomata mazedioid............................................................................................................................... 17 

17. Ascospores 4-celled, 11-13 × 6-8.5 µm, dark brown with lenticular cells; 

ascomatal wall melanized; thallus UV+ yellow (lichexanthone); 

south to Polk County .................................................Pyrgillus javanicus (Mont. & v.d. Bosch) Nyl. 

17. Ascospores submuriform, subspherical; ascomatal wall not 

melanized; one species in Costa Rica, second in New Guinea .............................. [Mazaediothecium] 

ACROCORDIELLA O. Eriksson 

When I re-examined material of Acrocordiella occulta, I was struck by the difference in the physes and in 

ascospore development from Requienella. Consequently I support the position that Eriksson has consistently 

and correctly maintained that Acrocordiella should have generic status. There is only a single species, A. 

occulta (Romell) O. Eriksson, known only from Europe. 

ANTHRACOTHECIUM Massal. 

1. Ascospores large, over 50 µm long........................................................................................................... 2 

2. Ascospores 8/ascus; ostiole apical ........................................................................................................ 3 

3. Ascospores muriform, 60-95 × 27-40 µm; peninsular 

Florida .........................................................................................A. prasinum (Eschw.) R. C. Harris 

3. Ascospores transversely septate to submuriform, 50-65 × 20-29 µm; 

Dade, Highlands and Seminole counties ....................................................... A. varians R. C. Harris 

2. Ascospores 2/ascus, 130-180(-225) × 38-50 µm; ostiole lateral; 

common, throughout the state .............................................................A. nanum (Zahlbr.) R. C. Harris 

88

1. Ascospores small, under 20 µm long ........................................................................................................ 4 

4. Ascospores flattened, lozenge-shaped, with four rows of two cells each .............................................. 5 

5. Hymenium inspersed; ascospores 11-13 × 6-8 µm; thallus UV-; 

north to Putnam County ...............................................................A. canellae-albae (Fée) Müll. Arg. 

5. Hymenium not inspersed; ascospores 14-18 × 7-9 µm; thallus UV+ 

yellow or UV-; Jefferson County............................ A. staurosporum (Tuck. ex Willey) R. C. Harris 

4. Ascospores globose to subglobose, cruciately divided into four cells, 

2-13 µm across or 12.5-15 × 10-13 µm; thallus UV+ yellow (lichexanthone); 

Franklin County, otherwise peninsular Florida .................................................A. subglobosum Riddle 

NOTES 

Anthracothecium staurosporum (Tuck. ex Willey) R. C. Harris 

In Harris (1989) the thallus is reported as UV-. I have since seen specimens from Texas and French 

Guiana in addition to the Florida collection which contain lichexanthone. Since they do not differ otherwise 

from more northern UV- material, I consider it to be minor chemical variation. 

DISTOPYRENIS Aptroot 

I am quite uncomfortable with Aptroot's choice of a generic type, D. americana. It is anomalous in terms 

of distribution (temperate), substrate (Betula), ascospores (almost mischoblastic with a median swelling) and 

possibly ascomatal structure. It raises the specter that Distopyrenis should be restricted to its type. The other 

species would then require a new generic name or could perhaps be dumped into the closely related 

Granulopyrenis. Only the single, somewhat aberrant species, D. submuriformis, is in Florida. 

1. Ascospores 2-celled; ascomata various.................................................................................................... 2 

2. Ostiole apical or ostiole orientation not known ...................................................................................... 3 

3. Growing on a variety of substrates other than Betula ....................................................................... 4 

4. Ascomata small, less than 0.5 mm diam.; ascospores without 

strong median septum.................................................................................................................. 5 

5. Ascospores with broadly rounded ends ................................................................................... 6 

6. Ascospores with a narrow ring in middle of septum where 

lumina meet (two dark dots in optical section),12-14 × 8-9.5 µm; 

ascomata apparently compound, mycoporoid or pyrenastroid?; 

Puerto Rico.................................................................................. [D. composita R. C. Harris] 

6. Ascospores with a narrow dark ring at outer ends of median septum 

(two dark dots in optical section), 12-15 × 7.5-9.5 µm; ascomata 

simple; Haiti.............................................................................. [Distopyrenis sp. Buck 9033] 

5. Ascospores tapering at ends, 15-18 × 8-10 µm; ascomata simple; 

ostiole orientation not evident; Venezuela.................................[Distopyrenis sp. Funk 6254-A] 

4. Ascomata larger, 0.6-0.8 mm diam.; ascospores with a dark 

median plate (euseptum?), ca. 14-16 × 7.7-8.5 µm; 

89

on Beaucarnia recurvata, Mexico.................................................................[D. fuliginosa Aptroot] 

3. Growing on smooth bark of Betula; ascospores appearing almost 

mischoblastic, often swollen in the middle, ca. 18-19 × 7 µm; 

eastern North America .................................................................................... [D. americana Aptroot] 

2. Ostiole lateral ........................................................................................................................................ 7 

7. Ascospores small, 18 × 8 µm or less ............................................................................................... 8 

8. Ascospores 13-18 × 7-8 µm; Texas, Dominican Republic, 

French Guiana ....................................................................................... [D. pachyospora Aptroot] 

8. Ascospores narrower, 14-17 × 5-6.5 µm; Cuba ............................. [Distopyrenis sp. Buck 23459] 

7. Ascospores relatively large, 16-19 × 9-10.5 µm; on bark of 

white oak; Mississippi.............................................................................. [D. quercicola R. C. Harris] 

1. Ascospores submuriform, 3 × 1-2 irregularly septate, 13-17 × 7-9 µm; 

ascoma with extensive clypeus elongated in direction of wood fibers; 

ostiole apical; on eroded wood; Monroe County .......................................D. submuriformis R. C. Harris 

NOTES 

Distopyrenis americana Aptroot 

Previously recorded from Iowa, New York and North Carolina, the range is extended to Wisconsin. 

WISCONSIN. Bayfield County: Rainbow Lake Wilderness, Chequamegon National Forest, NW of Muck 

Lake and SW of Beaver Lake, on yellow birch, 12 Aug 1992, Wetmore 71414 (MIN). 

Distopyrenis composita R. C. Harris, sp. nov. 

Ascomatis compositis, rare simplicibus, positione ostioli ignota, ascosporis late ellipsoideis annulo parvo 

centrali, 11.5-13.5 × 8-9.5 µm. 

Type. PUERTO RICO. Distr. Aguadilla: Montañas Ayamon, Bosque Estatal de Guajataca, at N end of forest on 

Hwy. 446, 200-250 m, along trail #1 to observation tower, mesic forest over limestone, 25 May 1989, Harris 

23966 (NY, holotype). 

Thallus indicated by slight discoloration of bark and black lines when in contact with other thalli. 

Ascomata flattened, often somewhat lumpy, orbicular to irregular, usually compound, rarely simple (young 

stage?), 0.2-0.5 mm diam. (individual ascomata ca. 0.1-2 mm diam.); clypeus usually covering several 

hymenia; ostiole orientation and mode of ascomatal joining unknown. Asci cylindrical to obclavate, with 

uniseriately to irregularly arranged ascospores. Ascospores broadly ellipsoidal with rounded to somewhat 

angular lumina, with a odd little dark ring in septum where the lumina come together, 11.5-13.5 × 8-9.5 µm. 

Microconidia not found. 

I apologize but I am simply unable to figure out the structure of the ascomata as they are so small. It will 

require microtome sections or an extremely lucky hand section to determine this character. However, the 

obviously compound nature of most of the ascomata conjoined with the small central ring in the ascospores 

sufficiently distinguishes this species. It is known only from the type collection. 

Distopyrenis fuliginosa Aptroot 

Microconidia from holotype, rod-shaped, 8-10 µm. Buck 4791 and Harris 15854 from the Dominican 

Republic cited by Aptroot as D. fuliginosa are D. pachyspora. Buck 9033 from Haiti is an undescribed species. 

90

Distopyrenis pachyospora Aptroot 

Aptroot described the ostiole as apical but my drawings from the type show it as strongly eccentric as it is 

in the other material cited here. Buck 4791 and Harris 15854 cited by Aptroot as D. fuliginosa are D. 

pachyospora. 

New to North America. TEXAS. Bastrop County: Bastrop State Park, just E of Bastrop, on Quercus, 19 

Mar 1976, Egan EL-8193 (MIN, NY); Robertson County: Boy Scouts of America Camp Arrowmoon, 6 mi E of 

Hearne off FM 2549, on Quercus, 10 Apr 1976, Egan EL-8355 (MIN). 

Distopyrenis quercicola R. C. Harris, sp. nov. 

Ostiolis excentris et ascosporis ovalibus sat grandis, 16-19 × 9-10.5 µm. 

Type. MISSISSIPPI. Smith County: Bienville National Forest, Marathon Lake Recreation Area, oak-pine woods 

on W shore of lake, on white oak, 30 Sep 1992, Harris 28782 (NY, holotype). 

Thallus indicated by whitening of bark. Ascomata mostly immersed, ± oval to pear-shaped, ca. 0.3 × 0.4- 

0.5 mm. Ostiole eccentric with a well developed gelatinous periphysoid plug. Asci obclavate, with no or poorly 

developed ocular chamber, with irregularly arranged ascospores. Ascospores oval with large rounded to 

obovoid lumina, euseptum not evident, 16-19 × 9-10.5 µm. Microconidia rod-shaped, 8-9 × 1 µm. 

Distopyrenis quercicola is similar to D. pachyspora in ascoma morphology but differs in the larger 

ascospores with larger lumina. It is known only from the type collection. 

Distopyrenis submuriformis R. C. Harris, sp. nov. 

Species unica ascosporis submuriformibus, 3 × 1-2 septatis, 13-17 × 7-9 µm et ascomatis similibus 

Pyrenographae. 

Type. FLORIDA. Monroe County: Everglades National Park, road W of West Lake through mangrove swamp 

and hammock to ocean, on old wood, 14 May 1967, Harris 2848-D (NY, holotype). 

Thallus indicated by pinkish white discoloration of wood. Ascomata oval to long oval, elongated with 

grain of wood, strongly melanized, sessile or on a small pedestal of wood (due to erosion of wood away from 

ascomata?), 1.2 × 0.7-1.0 mm. Ostiole central and apical. Asci cylindrical with uniseriate ascospores. 

Ascospores ellipsoidal, submuriform, 3 × 1-2 septate, 13-17 × 7-9 µm. Microconidia not found. 

This is a troubling species. The submuriform ascospores and Pyrenographa-like ascomata suggest 

placement in Pyrenographa. However, the ascospores have better developed endospore and a "Rinodinalike" 

2-celled stage in development unlike Pyrenographa where the young ascospores are relatively thin walled 

and do not form a strongly thickened endospore. Therefore, the species is provisionally assigned to 

Distopyrenis. The collection was made in 1967 providing some insight into how long it takes for systematic 

knowledge to accumulate to a point where some specimens can even be provisionally assigned to a family 

and genus. 

Distopyrenis sp. Buck 9033 

This is a rather poor collection and I am unable to determine ostiole orientation. The ascospores have a 

dark ring at the margin of the septum unlike D. composita where it is central. Its formal description awaits 

more adequate material as do two other neotropical species noted in the key. 

GRANULOPYRENIS Aptroot 

91

My first inclination was to lump Granulopyrenis, Distopyrenis and Parapyrenis. Aptroot distinguished 

Granulopyrenis on the basis of pigment granules in the endospore. In the broader context of the 

Pyrenulaceae this would not be significant. However, the ascospores are also distinctive in possessing a ± 

colorless inner endospore layer immediately adjacent to the lumen. This is also seen in Pyrenula rubrostoma 

R. C. Harris. In some ways it is more reasonable to lump and have only one genus which in terms of 

ascomatal morphology, biology, ecology and distribution don't make much sense rather than two such genera. 

In the context of the part of the Pyrenulaceae with short, coarse physes, this distinctive ascospore type can 

serve to define a genus to maintain a little controversy and encourage further investigation of generic limits. 

Aptroot's concept is expanded to include a species with submuriform ascospores. It seems odd that no 

species with 4-celled ascospores are known (possibly hidden in unsuspected non-lichen-forming genera?). 

Aptroot reported microconidia from only G. macrocarpoides but they are present and uniform in two other taxa. 

1. Ascospores 2-celled.................................................................................................................................. 2 

2. Ascomata solitary; ostiole apical or lateral ............................................................................................ 3 

3. Ostiole eccentric or lateral................................................................................................................. 4 

4. Ascus without ocular chamber (fide Aptroot); ascospores 

13-15 × 8-9 µm; Puerto Rico, Virgin Islands, Lesser Antilles .......................[G. antillensis Aptroot] 

4. Ascus with rounded ocular chamber (fide Aptroot); 

ascospores 15-17 × 8-10 µm; eastern North America .................. [G. hymnothora (Ach.) Aptroot] 

3. Ostiole apical; ascospores 16-21 × 11-13 µm; South Africa ............................................................... 

............................................................................................... [G. macrocarpoides (Zahlbr.) Aptroot] 

2. Ascomata compound, joined by fused necks with a single 

common ostiole; ascospores 16-19 × 7-9 µm; Bermuda 

and Brazil.................................................................................. [G. nigrescens (Müll. Arg.) R. C. Harris] 

1. Ascospores submuriform, ca. 4 × 1-2 irregularly septate, broadly elliptical, 

15-19 × 10-12 µm; ostiole lateral; microconida rod-shaped, 9-10 × 1 µm; 

Dominican Republic .............................................................................. [Granulopyrenis sp. Harris 19568] 

NOTES 

Granulopyrenis antillensis Aptroot 

This is maintained as distinct from G. hymnothora but I have not verified the ascus character used by 

Aptroot to separate them. The ascospore sizes overlap but those of 

G. antillensis seem slightly smaller. Microconidia are rod-shaped, 10-12 × 1 µm and were found in Seaver & 

Chardon 187 (NY) from Puerto Rico. Britton et al. 102 (NY) from St. Thomas cited by Aptroot as Distopyrenis 

americana is G. antillensis. 

Granulopyrenis hymnothora (Ach.) Aptroot 

Microconidia from the BM isotype are rod-shaped, 7-9 × 1 µm. 

Granulopyrenis macrocarpoides (Zahlbr.) Aptroot 

Microconidia from the holotype are 8-10 × 1 µm. 

Granulopyrenis nigrescens (Müll. Arg.) R. C. Harris, comb. nov. 

92

Tomasellia nigrescens Müll. Arg., Hedwigia 32: 135. 1893. Type. BRAZIL. Pará: Igarapé, Spruce, 

Lich. Amaz. And. 364 (G, isotype). 

Parapyrenis elongata Aptroot, Biblioth. Lichenol. 44: 100. 1991. Type. BERMUDA. Hall's Island, on 

Eugenia axillaris, 27 Aug-21 Sep 1912, Brown & Britton 885 (NY, holotype). 

Aptroot synonymized Tomasellia nigrescens with Granulopyrenis hymnothora but from my notes and 

drawings of the type (I have not re-examined it in 1995), it does not differ from Parapyrenis elongata. Study of 

the type of P. elongata shows a ± colorless inner layer of endospore and granules in the endospore typical of 

Granulopyrenis. 

Granulopyrenis sp. Harris 19568 

Although this species expands the circumscription of Granulopyrenis to include submuriform ascospores 

(the young 2-celled ascospores are identical in morphology to the mature ascsopores of the other taxa), the 

Dominican Republic collection is simply not ample enough for a type which would endure much in the way of 

future study. 

LITHOTHELIUM Müll. Arg. 

KEY TO LITHOTHELIUM IN FLORIDA 

1. Ascospores 17-20 × 7.5-9 µm; Duval County...................................................... L. illotum (Nyl.) Aptroot 

1. Ascospores 10-12 × 4-4.5 µm; Collier County............................................. L. microsporum R. C. Harris 

NOTES 

Lithothelium almbornii R. C. Harris & Aptroot 

A small collection from the Dominican Republic seems most closely related to 

L. almbornii previously known only from Africa. The Dominican collection differs in slightly broader ascospores 

and in having the upper part of the hymenium inspersed. Until better material is available it can be assigned 

here with a question mark. 

DOMINICAN REPUBLIC. Prov. La Romana: Isla Catalina, alt. 0-10 m, 18 Dec 1987, Zanoni et al. 37309[A] 

(NY). 

Lithothelium bahamense Riddle = Strigula bahamense 

Lithothelium concameratum (Stirton) Makhija et al. 

I have seen no material but the illustration in Makhija et al. (1994) suggests Anisomeridium. 

Lithothelium fugax Müll. Arg. 

Aptroot synonymized this taxon with L. obtectum (Müll. Arg.) Aptroot with pyrenastroid ascomata. 

However, the ascomata are fused as in L. cubanum (± astrothelioid) and the ascospores are larger, 16-18 × 7- 

8 µm. I would prefer to recognize it as a distinct species. It is only known from the African type collection. 

Lithothelium illotum (Nyl.) Aptroot 

Sphaeria (Metasphaeria) cavernosa Ellis & Everhart, J. Mycol. 1: 91. 1885. Type. GEORGIA. 

Darien, on bark of Taxodium distichum, Ravenel 703 (NY, holotype), syn. nov. 

The occurrence of Lithothelium illotum in North Florida is no surprise as it has been found in the 

Carolinas, Georgia, Louisiana and Mississippi. 

FLORIDA. Duval County: Livezey's [Creek], on Liriodendron, Calkins, NAL 196 (NY ex PAC). Additional 

record. CUBA. Camagüey Prov.: In the Cuitos Mts., midway between Camagüey and La Gloria, 19-21 Feb 

1941, Johnson s.n. (NY). 

Lithothelium microsporum R. C. Harris, sp. nov. 

93

Ascomatis singularibus ostiolis excentricis, hymenio non insperso et ascosporis minimis, hyalinis, 4locularibus, 

10-12 × 4-4.5 µm. 

Type. FLORIDA. Collier County: Fakahatchee Strand State Preserve, Royal Palm Hammock, 5.7 mi from S 

entrance on W. J. Janes Memorial Scenic Drive, hardwood-Taxodium swamp with Roystonea, on prop roots 

of Roystonea, 6 Dec 1992, Buck 22741 (NY, holotype). 

Thallus greenish gray, matte, Trentepohlia present. Ascomata solitary, not compound, orbicular to 

obovoid, ca. ½-_ immersed, 0.4-0.5 × 0.3-0.4 mm. Ostiole eccentric. Hymenium not inspersed. Hymenial gel 

IKI+ dirty pinkish orange. Asci cylindrical with ascospores uniseriate to subbiseriate. Ascospores colorless, 

ovoid, 4-celled, 10-12 × 4-4.5 µm. 

Lithothelium microsporum has the smallest ascospores in the genus. Closest in ascospore size is L. 

obtectum that has compound, pyrenastroid ascomata. I do not exclude the possibility that L. microscopicum is 

a form of L. obtectum behaving oddly at the edge of its range. Description at the specific level focuses further 

attention on the question. It is known only from the type collection. 

Lithothelium obtectum (Müll. Arg.) Aptroot 

The range of this species is extended to Venezuela. 

VENEZUELA. Amazonas: Dpto. Río Negro, along Río Mawarinuma, just outside Cañon Grande, vicinity of 

Neblina base camp, ca. 140 m, 00°50'N, 66°10'W, low primary forest on white sand, Jan 1985, Buck 12556 

(NY, VEN). 

Pleurotrema polysemum Nyl. (Lithothelium polysemum (Nyl.) Aptroot) 

This species has been an embarassment to many of us. I placed Pleurotrema in the Trypetheliaceae. 

Aptroot, in agreement with Eriksson (1981), removed it from that family but placement in the Pyrenulaceae is 

no improvement. I had the good fortune to have Sabine Huhndorf as a collegue for several years and she was 

working on Saccardoella. Several taxa described in lichen-forming genera have proven to belong to 

Saccardoella, a unitunicate, non-lichenized genus. Being marginally involved in her study, it struck me one 

day that the ascus of Pleurotrema was very similar to that of Saccardoella. Both she and Margaret Barr 

agreed and Barr has accepted Pleurotrema and the Pleurotremaceae in the unitunicates related to the 

Amphisphaeriaceae (Mycotaxon 51: 211. 1994) and "pure" lichenologists no longer have to worry about it. I 

feel this is a prime example of the sort of broad collaboration that will be needed to elaborate a well founded 

system for the ascomycetes. 

MYCOPYRENULA Vainio 

In 1989 I synonymized this genus with Pyrenula. It has typical Pyrenula type microconidia and 

ascospores, however its niche as a saprophytic? colonizer of smooth bark, rather short physes and ascus with 

a thin, ± uniform wall seem to set it apart. Further, as the collective genus Pyrenula will probably be 

dismantled into more natural groups, it does no harm to anticipate this event. There is only a single species, 

M. coryli (Massal.) Vainio. It is not known from North America. 

PARAPYRENIS Aptroot 

My first intuitive thought was to simply synonymize Parapyrenis with Distopyrenis. However, the type 

species P. aurora (Zahlbr.) Aptroot, is distinctive in asci with a uniformly thin wall and very short or no stipe, 

ascospores with a very strongly pigmented median plate and euseptum?, hard, strongly melanized ascomatal 

shell and Asian distribution. Pending further studies, I consider Parapyrenis to be monotypic. Parapyrenis 

elongata Aptroot is a Granulopyrenis. I have not seen material of the other two original species. From the 

94

illustration and distribution, P. conica Aptroot, may possibly remain in Parapyrenis. Parapyrenis guayaci (Fée) 

Aptroot with massive (for this group) ascospores seems of uncertain position but possibly is referable to 

Distopyrenis. Aptroot (Nova Hedwigia 60: 325-379. 1995) has described three additional species. There are 

two isotypes at NY of P. maclurae (Saulescu & Sandu) Aptroot from Romania, unfortunately neither containing 

the Parapyrenis. From the distribution and substrate it is unlikely to belong to the Pyrenulaceae. I have not 

seen material of P. maritima Aptroot and P. tecomatis (Berkeley & Curtis) Aptroot. From the illustrations, the 

former may be a Granulopyrenis (distinct layer shown around lumina), the latter, said to have a spinulose wall, 

that would be anomalous in the Pyrenulaceae, is of uncertain position. Perhaps, the "spines" are endospore 

granules and it is also a Granulopyrenis. 

PYRENOGRAPHA Aptroot 

This genus is a problem. Aptroot described the ascospores as 3-septate. Sabine Huhndorf, thinking she 

possibly had this from the West Indies, examined the isotype at NY and found that the ascomata matched the 

original description but the ascospores were submuriform. Investigation of the holotype (CBG) and another 

isotype (B) gave identical results. Further, some time previously I had examined isotype material of 

Phaeopeltosphaeria irregularis Wehmeyer. It is the same as all of the available material of Pyrenographa 

xylographoides Aptroot. Did Aptroot see only immature ascospores or is there another taxon with 3-septate 

ascospores (perhaps an uncited duplicate in his personal herbarium)? I have not seen material of the type 

species of Phaeopeltosphaeria, P. caudata Berlese & Peglion from Italy, which from the description does not 

seem likely to be congeneric with Pyrenographa, but this problem must await further study. Based on the 

designated holotype and the two isotypes of Pyrenographa xylographoides, a new combination is required, 

Pyrenographa irregularis (Wehmeyer) R. C. Harris, comb. nov. Phaeopeltosphaeria irregularis Wehmeyer 

in Martin, Pacific Science 2: 74. 1948. Type. ECUADOR. Galapagos: South Seymour Island, on dead, 

decorticated wood of Bursera graveolens, 6 Sep 1945, Martin 6251 (NY, isotype). Even if Aptroot can come 

up with material which matches his description, it is my understanding that the designated holotype can only 

be superseded through conservation. In the unlikely event that this should happen, the status of the genus 

and the disposition of P. irregularis will have to be reassessed. In addition to the Australian and Ecuadoran 

material, NY holds specimens determined by Wehmeyer from the Marshall Islands and the Tuamotu 

Archipelago. 

Aptroot emphasized the elongate ascoma but I feel this is merely a consequence of the substrate, bare, 

eroded wood. Species of Anisomeridium, Arthopyrenia and Julella growing on old wood show this same 

character of the clypeus elongated in the direction of the wood fibers. Retention of this genus is supported by 

the rather poorly developed endospore. Young ascospores apparently do not pass through a "Rinodina-like" 2celled 

stage resembling the mature stage of other taxa of Distopyrenis. Distopyrenis submuriformis which 

otherwise has similar ascomata and ascospores does have a "Rinodina-like" stage. The microconidia present 

in the NY isotype of Pyrenographa xylographoides, rod-shaped, 12-15 × 1 µm, are somewhat longer than 

those in the rest of the Pyrenulaceae with coarse physes. 

PYRENULA Ach. 

1. Ascospores muriform ................................................................................................................................ 2 

2. Ascospores 8/ascus (occasionally fewer due to abortion), under 75 µm long ...................................... 3 

3. Thallus some shade of red, orange or yellow (occasionally only over ascomata), 

KOH+ purple...................................................................................................................................... 4 

4. Thallus orange to yellow................................................................................................................ 5 

5. Ascospores with 4 rows of 4 locelli each, 13-23 × 9-13 µm; 

Seminole County south ....................................................P. ochraceoflava (Nyl.) R. C. Harris 

95

5. Ascospores with 6-8 rows of 4-8 locelli each, 

23-35 × 11-17 µm; peninsular Florida ..........................P. ochraceoflavens (Nyl.) R. C. Harris 

4. Thallus red; hymenium inspersed; ascospores 30-42 × 15-20 µm; 

Dade and Monroe counties ................................................ P. cruentata (Müll. Arg.) R. C. Harris 

3. Thallus not red, orange or yellow, KOH- ........................................................................................... 6 

6. Ascomata fused in groups by their eccentric or lateral ostioles 

(pyrenastroid), or pseudostromatic shells fused completely 

(melanothecoid)............................................................................................................................. 7 

7. Pseudostromata pyrenastroid; pseudostromatal shells not 

completely fused into a flat Arthonia like plate ........................................................................ 8 

8. Ascospores over 45 × 18 µm; ascomata often only at an angle, 

sometimes solitary............................................................................................................... 9 

9. Ascospores 45-65 × 18-27 µm; throughout Florida..............................P. ravenelii Tuck. 

9. Ascospores 65-85 × 25-32 µm (60-72 × 25-30 µm in 

Florida specimen); Columbia County ......................................................Pyrenula sp. 47 

8. Ascospores 33-42 × 13-18 µm; ascomata usually 

distinctly recumbent; Dade, Monroe and Sarasota 

counties .............................................................................. P. astroidea (Fée) R. C. Harris 

7. Pseudostromata melanothecoid, looking much like an Arthonia 

except for the central ostiole; margins crenulate; ascomata recumbent, 

fused at the single ostiole; ascospores with 8 rows of locelli, 

33-40 × 12-14 µm; Key Largo, Monroe County .............................................Pyrenula sp. 208a 

6. Ascomata solitary, ostioles apical ............................................................................................... 10 

10. Thallus UV-............................................................................................................................ 11 

11. Postmature ascospores empty, collapsing...................................................................... 12 

12. Ascospores over 45 × 16 µm .................................................................................... 13 

13. Locelli smaller, spherical, in 8-10 rows of up to 10 each; 

ascospores 45-62 × 16-30 µm ............................................................................ 14 

14. Ascospores (45-) 50-65 × (18-) 20-26 µm, relatively 

narrow with pointed ends; Columbia and Volusia 

counties ....................................................... P. mucosa (Vainio) R. C. Harris 

14. Ascospores 50-70 × 24 -30 µm, broad with rounded end; 

known only from Mississippi............................................................................. . 

............................................................... [P. pyrenuloides (Mont.) R. C. Harris] 

13. Locelli large, often angular, in 8 rows of up to ca. 6 each; 

ascospores 45-60 × 16-22(-25) µm; very common, 

throughout the state ..............................................................P. leucostoma Ach. 

96

12. Ascospores 30-42(-53) × 11-15 µm; south to Sarasota 

County ...................................................................................... P. thelomorpha Tuck. 

11. Postmature ascospores filled with red or colorless oil substance................................... 15 

15. Oily substance red; ascospores 40-53 × 17-22 µm; 

Dade and Monroe counties ..................................P. macularis (Zahlbr.) R. C. Harris 

15. Oily substance colorless; ascospores 37-51 × 14-19 µm; 

Keys..........................................................................................P. oleosa R. C. Harris 

10. Thallus white, UV+ yellow (lichexanthone); ascospores 14-23 × 8-13 µm; 

Dade, Monroe and Pinellas counties........................................ P. confinis (Nyl.) R. C. Harris 

2. Ascospores (2-)4(-6)/ascus, 100-150(-190) × 30-41(-48) µm; ostioles lateral, 

sometimes several fused; south to Highlands County ....................... P. falsaria (Zahlbr.) R. C. Harris 

1. Ascospores transversely septate or rarely with a few cells longitudinally divided ................................... 16 

16. Ostiole lateral or lateral and several fused to form compound ascomata......................................... 17 

17. Ascospores with terminal lumina elongate and directly against exospore ................................. 18 

18. Ascomata solitary................................................................................................................. 19 

19. Ascospores less than 35 µm long ................................................................................... 20 

20. Ascospores 25-32 × 10-13 µm; Dade and Monroe 

counties ..............................................................P. cuyabensis (Malme) R. C. Harris 

20. Ascospores 18-22(-25) × 7-9 µm; Dade and Monroe 

counties ......................................................... P. subferruginea (Malme) R. C. Harris 

19. Ascospores 45-65 × 20-25 µm, often with 2-3 cells longitudinally 

sudivided; Dade County ............................................................P. erumpens R. C. Harris 

18. Several ascomata joined by fused ostioles; ascospores 

17-21 × 7-7.5 µm; Florida, without locality, 1878, Austin.......................................................... 

......................................................................................... P. personata (Malme) R. C. Harris 

17. Ascospores with terminal lumina separated from exospore 

by a layer of endospore.............................................................................................................. 21 

21. Ostiole not red ringed ........................................................................................................... 22 

22. Several ascomata joined by fused ostioles .................................................................... 23 

23. Ascospores 21-25 × 8-10 µm; numerous crystals around ostiole; 

Collier, Dade and Monroe counties .....................P. cubana (Müll. Arg.) R. C. Harris 

23. Ascospores 16-21 × 6-8 µm; crystals absent; 

throughout Florida .......................................... P. septicollaris (Eschw.) R. C. Harris 

22. Ascomata solitary........................................................................................................... 24 

24. Hymenium not inspersed.......................................................................................... 25 

97

25. Ascospores 18-25 × 8-10 µm; hymenial gel IKI+ blue-green 

or IKI-; throughout Florida ..........................................P. microtheca R. C. Harris 

25. Ascospores 37-45 × 15-22 µm; hymenial gel IKI+ orangish; 

Dade and Monroe counties .........................P. martinicana (Vainio) R. C. Harris 

24. Hymenium inspersed, gel IKI+ orangish; ascospores 13-17 × 

5-6 µm; Polk County...............................................................P. wheeleri R. C. Harris 

21. Ostiole surrounded by bright red ring; hymenium inspersed to top of asci; 

ascospores dark brown with darker bands between cells, 18-24 × 10-14 µm; 

Georgia, not yet found from Florida................................................. [P. wetmorei R. C. Harris] 

16. Ostiole apical, erect (pseudostromatic shells may be laterally fused 

to form extensive pseudostromata in P. anomala)............................................................................. 26 

26. Ascospores elliptical or fusiform, not tapering to a tail .................................................................. 27 

27. Ascospores over 21 µm long (if thallus ecorticate see P. microcarpa) ................................... 28 

28. Thallus orange or red at least near ascomata................................................................... 29 

29. Thallus orange, pseudocyphellate; ascospores 26-42 × 12-15 µm; 

Dade, Monroe and Sarasota counties.................................................P. cerina Eschw. 

29. Thallus red (in shade forms pigmented only near ascomata); 

hymenium heavily inspersed; ascospores 27-35 × 13-17 µm; 

very common, throughout the state.......................................P. cruenta (Mont.) Vainio 

28. Thallus greenish when fresh, whitish to brown in the herbarium ...................................... 30 

30. Postmature ascospores with red or colorless oily substance 31 

31. Oily substance red; ascospores 4- or 6-celled, 

28-34 × 12-15 µm; very common, throughout the 

state.................................................................P. concatervans (Nyl.) R. C. Harris 

31. Oily substance colorless; ascospores 4-celled, 

28-35(-40) × 10-15(-17) µm; throughout the 

state.........................................................................................P. quassiaecola Fée 

30. Postmature ascospores empty, collapsing................................................................... 32 

32. Ascospores with rounded ends; hymenium not inspersed; 

hymenial gel IKI+ orange....................................................................................... 33 

33. Ascospores reaching 45 × 18 µm..................................................................... 34 

34. Thallus UV-; ascospores 27-38 × 12-18 µm; 

gelatinous sheath lacking dark terminal caps; 

not common, south to Sarasota County......................................................... 

...........................................................................P. punctella (Nyl.) Trevisan 

34. Thallus UV+ yellow (lichexanthone) or UV-; ascospores 

36-45 × 12-17 µm; ascospore sheath gelatinous with 

dark terminal caps; Holmes County ...........................P. caryae R. C. Harris 

98

33. Ascospores 44-50(-56) × 17-24 µm; on Ficus; 

Collier County........................................................................Pyrenula sp. 29916 

32. Ascospores apiculate or at least pointed at one or both ends, 

rather dark brown when mature, 22-29 × 9-11 µm; hymenium 

inspersed only in a layer just below ostiole; hymenial gel IKI-; 

Collier County.......................................................................P. acutalis R. C. Harris 

27. Ascospores less than 21 µm long (to 25 µm in P. microcarpa)............................................... 35 

35. Ascospores with terminal lumina directly against exospore; 

hymenial gel IKI+ blue-green............................................................................................. 36 

36. Hymenium inspersed................................................................................................... 37 

37. Thallus UV-; ascomata 0.3-0.6 mm across; ascospores 

often citriform, 14-18 × 8-10 µm; south to Lake County............................................ 

........................................................................................P. citriformis R. C. Harris 

37. Thallus UV+ yellow (lichexanthone); ascospores 

13-22 × 8-11 µm; northern Florida............................................................................. 

.................................................................P. pseudobufonia (Rehm) R. C. Harris 

36. Hymenium not inspersed............................................................................................. 38 

38. Thallus UV+ yellow (lichexanthone); pseudostromatal 

wall lacking crystals; ascospores 18-21 × 10-13 µm; 

Dade and Monroe counties ....................................................P. cocoes Müll. Arg. 

38. Thallus UV-; pseudostromatal wall containing 

colorless crystals; ascospores 18-20 × 11-12 µm; 

Dade County.................................................. P. nitidula (Bresadola) R. C. Harris 

35. Terminal lumina separated from exospore by an obvious layer 

of endospore .................................................................................................................... 39 

39. Ascomata fused laterally to form an extensive pseudostromata 

(melanothecoid).......................................................................................................... 40 

40. Ascospores more or less uniformly colored; thallus usually 

tan to yellow brown.............................................................................................. 41 

41. Ascospores with thin endospore, 17-21 × 7-9 µm......................................... 42 

42. Pseudostromata raised, not forming extensive 

network,brown, with crowded ascomata; 

common, throughout the state ......................... P. anomala (Ach.) Vainio 

42. Pseudostromata flush with thallus, forming 

an extensive network, shiny black, with 

scattered ascomata; Dade and Monroe 

counties .........................................................................Pyrenula sp. 415 

41. Ascospores with thicker endospore, 

99

18-21 × 8-11 µm; Dade County...........................................Pyrenula sp. 6121 

40. Ascospores with a distinct dark brown median plate or 

sometimes in old spores plates between all four lumina, 

18-20 × 7-8 µm; thallus brown to blackish brown; on Ilex; 

Liberty County ............................................................P. atrolaminata R. C. Harris 

39. Ascomata solitary....................................................................................................... 43 

43. Thallus corticate ................................................................................................... 44 

44. Ascomata large, 0.7-1.5 mm across, flattened conical ................................. 45 

45. Hymenium not inspersed; ascomata with a thin 

translucent thalline covering................................................................... 46 

46. Ascospores 15-19 × 6-8 µm; south to 

Lake County .........................................P. santensis (Nyl.) Müll. Arg. 

46. Ascospores 12.5-15.5 × 4.5-5 µm; 

Collier County .......................................................Pyrenula sp. 30313 

45. Hymenium heavily inspersed; ascospores 

15.5-21 × 5.5-8 µm; ascomata naked, shiny; 

very common, throughout the state............................................................ 

................................................................ P. mamillana (Ach.) Trevisan 

44. Ascomata smaller, to 0.7 µm......................................................................... 47 

47. Endospore developed ............................................................................. 48 

48. Endospore uniform........................................................................... 49 

49. Hymenium inspersed; ascospores 

13-15 × 4.5-6 µm; Taylor to Polk County........................................ 

.......................................................................P. laetior Müll. Arg. 

49. Hymenium not inspersed; ascospores 

13-16 × 4.5-6 µm; throughout the state.......................................... 

.................................................................P. aspistea (Ach.) Ach. 

48. Endospore with a distinct, additional layer around 

lumina or older ascospores with darker bands 

between the lumina .......................................................................... 50 

50. Endospore with additional layer around lumina; 

ascospores 17-19 × 7.5-9 µm; ostiole usually 

red pigmented, KOH+ blue; Columbia, Dade 

and Seminole counties ................................................................... 

........................................................ P. rubrostoma R. C. Harris 

50. Endospore with darker bands between the 

lumina; ascospores 15-19 × 6-7.5 µm; 

ostiole not pigmented; Sarasota County ........................................ 

........................................................ P. confoederata R. C. Harris 

100

47. Endospore very reduced; ascospores 14-17 × 

5-7 µm; Dade County ....................................... P. tenuisepta R. C. Harris 

43. Thallus ecorticate, white; ascospores 19-25 × 8-12 µm; 

hymenial gel IKI-; throughout the state...........................P. microcarpa Müll. Arg. 

26. Ascospores fusiform-clavate, tapering to a long tail, in a single 

bundle in the ascus, 7-10-celled, 60-70 × 4.5-5 µm; asci markedly 

clavate; hymenium inspersed, IKI-; thallus UV-, not pseudocyphellate; 

on shrub at edge of salt marsh, Collier County ................................................Pyrenula sp. 30018 

Pyrenula atrolaminata R. C. Harris, sp. nov. 

NOTES 

Subsimilis P. anomalae sed ascosporis lamina media atra vel vetustis omnibus laminis atris, 18-20 × 7-8 

µm et thallo castaneo atranti. 

Type. FLORIDA. Liberty County: Whitehead Lake Campground, Apalachicola National Forest, cypress swamp 

along Ochlockonee River, on Ilex, 11 Apr 1967, Harris 1454 (NY, holotype; MSC, isotype). 

Thallus chestnut brown initially but soon blackening to blackish brown, corticate, not pseudocyphellate, 

UV-. Ascomata partly solitary, hemispherical, ca. 0.5 mm diam. but mostly laterally aggregated into slightly 

raised, blackish pseudostromata containing several to ca. 30 ascomata, initially appearing pruinose due to 

disrupted bark cells; ostiole dark; crystals lacking. Hymenium not inspersed; hymenial gel IKI-. Ascospores 

subbiseriate, 4-celled, 18-20 × 7-8 µm, with a dark brown median plate and occasionally in older ascospores 

all three septa with brown plates. 

The epithet is given for the ascospores with dark layers between lumina but could also be applied to the 

dark thallus. I am unaware of any other species in the genus with this type of ascospore. Externally one 

would think this to be a dark thallus form of Pyrenula anomala. It is known from only a single site in the 

Apalachicola region, notorious for its high rate of endemism. 

Additional specimen. As type, Harris 1438 (NY). 

Pyrenula caryae R. C. Harris, sp. nov. 

Similissima P. punctellae sed thallo ceraceo saepe lichexanthone continente, ascosporis operculis 

brunneis et majoribus, (30-)36-45(-50) × 12-17(-21) µm et saepe ad Caryam crescente. 

Type. MISSOURI. Iron County: St. Francis Mountains, Clark National Forest, along Co. Rd. N just N of 

Reynolds Co. line, ca. 335 m, 37°40'N, 90°47'W, at 2 m height on a 6" diam. Carya, 13 Oct 1993, Harris 

31124 (holotype NY; isotypes H, CBG, MOR). 

Thallus usually yellowish or yellowish tan but occasionally green, corticate, somewhat shiny, usually with 

a waxy look, pseudocyphellate, UV+ yellow (lichexanthone) or less often UV-. Ascomata flask-shaped, 0.5-0.8 

mm diam., immersed, often entirely covered by thallus with only the pale ostiole visible, in some (with age?) 

the black ascomatal apex exposed; numerous colorless crystals in upper ascomatal wall. Hymenium not 

101

inspersed; hymenial gel IKI-. Ascospores subbiseriate to uniseriate, 4-celled, (30-)36-45(-50) × 12-17(-21) 

µm, with a darker pigmented median band and with a brownish apical cap at both ends (outside the exospore) 

which in fresh material is often reflexed looking like a little sombrero, gelatinous sheath otherwise not readily 

evident. 

Harris (1989) cited some of the collections now recognized as P. caryae under P. punctella (Nyl.) 

Trevisan where it was noted "The taxonomy of P. punctella requires further work...". Pyrenula caryae was 

initially segregated when the presence of lichexanthone was noticed. However, closer study revealed several 

correlated characters, the "waxy" thallus, larger ascospores with the very odd apical caps and a strong 

tendency to grow on Carya. Eleven of the fifteen known collections are from Carya and the species has been 

named for this preference. The substrate of the other four is neither noted on the label nor readily 

determinable. The ascospores of P. punctella are 27-38 × 12-18 µm and lack the apical caps but may have 

an apical dark spot in the exospore or are weakly apiculate with the apiculus often darkening. Also older 

ascospores tend to have a paler zone around the locules not seen in P. caryae. Pyrenula punctella is still 

probably not homogeneous. The Coastal Plain collections, mostly on Fraxinus or Magnolia, have a reddish 

almost endophloeodal thallus, interior collections, mostly on Fagus, have a whitish or grayish dull epiphloeodal 

thallus but I am not as yet prepared to separate them at the species level in a group where the intimate 

relation between lichen and substrate give rise to great differences in aspect. Three collections of P. caryae 

seem to lack lichexanthone entirely. In fact one of the three collections made at the type locality lacks 

lichexanthone. Overall it seems present only in low quantities as the fluorescence is quite weak. Presence or 

absence of lichexanthone is sometimes recognized at the specific rank but the unique ascospores allow one 

to dismiss such a treatment here. The known distribution of P. caryae may also be distinctive, running in an 

arc around the Southern Appalachians from North Carolina to Missouri mainly on the Piedmont although like 

P. caryae several also extend into northern Florida. The elevations noted are between 35 m and 900 m. 

Gomphillus americanus also fits this pattern and also possibly Cladonia caroliniana Tuck. s. lat. (if usnic acid 

only strain is included) and Cresponea premnea (Ach.) Egea & Torrente var. saxicola (Leighton) Egea & 

Torrente. 

Additional specimens. ALABAMA. Cleburne County: 2 mi NE of Heflin, 23 Oct 1964, Johnson 

(NY);Tuscaloosa County: near Rickey, 24 Oct 1964, Johnson (NY). ARKANSAS. Washington/Crawford County: 

Devil's Den State Park, 9 Jun 1961, Johnson (NY). FLORIDA. Holmes County: along St. Rd. 2 just E of 

Choctawhatchee River, 10 Dec 1993, Buck 24686 (NY). GEORGIA. Towns/Rabun County: Chattahooche 

National Forest, along Appalachian Trail between Dick's Creek Gap and Little Bald Knob, 825-900 m, 21 Sep 

1992, Harris 28115 (NY). MISSISSIPPI. Itawamba County: Donivan Slough, Mile 283.3 along Natchez Trace, c. 

35 m, 28 Sep 1992, Harris 28633 (NY); Lee County: near Tupelo, 31 Dec 1952, Johnson 1041A, 1046B (NY); 

Pearl River County: 1 mi S of Carriere, 14 Jun 1939, Johnson 1118 (NY). MISSOURI. Carter County: Bluff on E 

side of Current River across from Big Spring S of Van Buren, 23 Sep 1990, Harris 25633 (NY); Iron County: as 

type, Harris 31190, 31204 (NY). NORTH CAROLINA. Graham County: Horse Cove Campground, Nantahala 

National Forest, 3 Oct 1987, Buck 15387 (NY); Haywood County: Great Somky Mtns. Natl. Park, Big Creek, 

45 mi ESE of Knoxville, 2000-2300 ft, 25 Aug 1977, Wetmore 29850 (MIN); Macon County: McDowell 

Mountain, Nantahala National Forest, 640-945 m, oak-hickory woods, 12 Aug 1994, Harris 33192 (NY). 

SOUTH CAROLINA. Chester County: Chester, [Green?] (NY). 

Pyrenula concastroma R. C. Harris, sp. nov. 

Characteribus internis ut P. concatervanti sed ascomatis inclusis in pseudostromatis trypethelioidibus. 

Type. PUERTO RICO. Salinas de Guánica, 27 Dec 1930, Britton & Britton 9602 (NY, holotype). 

Thallus grayish brown, corticate, not pseudocyphellate, UV-. Pseudostromata sessile, constricted at 

base, mostly oval, ca. 2 × 1 mm, occasionally fused into larger groups, brown, filled with large oxalate(?) 

crystals and a whitish powder bleeding yellow in KOH. Ascomata many/pseudostroma with apical ostioles. 

Hymenium not inspersed. Hymenial gel IKI+ orangish. Asci with 8 biseriate ascospores. Ascospores 4- 

102

celled, becoming filled with reddish oily substance as contents disintegrate, 31-40 × 15-16µm. 

The epithet is derived from the bastardization of the epithet of the sister species "concatervans" and 

"stroma" as the ascospores, etc., are identical to those of P. concatervans and the ascomata are included in 

pseudostromata. This assumes that ascospores types are conservative and define clades although there is 

no independent proof for this. In previous generations, P. concastroma would have been placed in 

Melanotheca, a genus I have abandoned as polyphyletic, containing taxa derived independently in several 

ascospore defined clades. Pyrenula concastroma is virtually identical to P. wrightii which differs in having a 

colorless oily substance in postmature ascospores (as P. quassiaecola differs from P. concatervans). This 

raises the question, does the color of the oily substance mean anything? Have two taxa, differing mainly in the 

color of the contents of postmature ascospores, evolved identical, highly unusual for the Pyrenulaceae, 

pseudostromata? This would seem to be a question which the friendly neighborhood molecular biologist is 

best equipped to answer. Pyrenula concastroma is known only from the type. 

Pyrenula concatervans (Nyl.) R. C. Harris 

Pyrenula crystalligera H. Magn., Ark. Bot. n.s. 3(3-4): 239. 1956. Type. [HAWAIIAN ISLANDS.] Iles 

Sandwich. Voyage de M. J. Remy, 1851-1855. Herb. Mus. Paris, misit cel. Nylander, 1860 (FH- 

Tuck 4107, holotype), syn. nov. 

Pyrenula confinis (Nyl.) R. C. Harris, comb. nov. 

Verrucaria confinis Nyl., Ann. Sci. Nat. Bot. sér. 4, 3: 174. 1855. Anthracothecium confine (Nyl.) 

Müll. Arg., Linnaea 63: 45. 1880. Type. CHILE. Ad corticem eadem ac Glyphis cicatricosa (H-Nyl 


Verrucaria albescens Nyl., Expos. syn. pyrenoc. 50. 1858. Anthracothecium albescens (Nyl.) Müll. 

Arg., Linnaea 64: 45. 1880. Type. GUYANA BRITANNICA. Ad cort. Spondias lutea L. (H-Nyl 1285, 

holotype). 

Anthracothecium corticatum Müll. Arg., Bull. Soc. Roy. Bot. Belgique 30: 96. 1891. Pyrenula 

corticata (Müll. Arg.) R. C. Harris, Mem. New York Bot. Gard. 49: 89. 1989. Type. COSTA RICA. 

San José, 5300 ft., 1890, Pittier (MSC, holotype fragment ex G). 

Harris (1989) declined to take up the epithet confinis as the holotype had not been located. Singh (1985) 

incorrectly thought a specimen from French Guiana to be the type. In 1994 I located the holotype in Helsinki. 

Nylander had apparently intended to name the species Verrucaria concinna but never published this name 

(possibly due to the existence of Verrucaria concinna Borrer). Nylander had not changed the name on the 

specimen and it was quite naturally filed under this name in his herbarium. I have now studied the specimen 

and it agrees with the taxon previously treated by me as Pyrenula corticata. 

Pyrenula cruenta (Mont.) Vainio 

Verrucaria circumrubens Nyl., Flora 50: 196. 1867, nom. nud., Syn. lich. Nov. Caled. 90. 1868. 

Pyrenula circumrubens (Nyl.) B. de Lesd., Bull. Soc. Bot. France 62: 463. 1910. Type. [NEW 

CALEDONIA. Loyalty Islands:] Lifu, 1864 Thiebaut (H-Nyl 1418, holotype), syn. nov. 

Pyrenula rubromaculata Vainio, Bot. Mag. Tokyo 35: 77. 1921. Lectotype (selected here). JAPAN. 

Prov. Kii: on Taonabo japonica, 30 Dec 1911, Yasuda 259 (Tur-Vainio 31242). Syntype. as 

lectotype, Yasuda 262 (TUR-Vainio 31243), syn. nov. 

Pyrenula laetior Müll. Arg. 

Bot. Jahrb. Syst. 6: 413. 1885. Type. CUBA. Wright, Verr. Cub. 51 (FH-Tuck 4108, isotype). 

New to North America. FLORIDA. Citrus County: Mason Creek, ca. 2 mi SW of Homossasa, coastal 

hammock, elev. 2 m, on trunk of Ilex cassine, 10 May 1994, Griffin 2944 (FLAS, NY); Polk County: Nalcrest, 

on fallen oak branch, 11 Mar 1989, Wheeler (NY); Taylor County: Aucilla Wildlife Management Area, Aucilla 

Sinks, S of Goose Pasture Road along Florida Trail, ca. 4 mi N of U.S. Hwy. 98, on young Carya, 12 Dec 

1993, Harris 32285 (NY). 

The ascospores and asci resemble those of Pyrenula aspistea (Ach.) Ach. which differs in having the 

hymenium not inspersed. Pyrenula laetior is also close to Pyrenula velata Müll. Arg. from Brazil which differs 

103

in IKI+ blue hymenium and larger ascospores (_ = 16.4 × 6.1 µm). The Cuban species Pyrenula gregantula 

Müll. Arg. may be synonymous but is tentatively separated by slightly broader ascospores (_ = 6.0 µm) and 

distinctly papillate ascomata. Pyrenula laetior was included in Some Florida Lichens as sp. 11-3-89. 

Pyrenula laevigata (Pers.) Arnold 

A new character for this species has been brought to my attention by J. Etayo. Addition of KOH to a 

section of the hymenium gives a fleeting sky-blue reaction. As far as I know this is unique. 

Pyrenula macularis (Zahlbr.) R. C. Harris 

Bottaria albidopallens Vainio, Ann. Acad. Sci. Fenn., ser. A, 15(6): 328. 1921. Anthracothecium 

albidopallens (Vainio) Zahlbr., Cat. lich. univ. 8: 121. 1931. Type. PHILIPPINE ISLANDS. Luzon: 

Manila, Dec 1912, Merrill, Bur. Sci. 8495 (F, isotype), syn. nov. 

Although Vainio's name is older I refrain from taking it up as there are other possible synonyms older yet 

whose types have not been examined. There is no point in changing the name several times. Also the type of 

Verrucaria analepta var. americana Ach., Anthracothecium americanum (Ach.) Müll. Arg. belongs here. The 

epithet cannot be taken up in Pyrenula due to the existence of Pyrenula americana (Massal.) Trevisan [= 

Ditremis americana (Massal.) R. C. Harris]. This is just as well as the species is by no means strictly 

American. Details of the typification will be dealt with elsewhere. 

Pyrenula mamillana (Ach.) Trevisan 

Pyrenula marginata Hook. in Kunth, Syn. pl. 20. 1822, syn. nov. 

As I have long suspected, but not been able to prove due to the unavailability of the type, the oldest name 

for the species treated in Harris (1989) as P. marginata is indeed P. mamillana. The typification, etc., will be 

dealt with elsewhere. 

Pyrenula mucosa (Vainio) R. C. Harris, comb. nov. 

Bottaria (Anthracothecium) mucosa Vainio, Hedwigia 38: (257). 1899. Type. GUADELOUPE. 

Mouteran, 250 m, sur un vieux pied d'Inga laurifolia, Duss 501 (TUR-Vainio 30963, holotype) 

[mixed with Pyrenula nitidula (Bresadola) R. C. Harris]. 

This seems to be a member of a complex of taxa which differ mainly in ascospore size. The ascospores 

of this complex seem to shrivel with age, not developing either reddish or colorless oily material as in some 

otherwise similar species. Pyrenula pyrenuloides has larger, mostly blunt ascospores. Also similar is 

Pyrenula dissimulans (Müll. Arg.) R. C. Harris, comb. nov., Pleurothelium dissimulans Müll. Arg., Bot. 

Jahrb. Syst. 6: 387. 1885. Type. CUBA. Wright, Verr. Cub. 213 (G, holotype; NY, isotype). Synonym: Bottaria 

libricola f. nana Vainio, Mycologia 21: 39. 1929. PUERTO RICO. on open, dry hilltop near Yauco, 30 Dec 1915, 

Fink 1669 (MICH, NY, isotypes). Pyrenula dissimulans was referred to Pleurothelium as a few of the ostioles 

are slighty eccentric. Basically it seems to have upright ostioles. The ascospores are slightly smaller and 

broader in proportion (35-50 × 17-22 µm) than those of P. mucosa. Also the locelli are mostly larger and fewer 

in number. Pyrenula dissimulans has not yet been found in North America. However, to complicate matters 

even further there are two collections from Florida (Dade and Union counties) with rather small ascospores but 

not the spore type of P. dissimulans. I tentatively refer them to P. mucosa. There is also a "wild card" name 

whose type I have not seen, Pyrenula libricola Fée. It is readily apparent that further work is needed and that 

the situation is unstable. 

Pyrenula nitidula (Bresadola) R. C. Harris ined. 

The combination will be made elsewhere. The type is from Puerto Rico. It is also known from 

Guadeloupe (type of Pyrenula subsimplex Vainio) and French Guiana. 

New to North America. FLORIDA. Dade County: Deering Hammock, Cutler, on live oak, 15 Dec 1919, N. 

L. & E. G. Britton 724 (NY). 

Pyrenula occidentalis (R. C. Harris) R. C. Harris 

Pyrenula harrisii Hafellner & Kalb, Herzogia 9: 85. 1992. Type. MADEIRA. An der Strasse zwischen 

Ponta do Pargo und Porto Moniz, bei Serrado. An Alleebäumen (Acer pseudoplatanus). 530 m. 

104

5.VIII.1990, Kalb 23189, not seen, syn. nov. 

Although this has nothing to do with Florida, it gives me some pleasure to dispose of this name as I do 

not feel any need of advertising. Pyrenula occidentalis, not occurring in eastern North America, was not 

included in Harris (1989) which seems to have been Hafellner and Kalb's source of information. The species 

is oceanic occurring in the Pacific Nortwest, Great Britain, Scandinavia and South Africa. Its occurrence in 

Madeira is not the least unexpected. 

Pyrenula punctella (Nyl.) Trevisan 

Pyrenula ferax Müll. Arg., Bot. Jahrb. Syst. 6: 414. 1885. Type. CUBA. Wright, Verr. Cub. 81, 82 

(FH-Tuck 4108, isosyntypes), syn. nov. 

Pyrenula pyrenuloides (Mont.) R. C. Harris 

Harris (1989) tentatively reported this taxon from several southern states noting that the ascospores were 

smaller than those of the type and other specimens from French Guiana. Having re-examined the type of 

Bottaria mucosa Vainio, I now prefer to refer the smaller spored collections to this species. Only one 

collection from Mississippi seems to have the very broad, stumpy ascospores of P. pyrenuloides. 

Pyrenula wrightii (Müll. Arg.) R. C. Harris, comb. nov. 

Melanotheca wrightii Müll. Arg., Bot. Jahrb. Syst. 6: 396. 1885. Type. CUBA. Wright, Verr. Cub. 

231(NY, isosyntype), Verr. Lich. Cub. II. 679 (H, isosyntype). 

This species has ascospores and other internal features identical to those of P. quassiaecola Fée 

differing in having the ascomata in pseudostromata filled with oxalate(?) crystals and whitish powder bleeding 

yellow in KOH. It differs from P. concastroma only in having colorless rather than reddish "oil" in the 

postmature ascospores. It is known only from the type. See also discussion of P. concastroma. 

Pyrenula sp. 47 

Discussed by Harris (1989: 102). The ascomata of the Florida collection show considerable fusion and 

are superficially indistinguishable from P. ravenelii. Previously known from Louisiana and Misssissippi, its 

occurrence in northernmost Florida is no surprise. 

Pyrenula sp. 208a 

Although extremely distinctive this species is not described as the available material is scant. It is 

included as part of my ongoing campaign to convince colleagues that ascomatal type in the Pyrenulaceae 

does not seem to have any meaning above the species level. This species combines features of both the 

melanothecoid and pyrenastroid types. The ascomata are recumbent, fused at the single ostiole but the 

pseudostromatic shells are fused into a single melanized mass as in the species formerly referred to 

Melanotheca Fée. I suppose in Zahlbruckner's system the former would have outweighed the latter and the 

species would have been placed in Parmentaria Fée. 

Pyrenula sp. 6121 

Superficially this species is similar to P. anomala but seemingly the ascomata are larger and although the 

pseudostromata are similar sized, there are fewer ascomata making them up. The ascospore difference is 

fairly slight and I have not yet done the statistics but my impression is that the greater ascospore width of sp. 

6121 will be the most reliable character for separation. I have not given the taxon a name as it is fairly 

widespread and surely has one elsewhere. Melanotheca arthonioides var. machaerii Malme from Brazil, if the 

type matches, seems to be this and could be elevated in rank. In Florida it is known from only two sites in 

Everglades National Park collected by Johnson in 1965. 

PYRGILLUS Nyl. 

The genus has only a single species in Florida. See the generic key for a diagnosis. 

105

REQUIENELLACEAE Boise 

The Requienellaceae are recognized here in the restricted, original sense as discussed above under 

Pyrenulaceae. While comfortable with the family, I have never been satisfied with the species level taxonomy. 

However, as Boise had decided, not readily changeable, opinions, I kept my mouth shut until now. I have 

always felt that R. subcollapsa and Acrocordiella occulta (see Pyrenulaceae) merited recognition. 

REQUIENELLA Fabre 

1. Young ascospores ± thin walled................................................................................................................ 2 

2. Ascospores with cells densely filled with small oil droplets, 6-celled, 

ca. 29-33 × 10-12 µm; ostiolar area inspersed with large oil drops; 

Europe.....................................................................................[Requienella seminuda (Persoon) Boise] 

2. Ascospores with 1-few oil droplets/masses, 8-celled, sometimes with 

one cell longitudinally divided, ca. 36-48 × 11-13 µm; Europe ................................................................ 

................................................................................................[Requienella lichenopsis (Massal.) Boise] 

1. Young ascospores with a well developed "cap" of endospore in the 

terminal cells; mature ascospores 4-celled, occasionally with 1-2 cells 

longitudinally subdivided, ca. 25-29 × 8-10 µm; on soft barked deciduous 

trees, eastern North America ............................. [Requienella subcollapsa (Ellis & Everhart) R. C. Harris] 

NOTES 

Requienella subcollapsa (Ellis & Everhart) R. C. Harris, comb. nov. 

Lophiostoma subcollapsa Ellis & Everhart, J. Mycol. 2: 100. 1886. Type. NEW JERSEY. Newfield, 

on outer bark of living Nyssa, Jun 1886, Ellis (NY, holotype). 

I am preliminarily assuming that there is only a single species in North Americ but I have not yet reexamined 

all the specimens cited by Boise as R. seminuda so that this assumption may prove incorrect. 

TRYPETHELIACEAE Zenker 

Harris (Acta Amazonica 14(1/2, Supl.): 55-80. 1984[1986]) included the genus Pleurotrema Müll. Arg. in 

the Trypetheliaceae. It has been since removed to its own family, Pleurotremataceae, in the non-lichenized 

unitunicate ascomycetes. (See Pyrenulaceae for more detailed comments). 

Aptroot (Biblioth. Lichenol. 44, 1991) recognized two additional genera of Trypetheliaceae, Megalotremis 

Aptroot and Ornatopyrenis Aptroot. Unfortunaely Aptroot's descriptions of ascus and hamathecium are not 

adequate and he did not trouble with conidia. I have not seen the types of either genera. 

Aptroot's hesitation in describing Megalotremis was well founded. Megalotremis, based on M. verrucosa 

(Makhija & Patwardhan) Aptroot, is certainly a synonym of Anisomeridium (Aptroot's drawings show both 

Anisomeridium-type ascospores and ascus tip.) Two-celled ascospores are unknown in the Trypetheliaceae. 

Anisomeridium, with a handful of exceptions, has 2-celled ascospores and species having asci with only 2 or 4 

ascospores are not uncommon. The thallus of M. verrucosa is corticate but corticate thalli are also seen in 

Anisomeridium (A. glaucescens and A. clandestinum). The second species, M. biocellata Aptroot, is almost 

certainly a species of Arthopyrenia (probably the same as Tomasellia dispora Müll. Arg., see Arthopyrenia). 

Most species of Arthopyrenia show some constriction of the ascospore wall ± grading into the "biocellate" type 

("porospore" in Vainio's terminology). (There are probably ten or so species with the porospore ascospore 

type.) Aptroot (Nova Hedwigia 60: 338. 1995) has recognized a third species, M. megalospora (Vainio) 

106

Aptroot, for which I have seen the type and a couple additional specimens. It belongs in Anisomeridium. The 

asci and physes are clearly the same as those seen in Monoblastiaceae. The ascospores of M. verrucosa 

and M. megalospora contain colorless crystals. (Eriksson also sees them as crystals but Aptroot interprets 

them as folds in the ascospore wall.) This character is seen in scattered species with large ascospores in the 

Monoblastiaceae, Pyrenulaceae and Trypetheliaceae and seems to have no lower level taxonomic 

significance. Possibly it is some storage product crystallizing which might provide support for ordinal 

relationship if biochemical analysis were to show it was unique or unusual. 

The disposition of Ornatopyrenis is more problematical. Ornamented ascospores are otherwise 

unknown in the entire family Trypetheliaceae but the pale brown, ornamented ascospores do suggest 

Mycomicrothelia (Arthopyreniaceae). Hawksworth's (1985) photo of the type, Microthelia queenslandica Müll. 

Arg., shows dark septa which also seem typical of Mycomicrothelia. Hawksworth's photo and Aptroot's 

drawings both show distoseptation which does not fit well with Mycomicrothelia as defined by Hawksworth. 

However, Aptroot described Mycomicrothelia rubrostoma with somewhat distoseptate ascospores although his 

examination was apparently so casual that he did not realize this. The ascospores of M. rubrostoma are 

similar to those in the porospore Arthopyreniae mentioned above, only brown. Thus, I am not uncomfortable 

with placement in the Arthopyreniaceae. Also the ascospores of Ornatopyrenis may become submuriform. I 

have already (1989) introduced a muriform-spored species into Mycomicrothelia, M. decipiens (Müll. Arg.) R. 

C. Harris. Until such time as I can study material of Ornatopyrenis queenslandica, I suggest Ornatopyrenis be 

placed tentatively as a synonym of Mycomicrothelia. 

KEY TO GENERA 

1. Thallus not corticate, whitish ..................................................................................................................... 2 

2. Ascospores with well developed endospore developed, lumina angular, 

4-celled, 20-27 × 6-8 µm; hymenium inspersed, also often yellow 

pigmented, KOH+ red; throughout the state ........................... Pseudopyrenula subnudata Müll. Arg. 

2. Ascospores with endospore not developed, lumina cylindrical, 

with 4-8 transverse septa or muriform.............................................................................. Polymeridium 

1. Thallus corticate, green, tan, brown or orange.......................................................................................... 3 

3. Ascomata solitary or in pseudostromata, not fused and sharing a common ostiole 

or ostiolar plate...................................................................................................................................... 4 

4. Stromata solitary or in pseudostromata which are not brown, and slightly 

shiny , if containing KOH+ purple pigment, then pseudostroma pruinose and 

ascospores 9-13 transversely septate; pseudostromal wall not composed 

of brown, convoluted hyphae............................................................................................................. 5 

5. Ascospores muriform .................................................................................................................... 6 

6. Ostiole apical..................................................................................................................Laurera 

6. Ostiole eccentric or lateral............................................................... [Campylothelium Müll. Arg.] 

5. Ascospores transversely septate .................................................................................................. 7 

7. Ascospores less than 100 µm long or if longer, then multiseptate.............................................. 

................................................................................................................... Trypethelium s. lat. 

7. Ascospores mostly more than 100 µm long, 3-5-septate..................[Architrypethelium Aptroot] 

4. Ascomata in brown, usually slightly shiny psedostromata containing 

yellow or orange pigments, KOH- or KOH+ red; pseudostromal 

wall composed of brown, jigsaw puzzle-like hyphae; ascospores 

107

transversely septate or muriform.........................................................................................Bathelium 

3. Ascomata compound, solitary or in pseudostromata, several hymenia with 

a common ostiole or ostiolar plate ........................................................................................................ 8 

8. Ascospores with well developed endospore ..................................................................................... 9 

9. Ascospores transversely septate or with a few cells longitudinally 

divided .......................................................................................................................Astrothelium 

9. Ascospores muriform .................................................................................[Cryptothelium Massal.] 

8. Ascospores without obvious endospore, submuriform; Brazil.....................[Exiliseptum R. C. Harris] 

ASTROTHELIUM Eschw. 

Astrothelium as presently constituted is polyphyletic with astrothelioid morphs from several different 

clades within the Trypetheliaceae. The clade containing the type species, A. conicum Eschw., is probably 

sufficiently distinct that Astrothelium in a restricted sense will be retained. "Species" based on presence or 

absence of lichexanthone will be abandoned as in Polymeridium. Some astrothelioid morphs, e.g., A. 

variolosum, confusum, do not seem to merit recognition even at the species level. Others will probably be 

recognized in Laurera along with much of the rest of the family. 

1. Ascomatal warts with orange pigmented tips or containing orange 

or brownish pigments, KOH+ purple or reddish ....................................................................................... 2 

2. Ascospores 4-celled, IKI-; pigment orange, KOH+ purple .................................................................... 3 

3. Thallus UV+ yellow (lichexanthone) .................................................................................................. 4 

4. Ascospores 28-35 × 10-13 µm; throughout the state ...............................A. versicolor Müll. Arg. 

4. Ascospores 21-28 × 7-9 µm; throughout the state .................................A. galbineum Krempelh. 

3. Thallus UV-; ascospores 23-30 × 8-10 µm; throughout the state......................................................... 

..............................................................................................A. cinnamomeum (Eschw.) Müll. Arg. 

2. Ascospores 10-celled, IKI+ violet, 90-110 × 22-28 µm; pigment 

around ostiolar neck brown, KOH+ dull reddish; thallus UV-; 

hymenium not inspersed except in ostiolar neck; Collier County................ A. diplocarpum Nyl. s. lat. 

1. Ascomatal warts covered by thallus or black where exposed, KOH-.......................................................... 

.................................................................................................................Trypethelium variolosum Ach. 

The astrothelioid morphs of this morphologically and chemically variable species have been recognized 

as follows. Some may find it more satisfying to continue to do so. 

A. Thallus UV+ yellow (lichexanthone); ascospores 20-25 × 7.5-10 µm; 

Franklin, Liberty and Monroe counties (intergrades with 

Trypethelium ochroleucum).....................................................................A. variolosum (Ach.) Müll Arg . 

A. Thallus UV-; ascospores 20-27 × 8-10 µm; Dade County 

(intergrades with Trypethelium nitidiusculum) .....................................................A. confusum Müll. Arg. 

Astrothelium diplocarpum Nyl. s. lat. 

NOTES 

108

The type collection has some ascospores with 1-several cells longitudinally divided and for that reason 

the species was placed in Cryptothelium by Zahlbruckner. This, then, is another of those intermediate taxa 

that suggest that generic separation between transversely septate and muriform ascospore types is 

meaningless. The Florida collection has transverse septa only but it resembles A. diplocarum in large 

ascomata, brown pigment (KOH+ reddish), and IKI+ violet ascospores that resemble the non-muriform ones in 

the type. In addition to the type from Colombia I have seen only two other collections (Costa Rica and 

Ecuador). Since is difficult to adequately interpret the ascospore variation from so few collections, I am 

tentatively including the Florida collection in A. diplocarpum. 

New to North America. FLORIDA. Collier County: Fakahatchee Strand State Preserve, Royal Palm 

Hammock, 5.7 mi from S entrance on W. J. Janes Memorial Scenic Drive, hardwood-Taxodium swamp with 

Roystonea, on Ilex, 6 Dec 1992, Harris 29938 (FLAS, NY). 

BATHELIUM Ach. 

Methodus 111, pl. 8, f. 3. 1803. Type. Bathelium mastoideum Ach. 

Bathelium mastoideum has been generally misapplied to a North American species with 4-celled 

ascospores but the type has muriform ascospores and ascomatal anatomy of the species known as Laurera 

madreporiformis (Eschw.) Riddle (see below). I have always been puzzled by Acharius' illustration as it has no 

resemblance to Trypethelium mastoideum auct. The North Italian lichenologists Massalongo (1860) and 

Trevisan (1853) correctly understood Bathelium to have muriform ascospores as did Müller initially. I do not 

understand why he switched his concept to a 4-celled taxon. It was unfortunate since Zahlbruckner followed 

this later misinterpretation and dragged 75 years of lichenologists into error. Dodge attempted to use 

Bathelium as a segregate genus for 3-septate species of Trypethelium (a sure guarantee of error) but this was 

never accepted. Fortunately the resurrection of Bathelium does not cause nomenclatural problems since B. 

mastoideum typifies a distinctive group of taxa that deserve generic recognition. Trypethelium s. str. 

encompasses only a few species. The central group, the majority of the family, will eventually be recognized 

in Laurera. Jørgensen and Santesson recently proposed Laurera for conservation with a new type, L. 

megasperma (Mont.) Riddle. Fortunately it is congeneric with the original holotype, L. varia (Fée) Zahlbr. 

although they could not have known that. Such haphazard conservation, changing type species without an 

intimate knowledge of the group and possible consequences, is foolish and should be firmly discouraged. 

Fortuitously, in this case, it does not cause problems. 

Bathelium is distinguished from Laurera and Trypethelium by pseudostromatal characters. The 

pseudostromata are somewhat raised to sessile, globose to elongated and irregular in outline, with one to 

many ascomata, brown, often shiny, rarely pruinose, with a very distinctive outer layer of brownish short celled, 

intricately entwined and branched hyphae forming a jigsaw puzzle-like pattern, pseudostromatal "medulla" 

filled with oxalate? and pigment crystals or granules, most often KOH+ red and a cartilaginous, amorphous 

cortex-like layer surrounding the individual ascomata within the pseudostroma. The ascospores range from 

relatively small (20-30 µm) and transversely 3-septate to very large (200-300 µm) and muriform. Nine species 

are currently included but I would anticipate additional taxa being added. There are two species in North 

America, both occur in Florida. 

1. Ascospores transversely septate, 4-6-celled; hymenium not inspersed ................................................... 2 

2. Ascospores 4-celled.............................................................................................................................. 3 

3. Ascospores 28 × 9 µm or less........................................................................................................... 4 

4. Pseudostromatal pigment KOH bleeding yellow; ascospores 

18-28 × 6-9 µm; throughout the state..................................B. carolinianum (Tuck.) R. C. Harris 

4. Pseudostromatal pigment bleeding red in KOH; ascospores 

20-27 × 8-9 µm; West Indies, Central and South America.............................................................. 

.............................................................................................. [B.degenerans (Vainio) R. C. Harris] 

109

3. Ascospores 35-50 × 12-15 µm; pseudostromata brown, ± shiny 

with medulla KOH+ red; Brazil .............................................[B. endochryseum (Vainio) R. C. Harris] 

2. Ascospores 6-celled, 25-32 × 6-9 µm; pseudostromata ± sessile, solitary 

or 2-3 fused, brown, often somewhat pruinose around ostiole; medulla 

orange-brown to brown in section, KOH+ scarlet; Southeast Asia ........................................................ 

............................................................................................... [B. albidoporum (Mak. & Pat.) R. C. Harris] 

1. Ascospores muriform; hymenium inspersed............................................................................................. 5 

5. Thallus ± smooth................................................................................................................................... 6 

6. Ascospores mostly over 50 µm long; Africa ...................................................................................... 7 

7A. Ascospores 44-80 µm long .........................................................................[B. mastoideum Ach.] 

7B. Ascospores 100-120 µm long .................................................... [B. lineare (Dodge) R. C. Harris] 

7C. Ascospores 130-200 µm long............................................ [B. sphaericum (Dodge) R. C. Harris] 

6. Ascospores 40-50 × 12-17 µm, with 10-11 transverse septa; 

throughout Florida ............................................................... B. madreporiforme (Eschw.) Trevisan 

5. Thallus with subglobose to subdactyliform areoles filled with white 

oxalate? crystals and a few bark cells; ascomata nestled between 

areoles, brown, with orange crystals in medulla, KOH+ scarlet; 

ascospores 90-140 × 27-35 µm; India............................... [B. tuberculosum (Mak. & Pat.) R. C. Harris] 

NOTES 

Bathelium albidoporum (Makhija & Patwardhan) R. C. Harris, comb. nov. 

Trypethelium albidoporum Makhija & Patwardhan, Internatl. J. Mycol. Lichenol. 5: 238. 1992. 

Type. [VIETNAM]. Tu-phap, Tonkin, Balansa s.n. (BM, holotype). 

Additional specimens. BURMA. Kachin State: along Ledo Road, 21 mi S of Bhamo, 650 ft, Apr 

1945, McMillen 245j (US). CHINA. Hainan: Ling-shui, Deng 1991 (BPI). 

Bathelium carolinianum (Tuck.) R. C. Harris, comb. nov. 

Trypethelium carolinianum Tuck., Amer. Jour. Sci. Arts, ser. 2, 25: 429. 1858. Type. SOUTH 

CAROLINA. 1851, Ravenel 298 (FH-Tuck 3952, lectotype, selected here). 

This seems to be the oldest name for the taxon previously known as Trypethelium mastoideum. Several 

older names have been cited as synonyms. The only one of these I have not re-examined is T. feei Meissner 

ex Fée. Trypethelium phlyctaena Fée and T. quassiaecola Fée are synonymous with T. ochroleucum. 

Trypethelium scoria Fée is apparently a good species of Trypethlium s. lat. 

Bathelium degenerans (Vainio) R. C. Harris, comb. nov. 

Pseudopyrenula degenerans Vainio, J. Bot. 34: 292. 1896. Trypethelium degenerans (Vainio) 

Zahlbr., Cat. lich. univ. 1: 490. 1922. Type. DOMINICA. Laudat, 1700 ft, Elliot s.n. (BM, isotype). 

This species is very close to B. carolinianum but the pseudostromatal pigment reacts KOH+ red. The 

pigments need to be analyzed. The KOH reaction may be due to differing pigment concentrations. I have 

seen this in other species where a reaction may vary from yellow to red depending on concentration. For the 

present I am maintaining them as distinct as there is little overlap in distribution. Bathelium degenerans is not 

known from North America but is not uncommon in the Caribbean and Mexico, also known from Brazil, Costa 

Rica and Venezuela. 

Bathelium endochryseum (Vainio) R. C. Harris, comb. nov. 

110

Pseudopyrenula endochrysea Vainio, Étud. class. lich. Brésil 2: 206. 1890. Type. BRAZIL. Minas 

Gerais: Caraça, 1885, Vainio, Lich. bras. 1157 (BM, M, isotypes). 

Bathelium endochryseum is known only from Brazil. 

Bathelium lineare (Dodge) R. C. Harris, comb. nov. 

Polyblastiopsis linearis Dodge, Ann. Missouri Bot. Gard. 40: 276. 1953. Laurera linearis (Dodge) 

Letrouit-Galinou, Rev. Bryol. Lichénol. 36: 253. 1957. Type. SIERRA LEONE. Njala (Kori), on 

Citrus aurantifolius, 27 Nov 1951, Deighton M4634 (FH-Dodge, holotype; BM, isotype). 

Bathelium madreporiforme (Eschw.) Trevisan 

Trypethelium madreporiforme Eschw., Syst. lich. 24, f. 24a-c. 1824. Laurera madreporiformis 

(Eschw.) Riddle in Howe, Torreya 16: 50. 1916. BRAZIL. Serra dos Montes altos, Martius, (M, 

holotype?). 

Trypethelium madreporiforme f. pruinosa Malme, Ark. bot. 19(1): 23. 1924. Type. BRAZIL. 

Mato Grosso: Cuyaba, in cort. Macherii, 28 Mar 1893, Malme, Regn. Lich. 2560A (S, lectotype, 

selected here). 

This is the only taxon of Bathelium with muriform ascospores in the New World. There is a disjunct 

record from Burma. The center of distribution is Africa. It is interesting that ascospores of Burman and North 

American collections are slightly smaller than in South American collections. Ascospore size variation in the 

group with muriform ascospores is almost continuous and requires more sophisticated reanalysis. The ranges 

for B. madreporiforme and B. mastoideum overlap considerably and it may prove necessary to synonymize 

them. 

Additional specimen. BURMA. Kachin State: along Ledo Road, 21 mi S of Bhamo, 650 ft, Apr 1945, 

McMillen 245j (US, mixed in collection of B. albidoporum). 

Bathelium mastoideum Ach. 

Methodus 111, pl. 8, f. 2-a-c. Type. SIERRA LEONE. Afzelius (S, lectotype; H-Ach 879-E, 

isolectotype). 

Trypethelium marginatum Fée, Ann. Sci. Nat. Bot. 23: 433, pl. 12, f. 2A-E. 1831. Laurera 

marginata (Fée) Dodge, Ann. Missouri Bot. Gard. 40: 297. 1953. Type. SENEGAL. Presque Ile 

du cap vert, Perrottet (G, holotype). 

Thelenella (Laurera) elegans Vainio, Bull. Soc. Broteriana sér. 2, 6: 174. 1930. Laurera elegans 

(Vainio) Zahlbr., Cat. lich. univ. 10: 103. 1938. Type. MOZAMBIQUE. Palma, 1916, Pires de Lima 

94 (TUR-Vainio 34704, holotype). 

?Laurera submadreporiformis Abbayes, Bull. Inst. Franç. Afr. Noire 15: 52. 1953. Type not 

seen. 

Polyblastiopsis pyriformis Dodge, Ann. Missouri Bot. Gard. 40: 276. 1953. Laurera pyriformis 

(Dodge) Letrouit-Galinou, Rev. Bryol. Lichénol. 26: 252. 1957. Type. SIERRA LEONE. Njala (Kori), 

on trunk of Anisophyllea laurina, 26 Oct 1951, Deighton M4404 (FH-Dodge, holotype; BM, 

isotype). 

Since none of the names cited here which are replaced by Bathelium mastoideum have ever appeared in 

the literature more than a couple of times, the change in name is of no consequence. The species is known 

only from Africa. Before examining the Acharian material I was puzzled by Acharius's illustration. Now it 

makes perfect sense and represents a typical species of the muriform-spored group of Bathelium. 

Bathelium sphaericum (Dodge) R. C. Harris, comb. nov. 

Polyblastiopsis sphaerica Dodge, Ann. Missouri Bot. Gard. 40: 277. 1953. Laurera sphaerica 

(Dodge) Letrouit-Galinou, Rev. Bryol. Lichénol. 26: 254. 1957. Type. SIERRA LEONE. Njala (Kori), 

on Orthocosmus africanus, 7 Apr 1952, Deighton M4795 (FH-Dodge, holotype; BM, isotype). 

Bathelium tuberculosum (Makhija & Patwardhan) R. C. Harris, comb. nov. 

Laurera tuberculosa Makhija & Patwardhan, Mycotaxon 31: 584. 1988. Type. INDIA. Karnataka, 

Sirsi-Kumtha Road, 48 km from Sirsi, 12 Dec 1974, Kulkarni 74.2689 (not seen). 

Additional specimens. INDIA. Karnataka, near Hebri, humid rain forest, elev. 50 m, 2 Feb 1977, 

111

Hale 48878 (US); South Canara, remnants of evergreen forest along road E of Ujire (Udipi 

area), 50 m, 22 Feb 1978, Hale 50546 (US). 

LAURERA Reichenb. 

Eriksson has suggested several times that Cryptothelium should be synonymized here. I agree. I have 

not formally done so since most of the family ends up in Laurera and I am waiting to make the transfers as a 

package in an organized fashion. Many species of Campylothelium belong here although the type, C. 

puiggarii Müll. Arg., is an oddball and might just be worth recognizing in its own genus. 

1. Pseudostromata poorly developed, brownish, almost pyrenuloid, KOH-; 

ascospores 8/ascus, 90-130 × 20-30 µm; Franklin County.......................................................................... 

.................................................................................................. L. subdisjuncta (Müll. Arg.) R. C. Harris 

1. Ascomatal warts mostly solitary, covered by yellow green thallus 

except at apex which usually is black with a whitish ring; ascospores 

4/ascus, 200-270 × 30-40 µm, with 50-70 transverse septa; 

hymenium inspersed; peninsular Florida................................................. L. megasperma (Mont.) Riddle 

POLYMERIDIUM (Müll. Arg.) R. C. Harris 

1. Ascospores transversely septate .............................................................................................................. 2 

2. Ascospores 4-celled............................................................................................................................... 3 

3. Thallus UV+ yellow (lichexanthone); ascospores 22-32 × 7-10 µm; 

hymenium not inspersed; common throughout the state .............. P. catapastum (Nyl.) R. C. Harris 

3. Thallus UV-......................................................................................................................................... 4 

4A. Ascospores 13-15.3-17 × 4-5-6 µm; Polk and Sarasota counties .................................................. 

.................................................................................................P. contendens (Nyl.) R. C. Harris 

4B. Ascospores 16-18.5-21 × 5-5.8-7 µm; throughout the state ........................................................... 

...............................................................................................P. subcinereum (Nyl.) R. C. Harris 

4C. Ascospores 19-22.3-25 × 5.5-6.6-7.5 µm; Dade County ................................................................ 

............................................................................................. P. albidum (Müll. Arg.) R. C. Harris 

2. Ascospores 5-11-celled......................................................................................................................... 5 

5. Ascospores (5-)6-8(-9)-celled, 18-22.5-27 × 4-5.5-7 µm; throughout the state .................................. 

...........................................................................................P. quinqueseptatum (Nyl.) R. C. Harris 

5. Ascospores (6-)8-10(-11)-celled, 24-32.3-40 × 5.5-7-8.5 µm; Dade County....................................... 

.......................................................................................P. albocinereum (Krempelh.) R. C. Harris 

1. Ascospores muriform, amyloid, 39-54 × 15-19 µm, (9-)11(-13) × 1-4-septate 

thallus UV+ yellow (lichexanthone) or UV-; hymenium not inspersed; 

south to Marion County ......................................................................... P. proponens (Nyl.) R. C. Harris 

PSEUDOPYRENULA Müll. Arg. 

Pseudopyrenula subnudata Müll. Arg. (1883) antedates P. subgregaria Müll. Arg. (1885). There is only 

the one species in North America. 

112

TRYPETHELIUM Sprengel 

1. Thallus or ascomatal wart KOH+ purple ................................................................................................... 2 

2. Thallus usually yellowish or brownish, KOH-; ascomata aggregated 

in raised pseudostromata which are often constricted at base, 

contents KOH+ purple.......................................................................................................................... 3 

3. Ascospores 10-14-celled, 40-50 × 9-12 µm; powder in pseudostromata 

yellow to tan; pseudostromata and/or thallus often pruinose; 

peninsular Florida..............................................................................................T. eluteriae Sprengel 

3. Ascospores 13-16-celled, 60-75(-85) × 11-12 µm; powder in 

pseudostromata orange; pseudostromata orange to brown, 

not pruinose; Alachua, Gilchrist, Polk and Sarasota counties.............................................................. 

................................................................................................................. T. subeluteriae Mak. & Pat. 

2. Thallus orange; ascomata mostly immersed, solitary and scattered 

to aggregated; ascospores 4-celled, 21-28 × 8-9 µm; peninsular Florida ............................................... 

.................................................................................................................. T. aeneum (Eschw.) Zahlbr. 

1. Thallus and warts KOH- ............................................................................................................................ 4 

4. Ascomata covered by thallus or in pseudostroamta ......................................................................... 5 

5. Ascospores 4-celled, IKI- .............................................................................................................. 6 

6. Ascospores smaller, less than 20-27 × 7.5-10 µm long .............................................................. 

....................................................................................................Trypethelium variolosum Ach. 

The trypethelioid morphs with pseudostromata containing 1-many simple ascomata have been 

recognized as follows. Some may find it more satisfying to continue to do so. 

A. Thallus UV+ yellow (lichexanthone);throughout the state 

........................................................................................................ T. ochroleucum (Eschw.) Nyl. 

A. Thallus UV-; throughout the state.................................................T. nitidiusculum (Nyl.) R. C. Harris 

6. Ascospores large, 35-43 × 12-15 µm; ascomata solitary or 

grouped, covered by yellow green thallus except at apex which 

is black with a pale ring; hymenium not inspersed; peninsular Florida ....................................... 

........................................................................ T. floridanum (Zahlbr. ex Choisy) R. C. Harris 

5. Ascospores 8-12-celled, 38-52 × 7-10 µm, IKI+ weak violet; 

pseudostromata not raised; south to Marion County........................T. virens Tuck. ex Michener 

4. Ascomata black, naked, solitary or aggregated; hymenium inspersed; 

ascospores 4-celled, 20-26 × 6-8 µm; throughout the state................................................................ 

............................................................................................................T. tropicum (Ach.) Müll. Arg. 

NOTES 

Trypethelium aeneum (Eschw.) Zahlbr. 

Trypethelium ferrugineum Müll. Arg., Bot. Jahrb. Syst. 6: 392. 1885. Type. CUBA. Wright, Verr. 

Cub. 157, Lich. Cub. II, 593, 594 (FH-Tuck 3972, isosyntypes), syn. nov. 

Trypethelium ferrugineum β. inornatum Müll. Arg., Bot. Jahrb. Syst. 6: 392. 1885. Type. CUBA. 

113

Wright, Verr. Cub. 157 (G, holotype; US, isotype), syn. nov. 

Trypethelium ferrugineum represents the astrothelioid morph of T. aeneum analogous to "Astrothelium" 

confusum or variolosum in the Trypethelium variolosum aggregate. It has never been combined in 

Astrothelium and since I believe these astrothelioid morphs to be of little significance, I will not do so. Others 

may wish to. If one were a real hardcore lumper, T. aeneum, could also be included in the T. variolosum 

group as chemotype with anthraquinone pigments instead of lichexanthone. 

Trypethelium subeluteriae Makhija & Patwardhan 

Trypethelium eluteriae var. polystomum Malme, Ark. Bot. 19(1): 25. 1924. Type. BRAZIL. Mato 

Grosso: Buriti, in cort. Platypodii, 16 Jun 1894, Malme s.n. (S, lectotype, selected here), syn. 

nov. 

This taxon was noted in Harris (1986) and in "Some Florida Lichens" as Trypethelium sp. aff. eluteriae. 

Unfortunately Makhija and Patwardhan ran across a really scrappy specimen from Sri Lanka that I had so 

annotated and used it as the type for this widespread species. Infinitely better material was available. The 

species is not confined to Sri Lanka as they would imply. It is known from Australia, Central America (Costa 

Rica), India, Indochina, Indonesia, Papua New Guinea, Philippines, South America (Bolivia, Brazil, Paraguay, 

Venezuela), West Indies (Cuba, Dominican Republic) and Florida. 

New to North America. FLORIDA. Alachua County: Paynes Prairie State Preserve, Bolen Bluff Trail off US 

441, ca. 3.5 mi N of main entrance, 29°34'N, 82°21'W; dry hardwoods with Sabal pametto, 1 Dec 1992, Buck 

22561; Lafayette County: along Fla Hwy 51, 5.7 mi S of US 27 at Mayo, 29°59'N, 83°14'W; hardwood swamp, 

29 Nov 1994, Buck 27089; Polk County: along US 17, 2 mi N of Ft. Meade, 2 Feb 1965, Johnson 6205, 6209; 

Sarasota County: Myakka State Park, 31 Jan 1965, Johnson 6186 (all NY). 

Trypethelium tropicum (Ach.) Müll. Arg. 

A collection from Lake County (Harris 23579, NY) has an unusual, thick, bullate-rugose areolate thallus 

with a very thick medulla filled with white oxalate? crystals. The ascomata, hamathecium and ascospores are 

perfectly typical T. tropicum. I have speculated that thallus appearance is often substrate dependent. This 

collection is on Acer rubrum which is a common substrate for "typical" T. tropicum, so that doesn't seem to be 

the case here. I include this comment so that I won't in future just sweep it under the rug. 

Trypethelium variolosum Ach. 

After 20 years and hundreds, if not thousands, of specimens I have come to the conclusion that there is a 

complete series of intermediates between the trypethelioid and astrothelioid morphs in this aggregate species. 

The astrothelioid ascoma in this taxon seems to be derived from the invagination of the ascomatal wall (not 

fusion of individual ascomata). All stages can be found, from basal, incomplete, non-melanized invaginations 

to complete, melanized dividing walls. Since I am tired of flipping a coin to decide if the invaginations are 

complete enough to warrant placement in Astrothelium s. lat. or not (there is even some variation on a single 

thallus) and since the amount of invagination seems to have no predictive value, I have decided to merge the 

two "morphs". Also, I have concluded that here (and throughout the family) the presence or absence of 

lichexanthone should not be used as a specific character (Harris, 1993). (Almost every lichexanthone 

containing taxon has some lichexanthone deficient collections.) Further, some collections in this group could 

be segregated on the basis of a large amount of cartilaginous tissue in the pseudostromata as T. leprieurii. 

Again, I find a complete gradation from essentially none to comprising most of the pseudostroma. Two further 

characters which do not seem to correlate in any meaningful way are hymenial inspersion and oxalate crystals 

in the ascomatal wall. As far as I can tell there is not the slightest correlation between any of these characters 

and ecology or geographic distribution. This complex seems to represent one of those cases where the 

extremes are distinct but the middle ground is vast and messy. Using the above characters in all possible 

combinations (most of which I have run across) one could recognize dozens of "species". The oldest name 

for a member of this aggregate is Trypethelium variolosum Ach., an astrothelioid morph containing 

lichexanthone. Trypethelium variolosum occurs throughout Florida. The following segregates have previously 

been recognized and some may wish to continue to do so. 

1). Trypethelium ochroleucum (Eschw.) Nyl. - Asomata solitary to aggregated in trypethelioid 

pseudostromata; lichexantone present; common, throughout Florida. Those who wish to recognize this 

commonly used name will have to protect it against a couple of older Fée names. 

114

2). Trypethelium variolosum Ach., Astrothelium variolosum (Ach.) Müll. Arg. - Ascomata astrotheliod; 

lichexanthone present; throughout Florida. 

3). Trypethelium nitidiusculum (Nyl.) R. C. Harris - Ascomata solitary to aggregated in trypethelioid 

pseudostromata; lichexanthone absent; common, throughout Florida. Again there are older names but 

nitidiusculum has been taken up so recently (1986) that I see no point in protecting it. 

4). Astrothelium confusum Müll. Arg. - Ascomata astrothelioid; lichexanthone absent; Dade County. 

5). Trypethelium leprieurii Mont. - Ascomata trypethelioid with much cartilaginous tissue in the 

pseudostromata; lichexanthone present. I have never segregated any North American material into this 

morph but I don't doubt that one could. My awareness of this morphotype is fairly recent. 

MONOBLASTIACEAE W. Watson 

New Phytol. 28: 106. 1929. Type. Monoblastia Riddle 

Acrocordiaceae Oxner ex Barr, Mycotaxon 29: 505. 1987. Type. Acrocordia Massal. 

Additional evidence has been found to support the recognition of the Monoblastiaceae. Sabine Huhndorf, 

attempting to place the genus Ascocratera Kohlmeyer, examined the asci of a number of species for 

fluorescent structures in the ascus tip. Those taxa examined uniformly had either a fluorescent ring or 

fluorescent cap. Two species of Acrocordia, two species of Monoblastia and two of Anisomeridium had a 

fluorescent cap. Four species of Anisomeridium had fluorescent rings. Those with caps included the type 

species of Acrocordia, Anisomeridium and Monoblastia. The type species of Ditremis had a distinct ring. 

(Ascocratera had a ring but considering all characters one reaches the impasse that there is no definitive 

reason to either include it in or exclude it from the Monoblastiaceae). 

KEY TO GENERA 

1. Ascospores 2-4-celled, granular ornamented or smooth .......................................................................... 2 

2. Asci various with narrow ocular chamber; ascospores in a single row, 

in two rows or irregularly arranged, 2-4-celled, colorless or brown, 

ornamented or smooth; more common in tropical regions ....................................................................3 

3. Microconidia and macroconidia borne in pycnidia......................................................Anisomeridium 

3. Microconidia pycnidial; macroconidia in campylidia ....................................................... [Musaespora] 

2. Asci cylindrical with very broad ocular chamber; ascospores 

in a single row in the ascus, colorless, 2-celled, oblong, 

granular ornamented; temperate regions ............................................................................. Acrocordia 

1. Ascospores simple, ornamentation wart-like or peg-like........................................................ Monoblastia 

ACROCORDIA Massal. 

1. Growing on bark ........................................................................................................................................ 2 

2. Ascospores larger, 18-60 × 9-23 µm..................................................................................................... 3 

3. Ascospores 18-27 × 9-13 µm; ostiole usually apical; 

Duval County......................................................................................... A. gemmata (Ach.) Massal. 

3. Ascospores 33-48(-60) × 15-23 µm; ostiole usually eccentric; 

eastern North America ...............................................................[A. megalospora (Fink) R. C. Harris] 

115

2. Ascospores small, 11-17 × 6-9.5 µm; eastern North 

America ....................................................................................................[A. cavata (Ach.) R. C. Harris] 

1. Growing on calcareous rock; ascospores 13-17 × 7-8.5 µm; 

Michigan ............................................................................................................. [A. conoidea (Fr.) Körber] 

NOTES 

Acrocordia gemmata (Ach.) Massal 

The Florida collection is the only North American record and was not included in my thesis. One wishes 

it were made by someone other than Calkins who does not have a good track record for relaibility. 

FLORIDA. Duval County: Jacksonville, on cypress, Calkins, N.A.L 238 (NY). 

ANISOMERIDIUM (Müll. Arg.) Choisy, nom. cons. 

Icon. Lich. Univ. fasc. 3 (unpaged). 1926. Arthopyrenia sect. Anisomeridium Müll. Arg., Flora 66: 

290. 1883. Lectotype (Riedl, 1963). Arthopyrenia xylogena Müll. Arg. (= Anisomeridium 

subnectendum (Nyl.) R. C. Harris). 

Lembidium Körber, Syst. Lich. Germ. 358. 1855, nom. rejic. Type. Lembidium polycarpum Körber (= 

Anisomeridium biforme (Borrer) R. C. Harris). 

Ditremis Clements, Genera of Fungi 41. 1909. Type. Pleurotrema inspersum Müll. Arg. (= 

Anisomeridium americanum (Massal.) R. C. Harris). 

Sarcinulella Sutton & Alcorn, Mycotaxon 16: 557. 1983, nom. illeg.? Type. Sarcinulella banksiae Sutton & 

Alcorn (= Anisomeridium nyssaegenum (Ellis & Everhart) R. C. Harris?). 

Megalotremis Aptroot, Biblioth. Lichenol. 44: 124. 1991. Type. Ditremis verrucosa Makhija & Patwardhan 

(= Anisomeridium verrucosum (Makhija & Patwardhan) R. C. Harris). 

I had originally intended to recognize both Anisomeridium and Ditremis in this volume, based partly on 

evidence from fluorescence studies of ascus tips by Sabine Huhndorf. Unfortunately Murphy's Law applies 

and the new data does not correlate well with other characters. I am confident that with more species studied 

for tedious and often hard to interpret ascus and anamorph characters, it will be possible to subdivide 

Anisomeridium s. lat. into several genera. The largest group evident at this time is that centered around the 

type of Anisomeridium, with thallus lacking lichexanthone, apical ostioles, clavate or saccate asci with a 

fluorescent cap, blunt, ± ovoid, often thick-walled ascospores and globose microconidia. The other, less 

cohesive, group centers around the type of Ditremis with thallus sometimes with lichexanthone, often eccentric 

ostioles, cylindrical asci with a fluorescent ring? (only one sampled), ascospores with ± pointed ends and 

oblong or elliptical microconidia. Anisomeridium biforme has a fluorescent cap and redisposition into 

Acrocordia should be considered. Unfortunately the ubiquitous A. nyssaegenum seems to stand alone. It is 

also possible that some taxa may be removed to Musaespora. 

Being unable to recognize two genera at this time, I have decided to accept the decision of the North 

European lichen oligarchs to force their will on the rest of the world and conserve Anisomeridium. (There are 

only three species in North Europe but 100+ in the rest of the world. Makhija & Patwardhan (1990) recognized 

18 species of Ditremis for India alone. As will be seen from the general treatment below, I was in fact very 

conservative in making combinations in Ditremis.) When I fought for Ditremis in "Some Florida Lichens", I was 

aware that this was not a battle I would win but I felt obliged to protest the rape of the International Code of 

Nomenclature by nomenclatural Luddites fearing change (mostly the result of a shift from an artificial system 

to a phylogentic system, already too long postponed in mycology). Such unfortunate precedents may, 

however, help win the war in St. Louis. 

KEY TO ANISOMERIDIUM IN FLORIDA 

116

1. Ascomata present; pycnidia present or absent, never with long, hairlike beaks....................................... 2 

2. Thallus UV-............................................................................................................................................ 3 

3. Ascospores 2-celled; ostiole apical or eccentric; growing on bark....................................................4 

4. Ostiole apical................................................................................................................................. 5 

5. Ascospores 12-24 × 4.5-8.5 µm ................................................................................................ 6 

6. Asci slender, cylindrical to narrowly clavate .......................................................................... 7 

7. Ascospores slender, narrowly ovate, fusiform to almost 

cylindrical (often slightly bent in A.tamarindi) .................................................................... 8 

8. Microconidia elliptical, 2.5-1.5 × 2 µm; macroconidia ± oblong, 

5-7 × 2.5-3 µm; ascospores narrowly ovoid, with weakly 

submedian septum; 12-18(-21) × 4.5-6.5 µm; neotropical 

north to Lake Couny............................................... A. subprostans (Nyl.) R. C. Harris 

8. Microconidia orbicular; macroconidia elliptical, 

4-7 × 2.5-3.5 µm; ascospores 15-22(-24) × 

4.5-5.5(-6) µm, often slightly bent; north to 

Volusia County .............................................................A. tamarindi (Fée) R. C. Harris 

7. Ascospores broadly elliptical to ovoid, rarely narrowly ovoid, 

with septum slightly submedian or ± median,10-18 × 4.5-7 µm; 

microconidia oblong or elliptical, 2.5-3 × 1.5-2 µm or globose, 

ca. 2 µm diam.; macroconidia elliptical or oblong, 5-7 × 3-4 µm; 

throughout Florida .......................................................... A. biforme (Borrer) R. C. Harris 

6. Asci ovate or obovate to narrowly ovate or obovate.............................................................. 9 

9. Ascospores relatively broad, 5.5 µm or more.................................................................. 10 

10. Ascus wall and ascospore wall thick and unevenly thickened; 

ascospores ovoid, with markedly submedian septum, 

15-22 × 6.5-8.5 µm; microconidia globose, ca. 2 µm diam; 

macroconidia oblong or ovoid, 5-7 × 2.5-3 µm; Dade 

County ..........................................................A. anisolobum (Müll. Arg.) R. C. Harris 

10. Ascus wall and ascospore wall relatively thin and even; 

ascospores 18-22 × 5.5-7 µm, ± fusiform, often slightly 

bent, with slightly submedian septum; microconidia globose 

or elliptical, ca. 2 µm diam. or 2-3 × 1.5-2 µm; Seminole 

County ..........................................................A. excaecariae (Müll. Arg.) R. C. Harris 

9. Ascospores relatively narrow, 14-20 × 4.5-6 µm, ovoid, with 

submedian septum, often becoming 4-celled; micropycnidia 

beaked ("Sarcinulella"); microconidia oblong or narrowly elliptical, 

2-3 × 1-1.5 µm; Columbia County ....................................................................................... 

..............................................................A. nyssaegenum (Ellis & Everhart) R. C. Harris 

117

5. Ascospores small, 9-13(-15) × 4-5 µm, ovoid, with septum 

median to slightly submedian; ascomata small, 0.2-0.3(-0.4) mm 

diam.; microconidia oblong or elliptical, Duval, Liberty and Seminole 

counties ..................................................................................... A. albisedum (Nyl.) R. C. Harris 

4. Ostiole eccentric............................................................................................................................ 11 

11. Ascospores granular ornamented, fusiform, with septum moderately 

submedian, 19-27(-30) × 5.5-7(-8) µm; microconidia oblong or 

elliptical, 2.5-3 × 1.5-2 µm; ascomata usually naked; Collier and 

Monroe counties................................................................... A. terminatum (Nyl.) R. C. Harris 

11. Ascospores smooth, ± ovoid ................................................................................................... 12 

12. Ascospore septum distinctly submedian; ascospores 

32-40(-45) × 13.5-18 µm; microconidia subglobose, 

ca. 2-2.5 µm diam.; macroconidia elliptical or oblong, 

5-7 × 2.5-4 µm; ascomata covered by thallus; Duval, 

Franklin and Marion counties ...........................................................A. griffinii R. C. Harris 

12. Ascospore septum ± median; ascospores 17-23 × 7-9 µm; 

pycnidia not found; Levy County..................................Anisomeridium sp. Harris 29392-A 

3. Ascospores 4-celled or if 2-celled, growing on rock; ostiole always apical 

(2-celled, on rock, see also Pyrenocollema and Zwackhiomyces) ................................................... 13 

13. Ascospores 4-celled.................................................................................................................... 14 

14. Growing on bark .................................................................................................................... 15 

15. Ascospores soon 4-celled ............................................................................................... 16 

16. Ascospores 21-24 × 6-8 µm; microconidia elliptical, 3-4 × 

1.5-2 µm; Columbia County........................................ A. quadrococcum R. C. Harris 

16. Ascospores16-20 × 3.5-4.5 µm; microconidia ± globose, 

ca. 2 µm diam.; macroconidia oblong, 5-6 × 2-3 µm; Louisiana, 

not yet found in Florida............................ [A. quaternarium (R. C. Harris) R. C. Harris] 

15. Ascospores tardily 4-celled, ovoid, 14-20 × 4.5-6 µm; 

Columbia County...................................... A. nyssaegenum (Ellis & Everh.) R. C. Harris 

14. Growing on sandstone; ascospores tardily 4-celled, ovoid, with first 

septum submedian, smooth, 16-22 × 6.5-8 µm; microconidia narrowly 

elliptical, 4-5 × 1.5-2 µm; macroconidia ovoid or elliptical, 5-7 × 2.5-3 µm; 

Massachusetts to Illinois south to southern Georgia and Louisiana 

but not yet found in Florida .................................................... [A. distans (Willey) R. C. Harris] 

13. Ascospores 2-celled.................................................................................................................... 17 

17. Growing on limestone; ascospores narrowly ovoid to 

fusiform, with submedian septum, smooth, 20-25 × 

6.5-8.5 µm; microconidia elliptical, 3-5 × 1-1.5 µm; 

macroconidia broadly elliptical, 5-6 × 4-4.5 µm; Dade 

and Monroe counties........................................................ A. finkii (R. C. Harris) R. C. Harris 

17. Growing on sandstone; "immature" forms, see couplet 

118

14 above.................................................................................. [A. distans (Willey) R. C. Harris] 

2. Thallus UV+ yellow (lichexanthone) ..................................................................................................... 18 

18. Ascospores over 20 µm long.......................................................................................................... 19 

19. Ostiole eccentric, ringed with orange pigment, KOH+ purple; 

ascospores biseriate, fusiform, with septum submedian, 

smooth, 37-52 × 12-20 µm; microconidia oblong, 3-4 × 1.2-1.5 µm; 

macroconidia broadly elliptical to ovoid, 30-45 × 15-20 µm; 

Dade County............................................................................A. aureopunctatum R. C. Harris 

19. Ostiole apical, unpigmented; ascospores mostly uniseriate, 

± fusiform with one end often more rounded, with septum 

median or submedian, smooth, 24-33 × 8-12 µm; 

microconidia elliptical, 3-5 × 1.5-2 µm; macroconidia 

broadly ovoid to ovoid, thick-walled,15-22 × 9-16 µm; 

peninsular Florida............................................................. A. ambiguum (Zahlbr.) R. C. Harris 

18. Ascospores less than 20 µm long .................................................................................................. 20 

20. Ascospores colorless ............................................................................................................... 21 

21. Ascospores fusiform, with slightly submedian septum, smooth, 

15-20 × 5-6.5 µm; macroconidia cylindrical 7.5-12 × 2-2.5 µm; 

microconidia elliptical to subglobose, 2.5-3 × 1.5-2 µm or ca. 2 µm diam.; 

peninsular Florida, Lousiana ................................... A. tuckerae (R. C. Harris) R. C. Harris 

21. Ascospores broadly elliptical, not constricted at median septum, 

smooth, (10-)12-15 × (6-)7.5-9 µm; thallus ± corticate; microconidia 

broadly elliptical 3 × 1.5-2 µm; Okefenokee Swamp, Georgia and 

Polk County .............................................................................. A. biformoides R. C. Harris 

20. Ascospores soon dark brown, elliptical or ends slightly pointed, 

with septum ± median, granular ornamented, uniseriate in ascus, 

16-20 × 8-9 µm; thallus white, UV+ yellow (lichexanthone); ostiole apical; 

Collier County........................................................................... A. phaeospermum R. C. Harris 

1. Ascomata not known; pycnidia with hair-like beaks, to 1 cm long; macroconidia 

ellipsoid to narrowly ellipsoid, 5-7 × 2-2.5 µm, in ± packets surrounded by a 

gelatinous sheath; Collier County...........................................................Anisomeridium sp. Buck 22812 

NOTES 

Anisomeridium albisedum (Nyl.) R. C. Harris 

in Egan, Bryologist 90: 163. 1987. Verrucaria viridiseda f. albiseda Nyl., Lich. Japon. 108. 

1890. Porina viridiseda f. albiseda (Nyl.) Zahlbr., Cat. lich. univ. 1: 409. 1922. Ditremis 

albiseda (Nyl.) R. C. Harris, Some Florida Lichens 31. 1990. Type. FLORIDA. Calkins (H- Nyl 

717, lectotype; MICH, US, isolectotypes). 

Anisomeridium ambiguum (Zahlbr.) R. C. Harris 

in Egan, Bryologist 90: 163. 1987. Arthopyrenia (Acrocordia) ambigua Zahlbr., Ann. Mycol. 3: 

35. 1935. Ditremis ambigua (Zahlbr.) R. C. Harris, Some Florida Lichens 31. 1990. Type. FLORIDA. 

Seminole County: Sanford, on elm, May 1923, Rapp 142 (W-2452, holotype). 

119

Anisomeridium anisolobum (Müll. Arg.) Aptroot 

Biblioth. Lichenol. 57: 21. 1995. Arthopyrenia anisoloba Müll. Arg., Flora 66: 305. 1883. Ditremis 

anisoloba (Müll. Arg.) R. C. Harris, Some Florida Lichens 31. 1990. Type. BRAZIL. São Paulo: 

Apiaí, 1882, Puiggari (G, holotype). 

Arthopyrenia feeana Müll. Arg., Mém. Soc. Phys. Hist. Genève 30(3): 28. 1888. Anisomeridium 

feeanum (Müll. Arg.) R. C. Harris in Tucker & Harris, Bryologist 83: 4. 1980. Type. WEST INDIES. 

On Croton cascarillae, ex hb. Fée (G, holotype). 

Arthopyrenia (Mesopyrenia) knysana Zahlbr., Ann. Crypt. Exot. 5: 200. 1932. Type. SOUTH 

AFRICA. Cape Province: Knysna, on bark of Acacia melanoxylon, v. d. Byl 916 (LD, isotype). 

Anisomeridium aureopunctatum R. C. Harris, nom. nov. 

Ditremis macrospora R. C. Harris, Some Florida Lichens 33. 1990. Type. FLORIDA. Dade 

County: Everglades National Park, NE of Pa-hay-okee Overlook, 13 May 1967, Harris 2791-A 

(MSC, holotype), non Makhija & Patwardhan, Biovigyanam 16: 19. 1990 (before? 21 

December). 

Anisomeridium biforme (Borrer) R. C. Harris 

in V_zda, Lich. sel. exs. fasc. 61: 1305. 1978. Verrucaria biformis Borrer in Hooker & Sowerby, 

Suppl. Engl. Bot. 1: pl. 2617, f. 1. 1831. Arthopyrenia biformis (Borrer) Massal., Framm. Lich. 

26. 1855. Ditremis biformis (Borrer) R. C. Harris, Some Florida Lichens 32. 1990. Type not 

seen. 

Verrucaria conformis Nyl., Flora 47: 357. 1864; Arthopyrenia conformis (Nyl.) Müll. Arg., Bull. 

Herb. Boissier 2(app. 1): 90. 1894. Type. FRANCE. Finistère, Brest, Crouan 28 (H-Nyl 1653, 

lectotype). 

Verrucaria alboatra Krempelh. ex Nyl., Flora 50: 8. 1867. Arthopyrenia alboatra (Krempelh. ex 

Nyl.) Müll. Arg., Flora 66: 306. 1883. Type. INDIA. West Bengal, Calcutta Botanic Gardens, 

1866, Kurz 10 (H-NYL 1660, lectotype; BM, M, isolectotypes). 

Arthopyrenia brachyspora Müll. Arg., Nuovo Giorn. Bot. Ital. 21: 52. 1889. Type. ARGENTINA. Isla 

de los Estados, Spegazzini (G, holotype). 

Arthopyrenia parvula Zahibr., Beih. Bot. Centralbl. 13: 149. 1902. Type. UNITED STATES. 

California: Santa Monica Mountains, Malibu Canyon, Aug. 1898, Hasse 759 (FH, MIN, US, 

isotypes). 

Didymella microspora Vainio, Ann. Acad. Sci. Fenn. A 19(15): 15.1923. Type. PHILIPPINE ISLANDS. 

Comiran Island: Sulu Sea, Sep 1910, Merrill, Bur. Sci. 7165 (TUR-Vainio 32293, holotype). 

Arthopyrenia amaura Zahlbr. in Handel-Mazzetti, Symb. Sinic. 3: 19. 1930. Type. CHINA. Hunan, 

near Tschangscha, 10 Dec 1917, Handel-Mazzetti 11426 (US, isotype). 

Didymella laurilae Räsänen, Ann. Bot. Soc. Zool. Bot. Fenn. Vanamo 12(1): 204. 1939. Type. 

FINLAND. Par. Kaukola, Heinsalmi, in ripa boreale lacus Ruokajärvi, ad lignum madidum in 

aquam dejectum in luco ripario, 25 Jul 1935, Laurila (H, lectotype; H, isolectotype). 

Anisomeridium biformoides R. C. Harris, sp. nov. 

Similis A. biformis quoad characteribus microscopicis differt thallo UV+ aureo. 

Type. GEORGIA. Charleton County: Okefenokee National Wildlife Refuge, Chesser Island, 15 Nov 1989, 

Wetmore 64362 (NY, holotype; MIN, isotype, not seen). 

Thallus pale gray, endophloeodal, UV+ yellow (lichexanthone). Ascomata immersed, tip exposed, 

subglobose, ca. 0.3-0.5 mm diam.; clypeus apical and shield shaped to hemispherical, occasionally slightly 

laterally expanded. Asci cylindrical, ca. 120-135 × 12-15 µm, with eight uniseriate spores. Ascospores broadly 

elliptical, not constricted at median septum, smooth, (10-)12-15 × (6-)7.5-9 µm; microconidia elliptical, 3 × 1.5- 

2 µm; macroconidia not found. 

120

This species was treated in 1990 as Ditremis sp. Wheeler 15-4-89 from Polk County. Better material 

collected by Wetmore in Georgia makes its recognition here warranted. 

Additional specimen. FLORIDA. Polk County: Nalcrest, W side of recreation area, on fallen oak branch, 15 

Apr 1989, Wheeler s.n. (NY). 

Anisomeridium distans (Willey) R. C. Harris 

in Egan, Bryologist 90: 163. 1987. Verrucaria distans Willey, Enum. Lich. New Bedford 38. 

1892. Arthopyrenia distans (Willey) Zahlbr., Cat. Lich. Univ. 1: 276. 1921. Ditremis distans 

(Willey) R. C. Harris, Some Florida Lichens 32. 1990. Type. MASSACHUSETTS. Bristol County: 

New Bedford, 1866, Willey (MICH, isotype). 

Anisomeridium excaecariae (Müll. Arg.) R. C. Harris, comb. nov. 

Sagedia excaecariae Müll. Arg., Flora 63: 289. 1880. Arthopyrenia excaecariae (Müll. Arg.) 

Müll. Arg., Flora 66: 305. 1883. Type. ARGENTINA. Buenos Aires, on Excaecaria biglandulosa, 

Schnyder 2 (G, holotype). 

Arthopyrenia incurva Müll. Arg., Flora 66: 305. 1883. Type. BRAZIL. São Paulo: Apiaí, Mar 

1880, Puiggari 138.b pr. p. (G, holotype). 

Arthopyrenia sanfordensis Zahlbr., Ann. Mycol. 33: 33. 1935. Anisomeridium sanfordense 

(Zahlbr.) R. C. Harris in Egan, Bryologist 90: 163. 1987. Ditremis sanfordensis (Zahlbr.) R. C. 

Harris, Some Florida Lichens 34. 1990. Type. FLORIDA. Seminole County: Sanford, on 

persimmon, Apr 1932, Rapp 134 (W-2460, holotype). 

Anisomeridium finkii (R. C. Harris) R. C. Harris, comb. nov. 

Ditremis finkii R. C. Harris, Some Florida Lichens 32. 1990. Type. PUERTO RICO. Arecibo District: 

Manati, 12 Jan 1916, Fink 2052 (US, holotype; FH, isotype). 

Anisomeridium griffinii R. C. Harris, sp. nov. 

Thallo albo, UV-, ostiolo eccentrico, ascosporis ovoideis vel interdum late fusiformibus, laevibus, septis 

distincte submedianis, 32-38 × 13.5-18 µm et macroconidiis ovoideis, 5-7 × 2.5-4 µm. 

Type. FLORIDA. Marion County: ca. 0.5 mi S of U.S. 40 & 1 mi W of Oklawaha River bridge, mesic hammock 

of Liquidambar styraciflua, Celtis laevigata, Sapindus marginata & Quercus virginiana, on trunk of Ulmus 

crassifolia, 23 May 1994, Griffin 3217 (NY, holotype; ESS, FLAS, H, isotypes). 

Thallus white, endophloeodal, UV-. Ascomata covered by thin layer of thallus, ± lageniform, ca. 0.5-0.6 × 

0.4 mm; ostiole eccentric to lateral; wall entirely melanized or only to base of hamathecium. Asci cylindrical to 

subclavate, 150-210 × 30-40 µm, with eight biseriate spores. Ascospores ovoid or somewhat fusiform, slightly 

constricted at the markedly submedian septum, smooth, 32-40 × 12-18 µm; microconidia subglobose, ca. 2 

µm diam.; macroconidia ovoid, 5-7 × 2.5-4 µm. 

It is a genuine pleasure to name this species to honor its collector, Dana Griffin, III, who has unstintingly 

supported work on Florida lichens. Anisomeridium griffinii seems closest to Asian A. palvanum, differing in 

larger ascospores. In Florida it might be confused with A. ambiguum which differs in UV+ thallus and apical 

ostiole. Anisomeridium griffini was treated in 1990 as Ditremis sp. 11288 and Ditremis sp. Buck B724 (ostiole 

orientation misunderstood). 

Additional specimens. FLORIDA. Duval County: Jacksonville Beach, flood plain forest W and NW of 23rd 

& 24th St. N., between 9th & 11th Ave. N., on Liquidambar, 23 Dec 1975, Buck B724; Franklin County: 

Apalachicola National Forest, Hickory Landing Rec. Area, ca. 2 mi S of Sumatra, hardwoods along Owl Creek, 

23 May 1976, Harris 11288 (both NY). 

121

Anisomeridium nyssaegenum (Ellis & Everhart) R. C. Harris 

Evansia 2: 44. 1985. Zignoella nyssaegena Ellis & Everhart, Proc. Acad. Sci. Nat. Philadelphia 

1893. 137. 1893. Ditremis nyssaegena (Ellis & Everhart) R. C. Harris, Some Florida Lichens 3. 

1990. Type. NEW JERSEY. [Gloucester County:] Newfield, on Nyssa multiflora, 25 Nov 1892, Elis 

(NY, holotype). 

Apiospora polypori Ellis & Everhart, N. Amer. pyrenomyc. 33. 1892. Type. DELAWARE. Newark, 

on old Fomes applanatus, 7 Nov 1890, Commons 1713 (NY, holotype), syn. nov. 

Melanopsamma corticola Ellis & Everhart, Proc. Acad, Nat. Sci. Philadelphia 1893: 129. 1893. 

Type. NEW JERSEY. [Gloucester County:] Newfield, on outer bark of Quercus alba, 7 Jul 1890, 

Ellis (NY, holotype), syn. nov. 

Thelidium juistense Erichsen, Mitt. Inst. Allg. Bot. Hamburg 10: 403. 1939. Anisomeridium 

juistense (Erichsen) R. C. Harris in D. Hawksw. et al., Lichenologist 12: 106. 1980. Type. 

GERMANY. Insel Juist, on Sambucus nigra, 23 Jun 1937, Erichsen (US, isotype). 

Arthopyrenia willeyana R. C. Harris, Michigan Bot. 12: 16. 1973. Type. MICHIGAN. Iosco County: 

N of Tawas City, on Populus tremuloides, 18 Sep 1965, Harris 818 (MSC, holotype; B, BM, 

CAN, COLO, DUKE, H, LD, MIN, NY, S, TNS, UPS, US, V_zda, WIS, isotypes). 

Apiospora polypori is a year older than Zignoella nyssaegena and I believe that Margaret Barr plans to 

take up the epithet. However, since I have already changed the name of this taxon so many times it is almost 

embarassing, since this is such a weedy species there are probably yet older epithets hidden in equally 

improbable genera, and since there is some possibility that the species may be segregated out of 

Anisomeridium, I plan to wait until the dust settles before accepting this or any further nomenclatural change. 

Anisomeridium nyssaegenum does not fit neatly into either of the main groups in Anisomeridium (see generic 

discussion above). It also has beaked pycnidia which have been given the illegitimate generic name 

Sarcinulella Sutton & Alcorn. There are several other species with beaked pycnidia but it remains to be studied 

whether a group so defined can be given generic recognition. 

Anisomeridium phaeospermum R. C. Harris, sp. nov. 

Species unica ostiolo apicali et ascosporis fuscis, granulis ornatis, 16-20 × 8-9 µm. 


along Florida Trail N of hwy., 25°50'N, 81°03'W, Taxodium swamp, on Taxodium, 9 Dec 1992, Harris 30189 

(NY, holotype). 

Thallus white, endophloeodal, UV+ yellow (lichexanthone). Ascomata mostly immersed, tip exposed, 

subglobose, sometimes with clypeus laterally expanded, ca. 0.5 mm diam. Asci cylindrical, ca. 140-150 × 15- 

16 µm, with eight uniseriate spores. Ascospores brown, elliptical, slightly constricted at the ± median septum, 

granular ornamented, 16-20 × 8-9 µm; conidia not found. 

The other species of Anisomeridium with brown ascospores differ in having an eccentric ostiole. 

Anisomeridium phaeospermum is known only from the type collection. 

Anisomeridium quadrococcum R. C. Harris, sp. nov. 

Thallo dilute griseo UV-, ostiolo apicale, ascosporis 3-septatis, laevibus, 21-24 × 5.5-8 µm et 

microconidiis oblongis, 3-4 × 1.5 µm. 

Type. FLORIDA. Columbia County: Ichetucknee Springs State Park, Fig Island, 29°58'N, 82°45'W, moist 

hardwoods on small limestone island, on decorticate wood, 2 Dec 1992, Buck 22601 (NY, holotype; FLAS, 

isotype). 

Thallus pale gray, endophloeodal, UV-. Ascomata ca. ½-¾ immersed, globose, ca. 0.2-0.25 mm diam.; 

ostiole apical; melanized wall entire or lacking at base; clypeus not expanded laterally. Asci cylindrical to 

122

subclavate, 180-220 × 45-55 µm, with eight biseriate spores. Ascospores 4-celled, end cells smaller, fusiform, 

not constricted at septa, smooth, 21-26 × (5.5-)6.5-8 µm; microconidia oblong, 3-4 × 1.5 µm; macroconidia not 

found. 

Corticolous species of Anisomeridium with 4-celled ascospores are rare, A. quaternarium has smaller 

ascospores and A. nyssaegenum has mostly 2-celled, only tardily 4-celled ascospores. 

Additional specimen. GEORGIA. Charleton County: Okefenokee National Wildlife Refuge, Chesser Island, 

15 Nov 1989, Wetmore 64357 (MIN). 

Anisomeridium quaternarium (R. C. Harris) R. C. Harris, comb. nov. 

Ditremis quaternaria R. C. Harris, Some Florida Lichens 34. 1990. Type. LOUISIANA. Plaquemines 

Parish: Pointe a la Hache, on trunk of Populus, 5 Jun 1976, Sérusiaux 1708 (NY, holotype). 

Anisomeridium subprostans (Nyl.) R. C. Harris 

in Tucker & Harris, Bryologist 83: 4. 1980. Verrucaria subprostans Nyl., Expos. syn. pyrenocarp. 

56. 1858. Arthopyrenia subprostans (Nyl.) Müll. Arg., Flora 66: 317. 1883. Ditremis subprostans 

(Nyl.) R. C. Harris, Some Florida Lichens 34. 1990. Type. SOUTH CAROLINA. On Taxodium, 

Ravenel, comm. Tuckerman 75, (H-NYL Pm. 7378, holotype?). 

Anisomeridium tamarindi (Fée) R. C. Harris 

in Tucker & Harris, Bryologist 83: 4. 1980. Verrucaria tamarindi Fée, Essai crypt. écorc. suppl. 

85. 1837. Porina tamarindi (Fée) Müll. Arg., Mém. Soc. Phys. Genève 30: 24. 1888. Ditremis 

tamarindi (Fee) R. C. Harris, Some Florida Lichens 34. 1990. Type. "ad cortices Tamarindi 

officinalis L. Antillarum" (probably GUADELOUPE, see Fée, loc. cit, p. 165). (G, lectotype; G, H- Nyl 

748. isolectotypes). 

Verrucaria vaga Nyl., Expos. syn. pyrenocarp. 55. 1858. Porina vaga (Nyl.) Müll. Arg., Hedwigia 

31: 287. 1892. Type. COMORO ISLANDS. Mayotte, Boivin (H-Nyl 750, holoype). 

Porina elaeidis Vainio in Hiern et al., Cat. Afr. pl. 2(2): 458.1901. Type. ANGOLA. Ambriz: Ad 

Elaeidis guineensis truncum, Nov 1853, Welwitsch 360 (BM, isotype). 

Tomasellia (Syngenosorus) africana Zahlbr., Ann. Cryptog. Exot. 5: 203. 1932. Type. SOUTH 

AFRICA. Knysna, v. d. Byl 749 (LD, isotype). 

Anisomeridium terminatum (Nyl.) R. C. Harris, comb. nov. 

Verrucaria terminata Nyl., Bull. Soc. Linn. Normandie sér. 2, 7: 180. 1873. Arthopyrenia terminata 

(Nyl.) Müll. Arg., Flora 73: 345. 1890. Ditremis terminata (Nyl.) R. C. Harris in Egan, Bryologist 

90: 163. 1987. Type. ANDAMAN ISLANDS. 1867, Kurz 23 (H-Nyl 725, lectotype; BM, M, 

isolectotypes). 

Verrucaria interspersa Nyl., Bull. Soc. Linn. Normandie sér. 2, 7: 181. 1873; Porina interspersa 

(Nyl.) Zahlbr., Cat. Lich. Univ. 1: 387. 1922. Anisomeridium interspersum (Nyl.) Makhija et al., 

J. Econ. Tax. Bot. 18: 544. 1994. Type. ANDAMAN ISLANDS. 1867, Kurz 3 (H-Nyl 726, lectotype). 

Arthopyrenia (sect. Polymeridium) glaucina Müll. Arg., Bot. Jahrb. Syst. 6: 406. 1885. Type. 

CUBA. Wright, Lich. Cub. ser. 2: 647 (G, holotype; US, isotype) . 

Tomasellia queenslandica Müll. Arg., Australas. Ass. Adv. Sci. Rep. 6: 460. 1895. Type. 

AUSTRALIA. Queensland, 1887, Knight 46 (G, holotype) . 

Arthopyrenia anacardii Vainio, Ann. Acad. Sci. Fenn. ser. A, 6(7): 209. 1915. Type. GUADELOUPE. 

Basse Terre, sur un Anacardium occidentale, Duss 498 (TUR-Vainio 32131, holotype; NY, 

isotype). 

Arthopyrenia insularis Vainio, Ann. Acad. Sci. Fenn. ser. A, 6(7): 208. 1915. Type. WEST INDIES. 

Insulis Danicis, 1905-6, Boergesen (TUR-Vainio 32116, holotype). 

Arthopyrenia subinsularis Vainio, Ann. Acad. Sci. Fenn. ser. A, 6(7): 208. 1915. Type. WEST 

INDIES. St. Croix, Mount Eagle, 20 Jan 1906, Boergesen (TUR-Vainio 32117, holotype). 

Verrucaria terminata and V. interspersa were published simultaneously. I did not formally 

123

select between them in "Some Florida Lichens" but the choice was implied in the citation of 

synonymy. The choice of the name Verrucaria terminata is formally endorsed here. 

Anisomeridium tuckerae R. C. Harris 

in Tucker & Harris, Bryologist 83: 4. 1980. Ditremis tuckerae (R. C. Harris) R. C. Harris, Some 

Florida Lichens 35. 1990.Type. LOUISIANA. East Baton Rouge Parish: hardwood forest at Burden 

Plantation, Essen Lane, Baton Rouge, Tucker 10003 (LSU, holotype) . 

MONOBLASTIA Riddle 

1. Ostiole strongly eccentric to lateral ........................................................................................................... 2 

2. Ascospores 30 µm long or less, 8/ascus, very strongly ornamented to 

rather strongly ornamented ................................................................................................................... 3 

3. Ascospores globose to long-ellipsoid................................................................................................ 4 

4. Ascospores ellipsoid to long-ellipsoid, 18-24 × 8-10 µm, 

ornamentation moderately granular/papillate; Puerto Rico, 

Mississippi ........................................................................................[M. borinquensis R. C. Harris] 

4. Ascospores globose to oblong or rarely ellipsoidal (in the same 

ascoma), 11-15 µm in diam. or 14-18 × 7-10 µm; southern Coastal 

Plain, French Guiana, French Antilles................................................................. M. rappii Zahlbr. 

3. Ascospores cylindrical to fusiform, 24-30 × 8-9 µm; Suwanee 

County ............................................................................................................. M. buckii R. C. Harris 

2. Ascospores over 30 µm long, weakly ornamented ............................................................................... 5 

5. Ascospores initially 8/ascus, often 4-6 aborting, 

32-36(-50) × 15-17(-20) µm; Cuba .....................................................................[M. palmicola Riddle] 

5. Ascospores initially 4/ascus, occasionally 1-2 aborting, 

45-50(-67) × 17-21 µm; Dominican Republic .......................................[M. quisqueyana R. C. Harris] 

1. Ostiole apical to slightly eccentric ............................................................................................................. 6 

6. Growing on bark .................................................................................................................................... 7 

7. Ascomata pallid, not melanized; ascospores ornamented 

with short spines, 9-13 × 7-9 µm; French Guiana, Papua 

New Guinea....................................................................................................... [M. pellucida Aptroot] 

7. Ascomata dark brown-black, melanized; ascospores 

ornamented with low, flat warts, 32-45 × 17-22 µm; 

Collier County..............................................................................................M. cypressi R. C. Harris 

6. Growing on limestone; ascospores strongly warted, 19-25(-28) × 

8-11 µm; Cuba, Haiti, Puerto Rico............................................ [M. echinulospora (Riddle) R. C. Harris] 

124

Monoblastia borinquensis R. C. Harris, sp. nov. 

NOTES 

Ascomatis ostiolis valde excentricis vel lateralibus et ascosporis octonis, ellipsoideis, verruculosis, 18-24 

× 8-10 µm. 

Type. PUERTO RICO. Distr. Arecibo, ca. i km N of ciales on Hwy. 149, 220 m, humid forest over limestone in 

haystack hills, 11 Jun 1988, Harris 22615 (NY, holotype). 

Thallus olivaceous or grayish, not corticate, UV-. Ascomata flask-shaped with subspherical body and 

projecting ostiolar neck, 0.5-0.7 mm in diam. (without neck), ca. ¾ immersed; ostiole strongly eccentric to 

lateral. Asci cylindrical with 8 uniseriate ascospores. Ascospores ellipsoidal, with verruculose to papillate 

ornamentation, 18-24 × 8-10(-11) µm; microconidia oblong, 3-4 1.5 µm. 

Monoblastia borinquensis is close to M. buckii but differs in shorter, broader ascospores. With additional 

material this difference may prove meaningless. The ascospore ornamentation of the Mississippi collection is 

weaker than in the type but I can find no other differences. 

Additional specimen. MISSISSIPPI. Sharkey County: Delta National Forest, along road #707, 9 mi NE of 

Valley Park, Jul 1978, Johnson s.n. (NY). 

Monoblastia cypressi R. C. Harris, sp. nov. 

Ascomatis ostiolis apicalibus et ascosporis octonis, oblongi-ellipsoideis, verrucis latis, 32-47 × 17-22 µm. 


along Florida Trail N of highway, Taxodium swamp, on Taxodium, 9 Dec 1992, Harris 30194 (NY, holotype). 

Thallus whitish, thin (bark showing through), not corticate, UV-. Ascomata subspherical with apical 

ostiolar papilla, 0.7-1.0 mm diam., ¾ to entirely immersed; ostiole apical. Asci cylindrical with eight uniseriate 

ascospores. Ascospores oblong-ellipsoidal with strong oranamentation of low, broad warts, 31-47 × 17-22 

µm; microconidia and macroconidia not found. 

Monoblastia cypressi has the largest ascospores of the eight-spored species. It is known only from the 

type. 

Monoblastia quisqueyana R. C. Harris, sp. nov. 

Ascomatis ostiolis valde excentricis et ascosporis quaternis, oblongi-ellipsoideis, leniter papillatis, 45-67 × 

17-22 µm. 

Type. DOMINICAN REPUBLIC. Prov. La Altagracia: 1.2 km S of El Macao along beach road, 18°46'N, 68°31'W, 

sea level, red mangrove swamp, on Rhizophora mangle, 22 Jan 1991, Harris 26612 (NY, holotype). 

Thallus pale gray, matte, not corticate, UV-. Ascomata ± spherical, occasionally with slightly projecting 

ostiolar neck, 0.7-1.0 mm diam., ca. ½-¾ immersed; ostiole strongly eccentric. Asci cylindrical with 4 

uniseriate ascospores. Ascospores oblong-ellipsoid with weak papillate ornamentation, 45-67 × 17-22 µm; 

microconidia and macroconidia not found. 

Other species of Monoblastia may have fewer than eight spores per ascus due to abortion (young asci 

with eight spores always present) but as far as I can tell M. quisqueyana is the only one which has only four 

initially. It is known only from the type. 

125

MUSAESPORA Aptroot & Sipman 

Aptroot and Sipman placed Musaespora in the Aspidotheliaceae (= Thelenellaceae) but the asci do not 

agree with those of that family and Musaespora seems better assigned to the Monoblastiaceae. In fact there 

are several taxa with corticate thalli that I am carrying in Anisomeridium which, when that genus is broken up, 

may be appropriately placed in Musaespora. I have seen material of an additional, probably undescribed, 

corticolous species with campylidia from Costa Rica. I need to go back and search for campylidia on the thalli 

of some other species which have a similar ascospore and ascus type, although I am not completely 

convinced a campylidiate anamorph alone should determine generic status. 

Musaespora epiphylla (R. Sant.) R. C. Harris, comb. nov. 

Astrothelium epiphyllum R. Sant., Symb. Bot. Upsal. 12(1): 131. 1952. Type. SUMATRA. Oostkust 

v. Sumatra, near Besitang, (near boundary of Atjeh), alt. c. 30 m, 1926, Palm 17b (UPS, 

holotype). 

?Musaespora multilocularis Aptroot & Sipman, Lichenologist 25: 127. 1993. Type. PAPUA NEW 

GUINEA (not seen). 

Musaespora gigas (Zahlbr.) R. C. Harris, comb. nov. 

Mycoporellum gigas Zahlbr., Ann. Crypt. Exot. 1: 117. 1928. Type. INDONESIA. Java: Pati, 

Seubert (US, isotype). 

Musaespora corticola Aptroot & Sipman, Lichenologist 25: 127. 1993. Type. INDONESIA. West 

Java (not seen). 

APPENDIX 

ANISOMERIDIUM, A GENERAL KEY TO SPECIES 

WARNING. The following key is only a very rough approximation. Anisomeridium s. lat. is a very speciose 

taxon and perhaps only 50-75% of the species may be included here. I am well aware illustrations are a vital 

necessity but time and money simply have not been available. Much more data must be gathered on conidia 

and the user is informed that in many cases conidia are known for a taxon from only a single instance and 

may not belong to the species to which they are assigned. Repeated co-occurrences can provide some 

confidence but culture from ascospores is the only definitive proof. The utility as characters of lichexanthone 

and ascospore ornamentation also needs to be validated. 

1. Growing on bark or wood .......................................................................................................................... 2 

2. Ascospores 2-celled; ostiole apical or eccentric ................................................................................... 3 

3. Thallus endophloeodal, not corticate, usually dull, usually whitish or grayish, 

less commonly greenish; ascomata immersed or not...................................................................... 4 

4. Thallus UV-................................................................................................................................... 5 

5. Ostiole apical............................................................................................................................ 6 

6. Asci with 8 spores ............................................................................................................... 7 

7. Ascospores 35 µm or less in length ............................................................................... 8 

8. Ascospores smooth [a complex of intergrading species 

containing the type of Anisomeridium]........................................................................ 9 

126

9. Ascospores biseriate or irregularly arranged in ascus ......................................... 10 

10. Ascospores less than 8 µm broad.................................................................. 11 

11. Macropycnidia not beaked ....................................................................... 12 

12. Ascospores relatively long, mostly over 

15 µm long........................................................................................... 13 

13. Ascospores remaining 2-celled; microconidia 

mostly globose ............................................................................. 14 

14. Asci slender, cylindrical or narrowly 

clavate ..................................................................................... 15 

15. Microconidia elliptical, 2.5-1.5 × 2 µm; 

macroconidia ± oblong, 5-7 × 2.5-3 

µm; ascospores narrowly ovoid, with 

weakly submedian septum; 12-18(-21) 

× 4.5-6.5 µm; neotropical........................................................ 

........................................A. subprostans (Nyl.) R. C. Harris 

15. Microconidia globose, ca. 2 µm diam.; 

macroconidia elliptical, 4-7 × 2.5-3.5 

µm; ascospores 15-22(-24) × 4.5- 

5.5(-6) µm, fusiform, often slightly 

bent; pantropical/subtropical ................................................ 

............................................ A. tamarindi (Fée) R. C. Harris 

14. Asci broader, ovoid or obovoid, broadly 

cylindrical or narrowly ovoid or 

narrowly obovoid .................................................................... 16 

16. Thallus poorly developed, 

endophloeodal................................................................... 17 

17. Ascospores over 5 µm in width ................................... 18 

18. Ascospore wall usually 

unevenly thickened; 

ascospores ovoid, with 

markedly submedian 

septum, 14-18 × 6-8 µm; 

microconida globose, ca. 2 µm 

diam., macroconidia oblong or 

ovoid, 5-7 × 2.5-3 µm; 

pantropical/subtropical .............................................. 

A. anisolobum (Müll. Arg.) Aptroot 

18. Ascospore wall relatively thin, 

even; ascospores 18-22 × 

5.5-7 µm, ± fusiform, often 

slightly bent, with slightly 

submedian septum; 

127

128 

microconidia globose or 

elliptical, ca. 2 µm diam. or 

2-3 × 1.5-2 µm; neotropical ....................................... 

A. excaecariae (Müll. Arg.) R. C. Harris 

17. Ascospores 12-17(-19) × 3.5-4.7 

µm; microconidia narrowly ovoid, 

4-5 × 1-1.3 µm; macroconidia 

oblong, 9-10.5 × 2-2.5 µm; 

Scotland & Wales............................................................ 

A. viridescens (Coppins) R. C. Harris 

16. Thallus well developed, epiphloeodal; 

ascospores 12-18 × 5-7.5 µm; 

macroconidia globose, ca. 2 µm diam.; 

microconidia globose, 2.5-4 µm diam.; 

Illinois, Massachusetts, Ohio................................................. 

A. leucochlorum (Müll. Arg.) R. C. Harris 

13. Ascospores 14-20 × 4.5-6 µm, ovoid, with 

submedian septum, often becoming 4-celled; 

micropycnidia beaked ("Sarcinulella"); 

microconidia oblong or narrowly elliptical, 

2-3 × 1-1.5 µm; temperate North America 

and Europe........................................................................................ 

................................ A. nyssaegenum (Ellis & Everhart) R. C. Harris 

12. Ascospores relatively short, less than 15 µm ..................................... 19 

19. Septum weakly submedian to median ......................................... 20 

20. Ascospores 10-18 × 4.5-7 µm, broadly 

elliptical to ovoid, rarely narrowly ovoid, with 

septum slightly submedian or ± median; 

microconidia oblong or elliptical, 2.5-3 × 

1.5-2 µm or globose, ca. 2 µm diam.; 

macroconidia elliptical or oblong, 5-7 × 

3-4 µm; cosmopolitan.................................................................. 

................................................. A. biforme (Borrer) R. C. Harris 

20. Ascospores 9-13(-15) × 4-5 µm, ovoid, 

with septum median to slightly submedian; 

ascomata small 0.2-0.3(-0.4) mm diam.; 

microconidia oblong or elliptical, 3-4 × 1.5-2 

µm; Florida ................................ A. albisedumi (Nyl.) R. C. Harris 

19. Ascospore septum markedly submedian; 

ascospores ovoid, 12-15 × 6-8 µm; 

microconidia globose, ca. 2 µm; New 

Zealand ................................... A. subatomaria (Knight) R. C. Harris 

11. Macropycnida with a long, hair-like beak, extruding

conidia in a single row; macroconida oblong, 

± constricted in middle, 8-10 × 4-5 µm; 

ascospores ovoid, with submedian septum, 

12-17 × 5-6 µm; Hong Kong................................A. throwerae R. C. Harris 

10. Ascospores mostly more than 8.5 µm broad................................................. 21 

21. Ascospores 8-12 µm broad ..................................................................... 22 

22. Ascospores fusiform; septum slightly submedian 

or median ........................................................................................... 23 

23. Ascospores 22-28 × (7-)8-9.5 µm; Philippine 

Islands ................................. A. sphaerocarpum (Vainio) R. C. Harris 

23. Ascospores 26-32(-34) × 8.5-11.5 µm; Andaman 

Islands ..............................................A. subnexum (Nyl.) R. C. Harris 

22. Ascospores ovoid to broadly ovoid, with 

septum strongly submedian, 18-23(-28) × 

9-14 µm; microconidia globose, ca. 2 µm diam.; 

neotropical................................ A. leptospermum (Zahlbr.) R. C. Harris 

21. Ascospores 23-32 × 12-18 µm broad, ovoid to broadly 

ovoid, with strongly submedian septum; pantropical................................... 

...........................................................A. subnectendum (Nyl.) R. C. Harris 

9. Ascospores uniseriate.......................................................................................... 24 

24. Ascospores 10-18 × 4.5-7 µm, elliptical or ovoid, with 

septum slightly submedian or ± median; microconidia oblong 

or elliptical, 2.5-3 × 1.5-2 µm or globose, ca. 2 µm diam.; 

macroconidia elliptical or oblong, 5-7 × 3-4 µm; 

cosmopolitan ..................................................... A. biforme (Borrer) R. C. Harris 

24. Ascospores 17-22 × 9-9.5 µm, ovoid, with submedian 

septum; macroconidia oblong or elliptical, 5-6 × 2.5-3 µm; 

Philippine Islands ............................................A. consimile (Vainio) R. C. Harris 

8. Ascospores granular ornamented, ovoid to ± pyriform, with 

septum very markedly submedian, biseriate/irregular in ascus, 

26-33 × 16-18 µm; microconidia globose, ca. 2 µm diam.; 

Brazil...................................................................... A. adnexa (Müll. Arg.) R. C. Harris 

7. Ascospores 37-52 × 17-24 µm, broadly elliptical, with ± median septum, 

smooth?; New Zealand .......................................... A. magnospora (Knight) D. Hawksw. 

6. Asci with 2-4 spores .......................................................................................................... 25 

25. Ascospores 4/ascus, fusiform, thick-walled, with ± median 

septum, strongly ornamented, 55-85 × 20-28 µm; 

microconidia elliptical or oblong, 22-3 × 1.5 µm; Cuba, 

Puerto Rico, St. Eustatius ...................................... A. excellens (Müll. Arg.) R. C. Harris 

129

25. Ascospores 2-4/ascus, broadly elliptical, relatively thin-walled, 

with ± median, smooth? septum, 62-82 × (22-)27-33 µm; 

Jamaica?.................................................................. A. planiusculum (Nyl.) R. C. Harris 

5. Ostiole eccentric..................................................................................................................... 26 

26. Ascospores colorless, ornamented or smooth................................................................. 27 

27. Asci with 8 spores ....................................................................................................... 28 

28. Ascospores smooth................................................................................................ 29 

29. Ascospores 7-11 µm in width, fusiform; septum ± median ............................... 30 

30. Ascospores 30-35(-40) × 7-11 µm; microconidia elliptical, 

3 × 1.5-2 µm; macroconidia narrowly ovoid, 11-20 × 

5.5-7 µm; Andaman Islands, India ................................................................... 

.....................................A. complanatum (Makhija & Patwardhan) R. C. Harris 

30. Ascospores 17-29 × 7-8 µm ......................................................................... 31 

31. Ascospores 23-29 × 7.5-8 µm; microconidia 

elliptical, 2.5 × 1.5 µm; New Caledonia ...................................................... 

.......................................................A. polycarpum (Müll. Arg.) R. C. Harris 

31. Ascospores 17-23 × 7-9 µm; pycnidia not found; 

Florida ................................................. Anisomeridium sp. Harris 29392-A 

29. Ascospores 9-18 µm in width, elliptical or broadly ovoid................................... 32 

32. Ascospores 22-30(-33) × 9-14 µm, with slightly submedian 

septum; macroconidia broadly ovoid or elliptical, 4-6 × 

3-4 µm; Philippine Islands, New Caledonia, Andaman 

Islands ........................................................A. palavanum (Vainio) R. C. Harris 

32. Ascospores 32-40(-45) × 13.5-18 µm; microconidia 

subglobose, ca. 2-2.5 µm diam.; macroconidia elliptical or 

oblong, 5-7 × 2.5-4 µm; Florida .......................................A. griffinii R. C. Harris 

28. Ascospores granular ornamented .......................................................................... 33 

33. Ascospores fusiform to narrowly ovoid ............................................................. 34 

34. Ascospores 19-42 × 5.5-12 µm, fusiform, with septum 

± submedian; pantropcical/subtropical......................................................... 35 

35. Ascospores 26-35(-42) × 7.5-10(-12) µm................................................... 

........................................................ A. americanum (Massal.) R. C. Harris 

35. Ascospores 19-27(-30) × 5.5-7(-8) µm; 

microconidia oblong or elliptical, 2.5-3 × 

1.5-2 µm ................................................. A. terminatum (Nyl.) R. C. Harris 

34. Ascospores 18-21 × 7-9 µm, narrowly ovoid, with 

septum submedian; New Zealand..................................................................... 

.................................................................. A. subbiforme (Knight) R. C. Harris 

130

33. Ascospores broadly elliptical, with moderately submedian 

septum, 17-21 × 8-10 µm; Hawaiian Islands.......................................................... 

................................................................................A. rockii (Zahlbr.) R. C. Harris 

27. Asci with 2-4 spores .................................................................................................... 36 

36. Asci with (2-)4 spores; ascospores fusiform, constricted at 

± median septum..................................................................................................... 37 

37. Ascospores ornamented, 55-85 × 20-28 µm; microconidia 

elliptical or oblong, 2-3 × 1.5 µm; Cuba, Puerto Rico, 

St. Eustatius ................................................ A. excellens (Müll. Arg.) R. C. Harris 

37. Ascospores smooth, 52-68 × 14-20 µm; microconida 

± globose, ca. 2 µm diam.; Andaman Islands ....................................................... 

...........................................A. tarmugliense (Makhija & Patwardhan) R. C. Harris 

36. Asci with 2 spores; ascospores broadly elliptical or oblong, 

constricted at ± median septum, granular ornamented; 52-74 × 

21-27 µm; Trinidad, Cuba, Dominican Republic ......................................................... 

..........................................................................A. megalosprum (Vainio) R. C. Harris 

26. Ascospores brown, strongly granular ornamented, oblong-ovoid, with ± 

median septum, 27-36 × 9-11 µm; ascomata emergent, with aspect 

of A. inspersum; microconidia elliptical, 3 × 1.5 µm; 

Cuba...................................................................................Anisomeridium sp. Harris 14615 

4. Thallus UV+ yellow (lichexanthone) ........................................................................................... 38 

38. Ostiole apical; ascospore septum submedian or not .......................................................... 39 

39. Ascospores colorless, not ornamented.......................................................................... 40 

40. Ascospores over 8 µm in width, mostly uniseriate in ascus ...................................... 41 

41. Ascospores ± fusiform with one end often more rounded, with 

septum median or submedian, smooth, 24-33 × 8-12 µm; 

microconidia elliptical, 3-5 × 1.5-2 µm; macroconidia broadly 

ovoid to ovoid, thick-walled,15-22 × 9-16 µm; French Guiana, 

Jamaica, Florida, Seychelles.............................. A. ambiguum (Zahlbr.) R. C. Harris 

41. Ascospores elliptical to broadly elliptical, with septum ± median, 

25-30 × 9-13 µm; microconidia elliptical, 3-4 × 1.5-2 µm; 

New Caledonia, Queensland......................................A. consobrinum (Nyl.) Aptroot 

40. Ascospores less than 7 µm in width, biseriate in ascus ............................................ 42 

42. Macroconidia cylindrical, 7.5-12 × 2-2.5 µm; microconidia 

elliptical to subglobose, 2.5-3 × 1.5-2 or ca. 2 µm diam.; 

ascospores fusiform, with slightly submedian septum 15-20 × 

5-6.5 µm; Florida ...........................................A. tuckerae (R. C. Harris) R. C. Harris 

42. Macroconidia oblong, 6-8 × 2-2.5 µm; microconidia elliptical 

to subglobose, 2.5-3 × 1.5-2 µm; ascospores narrowly fusiform, 

131

with slightly submedian septum, 14-19 × 4-5.5 µm; São 

Tomé .....................................................................A. albidoatrum (Nyl.) R. C. Harris 

39. Ascospores brown, ornamented, elliptical to broadly 

elliptical, with median septum, 17-20 × 8-9 µm; 

Florida ...............................................................................A. phaeospermum R. C. Harris 

38. Ostiole eccentric; ascospores broadly fusiform, with submedian 

septum, smooth .................................................................................................................. 43 

43. Ascospores colorless .................................................................................................... 44 

44. Ascospores 13 µm or more in width......................................................................... 45 

45. Ostiole not ringed with orange pigment.............................................................. 46 

46. Ascospores (37-)42-53 × 13-17 µm, with markedly 

submedian septum; Dominican Republic, Puerto 

Rico ........................................................... Anisomeridium sp. Harris 20160-A 

46. Ascospores 30-40 × 13-16.5(-18) µm; microconidia 

oblong, 3-4 × 1.5-2 µm; India ........................................................................... 

..................................... A. flavopallidum (Makhija & Patwardhan) R. C. Harris 

45. Ostiole ringed with orange pigment, KOH+ purple; 

ascospores fusiform, with septum submedian, 37-52 × 

12-20 µm; microconidia oblong, 3-4 × 1.2-1.5 µm; 

macroconidia broadly elliptical to ovoid, 30-45 × 

15-20 µm; Florida ..................................................A. aureopunctatum R. C. Harris 

44. Ascospores 13 µm or less in width........................................................................... 47 

47. Ascomata exposed, with a broad, melanized clypeus; 

ascospores 20-27 × 8.5-11 µm; Cuba..................................................................... 

....................................................................A. angulosum (Müll. Arg.) R. C. Harris 

47. Ascomata mostly hidden, immersed, with poorly developed 

clypeus; ascospores 23-34 × 9-13 µm; Asia, Pacific............................................... 

...............................................................................A. ubiana (Vainio) R. C. Harris 

43. Ascospores brown, broadly elliptical, with ± median septum 

9-11 × 5-6.5 µm; microconidia oblong, 2-3.5 × 1-1.5 µm; 

South Africa...................................................................A. uniseriale (Zahlbr.) R. C. Harris 

3. Thallus mostly epiphloeodal, corticate, usually ± glossy, usually greenish or grayish; 

ascomata immersed (except A. truncatum) ................................................................................... 48 

48. Thallus (cortex or medulla) UV+ yellow (lichexanthone); ascospores mostly 

ovoid to broadly ovoid, constricted at markedly submedian septum, smooth.......................... 49 

49. Ascospores 45 × 18 µm or larger ........................................................................................ 50 

50. Medulla UV+ yellow; cortex UV-; ascospores (48-)55-65(-70) 

× 25-30(35) µm; microconidia elliptical, 3 × 1.5-2 µm; 

French Guiana, Trinidad, St. Vincent ................................A. infernale (Mont.) R. C. Harris 

132

50. Medulla UV-; cortex UV+ yellow; ascospores 45-57 × 18-26 µm; 

Puerto Rico..................................................................................... A. elegans R. C. Harris 

49. Ascospores 45 × 18 µm or smaller...................................................................................... 51 

51. Ascospores with septum median to slightly submedian................................................ 52 

52. Ascospores with rounded ends, broadly elliptical, 

(10-)12-15 × 7-9 µm; microconidia broadly elliptical, 

3 × 1.5-2 µm; Florida and Georgia ................................... A. biformoides R. C. Harris 

52. Ascospores with ends somewhat pointed, 31-41 × 13-17 µm; 

microconidia oblong or ovoid, 3-4 × 1.5-2 µm; India ................................................... 

......................................................... A. indicum (Makhija & Patwardhan) R. C. Harris 

51. Ascospores with septum markedly submedian, ends rounded, 

34-42 × 16-18 µm; ?microconidia globose, 3-4 µm diam.; 

Dominican Republic .......................................................Anisomeridium sp. Harris 16267 

48. Thallus UV-............................................................................................................................... 53 

53. Ascomata and pycnidia solitary.......................................................................................... 54 

54. Ostiole eccentric; ascospores smooth .......................................................................... 55 

55. Asci with eight spores.............................................................................................. 56 

56. Spores in a single row in the ascus.................................................................... 57 

57. Ascospores fusiform with pointed ends; septum median............................. 58 

58. Ascospores over 50 µm long.................................................................. 59 

59. Ascospores 84-117 × 22-27 µm; microconidia 

oblong, 3-5 × 1.5-2 µm; French Guiana and 

Trinidad .......................................................A. nemorosum R. C. Harris 

59. Ascospores 52-83 × 16-21 µm; microconidia 

oblong, 2.5-3.5 × 1.5 µm; India............................................................... 

................................A. immersum (Makhija & Patwardhan) R. C. Harris 

58. Ascospores 30-37 × 9-11 µm; microconidia 3-4 × 

1.5 µm; Cuba................................ A. glaucescens (Müll. Arg.) R. C. Harris 

57. Ascospores with rounded ends, broadly elliptical, with 

median septum, 31-46 × 14-21 µm; macroconidia 

broadly elliptical, 5.5-7.5 × 3-4 µm; Trinidad and 

Dominica ......................................................... Anisomeridium sp. Hale 38240 

56. Spores in two rows or irregularly arrranged in the ascus .................................. 60 

60. Ascospores ovoid, ends rounded, with septum markedly 

submedian, 40-45 × 15-19 µm; French Guiana ................................................ 

......................................................................... Anisomeridium sp. Buck 18364 

60. Ascospores broadly fusiform with ± acute ends, with 

median septum, 57-76 × 22-27 µm; New 

133

Caledonia .......................................................A. holopolium (Nyl.) R. C. Harris 

55. Asci with 1-4 spores ................................................................................................ 61 

61. Asci with 1 spore; ascospores fusiform, constricted at ± median 

septum, ends ± rounded, 200-225 × 52-62 µm; microconidia 

oblong, 3-4 × 1.5-2 µm; India .................................................................................. 

............................................. A. monosporum (Makhija & Patwarhan) R. C. Harris 

61. Asci with 4 spores; ascospores fusiform, constricted at ± median 

septum, ends pointed, 58-70 × 12-17 µm; Andaman Islands and 

Australia ...................................A. verrucosa (Makhija & Patwardhan) R. C. Harris 

54. Ostiole apical; ascospores smooth or ornamented....................................................... 62 

62. Ascomata emergent, flat-topped; thallus very well developed, 

greenish tan, Pyrenula-like; ascospores narrowly ovoid, with 

septum submedian, strongly granular oranamented, uniseriate in 

ascus; Brazil ..........................................................A. truncatum (Müll. Arg.) . C. Harris 

62. Ascomata completely immersed; thallus olivaceous, almost 

Clathroporina-like; ascospores broadly ovoid, ± constricted 

at the markedly submedian septum, smooth, subbiseriate in 

ascus, 42-52 × 20-24 µm; Sri Lanka .............................................................................. 

...............................................................................A. nidulans (Müll. Arg.) R. C. Harris 

53. Ascomata and pycnidia several under a single common clypeus; 

thallus thick, to 75 µm, with thick cortex, to 25 µm; ascospores 

fusiform, slightly constricted at weakly submedian septum, smooth, 

15-23 × 4.5-5.5 µm; microconidia elliptical, 3 × 1.5-2 µm; 

macroconidia elliptical, 5-6 × 2-2.5 µm; Dominican 

Republic ..................................................................................... A. stromaticum R. C. Harris 

2. Ascospores 4-celled; ostiole apical ..................................................................................................... 63 

63. Ascospores soon 4-celled ............................................................................................................. 64 

64. Ascospores 21-24 × 6-8 µml; microconidia elliptical, 3-4 × 1.5-2 µm; 

Florida and Georgia....................................................................... A. quadrococcum R. C. Harris 

64. Ascospores16-20 × 3.5-4.5 µm; microconidia ± globose, 

ca. 2 µm diam.; macroconidia oblong, 5-6 × 2-3 µm; 

Louisiana ................................................................... A. quaternarium (R. C. Harris) R. C. Harris 

63. Ascospores tardily 4-celled, ovoid, 14-20 × 4.5-6 µm; eastern 

North America .............................................................A. nyssaegenum (Ellis & Everh.) R. C. Harris 

1. Growing on rock, soil, liverworts or leaves .............................................................................................. 65 

65. Growing on rock or compacted soil (in part modified from McCarthy (1993).................................... 66 

66. Ascospores 2-celled..................................................................................................................... 67 

67. Growing on non-calcareous rock ............................................................................................ 68 

68. North Temperate taxa ........................................................................................................ 69 

134

69. Growing in aquatic or semiaquatic habitats................................................................... 70 

70. Thallus well developed, often with black hypothallus; ascospores 

ovoid, with septum submedian, smooth, (11-)13-18 × 5.5-7 µm; 

microconidia oblong, 4-5 × 1.5-2 µm; eastern North America, 

Europe...............................................................A. carinthiacum (Steiner) R. C. Harris 

70. Thallus scant or not evident; ascospores tardily 4-celled 

(see below)................................................................. A. distans (Willey) R. C. Harris 

69. Growing on dry rock; ascospores ovoid, with submedian septum, 

smooth, 13-15 × 6-6.5 µm; Virgin Islands ......................................................................... 

.....................................................................................A. antillarum (Vainio) R. C. Harris 

68. Australasian/Pacific taxa .................................................................................................... 71 

71. Thallus 60-140 µm thick, pale gray; perithecia 0.17-0.32 mm diam.; 

clypeus 30-45 µm thick; New Zealand .............................................................................. 

..............................................................................A. laevigatum (McCarthy) R. C. Harris 

71. Thallus 30-60 µm thick, pale gray-green to pale brown gray; 

perithecia 0.24-0.44 mm diam.; clypeus 45-65 µm thick; 

Hawaiian..................................................................A. pacificum (McCarthy) R. C. Harris 

67. Growing on limestone.............................................................................................................. 72 

72. Ascospores 20-27 × 6-8.5 µm, narrowly ovoid to fusiform, 

with submedian septum, smooth; ascomata solitary; microconidia 

elliptical 3-5 × 1-1.5 µm; macroconidia broadly elliptical,5-6 × 4-4.5 µm; 

Florida and Puerto Rico..................................................... A. finkii (R. C. Harris) R. C. Harris 

72. Ascospores 11-13 × 3.5-4.5 µm, fusiform, with septum ± median, 

smooth; ascomata initially solitary, later forming in clusters; microconidia 

oblong, 3 × 1.5 µm; Puerto Rico..........................................................A. gregale R. C. Harris 

66. Ascospores 4-celled..................................................................................................................... 73 

73. Ascospores tardily 4-celled, ovoid, 16-22 × 6.5-8 µm; microconidia 

narrowly elliptical, 4-5 × 1.5-2 µm; macroconidia ovoid or narrowly 

elliptical, 5-7 × 2.5-3 µm; eastern North America. ....................... A. distans (Willey) R. C. Harris 

73. Ascospores soon 4-celled, broadly fusiform, 21-36 × 7.5-13 µm; 

microconidia bacilliform 2.5-4 × 0.7 µm; Australia........................................................................ 

..........................................................................................A. australiense (McCarthy) R. C. Harris 

65. Growing on bryophytes or leaves ...................................................................................................... 74 

74. Growing on leaves........................................................................................................................ 75 

75. Ostiole apical........................................................................................................................... 76 

76. Ascomata dark; ascospores elliptical, with ± submedian septum, 

smooth, 13-14.5 × 4.5-5.5 µm; pycnidia with apical protruberances 

in a ± stellate arrangement; macroconidia? 4.5-5 × 3.5-4.5 µm; 

pantropical...................................................................................A. foliicola R. Sant. & Tibell 

76. Ascomata pale; ascospores ovoid, septum slightly supramedian, 

14-20 × 5.5-6.5 µm; Tanzania .........................................................Porinula tanzanica V_zda 

135

(I have not seen material. Pale ascomata are known in Monoblastiaceae and 

have not been given generic recognition. In all probability this will be 

transferred to Anisomeridium.) 

75. Ostiole eccentric; ascospores elliptical, with septum ± median, 

30-40 × 10-12 µm; microconidia broadly elliptical or ovoid, 

4.5-5 × 2.5-3.5 µm; Zaire...................................................... A. epiphyllum (V_zda) R. C. Harris 

74. Growing on liverworts; ostiole apical; ascospores broadly fusifrom to 

subclavate, with ± median septum, smooth, 33-42 × 10-12.5 µm; 

macroconidia forming cirrhi?, embedded in gelatin, oblong, 

5-7 × 3.5-4.5 µm; Brazil ................................................. A. endobrya (Döbbler & Poelt) R. C. Harris 

NOTES 

Anisomeridium adnexum (Müll. Arg.) R. C. Harris, comb. nov. 

Arthopyrenia adnexa Müll. Arg., Flora 66: 304. 1883. Leiophloea adnexa (Müll. Arg.) Riedl, 

Sydowia 16: 266. 1962(1963). Type. BRAZIL. São Paulo: Apiaí, Sep, Puiggari 240 (G, lectotype, 

selected here), May, Puiggari 1007 (G, syntype). The other syntype, Glaziou 5094, is A. 

leptospermum. 

Verrucaria thelena var. albidior Nyl. 

Ann. Sci. Nat. Bot. sér. 4, 20. 254. 1863. Type. NICARAGUA. Verr. 22 ex hb. Tuckerman (H-Nyl 

714, lectotype, Hawskworth, 1985); HONG KONG. Verr. 21 ex hb. Tuckerman (H-Nyl 715, 

syntype). 

Hawksworth (1985) suggested the lectotype might belong in Anisomeridium. The structure of 

the ascoma and rather coarse physes suggest it is better included in Arthopyrenia. Its exact 

disposition awaits further work on tropical Arthopyreniae. The syntype is a Mycomicrothelia in 

the group with thickened endospore. Its disposition is also pending more study. 

Anisomeridium albidoatrum (Nyl.) R. C. Harris, comb. nov. 

Verrucaria albidoatra Nyl., Flora 69: 178. 1886. Arthopyrenia albidoatra (Nyl.) Zahlbr., Cat. lich. 

univ. 1: 300. 1921. Type. SÃO TOMÉ. Altit. 20 m, Moller (H-Nyl 827, lectotype (selected here); H-Nyl 

826, syntype). 

Anisomeridium americanum (Massal.) R. C. Harris, comb. nov. 

Arthopyrenia americana Massal., Ric. auton. lich. crost. 170, f. 341. 1852. Type. "Vive n'ell 

America sui tronchi degli alberi."[ex hb. Fee as Pyrenula nitida v. americana Fee] (VER, 

holotype). 

Verrucaria limitans Nyl., Flora 49: 295. 1866. Arthopyrenia limitans (Nyl.) Müll. Arg., Flora 6: 

306. 1883. Type. CUBA. [Wright] (H-Nyl 1714, holotype; M, isotypes). 

Arthopyrenia ceylonensis Müll. Arg., Flora 66: 289. 1883. Type. CEYLON. 1876, Thwaites 174 (G, 

lectotype; G, isolectotypes). 

Pleurotrema inspersum Müll. Arg., Bot. Jahrb. Syst. 6: 389. 1885. Ditremis inspersa (Müll. 

Arg.) Clements, The Genera of Fungi 173. 1909. Type. CUBA. Wright, Verr. Cub. 123 (H-Nyl 

1715, FH-Tuck 3948, US, isotypes) . 

Pleurotrema pyrenuloides Müll. Arg., Rep. Meetings Australas. Assoc. Advancem. Sci. 6: 462. 

1895. Ditremis pyrenuloides (Müll. Arg.) Makhija & Patwardhan, Biovigyanam 16: 22. 1990. 

Type. AUSTRALIA. Queensland, Knight 49 (G, lectotype, selected here). 

Didymella labiata Vainio, Hedwigia 38(Beibl.): 258. 1899. Type. GUADELOUPE. Basse Terre, 

Duss 503 (TUR-Vainio 32200, holotype). 

Arthopyrenia media Harmand, Bull. Soc. Sci. Nancy, sér. 3, 12: 130. 1911. Type. NEW CALEDONIA. 

Pionnier (DUKE, holotype). 

136

Anisomeridium angulosum (Müll. Arg.) R. C. Harris, comb. nov. 

Tomasellia (sect. Syngenosorus) angulosa Müll. Arg., Bot. Jahrb. Syst. 6: 397. 1885. Type. CUBA. 

Wright, Verr. Cub. 127 (G, holotype). 

Anisomeridium antillarum (Vainio) R. C. Harris, comb. nov. 

Arthopyrenia antillarum Vainio, Ann. Acad. Sci. Fenn. ser. A, 6(7): 207. 1915. Type. VIRGIN 

ISLANDS. St. John: Macombo, 20 Mar 1906, Boergesen (TUR-Vainio 32115). 

Anisomeridium australiense (McCarthy) R. C. Harris, comb. nov. 

Ditremis australiensis Mc Carthy, Austral. Syst. Bot. 5: 125. 1992. Type. AUSTRALIA. New South 

Wales: Newcastle region, 3 kn Se of Stroud Road, 1 km S of Stroud Mt., creek beside Ducks 

Hill road, on deeply shaded semi-aquatic and aquatic rhyolite, 22 Apr 1990, McCarthy 366 

(MEL, holotype). 

Ditremis bispora Makhija & Patwardhan, 

Biovigyanam 16: 14. 1990. Type. ANDAMAN ISLANDS. South Andaman:, Baratang Islands, 

Baludera, 23 Feb 1985, Sethy & Nagarkar 85.795 (AMH, holotype). 

= Arthopyrenia 

Anisomeridium carinthiacum (Steiner) R. C. Harris 

in Egan, Bryologist 90: 163. 1987. Arthopyrenia carinthiaca Steiner, Österr. Bot. Z. 63: 335. 1913. 

Type. AUSTRIA. Carinthia, near Klagenfurt, Steiner (US, isosyntype). 

Arthopyrenia dimidiata Fink in Hedrick, Mycologia 25: 306. 1934. Type. CONNECTICUT. Litchfield 

County: near Ellsworth, 1895, Green (MICH, holotype). 

Anisomeridium complanatum (Makhija & Patwardhan) R. C. Harris, comb. nov. 

Ditremis complanata Makhija & Patwardhan, Biovigyanam 16: 15. 1990. Type. ANDAMAN ISLANDS. 

South Andaman: Wimbeliganj, Kalatang, 16 Feb 1985, Sethy & Nagarkar 85.145 (AMH, 

holotype). 

Ditremis aggregata Makhija & Patwardhan, Biovigyanam 16: 13. 1990. Type. INDIA. Kerala, 

Sabrimalai Hills, Patnamathatta to Sabrimalai, 29 Jan 1983, Patwardhan & Sethy 83.470 (AMH, 

holotype). 

Ditremis harrisii Makhija & Patwardhan, Biovigyanam 16: 17. 1990. Type. INDIA. Karnataka, 

Sirsi-Kumtha Road, Ammanhalli, 12 Dec 1974, Nagarkar & Patwardhan 74.2619 (AMH, 

holotype). 

Ditremis karnatakensis Makhija & Patwardhan, Biovigyanam 16: 18. 1990. Type. INDIA. Karnataka, 

Sirsi-Kumtha Road, Devimane ghat, 12 Dec 1974, Kulkarni 74.2648 (AMH, holotype). 

Ditremis muelleri Makhija & Patwardhan, Biovigyanam 16: 20. 1990. Type. INDIA. Karnataka, 

Hosar Ghat, 26 Dec 1981, Nagarkar & Makhija 81.664 (AMH, holotype). 

Ditremis straminicolor Makhija & Patwardhan, Biovigyanam 16: 22. 1990. Type. ANDAMAN 

ISLANDS. South Andaman: Baratang Island, Wrafter's Creek, 21 Feb 1985, Patwardhan & Sethy 

85.534 (AMH, holotype). 

Anisomeridium complanatum is the most common species of the genus in India. The multiple 

names above are based on minor, ecologically induced variations. The species almost entirely 

replaces pantropical A. americanum in India and is barely seperable from it by the smooth 

ascospores and usually a very well developed, hard clypeus. The epithet "complanatum" has 

been selected from among the simultaneously published names since the complanate aspect 

of the ascomata due to the broad clypeus the most characteristic feature of the taxon. 

Anisomeridium consimile (Vainio) R. C. Harris, comb. nov. 

Arthopyrenia consimilis Vainio, Ann. Acad. Sci. Fenn. ser. A, 19(15): 14. 1923. Type. PHILIPPINE 

ISLANDS. Palawan: Taytay, on Pongamia mitis on seashore, Apr 1913, Merrill, Bur. Sci. 9053 

(TUR-Vainio 32110). 

137

Anisomeridium consobrinum (Nyl.) Aptroot 

Biblioth. Lichenol. 57: 21. 1995. Verrucaria consobrina Nyl., Ann. Sci. Nat. Bot. sér. 4, 15: 53. 

1861. Arthopyrenia consobrina (Nyl.) Müll. Arg., Flora 66: 317. 1883. Type. NEW CALEDONIA. 

Viellard 1838 (H-Nyl 1666, holotype; G, isotype). 

Anisomeridium elegans R. C. Harris, sp. nov. 

Thallo corticato, glauco, UV+ aureo, ostiolo eccentrico, ascosporis ovoideis, septis valde submedianis, 

laevibus, 51-57 × 23-26 µm 

Type. PUERTO RICO. Distr. Mayagüez: Maricao State Forest, just NE of Universidad Católica Estación 

Biologica, along Hwy 120 at km 15.8, along trail on hill top, ca. 700 m, semi-dry forest, 5 Jun 1988, Harris 

22280 (NY, holotype). 

Thallus corticate, epiphloeodal, glaucous green, glossy, UV+ yellow (lichexanthone), sometimes weakly 

so. Ascomata immersed, covered by thallus and only visible as a weakly raised bump, lageniform, 0.5-0.8 × 

0.4-0.5 mm; ostiole eccentric or lateral; wall entirely melanized or open below. Asci cylindrical to subclavate, 

ca. 200-300 × 40-55 µm, with eight subbiseriate spores. Ascospores ovoid, slightly constricted at markedly 

submedian septum, smooth, 45-57 × 18-26 µm; pycnidia not found. 

Anisomeridium elegans is related to A. infernale and A. nemorosum with similarly corticate thalli, but 

differs in size of ascomata and asci and in size and shape of ascospores. 

Additional specimen. PUERTO RICO. As type, ca. 600 m, moist forest on slope with tree ferns and palms, 

5 Jun 1988, Harris 22319 (NY). 

Anisomeridium endobryum (Döbbler & Poelt) R. C. Harris, comb. nov. 

Arthopyrenia endobrya Döbbler & Poelt, Pl. Syst. Evol. 138: 276. 1981. Type. BRAZIL. São 

Paulo: Ilha Comprida bei Cananeia, 15 Jul 1979, Kalb & Poelt (M, isotype). 

Anisomeridium epiphyllum (V_zda) R. C. Harris, comb. nov. 

Pleurotrema epiphyllum V_zda, Folia Geobot. Phytotax. 13: 99. 1978. Ditremis epiphylla (V_zda) 

Makhija & Patwardhan, Makhija & Patwardhan, Biovigyanam 16: 15. 1990. Type. ZAIRE (not 

seen). 

Anisomeridium excellens (Nyl.) R. C. Harris, comb. nov. 

Verrucaria excellens Müll. Arg., Bot. Jahrb. Syst. 6: 405. 1885. Type. CUBA. Wright, Verr. 

Cub. 79 (G, lectotype (selected here); UPS, US, isolectotypes); Verr. Cub. 78 (G, syntype). 

Anisomeridium flavopallidum (Makhija & Patwardhan) R. C. Harris, comb. nov. 

Ditremis flavopallida Makhija & Patwardhan, Biovigyanam 16: 16. 1990. Type. INDIA. Maharastra, 

Karnala, on the way to Ransai, 27 Aug 1974, Patwardhan & Nagarkar 74.431 (AMH, holotype). 

Anisomeridium foliicola R. Sant. & Tibell 

This lichen is apparently often produces only its very characteristic pycnidia. Santesson and Tibell 

suggested that they might be producing microconidia. From the size and shape of the conidia in Costa Rican 

material distributed by Lücking, I consider them to be macroconidia. This would suggest that since ascomata 

are rare, dispersal is via these macroconidia. It is also noteworthy that the pycnidia of A. foliicola have a well 

developed periphysoid plug which apparently relaeases the conidia single file. A similar feature is described 

for the possibly synonymous and therefore illegitimate pycnidial genus Compsosporiella Sanarkan & Sutton. 

Anisomeridium glaucescens (Müll. Arg.) R. C. Harris, comb. nov. 

138

Arthopyrenia glaucescens Müll. Arg., Bot. Jahrb. Syst. 6: 405. 1885. Type. CUBA. Wright, 

Verr. Cub. 77 (G, holotype; US, isotype). 

Anisomeridium gregale R. C. Harris, sp. nov. 

Thallo endolithico, glauco, UV-, ascomata aggregata, ostiolo apicale et ascosporis parvis, anguste 

ovoideis, 11-13 × 3.5-4.5 µm. 

Type. PUERTO RICO. Distr. Arecibo: ca. 1 km N of Ciales on Hwy. 149, ca. 220 m, humid forest over limestone 

in haystack hills, on limestone, 11 Jun 1988, Harris 22626 (NY, holotype) 

Thallus endolithic in limestone, glaucous, UV-. Ascomata complanate, ca. 0.3 mm diam., initially solitary, 

later forming small groups ca. 1.0 mm across (regeneration ?); ostiole apical; open below; clypeus spreading 

laterally. Asci cylindrical, ca. 55 × 8 µm, with eight biseriate spores. Ascospores narrowly ovoid, weakly 

constriicted at slightly submdian septum, smooth, 11-13 × 3.5-4.5 µm; microconidia elliptical or ovoid, 3 × 1.5 

µm, macroconidia not found. 

Microscopically A. gregale is similar to the corticolous A. albisedum but seems to be distinct in growing 

on rock, in aggregated ascomata and in spreading clypeus. It is known only from the type collection. 

Anisomeridium holopolium (Nyl.) R. C. Harris, comb. nov. 

Verrucaria holopolia Nyl., Flora 50: 196. 1867, nom. nud. Bull. Soc. Linn. Normandie sér. 2, 

2: 131. 1868. Type. NEW CALEDONIA. Loyalty Islands: Lifou, 1864, Deplanche (H-Nyl 1671, 

lectotype (selected here); H-Nyl 1670, syntype). 

Anisomeridium immersum (Makhija & Patwardhan) R. C. Harris, comb. nov. 

Ditremis immersa Makhija & Patwarhan, Biovigyanam 16: 17. 1990. Type. INDIA. Kerala, 

Sabrimalai Hills, Patanamthatta to Sabrimalai, 29 Jan 1983, Makhija & Kekre 83.431 (AMH, 

holotype). 

Additional specimen. INDIA. Karnataka, 38-48 km from Sirsi, Kumala-Sirsi Rd., Devimane 

Ghat, evergreen forest, 26 Feb 1977, Hale 48030 (US). 

Anisomeridium indicum (Makhija & Patwardhan) R. C. Harris, comb. nov. 

Ditremis indica Makhija & Patwardhan, Biovigyanam 16: 18. 1990. Type. INDIA. Maharastra, 

Karnala forest, 28 Aug 1974, Nagarkar & Prabhu 74.456 (AMH, holotype). 

Anisomeridium infernale (Mont.) R. C. Harris, comb. nov. 

Verrucaria infernalis Mont., Ann. Sci. Nat. Bot. sér. 3, 16: 68. 1851. Arthopyrenia infernalis 

(Mont.) Müll. Arg., Flora 68: 262. 1885. Type. FRENCH GUIANA. Leprieur 725 (PC, syntype). 

Pleurotrema anisomerum Müll. Arg., Flora 68: 251. 1885. Ditremis anisomera (Müll. Arg.) 

Makhija & Patwardhan, Biovigyanam 16: 14. 1990. Type. FRENCH GUIANA. Leprieur 1248 (G, 

holotype). 

Arthopyrenia infernalis var. rhychostoma Vainio, Proc. Amer. Acad. Arts Sci. 58: 147. 1923. Type. 

TRINIDAD. Sangre Grande, 1912-1913, Thaxter 73 (FH, isotype). 

Verrucaria clandestina sensu Mont., Ann. Sci. Nat. Bot. sér. 2, 19: 59. 1843. (non Verrucaria 

clandestina (Ach.) Mont. = Myriotrema). 

Trypethelium karnatakense R. C. Harris, nom. nov. 

Pleurotrema megaspermum Makhija & Patwardhan, Biovigyanam 16: 24. 1990. Type. INDIA. 

Karnataka, Agumbe, 16 Dec 1974, Kulkarni 74.2991 (AMH, holotype), non Trypethelium 

megaspermum Mont. 

Anisomeridium laevigatum (McCarthy) R. C. Harris, comb. nov. 

139

Ditremis laevigata McCarthy, Muelleria 8: 1: 1993. Type. NEW ZEALAND (not seen). 

Anisomeridium leptospermum (Zahlbr.) R. C. Harris, stat. et comb. nov. 

Arthopyrenia adnexa var. leptosperma Zahlbr. in Skottsberg, Nat. hist. Juan Fernandez 2: 320. 

1924. Type. CHILE. Juan Fernández, Masafuera, Quebrada de la Lobería, 300 m, auf alter 

Lumaborke, 17 Feb 1917, C. & I. Skottsberg (UPS, holotype). 

Anisomeridium leucochlorum (Müll. Arg.) R. C. Harris 

in Egan, Bryologist 90: 163. 1987. Arthopyrenia leucochlora Müll. Arg., F!ora 66: 287. 1883. 

Ditremis leucochlora (Müll. Arg.) R. C. Harris, Some Florida Lichens 33. 1990. Type. 

MASSACHUSETTS. New Bedford, on Acer, Willey (MICH, MIN, US, isotypes) 

Ditremis macrospora Makhija & Patwardhan 

Biovigyanam 16: 19. 1990. Type. INDIA. Agasthi Hills, Upper Kodayar, 24 Jan 1983, Sethy & 

Kekre 83.380 (AMH, holotype). 

= Arthopyrenia 

Ditremis macrospora R. C. Harris = Anisomeridium aureopunctatum 

Anisomeridium magnosporum (Knight) D. Hawksw. 

in Galloway, New Zealand J. Bot. 21: 195. 1983. Verrucaria magnospora Knight, Trans. Linn. 

Soc. Lond. 23: 99. 1860. Type. NEW ZEALAND (not seen). 

Verrucaria magnifica Nyl., Lich. Nov. Zel. 132. 1888. Microthelia magnifica (Nyl.) Müll. Arg., Bull. 

Soc. Bot. Belgique 31: 41. 1892. Type. NEW ZEALAND. Knight 349 (H-Nyl 882, holotype?). 

Anisomeridium megalosporum (Vainio) R. C. Harris, comb. nov. 

Didymosphaeria megalospora Vainio, Proc. Amer. Acad. Arts Sci. 58: 147. 1923. Megalotremis 

megalospora (Vainio) Aptroot, Nova Hedwigia 60: 338. 1995. Type. TRINIDAD. St. Ann' Valley, 

1912-13, Thaxter 99 (TUR-Vainio 32490, lectotype, Aptroot, 1995). 

Anisomeridium monosporum (Makhija & Patwardhan) R. C. Harris, comb. nov. 

Ditremis monospora Makhija & Patwardhan, Biovigyanam 16: 20. 1990. Type. INDIA. Karnataka, 

Kudremukha, 28 Jan 1980, Nagarkar & Daya Rade 80.291 (AMH, holotype). 

Anisomeridium nemorosum R. C. Harris, sp. nov. 

Thallo corticato, epiphloeodo, viridescente, UV-, ascomatis magnis, thallo tectis, ostiolo laterale et 

ascosporis magnis, fusiformibus, laevibus, 100-117 × 23-27 µm. 

Type. FRENCH GUIANA. St-Laurent-du-Moroni: Canton de Maripasoula, Commune de Saül, ca. 6 km N of Saül, 

vicinity of Eaux Claires, along Route de Bélizon N of Eaux Claires beyond third bridge, ca. 200 m, 3°37'N, 

53°12'W, moist tropical forest over quartz and granite, 4 Sep 1994, Harris 33592 (NY, holotype; B, CAY, US, 

isotypes). 

Thallus corticate, epiphloeodal, glossy, greenish, UV-. Ascomata immersed, covered by thallus and 

visible as slightly raised lageniform bumps, large, ca. 1.5-2.0 × 1.0 mm( including clypeus); ostiole lateral; 

clypeus melanized above, broadly expanded. Asci cylindrical, ca. 500 × 50 µm, with eight obliquely uniseriate 

spores. Ascospores fusiform, ends distinctly pointed, constricted at ± median septum, smooth, 84-117 × 22- 

27 µm; microconidia oblong, 3-5 × 1.5-2 µm; macroconidia not found. 

It is possible that this species is the same as Ditremis corticata Makhija & Patwardhan (Mycotaxon 31: 

468. 1988). However, I have not seen material of this taxon and considering Makhija and Patwardhan's track 

record, this may not even be an Anisomeridium. Additionally the ascospores are reported to be narrower, 

140

16.5-23 µm. As I have unpacked only a part of my French Guiana collections, I would guess that A. 

nemorosum will prove to be rather weedy on trunks in tropical moist forests. 

Additional specimens. FRENCH GUIANA. As type, Sentier Botanique, 200-300 m, 13-14 Nov 1990, Buck 

18822, 300-400 m, 7 Sep 1994, Harris 33819 (both NY). TRINIDAD. Cats Hill Road, ca. 6 mi from junction with 

Rio Claro-Guayaguare road, virgin lowland rain forest, ca. 200 ft., 18 Apr 1972, Hale 38195 (US). 

Anisomeridium nidulans (Müll. Arg.) R. C. Harris, comb. nov. 

Arthopyrenia nidulans Müll. Arg., Flora 68: 326. 1885. Type. SRI LANKA. Nieter, ex hb. Hampe, 

1877 (G, holotype). 

This is keyed out as having an apical ostiole. It is very difficult to determine the actual orientation 

and may actually be somewhat eccentric. 

Anisomeridium pacificum (McCarthy) R. C. Harris, comb. nov. 

Ditremis pacifica McCarthy, Muelleria 8: 3. 1993. Type. HAWAIIAN ISLANDS. Oahu (not seen). 

Anisomeridium palavanum (Vainio) R. C. Harris, comb. nov. 

Microthelia palavana Vainio, Ann. Acad. Sci. Fenn. ser. A., 15(6): 347. 1921. Type. PHILIPPINE 

ISLANDS. Palawan: Taytay, Apr 1913, Merrill, Bur. Sci. 9036 (BM, NY, isotypes). 

Arthopyrenia pongamiae Vainio, Ann. Acad. Sci. Fenn. ser. A, 19(15): 13. 1923. Leiophloea 

pongamiae (Vainio) Riedl, Sydowia 16: 271. 1962 [1963]. Type. PHILIPPINE ISLANDS. Palawan: 

Taytay, on Pongamia mitis, Apr 1913, Merrill, Bur. Sci. 9035 (TUR-Vainio 32111, holotype; BM, 

NY, isotypes). 

Ditremis andamanica Makhija & Patwardhan, Biovigyanam 16: 14. 1990. Type. ANDAMAN ISLANDS. 

North Andaman: Diglipur Range, Kalara, 4 Jan 1986, Sethy & Patwardhan 86.383 (AMH, 

holotype). 

Ditremis punctata Makhija & Patwarhan, Biovigyanam 16: 21. 1990. Type. ANDAMAN ISLANDS. 

Little Andaman: South Bay, 3 Mar 1985, Nagarkar & Patwarhan 85. 1026 (AMH, holotype). 

Anisomeridium planiusculum (Nyl.) R. C. Harris, comb. nov. 

Verrucaria planiuscula Expos. syn. pyrenocarp. 58. 1858. Arthopyrenia planiuscula (Nyl.) 

Zahlbr., Cat. lich. univ. 1:310. 1921. Type. JAMAICA? Ad cort. Quassia excelsa in hb. Fée (H- Nyl 


Anisomeridium polycarpum (Müll. Arg.) R. C. Harris, comb. nov. 

Pleurotrema polycarpum Müll. Arg., J. Bot. (Morot) 7: 109. 1893. Ditremis polycarpa (Müll. 

Arg.) Makhija & Patwardhan, Biovigyanam 16: 21. 1990. Type. NEW CALEDONIA. Cascade de la 

Ferme-Modèle, 1892, Balansa (G, holotype). 

Anisomeridium rockii (Zahlbr.) R. C. Harris, comb. nov. 

Pleurotrema rockii Zahlbr., Ann. Mycol. 10: 362. 1912. Ditremis rockii (Zahlbr.) Makhija & 

Patwardhan, Biovigyanam 16: 22. 1990. Type. HAWAIIAN ISLANDS. Lanai, Koela Valley, Rock 132 

(FH, isotype). 

Anisomeridium sphaerocarpum (Vainio) R. C. Harris, comb. nov. 

Didymella sphaerocarpa Vainio, Ann. Acad. Sci. Fenn. ser. A, 19(15): 14. 1923. Type. PHILIPPINE 

ISLANDS. Luzon: Benguet Province, Pauai, May 1909, Merrill, Bur. Sci. 6649 (TUR-Vainio 32297, 

holotype). 

Anisomeridium stromaticum R. C. Harris, sp. nov. 

141

Species distincta thallo crasso corticato, UV-, ascomatis pluribus subtus clypeo unico, pycnidiis pluribibus 

subtus clypeo unico, ascosporis fusiformibus septo submediano laevibus, 15-23 × 4.5-5.5 µm. 

Type. DOMINICAN REPUBLIC. Prov. Monte Cristi: El Morro de Monte Cristi, 237 m, limestone mesa by sea, 

19°54'N, 71°39'W, 8 Jan 1987, Harris 19534 (NY, holotype; JBSD, isotype). 

Thallus epiphloeodal, thick (to ca. 75 µm), corticate, gray, ± dull, UV-; cortex to 25 µm in places. 

Ascomata mostly immersed in bark, several under a single clypeus, ca. 0.3 mm diam., occasionally solitary, 

then larger, ca. 0.7 mm diam.; ostiole apical; wall not melanized; clypei often fusing in more extensive, black, 

irregular pseudostromata, 3 × 1-2 mm. Asci cylindrical, ca. 85 × 12 µm, with eight biseriate spores. 

Ascospores fusiform, slightly constricted at ± submedian septum, smooth, 15-23 × 4.5-5.5 µm; pycnidia as in 

ascomata, usually several under a single clypeus; microconidia elliptical, 3 × 1.5-2 µm; macroconidia elliptical, 

5-6 × 2-2.5 µm. 

This is a very peculiar species and I am not entirely sure that it is not an abnormal form of some other 

species. A somewhat similar situation occurs in the type of A. polycarpum which I interpreted as the 

ascomata growing in abnormal proximity. I may have been wrong and the type should be re-examined. 

However, in A. stromaticum the fact that both micropycnidia and macorpycnidia are clustered under a single 

clypeus lends support to interpreting this as the normal state for this species. The thick, corticate thallus is 

also distinctive and somewhat resembles that of A. glaucescens which differs in larger ascospores. 

Anisomeridium subatomarium (Knight) R. C. Harris, comb. nov. 

Verrucaria subatomaria Nyl. ex Knight, Trans. Proc. New Zealand Inst. 15: 356. 1883. 

Arthopyrenia subatomaria (Knight) Müll. Arg., Bull. Soc. Bot. Belgique 31: 40. 1892. Type. NEW 

ZEALAND. Knight 14 (H-Nyl 884, isotype?). 

Anisomeridium subbiforme (Knight) R. C. Harris, comb. nov. 

Verrucaria subbiformis Knight, Trans. Proc. New Zealand Inst. 8: 326. 1875. Arthopyrenia 

subbiformis (Knight) Müll. Arg., Bull. Soc. Bot. Belgique 31: 40. 1892. Type. NEW ZEALAND. 

Knight (G, M, UPS, isolectotypes?). 

Anisomeridium subnectendum (Nyl.) R. C. Harris, comb. nov. 

Verrucaria subnectenda Nyl., J. Linn. Soc., Bot. 20: 61. 1883. Arthopyrenia subnectenda (Nyl.) 

Zahlbr., Cat. lich. univ. 1: 327. 1922. Type. MALAYSIA. Malacca, 1864, Maingay 161 (H-Nyl 835, 

holotype; BM, isotype). 

Verrucaria anisomera Nyl., Flora 56: 364. 1876, nom. nud. "Type". CUBA. Wright, Verr. Cub. 105 

(H-Nyl 1673). 

Arthopyrenia xylogena Müll. Arg., Flora 66: 290. 1883. Leiophloea xylogena (Müll. Arg.) Riedl, 

Sydowia 16: 270. 1962 [1963].Type. BRAZIL. São Paulo: Apiaí, Puiggari 494 (G, holotype). 

Arthopyrenia exstans Müll. Arg., Rep. & Meetings Australas. Assoc. Advancem. Sci. 6: 451. 1895. 

Type. AUSTRALIA. Queensland, 1887, Knight 43 (G, holotype) 

Anisomeridium subnexum (Nyl.) R. C. Harris, comb. nov. 

Verrucaria subnexa Nyl., Bull. Soc. Linn. Normandie, sér. 2, 7: 181. 1873. Arthopyrenia subnexa 

(Nyl.) Müll. Arg., Hedwigia 30: 188. 1891. Type. ANDAMAN ISLANDS. 1867, Kurz 75 (H-Nyl 834, 

holotype; M, isotype). 

Anisomeridium tarmugliense (Makhija & Patwarhan) R. C. Harris, comb. nov. 

Ditremis tarmugliensis Makhija & Patwarhan, Biovigyanam 16: 22. 1990. Type. ANDAMAN ISLANDS. 

South Andaman: Wandoor, Tarmugli Island, Patwardhan & Nagarkar 85.1890 (AMH, holotype). 

Anisomeridium throwerae R. C. Harris, sp. nov. 

142

Species distincta macropycnidiis rostratis, rostro capillaceo ad 1.5 mm, macroconidiis oblongis in medio 

constrictis, 8-10 × 4-5 µm, ascosporis 12-17 × 5-6.5 µm. 

Type. HONG KONG. New Territories: near Cheung Uk, 21 oct 1977, Thrower 2900 (NY, holotype). 

Thallus gray green, endophloeodal, UV-. Ascomata immersed, subglobose, ca. 0.2 mm diam.; ostiole 

apical. Asci cylincrical, ca. 45 × 9 µm, with eight biseriate spores. Ascospores ovoid, slightly constricted at 

submedian septum, 12-17 × 5-6.5 µm; macropycnidia with a long, to 1.5 mm, beak through which the 

macroconidia are extruded in single file; macroconidia oblong, constricted in middle, 8-10 × 4-5 µm. 

The micropycnidia associated with A. nyssaegenum have a short beak. Similar long-beaked 

macropycnidia are known in Anisomeridium sp. Buck 22812 for which no teleomorph is present and 1 or 2 

undescribed species from Brazil. Also, from the illustration, it seems worth investigating Compsosporiella 

Sankaran & Sutton (Mycol. Res. 93: 1289. 1991) as a possible macropycnidial state of Anisomeridium. 

Anisomeridium truncatum (Müll. Arg.) R. C. Harris, comb. nov. 

Arthopyrenia truncata Müll. Arg., Flora 66: 289. 1883. Leiophloea truncata (Müll. Arg.) Rield, 

Sydowia 16: 273. 1962 [1963]. Type. BRAZIL. São Paulo: Apiaí, Puiggari 249 (G, holotype). 

Anisomeridium ubianum (Vainio) R. C. Harris, comb. nov. 

Arthopyrenia ubiana Vainio, Ann. Acad. Sci. Fenn. ser. A, 19(15): 12. 1923. Type. PHILIPPINES. 

Ubian Island, 12 Oct 1906, Merrill, Bur. Sci. 5403 (TUR-Vainio 32109, holotype; BM, isotype). 

Verrucaria consobrina var. glaucescens Nyl., Bull. Soc. Linn. Normandie, sér. 2, 2: 131. 1868. 

Arthopyrenia consobrina var. glaucescens (Nyl.) Zahlbr., Cat. lich. univ. 1: 323. 1922. Type. 

NEW CALEDONIA. Loyalty Islands: Lifou, 1864, Deplanche (H-Nyl 1664, lectotype (selected here); 

H-Nyl 1665, syntype?). 

Ditremis microcarpa Makhija & Patwarhan, Biovigyanam 16: 19. 1990. Type. ANDAMAN ISLANDS. 

Middle Andaman: Long Island Range, Elphiston Bay, 23 Dec 1985, Nagarkar & Patwardhan 

85.2219 (AMH, holotype). 

Anisomeridium uniseriale (Zahlbr.) R. C. Harris, comb. nov. 

Microthelia uniserialis Zahlbr., Ann. Cryptog. Exot. 5: 199. 1932. Pleurotrema uniseriale (Zahlbr.) 

D. Hawksw., Bull. Brit. Mus. Nat. Hist. Bot. 14(2): 171. 1985. Type. SOUTH AFRICA. Knysna, Hof 

van Eden, v. d. Byl 869 (LD, isotype). 

Hawksworth (1985) provided a modern description and photographs of habit and section of 

ascoma. 

Anisomeridium verrucosum (Makhija & Patwardhan) R. C. Harris, comb. nov. 

Ditremis verrucosa Makhija & Patwardhan, Mycotaxon 31: 469. 1988. Megalotremis verrucosa 

(Makhija & Patwardhan) Aptroot, Biblioth. Lichenol. 44: 126. 1991. Type. ANDAMAN ISLANDS (not 

seen). 

Anisomeridium viridescens (Coppins) R. C. Harris, comb. nov. 

Arthopyrenia viridescens Coppins, Lichenologist 20: 322. 1988. Type. SCOTLAND. Argyll Main, 

Taynult, Glen Nant prope Tailor's Leap, 27/01.27, alt. 45 m, ad corticem laevem Coryli, 3 

August 1980, B. J. Coppins 8031 (NY, isotype). 

When Coppins first sent a specimen of this species I studied it rather casually and moved on 

since it was not an American taxon but the presence of macroconidia in an Arthopyrenia was an 

anomaly which kept nagging in the back of my mind. After more careful study of the ascus and 

hamathecium, the anomaly was resolved as both proved to be of the Anisomeridium- type. 

STRIGULACEAE Fr. 

143

Syst. Orb. Veg. 1: 110. 1825. Type. Strigula Fr. 

Phyllobatheliaceae Bitter & Schilling, Hedwigia 67: 272, 291. 1927. Type. Phyllobathelium (Müll. Arg.) 

Müll. Arg., syn. nov. 

The Strigulaceae are constituted in a different manner from Harris (1975). Eriksson (1981) pointed out 

the family should be restricted excluding the Monoblastiaceae, Pleurotremataceae and Xanthopyreniaceae, a 

proposal supported by more recent studies. The Phyllobatheliaceae are placed in synonymy here as they 

have the same ascus type, apically free physes, ellipsoidal microconidia, septate macroconidia with gelatinous 

appendages and trentepohlioid photobiont. 

1. Ascomata solitary, not surrounded by pseudostromatic wart filled with dark 

granular material; pycnidia solitary; macroconidia usually with eccentric attachment 

and bipolar gelatinous appendages (except species with muriform ascospores?); 

on leaves, bark, wood or rock ........................................................................................................Strigula 

1. Ascomata 1-several, surrounded by a pseudostromatic wart filled with dark 

granular material; pycnidia aggregated; macroconidia with basal attachment 

and an apical gelatinous cap; on leaves ..........................................................................[Phyllobathelium] 

PHYLLOBATHELIUM (Müll. Arg.) Müll. Arg. 

Phyllobathelium is not known from North America. Following Santesson (1952) and Lücking (1992) I 

would recognize three foliicolous and one corticolous species. Mayrhofer (1987) added a corticolous and a 

saxicolous species. On this account I initially considered uniting Strigula and Phyllobathelium. However, 

examination of the species recognized by Santesson and Lücking revealed a number of characters that 

support recognition of both genera, pseudostromatic ascomatal warts, aggregated pycnidia and basally 

attached macroconidia (illustrated by V_zda, 1984) with an apical cap (not tapering appendages as in 

Strigula). The two non-foliicolous species recognized by Mayrhofer are transferred to Strigula below. Although 

Mayrhofer understood their general relationship, I suspect Strigula did not occur to him since in 1987 the 

present day plethora of muriform-spored species were as yet unrecognized. 

STRIGULA Fr. 

Raciborskiella Höhnel, Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturw. Cl., Abt. 1 ser. 1, 118: 1176. 1909. 

Type. Clypeolum talumae Racib. (= R. janeirensis (Müll. Arg.) R. Sant.). 

Dichoporis Clements, Genera of Fungi 40, 173. 1909. Type. Dichoporis schizospora (Vainio) Clements (= 

Strigula mediterranea Etayo). 

Diporina Clements, Genera of fungi 40, 173. 1909. Type. Diporina subsimplicans (Nyl.) Clements (= 

Strigula subsimplicans (Nyl.) R. C. Harris in Hafellner & Kalb). 

Discosiella H. & P. Sydow, Leafl. Philipp. Bot. 5: 1546. 1912, nom. illeg. (macroconidia) Type. Discosiella 

cylindrospora H. & P. Sydow (= Strigula sp.). 

Phylloporis Clements, Genera of Fungi 41, 173. 1909. Type. Phylloporis phyllogena (Müll. Arg.) Clements (= 

Strigula phyllogena (Müll. Arg.) R. C. Harris). 

Sagediomyces Ciferri & Tomaselli, Ist. Bot. Reale Univ. Reale Lab. Crittog. Pavia Atti ser. 5, 10: 30, 57. 

1953. Type. Sagediomyces affinis (Massal.) Ciferri & Tomaselli (= Strigula affinis (Massal.) 

R. C. Harris). 

Strigulomyces Ciferri & Tomaselli, Ist. Bot. Reale Univ. Reale Lab. Crittog. Pavia Atti ser. 5, 10, 34, 61. 

1953, nom. illeg. Type. Strigulomyces elegans (Fée) Ciferri & Tomaselli (= Strigula smaragdula Fr.) 

Shanoria Anahosur, Nova Hedwigia 14: 405. 1967[1968], nom. illeg. (macroconidia) Type. Shanoria 

indica Anahosur (= Strigula nitidula Mont.?). 

Specific limits in Strigula pose a number of problems. The corticolous and saxicolous taxa have the usual 

problem of overlapping ascospore and macroconidia sizes and possible significance of spore arrangement in 

the asci. The foliicolous taxa seem to me (a nonspecialist) based more on the aspect or pattern of growth of 

144

the species or strain of the photobiont and relatively little on "mycological" characters (e.g., Lücking, 1992). I 

feel that Santesson (1952) set the pattern for the recognition of what I would consider microspecies and this 

has been maintained or exaggerated since. It is thus no surprise that a seeming generic shift in photobiont has 

led to the recognition of Phylloporis. (The generic limits in the Trentepohliaceae are in flux and this difference 

may not exist.) Further the supra-/subcuticular "thallus" is a character of the photobiont as probably is the 

occurence of Raciborskiella on the underside of leaves. The asci, ascospores, microconidia and macroconidia 

are identical and thus I can see no shred of mycological support for the recognition of either genus. (The 

mycobiont characters of some of the species currently assigned to different genera seem so similar I think it 

would be worth using molecular techniques to verify whether they are or are not the same fungus associated 

with different algae.) Also the tide seems to be against granting even more striking photobiont differences 

generic status, e.g., Lobariaceae and most recently, Hymeneliaceae (Lutzoni & Brodo, 1995). 

Several species which do not occur in Florida or even in North America are described below as they 

provide new and interesting information for our understanding of the genus, corticolous species with black 

hypothallus, compound ascomata, lateral ostiole, muriform-spored taxa, beaked macropycnidia, fragmenting 

macroconidia and macroconida basally attached with apical or basal gealtinous caps. 

KEY TO NORTH AMERICAN SPECIES 

1. Growing on bark, wood or rock ................................................................................................................. 2 

2. Ascospores transversely 1-7-septate.................................................................................................... 3 

3. Ascospores 1-septate ....................................................................................................................... 4 

4. Growing on bark or wood .............................................................................................................. 5 

5. Black hypothallus absent........................................................................................................... 6 

6. Ascomata unilocular.............................................................................................................. 7 

7. Ascospores 12-25 × 4-5.5 µm ........................................................................................... 8 

8. Ascospores 12-17 × 4.5-5 µm; macroconidia 7-9 × 2-2.5 µm; 

throughout Florida ....................................................... S. viridiseda (Nyl.) R. C. Harris 

8. Ascospores 15-25 × 4-5.5 µm; macroconidia 14-19 × 3-4.5 µm; 

throughout Florida ................................................................ S. americana R. C. Harris 

7. Ascospores smaller, 8-12 × 2.5-3 µm; peninsular 

Florida ...................................................................................S. phaea (Ach.) R. C. Harris 

6. Ascomata 2-3-locular; ascospores 7-10(-12) × 3-4.5 µm; 

Dade County................................................................................. S. connivens R. C. Harris 

5. Black hypothallus present; thallus white punctate; ascospores 

17-22 × 5-6 µm; South Carolina .....................................................[S. hypothallina R. C. Harris] 

4. Growing on limestone.................................................................................................................... 9 

9. Ascospores 11-15(-18) × 3.5-5 µm; Dade and Monroe 

counties .................................................................................S. bermudana (Nyl.) R. C. Harris 

9. Ascospores 18-22 × 5-7.5 µm; Monroe County 

(Key West) ............................................................................. S. wilsonii (Riddle) R. C. Harris 

145

3. Ascospores 3-7-septate .................................................................................................................. 10 

10. Ascospores 7-septate, 24-42 × 5-7.5 µm; macroconida 8-celled, 

23-30 × 3.5-5 µm, often over mosses; northeastern North America, 

Europe...................................................................................... [S. stigmatella (Ach.) R. C. Harris] 

10. Ascospores 3-septate, 14-17 × 3.5-5(-6) µm; macroconidia 

4-celled, 13-17 × 3-3.5 µm, Louisiana, Minnesota, Missouri, 

Ontario, Europe ........................................................................[S. jamesii (Swinsc.) R. C. Harris] 

2. Ascospores with at least 1 or 2 cells longitudinally sepate, submuriform to muriform ........................ 11 

11. Ascospores submuriform, 20-33 × 6-10 µm.................................................................................. 12 

12. Ascospores 26-33 × 8-10 µm, 7(-8)-septate with 2-6 cells 

1-2 longitudinally septate; macroconidia not found; Gulf 

Coast of Florida...............................................................................S. griseonitens R. C. Harris 

12. Ascospores 20-27 × 6-7.5 µm, 5-7-septate with only 1-few cells 

longitudinally septate; mostly midwestern but a single dubious 

record from Duval County ....................................S. submuriformis (R. C. Harris) R. C. Harris 

11. Ascospores densely muriform, 35-45 × 13-17 µm; macroconidia 

not found; ascomata immersed; wall colorless except around 

ostiole; Georgia and Louisiana............................................................. [S. laceribracae R. C. Harris] 

1. Growing on leaves (mostly modified from Santesson, 1952).................................................................. 13 

13. Ascospores 1-septate; macroconidia bacillar, 1-septate or simple................................................... 14 

14. Ascospores over 11 × 4 µm ......................................................................................................... 15 

15. Ascospores with cells relatively equal in size, 14-22(-24) × 4-5.5 µm; 

common, throughout Florida ..........................................................................S. smaragdula Fr. 

15. Ascospores with upper cell enlarged, 11-15 × 4-4.5 µm; 

Dade County...................................................................................S. subelegans Vainio s. lat. 

14. Ascospores smaller, 8-10(-12) × (2-)2.5-3 µm; macroconidia simple; 

Seminole and Washington counties...................................................................... S. nitidula Mont. 

13. Ascospores 3-septate, (12-)15-24 × 3.5-6 µm; macroconidia filiform, 

3-7(-9)-septate;not common, throughout Florida ...................................... S. complanata (Fée) Mont. 

NOTES 

Strigula albicascens (Nyl.) R. C. Harris, comb. nov. 

Verrucaria albicascens Nyl., Lich. Nov. Zel. 129. 1888. Porina albicascens (Nyl.) Müll. Arg., Bull. 

Soc. Bot. Belgique 31: 38. 1892. Type. NEW ZEALAND. 1867, Knight 318 (H-Nyl 1534[A], 

lectotype selected here), 16 Jan 1882, Knight (H-Nyl 1983, syntype). 

This species known to me only from the type collections has whitish or grayish, endophloeodal thallus, 

unilocular ascomata with pale brown walls and 3-7-septate ascospores, 17-30 × 5-7 µm. 

Strigula albolinitum (Nyl.) R. C. Harris, comb. nov. 

Verrucaria albolinita Nyl., Lich. ins. guin. 38. 1889. Microglaena albolinita (Nyl.) Zahlbr., Cat. lich. 

146

univ. 1: 188. 1921. Phyllobathelium albolinitum (Nyl.) H. Mayrh., Biblioth. Lichenol. 26: 79. 

1987. Type. INSULA PRINCIPE. 1887, Fr. Quintas (H-Nyl 1588, holotype). 

See Mayrhofer (1987) for description. It is a little anomalous in either Phyllobathelium or Strigula. The 

immersed, pale walled ascomata fit best in Strigula but the large (ca. 40 × 16 µm) muriform macroconidia are 

apparently basally attached and have gelatinous caps at both ends more like Phyllobathelium. 

Strigula bahamensis (Riddle) R. C. Harris, comb. nov. 

Lithothelium bahamense Riddle in Britton & Millspaugh, Bahama Fl. 532. 1920. Type. BAHAMAS. 

Watling's Island: vicinity of Cockburn Town, on limestone, 13 Mar 1907, Britton & Millspaugh 

6127 (FH, holotype). 

Aptroot accepted this species in Lithothelium. He misread the the original description and cited Britton & 

Millspaugh 6129 as the type. This is of no importance since both 6127 and 6129 represent the same species 

of Strigula. The FH holotype has asci and ascospores of the Strigula-type and Britton & Millspaugh 6129 (NY) 

has two types of pycnidia, one with elliptical-oblong microconidia, the other with 5-7-septate macroconidia, 

both typical of Strigula, not Lithothelium, which has filiform microconidia and no known macroconidia. The 

reason that Aptroot did not see NY material is that it had already been filed in Strigula before he undertook his 

study and was not included in the loan for Lithothelium. 

Strigula bermudana (Tuck. ex Nyl.) R. C. Harris, comb. nov. 

Verrucaria bermudana Tuck. ex Nyl., Sert. lich. trop. 43. 1891. Porina bermudana (Nyl.) Zahlbr., 

Cat. lich. univ. 1: 367. 1922. Type. BERMUDA. 1881, Farlow 31 (H-Nyl 985, holotype). 

I had finished the notes for Strigula and quite contentedly synonymized Verrucaria bermudana with 

Strigula viridiseda as I had done in my thesis when I had a sudden attack of "What was I thinking?". Not only 

is the material on limestone (not bark) but the ascomata are considerably larger (0.5-1.0 mm vs. 0.3-0.4 mm). 

Therefore the saxicolous material is here recognized as a distinct species. All the problems are not solved, 

however, as the ascospore size intergrades with that of S. wilsonii. This problem requires an extensive, more 

sophisticated re-analysis of the data. The type and other tropical material have the ascospores uniseriately 

arranged. In Buck 27425 (NY) from Suwanee County the ascospores are biseriate and the paraphyses are 

more branched and may represent an undescribed species. 

Strigula cinefaciens (Nyl.) R. C. Harris, comb. nov. 

Verrucaria cinefaciens Nyl., Bull. Soc. Linn. Normandie sér. 2, 7: 180. 1873. Arthopyrenia 

cinefaciens (Nyl.) Zahlbr., Cat. lich. univ. 1: 320. 1921. Type. ANDAMAN ISLANDS. 1867, Kurz 40 

(H-Nyl 728, holotype; M, isotype). 

This corticolous species has a shiny gray thallus with black dots formed from a black hypothallus which 

here and there penetrates the thallus. This type of thallus is well known for the foliicolous taxa (previously 

placed in both Strigula and Phylloporis) but I believe this is the first corticolous representative. To my mind it is 

one more link binding together a broadly defined genus Strigula. The ascospores are 2-celled, ca. 14-15 × 4-5 

µm, tending to break into part spores. 

Strigula connivens R. C. Harris, sp. nov. 

Species unica ascocarpis compositis, 2-3 loculis ostiolo unico et ascosporis 1-septatis, 

7-10(-12) × 3-4.5 µm. 

Type. FLORIDA. Dade County: Cutler, 1898, Thaxter 19 (MSC, holotype). 

Thallus corticolous, greenish gray, well developed, endophloeodal. Ascomata compound, usually with 

two chambers, occasionally three, rarely uniloculate, complanate, 0.3-0.4 mm long, slightly less in width; 

chambers connected above, sharing a single ostiole; wall thinner or lacking below. Asci narrowly elliptical, 45- 

50 × 7-8 µm, with eight subbiseriate to biseriate spores. Ascospores, narrowly ovate, 2-celled, slightly 

constricted at septum, 7-10(-12) × 3-4.5 µm. 

147

If I had seen only a single collection of this I probably would have ignored it as an ascomatal aberration 

with sterile tissue or bark invading the centrum as is occasionally seen in Pyrenulaceae or Trypetheliaceae. 

However, it does seem to be a genuinely compound ascomatal type which expands the limits of variation in 

Strigula. 

Additional specimens. FLORIDA. Dade County: Cutler, Thaxter 17a (MICH), Snapper Hammock, 11 Dec 

1919, Britton & Britton 806 (FH, NY). 

Strigula fracticonidia R. C. Harris, sp. nov. 

Thallo corticola albo, ascosporis submuriformibus, 7 × 1-septatis, 25-27 × 8-9 µm et macroconidiis 3septatis, 

ca. 19 × 3 µm, in partibus rupentibus. 

Type. NEW ZEALAND. South Island: Marlborough, 9.5 km NW of Kaikoura, Mt. Fyffe S. F., 4 km SW of summit 

of Mt. Fyffe, along creek, alt. 180-330 m, on trunk of Melicytus ramiflora, 6 Feb 1981, Tibell 10790 (UPS, 

holotype). 

Thallus corticolous, white, not corticate, endophloeodal. Ascomata half immersed, subglobose, 0.25-0.4 

mm diam.; wall pale below with dark, poorly developed clypeus. Asci cylindrical, ca. 90 × 20 µm, with eight 

biseriate spores. Ascospores fusiform, submuriform, 7 × 1-septate, 25-27 × 8-9 µm. Macroconidia 3-septate, 

ca. 19 × 3 µm, fragmenting into four part spores. Microconidia oblong to ovate, 3-4.5 × 1.5-2 µm. 

Taxa with the ascospores fragmenting into part spores are well known in both foliicolous and corticolous 

species of Strigula but I do not know of any other species where the macroconidia break apart. The 

ascospores of S. fracticonidia are close to those of S. griseonitens which differs in having a corticate thallus 

and completely immersed ascomata. Strigula fracticonidia is known only from the type collection. 

Strigula griseonitens R. C. Harris, sp. nov. 

Thallo griseo corticato, ascomatis in cortice immersis parietibus hyalinis vel pallidis et ascosporis 

submuriformibus, 7(-8)-septatis, 3-6 cellulis longitudinaliter 1-2-septatis, 26-33 × 8-10 µm. 

Type. FLORIDA. Citrus County: Withlacoochee State Forest, along Florida Trail N to Brush Sink from Co. Rd. 

480, 1 mi E of Co. Rd. 491, ca. 11 mi SW of Inverness, 28°42'N, 82°26'W, oak-pine woods, 6 Dec 1993, 

Harris 31825 (NY, holotype). 

Thallus corticolous, gray, ± shiny, with prosoplectenchymatous cortex, 10-30 µm thick. Ascomata 

immersed, subglobose, 0.3-0.4 mm diam; wall colorless below, expanded and streaked with pale brown 

around ostiole. Asci cylindrical, 100-130 × 15-20 µm, with eight biseriate spores. Ascospores submuriform, 

7(-8)-septate with 3-6 cells longitudinally 1-2-septate, 26-33 × 8-10 µm. Microconidia oblong, 4-5 × 1.5-2 µm. 

Macroconidia not found. 

Strigula griseonitens, named for the conspicuous thallus, is separated from S. submuriformis by larger, 

more divided ascospores, immersed ascomata and the shiny, corticate thallus. Muriform-spored species of 

Strigula seem rather trendy just now. I suppose as a result of more natural families and genera being 

recognized since some of them have been transferred from other genera where they were misplaced due to 

an artificial system based almost entirely on ascospore septation and color. Ascospores in Strigula are now 

allowed to range from 1-septate through submuriform to very strongly muriform. In addition to S. 

submuriformis, known since 1967, three recently described species occur on rock, S. porinoides Canals et al. 

with submuriform ascospores from Spain and Sicily, S. australiensis McCarthy and S. johnsonii McCarthy with 

more strongly muriform ascospores from Australia and New Zealand, and one on bark, S. lateralis Aptroot & 

v.d. Boom from Europe. Also on bark are S. fracticonidia from New Zealand, S. griseonitens, S. laceribracae 

from North America, and S. rostrata from Madagascar, newly described here. Strigula albolinita, S. obtecta 

148

and S. tagananae are transferred from other genera. 

Additional specimens. FLORIDA. Citrus County: as type, Harris 31810; Hillsborough County: Hillsborough 

River State Park, Rapids Trail along river, 28°09'N, 82°14'W, mixed harwoods, 3 Dec 1992, Harris 29647; 

Liberty County: Apalachicola National Forest, mature Taxodium dome 2.7 mi NW of Sumatra on Fla. Hwy. 

379, 28 Dec 1990, Harris 26104; Sarasota County: Myakka River State Park, 7 Mar 1937, Romer s.n. 

MISSISSIPPI. Jasper County: Bienville National Forest, Forest Service Rd. 506-3 at E edge of forest by small 

cemetery, 32°12'N, 89°13'W, ca. 105 m, dry oak woods on sand, 30 Sep 1992, Harris 28864 (all NY). 

Strigula hypothallina R. C. Harris, sp. nov. 

Thallo corticola nitido albo punctato, hypothallo nigro et ascosporis 1-septatis, 17-22 × 5-6 µm. 

Type. SOUTH CAROLINA. Berkeley County: Francis Marion National Forest, Guilliard Lake Campground, on Ilex, 

24 May 1967, Harris 3201-A (MSC, holotype). 

Thallus light brown, shiny, with scattered white dots; hypothallus black, visible at thallus margin or in 

section. Ascomata complanate to hemispherical, immersed, 0.5-0.8 mm diam.; wall black, sometimes 

merging with hypthallus, lacking below. Asci cylindrical, 90-120 × 9-10 µm, with eight uniseriate spores. 

Ascospores narrowly ovate, with ± pointed ends, 2-celled, slightly constricted at septum, 17-22 × 5-6 µm. 

Microconidia and macroconidia not found. 

Strigula cinefaciens is the only other corticolous species with a black hypothallus (not uncommon for 

foliicolous taxa). It is distinguished by black not white dotted thallus. Although known only from the type, I 

describe it to validate a name from my thesis and to expand the parallel character set shared by foliicolous 

and corticolous taxa. 

Strigula indutula (Nyl.) R. C. Harris, comb. nov. 

Verrucaria indutula Nyl., Lich. Nov. Zel. 129. 1888. Porina indutula (Nyl.) Müll. Arg., Bull. Herb. 

Boissier 2, App. 1: 92. 1894. Type. NEW ZEALAND. Knight (H-Nyl 1981, holotype). 

As Nylander pointed out this species is close to S. albicascens differing in better developed thallus and 

smaller ascospores, 5-7-septate, 16-20 × 4-5 µm. 

Strigula jamesii (Swinsc.) R. C. Harris 

I originally treated this species and our American material as S. affinis (Massal.) R. C. Harris but it was 

pointed out to me by Brian Coppins, I believe, that the two are distinct. At the time I accepted this but did not 

re-analyze the situation. I have now done so as far as possible with limited material. The ascospores are not 

much different if one believes the published ranges, S. jamesii 12-20(-22) × 2-3.5 µm (Purvis et al., 1992), S. 

affinis 12-20 × 3-6 µm (Roux & Bricaud, 1993). The macroconidia are also close, 13-17 × 3-3.5 µm for S. 

jamesii, 13-18 × 3-5 µm for S. affinis. The distinctions seem to lie elsewhere, S. affinis has a well developed, 

often rimose thallus and larger, more complanate ascomata while S. jamesii has a thin, continuous thallus with 

small, more distinctly hemisphaerical ascomata. The ascospore width given by Purvis et al. (1992) for S. 

jamesii does not agree with my observations which include the type. I would suggest these ascospore ranges 

for the two species, S. jamesii 13-17 × 4-5 µm and S. affinis 15-22 × 4.5-6 µm. According to my analysis 

American material is correctly placed in S. jamesii. 

Strigula laceribracae R. C. Harris, sp. nov. 

Thallo corticola albo, ascomatis immersis pariete hyalino vel pallido et ascosporis dense muriformibus, 

11-15 × 1-5-septatis, 35-45 × 13-17 µm. 

Type. GEORGIA. Murray County: Chattahoochee National Forest, Cohutta Wilderness, Tearbritches Trail from 

Forest Service Rd. 68 to Bald Mountain, 34°52'N, 84°38'W, 1100-1200 m, mesic oak woods, 22 Sep 1992, 

Harris 28170 (NY, holotype). 

149

Thallus corticolous, white, thin, ecorticate, mostly endophloeodal. Ascomata immersed, subglobose, 0.3- 

0.5 mm diam.; wall colorless below, expanded and streaked with brown around ostiole. Asci broadly ovate to 

obpyriform, 110-130 × 40-60 µm, with eight subbiseriately to irregularly arranged spores. Ascospores broadly 

fusiform, narrower at one end, densely muriform, 10-15 × 1-5-septate, 35-45 × 8-10 µm. Microconidia and 

macroconidia not found. 

Strigula laceribracae, named in "dog Latin" for the place of collection, is a very inconspicuous species as 

the white thallus is close to the color of the oak bark and the immersed ascomata are visible only as minute 

dark dots. The densely muriform ascospores are diagnostic among North American Strigulae. 

Additional specimen. LOUISIANA. East Baton Rouge Parish: Burden Research Plantation, Essen Lane, 

Baton Rouge, bottomland hardwood forest, 27 Mar 1980, Tucker 20899C (LSU). 

Strigula multipunctata (R. Sant.) R. C. Harris, comb. nov. 

Porina multipunctata R. Sant., Symb. Bot. Upsal. 12(1): 216. 1952. Phylloporis multipunctata (R. 

Sant.) V_zda, Folia Geobot. Phytotax. 19: 183. 1984. Type not seen. 

Strigula obducta (Müll. Arg.) R. C. Harris, comb. nov. 

Phylloporina obducta Müll. Arg., Flora 73: 198. 1890. Phylloporis obducta (Müll. Arg.) R. Sant. 

& Tibell, Austrobaileya 2: 539. 1988. Type not seen. 

Strigula obtecta (Vainio) R. C. Harris, comb. nov. 

Thelenella obtecta Vainio, Étud. class. lich. Brésil 2: 218. 1890. Polyblastiopsis obtecta (Vainio) 

Zahlbr., Cat. lich. univ. 1: 351. 1922. Phyllobathelium obtectum (Vainio) H. Mayrh., Biblioth. 

Lichenol. 26: 82. 1987. Type. BRAZIL. Minas Gerais: Sitio, 1885, Vainio, Lich. Brasil. exs. 718 

(BM, isotype). 

This species has strongly muriform ascospores. See Mayrhofer (1987) for description and illustration. 

Strigula phaea (Ach.) R. C. Harris 

Porina mundula Müll. Arg., Bot. Jahrb. Syst. 6: 402. 1885. Type. CUBA. Wright, Verr. Cub. 212 

(NY, isotype). 

Some forms of this corticolous species (e.g., the type of Porina mundula) have a very similar aspect to 

taxa assigned to Phylloporis with conical ascomata and weak basal fringe. I suppose some would be happy to 

see all corticolous and saxicolous Strigula placed in Phylloporis. 

Strigula phyllogena (Müll. Arg.) R. C. Harris, comb. nov. 

Porina phyllogena Müll. Arg., Flora 66: 335. 1883. Phylloporis phyllogena (Müll. Arg.) Clements, 

Genera of Fungi 173. 1909. Type not seen. 

Strigula platypoda (Müll. Arg.) R. C. Harris, comb. nov. 

Porina platypoda Müll. Arg., Flora 66: 335. 1883. Phylloporis platypoda (Müll. Arg.) V_zda, Folia 

Geobot. Phytotax. 19: 185. 1984. Type not seen. 

Strigula rostrata R. C. Harris & Aptroot, sp. nov. 

Species propria corticola macropycnidiis rostratis, macroconidiis submuriformibus 15-18 × 7-8 µm et 

ascosporis submuriformibus 27-30 × 7-8 µm. 

Type. MADAGASCAR. Tamatave, Perinet (=Andasibe), primary montane forest, 950 m, 48°16'E, 18°56'S, 10 

May 1984, Aptroot 13397 & Hensen (hb. Aptroot, holotype). 

Thallus corticolous, white, not corticate, mostly endophloeodal. Ascomata flattened conical, sessile, ca. 

150

1.0 mm diam.; wall black, thinner below. Asci cylindrical ca. 90-105 × 11-13 µm, initially with 8 uniseriate 

spores but several usually aborting (probably not typical?). Ascospores submuriform, 4-5 × 1-septate, 27-30 × 

7-8 µm. Macropycnidia with a short, thick beak, 0.3-0.4 mm diam. basally, 0.2-0.3 mm high. Macroconidia 

often extruded as a cirrus, submuriform, 15-18 × 7-8 µm, basally attached with a gelatinous basal cap. 

Microconida oblong to ovate, 3-4 × 1-2 µm. 

This is rather distant from the subject of Florida lichens but it is such an unusual species (Madagascar 

strikes again) that I could not resist. Long-beaked macropycnidia occur in Anisomeridium but this is the first 

known occurrence in Strigula. The basally attached, submuriform macroconidia are also unusual. It may turn 

out that only the transversely septate macroconidia are eccetrically attached. 

Additional specimen (macropycnidia only). MADAGASCAR. As type, Aptroot 13395 & Hensen (hb. Aptroot). 

Strigula smaragdula Fr.: Fr. 

Since it is a sanctioned name, S. smaragdula Fr. is conserved over the well known S. elegans (Fée) Müll. 

Arg. 

Strigula subelegans Vainio s. lat. 

The name is used in a broad sense since the only Florida collection has rather small ascospores, 11-15 × 

4-4.5 µm. Santesson (1952) gave 17-25 × 4-7 µm. The material otherwise seems a reasonable match for 

Santesson's illustration and has a bluish tinge as emphasized by Lücking (1992). In view of my opinions on 

species limits in foliicolous Strigula (see family discussion) I am in no position to suggest this is an 

undescribed species. 

New to North America. FLORIDA. Dade County: Brickell Hammock, on leaves of Gymnanthes lucida, 16 

Dec 1919, Britton & Britton 690 (NY). 

Strigula sychnogonioides (Nitschke) R. C. Harris 

This was described in the monotypic genus Geisleria Nitschke. Largely on the basis of apically free 

physes, lacking information on conidial types, I included it as a synonym of Strigula. On re-examination of old 

material I would exclude it from Strigula. Ernst (Herzogia 9: 321-337. 1993) provided a detailed discussion of 

its ecology but no help at all as to its taxonomic position even though she had abundant fresh material. My 

first guess having proved wrong I venture a second (probably no better) that Geisleria sychnogonioides 

Nitschke is an algal parasite belonging somewhere in or near the Clypeosphaeriaceae in the Xylariales s. lat. 

Strigula tagananae (Harmand) R. C. Harris, comb. nov. 

Verrucaria tagananae Harmand in Pitard & Harmand, Bull. Soc. Bot. Fr. 58(Mém. 22): 68. 1911. 

Polyblastiopsis tagananae (Harmand) Zahlbr., Cat. lich. univ. 1: 352. 1922. Type. CANARY 

ISLANDS. Tenerife: Vueltas de Taganana, 500 m, sur Ilex canariensis, Pitard 1691 (DUKE, 

holotype). 

Verrucaria tagananae is placed in Strigula with some hesitation. The photobiont, apically free physes, 

asci and microconidia are in agreement. However, the ostiole is lateral, a feature otherwise unknown in the 

family. It is known to me only from the type. Perhaps additional material will provide better evidence as to its 

placement. 

Strigula viridiseda (Nyl.) R. C. Harris 

This is at the center of a complex of poorly resolved taxa. In 1975 I included saxicolous material now 

removed to S. bermudana. The ascospore size range overlaps that of S. americana and determination is 

often difficult. The possibility exists that the recognition of a third species intermediate in ascospore and 

macroconidia size might resolve the problem. Also there is a collection from Ontario (Wong 3811, CANL) with 

ascospores, 12-16 × 4-5 µm, in the range of S. viridiseda but so far out of geographic range that I am 

undecided as to its disposition without further study. 

Strigula viridis (Lücking) R. C. Harris, comb. nov. 

151

Phylloporis viridis Lücking, Nova Hedwigia 52: 276. 1991. Type not seen. 

THELENELLACEAE O. Eriksson ex H. Mayrh. 

Biblioth. Lichenol. 26: 16. 1987. Type. Thelenella Nyl. 

Microglaenaceae Servít, Studia Bot. _ech. 7: 108. 1946 [illegitimate, based on illegitimate name]. Type. 

Microglaena Körber (1855) [non Microglena Ehrenberg, 1832] (= Thelenella Nyl.). 

Aspidotheliaceae Räsänen ex J. C. David & D. Hawksw., Systema Ascomycetum 10: 13. 1991. Type. 

Aspidothelium Vainio 

The features which unite the members of this family are fissitunicate? asci with an ocular chamber and a 

cone shaped plug or ring above it in the ascus tip staining with phloxine/KOH, quite irregular, short celled, 

branched and interconnected physes, periphysoids, ascomatal wall often not melanized and a 

chlorococcalean photobiont or non-lichenized. Conidia seem unusually hard to find but Thelenella and 

Celothelium have arcuate, ± filiform microconidia. I have not found microconidia for "Aspidothelium" but 

Santesson (1952) reported oblong, 3-4 × 1 µm conidia for A. cinerascens. I have found elliptic oblong 

macroconidia in one specimen of "Aspidothelium" and one of Chromatochlamys but more occurrences are 

needed to be convincing. The ascus type is similar to that of the Requienellaceae but I can see no other 

features in common and in fact see no close relationships to any lichen-forming loculoascomycete family. The 

unusual ascus and photobiont suggest the Thelenellaceae represent an independent (ancient?) adoption of 

the lichen lifestyle. 

Three genera were assigned to the Thelenellaceae by Mayrhofer. Due to lack of adequte material for 

study I am temporarily recognizing Chromatochlamys as distinct. Julella is not lichen-forming, has a different 

ascus type and is clearly a member of the Arthopyreniaceae. Aspidothelium is a synonym of Thelenella. 

Aptroot and Sipman (1993) assigned Musaespora Aptroot & Sipman to the Aspidotheliaceae (correctly 

realizing that the Aspidotheliaceae hardly differed from the Thelenellaceae). However, Musaespora has a 

trentepohlioid photobiont, a different ascus type and ellipsoid to globose microconidia. Ascospore type and 

microconidia suggest placement in the Monoblastiaceae and I tentatively assign it there pending further study 

as I have only limited material at present. Celothelium is placed here provisionally, perhaps more out of 

expedience than anything else. Although the ascomatal type is very different from Thelenella, the ascus has a 

conical ring staining with phloxine/KOH, arcuate microconidia (illustrated by Aguirre, 1991, for C. stenobelum) 

and thin, irregular, short-celled physes. The photobiont is also a problem for me but Celothelium may not be 

lichen-forming which eases my conscience somewhat. (The Dade County collection of Celothelium 

aciculiferum is on lignum and no algae are present.) The key to genera below is artificial as to separating 

Thelenella and Chromatochlamys since I cannot at this time come up with any natural characters. 

1. Growing on bark, rock or leaves ............................................................................................................... 2 

2. Ascospores elliptical to fusiform, 3-septate to muriform ........................................................Thelenella 

2. Ascospores filiform, transversely septate........................................................................... Celothelium 

1. Growing on mosses and plant remains........................................................................ [Chromatochlamys] 

CELOTHELIUM Massal. 

Type. Verrucaria socialis sensu Montagne. (Celothelium leprieurii Trevisan?) 

?Trichotrema Clements, Genera of Fungi 41. 1909. Type. Pleurothrema trichosporum Müll. Arg. 

Celothelium was resurrected by Aguirre (1991). Her treatment is not up to the level of that for 

Leptorhaphis. She proposed to typify Verrucaria socialis Zenker (1829) with a Leprieur collection. Leprieur 

collected in French Guiana from 1830-1869 so that this is not appropriate. However, Massalongo in the 

original description designated the type species as "Verrucaria socialis (Montagn.) Zenk. ?". (Montagne (1851) 

used the name Verrucaria socialis Zenker for some Leprieur collections but with no description.) I interpret 

152

this to mean Massalongo intended (in modern terms)Verrucaria socialis sensu Montagne. This is supported 

by Trevisan (Flora 44: 25. 1861) using the name (nomen novum?) Celothelium leprieurii in trying to sort out 

the confusion of names involved with Verrucaria socialis which he says can only be solved by examination of 

Zenker's original specimen. This view is also supported by the fact that the combination Celothelium socialis 

was not used by Massalongo. I suggest that Celothelium should be typified with Verrucaria socialis sensu 

Montagne (not Zenker) based on a specimen in Massalongo's herbarium. If such does not exist, then 

Aguirre's lectotype fortuitously turns out to be acceptable (not for Zenker's name as she proposed but for what 

probably should be called Celothelium leprieurii Trevisan). Because of inadequate material Aguirre declined 

treating C. socialis/leprieurii and I have seen no material. (The epithet is younger than aciculiferum but could 

replace cinchonarum.) 

Trichotrema is placed in synonymy with question since the holotype specimen of the generitype has been 

destroyed by Indian lichenologists and failing the discovery of an isotype the application of the name will have 

to fixed through neotypification. 

Aguirre recognized three species in the Neotropics. I would recognize only two. The ascospore 

character proposed to separate C. dominicanum (Vainio) Aguirre in the key is not supported by the data in her 

descriptions. I consider it a synonym of C. cinchonarum. Integration of the species of Trichotrema into 

Celothelium will probably add to the number of species recognized. I have specimens at NY of C. aciculiferum 

from Florida, Cuba, Dominican Republic, Jamaica, Puerto Rico, and Guyana and C. cinchonarum from Cuba, 

Jamaica, Puerto Rico, Trinidad and Ecuador. 

1. Ascomata solitary or several covered by a common clypeus; ostioles 

eccentric; asci strongly slanted to almost horizontally arranged; 

ascospores ca. 65-100 × 2 µm; microconidia filiform, often arcuate, 

10-17 × 0.5 µm; Collier and Dade counties ................................................. C. aciculiferum (Nyl.) Vainio 

1. Ascomata solitary or fused laterally (± melanothecoid); ostioles apical; 

asci arranged ± vertically; ascospores ca. 80-110 × 2 µm; microconidia 

filiform, often arcuate, 10-13 × 0.5 µm ............................................... [C. cinchonarum (Müll. Arg.) Vainio] 

NOTES 

Celothelium aciculiferum (Nyl.) Vainio 

New to North America. FLORIDA. Collier County: Collier-Seminole State Park, just W of boat basin, 

29°59'N, 81°35'W, mangroves at edge of salt marsh, 7 Dec 1992, Harris 30060; Dade County: Everglades 

National Park, vicinity of Pinelands Trail, 2 Jan 1976, Buck B840 (both NY). 

CHROMATOCHLAMYS Trevisan 

I have not had sufficient material to make thorough studies to resolve the status of Chromatochlamys. 

Preliminarily the asci seem to show the same staining characteristics as Thelenella. Mayrhofer separated 

them on presence (Chromatochlamys) or absence (Thelenella) of an ocular chamber. However, fresh 

specimens of most Thelenella species show an obvious ocular chamber as do older specimens if stained and 

treated with KOH. Thelenella was originally said to lack perphysoids but this was later changed so that there 

is no difference on this score. The physes seem a little coarser and contain non-staining, refractive oil 

droplets. Is this an indication that generic status is warranted? Further studies are needed. 

There is only a single species in North America, C. muscorum (Fr.) H. Mayrh. & Poelt with two varieties, 

muscorum and octosporum (Nyl.) H. Mayrh. & Poelt. I have seen var. muscorum with 2-4 spores per ascus 

from scattered localities (Colorado, Minnesota, North Carolina). The var. octosporum is included in Egan 

(1987) based on a personal communication from Hawksworth that Pyrenidium octosporum Looman from 

Saskatchewan is this taxon. 

153

THELENELLA Nyl. 

Mém. Soc. Sci. Nat. Cherbourg 3: 193. 1855. Type. T. modesta (Nyl.) Nyl. 

Microglaena Körber, Syst. Lich. German. 388. 1855. Type. M. wallrothiana Körber (= Thelenella modesta 

(Nyl.) Trevisan). 

Luykenia Trevisan, Conspect. Verruc. 19. 1860. Type. L. modesta (Nyl.) Trevisan 

Aspidothelium Vainio, Étud. class. lich. Brésil 2: 189. 1890. Type. A. cinerascens Vainio. 

Aspidopyrenium Vainio, Étud. class. lich. Brésil 2: 190. 1890. Type. A. insigne Vainio (= Aspidothelium 

fugiens (Müll. Arg.) R. Sant. fide Santesson, 1952). 

Secolegiella (Müll. Arg.) Müll. Arg., Hedwigia 34: 205. 1896. Type. Lecania fugiens Müll. Arg. 

Patellonectria Spegazzini, Bol. Acad. Ci. Córdoba 23: 477. 1919. Type. P. puiggari Spegazzini (= 

Aspidothelium cinerascens Vainio fide Santesson, 1952). 

Porinopsis Malme, Ark. Bot. 22A(6): 3. 1928. Type. P. geminipara Malme 

Thelenella and Aspidothelium are identical in all important characters as noted above, oddly staining 

ascus, fleshy ascoma with periphysoids, chlorococcalean photobiont, etc. Aspidothelium may have ascomata 

ornamented with a apical disk, warts or setae but they may also be smooth as in Thelenella. Also, it would not 

seem to take much of an expansion to go from the thick apical tissue present in most Thelenella ascomata to 

achieve apical warts or an apical disk. Some taxa of Aspidothelium have mainly transversely septate spores 

but Santesson (1952) noted that they may have one-few longitudinal septa. Both commonly have the area 

around the ostiole tinted greenish. Both often have much of the centrum taken up with spent asci. Both can 

be foliicolous or corticolous although Aspidothelium is not known from rock. Overall, I feel the similarities are 

fundamental, the differences unimportant and therefore synonymize them. 

KEY TO NORTH AMERICAN TAXA 

(Some data taken from Mayrhofer, 1987) 

1. Ascospores colorless ................................................................................................................................ 2 

2. Growing on bark or leaves .................................................................................................................... 3 

3. Growing on bark ................................................................................................................................ 4 

4. Ascospores muriform .................................................................................................................... 5 

5. Ascospores basically 8/ascus ................................................................................................... 6 

6. Ascospores 11-17 µm wide................................................................................................... 7 

7. Asci obclavate to pyriform, broadest toward the base; 

ascospores 30-38 × 13-15 µm, 7-9 × 2-4-septate; 

Dade and Seminole counties ............................................................T. rappii R. C. Harris 

7. Asci cylindrical or broadest in middle; ascospores 25-42 × 11-17 µm, 

7-9 × 2-3-septate; Europe, Califormia, Minnesota, Australia .............................................. 

........................................................................................................[T. modesta (Nyl.) Nyl.] 

6. Ascospores 7.5-11 µm wide.................................................................................................. 8 

8. Ascospores 7-11 × 1(-2)-septate, 24-34 × 7.5-10 µm; 

Michigan, Minnesota ................................................... [T. pertusariella (Nyl.) Vainio s. lat.] 

8. Ascospores 6-7 × 1-2-septate, 20-30 × 7-11 µm; 

California, Australia ..........................................[T. sychnogonioides (Zahlbr.) R. C. Harris] 

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5. Ascospores basically 4/ascus, 11-13 × 1-2-septate, with large cells, 

53-63 × 16-18 µm; ascomata pallid to dark brown, ± hemispherical, 

smooth; Collier County.................................................... T. cinerascens (Vainio) R. C. Harris 

4. Ascospores transversely 11-17-septate, 2-4/ascus, 75-100 × 17-20 µm; 

ascomata pallid to brownish, smooth; on branches; locally common, 

Collier County....................................................................... T. geminipara (Malme) R. C. Harris 

3. Growing on leaves; ascospores 8/ascus, transversely septate, 50-90 × 12-21 µm 

(Santesson, 1952); in Egan (1987) as fide Tucker, not seen.............................................................. 

.........................................................................................................[A. fugiens (Müll. Arg.) R. Sant.] 

2. Growing on rock .................................................................................................................................... 9 

9. Ascospores muriform ...................................................................................................................... 10 

10. Ascomata without dark brown clypeus (involucrellum) ............................................................. 11 

11. Ascospores oblong-ellipsoidal, ± tapered at one end, 

over 20 µm long................................................................................................................... 12 

12. Ascospores medium sized, 20-36 × 8-13 µm................................................................ 13 

13. Thallus thin, continuous to ± rimose........................................................................ 14 

14. Ascospores 24-32 × 9-13 µm, 5-7 × 2-3-septate............................................... 15 

15. Thallus light brown, olive brown to olive green; 

ascospores (24-)27-32 × 9-13 µm, 6-7 × 2-septate; 

New Jersey, Oklahoma, Neotropics, Africa, 

China, Australia ..........................................[T. brasiliensis (Müll. Arg.) Vainio] 

15. Thallus pale ocher to yellow brown; 

ascospores 24-30 × 9-13 µm, 5-7 × 2-3-septate; 

Arctic ............................................................[T. sordidula (Th. Fr.) H. Mayrh.] 

14. Ascospores 20-27 × 8-11 µm, 6-9 × 1-2-septate; 

Louisiana, Puerto Rico ................................................ [T. sastreana R. C. Harris] 

13. Thallus thick, rimose-areolate; ascospores 24-36 × 9-13 µm, 

6-8 × (1-)2-3-septate; California, Macaronesia, SW Europe....................................... 

......................................................................................[T. inductula (Nyl.) H. Mayrh.] 

12. Ascospores larger, 30-45 × 12-19 µm, 7-9 × 3-septate; 

Georgia, Neotropics, South Africa, Australia, New Zealand.............................................. 

..............................................................................................[T. luridella (Nyl.) H. Mayrh.] 

11. Ascospores broadly ellipsoidal, small, to 20 × 8-11 µm, 

6-7 × 2-3-septate; New York ..............................................................[T. humilis R. C. Harris] 

10. Ascomata with an open dark brown clypeus (involucrellum); 

ascospores 35-37 × 14-19 µm, 6-8 × (1-)2-3-septate; 

California, Baja California.............................................................................[T. weberi H. Mayrh.] 

155

9. Ascospores 4-celled, 18-23 × 6.5-7.5 µm; thallus scanty; on pebble; 

Puerto Rico........................................................................................... [Thelenella sp. Buck 18291A] 

1. Ascospores soon brown, 25-40 × 10-17 µm, 6-8 × 2-3-septate; on bark; 

California ..................................................................................................... [T. hassei (Zahlbr.) H. Mayrh.] 

NOTES 

Thelenella brasiliensis (Müll. Arg.) Vainio 

I have placed some specimens with smaller ascospores in a new species, T. sastreana, although its 

range overlaps that given by Mayrhofer. In doing this revision I have gotten the impression that he must have 

measured immature ascospores and thus his ranges go too low. The Oklahoma collection seems typical but 

the New Jersey collection is ecologically anomalous, semiaquatic, growing with Ionaspis lacustris (With.) 

Lutzoni. Thelenella inductula "differs" in thicker thallus. Further study should be undertaken to determine if it 

is distinct from brasiliensis. 

New to North America. NEW JERSEY. Hunterdon County: [Mitchell property between Upper Creek and 

Pine Hill Roads,] on submerged shale, Nov 1987, Aptroot 214. OKALAHOMA. Cherokee County: along Terapin 

Creek ca. 6 mi N of St. Rd. 82 on St. Rd. 100, 23 Apr 1988, Harris 21384 (both NY). 

Thelenella cinerascens (Vainio) R. C. Harris, comb. nov. 

Aspidothelium cinerascens Vainio, Étud. class. lich. Brésil 2: 189. 1890. Type. BRAZIL. Rio de 

Janeiro, 1885, Vainio, Lich. Bras. 215 (not seen). 

The North American material is somewhat anomalous in having the cells of the ascospores rather large. 

Additional Florida specimens and more comparative specimens from the Neotropics may show that North 

American material should be given taxonomic recognition. 

New to North America. FLORIDA. Collier County: Fakahatchee Strand State Preserve, 1.2 mi from N 

entrance of W. J. Janes Memorial Scenic Drive, open Taxodium swamp, 6 Dec 1992, Harris 29959-A, Collier- 

Seminole State Park, just W of boat basin, mangroves at edge of salt marsh, 7 Dec 1992, Harris 30049-G, 

30067-A (all NY). 

Thelenella cuyabense Malme 

I have not seen material of this species from Brazil. McCarthy (Lichenologist 27: 345. 1995) has 

transferred it to Aspidothelium. It is possible that it is an older name for T. rappii but time has not allowed 

examination of the type of T. cuyabense. 

Thelenella fugiens (Müll. Arg.) R. C. Harris, comb. nov. 

Lecania fugiens Müll. Arg., Lich. epiphylli novi 3. 1890. Type. BRAZIL. Apiahy, 1881-82, Puiggari 

250 (not seen). 

Thelenella geminipara (Malme) R. C. Harris, comb. nov. 

Porinopsis geminipara Malme, Ark. Bot. 22A(6): 3. 1928. Lectotype (Santesson, 1952). 

PARAGUAY. Colonia Risso prope Río Apa, Malme, Regn. Lich. 1856 (not seen). 

This name is applied to Florida material in a broad sense. According to Santesson (1952) the ascomata 

may be expanded at the apex, warted or smooth. The Florida collections have smooth, rounded ascomata. I 

have seen almost no comparative material. A specimen from Ecuador has a broad disk-like expansion but the 

ascospores are quite different from Florida specimens. Once again there is the possibility that North American 

material may represent an undescribed species. It seems locally abundant in the western Everglades. 


Hammock, on Ilex, 6 Dec 1992, Harris 29929, Collier-Seminole State Park, just W of boat basin, at edge of 

salt marsh, 7 Dec 1992, Harris 30050, Big Cypress National Preserve, Oasis Ranger Station along U.S. Hwy. 

41, along Florida Trail N of hwy., Taxodium swamp, 9 Dec 1992, Harris 30175 (all NY). 

156

Thelenella humilis R. C. Harris, sp. nov. 

Thallo epilithico, disperso-areolato, glauco, ascomatis unicis in areolis immersis et ascosporis ellipsoideis 

vel late ellipsoideis, 18-20 × 8-11 µm. 

Type. NEW YORK. Warren County: Crane Mountain, 43°32'N, 73°57'45"W, 2100-3100 ft, on boulder in trail, 18 

Sep 1993, Harris 30577 (NY, holotype). 

Thallus epilithic, dispersed areolate with irregular to orbicular areoles, ca. 0.5-1.0 mm diam., gray green 

when wet, paler dry. Ascomata subglobose, immersed singly in areoles with only blackish tip visible, ca. 0.25 

mm diam. ( fertile areoles ca 1.0 mm diam.); wall colorless except greenish around ostiole. Asci cylindrical, ca. 

120 × 17 µm, with 8 uniseriate ascospores. Ascospores ellipsoidal to broadly ellipsoidal, 6-7 × 2-3-septate, 

18-20 × 8-11 µm; conidia not found. 

The epithet is given as it is an inconspicuous species with small areoles and small ascospores. 

Thelenella humilis is distinctive in the greenish thallus color (mostly shades of brown or whitish in other 

species) and in the small, smoothly rounded ellipsoidal ascospores. It is known only from the type collection. 

Thelenella luridella (Nyl.) H. Mayrh. 

This species may prove to be more widely distributed in acid rock glades in the Southeast. 

New to North America. GEORGIA. Coffee County: Broxton Rocks Ecological Preserve, 9 mi NE of Broxton, 

3 mi S of Ocmulgee River, 31°44'N, 82°45'W, on sandstone near falls, 16-17 Dec 1993, Harris 32595, 32596; 

Jeff Davis County: 0.4 mi E of Coffee County line on Ga. Hwy. 107, ca. 11 mi NE of Broxton, ca. 2 mi S of 

Ocmulgee River, 31°45'N, 82°43'W, dissected sandstone outcrop, 5 Feb 1995, Buck 27509 (all NY). 

Thelenella pertusariella (Nyl.) Vainio s. lat. 

The North American specimens have more septate ascospores than reported for European material. 

However, the ascospore size fits and as I have seen nothing from Europe for comparison the exact status of 

American specimens is left for future study. 

New to North America. MICHIGAN. Baraga County: moist woods along Sturgeon Rver ca. 0.25 mi N of 

campground, 5 mi NE of Sidnaw, on Acer negundo, 6 Sep 1972, Harris 8115 (NY). MINNESOTA. Cass County: 

4 mi S of Whipholt (11 mi SE of Walker), on quaking aspen, 14 Aug 1976, Wetmore 26513 (mixed with and 

distributed as Strangospora pinicola, NY). 

Thelenella rappii R. C. Harris, sp. nov. 

Similis T. modestae sed ascis obclavatis vel pyriformibus, 85-110 × 30-35 µm et ascosporis ovoideis, 7-9 

× 2-4-septatis, 30-38 × 13-15 µm. 

Type. FLORIDA. Seminole County: Sanford, on Hicoria, Apr 1916, Rapp (FLAS F29437, holotype; FH, isotype, 

both annotated by G. K. Merrill as Clathroporina obtecta sp. nov.). 

Thallus epiphloeodal, whitish or olivaceous, thin, continuous. Ascomata emergent, covered by thallus, 

subglobose, 0.25-0.4 mm diam.; wall pale or darkened above. Asci obclavate to pyriform, 85-110 × 30-35 µm, 

with eight irregularly arranged ascospores. Ascospores ovoid, 7-9 × 2-4-septate, 30-38 × 13-15 µm; 

microconidia not found. 

Thelenella rappi is close to T. modesta differing in ascus shape (obclavate vs. ± cylindrical) and 

distribution (tropical vs. temperate). It is possible a larger suite of specimens will lead to synonymization. 

Additional specimen. FLORIDA. Dade County: Everglades National Park, just N of Paurotis Pond, 11 May 

1967, Harris 2650 (NY). 

157

Thelenella sastreana R. C. Harris, sp. nov. 

Thelenellae brasiliensis similis ascosporis minoribus, 7-8 × 1-2-septatis, 24-27 × 9-11 µm. 

Type. PUERTO RICO. Distr. Mayagüez: Maricao State Forest, near Maricao Recreation Area, Camino del 

Guama and along Hwy. 120 between Kms 16 and 17, ca. 1700 m, relatively recent roadcut and roadside, 27 

May 1989, Buck 17213 (NY, holotype). 

Thallus thin, continuous, tan. Ascomatal warts ± complanate, orbicular, 0.3-0.5 mm diam., blackish 

around ostiole. Ascomata subglobose, 0.2-0.3 mm diam.; wall colorless except greenish around ostiole. Asci 

subcylindrical, slightly widened near middle, 85-130 × 15-20 µm, with 8 mostly biseriate ascospores. 

Ascospores colorless, with ± pointed ends, tapered to lower end, 6-9 × 1-2-septate, 20-27 × 8-11 µm; conidia 

not found. 

This species is named for Dra. Inés Sastre-De Jesús who has facilitated our work in Puerto Rico and is a 

most amiable companion in the field. Thelenella sastreana is close to T. brasiliensis. The Caribbean 

population seems to have consistently smaller ascospores. The collection sites in Puerto Rico may indicate 

this is something of a pioneer species able to take advantage of anthropogenic disturbance. The collection 

from Louisiana differs from the type in longer asci and in having little visible thallus. 

Additional specimens. LOUISIANA. Natchitoches Parish: Longleaf Trail Vista, ca. 35 mi WNW of 

Alexandria, sandstone cliffs, 28 May 1976, Harris 11445-A. PUERTO RICO. As type, Buck 17221; Distr. 

Ponce/Arecibo: SW slope of Cerro Morales, 1700-2100 ft., along road off Hwy. 531 at ca. Km 3.3, moist 

roadside, 2 Jun 1989, Buck 17384 (all NY). 

Thelenella sychnogonioides (Zahlbr.) R. C. Harris, comb. nov. 

Microglaena sychnogonioides Zahlbr., Beih. Bot. Centralbl. 13: 151. 1902. Type. CALIFORNIA. 

Santa Monica Mts., on Quercus agrifolia, Aug 1900, Hasse 776 (W-7057, lectotype, selected 

here). 

Thelenella harrisii H. Mayrh., Biblioth. Lichenol. 26: 36. 1987. Type. CALIFORNIA. Santa Monica 

Range, on Quercus agrifolia, 1907, Hasse 800 (W, holotype, not seen), syn nov. 

The lectotype above could be called a holotype as it was annotated by Zahlbruckner but since there is 

some confusion, I have chosen to designate a lectotype. Mayrhofer (1987) placed M. sychnogonioides in 

synonymy with Thelenella hassei choosing as "holotype" a specimen which does not fit Zahlbruckner's 

protologue, "sporis ... hyalinis ... loculis subcubicis, verticalibus 7-8, horizontalibus 2 ... 20-34µ longis et 9-11 µ 

latis.". (T. hassei has larger, brown ascospores.) The reasons for this seem to be twofold. The specimen 

designated as the lectotype is numbered "776" while that cited in the protologue is "778". However, 

Zahlbruckner's handwriting may have been bad, the printer had a dyslexic typesetter, or no one bothered to 

proofread the text, since miscitations of collection numbers are common in Zahlbruckner's work. (There are 

three or four in a single paper on Rapp's Florida lichens.) Since the specimen was annoted by Zahlbruckner 

and matches the protologue, I don't see the number confusion as a problem. (Nor did Mayrhofer who cited 

776 at FH as an "isotype" of 778 at W.) Material distributed by Hasse as 776 (not seen by Zahlbruckner) is T. 

hassei. It may be noted here that the numbers on Hasse collections are not collection numbers but species or 

herbarium numbers so that specimens collected at different places and times will carry the same number. The 

other reason for Mayrhofer's treatment is that the lectotype specimen probably was not available to him. It 

was included in a loan to Imshaug and his loans have only very recently been returned. It is something of a 

relief to be able to dispose of T. harrisii as it is rather an embarassment to have a Californian species named 

for one who professes to beleive that North America ends at the Mississippi River. 

Thelenella trichothelioides (Sérusiaux & V_zda) R. C. Harris, comb. nov. 

Aspidothelium trichotheliodes Sérusiaux & V_zda, Lejeunia n.s. 90: 4. 1978. Type. ZAIRE. Dist. 

158

Afro-montagnard Centre-oriental: env. du km 42 de la route Bukavu-Walikale, alt. env. 2300 m, 

épiphylle sur Connarus longestipitatus, Lambinon 71/1178 (LG, holotype, not seen). 

Thelenella sp. Buck 18291A 

Although known only from a small collection from Puerto Rico, this species is included as it extends the 

limits of Thelenella. The ascospores are transversely 3-septate and the thallus is scanty but in other respects 

it seems to be a typical Thelenella. The ascospores although small overlap the lower end of the ranges of 

several muriform-spored species. 

PUERTO RICO. Distr. Ponce: Caribbean National Forest, Toro Negro Division, along Hwy. 143 at Km 20.5, 

ca. 100 m, on roadside pebble, 4 Nov 1990, Buck 18291A (NY). 

TRICHOTHELIALES 

TRICHOTHELIACEAE (Müll. Arg.) Bitter & Schilling 

Since my brush with this family in my student days I have tended to avoid it being aware of the problems 

involved with defining species. Recently it has received considerable attention. A family treatment for the 

Guianas (Aptroot & Sipman, Flora of the Guianas, Series E, fasc. 2, 1993), saxicolous Southern Hemisphere 

taxa (McCarthy, Biblioth. Lichenol. 52, 1992), Clathroporina (McCarthy, Lichenologist 27: 321-350. 1995) and 

a revision of the genera (Hafellner & Kalb, Biblioth. Lichenol. 57: 161-186. 1995). 

The first, the treatment for the Guianas, is rather unfortunate. For historical reasons and differing 

approaches by the two authors, this is the kind of publication that gives a bad name to mycological αtaxonomists 

struggling to deal with biodiversity before it is gone. It leaves us open to criticism (and denial of 

funding) by those employing more sophisticated techniques. Three "species" concepts for ascospore variation 

are used in the work without adequate warning or explanation. Following Santesson (1952) Sipman divided 

foliicolous Porina into very narrowly defined species. In contrast Aptroot used a very broad concept for the 

corticolous species, with what I would consider an unacceptably large amount of variation. A third treatment of 

ascospore variation is found in Trichothelium where a distinct morphotype is assigned to the varietal level. I 

think that as α-taxonomists we have an obligation to be consistent in our species concepts within a group or, if 

not, then to provide some "hard" evidence as to why not. The treatment is also flawed in more obvious ways, 

types of taxa described from the Guianas have not been studied and names have been used with little logic 

(see criticisms by McCarthy, 1995). Such an interesting and complex family deserves better than this hasty 

attempt. 

McCarthy has provided two good, well illustrated α-taxonomic revisions. As will be seen below I do not 

agree with him in all details (mostly in regard to recognition of North American populations which consistently 

differ from his broader concepts). Thus I find Aptroot's comment in his review of McCarthy (1992) (Bryologist 

97: 467. 1994) most amusing: "However, the species concept applied is rather narrow and some specimens 

seem to represent transitional forms." I suspect that McCarthy's approach may well represent the mean 

between the lumper and the splitter and therefore be the most useful to the general lichenologist trying to 

name a specimen. 

For those who dislike change, the paper by Hafellner and Kalb is most unfortunate, not only in itself, but 

worse it has encouraged me to go even further in rearranging the Trichotheliaceae. They used the 

pigmentation of the ascomatal wall to divide Porina s. lat. into Porina and Pseudosagedia. These groups have 

been long recognized with varying degrees of clarity but mostly nomenclatural problems have led to the 

retention of a broad Porina. Hafellner and Kalb have cut the Gordian Knot. Below I have taken a different 

approach that essentially raises to the generic level the major groups recognized by Santesson (1952), uniting 

the P. nitidula group (Pseudosagedia) with Trichothelium, removing the P. phyllogena group to Strigula and 

adding a redefined Clathroporina. Amazing as it seems, this does not require the description of any new 

genera. 

Looking first at the taxa with blackish ascomata containing the pigment Pseudosagedia-violet, found in 

both Pseudosagedia and Trichothelium, the question is "Are setose ascomata a generic character?". I do not 

believe so. Hafellner and Kalb admitted there is an intermediate species with "bumps" on the ascomata (I 

commend their honesty, I wouldn't have known). The tendency to develop ascomatal appendages in 

159

foliicolous species may be a general phenomenon. It is seen in the P. leptalea group (Segestria/Polycornum) 

and in the unrelated Thelenellaceae. Further, both the setose and non-setose species of Trichothelium share 

the tendency towards long, multiseptate ascospores, only very rarely becoming somewhat muriform and 

growing if not on leaves, in wet, even semiaquatic habitats. The presence of setae has no predictive value 

and even if it were to be demonstrated that the setose taxa form a monophyletic clade, I would be reluctant to 

recognize more than a single genus, Trichothelium. The problem of how to deal with a single species or group 

with a single strikingly different character has always been a problem, e.g., placement of the compound-leaved 

Acer negundo. It is my understanding the theory (but perhaps not the practice) of modern phylogenetic 

approaches is that the preponderance of evidence should outweigh any single character, e.g., the 

abandonment of distinctive single character lichen genera by the merger of Coelocaulon into Cetraria or the 

realignment of Ionaspis/Hymenelia (abandoning the photobiont character). With the exception of the 

foliicolous species, the distribution of Trichothelium is amphitropical. I also include Zamenhofia in 

Trichothelium. It was tentatively maintained by Hafellner and Kalb on the basis of lacking a chitinoid ascus 

ring and lacking pigments. I am very hesitant to give possible "loss mutations" generic status. Also the assay 

used for the pigments was the crude application of KOH, etc., and also there seems to be some disagreement 

, "... some species with a layer reacting K+ red" (Purvis et al., 1992). There is general agreement among 

recent studies that the original characters for Zamenhofia, isidia and periphyses, are meaningless at the 

generic level. (I do part company with McCarthy (1993) who dismissed the use of isidia at any level. I agree in 

a couple of the cases he cited that the "isidia" are due to abnormal growth of the photobiont but isidiate 

species are not uncommon, especially in Clathroporina where sterile isidiate thalli are common and isidia may 

be the main means of reproduction in tropical wet forests.) 

When it comes to Hafellner and Kalb's residual Porina, I would move in the other direction, splitting rather 

than lumping. Part of the impetus for this approach came from McCarthy's (1995) discovery that if one 

merges Clathroporina with Porina (as he would and I whole-heartedly agree), the older name is Clathroporina. 

The groups involved are those centered around P. lectissima, P. nucula and Clathroporina eminentior. The 

first, for which the name Segestria Fr. is available, can be separated by lack of oxalate crystals in thallus and 

ascomata, small, ± naked ascomata, small ascospores and corticolous/saxicolous species basically 

amphitropical. Hafellner and Kalb pointed out two species intermediate between this group and the P. nucula 

group. I suggest that they are actually reduced members (in adaptation to living on leaves) of the P. nucula 

group. The P. nucula group is separated from Clathroporina not on ascospore sepatation (transverse vs. 

muriform) but on thallus characters which lead to very different aspects. Clathroporina has a smooth, often 

shiny, highly organized thallus with a cortex, photobiont ± organized in vertical rows, a layer of oxalate crystals 

and a dark hypothallus. The thallus is not intimately involved with the substrate but separated by the 

hypothallus and often peels away becoming almost foliose. The ascomata are ± immersed, sessile or 

protruding from the lower side of the thallus (well illustrated by McCarthy, 1995). To Clathroporina as defined 

by its type, C. eminentior, I would add the P. mastoidea group, differing in having transversely septate 

ascospores. Hafellner and Kalb recognized Clathroporina on ascus characters: lack of chitinoid ring and 

presence of a constriction near the tip in young asci. I consider usage of such characters as grasping at 

straws but they could be right, in which case a new home would have to be found for the P. mastoidea group. 

Porina nucula, the type of Porina, has a less organized, often verrucose thallus lacking a dark hypothallus with 

sessile ascomata, emergent and often constricted at the base. Again ascospore septation is not a character 

as several species with muriform ascospores are referable to Porina in this restricted sense. These genera 

are also supported by ecology and distribution. Clathroporina is mainly a genus of wet, dark, mostly tropical 

forests. It is one of the few groups, like Phyllopsora and Thelotremataceae, that are adapted to the lower 

levels of tropical rainforest and Caribbean wet broadleaf forest. (Is the development of a hypothallus an 

adaptation to this habitat?) Porina may be basically subtropical, adapted to more open forests with higher light 

and probably less moisture. 

The generic definitions suggested here are rather crude but by using them as a hypothesis, additional 

characters can be examined in detail to evaluate their utility. It is possible that conidial characters may support 

the new generic arrangement. At this time there is too little data about microconidia. Macroconidia are 

reported (Purvis et al. 1992) for two species referred here to Trichothelium. They may also occur in Segestria 

leptalea but I need several more occurrences to hypothesize an anamorphic connection. Ascospore 

tendencies also give some support, large with a strong tendency to become muriform in Clathroporina, 

medium to large with little tendency to become muriform in Porina, small, few septate in Segestria, long, 

160

multiseptate in Trichothelium. Ascomatal setae are confined to Segestria and Trichothelium. Clathroporina 

also differs in having more organized isidia, the ostiolar area is often black and there are no foliicolous 

species. 

The recognition of these four genera, Clathroporina, Porina, Segestria and Trichothelium, is somewhat 

analogous to generic segregation in Parmelia, distinctive in aspect (characters of the association) but ± 

uniform in characters of the asci and ascospores. I feel that the genera thus constituted have considerable 

predictive value regarding probable ascospore type, ecology and distribution. There seem to be four basic 

thallus/ascomata paradigms with rather continuous ascospore variation within each. This can be seen in the 

specific keys below where ascomata play only a small role. I simply do not understand the ascospore 

variation in the Trichotheliaceae. As pointed out by Aptroot and Sipman, it seems to encompass a different 

order of magnitude from that which mycologists are accustomed. In fact I consider this one of the distinctive 

features defining Trichotheliaceae. However, no one, myself included, has put the time and effort in to 

analyzing the situation. This leaves a situation where "species" are intuitively defined and one person's 

speculation is a good as another's. I consider my treatment below speculative at best and subject to 

considerable correction. 

Most of the species treated by McCarthy (1995) remain in Clathroporina under the new circumscription. 

Porina tahitensis (Räsänen) McCarthy and Clathroporina nuculastrum Müll. Arg. belong in Porina as defined 

here. 

Caribbean Porina firmula Müll. Arg. and P. subfirmula Riddle have very odd, placodioid thalli and may be 

considered for generic recognition. Two other species that may be referable to Segestria but are anomalous 

because of the pale pinkish ascomata and long, multiseptate ascospores and occur in eastern North America 

are an isidiate taxon reported from Long Island, New York reported by Brodo as Porina nucula (Porina sp. 4 in 

thesis), and a non-isidiate one from New Hampshire (Porina sp. 2 in thesis). I have not had time to reexamine 

the material, especially with regard to generic disposition, and as they are both very rare, I have 

decided to omit them here. 

KEY TO GENERA 

1. Ascomata usually small, essentially naked, with at most a thin covering of thallus 

(occasionally immersed in substrate); ascomata and thallus lacking oxalate 

crystals; ascomatal wall either some shade of red (Porina-yellow, Sagedia-red) or 

brown to blackish, often with blue or violet tones (Pseudosagedia-violet); some 

species with setose ascomata; muriform ascospores very rare; considerable 

diversity on leaves; distribution, except for foliicolous species, amphitropical .......................................... 2 

2. Ascomata blackish; wall often with blue or violet tints, without orange or 

red tints, KOH+ purplish or KOH-(Pseudosagedia-violet); with a very 

strong tendency for muitiseptate, elongate (even filiform) ascospores; 

setose species not uncommon........................................................................................ Trichothelium 

2. Ascomata reddish, sometimes darkening to black, lacking blue or violet 

tints, KOH+ more intensely orange or red (Porina-yellow); ascospores 

short, mostly 3-7-septate; setose species rare .......................................................................Segestria 

1. Ascomata usually rather large, surrounded by well developed thalline wart or 

immersed in thallus; wart and thallus containing oxalate crystals; ascomatal wall 

some shade of yellow or red (Porina-yellow); ascomata not setose; muriform 

ascospores not uncommon; few species on leaves; distribution essentially 

tropical and subtropical ............................................................................................................................. 3 

3. Black hypothallus lacking; thallus firmly attached to substrate, poorly 

organized, often verrucose; ascomata superficial, often somewhat constricted 

at base; few muriform-spored species; some follicolous species known; mainly 

species of open, drier? forest or scrub habitats, perhaps mainly subtropical ...............................Porina 

161

3. Black hypothallus usually well developed; thallus often separating from 

substrate, usually well organized into cortical, algal and oxalate crystal 

layers, smooth, continous, shiny; ascomata mostly immersed to weakly 

emergent, sometimes forming spherical warts projected below the almost 

foliose thallus; ascospores large, often muriform; no foliicolous species known; 

mostly species of shaded, closed, wet, tropical forests ................................................... Clathroporina 

CLATHROPORINA Müll. Arg. 

1. Thallus not isidiate..................................................................................................................................... 2 

2. Ascospores 8-celled, 40-55 × 7.5-9(-11) µm; Dade and Seminole 

counties ......................................................................................C. subpungens (Malme) R. C. Harris 

2. Ascospores 8-celled, 35-50 × 5-7 µm; Collier, Dade and Monroe 

counties ..............................................................................................C. tetracerae (Ach.) R. C. Harris 

1. Thallus isidiate; ascospores 8-celled, 32-48 × 5-6(-7.5) µm; Dade, Monroe 

and Polk counties ................................................................................................ C. isidiifera R. C. Harris 

NOTES 

Clathroporina confinis Müll. Arg. = Porina nuculastrum (Müll. Arg.) R. C. Harris 

Clathroporina isidiifera R. C. Harris, sp. nov. 

Similis C. tetracerae sed thallo isidiifero, ascosporis 8-loculis, 36-48 × 5-6 µm. 

Type. FLORIDA. Polk County: Nalcrest, scrub W of recreation area, 28 Jan 1989, Wheeler s.n. (NY, holotype). 

Thallus light green fresh, grayer in herbarium, shiny, continuous, not separating from substrate, with a 

dark brown to black hypothallus, isidiate; isidia cylindrical to slightly clavate, occasionally branched, with a ca. 

10-15 µm cortical/algal layer and a ca. 25 µm diam. core of oxalate crystals. Ascomata subglobose, 0.2-0.3 

mm diam., immersed or in somewhat emergent warts; wart 0.4-0.7 mm diam.; wall colorless below to 

yellowish or orange-brown around ostiole. Asci cylindrical, ca. 140 × 12 µm, with eight obliquely ± biseriate 

spores. Ascospores cylindrical fusiform, mostly ± equally tapered at either end, 8-celled, 32-48 × 5-6(-7.5) µm; 

microconidia not found. 

There seem to be rather few isidiate species described. Based on the literature I originally identified this 

material as Porina isidiophora Vainio but having seen specimens at Turku, I agree with McCarthy that the 

"isidia" in that taxon are abnormal outgrowths of Trentepohlia. The same is true of Porina granulifera Vainio. 

The isidia of C. isidiifera are true isidia, constricted at the base with a organization similar to that of the thallus. 

Porina conspersa Malme from Brazil may be close (I have not seen material) but the ascospores are 

described as 50-65 × 7-9 µm. 

Additional specimens. (With ascomata) FLORIDA. Dade County: Deering Hammock, Cutler, Dec 1921, 

Small et al.10279, Snapper Hammock, 11 Dec 1919, Britton & Britton 793; Monroe County: S part of Big Pine 

Key, 8 May 1919, Small 9125; Polk County: as type, 28 Jan, 20, 21 May 1989, Wheeler s.n. (Sterile) Dade 

County: Hammer Key, Everglades, 12 May 1918, Small 8609, Royal Palm Hammock, 27 Apr 1918, Small 

8523; Polk County: as type, 5 Feb, 11 Feb, 22 Apr, 6 May 1989, Wheeler s.n. (all NY). 

Clathroporina mastoidea (Ach.) R. C. Harris, comb. nov. 

162

Pyrenula mastoidea Ach., Ges. Naturf. Berlin Mag. Neusten Entdeck. Gesammten Naturk. 6:16. 

1814. Porina mastoidea (Ach.) Müll. Arg., Bot. Jahrb. Syst. 6: 399. 1885. Type. PHILIPPINES. 

Manila (UPS, lectotype, Malme, 1929, not seen; isolectotype, H-Ach 830-A). 

Although reported from North America, all the material which I have examined has ascospores smaller 

than those of the type of C. mastoidea. The epithet mastoidea has been widely misapplied in North America, 

usually to taxa in the Porina nucula group. 

Clathroporina subpungens (Malme) R. C. Harris, comb. nov. 

Porina subpungens Malme, Ark. Bot. 23A(1): 17. 1929. Type. PARAGUAY. Colonia Risso, Guarda 

Francia, 16 Oct 1893, Malme, Regn. Lich. 1927 (S, lectotype, Mccarthy, 1993). 

McCarthy (1993) placed this name in synonymy with Porina mastoidea (Ach.) Müll. Arg., that has larger 

ascospores (50-62 × 8-9(-10.5) µm in lectotype). On a global scale McCarthy may well be correct but since 

the lectotype of P. mastoidea is Asian and all the North American material has smaller ascospores, and 

because a name for a taxon so defined is available, I have chosen to take up Malme's name pending a careful 

statistical study of ascospore variation or perhaps molecular study. Although this species and the next were 

included in my thesis, this is the first published record of their occurrence in North America. 

New to North America. FLORIDA. Dade County: Royal Palm State Park, 6 Dec 1919, Britton & Britton 763 

(FH, NY), hammock, E end of Long [Pine] Key, Everglades, Small 8397 (NY); Seminole County: Sanford, on 

oak, Feb 1919, Rapp 746 (MICH). 

Clathroporina tetracerae (Ach.) R. C. Harris, comb. nov. 

Verrucaria tetracerae Ach., Methodus 121. 1803. Porina tetracerae (Ach.) Müll. Arg., Bot. Jahrb. 

Syst. 6: 401. 1885. Type. GUINEA. (H-Ach 838-C, lectotype, selected here). 

McCarthy (1993) cited H-Ach 838 as the holotype. It is mixed: 838-A is a Pyrenula. 838-B is internally 

identical to 838-C but has ascomata ca. 2× as large. This refinement of the lectotypification is purely 

precautionary and has no effect on the circumscription at this time. Material at S also contains a mixture 

pieces with large and small ascomata. 

Clathroporina tetracerae seems to have been more common in the past than it is now. It will be 

interesting to see if this is lack of collecting or loss of habitat as Clathroporina, with its preference for rain 

forests, is likely to be the most ecologically sensitive member of the Trichotheliaceae. 


Hammock, 6 Dec 1992, Harris 29932 (NY); Dade County: Cocoanut Grove, 1898, Thaxter 277 (FH), 287 

(MICH), Deering Hammock, Cutler, 29 Apr 1918, Small 8352 (NY), Hattie Bauer Hammock, Small & Mosier 

5954 (NY), Royal Palm Hammock State Park, 6 Dec 1919, Britton & Britton 757 (NY), Snapper Hammock, 11 

Dec 1919, Britton & Britton 800 (FH, NY); Monroe County: Key Largo, Apr 1921, Kelly (MICH). 

PORINA Müll. Arg., nom. cons. 

1. Thallus and ascomatal warts not isidiate .................................................................................................. 2 

2. Ascospores transversely septate .......................................................................................................... 3 

3. Growing on bark ................................................................................................................................ 4 

4. Ascospores fusiform to fusiform-cyllindrical, 7(-9)-septate ........................................................... 5 

5. Ascospores larger, over 40 × 8 µm ........................................................................................... 6 

6. Ascospores less than 60 µm long ......................................................................................... 7 

7. Ascospores broad, with rounded ends, 40-52 × 11-14 µm, 

8-celled; Dade and Lee? counties.............................................................. P. nucula Ach. 

163

7. Ascospores narrower, with pointed ends, 42-53(-61) 

× (7-)8-11(-12) µm, 8(-10)-celled; eastern Florida, 

Citrus to Dade County .................................................................................Porina sp. 674 

6. Ascospores 60-75 × 10-13(-15) µm, 8-celled, ends pointed, 

± evenly tapered at both ends; ascomatal warts rather large, 

ca. 0.6 mm diam.; Collier, Dade, Putnam and Seminole 

counties ................................................................................................................Porina sp. 6 

5. Ascospores small, 25-33 × 5-6 µm, 8-celled; ascomatal warts small, 

ca. 0.3 mm diam.; Collier County ....................................................................Porina sp. 30308 

4. Ascospores clavate with long tapering tail, (9-)11-13-septate, 

85-110 × 12-14 µm; common, Texas to North Carolina south to 

Seminole County in Florida .................................................... P. heterospora (Fink) R. C. Harris 

3. Growing on HCl- rock; thallus relatively smooth; asci ca. 130 × 20 µm; 

ascospores 8-celled, fusiform, 43-49 × 8-9 µm; Hillsborough County................................................ 

.....................................................................................................................Porina sp. Buck 22669 

2. Ascospores fusiform, muriform, 9-13-septate with few to most cells longitudinally 

or obliquely 1-2-septate, 55-80 × 13-18 µm, usually with well developed gel sheath; 

Dade and Lee? counties .......................................................P. nuculastrum (Müll. Arg.) R. C. Harris 

1. Thallus and/or ascomatal warts isidiate; ascospores 8-celled, 25-47(-57) × 6.5-8 µm; 

North Florida, Louisiana and South Carolina ...................................................... P. scabrida R. C. Harris 

NOTES 

Porina heterospora (Fink) R. C. Harris 

McCarthy (1993) placed this taxon as a synonym of Porina guaranitica Malme. Although I am generally 

rather broad minded at the species level, I cannot bring myself to accept this disposition. I have examined the 

lectotype of P. guaranitica. It is on rock with a very smooth pinkish thallus. Porina heterospora occurs on bark 

and has a verrucose, usually tan thallus. One might be able to dismiss the difference as a consequence of the 

semiaquatic habitat of P. guarantica and I would probably be willing to do so if the ascospore size ranges were 

more congruent. Based on ten ascospores from the lectotype of P. guaranitica the mean ascospore length is 

65.1 µm (range 57-71 µm) with a mean number of 9.5 septa. A small sample of P. heterospora gives 87.3 µm 

(range 60-112 µm) and 11.1 septa. McCarthy gave a mean ascospore size of 71.5 µm, presumably based on 

a mixture of P. guaranitica and P. heterospora but still well under that of North American collections. The form 

of the ascospores is different also. Both are clavate but P. heterospora has a longer tail, seemingly 

accounting for the greater ascospore length. Therefore, on the basis of larger, more sepate, longer tailed 

ascospores, thallus characters and corticolous substrate, I am maintaining P. heterospora as a distinct 

species. This would seem to be the most common species of Porina in the southern Coastal Plain. NY holds 

specimens of typical P. heterospora from Cuba and Venezuela but none seen as yet from Brazil. Since the 

epithet heterospora has a history of usage both in North America and Great Britain, even if it were a synonym, 

I suspect it would be conserved. 

Porina nucula Ach. 

McCarthy (1993) cited H-Ach 731 as the holotype of this name. However, this consists of two nonidentical 

pieces. H-Ach 731-A should be considered the lectotype. H-Ach 731-B is Porina mastoidea s. lat. A 

narrow species concept is used here, including only those collections with rather broad, rather short 

ascospores with rather blunt ends. Obviously the break between this circumscription and other putative taxa 

is not as tidy as one might like. However, I prefer to cut too finely rather than lose information in too broad a 

164

junk heap. These variants (sp. 6, sp. 674, sp. Buck 22669) have not been named as there are many 

uninvestigated names potentially available. They vary mainly in ascospore shape (more acutely pointed) and 

ascospore size. Buck 22669 grew on rock and may possibly be referable to Porina simulans Malme sensu 

McCarthy. 

Porina nuculastrum (Müll. Arg.) R. C. Harris, comb. nov. 

Clathroporina nuculastrum Müll. Arg., Flora 67: 618. 1884. Type. MADAGASCAR. Nossi-bé, 

Hildebrandt, comm. Rensch, 1884 (G, holotype). 

Since muriform ascospores have obviously evolved several times in the Trichotheliaceae, this character 

cannot be used to form a monophyletic genus. As Müller's name indicates, Clathroporina nuculastrum is most 

closely related to Porina nucula (they are externally indistinguishable) and the name is accordingly transferred 

to Porina. Two other species with submuriform/muriform ascospores belong in Porina as defined here, Porina 

ocellata (Malme) McCarthy (1993) and P. tahitensis (Räsänen) McCarthy (1995). See McCarthy (1995) for a 

list of synonyms of Porina nuculastrum, that was previously recorded from North America under one of these, 

Clathroporina confinis Müll. Arg. McCarthy (1995) incorrectly mapped the Florida collections in the central 

part of the state. I am unaware of any recent collections. 

FLORIDA. "East Florida", 1877, Smith & Austin (FH-Tuck 3929, MICH, NY), "Southeastern Florida", Smith 

(MICH, holotype of Polyblastiopsis floridana Fink in Hedrick); Dade County: Cocoanut Grove, 1898, Thaxter 

330 (FH), 296, 326 (MICH). 

Porina scabrida R. C. Harris, sp. nov. 

Subsimilis P. nuculae sed thallo isidiato et ascosporis minoribus, 37-42 × 5.5-7.5 µm, extremiis 

acutioribus. 

Type. FLORIDA. Columbia County: W side of Woodland Avenue, off Co. Rd. 138, 0.5 mi W of US 27, 29°51'N, 

82°39'W, hardwood forest, on Fraxinus, 5 Dec 1994, Buck 27418 (NY, holotype). 

Thallus pale olive tan, verrucose, dull to slightly shiny, densely to sparsely isidiate or occasionally nonisidiate; 

isidia cylindrical, occasionally branched, often rather irregular, consisting mostly of the photobiont with 

an external layer one cell thick. Ascomatal warts sessile, complanate to hemispherical, sometimes slightly 

constricted at base, 0.5-0.7 mm diam., often bearing cylindrical isidia. Asci cylindrical, ca. 120 × 17 µm, with 

eight obliquely subbiseriate spores. Ascospores fusiform-cylindrical, 8-celled, 35-47(-57) × 5.5-8 µm, both 

ends pointed, tapering ± evenly. Microconidia not found. 

Lacking isidia, the ascospore size would distinguish this taxon in North America. Even if P. scabrida 

were considered an aberrantly isidiate morph, there is no obvious non-isidiate taxon to refer it to. The thallus is 

sometimes lacking isidia, especially when ascomata are abundant, but one can then usually find them on the 

ascomatal warts. This is apparently a species of the southern Coastal Plain with its southern limit in Dixie and 

Putnam counties, Florida. 

Additional specimens. FLORIDA. Dixie County: Steinhatchee Wildlfie Management Area, Pumpkin Swamp, 

ca. 7 mi NW of Cross City, 29°41'N, 83°13'W, hardwood-Taxodium swamp, on Nyssa, 4 Dec 1993, Harris 

31539-A (NY); Lafayette County: along Co. Rd. 357, 6.4 mi SE of Fla. Hwy. 51, 29°51'N, 83°15'W, hardwood- 

Taxodium swamp, 2 Dec 1994, Buck 27291 (NY); Leon County: Apalachicola Natl. Forest, Rocky Bluff Scenic 

Area, ca. 3 mi S of Bloxham, mesic hardwoods, on Quercus, 4 Dec 1988, Harris 23411 (NY), swamp forest E 

of Natural Bridge Battlfield Monument, 6 mi E of Woodville, on Taxodium, 1 Jan 1991, Harris 26227 (NY); 

Putnam County: Ocala Natl. Forest, old Johnson Field Campground along Oklawaha River, just W of US Hwy. 

19, swamp forest, 7 Dec 1988, Buck 16807 (NY); Suwannee County: Ichetucknee Springs State Park, near N 

entrance, 29°59'N, 82°46'W, wet hardwoods, 2 Dec 1992, Buck 22582 (NY). LOUISIANA. East Baton Rouge 

Parish: Centurion Development, 4 mi E of Baton Rouge, 29 May 1976, Wetmore 24892 (MIN); Rapides 

Parish: Magnolia Recreation Area, Kisatchie Natl. Forest, 12 mi SW of Alexandria, ca. 70 m, mixed deciduous 

forest, 31 Dec 1969, Egan EL-7305, 7339, 7350 (MIN); St. Martin Parish: near St. Martinville, on Carya 

165

aquatica, 15 Jan 1894, Langlois (NY). SOUTH CAROLINA. Santee Canal, Ravenel, Rel. Tuck. 136 (NY). 

Porina sp. 30308 

Known from a single Collier County collection, this taxon is distinct in its small ascomata and ascospores. 

On this basis it is similar to P. curtula Malme but lacks the dark ostiolar area emphasized by McCarthy (1993). 

SEGESTRIA Fr. 

Type. S. lectissima Fr. 

Polycornum Malcolm & V_zda, Folia Geobot. Phytotax. 30: 91: 1995. Type. P. rubrofuscum 

Malcolm & V_zda. 

As with Trichothelium, I do not regard the setose ascomata of Polycornum sufficient to define a genus. 

KEY TO SEGESTRIA IN EASTERN NORTH AMERICA 

1. Growing on bark or rock............................................................................................................................ 2 

2. Ascospores 4-celled, fusiform ............................................................................................................... 3 

3. Ascospores 16-23 × 3-5 µm (Purvis et al., 1992); on bark and rock; 

Leon and Liberty counties ................................................S. leptalea (Durieu & Mont.) R. C. Harris 

3. Ascospores 20-30 × 5-8 µm (Purvis et al., 1992); on rock; 

New Hampshire, Mississippi and New York ............................................................[S. lectissima Fr.] 

2. Ascospores 8-celled, clavate, 37-43 × 7-8 µm; Tennessee 

(Porina sp. 5 in thesis)...................................................................................................... [Segestria sp.] 

1. Growing on leaves..................................................................................................................................... 3 

4. Ascospores 4-celled, 14-20 × 2.5-4 µm (Santesson, 1952); Escambia and 

Seminole counties ........................................................................... S. rubentior (Stirton) R. C. Harris 

4. Ascospores 8-celled, 24-31 × 3-4 µm (Santesson, 1952); Dade County 

................................................................................................... S. octomera (Müll. Arg.) R. C. Harris 

NOTES 

Segestria leptalea (Durieu & Mont.) R. C. Harris, comb. nov. 

Biatora leptalea Durieu & Mont., Expl. sci. Algérie 1: 268. 1847. Porina leptalea (Durieu & Mont.) 

A. L. Sm., Monogr. Brit. lich. 2: 333. 1911. Type not seen. 

?Porina olivacea var. microspora Fink in Hedrick, Mycologia 25: 308. 1933. Type. NEW 

HAMPSHIRE. Lonely Lake, on Ilex, Farlow (MICH, Fink hb. 11,559, holotype). 

Eastern North American specimens on both bark and rock have oblong or rod-shaped microconidia, 3-6 

× 0.5-1.0 µm, unlike European which are said (Purvis et al., 1992) to be 1.7-2.5 × 1 µm. This was noticed by 

Harris (1975) and Fink's name was informally raised to the species level for eastern American material. 

However, whether there are any correlated differences remains to be studied. It may be that it should be 

treated as only a minor population variant. In addition to Florida and New Hampshire collections on bark it is 

known from Illinois and Kentucky on rock. 

Segestria mammillosa Th. Fr. 

Although distinctive in occurring on bryophytes and detritus, the lack of oxalate crystals, pigmentation and 

small 4-celled ascospores suggest placement here. It is known from western North America. 

Segestria octomera (Müll. Arg.) R. C. Harris, comb. nov. 

Phylloporina octomera Müll. Arg., Flora 73: 198. 1890. Porina octomera (Müll. Arg.) Schilling, 

166

Hedwigia 67: 274. 1927. Type not seen. 

This was known to me in 1975 from a single Thaxter collection from Dade County. I have lost track of 

the exact data. 

Segestria rubentior (Stirton) R. C. Harris, comb. nov. 

Verrucaria rubentior Striton, Proc. Philos. Soc. Glasgow 11: 107. 1878. Porina rubentior (Stirton) 

Müll. Arg., Flora 66: 334. 1883. Type not seen. 

New to North America. FLORIDA. Escambia County: along Pineville Road at Brushy Creek, ca. 2 mi W of 

Co. Rd. 97A, 30°48'N, 87°33'W, on bambusoid grass culm, 8 Dec 1993, Buck 24587 (NY); Seminole County: 

Sanford, on Sabal leaf, May 1931, Rapp (NY). 

TRICHOTHELIUM Müll. Arg. 

Type. T. epiphyllum Müll. Arg. 

Pseudosagedia (Müll. Arg.) Choisy, Bull. Mens. Soc. Linn. Soc. Bot. Lyon 18: 107. 1949. 

Arthopyrenia sect. Pseudosagedia Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 16: 428. 

1862. Lectotype (Hafellner & Kalb, 1995): Arthopyrenia carpinea (Pers. ex Ach.) Müll. Arg. (= 

Trichothelium aeneum (Wallr.) R. C. Harris). 

Zamenhofia Clauzade & Roux, Bull. Soc. Bot. Centre-Ouest, n.s., Num. Spéc. 7: 824. 1985. Type. 

Z. coralloidea (P. James) Clauzade & Roux 

It should be noted that if the typification of Arthopyrenia proposed elsewhere in this work is not accepted 

Trichothelium/Pseudosagedia may require conservation over Leiophloea. The rationale for uniting 

Pseudosagedia and Trichothelium is included in the discussion of the family above. I have no opinions about 

subgenus Limosagedia Hafellner & Kalb and therefore do not transfer it to Trichothelium. If I were to erect 

subgenera, I probably would create one for the distinctive T. crocynioides. It could well be that Trichothelium 

can be divided into smaller homogeneous genera but I feel that this has to await a critical re-evaluation of this 

whole group. 

KEY TO TRICHOTHELIUM IN EASTERN NORTH AMERICA 

1. Growing on bark or rock............................................................................................................................ 2 

2. Ascomata not setose............................................................................................................................. 3 

3. Thallus not isidiate............................................................................................................................. 4 

4. Thallus without obvious hypothallus .............................................................................................. 5 

5. Ascospores not filiform, 4-13-celled.......................................................................................... 6 

6. Ascospores 4-celled.............................................................................................................. 7 

7. Growing on bark; ascospores 20-24 × 4-5 µm; Polk and 

Seminole counties........................................................... T. aeneum (Wallr.) R. C. Harris 

7. Growing on rock; ascospores 18-25 × 4-5 µm; 

Massachusetts, New York............................................ [T. chloroticum (Ach.) R. C. Harris] 

6. Ascospores 8-13-celled......................................................................................................... 8 

8. Growing on bark ................................................................................................................ 9 

9. Ascospores clavate-fusiform or cylindrical; ascomatal wall 

brown or blue or violet in section................................................................................ 10 

167

10. Ascospores clavate-fusiform, 8-13-celled ............................................................. 11 

11. Ascospores 8-celled........................................................................................ 12 

12. Ascospores (32-)38-50 × 5.5-7.5 µm; common, 

throughout Florida ........................... T. cestrense (Michener) R. C. Harris 

12. Ascospores 28-35 × 4-6.5 µm; rare, Dade, 

Lee and Seminole counties ...................................... T. cestrense complex 

11. Ascospores 8-11(-13)-celled, 47-65 × 5-7.5 µm; 

rare, throughout Florida................................................. T. cestrense complex 

10. Ascospores narrowly cylindrical, 8-13-celled, 

(34-)50-67 × 3.5-6 µm; central Florida ................................ T. cestrense complex 

9. Ascospores fusiform, 8(-9)-celled, 30-38 × 6-9 µm; ascomatal 

wall blue-green above in section, pale below; ascomata 

0.3-0.4 mm diam., Palm Beach County ........................Trichothelium sp. Thaxter 92 

8. Growing on rock, often seasonally inundated; ascospores 

fusiform-clavate to fusiform, 8(-12)-celled, 32-45(-48) × 5-8(-10) µm; 

Kentucky, Massachusetts, Missouri, North Carolina and Virginia....................................... 

..................................................................................... [T. guentheri (Flotow) R. C. Harris] 

5. Ascospores filiform, 14-21-celled, 100-140 × 3-5 µm; 

common, south to Polk and Sarasota counties........................................................................... 

........................................................................T. rhaphidospermum (Müll. Arg.) R. C. Harris 

4. Thallus soft, dull, on a black byssoid hypothallus; ascospores 

narrowly fusiform to cylindrical, 4(-7)-celled, 20-35 × 2.5-3.5 µm; 

on Taxodium; Lee and Polk counties .............................................. T. crocynioides R. C. Harris 

3. Thallus isidiate; isidia crowded, branched and irregular; ascomata 

± immersed and hidden by isidia; ascospores clavate-fusiform, 

8(-9)-celled, 43-54 × 6-7.5 µm; on bark; Illinois...........................................[T. isidiatum R. C. Harris] 

2. Ascomata setose; on bark; ascospores narrowly fusiform, 15-25-septate, 

with some cells longitudinally septate, 100-124 × 8-14 µm (Santesson, 1952); 

Marion County ................................................................................T. horridulum (Müll. Arg.) R. Sant. 

1. Growing on leaves or culms of bambusoid grasses; thallus a thin olive-tan film .................................... 13 

13. Ascomata not setose......................................................................................................................... 14 

14. Ascospores fusiform, 4-6-celled, less than 32 µm long .............................................................. 15 

15. Ascospores 4-celled, 18-25 × 4-5 µm (Santesson, 1952); 

Dade County....................................................................... T. thaxteri (R. Sant.) R. C. Harris 

15. Ascospores 6-celled, 20-28(-32) × 3.5-5.5 µm (Santesson, 1952); 

Alachua (on bambusoid), Collier and Dade counties ................................................................ 

........................................................................................ T. nitidulum (Müll. Arg.) R. C. Harris 

14. Ascospores clavate, 8-celled, 38-42 × 7.5-8 µm; ascomatal wall 

168

dark green-blue in section; on bambusoid grass culm.................................................................... 

.................................................................................T. cestrense (Michener) R. C. Harris s. lat. 

13. Ascomata setose; ascospores fusiform, 8-celled, (25-)36-50 × 4-6 µm 

(Santesson, 1952); Louisiana..........................................................................[T. epiphyllum Müll. Arg.] 

NOTES 

Trichothelium aeneum (Wallr.) R. C. Harris, comb. nov. 

Verrucaria aenea Wallr., Fl. Crypt. Germ. 3: 299. 1831. Porina aenea (Wallr.) Zahlbr., Cat. lich. 

univ. 1: 363. 1922. Pseudosagedia aenea (Wallr.) Hafellner & Kalb, Biblioth. Lichenol. 57: 

172. 1995. Type not seen. 

?Porina amygdalina Müll. Arg., Hedwigia 34: 34. 1895. Type. LOUISIANA. Pointe à la Hache, 

Eckfeldt [Langlois] 797, not seen. 

Trichothelium cestrense (Michener) R. C. Harris, comb. nov. 

Verrucaria cestrensis Tuck. ex Michener in Darlington, Fl. cestr. ed. 3: 452. 1853. Porina 

cestrensis (Michener) Müll. Arg., Flora 64: 338. 1883. Type. PENNSYLVANIA. Chester, Michener 

204 (FH-Tuck 3986, isotype?). 

Porina salicina Müll. Arg., Hedwigia 34: 34. 1895. Type. LOUISIANA. Around New Orleans in 

nearby woods, 24 Nov 1884, Langlois 796 [erroneously attributed to Eckfeldt in original 

publication] (PH, isotype?). 

Porina cestrensis var. platyspora Fink in Hedrick, Mycologia 25: 308. 1933. Type. INDIANA. Near 

Scipio, 8 Feb 1909, Fink (MICH, Fink hb. 8,889). 

In one of my more egregious blunders (as noted by McCarthy, 1993) I misinterpreted the typification of 

Verrucaria pulla Ach. (MülIer made the same mistake.) Instead of referring back to the Lichenographis 

universalis (1810), I looked at the Synopsis methodus lichenum (1814). Different specimens are cited in the 

two works. The first (the holotype, H-Ach 799-A) turns out to be a common (but until now nameless) 

Caribbean Arthopyrenia. The second (BM, H-Ach 799-B) on Dirca is T. cestrense. 

Trichothelium cestrense complex 

This group is one of the reasons that any sane person would avoid this genus. There is a virtually 

complete series of intermediates in ascospore shape and size between T. cestrense and T. 

rhaphidospermum. There are some with cestrense size ascospores but rod-shaped (similar to Porina borreri 

var. leptospora (Nyl.) D. Hawksw.), there are longer acicular ascospores but considerably shorter than 

rhaphidospermum and there are various sizes of fusiform-clavate ascospores longer and with more septa than 

"typical" cestrense. (Brodo's 1968 report of Porina hibernica P. James & Swinscow is based on some of the 

longer spored morphs of this complex . As far as I know P. hibernica does not occur in North America.) I have 

been aware of this mess since my student days but since it was only peripheral to my thesis, I made no 

attempt to resolve it. I still have not had the time (or energy) to try to find some pattern. I hope with new 

computer graphics and statistical analyses that eventually it can be sorted out. In addition to the variation I was 

aware of 20 years ago, the recent emphasis on the color of the ascomatal wall has added yet another 

dimension. The color of the wall in section in this group seems to be either brown or blue-violet. As it does 

not seem to correlate with any of the ascospore variations, it doesn't seem helpful at this time. Also I have no 

idea how specimen drying/preservation affects this character. In Florida there is somewhat less variation than 

farther north and the variation falls into two groups as indicated in the key. A third variant with ascospores 

smaller than usual may be the European "Porina" borreri but I do not have material to resolve this question at 

the moment. 

Trichothelium chloroticum (Ach.) R. C. Harris, comb. nov. 

Verrucaria chlorotica Ach., Lichenogr. univ. 283. 1810. Porina chlorotica (Ach.) Müll. Arg., Rev. 

Mycol. (Toulouse) 6: 20. 1884. Pseudosagedia chlorotica (Ach.) Hafellner & Kalb, Biblioth. 

Lichenol. 57: 172. 1995. Type not seen. 

169

Trichothelium crocynioides R. C. Harris, sp. nov. 

Species unica thallo subbyssoideo, pagina thalli hyphis irregularibus erectis, hypothallo nigro byssoideo 

et ascosporis angustis, 3-septatis, 20-30 × 3-4 µm. 

Type. FLORIDA. Highlands County: Highlands Hammock State Park, swampy area near stream, on cypress, 22 

Dec 1965, Wetmore 13907 (NY, holotype; FLAS, MIN & to be distributed, isotypes). 

Thallus light olive gray, continuous, smooth, dull to almost byssoid in appearance due to short, irregular 

erect hyphae projecting from thallus surface; hypothallus conspicuous, black, byssoid, composed of loose 

interconnected dark brown or gray brown hyphae. Ascomata black, immersed, subglobose, ca. 0.2-0.3 mm 

diam.; wall blackish brown, sometimes with purplish tint, reddish purple in HNO3. Asci cylindrical, 95-110 × 8-9 

µm, with eight subbiseriate spores. Ascospores narrowly fusiform to cylindrical, 4(-7)-celled, 20-35 × 2.5-3.5 

µm; microconidia not found. 

Trichothelium crocynioides is named for its superficial resemblance to Crocynia pyxinoides Nyl. The very 

distinctive thallus with erect, projecting hyphae and byssoid hypothallus may be worthy of generic or 

subgeneric recognition. I prefer to wait until the variation in Trichothelium is more fully understood. Thus far 

all specimens are from Taxodium. 

Additional specimens. FLORIDA. Lee County: Caloosahatchee River State Recreation Area, along Co. Rd. 

78 ca. 3 mi E of Telegraph Creek, 26°43'N, 81°39'W, Taxodium swamp, Buck 22994, Harris 30215, 30246 

(NY). 

Trichothelium guentheri (Flotow) R. C. Harris, comb. nov. 

Verrucaria guentheri Flotow, Bot. Zeitung 8: 575. 1850. Porina guentheri (Flotow) Zahlbr., Cat. 

lich. univ. 1: 384. 1922. Pseudosagedia guentheri (Flotow) Hafellner & Kalb, Biblioth. Lichenol. 

57: 172. 1995. Type not seen. 

This is treated in Europe as a highly variable species. The inclusion of North American collections does 

not appreciably add to the problem. Specimens from Kentucky, Missouri, North Carolina and Virginia are fairly 

homogeneous with clavate-fusiform, 8(-9)-celled ascospores, 32-45 × 5-6 µm. Collections from Maine and 

Massachusetts have larger, more septate ascospores and may prove closer to Porina grandis (Körber) Zahlbr. 

(spp. 7, 8, 9 of my thesis and reported for North America by McCarthy (1993) on this basis). The species is 

apparently mostly semiaquatic except for the Missouri collection which from dry rock. 

Porina hibernica James & Swinscow 

North American records are collections of the Trichothelium cestrense complex lacking isidia and having 

smaller ascomata than P. hibernica. 

Trichothelium isidiatum R. C. Harris, sp. nov. 

Similis T. cestrensi sed thallo dense isidiato et ascosporis fusiformi-clavati, 7-septatis, 43-54 × 6-7.5 µm. 

Type. ILLINOIS. Jackson County: Shawnee National Forest, Pomona Natural Bridge, ca. 8 mi S of 

Murphysboro, ca. 37°38'N, 89°20'W, beech-maple woods, at base of oak, 15 Oct 1993, Harris 31265 (NY, 

holotype). 

Thallus dark gray green, continous but mostly concealed by dense isidia; isidia ± cylindrical, irregular, 

occasionally branched, with a thin, papillose? cortex. Ascomata immersed, almost concealed by isidia, black, 

subglobose, ca. 0.3-0.4 mm diam.; wall brown above with bluish-violet tints in lower part, pale below. Asci 

fusiform-cylindrical, ca. 130-160 × 15-17 µm, with eight subbiseriate spores. Ascospores fusiform-clavate, 8(- 

9)-celled, 43-54 × 6-7.5 µm. Microconidia not found. 

170

Trichothelium isidiatum could be referred to Zamenhofia if one accepts this genus. It is rather similar in 

aspect to Zamenhofia hibernica. It does not have periphyses as far as I can tell. There seems to be one 

isidiate species per genus in North America, Clathroporina isidiifera, Porina scabrida, Segestria sp. (thesis 

Porina sp. 4) and Trichothelium isidiatum. The last is known only from the type collection. 

Trichothelium lineare (Leighton) R. C. Harris, comb. nov. 

Verrucaria linearis Leighton, Brit. angiocarp. lich. 52. 1851. Porina linearis (Leighton) Zahlbr., Cat. 

lich. univ. 1: 391. 1922. Pseudosagedia linearis (Leighton) Hafellner & Kalb, Biblioth. 

Lichenol. 57: 174. 1995. Type not seen. 

Trichothelium lineare is reported from North America but not known in the East. 

Trichothelium nitidulum (Müll. Arg.) R. C. Harris, comb. nov. 

Porina nitidula Müll. Arg., Flora 66: 336. 1883. Pseudosagedia nitidula (Müll. Arg.) Hafellner & 

Kalb, Biblioth. Lichenol. 57: 173. 1995. Type not seen. 

Trichothelium rhaphidospermum (Müll. Arg.) R. C. Harris, comb. nov. 

Porina rhaphidosperma Müll. Arg., Hedwigia 34: 35. 1895. Type. MASSACHUSETTS. On hemlock, 

Willey (FH-Tuck 3988, US, isotypes). 

Trichothelium thaxteri (R. Sant.) R. C. Harris, comb. nov. 

Porina thaxteri R. Sant., Symb. Bot. Upsal. 12(1): 218. 1952. Pseudosagedia thaxteri (R. Sant.) 

Hafellner & Kalb, Biblioth. Lichenol. 57: 173. 1995. Type. FLORIDA. Dade County: Cutler, 

Thaxter 421 (MICH, isotype). 

Trichothelium sp. Thaxter 92 

This taxon is possibly related to the foliicolous Porina moralesiae Lücking but I have no comparative 

material nor have I re-examined the specimens studied for my thesis. It was included as Porina sp. 1. 

INDEX TO SCIENTIFIC NAMES 

Names in boldface represent new nomenclature; names in italics are synonyms, and names in regular typeface 

are accepted names. 

Acrocordia............................................................................122 

Acrocordia cavata................................................................122 

Acrocordia conoidea............................................................122 

Acrocordia gemmata ...........................................................122 

Acrocordia megalospora .....................................................122 

Acrocordiaceae....................................................................121 

Acrocordiella..........................................................................93 

Acrocordiella occulta .............................................................93 

Amandinea.............................................................................45 

Amandinea polyspora............................................................46 

Amandinea punctata .............................................................46 

Anisomeridium.....................................................................123 

Anisomeridium adnexum..................................................143 

Anisomeridium albidoatrum.............................................144 

Anisomeridium albisedum...................................................126 

Anisomeridium ambiguum...................................................126 

Anisomeridium americanum ............................................144 

Anisomeridium angulosum ..............................................144 

Anisomeridium anisolobum...........................................69, 126 

Anisomeridium antillarum ................................................144 

Anisomeridium aureopunctatum .....................................127 

Anisomeridium holopolium ..............................................147 

Anisomeridium immersum ...............................................147 

Anisomeridium indicum....................................................147 

Anisomeridium infernale ..................................................147 

Anisomeridium interspersum...............................................130 

Anisomeridium juistense .....................................................129 

171 

Anisomeridium australiense............................................ 144 

Anisomeridium biforme....................................................... 127 

Anisomeridium biformoides ............................................ 127 

Anisomeridium carinthiacum .............................................. 144 

Anisomeridium complanatum ......................................... 145 

Anisomeridium consimile ................................................ 145 

Anisomeridium consobrinum .............................................. 145 

Anisomeridium distans ....................................................... 127 

Anisomeridium elegans ................................................... 145 

Anisomeridium endobryum ............................................. 146 

Anisomeridium epiphyllum ................................................. 146 

Anisomeridium excaecariae ............................................ 128 

Anisomeridium excellens ................................................ 146 

Anisomeridium feeanum..................................................... 126 

Anisomeridium finkii......................................................... 128 

Anisomeridium flavopallidum.......................................... 146 

Anisomeridium foliicola....................................................... 146 

Anisomeridium glaucescens........................................... 146 

Anisomeridium gregale .................................................... 146 

Anisomeridium griffinii..................................................... 128 

Anisomeridium laevigatum .............................................. 147 

Anisomeridium leptospermum........................................ 147 

Anisomeridium leucochlorum ............................................. 147 

Anisomeridium magnosporum............................................ 148 

Anisomeridium megalosporum....................................... 148 

Anisomeridium monosporum.......................................... 148

Anisomeridium nemorosum.............................................148 

Anisomeridium nidulans...................................................148 

Anisomeridium nyssaegenum.............................................128 

Anisomeridium pacificum.................................................149 

Anisomeridium palavanum...............................................149 

Anisomeridium phaeospermum ......................................129 

Anisomeridium planiusculum ..........................................149 

Anisomeridium polycarpum .............................................149 

Anisomeridium quadrococcum .......................................129 

Anisomeridium quaternarium ..........................................130 

Anisomeridium rockii........................................................149 

Anisomeridium sanfordense................................................128 

Anisomeridium sphaerocarpum ......................................149 

Anisomeridium stromaticum............................................149 

Anisomeridium subatomarium ........................................150 

Anisomeridium subbiforme..............................................150 

Anisomeridium subnectendum........................................150 

Anisomeridium subnexum ...............................................150 

Anisomeridium subprostans................................................130 

Anisomeridium tamarindi.....................................................130 

Anisomeridium tarmugliense...........................................150 

Anisomeridium terminatum..............................................130 

Anisomeridium throwerae ................................................150 

Anisomeridium truncatum................................................151 

Anisomeridium tuckerae......................................................131 

Anisomeridium ubianum ..................................................151 

Anisomeridium uniseriale.................................................151 

Anisomeridium verrucosum.............................................151 

Anisomeridium viridescens..............................................151 

Anthracothecium....................................................................93 

Anthracothecium albescens................................................109 

Anthracothecium albidopallens...........................................110 

Anthracothecium americanum ............................................110 

Anthracothecium canellae-albae...........................................94 

Anthracothecium columellatum.............................................87 

Anthracothecium confine.....................................................109 

Anthracothecium corticatum................................................109 

Anthracothecium nanum .......................................................94 

Anthracothecium prasinum .............................................87, 93 

Anthracothecium staurosporum ............................................94 

Anthracothecium subglobosum.............................................94 

Anthracothecium varians.......................................................93 

Anzia ......................................................................................40 

Anzia ornata...........................................................................40 

Apiospora polypori...............................................................128 

Architrypethelium.................................................................114 

Arthonia tumidula...................................................................68 

Arthoniales...............................................................................2 

Arthopyrenia ..........................................................................75 

Arthopyrenia sect. Anisomeridium .....................................123 

Arthopyrenia sect. Pseudosagedia ....................................176 

Arthopyrenia adnexa var. leptosperma...............................147 

Arthopyrenia adnexa ...........................................................143 

Arthopyrenia albidoatra .......................................................144 

Arthopyrenia alboatra..........................................................127 

Arthopyrenia amaura...........................................................127 

Arthopyrenia ambigua .........................................................126 

Arthopyrenia americana ......................................................144 

Arthopyrenia megalospora..............................................62, 82 

Arthopyrenia minor ................................................................82 

Arthopyrenia mycoporoides ..................................................69 

Arthopyrenia nidulans..........................................................148 

Arthopyrenia nieteriana .........................................................80 

Arthopyrenia nylanderi ..........................................................72 

172 

Arthopyrenia anacardii........................................................ 130 

Arthopyrenia analepta........................................................... 78 

Arthopyrenia anisoloba....................................................... 126 

Arthopyrenia annulata .......................................................... 79 

Arthopyrenia antecellens................................................ 67, 79 

Arthopyrenia antillarum....................................................... 144 

Arthopyrenia atomarioides.............................................. 62, 79 

Arthopyrenia atractospora .............................................. 62, 79 

Arthopyrenia bifera ......................................................... 79, 82 

Arthopyrenia biformis.......................................................... 127 

Arthopyrenia brachyspora .................................................. 127 

Arthopyrenia carinthiaca..................................................... 144 

Arthopyrenia carpinea ........................................................ 176 

Arthopyrenia cedrina............................................................. 79 

Arthopyrenia cerasi............................................................... 79 

Arthopyrenia ceuthocarpoides.............................................. 72 

Arthopyrenia ceylonensis ................................................... 144 

Arthopyrenia cinchonae........................................................ 80 

Arthopyrenia cinefaciens .................................................... 155 

Arthopyrenia cineropruinosa................................................. 79 

Arthopyrenia confluens................................................ 69, 80 

Arthopyrenia conformis....................................................... 127 

Arthopyrenia consanguinea.................................................. 69 

Arthopyrenia consimilis....................................................... 145 

Arthopyrenia consobrina var. glaucescens........................ 151 

Arthopyrenia consobrina..................................................... 145 

Arthopyrenia convexella ....................................................... 79 

Arthopyrenia degelii ........................................................... 80 

Arthopyrenia dimidiata........................................................ 144 

Arthopyrenia distans........................................................... 128 

Arthopyrenia endobrya ....................................................... 146 

Arthopyrenia epigloea........................................................... 72 

Arthopyrenia esenbeckiana......................................... 69, 81 

Arthopyrenia exasperata.................................................... 81 

Arthopyrenia excaecariae................................................... 128 

Arthopyrenia exstans.......................................................... 150 

Arthopyrenia fallax ................................................................ 79 

Arthopyrenia fallax f. crategina............................................. 79 

Arthopyrenia fallax var. conspurcata.................................... 80 

Arthopyrenia feeana ........................................................... 126 

Arthopyrenia floridana .......................................................... 62 

Arthopyrenia fraxini......................................................... 62, 81 

Arthopyrenia glaucescens .................................................. 146 

Arthopyrenia glaucina......................................................... 130 

Arthopyrenia griseola............................................................ 67 

Arthopyrenia incurva........................................................... 128 

Arthopyrenia infernalis........................................................ 147 

Arthopyrenia infernalis var. rhychostoma........................... 147 

Arthopyrenia insularis ......................................................... 130 

Arthopyrenia japonica........................................................... 80 

Arthopyrenia knysana......................................................... 126 

Arthopyrenia lapponina................................................... 78, 81 

Arthopyrenia leucochlora.................................................... 147 

Arthopyrenia ligustri.............................................................. 80 

Arthopyrenia limitans .......................................................... 144 

Arthopyrenia lyrata................................................................ 81 

Arthopyrenia majuscula ........................................................ 82 

Arthopyrenia malaccitula ...................................................... 82 

Arthopyrenia media............................................................. 144 

Arthopyrenia oblongens..................................................... 82 

Arthopyrenia padi............................................................ 63, 82 

Arthopyrenia parvula........................................................... 127 

Arthopyrenia pinicola ............................................................ 79 

Arthopyrenia planior.............................................................. 82 

Arthopyrenia planipes........................................................... 80

Arthopyrenia planiuscula.....................................................149 

Arthopyrenia planorbiculata ..................................................82 

Arthopyrenia planorbis ..........................................................82 

Arthopyrenia plumbaria .........................................................83 

Arthopyrenia pongamiae .....................................................149 

Arthopyrenia punctiformis .....................................................63 

Arthopyrenia quassiaecola....................................................63 

Arthopyrenia rappii ................................................................83 

Arthopyrenia rhyponta.....................................................63, 83 

Arthopyrenia salicis ...............................................................83 

Arthopyrenia sanfordensis .................................................128 

Arthopyrenia sanfordensis var. cedrina ...............................62 

Arthopyrenia saxicola ...........................................................63 

Arthopyrenia stigmatella var. elabens ..................................79 

Arthopyrenia subantecellens ................................................67 

Arthopyrenia subatomaria ..................................................150 

Arthopyrenia subbiformis ...................................................150 

Arthopyrenia subinsularis ...................................................130 

Arthopyrenia subnectenda .................................................150 

Arthopyrenia subnexa ........................................................150 

Arthopyrenia subprostans ..................................................130 

Arthopyrenia taxodii............................................................83 

Arthopyrenia terminata .......................................................130 

Arthopyrenia texensis............................................................83 

Arthopyrenia transposita ......................................................80 

Arthopyrenia truncata .........................................................151 

Arthopyrenia ubiana ...........................................................151 

Arthopyrenia viridescens ....................................................151 

Arthopyrenia willeyana .......................................................129 

Arthopyrenia xylogena ................................................123, 150 

Arthopyrenia zonata .............................................................73 

Arthopyreniaceae ..................................................................75 

Arthothelium tumidulum ........................................................68 

Ascocratera..........................................................................121 

Aspidopyrenium ..................................................................162 

Aspidotheliaceae ................................................................160 

Aspidothelium .....................................................................162 

Aspidothelium cinerascens ................................................165 

Aspidothelium trichotheliodes ............................................167 

Astrothelium.........................................................................114 

Astrothelium cinnamomeum................................................114 

Astrothelium confusum ...............................................115, 121 

Astrothelium diplocarpum....................................................115 

Astrothelium epiphyllum ......................................................133 

Astrothelium galbineum.......................................................114 

Astrothelium variolosum .............................................115, 121 

Astrothelium versicolor........................................................114 

Athrismidium..........................................................................65 

Bactrospora .............................................................................3 

Bactrospora brevispora ...........................................................4 

Bactrospora denticulata.......................................................3, 4 

Bactrospora dryina .................................................................4 

Bactrospora integrispora ........................................................4 

Bactrospora macrospora.........................................................4 

Bactrospora mesospora ..........................................................3 

Bactrospora myriadea .............................................................3 

Bactrospora nematospora.......................................................4 

Baeomyces ............................................................................28 

Baeomyces roseus ................................................................28 

Baeomycetaceae...................................................................28 

Canoparmelia caroliniana .....................................................40 

Canoparmelia cryptochlorophaea .........................................40 

Canoparmelia martinicana ....................................................40 

Canoparmelia salacinifera ....................................................40 

Canoparmelia texana ............................................................40 

Catapyrenium ........................................................................56 

Catapyrenium squamulosum ................................................56 

173 

Bathelium ............................................................................ 115 

Bathelium albidoporum.................................................... 116 

Bathelium carolinianum ................................................... 116 

Bathelium degenerans ..................................................... 117 

Bathelium endochryseum................................................ 117 

Bathelium lineare .............................................................. 117 

Bathelium madreporiforme ................................................. 117 

Bathelium mastoideum ....................................................... 117 

Bathelium sphaericum ..................................................... 118 

Bathelium tuberculosum.................................................. 118 

Beckhausia ........................................................................... 65 

Biatora leptalea................................................................... 175 

Bottaria.................................................................................. 65 

Bottaria albidopallens ......................................................... 110 

Bottaria composita ................................................................ 68 

Bottaria libricola f. nana ...................................................... 110 

Bottaria mucosa.................................................................. 110 

Buellia ................................................................................... 45 

Buellia amphidexia................................................................ 46 

Buellia caloosensis ............................................................... 46 

Buellia catasema .................................................................. 46 

Buellia coccinea.................................................................... 46 

Buellia curatellae .................................................................. 45 

Buellia curtisii........................................................................ 45 

Buellia imshaugiana ............................................................. 45 

Buellia lauricassiae............................................................... 46 

Buellia leucomela.................................................................. 45 

Buellia melanochlora ............................................................ 46 

Buellia pachnidisca............................................................... 46 

Buellia placodiomorpha ........................................................ 51 

Buellia rappii ......................................................................... 46 

Buellia rubifaciens................................................................. 47 

Buellia stillingiana ................................................................. 46 

Buellia wheeleri..................................................................... 47 

Bulbothrix .............................................................................. 40 

Bulbothrix confoederata........................................................ 40 

Bulbothrix coronata............................................................... 40 

Bulbothrix goebelii ................................................................ 40 

Bulbothrix isidiza ................................................................... 40 

Bulbothrix laevigatula............................................................ 40 

Byssoloma............................................................................. 31 

Byssoloma leucoblepharum ................................................. 31 

Byssoloma pubescens....................................................... 31 

Byssoloma subdiscordans.................................................... 31 

Byssoloma tricholomum........................................................ 31 

Byssothecium ....................................................................... 74 

Caliciaceae ........................................................................... 22 

Caliciales............................................................................... 20 

Calicium ................................................................................ 22 

Calicium adspersum ............................................................. 22 

Calicium chlorosporum ......................................................... 22 

Calicium hyperelloides.......................................................... 22 

Calicium lenticulare............................................................... 22 

Calicium leucochlorum ......................................................... 22 

Calicium salicinum................................................................ 22 

Campylothelium .................................................................. 113 

Campylothelium decolorans ................................................. 87 

Canoparmelia........................................................................ 40 

Canoparmelia amazonica..................................................... 40 

Catapyrenium tuckermanii.................................................... 56 

Celothelium ......................................................................... 161 

Celothelium aciculiferum .................................................... 162 

Celothelium cinchonarum................................................... 162 

Celothelium dominicanum .................................................. 161 

Celothelium leprieurii .......................................................... 161 

Celothelium socialis............................................................ 161

Chaenotheca .........................................................................24 

Chaenotheca brunneola........................................................24 

Chaenotheca floridana........................................................24 

Chaenotheca hispidula....................................................24, 25 

Chaenothecopsis.............................................................26, 27 

Chaenothecopsis nana .........................................................26 

Chaenothecopsis norstictica .......................................26, 27 

Chaenothecopsis pilosa ..................................................26, 27 

Chaenothecopsis rappii................................................26, 27 

Chaenothecopsis savonica ...................................................26 

Chaetothyriales......................................................................56 

Chiodecton...............................................................................4 

Chiodecton malmei..............................................................2, 4 

Chiodecton perplexum ............................................................4 

Chlorodothis ..........................................................................65 

Chromatochlamys................................................................162 

Chromatochlamys muscorum .............................................162 

Chromatochlamys muscorum var. octosporum ..................162 

Cifferiolichen..........................................................................75 

Cladina...................................................................................32 

Cladina evansii ......................................................................32 

Cladina rangiferina ................................................................32 

Cladina sandstedei................................................................32 

Cladina subtenuis..................................................................32 

Cladina subtenuis f. cinerea..................................................32 

Cladonia.................................................................................33 

Cladonia abbreviatula ...........................................................33 

Cladonia anitae......................................................................33 

Cladonia bacillaris .................................................................33 

Cladonia beaumontii..............................................................34 

Cladonia botryocarpa ............................................................34 

Cladonia buckii ......................................................................34 

Cladonia caroliniana..............................................................34 

Cladonia chlorophaea ...........................................................35 

Cladonia cristatella ................................................................33 

Cladonia cryptochlorophaea .................................................35 

Cladonia cylindrica ................................................................35 

Cladonia didyma var. didyma................................................33 

Cladonia didyma var. vulcanica ............................................33 

Cladonia dimorphoclada .......................................................34 

Cladonia floridana .................................................................34 

Cladonia grayi..................................................................34, 35 

Cladonia hypoxantha.............................................................33 

Cladonia incrassata...............................................................33 

Cladonia leporina ..................................................................33 

Cladonia nana .......................................................................36 

Cladonia pachycladodes .......................................................34 

Cladonia perforata.................................................................34 

Cladonia peziziformis ............................................................35 

Cladonia polycarpia...............................................................34 

Cladonia polycarpoides.........................................................34 

Cladonia prostrata .................................................................36 

Cladonia ramulosa ................................................................35 

Cladonia rappii var. exilior.....................................................35 

Cladonia rappii var. rappii .....................................................35 

Cladonia ravenelii..................................................................33 

Cladonia santensis ................................................................34 

Cladonia simulata..................................................................35 

Cladonia sobolescens ...........................................................35 

Cladonia strepsilis .................................................................35 

Diporina subsimplicans .......................................................152 

Dirinaria .................................................................................47 

Dirinaria aegialita...................................................................47 

Dirinaria applanata ................................................................47 

Dirinaria confluens.................................................................47 

Dirinaria confusa ...................................................................47 

Dirinaria leopoldii...................................................................47 

Dirinaria papillulifera..............................................................47 

174 

Cladonia subradiata.............................................................. 35 

Cladonia subsetacea ............................................................ 34 

Cladoniaceae........................................................................ 32 

Clathroporina ...................................................................... 171 

Clathroporina amygdalina .................................................... 88 

Clathroporina confinis......................................................... 171 

Clathroporina exiguella......................................................... 88 

Clathroporina isidiifera .................................................... 171 

Clathroporina mastoidea ................................................. 172 

Clathroporina nuculastrum ................................................. 174 

Clathroporina subpungens.............................................. 172 

Clathroporina tetracerae.................................................. 172 

Clypeolum talumae............................................................. 152 

Clypeopyrenis ....................................................................... 93 

Clypeosphaeriaceae ........................................................... 159 

Collema ................................................................................. 28 

Collema callibotrys................................................................ 29 

Collema conglomeratum....................................................... 29 

Collema furfurceum var. luzone ........................................... 29 

Collema leptaleum................................................................ 29 

Collema pulcellum var. pulcellum......................................... 28 

Collema pulcellum var. leucopeplum ................................... 28 

Collema pulcellum var. subnigrescens ................................ 28 

Collema pustulatum.............................................................. 29 

Collema subflaccidum .......................................................... 29 

Collemataceae...................................................................... 28 

Collemopsidium .................................................................... 71 

Collemopsidium atlanticum .................................................. 72 

Collemopsidium iocarpum .................................................... 72 

Compsosporiella ................................................................. 146 

Cresponea .............................................................................. 4 

Cresponea chloroconia........................................................... 5 

Cresponea flava...................................................................... 5 

Cresponea proximata ............................................................. 4 

Cryptothelium.............................................................. 114, 118 

Cyrtidula ................................................................................ 65 

Cyrtidula hippocastani ....................................................... 65 

Cyrtidula macularis ............................................................... 65 

Cyrtidula pteleodes ............................................................... 65 

Dacampia.............................................................................. 74 

Dacampiaceae...................................................................... 74 

Dermatina ohiensis............................................................... 69 

Dermatina pyrenocarpa ........................................................ 68 

Dermatocarpella ................................................................... 56 

Dibaeis baeomyces .............................................................. 28 

Dichoporis ........................................................................... 152 

Dichoporis schizospora....................................................... 152 

Didymella gigantea ............................................................... 82 

Didymella labiata ................................................................ 144 

Didymella laurilae ............................................................... 127 

Didymella microspora ......................................................... 127 

Didymella sphaerocarpa..................................................... 149 

Didymellopsis........................................................................ 71 

Didymosphaeria megalospora............................................ 148 

Diploschistes......................................................................... 16 

Diploschistes hypoleucus ..................................................... 16 

Diploschistes muscorum....................................................... 16 

Diporina............................................................................... 152 

Dirinaria picta ........................................................................ 47 

Dirinaria purpurascens ......................................................... 47 

Discosiella........................................................................... 152 

Discosiella cylindrospora.................................................... 152 

Distopyrenis .......................................................................... 94 

Distopyrenis americana ........................................................ 95 

Distopyrenis composita ..................................................... 95 

Distopyrenis fuliginosa.......................................................... 96

Distopyrenis pachyospora .....................................................96 

Distopyrenis quercicola......................................................96 

Distopyrenis submuriformis ..............................................96 

Ditremis................................................................................123 

Ditremis aggregata ..............................................................145 

Ditremis albiseda.................................................................126 

Ditremis ambigua.................................................................126 

Ditremis andamanica...........................................................149 

Ditremis anisoloba...............................................................126 

Ditremis anisomera .............................................................147 

Ditremis australiensis ..........................................................144 

Ditremis biformis..................................................................127 

Ditremis bispora...................................................................144 

Ditremis complanata............................................................145 

Ditremis distans ...................................................................128 

Ditremis epiphylla ................................................................146 

Ditremis finkii .......................................................................128 

Ditremis flavopallida ............................................................146 

Ditremis harrisii....................................................................145 

Ditremis immersa.................................................................147 

Ditremis indica .....................................................................147 

Ditremis inspersa.................................................................144 

Ditremis karnatakensis........................................................145 

Ditremis laevigata................................................................147 

Ditremis leucochlora............................................................147 

Ditremis macrospora ...................................................127, 147 

Ditremis microcarpa ............................................................151 

Ditremis monospora ............................................................148 

Ditremis muelleri..................................................................145 

Ditremis nyssaegena...........................................................128 

Ditremis pacifica ..................................................................149 

Ditremis polycarpa...............................................................149 

Ditremis punctata.................................................................149 

Ditremis pyrenuloides..........................................................144 

Ditremis quaternaria ............................................................130 

Ditremis rockii ......................................................................149 

Ditremis sanfordensis..........................................................128 

Ditremis straminicolor..........................................................145 

Ditremis subprostans...........................................................130 

Ditremis tamarindi................................................................130 

Ditremis tarmugliensis.........................................................150 

Ditremis terminata ...............................................................130 

Ditremis tuckerae.................................................................131 

Ditremis verrucosa.......................................................123, 151 

Endocarpon ...........................................................................57 

Endocarpon pusillum.............................................................57 

Enterographa...........................................................................5 

Enterographa lecanoroides.....................................................5 

Enterographa quassiaecola ....................................................5 

Eopyrenula.......................................................................74, 92 

Eopyrenula intermedia ..........................................................74 

Eopyrenula leucoplaca ..........................................................74 

Eopyrenula parvispora ..........................................................74 

Exiliseptum ..........................................................................114 

Foraminella............................................................................40 

Geisleria...............................................................................159 

Geisleria sychnogonioides ..................................................159 

Hafellia bahiana.....................................................................45 

Hafellia bahiana var. pleiotropa ............................................46 

Hafellia callispora ..................................................................45 

Hafellia parastata ..................................................................45 

Heterodermia.........................................................................47 

Heterodermia albicans ..........................................................47 

Heterodermia barbifera .........................................................48 

Heterodermia casarettiana....................................................48 

Heterodermia crocea.............................................................48 

Heterodermia echinata..........................................................48 

175 

Glyphis .................................................................................... 9 

Glyphis cicatricosa .................................................................. 9 

Granulopyrenis...................................................................... 97 

Granulopyrenis antillensis .................................................... 98 

Granulopyrenis hymnothora ................................................. 98 

Granulopyrenis macrocarpoides .......................................... 98 

Granulopyrenis nigrescens ............................................... 98 

Graphidaceae ......................................................................... 8 

Graphidales............................................................................. 8 

Graphina ................................................................................. 9 

Graphina abaphoides ............................................................. 9 

Graphina adscribens............................................................. 11 

Graphina anguina ................................................................. 10 

Graphina antillarum .............................................................. 10 

Graphina columbina.............................................................. 10 

Graphina columbina ............................................................. 11 

Graphina cypressi ................................................................... 9 

Graphina floridana ................................................................ 10 

Graphina incrustans.............................................................. 10 

Graphina intertexta ............................................................... 11 

Graphina leuconephela ........................................................ 10 

Graphina marescens ............................................................ 10 

Graphina parilis..................................................................... 10 

Graphina peplophora.............................................................. 9 

Graphina platyleuca.............................................................. 11 

Graphina scolecitis ............................................................... 10 

Graphina subvirginalis ...................................................... 9, 11 

Graphina virginea ................................................................... 9 

Graphina xylophaga.............................................................. 10 

Graphis.................................................................................. 11 

Graphis afzelii ....................................................................... 11 

Graphis anfractuosa ............................................................. 12 

Graphis anguinoides....................................................... 10, 13 

Graphis caesiella .................................................................. 12 

Graphis candidata................................................................. 11 

Graphis chromothecia........................................................... 12 

Graphis desquamescens...................................................... 12 

Graphis grammatis ............................................................... 13 

Graphis haleana ................................................................... 12 

Graphis illiterata.................................................................... 13 

Graphis insidiosa .................................................................. 13 

Graphis inversa..................................................................... 12 

Graphis leptocarpa ............................................................... 12 

Graphis librata....................................................................... 12 

Graphis lineola...................................................................... 11 

Graphis lucifica ..................................................................... 12 

Graphis lumbricina................................................................ 12 

Graphis pavoniana................................................................ 11 

Graphis rufula ....................................................................... 13 

Graphis scripta...................................................................... 11 

Graphis striatula.................................................................... 12 

Graphis subelegans.............................................................. 12 

Graphis subnitidula ............................................................... 13 

Graphis subparilis ................................................................. 13 

Graphis turbulenta ................................................................ 13 

Gyrostomum.......................................................................... 13 

Gyrostomum scyphuliferum.................................................. 13 

Hafellia ............................................................................ 45, 47 

Heterodermia galactophylla.................................................. 48 

Heterodermia japonica ......................................................... 48 

Heterodermia leucomelos..................................................... 48 

Heterodermia obscurata ....................................................... 48 

Heterodermia speciosa......................................................... 47 

Hyperphyscia ........................................................................ 48 

Hyperphyscia adglutinata ..................................................... 48 

Hyperphyscia minor .............................................................. 48 

Hyperphyscia syncolla .......................................................... 48 

Hypotrachyna ........................................................................ 41

Hypotrachyna livida ...............................................................41 

Hypotrachyna osseoalba.......................................................41 

Hypotrachyna pustulifera.......................................................41 

Icmadophilaceae ...................................................................28 

Imshaugia ..............................................................................41 

Imshaugia aleurites ...............................................................41 

Jarxia .....................................................................................60 

Jarxia ilicicola ......................................................................60 

Jarxia thelenula .....................................................................60 

Julella.....................................................................................84 

Julella asema .......................................................................86 

Julella avicenniae ..................................................................86 

Julella buxi .............................................................................86 

Julella dactylospora...............................................................87 

Julella decolorans ...............................................................87 

Julella dispora......................................................................87 

Julella exiguella ...................................................................87 

Julella fallaciosa ....................................................................87 

Julella geminella..................................................................87 

Julella leopoldina...................................................................87 

Julella luzonensis ..................................................................87 

Julella macrospora ................................................................88 

Julella manakonensis............................................................88 

Julella phycophila ..................................................................88 

Julella sericea........................................................................88 

Julella sublactea....................................................................88 

Julella taxodii .......................................................................88 

Julella variiformis ................................................................89 

Julella zenkeriana..................................................................90 

Jullela vitrispora.....................................................................90 

Lachrymospora......................................................................93 

Lauderlindsaya ......................................................................57 

Laurera ................................................................................118 

Laurera elegans...................................................................117 

Laurera linearis....................................................................117 

Laurera madreporiformis.....................................................117 

Laurera marginata ...............................................................117 

Laurera megasperma ....................................................90, 118 

Laurera pyriformis................................................................118 

Laurera sphaerica ...............................................................118 

Laurera subdisjuncta ...........................................................118 

Laurera submadreporiformis...............................................118 

Laurera tuberculosa ............................................................118 

Lecanactis................................................................................5 

Lecanactis epileuca.................................................................5 

Lecanactis subattingens..........................................................6 

Lecania fugiens ...................................................................165 

Lecanographa..........................................................................2 

Lecanorales ...........................................................................28 

Leiophloea .............................................................................75 

Leiophloea adnexa..............................................................143 

Leiophloea pongamiae........................................................149 

Leiophloea truncata.............................................................151 

Leiophloea xylogena ...........................................................150 

Lembidium ...........................................................................123 

Lembidium polycarpum .......................................................123 

Microtheliopsidaceae.............................................................56 

Monoblastia .........................................................................131 

Monoblastia borinquensis................................................132 

Monoblastia buckii...............................................................131 

Monoblastia cypressi ........................................................132 

Monoblastia echinulospora .................................................131 

Monoblastia palmicola.........................................................131 

Monoblastia pellucida..........................................................131 

Monoblastia quisqueyana.................................................132 

Monoblastia rappii ...............................................................131 

Monoblastiaceae .................................................................121 

176 

Leotiales................................................................................ 28 

Leptogium ............................................................................. 29 

Leptogium adpressum .......................................................... 30 

Leptogium austroamericanum.............................................. 29 

Leptogium azureum .............................................................. 29 

Leptogium chloromelum ....................................................... 30 

Leptogium corticola............................................................... 30 

Leptogium cyanescens......................................................... 29 

Leptogium floridanum ........................................................... 30 

Leptogium fusisporum .......................................................... 30 

Leptogium isidiosellum ......................................................... 29 

Leptogium juniperinum ......................................................... 29 

Leptogium marginellum ........................................................ 29 

Leptogium microstictum........................................................ 30 

Leptogium milligranum ......................................................... 29 

Leptogium phyllocarpum ...................................................... 30 

Leptogium sessile ................................................................. 30 

Leptogium stipitatum............................................................. 30 

Leptogium tenuissimum........................................................ 30 

Leptorhaphis ......................................................................... 60 

Leptorhaphis amygdali.......................................................... 60 

Leptorhaphis parameca........................................................ 60 

Lichen analeptus................................................................... 78 

Lichenothelia......................................................................... 73 

Lithothelium........................................................................... 98 

Lithothelium almbornii........................................................... 98 

Lithothelium bahamense .............................................. 98, 155 

Lithothelium concameratum ................................................. 99 

Lithothelium fugax................................................................. 99 

Lithothelium illotum............................................................... 99 

Lithothelium microsporum ................................................ 99 

Lithothelium obtectum........................................................... 99 

Lithothelium polysemum....................................................... 99 

Lophiostoma subcollapsa................................................... 112 

Luykenia.............................................................................. 162 

Mazaediothecium.................................................................. 93 

Mazosia................................................................................... 5 

Mazosia ocellata ..................................................................... 5 

Megalotremis .............................................................. 112, 123 

Megalotremis biocellata...................................................... 113 

Megalotremis megalospora ................................................ 148 

Megalotremis verrucosa ............................................. 112, 151 

Melanommatales .................................................................. 91 

Melanopsamma corticola.................................................... 129 

Melanotheca esenbeckiana.................................................. 81 

Melanotheca wrightii........................................................... 111 

Microglaena ........................................................................ 162 

Microglaena albolinita......................................................... 155 

Microglaena sychnogonioides............................................ 167 

Microglaena wallrothiana.................................................... 162 

Microglaenaceae ................................................................ 160 

Microthelia magnifica .......................................................... 148 

Microthelia palavana........................................................... 149 

Microthelia queenslandica.................................................. 113 

Microthelia uniserialis ......................................................... 151 

Musaespora ........................................................................ 133 

Musaespora corticola.......................................................... 133 

Musaespora epiphylla ...................................................... 133 

Musaespora gigas............................................................. 133 

Musaespora multilocularis.................................................. 133 

Mycocaliciaceae.................................................................... 26 

Mycocalicium................................................................... 26, 27 

Mycocalicium albonigrum ..................................................... 26 

Mycocalicium calicioides ...................................................... 27 

Mycocalicium rappii .............................................................. 27 

Mycocalicium ravenelii.................................................... 26, 27

Mycocalicium subtile..............................................................26 

Mycocifferia............................................................................75 

Mycoglaena collosporella......................................................88 

Mycoglaena yasudae ............................................................87 

Mycomicrothelia.....................................................................90 

Mycomicrothelia oleosa.........................................................90 

Mycomicrothelia rubrostoma .................................................90 

Mycomicrothelia subfallens ...................................................90 

Mycomicrothelia willeyana.....................................................90 

Mycoporaceae .......................................................................64 

Mycoporellum ..................................................................64, 65 

Mycoporellum californicum....................................................68 

Mycoporellum deserticola .....................................................69 

Mycoporellum difforme ..........................................................69 

Mycoporellum eschweileri .....................................................69 

Mycoporellum gigas ............................................................133 

Mycoporellum hassei.............................................................69 

Mycoporellum lacteum ..........................................................69 

Mycoporellum lahmii..............................................................69 

Mycoporellum leucoplacum...................................................69 

Mycoporellum sparsellum .....................................................70 

Mycoporopsis.........................................................................64 

Mycoporopsis antecellens.....................................................67 

Mycoporopsis californica.......................................................68 

Mycoporopsis deserticola......................................................69 

Mycoporopsis exigua.............................................................69 

Mycoporopsis leucoplaca......................................................69 

Mycoporopsis rappii ........................................................70, 83 

Mycoporum ............................................................................65 

Mycoporum acervatum .......................................................66 

Mycoporum antecellens......................................................67 

Mycoporum buckii ...............................................................68 

Mycoporum californicum....................................................68 

Mycoporum compositum .......................................................68 

Mycoporum difforme..............................................................69 

Mycoporum eschweileri......................................................69 

Mycoporum integrum.............................................................68 

Mycoporum lacteum............................................................69 

Mycoporum mycoporoides.................................................69 

Mycoporum ohiense ..............................................................69 

Mycoporum pycnocarpoides .................................................70 

Mycoporum pycnocarpum var. ohiense................................69 

Mycoporum pycnocarpum.....................................................68 

Mycoporum sparsellum .........................................................70 

Mycoporum uniloculatum...................................................70 

Mycopyrenula ......................................................................100 

Mycopyrenula coryli .............................................................100 

Mycosphaerellaceae .............................................................56 

Myelochroa ............................................................................41 

Myelochroa aurulenta............................................................41 

Myriangium duriaei ................................................................28 

Myriotrema.............................................................................16 

Myriotrema bahianum............................................................17 

Myriotrema compunctum.......................................................16 

Myriotrema erodens...............................................................17 

Ornatopyrenis ........................................................................90 

Parapyrenis..........................................................................100 

Parapyrenis aurora ..............................................................100 

Parapyrenis conica ..............................................................100 

Parapyrenis elongata ............................................................98 

Parapyrenis guayaci ............................................................100 

Parapyrenis maritima...........................................................100 

Parapyrenis tecomatis.........................................................100 

Parmeliaceae.........................................................................38 

Parmelinopsis ........................................................................41 

Parmelinopsis cryptochlora ...................................................41 

Parmelinopsis horrescens.....................................................41 

177 

Myriotrema glaucescens....................................................... 16 

Myriotrema glauculum .......................................................... 17 

Myriotrema peninsulae ......................................................... 17 

Myriotrema reclusum ............................................................ 17 

Myriotrema rugiferum............................................................ 17 

Myriotrema subcompunctum ................................................ 16 

Myriotrema terebratulum....................................................... 17 

Myriotrema wightii ................................................................. 17 

Nadvornikia ........................................................................... 17 

Nadvornikia hawaiiensis....................................................... 17 

Nadvornikia sorediata........................................................... 17 

Naetrocymbaceae ............................................................... 59 

Naetrocymbe......................................................................... 61 

Naetrocymbe atomarioides................................................ 62 

Naetrocymbe atractospora ................................................ 62 

Naetrocymbe cedrina ......................................................... 62 

Naetrocymbe fraxini ........................................................... 62 

Naetrocymbe massalongiana............................................ 62 

Naetrocymbe megalospora................................................ 62 

Naetrocymbe punctiformis ................................................ 63 

Naetrocymbe quassiaecola ............................................... 63 

Naetrocymbe rhyponta....................................................... 63 

Naetrocymbe saxicola........................................................ 63 

Neocatapyrenium.................................................................. 57 

Normandina pulchella........................................................... 57 

Ocellularia ............................................................................. 17 

Ocellularia americana........................................................... 18 

Ocellularia auberiana............................................................ 18 

Ocellularia cavata ................................................................. 18 

Ocellularia concolor .............................................................. 18 

Ocellularia emersa.......................................................... 17, 18 

Ocellularia fissa .................................................................... 18 

Ocellularia granulosa............................................................ 17 

Ocellularia leiostoma ............................................................ 18 

Ocellularia praestans............................................................ 18 

Ocellularia retispora.............................................................. 18 

Ocellularia sanfordiana......................................................... 18 

Ocellularia stictidea............................................................... 16 

Ochrolechia........................................................................... 51 

Ochrolechia africana............................................................. 51 

Ochrolechia antillarum.......................................................... 51 

Opegrapha .............................................................................. 6 

Opegrapha astraea................................................................. 8 

Opegrapha atra....................................................................... 6 

Opegrapha aurantiaca............................................................ 6 

Opegrapha candida ................................................................ 6 

Opegrapha cypressi................................................................ 6 

Opegrapha filicina................................................................... 8 

Opegrapha leucoplaca ........................................................... 7 

Opegrapha longissima............................................................ 7 

Opegrapha varia ..................................................................... 7 

Opegrapha viridis.................................................................... 7 

Opegrapha vulgata ................................................................. 7 

Opegraphaceae ...................................................................... 2 

Parmelinopsis minarum ........................................................ 41 

Parmelinopsis spumosa ....................................................... 41 

Parmeliopsis ......................................................................... 41 

Parmeliopsis subambigua .................................................... 41 

Parmotrema .......................................................................... 41 

Parmotrema crinitum ............................................................ 42 

Parmotrema cristiferum ........................................................ 42 

Parmotrema dilatatum .......................................................... 42 

Parmotrema dominicanum ................................................... 42 

Parmotrema endosulphureum.............................................. 41 

Parmotrema gardeneri.......................................................... 42 

Parmotrema hypoleucinum................................................... 42

Parmotrema mellissii .............................................................42 

Parmotrema michauxianum ..................................................42 

Parmotrema perforatum ........................................................42 

Parmotrema praesorediosum................................................42 

Parmotrema rampoddense ...................................................42 

Parmotrema rigidum..............................................................42 

Parmotrema rubifaciens ........................................................42 

Parmotrema sulphuratum......................................................42 

Parmotrema tinctorum...........................................................41 

Parmotrema ultralucens ........................................................42 

Parmotrema xanthinum .........................................................42 

Parmotrema zollingeri ...........................................................42 

Patellonectria.......................................................................162 

Peltosphaeria.........................................................................84 

Peltosphaeria californica .......................................................85 

Peltosphaeria sandwichensis................................................84 

Pertusaria ..............................................................................51 

Pertusaria amara ...................................................................52 

Pertusaria copiosa.................................................................51 

Pertusaria epixantha .............................................................52 

Pertusaria expolita.................................................................52 

Pertusaria floridana ...............................................................51 

Pertusaria hypothamnolica....................................................51 

Pertusaria iners .....................................................................52 

Pertusaria leucostoma...........................................................52 

Pertusaria multipunctoides....................................................52 

Pertusaria obruta ...................................................................53 

Pertusaria paratuberculifera..................................................52 

Pertusaria pulchella...............................................................51 

Pertusaria pustulata ..............................................................53 

Pertusaria sinusmexicani ......................................................52 

Pertusaria subpertusa ...........................................................53 

Pertusaria tetrathalamia ........................................................52 

Pertusaria texana ..................................................................52 

Pertusaria trachythallina........................................................52 

Pertusaria velata....................................................................52 

Pertusaria ventosa.................................................................51 

Pertusaria virensica...............................................................52 

Pertusaria xanthodes ............................................................53 

Pertusariaceae ......................................................................51 

Pertusariales..........................................................................51 

Phaeocalicium .......................................................................28 

Phaeocalicium polyporaeum.................................................28 

Phaeographina ......................................................................13 

Phaeographina asteroides ....................................................13 

Phaeographina caesiopruinosa ............................................14 

Phaeographina explicans......................................................14 

Phaeographina quassiaecola................................................14 

Phaeographis ........................................................................14 

Phaeographis dendritica .......................................................15 

Phaeographis erumpens .......................................................15 

Phaeographis haematites .....................................................14 

Phaeographis inusta..............................................................14 

Phaeographis lobata .............................................................15 

Phaeographis multicolor........................................................15 

Pleomassariaceae.................................................................74 

Pleosporales..........................................................................59 

Pleurothelium dissimulans ..................................................110 

Pleurotrema .........................................................................112 

Pleurotrema anisomerum....................................................147 

Pleurotrema epiphyllum ......................................................146 

Pleurotrema inspersum ...............................................123, 144 

Pleurotrema megaspermum................................................147 

Pleurotrema polycarpum .....................................................149 

Pleurotrema polysemum .......................................................99 

Pleurotrema pyrenuloides ...................................................144 

Pleurotrema rockii................................................................149 

Pleurotrema uniseriale ........................................................151 

178 

Phaeographis punctiformis................................................... 15 

Phaeographis sericea........................................................... 14 

Phaeographis subfulgurata .................................................. 15 

Phaeographis subtigrina....................................................... 15 

Phaeopeltosphaeria irregularis .......................................... 100 

Phaeophyscia ....................................................................... 48 

Phaeophyscia cernohorskyi.................................................. 48 

Phaeophyscia chloantha ...................................................... 48 

Phaeophyscia ciliata ............................................................. 48 

Phaeophyscia pusilloides ..................................................... 48 

Phaeophyscia rubropulchra.................................................. 48 

Phyllobatheliaceae.............................................................. 151 

Phyllobathelium................................................................... 152 

Phyllobathelium albolinitum................................................ 155 

Phyllobathelium obtectum .................................................. 158 

Phylloporina obducta .......................................................... 158 

Phylloporina octomera........................................................ 176 

Phylloporis .......................................................................... 152 

Phylloporis multipunctata.................................................... 158 

Phylloporis obducta ............................................................ 158 

Phylloporis phyllogena................................................ 152, 159 

Phylloporis platypoda ......................................................... 159 

Phylloporis viridis ................................................................ 160 

Phyllopsora ........................................................................... 36 

Phyllopsora buettneri var. glauca ......................................... 38 

Phyllopsora buettneri var. munda......................................... 38 

Phyllopsora confusa ............................................................. 36 

Phyllopsora corallina var. corallina................................. 36, 37 

Phyllopsora corallina var. glaucella...................................... 38 

Phyllopsora corallina var. ochroxantha ................................ 38 

Phyllopsora corallina var. rappiana ...................................... 37 

Phyllopsora corallina var. santensis..................................... 38 

Phyllopsora furfuracea.......................................................... 37 

Phyllopsora halei................................................................... 37 

Phyllopsora isidiolyta ............................................................ 37 

Phyllopsora kalbii.................................................................. 37 

Phyllopsora parvifolia var. breviuscula........................... 36, 37 

Phyllopsora parvifolia var. parvifolia..................................... 36 

Phyllopsora parvifoliella........................................................ 36 

Physcia.................................................................................. 49 

Physcia aipolia ...................................................................... 49 

Physcia americana................................................................ 49 

Physcia atrostriata ................................................................ 49 

Physcia crispa....................................................................... 49 

Physcia neogaea .................................................................. 49 

Physcia poncinsii .................................................................. 49 

Physcia pumilior.................................................................... 49 

Physcia sorediosa................................................................. 49 

Physcia undulata................................................................... 49 

Physciaceae.......................................................................... 44 

Physma byrsaeum ................................................................ 28 

Pilocarpaceae ....................................................................... 30 

Pleomassaria ........................................................................ 74 

Pleurotremataceae ............................................................. 112 

Polyblastia dispora................................................................ 87 

Polyblastia fallaciuscula........................................................ 87 

Polyblastia transwaalensis ................................................... 88 

Polyblastiopsis alboatra........................................................ 87 

Polyblastiopsis dispora ......................................................... 87 

Polyblastiopsis fallaciuscula ................................................. 87 

Polyblastiopsis geminella ..................................................... 87 

Polyblastiopsis linearis ....................................................... 117 

Polyblastiopsis obtecta ....................................................... 158 

Polyblastiopsis pyriformis ................................................... 118 

Polyblastiopsis rappii ............................................................ 87

Polyblastiopsis sphaerica....................................................118 

Polyblastiopsis tagananae ..................................................160 

Polycornum..........................................................................175 

Polymeridium.......................................................................118 

Polymeridium albidum .........................................................119 

Polymeridium albocinereum................................................119 

Polymeridium catapastum...................................................118 

Polymeridium contendens...................................................119 

Polymeridium proponens.....................................................119 

Polymeridium quinqueseptatum..........................................119 

Polymeridium subcinereum.................................................119 

Polypyrenula ..........................................................................92 

Porina...................................................................................172 

Porina aenea .......................................................................178 

Porina albicascens ..............................................................154 

Porina amygdalina...............................................................178 

Porina cestrensis.................................................................178 

Porina cestrensis var. platyspora........................................178 

Porina chlorotica..................................................................179 

Porina concamerata ..............................................................80 

Porina elaeidis.....................................................................130 

Porina guaranitica ...............................................................173 

Porina guentheri ..................................................................179 

Porina heterospora..............................................................173 

Porina hibernica...................................................................180 

Porina indutula.....................................................................157 

Porina interspersa ...............................................................130 

Porina leptalea.....................................................................175 

Porina linearis......................................................................180 

Porina mastoidea.................................................................172 

Porina multipunctata............................................................158 

Porina mundula ...................................................................158 

Porina nitidula......................................................................180 

Porina nucula.......................................................................174 

Porina nuculastrum...........................................................174 

Porina octomera ..................................................................176 

Porina olivacea var. microspora..........................................175 

Porina phyllogena................................................................159 

Porina platypoda..................................................................159 

Porina plumbaria ...................................................................83 

Porina rhaphidosperma.......................................................180 

Porina rubentior...................................................................176 

Porina salicina .....................................................................178 

Porina scabrida..................................................................174 

Porina subpungens .............................................................172 

Porina tamarindi ..................................................................130 

Porina tetracerae.................................................................172 

Porina thaxteri .....................................................................180 

Porina vaga..........................................................................130 

Porina viridiseda f. albiseda ................................................126 

Porinopsis............................................................................162 

Porinopsis geminipara.........................................................165 

Pseudoparmelia.....................................................................43 

Pyrenula cuyabensis ...........................................................103 

Pyrenula dissimulans........................................................110 

Pyrenula erumpens .............................................................103 

Pyrenula falsaria..................................................................102 

Pyrenula ferax .....................................................................111 

Pyrenula harrisii...................................................................110 

Pyrenula herrei ......................................................................83 

Pyrenula laetior....................................................................109 

Pyrenula laevigata ...............................................................109 

Pyrenula leucostoma...........................................................102 

Pyrenula macularis..............................................................110 

Pyrenula mamillana.............................................................110 

Pyrenula marginata .............................................................110 

Pyrenula martinicana...........................................................103 

179 

Pseudoparmelia sphaerospora ............................................ 43 

Pseudopyrenula .................................................................. 119 

Pseudopyrenula degenerans ............................................. 117 

Pseudopyrenula endochrysea............................................ 117 

Pseudopyrenula subgregaria ............................................. 119 

Pseudopyrenula subnudata................................................ 119 

Pseudosagedia ................................................................... 176 

Pseudosagedia aenea........................................................ 178 

Pseudosagedia chlorotica .................................................. 179 

Pseudosagedia guentheri................................................... 179 

Pseudosagedia linearis ...................................................... 180 

Pseudosagedia nitidula ...................................................... 180 

Pseudosagedia thaxteri ...................................................... 180 

Punctelia ............................................................................... 43 

Punctelia rudecta .................................................................. 43 

Pyrenidium ............................................................................ 74 

Pyrenidium octosporum...................................................... 162 

Pyrenocollema ...................................................................... 71 

Pyrenocollema atlanticum ................................................. 72 

Pyrenocollema caesium........................................................ 72 

Pyrenocollema ceuthocarpoides ...................................... 72 

Pyrenocollema epigloeum ................................................. 72 

Pyrenocollema farlowii....................................................... 72 

Pyrenocollema prospersellum .............................................. 72 

Pyrenocollema saxicola........................................................ 63 

Pyrenocollema strontianense ............................................... 73 

Pyrenocollema tichothecioides............................................. 72 

Pyrenocollema tremelloides ................................................. 72 

Pyrenocollema zonatum..................................................... 73 

Pyrenographa ..................................................................... 100 

Pyrenographa irregularis ................................................. 100 

Pyrenographa xylographoides............................................ 100 

Pyrenowilmsia....................................................................... 93 

Pyrenula .............................................................................. 101 

Pyrenula acutalis................................................................. 104 

Pyrenula anomala............................................................... 105 

Pyrenula aspistea ............................................................... 106 

Pyrenula astroidea.............................................................. 101 

Pyrenula atrolaminata ...................................................... 107 

Pyrenula caryae................................................................. 107 

Pyrenula cerina ................................................................... 104 

Pyrenula circumrubens....................................................... 109 

Pyrenula citriformis ............................................................. 104 

Pyrenula cocoes ................................................................. 105 

Pyrenula concastroma ..................................................... 108 

Pyrenula concatervans ....................................................... 109 

Pyrenula confinis .............................................................. 109 

Pyrenula confoederata........................................................ 106 

Pyrenula corticata ............................................................... 109 

Pyrenula cruenta................................................................. 109 

Pyrenula cruentata.............................................................. 101 

Pyrenula crystalligera ......................................................... 109 

Pyrenula cubana ................................................................. 103 

Pyrenula mastoidea............................................................ 172 

Pyrenula microcarpa........................................................... 106 

Pyrenula microtheca ........................................................... 103 

Pyrenula mucosa .............................................................. 110 

Pyrenula nitidula ................................................................. 110 

Pyrenula occidentalis.......................................................... 110 

Pyrenula ochraceoflava ...................................................... 101 

Pyrenula ochraceoflavens .................................................. 101 

Pyrenula oleosa .................................................................. 102 

Pyrenula personata............................................................. 103 

Pyrenula pseudobufonia..................................................... 105 

Pyrenula punctella .............................................................. 111 

Pyrenula punctiformis var. cinereopruinosa f. buxicola ....... 79 

Pyrenula punctiformis var. cinereopruinosa f. hederae ....... 79

Pyrenula punctiformis var. cinereopruinosa f. pinicola.........79 

Pyrenula punctiformis var. fallax f. betulae ..........................79 

Pyrenula pyrenuloides.........................................................111 

Pyrenula quassiaecola ........................................................104 

Pyrenula ravenelii ................................................................101 

Pyrenula rubromaculata ......................................................109 

Pyrenula rubrostoma ...........................................................106 

Pyrenula santensis ..............................................................105 

Pyrenula septicollaris ..........................................................103 

Pyrenula subferruginea .......................................................103 

Pyrenula tenuisepta.............................................................106 

Pyrenula thelomorpha .........................................................102 

Pyrenula wetmorei...............................................................103 

Pyrenula wheeleri ................................................................103 

Pyrenula wrightii................................................................111 

Pyrenula zwackhii..................................................................67 

Pyrenulaceae...................................................................74, 91 

Pyrenyllium ............................................................................75 

Pyrgillus ...............................................................................111 

Pyrgillus javanicus.................................................................93 

Pyxine ....................................................................................50 

Pyxine albovirens ..................................................................50 

Pyxine berteriana...................................................................50 

Pyxine caesiopruinosa ..........................................................50 

Pyxine cocoes........................................................................50 

Pyxine coralligera ..................................................................50 

Pyxine eschweileri .................................................................50 

Pyxine sorediata ....................................................................50 

Pyxine subcinerea .................................................................50 

Raciborskiella ......................................................................152 

Relicina ..................................................................................43 

Relicina abstrusa...................................................................43 

Relicina eximbricata ..............................................................43 

Requienella..........................................................................112 

Requienella lichenopsis ......................................................112 

Requienella seminuda.........................................................112 

Requienella subcollapsa ..................................................112 

Requienellaceae..................................................................112 

Rimelia...................................................................................43 

Rimelia reticulata...................................................................43 

Rimelia subisidiosa................................................................43 

Rimeliella ...............................................................................43 

Rimeliella neotropica.............................................................43 

Rimeliella subtinctoria ...........................................................43 

Rinodina.................................................................................50 

Rinodina applanata ...............................................................50 

Rinodina granuligera .............................................................51 

Rinodina lepida......................................................................51 

Sagedia excaecariae...........................................................128 

Sagedia massalongiana........................................................62 

Sagedia nylanderi..................................................................72 

Sagedia nylanderi β. hudsoniana..........................................72 

Sagediomyces .....................................................................152 

Strigula nitidula ....................................................................152 

Strigula obducta ................................................................158 

Strigula obtecta..................................................................158 

Strigula phaea .....................................................................158 

Strigula phyllogena ...................................................152, 159 

Strigula platypoda .............................................................159 

Strigula rostrata.................................................................159 

Strigula smaragdula ....................................................152, 159 

Strigula stigmatella ..............................................................154 

Strigula subelegans.............................................................159 

Strigula subsimplicans.........................................................152 

Strigula sychnogonioides ....................................................159 

Strigula tagananae ............................................................160 

Strigula viridis....................................................................160 

180 

Sagediomyces affinis.......................................................... 152 

Santessoniolichen................................................................. 61 

Sarcinulella ......................................................................... 123 

Sarcinulella banksiae.......................................................... 123 

Sarcographa ......................................................................... 14 

Sarcographa intricans........................................................... 14 

Sarcographa labyrinthica...................................................... 14 

Sarcographa medusulina...................................................... 15 

Sarcographa tricosa.............................................................. 14 

Sarcographia labyrinthica ..................................................... 15 

Schismatomma ....................................................................... 8 

Schismatomma rappii ......................................................... 5, 8 

Schizoxylon berkelyeanum ..................................................... 4 

Sciodothis.............................................................................. 65 

Sclerophyton ........................................................................... 8 

Sclerophyton elegans ............................................................. 8 

Sclerophyton inscriptum ......................................................... 8 

Scleropyrenium ..................................................................... 57 

Secolegiella......................................................................... 162 

Segestria ............................................................................. 175 

Segestria lectissima............................................................ 175 

Segestria leptalea ............................................................. 175 

Segestria mammillosa ........................................................ 176 

Segestria octomera .......................................................... 176 

Segestria rubentior........................................................... 176 

Shanoria.............................................................................. 152 

Shanoria indica ................................................................... 152 

Sphaeria cavernosa.............................................................. 99 

Sphinctrina ............................................................................ 25 

Sphinctrina tubiformis ........................................................... 25 

Sphinctrinaceae .................................................................... 25 

Sphinctrinella ........................................................................ 27 

Splanchnonema.................................................................... 74 

Sporoschizon ........................................................................ 61 

Strigula ................................................................................ 152 

Strigula affinis ..................................................................... 152 

Strigula albicascens ......................................................... 154 

Strigula albolinitum .......................................................... 155 

Strigula bahamensis......................................................... 155 

Strigula bermudana .......................................................... 155 

Strigula cinefaciens.......................................................... 155 

Strigula connivens............................................................ 155 

Strigula elegans .................................................................. 159 

Strigula fracticonidia ........................................................ 156 

Strigula griseonitens ........................................................ 156 

Strigula hypothallina......................................................... 157 

Strigula indutula................................................................ 157 

Strigula jamesii.................................................................... 157 

Strigula laceribracae ........................................................ 158 

Strigula mediterranea ......................................................... 152 

Strigula multipunctata...................................................... 158 

Strigula viridiseda ............................................................... 160 

Strigulaceae ........................................................................ 151 

Strigulomyces ..................................................................... 152 

Strigulomyces elegans ....................................................... 152 

Syngenosorus....................................................................... 65 

Thelenella ........................................................................... 162 

Thelenella brasiliensis ........................................................ 164 

Thelenella cinerascens .................................................... 165 

Thelenella cuyabense......................................................... 165 

Thelenella elegans.............................................................. 117 

Thelenella fugiens............................................................. 165 

Thelenella geminipara ...................................................... 165 

Thelenella harrisii................................................................ 167 

Thelenella hassei................................................................ 164

Thelenella humilis..............................................................165 

Thelenella inductula ............................................................164 

Thelenella luridella ..............................................................166 

Thelenella modesta .....................................................162, 163 

Thelenella obtecta ...............................................................158 

Thelenella pertusariella .......................................................166 

Thelenella rappii ................................................................166 

Thelenella sastreana .........................................................166 

Thelenella sordidula ............................................................164 

Thelenella sychnogonioides ............................................167 

Thelenella trichothelioides ...............................................167 

Thelenella weberi ................................................................164 

Thelenellaceae ....................................................................160 

Thelidium ...............................................................................57 

Thelidium farlowii...................................................................72 

Thelidium juistense..............................................................129 

Thelidium minutulum .............................................................57 

Thelidium nylanderi ...............................................................72 

Thelidium nylanderi var. hudsoniana ....................................72 

Thelomma..............................................................................25 

Thelomma carolinianum........................................................25 

Thelotrema.............................................................................19 

Thelotrema adjectum.............................................................19 

Thelotrema alborosellum.......................................................19 

Thelotrema defectum ............................................................20 

Thelotrema dilatatum.............................................................19 

Thelotrema eximium..............................................................19 

Thelotrema floridense............................................................19 

Thelotrema halei....................................................................19 

Thelotrema lacteum...............................................................20 

Thelotrema lathraeum ...........................................................19 

Thelotrema lepadinum...........................................................20 

Thelotrema leprocarpum .......................................................20 

Thelotrema monospermum ...................................................20 

Thelotrema platycarpoides....................................................20 

Thelotrema porinoides...........................................................19 

Thelotrema santense.............................................................19 

Thelotrema subtile.................................................................19 

Thelotremataceae..................................................................16 

Tomasellia .............................................................................64 

Tomasellia africana .............................................................130 

Tomasellia americana ...........................................................64 

Tomasellia angulosa ...........................................................144 

Tomasellia cubana ................................................................69 

Tomasellia dispora ..............................................................113 

Tomasellia eschweileri ..........................................................69 

Tomasellia esenbeckiana................................................69, 81 

Tomasellia exumana .............................................................69 

Tomasellia lactea...................................................................69 

Tomasellia leucostoma....................................................69, 80 

Tomasellia macularis ..........................................................65 

Tomasellia nigrescens...........................................................98 

Tomasellia queenslandica...................................................130 

Usnea dimorpha ....................................................................44 

Usnea mutabilis.....................................................................43 

Usnea perplectata .................................................................43 

Usnea rubicunda ...................................................................44 

Usnea strigosa.......................................................................43 

Usnea subscabrosa...............................................................44 

Usnea trichodea ....................................................................44 

Verrucaria ..............................................................................58 

Verrucaria*Leiophloea...........................................................75 

Verrucaria aenea.................................................................178 

Verrucaria albescens...........................................................109 

Verrucaria albicascens........................................................154 

Verrucaria albidoatra...........................................................144 

Verrucaria alboatra..............................................................127 

181 

Tomasellia sparsella............................................................. 70 

Trichotheliaceae ................................................................. 168 

Trichotheliales..................................................................... 168 

Trichothelium ...................................................................... 176 

Trichothelium aeneum ............................................. 176, 178 

Trichothelium cestrense .................................................. 178 

Trichothelium chloroticum .............................................. 179 

Trichothelium crocynioides............................................. 179 

Trichothelium guentheri................................................... 179 

Trichothelium horridulum.................................................... 178 

Trichothelium isidiatum ................................................... 180 

Trichothelium lineare........................................................ 180 

Trichothelium nitidulum ................................................... 180 

Trichothelium rhaphidospermum ................................... 180 

Trichothelium thaxteri ...................................................... 180 

Trypetheliaceae .................................................................. 112 

Trypethelium ....................................................................... 119 

Trypethelium aeneum ......................................................... 120 

Trypethelium albidoporum.................................................. 116 

Trypethelium carolinianum ................................................. 116 

Trypethelium degenerans................................................... 117 

Trypethelium eluteriae ........................................................ 119 

Trypethelium eluteriae var. polystomum ............................ 120 

Trypethelium exiguellum....................................................... 87 

Trypethelium feei ................................................................ 117 

Trypethelium ferrugineum................................................... 120 

Trypethelium ferrugineum β. inornatum ............................. 120 

Trypethelium floridanum ..................................................... 120 

Trypethelium karnatakense ............................................. 147 

Trypethelium leprieurii ........................................................ 121 

Trypethelium madreporiforme ............................................ 117 

Trypethelium madreporiforme f. pruinosa .......................... 117 

Trypethelium marginatum................................................... 117 

Trypethelium nitidiusculum......................................... 120, 121 

Trypethelium ochroleucum ......................................... 120, 121 

Trypethelium phlyctaena .................................................... 117 

Trypethelium quassiaecola................................................. 117 

Trypethelium scoria............................................................. 117 

Trypethelium subeluteriae .................................................. 120 

Trypethelium tropicum ........................................................ 120 

Trypethelium variolosum ............................................ 114, 121 

Trypethelium virens............................................................. 120 

Tuckermanopsis.................................................................... 43 

Tuckermanopsis fendleri ...................................................... 43 

Tuckermanopsis viridis ......................................................... 43 

Tylophoron ............................................................................ 25 

Tylophoron moderatum......................................................... 25 

Tylophoron protrudens.......................................................... 25 

Usnea.................................................................................... 43 

Usnea baileyi ........................................................................ 43 

Usnea ceratina...................................................................... 43 

Verrucaria alboatra var. detergens....................................... 80 

Verrucaria albolinita............................................................ 155 

Verrucaria analepta var. americana ................................... 110 

Verrucaria anisomera ......................................................... 150 

Verrucaria antecellens.......................................................... 67 

Verrucaria bermudana........................................................ 155 

Verrucaria biformis.............................................................. 127 

Verrucaria calkinsiana .......................................................... 59 

Verrucaria cerasi................................................................... 79 

Verrucaria cestrensis.......................................................... 178 

Verrucaria chlorotica........................................................... 179 

Verrucaria cinchonae............................................................ 80 

Verrucaria cinefaciens ........................................................ 155 

Verrucaria cinereopruinosa .................................................. 79

Verrucaria circumrubens .....................................................109 

Verrucaria clandestina.........................................................147 

Verrucaria concamerata........................................................80 

Verrucaria confinis...............................................................109 

Verrucaria conformis ...........................................................127 

Verrucaria consobrina .........................................................145 

Verrucaria consobrina var. glaucescens.............................151 

Verrucaria distans................................................................127 

Verrucaria epidermidis var. fallax..........................................79 

Verrucaria epidermidis var. γ. quassiaecola .........................63 

Verrucaria epigloea ...............................................................72 

Verrucaria excellens............................................................146 

Verrucaria fallax.....................................................................79 

Verrucaria geminella .............................................................87 

Verrucaria guentheri............................................................179 

Verrucaria hippocastani ........................................................65 

Verrucaria holopolia ............................................................147 

Verrucaria indutula ..............................................................157 

Verrucaria infernalis ............................................................147 

182 

Verrucaria interspersa ........................................................ 130 

Verrucaria lactea................................................................... 69 

Verrucaria limitans .............................................................. 144 

Verrucaria linearis............................................................... 180 

Verrucaria magnifica........................................................... 148 

Verrucaria magnospora ...................................................... 148 

Verrucaria majuscula............................................................ 82 

Verrucaria malaccitula .......................................................... 82 

Verrucaria mamillaris............................................................ 59 

Verrucaria muralis................................................................. 58 

Verrucaria planiuscula ........................................................ 149 

Verrucaria planorbis.............................................................. 82 

Verrucaria plumbaria ............................................................ 83 

Verrucaria prostans .............................................................. 80 

Verrucaria punctiformis......................................................... 63 

Verrucaria rhyponta .............................................................. 63 

Verrucaria riddleana ........................................................... 59 

Verrucaria rubentior............................................................ 176 

Verrucaria socialis .............................................................. 161 

Verrucaria stigmatella var. lactea ......................................... 69 

Verrucaria subantecellens.................................................... 67 

Verrucaria subatomaria ...................................................... 150 

Verrucaria subbiformis........................................................ 150 

Verrucaria sublactea............................................................. 88 

Verrucaria subnectenda ..................................................... 150 

Verrucaria subnexa............................................................. 150 

Verrucaria subprostans ...................................................... 130 

Verrucaria subpunctiformis................................................... 62 

Verrucaria tagananae ......................................................... 160 

Verrucaria tamarindi ........................................................... 130 

Verrucaria terminata ........................................................... 130 

Verrucaria tetracerae.......................................................... 172 

Verrucaria thelena var. albidior .......................................... 143 

Verrucaria vaga................................................................... 130 

Verrucaria viridiseda f. albiseda ......................................... 126 

Verrucariaceae...................................................................... 56 

Verrucariales......................................................................... 56 

Vulpicida................................................................................ 44 

Vulpicida viridis ..................................................................... 44 

Xanthoparmelia..................................................................... 44 

Xanthoparmelia conspersa................................................... 44 

Xanthopyrenia....................................................................... 72 

Xanthopyreniaceae............................................................... 70 

Zamenhofia ......................................................................... 176 

Zignoella nyssaegena......................................................... 128 

Zwackhiomyces..................................................................... 73 

Zwackhiomyces arenicola ................................................. 73

PART 1

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