PART 1
PART 1
PART 1
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<strong>PART</strong> 1<br />
INTRODUCTION<br />
Sadly this introduction is a litany of apologies. This second part appears almost five years to the day<br />
after the publication of the first, "Some Florida Lichens". Its publication is occasioned by the Third Tuckerman<br />
Workshop in Gainesville, Florida as the first was by the meeting of The American Bryological and<br />
Lichenological Society in Wakulla Springs, Florida. Obviously I had hoped to have a second part finished<br />
much earlier but without an absolute deadline such as the Tuckerman Workshop, procrastination rules. I<br />
badly underestimated the time necessary to produce the manuscript, deal with accumulated specimens,<br />
changes and errors. As a result much has been left unfinished. The presentation is only a skeleton intended<br />
to document Florida's lichen diversity, provide names by which it can be manipulated and encourage further<br />
investigations. It is definitely not "user friendly", especially for nonprofessionals. Descriptions, bibliography,<br />
illustrations and maps are not included although a glimpse of the future is included in the form of computer<br />
generated maps for Calicium. Human limitations have also led me to confine the scope primarily to materials<br />
held by The New York Botanical Garden. Study of specimens from other herbaria and new collections will<br />
make this preliminary survey obsolete within a matter of months. (A prime example is the Graphidaceae<br />
where the species curve shows no sign of leveling off.) This will be exacerbated by the rapidly changing<br />
generic concepts in crustose lichens resulting from release from the constraints of artificial systems and the<br />
introduction of new characters and the techniques to study them.<br />
"More Florida Lichens" consists of two parts. The first is mostly the keys from "Some Florida Lichens"<br />
with corrections and a few additions plus a few new family treatments. It is assumed that most users of "More<br />
Florida Lichens" have the first part and therefore, in order to save space, the reprinted keys are in 8pt type with<br />
additions and corrections in bold face. The second part is an expanded treatment of the pyrenolichens. It is<br />
given a separate section and called a" tour" since it is not confined to Florida taxa and like a tour covers<br />
whatever seems relevant, hitting the high points and avoiding the slums. It is essentially a complete treatment<br />
of the corticolous pyrenolichens for eastern North America. However, for some families that are special<br />
favorites, the digressions are global.<br />
As always I wish to thank my companion, Bill Buck, for moral support as well as editing and indexing.<br />
Dana Griffin, III is thanked for arranging many pleasant field trips and making numerous interesting specimens<br />
available. I am grateful to the Florida Department of Natural Resources for permission to collect in a number<br />
of state parks and preserves.<br />
ARTHONIALES<br />
1
OPEGRAPHACEAE Stizenb.<br />
KEY TO GENERA<br />
1. Exciple or hypothecium dark; exciple not byssoid; hypothecium, if dark, usually<br />
confluent with exciple ................................................................................................................................ 2<br />
2. Ascomata not aggregated, not in raised stroma-like structures............................................................3<br />
3. Ascoma elongated, lirelliform; ascospores with a gelatinous sheath............................................... 4<br />
4. Ascoma not pruinose; ascus of vulgata-type ............................................................. Opegrapha<br />
4. Ascoma pruinose; ascus of grumulosa-type ........................................................ [Lecanographa]<br />
3. Ascoma rounded, apothecioid; ascospores with or without a sheath .............................................. 5<br />
5. Asci and ascospores not fragile; paraphysoids tightly interconnected,<br />
not separating easily; ascus abietina- or grumulosa-type ............................................................ 6<br />
6. Ascus abietina-type, endoascus strongly hemiamyloid with conspicuous<br />
apical ring; ascospores without sheath ................................................................................... 7<br />
7. Ascoma not pruinose or disk with bright colored pruina<br />
[orange, yellow or greenish], KOH+ yellow or purple; exciple<br />
carbonaceous, hyphae not distinguishable in KOH............................................Cresponea<br />
7. Ascoma pruinose; pruina the same color as thallus, KOH-;<br />
exciple dark brown, hyphae in outer part distinguishable in KOH ......................Lecanactis<br />
6. Ascus grumulosa-type, endoascus weakly hemiamyloid with apical<br />
ring little or not differentiated; ascospores with a sheath ................................. [Lecanographa]<br />
5. Asci and ascospores fragile; paraphysoids dichotomously branched,<br />
easily separating; ascus of Bactrospora-type...........................................................Bactrospora<br />
2. Ascomata aggregated or in raised stroma-like structures..................................................................... 8<br />
8. Ascoma perithecioid, in weakly raised stroma-like structures;<br />
ascospores 4-celled, 23-27 × 2-3 µm; roccellic acid;<br />
Duval and Putnam counties ......................................................................Chiodecton malmei Thor<br />
8. Ascomata with exposed disk, not perithecioid; PD+ or C+<br />
(protocetraric acid, psoromic acid or erythrin ..................................................................................... 9<br />
9. Thallus C-, PD+ yellow or red orange .......................................................................................... 10<br />
10. Thallus PD+ red orange (protocetraric acid); ascomata asteriform,<br />
in raised "stroma" which is constricted at base; disk brown,<br />
white pruinose; ascospores 4-celled, 50-70 × 4 µm;<br />
peninsular Florida............................Syncesia [treated in 1990 as Chiodecton sp. Buck 3254]<br />
10. Thallus PD+ yeliow (psoromic acid); ascomata round,<br />
in rows .................................................................................Enterographa quassiaecola Fée<br />
9. Thallus C+ red (erythrin), PD-; ascomata asteriform in raised "stroma"<br />
which is constricted at base; disk light to dark brown, white pruinose;<br />
ascospores 4-celled, tapered, 50-70 × 4-5 µm; Duval County ........................................................<br />
2
..............................................................Syncesia [treated in 1990 as Chiodecton perplexum Nyl.]<br />
1. Exciple not evident or byssoid (when pruina dissolved in KOH); hypothecium colorless<br />
or light brown and small, confined to base of ascoma ............................................................................ 11<br />
11. Hypothecium light brown, small, only at base of ascoma.............................................................. 12<br />
12. Exciple not evident; epithecium not byssoid; ascomata immersed, asteriform;<br />
disk exposed, orange or red brown; ascospores 4-celled, 15-20 × 5 µm;<br />
psoromic acid; Duval, Leon and St. Lucie counties ....................................................................<br />
................................................................................ Sclerophyton inscriptum (Nyl.) Müll. Arg<br />
12. Ascomata densely pruinose; exciple and epithecium pale, byssoid<br />
hypothecium brown; ascospores 4-celled, ± clavate, ca. 24-31 × 6 µm;<br />
Bay County.......................................................................................... Genus? sp. Harris 35756<br />
11. Hypothecium colorless ................................................................................Enterographa/Mazosia<br />
BACTROSPORA Massal.<br />
Since "Some Florida Lichens" was published Egea and Torrente have revised the genus<br />
(Lichenologist 25: 211-255. 1993). Their study has resulted in the reduction of one of the Florida species to<br />
synonymy and the addition of one species to the mycota.<br />
1. Ascospores narrow, 3-4 µm wide.............................................................................................................. 2<br />
2. Ascospores long, 35-130 µm ................................................................................................................ 3<br />
3. Ascospores 12-26-celled; exciple or exciple and epithecium containing<br />
small granules; tissues IKI-, KOH/IKI + bluish................................................................................... 4<br />
4. Ascomatal margin smooth; ascospores 12-20-septate,<br />
47-85(-90) × 2-3(-3.5) µm; Collier, Duval, Lee, Levy and<br />
Polk counties ........................................................................ B. myriadea (Fée) Egea & Torrente<br />
4. Ascomatal margin with sparse, small teeth; ascospores<br />
70-130 × 2-2.5(-3) µm; Dade, Levy and Monroe<br />
counties ........................................................................ B. denticulata (Vainio) Egea & Torrente<br />
3. Ascospores 10-celled, 35-50 × 3 µm; exciple and epithecium lacking<br />
granules; parts of exciple and base of hymenium IKI+ bluish,<br />
KOH/IKI+ deep blue; thallus IKI+ bluish; common, south to<br />
Sarasota County......................................................................................B. mesospora R. C. Harris<br />
2. Ascospores short, (4-)6-celled, 22-27 × 3-4 µm; on lignum or bark<br />
of Taxodium; Franklin, Liberty and Wakulla counties...................................B. brevispora R. C. Harris<br />
1. Ascospores broad, 18-20-celled, 80-90 × 7-10 µm; Calhoun,<br />
Duval and Nassau counties........................................................................... B. macrospora R. C. Harris<br />
NOTES<br />
Bactrospora denticulata (Vainio) Egea & Torrente<br />
3
Bactrospora integrispora Seaver, Mycologia 16: 8. 1924. Type. VIRGIN ISLANDS. St. Thomas: on<br />
decorticated wood, 10 Mar 1923, Seaver 754 (NY, holotype), syn. nov.<br />
Seaver's name, appearing in a mycological journal, was not picked up by Zahlbruckner and thus not<br />
included by Egea & Torrente. (Also omitted, Seaver's report of Bactrospora dryina (Ach.) Massal. from North<br />
Dakota (Mycologia 1: 112. 1909.) is Schizoxylon berkelyeanum (Dur. & Lév.) Fuckel s. lat.) The Aptroot<br />
specimen from Key Largo was incorrectly assigned to Dade County instead of Monroe County in Egea and<br />
Torrente (1993).<br />
Bactrospora macrospora R. C. Harris<br />
Henssen in Henssen and Thor (1990) transferred this species to Melampylidium Stirton. I am<br />
uncomfortable with this on several grounds. First, the decision is based on material from North Carolina which<br />
the authors admit differs from my description of B. macrosporum. Second, I am cited for pointing out the<br />
different ascospore type. This is true but this was before Egea and Torrente's (1993) treatment of<br />
Bactrospora and I was simply ignorant of the range of ascospore variation in the genus. I did comment on the<br />
identity of the ascomatal type within Bactrospora. Third, Egea and Torrente (1995) present evidence for<br />
synonymizing Melampylidium with Bactrospora. Until more convincing evidence for recognizing<br />
Melampylidium and B. macrospora as a member of the genus, I prefer to retain the species in Bactrospora.<br />
Bactrospora nematospora R. C. Harris = B. myriadea<br />
CHIODECTON Ach.<br />
Only a single species, C. malmei Thor, is in North America, and it is diagnosed in the key to genera.<br />
Two other species C. perplexum and sp. Buck 3254 would seem to belong in Syncesia Taylor (Tehler, 1993)<br />
but there is as yet no treatment of the species.<br />
CRESPONEA Egea & Torrente<br />
This genus was recently separated from Lecanactis and revised by Egea and Torrente (Mycotaxon<br />
48: 301-331. 1993). They reported C. proximata from Florida.<br />
1. Ascospores 3-4(-5)-septate, 15-22(-24) × (4-)4.5-5.5 µm; margin black, raised,<br />
cracked; disk chartreuse pruinose; Alachua County........................... C. flava (Vainio) Egea & Torrente<br />
1. Ascospores 5-7(-8)-septate, 25-38(-40) × 5-7(-7.5) µm; margin black, thick,<br />
slightly raised, ± smooth; disk chartreuse pruinose; Monroe County (Keys)...............................................<br />
....................................................................................................... C. proximata (Nyl.) Egea & Torrente<br />
NOTES<br />
Cresponea flava (Vainio) Egea & Torrente<br />
This species is close to C. chloroconia (Tuck.) Egea & Torrente. They annotated a Calkins collection,<br />
purportedly from Florida, as C. chloroconia but did not cite it in the publication. The Cresponea is associated<br />
with sterile Chaenotheca furfuracea (L.) Tibell and is surely not from Florida.<br />
Cresponea flava is new to North America. FLORIDA. Alachua County: Paynes Prairie State Preserve,<br />
vicinity of Alachua Sink, 29°37'N, 82°18'W, mixed hardwoods over limestone, 1 Dec 1992, Harris 29432 (NY).<br />
ENTEROGRAPHA Fée & MAZOSIA Massal.<br />
4
1. Ascomata immersed or emergent; not resembling Byssoloma; margin not white .................................... 2<br />
2. Growing on bark .................................................................................................................................... 3<br />
3. Ascomata immersed rounded or lirelline.......................................................................................... 4<br />
4. Ascospores 8-celled...................................................................................................................... 5<br />
5. Ascomata rounded, arranged in short rows or irregular groups,<br />
surrounding area somewhat raised in age; psoromic acid;<br />
ascospores 34-37 × 5-6 µm, with thick gelatinous sheath;<br />
Monroe County (Keys)...............................................................................E. quassiaecola Fée<br />
5. Ascomata solitary, immersed, with slightly raised margin, lirelliform<br />
to weakly asteriform; disk yellowish; ascospores 26-35 × 4.5 µm;<br />
TLC not done; Collier County ....................................................Enterographa sp. Buck 22882<br />
4. Ascospores 4-celled; ascomata scattered; no substances;<br />
throughout Florida ................................................................ Mazosia ocellata (Nyl.) R. C. Harris<br />
3. Ascomata emergent, Lecanora-like or weakly lirelline, with distinct thalline margin;<br />
disk pale yellowish to brown, weakly pruinose; ascospores 7-8-celled, median cells<br />
occasionally somewhat swollen, 30-45 × 3-4 µm; psoromicacid; Franklin, Gulf and<br />
Okaloosa counties................................................................................E. lecanoroides R. C. Harris<br />
2. Growing or limestone; thallus thick, pale buff brown; ascomata immersed;<br />
disk dark, rounded; ascospores 20-24 × 5-5.5 µm; Monroe County<br />
(Key West) ........................................................................................... Enterographa sp. Thaxter 415<br />
1. Ascomata minute, resembling Byssoloma; margin white; disk brown,<br />
white pruinose; ascospores 6(-7)-celled,18-22 × 3-3.5 µm; Seminole<br />
County ...........................................................Enterographa [Schismatomma rappii (Zahlbr.) R C. Harris]<br />
LECANACTIS Fr.<br />
1. Ascospores 6-8-celled, 30-40 × 3-4 µm; ascomata gray brown, white pruinose,<br />
strongly constricted at base, round, irregular or slightly elongate; schizopeltic acid;<br />
Duval County.......................................................................................................L. epileuca (Nyl.) Tehler<br />
1. Ascospores 4-celled, 20-24 × 4 µm; ascomata as above; schizopeltic acid;<br />
Duval County................................................................................................... Lecanactis sp. Calkins 52<br />
NOTES<br />
Lecanactis subattingens (Nyl.) R. C. Harris<br />
= L. epileuca (Nyl.) Tehler. See Egea & Torrente (Biblioth. Lichenol. 54: 1-205. 1994).<br />
OPEGRAPHA Ach.<br />
Although badly in need of revision, the key to Opegrapha is merely reprinted from 1990.<br />
1. Ascospores 4-celled.................................................................................................................................. 2<br />
5
2. Lirellae black or white pruinose, not pigmented .................................................................................... 3<br />
3. Lirellae sessile, black, shiny, not pruinose .......................................................................................... 4<br />
4. Ascospores rounded at one or both ends ....................................................................................... 5<br />
5. Lirellae smaller, short, fusiform or once branched, rarely asteriform ..........................................6<br />
6. Hymenium not inspersed; ascospores 13-15 × 4-5 µm;<br />
south to Manatee County .....................................................................................O. atra Pers.<br />
6. Hymenium inspersed; disk exposed; ascospores<br />
13-15 × 4-4.5 µm; Duval County ................................................. Opegrapha sp. Calkins 604<br />
5. Lirellae coarse, elongate, branched; ascospores 13-15 × 4-5 µm;<br />
Collier, Dade and Duval counties .............................................................. O. candida Müll. Arg.<br />
4. Ascospores tapering at both ends, 18-19 ×<br />
4-4.5 µm; lirellae small; disk not exposed;<br />
Okeechobee and Polk counties ...........................................................Opegrapha sp. Britton 819<br />
3. Lirellae initially immersed in bark of Taxodium; disk a strongly flexuose<br />
slit becoming quite broadly exposed; margin thick, black or densely white<br />
pruinose; ascospores 25-28 × 4-5 µm, cells ± equal; Franklin and Liberty<br />
counties ......................................................................................................... O. cypressi R. C. Harris<br />
2. Lirellae encrusted with bright orange pigment, short, straight;<br />
ascospores 17-20 × 4 µm; Collier County ................................................... O. aurantiaca B. de Lesd.<br />
1. Ascospores 6-14-celled............................................................................................................................. 7<br />
7. Growing on bark or wood ...................................................................................................................... 8<br />
8. Disk not covered by dense white layer .............................................................................................. 9<br />
9. Disk broadly expanded, red brown.............................................................................................. 10<br />
10. Ascospores 5-8-celled, 16-20 × 4-5 µm; lirellae long, flexuose,<br />
branched; margin not carbonized below hymenium; south to<br />
Polk County ....................................................................................... O. longissima Müll. Arg.<br />
10. Ascospores 9-12-celled, 38-50 × 4 µm; lirellae short, orbicular<br />
or elongate and strongly flexuose; margin carbonized below<br />
hymenium; Dade County...................................................................Opegrapha sp. Britton 2<br />
9. Disk hidden or if exposed, then black ......................................................................................... 11<br />
11. Ascospores 6-celled; lirellae fusiform or once branched ...................................................... 12<br />
12. Ascospores narrow, 3-4 µm wide; disk hidden................................................................. 13<br />
13. Thallus whitish, continuous or rimose,<br />
sometimes scanty; ascospores 24-30 × 3-4 µm;<br />
south to Polk County ............................................................................ O. vulgata Ach.<br />
13. Thallus white, thick, matte, almost byssoid;<br />
6
ascospores 18-24 × 3 µm; Louisiana,<br />
not yet found in Florida...........................................................[O. leucoplaca Müll. Arg.]<br />
12. Ascospores broader, 4.5-6 µm wide; disk becoming exposed, black .............................. 14<br />
14. Hymenium not inspersed; ascospores18-20 × 5-6 µm;<br />
Duval County........................................................................................... O. varia Pers.<br />
14. Hymenium inspersed; ascospores 20-27 × 4.5-5 µm;<br />
Southwest Florida, Mar 1878, Smith & Austin 82.............................Opegrapha sp. 82<br />
11. Ascospores 8-14-celled......................................................................................................... 15<br />
15. Ascospores narrow, 3-4 µm wide.................................................................................... 16<br />
16. Ascospores 12-14-celled, 30-55 × 3-4 µm; thallus dark;<br />
disk hidden; lirellae fusiform or once branched; south to<br />
Polk County ..................................................... O. viridis (Ach.) Behlen & Desberger<br />
16. Ascospores 8-celled, 32-43 × 3 µm; thallus pale;<br />
disk hidden; lirellae branched, flaring out at base;<br />
Dade and Monroe counties ................................................ Opegrapha sp. Small 471<br />
15. Ascospores broader, 5-7 µm wide .................................................................................. 17<br />
17. Hymenium not inspersed; disk becoming exposed;<br />
lirellae small, fusiform to linear, rarely once branched;<br />
ascospores 7-8-celled, 35-40 × 5-7 µm; Dade County..................................................<br />
.........................................................................................Opegrapha sp. Britton 799<br />
17. Hymenium inspersed; disk hidden; lirellae coarse,<br />
simple to weakly asteriform; ascospores 7-8-celled,<br />
40-45 × 5-6.5 µm; Collier County .....................................Opegrapha sp. Britton 652<br />
8. Disk becoming broadly expanded, covered by a dense layer of white<br />
material; margin black, usually lacking at base; lirellae long, flexuose,<br />
branched; ascospores rarely produced?, (4-)6-celled, ca. 24 × 5 µm;<br />
Duval, Polk and Putnam counties ........................................................................... O. astraea Tuck.<br />
7. Growing on bamboo culms or rock ...................................................................................................... 18<br />
18. Growing on bamboo culms (normally a foliicolous species);<br />
ascospores narrow, 4-6-celled, 18-25 × 3-4 µm; Louisiana,<br />
not seen from Florida ..............................................................................................[O. filicina Mont.]<br />
18. Growing on limestone; lirellae sessile, fusiform to linear,<br />
not branched; ascospores 5-6-celled, 20-22 × 4 µm; Keys................................................................<br />
..............................................................................................................Opegrapha sp. Britton 869<br />
SCHISMATOMMA Massal.<br />
Tehler (1993) has removed S. rappii, the only Florida species, from Schismatomma s. str. and<br />
suggested that it be placed provisionally in Enterographa s. lat.<br />
7
SCLEROPHYTON Eschw.<br />
Only the single species, S. inscriptum (Nyl.) Müll. Arg., is in Florida. Egea & Torrente (Bryologist 98:<br />
207-217. 1995) have revised Sclerophyton s. lat. for North America. They provide excellent illustrations. In<br />
1990 I suggested that Sclerophyton should be restricted to the hyaline spored species with dendroid,<br />
immersed ascomata and I am still of this opinion. Egea and Torrente retained the old, broad definition,<br />
recognizing that there are two distinct groups in the genus so constituted, but provided no reasons as to why<br />
they should be joined in a single genus. Superficially S. inscriptum (North America) and S. elegans Eschw.<br />
(Brazil) are identical and both contain psoromic acid. My first thought was to synonymize them. Egea and<br />
Torrente suggested that they are conspecific but did not formally make the synonymy. However, examination<br />
of an isotype of S. elegans (H-Nyl 4583) revealed that the ascospores are slightly smaller (13-14.5 × 4-5 µm)<br />
than in S. inscriptum, (15-20 × 5 µm). Pending study of more material, I agree the two should be kept<br />
separate.<br />
GRAPHIDALES<br />
GRAPHIDACEAE Dumortier<br />
I had hoped to completely revise this family. However, I have had to be content with a few minor additions and corrections. I<br />
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<br />
speciose families among the lichen-forming fungi in Florida.<br />
KEY TO GENERA<br />
1. Ascomata immersed to sessile, mostly elongated with a narrow, often slit-like disk;<br />
not aggregated in a pseudostroma ......................................................................................................................................................... 2<br />
2. Spores colorless................................................................................................................................................................................... 3<br />
3. Spores transversely septate...................................................................................................................................................Graphis<br />
3. Spores submuriform to densely muriform........................................................................................................................... Graphina<br />
2. Spores brown ....................................................................................................................................................................................... 4<br />
4. Spores transversely septate....................................................................................................................................... Phaeographis<br />
4. Spores submuriform to densely muriform................................................................................................................ Phaeographina<br />
1. Ascomata with brown disk, cup-like or aggregated in a pseudostroma................................................................................................. 6<br />
5. Ascomata cuplike with a thick raised margin; disk brown; hymenium not inspersed;<br />
ascospores colorless to weakly tinted, muriform with 9-11 longitudinal cells,<br />
33-48 × 13-16 µm; no substances; throughout Florida......................................................Gyrostomum scyphuliferum (Ach.) Nyl.<br />
5. Ascomata aggregated in a pseudostroma.......................................................................................................................................... 6<br />
6. Spores muriform, colorless becoming brownish in age, 35-50 × 8-9 µm;<br />
pseudostroma well developed, usually gray pruinose; hymenium not inspersed;<br />
no substances; throughout Florida......................................................................................................... Glyphis cicatricosa Ach.<br />
6. Spores transversely 4-6-celled, brown; pseudostroma poorly developed,<br />
usually whitish............................................................................................................................................................. Sarcographa<br />
GLYPHIS Ach.<br />
Only a single species in North America. Redinger (1934) in his treatment for Brazil synonymized the names which also appear in<br />
the North American list, choosing Glyphis cicatricosa Ach. from among the competing synonyms. See generic key for diagnosis.<br />
GRAPHINA Müll. Arg.<br />
1. Lirellae not carbonized ............................................................................................................................................................................ 2<br />
8
2. Ascospores 1(-2)/ascus....................................................................................................................................................................... 3<br />
3. Norstictic acid present; ascospores 80-115 × 24-38 µm;<br />
Alachua, Duval, Levy and Putnam counties ........................................................................................................Graphina n. sp.<br />
3. No lichen substances detected;thallus rough, with a lumpy, scurfy look;<br />
lirellae crowded, short and irregular; disk often exposed; ascospores<br />
85-105 × 20-28 µm; south to Marion County ................................................................................................ G. cypressi Müll. Arg.<br />
2. Ascospores 2-8/ascus.......................................................................................................................................................................... 4<br />
4. Thallus UV-........................................................................................................................................................................................ 5<br />
5. Ascospores longer than 40 µm and/or wider than 20 µm............................................................................................................. 6<br />
6. Lichen substances present (usually PD+) ................................................................................................................................. 7<br />
7. Protocetraric acid; thallus pale gray; lirellae white, emergent; ascospores<br />
2-4/ascus, 110-140 × 15-17 µm; south to Lake County.......................................................G. abaphoides (Nyl.) Müll. Arg.<br />
7. Norstictic acid or stictic acid satellites; ascospores at least initially 8/ascus ....................................................................... 8<br />
8. Stictic acid satellites; lirellae paler than thallus ................................................................................................................. 9<br />
9. Hypostictic and hyposalazinic acids; thallus dull, pale green gray; lirellae<br />
white, immersed, elongate and branched; ascospores 40-70 × 13-17 µm;<br />
north to Lake Okeechobee........................................................................................... G. virginea (Eschw.) Müll. Arg.<br />
9. Constictic acid; lirellae "mealy" due to multiple crumbling layers of exciple;<br />
ascospores 35-78 × 18-36 µm [2-4/ascus according to Wirth & Hale<br />
but U.S. material initially 8/ascus]; Seminole County ....................................................... G. peplophora Wirth & Hale<br />
8. Norstictic acid; lirellae concolorous with thallus, crowded, irregular;<br />
ascospores 60-85 × 13-17 µm; Duval and Levy counties.........................................G. subvirginalis (Nyl.) Müll. Arg.<br />
6. Lichen substances absent; lirellae crowded, irregular with white crumbling exciple;<br />
ascospores IKI-, 35-78 × 18-36 µm; Seminole County............................................................... G. leuconephela auct. Amer.<br />
5. Ascospores shorter than 40 µm and/or narrower than 20 µm.................................................................................................... 10<br />
10. Ascospores 1-2 × 4(-6)-celled, 15-20 × 7-9 µm .................................................................................................................... 11<br />
11. Lirellae fissurine; disk not exposed; no substances;<br />
peninsular Florida..............................................................................................................G. incrustans (Fée) Müll. Arg.<br />
11. Lirellae long, regularly branched; disk exposed, reddish; norstictic acid;<br />
Collier County ................................................................................................................ G. floridana (Tuck.) R. C. Harris<br />
10. Ascospores more than 1-2 × 4-celled, over 20 µm long....................................................................................................... 12<br />
12. Thallus PD- (no subsances)............................................................................................................................................ 13<br />
13. Exciple well developed, yellow to orange, darker in KOH;<br />
lirellae raised, elongate, flexuose; ascospores 2-3 × 6-8-celled,<br />
22-32 × 10-11 µm; Duval County ........................................................................................ G. scolecitis (Tuck.) Fink<br />
13. Exciple not evident; lirellae short,crowded; disk becoming exposed, white<br />
pruinose; ascospores 2-4 × 6-8-celled, 22-24 × 7-9 µm; Marion County....................................Graphina sp. 23599<br />
12. Thallus and lirellae PD+ yellow (psoromic acid); ascospores elliptical,<br />
IKI+ dark violet, 20-30(-40) × 13-19 µm; often sterile but combination<br />
of fissurine lirellae and psoromic acid is diagnostic; south to Lake<br />
County.......................................................................................................................G. columbina (Tuck.) Wirth & Hale<br />
4. Thallus UV+ yellow (lichexanthone); lirellae imnersed; disk pale red brown;<br />
margin white, patchily developed; ascospores 35-50 × 12-15 µm, found only<br />
in Liberty County specimen, sterile from Duval and Wakulla counties but could be<br />
Graphis "anguinoides"....................................................................................................................... G. anguina (Mont.) Müll. Arg.<br />
1. Lirellae carbonized, at least patchily so or at apices of exciple............................................................................................................ 14<br />
9
14. Ascospores 2-8/ascus...................................................................................................................................................................... 15<br />
15. Exciple carbonized to base of hymenium ................................................................................................................................... 16<br />
16. Exciple not striate ................................................................................................................................................................... 17<br />
17. Exciple covered with thin white layer; salazinic acid; ascospores 8/ascus,<br />
1-2 × 4-5-celled, 12-16 × 5-8 µm; Dade County ................................................................ G. marescens (Fée) Müll. Arg.<br />
17. Exciple naked, black; lirellae elliptical toweakly elongate with broad<br />
red brown disk; ascospores 2-4/ascus, 35-50 × 17-25 µm;<br />
no substances; Lake County..................................................................................................................Graphina sp. 1271<br />
16. Exciple becoming striate, bleeding purplish in KOH; norstictic acid;<br />
ascospores 2-4 × 6-8-celled, 20-35 × 10-15 µm; Dade County......................................................Graphina sp. Britton 680<br />
15. Exciple only carbonized at apices, becoming striate.................................................................................................................. 18<br />
18. Ascospores 2-4/ascus, 30-50 × 20-27 µm; norstictic acid;<br />
exciple carbonized almost to base but inner part yellowish;<br />
Florida, without locality ...............................................................................................G. antillarum (Vainio) Zahlbr.<br />
18. Ascospores 8/ascus, 30-40 × 15-17 µm; stictic acid; Louisiana,<br />
not yet known from Florida ...................................................................................... [G. parilis (Krempelh.) Müll. Arg.]<br />
14. Ascospores 1/ascus ......................................................................................................................................................................... 19<br />
19. No substances; ascospores 65-90 × 20-30 µm; lirellae semi-immersed;<br />
disk narrow, not pruinose; margin black, shiny, not striate, open below;<br />
throughout Florida ................................................................................................................................G. xylophaga R. C. Harris<br />
19. Protocetraric acid; ascospores 120-165 × 35-40 µm; lirellae immersed,<br />
disk broad, whitish pruinose; margin pruinose, often only patchily carbonized;<br />
Collier County ....................................................................................................................................G. platyleuca (Nyl.) Zahlbr.<br />
NOTES<br />
Graphina adscribens Nyl.) Müll. Arg.<br />
I have now examined the type of this species (H-Nyl 6824) and it does not match anything in North America. The Florida<br />
collection in Nylander's herbarium under this name is Graphina scolecitis. Thus Graphina adscribens can be excluded from the<br />
North American mycota with confidence.<br />
Graphina columbina (Tuck.) Wirth & Hale<br />
The lead for this species was inadvertenly omitted from "Some Florida Lichens".<br />
Graphina platyleuca (Nyl.) Zahlbr.<br />
New to North America. FLORIDA. Collier County: Collier-Seminole State Park, just W of boat basin, 25°59'N, 81°35'W,<br />
mangroves at edge of salt marsh, 7 Dec 1992, Harris 30065 (NY).<br />
Graphina subvirginalis (Nyl.) Müll. Arg.<br />
I was badly confused in 1990, mixing two obviously distinct taxa. Graphina subvirginalis has a variable number of<br />
ascospores, 2-8/ascus (type, H-Nyl 6851) and another undescribed species has only 1/ascus. I have not had time to re-examine<br />
material but I presume that Graphina intertexta (Müll. Arg.) R. C. Harris will prove to be a synonym of G. subvirginalis.<br />
GRAPHIS Adanson<br />
1. Exciple carbonized; spores 4-celled only in G. afzelii............................................................................................................................. 2<br />
2. Margin not striate; hymenium inspersed or not ................................................................................................................................... 3<br />
3. Ascomata sessile, covered with thin white layer; exciple well developed below hymenium .......................................................... 4<br />
4. Lirellae C+ red (lecanoric acid); ascospores 4-celled, 16-23 × 6-9 µm;<br />
common, throughout the state .................................................................................................................................G. afzelii Ach.<br />
4. Lirellae and thallus UV+ yellow (lichexanthone), PD+ red (protocetraric acid);<br />
ascospores ca. 22-23-celled, 60-95 × 7-9 µm; Highlands County .....................................................................G. candidata Nyl<br />
10
3. Ascomata immersed or sessile, black, not covered by a white layer.............................................................................................. 5<br />
5. Epihymenium colorless, KOH- ..................................................................................................................................................... 6<br />
6. Exciple lacking below hymenium ............................................................................................................................................. 7<br />
7. Lichen substances absent.................................................................................................................................................... 8<br />
8. Ascospores 8-11-celled, 25-40 µm long .......................................................................................................................... 9<br />
9. Disk not exposed; exciple carbonized to base of hymenium;<br />
hymenium not inspersed; ascospores 8-10-celled, 25-40 × 7-9 µm;<br />
Columbia, Duval and Lake counties ..................................................................................................... G. lineola Ach.<br />
9. Disk becoming exposed, whitish or bluish pruinose; exciple<br />
mostly not carbonized to base; hymenium not inspersed;<br />
ascospores 8-11-celled, 30-40 × 7-9 µm; Duval County ............................................................... G. scripta (L.) Ach.<br />
8. Ascospores 11-13-celled, 50-70 × 8-11 µm; hymenium inspersed;<br />
Duval, Franklin and Highlands counties .............................................................................................. G. pavoniana Fée<br />
7. Norstictic, stictic or protocetraric acids............................................................................................................................... 10<br />
10. Norstictic acid only....................................................................................................................................................... 11<br />
11. Lirellae flexuose, branching, sessile to semi-immersed;<br />
hymenium not inspersed; ascospores 6-8-celled, 18-35 × 5-9 µm;<br />
peninsular Florida............................................................................................................................ G. librata Knight<br />
11. Lirellae short, straight, immersed; hymenium inspersed;<br />
ascospores 8(-9)-celled, 30-35 × 8-9 µm; Dade County..................................................... Graphis sp. Britton 664<br />
10. Stictic acid (with or without norstictic acid) or protocetraric acid.................................................................................. 12<br />
12. Lirellae immersed; margin and disk white pruinose; hymenium<br />
not inspersed; ascospores 8-11-celled, 25-35 × 7-8 µm; stictic acid<br />
or protocetraric acid; Dade and Palm Beach counties ....................................................... G. caesiella Vainio s. lat<br />
12. Lirellae semi-immersed, black; margin and disk not pruinose;<br />
hymenium inspersed; ascospores 10-11-celled 30-40 × 7-8 µm;<br />
norstictic plus stictic acid; Dade County.............................................................................. G. leptocarpa Fée s. lat.<br />
6. Exciple entire, carbonized below hymenium.......................................................................................................................... 13<br />
13. Ascospores 11-15-celled, 50-60 × 8-10 µm..................................................................................................................... 14<br />
14. Hymenium inspersed; lirellae mostly immersed; norstictic acid (rarely<br />
acid deficient); throughout the state................................................................................................. Graphis sp. 23509<br />
14. Hymenium not inspersed; lirellae sessile; no substances; Polk County................................. G. anfractuosa Eschw.<br />
13. Ascospores 6-8-celled, 18-25 × 6-7 µm; hymenium inspersed; norstictic acid;<br />
Duval and Polk counties......................................................................................G. desquamescens (Fée) Zahlbr. s. lat<br />
5. Epihymenium orange, KOH+ purple; margin not striate; disk exposed, orange....................................................................... 15<br />
15. Exciple closed; hymenium yellowish inspersed; ascospores 8-celled,<br />
27-34 × 7-8 µm; norstictic acid; Dade County..........................................................................G. chromothecia R. C. Harris<br />
15. Exciple lacking below; hymenium not inspersed; ascospores 8-celled,<br />
23-30 × 6-7 µm; no substances; Franklin and Liberty counties..........................................................G. inversa R. C. Harris<br />
2. Margin striate; hymenium not inspersed............................................................................................................................................ 16<br />
16. Thallus UV+ yellow (lichexanthone)............................................................................................................................................. 17<br />
17. Lirellae narrow, flexuose, branched; margin weakly striate; ascospores<br />
6-8-celled, 20-30 × 7-9 µm; south to Polk County ................................................................................ G. lucifica R. C. Harris<br />
17. Lirellae rather broad, short, often straight, sparsely branched; margin<br />
strongly striate; ascospores 11-14-celled, 50-75 × 9-12 µm;<br />
Franklin and Liberty counties ............................................................................................................... G. haleana R. C. Harris<br />
11
16. Thallus UV-................................................................................................................................................................................... 18<br />
18. Ascospores 14-19-celled, 55-80 µm long .............................................................................................................................. 19<br />
19. Stictic acid; exciple strongly developed below; ascospores ca. 14-celled,<br />
55 × 8-9 µm; Southwest Florida, Mar 1878, Smith & Austin 91.................................................................. Graphis sp. 91<br />
19. Norstictic acid; exciple barely closed below; ascospores 16-19-celled,<br />
70-80 × 10-11 µm; Louisiana, not yet known from Florida ..................................................[G. lumbricina Vainio s. lat.]<br />
18. Ascospores 8-12-celled, 20-45 µm long ................................................................................................................................ 20<br />
20. Margin carbonized to base of hymenium.......................................................................................................................... 21<br />
21. No substances; ascospores 10-12-celled, 35-45 × 7-9 µm;<br />
Duval and Lake counties....................................................................................................G. striatula (Ach.) Sprengel<br />
21. Stictic acid; ascospores 8-celled, 23-28 × 6-7 µm; Leon County .................................................... Graphis sp. 25050<br />
20. Margin carbonized above, orange below; ascospores 9-10-celled,<br />
25-27 × 6-7 µm often sterile; no substances; Duval and Marion counties ............................................G. subelegans Nyl.<br />
1. Exciple not carbonized; ascospores 2-4-celled .................................................................................................................................... 22<br />
.<br />
22. Ascospores 4-celled; no substances ............................................................................................................................................... 23<br />
23. Lirellae fissurine; exciple not evident.......................................................................................................................................... 24<br />
24. Margin not swollen; ascospores to 17 × 7 µm ....................................................................................................................... 25<br />
25. Thallus white, ecorticate; lirellae not gaping, minute, visible only as<br />
thin lines; ascospores 15-17 × 6-7 µm; Duval and Polk counties.................................................G. illiterata R. C. Harris<br />
25. Thallus greenish, with shiny prosoplectenchymatous cortex;<br />
lirellae gaping; ascospores 11-13 × 5-6 µm; Levy County.................................................... G. subnitidula Nyl. ex Tuck.<br />
24. Margin swollen; ascospores 17-20 × 8-10 µm; Duval, Lake and<br />
Marion counties ......................................................................................................... G. insidiosa (Knight & Mitten) Hook. f.<br />
23. Lirellae long, dichotomously branched; disk hidden to exposed;<br />
exciple orangish, well developed laterally or at base ................................................................................................................. 26<br />
26. Disk not exposed; exciple well developed laterally ............................................................................................................... 27<br />
27. Margin not striate; ascospores 15-17 × 6-8 µm;<br />
Duval and Liberty counties........................................................................................................................G. subparilis Nyl.<br />
27. Margin striate; ascospores 22-24 × 10-13 µm; East Florida, 1877, Smith..................................................G. rufula Mont.<br />
26. Disk exposed; "exciple" well developed below hymenium;<br />
ascospores 10-13 × 5 µm; Collier, Dade and Osceola counties .................................................................G. grammatis Fée<br />
22. Ascospores 2-celled or 8-9-celled ................................................................................................................................................... 28<br />
28. Ascospores 2-celled; norstictic acid; thallus whitish, shiny, UV-; lirellae crowded, initially<br />
with closed ± pruinose lips, later disk exposed, pale, pruinose; Duval County................................................ G. turbulenta Nyl.<br />
28. Ascospores 8-9-celled, 21-31 × 7.5-9 µm; thallus whitish, dull, UV+ yellow (lichexanthone);<br />
lirellae imnersed; disk pale red brown; margin white, patchily developed; Bay, Jefferson and<br />
Lafayette counties ......................................................................................................................................G. anguinoides ined.<br />
NOTES<br />
Graphis "anguinoides" ined.<br />
This species is superficially indistinguishable from Graphina anguina. I had actually placed a couple specimens in this<br />
species on the external characters of ascoma and UV+ thallus. When I checked the ascospores of some recent collections, I<br />
discovered a taxon with transversely septate, not muriform, ascospores. To my mind this is another nail in the coffin of the<br />
ascospore genera, Graphis vs. Graphina. It is inconceivable to me that genera defined on the single character of ascospore<br />
septation should have evolved parallel species with identical multicharacter complexes of thallus and ascoma.<br />
12
GYROSTOMUM Fr.<br />
Only the single species, G. scyphuliferum (Ach.) Nyl. in North America although there are two additional apparently undescribed<br />
species in Puerto Rico. See generic key for diagnosis.<br />
PHAEOGRAPHINA Müll. Arg.<br />
1. Exciple not carbonized ............................................................................................................................................................................ 2<br />
2. Disk initially hidden, finally exposed; margin white, separating from thallus;<br />
margin and disk densely white pruinose; hymenium not inspersed; ascospores<br />
8/ascus, 2-4 × 8-celled, 27-35 × 10-13 µm; no substances; Highlands and<br />
Polk counties ........................................................................................................................................................... P. asteroides Fink<br />
2. Disk exposed from beginning; disk weakly white pruinose; hymenium not<br />
inspersed; ascospores 8/ascus, 1-2 × 6-celled, 20-30 × 10-11 µm;<br />
norstictic acid; Highlands County.............................................................................................................Phaeographina sp. 18150B<br />
1. Exciple carbonized, at least at apices..................................................................................................................................................... 3<br />
3. Lirellae immersed or if emergent, not white......................................................................................................................................... 4<br />
4. Disk broad, bluish pruinose or not pruinose .................................................................................................................................... 5<br />
5. Lirellae immersed; disk not or weakly pruinose; exciple usually carbonized below;<br />
hymenium inspersed; ascospores 8/ascus, 2-4 × 6-8-celled, 24-30 × 9-12 µm;<br />
no substances; Duval and Lake counties ..........................................................................................................P. explicans Fink<br />
5. Lirellae emergent, large; disk usually bluish pruinose; exciple carbonized only<br />
at apex; hymenium inspersed; ascospores 8/ascus, muriform, 65-75 × 16-23 µm;<br />
no substances; south to Highlands County ..........................................................................P. caesiopruinosa (Fée) Müll. Arg.<br />
4. Disk narrow, white pruinose; margin black, carbonized only at edges; hymenium<br />
inspersed; ascospores 2-6/ascus, very variable, 35-50 × 13-24 µm; norstictic acid;<br />
Keys ..............................................................................................................................................Phaeographina sp. Small 10114<br />
3. Lirellae strongly emergent with thin white covering, cup-shaped, oval to weakly elongate<br />
and once branched; margin and base thick, heavily carbonized; hymenium inspersed;<br />
ascospores 8/ascus, 100-110 × 24-27 µm; no substances; Dade County............................. P. quassiaecola (Fée) Müll Arg. s. lat.<br />
PHAEOGRAPHIS Müll. Arg. & SARCOGRAPHA Fée<br />
1. Lirellae red; exciple not carbonized; ascospores 6-10-celled, 21-35 × 9-11 µm;<br />
Pinellas and Sarasota counties...............................................................................................................P. haematites (Fée) Müll. Arg.<br />
1. Lirellae white, brown or blackish ............................................................................................................................................................. 2<br />
2. Ascospores 4-celled ............................................................................................................................................................................. 3<br />
3. Ascomata not erumpent ................................................................................................................................................................... 4<br />
4. Hymenium inspersed..................................................................................................................................................................... 5<br />
5. Exciple entire, carbonized below .............................................................................................................................................. 6<br />
6. Lirellae simple to loosely asteriform, not surrounded by whitish "stroma";<br />
ascospores 15-16 × 5-6 µm; no substances or rarely norstictic acid;<br />
south to Polk County .............................................................................................................P. sericea (Eschw.) Müll. Arg.<br />
6. Lirellae densely branched, surrounded by a weakly raised whitish "stroma"....................................................................... 7<br />
7. No substances; ascospores17-21 × 7-8.5 µm; Highlands County ..................... Sarcographa tricosa (Ach.) Müll. Arg.<br />
7. Stictic acid agg.; ascospores 16-21 × 5.5-6 µm; Dade County......................................................................................<br />
.....................................................................................................................Sarcographa labyrinthica (Ach.) Müll. Arg.<br />
5. Exciple carbonized only at apex; hymenium inspersed; ascospores 16-21 × 6-10 µm;<br />
13
no substances; not yet found in Florida .........................................................................................[P. inusta (Ach.) Müll. Arg.]<br />
4. Hymenium not inspersed............................................................................................................................................................... 8<br />
8. Lirellae densely branched or crowded and irregular, surrounded by<br />
distinct whitish area ................................................................................................................................................................... 9<br />
9. Norstictic acid; whitish area weakly raised, not mealy; ascospores 15-20 × 5-6 µm;<br />
Polk and Seminole counties.......................................... Sarcographa intricans sensu Harris 1990 non (Nyl.) Müll. Arg.<br />
9. No substances; lirellae in mealy decorticate patches, brown, irregular,<br />
crowded ; ascospores 13-16 × 5-6 µm; Puntnam County .............................................................Phaeographis sp. 23697<br />
8. Lirellae simple to loosely branched, not surrounded by whitish area;<br />
ascospores 15-18 × 5-6 µm; norstictic acid or norstictic acid plus<br />
stictic acid or rarely acid deficient; Duval, Polk and Volusia counties.......................................P. subtigrina (Vainio) Zahlbr.<br />
3. Ascomata erumpent with more or less erect bark flaps ................................................................................................................ 10<br />
10. No substances; hymenium inspersed; ascospores 16-20 × 7-8 µm;<br />
Volusia County...............................................................................................................................P. subfulgurata (Nyl.) Zahlbr.<br />
10. Norstictic acid; hymenium not inspersed; ascospores 19-21 × 7-8 µm;<br />
Florida, without locality, Wilson...............................................................................................................Phaeographis sp. 1233<br />
2. Ascospores 6-10-celled....................................................................................................................................................................... 11<br />
11. Ascomata round to slightly elongated, not branched .................................................................................................................. 12<br />
12. Thallus UV+ yellow (lichexanthone); margin not raised; ascospores 6-celled,<br />
18-24 × 6-7 µm; south to Polk County............................................................................ P. punctiformis (Eschw.) Müll. Arg.<br />
12. Thallus UV-(no substances); margin often erect, jagged; ascospores 8-10-celled,<br />
25-40 × 7-10 µm; south to Polk County......................................................................................P. lobata (Eschw.) Müll. Arg.<br />
11. Ascomata elongate, often branched ............................................................................................................................................ 13<br />
13. Exciple not carbonized; hymenium not inspersed ................................................................................................................. 14<br />
14. Margin not striate............................................................................................................................................................... 15<br />
15. Lirellae simple or sparingly branched; disk weakly white pruinose;<br />
ascospores (4-)6-celled, 20-25 × 7-9 µm; norstictic acid;<br />
Highlands and Volusia counties.......................................................................................... Phaeographis sp. 23787B<br />
15. Lirellae much branched, angular, crowded, surrounded by poorly<br />
developed whitish area; disks bluish white pruinose; ascospores<br />
(4-)6-celled, 22-24 × 6-7 µm; stictic acid; Highlands, Lake and<br />
Volusia counties ....................................................................................... Sarcographa medusulina (Nyl.) Müll. Arg.<br />
14. Margin striate; lirellae long, flexuose, branched; disk becoming<br />
exposed, brown; ascospores 6-celled, 25-27 × 9-10 µm;<br />
Florida, without locality, Wilson......................................................................................................Phaeographis sp. 1259<br />
13. Exciple carbonized, at least at apex ..................................................................................................................................... 16<br />
15. Ascomata not erumpent ................................................................................................................................................... 17<br />
17. Hymenium inspersed with orange pigment, KOH+ dull purple;<br />
lirellae black, short, little branched; exciple carbonized at apex;<br />
ascospores 8-celled, 30-40 × 8-11 µm; norstictic acid;<br />
Monroe County ..................................................................................................................... P. multicolor R. C. Harris<br />
17. Hymenium colorless, KOH-; exciple entire; ascospores 6-8-celled,<br />
26-42 × 8-11 µm; norstictic acid; not yet found in Florida..............................................[P. dendritica (Ach.) Müll. Arg.]<br />
16. Ascomata erumpent with distinct erect bark flaps; exciple entire;<br />
hymenium inspersed; thallus often with a reddish tinge; ascospores<br />
6-7-celled, 27-32 × 8-9 µm; no substances; northern Florida ............................................ P. erumpens (Nyl.) Müll. Arg.<br />
NOTES<br />
14
Sarcographia labyrinthica (Ach.) Müll. Arg.<br />
Restored to the North American mycota. FLORIDA. Dade County: Everglades National Park, Mahogany Hammock, on large vine,<br />
2 Jan 1976, Buck B798 (NY).<br />
THELOTREMATACEAE (Nyl.) Stizenb.<br />
KEY TO GENERA<br />
1. Growing on bark ...................................................................................................................................................................................... 2<br />
2. Ascoma an apothecium........................................................................................................................................................................ 3<br />
3. Upper hymenium and ascus sheath I-; paraphyses mostly simple ................................................................................................. 4<br />
4. Periphysoids absent; columella present or absent....................................................................................................................... 5<br />
5. Exciple colorless to pale reddish brown; columellar structures,<br />
if present, noncarbonized..........................................................................................................................................Myriotrema<br />
5. Exciple carbonized; carbonized columella often present .........................................................................................Ocellularia<br />
4. Exciple with periphysoids on the inner face; columella always lacking.......................................................................Thelotrema<br />
3. Upper hymenium and ascus sheath I+ blue green; paraphyses somewhat<br />
anastomosed below, strongly so above the asci; apothecia immersed; hymenium<br />
separated from bark by thick layer of colorless crystals; periphysoids absent;<br />
ascospores colorless, vermiform, 30-40-celled, I-, ca. 85-120 × 7-9 µm;<br />
Seminole County ............................................................................................."Ocellularia" stictidea (Nyl.) V_zda ex Gilenstam<br />
2. Ascoma a mazaedium........................................................................................................................................................Nadvornikia<br />
1. Growing on soil or over Cladonia; spores brown, muriform ..............................................................................................Diploschistes<br />
DIPLOSCHISTES Norman<br />
1. Growing on Cladonia, at least initially; ascospores 4/ascus, 18-32 × 6-15 µm;<br />
lecanoric and diploschistesic acids; Polk County .................................................................................. D. muscorum (Scop.) R. Sant.<br />
1. Growing on soil, forming a continuous crust; ascospores 6-8/ascus, 20-28 × 8-14 µm;<br />
gyrophoric acid plus small amount of lecanoric acid; Gadsden County ............................................................ D. hypoleucus Zahlbr.<br />
MYRIOTREMA Fée<br />
1. Thallus not sorediate ............................................................................................................................................................................... 2<br />
2. Ascospores brown, submuriform to densely muriform ........................................................................................................................ 3<br />
3. Spores 8/ascus, submuriform .......................................................................................................................................................... 4<br />
4. Thallus PD+ orange or red............................................................................................................................................................ 5<br />
5. Thallus PD+ orange (stictic acid agg.)...................................................................................................................................... 6<br />
6. Ascomata rounded, not crowded .......................................................................................................................................... 7<br />
7. Norstictic acid lacking; ascomata round, slightly raised; disk not pulling<br />
away from thallus; Bay, Jackson, Liberty, Sarasota and Wakulla<br />
counties ........................................................................................................................... M. subcompunctum (Nyl.) Hale<br />
7. Norstictic acid in addition to stictic acid; ascomata smaller, more<br />
irregular; disk pulling away from thallus, Dade, Lake, Monroe and<br />
Seminole counties ................................................................................................................ M. compunctum (Ach.) Hale<br />
6. Ascomata angular, crowded, often in whitish, decorticate areas;<br />
15
ascospores 1-2 × 4-celled, 13-16 × 7-10 µm; throughout Florida .............................................M. glaucescens (Nyl.) Hale<br />
5. Thallus PD+ red (protocetraric acid); ascomata large, emergent with<br />
a simple to divided columella; ascospores 1-3 × 5-6-celled, 16-24 × 12-14 µm;<br />
Collier, Dade, Lee and Lake counties............................................................................................... M. bahianum (Ach.) Hale<br />
4. Thallus PD- (no substances), with large polygonal crystals, usually with<br />
pockets of bright red pigment; ascospores 2 × 4-celled, 14-18 × 10-14 µm;<br />
throughout Florida ....................................................................................................................................M. wightii (Taylor) Hale<br />
3. Spores 1-2/ascus, densely muriform, 70-130 × 20-30 µm; thallus PD+ orange<br />
(stictic acid agg.); common, south to Highalnds County ...............................................................M. reclusum (Krempelh.) Hale<br />
2. Ascospores colorless; apothecia smaller ........................................................................................................................................... 8<br />
8. Spores submuriform........................................................................................................................................................................ 9<br />
9. Apothecia small, 0.3-0.4 mm across, immersed; thallus light gray with<br />
numerous crystals; psoromic acid; ascospores 1-3 × 3-5-celled,<br />
9-20 × 4-6 µm; south to Hillsborough County .....................................................................................M. rugiferum (Harm.) Hale<br />
9. Apothecia larger, to 1.0 man, emergent, with broad rounded pore;<br />
thallus pale tan with numerous crystals; stictic acid agg. or acid deficient;<br />
ascospores 1-2 × 6-7-celled, 18-25 × 6-8 µm; throughout Florida ................................................... M. peninsulae R. C. Harris<br />
8. Spores transversely septate.......................................................................................................................................................... 10<br />
10. Cortex UV+ yellow, medulla PD- (lichexanthone, hypoprotocetraric acid);<br />
ascospores 4-celled, 10-12 × 5-6 µm; Dade and Monroe counties ................................................... M. glauculum (Nyl.) Hale<br />
10. Cortex UV-, medulla PD+ yellow (psoromic acid); ascospores 3-4-celled,<br />
9-15 × 4-7 µm; Alachua County....................................................................................................... M. terebratulum (Nyl.) Hale<br />
1. Thallus sorediate; protocetraric acid; ascospores colorless, 6-10-celled,<br />
42-46 × 9-11 µm; throughout Florida ................................................................................................................ M. erodens R. C. Harris<br />
NADVORNIKIA Tibell<br />
1. Thallus sorediate; throughout Florida ..............................................................................................................N. sorediata R. C. Harris<br />
1. Thallus not sorediate; Highlands County..................................................................................................N. hawaiiensis (Tuck.) Tibell<br />
OCELLULARIA G. Meyer<br />
1. Ascospores colorless .............................................................................................................................................................................. 2<br />
2. Ascospores transversely septate ......................................................................................................................................................... 3<br />
3. Ascoma smaller; medulla C-; ascospores under 50 µm long........................................................................................................... 4<br />
4. Thallus PD+ yellow or PD- ............................................................................................................................................................ 5<br />
5. Thallus PD+ yellow (psoromic acid).......................................................................................................................................... 6<br />
6. Columella present .................................................................................................................................................................. 7<br />
7. Ascospores 6-8-celled, 17-24 × 6-7 µm; apothecial margin<br />
not erect; pore usually small; columella undivided; common,<br />
throughout Florida ................................................................................................................ O. granulosa (Tuck.) Zahlbr.<br />
7. Ascospores 8(-9)-celled, 23-33 × 5-6 µm; apothecial margin<br />
erect; disk ± broad; columella becoming reticulate; rare,<br />
Collier County .............................................................................................................O. emersa (Krempelh.) Müll. Arg.<br />
6. Columella absent; ascospores 6-8-celled, 25-32 × 9-11 µm;<br />
Monroe County (Keys)..................................................................................................................Ocellularia sp. Small 9122<br />
5. Thallus PD- (no substances); apothecia emergent; rim often eroding<br />
16
to expose black rim around pore; columella present, black ..................................................................................................... 8<br />
8. Ascospores 4-6-celled, 13-20 × 5-8 µm; thallus dull, ecorticate;<br />
Hillsborough County ............................................................................................................ O. concolor Meyen & Flotow<br />
8. Ascospores 4-celled, 13-17 × 6-7 µm; thallus ± glossy, thinly<br />
corticate; Seminole County ...................................................................................................Ocellularia sp. Schallert 1664<br />
4. Thallus PD+ red (cinchonarum unknowns); apothecia emergent with round<br />
pore; columella present; ascospores 9-11-celled, 27-37 × 7.5-8.5 µm;<br />
Duval County ...................................................................................................................................... O. cavata (Ach.) Müll. Arg.<br />
3. Ascoma large, emergent, hemispherical; medulla pink, C+ pink (gyrophoric<br />
acid); ascospores very large, 19-23-celled, 100-130 × 20-35 µm; south to<br />
Lake County...........................................................................................................................................................O. americana Hale<br />
2. Ascospores densely muriform, 1-2/ascus, over 100 µm long ............................................................................................................. 9<br />
9. Thallus PD- (no substances), dull, usually roughened with small warts;<br />
medulla not conspicuous; south to Seminole County........................................................................O. sanfordiana (Zahlbr.) Hale<br />
9. Thallus PD+ orange (praestans unknown), shiny, very irregular; medulla<br />
conspicuous; Duval, Gulf and Nassau counties ..............................................................................O. praestans (Müll. Arg.) Hale<br />
1. Ascospores brown................................................................................................................................................................................. 10<br />
10. Ascospores submuriform ................................................................................................................................................................ 11<br />
11. Thallus PD- (no substances); ascospores 2 × 4-celled, 16-18 × 8-11 µm,<br />
with reticulate ornamentation; columella absent; Columbia and<br />
Wakulla counties ..................................................................................................................................O. retispora R. C. Harris<br />
11. Thallus PD+ yeiiow psoromic acid); ascospores 2 × 4-celled,<br />
13-16 × 8-10 µm, with weak striate ornamentation; known only<br />
from Louisiana ................................................................................................................................................[O. fissa (Nyl.) Hale]<br />
10. Ascospores transversely 4-celled ................................................................................................................................................... 12<br />
12. Apothecia immersed with small pore; columella undivided, not carbonized;<br />
pycnidia conspicuous, orange; hypoprotocetraric acid; ascospores<br />
15-18 × 8-10 µm, weakly granular or rugose warted; Columbia,<br />
Franklin, Leon, Liberty and Wakulla counties........................................................................O. leiostoma (Tuck.) R. C. Harris<br />
12. Apothecia immersed to emergent with broadly exposed disk; columella<br />
divided, carbonized; pycnidia not conspicuous; psoromic acid;<br />
ascospores 9-14 × 4-8 µm, smooth; Hillsborough County ......................................................... O. auberiana (Mont.) Hale<br />
NOTES<br />
Ocellularia auberiana (Mont.) Hale<br />
I excluded this species from North America in "Some Florida Lichens" as I could find no supporting specimen. I have since<br />
collected it. FLORIDA. Hillsborough County: Hillsborough River State Park, 28°09'N, 82°14'W, on oak, 3,4 Dec 1992, Harris 29684, 29722<br />
(NY).<br />
Ocellularia concolor Meyen & Flotow<br />
New to North America. FLORIDA. Hillsborough County: Hillsborough River State Park, 28°09'N, 82°14'W, on Carya and<br />
Liquidamabar, 3 Dec 1992, Harris 29680, 29686 (NY).<br />
Ocellularia emersa (Krempelh.) Müll. Arg.<br />
New to North America. FLORIDA. Collier County: Fakahatchee Strand State Preserve, Royal Palm Hammock, 26°01'N, 81°25'W,<br />
Buck 22763 (NY).<br />
THELOTREMA Ach.<br />
1. Thallus isidiate......................................................................................................................................................................................... 2<br />
2. Thallus PD+ yellow (psoromic acid), thinner ...................................................................................................................................... 3<br />
17
3. Ascospores brown, 2-celled, 8-11 × 5-6 µm, reticulately ornamented;<br />
isidia thick, long, often recumbent, "branching" by regeneration;<br />
apothecia emergent, doughnut-like; Liberty County.................................................................................. T. eximium R. C. Harris<br />
3. Ascospores colorless, submuriform, 4 × 4-celled, 15-21 × 9-12 µm;<br />
isidia sparse, initially globose; apothecia emergent, resembling Porina nucula Ach.;<br />
unknown substance in addition to psoromic acid; Leon, Okaloosa<br />
and Santa Rosa counties .............................................................................................................. T. halei (Tuck. & Mont.) Zahlbr.<br />
2. Thallus PD- (no substances), thick, filled with crystals, often separating<br />
from the substrate; isidia sparse, coarse; ascospores brown, submuriform,<br />
16-28 × 7-12 µm; south to Sarasota County...........................................................................................................T. santense Tuck.<br />
1. Thallus not isidiate................................................................................................................................................................................... 4<br />
4. Ascospores colorless .......................................................................................................................................................................... 5<br />
5. Ascospores transversely septate .................................................................................................................................................... 6<br />
6. Thallus PD- (no substances); ascomata small, immersed ......................................................................................................... 7<br />
7. Ascospores purple with iodine ................................................................................................................................................ 8<br />
8. Apothecia 0.4-0.6 mm across; thallus poorly developed, not corticate;<br />
medulla not obvious; ascospores 9-15-celled, 25-40 × 6-8 µm;<br />
northern Florida ...........................................................................................................................................T. subtile Tuck.<br />
8. Apothecia 0.2-0.35 m across; thallus usually well developed,<br />
weakly corticate; medulla thick; ascospores 8-13-celled, 25-45 × 6-8 µm;<br />
south to Polk County ............................................................................................................................. T. lathraeum Tuck.<br />
7. Ascospores colorless with iodine, 11-14-celled, 24-38 × 6-7 µm;<br />
apothecial disk mostly flush with thallus; Lake County ..........................................................T. alborosellum (Nyl.) Tuck .?<br />
6. Thallus PD+ yellow or orange; apothecia larger......................................................................................................................... 9<br />
9. Thallus PD+ orange (stictic acid agg.).................................................................................................................................. 10<br />
10. Apothecia with thick, recurved and split margin; ascospores 7-10-celled,<br />
30 × 6-8 µm; Florida Panhandle ......................................................................................... T. dilatatum (Müll. Arg.) Hale<br />
10. Apothecia emergent with broad round pore; exciple free from margin<br />
forming a conspicuous inner ring; ascospores to 30-celled,<br />
120-140 × 15-20 µm; exact locality unknown, known only from<br />
type of Ocellularia floridensis ......................................................................................T. porinoides Mont. & v.d. Bosch<br />
9. Thallus PD+ yellow (psoromic acid); apothecia emergent, round with broad<br />
round pore and smooth margin; exciple separating from margin; ascospores<br />
4-6-celled, 13-18 × 6-7 µm; Franklin and Liberty counties .............................................................T. floridense R. C. Harris<br />
5. Spores submuriform to muriform; thallus PD- (no substances) ................................................................................................... 11<br />
11. Ascospores large, over 50 µm long, 1-4/ascus ...................................................................................................................... 12<br />
12. Ascospores thick walled, 2-4/ascus, 65-90 × 15-20 µm ................................................................................................... 13<br />
13. Apothecia immersed when young, becoming urceolate with low,<br />
thick margin; exciple coarse, pinkish tan, often filling pore and<br />
covering the disk; Panhandle ...............................................................................................................T. adjectum Nyl.<br />
13. Apothecia with large, open pore; margin regular, thinner;<br />
exciple fragile, whitish, free from margin; Duval County ....................................................... T. lepadinum (Ach.) Ach.<br />
12. Ascospores thin walled, IKI-, 1-2/ascus, 50-80 × 15-20 µm; apothecia<br />
white pruinose with erect, jagged margin; south to Hillsborough<br />
County ......................................................................................................................................T. leprocarpum (Nyl.) Tuck.<br />
11. Ascospores 25-35(-45) × 12-15(-18) µm, initially 8/ascus but several may abort;<br />
apothecia small, immersed, radially striate and lacerate; Citrus, Columbia,<br />
Jackson, Leon and Union counties ................................................................................... T. defectum Hale ex R. C. Harris<br />
18
4. Ascospores light to dark brown......................................................................................................................................................... 14<br />
14. Ascospores 5-10 transversely septate......................................................................................................................................... 15<br />
15. Ascospores 6-celled, 17-20 × 6-7 µm; stictic acid agg.;<br />
apothecia with rim recurved and splitting; disk white<br />
pruinose; Collier County..................................................................................................................T. platycarpoides Tuck.<br />
15. Ascospores 7-11-celled, 33-60 × 6-10 µm; no substances;<br />
apothecia emergent with broad, round pores; exciple pulling<br />
away from margin; Santa Rosa and Seminole counties....................................................................... T. lacteum Krempelh.<br />
14. Ascospores densely muriform, 1/ascus, 105-140 × 24-30 µm;<br />
no substances; apothecia emergent with broad, round pores;<br />
exciple pulling away from margin; Dixie, Liberty, Marion<br />
and Santa Rosa counties ............................................................................................................. T. monospermum R. C. Harris<br />
NOTES<br />
Thelotrema platycarpoides Tuck.<br />
Thelotrema platycarpoides was originally described from Cuba. Hale (1981) was in error describing the ascospores as "3-4loculate".<br />
Tuckerman described them as 6-celled and an examination of an isolectotype (BUF) corroborates this. (The very similar T.<br />
platycarpum Tuck. does have 4-celled ascospores and also differs in larger apothecia.)<br />
New to North America. FLORIDA. Collier County, Fakahatchee Strand State Preserve, Royal Palm Hammock, 26°01'N, 81°25'W,<br />
6 Dec 1992, Harris 29940 (NY).<br />
"CALICIALES"<br />
"Calciales" are used here in an informal, highly artificial, sense to include not only the Caliciaceae and<br />
Mycocaliciaceae but mazaedioid genera belonging to other families or of uncertain position. Mycocaliciaceae<br />
and Caliciaceae are clearly not at all closely related. The familial disposition of Heterocyphelium Vainio and<br />
Tylophoron Nyl. is uncertain. The first is monotypic and treated in the generic key. The second is left in the<br />
Caliciaceae until its relationships are better understood.<br />
KEY TO GENERA<br />
1. Apothecia stalked or if almost sessile, then very small ............................................................................ 2<br />
2. Asci disintegrating, forming a mazaedium ........................................................................................... 3<br />
3. Ascospores simple ............................................................................................................................ 4<br />
4. Ascospores small (5 µm or less), more or less globose, pale,<br />
with weak ornamentation formed by cracking of exospore;<br />
lichenized, on bark or leaf bases of Sabal................................................................ Chaenotheca<br />
4. Ascospores larger (to 17µm), lemon-shaped, dark,<br />
with strong reticulate ribbing; apothecia top-shaped with short,<br />
thick stalk; parasymbiont on Pertusaria; so far only Gulf Coast<br />
from Dixie to Collier County.........................................................Sphinctrina tubiformis Massal.<br />
3. Ascopores 2-celled, brown .................................................................................................. Calicium<br />
2. Asci persistent, not forming mazaedium (Mycocaliciaceae) ................................................................5<br />
5. Growing on old wood or bark ............................................................................................................ 6<br />
6. Stalk dark, usually of more or less parallel hyphae;<br />
19
ascospores simple with pointed ends; ascus tip not<br />
pierced by narrow canal ...........................................................................................Mycocalicium<br />
6. Stalk pale, usually of irregularly arranged hyphae<br />
(very short in C. pilosa); ascospores simple or 2-celled,<br />
usually with rounded ends but sometimes as in Mycocalicium;<br />
ascus tip pierced by a narrow canal .................................................................. Chaenothecopsis<br />
5. Growing on the polypore, Trichaptum biforme (Fr.) Ryv.;<br />
ascospores 2-celled, pale, 10-14 × 3-5 µm;<br />
northern Florida .............................................................Phaeocalicium polyporaeum (Nyl.) Tibell<br />
1. Apothecia immersed or sessile, large ...................................................................................................... 7<br />
7. Ascospores 3-4-celled............................................................................................................................ 8<br />
8. Thallus UV+ yellow (lichexanthone); ascoma formed of a lower<br />
perithecioid chamber and upper cup-shaped chamber holding loose<br />
ascopores; ascospores 4-celled...........................................................................................................<br />
............................................... Pyrgillus javanicus (Mont. & v.d. Bosch) Nyl. (See Pyrenulaceae)<br />
8. Thallus UV-; ascoma not 2-chambered; ascospores usually 3-celled,<br />
with middle cell largest; no substances; Seminole County..................................................................<br />
................................................................................. Heterocyphelium leucampyx (Tuck.) Vainio<br />
7. Ascospores 2-celled............................................................................................................................... 9<br />
9. Apothecial margin and thallus C-, KOH+ yellow or red, PD+ orange<br />
(norstictic or stictic acid)................................................................................................................... 10<br />
10. Norstictic acid agg. present; ascospores 13-17 × 8-9 µm (Tibell,<br />
1976); peninsular Florida south to Highlands County ....................................................................<br />
....................................................................................... Thelomma carolinianum (Tuck.) Tibell<br />
10. Stictic acid agg. present; ascospores 7-9 × 5-6 µm (Tibell,1987) ..................................................<br />
...........................................................................................Nadvornikia (See Thelotremataceae)<br />
9. Apothecial margin C+ red (lecanoric acid), KOH-, PD-;<br />
ascospores with very dark middle region ......................................................................... Tylophoron<br />
CALICIACEAE Chev.<br />
CALICIUM Pers.<br />
1. Lowerside of capitulum white pruinose or not pruinose ............................................................................ 2<br />
2. Apothecia relatively large, over 0.5 mm tall............................................................................................ 3<br />
3. Thallus KC+ orange (xanthones), areolate and yellowish or more<br />
often in Florida material not evident; stalk IKI-; ascospores<br />
10.5-14 × 4.5-7.5 µm (Tibell, 1987); common, peninsular Florida ................... C. hyperelloides Nyl.<br />
3. Thallus KC- (no substances), granular; stalk IKI+ blue;<br />
ascospores 9-14 × 4-7.5 µm (Tibe11, 1987); Bradford,<br />
Leon and Marion counties .................................................................................... C. lenticulare Ach.<br />
20
2. Apothecia minute, ca. 0.2 mm tall; capitulum with faint yellow pruina?;<br />
ascospores with pointed ends, strongly spirally ornamented, 10-13 × 4.5-6 µm;<br />
on twigs; Citrus and Hamilton counties ........................................................ C. adspersum Pers. s. lat.<br />
1. Lowerside of capitulum brown pruinose; ascospores with spiral<br />
ornamentation ........................................................................................................................................... 4<br />
4. Spores larger, mostly over 11 × 6 µm ................................................................................................... 5<br />
5. Thallus UV+ yellow (lichexanthone), KC+ weak orange, with same<br />
unknown as Florida C. salicinum, not shiny; ascospores<br />
9-14 × 5.5-7 µm (Tibell, 1987); peninsular Florida ................................ C. chlorosporum F. Wilson<br />
5. Thallus UV-, KC+ red-orange (xanthones); ascospores 12.5-15.5<br />
× 7-8 µm; Gulf Coast from Taylor to Collier County ...................................... C. leucochlorum Tuck.<br />
4. Spores small, 7-11 × 3.5-5 µm (Tibell, 1987); thallus with unknown<br />
substance; Bay and Lake counties.................................................................C. salicinum Pers. s. lat.<br />
NOTES<br />
Calicium adspersum Pers. s. lat.<br />
Tibell (1987) considered this a variable species. Whether it is commodious enough to encompass the<br />
Florida material is open to question. The two collections, although very minute in stature, are placed here<br />
because of the faint yellow pruina and the rather narrow, mostly pointed ascospores. Its ecological<br />
preferences may prove to be significant, one collection is from small oak twigs, the other from a stem of the<br />
shrub Ceratiola.<br />
Calicium salicinum Pers. s. lat.<br />
The chemistry of this group is highly problematical. Tibell (1987) reported norstictic acid. In northern<br />
North American, European, Andean South American and Australian material I find a spot that runs much like<br />
norstictic acid but seems to differ in color and shape. I get no red needles with KOH on thallus sections. I<br />
regard norstictic acid as possible but unlikely. The Florida specimens and another from Puerto Rico contain a<br />
different substance in high concentration. It has some of the characteristics of and is possibly a dibenzofuran.<br />
This same substance is also found in C. chlorosporum accompanied by lichexanthone. Since the lowland<br />
neotropical population of C. salicinum is chemically distinct its taxonomic status should perhaps be reviewed.<br />
21
CHAENOTHECA Th. Fr.<br />
1. Base of capitulum brown to whitish, not pruinose; photobiont<br />
Chlorococcalean........................................................................................................................................ 2<br />
2. Thallus PD+ yellow (unknown); apothecia relatively tall and slender;<br />
ascospores 2.7-4.3 µm in diameter, more or less smooth (Tibell,1980);<br />
common, penisular Florida.....................................................................C. brunneola (Ach.) Müll. Arg.<br />
2. Thallus PD+ red-orange (virensic acid); apothecia relatively short and<br />
stout; ascospores 4-5 µm in diameter, rather coarsely ornamented;<br />
Hillsborough, Levy, Putnam and Sarasota counties.........................................C. floridana R. C. Harris<br />
1. Base of capitulum yellow pruinose; photobiont Trentepohlia;<br />
ascospores 3-4 µm in diam.; Bay County............................................. C. hispidula (Ach.) Zahlbr. s. lat.<br />
NOTES<br />
Chaenotheca brunneola (Ach.) Müll. Arg. s. lat.<br />
I cannot make out the chemistry reported by Tibell (1980,1987) for the brunneola-hygrophilastemonea<br />
complex. Most Florida collections of C. brunneola contain an unknown identical to that found in<br />
22
three collections of C. stemonea tested (New York, Minnesota, Sweden). It runs about the same as<br />
baeomyesic acid but the spot characteristics are different. Three collections of C. brunneola (Florida, Maine &<br />
Sweden) contain no detectable substances, four contain baeomycesic and squamatic acids (Florida, New<br />
York, North Carolina & Sweden). Tibell reported baeomycesic acid in1980 but rejected it in 1987. It seems<br />
probable that his unknowns B & C are the same as the single unknown spot that I get with my TLC system.<br />
However, it is clear that more work needs to be done to confirm this. The baeomycesic acid strain is known<br />
from a single collection at Gold Head Branch State Park (Clay County) where the unknown substance strain<br />
also occurs. Most specimens checked had asci in chains. Collections from Duval and Hernando counties had<br />
asci arising singly.<br />
Chaenotheca floridana R. C. Harris, sp. nov<br />
Ab aliis acidum virensicum continentia, ascosporis grosse ornamentis, 4-5 µm diametro et apothecis<br />
pallidis brevibus et crassiusculis differt.<br />
Type. FLORIDA. Putnam County: Ocala National Forest, old Johnson Field Campground along Oklawaha River<br />
just W of U.S. Hwy. 19, swamp forest, 7 Dec 1988, Buck 16798 (NY, holotype; B, UPS, isotypes).<br />
Thallus whitish, granular leprose, PD+ red-orange, virensic acid, unknown (tr.), protocetraric acid (tr.).<br />
Phycobiont trebouxioid. Apothecia relatively short and stout, to 1 mm tall. Stalk to 0.3 mm across at base<br />
tapering to ca. 0.1 mm, light orange brown (more or less concolorous with spore mass), appearing white<br />
pruinose due to wefts of loose hyphae or occasionally covered with thallus particles. Capitulum obconical to<br />
globose, to 0.5 mm in diameter. Asci not in chains. Ascospores globose or appearing slightly angular due to<br />
coarse plate-like ornamentation, 4-5 µm in diameter, yellow brown.<br />
I originally included this species in C. brunneola s. lat. until I studied its chemistry. A subsequent reevaluation<br />
showed that it also differs in the pale, shorter apothecia and slightly larger, coarsely ornamented<br />
ascospores.<br />
Additional specimens. FLORIDA. Hillsborough County: Hillsborough River State Park, Rapids Trail<br />
along river, 28°09'N, 82°14W, on trunk of Sabal, 3 Dec 1992, Buck 22641; Levy County: Cedar Key Scrub<br />
State Preserve, along Co. Rd. 347 ca. 1.5 mi N of Fla. Hwy. 24, on Sabal leaf bases, 30 Nov 1992, Buck<br />
22425, 22434, 22436; Sarasota County: Myakka River State Park, along Fla. Hwy. 72 ca. 0.7 mi SE of park<br />
entrance, on trunk of Sabal, 5 Dec 1992, Buck 22717, 22731 (all NY).<br />
Chaenotheca hispidula (Ach.) Zahlbr. s. lat.<br />
The single specimen varies from the North American norm in having the asci in chains and in smaller<br />
ascospores reaching only 4 µm which is below the minimum given by Tibell (1980). It is possible that the<br />
deviation is clinal as the locality is considerably south of the range given by Tibell.<br />
THELOMMA Massal.<br />
Thelomma carolinianum (Tuck.) Tibell is endemic to the Coastal Plain. The other species of the<br />
genus occur in western North America and Europe. See key to genera.<br />
TYLOPHORON Nyl. ex Stizenb.<br />
1. Thallus and ascomatal margin C+ red (lecanoric acid); ascomata<br />
0.6-1.2 mm diam. (Tibell, 1982); ascospores 12-15 × 6.5-8 µm;<br />
sporodochia white; conidiospores 1-2(-4)-celled, colorless,<br />
rough-walled; throughout Florida.................................................................................. T. protrudens Nyl.<br />
1. Thallus C-; ascomatal margin C+ red (lecanoric acid, 2'-O-methylperlatolic<br />
acid); ascomata 0.4-0.8 mm diam. (Tibell, 1982); ascospores 10-14 × 5-6 µm; sporodochia<br />
23
own (not known in Florida); conidiospores simple,<br />
brown; Collier County ...................................................................................................T. moderatum Nyl.<br />
NOTES<br />
Tibell (1991) described the sporodochial anamorph of T. moderatum but did not mention one for T.<br />
protrudens. However, Florida material always has conspicuous sporodochia. In fact, seven of the ten Florida<br />
collections consist of sporodochia only. The same anamorph was also found on Lindig 1 from Colombia. The<br />
number of cells in a conidium is apparently determined by how the hyphae break to form the chlamydosporelike<br />
conidia. In T. moderatum the sporodochia apparently do not occur on the same thallus as the ascomata.<br />
However, to my eye two collections of T. protrudens have both on the same thallus. There is only a single<br />
chemotype in Florida as neither lichexanthone nor the red ascomatal pigment was detected. Tylophoron<br />
protrudens is probably common but not often collected by lichenologists as it somewhat resembles members<br />
of the Aphyllophorales. It might also be confused with Cryptothecia striata Thor (Chiodecton montagnei auct.)<br />
that also seems sterile and is C+. A section of a sporodochium with conidiospores readily identifies the<br />
Tylophoron.<br />
SPHINCTRINACEAE Choisy<br />
SPHINCTRINA Fr.<br />
Only S. tubiformis is known from Florida. See key to genera.<br />
MYCOCALICIACEAE A. Schmidt<br />
KEY TO CHAENOTHECOPSIS AND MYCOCALICIUM<br />
1. Apothecia and thallus lacking lichen substances; ascospores simple,<br />
elliptical to fusiform.................................................................................................................................... 2<br />
2. Apothecia distinctly stalked, not hairy.................................................................................................... 3<br />
3. Exciple and stalk brown or stalk almost colorless; exciple not<br />
incurved and rugose......................................................................................................................... 4<br />
4. Stalk brown to dark brown; stalk hyphae more or less parallel .................................................... 5<br />
5. Cells of lower part of exciple not enlarged, not paraplectenchymatous .................................. 6<br />
6. Apothecia relatively broad and flat, ± conical, often<br />
2-3/stalk; stalk relatively short and stout; ascospores<br />
6-8 × 2.5-3.5 µm; on old wood; common, south to<br />
Palm Beach County ............................................... Mycocalicium subtile (Pers.) Szatala<br />
6. Apothecia narrowly obconical or obpyriform, often 2-3/stalk;<br />
stalk long, slender and flexuous; ascospores pale, 6-7 × 2.5-3 µm;<br />
rare, Washington County ................................Chaenothecopsis savonica (Räs.) Tibell?<br />
5. Cells of lower part of exciple enlarged and paraplectenchymatous;<br />
ascospores 6-8 × 2.5-3.5 µm; on old wood; Citrus and Sumter counties.................................<br />
24
.................................................................................... Mycocalicium albonigrum (Nyl.) Fink<br />
4. Stalk almost colorless to pale olive or pale brown; hyphae irregularly arranged.......................... 7<br />
7. Ascospores arranged end to end ............................................................................................. 8<br />
8. On Sabal palmetto; ascospores 7-9 × 3-4 um; common,<br />
south to Collier County ............................... Chaenothecopsis rappii (Nadv.) R. C. Harris<br />
8. On pine bark; ascospores 6-7 × 2.5-4 µm; rare,<br />
Franklin County ........................................................................Chaenothecopsis sp. 1393<br />
7. Ascospores obliquely arranged, 7-8 × 3.5-4 µm; on Quercus;<br />
rare, Citrus County .................................................................... Chaenothecopsis nana Tibell<br />
3. Exciple and stalk reddish, redder in KOH; exciple incurved, longitudinally<br />
rugose; ascospores 8-11 × 4.5-5.5 µm; on old wood; Santee Canal,<br />
South Carolina, not yet found in Florida ...................... [Mycocalicium ravenelii (Tuck. ex Nyl.) Fink]<br />
2. Apothecia sessile or with a very short stalk, hairy due to projecting<br />
excipular hyphae; ascospores simple; growing on thallus of sterile<br />
Tylophoron protrudens; Wakulla County................................Chaenothecopsis pilosa Tibell & Kalb<br />
1. Apothecia containing norstictic acid; ascospores 2-celled, ± cylindrical<br />
with rounded ends, 7-11 × 3-3.5 µm; on old wood; Bradford County and<br />
Okefenokee Swamp, Georgia............................................... Chaenothecopsis norstictica R. C. Harris<br />
NOTES<br />
CHAENOTHECOPSIS Vainio<br />
Chaenothecopsis norstictica R. C. Harris, sp. nov.<br />
Unica Mycocalicacearum acidum norsticticum continens.<br />
Type. GEORGIA. Charlton County: Okefenokee National Wildlife Refuge, Suwanee Canal Recreation Area,<br />
along "Swamp Trail" on Chesser Island, swamp forest, 12 Dec 1987, Buck 15462 (NY, holotype).<br />
Thallus not evident, on lignum, no algae seem to be associated with stalk bases. Cladonia ravenelii<br />
Tuck. is also on the wood but the Chaenothecopsis does not seem to be directly associated with it. Apothecia<br />
dark brown, slightly shiny, ca. 0.9 mm tall. Stalk dark brown; hyphae seem more or less parallel but hard to<br />
tell. Capitulum obconical, ca. 0.3-0.4 mm in diameter. Exciple and upper stalk bleeding yellow in KOH with<br />
subsequent formation of typical crystals of norstictic acid (confirmed by TLC), norstictic acid apparently<br />
present in substantial concentration. Asci with tip pierced by narrow canal. Ascospores pale brown, more or<br />
less cylindrical with rounded ends, 2-celled, 7-11 × 3-3.5 µm; septum dark.<br />
Chaenothecopsis norstictica is apparently unique among the Mycocaliciaceae in producing a<br />
depsidone. The chemistry of the family otherwise seems confined to various unknown pigments or rarely the<br />
pulvinic acid group.<br />
Additional specimens. FLORIDA. Bradford County: Lake Butler Wildlife Management Area, Raiford<br />
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<br />
swamp, 3 Dec 1994, Buck 27333, 27343, 27348 (NY).<br />
25
Chaenothecopsis pilosa Tibell & Kalb, Nova Hedwigia 60: 211. 1995.<br />
New to North America. FLORIDA. Wakulla County: Apalachicola National Forest, along Forest Service<br />
Rd. 309 at Lost Creek just S of Leon Co. line, 5.6 mi W of Fla. Hwy. 267, swamp forest, on Tylophoron<br />
protrudens on Fraxinus, 4 Dec 1988, Harris 23372 (NY).<br />
Chaenothecopsis rappii (Nádv.) R. C. Harris, comb. nov.<br />
Mycocalicium rappii Nádv., Ann. Mycol. 40: 139. 1942.<br />
This species has simple pointed ascospores as in Mycocalicium but Tibell includes other species with this<br />
ascospore type in Chaenothecopsis. The stalk is pale with irregularly arranged hyphae and the ascus tip has<br />
a narrow canal which are considered by Tibell, 1987) as better indicators of generic placement if one wishes to<br />
recognize Chaenothecopsis as distinct from Mycocalicium. It would be simpler not to.<br />
MYCOCALICIUM Vainio<br />
Two species are known from Florida. See key above. Mycocalicium ravenelii is close to M. calicioides<br />
(Nádv.) Tibell. Both seem to me quite distinct from M. subtile, the generitype, and probably should be<br />
segregated into Sphinctrinella Nádv. (type, S. calicioides).<br />
PHAEOCALICIUM A. Schmidt<br />
Only the single species, P. polyporaeum (Nyl.) Tibell, occurs in Florida and is on the polypore, Tricaptum<br />
biforme (Fr.) Ryv. (Polyporus pargamenus Fr.). See key to genera.<br />
LEOTIALES<br />
BAEOMYCETACEAE Dumortier<br />
(incl. Icmadophilaceae Triebel)<br />
I do not think that recognition of Icmadophilaceae is warrented and does not seem to be in line with<br />
general practice in the leotioid ascomycetes. The generic split between Baeomyces Pers. and Dibaeis<br />
Clements is well supported. Dibaeis baeomyces (L. fil.) Rambold & Hertel ( syn. Baeomyces roseus Pers.) is<br />
the only species in Florida. It is obviously at the southern edge of its North American range. It was collected<br />
on a road bank in Escambia County and disintegrated sandstone at Rock Hill, Washington County.<br />
LECANORALES<br />
COLLEMATACEAE Zenker<br />
KEY TO GENERA<br />
1. Ascospores septate; thallus without reticulate maculae......................................................................................................................... 2<br />
2. Thallus with cortex...............................................................................................................................................................Leptogium<br />
2. Thallus without cortex.............................................................................................................................................................. Collema<br />
1. Ascospores simple; thallus with reticulate markings (maculae) due to absence of photobiont;<br />
apothecia large, with very thick wrinkled margin; thallus tomentose below;<br />
reported by Moore (1970) from Citrus County, not seen by me ......................................................... Physma byrsaeum (Ach.) Tuck.<br />
26
COLLEMA Wigg.<br />
[The non-lichenized loculoascomycete, Myriangium duriaei Montagne & Berkeley forms small cushions with apothecium-like ascomata<br />
and can in the field be confused with a tiny Collema. The brown, cellular tissue, lack of photobiont and globose asci with muriform<br />
ascospores readily distinguish it in the lab.]<br />
1. Lobes large and broad ............................................................................................................................................................................ 2<br />
2. Lobes longitudinally ridged and pustulate, isidiate or not.................................................................................................................... 3<br />
3. Isidia absent (C. pulcellum Ach. s. lat.)............................................................................................................................................. 4<br />
4. Ascospores narrower, under 5 µm wide ....................................................................................................................................... 5<br />
5. Apothecial disks strongly white pruinose; south to Polk County .................................................................................................<br />
.........................................................................................................................C. pulcellum var. leucopeplum (Tuck.) Degel.<br />
5. Apothecial disks not pruinose; Duval County ......................................................................C. pulcellum Ach. var. pulcellum<br />
4. Ascospores broad, over 5 µm wide; apothecial disk occasionally pruinose;<br />
north to Flagler County.............................................................................C. pulcellum var. subnigrescens (Müll. Arg.) Degel.<br />
3. Isidia present, mainly on ridges and pustules; apothecial disk pruinose; proper exciple<br />
paraplectenchymatous; Citrus, Collier, Hernando and Levy counties (sterile),<br />
"S.W. Florida" (with apothecia) ..............................................................C. furfurceum (Arn.) Du Rietz var. luzone (Räs.) Degel.<br />
2. Lobes not ridged or pustulate, with ± globose isidia; a single scrappy<br />
collection from Madison County tentatively assigned here ........................................................................... C. subflaccidum Degel.<br />
1. Lobes narrow, often anastomosing......................................................................................................................................................... 6<br />
6. Ascospores rod-shaped with more than 2 cells or if broader, submuriform ...................................................................................... 7<br />
7. Apothecia subglobose, mostly immersed in the thallus; ascospores globose to ellipsoid, submuriform ..................................... 8<br />
8. Thallus foliose; proper exciple paraplectenchymatous; ascospores ellipsoid,<br />
2-3 × 4(-6)-celled, 20-40 × 11-15 µm (Degelius); Dade County.................................................................... C. pustulatum Ach.<br />
8. Thallus subfoliose; proper exciple prosoplectenchymatous; ascospores globose<br />
to subglobose, 2 × 3-4-celled, 8.5-13 µm in diameter or 10.5-17 × 8.5-10.5 µm;<br />
Jackson County and reported by Degelius (1974) from Caloosa River ......................................................C. callibotrys Tuck.<br />
7. Apothecia cup-shaped, sessile to stalked; ascospores narrow, ca. 2 µm wide,<br />
transversely septate with blunt ends; Leon and Seminole counties ................................................................ C. leptaleum Tuck.<br />
6. Ascospores 2-celled, 8.5-21 × 3-6.5 µm; thallus lobes small and crowded;<br />
apothecia usually numerous; rare, Citrus and Hillsborough counties......................................................C. conglomeratum Hoffm.<br />
LEPTOGIUM (Ach.) Gray<br />
Since the bulk of NY's collections of Leptogium are on loan and the new treatment, when it appears, is likely to encompass<br />
considerable change, I include Moore's key for completeness and convenience.<br />
1. Thallus larger, clearly foliose................................................................................................................................................................... 2<br />
2. Thallus isidiate...................................................................................................................................................................................... 3<br />
3. Thallus wrinkled................................................................................................................................................................................. 4<br />
4. Lobes distinct and not anastamosing, the margins not thickened; isidia not granular ................................................................ 5<br />
5. Thallus longitudinally wrinkled; isidia concentrated along the ridges;<br />
throughout Florida ......................................................................................................................L. isidiosellum (Riddle) Sierk<br />
5. Thallus irregularly wrinkled; isidia marginal or scattered on the surface ................................................................................. 6<br />
6. Isidia mostly marginal and associated with apothecia; on hardwood trees;<br />
throughout Florida ............................................................................................................. L. marginellum (Swartz) S. Gray<br />
27
6. Isidia scattered on the surface of the thallus; on trees and rocks; throughout<br />
the state .................................................................................................................... L. austroamericanum (Malme) Dodge<br />
4. Lobes irregularly anastamosing, the margins thickened; isidia becoming granular;<br />
scattered throughout Florida .........................................................................................................................L. milligranum Sierk<br />
3. Thallus not wrinkled; throughout Florida.................................................................................... L. cyanescens (Rabenh.) Körber<br />
2. Thallus not isidiate................................................................................................................................................................................ 7<br />
7. Thallus smooth, not wrinkled............................................................................................................................................................ 8<br />
8. Thallus not pitted .......................................................................................................................................................................... 9<br />
9. Thallus blue-gray, lobes 1-5 mm broad; scattered throughout Florida ......................................... L. azureum (Swartz) Mont.<br />
9. Thallus gray-brown, lobes 2-3 mm broad; Duval County?, reported by Sierk<br />
on the basis of a Calkins collection..........................................................................................................L. juniperinum Tuck.<br />
8. Thallus pitted; peninsular Florida ........................................................................................................... L. microstictum Vainio<br />
7. Thallus wrinkled.............................................................................................................................................................................. 10<br />
10. Spores acicular or fusiform ...................................................................................................................................................... 11<br />
11. Spores acicular, 50-85 µm long; thallus gray; Sarasota and Seminole counties.................................... L. adpressum Nyl.<br />
11. Spores fusiform, 25-40 µm long; thallus usually brownish; peninsular Florida<br />
north to Levy County...............................................................................................................L. fusisporum (Tuck.) Dodge<br />
10. Spores ellipsoid or subfusiform................................................................................................................................................ 12<br />
12. Lobes distinct, not anastamosing........................................................................................................................................ 13<br />
13. Thallus naked above, shiny, with large pustules; cited by Moore<br />
from four counties, not found by me nor have I seen her material,<br />
Florida records possibly equal the next species............................................................. L. corticola (Taylor) Tuck.<br />
13. Thallus sparsely hairy above; hairs short with cylindrical cells; thallus naked<br />
below, dull, with small pustules ................................................................................................ Leptogium sp. 31815<br />
12. Lobes irregularly anastomosing .......................................................................................................................................... 14<br />
14. Thallus heavily wrinkled longitudinally, lobes narrow and<br />
elongate or inflated and hollow ................................................................................................................................ 15<br />
15. Apothecia not at the ends of hollow lobes; cortical cells<br />
usually more than 5 µm wide.............................................................................................................................. 16<br />
16. Thallus thickened but not warty, generally appearing gray<br />
or gray-green ................................................................................................................................................. 17<br />
17. Thallus gray; thalloid exciple lobulate; scattered throughout<br />
the state ................................................................................................... L. phyllocarpum (Pers.) Mont.<br />
17. Thallus green or gray-green; thalloid exciple not lobulate;<br />
scattered throughout the peninsula................................................................L. chloromelum (Ach.) Nyl.<br />
16. Thallus warty, usually appearing bluish; north to Lake<br />
and Seminole counties..................................................................................................... L. floridanum Sierk<br />
15. Apothecia at the ends of elongate hollow lobes; cortical cell<br />
usually less than 5 µm wide; peninsular Florida from Levy and<br />
Marion counties south to Palm Beach County......................................................................L. stipitatum Vainio<br />
14. Thallus wrinkled but flattened, lobes rounded; Lake, Polk and<br />
Seminole counties ............................................................................................................................ L. sessile Vainio<br />
1. Thallus small, appearing almost crustose; reported from Seminole County by Sierk ....................L. tenuissimum (Dickson) Körber<br />
28
PILOCARPACEAE Zahlbr.<br />
There are more Florida species of this family in genera other than Byssoloma. I have chosen to treat<br />
only Byssoloma at this time since the most common North American species has not been previously<br />
recognized. Further, the generic limits in the family are not clear to me at this time and the other North<br />
American species seem to fall into groups which may merit generic status. The key owes much to Santesson<br />
(1952).<br />
BYSSOLOMA Trevisan<br />
1. Thallus UV-; hypothecium dark brown; mostly on leaves.......................................................................... 2<br />
2. Margin whitish, KOH-; disk brownish to black ....................................................................................... 3<br />
3. Disk yellow brown to dirty brown, often mottled; exciple pale;<br />
ascospores10-18 × 2.5-4 µm; on leaves and bark; common,<br />
south to Polk County .....................................................................B. leucoblepharum (Nyl.) Vainio<br />
3. Disk black; exciple often greenish tinted; ascospores<br />
10-17 × 3-5 µm; on leaves; Lake, Seminole, Taylor,<br />
and Wakulla counties ................................................................. B. subdiscordans (Nyl.) P. James<br />
2. Margin yellow brown, KOH+ yellow; disk brown to brownish black;<br />
ascospores 9-15 × 3-4 µm; on leaves; Highlands,<br />
Marion and Seminole counties ........................................................... B. tricholomum (Mont.) Zahlbr.<br />
1. Thallus UV+ orange (xanthones); hypothecium yellowish; byssoid margin<br />
pale; disk yellowish becoming yellow brown, often mottled; ascospores<br />
10-15 × 2.5-4 µm; on bark; common, south to Polk County........................................................................<br />
........................................................................................................B. pubescens V_zda ex R. C. Harris<br />
NOTES<br />
Byssoloma pubescens V_zda ex R. C. Harris, sp. nov.<br />
Superficie similis B. leucoblephari sed thallo UV+ aurantiaco et hypothecio pallido.<br />
Type. FLORIDA. Seminole County: Sanford, on oak, May 1911, Rapp (NY, holotype).<br />
Thallus corticolous, gray green, dull, smooth, continuous, UV+ orange (xanthones). Apothecia<br />
sessile, orbicular, flattened, tightly adnate, 0.2-0.7 mm diam.; byssoid margin moderately to slightly developed,<br />
usually not spreading, whitish, KOH-; disk pale whitish yellow to dirty yellowish, often mottled with darker<br />
yellow brown, sometimes entirely darkening to dirty brown. Epithecium ± colorless in section. Hymenium<br />
nearly colorless or yellowish. Hypothecium nearly colorless or yellowish, rarely light brownish. Ascospores 4celled,<br />
10-15 × 2.5-4 µm. Microconidia lageniform, 3-4 × 1.5 µm.<br />
This very commonly collected species sometimes grows mixed with the rather similar B.<br />
leucoblepharum for which it has been mistaken until recently. Byssoloma pubescens is easily distinguished<br />
under the ultraviolet lamp by the intense orange fluorescence of the thallus. Also under the dissecting<br />
microscope the absence of a dark hypothecium is readily evident. The taxon was first recognized by V_zda 30<br />
years ago when he annotated the type specimen with this name. I believe all other species of the genus have<br />
a dark hypothecium. This raises the question as to generic limits, which in the Pilocarpaceae are, I think,<br />
rather fuzzy. Byssoloma, apparently can have an essentially non-byssoid margin but I am unaware of the<br />
29
everse, i.e., anything other than Byssoloma with a byssoid margin. Thus far B. pubescens is always on bark,<br />
ranging from tiny twigs to large tree trunks, mainly in mesic or swamp forest habitats. In North America it is a<br />
species of the southern Coastal Plain and is also known from Cuba and Puerto Rico.<br />
Additional specimens. FLORIDA. Bay County: N of Co. Rd. 388 at Econfina Creek, 7.9 mi W of U.S.<br />
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,<br />
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,<br />
30°16'N, 85°10'W, 11 Dec 1993, Buck 24769; Dixie County: Steinhatchee Wildlife Management Area,<br />
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<br />
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,<br />
83°04'W, 4 Dec 1993, Buck 24354; Duval County: Big Talbot Island St. Park, ca. 0.7 mi S of northern<br />
boundary, on Lyonia ferruginea, 17 Dec 1987, Buck 15513, Jacksonville, Calkins; Escambia County: along Co.<br />
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<br />
24550, on Ilex, Harris 31933; Franklin County: Apalachicola National Forest, Hickory Landing Rec. Area, ca. 2<br />
mi S of Sumatra, 23 May 1976, Harris 11304; Gilchrist County: Waccasassa Flats, along Co. Rd. 232, ca. 7.5<br />
mi N of Trenton, 29°44'N. 82°48'W, 5 Dec 1993, Buck 24415; Hamilton County: Bee Haven Bay, N of Co. Rd.<br />
6, E of Jasper, 30°30'N, 82°45'W, on branches of Taxodium,15 Dec 1993, Harris 32472, Holton Creek Wildlife<br />
Management Area, 30°26'N, 83°04'W, on Quercus, 5 Dec 1994, Harris 36099; Jackson County: Black Spring,<br />
ca. 6 mi S of Marianna, 3 Dec 1988, Harris 23330, 23346; Jefferson County: E of Fla. 59, 8.4 mi S of Wacissa,<br />
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,<br />
29°51'N, 83°15'W, 2 Dec 1994, Buck 27308; Lake County: Ocala National Forest, ca. 2 mi SE of Alexander<br />
Springs, on Magnolia virginiana, 6 Dec 1988, Buck 16735; Liberty County: Apalachicola National Forest,<br />
Ochlockonee River at Forest Serv. Rd. 13 crossing, 3 May 1990, Harris 25026; Madison County: 0.3 mi on dirt<br />
road W of Co. Rd. 150, NNE of Greenville, 30°36'N, 83°33'W, on Quercus, 14 Dec 1993, Buck 24888, Harris<br />
32369, 32382; Marion County: Ocala National Forest, ca. 0.75 mi NE of Doe Lake, 11 Dec 1988, Buck 16901,<br />
just E of Oklawaha River of Fla. Hwy. 316, 10 Aug 1985, Harris 18036; Polk County: Nalcrest, on Ilex, 28 May<br />
1989, Wheeler s.n.; Santa Rosa County: Blackwater River State Forest, Jackson Red Ground Trail from Red<br />
rock Rd., ca. 12 mi NE of Milton, 30°47'N, 86°53'W, 9 Dec 1993, Buck 24623, 24624, Harris 31990, 32002;<br />
Wakulla County: Edward Ball Wakulla Springs State Park, ca. 30°14'N, 84°18'W, on bamboo culm, 26-27 Dec<br />
1990, Buck 18991-A. GEORGIA. Charlton County: Okefenokee National Wildlife Refuge, N edge of Middle<br />
Island, 22 Nov 1989, Wetmore 65012; Coffee County: Broxton Rocks Ecological Preserve, on dead azalea,<br />
16-17 Dec 1993, Harris 32722, Ricketson Tract, on Vitis, 4 Feb 1995, Buck 27481. MISSISSIPPI. Winston<br />
County: Tombigbee National Forest, Choctaw Wildlife Area, 33°16'N, 89°06'W, on Magnolia, 28 Sep 1992,<br />
Harris 28661. NORTH CAROLINA. Dare Couny: Kitty Hawk, Nearing s.n., Nags Head Woods Ecological<br />
Preserve, 35°58'N, 75°06'W, on Liquidambar, 29 Sep 1993, Harris 31031 (all NY).<br />
CUBA. Prov. Santiago de Cuba: vicinity of Museo de la Francesca, "La Isabelita," ca. 2 km SE of La<br />
Gran Piedra, ca. 1000 m, 7 Apr 1982, Harris 14599 (HAC, NY). PUERTO RICO. Distr. Mayagüez: Maricao State<br />
Forest, Area de Recreo "Lookout," along Hwy 120 at Km 13.9, ca. 800 m, 28 May 1989, Buck 17241, 17265<br />
(all NY).<br />
CLADONIACEAE Zenker<br />
CLADINA (Nyl.) Nyl.<br />
1. Podetia yellowish, usnic acid present; PD+ red or PD-; branching dichotomous.................................................................................. 2<br />
2. Colonies without dense rounded heads; PD+ red (fumarprotocetraric acid);<br />
throughout Florida ................................................................................................................... C. subtenuis (des Abb.) Hale & Culb.<br />
2. Colonies with dense rounded heads; PD- (atranorin and perlatolic acid);<br />
throughout Florida ........................................................................................................................ C. evansii (des Abb.) Hale & Culb.<br />
1. Podetia not yellowish, usnic acid absent; PD+ red, fumarprotocetraric acid present............................................................................ 3<br />
3. Atranorin present; branching dichotomous or whorled........................................................................................................................ 4<br />
4. Branching dichotomous; pycnidial gel reddish; Everglades and Panhandle<br />
extending into southern Georgia....................................................................................................... C. sandstedei (des Abb.) Ahti<br />
4. Branching whorled; pycnidial gel colorless; Duval County (I have come to<br />
30
doubt the occurrence of this species in Florida. It needs to be confirmed<br />
with recent collections) ...................................................................................................................................[C. rangiferina (L.) Nyl.]<br />
3. Atranorin absent; branching dichotomous; Bay, Franklin and Hamilton counties ....................... C. subtenuis f. cinerea (Ahti) Ahti<br />
CLADONIA Hill ex Browne<br />
1. Apothecia and/or pycnidia red................................................................................................................................................................. 2<br />
2. Podetia simple or sparsely branched................................................................................................................................................... 3<br />
3. Podetia corticate, not sorediate/decorticate .................................................................................................................................... 4<br />
4. Thallus KOH+, PD+ deep yellow (thamnolic acid)........................................................................................................................ 5<br />
5. Usnic acid absent; podetia gray................................................................................................................................................ 6<br />
6. Podetia well developed, often branched; squamules small, often dissolving into<br />
sorediate masses; didymic acid; throughout Florida ................................................................................. C. ravenelii Tuck.<br />
6. Podetia poorly developed; squamules large, at most marginally sorediate,<br />
with broad yellow band below; pigments; throughout Florida ..............................................................C. hypoxantha Tuck.<br />
5. Usnic acid present; podetia yellowish, short, turbinate; didymic and squamatic acids<br />
or in one collection from Hillsborough County didymic acid is replaced by grayanic<br />
acid; often on Taxodium; throughout Florida .......................................................................................... C. abbreviatula Merr.<br />
4. Thallus KOH-, PD-; usnic acid present ......................................................................................................................................... 7<br />
7. Barbatic acid; squamules esorediate; northern tier of counties ................................................................. C. cristatella Tuck.<br />
7. Grayanic and squamatic acids; squamules often marginally sorediate;<br />
Lake, Liberty and Polk counties ........................................................................................................... C. anitae Culb. & Culb.<br />
3. Podetia sorediate or becoming mostly decorticate.......................................................................................................................... 8<br />
8. Barbatic and/or thamnolic acid...................................................................................................................................................... 9<br />
9. Soredia granular...................................................................................................................................................................... 10<br />
10. Podetia becoming decorticate; exposed stereome translucent; didymic acid;<br />
throughout Florida ............................................................................................................................................................. 11<br />
11. Barbatic acid........................................................................................................ C. didyma (Fée) Vainio var. didyma<br />
11. Thamnolic acid, often with accessory barbatic acid ......................C. didyma var. vulcanica (Zoll. & Moritzi) Vainio<br />
10. Podetia mostly corticate with a few sorediate patches....................................................................see C. ravenelii above<br />
9. Soredia farinose; barbatic acid; exposed stereome opaque; Leon County................................................... C. bacillaris Nyl.<br />
8. Squamatic, didymic and usnic acids; squamules abundantly sorediate;<br />
podetia very short; south to Polk County ......................................................................................................C. incrassata Flörke<br />
2. Podetia abundantly branched, forming lax, tangled colonies on sand; usnic, didymic,<br />
baeomycesic and squamatic acids; throughout Florida ................................................................................................C. leporina Fr.<br />
1. Apothecia and/or pycnidia brown or pallid............................................................................................................................................ 12<br />
12. Squamules smaller, not loose from substrate and curling into balls.............................................................................................. 13<br />
13. Podetia repeatedly dichotomously branched, "shrubby"; thallus PD-<br />
or PD+ yellow (baeomycesic acid); usnic acid present .............................................................................................................. 14<br />
14. Cortex lacking or poorly developed........................................................................................................................................ 15<br />
15. Podetia greenish with poorly developed cortex; podetial wall thin; cartilaginous<br />
tissue more continuous; Rock Hill Preserve,Washington County................................................................................... 16<br />
31
16. Thallus coarse, generally somewhat upright to sprawling ;<br />
tips abruptly narrowed, generally not curved downward;<br />
usnic acid only in Florida material .................................................................C. caroliniana Schweinitz ex Tuck.<br />
16. Thallus thin; tips gradually narrowing and downward pointing;<br />
usnic acid only in Florida material .................................................................................. C. dimorphoclada Robbins<br />
15. Podetia gray-white, ecorticate; podetial wall very thick, with scattered<br />
cartilaginous strands; south to Broward County ........................................................................ C. pachycladodes Vainio<br />
14. Cortex well developed............................................................................................................................................................ 17<br />
17. Podetia coarse, erect, hollow, perforate; PD-, squamatic acid;<br />
Highlands and Okaloosa counties ........................................................................................................ C. perforata Evans<br />
17. Podetia delicate, in lax, tangled, recumbent colonies; PD+ yellow<br />
(baeomycesic and squamatic acids); north to Polk County......................................... C. subsetacea Robbins ex Evans<br />
13. Podetia simple or sparingly branched........................................................................................................................................ 18<br />
18. Thallus PD-............................................................................................................................................................................ 19<br />
19. Podetia not forming cups ................................................................................................................................................ 20<br />
20. Squamules coarse, maculate; podetia proliferating, perforate,<br />
broadly expanded above; numerous apothecia on ultimate<br />
proliferations; squamatic acid; on Taxodium; Franklin and<br />
Liberty counties........................................................................................................................... C. buckii R. C. Harris<br />
20. Squamules thinner, emaculate; podetia mostly simple, mostly decorticate,<br />
narrower than the apothecia; barbatic acid; Duval and Seminole counties...................................................................<br />
................................................................................................................................... C. botryocarpa Merr. ex Sandst.<br />
19. Podetia forming broad cups, coarsely sorediate/areolate; grayanic acid;<br />
northern Florida .........................................................................................................................C. grayi Merr. ex Sandst.<br />
18. Thallus PD+ yellow, orange or orange-red ........................................................................................................................... 21<br />
21. Thallus PD+ yellow or orange .......................................................................................................................................... 22<br />
22. Podetia not forming cups ........................................................................................................................................... 23<br />
23. Squamules linear, large; podetia rarely squamulose, broader at top;<br />
norstictic acid or norstictic and stictic acids ......................................................................................................... 24<br />
24. Norstictic and stictic acids; south to Polk County ..............................................................C. polycarpia Merr.<br />
24. Norstictic acid ± atranorin; northern Florida.................................................................. C. polycarpoides Nyl.<br />
23. Squamules irregular, small; podetia often squamulose; thamnolic or<br />
baeomycesic acid ................................................................................................................................................. 25<br />
25. Thallus KOH+ deep yellow (thamnolic and barbatic acids)........................................................................... 26<br />
26. Cortex dispersed, coarsely areolate/squamulose; podetia<br />
short, rarely branched; central Florida ......................................................................... C. santensis Tuck.<br />
26. Cortex continuous; podetia taller, with few squamules,<br />
commonly branched, especially above; south to Polk County .................................... C. floridana Vainio<br />
25. Thallus KOH- ................................................................................................................................................. 27<br />
27. Squamules and podetia C-, KC-(baeomycesic and<br />
squamatic acids); podetia densely squamulose;<br />
south to Charlotte County..................................................................... C. beaumontii (Tuck.) Vainio<br />
27. Squamules and podetia C+, KC+ blue-green (strepsilin,<br />
baeomycesic and squamatic acids); podetia usually not<br />
densely squamulose, often lacking; rare, Holmes and<br />
Washington counties ........................................................................... C. strepsilis (Ach.) Grognot<br />
32
22. Podetia forming abruptly flared, centrally proliferating cups;<br />
psoromic acid; north to Seminole County..........................................................................C. rappii Evans var. rappii<br />
21. Thallus PD+ orange-red (fumarprotocetraric acid).......................................................................................................... 28<br />
28. Podetia forming broad cups ....................................................................................................................................... 29<br />
29. Cups not centrally proliferating, sorediate/areolate.............................................................................................. 30<br />
30. Fumarprotocetraric and either grayanic or cryptochlorophaeic acids ............................................................ 31<br />
31. Grayanic acid (this strain may not be in Florida) .................................................C. grayi Merr. ex Sandst.<br />
31. Cryptochlorophaeic acid; rare, Franklin County ..................................C. cryptochlorophaea Asahina<br />
30. Fumarprotocetraric acid only; reported from peninsular Florida<br />
by Moore, NY holds a couple of dubious records................................. C. chlorophaea (Sommerf.) Sprengel<br />
29. Cups centrally proliferating, abruptly flared; smoothly<br />
corticate; south to Polk County..........................................................C. rappii Evans var. exilior (Abbayes) Ahti<br />
28. Podetia not forming cups or forming very narrow, irregular cups ............................................................................ 32<br />
32. Podetia not sorediate ........................................................................................................................................... 33<br />
33. Podetia simple, short; apothecia much exceeding stalk or<br />
podetia gradually swelling to the apothecia.................................................................................................... 34<br />
34. Squamules small, unlobed, appressed, imbricate;<br />
apothecia exceeding stalk; south to Polk County................................... C. peziziformis (With.) Laundon<br />
34. Squamules larger, somewhat lobed, usually recurved;<br />
apothecia ± equal to stalk; Duval County, a somewhat<br />
dubious Calkins collection....................................................................................... C. sobolescens Nyl.<br />
33. Podetia taller, branched, often with poorly developed cups;<br />
squamules lobed, erect to recurved; south to Polk County............................................... C. simulata Robbins<br />
31. Podetia sorediate ................................................................................................................................................. 35<br />
35. Primary squamules at most marginally sorediate........................................................................................... 36<br />
36. Podetia awl-shaped or irregular; soredia farinose or<br />
granular; grayanic acid lacking [C. ochrochlora Flörke<br />
will key here. There is no trustworthy record for<br />
the state] .................................................................................................................................................... 37<br />
37. Podetia with farinose to granular soredia above,<br />
granular soredia mixed with isidioid squamules below;<br />
cortex dissolved except for basal collar; throughout<br />
Florida....................................................................................................C. subradiata (Vainio) Sandst.<br />
37. Podetia often mostly or partly corticate with patches<br />
of coarse soredia; exposed stereome translucent;<br />
throughout Florida ..........................................................................C. ramulosa (With.) Laundon s.lat.<br />
36. Podetia short, stubby with coarse granular soredia;<br />
containing grayanic acid; rare, Union County...........................................C. cylindrica (Evans) Evans<br />
35. Primary squamules dissolving into a sorediate crust;<br />
podetia tiny, pointed; Franklin County.......................................................................................... C. nana Vainio<br />
12. Squamules very large, curled into balls when dry; usually without podetia;<br />
on sand; south to Polk County ............................................................................................................................ C. prostrata Evans<br />
BIATORACEAE Stizenb.<br />
33
PHYLLOPSORA Müll. Arg.<br />
Note: This treatment mainly follows Brako (1990). Sterile isidiate crusts which are doubtfully<br />
squamulose should also be sought in the key to sterile isidiate crusts.<br />
1. Thallus PD- ............................................................................................................................................... 2<br />
2. Prothallus pale (in North America rarely darkening in old, abused specimens).................................... 3<br />
3. Squamules with marginal proliferations ± parallel to the substrate, flattened<br />
to some degree and not distinctly cylindrical; proliferations initially simple but<br />
usually repeatedly branched or proliferate ....................................................................................... 4<br />
4. Squamules large; lobes more than 0.5 mm wide .......................................................................... 5<br />
5. Marginal proliferations usually abundant, narrow, mostly ecorticate,<br />
forming a distinct white fringe; cortex less than 50 µm thick; ascospores<br />
ellipsoid to short fusiform, 8-12.5 × 2-3 µm; south to Highlands<br />
County ................................................................P. parvifolia (Pers.) Müll. Arg. var. parvifolia<br />
5. Marginal proliferations scarce, broader, similar to primary squamules;<br />
cortex over 50 µm thick; ascospores ovoid, 7-10.5 × 2.5-4 µm; Alabama,<br />
not yet known from Florida........................................ [P. parvifolia var. breviuscula (Nyl.) Brako]<br />
4. Squamules smaller, 0.1-0.5 mm wide .......................................................................................... 6<br />
6. Squamules adnate; proliferations narrow in respect to squamule,<br />
initially ± ovate becoming elongated, with constrictions or branched ......................................7<br />
7. Cortex type 2; very irregular, ± non-linear proliferations usually<br />
abundant and forming overlapping layers; ascospores 8.5-12<br />
× 2.5-3 µm; common, south to Hillsborough County..............................................................<br />
........................................................................................... P. confusa Swinscow & Krog<br />
7. Cortex type 1; proliferations less abundant less commonly elongated;<br />
ascospores 7.5-11.5 × 2-3 µm; reported by Brako from Alachua<br />
and Duval counties.................................... P. corallina (Eschw.) Müll. Arg. var. corallina<br />
6. Squamules erect; proliferations broad in respect to squamule or<br />
squamules adnate; proliferations coarse and inflated-looking, not<br />
much elongated ....................................................................................................................... 8<br />
8. Squamules ± erect with broad, flat lobules; ascospores<br />
4-6.5 × 2-3 µm; Lake County ............................................. P. parvifoliella (Nyl.) Müll. Arg.<br />
8. Squamules closely adnate, often contiguou and forming ±<br />
continuous crust; proliferations coarse, appearing inflated;<br />
an abberant population from Sharkey County, Mississippi,<br />
not yet known from Florida; referred in hb. by Brako<br />
to .............................................................................................. [cf. P. corallina var. corallina]<br />
3. Squamules with ± globose to cylindrical isidia or squamules lacking isidia<br />
and marginal proliferations .............................................................................................................. 9<br />
9. Thallus isidiate ............................................................................................................................ 10<br />
10. Squamules medium, 0.1-0.5 mm wide ................................................................................. 11<br />
34
11. Isidia cylindrical .............................................................................................................. 12<br />
12. Isidia common, often obpyriform, constricted at base<br />
and broadest just above base; atranorin and parvifoliin;<br />
ascospores 6.5-10 × 2.5-3.5 µm; south to<br />
Sarasota County .................................................... P. corallina var. rappiana Brako<br />
12. Isidia rare, cylindrical; squamules mostly with marginal<br />
proliferations; no substances; seen by me only from<br />
South Carolina........................................[P. corallina (Eschw.) Müll. Arg. var. corallina]<br />
11. Isidia nearly globose or irregular, not much elongated or weakly<br />
elongated by buddiing; no substances; ascospores 6.5-12.5<br />
× 2-3.5 µm; Louisiana and Oklahoma, not yet known from<br />
Florida ....................................................................................................... [P. kalbii Brako]<br />
10. Squamules/areoles minute, ca. 0.1 mm wide; isidia long,<br />
cylindrical, tangled; no substances; ascospores<br />
7.5-12.5 × 2-3.5 µm; Leon County .............................................P. isidiolyta (Vainio) Riddle<br />
9. Thallus not isidiate or lobulate; see couplet 5 ................................. [P. parvifolia var. breviuscula]<br />
2. Prothallus dark, abundant ................................................................................................................. 13<br />
13. Squamules minute, inconspicuous; thallus dominated by long<br />
slender isidia; furfuracein; ascospores 6-12 × 2-2.5 µm;<br />
Dade and Seminole Counties ...........................................................P. furfuracea (Pers.) Zahlbr.<br />
13. Squamules/areoles large, tightly adjoined forming an almost<br />
continuous crust, separated by white lines formed by fibrils,<br />
isidiate; isidia sparse, short, cylindrical; atranorin; Louisiana,<br />
not yet known from Florida ........................................................................[P. halei (Tuck.) Zahlbr.]<br />
1. Thallus PD+ orange (argopsin agg. or pannarin) .................................................................................... 14<br />
14. Thallus isidiate, not pruinose (P. corallina (Eschw.) Müll. Arg. s. lat.)............................................... 15<br />
15. Containing argopsin agg. as major substances .......................................................................... 16<br />
16. Lacking phyllopsorins; ascospores 7-13 × 2.5-3 µm;<br />
common, south to Sarasota County........................P. corallina var. santensis (Tuck.) Brako<br />
16. Containiing phyllopsorins; ascospores 6.5-12 × 2.5-3.5 µm;<br />
reported by Brako from Florida without locality (prob. Lake<br />
County) ...............................................................P. corallina var. ochroxantha (Nyl.) Brako<br />
15. Containing vicanicin as major substance with traces of<br />
norvicanicin and argopsin; ascospores 7-11 × 2.5-3 µm;<br />
reported by Brako from Dade and Seminole<br />
counties ........................................................................P. corallina var. glaucella (Vainio) Brako<br />
14. Thallus not isidiate, pruinose; squamules large<br />
(P. buettneri (Müll. Arg.) Zahlbr. s. lat.) ............................................................................................ 17<br />
35
17. Containing argopsin agg.; ascospores 7-12 × 2.5-3 µm;<br />
Wakulla County ...................................................P. buettneri var. glauca (B. de Lesd.) Brako<br />
17. Containing pannarln, phyllopsorin and zeorin; ascospores<br />
10-12 × 2.5-3 µm; reported by Brako from Louisiana,<br />
not yet found in Florida.................................................. [P. buettneri var. munda (Malme) Brako]<br />
NOTES<br />
Phyllopsora buettneri (Müll. Arg.) Zahlbr. var. glauca (B. de Lesd.) Brako<br />
This variety of P. buettneri has not been reported from North America previously. The specimen was<br />
determined by Lois Brako.<br />
FLORIDA. Wakulla County: Swamp forest, Lost Creek at Forest Service Rd. 309, just S of Leon County<br />
line, 4 Dec 1988, Harris 23375 (NY).<br />
PARMELIACEAE Zenker<br />
KEY TO THE GENERA<br />
1. Thallus foliose; central strand absent ..................................................................................................................................................... 2<br />
2. Lobes ciliate on margins or in axils...................................................................................................................................................... 3<br />
3. Cilia not swollen at base ................................................................................................................................................................... 4<br />
4. Lobes broad, 4-20 mm wide; cilia usually conspicuous ............................................................................................................... 5<br />
5. Underside pale at margin, black in center, with only one type of rhizine ................................................................................. 6<br />
6. Cortex not or only weakly maculate, not or only slightly cracked;<br />
rhizines lacking in marginal zone; chemistry various......................................................................................... Parmotrema<br />
6. Cortex maculate and abundantly cracked; rhizines usually reaching margin;<br />
salazinic acid .............................................................................................................................................................. Rimelia<br />
5. Underside uniformly pale to medium brown, with numerous short rhizines<br />
and fewer long ones ; Florida species isidiate; rare (previously<br />
included in Parmotrema) ......................................................................................................................................... Rimeliella<br />
4. Lobes narrow, 0.5-6.0 mm wide; cilia inconspicuous................................................................................................................... 7<br />
7. Medulla yellow, especially below soralia, C+ yellow (pigments, terpenoids);<br />
soralia pustular, breaking down to form coarse soredia; south to Seminole County..................................................................<br />
................................................................................................................................Myelochroa aurulenta (Tuck.) Elix & Hale<br />
7. Medulla white, C+ pink (gyrophoric or hiasic acid complex) ............................................................................. Parmelinopsis<br />
3. Cilia swollen at base (bulbate) ......................................................................................................................................................... 8<br />
8. Cortex gray (atranorin) ................................................................................................................................................. Bulbothrix<br />
8. Cortex yellow (usnic acid) ................................................................................................................................................ Relicina<br />
2. Lobes not ciliate on margins or in axils................................................................................................................................................ 9<br />
9. Thallus not pseudocyphellate......................................................................................................................................................... 10<br />
10. Cortex or medulla yellow.......................................................................................................................................................... 11<br />
11. Lobes narrow, 0.5-6.0 mm wide.......................................................................................................................................... 12<br />
12. Cortex yellow (usnic acid) ............................................................................................................................................... 13<br />
13. Growing on bark; thallus pustulate; pustules breaking down<br />
to form coarse soredia; divaricatic acid; south to Lake<br />
County................................................................................................................ Parmeliopsis subambigua Gyelnik<br />
13. Growing on sandstone; isidiate; stictic acid agg.; known only from<br />
Rock Hill, Washington County, new to Florida ......................................Xanthoparmelia conspersa (Ach.) Hale<br />
36
12. Medulla yellow; thallus not sorediate or isidiate ............................................................................................................. 14<br />
14. Medulla C+, KC+ orange (pigments); apothecia laminal; common,<br />
throughout Florida ............................................................................... Pseudoparmelia sphaerospora (Nyl.) Hale<br />
14. Medulla C-, KC- (vulpinic acid); apothecia marginal;<br />
Jackson and Walton counties....................................................... Vulpicida viridis (Schweinitz) Mattson & Lai<br />
11. Lobes broad, 4-20 mm wide............................................................................................................................... Parmotrema<br />
10. Cortex gray or brownish; medulla white...................................................................................................................................... 15<br />
15. Lobes narrow, 0.5-6.0 mm wide............................................................................................................................................ 16<br />
16. Thallus with "normal" rhizines, thick felty rhizine mat lacking.......................................................................................... 17<br />
17. Cortex KOH+ yellow; pycnidia laminal........................................................................................................................ 18<br />
18. Underside with inconspicuous pale to brown, simple rhizines............................................................................. 19<br />
19. Cortex KOH+ pale yellow (atranorin) .......................................................................................... Canoparmelia<br />
19. Cortex KOH+ deep yellow, PD+ deep yellow (thamnolic acid);<br />
isidiate; lobes linear; cortex shiny; Duval County.................................Imshaugia aleurites (Ach.) Meyer<br />
18. Underside with conspicuous dark, dichotomously branched rhizines;<br />
medulla KC+ purple (lividic acid agg.) .............................................................................................. Hypotrachyna<br />
17. Cortex KOH-; pycnidia conspicuous, marginal; fatty acids;<br />
on pine; south to Lake County ......................................................................... Tuckermanopsis fendleri (Nyl.) Hale<br />
16. Thallus with a thick spongy mat of rhizines below; marginally isidiate;<br />
divaricatic acid; rare, Escambia County, new to Florida ..........................................Anzia ornata (Zahlbr.) Asahina<br />
15. Lobes broad, 4-20 mm wide................................................................................................................................. Parmotrema<br />
9. Thallus white pseudocyphellate, isidiate; medulla C+ red (lecanoric acid);<br />
south to Seminole County ...............................................................................................................Punctelia rudecta (Ach.) Hale<br />
1. Thallus fruticose with a central solid cartilaginous strand ............................................................................................................. Usnea<br />
ANZIA Stizenb.<br />
Anzia ornata (Zahlbr.) Asahina has previously been reported from Alabama and North Carolina. Thus, its occurrence in<br />
northernmost Florida is not especially notable. FLORIDA. Escambia County: along Co. Rd. 99A at Brushy Creek, 4.3 mi W of St.<br />
Rd. 97 at Walnut Hill, 30°53'N, 87°32'W, floodplain woods, on Magnolia, 8 Dec 1993, Harris 31913 (NY).<br />
BULBOTHRIX Hale<br />
1. Thallus not isidiate; apothecia present ................................................................................................................................................... 2<br />
2. Apothecia lacking bulbate rhizines (ecoronate); medulla C+ red (lecanoric acid);<br />
south to Polk County ............................................................................................................................ B. confoederata (Culb.) Hale<br />
2. Exciple with bulbate rhizines (coronate); medulla C+ pink (gyrophoric acid);<br />
Texas, not yet known from Florida ................................................................................................................. [B. coronata (Fée) Hale]<br />
1. Thallus isidiate; apothecia absent........................................................................................................................................................... 3<br />
3. Medulla C+ pink to red, KOH-; lobes linear ......................................................................................................................................... 4<br />
4. Gyophoric acid present; south to Polk County.........................................................................................B. goebelii (Zenker) Hale<br />
4. Lecanoric acid present; south to Lake County ........................................................................................ B. laevigatula (Nyl.) Hale<br />
3. Medulla C-, KOH+ red (salazinic acid); lobes short, truncate to rounded;<br />
isidia often globose, brown; south to Polk County...............................................................................................B. isidiza (Nyl.) Hale<br />
NOTES<br />
37
Bulbothrix isidiza (Nyl.) Hale<br />
In the previous edition I speculated that this species might be recently introduced. However, an examination of Moore's specimens<br />
of Canoparmelia salacinifera revealed several collections of B. isidiza, putting an end to this notion.<br />
CANOPARMELIA Elix & Hale<br />
1. Thallus isidiate........................................................................................................................................................................................ 2<br />
2. Medulla PD+ orange or red................................................................................................................................................................. 3<br />
3. Medulla KOH- (protocetraric acid) .................................................................................................................................................. 4<br />
4. Isidia globose to cylindrical; throughout Florida....................................................................... C. amazonica (Nyl.) Elix & Hale<br />
4. Isidia inflated, somewhat lobulate; Monroe County (Keys) ................................................... C. martinicana (Nyl.) Elix & Hale<br />
3. Medulla KOH+ red (saiazinic acid); lobe tips unmarked;<br />
south to Highlands County ....................................................................................................... C. salacinifera (Hale) Elix & Hale<br />
2. Medulla PD- (perlatolic acid); lobe tips with reticulate<br />
white maculae; throughout Florida...................................................................................................C. caroliniana (Nyl.) Elix & Hale<br />
1. Thallus sorediate ..................................................................................................................................................................................... 5<br />
5. Soralia capitate on upturned lobules; medulla KC+ purple (cryptochlorophaeic acid);<br />
throughout Florida .............................................................................................................C. cryptochlorophaea (Hale) Elix & Hale<br />
5. Soralia pustular, laminal; medulla KC+ pinkish (divaricatic acid); Leon and<br />
Seminole counties according to Moore, not seen by me..................................................................... C. texana (Tuck.) Elix & Hale<br />
See Parmeliopsis.<br />
FORAMINELLA Fricke-Meyer<br />
HYPOTRACHYNA (Vainio) Hale<br />
1. Thallus pustular-sorediate....................................................................................................................................................................... 2<br />
2. Thallus UV+ yellow (lichexanthone); very common, south to<br />
Polk County ................................................................................................................................. H. osseoalba (Vainio) Park & Hale<br />
2. Thallus UV- (atranorin); Duval and Wakulla counties ........................................................................ H. pustulifera (Hale) Skorepa<br />
1. Thallus not pustulate-sorediate, UV-; apothecia common; south to Lake County.............................................. H. livida (Taylor) Hale<br />
IMSHAUGIA Fricke-Meyer<br />
Only I. aleurites (Ach.) Meyer known from Florida. Previously treated as Parmeliopsis.<br />
MYELOCHROA (Asahina) Elix & Hale<br />
Only the single species M. aurulenta (Tuck.) Elix & Hale is known from Florida.<br />
PARMELINOPSIS Elix & Hale<br />
1. Thallus isidiate......................................................................................................................................................................................... 2<br />
2. Isidia usually tipped with ciiia; hiasic acid complex; south to<br />
Pinellas County............................................................................................................................ P. horrescens (Taylor) Elix & Hale<br />
2. Isidia usually eciliate; gyrophoric acid and a UV+ white unknown;<br />
northern part of the state ................................................................................................................. P. minarum (Vainio) Elix & Hale<br />
1. Thallus sorediate; gyrophoric acid and a UV+ white unknown .............................................................................................................. 3<br />
3. Soredia coarse, derived from pustules; Marion, Wakulla and<br />
Walton counties........................................................................................................................... P. spumosa (Asahina) Elix & Hale<br />
38
3. Soredia farinose, not derived from pustules; Leon County .....................................................P. cryptochlora (Vainio) Elix & Hale<br />
PARMELIOPSIS (Stizenb.) Nyl<br />
Only the single species P. subambigua Gyelnik is known from Florida. It was treated in the previous edition as Foraminella.<br />
PARMOTREMA Massal.<br />
1. Lobes lacking marginal cilia .................................................................................................................................................................... 2<br />
2. Thallus isidiate...................................................................................................................................................................................... 3<br />
3. Medulla white, C+ red (lecanoric acid); common, throughout Florida .................................................... P. tinctorum (Nyl. ) Hale<br />
3. Medulla yellow. C+ pink (gyrophoric acid, ± echinocarpic acid);<br />
throughout Florida ...................................................................................................................... P. endosulphureum (Hillm.) Hale<br />
2. Thallus sorediate or lacking both soredia and isidia ........................................................................................................................... 4<br />
4. Thallus sorediate .............................................................................................................................................................................. 5<br />
5. Medulla KOH-, PD+ red or PD-..................................................................................................................................................... 6<br />
6. Medulla PD+ red (protocetraric acid)........................................................................................................................................ 7<br />
7. Soralia white ........................................................................................................................................................................... 8<br />
8. Protocetraric acid only; common, throughout Florida.................................................... P. gardeneri (Dodge) Sérusiaux<br />
8. Protocetraric acid with echinocarpic acid, usnic acid and unknowns;<br />
common, throughout Florida ......................................................................................................P. dilatatum (Vainio) Hale<br />
7. Soralia yellowish (usnic acid); south to Polk County,<br />
reported by Moore, not seen by me ....................................................................................... P. dominicanum (Vainio) Hale<br />
6. Medulla PD- (fatty acids); generally smaller than most Parmotrema;<br />
common, throughout Florida ....................................................................................................P. praesorediosum (Nyl.) Hale<br />
5. Medulla KOH+ red, PD+ orange ................................................................................................................................................... 9<br />
9. Salazinic acid present; common, Peninsular Florida.................................................................... P. cristiferum (Taylor) Hale<br />
9. Norstictic acid present; Dade and Volusia counties ........................................................................P. rubifaciens (Hale) Hale<br />
4. Thallus not sorediate; medulla PD+ red (protocetraric acid); conidia sublageniform;<br />
Dade and Monroe counties.......................................................................................................................P. zollingeri (Hepp) Hale<br />
1. Lobes with marginal cilia ....................................................................................................................................................................... 10<br />
10. Thallus isidiate................................................................................................................................................................................. 11<br />
11. Cortex or medulla yellow........................................................................................................................................................... 12<br />
12. Cortex yellowish (usnic acid); medulla white; medulla C- (fatty acids)<br />
or C+ pink (gyrophoric acid, i.e., including P. madagascariaceum (Hue) Hale);<br />
south to Marion County ........................................................................................................P. xanthinum (Müll. Arg.) Hale<br />
12. Medulla bright yellow (vulpinic acid); common, Peninsular<br />
Florida...................................................................................................................... P. sulphuratum (Nees & Flotow) Hale<br />
11. Cortex gray; medulla white......................................................................................................................................................... 13<br />
13. Medulla KOH+ yellow or red, UV- or UV+ yellow in lower part ............................................................................................ 14<br />
14. Medulla KOH+ yellow (stictic acid agg.); throughout Florida ........................................................P. crinitum (Ach.) Hale<br />
14. Medulla KOH+ red (salazinic acid); lower medulla UV+ yellow<br />
(lichexanthone); south to Marion County .................................................................................P. ultralucens (Krog) Hale<br />
39
13. Medulla KOH-, UV+ bright blue white (alectoronic acid); isidia often ciliate;<br />
south to Lake County........................................................................................................................ P. mellissii (Dodge) Hale<br />
2. Thallus sorediate or neither sorediate nor isidiate................................................................................................................................ 15<br />
15. Thallus sorediate ............................................................................................................................................................................. 16<br />
16. Medulla KOH-, UV+ bright blue white (alectoronic acid); cortex not maculate;<br />
common throughout Florida ............................................................................................................ P. rampoddense (Nyl.) Hale<br />
16. Medulla KOH+ yellow (stictic acid agg.); cortex maculate;<br />
common throughout Florida ......................................................................................................P. hypoleucinum (Steiner) Hale<br />
15. Thallus not sorediate....................................................................................................................................................................... 17<br />
17. Cortex maculate; medulla PD+ yellow (norstictic acid) or PD-................................................................................................. 18<br />
18. Norstictic acid only present (rarely acid deficient);<br />
south to Highlands County ....................................................................................................P. perforatum (Jacq.) Massal<br />
18. Alectoronic acid with or without norstictic acid present;<br />
common, throughout Florida ..........................................................................................................P. rigidum (Lynge) Hale<br />
17. Cortex not maculate; medulla PD+ red (protocetraric acid); conidia rod-shaped;<br />
throughout Florida .................................................................................................................... P. michauxianum (Zahlbr.) Hale<br />
PSEUDOPARMELIA Lynge<br />
Only the single species P. sphaerospora (Nyl.) Hale is known from Florida.<br />
PUNCTELIA Krog<br />
Only the single species P. rudecta (Ach.) Krog is known from Florida.<br />
RELICINA (Hale & Kurok.) Hale<br />
1. Thallus isidiate; medulla KOH+ red (norstictic/salazinic acid agg.);<br />
Baker County, reported by Moore, not seen by me, but recently collected<br />
at Broxton Rocks, Coffee county, Georgia..................................................................................................... R. abstrusa (Vainio) Hale<br />
1. Thallus not isidiate or sorediate; medulla KOH-, PD+ red (fumarprotocetraric acid agg.);<br />
Monroe County (Keys)............................................................................................................................ R. eximbricata (Gyelnik) Hale<br />
RIMELIA Hale & Fletcher<br />
1. Thallus sorediate; throughout Florida .........................................................................................R. reticulata (Taylor) Hale & Fletcher<br />
1. Thallus isidiate; south to Polk County............................................................................... R. subisidiosa (Müll. Arg.) Hale & Fletcher<br />
RIMELIELLA Kurokawa<br />
1. Norlobaridone present; Bradford County........................................................................................R. subtinctoria (Zahlbr.) Kurok.<br />
1. Norlobaridone absent; Bradford and Wakulla counties ......................................................R. neotropica (Kurok. & Hale) Kurok.<br />
TUCKERMANOPSIS Gyelnik<br />
Only T. fendleri (Nyl.) Hale occurs in Florida. Tuckermanopsis viridis has been removed to Vulpicida. See key to genera.<br />
USNEA Dill. ex Adanson<br />
1. Cortex, medulla or axis red or pink ......................................................................................................................................................... 2<br />
2. Cortex not red; medulla or axis red or pink.......................................................................................................................................... 3<br />
3. Main axes hollow; medulla red or pink............................................................................................................................................. 4<br />
4. Medulla usually red; norstictic acid present; sorediate-isidiate; south to<br />
40
Lake County U. baileyi (Stirton) Zahlbr.<br />
4. Medulla usually pink; diffractaic acid; isidiate-sorediate or sorediate; south<br />
to Lee County ..............................................................................................................................................U. perplectata Motyka<br />
3. Main axes solid................................................................................................................................................................................. 5<br />
5. Thallus shrubby to pendant without numerous perpendicular branchlets;<br />
apothecia rare; medulla pink or axis red; isidiate-sorediate......................................................................................................... 6<br />
6. Axis reddish; fatty acids; south to Lake Okeechobee................................................................................ U. mutabilis Stirton<br />
6. Axis unpigmented; medulla pale pink; diffractaic acid; branches<br />
often coarse; Duval County..............................................................................................................................U. ceratina Ach.<br />
5. Thallus shrubby with numerous perpendicular branchlets; main branches often<br />
ending in apothecia; not sorediate or isidiate; medulla red; chemistry various;<br />
south to Lake Okeechobee ................................................................................................................U. strigosa (Ach.) A. Eaton<br />
2. Cortex red mottled; medulla and cortex not pigmented; shrubby to subpendant;<br />
isidiate or isidiate-sorediate; stictic acid agg. (or in one collection from Hamilton<br />
County galbinic acid agg.); south to Lake County .............................................................................................. U. rubicunda Stirton<br />
1. Cortex, medulla or axis not red or pink ................................................................................................................................................... 7<br />
7. Soredia and/or isidia present .............................................................................................................................................................. 8<br />
8. Thallus shruby or subpendant.......................................................................................................................................................... 9<br />
9. Medulla loose .............................................................................................................................................................................. 10<br />
10. Thallus not inflated; salazinic acid ....................................................................................................................................... 11<br />
11. Isidiate-sorediate; cortex with numerous raised pseudocyphellae;<br />
Lake and Leon counties ...........................................................................................................................Usnea sp. 23493<br />
11. Isidiate; cortex smooth and shiny; Dade County ............................................................................ Usnea sp. Small 5853<br />
10. Thallus inflated and somewhat articulate; isidiate-sorediate;<br />
protocetraric acid; Lake County ...................................................................................................................Usnea sp. 23508<br />
9. Medulla compact; isidiate-sorediate; protocetraric acid; south to<br />
Lake County................................................................................................................................ U. subscabrosa Nyl. ex Motyka<br />
8. Thallus pendant, often with numerous perpendicular branchlets;<br />
galbinic acid agg.; south to Polk County......................................................................................U. dimorpha (Müll. Arg.) Motyka<br />
7. Soredia and isidia absent; thallus long, pendent; axis brownish; diffractaic acid<br />
and/or constictic acid; south to Polk County............................................................................................................U. trichodea Ach.<br />
VULPICIDA Mattson & Lai<br />
Only a single species, V. viridis (Schweinitz) Mattson & Lai in Florida. It was included in Tuckermanopsis in the previous<br />
edition. See key to genera.<br />
XANTHOPARMELIA (Vainio) Hale<br />
Since non-calcareous rock is almost entirely lacking in Florida, it is only with recent access to the Rock Hill Preserve that<br />
this genus has been added to the lichenota. Only X. conspersa (Ach.) Hale is known from the state. Many thanks to Angus<br />
Gholson for guiding Bill Buck and myself around the site.<br />
PHYSCIACEAE Zahlbr.<br />
KEY TO THE GENERA<br />
1. Thallus foliose.......................................................................................................................................................................................... 2<br />
41
2. Rhizines present.................................................................................................................................................................................. 3<br />
3. Upper cortex paraplectenchymatous; lower cortex present .......................................................................................................... 4<br />
4. Medulla white (bright red orange in Phaeophyscia rubropulchra), PD-; cortex UV-;<br />
apothecia with thalline margin..................................................................................................................................................... 5<br />
5. Cortex KOH+ yellow .....................................................................................................................................................Physcia<br />
5. Cortex KOH- ......................................................................................................................................................Phaeophyscia<br />
4. Medulla often orangish or yellowish, if white then cortex UV+ yellow<br />
or medulla PD+ orange; apothecia without thalline margin...............................................................................................Pyxine<br />
3. Upper cortex prosoplectenchymatous; lower cortex often lacking............................................................................Heterodermia<br />
2. Rhizines lacking or very sparse and poorly developed ...................................................................................................................... 6<br />
6. Divaricatic acid or sekikaic acid agg. present; underside black; lobes separable from<br />
the substrate; hypothecium brown .......................................................................................................................................Dirinaria<br />
6. No substances present; lobes entirely adnate to the substrate; hypothecium pale.................................................. Hyperphyscia<br />
1. Thallus crustose ...................................................................................................................................................................................... 7<br />
7. Ascospores with irregularly thickened walls ....................................................................................................................................... 8<br />
8. Ascospores small, under 30 µm long; hymenium not inspersed.............................................................................Rinodina s. lat.<br />
8. Ascospores larger, over 30 µm long; hymenium inspersed ................................................................................................ Hafellia<br />
7. Ascospores with uniformly thin walls or somewhat thickened only at<br />
septum (Buellia imshaugiana)............................................................................................................................................................. 9<br />
9. Conidiospores oblong or rod shaped......................................................................................................................................Buellia<br />
9. Conidiospores filiform, curved.........................................................................................................................................Amandinea<br />
BUELLIA De Notaris<br />
including AMANDINEA Scheid. & Mayrh. & HAFELLIA Kalb et al.<br />
1. Thallus not sorediate ............................................................................................................................................................................... 2<br />
2. Thallus PD- or PD+ yellow (norstictic or baeomycesic acid).............................................................................................................. 3<br />
3. Ascospores 8/ascus ........................................................................................................................................................................ 4<br />
4. Ascospores 15 µm or more long and 5 µm or more wide........................................................................................................... 5<br />
5. Apothecial disk naked ............................................................................................................................................................. 6<br />
6. Ascospores 2-celled ............................................................................................................................................................ 7<br />
7. Norstictic acid present; ascospore wall not or weakly ornamented ............................................................................... 8<br />
8. Hymenium without oil droplets or rarely with some oil in upper part.......................................................................... 9<br />
9. Ascospore wall thickened at septum; ascospores<br />
18-22 × 7-8.5(-10) µm; common, throughout Florida................................................ B. imshaugiana R. C. Harris<br />
9. Ascospore wall not thickened at septum; ascospores<br />
16-23 × 6-9 µm; common, south to Polk County..........................................................B. curtisii (Tuck.) Imshaug<br />
8. Hymenium with numerous oil droplets throughout ................................................................................................... 10<br />
10. Ascospores 14-22 × 6-8 µm ................................................................................................................................ 11<br />
11. Hymenium greenish above, KOH+ purplish; common,<br />
throughout Florida ..................................................................................... Hafellia bahiana (Malme) Sheard<br />
11. Hymenium brownish above, KOH-; throughout Florida.................................................. B. curatellae Malme<br />
42
10. Ascospores 22-41 × 11-21 µm............................................................................................................................ 12<br />
12. Ascospores 22-31 × 11-13 µm, darkly pigmented, wall pitted;<br />
northern Florida ..............................................................Hafellia callispora (Knight) H. Mayrh. & Sheard<br />
12. Ascospores 30-41 × 16-21 µm, lightly pigmented;<br />
wall not pitted; south to Sarasota County ...................................................... Hafellia parastata (Nyl.) Kalb<br />
7. Norstictic acid lacking; ascospore wall strongly ornamented;<br />
ascospores 14-20 × 8-11 µm; peninsular Florida .......................................................................B. leucomela Imshaug<br />
6. Ascospores becoming 4-celled, 17-21 × 6-8 µm; norstictic acid;<br />
north to Seminole County..................................................................................................B. lauricassiae (Fée) Müll. Arg.<br />
5. Apothecial disk yellow-green pruinose; thallus yellowish (xanthones);<br />
ascospores 19-28 × 8-10 µm; reported by Imshaug (1955) but<br />
not seen ................................................................................................................. B. melanochlora (Krempelh.) Müll. Arg .<br />
4. Ascospores 15 µm or less long and mostly 5 µm or less wide................................................................................................. 13<br />
13. Thallus not pigmented......................................................................................................................................................... 14<br />
14. Hymenium without oil droplets ...................................................................................................................................... 15<br />
15. Exciple at most paler brown within.......................................................................................................................... 16<br />
16. Lichen substances present (xanthones, atranorin, baeomycesic,<br />
barbatic, norstictic acids).................................................................................................................................... 17<br />
17. Containing other substances; exciple not paler within ................................................................................ 18<br />
18. Thallus PD-............................................................................................................................................. 19<br />
19. Thallus UV+ (lichexanthone and barbatic acid);<br />
disk not pruinose; ascospores 10-12.5 × 4-5.5 µm;<br />
throughout Florida .................................................................................... B. catasema (Tuck.) Tuck.<br />
19. Thallus UV-, KC+ orange (xanthones); disk white<br />
pruinose; ascospores 12-15 × 4.5-6 µm; Franklin<br />
and Seminole counties..........................................................................B. pachnidisca R. C. Harris<br />
18. Thallus PD+ yellow (baeomycesic and squamatic acids)<br />
(see below) ............................................................................................................. B. caloosensis Tuck.<br />
17. Atranorin and norstictic acid or atranorin only; exciple paler<br />
within; ascospores 10-15 × 5-7 µm; common, throughout<br />
Florida.......................................................................................................................... B. stillingiana Steiner<br />
16. Lichen substances lacking; tips of paraphyses swollen,<br />
dark-capped; ascospores 9-16 × 6-7 µm; throughout<br />
Florida ............................................................................... Amandinea punctata (Hoffm.) Coppins & Scheid.<br />
15. Exciple with a strongly differentiated colorless layer; ascospores<br />
9-12 × 4-5 µm; south to Lake County........................................................B. amphidexia Imshaug ex R. C. Harris<br />
14. Hymenium with abundant oil droplets; ascospores 9-13 × 4-5 µm;<br />
atranorin; Duval, Franklin, Lake and Seminole counties.........................................B. rappii Imshaug ex R. C. Harris<br />
13. Thallus containing bright pink or red pigment..................................................................................................................... 20<br />
20. Thallus thick, bullate, shiny, PD-; pigment red (xanthones and<br />
chiodectonic acid); medulla ascospores 8.5-12 × 4-5.5 µm; Dade<br />
and Liberty counties .............................................................................................................. B. coccinea (Fée) Aptroot<br />
20. Thallus thin, weakly areolate, PD+ yellow (baeomycesic and<br />
squamatic acids); pigment pink; ascospores 9-13 × 4-5.5 µm;<br />
Duval, Lake, Volusia and Wakulla counties ................................................................................. B. caloosensis Tuck.<br />
3. Ascospores more than 8/ascus..................................................................................................................................................... 21<br />
43
21. Hymenium with abundant oil droplets; ascospores 12-16/ascus;<br />
upper hymenium KOH+ purplish; south to Polk<br />
County............................................................................Hafellia bahiana (Malme) Sheard var. pleiotropa (Malme) Sheard<br />
21. Hymenium without oil droplets; ascospores 24-32/ascus; upper<br />
hymenium KOH-; south to Polk County.....................................................................Amandinea polyspora (Willey) ined.<br />
2. Thallus PD+ orange-red (fumarprotocetraric acid); ascospores 13.5-16.5 × 6.5-7.5 µm;<br />
Franklin, Levy, Liberty, Santa Rosa and Wakulla counties .................................................................... B. rubifaciens R. C. Harris<br />
1. Thallus with yellow soredia (KC+ orange) in punctiform to diffuse, confluent soralia;<br />
ascospore wall strongly ornamented; ascospores 16-19 × 9-11.5 µm; south to Polk County........................ B. wheeleri R. C. Harris<br />
DIRINARIA (Tuck.) Clements<br />
1. Thallus not sorediate, pustulate or isidiate ............................................................................................................................................. 2<br />
2. Apothecial disk with white pruina or pruina lacking ............................................................................................................................ 3<br />
3. Sekikaic acid agg. present; throughout Florida ................................................................................................D. confusa Awasthi<br />
3. Divaricatic acid present; south to Polk County ..................................................................................... D. confluens (Fr.) Awasthi<br />
2. Apothecial disk with purple pruina; divaricatic acid; throughout Florida<br />
......................................................................................................................................................... D. purpurascens (Vainio) Moore<br />
1. Thallus sorediate, pustulate or isidiate ................................................................................................................................................... 4<br />
4. Medulla white....................................................................................................................................................................................... 5<br />
5. Sorediate, pustulate or isidiate-pustulate ....................................................................................................................................... 6<br />
6. Sorediate, soralia mostly orbicular, eroded to capitate; soredia farinose ................................................................................. 7<br />
7. Lobes plane, not confluent, irregularly branched; apices truncate; thalli mostly<br />
smaller than next; divaricatic acid; throughout Florida .................................................. D. picta (Swartz) Clements & Shear<br />
7. Lobes often convex, confluent; apices flabellate;<br />
divaricatic acid; throughout Florida ................................................................................................D. applanata (Fée) Awasthi<br />
6. Pustulate or isidiate-pustulate; divaricatic acid; throughout Florida...................................................D. aegialita (Ach.) Moore<br />
5. Isidiate; isidia subglobose to cylindrical, not forming pustules or soredia<br />
divaricatic acid; Citrus and Okeechobee counties.......................................................................... D. papillulifera (Nyl.) Awasthi<br />
4. Medulla red; divaricatic acid; Levy and Seminole counties....................................................................D. leopoldii (Stein) Awasthi<br />
HAFELLIA Kalb, Mayrhofer & Scheidegger<br />
The treatment follows Sheard (Bryologist 95: 79-87. 1992) although I am somewhat uncomfortable about including Buellia bahaiana<br />
in Hafellia. The three species and one variety known from Florida are included in the key to Buellia above.<br />
HETERODERMIA Trevisan<br />
1. Thallus lobes shorter, not strap-like, mostly closely adnate to substrate............................................................................................... 2<br />
2. Lobes sorediate or isidiate; apothecia rare......................................................................................................................................... 3<br />
3. Sorediate .......................................................................................................................................................................................... 4<br />
4. Underside corticate ...................................................................................................................................................................... 5<br />
5. Salazinic acid and unknown PQ4 present; common, north<br />
of the Everglades.........................................................................................................H. albicans (Pers.) Swinscow & Krog<br />
5. Salazinic acid and unknown PQ4 absent; common, south<br />
to Highlands County ................................................................................................................H. speciosa (Wulfen) Trevisan<br />
4. Underside not corticate ................................................................................................................................................................ 6<br />
44
6. Underside white or purplish gray in the center, KOH- ............................................................................................................. 7<br />
7. Thallus coarse with coarse marginal soredia; rhizines dark................................................................................................. 8<br />
8. Underside white or weakly purplish toward the center,<br />
not yellowish toward tips; norstictic acid rarely present;<br />
south to Lake Okeechobee ..................................................................................H. japonica (Sato) Swinscow & Krog<br />
8. Underside purplish, yellowish toward tips, ultimate parts<br />
white; norstictic acid present; Marion County ...........................................................H. casarettiana (Massal.) Trevisan<br />
7. Thallus delicate with fine soredia in terminal, somewhat labrose, soralia;<br />
rhizines pale; norstictic acid lacking in N. American material;<br />
Lake and Marion counties ..............................................................................................H. galactophylla (Tuck.) W. Culb.<br />
6. Underside orange, KOH+ purple; south to Lake Okeechobee ................................................. H. obscurata (Nyl.) Trevisan<br />
3. Isidiate; underside orange, KOH+ purple; Leon, Marion and Okaloosa counties ....................................... H. crocea R. C. Harris<br />
2. Lobes neither sorediate nor isidiate; apothecia present...................................................................................................................... 9<br />
9. Rhizines simple; norstictic acid absent; Panhandle ......................................................................... H. echinata (Taylor) W. Culb.<br />
9. Rhizines branched; norstictic acid usually present; reported by Kurokawa<br />
from Dade County, not seen by me .................................................................................................H. barbifera (Nyl.) K. P. Singh<br />
1. Thallus lobes narrow, long, strap-like, subfruticose, loosely attached to substrate;<br />
tips of lobes often sorediate below; with or without salazinic acid; Marion County<br />
and south, reported by Moore, not seen .......................................................................................................... H. leucomelos (L.) Poelt<br />
HYPERPHYSCIA Müll. Arg.<br />
1. Thallus sorediate ..................................................................................................................................................................................... 2<br />
2. Lower surface pale, conspicuous black hypothallus lacking;<br />
soralia marginal; south to Polk County ..................................................................................H. adglutinata (Flörke) Mayrh. & Poelt<br />
2. Lower surface dark, conspicuous black hypothallus present;<br />
soralia laminal; Dade County ..........................................................................................................................H. minor (Fée) Awasthi<br />
1. Thallus esorediate; Duval and Levy counties, reported by Moore, not seen ......................................H. syncolla (Tuck. ex Nyl.) Kalb<br />
PHAEOPHYSCIA Moberg<br />
1. Thallus sorediate; apothecia usually lacking .......................................................................................................................................... 2<br />
2. Medulla white........................................................................................................................................................................................ 3<br />
3. Tips of lobes naked .......................................................................................................................................................................... 4<br />
4. Soralia capitate, borne on raised, lateral lobes; underside dark; Panhandle<br />
(fide Esslinger, not seen by me).....................................................................................................P. pusilloides (Zahlbr.) Essl.<br />
4. Soralia marginal; underside pale; Baker County (fide Esslinger, not<br />
seen by me) ..................................................................................................................................... P. chloantha (Ach.) Moberg<br />
3. Tips of lobes with erect, colorless hairs; St. Johns County ............................................................. P. cernohorskyi (Nádv.) Essl.<br />
2. Medulla orange-red; Panhandle fide Esslinger, not seen by me ....................................................... P. rubropulchra (Degel.) Essl.<br />
1. Thallus not sorediate; apothecia present; Panhandle .................................................................................. P. ciliata (Hoffm.) Moberg<br />
PHYSCIA (Schreber) Michaux<br />
1. Thallus not sorediate ............................................................................................................................................................................... 2<br />
2. Lobes white spotted, including lobe ends; zeorin present or absent; ascospores Physcia-<br />
or Pachysporaria-type .......................................................................................................................................................................... 3<br />
45
3. Spores Physcia-type; zeorin present ................................................................................................................................................ 4<br />
4. Lobes narrow, often convex, not confluent, branching irregular; apothecial<br />
disk not or only weakly pruinose; margin thin, smooth, not raised;<br />
ascospores small, 15-17 × 6-7.5 µm; Calhoun, Dixie, Gadsden<br />
and Hamilton counties..............................................................................................................................P. pumilior R. C. Harris<br />
4. Lobes broader, flatter, often confluent, branching more nearly dichotomous;<br />
apothecial disk normally strongly pruinose; margin thick, often becoming<br />
crenulate, usually strongly raised; ascospores larger, 18-25 × 8-12 µm;<br />
mostly north of Mason-Dixon line..........................................................................................[P. aipolia (Ehrh. ex Hampe) Fürnr.]<br />
3. Spores Pachysporaria-type, 19-20 × 8-9 µm; zeorin absent or trace;<br />
Dade County and Florida Keys ................................................................................................................... Physcia sp. Small 7569<br />
2. Lobes not white spotted or if white spotted then only in the older parts,<br />
not on lobe ends; zeorin absent; apothecia with rhizines on back; ascospores<br />
Pachysporaria-type, 20-25 × 8-11 µm; throughout Florida............................................................................ P. neogaea R. C. Harris<br />
1. Thallus sorediate ..................................................................................................................................................................................... 5<br />
5. Soralia mainly marginal....................................................................................................................................................................... 6<br />
6. Underside dark or pale, not striate; lower cortex paraplectenchymatous ..................................................................................... 7<br />
7. Underside pale; soralia marginal, not pustular ............................................................................................................................ 8<br />
8. Medulla KOH-; zeorin absent; thallus not "frosted", thin and fragile;<br />
Lake County, "SW Florida" and Louisiana...........................................................................................................P. crispa Nyl.<br />
8. Medulla KOH+ yellow; zeorin present; thallus "frosted", thicker;<br />
margins often undulate; Collier and Hillsborough counties ..............................................................P. undulata Moberg<br />
7. Underside dark; soralia marginal, often pustular; soredia coarse;<br />
zeorin present; throughout Florida ..................................................................................................P. sorediosa (Vainio) Lynge<br />
6. Underside pale at tips with dark longitudinal lines; lower cortex more or less<br />
prosoplectenchymatous; lobe margins pruinose especially at tips; common,<br />
throughout Florida ..........................................................................................................................................P. atrostriata Moberg<br />
5. Soralia mainly laminal; underside pale ............................................................................................................................................... 9<br />
9. Lower cortex prosoplectenchymatous; zeorin absent, unknown terpenoid present;<br />
northern Florida .................................................................................................................................................P. americana Merr.<br />
9. Lower cortex paraplectenchymatous; zeorin present; Columbia and Dade counties................................ P. poncinsii Hue<br />
NOTES<br />
Physcia poncinsii Hue<br />
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,<br />
29°51'N, 82°39'W, on Berchemia scandens, 5 Dec 1994, Harris 36037 (NY); Dade County: Coral Gables, Fairchild Tropical Garden,<br />
4 Nov 1982, Tucker 25266 (LSU).<br />
Physcia undulata Moberg<br />
New to North America. FLORIDA. Collier County: Fakahatchee Strand State Preserve, Royal Palm Hammock, on Ilex, 6 Dec<br />
1992, Harris 29942; Hillsborough County: Hillsborough River State Park, on Carya, 3 Dec 1992, Harris 29691. LOUISIANA. West<br />
Feliciana Parish: Locust Grove Cemetery, ca. 4 mi from Bains, on grave stone, 13 Sep 1975, Buck B596 (all NY).<br />
PYXINE Fr.<br />
1. Thallus UV-; medulla PD+ orange or PD-............................................................................................................................................... 2<br />
2. Medulla white or tinted yellow or orange, PD+ orange (testacein agg.) ............................................................................................. 3<br />
3. Lobes with marginal dactyls (isidiate/pustulate) ................................................................................................................................ 4<br />
4. Dactyls fragile, almost immediately breaking down into granular soredia;<br />
46
PD+ dirty orangish; ascospores 4-celled; throughout Florida ....................................................... P. eschweileri (Tuck.) Vainio<br />
4. Dactyls coarse, not becoming sorediate (may be broken in handling);<br />
PD+ red orange; ascospores 2-celled; Walton County............................................................................... P. coralligera Malme<br />
3. Lobes with laminal soralia; ascospores 2-celled; Leon, Marion and<br />
Walton counties......................................................................................................................... P. eschweileri (Tuck.) Vainio s. lat.<br />
2. Medulla yellow-orange, PD-; soredia marginal; not yet definitely known from Florida ...............................[P. sorediata (Ach.) Mont.]<br />
1. Thallus UV+ yellow (lichexanthone); medulla PD- ................................................................................................................................. 5<br />
5. Soredia or dactyls present................................................................................................................................................................... 6<br />
6. Medulla white or pale yellow, KOH+ yellow or KOH-...................................................................................................................... 7<br />
7. Medulla white; Peninsular Florida......................................................................................................... P. cocoes (Swartz) Nyl.<br />
7. Medulla yellow; Alachua, Dixie and Hamilton counties ............................................................................P. subcinerea Stirton<br />
6. Medulla orange to pale salmon, KOH+ purple ................................................................................................................................ 8<br />
8. Lobes with laminal soralia; throughout Florida ............................................................P. caesiopruinosa (Nyl.) Imshaug s. lat.<br />
8. Lobes with marginal dactyls; south to Seminole County............................................ P. caesiopruinosa (Nyl.) Imshaug s. str.<br />
5. Soredia and dactyls lacking; medulla yellow; Lake Okeechobee southward....................................... P. berteriana (Fée) Imshaug<br />
NOTES<br />
Pyxine caesiopruinosa (Nyl ) Imshaug<br />
As noted in 1991 there are two taxa included under this name which are readily separable by their asexual reproductive structures<br />
and terpene patterns. I have now had a chance to examine Nylander's type. Although fragmentary it is clearly the taxon with marginal<br />
dactyls. It still remains to examine the type of<br />
P. albovirens (G. Meyer) Aptroot. Aptroot's illustration (not type) of P. albovirens is clearly the laminal soraliate taxon which is also in the<br />
West Indies but I see no point in taking up albovirens until the correct application of the name is determined.<br />
RINODINA (Ach.) S. Gray<br />
1. Thallus white, gray or brownish............................................................................................................................................................... 2<br />
2. Thallus gray or brownish; apothecial margin often poorly developed; no lichen substances ........................................................... 3<br />
3. Thallus lacking isidioid granules ..................................................................................................................................................... 4<br />
4. Apothecia sessile, sometimes constricted below; thallus usually somewhat<br />
areolate, often poorly developed, not shiny................................................................................................................................ 5<br />
5. Ascospore lumina not constricted in the middle; apothecia with at least<br />
remnants of thalline margin...................................................................................................................................................... 6<br />
6. Ascospore lumina globose to slightly longer than broad;<br />
ascospores 16-18 × 7-8 µm; south to Polk County..........................................................................R. applanata H. Magn.<br />
6. Ascospore lumina distinctly longer than broad; ascospores 15-18 × 6-7.5 µm;<br />
Leon and Volusia counties.................................................................................................................. Rinodina sp. 23268<br />
5. Ascospore lumina elongated, constricted in the middle, bowling pin-<br />
or dumbell-shaped; ascospores 17-20 × 7.5-9 µm; apothecia without<br />
thalline margin; Sarasota County......................................................................................... Buellia placodiomorpha Vainio<br />
4. Apothecia immersed in continuous, smooth, shiny thallus; ascospores<br />
with subglobose lumina, 15-19 × 9-10 µm; Dade County............................................................... Rinodina sp. Tucker 25286<br />
3. Thallus tan, with isidioid granules; ascospores with globose lumina,<br />
16-19 × 9-10 µm; Polk and Dade counties ......................................................................................... Rinodina sp. Tucker 25528<br />
2. Thallus and apothecial margin white or pale gray, well developed,<br />
resembling a Lecanora; ascospores Physcia-type, 18-20 × 9-11 µm;<br />
atranorin, zeorin; Duval, Lake and Seminole counties.................................................................................R. granuligera H. Magn.<br />
47
1. Thallus and apothecial margin greenish yellow; ascospores Pachysporaria-type,<br />
20-27 × 11-13 µm; central Florida ...................................................................................................................R. lepida (Nyl.) Müll. Arg.<br />
PERTUSARIALES<br />
PERTUSARIACEAE Körber<br />
OCHROLECHIA Massal.<br />
1. Thallus not pustulate-isidiate, smooth to rugose, rugae often vermiform;<br />
apothecial cortex C-, medulla C+ red (gyrophoric acid with accessory<br />
lichexanthone and methylhiascic acids); throughout the state...................................................................................O. africana Vainio<br />
1. Thallus pustulate-isidiate; pustules and thalline rugae very irregular, occasionally<br />
breaking down into a granular sorediose crust; apothecia mostly without<br />
cortex; apothecial medulla C+ red, (gyrophoric acid and 5-O-methylhiascic acid);<br />
Liberty and Marion counties ................................................................................................................................... O. antillarum Brodo<br />
PERTUSARIA DC.<br />
1. Ascomata discoid, lecanorate, often heavily pruinose or sorediate above or sterile and<br />
soraliate; ascospores l-2/ascus when present [if photobiont is Trentepohlia or chemistry<br />
is not one of those below, these will be treated in key to sterile sorediate crusts]................................................................................ 2<br />
2. Thallus UV+ yellow (lichexanthone)..................................................................................................................................................... 3<br />
3A. Lecanoric acid; wart C+ red; often fertile; south to Lake County....................................................................P. pulchella Malme<br />
3B. Hypothamnolic acid; wart KOH+ lavender, KC+ red, PD-;<br />
sometimes fertile; Duval, Liberty and Walton counties ......................................................................P. hypothamnolica Dibben<br />
3C. Picrolichenic acid; wart KOH-, C-, KC+ violet, PD-;<br />
sterile; Putnam and Wakulla counties ...............................................................................................................P. ventosa Malme<br />
3D. Baeomycesic acid, unknowns, squamatic acid (tr.);<br />
wart KOH-, C-, KC-, PD+ yellow; often fertile;<br />
south to Lee County ........................................................................................................................................ P. floridana Dibben<br />
3E. Haemathamnolic acid; wart KOH+ yellow becoming brownish,<br />
C-, KC-, PD+ orange; rarely fertile; throughout the state ..............................................................................P. copiosa Erichsen<br />
3F. Thamnolic acid; wart KOH+ yellow, C-, KC-, PD+ orange;<br />
south to Lake County.....................................................................................................................................Pertusaria sp. 25229<br />
3G. Stictic acid; wart KOH+ yellow, C-, KC-, PD+ pink-orange;<br />
Marion County .....................................................................................................................................Pertusaria sp. Buck 16889<br />
2. Thallus UV- or UV+ pinkish [if appropriate chemistry is not found, specimen is either a<br />
lichexanthone deficient form of the above group or not a Pertusaria] ............................................................................................... 4<br />
4. Thallus UV-, cortical xanthone lacking; usually fertile ..................................................................................................................... 5<br />
5A. Lecanoric acid; wart C+ red; south to Orange County........................................................................... P. velata (Turner) Nyl.<br />
5B. Picrolichenic acid; wart KOH-, C-, KC+ violet, PD-; Duval County.......................................................... P. amara (Ach.) Nyl.<br />
5C. Thamnolic acid; wart KOH+ yellow, C-, KC-, PD+ orange;<br />
Volusia County..................................................................................................................................P. trachythallina Erichsen<br />
5D. Fumarprotocetraric acid; wart KOH-, C-, KC-, PD+<br />
orange-red; Leon and Wakulla counties........................................................................................ P. multipunctoides Dibben<br />
4. Thallus UV+ pinkish (coronatone); stictic acid agg.; wart KOH+ yellow, C-, KC-,<br />
PD+ pink-orange; sterile; Duval, Highlands, Liberty and Nassau counties ............................................... P. expolita R. C. Harris<br />
1. Apothecia with small pore-like disk......................................................................................................................................................... 6<br />
6. Ascospores 8/ascus ............................................................................................................................................................................. 7<br />
7. Cortex UV+ orange, pink or UV- ...................................................................................................................................................... 8<br />
8. Cortex UV+ orange (thiophaninic acid)......................................................................................................................................... 9<br />
48
9. Medulla PD+ orange ............................................................................................................................................................... 10<br />
10. Stictic acid agg. present; ostioles often yellow and raised;<br />
ascospores 45-75 × 21-33 µm; throughout the state.............................................................................P. texana Müll. Arg.<br />
10. Virensic and an unknown (PD-) present; ostioles not yellow and raised;<br />
ascospores 60-70 × 28-35 µm; Highlands and Okaloosa counties.............................................. P. virensica R. C. Harris<br />
9. Medulla PD-............................................................................................................................................................................. 11<br />
11. Containing variolaric acid; ascospores basally biseriate to uniseriate,<br />
65-90 × 30-40 µm; ostiolar area often depressed and yellow;<br />
Duval, Leon and Liberty counties.................................................................................................P. epixantha R. C. Harris<br />
11. Containing unknown substance (perlatolic/confluentic acid group);<br />
ascospores uniseriate, 55-60 × 25-28 µm; ostiolar area not yellow;<br />
Duval County .......................................................................................................................................Pertusaria sp. 23902<br />
8. Cortex UV+ pinkish (coronatone) or UV- .................................................................................................................................... 12<br />
12. Cortex UV-; medulla PD- (unknown substance); ascospores uniseriate, rough,<br />
70-95 × 30-40 µm; south to Highlands County ........................................................................................ P. iners R. C. Harris<br />
12. Cortex UV+ pinkish (coronatone); medulla PD+ pink-orange (stictic acid agg.);<br />
apothecial warts very irregular; ascospores uniseriate, smooth,<br />
50-60 × 27-30 µm; Lake County.............................................................................................................Pertusaria sp. 23573<br />
7. Cortex UV+ yellow (lichexanthone); medulla K-, P- (4-O-methylperlatolic acid);<br />
ascospores uniseriate, 55-100 × 28-40 µm; south to Marion County.................................................P. paratuberculifera Dibben<br />
6. Ascospores basically 2 or 4/ascus (1-5) ............................................................................................................................................... 13<br />
13. Ascospores 4/ascus ......................................................................................................................................................................... 14<br />
14. Cortex UV+ pinkish or UV-.......................................................................................................................................................... 15<br />
15. Medulla PD+ pink-orange (stictic acid agg.); hymenium not inspersed................................................................................ 16<br />
16. Ascospores smooth, 50-100 × 23-43 µm; Duval County (reported by Dibben<br />
from Lee and Seminole counties also) ............................................................................. P. leucostoma (Bernh.) Massal.<br />
16. Ascospores rough, 75-130 × 27-43 µm; throughout the state...................................................P. tetrathalamia (Fée) Nyl.<br />
15. Medulla PD- (traces of unknown); hymenium densely inspersed;<br />
ascospores smooth, 63-100 × 30-45 µm; throughout the state...................................................... P. sinusmexicani Dibben<br />
14. Cortex UV+ orange (xanthone X-l); medulla PD+ pink-orange (stictic acid agg.);<br />
ascospores smooth, 90-105 × 30-35 µm; St. Lucie and Pinellas counties ........................................ Pertusaria sp. Britton 829<br />
13. Ascospores 2/ascus ......................................................................................................................................................................... 17<br />
17. Medulla PD+ orange .................................................................................................................................................................... 18<br />
18. Medulla PD+ pink-orange (stictic acid agg.).......................................................................................................................... 19<br />
19. Ascospores rough, 55-125 × 28-45 µm; ostiole pale; cortical xanthone<br />
thiophaninic acid; throughout the state ......................................................................................... P. xanthodes Müll. Arg.<br />
19. Ascospores smooth, 65-135 × 27-40 µm; ostiole dark; cortical xanthone,<br />
xanthone X-l; reported by Dibben from Duval and Seminole counties .......................................P. pustulata (Ach.) Duby<br />
18. Medulla PD+ orange-red (fumarprotocetraric acid); cortex UV-<br />
(unknown xanthone); ascospores somewhat roughened,<br />
75-135 × 30-45 µm; Lake and Marion counties .....................................................................................P. subpertusa Brodo<br />
17. Medulla PD- (variolaric acid); cortex UV- (no xanthone); apothecia mostly<br />
immersed; ascospores somewhat roughened, 100-120 × 30-40 µm; Marion<br />
and Wakulla counties...................................................................................................................................P. obruta R. C. Harris<br />
49
<strong>PART</strong> 2<br />
THE 10¢ TOUR OF THE PYRENOLICHENS<br />
A TWENTY-YEAR PERSPECTIVE<br />
In 1975 I submitted my thesis which, while dealing primarily with Arthopyrenia s. lat., also included a<br />
primitive attempt to break away from Zahlbruckner's system, then in use. Over the ensuing 20 years my<br />
mistakes as regards Arthopyrenia and the more general system have become evident, new ideas and<br />
information have become available, critical specimens have been examined and a large number of new<br />
records have accumulated. This anniversary seems an appropriate time to try again. The system proposed is<br />
not intended in any way to be final but only an interim report. As noted in the introduction the title "tour" is<br />
given to this part so that I am free to pick and choose what is treated. This is not the proverbial 50¢ tour, only<br />
the 10¢ version, since I badly miscalculated the amount of errors, changes and specimens to be waded<br />
through. The most interesting part, the higher categories, is left undone. They naturally fall at the end of such<br />
a project since I have worked from the bottom (species) upwards. I do not like the current tendency to assign<br />
new genera, etc., to "incertae sedis". This conveys absolutely no information. It is useless. It is an admission<br />
of failure, of lack of general knowledge, of laziness, which one is surely forced to admit to on occasion but<br />
should not become a way of life. Given a choice, I have chosen to err on the side of controversy. This, I<br />
50
hope, leads to more intensive study and ultimately to a better result. Nothing is set in stone. I do not consider<br />
some of the groups treated here to be lichens. Not only do I feel there is no consistent association with a<br />
photobiont, they fail the basic definition that requires some visible morphological evidence of interaction<br />
between fungus and photobiont. I have omitted for now perithecoid forms belonging to the Gyalectales s. lat.<br />
and Lecanorales. Also the non-lichen Mycoglaena, Micropeltidaceae, is omitted. The following table shows<br />
the changes over 20 years in the systematic arrangement of the corticolous pyrenolichens. Due to time<br />
constraints I have not been able to work through the mostly saxicolous Verrucariaceae nor the perithecioid<br />
genera of parasites/ parasymbionts, some of which probably are confamilial with lichen-forming taxa.<br />
ZAHLBRUCKNER (1926)<br />
PYRENULACEAE<br />
Anthracothecium<br />
Arthopyreneiella<br />
Arthopyrenia<br />
Belonia<br />
Blastodesmia<br />
Clathroporina<br />
Coccotrema<br />
Leptorhaphis<br />
Microthelia<br />
Monoblastia<br />
Polyblastiopsis<br />
Porina<br />
Pseudopyrenula<br />
Pyrenula<br />
Stereochlamys<br />
Thelopsis<br />
TRYPETHELIACEAE<br />
Bottaria<br />
Laurera<br />
Melanotheca<br />
Tomasellia<br />
Trypethelium<br />
HARRIS (1974)<br />
PLEOSPORACEAE<br />
Arthopyrenia<br />
Blastodesmia<br />
Leptorhaphis<br />
Microthelia<br />
Mycomicrothelia?<br />
Mycoglaena?<br />
Mycoporum<br />
Polyblastiopsis<br />
Sporoschizon<br />
Tomasellia<br />
PYRENULACEAE<br />
Anthracothecium<br />
Bottaria<br />
Eopyrenula<br />
Lithothelium<br />
Melanotheca<br />
Microglaena s. str.<br />
Parathelium<br />
Parmentaria<br />
Plagiocarpa<br />
Pleurotheliopsis<br />
Pyrenastrum<br />
PARATHELIACEAE<br />
Campylothelium<br />
Parathelium<br />
Plagiotrema<br />
Pleurotrema<br />
HARRIS (1995)<br />
LOCULOASCOMYCETES<br />
PLEOSPORALES<br />
NAETROCYMBACEAE<br />
Jarxia<br />
Leptorhaphis<br />
Naetrocymbe<br />
Tomasellia<br />
MICROPELTIDACEAE<br />
Mycoglaena<br />
MYCOPORACEAE<br />
Mycoporum<br />
ARTHOPYRENIACEAE<br />
Arthopyrenia<br />
Julella<br />
Mycomicrothelia<br />
XANTHOPYRENIACEAE<br />
Pyrenocollema<br />
Zwackhiomyces<br />
ZAHLBRUCKNER (1926)<br />
Pleurothelium<br />
ASTROTHELIACEAE<br />
Astrothelium<br />
Cryptothelium<br />
51<br />
Lithothelium<br />
Parmentaria<br />
Pyrenastrum<br />
STRIGULACEAE
Haplopyrenula<br />
Micropyrenula<br />
Microtheliopsis<br />
Phyllobathelium<br />
Phylloblastia<br />
Phylloporina<br />
Raciborskiella<br />
Strigula<br />
Trichothelium<br />
XANTHOPYRENIACEAE<br />
Xanthopyrenia<br />
MYCOPORACEAE<br />
Asteroporum<br />
Dermatina<br />
Mycoporellum<br />
HARRIS (1975)<br />
PYRENULACEAE<br />
Pyrenula<br />
STRIGULACEAE<br />
Acrocordia<br />
Anisomeridium<br />
Monoblastia<br />
Pleurotrema<br />
Pyrenocollema<br />
Strigula<br />
TRICHOTHELIACEAE<br />
Clathroporina<br />
Porina<br />
Trichothelium<br />
TRYPETHELIACEAE<br />
Astrothelium<br />
Campylothelium<br />
Cryptothelium<br />
Laurera<br />
Polymeridium<br />
Pseudopyrenula<br />
Trypethelium<br />
* Genera in square brackets probably doomed to synonymy.<br />
VERRUCARIALES<br />
52<br />
HARRIS (1995)<br />
DACAMPIACEAE<br />
Dacampia<br />
Eopyrenula<br />
MELANOMMATALES s. lat.<br />
THELENELLACEAE<br />
?Celothelium<br />
[Chromatochlamys]*<br />
Thelenella<br />
STRIGULACEAE<br />
Phyllobathelium<br />
Strigula<br />
PYRENULACEAE<br />
Acrocordiella<br />
Anthracothecium<br />
Clypeopyrenis<br />
Distopyrenis<br />
Granulopyrenis<br />
Lachrymospora<br />
Lithothelium<br />
Mazaediothecium<br />
Mycopyrenula<br />
Parapyrenis<br />
Polypyrenula<br />
Pyrenographa<br />
Pyrenowilmsia<br />
Pyrenula<br />
Pyrgillus<br />
TRYPETHELIACEAE<br />
Architrypethelium<br />
Astrothelium<br />
Bathelium<br />
[Campylothelium]<br />
[Cryptothelium]<br />
Exiliseptum<br />
Laurera<br />
Polymeridium<br />
Pseudopyrenula<br />
Trypethelium<br />
HYMENOASCOMYCETES<br />
TRICHOTHELIALES<br />
TRICHOTHELIACEAE<br />
Clathroporina<br />
Porina<br />
Segestria<br />
Trichothelium<br />
XYLARIALES<br />
PLEUROTREMATACEAE<br />
Pleurotrema, etc. (see Barr, 1994)<br />
I have not had time to deal with this order in a thorough fashion, treating only the rather few Florida<br />
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<br />
Mycosphaerellaceae should be transferred to the Microtheliopsidaceae that is more closely allied to<br />
Chaetothyriales than Verrucariales.<br />
VERRUCARIACEAE Zenker<br />
1. Thallus crustose, on rock; ascospores simple or 2-celled ........................................................................ 2<br />
2. Ascospores simple ................................................................................................................ Verrucaria<br />
2. Ascospores 2-celled............................................................................................................... Thelidium<br />
1. Thallus squamulose, on bark, mosses, lichens or rock; ascspores simple,<br />
transversely septate or muriform.............................................................................................................. 3<br />
3. Thallus of large to small, brown or tan, ± orbicular or irregular squamules,<br />
without a raised margin, not sorediate; on bark, soil or rock................................................................ 4<br />
4. Ascospores simple, 8/ascus; hymenial algae absent; on bark or soil........................ Catapyrenium<br />
4. Ascospores muriform, 2/ascus; hymenial algae present; on limestone<br />
or cement ...................................................................................................................... Endocarpon<br />
3. Thallus of tiny, green gray, orbicular or ± irregular squamules, usually with<br />
a raised margin or marginal soralia; over bark, mosses or other lichens;<br />
ascomata very rare; Dixie, Marion and Suwannee counties ...................................................................<br />
................................................................................................... Normandina pulchella (Borrer) Nyl.<br />
CATAPYRENIUM Flotow<br />
1. Growing on calcareous soil; thallus usually quite orbicular,<br />
not imbricate,brown; Gadsden County.................................................... C. squamulosum (Ach.) Breuß<br />
1. Growing on bark (occasionally on shaded limestone);<br />
thallus pale, tan or grayish, imbricate when well developed;<br />
Leon County .........................................................................................C. tuckermanii (Mont.) Thomson<br />
NOTES<br />
I thank Othmar Breuß for determining C. squamulosum. Both species known from Florida belong to the<br />
segregate genus Dermatocarpella Harada (1993). Based on my limited knowledge of this group, Harada's<br />
genera seem tenable but the European arbiters have not yet spoken. The combination for tuckermanii has not<br />
been made however Dermatocarpella squamulosa (Ach.) Harada is available. The genus just barely makes it<br />
into northern Florida. Harada divides Catapyrenium as follows.<br />
1. Exciple colorless........................................................................................................................................ 2<br />
2. Pycnidia of Xanthoria-type; lobes round; attached to substrate by<br />
rhizohyphae over almost the whole lower surface ...........................................Dermatocarpella Harada<br />
2. Pycnidia of Staurothele-type; lobes linear, canaliculate below,<br />
basally attached, almost stipitate; lower surface naked ................................. Neocatapyrenium Harada<br />
1. Exciple brown to almost black................................................................................................................... 3<br />
3. Pycnidia unknown; exciple brown; upper surface ± scabrose, dull;<br />
upper cortex paraplectenchymatous; on soil or bark ................................................Catapyrenium Ach.<br />
3. Pycnidia of the Staurothele-type; exciple almost black; upper<br />
53
surface smooth, ± glossy; upper cortex pachydermatous; on<br />
soil or rock ......................................................................................................... Scleropyrenium Harada<br />
ENDOCARPON Hedwig<br />
1. On mortar between bricks (HCl+); thallus brown, relatively undivided;<br />
ascospores 23-31 × 11-16 µm; Dade County.............................................. Endocarpon sp. Calkins 147<br />
1. On HCl- rock along river; thallus light gray, becoming lobate;<br />
ascospores 29-41 × 11-16 µm; Hillsborough County .................................. Endocarpon sp. Buck 22648<br />
NOTES<br />
Calkins 147 resembles some forms assigned to E. pusillum Hedwig but the upper cortex is composed of<br />
relatively thin walled hyphae. Buck 22648 is smaller and more divided than E. pusillum and also differs in thin<br />
walled cortical cells. There are several Cuban species but I have not seen adequate material. Endocarpon is<br />
in need of a careful revision in the Americas.<br />
NORMANDINA Nyl.<br />
Normandina pulchella (Borrer) Nyl. is probably undercollected because of its small size and is more<br />
common in Florida than the few specimens would indicate. Applying Occam's Razor I prefer not to recognize<br />
Lauderlindsaya J. David & D. Hawksw. (= ascomata of Normandina). Three similar<br />
squamulose/subsquamulose species with ascomata seems to me a simpler explanation than three similar<br />
species each with three similar parasymbionts. There is no "hard" data such as attempts to recombine<br />
ascospores with isolated algae or comparative molecular studies of ascospores and vegetative hyphae.<br />
THELIDIUM Massal.<br />
1. Thallus epilithic, ± continuous to areolate, greenish; ascomata globose,<br />
sessile, ca. 0.2 mm diam., wall brown-black; ascospores 2-celled,<br />
20-23 × (8-)9-11(-13.5) µm; on natural limestone and cement rubble;<br />
Clay, Collier and Taylor counties..................................................................... Thelidium sp. Buck 22736<br />
1. Thallus mostly endolithic, greenish?; ascomata immersed, subglobose,<br />
clypeate; exciple colorless; ascospores 2-celled, 17-20 × 7.5-8.5 µm;<br />
Monroe County (Key West) .............................................................................Thelidium sp. Thaxter 403<br />
NOTES<br />
Buck 22736 may be referable to T. minutulum Körber but comparative material was not available.<br />
VERRUCARIA Ach.<br />
1. Ascomata deeply immersed in limestone; black clypeus restricted to a disk<br />
or cap around ostiole; ascomatal wall colorless........................................................................................ 2<br />
2. Ascomata numerous, crowded; clypeus a small (ca. 0.2 mm) flat disk;<br />
thallus endolithic, white; all collections sterile with immature asci;<br />
periphyses very abundant and conspicuous; Collier, Dade, Monroe<br />
and Taylor counties ..................................................................................... Verrucaria sp. Britton 861<br />
54
2. Ascomata scattered; clypeus large (0.5-0.7 mm), flat to hemispherical,<br />
(depending on erosion of surrounding limestone), often with a distinct<br />
papilla; thallus endolithic, white; ascospores 17-22(-24) × 9-11 µm;<br />
Monroe County (Key West) ............................................................................. V. riddleana R. C. Harris<br />
1. Ascomata sessile or with base immersed in thallus or limestone; black<br />
clypeus hemispherical; ascomatal wall colorless or dark.......................................................................... 3<br />
3. Ascospores larger, 15-30 × (7-)9-16 µm; thallus endolithic or<br />
epilithic; ascomata 0.35-0.6 mm diam. ................................................................................................. 4<br />
4. Thallus white, endolithic or epilithic ................................................................................................... 5<br />
5. Exciple dark; clypeus not broadly spreading; ascomata<br />
0.4-0.5 mm diam.; ostiolar area depressed; ascospores<br />
21-30 × 11-14 µm; Dade and Monroe counties.................................. Verrucaria sp. Britton 870<br />
5. Exciple pale; clypeus spreading, 0.5-0.6 mm diam.;<br />
ascospores 2-28 × 10.5-14.5 µm; Dade and<br />
Monroe counties................................................................................ Verrucaria sp. Thaxter 397<br />
4. Thallus green gray or olive green to brown green............................................................................. 6<br />
6. Ascospores 15-23 × (7-)9-11 µm; thallus olive or brown<br />
green, areolate; ascomata 0.35-0.4 mm diam.; Taylor<br />
County ................................................................................................ Verrucaria sp. Buck 24842<br />
6. Ascospores 23-27 × 11-12.5 µm; thallus green gray, rimose<br />
or areolate; ascomata ca. 0.4 mm diam.; Taylor and<br />
Wakulla counties ................................................................................ Verrucaria sp. Buck 18979<br />
3. Ascospores small, 9-16 × 4-6.5 µm; thallus epilithic, tan to brown;<br />
ascomata 0.25-0.3 mm diam. ............................................................................................................... 7<br />
7. Ascospores 9-12 × (4-)5-5.5(-6.5) µm; exciple usually dark;<br />
Dade, Hillsborough and Monroe counties .............................................. Verrucaria sp. Buck 22655<br />
7. Ascospores 13-16 × 5.5-6.5 µm; exciple pale;<br />
Dade County............................................................................................ Verrucaria sp. Britton 741<br />
NOTES<br />
There is so little known about subtropical/tropical Verrucaria that I have not ventured to describe any new<br />
taxa. Only three species have Caribbean types and I have seen material of only one of these. One could<br />
shoehorn some of the Florida taxa into European species, i.e., Britton 870 into V. muralis Ach. or Thaxter 397<br />
into V. calkinsiana Servít, but in my present state of ignorance it doesn't seem worth it. This has obviously<br />
been the practice in the West Indies.<br />
Verrucaria riddleana R. C. Harris, nom. nov.<br />
Verrucaria mamillaris Riddle in Britton & Millspaugh, Bahama fl. 522. 1920 (non Hepp in Zollinger,<br />
1854). Type. BAHAMAS. Watling's Island: vicinity of Cockburn Town, 13 Mar 1917, Britton &<br />
Millspaugh 6130, not seen.<br />
New to North America. I have not seen the type collection but two "paratypes" agree with the Florida<br />
material. It is no surprise to find a species desribed from the Bahamas on Key West. I have no recent<br />
collections. FLORIDA. Monroe County: Key West, Feb 1898, Thaxter 395, 420, s.n. (NY).<br />
55
PLEOSPORALES<br />
NAETROCYMBACEAE Höhnel ex R. C. Harris, fam. nov.<br />
Naetrocymbeen Höhnel, Sitzungsber. Kaiserl. Akad. Wiss. Math.-Naturwiss. Cl., Abt. 1, 118: 1200.<br />
1909, nom. inval. [not in Latin or form complying with ICBN].<br />
Familia Pleosporalium pseudoparaphysibus peculiaribus areis refractivis ad extremos cellulorum, ascis<br />
obpyriformibus loculo apicali lato, plerumque in sectione subtriangulari, ascosporis hyalinis vel brunneis,<br />
plerumque granularibus ornamentatis et microconidiis breviter bacillaribus.<br />
Type. Naetrocymbe Körber.<br />
Eriksson (1981) had an extensive description of the type species of Naetrocymbe as N. fumago where he<br />
illustrated the odd physes with the ends of the cells usually ± refractive and differentially staining. It is also<br />
described under Arthopyrenia rhyponta. The validation of this family is necessitated due to facing up to the<br />
fact that Arthopyrenia s. lat. contains two major disparate elements. This is more extensively discussed under<br />
Arthopyreniaceae. In addition to the short-celled pseudoparaphyses, the group is supported by the obpyriform<br />
asci with a rather distinctive apical region (lacking a nasse often seen in Arthopyreniaceae), ornamented<br />
ascospores, short rod-shaped microconidia and a mainly temperate/boreal distribution. Currently I accept four<br />
genera in the Naetrocymbaceae, Jarxia, Naetrocymbe and Tomasellia with reasonable confidence.<br />
Leptorhaphis is placed here more or less by default. The physes are not quite right, the asci are more oblong<br />
although they seem to have a similar apical region and macroconidia are known. As far as I have been able<br />
to ascertain over the years none of the members of this family are lichen-forming. In my opinion it makes<br />
about as much sense to include Peziza or Sordaria in checklists of lichens as it does to include members of<br />
the Naetrocymbaceae.<br />
KEY TO THE GENERA<br />
1. Ascospores colorless (sometimes weakly tinted when postmature)......................................................... 2<br />
2. Ascomata with a single chamber .......................................................................................................... 3<br />
3. Ascospores soleiform, 1-septate to submuriform ........................................................ Naetrocymbe<br />
3. Ascospores narrowly fusiform arcuate, 1-5-septate..................................................... Leptorhaphis<br />
2. Ascomata compound with several/many centra under a single<br />
clypeus; ascospores 4-celled, 14-20 × 4-5.5 µm; Duval<br />
County ............................................................................. Tomasellia americana (Willey) R. C. Harris<br />
1. Ascospores brown, 1-septate (rarely 3-septate when postmature);<br />
tropical/subtropical (but poorly known)..............................................................................................Jarxia<br />
JARXIA D. Hawksw.<br />
Stud. Mycol. 31: 93. 1989.<br />
Jarxia is distinguished from Mycomicrothelia in coarser physes with refractive areas at the cell ends,<br />
obpyriform asci and differently proportioned, thinner walled ascospores. The genus is probably overlooked<br />
due to its small ascomata and likely to be more common than the few collections indicate.<br />
1. Ascospores (23-)25-28 × (5.5-)6.5-7(-8.5) µm; on Ilex;<br />
56
Duval County...................................................................................J. thelenula (Müll. Arg.) D. Hawksw.<br />
1. Ascospores 17-21 × 5-7 µm; on Ilex; Monroe and<br />
Duval counties....................................................................................................... J. ilicicola R. C. Harris<br />
Jarxia ilicicola R. C. Harris, sp. nov.<br />
NOTES<br />
Jarxiae thelenulae similis sed ascosporis minoribus, 17-21 × 5-7 µm.<br />
Type. FLORIDA. [Duval County: Jacksonville?] on Ilex cassine, Calkins, Fla. Lich. 152 (NY, holotype).<br />
Thallus indicated by grayish blotch on bark, with blackish antagonism zone. Ascomata scattered,<br />
hemispherical to subglobose, superficial to ca. ½ immersed, 0.15-0.2 mm diam.; wall not melanized at base.<br />
Asci obpyriform, ca. 60 × 20-30 µm, with irregularly arranged ascospores. Ascospores yellow-brown, 2-celled,<br />
rarely? 4-celled in age, 17-21 × 5-7 µm, finely granular ornamented. Microconidia not found.<br />
Additional specimens. FLORIDA. Duval County: Ft. George [Island], on Ilex, Calkins, N. Amer. Lich. 280;<br />
Monroe County: hammock, S end of Big Pine Key, 11 Dec 1921, Small et al. 10159 (both NY).<br />
Jarxia thelenula (Müll. Arg.) D. Hawksw.<br />
New to North America. Reported by Hawksworth from Cuba and the Dominican Republic.<br />
FLORIDA. Duval County: Jacksonville, [on Ilex], Calkins, N. Amer. Lich. 255 (NY). TEXAS. Webb County:<br />
University of Wisconsin monkey colony, 30 mi N of Loredo on U.S. Hwy. 83, 2 Dec 1972, Iltis 27420 (MIN).<br />
LEPTORHAPHIS Körber<br />
The genus is almost entirely northern with only a single scrappy collection from Florida. It seems close to<br />
L. amygdali (Massal.) Zwackh and L. parameca (Massal.) Körber (both on Prunus) in having a broad basal<br />
fringe, IKI+ bluish hymenial gel and similar ascospore shape. However, the ascospores are broader, 37-47 ×<br />
4-5.5 µm, and it does not occur on Prunus. Better material is need before it is formally described. For the<br />
northern species the excellent revision by Aguirre-Hudson (Bull. Br. Mus. Nat. Hist. (Bot.) 21: 85-192. 1991)<br />
should be consulted. Otherwise a pyrenocarpous specimen with filiform ascospores from Florida is probably a<br />
Celothelium.<br />
FLORIDA. Monroe County: hammock, S end of Big Pine Key, 11 Dec 1921, Small et al. 10154-D (NY).<br />
NAETROCYMBE Körber<br />
Santessoniolichen Tomaselli & Cifferi, Arch. Bot. (Forli) 28: 5. 1952. ≡ Santessoniomyces Cifferi &<br />
Tomaselli, Ist. Bot. Reale Univ. Reale Lab. Crittog. Pavia Atti, ser. 5, 10: 29,57. 1953. Type. S.<br />
punctiformis ("Fr.") sensu Jatta [sic].<br />
?Sporoschizon Riedl, Sydowia 14: 334. 1960. Type. Sporoschizon petrakianum Riedl (= Naetrocymbe<br />
punctiformis s. lat.).<br />
My previous intent was to recognize N. rhyponta as a monotypic Arthopyrenia separate from the<br />
punctiformis-group (which I have occasionally annotated as Santessoniolichen). The "retypification" of<br />
Arthopyrenia has changed that. Also, I have recently concluded that the differences are minor. The most<br />
obvious being the dark brown subiculum usually present in N. rhyponta. Foucard (1992) indicated from his<br />
study of ample material that it probably is of no taxonomic significance. Eriksson (1987) recognized the type<br />
of Naetrocymbe, N. fumago, as belonging in Arthopyrenia s. lat. and Foucard (1992) further considered it to be<br />
synonymous with Arthopyrenia rhyponta. Although I have not seen type material of N. fumago, I concur with<br />
57
their decisions and the resurrection of the genus and family is the result. Sporoschizon is synonymized with<br />
question as I have not seen the holotype of Sporoschizon petrakianum but an isotype (Krypt. Vind. 4454, S)<br />
does not show any ascospore fragmentation which was the basis of Riedl's genus and is synonymous with<br />
Naetrocymbe punctiformis s. lat. I am making only such combinations in Naetrocymbe as affect North<br />
America (although it is my hope they will not be included in the checklist). I hope this is the last time I have to<br />
deal with this peripheral non-lichenized taxon. It will be omitted from a final treatment of Florida lichens.<br />
KEY TO NAETROCYMBE IN NORTH AMERICA<br />
1. Ascospores 2-4-celled............................................................................................................................... 2<br />
2. Ascopsores 2-celled (rarely 4-celled when postmature) ....................................................................... 3<br />
3. Physes persistent .............................................................................................................................. 4<br />
4. Ascospores smaller, 26 µm or less long ....................................................................................... 5<br />
5. Ascomata superficial, flattened, often surrounded by dark hyphal ring..................................... 6<br />
6. Ascospores 19-25 × 5.5-7.5 µm; on Fraxinus; Maine,<br />
Minnesota and Vermont ...................................................... [N. fraxini (Massal.) R. C. Harris]<br />
6. Ascospores 15-20 × 4.5-5.5(-6.5) µm; on a variety of<br />
smooth barks; northern U.S. (southern limit in East is<br />
Long Island)................................................................... [N. punctiformis (Pers.) R. C. Harris]<br />
5. Ascomata at least initially immersed, ± globose ....................................................................... 7<br />
7. Ascospores 22-26 × 6-7.5 µm; known only from Florida, Duval,<br />
Gilchrist, Levy and Seminole counties ...............................N. cedrina (Zahlbr.) R. C. Harris<br />
7. Ascospores 16-22 × 4-5 µm; asci slender, 55-70 × 12-16 µm;<br />
rare, Seminole County and Alabama ........................N. atractospora (Zahlbr.) R. C. Harris<br />
4. Ascospores large, 25-32 × 6-8 µm; ascomata superficial, flattened;<br />
Maine, Europe .................................................................... [N. megalospora (Lönnr.) R. C. Harris]<br />
3. Physes dissolving with only fragments remaining; ascospores<br />
14.5-17 × 4-5 µm; Monroe County (Keys) ..................................N. quassiaecola (Fée) R. C. Harris<br />
2. Ascospores 4-celled.............................................................................................................................. 8<br />
8. Ascomata resting in extensive dark brown subiculum,<br />
0.2-0.3 mm diam.; ascospores 17-20 × 5.5-7 µm;<br />
northern U.S.? (no "good" American material seen) ........................ [N. rhyponta (Ach.) R. C. Harris]<br />
8. Ascomata not surrounded by a subiculum, 0.1-0.2<br />
mm diam.; ascospores 17-21 × 5-6 µm; subtropical,<br />
Duval County....................................................................N. atomarioides (Müll. Arg.) R. C. Harris<br />
1. Ascospores 6-celled, 20-23 × 5-5.5 µm; physes persistent;<br />
Duval County............................................................................................. Naetrocymbe sp. Calkins 157<br />
NOTES<br />
Naetrocymbe atomarioides (Müll. Arg.) R. C. Harris, comb. nov.<br />
58
Arthopyrenia atomarioides Müll. Arg., Bot. Jahrb. Syst. 6: 406. 1885. Type. CUBA. Wright, Lich.<br />
Cub. II. 629 (G, holotype).<br />
Verrucaria subpunctiformis Nyl., Lich. ins. guin. 51. 1889. Arthopyrenia floridana Zahlbr., Cat. lich.<br />
univ. 1: 332. 1922, nom. nov. Type. FLORIDA. [Duval County:] Ft. George, 1887, Calkins<br />
(FH, MICH, NY, US, isotypes?).<br />
Naetrocymbe atractospora (Zahlbr.) R. C. Harris, comb. nov.<br />
Arthopyrenia atractospora Zahlbr., Ann. Mycol. 33: 33. 1935. Type. FLORIDA. [Seminole County:]<br />
Sanford, on Quercus, Jul 1928, Rapp137 (W 2451, holotype).<br />
Additional specimen. ALABAMA. Baldwin County: Yupon Point, 1 Mar 1925, Evans 210 (MICH).<br />
Naetrocymbe cedrina (Zahlbr.) R. C. Harris, stat. et comb. nov.<br />
Arthopyrenia sanfordensis var. cedrina Zahlbr., Ann. Mycol. 33: 33. 1935. Type. FLORIDA.<br />
[Seminole County] Sanford, on Taxodium, Oct 1922, Rapp 135 (W 2459, holotype).<br />
The type and the Gilchrist County collection are on Taxodium, the Duval County and Alabama collections<br />
on Myrica. Microscopically rather similar to N. fraxini, this species is distinct in the smaller more globose<br />
ascomata, substrate and distribution.<br />
Naetrocymbe fraxini (Massal.) R. C. Harris, comb. nov.<br />
Arthopyrenia fraxini Massal., Ric. auton. lich. crost.167, f. 333. 1852. Type. "Ad truncos Fraxini<br />
ubique in regno Veneto. Massal.", Massal. Lich. Ital. 298A (BM, lectotype, Coppins, 1988, not<br />
seen by me).<br />
Naetrocymbe megalospora (Lönnr.) R. C. Harris, comb. nov.<br />
Arthopyrenia megalospora Lönnr., Flora 41: 634. 1858. Type. SWEDEN. Fries, Lich. Suec. 244<br />
(FH-Tuck 4067, isolectotype, Harris, 1975).<br />
Foucard (Graphis Scripta 4: 49-60. 1992) has placed this in synonymy with N. fraxini on no better<br />
grounds than those by which I recognize it, speculation. The ascospores seem significantly larger to me. I<br />
prefer to keep the question open and therfore make the combination in Naetrocymbe.<br />
Naetrocymbe massalongiana (Hepp) R. C. Harris, comb. nov.<br />
Sagedia massalongiana Hepp, Flecht. Eur. 444. 1857. Type. SWITZERLAND. Zurich and Liestal,<br />
Hepp (NY).<br />
This species has been previously synonymized with N. saxicola (see below) but the asci and ascospores<br />
are larger than in the probable type of N. saxicola (Massalongo, Lich. Ital. 348). The asci are 75-85 × 20-25<br />
µm vs. ca. 50 × 20 µm and are less distinctly pyriform than in N. saxicola. The ascospores are 18-25 × 6-7(-8)<br />
µm vs. 16-20 × 4-5 µm. A broader study may find these differences trivial and recognition here is mainly<br />
intended as a stimulus for such a study. There are apparently two colections distributed in Hepp 444 and I<br />
have no way of making an intellegent lectotype. Hepp's description is also apparently mixed. The illustration<br />
of the ascus and the measurements of ascus and ascospores agree with the Naetrocymbe. The 4-celled<br />
ascospores would seem to belong to Trichothelium lineare which is mixed in one of the NY packets of Hepp<br />
444. Its previous synonymization with N. saxicola would seem to indicate maintaining the name for the<br />
Naetrocymbe. It is not known from North America.<br />
Naetrocymbe punctiformis (Pers.) R. C. Harris, comb. nov.<br />
Verrucaria punctiformis Pers., Ann. Bot. (Usteri) 11: 19. 1794. Arthopyrenia punctiformis auct.<br />
non Massal. Type. "Verrucaria punctiformis", sine loca in herb. Persoon (L, lectotype, Riedl,<br />
1977, not seen).<br />
Arthopyrenia padi Rabenh., Lich. Eur., fasc. 14, 390. 1858. Type. "an Prunus Padus bei<br />
Hermsdorf in Bilaer Grunde in der sächse Schweiz." (MICH, isotype).<br />
One of the benefits of the removal of this species to a genus other than Arthopyrenia is that it allows<br />
Persoon's sacred, although commonly misapplied, epithet to be taken up again legitimately. (Because of the<br />
existence of Arthopyrenia punctiformis Massal. which excluded Persoon's name, I have been using A. padi<br />
59
Rabenh.) Foucard (Graphis Scripta 4: 53. 1992) designated a neotype. However, Riedl (Sydowia 29: 115-<br />
138. 1977) described Persoon's material at great length and provided a photograph of its label. Until there is<br />
some statement by Prof. Riedl that this Persoon material is lost or destroyed, I consider the neotype to be<br />
superseded.<br />
Naetrocymbe quassiaecola (Fée) R. C. Harris, comb. nov.<br />
Verrucaria epidermidis var. γ. quassiaecola Fée, Essai crypt. écorc. 84. 1825. Arthopyrenia<br />
quassiaecola (Fée) Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 30(3): 26. 1888. Type.<br />
AMERICA. On Quassia excelsa (G, lectotype; G, isolectotype).<br />
The ascospores of Florida material are slightly larger than those of the type. There is a single specimen<br />
from Puerto Rico. The physes dissolve leaving only fragments embedded in gel as in Arthopyrenia salicis<br />
Massal. s. lat. I include quassiaecola in Naetrocymbe but suspect that the dissolving hamathecium indicates<br />
that it really belongs elsewhere but have no suggestions at this time.<br />
Naetrocymbe rhyponta (Ach.) R. C. Harris, comb. nov.<br />
Verrucaria rhyponta Ach., Kongl. Vetensk. Akad. Nya Handl. 30: 150. 1809. Arthopyrenia<br />
rhyponta (Ach.) Massal., Ric. auton. lich. crost. 166. 1852. Type. SWEDEN. On Fraxinus (H-Ach<br />
772, the three left hand pieces, lectotype, selected here).<br />
Naetrocymbe saxicola (Massal.) R. C. Harris, comb. nov.<br />
Arthopyrenia saxicola Massal., Symm. lich. nov. 107. 1855. Pyrenocollema saxicola (Massal.)<br />
Coppins in Coppins et. al., Lichenologist 24: 368. 1992. Type. ITALY. Massalongo, Lich. Ital. 348<br />
(NY, isotype?).<br />
Although not in North America, I treated this species briefly in my thesis, placing it in Pyrenocollema, and<br />
now wish to correct my misunderstanding of its taxonomic position. I was uneasy about its disposition and left<br />
it to Coppins to do the dirty work of making the transfer. In restudying some of the problems in Pyrenocollema<br />
for this volume, I realized that the asci were pyriform and that the hamathecium and microconidia were of the<br />
Naetrocymbe-type. I do not know what the biology of this species is. It may be a parasite as it grows with<br />
cyanobacteria on limestone with no obvious thallus formation. (This association with cyanobacteria suggested<br />
Pyrenocollema.) I have rummaged briefly for an appropriate parasite/parasymbiont genus but none is<br />
immediately obvious. Careful morphological studies combined with the different? biology of N. saxicola and N.<br />
massalongiana (segregated from N. saxicola above) may well find that generic recognition is indicated.<br />
TOMASELLIA Massal.<br />
The genus is treated in a narrower sense than previously. Most of the species have been transferred to<br />
Mycoporum (see Mycoporaceae for discussion). There is only a single species in Florida, T. americana<br />
(Willey) R. C. Harris, based on a single Calkins collection from Duval County which is, thus, unfortunately a<br />
little suspect.<br />
MYCOPORACEAE Zahlbr.<br />
This group has turned out to be one of my major early failures. In my thesis work I was so impressed by<br />
the compound nature of the ascomata that I did not look closely enough at the characters of the ascus and<br />
hamathecium. In re-evaluating this group I have come to the conclusion that most of the taxa referred to<br />
Mycoporellum Müll. Arg. and Mycoporopsis Müll. Arg. are not congeneric with Tomasellia Massal. and, at least<br />
tentatively, not confamilial. In 1975 I declined to recognize 2-celled versus 4-celled ascospores as a generic<br />
difference. I have since found taxa with ascospores intermediate between transversely septate and muriform.<br />
Further, on the basis of ascus and hamathecium I believe that two species with simple, uniloculate ascomata<br />
should be placed in this group also. Assuming all transversely sepatate species are placed in one genus,<br />
traditional mycological taxonomy would recognize four genera, Mycoporum (compound ascomata, muriform<br />
ascospores), Mycoporellum (compound ascomata, transversely septate ascospores), unnamed (simple<br />
60
ascomata, muriform ascospores) and unnamed (simple ascomata, transversely septate ascospores).<br />
However, I do not feel that these distinctions are supported by any additional characters and have no<br />
predictive value. Therefore, I recognize only the single genus Mycoporum Flotow ex Nyl. as have Eriksson<br />
(1981) and Barr (1987), but with a broader circumscription.<br />
Both Mycoporum and Tomasellia have compound ascomata with several to many locules, each with its<br />
own ostiole (except for the ostioles the ascomata superficially resemble those of Arthonia and have often been<br />
confused). The asci of Mycoporum are obpyriform to obclavate with the wall in the upper third very much<br />
thickened and with virtually no stipe or with a broad, short stipe. Those of Tomasellia arthonioides (Massal.)<br />
Massal. (the generitype) are ± cylindrical with relatively even wall thickenings and a short, broad stipe. The<br />
hamathecium in Mycoporum consists of very irregular, slightly gelatinizing and somewhat disarticulating<br />
physes which are almost entirely obscured by irregular oil droplets. (If you can make out the physes clearly, it<br />
probably isn't a Mycoporum). That of Tomasellia consists of narrower, more regular physes which may also<br />
gelatinize somewhat, having a refractive body at one or both ends of the cells as in Arthopyrenia rhyponta<br />
(Ach.) Massal. and lacks oil droplets. The ascospores of Mycoporum are thin walled, often collapsing, and<br />
tend to turn yellow brown with a conspicuous granular ornamentation when old. Those of Tomasellia are<br />
comparatively thicker walled and do not readily become brownish and ornamented. The microconidia in both<br />
genera are short, ca. 5 µm, rod-like or oblong. Possible oblong macroconidia are found in M. buckii assigned<br />
to Mycoporum with some reservation. The biological status of Mycoporum is unclear. I was incorrect in<br />
suggesting Trentepohlia as a photobiont. There is no morphologically differentiated lichen thallus although in<br />
section thick walled superficial hyphae are present which become brown and more convoluted near the<br />
ascomata. Some species seem not to be lichen-forming. Others usually seem to have a cover of "nonstandard"<br />
algae (i.e., not similar to known photobionts). It may be that Mycoporum is weakly or facultatively<br />
lichen-forming. If so, it represents another independent development of the lichen lifestyle. Tomasellia seems<br />
definitely not lichen-forming.<br />
Purvis et al. (1992) include two species in Mycoporum, M. hippocastani (DC.) Coppins and M. quercus<br />
(Massal.) Müll. Arg., both of which seem to me to belong elsewhere. They have ± spherical asci, ±<br />
paraplectenchymatous hamathecial tissue and the ascoma is not of the "clypeal" type but a stroma pushing<br />
through the upper layers of bark, possibly non-ostiolate. My current thought is that these two species should<br />
be referred to Cyrtidula Minks. Vainio (1921) emended the genus retaining only one of the original species<br />
and that can serve as a lectotypification with C. pteleodes (Ach.) Minks (= hippocastani). The combination C.<br />
quercus (Massal.) Minks exists. The other combination seems not to have been made. Cyrtidula<br />
hippocastani (DC.) R. C. Harris, comb. nov. (Verrucaria hippocastani DC., Fl. franç. 6: 172. 1815.). I have<br />
no records of either from North America.<br />
Of the species referred to Tomasellia in addition to the type, T. arthonioides, I would retain T. americana<br />
(Minks ex Willey) R. C. Harris, T. diffusa (Leighton) Lahm., T. gelatinosa (Chev.) Zahlbr. and T. macularis<br />
(Minks ex Willey) R. C. Harris, comb. nov., (Cyrtidula macularis Minks ex Willey, Enum. lich. New Bedford<br />
33. 1892. Type. MASSACHUSETTS. New Bedford, Willey, (MICH, isosyntype), treated in 1975 but combination<br />
not validly published). Tomasellia americana and T. macularis are maintained provisionally pending reexamination<br />
of type material. All four differ somewhat from the type species in more obclavate asci and are<br />
kept in Tomasellia on the basis of compound ascoma and hamathecial type. The genus Athrismidium Trevisan<br />
(syn. Beckhausia Hampe ex Körber, nom. inval.), typified with A. gelatinosa, is available for the others if one<br />
wishes to restrict Tomasellia to its type species. Tomasellia diffusa was included in Melanthecopsis Dodge.<br />
However, the type of this genus, M. follmanii Dodge, is a Pyrenula, close to or synonymous with P. cubana<br />
(Müll. Arg.) R. C. Harris. Melanothecopsis sinensis (Krempelh.) Dodge is a coelomycete with brown 2-celled<br />
conidiospores. The other two species, M. coarctella (Stirton) Dodge and M. subpuncta (Nyl.) Dodge are<br />
unknown to me. Tomasellia americana is treated in the Naetrocymbaceae, T. arthonioides and T. diffusa do<br />
not occur in North America, T. gelatinosa is known from Maine and Newfoundland and T. macularis from<br />
Massachusetts.<br />
The following are possible generic synonyms of Mycoporum but cannot be placed definitively at this time<br />
as I have not seen type material, Mycoporopsis Müll. Arg., lectotype (Riedl, 1962), Mycoporum sorenocarpum<br />
Knight, and Nothostroma Clements, holotype, Mycoporopsis roseola Müll. Arg.<br />
MYCOPORUM Flotow ex Nyl., nom. cons.<br />
61
Mém. Soc. Sci. Nat. Cherbourg 3: 186. 1855. Type. M. elabens Flotow ex Nyl.<br />
Bottaria Massal., Misc. lichenol. 43. 1856. Holotype. B. composita Massal.<br />
Mycoporellum Müll. Arg., Rev. Mycol. (Toulouse) 6: 14. 1884. Lectotype (Riedl, 1962). Mycoporum<br />
sparsellum Nyl.<br />
Syngenosorus Trevisan, Conspect. Verruc. 15. 1860. Lectotype (chosen here). Melanotheca arthoniella Nyl.<br />
Chlorodothis Clements, Genera of fungi 50. 1909. Holotype. Mycoporellum lahmii Müll. Arg. (= Mycoporum<br />
lacteum)<br />
Sciodothis Clements, Genera of Fungi 50. 1909. Holotype. Mycoporopsis leucoplaca Müll. Arg. (=<br />
Mycoporum lacteum)<br />
1. Ascoma compound, with several to many chambers, each with a pore,<br />
each containing many asci ........................................................................................................................ 2<br />
2. Ascospores transversely septate .......................................................................................................... 3<br />
3. Ascospores mostly 2-celled, some becoming 4-celled when old...................................................... 4<br />
4. Ascospores fusiform, relatively slender, 2.5-4 times longer than broad........................................ 5<br />
5. Ascospores larger, over 15 × 5 µm; ascoma without well developed<br />
brown basal layer; macroconidia not known ............................................................................. 6<br />
6. Ascospores 17-20(-22) × 5-6(-7) µm; very common, throughout<br />
Florida .................................................................... M. eschweileri (Müll. Arg.) R. C. Harris<br />
6. Ascospores 20-27 × 6.5-8 µm; not uncommon, throughout<br />
Florida ...................................................................................M. lacteum (Ach.) R. C. Harris<br />
5. Ascospores small, 10-13 × 3-3.5 µm; ascoma with a distinct brown<br />
basal layer; macroconidia oblong, ca. 5 × 2 µm; on Taxodium or<br />
old wood; Collier and Duval counties ...................................................... M. buckii R. C. Harris<br />
4. Ascospores relatively short and "fat", 1.5-2.5 times longer than broad,<br />
17-22(-25) × 7.5-10 µm; upper cell markedly shorter than lower;<br />
common, throughout Florida ............................................................................M. sparsellum Nyl.<br />
3. Ascospores mostly 4-celled, only immature ascospores 2-celled..................................................... 7<br />
7. Ascospores 17-22 × 5.5-7 µm; physes obscured by oil;<br />
rare, Citrus and Escambia counties ..................................M. californicum (Zahlbr.) R. C. Harris<br />
7. Ascospores 15-17 × 4-5.5 µm; physes not obscured by<br />
oil; rare, Duval County............................................. Tomasellia americana (Willey) R. C. Harris<br />
2. Ascospores muriform ............................................................................................................................ 8<br />
8. Ascospores over 30 × 12 µm ........................................................................................................... 9<br />
9. Ascospores 30-38(-43) × 12-17(-18) µm, with 8-9 rows of locelli;<br />
Franklin, Gilchrist and Okaloosa counties.......................M. compositum (Massal.) R. C. Harris<br />
9. Ascospores (36-)39-48(-55) × (14-)16-21(-24) µm, with (9-)10(-11)<br />
rows of locelli; Duval County ........................................................ M. pycnocarpoides Müll. Arg.<br />
8. Ascospores 23-28(-30) × 9-12 µm, with 8 rows of locelli; not uncommon,<br />
northern Florida ........................................................................................M. acervatum R. C. Harris<br />
62
1. Ascoma simple, with only a single chamber ........................................................................................... 10<br />
10. Ascospores mostly 2-celled, 4-celled and brownish in old age,<br />
28-40 × 8-12 µm; Lake County.......................................................M. antecellens (Nyl.) R. C. Harris<br />
10. Ascospores muriform, 22-37 × 7-9.5 µm, with 8-9 rows of locelli;<br />
not seen to become brownish; northern Florida.................................... M. uniloculatum R. C. Harris<br />
Mycoporum acervatum R. C. Harris, sp. nov.<br />
NOTES<br />
Similis M. compositi et M. pycnocarpoidis sed ascosporis minoribus, 24-29 × 9-11 µm.<br />
Type. FLORIDA. Clay County: Gold Head Branch State Park, vicinity of Loblolly Trail, mixed hardwoods along<br />
stream, 29 Nov 1992, Harris 29260 (NY, holotype).<br />
Thallus indicated by whitish blotch on bark, in type patchily covered by small clumps of algae. Ascomata<br />
orbicular to ± irregular in outline, flattened, 0.5-1.0 mm diam. Asci obpyriform, 60-75 × 30-40 µm, with<br />
ascospores irregularly arranged. Ascospores oblong-elliptical with rounded ends, slightly constricted at<br />
median septum, colorless (no brown ascospores observed), muriform, with 8 rows of 2-6 cells, 23-28(-30) × 9-<br />
12 µm. Pycnidia with microconidia, oblong, 3 × 1.5 µm.<br />
Mycoporum acervatum is distinguished from its near relatives M. compositum, M. elabens and M.<br />
pycnocarpoides only on ascospore size. Since it has no really distinctive character, I have given it an epithet<br />
which could apply to most of the genus, meaning "in small heaps" in reference to the appearance of the<br />
compound ascoma. I am not really sure how to interpret the ascospore variation in this group. We have no<br />
theoretical basis. Why should basically similar ascospores vary in size? Is it the result of random mutation<br />
fixed in isolated populations? Probably not in this case as compositum and pycnocarpoides co-occur widely in<br />
the Neotropics and Africa and both occur with acervatum in Florida. Is there some microhabitat or dispersal<br />
advantage related to size? Is it just minor genetic "noise", e.g., big vs. little humans? If the latter, then all four<br />
should probably be lumped into a single species. I am not even sure how one would go about addressing this<br />
problem. This question aside, M. acervata does differ in a minor way from compositum and pycnocarpoides in<br />
that I have never observed brown ascospores which are common at maturity in those species. Mycoporum<br />
acervatum is closest in ascospore size to the European M. elabens but the latter has larger, hence fewer<br />
locelli per row.<br />
Additional specimens (a few from old notes not re-examined 1995). ALABAMA. Conecuh County: Brooklyn,<br />
1860-61, Beaumont 218 (FH-Tuck 3735). FLORIDA. Bradford County: Montgomery Conference Center, 4 mi<br />
NW of Keystone Heights, on branches of Quercus margaretta, 11 Feb 1993, Griffin 390 (FLAS); Duval County:<br />
Jacksonville, Calkins, NAL 78 (FH), NAL 163 (NY); Leon County: Apalachicola National Forest, Leon Sinks<br />
Geological Area, oak-pine woods, on Quercus, 2 Dec 1988, Harris 23272 (NY); Marion County: Ocala, 31 Feb<br />
1891, Underwood 1843 (NY); Walton County: Lakewood, Lakewood Park, highest point in Florida, 345 ft, oakhardwood<br />
forest, on Quercus, 30 Nov 1988, Harris 23077 (NY). SOUTH CAROLINA. without location, Ravenel<br />
(FH-Tuck 3735).<br />
Mycoporum antecellens (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria antecellens Nyl., Flora 49: 86. 1866. Arthopyrenia antecellens (Nyl.) Müll. Arg., Flora<br />
53: 485. 1870. Mycoporopsis antecellens (Nyl.) Riedl, Sydowia 15: 268. 1961[1962], nom.<br />
inval., (basionym not cited). Type. ENGLAND. Sussex: Tilgate Forest, Larblastier 77 (H-Nyl<br />
767, holotype).<br />
Pyrenula zwackhii Hepp, Fl. Eur. 954. 1867. Type. GERMANY. Near Heidelberg, Zwackh (FH, FH-<br />
Tuck 4066, NY, isotypes).<br />
63
Verrucaria subantecellens Nyl., Flora 59: 364. 1876, nom. nud. Arthopyrenia subantecellens<br />
Müll. Arg., Bot. Jahrb. Syst. 6: 403. 1885. Type. CUBA. Wright, Verr. Cub. 2 (MSC, holotype<br />
fragment; H-Nyl 769, isotype).<br />
Arthopyrenia griseola Müll. Arg., Rev. Mycol. (Toulouse) 10: 182. 1888. Type. PARAGUAY.<br />
Paraguarí, Balansa (G, holotype).<br />
Presumably on the basis of the ascospores that soon become 4-celled, brownish and granular<br />
ornamented, Riedl removed this species from Arthopyrenia to Mycoporopsis even though the ascomata are<br />
uniloculate. Placement in Mycoporum ( incl. Mycoporopsis sensu Riedl) is further supported by the large<br />
ascospores, large asci, thick walled in the upper part and the characters of the hamathecium. Purvis et al.<br />
(1992) spelled the epithet "antecellans". All the specimens in Nylander's herbarium have the "-ens" spelling<br />
and my small Latin dictionary gives "antecello, -ere" the participle of which should end in "-ens". The British are<br />
noted scholars of Latin and perhaps there is some linguistic point I have missed. However, until I am<br />
corrected, I will spell the epithet "antecellens".<br />
Mycoporum buckii R. C. Harris, sp. nov.<br />
Similis M. sparselli sed ascomatis planioribus, strato brunneo sub hymeniis, ascosporis minoribus, 9.5-12<br />
× 2.5-3 µm et macroconidiis oblongiis, ca. 5 × 2 µm.<br />
Type. FLORIDA. Duval County: Flood plain forest E of San Pablo River (Intracoastal Waterway), W of 17th St.<br />
N., dead portion of trunk of oak, 19 Jun 1975, Buck B175 (NY, holotype; MICH, isotype? not seen).<br />
Thallus a whitish blotch on substrate. Ascomata black, superficial, compound, multiostiolate, with many<br />
chambers, orbicular to elliptical (elongated with the grain of the wood), 0.5-1.0 mm diam, or to ca. 0.5 × 1.0<br />
mm, with a brown basal layer. Asci obclavate to ± fusiform, 40-55 × 10-14 µm, with ascospores weakly<br />
biseriately to irregularly arranged. Ascospores fusiform, colorless, 2-celled, 10-13 × 3-3.5 µm. Microconidia<br />
not found. Pycnidia with macroconidia, oblong, colorless, ca. 5 × 2 µm.<br />
Mycoporum buckii is anomalous in several ways, the narrow asci which are not basally swollen, the well<br />
developed brown layer at the base of the ascoma and the presence of what can be interpreted as<br />
macroconidia. However, I would not feel comfortable creating a new genus and believe it is best<br />
accommodated in Mycoporum pending more thorough investigation. It is easily recognized by the small<br />
ascospores. Some species of Strigula have ascospores this small but have small, uniloculate ascomata. In<br />
addition to the cited collections it has been collected in Belize by Mason Hale.<br />
Additional specimens. FLORIDA. Collier County: Fakahatchee Strand State Preserve, 1.2 mi from N<br />
entrance on W. J. James Memorial Scenic Drive, open Taxodium swamp, on trunk of Taxodium, 6 Dec 1992,<br />
Buck 22817 (NY). DOMINICAN REPUBLIC. Prov. Monte Cristi: SE of El Morro de Monte Cristi, ca. 20 m, low dry<br />
forest near sea, 9 Jan 1987, Harris 19603 (JBSD, NY).<br />
Mycoporum californicum (Zahlbr.) R. C. Harris, comb. nov.<br />
Mycoporellum californicum Zahlbr., Ann. Mycol. 10: 363. 1912. Mycoporopsis californica (Zahlbr.)<br />
Riedl, Sydowia 16: 216. 1962[1963]. Tomasellia californica (Zahlbr.) R. C. Harris in Tucker &<br />
Jordan, Wasmann J. Biol. 36: 87. 1979, comb. inval. (33.2). Type. CALIFORNIA. Santa Monica<br />
Mtns., Hasse 1003 (FH, isotype).<br />
This species is separated from M. eschweileri, that has similar sized ascospores, by the early<br />
development of 4-celled ascospores, and from Tomasellia americana by larger ascospores and hamathecium<br />
obscured by oil droplets. Its distribution is mainly western. The easternmost record previously was from<br />
Texas. However, a number of ± western species are turning up in northern Florida.<br />
64
Mycoporum compositum (Massal.) R. C. Harris<br />
in Egan, Bryologist 90: 164. 1987. Bottaria composita Massal., Misc. lichenol. 43. 1856. Type. ad<br />
cort. Chinae [comm. Hampe] (VER, holotype).<br />
Mycoporum pycnocarpum Nyl., Flora 41: 38. 1858 (lapsus pyrenocarpa). Dermatina pyrenocarpa<br />
(Nyl.) Zahlbr., Cat. lich. univers. 1: 550. 1922. Type. MEXICO. Sierra de Cristóbal, 1855, Fr.<br />
Müller (H-Nyl 4223, holotype; MICH, isotype?).<br />
Arthonia tumidula Leighton, J. Linn. Soc., Bot. 9: 200. 1867. Arthothelium tumidulum (Leighton)<br />
Zahlbr., Cat. lich. univers. 2: 137. 1922. Type. [CANADA.] St. Mary's, Lake Superior, Richardson<br />
158 (BM, lectotype, selected by Makhija & Patwardhan, 1995; BM, isolectotype (neither seen);<br />
NY, isolectotype).<br />
Mycoporum integrum Vainio, Mycologia 21: 40. 1929. Type. PUERTO RICO. On bark near Aibonito,<br />
3 Jan 1916, Fink 1800 (FH(2), MICH-Fink 8633, NY, "isolectotypes". (I have not examined<br />
material in Vainio's herbarium but his published ascospore size agrees with M. compositum.<br />
Other duplicates (MICH-Fink 8953, NY) have the larger ascospores of M. pycnocarpoides. The<br />
official designation of a lectotype can wait pending examination of material at Turku.)<br />
Mycoporum pycnocarpum var. ohiense Nyl., Mém. Soc. Sci. Nat. Cherbourg 5:135. 1867, nom.<br />
nud. Mycoporum ohiense Nyl. ex Fink, Lich. Fl. U.S. 67. 1935. Dermatina ohiensis (Fink)<br />
Zahlbr., Cat. lich. univ. 10: 108. 1938. Lectotype (Harris, 1973). OHIO. Lea (ex hb. Hooker) (H-<br />
Nyl 4234).<br />
Tomasellia cubana Müll. Arg.<br />
Bot. Jahrb. Syst. 6: 397. 1885. Type. CUBA. Wright, Verr. Cub. 126 (NY, isotype).<br />
This name is a synonym of Arthopyrenia consanguinea Müll. Arg.<br />
Mycoporum difforme Minks, Mycoporellum difforme (Minks) Fink<br />
Described from Massachusetts but I have not been able to locate type. From the description by Riedl<br />
(1961) it is a synonym of M. lacteum and should be removed from the checklist.<br />
Mycoporum eschweileri (Müll. Arg.) R. C. Harris, comb. nov.<br />
Mycoporellum eschweileri Müll. Arg., Flora 71: 526. 1888, Tomasellia eschweileri (Müll. Arg.) R.<br />
C. Harris in Tucker & Harris, Bryologist 83: 19. 1980. Type. BRAZIL. Bahia (G, holotype).<br />
Tomasellia esenbeckiana (Fée) Müll. Arg. = Arthopyrenia esenbeckiana<br />
Tomasellia exumana Riddle<br />
in Britton & Millspaugh, Bahama Fl. 528. 1920. Type. BAHAMAS. Great Exuma: on bark near<br />
Georgetown, 22-28 Feb 1905, Britton & Millspaugh 3135 (NY, isotype).<br />
This is a synonym of Anisomeridium anisolobum (Müll. Arg.) Aptroot.<br />
Mycoporum lacteum (Ach.) R. C. Harris, comb. nov.<br />
Verrucaria stigmatella var. lactea Ach., Lichenogr. universalis 227. 1810. Verrucaria lactea (Ach.)<br />
Eschw. in Martius, Fl. bras. enum. pl. 1: 126. 1833. Mycoporellum lacteum (Ach.) Zahlbr., Cat.<br />
lich. univers. 1: 554. 1922. Tomasellia lactea (Ach.) R. C. Harris in Tucker & Harris, Bryologist<br />
83: 19. 1980. Type. Amer. meridion., in Copaifera, [Swartz], (H-Ach 779A, holotype; BM-Ach,<br />
UPS, isotypes).<br />
Mycoporopsis leucoplaca Müll. Arg., Flora 69: 316. 1886. Mycoporellum leucoplacum (Müll. Arg.)<br />
Zahlbr., Nat. Pflanzenfam. 1(1*): 78. 1903. Type. BRAZIL. Minaes Geraes, Glaziou (FH, FH-<br />
Tayl 118, US, isotypes).<br />
Mycoporellum lahmii Müll. Arg., Flora 70: 78. 1887. Type. SOUTH AFRICA. Transvaal: near<br />
Lydenburg, Wilms, comm. Lahm 61 (G, holotype).<br />
?Mycoporum difforme Minks ex Willey, Enum. lich. New Bedford 32. 1892. Type not seen.<br />
Mycoporopsis exigua Zahlbr., Sitzungsber. K. Akad. Wiss. Wien. Math.-Naturw. Cl. 111(1): 360,<br />
pl. 2, f. 24-26. 1902. Type. BRAZIL. Near Theresiopolis, Höhnel, Lich. Bras. 159 (W, holotype).<br />
Mycoporellum hassei Zahlbr. in Hasse, Bryologist 15: 46. 1912. Type. CALIFORNIA. Catalina<br />
65
Island: near Avalon, Hasse 1321 (W, holotype; FH, isotype).<br />
Mycoporellum deserticola Fink in Hedrick, Mycologia 22:248. 1930. Mycoporopsis deserticola<br />
(Fink) Riedl, Sydowia 16: 218. 1962[1963]. Type. PUERTO RICO. Yauco, 1 Jan 1916, Fink<br />
1688 (MICH, holotype).<br />
Tomasellia leucostoma Müll. Arg.<br />
See Arthopyrenia confluens R. C. Harris<br />
Mycoporum mycoporoides (Müll. Arg.) R. C. Harris, comb. nov.<br />
Arthopyrenia mycoporoides Müll. Arg., Rev. Mycol. (Toulouse) 10: 4. 1888. Type. URUGUAY.<br />
Montevideo, 1887, Arechavaleta (G, holotype).<br />
This species has uniloculate ascomata and in hamathecial and ascal characters seems very close to M.<br />
antecellens. In view of the epithet Müller had some inklng of its generic affinities. It differs in having 6-celled<br />
ascospores.<br />
Mycoporum pycnocarpoides Müll. Arg.<br />
Bot. Jahrb. Syst. 20: 290. 1895. Type. TANZANIA? Usambara, im Lutindi-Hochwald, Holst 3373 (G,<br />
holotype).<br />
Mycoporopsis rappii (Zahlbr.) Riedl = Melaspilea Nyl. s. lat. (see Arthopyrenia).<br />
Mycoporum sparsellum Nyl.<br />
Ann. Sci. Nat. Bot. sér. 5, 7: 343. 1867. Mycoporellum sparsellum (Nyl.) Müll. Arg., Rev. Mycol.<br />
(Toulouse) 4: 14. 1884. Tomasellia sparsella (Nyl.) R. C. Harris in Tucker & Jordan, Wasmann<br />
J. Biol. 36: 87. 1979 comb. inval (33.2). Type. COLOMBIA. Monte del Morro, Lindig (H-Nyl 4292,<br />
lectotype, selected here).<br />
Mycoporum uniloculatum R. C. Harris, sp. nov.<br />
Quoad ascos et ascosporas similis M. acervatae sed ascomatis uniloculatis, quoad ascomatas similis M.<br />
antecellentis sed ascosporis muriformibus.<br />
Type. FLORIDA. Liberty County: Apalachicola National Forest, pine-oak scrub S of Forest Serv. Rd. 113, 0.8 mi<br />
W of County Road 12, on small branches of Quercus, 6 May 1990, Harris 25258 (NY, holotype).<br />
Thallus indicated by pale olive brown blotch apparently due to a supeficial layer of algae in clumps that do<br />
not seem to be surrounded by hyphae. Ascomata orbicular, flattened, 0.1-0.2 mm diam., with a single<br />
hymenium, with a weak clypeal ring. Physes with short, irregular cells, mostly obscured by oil droplets. Asci<br />
obpyriform, ca. 60-75 × 30-35 µm, with irregularly arranged ascospores. Ascospores fusiform, constricted at<br />
median septum, colorless, muriform, with (6-)8 rows of 1-4 locelli, 22-32 × 7-9.5 µm. Pycnidia with short rodlike<br />
microconidia, 3-5 × 1 µm.<br />
As noted in the introduction this species is included in Mycoporum on the basis of features of ascus and<br />
hamathecium. In ascospore size M. uniloculata is close to M. acervata but has a narrower, more pointed,<br />
fusiform shape and fewer cells in a row. The general shape and size of the ascospores is close to M.<br />
antecellens except that they are muriform. All the collections are on oak twigs or small branches from pineoak<br />
scrub habitat. It is probably common but overlooked.<br />
Additional specimens. FLORIDA. Clay County: Gold Head Branch State Park, along Florida Trail from<br />
cabins to clay pit in SW corner of park, pine-oak scrub, 29 Nov 1992, Harris 29241; Marion County: Ocala<br />
National Forest, Huges Island, slightly moister oak woods surrounded by dry sand pine scrub, 22 Apr 1967,<br />
Harris 2007-A; Okaloosa County: 1.2 mi W of Fla. Hwy. 85 on Antioch Road (Co. Rd. 4), ca. 4 mi S of<br />
Crestview, oak-pine scrub, 5 May 1990, Buck 17917 (all NY).<br />
66
XANTHOPYRENIACEAE Zahlbr.<br />
It has been suggested that this family should be synonymized with the Arthopyreniaceae. I prefer to keep<br />
it separate on the basis of physes which are more intricately and densely branched and interconnected,<br />
differently shaped asci, differently shaped ascospores and elliptical (not rod-shaped) microconidia. The<br />
differences do not seem major but I think that the relationships of the Xanthopyreniaceae probably lie<br />
elsewhere in some "non-lichenized" family. Also since I have invested considerable energy in making the<br />
Arthopyreniaceae homogeneous, I am reluctant to "mess it up". The inclusion of Zwackhiomyces is basically<br />
arbitrary. There are few useful characters avaiable to place these simple/highly reduced taxa. Essentially<br />
there are no characters which would exclude Zwackhiomyces from the Xanthopyreniaceae but equally there<br />
are no unique shared characters to support inclusion. Collemopsidium Nyl. and Didymellopsis (Saccardo)<br />
Clements & Shear are also homeless and if the positioning of Zwackhiomyces in the Xanthopyreniaceae<br />
works out, perhaps, they too could be dumped here.<br />
1. Associated with cyanobacteria; pigment in ascomatal wall amorphous, located<br />
in the hyphal cell wall; ascospores thin-walled, readily collapsing, not becoming<br />
brownish and granular-roughened .....................................................................................Pyrenocollema<br />
1. Parasitic or parasymbiontic on lichens with green algal photobiont; pigment in<br />
ascomatal wall fine granular, located between hyphae; ascospores with thicker<br />
wall, not readily collapsing (until overmature), becoming brownish and granular<br />
ornamented ..................................................................................................................... [Zwackhiomyces]<br />
PYRENOCOLLEMA Reinke<br />
In my thesis I pointed out the problems involved in defining species in Pyrenocollema. They have not<br />
been resolved and even seem to have worsened. The major unresolved questions are: does the<br />
host/photobiont matter (and therefore does the rock substrate matter, i.e., preference of the cyanobacteria) or<br />
should the taxonomy be based entirely on fungal characters?<br />
1. Fresh water aquatics, semi-aquatics or on rock in moist places...............................................................2<br />
2. Photobiont blue-green in color; on calcareous rock .............................................................................. 3<br />
3. Ascomata sessile to semi-immersed, ascospores 17-23 × 8-11 µm;<br />
eastern United States ..................................................[P. prospersellum (Nyl.) R. C. Harris]<br />
3. Ascomata immersed, with a broad clypeus; ascospores 18-23 × 8-9.5 µm;<br />
Bermuda............................................................................................ [P. farlowii (Riddle) R. C. Harris<br />
2. Photobiont yellow brown to brown due to colored sheath (Gleocapsa-like)<br />
or thallus dark brown ............................................................................................................................ 4<br />
4. Growing on limestone; ascspores 16-18(-20) × 5-7 µm; Monroe County<br />
(Key West), Puerto Rico.............................................................P. atlanticum (Vainio) R. C. Harris<br />
4. Growing on a variety of rock, usually ± aquatic; ascospores<br />
(18-)20-27 × 8-11 µm; temperate North America, Europe ..................................................................<br />
.....................................................................................[P. tichothecioides (Arnold) R. C. Harris s. lat.]<br />
1. Marine, in tidal zone ....................................................................... [P. halodytes (Nyl.) R. C. Harris s. lat.]<br />
[In my thesis I took a very broad view of this species. It is divided in Purvis et al. (1992) as follows.<br />
I am not sure at this time how this view applies in North America.<br />
A. On calcareous rocks or shells; clypeus absent or not spreading;<br />
ascomata not cracked around ostiole ................................................................................................B<br />
67
B. Ascomata sessile and flattened, or immersed and globose, 0.2-0.25 mm<br />
diam.; on calcareous rocks or shells................................................P. halodytes (Nyl.) R. C. Harris<br />
B. Ascomata immersed or protruding, 1-1.5 mm diam.; on chalk or<br />
limestone................................................................................. P. sublitorale (Leighton) R. C. Harris<br />
A. On siliceous rock; clypeus laterally spreading; ascomata cracked<br />
around ostiole; ascospores 15-20 × 6-9 µm.................................... A. orustense (Erichsen) Fletcher]<br />
NOTES<br />
Pyrenocollema atlanticum (Vainio) R. C. Harris, comb. nov.<br />
Collemopsidium atlanticum Vainio, Mycologia 21: 34. 1929. Type. PUERTO RICO. Yauco, 28 Dec<br />
1915, Fink 1402 (MICH, isotype).<br />
The material of Collemopsidium iocarpum (Nyl.) Nyl., the type species of the genus, was still out on loan<br />
when I was in Helsinki so that I am unable to say if Collemopsidium Nyl. (1881) and Pyrenocollema Reinke<br />
(1895) are synonymous. If they should prove to be, although Collemopsidium has priority, I assume<br />
Pyrenocollema would be conserved since it is thoroughly embedded in both North American and European<br />
literature. The isotype of C. atlanticum consists mostly of a species of Lichenaceae. There are, however, a<br />
few patches of pale bluish "thallus" supporting the conclusion drawn from microscopic characters to include<br />
the Florida and recent Puerto Rican specimens in this species. Pyrenocollema atlanticum is quite similar in<br />
aspect to P. caesium but is distinguished by the smaller ascospores.<br />
New to North America. FLORIDA. Monroe County: Key West, Feb 1898, Thaxter 412. PUERTO RICO.<br />
Reserva Forestal Guánica, along Hwy. 333 at Km 4.8, coastal scrub forest, 26 May 1989, Buck 17135 (both<br />
NY).<br />
Pyrenocollema caesium (Nyl.) R. C. Harris<br />
Sagedia nylanderi Hepp, Flecht. Eur. 440. 1857. Thelidium nylanderi (Hepp) Lönnroth, Öfvers.<br />
Förh. Kongl. Svenska Vetensk.-Akad. 15: 284. 1859. Arthopyrenia nylanderi (Hepp) Riedl,<br />
Linzer Biol. Beitr. 22: 154. 1990. Type. SWITZERLAND. "An Alpenfindlingen, am Ufar bei Zürich",<br />
Hepp. (NY, US, isotypes).<br />
Pyrenocollema ceuthocarpoides (Müll. Arg.) R. C. Harris, comb. nov.<br />
Arthopyrenia ceuthocarpoides Müll. Arg., Flora 67: 468. 1884. Type. BRAZIL. São Paulo: "Ad saxa<br />
gneissiaca rivuli Iguape prope Apiahy.", Puiggari (G, holotype).<br />
Pyrenocollema epigloeum (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria epigloea Nyl. Flora 69: 464. 1886. Arthopyrenia epigloea (Nyl.) Zahlbr., Cat. lich. univ.<br />
1: 279. 1921. ≡ Pyrenocollema tremelloides Reinke, Jahrb. Wiss. Bot. 28: 461. 1886.Type.<br />
"Supra Nostoc saxa dolomitica incolens inundata fluminis Narenta in Herzegovinia (Lojka)",<br />
Lojka Lich. univ. 249 (MICH, NY, isotypes).<br />
This is the type of the genus but I far as I know the oldest epithet has not been combined in<br />
Pyrenocollema previously.<br />
Pyrenocollema farlowii (Riddle) R. C. Harris, comb. nov.<br />
Thelidium farlowii Riddle, Bull. Torrey Bot. Club 43: 147. 1916. Type. BERMUDA. 1881, Farlow<br />
(MICH, NY, isotypes?).<br />
Pyrenocollema prospersellum (Nyl.) R. C. Harris<br />
Sagedia nylanderi β. hudsoniana Hepp, Flecht. Eur. 945. 1867. Thelidium nylanderi var.<br />
hudsoniana (Hepp) Zahlbr., Cat. lich. univ. 1: 135. 1921. Type. [NEW JERSEY.] "Auf<br />
Serpentinblöcken am rechten Ufer des Hudson, in der Nähe von Hoboken bei Newyork",<br />
1861, Uhl (NY, US, isotypes).<br />
68
Pyrenocollema tichothecioides (Arn.) R. C. Harris s. lat.<br />
Since my thesis I have examined the type material of this species, the type of Xanthopyrenia Bachm.,<br />
and it is clearly congeneric with Pyrenocollema. This is where most of the obvious problems occur. Michigan<br />
material cited in my thesis has an ascospore size nearer that of P. strontianense (Swinscow) R. C. Harris but<br />
are on calcareous rock. Material reported from Louisiana on sandstone is not available but more recent<br />
collections seem to be better placed in or near Lichenothelia D. Hawksw. A recent collection on sandstone<br />
from Mississippi is closer to P. strontianense but apparently differs in asomatal characters. Specimens from<br />
the West seem to be closer to P. tichothecioides. Until a large amount of material is amassed and studied at<br />
one time, these problems cannot be resolved.<br />
Pyrenocollema zonatum (Müll. Arg.) R. C. Harris, comb. nov.<br />
Arthopyrenia zonata Müll. Arg., Flora 71: 549. 1888. Type. BRAZIL. "Saxicola ad flumen Iguape<br />
prope Iporanga in Brasilia meridionali", Puiggari 3025, 3027 (G, syntypes; BM, M, isosyntypes).<br />
ZWACKHIOMYCES Grube & Hafellner<br />
The species described here as Z. arenicola has the ascomatal pigmentation, physis type, ascospore<br />
type and lifestyle of Zwackhiomyces. Not having seen verified material of a real Zwackhiomyces, I am not<br />
sure about the ascus type. The apex is not in agreement with Grube's illustrations and the ascus is broadest<br />
at or below the middle. Also the ascospores have a diffuse gel sheath. I would not like to think another genus<br />
is needed but the possibility exists.<br />
1. Ascospores 28-37 × 10-13 µm; on unidentifiable areolate thallus or without<br />
obvious host; southern Georgia ......................................................................... [Z. arenicola R. C. Harris]<br />
1. Ascospores 24-27 × 7.5-8.5 µm; without obvious host but green algae present;<br />
Louisiana ..............................................................................................[Zwackhiomyces sp. Harris 11406]<br />
Zwackhiomyces arenicola R. C. Harris, sp. nov.<br />
NOTES<br />
Ascomatis in areolis lichenis ignoti vel super saxum arenaceum nudum et ascosporis 29-37 × 9-13 µm.<br />
Type. GEORGIA. Johnson County: 2 mi N of Adrian on Georgia Highway 15, 32°34'N, 82°35'W, ca. 80 m, small<br />
Altamaha Grit sandstone outcrop on slope, 18 Mar 1995, Harris 36282 (NY, holotype)<br />
Ascomata initially immersed in and then erupting through small, white, unidentifiable areoles on moist<br />
sandstone or growing sessile and naked directly on moist sandstone with a layer of green algae, subglobose,<br />
ca. 0.2-0.35 mm diam.; when growing immersed, ascomata often surrounded by a layer of paler brown<br />
hyphae, when growing exposed, ascomata black and glossy; ascomatal wall pale below when immersed,<br />
entirely dark when exposed. Asci elliptical to pyriform, ca. 110-135 × 30-45 µm, with eight irregularly arranged<br />
spores. Ascospores 28-37 × 10-13 µm, 2-celled, becoming 4-celled when overmature, ovoid, constricted at<br />
septum, becoming brownish and granular ornamented with age, with a watery gel sheath; upper cell slightly<br />
shorter and broader than the lower; pycnidia not found.<br />
All the known collections are from Altamaha Grit which is proving to be a habitat for many interesting<br />
lichens. The species is so inconspicuous that its absence from similar outcrops elsewhere in the South may<br />
be a collection (or lack of) phenomenon. A very similar species differing only in smaller ascospores occurs on<br />
sandstone in Louisiana. Assuming Z. arenicola is correctly assigned to genus, it is distinctive in the larger<br />
ascospores. Other species of Zwackhiomyces are reported by Grube (Nova Hedwigia 51: 283-360. 1990)<br />
from Clauzadea, Lecanora, Mycobilimbia and Porpidia. I am unable to determine the host for Z. arenicola,<br />
partly I believe, since the habitat is sufficiently wet that the lichen does not form a typical thallus.<br />
69
Additional speciemns. GEORGIA. Coffee County: Broxton Rocks Ecological Preserve, Ricketson Tract, ca.<br />
10 mi NE of Broxton, ca. 3 mi S of Ocmulgee River, 31°44'N, 82°44'W, ca. 70 m, extensive, dissected<br />
sandstone outcrop [Altamaha Grit], 4 Feb 1995, Buck 27461, 27495; Johnson County: as type, Buck 27570;<br />
Washington County: ca. 3 mi N of Harrison along County Road 206 (Peacock Road), near headwaters of<br />
Rocky Creek, 32°53'N, 82°44'W, ca. 140 m, Altamaha Grit sandstone flats, 18 Mar 1995, Buck 27543 (all NY).<br />
DACAMPIACEAE Körber<br />
EOPYRENULA R. C. Harris<br />
When I described this genus I placed it in the Pyrenulaceae, largely on the basis of IKI+ blue hymenial<br />
gel, long rod-shaped microconidia and brown ascospores. It was given the generic name with youthful<br />
naiveté, speculating that the genus might be ancestral to Pyrenula. Aptroot (1991) placed Eopyrenula in the<br />
Pleomassariaceae. This was accepted by Barr (Mycotaxon 49: 129-142. 1993). Eriksson (Lichenologist 25:<br />
310. 1993) was dubious about this placement. Personally I find it absurd. The type species of Pleomassaria<br />
and Splanchnonema have massive clavate asci with thick tips with indistinct, rounded ocular chambers ( with<br />
an apical nasse? in Pleomassaria siparia (Berk. & Br.) Sacc.), very coarse physes and large ascospores with<br />
thick gelatinous sheaths. Eopyrenula has smaller cylindrical asci with thin walls evenly thickened in water<br />
mounts, with a round or elongate ocular chamber in KOH, physes ca. ½ the width of those in<br />
Pleomassariaceae and medium sized ascospores without a gelatinous sheath. The only character I can see<br />
linking Eopyrenula to the Pleomassariaceae is the production of brown, septate macroconidia. Casting<br />
around for a better home for Eopyrenula, I ran across notes made for another purpose on Byssothecium<br />
Fuckel and Dacampia Massal., both placed in the Dacampiaceae. Eopyrenula has distinctive ascospores with<br />
the end cells paler than the middle cells. This seems to be due to a two layered spore wall, the inner pale, the<br />
outer dark. The outer layer apparently forms from the center outwards not covering the end cells which are<br />
therefore pale. This also seems to be the case in Byssothecium and possibly Dacampia, both of which have<br />
pale end cells. Both genera lack a gelatinous ascospore sheath as in Eopyrenula. The asci are ± cylindrical<br />
with ocular chamber similar to Eopyrenula. The pseudoparaphyses are slender as in Eopyrenula. For these<br />
reasons I suggest placing Eopyrenula in the Dacampiaceae. Barr described the ascomatal wall of<br />
Dacampiaceae as three layered with the outer pseudoparenchymatous. The ascomatal wall of Eopyrenula is<br />
of textura intricata. However, to my eye, the wall of Byssothecium circinans Fuckel also consists of textura<br />
intricata. Some Dacampiaceae have a pore in the ascospore septa which is not found in Eopyrenula. Also<br />
Eopyrenula-type macroconidia do not seem to be known from the Dacampiaceae. Even with these<br />
anomalies, Dacampiaceae seem to me a much better fit than Pleomassariaceae or Pyrenulaceae.<br />
Eopyrenula is well lichenized with Trentepohlia and would seem to represent another independent line of<br />
lichenization. In the Dacampiaceae Dacampia and Pyrenidium Nyl. are lichen parasites on a wide variety of<br />
hosts.<br />
There are three species of Eopyrenula known to me from North America (none of which occur in Florida),<br />
E. intermedia Coppins ex Aptroot in eastern North America, E. parvispora R. C. Harris & Aptroot in the Pacific<br />
Northwest and an apparently undescribed species on spruce from Minnesota. The potentially undescribed<br />
taxon may be the European E. leucoplaca (Wallr.) R. C. Harris as it has 2-celled macroconidia but the<br />
substrate is unique and I have almost no material of E. leucoplaca for comparison at this time.<br />
Type. Arthopyrenia Massal.<br />
ARTHOPYRENIACEAE W. Watson<br />
70
The family is defined here more narrowly than generally used in the last decade. The mechanics of this<br />
are discussed under Arthopyrenia and see also Naetrocymbaceae. Jarxia, Leptorhaphis and Naetrocymbe<br />
(Arthopyrenia rhyponta-punctiformis group) are removed to the Naetrocymbaceae. The restricted family is<br />
held together by clypeate ascomata, ascomatal wall KOH+ dark green [in the three species (incl. A. lapponina)<br />
studied by Foucard (1992); I have not assessed this character], pseudoparaphyses (cellular<br />
pseudoparaphyses), asci in a basal layer, asci fissitunicate, cylindrical to clavate with an apical nasse,<br />
ascospores colorless or brown, often granular ornamented, septum or initial septum submedian, 2-celled to<br />
muriform, pycnidia with rod shaped to filiform microconidia, occasionally associated with Trentepohlia, growing<br />
on bark, less commonly old wood and highest diversity in the tropics. The physes are often long and slender<br />
(easily distinguished from the sparse, coarse physes of the Naetrocymbaceae) and easily confused with<br />
paraphysoids (trabecular pseudoparaphyses) suggestive of the Melanommatales but they clearly taper from<br />
broad anastomosing bases. The cylindrical to clavate, stipitate asci (markedly obovate and nearly estipitate in<br />
Naetrocymbaceae) to some extent also suggest Melanommatales but the apical nasse would seem be<br />
anomalous in that order. Placement in the Pleosporales is also supported by the basal (rather than peripheral)<br />
origin of the asci. It does not seem unreasonable to include Julella in Arthopyrenia but more study is needed<br />
as there are no really clear intermediates and there are unresolved generic problems within Julella. I do not<br />
consider the vast majority of the family to be lichenized but a few, e.g., Arthopyrenia lyrata, seem consistently<br />
associated with Trentepohlia and perhaps continued treatment by lichenists is not inappropriate.<br />
1. Ascospores colorless ................................................................................................................................ 2<br />
2. Ascospores 2-4-celled.......................................................................................................Arthopyrenia<br />
2. Ascospores submuriform to muriform..........................................................................................Julella<br />
1. Ascospores brown, 2-celled to submuriform, usually granular ornamented ..................Mycomicrothelia<br />
ARTHOPYRENIA Massal.<br />
Ric. auton. lic. crost. 165. 1852. Lectotype (Fink, 1910). Arthopyrenia analepta (Ach.) Massal.<br />
≡ Leiophloea (Ach.) S. Gray, Nat. arr. Brit. pl. 1: 495. 1821. Verrucaria*Leiophloea Ach., Methodus<br />
Suppl. 24. 1803. Lectotype (Riedl, 1962). L. analepta (Ach.) S. Gray<br />
≡ Pyrenyllium Clements, Genera fungi 41, 173. 1909. Holotype. P. analeptum (Ach.) Clements<br />
Cifferiolichen Tomaselli in Tomaselli & Ciferri, Arch. Bot. (Forlì) 28: 4. 1952. ≡ Mycocifferia Tomaselli in<br />
Ciferri & Tomaselli, Ist. Bot. Reale Univ. Reale Lab. Crittog. Pavia Atti, ser. 5, 10: 28, 56. 1953. Holotype.<br />
Arthopyrenia lapponina Anzi (= A. analepta).<br />
In my unpublished thesis I made three major mistakes regarding Arthopyrenia, 1) Not splitting out the<br />
taxa here referred to Naetrocymbe (I came close, see Harris, 1975: 39), 2) Interpreting Lichen analeptus Ach.<br />
as illegitimate, 3) Accepting Riedl's lectotypification (If it is maintained, the nomenclatural changes consequent<br />
to the division of Arthopyrenia are distinctly unpleasant. Cifferiolichen is the next available name for the A.<br />
lapponina group.). As regards the first, I placed too much emphasis on microconidia. Arthopyrenia fraxini<br />
Massal. has the longer rod-shaped microconidia typical of A. analepta and I thought it represented a link<br />
between the two genera. It does not. All the other characters are typical for Naetrocymbe. The microconidia<br />
are anomalous. (With 20 years of retrospect the ascus and hamathecium characters seem extremely strong<br />
and necessitate the recognition of two families as is done in this work.) This division of the genus reopens the<br />
can of worms involved in typifying Arthopyrenia. Lichen analeptus is not illegitimate even though Acharius cited<br />
Verrucaria olivacea Pers. in synonymy since the name Lichen olivaceus (Melanelia olivacea (L.) Essl.) had<br />
already been used in the genus Lichen in the Lichenographiae sueciae prodromus. This makes possible the<br />
acceptance of Fink's typification of the genus. In 1975 I accepted Rield's choice of A. rhyponta (Ach.) Massal.<br />
as a temporary expedient, thinking that I would change it as soon as possible and believing that no one else<br />
would seriously endorse such an atypical species as the generitype. I erred grievously on both counts. In<br />
1994 I was finally able to study the relevant Acharian material in Helsinki. When I discovered that a specimen<br />
of the taxon currently known as A. lapponina was the obvious lectotype for A. analepta, the way was cleared to<br />
71
preserve Arthopyrenia in the broadest possible sense. Although Fink's typification may have been<br />
mechanical, it is not required that it be rejected if it is the most logical, preserving current usage. [If this is<br />
unacceptable to some, then consider this a proposal to overturn Riedl's lectotype and to reselect A. analepta.]<br />
The problem remains to decide whether Acharius was describing a new species or creating a new name for<br />
Verrucaria olivacea Pers. There is no direct indication in his work itself. In the interests of stability, I have<br />
chosen to consider Lichen analeptus Ach. as a new species. If this is totally unpalatable, then, since<br />
Arthopyrenia in this sense would require conservation no matter what (see below), it could be lectotypified with<br />
A. cerasi (Schrader) Massal. which would have the same effect. There is an appropriate Schrader collection<br />
in H-Ach of A. cerasi in the accepted sense. Either way the bulk of the taxa in temperate, subtropical and<br />
tropical regions are retained in Arthopyrenia. (In North America seven taxa are transferrred to Naetrocymbe<br />
but only N. punctiformis is represented by more than a few records so that the real effect is very minimal.)<br />
Arthopyrenia cinchonae is one of the most commonly collected pansubtropical/pantropical species, likewise A.<br />
analepta (syn. A. lapponina) in temperate regions and A. plumbaria in the Pacific Northwest, etc. In boreal<br />
regions Naetrocymbe is perhaps more important and names will have to change in the British and<br />
Scandinavian mycotas unless it is decided to ignore the differences and retain a broad view of Arthopyrenia.<br />
(Arthopyrenia seems a possible test case for the meaning of "Names in Common Usage". Will a small area<br />
with an essentially depauperate (but "well known") mycota sieze precedence over the rest of the world to<br />
retain Arthopyrenia in the sense of Naetrocymbe?) This typification allows the family name Arthopyreniaceae<br />
to be retained for three large well known genera, Arthopyrenia, Julella and Mycomicrothelia, that have been<br />
assigned to this family for some years already. This is also an "economical" solution as it does not require<br />
many new names, especially at the familial and generic levels (except for the validation of the previously<br />
proposed Naetrocymbaceae). A side benefit of realizing Lichen analeptus is not illegitimate and obligately<br />
typified with Verrucaria olivacea Pers. [= Porina s. lat., V. olivacea is not to be typified by the type of the<br />
misapplied name cited by Persoon (Art. 33 Note 2) as Hawksworth (1985: 133) proposed] is that Leiophloea<br />
(Ach.) S. Gray (1821) and Pyrenillium Clements (1909) do not have priority over Pseudosagedia (Müll. Arg.)<br />
M. Choisy (1949), recently resurrected by Hafellner and Kalb. However, since Riedl (1962) lectotypified<br />
Leiophloea with Lichen analeptus, Arthopyrenia will have to be conserved over Leiophloea or A. cerasi chosen<br />
as conserved type.<br />
In my thesis I included a small group of species in Arthopyrenia with some hesitation at the end of the<br />
genus as the "Arthopyrenia bifera group". The ascospores were distinctive in having ring-like wall thickenings<br />
forming partial subdivisions of the cells (and in postmature ascospores often ultimately a complete septum).<br />
Aptroot refers to this type as "biocellate", Vainio as "porospore". Over the succeeding years I have come to<br />
the conclusion that this is not significant but is in fact a more extreme expression of the general tendency for<br />
the ascospore cells to be medially constricted in Arthopyrenia. This is most conspicuous in A.<br />
cinereopruinosa, A. lyrata and A. plumbaria but shows up to some extent even in such "typical" species as A.<br />
analepta and A. cinchonae. Within the Arthopyreniaceae one might wish to give this ascospore type generic<br />
recognition but there is no correlation with any other feature and the other characters agree with those of<br />
Arthopyrenia, so I prefer to maintain a single genus. The same ascospore type occurs also in<br />
Mycomicrothelia suggesting that it may be a familial tendency.<br />
KEY TO ARTHOPYRENIA IN NORTH AMERICA<br />
1. Ascospores 2-celled (rarely 4-celled when postmature) ........................................................................... 2<br />
2. Ascospore cells not pinched in nor wall with internal ring-like thickenings............................................ 3<br />
3. Ascospores narrowly ovate or narrowly elliptical, length/width ratio<br />
2.5:1 or greater................................................................................................................................. 4<br />
4. Ascospores mostly more than 20 × 7 µm..................................................................................... 5<br />
5. Ascomata superficial, complanate to hemispherical, with or without<br />
72
a basal fringe; ascospores 20-30 × 7-9.5(-11) µm; microconidia<br />
4-5 × 0.5 µm; on a wide variety of smooth barks, very common,<br />
throughout Florida ................................................................... A. cinchonae (Ach.) Müll. Arg.<br />
5. Ascomata immersed, with a broad clypeus; ascospores 20-28(-32) ×<br />
6-8(-9.5) µm; microconidia 7-8 × 0.5 µm; on Taxodium; Dade and<br />
Seminole counties ................................................................................. A. taxodii R. C. Harris<br />
4. Ascospores mostly less than 23 × 7.5 µm.................................................................................... 6<br />
6. Ascospores 15-23 × 5-7.5 µm .................................................................................................. 7<br />
7. Ascomata usually complanate with a broad clypeus, often fusing<br />
with others; ascospores 17-23 × 5-7 µm; microconidia arcuate,<br />
20-27 × 0.5 µm; subtropical; Dade, Duval, Monroe and Sarasota<br />
counties .................................................................................A. planorbis (Ach.) Müll. Arg.<br />
7. Ascomata hemispherical to subglobose, with weak basal fringe;<br />
ascospores 15-22 × 5.5-7.5 µm; microconidia 7-10 × 0.5 µm;<br />
Massachusetts, Newfoundland, Europe.....................................[A. analepta (Ach.) Massal.]<br />
6. Ascospores 12-17 × 4-5.5 µm; microconidia 4-7 × 0.5 µm; rare,<br />
Liberty County ........................................................................................A. minor R. C. Harris<br />
3. Ascospores ovate, length/width ratio 1.5-2:1; 13-15 × 7-8(-10) µm;<br />
cells markedly unequal; microconidia 8-10 × 0.5 µm; Lake, Levy,<br />
Seminole and Wakulla counties...............................................................A. oblongens R. C. Harris<br />
2. Ascospore cell(s) pinched in ± mid cell or wall with internal ring-like thickenings<br />
as if additional septa were forming, often near apices (ultimately 4-celled in<br />
postmature stage) ................................................................................................................................. 8<br />
8. Ascospore wall with granular ornamentation, often with an internal<br />
ring-like thickening; southern ............................................................................................................ 9<br />
9. Ascospores 4-8/ascus................................................................................................................. 10<br />
10. Ascospore wall usually without internal thickening, merely<br />
constricted; ascospores only very rarely 4-celled when postmature,<br />
18-30 × 6.5-9.5(-12) µm; microconidia 6-9 × 0.5 µm; always?<br />
associated with Trentepohlia; throughout Florida...................................... A. lyrata R. C. Harris<br />
10. Ascospore wall with an internal ring-like ridge in both cells<br />
which forms a complete septum in postmature spores;<br />
ascospores 27-37 × 10-12 µm; microconidia 6-8 × 0.5 µm;<br />
Monroe County...............................................................................A. majuscula (Nyl.) Zahlbr.<br />
9. Ascospores 2/ascus, 37-48 × 15-16 µm; ostiole occasionally<br />
eccentric; microconidia 5-10 × 0.5 µm; common, throughout<br />
Florida ..............................................................................................A. malaccitula (Nyl.) Zahlbr.<br />
8. Ascospore wall smooth and lacking any internal thickening; northern............................................ 11<br />
11. Ascospores smaller, 12-15(-18) × 4-5 µm; spore cells weakly<br />
constricted; microconidia 6-8 × 1 µm; Massachusetts, Tennessee and<br />
73
West Virginia ...............................................................................................[A. degelii R. C. Harris]<br />
11. Ascospores (13-)15-26 × 4.5-9 µm; cells usually with stong constrictions................................ 12<br />
12. Ascospores with both ends rounded, 18-22 × 7-8(-9) µm; upper<br />
cell usually somewhat larger; asci 60-80 × 17-22 µm; microconidia<br />
10-12 × 0.5 µm; rare, Vermont, Newfoundland, California and<br />
Oregon .........................................................................[A. cinereopruinosa (Schaerer) Massal.]<br />
12. Ascospores with one or both ends pointed, (13-)15-20 × 4.5-6.5(-7.5) µm;<br />
cells ± equal; asci 75-110 × 12-18 µm; microconidia 10-12(-15) × 0.5 µm;<br />
Pacific Northwest...............................................................[A. plumbaria (Stizenb.) R. C. Harris]<br />
1. Ascospores 4-celled................................................................................................................................ 13<br />
13. Ascospores less than 25 µm long ..................................................................................................... 14<br />
14. Ascomata immersed, the tip surrounded by a broad clypeus often<br />
confluent with others; ostiole surrounded by a whitish ring;<br />
ascospores 17-22 × 6-8 µm; microconidia 6-9 × 0.5 µm;<br />
Dade and Monroe counties ......................................................................A. confluens R. C. Harris<br />
14. Ascomata lacking a broad clypeus, not confluent; ostiole not<br />
surrounded by a whitish ring; ascospores 17-23 × 6-7.5 µm;<br />
microconidia 10-13 × 0.5 µm; reported from British Columbia<br />
by Noble et al., 1987............................................................................[A. cerasi (Schrader) Massal.]<br />
13. Ascospores 28-40(-45) × 9-12(-15) µm, with distinct granular ornamentation<br />
and thick gelatinous epispore; microconidia 9-12 × 0.5 µm; Collier, Dade<br />
and Monroe counties ................................................................................. A. exasperata R. C. Harris<br />
NOTES<br />
Much of the information for this genus comes from my unpulished thesis (1975). To avoid possible<br />
confusion I have tried to account for all of the names included in it. The synonymy from the thesis and that<br />
accumulated in the 20 years since is also given.<br />
Arthopyrenia analepta (Ach.) Massal.<br />
Ric. auton. lic. crost. 165. 1852. Lichen analeptus Ach., Lichenogr. suec. prodr. 15. 1798. Type.<br />
SWEDEN. (H-Ach 796-B, lectotype, selected here).<br />
Arthopyrenia lapponina Anzi, Comment. Soc. Crittogam. Ital. 2: 25. 1864. Type. ITALY. Bormio,<br />
Anzi, Lich. Lang. 347 (FH-Tuck 4084, isotype).<br />
Verrucaria epidermidis var. fallax Nyl, Bot. Notiser 1852: 178. 1852. Verrucaria fallax (Nyl.) Nyl.,<br />
Flora 55: 363. 1872. Arthopyrenia fallax (Nyl.) Arnold, Verhandl. Zool.-Bot. Ges. Wien 23:<br />
505. 1873. Type. FINLAND. Holmia, Nylander (H-Nyl 955, holotype).<br />
Pyrenula punctiformis var. fallax f. betulae Hepp, Flecht. Eur. 452. 1857. Type. SWITZERLAND.<br />
Zürich, Hepp (FH, isotype)<br />
Arthopyrenia fallax f. crategina Steiner in Zahlbr., Ann. Naturhist. Hofmus. Wien 24: 283. 1911.<br />
Type. YUGOSLAVIA. "Ad litus meridionalis lacus "Veldser See", Steiner, Krypt. Vind. 1763a<br />
(FH, US, isotypes).<br />
The material in the Acharian herbarium under Verrucaria analepta contains three pieces of bark labelled<br />
"Suecia" (marked A, B, C) and therefore candidates for the lectotype. Pieces A and C lack ascospores but the<br />
hamathecium is of the "Naetrocymbe-type". Piece C also has microconidia in accord with Naetrocymbe.<br />
Piece B has hamathecium, asci, ascospores and microconidia identical to those of the species currently<br />
known as Arthopyrenia lapponina. I have selected Piece B as the lectotype as it contains good material of<br />
both teleomorph and anamorph and to stabilize the usage of Arthopyrenia in the broadest possible sense (see<br />
74
above). The selection of piece B is further endorsed by the fact that the only specimen cited by Massalongo<br />
under A. analepta (Schaerer, Lich. Helv. 287) is also this species. Some probably have not gotten over the<br />
loss of the epithet "fallax" and may be upset by this further name change but I remind them that the epithet<br />
"analepta" was widely used in the past in this sense. (Of the three older European collections at NY not<br />
previously annotated (Czechoslovakia, England and Switzerland), two were this species and one A.<br />
cinereopruinosa.)<br />
Arthopyrenia annulata R. C. Harris ined. = A. majuscula<br />
Arthopyrenia antecellens (Nyl.) Arnold<br />
Transferred in this publication to Mycoporum.<br />
Arthopyrenia atomarioides Müll. Arg. = Naetrocymbe<br />
Arthopyrenia atractospora Zahlbr. = Naetrocymbe<br />
Arthopyrenia bifera Zahlbr. = A. malaccitula<br />
Arthopyrenia cedrina (Zahlbr.) ined. = Naetrocymbe<br />
Arthopyrenia cerasi (Schrader) Massal.<br />
Ric. auton. lic. crost. 167. 1852. Verrucaria cerasi Schrader, Ann. bot. (Usteri) 22: 86. 1797. Type<br />
not seen.<br />
Arthopyrenia cineropruinosa (Schaerer) Massl.<br />
Symm. lich. nov. 117. 1855. Verrucaria cinereopruinosa Schaerer, Lich.helv. spic. sect. 6, 343.<br />
1836. Type not seen.<br />
Pyrenula punctiformis var. cinereopruinosa f. hederae Hepp, Flecht. Eur. 105. 1853. Type.<br />
SWITZERLAND. Zürich, Hepp (FH, FH-Tuck 4073, isotypes).<br />
Pyrenula punctiformis var. cinereopruinosa f. pinicola Hepp, Flecht. Eur. 106. 1853. Arthopyrenia<br />
pinicola (Hepp) Massal., Symm. lich. nov. 118. 1855. Type. SWITZERLAND. Zürich, Hepp<br />
(FH, FH-Tuck 4073, isotypes).<br />
Arthopyrenia stigmatella var. elabens Massal., Symm. lich. nov. 120. 1855. Type. ITALY. "Ad<br />
truncos pinorum in Prov. Patavinia (Rua)", Massal. Lich. Ital. 202 (FH, FH-Tuck 4077, isotypes).<br />
Pyrenula punctiformis var. cinereopruinosa f. buxicola Hepp ex Rabenh., Lich. Eur. 630. 1862,<br />
nom. inval. sine descr.<br />
Arthopyrenia convexella Müll. Arg., Flora 66: 288. 1883. Type. BRAZIL. São Paulo: Apiaí, Dec<br />
1879, Puiggari 1004 (G, holotype).<br />
Arthopyrenia transposita Müll. Arg., Bull. Soc. Bot. Belgique 31: 39. 1892. Type. NEW ZEALAND.<br />
1883, Knight (G, holotype).<br />
Arthopyrenia fallax var. conspurcata Steiner in Beck & Zahlbr., Ann. Naturhist. Hofmus. Wien 12:<br />
94. 1898. Type. AUSTRIA. Carinthia, near Klagenfurt, Steiner, Krypt. Vind. 269 (US, isotype).<br />
Arthopyrenia ligustri Britz., Hedwigia 43 (Beibl.): (37). 1904, hom. illeg. Type. GERMANY.<br />
Augsburg, westliche Hugelzug, Britzelmayer (US, isotype).<br />
Arthopyrenia japonica Vainio, Bot. Mag. (Tokyo) 32: 163. 1918. Type. JAPAN. Izumo Prov., 1<br />
Mar 1914, Yasuda 101 (TUR-Vainio 32112, holotype).<br />
Arthopyrenia cinchonae (Ach.) Müll. Arg.<br />
Flora 66: 287. 1883. Verrucaria cinchonae Ach., Syn. meth. lich. 90. 1814. Type. AMERICA. On<br />
Cinchona officinalis (H-Ach, holotype).<br />
Verrucaria prostans Mont., Ann. Sci. Nat. Bot. sér. 2, 19: 53. 1843. Type. FRENCH GUIANA.<br />
Leprieur 215 (PC-Mont, holotype).<br />
Verrucaria alboatra var. detergens Nyl., Flora 52: 125. 1869. Type. BRAZIL. Prope Rio de Janeiro,<br />
Glaziou 1915 (M, isotype).<br />
75
Verrucaria concamerata Stirton, Proc. Phil. Soc. Glasgow 13: 192. 1881. Porina concamerata<br />
(Stirton) Zahlbr., Cat. lich. univ. 1: 377. 1922. Type. ASSAM. Watt (BM, isotype).<br />
Arthopyrenia nieteriana Müll. Arg., Flora 66: 288. 1883. Type. CEYLON. Nieter 30, ex hb. Hampe,<br />
1877 (G, holotype).<br />
Arthopyrenia planipes Müll. Arg., Flora 73: 345. 1890. Type. AFRICA TROPICO-ORIENTALI. Rev. J.<br />
Hannington (BM ex K, holotype).<br />
As indicated by the number of synonyms and their wide distribution A. cinchonae is common,<br />
conspicuous and widespread. It is one of the most common species of the oak-pine woods of Florida, mainly<br />
on oak but not at all particular as to substrate. Over the years I have seen hundreds of specimens from drier<br />
areas of both Neo- and Paleotropics.<br />
Arthopyrenia confluens R. C. Harris, nom. nov.<br />
Tomasellia (sect. Oligomeris) leucostoma Müll. Arg., Flora 68: 257. 1885. [non Arthopyrenia<br />
leucostoma (Ach.) Massal. (1852)] Type. "in Cort. Cascarillae ex hb. Hampe, 1877". (G,<br />
holotype).<br />
Arthopyrenia degelii R. C. Harris, sp. nov.<br />
Ascis sat parvis, 48-60 × 11-13 µm et ascosporis parvis, bilocularibus, uno cellulo vel ambis cellulis in<br />
medio constrictis, 12-15 × 4-5 µm.<br />
Type. TENNESSEE. Great Smoky Mountains, Cherokee Orchard, on Hamamelis, 11 Sep 1939, Degelius (US,<br />
holotype; FH, US, isotypes).<br />
Forming brown, rarely tan, blotches on smooth bark, often Hamamelis. Ascomata superficial,<br />
hemispherical to complanate, 0.3-0.5 mm diam., clypeate, not melanized below. Asci narrowly elliptical, 48-60<br />
× 11-13 µm, with 8 biseriate to subbiseriate spores. Ascospores narrowly elliptical, 2-celled, on or both cells<br />
constricted near the middle (spore outline sinuose), 12-15(-18) × 4-5 µm, with a thin sheath. Microconidia rodshaped,<br />
6-8 × 1 µm.<br />
Arthopyrenia degelii is characterized by the sinuose ascospore outline, small ascospores, small, narrow<br />
asci and small hemispherical ascomata. Arthopyrenia minor is similar in ascospore size but the ascospores<br />
do not have median constrictions. Arthopyrenia degelii seems ± Appalachian in distribution.<br />
Additional specimens. MASSACHUSETTS. Bristol County: New Bedford, Willey (US). NORTH CAROLINA.<br />
Gaston County: Crowder's Mountain, 11 Aug 1886, Green (NY); Haywood County: Great Smoky Mtns. Natl.<br />
Park, at Big Creek, 45 mi ESE of Knoxville, 2000-2300 ft, on Liriodendron, 25 Aug 1977, Wetmore 29846<br />
(MIN, NY); Macon/Jackson County: Whiteside Mountain, ca. 5.5 mi E of Highlands, 4400-4930 ft, on<br />
Hamamelis, 10 Jun 1981, Harris 13756 (NY). TENNESSEE. Carter County: Doe River Gorge, ca. 1 mi N of U.S.<br />
Hwy. 19E, ca. 5 mi NW of town of Roan Mountain, ca. 800 m, on Hamamelis, 26 Sep 1993, Harris 30978<br />
(NY). WEST VIRGINIA. sine loc., on Hamamelis, Calkins, (FH, NY); Fayette County: sine loc., on Hamamelis,<br />
Nuttall 85 (FH), Nuttall s.n. (MICH, NY); Greenbrier County: Greenbrier Run, 1 Jun 1880, coll. unknown (FH);<br />
Taylor County: Grafton, on Hamamelis, 17 Dec 1880, coll. unknown (MSC).<br />
Arthopyrenia esenbeckiana (Fée) R. C. Harris, comb. nov.<br />
Melanotheca esenbeckiana Fée, Essai crypt. écorc. suppl. 74. 1837. Tomasellia esenbeckiana<br />
(Fée) Müll. Arg. Type. ANTILLES. on Exostemma, (G, lectotype, selected here).<br />
I am still of the opinion that this species does not occur in North America as I suggested in Egan (1987).<br />
It has nothing to do with either Melanotheca or Tomasellia. The characters of microconidia, ascus and<br />
hamathecium suggest placement in Arthopyrenia as defined in this work. The compound ascoma is<br />
anomalous although A. planorbis (Ach.) Müll. Arg. has a strong tendency for the clypei to fuse laterally. If one<br />
saw only ascospores and asci, one would identify it as A. cinchonae (Ach.) Müll. Arg.<br />
76
Arthopyrenia exasperata R. C. Harris, sp. nov.<br />
Primo aspectu Polymeridi catapasti similis sed differt ascosporis exasperatis, 36-44 ×<br />
12.5-15 µm, perisporio crasso et microconidiis longioribus, 9-12 × 0.5 µm.<br />
Type. FLORIDA. Collier County: Big Cypress National Preserve, Oasis Ranger Station along U.S. Hwy. 41,<br />
along Florida Trail N of hwy., 25°50'N, 81°03'W, Taxodium swamp, 9 Dec 1992, Buck 22976 (NY, holotype).<br />
Forming whitish patches on bark. Ascomata hemispherical to subglobose, mostly immersed, 0.4-0.6 mm<br />
diam., with slightly spreading clypeus or not, little or not melanized below. Asci cylindrical, 140-160 × 20-28<br />
µm, with 8 biseriate to nearly uniseriate spores. Ascospores narrowly elliptical to narrowly ovate, 4-celled, 28-<br />
40(-45) × 9-12(-15) µm, distinctly granular ornamented, with a thick gelatinous sheath. Microconidia 9-12 ×<br />
0.5 µm.<br />
In 1975 I placed this species in Polymeridium (Trypetheliaceae) but when I revised that genus (Harris,<br />
1993) I realized that it did not have Trypetheliaceae type of ascus and hamathecium but that of<br />
Arthopyreniaceae. In 1975 I designated a different type collection but since I have not re-examined the<br />
material at MSC, I have chosen a recently collected and studied specimen. There is some possibility that the<br />
MSC material is mixed and the specimens cited below need to be restudied.<br />
Additional specimens. FLORIDA. Dade County: Everglades National Park, NE of Pa-hay-okee Overlook,<br />
Harris 2814-B, 2829-A, 2836-C (MSC); Monroe County: Key West, Jan 1898, Thaxter 122 (MICH).<br />
Arthopyrenia fraxini Massal. = Naetrocymbe<br />
Arthopyrenia lapponina Anzi = Arthopyrenia analepta<br />
Arthopyrenia lyrata R. C. Harris<br />
in Tucker & Harris, Bryologist 83: 6. 1980.<br />
Arthopyrenia majuscula (Nyl.) Zahlbr.<br />
Cat. lich. univ. 1: 332. 1922. Verrucaria majuscula Nyl., Ann. Sci. Nat. Bot. sér. 4, 20: 253. 1864.<br />
Type. BONIN ISLANDS. Wright (H-Nyl 824, holotype; BM, isotype).<br />
This was treated in my thesis as Arthopyrenia annulata ined. Any specimens so annotated are to be<br />
referred here.<br />
Arthopyrenia malaccitula (Nyl.) Zahlbr.<br />
Cat. lich. univ. 1: 284. 1921. Verrucaria malaccitula Nyl., J. Linn. Soc., Bot. 20: 61. 1883. Type.<br />
MALACCA. Water Islands, 1864, Maingay 191 (H-Nyl 1706, holotype; BM, isotype).<br />
Arthopyrenia bifera Zahlbr., Ann. Mycol. 33: 34. 1935. Type. FLORIDA. [Seminole County:] Sanford,<br />
27 Apr 1929, Rapp 80 [publ. as 86] (W, holotype).<br />
Didymella gigantea Räsänen, Arch. Soc. Zool. Bot. Fenn. "Vanamo" 3: 89. 1949. Type. BORNEO.<br />
Sandakan, Myburgh, 1921, Elmer 20019 (H, lectotype, selected here; H, isolectotype).<br />
Arthopyrenia megalospora Lönnr. = Naetrocymbe<br />
Arthopyrenia minor R. C. Harris<br />
in Tucker & Harris, Bryolgoist 83: 7. 1980.<br />
Arthopyrenia oblongens R. C. Harris, sp. nov.<br />
Species distincta sporis ovatis, bilocularibus, cellulis insigniter inaequalibus, 13-15 × 7-8 µm.<br />
77
Type. FLORIDA. Sanford, Apr 1930, Rapp (MICH, holotype; LD, isotype).<br />
Forming a whitish blotch on bark of ericaceous shrubs. Ascomata ± subglobose, immersed, 0.2-0.3 mm<br />
diam., with a broad, elongate clypeus, 0.3-0.5 × 0.7-1.0 mm, little or not melanized below. Asci narrowly<br />
elliptical to cylindrical, 75-110 × 14-16 µm, with 8 uniseriate to subbiseriate spores. Ascospores ovate, 2celled,<br />
13-15(-17) × 7-8.5(-10) µm, smooth; cells markedly unequal, the lower narrower and shorter.<br />
Microconidia rod-shaped, 8-12 × 0.5-1 µm.<br />
The broad ascospores with strongly unequal cells are anomalous in Arthopyrenia and more suggestive of<br />
Anisomeridium but hamathecium, ascus and microconidium type place it here. It may be possibly related to A.<br />
analepta having evolved unusually broad ascospores. As far as I can tell A. oblongens is restricted to<br />
ericaceous substrates and is known only from Florida.<br />
Additional specimens. FLORIDA. Lake County: Ocala National Forest, ca. 2 mi W of Alexander Springs,<br />
dwarf oak scrub, on Lyonia, 6 Dec 1988, Harris 23620 (NY); Levy County: Cedar Key Scrub State Preserve,<br />
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<br />
22421, 22423, 22453 (NY), Harris 29357 (FLAS, NY); Seminole County: Sanford, 28 Jun 1928, Rapp 709<br />
(MICH); Wakulla County: Apalachicola National Forest, Bradwell Bay Wilderness, along Apalachicola Trail at<br />
Sopchoppy River, 29 Nov 1988, Harris 22979 (NY).<br />
Arthopyrenia padi Rabenh. = Naetrocymbe punctiformis<br />
Arthopyrenia planorbis (Ach.) Müll. Arg.<br />
Mém. Soc. Phys. Genève 30: 27. 1888. Verrucaria planorbis Ach., Syn. meth. lich. 92. 1814.<br />
Type. "in America ad Crotonis Cascarillae". (H-Ach, holotype).<br />
Arthopyrenia planior Müll. Arg., Bot. Jahrb. Syst. 6: 404. 1885. Type. CUBA. Wright, Lich. Cub.<br />
II: 627a, 627b (G, syntypes), 627b (US, isosyntype).<br />
Arthopyrenia planorbiculata Müll. Arg., Bot. Jahrb. Syst. 6: 405. 1885. Type. CUBA. Wright, Verr.<br />
Cub. 64 (G, holotype; US, isotype).<br />
Arthopyrenia plumbaria (Stizenb.) R. C. Harris<br />
in Egan, Bryologist 90: 163. 1987. Verrucaria plumbaria Stizenb. in Hasse, Erythrea 3: 44. 1895.<br />
Porina plumbaria (Stizenb.) Hasse, Contr. U. S. Natl. Herb. 17: 12. 1913. Type not seen. Pyrenula<br />
herrei Fink in Hedrick, Mycologia 25: 309. 1933. Type. CALIFORNIA. Santa Cruz Peninsula, Herre<br />
(MICH, holotype).<br />
Arthopyrenia rappii Zahlbr., Ann. Mycol. 33: 34. 1935. Mycoporopsis rappii (Zahlbr.) Riedl, Sydowia<br />
16: 272. 1962[1963]. Type. FLORIDA. Seminole County, Fort Reed, on Hicoria, Jan 1929, Rapp<br />
138 [publ. as 183] (W, holotype).<br />
This is a species of Melaspilea Nyl. s. lat.<br />
Arthopyrenia rhyponta (Ach.) Massal. = Naetrocymbe<br />
Arthopyrenia salicis Massal.<br />
Position uncertain. A related species is included in Naetrocymbe as a temporary expedient. Like<br />
Naetrocymbe it is not lichen-forming and the most satisfactory solution as far as I am concerned would be to<br />
exclude if from the checklist as having no relevance for lichenologists. I wish I had the "silver bullet" to kill off<br />
the perpetually "undead" irrelevancies in checklists.<br />
Arthopyrenia taxodii R. C. Harris, sp. nov.<br />
Sat similis Arthopyreniae cinchonae quoad ascos et ascosporas sed differt ascomatis in corticem Taxodii<br />
78
immersis clypeo lato et microconidiis 7-8 × 0.5 µm.<br />
Type. TEXAS. Houston, on Taxodium distichum, 1869, Ravenel 60 (FH-Tuck 4022, holotype; US, isotype; FH,<br />
MICH, PH, US, isotypes?).<br />
Forming a whitish blotch on bark of Taxodium. Ascomata subglobose, immersed, 0.2-0.3 mm diam. with<br />
a broad clypeus; wall poorly melanized. Asci cylindrical, 100-130 × 14-20 µm, with 8 nearly uniseriate to<br />
subbiseriate spores. Ascospores narrowly ovate to narrowly elliptical, 20-28(-32) × 6-8(-9.5) µm, smooth.<br />
Microconidia 7-8 × 0.5 µm.<br />
Arthopyrenia taxodii is distinctive in the immersed ascomata and substrate. Most species of<br />
Arthopyrenia occur on smooth bark of hardwoods. It seems closer to the "Peltosphaeria" group of Julella than<br />
to the rest of Arthopyrenia.<br />
Additional specimens. FLORIDA. Duval County: Jacksonville, [Calkins] (MICH); Seminole County: Sanford,<br />
Rapp s.n. (FLAS), 11 May 1932, Rapp 29 (FLAS), Upsala, 11 May 1932, Rapp 2014 (FLAS, MICH).<br />
Arthopyrenia texensis (Cooke) D. Hawksw.<br />
I have not seen the specimen involved and since Hawksworth's concept of Arthopyrenia was very broad,<br />
I cannot place the species even as to genus.<br />
JULELLA Fabre<br />
Just as I was nearly finished with this volume, Aptroot and v.d. Boom published a paper with notes on<br />
Julella (Mycotaxon 56:1-8. 1995). We have reviewed essentially the same group of species and come up with<br />
very different results. Depending on one's view point, Aptroot and v.d. Boom have indulged in an orgy of<br />
"lumping" or I have indulged in an orgy of "splitting".<br />
As with other muriform-spored taxa, the possibility exists that Julella should be merged with its<br />
transversely septate sister taxon, Arthopyrenia. I am reluctant to take this step as Julella is probably<br />
heterogenous. For instance, I am sorely tempted to reinstate Peltosphaeria Berlese and some of the tropical<br />
taxa seem to form a relatively distinct clade. Further, I am not happy with the specific level taxonomy<br />
presented below. Many of the specimens come from dry, harsh habitats and "external" characters of clypeus<br />
and ascomatal immersion are highly variable. Also as with many muriform-spored taxa, one is never quite<br />
sure ascospore differences are due to genetics or age. Even worse, the microconidia, which I normally<br />
consider a highly conservative character, don't seem to provide much help. I suspect that there are more host<br />
specific microspecies but I do not now have either enough specimens or enough time to make sense of them.<br />
A few names that have been placed in Julella (or Peltosphaeria) whose types I have not examined are<br />
included in the key or commented on in the notes despite this basic ignorance.<br />
1. Asci with 8 ascospores (rarely 1-several aborting).................................................................................... 2<br />
2. Ascospores remaining colorless ........................................................................................................... 3<br />
3. Asci clavate to pyriform with biseriately to irregularly arranged spores.............................................4<br />
4. Asci narrower clavate or short-cylindrical...................................................................................... 5<br />
5. Ascospores 24 µm long or longer ............................................................................................. 6<br />
79
6. Microconidia known............................................................................................................... 7<br />
7. Microconidia filiform, curved, 16-27 µm long; ascospores<br />
24-34 × 8.5-11 µm, with 8-10 rows of cells; Collier, Polk<br />
and Washington counties......................................................... J. variiformis R. C. Harris<br />
7. Microconidia rod-shaped, 6-9 µm long; ascospores<br />
24-33 × 10-15 µm, with 8(-10) rows of cells;<br />
Dade County.................................................................................... J. asema R. C. Harris<br />
6. Microconidia not known; ascospores 18-32 × 10-15 µm,<br />
5-7 × 2-3-septate; Hawaiian Islands; no material seen ...........................................................<br />
..................................................................................... [Peltosphaeria sandwichensis Petrak]<br />
5. Ascospores 25 µm long or shorter ............................................................................................ 7<br />
7. Microconidia known............................................................................................................... 8<br />
8. Ascospores 22-25 × 11-14 µm, rather regularly 5-6 × 2-3-septate;<br />
microconidia 5-8 µm long; oceanic?, Europe .......................[J. sericea (Massal.) Coppins]<br />
8. Ascospores 17-23 × 7-9 µm, irregularly submuriform to muriform,<br />
4-6 × 1-2-septate; microconidia 6-10 µm long; a single questionable<br />
Calkins record from Duval County ...................................J. fallaciosa (Arn.) R. C. Harris<br />
7. Microconidia not known; ascospores 13-19 × 7-9 µm,<br />
3-5 × 1-2-septate; California; no material seen .................. [Peltosphaeria californica Petrak]<br />
4. Asci broad, mostly pyriform but occasionally long-ellipsoidal;<br />
ascospores 26-38 × 12-16 µm, with 8-10 rows of cells;<br />
microconidia 10-20 µm long; peninsular Florida ..........................J. sublactea (Nyl.) R. C. Harris<br />
3. Asci long-cylindrical, 200-300 × 20-25 µm with uniseriate spores..................................................... 9<br />
9. Ascospores larger, 24 × 9 µm or over ......................................................................................... 10<br />
10. Ascospores large, 29-46 × (9.5-)11-14 µm ........................................................................ 11<br />
11. Ascospores 36-46 × 12-13(-15) µm, rather irregularly<br />
7-11 × 1-2-septate, with large cells; microconidia 6-7 µm<br />
long; on bark of Taxodium; Collier, Dade, Duval and<br />
Wakulla counties .........................................................................J. taxodii R. C. Harris<br />
11. Ascospores 29-37 × 11-14 µm, regularly 7-11 × 2-4-septate,<br />
with smaller cells; microconidia 10-11 µm long; on decorticate<br />
stems of shrubs; California ............................... [J. vitrispora (Cooke & Harkness) Barr]<br />
10. Ascospores smaller, 24-30(-35) × 9-12 µm (NB: 3 choices below).................................... 12<br />
12. Microconidia 5-6 µm long; ascospores 24-30 × 9-11 µm,<br />
with 8-9 rows of cells; on Xanthoxylum; Flagler County...................Julella sp. 2485-B<br />
12. Microconidia 8-9 µm long; ascospores 24-35 × 11-12.5 µm,<br />
with 6-8 rows of cells; Namibia.................................................................[Harris 18533]<br />
12. Microconidia 15-17 µm, curved; ascospores 25-30 × 9-12 µm,<br />
with 7-8 rows of cells; Namibia.................................................................[Harris 18519]<br />
80
9. Ascospores small, 16-22 × 8-9 µm; ascomata on old wood, raised,<br />
± constricted at base; clypeus elongated in direction of grain;<br />
Dominican Republic ........................................................................................[Julella sp. 19512-A]<br />
2. Ascospores becoming brown, ± biseriate, 28-36 × 12-16 µm; ascomata<br />
pyriform, immersed; anamorph in culture a Phoma;on intertidal<br />
mangrove wood; India, Australia .............................................................[?J. avicenniae (Borse) Hyde]<br />
1. Asci with 2 or 4 ascospores..................................................................................................................... 13<br />
13. Asci with 2 ascopores........................................................................................................................ 14<br />
14. Ascospores over 35 × 15 µm ...................................................................................................... 15<br />
15. Growing on a wide variety of bark in dry habitats; ascoma not<br />
strongly immersed; clypei often fused; ostioles often surrounded<br />
by a whitish ring; ascospores 40-51 × 17-21 µm, with 9-11 rows<br />
of cells; Dade and Seminole counties, also "Caloosa River",<br />
Austin ...................................................................................J. geminella (Nyl.) R. C. Harris<br />
15. Growing on Buxus sempervirens in Europe; ascoma immersed;<br />
clypei may be fused; ascospores (26-)40-50 × (10-)16-20 µm,<br />
with (4-)16 rows of cells (Barr, 1986) .............................................................. [J. buxi Fabre]<br />
14. Ascospores 27-35 × 10-12.5 µm, with 8-9 rows of cells;<br />
microconidia 14-17 × 0.5 µm; Texas ....................................... [J. dispora (Müll. Arg.) R. C. Harris]<br />
13. Asci with 4 ascospores...................................................................................................................... 16<br />
16. Ostiole apical, colorless............................................................................................................... 17<br />
17. Ascospores 30-36 × 12-15 µm, with 7-9 rows of cells; ascomata<br />
± superficial, with narrow hyphal ring; microconidia 15-18 × 0.5 µm;<br />
eastern North America and Europe................................................... [J. lactea (Massal.) Barr]<br />
17. Ascospores 30-36 × 14-17 µm with 6 rows of cells; ascomata<br />
immersed under a broad hyphal ring (thin clypeus?); microconidia<br />
7.5-11 × 0.5 µm; Australia ................................................... [J. exiguella (Bailey) R. C. Harris]<br />
16. Ostiole lateral, red pigmented, olive brown in KOH; ascospores<br />
40-50(-62) × 15-18(-22) µm, with 8-10 rows of cells; Bahamas<br />
and Puerto Rico....................................................................... [J. declorans (Riddle) R. C. Harris]<br />
Julella asema R. C. Harris, sp. nov.<br />
NOTES<br />
Julellae sublacteae similis sed ascis angustioribus et J. variiformis sed ascosporis latioribus, 24-33 × 10-<br />
15 µm, 7(-9) × 2-4-septatis, microconidiis 7-9 µm longis.<br />
Type. FLORIDA. Dade County: Everglades National Park, just N of Paurotis Pond, Paurotis-buttonwood<br />
hammocks surrounded by red mangrove, on Rhizophora mangle, 11 May 1967, Harris 2663-A (NY, holotype).<br />
81
Thallus indicated by whitish discoloration of bark. Ascomata orbicular, initially immersed and flat,<br />
becoming hemispherical, 0.5 mm diam., with or whithout a dark surrounding clypeoid ring; ostiole sometimes<br />
surrounded with pale ring. Asci oval to clavate, with 8 ascospores ± biseriately arranged. Ascospores with one<br />
end narrower, often slightly constricted at median septum, 7(-9) × 2-4-septate, 24-33 × 10-15 µm, often with a<br />
gel sheath. Microconidia rod-shaped, 6-7 µm.<br />
This species is perhaps poorly defined and rather characterless which is reflected in the epithet meaning<br />
"without a mark or sign". In ascospore size it is not very different from J. sublactea and J. variiformis, J.<br />
sublactea differs in broad pyriform asci, J. variiformis has narrower ascospores, both have longer<br />
microconidia. Dominican and Venezuelan material seems to agree quite well.<br />
Additional specimens. FLORIDA. Dade County: Coconut Grove, on mango, 23 Dec 1919, Britton & Britton<br />
665. DOMINICAN REPUBLIC. Prov. La Romana: Isla Catalina, 0-10 m, 18 Dec 1987, Zanoni et al. 37318[D].<br />
VENEZUELA. Falcón: Península de Paraguaná, Cumbre de Cerro Colorado, zona xerófita, 200 m, López &<br />
Wingfield 22504 (all NY).<br />
Julella avicenniae (Borse) Hyde<br />
This species is retained in Julella with considerable hesitation. The brown ascospores and Phoma<br />
anamorph are anomalous.<br />
Julella dactylospora Rehm<br />
Hedwigia 40: 110. 1901.<br />
= Anthracothecium columellatum (Vainio) Zahlbr. fide Aptroot & v. d. Boom (1995).<br />
Julella decolorans (Riddle) R. C. Harris, comb. nov.<br />
Campylothelium decolorans Riddle in Britton & Millspaugh, Bahama Fl. 532. 1920. Type.<br />
BAHAMAS. Exuma Chain: Cay N of Wide Opening, on Drypetes, 18 Feb 1905, Britton &<br />
Millspaugh 2795 (FH, holotype; NY, isotype).<br />
The red pigmented ostiole and ± 4-spored asci make this a very distinctive species. The asci are initially<br />
8-spored but 4-6 routinely abort with most commonly 4 surviving although other numbers are encountered. It<br />
seems to be a dry forest species and I would expect it to turn up in Florida.<br />
Julella dispora (Müll. Arg.) R. C. Harris, comb. nov.<br />
Polyblastia dispora Müll. Arg., Flora 60: 79. 1877. Polyblastiopsis dispora (Müll. Arg.) Zahlbr., Cat.<br />
lich. univ. 1: 348. 1922. Type. TEXAS. Near Dallas, Boll (US, isotype).<br />
?Polyblastia fallaciuscula Müll. Arg., Bot. Jahrb. Syst. 6: 407. 1885. Polyblastiopsis fallaciuscula<br />
(Müll. Arg.) Zahlbr., Cat. lich. univ. 1: 349. 1922. Type. CUBA. Wright, Verr. Cub. 131.<br />
The fragment of the holotype of Polyblastia fallaciuscula I have seen lacked ascospores. It is tentatively<br />
assigned here based on the ascospore size given by Müller, 26-30 × 12-15 µm. Geographically it is more<br />
likely to be an immature specimen of J. geminella.<br />
Julella exiguella (Bailey) R. C. Harris, comb. nov.<br />
Trypethelium exiguellum Stirton ex Bailey, Queensland Agric. J. 5: 39. 1899. Type. AUSTRALIA.<br />
Queensland: Thursday Island, Jun 1897, Bailey s.n. (BM, holotype?).<br />
Julella fallaciosa (Arn.) R. C. Harris<br />
Mycoglaena yasudae Vainio, Bot. Mag. (Tokyo) 35: 75. 1921. Type. JAPAN. Prov. Kozuke, 7 May<br />
1918, Yasuda 238 (TUR-Vainio 31023), syn. nov.<br />
Julella geminella (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria geminella Nyl., Expos. syn. pyrenocarp. 40. 1858. Polyblastiopsis geminella (Nyl.)<br />
Zahlbr., Cat. lich. univ. 1: 349. 1922. Type. "Ad cortices in Mexico." (not seen).<br />
82
?Polyblastiopsis alboatra Zahlbr. in Rechinger, Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr.<br />
81: 230. 1907. Type. SAMOA. Upolu, auf Clerodendron im Mangrove Sumpfe bei Mulinuu,<br />
Rechinger 2797 (not seen).<br />
Polyblastiopsis rappii Zahlbr., Ann. Mycol. 33: 36. 1935. Type. FLORIDA. [Seminole County:]<br />
Sanford, on Amorpha fruticosa, Apr 1933, Rapp 133 (W, holotype; FLAS F-27978, isotype).<br />
Although I have not seen the types of Verrucaria geminella or Polyblastiopsis atroalba, the 2-spored asci<br />
and ascospore size suggest this disposition. Julella geminella is apparently pantropical, with specimens seen<br />
from Florida, Bermuda, Puerto Rico, Bequia, St. Lucia, Tanzania, India, Hong Kong and Papua New Guinea.<br />
Julella leopoldina Rehm, Ann. Mycol. 6: 319. 1908.<br />
From the description I suspect this is an Anthracothecium or Pyrenula.<br />
Julella luzonensis P. Henning<br />
Hedwigia 47: 257. 1908.<br />
= Pyrenula sp. (Harris, 1989) or Anthracothecium prasinum (Eschw.) R. C. Harris (Aptroot & v.d. Boom,<br />
1995). The type collection may be mixed and the typification needs to be carefully investigated.<br />
Julella macrospora Döbbler, Mitt. Bot. München 14: 191. 1978.<br />
Julella phycophila Döbbler, Mitt. Bot. München 14: 193. 1978.<br />
Both are described as being aparaphysate. They certainly have nothing to do with Julella. However, I<br />
am not aware that they have been transferred elsewhere.<br />
Julella manakonensis Kern, Ber. Schweiz Bot. Ges. 69: 282. 1959.<br />
From the description and illustrations I surmise that this is referable to either Anthracothecium or<br />
Pyrenula.<br />
Julella sericea (Massal.) Coppins<br />
Purvis et al. (1992) and Aptroot and v.d. Boom (1995) suggest that J. sericea might be synonymous with<br />
J. fallaciosa. From the few specimens of J. sericea (including Massal. 262) I have studied, the ascospores<br />
seem broader, more ellipsoidal, more divided and more regularly divided. The distribution seems<br />
Mediterranean/ oceanic. I am inclined to keep the taxa separate.<br />
Julella sublactea (Nyl.) R. C. Harris<br />
Verrucaria sublactea Nyl., Flora 69: 464. 1886. Type. CORFU. S. Nicolo[?], an junger Oelbäume,<br />
May 1886, P. Sydow s.n. (NY, isotype?).<br />
Polyblastia transwaalensis Müll. Arg., Flora 70: 79. 1887. Type. SOUTH AFRICA. Transvaal:<br />
Lydenburg, Wilms, comm. Lahm no. 52 (G, holotype).<br />
Mycoglaena collosporella Vainio, Bol. Soc. Broteriana, sér. 2, 6: 176. 1930. Type. MOZAMBIQUE.<br />
Palma, 1916, Pires de Lima 774 (TUR-Vainio 34328, holotype).<br />
Clathroporina exiguella Zahlbr., Ann. Mycol. 33: 38. 1935. Type. FLORIDA. [Seminole County:]<br />
Sanford, on Garberia fruticosa, Oct 1932, Rapp 119 (FLAS F-30032, isotype).<br />
Clathroporina amygdalina sensu Fink, Lich. Fl. U.S. 56. 1935.<br />
This species is distinctive in having a whitish thallus and long, unusually variable microconidia. I had<br />
originally thought that Polyblastia alba Müll. Arg. from South Africa was synonymous but the type has slightly<br />
smaller ascospores and microconidia only 8-10 µm long. It antedates Verrucaria sublactea by a few months<br />
and would have priority if my present stance turns out to be incorrect. McCarthy (1995) suggested that<br />
Clathroporina exiguella should be referred to Thelenella. One collection from Puerto Rico and one from<br />
Venezuela seem referable to P. alba.<br />
Julella taxodii R. C. Harris, sp. nov.<br />
Ascomatis immersis clypeis grandis, 1-1.5 mm diam., ascis anguste cylindraceis, 220-300 × 20-25 µm,<br />
ascosporis uniseriatis, 9-10 × 1-2-septatis, 36-46 × 12-15 µm, microconidiis bacilliformibus, 6-7 µm et ad<br />
Taxodium crescentibus.<br />
83
Type. FLORIDA. Dade County: Everglades National Park, cypress dome between Mahogany Hammock and<br />
Pa-hay-okee Overlook, on Taxodium, 16 May 1967, Harris 2947-A (NY, holotype).<br />
Thallus not evident or a whitish discoloration of the bark. Ascomata immersed to somewhat emergent<br />
with age, ± spherical, 0.45-0.7 mm diam., usually with a broad clypeus, 0.6-1.8 mm diam. Asci narrowly<br />
cylindrical, 200-300 × 20-25 µm, often with a long stipe, with 8 uniseriate ascospores. Ascospores fusiform,<br />
rather irregularly 9-10 × 1-2-septate with large cells, 36-46 × 12-13(-15) µm, often with an obvious gel sheath.<br />
Microconidia rod-shaped, 5-7 × 0.5 µm.<br />
Julella taxodii is conspicuous on Taxodium due to the very broad clypeus and is probably locally<br />
common. It was widely distributed by Calkina under the name Pyrenula subcynerea. Its nearest relative is<br />
probably J. vitrispora from which it differs in larger ascospores and shorter microconidia. There are two<br />
Greater Antillean collections which are very close, differing mainly in substrate. They are included here<br />
tentatively.<br />
Additional specimens. FLORIDA. Collier County: Big Cypress National Preserve, Oasis Ranger Station<br />
along U.S. Hwy. 41, along Florida Trail N of hwy., 9 Dec 1992, Buck 22957 (NY); Dade County: NE of Pa-hayokee<br />
Overlook, 3 May 1967, Harris 2819, 2832, S of Pa-hay-okee Overlook, 12 May 1967, Harris 2779-A<br />
(NY); Duval County: Jacksonville, Calkins (NY); Wakulla County: Appalachicola National Forest, along Co. Rd.<br />
368, 3.6 mi from Co. Rd. 319 at Crawfordsville, 29 Nov 1988, Harris 22948 (NY). GEORGIA. Ware County:<br />
Okefenokee National Wildlife Refuge, N end of Cedar Hammock Canal, 3 mi NW of boat landing, 25 Nov<br />
1989, Wetmore 65182 (MIN, NY), 1 mi W of Chesser Island along boat trail, 29 Nov 1989, Wetmore 65450<br />
(MIN) (all on Taxodium).<br />
Not on Taxodium. DOMINICAN REPUBLIC. Prov. Monte Cristi: El Morro de Monte Cristi, 237 m, limestone<br />
mesa by sea, 19°54'N, 71°39'W, 8 Jan 1987, Buck 13947 (NY). PUERTO RICO. Bosque Estatal de Guajataca,<br />
Vereda Juan Pérez near S end of reserve, 18°24'N, 66°58'W, ca. 300 m, humid forest, 17 Jan 1992, Harris<br />
27876 (NY).<br />
Julella variiformis R. C. Harris, sp. nov.<br />
Ascomatis hemisphaericis, ascosporis 25-33 × 9-10.5 µm, 7 × 1-3-septatis et microconidiis filiformibus,<br />
arcuatis, ca. 22 µm longis.<br />
Type. PUERTO RICO. Mun. Cabo Rojo: Sierra Bermeja, Cerro Mariquita, from W end of Laguna Cartagena to<br />
summit, on cactus, ca. 18°00'N, 67°07'W, 35-300 m, dry forest over chert, 15 Jan 1992, Harris 27732 (NY,<br />
holotype).<br />
Thallus indicated by whitish or grayish area of disrupted bark. Ascomata extremely variable, from forms<br />
with immersed ascomata and the clypeus flush with the bark surface and a whitish ring around the ostiole<br />
(young forms?) to forms ± superficial and hemispherical with a dark hyphal, clypeoid ring well developed to<br />
absent, on barks which erode occasionally and thus raised on pedestals of bark held together by the ascoma,<br />
black, centrum ca. 0.5 mm diam. but clypeus to 2 mm diam. and irregular groups of fused clypei to several<br />
mm; clypeus dense to diffuse hyphae in upper bark layer. Asci fusiform to ± clavate, with 8 mostly biseriate<br />
ascospores. Ascospores with long-ovoid or short-cylindric, 7 × 1-3-septate, 24-34 × 8.5-11 µm, often with an<br />
obvious gel sheath. Microconidia filiform, arcuate, (16-)22-27 × 0.5 µm.<br />
As noted above in this circumscription the ascomata are extremely variable and hence the specific<br />
epithet. The type is a form with superficial hemispherical ascomata. It is certainly possible that several<br />
microspecies are included. The characters that hold the putative taxon together are the filiform microconidia<br />
and the medium sized, rather narrow ascospores. Julella variiformis may be difficult to separate from J.<br />
asema without microconidia and one has to depend on ascospore width. Julella variiformis seems to be a<br />
species of exposed, often dry, scrub habitats.<br />
84
Additional specimens. FLORIDA. Collier County: Big Cypress National Preserve, Oasis Ranger Station on<br />
U.S. Hwy. 41, along Florida Trail N of hwy., on Bumelia, 9 Dec 1992, Harris 30153; Polk County: Nalcrest, 12<br />
Mar 1989, Wheeler s.n.; Washington County: Rock Hill Preserve, 4.1 mi SE of Chipley, Griffin 2919[A].<br />
BAHAMAS. Cat Isalnd: The Bight and Vicinity, on Coccoloba krugii, 1-6 Mar 1907, Britton & Millspaugh 5922.<br />
PUERTO RICO. Reserva Forestal Guánica, Bahía de la Ballena at Km 9.5 on Hwy. 333, ± sea level, dry coastal<br />
scrub over rough limestone, 26 May 1989, Harris 23984, 23985, 24013; Vereda El Ojo de Agua at Km 3.8 on<br />
Hwy. 334, ca. 600 ft, semi-dry scrub, 1 Jun 1989, Harris 24152; Maricao State Forest, Santana Ridge,<br />
secondary moist scrub, 27 May 1989, Harris 24044; Bosque Estatal de Susúa, S edge of reserve, ca. 275 m,<br />
dry scrub forest over exposed serpentine, 13 Jan 1992, Buck 21268; as type, Harris 27712-A (all NY).<br />
Jullela vitrispora (Cooke & Harkness) Barr<br />
E. Britton 4367 (NY) from Puerto Rico on Byrsonema has ascospores which fall into the range of J.<br />
vitrispora but differs in fewer, less regular longitudinal septations and in microconidia only 4-6 µm long. If<br />
future collections maintain these differences, it should be given taxonomic recognition.<br />
Julella zenkeriana P. Henning<br />
Bot. Jahrb. Syst. 38: 127. 1905.<br />
= Laurera megasperma (Mont.) Riddle fide Aptroot & v.d. Boom (1995).<br />
MYCOMICROTHELIA Keissler<br />
?Ornatopyrenis Aptroot, Biblioth. Lichenol. 44: 127. 1991. Type. Microthelia queenslandica Müll. Arg.<br />
Ornatopyrenis was assigned by Aptroot to the Trypetheliaceae. I have seen no material. The reasons<br />
for tentatively assigning it to synonymy of Mycomicrothelia are discussed in the introduction to the<br />
Trypetheliaceae.<br />
KEY TO FLORIDA SPECIES<br />
1. Ascospores 13.5-16 × 6-7 µm (Hawksworth, 1985); on a variety of trees,<br />
perhaps favoring Myrica;common, throughout Florida.................M. subfallens (Müll. Arg.) D. Hawksw.<br />
1. Ascospores 17-20(-21) × 7-8(-9) µm (Hawksworth, 1985); on Fraxinus;<br />
Duval, Liberty and Sarasota counties............................................ M. willeyana (Müll. Arg.) D. Hawksw.<br />
NOTES<br />
Mycomicrothelia oleosa Aptroot<br />
This species is unique in having an inspersed hymenium. Its range is extended to Brazil. Brako 5640<br />
shows some endospore development forming a ring near the ends of the ascospores forming a rounded<br />
chamber as in some species of the sister genus Arthopyrenia. In Arthopyrenia these endospore rings often go<br />
on to form septa but that was not observed here.<br />
BRAZIL. Mato Grosso: Serra do Cachimbo, 763 km N of Cuiabá on Cuiabá-Santarém highway (Br-163),<br />
ca. 400 m, 09°35'S, 54°55'W, 22 Apr 1983, Brako 5640; Pará: Serra do Cachimbo, Aeroporta Cachimbo, 20<br />
km N of border with Mato Grosso on Cuiabá-Santarém highway (Br-163), 430-480m, 09°22'S, 54°54'W, 27<br />
Apr 1983, Brako 6208, near Base Aerea do Cachimbo, along Cuiabá-Santarém highway (Br-163), 430-480 m,<br />
ca. 09°22'S, 54°54'W, 28 Apr 1983, Brako 6323 (all INPA and NY).<br />
Mycomicrothelia rubrostoma Aptroot<br />
Aptroot either did not notice or ignored the fact that this species has well developed endospore (to 4 µm)<br />
giving the locules a bowling pin shape. Since all the other characters seem to agree with Mycomicrothelia, I<br />
am satisfied that it is correctly placed. There is a second species with this ascospore type in Brazil with<br />
shorter, elliptical to subspherical ascospores. It is left undescribed pending more ample material.<br />
85
MELANOMMATALES s. lat.<br />
PYRENULACEAE Rabenh.<br />
Since my treatment of Pyrenulaceae in eastern North America (Mem. New York Bot. Gard. 49: 74-107.<br />
1989) and the preliminary treatment in "Some Florida Lichens", Aptroot has published a treatment of the<br />
"peripheral" genera of Pyrenulaceae and Requienellaceae (Biblioth. Lichenol. 44: 1-178. 1991). Unfortunately<br />
this work is flawed by numerous careless errors and poor observation. An extensive critique of the<br />
morphological characters and cladistic results was published by Eriksson (Lichenologist 25: 307-311. 1993).<br />
(From the point of view of a cladistic-know-nothing, Aptroot's errors, especially incorrect generic and familial<br />
placements, would seem to make many of Eriksson's comments on the cladistic trees rather irrelevant.) In<br />
spite of these problems much of the skeleton is sound and should not be discarded. Further, as a result of reexamining<br />
some of the material studied by Aptroot, I have modified some of my ideas regarding the familial<br />
placement of some of these genera. I had believed that the taxa with short, coarse, essentially unbranched<br />
physes should be referred to the Requienellaceae. Restudy of hamathecial and ascal characters in Pyrenula<br />
have led me to think that these are differences of degree and not kind. Requienella has a unique ascus type<br />
with a "tholus" staining with various dyes (Boise, 1986). The taxa previously thought to belong to the<br />
Requienellaceae also have rod-shaped microconidia, shorter than the filiform, often curved microconidia of the<br />
Pyrenulaceae but, again, it seems to be a matter of degree. Requienellaceae sensu Aptroot seem to be nonlichen-forming.<br />
I surmise this may be an adaptation to xeric habitats similar to Polymeridium in the<br />
Trypetheliaceae. As a consequence I am restricting the Requienellaceae to Requienella, transferring the<br />
other genera placed in the family by Aptroot to the Pyrenulaceae along with the resurrected genera,<br />
Acrocordiella and Mycopyrenula. Another possibility, of course, would be to erect another family to<br />
accommodate Distopyrenis, Granulopyrenis, Lachrymospora and Pyrenographa and possibly Acrocordiella<br />
and Parapyrenis. The uniformly rod-shaped microconidia of the first four (not filiform as in Pyrenulaceae s.<br />
str.) make this a tempting solution (microconidia not known for the last two). Perhaps adding to the confusion,<br />
there are a wide variety of ascomatal types, pyrenuloid, parathelioid, pyrenastroid and possibly even<br />
mycoporoid, none of which seems to align with ascospore type. There are so few specimens known it seems<br />
best to more or less maintain the status quo pending more material and, perhaps, even molecular studies<br />
which would seem useful in dealing with what seems to be a highly reduced group.<br />
In 1989 when I abandoned ascomatal orientation and arrangement as generic characters, I did not<br />
mention the case where many ascomata with apical ostioles are included within a pseudostroma filled with<br />
pigment and/or oxalate(?) crystals. This type is very common in the Trypetheliaceae but very rare in the<br />
Pyrenulaceae. To repair the previous omission two such taxa are included below although neither is reported<br />
from Florida, P. concastroma and P. wrightii.<br />
KEY TO GENERA OF PYRENULACEAE AND REQUIENELLACEAE<br />
1. Ascomata perithecioid............................................................................................................................... 2<br />
2. Ascospores some shade of brown, mostly thick walled........................................................................ 3<br />
3. Ascospores 4-celled to muriform; physes long and slender<br />
(except in Acrocordiella and Mycopyrenula which have 4-celled<br />
ascospores and are non-lichenized) ................................................................................................ 4<br />
4. Lichenized; thallus usually conspicuous........................................................................................ 5<br />
86
5. Endospore absent or poorly developed..................................................................................... 6<br />
6. Ascomata with an outer pseudostromatic shell; ascospores<br />
dark brown, relatively thin walled (endospore poorly<br />
developed); lumina nearly cylindrical or cubical; mainly<br />
subtropical and tropical .............................................................................. Anthracothecium<br />
6. Ascomata lacking outer pseudostromatic shell; ascospores<br />
medium brown or yellow brown; thin walled (endospore not<br />
readily detectable); temperate North America and Europe<br />
............................................................................................ [Eopyrenula, see Dacampiaceae]<br />
5. Endospore well developed; various shades of brown;<br />
lumina rounded; mostly distoseptate............................................................................Pyrenula<br />
4. Not lichenized; no thallus evident.................................................................................................. 7<br />
7. Ascospores 4-celled; physes sparse, coarse; Europe .............................................................. 8<br />
8. Ascomata immersed in twigs of Ribes ............................................................. [Acrocordiella]<br />
8. Ascomata superficial on young bark of Betulaceae ........................................[Mycopyrenula]<br />
7. Ascospores 3-7 transversely septate to submuriform............................................................... 9<br />
9. Occurring in Europe and eastern North America; ascospores<br />
not distoseptate with a single eccentric euseptum; physes<br />
relatively long, abundant; ascomata large, immersed in soft<br />
bark of deciduous trees.................................................................... [Requienella, see below]<br />
9. Occurring in the West Indies; ascospores with 4 distosepta and<br />
a single terminal euseptum; known only from the type (not seen<br />
by me, placed only provisionally in Pyrenulaceae) .......................................... [Polypyrenula]<br />
3. Ascospores 2-celled or rarely submuriform (no 4-celled ascospores<br />
yet known in this group); physes coarse, relatively short, rather abruptly<br />
tapering; non-lichenized except for Clypeopyrenis; microconidia rod-shaped,<br />
to 15 µm, longer in Clypeopyrenis................................................................................................... 10<br />
10. Ascospores with well developed endospore, larger; not<br />
lichenized; physes coarse ........................................................................................................... 11<br />
11. Ascospores 2-celled.............................................................................................................. 12<br />
12. Ascospores elliptical, oblong or subspherical; septum median........................................ 13<br />
13. Ascospores with unlayered endospore without pigment granules ............................. 14<br />
14. Ascus tip thickened, with a distinct apical chamber (may be<br />
necessary to use stain + KOH) .......................................................... Distopyrenis<br />
14. Ascus wall uniform, thin, without obvious apical chamber;<br />
Indonesia, Australia?...........................................................................[Parapyrenis]<br />
13. Ascospores with an inner colorless layer surrounding the lumina;<br />
pigment granules in the endospore......................................................[Granulopyrenis]<br />
12. Ascospores teardrop-shaped, with septum strongly eccentric;<br />
87
Madagascar............................................................................................... [Lachrymospora]<br />
11. Ascospores submuriform ...................................................................................................... 15<br />
15. Ascoma highly melanized, elongated and somewhat<br />
graphid-like in aspect; "Pacific", from Galapagos to<br />
Australia ......................................................................................................[Pyrenographa]<br />
15. Ascoma with eccentric ostiole, not resembling a graphid .........................[Granulopyrenis]<br />
10. Ascospores 2-celled, with only the slightest hint of endospore<br />
at the median septum, ca. 7 × 2.5-3 µm; thallus well developed,<br />
lichenized; physes slender, Pyrenula-type; West Indies,<br />
Central America and northern South America ........................................................ [Clypeopyrenis]<br />
2. Ascospores colorless .......................................................................................................................... 16<br />
16. Ascospores ovate, 4-celled; ascomata not pigmented, simple or<br />
compound (astrothelioid or pyrenastroid); ostiole apical or lateral.............................. Lithothelium<br />
16. Ascospores ± cylindrical, 4-celled, often bent, ca. 15-21 × 3-4.5 µm;<br />
ascoma not compound, with apical ostiole covered with orange<br />
pigment, KOH+ purple; South Africa .......................................................................[Pyrenowilmsia]<br />
1. Ascomata mazedioid............................................................................................................................... 17<br />
17. Ascospores 4-celled, 11-13 × 6-8.5 µm, dark brown with lenticular cells;<br />
ascomatal wall melanized; thallus UV+ yellow (lichexanthone);<br />
south to Polk County .................................................Pyrgillus javanicus (Mont. & v.d. Bosch) Nyl.<br />
17. Ascospores submuriform, subspherical; ascomatal wall not<br />
melanized; one species in Costa Rica, second in New Guinea .............................. [Mazaediothecium]<br />
ACROCORDIELLA O. Eriksson<br />
When I re-examined material of Acrocordiella occulta, I was struck by the difference in the physes and in<br />
ascospore development from Requienella. Consequently I support the position that Eriksson has consistently<br />
and correctly maintained that Acrocordiella should have generic status. There is only a single species, A.<br />
occulta (Romell) O. Eriksson, known only from Europe.<br />
ANTHRACOTHECIUM Massal.<br />
1. Ascospores large, over 50 µm long........................................................................................................... 2<br />
2. Ascospores 8/ascus; ostiole apical ........................................................................................................ 3<br />
3. Ascospores muriform, 60-95 × 27-40 µm; peninsular<br />
Florida .........................................................................................A. prasinum (Eschw.) R. C. Harris<br />
3. Ascospores transversely septate to submuriform, 50-65 × 20-29 µm;<br />
Dade, Highlands and Seminole counties ....................................................... A. varians R. C. Harris<br />
2. Ascospores 2/ascus, 130-180(-225) × 38-50 µm; ostiole lateral;<br />
common, throughout the state .............................................................A. nanum (Zahlbr.) R. C. Harris<br />
88
1. Ascospores small, under 20 µm long ........................................................................................................ 4<br />
4. Ascospores flattened, lozenge-shaped, with four rows of two cells each .............................................. 5<br />
5. Hymenium inspersed; ascospores 11-13 × 6-8 µm; thallus UV-;<br />
north to Putnam County ...............................................................A. canellae-albae (Fée) Müll. Arg.<br />
5. Hymenium not inspersed; ascospores 14-18 × 7-9 µm; thallus UV+<br />
yellow or UV-; Jefferson County............................ A. staurosporum (Tuck. ex Willey) R. C. Harris<br />
4. Ascospores globose to subglobose, cruciately divided into four cells,<br />
2-13 µm across or 12.5-15 × 10-13 µm; thallus UV+ yellow (lichexanthone);<br />
Franklin County, otherwise peninsular Florida .................................................A. subglobosum Riddle<br />
NOTES<br />
Anthracothecium staurosporum (Tuck. ex Willey) R. C. Harris<br />
In Harris (1989) the thallus is reported as UV-. I have since seen specimens from Texas and French<br />
Guiana in addition to the Florida collection which contain lichexanthone. Since they do not differ otherwise<br />
from more northern UV- material, I consider it to be minor chemical variation.<br />
DISTOPYRENIS Aptroot<br />
I am quite uncomfortable with Aptroot's choice of a generic type, D. americana. It is anomalous in terms<br />
of distribution (temperate), substrate (Betula), ascospores (almost mischoblastic with a median swelling) and<br />
possibly ascomatal structure. It raises the specter that Distopyrenis should be restricted to its type. The other<br />
species would then require a new generic name or could perhaps be dumped into the closely related<br />
Granulopyrenis. Only the single, somewhat aberrant species, D. submuriformis, is in Florida.<br />
1. Ascospores 2-celled; ascomata various.................................................................................................... 2<br />
2. Ostiole apical or ostiole orientation not known ...................................................................................... 3<br />
3. Growing on a variety of substrates other than Betula ....................................................................... 4<br />
4. Ascomata small, less than 0.5 mm diam.; ascospores without<br />
strong median septum.................................................................................................................. 5<br />
5. Ascospores with broadly rounded ends ................................................................................... 6<br />
6. Ascospores with a narrow ring in middle of septum where<br />
lumina meet (two dark dots in optical section),12-14 × 8-9.5 µm;<br />
ascomata apparently compound, mycoporoid or pyrenastroid?;<br />
Puerto Rico.................................................................................. [D. composita R. C. Harris]<br />
6. Ascospores with a narrow dark ring at outer ends of median septum<br />
(two dark dots in optical section), 12-15 × 7.5-9.5 µm; ascomata<br />
simple; Haiti.............................................................................. [Distopyrenis sp. Buck 9033]<br />
5. Ascospores tapering at ends, 15-18 × 8-10 µm; ascomata simple;<br />
ostiole orientation not evident; Venezuela.................................[Distopyrenis sp. Funk 6254-A]<br />
4. Ascomata larger, 0.6-0.8 mm diam.; ascospores with a dark<br />
median plate (euseptum?), ca. 14-16 × 7.7-8.5 µm;<br />
89
on Beaucarnia recurvata, Mexico.................................................................[D. fuliginosa Aptroot]<br />
3. Growing on smooth bark of Betula; ascospores appearing almost<br />
mischoblastic, often swollen in the middle, ca. 18-19 × 7 µm;<br />
eastern North America .................................................................................... [D. americana Aptroot]<br />
2. Ostiole lateral ........................................................................................................................................ 7<br />
7. Ascospores small, 18 × 8 µm or less ............................................................................................... 8<br />
8. Ascospores 13-18 × 7-8 µm; Texas, Dominican Republic,<br />
French Guiana ....................................................................................... [D. pachyospora Aptroot]<br />
8. Ascospores narrower, 14-17 × 5-6.5 µm; Cuba ............................. [Distopyrenis sp. Buck 23459]<br />
7. Ascospores relatively large, 16-19 × 9-10.5 µm; on bark of<br />
white oak; Mississippi.............................................................................. [D. quercicola R. C. Harris]<br />
1. Ascospores submuriform, 3 × 1-2 irregularly septate, 13-17 × 7-9 µm;<br />
ascoma with extensive clypeus elongated in direction of wood fibers;<br />
ostiole apical; on eroded wood; Monroe County .......................................D. submuriformis R. C. Harris<br />
NOTES<br />
Distopyrenis americana Aptroot<br />
Previously recorded from Iowa, New York and North Carolina, the range is extended to Wisconsin.<br />
WISCONSIN. Bayfield County: Rainbow Lake Wilderness, Chequamegon National Forest, NW of Muck<br />
Lake and SW of Beaver Lake, on yellow birch, 12 Aug 1992, Wetmore 71414 (MIN).<br />
Distopyrenis composita R. C. Harris, sp. nov.<br />
Ascomatis compositis, rare simplicibus, positione ostioli ignota, ascosporis late ellipsoideis annulo parvo<br />
centrali, 11.5-13.5 × 8-9.5 µm.<br />
Type. PUERTO RICO. Distr. Aguadilla: Montañas Ayamon, Bosque Estatal de Guajataca, at N end of forest on<br />
Hwy. 446, 200-250 m, along trail #1 to observation tower, mesic forest over limestone, 25 May 1989, Harris<br />
23966 (NY, holotype).<br />
Thallus indicated by slight discoloration of bark and black lines when in contact with other thalli.<br />
Ascomata flattened, often somewhat lumpy, orbicular to irregular, usually compound, rarely simple (young<br />
stage?), 0.2-0.5 mm diam. (individual ascomata ca. 0.1-2 mm diam.); clypeus usually covering several<br />
hymenia; ostiole orientation and mode of ascomatal joining unknown. Asci cylindrical to obclavate, with<br />
uniseriately to irregularly arranged ascospores. Ascospores broadly ellipsoidal with rounded to somewhat<br />
angular lumina, with a odd little dark ring in septum where the lumina come together, 11.5-13.5 × 8-9.5 µm.<br />
Microconidia not found.<br />
I apologize but I am simply unable to figure out the structure of the ascomata as they are so small. It will<br />
require microtome sections or an extremely lucky hand section to determine this character. However, the<br />
obviously compound nature of most of the ascomata conjoined with the small central ring in the ascospores<br />
sufficiently distinguishes this species. It is known only from the type collection.<br />
Distopyrenis fuliginosa Aptroot<br />
Microconidia from holotype, rod-shaped, 8-10 µm. Buck 4791 and Harris 15854 from the Dominican<br />
Republic cited by Aptroot as D. fuliginosa are D. pachyspora. Buck 9033 from Haiti is an undescribed species.<br />
90
Distopyrenis pachyospora Aptroot<br />
Aptroot described the ostiole as apical but my drawings from the type show it as strongly eccentric as it is<br />
in the other material cited here. Buck 4791 and Harris 15854 cited by Aptroot as D. fuliginosa are D.<br />
pachyospora.<br />
New to North America. TEXAS. Bastrop County: Bastrop State Park, just E of Bastrop, on Quercus, 19<br />
Mar 1976, Egan EL-8193 (MIN, NY); Robertson County: Boy Scouts of America Camp Arrowmoon, 6 mi E of<br />
Hearne off FM 2549, on Quercus, 10 Apr 1976, Egan EL-8355 (MIN).<br />
Distopyrenis quercicola R. C. Harris, sp. nov.<br />
Ostiolis excentris et ascosporis ovalibus sat grandis, 16-19 × 9-10.5 µm.<br />
Type. MISSISSIPPI. Smith County: Bienville National Forest, Marathon Lake Recreation Area, oak-pine woods<br />
on W shore of lake, on white oak, 30 Sep 1992, Harris 28782 (NY, holotype).<br />
Thallus indicated by whitening of bark. Ascomata mostly immersed, ± oval to pear-shaped, ca. 0.3 × 0.4-<br />
0.5 mm. Ostiole eccentric with a well developed gelatinous periphysoid plug. Asci obclavate, with no or poorly<br />
developed ocular chamber, with irregularly arranged ascospores. Ascospores oval with large rounded to<br />
obovoid lumina, euseptum not evident, 16-19 × 9-10.5 µm. Microconidia rod-shaped, 8-9 × 1 µm.<br />
Distopyrenis quercicola is similar to D. pachyspora in ascoma morphology but differs in the larger<br />
ascospores with larger lumina. It is known only from the type collection.<br />
Distopyrenis submuriformis R. C. Harris, sp. nov.<br />
Species unica ascosporis submuriformibus, 3 × 1-2 septatis, 13-17 × 7-9 µm et ascomatis similibus<br />
Pyrenographae.<br />
Type. FLORIDA. Monroe County: Everglades National Park, road W of West Lake through mangrove swamp<br />
and hammock to ocean, on old wood, 14 May 1967, Harris 2848-D (NY, holotype).<br />
Thallus indicated by pinkish white discoloration of wood. Ascomata oval to long oval, elongated with<br />
grain of wood, strongly melanized, sessile or on a small pedestal of wood (due to erosion of wood away from<br />
ascomata?), 1.2 × 0.7-1.0 mm. Ostiole central and apical. Asci cylindrical with uniseriate ascospores.<br />
Ascospores ellipsoidal, submuriform, 3 × 1-2 septate, 13-17 × 7-9 µm. Microconidia not found.<br />
This is a troubling species. The submuriform ascospores and Pyrenographa-like ascomata suggest<br />
placement in Pyrenographa. However, the ascospores have better developed endospore and a "Rinodinalike"<br />
2-celled stage in development unlike Pyrenographa where the young ascospores are relatively thin walled<br />
and do not form a strongly thickened endospore. Therefore, the species is provisionally assigned to<br />
Distopyrenis. The collection was made in 1967 providing some insight into how long it takes for systematic<br />
knowledge to accumulate to a point where some specimens can even be provisionally assigned to a family<br />
and genus.<br />
Distopyrenis sp. Buck 9033<br />
This is a rather poor collection and I am unable to determine ostiole orientation. The ascospores have a<br />
dark ring at the margin of the septum unlike D. composita where it is central. Its formal description awaits<br />
more adequate material as do two other neotropical species noted in the key.<br />
GRANULOPYRENIS Aptroot<br />
91
My first inclination was to lump Granulopyrenis, Distopyrenis and Parapyrenis. Aptroot distinguished<br />
Granulopyrenis on the basis of pigment granules in the endospore. In the broader context of the<br />
Pyrenulaceae this would not be significant. However, the ascospores are also distinctive in possessing a ±<br />
colorless inner endospore layer immediately adjacent to the lumen. This is also seen in Pyrenula rubrostoma<br />
R. C. Harris. In some ways it is more reasonable to lump and have only one genus which in terms of<br />
ascomatal morphology, biology, ecology and distribution don't make much sense rather than two such genera.<br />
In the context of the part of the Pyrenulaceae with short, coarse physes, this distinctive ascospore type can<br />
serve to define a genus to maintain a little controversy and encourage further investigation of generic limits.<br />
Aptroot's concept is expanded to include a species with submuriform ascospores. It seems odd that no<br />
species with 4-celled ascospores are known (possibly hidden in unsuspected non-lichen-forming genera?).<br />
Aptroot reported microconidia from only G. macrocarpoides but they are present and uniform in two other taxa.<br />
1. Ascospores 2-celled.................................................................................................................................. 2<br />
2. Ascomata solitary; ostiole apical or lateral ............................................................................................ 3<br />
3. Ostiole eccentric or lateral................................................................................................................. 4<br />
4. Ascus without ocular chamber (fide Aptroot); ascospores<br />
13-15 × 8-9 µm; Puerto Rico, Virgin Islands, Lesser Antilles .......................[G. antillensis Aptroot]<br />
4. Ascus with rounded ocular chamber (fide Aptroot);<br />
ascospores 15-17 × 8-10 µm; eastern North America .................. [G. hymnothora (Ach.) Aptroot]<br />
3. Ostiole apical; ascospores 16-21 × 11-13 µm; South Africa ...............................................................<br />
............................................................................................... [G. macrocarpoides (Zahlbr.) Aptroot]<br />
2. Ascomata compound, joined by fused necks with a single<br />
common ostiole; ascospores 16-19 × 7-9 µm; Bermuda<br />
and Brazil.................................................................................. [G. nigrescens (Müll. Arg.) R. C. Harris]<br />
1. Ascospores submuriform, ca. 4 × 1-2 irregularly septate, broadly elliptical,<br />
15-19 × 10-12 µm; ostiole lateral; microconida rod-shaped, 9-10 × 1 µm;<br />
Dominican Republic .............................................................................. [Granulopyrenis sp. Harris 19568]<br />
NOTES<br />
Granulopyrenis antillensis Aptroot<br />
This is maintained as distinct from G. hymnothora but I have not verified the ascus character used by<br />
Aptroot to separate them. The ascospore sizes overlap but those of<br />
G. antillensis seem slightly smaller. Microconidia are rod-shaped, 10-12 × 1 µm and were found in Seaver &<br />
Chardon 187 (NY) from Puerto Rico. Britton et al. 102 (NY) from St. Thomas cited by Aptroot as Distopyrenis<br />
americana is G. antillensis.<br />
Granulopyrenis hymnothora (Ach.) Aptroot<br />
Microconidia from the BM isotype are rod-shaped, 7-9 × 1 µm.<br />
Granulopyrenis macrocarpoides (Zahlbr.) Aptroot<br />
Microconidia from the holotype are 8-10 × 1 µm.<br />
Granulopyrenis nigrescens (Müll. Arg.) R. C. Harris, comb. nov.<br />
92
Tomasellia nigrescens Müll. Arg., Hedwigia 32: 135. 1893. Type. BRAZIL. Pará: Igarapé, Spruce,<br />
Lich. Amaz. And. 364 (G, isotype).<br />
Parapyrenis elongata Aptroot, Biblioth. Lichenol. 44: 100. 1991. Type. BERMUDA. Hall's Island, on<br />
Eugenia axillaris, 27 Aug-21 Sep 1912, Brown & Britton 885 (NY, holotype).<br />
Aptroot synonymized Tomasellia nigrescens with Granulopyrenis hymnothora but from my notes and<br />
drawings of the type (I have not re-examined it in 1995), it does not differ from Parapyrenis elongata. Study of<br />
the type of P. elongata shows a ± colorless inner layer of endospore and granules in the endospore typical of<br />
Granulopyrenis.<br />
Granulopyrenis sp. Harris 19568<br />
Although this species expands the circumscription of Granulopyrenis to include submuriform ascospores<br />
(the young 2-celled ascospores are identical in morphology to the mature ascsopores of the other taxa), the<br />
Dominican Republic collection is simply not ample enough for a type which would endure much in the way of<br />
future study.<br />
LITHOTHELIUM Müll. Arg.<br />
KEY TO LITHOTHELIUM IN FLORIDA<br />
1. Ascospores 17-20 × 7.5-9 µm; Duval County...................................................... L. illotum (Nyl.) Aptroot<br />
1. Ascospores 10-12 × 4-4.5 µm; Collier County............................................. L. microsporum R. C. Harris<br />
NOTES<br />
Lithothelium almbornii R. C. Harris & Aptroot<br />
A small collection from the Dominican Republic seems most closely related to<br />
L. almbornii previously known only from Africa. The Dominican collection differs in slightly broader ascospores<br />
and in having the upper part of the hymenium inspersed. Until better material is available it can be assigned<br />
here with a question mark.<br />
DOMINICAN REPUBLIC. Prov. La Romana: Isla Catalina, alt. 0-10 m, 18 Dec 1987, Zanoni et al. 37309[A]<br />
(NY).<br />
Lithothelium bahamense Riddle = Strigula bahamense<br />
Lithothelium concameratum (Stirton) Makhija et al.<br />
I have seen no material but the illustration in Makhija et al. (1994) suggests Anisomeridium.<br />
Lithothelium fugax Müll. Arg.<br />
Aptroot synonymized this taxon with L. obtectum (Müll. Arg.) Aptroot with pyrenastroid ascomata.<br />
However, the ascomata are fused as in L. cubanum (± astrothelioid) and the ascospores are larger, 16-18 × 7-<br />
8 µm. I would prefer to recognize it as a distinct species. It is only known from the African type collection.<br />
Lithothelium illotum (Nyl.) Aptroot<br />
Sphaeria (Metasphaeria) cavernosa Ellis & Everhart, J. Mycol. 1: 91. 1885. Type. GEORGIA.<br />
Darien, on bark of Taxodium distichum, Ravenel 703 (NY, holotype), syn. nov.<br />
The occurrence of Lithothelium illotum in North Florida is no surprise as it has been found in the<br />
Carolinas, Georgia, Louisiana and Mississippi.<br />
FLORIDA. Duval County: Livezey's [Creek], on Liriodendron, Calkins, NAL 196 (NY ex PAC). Additional<br />
record. CUBA. Camagüey Prov.: In the Cuitos Mts., midway between Camagüey and La Gloria, 19-21 Feb<br />
1941, Johnson s.n. (NY).<br />
Lithothelium microsporum R. C. Harris, sp. nov.<br />
93
Ascomatis singularibus ostiolis excentricis, hymenio non insperso et ascosporis minimis, hyalinis, 4locularibus,<br />
10-12 × 4-4.5 µm.<br />
Type. FLORIDA. Collier County: Fakahatchee Strand State Preserve, Royal Palm Hammock, 5.7 mi from S<br />
entrance on W. J. Janes Memorial Scenic Drive, hardwood-Taxodium swamp with Roystonea, on prop roots<br />
of Roystonea, 6 Dec 1992, Buck 22741 (NY, holotype).<br />
Thallus greenish gray, matte, Trentepohlia present. Ascomata solitary, not compound, orbicular to<br />
obovoid, ca. ½-_ immersed, 0.4-0.5 × 0.3-0.4 mm. Ostiole eccentric. Hymenium not inspersed. Hymenial gel<br />
IKI+ dirty pinkish orange. Asci cylindrical with ascospores uniseriate to subbiseriate. Ascospores colorless,<br />
ovoid, 4-celled, 10-12 × 4-4.5 µm.<br />
Lithothelium microsporum has the smallest ascospores in the genus. Closest in ascospore size is L.<br />
obtectum that has compound, pyrenastroid ascomata. I do not exclude the possibility that L. microscopicum is<br />
a form of L. obtectum behaving oddly at the edge of its range. Description at the specific level focuses further<br />
attention on the question. It is known only from the type collection.<br />
Lithothelium obtectum (Müll. Arg.) Aptroot<br />
The range of this species is extended to Venezuela.<br />
VENEZUELA. Amazonas: Dpto. Río Negro, along Río Mawarinuma, just outside Cañon Grande, vicinity of<br />
Neblina base camp, ca. 140 m, 00°50'N, 66°10'W, low primary forest on white sand, Jan 1985, Buck 12556<br />
(NY, VEN).<br />
Pleurotrema polysemum Nyl. (Lithothelium polysemum (Nyl.) Aptroot)<br />
This species has been an embarassment to many of us. I placed Pleurotrema in the Trypetheliaceae.<br />
Aptroot, in agreement with Eriksson (1981), removed it from that family but placement in the Pyrenulaceae is<br />
no improvement. I had the good fortune to have Sabine Huhndorf as a collegue for several years and she was<br />
working on Saccardoella. Several taxa described in lichen-forming genera have proven to belong to<br />
Saccardoella, a unitunicate, non-lichenized genus. Being marginally involved in her study, it struck me one<br />
day that the ascus of Pleurotrema was very similar to that of Saccardoella. Both she and Margaret Barr<br />
agreed and Barr has accepted Pleurotrema and the Pleurotremaceae in the unitunicates related to the<br />
Amphisphaeriaceae (Mycotaxon 51: 211. 1994) and "pure" lichenologists no longer have to worry about it. I<br />
feel this is a prime example of the sort of broad collaboration that will be needed to elaborate a well founded<br />
system for the ascomycetes.<br />
MYCOPYRENULA Vainio<br />
In 1989 I synonymized this genus with Pyrenula. It has typical Pyrenula type microconidia and<br />
ascospores, however its niche as a saprophytic? colonizer of smooth bark, rather short physes and ascus with<br />
a thin, ± uniform wall seem to set it apart. Further, as the collective genus Pyrenula will probably be<br />
dismantled into more natural groups, it does no harm to anticipate this event. There is only a single species,<br />
M. coryli (Massal.) Vainio. It is not known from North America.<br />
PARAPYRENIS Aptroot<br />
My first intuitive thought was to simply synonymize Parapyrenis with Distopyrenis. However, the type<br />
species P. aurora (Zahlbr.) Aptroot, is distinctive in asci with a uniformly thin wall and very short or no stipe,<br />
ascospores with a very strongly pigmented median plate and euseptum?, hard, strongly melanized ascomatal<br />
shell and Asian distribution. Pending further studies, I consider Parapyrenis to be monotypic. Parapyrenis<br />
elongata Aptroot is a Granulopyrenis. I have not seen material of the other two original species. From the<br />
94
illustration and distribution, P. conica Aptroot, may possibly remain in Parapyrenis. Parapyrenis guayaci (Fée)<br />
Aptroot with massive (for this group) ascospores seems of uncertain position but possibly is referable to<br />
Distopyrenis. Aptroot (Nova Hedwigia 60: 325-379. 1995) has described three additional species. There are<br />
two isotypes at NY of P. maclurae (Saulescu & Sandu) Aptroot from Romania, unfortunately neither containing<br />
the Parapyrenis. From the distribution and substrate it is unlikely to belong to the Pyrenulaceae. I have not<br />
seen material of P. maritima Aptroot and P. tecomatis (Berkeley & Curtis) Aptroot. From the illustrations, the<br />
former may be a Granulopyrenis (distinct layer shown around lumina), the latter, said to have a spinulose wall,<br />
that would be anomalous in the Pyrenulaceae, is of uncertain position. Perhaps, the "spines" are endospore<br />
granules and it is also a Granulopyrenis.<br />
PYRENOGRAPHA Aptroot<br />
This genus is a problem. Aptroot described the ascospores as 3-septate. Sabine Huhndorf, thinking she<br />
possibly had this from the West Indies, examined the isotype at NY and found that the ascomata matched the<br />
original description but the ascospores were submuriform. Investigation of the holotype (CBG) and another<br />
isotype (B) gave identical results. Further, some time previously I had examined isotype material of<br />
Phaeopeltosphaeria irregularis Wehmeyer. It is the same as all of the available material of Pyrenographa<br />
xylographoides Aptroot. Did Aptroot see only immature ascospores or is there another taxon with 3-septate<br />
ascospores (perhaps an uncited duplicate in his personal herbarium)? I have not seen material of the type<br />
species of Phaeopeltosphaeria, P. caudata Berlese & Peglion from Italy, which from the description does not<br />
seem likely to be congeneric with Pyrenographa, but this problem must await further study. Based on the<br />
designated holotype and the two isotypes of Pyrenographa xylographoides, a new combination is required,<br />
Pyrenographa irregularis (Wehmeyer) R. C. Harris, comb. nov. Phaeopeltosphaeria irregularis Wehmeyer<br />
in Martin, Pacific Science 2: 74. 1948. Type. ECUADOR. Galapagos: South Seymour Island, on dead,<br />
decorticated wood of Bursera graveolens, 6 Sep 1945, Martin 6251 (NY, isotype). Even if Aptroot can come<br />
up with material which matches his description, it is my understanding that the designated holotype can only<br />
be superseded through conservation. In the unlikely event that this should happen, the status of the genus<br />
and the disposition of P. irregularis will have to be reassessed. In addition to the Australian and Ecuadoran<br />
material, NY holds specimens determined by Wehmeyer from the Marshall Islands and the Tuamotu<br />
Archipelago.<br />
Aptroot emphasized the elongate ascoma but I feel this is merely a consequence of the substrate, bare,<br />
eroded wood. Species of Anisomeridium, Arthopyrenia and Julella growing on old wood show this same<br />
character of the clypeus elongated in the direction of the wood fibers. Retention of this genus is supported by<br />
the rather poorly developed endospore. Young ascospores apparently do not pass through a "Rinodina-like" 2celled<br />
stage resembling the mature stage of other taxa of Distopyrenis. Distopyrenis submuriformis which<br />
otherwise has similar ascomata and ascospores does have a "Rinodina-like" stage. The microconidia present<br />
in the NY isotype of Pyrenographa xylographoides, rod-shaped, 12-15 × 1 µm, are somewhat longer than<br />
those in the rest of the Pyrenulaceae with coarse physes.<br />
PYRENULA Ach.<br />
1. Ascospores muriform ................................................................................................................................ 2<br />
2. Ascospores 8/ascus (occasionally fewer due to abortion), under 75 µm long ...................................... 3<br />
3. Thallus some shade of red, orange or yellow (occasionally only over ascomata),<br />
KOH+ purple...................................................................................................................................... 4<br />
4. Thallus orange to yellow................................................................................................................ 5<br />
5. Ascospores with 4 rows of 4 locelli each, 13-23 × 9-13 µm;<br />
Seminole County south ....................................................P. ochraceoflava (Nyl.) R. C. Harris<br />
95
5. Ascospores with 6-8 rows of 4-8 locelli each,<br />
23-35 × 11-17 µm; peninsular Florida ..........................P. ochraceoflavens (Nyl.) R. C. Harris<br />
4. Thallus red; hymenium inspersed; ascospores 30-42 × 15-20 µm;<br />
Dade and Monroe counties ................................................ P. cruentata (Müll. Arg.) R. C. Harris<br />
3. Thallus not red, orange or yellow, KOH- ........................................................................................... 6<br />
6. Ascomata fused in groups by their eccentric or lateral ostioles<br />
(pyrenastroid), or pseudostromatic shells fused completely<br />
(melanothecoid)............................................................................................................................. 7<br />
7. Pseudostromata pyrenastroid; pseudostromatal shells not<br />
completely fused into a flat Arthonia like plate ........................................................................ 8<br />
8. Ascospores over 45 × 18 µm; ascomata often only at an angle,<br />
sometimes solitary............................................................................................................... 9<br />
9. Ascospores 45-65 × 18-27 µm; throughout Florida..............................P. ravenelii Tuck.<br />
9. Ascospores 65-85 × 25-32 µm (60-72 × 25-30 µm in<br />
Florida specimen); Columbia County ......................................................Pyrenula sp. 47<br />
8. Ascospores 33-42 × 13-18 µm; ascomata usually<br />
distinctly recumbent; Dade, Monroe and Sarasota<br />
counties .............................................................................. P. astroidea (Fée) R. C. Harris<br />
7. Pseudostromata melanothecoid, looking much like an Arthonia<br />
except for the central ostiole; margins crenulate; ascomata recumbent,<br />
fused at the single ostiole; ascospores with 8 rows of locelli,<br />
33-40 × 12-14 µm; Key Largo, Monroe County .............................................Pyrenula sp. 208a<br />
6. Ascomata solitary, ostioles apical ............................................................................................... 10<br />
10. Thallus UV-............................................................................................................................ 11<br />
11. Postmature ascospores empty, collapsing...................................................................... 12<br />
12. Ascospores over 45 × 16 µm .................................................................................... 13<br />
13. Locelli smaller, spherical, in 8-10 rows of up to 10 each;<br />
ascospores 45-62 × 16-30 µm ............................................................................ 14<br />
14. Ascospores (45-) 50-65 × (18-) 20-26 µm, relatively<br />
narrow with pointed ends; Columbia and Volusia<br />
counties ....................................................... P. mucosa (Vainio) R. C. Harris<br />
14. Ascospores 50-70 × 24 -30 µm, broad with rounded end;<br />
known only from Mississippi............................................................................. .<br />
............................................................... [P. pyrenuloides (Mont.) R. C. Harris]<br />
13. Locelli large, often angular, in 8 rows of up to ca. 6 each;<br />
ascospores 45-60 × 16-22(-25) µm; very common,<br />
throughout the state ..............................................................P. leucostoma Ach.<br />
96
12. Ascospores 30-42(-53) × 11-15 µm; south to Sarasota<br />
County ...................................................................................... P. thelomorpha Tuck.<br />
11. Postmature ascospores filled with red or colorless oil substance................................... 15<br />
15. Oily substance red; ascospores 40-53 × 17-22 µm;<br />
Dade and Monroe counties ..................................P. macularis (Zahlbr.) R. C. Harris<br />
15. Oily substance colorless; ascospores 37-51 × 14-19 µm;<br />
Keys..........................................................................................P. oleosa R. C. Harris<br />
10. Thallus white, UV+ yellow (lichexanthone); ascospores 14-23 × 8-13 µm;<br />
Dade, Monroe and Pinellas counties........................................ P. confinis (Nyl.) R. C. Harris<br />
2. Ascospores (2-)4(-6)/ascus, 100-150(-190) × 30-41(-48) µm; ostioles lateral,<br />
sometimes several fused; south to Highlands County ....................... P. falsaria (Zahlbr.) R. C. Harris<br />
1. Ascospores transversely septate or rarely with a few cells longitudinally divided ................................... 16<br />
16. Ostiole lateral or lateral and several fused to form compound ascomata......................................... 17<br />
17. Ascospores with terminal lumina elongate and directly against exospore ................................. 18<br />
18. Ascomata solitary................................................................................................................. 19<br />
19. Ascospores less than 35 µm long ................................................................................... 20<br />
20. Ascospores 25-32 × 10-13 µm; Dade and Monroe<br />
counties ..............................................................P. cuyabensis (Malme) R. C. Harris<br />
20. Ascospores 18-22(-25) × 7-9 µm; Dade and Monroe<br />
counties ......................................................... P. subferruginea (Malme) R. C. Harris<br />
19. Ascospores 45-65 × 20-25 µm, often with 2-3 cells longitudinally<br />
sudivided; Dade County ............................................................P. erumpens R. C. Harris<br />
18. Several ascomata joined by fused ostioles; ascospores<br />
17-21 × 7-7.5 µm; Florida, without locality, 1878, Austin..........................................................<br />
......................................................................................... P. personata (Malme) R. C. Harris<br />
17. Ascospores with terminal lumina separated from exospore<br />
by a layer of endospore.............................................................................................................. 21<br />
21. Ostiole not red ringed ........................................................................................................... 22<br />
22. Several ascomata joined by fused ostioles .................................................................... 23<br />
23. Ascospores 21-25 × 8-10 µm; numerous crystals around ostiole;<br />
Collier, Dade and Monroe counties .....................P. cubana (Müll. Arg.) R. C. Harris<br />
23. Ascospores 16-21 × 6-8 µm; crystals absent;<br />
throughout Florida .......................................... P. septicollaris (Eschw.) R. C. Harris<br />
22. Ascomata solitary........................................................................................................... 24<br />
24. Hymenium not inspersed.......................................................................................... 25<br />
97
25. Ascospores 18-25 × 8-10 µm; hymenial gel IKI+ blue-green<br />
or IKI-; throughout Florida ..........................................P. microtheca R. C. Harris<br />
25. Ascospores 37-45 × 15-22 µm; hymenial gel IKI+ orangish;<br />
Dade and Monroe counties .........................P. martinicana (Vainio) R. C. Harris<br />
24. Hymenium inspersed, gel IKI+ orangish; ascospores 13-17 ×<br />
5-6 µm; Polk County...............................................................P. wheeleri R. C. Harris<br />
21. Ostiole surrounded by bright red ring; hymenium inspersed to top of asci;<br />
ascospores dark brown with darker bands between cells, 18-24 × 10-14 µm;<br />
Georgia, not yet found from Florida................................................. [P. wetmorei R. C. Harris]<br />
16. Ostiole apical, erect (pseudostromatic shells may be laterally fused<br />
to form extensive pseudostromata in P. anomala)............................................................................. 26<br />
26. Ascospores elliptical or fusiform, not tapering to a tail .................................................................. 27<br />
27. Ascospores over 21 µm long (if thallus ecorticate see P. microcarpa) ................................... 28<br />
28. Thallus orange or red at least near ascomata................................................................... 29<br />
29. Thallus orange, pseudocyphellate; ascospores 26-42 × 12-15 µm;<br />
Dade, Monroe and Sarasota counties.................................................P. cerina Eschw.<br />
29. Thallus red (in shade forms pigmented only near ascomata);<br />
hymenium heavily inspersed; ascospores 27-35 × 13-17 µm;<br />
very common, throughout the state.......................................P. cruenta (Mont.) Vainio<br />
28. Thallus greenish when fresh, whitish to brown in the herbarium ...................................... 30<br />
30. Postmature ascospores with red or colorless oily substance 31<br />
31. Oily substance red; ascospores 4- or 6-celled,<br />
28-34 × 12-15 µm; very common, throughout the<br />
state.................................................................P. concatervans (Nyl.) R. C. Harris<br />
31. Oily substance colorless; ascospores 4-celled,<br />
28-35(-40) × 10-15(-17) µm; throughout the<br />
state.........................................................................................P. quassiaecola Fée<br />
30. Postmature ascospores empty, collapsing................................................................... 32<br />
32. Ascospores with rounded ends; hymenium not inspersed;<br />
hymenial gel IKI+ orange....................................................................................... 33<br />
33. Ascospores reaching 45 × 18 µm..................................................................... 34<br />
34. Thallus UV-; ascospores 27-38 × 12-18 µm;<br />
gelatinous sheath lacking dark terminal caps;<br />
not common, south to Sarasota County.........................................................<br />
...........................................................................P. punctella (Nyl.) Trevisan<br />
34. Thallus UV+ yellow (lichexanthone) or UV-; ascospores<br />
36-45 × 12-17 µm; ascospore sheath gelatinous with<br />
dark terminal caps; Holmes County ...........................P. caryae R. C. Harris<br />
98
33. Ascospores 44-50(-56) × 17-24 µm; on Ficus;<br />
Collier County........................................................................Pyrenula sp. 29916<br />
32. Ascospores apiculate or at least pointed at one or both ends,<br />
rather dark brown when mature, 22-29 × 9-11 µm; hymenium<br />
inspersed only in a layer just below ostiole; hymenial gel IKI-;<br />
Collier County.......................................................................P. acutalis R. C. Harris<br />
27. Ascospores less than 21 µm long (to 25 µm in P. microcarpa)............................................... 35<br />
35. Ascospores with terminal lumina directly against exospore;<br />
hymenial gel IKI+ blue-green............................................................................................. 36<br />
36. Hymenium inspersed................................................................................................... 37<br />
37. Thallus UV-; ascomata 0.3-0.6 mm across; ascospores<br />
often citriform, 14-18 × 8-10 µm; south to Lake County............................................<br />
........................................................................................P. citriformis R. C. Harris<br />
37. Thallus UV+ yellow (lichexanthone); ascospores<br />
13-22 × 8-11 µm; northern Florida.............................................................................<br />
.................................................................P. pseudobufonia (Rehm) R. C. Harris<br />
36. Hymenium not inspersed............................................................................................. 38<br />
38. Thallus UV+ yellow (lichexanthone); pseudostromatal<br />
wall lacking crystals; ascospores 18-21 × 10-13 µm;<br />
Dade and Monroe counties ....................................................P. cocoes Müll. Arg.<br />
38. Thallus UV-; pseudostromatal wall containing<br />
colorless crystals; ascospores 18-20 × 11-12 µm;<br />
Dade County.................................................. P. nitidula (Bresadola) R. C. Harris<br />
35. Terminal lumina separated from exospore by an obvious layer<br />
of endospore .................................................................................................................... 39<br />
39. Ascomata fused laterally to form an extensive pseudostromata<br />
(melanothecoid).......................................................................................................... 40<br />
40. Ascospores more or less uniformly colored; thallus usually<br />
tan to yellow brown.............................................................................................. 41<br />
41. Ascospores with thin endospore, 17-21 × 7-9 µm......................................... 42<br />
42. Pseudostromata raised, not forming extensive<br />
network,brown, with crowded ascomata;<br />
common, throughout the state ......................... P. anomala (Ach.) Vainio<br />
42. Pseudostromata flush with thallus, forming<br />
an extensive network, shiny black, with<br />
scattered ascomata; Dade and Monroe<br />
counties .........................................................................Pyrenula sp. 415<br />
41. Ascospores with thicker endospore,<br />
99
18-21 × 8-11 µm; Dade County...........................................Pyrenula sp. 6121<br />
40. Ascospores with a distinct dark brown median plate or<br />
sometimes in old spores plates between all four lumina,<br />
18-20 × 7-8 µm; thallus brown to blackish brown; on Ilex;<br />
Liberty County ............................................................P. atrolaminata R. C. Harris<br />
39. Ascomata solitary....................................................................................................... 43<br />
43. Thallus corticate ................................................................................................... 44<br />
44. Ascomata large, 0.7-1.5 mm across, flattened conical ................................. 45<br />
45. Hymenium not inspersed; ascomata with a thin<br />
translucent thalline covering................................................................... 46<br />
46. Ascospores 15-19 × 6-8 µm; south to<br />
Lake County .........................................P. santensis (Nyl.) Müll. Arg.<br />
46. Ascospores 12.5-15.5 × 4.5-5 µm;<br />
Collier County .......................................................Pyrenula sp. 30313<br />
45. Hymenium heavily inspersed; ascospores<br />
15.5-21 × 5.5-8 µm; ascomata naked, shiny;<br />
very common, throughout the state............................................................<br />
................................................................ P. mamillana (Ach.) Trevisan<br />
44. Ascomata smaller, to 0.7 µm......................................................................... 47<br />
47. Endospore developed ............................................................................. 48<br />
48. Endospore uniform........................................................................... 49<br />
49. Hymenium inspersed; ascospores<br />
13-15 × 4.5-6 µm; Taylor to Polk County........................................<br />
.......................................................................P. laetior Müll. Arg.<br />
49. Hymenium not inspersed; ascospores<br />
13-16 × 4.5-6 µm; throughout the state..........................................<br />
.................................................................P. aspistea (Ach.) Ach.<br />
48. Endospore with a distinct, additional layer around<br />
lumina or older ascospores with darker bands<br />
between the lumina .......................................................................... 50<br />
50. Endospore with additional layer around lumina;<br />
ascospores 17-19 × 7.5-9 µm; ostiole usually<br />
red pigmented, KOH+ blue; Columbia, Dade<br />
and Seminole counties ...................................................................<br />
........................................................ P. rubrostoma R. C. Harris<br />
50. Endospore with darker bands between the<br />
lumina; ascospores 15-19 × 6-7.5 µm;<br />
ostiole not pigmented; Sarasota County ........................................<br />
........................................................ P. confoederata R. C. Harris<br />
100
47. Endospore very reduced; ascospores 14-17 ×<br />
5-7 µm; Dade County ....................................... P. tenuisepta R. C. Harris<br />
43. Thallus ecorticate, white; ascospores 19-25 × 8-12 µm;<br />
hymenial gel IKI-; throughout the state...........................P. microcarpa Müll. Arg.<br />
26. Ascospores fusiform-clavate, tapering to a long tail, in a single<br />
bundle in the ascus, 7-10-celled, 60-70 × 4.5-5 µm; asci markedly<br />
clavate; hymenium inspersed, IKI-; thallus UV-, not pseudocyphellate;<br />
on shrub at edge of salt marsh, Collier County ................................................Pyrenula sp. 30018<br />
Pyrenula atrolaminata R. C. Harris, sp. nov.<br />
NOTES<br />
Subsimilis P. anomalae sed ascosporis lamina media atra vel vetustis omnibus laminis atris, 18-20 × 7-8<br />
µm et thallo castaneo atranti.<br />
Type. FLORIDA. Liberty County: Whitehead Lake Campground, Apalachicola National Forest, cypress swamp<br />
along Ochlockonee River, on Ilex, 11 Apr 1967, Harris 1454 (NY, holotype; MSC, isotype).<br />
Thallus chestnut brown initially but soon blackening to blackish brown, corticate, not pseudocyphellate,<br />
UV-. Ascomata partly solitary, hemispherical, ca. 0.5 mm diam. but mostly laterally aggregated into slightly<br />
raised, blackish pseudostromata containing several to ca. 30 ascomata, initially appearing pruinose due to<br />
disrupted bark cells; ostiole dark; crystals lacking. Hymenium not inspersed; hymenial gel IKI-. Ascospores<br />
subbiseriate, 4-celled, 18-20 × 7-8 µm, with a dark brown median plate and occasionally in older ascospores<br />
all three septa with brown plates.<br />
The epithet is given for the ascospores with dark layers between lumina but could also be applied to the<br />
dark thallus. I am unaware of any other species in the genus with this type of ascospore. Externally one<br />
would think this to be a dark thallus form of Pyrenula anomala. It is known from only a single site in the<br />
Apalachicola region, notorious for its high rate of endemism.<br />
Additional specimen. As type, Harris 1438 (NY).<br />
Pyrenula caryae R. C. Harris, sp. nov.<br />
Similissima P. punctellae sed thallo ceraceo saepe lichexanthone continente, ascosporis operculis<br />
brunneis et majoribus, (30-)36-45(-50) × 12-17(-21) µm et saepe ad Caryam crescente.<br />
Type. MISSOURI. Iron County: St. Francis Mountains, Clark National Forest, along Co. Rd. N just N of<br />
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<br />
31124 (holotype NY; isotypes H, CBG, MOR).<br />
Thallus usually yellowish or yellowish tan but occasionally green, corticate, somewhat shiny, usually with<br />
a waxy look, pseudocyphellate, UV+ yellow (lichexanthone) or less often UV-. Ascomata flask-shaped, 0.5-0.8<br />
mm diam., immersed, often entirely covered by thallus with only the pale ostiole visible, in some (with age?)<br />
the black ascomatal apex exposed; numerous colorless crystals in upper ascomatal wall. Hymenium not<br />
101
inspersed; hymenial gel IKI-. Ascospores subbiseriate to uniseriate, 4-celled, (30-)36-45(-50) × 12-17(-21)<br />
µm, with a darker pigmented median band and with a brownish apical cap at both ends (outside the exospore)<br />
which in fresh material is often reflexed looking like a little sombrero, gelatinous sheath otherwise not readily<br />
evident.<br />
Harris (1989) cited some of the collections now recognized as P. caryae under P. punctella (Nyl.)<br />
Trevisan where it was noted "The taxonomy of P. punctella requires further work...". Pyrenula caryae was<br />
initially segregated when the presence of lichexanthone was noticed. However, closer study revealed several<br />
correlated characters, the "waxy" thallus, larger ascospores with the very odd apical caps and a strong<br />
tendency to grow on Carya. Eleven of the fifteen known collections are from Carya and the species has been<br />
named for this preference. The substrate of the other four is neither noted on the label nor readily<br />
determinable. The ascospores of P. punctella are 27-38 × 12-18 µm and lack the apical caps but may have<br />
an apical dark spot in the exospore or are weakly apiculate with the apiculus often darkening. Also older<br />
ascospores tend to have a paler zone around the locules not seen in P. caryae. Pyrenula punctella is still<br />
probably not homogeneous. The Coastal Plain collections, mostly on Fraxinus or Magnolia, have a reddish<br />
almost endophloeodal thallus, interior collections, mostly on Fagus, have a whitish or grayish dull epiphloeodal<br />
thallus but I am not as yet prepared to separate them at the species level in a group where the intimate<br />
relation between lichen and substrate give rise to great differences in aspect. Three collections of P. caryae<br />
seem to lack lichexanthone entirely. In fact one of the three collections made at the type locality lacks<br />
lichexanthone. Overall it seems present only in low quantities as the fluorescence is quite weak. Presence or<br />
absence of lichexanthone is sometimes recognized at the specific rank but the unique ascospores allow one<br />
to dismiss such a treatment here. The known distribution of P. caryae may also be distinctive, running in an<br />
arc around the Southern Appalachians from North Carolina to Missouri mainly on the Piedmont although like<br />
P. caryae several also extend into northern Florida. The elevations noted are between 35 m and 900 m.<br />
Gomphillus americanus also fits this pattern and also possibly Cladonia caroliniana Tuck. s. lat. (if usnic acid<br />
only strain is included) and Cresponea premnea (Ach.) Egea & Torrente var. saxicola (Leighton) Egea &<br />
Torrente.<br />
Additional specimens. ALABAMA. Cleburne County: 2 mi NE of Heflin, 23 Oct 1964, Johnson<br />
(NY);Tuscaloosa County: near Rickey, 24 Oct 1964, Johnson (NY). ARKANSAS. Washington/Crawford County:<br />
Devil's Den State Park, 9 Jun 1961, Johnson (NY). FLORIDA. Holmes County: along St. Rd. 2 just E of<br />
Choctawhatchee River, 10 Dec 1993, Buck 24686 (NY). GEORGIA. Towns/Rabun County: Chattahooche<br />
National Forest, along Appalachian Trail between Dick's Creek Gap and Little Bald Knob, 825-900 m, 21 Sep<br />
1992, Harris 28115 (NY). MISSISSIPPI. Itawamba County: Donivan Slough, Mile 283.3 along Natchez Trace, c.<br />
35 m, 28 Sep 1992, Harris 28633 (NY); Lee County: near Tupelo, 31 Dec 1952, Johnson 1041A, 1046B (NY);<br />
Pearl River County: 1 mi S of Carriere, 14 Jun 1939, Johnson 1118 (NY). MISSOURI. Carter County: Bluff on E<br />
side of Current River across from Big Spring S of Van Buren, 23 Sep 1990, Harris 25633 (NY); Iron County: as<br />
type, Harris 31190, 31204 (NY). NORTH CAROLINA. Graham County: Horse Cove Campground, Nantahala<br />
National Forest, 3 Oct 1987, Buck 15387 (NY); Haywood County: Great Somky Mtns. Natl. Park, Big Creek,<br />
45 mi ESE of Knoxville, 2000-2300 ft, 25 Aug 1977, Wetmore 29850 (MIN); Macon County: McDowell<br />
Mountain, Nantahala National Forest, 640-945 m, oak-hickory woods, 12 Aug 1994, Harris 33192 (NY).<br />
SOUTH CAROLINA. Chester County: Chester, [Green?] (NY).<br />
Pyrenula concastroma R. C. Harris, sp. nov.<br />
Characteribus internis ut P. concatervanti sed ascomatis inclusis in pseudostromatis trypethelioidibus.<br />
Type. PUERTO RICO. Salinas de Guánica, 27 Dec 1930, Britton & Britton 9602 (NY, holotype).<br />
Thallus grayish brown, corticate, not pseudocyphellate, UV-. Pseudostromata sessile, constricted at<br />
base, mostly oval, ca. 2 × 1 mm, occasionally fused into larger groups, brown, filled with large oxalate(?)<br />
crystals and a whitish powder bleeding yellow in KOH. Ascomata many/pseudostroma with apical ostioles.<br />
Hymenium not inspersed. Hymenial gel IKI+ orangish. Asci with 8 biseriate ascospores. Ascospores 4-<br />
102
celled, becoming filled with reddish oily substance as contents disintegrate, 31-40 × 15-16µm.<br />
The epithet is derived from the bastardization of the epithet of the sister species "concatervans" and<br />
"stroma" as the ascospores, etc., are identical to those of P. concatervans and the ascomata are included in<br />
pseudostromata. This assumes that ascospores types are conservative and define clades although there is<br />
no independent proof for this. In previous generations, P. concastroma would have been placed in<br />
Melanotheca, a genus I have abandoned as polyphyletic, containing taxa derived independently in several<br />
ascospore defined clades. Pyrenula concastroma is virtually identical to P. wrightii which differs in having a<br />
colorless oily substance in postmature ascospores (as P. quassiaecola differs from P. concatervans). This<br />
raises the question, does the color of the oily substance mean anything? Have two taxa, differing mainly in the<br />
color of the contents of postmature ascospores, evolved identical, highly unusual for the Pyrenulaceae,<br />
pseudostromata? This would seem to be a question which the friendly neighborhood molecular biologist is<br />
best equipped to answer. Pyrenula concastroma is known only from the type.<br />
Pyrenula concatervans (Nyl.) R. C. Harris<br />
Pyrenula crystalligera H. Magn., Ark. Bot. n.s. 3(3-4): 239. 1956. Type. [HAWAIIAN ISLANDS.] Iles<br />
Sandwich. Voyage de M. J. Remy, 1851-1855. Herb. Mus. Paris, misit cel. Nylander, 1860 (FH-<br />
Tuck 4107, holotype), syn. nov.<br />
Pyrenula confinis (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria confinis Nyl., Ann. Sci. Nat. Bot. sér. 4, 3: 174. 1855. Anthracothecium confine (Nyl.)<br />
Müll. Arg., Linnaea 63: 45. 1880. Type. CHILE. Ad corticem eadem ac Glyphis cicatricosa (H-Nyl<br />
1250, holotype).<br />
Verrucaria albescens Nyl., Expos. syn. pyrenoc. 50. 1858. Anthracothecium albescens (Nyl.) Müll.<br />
Arg., Linnaea 64: 45. 1880. Type. GUYANA BRITANNICA. Ad cort. Spondias lutea L. (H-Nyl 1285,<br />
holotype).<br />
Anthracothecium corticatum Müll. Arg., Bull. Soc. Roy. Bot. Belgique 30: 96. 1891. Pyrenula<br />
corticata (Müll. Arg.) R. C. Harris, Mem. New York Bot. Gard. 49: 89. 1989. Type. COSTA RICA.<br />
San José, 5300 ft., 1890, Pittier (MSC, holotype fragment ex G).<br />
Harris (1989) declined to take up the epithet confinis as the holotype had not been located. Singh (1985)<br />
incorrectly thought a specimen from French Guiana to be the type. In 1994 I located the holotype in Helsinki.<br />
Nylander had apparently intended to name the species Verrucaria concinna but never published this name<br />
(possibly due to the existence of Verrucaria concinna Borrer). Nylander had not changed the name on the<br />
specimen and it was quite naturally filed under this name in his herbarium. I have now studied the specimen<br />
and it agrees with the taxon previously treated by me as Pyrenula corticata.<br />
Pyrenula cruenta (Mont.) Vainio<br />
Verrucaria circumrubens Nyl., Flora 50: 196. 1867, nom. nud., Syn. lich. Nov. Caled. 90. 1868.<br />
Pyrenula circumrubens (Nyl.) B. de Lesd., Bull. Soc. Bot. France 62: 463. 1910. Type. [NEW<br />
CALEDONIA. Loyalty Islands:] Lifu, 1864 Thiebaut (H-Nyl 1418, holotype), syn. nov.<br />
Pyrenula rubromaculata Vainio, Bot. Mag. Tokyo 35: 77. 1921. Lectotype (selected here). JAPAN.<br />
Prov. Kii: on Taonabo japonica, 30 Dec 1911, Yasuda 259 (Tur-Vainio 31242). Syntype. as<br />
lectotype, Yasuda 262 (TUR-Vainio 31243), syn. nov.<br />
Pyrenula laetior Müll. Arg.<br />
Bot. Jahrb. Syst. 6: 413. 1885. Type. CUBA. Wright, Verr. Cub. 51 (FH-Tuck 4108, isotype).<br />
New to North America. FLORIDA. Citrus County: Mason Creek, ca. 2 mi SW of Homossasa, coastal<br />
hammock, elev. 2 m, on trunk of Ilex cassine, 10 May 1994, Griffin 2944 (FLAS, NY); Polk County: Nalcrest,<br />
on fallen oak branch, 11 Mar 1989, Wheeler (NY); Taylor County: Aucilla Wildlife Management Area, Aucilla<br />
Sinks, S of Goose Pasture Road along Florida Trail, ca. 4 mi N of U.S. Hwy. 98, on young Carya, 12 Dec<br />
1993, Harris 32285 (NY).<br />
The ascospores and asci resemble those of Pyrenula aspistea (Ach.) Ach. which differs in having the<br />
hymenium not inspersed. Pyrenula laetior is also close to Pyrenula velata Müll. Arg. from Brazil which differs<br />
103
in IKI+ blue hymenium and larger ascospores (_ = 16.4 × 6.1 µm). The Cuban species Pyrenula gregantula<br />
Müll. Arg. may be synonymous but is tentatively separated by slightly broader ascospores (_ = 6.0 µm) and<br />
distinctly papillate ascomata. Pyrenula laetior was included in Some Florida Lichens as sp. 11-3-89.<br />
Pyrenula laevigata (Pers.) Arnold<br />
A new character for this species has been brought to my attention by J. Etayo. Addition of KOH to a<br />
section of the hymenium gives a fleeting sky-blue reaction. As far as I know this is unique.<br />
Pyrenula macularis (Zahlbr.) R. C. Harris<br />
Bottaria albidopallens Vainio, Ann. Acad. Sci. Fenn., ser. A, 15(6): 328. 1921. Anthracothecium<br />
albidopallens (Vainio) Zahlbr., Cat. lich. univ. 8: 121. 1931. Type. PHILIPPINE ISLANDS. Luzon:<br />
Manila, Dec 1912, Merrill, Bur. Sci. 8495 (F, isotype), syn. nov.<br />
Although Vainio's name is older I refrain from taking it up as there are other possible synonyms older yet<br />
whose types have not been examined. There is no point in changing the name several times. Also the type of<br />
Verrucaria analepta var. americana Ach., Anthracothecium americanum (Ach.) Müll. Arg. belongs here. The<br />
epithet cannot be taken up in Pyrenula due to the existence of Pyrenula americana (Massal.) Trevisan [=<br />
Ditremis americana (Massal.) R. C. Harris]. This is just as well as the species is by no means strictly<br />
American. Details of the typification will be dealt with elsewhere.<br />
Pyrenula mamillana (Ach.) Trevisan<br />
Pyrenula marginata Hook. in Kunth, Syn. pl. 20. 1822, syn. nov.<br />
As I have long suspected, but not been able to prove due to the unavailability of the type, the oldest name<br />
for the species treated in Harris (1989) as P. marginata is indeed P. mamillana. The typification, etc., will be<br />
dealt with elsewhere.<br />
Pyrenula mucosa (Vainio) R. C. Harris, comb. nov.<br />
Bottaria (Anthracothecium) mucosa Vainio, Hedwigia 38: (257). 1899. Type. GUADELOUPE.<br />
Mouteran, 250 m, sur un vieux pied d'Inga laurifolia, Duss 501 (TUR-Vainio 30963, holotype)<br />
[mixed with Pyrenula nitidula (Bresadola) R. C. Harris].<br />
This seems to be a member of a complex of taxa which differ mainly in ascospore size. The ascospores<br />
of this complex seem to shrivel with age, not developing either reddish or colorless oily material as in some<br />
otherwise similar species. Pyrenula pyrenuloides has larger, mostly blunt ascospores. Also similar is<br />
Pyrenula dissimulans (Müll. Arg.) R. C. Harris, comb. nov., Pleurothelium dissimulans Müll. Arg., Bot.<br />
Jahrb. Syst. 6: 387. 1885. Type. CUBA. Wright, Verr. Cub. 213 (G, holotype; NY, isotype). Synonym: Bottaria<br />
libricola f. nana Vainio, Mycologia 21: 39. 1929. PUERTO RICO. on open, dry hilltop near Yauco, 30 Dec 1915,<br />
Fink 1669 (MICH, NY, isotypes). Pyrenula dissimulans was referred to Pleurothelium as a few of the ostioles<br />
are slighty eccentric. Basically it seems to have upright ostioles. The ascospores are slightly smaller and<br />
broader in proportion (35-50 × 17-22 µm) than those of P. mucosa. Also the locelli are mostly larger and fewer<br />
in number. Pyrenula dissimulans has not yet been found in North America. However, to complicate matters<br />
even further there are two collections from Florida (Dade and Union counties) with rather small ascospores but<br />
not the spore type of P. dissimulans. I tentatively refer them to P. mucosa. There is also a "wild card" name<br />
whose type I have not seen, Pyrenula libricola Fée. It is readily apparent that further work is needed and that<br />
the situation is unstable.<br />
Pyrenula nitidula (Bresadola) R. C. Harris ined.<br />
The combination will be made elsewhere. The type is from Puerto Rico. It is also known from<br />
Guadeloupe (type of Pyrenula subsimplex Vainio) and French Guiana.<br />
New to North America. FLORIDA. Dade County: Deering Hammock, Cutler, on live oak, 15 Dec 1919, N.<br />
L. & E. G. Britton 724 (NY).<br />
Pyrenula occidentalis (R. C. Harris) R. C. Harris<br />
Pyrenula harrisii Hafellner & Kalb, Herzogia 9: 85. 1992. Type. MADEIRA. An der Strasse zwischen<br />
Ponta do Pargo und Porto Moniz, bei Serrado. An Alleebäumen (Acer pseudoplatanus). 530 m.<br />
104
5.VIII.1990, Kalb 23189, not seen, syn. nov.<br />
Although this has nothing to do with Florida, it gives me some pleasure to dispose of this name as I do<br />
not feel any need of advertising. Pyrenula occidentalis, not occurring in eastern North America, was not<br />
included in Harris (1989) which seems to have been Hafellner and Kalb's source of information. The species<br />
is oceanic occurring in the Pacific Nortwest, Great Britain, Scandinavia and South Africa. Its occurrence in<br />
Madeira is not the least unexpected.<br />
Pyrenula punctella (Nyl.) Trevisan<br />
Pyrenula ferax Müll. Arg., Bot. Jahrb. Syst. 6: 414. 1885. Type. CUBA. Wright, Verr. Cub. 81, 82<br />
(FH-Tuck 4108, isosyntypes), syn. nov.<br />
Pyrenula pyrenuloides (Mont.) R. C. Harris<br />
Harris (1989) tentatively reported this taxon from several southern states noting that the ascospores were<br />
smaller than those of the type and other specimens from French Guiana. Having re-examined the type of<br />
Bottaria mucosa Vainio, I now prefer to refer the smaller spored collections to this species. Only one<br />
collection from Mississippi seems to have the very broad, stumpy ascospores of P. pyrenuloides.<br />
Pyrenula wrightii (Müll. Arg.) R. C. Harris, comb. nov.<br />
Melanotheca wrightii Müll. Arg., Bot. Jahrb. Syst. 6: 396. 1885. Type. CUBA. Wright, Verr. Cub.<br />
231(NY, isosyntype), Verr. Lich. Cub. II. 679 (H, isosyntype).<br />
This species has ascospores and other internal features identical to those of P. quassiaecola Fée<br />
differing in having the ascomata in pseudostromata filled with oxalate(?) crystals and whitish powder bleeding<br />
yellow in KOH. It differs from P. concastroma only in having colorless rather than reddish "oil" in the<br />
postmature ascospores. It is known only from the type. See also discussion of P. concastroma.<br />
Pyrenula sp. 47<br />
Discussed by Harris (1989: 102). The ascomata of the Florida collection show considerable fusion and<br />
are superficially indistinguishable from P. ravenelii. Previously known from Louisiana and Misssissippi, its<br />
occurrence in northernmost Florida is no surprise.<br />
Pyrenula sp. 208a<br />
Although extremely distinctive this species is not described as the available material is scant. It is<br />
included as part of my ongoing campaign to convince colleagues that ascomatal type in the Pyrenulaceae<br />
does not seem to have any meaning above the species level. This species combines features of both the<br />
melanothecoid and pyrenastroid types. The ascomata are recumbent, fused at the single ostiole but the<br />
pseudostromatic shells are fused into a single melanized mass as in the species formerly referred to<br />
Melanotheca Fée. I suppose in Zahlbruckner's system the former would have outweighed the latter and the<br />
species would have been placed in Parmentaria Fée.<br />
Pyrenula sp. 6121<br />
Superficially this species is similar to P. anomala but seemingly the ascomata are larger and although the<br />
pseudostromata are similar sized, there are fewer ascomata making them up. The ascospore difference is<br />
fairly slight and I have not yet done the statistics but my impression is that the greater ascospore width of sp.<br />
6121 will be the most reliable character for separation. I have not given the taxon a name as it is fairly<br />
widespread and surely has one elsewhere. Melanotheca arthonioides var. machaerii Malme from Brazil, if the<br />
type matches, seems to be this and could be elevated in rank. In Florida it is known from only two sites in<br />
Everglades National Park collected by Johnson in 1965.<br />
PYRGILLUS Nyl.<br />
The genus has only a single species in Florida. See the generic key for a diagnosis.<br />
105
REQUIENELLACEAE Boise<br />
The Requienellaceae are recognized here in the restricted, original sense as discussed above under<br />
Pyrenulaceae. While comfortable with the family, I have never been satisfied with the species level taxonomy.<br />
However, as Boise had decided, not readily changeable, opinions, I kept my mouth shut until now. I have<br />
always felt that R. subcollapsa and Acrocordiella occulta (see Pyrenulaceae) merited recognition.<br />
REQUIENELLA Fabre<br />
1. Young ascospores ± thin walled................................................................................................................ 2<br />
2. Ascospores with cells densely filled with small oil droplets, 6-celled,<br />
ca. 29-33 × 10-12 µm; ostiolar area inspersed with large oil drops;<br />
Europe.....................................................................................[Requienella seminuda (Persoon) Boise]<br />
2. Ascospores with 1-few oil droplets/masses, 8-celled, sometimes with<br />
one cell longitudinally divided, ca. 36-48 × 11-13 µm; Europe ................................................................<br />
................................................................................................[Requienella lichenopsis (Massal.) Boise]<br />
1. Young ascospores with a well developed "cap" of endospore in the<br />
terminal cells; mature ascospores 4-celled, occasionally with 1-2 cells<br />
longitudinally subdivided, ca. 25-29 × 8-10 µm; on soft barked deciduous<br />
trees, eastern North America ............................. [Requienella subcollapsa (Ellis & Everhart) R. C. Harris]<br />
NOTES<br />
Requienella subcollapsa (Ellis & Everhart) R. C. Harris, comb. nov.<br />
Lophiostoma subcollapsa Ellis & Everhart, J. Mycol. 2: 100. 1886. Type. NEW JERSEY. Newfield,<br />
on outer bark of living Nyssa, Jun 1886, Ellis (NY, holotype).<br />
I am preliminarily assuming that there is only a single species in North Americ but I have not yet reexamined<br />
all the specimens cited by Boise as R. seminuda so that this assumption may prove incorrect.<br />
TRYPETHELIACEAE Zenker<br />
Harris (Acta Amazonica 14(1/2, Supl.): 55-80. 1984[1986]) included the genus Pleurotrema Müll. Arg. in<br />
the Trypetheliaceae. It has been since removed to its own family, Pleurotremataceae, in the non-lichenized<br />
unitunicate ascomycetes. (See Pyrenulaceae for more detailed comments).<br />
Aptroot (Biblioth. Lichenol. 44, 1991) recognized two additional genera of Trypetheliaceae, Megalotremis<br />
Aptroot and Ornatopyrenis Aptroot. Unfortunaely Aptroot's descriptions of ascus and hamathecium are not<br />
adequate and he did not trouble with conidia. I have not seen the types of either genera.<br />
Aptroot's hesitation in describing Megalotremis was well founded. Megalotremis, based on M. verrucosa<br />
(Makhija & Patwardhan) Aptroot, is certainly a synonym of Anisomeridium (Aptroot's drawings show both<br />
Anisomeridium-type ascospores and ascus tip.) Two-celled ascospores are unknown in the Trypetheliaceae.<br />
Anisomeridium, with a handful of exceptions, has 2-celled ascospores and species having asci with only 2 or 4<br />
ascospores are not uncommon. The thallus of M. verrucosa is corticate but corticate thalli are also seen in<br />
Anisomeridium (A. glaucescens and A. clandestinum). The second species, M. biocellata Aptroot, is almost<br />
certainly a species of Arthopyrenia (probably the same as Tomasellia dispora Müll. Arg., see Arthopyrenia).<br />
Most species of Arthopyrenia show some constriction of the ascospore wall ± grading into the "biocellate" type<br />
("porospore" in Vainio's terminology). (There are probably ten or so species with the porospore ascospore<br />
type.) Aptroot (Nova Hedwigia 60: 338. 1995) has recognized a third species, M. megalospora (Vainio)<br />
106
Aptroot, for which I have seen the type and a couple additional specimens. It belongs in Anisomeridium. The<br />
asci and physes are clearly the same as those seen in Monoblastiaceae. The ascospores of M. verrucosa<br />
and M. megalospora contain colorless crystals. (Eriksson also sees them as crystals but Aptroot interprets<br />
them as folds in the ascospore wall.) This character is seen in scattered species with large ascospores in the<br />
Monoblastiaceae, Pyrenulaceae and Trypetheliaceae and seems to have no lower level taxonomic<br />
significance. Possibly it is some storage product crystallizing which might provide support for ordinal<br />
relationship if biochemical analysis were to show it was unique or unusual.<br />
The disposition of Ornatopyrenis is more problematical. Ornamented ascospores are otherwise<br />
unknown in the entire family Trypetheliaceae but the pale brown, ornamented ascospores do suggest<br />
Mycomicrothelia (Arthopyreniaceae). Hawksworth's (1985) photo of the type, Microthelia queenslandica Müll.<br />
Arg., shows dark septa which also seem typical of Mycomicrothelia. Hawksworth's photo and Aptroot's<br />
drawings both show distoseptation which does not fit well with Mycomicrothelia as defined by Hawksworth.<br />
However, Aptroot described Mycomicrothelia rubrostoma with somewhat distoseptate ascospores although his<br />
examination was apparently so casual that he did not realize this. The ascospores of M. rubrostoma are<br />
similar to those in the porospore Arthopyreniae mentioned above, only brown. Thus, I am not uncomfortable<br />
with placement in the Arthopyreniaceae. Also the ascospores of Ornatopyrenis may become submuriform. I<br />
have already (1989) introduced a muriform-spored species into Mycomicrothelia, M. decipiens (Müll. Arg.) R.<br />
C. Harris. Until such time as I can study material of Ornatopyrenis queenslandica, I suggest Ornatopyrenis be<br />
placed tentatively as a synonym of Mycomicrothelia.<br />
KEY TO GENERA<br />
1. Thallus not corticate, whitish ..................................................................................................................... 2<br />
2. Ascospores with well developed endospore developed, lumina angular,<br />
4-celled, 20-27 × 6-8 µm; hymenium inspersed, also often yellow<br />
pigmented, KOH+ red; throughout the state ........................... Pseudopyrenula subnudata Müll. Arg.<br />
2. Ascospores with endospore not developed, lumina cylindrical,<br />
with 4-8 transverse septa or muriform.............................................................................. Polymeridium<br />
1. Thallus corticate, green, tan, brown or orange.......................................................................................... 3<br />
3. Ascomata solitary or in pseudostromata, not fused and sharing a common ostiole<br />
or ostiolar plate...................................................................................................................................... 4<br />
4. Stromata solitary or in pseudostromata which are not brown, and slightly<br />
shiny , if containing KOH+ purple pigment, then pseudostroma pruinose and<br />
ascospores 9-13 transversely septate; pseudostromal wall not composed<br />
of brown, convoluted hyphae............................................................................................................. 5<br />
5. Ascospores muriform .................................................................................................................... 6<br />
6. Ostiole apical..................................................................................................................Laurera<br />
6. Ostiole eccentric or lateral............................................................... [Campylothelium Müll. Arg.]<br />
5. Ascospores transversely septate .................................................................................................. 7<br />
7. Ascospores less than 100 µm long or if longer, then multiseptate..............................................<br />
................................................................................................................... Trypethelium s. lat.<br />
7. Ascospores mostly more than 100 µm long, 3-5-septate..................[Architrypethelium Aptroot]<br />
4. Ascomata in brown, usually slightly shiny psedostromata containing<br />
yellow or orange pigments, KOH- or KOH+ red; pseudostromal<br />
wall composed of brown, jigsaw puzzle-like hyphae; ascospores<br />
107
transversely septate or muriform.........................................................................................Bathelium<br />
3. Ascomata compound, solitary or in pseudostromata, several hymenia with<br />
a common ostiole or ostiolar plate ........................................................................................................ 8<br />
8. Ascospores with well developed endospore ..................................................................................... 9<br />
9. Ascospores transversely septate or with a few cells longitudinally<br />
divided .......................................................................................................................Astrothelium<br />
9. Ascospores muriform .................................................................................[Cryptothelium Massal.]<br />
8. Ascospores without obvious endospore, submuriform; Brazil.....................[Exiliseptum R. C. Harris]<br />
ASTROTHELIUM Eschw.<br />
Astrothelium as presently constituted is polyphyletic with astrothelioid morphs from several different<br />
clades within the Trypetheliaceae. The clade containing the type species, A. conicum Eschw., is probably<br />
sufficiently distinct that Astrothelium in a restricted sense will be retained. "Species" based on presence or<br />
absence of lichexanthone will be abandoned as in Polymeridium. Some astrothelioid morphs, e.g., A.<br />
variolosum, confusum, do not seem to merit recognition even at the species level. Others will probably be<br />
recognized in Laurera along with much of the rest of the family.<br />
1. Ascomatal warts with orange pigmented tips or containing orange<br />
or brownish pigments, KOH+ purple or reddish ....................................................................................... 2<br />
2. Ascospores 4-celled, IKI-; pigment orange, KOH+ purple .................................................................... 3<br />
3. Thallus UV+ yellow (lichexanthone) .................................................................................................. 4<br />
4. Ascospores 28-35 × 10-13 µm; throughout the state ...............................A. versicolor Müll. Arg.<br />
4. Ascospores 21-28 × 7-9 µm; throughout the state .................................A. galbineum Krempelh.<br />
3. Thallus UV-; ascospores 23-30 × 8-10 µm; throughout the state.........................................................<br />
..............................................................................................A. cinnamomeum (Eschw.) Müll. Arg.<br />
2. Ascospores 10-celled, IKI+ violet, 90-110 × 22-28 µm; pigment<br />
around ostiolar neck brown, KOH+ dull reddish; thallus UV-;<br />
hymenium not inspersed except in ostiolar neck; Collier County................ A. diplocarpum Nyl. s. lat.<br />
1. Ascomatal warts covered by thallus or black where exposed, KOH-..........................................................<br />
.................................................................................................................Trypethelium variolosum Ach.<br />
The astrothelioid morphs of this morphologically and chemically variable species have been recognized<br />
as follows. Some may find it more satisfying to continue to do so.<br />
A. Thallus UV+ yellow (lichexanthone); ascospores 20-25 × 7.5-10 µm;<br />
Franklin, Liberty and Monroe counties (intergrades with<br />
Trypethelium ochroleucum).....................................................................A. variolosum (Ach.) Müll Arg .<br />
A. Thallus UV-; ascospores 20-27 × 8-10 µm; Dade County<br />
(intergrades with Trypethelium nitidiusculum) .....................................................A. confusum Müll. Arg.<br />
Astrothelium diplocarpum Nyl. s. lat.<br />
NOTES<br />
108
The type collection has some ascospores with 1-several cells longitudinally divided and for that reason<br />
the species was placed in Cryptothelium by Zahlbruckner. This, then, is another of those intermediate taxa<br />
that suggest that generic separation between transversely septate and muriform ascospore types is<br />
meaningless. The Florida collection has transverse septa only but it resembles A. diplocarum in large<br />
ascomata, brown pigment (KOH+ reddish), and IKI+ violet ascospores that resemble the non-muriform ones in<br />
the type. In addition to the type from Colombia I have seen only two other collections (Costa Rica and<br />
Ecuador). Since is difficult to adequately interpret the ascospore variation from so few collections, I am<br />
tentatively including the Florida collection in A. diplocarpum.<br />
New to North America. FLORIDA. Collier County: Fakahatchee Strand State Preserve, Royal Palm<br />
Hammock, 5.7 mi from S entrance on W. J. Janes Memorial Scenic Drive, hardwood-Taxodium swamp with<br />
Roystonea, on Ilex, 6 Dec 1992, Harris 29938 (FLAS, NY).<br />
BATHELIUM Ach.<br />
Methodus 111, pl. 8, f. 3. 1803. Type. Bathelium mastoideum Ach.<br />
Bathelium mastoideum has been generally misapplied to a North American species with 4-celled<br />
ascospores but the type has muriform ascospores and ascomatal anatomy of the species known as Laurera<br />
madreporiformis (Eschw.) Riddle (see below). I have always been puzzled by Acharius' illustration as it has no<br />
resemblance to Trypethelium mastoideum auct. The North Italian lichenologists Massalongo (1860) and<br />
Trevisan (1853) correctly understood Bathelium to have muriform ascospores as did Müller initially. I do not<br />
understand why he switched his concept to a 4-celled taxon. It was unfortunate since Zahlbruckner followed<br />
this later misinterpretation and dragged 75 years of lichenologists into error. Dodge attempted to use<br />
Bathelium as a segregate genus for 3-septate species of Trypethelium (a sure guarantee of error) but this was<br />
never accepted. Fortunately the resurrection of Bathelium does not cause nomenclatural problems since B.<br />
mastoideum typifies a distinctive group of taxa that deserve generic recognition. Trypethelium s. str.<br />
encompasses only a few species. The central group, the majority of the family, will eventually be recognized<br />
in Laurera. Jørgensen and Santesson recently proposed Laurera for conservation with a new type, L.<br />
megasperma (Mont.) Riddle. Fortunately it is congeneric with the original holotype, L. varia (Fée) Zahlbr.<br />
although they could not have known that. Such haphazard conservation, changing type species without an<br />
intimate knowledge of the group and possible consequences, is foolish and should be firmly discouraged.<br />
Fortuitously, in this case, it does not cause problems.<br />
Bathelium is distinguished from Laurera and Trypethelium by pseudostromatal characters. The<br />
pseudostromata are somewhat raised to sessile, globose to elongated and irregular in outline, with one to<br />
many ascomata, brown, often shiny, rarely pruinose, with a very distinctive outer layer of brownish short celled,<br />
intricately entwined and branched hyphae forming a jigsaw puzzle-like pattern, pseudostromatal "medulla"<br />
filled with oxalate? and pigment crystals or granules, most often KOH+ red and a cartilaginous, amorphous<br />
cortex-like layer surrounding the individual ascomata within the pseudostroma. The ascospores range from<br />
relatively small (20-30 µm) and transversely 3-septate to very large (200-300 µm) and muriform. Nine species<br />
are currently included but I would anticipate additional taxa being added. There are two species in North<br />
America, both occur in Florida.<br />
1. Ascospores transversely septate, 4-6-celled; hymenium not inspersed ................................................... 2<br />
2. Ascospores 4-celled.............................................................................................................................. 3<br />
3. Ascospores 28 × 9 µm or less........................................................................................................... 4<br />
4. Pseudostromatal pigment KOH bleeding yellow; ascospores<br />
18-28 × 6-9 µm; throughout the state..................................B. carolinianum (Tuck.) R. C. Harris<br />
4. Pseudostromatal pigment bleeding red in KOH; ascospores<br />
20-27 × 8-9 µm; West Indies, Central and South America..............................................................<br />
.............................................................................................. [B.degenerans (Vainio) R. C. Harris]<br />
109
3. Ascospores 35-50 × 12-15 µm; pseudostromata brown, ± shiny<br />
with medulla KOH+ red; Brazil .............................................[B. endochryseum (Vainio) R. C. Harris]<br />
2. Ascospores 6-celled, 25-32 × 6-9 µm; pseudostromata ± sessile, solitary<br />
or 2-3 fused, brown, often somewhat pruinose around ostiole; medulla<br />
orange-brown to brown in section, KOH+ scarlet; Southeast Asia ........................................................<br />
............................................................................................... [B. albidoporum (Mak. & Pat.) R. C. Harris]<br />
1. Ascospores muriform; hymenium inspersed............................................................................................. 5<br />
5. Thallus ± smooth................................................................................................................................... 6<br />
6. Ascospores mostly over 50 µm long; Africa ...................................................................................... 7<br />
7A. Ascospores 44-80 µm long .........................................................................[B. mastoideum Ach.]<br />
7B. Ascospores 100-120 µm long .................................................... [B. lineare (Dodge) R. C. Harris]<br />
7C. Ascospores 130-200 µm long............................................ [B. sphaericum (Dodge) R. C. Harris]<br />
6. Ascospores 40-50 × 12-17 µm, with 10-11 transverse septa;<br />
throughout Florida ............................................................... B. madreporiforme (Eschw.) Trevisan<br />
5. Thallus with subglobose to subdactyliform areoles filled with white<br />
oxalate? crystals and a few bark cells; ascomata nestled between<br />
areoles, brown, with orange crystals in medulla, KOH+ scarlet;<br />
ascospores 90-140 × 27-35 µm; India............................... [B. tuberculosum (Mak. & Pat.) R. C. Harris]<br />
NOTES<br />
Bathelium albidoporum (Makhija & Patwardhan) R. C. Harris, comb. nov.<br />
Trypethelium albidoporum Makhija & Patwardhan, Internatl. J. Mycol. Lichenol. 5: 238. 1992.<br />
Type. [VIETNAM]. Tu-phap, Tonkin, Balansa s.n. (BM, holotype).<br />
Additional specimens. BURMA. Kachin State: along Ledo Road, 21 mi S of Bhamo, 650 ft, Apr<br />
1945, McMillen 245j (US). CHINA. Hainan: Ling-shui, Deng 1991 (BPI).<br />
Bathelium carolinianum (Tuck.) R. C. Harris, comb. nov.<br />
Trypethelium carolinianum Tuck., Amer. Jour. Sci. Arts, ser. 2, 25: 429. 1858. Type. SOUTH<br />
CAROLINA. 1851, Ravenel 298 (FH-Tuck 3952, lectotype, selected here).<br />
This seems to be the oldest name for the taxon previously known as Trypethelium mastoideum. Several<br />
older names have been cited as synonyms. The only one of these I have not re-examined is T. feei Meissner<br />
ex Fée. Trypethelium phlyctaena Fée and T. quassiaecola Fée are synonymous with T. ochroleucum.<br />
Trypethelium scoria Fée is apparently a good species of Trypethlium s. lat.<br />
Bathelium degenerans (Vainio) R. C. Harris, comb. nov.<br />
Pseudopyrenula degenerans Vainio, J. Bot. 34: 292. 1896. Trypethelium degenerans (Vainio)<br />
Zahlbr., Cat. lich. univ. 1: 490. 1922. Type. DOMINICA. Laudat, 1700 ft, Elliot s.n. (BM, isotype).<br />
This species is very close to B. carolinianum but the pseudostromatal pigment reacts KOH+ red. The<br />
pigments need to be analyzed. The KOH reaction may be due to differing pigment concentrations. I have<br />
seen this in other species where a reaction may vary from yellow to red depending on concentration. For the<br />
present I am maintaining them as distinct as there is little overlap in distribution. Bathelium degenerans is not<br />
known from North America but is not uncommon in the Caribbean and Mexico, also known from Brazil, Costa<br />
Rica and Venezuela.<br />
Bathelium endochryseum (Vainio) R. C. Harris, comb. nov.<br />
110
Pseudopyrenula endochrysea Vainio, Étud. class. lich. Brésil 2: 206. 1890. Type. BRAZIL. Minas<br />
Gerais: Caraça, 1885, Vainio, Lich. bras. 1157 (BM, M, isotypes).<br />
Bathelium endochryseum is known only from Brazil.<br />
Bathelium lineare (Dodge) R. C. Harris, comb. nov.<br />
Polyblastiopsis linearis Dodge, Ann. Missouri Bot. Gard. 40: 276. 1953. Laurera linearis (Dodge)<br />
Letrouit-Galinou, Rev. Bryol. Lichénol. 36: 253. 1957. Type. SIERRA LEONE. Njala (Kori), on<br />
Citrus aurantifolius, 27 Nov 1951, Deighton M4634 (FH-Dodge, holotype; BM, isotype).<br />
Bathelium madreporiforme (Eschw.) Trevisan<br />
Trypethelium madreporiforme Eschw., Syst. lich. 24, f. 24a-c. 1824. Laurera madreporiformis<br />
(Eschw.) Riddle in Howe, Torreya 16: 50. 1916. BRAZIL. Serra dos Montes altos, Martius, (M,<br />
holotype?).<br />
Trypethelium madreporiforme f. pruinosa Malme, Ark. bot. 19(1): 23. 1924. Type. BRAZIL.<br />
Mato Grosso: Cuyaba, in cort. Macherii, 28 Mar 1893, Malme, Regn. Lich. 2560A (S, lectotype,<br />
selected here).<br />
This is the only taxon of Bathelium with muriform ascospores in the New World. There is a disjunct<br />
record from Burma. The center of distribution is Africa. It is interesting that ascospores of Burman and North<br />
American collections are slightly smaller than in South American collections. Ascospore size variation in the<br />
group with muriform ascospores is almost continuous and requires more sophisticated reanalysis. The ranges<br />
for B. madreporiforme and B. mastoideum overlap considerably and it may prove necessary to synonymize<br />
them.<br />
Additional specimen. BURMA. Kachin State: along Ledo Road, 21 mi S of Bhamo, 650 ft, Apr 1945,<br />
McMillen 245j (US, mixed in collection of B. albidoporum).<br />
Bathelium mastoideum Ach.<br />
Methodus 111, pl. 8, f. 2-a-c. Type. SIERRA LEONE. Afzelius (S, lectotype; H-Ach 879-E,<br />
isolectotype).<br />
Trypethelium marginatum Fée, Ann. Sci. Nat. Bot. 23: 433, pl. 12, f. 2A-E. 1831. Laurera<br />
marginata (Fée) Dodge, Ann. Missouri Bot. Gard. 40: 297. 1953. Type. SENEGAL. Presque Ile<br />
du cap vert, Perrottet (G, holotype).<br />
Thelenella (Laurera) elegans Vainio, Bull. Soc. Broteriana sér. 2, 6: 174. 1930. Laurera elegans<br />
(Vainio) Zahlbr., Cat. lich. univ. 10: 103. 1938. Type. MOZAMBIQUE. Palma, 1916, Pires de Lima<br />
94 (TUR-Vainio 34704, holotype).<br />
?Laurera submadreporiformis Abbayes, Bull. Inst. Franç. Afr. Noire 15: 52. 1953. Type not<br />
seen.<br />
Polyblastiopsis pyriformis Dodge, Ann. Missouri Bot. Gard. 40: 276. 1953. Laurera pyriformis<br />
(Dodge) Letrouit-Galinou, Rev. Bryol. Lichénol. 26: 252. 1957. Type. SIERRA LEONE. Njala (Kori),<br />
on trunk of Anisophyllea laurina, 26 Oct 1951, Deighton M4404 (FH-Dodge, holotype; BM,<br />
isotype).<br />
Since none of the names cited here which are replaced by Bathelium mastoideum have ever appeared in<br />
the literature more than a couple of times, the change in name is of no consequence. The species is known<br />
only from Africa. Before examining the Acharian material I was puzzled by Acharius's illustration. Now it<br />
makes perfect sense and represents a typical species of the muriform-spored group of Bathelium.<br />
Bathelium sphaericum (Dodge) R. C. Harris, comb. nov.<br />
Polyblastiopsis sphaerica Dodge, Ann. Missouri Bot. Gard. 40: 277. 1953. Laurera sphaerica<br />
(Dodge) Letrouit-Galinou, Rev. Bryol. Lichénol. 26: 254. 1957. Type. SIERRA LEONE. Njala (Kori),<br />
on Orthocosmus africanus, 7 Apr 1952, Deighton M4795 (FH-Dodge, holotype; BM, isotype).<br />
Bathelium tuberculosum (Makhija & Patwardhan) R. C. Harris, comb. nov.<br />
Laurera tuberculosa Makhija & Patwardhan, Mycotaxon 31: 584. 1988. Type. INDIA. Karnataka,<br />
Sirsi-Kumtha Road, 48 km from Sirsi, 12 Dec 1974, Kulkarni 74.2689 (not seen).<br />
Additional specimens. INDIA. Karnataka, near Hebri, humid rain forest, elev. 50 m, 2 Feb 1977,<br />
111
Hale 48878 (US); South Canara, remnants of evergreen forest along road E of Ujire (Udipi<br />
area), 50 m, 22 Feb 1978, Hale 50546 (US).<br />
LAURERA Reichenb.<br />
Eriksson has suggested several times that Cryptothelium should be synonymized here. I agree. I have<br />
not formally done so since most of the family ends up in Laurera and I am waiting to make the transfers as a<br />
package in an organized fashion. Many species of Campylothelium belong here although the type, C.<br />
puiggarii Müll. Arg., is an oddball and might just be worth recognizing in its own genus.<br />
1. Pseudostromata poorly developed, brownish, almost pyrenuloid, KOH-;<br />
ascospores 8/ascus, 90-130 × 20-30 µm; Franklin County..........................................................................<br />
.................................................................................................. L. subdisjuncta (Müll. Arg.) R. C. Harris<br />
1. Ascomatal warts mostly solitary, covered by yellow green thallus<br />
except at apex which usually is black with a whitish ring; ascospores<br />
4/ascus, 200-270 × 30-40 µm, with 50-70 transverse septa;<br />
hymenium inspersed; peninsular Florida................................................. L. megasperma (Mont.) Riddle<br />
POLYMERIDIUM (Müll. Arg.) R. C. Harris<br />
1. Ascospores transversely septate .............................................................................................................. 2<br />
2. Ascospores 4-celled............................................................................................................................... 3<br />
3. Thallus UV+ yellow (lichexanthone); ascospores 22-32 × 7-10 µm;<br />
hymenium not inspersed; common throughout the state .............. P. catapastum (Nyl.) R. C. Harris<br />
3. Thallus UV-......................................................................................................................................... 4<br />
4A. Ascospores 13-15.3-17 × 4-5-6 µm; Polk and Sarasota counties ..................................................<br />
.................................................................................................P. contendens (Nyl.) R. C. Harris<br />
4B. Ascospores 16-18.5-21 × 5-5.8-7 µm; throughout the state ...........................................................<br />
...............................................................................................P. subcinereum (Nyl.) R. C. Harris<br />
4C. Ascospores 19-22.3-25 × 5.5-6.6-7.5 µm; Dade County ................................................................<br />
............................................................................................. P. albidum (Müll. Arg.) R. C. Harris<br />
2. Ascospores 5-11-celled......................................................................................................................... 5<br />
5. Ascospores (5-)6-8(-9)-celled, 18-22.5-27 × 4-5.5-7 µm; throughout the state ..................................<br />
...........................................................................................P. quinqueseptatum (Nyl.) R. C. Harris<br />
5. Ascospores (6-)8-10(-11)-celled, 24-32.3-40 × 5.5-7-8.5 µm; Dade County.......................................<br />
.......................................................................................P. albocinereum (Krempelh.) R. C. Harris<br />
1. Ascospores muriform, amyloid, 39-54 × 15-19 µm, (9-)11(-13) × 1-4-septate<br />
thallus UV+ yellow (lichexanthone) or UV-; hymenium not inspersed;<br />
south to Marion County ......................................................................... P. proponens (Nyl.) R. C. Harris<br />
PSEUDOPYRENULA Müll. Arg.<br />
Pseudopyrenula subnudata Müll. Arg. (1883) antedates P. subgregaria Müll. Arg. (1885). There is only<br />
the one species in North America.<br />
112
TRYPETHELIUM Sprengel<br />
1. Thallus or ascomatal wart KOH+ purple ................................................................................................... 2<br />
2. Thallus usually yellowish or brownish, KOH-; ascomata aggregated<br />
in raised pseudostromata which are often constricted at base,<br />
contents KOH+ purple.......................................................................................................................... 3<br />
3. Ascospores 10-14-celled, 40-50 × 9-12 µm; powder in pseudostromata<br />
yellow to tan; pseudostromata and/or thallus often pruinose;<br />
peninsular Florida..............................................................................................T. eluteriae Sprengel<br />
3. Ascospores 13-16-celled, 60-75(-85) × 11-12 µm; powder in<br />
pseudostromata orange; pseudostromata orange to brown,<br />
not pruinose; Alachua, Gilchrist, Polk and Sarasota counties..............................................................<br />
................................................................................................................. T. subeluteriae Mak. & Pat.<br />
2. Thallus orange; ascomata mostly immersed, solitary and scattered<br />
to aggregated; ascospores 4-celled, 21-28 × 8-9 µm; peninsular Florida ...............................................<br />
.................................................................................................................. T. aeneum (Eschw.) Zahlbr.<br />
1. Thallus and warts KOH- ............................................................................................................................ 4<br />
4. Ascomata covered by thallus or in pseudostroamta ......................................................................... 5<br />
5. Ascospores 4-celled, IKI- .............................................................................................................. 6<br />
6. Ascospores smaller, less than 20-27 × 7.5-10 µm long ..............................................................<br />
....................................................................................................Trypethelium variolosum Ach.<br />
The trypethelioid morphs with pseudostromata containing 1-many simple ascomata have been<br />
recognized as follows. Some may find it more satisfying to continue to do so.<br />
A. Thallus UV+ yellow (lichexanthone);throughout the state<br />
........................................................................................................ T. ochroleucum (Eschw.) Nyl.<br />
A. Thallus UV-; throughout the state.................................................T. nitidiusculum (Nyl.) R. C. Harris<br />
6. Ascospores large, 35-43 × 12-15 µm; ascomata solitary or<br />
grouped, covered by yellow green thallus except at apex which<br />
is black with a pale ring; hymenium not inspersed; peninsular Florida .......................................<br />
........................................................................ T. floridanum (Zahlbr. ex Choisy) R. C. Harris<br />
5. Ascospores 8-12-celled, 38-52 × 7-10 µm, IKI+ weak violet;<br />
pseudostromata not raised; south to Marion County........................T. virens Tuck. ex Michener<br />
4. Ascomata black, naked, solitary or aggregated; hymenium inspersed;<br />
ascospores 4-celled, 20-26 × 6-8 µm; throughout the state................................................................<br />
............................................................................................................T. tropicum (Ach.) Müll. Arg.<br />
NOTES<br />
Trypethelium aeneum (Eschw.) Zahlbr.<br />
Trypethelium ferrugineum Müll. Arg., Bot. Jahrb. Syst. 6: 392. 1885. Type. CUBA. Wright, Verr.<br />
Cub. 157, Lich. Cub. II, 593, 594 (FH-Tuck 3972, isosyntypes), syn. nov.<br />
Trypethelium ferrugineum β. inornatum Müll. Arg., Bot. Jahrb. Syst. 6: 392. 1885. Type. CUBA.<br />
113
Wright, Verr. Cub. 157 (G, holotype; US, isotype), syn. nov.<br />
Trypethelium ferrugineum represents the astrothelioid morph of T. aeneum analogous to "Astrothelium"<br />
confusum or variolosum in the Trypethelium variolosum aggregate. It has never been combined in<br />
Astrothelium and since I believe these astrothelioid morphs to be of little significance, I will not do so. Others<br />
may wish to. If one were a real hardcore lumper, T. aeneum, could also be included in the T. variolosum<br />
group as chemotype with anthraquinone pigments instead of lichexanthone.<br />
Trypethelium subeluteriae Makhija & Patwardhan<br />
Trypethelium eluteriae var. polystomum Malme, Ark. Bot. 19(1): 25. 1924. Type. BRAZIL. Mato<br />
Grosso: Buriti, in cort. Platypodii, 16 Jun 1894, Malme s.n. (S, lectotype, selected here), syn.<br />
nov.<br />
This taxon was noted in Harris (1986) and in "Some Florida Lichens" as Trypethelium sp. aff. eluteriae.<br />
Unfortunately Makhija and Patwardhan ran across a really scrappy specimen from Sri Lanka that I had so<br />
annotated and used it as the type for this widespread species. Infinitely better material was available. The<br />
species is not confined to Sri Lanka as they would imply. It is known from Australia, Central America (Costa<br />
Rica), India, Indochina, Indonesia, Papua New Guinea, Philippines, South America (Bolivia, Brazil, Paraguay,<br />
Venezuela), West Indies (Cuba, Dominican Republic) and Florida.<br />
New to North America. FLORIDA. Alachua County: Paynes Prairie State Preserve, Bolen Bluff Trail off US<br />
441, ca. 3.5 mi N of main entrance, 29°34'N, 82°21'W; dry hardwoods with Sabal pametto, 1 Dec 1992, Buck<br />
22561; Lafayette County: along Fla Hwy 51, 5.7 mi S of US 27 at Mayo, 29°59'N, 83°14'W; hardwood swamp,<br />
29 Nov 1994, Buck 27089; Polk County: along US 17, 2 mi N of Ft. Meade, 2 Feb 1965, Johnson 6205, 6209;<br />
Sarasota County: Myakka State Park, 31 Jan 1965, Johnson 6186 (all NY).<br />
Trypethelium tropicum (Ach.) Müll. Arg.<br />
A collection from Lake County (Harris 23579, NY) has an unusual, thick, bullate-rugose areolate thallus<br />
with a very thick medulla filled with white oxalate? crystals. The ascomata, hamathecium and ascospores are<br />
perfectly typical T. tropicum. I have speculated that thallus appearance is often substrate dependent. This<br />
collection is on Acer rubrum which is a common substrate for "typical" T. tropicum, so that doesn't seem to be<br />
the case here. I include this comment so that I won't in future just sweep it under the rug.<br />
Trypethelium variolosum Ach.<br />
After 20 years and hundreds, if not thousands, of specimens I have come to the conclusion that there is a<br />
complete series of intermediates between the trypethelioid and astrothelioid morphs in this aggregate species.<br />
The astrothelioid ascoma in this taxon seems to be derived from the invagination of the ascomatal wall (not<br />
fusion of individual ascomata). All stages can be found, from basal, incomplete, non-melanized invaginations<br />
to complete, melanized dividing walls. Since I am tired of flipping a coin to decide if the invaginations are<br />
complete enough to warrant placement in Astrothelium s. lat. or not (there is even some variation on a single<br />
thallus) and since the amount of invagination seems to have no predictive value, I have decided to merge the<br />
two "morphs". Also, I have concluded that here (and throughout the family) the presence or absence of<br />
lichexanthone should not be used as a specific character (Harris, 1993). (Almost every lichexanthone<br />
containing taxon has some lichexanthone deficient collections.) Further, some collections in this group could<br />
be segregated on the basis of a large amount of cartilaginous tissue in the pseudostromata as T. leprieurii.<br />
Again, I find a complete gradation from essentially none to comprising most of the pseudostroma. Two further<br />
characters which do not seem to correlate in any meaningful way are hymenial inspersion and oxalate crystals<br />
in the ascomatal wall. As far as I can tell there is not the slightest correlation between any of these characters<br />
and ecology or geographic distribution. This complex seems to represent one of those cases where the<br />
extremes are distinct but the middle ground is vast and messy. Using the above characters in all possible<br />
combinations (most of which I have run across) one could recognize dozens of "species". The oldest name<br />
for a member of this aggregate is Trypethelium variolosum Ach., an astrothelioid morph containing<br />
lichexanthone. Trypethelium variolosum occurs throughout Florida. The following segregates have previously<br />
been recognized and some may wish to continue to do so.<br />
1). Trypethelium ochroleucum (Eschw.) Nyl. - Asomata solitary to aggregated in trypethelioid<br />
pseudostromata; lichexantone present; common, throughout Florida. Those who wish to recognize this<br />
commonly used name will have to protect it against a couple of older Fée names.<br />
114
2). Trypethelium variolosum Ach., Astrothelium variolosum (Ach.) Müll. Arg. - Ascomata astrotheliod;<br />
lichexanthone present; throughout Florida.<br />
3). Trypethelium nitidiusculum (Nyl.) R. C. Harris - Ascomata solitary to aggregated in trypethelioid<br />
pseudostromata; lichexanthone absent; common, throughout Florida. Again there are older names but<br />
nitidiusculum has been taken up so recently (1986) that I see no point in protecting it.<br />
4). Astrothelium confusum Müll. Arg. - Ascomata astrothelioid; lichexanthone absent; Dade County.<br />
5). Trypethelium leprieurii Mont. - Ascomata trypethelioid with much cartilaginous tissue in the<br />
pseudostromata; lichexanthone present. I have never segregated any North American material into this<br />
morph but I don't doubt that one could. My awareness of this morphotype is fairly recent.<br />
MONOBLASTIACEAE W. Watson<br />
New Phytol. 28: 106. 1929. Type. Monoblastia Riddle<br />
Acrocordiaceae Oxner ex Barr, Mycotaxon 29: 505. 1987. Type. Acrocordia Massal.<br />
Additional evidence has been found to support the recognition of the Monoblastiaceae. Sabine Huhndorf,<br />
attempting to place the genus Ascocratera Kohlmeyer, examined the asci of a number of species for<br />
fluorescent structures in the ascus tip. Those taxa examined uniformly had either a fluorescent ring or<br />
fluorescent cap. Two species of Acrocordia, two species of Monoblastia and two of Anisomeridium had a<br />
fluorescent cap. Four species of Anisomeridium had fluorescent rings. Those with caps included the type<br />
species of Acrocordia, Anisomeridium and Monoblastia. The type species of Ditremis had a distinct ring.<br />
(Ascocratera had a ring but considering all characters one reaches the impasse that there is no definitive<br />
reason to either include it in or exclude it from the Monoblastiaceae).<br />
KEY TO GENERA<br />
1. Ascospores 2-4-celled, granular ornamented or smooth .......................................................................... 2<br />
2. Asci various with narrow ocular chamber; ascospores in a single row,<br />
in two rows or irregularly arranged, 2-4-celled, colorless or brown,<br />
ornamented or smooth; more common in tropical regions ....................................................................3<br />
3. Microconidia and macroconidia borne in pycnidia......................................................Anisomeridium<br />
3. Microconidia pycnidial; macroconidia in campylidia ....................................................... [Musaespora]<br />
2. Asci cylindrical with very broad ocular chamber; ascospores<br />
in a single row in the ascus, colorless, 2-celled, oblong,<br />
granular ornamented; temperate regions ............................................................................. Acrocordia<br />
1. Ascospores simple, ornamentation wart-like or peg-like........................................................ Monoblastia<br />
ACROCORDIA Massal.<br />
1. Growing on bark ........................................................................................................................................ 2<br />
2. Ascospores larger, 18-60 × 9-23 µm..................................................................................................... 3<br />
3. Ascospores 18-27 × 9-13 µm; ostiole usually apical;<br />
Duval County......................................................................................... A. gemmata (Ach.) Massal.<br />
3. Ascospores 33-48(-60) × 15-23 µm; ostiole usually eccentric;<br />
eastern North America ...............................................................[A. megalospora (Fink) R. C. Harris]<br />
115
2. Ascospores small, 11-17 × 6-9.5 µm; eastern North<br />
America ....................................................................................................[A. cavata (Ach.) R. C. Harris]<br />
1. Growing on calcareous rock; ascospores 13-17 × 7-8.5 µm;<br />
Michigan ............................................................................................................. [A. conoidea (Fr.) Körber]<br />
NOTES<br />
Acrocordia gemmata (Ach.) Massal<br />
The Florida collection is the only North American record and was not included in my thesis. One wishes<br />
it were made by someone other than Calkins who does not have a good track record for relaibility.<br />
FLORIDA. Duval County: Jacksonville, on cypress, Calkins, N.A.L 238 (NY).<br />
ANISOMERIDIUM (Müll. Arg.) Choisy, nom. cons.<br />
Icon. Lich. Univ. fasc. 3 (unpaged). 1926. Arthopyrenia sect. Anisomeridium Müll. Arg., Flora 66:<br />
290. 1883. Lectotype (Riedl, 1963). Arthopyrenia xylogena Müll. Arg. (= Anisomeridium<br />
subnectendum (Nyl.) R. C. Harris).<br />
Lembidium Körber, Syst. Lich. Germ. 358. 1855, nom. rejic. Type. Lembidium polycarpum Körber (=<br />
Anisomeridium biforme (Borrer) R. C. Harris).<br />
Ditremis Clements, Genera of Fungi 41. 1909. Type. Pleurotrema inspersum Müll. Arg. (=<br />
Anisomeridium americanum (Massal.) R. C. Harris).<br />
Sarcinulella Sutton & Alcorn, Mycotaxon 16: 557. 1983, nom. illeg.? Type. Sarcinulella banksiae Sutton &<br />
Alcorn (= Anisomeridium nyssaegenum (Ellis & Everhart) R. C. Harris?).<br />
Megalotremis Aptroot, Biblioth. Lichenol. 44: 124. 1991. Type. Ditremis verrucosa Makhija & Patwardhan<br />
(= Anisomeridium verrucosum (Makhija & Patwardhan) R. C. Harris).<br />
I had originally intended to recognize both Anisomeridium and Ditremis in this volume, based partly on<br />
evidence from fluorescence studies of ascus tips by Sabine Huhndorf. Unfortunately Murphy's Law applies<br />
and the new data does not correlate well with other characters. I am confident that with more species studied<br />
for tedious and often hard to interpret ascus and anamorph characters, it will be possible to subdivide<br />
Anisomeridium s. lat. into several genera. The largest group evident at this time is that centered around the<br />
type of Anisomeridium, with thallus lacking lichexanthone, apical ostioles, clavate or saccate asci with a<br />
fluorescent cap, blunt, ± ovoid, often thick-walled ascospores and globose microconidia. The other, less<br />
cohesive, group centers around the type of Ditremis with thallus sometimes with lichexanthone, often eccentric<br />
ostioles, cylindrical asci with a fluorescent ring? (only one sampled), ascospores with ± pointed ends and<br />
oblong or elliptical microconidia. Anisomeridium biforme has a fluorescent cap and redisposition into<br />
Acrocordia should be considered. Unfortunately the ubiquitous A. nyssaegenum seems to stand alone. It is<br />
also possible that some taxa may be removed to Musaespora.<br />
Being unable to recognize two genera at this time, I have decided to accept the decision of the North<br />
European lichen oligarchs to force their will on the rest of the world and conserve Anisomeridium. (There are<br />
only three species in North Europe but 100+ in the rest of the world. Makhija & Patwardhan (1990) recognized<br />
18 species of Ditremis for India alone. As will be seen from the general treatment below, I was in fact very<br />
conservative in making combinations in Ditremis.) When I fought for Ditremis in "Some Florida Lichens", I was<br />
aware that this was not a battle I would win but I felt obliged to protest the rape of the International Code of<br />
Nomenclature by nomenclatural Luddites fearing change (mostly the result of a shift from an artificial system<br />
to a phylogentic system, already too long postponed in mycology). Such unfortunate precedents may,<br />
however, help win the war in St. Louis.<br />
KEY TO ANISOMERIDIUM IN FLORIDA<br />
116
1. Ascomata present; pycnidia present or absent, never with long, hairlike beaks....................................... 2<br />
2. Thallus UV-............................................................................................................................................ 3<br />
3. Ascospores 2-celled; ostiole apical or eccentric; growing on bark....................................................4<br />
4. Ostiole apical................................................................................................................................. 5<br />
5. Ascospores 12-24 × 4.5-8.5 µm ................................................................................................ 6<br />
6. Asci slender, cylindrical to narrowly clavate .......................................................................... 7<br />
7. Ascospores slender, narrowly ovate, fusiform to almost<br />
cylindrical (often slightly bent in A.tamarindi) .................................................................... 8<br />
8. Microconidia elliptical, 2.5-1.5 × 2 µm; macroconidia ± oblong,<br />
5-7 × 2.5-3 µm; ascospores narrowly ovoid, with weakly<br />
submedian septum; 12-18(-21) × 4.5-6.5 µm; neotropical<br />
north to Lake Couny............................................... A. subprostans (Nyl.) R. C. Harris<br />
8. Microconidia orbicular; macroconidia elliptical,<br />
4-7 × 2.5-3.5 µm; ascospores 15-22(-24) ×<br />
4.5-5.5(-6) µm, often slightly bent; north to<br />
Volusia County .............................................................A. tamarindi (Fée) R. C. Harris<br />
7. Ascospores broadly elliptical to ovoid, rarely narrowly ovoid,<br />
with septum slightly submedian or ± median,10-18 × 4.5-7 µm;<br />
microconidia oblong or elliptical, 2.5-3 × 1.5-2 µm or globose,<br />
ca. 2 µm diam.; macroconidia elliptical or oblong, 5-7 × 3-4 µm;<br />
throughout Florida .......................................................... A. biforme (Borrer) R. C. Harris<br />
6. Asci ovate or obovate to narrowly ovate or obovate.............................................................. 9<br />
9. Ascospores relatively broad, 5.5 µm or more.................................................................. 10<br />
10. Ascus wall and ascospore wall thick and unevenly thickened;<br />
ascospores ovoid, with markedly submedian septum,<br />
15-22 × 6.5-8.5 µm; microconidia globose, ca. 2 µm diam;<br />
macroconidia oblong or ovoid, 5-7 × 2.5-3 µm; Dade<br />
County ..........................................................A. anisolobum (Müll. Arg.) R. C. Harris<br />
10. Ascus wall and ascospore wall relatively thin and even;<br />
ascospores 18-22 × 5.5-7 µm, ± fusiform, often slightly<br />
bent, with slightly submedian septum; microconidia globose<br />
or elliptical, ca. 2 µm diam. or 2-3 × 1.5-2 µm; Seminole<br />
County ..........................................................A. excaecariae (Müll. Arg.) R. C. Harris<br />
9. Ascospores relatively narrow, 14-20 × 4.5-6 µm, ovoid, with<br />
submedian septum, often becoming 4-celled; micropycnidia<br />
beaked ("Sarcinulella"); microconidia oblong or narrowly elliptical,<br />
2-3 × 1-1.5 µm; Columbia County .......................................................................................<br />
..............................................................A. nyssaegenum (Ellis & Everhart) R. C. Harris<br />
117
5. Ascospores small, 9-13(-15) × 4-5 µm, ovoid, with septum<br />
median to slightly submedian; ascomata small, 0.2-0.3(-0.4) mm<br />
diam.; microconidia oblong or elliptical, Duval, Liberty and Seminole<br />
counties ..................................................................................... A. albisedum (Nyl.) R. C. Harris<br />
4. Ostiole eccentric............................................................................................................................ 11<br />
11. Ascospores granular ornamented, fusiform, with septum moderately<br />
submedian, 19-27(-30) × 5.5-7(-8) µm; microconidia oblong or<br />
elliptical, 2.5-3 × 1.5-2 µm; ascomata usually naked; Collier and<br />
Monroe counties................................................................... A. terminatum (Nyl.) R. C. Harris<br />
11. Ascospores smooth, ± ovoid ................................................................................................... 12<br />
12. Ascospore septum distinctly submedian; ascospores<br />
32-40(-45) × 13.5-18 µm; microconidia subglobose,<br />
ca. 2-2.5 µm diam.; macroconidia elliptical or oblong,<br />
5-7 × 2.5-4 µm; ascomata covered by thallus; Duval,<br />
Franklin and Marion counties ...........................................................A. griffinii R. C. Harris<br />
12. Ascospore septum ± median; ascospores 17-23 × 7-9 µm;<br />
pycnidia not found; Levy County..................................Anisomeridium sp. Harris 29392-A<br />
3. Ascospores 4-celled or if 2-celled, growing on rock; ostiole always apical<br />
(2-celled, on rock, see also Pyrenocollema and Zwackhiomyces) ................................................... 13<br />
13. Ascospores 4-celled.................................................................................................................... 14<br />
14. Growing on bark .................................................................................................................... 15<br />
15. Ascospores soon 4-celled ............................................................................................... 16<br />
16. Ascospores 21-24 × 6-8 µm; microconidia elliptical, 3-4 ×<br />
1.5-2 µm; Columbia County........................................ A. quadrococcum R. C. Harris<br />
16. Ascospores16-20 × 3.5-4.5 µm; microconidia ± globose,<br />
ca. 2 µm diam.; macroconidia oblong, 5-6 × 2-3 µm; Louisiana,<br />
not yet found in Florida............................ [A. quaternarium (R. C. Harris) R. C. Harris]<br />
15. Ascospores tardily 4-celled, ovoid, 14-20 × 4.5-6 µm;<br />
Columbia County...................................... A. nyssaegenum (Ellis & Everh.) R. C. Harris<br />
14. Growing on sandstone; ascospores tardily 4-celled, ovoid, with first<br />
septum submedian, smooth, 16-22 × 6.5-8 µm; microconidia narrowly<br />
elliptical, 4-5 × 1.5-2 µm; macroconidia ovoid or elliptical, 5-7 × 2.5-3 µm;<br />
Massachusetts to Illinois south to southern Georgia and Louisiana<br />
but not yet found in Florida .................................................... [A. distans (Willey) R. C. Harris]<br />
13. Ascospores 2-celled.................................................................................................................... 17<br />
17. Growing on limestone; ascospores narrowly ovoid to<br />
fusiform, with submedian septum, smooth, 20-25 ×<br />
6.5-8.5 µm; microconidia elliptical, 3-5 × 1-1.5 µm;<br />
macroconidia broadly elliptical, 5-6 × 4-4.5 µm; Dade<br />
and Monroe counties........................................................ A. finkii (R. C. Harris) R. C. Harris<br />
17. Growing on sandstone; "immature" forms, see couplet<br />
118
14 above.................................................................................. [A. distans (Willey) R. C. Harris]<br />
2. Thallus UV+ yellow (lichexanthone) ..................................................................................................... 18<br />
18. Ascospores over 20 µm long.......................................................................................................... 19<br />
19. Ostiole eccentric, ringed with orange pigment, KOH+ purple;<br />
ascospores biseriate, fusiform, with septum submedian,<br />
smooth, 37-52 × 12-20 µm; microconidia oblong, 3-4 × 1.2-1.5 µm;<br />
macroconidia broadly elliptical to ovoid, 30-45 × 15-20 µm;<br />
Dade County............................................................................A. aureopunctatum R. C. Harris<br />
19. Ostiole apical, unpigmented; ascospores mostly uniseriate,<br />
± fusiform with one end often more rounded, with septum<br />
median or submedian, smooth, 24-33 × 8-12 µm;<br />
microconidia elliptical, 3-5 × 1.5-2 µm; macroconidia<br />
broadly ovoid to ovoid, thick-walled,15-22 × 9-16 µm;<br />
peninsular Florida............................................................. A. ambiguum (Zahlbr.) R. C. Harris<br />
18. Ascospores less than 20 µm long .................................................................................................. 20<br />
20. Ascospores colorless ............................................................................................................... 21<br />
21. Ascospores fusiform, with slightly submedian septum, smooth,<br />
15-20 × 5-6.5 µm; macroconidia cylindrical 7.5-12 × 2-2.5 µm;<br />
microconidia elliptical to subglobose, 2.5-3 × 1.5-2 µm or ca. 2 µm diam.;<br />
peninsular Florida, Lousiana ................................... A. tuckerae (R. C. Harris) R. C. Harris<br />
21. Ascospores broadly elliptical, not constricted at median septum,<br />
smooth, (10-)12-15 × (6-)7.5-9 µm; thallus ± corticate; microconidia<br />
broadly elliptical 3 × 1.5-2 µm; Okefenokee Swamp, Georgia and<br />
Polk County .............................................................................. A. biformoides R. C. Harris<br />
20. Ascospores soon dark brown, elliptical or ends slightly pointed,<br />
with septum ± median, granular ornamented, uniseriate in ascus,<br />
16-20 × 8-9 µm; thallus white, UV+ yellow (lichexanthone); ostiole apical;<br />
Collier County........................................................................... A. phaeospermum R. C. Harris<br />
1. Ascomata not known; pycnidia with hair-like beaks, to 1 cm long; macroconidia<br />
ellipsoid to narrowly ellipsoid, 5-7 × 2-2.5 µm, in ± packets surrounded by a<br />
gelatinous sheath; Collier County...........................................................Anisomeridium sp. Buck 22812<br />
NOTES<br />
Anisomeridium albisedum (Nyl.) R. C. Harris<br />
in Egan, Bryologist 90: 163. 1987. Verrucaria viridiseda f. albiseda Nyl., Lich. Japon. 108.<br />
1890. Porina viridiseda f. albiseda (Nyl.) Zahlbr., Cat. lich. univ. 1: 409. 1922. Ditremis<br />
albiseda (Nyl.) R. C. Harris, Some Florida Lichens 31. 1990. Type. FLORIDA. Calkins (H- Nyl<br />
717, lectotype; MICH, US, isolectotypes).<br />
Anisomeridium ambiguum (Zahlbr.) R. C. Harris<br />
in Egan, Bryologist 90: 163. 1987. Arthopyrenia (Acrocordia) ambigua Zahlbr., Ann. Mycol. 3:<br />
35. 1935. Ditremis ambigua (Zahlbr.) R. C. Harris, Some Florida Lichens 31. 1990. Type. FLORIDA.<br />
Seminole County: Sanford, on elm, May 1923, Rapp 142 (W-2452, holotype).<br />
119
Anisomeridium anisolobum (Müll. Arg.) Aptroot<br />
Biblioth. Lichenol. 57: 21. 1995. Arthopyrenia anisoloba Müll. Arg., Flora 66: 305. 1883. Ditremis<br />
anisoloba (Müll. Arg.) R. C. Harris, Some Florida Lichens 31. 1990. Type. BRAZIL. São Paulo:<br />
Apiaí, 1882, Puiggari (G, holotype).<br />
Arthopyrenia feeana Müll. Arg., Mém. Soc. Phys. Hist. Genève 30(3): 28. 1888. Anisomeridium<br />
feeanum (Müll. Arg.) R. C. Harris in Tucker & Harris, Bryologist 83: 4. 1980. Type. WEST INDIES.<br />
On Croton cascarillae, ex hb. Fée (G, holotype).<br />
Arthopyrenia (Mesopyrenia) knysana Zahlbr., Ann. Crypt. Exot. 5: 200. 1932. Type. SOUTH<br />
AFRICA. Cape Province: Knysna, on bark of Acacia melanoxylon, v. d. Byl 916 (LD, isotype).<br />
Anisomeridium aureopunctatum R. C. Harris, nom. nov.<br />
Ditremis macrospora R. C. Harris, Some Florida Lichens 33. 1990. Type. FLORIDA. Dade<br />
County: Everglades National Park, NE of Pa-hay-okee Overlook, 13 May 1967, Harris 2791-A<br />
(MSC, holotype), non Makhija & Patwardhan, Biovigyanam 16: 19. 1990 (before? 21<br />
December).<br />
Anisomeridium biforme (Borrer) R. C. Harris<br />
in V_zda, Lich. sel. exs. fasc. 61: 1305. 1978. Verrucaria biformis Borrer in Hooker & Sowerby,<br />
Suppl. Engl. Bot. 1: pl. 2617, f. 1. 1831. Arthopyrenia biformis (Borrer) Massal., Framm. Lich.<br />
26. 1855. Ditremis biformis (Borrer) R. C. Harris, Some Florida Lichens 32. 1990. Type not<br />
seen.<br />
Verrucaria conformis Nyl., Flora 47: 357. 1864; Arthopyrenia conformis (Nyl.) Müll. Arg., Bull.<br />
Herb. Boissier 2(app. 1): 90. 1894. Type. FRANCE. Finistère, Brest, Crouan 28 (H-Nyl 1653,<br />
lectotype).<br />
Verrucaria alboatra Krempelh. ex Nyl., Flora 50: 8. 1867. Arthopyrenia alboatra (Krempelh. ex<br />
Nyl.) Müll. Arg., Flora 66: 306. 1883. Type. INDIA. West Bengal, Calcutta Botanic Gardens,<br />
1866, Kurz 10 (H-NYL 1660, lectotype; BM, M, isolectotypes).<br />
Arthopyrenia brachyspora Müll. Arg., Nuovo Giorn. Bot. Ital. 21: 52. 1889. Type. ARGENTINA. Isla<br />
de los Estados, Spegazzini (G, holotype).<br />
Arthopyrenia parvula Zahibr., Beih. Bot. Centralbl. 13: 149. 1902. Type. UNITED STATES.<br />
California: Santa Monica Mountains, Malibu Canyon, Aug. 1898, Hasse 759 (FH, MIN, US,<br />
isotypes).<br />
Didymella microspora Vainio, Ann. Acad. Sci. Fenn. A 19(15): 15.1923. Type. PHILIPPINE ISLANDS.<br />
Comiran Island: Sulu Sea, Sep 1910, Merrill, Bur. Sci. 7165 (TUR-Vainio 32293, holotype).<br />
Arthopyrenia amaura Zahlbr. in Handel-Mazzetti, Symb. Sinic. 3: 19. 1930. Type. CHINA. Hunan,<br />
near Tschangscha, 10 Dec 1917, Handel-Mazzetti 11426 (US, isotype).<br />
Didymella laurilae Räsänen, Ann. Bot. Soc. Zool. Bot. Fenn. Vanamo 12(1): 204. 1939. Type.<br />
FINLAND. Par. Kaukola, Heinsalmi, in ripa boreale lacus Ruokajärvi, ad lignum madidum in<br />
aquam dejectum in luco ripario, 25 Jul 1935, Laurila (H, lectotype; H, isolectotype).<br />
Anisomeridium biformoides R. C. Harris, sp. nov.<br />
Similis A. biformis quoad characteribus microscopicis differt thallo UV+ aureo.<br />
Type. GEORGIA. Charleton County: Okefenokee National Wildlife Refuge, Chesser Island, 15 Nov 1989,<br />
Wetmore 64362 (NY, holotype; MIN, isotype, not seen).<br />
Thallus pale gray, endophloeodal, UV+ yellow (lichexanthone). Ascomata immersed, tip exposed,<br />
subglobose, ca. 0.3-0.5 mm diam.; clypeus apical and shield shaped to hemispherical, occasionally slightly<br />
laterally expanded. Asci cylindrical, ca. 120-135 × 12-15 µm, with eight uniseriate spores. Ascospores broadly<br />
elliptical, not constricted at median septum, smooth, (10-)12-15 × (6-)7.5-9 µm; microconidia elliptical, 3 × 1.5-<br />
2 µm; macroconidia not found.<br />
120
This species was treated in 1990 as Ditremis sp. Wheeler 15-4-89 from Polk County. Better material<br />
collected by Wetmore in Georgia makes its recognition here warranted.<br />
Additional specimen. FLORIDA. Polk County: Nalcrest, W side of recreation area, on fallen oak branch, 15<br />
Apr 1989, Wheeler s.n. (NY).<br />
Anisomeridium distans (Willey) R. C. Harris<br />
in Egan, Bryologist 90: 163. 1987. Verrucaria distans Willey, Enum. Lich. New Bedford 38.<br />
1892. Arthopyrenia distans (Willey) Zahlbr., Cat. Lich. Univ. 1: 276. 1921. Ditremis distans<br />
(Willey) R. C. Harris, Some Florida Lichens 32. 1990. Type. MASSACHUSETTS. Bristol County:<br />
New Bedford, 1866, Willey (MICH, isotype).<br />
Anisomeridium excaecariae (Müll. Arg.) R. C. Harris, comb. nov.<br />
Sagedia excaecariae Müll. Arg., Flora 63: 289. 1880. Arthopyrenia excaecariae (Müll. Arg.)<br />
Müll. Arg., Flora 66: 305. 1883. Type. ARGENTINA. Buenos Aires, on Excaecaria biglandulosa,<br />
Schnyder 2 (G, holotype).<br />
Arthopyrenia incurva Müll. Arg., Flora 66: 305. 1883. Type. BRAZIL. São Paulo: Apiaí, Mar<br />
1880, Puiggari 138.b pr. p. (G, holotype).<br />
Arthopyrenia sanfordensis Zahlbr., Ann. Mycol. 33: 33. 1935. Anisomeridium sanfordense<br />
(Zahlbr.) R. C. Harris in Egan, Bryologist 90: 163. 1987. Ditremis sanfordensis (Zahlbr.) R. C.<br />
Harris, Some Florida Lichens 34. 1990. Type. FLORIDA. Seminole County: Sanford, on<br />
persimmon, Apr 1932, Rapp 134 (W-2460, holotype).<br />
Anisomeridium finkii (R. C. Harris) R. C. Harris, comb. nov.<br />
Ditremis finkii R. C. Harris, Some Florida Lichens 32. 1990. Type. PUERTO RICO. Arecibo District:<br />
Manati, 12 Jan 1916, Fink 2052 (US, holotype; FH, isotype).<br />
Anisomeridium griffinii R. C. Harris, sp. nov.<br />
Thallo albo, UV-, ostiolo eccentrico, ascosporis ovoideis vel interdum late fusiformibus, laevibus, septis<br />
distincte submedianis, 32-38 × 13.5-18 µm et macroconidiis ovoideis, 5-7 × 2.5-4 µm.<br />
Type. FLORIDA. Marion County: ca. 0.5 mi S of U.S. 40 & 1 mi W of Oklawaha River bridge, mesic hammock<br />
of Liquidambar styraciflua, Celtis laevigata, Sapindus marginata & Quercus virginiana, on trunk of Ulmus<br />
crassifolia, 23 May 1994, Griffin 3217 (NY, holotype; ESS, FLAS, H, isotypes).<br />
Thallus white, endophloeodal, UV-. Ascomata covered by thin layer of thallus, ± lageniform, ca. 0.5-0.6 ×<br />
0.4 mm; ostiole eccentric to lateral; wall entirely melanized or only to base of hamathecium. Asci cylindrical to<br />
subclavate, 150-210 × 30-40 µm, with eight biseriate spores. Ascospores ovoid or somewhat fusiform, slightly<br />
constricted at the markedly submedian septum, smooth, 32-40 × 12-18 µm; microconidia subglobose, ca. 2<br />
µm diam.; macroconidia ovoid, 5-7 × 2.5-4 µm.<br />
It is a genuine pleasure to name this species to honor its collector, Dana Griffin, III, who has unstintingly<br />
supported work on Florida lichens. Anisomeridium griffinii seems closest to Asian A. palvanum, differing in<br />
larger ascospores. In Florida it might be confused with A. ambiguum which differs in UV+ thallus and apical<br />
ostiole. Anisomeridium griffini was treated in 1990 as Ditremis sp. 11288 and Ditremis sp. Buck B724 (ostiole<br />
orientation misunderstood).<br />
Additional specimens. FLORIDA. Duval County: Jacksonville Beach, flood plain forest W and NW of 23rd<br />
& 24th St. N., between 9th & 11th Ave. N., on Liquidambar, 23 Dec 1975, Buck B724; Franklin County:<br />
Apalachicola National Forest, Hickory Landing Rec. Area, ca. 2 mi S of Sumatra, hardwoods along Owl Creek,<br />
23 May 1976, Harris 11288 (both NY).<br />
121
Anisomeridium nyssaegenum (Ellis & Everhart) R. C. Harris<br />
Evansia 2: 44. 1985. Zignoella nyssaegena Ellis & Everhart, Proc. Acad. Sci. Nat. Philadelphia<br />
1893. 137. 1893. Ditremis nyssaegena (Ellis & Everhart) R. C. Harris, Some Florida Lichens 3.<br />
1990. Type. NEW JERSEY. [Gloucester County:] Newfield, on Nyssa multiflora, 25 Nov 1892, Elis<br />
(NY, holotype).<br />
Apiospora polypori Ellis & Everhart, N. Amer. pyrenomyc. 33. 1892. Type. DELAWARE. Newark,<br />
on old Fomes applanatus, 7 Nov 1890, Commons 1713 (NY, holotype), syn. nov.<br />
Melanopsamma corticola Ellis & Everhart, Proc. Acad, Nat. Sci. Philadelphia 1893: 129. 1893.<br />
Type. NEW JERSEY. [Gloucester County:] Newfield, on outer bark of Quercus alba, 7 Jul 1890,<br />
Ellis (NY, holotype), syn. nov.<br />
Thelidium juistense Erichsen, Mitt. Inst. Allg. Bot. Hamburg 10: 403. 1939. Anisomeridium<br />
juistense (Erichsen) R. C. Harris in D. Hawksw. et al., Lichenologist 12: 106. 1980. Type.<br />
GERMANY. Insel Juist, on Sambucus nigra, 23 Jun 1937, Erichsen (US, isotype).<br />
Arthopyrenia willeyana R. C. Harris, Michigan Bot. 12: 16. 1973. Type. MICHIGAN. Iosco County:<br />
N of Tawas City, on Populus tremuloides, 18 Sep 1965, Harris 818 (MSC, holotype; B, BM,<br />
CAN, COLO, DUKE, H, LD, MIN, NY, S, TNS, UPS, US, V_zda, WIS, isotypes).<br />
Apiospora polypori is a year older than Zignoella nyssaegena and I believe that Margaret Barr plans to<br />
take up the epithet. However, since I have already changed the name of this taxon so many times it is almost<br />
embarassing, since this is such a weedy species there are probably yet older epithets hidden in equally<br />
improbable genera, and since there is some possibility that the species may be segregated out of<br />
Anisomeridium, I plan to wait until the dust settles before accepting this or any further nomenclatural change.<br />
Anisomeridium nyssaegenum does not fit neatly into either of the main groups in Anisomeridium (see generic<br />
discussion above). It also has beaked pycnidia which have been given the illegitimate generic name<br />
Sarcinulella Sutton & Alcorn. There are several other species with beaked pycnidia but it remains to be studied<br />
whether a group so defined can be given generic recognition.<br />
Anisomeridium phaeospermum R. C. Harris, sp. nov.<br />
Species unica ostiolo apicali et ascosporis fuscis, granulis ornatis, 16-20 × 8-9 µm.<br />
Type. FLORIDA. Collier County: Big Cypress National Preserve, Oasis Ranger Station along U.S. Hwy. 41,<br />
along Florida Trail N of hwy., 25°50'N, 81°03'W, Taxodium swamp, on Taxodium, 9 Dec 1992, Harris 30189<br />
(NY, holotype).<br />
Thallus white, endophloeodal, UV+ yellow (lichexanthone). Ascomata mostly immersed, tip exposed,<br />
subglobose, sometimes with clypeus laterally expanded, ca. 0.5 mm diam. Asci cylindrical, ca. 140-150 × 15-<br />
16 µm, with eight uniseriate spores. Ascospores brown, elliptical, slightly constricted at the ± median septum,<br />
granular ornamented, 16-20 × 8-9 µm; conidia not found.<br />
The other species of Anisomeridium with brown ascospores differ in having an eccentric ostiole.<br />
Anisomeridium phaeospermum is known only from the type collection.<br />
Anisomeridium quadrococcum R. C. Harris, sp. nov.<br />
Thallo dilute griseo UV-, ostiolo apicale, ascosporis 3-septatis, laevibus, 21-24 × 5.5-8 µm et<br />
microconidiis oblongis, 3-4 × 1.5 µm.<br />
Type. FLORIDA. Columbia County: Ichetucknee Springs State Park, Fig Island, 29°58'N, 82°45'W, moist<br />
hardwoods on small limestone island, on decorticate wood, 2 Dec 1992, Buck 22601 (NY, holotype; FLAS,<br />
isotype).<br />
Thallus pale gray, endophloeodal, UV-. Ascomata ca. ½-¾ immersed, globose, ca. 0.2-0.25 mm diam.;<br />
ostiole apical; melanized wall entire or lacking at base; clypeus not expanded laterally. Asci cylindrical to<br />
122
subclavate, 180-220 × 45-55 µm, with eight biseriate spores. Ascospores 4-celled, end cells smaller, fusiform,<br />
not constricted at septa, smooth, 21-26 × (5.5-)6.5-8 µm; microconidia oblong, 3-4 × 1.5 µm; macroconidia not<br />
found.<br />
Corticolous species of Anisomeridium with 4-celled ascospores are rare, A. quaternarium has smaller<br />
ascospores and A. nyssaegenum has mostly 2-celled, only tardily 4-celled ascospores.<br />
Additional specimen. GEORGIA. Charleton County: Okefenokee National Wildlife Refuge, Chesser Island,<br />
15 Nov 1989, Wetmore 64357 (MIN).<br />
Anisomeridium quaternarium (R. C. Harris) R. C. Harris, comb. nov.<br />
Ditremis quaternaria R. C. Harris, Some Florida Lichens 34. 1990. Type. LOUISIANA. Plaquemines<br />
Parish: Pointe a la Hache, on trunk of Populus, 5 Jun 1976, Sérusiaux 1708 (NY, holotype).<br />
Anisomeridium subprostans (Nyl.) R. C. Harris<br />
in Tucker & Harris, Bryologist 83: 4. 1980. Verrucaria subprostans Nyl., Expos. syn. pyrenocarp.<br />
56. 1858. Arthopyrenia subprostans (Nyl.) Müll. Arg., Flora 66: 317. 1883. Ditremis subprostans<br />
(Nyl.) R. C. Harris, Some Florida Lichens 34. 1990. Type. SOUTH CAROLINA. On Taxodium,<br />
Ravenel, comm. Tuckerman 75, (H-NYL Pm. 7378, holotype?).<br />
Anisomeridium tamarindi (Fée) R. C. Harris<br />
in Tucker & Harris, Bryologist 83: 4. 1980. Verrucaria tamarindi Fée, Essai crypt. écorc. suppl.<br />
85. 1837. Porina tamarindi (Fée) Müll. Arg., Mém. Soc. Phys. Genève 30: 24. 1888. Ditremis<br />
tamarindi (Fee) R. C. Harris, Some Florida Lichens 34. 1990. Type. "ad cortices Tamarindi<br />
officinalis L. Antillarum" (probably GUADELOUPE, see Fée, loc. cit, p. 165). (G, lectotype; G, H- Nyl<br />
748. isolectotypes).<br />
Verrucaria vaga Nyl., Expos. syn. pyrenocarp. 55. 1858. Porina vaga (Nyl.) Müll. Arg., Hedwigia<br />
31: 287. 1892. Type. COMORO ISLANDS. Mayotte, Boivin (H-Nyl 750, holoype).<br />
Porina elaeidis Vainio in Hiern et al., Cat. Afr. pl. 2(2): 458.1901. Type. ANGOLA. Ambriz: Ad<br />
Elaeidis guineensis truncum, Nov 1853, Welwitsch 360 (BM, isotype).<br />
Tomasellia (Syngenosorus) africana Zahlbr., Ann. Cryptog. Exot. 5: 203. 1932. Type. SOUTH<br />
AFRICA. Knysna, v. d. Byl 749 (LD, isotype).<br />
Anisomeridium terminatum (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria terminata Nyl., Bull. Soc. Linn. Normandie sér. 2, 7: 180. 1873. Arthopyrenia terminata<br />
(Nyl.) Müll. Arg., Flora 73: 345. 1890. Ditremis terminata (Nyl.) R. C. Harris in Egan, Bryologist<br />
90: 163. 1987. Type. ANDAMAN ISLANDS. 1867, Kurz 23 (H-Nyl 725, lectotype; BM, M,<br />
isolectotypes).<br />
Verrucaria interspersa Nyl., Bull. Soc. Linn. Normandie sér. 2, 7: 181. 1873; Porina interspersa<br />
(Nyl.) Zahlbr., Cat. Lich. Univ. 1: 387. 1922. Anisomeridium interspersum (Nyl.) Makhija et al.,<br />
J. Econ. Tax. Bot. 18: 544. 1994. Type. ANDAMAN ISLANDS. 1867, Kurz 3 (H-Nyl 726, lectotype).<br />
Arthopyrenia (sect. Polymeridium) glaucina Müll. Arg., Bot. Jahrb. Syst. 6: 406. 1885. Type.<br />
CUBA. Wright, Lich. Cub. ser. 2: 647 (G, holotype; US, isotype) .<br />
Tomasellia queenslandica Müll. Arg., Australas. Ass. Adv. Sci. Rep. 6: 460. 1895. Type.<br />
AUSTRALIA. Queensland, 1887, Knight 46 (G, holotype) .<br />
Arthopyrenia anacardii Vainio, Ann. Acad. Sci. Fenn. ser. A, 6(7): 209. 1915. Type. GUADELOUPE.<br />
Basse Terre, sur un Anacardium occidentale, Duss 498 (TUR-Vainio 32131, holotype; NY,<br />
isotype).<br />
Arthopyrenia insularis Vainio, Ann. Acad. Sci. Fenn. ser. A, 6(7): 208. 1915. Type. WEST INDIES.<br />
Insulis Danicis, 1905-6, Boergesen (TUR-Vainio 32116, holotype).<br />
Arthopyrenia subinsularis Vainio, Ann. Acad. Sci. Fenn. ser. A, 6(7): 208. 1915. Type. WEST<br />
INDIES. St. Croix, Mount Eagle, 20 Jan 1906, Boergesen (TUR-Vainio 32117, holotype).<br />
Verrucaria terminata and V. interspersa were published simultaneously. I did not formally<br />
123
select between them in "Some Florida Lichens" but the choice was implied in the citation of<br />
synonymy. The choice of the name Verrucaria terminata is formally endorsed here.<br />
Anisomeridium tuckerae R. C. Harris<br />
in Tucker & Harris, Bryologist 83: 4. 1980. Ditremis tuckerae (R. C. Harris) R. C. Harris, Some<br />
Florida Lichens 35. 1990.Type. LOUISIANA. East Baton Rouge Parish: hardwood forest at Burden<br />
Plantation, Essen Lane, Baton Rouge, Tucker 10003 (LSU, holotype) .<br />
MONOBLASTIA Riddle<br />
1. Ostiole strongly eccentric to lateral ........................................................................................................... 2<br />
2. Ascospores 30 µm long or less, 8/ascus, very strongly ornamented to<br />
rather strongly ornamented ................................................................................................................... 3<br />
3. Ascospores globose to long-ellipsoid................................................................................................ 4<br />
4. Ascospores ellipsoid to long-ellipsoid, 18-24 × 8-10 µm,<br />
ornamentation moderately granular/papillate; Puerto Rico,<br />
Mississippi ........................................................................................[M. borinquensis R. C. Harris]<br />
4. Ascospores globose to oblong or rarely ellipsoidal (in the same<br />
ascoma), 11-15 µm in diam. or 14-18 × 7-10 µm; southern Coastal<br />
Plain, French Guiana, French Antilles................................................................. M. rappii Zahlbr.<br />
3. Ascospores cylindrical to fusiform, 24-30 × 8-9 µm; Suwanee<br />
County ............................................................................................................. M. buckii R. C. Harris<br />
2. Ascospores over 30 µm long, weakly ornamented ............................................................................... 5<br />
5. Ascospores initially 8/ascus, often 4-6 aborting,<br />
32-36(-50) × 15-17(-20) µm; Cuba .....................................................................[M. palmicola Riddle]<br />
5. Ascospores initially 4/ascus, occasionally 1-2 aborting,<br />
45-50(-67) × 17-21 µm; Dominican Republic .......................................[M. quisqueyana R. C. Harris]<br />
1. Ostiole apical to slightly eccentric ............................................................................................................. 6<br />
6. Growing on bark .................................................................................................................................... 7<br />
7. Ascomata pallid, not melanized; ascospores ornamented<br />
with short spines, 9-13 × 7-9 µm; French Guiana, Papua<br />
New Guinea....................................................................................................... [M. pellucida Aptroot]<br />
7. Ascomata dark brown-black, melanized; ascospores<br />
ornamented with low, flat warts, 32-45 × 17-22 µm;<br />
Collier County..............................................................................................M. cypressi R. C. Harris<br />
6. Growing on limestone; ascospores strongly warted, 19-25(-28) ×<br />
8-11 µm; Cuba, Haiti, Puerto Rico............................................ [M. echinulospora (Riddle) R. C. Harris]<br />
124
Monoblastia borinquensis R. C. Harris, sp. nov.<br />
NOTES<br />
Ascomatis ostiolis valde excentricis vel lateralibus et ascosporis octonis, ellipsoideis, verruculosis, 18-24<br />
× 8-10 µm.<br />
Type. PUERTO RICO. Distr. Arecibo, ca. i km N of ciales on Hwy. 149, 220 m, humid forest over limestone in<br />
haystack hills, 11 Jun 1988, Harris 22615 (NY, holotype).<br />
Thallus olivaceous or grayish, not corticate, UV-. Ascomata flask-shaped with subspherical body and<br />
projecting ostiolar neck, 0.5-0.7 mm in diam. (without neck), ca. ¾ immersed; ostiole strongly eccentric to<br />
lateral. Asci cylindrical with 8 uniseriate ascospores. Ascospores ellipsoidal, with verruculose to papillate<br />
ornamentation, 18-24 × 8-10(-11) µm; microconidia oblong, 3-4 1.5 µm.<br />
Monoblastia borinquensis is close to M. buckii but differs in shorter, broader ascospores. With additional<br />
material this difference may prove meaningless. The ascospore ornamentation of the Mississippi collection is<br />
weaker than in the type but I can find no other differences.<br />
Additional specimen. MISSISSIPPI. Sharkey County: Delta National Forest, along road #707, 9 mi NE of<br />
Valley Park, Jul 1978, Johnson s.n. (NY).<br />
Monoblastia cypressi R. C. Harris, sp. nov.<br />
Ascomatis ostiolis apicalibus et ascosporis octonis, oblongi-ellipsoideis, verrucis latis, 32-47 × 17-22 µm.<br />
Type. FLORIDA. Collier County: Big Cypress National Preserve, Oasis Ranger Station along U.S. Hwy. 41,<br />
along Florida Trail N of highway, Taxodium swamp, on Taxodium, 9 Dec 1992, Harris 30194 (NY, holotype).<br />
Thallus whitish, thin (bark showing through), not corticate, UV-. Ascomata subspherical with apical<br />
ostiolar papilla, 0.7-1.0 mm diam., ¾ to entirely immersed; ostiole apical. Asci cylindrical with eight uniseriate<br />
ascospores. Ascospores oblong-ellipsoidal with strong oranamentation of low, broad warts, 31-47 × 17-22<br />
µm; microconidia and macroconidia not found.<br />
Monoblastia cypressi has the largest ascospores of the eight-spored species. It is known only from the<br />
type.<br />
Monoblastia quisqueyana R. C. Harris, sp. nov.<br />
Ascomatis ostiolis valde excentricis et ascosporis quaternis, oblongi-ellipsoideis, leniter papillatis, 45-67 ×<br />
17-22 µm.<br />
Type. DOMINICAN REPUBLIC. Prov. La Altagracia: 1.2 km S of El Macao along beach road, 18°46'N, 68°31'W,<br />
sea level, red mangrove swamp, on Rhizophora mangle, 22 Jan 1991, Harris 26612 (NY, holotype).<br />
Thallus pale gray, matte, not corticate, UV-. Ascomata ± spherical, occasionally with slightly projecting<br />
ostiolar neck, 0.7-1.0 mm diam., ca. ½-¾ immersed; ostiole strongly eccentric. Asci cylindrical with 4<br />
uniseriate ascospores. Ascospores oblong-ellipsoid with weak papillate ornamentation, 45-67 × 17-22 µm;<br />
microconidia and macroconidia not found.<br />
Other species of Monoblastia may have fewer than eight spores per ascus due to abortion (young asci<br />
with eight spores always present) but as far as I can tell M. quisqueyana is the only one which has only four<br />
initially. It is known only from the type.<br />
125
MUSAESPORA Aptroot & Sipman<br />
Aptroot and Sipman placed Musaespora in the Aspidotheliaceae (= Thelenellaceae) but the asci do not<br />
agree with those of that family and Musaespora seems better assigned to the Monoblastiaceae. In fact there<br />
are several taxa with corticate thalli that I am carrying in Anisomeridium which, when that genus is broken up,<br />
may be appropriately placed in Musaespora. I have seen material of an additional, probably undescribed,<br />
corticolous species with campylidia from Costa Rica. I need to go back and search for campylidia on the thalli<br />
of some other species which have a similar ascospore and ascus type, although I am not completely<br />
convinced a campylidiate anamorph alone should determine generic status.<br />
Musaespora epiphylla (R. Sant.) R. C. Harris, comb. nov.<br />
Astrothelium epiphyllum R. Sant., Symb. Bot. Upsal. 12(1): 131. 1952. Type. SUMATRA. Oostkust<br />
v. Sumatra, near Besitang, (near boundary of Atjeh), alt. c. 30 m, 1926, Palm 17b (UPS,<br />
holotype).<br />
?Musaespora multilocularis Aptroot & Sipman, Lichenologist 25: 127. 1993. Type. PAPUA NEW<br />
GUINEA (not seen).<br />
Musaespora gigas (Zahlbr.) R. C. Harris, comb. nov.<br />
Mycoporellum gigas Zahlbr., Ann. Crypt. Exot. 1: 117. 1928. Type. INDONESIA. Java: Pati,<br />
Seubert (US, isotype).<br />
Musaespora corticola Aptroot & Sipman, Lichenologist 25: 127. 1993. Type. INDONESIA. West<br />
Java (not seen).<br />
APPENDIX<br />
ANISOMERIDIUM, A GENERAL KEY TO SPECIES<br />
WARNING. The following key is only a very rough approximation. Anisomeridium s. lat. is a very speciose<br />
taxon and perhaps only 50-75% of the species may be included here. I am well aware illustrations are a vital<br />
necessity but time and money simply have not been available. Much more data must be gathered on conidia<br />
and the user is informed that in many cases conidia are known for a taxon from only a single instance and<br />
may not belong to the species to which they are assigned. Repeated co-occurrences can provide some<br />
confidence but culture from ascospores is the only definitive proof. The utility as characters of lichexanthone<br />
and ascospore ornamentation also needs to be validated.<br />
1. Growing on bark or wood .......................................................................................................................... 2<br />
2. Ascospores 2-celled; ostiole apical or eccentric ................................................................................... 3<br />
3. Thallus endophloeodal, not corticate, usually dull, usually whitish or grayish,<br />
less commonly greenish; ascomata immersed or not...................................................................... 4<br />
4. Thallus UV-................................................................................................................................... 5<br />
5. Ostiole apical............................................................................................................................ 6<br />
6. Asci with 8 spores ............................................................................................................... 7<br />
7. Ascospores 35 µm or less in length ............................................................................... 8<br />
8. Ascospores smooth [a complex of intergrading species<br />
containing the type of Anisomeridium]........................................................................ 9<br />
126
9. Ascospores biseriate or irregularly arranged in ascus ......................................... 10<br />
10. Ascospores less than 8 µm broad.................................................................. 11<br />
11. Macropycnidia not beaked ....................................................................... 12<br />
12. Ascospores relatively long, mostly over<br />
15 µm long........................................................................................... 13<br />
13. Ascospores remaining 2-celled; microconidia<br />
mostly globose ............................................................................. 14<br />
14. Asci slender, cylindrical or narrowly<br />
clavate ..................................................................................... 15<br />
15. Microconidia elliptical, 2.5-1.5 × 2 µm;<br />
macroconidia ± oblong, 5-7 × 2.5-3<br />
µm; ascospores narrowly ovoid, with<br />
weakly submedian septum; 12-18(-21)<br />
× 4.5-6.5 µm; neotropical........................................................<br />
........................................A. subprostans (Nyl.) R. C. Harris<br />
15. Microconidia globose, ca. 2 µm diam.;<br />
macroconidia elliptical, 4-7 × 2.5-3.5<br />
µm; ascospores 15-22(-24) × 4.5-<br />
5.5(-6) µm, fusiform, often slightly<br />
bent; pantropical/subtropical ................................................<br />
............................................ A. tamarindi (Fée) R. C. Harris<br />
14. Asci broader, ovoid or obovoid, broadly<br />
cylindrical or narrowly ovoid or<br />
narrowly obovoid .................................................................... 16<br />
16. Thallus poorly developed,<br />
endophloeodal................................................................... 17<br />
17. Ascospores over 5 µm in width ................................... 18<br />
18. Ascospore wall usually<br />
unevenly thickened;<br />
ascospores ovoid, with<br />
markedly submedian<br />
septum, 14-18 × 6-8 µm;<br />
microconida globose, ca. 2 µm<br />
diam., macroconidia oblong or<br />
ovoid, 5-7 × 2.5-3 µm;<br />
pantropical/subtropical ..............................................<br />
A. anisolobum (Müll. Arg.) Aptroot<br />
18. Ascospore wall relatively thin,<br />
even; ascospores 18-22 ×<br />
5.5-7 µm, ± fusiform, often<br />
slightly bent, with slightly<br />
submedian septum;<br />
127
128<br />
microconidia globose or<br />
elliptical, ca. 2 µm diam. or<br />
2-3 × 1.5-2 µm; neotropical .......................................<br />
A. excaecariae (Müll. Arg.) R. C. Harris<br />
17. Ascospores 12-17(-19) × 3.5-4.7<br />
µm; microconidia narrowly ovoid,<br />
4-5 × 1-1.3 µm; macroconidia<br />
oblong, 9-10.5 × 2-2.5 µm;<br />
Scotland & Wales............................................................<br />
A. viridescens (Coppins) R. C. Harris<br />
16. Thallus well developed, epiphloeodal;<br />
ascospores 12-18 × 5-7.5 µm;<br />
macroconidia globose, ca. 2 µm diam.;<br />
microconidia globose, 2.5-4 µm diam.;<br />
Illinois, Massachusetts, Ohio.................................................<br />
A. leucochlorum (Müll. Arg.) R. C. Harris<br />
13. Ascospores 14-20 × 4.5-6 µm, ovoid, with<br />
submedian septum, often becoming 4-celled;<br />
micropycnidia beaked ("Sarcinulella");<br />
microconidia oblong or narrowly elliptical,<br />
2-3 × 1-1.5 µm; temperate North America<br />
and Europe........................................................................................<br />
................................ A. nyssaegenum (Ellis & Everhart) R. C. Harris<br />
12. Ascospores relatively short, less than 15 µm ..................................... 19<br />
19. Septum weakly submedian to median ......................................... 20<br />
20. Ascospores 10-18 × 4.5-7 µm, broadly<br />
elliptical to ovoid, rarely narrowly ovoid, with<br />
septum slightly submedian or ± median;<br />
microconidia oblong or elliptical, 2.5-3 ×<br />
1.5-2 µm or globose, ca. 2 µm diam.;<br />
macroconidia elliptical or oblong, 5-7 ×<br />
3-4 µm; cosmopolitan..................................................................<br />
................................................. A. biforme (Borrer) R. C. Harris<br />
20. Ascospores 9-13(-15) × 4-5 µm, ovoid,<br />
with septum median to slightly submedian;<br />
ascomata small 0.2-0.3(-0.4) mm diam.;<br />
microconidia oblong or elliptical, 3-4 × 1.5-2<br />
µm; Florida ................................ A. albisedumi (Nyl.) R. C. Harris<br />
19. Ascospore septum markedly submedian;<br />
ascospores ovoid, 12-15 × 6-8 µm;<br />
microconidia globose, ca. 2 µm; New<br />
Zealand ................................... A. subatomaria (Knight) R. C. Harris<br />
11. Macropycnida with a long, hair-like beak, extruding
conidia in a single row; macroconida oblong,<br />
± constricted in middle, 8-10 × 4-5 µm;<br />
ascospores ovoid, with submedian septum,<br />
12-17 × 5-6 µm; Hong Kong................................A. throwerae R. C. Harris<br />
10. Ascospores mostly more than 8.5 µm broad................................................. 21<br />
21. Ascospores 8-12 µm broad ..................................................................... 22<br />
22. Ascospores fusiform; septum slightly submedian<br />
or median ........................................................................................... 23<br />
23. Ascospores 22-28 × (7-)8-9.5 µm; Philippine<br />
Islands ................................. A. sphaerocarpum (Vainio) R. C. Harris<br />
23. Ascospores 26-32(-34) × 8.5-11.5 µm; Andaman<br />
Islands ..............................................A. subnexum (Nyl.) R. C. Harris<br />
22. Ascospores ovoid to broadly ovoid, with<br />
septum strongly submedian, 18-23(-28) ×<br />
9-14 µm; microconidia globose, ca. 2 µm diam.;<br />
neotropical................................ A. leptospermum (Zahlbr.) R. C. Harris<br />
21. Ascospores 23-32 × 12-18 µm broad, ovoid to broadly<br />
ovoid, with strongly submedian septum; pantropical...................................<br />
...........................................................A. subnectendum (Nyl.) R. C. Harris<br />
9. Ascospores uniseriate.......................................................................................... 24<br />
24. Ascospores 10-18 × 4.5-7 µm, elliptical or ovoid, with<br />
septum slightly submedian or ± median; microconidia oblong<br />
or elliptical, 2.5-3 × 1.5-2 µm or globose, ca. 2 µm diam.;<br />
macroconidia elliptical or oblong, 5-7 × 3-4 µm;<br />
cosmopolitan ..................................................... A. biforme (Borrer) R. C. Harris<br />
24. Ascospores 17-22 × 9-9.5 µm, ovoid, with submedian<br />
septum; macroconidia oblong or elliptical, 5-6 × 2.5-3 µm;<br />
Philippine Islands ............................................A. consimile (Vainio) R. C. Harris<br />
8. Ascospores granular ornamented, ovoid to ± pyriform, with<br />
septum very markedly submedian, biseriate/irregular in ascus,<br />
26-33 × 16-18 µm; microconidia globose, ca. 2 µm diam.;<br />
Brazil...................................................................... A. adnexa (Müll. Arg.) R. C. Harris<br />
7. Ascospores 37-52 × 17-24 µm, broadly elliptical, with ± median septum,<br />
smooth?; New Zealand .......................................... A. magnospora (Knight) D. Hawksw.<br />
6. Asci with 2-4 spores .......................................................................................................... 25<br />
25. Ascospores 4/ascus, fusiform, thick-walled, with ± median<br />
septum, strongly ornamented, 55-85 × 20-28 µm;<br />
microconidia elliptical or oblong, 22-3 × 1.5 µm; Cuba,<br />
Puerto Rico, St. Eustatius ...................................... A. excellens (Müll. Arg.) R. C. Harris<br />
129
25. Ascospores 2-4/ascus, broadly elliptical, relatively thin-walled,<br />
with ± median, smooth? septum, 62-82 × (22-)27-33 µm;<br />
Jamaica?.................................................................. A. planiusculum (Nyl.) R. C. Harris<br />
5. Ostiole eccentric..................................................................................................................... 26<br />
26. Ascospores colorless, ornamented or smooth................................................................. 27<br />
27. Asci with 8 spores ....................................................................................................... 28<br />
28. Ascospores smooth................................................................................................ 29<br />
29. Ascospores 7-11 µm in width, fusiform; septum ± median ............................... 30<br />
30. Ascospores 30-35(-40) × 7-11 µm; microconidia elliptical,<br />
3 × 1.5-2 µm; macroconidia narrowly ovoid, 11-20 ×<br />
5.5-7 µm; Andaman Islands, India ...................................................................<br />
.....................................A. complanatum (Makhija & Patwardhan) R. C. Harris<br />
30. Ascospores 17-29 × 7-8 µm ......................................................................... 31<br />
31. Ascospores 23-29 × 7.5-8 µm; microconidia<br />
elliptical, 2.5 × 1.5 µm; New Caledonia ......................................................<br />
.......................................................A. polycarpum (Müll. Arg.) R. C. Harris<br />
31. Ascospores 17-23 × 7-9 µm; pycnidia not found;<br />
Florida ................................................. Anisomeridium sp. Harris 29392-A<br />
29. Ascospores 9-18 µm in width, elliptical or broadly ovoid................................... 32<br />
32. Ascospores 22-30(-33) × 9-14 µm, with slightly submedian<br />
septum; macroconidia broadly ovoid or elliptical, 4-6 ×<br />
3-4 µm; Philippine Islands, New Caledonia, Andaman<br />
Islands ........................................................A. palavanum (Vainio) R. C. Harris<br />
32. Ascospores 32-40(-45) × 13.5-18 µm; microconidia<br />
subglobose, ca. 2-2.5 µm diam.; macroconidia elliptical or<br />
oblong, 5-7 × 2.5-4 µm; Florida .......................................A. griffinii R. C. Harris<br />
28. Ascospores granular ornamented .......................................................................... 33<br />
33. Ascospores fusiform to narrowly ovoid ............................................................. 34<br />
34. Ascospores 19-42 × 5.5-12 µm, fusiform, with septum<br />
± submedian; pantropcical/subtropical......................................................... 35<br />
35. Ascospores 26-35(-42) × 7.5-10(-12) µm...................................................<br />
........................................................ A. americanum (Massal.) R. C. Harris<br />
35. Ascospores 19-27(-30) × 5.5-7(-8) µm;<br />
microconidia oblong or elliptical, 2.5-3 ×<br />
1.5-2 µm ................................................. A. terminatum (Nyl.) R. C. Harris<br />
34. Ascospores 18-21 × 7-9 µm, narrowly ovoid, with<br />
septum submedian; New Zealand.....................................................................<br />
.................................................................. A. subbiforme (Knight) R. C. Harris<br />
130
33. Ascospores broadly elliptical, with moderately submedian<br />
septum, 17-21 × 8-10 µm; Hawaiian Islands..........................................................<br />
................................................................................A. rockii (Zahlbr.) R. C. Harris<br />
27. Asci with 2-4 spores .................................................................................................... 36<br />
36. Asci with (2-)4 spores; ascospores fusiform, constricted at<br />
± median septum..................................................................................................... 37<br />
37. Ascospores ornamented, 55-85 × 20-28 µm; microconidia<br />
elliptical or oblong, 2-3 × 1.5 µm; Cuba, Puerto Rico,<br />
St. Eustatius ................................................ A. excellens (Müll. Arg.) R. C. Harris<br />
37. Ascospores smooth, 52-68 × 14-20 µm; microconida<br />
± globose, ca. 2 µm diam.; Andaman Islands .......................................................<br />
...........................................A. tarmugliense (Makhija & Patwardhan) R. C. Harris<br />
36. Asci with 2 spores; ascospores broadly elliptical or oblong,<br />
constricted at ± median septum, granular ornamented; 52-74 ×<br />
21-27 µm; Trinidad, Cuba, Dominican Republic .........................................................<br />
..........................................................................A. megalosprum (Vainio) R. C. Harris<br />
26. Ascospores brown, strongly granular ornamented, oblong-ovoid, with ±<br />
median septum, 27-36 × 9-11 µm; ascomata emergent, with aspect<br />
of A. inspersum; microconidia elliptical, 3 × 1.5 µm;<br />
Cuba...................................................................................Anisomeridium sp. Harris 14615<br />
4. Thallus UV+ yellow (lichexanthone) ........................................................................................... 38<br />
38. Ostiole apical; ascospore septum submedian or not .......................................................... 39<br />
39. Ascospores colorless, not ornamented.......................................................................... 40<br />
40. Ascospores over 8 µm in width, mostly uniseriate in ascus ...................................... 41<br />
41. Ascospores ± fusiform with one end often more rounded, with<br />
septum median or submedian, smooth, 24-33 × 8-12 µm;<br />
microconidia elliptical, 3-5 × 1.5-2 µm; macroconidia broadly<br />
ovoid to ovoid, thick-walled,15-22 × 9-16 µm; French Guiana,<br />
Jamaica, Florida, Seychelles.............................. A. ambiguum (Zahlbr.) R. C. Harris<br />
41. Ascospores elliptical to broadly elliptical, with septum ± median,<br />
25-30 × 9-13 µm; microconidia elliptical, 3-4 × 1.5-2 µm;<br />
New Caledonia, Queensland......................................A. consobrinum (Nyl.) Aptroot<br />
40. Ascospores less than 7 µm in width, biseriate in ascus ............................................ 42<br />
42. Macroconidia cylindrical, 7.5-12 × 2-2.5 µm; microconidia<br />
elliptical to subglobose, 2.5-3 × 1.5-2 or ca. 2 µm diam.;<br />
ascospores fusiform, with slightly submedian septum 15-20 ×<br />
5-6.5 µm; Florida ...........................................A. tuckerae (R. C. Harris) R. C. Harris<br />
42. Macroconidia oblong, 6-8 × 2-2.5 µm; microconidia elliptical<br />
to subglobose, 2.5-3 × 1.5-2 µm; ascospores narrowly fusiform,<br />
131
with slightly submedian septum, 14-19 × 4-5.5 µm; São<br />
Tomé .....................................................................A. albidoatrum (Nyl.) R. C. Harris<br />
39. Ascospores brown, ornamented, elliptical to broadly<br />
elliptical, with median septum, 17-20 × 8-9 µm;<br />
Florida ...............................................................................A. phaeospermum R. C. Harris<br />
38. Ostiole eccentric; ascospores broadly fusiform, with submedian<br />
septum, smooth .................................................................................................................. 43<br />
43. Ascospores colorless .................................................................................................... 44<br />
44. Ascospores 13 µm or more in width......................................................................... 45<br />
45. Ostiole not ringed with orange pigment.............................................................. 46<br />
46. Ascospores (37-)42-53 × 13-17 µm, with markedly<br />
submedian septum; Dominican Republic, Puerto<br />
Rico ........................................................... Anisomeridium sp. Harris 20160-A<br />
46. Ascospores 30-40 × 13-16.5(-18) µm; microconidia<br />
oblong, 3-4 × 1.5-2 µm; India ...........................................................................<br />
..................................... A. flavopallidum (Makhija & Patwardhan) R. C. Harris<br />
45. Ostiole ringed with orange pigment, KOH+ purple;<br />
ascospores fusiform, with septum submedian, 37-52 ×<br />
12-20 µm; microconidia oblong, 3-4 × 1.2-1.5 µm;<br />
macroconidia broadly elliptical to ovoid, 30-45 ×<br />
15-20 µm; Florida ..................................................A. aureopunctatum R. C. Harris<br />
44. Ascospores 13 µm or less in width........................................................................... 47<br />
47. Ascomata exposed, with a broad, melanized clypeus;<br />
ascospores 20-27 × 8.5-11 µm; Cuba.....................................................................<br />
....................................................................A. angulosum (Müll. Arg.) R. C. Harris<br />
47. Ascomata mostly hidden, immersed, with poorly developed<br />
clypeus; ascospores 23-34 × 9-13 µm; Asia, Pacific...............................................<br />
...............................................................................A. ubiana (Vainio) R. C. Harris<br />
43. Ascospores brown, broadly elliptical, with ± median septum<br />
9-11 × 5-6.5 µm; microconidia oblong, 2-3.5 × 1-1.5 µm;<br />
South Africa...................................................................A. uniseriale (Zahlbr.) R. C. Harris<br />
3. Thallus mostly epiphloeodal, corticate, usually ± glossy, usually greenish or grayish;<br />
ascomata immersed (except A. truncatum) ................................................................................... 48<br />
48. Thallus (cortex or medulla) UV+ yellow (lichexanthone); ascospores mostly<br />
ovoid to broadly ovoid, constricted at markedly submedian septum, smooth.......................... 49<br />
49. Ascospores 45 × 18 µm or larger ........................................................................................ 50<br />
50. Medulla UV+ yellow; cortex UV-; ascospores (48-)55-65(-70)<br />
× 25-30(35) µm; microconidia elliptical, 3 × 1.5-2 µm;<br />
French Guiana, Trinidad, St. Vincent ................................A. infernale (Mont.) R. C. Harris<br />
132
50. Medulla UV-; cortex UV+ yellow; ascospores 45-57 × 18-26 µm;<br />
Puerto Rico..................................................................................... A. elegans R. C. Harris<br />
49. Ascospores 45 × 18 µm or smaller...................................................................................... 51<br />
51. Ascospores with septum median to slightly submedian................................................ 52<br />
52. Ascospores with rounded ends, broadly elliptical,<br />
(10-)12-15 × 7-9 µm; microconidia broadly elliptical,<br />
3 × 1.5-2 µm; Florida and Georgia ................................... A. biformoides R. C. Harris<br />
52. Ascospores with ends somewhat pointed, 31-41 × 13-17 µm;<br />
microconidia oblong or ovoid, 3-4 × 1.5-2 µm; India ...................................................<br />
......................................................... A. indicum (Makhija & Patwardhan) R. C. Harris<br />
51. Ascospores with septum markedly submedian, ends rounded,<br />
34-42 × 16-18 µm; ?microconidia globose, 3-4 µm diam.;<br />
Dominican Republic .......................................................Anisomeridium sp. Harris 16267<br />
48. Thallus UV-............................................................................................................................... 53<br />
53. Ascomata and pycnidia solitary.......................................................................................... 54<br />
54. Ostiole eccentric; ascospores smooth .......................................................................... 55<br />
55. Asci with eight spores.............................................................................................. 56<br />
56. Spores in a single row in the ascus.................................................................... 57<br />
57. Ascospores fusiform with pointed ends; septum median............................. 58<br />
58. Ascospores over 50 µm long.................................................................. 59<br />
59. Ascospores 84-117 × 22-27 µm; microconidia<br />
oblong, 3-5 × 1.5-2 µm; French Guiana and<br />
Trinidad .......................................................A. nemorosum R. C. Harris<br />
59. Ascospores 52-83 × 16-21 µm; microconidia<br />
oblong, 2.5-3.5 × 1.5 µm; India...............................................................<br />
................................A. immersum (Makhija & Patwardhan) R. C. Harris<br />
58. Ascospores 30-37 × 9-11 µm; microconidia 3-4 ×<br />
1.5 µm; Cuba................................ A. glaucescens (Müll. Arg.) R. C. Harris<br />
57. Ascospores with rounded ends, broadly elliptical, with<br />
median septum, 31-46 × 14-21 µm; macroconidia<br />
broadly elliptical, 5.5-7.5 × 3-4 µm; Trinidad and<br />
Dominica ......................................................... Anisomeridium sp. Hale 38240<br />
56. Spores in two rows or irregularly arrranged in the ascus .................................. 60<br />
60. Ascospores ovoid, ends rounded, with septum markedly<br />
submedian, 40-45 × 15-19 µm; French Guiana ................................................<br />
......................................................................... Anisomeridium sp. Buck 18364<br />
60. Ascospores broadly fusiform with ± acute ends, with<br />
median septum, 57-76 × 22-27 µm; New<br />
133
Caledonia .......................................................A. holopolium (Nyl.) R. C. Harris<br />
55. Asci with 1-4 spores ................................................................................................ 61<br />
61. Asci with 1 spore; ascospores fusiform, constricted at ± median<br />
septum, ends ± rounded, 200-225 × 52-62 µm; microconidia<br />
oblong, 3-4 × 1.5-2 µm; India ..................................................................................<br />
............................................. A. monosporum (Makhija & Patwarhan) R. C. Harris<br />
61. Asci with 4 spores; ascospores fusiform, constricted at ± median<br />
septum, ends pointed, 58-70 × 12-17 µm; Andaman Islands and<br />
Australia ...................................A. verrucosa (Makhija & Patwardhan) R. C. Harris<br />
54. Ostiole apical; ascospores smooth or ornamented....................................................... 62<br />
62. Ascomata emergent, flat-topped; thallus very well developed,<br />
greenish tan, Pyrenula-like; ascospores narrowly ovoid, with<br />
septum submedian, strongly granular oranamented, uniseriate in<br />
ascus; Brazil ..........................................................A. truncatum (Müll. Arg.) . C. Harris<br />
62. Ascomata completely immersed; thallus olivaceous, almost<br />
Clathroporina-like; ascospores broadly ovoid, ± constricted<br />
at the markedly submedian septum, smooth, subbiseriate in<br />
ascus, 42-52 × 20-24 µm; Sri Lanka ..............................................................................<br />
...............................................................................A. nidulans (Müll. Arg.) R. C. Harris<br />
53. Ascomata and pycnidia several under a single common clypeus;<br />
thallus thick, to 75 µm, with thick cortex, to 25 µm; ascospores<br />
fusiform, slightly constricted at weakly submedian septum, smooth,<br />
15-23 × 4.5-5.5 µm; microconidia elliptical, 3 × 1.5-2 µm;<br />
macroconidia elliptical, 5-6 × 2-2.5 µm; Dominican<br />
Republic ..................................................................................... A. stromaticum R. C. Harris<br />
2. Ascospores 4-celled; ostiole apical ..................................................................................................... 63<br />
63. Ascospores soon 4-celled ............................................................................................................. 64<br />
64. Ascospores 21-24 × 6-8 µml; microconidia elliptical, 3-4 × 1.5-2 µm;<br />
Florida and Georgia....................................................................... A. quadrococcum R. C. Harris<br />
64. Ascospores16-20 × 3.5-4.5 µm; microconidia ± globose,<br />
ca. 2 µm diam.; macroconidia oblong, 5-6 × 2-3 µm;<br />
Louisiana ................................................................... A. quaternarium (R. C. Harris) R. C. Harris<br />
63. Ascospores tardily 4-celled, ovoid, 14-20 × 4.5-6 µm; eastern<br />
North America .............................................................A. nyssaegenum (Ellis & Everh.) R. C. Harris<br />
1. Growing on rock, soil, liverworts or leaves .............................................................................................. 65<br />
65. Growing on rock or compacted soil (in part modified from McCarthy (1993).................................... 66<br />
66. Ascospores 2-celled..................................................................................................................... 67<br />
67. Growing on non-calcareous rock ............................................................................................ 68<br />
68. North Temperate taxa ........................................................................................................ 69<br />
134
69. Growing in aquatic or semiaquatic habitats................................................................... 70<br />
70. Thallus well developed, often with black hypothallus; ascospores<br />
ovoid, with septum submedian, smooth, (11-)13-18 × 5.5-7 µm;<br />
microconidia oblong, 4-5 × 1.5-2 µm; eastern North America,<br />
Europe...............................................................A. carinthiacum (Steiner) R. C. Harris<br />
70. Thallus scant or not evident; ascospores tardily 4-celled<br />
(see below)................................................................. A. distans (Willey) R. C. Harris<br />
69. Growing on dry rock; ascospores ovoid, with submedian septum,<br />
smooth, 13-15 × 6-6.5 µm; Virgin Islands .........................................................................<br />
.....................................................................................A. antillarum (Vainio) R. C. Harris<br />
68. Australasian/Pacific taxa .................................................................................................... 71<br />
71. Thallus 60-140 µm thick, pale gray; perithecia 0.17-0.32 mm diam.;<br />
clypeus 30-45 µm thick; New Zealand ..............................................................................<br />
..............................................................................A. laevigatum (McCarthy) R. C. Harris<br />
71. Thallus 30-60 µm thick, pale gray-green to pale brown gray;<br />
perithecia 0.24-0.44 mm diam.; clypeus 45-65 µm thick;<br />
Hawaiian..................................................................A. pacificum (McCarthy) R. C. Harris<br />
67. Growing on limestone.............................................................................................................. 72<br />
72. Ascospores 20-27 × 6-8.5 µm, narrowly ovoid to fusiform,<br />
with submedian septum, smooth; ascomata solitary; microconidia<br />
elliptical 3-5 × 1-1.5 µm; macroconidia broadly elliptical,5-6 × 4-4.5 µm;<br />
Florida and Puerto Rico..................................................... A. finkii (R. C. Harris) R. C. Harris<br />
72. Ascospores 11-13 × 3.5-4.5 µm, fusiform, with septum ± median,<br />
smooth; ascomata initially solitary, later forming in clusters; microconidia<br />
oblong, 3 × 1.5 µm; Puerto Rico..........................................................A. gregale R. C. Harris<br />
66. Ascospores 4-celled..................................................................................................................... 73<br />
73. Ascospores tardily 4-celled, ovoid, 16-22 × 6.5-8 µm; microconidia<br />
narrowly elliptical, 4-5 × 1.5-2 µm; macroconidia ovoid or narrowly<br />
elliptical, 5-7 × 2.5-3 µm; eastern North America. ....................... A. distans (Willey) R. C. Harris<br />
73. Ascospores soon 4-celled, broadly fusiform, 21-36 × 7.5-13 µm;<br />
microconidia bacilliform 2.5-4 × 0.7 µm; Australia........................................................................<br />
..........................................................................................A. australiense (McCarthy) R. C. Harris<br />
65. Growing on bryophytes or leaves ...................................................................................................... 74<br />
74. Growing on leaves........................................................................................................................ 75<br />
75. Ostiole apical........................................................................................................................... 76<br />
76. Ascomata dark; ascospores elliptical, with ± submedian septum,<br />
smooth, 13-14.5 × 4.5-5.5 µm; pycnidia with apical protruberances<br />
in a ± stellate arrangement; macroconidia? 4.5-5 × 3.5-4.5 µm;<br />
pantropical...................................................................................A. foliicola R. Sant. & Tibell<br />
76. Ascomata pale; ascospores ovoid, septum slightly supramedian,<br />
14-20 × 5.5-6.5 µm; Tanzania .........................................................Porinula tanzanica V_zda<br />
135
(I have not seen material. Pale ascomata are known in Monoblastiaceae and<br />
have not been given generic recognition. In all probability this will be<br />
transferred to Anisomeridium.)<br />
75. Ostiole eccentric; ascospores elliptical, with septum ± median,<br />
30-40 × 10-12 µm; microconidia broadly elliptical or ovoid,<br />
4.5-5 × 2.5-3.5 µm; Zaire...................................................... A. epiphyllum (V_zda) R. C. Harris<br />
74. Growing on liverworts; ostiole apical; ascospores broadly fusifrom to<br />
subclavate, with ± median septum, smooth, 33-42 × 10-12.5 µm;<br />
macroconidia forming cirrhi?, embedded in gelatin, oblong,<br />
5-7 × 3.5-4.5 µm; Brazil ................................................. A. endobrya (Döbbler & Poelt) R. C. Harris<br />
NOTES<br />
Anisomeridium adnexum (Müll. Arg.) R. C. Harris, comb. nov.<br />
Arthopyrenia adnexa Müll. Arg., Flora 66: 304. 1883. Leiophloea adnexa (Müll. Arg.) Riedl,<br />
Sydowia 16: 266. 1962(1963). Type. BRAZIL. São Paulo: Apiaí, Sep, Puiggari 240 (G, lectotype,<br />
selected here), May, Puiggari 1007 (G, syntype). The other syntype, Glaziou 5094, is A.<br />
leptospermum.<br />
Verrucaria thelena var. albidior Nyl.<br />
Ann. Sci. Nat. Bot. sér. 4, 20. 254. 1863. Type. NICARAGUA. Verr. 22 ex hb. Tuckerman (H-Nyl<br />
714, lectotype, Hawskworth, 1985); HONG KONG. Verr. 21 ex hb. Tuckerman (H-Nyl 715,<br />
syntype).<br />
Hawksworth (1985) suggested the lectotype might belong in Anisomeridium. The structure of<br />
the ascoma and rather coarse physes suggest it is better included in Arthopyrenia. Its exact<br />
disposition awaits further work on tropical Arthopyreniae. The syntype is a Mycomicrothelia in<br />
the group with thickened endospore. Its disposition is also pending more study.<br />
Anisomeridium albidoatrum (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria albidoatra Nyl., Flora 69: 178. 1886. Arthopyrenia albidoatra (Nyl.) Zahlbr., Cat. lich.<br />
univ. 1: 300. 1921. Type. SÃO TOMÉ. Altit. 20 m, Moller (H-Nyl 827, lectotype (selected here); H-Nyl<br />
826, syntype).<br />
Anisomeridium americanum (Massal.) R. C. Harris, comb. nov.<br />
Arthopyrenia americana Massal., Ric. auton. lich. crost. 170, f. 341. 1852. Type. "Vive n'ell<br />
America sui tronchi degli alberi."[ex hb. Fee as Pyrenula nitida v. americana Fee] (VER,<br />
holotype).<br />
Verrucaria limitans Nyl., Flora 49: 295. 1866. Arthopyrenia limitans (Nyl.) Müll. Arg., Flora 6:<br />
306. 1883. Type. CUBA. [Wright] (H-Nyl 1714, holotype; M, isotypes).<br />
Arthopyrenia ceylonensis Müll. Arg., Flora 66: 289. 1883. Type. CEYLON. 1876, Thwaites 174 (G,<br />
lectotype; G, isolectotypes).<br />
Pleurotrema inspersum Müll. Arg., Bot. Jahrb. Syst. 6: 389. 1885. Ditremis inspersa (Müll.<br />
Arg.) Clements, The Genera of Fungi 173. 1909. Type. CUBA. Wright, Verr. Cub. 123 (H-Nyl<br />
1715, FH-Tuck 3948, US, isotypes) .<br />
Pleurotrema pyrenuloides Müll. Arg., Rep. Meetings Australas. Assoc. Advancem. Sci. 6: 462.<br />
1895. Ditremis pyrenuloides (Müll. Arg.) Makhija & Patwardhan, Biovigyanam 16: 22. 1990.<br />
Type. AUSTRALIA. Queensland, Knight 49 (G, lectotype, selected here).<br />
Didymella labiata Vainio, Hedwigia 38(Beibl.): 258. 1899. Type. GUADELOUPE. Basse Terre,<br />
Duss 503 (TUR-Vainio 32200, holotype).<br />
Arthopyrenia media Harmand, Bull. Soc. Sci. Nancy, sér. 3, 12: 130. 1911. Type. NEW CALEDONIA.<br />
Pionnier (DUKE, holotype).<br />
136
Anisomeridium angulosum (Müll. Arg.) R. C. Harris, comb. nov.<br />
Tomasellia (sect. Syngenosorus) angulosa Müll. Arg., Bot. Jahrb. Syst. 6: 397. 1885. Type. CUBA.<br />
Wright, Verr. Cub. 127 (G, holotype).<br />
Anisomeridium antillarum (Vainio) R. C. Harris, comb. nov.<br />
Arthopyrenia antillarum Vainio, Ann. Acad. Sci. Fenn. ser. A, 6(7): 207. 1915. Type. VIRGIN<br />
ISLANDS. St. John: Macombo, 20 Mar 1906, Boergesen (TUR-Vainio 32115).<br />
Anisomeridium australiense (McCarthy) R. C. Harris, comb. nov.<br />
Ditremis australiensis Mc Carthy, Austral. Syst. Bot. 5: 125. 1992. Type. AUSTRALIA. New South<br />
Wales: Newcastle region, 3 kn Se of Stroud Road, 1 km S of Stroud Mt., creek beside Ducks<br />
Hill road, on deeply shaded semi-aquatic and aquatic rhyolite, 22 Apr 1990, McCarthy 366<br />
(MEL, holotype).<br />
Ditremis bispora Makhija & Patwardhan,<br />
Biovigyanam 16: 14. 1990. Type. ANDAMAN ISLANDS. South Andaman:, Baratang Islands,<br />
Baludera, 23 Feb 1985, Sethy & Nagarkar 85.795 (AMH, holotype).<br />
= Arthopyrenia<br />
Anisomeridium carinthiacum (Steiner) R. C. Harris<br />
in Egan, Bryologist 90: 163. 1987. Arthopyrenia carinthiaca Steiner, Österr. Bot. Z. 63: 335. 1913.<br />
Type. AUSTRIA. Carinthia, near Klagenfurt, Steiner (US, isosyntype).<br />
Arthopyrenia dimidiata Fink in Hedrick, Mycologia 25: 306. 1934. Type. CONNECTICUT. Litchfield<br />
County: near Ellsworth, 1895, Green (MICH, holotype).<br />
Anisomeridium complanatum (Makhija & Patwardhan) R. C. Harris, comb. nov.<br />
Ditremis complanata Makhija & Patwardhan, Biovigyanam 16: 15. 1990. Type. ANDAMAN ISLANDS.<br />
South Andaman: Wimbeliganj, Kalatang, 16 Feb 1985, Sethy & Nagarkar 85.145 (AMH,<br />
holotype).<br />
Ditremis aggregata Makhija & Patwardhan, Biovigyanam 16: 13. 1990. Type. INDIA. Kerala,<br />
Sabrimalai Hills, Patnamathatta to Sabrimalai, 29 Jan 1983, Patwardhan & Sethy 83.470 (AMH,<br />
holotype).<br />
Ditremis harrisii Makhija & Patwardhan, Biovigyanam 16: 17. 1990. Type. INDIA. Karnataka,<br />
Sirsi-Kumtha Road, Ammanhalli, 12 Dec 1974, Nagarkar & Patwardhan 74.2619 (AMH,<br />
holotype).<br />
Ditremis karnatakensis Makhija & Patwardhan, Biovigyanam 16: 18. 1990. Type. INDIA. Karnataka,<br />
Sirsi-Kumtha Road, Devimane ghat, 12 Dec 1974, Kulkarni 74.2648 (AMH, holotype).<br />
Ditremis muelleri Makhija & Patwardhan, Biovigyanam 16: 20. 1990. Type. INDIA. Karnataka,<br />
Hosar Ghat, 26 Dec 1981, Nagarkar & Makhija 81.664 (AMH, holotype).<br />
Ditremis straminicolor Makhija & Patwardhan, Biovigyanam 16: 22. 1990. Type. ANDAMAN<br />
ISLANDS. South Andaman: Baratang Island, Wrafter's Creek, 21 Feb 1985, Patwardhan & Sethy<br />
85.534 (AMH, holotype).<br />
Anisomeridium complanatum is the most common species of the genus in India. The multiple<br />
names above are based on minor, ecologically induced variations. The species almost entirely<br />
replaces pantropical A. americanum in India and is barely seperable from it by the smooth<br />
ascospores and usually a very well developed, hard clypeus. The epithet "complanatum" has<br />
been selected from among the simultaneously published names since the complanate aspect<br />
of the ascomata due to the broad clypeus the most characteristic feature of the taxon.<br />
Anisomeridium consimile (Vainio) R. C. Harris, comb. nov.<br />
Arthopyrenia consimilis Vainio, Ann. Acad. Sci. Fenn. ser. A, 19(15): 14. 1923. Type. PHILIPPINE<br />
ISLANDS. Palawan: Taytay, on Pongamia mitis on seashore, Apr 1913, Merrill, Bur. Sci. 9053<br />
(TUR-Vainio 32110).<br />
137
Anisomeridium consobrinum (Nyl.) Aptroot<br />
Biblioth. Lichenol. 57: 21. 1995. Verrucaria consobrina Nyl., Ann. Sci. Nat. Bot. sér. 4, 15: 53.<br />
1861. Arthopyrenia consobrina (Nyl.) Müll. Arg., Flora 66: 317. 1883. Type. NEW CALEDONIA.<br />
Viellard 1838 (H-Nyl 1666, holotype; G, isotype).<br />
Anisomeridium elegans R. C. Harris, sp. nov.<br />
Thallo corticato, glauco, UV+ aureo, ostiolo eccentrico, ascosporis ovoideis, septis valde submedianis,<br />
laevibus, 51-57 × 23-26 µm<br />
Type. PUERTO RICO. Distr. Mayagüez: Maricao State Forest, just NE of Universidad Católica Estación<br />
Biologica, along Hwy 120 at km 15.8, along trail on hill top, ca. 700 m, semi-dry forest, 5 Jun 1988, Harris<br />
22280 (NY, holotype).<br />
Thallus corticate, epiphloeodal, glaucous green, glossy, UV+ yellow (lichexanthone), sometimes weakly<br />
so. Ascomata immersed, covered by thallus and only visible as a weakly raised bump, lageniform, 0.5-0.8 ×<br />
0.4-0.5 mm; ostiole eccentric or lateral; wall entirely melanized or open below. Asci cylindrical to subclavate,<br />
ca. 200-300 × 40-55 µm, with eight subbiseriate spores. Ascospores ovoid, slightly constricted at markedly<br />
submedian septum, smooth, 45-57 × 18-26 µm; pycnidia not found.<br />
Anisomeridium elegans is related to A. infernale and A. nemorosum with similarly corticate thalli, but<br />
differs in size of ascomata and asci and in size and shape of ascospores.<br />
Additional specimen. PUERTO RICO. As type, ca. 600 m, moist forest on slope with tree ferns and palms,<br />
5 Jun 1988, Harris 22319 (NY).<br />
Anisomeridium endobryum (Döbbler & Poelt) R. C. Harris, comb. nov.<br />
Arthopyrenia endobrya Döbbler & Poelt, Pl. Syst. Evol. 138: 276. 1981. Type. BRAZIL. São<br />
Paulo: Ilha Comprida bei Cananeia, 15 Jul 1979, Kalb & Poelt (M, isotype).<br />
Anisomeridium epiphyllum (V_zda) R. C. Harris, comb. nov.<br />
Pleurotrema epiphyllum V_zda, Folia Geobot. Phytotax. 13: 99. 1978. Ditremis epiphylla (V_zda)<br />
Makhija & Patwardhan, Makhija & Patwardhan, Biovigyanam 16: 15. 1990. Type. ZAIRE (not<br />
seen).<br />
Anisomeridium excellens (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria excellens Müll. Arg., Bot. Jahrb. Syst. 6: 405. 1885. Type. CUBA. Wright, Verr.<br />
Cub. 79 (G, lectotype (selected here); UPS, US, isolectotypes); Verr. Cub. 78 (G, syntype).<br />
Anisomeridium flavopallidum (Makhija & Patwardhan) R. C. Harris, comb. nov.<br />
Ditremis flavopallida Makhija & Patwardhan, Biovigyanam 16: 16. 1990. Type. INDIA. Maharastra,<br />
Karnala, on the way to Ransai, 27 Aug 1974, Patwardhan & Nagarkar 74.431 (AMH, holotype).<br />
Anisomeridium foliicola R. Sant. & Tibell<br />
This lichen is apparently often produces only its very characteristic pycnidia. Santesson and Tibell<br />
suggested that they might be producing microconidia. From the size and shape of the conidia in Costa Rican<br />
material distributed by Lücking, I consider them to be macroconidia. This would suggest that since ascomata<br />
are rare, dispersal is via these macroconidia. It is also noteworthy that the pycnidia of A. foliicola have a well<br />
developed periphysoid plug which apparently relaeases the conidia single file. A similar feature is described<br />
for the possibly synonymous and therefore illegitimate pycnidial genus Compsosporiella Sanarkan & Sutton.<br />
Anisomeridium glaucescens (Müll. Arg.) R. C. Harris, comb. nov.<br />
138
Arthopyrenia glaucescens Müll. Arg., Bot. Jahrb. Syst. 6: 405. 1885. Type. CUBA. Wright,<br />
Verr. Cub. 77 (G, holotype; US, isotype).<br />
Anisomeridium gregale R. C. Harris, sp. nov.<br />
Thallo endolithico, glauco, UV-, ascomata aggregata, ostiolo apicale et ascosporis parvis, anguste<br />
ovoideis, 11-13 × 3.5-4.5 µm.<br />
Type. PUERTO RICO. Distr. Arecibo: ca. 1 km N of Ciales on Hwy. 149, ca. 220 m, humid forest over limestone<br />
in haystack hills, on limestone, 11 Jun 1988, Harris 22626 (NY, holotype)<br />
Thallus endolithic in limestone, glaucous, UV-. Ascomata complanate, ca. 0.3 mm diam., initially solitary,<br />
later forming small groups ca. 1.0 mm across (regeneration ?); ostiole apical; open below; clypeus spreading<br />
laterally. Asci cylindrical, ca. 55 × 8 µm, with eight biseriate spores. Ascospores narrowly ovoid, weakly<br />
constriicted at slightly submdian septum, smooth, 11-13 × 3.5-4.5 µm; microconidia elliptical or ovoid, 3 × 1.5<br />
µm, macroconidia not found.<br />
Microscopically A. gregale is similar to the corticolous A. albisedum but seems to be distinct in growing<br />
on rock, in aggregated ascomata and in spreading clypeus. It is known only from the type collection.<br />
Anisomeridium holopolium (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria holopolia Nyl., Flora 50: 196. 1867, nom. nud. Bull. Soc. Linn. Normandie sér. 2,<br />
2: 131. 1868. Type. NEW CALEDONIA. Loyalty Islands: Lifou, 1864, Deplanche (H-Nyl 1671,<br />
lectotype (selected here); H-Nyl 1670, syntype).<br />
Anisomeridium immersum (Makhija & Patwardhan) R. C. Harris, comb. nov.<br />
Ditremis immersa Makhija & Patwarhan, Biovigyanam 16: 17. 1990. Type. INDIA. Kerala,<br />
Sabrimalai Hills, Patanamthatta to Sabrimalai, 29 Jan 1983, Makhija & Kekre 83.431 (AMH,<br />
holotype).<br />
Additional specimen. INDIA. Karnataka, 38-48 km from Sirsi, Kumala-Sirsi Rd., Devimane<br />
Ghat, evergreen forest, 26 Feb 1977, Hale 48030 (US).<br />
Anisomeridium indicum (Makhija & Patwardhan) R. C. Harris, comb. nov.<br />
Ditremis indica Makhija & Patwardhan, Biovigyanam 16: 18. 1990. Type. INDIA. Maharastra,<br />
Karnala forest, 28 Aug 1974, Nagarkar & Prabhu 74.456 (AMH, holotype).<br />
Anisomeridium infernale (Mont.) R. C. Harris, comb. nov.<br />
Verrucaria infernalis Mont., Ann. Sci. Nat. Bot. sér. 3, 16: 68. 1851. Arthopyrenia infernalis<br />
(Mont.) Müll. Arg., Flora 68: 262. 1885. Type. FRENCH GUIANA. Leprieur 725 (PC, syntype).<br />
Pleurotrema anisomerum Müll. Arg., Flora 68: 251. 1885. Ditremis anisomera (Müll. Arg.)<br />
Makhija & Patwardhan, Biovigyanam 16: 14. 1990. Type. FRENCH GUIANA. Leprieur 1248 (G,<br />
holotype).<br />
Arthopyrenia infernalis var. rhychostoma Vainio, Proc. Amer. Acad. Arts Sci. 58: 147. 1923. Type.<br />
TRINIDAD. Sangre Grande, 1912-1913, Thaxter 73 (FH, isotype).<br />
Verrucaria clandestina sensu Mont., Ann. Sci. Nat. Bot. sér. 2, 19: 59. 1843. (non Verrucaria<br />
clandestina (Ach.) Mont. = Myriotrema).<br />
Trypethelium karnatakense R. C. Harris, nom. nov.<br />
Pleurotrema megaspermum Makhija & Patwardhan, Biovigyanam 16: 24. 1990. Type. INDIA.<br />
Karnataka, Agumbe, 16 Dec 1974, Kulkarni 74.2991 (AMH, holotype), non Trypethelium<br />
megaspermum Mont.<br />
Anisomeridium laevigatum (McCarthy) R. C. Harris, comb. nov.<br />
139
Ditremis laevigata McCarthy, Muelleria 8: 1: 1993. Type. NEW ZEALAND (not seen).<br />
Anisomeridium leptospermum (Zahlbr.) R. C. Harris, stat. et comb. nov.<br />
Arthopyrenia adnexa var. leptosperma Zahlbr. in Skottsberg, Nat. hist. Juan Fernandez 2: 320.<br />
1924. Type. CHILE. Juan Fernández, Masafuera, Quebrada de la Lobería, 300 m, auf alter<br />
Lumaborke, 17 Feb 1917, C. & I. Skottsberg (UPS, holotype).<br />
Anisomeridium leucochlorum (Müll. Arg.) R. C. Harris<br />
in Egan, Bryologist 90: 163. 1987. Arthopyrenia leucochlora Müll. Arg., F!ora 66: 287. 1883.<br />
Ditremis leucochlora (Müll. Arg.) R. C. Harris, Some Florida Lichens 33. 1990. Type.<br />
MASSACHUSETTS. New Bedford, on Acer, Willey (MICH, MIN, US, isotypes)<br />
Ditremis macrospora Makhija & Patwardhan<br />
Biovigyanam 16: 19. 1990. Type. INDIA. Agasthi Hills, Upper Kodayar, 24 Jan 1983, Sethy &<br />
Kekre 83.380 (AMH, holotype).<br />
= Arthopyrenia<br />
Ditremis macrospora R. C. Harris = Anisomeridium aureopunctatum<br />
Anisomeridium magnosporum (Knight) D. Hawksw.<br />
in Galloway, New Zealand J. Bot. 21: 195. 1983. Verrucaria magnospora Knight, Trans. Linn.<br />
Soc. Lond. 23: 99. 1860. Type. NEW ZEALAND (not seen).<br />
Verrucaria magnifica Nyl., Lich. Nov. Zel. 132. 1888. Microthelia magnifica (Nyl.) Müll. Arg., Bull.<br />
Soc. Bot. Belgique 31: 41. 1892. Type. NEW ZEALAND. Knight 349 (H-Nyl 882, holotype?).<br />
Anisomeridium megalosporum (Vainio) R. C. Harris, comb. nov.<br />
Didymosphaeria megalospora Vainio, Proc. Amer. Acad. Arts Sci. 58: 147. 1923. Megalotremis<br />
megalospora (Vainio) Aptroot, Nova Hedwigia 60: 338. 1995. Type. TRINIDAD. St. Ann' Valley,<br />
1912-13, Thaxter 99 (TUR-Vainio 32490, lectotype, Aptroot, 1995).<br />
Anisomeridium monosporum (Makhija & Patwardhan) R. C. Harris, comb. nov.<br />
Ditremis monospora Makhija & Patwardhan, Biovigyanam 16: 20. 1990. Type. INDIA. Karnataka,<br />
Kudremukha, 28 Jan 1980, Nagarkar & Daya Rade 80.291 (AMH, holotype).<br />
Anisomeridium nemorosum R. C. Harris, sp. nov.<br />
Thallo corticato, epiphloeodo, viridescente, UV-, ascomatis magnis, thallo tectis, ostiolo laterale et<br />
ascosporis magnis, fusiformibus, laevibus, 100-117 × 23-27 µm.<br />
Type. FRENCH GUIANA. St-Laurent-du-Moroni: Canton de Maripasoula, Commune de Saül, ca. 6 km N of Saül,<br />
vicinity of Eaux Claires, along Route de Bélizon N of Eaux Claires beyond third bridge, ca. 200 m, 3°37'N,<br />
53°12'W, moist tropical forest over quartz and granite, 4 Sep 1994, Harris 33592 (NY, holotype; B, CAY, US,<br />
isotypes).<br />
Thallus corticate, epiphloeodal, glossy, greenish, UV-. Ascomata immersed, covered by thallus and<br />
visible as slightly raised lageniform bumps, large, ca. 1.5-2.0 × 1.0 mm( including clypeus); ostiole lateral;<br />
clypeus melanized above, broadly expanded. Asci cylindrical, ca. 500 × 50 µm, with eight obliquely uniseriate<br />
spores. Ascospores fusiform, ends distinctly pointed, constricted at ± median septum, smooth, 84-117 × 22-<br />
27 µm; microconidia oblong, 3-5 × 1.5-2 µm; macroconidia not found.<br />
It is possible that this species is the same as Ditremis corticata Makhija & Patwardhan (Mycotaxon 31:<br />
468. 1988). However, I have not seen material of this taxon and considering Makhija and Patwardhan's track<br />
record, this may not even be an Anisomeridium. Additionally the ascospores are reported to be narrower,<br />
140
16.5-23 µm. As I have unpacked only a part of my French Guiana collections, I would guess that A.<br />
nemorosum will prove to be rather weedy on trunks in tropical moist forests.<br />
Additional specimens. FRENCH GUIANA. As type, Sentier Botanique, 200-300 m, 13-14 Nov 1990, Buck<br />
18822, 300-400 m, 7 Sep 1994, Harris 33819 (both NY). TRINIDAD. Cats Hill Road, ca. 6 mi from junction with<br />
Rio Claro-Guayaguare road, virgin lowland rain forest, ca. 200 ft., 18 Apr 1972, Hale 38195 (US).<br />
Anisomeridium nidulans (Müll. Arg.) R. C. Harris, comb. nov.<br />
Arthopyrenia nidulans Müll. Arg., Flora 68: 326. 1885. Type. SRI LANKA. Nieter, ex hb. Hampe,<br />
1877 (G, holotype).<br />
This is keyed out as having an apical ostiole. It is very difficult to determine the actual orientation<br />
and may actually be somewhat eccentric.<br />
Anisomeridium pacificum (McCarthy) R. C. Harris, comb. nov.<br />
Ditremis pacifica McCarthy, Muelleria 8: 3. 1993. Type. HAWAIIAN ISLANDS. Oahu (not seen).<br />
Anisomeridium palavanum (Vainio) R. C. Harris, comb. nov.<br />
Microthelia palavana Vainio, Ann. Acad. Sci. Fenn. ser. A., 15(6): 347. 1921. Type. PHILIPPINE<br />
ISLANDS. Palawan: Taytay, Apr 1913, Merrill, Bur. Sci. 9036 (BM, NY, isotypes).<br />
Arthopyrenia pongamiae Vainio, Ann. Acad. Sci. Fenn. ser. A, 19(15): 13. 1923. Leiophloea<br />
pongamiae (Vainio) Riedl, Sydowia 16: 271. 1962 [1963]. Type. PHILIPPINE ISLANDS. Palawan:<br />
Taytay, on Pongamia mitis, Apr 1913, Merrill, Bur. Sci. 9035 (TUR-Vainio 32111, holotype; BM,<br />
NY, isotypes).<br />
Ditremis andamanica Makhija & Patwardhan, Biovigyanam 16: 14. 1990. Type. ANDAMAN ISLANDS.<br />
North Andaman: Diglipur Range, Kalara, 4 Jan 1986, Sethy & Patwardhan 86.383 (AMH,<br />
holotype).<br />
Ditremis punctata Makhija & Patwarhan, Biovigyanam 16: 21. 1990. Type. ANDAMAN ISLANDS.<br />
Little Andaman: South Bay, 3 Mar 1985, Nagarkar & Patwarhan 85. 1026 (AMH, holotype).<br />
Anisomeridium planiusculum (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria planiuscula Expos. syn. pyrenocarp. 58. 1858. Arthopyrenia planiuscula (Nyl.)<br />
Zahlbr., Cat. lich. univ. 1:310. 1921. Type. JAMAICA? Ad cort. Quassia excelsa in hb. Fée (H- Nyl<br />
732, holotype).<br />
Anisomeridium polycarpum (Müll. Arg.) R. C. Harris, comb. nov.<br />
Pleurotrema polycarpum Müll. Arg., J. Bot. (Morot) 7: 109. 1893. Ditremis polycarpa (Müll.<br />
Arg.) Makhija & Patwardhan, Biovigyanam 16: 21. 1990. Type. NEW CALEDONIA. Cascade de la<br />
Ferme-Modèle, 1892, Balansa (G, holotype).<br />
Anisomeridium rockii (Zahlbr.) R. C. Harris, comb. nov.<br />
Pleurotrema rockii Zahlbr., Ann. Mycol. 10: 362. 1912. Ditremis rockii (Zahlbr.) Makhija &<br />
Patwardhan, Biovigyanam 16: 22. 1990. Type. HAWAIIAN ISLANDS. Lanai, Koela Valley, Rock 132<br />
(FH, isotype).<br />
Anisomeridium sphaerocarpum (Vainio) R. C. Harris, comb. nov.<br />
Didymella sphaerocarpa Vainio, Ann. Acad. Sci. Fenn. ser. A, 19(15): 14. 1923. Type. PHILIPPINE<br />
ISLANDS. Luzon: Benguet Province, Pauai, May 1909, Merrill, Bur. Sci. 6649 (TUR-Vainio 32297,<br />
holotype).<br />
Anisomeridium stromaticum R. C. Harris, sp. nov.<br />
141
Species distincta thallo crasso corticato, UV-, ascomatis pluribus subtus clypeo unico, pycnidiis pluribibus<br />
subtus clypeo unico, ascosporis fusiformibus septo submediano laevibus, 15-23 × 4.5-5.5 µm.<br />
Type. DOMINICAN REPUBLIC. Prov. Monte Cristi: El Morro de Monte Cristi, 237 m, limestone mesa by sea,<br />
19°54'N, 71°39'W, 8 Jan 1987, Harris 19534 (NY, holotype; JBSD, isotype).<br />
Thallus epiphloeodal, thick (to ca. 75 µm), corticate, gray, ± dull, UV-; cortex to 25 µm in places.<br />
Ascomata mostly immersed in bark, several under a single clypeus, ca. 0.3 mm diam., occasionally solitary,<br />
then larger, ca. 0.7 mm diam.; ostiole apical; wall not melanized; clypei often fusing in more extensive, black,<br />
irregular pseudostromata, 3 × 1-2 mm. Asci cylindrical, ca. 85 × 12 µm, with eight biseriate spores.<br />
Ascospores fusiform, slightly constricted at ± submedian septum, smooth, 15-23 × 4.5-5.5 µm; pycnidia as in<br />
ascomata, usually several under a single clypeus; microconidia elliptical, 3 × 1.5-2 µm; macroconidia elliptical,<br />
5-6 × 2-2.5 µm.<br />
This is a very peculiar species and I am not entirely sure that it is not an abnormal form of some other<br />
species. A somewhat similar situation occurs in the type of A. polycarpum which I interpreted as the<br />
ascomata growing in abnormal proximity. I may have been wrong and the type should be re-examined.<br />
However, in A. stromaticum the fact that both micropycnidia and macorpycnidia are clustered under a single<br />
clypeus lends support to interpreting this as the normal state for this species. The thick, corticate thallus is<br />
also distinctive and somewhat resembles that of A. glaucescens which differs in larger ascospores.<br />
Anisomeridium subatomarium (Knight) R. C. Harris, comb. nov.<br />
Verrucaria subatomaria Nyl. ex Knight, Trans. Proc. New Zealand Inst. 15: 356. 1883.<br />
Arthopyrenia subatomaria (Knight) Müll. Arg., Bull. Soc. Bot. Belgique 31: 40. 1892. Type. NEW<br />
ZEALAND. Knight 14 (H-Nyl 884, isotype?).<br />
Anisomeridium subbiforme (Knight) R. C. Harris, comb. nov.<br />
Verrucaria subbiformis Knight, Trans. Proc. New Zealand Inst. 8: 326. 1875. Arthopyrenia<br />
subbiformis (Knight) Müll. Arg., Bull. Soc. Bot. Belgique 31: 40. 1892. Type. NEW ZEALAND.<br />
Knight (G, M, UPS, isolectotypes?).<br />
Anisomeridium subnectendum (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria subnectenda Nyl., J. Linn. Soc., Bot. 20: 61. 1883. Arthopyrenia subnectenda (Nyl.)<br />
Zahlbr., Cat. lich. univ. 1: 327. 1922. Type. MALAYSIA. Malacca, 1864, Maingay 161 (H-Nyl 835,<br />
holotype; BM, isotype).<br />
Verrucaria anisomera Nyl., Flora 56: 364. 1876, nom. nud. "Type". CUBA. Wright, Verr. Cub. 105<br />
(H-Nyl 1673).<br />
Arthopyrenia xylogena Müll. Arg., Flora 66: 290. 1883. Leiophloea xylogena (Müll. Arg.) Riedl,<br />
Sydowia 16: 270. 1962 [1963].Type. BRAZIL. São Paulo: Apiaí, Puiggari 494 (G, holotype).<br />
Arthopyrenia exstans Müll. Arg., Rep. & Meetings Australas. Assoc. Advancem. Sci. 6: 451. 1895.<br />
Type. AUSTRALIA. Queensland, 1887, Knight 43 (G, holotype)<br />
Anisomeridium subnexum (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria subnexa Nyl., Bull. Soc. Linn. Normandie, sér. 2, 7: 181. 1873. Arthopyrenia subnexa<br />
(Nyl.) Müll. Arg., Hedwigia 30: 188. 1891. Type. ANDAMAN ISLANDS. 1867, Kurz 75 (H-Nyl 834,<br />
holotype; M, isotype).<br />
Anisomeridium tarmugliense (Makhija & Patwarhan) R. C. Harris, comb. nov.<br />
Ditremis tarmugliensis Makhija & Patwarhan, Biovigyanam 16: 22. 1990. Type. ANDAMAN ISLANDS.<br />
South Andaman: Wandoor, Tarmugli Island, Patwardhan & Nagarkar 85.1890 (AMH, holotype).<br />
Anisomeridium throwerae R. C. Harris, sp. nov.<br />
142
Species distincta macropycnidiis rostratis, rostro capillaceo ad 1.5 mm, macroconidiis oblongis in medio<br />
constrictis, 8-10 × 4-5 µm, ascosporis 12-17 × 5-6.5 µm.<br />
Type. HONG KONG. New Territories: near Cheung Uk, 21 oct 1977, Thrower 2900 (NY, holotype).<br />
Thallus gray green, endophloeodal, UV-. Ascomata immersed, subglobose, ca. 0.2 mm diam.; ostiole<br />
apical. Asci cylincrical, ca. 45 × 9 µm, with eight biseriate spores. Ascospores ovoid, slightly constricted at<br />
submedian septum, 12-17 × 5-6.5 µm; macropycnidia with a long, to 1.5 mm, beak through which the<br />
macroconidia are extruded in single file; macroconidia oblong, constricted in middle, 8-10 × 4-5 µm.<br />
The micropycnidia associated with A. nyssaegenum have a short beak. Similar long-beaked<br />
macropycnidia are known in Anisomeridium sp. Buck 22812 for which no teleomorph is present and 1 or 2<br />
undescribed species from Brazil. Also, from the illustration, it seems worth investigating Compsosporiella<br />
Sankaran & Sutton (Mycol. Res. 93: 1289. 1991) as a possible macropycnidial state of Anisomeridium.<br />
Anisomeridium truncatum (Müll. Arg.) R. C. Harris, comb. nov.<br />
Arthopyrenia truncata Müll. Arg., Flora 66: 289. 1883. Leiophloea truncata (Müll. Arg.) Rield,<br />
Sydowia 16: 273. 1962 [1963]. Type. BRAZIL. São Paulo: Apiaí, Puiggari 249 (G, holotype).<br />
Anisomeridium ubianum (Vainio) R. C. Harris, comb. nov.<br />
Arthopyrenia ubiana Vainio, Ann. Acad. Sci. Fenn. ser. A, 19(15): 12. 1923. Type. PHILIPPINES.<br />
Ubian Island, 12 Oct 1906, Merrill, Bur. Sci. 5403 (TUR-Vainio 32109, holotype; BM, isotype).<br />
Verrucaria consobrina var. glaucescens Nyl., Bull. Soc. Linn. Normandie, sér. 2, 2: 131. 1868.<br />
Arthopyrenia consobrina var. glaucescens (Nyl.) Zahlbr., Cat. lich. univ. 1: 323. 1922. Type.<br />
NEW CALEDONIA. Loyalty Islands: Lifou, 1864, Deplanche (H-Nyl 1664, lectotype (selected here);<br />
H-Nyl 1665, syntype?).<br />
Ditremis microcarpa Makhija & Patwarhan, Biovigyanam 16: 19. 1990. Type. ANDAMAN ISLANDS.<br />
Middle Andaman: Long Island Range, Elphiston Bay, 23 Dec 1985, Nagarkar & Patwardhan<br />
85.2219 (AMH, holotype).<br />
Anisomeridium uniseriale (Zahlbr.) R. C. Harris, comb. nov.<br />
Microthelia uniserialis Zahlbr., Ann. Cryptog. Exot. 5: 199. 1932. Pleurotrema uniseriale (Zahlbr.)<br />
D. Hawksw., Bull. Brit. Mus. Nat. Hist. Bot. 14(2): 171. 1985. Type. SOUTH AFRICA. Knysna, Hof<br />
van Eden, v. d. Byl 869 (LD, isotype).<br />
Hawksworth (1985) provided a modern description and photographs of habit and section of<br />
ascoma.<br />
Anisomeridium verrucosum (Makhija & Patwardhan) R. C. Harris, comb. nov.<br />
Ditremis verrucosa Makhija & Patwardhan, Mycotaxon 31: 469. 1988. Megalotremis verrucosa<br />
(Makhija & Patwardhan) Aptroot, Biblioth. Lichenol. 44: 126. 1991. Type. ANDAMAN ISLANDS (not<br />
seen).<br />
Anisomeridium viridescens (Coppins) R. C. Harris, comb. nov.<br />
Arthopyrenia viridescens Coppins, Lichenologist 20: 322. 1988. Type. SCOTLAND. Argyll Main,<br />
Taynult, Glen Nant prope Tailor's Leap, 27/01.27, alt. 45 m, ad corticem laevem Coryli, 3<br />
August 1980, B. J. Coppins 8031 (NY, isotype).<br />
When Coppins first sent a specimen of this species I studied it rather casually and moved on<br />
since it was not an American taxon but the presence of macroconidia in an Arthopyrenia was an<br />
anomaly which kept nagging in the back of my mind. After more careful study of the ascus and<br />
hamathecium, the anomaly was resolved as both proved to be of the Anisomeridium- type.<br />
STRIGULACEAE Fr.<br />
143
Syst. Orb. Veg. 1: 110. 1825. Type. Strigula Fr.<br />
Phyllobatheliaceae Bitter & Schilling, Hedwigia 67: 272, 291. 1927. Type. Phyllobathelium (Müll. Arg.)<br />
Müll. Arg., syn. nov.<br />
The Strigulaceae are constituted in a different manner from Harris (1975). Eriksson (1981) pointed out<br />
the family should be restricted excluding the Monoblastiaceae, Pleurotremataceae and Xanthopyreniaceae, a<br />
proposal supported by more recent studies. The Phyllobatheliaceae are placed in synonymy here as they<br />
have the same ascus type, apically free physes, ellipsoidal microconidia, septate macroconidia with gelatinous<br />
appendages and trentepohlioid photobiont.<br />
1. Ascomata solitary, not surrounded by pseudostromatic wart filled with dark<br />
granular material; pycnidia solitary; macroconidia usually with eccentric attachment<br />
and bipolar gelatinous appendages (except species with muriform ascospores?);<br />
on leaves, bark, wood or rock ........................................................................................................Strigula<br />
1. Ascomata 1-several, surrounded by a pseudostromatic wart filled with dark<br />
granular material; pycnidia aggregated; macroconidia with basal attachment<br />
and an apical gelatinous cap; on leaves ..........................................................................[Phyllobathelium]<br />
PHYLLOBATHELIUM (Müll. Arg.) Müll. Arg.<br />
Phyllobathelium is not known from North America. Following Santesson (1952) and Lücking (1992) I<br />
would recognize three foliicolous and one corticolous species. Mayrhofer (1987) added a corticolous and a<br />
saxicolous species. On this account I initially considered uniting Strigula and Phyllobathelium. However,<br />
examination of the species recognized by Santesson and Lücking revealed a number of characters that<br />
support recognition of both genera, pseudostromatic ascomatal warts, aggregated pycnidia and basally<br />
attached macroconidia (illustrated by V_zda, 1984) with an apical cap (not tapering appendages as in<br />
Strigula). The two non-foliicolous species recognized by Mayrhofer are transferred to Strigula below. Although<br />
Mayrhofer understood their general relationship, I suspect Strigula did not occur to him since in 1987 the<br />
present day plethora of muriform-spored species were as yet unrecognized.<br />
STRIGULA Fr.<br />
Raciborskiella Höhnel, Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturw. Cl., Abt. 1 ser. 1, 118: 1176. 1909.<br />
Type. Clypeolum talumae Racib. (= R. janeirensis (Müll. Arg.) R. Sant.).<br />
Dichoporis Clements, Genera of Fungi 40, 173. 1909. Type. Dichoporis schizospora (Vainio) Clements (=<br />
Strigula mediterranea Etayo).<br />
Diporina Clements, Genera of fungi 40, 173. 1909. Type. Diporina subsimplicans (Nyl.) Clements (=<br />
Strigula subsimplicans (Nyl.) R. C. Harris in Hafellner & Kalb).<br />
Discosiella H. & P. Sydow, Leafl. Philipp. Bot. 5: 1546. 1912, nom. illeg. (macroconidia) Type. Discosiella<br />
cylindrospora H. & P. Sydow (= Strigula sp.).<br />
Phylloporis Clements, Genera of Fungi 41, 173. 1909. Type. Phylloporis phyllogena (Müll. Arg.) Clements (=<br />
Strigula phyllogena (Müll. Arg.) R. C. Harris).<br />
Sagediomyces Ciferri & Tomaselli, Ist. Bot. Reale Univ. Reale Lab. Crittog. Pavia Atti ser. 5, 10: 30, 57.<br />
1953. Type. Sagediomyces affinis (Massal.) Ciferri & Tomaselli (= Strigula affinis (Massal.)<br />
R. C. Harris).<br />
Strigulomyces Ciferri & Tomaselli, Ist. Bot. Reale Univ. Reale Lab. Crittog. Pavia Atti ser. 5, 10, 34, 61.<br />
1953, nom. illeg. Type. Strigulomyces elegans (Fée) Ciferri & Tomaselli (= Strigula smaragdula Fr.)<br />
Shanoria Anahosur, Nova Hedwigia 14: 405. 1967[1968], nom. illeg. (macroconidia) Type. Shanoria<br />
indica Anahosur (= Strigula nitidula Mont.?).<br />
Specific limits in Strigula pose a number of problems. The corticolous and saxicolous taxa have the usual<br />
problem of overlapping ascospore and macroconidia sizes and possible significance of spore arrangement in<br />
the asci. The foliicolous taxa seem to me (a nonspecialist) based more on the aspect or pattern of growth of<br />
144
the species or strain of the photobiont and relatively little on "mycological" characters (e.g., Lücking, 1992). I<br />
feel that Santesson (1952) set the pattern for the recognition of what I would consider microspecies and this<br />
has been maintained or exaggerated since. It is thus no surprise that a seeming generic shift in photobiont has<br />
led to the recognition of Phylloporis. (The generic limits in the Trentepohliaceae are in flux and this difference<br />
may not exist.) Further the supra-/subcuticular "thallus" is a character of the photobiont as probably is the<br />
occurence of Raciborskiella on the underside of leaves. The asci, ascospores, microconidia and macroconidia<br />
are identical and thus I can see no shred of mycological support for the recognition of either genus. (The<br />
mycobiont characters of some of the species currently assigned to different genera seem so similar I think it<br />
would be worth using molecular techniques to verify whether they are or are not the same fungus associated<br />
with different algae.) Also the tide seems to be against granting even more striking photobiont differences<br />
generic status, e.g., Lobariaceae and most recently, Hymeneliaceae (Lutzoni & Brodo, 1995).<br />
Several species which do not occur in Florida or even in North America are described below as they<br />
provide new and interesting information for our understanding of the genus, corticolous species with black<br />
hypothallus, compound ascomata, lateral ostiole, muriform-spored taxa, beaked macropycnidia, fragmenting<br />
macroconidia and macroconida basally attached with apical or basal gealtinous caps.<br />
KEY TO NORTH AMERICAN SPECIES<br />
1. Growing on bark, wood or rock ................................................................................................................. 2<br />
2. Ascospores transversely 1-7-septate.................................................................................................... 3<br />
3. Ascospores 1-septate ....................................................................................................................... 4<br />
4. Growing on bark or wood .............................................................................................................. 5<br />
5. Black hypothallus absent........................................................................................................... 6<br />
6. Ascomata unilocular.............................................................................................................. 7<br />
7. Ascospores 12-25 × 4-5.5 µm ........................................................................................... 8<br />
8. Ascospores 12-17 × 4.5-5 µm; macroconidia 7-9 × 2-2.5 µm;<br />
throughout Florida ....................................................... S. viridiseda (Nyl.) R. C. Harris<br />
8. Ascospores 15-25 × 4-5.5 µm; macroconidia 14-19 × 3-4.5 µm;<br />
throughout Florida ................................................................ S. americana R. C. Harris<br />
7. Ascospores smaller, 8-12 × 2.5-3 µm; peninsular<br />
Florida ...................................................................................S. phaea (Ach.) R. C. Harris<br />
6. Ascomata 2-3-locular; ascospores 7-10(-12) × 3-4.5 µm;<br />
Dade County................................................................................. S. connivens R. C. Harris<br />
5. Black hypothallus present; thallus white punctate; ascospores<br />
17-22 × 5-6 µm; South Carolina .....................................................[S. hypothallina R. C. Harris]<br />
4. Growing on limestone.................................................................................................................... 9<br />
9. Ascospores 11-15(-18) × 3.5-5 µm; Dade and Monroe<br />
counties .................................................................................S. bermudana (Nyl.) R. C. Harris<br />
9. Ascospores 18-22 × 5-7.5 µm; Monroe County<br />
(Key West) ............................................................................. S. wilsonii (Riddle) R. C. Harris<br />
145
3. Ascospores 3-7-septate .................................................................................................................. 10<br />
10. Ascospores 7-septate, 24-42 × 5-7.5 µm; macroconida 8-celled,<br />
23-30 × 3.5-5 µm, often over mosses; northeastern North America,<br />
Europe...................................................................................... [S. stigmatella (Ach.) R. C. Harris]<br />
10. Ascospores 3-septate, 14-17 × 3.5-5(-6) µm; macroconidia<br />
4-celled, 13-17 × 3-3.5 µm, Louisiana, Minnesota, Missouri,<br />
Ontario, Europe ........................................................................[S. jamesii (Swinsc.) R. C. Harris]<br />
2. Ascospores with at least 1 or 2 cells longitudinally sepate, submuriform to muriform ........................ 11<br />
11. Ascospores submuriform, 20-33 × 6-10 µm.................................................................................. 12<br />
12. Ascospores 26-33 × 8-10 µm, 7(-8)-septate with 2-6 cells<br />
1-2 longitudinally septate; macroconidia not found; Gulf<br />
Coast of Florida...............................................................................S. griseonitens R. C. Harris<br />
12. Ascospores 20-27 × 6-7.5 µm, 5-7-septate with only 1-few cells<br />
longitudinally septate; mostly midwestern but a single dubious<br />
record from Duval County ....................................S. submuriformis (R. C. Harris) R. C. Harris<br />
11. Ascospores densely muriform, 35-45 × 13-17 µm; macroconidia<br />
not found; ascomata immersed; wall colorless except around<br />
ostiole; Georgia and Louisiana............................................................. [S. laceribracae R. C. Harris]<br />
1. Growing on leaves (mostly modified from Santesson, 1952).................................................................. 13<br />
13. Ascospores 1-septate; macroconidia bacillar, 1-septate or simple................................................... 14<br />
14. Ascospores over 11 × 4 µm ......................................................................................................... 15<br />
15. Ascospores with cells relatively equal in size, 14-22(-24) × 4-5.5 µm;<br />
common, throughout Florida ..........................................................................S. smaragdula Fr.<br />
15. Ascospores with upper cell enlarged, 11-15 × 4-4.5 µm;<br />
Dade County...................................................................................S. subelegans Vainio s. lat.<br />
14. Ascospores smaller, 8-10(-12) × (2-)2.5-3 µm; macroconidia simple;<br />
Seminole and Washington counties...................................................................... S. nitidula Mont.<br />
13. Ascospores 3-septate, (12-)15-24 × 3.5-6 µm; macroconidia filiform,<br />
3-7(-9)-septate;not common, throughout Florida ...................................... S. complanata (Fée) Mont.<br />
NOTES<br />
Strigula albicascens (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria albicascens Nyl., Lich. Nov. Zel. 129. 1888. Porina albicascens (Nyl.) Müll. Arg., Bull.<br />
Soc. Bot. Belgique 31: 38. 1892. Type. NEW ZEALAND. 1867, Knight 318 (H-Nyl 1534[A],<br />
lectotype selected here), 16 Jan 1882, Knight (H-Nyl 1983, syntype).<br />
This species known to me only from the type collections has whitish or grayish, endophloeodal thallus,<br />
unilocular ascomata with pale brown walls and 3-7-septate ascospores, 17-30 × 5-7 µm.<br />
Strigula albolinitum (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria albolinita Nyl., Lich. ins. guin. 38. 1889. Microglaena albolinita (Nyl.) Zahlbr., Cat. lich.<br />
146
univ. 1: 188. 1921. Phyllobathelium albolinitum (Nyl.) H. Mayrh., Biblioth. Lichenol. 26: 79.<br />
1987. Type. INSULA PRINCIPE. 1887, Fr. Quintas (H-Nyl 1588, holotype).<br />
See Mayrhofer (1987) for description. It is a little anomalous in either Phyllobathelium or Strigula. The<br />
immersed, pale walled ascomata fit best in Strigula but the large (ca. 40 × 16 µm) muriform macroconidia are<br />
apparently basally attached and have gelatinous caps at both ends more like Phyllobathelium.<br />
Strigula bahamensis (Riddle) R. C. Harris, comb. nov.<br />
Lithothelium bahamense Riddle in Britton & Millspaugh, Bahama Fl. 532. 1920. Type. BAHAMAS.<br />
Watling's Island: vicinity of Cockburn Town, on limestone, 13 Mar 1907, Britton & Millspaugh<br />
6127 (FH, holotype).<br />
Aptroot accepted this species in Lithothelium. He misread the the original description and cited Britton &<br />
Millspaugh 6129 as the type. This is of no importance since both 6127 and 6129 represent the same species<br />
of Strigula. The FH holotype has asci and ascospores of the Strigula-type and Britton & Millspaugh 6129 (NY)<br />
has two types of pycnidia, one with elliptical-oblong microconidia, the other with 5-7-septate macroconidia,<br />
both typical of Strigula, not Lithothelium, which has filiform microconidia and no known macroconidia. The<br />
reason that Aptroot did not see NY material is that it had already been filed in Strigula before he undertook his<br />
study and was not included in the loan for Lithothelium.<br />
Strigula bermudana (Tuck. ex Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria bermudana Tuck. ex Nyl., Sert. lich. trop. 43. 1891. Porina bermudana (Nyl.) Zahlbr.,<br />
Cat. lich. univ. 1: 367. 1922. Type. BERMUDA. 1881, Farlow 31 (H-Nyl 985, holotype).<br />
I had finished the notes for Strigula and quite contentedly synonymized Verrucaria bermudana with<br />
Strigula viridiseda as I had done in my thesis when I had a sudden attack of "What was I thinking?". Not only<br />
is the material on limestone (not bark) but the ascomata are considerably larger (0.5-1.0 mm vs. 0.3-0.4 mm).<br />
Therefore the saxicolous material is here recognized as a distinct species. All the problems are not solved,<br />
however, as the ascospore size intergrades with that of S. wilsonii. This problem requires an extensive, more<br />
sophisticated re-analysis of the data. The type and other tropical material have the ascospores uniseriately<br />
arranged. In Buck 27425 (NY) from Suwanee County the ascospores are biseriate and the paraphyses are<br />
more branched and may represent an undescribed species.<br />
Strigula cinefaciens (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria cinefaciens Nyl., Bull. Soc. Linn. Normandie sér. 2, 7: 180. 1873. Arthopyrenia<br />
cinefaciens (Nyl.) Zahlbr., Cat. lich. univ. 1: 320. 1921. Type. ANDAMAN ISLANDS. 1867, Kurz 40<br />
(H-Nyl 728, holotype; M, isotype).<br />
This corticolous species has a shiny gray thallus with black dots formed from a black hypothallus which<br />
here and there penetrates the thallus. This type of thallus is well known for the foliicolous taxa (previously<br />
placed in both Strigula and Phylloporis) but I believe this is the first corticolous representative. To my mind it is<br />
one more link binding together a broadly defined genus Strigula. The ascospores are 2-celled, ca. 14-15 × 4-5<br />
µm, tending to break into part spores.<br />
Strigula connivens R. C. Harris, sp. nov.<br />
Species unica ascocarpis compositis, 2-3 loculis ostiolo unico et ascosporis 1-septatis,<br />
7-10(-12) × 3-4.5 µm.<br />
Type. FLORIDA. Dade County: Cutler, 1898, Thaxter 19 (MSC, holotype).<br />
Thallus corticolous, greenish gray, well developed, endophloeodal. Ascomata compound, usually with<br />
two chambers, occasionally three, rarely uniloculate, complanate, 0.3-0.4 mm long, slightly less in width;<br />
chambers connected above, sharing a single ostiole; wall thinner or lacking below. Asci narrowly elliptical, 45-<br />
50 × 7-8 µm, with eight subbiseriate to biseriate spores. Ascospores, narrowly ovate, 2-celled, slightly<br />
constricted at septum, 7-10(-12) × 3-4.5 µm.<br />
147
If I had seen only a single collection of this I probably would have ignored it as an ascomatal aberration<br />
with sterile tissue or bark invading the centrum as is occasionally seen in Pyrenulaceae or Trypetheliaceae.<br />
However, it does seem to be a genuinely compound ascomatal type which expands the limits of variation in<br />
Strigula.<br />
Additional specimens. FLORIDA. Dade County: Cutler, Thaxter 17a (MICH), Snapper Hammock, 11 Dec<br />
1919, Britton & Britton 806 (FH, NY).<br />
Strigula fracticonidia R. C. Harris, sp. nov.<br />
Thallo corticola albo, ascosporis submuriformibus, 7 × 1-septatis, 25-27 × 8-9 µm et macroconidiis 3septatis,<br />
ca. 19 × 3 µm, in partibus rupentibus.<br />
Type. NEW ZEALAND. South Island: Marlborough, 9.5 km NW of Kaikoura, Mt. Fyffe S. F., 4 km SW of summit<br />
of Mt. Fyffe, along creek, alt. 180-330 m, on trunk of Melicytus ramiflora, 6 Feb 1981, Tibell 10790 (UPS,<br />
holotype).<br />
Thallus corticolous, white, not corticate, endophloeodal. Ascomata half immersed, subglobose, 0.25-0.4<br />
mm diam.; wall pale below with dark, poorly developed clypeus. Asci cylindrical, ca. 90 × 20 µm, with eight<br />
biseriate spores. Ascospores fusiform, submuriform, 7 × 1-septate, 25-27 × 8-9 µm. Macroconidia 3-septate,<br />
ca. 19 × 3 µm, fragmenting into four part spores. Microconidia oblong to ovate, 3-4.5 × 1.5-2 µm.<br />
Taxa with the ascospores fragmenting into part spores are well known in both foliicolous and corticolous<br />
species of Strigula but I do not know of any other species where the macroconidia break apart. The<br />
ascospores of S. fracticonidia are close to those of S. griseonitens which differs in having a corticate thallus<br />
and completely immersed ascomata. Strigula fracticonidia is known only from the type collection.<br />
Strigula griseonitens R. C. Harris, sp. nov.<br />
Thallo griseo corticato, ascomatis in cortice immersis parietibus hyalinis vel pallidis et ascosporis<br />
submuriformibus, 7(-8)-septatis, 3-6 cellulis longitudinaliter 1-2-septatis, 26-33 × 8-10 µm.<br />
Type. FLORIDA. Citrus County: Withlacoochee State Forest, along Florida Trail N to Brush Sink from Co. Rd.<br />
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,<br />
Harris 31825 (NY, holotype).<br />
Thallus corticolous, gray, ± shiny, with prosoplectenchymatous cortex, 10-30 µm thick. Ascomata<br />
immersed, subglobose, 0.3-0.4 mm diam; wall colorless below, expanded and streaked with pale brown<br />
around ostiole. Asci cylindrical, 100-130 × 15-20 µm, with eight biseriate spores. Ascospores submuriform,<br />
7(-8)-septate with 3-6 cells longitudinally 1-2-septate, 26-33 × 8-10 µm. Microconidia oblong, 4-5 × 1.5-2 µm.<br />
Macroconidia not found.<br />
Strigula griseonitens, named for the conspicuous thallus, is separated from S. submuriformis by larger,<br />
more divided ascospores, immersed ascomata and the shiny, corticate thallus. Muriform-spored species of<br />
Strigula seem rather trendy just now. I suppose as a result of more natural families and genera being<br />
recognized since some of them have been transferred from other genera where they were misplaced due to<br />
an artificial system based almost entirely on ascospore septation and color. Ascospores in Strigula are now<br />
allowed to range from 1-septate through submuriform to very strongly muriform. In addition to S.<br />
submuriformis, known since 1967, three recently described species occur on rock, S. porinoides Canals et al.<br />
with submuriform ascospores from Spain and Sicily, S. australiensis McCarthy and S. johnsonii McCarthy with<br />
more strongly muriform ascospores from Australia and New Zealand, and one on bark, S. lateralis Aptroot &<br />
v.d. Boom from Europe. Also on bark are S. fracticonidia from New Zealand, S. griseonitens, S. laceribracae<br />
from North America, and S. rostrata from Madagascar, newly described here. Strigula albolinita, S. obtecta<br />
148
and S. tagananae are transferred from other genera.<br />
Additional specimens. FLORIDA. Citrus County: as type, Harris 31810; Hillsborough County: Hillsborough<br />
River State Park, Rapids Trail along river, 28°09'N, 82°14'W, mixed harwoods, 3 Dec 1992, Harris 29647;<br />
Liberty County: Apalachicola National Forest, mature Taxodium dome 2.7 mi NW of Sumatra on Fla. Hwy.<br />
379, 28 Dec 1990, Harris 26104; Sarasota County: Myakka River State Park, 7 Mar 1937, Romer s.n.<br />
MISSISSIPPI. Jasper County: Bienville National Forest, Forest Service Rd. 506-3 at E edge of forest by small<br />
cemetery, 32°12'N, 89°13'W, ca. 105 m, dry oak woods on sand, 30 Sep 1992, Harris 28864 (all NY).<br />
Strigula hypothallina R. C. Harris, sp. nov.<br />
Thallo corticola nitido albo punctato, hypothallo nigro et ascosporis 1-septatis, 17-22 × 5-6 µm.<br />
Type. SOUTH CAROLINA. Berkeley County: Francis Marion National Forest, Guilliard Lake Campground, on Ilex,<br />
24 May 1967, Harris 3201-A (MSC, holotype).<br />
Thallus light brown, shiny, with scattered white dots; hypothallus black, visible at thallus margin or in<br />
section. Ascomata complanate to hemispherical, immersed, 0.5-0.8 mm diam.; wall black, sometimes<br />
merging with hypthallus, lacking below. Asci cylindrical, 90-120 × 9-10 µm, with eight uniseriate spores.<br />
Ascospores narrowly ovate, with ± pointed ends, 2-celled, slightly constricted at septum, 17-22 × 5-6 µm.<br />
Microconidia and macroconidia not found.<br />
Strigula cinefaciens is the only other corticolous species with a black hypothallus (not uncommon for<br />
foliicolous taxa). It is distinguished by black not white dotted thallus. Although known only from the type, I<br />
describe it to validate a name from my thesis and to expand the parallel character set shared by foliicolous<br />
and corticolous taxa.<br />
Strigula indutula (Nyl.) R. C. Harris, comb. nov.<br />
Verrucaria indutula Nyl., Lich. Nov. Zel. 129. 1888. Porina indutula (Nyl.) Müll. Arg., Bull. Herb.<br />
Boissier 2, App. 1: 92. 1894. Type. NEW ZEALAND. Knight (H-Nyl 1981, holotype).<br />
As Nylander pointed out this species is close to S. albicascens differing in better developed thallus and<br />
smaller ascospores, 5-7-septate, 16-20 × 4-5 µm.<br />
Strigula jamesii (Swinsc.) R. C. Harris<br />
I originally treated this species and our American material as S. affinis (Massal.) R. C. Harris but it was<br />
pointed out to me by Brian Coppins, I believe, that the two are distinct. At the time I accepted this but did not<br />
re-analyze the situation. I have now done so as far as possible with limited material. The ascospores are not<br />
much different if one believes the published ranges, S. jamesii 12-20(-22) × 2-3.5 µm (Purvis et al., 1992), S.<br />
affinis 12-20 × 3-6 µm (Roux & Bricaud, 1993). The macroconidia are also close, 13-17 × 3-3.5 µm for S.<br />
jamesii, 13-18 × 3-5 µm for S. affinis. The distinctions seem to lie elsewhere, S. affinis has a well developed,<br />
often rimose thallus and larger, more complanate ascomata while S. jamesii has a thin, continuous thallus with<br />
small, more distinctly hemisphaerical ascomata. The ascospore width given by Purvis et al. (1992) for S.<br />
jamesii does not agree with my observations which include the type. I would suggest these ascospore ranges<br />
for the two species, S. jamesii 13-17 × 4-5 µm and S. affinis 15-22 × 4.5-6 µm. According to my analysis<br />
American material is correctly placed in S. jamesii.<br />
Strigula laceribracae R. C. Harris, sp. nov.<br />
Thallo corticola albo, ascomatis immersis pariete hyalino vel pallido et ascosporis dense muriformibus,<br />
11-15 × 1-5-septatis, 35-45 × 13-17 µm.<br />
Type. GEORGIA. Murray County: Chattahoochee National Forest, Cohutta Wilderness, Tearbritches Trail from<br />
Forest Service Rd. 68 to Bald Mountain, 34°52'N, 84°38'W, 1100-1200 m, mesic oak woods, 22 Sep 1992,<br />
Harris 28170 (NY, holotype).<br />
149
Thallus corticolous, white, thin, ecorticate, mostly endophloeodal. Ascomata immersed, subglobose, 0.3-<br />
0.5 mm diam.; wall colorless below, expanded and streaked with brown around ostiole. Asci broadly ovate to<br />
obpyriform, 110-130 × 40-60 µm, with eight subbiseriately to irregularly arranged spores. Ascospores broadly<br />
fusiform, narrower at one end, densely muriform, 10-15 × 1-5-septate, 35-45 × 8-10 µm. Microconidia and<br />
macroconidia not found.<br />
Strigula laceribracae, named in "dog Latin" for the place of collection, is a very inconspicuous species as<br />
the white thallus is close to the color of the oak bark and the immersed ascomata are visible only as minute<br />
dark dots. The densely muriform ascospores are diagnostic among North American Strigulae.<br />
Additional specimen. LOUISIANA. East Baton Rouge Parish: Burden Research Plantation, Essen Lane,<br />
Baton Rouge, bottomland hardwood forest, 27 Mar 1980, Tucker 20899C (LSU).<br />
Strigula multipunctata (R. Sant.) R. C. Harris, comb. nov.<br />
Porina multipunctata R. Sant., Symb. Bot. Upsal. 12(1): 216. 1952. Phylloporis multipunctata (R.<br />
Sant.) V_zda, Folia Geobot. Phytotax. 19: 183. 1984. Type not seen.<br />
Strigula obducta (Müll. Arg.) R. C. Harris, comb. nov.<br />
Phylloporina obducta Müll. Arg., Flora 73: 198. 1890. Phylloporis obducta (Müll. Arg.) R. Sant.<br />
& Tibell, Austrobaileya 2: 539. 1988. Type not seen.<br />
Strigula obtecta (Vainio) R. C. Harris, comb. nov.<br />
Thelenella obtecta Vainio, Étud. class. lich. Brésil 2: 218. 1890. Polyblastiopsis obtecta (Vainio)<br />
Zahlbr., Cat. lich. univ. 1: 351. 1922. Phyllobathelium obtectum (Vainio) H. Mayrh., Biblioth.<br />
Lichenol. 26: 82. 1987. Type. BRAZIL. Minas Gerais: Sitio, 1885, Vainio, Lich. Brasil. exs. 718<br />
(BM, isotype).<br />
This species has strongly muriform ascospores. See Mayrhofer (1987) for description and illustration.<br />
Strigula phaea (Ach.) R. C. Harris<br />
Porina mundula Müll. Arg., Bot. Jahrb. Syst. 6: 402. 1885. Type. CUBA. Wright, Verr. Cub. 212<br />
(NY, isotype).<br />
Some forms of this corticolous species (e.g., the type of Porina mundula) have a very similar aspect to<br />
taxa assigned to Phylloporis with conical ascomata and weak basal fringe. I suppose some would be happy to<br />
see all corticolous and saxicolous Strigula placed in Phylloporis.<br />
Strigula phyllogena (Müll. Arg.) R. C. Harris, comb. nov.<br />
Porina phyllogena Müll. Arg., Flora 66: 335. 1883. Phylloporis phyllogena (Müll. Arg.) Clements,<br />
Genera of Fungi 173. 1909. Type not seen.<br />
Strigula platypoda (Müll. Arg.) R. C. Harris, comb. nov.<br />
Porina platypoda Müll. Arg., Flora 66: 335. 1883. Phylloporis platypoda (Müll. Arg.) V_zda, Folia<br />
Geobot. Phytotax. 19: 185. 1984. Type not seen.<br />
Strigula rostrata R. C. Harris & Aptroot, sp. nov.<br />
Species propria corticola macropycnidiis rostratis, macroconidiis submuriformibus 15-18 × 7-8 µm et<br />
ascosporis submuriformibus 27-30 × 7-8 µm.<br />
Type. MADAGASCAR. Tamatave, Perinet (=Andasibe), primary montane forest, 950 m, 48°16'E, 18°56'S, 10<br />
May 1984, Aptroot 13397 & Hensen (hb. Aptroot, holotype).<br />
Thallus corticolous, white, not corticate, mostly endophloeodal. Ascomata flattened conical, sessile, ca.<br />
150
1.0 mm diam.; wall black, thinner below. Asci cylindrical ca. 90-105 × 11-13 µm, initially with 8 uniseriate<br />
spores but several usually aborting (probably not typical?). Ascospores submuriform, 4-5 × 1-septate, 27-30 ×<br />
7-8 µm. Macropycnidia with a short, thick beak, 0.3-0.4 mm diam. basally, 0.2-0.3 mm high. Macroconidia<br />
often extruded as a cirrus, submuriform, 15-18 × 7-8 µm, basally attached with a gelatinous basal cap.<br />
Microconida oblong to ovate, 3-4 × 1-2 µm.<br />
This is rather distant from the subject of Florida lichens but it is such an unusual species (Madagascar<br />
strikes again) that I could not resist. Long-beaked macropycnidia occur in Anisomeridium but this is the first<br />
known occurrence in Strigula. The basally attached, submuriform macroconidia are also unusual. It may turn<br />
out that only the transversely septate macroconidia are eccetrically attached.<br />
Additional specimen (macropycnidia only). MADAGASCAR. As type, Aptroot 13395 & Hensen (hb. Aptroot).<br />
Strigula smaragdula Fr.: Fr.<br />
Since it is a sanctioned name, S. smaragdula Fr. is conserved over the well known S. elegans (Fée) Müll.<br />
Arg.<br />
Strigula subelegans Vainio s. lat.<br />
The name is used in a broad sense since the only Florida collection has rather small ascospores, 11-15 ×<br />
4-4.5 µm. Santesson (1952) gave 17-25 × 4-7 µm. The material otherwise seems a reasonable match for<br />
Santesson's illustration and has a bluish tinge as emphasized by Lücking (1992). In view of my opinions on<br />
species limits in foliicolous Strigula (see family discussion) I am in no position to suggest this is an<br />
undescribed species.<br />
New to North America. FLORIDA. Dade County: Brickell Hammock, on leaves of Gymnanthes lucida, 16<br />
Dec 1919, Britton & Britton 690 (NY).<br />
Strigula sychnogonioides (Nitschke) R. C. Harris<br />
This was described in the monotypic genus Geisleria Nitschke. Largely on the basis of apically free<br />
physes, lacking information on conidial types, I included it as a synonym of Strigula. On re-examination of old<br />
material I would exclude it from Strigula. Ernst (Herzogia 9: 321-337. 1993) provided a detailed discussion of<br />
its ecology but no help at all as to its taxonomic position even though she had abundant fresh material. My<br />
first guess having proved wrong I venture a second (probably no better) that Geisleria sychnogonioides<br />
Nitschke is an algal parasite belonging somewhere in or near the Clypeosphaeriaceae in the Xylariales s. lat.<br />
Strigula tagananae (Harmand) R. C. Harris, comb. nov.<br />
Verrucaria tagananae Harmand in Pitard & Harmand, Bull. Soc. Bot. Fr. 58(Mém. 22): 68. 1911.<br />
Polyblastiopsis tagananae (Harmand) Zahlbr., Cat. lich. univ. 1: 352. 1922. Type. CANARY<br />
ISLANDS. Tenerife: Vueltas de Taganana, 500 m, sur Ilex canariensis, Pitard 1691 (DUKE,<br />
holotype).<br />
Verrucaria tagananae is placed in Strigula with some hesitation. The photobiont, apically free physes,<br />
asci and microconidia are in agreement. However, the ostiole is lateral, a feature otherwise unknown in the<br />
family. It is known to me only from the type. Perhaps additional material will provide better evidence as to its<br />
placement.<br />
Strigula viridiseda (Nyl.) R. C. Harris<br />
This is at the center of a complex of poorly resolved taxa. In 1975 I included saxicolous material now<br />
removed to S. bermudana. The ascospore size range overlaps that of S. americana and determination is<br />
often difficult. The possibility exists that the recognition of a third species intermediate in ascospore and<br />
macroconidia size might resolve the problem. Also there is a collection from Ontario (Wong 3811, CANL) with<br />
ascospores, 12-16 × 4-5 µm, in the range of S. viridiseda but so far out of geographic range that I am<br />
undecided as to its disposition without further study.<br />
Strigula viridis (Lücking) R. C. Harris, comb. nov.<br />
151
Phylloporis viridis Lücking, Nova Hedwigia 52: 276. 1991. Type not seen.<br />
THELENELLACEAE O. Eriksson ex H. Mayrh.<br />
Biblioth. Lichenol. 26: 16. 1987. Type. Thelenella Nyl.<br />
Microglaenaceae Servít, Studia Bot. _ech. 7: 108. 1946 [illegitimate, based on illegitimate name]. Type.<br />
Microglaena Körber (1855) [non Microglena Ehrenberg, 1832] (= Thelenella Nyl.).<br />
Aspidotheliaceae Räsänen ex J. C. David & D. Hawksw., Systema Ascomycetum 10: 13. 1991. Type.<br />
Aspidothelium Vainio<br />
The features which unite the members of this family are fissitunicate? asci with an ocular chamber and a<br />
cone shaped plug or ring above it in the ascus tip staining with phloxine/KOH, quite irregular, short celled,<br />
branched and interconnected physes, periphysoids, ascomatal wall often not melanized and a<br />
chlorococcalean photobiont or non-lichenized. Conidia seem unusually hard to find but Thelenella and<br />
Celothelium have arcuate, ± filiform microconidia. I have not found microconidia for "Aspidothelium" but<br />
Santesson (1952) reported oblong, 3-4 × 1 µm conidia for A. cinerascens. I have found elliptic oblong<br />
macroconidia in one specimen of "Aspidothelium" and one of Chromatochlamys but more occurrences are<br />
needed to be convincing. The ascus type is similar to that of the Requienellaceae but I can see no other<br />
features in common and in fact see no close relationships to any lichen-forming loculoascomycete family. The<br />
unusual ascus and photobiont suggest the Thelenellaceae represent an independent (ancient?) adoption of<br />
the lichen lifestyle.<br />
Three genera were assigned to the Thelenellaceae by Mayrhofer. Due to lack of adequte material for<br />
study I am temporarily recognizing Chromatochlamys as distinct. Julella is not lichen-forming, has a different<br />
ascus type and is clearly a member of the Arthopyreniaceae. Aspidothelium is a synonym of Thelenella.<br />
Aptroot and Sipman (1993) assigned Musaespora Aptroot & Sipman to the Aspidotheliaceae (correctly<br />
realizing that the Aspidotheliaceae hardly differed from the Thelenellaceae). However, Musaespora has a<br />
trentepohlioid photobiont, a different ascus type and ellipsoid to globose microconidia. Ascospore type and<br />
microconidia suggest placement in the Monoblastiaceae and I tentatively assign it there pending further study<br />
as I have only limited material at present. Celothelium is placed here provisionally, perhaps more out of<br />
expedience than anything else. Although the ascomatal type is very different from Thelenella, the ascus has a<br />
conical ring staining with phloxine/KOH, arcuate microconidia (illustrated by Aguirre, 1991, for C. stenobelum)<br />
and thin, irregular, short-celled physes. The photobiont is also a problem for me but Celothelium may not be<br />
lichen-forming which eases my conscience somewhat. (The Dade County collection of Celothelium<br />
aciculiferum is on lignum and no algae are present.) The key to genera below is artificial as to separating<br />
Thelenella and Chromatochlamys since I cannot at this time come up with any natural characters.<br />
1. Growing on bark, rock or leaves ............................................................................................................... 2<br />
2. Ascospores elliptical to fusiform, 3-septate to muriform ........................................................Thelenella<br />
2. Ascospores filiform, transversely septate........................................................................... Celothelium<br />
1. Growing on mosses and plant remains........................................................................ [Chromatochlamys]<br />
CELOTHELIUM Massal.<br />
Type. Verrucaria socialis sensu Montagne. (Celothelium leprieurii Trevisan?)<br />
?Trichotrema Clements, Genera of Fungi 41. 1909. Type. Pleurothrema trichosporum Müll. Arg.<br />
Celothelium was resurrected by Aguirre (1991). Her treatment is not up to the level of that for<br />
Leptorhaphis. She proposed to typify Verrucaria socialis Zenker (1829) with a Leprieur collection. Leprieur<br />
collected in French Guiana from 1830-1869 so that this is not appropriate. However, Massalongo in the<br />
original description designated the type species as "Verrucaria socialis (Montagn.) Zenk. ?". (Montagne (1851)<br />
used the name Verrucaria socialis Zenker for some Leprieur collections but with no description.) I interpret<br />
152
this to mean Massalongo intended (in modern terms)Verrucaria socialis sensu Montagne. This is supported<br />
by Trevisan (Flora 44: 25. 1861) using the name (nomen novum?) Celothelium leprieurii in trying to sort out<br />
the confusion of names involved with Verrucaria socialis which he says can only be solved by examination of<br />
Zenker's original specimen. This view is also supported by the fact that the combination Celothelium socialis<br />
was not used by Massalongo. I suggest that Celothelium should be typified with Verrucaria socialis sensu<br />
Montagne (not Zenker) based on a specimen in Massalongo's herbarium. If such does not exist, then<br />
Aguirre's lectotype fortuitously turns out to be acceptable (not for Zenker's name as she proposed but for what<br />
probably should be called Celothelium leprieurii Trevisan). Because of inadequate material Aguirre declined<br />
treating C. socialis/leprieurii and I have seen no material. (The epithet is younger than aciculiferum but could<br />
replace cinchonarum.)<br />
Trichotrema is placed in synonymy with question since the holotype specimen of the generitype has been<br />
destroyed by Indian lichenologists and failing the discovery of an isotype the application of the name will have<br />
to fixed through neotypification.<br />
Aguirre recognized three species in the Neotropics. I would recognize only two. The ascospore<br />
character proposed to separate C. dominicanum (Vainio) Aguirre in the key is not supported by the data in her<br />
descriptions. I consider it a synonym of C. cinchonarum. Integration of the species of Trichotrema into<br />
Celothelium will probably add to the number of species recognized. I have specimens at NY of C. aciculiferum<br />
from Florida, Cuba, Dominican Republic, Jamaica, Puerto Rico, and Guyana and C. cinchonarum from Cuba,<br />
Jamaica, Puerto Rico, Trinidad and Ecuador.<br />
1. Ascomata solitary or several covered by a common clypeus; ostioles<br />
eccentric; asci strongly slanted to almost horizontally arranged;<br />
ascospores ca. 65-100 × 2 µm; microconidia filiform, often arcuate,<br />
10-17 × 0.5 µm; Collier and Dade counties ................................................. C. aciculiferum (Nyl.) Vainio<br />
1. Ascomata solitary or fused laterally (± melanothecoid); ostioles apical;<br />
asci arranged ± vertically; ascospores ca. 80-110 × 2 µm; microconidia<br />
filiform, often arcuate, 10-13 × 0.5 µm ............................................... [C. cinchonarum (Müll. Arg.) Vainio]<br />
NOTES<br />
Celothelium aciculiferum (Nyl.) Vainio<br />
New to North America. FLORIDA. Collier County: Collier-Seminole State Park, just W of boat basin,<br />
29°59'N, 81°35'W, mangroves at edge of salt marsh, 7 Dec 1992, Harris 30060; Dade County: Everglades<br />
National Park, vicinity of Pinelands Trail, 2 Jan 1976, Buck B840 (both NY).<br />
CHROMATOCHLAMYS Trevisan<br />
I have not had sufficient material to make thorough studies to resolve the status of Chromatochlamys.<br />
Preliminarily the asci seem to show the same staining characteristics as Thelenella. Mayrhofer separated<br />
them on presence (Chromatochlamys) or absence (Thelenella) of an ocular chamber. However, fresh<br />
specimens of most Thelenella species show an obvious ocular chamber as do older specimens if stained and<br />
treated with KOH. Thelenella was originally said to lack perphysoids but this was later changed so that there<br />
is no difference on this score. The physes seem a little coarser and contain non-staining, refractive oil<br />
droplets. Is this an indication that generic status is warranted? Further studies are needed.<br />
There is only a single species in North America, C. muscorum (Fr.) H. Mayrh. & Poelt with two varieties,<br />
muscorum and octosporum (Nyl.) H. Mayrh. & Poelt. I have seen var. muscorum with 2-4 spores per ascus<br />
from scattered localities (Colorado, Minnesota, North Carolina). The var. octosporum is included in Egan<br />
(1987) based on a personal communication from Hawksworth that Pyrenidium octosporum Looman from<br />
Saskatchewan is this taxon.<br />
153
THELENELLA Nyl.<br />
Mém. Soc. Sci. Nat. Cherbourg 3: 193. 1855. Type. T. modesta (Nyl.) Nyl.<br />
Microglaena Körber, Syst. Lich. German. 388. 1855. Type. M. wallrothiana Körber (= Thelenella modesta<br />
(Nyl.) Trevisan).<br />
Luykenia Trevisan, Conspect. Verruc. 19. 1860. Type. L. modesta (Nyl.) Trevisan<br />
Aspidothelium Vainio, Étud. class. lich. Brésil 2: 189. 1890. Type. A. cinerascens Vainio.<br />
Aspidopyrenium Vainio, Étud. class. lich. Brésil 2: 190. 1890. Type. A. insigne Vainio (= Aspidothelium<br />
fugiens (Müll. Arg.) R. Sant. fide Santesson, 1952).<br />
Secolegiella (Müll. Arg.) Müll. Arg., Hedwigia 34: 205. 1896. Type. Lecania fugiens Müll. Arg.<br />
Patellonectria Spegazzini, Bol. Acad. Ci. Córdoba 23: 477. 1919. Type. P. puiggari Spegazzini (=<br />
Aspidothelium cinerascens Vainio fide Santesson, 1952).<br />
Porinopsis Malme, Ark. Bot. 22A(6): 3. 1928. Type. P. geminipara Malme<br />
Thelenella and Aspidothelium are identical in all important characters as noted above, oddly staining<br />
ascus, fleshy ascoma with periphysoids, chlorococcalean photobiont, etc. Aspidothelium may have ascomata<br />
ornamented with a apical disk, warts or setae but they may also be smooth as in Thelenella. Also, it would not<br />
seem to take much of an expansion to go from the thick apical tissue present in most Thelenella ascomata to<br />
achieve apical warts or an apical disk. Some taxa of Aspidothelium have mainly transversely septate spores<br />
but Santesson (1952) noted that they may have one-few longitudinal septa. Both commonly have the area<br />
around the ostiole tinted greenish. Both often have much of the centrum taken up with spent asci. Both can<br />
be foliicolous or corticolous although Aspidothelium is not known from rock. Overall, I feel the similarities are<br />
fundamental, the differences unimportant and therefore synonymize them.<br />
KEY TO NORTH AMERICAN TAXA<br />
(Some data taken from Mayrhofer, 1987)<br />
1. Ascospores colorless ................................................................................................................................ 2<br />
2. Growing on bark or leaves .................................................................................................................... 3<br />
3. Growing on bark ................................................................................................................................ 4<br />
4. Ascospores muriform .................................................................................................................... 5<br />
5. Ascospores basically 8/ascus ................................................................................................... 6<br />
6. Ascospores 11-17 µm wide................................................................................................... 7<br />
7. Asci obclavate to pyriform, broadest toward the base;<br />
ascospores 30-38 × 13-15 µm, 7-9 × 2-4-septate;<br />
Dade and Seminole counties ............................................................T. rappii R. C. Harris<br />
7. Asci cylindrical or broadest in middle; ascospores 25-42 × 11-17 µm,<br />
7-9 × 2-3-septate; Europe, Califormia, Minnesota, Australia ..............................................<br />
........................................................................................................[T. modesta (Nyl.) Nyl.]<br />
6. Ascospores 7.5-11 µm wide.................................................................................................. 8<br />
8. Ascospores 7-11 × 1(-2)-septate, 24-34 × 7.5-10 µm;<br />
Michigan, Minnesota ................................................... [T. pertusariella (Nyl.) Vainio s. lat.]<br />
8. Ascospores 6-7 × 1-2-septate, 20-30 × 7-11 µm;<br />
California, Australia ..........................................[T. sychnogonioides (Zahlbr.) R. C. Harris]<br />
154
5. Ascospores basically 4/ascus, 11-13 × 1-2-septate, with large cells,<br />
53-63 × 16-18 µm; ascomata pallid to dark brown, ± hemispherical,<br />
smooth; Collier County.................................................... T. cinerascens (Vainio) R. C. Harris<br />
4. Ascospores transversely 11-17-septate, 2-4/ascus, 75-100 × 17-20 µm;<br />
ascomata pallid to brownish, smooth; on branches; locally common,<br />
Collier County....................................................................... T. geminipara (Malme) R. C. Harris<br />
3. Growing on leaves; ascospores 8/ascus, transversely septate, 50-90 × 12-21 µm<br />
(Santesson, 1952); in Egan (1987) as fide Tucker, not seen..............................................................<br />
.........................................................................................................[A. fugiens (Müll. Arg.) R. Sant.]<br />
2. Growing on rock .................................................................................................................................... 9<br />
9. Ascospores muriform ...................................................................................................................... 10<br />
10. Ascomata without dark brown clypeus (involucrellum) ............................................................. 11<br />
11. Ascospores oblong-ellipsoidal, ± tapered at one end,<br />
over 20 µm long................................................................................................................... 12<br />
12. Ascospores medium sized, 20-36 × 8-13 µm................................................................ 13<br />
13. Thallus thin, continuous to ± rimose........................................................................ 14<br />
14. Ascospores 24-32 × 9-13 µm, 5-7 × 2-3-septate............................................... 15<br />
15. Thallus light brown, olive brown to olive green;<br />
ascospores (24-)27-32 × 9-13 µm, 6-7 × 2-septate;<br />
New Jersey, Oklahoma, Neotropics, Africa,<br />
China, Australia ..........................................[T. brasiliensis (Müll. Arg.) Vainio]<br />
15. Thallus pale ocher to yellow brown;<br />
ascospores 24-30 × 9-13 µm, 5-7 × 2-3-septate;<br />
Arctic ............................................................[T. sordidula (Th. Fr.) H. Mayrh.]<br />
14. Ascospores 20-27 × 8-11 µm, 6-9 × 1-2-septate;<br />
Louisiana, Puerto Rico ................................................ [T. sastreana R. C. Harris]<br />
13. Thallus thick, rimose-areolate; ascospores 24-36 × 9-13 µm,<br />
6-8 × (1-)2-3-septate; California, Macaronesia, SW Europe.......................................<br />
......................................................................................[T. inductula (Nyl.) H. Mayrh.]<br />
12. Ascospores larger, 30-45 × 12-19 µm, 7-9 × 3-septate;<br />
Georgia, Neotropics, South Africa, Australia, New Zealand..............................................<br />
..............................................................................................[T. luridella (Nyl.) H. Mayrh.]<br />
11. Ascospores broadly ellipsoidal, small, to 20 × 8-11 µm,<br />
6-7 × 2-3-septate; New York ..............................................................[T. humilis R. C. Harris]<br />
10. Ascomata with an open dark brown clypeus (involucrellum);<br />
ascospores 35-37 × 14-19 µm, 6-8 × (1-)2-3-septate;<br />
California, Baja California.............................................................................[T. weberi H. Mayrh.]<br />
155
9. Ascospores 4-celled, 18-23 × 6.5-7.5 µm; thallus scanty; on pebble;<br />
Puerto Rico........................................................................................... [Thelenella sp. Buck 18291A]<br />
1. Ascospores soon brown, 25-40 × 10-17 µm, 6-8 × 2-3-septate; on bark;<br />
California ..................................................................................................... [T. hassei (Zahlbr.) H. Mayrh.]<br />
NOTES<br />
Thelenella brasiliensis (Müll. Arg.) Vainio<br />
I have placed some specimens with smaller ascospores in a new species, T. sastreana, although its<br />
range overlaps that given by Mayrhofer. In doing this revision I have gotten the impression that he must have<br />
measured immature ascospores and thus his ranges go too low. The Oklahoma collection seems typical but<br />
the New Jersey collection is ecologically anomalous, semiaquatic, growing with Ionaspis lacustris (With.)<br />
Lutzoni. Thelenella inductula "differs" in thicker thallus. Further study should be undertaken to determine if it<br />
is distinct from brasiliensis.<br />
New to North America. NEW JERSEY. Hunterdon County: [Mitchell property between Upper Creek and<br />
Pine Hill Roads,] on submerged shale, Nov 1987, Aptroot 214. OKALAHOMA. Cherokee County: along Terapin<br />
Creek ca. 6 mi N of St. Rd. 82 on St. Rd. 100, 23 Apr 1988, Harris 21384 (both NY).<br />
Thelenella cinerascens (Vainio) R. C. Harris, comb. nov.<br />
Aspidothelium cinerascens Vainio, Étud. class. lich. Brésil 2: 189. 1890. Type. BRAZIL. Rio de<br />
Janeiro, 1885, Vainio, Lich. Bras. 215 (not seen).<br />
The North American material is somewhat anomalous in having the cells of the ascospores rather large.<br />
Additional Florida specimens and more comparative specimens from the Neotropics may show that North<br />
American material should be given taxonomic recognition.<br />
New to North America. FLORIDA. Collier County: Fakahatchee Strand State Preserve, 1.2 mi from N<br />
entrance of W. J. Janes Memorial Scenic Drive, open Taxodium swamp, 6 Dec 1992, Harris 29959-A, Collier-<br />
Seminole State Park, just W of boat basin, mangroves at edge of salt marsh, 7 Dec 1992, Harris 30049-G,<br />
30067-A (all NY).<br />
Thelenella cuyabense Malme<br />
I have not seen material of this species from Brazil. McCarthy (Lichenologist 27: 345. 1995) has<br />
transferred it to Aspidothelium. It is possible that it is an older name for T. rappii but time has not allowed<br />
examination of the type of T. cuyabense.<br />
Thelenella fugiens (Müll. Arg.) R. C. Harris, comb. nov.<br />
Lecania fugiens Müll. Arg., Lich. epiphylli novi 3. 1890. Type. BRAZIL. Apiahy, 1881-82, Puiggari<br />
250 (not seen).<br />
Thelenella geminipara (Malme) R. C. Harris, comb. nov.<br />
Porinopsis geminipara Malme, Ark. Bot. 22A(6): 3. 1928. Lectotype (Santesson, 1952).<br />
PARAGUAY. Colonia Risso prope Río Apa, Malme, Regn. Lich. 1856 (not seen).<br />
This name is applied to Florida material in a broad sense. According to Santesson (1952) the ascomata<br />
may be expanded at the apex, warted or smooth. The Florida collections have smooth, rounded ascomata. I<br />
have seen almost no comparative material. A specimen from Ecuador has a broad disk-like expansion but the<br />
ascospores are quite different from Florida specimens. Once again there is the possibility that North American<br />
material may represent an undescribed species. It seems locally abundant in the western Everglades.<br />
New to North America. FLORIDA. Collier County: Fakahatchee Strand State Preserve, Royal Palm<br />
Hammock, on Ilex, 6 Dec 1992, Harris 29929, Collier-Seminole State Park, just W of boat basin, at edge of<br />
salt marsh, 7 Dec 1992, Harris 30050, Big Cypress National Preserve, Oasis Ranger Station along U.S. Hwy.<br />
41, along Florida Trail N of hwy., Taxodium swamp, 9 Dec 1992, Harris 30175 (all NY).<br />
156
Thelenella humilis R. C. Harris, sp. nov.<br />
Thallo epilithico, disperso-areolato, glauco, ascomatis unicis in areolis immersis et ascosporis ellipsoideis<br />
vel late ellipsoideis, 18-20 × 8-11 µm.<br />
Type. NEW YORK. Warren County: Crane Mountain, 43°32'N, 73°57'45"W, 2100-3100 ft, on boulder in trail, 18<br />
Sep 1993, Harris 30577 (NY, holotype).<br />
Thallus epilithic, dispersed areolate with irregular to orbicular areoles, ca. 0.5-1.0 mm diam., gray green<br />
when wet, paler dry. Ascomata subglobose, immersed singly in areoles with only blackish tip visible, ca. 0.25<br />
mm diam. ( fertile areoles ca 1.0 mm diam.); wall colorless except greenish around ostiole. Asci cylindrical, ca.<br />
120 × 17 µm, with 8 uniseriate ascospores. Ascospores ellipsoidal to broadly ellipsoidal, 6-7 × 2-3-septate,<br />
18-20 × 8-11 µm; conidia not found.<br />
The epithet is given as it is an inconspicuous species with small areoles and small ascospores.<br />
Thelenella humilis is distinctive in the greenish thallus color (mostly shades of brown or whitish in other<br />
species) and in the small, smoothly rounded ellipsoidal ascospores. It is known only from the type collection.<br />
Thelenella luridella (Nyl.) H. Mayrh.<br />
This species may prove to be more widely distributed in acid rock glades in the Southeast.<br />
New to North America. GEORGIA. Coffee County: Broxton Rocks Ecological Preserve, 9 mi NE of Broxton,<br />
3 mi S of Ocmulgee River, 31°44'N, 82°45'W, on sandstone near falls, 16-17 Dec 1993, Harris 32595, 32596;<br />
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<br />
Ocmulgee River, 31°45'N, 82°43'W, dissected sandstone outcrop, 5 Feb 1995, Buck 27509 (all NY).<br />
Thelenella pertusariella (Nyl.) Vainio s. lat.<br />
The North American specimens have more septate ascospores than reported for European material.<br />
However, the ascospore size fits and as I have seen nothing from Europe for comparison the exact status of<br />
American specimens is left for future study.<br />
New to North America. MICHIGAN. Baraga County: moist woods along Sturgeon Rver ca. 0.25 mi N of<br />
campground, 5 mi NE of Sidnaw, on Acer negundo, 6 Sep 1972, Harris 8115 (NY). MINNESOTA. Cass County:<br />
4 mi S of Whipholt (11 mi SE of Walker), on quaking aspen, 14 Aug 1976, Wetmore 26513 (mixed with and<br />
distributed as Strangospora pinicola, NY).<br />
Thelenella rappii R. C. Harris, sp. nov.<br />
Similis T. modestae sed ascis obclavatis vel pyriformibus, 85-110 × 30-35 µm et ascosporis ovoideis, 7-9<br />
× 2-4-septatis, 30-38 × 13-15 µm.<br />
Type. FLORIDA. Seminole County: Sanford, on Hicoria, Apr 1916, Rapp (FLAS F29437, holotype; FH, isotype,<br />
both annotated by G. K. Merrill as Clathroporina obtecta sp. nov.).<br />
Thallus epiphloeodal, whitish or olivaceous, thin, continuous. Ascomata emergent, covered by thallus,<br />
subglobose, 0.25-0.4 mm diam.; wall pale or darkened above. Asci obclavate to pyriform, 85-110 × 30-35 µm,<br />
with eight irregularly arranged ascospores. Ascospores ovoid, 7-9 × 2-4-septate, 30-38 × 13-15 µm;<br />
microconidia not found.<br />
Thelenella rappi is close to T. modesta differing in ascus shape (obclavate vs. ± cylindrical) and<br />
distribution (tropical vs. temperate). It is possible a larger suite of specimens will lead to synonymization.<br />
Additional specimen. FLORIDA. Dade County: Everglades National Park, just N of Paurotis Pond, 11 May<br />
1967, Harris 2650 (NY).<br />
157
Thelenella sastreana R. C. Harris, sp. nov.<br />
Thelenellae brasiliensis similis ascosporis minoribus, 7-8 × 1-2-septatis, 24-27 × 9-11 µm.<br />
Type. PUERTO RICO. Distr. Mayagüez: Maricao State Forest, near Maricao Recreation Area, Camino del<br />
Guama and along Hwy. 120 between Kms 16 and 17, ca. 1700 m, relatively recent roadcut and roadside, 27<br />
May 1989, Buck 17213 (NY, holotype).<br />
Thallus thin, continuous, tan. Ascomatal warts ± complanate, orbicular, 0.3-0.5 mm diam., blackish<br />
around ostiole. Ascomata subglobose, 0.2-0.3 mm diam.; wall colorless except greenish around ostiole. Asci<br />
subcylindrical, slightly widened near middle, 85-130 × 15-20 µm, with 8 mostly biseriate ascospores.<br />
Ascospores colorless, with ± pointed ends, tapered to lower end, 6-9 × 1-2-septate, 20-27 × 8-11 µm; conidia<br />
not found.<br />
This species is named for Dra. Inés Sastre-De Jesús who has facilitated our work in Puerto Rico and is a<br />
most amiable companion in the field. Thelenella sastreana is close to T. brasiliensis. The Caribbean<br />
population seems to have consistently smaller ascospores. The collection sites in Puerto Rico may indicate<br />
this is something of a pioneer species able to take advantage of anthropogenic disturbance. The collection<br />
from Louisiana differs from the type in longer asci and in having little visible thallus.<br />
Additional specimens. LOUISIANA. Natchitoches Parish: Longleaf Trail Vista, ca. 35 mi WNW of<br />
Alexandria, sandstone cliffs, 28 May 1976, Harris 11445-A. PUERTO RICO. As type, Buck 17221; Distr.<br />
Ponce/Arecibo: SW slope of Cerro Morales, 1700-2100 ft., along road off Hwy. 531 at ca. Km 3.3, moist<br />
roadside, 2 Jun 1989, Buck 17384 (all NY).<br />
Thelenella sychnogonioides (Zahlbr.) R. C. Harris, comb. nov.<br />
Microglaena sychnogonioides Zahlbr., Beih. Bot. Centralbl. 13: 151. 1902. Type. CALIFORNIA.<br />
Santa Monica Mts., on Quercus agrifolia, Aug 1900, Hasse 776 (W-7057, lectotype, selected<br />
here).<br />
Thelenella harrisii H. Mayrh., Biblioth. Lichenol. 26: 36. 1987. Type. CALIFORNIA. Santa Monica<br />
Range, on Quercus agrifolia, 1907, Hasse 800 (W, holotype, not seen), syn nov.<br />
The lectotype above could be called a holotype as it was annotated by Zahlbruckner but since there is<br />
some confusion, I have chosen to designate a lectotype. Mayrhofer (1987) placed M. sychnogonioides in<br />
synonymy with Thelenella hassei choosing as "holotype" a specimen which does not fit Zahlbruckner's<br />
protologue, "sporis ... hyalinis ... loculis subcubicis, verticalibus 7-8, horizontalibus 2 ... 20-34µ longis et 9-11 µ<br />
latis.". (T. hassei has larger, brown ascospores.) The reasons for this seem to be twofold. The specimen<br />
designated as the lectotype is numbered "776" while that cited in the protologue is "778". However,<br />
Zahlbruckner's handwriting may have been bad, the printer had a dyslexic typesetter, or no one bothered to<br />
proofread the text, since miscitations of collection numbers are common in Zahlbruckner's work. (There are<br />
three or four in a single paper on Rapp's Florida lichens.) Since the specimen was annoted by Zahlbruckner<br />
and matches the protologue, I don't see the number confusion as a problem. (Nor did Mayrhofer who cited<br />
776 at FH as an "isotype" of 778 at W.) Material distributed by Hasse as 776 (not seen by Zahlbruckner) is T.<br />
hassei. It may be noted here that the numbers on Hasse collections are not collection numbers but species or<br />
herbarium numbers so that specimens collected at different places and times will carry the same number. The<br />
other reason for Mayrhofer's treatment is that the lectotype specimen probably was not available to him. It<br />
was included in a loan to Imshaug and his loans have only very recently been returned. It is something of a<br />
relief to be able to dispose of T. harrisii as it is rather an embarassment to have a Californian species named<br />
for one who professes to beleive that North America ends at the Mississippi River.<br />
Thelenella trichothelioides (Sérusiaux & V_zda) R. C. Harris, comb. nov.<br />
Aspidothelium trichotheliodes Sérusiaux & V_zda, Lejeunia n.s. 90: 4. 1978. Type. ZAIRE. Dist.<br />
158
Afro-montagnard Centre-oriental: env. du km 42 de la route Bukavu-Walikale, alt. env. 2300 m,<br />
épiphylle sur Connarus longestipitatus, Lambinon 71/1178 (LG, holotype, not seen).<br />
Thelenella sp. Buck 18291A<br />
Although known only from a small collection from Puerto Rico, this species is included as it extends the<br />
limits of Thelenella. The ascospores are transversely 3-septate and the thallus is scanty but in other respects<br />
it seems to be a typical Thelenella. The ascospores although small overlap the lower end of the ranges of<br />
several muriform-spored species.<br />
PUERTO RICO. Distr. Ponce: Caribbean National Forest, Toro Negro Division, along Hwy. 143 at Km 20.5,<br />
ca. 100 m, on roadside pebble, 4 Nov 1990, Buck 18291A (NY).<br />
TRICHOTHELIALES<br />
TRICHOTHELIACEAE (Müll. Arg.) Bitter & Schilling<br />
Since my brush with this family in my student days I have tended to avoid it being aware of the problems<br />
involved with defining species. Recently it has received considerable attention. A family treatment for the<br />
Guianas (Aptroot & Sipman, Flora of the Guianas, Series E, fasc. 2, 1993), saxicolous Southern Hemisphere<br />
taxa (McCarthy, Biblioth. Lichenol. 52, 1992), Clathroporina (McCarthy, Lichenologist 27: 321-350. 1995) and<br />
a revision of the genera (Hafellner & Kalb, Biblioth. Lichenol. 57: 161-186. 1995).<br />
The first, the treatment for the Guianas, is rather unfortunate. For historical reasons and differing<br />
approaches by the two authors, this is the kind of publication that gives a bad name to mycological αtaxonomists<br />
struggling to deal with biodiversity before it is gone. It leaves us open to criticism (and denial of<br />
funding) by those employing more sophisticated techniques. Three "species" concepts for ascospore variation<br />
are used in the work without adequate warning or explanation. Following Santesson (1952) Sipman divided<br />
foliicolous Porina into very narrowly defined species. In contrast Aptroot used a very broad concept for the<br />
corticolous species, with what I would consider an unacceptably large amount of variation. A third treatment of<br />
ascospore variation is found in Trichothelium where a distinct morphotype is assigned to the varietal level. I<br />
think that as α-taxonomists we have an obligation to be consistent in our species concepts within a group or, if<br />
not, then to provide some "hard" evidence as to why not. The treatment is also flawed in more obvious ways,<br />
types of taxa described from the Guianas have not been studied and names have been used with little logic<br />
(see criticisms by McCarthy, 1995). Such an interesting and complex family deserves better than this hasty<br />
attempt.<br />
McCarthy has provided two good, well illustrated α-taxonomic revisions. As will be seen below I do not<br />
agree with him in all details (mostly in regard to recognition of North American populations which consistently<br />
differ from his broader concepts). Thus I find Aptroot's comment in his review of McCarthy (1992) (Bryologist<br />
97: 467. 1994) most amusing: "However, the species concept applied is rather narrow and some specimens<br />
seem to represent transitional forms." I suspect that McCarthy's approach may well represent the mean<br />
between the lumper and the splitter and therefore be the most useful to the general lichenologist trying to<br />
name a specimen.<br />
For those who dislike change, the paper by Hafellner and Kalb is most unfortunate, not only in itself, but<br />
worse it has encouraged me to go even further in rearranging the Trichotheliaceae. They used the<br />
pigmentation of the ascomatal wall to divide Porina s. lat. into Porina and Pseudosagedia. These groups have<br />
been long recognized with varying degrees of clarity but mostly nomenclatural problems have led to the<br />
retention of a broad Porina. Hafellner and Kalb have cut the Gordian Knot. Below I have taken a different<br />
approach that essentially raises to the generic level the major groups recognized by Santesson (1952), uniting<br />
the P. nitidula group (Pseudosagedia) with Trichothelium, removing the P. phyllogena group to Strigula and<br />
adding a redefined Clathroporina. Amazing as it seems, this does not require the description of any new<br />
genera.<br />
Looking first at the taxa with blackish ascomata containing the pigment Pseudosagedia-violet, found in<br />
both Pseudosagedia and Trichothelium, the question is "Are setose ascomata a generic character?". I do not<br />
believe so. Hafellner and Kalb admitted there is an intermediate species with "bumps" on the ascomata (I<br />
commend their honesty, I wouldn't have known). The tendency to develop ascomatal appendages in<br />
159
foliicolous species may be a general phenomenon. It is seen in the P. leptalea group (Segestria/Polycornum)<br />
and in the unrelated Thelenellaceae. Further, both the setose and non-setose species of Trichothelium share<br />
the tendency towards long, multiseptate ascospores, only very rarely becoming somewhat muriform and<br />
growing if not on leaves, in wet, even semiaquatic habitats. The presence of setae has no predictive value<br />
and even if it were to be demonstrated that the setose taxa form a monophyletic clade, I would be reluctant to<br />
recognize more than a single genus, Trichothelium. The problem of how to deal with a single species or group<br />
with a single strikingly different character has always been a problem, e.g., placement of the compound-leaved<br />
Acer negundo. It is my understanding the theory (but perhaps not the practice) of modern phylogenetic<br />
approaches is that the preponderance of evidence should outweigh any single character, e.g., the<br />
abandonment of distinctive single character lichen genera by the merger of Coelocaulon into Cetraria or the<br />
realignment of Ionaspis/Hymenelia (abandoning the photobiont character). With the exception of the<br />
foliicolous species, the distribution of Trichothelium is amphitropical. I also include Zamenhofia in<br />
Trichothelium. It was tentatively maintained by Hafellner and Kalb on the basis of lacking a chitinoid ascus<br />
ring and lacking pigments. I am very hesitant to give possible "loss mutations" generic status. Also the assay<br />
used for the pigments was the crude application of KOH, etc., and also there seems to be some disagreement<br />
, "... some species with a layer reacting K+ red" (Purvis et al., 1992). There is general agreement among<br />
recent studies that the original characters for Zamenhofia, isidia and periphyses, are meaningless at the<br />
generic level. (I do part company with McCarthy (1993) who dismissed the use of isidia at any level. I agree in<br />
a couple of the cases he cited that the "isidia" are due to abnormal growth of the photobiont but isidiate<br />
species are not uncommon, especially in Clathroporina where sterile isidiate thalli are common and isidia may<br />
be the main means of reproduction in tropical wet forests.)<br />
When it comes to Hafellner and Kalb's residual Porina, I would move in the other direction, splitting rather<br />
than lumping. Part of the impetus for this approach came from McCarthy's (1995) discovery that if one<br />
merges Clathroporina with Porina (as he would and I whole-heartedly agree), the older name is Clathroporina.<br />
The groups involved are those centered around P. lectissima, P. nucula and Clathroporina eminentior. The<br />
first, for which the name Segestria Fr. is available, can be separated by lack of oxalate crystals in thallus and<br />
ascomata, small, ± naked ascomata, small ascospores and corticolous/saxicolous species basically<br />
amphitropical. Hafellner and Kalb pointed out two species intermediate between this group and the P. nucula<br />
group. I suggest that they are actually reduced members (in adaptation to living on leaves) of the P. nucula<br />
group. The P. nucula group is separated from Clathroporina not on ascospore sepatation (transverse vs.<br />
muriform) but on thallus characters which lead to very different aspects. Clathroporina has a smooth, often<br />
shiny, highly organized thallus with a cortex, photobiont ± organized in vertical rows, a layer of oxalate crystals<br />
and a dark hypothallus. The thallus is not intimately involved with the substrate but separated by the<br />
hypothallus and often peels away becoming almost foliose. The ascomata are ± immersed, sessile or<br />
protruding from the lower side of the thallus (well illustrated by McCarthy, 1995). To Clathroporina as defined<br />
by its type, C. eminentior, I would add the P. mastoidea group, differing in having transversely septate<br />
ascospores. Hafellner and Kalb recognized Clathroporina on ascus characters: lack of chitinoid ring and<br />
presence of a constriction near the tip in young asci. I consider usage of such characters as grasping at<br />
straws but they could be right, in which case a new home would have to be found for the P. mastoidea group.<br />
Porina nucula, the type of Porina, has a less organized, often verrucose thallus lacking a dark hypothallus with<br />
sessile ascomata, emergent and often constricted at the base. Again ascospore septation is not a character<br />
as several species with muriform ascospores are referable to Porina in this restricted sense. These genera<br />
are also supported by ecology and distribution. Clathroporina is mainly a genus of wet, dark, mostly tropical<br />
forests. It is one of the few groups, like Phyllopsora and Thelotremataceae, that are adapted to the lower<br />
levels of tropical rainforest and Caribbean wet broadleaf forest. (Is the development of a hypothallus an<br />
adaptation to this habitat?) Porina may be basically subtropical, adapted to more open forests with higher light<br />
and probably less moisture.<br />
The generic definitions suggested here are rather crude but by using them as a hypothesis, additional<br />
characters can be examined in detail to evaluate their utility. It is possible that conidial characters may support<br />
the new generic arrangement. At this time there is too little data about microconidia. Macroconidia are<br />
reported (Purvis et al. 1992) for two species referred here to Trichothelium. They may also occur in Segestria<br />
leptalea but I need several more occurrences to hypothesize an anamorphic connection. Ascospore<br />
tendencies also give some support, large with a strong tendency to become muriform in Clathroporina,<br />
medium to large with little tendency to become muriform in Porina, small, few septate in Segestria, long,<br />
160
multiseptate in Trichothelium. Ascomatal setae are confined to Segestria and Trichothelium. Clathroporina<br />
also differs in having more organized isidia, the ostiolar area is often black and there are no foliicolous<br />
species.<br />
The recognition of these four genera, Clathroporina, Porina, Segestria and Trichothelium, is somewhat<br />
analogous to generic segregation in Parmelia, distinctive in aspect (characters of the association) but ±<br />
uniform in characters of the asci and ascospores. I feel that the genera thus constituted have considerable<br />
predictive value regarding probable ascospore type, ecology and distribution. There seem to be four basic<br />
thallus/ascomata paradigms with rather continuous ascospore variation within each. This can be seen in the<br />
specific keys below where ascomata play only a small role. I simply do not understand the ascospore<br />
variation in the Trichotheliaceae. As pointed out by Aptroot and Sipman, it seems to encompass a different<br />
order of magnitude from that which mycologists are accustomed. In fact I consider this one of the distinctive<br />
features defining Trichotheliaceae. However, no one, myself included, has put the time and effort in to<br />
analyzing the situation. This leaves a situation where "species" are intuitively defined and one person's<br />
speculation is a good as another's. I consider my treatment below speculative at best and subject to<br />
considerable correction.<br />
Most of the species treated by McCarthy (1995) remain in Clathroporina under the new circumscription.<br />
Porina tahitensis (Räsänen) McCarthy and Clathroporina nuculastrum Müll. Arg. belong in Porina as defined<br />
here.<br />
Caribbean Porina firmula Müll. Arg. and P. subfirmula Riddle have very odd, placodioid thalli and may be<br />
considered for generic recognition. Two other species that may be referable to Segestria but are anomalous<br />
because of the pale pinkish ascomata and long, multiseptate ascospores and occur in eastern North America<br />
are an isidiate taxon reported from Long Island, New York reported by Brodo as Porina nucula (Porina sp. 4 in<br />
thesis), and a non-isidiate one from New Hampshire (Porina sp. 2 in thesis). I have not had time to reexamine<br />
the material, especially with regard to generic disposition, and as they are both very rare, I have<br />
decided to omit them here.<br />
KEY TO GENERA<br />
1. Ascomata usually small, essentially naked, with at most a thin covering of thallus<br />
(occasionally immersed in substrate); ascomata and thallus lacking oxalate<br />
crystals; ascomatal wall either some shade of red (Porina-yellow, Sagedia-red) or<br />
brown to blackish, often with blue or violet tones (Pseudosagedia-violet); some<br />
species with setose ascomata; muriform ascospores very rare; considerable<br />
diversity on leaves; distribution, except for foliicolous species, amphitropical .......................................... 2<br />
2. Ascomata blackish; wall often with blue or violet tints, without orange or<br />
red tints, KOH+ purplish or KOH-(Pseudosagedia-violet); with a very<br />
strong tendency for muitiseptate, elongate (even filiform) ascospores;<br />
setose species not uncommon........................................................................................ Trichothelium<br />
2. Ascomata reddish, sometimes darkening to black, lacking blue or violet<br />
tints, KOH+ more intensely orange or red (Porina-yellow); ascospores<br />
short, mostly 3-7-septate; setose species rare .......................................................................Segestria<br />
1. Ascomata usually rather large, surrounded by well developed thalline wart or<br />
immersed in thallus; wart and thallus containing oxalate crystals; ascomatal wall<br />
some shade of yellow or red (Porina-yellow); ascomata not setose; muriform<br />
ascospores not uncommon; few species on leaves; distribution essentially<br />
tropical and subtropical ............................................................................................................................. 3<br />
3. Black hypothallus lacking; thallus firmly attached to substrate, poorly<br />
organized, often verrucose; ascomata superficial, often somewhat constricted<br />
at base; few muriform-spored species; some follicolous species known; mainly<br />
species of open, drier? forest or scrub habitats, perhaps mainly subtropical ...............................Porina<br />
161
3. Black hypothallus usually well developed; thallus often separating from<br />
substrate, usually well organized into cortical, algal and oxalate crystal<br />
layers, smooth, continous, shiny; ascomata mostly immersed to weakly<br />
emergent, sometimes forming spherical warts projected below the almost<br />
foliose thallus; ascospores large, often muriform; no foliicolous species known;<br />
mostly species of shaded, closed, wet, tropical forests ................................................... Clathroporina<br />
CLATHROPORINA Müll. Arg.<br />
1. Thallus not isidiate..................................................................................................................................... 2<br />
2. Ascospores 8-celled, 40-55 × 7.5-9(-11) µm; Dade and Seminole<br />
counties ......................................................................................C. subpungens (Malme) R. C. Harris<br />
2. Ascospores 8-celled, 35-50 × 5-7 µm; Collier, Dade and Monroe<br />
counties ..............................................................................................C. tetracerae (Ach.) R. C. Harris<br />
1. Thallus isidiate; ascospores 8-celled, 32-48 × 5-6(-7.5) µm; Dade, Monroe<br />
and Polk counties ................................................................................................ C. isidiifera R. C. Harris<br />
NOTES<br />
Clathroporina confinis Müll. Arg. = Porina nuculastrum (Müll. Arg.) R. C. Harris<br />
Clathroporina isidiifera R. C. Harris, sp. nov.<br />
Similis C. tetracerae sed thallo isidiifero, ascosporis 8-loculis, 36-48 × 5-6 µm.<br />
Type. FLORIDA. Polk County: Nalcrest, scrub W of recreation area, 28 Jan 1989, Wheeler s.n. (NY, holotype).<br />
Thallus light green fresh, grayer in herbarium, shiny, continuous, not separating from substrate, with a<br />
dark brown to black hypothallus, isidiate; isidia cylindrical to slightly clavate, occasionally branched, with a ca.<br />
10-15 µm cortical/algal layer and a ca. 25 µm diam. core of oxalate crystals. Ascomata subglobose, 0.2-0.3<br />
mm diam., immersed or in somewhat emergent warts; wart 0.4-0.7 mm diam.; wall colorless below to<br />
yellowish or orange-brown around ostiole. Asci cylindrical, ca. 140 × 12 µm, with eight obliquely ± biseriate<br />
spores. Ascospores cylindrical fusiform, mostly ± equally tapered at either end, 8-celled, 32-48 × 5-6(-7.5) µm;<br />
microconidia not found.<br />
There seem to be rather few isidiate species described. Based on the literature I originally identified this<br />
material as Porina isidiophora Vainio but having seen specimens at Turku, I agree with McCarthy that the<br />
"isidia" in that taxon are abnormal outgrowths of Trentepohlia. The same is true of Porina granulifera Vainio.<br />
The isidia of C. isidiifera are true isidia, constricted at the base with a organization similar to that of the thallus.<br />
Porina conspersa Malme from Brazil may be close (I have not seen material) but the ascospores are<br />
described as 50-65 × 7-9 µm.<br />
Additional specimens. (With ascomata) FLORIDA. Dade County: Deering Hammock, Cutler, Dec 1921,<br />
Small et al.10279, Snapper Hammock, 11 Dec 1919, Britton & Britton 793; Monroe County: S part of Big Pine<br />
Key, 8 May 1919, Small 9125; Polk County: as type, 28 Jan, 20, 21 May 1989, Wheeler s.n. (Sterile) Dade<br />
County: Hammer Key, Everglades, 12 May 1918, Small 8609, Royal Palm Hammock, 27 Apr 1918, Small<br />
8523; Polk County: as type, 5 Feb, 11 Feb, 22 Apr, 6 May 1989, Wheeler s.n. (all NY).<br />
Clathroporina mastoidea (Ach.) R. C. Harris, comb. nov.<br />
162
Pyrenula mastoidea Ach., Ges. Naturf. Berlin Mag. Neusten Entdeck. Gesammten Naturk. 6:16.<br />
1814. Porina mastoidea (Ach.) Müll. Arg., Bot. Jahrb. Syst. 6: 399. 1885. Type. PHILIPPINES.<br />
Manila (UPS, lectotype, Malme, 1929, not seen; isolectotype, H-Ach 830-A).<br />
Although reported from North America, all the material which I have examined has ascospores smaller<br />
than those of the type of C. mastoidea. The epithet mastoidea has been widely misapplied in North America,<br />
usually to taxa in the Porina nucula group.<br />
Clathroporina subpungens (Malme) R. C. Harris, comb. nov.<br />
Porina subpungens Malme, Ark. Bot. 23A(1): 17. 1929. Type. PARAGUAY. Colonia Risso, Guarda<br />
Francia, 16 Oct 1893, Malme, Regn. Lich. 1927 (S, lectotype, Mccarthy, 1993).<br />
McCarthy (1993) placed this name in synonymy with Porina mastoidea (Ach.) Müll. Arg., that has larger<br />
ascospores (50-62 × 8-9(-10.5) µm in lectotype). On a global scale McCarthy may well be correct but since<br />
the lectotype of P. mastoidea is Asian and all the North American material has smaller ascospores, and<br />
because a name for a taxon so defined is available, I have chosen to take up Malme's name pending a careful<br />
statistical study of ascospore variation or perhaps molecular study. Although this species and the next were<br />
included in my thesis, this is the first published record of their occurrence in North America.<br />
New to North America. FLORIDA. Dade County: Royal Palm State Park, 6 Dec 1919, Britton & Britton 763<br />
(FH, NY), hammock, E end of Long [Pine] Key, Everglades, Small 8397 (NY); Seminole County: Sanford, on<br />
oak, Feb 1919, Rapp 746 (MICH).<br />
Clathroporina tetracerae (Ach.) R. C. Harris, comb. nov.<br />
Verrucaria tetracerae Ach., Methodus 121. 1803. Porina tetracerae (Ach.) Müll. Arg., Bot. Jahrb.<br />
Syst. 6: 401. 1885. Type. GUINEA. (H-Ach 838-C, lectotype, selected here).<br />
McCarthy (1993) cited H-Ach 838 as the holotype. It is mixed: 838-A is a Pyrenula. 838-B is internally<br />
identical to 838-C but has ascomata ca. 2× as large. This refinement of the lectotypification is purely<br />
precautionary and has no effect on the circumscription at this time. Material at S also contains a mixture<br />
pieces with large and small ascomata.<br />
Clathroporina tetracerae seems to have been more common in the past than it is now. It will be<br />
interesting to see if this is lack of collecting or loss of habitat as Clathroporina, with its preference for rain<br />
forests, is likely to be the most ecologically sensitive member of the Trichotheliaceae.<br />
New to North America. FLORIDA. Collier County: Fakahatchee Strand State Preserve, Royal Palm<br />
Hammock, 6 Dec 1992, Harris 29932 (NY); Dade County: Cocoanut Grove, 1898, Thaxter 277 (FH), 287<br />
(MICH), Deering Hammock, Cutler, 29 Apr 1918, Small 8352 (NY), Hattie Bauer Hammock, Small & Mosier<br />
5954 (NY), Royal Palm Hammock State Park, 6 Dec 1919, Britton & Britton 757 (NY), Snapper Hammock, 11<br />
Dec 1919, Britton & Britton 800 (FH, NY); Monroe County: Key Largo, Apr 1921, Kelly (MICH).<br />
PORINA Müll. Arg., nom. cons.<br />
1. Thallus and ascomatal warts not isidiate .................................................................................................. 2<br />
2. Ascospores transversely septate .......................................................................................................... 3<br />
3. Growing on bark ................................................................................................................................ 4<br />
4. Ascospores fusiform to fusiform-cyllindrical, 7(-9)-septate ........................................................... 5<br />
5. Ascospores larger, over 40 × 8 µm ........................................................................................... 6<br />
6. Ascospores less than 60 µm long ......................................................................................... 7<br />
7. Ascospores broad, with rounded ends, 40-52 × 11-14 µm,<br />
8-celled; Dade and Lee? counties.............................................................. P. nucula Ach.<br />
163
7. Ascospores narrower, with pointed ends, 42-53(-61)<br />
× (7-)8-11(-12) µm, 8(-10)-celled; eastern Florida,<br />
Citrus to Dade County .................................................................................Porina sp. 674<br />
6. Ascospores 60-75 × 10-13(-15) µm, 8-celled, ends pointed,<br />
± evenly tapered at both ends; ascomatal warts rather large,<br />
ca. 0.6 mm diam.; Collier, Dade, Putnam and Seminole<br />
counties ................................................................................................................Porina sp. 6<br />
5. Ascospores small, 25-33 × 5-6 µm, 8-celled; ascomatal warts small,<br />
ca. 0.3 mm diam.; Collier County ....................................................................Porina sp. 30308<br />
4. Ascospores clavate with long tapering tail, (9-)11-13-septate,<br />
85-110 × 12-14 µm; common, Texas to North Carolina south to<br />
Seminole County in Florida .................................................... P. heterospora (Fink) R. C. Harris<br />
3. Growing on HCl- rock; thallus relatively smooth; asci ca. 130 × 20 µm;<br />
ascospores 8-celled, fusiform, 43-49 × 8-9 µm; Hillsborough County................................................<br />
.....................................................................................................................Porina sp. Buck 22669<br />
2. Ascospores fusiform, muriform, 9-13-septate with few to most cells longitudinally<br />
or obliquely 1-2-septate, 55-80 × 13-18 µm, usually with well developed gel sheath;<br />
Dade and Lee? counties .......................................................P. nuculastrum (Müll. Arg.) R. C. Harris<br />
1. Thallus and/or ascomatal warts isidiate; ascospores 8-celled, 25-47(-57) × 6.5-8 µm;<br />
North Florida, Louisiana and South Carolina ...................................................... P. scabrida R. C. Harris<br />
NOTES<br />
Porina heterospora (Fink) R. C. Harris<br />
McCarthy (1993) placed this taxon as a synonym of Porina guaranitica Malme. Although I am generally<br />
rather broad minded at the species level, I cannot bring myself to accept this disposition. I have examined the<br />
lectotype of P. guaranitica. It is on rock with a very smooth pinkish thallus. Porina heterospora occurs on bark<br />
and has a verrucose, usually tan thallus. One might be able to dismiss the difference as a consequence of the<br />
semiaquatic habitat of P. guarantica and I would probably be willing to do so if the ascospore size ranges were<br />
more congruent. Based on ten ascospores from the lectotype of P. guaranitica the mean ascospore length is<br />
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<br />
(range 60-112 µm) and 11.1 septa. McCarthy gave a mean ascospore size of 71.5 µm, presumably based on<br />
a mixture of P. guaranitica and P. heterospora but still well under that of North American collections. The form<br />
of the ascospores is different also. Both are clavate but P. heterospora has a longer tail, seemingly<br />
accounting for the greater ascospore length. Therefore, on the basis of larger, more sepate, longer tailed<br />
ascospores, thallus characters and corticolous substrate, I am maintaining P. heterospora as a distinct<br />
species. This would seem to be the most common species of Porina in the southern Coastal Plain. NY holds<br />
specimens of typical P. heterospora from Cuba and Venezuela but none seen as yet from Brazil. Since the<br />
epithet heterospora has a history of usage both in North America and Great Britain, even if it were a synonym,<br />
I suspect it would be conserved.<br />
Porina nucula Ach.<br />
McCarthy (1993) cited H-Ach 731 as the holotype of this name. However, this consists of two nonidentical<br />
pieces. H-Ach 731-A should be considered the lectotype. H-Ach 731-B is Porina mastoidea s. lat. A<br />
narrow species concept is used here, including only those collections with rather broad, rather short<br />
ascospores with rather blunt ends. Obviously the break between this circumscription and other putative taxa<br />
is not as tidy as one might like. However, I prefer to cut too finely rather than lose information in too broad a<br />
164
junk heap. These variants (sp. 6, sp. 674, sp. Buck 22669) have not been named as there are many<br />
uninvestigated names potentially available. They vary mainly in ascospore shape (more acutely pointed) and<br />
ascospore size. Buck 22669 grew on rock and may possibly be referable to Porina simulans Malme sensu<br />
McCarthy.<br />
Porina nuculastrum (Müll. Arg.) R. C. Harris, comb. nov.<br />
Clathroporina nuculastrum Müll. Arg., Flora 67: 618. 1884. Type. MADAGASCAR. Nossi-bé,<br />
Hildebrandt, comm. Rensch, 1884 (G, holotype).<br />
Since muriform ascospores have obviously evolved several times in the Trichotheliaceae, this character<br />
cannot be used to form a monophyletic genus. As Müller's name indicates, Clathroporina nuculastrum is most<br />
closely related to Porina nucula (they are externally indistinguishable) and the name is accordingly transferred<br />
to Porina. Two other species with submuriform/muriform ascospores belong in Porina as defined here, Porina<br />
ocellata (Malme) McCarthy (1993) and P. tahitensis (Räsänen) McCarthy (1995). See McCarthy (1995) for a<br />
list of synonyms of Porina nuculastrum, that was previously recorded from North America under one of these,<br />
Clathroporina confinis Müll. Arg. McCarthy (1995) incorrectly mapped the Florida collections in the central<br />
part of the state. I am unaware of any recent collections.<br />
FLORIDA. "East Florida", 1877, Smith & Austin (FH-Tuck 3929, MICH, NY), "Southeastern Florida", Smith<br />
(MICH, holotype of Polyblastiopsis floridana Fink in Hedrick); Dade County: Cocoanut Grove, 1898, Thaxter<br />
330 (FH), 296, 326 (MICH).<br />
Porina scabrida R. C. Harris, sp. nov.<br />
Subsimilis P. nuculae sed thallo isidiato et ascosporis minoribus, 37-42 × 5.5-7.5 µm, extremiis<br />
acutioribus.<br />
Type. FLORIDA. Columbia County: W side of Woodland Avenue, off Co. Rd. 138, 0.5 mi W of US 27, 29°51'N,<br />
82°39'W, hardwood forest, on Fraxinus, 5 Dec 1994, Buck 27418 (NY, holotype).<br />
Thallus pale olive tan, verrucose, dull to slightly shiny, densely to sparsely isidiate or occasionally nonisidiate;<br />
isidia cylindrical, occasionally branched, often rather irregular, consisting mostly of the photobiont with<br />
an external layer one cell thick. Ascomatal warts sessile, complanate to hemispherical, sometimes slightly<br />
constricted at base, 0.5-0.7 mm diam., often bearing cylindrical isidia. Asci cylindrical, ca. 120 × 17 µm, with<br />
eight obliquely subbiseriate spores. Ascospores fusiform-cylindrical, 8-celled, 35-47(-57) × 5.5-8 µm, both<br />
ends pointed, tapering ± evenly. Microconidia not found.<br />
Lacking isidia, the ascospore size would distinguish this taxon in North America. Even if P. scabrida<br />
were considered an aberrantly isidiate morph, there is no obvious non-isidiate taxon to refer it to. The thallus is<br />
sometimes lacking isidia, especially when ascomata are abundant, but one can then usually find them on the<br />
ascomatal warts. This is apparently a species of the southern Coastal Plain with its southern limit in Dixie and<br />
Putnam counties, Florida.<br />
Additional specimens. FLORIDA. Dixie County: Steinhatchee Wildlfie Management Area, Pumpkin Swamp,<br />
ca. 7 mi NW of Cross City, 29°41'N, 83°13'W, hardwood-Taxodium swamp, on Nyssa, 4 Dec 1993, Harris<br />
31539-A (NY); Lafayette County: along Co. Rd. 357, 6.4 mi SE of Fla. Hwy. 51, 29°51'N, 83°15'W, hardwood-<br />
Taxodium swamp, 2 Dec 1994, Buck 27291 (NY); Leon County: Apalachicola Natl. Forest, Rocky Bluff Scenic<br />
Area, ca. 3 mi S of Bloxham, mesic hardwoods, on Quercus, 4 Dec 1988, Harris 23411 (NY), swamp forest E<br />
of Natural Bridge Battlfield Monument, 6 mi E of Woodville, on Taxodium, 1 Jan 1991, Harris 26227 (NY);<br />
Putnam County: Ocala Natl. Forest, old Johnson Field Campground along Oklawaha River, just W of US Hwy.<br />
19, swamp forest, 7 Dec 1988, Buck 16807 (NY); Suwannee County: Ichetucknee Springs State Park, near N<br />
entrance, 29°59'N, 82°46'W, wet hardwoods, 2 Dec 1992, Buck 22582 (NY). LOUISIANA. East Baton Rouge<br />
Parish: Centurion Development, 4 mi E of Baton Rouge, 29 May 1976, Wetmore 24892 (MIN); Rapides<br />
Parish: Magnolia Recreation Area, Kisatchie Natl. Forest, 12 mi SW of Alexandria, ca. 70 m, mixed deciduous<br />
forest, 31 Dec 1969, Egan EL-7305, 7339, 7350 (MIN); St. Martin Parish: near St. Martinville, on Carya<br />
165
aquatica, 15 Jan 1894, Langlois (NY). SOUTH CAROLINA. Santee Canal, Ravenel, Rel. Tuck. 136 (NY).<br />
Porina sp. 30308<br />
Known from a single Collier County collection, this taxon is distinct in its small ascomata and ascospores.<br />
On this basis it is similar to P. curtula Malme but lacks the dark ostiolar area emphasized by McCarthy (1993).<br />
SEGESTRIA Fr.<br />
Type. S. lectissima Fr.<br />
Polycornum Malcolm & V_zda, Folia Geobot. Phytotax. 30: 91: 1995. Type. P. rubrofuscum<br />
Malcolm & V_zda.<br />
As with Trichothelium, I do not regard the setose ascomata of Polycornum sufficient to define a genus.<br />
KEY TO SEGESTRIA IN EASTERN NORTH AMERICA<br />
1. Growing on bark or rock............................................................................................................................ 2<br />
2. Ascospores 4-celled, fusiform ............................................................................................................... 3<br />
3. Ascospores 16-23 × 3-5 µm (Purvis et al., 1992); on bark and rock;<br />
Leon and Liberty counties ................................................S. leptalea (Durieu & Mont.) R. C. Harris<br />
3. Ascospores 20-30 × 5-8 µm (Purvis et al., 1992); on rock;<br />
New Hampshire, Mississippi and New York ............................................................[S. lectissima Fr.]<br />
2. Ascospores 8-celled, clavate, 37-43 × 7-8 µm; Tennessee<br />
(Porina sp. 5 in thesis)...................................................................................................... [Segestria sp.]<br />
1. Growing on leaves..................................................................................................................................... 3<br />
4. Ascospores 4-celled, 14-20 × 2.5-4 µm (Santesson, 1952); Escambia and<br />
Seminole counties ........................................................................... S. rubentior (Stirton) R. C. Harris<br />
4. Ascospores 8-celled, 24-31 × 3-4 µm (Santesson, 1952); Dade County<br />
................................................................................................... S. octomera (Müll. Arg.) R. C. Harris<br />
NOTES<br />
Segestria leptalea (Durieu & Mont.) R. C. Harris, comb. nov.<br />
Biatora leptalea Durieu & Mont., Expl. sci. Algérie 1: 268. 1847. Porina leptalea (Durieu & Mont.)<br />
A. L. Sm., Monogr. Brit. lich. 2: 333. 1911. Type not seen.<br />
?Porina olivacea var. microspora Fink in Hedrick, Mycologia 25: 308. 1933. Type. NEW<br />
HAMPSHIRE. Lonely Lake, on Ilex, Farlow (MICH, Fink hb. 11,559, holotype).<br />
Eastern North American specimens on both bark and rock have oblong or rod-shaped microconidia, 3-6<br />
× 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<br />
Harris (1975) and Fink's name was informally raised to the species level for eastern American material.<br />
However, whether there are any correlated differences remains to be studied. It may be that it should be<br />
treated as only a minor population variant. In addition to Florida and New Hampshire collections on bark it is<br />
known from Illinois and Kentucky on rock.<br />
Segestria mammillosa Th. Fr.<br />
Although distinctive in occurring on bryophytes and detritus, the lack of oxalate crystals, pigmentation and<br />
small 4-celled ascospores suggest placement here. It is known from western North America.<br />
Segestria octomera (Müll. Arg.) R. C. Harris, comb. nov.<br />
Phylloporina octomera Müll. Arg., Flora 73: 198. 1890. Porina octomera (Müll. Arg.) Schilling,<br />
166
Hedwigia 67: 274. 1927. Type not seen.<br />
This was known to me in 1975 from a single Thaxter collection from Dade County. I have lost track of<br />
the exact data.<br />
Segestria rubentior (Stirton) R. C. Harris, comb. nov.<br />
Verrucaria rubentior Striton, Proc. Philos. Soc. Glasgow 11: 107. 1878. Porina rubentior (Stirton)<br />
Müll. Arg., Flora 66: 334. 1883. Type not seen.<br />
New to North America. FLORIDA. Escambia County: along Pineville Road at Brushy Creek, ca. 2 mi W of<br />
Co. Rd. 97A, 30°48'N, 87°33'W, on bambusoid grass culm, 8 Dec 1993, Buck 24587 (NY); Seminole County:<br />
Sanford, on Sabal leaf, May 1931, Rapp (NY).<br />
TRICHOTHELIUM Müll. Arg.<br />
Type. T. epiphyllum Müll. Arg.<br />
Pseudosagedia (Müll. Arg.) Choisy, Bull. Mens. Soc. Linn. Soc. Bot. Lyon 18: 107. 1949.<br />
Arthopyrenia sect. Pseudosagedia Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 16: 428.<br />
1862. Lectotype (Hafellner & Kalb, 1995): Arthopyrenia carpinea (Pers. ex Ach.) Müll. Arg. (=<br />
Trichothelium aeneum (Wallr.) R. C. Harris).<br />
Zamenhofia Clauzade & Roux, Bull. Soc. Bot. Centre-Ouest, n.s., Num. Spéc. 7: 824. 1985. Type.<br />
Z. coralloidea (P. James) Clauzade & Roux<br />
It should be noted that if the typification of Arthopyrenia proposed elsewhere in this work is not accepted<br />
Trichothelium/Pseudosagedia may require conservation over Leiophloea. The rationale for uniting<br />
Pseudosagedia and Trichothelium is included in the discussion of the family above. I have no opinions about<br />
subgenus Limosagedia Hafellner & Kalb and therefore do not transfer it to Trichothelium. If I were to erect<br />
subgenera, I probably would create one for the distinctive T. crocynioides. It could well be that Trichothelium<br />
can be divided into smaller homogeneous genera but I feel that this has to await a critical re-evaluation of this<br />
whole group.<br />
KEY TO TRICHOTHELIUM IN EASTERN NORTH AMERICA<br />
1. Growing on bark or rock............................................................................................................................ 2<br />
2. Ascomata not setose............................................................................................................................. 3<br />
3. Thallus not isidiate............................................................................................................................. 4<br />
4. Thallus without obvious hypothallus .............................................................................................. 5<br />
5. Ascospores not filiform, 4-13-celled.......................................................................................... 6<br />
6. Ascospores 4-celled.............................................................................................................. 7<br />
7. Growing on bark; ascospores 20-24 × 4-5 µm; Polk and<br />
Seminole counties........................................................... T. aeneum (Wallr.) R. C. Harris<br />
7. Growing on rock; ascospores 18-25 × 4-5 µm;<br />
Massachusetts, New York............................................ [T. chloroticum (Ach.) R. C. Harris]<br />
6. Ascospores 8-13-celled......................................................................................................... 8<br />
8. Growing on bark ................................................................................................................ 9<br />
9. Ascospores clavate-fusiform or cylindrical; ascomatal wall<br />
brown or blue or violet in section................................................................................ 10<br />
167
10. Ascospores clavate-fusiform, 8-13-celled ............................................................. 11<br />
11. Ascospores 8-celled........................................................................................ 12<br />
12. Ascospores (32-)38-50 × 5.5-7.5 µm; common,<br />
throughout Florida ........................... T. cestrense (Michener) R. C. Harris<br />
12. Ascospores 28-35 × 4-6.5 µm; rare, Dade,<br />
Lee and Seminole counties ...................................... T. cestrense complex<br />
11. Ascospores 8-11(-13)-celled, 47-65 × 5-7.5 µm;<br />
rare, throughout Florida................................................. T. cestrense complex<br />
10. Ascospores narrowly cylindrical, 8-13-celled,<br />
(34-)50-67 × 3.5-6 µm; central Florida ................................ T. cestrense complex<br />
9. Ascospores fusiform, 8(-9)-celled, 30-38 × 6-9 µm; ascomatal<br />
wall blue-green above in section, pale below; ascomata<br />
0.3-0.4 mm diam., Palm Beach County ........................Trichothelium sp. Thaxter 92<br />
8. Growing on rock, often seasonally inundated; ascospores<br />
fusiform-clavate to fusiform, 8(-12)-celled, 32-45(-48) × 5-8(-10) µm;<br />
Kentucky, Massachusetts, Missouri, North Carolina and Virginia.......................................<br />
..................................................................................... [T. guentheri (Flotow) R. C. Harris]<br />
5. Ascospores filiform, 14-21-celled, 100-140 × 3-5 µm;<br />
common, south to Polk and Sarasota counties...........................................................................<br />
........................................................................T. rhaphidospermum (Müll. Arg.) R. C. Harris<br />
4. Thallus soft, dull, on a black byssoid hypothallus; ascospores<br />
narrowly fusiform to cylindrical, 4(-7)-celled, 20-35 × 2.5-3.5 µm;<br />
on Taxodium; Lee and Polk counties .............................................. T. crocynioides R. C. Harris<br />
3. Thallus isidiate; isidia crowded, branched and irregular; ascomata<br />
± immersed and hidden by isidia; ascospores clavate-fusiform,<br />
8(-9)-celled, 43-54 × 6-7.5 µm; on bark; Illinois...........................................[T. isidiatum R. C. Harris]<br />
2. Ascomata setose; on bark; ascospores narrowly fusiform, 15-25-septate,<br />
with some cells longitudinally septate, 100-124 × 8-14 µm (Santesson, 1952);<br />
Marion County ................................................................................T. horridulum (Müll. Arg.) R. Sant.<br />
1. Growing on leaves or culms of bambusoid grasses; thallus a thin olive-tan film .................................... 13<br />
13. Ascomata not setose......................................................................................................................... 14<br />
14. Ascospores fusiform, 4-6-celled, less than 32 µm long .............................................................. 15<br />
15. Ascospores 4-celled, 18-25 × 4-5 µm (Santesson, 1952);<br />
Dade County....................................................................... T. thaxteri (R. Sant.) R. C. Harris<br />
15. Ascospores 6-celled, 20-28(-32) × 3.5-5.5 µm (Santesson, 1952);<br />
Alachua (on bambusoid), Collier and Dade counties ................................................................<br />
........................................................................................ T. nitidulum (Müll. Arg.) R. C. Harris<br />
14. Ascospores clavate, 8-celled, 38-42 × 7.5-8 µm; ascomatal wall<br />
168
dark green-blue in section; on bambusoid grass culm....................................................................<br />
.................................................................................T. cestrense (Michener) R. C. Harris s. lat.<br />
13. Ascomata setose; ascospores fusiform, 8-celled, (25-)36-50 × 4-6 µm<br />
(Santesson, 1952); Louisiana..........................................................................[T. epiphyllum Müll. Arg.]<br />
NOTES<br />
Trichothelium aeneum (Wallr.) R. C. Harris, comb. nov.<br />
Verrucaria aenea Wallr., Fl. Crypt. Germ. 3: 299. 1831. Porina aenea (Wallr.) Zahlbr., Cat. lich.<br />
univ. 1: 363. 1922. Pseudosagedia aenea (Wallr.) Hafellner & Kalb, Biblioth. Lichenol. 57:<br />
172. 1995. Type not seen.<br />
?Porina amygdalina Müll. Arg., Hedwigia 34: 34. 1895. Type. LOUISIANA. Pointe à la Hache,<br />
Eckfeldt [Langlois] 797, not seen.<br />
Trichothelium cestrense (Michener) R. C. Harris, comb. nov.<br />
Verrucaria cestrensis Tuck. ex Michener in Darlington, Fl. cestr. ed. 3: 452. 1853. Porina<br />
cestrensis (Michener) Müll. Arg., Flora 64: 338. 1883. Type. PENNSYLVANIA. Chester, Michener<br />
204 (FH-Tuck 3986, isotype?).<br />
Porina salicina Müll. Arg., Hedwigia 34: 34. 1895. Type. LOUISIANA. Around New Orleans in<br />
nearby woods, 24 Nov 1884, Langlois 796 [erroneously attributed to Eckfeldt in original<br />
publication] (PH, isotype?).<br />
Porina cestrensis var. platyspora Fink in Hedrick, Mycologia 25: 308. 1933. Type. INDIANA. Near<br />
Scipio, 8 Feb 1909, Fink (MICH, Fink hb. 8,889).<br />
In one of my more egregious blunders (as noted by McCarthy, 1993) I misinterpreted the typification of<br />
Verrucaria pulla Ach. (MülIer made the same mistake.) Instead of referring back to the Lichenographis<br />
universalis (1810), I looked at the Synopsis methodus lichenum (1814). Different specimens are cited in the<br />
two works. The first (the holotype, H-Ach 799-A) turns out to be a common (but until now nameless)<br />
Caribbean Arthopyrenia. The second (BM, H-Ach 799-B) on Dirca is T. cestrense.<br />
Trichothelium cestrense complex<br />
This group is one of the reasons that any sane person would avoid this genus. There is a virtually<br />
complete series of intermediates in ascospore shape and size between T. cestrense and T.<br />
rhaphidospermum. There are some with cestrense size ascospores but rod-shaped (similar to Porina borreri<br />
var. leptospora (Nyl.) D. Hawksw.), there are longer acicular ascospores but considerably shorter than<br />
rhaphidospermum and there are various sizes of fusiform-clavate ascospores longer and with more septa than<br />
"typical" cestrense. (Brodo's 1968 report of Porina hibernica P. James & Swinscow is based on some of the<br />
longer spored morphs of this complex . As far as I know P. hibernica does not occur in North America.) I have<br />
been aware of this mess since my student days but since it was only peripheral to my thesis, I made no<br />
attempt to resolve it. I still have not had the time (or energy) to try to find some pattern. I hope with new<br />
computer graphics and statistical analyses that eventually it can be sorted out. In addition to the variation I was<br />
aware of 20 years ago, the recent emphasis on the color of the ascomatal wall has added yet another<br />
dimension. The color of the wall in section in this group seems to be either brown or blue-violet. As it does<br />
not seem to correlate with any of the ascospore variations, it doesn't seem helpful at this time. Also I have no<br />
idea how specimen drying/preservation affects this character. In Florida there is somewhat less variation than<br />
farther north and the variation falls into two groups as indicated in the key. A third variant with ascospores<br />
smaller than usual may be the European "Porina" borreri but I do not have material to resolve this question at<br />
the moment.<br />
Trichothelium chloroticum (Ach.) R. C. Harris, comb. nov.<br />
Verrucaria chlorotica Ach., Lichenogr. univ. 283. 1810. Porina chlorotica (Ach.) Müll. Arg., Rev.<br />
Mycol. (Toulouse) 6: 20. 1884. Pseudosagedia chlorotica (Ach.) Hafellner & Kalb, Biblioth.<br />
Lichenol. 57: 172. 1995. Type not seen.<br />
169
Trichothelium crocynioides R. C. Harris, sp. nov.<br />
Species unica thallo subbyssoideo, pagina thalli hyphis irregularibus erectis, hypothallo nigro byssoideo<br />
et ascosporis angustis, 3-septatis, 20-30 × 3-4 µm.<br />
Type. FLORIDA. Highlands County: Highlands Hammock State Park, swampy area near stream, on cypress, 22<br />
Dec 1965, Wetmore 13907 (NY, holotype; FLAS, MIN & to be distributed, isotypes).<br />
Thallus light olive gray, continuous, smooth, dull to almost byssoid in appearance due to short, irregular<br />
erect hyphae projecting from thallus surface; hypothallus conspicuous, black, byssoid, composed of loose<br />
interconnected dark brown or gray brown hyphae. Ascomata black, immersed, subglobose, ca. 0.2-0.3 mm<br />
diam.; wall blackish brown, sometimes with purplish tint, reddish purple in HNO3. Asci cylindrical, 95-110 × 8-9<br />
µm, with eight subbiseriate spores. Ascospores narrowly fusiform to cylindrical, 4(-7)-celled, 20-35 × 2.5-3.5<br />
µm; microconidia not found.<br />
Trichothelium crocynioides is named for its superficial resemblance to Crocynia pyxinoides Nyl. The very<br />
distinctive thallus with erect, projecting hyphae and byssoid hypothallus may be worthy of generic or<br />
subgeneric recognition. I prefer to wait until the variation in Trichothelium is more fully understood. Thus far<br />
all specimens are from Taxodium.<br />
Additional specimens. FLORIDA. Lee County: Caloosahatchee River State Recreation Area, along Co. Rd.<br />
78 ca. 3 mi E of Telegraph Creek, 26°43'N, 81°39'W, Taxodium swamp, Buck 22994, Harris 30215, 30246<br />
(NY).<br />
Trichothelium guentheri (Flotow) R. C. Harris, comb. nov.<br />
Verrucaria guentheri Flotow, Bot. Zeitung 8: 575. 1850. Porina guentheri (Flotow) Zahlbr., Cat.<br />
lich. univ. 1: 384. 1922. Pseudosagedia guentheri (Flotow) Hafellner & Kalb, Biblioth. Lichenol.<br />
57: 172. 1995. Type not seen.<br />
This is treated in Europe as a highly variable species. The inclusion of North American collections does<br />
not appreciably add to the problem. Specimens from Kentucky, Missouri, North Carolina and Virginia are fairly<br />
homogeneous with clavate-fusiform, 8(-9)-celled ascospores, 32-45 × 5-6 µm. Collections from Maine and<br />
Massachusetts have larger, more septate ascospores and may prove closer to Porina grandis (Körber) Zahlbr.<br />
(spp. 7, 8, 9 of my thesis and reported for North America by McCarthy (1993) on this basis). The species is<br />
apparently mostly semiaquatic except for the Missouri collection which from dry rock.<br />
Porina hibernica James & Swinscow<br />
North American records are collections of the Trichothelium cestrense complex lacking isidia and having<br />
smaller ascomata than P. hibernica.<br />
Trichothelium isidiatum R. C. Harris, sp. nov.<br />
Similis T. cestrensi sed thallo dense isidiato et ascosporis fusiformi-clavati, 7-septatis, 43-54 × 6-7.5 µm.<br />
Type. ILLINOIS. Jackson County: Shawnee National Forest, Pomona Natural Bridge, ca. 8 mi S of<br />
Murphysboro, ca. 37°38'N, 89°20'W, beech-maple woods, at base of oak, 15 Oct 1993, Harris 31265 (NY,<br />
holotype).<br />
Thallus dark gray green, continous but mostly concealed by dense isidia; isidia ± cylindrical, irregular,<br />
occasionally branched, with a thin, papillose? cortex. Ascomata immersed, almost concealed by isidia, black,<br />
subglobose, ca. 0.3-0.4 mm diam.; wall brown above with bluish-violet tints in lower part, pale below. Asci<br />
fusiform-cylindrical, ca. 130-160 × 15-17 µm, with eight subbiseriate spores. Ascospores fusiform-clavate, 8(-<br />
9)-celled, 43-54 × 6-7.5 µm. Microconidia not found.<br />
170
Trichothelium isidiatum could be referred to Zamenhofia if one accepts this genus. It is rather similar in<br />
aspect to Zamenhofia hibernica. It does not have periphyses as far as I can tell. There seems to be one<br />
isidiate species per genus in North America, Clathroporina isidiifera, Porina scabrida, Segestria sp. (thesis<br />
Porina sp. 4) and Trichothelium isidiatum. The last is known only from the type collection.<br />
Trichothelium lineare (Leighton) R. C. Harris, comb. nov.<br />
Verrucaria linearis Leighton, Brit. angiocarp. lich. 52. 1851. Porina linearis (Leighton) Zahlbr., Cat.<br />
lich. univ. 1: 391. 1922. Pseudosagedia linearis (Leighton) Hafellner & Kalb, Biblioth.<br />
Lichenol. 57: 174. 1995. Type not seen.<br />
Trichothelium lineare is reported from North America but not known in the East.<br />
Trichothelium nitidulum (Müll. Arg.) R. C. Harris, comb. nov.<br />
Porina nitidula Müll. Arg., Flora 66: 336. 1883. Pseudosagedia nitidula (Müll. Arg.) Hafellner &<br />
Kalb, Biblioth. Lichenol. 57: 173. 1995. Type not seen.<br />
Trichothelium rhaphidospermum (Müll. Arg.) R. C. Harris, comb. nov.<br />
Porina rhaphidosperma Müll. Arg., Hedwigia 34: 35. 1895. Type. MASSACHUSETTS. On hemlock,<br />
Willey (FH-Tuck 3988, US, isotypes).<br />
Trichothelium thaxteri (R. Sant.) R. C. Harris, comb. nov.<br />
Porina thaxteri R. Sant., Symb. Bot. Upsal. 12(1): 218. 1952. Pseudosagedia thaxteri (R. Sant.)<br />
Hafellner & Kalb, Biblioth. Lichenol. 57: 173. 1995. Type. FLORIDA. Dade County: Cutler,<br />
Thaxter 421 (MICH, isotype).<br />
Trichothelium sp. Thaxter 92<br />
This taxon is possibly related to the foliicolous Porina moralesiae Lücking but I have no comparative<br />
material nor have I re-examined the specimens studied for my thesis. It was included as Porina sp. 1.<br />
INDEX TO SCIENTIFIC NAMES<br />
Names in boldface represent new nomenclature; names in italics are synonyms, and names in regular typeface<br />
are accepted names.<br />
Acrocordia............................................................................122<br />
Acrocordia cavata................................................................122<br />
Acrocordia conoidea............................................................122<br />
Acrocordia gemmata ...........................................................122<br />
Acrocordia megalospora .....................................................122<br />
Acrocordiaceae....................................................................121<br />
Acrocordiella..........................................................................93<br />
Acrocordiella occulta .............................................................93<br />
Amandinea.............................................................................45<br />
Amandinea polyspora............................................................46<br />
Amandinea punctata .............................................................46<br />
Anisomeridium.....................................................................123<br />
Anisomeridium adnexum..................................................143<br />
Anisomeridium albidoatrum.............................................144<br />
Anisomeridium albisedum...................................................126<br />
Anisomeridium ambiguum...................................................126<br />
Anisomeridium americanum ............................................144<br />
Anisomeridium angulosum ..............................................144<br />
Anisomeridium anisolobum...........................................69, 126<br />
Anisomeridium antillarum ................................................144<br />
Anisomeridium aureopunctatum .....................................127<br />
Anisomeridium holopolium ..............................................147<br />
Anisomeridium immersum ...............................................147<br />
Anisomeridium indicum....................................................147<br />
Anisomeridium infernale ..................................................147<br />
Anisomeridium interspersum...............................................130<br />
Anisomeridium juistense .....................................................129<br />
171<br />
Anisomeridium australiense............................................ 144<br />
Anisomeridium biforme....................................................... 127<br />
Anisomeridium biformoides ............................................ 127<br />
Anisomeridium carinthiacum .............................................. 144<br />
Anisomeridium complanatum ......................................... 145<br />
Anisomeridium consimile ................................................ 145<br />
Anisomeridium consobrinum .............................................. 145<br />
Anisomeridium distans ....................................................... 127<br />
Anisomeridium elegans ................................................... 145<br />
Anisomeridium endobryum ............................................. 146<br />
Anisomeridium epiphyllum ................................................. 146<br />
Anisomeridium excaecariae ............................................ 128<br />
Anisomeridium excellens ................................................ 146<br />
Anisomeridium feeanum..................................................... 126<br />
Anisomeridium finkii......................................................... 128<br />
Anisomeridium flavopallidum.......................................... 146<br />
Anisomeridium foliicola....................................................... 146<br />
Anisomeridium glaucescens........................................... 146<br />
Anisomeridium gregale .................................................... 146<br />
Anisomeridium griffinii..................................................... 128<br />
Anisomeridium laevigatum .............................................. 147<br />
Anisomeridium leptospermum........................................ 147<br />
Anisomeridium leucochlorum ............................................. 147<br />
Anisomeridium magnosporum............................................ 148<br />
Anisomeridium megalosporum....................................... 148<br />
Anisomeridium monosporum.......................................... 148
Anisomeridium nemorosum.............................................148<br />
Anisomeridium nidulans...................................................148<br />
Anisomeridium nyssaegenum.............................................128<br />
Anisomeridium pacificum.................................................149<br />
Anisomeridium palavanum...............................................149<br />
Anisomeridium phaeospermum ......................................129<br />
Anisomeridium planiusculum ..........................................149<br />
Anisomeridium polycarpum .............................................149<br />
Anisomeridium quadrococcum .......................................129<br />
Anisomeridium quaternarium ..........................................130<br />
Anisomeridium rockii........................................................149<br />
Anisomeridium sanfordense................................................128<br />
Anisomeridium sphaerocarpum ......................................149<br />
Anisomeridium stromaticum............................................149<br />
Anisomeridium subatomarium ........................................150<br />
Anisomeridium subbiforme..............................................150<br />
Anisomeridium subnectendum........................................150<br />
Anisomeridium subnexum ...............................................150<br />
Anisomeridium subprostans................................................130<br />
Anisomeridium tamarindi.....................................................130<br />
Anisomeridium tarmugliense...........................................150<br />
Anisomeridium terminatum..............................................130<br />
Anisomeridium throwerae ................................................150<br />
Anisomeridium truncatum................................................151<br />
Anisomeridium tuckerae......................................................131<br />
Anisomeridium ubianum ..................................................151<br />
Anisomeridium uniseriale.................................................151<br />
Anisomeridium verrucosum.............................................151<br />
Anisomeridium viridescens..............................................151<br />
Anthracothecium....................................................................93<br />
Anthracothecium albescens................................................109<br />
Anthracothecium albidopallens...........................................110<br />
Anthracothecium americanum ............................................110<br />
Anthracothecium canellae-albae...........................................94<br />
Anthracothecium columellatum.............................................87<br />
Anthracothecium confine.....................................................109<br />
Anthracothecium corticatum................................................109<br />
Anthracothecium nanum .......................................................94<br />
Anthracothecium prasinum .............................................87, 93<br />
Anthracothecium staurosporum ............................................94<br />
Anthracothecium subglobosum.............................................94<br />
Anthracothecium varians.......................................................93<br />
Anzia ......................................................................................40<br />
Anzia ornata...........................................................................40<br />
Apiospora polypori...............................................................128<br />
Architrypethelium.................................................................114<br />
Arthonia tumidula...................................................................68<br />
Arthoniales...............................................................................2<br />
Arthopyrenia ..........................................................................75<br />
Arthopyrenia sect. Anisomeridium .....................................123<br />
Arthopyrenia sect. Pseudosagedia ....................................176<br />
Arthopyrenia adnexa var. leptosperma...............................147<br />
Arthopyrenia adnexa ...........................................................143<br />
Arthopyrenia albidoatra .......................................................144<br />
Arthopyrenia alboatra..........................................................127<br />
Arthopyrenia amaura...........................................................127<br />
Arthopyrenia ambigua .........................................................126<br />
Arthopyrenia americana ......................................................144<br />
Arthopyrenia megalospora..............................................62, 82<br />
Arthopyrenia minor ................................................................82<br />
Arthopyrenia mycoporoides ..................................................69<br />
Arthopyrenia nidulans..........................................................148<br />
Arthopyrenia nieteriana .........................................................80<br />
Arthopyrenia nylanderi ..........................................................72<br />
172<br />
Arthopyrenia anacardii........................................................ 130<br />
Arthopyrenia analepta........................................................... 78<br />
Arthopyrenia anisoloba....................................................... 126<br />
Arthopyrenia annulata .......................................................... 79<br />
Arthopyrenia antecellens................................................ 67, 79<br />
Arthopyrenia antillarum....................................................... 144<br />
Arthopyrenia atomarioides.............................................. 62, 79<br />
Arthopyrenia atractospora .............................................. 62, 79<br />
Arthopyrenia bifera ......................................................... 79, 82<br />
Arthopyrenia biformis.......................................................... 127<br />
Arthopyrenia brachyspora .................................................. 127<br />
Arthopyrenia carinthiaca..................................................... 144<br />
Arthopyrenia carpinea ........................................................ 176<br />
Arthopyrenia cedrina............................................................. 79<br />
Arthopyrenia cerasi............................................................... 79<br />
Arthopyrenia ceuthocarpoides.............................................. 72<br />
Arthopyrenia ceylonensis ................................................... 144<br />
Arthopyrenia cinchonae........................................................ 80<br />
Arthopyrenia cinefaciens .................................................... 155<br />
Arthopyrenia cineropruinosa................................................. 79<br />
Arthopyrenia confluens................................................ 69, 80<br />
Arthopyrenia conformis....................................................... 127<br />
Arthopyrenia consanguinea.................................................. 69<br />
Arthopyrenia consimilis....................................................... 145<br />
Arthopyrenia consobrina var. glaucescens........................ 151<br />
Arthopyrenia consobrina..................................................... 145<br />
Arthopyrenia convexella ....................................................... 79<br />
Arthopyrenia degelii ........................................................... 80<br />
Arthopyrenia dimidiata........................................................ 144<br />
Arthopyrenia distans........................................................... 128<br />
Arthopyrenia endobrya ....................................................... 146<br />
Arthopyrenia epigloea........................................................... 72<br />
Arthopyrenia esenbeckiana......................................... 69, 81<br />
Arthopyrenia exasperata.................................................... 81<br />
Arthopyrenia excaecariae................................................... 128<br />
Arthopyrenia exstans.......................................................... 150<br />
Arthopyrenia fallax ................................................................ 79<br />
Arthopyrenia fallax f. crategina............................................. 79<br />
Arthopyrenia fallax var. conspurcata.................................... 80<br />
Arthopyrenia feeana ........................................................... 126<br />
Arthopyrenia floridana .......................................................... 62<br />
Arthopyrenia fraxini......................................................... 62, 81<br />
Arthopyrenia glaucescens .................................................. 146<br />
Arthopyrenia glaucina......................................................... 130<br />
Arthopyrenia griseola............................................................ 67<br />
Arthopyrenia incurva........................................................... 128<br />
Arthopyrenia infernalis........................................................ 147<br />
Arthopyrenia infernalis var. rhychostoma........................... 147<br />
Arthopyrenia insularis ......................................................... 130<br />
Arthopyrenia japonica........................................................... 80<br />
Arthopyrenia knysana......................................................... 126<br />
Arthopyrenia lapponina................................................... 78, 81<br />
Arthopyrenia leucochlora.................................................... 147<br />
Arthopyrenia ligustri.............................................................. 80<br />
Arthopyrenia limitans .......................................................... 144<br />
Arthopyrenia lyrata................................................................ 81<br />
Arthopyrenia majuscula ........................................................ 82<br />
Arthopyrenia malaccitula ...................................................... 82<br />
Arthopyrenia media............................................................. 144<br />
Arthopyrenia oblongens..................................................... 82<br />
Arthopyrenia padi............................................................ 63, 82<br />
Arthopyrenia parvula........................................................... 127<br />
Arthopyrenia pinicola ............................................................ 79<br />
Arthopyrenia planior.............................................................. 82<br />
Arthopyrenia planipes........................................................... 80
Arthopyrenia planiuscula.....................................................149<br />
Arthopyrenia planorbiculata ..................................................82<br />
Arthopyrenia planorbis ..........................................................82<br />
Arthopyrenia plumbaria .........................................................83<br />
Arthopyrenia pongamiae .....................................................149<br />
Arthopyrenia punctiformis .....................................................63<br />
Arthopyrenia quassiaecola....................................................63<br />
Arthopyrenia rappii ................................................................83<br />
Arthopyrenia rhyponta.....................................................63, 83<br />
Arthopyrenia salicis ...............................................................83<br />
Arthopyrenia sanfordensis .................................................128<br />
Arthopyrenia sanfordensis var. cedrina ...............................62<br />
Arthopyrenia saxicola ...........................................................63<br />
Arthopyrenia stigmatella var. elabens ..................................79<br />
Arthopyrenia subantecellens ................................................67<br />
Arthopyrenia subatomaria ..................................................150<br />
Arthopyrenia subbiformis ...................................................150<br />
Arthopyrenia subinsularis ...................................................130<br />
Arthopyrenia subnectenda .................................................150<br />
Arthopyrenia subnexa ........................................................150<br />
Arthopyrenia subprostans ..................................................130<br />
Arthopyrenia taxodii............................................................83<br />
Arthopyrenia terminata .......................................................130<br />
Arthopyrenia texensis............................................................83<br />
Arthopyrenia transposita ......................................................80<br />
Arthopyrenia truncata .........................................................151<br />
Arthopyrenia ubiana ...........................................................151<br />
Arthopyrenia viridescens ....................................................151<br />
Arthopyrenia willeyana .......................................................129<br />
Arthopyrenia xylogena ................................................123, 150<br />
Arthopyrenia zonata .............................................................73<br />
Arthopyreniaceae ..................................................................75<br />
Arthothelium tumidulum ........................................................68<br />
Ascocratera..........................................................................121<br />
Aspidopyrenium ..................................................................162<br />
Aspidotheliaceae ................................................................160<br />
Aspidothelium .....................................................................162<br />
Aspidothelium cinerascens ................................................165<br />
Aspidothelium trichotheliodes ............................................167<br />
Astrothelium.........................................................................114<br />
Astrothelium cinnamomeum................................................114<br />
Astrothelium confusum ...............................................115, 121<br />
Astrothelium diplocarpum....................................................115<br />
Astrothelium epiphyllum ......................................................133<br />
Astrothelium galbineum.......................................................114<br />
Astrothelium variolosum .............................................115, 121<br />
Astrothelium versicolor........................................................114<br />
Athrismidium..........................................................................65<br />
Bactrospora .............................................................................3<br />
Bactrospora brevispora ...........................................................4<br />
Bactrospora denticulata.......................................................3, 4<br />
Bactrospora dryina .................................................................4<br />
Bactrospora integrispora ........................................................4<br />
Bactrospora macrospora.........................................................4<br />
Bactrospora mesospora ..........................................................3<br />
Bactrospora myriadea .............................................................3<br />
Bactrospora nematospora.......................................................4<br />
Baeomyces ............................................................................28<br />
Baeomyces roseus ................................................................28<br />
Baeomycetaceae...................................................................28<br />
Canoparmelia caroliniana .....................................................40<br />
Canoparmelia cryptochlorophaea .........................................40<br />
Canoparmelia martinicana ....................................................40<br />
Canoparmelia salacinifera ....................................................40<br />
Canoparmelia texana ............................................................40<br />
Catapyrenium ........................................................................56<br />
Catapyrenium squamulosum ................................................56<br />
173<br />
Bathelium ............................................................................ 115<br />
Bathelium albidoporum.................................................... 116<br />
Bathelium carolinianum ................................................... 116<br />
Bathelium degenerans ..................................................... 117<br />
Bathelium endochryseum................................................ 117<br />
Bathelium lineare .............................................................. 117<br />
Bathelium madreporiforme ................................................. 117<br />
Bathelium mastoideum ....................................................... 117<br />
Bathelium sphaericum ..................................................... 118<br />
Bathelium tuberculosum.................................................. 118<br />
Beckhausia ........................................................................... 65<br />
Biatora leptalea................................................................... 175<br />
Bottaria.................................................................................. 65<br />
Bottaria albidopallens ......................................................... 110<br />
Bottaria composita ................................................................ 68<br />
Bottaria libricola f. nana ...................................................... 110<br />
Bottaria mucosa.................................................................. 110<br />
Buellia ................................................................................... 45<br />
Buellia amphidexia................................................................ 46<br />
Buellia caloosensis ............................................................... 46<br />
Buellia catasema .................................................................. 46<br />
Buellia coccinea.................................................................... 46<br />
Buellia curatellae .................................................................. 45<br />
Buellia curtisii........................................................................ 45<br />
Buellia imshaugiana ............................................................. 45<br />
Buellia lauricassiae............................................................... 46<br />
Buellia leucomela.................................................................. 45<br />
Buellia melanochlora ............................................................ 46<br />
Buellia pachnidisca............................................................... 46<br />
Buellia placodiomorpha ........................................................ 51<br />
Buellia rappii ......................................................................... 46<br />
Buellia rubifaciens................................................................. 47<br />
Buellia stillingiana ................................................................. 46<br />
Buellia wheeleri..................................................................... 47<br />
Bulbothrix .............................................................................. 40<br />
Bulbothrix confoederata........................................................ 40<br />
Bulbothrix coronata............................................................... 40<br />
Bulbothrix goebelii ................................................................ 40<br />
Bulbothrix isidiza ................................................................... 40<br />
Bulbothrix laevigatula............................................................ 40<br />
Byssoloma............................................................................. 31<br />
Byssoloma leucoblepharum ................................................. 31<br />
Byssoloma pubescens....................................................... 31<br />
Byssoloma subdiscordans.................................................... 31<br />
Byssoloma tricholomum........................................................ 31<br />
Byssothecium ....................................................................... 74<br />
Caliciaceae ........................................................................... 22<br />
Caliciales............................................................................... 20<br />
Calicium ................................................................................ 22<br />
Calicium adspersum ............................................................. 22<br />
Calicium chlorosporum ......................................................... 22<br />
Calicium hyperelloides.......................................................... 22<br />
Calicium lenticulare............................................................... 22<br />
Calicium leucochlorum ......................................................... 22<br />
Calicium salicinum................................................................ 22<br />
Campylothelium .................................................................. 113<br />
Campylothelium decolorans ................................................. 87<br />
Canoparmelia........................................................................ 40<br />
Canoparmelia amazonica..................................................... 40<br />
Catapyrenium tuckermanii.................................................... 56<br />
Celothelium ......................................................................... 161<br />
Celothelium aciculiferum .................................................... 162<br />
Celothelium cinchonarum................................................... 162<br />
Celothelium dominicanum .................................................. 161<br />
Celothelium leprieurii .......................................................... 161<br />
Celothelium socialis............................................................ 161
Chaenotheca .........................................................................24<br />
Chaenotheca brunneola........................................................24<br />
Chaenotheca floridana........................................................24<br />
Chaenotheca hispidula....................................................24, 25<br />
Chaenothecopsis.............................................................26, 27<br />
Chaenothecopsis nana .........................................................26<br />
Chaenothecopsis norstictica .......................................26, 27<br />
Chaenothecopsis pilosa ..................................................26, 27<br />
Chaenothecopsis rappii................................................26, 27<br />
Chaenothecopsis savonica ...................................................26<br />
Chaetothyriales......................................................................56<br />
Chiodecton...............................................................................4<br />
Chiodecton malmei..............................................................2, 4<br />
Chiodecton perplexum ............................................................4<br />
Chlorodothis ..........................................................................65<br />
Chromatochlamys................................................................162<br />
Chromatochlamys muscorum .............................................162<br />
Chromatochlamys muscorum var. octosporum ..................162<br />
Cifferiolichen..........................................................................75<br />
Cladina...................................................................................32<br />
Cladina evansii ......................................................................32<br />
Cladina rangiferina ................................................................32<br />
Cladina sandstedei................................................................32<br />
Cladina subtenuis..................................................................32<br />
Cladina subtenuis f. cinerea..................................................32<br />
Cladonia.................................................................................33<br />
Cladonia abbreviatula ...........................................................33<br />
Cladonia anitae......................................................................33<br />
Cladonia bacillaris .................................................................33<br />
Cladonia beaumontii..............................................................34<br />
Cladonia botryocarpa ............................................................34<br />
Cladonia buckii ......................................................................34<br />
Cladonia caroliniana..............................................................34<br />
Cladonia chlorophaea ...........................................................35<br />
Cladonia cristatella ................................................................33<br />
Cladonia cryptochlorophaea .................................................35<br />
Cladonia cylindrica ................................................................35<br />
Cladonia didyma var. didyma................................................33<br />
Cladonia didyma var. vulcanica ............................................33<br />
Cladonia dimorphoclada .......................................................34<br />
Cladonia floridana .................................................................34<br />
Cladonia grayi..................................................................34, 35<br />
Cladonia hypoxantha.............................................................33<br />
Cladonia incrassata...............................................................33<br />
Cladonia leporina ..................................................................33<br />
Cladonia nana .......................................................................36<br />
Cladonia pachycladodes .......................................................34<br />
Cladonia perforata.................................................................34<br />
Cladonia peziziformis ............................................................35<br />
Cladonia polycarpia...............................................................34<br />
Cladonia polycarpoides.........................................................34<br />
Cladonia prostrata .................................................................36<br />
Cladonia ramulosa ................................................................35<br />
Cladonia rappii var. exilior.....................................................35<br />
Cladonia rappii var. rappii .....................................................35<br />
Cladonia ravenelii..................................................................33<br />
Cladonia santensis ................................................................34<br />
Cladonia simulata..................................................................35<br />
Cladonia sobolescens ...........................................................35<br />
Cladonia strepsilis .................................................................35<br />
Diporina subsimplicans .......................................................152<br />
Dirinaria .................................................................................47<br />
Dirinaria aegialita...................................................................47<br />
Dirinaria applanata ................................................................47<br />
Dirinaria confluens.................................................................47<br />
Dirinaria confusa ...................................................................47<br />
Dirinaria leopoldii...................................................................47<br />
Dirinaria papillulifera..............................................................47<br />
174<br />
Cladonia subradiata.............................................................. 35<br />
Cladonia subsetacea ............................................................ 34<br />
Cladoniaceae........................................................................ 32<br />
Clathroporina ...................................................................... 171<br />
Clathroporina amygdalina .................................................... 88<br />
Clathroporina confinis......................................................... 171<br />
Clathroporina exiguella......................................................... 88<br />
Clathroporina isidiifera .................................................... 171<br />
Clathroporina mastoidea ................................................. 172<br />
Clathroporina nuculastrum ................................................. 174<br />
Clathroporina subpungens.............................................. 172<br />
Clathroporina tetracerae.................................................. 172<br />
Clypeolum talumae............................................................. 152<br />
Clypeopyrenis ....................................................................... 93<br />
Clypeosphaeriaceae ........................................................... 159<br />
Collema ................................................................................. 28<br />
Collema callibotrys................................................................ 29<br />
Collema conglomeratum....................................................... 29<br />
Collema furfurceum var. luzone ........................................... 29<br />
Collema leptaleum................................................................ 29<br />
Collema pulcellum var. pulcellum......................................... 28<br />
Collema pulcellum var. leucopeplum ................................... 28<br />
Collema pulcellum var. subnigrescens ................................ 28<br />
Collema pustulatum.............................................................. 29<br />
Collema subflaccidum .......................................................... 29<br />
Collemataceae...................................................................... 28<br />
Collemopsidium .................................................................... 71<br />
Collemopsidium atlanticum .................................................. 72<br />
Collemopsidium iocarpum .................................................... 72<br />
Compsosporiella ................................................................. 146<br />
Cresponea .............................................................................. 4<br />
Cresponea chloroconia........................................................... 5<br />
Cresponea flava...................................................................... 5<br />
Cresponea proximata ............................................................. 4<br />
Cryptothelium.............................................................. 114, 118<br />
Cyrtidula ................................................................................ 65<br />
Cyrtidula hippocastani ....................................................... 65<br />
Cyrtidula macularis ............................................................... 65<br />
Cyrtidula pteleodes ............................................................... 65<br />
Dacampia.............................................................................. 74<br />
Dacampiaceae...................................................................... 74<br />
Dermatina ohiensis............................................................... 69<br />
Dermatina pyrenocarpa ........................................................ 68<br />
Dermatocarpella ................................................................... 56<br />
Dibaeis baeomyces .............................................................. 28<br />
Dichoporis ........................................................................... 152<br />
Dichoporis schizospora....................................................... 152<br />
Didymella gigantea ............................................................... 82<br />
Didymella labiata ................................................................ 144<br />
Didymella laurilae ............................................................... 127<br />
Didymella microspora ......................................................... 127<br />
Didymella sphaerocarpa..................................................... 149<br />
Didymellopsis........................................................................ 71<br />
Didymosphaeria megalospora............................................ 148<br />
Diploschistes......................................................................... 16<br />
Diploschistes hypoleucus ..................................................... 16<br />
Diploschistes muscorum....................................................... 16<br />
Diporina............................................................................... 152<br />
Dirinaria picta ........................................................................ 47<br />
Dirinaria purpurascens ......................................................... 47<br />
Discosiella........................................................................... 152<br />
Discosiella cylindrospora.................................................... 152<br />
Distopyrenis .......................................................................... 94<br />
Distopyrenis americana ........................................................ 95<br />
Distopyrenis composita ..................................................... 95<br />
Distopyrenis fuliginosa.......................................................... 96
Distopyrenis pachyospora .....................................................96<br />
Distopyrenis quercicola......................................................96<br />
Distopyrenis submuriformis ..............................................96<br />
Ditremis................................................................................123<br />
Ditremis aggregata ..............................................................145<br />
Ditremis albiseda.................................................................126<br />
Ditremis ambigua.................................................................126<br />
Ditremis andamanica...........................................................149<br />
Ditremis anisoloba...............................................................126<br />
Ditremis anisomera .............................................................147<br />
Ditremis australiensis ..........................................................144<br />
Ditremis biformis..................................................................127<br />
Ditremis bispora...................................................................144<br />
Ditremis complanata............................................................145<br />
Ditremis distans ...................................................................128<br />
Ditremis epiphylla ................................................................146<br />
Ditremis finkii .......................................................................128<br />
Ditremis flavopallida ............................................................146<br />
Ditremis harrisii....................................................................145<br />
Ditremis immersa.................................................................147<br />
Ditremis indica .....................................................................147<br />
Ditremis inspersa.................................................................144<br />
Ditremis karnatakensis........................................................145<br />
Ditremis laevigata................................................................147<br />
Ditremis leucochlora............................................................147<br />
Ditremis macrospora ...................................................127, 147<br />
Ditremis microcarpa ............................................................151<br />
Ditremis monospora ............................................................148<br />
Ditremis muelleri..................................................................145<br />
Ditremis nyssaegena...........................................................128<br />
Ditremis pacifica ..................................................................149<br />
Ditremis polycarpa...............................................................149<br />
Ditremis punctata.................................................................149<br />
Ditremis pyrenuloides..........................................................144<br />
Ditremis quaternaria ............................................................130<br />
Ditremis rockii ......................................................................149<br />
Ditremis sanfordensis..........................................................128<br />
Ditremis straminicolor..........................................................145<br />
Ditremis subprostans...........................................................130<br />
Ditremis tamarindi................................................................130<br />
Ditremis tarmugliensis.........................................................150<br />
Ditremis terminata ...............................................................130<br />
Ditremis tuckerae.................................................................131<br />
Ditremis verrucosa.......................................................123, 151<br />
Endocarpon ...........................................................................57<br />
Endocarpon pusillum.............................................................57<br />
Enterographa...........................................................................5<br />
Enterographa lecanoroides.....................................................5<br />
Enterographa quassiaecola ....................................................5<br />
Eopyrenula.......................................................................74, 92<br />
Eopyrenula intermedia ..........................................................74<br />
Eopyrenula leucoplaca ..........................................................74<br />
Eopyrenula parvispora ..........................................................74<br />
Exiliseptum ..........................................................................114<br />
Foraminella............................................................................40<br />
Geisleria...............................................................................159<br />
Geisleria sychnogonioides ..................................................159<br />
Hafellia bahiana.....................................................................45<br />
Hafellia bahiana var. pleiotropa ............................................46<br />
Hafellia callispora ..................................................................45<br />
Hafellia parastata ..................................................................45<br />
Heterodermia.........................................................................47<br />
Heterodermia albicans ..........................................................47<br />
Heterodermia barbifera .........................................................48<br />
Heterodermia casarettiana....................................................48<br />
Heterodermia crocea.............................................................48<br />
Heterodermia echinata..........................................................48<br />
175<br />
Glyphis .................................................................................... 9<br />
Glyphis cicatricosa .................................................................. 9<br />
Granulopyrenis...................................................................... 97<br />
Granulopyrenis antillensis .................................................... 98<br />
Granulopyrenis hymnothora ................................................. 98<br />
Granulopyrenis macrocarpoides .......................................... 98<br />
Granulopyrenis nigrescens ............................................... 98<br />
Graphidaceae ......................................................................... 8<br />
Graphidales............................................................................. 8<br />
Graphina ................................................................................. 9<br />
Graphina abaphoides ............................................................. 9<br />
Graphina adscribens............................................................. 11<br />
Graphina anguina ................................................................. 10<br />
Graphina antillarum .............................................................. 10<br />
Graphina columbina.............................................................. 10<br />
Graphina columbina ............................................................. 11<br />
Graphina cypressi ................................................................... 9<br />
Graphina floridana ................................................................ 10<br />
Graphina incrustans.............................................................. 10<br />
Graphina intertexta ............................................................... 11<br />
Graphina leuconephela ........................................................ 10<br />
Graphina marescens ............................................................ 10<br />
Graphina parilis..................................................................... 10<br />
Graphina peplophora.............................................................. 9<br />
Graphina platyleuca.............................................................. 11<br />
Graphina scolecitis ............................................................... 10<br />
Graphina subvirginalis ...................................................... 9, 11<br />
Graphina virginea ................................................................... 9<br />
Graphina xylophaga.............................................................. 10<br />
Graphis.................................................................................. 11<br />
Graphis afzelii ....................................................................... 11<br />
Graphis anfractuosa ............................................................. 12<br />
Graphis anguinoides....................................................... 10, 13<br />
Graphis caesiella .................................................................. 12<br />
Graphis candidata................................................................. 11<br />
Graphis chromothecia........................................................... 12<br />
Graphis desquamescens...................................................... 12<br />
Graphis grammatis ............................................................... 13<br />
Graphis haleana ................................................................... 12<br />
Graphis illiterata.................................................................... 13<br />
Graphis insidiosa .................................................................. 13<br />
Graphis inversa..................................................................... 12<br />
Graphis leptocarpa ............................................................... 12<br />
Graphis librata....................................................................... 12<br />
Graphis lineola...................................................................... 11<br />
Graphis lucifica ..................................................................... 12<br />
Graphis lumbricina................................................................ 12<br />
Graphis pavoniana................................................................ 11<br />
Graphis rufula ....................................................................... 13<br />
Graphis scripta...................................................................... 11<br />
Graphis striatula.................................................................... 12<br />
Graphis subelegans.............................................................. 12<br />
Graphis subnitidula ............................................................... 13<br />
Graphis subparilis ................................................................. 13<br />
Graphis turbulenta ................................................................ 13<br />
Gyrostomum.......................................................................... 13<br />
Gyrostomum scyphuliferum.................................................. 13<br />
Hafellia ............................................................................ 45, 47<br />
Heterodermia galactophylla.................................................. 48<br />
Heterodermia japonica ......................................................... 48<br />
Heterodermia leucomelos..................................................... 48<br />
Heterodermia obscurata ....................................................... 48<br />
Heterodermia speciosa......................................................... 47<br />
Hyperphyscia ........................................................................ 48<br />
Hyperphyscia adglutinata ..................................................... 48<br />
Hyperphyscia minor .............................................................. 48<br />
Hyperphyscia syncolla .......................................................... 48<br />
Hypotrachyna ........................................................................ 41
Hypotrachyna livida ...............................................................41<br />
Hypotrachyna osseoalba.......................................................41<br />
Hypotrachyna pustulifera.......................................................41<br />
Icmadophilaceae ...................................................................28<br />
Imshaugia ..............................................................................41<br />
Imshaugia aleurites ...............................................................41<br />
Jarxia .....................................................................................60<br />
Jarxia ilicicola ......................................................................60<br />
Jarxia thelenula .....................................................................60<br />
Julella.....................................................................................84<br />
Julella asema .......................................................................86<br />
Julella avicenniae ..................................................................86<br />
Julella buxi .............................................................................86<br />
Julella dactylospora...............................................................87<br />
Julella decolorans ...............................................................87<br />
Julella dispora......................................................................87<br />
Julella exiguella ...................................................................87<br />
Julella fallaciosa ....................................................................87<br />
Julella geminella..................................................................87<br />
Julella leopoldina...................................................................87<br />
Julella luzonensis ..................................................................87<br />
Julella macrospora ................................................................88<br />
Julella manakonensis............................................................88<br />
Julella phycophila ..................................................................88<br />
Julella sericea........................................................................88<br />
Julella sublactea....................................................................88<br />
Julella taxodii .......................................................................88<br />
Julella variiformis ................................................................89<br />
Julella zenkeriana..................................................................90<br />
Jullela vitrispora.....................................................................90<br />
Lachrymospora......................................................................93<br />
Lauderlindsaya ......................................................................57<br />
Laurera ................................................................................118<br />
Laurera elegans...................................................................117<br />
Laurera linearis....................................................................117<br />
Laurera madreporiformis.....................................................117<br />
Laurera marginata ...............................................................117<br />
Laurera megasperma ....................................................90, 118<br />
Laurera pyriformis................................................................118<br />
Laurera sphaerica ...............................................................118<br />
Laurera subdisjuncta ...........................................................118<br />
Laurera submadreporiformis...............................................118<br />
Laurera tuberculosa ............................................................118<br />
Lecanactis................................................................................5<br />
Lecanactis epileuca.................................................................5<br />
Lecanactis subattingens..........................................................6<br />
Lecania fugiens ...................................................................165<br />
Lecanographa..........................................................................2<br />
Lecanorales ...........................................................................28<br />
Leiophloea .............................................................................75<br />
Leiophloea adnexa..............................................................143<br />
Leiophloea pongamiae........................................................149<br />
Leiophloea truncata.............................................................151<br />
Leiophloea xylogena ...........................................................150<br />
Lembidium ...........................................................................123<br />
Lembidium polycarpum .......................................................123<br />
Microtheliopsidaceae.............................................................56<br />
Monoblastia .........................................................................131<br />
Monoblastia borinquensis................................................132<br />
Monoblastia buckii...............................................................131<br />
Monoblastia cypressi ........................................................132<br />
Monoblastia echinulospora .................................................131<br />
Monoblastia palmicola.........................................................131<br />
Monoblastia pellucida..........................................................131<br />
Monoblastia quisqueyana.................................................132<br />
Monoblastia rappii ...............................................................131<br />
Monoblastiaceae .................................................................121<br />
176<br />
Leotiales................................................................................ 28<br />
Leptogium ............................................................................. 29<br />
Leptogium adpressum .......................................................... 30<br />
Leptogium austroamericanum.............................................. 29<br />
Leptogium azureum .............................................................. 29<br />
Leptogium chloromelum ....................................................... 30<br />
Leptogium corticola............................................................... 30<br />
Leptogium cyanescens......................................................... 29<br />
Leptogium floridanum ........................................................... 30<br />
Leptogium fusisporum .......................................................... 30<br />
Leptogium isidiosellum ......................................................... 29<br />
Leptogium juniperinum ......................................................... 29<br />
Leptogium marginellum ........................................................ 29<br />
Leptogium microstictum........................................................ 30<br />
Leptogium milligranum ......................................................... 29<br />
Leptogium phyllocarpum ...................................................... 30<br />
Leptogium sessile ................................................................. 30<br />
Leptogium stipitatum............................................................. 30<br />
Leptogium tenuissimum........................................................ 30<br />
Leptorhaphis ......................................................................... 60<br />
Leptorhaphis amygdali.......................................................... 60<br />
Leptorhaphis parameca........................................................ 60<br />
Lichen analeptus................................................................... 78<br />
Lichenothelia......................................................................... 73<br />
Lithothelium........................................................................... 98<br />
Lithothelium almbornii........................................................... 98<br />
Lithothelium bahamense .............................................. 98, 155<br />
Lithothelium concameratum ................................................. 99<br />
Lithothelium fugax................................................................. 99<br />
Lithothelium illotum............................................................... 99<br />
Lithothelium microsporum ................................................ 99<br />
Lithothelium obtectum........................................................... 99<br />
Lithothelium polysemum....................................................... 99<br />
Lophiostoma subcollapsa................................................... 112<br />
Luykenia.............................................................................. 162<br />
Mazaediothecium.................................................................. 93<br />
Mazosia................................................................................... 5<br />
Mazosia ocellata ..................................................................... 5<br />
Megalotremis .............................................................. 112, 123<br />
Megalotremis biocellata...................................................... 113<br />
Megalotremis megalospora ................................................ 148<br />
Megalotremis verrucosa ............................................. 112, 151<br />
Melanommatales .................................................................. 91<br />
Melanopsamma corticola.................................................... 129<br />
Melanotheca esenbeckiana.................................................. 81<br />
Melanotheca wrightii........................................................... 111<br />
Microglaena ........................................................................ 162<br />
Microglaena albolinita......................................................... 155<br />
Microglaena sychnogonioides............................................ 167<br />
Microglaena wallrothiana.................................................... 162<br />
Microglaenaceae ................................................................ 160<br />
Microthelia magnifica .......................................................... 148<br />
Microthelia palavana........................................................... 149<br />
Microthelia queenslandica.................................................. 113<br />
Microthelia uniserialis ......................................................... 151<br />
Musaespora ........................................................................ 133<br />
Musaespora corticola.......................................................... 133<br />
Musaespora epiphylla ...................................................... 133<br />
Musaespora gigas............................................................. 133<br />
Musaespora multilocularis.................................................. 133<br />
Mycocaliciaceae.................................................................... 26<br />
Mycocalicium................................................................... 26, 27<br />
Mycocalicium albonigrum ..................................................... 26<br />
Mycocalicium calicioides ...................................................... 27<br />
Mycocalicium rappii .............................................................. 27<br />
Mycocalicium ravenelii.................................................... 26, 27
Mycocalicium subtile..............................................................26<br />
Mycocifferia............................................................................75<br />
Mycoglaena collosporella......................................................88<br />
Mycoglaena yasudae ............................................................87<br />
Mycomicrothelia.....................................................................90<br />
Mycomicrothelia oleosa.........................................................90<br />
Mycomicrothelia rubrostoma .................................................90<br />
Mycomicrothelia subfallens ...................................................90<br />
Mycomicrothelia willeyana.....................................................90<br />
Mycoporaceae .......................................................................64<br />
Mycoporellum ..................................................................64, 65<br />
Mycoporellum californicum....................................................68<br />
Mycoporellum deserticola .....................................................69<br />
Mycoporellum difforme ..........................................................69<br />
Mycoporellum eschweileri .....................................................69<br />
Mycoporellum gigas ............................................................133<br />
Mycoporellum hassei.............................................................69<br />
Mycoporellum lacteum ..........................................................69<br />
Mycoporellum lahmii..............................................................69<br />
Mycoporellum leucoplacum...................................................69<br />
Mycoporellum sparsellum .....................................................70<br />
Mycoporopsis.........................................................................64<br />
Mycoporopsis antecellens.....................................................67<br />
Mycoporopsis californica.......................................................68<br />
Mycoporopsis deserticola......................................................69<br />
Mycoporopsis exigua.............................................................69<br />
Mycoporopsis leucoplaca......................................................69<br />
Mycoporopsis rappii ........................................................70, 83<br />
Mycoporum ............................................................................65<br />
Mycoporum acervatum .......................................................66<br />
Mycoporum antecellens......................................................67<br />
Mycoporum buckii ...............................................................68<br />
Mycoporum californicum....................................................68<br />
Mycoporum compositum .......................................................68<br />
Mycoporum difforme..............................................................69<br />
Mycoporum eschweileri......................................................69<br />
Mycoporum integrum.............................................................68<br />
Mycoporum lacteum............................................................69<br />
Mycoporum mycoporoides.................................................69<br />
Mycoporum ohiense ..............................................................69<br />
Mycoporum pycnocarpoides .................................................70<br />
Mycoporum pycnocarpum var. ohiense................................69<br />
Mycoporum pycnocarpum.....................................................68<br />
Mycoporum sparsellum .........................................................70<br />
Mycoporum uniloculatum...................................................70<br />
Mycopyrenula ......................................................................100<br />
Mycopyrenula coryli .............................................................100<br />
Mycosphaerellaceae .............................................................56<br />
Myelochroa ............................................................................41<br />
Myelochroa aurulenta............................................................41<br />
Myriangium duriaei ................................................................28<br />
Myriotrema.............................................................................16<br />
Myriotrema bahianum............................................................17<br />
Myriotrema compunctum.......................................................16<br />
Myriotrema erodens...............................................................17<br />
Ornatopyrenis ........................................................................90<br />
Parapyrenis..........................................................................100<br />
Parapyrenis aurora ..............................................................100<br />
Parapyrenis conica ..............................................................100<br />
Parapyrenis elongata ............................................................98<br />
Parapyrenis guayaci ............................................................100<br />
Parapyrenis maritima...........................................................100<br />
Parapyrenis tecomatis.........................................................100<br />
Parmeliaceae.........................................................................38<br />
Parmelinopsis ........................................................................41<br />
Parmelinopsis cryptochlora ...................................................41<br />
Parmelinopsis horrescens.....................................................41<br />
177<br />
Myriotrema glaucescens....................................................... 16<br />
Myriotrema glauculum .......................................................... 17<br />
Myriotrema peninsulae ......................................................... 17<br />
Myriotrema reclusum ............................................................ 17<br />
Myriotrema rugiferum............................................................ 17<br />
Myriotrema subcompunctum ................................................ 16<br />
Myriotrema terebratulum....................................................... 17<br />
Myriotrema wightii ................................................................. 17<br />
Nadvornikia ........................................................................... 17<br />
Nadvornikia hawaiiensis....................................................... 17<br />
Nadvornikia sorediata........................................................... 17<br />
Naetrocymbaceae ............................................................... 59<br />
Naetrocymbe......................................................................... 61<br />
Naetrocymbe atomarioides................................................ 62<br />
Naetrocymbe atractospora ................................................ 62<br />
Naetrocymbe cedrina ......................................................... 62<br />
Naetrocymbe fraxini ........................................................... 62<br />
Naetrocymbe massalongiana............................................ 62<br />
Naetrocymbe megalospora................................................ 62<br />
Naetrocymbe punctiformis ................................................ 63<br />
Naetrocymbe quassiaecola ............................................... 63<br />
Naetrocymbe rhyponta....................................................... 63<br />
Naetrocymbe saxicola........................................................ 63<br />
Neocatapyrenium.................................................................. 57<br />
Normandina pulchella........................................................... 57<br />
Ocellularia ............................................................................. 17<br />
Ocellularia americana........................................................... 18<br />
Ocellularia auberiana............................................................ 18<br />
Ocellularia cavata ................................................................. 18<br />
Ocellularia concolor .............................................................. 18<br />
Ocellularia emersa.......................................................... 17, 18<br />
Ocellularia fissa .................................................................... 18<br />
Ocellularia granulosa............................................................ 17<br />
Ocellularia leiostoma ............................................................ 18<br />
Ocellularia praestans............................................................ 18<br />
Ocellularia retispora.............................................................. 18<br />
Ocellularia sanfordiana......................................................... 18<br />
Ocellularia stictidea............................................................... 16<br />
Ochrolechia........................................................................... 51<br />
Ochrolechia africana............................................................. 51<br />
Ochrolechia antillarum.......................................................... 51<br />
Opegrapha .............................................................................. 6<br />
Opegrapha astraea................................................................. 8<br />
Opegrapha atra....................................................................... 6<br />
Opegrapha aurantiaca............................................................ 6<br />
Opegrapha candida ................................................................ 6<br />
Opegrapha cypressi................................................................ 6<br />
Opegrapha filicina................................................................... 8<br />
Opegrapha leucoplaca ........................................................... 7<br />
Opegrapha longissima............................................................ 7<br />
Opegrapha varia ..................................................................... 7<br />
Opegrapha viridis.................................................................... 7<br />
Opegrapha vulgata ................................................................. 7<br />
Opegraphaceae ...................................................................... 2<br />
Parmelinopsis minarum ........................................................ 41<br />
Parmelinopsis spumosa ....................................................... 41<br />
Parmeliopsis ......................................................................... 41<br />
Parmeliopsis subambigua .................................................... 41<br />
Parmotrema .......................................................................... 41<br />
Parmotrema crinitum ............................................................ 42<br />
Parmotrema cristiferum ........................................................ 42<br />
Parmotrema dilatatum .......................................................... 42<br />
Parmotrema dominicanum ................................................... 42<br />
Parmotrema endosulphureum.............................................. 41<br />
Parmotrema gardeneri.......................................................... 42<br />
Parmotrema hypoleucinum................................................... 42
Parmotrema mellissii .............................................................42<br />
Parmotrema michauxianum ..................................................42<br />
Parmotrema perforatum ........................................................42<br />
Parmotrema praesorediosum................................................42<br />
Parmotrema rampoddense ...................................................42<br />
Parmotrema rigidum..............................................................42<br />
Parmotrema rubifaciens ........................................................42<br />
Parmotrema sulphuratum......................................................42<br />
Parmotrema tinctorum...........................................................41<br />
Parmotrema ultralucens ........................................................42<br />
Parmotrema xanthinum .........................................................42<br />
Parmotrema zollingeri ...........................................................42<br />
Patellonectria.......................................................................162<br />
Peltosphaeria.........................................................................84<br />
Peltosphaeria californica .......................................................85<br />
Peltosphaeria sandwichensis................................................84<br />
Pertusaria ..............................................................................51<br />
Pertusaria amara ...................................................................52<br />
Pertusaria copiosa.................................................................51<br />
Pertusaria epixantha .............................................................52<br />
Pertusaria expolita.................................................................52<br />
Pertusaria floridana ...............................................................51<br />
Pertusaria hypothamnolica....................................................51<br />
Pertusaria iners .....................................................................52<br />
Pertusaria leucostoma...........................................................52<br />
Pertusaria multipunctoides....................................................52<br />
Pertusaria obruta ...................................................................53<br />
Pertusaria paratuberculifera..................................................52<br />
Pertusaria pulchella...............................................................51<br />
Pertusaria pustulata ..............................................................53<br />
Pertusaria sinusmexicani ......................................................52<br />
Pertusaria subpertusa ...........................................................53<br />
Pertusaria tetrathalamia ........................................................52<br />
Pertusaria texana ..................................................................52<br />
Pertusaria trachythallina........................................................52<br />
Pertusaria velata....................................................................52<br />
Pertusaria ventosa.................................................................51<br />
Pertusaria virensica...............................................................52<br />
Pertusaria xanthodes ............................................................53<br />
Pertusariaceae ......................................................................51<br />
Pertusariales..........................................................................51<br />
Phaeocalicium .......................................................................28<br />
Phaeocalicium polyporaeum.................................................28<br />
Phaeographina ......................................................................13<br />
Phaeographina asteroides ....................................................13<br />
Phaeographina caesiopruinosa ............................................14<br />
Phaeographina explicans......................................................14<br />
Phaeographina quassiaecola................................................14<br />
Phaeographis ........................................................................14<br />
Phaeographis dendritica .......................................................15<br />
Phaeographis erumpens .......................................................15<br />
Phaeographis haematites .....................................................14<br />
Phaeographis inusta..............................................................14<br />
Phaeographis lobata .............................................................15<br />
Phaeographis multicolor........................................................15<br />
Pleomassariaceae.................................................................74<br />
Pleosporales..........................................................................59<br />
Pleurothelium dissimulans ..................................................110<br />
Pleurotrema .........................................................................112<br />
Pleurotrema anisomerum....................................................147<br />
Pleurotrema epiphyllum ......................................................146<br />
Pleurotrema inspersum ...............................................123, 144<br />
Pleurotrema megaspermum................................................147<br />
Pleurotrema polycarpum .....................................................149<br />
Pleurotrema polysemum .......................................................99<br />
Pleurotrema pyrenuloides ...................................................144<br />
Pleurotrema rockii................................................................149<br />
Pleurotrema uniseriale ........................................................151<br />
178<br />
Phaeographis punctiformis................................................... 15<br />
Phaeographis sericea........................................................... 14<br />
Phaeographis subfulgurata .................................................. 15<br />
Phaeographis subtigrina....................................................... 15<br />
Phaeopeltosphaeria irregularis .......................................... 100<br />
Phaeophyscia ....................................................................... 48<br />
Phaeophyscia cernohorskyi.................................................. 48<br />
Phaeophyscia chloantha ...................................................... 48<br />
Phaeophyscia ciliata ............................................................. 48<br />
Phaeophyscia pusilloides ..................................................... 48<br />
Phaeophyscia rubropulchra.................................................. 48<br />
Phyllobatheliaceae.............................................................. 151<br />
Phyllobathelium................................................................... 152<br />
Phyllobathelium albolinitum................................................ 155<br />
Phyllobathelium obtectum .................................................. 158<br />
Phylloporina obducta .......................................................... 158<br />
Phylloporina octomera........................................................ 176<br />
Phylloporis .......................................................................... 152<br />
Phylloporis multipunctata.................................................... 158<br />
Phylloporis obducta ............................................................ 158<br />
Phylloporis phyllogena................................................ 152, 159<br />
Phylloporis platypoda ......................................................... 159<br />
Phylloporis viridis ................................................................ 160<br />
Phyllopsora ........................................................................... 36<br />
Phyllopsora buettneri var. glauca ......................................... 38<br />
Phyllopsora buettneri var. munda......................................... 38<br />
Phyllopsora confusa ............................................................. 36<br />
Phyllopsora corallina var. corallina................................. 36, 37<br />
Phyllopsora corallina var. glaucella...................................... 38<br />
Phyllopsora corallina var. ochroxantha ................................ 38<br />
Phyllopsora corallina var. rappiana ...................................... 37<br />
Phyllopsora corallina var. santensis..................................... 38<br />
Phyllopsora furfuracea.......................................................... 37<br />
Phyllopsora halei................................................................... 37<br />
Phyllopsora isidiolyta ............................................................ 37<br />
Phyllopsora kalbii.................................................................. 37<br />
Phyllopsora parvifolia var. breviuscula........................... 36, 37<br />
Phyllopsora parvifolia var. parvifolia..................................... 36<br />
Phyllopsora parvifoliella........................................................ 36<br />
Physcia.................................................................................. 49<br />
Physcia aipolia ...................................................................... 49<br />
Physcia americana................................................................ 49<br />
Physcia atrostriata ................................................................ 49<br />
Physcia crispa....................................................................... 49<br />
Physcia neogaea .................................................................. 49<br />
Physcia poncinsii .................................................................. 49<br />
Physcia pumilior.................................................................... 49<br />
Physcia sorediosa................................................................. 49<br />
Physcia undulata................................................................... 49<br />
Physciaceae.......................................................................... 44<br />
Physma byrsaeum ................................................................ 28<br />
Pilocarpaceae ....................................................................... 30<br />
Pleomassaria ........................................................................ 74<br />
Pleurotremataceae ............................................................. 112<br />
Polyblastia dispora................................................................ 87<br />
Polyblastia fallaciuscula........................................................ 87<br />
Polyblastia transwaalensis ................................................... 88<br />
Polyblastiopsis alboatra........................................................ 87<br />
Polyblastiopsis dispora ......................................................... 87<br />
Polyblastiopsis fallaciuscula ................................................. 87<br />
Polyblastiopsis geminella ..................................................... 87<br />
Polyblastiopsis linearis ....................................................... 117<br />
Polyblastiopsis obtecta ....................................................... 158<br />
Polyblastiopsis pyriformis ................................................... 118<br />
Polyblastiopsis rappii ............................................................ 87
Polyblastiopsis sphaerica....................................................118<br />
Polyblastiopsis tagananae ..................................................160<br />
Polycornum..........................................................................175<br />
Polymeridium.......................................................................118<br />
Polymeridium albidum .........................................................119<br />
Polymeridium albocinereum................................................119<br />
Polymeridium catapastum...................................................118<br />
Polymeridium contendens...................................................119<br />
Polymeridium proponens.....................................................119<br />
Polymeridium quinqueseptatum..........................................119<br />
Polymeridium subcinereum.................................................119<br />
Polypyrenula ..........................................................................92<br />
Porina...................................................................................172<br />
Porina aenea .......................................................................178<br />
Porina albicascens ..............................................................154<br />
Porina amygdalina...............................................................178<br />
Porina cestrensis.................................................................178<br />
Porina cestrensis var. platyspora........................................178<br />
Porina chlorotica..................................................................179<br />
Porina concamerata ..............................................................80<br />
Porina elaeidis.....................................................................130<br />
Porina guaranitica ...............................................................173<br />
Porina guentheri ..................................................................179<br />
Porina heterospora..............................................................173<br />
Porina hibernica...................................................................180<br />
Porina indutula.....................................................................157<br />
Porina interspersa ...............................................................130<br />
Porina leptalea.....................................................................175<br />
Porina linearis......................................................................180<br />
Porina mastoidea.................................................................172<br />
Porina multipunctata............................................................158<br />
Porina mundula ...................................................................158<br />
Porina nitidula......................................................................180<br />
Porina nucula.......................................................................174<br />
Porina nuculastrum...........................................................174<br />
Porina octomera ..................................................................176<br />
Porina olivacea var. microspora..........................................175<br />
Porina phyllogena................................................................159<br />
Porina platypoda..................................................................159<br />
Porina plumbaria ...................................................................83<br />
Porina rhaphidosperma.......................................................180<br />
Porina rubentior...................................................................176<br />
Porina salicina .....................................................................178<br />
Porina scabrida..................................................................174<br />
Porina subpungens .............................................................172<br />
Porina tamarindi ..................................................................130<br />
Porina tetracerae.................................................................172<br />
Porina thaxteri .....................................................................180<br />
Porina vaga..........................................................................130<br />
Porina viridiseda f. albiseda ................................................126<br />
Porinopsis............................................................................162<br />
Porinopsis geminipara.........................................................165<br />
Pseudoparmelia.....................................................................43<br />
Pyrenula cuyabensis ...........................................................103<br />
Pyrenula dissimulans........................................................110<br />
Pyrenula erumpens .............................................................103<br />
Pyrenula falsaria..................................................................102<br />
Pyrenula ferax .....................................................................111<br />
Pyrenula harrisii...................................................................110<br />
Pyrenula herrei ......................................................................83<br />
Pyrenula laetior....................................................................109<br />
Pyrenula laevigata ...............................................................109<br />
Pyrenula leucostoma...........................................................102<br />
Pyrenula macularis..............................................................110<br />
Pyrenula mamillana.............................................................110<br />
Pyrenula marginata .............................................................110<br />
Pyrenula martinicana...........................................................103<br />
179<br />
Pseudoparmelia sphaerospora ............................................ 43<br />
Pseudopyrenula .................................................................. 119<br />
Pseudopyrenula degenerans ............................................. 117<br />
Pseudopyrenula endochrysea............................................ 117<br />
Pseudopyrenula subgregaria ............................................. 119<br />
Pseudopyrenula subnudata................................................ 119<br />
Pseudosagedia ................................................................... 176<br />
Pseudosagedia aenea........................................................ 178<br />
Pseudosagedia chlorotica .................................................. 179<br />
Pseudosagedia guentheri................................................... 179<br />
Pseudosagedia linearis ...................................................... 180<br />
Pseudosagedia nitidula ...................................................... 180<br />
Pseudosagedia thaxteri ...................................................... 180<br />
Punctelia ............................................................................... 43<br />
Punctelia rudecta .................................................................. 43<br />
Pyrenidium ............................................................................ 74<br />
Pyrenidium octosporum...................................................... 162<br />
Pyrenocollema ...................................................................... 71<br />
Pyrenocollema atlanticum ................................................. 72<br />
Pyrenocollema caesium........................................................ 72<br />
Pyrenocollema ceuthocarpoides ...................................... 72<br />
Pyrenocollema epigloeum ................................................. 72<br />
Pyrenocollema farlowii....................................................... 72<br />
Pyrenocollema prospersellum .............................................. 72<br />
Pyrenocollema saxicola........................................................ 63<br />
Pyrenocollema strontianense ............................................... 73<br />
Pyrenocollema tichothecioides............................................. 72<br />
Pyrenocollema tremelloides ................................................. 72<br />
Pyrenocollema zonatum..................................................... 73<br />
Pyrenographa ..................................................................... 100<br />
Pyrenographa irregularis ................................................. 100<br />
Pyrenographa xylographoides............................................ 100<br />
Pyrenowilmsia....................................................................... 93<br />
Pyrenula .............................................................................. 101<br />
Pyrenula acutalis................................................................. 104<br />
Pyrenula anomala............................................................... 105<br />
Pyrenula aspistea ............................................................... 106<br />
Pyrenula astroidea.............................................................. 101<br />
Pyrenula atrolaminata ...................................................... 107<br />
Pyrenula caryae................................................................. 107<br />
Pyrenula cerina ................................................................... 104<br />
Pyrenula circumrubens....................................................... 109<br />
Pyrenula citriformis ............................................................. 104<br />
Pyrenula cocoes ................................................................. 105<br />
Pyrenula concastroma ..................................................... 108<br />
Pyrenula concatervans ....................................................... 109<br />
Pyrenula confinis .............................................................. 109<br />
Pyrenula confoederata........................................................ 106<br />
Pyrenula corticata ............................................................... 109<br />
Pyrenula cruenta................................................................. 109<br />
Pyrenula cruentata.............................................................. 101<br />
Pyrenula crystalligera ......................................................... 109<br />
Pyrenula cubana ................................................................. 103<br />
Pyrenula mastoidea............................................................ 172<br />
Pyrenula microcarpa........................................................... 106<br />
Pyrenula microtheca ........................................................... 103<br />
Pyrenula mucosa .............................................................. 110<br />
Pyrenula nitidula ................................................................. 110<br />
Pyrenula occidentalis.......................................................... 110<br />
Pyrenula ochraceoflava ...................................................... 101<br />
Pyrenula ochraceoflavens .................................................. 101<br />
Pyrenula oleosa .................................................................. 102<br />
Pyrenula personata............................................................. 103<br />
Pyrenula pseudobufonia..................................................... 105<br />
Pyrenula punctella .............................................................. 111<br />
Pyrenula punctiformis var. cinereopruinosa f. buxicola ....... 79<br />
Pyrenula punctiformis var. cinereopruinosa f. hederae ....... 79
Pyrenula punctiformis var. cinereopruinosa f. pinicola.........79<br />
Pyrenula punctiformis var. fallax f. betulae ..........................79<br />
Pyrenula pyrenuloides.........................................................111<br />
Pyrenula quassiaecola ........................................................104<br />
Pyrenula ravenelii ................................................................101<br />
Pyrenula rubromaculata ......................................................109<br />
Pyrenula rubrostoma ...........................................................106<br />
Pyrenula santensis ..............................................................105<br />
Pyrenula septicollaris ..........................................................103<br />
Pyrenula subferruginea .......................................................103<br />
Pyrenula tenuisepta.............................................................106<br />
Pyrenula thelomorpha .........................................................102<br />
Pyrenula wetmorei...............................................................103<br />
Pyrenula wheeleri ................................................................103<br />
Pyrenula wrightii................................................................111<br />
Pyrenula zwackhii..................................................................67<br />
Pyrenulaceae...................................................................74, 91<br />
Pyrenyllium ............................................................................75<br />
Pyrgillus ...............................................................................111<br />
Pyrgillus javanicus.................................................................93<br />
Pyxine ....................................................................................50<br />
Pyxine albovirens ..................................................................50<br />
Pyxine berteriana...................................................................50<br />
Pyxine caesiopruinosa ..........................................................50<br />
Pyxine cocoes........................................................................50<br />
Pyxine coralligera ..................................................................50<br />
Pyxine eschweileri .................................................................50<br />
Pyxine sorediata ....................................................................50<br />
Pyxine subcinerea .................................................................50<br />
Raciborskiella ......................................................................152<br />
Relicina ..................................................................................43<br />
Relicina abstrusa...................................................................43<br />
Relicina eximbricata ..............................................................43<br />
Requienella..........................................................................112<br />
Requienella lichenopsis ......................................................112<br />
Requienella seminuda.........................................................112<br />
Requienella subcollapsa ..................................................112<br />
Requienellaceae..................................................................112<br />
Rimelia...................................................................................43<br />
Rimelia reticulata...................................................................43<br />
Rimelia subisidiosa................................................................43<br />
Rimeliella ...............................................................................43<br />
Rimeliella neotropica.............................................................43<br />
Rimeliella subtinctoria ...........................................................43<br />
Rinodina.................................................................................50<br />
Rinodina applanata ...............................................................50<br />
Rinodina granuligera .............................................................51<br />
Rinodina lepida......................................................................51<br />
Sagedia excaecariae...........................................................128<br />
Sagedia massalongiana........................................................62<br />
Sagedia nylanderi..................................................................72<br />
Sagedia nylanderi β. hudsoniana..........................................72<br />
Sagediomyces .....................................................................152<br />
Strigula nitidula ....................................................................152<br />
Strigula obducta ................................................................158<br />
Strigula obtecta..................................................................158<br />
Strigula phaea .....................................................................158<br />
Strigula phyllogena ...................................................152, 159<br />
Strigula platypoda .............................................................159<br />
Strigula rostrata.................................................................159<br />
Strigula smaragdula ....................................................152, 159<br />
Strigula stigmatella ..............................................................154<br />
Strigula subelegans.............................................................159<br />
Strigula subsimplicans.........................................................152<br />
Strigula sychnogonioides ....................................................159<br />
Strigula tagananae ............................................................160<br />
Strigula viridis....................................................................160<br />
180<br />
Sagediomyces affinis.......................................................... 152<br />
Santessoniolichen................................................................. 61<br />
Sarcinulella ......................................................................... 123<br />
Sarcinulella banksiae.......................................................... 123<br />
Sarcographa ......................................................................... 14<br />
Sarcographa intricans........................................................... 14<br />
Sarcographa labyrinthica...................................................... 14<br />
Sarcographa medusulina...................................................... 15<br />
Sarcographa tricosa.............................................................. 14<br />
Sarcographia labyrinthica ..................................................... 15<br />
Schismatomma ....................................................................... 8<br />
Schismatomma rappii ......................................................... 5, 8<br />
Schizoxylon berkelyeanum ..................................................... 4<br />
Sciodothis.............................................................................. 65<br />
Sclerophyton ........................................................................... 8<br />
Sclerophyton elegans ............................................................. 8<br />
Sclerophyton inscriptum ......................................................... 8<br />
Scleropyrenium ..................................................................... 57<br />
Secolegiella......................................................................... 162<br />
Segestria ............................................................................. 175<br />
Segestria lectissima............................................................ 175<br />
Segestria leptalea ............................................................. 175<br />
Segestria mammillosa ........................................................ 176<br />
Segestria octomera .......................................................... 176<br />
Segestria rubentior........................................................... 176<br />
Shanoria.............................................................................. 152<br />
Shanoria indica ................................................................... 152<br />
Sphaeria cavernosa.............................................................. 99<br />
Sphinctrina ............................................................................ 25<br />
Sphinctrina tubiformis ........................................................... 25<br />
Sphinctrinaceae .................................................................... 25<br />
Sphinctrinella ........................................................................ 27<br />
Splanchnonema.................................................................... 74<br />
Sporoschizon ........................................................................ 61<br />
Strigula ................................................................................ 152<br />
Strigula affinis ..................................................................... 152<br />
Strigula albicascens ......................................................... 154<br />
Strigula albolinitum .......................................................... 155<br />
Strigula bahamensis......................................................... 155<br />
Strigula bermudana .......................................................... 155<br />
Strigula cinefaciens.......................................................... 155<br />
Strigula connivens............................................................ 155<br />
Strigula elegans .................................................................. 159<br />
Strigula fracticonidia ........................................................ 156<br />
Strigula griseonitens ........................................................ 156<br />
Strigula hypothallina......................................................... 157<br />
Strigula indutula................................................................ 157<br />
Strigula jamesii.................................................................... 157<br />
Strigula laceribracae ........................................................ 158<br />
Strigula mediterranea ......................................................... 152<br />
Strigula multipunctata...................................................... 158<br />
Strigula viridiseda ............................................................... 160<br />
Strigulaceae ........................................................................ 151<br />
Strigulomyces ..................................................................... 152<br />
Strigulomyces elegans ....................................................... 152<br />
Syngenosorus....................................................................... 65<br />
Thelenella ........................................................................... 162<br />
Thelenella brasiliensis ........................................................ 164<br />
Thelenella cinerascens .................................................... 165<br />
Thelenella cuyabense......................................................... 165<br />
Thelenella elegans.............................................................. 117<br />
Thelenella fugiens............................................................. 165<br />
Thelenella geminipara ...................................................... 165<br />
Thelenella harrisii................................................................ 167<br />
Thelenella hassei................................................................ 164
Thelenella humilis..............................................................165<br />
Thelenella inductula ............................................................164<br />
Thelenella luridella ..............................................................166<br />
Thelenella modesta .....................................................162, 163<br />
Thelenella obtecta ...............................................................158<br />
Thelenella pertusariella .......................................................166<br />
Thelenella rappii ................................................................166<br />
Thelenella sastreana .........................................................166<br />
Thelenella sordidula ............................................................164<br />
Thelenella sychnogonioides ............................................167<br />
Thelenella trichothelioides ...............................................167<br />
Thelenella weberi ................................................................164<br />
Thelenellaceae ....................................................................160<br />
Thelidium ...............................................................................57<br />
Thelidium farlowii...................................................................72<br />
Thelidium juistense..............................................................129<br />
Thelidium minutulum .............................................................57<br />
Thelidium nylanderi ...............................................................72<br />
Thelidium nylanderi var. hudsoniana ....................................72<br />
Thelomma..............................................................................25<br />
Thelomma carolinianum........................................................25<br />
Thelotrema.............................................................................19<br />
Thelotrema adjectum.............................................................19<br />
Thelotrema alborosellum.......................................................19<br />
Thelotrema defectum ............................................................20<br />
Thelotrema dilatatum.............................................................19<br />
Thelotrema eximium..............................................................19<br />
Thelotrema floridense............................................................19<br />
Thelotrema halei....................................................................19<br />
Thelotrema lacteum...............................................................20<br />
Thelotrema lathraeum ...........................................................19<br />
Thelotrema lepadinum...........................................................20<br />
Thelotrema leprocarpum .......................................................20<br />
Thelotrema monospermum ...................................................20<br />
Thelotrema platycarpoides....................................................20<br />
Thelotrema porinoides...........................................................19<br />
Thelotrema santense.............................................................19<br />
Thelotrema subtile.................................................................19<br />
Thelotremataceae..................................................................16<br />
Tomasellia .............................................................................64<br />
Tomasellia africana .............................................................130<br />
Tomasellia americana ...........................................................64<br />
Tomasellia angulosa ...........................................................144<br />
Tomasellia cubana ................................................................69<br />
Tomasellia dispora ..............................................................113<br />
Tomasellia eschweileri ..........................................................69<br />
Tomasellia esenbeckiana................................................69, 81<br />
Tomasellia exumana .............................................................69<br />
Tomasellia lactea...................................................................69<br />
Tomasellia leucostoma....................................................69, 80<br />
Tomasellia macularis ..........................................................65<br />
Tomasellia nigrescens...........................................................98<br />
Tomasellia queenslandica...................................................130<br />
Usnea dimorpha ....................................................................44<br />
Usnea mutabilis.....................................................................43<br />
Usnea perplectata .................................................................43<br />
Usnea rubicunda ...................................................................44<br />
Usnea strigosa.......................................................................43<br />
Usnea subscabrosa...............................................................44<br />
Usnea trichodea ....................................................................44<br />
Verrucaria ..............................................................................58<br />
Verrucaria*Leiophloea...........................................................75<br />
Verrucaria aenea.................................................................178<br />
Verrucaria albescens...........................................................109<br />
Verrucaria albicascens........................................................154<br />
Verrucaria albidoatra...........................................................144<br />
Verrucaria alboatra..............................................................127<br />
181<br />
Tomasellia sparsella............................................................. 70<br />
Trichotheliaceae ................................................................. 168<br />
Trichotheliales..................................................................... 168<br />
Trichothelium ...................................................................... 176<br />
Trichothelium aeneum ............................................. 176, 178<br />
Trichothelium cestrense .................................................. 178<br />
Trichothelium chloroticum .............................................. 179<br />
Trichothelium crocynioides............................................. 179<br />
Trichothelium guentheri................................................... 179<br />
Trichothelium horridulum.................................................... 178<br />
Trichothelium isidiatum ................................................... 180<br />
Trichothelium lineare........................................................ 180<br />
Trichothelium nitidulum ................................................... 180<br />
Trichothelium rhaphidospermum ................................... 180<br />
Trichothelium thaxteri ...................................................... 180<br />
Trypetheliaceae .................................................................. 112<br />
Trypethelium ....................................................................... 119<br />
Trypethelium aeneum ......................................................... 120<br />
Trypethelium albidoporum.................................................. 116<br />
Trypethelium carolinianum ................................................. 116<br />
Trypethelium degenerans................................................... 117<br />
Trypethelium eluteriae ........................................................ 119<br />
Trypethelium eluteriae var. polystomum ............................ 120<br />
Trypethelium exiguellum....................................................... 87<br />
Trypethelium feei ................................................................ 117<br />
Trypethelium ferrugineum................................................... 120<br />
Trypethelium ferrugineum β. inornatum ............................. 120<br />
Trypethelium floridanum ..................................................... 120<br />
Trypethelium karnatakense ............................................. 147<br />
Trypethelium leprieurii ........................................................ 121<br />
Trypethelium madreporiforme ............................................ 117<br />
Trypethelium madreporiforme f. pruinosa .......................... 117<br />
Trypethelium marginatum................................................... 117<br />
Trypethelium nitidiusculum......................................... 120, 121<br />
Trypethelium ochroleucum ......................................... 120, 121<br />
Trypethelium phlyctaena .................................................... 117<br />
Trypethelium quassiaecola................................................. 117<br />
Trypethelium scoria............................................................. 117<br />
Trypethelium subeluteriae .................................................. 120<br />
Trypethelium tropicum ........................................................ 120<br />
Trypethelium variolosum ............................................ 114, 121<br />
Trypethelium virens............................................................. 120<br />
Tuckermanopsis.................................................................... 43<br />
Tuckermanopsis fendleri ...................................................... 43<br />
Tuckermanopsis viridis ......................................................... 43<br />
Tylophoron ............................................................................ 25<br />
Tylophoron moderatum......................................................... 25<br />
Tylophoron protrudens.......................................................... 25<br />
Usnea.................................................................................... 43<br />
Usnea baileyi ........................................................................ 43<br />
Usnea ceratina...................................................................... 43<br />
Verrucaria alboatra var. detergens....................................... 80<br />
Verrucaria albolinita............................................................ 155<br />
Verrucaria analepta var. americana ................................... 110<br />
Verrucaria anisomera ......................................................... 150<br />
Verrucaria antecellens.......................................................... 67<br />
Verrucaria bermudana........................................................ 155<br />
Verrucaria biformis.............................................................. 127<br />
Verrucaria calkinsiana .......................................................... 59<br />
Verrucaria cerasi................................................................... 79<br />
Verrucaria cestrensis.......................................................... 178<br />
Verrucaria chlorotica........................................................... 179<br />
Verrucaria cinchonae............................................................ 80<br />
Verrucaria cinefaciens ........................................................ 155<br />
Verrucaria cinereopruinosa .................................................. 79
Verrucaria circumrubens .....................................................109<br />
Verrucaria clandestina.........................................................147<br />
Verrucaria concamerata........................................................80<br />
Verrucaria confinis...............................................................109<br />
Verrucaria conformis ...........................................................127<br />
Verrucaria consobrina .........................................................145<br />
Verrucaria consobrina var. glaucescens.............................151<br />
Verrucaria distans................................................................127<br />
Verrucaria epidermidis var. fallax..........................................79<br />
Verrucaria epidermidis var. γ. quassiaecola .........................63<br />
Verrucaria epigloea ...............................................................72<br />
Verrucaria excellens............................................................146<br />
Verrucaria fallax.....................................................................79<br />
Verrucaria geminella .............................................................87<br />
Verrucaria guentheri............................................................179<br />
Verrucaria hippocastani ........................................................65<br />
Verrucaria holopolia ............................................................147<br />
Verrucaria indutula ..............................................................157<br />
Verrucaria infernalis ............................................................147<br />
182<br />
Verrucaria interspersa ........................................................ 130<br />
Verrucaria lactea................................................................... 69<br />
Verrucaria limitans .............................................................. 144<br />
Verrucaria linearis............................................................... 180<br />
Verrucaria magnifica........................................................... 148<br />
Verrucaria magnospora ...................................................... 148<br />
Verrucaria majuscula............................................................ 82<br />
Verrucaria malaccitula .......................................................... 82<br />
Verrucaria mamillaris............................................................ 59<br />
Verrucaria muralis................................................................. 58<br />
Verrucaria planiuscula ........................................................ 149<br />
Verrucaria planorbis.............................................................. 82<br />
Verrucaria plumbaria ............................................................ 83<br />
Verrucaria prostans .............................................................. 80<br />
Verrucaria punctiformis......................................................... 63<br />
Verrucaria rhyponta .............................................................. 63<br />
Verrucaria riddleana ........................................................... 59<br />
Verrucaria rubentior............................................................ 176<br />
Verrucaria socialis .............................................................. 161<br />
Verrucaria stigmatella var. lactea ......................................... 69<br />
Verrucaria subantecellens.................................................... 67<br />
Verrucaria subatomaria ...................................................... 150<br />
Verrucaria subbiformis........................................................ 150<br />
Verrucaria sublactea............................................................. 88<br />
Verrucaria subnectenda ..................................................... 150<br />
Verrucaria subnexa............................................................. 150<br />
Verrucaria subprostans ...................................................... 130<br />
Verrucaria subpunctiformis................................................... 62<br />
Verrucaria tagananae ......................................................... 160<br />
Verrucaria tamarindi ........................................................... 130<br />
Verrucaria terminata ........................................................... 130<br />
Verrucaria tetracerae.......................................................... 172<br />
Verrucaria thelena var. albidior .......................................... 143<br />
Verrucaria vaga................................................................... 130<br />
Verrucaria viridiseda f. albiseda ......................................... 126<br />
Verrucariaceae...................................................................... 56<br />
Verrucariales......................................................................... 56<br />
Vulpicida................................................................................ 44<br />
Vulpicida viridis ..................................................................... 44<br />
Xanthoparmelia..................................................................... 44<br />
Xanthoparmelia conspersa................................................... 44<br />
Xanthopyrenia....................................................................... 72<br />
Xanthopyreniaceae............................................................... 70<br />
Zamenhofia ......................................................................... 176<br />
Zignoella nyssaegena......................................................... 128<br />
Zwackhiomyces..................................................................... 73<br />
Zwackhiomyces arenicola ................................................. 73