A TAXONOMIC RE‘flSlON OF THE GENUS
ARYHOPYRENEA MASSAL. S. LAT.
(ASCOMYCETFS) IN NORTH AMERl‘CA

Mssertation for the Degree of Ph. D.
MiCHlGAN STATE UNIVERSHY
RICHARD CLINTON HARRlS
197.5

 

 

This is to certify that the

thesis entitled

A TAXONOMIC REVISION OF THE GENUS
ARTHOPYRENIA MASSAL. S. LAT.
(ASCOMYCETES) IN NORTH AMERICA

presented by

RI CHARD CLI NTON HARRIS

has been accepted towards fulfillment
of the requirements for

. . . Botan
Ph D Jegree in y

 

/%-\ flflflfl
/ Wk

Major professor

Date 3 July 1975

 

0-7 639 ,'

    
       
   

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'4 ABSTRACT

A TAXONOMIC REVISION OF THE GBNUS
ARTHOPYRENIA MASSAL. S. LAT. (ASCOMYCETES) IN NORTH AMERICA
BY

Richard Clinton Harris

While studying large numbers of specimens of the pyrenolichen genus

Arthopyrenia preparatory to a monograph of the North American species,

 

it became clear that this was impossible without first clarifying famil-
ial and generic boundaries within the pyrenolichens. For this purpose
a wide variety of additional material was examined and pertinent char-
acters were recorded with drawings. These morphological studies, and
chemical studies where relevant, have resulted in suggestions for minor
reorganization of the families of pyrenolichens and in a drastic revi-
sion of generic boundaries. After a preliminary assessment of the tax-
onomic characters used, a key is given to the families and the genera
included in them are listed. This is followed by a key to the genera
of hyalodidymous and hyalophragmous pyrenolichens of North America
(excluding Verrucariaceae). In the subsequent systematic section
Arthopyrenia is redefined and many species are placed in segregate
genera and combined with species described in other genera resulting in

the following groupings: Pleosporaceae Wint. (Arthopyrenia Massal.,

 

Mycoglaena v. Hahn. and Tomasellia Massal.), Strigulaceae Fr. (Acro-

 

cordia Massal., Anisomeridium (Mull. Arg.) Choisy, Pleurotrema Mull.

 

Arg.,_£yrenocollema Reinke and Strigula Fr.) and Trypetheliaceae Eschw.

 

(Polymeridium (Mfill. Arg.) R. C. Harris and Pseudopyrenula Mull. Arg.).

 

Richard Clinton Harris

Keys, short descriptions, discussions, illustrations and lists of
exsiccati examined and specimens seen are provided for the 85 species
included in the genera above. Distribution maps are provided for the
more common species. A subsequent section contains keys and brief dis-
cussions for the North American species of other genera with colorless
transversely septate spores not directly involved in the revision of
Arthopyrenia: Pyrenulaceae Rabenh. (Lithothelium Mull. Arg. and
Plagiocarpa R. C. Harris), Trichotheliaceae (Mull. Arg.) Bitt. & Schill.
ig_Schill. (Porina Mull. Arg. and Trichothelium Mull. Arg.), Trypethel-
iaceae Eschw. (Astrothelium Eschw., Trypethelium Spreng. and an unde-
scribed genus) and Thelopsis Nyl. whose familial position is uncetain.
These genera contain 47 North American species, although many of them
are left unnamed. Excluded taxa, the disposition of relevant names
presently included in the North American lichen flora, and exsiccati
examined are listed.

No new genera were described but one section is raised to the
rank of genus, Polymeridium (Mull. Arg.) R. C. Harris. Fifteen new

species are described: Anisomeridium finkii, A. macrosporum, A. tuckeri,

 

Arthopyrenia annulata, A, degelii, A, herrei, A, lyrata, A, minor,
A, oblongens, A, taxodii, Mycoglaena wetmorei, Polymeridium exasperatum,
Pyrenocollema imshaugii, Strigula connivens and §, hypothallina.

TWo new names are provided to avoid creating homonyms: Arthopyrenia

 

confluens for Tomasellia leucostoma Mull. Arg. and.Strigula americana

 

for Arthopyrenia tenuis R. C. Harris. Forty-six new combinations are
proposed: Acrocordia meqalospora (Fink) R. C. Harris, Anisomeridium
adnexum (Mull. Arg.) R. C. Harris, A, albisedum (Nyl.) R. C. Harris,

A, ambiggum (Zahlbr.) R. C. Harris,

Richard Clinton Harris

Anisomeridium biforme (Borr.) R. C. Harris, A. carinthiacum (J. Stein.)

 

 

R. C. Harris, A. distans (Willey) R. C. Harris, A. feeanum (Mull. Arg.)

R. C. Harris, A. leucochlorum (Mull. Arg.) R. C. Harris, A; sanfordense

 

 

(Zahlbr.) R. C. Harris, A. subprostans (Nyl.) R. C. Harris, A. tamarindi

 

(Fee) R. C. Harris, A. willeyanum (R. C. Harris) R. C. Harris,

 

Arthopyrenia cedrina (Zahlbr.) R. C. Harris,_A. plumbaria (Stizenb. in

 

Hasse) R. C. Harris, Pleurotrema anacardii (Vain.) R. C. Harris,

 

Polymeridium albidum (M611. Arg.) R. C. Harris,_§. catapastum (Nyl.)

 

 

R. C. Harris,_§. contendens (Nyl.) R. C. Harris,_§. pleiomerellum (Mull.

 

 

Arg.) R. C. Harris, 2. guinqueseptatum (Nyl.) R. C. Harris,_§. subciner-

 

eum (Nyl.) R. C. Harris, Porina heterospora (Fink in Hedr.) R. C. Harris,

 

.E’ microspora (Fink in Hedr.) R. C. Harris, Pyrenocollema caesia (Nyl.)

 

 

R. C. Harris, E. epigloea (Nyl.) R. C. Harris, 2. halodytes (Nyl.) R. C.

Harris, 3. prospersella (Nyl.) R. C. Harris, 3. saxicola (Massal.) R. C.

 

Harris, 3. tichothecioides (Arn.) R. C. Harris, Strigula affinis

 

 

 

(Massal.) R. C. Harris, é, phaea (Ach.) R. C. Harris, §, stigmatella

(Ach.) R. C. Harris, g, submuriformis (R. C. Harris) R. C. Harris,

 

g, sychnogonioides (Nitschke in_Rabenh.) R. C. Harris, g. taylori (Nyl.)

 

R. C. Harris, §, viridiseda (Nyl.) R. C. Harris, g. wilsonii (Ridd.)

 

R. C. Harris, Tomasellia americana (Minks ex_Willey) R. C. Harris,

 

.2. californica (Zahlbr.) R. C. Harris, I, eschweileri (Mfill. Arg.) R. C.

 

 

Harris, 2, lactea (Ach.) R. C. Harris, 2, macularis (Minks ex Willey)

 

R. C. Harris, 2. sparsella (Nyl.) R. C. Harris and Trypethelium

floridanum (Zahlbr. ex Choisy) R. C. Harris.

 

A TAXONOMIC REVISION OF THE GENUS

ARTHOPYRENIA MASSAL. S. LAT. (ASCOMYCETES) IN NORTH AMERICA

BY

Richard Clinton Harris

A DISSERTATION

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Botany and Plant Pathology

1975

ACKNOWLEDGEMENTS

Above all I would like to thank Dr. Henry Imshaug for his invaluable
advice on nomenclatural and editorial problems. Special thanks go to
Dr. Howard Crum and Dr. Rogers McVaugh for support, both spiritual and
financial, during the past year which has made completion of this thesis
possible. I would also like to thank the curators and institutions from
which material was borrowed, especially Peter James, British Museum, for
the loan of Acharian specimens, 0. Vitikainen, Helsinki, for his help
with Nylander types and Dr. Shirley Tucker, Louisiana State University,
for lending me her very interesting unidentified personal collections.

Finally, I wish to thank W. R. Buck for helping with some of the typing.

ii

Section

II.

III.

TABLE OF CONTENTS

INTRODUCTION OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO

METHODS O00......O...OOOOCCOOOOOOOOOOOOOOOOOOOOOOOOO

TAXONODQIC CI—{ARACTERS 0.0.0.000...OOOOCOOOOOOOOOOOOOO

A. Thallus ........................................
B. Phycobiont .....................................
C. Ascocarp .......................................
D. MmaHWI”.H.u.u.u.n.n.u.u.n.n.u.n.
E. Asci ...........................................
F. Ascospores .....................................
G. Microconidia ...................................
H. Macroconidia ...................................

I. LiChen SUbStanceS 0.0......OOOOOOOOOOOOOOOOOOOOO

A SUGGESTED REORGANIZATION OF THE FAMILIES

OF PYRENOLICHENS

SYSTEMATIC SECTION

A. A key to the genera of hyalodidymous and hyalo-
phragmous pyrenolichens of North America .....

B. Pleosporaceae Wint.

l. Arthopyrenia Massal.

 

iii

Page

12

14

17

2O

21

21

23

29

29

34

35

VI.

2.

3.

4.

Arthopyrenia bifera group ..................

 

mcoglaena v. Hahn. OOOOOOOOOOOOOOOOOOOOOOO

 

Tmasellia Massal. OOOOOOOOOOOOOOOOOOOOOOOO

 

Strigulaceae Fr.

1.

2.

5.

Acrocordia massal. OOOOOOOOOOOOOOOOOOOOOOOO

Anisomeridium

 

(Mull. Arg.) Choisy ..........

Pleurotrerna Mull. Arg. COOOOOOOCCOOCCCOCOOO

 

PyrenOCOllerna Reime OOOOOOOOOOOOOOOOOOOOOOO

 

Strigula Fr.

Trypetheliaceae Eschw. ........................

l.

2.

Polymeridium (Mull. Arg.) R. C. Harris .....

 

Pseudopyrenula Mull. Arg. .................

SOME ADDITIONAL HYALINE SPORED PYRENOLICHENS .......

A.

Pyrenulaceae Rabenh. ..........................

l.

2.

Lithothelium Mall. Arg. OOOOOOOOOOOOOOOOOOO

 

Plagiocarpa R.

 

C. Harris OOOOOOOOOOOOOOOOOO

Trichotheliaceae (Mull. Arg.) Bitt. & Schill.

l.

2.

Porina Mull. Arg. COO...OOOOOOOOOOOOOOOOOOO

TriChOthelim Mfillo Mg. OOOOOOOOOOOOOOOOOO

 

Trypetheliaceae Eschw. ........................

l.

2.

3.

AstrothEJ—ium ESChWo OOOOOOOOOOOOOOOOOOOOOOO

 

Trypethelium Spreng. ......................

 

Undescribed genus with parathelioid
ascocarps 00.000.00.00...COCO-0.0.00.0...O

Family urlcertain OOOOOOOOOOOOOOOOOOOO00.0.00...O

l.

Thelopsis Nyl.

iv

69

74

78

9O

92

97

118

121

131

148

150

160

163

163

164

165

166

166

179

180

180

183

189

189

189

VII.

VIII.

XI.

XII.

EXCLUDED TAXA COCOOOOOCCCOOOOOOOOOCOO.COOOOOIOOOOOOO

A. BXClUdEd generic names O..COOOOOOOOOOCOOCOOOOOOI

B. Excluded specific names ........................

DISPOSITION OF THE PYRENOLICHEN NAMES INCLUDED IN
HALE AND CULBERSON (1970) ACCORDING TO THE
TREAWT OF TI-{Is WORK .C...‘.......C.....00......

ALPHABETICAL LIST OF EXSICCATI EXAMINED ............

LITERATIJRE CITED 0......00.0.0.0...OOOOOCOOOOOOOOOOO

mDEXTOTAXA OOOOOCOCOOCOOOOOOCOOOO00.000.00.000...

FIGIJRES O00......0......0....OOOCOOOOOOOOOOOOOOOOOOO

192

192

194

195

199

203

208

214

LIST OF FIGURES

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Figures Page
1-6. Arthopyrenia antecellens (Nyl.) Arn. .............. 215
7-11. Arthopyrenia atomarioides Mull. Arg. .............. 215
12-15. Arthopyrenia atractospora Zahlbr. ..... ...... ...... 215
16-19. Arthopyrenia cedrina (Zahlbr.) R. C. Harris ........ 215
20-23. Arthopyrenia cerasi (Schrad.) Massal. ............. 217
24-30. Arthopyrenia cinchonae (Ach.) Mfill. Arg. ....... ... 217
31-35. Arthopyrenia cinereopruinosa (Schaer.) Massal. .... 217
36-39. Arthopyrenia confluens R. C. Harris ................ 219
40-43. Arthopyrenia degelii R. C. Harris ... ..... .......... 219
44-48. Arthopyrenia fraxini Massal. ...................... 219
49-52. Arthopyrenia herrei R. C. Harris ................... 219
53-59. Arthopyrenia lapponina Anzi ........................ 221
60-64. Arthopyrenia megalospora Lannr. ................... 221
65-68. Arthopyrenia minor R. C. Harris .................... 221
69-72. Arthopyrenia oblongens Zahlbr. 2x R. C. Harris ..... 223

 

73-79. Arthopyrenia padi Rabenh. ......................... 223

 

80-87. Arthopyrenia_p1anorbis (Ach.) Mull. Arg. .......... 223

 

88-92. Arthgpyreniagplumbaria (Stizenb.) R. C. Harris ..... 225

 

93-96. Arthopyrenia rhyponta (Ach.) Massal. .............. 225

 

97-100. Arthopyrenia salicis Massal. ...................... 225

 

 

101-104. Arthopyrenia taxodii R. C. Harris ................. 225
105-108. Arthopyrenia annulata R. C. Harris ................. 227

 

vi

109-113.
114-118.
119-125.
126-130.
131-134.
135-137.
138-140.
141-144.
145-149.
150-155.
156-159.
160-166.
167-169.
170-173.
174-178.
179-182.
183-186.
187-190.
191-194.
195-199.
200-207.
208-212.
213-218.
219-224.
225-229.
230-235.

236—241.

Arthopyrenia bifera Zahlbr.

 

Arthopyrenia Hrata R. C. Harris ......OOOOCOOOCOOOO

 

Mycoglaena

Mygoglaena

meridionalis (Zahlbr.) Szat.

myricae (Nyl.) R. C. Harris ..

 

Mycoglaena

quercicola R. C. Harris ......

 

Mycoglaena

wetmorei R. C. Harris ........

 

Tomasellia

americana (Willey) R. C. Harris .........

 

Tomasellia

arthonioides (Massal.) Massal.

 

Tomasellia

californica (Zahlbr.) R. C. Harris ......

 

Tomasellia

eSchweileri (Mull. Arg.) R. C.

 

Tomasellia

gelatinosa (Chev.) Zahlbr. ..

 

Tomasellia

lactea (Ach.) R. C. Harris ...

 

Tomasellia

Harris ...

macularis (Willey) R. C. Harris .........

 

Tomasellia

sparsella (Nyl.) R. C. Harris

 

Acrocordia

cavata (Ach.) R. C. Harris ...

 

Acrocordia

Acrocordia

conoidea (Fr.) Korb.

megalospora (Fink) R. C. Harris .........

 

Anisomeridium

Anisomeridium

Anisomeridium

 

Anisomeridium

 

Anisomeridium

carinthiacum (J. Stein.) R.

 

Anisomeridium

 

Anisomeridium

 

Anisomeridium

finkii R. C.

 

Anisomeridium

 

Anisomeridium

macrosporum R. C. Harris ..

 

vii

adnexum (Mull. Arg.) R. C. Harris ....
albisedum (Nyl.) R. C. Harris ........
ambiguum (Zahlbr.) R. C. Harris ......

biforme (Borr.) R. C. Harris .........

C. Harris

distans (Willey) R. C. Harris . . . . C . . .
feeanum (Mull. Arg.) R. C. Harris ....
Harris ..................

leucochlorum (Mfill. Arg.) R. C. Harris

227

229

229

229

231

231

231

231

231

233

233

233

235

235

235

237

237

239

239

239

241

241

241

243

243

243

245

242-245.

246-250.

251-256.

257-261.

262-265.

266-269.

270-272.

273-2740

275-278.

279-283.

284-288.

289-292.

293-295.

296-300.

301-308.

309-314.

315.

316-320.

321-322.

323-325.

328-336.

337-3410

346-348.

349-353.

354-359.

Anisomeridium sanfordense (Zahlbr.) R. C. Harris ...

Anisomeridium subprostans (Nyl.) R. C. Harris ......

Anisomeridium tamarindi (Fee) R. C. Harris .........

 

Anisomeridium tuckeri R. C.

Harris 00.000.000.000...

 

Anisomeridium willeyanum (R. C. Harris) R. C. Harris

 

Pleurotrema anacardii (Vain.) R. C. Harris .........

 

Pleurotrema inspersum Mull. Arg. ...................

 

Pyrenocollema caesia (Nyl.) R. C. Harris ...........

 

Pyrenocollema epigloea (Nyl.) R. C. Harris .........

Pyrenocollema halodytes (Nyl.) R. C. Harris ........

 

Pyrenocollema imshaugii R. C. Harris ...............

 

Pyrenocollema prospersella (Nyl.) R. C. Harris .....

 

Pyrenocollema saxicola (Massal.) R. C. Harris ......

 

Pyrenocollema tichothecioides (Arn.) R. C. Harris ..

 

Strigula

affinis (Massal.) R. C. Harris ............

 

Strigula

americana R. C. Harris 00000000000000.0000.

 

Strigula

complanata (Fee) Mont.

 

Strigula

connivens R. C. Harris 00.000.000.000000000

 

Strigula

Strigula

elegans (Fee) Mfill. Arg.

hypothallina R. C. Harris .................

 

Strigula

nitidula Mont.

 

Strigula

phaea (ACho) R. C. Harris 0.00.00.00.00...-

 

Strigula

Strigula

stigmatella (Ach.) R. C. Harris ...........

submuriformis (R. C. Harris) R. C. Harris

 

Strigula
Strigula

Strigula

sychnogonioides (Nitschke) R. C. Harris ...
taylori (Nyl.) R. C. Harris ...............

viridiseda (Nyl.) R. C. Harris ............

viii

245

247

247

247

247

249

249

249

249

251

251

251

251

253

253

253

255

255

255

255

255

255

257

257

257

257

259

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

ix

360-366. Strigula wilsonii (Ridd.) R. C. Harris ............. 259
367-371. Polymeridium albidum (Mull. Arg.) R. C. Harris ..... 259
372-378. Polymeridium catapastum (Nyl.) R. C. Harris .. ...... 261
379-382. Polymeridium contendens (Nyl.) R. C. Harris ........ 261
383-387. Polymeridium exasperatum R. C. Harris .............. 261
388-391. Polymeridium pleiomerellum (Mfill. Arg.) R. C. Harris 263
392-395. Polymeridium guingueseptatum (Nyl.) R. c. Harris ... 263
396-399. Polymeridium subcinereum (Nyl.) R. C. Harris ....... 263
400-405. Pseudopyrenula subgregaria M611. Arg. .............. 263
406. Arthopyrenia cinchonae (Ach.) Mfill. Arg. .......... 264
407. Arthopyrenia fraxini Massal. ...................... 265
408. Arthopyrenia padi Rabenh. ......................... 266
409. Arthopyrenia planorbis (Ach.) Mfill. Arg. .......... 267
410. Arthopyrenia plumbaria (Stizenb.) R. C. Harris ..... 268
411. Arthopyrenia bifera Zahlbr. ....................... 269
412. Arthopyrenia lyrata R. C. Harris ................... 270
413. Tomasellia eschweileri (Mull. Arg.) R. C. Harris ... 271
414. Tomasellia lactea (Ach.) R. C. Harris .............. 272
415. Tomasellia sparsella (Nyl.) R. C. Harris ........... 273
416. Acrocordia cavata (Ach.) R. C. Harris .............. 274
417. Anisomeridium biforme (Borr.) R. C. Harris ......... 275
418. Anisomeridium tamarindi (Fee) R. C. Harris ......... 276
419. Anisomeridium tuckeri R. C. Harris ................. 277
420. Anisomeridium willeyanum (R. C. Harris) R. C. Harris 278
421. Pyrenocollema halodytes (Nyl.) R. C. Harris ........ 279
422. Strigula americana R. C. Harris .................... 280

423.

424.

425.

426.

427.

4280

429.

430.

Strigula complanata (Fee) Mont. ...................

 

Strigula elegans (Fee) M611. Arg. .................

 

Strigula phaea (Ach.) R. C. Harris .................

 

Strigula viridiseda (Nyl.) R. C. Harris ............

 

Strigula wilsonii (Ridd.) R. C. Harris .............

 

Polymeridium catapastum (Nyl.) R. C. Harris ........

 

Polymeridium quinqueseptatum (Nyl.) R. C. Harris ...

 

Pseudopyrenula subgregaria Mfill. Arg.

 

281

282

283

284

285

286

287

288

r-.
s..
‘50 1
.
'4
....
A.
u

 

 

I . INTRODUCTION

This study was begun with the intention of monographing the genus

Arthopyrenia in North America. After several years of study it became

 

clear that Arthopyrenia was not a single genus. This view was strongly
supported by the developmental studies of Janex-Favre (1971) which
indicated that some of the species included in the genus belong to very
different groups of Ascomycetes. As a result it became necessary to
define the genera involved before any monographic studies could be
attempted. Thus the direction of my study was modified to provide a
foundation on which future monographic studies may be constructed.
Since definitions of generic limits require a knowledge of related
genera, the study was broadened to include the other genera of pyreno-
lichens with hyaline spores. The preliminary results of these supple-
mentary investigations are presented in the form of keys to provide a
context for the dismemberment of Arthopyrenia. They also provide a
rather comprehensive manual by which the more common pyrenolichens with
colorless, transversely septate spores may be identified. Ultimately I
ended up making some preliminary studies in almost all of the genera of
pyrenolichens (mostly excluding the Verrucariaceae). This has meant
almost ten years of work and examination of approximately 7000 specimens.
The tentative results of this overview are embodied in a suggested re-
organization of the families of pyrenolichens.

This paper is in no way intended to be final or complete. It is
intended to place my ideas on the systematics of the pyrenolichens
before the scientific community to serve as a basis for discussion.

Nor is it complete in its coverage of the species of Arthopyrenia 5. lat.

in North America. Specimens from several large herbaria remain to be

1.
fi'

:fi
r..'.

 

 

studied. Also a number of presumed new species have not been included
since they are represented by only one or two collections. I feel that
this is justified since I hope to go on from this starting point to
monograph each of the segregate genera separately. Also the probability
of these rare species being encountered is very low. Undoubtedly there
will be some changes in specific names in the future as I have not ex-
amined the types of a large number of tropical American species (Brazil
especially has many species in common with the southern United States).
Thus at the familial and generic levels this study is intended to serve
as a foundation for future work, but at the specific level it is best

described as a progress report.

Arthopyrenia, as a genus, has in the past been defined mainly by

 

three characters, perithecial ascocarp, colorless transversely septate
spores and branched paraphyses which are usually persistent. It has
grown by a process of accretion to an unwieldy size of c. 300 species.
This alone might lead one to suspect the naturalness of the genus.

That the unnaturalness of this assemblage was in part realized by
earlier workers seems to be indicated by the often extensive generic
synonymies. Thus I am not so much in the position of creating new
genera but in one of redefining already published taxa. Keissler (1936-
38) made a beginning in separating out the species associated with blue-
green algae. Riedl (1962) has tried to reduce the size of the genus by
dividing it on the basis of spore septation. Recent European workers

have re-established Acrocordia (VEZda. 1968; Poelt, 1969). VEzda (1968)

 

has very clearly pointed out the need for segregate genera from

Arthopyrenia.

In establishing these segregate genera I have taken as a first

principlerthat they should be as homogeneous as possible. I have

retained in Arthopyrenia any species whose position is in doubt. Thus

 

Arthgpyrenia itself is still somewhat heterogeneous. I suspect further

 

work may lead to the segregation of at least one additional genus

(Arthopyrenia bifera group). The splitting up of Arthopyrenia has re-

 

sulted in a very large number of name changes, however I feel that
these are fully justified by the distinctness and homogeneity of the
segregates.

The terminology used here is essentially that used in Harris
(1973). Any modifications or additional terms are included in the dis-
cussions of the appropriate structures in the section on taxonomic
characters. Since the pyrenolichens include ascolocular, semi-asco-
hymenial and ascohymenial members, I have chosen to use a single rela-
tively neutral set of terms throughout rather than changing with each
group even though some precision is lost. In discussing ascus types and
ascocarp types I follow the terminology of Chadefaud (1973) and Janex-
Favre (1971), mostly translated into English but occasionally left in
the original French where this seems appropriate. Spore shapes are
described in terms of plane figures based oneuioptical section.

Although this study was primarily centered on North America north
of Mexico, a considerable amount of European, West Indian and South
American material was examined. For common European species or where
it has otherwise seemed useful some species not occurring in North
America have been included. Species with only a few collections from
North America have not been mapped nor have those whose distribution is
essentially unchanged from that shown in Harris (1973). In the case of

very common species only selected specimens have been cited.

The abbreviations for journal titles have been taken from
Lawrence E: 21. (1968), those of exsiccati from Lynge (1920-22, 1939) or
follow the pattern of these works. Acronyms for herbaria in specimen
citations are the standard ones of Lanjouw & Stafleu (1964). Herbaria
of special significance kept separately from the general collections are
indicated by the acronym of the institution plus the abbreviated name of
the original owner of the collection. In the case of the Tuckerman and
Nylander herbaria a sheet or specimen number is also included to facili-
tate location of the specimens. Specimens have been seen from the
following herbaria: CAN, FH, G, H, LD, LSU, M, MICH, MSC, OSC, PC, 3,

TUR, UMBS, UPS, US, US 339 M0, w & WIS.

II. METHODS

The methods of study used were mainly those outlined in Harris
(1973). The only major change was that the critical characters have
been recorded in drawings for the majortity of specimens examined using
a Wild drawing tube. This procedure, although very tedious and time
consuming, has a number of very definite benefits. It provides a per-
manent record of a collection's characteristics and greatly reduces the
necessity for re-examination with consequent destruction of additional
material. This is most important in preserving material for future study
in these groups where a type collection may consist of only a few tiny
perithecia. Secondly it forces one to examine each specimen very
closely. These advantages are obvious. More important, however, is
that the large amount of time necessarily spent at the microscope pro-
vides the time to consider each specimen and its relations to other
specimens very carefully. As a result of this procedure my taxonomic
concepts evolved through the relating of specimens to a few common
species for which I had abundant material. These species thus served
as nuclei around which the segregate genera have formed. The collec-
tions were not actually completely sorted into species until late in
the study. Finally, these camera lucida drawings have been directly
utilized for the illustrations, either by direct tracings in the case
of the asci and ascocarp cross sections or by tracings with a panto-
graph in order to enlarge them in the case of the ascospores and
conidia.

A minor change in method is the increased use of two stains.

Phloxine (1-2 percent in water) mixed with 15-20 percent aqueous

6
potassium hydroxide (KOH) was used to stain thin sections of the thallus,
especially in old herbarium specimens, to check the presence or absence
of algae. The contents of the algal cells usually stain more heavily
than the surrounding bark cells. Additional contrast between bark and
algal cells may be obtained by washing with KOH. Chlorazol Black
( 1-2 percent in water) mixed with KOH on a microscope slide immediate-
ly before use has proved to be the most satisfactory stain for the

chitinoid ring in the ascus tip of Porina and Trichothelium.

 

III. TAXONOMIC CHARACTERS

AM

In the majority of the Pleosporaceae and Strigulaceae the thallus
is mostly just a lighter blotch on the surface of the substrate and
provides little or no information of taxonomic value. In some species
of Strigula, especially the foliicolous ones, the growth habit, color,
presence of punctation and presence of hypothallus are useful in dis-
tinguishing species. In the Pyrenulaceae, Trichotheliaceae and Trype-
theliaceae the thallus may provide as much taxonomic information as the
characters of the ascocarp, asci and spores. There is considerable
variation in such characters as texture, color, pruinosity, punctation
and crystalline inclusions. In Porina there are even some species
which produce isidia. Interestingly enough pyrenolichens seem not to

produce soredia. The only instance I am aware of is Normandina

 

pulchella (Borr.) Nyl.

B. Phycobiont

 

Among the pyrenolichens there seems to me to be a high correlation
between the nature of the phycobiont and mycological characters at the
familial level or occasionally even at the generic or specific level.
Those genera which I consider to belong in the Pleosporaceae have only

a very few species which seem to be associated with Trentepohlia, the

 

majority are non-lichenized. Ahmadjian (1958, 1967) has reported six

genera of the Chlorophyceae (but not Trentepohlia) and one of the

 

Xanthophyceae from various members of the Verrucariaceae. The Stri-

gulaceae, Trypetheliaceae and Trichotheliaceae are associated with

 

 

8

Trentepohlia with a few exceptions. Foliicolous species of Strigula

 

are associated with Cephaleuros, while those of Porina are associated

 

with Phycopeltis. A few species in the Strigulaceae and Trypetheliaceae

 

seem to be non-lichenized. However, this is not certain since they are
tropical species known from very few collections, often in poor condi-

tion. It is quite possible that Trentepohlia could be found in fresh

 

material. A few species in the Strigulaceae are associated with various
blue-green algae. They have been placed in a single genus which is
distinct on mycological grounds as well. They seem to represent an
evolutionary line line adapted to aquatic and semi-aquatic habitats.
It could be possible that this association with blue-green algae is
part of the adaptation. There are, however, a few semi-aquatic species

of Anisomeridium associated with Trentepohlia.

 

 

C. Ascocarp

The ascocarp and associated modifications of the thallus are of
relatively little use at the generic and specific levels in the Pleo-
sporaceae and Strigulaceae. In the past much has been made of whether
the carbonized wall layer is continuous beneath the ascocarp or not.

A given species may have a definite tendency in this regard but it is
subject to considerable modification due to the nature of the substrate,
age, and amount of erosion of the upper layers of thallus and bark.
Ascocarp size and shape are of some use but are often quite variable
within a species. The amount the ascocarp is immersed in the substrate
or covered by the thallus is occasionally useful but is also subject to
environmental modification. Although the above characters are compara-
tively easily observed, I have tended to avoid using them in keys due

to their variability.

9

Obvious exceptions to the above are when the position of the
ostiole is eccentric or lateral (parathelioid ascocarp), when two or
more ascocarps are joined by the fusion of their ostiolar necks (astro-
thelioid ascocarp) or when fusion is even more complete so that a com-
pound ascocarp is formed with the ostioles separate (mycoporoid asco-
carp) or joined (astrothelioid ascocarp). The parathelioid and astro-
thelioid types are lacking in the Pleosporaceae and Trichotheliaceae,
rare in the Strigulaceae, but fairly common in the Pyrenulaceae and

TryPetheliaceae. These two types of ascocarps have been used in the
Past to define families (Zahlbruckner, 1926) but it is clear that such
falnilies are unnatural and the genera involved are better placed with
their erect ostioled relatives.

The color of the outer. layers of the ascocarp wall is more or

1938 uniformly brown to brownish black in the majority of the pyreno-
1ichens but species of Mycoilaena, Porina and Thelopsis provide notable
eXCeptions. The color in these may be blue green, yellowish to reddish,
pallid or even colorless. The color of the ascocarp is taxonomically
important in these genera.

Although only a few species have been studied it seems as if the
manner in which the ascocarp develops may be distinctive at the fami-
lial level (Janex-Favre, 1971). Arthopyrenia 5. str. has typical asco-
lOcular development as one would expect in a member of the Pleosporaceae.
Me"Ibers of the Verrucariaceae (except Dermatocarpon) have a develop-
mental type combining features of both ascolocular and ascohymenial

asCocarps. Chadefaud (1973) has referred to them as semi-ascohymenial.

D‘erfiitocarpon is wholly ascohymenial (Janex-Favre, 1971). Acrocordia

 

Mel; and Pyrenocollema halodytes, both in the Strigulaceae, have

 

..--i

o.

C...

 

~.

,‘-

‘.
‘.

u

- t‘

 

 

10

ascohymenial development but lack any ostiolar apparatus. Janex-Favre
refers to these as perithecioid ascocarps. Pyrenula nitida and her
Porina sp. are also ascohymenial but with an ostiolar apparatus with
periphyses and are indicated as possibly more closely related to
primitive discomycetes than to the non-lichenized ascohymenial pyreno-
*mycetes (Janex-Favre, 1971).

I have previously used the terms involucrellum and exciple
(Harris, 1973). The work of Janex-Favre has shown that the situation
is more complex, but I feel that the terms are still useful. The invo-
lucrellum is the primary envelope which involves the thallus and bark in
its outer layers as in Pgenula nitida and her Porina sp. Or perhaps,
as in Acrocordia conoidea and Pyrenocollema halodytes, the thalline
e1'1Vv5310pe could be considered to be equivalent to the involucrellum.

In fact these two envelopes may be different expressions of the same
Stmcture. The involucrellum could also be regarded, in my opinion, as
a Very much reduced pseudostroma since it contains bark and algal cells.
The exciple would then usually be either the secondary envelope and/or
the carpocentral envelope.

The involucrellum, carbonized or not, often contains colorless
Crystals (Pyrenulaceae, Trichotheliaceae and Trypetheliaceae). In the
Pyrenulaceae crystals are most often produced in and around the ostiolar
apparatus or in the ascocarp wall. Production of crystals, although
0Cicasionally variable, seems on the whole to be a reliable specific

Character.

The well developed ostiolar apparatus in the Pyrenulaceae seems to

me to be of biological significance. In many species, especially those

With parathelioid and astrothelioid ascocarps, the ostiolar apparatus

11

is large and heavily gelatinized and in fact may control spore libera-
'ti43r1 from the ascocarp. When wet the diameter of the canal through it
seaemns less than the diameter of the spores. This combined with the
fact that the neck of some ascocarps is several times longer than the
aasn:j_ leads me to speculate that the spores are released within the
ascocarp. Granted that the bitunicate asci of this group are extensible,
but not only would they have to extend several times their own initial
ILEflfigrth [ and none of the illustrations by Swinscow (1966) or Morgan-
Jones (1972) show any great amount of extension], they would have to
force their way through the narrow canal of the ostiolar apparatus.
The hymenium contains much gelatinous material and the secondary enve-
lope is free from the primary envelope except at the tip and contracts
and expands with changes in moisture. Perhaps the pressure developed
in these changes expels the spores. However I suspect that the final
iWQDEFtus which liberates the spore is given by the ostiolar apparatus
Wich, pops it out in the same way a bean squeezed between two fingers
ma)’ toe shot some distance. Thus it seems possible that in the Pyrenu-
lac:eae the ascocarp has taken over the function of the ascus in spore
lihkeration. Indeed it seems to be one of the characters which help
dififerentiate the Pyrenulaceae from the Trypetheliaceae, since in the
fcnflner the secondary envelope commonly separates from the primary enve-
14396, I have never observed this phenomenon in the latter. Nor have I
Observed any well developed ostiolar apparatus in the Trypetheliaceae.
As indicated in the section on lichen substances, various pig-

m‘Efnts are not uncommonly deposited within the outer layers of the

iHVOlucrellum or pseudostroma in the Pyrenulaceae, Trichotheliaceae and

Trypetheliaceae.

12

In the Trypetheliaceae and to a lesser extent in the Pyrenulaceae
tjiez.ascocarps may be aggregated into small or large groups within
{asuiexdostromata. These pseudostromata may be little different from the
Stirurounding thallus, e.g., Trypethelium virens, or may form warts much
cijrffierent from the thallus, e.g., Trypethelium eluteriae or Laurera spp.
4A£3 61 result species in the Trypetheliaceae can often be identified with
tile: dissecting microscope alone, whereas in other pyrenolichen families

SENDInes and even conidia are required for a positive identification.

D. Hymenium‘

The hymenium consists of interascal threads of various sorts and
thfi aasci, often surrounded by abundant gelatinous material. I have de-
Ciiieaci to follow Poelt (1974) in using paraphyses as a neutral term for
th€3 :interascal threads. More precisely however those in ascolocular
asCKDczarps, e.g., Arthopyrenia 5. str., are pseudoparaphyses. Those

ass'>O<:iated with ascohymenial ascocarps, e.g., Acrocordia, are true

 

parnaldhyses (Janex-Favre, 1971). Members of the Verrucariaceae have

botfli pseudoparaphyses and true paraphyses or only pseudoparaphyses.
The amount of branching and anastomosing of the paraphyses is

geruarally a useful character, although not one which lends itself to

e35?! description or illustration. In some species of Arthopyrenia and

 

915L of those of Tomasellia the paraphyses are quite broad, much branched
and consist of short cells so that the hymenium appears cellular. In
the Verrucariaceae the paraphyses are lacking or very soon disappear.
In the remaining pyrenolichens the paraphyses are slender and thread-

]ike with relatively long cells. In Strigula there is very little

trenching and this is helpful in recognizing members of this genus.

...

 

 

..
..-

 

 

13

13162 paraphyses in members of the Trichotheliaceae are essentially un-
branched although an occasional dichotomy may be found. In the Trype-
theliaceae on the other hand it is quite the opposite since the regular
net-like branching and anastomosing seems to characterize the family.

The gelatin surrounding the paraphyses and asci often reacts with
iodine, giving blue green, sordid or orangish colors. As far as I know
aJJI <of the members of the Verrucariaceae exhibit this reaction. A size-
able percentage of the Pyrenulaceae react as do a small number of the
Pleosporaceae. This seems to be a useful character in recognizing
SpeCies in these families. The Strigulaceae, Trichotheliaceae and
T1TYpetheliaceae have not yet been found to have any hymenial reactions
With iodine.

In addition to the gelatinous matrix the hymenium may contain
Other substances dispersed through it. The presence of these substances
appears to be a good specific character in most cases. In the Pyrenu-
laCREEae they are usually in the form of small oil-like droplets. In the
TrYpetheliaceae, at least in herbarium specimens, they in the form of
col<Drless amorphous granules which liquify on addition of KOH. This
difference between the types of materials inspersing the hymenium
seefils to be quite reliable in distinguishing the 'two families. The
other families of pyrenolichens may have a few oil-like droplets in the
k‘Ymenium, they are never abundant enough or variable enough to provide

anYtaxonomic information.

In Psuedopyrenula the outer layer of the hymenium contains a

yellowish pigment which turns red in KOH. The amount present is often
Variable and thus difficult to detect in some cases. An attempt to

correlate the presence or absence of this pigment with spore size led

 

 

 

 

 

14

to chaos. More study is required to assess the taxonomic worth of this

pigment. Temporarily I do not attach much importance to the pigment at

the species level, although its presence is diagnostic for the genus.

E. Asci

With few exceptions the fundamental ascus type of the pyrenolichens
has been found to be the bitunicate type (Richardson & Morgan-Jones,

1964; Swinscow, 1965b, 1966, 1967; Morgan-Jones & Swinscow, 1965; Ve'zda,

1968; Janex-Favre, 1971; Morgan-Jones, 1972). In those most closely

Stuclied they have proved to be bitunicate-nassascé (Janex-Favre, 1971).
ThuS the asci have an exoascus and endoascus which are not permanently

fUSEd together and the ocular chamber contains a nassé [see Chadefaud

”—9 73) for a summary of ascus types]. In the process of spore liberation

the more fragile exoascus ruptures and the endoascus increases in

length and releases the spores. A number of illustrations in the papers

referred to above show the exoascus breaking some distance from the tip

of the ascus leaving a thimble-shaped cap of exoascus on the extended

erlcloascus. This troubles me somewhat, since if the spores are to be

released through a pore or break in the tip of the endoascus, this cap

would almost surely interfere. In the perithecial type of ascocarp I

can visualize no ready mechanism for the endoascus to rid itself of

this cap. The only obvious method would be for it to be carried away

by the discharge of the spores, but that would surely reduce the distance

which the spores could be discharged. I have never observed any such

thimble-shaped caps left in the hymenium. Another possibility is that

the endoascus simply disintegrates releasing the spores within the asco-

carp and then the spores are released from the ascocarp by some means

15

other than active discharge from the ascus. In the Pyrenulaceae there

are some indications that this what happens (see under discussion of

ascocarp). In view of the difficulties which an exoascus cap might

present to spore release it seems to me more probable that the brittle

exoascus has been broken in an unnatural manner in the process of pre-

paring the material for study. This, of course, would not change the

interpretation that these asci are bitunicate.

In the Trichotheliaceae the ascus seems to be fundamentally dis-

tinct from those previously discussed. Janex-Favre (1971) has inter-

preted the ascus in a species of Porina as being of the archeascé type,

laCking a nassé, but bitunicate with the endoascus reduced mainly to

the apical dome. Janex-Favre also reports a raised ring-like thickening

at the tip of the exoascus. I have examined many species of Porina and

two of Trichothelium and have seen no indications that the ascus is

functionally bitunicate. There is, however, in most species a very

distinct ring in the tip of the ascus. It is often visible without

Staiming due to its refractive properties differing from those of the

Surrounding asucs wall. It can readily be stained with Congo Red or

ChlOrazol Black and thus is of the chitinoid type.

Janex-Favre (1971,
f.

51,C, D) illustrates it as being raised and thicker than the rest of

the outer layer of the exoascus. As far as I am able to determine it is

flllsh with the rest of the exoascus and included within it (or if the

exOascus were interpreted as being very thin, perhaps in the outermost ,

lfiller of the endoascus). Since it does seem to be part of the exoascus

it is not an apical ring in the sense of Chadefaud and his students, as

the latter structure is a part of the endoascus. It is very similar to

the upper chitinoid ring I have seen in the ascus apex of Nectria

 

y
.

16

eqoijsphaeria (Tode gx_Fr.) Fr. The lower ring is clearly part of the

 

enndkoascus but the upper seems to occupy a position like that in Porina.
I}: ciiffers in being broader. It is possible that the Porina type of

anscnas could be derived from that of Nectria episphaeria by reduction of

 

'tlie: endoascus and loss of the lower ring. This possibility taken to-
ggerttuer with the spore type and the tendency for Porina to have brightly
cxolxored ascocarps, as in the Nectriales, tempts me to wonder if perhaps
tile: Imichotheliaceae might not have its origins in the Nectriales. If
tfxiss were true then the Trichotheliaceae would seem to be only distantly
r€¥larted to the other pyrenolichens. Janex-Favre (1971) described the
aScocarp development of Porina as being very similar to that of Pyrenula
“filiJZTl would argue against a relationship with the Nectriales. However
‘it Eneems to be a possibility which should be investigated.

Some species of Microglaena and Thrombium epigaeum have an amyloid

 

apiiteal ring. Species of Microglaena have been shown to have bitunicate

 

ascgi (Morgan-Jones & Swinscow, 1965). Thus the asci W0U1d seem to be
Of ‘tlie archeascé type with an amyloid ring and bitunicate dehiscence.
A fourth type of ascus is found in Thelopsis and Belonia. The asci
9:53 \rery thin walled, more or less as in Porina, but lacking a chitinoid
firm; in the ascus apex. The hymenial gelatin reacts with iodine which
61533 seems to indicate their removal from the Trichotheliaceae.

Thus the bulk of the pyrenolichens, Pleosporaceae, Pyrenulaceae,
Strigulaceae and Trypetheliaceae have the same basic ascus type. There
599m to be a few smaller groups which may in part be defined by ascus
type. These are the Trichotheliaceae plus two groups of uncertain

POSition.

17

Within the larger groups the asci are useful in defining genera and
species. The Trypetheliaceae are largely characterized by asci with
broad, shallow ocular chambers. The long cylindrical asci with an al-

most subglobose ocular chamber are diagnostic for the genus Acrocordia.

 

Ascus shape is often useful. Typical pleosporoid asci, swollen at the

base, are almost entirely confined to Arthopyrenia s. str. and Toma-

 

sellia. Within Arthopyrenia there is a transition from this type

 

(i- fraxini, A. punctiformis) to nearly cylindrical asci (A. lapponina).

 

The long flexuous ascus base found in most species of Pyrenocollema is

 

""3176? or less diagnostic for the genus. Ascus size may occasionally be
IJSGEfHJl in separating closely related species but as a general rule it
513 ,Ifielated to spore size. Thus ascus size is not included in the

descriptions unless it is genuinely useful in distinguishing species.

F. Ascospores

 

Without any doubt, although it means much tedious microscopic work,
the* Eascospores provide many of the most reliable characters for identi-
fiYirhg and classifying taxa of the pyrenolichens. They have been one of
tIKe rnost important characters used in the formation of genera (Zahl-
bru<aner, 1926), based primarily on color, septation and wall thicken-
iIKQS. This dependence on spore characters to the exclusion of others

luas in some cases led to fairly natural groupings, e.g., Pyrenula, but

1“ Others to largely unnatural assemblages, e.g., Arthopyrenia 5. lat.

 

when assigning a pyrenolichen to a genus one must weigh all available

data. .A case in point is Plagiocarpa hyalospora which has colorless

 

flxmes with very little wall thickening. 0n the basis of this type of

flxme one would not associate the species with Pyrenula. However it

18

ruas the same type of ascocarp, hymenium and conidia which are particular
-tc> Eyrenula and related genera. Further the species is included in the
,saunezgenus with brown spored species on the basis of an unusual asco-

cuaijp type and a unique ascus type.

In addition to the spore characters mentioned above, size and

sriaipe are useful at various levels. It is interesting to note that the

learnger spores are found in lichenized genera, especially Acrocordia

 

arici Anisomeridium. The non-lichenized genera Arthopyrenia and Tomasellia

 

'tearnd to have rather small spores mostly less than 25 u in length. The

ruinfloer of spores in an ascus is usually eight although one or two often

dC> rust develop. A regular decrease in spore number is found in many

genlenra but seems of no significance above the specific level. It is

uSmallyaccompanied by an increase in spore size. Polyspory is very

133513 in the pyrenolichens (Thelopsis), and apparently lacking in the
Inaill families.

However, the separation of the spores into two part
SENDIwes while still in the ascus (Strigula) gives rise to a kind of
polySpory different from the usual sort.

In the smaller spored species the only layers of the spore wall
(Cnladefaud, 1969) which are readily discernible are the perisporal
formation (ectospore + perispore) and the episporal formation (exo-
Spere + epispore). In the Pyrenulaceae and Trypetheliaceae the endo-

SPOral formation (mesospore + endospore) is very well developed and is

in part diagnostic of these families. In fact the endospore is so well

developed that Morgan-Jones (1972, 1973) has interpreted this structure
as forming endoascosporic cells or endospores which could possibly be
set free to act as secondary diaspores. I personally doubt that this

happens. I have examined hundreds of specimens of members of these '

 

...
b.
-
.
s
s
-_,
‘.
'~
~ 1
‘.
'-
I
-

...‘
.‘
,
1
4

,-

 

19

auad never seen these endospores released in the manner illustrated by

Pkoxxgan-Jones. The endospore layer can be partially freed from the

nmesnospore but only with considerable difficulty and not in all species.

III .a.number of species of Pyrenula I have found spores germinated in the

laynruenium in the manner illustrated by Pyatt (1974) by a germ tube at

eaixtfiaer end of the spore. Some species of Anthracothecium and Parmentaria

 

tuaxreemuriform spores sufficently large that the spores can be collected

frrcxu the bark on to which they have been expelled. Such spores are

<oiftnen found to have begun germination by means of numerous germ tubes

Cnreu: the entire surface of the spore. One species of Parmentaria in

aChdjttion to producing germ tubes was found to produce internally a

Jiiyear of conidiophores with conidia similar to those found in normal

Pynzrxidia on the thallus. This suggests that the microconidia, at least

lrl tfliis instance, function as asexual diaspores.

In order to simlify Chadefaud's terminology somewhat, I will refer
tC’ tine perisporal formation by its most obvious layer, the perispore.
Inthe past I and others have referred to the wall thickenings in the

SEXIEes of the Pyrenulaceae and Trypetheliaceae as endospore. It seems

(ilear‘that it is mainly the mesospore which is thickened (Rudolph &

Giesy, 1966; Chadefaud, 1969; Janex-Favre, 1971). It seems wise to

make this distinction since the mesospore layer and the endospore layer

are often different in color and texture.

The perispore is often rather gelatinous and may swell considerably

in water or KOH. In some groups, however, it seems to be very reduced

c>I‘perhaps absent, at least it is not readily discernable. Thus the

degree of development of the perispore seems taxonomically useful.

A number of scattered species and even an entire genus (Acrocordia)

 

have spores with ornamented walls. The ornamentation seems in most

20

cnases to consist of granules trapped between the exospore and perispore

(or? perhaps projections from the exospore. Another type seems to con-

ssijst of small holes in the perispore layer, which when stained, look

riatiher like the granular type of ornamentation. The presence anf type

<31? ornamentation is often of considerable taxonomic value.

G. Microconidia

Conidia, both microconidia and macroconidia, have proved to be

e)<t:remely useful at all levels of classification. Although all are

a~pparently of the same basic type with simple conidiophores, there is

CCNfisxiderable variation in shape and size. The microconidia could also

lb€3 tuermed spermatia. The pycnidia producing them are most commonly

fcnaruj at the margin of the thallus, often in areas without mature asco-

cinIDs, and they can be found in the majority of specimens. Their loca-

ticnl and rather consistent presence suggest that they might be func-

tiCHlal spermatia. In species also producing macroconidia both types

are: found on the same thallus. Occasionally a pycnidium is found pro-

du<Zing both microconidia and macroconidia.

The pycnidia in most species are more or less globose with a
Simple chamber but in the Trypetheliaceae the surface area producing
COnidia may be increased by folding.

In some ways it is unfortunate that the microconidia are so useful.
The botanist with only a few specimens to determine is not likely to make

the often tedious search for pycnidia, but often microconidia are the

most satisfactory character to provide a determination.

21

H. Macroconidia

Ddacroconidia are produced in pycnidia similar to those producing
mirzr13c20nidia but which are usually somewhat larger. The conidia are
prcxitlc:ed singly at the tip of staff shaped conidiophores. They probably
serwree as asexual diaspores.

Pdacroconidia are known with certainty in the colorless spored
pyrwerlcolichens only from two genera, Anisomeridium and Strigula, both in
due £3t;rigulaceae. In Anisomeridium they are non-septate and resemble
1611363 ‘versions of the microconidia. In Strigula the macroconidia are
apparently patterned on the ascospores and have as many cells as the
aSCCHspxores with the exception of Strigula complanata which has more.

In EEFidsomeridium macroconidia have been found in relatively few species

while they are known from all species of Strigula except _£_3_. sychnogoni-

 

oides .
____~_

Tflue macroconidia are taxonomically most useful at the generic

levell, Some of the small spored species of Anisomeridium and Strigula

are sSuperficially very similar but are easily distinguished if macro-

Conldia can be found.

I. Lichen Substances

Lichen substances, although rarely present, are very useful taxon-
omiCally primarily at the specific level. They are not as widespread
and varied in the pyrenolichens as in many other groups of lichens.

The non-lichenized genera belonging to the Pleosporaceae, as expected,
PBVe not been found to produce any lichen substances. In the Verru-

cariaceae I have found zeorin in Normandina. Lichexanthone seems not

 

22
uncommon in the Pyrenulaceae and Trypetheliaceae and occurs in a single
genus of the Strigulaceae (Anisomeridium). Anthraquinones, commonly
parietin, are found in a number of the Pyrenulaceae and Trypetheliaceae

and in one species of the genus Anisomeridium. They are often present

 

in the thallus, over the entire surface as in Pgenula cerina and

 

Trypethelium aeneum, or may occur only over the ascocarps as in Astro-

thelium or in only a small area around the ostiole as in Anisomeridium.

 

Anthraquinone-like pigments may also be present inside the ascocarp

wall as in Pyrenula nitida and Pseudopyrenula spp. The pseudostromata

 

in members of the Trypetheliaceae often contain layers of anthraquinone
Pigments. The Pyrenulaceae and Trypetheliaceae also produce a variety
0f pigments of an unknown nature, usually in relation to the ascocarps.
The oddest case, perhaps, is in a few species of Eyrenula and one of

CEIIT‘EXlO‘thelium where the mouth of the ostiole (but not the surrounding
thallus) is pigmented. As indicated above these various lichen sub-

StanCeS are useful at the specific level and when their distribution is

fully known may have implications at higher levels.

IV. A SUGGESTED REORGANIZATION OF THE FAMILIES OF PYRBNOLICHENS

.Since my ideas on the number and composition of the families of

pyrenolichens do not coincide exactly with any previous treatments

“ZaddjLIDruCkner, 1926; Hale & Culberson, 1970; Poelt, 1974), I have sum-

mec1 11}: their definitions in the key below and their tentative composi-

ticnu. :in a list following the key.

A few of the more interesting prob-

lenus Eire discussed at the end of this section.

Pkaraphyses gelatinized; hymenial gelatin IKI+ pink

(Drxange, orange red or bluish; periphyses usually pre-
sent; microconidia elliptical, oblong or cylindrical;
Imacroconidia not known; almost all species growing on

on rOCk or $011 ......OOOOOOOOOOO......OOOOOOOOOOOOO vermcariaceae

Paraphyses usually persistent (if gelatinizing, then
hanenial gelatin IUCL—); hymenial gelatin IKI- or IKI+;
PKEriphyses rarely present; microconidia various; macro-
Ccnnidia present in some genera; more commonly on sub-

EFtrates other than rock and soil ........................... 2

2. Ascus tip thin with a chitinoid ring in the tip of

the exoascus (ring not evident in Clathroporina and
some large spores species of Porina); ascocarp wall
often brightly colored; paraphyses unbranched;

hymenial gelatin IKI-; microconidia elliptical, ob-

long or rod-shaped; macroconidia not known ........

....O.........OOOOOOOOOOOOOOOOOO0.00.00.00.00. TriChOthelj-aceae

23

24

2. Ascus tip lacking a chitinoid ring in the exoascus ...... 3

lxscus with amyloid apical apparatus ...................

- -................................... Microglaena pr. p. & Thrombium

Iksxnxs lacking amyloid apical apparatus ..................... 4

4i.. Spore wall thickened or if spores muriform and with-

out obviously thickened walls, then with well devel-
oped thallus, ascocarp type and chemical characters
oprmmflaorTqmmmdimlu.u.u.u.n.u.n.n.u..S

Spore wall not thickened; thallus mostly not well

developed 00000000000000.00000000000000 ...... 00000000... 6

Pdixzroconidia filiform; hymenial gelatin commonly IKI+;
Espxares mostly some shade of brown, rarely colorless,
11(151 ascocarps rarely aggregated and included in a

psuedostroma (trypethelioid); macroconidia known in one

gfanus (Eopyrenula) ........... ........ .. ..... ......... Pyrenulaceae
I“hieroconidia rod-shaped; hymenial gelatin IKI-; spores

“Kbstly colorless, rarely brown, occasionally IKI+ violet;

iiscocarps often aggregated and included in a pseudo-

Stroma° macroconidia not known .................... Tr etheliaceae
, yp

6. Ascus not thickened at the tip; hymenial gelatin or

ascus sheath commonly IKI+; periphyses present or

not 000000000000.0.000.0000000000000000000. TrlelOpSiS&Be]-0nia

Ascus thickened at the tip; hymenial gelatin mostly

IKI’; periphyses absent 00000000000000000000000.00000000 7

I
.

 

 

 

 

 

 

 

 

 

 

7.

25

.Microconidia relatively broad, orbicular, elliptical or

<jblong; macroconidia present in two genera; hymenial

glelatine IKI-; spores colorless; almost all species

lgichenized; growing on a wide variety of substrates ...

--...0000000000 00000 0000000000000.0000000000000000000. Strigulaceae

7. LPchroconidia narrower,

short to long rods or rarely

ifjliform; macroconidia not known; hymenial gelatin

nnrostly IKI-, rarely IKI+; spores colorless or brown;

rnrbstly non-lichenized; more or less restricted to bark

and. WOOd 000000000.0000000000..000000000.000000000000 Pleosporaceae

Composition of the families of pyrenolichens

PLEI OS PORACEAE

ArthoDYrenia
BlastOdesmia
Leptorh aphis
MicrOthelia
MYcomiczrothelia?
Mycog laena?
MYC-Oporum
POlelastiopsis
SPOrOSchizon
Tomasellia

PYRENULACBAE

Antihraoothecium
Bottaria
E0Pyrenula
Lithothelium
Nelanotheca
MiCrOglaena 5. str.
Parathelium
Parmentaria.
Plagiocarpa
pleurotheliopsis
Pyrenastrum1
Eyrenula

STRIGULACEAE

Acrocordia
Anisomeridium
Monoblastia
Pleurotrema
Pyrenocollema
Strigula

TRICHOTHELIACEAE

Clathroporina
Porina
Trichothelium

TRYPETHBLIACEAE

Astrothelium
Campylothelium
Cryptothelium
Laurera
Polymeridium
Pseudopyrenula
Trypethelium

 

 

 

 

. ,
u »
-.
'v
-‘ 0‘
<.
‘n ‘
\
‘~
’_\
‘,
.

 

l \
\

I

 

26

VERRUCARIACEZAB GENBRA OF UNCERTAIN POSITION WHICH MAY

As in Poelt (1974) with the CONSTITUTE SEPARATE FAMILIES

addition of l) Belonia, Thelopsis

Microtheliopsis 2) Microglaena pr. p., Thrombium
Normandina

Psoroglaena 3) Aspidothelium?, Phyllobathelium

In addition to the discussion of each family in the taxonomic

section I will mention a few of the more interesting problems here. As

indicated in Harris (1973) Microfiaena is heterogeneous. The type

SPeCies, fl. modesta (Nyl.) A.L. Sm., does not have the amyloid ring

found in some of the other species. I have recently located microcon-

idia in several species including fl. modesta and fl. hassei Zahlbr. and

they are filiform as in grenula. In Micrmlaena hassei, which also

lacks an amyloid ring, the spores are brown which also suggests the

PYrenulaceae. Therefore I feel that MicroLlaena 5. str. can be tenta-

tively assigned to the Pyrenulaceae. The species of Microglaena with

an am){loid ring require a new generic name. They are probably related

’50 Thrombium epiglaewn which also has this feature. Ve'zda (1968) has

3150 reported an amyloid apical apparatus in Geisleria. I have ex-

amiHEd a number of specimens of the type species, g. sychnogonioides

 

Nitschke, including the type collection, and have not been able to de-

tect an amyloid ring with either IKI or Melzer's reagent.

The Laureraceae Ve'zda ad int. (Poelt, 1974), created for genera

with muriform spores with thin spore walls, is I feel unnatural. There

Seems to be a tendency for the reduction and loss of mesospore thickening

in the evolution of muriform spores from septate spores with thickened

mesospore. In Anthracothecium and Parmentaria there are several species

27

which lack the characteristic thick spore walls. Also in the Trypethe-
liaceae there are several species with submuriform spores which connect
irypethelium and Laurera. Thus I feel that the genera placed in the
Laureraceae are better assigned to either the Pyrenulaceae or Trype-

theliaceae with the exception of Phyllobathelium of whose position I am

 

unsure.

VEZda (1968) has suggested that perhaps Thelopsis should be inclu-
ded in the Thelotremataceae. The genus seems to me to be out of place
in that family but I do not have enough evidence to suggest any other
placement. Thelopsis seems to me to be closely related to Belonia,
previously included by Ve'zda (1968) in the Porinaceae (= Trichotheliac-
eae), but seems to be excluded from that family on the basis of ascus
type and hymenial reaction with iodine. Not all members of the Tricho-
theliaceae possess the chitinoid ring in the ascus tip. It is lacking
in gathroporina and in some species of Porina. However the similarity .
of thallus and ascocarp type taken together with a series of submuri-

form Spored species linking Clathroporina to Porina leaves no doubt

 

that they are closely related.

The Pyrenulaceae and Trypetheliaceae show a considerable number of
parallel trends, astrothelioid and parathelioid ascocarps, inclusion of
ascocarps in pseudostromata, production of lichexanthone and various
pigments, thickened mesospore wall, well developed endospore in many
SPECies and similar ecology and distribution. Thus it is possible that
the families could be combined. However for the time being I feel that
the two families can be maintained on the basis of the differences in
Spore type (color and shape of lumina), paraphysis type, iodine reac-

tions in the hymenium and possibly in the method of spore liberation.

28

Although Poelt (1974) lists Normandina among genera of uncertain
position, I feel it is clearly a member of the Verrucariaceae. The
systematic position will be discussed in detail in another paper

(Pharris & Weber, in preparation).

V. SYSTEMATIC SECTION

A. A key to the genera of hyalodidymous and

hyalophragmous pyrenolichens of North America

Paraphyses gelatinized; hymenial gelatin IKI+ pinkish
(orange, orange red or bluish; almost always growing on

Jrock ............ ............. Verrucariaceae (not treated further)

I?araphyses persistent (or if gelatinizing, then hymenial

cgelatin IKI- and not growing on rock); growing on bark,

<>ld wood, leaves, bryophytes and rock . ................ ..... 2
2. GrOWing on leaves cocooooooo ooooooooooooooooooo 00000.... 3
2.. Growing on some other substrate .... ....... ............. 4

‘Asnzi thickened at the tip, lacking a chitinoid ring at

tilee tip of the exoascus; paraphyses sparsely branched

..--ooooooooooooo ooooooo ooooooooooooooo ooooooooo 00.000000. Stfigllla

Ascxi not thickened at the tip, with a chitinoid ring at

the? ‘tip of the exoascus; paraphyses unbranched .. ..... ...... Porina

4° .Ascocarp with stiff black hairs; ascus with a

(fliitinoid ring at the tip of the exoascus ....... Trichothelium

 

4° .Ascocarp smooth, without hairs . ........ ................ 5

AScOcarp with more than one chamber; ostioles separate
or'Illnited .................. ...... ......... ......... ........ 6
ASCOcarp with only one chamber (ascocarps may be aggre-

gatEd within a pseudostroma, but not fused into a

compound ascocarp .......... ......... ...... ......... ........ 9

29

"

 

3O

 

6. Ostioles separate .................................. Tomasellia

6. Ostioles united in a common disk or into a common

Opening 0.0000000000000000000000.00.00.0.000000000000000 7

Paraphyses mostly unbranched; hymenial gelatin IKI+ blue

 

green becoming orangish .............................. Lithothelium
7. Paraphyses branched and anastomosed; hymenial gelatin

8

IKI- 00.000000.0000000000000000000000000.0000000000000000...

£3. Spores with 4 or more cells; ascocarp very carbon-
aceous and hard, often pigmented outside ......... Astrothelium

£3. Spores with 2 cells, small, 8-12 x 3-4 p; ascocarp

not carbonaceous, rather soft .. ............... ....... Strigula

9- Ckstioles eccentric or lateral .............................. lO

9. OStiOles erect 00000000000000.000000000000000000000.00000000 l2

11)- Paraphyses mostly unbranched; hymenial gelatin

IKI+ blue green becoming orangish; microconidia

filiform 00000000000000.000000000.0000000000000000 Plagiocarpa
lO- Paraphyses branched and anastomosed; hymenial gela-

tin IKI-; microconidia rod-shaped ..................... 11

ll' SFKDres IKI-; mesospore not thickened ................. Pleurotrema

 

11' SFKJres IKI+ violet; mesospore thickened from the

beQimning ....................... Unnamed genus of Trypetheliaceae

12- Ascocarp wall blue green, HNO3+ reddish .............. l3

12. Ascocarp wall some other color ....................... l4

 

y
..e

....s

 
 

31

1;3- ,Asci thin tipped with a chitinoid ring at the apex of
the exoascus; paraphyses unbranched .. ...... .. ..... ........ Porina
1g3- Asci relatively thick tipped, lacking a chitinoid ring;

paraphyses branched ........ .......... . ...... . ...... ... Mycoglaena

 

$14. Asci with more than eight spores, thin tipped;

hymenial gelatin or ascus sheath IKI+ ............. Thelopsis

3.4. Asci with eight or fewer spores .... ........ .......... 15
15. 53;)cmes long cylindrical to filiform ....... ............ ..... 16
15 O Spores fUSifOI'm t0 OVal o o o o o o ooooooo o oooooooooooooooooo o o o o 18

115. Paraphyses branched and anastomosed; ascus tip
often appearing truncated; ascocarps simple or

multilocular ... ............... ......... Leptorhaphis 5. lat.

 

1€3- Paraphyses mostly unbranched . ........ ................ l7

17- «AEM2us with chitinoid ring at the tip of the exoascus ...... Porina
173 1A£M2us tip lacking a chitinoid ring ...... ..... ........... Belonia’
113- Mesospore thickened from the beginning; spore
lumina rounded or angular ............................ 19
18- Mesospore not thickened except in old age; lumina
cylindrical .......................................... 2O
\

 

(Hie only species of Belonia known from North America,
2? EESEEE§2§_Fink in_Hedr., is a non-lichenized fungus belonging to
the OStropaceae. Thus the genus should be removed from the North

American flora. Details will be published in Harris (1975).

.1E9-

lf9.

21.

21.

230

32

Thallus well developed, usually cartilaginous and shiny;

ascocarps mostly clustered in pseudostromata, but

scattered singly in some species .......... ........ .. Trypethelium
Thallus poorly developed, never cartilaginous, rarely
lion-lichenized; ascocarps never clustered within

gaseudostromata ........ ....... . ............. ....... Pseudopyrenula

220. Phycobiont a blue-green alga, blue-green, purplish
or yellowish brown in color; mostly on calcareous
rock, aquatic or in moist situations .......... Pyrenocollema
2H3. Phycobiont a green alga, usually Trentepohlia, or
non-lichenized; mostly on bark and old wood but

also on rock and soil, rarely aquatic . ......... ...... 21

Asci with a chitinoid ring at the tip of the exoascus;
paraphyses not branChEd ............O..............00...... porina
Asci lacking a chitinoid ring; paraphyses branched and

aIlastornosed except in Strigula ............................ 22

:32. Paraphyses mostly with only a few branches; macro-

conidia commonly present, septate, fusiform, cylin-

drical or rarely filiform; asci usually cylindrical;

spores often uniseriate ............................ Strigula
22. Paraphyses branched and anastomosed; macroconidia

rarely produced, non-septate when present; asci

often broader at one end; spores uniseriate or not ... 23

Spores oval to elliptic; spore wall ornamented with

Small granules, uniseriate ............................ Acrocordia

2L3-

25.

25.

33

Spores fusiform to ovate, most commonly with one half
of the spore shorter and/or narrower than the other, if
oval then not ornamented and uniseriate; often biseriate

or irregularly arranged in the ascus ........ ........ ...... 24

24. Microconidia globose to elliptical; spores 2(-4)-

celled, septum eccentric; Trentepohlia always

 

present ........... ........ .......... ...... .... Anisomeridium

 

224. Microconidia short or long rods, rarely filiform;
spores 2-many-celled, septum eccentric or not;

Often non-liCheniZEd o ooooo 0000...... ooooooooo 00...... 25

Eharaphyses rather loosely and reticulately branched;
liynnenium often inspersed; lichenized or not; spores
‘4-qnany-celled, endospore gradually filling lumen in

Emge, not becoming brownish or ornamented ... ....... .. Polymeridium

 

Ekaraphyses thick, short celledand appearing almost
Fnarenchyma-like to slender and thread-like or gelatin-
jJaing, but not loosely and regularly reticulate; very
rEtrely lichenized; endospore not filling spore lumen
jfl age, but spore wall may become brownish, thickened

anti ornamented in age .............. ................ ....... 26

26. Spores 2-celled, breaking into part spores at
maturity ........................... ....... ..... §poroschizon
26. Spores 2-many-celled, not breaking into part

spores at maturity . .............. .............. Arthopyrenia

 

34

B. PLEOSPORACEAE Wint.

Holotype: Pleospora Rabenh.

Mycoporaceae Zahlbr., Nat. Pflanzenfam. 1(1'): 77. 1903.

Holotype: Mycoporum Flot. _e_x_ Nyl.

Arthopyreniaceae w. Wats., New Phytol. 28: 107. l929. Holotype:

Arthopyrenia Massal.

Thallus mostly indicated by modification of the substrate, rarely

forming a superficial subicular layer of hyphae or in one case, epi-
lithic. A few species constantly or occasionally associated with

 

Trentepohlia, but most are probably saprophytic or possibly even

parasitic.

Ascocarps ascolocular in the two species studied by Janex-Favre

(1971 ) . Ascocarps simple or compound, when compound each locule with

its own ostiole. Ostioles erect or rarely eccentric in a few species

prOViSionally included here. Interascal filaments in the species

Studied by Janex-Favre (1971) are pseudoparaphyses. Pseudoparaphyses
branched and anastomosed, thick and irregular with short cells to
Slender and regular with relatively long cells, often somewhat gelatin-
ized or rarely completely so. Asci bitunicate (Richardson & Morgan-
Jones, 1964), bitunicate-nassascé (Janex-Favre, 1971), often broadest at
the base (Pleosporales type, Janex-Favre, 1971). Spores colorless or
brown, 2—celled to muriform, usually 8/ascus but rarely 2-4/aSCUS3
Spore Wall granular ornamented from the beginning in some, becoming so
in old age in a number; perispore usuall well developed.

Microconidia thin, rod-like, linear or filiform.

The above synonymy and diagnosis of the Pleosporaceae are not in-

tended to be comprehensive but only to include the taxa which have been

 

 

35

previously considered to be lichens. A few do indeed seem to be lichen-

ized but many are found only on smooth, thin bark where the hyphae might

well be able to invade living tissues and I suspect that they are para-

sitic, at least in part.

The species which I consider to be primitive are easily recognized

as members of this family by their ovate, thick tipped asci and coarse,

almost parenchyma-like paraphyses. More advanced species approach mem-

bers of the Strigulaceae or Trypetheliaceae on the basis of microconidial

type, lack of mesospore thickening at any stage, hymenial type and ab-

sence of Trentepohlia.

Arthopfienia and Tomasellia seem to be very closely related but

perhaps Mycoglaena should be included in some other family. However,

due to my lack of knowledge of the families of non-lichenized pyrenomy-

Cetes, Mycoglaena is placed in the Pleosporaceae more or less by default.

It seems noteworthy that the parathelioid and astrothelioid type of
aSCocarp is lacking in the Pleosporaceae and is confined to the lichen-

ized DYrenomycetes, whereas the Tomasellia type of ascocarp is lacking

in the lichenized families. Also, there is no development of pseudo-

Stromata as in the Trypetheliaceae or Pyrenulaceae.

1 . ARTHOPYRENIA Mas sal .

Ricerch. Auton. Lich. 165. 1852. Lectotype (Riedl, 1962): Verrucaria

 

rh onta Ach.

Marthopyrenia Keissl., Ann. Naturhist. Hofmus. Wien 34: 17. 1921.

H010type: fl. sorbi Keissl.

Cif

 

erriolichen Tomas. in Tomas. & Cif., Arch. Bot. (Forli) 28: 4.

1952- sflycociferria Tomas. i_nCif. & Tomas., Ist. Bot. Reale Univ.

 

36

Reale Lab. Crittog. Pavia Atti, ser. S, 10: 28, 56. 1953. Holotype:

Arthopyrenia lapponina Anzi.

Jatteolichen Tomas. & Cif., Arch. Bot. (Forli) 28: 6. 1952.

ggiatteomyces Cif. & Tomas., Ist. Bot. Reale Univ. Reale Lab. Crittog.

Pavia Atti, ser. 5, 10: 34, 61. 1953. Holotype: Verrucaria

erenastrella Nyl .

Santessoniolichen Tomas. & Cif., Arch. Bot. (Forli) 28: 5. 1952.

ESantessoniomyces Cif. & Tomas., Ist. Bot. Reale Univ. Reale Lab.

Crittog. Pavia Atti, ser. 5, 10: 29, 57. 1953. Holotype: §_. puncti-

formis ("Fr.") sensu Jatta [sic].

Giacominia Cif. & Tomas., Ist. Bot. Reale Univ. Reale Lab. Crittog.

Pavia Atti, ser. 5, 10: 29, 57. 1953. Holotype: Arthopyrenia parolinii
Beltram.

Pfiycoarthopyrenia Cif. & Tomas., Ist. Bot. Reale Univ. Reale Lab.

Crittoq. Pavia Atti, ser. 5, 10: 29, S7. 1953. horn. illeg. Holotype:

W analeptella Nyl.

Thallus endophloeodal, indicated by discoloration of the substrate,
Usually whitish or grayish, rarely darker, in a few species consisting

of an epiphloeodal dark brown subicular layer of hyphae, or epilithic in

a Single species. Phycobiont, when present, Trentepohlia.

ASCocarp with brown to blackish wall, usually lacking below. Osti-

ole erect or rarely eccentric. Paraphyses thick and irregular with
Short CIslls, often appearing almost parenchyma-like to slender and regu-

1ar With relatively long cells, sometimes partially gelatinized, rarely

Completfifly so. Asci various, ovate to narrowly ovate, narrowly ellipti-

cal, narrowly obovate or cylindrical; tip often thickened, with or with-

0Ut an obvious ocular chamber. Spores typically 8/ascus, often fewer

37
due to random abortion, rarely 2-4/ascus, mostly 2-4-celled but up to
8-ceflled, constricted at the septa; spore wall ornamented from the be-
ngIning in a few species, becoming ornamented in old age in a number;
puertispore usually evident, frequently well developed.
Microconidia commonly rod-like or linear, rarely filiform.
Habitat mostly on bark, often smooth, young bark, less commonly on
ecnaxrticate wood and on rock in a single species.
The typification of Arthopyrenia is a difficult problem which is

ccxnpilicated by the fact that Lichen analeptus Ach. (Arthopyrenia

 

arualtapta (Ach.) Massal.) is a superfluous name for Verrucaria olivacea

 

Peres. (= Porina olivacea (Pers.) A. L. Sm.). Thus genera which have

beerl typified with A. analepta are nomenclatural synonyms of Porina.

This; iiicludes Pyrenillium Clem. and Arthopyreniomyces Cif. & Tomas., as
well eas Leiophloea (Ach.) S. Gray if Riedl's (1962) lectotype is accepted.
The Exirliest typification of Arthopyrenia was with A. analepta by Th.
Fries 3J1 1861. This species was also chosen by Fink in 1910. However,
accepteulce of this priority would lead to the synonymization of fiEEEQf
£§3§§EEE_i~ith Boring, which certainly was not intended by either Th. Fries
or Fink ‘who could not anticipate the results of future nomenclatural

rule5° Thus, I feel that this is exactly the sort of situation covered

by Par- 4.f. of the Guide for the Determination of Types (Stafleu, et al.,
1972) and that the earliest lectotypification should not be accepted.
Clemenths (1909) chose A. pyrenuloides (Fee) Mull. Arg. as the type, but
this WEHS not one of the original species. Thus, Riedl's (1962) lecto-

type,;é’ rhyponta, has priority and must be accepted. In a similar

manner one could reject Riedl's typification of Leiophloea, but I leave

 

that tO others since I have no interest in resurrecting the genus.

 

38

 

Leigphloea has no history of usage and its synonymization with Porina is

of little significance. Also, it leaves Arthopyrenia as the oldest name

for the genus as treated here, including species with 2-celled and multi-

cel led spores.

The genus Arthgpyrenia in the sense which I accept it is primarily
non—lichenized although there are some exceptions. The bulk of the

lichenized species are those whose position is unclear to me at this time

and remain in Arthopyrenia by default. Others, a very few, seem to be

lichenized species of Arthopyrenia in the strict sense. The most inter-

esting of these is the West Coast population of _A_. padi which is con-
sistently associated with Trentepohlia, while collections from the rest

Of its extensive range seem to be non-lichenized. Another is the mari-

time species _A_. herrei which has a well developed thallus remininscent

0f firenocollema but on the basis of mycological characters seems to

bE10ng in Arthopyrenia. Of course, it is possible that it is a para-

Symbiont on another sterile lichen.

In addition to the absence of Trentgmhlia, the genus tends to be

marked by the thin microconidia which range from short and rod-like to

filiform- Many of the species have asci which are broadest at the base,

a . . . . . .
CharaCter which 15 relatively rare in the lichenized genera. However,

in the more advanced species the asci become cylindrical or broader at

the tip. This change in ascus type is more or less correlated with the

ParaphySis type which in the more primitive species is thick, irregular

and Short celled becoming slender and thread-like in the more advanced

ones. The spores are initially hyaline but in many species very old

spores become tinted and ornamented. The perispore is often well devel-

OPEd-' These characters taken together have been used to define

39

Arthopyrenia in a rather broad sense. The genus could be divided further

as Riedl (1962) has done by separating species with 2—ce11ed spores from

those with 4- or more celled spores. As I indicated in 1973, I feel that

this is artificial, separating closely related species as A. lapponina

from A. cerasi and A. padi from the A. persoonii group. A more natural

method would be to recognize two genera based on asci, paraphyses and

microconidia, one centering around A. padi the other around A. lapponina.

This would separate the species I consider to be primitive from the ones

I consider more advanced. However, the distinction is not sharp since a

few species, such a A. fraxini and A. antecellens, are rather intermed-

iate and, therefore, I prefer to maintain a single genus. - The separation

Of Ethopyrenia from Tomasellia and Polymeridium is discussed under the

latter genera.

l. Spores 4’Celled 0000. oooooooooooooooooooooooooooooooo 00.0.00 2
1' SPKlres 2-celled, rarely 4-celled in old age ................ 5
2- Thallus dark brown; hyphae thick, epiphloeodal;
spores 17-20 x 5.5-7 p .......................... [fir rhyponta]
2- Thallus whitish, grayish or little different from
the surrounding bark .. .......... ..... ........... ..... 3
3.

ASC1<Dcarp immersed, the tip surrounded by a broad, thin
Shiiéld often confluent with other shields; ostiole often
SUIUCQunded by a whitish ring; spores 17-22 x 6-8 p;

microconidia 6-9 le 0.00.00.00.00. ........ 00-00000... £0 COUfluenS

3. Ascocarp without a broad, thin shield, not confluent;

OStiole without a whitish ring ... ................ .......... 4

4O

4. Asci ovate; spores 17-21 X 5—6 p .............. A, atomarioides

 

4. Asci narrowly ovate to cylindrical, 60-90 X 17-20 u;

spores 17-23 x 6-7.5 u .. ......................... [A3 cerasi]

55- Spore wall constricted in the middle of one or both cells

or mesospore thickened as if additional septa were form-

ing, becoming 4-CellEd in Old age .0 ...... ... ...... ......00. 6
55. Spore wall not constricted in the middle of the cells;
Inesospore not thickened ........... ..... ..... .......... ..... ll

6. Spore wall ornamented; mesospore often forming a

ring cutting off a subchamber; North Carolina to

Florida and Texas (A. bifera group) ................... 7
6. Spore wall not ornamented; no subchamber partially

cut off; West Virginia and Tennessee to Newfoundland

or western ......OOOOOOOOOOO00.000.00.00...00.0.0000... 9

7' SFKJres 2/ascus, 37-48 x lS-lS‘u; ostiole occasionally
eccentric ......OOOOCO ..... .............OOOOOOOOOOOOCOCOOO A. bifera
7’ SPCnres 4-8/ascus .................. .................. ....... 8

8- Spore wall with a flat-topped, ring—like ridge with-

in each cell which in old age forms a complete

septum; spores 27-37 x lO-12 p .... ........ ........ A, annulata
8. Spore wall usually not thickened, merely constricted

at the middle of each cell; spores very rarely 4-

CGIlEd in Old age, 18-30 X 605-905(’12) p 0.0.00.0... fir lzrata

9- Spores 12-lS(-18) x 4-5 11; constrictions rather weak .6: degelii

9.

11.

ll.

l3.

13.

41

Spores larger, (13-)lS-26 X 4.5-9 p; wall rather strong-

ly constricted ......OOOOOOOOOOOOO......OCOCOOOO......C..... lo

10. Spores with both ends rounded, upper cell usually
somewhat larger, 18-22 X 7-8(-9) u; asci mostly

elliptical or obovate, 60—80 X l7—22 u .... A. cinereopruinosa

 

10. Spores with one or both ends pointed, cells approx-
imately equal, (134)15-20 X 4.5-6.S(-7.5) u; asci
mostly narrowly elliptical, narrowly ovate or nar-

rowly obovate, 75-llO X 12-18 p .... ...... ....... A, plumbaria

Thallus well developed, brown, epilithic; spores l4-l9 X
405-5051], 000....... ................................ 0.... A. herrei

TFhallus poorly developed, endophloeodal ................... 12

12. Paraphyses completely or almost completely gelatin-

ized; spores (12-)lS-22 X 4.5-6 u; asci ovate to

narrowly ovate, 35—50 X 15—20 p ................... A, salicis
112. Paraphyses persistent, occasionally somewhat gelat-

inized or embedded in much gelatin but not disap-

pearing ......OOOOOOOOOOOOOOO..... ..... ............... 13

Spores ovate, length/width ratio l.5:l-2:l, cells mark-
edly unequal, 13-15 X 7-8(-10) u ......0.........00... f... Oblongens
E5pores narrowly ovate or narrowly elliptical, length/

”Width ratio 2.5:1 or greater .. ......... ......... ......... l4

14. Asci cylindrical, narrowly elliptical or narrowly
Obovate ............ ..... ............................. 15

l4. Asci narrowly ovate, broadest below the middle ....... l9

42

15. Spores mostly less than 23 u long and 7.5 p wide .......... 16

15. Spores mostly more than 20 u long and 7‘p wide .. .......... 18
16. Spores 12-17X4-5.5p ......0....0. ..... 00.....0.... A. minor
16. Spores 15'23X5'705u 0.0.00... ........... ......0.... 17

137- Spores 15-22 X 5.5-7.5 u; ascocarp wall not extended
outward into a broad shield; Massachusetts and Europe .....
............... ....................... ................ A, lapponina
137. Spores l7-23 X 5-7 p; ascocarp wall extended outward

forming a broad shield; Florida and the West Indies .. A, planorbis

118. Ascocarp immersed, tip surrounded by a broad, thin

hyphal ring; spores 20-28(-32) X 6—8(-9.5) u; on

Taxodium ................. ..... .................... A, taxodii
li3. Ascocarp superficial, hemispherical, no hyphal ring

present; spores 20-30 x 7-9.5(-ll) u; on a wide

variety of smooth, thin barks ................... A, cinchonae

19' SFXDres larger, (21-)25-4O p in length ..................... 20

19° SEKDres smaller, 25 p or less in length .................... 22

20- Spores broader, (25-)28-4O x 8-12(-l4) 11., becoming
4-celled, brownish and ornamented with age .... A, antecellens
2C)- Spores narrower, 6-8 p in width, not obviously

becoming 4-celled, brownish and ornamented ........... 21

21‘ ASCocarps subglobose, immersed, 0.1-0.15 mm in diameter;
Spores 21-33 X 6-7.5 p; Florida and Alabama ............ A, cedrina

21- Ascocarps flattened, superficial, 0.4-0.6 mm in diameter;

 

43

spores 25-32 x 6-8 p; Maine ... ................. ... A. megalospora

 

22. Spores larger, 19-25 X 5.5-7.5 11.; ascocarps hemi-
spherical or flattened, 0.3-0.5 mm in diameter; north-
eastern United States and Europe ..................... . A. fraxini

22- Spores smaller, 15-22 X 4'505 p .0000 ................. 0.0.0 23

23. Asci slender, 55-70 x 12-16 11; Florida and

Alabama . ............................. ....... A. atractospora
23. Asci shorter and stouter, 30-60 x 15-24 1).; spores

15-20 x 4.5-5.5(-6.5) 1.1.; widespread northward,

Maine to British Columbia ....................... ..... A. padi

BLT-hOpyrenia antecellens (Nyl.) Arn.

Flora S3: 485. 1870. Verrucaria antecellens Nyl., Flora 49: 86.

1866- flycoporopsis antecellens (Nyl.) Riedl, Sydowia 15: 268. 1962

m- inval. Holotype: England, Sussex, Tilgate Forest, Larbalestier

fl (H-NYL 767).

 

menula zwackhii Hepp, F1. Eur. 954. 1867. Type collection:

Germany , near Heidelberg, Zwackh. Isotypes in FH & FH-TUCK 4066.

Ascocarps superficial, hemispherical to flattened, 0.3-0.6 mm in
diamEter; ascocarp wall brown, not heavily carbonized, lacking below.
Paraphyses thick, up to 3 u, somewhat irregular. Asci ovate to narrowly
ovate, (65-)80-105 x (20-)27_35(-4o) p. Spores irregularly arranged,
narrowly ovate, 2-celled, becoming 4-celled, brownish and ornamented in

Old age; perispore usually not obvious; (25-)28-40 x 8-12(-14) u.

Microconidia short, rod-like, 3-4 x l u.

44

Habitat on smooth bark.

Zahlbruckner (1921-22) incorrectly placed Pyrenula zwackhii as a

synonym of Thelidium zwackhii (Hepp) Massal. based on Sagedia zwackhii

 

Hepp, a different independently published species.
Arthopyrenia antecellens has not previously been reported from

North America, but it is easily recognized by its large spores and ovate

asci. It seems nearest to the A. padi group but the paraphyses are

somewhat intermediate with those of the A. lapponina group. Based on

specimens which I have verified, A. antecellens has a rather oceanic

 

distribution both in North America and Europe.
Illustrations: Figs. 1-6.
Iixsiccati examined: Hepp 954 (FH, FH-TUCK 4066).
iipecimens seen: Canada. BRITISH COLUMBIA: Queen Charlotte

Islarnis, Graham Is., Brodo 11603, 12901a (CAN), Lyell Is., Brodo 11922

 

(CAN, IWSC). NEWFOUNDLAND: Bay of Islands, Lark Harbour, 1887 Waghorne

fl (MICH, US ex MO).

Urlited States. MAINE: Sagadahoc County, near Brunswick, 3.XI.1939

.EgflgiiEEi (U5: US ex MO). NORTH CAROLINA: Haywood County, Blue Ridge
Parkw3Y’ lookout at Graveyard Fields, Harris 3397-A (MSC). OREGON:
“Ll’Leloxcq 1871 Hall _(FH-TUCK 4048). WASHINGTON:

Jefferson County,

5 mijB (of Lake Quinault, Thcker 8365 (LSU).

W9. atomarioides Mull. Arg.

BCNZ. Jahrb. Syst. 6: 406. 1885. Holotype: Cuba, Wright Verr.

CUb- 11: 629 (G).

YErrucaria subpunctiformis Nyl., Lich. Ins. Guineens. 51. 1889.

Arthogyrenia floridana Zahlbr., Cat. Lich. Univ. 1: 332. 1922 E nom.

45

r1c3\r- Type collection?: Florida, Ft. George, 1887 Calkins. Isotypes? in

FH, MICH, US & us ex MO.

Ascocarps superficial or partly immersed, hemispherical, 0.1-0.2 mm
,irl cijLameter; ascocarp wall brown, lacking below. Paraphyses thick and
iirxreeggular, somewhat gelatinized. Asci ovate, 35-50 X 18-22 u. Spores
iirxreecgularly arranged, narrowly ovate, soon 4-ce11ed, a few becoming
6-cel1ed; perispore not obvious; 17-21 X 5-6 p.

Habitat on smooth bark.

Arthopyrenia atomarioides can be recognized by its small ascocarps,
a§3<3iL and 4-ce11ed spores. Nylander credited the type collection to
Iacflfif‘e-ldt, but I have seen no North American material except the single

CalICJ'Lns collection which is uniformly mixed with Tomasellia lactea.

 

Illustrations: Figs. 7-11.
fEElgfigfigpyrenia atractospora Zahlbr.

Ann. Mycol. 33: 33. 1935. Holotype: Florida, Sanford, VII.1928

$22 3.21m).

Ascocarps immersed, subglobose, c. 0.2 mm in diameter. Paraphyses
Scxnfifluhat gelatinized. Asci narrowly ovate, wall strongly thickened in
time, tip, 55-70 X 12-16 u. Spores irregularly arranged, narrowly ovate,
2"Celled, lower cell often somewhat longer and narrower; perispore not

Ob"’ious; 16-22 x 4-5 p.

Microconidia rod-like, 4-5 X 1 u.

Habitat on bark.

Arthopyrenia atractospora is characterized by slender asci with

the tip strongly thickened and by its slender spores. It is known from

46

on 1 y two collections .
Illustrations: Figs. 12-15.

Specimens seen: United States. ALABAMA: Baldwin County, Yupon

EijLr112, 1.III.1925 Evans 210 (MICH). FLORIDA: Seminole County, Sanford,

VII- 1928 Rapp 137 (W).

Pu:t:k1<3pyrenia cedrina (Zahlbr.) R. C. Harris stat. nov.

Arthopyrenia sanfordensis var. cedrina Zahlbr., Ann. Mycol.

323: :33. 1935. Holotype: Florida, Sanford, X.1922 Rapp 135 (W).

Thallus endophloeodal, Trentepohlia present.
Ascocarps semi-immersed, subglobose, c. 0.15 mm in diameter.

IParfiaphyses thick, up to 3 u in.width, irregular, somewhat gelatinized.

Ad3<3ii narrowly ovate, 60-90 x 20-25 u. Spores irregularly arranged,
nail:‘J’Jowly ovate, 2-ce11ed; perispore thin; 21-33 X 6-7.5 u.

Microconidia short, rod-like, c. 3 X 1 u.

Habitat on bark.

 

Arthopyrenia cedrina is not closely related to A, sanfordensis

Mfl‘jscfli is a member of the genus Anisomeridium. Arthopyrenia cedrina

jJS CDJite similar in paraphyses, asci and spores to A, fraxini but

diffiiers by its small subglobose ascocarps and thinner perispore.

3329Trtepohlia is present in both of the known collections.
Illustrations: Figs. 16-19.

Specimens seen: United States. ALABAMA: Baldwin County,

V01anta, 9.III.1925 Evans 258 (MICH). FLORIDA: Seminole County,

Sanford, X.1922 Rapp 135 (W).

-.
..
..

. _

‘0-0 ..
.

- ' "

.7 ‘—-

:'- ...

-.. _

"v,.

0 ~.'

 

 

 

 

47

ZX171:knopyrenia cerasi (Schrad.) Massal.

Ricerch. Auton. Lich. 167, f. 332. 1852. Verrucaria cerasi Schrad.,

 

,Axixu,. Bot. (Usteri) 22: 86. 1797. Type collection not seen.

Ascocarps superficial, hemispherical to flattened, (0.2-)0.4-0.5 mm
ir1 (fl;iameter; ascocarp wall lacking below. Paraphyses slender, regular
aIICi 1:hread-like. Asci narrowly ovate to narrowly elliptic, rarely
rmaxrir<3wly obovate, 60-90 x 17-20 p. Spores biseriate to irregularly
alrxreardged, narrowly ovate, 4-ce11ed; perispore often rather thick;
17—23 x 6-7.5 u (excl. perispore).

Microconidia linear, 10-13 x l u.

Habitat on smooth bark.

Arthopyrenia cerasi seems to me to be rather closely related to

 

ii- ldapponina, differing in having 4-ce11ed spores. Although reported

firCDnn bkmth America, I have not been able to verify any specimens of it
£3r1C1 Lit should be deleted from the North American flora.
Illustrations: Figs. 20-23.
Exsiccati examined: Anzi Ven. 130 (FH-TUCK 4077); Harm. Loth. 450
(PH); Hepp 457 (FH, FH-TUCK 4081); Malbr. 400 (MSC); Mass. 106 (FH-TUCK
4081), 219 (FH-TUCK 4083); Norrl. 393 (MSC); Nyl. Pyr. 50 (FH-TUCK
4081); Rab. 145 (MICH, FH-TUCK 4082); Samp. 22 (MSC).
EEZEiygpyrenia cinchonae (Ach.) Mull. Arg.

Flora 66: 287. 1883. Verrucaria cinchonae Ach., Synops. Lich. 90.

 

1814. Holotype: America, on Cinchona officinalis (H-ACH).

Verrucaria prostans Mont., Ann. Sci. Nat. Bot., ser. 2, 19: 53.

 

1843. Holotype: French Guiana, Leprieur 215 (PC-MONT).

 

Verrucaria alboatra var. detergens Nyl., Flora 52: 125. 1869.

 

-..
l.
.
-

 

 

I

, .

 

I

48

ijijéi collection: Brazil, near Rio de Janeiro, Glaziou 1915. Isotype

.iri D4-

Ascocarps superficial, hemispherical to flattened, 0.4-0.6 mm in
<dj_ELnn€eter; ascocarp wall lacking below. Paraphyses slender, regular and
tlixreeaad-like. Asci mostly narrowly obovate, rarely almost cylindrical;
<ocru1]_ear chamber mostly not evident; (85-)100-125 X 17-22 u. Spores bi-
seexriLcate, subbiseriate or almost uniseriate, occasionally only 4/ascus,
inélxrxrcowly ovate, 2-celled, lower cell occasionally slightly constricted
ir1 1:}1e middle; perispore well developed; 20-30 X 7-9.5(-1l) u.

Microconidia rod-like, 4-5 x 1 u.

Habitat on smooth bark.

Arthopyrenia cinchonae can be recognized by the rather large asco-

 

<2511TID£3, asci and spores. However, it has commonly been confused with
51‘ Zlgfirata, which has more nearly cylindrical asci, not broadening to
‘tk‘ei “tip, ornamented spores with one or both cells constricted near the

”ELCiCilcn longer microconidia and is always associated with Trentepohlia.

Arthopyrenia cinchonae is widespread and often very abundant in the

 

SOLltheastern United States occurring as far north as New Jersey. Of all
thEE onrenolichens it most truely deserves the title of "weed". It is
£3143C> known from Brazil, French Guiana, Mexico and the West Indies
(Befiiamas, Bermuda, Cuba, Dominica, and Puerto Rico).
Illustrations: Figs. 24-30. Distribution: Fig. 406.
Exsiccati examined: Rel. Tuck. 130 (FH, MICH, MSC, US).
Selected specimens seen: United States. ALABAMA: Baldwin County,
Volanta, 20.III.1925'§2§A§.;AQ (MICH); Cleburne County, N of Heflin,
Harris 1328-A, 1338-A, 1392, 1406, l407-A, 1425 (MSC); Lawrence County,

Mmflton, 1874 Peters (FH-TUCK 4025). FLORIDA: Alachua County,

 

1
I
I
l

D

 

' 0
.-._

' o
.. .
—
A.

2"
d

J"

u.

'0._

0" r

'4

. s

u

9‘-
~~

-.

 

 

 

 

 

 

‘.
.»

 

 

49

Gainesville, IV.1915 Nelson 274 (FH); Baker County, Osceola Nat. Forest,
Harris 3127, 3129-B (MSC); Dade County, Everglades Nat. Park, between

Mahogany Hammock and Pa-hay-okee Overlook, Harris 29l6-C, 2946-C, 2966-A,

 

2968—A (MSC), near Pa-hay-okee Overlook, Harris 27l7-A, 2718, 2729,

 

274l-A, 2742, 2753-8, 2757-A, 2763, 2765-B, 2795-A, 2829-B, 2836-8 (msc),

 

near Pinelands Trail Area, Harris 2879-A, 2884-A, 2897-B, 2898-B,

 

@05—8, 2911-E, 2986-E, 2988-A, 3017 (MSC), w. Palm Beach, 1897 Thaxter

5‘0 (FH); Duval County, Bayard, IV.1910 Bussey (FH, US), Jacksonville,

gal-kins (FH, MICH, US); Flagler County, Flagler St. Park, Harris 2481-A,

 

w, M (MSC); Franklin County, W of St. Teresa, Harris l741-E

 

(MSC); Liberty County, Apalachicola Nat. Forest, Whitehead Lake Camp-
ground, Harris 1456-8, 1583-D, 1785 (nsc); Marion County, 0ca1a Nat.
FCDJTest, S of Long Pond, Harris 1803-A (MSC), NW of Big Bass Lake, Harris

%r 2054, 2060-A (MSC); Sarasota County, Myakka River St. Park,

N 2586, 2623-A (MSC); Seminole County, Sanford, AV.1908 Rapp (FH),
11- IV.l9l4 Rapp (FH); St. Johns County, Six Mile Creek, Calkins (FH,

MICH, MSC, US); Volusia County, Daytona [= Daytona Beach], 1898

W 350 (MICH); Wakulla County, St. Marks Nat. Wildlife Refuge,
N 1733-B (MSC). KENTUCKY: MCCreary County, SW of Corbin, Harris
% (MSC). LOUISIANA: E. Feliciana Parish, Felixville. Tucker.

% (LSU); Iberia Parish, Avery Is., 11.1915 MCIllenny (FH-RIDD);

St. Landry parish, Grand Coteau, 6.VI.1894 Langlois (vs); St. Martin
pafish, Breaux Bridge, 12.XII.1893 Langlois (US), St. Martinville,
8~VII.1892 Langlois (US); St. Tammany parish, Covington, 16.Iv.1894

L. anglois 9_40_ (US), near Tallisheek, Tucker 10302 (LSU); Washington
Parish, E. of Mount Hermon, Tucker 10255 (LSU). NEW JERSEY: Atlantic

County, Atlantic City, l2.VIII. 1887 Green (MICH); Camden County, Atco,

u
_
A
~'.

’---._

 

 

50

:1535322 Eckfeldt (CAN). SOUTH CAROLINA: County unknown, Santee Canal, 1857
Ileaxrtanel (FH,MICH, MSC, US); Aiken County, Aiken, Ravenel (FH); Berkeley

CZc>11r1ty, Francis Marion Nat. Forest, Bonneau Campground, Harris 3157-A

 

(b453CZ), Guilliard Lake Campground, Harris 3211-A (MSC); Pickens County,
Dit:- Pinnacle, 10.VII.1886 Green (US). TENNESSEE: Wilson County,

(Zeecieaxs.of Lebanon St. Park, Harris 1318-A (MSC). TEXAS: sine loc.,

Hall (US).

iiETtiklopyrenia cinereopruinosa (Schaer.) Massal.
Symmict. Lich. 117. 1855. Verrucaria cinereopruinosa Schaer.,
IJj—Cifi. Helvet. Spici1., sect. 6: 343. 1836. Type collection not seen.
Pyrenula punctiformis var. cinereopruinosa f. hederae Hepp, FleCht.

13111?.. 105. 1853. Type collection: Switzerland, Zfirich, Hepp. Isotypes

in PB & FH-TUCK 4073.

Pyrenula punctiformis var. cinereopruinosa f. pinicola Hepp,
I?1—Efl:ht, Eur. 106. 1853. Arthopyrenia pinicola (Hepp) Massal., Symmict.
Ifjvcfln. 118. 1855. Type collection: Switzerland, Zurich, Hepp, Isotypes
in PH & FH-TUCK 4073.

Arthgpyrenia stigmatella var. elabens Massal., Symmict. Lich. 120.
153EN3. Type collection: Mass. 202, Italy, "ad truncos pinorum in Prov.
Patavinia (Rua)". Isotypes in FH & FH-TUCK 4077.

Pyrenula punctiformis var. cinereopruinosa f. buxicola Hepp ex

Rabenh., Lich. Eur. 630. 1862. nom. inval. sin. descr.
Arthopyrenia fallaX var. conspurcata J. Stein. iA_Beck & Zahlbr.,
Ann, Naturhist. Hofmus. Wien 12: 94. 1898. Type collection: Krypt.
Vind. 269, Austria, Carinthia, woods below castle Hornstein near

Iflagenfurt, g, Steiner. Isotype in US.

51

Arthopyrenia ligustri Britz., Hedwigia (Beib1.): (37). 1904.

Idcnnn- illeg. Syntype collection: Britz. 385, Germany, Augsburg,

vveassi:liche Hugelzug, Britzelmayer. Isotype in US ex MO.

Ascocarps superficial to semi-immersed, hemisphereical to flattened,
(D- 22-(3.3(-0.4) mm in diameter; ascocarp wall lacking below. Paraphyses
s]_eerdkjer, regular and thread-like. Asci mostly elliptical or obovate,
:rEixreeiLy narrowly so, 60-80 X 17-22 u. Spores irregularly arranged,
1161:?1rxowly ovate to narrowly elliptical, 2-celled, one or both cells
S<>nneendmat constricted near the middle, upper cell often broader and
:LCDIlggeer, rarely becoming 4-celled; perispore usually well developed;
18—22 x 7-8(-9) 11.

Microconidia linear, 10-12 X l p.

Habitat on smooth bark.

The transfer of Verrucaria cinereopruinosa to Arthopyrenia was

 

attributed by Zahlbruckner (1921-22) to Korber, but it seems fairly

Clear that Massalongo's publication was earlier.
Arthopyrenia cinereopruinosa seems fairly closely related to

£EF :Lapponina but differs in that the spore cells are constricted in the

““jJflxile. Arthopyrenia plumbaria is very closely related to A, cinereo-

EEZEEL22§2,3nd is distinguished by its more slender asci and spores and
1J1 llaving rather pointed spores. Arthopyrenia cinereopruinosa is
CCmHnon in Europe but apparently rare in North America where its

distribution seems to be oceanic.
Illustrations: Figs. 31-35.
Exsiccati examined: Anzi Ven. 129 (FH-TUCK 4077); Arn. 1262,
1740 (MICH); Arn. Mon. 420, 449 (MICH); Britz. 51, 385, 386 pr. p.

(US eX MO); Desm. ed. II, ser. II: 598 (FH); Erb. II: 419 (FH);

52

Hepp 105, 106, 455 (FH, FH-TUCK 4073), 107 (FH, FH-TUCK 4076), 456 (FH-
TUCK 4066); Krypt. Vind. 269 (US); Leight. 197 (FH, US); Malbr. 199 (MSC);
Mass- 197, 199, 200, 203 (FH-TUCK 4077), 198, 201 (FH), 202 (FH, FH-TUCK
14(3'7”7'); Moug. 364 pr. p. (FH-TUCK 4069); Rab. 328, 630, 659 (MICH, FH-
fTLJCZI<Z 4073, 4076); Schaer. 287 pr. p. (FH-TUCK 4072); Trev. 29, 41, 42,

4:3 (ZFH).

Specimens seen: Canada. NEWFOUNDLAND: Bay of Islands, John's

136351<:}1, Waghorne 367 (US ex MO), Middle Arm, Waghorne 463 (MICH).

 

United States. CALIFORNIA: Santa Cruz County, Stevens Creek
(:Eifizgkon, 24.XI.1906 Herre (FH). OREGON: County unknown, Upper Willamette
Valley, Sipe 716 (MICH). VERMONT: Addison County, Goshen, 8. III.1925

Eton 2312 (m).

fizflEjEilppyrenia confluens R. C. Harris nom. nov.

Tomasellia (s. Oligomeris) leucostoma Mfill. Arg., Flora 68: 257.

 

163855. (non Arthopyrenia leucostoma (Fee) Massal., Ricerch. Auton. Lich.

{167C). 1852). Holotype: "in Cort. Cascarillae ex hb. Hampe, 1877". (G).

Ascocarps solitary or grouped, immersed, hymenium pyriform to
)1efintispherical, 0.1-0.2 mm in diameter; ascocarp wall poorly developed,
lacIking below; tip of ascocarp surrounded by a broad, thin, black ring,

O'2‘~O.3 mm in width, often confluent with the rings of other ascocarps

folfining groups of 2-10 ascocarps. Ostiole often depressed, surrounded

k?! a whitish ring. Paraphyses slender, regular and thread-like. Asci
m0Stly narrowly ovate, rarely almost cylindrical, 60-105 X 16-26 p.
SPores irregularly arranged to biseriate, narrowly ovate to narrowly
elliptical, 4-celled; perispore well developed; 17-22 X 6-8 p.

Microconidia linear, 6-9 x l u.

53

Habitat on bark.

Although A, confluens at first glance has the appearance of a

Ikornzasellia, the structure of the ascocarp is not truely compound as in

 

iikiaat: genus. Numerous solitary, simple ascocarps can be found and the
ggxrc>L1ps are formed by the fusion of the thin ring—like shields sur-
ITCJLlrlding the tips of the ascocarps. Also the slender thread-like
EDELxreaphyses would exclude this species from Tomasellia. Arthopyrenia

<2c1rljfluens has been found in southern Florida and the Bahamas.

 

Illustrations: Figs. 36-39.

Specimens seen: United States. FLORIDA: County unknown, Constant
Springs, XII.1880 Hitchcock (US ex MO); Monroe County, Key West, I. 1898

T‘haxter 122 (MICH, MSC), II.1898 ‘I'haxter 301, 324a (MICH).

 

 

fiiilgklopyrenia degelii R. C. Harris sp. nov.

Arthopyrenia ascis sat parvis, 48-60 X 11-13 u et sporis parvis,

 

bilocularibus cum uno vel ambis cellulis in medio constrictis, 12-15
)C ‘qw-S p.
Holotype: Tennessee, Great Smoky Mountains, Cherokee Orchard,

:L]--IIX.1939 Degelius (US). Isotypes in FH & US eX MO.

Ascocarps superficial, hemispherical to flattened, 0.3-0.5 mm in
(iiiflneter; ascocarp wall lacking below. Paraphyses thread-like, regular,
upto 2‘p in width. Asci narrowly elliptical, 48-60 X 11-13 u. Spores
biSeriate to subbiseriate, narrowly elliptical, 2-celled, one or both

Cells constricted near the middle; perispore thin; 12-15(-18) x 4-5 p.

Microconidia rod-like, 6-8 x 1.5 u.

Habitat on smooth bark, possibly always Hamamelis.

54

The type of paraphyses and spores shown by A, degelii would seem to
‘place it near A, cinereopruinosa. The constriction of the spore cells
:is often slight and the species could be confused with A, migor, but the
huroader microconidia of the former along with the differences in geo-
<giraphy and substrate help to separate them if the constrictions are
cnrerlooked. Arthopyrenia degelii has been most frequently collected in
UVEast Virginia but is also known from Tennessee and Massachusetts.

Illustrations: Figs. 40-43.

Specimens seen: United States. MASSACHUSETTS: Bristol County,
rqewv Bedford, Willey (US). TENNESSEE: Sevier County, Cherokee Orchard,
ll.Ix.1939 Degelius (FH, US, US ex MO). WEST VIRGINIA: 5.3363139,
SkilJcins (FH); County unknown, Benaverti, 31.XII.1880, coll. unknown
79x9 (MSC); Fayette County, 3133139., 1893 Nuttall _8_5 (FH, MICH);
<3I‘eenbrier County, Greenbrier Run, 1.VI.1880 [EEEQE?] (FH); Taylor

County, Grafton, 17.XII.1880 coll. unknown (Msc).

.fflaihopyrenia fraxini Massal.

Ricerch. Auton. Lich. 167, f. 333. 1852. Type collection not seen.

Ascocarps superficial, hemispherical to flattened, 0.3-0.5 mm in
ciiameter; ascocarp wall lacking below. Paraphyses quite slender, but
J:‘Eather irregular and tending to gelatinize. Asci ovate to narrowly
C”iate, 60-90 X l6-25(-30) u. Spores irregularly arranged, narrowly
‘Ovate, rarely narrowly elliptical, 2-ce11ed; perispore often rather
thick; 19-25 X 5.5-7.5 u.

Microconidia rod-like, 4-5 x l u.

Habitat on smooth bark, most often on young bark of Fraxinus.

The spores of A, fraxini approach those of A. lapponina in size

55

and shape, but the former has ovate asci, shorter macroconidia and less

iregular paraphyses. Arthopyrenia megalospora may be a large form of
‘tfuis species and is discussed further under that species. It is largely

tjieashape of the asci and the small microconidia which are diagnostic

for A. fraxini.
Illustrations: Figs. 44-48; Harris (1973), figs. 8-9. Distribution:

Fig. 407.

Exsiccati examined: Anzi It. 385 (FH-TUCK 4071); Fellm. 222 (FH-

TUCK 4072); Fr. 242 (FH-TUCK 4067); Hepp 453 (FH pr. p., FH-TUCK 4066);

K33Y13t. Vind. 268b (FH), 1523 (US); Mass. 184 pr. p. (FH-TUCK 4078),
298A, 8, 299 (FH-TUCK 4070); Merr. I: 281 (us), II: 20 (MICH, MSC, US,

US ex MO, WIS); Rab. 146 (MICH, FH-TUCK 4071), 658 (MICH, FH-‘I’UCK 4078).

Additional specimens seen (notejjxxiin Harris, 1973): United

States. MAINE: Knox County, Rockport, 20.IV.1910 Merrill (FH).

 

MINNESOTA: Chicago County, N shore of Clear Lake, Tucker 1529-B (LSU).

‘VEHRMONT: Addison County, Goshen, 3.III.1926 Dutton 2454 (PH).

-£Eithopyrenia herrei R. C. Harris sp. nov.

Arthopyrenia thallo epilithico, fusco et gelatinoso, ascis

g1Tacilibus, 65-85 X 12-15(-20)'p et sporis parvis bilocularibus,

l41,-19 x 4.5-5.5 11.

Holotype: California, San Francisco County, San Francisco, POint
LObos, 19.VII.1906 Herre 889 (FH). Isotypes in FH, MICH & US.

Thallus well developed, epilithic, brown to dark brown, gelatinous

Mmen wet. Phycobiont apparently Trentepohlia.

Ascocarps very numerous, semi-immersed, subglobose, 0.2-0.3 mm in

56

diameter; ascocarp wall lacking below. Paraphyses thread-like but

.rather irregular and short celled, up to 2.5-3 u in width. Asci
Idarrowly ovate to cylindrical; wall strongly thickened at the apex;

651—85 X 12-15(-20) u. Spores irregularly arranged, narrowly ovate,

2-«:elled; perispore not obvious; 14-19 x 4.5-5.5 u.
Microconidia rod-like, 4-5 x l u.
Habitat on maritime rock (sandstone).

Although A, herrei is known from only a single collection, there is

anngble material and it is a most unusual species. As a rule the genus

EEEtfilopyrenia is corticolous and non-lichenized, while A, herrei is saxi-

 

CCfiLCNJs and has a well developed thallus. It is possible that it is a

Exarfasymbiont on an otherwise sterile lichen thallus, but without addi-
‘ticnlal material the problem cannot be resolved.

Illustrations: Figs. 49-52.

EEEiglopyrenia lapponina Anzi

Comment. Soc. Crittogam. Ital. 2: 25. 1864. Type collection:

‘Arnli Lang. 347, Italy, Bormio, Agzi. Isotype in FH-TUCK 4084.
Verrucaria epidermidis var. fallax Nyl., Bot. Notiser 1852: 178,
pl. 1, f. 12b. 1852. Verrucaria fallax (Nyl.) Nyl., Flora 55: 363.
18'72, Arthopyrenia fallax (Nyl.) Arn., Verhandl. Zool.-Bot. Ges. Wien
23: 505. 1873. Leiophloea fallax (Nyl.) Riedl, Sydowia 23: 234. 1971.
HOlotype: Finland, Holmia, Nylander (H-NYL 955).
Pyrenula punctiformis var. fallax f. betulae Hepp, Flecht. Eur.

452. 1857. Type collection: Switzerland, Zfirich, Hepp. Isotype in FH.

Arthopyrenia fallax f. crategina J. Stein. $2 Zahlbr., Ann.

Naturhist. Hofmus. Wien 24: 283. 1911. Type collection: Krypt. Vind.

57

1763a, Yugoslavia, "ad litus meridionalis lacus "Veldeser See", Steiner.
LIsotypes in FH & US.

Ascocarps superficial, hemispherical to flattened, 0.3-0.6 mm in
Chiameter; ascocarp wall lacking below. Paraphyses slender, regular and
tliread-like. Asci narrowly elliptical, 75-90(-110) X 15-23 u. Spores
laigseriate to subbiseriate, narrowly ovate, 2-celled; perispore well
developed; 15-22 X 5.5-7.5 11..

Microconidia linear, 7-10 x 1 p.

Habitat on smooth bark.

It is somewhat unfortunate that such a well known name as A. fallax
SFNDLle be superseded but it probably is just as well since the name is
EUnCnig the most misused names. Arthopyrenia lapponina is characterized
1:y ‘the large ascocarps, slender paraphyses, narrowly elliptical asci
aFKi medium length microconidia. The species is common in Europe but is
kxlcwwn from North America in the typical form by only a single collection.
ITNEre is a small handful of collections which differ slightly from
£l~ lapponina but they have not been treated.

Illustrations: Figs. 53-59; Richardson & Morgan-Jones (1964),
fig. 1.

Exsiccati examined: Anzi Lang. 347 (FH-TUCK 4084); Claud. 50 (FH);
. II: 930 (US ex MO); Fellm. 221 (FH-TUCK 4072); Fr. 244 pr. p. (FH-
TIJCK.4067); Harm. 220 (FH); Hepp 450 (FH, FH-TUCK 4080), 451 (FH, FH-
TUCK 4066), 452 (FH), 453 pr. p. (FH); Krypt. Vind. 268a (FH,US), 1763a
(FH, US), 1763b (MSC); Malbr. 99 (Msc); Mass. 185, 186 (FH-TUCK 4071);
MOug. 364 pr. p (FH-TUCK 4069); Nyl. Par. 148 (FH-TUCK 4068); Rel. Suz.

6 (MSC); Schaer. 287 (FH-TUCK 4072); Stenh. 89 (FH-TUCK 4067, MICH, US).

58

Specimens seen: United States. NEW HAMPSHIRE: County unknown,

Shelburne Lake, Gentian Point, 1886 [Farlowl (FH).

Aurthopyrenia megalospora LOnnr.

Flora 41: 634. 1858. Lectotype collection: Fr. 244, Sweden.

Isolectotype in FH-TUCK 4067.

Ascocarp flattened, orbicular to elliptical in outline, 0.4-0.6 mm
ir1 diameter or length, sometimes less in width. Paraphyses slender but
{Yitflier irregular and someWhat gelatinized. Asci narrowly ovate, 85-100
>< 2&4-27‘u. Spores irregularly arranged, narrowly ovate, 2-celled;
FKEIIispore well developed; 25-32 X 6-8 u.

Microconidia rod-like, c. 5 X l u.

Habitat on smooth bark.

As Keissler (1936-38) has pointed out the two numbers of Fries'
exSiccatus cited by Lannroth were reversed in the original publication,
but it is clear that he meant var. okto apply to Fr. 244 and var./9to
apply to Fr. 242. The situation is further complicated since Fr. 244
143 a mixed collection of A, megalospora and A, lapponina. Apparently
tile collection examined by Keissler contained only A, lapponina and he
t1ierefore abandoned A, megalospora. However Since there is material
W1iich clearly fits LOnnroth's description, the name should be retained.
Fries 242 (var. fl) is A. fraxini.

Arthopyrenia megalospora is very similar to A, fraxini, differing

Only by larger asci and spores. It is possible that further study may
reveal that the two taxa are in fact extremes of the same species.

Material assigned to this species comes from Maine and Europe (Sweden

and England).

59

Illustrations: Figs. 60-64.
Exsiccati examined: Fr. 244 pr. p. (FH-TUCK 4067).
Specimens seen: United States. MAINE: Knox County, Warren,

Stunnit of Mt. Pleasant, 20.X.1912 Merri11 (FH).

lkrfthopyrenia minor R. C. Harris sp. nov.
Arthopyrenia sporis parvis, anguste ovatis, 12-15 x 4-4.5 u et
casu2is gracilibus, anguste ellipticis vel cylindraceis, 71-89 x 13-14 u.
Holotype: Florida, Liberty County, Apalachicola Nat. Forest,

1Nhi‘tehead Lake Campground, Harris 1545 (MSC). Isotype in H.

 

Ascocarps hemispherical to subglobose, superficial to semi-immersed,
O-25-0.5 mm in diameter; ascocarp wall thinner or lacking below. Para-
Ffllywses slender, regular and thread-like. Asci slender, narrowly ellipti-
Cxa]_ to cylindricla, ocular chamber usually distinct, (SO-)60-90 X
(S9-)12-14 u. Spores subbiseriate, narrowly ovate, 2-celled; perispore
thin; 12-17 x 4-5.5 p.

Microconidia rod-like, 4-7 X l u.

Habitat on bark.

Arthopyrenia minor is not a very distinctive species. It differs
flTom the majority of the American species by its smaller asci and
SEMores. It differs from A, degelii, which also has small spores, in

E5howing no signs of any constriction of the spore cells. Arthopyrenia

 

Eninor seems to be restricted to Florida, Louisiana and Texas.

Illustrations: Figs. 65-68.

Specimens seen: United States. FLORIDA: sine loc., Eckfeldt

 

(US); Liberty County, Apalachicola Nat. Forest, Whitehead Lake Camp-

ground, Harris 1498, 1545 (MSC). LOUISIANA: Lafayette/St. Martin

 

6O

Parish, Bayoue Tortue, 15.IV.1895 Langlois (US); St. Landry Parish,

Grand Coteau, 6.VI.1894 Langlois (US). TEXAS: sine loc., Hall (US).

lkrthopyrenia oblongens Zahlbr. EE R. C. Harris sp. nov.

 

Species distincta sporis ovatis, bilocularibus, cellulis insigniter
iluaequalibus, 13-15 X 7-8 p.

Holotype: Florida, Sanford, IV.1930 Rapp (MICH). Isotype in LD.

Ascocarps immersed, hymenium subglobose, 0.2-0.3 mm in diameter;
aimsocarp wall extended outward above forming an elongate shield,
c:- O.3-0.5 X 0.7-1.0 mm, thinner or lacking below. Paraphyses slender,
ITEgndlar and thread-like. Asci narrowly elliptical to cylindrical,
'75-110 X 14-16'p. Spores uniseriate to subbiseriate, ovate, 2-Ce11ed,
tile: cells markedly unequal with the lower narrower and shorter; peri-
SENDre not obvious; 13-15 x 7-8(-10) p.

Microconidia linear, 8-10 x 1 u.

Habitat on bark, both known collections on Andromeda (probably
== Lyonia).

The broad spores with unequal cells of this species would at first

Enggest the genus Anisomeridium, but the linear microconidia and absence

 

(If Trentepohlia lead me to place it in Arthopyrenia, especially Since

 

 

nmicroconidial shape seems to be a more stable character than spore
Shape.

Illustrations: Figs. 69-72.

Specimens seen: United States. Florida: Seminole County,

Sanford, 28.VI.1928 Rapp 709 (MICH), IV.1930 Rapp (MICH, LD).

61

Arth opyrenia padi Rabenh.

 

Lich. Eur., fasc. 14, 390. 1858. Type collection: "An Prunus
Padus bei Hermsdorf in Bilaer Grunde in der sachse Schweiz". Isotype

in MICH.

Ascocarps superficial, flattened to hemispherical, 0.1-0.3 mm in
diameter; ascocarp wall lacking below. Paraphyses thick, up to 2-4 p
in width, irregular, with short cells and often appearing rather paren-
ChYma-like, occasionally somewhat gelatinized. Asci ovate to narrowly
ovate, rarely narrowly elliptical, 2-celled, rarely 4-ce11ed in old
age; perispore thin; 15-20 X 4.5-5.5(-6.5) 11.

Microconidia short, rod-like, 3-4 X 1 1;.

Habitat on smooth bark.

In 1973 I treated this species under the name A. punctiformis

 

Massal. , mainly because I had no other name for it and the name had
b . . . .
een Commonly used in this sense. However, Since so much confu51on and

m‘ - . . . .
lslnterpretation surrounds A. pgnctiformis, Since I have not seen the

 

type and since A. padi is almost as old a name and is typifiable, I
have chosen to adopt A. padi. Further the European concept of

A. . . . .
‘ Ennctiformis 5. lat. (including A. analepta Auct. and A. atomaria

A
net. ) encompasses what I believe to be two distinct species, A. padi

a
nd 5.- salicis. Arthopyrenia padi differs from A. salicis in having

 

 

perSistent, or mostly so, paraphyses and in having generally longer and
hatrower spores. There are some specimens which are intermediate in
One or the other of these characters and their assignment to a species
is somewhat arbitrary. As a general rule I have used the gelatinization
Qf the paraphyses as the deciding character. Others may wish to do

cherwise.

62

I take a very broad view of A. Bali—iv since to do otherwise would
result in the recognition of numerous, barely distinguishable popula-
tions as microspecies. One of the more distinctive populations is the
one occurring on the West Coast. The asci and spores are somewhat

larger than normal for other regions and Trentepohlia is present in the

 

thallus. However I am not inclined to give this population any
taxonomic recognition.

Arthopyrenia padi is broadly distributed in temparate regions,

 

inCluding both Northern and Southern Hemispheres. In North America it
is the only species which is more or less uniformly distributed from
coast to coast in the northern states.

Illustrations: Figs. 73-79; Harris (1973), figs. 9-10; Richardson
& Morgan-Jones (1964), fig. 2; ve’zda (1968), fig. 8. Distribution:
fig. 408.

Exsiccati examined: Britz. 185 (US ex MO), 218 (MICH, US ex MO),
386 pr. p., 388 (US ex MO); Fr. 243 (FH—TUCK 4088); Kern. 2774 (US ex MO);
KIth. Vind. 468b (Us), 1523 (FH), 1763b (US); Lich. Colo. 67 (MICH, MSC);
M9119- 557 (MSC); Fis’ut 77 (MSC); Rab. 390 (MICH), 476 (FH-TUCK 4070);
Stenh. 180 (FH, FH-TUCK 4067, MICH, US).

Selected specimens seen: Canada. BRITISH COLUMBIA: Queen

ChaI‘lotte Islands, Graham Is., Brodo 11620, 11650, 12915 (CAN),

 

M‘u‘rChison Is., Brodo 11846 (CAN). ONTARIO: Carleton County, Ottawa,

 

30‘IV.1897 Macoun 462 (CAN, FH, US).

United States. ARIZONA: Cochise County, Barfoot Park, Weber &

W (MICH, MSC). CALIFORNIA: San Diego County, San Diego, 1883

w 28_ (US). CONNECTICUT: Middlesex County, Killingworth, 1876

% (FH). IOWA: Franklin County, Beeds Lake St. Park, Wetmore 17489,

 

 

63

174K96 (MIN). MAINE: Hancock County, Mt. Desert Is., Salisbury Cove,
2CVIII.1922 EAEEE_AQ§ (MICH, US). MASSACHUSETTS: Bristol County, New
‘BedfOrd, Willey (US); Essex County, Magnolia, IV.1896 [Farlow] (FH);
Ifliddlesex County, Newton, IV.1893 Farlow (FH). MICHIGAN: Alger County,

EI<of Kingston Lake, Harris 661 (MSC), Laughing Whitefish Point, Lowe

 

2£1§2_(FH, MICH); Charlevoix County, Beaver Is., Imshaug 27485 (MSC);
Ktnveenaw County, Isle Royale, Angleworm Lake, Mc Farlin & Brown 580a
(Ifli, MICH). MINNESOTA: County unknown, Minneapolis, 1896 Eégk A;
(DTICH). NEW YORK: Suffolk County, Gardiner's Is., Brodo 2375 (MSC).
SCXJTH DAKOTA: Custer County, 3 mi E of Sylvan Lake, Wetmore 7371 (MSC);
lemurence County, Terry Peak, Wetmore 9349 (CAN, MSC, WIS); Meade County,

11) mi E of Lead, Wetmore 10524 (MSC). WASHINGTON: Jefferson County,

 

almong Hoh River, TUCker 8311 (LSU); Pierce County, c. 15 mi W of

 

C:hinook Pass, Furlow 291 (MSC).

fgzghopyrenia planorbis (Ach.) Mull. Arg.

 

Mém. Soc. Phys. Geneve 30: 27. 1888. Verrucaria planorbis Ach.,
Slqlops. Lich. 92. 1814. Holotype: "in America ad Crotonis Cascarillae".
(H-ACH).

Arthopyrenia fallacior Mfill. Arg., Bot. Jahrb. Syst. 6: 404. 1885.

 

rEli/De collection: Cuba, Wright Verr. Cub. 129. Isotype in US.

Arthopyrenia planior Mfill. Arg., Bot. Jahrb. Syst. 6: 404. 1885.

 

SYUtype collection: Cuba, Wright Verr. Cub. 627b. Isosyntype in US.
Arthopyrenia planorbiculata Mfill. Arg., Bot. Jahrb. Syst. 6: 405.

1885. Type collection: Cuba, wright Verr. Cub. 64. Isotype in US.

Ascocarp semi-immersed to superficial, flattened to hemispherical,

64

hymnenium mostly pyriform to subglobose, c. 0.2 mm in diameter; ascocarp

vwall usually extended outwards above forming a broad shield up to 1 mm

5J1 diameter, lacking below. Paraphyses slender, regular and thread-like.

,Ascfi.narrow1y elliptical, 55-100 X 12-17(-20) u. Spores subbiseriate to

lxiseriate, narrowly ovate, 2-ce11ed, rarely 4-ce11ed; perispore thin;

17-23 x 5-711.

Microconidia filiform, usually curved, 20-27 x l u.

Habitat on bark.

Arthopyrenia planorbis is characterized by the broad shield formed
1:y the ascocarp wall and by the filiform microconidia. At first I was
iJiclined to recognize A, planorbiculata (including A, planior) as a
Skeparate species on the basis of smaller spores and longer microconidia,
knit the amount of overlap in both characters has finally led me to
Syrxxmmdze them. Arthopyrenia planorbis occurs in southernmost Florida
arnj the West Indies (Bahamas, Cuba, Dominican Republic, Grand Cayman,
Jamaica and Puerto Rico).

Illustrations: Figs. 80-87. Distribution: Fig. 409.

Specimens seen: United States. FLORIDA: Dade County, Coconut
(Stove, 1898 Thaxter 275 (FH), Everglades Nat. Park, SW of Pineland
Trail Area, Harris 2890 (MSC), Lemon City, Simpson's Hammock, 22.II.1917

fiafford 1753b (US); Monroe County, Big Pine Key, Hale 36816 (US), No

 

Name Key, Hale 36942 (US).

figfihopyrenia plumbaria (Stizenb. i2 Hasse) R. C. Harris comb. nov.
Verrucaria plumbaria Stizenb. i2 Hasse, Erythrea 3: 44. 1895.
EEgina_plumbaria (Stizenb. £2 Hasse) Hasse, Contr. U. S. Natl. Herb.

17: 12. 1913. Type collection not seen.

65

Pyrenula herrei Fink $2 Hedrick, Mycologia 25: 309. 1933. Holotype:

 

Ckilifornia, Santa Cruz Peninsula, Herre (MICH, Fink hb. 13,044).

Ascocarps superficial to slightly immersed, hemispherical, 0.3-
(3-4(-O.5) mm in diameter; ascocarp wall thinner or lacking below.
Paraphyses slender, regular and thread-like. Asci narrowly elliptical,
Ilarrowly ovate or narrowly obovate, 75-110 x 12-18 u. Spores uniseriate
'to biseriate, narrowly elliptical to narrowly ovate with one or both
sands pointed, 2-ce11ed, rarely 4-celled in old age, cells approximately
Eflqual in size, cells constricted near the middle or rarely two con-
strictions per cell; perispore well developed; (l3-)15-20 x 4.5-6.5 u.

Microconidia linear, 10-12(-15) x l u.

Habitat on smooth bark.

Arthopyrenia plumbaria is easily recognized by the spores with

 

CRonstricted spore cells. It is closely related to A, cinereopruinosa

 

thit differs in its more slender spores with equal cells and pointed
Enids and in longer and more slender asci. Misdeterminations of this

£Species account for the vast majority of the Arthopyrenia reports from

 

tflie West Coast. Arthopyrenia plumbaria ranges from California to

 

'British Columbia but never very far inland. It seems to be common.
Illustrations: Figs. 88-92. Distribution: Fig. 410.
Selected specimens seen: Canada. BRITISH COLUMBIA: Queen

Charlotte Islands, Graham Is., Brodo 10163 (CAN, MSC), Moresby Is.,

Egodo 10763, 12884 (CAN); Vancouver Island, Sidney, 19.VII.1914

Macoun (CAN, FH) .

United States. CALIFORNIA: Los Angeles County, Catalina Is.,

IX.1912 Hasse (ABSL, FH, US), Santa Monica Mtns., Sullivans Canyon,

66

V]I[.1913 Hasse (FH), Topanga Canyon, VII.1914 Hasse (FH); Napa County,
ZZIni W of Lake Berryessa, Tucker 6421-B (LSU); Santa Clara County,

.Santa Cruz Mtns., Los Gatos, 29.XI.1907 Herre 1175 (ABSL, FH, US),

 

Saxatoga, Toyon Lodge, 31.V.1945 Herre (MICH); Santa Cruz County, Santa
(lruz Peninsula, Herre (MICH). OREGON: Benton County, Adair Tract St.
Ifiorest, Pike 3041 (OSC), Corvallis, II.1931 Plitt (US). WASHINGTON:

,Iefferson County, along Hoh River, Tucker 8311 (LSU).

 

fgrthopyrenia rhyponta (Ach.) Massal.

Ricerch. Auton. Lich. 166, f. 329. 1852. Verrucaria rhyponta Ach.,
IQongl. Vetensk. Akad. Nya Handl. 30: 150. 1809. Type collection:
Ehueden, coll. unknown. Isotype? in BM-ACH 289.

Thallus brown to blackish, epiphloeodal, consisting of thick dark
brown hyphae.

Ascocarps more or less immersed in the thallus, hemisphereical,
0.2-0.3 mm in diameter. Paraphyses thick, irregular with short cells,
<then mostly gelatinized. Asci narrowly ovate, 40-75 x 15-20 u. Spores
lirregularly arranged, narrowly ovate, 4-celled; perispore thin;

17-20 X 5.5-7 1)..

Microconidia rod-like, 4-5 X l p.

Arthopyrenia rhyponta is distinguished by its dark brown thallus
and 4-ce11ed Spores. It seems somewhat related to 53.22913 It has
been reported from North America, but I have not been able to verify

any specimens of it and it should be deleted from the North American

flora.
Illustrations: Figs. 93-96.

Exsiccati examined: Anzi Lang. 471 (FH-TUCK 4088); Anzi Ven. 121

67

(Eii-TUCK 4088); Kern. 3136 (FH, US ex MO); Krypt. Vind. 1021 (FH);

bkmdg. 557 (FH-TUCK 4087).

_ߤihopyrenia salicis Massal.

Ricerch. Auton. Lich. 169, f. 336. 1852. Type collection not
seen.

Ascocarps superficial, hemispherical to flattened, 0.15-0.25(-0.3)
nun in diameter; ascocarp wall lacking below. Paraphyses more or less
menpletely gelatinized; gelatin occasionally IKI+ pale violet. Asci
<3vate to narrowly ovate, 35-50 X 15-20 u. Spores irregularly arranged,
ruarrowly ovate, mostly with rather blunt ends, 2-ce11ed; perispore thin;
(12-)15-22 X 4.5-6 u.

Microconidia short, narrowly elliptical, 2-3 X 1 u.

Habitat on smooth bark.

Arthopyrenia salicis is recognized by its gelatinized paraphyses,
SHnall, usually ovate, asci and rather small spores which are somewhat
llroader in proportion to length than those of the rather similar 53.229l'
Elle relation of these two species is discussed further under the latter.

fifigthopyrenia salicis is common in Europe but rare in North America and
11as not been previously reported.

Illustrations: Figs. 97-100.

Exsiccati examined: Erb. II: 120 (FH); Kern. 3534 (FH, US ex MO);
Kiypt. Vind. 468 (FH, US), 468c (FH); Leight. 288 (FH), 344 (FH, us);
Mass. 127 (FH-TUCK 4085), 201 (FH-TUCK 4077); Norrl. 149 (FH-TUCK 4068,
NBC); Rab. 203 (MICH), 629 (FH-TUCK 4070, MICH), 943 (MICH).

Specimens seen: United States. ALABAMA: Cleburne County, N of

Heflin, Harris 1348 (MSC). LOUISIANA: St. Helena Parish, NE of

68

Greungeville, Tucker 10165 (LSU). MASSACHUSETTS: Bristol County, New

 

Bedford, 1883 Willey (FH).

.Arthopyrenia taxodii R. C. Harris Sp. nov.

Arthopyrenia ascocarpis immersis clypeo lato, ascis cylindraceis,

 

105-130 x 15-20 u et microconidiis linearibus, 7-8 x 1 u.
Holotype: Texas, Houston, 1869 Ravenel 69_(FH-TUCK 4022).

IIsotypes in FH, MICH & US.

Ascocarps immersed, hymenium pyriform, 0.2-0.3 mm in diameter; tip
<Df ascocarp surrounded by a broad, thin black ring; ascocarp wall
Exoorly developed, lacking below. Paraphyses slender and regular. Asci
<2ylindrical, 100-130 X 14-20 u. Spores subbiseriate, often almost
Lnuiseriate, narrowly ovate to narrowly elliptical, 20-28(-32) x
6-8(-9.5) 11.

Microconidia linear, 7-8 x l u.

Habitat on soft bark of Taxodium.

‘Arthopyrenia taxodii is distinctive in its substrate preference,

Stince as a whole the genus Arthopyrenia occurs on thin smooth barks.

 

'The spores are reminiscent 0f.flr cinchonae, but the asci are more
Cylindrical and the ascocarp is immersed and the ostiole is surrounded
bY'a broad hyphal ring. The species is known only from Texas and
Florida.

Illustrations: Figs. 101-104.

Specimens seen: United States. FLORIDA: County unknown, Upsala,
S.XI.1932 Rgpp_(MICH); Duval County, Jacksonville, [Calkins?] (MICH).

TEXAS: Harris County, Houston, 1869 Ravenel 69_(FH, FH-TUCK 4022,

MICH, US).

69

2. ARTHOPYRENIA BIFERA GROUP

This group, although placed in Arthopyrenia, is kept separate since

 

thezspecies included seem to be more closely related to one another than
‘to the rest of the genus. Their 2-celled spores tend to have the Spore
\Nall slightly thickened forming a partial subdivision of the spore
lumen. Ultimately these thickenings form additional septa giving rise
'to 4-celled spores. The spore wall is usually strongly ornamented and
tile perispore is usually quite thick. There is also a tendency for
Inaduction in spore number and even in those which normally have 8 spores
fixer ascus occasional collections are found with four. Two of the species
Filaced here have a strong tendency for the ostiole to be eccentric. The
Inicroconidia are linear. Several of the species seem to be consistently
Eissociated with Trentepohlia. All of them are tropical or subtropical
1J1 distribution.

Many of the above characters are anomalous in Arthopyrenia and are

 

IYather more suggestive of the Trypetheliaceae. At the present time my
Llhderstanding of this group of species is not sufficiently great that I
VWish to take any major action in regard to them. It is possible that

tlhey might be included in Polymeridium or constitute a new genus

 

-related to it or, perhaps, a new genus of the Pleosporaceae. Hopefully
fUrther investigations of tropical pyrenolichens will bring to light
the connecting links needed to clarify the position of this group.

In addition to the species treated here, the group includes two
West Indian species, A, porospora Vain. from Dominica and an undescribed

Species from Jamaica with eccentric ostioles and two spores per ascus.

7O

Arijiopyrenia annulata R. C. Harris sp. nov.
Species distintissima sporis bilocularibus, cellulis intra
(annulo elevato, vel tarde 4-locularibus, asperis, 27-37 x 10-1211.
Holotype: Florida, Upper Matecumbe Key, 13.XII.1919 Britton &

Eygitton 846 (FH-RIDD).

Ascocarps immersed, hymenium flattened hemispherical, (0.2-)0.3-
(3.4 mm in diameter; tip of ascocarp surrounded by a broad, thin hyphal
.ring; ascocarp wall thin, lacking below. Paraphyses slender, regular
amid thread-like. Asci narrowly obovate, (80-)105-125 X 25-30 u. Spores
‘4-8/ascus, irregularly arranged, narrowly elliptical to narrowly ovate,
22-celled, each cell partially subdivided by a flat topped ring-like
tfliickening of the wall, becoming 4-ce11ed; spore wall strongly granular
<1rnamented; perispore thick, up to 2 M; 27-37 X 10-12 p (excluding
perispore) .

Microconidia linear, 6-8 X 1 p.

Habitat on bark.

This species is unique in the peculiar ring-like thickenings
Eiround the inside of the spore wall which eventually become septa
<iividing the spore into four cells. The ascocarp type and strongly
ornamented spores are also distinctive. In addition to the type col-
lection from Florida, the species is known from the Bahamas and the
West Indies.

Illustrations: Figs. 105-108.

Specimens seen: United States. FLORIDA: Monroe County, Upper
Matecumbe Key, l3.XII.l9l9 Britton & Britton 846 (FH-RIDD).

West Indies. BAHAMAS: New Providence, 9.XII.1918 Brace 9759

 

71

(FWi-RIDD). CUBA: Isle of Pines, Vivijagua, Britton & Wilson 15596 (FH-

 

RIIND). PUERTO RICO: Mayagfiez District, Yauco, 29.XII.1915 Fink 1482

(US).

.flgjhopyrenia bifera Zahlbr.

Ann. Mycol. 33: 34. 1935. Holotype: Florida, Sanford, 27.IV.1929

Rapp £39. [86 in orig.l (w).

Thallus poorly developed but Trentepohlia usually detectable.

Ascocarps mostly immersed, subglobose to hemispherical, 0.2-0.4 mm
1J1 diameter; ascocarp wall thinner or lacking below. Ostiole often
irather eccentric. Paraphyses slender, regular and thread-like. Asci
ruarrowly elliptical to elliptical, 75-105(-120) X (20-)25-35 u. Spores
z3/ascus, narrowly elliptical, 2-Celled, with a slightly thickened area
(bf the wall outlining a smaller subchamber, ultimately becoming 4-celled;
Spore wall strongly granular ornamented; perispore well developed;

137-48 x 15-16 u (excluding perispore).

Microconidia rod-like to linear, 5-10 x l u.

Habitat on bark.

The asci with only two spores, the peculiar thickenings of the
8pore wall and the strong ornamentation of the spore wall make A. bifera
One of the most distinctive pyrenolichens. It is common in Florida and
also known from Alabama, Louisiana and Texas.

Illustrations: Figs. 109-113. Distribution: Fig. 411.

Specimens seen: United States. ALABAMA: .§£23.122'1 Wilson (FH-
TUCK 4042); Baldwin County, Volanta, 21.II.1925 Eygggfli§§_(MICH).

FLORIDA: Dade County, Everglades Nat. Park, near Pa-hay-okee Overlook,

72

Harris 2744-8, 2749, 2751, 2757-8, 2826-G, 2829-8, 2916-8, 2947-8,

2968-8, 2971-B (MSC), N of Paurotis Pond, Harris 2672-8 (MSC), SW of

 

Pineland Trail Area, Harris 2875-C, 2882-B (MSC); Liberty County,
Apalachicola Nat. Forest, Porter Lake Picnic Area, Harris 1637 (MSC),
Whitehead Lake Campground, Harris l471-A, 1486, 1490, 1575-A, 1717
(MSC); Orange County, Camp Wewa, Schallert (F); Seminole County,
Sanford, 27°IV°1929.§EEEH§9 (W). LOUISIANA: St. Tammany Parish, Abita,

30.VII.1898 Langlois (US). TEXAS: sine loc., Calkins (US).

,Arthopyrenia lyrata R. C. Harris sp. nov.

Species distincta sporis bilocularibus cum uno vel ambis cellulis
in.medio parum constrictis, asperis, 21-29 X 8.5-10 u, microconidiis
liJiearibus, 6.5-9 X 1‘“ et algis ad Trentepohliam pertinentibus.

Holotype: Merr. II: 38, Florida, Daytona, 18.11.1911 Merrill

(PESC). Isotypes in MICH, US, US ex MO.

Ascocarps semi-immersed, less commonly immersed or superficial,
hemispherical, 0.3-0.5 mm in diameter; ascocarp wall usually lacking
bEilcmw. Paraphyses slender, regular and thread-like. Asci narrowly
eliljgotical to elliptical, usually with a distinct ocular chamber;
‘75-1420(-13o) x 20-30 p. Spores irregularly arranged to biseriate,
rk313rkowly elliptical or narrowly ovate, 2-celled, with one or both cells
“Kaeflily'constricted near the middle, rarely becoming 4-ce11ed; spore

' Weill granular ornamented; perispore well developed; 18-30 x 6.5-9.5(-12)
11- ( excluding perispore) .

Microconidia linear, 6-9 x l u.

Arthopyrenia lyrata has been previously confused with A, 222229222

lmdt can be separated from it and other similar species by the

73

ornamented spores, constricted spore cells and the presence of Trente-

Arthopyrenia lyrata is almost as uiquitous in the southeastern

pohlia.

United States as A, cinchonae and known from Alabama, Florida, Louisiana,

North Carolina, Texas and Mexico.

Illustrations: Figs. 114-118. Distribution: Fig. 412.

Exsiccati examined: Cum. 1: 180 (CAN, MSC, US, US ex MO); Cum. II:

110 (MICH, FH-RIDD); Merr. II: 38 (MICH, MSC, US, US ex M0).

Selected specimens seen: United States. ALABAMA: Baldwin County,

Fairhope, 26.V.l925 Evans 456 (MICH); Mobile County, Mobile, 7.X.1870

Mohr (US). FLORIDA: Dade County, Everglades Nat. Park, N of Paurotis

Pond, Harris 2663-A, 2686-C (MSC); Duval County, Jacksonville, Calkins
UWICH); Monroe County, Key Wesc, 11.1898 Thaxter 361 (MICH); Sarasota
CkNinty, Myakka River St. Park, Harris 2584-D (MSC); Seminole County,
Séuiford, XI.1910 BEER.§Z (FH); Volusia County, Daytona [= Daytona

Beach], 18.11.1911 Merrill (MICH, MSC, US, US ex MO), 1898 Thaxter 25

(PH), 337, 338 (MICH). LOUISIANA: E. Baton Rouge Parish, 1.5 mi S of

Baitnon Rouge, Tucker 8846-A (LSU); Iberia Parish, Avery Is., 11.1915

LICIllenny (FH-RIDD); Sabine parish, 3.5 mi N of Hornbeck, Tucker 7026

(LSU); St. Landry parish, St. Leo, 21.III.1895 Langlois (US); St. Martin
Parish, St. Martinville, 26.111.1894 Langlois (US); w. Feliciana parish,
Tunica Hills, Tucker 8417 (LSU). NORTH CAROLINA: Polk County, Tryon,
24-VI.1893 fl (CAN, FH-RIDD, MICH, MSC, US, US ex MO). TEXAS:
A*rElhsas/San Patricio County, Aransas Pass, 9.V.l913 Orcutt (ABSL, FH,
NtICHq); Brazos County, 5 mi E of Bryan, Darrow 4842 (US); Burnet County,

<3r1 St. Road 1604 4 mi W of old Blanco Rd., Fox T-44 (US); Cameron

C10Unty, 2 mi W of Boca Chica, F. Brodo 26H (CAN), NE of Brownsville,

Egdnyon 4144a (US ex MO); Kenedy County, 10 mi S of Armstrong, F. Brodo

74

EB (CAN); Nueces County, Corpus Christi, 1869 Ravenel §§_(US); San

Jacinto County, Little Thicket Sanctuary, 25.VIII.1967 Arnold (CAN).

3. MYCOGLAENA v. HOhn.
Sitzungsber. Kaiserl. Akad. Wiss. Math.-Naturwiss. C1., Abt. l. 188:

1210. 1909. Holotype: Verrucaria subcoerulescens Nyl.

?Arthopyreniella J. Stein. $2 Zahlbr., Ann. Naturhist. Hofmus.

Ist. Bot. Reale Univ. Reale

24: 284. 1911. EEBertossia Cif. & Tomas.,

Lab. Crittog. Pavia Atti, ser. 5, 10: 29, 56. 1953. Holotype:

Arthopyrenia cinersescens Massal. (see Harris, 1973).

Ascocarps flattened, superficial; ascocarp wall green or greenish

lilack, HN03+ reddish, lacking below. Hymenial gelatin IK1-. Paraphyses

CXEten rather thick, only slightly branched to branched and anastomosed.

Asx2i cylindrical, often with truncate apex, usually with a small ocular

Ckuamber. Spores colorless, 2-celled to muriform; perispore not evident.

Microconidia rod-like
Habitat on smooth bark, especially Myrica and Quercus.

For a discussion of the circumsription of this genus see Harris

(159'73). The color of the ascocarp wall and the usually truncate tipped

as“:j- are diagnostic for the genus. Detailed comments and specimen

CitEi‘tions (except for M, wetmorei) should be sought in Harris (1973).

1“ Spores 2-celled, 17-23 x 6-8 p ........................ M, wetmorei
1“ Spores 4-celled to submuriform ............................. 2

2. Spores with 4-6(-7) cells, two or more cells longi-

 

tudinally septate, 18-26 X (7-)8-10 p ......... M, meridionalis

 

75

2. Spores mostly with only 4-6 cells, rarely a single

cell longitudinally septate . ..................... ...... 3
3. Spores pointed at both ends, 4-ce11ed, one of the median
cells enlarged, 18-23 X 6-8 u; on Myrica, rarely on
A_b_i_€_3§ .. .............................................. .. M. myricae
3. Spores with more rounded ends, 4-6-celled, cells all
approximately equal, 18-30 X (5-)6-8(-9) u; on
Quercus ........... ........ ... ...... . ....... ......... M, quercicola

,Mycoglaena meridionalis (Zahlbr.) Szat. 12 Degen
Fl. Velebitica 3: 310. 1938. Polyblastiopsis meridionalis Zahlbr.,
Raul. Naturhist. Hofmus. 23: 223. 1909. Type collection: Krypt. Vind.
11546, "Hungaria: ad ramulos Fraxini orni in Valle "Skurinjatal" prope
FiJime, Schuler". Isotypes in FH & US.
Arthopyrenia peranomala Zahlbr., Ann. Mycol. 7: 472. 1909.
HOlotype: Albania, Boschoholz im Ragusaner Hafen zum Verhaus

ausEggeschiffts, l8.X.1907 Latzel 589C (W).

Spores ovate to elliptical, submuriform, 3-5(-6) transverse septa,
1‘5 cells with 1(-2) longitudinal septa, 18-26 x (7-)8-10 u.
Illustrations: Figs 119-125; Harris (1973), fig. 32.

iExsiccati examined: Krypt. Vind. 1646 (FH, US); Mass. 43 (FH-TUCK
4£84); Merr. II: 93 (MICH, MSC, US, US ex MO).

Additional specimens seen (not cited in Harris, 1973): United

Stiates. CONNECTICUT: Litchfield County, Ellsworth, 27.VIII.1884

% (US).

 

76

Mycoglaena myricae (Nyl.) R. C. Harris
12: 29. 1973. Verrucaria myricae Nyl., Flora 52: 297.

Michigan Bot.
France, Brest, 1868 Crouan (H-NYL 799).

1869. Holotype:
Verrucaria epidermidis var. aeruginella Nyl., Not. Séllsk. Fauna

Fl. Fenn. FOrh. 8: 173. 1866. Holotype: Finland, Kola, 1863 Fellman

Verrucaria fallacissima Nyl., Not. Sdllsk. Fauna Fl. Fenn. FOrh.

Finland, Kola, 1861 Fellman (H) nom. illeg.

8: 173. 1866. Holotype:

Spores narrowly ovate with rather pointed ends, 4-celled, one of

the median cells swollen, 18-23 x 6-8 p.
Therspecimen on which Nylander based Verrucaria fallacissima is one

Of M, myricae. However he included Verrucaria frankliniana Leight.
E== Polyblastiopsis fallaciosa (Arn.) Zahlbr.] in synonymy and thus
Unfortunately Vainio (1921)

.K} fallacissima is a superfluous name.

bEused his concept of Mycoglaena fallaciosa (Arn.) Vain. on this same
SPKECimen. This has led to some confusion in regards to the interpre-
No additional

tEItxion of Polyblastiopsis fallaciosa (Riedl, 1971).

American material has been seen.
Illustrations: Figs. 126-130; Harris (1973), fig. 33.
Exsiccati examined: Krypt. Vind. 861 (FH), 3477 (US); Magn. 127

(FH); Mig. 100 (MICH, MSC); Zw. 1197 (MICH).

£1lfiszégglaena guercicola R. C. Harris
1895

Michigan Bot. 12: 30. 1973.

Holotype: Iowa, Bremer County,

Eiiigi (US). Isotypes in CAN & US.

77

Spores narrowly elliptical to narrowly ovate, 4-6-celled, cells

approximately equal in size, occasionally with one cell longitudinally

septate, 18-30 x (5-)6-8(-9) u.

Illustrations: Figs. 131-134; Harris (1973), fig. 34.

Exsiccati examined: Cum. I: 249 (CAN, MICH, MSC, US, US ex MO, WIS);

II: 189 (ABSL, FH-RIDD, MICH, MSC, US).

Additional specimens seen (not cited in Harris, 1973): United

States. MISSOURI: Saline County, Emma, 19.VI.1900 Demetrio (MICH).

Mycoglaena wetmorei R. C. Harris sp. nov.

Mycoglaena sporis bilocularibus, 17-23 X 6-8 p.

 

Holotype: Texas, Brewster County, Big Bend Nat. Park, N slope of

Ehuory Peak, 6300 ft., Wetmore 19648-8 (MIN).

Thallus indicated by a lighter area on the bark, no algae seen.

Ascocarps flattened, orbicular, c. 0.3 mm in diameter, occasionally

depressed in the center. Paraphyses little branched, rather thick,

C‘ lB‘p, with scattered swollen cells, almost moniliform in places.

A5“3j- cylindrical or narrowly ovate, 70-90 X 12-15 u. Spores narrowly

vaatKE, 2-celled, very rarely 4-celled, upper cell broader than the lower;
perispore not obvious; 17-23 x 6-8 11.
Although this species is known only from the type collection, I

feiltl it should be described since it is the first 2-celled Mycoglaena

 

t(3 <Zome to my attention. Among the ten or so ascocarps examined a
Slngle 4-celled spore was seen.

Illustrations: Figs. 135-137.

78

4. TOMASELLIA Massal.
Flora 39: 283. 1856. EETomaselliomyces Cif. & Tomas., Ist. Bot. Reale
Univ. Reale Lab. Crittog. Pavia Atti, ser. 5, 10: 35, 62. 1953.
Holotype: Arthopyrenia arthonioides Massal.
Athrismidium Trev., Consp. Verruc. 15. 1860. Lectotype: Arthonia
gelatinosa Chev.

Beckhausia Hampe ex KOrb., Parerga Lich. 396. 1865. 32m, EEX§£°.lE
gyggg. Holotype: B, nitida Hampe g5 KBrb. (= Tomasellia gelatinosa).
Mycoporellum Mull. Arg., Rev. Mycol. (Toulouse) 6: 14. 1884.

Lectotype (Riedl, 1962): Mycoporum sparsellum Nyl.
Mycoporopsis Mfill. Arg., Flora 68: 514. 1885. Lectotype (Riedl,
1962): Mycoporum sorenocarpum Knight
Chlorodothis C1em., Genera of Fungi 50. 1909. Holotype:
fibgcoporellum lahmii Mull. Arg.
Nothostroma C1em., Genera of Fungi 50. 1909. Holotype:
Eggsgoporopsis roseola Mfill. Arg.
Sciodothis C1em., Genera of Fungi 50. 1909. Holotype:

Ehfisagporopsis leucoplaca Mfill. Arg.

Thallus indicated by discoloration of substrate, whitish, grayish
017 1Drownish, rarely dark brown, non-lichenized.

Ascocarps flattened, orbicular, elliptical or irregular in outline,
C10mpound with few to many chambers, each chamber with an ostiole;
Cheimbers separated by colorless or partially carbonized tissue. Hymen-
ifil gelatin IKI-. Paraphyses with short, rather irregular cells, often
a“ppearing rather parenchyma-like. Asci mostly narrowly ovate or ovate,

(*3Casionally narrowly elliptical or elliptical, ocular chamber mostly

79

not evident. Spores colorless when young, mostly becoming tinted

yellow brown in age, mostly irregularly arranged, narrowly ovate or

ovate, 2-4-celled, usually constricted at the septa; perispore usually

evident, thin to moderately thick; spore wall in most species becoming

strongly granular ornamented in old age.

Microconidia rod-like.

Habitat on bark, usually smooth young bark, rarely on ecorticate

wood or palmetto petioles.

Tomasellia was related by Massalongo (1856) to genera in the

 

Trypetheliaceae. This was reinforced by Trevisan (1861) who created a
SEparate section of the family for Tomasellia 5. lat. Zahlbruckner

(1926) and Keissler (1936-38) maintained this disposition of the genus.
However, as Miiller 8 von Arx (1962) and ve’zda (1968) pointed out,

lkgnasellia is closely related to Arthopyrenia and has no relationship to

 

13163 Trypetheliaceae but is a member of the Pleosporaceae (Mfiller & von

Alix, 1962). Further, study of Tomasellia in relation to Mycoporellum

arkj Aycoporopsis has led me to the conclusion that all three are con-

 

geniemfirn They all have the same type of ascocarps, paraphyses and asci.

wi?t}1 the exception of T. arthonioides and T, sparsella, all their

SpeCries known to me have spores which sooner or later become 4-celled

and-jpale yellow brown with an ornamented spore wall. Tomasellia, in my

nuirkd, is separated from such species of Arthopyrenia as A, padi

fi‘ irhyponta and the A, persoonii group only by having more than one

Chamber in the ascocarp. Riedl (1962) separates Mycoporellum and

ESIFZOEoropsis from Arthopyrenia by whether the spores become colored or

not. I have seen occasional colored spores in most of the common species

(3f Arthopyrenia. I personally find that if one wishes to recognize

80

separate genera, it is much simpler and more useful to do so on the basis

of ascocarp structure rather than on the color of the spores in old age.

Thus Tomasellia is somewhat artificial but, since the group is homo-

 

geneous and its recognition reduces the unwieldy size of Arthopyrenia,

I feel justified in recognizing the genus.
The maintenance of separate genera for 2-ce11ed and 4-celled

Spored species also seems unwarranted as I have found a few 4-celled

Spores in most of the species previously considered 2-celled. The only

 

species which consistently have only 2-celled spores are T. arthonioides

Thus if one wishes to recognize separate genera on

and T. sparsella.

the basis of spore septation, Tomasellia would be the correct name for
Species with 2-celled spores and Athrismidium for those becoming
‘Q-celled (Athrismidium had two original Species but one was included
knith a question mark so that Arthonia gelatinosa is made the lectotype).

1: feel that such a distinction is unecessary and prefer to recognize

Offily a single genus.
There is a rather interesting problem relating to the recognition

(Df: species within Tomasellia. My species concepts are broader than those

01? Iiiedl (1962, 1963) and are based primarily on spore characters. In

SCNTNE species 4-celled spores develop very early and the majority of the

SEDCHTes are 4-celled. In others 4-celled spores develop very late and

tilefll only in small numbers. For all practical purposes T. gelatinosa

differs from T. lactea and '_I'_. californica from _T_. eschweileri only

lr1 the time at which the Spores become 4-celled. I cannot help wonder-

11mg if this might not be under simple genetic control. If this were
'ttlmal think one might be justified in treating them as forms of a

Single species. However since there is no direct evidence to hand,

81

I provisionally maintain them as separate species.

Taken as a whole Tomasellia seems to be highly oceanic in distribu-

 

tion with only a few collections of a few species occurring in con-

tinental areas.

Spores very soon 4-celled . ................ . ......... ....... 2
Spores 2-3- or tardily 4-celled .. .................... ...... 4
2. Spores 15.17 X 4-505 11. 0.0.0....0 ..... 0 0000000000 . E. americana

2. Spores 17 u or more in length and 5.5 u or more in

 

 

Width 00...... ......... .... OOOOOOOOOOOOOOOOOOOOOOOO ...... 3
Spores 17-22 x 5.5-7 u ...... ................. ...... T. californica
Spores 20-28 x 7-8 p ............................ .... g, gelatinosa
4. Spores constantly 3-ce11ed .................... ..._T. macularis

4. Spores mostly 2-celled, often tardily 4-celled ......... 5

Spores small, constantly 2-celled, 12-15 X 4.5-5 u .....

. .......... . ................................... .. LT. arthonioides]

 

Spores larger, over 17 u in length ....... ....... ........... 6

6. Spores 1.5-2.5 times longer than broad, l7-22(-25)
X705-10u 0.... ..... 0 0000000000000000 0.0.0.0.... 2. SEarsella

6. (Spores 2.5-4 times longer than broad, 17-27 x 5-8 p .... 7

Spores 20-27 X 605-8 1.1. .000.0.0.0.........0...00.00.0...0 2:. lactea.

Spores l7-20(-22) x 5-6(-7) u ...................... T. eschweileri

 

82

Tomasellia americana (Minks EA Willey) R. C. Harris comb. nov.

Cyrtidula americana Minks §§_Willey, Enum. Lich. New Bedford 33.

1892. Syntype collection: Massachusetts, New Bedford, Willey.

Iscsyntype in MICH.
Ascocarps orbicular to elliptical, 0.25-0.7 mm in diameter or

lenigth, occasionally somewhat less in width, 0.05-0.l mmin height, with
:fevv to many locules. Asci ovate to narrowly ovate or narrowly elliptic-

al., 35-60 x 14-18 u. Spores irregularly arranged, narrowly ovate,

mcnsi:ly 4-celled; perispore thin; 15-17 x 4-5.5 p.
Prunus and Vitex.

Habitat on smooth bark of Cornus, Hamamelis,
Tomasellia americana has the smallest spores of any of the species

VW11:}1 4-ce11ed spores. It is known from only a few collections from

Massachusetts and one from Yugoslavia.

Illustrations: Figs. 138-140.

vezda Sel. 701 (05).

Exsiccati examined:
MASSACHUSETTS:

United States.

Bristol County,

Specimens seen:
New Bedford, Willey (MICH), Willey 742 (FH-TUCK 3734, 3735), Willey 857

(PH-TUCK 3731).

Ezifléfliellia arthonioides (Massal.) Massal.
Arthopyrenia arthonioides Massal., Ricerch.

Flora 39: 283. 1856.

Aultan. Lich. 169, f. 339. 1852. Type collection not seen.
0.9-2.0 mm in diameter, 0.05-0.1 mm in height,

Ascocarps orbicular,
Asci narrowly ovate to narrowly elliptical,

mos‘Ltly with many locules.
Spores irregularly arranged to subbiseriate, narrowly

4‘5~70 x 13-15 N-
C“Vette, 2—celled, lower cell narrower and longer; perispore not

1'
\
..
0
c
-
o .
‘0 .-
.. ...»
u
a-..
. -

83

evident; 12-15 X 4.5-5 0.

Microconidia rod-like, 3-5 X 1 u.

Habitat on smooth bark, especially young Fraxinus.

Tomasellia arthonioides is included since it is the type species
<of the genus. It has not yet been found in North America.

Illustrations: Figs. l4l-144; vezda (1968), fig. 10.

Ttxncisellia californica (Zahlbr.) R. C. Harris comb. nov.
Mycoporellum californicum Zahlbr., Ann. Mycol. 10: 363. 1912.
Myxchporopsis californica (Zahlbr.) Riedl, Sydowia 16: 216. 1963.

TYIDGE collection: California, Santa Monica Mtns., Hasse 1003. Isotype

 

irl IFH.

Ascocarps orbicular, elliptical or irregular in outline, 0.3-0.7 mm
irl ciiameter or length, often c. 0.1 mm less in width, 0.05-0.1 mm in
hEILQflQt, with few to many locules. Asci mostly Ovate or elliptical, less
Cornmonly narrowly ovate or narrowly elliptical, 45-60 x (15-)20-27 u.
SFKDITES irregularly arranged, narrowly ovate, mostly 4-celled; perispore
tffirls 17-22 x 5.5-7,1.

,Habitat on smooth bark.
_Tpmasellia californica has about the same spore size as

‘- £E§ghweileri but the latter has predominantly 2-celled spores.

W“ .
JItll the exception of a single collection from Texas, T. californica

 

i . . ..
£3 ICnown only from California and Hawaii.

Illustrations: Figs. 145-149; Richardson & Morgan-Jones (1964),

fig. 7

Exsiccati examined: Merr. I: 156 (FH, US).

4..
n-

0 H.
~< \—
'----_
~
‘- --

., ~‘

 

0“-

s
.

‘I
x
Q
.3 -
\“
‘-
Q.‘ .
‘u

-,
I

H.
1

.
“‘-

 

 

I

4
1"
/

84

Specimens seen: United States. CALIFORNIA: Los Angeles County,
Catalina Island, IV.1915 Hgggg (FH), Santa Monica Canyon, V.1914 EEEEE
(FH), VII.1914 Hgggg (FH), Santa Monica Mtns., 1906 Hgggg (FH), 1913
Idasse (FH), 111°1915.§§§§§ (FH, US). HAWAII: Oahu, Punchbowl,

25.]151895 Hellgg (FH), 1.1V.1895 Heller 2051 (FH, MSC). TEXAS: Dallas

 

County, Dallas, 1879 [Hall] (FH-TUCK 3734).

Ttxncisellia eschweileri (Mfill. Arg.) R. C. Harris comb. nov.

Mycoporellum eschweileri Mfill. Arg., Flora 71: 526. 1888.

 

HcfiLCDtype: Brazil, Bahia (G).

Ascocarps orbicular to irregular, 0.3-0.6(-0.8) mm in diameter or
lenuggth, often somewhat less in width, 0.06-0.12 mm in height , with few
tC> niany locules. Asci narrole ovate, 42-65 X 20-23(-26) u. Spores
irregularly arranged, narrowly ovate, mostly 2-celled, a few becoming
4‘CZ‘Eflled; perispore thin; 17-20 X 5-6(-7) u.

Iflicroconidia rod-like, 4-5 x l u.

liabitat on bark and rarely on palmetto petioles.

TThe spore size of T. eschweileri overlaps that of T. lactea and

 

Occasional collections are difficult to place. It differs from

IS

° .E§3$;£Q£A122_in spore septation. The Iowa and Minnesota collections
d' . .
l:E:E-:"‘-“r in that the spore wall is very thin and the spores tend to

c
Ollapse, but I am not inclined to recognize them as a separate taxon

01‘) U). . . . .
is baSis. They were included in Harris (1973) as Mycoporellum sp.

 

T
‘££E§E§gllia eschweileri has been collected in the southeastern United
S
tiitkas, the Hawaiian Islands, Brazil and the West Indies (Cuba, Grand
a‘yman, Haiti, Jamaica, Puerto Rico, Martinique and St. Lucia). It

t1
Ens not been reported from North America preViOUSlY-

85

Illustrations: Figs. 150-155. Distribution: Fig. 413.

Exsiccati examined: Merr. I: 156 (CAN, FH-RIDD, US).

Specimens seen: United States. ALABAMA: Baldwin County, Fish
River, 21.111.1925 §y§A§_§2§_(MICH). FLORIDA: Dade County, Everglades

phat. Park, near Pa-hay-okee Overlook, Harris 2730-8, 2815-C, 2840-8

 

(DESC), near Pineland Trail Area, Harris 2888-8, 2889-A, 29ll-F (MSC);
.Dtnfal County, Jacksonville, Calkins (FH, US); Flagler County, Flagler
St;. Park, Harris 2500-8, 2502-8 (MSC); Sarasota County, Myakka River
StL. Park, Harris 2663-C (MSC); Seminole County, Sanford, 5.VI.1915

RaIEEZ 214 (FH); Volusia County, Tomoka St. Park, Harris 2319 (MSC).

 

ILAVVIKII: Oahu, Punchbowl, 25.111.1895 Heller (FH), l.IV.1895 Heller
(CAN, FH-RIDD, US, US ex MO). IOWA: Bremer County, VIII.1895 Fink
(NEICIH). KENTUCKY: Rockcastle County, Conway, 8.1V.1927 Fink (MICH).

LOUISIANA: St. Martin parish, 1889 Langlois 957 (US). MlNNESOTA:

 

811163 Earth County, Mankato, 24.VI.1899 Fink 29_(MIN). TEXAS: Cameron

County, 2 mi N of Olmito, Runyon 3814 (US ex M0).

 

gE§E§E§§llia gelatinosa (Chev.) Zahlbr.

(Sat. Lich. Univ. 1: 474. 1922. Arthonia gelatinosa Chev., J. Phys.

 

Chint- Hist. Nat. Arts 94: 54. 1822. Type collection not seen.

Ascocarps orbicular, 0.4-1.0 mm in diameter, 0.07-0.15 mm in
heiiSflfit, mostly with numerous locules. Asci ovate to almost elliptical,
SCLCE35 x 24—30 u. Spores irregularly arranged, narrowly ovate, mostly
4‘Qelled; perispore thin; 20-28 X 7-8 1)..

Habitat on smooth bark.

Tomasellia gelatinosa differs from T. lactea in having predominantly

 

. .—
...,
u- -.

5v~..

 

 

86

4-Celled spores and from T. californica by larger spores, although
there is some overlap in spore size. Tomasellia gelatinosa has not
previously been reported from North America.
Illustrations: Figs. 156-159.
Exsiccati examined: Magn. 377 (US); Malbr. 250 (MSC); Rab. 780
(PTi-TUCK 3734); Trev. 54 (FH).
Specimens seen: Canada. NEWFOUNDLAND: Bay of Islands, Lark
Pharioour, 19.111.1887 Waghorne 576 (US ex MO).
United States. MAINE: Knox County, Rockland, 15.1V.1910 Merri11

CFPI).

EggnEisellia lactea (Ach.) R. C. Harris comb. nov.

Verrucaria stigmatella var. lactea Ach., Lich. Univ. 277. 1810.
.Xfiflflfucaria lactea (Ach.) Eschw. ig_Martius, F1. Brasil. 1: 126. 1833.
EQZEEEflggrellum lacteum (Ach.) Zahlbr., Cat. Lich. Univ. 1: 554. 1922.
TYTDGE collection: "India Occid., Swartz". Isotypes in BM-ACH & UPS.

NWCoporopsis leucoplaca Mull. Arg., Flora 69: 316. 1886.
y§§2£flggrellum leucoplacum (Mfill. Arg.) Zahlbr., Nat. Pflanzenfam.
1(1‘1): 78. 1903. Type collection: Brazil, Minas Geraes, Glaziou.
ISOtypes in FH, FH-TAYL 118 8 US.

jMycoporellum hassei Zahlbr. i2 Hasse, Bryologist 15: 46. 1912.
Holetype: California, Catalina Island, near Avalon, Hasse 1321 (W).
ISOtypes in FH.

flycoporellum deserticola Fink £2 Hedrick, Mycologia 22: 248. 1930.
EEXEEE‘ELoropsis deserticola (Fink $2 Hedr.) Riedl, Sydowia 16: 218. 1963.

1101-<3type: Puerto Rico, Yauco, 1.1.1916 Fink 1688 (MICH).

87

Ascocarps orbicular, elliptical or irregular, 0.3-0.6 mm in diameter
or length, Often less in width, 0.09-0.13 mm in height, with few to many
locules. Asci ovate to elliptical or narrowly ovate to narrowly ellipti-
cal, 50-75(-100) X 20-27(-30) u. Spores irregularly arranged, narrowly
(ovate, mostly 2-celled but a few 4-Ce11ed; perispore thin to moderately
tfliick; 20-27 x 6.5-8 u.

Microconidia rod-like, 4-5 x l u.

Habitat on bark and wood.

Riedl (1962) excluded this species since the specimen in Helsinki

MESS; apparently a poorly developed Arthopyrenia. However other isotypes
ccarrtain well developed material of this species. Mycoporellum difforme
(de;rMS 12_Willey) Fink was described from Massachusetts, but I have not
hxaeerlable to examine the original material. I presume from Riedl's
(119652) description that it will prove to be identical with T. lactea.
“3163 original material of Mycoporopsis leucoplaca has asci larger than
SSEEHH normal for T. lactea but agrees in its other characters, so I have
tentatively placed it in synonymy.

Tomasellia lactea is the most widely distributed species in the

 

geenus. In North America it occurs from Massachusetts to Texas and in
Cal"ifornia. It is also known to me from Brazil, Puerto Rico and in
BumDpe from England, Ireland and Tenerife.

Illustration: Figs. 160-166. Distribution: Fig. 414.

Exsiccati examined: Rel. Hasse 113 (FH).

Specimens seen: United States. ALABAMA: Baldwin County, Bailey's
Beach, 8.111.1925 mg (MICH). CALIFORNIA: Los Angeles County,
Catalina Island, V.l9ll 331533 (FH), 1912 flags}: (us), beach 5 of Avalon,

‘V‘Zl9ll Hasse (FH, W), near "Clifton by the Sea," XII.1907 Hasse (FH),

.-.

u....

'II

II‘

~—

 

\
\
~

88

Santa Monica Mtns., Hasse (MICH); San Diego County, San Diego, 111.1885
Farlow (FH, MICH). FLORIDA: Dade County, Everglades Nat. Park, between

Mahagony Hammock and Pa-hay-okee Overlook, Harris 2916-A (MSC), near

 

Pa-hay-okee Overlook, Harris 2744-A, 2757-D, 2762, 2813-B, 2829-F (MSC),

ruear Pineland Trail Area, Harris 2875-B, 2896-B, 2898-A (MSC); Duval
Chaunty, Ft. George Island, Calkins (MSC), Jacksonville, Calkins (FH);
Fflqagler County, Flagler St. Park, Harris 2478-A, 2486-A (MSC); Liberty
(:cnlnty, Apalachicola Nat. Forest, Whitehead Lake campground, Harris 1448,
1488 (MSC); Seminole County, Sanford, 27.IV.1914 my (FH), 5.V.19l4
RaEE 1_3 (FH), v.19l9 EBB (US). MASSACHUSETTS: Bristol County, New

Bedford, 1881 Willey 850d (FH-TUCK 3731). SOUTH CAROLINA: Berkeley

 

<3cnqunty, Francis Marion Nat. Forest, Guilliard Lake campground, Harris

3166 (MSC);Charleston County, near Charleston, Ravenel 347 (PH-TUCK 4050).

.EEEEEggellia macularis (Minks ex Willey) R. C. Harris comb. nov.
Cyrtidula macularis Minks ex Willey, Enum. Lich. New Bedford 33.
ley922. Syntype collection: Massachusetts, New Bedford, Willey. Iso—

syntYpe in MICH. .

Ascocarps orbicular to elongated and irregular, 0.4-0.6 mm in
diameter to 1.2 mm long and 0.4 mm wide, 0.07-0.12 m in height, with
few"to many locules. Asci ovate to elliptical, 45-65 x 27-33 p. Spores
irreEgularly arranged, narrowly ovate to ovate, 3-ce11ed, upper cell the
lair‘gest; perispore moderately thick, 22-26 x 8-12 u.

Habitat on smooth bark of Hamamelis, Pinus and Egugus.

Tomasellia macularis is quite unusual in that the spores are 3-

 

‘:eiLled. A few 3-celled spores may be found in other species but they

ax“? transitional to 4-celled spores. No 4-celled spores have been found

 

 

 

7v
.7.
'\
s.
~-
s"
~
a

 

.x~‘

 

89
in T. macularis.
Illustration: Figs. 167-169.
Specimens seen: United States. MASSACHUSETTS: Bristol County,

IVew Bedford, 1881 Willey 850a, 850C (FH-TUCK 3731), 1885 Willey 972

 

 

(FH-TUCK 3735).

Itxnasellia sparsella (Nyl.) R. C. Harris comb. nov.

 

Mycoporum sparsellum Nyl., Ann. Sci. Nat. Bot. ser. 5, 7: 343. 1867.

 

Eggcnoporellum sparsellum (Nyl.) Mull. Arg., Rev. Mycol. (Toulouse) 4: l4.

li3Eh4. Lectotype: Columbia, Monte del Morro, Lindig (H-NYL 4292).

Ascocarps orbicular, elliptical or irregular in outline, 0.4-0.6 mm
irl (fliameter or length, often somewhat less in width, 0.08-0.2 mm in
liexigght, with relatively few locules (up to c. 15). Asci ovate or ellip-
tixzeal, 55-85 x 24-35 u. Spores irregularly arranged, ovate, 2-celled;
Perispore thin; 17-22(-25) x 7.5-10 u.

,Microconidia rod-like, 4-5 x 111.

Tomasellia sparsella is distinguished by its relatively short but

 

brVDEui spores which are ovate (length/width ratio 2-2.5:l) rather than
n913170le ovate (length/width ratio 2.5-4:l) as in the rest of the species
trweérted. It is known only from Florida in North America but also has
ben3r1 found in Columbia, Guatemala and the West Indes (Cuba, Dominica,
Grand Cayman and Jamaica).

Illustration: Figs. 170-173. Distribution: Fig. 415.

Specimens seen: United States. FLORIDA: Dade County, Everglades
Nat.

Park, between Mahagony Hammock and Pa-hay-okee Overlook, Harris

E§2§§i_(MSC), near Pa-hay-okee Overlook, Harris 2799, 2815-A, 2820 (MSC),

 

heEerineland Trail Area, Harris 2911-6 (MSC); Flagler County, Flagler

 

 

.0.

(I!

-.

 

 

I

II

90

St. Park, Harris 2484-8 (MSC); Marion County, Ocala Nat. Forest, 5 of

 

Long Pond, Harris 1819 (MSC); Seminole County, Sanford, Rapp (MICH),

 

1.1912 RaER (FH, US), 25.V.1914 Rapp (FH).

C. STRIGULACEAE Fr.
Syst. Orb. Veg. 1: 110. 1825. Holotype: Strigula Fr.
Xanthopyreniaceae Zahlbr., Nat. Pflanzenfam., ed. 2, 8: 91. 1926.

Holotype: Xanthopyrenia Bachm.

 

Monoblastiaceae W. Wats., New Phytol. 28: 106. 1929. Holotype:

Monoblastia Ridd.

 

Pleurotremataceae W. Wats., New Phytol. 28: 112. 1929. Holotype:

Pleurotrema Mfill. Arg.

 

Acrocordiaceae Oxn., Flora Lishainikiv Ukraini 1: 144. 1956.

nom. inval. descr. ross.

 

Thallus usually well developed, mostly within the substrate but
occasionally mostly superficial, subcuticular in the foliicolous species.

Most species associated with Trentepohlia, foliicolous species with

 

Cephaleuros, species of Pyrenocollema with various blue—green algae.

 

 

Ascocarp perithecioid lacking an ostiolar apparatus in the two
species studied by Janex-Favre (1971), unknown for the rest but presum-
ably similar. Ascocarps not fused by their necks except in Strigula
connivens. Ostiole erect in all except Pleurotrema, Monoblastia
palmicola and Anisomeridium mécgosporum. Interascal filaments in the
species studied by Janex-Favre (1971) were found to be true paraphyses.
Paraphyses slender, thread-like, branched and anastomosed in most, only
sparsely so in Strigula. Asci bitunicate (Richardson & Morgan-Jones,

1964; Swinscow, 1965b, 1967), nassascé (Magne, 1946), bitunicate-

91

rmssascé (Janex-Favre, 1971). Spores colorless, transversely septate
(submuriform in Strigula submuriformis, simple in Monoblastia palmicola),
eight per ascus (except for random abortion), occasionally dividing into
two part spores before or after release from the ascus, mostly without a
well developed perispore, with granular ornamentation in Acrocordia,

commonly in Pleurotrema, rarely in Anisomeridium.

 

Microconidia orbicular, elliptical or oblong.

Macroconidia known for most species of Strigula and some of Anigg-
meridium, cylindrical or fusiform and transversely septate in Strigula,
orbicular, elliptical or rarely cylindrical and non-septate in

Anisomeridium.

 

Lichen substances generally lacking except for a few species of
flflisomeridium which produce lichexanthone and one or more anthraquinones
in one species.

Habitats include bark, wood, leaves, bryophytes, rock and soil,

occasionally aquatic.

The six genera of the Strigulaceae contain the bulk of the species

removed from Arthopyrenia, some wrongly placed in Porina and Thelidium,

 

and even one from the Collemataceae. The genera are united on the basis
Of an aggregation of characters rather than any one outstanding feature.
These include similar ascocarp type and development (as far as is known),
generally similar asci and paraphyses and similar type of spores and
microconidia. It is of interest to note that the spores show relatively
little variation in comparison to the other families of pyrenolichens.
NO members have yet been found with truely muriform spores, acicular
'Spores or an increased or decreased number of spores in the ascus.

There is also little variation in the form of the ascocarp. Parathelioid

A“

'.’

_-

92

and astrothelioid types are rare and no instances of aggregation within

a pseudostroma are known, while these types are common in the Pyrenula-
ceae and Trypetheliaceae. Although the Strigulaceae may be lacking in
morphological versatility it is ecologically very versatile. The only
other pyrenolichen family capable of living in such a variety of habitats

is the Trichotheliaceae. The Pyrenulaceae (except Lithothelium) and

 

Trypetheliaceae are exclusively corticolous. Although a number of

species are tropical in distribution the Strigulaceae seem less exclu-
sively tropical than the Pyrenulaceae, Trichotheliaceae and Trypetheli-
aceae. The Strigulaceae do not seem to me to be closely related to any

of the other pyrenolichen families.

1. ACROCORDIA Massal.

Geneac. Lich. 17. 1854. Lectotype (Th. Fr., 1861): Lichen gemmatus Ach.

 

Acrocordiomyces Cif. & Tomas., Ist. Bot. Reale Univ. Reale Lab.

 

Crittog. Pavia Atti, ser. 5, 10: 26, 56. 1953. Holotype: Verrucaria

conoidea Fr.

Thallus well developed, mostly endophloeodal or endolithic.

Phycobiont Trentepohlia.

 

Ascocarp with outer layers of wall brown to brown black, occasion-
ally almost colorless in deeply embedded ascocarps. Ostiole erect or
sometimes eccentric. Paraphyses slender, rather loosely branched and
anastomosed forming a rather regular network. Asci cylindrical with a
hemispherical to almost subglobose ocular chamber which is capped by a
layer which is clearly distinct when stained (meniscus of Janex-Favre,

1971). Spores eight in the ascus, uniseriate, oblong to elliptical;

93

spore wall with granular ornamentation on the epispore (apparently
included in the perispore), perispore and granules disappearing in KOH.
Microconidia elliptical to oblong.
Habitat on bark, old wood and rock.

Acrocordia is a genus easily recognized by its distinctive ascus

 

and spore type. At present I would include only seven species, A, cavata,
A, conoidea, A, gemmata (Ach.) Massal., A, megalospora, A, salweyi
(leight.) A. L. Sm., Arthopyrenia subglobosa V5zda and possibly
Acrocordia macrospora Massal.

The distribution of the species is of some interest. Acrocordia
gemmata, macrospora, salweyi and subglobosa are strictly European.

Acrocordia gemmata has already been excluded from the North American

 

flora and all of the previous records of A, conoidea examined have

turned out to be either Verrucaria spp. or Thelidium spp. Thus A, con-

 

oidea is European with the exception of one collection from Chile
(Juan Fernandez Is.) and another in North America (Michigan). Acrocor-

dia megalospora is restricted to eastern North America. Therefore the

 

only widespread species is A, cavata, occurring in both Europe and
North America. No specimens of this genus have been verified from the
tropics nor, with a single exception, from the Southern Hemisphere.

Acrocordia is rather unique among the pyrenolichens in having an almost

 

entirely North Temperate distribution.

1. Growing on calcareous rock; spores 13-17 x 7-8.5 u .... A, conoidea

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

 

u.‘

 

I)

 

(In!

94

2. Spores 33-48(-60) x 15-23 u ... ............. ... A, megalospora

2. Spores 11-17 x 6-9.5 u ............................. A, cavata

Acrocordia cavata (Ach.) R. C. Harris l2_V5zda

 

Lich. Sel. 1229. 1974. Verrucaria cavata Ach., Syn. Lich. 91. 1814.

 

Arthopyrenia cavata (Ach.) R. C. Harris, Michigan Bot. 12: 11. 1973.

 

Holotype: Switzerland, [Schleieher21 442a (H-ACH).

Ascocarps 0.3-0.5 mm in diameter. Asci 80-115 x 9-10 p. Spores
11-17 x 6-9.5 u; spore wall with minute granular ornamentation.

Microconidia oblong, 3-6 x 1-2 u.

The relationship Of.£° cavata to other species in the genus is
discussed in Harris (1973). European specimens have been verified from
England, France, Germany, Sweden and Switzerland. The range in eastern
North America has been extended significantly by a collection of this
species from South Carolina.

Illustrations: Figs. l74-l78; Harris (1973), fig. 4.
Distribution: Fig. 416.

Exsiccati examined: Fr. 274 (MSC); Krypt. Vind. 2152 (FH, US);
Malbr. 398 (MSC); Rab. 89 (FH-TUCK 4062); vezda Sel. 1229.

Additional specimens seen (not cited in Harris, 1973): United
States. MICHIGAN: Baraga County, NE of Sidnaw, Harris 7827, 7878 (MSC);
Keweenaw County, Isle Royale, Conglomerate Bay, Wetmore 4214 (MSC) [sub

Bacidia rubella]. SOUTH CAROLINA: Berkeley County, Francis Marion Nat.

 

Forest, Guilliard Lake Campground, Harris 3193-B (MSC).

95

Acrocordia conoidea (Fr.) KOrb.

 

Syst. Lich. Germ. 358. 1855. Verrucaria conoidea Fr., Lichenogr.

 

Europ. Reform. 432. 1831. Type collection not seen.

Ascocarps 0.6-0.8 mm in diameter. Asci 120-140 x 9-11 u. Spores
13-17 x 7-8.5 u; spore wall with minute granular ornamentation. (In
part from VEZda, 1961).

Microconidia oblong to elliptical, 3-4 x 1.5-2 u.

Habitat on calcareous rock.

The spore measurements of the sole North American collection compare
very closely with those given by VEzda (1961) for var. conoidea.

However the range he gives seems somewhat low based on the comparatively
few specimens 1 have studied (15). The ascocarps are also slightly
smaller and less spreading at the base than in European material, but
the wall is not as incurved at the base as in var. EEEQE‘VEZda.

Acrocordia conoidea is currently known in North America only from

 

Michigan. All other North American collections under this name seen by
me have proved to belong to either Thelidium or Verrucaria. The Michigan
collection grew on a limestone boulder at the edge of Lake Huron,
certainly wetted by waves in rough weather.

Illustrations: Figs. 179-182; Vazda (1961), figs. 1-4; (1968),
fig. 31.

Exsiccati examined: Krypt. Vind. 376 (FH, US); Leight. 31 (US);
Malbr. 397 (MSC); Pi§ut 151 (MSC); Rab. 598 (US); VEzda Bohem. 62 (MSC).

Specimen seen: United States. MICHIGAN: Mackinaw County, shore

of Lake Huron at Bush Bay, Harris 9474 (MICH).

 

96

Acrocordia megalospora (Fink) R. C. Harris comb. nov.

 

Pyrenula megalospora Fink, Minnesota Bot. Stud. 2: 329. 1899.

 

Arthopyrenia macrospora Fink, Contr. U.S. Natl. Herb. 14: 237. 1910 BE

 

nom. nov., hom. illeg. (non A, macrospora J. Stein., Ann. Naturhist.

 

 

 

Hofmus. 23: 109. 1909). Arthopyrenia finkii Zahlbr., Cat. Lich. Univ.

 

1: 306. 1921°.EE nom. nov. Lectotype (Harris, 1973): Minnesota,

Mankato, Fink 209 (MICH). Isolectotype in MIN.

Ascocarps 0.5-1.0 mm in diameter. Ostiole often at the end of a
short neck, often displaced from the vertical. Asci 150-250 x 15-25 u.
Spores 33-48(-60) x 15-23 u; spore wall granular ornamented; perispore
rather thick.

Microconidia elliptical, 4-5 x 2 u.

The transfer Of Fink's original name was blocked in Arthopyrenia

 

by the existence of an earlier homonym, but there is no such obstacle

in Acrocordia. Further discussion of this species can be found in

 

Harris (1973).

Illustrations: Figs. 183-186; Harris (1973), fig. 6.
Distribution: Harris (1973), fig. 59.

Additional specimens seen (not cited in Harris, 1973): United
States. MAINE: Knox County, Rockland, l.IX.l9lO Merrill (FH).
MARYLAND: Baltimore County, The Caves, 1.111.1910 EAEEELng (US),

Catonsville, 14.1.1911 Plitt 402 (US), Muby Rd., 22.XII.1906 Plitt 119a

 

(us), Union Dam, 12.IX.1907 Plitt 208 (FH, Us). MASSACHUSETTS:
Bristol County, New Bedford, Willey (FH-TUCK 4025, US); Hampshire
County, Amherst, 1857 TUCkerman (FH-TUCK 4024); Middlesex County,

Cambridge, 1852 Tuckerman (FH-TUCK 4024), Newton, Farlow £6 (FH-TUCK

97

4047), Pepperell, VII.1909 Riddle (FH-RIDD); Norfolk County, Weymouth,
X1.1852 Tuckerman (FH-TUCK 4024). MICHIGAN: Jackson County, Race Rd.

E of Jackson, Harris 9759 (MICH). PENNSYLVANIA: Huntingdon County,

 

Penn. State College Nature Camp, Thomson 1789 (Thomson). VIRGINIA:

 

Page County, Little Stony Man, Imshaug 38670-B (MSC).

 

2. ANISOMERIDIUM (Mu11. Arg.) Choisy

Icon. Lich. Univ., fasc. 3 (sine pagin.). 1928. Arthopyrenia sect.

 

Anisomeridium Mfill. Arg., Flora 66: 290. 1883. Lectotype (Riedl, 1963):

 

Arthopyrenia xylogena Mfill. Arg.

 

 

Lembidium KOrb., Syst. Lich. Germ. 358. 1855 nom. rejic. [non
Lembidium Mitt., 1867 (Hepaticae)1. Holotype: Lembidium polycarpum

KOrb. (= Anisomeridium biforme).

 

Thallus whitish or grayish, endophloeodal or less commonly epi-

lithic or endolithic. Phycobiont Trentepohlia.

 

Ascocarps variable in size and shape, immersed or superficial;
outer layer of ascocarp wall dark brown or blackish, often lacking
below. Hymenial gelatin IKI-. Paraphyses thread-like, branched and
anastomosed. Asci mostly cylindrical or dactyloid, less commonly
narrowly elliptical, narrowly ovate or narrowly obovate to ovate or
obovate; normally with a small ocular chamber. Spores uniseriate, bi-
seriate or irregularly arranged, broadly elliptical, broadly ovate to
narrowly ovate or subcylindrical, mostly 2-celled, rarely 3-4-celled;
cells mostly unequal, often markedly so, lower cell shorter and nar-
rower than upper; spore wall quite thick, up to 2 u; perispore usually

not distinguish able; without ornamentation except in A, adnexum.

98

Microconidia orbicular, elliptical or narrowly elliptical.
Macroconidia simple, orbicular or elliptical.
Lichen substances: Lichexanthone in several species, antraquinone

pigments in A. macrospora.

 

Choisy's use of Anisomeridium as a genus is found in a key without

 

any indication of basionym or species included. It seems clear from the
context that he meant it as a genus and that he was dealing with the

sections of Arthopyrenia as recognized by Zahlbruckner (1921-22, 1926).

 

Acrocordia and Anisomeridium were given generic status by Choisy while

 

 

he retained the other sections in Arthopyrenia. Thus, even though there

 

is no specific reference to Mfiller's section, I feel fully justified in
treating Choisy's name as a change of status rather than a new genus.

Anisomeridium is to some extent composed of species which do not

 

fit into any of the other genera of the Strigulaceae. In spite of this

Anisomeridium seems to be a natural grouping. The orbicular microconidia,

 

simple macroconidia and presence of lichexanthone are diagnostic. The
tendency for the spore cells to be unequal in length and width is also
characteristic. The spore cells may be unequal in other genera of
pyrenolichens but in most cases the lower cell although narrower is

usually as long or longer than the upper. In Anisomeridium the lower

 

cell is usually the shorter. The genus most likely to be confused with

Anisomeridium is Strigula, but the almost unbranched paraphyses in the

 

latter distinguishes them rather easily. Anisomeridium is separated

 

from Acrocordia by the ascus and spore type and from Pyrenocollema by

 

 

the phycobiont and spore type.

Anisomeridium has its greatest diversity in subtropical regions.

 

Anisomeridium biforme is the most widely distributed species, growing

 

99

in temperate to tropical areas in both hemispheres. Of the other species

known to me only A, carinthiacum is found in both Europe and North

 

America. All the rest are American.

Growing on rock ..... ................................. ...... 2

Growing on bark or wood . ......................... . ...... ... 4

2. Spores becoming 4-celled, 16-22 x 6.5-8 u; not
aquatic . ........... . ......................... ....... A, distans

2. Spores remaining 2-celled .. .................... ........ 3

Spores (ll-)13-l8 x 5.5-7 u; more-or-less aquatic on non-

calcareous rock; northeastern U. S. ................ A. carinthiacum

 

Spores 20-25 x 6.5-8.5 u; on limestone, not aquatic;

southern Florida and West Indies .... ............. ........ A. finkii

4. Thallus UV+ yellow (lichexanthone) or rarely UV- in

A. aIleigL—lumo ooooooo o oooooooooooooooooooo 00.00.000.00... 5
4o Thallus UV- 0... ooooooooo coo-o. ooooooooooooooooooooooo o. 7

Spores small, (12-)15-20 x 5-6.5 u; macroconidia cylin-
drical, 7.5-12X2-205u O..0......CIOOOOOOOOOCOO00...... A. tUCkeri
Spores larger, over 25 u long; macroconidia orbicular to

elliptical OOOOOHOOOOOOOOOOIOOOOO ...... ......OOOOOOOOIOOOOOO 6

6. Spores 37-52 x 12-20 u; orange pigment around ostiole

KOH+ purple; macroconidia elliptical, 30-45 x 15-20 u ..

....... .......... ...... ....... ................... A, macrosporum
6. Spores 24-33 x (8-)9-13 u; macroconidia suborbicular

to broadly elliptical, 15-21 x 10-15 u ............. A, ambiguum

:11.

11.

100

Spore wall ornamented; spores broadly ovate, 25-33 x

15-18 u; spore cells strongly unequal ................... A, adnexum

Spore wall smooth . ........ ..... .................... ....... 8
8. Spores small, 9-13 x 4-5 u ....... ............. .... A, albisedum
8. Spores larger, over 10 u in length . ....... ............. 9

Spores becoming 4-ce11ed, 14-20 x 4.5-6 u; microconidia

elliptical; mainly northern (as far south as North

Carolina) ... ............ ..... ....................... A, willeyanum
Spores remaining 2-celled; microconidia mostly orbicular

(elliptical in A, subprostans); mostly in the coastal

 

plain (A, leucochlorum as far inland as southern Ohio

 

and Illinois) ... ..................... ........ ......... ..... 10

10. Asci slender, cylindrical or dactyloid .. .......... .... ll
10. Asci ovate or obovate to narrowly ovate or narrowly

obovate .... ..................................... ...... l3

Spores broadly elliptical to ovate, rarely narrowly

ovate, 10-18 x 4.5-7.5 u; common on both coasts, rarer

in the South . ............................. ............. A, biforme
Spores slender, narrowly ovate to almost cylindrical

(often slightly bent in A, tamarindi); coastal plain

and west Ind-es .00.... ........ 0...“...00 ....... .....O...... 12

12. Microconidia elliptical; macroconidia not known;

spores 12-18(-2l) x 4.5-6.5 u, not bent ....... A, subprostans

 

101

12. Microconidia orbicular; macroconidia elliptical,
4-7 x 2.5-3.5 u; spores 15-22(—24) x 4-5.5(-6) u,

often slightly bent ............................. A, tamarindi

l3. Spores 12-18 x 5-7.5 u; ascus wall thin and even; macro-
conidia glObose ................................... _A-O laucocrllorum

l3. Spores larger, 15-22 x 5.5-8.5 p .... ............ .......... 14

14. Ascus wall thick, thickened unevenly; spores 15-22 x
605-8051]. 0. oooooooooooooooooooooooooo ...-0.00.000. A. feeanum

14. Ascus wall relatively thin and even; spores 18-22 x

505‘?“ coo-coco... oooooooooo o... oooooooo 000.00 £- sanfordense

fgjjgsomeridium adnexum (Mfill. Arg.) R. C. Harris comb. nov.
Arthopyrenia adnexa MUll. Arg., Flora 66: 304. 1883. (adnaex err.
IEKEio). Leiophloea adnexa Riedl, Sydowia 16: 266. 1963. Lectotype:

Brazil, Apiahy, Puiggari 240 (G).

Asci ovate. Spores irregularly arranged, ovate; cells markedly
urleCIUal; 25-33 x 15-18 u; spore wall strongly ornamented.
Microconidia orbicular, c. 2 u in diameter.

The collection chosen as the lectotype is the one which Riedl (1963)

FEas <discussed in some detail. The syntype from Rio de Janeiro is another

8 . . . . . . . .
peGiles of Anisomeridium. Anisomeridium adnexum has not been found in

 

North America.

Illustration: Figs. 187-190.

Specimens seen: Brazil, Apiahy, Puiggari 240, 1007 (G).

fiEEL§9meridium albisedum (Nyl.) R. C. Harris comb. nov.

102

Verrucaria viridiseda f. albiseda Nyl., Lich. Japon. 108. 1890.

Porina viridiseda f. albiseda (Nyl.) Zahlbr., Cat. Lich. Univ. 1: 409.
1922. Lectotype: Florida, Calkins (H-NYL 717). Isolectotypes in

MICH? and US.

Thallus whitish, endophloeodal.

Ascocarps hemispherical to subglobose, small, 0.2-0.3(-0.4) mm in
diameter. Asci cylindrical, occasionally slightly broader toward the

base, 50-75(-85) u in length.
slightly constricted at the septum, 9-13 x 4-5 u.

Spores nearly uniseriate to subbiseriate,

narrowly ovate,
Microconidia narrowly elliptical to rectangular, 3.5-4.5 x 1.2-2 u.

Habitat on bark, usually Quercus but once on Myrica.

The other syntype of V, viridiseda f. albideda (H-NYL 718) is
Anisomeridium albisedum is the smallest spored species

gigrigula phaea.
Strigula phaea has spores similar in size and shape, but

irl the genus.
djgffers in having a darker thallus, unbranched paraphyses and 2-ce11ed

Anisomeridium albisedum is known only from northern and

macroconidia.

Central Florida.

Illustrations: Figs. 191-194.
United States. FLORIDA: sine loc., Calkins (FH,

Specimens seen:
Ii‘bfij, MICH, US); County unknown, Livezey's Creek, Calkins (MICH); Duval

County, Jacksonville, Calkins (FH, MICH, MSC); Liberty County, Apalach-

 

leciléi Nat. Forest, Whitehead Lake Campground, Harris 1549 (MSC); Seminole

c
Ounty, Sanford, X.1910 Rapp (FH), 31.IX.1912 Rapp (FH).

An'
“‘5EEEE§Qeridium ambiguum (Zahlbr.) R. C. Harris comb. nov.
,firthopyrenia (Acrocordia) ambigua Zahlbr., Ann. Mycol. 33: 35. 1935.

GEQt-‘Dtype: Florida, Sanford, V.l923 Rapp 142 (W, middle piece of three

<3
I1 Qard) .

103
Thallus whitish or grayish, UV+ yellow (lichexanthone) or rarely UV-.

Ascocarps immersed, hymenium ovate; ascocarp wall usually thick above

and extending outward to form a shield, thin or lacking below. Asci

cylindrical. Spores nearly uniseriate, ovate to narrowly ovate, ends

Often somewhat pointed, slightly constricted at the septum, 24—33 x

(8-)9-l3 u.

Microconidia elliptical to narrowly elliptical, 3-4 x 1.5-2 p.

Macroconidia suborbicular to broadly elliptical, 15-21 x 10-15 u.

Habitat on soft bark of a variety of trees.
The middle piece on the card is designated as the lectotype since

it seems to be the piece studied by Zahlbruckner. However, it is the

only specimen I have seen which I have seen whiCh lacks lichexanthone.
It is identical to lichexanthone containing specimens in all other

rwespects. The two syntypes on the card do contain lichexanthone.

Anisomeridium ambiguum is one of three closely related species hav-
iTKJ similar spores, asci and chemistry. Anisomeridium tuckeri has

Eflnailler spores and A, macrosporum larger. Anisomeridium ambiguum occurs
irl Ilorthwestern Florida and Jamaica.
Illustrations: Figs. 195-199.

Specimens seen: United States. FLORIDA: Putnam County, Oklawaha

Rjj’SEIT at Fla. Hwy 19, Harris 2200 (MSC); Seminole County, Sanford,
1C).\f-jl915 Rapp 167 (ABSL, US), V.l923 Rapp 142 (W), Sanford, Robinson

S .
ElrtLrlgg, 11.111.1915 Rapp 107 (US); Volusia County, Daytona [=Daytona

:B
each] , 1.1898 Thaxter _1_5_ (FH).

VVest Indies. JAMAICA: Clarendon Parish, N slope of Portland Ridge,

W 13264 (msc).

104
Anisomeridium biforme R. C. Harris comb. nov.
Verrucaria biformis Borr. ;2_Hook. & Sowerby, Suppl. Engl. Bot.

1: pl. 2617, f. 1. 1831. Arthopyrenia biformis (Borr.) Massal., Framm.

Lich. 26. 1855. Type collection not seen.
Arthopyrenia brachyspora Mfill. Arg., Nuovo Giorn. Bot. Ital.

21: 52. 1889. Holotype: Argentina, Isla de los Esados, 1888

Spegazzini (G).

Arthopyrenia parvula Zahlbr., Beih. Bot. Centralbl. 13: 149. 1902.

Leiophloea parvula (Zahlbr.) Riedl, Sydowia 16: 269. 1963. Type

collection: California, Santa Monica Mtns., Malibu Canyon, VIII.1898

Hasse 759. Isotypes in ABSL, FH & US.

Arthopyrenia amaura Zahlbr. lg Handel-Mazzetti, Symbol. Sinic.

3: 19. 1930. Type collection: China, Hunan Province, near Tschangscha,

11).XII.1917 Handel-Mazzetti 11426. Isotype in US.

Ascocarps hemispherical to globose, immersed or emergent,
O-3--O.6 mm in diameter. Asci dactyloid to cylindrical, 65-130 x
1C3-515 u. Spores subbiseriate to uniseriate, narrowly ovate to elliptic-
a]_, Sometimes slightly constricted at the septum, 10-17 x 5-811.
lflicroconidia orbicular, 2-3 p in diameter.
Pflacroconidia relatively rare, orbicular to elliptical or ovate,
3-{3 ll in diameter or 3.5-5 x 2.5-3 u.
I‘iabitat on bark or old wood.
lklthough I have not seen the original material of this species, I
lianree sseen several specimens collected and determined by Borrer. 'AAigg-

The: ~ .
“*SEEESE£umibiforme is apparently cosmopolitan in its distribution. In

Eici . .
<135tlon to numerous collections from Europe and North America I have

105

verified collections from Asia (China and India), New Zealand (Campbell
15.), Bermuda, the West Indies and southern South America (Falkland Is.,

Isla de los Estados). As would be expected there is some variation in

the spores from region to region. The most strikingly different popu-

lation is the one in eastern North America in which the spores are

almost always elliptical and uniseriately arranged in the ascus. This

has caused it to be confused with various Acrocordia species but the

lack of any ornamentation of the spores removes it from that genus.
However this type of spore is found occasionally in the southeastern

United States/West Indian and the western North American populations.

The southeastern/West Indian population has the smallest asci. Aniso-

Ineridium biforme in North America is mainly coastal with relatively

:few inland stations. One wonders if the gap in its distribution be-

tnveen Maryland and Florida is real or artificial.
Illustrations: Figs. 200-207; Harris (1973), figs. 2-3.

I{iczhardson & Morgan-Jones (1964), fig. 4. Distribution: Fig. 417.

Exsiccati examined: Anxi Ven. 132 (FH-TUCK 4062); Desm. ed. 11,

Seirfl- II: 400 (FH); Hepp 953 (FH, FH-TUCK 4065); Krypt. Vind. 375 (FH,
Us) ; Leight. 100 (FH, FH-TUCK 4064, US); Merr. I: 147 (CAN, FH, FH-
RIDD, MICH, US); Merr. I: 164 (CAN, FH, FH-RIDD, MICH, us); Nyl. Par.
91 (PH-TUCK 4065); Nyl. Par. 92 (FH-TUCK 4065); Rab. 483 (FH-TUCK 4065,

MICH); Rel. Hasse 79 (FH); Zahl. 190 (MICH, w).

£3e1ected specimens seen: Canada. BRITISH COLUMBIA: Vancouver

I
gland, Sidney, 19.x1.1912 Macoun (FH), Victoria, 28.1V.1914 Macoun

3l\4 FH).

‘United States. CALIFORNIA: Los Angeles County, Malibu Canyon,

\r
III~1898 Hasse 759 (ABSL, FH, US), Santa Catalina Island, IX.1912

106

Hasse (ABSL, FH), Santa Monica Canyon, V1.1914 Hasse (FH); Orange
County, Laguna Creek, 1.1.1908 Herre 1240 (FH), near Newport, 1911
Hasse (US); Santa Cruz County, Santa Cruz Peninsula, Hasse (MICH).

FLORIDA: Dade County, Everglades Nat. Park, S of Pa-hay-okee Overlook,

Harris 2730-A, 2826-E (MSC); Liberty County, Apalachicola Nat. Forest,
Porter Lake Picnic Area, Harris 1634 (MSC); Palm Beach County, W. Palm
Beach, Thaxter 21_(MICH); Seminole County, Sanford, 1.1909 Egpp_(FH),
4°II°1914.§EEE (FH). ILLINOIS: Cook County, Calkins (FH, MICH,
Thomson, US, US ex M0). KENTUCKY: McCreary County, SW of Corbin, Harris

1253 (MSC). LOUISIANA: E. Baton Rouge Parish, S of Baton Rouge, Tucker

7403-8, 8846, 8846-B (LSU); Lafayette Parish, Lafayette, Tucker 7164

 

(LSU); Plaquemines Parish, Pointe-a-la-Hache, 23.XII.1885 Langlois 796
(US). MAINE: Hancock County, Mt. Desert Island, Salisbury Cove,
113.VIII.1932.21£EE (US); Knox County, Rockland, 10.X.1909 Merri11 (CAN,
F11, FH-RIDD, MICH, US), Thomaston, 24.VII.1910 Merri11 (CAN, FH, FH-
IQJHJD, MICH, US), Warren, 21.VIII.1909 2113:.ggg (FH, US). MARYLAND:
Anne Arundel County, road to Elvaton, 11.VI.1910 ME (FH, US);
Ehal:timore County, road to Perry Hall, 7.VI.1910 Plitt 261a (FH, US).
NLASSACHUSETTS: Barnstable County, Chatham, Eggdg 4396 (CAN, us);
BriS'tol County, New Bedford, Willey (US); Middlesex County, Concord,
1863 Man}; (FH-TUCK 4024, us); Plymouth County, Wareham, 1919 Robbins
(Ffiij); Suffolk County, Waverly, 11°X°1894.§E£E (FH); Worchester County,
'8‘ I"lilford, X1.1852 Tuckerman 13 (FH-TUCK 4024). MICHIGAN: Washtenaw
(:CNJIthy, NW of Crooked Lake, Harris 869 (MSC). NEW HAMPSHIRE: White
Dqt33£3~ , Tuckerman (FH-TUCK 4024). NEW JERSEY: .§£Q2.lgg., 1876 Austin

63
‘~S¥§; (EH-TUCK 4025). OREGON: County unknown, sea coast, 2.XI.1881

P .
.IJEEEEEQIE g (FH, US). PENNSYLVANIA: Pike County, Bushkill Falls,

107

28.1V.1940 Nearing (WIS). WASHINGTON: San Jaun County, Friday Harbor,

28.VI.1906 Fink 40_(M1CH).

Anisomeridium carinthiacum (J. Stein.) R. C. Harris comb. nov.

Arthopyrenia carinthiaca J. Stein., Oesterr. Bot. Z. 63: 335. 1913.

("carintiaca", corr. Zahlbr., 1921). Lectotype not selected.

Isosyntype: Austria, near Klagenfurt, Steiner (US ex MO).

Arthopyrenia dimidiata Fink ;A_Hedrick, Mycologia 25: 306. 1934.

Holotype: Connecticut, near Ellsworth, 1895 Green (MICH, hb. Fink

15,469).

Thallus epilithic, whitish, greenish or tan; black hypothallus not

evident in some pecimens to dominant in others.

Ascocarps hemispherical to subglobose, semi-immersed, 0.2-0.3 mm in

Ctiameter. Asci narrowly elliptical to narrowly obovate; ascus wall

tfrickened at tip; 50-60 x 13-17 u. Spores irregularly arranged, ovate

tc> almost elliptical, (ll-)l3-18 x 5.5-7,1.
Microconidia narrowly elliptical, 4-5 x 1.5-2 u.

Habitat on non-calcareous rock along streams, probably occasionally

inLlrIdated.

This species is unusual in its habitat and in the variability of

:Ltss lhypothallus. There is no evident hypothallus in any other species

ch ifiifiisomeridium. In the Michigan collections the lichenized thallus

‘155 (DiFten restricted to isolated patches surrounded by black hypothallus.
33163 1”Typothallus is present but not well developed in the isosyntype of
£3- 5:51rinthiacum and is apparently lacking in the Connecticut collections.
'Inj‘a Semi-aquatic habitat suggests Pyrenocollema, but the phycobiont, asci

a
r1c1 Spores are not characteristic of that genus but rather of Anisomeridium.

108

Illustrations: Figs. 208-212.
Specimens seen: United States. CONNECTICUT: Litchfield County,
Aton Forest, Hale 137 (US), Ellsworth, 1895 Green (MICH). MICHIGAN:

Delta County, NE of Rapid River, Harris 4202-B, 4210-A (MSC).

 

Anisomeridium distans (Willey) R. C. Harris comb. nov.

 

Verrucaria distans Willey, Enum. Lich. New Bedford 38. 1892.

 

Arthopyrenia distans (Willey) Zahlbr., Cat. Lich. Univ. 1: 276. 1921.

 

Type collection: Massachusetts, New Bedford, head of Purchase St.,

1866 Willey. Isotype in MICH.

Thallus epilithic, whitish to brownish, thin.

Ascocarps subglobose, 0.1-0.3 mm in diameter. Asci narrowly
elliptical or narrowly obovate, c. 70-80 x 15-18 u. Spores irregularly
arranged, narrowly ovate to narrowly elliptical, 2-4-ce11ed, not con-
stricted at the septa, 16-22 X 6.5-8 u.

Microconidia narrowly elliptical, 4-5 x 1.5-2 u.

Macroconidia elliptical, 5-5.5 x 2.5-3 u.

Habitat on moist rocks but apparently not aquatic.

Anisomeridium distans is unusual in the genus in that it grows on
rock and has 4-celled spores. The phycobiont is hard to determine with

certainty. Trentepohlia is present in both collections but another alga,

 

probably a blue-green, is also present.

Illustrations: Figs. 213-218.

Specimens seen: United States. MASSACHUSETTS: Bristol County,
New Bedford, 1866 Willey (MICH). OHIO: Coshocton County, near Cosh-

octon, Moldenke 14399 (Us ex M0).

 

109

Anisomeridium feeanum (M611. Arg.) R. C. Harris comb. nov.

 

Arthopyrenia feeana Mfill. Arg., Mem. Soc. Phys. Hist. Geneve

 

30: 28. 1888. Holotype: [West Indies] on Croton cascarilla (G).

Ascocarp hemispherical to subglobose, semi-immersed to immersed;
ascocarp wall sometimes extended outwards above to form a shield, thin
or lacking below. Asci narrowly obovate or rarely narrowly elliptical,
60-80 x 13-20 u. Spores irregularly arranged, ovate, lower cell markedly
shorter, occasionally slightly consticted at the septum, 15-22 x 6.5-8.5 u.
Microconidia in one collection (Thaxter 22) orbicular, c. 2.5 p in

diameter, in another (Harris 1306) elliptical, c. 3 x 1.5-2 u.

 

Macroconidia elliptical, 5-6 x 2.5-3.5 u.
Habitat on various barks.

Anisomeridium feeanum is very close to A, biforme and along with

 

.5- leucochlorum and A, sanfordense forms an intergrading group of species

 

 

which are maintained as distinct on a provisional basis. Specimens
tentatively identified with A, feeanum come from Tennessee, Florida,
the West Indies and Hong Kong.

Illustrations: Figs. 219-224.

Specimens seen: United States. FLORIDA: Dade County, Coconut
Grove, 1898 Thaxter_22 (FH); Volusia County, Daytona Beach, Thaxter
(MICH). TENNESSEE: Wilson County, Cedars of Lebanon St. Park, Harris

1306 (MSC).

Anisomeridium finkii R. C. Harris sp. nov.

 

Anisomeridium ascocarpis in saxo calcareo immersis, sporis

 

20-24 x 6.5-8.5 u et microconidiis ellipticis, 3-5 x 1.5 u.

110

Holotype: Puerto Rico, Arecibo District, Manati, 12.1.1916 Fink

2052 (US). Isotype in FH.

Thallus endolithic, whitish.

Ascocarps immersed in Puerto Rican specimens, emergent in Florida
specimens, subglobose or with upper part of ascocarp wall extended
outward to form a shield; ascocarp wall thinner below. Paraphyses
branched and anasomosed, embedded in abundant hymenial gelatin. Asci
cylindrical, thickened at the tip with a small ocular chamber,

105-125 x 15-18 u. Spores subbiseriate, narrowly ovate, lower cell
usually shorter, 20-25 x 6.5-8.5 U.

Microconidia elliptical, 3-5 x 1-l.5 u.

Habitat on calcareous rock.

The collection in US was designated as the holotype Since the one
in Fink's herbarium contained only a discocarpous lichen associated with

a blue-green alga. The material was named Thelidium immersum by

 

Merrill. The other saxicolous species of Anisomeridium grow on non-

 

calcareous rock and are temperate in distribution.

Illustrations: Figs. 225-229.

Specimens seen: United States. FLORIDA: Dade County, Coconut
Grove, XII.1897 Thaxter (MICH).

West Indies. PUERTO RICO: Arecibo District, Manati, 12.1.1916
EEQEHggig (FH, US); Mayagfiez District, Yauco, 20.XII.1915 Eigkwligg

(FH).

Anisomeridium leucochlorum (Mall. Arg.) R. C. Harris comb. nov.

 

Arthopyrenia leucochlora Mfill. Arg., Flora 66: 287. 1883.

 

'.

s ‘

 

 

A‘aI! b. t -

r4,

111
Type collection: Massachusetts, New Bedford, Willey. Isotype? in US
and MICH.

Ascocarps often immersed, hemispherical to subglobose, ascocarp wall
lacking below, 0.2-0.3 mm in diameter. Asci narrowly obovate, ascus wall
rather uniform, c. 1.5-3 u thick, 53-72(-77) x 12-17(-20) u. Spores
irregularly arranged, narrowly ovate to ovate, 12-18 x 5-7.5 u.

Microconidia orbicular, 2-2.5 u in diameter.

Macroconidia orbicular, often slightly tinted, 2.5-4 u in diameter.

Habitat on bark.

This species is maintained with some hesitation, since it is vir-
tually identical to A, biforme in most respects. It differs in the size
and shape of the ascus and in the spore arrangement. It is possible
that the material placed here represents an extreme form of A, biforme,
and it is often difficult to place some specimens with certainty.

There are a number of specimens named "Pyrenula leucochlora" by
Willey, but only one, on maple, fits Mfiller's description of the thallus
which is rather peculiar and seems to me to be the result of a foreign

thallus overgrowing the Anisomeridium. All the other specimens seen

 

have a typical endophloeodal thallus. The other specimens collected by
Willey are all A, willeyanum.

Illustrations: Figs. 230-235.

Specimens seen: United States. CONNECTICUT: Middlesex County,
Old Saybrook, Eyggg & EEEEE ZZZ (FH). ILLINOIS: sine loc., Wolf (MICH,
US); Menard County, Athens, H2ll_(MICH). MASSACHUSETTS: Bristol County,
New Bedford, Willey (MICH, US, US ex MO). OHIO: Oxford County, Sloat's

Hill, 17.1V.1927 Fink (MICH).

 

Azll'..- ‘

112

Anisomeridium macrosporum R. C. Harris sp. nov.

 

Species distinctissima thallo UV+ aureo (lichexanthone), ostiolis
eccentricis pigmento aurantiaco circumdatis et sporis magnis, 39-53X
12-15 u.

Holotype: Florida, Dade County, Everglades Nat. Park, NE of

Pa-hay-okee Overlook, Harris 2791-A (MSC). Isotypes in BM, H, TNS, US,

 

Vézda.

Thallus white, UV+ yellow (lichexanthone).

Ascocarps mostly immersed, hemispherical to subglobose, 0.5-0.8 mm
in diameter, ascocarp wall often extended outwards above to form a shield,
thinner or lacking below; ostioles mostly eccentric, surrounded by a
small ring of orange pigment which is KOH+ purple. Asci cylindrical
with a distinct ocular chamber, 150-200 X 20-30 u. Spores subbiseriate
to biseriate, narrowly ovate, often with rather pointed ends, lower cell
markedly shorter, 37-52 X 12-20 u.

Microconidia elliptical, 2.5-4 X 1.2-2 p.

Macroconidia elliptical, 30-45 X 15-20 u.

Habitat on bark, including Taxodium.

This remarkable species is known only from a single small region of
the Everglades National Park but it is rather abundant there.

Illustrations: Figs. 236-241.

Specimens seen: United States. FLORIDA: Dade County, Everglades

Nat. Park, NE of Pa-hay-okee Overlook, Harris 2785, 2791-A, 2810, 2813-A,

 

Anisomeridium sanfordense (Zahlbr.) R. C. Harris comb. nov.

Arthopyrenia (Mesopyrenia) sanfordensis Zahlbr., Ann. Mycol. 33: 33.

 

 

 

113

1935. Holotype: Florida, Sanford, IV.1932 Rapp 134 ("139" in original

description), (W).

Ascocarps small, subglobose, 0.15-0.2 mm in diameter. Asci obovate,
50-63 X 16-23 u; ascocarp wall rather thin, 1-2 p thick. Spores irregu-
larly arranged, narrowly ovate with rather pointed ends, 18-22 X 5.5-7 u.

Microconidia orbicular, c. 2 u in diameter.

Habitat on bark (Diospyros).

Anisomeridium sanfordense is little different from A, leucochlorum,

 

hauling broader asci and longer spores. It is known only from the type

collection.

Illustrations: Figs. 242-245.

fggixsomeridium subprostans (Nyl.) R. C. Harris comb. nov.
Verrucaria subprostans Nyl., Expos. Syn. Pyrenoc. 56. 1858.
figzgrnopyrenia subprostans (Nyl.) Mu11. Arg., Flora 66: 317. 1883.

Hc310type: South Carolina, Ravenel (H-NYL Pm. 7378). Isotype in US.

,Ascocarps hemispherical to subglobose, 0.3-0.5(-0.6) mm in diameter,
aSCCXZEirp wall thinner or lacking below. Asci dactyloid to cylindrical,
75-14255 x 9-12 p. Spores uniseriate to biseriate, narrowly ovate, often
With -rYather pointed ends, slightly constricted at the septum, 12-18(-21) x
4.5-6- 5 ll-

IVIicroconidia narrowly elliptical to elliptical, 2.5-4 X 1.2-2 u.

Macroconidia elliptical, 6-6 x 2.5-5 u-

I‘Ieibitat on bark, especially EEEESEHEV

Nl’lander in his original description indicated the type as coming
frqn Te‘Xas. However, the specimen cited above bears notes in Nylander's

h .
andwxnlfaing which agree with the published description. As far as I

114

know the species is not found in Texas.

Anisomeridium subprostans differs from the closely related A, biforme

 

and A, tamarindi by its elliptical microconidia and its spore shape.
Illustrations: Figs. 246-250.
Specimens seen: United States. FLORIDA: Monroe County, Big Pine
,Key, 14.1V.1921 Kgliy (US). LOUISIANA: Parish unknown, Bayou Millieu,
26.X.1896 Langlois (US). SOUTH CAROLINA: _s__i£e__1£g., Ravenel (H-NYL, US).

West Indies. PUERTO RICO: San Juan District, Rio Piedras, 1.XII.1915

 

Fink 536 (MICH, US).

_égxisomeridium tamarindi (Fee) R. C. Harris comb. nov.

Verrucaria tamarindi Fée, Suppl. Essai Crypt. Ecorc. Off. 85. 1837.

 

fflgiiina tamarindi (Fee) MUll. Arg., Mém. Soc. Phys. Geneve 30: 24. 1888.
IKECTtotype: "ad cortices Tamarindi officinalis L. Antillarum" (Guadeloupe

fixi- Fée, loc. cit. p. 165) (G). Isolectotypes in G and H-NYL 748.

Ascocarps immersed, hemispherical, 0.4-0.6 mm in diameter; ascocarp
walJl forming a shield, lacking below. Asci dactyloid to cylindrical,
75-341() x 10-12 u. Spores uniseriate to biseriate, narrowly ovate to
OVatxa, longer spores often somewhat bent, 15-22(-24) x 4.5-5.5(-6) u.

.Pticroconidia orbicular, c. 2 u in diameter.
Pqéicroconidia elliptical, 4-7 X 2.5-3.5 u.
fiéibitat on bark, in the West Indies often on coconut palm.

:fEEfiisomeridium tamarindi differs from A, biforme by longer, narrower,

 

o
ften 1Dent spores and by elliptical macroconidia. The spores in some

c . .
Ollecztlnuons of A, tamarindi approach those 0f.fl: subprostans which has

 

Elli - . . . . . . . . . .
ptlcZal microconidia. Anisomeridium tamarindi is not uncommon in

 

_ F ‘
lor1¢h3_ and Louisiana. It also occurs in the Bahamas and the West Indies

 

175...? I. L .

.413.

 

115

(Bequia, Cuba, Dominica, Grand Cayman, Guadeloupe, St. Vincent and

Trinidad).
Illustrations: Figs. 251-256. Distribution: Fig. 418.

Specimens seen: United States. FLORIDA: County unknown, Pumpkin

Key, IV.1921 Kelly 8_9_, 148 (MICH); Dade County, Coconut Grove, 1898

Thaxter 313 (MICH), Everglades Nat. Park, NE of Pa—hay-okee Overlook,

Harris 2791-B (MSC); Monroe County, Key West, 11.1898 Thaxter 324c (MICH);

Palm Beach County, Palm Beach, 1898 Thaxter i8 (FH); Volusia County,

 

Tomoka St. Park, Harris 2280 (MSC). LOUISIANA: E. Baton Rouge Parish,

Baton Rouge, Tucker 9129 (LSU); St. Martin's Parish, Bois Charmant,

27.VI.1895 Langlois (US).

misuneridium tuckeri R. C. Harris sp. nov.

Sat similis A. ambigui sed sporis minoribus, 15-20 X 5-6 ‘11 et
macroconidiis cylindricis, 7.5-12 X 2-2.5 ll.
Louisiana, E. Baton Rouge Parish, Baton Rouge, LSU

Holotype:

PrOperty off Essen Lane, Tucker 10003 (LSU).

Thallus gray or whitish, UV+ yellow (lichexanthone).

Ascocarps mostly immersed, hemispherical, 0.5-0.6 mm in diameter;
aSCocarp wall often extending outward above to form a shield, lacking
bElOW- Asci cylindrical, 90-115 X 10-12 11. Spores uniseriate to sub-

biseriate, narrowly ovate, slightly constricted at the septum, (12-)15-20 X

5-6.5 11.
IVLiCroconidia elliptical to suborbicular, 2.5-3 X 1.5-2 11.
MaCroconidia cylindrical, 7.5-12 X 2-2.5 '0.

Habitat on various barks.

This interesting species is named for Dr. Shirley Tucker whose

116

diligent collecting has brought to light many novel pyrenolichens. The

presence of lichexanthone and the spore shape relate A, tuckeri to A.

ambiguum and A, macrosporum. While all the other species of the genus

have orbicular to elliptical macroconidia, A, tuckeri is unique in having

cylindrical macroconidia.
Illustrations: Figs. 257-261. Distribution: Fig. 419.

Specimens seen: United States. FLORIDA: Putnam County, Oklawaha

RjAIer at Fla. Hwy. 19, Harris 2266 (MSC); Seminole County, Sanford,

1£9.V,1919 RaEE (FH), VII.1920 Rapp 626 (FH, MICH). LOUISIANA: E. Baton

chage Parish, Baton Rouge, Tucker 10003 (LSU); Livingston Parish, 3 mi S

of? Livington, Tucker 7357 (LSU); St. James Parish, near Frenier, Koch

 

7CH58 (CAN); St. Martin Parish, Duchamps, 26.1X.1894 Langlois (US).

fglfigsomeridium willeyanum (R. C. Harris) R. C. Harris comb. nov.

Arthopyrenia willeyana R. C. Harris, Michigan Bot. 12: 16. 1973.
Phalcytype: Michigan, Iosco County, N of Tawas City, Harris §l§_(MSC).
Isotypes in B, BM, CAN, COLO, Duke, H, Harris, LD, MIN, 5, TNS, UPS,
US, ve'zda & WIS.

ESee Harris (1973) for a detailed description. Anisomeridium
3E£3£§2fi§ggm_is distinquished by its small, rather shiny ascocarps, nar-
rOWI)’ <3vate spores which often become 4-celled and elliptical micro-
conidjLER. The known range of the species has been extended southward by
a'C013Leection from Alabama. Two collections by Sandstede in Germany
consth1:Lrte the first records for Europe.

Illustrations: Figs. 262-265; Harris (1973), figs. 16-17.

D‘ -
lStrlbution: Fig. 420.

AC1Ciitional specimens seen (not Cited in Harris, 1973): United

 

 

117

States. ALABAMA: Cleburne County, N of Heflin, Harris 1330 (MSC).

CONNECTICUT: Litchfield County, Aton Forest, EBA-3.122. (US). DELAWARE:
Sussex County, 5 mi W of Georgetown, £11332 _9_5_4_2_ (CAN, MSC). ILLINOIS:
Cook County, Calkins (MICH). KENTUCKY: McCreary County, SW of Corbin,

Harris 1192 (MSC). MAINE: Waldo County, Freedom, Parlin 7534 (FH).

 

MASSACHUSETTS: Bristol County, Fairhaven, 1876 Willey (US), New Bedford,
Willey (FH-TUCK 4024, 4051, US); Middlesex County, Concord, 1863 Mary;
(FH-TUCK 4024), Wayland, IX. 1909 Riddle l8 (FH-RIDD);Suffolk County,
Newton, (Farlow) (FH). MICHIGAN: Baraga County, 5 mi NE of Sidnaw,

iarris 7839 (MSC); Jackson County, E of Leeke Lake, Harris 7712 (MSC);

 

Lenawee County, Bean Creek at Lime Creek Rd., Harris 7695, 7699 (MSC);
Mackinac County, 8 mi E of Naubinway, Harris 9627 (UMBS); Newaygo County,
ENE of White Cloud, Harris 7761, 8162 (MSC). MISSOURI: Saline County,
Davis Creek-Bottom, 12.VIII.1896 Demetrio (FH). NEW JERSEY: _s_i_n_e_ _10_c.,
1876 Austin 6% (FH-TUCK 4025). NEW YORK: Madison County, Bridgeport,
7- II-1959 _ngcgz (CAN); Saratoga/Washington County, Hudson Falls, IX.1919
m1“. (MICH); Suffolk County, Long Island, _B_r_‘_o_c_12 21$ (CAN, msc).
NORTI‘II CAROLINA: Transylvania County, Lookout over John Rock, Harris 3360
(MSC). OHIO: Lake County, Painesville, 8.X.1915 _Fi_nk__6_9_l_ (MICH).
PENNSYLVANIA: Clarion County, near Cooksburg, god—o _5_52_9_ (CAN);
Huntingdon County, Penn. St. Nature Camp, Thomson 1798 (Thomson).
RHODE ISLAND; _s_i_rle: 1&1.” Bennett (MICH). TENNESSEE: Wilson County,
Cedars of Lebanon St. Park, Harris 1304-C (MSC). VIRGINIA: Madison
Co‘mty, Stony Man, Imshaug 38551, 38569 (MSC); Page County, Big Meadows:
% 238;, gig (CAN), 9183 (CAN, MIN), Wetmore 15299 (MIN). WEST

V
RGINIA: Randolph County, Gaudineer Scenic Area, Harris 5974, 5977 (MSC).

‘- I}- J

118

3. PLEUROTREMA M1111. Arg.
Bot. Jahrb. Syst. 6: 388. 1885. Lectotype (Clements, 1909):

Parathelium polysemum Nyl. EPleurotrematomyces Cif. & Tomas., Ist. Bot.

 

Reale Univ. Reale Lab. Crittog. Pavia Atti, ser. 5, 14: 6. 1957.
Holotype: Parathelium polysemum Nyl.

Ditremis C1em., Genera of Fungi 41. 1909. Holotype: Pleurotrema

inspersum Miill . Arg. 5

I'm-‘1‘. =7'

Thallus whitish, cream color or grayish, endophloeodal.

Phycobiont Trentepohlia or apparently non-lichenized.

 

Ascocarps flattened, broadly elliptical or orbicular in outline,
HKDsdzly not much immersed; ascocarp wall brownish to blackish, thinner
0:? Jwacking below. Ostiole eccentric to lateral, occasionally at the
erki Cfi:a short projecting neck. Hymenial gelatin IKI-. Paraphyses
ifldxreaad-like, branched and anastomosed. Asci cylindrical with a small
OCHJJJar chamber. Spores uniseriate, rarely biseriate, ovate to narrowly
OVEitHE, often with somewhat pointed ends, 2-celled, cells mostly approxi-
matEJ—y equal, usually constricted at the septum; spore wall usually
graITttlar ornamented, at least in age; perispore thin, often not dis-
cernable.

Dflicroconidia narrowly elliptical to elliptical.

P‘Iabitat on bark.

Cllements (1909) chose P. polysemum (Nyl.) Mfill. Arg. as the type
of tjifie genus. However, at this time I do not know whether Mfiller ever
Saw ‘a- specimen of this species or whether he made the transfer solely
on 1:?“E basis of Nylander's description. If Mfiller had not seen any

matenrial of P. polysemum, I would recommend that P, inspersum should

119

be the lectotype of Pleurotrema. The choice of lectotype is of con-

siderable importance since I do not believe P, polysemum and

_E. inspersum to be congeneric. The bulk of the species previously

described belong with _P. inspersum and retypification, if possible,

would avoid a number of name changes. If Mfiller did have material of

_E. polysemum available when he described Pleurotrema, then the correct
name for the genus as accepted here is Ditremis, since in that case

there would seem to be no basis for overturning Clement's lectotypifi-

cation.

In addition to the eccentric ostiole, Pleurotrema can be distin-

guished by its cylindrical asci, uniseriate spores, spore shape and

usually ornamented Spores. All of the known species are tropical or

Subtropical in distribution. The genus has not been previously re-

ported from North America and is represented by only a single collection.

A few additional species with parathelioid ascocarps belonging to

other genera have been included in the key to avoid confusion.

1' Spores 2/ascus; ostioles mostly upright but more or less

......... Arthopyrenia bifera

 

eCcentric in some collections . ..... ..
1' Spores 8/ascus ........ ........... ..... . ...... ....... 2

2- filallus UV+ yellow .................. .......... ......... 3

2- quallus W- ....................0....................... 4
3.

Spores 2-ce11ed, 30-45 x 15-20 11; spore wall not orna-

Inerlted; orange pigment around ostiole, ...... ........... .

.....O........ ........ ................... AIfi—Smeridim maCrosporum

 

 

120

Spores 4-celled, 28-36 X 9-12 u; spore wall ornamented

.......................................... Polymeridium exasperatum

 

40 Spores 4-CEllEd .000000000000000000... POlymeridlum exasperatum

40 Spores 2-C€ll€d . oooooooooooooooooooooo 0.000.000.0000... S

Spores l7-25(-30) X 5.5-7.5 u ............... Pleurotrema anacardii

Spores 25-40 x 8-10 u ..................... [Pleurotrema inspersum]

aurotrema anacardii (Vain.) R. C. Harris comb. nov.

 

Arthgpyrenia (Acrocordia) anacardii Vain., Hedwigia 38(Beib1.):

 

53). 1899. Holotype: Guadeloupe, Basse-Terre, Duss 498 (TUR).

Thallus whitish or cream color. Trentepohlia present.

 

.Ascocarps flattened, elliptical, ovate or circular in outline, long
53 oriented toward the ostiole or perpendicular to it, 0.4-0.7 mm
3' and usually 0.05-0.1 mm less in width. Asci cylindrical, 110-170
)-15 u. Spores uniseriate, rarely biseriate, narrowly ovate, ends
frl rather pointed; spore wall of older spores granular ornamented;
5(-30) x 5.5-7.5 u.
I"licroconidia elliptical to narrowly elliptical, 2-4.5 x 1.2-2 0.
HEibitat on bark.
Eilgeurotrema anacardii is rather similar to E. inspersum but is
:Lfiir in all respects. In North America it is found only in southern-
IPlorida.
JILlustrations: Figs. 266-269.
ESIPecimens seen: United States. FLORIDA: Monroe County, Umbrella

: 8.1V.1916 Small 7598 (FH-RIDD, us).

 

 

121

West Indies. CUBA: sine loc., Wright (MSC ex G). PUERTO RICO:

suine loc., Seaver (MICH). ST. LUCIA: Quarter of Soufriere, summit area

cxf'Gros Piton, Imshaug 30198, 30210 (MSC).

Efileurotrema inspersum Mfill. Arg.

Bot. Jahrb. Syst. 6: 389. 1885. Type collection: Cuba, Wright

vrarr, Cub. 123. Isotypes in FH-TUCK 3948 & US.

 

Didymella labiata Vain., Hedwigia 38(Beib1.): (258). 1899.

Pkolxotype: Guadeloupe, Basse-Terre, Duss 503 (TUR).

Ascocarps flattened, mostly ovate in outline, the long axis oriented
toward the ostiole, 0.5-0.6 mm long, 0.1-0.2 mm less in width. Asci
Cl’l;indrical, 180-215 X 15-20 u. Spores uniseriate, narrowly ovate;
SENDITe wall ornamented, especially in older spores; 25-40 x 8-10 0.
Pleurotrema inspersum has not been collected in North America.

I: l'la‘ve verified specimens from Cuba, Guadeloupe and St. Lucia.

Illustrations: Figs. 270-272; Vezda (1968), fig. 32.

4. PYRENOCOLLEMA Reinke

{IaufiJdb. Wiss. Bot. 28: 461. 1895. Holotype: .2. tremelloides Reinke

(== 33. epigloea).

Xanthopyrenia Bachm., Nov. Acta Acad. Caes. Leop.-Carol. German.

 

khail. Cur. 105: 65. 1919. Holotype: Arthopyrenia tichothecioides Arn.
Pseudarthopyrenia Keissl., Revue Bryol. Lichénol. 8: 32. 1935.

HQlotype: Verrucaria caesia Nyl.

Thallus often well developed and epilithic, subfoliose in one species,

Or poorly developed and mostly within the substrate. Phycobionts

122

include a variety of blue-green algae, including Hyella, Nostoc and

 

some Gleocapsa-like algae.

Ascocarps usually rather small, 0.2-0.5 mm in diameter. Ascocarp
wall brown, flexible when wet except in _P_. halodytes which has a car-
bonized thalline envelope. Paraphyses rather sparse, branched and
anastomosed, embedded in an abundant gelatinous matrix. Asci mostly
ovate or obovate, occasionally almost cylindrical, with a long flexuous
base especially when young. Spores ovate with truncate ends to narrowly
ovate, upper cell usually broader and shorter, usually constricted at
the septum. Spore wall thin and the spores often collapsing, rarely
with a thin perispore. Spore contents often weakly orangish in IKI.

Microconidia elliptical to oblong.

Habitat on rock, mostly calcareous, and barnacles; marine, fresh

water aquatics or in other moist situations.

Eyrenocollema is a most remarkable genus, especially for the wide

 

variety of blue-green phycobionts and its preference for wet habitats.
RGinJce, Bachmann and Keissler erected their genera mainly on the basis
Of the type of algal associate but the combined genus is also distinctive
on rmycological grounds. The spore shape, thin spore wall, IKI+ orangish
Spore contents and the rather long, often flexuous ascus base are diag-
nostic for the genus. With the exception of P. halodytes, I am acquain-
ted with the species of the genus from only a handful of specimens.

This has made interpretation difficult. firenocollema epiiloea,

W and prospersella are virtually indistinguishable on the basis

 

of mYczological characters and are separated on the basis of thallus,
phycobiont and habitat characters. Swinscow (1965b) has taken a

dlfferent point of view and included collections with differing

123

phycobionts in P, caesia. Problems of this sort point up how little is

really known of the biology of these organisms. Before any final
taxonomy can be attempted more ecological data must be obtained and,

if possible, isolation of the phycobionts should be attempted.

it is

Of the previously described species included in Pyrenocollema,

clear from the descriptions that _P_. minutulum (Born.) Puym. and

_P_. neozelandicum Zahlbr. are discolichens. Pyrenocollema aquensis

Crozals & Dughi would seem to be excluded from the genus due to its
lack of paraphyses, lack of long ascus base and by the pointed spores.
Also in addition to the species included in the key, it seems likely
from Swinscow's (1967a, 1970) descriptions that Arthopyrenia stronianen-

Sis and A. monensis (Wheld.) Zahlbr. will prove to be members of this

genus.

1. Marine, in tidal or spray Zone .............................. 2
1- Fresh water aquatics, semi-aquatics or on rock in moist

3

places ......................................................

2 - Thallus thin, inconspicuous; algal cells small, up

to 12 p, in diameter (Swinscow); spores 12-20( -23)

X 5-8(-10) p. ..................................... P, halodytes
2 - ‘Ihallus thick, gelatinous when wet, conspicuous;

algal cells large, up to 25 p. in diameter; spores

14-18 X 7'9 11 0.00.00.00.0000000000.00.00.00.0.0.0 20 1111511811911

3. . . .
PhYcobiont a subfoliose spec1es of Nostoc; thallus

Similar to that of Collema; spores 18-21 X 8-9.5 11 .....

'0o................................................... [3. epigloea]

124

:3. Phycobiont not Nostoc; thallus crustose .................... 4

4. Algae blue-green in COlor .............................O S
4. Algae yellow-brown to brown due to colored sheath

(Gleocaesa-like) ....................................... 6

S. Spores 25-3OXlO-12.5u ............................... [BO caeSia]

5. Spores 17-23X8-llu ............................. B. prospersella

6. Spores with a thin perispore, 20-25 X 5-7 p ..... LE. saxicola]
6. Spores lacking an obvious perispore, (18-)20-27

X8-1lu .........I......................... _P_. tiChOtheCiOideS

,Eygrrenocollema caesia (Nyl.) R. C. Harris comb. nov.

Verrucaria caesia Nyl., Bot. Not. 1853: 162. 1853. Arthopyrenia

 

Casessia (Nyl.) Zahlbr., Cat. Lich. Univ. 1: 330. 1922. Type collection
not seen.

Thallus grayish to brownish, continuous, rimose or of separated
grénfililes. Phycobiont with blue-green colored cells in groups surrounded
by a sheath.

ARscocarps mostly emergent. Asci obovate. Spores 25-30 x 10-12.5 p.

Iiabitat on limestone.

Ifly concept of this species is based on a single specimen plus the
description by Swinscow (1965b). It differs from _P. prospersella in
having larger spores. However Swinscow places the lower limit of spore
lerkgtfli in P. caesia at 20 p which would overlap P, prospersella. In the

linuifued material I have examined there is no overlap and I recognize two

8 - . . . . .
Peches, at least temporarily. Pyrenocollema caes1a is Similar to

125

E1. t:ichothecioides in spore size but differs in phycobiont.

Pyreno-

col lema caesia has not been found in North America.

Illustrations: Figs. 273-274; Swinscow (1965b), fig. 8.

Specimen seen: France. VAR: Cap Brun, 1.1923 Crozals (US eX MO).

jiyfirreariocollema epigloea (Nyl.) R. C. Harris comb. nov.

Verrucaria epigloea Nyl., Flora 69: 464. 1886. Arthopyrenia
epigloea (Nyl.) Zahlbr., Cat. Lich. Univ. 1: 279. 1921.

gernelloides Reinke, Jahrb. wiss. Bot. 28: 461. 1895.

Pyrenocollema

Type collection:

1953:11:.. Univ. 249, "Supra Nostoc saxa dolomitica incolens inundata

fl‘Llrninis Narenta in Herzegovinia (Lojka)." fide Nyl. Isotype in MICH.

Thallus blackish, subfoliose, very similar to a primitive Collema,

PIEiKDist determined by the Nostoc phycobiont.

.Ascocarps immersed in the thallus. Asci narrowly obovate. Spores

18~21 x 8-9.5 u
IWicroconidia elliptical, c. 3 X 1.2 p.

Pyrenocollema epigloea is known only from the type collection. It
jLss That completely clear whether the last two numbers of Lojka's Licheno-
1:IVISECZaUniversalis (249 & 250) were ever officially distributed. Reinke's
Specimens had only the number without any data (Reinke, 1895). The
ESEDEEClimen in MICH has a handwritten label naming it as Verrucaria

e 0
‘~EEE£1;22§ and giving an abbreviated version of the collection data

CID‘Q‘ted from Nylander above.

The discovery that the mycobiont of g. epigloea was congeneric
Vwitfll that of_§, halodytes and related species was most startling to me.

tune thallus would indeed lead one to place P, epigloea in the

126

Collemataceae. However, if I were presented only with the ascocarp and

its contents 1 would be virtually unable to distinguish them from those

of P- imshaugii or P. prospersella. The microconidia are very similar

but slightly larger in the last two species. Thus I am not at all

hesitant in uniting P. epigloea with the crustose Species.

Keissler (1936-38) suggested that this species was a parasite on
the Nostoc, but in view of the constant association of the other species
Of the genus with blue-green algae and its similarities to other members

Of the Strigulaceae, I feel it is best treated as a true pyrenolichen

with an extraordinary phycobiont. The name arenocollema at first

Seems somewhat inappropriate but since the genus consists of pyreno-

lichens with blue-green phycobionts it is actually quite a descriptive

name,

Illustrations: Figs. 275-278; Reinke (1895), fig. 179.

Ewcollema halodytes (Nyl.) R. C. Harris comb. nov.

Verrucaria halodytes Nyl., Mem. Soc. Sci. Nat. Cherbourg 5: 142.
1875- Arthopyrenia halodytes (Nyl.) Arn., Ber. Bayer. Bot. Ges. 1,

Slip131.: 121. 1891. Type collection not seen.

Verrucaria consequella Nyl., Sert. Lich. Labuan & Singap. 43. 1891.
wpyrenia consequella (Nyl.) Zahlbr., Cat. Lich. Univ. 1: 276. 1921.
H

OlOtype: Massachusetts, Weymouth, 1890 Willey (H-NYL 984).

For additional synonymy see Swinscow (1965a).

Thallus usually immersed when on limestone or barnacles, epilithic,

thin, brown and shiny when on non-calcareous rock. Phycobiont Hyella

(Swinscow, 1965a) .

127

Ascocarps emergent or immersed. Asci variable, slightly ovate,
rueeaxrly cylindrical or slightly obovate. Spores 12-20(—23) X 5-8(-10)ll.

Microconidia elliptical, 2-3 X 1.2-2 0.

Habitat on rocks, calcareous or not and barnacles in the inter-
tidal zone.

This species is most easily recognized by its habitat. Pyreno-
collema imshaugii is also marine but has a much thicker thallus with
Iléllfggeer algae and slightly larger microconidia.

This is certainly the most common and widespread member of the
EJEBIIIJSS. Not unexpectedly there is some variation. The majority of the
<-‘-<>:Llections from California have larger spores than those from the East
C2C>Ei£3tz or the majority of European specimens. The range is, however,
GEEsteantially continuous and some European specimens have equally large
ESEDCIINES. Thus it seems unprofitable to give taxonomic recognition to
1:}163 (Californian population.

The distribution of E. halodytes is essentially cosmopolitan,
IDEEerhg absent only on tropical shores (Santesson, 1939). Most of the
636155ternrecords are contained in a thesis by R. M. Taylor (Michigan
Esilaitue University) on the ecology of marine lichens and will not be
repeated here.

Illustrations: Figs. 279-283; Swinscow (1965a), figs. 1-2.
Distribution: Fig. 421.

Exsiccati examined: Krypt. Vind. 469 (US), 469b (FH, US); Magn.

242 (MSC); Mig. 24 (MICH, MSC), 74 (MICH, MSC); ve’zda Sel. 130 (MSC).
Specimens seen: United States. CALIFORNIA: Humboldt County,
Thiinidad,‘2§£E§ (MSC); Los Angeles County, Santa Catalina Island, ESEEE

& Santesson L-42155 (FH); Monterey County, Carmel, Pt. Lobos, 10.1X.1932

128

Bonar (FH, MICH, US, US ex MO); San Francisco County, San Francisco,
Cliff House, 11.111.1935 £33231; (FH, MICH), Land's End, 1.1921 Gardener
(FH, MICH, US, US ex M0), 21.VIII.1932 Eggs—r (FH, MICH, MSC, US).
NIAIIVE: County unknown, Wood. Island near Brunswick, 28.VIII.1939
Degelius (US); Hancock County, Mt. Desert Island, 31.VII.1922 wig
(US ) ; Sagadahoc County Cundys Harbor, 28.VIII.1939 Degelius (US).
MASSACHUSETTS: Essex County, Magnolia, VIII.1903 [Farlow] (FH, MICH,
Msc) ; Norfolk County, Weymouth, Willey (H-NYL, MICH). RHODE ISLAND:
NeWport County, S of Newport, Wetmore 12317 (CAN, MIN). WASHINGTON:

Clal lam County, Crescent Beach, Pike 917 (OSC).

‘EXrenocollema imshaugii R. C. Harris sp. nov.

_P_. halodyti sat similis quoad characteres fungi sed differt thallo
CraSSiore cum cellulis algarum majoribus, usque 25 p. diam. et micro-
coniciiis parum majoribus, 3-4 X 1.2-2 1)..

Holotype: California, Mendocino County, Point Arena, on coastal

rocks near lighthouse, Imshalg 17678 (MSC). Isotype in H-

 

CEhallus thick, c. 0.1-0.2 mm, olivaceous to dark brown, shiny,
SCXHEEVVFIat translucent even when dry, more so when wet. Phycobiont a
blue“(green alga, blue gray in color with cells of varying size up to
25 1* jLn diameter, cells weakly arranged into vertical rows.

Ikscocarps subglobose, immersed, becoming emergent, black, 0.2-0.3 mm
ill Ciji~EEtmeter. Ascocarp wall carbonized below. Asci narrowly elliptical
.to I—:‘-'531.‘::‘row1y obovate, c. 80-85 X 17-22 u. Spores 14-18 X 7-9 p.

bflicroconidia elliptical, 3-4 X 1.2-2 0.

liabitat on non-calcareous rock in the intertidal zone? (barnacles

on the same piece of rock).

129

I had at first thought 2, imshaugii to be no more than an odd form
(>1? _2. halodytes but the distinctive thallus, phycobiont not Hyella and
1:?163 slight difference in microconidial size seem to warrent recognition
eat: 'the species level. The species is known only from the type locality.

Illustrations: Figs. 284-288.

Specimens seen: United States. CALIFORNIA: Mendocino County,

Pkojirrt Arena, Imshaug 17677 (MSC), 17678 (H, MSC).

 

_Efifxreeriocollema prospersella (Nyl.) R. C. Harris comb. nov.

Verrucaria prospersella Nyl., Lich. Japon. 108. 1890. Arthopyrenia

 

 

EE?<>£3;>erse11a (Nyl.) Zahlbr., Cat. Lich. Univ. 1: 288. 1921. Holotype:

IlJllixiois, Cook County, 10 mi from Chicago, Calkins (H-NYL 991).

Thallus gray, continuous to rimose, epilithic. Phycobiont with
CEiliLss blue green in color, in small groups but without an obvious sheath.

.Ascocarps globos, 0.2-0.25 mm in diameter. Asci slightly ovate to
elliptical. Spores 17-23 x 8-11 ,1.

Inicroconidia elliptical, 3.5-4 x 1.5-2 u.

Edabitat on calcareous rocks, possibly more or less aquatic.

jEyrenocollema prospersella is externally similar to P. caesia but

 

differs in havin smaller spores. It is known from a Belgian collectiOn
ill Eicidition to the type locality.

Itllustrations: Figs. 289-292.

£3pecimens seen: United States. ILLINOIS: Cook County, Calkins

(F
Ei’ DfiICH, MSC), 10 mi from Chicago, Calkins (H-NYL 991).

130

pyrenocollema saxicola (Massal.) R. C. Harris comb. nov.

Arthopyrenia saxicola Massal., Symmict. Lich. 107. 1855.

 

Leiophloea saxicola (Massal.) Riedl, Sydowia 23: 234. 1971. Type

 

col lection not seen.

Thallus mostly epilithic, brown to blackish, appearing speckled
due to uneven distribution of algae. Phycobiont yellow brown in color,
Gleocapsa-like.

Ascocarps globose, partially immersed in rock, c. 0.2 mm in dia-
meter. Spores narrowly elliptic, narrowly ovate, elliptic or ovate,
With rather blunt ends, 20-25 X 5-7 u, with a thin perispore.

This species is distinctive in the Gleocapsa-like phycobiont and
in the spores which are longer in proportion to width than is usual in
the genus. I have seen specimens from England and Switzerland but not
from North America.

Illustrations: Figs . 29 3-295 .

wollema tichothecioides (Arn.) R. C. Harris comb. nov.

 

firthopyrenia tichothecioides Arn., Flora 52: 268, pl. 8, f. 6.

 

1869 - Xanthopyrenia tichothecioides (Arn.) Bacm. , Nova Acta Acad. Caes.

 

L
er- ~Carol. German. Nat. Cur. 105: 65. 1919. Type collection not seen.

Thallus yellow brown to dark brown, epilithic, in drier situations
Cohsjesting of dispersed groups of algae, rimose to continuous and
some"What gelatinous in moister or aquatic habitats. Phycobiont yellow
brown in color, Gleocapsa-like.

Ascocarps globose, immersed in rock or superficial, 0.2-0.25 mm in

d.
lameter. Asci narrowly ovate, narrowly obovate to ovate, elliptic or

131

obovate. Spores (18-)20-27 X 8-11 11.

Microconidia elliptic to oblong, 3-5 X 1.2-1.5 11.

Habitat on calcareous rock in moist to aquatic situations.

Pyrenocollema tichothecioides is distinguished from B. caesia by
the associated alga, from the rest of the genus by its larger spores.
The aspect of the thallus is markedly different between drier and wetter
habitats. Possibly more than one species is involved. The aquatic
forms may prove to be referable to Arthgyrenia strontianenSis Swinsc.
The European specimens examined (Yugoslavia and France) seem to be
from drier habitats. One of the Michigan collections (1080-A) is from
a lake Shore (but not aquatic, associated with Verrucaria muralis Ach. ).
The other Michigan collections were aquatic.

Illustrations: Figs. 296-300.

Exsiccati examined: Krypt. Vind. 1356 (US).

Specimens seen: United States. MICHIGAN: Mackinac County, shore
Of Lake Huron W of Hessel, Harris 1080-A (MSC); Menominee County, N of
CSdar River, Harris 731-C, 749 (MSC).

Mexico. State unknown: Villas Cardinas, XI. 1910 Orcutt 166 (FH).

5, STRIGULA Fr.
S
YSt- Myc. 2: 535. 1823. Lectotype (Santesson, 1952): _S_. smaragdula Fr.
(= £- elegans (Fee) Miill. Arg.

geisleria Nitschke it}; Rabenh., Lich. Eur. 574. 1861. Holotype:

 

G.
‘ ELChnogonioides Nitschke _ig Rabenh.

Baciboskiella v. HOhn., Sitzungsber. Kaiserl. Akad. Wiss., Math.-

 

N
atklt‘wiss. C1., Abt. 1. ser. 1, 118: 1176. 1909. Holotype: Clypeolum

W Racib. (= 3. janeirensis (M1111. Arg.) Sant.).

132

Geisleriomyces Cif. & Tomas., 1st. Bot. Reale Univ. Reale Lab.

(Ixrjlttog. Pavia Atti, ser. 5, 10: 31, 59. 1953 nom. illeg. Holotype:

 

'53- _sychnogonioides Cif. & Tomas.
Sagediomyces Cif. & Tomas., Ist. Bot. Reale Univ. Reale Lab. Crittog.
I?ea\rjxa.Atti, ser. 5, 10: 30, 57. 1953. Holotype: S, affinis Cif. & Tomas.
Strigulomyces Cif. & Tomas., Ist. Bot. Reale Univ. Reale Lab.

(Zr?j_t:tog. Pavia Atti, ser. 5, 10: 34, 61. 1953. nom. illeg. Holotype:

 

é- elegans Cif. & Tomas.

See Santesson (1952) for additional synonymy.

Illailfilus whitish, grayish or greenish to dark green or brown, rarely
FNJIlczizate, endophloeodal, epiphloeodal, epiphyllous and subcuticular or
enCiOlithic; black hypothallus present in a few species. Phycobiont

£2532£2j:epohlia or Cephaleuros.

 

Ascocarps mostly hemispherical, less commonly subglobose, immersed
(Jr' SSLlperficial; outer layer of the wall dark brown to blackish, often
15“3L:fi4ng below or occasionally when the ascocarps are immersed the wall
Ina3? IDe colorless except for the exposed tip. Hymenial gelatin IKI-.
Pa“railpl'lyses thread-like, unbranched or occasionally dichotomously
brkar1Cihed above or rarely somewhat branched and anastomosed above. Asci
Cyrlelt‘ldrical to narrowly elliptical, rarely narrowly ovate; often with a
Sfiua]_3_ ocular chamber. Spores commonly uniseriate, occasionally sub-
biTSEEIiiate to biseriate, rarely irregularly arranged, mostly narrowly
Ointlea, less commonly ovate or elliptical, commonly 2-ce11ed but some
SpEECljoes 4-8-ce11ed and one submuriform; cells in 2-celled species often
EHJiSJPItly unequal with the lower cell shorter and narrower; spore wall

r1
0t; (Drnamented; perispore not distinguishable.

133

Microconidia narrowly elliptical to elliptical.

Macroconidia common, present in most collections, narrowly ellipti-
cal or rarely filiform, usually with as many cells as the ascospores,
rarely more.

The realization that Strigula encompasses many non-foliicolous
species was a very slow process for me, but the conclusion seems ines-
capable. The foliicolous and non-foliicolous species share a number of
unique characters, septate macroconidia, almost unbranched paraphyses,
Spores separating into part spores, punctation of the thallus and
Presence of a black hypothallus. Also the general shape of the asco-
Carps, asci, spores and microconidia is very similar. Ve'zda (1_n_ 1331;.)
has indicated some reluctance to merge the non-foliicolous species with
the foliicolous ones since the latter have a distinctive subcuticular

thallus. However, the Cephaleuros phycobiont is also subcuticular in

 

the free living state (Printz, 1940). Therefore, in my opinion, the
SubCIL‘lticular thallus of the foliicolous species of Strigula is merely a

resUCLt of the association with Cephaleuros and should not be considered

 

when delimiting genera. The combined taxon seems to me to be a very
hatL—‘Ji‘al one adapted to a wide variety of substrates and climates.
W seems to be mainly subtropical or tropical although a number
of species occur in temperate areas of the Northern Hemisphere. I have
not S:een any specimens of non-foliicolous species from the Southern
Hemisphere.

1..
CEnrowing on leaves (modified from Santesson, 1952) . . . . . . . . . . . 2

1. . .
GrOWIng on bark, wood, mosses, 5011 or rock ................. 4

134

2. Spores 4-ce11ed, rarely 2-ce11ed, fusiform to almost
bacillar, (12-)15-24 x 3.5-6 u; ascocarps 0.1-0.2 mm
in diarneter ........C............................ E. complanata

2. Spores 2-Celled, rarely 4-Celled ......O................ 3

Spores elliptical, breaking into two part spores,

8-10(-12) X (2-)2.5-3 u; ascocarps 0.4-0.6 mm in

diameter .............................................. S, nitidula
Spores fusiform, not breaking into part spores, l4-22(-24)

X 4-5.5 p; ascocarps 0.25-0.4 mm in diameter ........... S, elegans

4. Growing on rock or soil ................................ 5

4. Growing on bark, wood or mosses on bark ................ 8

Spores 4-Celled ............................................ 6

Spores 2-Celled ............................................ 7

6. Spores 15-21 X 4.5-7 u; on limestone or mortar ..... S. affinis
6. Spores larger, 18-30(-35) X 7-10 u; on soil ........

.......................................... EE. SYChnggoniOideS]

 

Spores 12-15 X 4.5-5.5 u ............................ S, viridiseda

 

Spores larger, 14-20 X 4.5-6 u ........................ S, wilsonii

8. Spores 4-8-C8118d 0r Smeurifom 000.000.000.00000000000 9

8. spores 2-Celled ........................................ ll

Spores 4-C€lled, 15-21 X 405-7 1]. 00000000000.00.00...000 é. affinis

Spores 6-8-celled or submuriform ........................... 10

 

....
..Cx

135

10. Spores often submuriform, 20-27 x 6-7.5 u; growing

on bark ...............O..................... -S_. Sllbmurifomis

 

10. Spores never submuriform, 24-42 X 5-7.5 u; on bryo-

phytes, less commonly directly on bark ........ S, stigmatella

 

ll, Ascocarp compound, with 2-3 chambers sharing a common
OStiOle; Spores small, 7-lO(-12) X 3‘405 n 0000000000 §f connivens

Ill. Ascocarp simple, with only a single chamber ............... l2

l2. Thallus with small white dots; black hypothallus

present; spores 17-22 X 5-6 p ............... S, hypothallina

 

12. Thallus not dotted; hypothallus lacking .............. 13

3.33- Asci narrowly ovate; spores usually breaking into two
part Spores, 21'32 X 4-505('7) u .0000000000000000000 LE0 tailori]
1 3.. Asci cylindrical to narrowly elliptical; spores not

breaking into part spores ................................. 14

14. Spores small, 9-11(-13) X 2.5-4 0, ends usually
rounded with upper cell often broader than lower ... S, phaea
l4. Spores larger, 12-25 X 4-5.5 u, usually with rather

pointed ends, cells not much different in width ...... 15

1J3. Spores 12-17 X 405.5 u 000.0...0.00.00.00.0000000... é: ViridiSEda

1&3. Spores 16-25 X 4-5.5 p .............................. S, americana

Eigifliflglgflg£££§£§ (Massal.) R. C. Harris comb. nov.

Sagedia affinis Massal., Memor. Lichenogr. 138, f. 169. 1853.

 

9% affinis (Massal.) Zahlbr., Oesterr. Bot. z. 51: 277. 1901.

136

,Arthopyrenia affinis (Massal.) Boist., Nouv. Flore Lich. 2: 278. 1903.

 

Type collection not seen.

Geisleria jamesii Swinsc., Lichenologist 3: 420. 1967. Holotype:

 

ISngland, Dorset, Brownsea Island, X1.1964 James (BM).

Ascocarps hemispherical to subglobose, 0.1-0.4 mm in diameter;
aascocarp wall sometimes extended outward above, often lacking below.
lksci narrowly elliptical to cylindrical, 60-100 x 10-15 0. Spores sub-
kxiseriate to biseriate, narrowly ovate to narrowly elliptical, 4-ce11ed,
sslxightly constricted at the septa, 15-21 X 4.5-7 u.

Microconidia narrowly elliptical to elliptical, 2.5-4 x 1-1.7 u.

Macroconidia narrowly elliptical, 4-ce11ed, 12-17(-20) X 3-5 0.

Habitat on bark or rarely on calcareous rock or mortar.

In 1973 I carelessly overlooked Boistel's earlier transfer of this

Species to Arthopyrenia and made an unnecessary transfer. I can see no

 

differences between Geisleria jamesii and S, affinis except substrate.

 

jljfii collection from Haiti is on limestone and has slightly broader
Spores and longer macroconidia than usual.

Illustrations: Figs. 301-308; Harris (1973), fig. 1; Swinscow
(1962), fig. 13, (1967b), figs. 2-4.

Exsiccati examined: Hepp 458 (FH); Mass. 350 (FH); Rab. 561 (MICH).

Specimens seen: Canada. ONTARIO: Carleton County, Ottawa,
12-IV.1895 Macoun (FH).

United States. LOUISIANA: E. Baton Rouge Parish, Baton Rouge,
W 9056-A (MSC).

West Indies. HAITI: Department de 1'Ouest, N of Foret des Pins,

Wetmore 2903 (MSC).

137

Strigula americana R. C. Harris nom. nov.

Arthopyrenia tenuis R. C. Harris, Michigan Bot. 12: 16. 1973.

 

(non Strigula tenuis Mfill. Arg., Flora 68: 344. 1885.). Holotype:

ZIowa, Fayette County, 1896 Fink (MSC). Isotypes in FH, MICH, MIN & US.

Ascocarps hemispherical or somewhat Conical, 0.3-0.5 mm in diameter,
Eiscocarp wall thinner or lacking below. Asci cylindrical, 70-135 X
£3-12 p. Spores uniseriate to biseriate, narrowly ovate with pointed
enuds, occasionally slightly bent, 2-celled, slightly constricted at the
septum, 16-25 X 4.5-5.5 11.

Microconidia narrowly elliptical to elliptical, 3-5 X 1.2-2 u.

Macroconidia narrowly elliptical, 2-ce11ed, 12-20 X 3-4.5 u.

Habitat on bark.

Strigula americana is closely related t0.§° viridiseda but differs
54r1 having larger spores. However in the southeastern United States
'“flieare their ranges overlap a number of collections have spores inter-
meEdiate in size. They have been assigned to S, americana. I previously
thOnght this species to have a rather northern distribution but it turns
cnit; to be fairly common in Florida and Louisiana.

Illustrations: Figs. 309-314; Harris (1973), figs. 13-15.
Distribution: Fig. 422.

Additional specimens seen (not cited in Harris, 1973): United
E}tfttes. ALABAMA: Cleburne County, N of Heflin, Harris 1403 (MSC).
IEL“3RIDA: Dade County, Coconut Grove, 1898 Thaxter 275 (FH), g§g_(MICH);

Duval County, Jacksonville, Calkins (MICH); Jackson County, Florida
(hiverns St. Park, Harris 1597 (MSC); Liberty County, Apalachicola Nat.

F0rest, Whitehead Lake Campground, Harris 1519 (MSC); Marion County,

 

138

Eton Creek at Fla. Hwy. 314, Harris 2405 (MSC); Seminole County,

 

Sanford, 1.1924 Rapp 177 (MICH). LOUISIANA: E. Baton Rouge Parish,

jBaton Rouge, Tucker 9035 (LSU); St. Martin Parish, Duchamps, 26.IX.1894

 

Langlois (us), John Durand's, 2.1.1894 Langlois 913 (US). NEW JERSEY:
IBergen County, Closter, 1877 Austin 747 (FH-TUCK 4049), 752 (FH-TUCK
41048). SOUTH CAROLINA: Berkeley County, Guilliard Lake Campground,

Iiarris 3193-A (MSC).

Eitrigula complanata (Fee) Mont.
Hist. Cuba [Sagra] Bot. P1. Cell. 140. 1842. Phyllocharis

czcxnplanata Fée, Essai Crypt. Ecorc. Off. XCIV, XCIX, pl. 2, f. 3. 1824.

 

T‘yPe collection not seen.

See Santesson (1952) for synonymy and description.

Strigula complanata in the United States is often sterile or pro-

 

Citu:ing only pycnidia. It is easily recognized by its 4-celled spores or
by the filifrom conidia.

Illustrations: Fig. 315. Distribution: Fig. 423.

Specimens seen (not cited by Santesson, 1952): United States.
FLORIDA: Jackson County, Florida Caverns St. Park, Harris 1598 (MSC).
LO'UIISIANA: E. Baton Rouge Parish, Baton Rouge, Tucker 8841, 9063 (MSC);

Livingston parish, 23.IX.l968 Exner (Msc).

Eitfliigglg connivens R. C. Harris sp. nov.
Species generis unica ascocarpis compositis, 2-3 loculis cum
OStiolo unico et sporis 7-10(-12) X 3-4.5 p.

Holotype: Florida, Dade County, Cutler, 1898 Thaxter l2 (MSC).

139

Thallus greenish gray, well developed, endophloeodal.

Ascocarps compound, usually with two chambers, occasionally three,
.rarely simple, 0.3-0.4 mm long, somewhat less in width; chambers con-
Iaected above, sharing a single ostiole; ascocarp wall thinner or lacking
loelow. Asci narrowly elliptical, 45-50 x 7-8 p. Spores subbiseriate to
kaiseriate, narrowly ovate, 2-celled, slightly constricted at the septum,
'7-10(-l2) X 3-4.5 u.

Microconidia narrowly elliptical to elliptical, 2.5-4 x 1.5-2 u.

Macroconidia narrowly elliptical, 2-celled, 6.5-7.5 X 2-3 p.

Habitat on bark.

This is the only species known from the Strigulaceae with astro-
't11ralioid ascocarp structure. In all other respects it is a typical
53tzrigula. Normally I would not describe a species on the basis of so
3_j:tt1e material, but I feel it is necessary to round out the circum-

S Cription of Striggla.
Illustrations: Figs. 316-320.
Specimens seen: United States. FLORIDA: County unknown, Snapper

Hammock, 11.XII.1919 Britton & Britton 806 (FH-RIDD); Dade County,

 

 

 

Cutler, 1898 'Ihaxter 17a (MICH), lg (Msc).

SiEEELgula elegans (Fée) Mfill. Arg.

Linnaea 43: 41. 1880. Phyllocharis elegans Fee, Essai Crypt.

 

ECOrc. Off. XCIV, (3, pl. 2, f. 7. 1824. Type collection not seen.

See Santesson (1952) for synonmy and description.

Strigula elegans is distinguished from S. complanata by its 2-ce11ed

 

SPores and from S. nitidula by larger spores which do not break up into

140

Ipart spores. Many of the specimens cited below were determined by
ESantesson after the publication of his work on foliicolous lichens.
Illustrations: Fig. 321-322; Vezda (1968), fig. 29.
IDistribution: Fig. 424.
Specimens seen (not cited in Santesson, 1952): United States.
IPLORIDA: Alachua County, Gainesville, 111.1924 Burger (MICH), 21.X.1907
Piampton (MICH), 20.VI.1922 O'Byrne (MICH); Dade County, Coconut Grove,

I.1898 Thaxter .111 (MICH, MSC), 397, 406, 409, 427 (MICH), Cutler,

 

I:(.1897 Thaxter 414 (MICH); Jackson County, Florida Caverns St. Park,

 

risirris 1600 (MSC); Lake County, Eustis, Thaxter (MICH, MSC); Marion
Clcnunty, Ocala, X.1897 Thaxter (MICH, MSC), Ocala Nat. Forest, Hughes

Jisiland, Harris 2000 (MSC); Orange County, Plymouth, 1914 Grossenbacker

 

(MICH); Palm Beach County, Palm Beach, Thaxter (MICH); Putnam County,

(Ikilawaha River at Fla. Hwy. 19, Harris 2260 (MSC); Volusia County,

 

Daytona [= Daytona Beach], 1898 Thaxter 422 (MICH). LOUISIANA:

 

VV- Feliciana Parish, TUnica bulls, Koch 7743 (MSC); Livingston Parish,

DeI’lham Springs, 30.V.1968 Evans (MICH, MSC).

Siii-iiigglghypothallina R. C. Harris sp. nov.

 

Striggla thallo nitido, albo-punctato, infra cum hypothallo nigro
et Sporis bicellularis, 17-22 X 5-6 11.
Holotype: South Carolina, Berkeley County, Francis Marion Nat.

FC”Test, Guilliard Lake Campground, Harris 3201-A (MSC).

 

Thallus light brown, shiny, with scattered white punctae, underlain
'bY a black hypothallus which is visible at the margins of the thallus.

Ascocarps flattened to hemispherical, immersed, 0.5-0.8 mm in

141

diameter, 0.2 mm in height; ascocarp wall sometimes indistinguishable

from the hypothallus, lacking below. Asci cylindrical, 90-120 X 9-10 u.

.Spores uniseriate, narrowly ovate, with rather pointed ends, 2-celled,
fslightly constricted at the septum, 17-22 x 5-6 p.

Habitat on bark (Ilex).

Although this species is known from only a single collection, I
Krenture to describe it since it helps to link the non-foliicolous and
fioliicolous species and to reinforce the concept that they constitute

51 single genus. Strigula nemathora Mont. has a similarly punctate

tiiadlus and S, subtilissima (Fee) Mfill. Arg., S, maculata (Cooke & Massee)

Sant. and S. melanobapha (Kremp.) Sant. have a well developed hypothallus.

 

Illustrations: Figs. 323-325.

Strigula nitidula Mont.
Hist. Cuba [Sagra] Bot. Pl. Cell. 139. 1842. Type collection not

ESExan. See Santesson (1952) for synonymy and description.

This is the only foliicolous Strigula known from the United States

'lr1 Vfliich the spores separate into two part spores. The species is new

tO'the United States flora and the single collection is on Magnolia

leaVes.
Illustrations: Figs. 326-327; vezda (1968), fig. 28.
Seminole County, Oviedo,

Specimen seen: United States. FLORIDA:

XI- 1922 Rapp (MICH).

mm (Ach.) R. C. Harris comb. nov.
Porina phaea (Ach.)

Verrucaria phaea Ach., Synops. Lich. 88. 1814.

142

jMfill. Arg., Flora 68: 261. 1885. Holotype: "India Occid. ad corticem
Cfinchonae?" (H-ACH). Isotype in S.

Porina cineriseda Mfill. Arg., Bot. Jahrb. Syst. 6: 402. 1885.

 

‘Verrucaria cineriseda Nyl., Flora 59: 364. 1876. nom. nud. Holotype:

 

(Suba, wright Verr. Cub. 10 (G).

Pyrenula virella Merr., Lich. Exs., ser. 1, 289. s. d. nom. nud.,

 

Ikcrocordia virella Merr., Lich. Exs., ser. 2, 5. 1925. nom. nud.
lkrthopyrenia virella (Merr.) Zahlbr., Krypt. Exs. Vindob. fasc. 32, 3141.

1933. norm. nud.

Thallus light green to dark green or dark olive or brown, at least
er1 part epiphloeodal.

Ascocarps hemispherical or conical, 0.2-0.3 mm in diameter; ascocarp
VVEill usually thinner or lacking below. Asci narrowly elliptical to
Ciauctyloid, 40-55(-65) X 5-8 p. Spores uniseriate to biseriate, narrowly
<3V13te or narrowly elliptical, 2-ce11ed, slightly constricted at the
septum, 9-11(-13) X 2.5-4 u.

Microconidia elliptical 2.5-4.5 X 1.2-2 u.

Macroconidia narrowly elliptical, 2-ce11ed, 6.5-10 x 2-3.5 u.

Habitat on bark.

This species can often be recognized by its dark colored, usually
e39i$fllloeodal thallus. Its small spores are also distinctive.

Illustrations: Figs. 328-336. Distribution: Fig. 425.

Exsiccati examined: Krypt. Vind. 3141 (US); Merr. I: 289 (US);
Merr. II: 5 (MICH, MSC, US, US ex MO).

Specimens seen: United States. FLORIDA: sine loc., Calkins (MSC);

DuVal County, Jacksonville, Calkins (US); Putnam County, Oklawaha River

at Fla. Hwy. 19, Harris 2217-D (MSC); Seminole County, Sanford, X.1907

 

143
Rapp (MICH, MSC, US, US ex MO), 3.1v.1914 Rapp (FH), XII.1914 Rapp_g§

(FH), 111.1916 Egpp_(MICH, US), V.l927 Egpp_(FH), Selman & Ridge (US).
GEORGIA: McIntosh County, Darien, Ravenel 647 (FH-TUCK 4049).

West Indies. BERMUDA: Somerville, 17.1.1916 Hervey (MICH).
JAMAICA: Parish of Trelawny, Barbecue Bottom, Imshaug 16064 (MSC).

PUERTO RICO: San Juan District, Rio Piedras, 18.1.1916 Fink 2171

(FH, MICH).

£Strigula stigmatella (Ach.) R. C. Harris comb. nov.

Lichen stigmatellus Ach., Lich. Suec. Prodr. 15. 1798. Lectotype:

Germany , Persoon (H-ACH) .

Verrucaria cinerea Pers., Ann. Bot. (Usteri) 7: 28, pl. 3, f. 6A.

3.794. nom. illeg. (non Humbolt, Flora Friburg. Specim. 44. 1793).

IEsotype? in H-ACH.
Sagedia faginea Schaer., Enum. Crit. Lich. Europ. 208. 1850.
liggrina faginea (Schaer.) Arn., Flora 68: 166. 1885. Arthopyrenia

faginea (Schaer.) Swinsc., Lichenologist 3: 72. 1965. Type collection

not seen.

Thallus whitish to green, endophloeodal or epiphloeodal.
Ascocarps subglobose to hemispherical, immersed or emergent, 0.2-

C3-44 mm in diameter; ascocarp wall brown above or almost entirely color-

16ass in immersed ascocarps. Asci cylindrical, 80-110 X 13-15 u. Spores

biseriate or subbiseriate, narrowly ovate with pointed ends, 8-celled,
$51ightly constricted at the septa, especially the middle one, 24-42 x

5‘7.5 u.

Macroconidia narrowly elliptical, 23-30 x 3.5-5 u.

144

Habitat on bark and mosses, in Michigan usually on Thuja.
Henry Imshaug reports that he could find no Swedish collection of

Lichen stigmatellus in Helsinki. However, specimens from Persoon and

 

Schrader were present. Both were cited in the original description.

Schrader's specimen apparently bears only macroconidia, while Persoon's
has both ascospores and macroconidia. Therefore, Persoon's specimen is
designated as the lectotype. Material of this species in the Acharian

collection at the British Museum (284-287) is a mixture of S. stigmatella

 

(macroconidia only) and Pyrenula coryli.

 

Strigula stigmatella is recognized by its 8-ce11ed spores larger

 

'than those of S, submuriformis with which it could be confused. The

 

species is apparently rather common in Michigan IEEiE swamps.
Illustrations: Figs. 337-341; Harris (1973), fig. 5; Swinscow
(1962), fig. 13; Ve‘s’zda (1968), fig. 27.
Exsiccati examined: Hepp 464, 708 (FH); Krypt. Vind. 180 (FH, US);
Inojk. Univ. 199 (FH-TUCK 4074); Rab. 579 (FH), 623 (MICH).
Additional specimens seen (not cited in Harris, 1973): United

531zates. MICHIGAN: Baraga County, NE of Sidnaw, Harris 7828, 7830, 7831,

 

LZEEQQ, 7977, 7978 (MSC); Cheboygan County, E of Pine Grove campground,

 

firms 7810 (MSC); Oscoda County, S of Mio, Harris 8289, 8364, 8371

 

(Msc).

5§Jgrigula submuriformis (R. C. Harris) R. C. Harris comb. nov.

 

Arthopyrenia submuriformis R. C. Harris, Michigan Bot. 12: 15.

 

1~973. Holotype: Pennsylvania, West Chester, VII.1898 Windle (MSC).
Isotypes in CAN, MICH & US as Cum. I: 299; in FH, FH-RIDD, MICH & MSC

as Cum. II: 250.

145
See Harris (1973) for description and discussion. Strigula

submuriformis is easily recognized by its 5-7 transversely septate

 

spores, usually with l or 2 cells longitudinally septate. I have

seen no additional material since 1973.

Illustrations: Figs. 342-345; Harris (1973), figs. 11-12.

SErigula sychnogonioides (Nitschke £2 Rabenh.) R. C. Harris comb. nov.
Geisleria Sychnogonioides Nitschke Ag Rabenh., Lich. Eur. 574. 1861.

Type collection: "an Erdwallen des Schiessplatzes bei Mfinster..."

Isotypes in BM & MICH.

See V5zda (1970) for description and discussion. The species is

<distinctive in being terricolous and in having rather large 4-celled

:Spores. I have examined as many specimens as possible in hopes of

:Einding conidia to confirm its position in Striggla. I have not been

aflole to find either micro- or macroconidia, but even so I feel the

Supecies can be included in Striggla on the basis of ascocarp, ascus and

ESpore characters. Strigula Sychnogonioides is not known from North

1anerica.

Illustrations: Figs. 346-348; Swinscow (1967b), figs. 1-2, Vazda

(1968), fig. 2.

Exsiccati examined: Rab. 574 (BM, MICH); V5zda Sel. 577 (US).

£§3§rigula taylori (Carroll 25 Nyl.) R. C. Harris comb. nov.

Verrucaria taylori Carroll 35 Nyl., Expos. Syn. Pyrenoc. 82. 1858.

Mém. Soc. Sci. Nat. Cherbourg 5: 337. 1857. nom. nud. Arthopyrenia

taylori (Carroll 31 Nyl.) Mudd, Manual Brit. Lich. 302. 1861. Porina

EM (Carroll 2g Nyl.) Swinsc., Lichenologist 2: 169. 1963. Type

146
collection?: Ireland, Cork, Dunscombe's Wood, 12.VIII.1856 Carroll.

Isotypes in BM g§_Kew.

See Swinscow (1963) for description and discussion. The asci tend
to be more ovate than in other species of Strigula. The separation of
the spores into two part spores at maturity is found otherwise only in

some foliicolous species. Strigula taylori has not been found in North

 

America.

Illustrations: Figs. 349-353; Swinscow (1963), fig. 2.

:Strigula viridiseda (Nyl.) R. C. Harris comb. nov.

 

Verrucaria viridiseda Nyl., Expos. Syn. Pyrenoc. 55. 1858. Porina

 

\Iiridiseda (Nyl.) Zahlbr., Cat. Lich. Univ. 1: 409. 1922. Holotype:

 

"Guyana gallica, Melinon" (H-NYL 719).

Verrucaria bermudana Tuck. §§_Nyl., Sert. Lich. Trop. 43. 1891.

 

flflielidium bermudanum (Thck. §§_Nyl.) Ridd., Bull. Torrey Bot. Club 43:

 

 

1446. 1916. Holotype: Bermuda, Walsingham, Farlow S; (H-NYL 985).

Isotypes in FH & FH-TUCK.

Thallus whitish to light green, endophloeodal or endolithic.

Ascocarps hemispherical to subglobose, immersed or emergent,
Oc.25-O.5 mm in diameter. Asci cylindrical, 65-105 x 6-10 p. Spores
nu<>st1y uniseriate, less often subbiseriate, narrowly ovate with some-
What pointed ends, 2-celled, slightly constricted at the septum, 12-17 X
4- 5-5 11.

Microconidia elliptical to narrowly elliptical, 2.5-4 X 1.2-2 u.

Macroconidia narrowly elliptical to elliptical, 2-celled, 10-13(-15) X

2.5-4.5 11-

Q0~

147

Habitat on bark, calcareous rock or shell.

The type of Verrucaria bermudana is on rock, but I can find no
cyther characters separating it from S. viridiseda. Strigula viridiseda
:is closely related to and intergrades with S, americana and S, wilsonii
150th of which have larger spores. Collections with spores somewhat
:intermediate in size occur and if on bark have been refered to S.

eunericana, if on rock to S, wilsonii.

Strigula viridiseda occurs in the southeastern United States and is

 

zipparently widespread in the West Indies (Bermuda, Dominican Republic,
LIamaica, Puerto Rico and Trinidad). Thaxter's collections are on an old
(nonch shell, the rest are on bark.

Illustrations: Figs. 354-359. Distribution: Fig. 426.

Specimens seen: United States. ALABAMA: Cleburne County, N of

Pkeflin, Harris 1321, 1336 (MSC). FLORIDA: Dade County, Coconut Grove,

 

>CI];1897 Thaxter 400 (MSC), 402, 414 (MICH); Seminole County, Sanford,

 

4.11.1914 Rapp (FH, us), XII.1915 Rapp (US), 25.v.1921 Rapp 177 (FH,

WHICH, US). LOUISIANA: Parish unknown, near New Orleans, Langlois (MICH).

Eiggigula wilsonii (Ridd.) R. C. Harris comb. nov.

Porina wilsonii Ridd. $g_Brit. & Millsp., Bahama F1. 523. 1920.

 

HOlotype: Cuba, Isle of Pines, Caleta Cocodrilos, N. Britton, Wilson
& Lfig Egg (FH-RIDD).

Porina subprospersella Vain., Mycologia 21: 39. 1929. Type
COllection: Puerto Rico, Yauco, 28°XII°1915.EEEE.1§2§° Isotype in
MICH.

Porina vainii Fink $2 Hedrick, Mycologia 22: 247. 1930. Holotype:

 

Puerto Rico, near Yauco, 28.XII.1915 Fink 1431 (MICH). Isotype in FH.

148

Thallus white to greenish, endolithic.

Ascocarps mostly immersed, subglobose or more less conical; asco-
<:arp wall well developed above and lacking below, or reduced to a small
.ring around the ostiole of deeply immersed ascocarps. Asci cylindrical,
f90-130(-l60) x 7-12 u. Spores uniseriate, narrowly ovate, sometimes
\Nith rather pointed ends, 2-ce11ed, slightly constricted at the septum,
.14-20 X 4.5-6 u.

Microconidia narrowly elliptical to elliptical, 4-5 X 1.2-2 u.

Macroconidia narrowly elliptical to elliptical, 2-ce11ed,
le-l3(-17) X 3-4.5 0.

Habitat on calcareous rock.

Strigula wilsonii is very close to being a saxicolous form of S.

 

amnericana, but the spores are in general shorter and broader in the

 

ffiormer. It also intergrades to some extent with the saxicolous forms

(bf S, viridiseda and occasional collections may be hard to place.

 

fiitrigula wilsonii is apparently very common in Puerto Rico and is also
kniown from Bermuda, Cuba and Trinidad. In the United States it is
Jrestricted to southernmost Florida.
Illustrations: Figs. 360-366. Distribution: Fig. 427.
Specimens seen: United States. FLORIDA: Dade County, Coconut
<3rove, XII.1897 Thaxter (MICH, MSC); Monroe County, Key West, 11.1898

Thaxter (MICH).

D. TRYPETHELIACEAE
Syst. Lich. 17. 1824. Holotype: Trypethelium Spreng. nom. cons.
Astrotheliaceae Zahlbr., Nat. Pflanzenfam. 1(1‘): 72. 1903.

Holotype: Astrothelium Eschw.

149

Cryptotheliaceae W. Wats., New Phytol. 28: 113. 1929. Holotype:
(Zryptothelium Massal.

Hyalophragmiaceae Ras., Acta Bot. Fenn. 33: 9, 23, 74. 1943. 22E:
.inval. descr. germ. Lectotype (Cooke & Hawksworth, 1970):
Pseudopyrenula Mfill. Arg.

Laureraceae Vezda SE Poelt SA_Ahmadjian & Hale, The Lichens 609.

1974. nom. inval. descr. eng.

 

Thallus mostly well developed, semi-endophloeodal, with a cortex-
Ilike upper layer of compacted hyphae and bark cells, sometimes with
eaxtensive deposits of crystals or pigment granules beneath the algal
Jxayer. Phycobiont Trentepohlia or in a few cases non-lichenized.
Ascocarps commonly simple, grouped and immersed in a pseudostroma
Vfliich may be differently colored from the thallus or may contain
Clrystals or pigment granules, sometimes ascocarps solitary, immersed or
:ENJperficial; less commonly with compound ascocarps possessing several
1'Iymenia sharing a common ostiole. Ostioles mostly erect, occasionally
Exzcentric or lateral. Hymenium commonly containing granules of an
Lnflknown substance; hymenial gelatin IKI-. Paraphyses branched and
Einastomosed, often in a regular and reticulate manner. Asci bitunicate
(Morgan-Jones, 1972), often with a broad, shallow ocular chamber.
Spores mostly colorless, very rarely brownish, 4-ce11ed to muriform,
Commonly eight/ascus but occasionally four or two/ascus; mesospore
thickened, rarely remaining unthickened, rarely IKI+ violet; perispore
Present, occasionally rather thick; a few species with granular orna-
mentation.

Microconidia rod-like.

150

Lichen substances: lichexanthone sporadically in most genera and
riumerous pigments, including anthraquinone pigments such as parietin
(.Santesson, 1970).

Habitat on bark and perhaps occasionally on ecorticate wood.

The most characteristic feature of the Trypetheliaceae is the
c:<olor1ess spores with thickened mesospore. The few brownish spored
ssIDecies known to me have been placed in this family on the basis of

1:11a11us and ascocarp morphology. Polymeridium lacks mesospore thickening

 

eazad a well developed thallus but seems best placed in the Trypetheliaceae
(Dru the basis of hymenial, microconidial and chemical characters, i.e.

eraspersed hymenia of the same type as Trypethelium, rod—shaped micro-

 

c:<3nidia and the presence of lichexanthone in some species. The same

1:){pe of poorly developed thallus is found in Pseudopyrenula which does

 

'Pleave the typical mesospore thickening.
The Trypetheliaceae are almost entirely subtropical and tropical

iLIu.distribution with only a single species Trypethelium virens reaching

 

ESCJuthern Canada. In comparison with the Strigulaceae and Trichotheliac-
€3€1e the range of substrates is very limited.

Astrothelium and Trypethelium are treated briefly in Appendix A.

 

 

lisieudopyrenula is included here with Polymeridium since the single

 

 

1\I<>J:th American species has previously been misidentified as various

Species of Arthopyrenia and also to provide some context for the under-

 

S'tanding of Polymeridium.

 

1" POLYMERIDIUM (Mfill. Arg.) R. C. Harris stat. nov.

fllrthopyrenia sect. Polymeridium Mfill. Arg., Flora 46: 317. 1883.

 

Leictotype (Riedl, 1962): Verrucaria contendens Nyl.

 

151

Thallus whitish, endophloeodal, not differentiated into layers as
in Trypethelium.

Ascocarps flattened, hemispherical or subglobose, immersed or

superficial; outer layer of ascocarp wall brown to blackish, often
lacking below. Hymenium inspersed with amorphous granules or not.
Asci cylindrical to narrowly elliptical or narrowly ovate, with a small,
shallow ocular chamber. Spores biseriate or irregularly arranged,
narrowly ovate or narrowly elliptical, 4-10-celled, cells approximately
equal, slightly constricted at the septa; spore wall mostly not ornament-
ed; perispore visible, occasionally rather thick, up to 2 u.

Microconidia rod-like, 5-12 x 111.

Lichen substances: Lichexanthone in the thallus of several species.

Polymeridium seems to me to be closely related to Pseudopyrenula,

 

with which it has been confused in the past. The main feature separ-
ating them is the lack of mesospore thickening in Polymeridium. Also

in Polymeridium the paraphyses tend to be less regularly reticulate,

 

the ascocarps tend to be smaller, more immersed and have a less heavily
carbonized wal 1 .

It is possible that some of the species placed in Polymeridium
txalong in the genus Arthopyrenia. Many species previously included in

tile sect. Polymeridium clearly have their nearest relatives in Artho-

 

.renia and are placed there, e.g., A, atomarioides and A, cerasi.

13?.Egentepohlia is not evident, the hymenium uninspersed, etc., the

 

deCision as to the correct genus is difficult. In these cases the
natlire of the old spores has proved useful. In Polymeridium the lumen
bexxmnes almost filled with additional layers of spore wall and the

Spcunes do not become tinted. In Arthopyrenia additional wall layers

 

152

are not usually added with age and the spores often become tinted and
ornamented in extreme old age.

As presently circumscribed Polymeridium does not include any species

 

with 2-celled spores. It is possible that the species with 2-celled

spores now included in the Arthopyrenia bifera group should actually be

 

included in Polymeridium. Their spores tend to have some mesospore

 

thickening, the microconidia are of the same type and they are often

asscociated with Trentepohlia. However, my knowledge of the group is

 

not such that I wish to make a decision at this time.

1. Thallus UV+ yellow; spores 4-celled ........................ 2

l0 Thallus UV“; Spores 4-lO-Celled ..0000000.00.000.00000000000 3

2. Spores smooth, 24-33 X 8-10 u; perispore thin ......

................................................. ._P-. catapastllm

 

2. Spores ornamented, 28-36 x 9-12 u; perispore thick

................................................ _P_. exasperatm

 

3. Spores 4-Celled ........................................O... 4

3. Spores more than 4-celled .................................. 7

4. Hymenium inspersed; spores 14-17 x 5-6 u ........ E. contendens

 

4. Hymenium not inspersed ................................. 5

5- Spores large, 28-36 X 9-12 0, ornamented; perispore

thiCk .............O................................ E. exasperatum

 

Spores smaller, less than 25'u long, not ornamented ........ 6

153

6. Spores l7-20(-22) X 5-6.5(-7) u; asci 70-85 p long;

perispore thin ................................. E. subcinereum

 

6. Spores 20-25 X 6-7.5 u; asci 90-125 u long; peri-

spore well developed ............................... P. albidum

7. Spores 6-8-celled, 18-27 X 5.5-7 p ............. E. guinqueseptatum

 

7. Spores 8-lO-Celled, 27-35 X 7-8 p. 0.0000000.0.0.00 E0 Bleiomerellum

 

Polymeridium albidum (Mfill. Arg.) R. C. Harris comb. nov.

 

Arthopyrenia albida Mull. Arg., Flora 67: 664. 1884. Lectotype:

 

Brazil, Cayeté, Martius (M).

Ascocarps subglobose, immersed or emergent, 0.3-0.4 mm in diameter.
Hymenium not inspersed. Asci narrowly elliptical, 90-125 X l6-22 u.
Spores biseriate, narrowly ovate, 4-celled; perispore well developed;
20-25 X 6-7.5 u (excluding perispore),

Microconidia linear, 9-lO X 111.

Habitat on bark and petiole of palmetto.

Polymeridium albidum is very similar to_§. subcinereum, differing
slightly in spore and ascus size. I have seen very few specimens of
either and it is possible that additional material might bridge the gap.

Illustrations: Figs. 367-371.

Specimens seen: United States. FLORIDA: Dade County, Everglades

Nat. Park, SW of Pineland Trail Area, Harris 2889-A (MSC). TEXAS:

 

sine data (MICH).

Polymeridium catapastum (Nyl.) R. C. Harris comb. nov.

Verrucaria catapasta Nyl., Acta Soc. Sci. Penn. 7: 488. 1863.

 

154

Arthopyrenia catapasta (Nyl.) Mull. Arg., Flora 66: 318. 1883. Type

 

collection: Colombia, Bogota, Tequendama, 2600 m, Lindig 2869.

 

Isotype in FH-TUCK 4086.

Byrenastrum album, verrucarioides Eschw. in Martius, Flora Brasil.

 

l: 147. 1833. Arthopyrenia tumida Mfill. Arg., Flora 67: 669. 1884.

 

Lectotype: Brazil, near Bahia, Martius (G).

Thallus white, usually well developed, UV+ yellow (lichexanthone).

Ascocarps subglobose to hemispherical, mostly immersed, 0.3-0.5 mm
in diameter; ascocarp wall often lacking below. Hymenium not inspersed.
Asci narrowly elliptical or elliptical, 80-130 X 20-25 u. Spores irregu-
larly arranged or less often biseriate, narrowly elliptical or narrowly
ovate, 4-celled; perispore relatively thin; 24-33 X 8-10 u (excluding
perispore).

Microconidia rod-like, 6-8 x l'u.

Habitat on bark, in Florida especially on Quercus.

This is the only North American species of Polymeridium which

 

regularly produces large amounts of lichexanthone in the thallus.

Polymeridium exasperatum may produce small amounts, but it is easily

 

distinguished by longer microconidia, ornamented spore wall and thicker
perispore. Polymeridium catapastum is very abundant locally in central
Florida. It is also known from Alabama, the Bahamas, Cuba, Grand Cayman
and Colombia.

Illustrations: Figs. 372-378. Distribution: Fig. 428.

Specimens seen: United States. ALABAMA: Baldwin County, Rock
Creek, 27.II.l925 §y§g§.lgg (MICH). FLORIDA: Dade County, junction of

Hwys. 31 & 74, IV.1970 Hale (US); Lake County, Bustis, 1898 Thaxter 339

 

15S

(MICH); Liberty County, Apalachicola Nat. Forest, Whitehead Lake Camp-

ground, Harris 1449, 1464, 1472 (MSC); Marion County, Ocala Nat. Forest,

 

NW of Big Bass Lake, Harris 2050-8, 2051 (MSC), Hopkins Prairie, Harris

 

2427-A, 2429-B, 2443, 2447, 2466-8, 2473-A (MSC), Hughes Island, Harris

 

 

 

1999, 2034, 2044 (MSC), s of Long Pond, Harris 1802-B, 1820, 1823, 1828,

 

 

 

1835, l842—B, 1843, 1866, 1867, 1870, 2325, 2329-A, 2333, 2353 (MSC),

 

 

 

 

Pats Island, Harris 2068, 2082, 2113 (MSC); Seminole County, Sanford,

 

 

4.X.l904 Rapp (FH), 1.1910 Rapp (FH), 15.X.l912 Rapp (FH); Volusia

County, Tomoka St. Park, Harris 2314, 2321 (MSC).

 

Polymeridium contendens (Nyl.) R. C. Harris comb. nov.

 

Verrucaria contendens Nyl., Acta Soc. Sci Fenn. 7: 492. 1863.

 

Arthopyrenia contendens (Nyl.) Mfill. Arg., Flora 66: 317. 1883.

 

Holotype: Colombia, San Antonio, Lindig 2877 (H-NYL 7331).

 

Ascocarps hemispherical, semi-immersed, c. 0.3 mm in diameter.
Hymenium heavily inspersed. Asci narrowly elliptical, 70-90 x 14-15 u.
Spores subbiseriate, narrowly elliptical or narrowly ovate, 4-celled;
perispore very thin; 14-17 x 5-6 p.

The heavily inspersed hymenium and small spores characterize this

rare species. Nylander in his original description cited Lindig 2877

 

as being from La Mesa but the specimen of this number in Nylander's
herbarium is labelled "San Antonio".

Illustrations: Figs. 379-382.

Specimen seen: United States. FLORIDA: Sarasota County, Myakka

River St. Park, Harris 2642 (MSC).

 

156

Polymeridium exasperatum R. C. Harris sp. nov.

 

Sat similis E. catapasti sed differt sporis exasperatis, perisporio
crassiore, usque 2‘0, et microconidiis longioribus, 9—12 x l p.
Holotype: Florida, Dade County, Everglades Nat. Park, NE of

Pa-hay-okee Overlook, Harris 2829-A (MSC). Isotype in H.

 

Thallus whitish, UV- or patchily UV+ yellowish (lichexanthone?).

Trentepohlia present or absent.

 

Ascocarps hemispherical to subglobose, mostly immersed, 0.4-0.6 mm
in diameter; ascocarp wall occasionally extending outward above, thinner
or lacking below. Ostiole often somewhat eccentric. Hymenium not in-
spersed. Asci narrowly elliptical, with a small ocular chamber, l40-l60
x 20-28 0. Spores biseriate to almost uniseriate, narrowly elliptical
to narrowly ovate, 4-celled; spore wall granular ornamented; perispore
thick, up to 2 p; 28-36 x 9-12 0 (excluding perispore).

Microconidia linear, 9-12 x l p.

Polymeridium exasperatum is the only species in the genus with

 

ornamented spores. The variable presence of lichexanthone and the size

and shape of the spores is very similar to E. catapastum. It is easily

 

distinguished, however, by the features noted in the diagnosis as well
as by the tendency for the ostiole to be eccentric and by larger asci.
It is known only from southernmost Florida.

Illustrations: Figs. 383-386.

Specimens seen: United States. FLORIDA: Dade County, Everglades

Nat. Park, NE of Pa-hay-okee Overlook, Harris 2814-B, 2829-A, 2836-C

 

(MSC); Monroe County, Key West, I.1898 Thaxter 122 (MICH).

 

157

Polymeridium pleiomerellum (Mull. Arg.) R. C. Harris comb. nov.

 

Arthopyrenia pleiomerella Mfill. Arg., Bot. Jahrb. Syst. 6: 406. 1885.

 

Type collection not seen.

Thallus whitish, endophloeodal. Trentepohlia apparently absent

 

in many collections.

Ascocarps hemispherical, mostly superficial but occasionally
immersed, 0.3-0.6 mm in diameter; ascocarp wall lacking below. Hymenium
heavily inspersed. Asci narrowly elliptical or narrowly obovate,
(80-)lOO-l20(-130) X 20-25 0. Spores subbiseriate to irregularly
arranged, narrowly elliptical to narrowly ovate, 8-lO-celled; perispore
usually well developed; 27-35 x 7-8 p (excluding perispore).

Microconidia linear, 7-ll x l p.

Habitat on smooth barks.

The spore size given by Muller in his original description is some-
what smaller than in the material I have seen but the number of cells

agrees. The name is therefore used provisionally. Polymeridium

 

 

pleiomerellum intergrades with E. quinqueseptatum, especially in

 

southern Florida where both occur. Typical E. pleiomerellum is 9-10-

 

celled but also commonly 8-celled while_§. quinqueseptatum is 6-7-
celled ranging up to 8-celled and rarely 9-celled. In southern Florida
many collections are 7-8(-9)-celled and I have more or less arbitrarily
separated them on the basis of spore length ( longer or shorter than

27'0). Material determined as E. pleiomerellum is confined to southern

 

Florida, Mexico and the West Indies while 2, quinqueseptatum ranges
more widely on the Coastal Plain.

Illustrations: Figs. 388-391.

158

Specimens seen: United States. FLORIDA: Dade County, Everglades

Nat. Park, near Pineland Trail Area, Harris 2875-A, 2883-D, 2887, 2896-A,

 

2900, 2908, 2911-0, 2986—F (MSC).

Mexico. GUERRERO: near Cruz Grande, Graham 1219-B (MSC).

 

West Indies. JAMAICA: Parish of St. Andrew, summit of Long Mtn.,

Imshaugle487, 13505-8, 13507, 13515-A (MSC). PUERTO RICO: Mayagfiez

 

District, Yauco, 29.XII.1915 Fink 1512 (PH).

Polymeridium quinqueseptatum (Nyl.) R. C. Harris comb. nov.

Verrucaria guingueseptata Nyl., Expos. Syn. Pyrenoc. 58. 1858.
Arthopyrenia quinqueseptata (Nyl.) Mull. Arg., Flora 68: 326. 1885.
Holotype: South Carolina, Ravenel (H-NYL 591).

Arthopyrenia comparatula Mull. Arg., Jahrb. Bot. Syst. 6: 406. 1885.

Holotype: Cuba, wright Verr. Cub. 152 (G).

Thallus whitish. Trentepohlia present or apparently absent.

 

Ascocarps hemispherical, superficial, 0.25-0.5(-0.6) mm in diameter;
ascocarp wall usually lacking below. Hymenium inspersed. Asci narrowly
elliptical to narrowly ovate, (65-)75-90 x (15-)18-21 p. Spores
narrowly ovate, 6-8(-9)-celled; perispore thin; l8-27 x 5.5-7 0
(excluding perispore).

Microconida linear 9-12 X l 0.

Habitat on smooth barks.

The distinction between this species and E._pleiomerellum is dis-

 

cussed under the latter. Polymeridium quinqueseptatum is less tropical
in distribution than most of the other species in the genus ranging from
South Carolina to Texas and south to southern Florida but is apparently

absent from the West Indies.

159

Illustrations: Figs. 392-395. Distribution: Fig. 429.
Specimens seen: United States. ALABAMA: Baldwin County, Volanta,

9.III.1925 Evans 262 (MICH); Lawrence County, Moulton, 1860 Beaumont 168,

 

228 (FH-TUCK 4023). FLORIDA: Baker County, Osceola Nat. Forest, Harris
3128—C (MSC); Dade County, Everglades Nat. Park, S of Pa-hay-okee Over-

look, Harris 2745-D (MSC), SW of Pineland Trail Area, Harris 2879-B,

 

 

29ll-B (MSC); Duval County, Jacksonville, Calkins (FH, US); Flagler

County, Flagler St. Park, Harris 2478-C (MSC); Liberty County, Apalacha-

 

cola Nat. Forest, Whitehead Lake Campground, Harris l456-A, 1557, 1575-B,

 

1576, 1726 (MSC); Marion County, Ocala Nat. Forest, S of Long Pond,

 

Harris 2345-A (MSC), Pats Island, Harris 2094-A (MSC); Monroe County,

 

 

Key West, 11.1898 Thaxter 299, 301, 361 (MICH); St. Johns County, Six

 

Mile Creek, Calkins (FH); Seminole County, Sanford, IV.1908 Rapp (FH).

LOUISIANA: E. Feliciana Parish, Felixville, TUCker 7762 (MSC); St.

 

Tammany Parish, Covington, 16.IV.1894 Langlois (US); Vernon Parish,

Kisatchie Nat. Forest, Fort Polk, Tucker 7875 (LSU). SOUTH CAROLINA:

 

County unknown, Santee Canal, 1857 Ravenel (FH, FH-TUCK 4023, MICH, MSC,
US); Berkeley County, Francis Marion Nat. Forest, Guilliard Lake Camp-

ground, Harris 3175-B (MSC). TEXAS: Harris County, Houston, coll.

 

unknown (FH-TUCK 4023).

Polymeridium subcinereum (Nyl.) R. C. Harris comb. nov.

 

Verrucaria subcinerea Nyl., Expos. Syn. Pyrenoc. 37. 1858.

 

Arthopyrenia subcinerea (Nyl.) Mfill. Arg., Flora 64: 318. 1883.
Holotype: Texas, 1850 wright (H-NYL 1529). Isotypes in FH-TUCK 4022
& US.

Arthopyrenia subimitans MGll. Arg., Bull. Soc. Bot. Belg. 32: 169.

 

160

1893. Type collection: Costa Rica, Boruca, Pittier 6287. Isotype in

 

US.

Ascocarps hemispherical or flattened, semi-immersed, 0.3-0.5 mm in
diameter. Hymenium not inspersed. Asci narrowly elliptical, 70-85
x 15-22 0. Spores biseriate or subbiseriate, narrowly elliptical to
narrowly ovate, 4-celled; perispore thin; l7-20(-22) x 5-6.5(-7) u.

Microconidia rod-like, 5-7 x l p.

This species is very similar to P. albidum but has smaller spores
and asci. It is known from Florida, Texas and Costa Rica.

Illustrations: Figs. 396—399.

Specimens seen: United States. FLORIDA: Orange County, Rock
Springs, 6.V.l921 Eglly 825 (US). TEXAS: County unknown, thickets of
the Blanco, 1850 Wright §§_(US); Hidalgo County ?, valley of the Rio

Grande below Donna, Wright (FH-TUCK 4023).

2, PSEUDOPYRENULA Mull. Arg.

Flora 66: 247. 1883. Lectotype (Clements, 1909): Verrucaria diluta

 

Fee.

Plagiotrema Mull. Arg., Bot. Jahrb. Syst. 6: 387. 1885. Holotype:

 

.3. cubanum Mull. Arg.

Thallus whitish, endophloeodal, not differentiated into layers as in

Trypethelium. Phycobiont Trentepohlia, but some species apparently not

 

lichenized.
Ascocarps flattened, hemispherical or subglobose, mostly superficial,
rarely immersed; ascocarp wall carbonaceous, often very hard and brittle,

often extended outward at the base, often lacking below. Hymenium often

161

inspersed with amorphous granules. Paraphyses branched and anastomosed,
often quite regularly and reticulately. Asci cylindrical or narrowly
elliptical, often with a broad, shallow ocular chamber. Spores ellipti-
cal to narrowly elliptical, 4-ce11ed; mesospore thickened so that the
lumina are angular or rounded in optical section; spore wall not orna-
mented; perispore thin.

Microconidia rod-like.

Lichen substances: Lichexanthone in one species and a yellowish
pigment, KOH+ red, in the hymenia of several species.

Pseudopyrenula is distinguished from Trypethelium mainly on the

 

basis of thallus and ascocarp characters. The thallus of Pseudopyrenula
does not have a cortex-like upper layer and the ascocarps are super-

ficial and often rather flattened. In Trypethelium the ascocarps are

 

immersed, either singlely in the thallus or grouped within pseudostromata.

The distinction of Pseudopyrenula from Polymeridium is discussed under

 

 

the latter genus.

There is only a single species of Pseudopyrenula in North America.

 

The previous reports of B. pupula (Ach.) Mull. Arg. were based on mis-

identified specimens of Trypethelium floridanum.

 

Pseudopyrenula subgregaria Mull. Arg.

 

Bot. Jahrb. Syst. 6: 408. 1885. Holotype: Cuba, Wright Verr. Cub.
80 (G). Isotype in US.

Pseudopyrenula confluens Merr. ig_Hedrick, Mycologia 22: 247. 1930.

 

Holotype: Puerto Rico, near Aibonito, 4.1.1916 Fink 1856 (MICH).

Isotype in FH.

162

Thallus whitish, endophloeodal, Trentepohlia present or not.

 

Ascocarps flattened to hemispherical, rarely subglobose, often
somewhat aggregated in irregular groups or rows, 0.3-0.7 mm in diameter;
ascocarp wall often exteneded outward at the base, thinner or lacking
below. Hymenium inspersed with amorphous granules and with a yellowish
pigment, KOH+ red, pigment rarely lacking. Asci narrowly elliptical,
75-95(-l20) x 15-20 0. Spores 20-27 x 6-8 p.

Microconidia rod-like, 5-8 x l 0.

Habitat on bark and old wood.

In general species in the genus Pseudopyrenula do not seem to me

 

at this time to be very distinct. There is considerable intergradation
in spore size and the presence or absence of the yellowish hymenial
pigment does not seem to correlate with other characters. Pseudopyrenula

subgregaria has the smallest spores in the genus and the presence of the

 

pigment is quite constant so that it is perhaps a little better defined
than most. It is apparently rather common in Florida and the West
Indies ( Cuba, Dominica, Dominican Republic, Grand Cayman, Jamaica and
Puerto Rico) and represented from Louisiana and Costa Rica by single
collections.

Illustrations: Figs. 400-405. Distribution: Fig. 430.

Specimens seen: United States. FLORIDA: sine loc., Shockley

 

(US); Dade County, Everglades Nat. Park, near Mahogany Hammock, Harris
2929-C (MSC), S of Pa-hay-okee Overlook, Harris 2719-H, 2741-B (MSC),

near Pineland Trail Area, Harris 2891, 2906, 3016, 3018 (MSC); Liberty

 

County, Apalachicola Nat. Forest, Whitehead Lake Campground, Harris

1499, 1515, 1521 (MSC), Porter Lake Picnic Area, Harris 1689 (MSC).

 

 

LOUISIANA: E. Baton Rouge Parish, Baton Rouge, Tucker 10005 (LSU).

 

VI. SOME ADDITIONAL HYALINE_SPORED PYRENOLICHENS

In order to provide some sort of preliminary manual for the trans-
versely septate, colorless spored pyrenolichens (excluding the Verru-
cariaceae) this appendix contains keys, nomenclatural citations and
brief statements of diagnostic characters and distribution. All of the
species known to me from the United States are included with the
exception of Porina, where a few rare species found only in southernmost
Florida have been omitted.

In this appendix sf. is used to indicate an identification based
almost entirely on the literature without examination of authentic
material, while §££° is used to indicate an unnamed species which seems
very closely related to a well known species and is only tentatively

maintained as a separate species.

A. PYRENULACEAE Rabenh.
Kryptog.-F1. von Sachsen 2: 42. 1870. Holotype: Pyrenula Ach.
Paratheliaceae Zahlbr., Nat. Pflanzenfam. 1(1‘): 71. 1903.

Holotype: Parathelium Nyl.

 

Gleophragmiaceae Rés., Acta Bot. Fenn. 33: 9, 22, 72. 1943. nom.

inval. descr. germ. Lectotype (Cooke & Hawksworth, 1970):

 

 

Mycopyrenula Vain.
Gloenodictyaceae Rés., Acta Bot. Fenn. 33: 9, 22, 73. 1943. nom.

inval. descr. germ. Lectotype (Cooke & Hawksworth, 1970): Bottaria

 

 

Massal.
Microglaenaceae Serv., Cesk. Lisejniky Celedi Verruc. 17. 1954.

Iiolotype: Microglaena Korb.

 

163

 

164

Cooke & Hawksworth (1970) attribute the first publication of the
name Pyrenulaceae to Zahlbruckner (1903-7), but a superficial search
has turned up an earlier usage by Rabenhorst and a more exhaustive
search might reveal even earlier ones.

Although the Pyrenulaceae is mainly comprised of brown spored
species, there are a few with colorless spores. They are included in
the Pyrenulaceae on the basis of ascocarp structure, hymenial characters

and type of microconidia.

1. LITHOTHELIUM Mull. Arg.

Bot. Jahrb. Syst. 6: 386. 1885. Holotype: I? cubanum Mull. Arg.

A compound ascocarp containing several hymenia sharing a single
ostiole, inspersed hymenium, hymenial gelatin IKI+ bluish to orangish

and filiform microconidia are diagnostic for Lithothelium.

 

Lithothelium cubanum Mfill. Arg.

Bot. Jahrb. Syst. 6: 386. 1885. Type collection: Cuba, Wright
Verr. Cub. II: 612. Isotype in FH-TUCK 3982.

Verrucaria falklandica Nyl., Lich. Fueg. & Patagon. 22. 1888.

Holotype: "Insula Falkland, calcicola, Rabenhorst fil." (H-NYL 1274).

 

Porina (sect. Sagedia) macrocarpa Ridd. lg Brit. & Millsp.,
Bahama F1. 523. 1920. Type collection not seen.

Lithothelium violascens Malme, Ark. Bot. 19(1): 16. 1924. Type
Collection: Malme Austroam. 25, Paraguay, Colonia Risso pr. Rio Apa,

IVlalme. Isotype in MSC.

165

The ascocarp wall is usually rather purplish in this species, which

led Malme to name it "violascens". Lithothelium has not yet been found

 

in North America but its rather broad subtropical distribution suggests
that it might well occur on limestone in southern Florida. The Falkland

Island locality seems somewhat doubtful.

2. PLAGIOCARPA R. C. Harris

Michigan Bot. 12: 34. 1973. Holotype: fig. septemseptata R. C. Harris.

 

See Harris (1973) for description and discussion.

Plagiocarpa hyalospgra (Nyl.) R. C. Harris
Michigan Bot. 12: 34. 1973. Verrucaria hyalospora Nyl., Mem. Soc.

Acad. Maine Loire 4: 48. 1858. Holotype: New England (H-NYL 1251).

See Harris (1973) for description and discussion.

In addition to the widespread E, hyalospora there are two other

 

colorless spored species which are very similar. They differ from

§g_hyalospora in having Pyrenula type asci, lacking the very distinctive

 

ocular chamber unique to Plagiocarpa. Their distribution is more

 

tropical but they agree with Plagiocarpa in all other respects. At the

 

present time I do not wish to describe them in Plagiocarpa or create a
new genus for them. The two species differ in spore size. The smaller
Spored species (17-20 x 7-10 0) is known from a few collections from
Louisiana and one from the West Indies (St. Lucia). The larger (23-29
X 10-12.5 u) is mixed with the original material of Anisomeridium

tamarindi (G) and probably comes from Guadeloupe.

166

B. TRICHOTHBLIACEAE (Mull. Arg.) Bitt. & Schill. in Schill
Hedwigia 67: 273. 1927. Pyrenulae subtr. Trichotheliae Mull. Arg.,

Bot. Jahrb. Syst. 6: 418. 1885. Holotype: Trichothelium Mfill. Arg.

 

Porinaceae W. Wats., New Phytol. 28: 108. 1929. Holotype: Porina
Mull. Arg.
Gloenoblastiaceae Ras., Acta Bot. Fenn. 33: 9, 22, 74. 1943. nom.

inval. diagn. germ. Lectotype (Cooke & Hawksworth, 1970):

 

 

StereochlamysfiMfill. Arg.

 

The Trichotheliaceae seem to form a homogeneous unit if one

accepts the definition proposed in this work. The asci are character-
istically rather truncate, the ascus tip is little thickened and most
species have a chitinoid ring in the apex of the exoascus. Other aids
to recognizing the family are the unbranched paraphyses, spores mostly
multiseptate, ascocarp wall often some color other than brown black and
abundant colorless crystals in the thallus, especially around the asco-
carp. The bulk of the family is subtropical or tropical. The majority
of the species are corticolous, but some species grow on rock, mosses or

leaves.

1. PORINA Mull. Arg.
Flora 66: 320. 1883. nom. cons. Lectotype: ‘P. nucula Ach.

Pseudosagedia (Mull. Arg.) Choisy, Bull. Mens. Soc. Linn. Lyon.

 

18: 107. 1949. Arthopyrenia sect. Pseudosagedia Mull. Arg., Mém. Soc.

 

 

Phys. Geneve 16: 428. 1862. Type not selected but all original species

have been referred to Porina.

167

Porina is one of the largest genera of pyrenolichens in North

America, but many of the species are rare. The genus presents a number

of taxonomic problems which will require considerable additional study.

Porina is most diverse in Florida, thus it is probable that some of the

unnamed species will prove to have been described in other regions of

tropical America.

3.

Growing on leaves (after Santesson, 1952) .................. 2

Growing on bark, wood, mosses or rock ...................... 4

2. Spores 8-celled, 24-31 x 3-4-0; ascocarps reddish
brown, constricted at the base .................... P. octomera
2. Spores 4-6—ce11ed; ascocarps black with a thin or

thick thallus covering ................................. 3

Spores 4-ce11ed, 18-25 X 4-5 p; thallus covering more or

less thick, ascocarp thus similar in color to thallus
°'°°°°°°'°"'°"°°°°'°°°°°°°°°'°'°'°'°°°'°"'°'°°°'°°°°.E‘ thaxteri
Spores 6-celled, 20-28 x 3.5-4.5 u; ascocarps blackish

to grayish, with a thin thallus covering .............. P. nitidula

4. Spores 4-Celled 0.000.000.0000.........OOOOOOOOOOOOOOCOO 5

4. spores more than 4-Celled 0.0.0..........OOOOOOOOOOOOOOO 12

Involucrellum black (lacking reddish tints even in
Section) ......O.’......OOCOCOOOOIOOO......OOOOOOOCOOOCCOCCO 6
Involucrellum orange red to reddish black (if necessary

examineat-lflin section) 0..........OOOOOOOOOOOOO......OOOOOO 8

7.

11.

11.

168

60 GrOWing 0n bark; Spores 16-24 X 405-5 11 0.0.0000... _P_. carEinea

6. GrOWing on rOCk 0......0.0.00.........OOOCOOOOOOOOOOOOCO 7

Microconidia oblong, 2-3 u long; spores 18-25 x 4.5-5 0

......OOOOOOOOOO......OOOOOOOOOOOOOO......OOOOOOOOOOO B. Chlorotica

 

Microconidia cylindrical, 5-10 u long; spores 18-25 x

5.5-711 ......OOCOOOOCCOOOOOOO......OOOOOOOOOOOO B. aff. Chlorotica

 

8. Growing on bark or on mosses and plant remains ......... 9

8. GrOWing on rOCk 00.....................OOOOOOOOOOOOOOCOO 11—

Growing on bryophytes and plant remains; involucrellum

purplish red in section; spores 25-40 x 4-6 p (Swinscow,

1962) ................................................ P. mamillosa
Growing on bark; involucrellum orange red to yellowish

in section ......OOOOOOOOOOO.........OOOOOOOOOOOOOOO00...... 10

10. Microconidia oblong, 2-3 u long; spores 16-23 x
3-5 p (Swinscow, 1962) ........................... P. leptalea
10. Microconidia cylindrical, c. 5 u long; spores

17-23 x 4.5-5.5 u .............................. £5 microspora

 

Spores 20-30(-40) x 5-8 p (Swinscow, 1962); micro-
conidia elliptical with pointed ends, 0. 1.5 p broad

0.0.0.0............OOOOOOOOOOOOOOOOOO......OOOOOOOOO £0 lectiSSj—ma

 

Spores 15-19 X 4.5-6 0; microconidia oblong l'p or

less broad 0.......O...........OOOOOOOOOOOOOOOO IE. aff. lectiSSj—ma

 

13.

13.

15.

15.

17.

17.

169
12. Ascocarp wall green; involucrellum absent; spores
7-9-celled, 28-38 x 6.5-9 0 ..................... Porina sp. 1
12. Ascocarp wall or involucrellum yellowish, reddish,

brown or blaCk ....................OOOOOOOOOOOOOOOOOO. l3

Involucrellum or ascocarp wall pink, yellowish or
reddish, sometimes darker colored around the ostiole ...... 14

Involucrellum or ascocarp wall dark brown to black ........ 23

14. Growing on rock; ascocarps pinkish orange, lacking
involucrellum; spores 9-15-celled, 55-75 X
705-911. O00.0.00.........OOOOOOOOO......OOOOOOOOO Porina Sp. 2

14. GrOWing on bark ....0...............OOOOOOOOOOCOOOOOOO 15

Thallus shiny, thin and fragile, with a definite black
hypothallus, often becoming detached from the substrate;
ostiole usually surrounded by a dark ring ................. l6
Thallus dull, without evident hypothallus and never

becoming detached from the substrate; ostiole not

normally surrounded by a dark ring ........................ 18

16. Thallus with scattered cylindrical isidia; spores
8-C€lled, 30-50X5-705p 000000000000.0.0....0.. Porina Sp. 3

16. |I‘lvlaJ-llls laCking iSidia O0............OOOOOOOOOOOOOOOOO l7

Spores 35-50 x 5-7.5 p, 8-celled ................ 2. cf. tetracerae

 

Spores 42-55 X 8'905 11, 8-C€ll€d .0000000000000000000 £0 SUbiungenS

 

19.

19.

21.

21.

23.

23.

25.

25.

170

18. Thallus covered with simple to coralloid isidia;
ascocarps immersed, flesh-colored; spores 9-18-
celled, 65-110 X 705-1211 ......OOOOOOOOOOOOOOOOO Porina Sp. 4

18. Trlallus not iSidiate ......OOOOOOO......OOOOOOOOOOOOOO 19

Spores dactyloid, much narrower at one end ................ 20

Spores narrowly elliptical or narrowly ovate .............. 21

20. Spores 11-15—ce11ed, 75-125 x 10-15 p; involucrellum
filled with crystals .......................... P, heterospora
20. Spores 8-ce11ed, 37-42 X 7-8 p; involucrellum not

filled With crystals ......OOOOOOOOOOO0.0.0.0.... Porina Sp. 5

Spores 35-6OX705-10p ..........OOOOOOOOOOOOOOOOOO £0 aff. nucula

Spores mostly over 10 p in width .......................... 22

22. Spores 45-55 x 10-14‘0 ............................. E. nucula

220 Spores 55-80X10-13p 000000....0000000.00.00... porina Sp. 6

GrOWing on rOCk 00.00.0000...000............OOOOOOOOOOOOOOO 24

GrOWing on bark ...O....OOOO0.............OOOOOOOOOOOIOOOOO 26

24. Spores 33-40 X 6.5-8(-10) p, 8(-12)-celled, cylin-
drical to narrowly ovate ........................ Porina sp. 7

24. Spores 6.5 u or less in width ........................ 25

Spores cylindrical, 30-47 x 4.5-6 0, 8(-l4)-ce11ed ... Porina sp. 8
Spores dactyloid, tapering to a long tail, 45-60 X 5-6 p,

8(-13)-Celled O0.0.000...0.000.000.0000.........OOOOOO Porina Sp. 9

171

26. Spores filiform, 100-140 x 3-5 0, up to 2l-ce11ed ....

O00.0.00.........OOOOOOOO......OOOOOOOOOOOOOO E. raphidospema

 

26. Spores 90 p or less in length ........................ 27

27. Spores 8(rare1y —10)-celled, narrowly ovate, 30-50 x

5-7.511 000............OO...00.00.0000...0.0.0.000... £0 cestrenSis

 

27. Spores (8-)10-18—ce11ed, cylindrical, dactyloid or
linear, mostly narrower at one end in the larger spored
forms, if only 8-celled, then the spores cylindrical,

40-9- X 3-7(-7.5) u ............... see discussion of P. cestrensis

 

Porina carpinea (Pers. ex Ach.) Zahlbr.

 

in_Engler & Prantl, Natfirl. Pflanzenfam. l(l‘): 66. 1903. Verrucaria

 

carpinea Pers. ex Ach., Meth. Lich. 120. 1803. Type collection not seen.

The small black ascocarps, 4-celled spores and corticolous habitat
are diagnostic. Specimens have been verified from British Columbia,

Florida and Texas.

Porina cestrensis (Tuck. ex Michener) Mfill. Arg.

 

Flora 64: 338. 1883. Verrucaria cestrensis Tuck. gx_Michener lg
w. Dar1., F1. Cest. ed. 3. 452. 1853. Lectotype(?): Pennsylvania,

Chester, Michener 204 (FH-TUCK 3986).

 

Porina cestrensis var. platyspora Fink in_Hedrick, Mycologia 25:

 

 

308. 1933. Holotype: Indiana, near Scipio, 8.II.1909 (MICH, hb. Fink

8,889).

Porina cestrensis is one of the commonest species in eastern North

 

America. Unfortunately it is involved in one of the worst taxonomic

172

problems in the genus. A majority of collections have uniformly 8-
celled, narrowly ovate spores, but a sizable number of collections have
spores which range, in different collections, from 8-celled and cylindri-
cal to linear, often with one end narrowed, and up to 18-celled. There
do not seem to be any immediately obvious discontinuities in this con-
tinuum although the various spore forms encompass a range of variation
not normally included in a single species. Swinscow (1962) includes
narrowly ovate and cylindrical spored forms as varieties of a single
species. Thus one might include the 8-celled cylindrical spored

forms in P. cestrensis proper. Some of the longest spores begin to

 

approach the very long filiform spores of P. rhaphidosperma. I am

 

tempted to speculate that the considerable number of specimens intermed-

iate between.§§ cestrensis and P. rhaphidosperma represent a hybrid

 

 

swarm. This would account for the lack of discontinuities and the extra-

ordinary variation in spore size, shape and septation. Porina hibernica

 

James & Swinsc., which is apparently a distinct species in Europe,

represents the upper end of the range of this swarm. Porina olivacea

 

(Pers.) A. L. Sm. is the European vicariant of P. cestrensis and differs

 

only in slightly smaller spores.

Porina cestrensis and its variants range from Massachusetts to Ohio

 

south to Florida and eastern Texas.

Porina chlorotica (Ach.) Mull. Arg.

Revue Mycol. (Toulouse) 6: 20. 1884. Verrucaria chlorotica Ach.,

 

Lich. Univ. 283. 1810. Type collection not seen.

This is the saxicolous form 0f.E° carpinea, both having black asco-

carps and 4-celled spores. It is known in North America only from

173

Massachusetts.

Porina aff. chlorotica

This is a marine or maritime species found so far only in British
Columbia and Alaska. I had originally included the western population
in E. chlorotica but the broader spores and longer microconidia suggest

it should be recognized as a separate species.

 

Porina heterospora (Fink ig_Hedrick) R. C. Harris stat. nov.

 

Porina nucula var. heterospora Fink 22 Hedrick, Mycologia 25: 308.

 

1933. Holotype: Florida, Calkins (MICH, hb. Fink 15,512).

Although clearly related to E. nucula, the spores are many times
longer and differently shaped. Swinscow's (1962) description of P.
nucula seems to be based, at least in part, on specimens of this species.

Porina heterospora is common in the southern part of the Coastal Plain.

 

Porina lectissima (Fr.) Zahlbr.
ig_Engler & Prantl, Natfirl. Pflanzenfam. 1(1‘): 66. 1903. Segestria

lectissima Fr., Syst. Orb. Veget. 287. 1825. Type collection not seen.

 

This species is recognized by its reddish ascocarp, 4-celled spores
and saxicolous habitat. The only verified specimen from North America

is from New Hampshire.

Porina aff. lectissima
Just as in E. chlorotica, the western population seems sufficiently

different from eastern North American and European material to warrant

174
recognition. The spores are smaller and the microconidia narrower. It

is represented by a single specimen from California.

Porina leptalea (Dur. & Mont.) A. L. Sm.

 

Monogr. Brit. Lich. 2: 333. 1911. Biatora leptalea Dur. & Mont.,

 

_ig Dur., Fl. d'Alger., Crypt. 1: 268. 1846-9. Type collection not seen.

Porina leptalea is related to P. lectissima but has smaller spores

 

and grows on bark. It is distinguished from E, microspora mainly by its

microconidia. New to North America, it is so far known only from Florida.

Porina mamillosa (Th. Fr.) Vain.

 

Acta Soc. Fauna Fl. Fenn. 49: 176. 1922. Segestria mamillosa Th. Fr.,

 

Lich. Arctoi 262. 1860.

This species is readily distinguished by its substrate, 4-celled
spores and purplish-red involucrellum. It has been reported by Anderson

(1967) from alpine areas of Colorado.

Porina microspora (Fink i2_Hedrick) R. C. Harris stat. nov.

Porina olivacea var. microspora Fink ig_Hedrick, Mycologia 25: 308.

 

1933. Holotype: New Hampshire, Chocorua, Lonely Lake, VIII.1918 Farlow

(MICH, hb. Fink 11,559). Isotypes in FH, FH-RIDD and MICH.

Fink, failing to notice the reddish tints in the involucrellum,
thought his species to be a variety of P. olivacea.‘ It is questionably
distinct from_§. leptalea which has shorter microconidia and brighter
reddish ascocarps. In other genera with which I am better acquainted,

this sort of difference in the microconidia is often significant, thus

 

175
I am recognizing the species, at least until I have had more experience

with the genus Porina. Porina microspora is known only from the type

 

 

collection.

Porina nitidula Mfill. Arg.

 

Flora 66: 336. 1883. Type collection not seen.

See Santesson (1952) for description and distingushing characters.

Porina nitidula is new to North America and was collected in Dade County,

 

Florida by Roland Thaxter.

Porina nucula Ach.

 

Syn. Lich. 112. 1814. Lectotype: Africa, Guinea (H-ACH).

I am indebted to Henry Imshaug for a preparation from the Acharian
specimen and to Mason Hale for a photograph of it. Porina nucula has
not been generally correctly interpreted. A number of closely related
species have the same thallus and ascocarp type and specific identifica-
tion depends on spore characters. The thallus is generally warted or
rugose and contains numerous colorless crystals; the upper part of the
ascocarp is yellowish to reddish, occasionally almost black around the
ostiole, and the involucrellum also contains numerous crystals. Porina
nucula is distinguished by its narrowly elliptic, 8-celled spores which
are 10 p or more in width. It is relatively rare in North America, known
from Florida and South Carolina. I have also verified a specimen from

the West Indies (St. Thomas). Brodo's (1968) report is all Porina sp. 4.

 

176

Porina aff. nucula

This taxon differs from P. nucula by its consistently narrower
spores. Further study is needed to determine at what taxonomic rank it
would be best recognized. It is rather common in the southern part of

the Coastal Plain.

Porina octomera (Mfill. Arg.) Schill.

 

Hedwigia 67: 274. 1927. Phylloporina octomera Mull. Arg., Flora

 

 

73: 198. 1890. Type collection not seen.

See Santesson (1952) for description and distinguishing characters.

Porina octomera is new to North America. It has only been collected

 

once in Dade County, Florida by Thaxter.

Porina rhaphidosperma Mull. Arg.

 

Hedwigia 34: 35. 1895. Type collection: Massachusetts, on hemlock,

Willey. Isotypes in FH-TUCK 3988 and US.

This species is immediately recognizable by its filiform spores.
For a discussion of the relationship of this species with the P.
cestrensis-complex, see under that species. Apparently not common,

verified specimens come from Florida, Kentucky and Massachusetts.

Porina subpungens Malme
Ark. Bot. 23A(1): 17. 1929. Isosyntype: Paraguay, Colonia Risso,

l6.X.1893 Malme (MSC).

The identification of_P. subpungens from Florida is somewhat tenta-

tive since the spores in the isosyntype are slightly broader, but the

177

specimens are otherwise identical. The shiny thallus, black hypothallus

and relatively broad spores characterize the species. Porina subpungens

 

has not previously been reported from North America.

Porina cfr. tetracerae (Afz. ex Ach.) Mu'll. Arg.

Bot. Jahrb. Syst. 6: 401. 1885. Verrucaria tetracerae Afz. _ex Ach.,

 

Meth. Lich. 121. 1803. Lectotype: Africa, Sierra Leone, ad corticem

Tetracerae, Afzelius (H-ACH).

 

Tentative recognition of this species in North America is based on
a preparation from the Acharian specimen by Dr. Imshaug and on Malme's
(1929) discussion of its relationship to P. subpungens, from which it
differs by its narrower spores. I have examined several specimens from

southern Florida and one from Puerto Rico.

Porina thaxteri Sant.

 

Syrnb. Bot. Upsal. 12(1): 218. 1952. Type collection not seen.

See Santesson (1952) for description and distinguishing characters.

prina thaxteri was described from Florida.

 

rina sp. 1
Unique among North American Porinas due to the green or blue-green
)r of the ascocarp. It has been collected in two localities in

hern Florida.

Esp. 2

This species is related to B. lectissima by the color of its

 

178
ascocarps and saxicolous habitat. It differs in having larger, multi-
septate spores. It is known from a single collection by Tuckermann from

the White Mountains of New Hampshire.

Porina sp. 3

One of two isidiate species in North America, it seems from Malme's
(1929) description closely related to P. conspersa Malme, differing by
8-celled, slightly smaller spores. It has been found several times in

southern Florida.

Porina sp. 4

The thallus is often so densely isidiate as to appear granulose,
which along with the palid ascocarps and large spores make this species
unique. It seems to have the same type of thallus as P. coralloidea
James but differs in ascocarp color and spores. It has been found only

on Long Island, New York, and was reported by Brodo (1968) as P. nucula.

Porina sp. 5

The spores of this species closely resemble those of P. cestrensis,
but it differs from that species in its reddish ascocarps (orange-yellow
in section). It is known from a single collection by Calkins in

Tennessee.

Porina sp. 6
Very similar to P. nucula externally, but it differs in having

larger, more pointed spores. It occurs rarely in central Florida.

 

179

Porina sp. 7, sp. 8 and sp. 9

These three taxa are all related to_P. guentheri (Flot.) Zahlbr.
The spores of P. ggentheri are 8—celled while the North American material
is variable with some spores 8-celled but ranging up to 12-14-celled.
They show the same sort of variation treated by Swinscow (1963) as var-
ieties of P. guentheri and, perhaps, should be treated as such. All are

known from only one or two collections from New England.

2, TRICHOTHELIUM Mull. Arg.
Bot. Jahrb. Syst. 6: 418. 1885. Holotype: Trichothelium epiphyllum

Mfill. Arg.

For additional synonymy and description see Santesson (1952). The
asci of both T. epiphyllum and T. horridulum have chitinoid rings in the

tip closely linking the genus with Porina.

Trichothelium horridulum (Mull. Arg.) Sant.
Symb. Bot. Upsal. 12(1): 278. 1952. Stereochlamys horridula

Mull. Arg., Flora 68: 344. 1885. Type collection not seen.

See Santesson (1952) for description and distinguishing characters.
Previously known only from Brazil, it was collected by Thaxter in Marion

County, Florida.

 

180

C. TRYPETHELIACEAE Eschw.

See pages 148-149 for synonymy and diagnosis.

1. ASTROTHELIUM Eschw.
Syst. Lich. 18. 1824. Lectotype (Massalongo, 1860): A. conicum Eschw.
EHeufleria Trev., Spighe e Paglie 19. 1853. Holotype:

A. conicum Eschw.

The genus Astrothelium was based on two syntypes, A. conicum and

 

A, isabellinum. Trevisan (1853) created a new genus Heufleria with only

 

a single species, Astrothelium conicum, but apparently made no mention

of A. isabellinum. The present day rules of nomenclature recommend that

 

the remaining species, A. isabellinum, should be the lectotype of

 

Astrothelium. However, Massalongo in 1860 was under no such constraints

 

and his explicit lectotypification of Astrothelium with A, conicum has

 

priority as far as I know. Nor, since Trevisan did not mention

A, isabellinum, does his creation of Heufleria constitute a schizotype

 

(Korf & Rogers, 1967). In 1973 Imshaug was unable to locate either of
the type specimens involved although others of Eschweiler's types were
found. Thus the application of the names is still somewhat uncertain
and it may eventually be necessary to designate neotypes for them.

Astrothelium differs from Trypethelium in having a multichambered

 

ascocarp with a single common ostiole. In the Pyrenulaceae this type
of ascocarp apparently results from the fusion of several ascocarps.

It is possible that in Astrothelium a compound ascocarp results from

 

invaginations of the wall within a single ascocarp. The ascocarps of

Trypethelium ochroleucum are often found partially divided into

 

181

chambers in a manner suggestive of Astrothelium. Also it seems to me
that various species of Astrothelium are more closely related to species

of Trypethelium than to each other. This suggests that the two genera

 

could be merged, however, further study is necessary before any final
decision.

The three species of Astrothelium occurring in North America are
closely related and have the stromata pigmented orange outside, at least
at the tip. Thin layer chromatography has shown that the major pigment
responsible is parietin. The species in this group are separated by
spore size and presence or absence of lichexanthone. The genus is not

included in Hale & Culberson (1970) but A, ochrothelizum was reported by

Herre (1942).

1. Thallus UV-, usually greenish or olivaceous, shiny;
Spores 23-30(-32) X 8-lO("l2) p ......0..00000.000.. A. CinnaInomem
l. Thallus UV+ yellow (licheXanthone), pale tan, usually

prUlnose ......OOOOOOOOOO00.00.....0.........OOOOOOOOOOOIOOO 2

20 Spores 21.-28 X 7-9(-10) p. 00000.00.....0000000 A. OChrOthElizum

2. Spores (26‘)28-35(-39) X 10-13 1]. .0..00.0000...00 A. verSiCOlOr

Astrothelium cinnamomeum (Eschw.) Mull. Arg.

Flora 68: 270. 1884. Pyrenastrum cinnamomeum Eschw. ig_Mart.,
Icon. Pl. Crypt. 18, pl. 9, f. 1. 1828. Holotype: Brazil, prope
Caiteté (M).

Astrothelium minus Mull. Arg., Bot. Jahrb. Syst. 6: 382. 1885.

Type collection: Cuba, Wright Verr. Cub. 235. Isotypes in FH-TUCK

3982 & US.

 

182

Astrofluflium minus var. nigratum Mull. Arg., Bot. Jahrb. Syst. 6.

Cuba, Wright Verr. Cub. II. 638. Isotype

382.1885. Type collection:

in FH-TUCK 3982.

The smfll spores and rather shiny thallus lacking lichexanthone

characterize the species. I have verified three collections from

norflMaXIF1orida, as well as collections from Brazil, Cuba, French

Guiana, Tobago and Trinidad.

Astrothelium ochrothelizum Mull. Arg.
Bot. Jahrb. Syst. 6: 382. 1885. Type collection: Cuba, Wright

Verr. Cub. 144. Isotype in FH-TUCK 3982.

Astrothelium conicum var. pallidum Mull. Arg., Bot. Jahrb. Syst.

wright Verr. Cub. 605. Isolecto-

6: 382. 1885. Lectotype collection:

types in FH-TUCK 3981 & US.

This species has the smallest spores of those containing liche-

xanthone. It is represented in North America by three collections made

in Florida by Rapp. It also occurs in Cuba and French Guiana.

Astrothelium versicolor Mull. Arg.
Puerto Rico, Sintensis 6 (0).

Flora 71: 495. 1888. Holotype:

This species is distinguished from A. ochrothelizum, which also

roduces lichexanthone, by its larger spores. It is known to me from

lorida and also British Guiana, French Guiana and Puerto Rico.

 

 

183

2. TRYPETHELIUM Spreng.
Anleitung Kenntn. Gewéchse 3: 350. 1804. nom. cons. (syn. prius

Bathelium Ach., Meth. Lich. 111. 1803). Holotype: '2. eluteriae Spreng.

Trypethelium seems to me to be distingushed from Pseudopyrenula

 

 

mainly by thallus characters. Trypethelium has a very well developed

 

thallus with a cortex-like upper layer. The thallus of Pseudopyrenula

 

lacks any such layer and may possibly be occasionally non-lichenized.

Also the ascocarp in Pseudopyrenula is flattened or conical and is not

 

immersed in a pseudostroma or in the thallus. Thus I feel a number of
species, such as P. calospora Mull. Arg. and_P. papulosa (Nyl.) Mull.

Arg., placed in Pseudopyrenula on the basis of having scattered ascocarps

 

not aggregated in a pseudostroma should be transferred to Trypethelium.

At one time I separated species of Trypethelium on the basis of

 

whether they had an inspersed hymenium and lacked crystals in the asco-
carp wall or whether they had an uninspersed hymenium and crystals were
present in the wall. While most collections belong to one type or the

other, I have found a few collections of T. ochroleucum which have both

 

an inspersed hymenium and a few crystals in the ascocarp wall. These

two types are also found in T. aeneum,_T. ochroleucum and Trypethelium

 

sp. 1. In the Pyrenulaceae the presence of oily substances in the hymen-
ium seems to be useful, but in view of the intermediates and the parallel
occurence in several species I feel it should not be stressed in these
cases in the Trypetheliaceae. I wonder if the oily substances in the
hymenium and the crystals in the ascocarp wall might not be different

expressions of the same metabolic pathway.

 

5.

184
Thallus entirely or partly orange pigmented, KOH+ purple;

Spores 21-28X8-905u 0............OOCOOOOOOOOOOO0.00.... 2. aenem

Thallus not orange pigmented, KOH- ......................... 2

2. Pseudostroma pigmented pale yellow to orangish within .. 3

2. Pseudostroma not pigmented within ...................... 4

Spores 4-celled, 18-28 x 6-9 0 (Johnson, 1959); pseudo-

stromata irregular, brownish, not constricted at the

base, pigment KOH- ................................... T. mastoideum
Spores up to 12-ce11ed, 40-50 x 9-12 p; pseudostromata

often concolorous with the thallus (may become brownish

or yellowish in age), cushion-shaped and often constricted

at the base, pigment KOH+ purple ...................... T. eluteriae

4. Thallus UV+ yellow (lichexanthone), usually appearing
pruinose; ascocarp often incompletely divided into

separate chambers (similar to Astrothelium); spores

 

4-C8118d, 20-25 X 7.5-10 p. .00....00000000....0.0 E0 OChrOleucm

 

4. mallus UV- ...00.1.0.0.........OOOOOOOOOOOO0.0.0.0000... 5

Ascocarps not immersed in thallus or pseudostromata,
black with whitish indented area around ostiole; spores
4-CElled, 20-26 X 6-8 n («3011115011, 1959) 0.00.000000.0000 E0 trOBicum

Ascocarp immersed in thallus or pseudostromata ............. 6

6. Spores 4-CellEd ..........0.....0.........OOOOOOOCOOCCOO 7

6. spores up to 1.2-celled 00.0.00.........OOOOOOOOOOOOOOOOO 8

 

185
7. Spores large, 35-43 x 12-15 u; thallus usually yellowish
or yellowish green; cortex-like layer of thallus extend-
ing beneath the ascocarp ............................. T. floridanum
7. Spores smaller, 20-27 x (7-)8-10 p; thallus usually
greenish to brownish; cortex-like layer of thallus not
extending beneath ascocarp; ascocarp wall often little

carbonized ...................................... Trypethelium sp. 1

 

8. Spores up to 12-celled, 38-52 x 7-10 0 (Johnson,

1959); thallus well developed or not, lacking any

obvious layer of whitish crystals .................... T. virens
8. Spores 6-7-celled, 39-45 x 10-12 p; thallus well

developed, with a layer of whitish crystals . Trypethelium sp. 2

 

Trypethelium aeneum (Eschw.) Zahlbr.

 

in Engler & Prantl, Natfirl. Pflanzenfam. l(l‘): 70. 1903.

Verrucaria aenea Eschw. in Mart., Icon. Pl. Cryptog. 2: 15, pl. 8, f. 3.

 

1828. Holotype: Brazil, Cayeté, Martius (M).

This species, like Pyrenula cerina Eschw. and some species of

 

Anthracotheciwn,producesperietin in the upper layers of the thallus.

 

The thallus may be completely orange but is often greenish with just a
few orangish patches. It is not uncommon in Florida and I have also seen

specimens from Brazil and Guatemala.

Trypethelium eluteriae Spreng.

 

Anleitung Kenntn. Gewachse 3: 351. 1804. Type collection not seen.

The pseudostromata filled with anthraquinone crystals (mainly

 

186

parietin) and the multilocular spores are diagnostic. Trypethelium
eluteriae is common in Florida, especially southward, and is less common
in Louisiana. I have also examined specimens from Mexico, the West Indies,

Brazil, Java and Australia.

Trypethelium floridanum (Zahlbr. ex Choisy) R. C. Harris comb. nov.
Astrothelium floridanum Zahlbr. gx_Choisy, Icon. Lich. Univ. pl. 5.

1928. Zahlbr. Lich. Rar. Exs. 241. 1927. nom. nud. Type collection:

 

(Zahlbr. Lich. Rar. 241) Florida, Miami, Plitt. Isotype in MICH.

Although Zahlbruckner never validly published Astrothelium floridanum,
I accept Choisy's illustration, which shows ascocarp cross-section, ascus,
paraphyses and spores with labels and scale, as valid publication. All
of the North American records of Pseudopyrenula pupula are misidentifica-
tions and are referable to this species. The ascocarps are usually
rather scattered and not aggregated into pseudostromata. The yellowish
thallus and white ring around the ostiole characterize the T. annulare
group to which 2. floridanum belongs, as does the continuation of the
cortex-like layer below the ascocarp. Although T. floridanum closely
resembles Fee's figure of T. annulare, Muller (1888) reports larger
spores (50-70 X 16-25 p) in Fee's material. Trypethelium floridanum is
rather common in Florida and a collection from British Honduras apparently

belongs here also.

Trypethelium mastoideum (Ach.) Ach.

Lich. Univers. 307, pl. 4, f. 9. 1810. Bathelium mastoideum Ach.,

Meth. Lich. 111. 1803. Type collection not seen.

187

Trypethelium scoria Fee, Essai Crypt. Ecorc. Off. 69. 1824.

 

Original material: "in America ad corticem Crotonis Cascarillae (G).

The pseudostromata of T. mastoideum are only slightly raised, brown-
ish outside and pale yellow within. This is one of the most common species

of Trypethelium in North America. It ranges from Virginia and Kentucky

 

to Oklahoma and southward to Florida and Texas. Specimens have also been

seen from Mexico and Cuba.

Trypethelium ochroleucum (Eschw.) Nyl.

 

Flora 52: 126. 1869. Verrucaria ochroleuca Eschw. in Mart., Icon.

 

P1. Cryptog. 2: 16, pl. 8, f. 3-4. 1828. Lectotype: Brazil, Cayeté,
Martius (M, with holotype of T. aeneum).

Trypethelium pallescens Fée, Ann. Sci. Nat. 23: 440, pl. 13, f. 3,

 

A-C. 1831. Lectotype (Mull. Arg., 1888): "Surinam ad corticem arboris
ignotae" (G). Isolectotype in G.

Trypethelium erubescens Kunze ex Fee, Ann. Sci. Nat. 23: 441, pl. 14,

 

f. l. 1831. Lectotype: "Surinam ad corticem arboris ignotae" (G, with
isolectotype of T. pallescens).

Pseudopyrenula portoricensis Hedrick, Mycologia 22: 248. 1930.

 

Holotype: Puerto Rico, near Mayaguez, Fink 1025 (MICH).

This is the only species of Trypethelium in North America which

 

produces lichexanthone. This very common species ranges from Tennessee
to Florida and Texas, also widespread in the West Indies and known to me

from Brazil and Paraguay.

 

188

Trypethelium virens Tuck. gx_Michen.
.in_w. Darl., F1. Cest. ed. 3. 453. 1853. Type collection not seen.

Trypethelium exocanthum Thck. ex_Nyl., Ann. Sci. Nat. Bot., ser. 4,

 

20: 258. 1863. Holotype: Louisiana, Hale (H-NYL 177).

See Harris (1973) for description.

I previously attributed this species to Tuckerman, but Darlington's
footnote on page 431 makes it clear that Michener supplied the section
on lichens. Also, on rereading, the descriptions seem different in style
from Tuckerman's.

Trypethelium virens is common throughout the eastern United States

 

and has not been found elsewhere as yet. In the northernmost part of its

range it often produces only pycnidia.

Trypethelium sp. 1

 

The rather widely scattered, immersed ascocarps and brownish or
greenish thallus distinguish this species. Malme distributed material of

this species as_T. catervarium (Fee) Tuck. (Malme Austroam. 30).

 

However, I have examined the original material Of.2° catervarium and it
is a depauperate specimen of Astrothelium variolosum (Ach.) Mfill. Arg.

Muller (1888) acknowledged the poor condition of the specimen but chose
to use the name anyway. He cited a number of synonyms, but I have not

yet had a chance to examine their types. I have seen specimens from

Florida, Texas, the West Indies and Brazil.

Trypethelium sp. 2

 

Superficially this species is similar to T. annulare but differs in

 

189

the spores, the lack of the cortex-like layer below the ascocarp and the
presence of a layer of whitish crystals in the thallus. It is known from

a single collection in Florida by Rapp.

3. UNDESCRIBED GENUS WITH PARATHELIOID ASCOCARPS

This genus is analogous to Campylothelium but with the spores only

 

transversely septate instead of muriform. Parts of the spore wall are

IKI+ violet. This reaction is found sporadically in other members of the

 

Trypetheliaceae, especially Laurera. The rather large (44-57 x 11-14 0),

8-celled spores might lead to confusion with Trypethelium virens, if the

 

eccentric ostiole is not noticed. I have seen one collection each from

Louisiana and North Carolina.

D. FAMILY UNCERTAIN

1. THELOPSIS Nyl.

Mém. Soc. Sci. Nat. Cherbourg 3: 194. 1855. nom. cons. (syn. prius

Sychnogonia K6rb., Syst. Lich. Germ. 332. 1855.). Holotype: ‘T. rubella

 

Nyl.

See VEZda (1968) for descriptions and synonymies. Thelopsis is the
only North American genus of pyrenolichens with multisporous asci. The

key is modified from Vézda (1968).

190

l. Ascocarp remaining covered by the thallus; spores 2-celled,
lZ‘lSXS'8u 00.000000000000000.0.0.000000000000000000000 2. iSiaca
l. Ascocarp more or less free from the thallus; spores 4-

celled or submuriform 000......O..........OOOOOOOOOOOIOOOOOO 2

2. Growing on mosses or plant remains; ascocarps black
and roughened; spores 4-celled, 11-20 x 4-7 p .... E} melathelia

2. Growing on bark; ascocarps reddish ..................... 3

30 Spores 4-6-CellEd, 12-18(-2l) X 5-6 1.1 0.0.00...0..0.0.000 20 [11138118
3. Spores submuriform with 3 transverse septa and 1-4 long-

itudinal septa, 9-14 X 5-7 p .............O........... 2. inordinata

Thelopsis inordinata Nyl.

Flora 50: 9. 1867. Type collection not seen.

This species, recognized by its submuriform spores and reddish asco-
carps, was previously known only from the type collection (VEZda, 1968).

I have seen one collection each from Tennessee and Louisiana.

Thelopsis isiaca Stizenb.
Ber. Thatigk. St. Gallishen Naturwiss. Ges. 1893-94: 262. 1895.

Type collection not seen.

The whitish or slightly yellowish immersed ascocarps distinguish T.

isiaca. It is known from California.

Thelopsis melathelia Nyl.

 

Flora 47: 358. 1864. Type collection not seen.

 

191

Thelopsis melathelia is the only species in the genus growing on

 

mosses and detritus. The species has not been previously reported from
North America and the single collection by Lowe from Isle Royale,

Michigan, is typical of the species.

Thelopsis rubella Nyl.

 

Mém. Soc. Sci. Nat. Cherbourg 3: 200. 1855. Type collection not seen.

Thelopsis rubella normally has only 4-celled spores. The single

 

North American collection from Texas differs in that many of the spores
become 6-ce11ed. However, it is otherwise identical to European collec-

tions of T. rubella. The species is new to North America.

VII. EXCLUDED TAXA

A. Excluded Generic Names

Celothelium Massal., Atti R. Ist. Venet., ser. 3, 5: 331. 1860.

 

This genus is treated as a synonym of Tomasellia by Zahlbruckner
(1921-22). Since I have not been able to examine material of the type

species I do not wish to comment on its disposition.

Inoderma (Ach.) S. F. Gray, A Natural Arrang. Brit. Plants 1: 498. 1821.

Verrucaria subg. Inoderma Ach., Lich. Univ. 294. 1810.

 

Although referred in part in Ainsworth (1971) to Arthopyrenia,
it has been lectotypified by Choisy (1954) with Verrucaria epigaea Ach.

and therefore is a synonym of Thrombium.

Leiophloea (Ach.) S. F. Gray, A Natural Arrang. Brit. Plants 1: 495. 1821.

 

Verrucaria subg. Leiophloeia Ach., Meth. Lich. Suppl. 24. 1803.

 

As currently lectotypified (Riedl, 1962), Leiophloea is a nomen-

clatural synonym of Porina (see discussion, p. 37).

,Magmopsis Nyl., Flora 58: 102. 1878.

Although listed by Zahlbruckner (1921-22) as a questionable

synonym of Arthopyrenia, Nylander related it to the Pyrenopsidaceae

 

in his original description and Willey (1892) transferred the type

species (M. pertenella Nyl.) to Pyrenopsis where Zahlbruckner retained

 

it. I have seen no material.

192

193

Mesopyrenia Choisy, Bull. Soc. Bot. France 78: 456. 1931.

 

There is no indication whether this is an elevation in rank of

Arthopyrenia sect. Mesopyrenia Mull. Arg. or whether it is a new genus

 

with a generico-specific description. In either case I am unable to

place the name at this time.

Prototylium Choisy, Icon. Lich. Univ., fasc. 2, pl. 25. 1929? (drawn

 

5 Nov. 1928).

Apparentlyintended as a genus similar to Pseudopyrenula except

 

that the conidia were ovate rather than rod-like. Since no species

were included I am unable to place the genus.

Pseudopyrenia Keissl.

 

This name is listed in Ainsworth (1971) as a synonym of Artho-

pyrenia and presumably is a typographic error for Pseudarthopyrenia

 

Keissl. (=_Pyrenocollema).

 

Pyrenillium Clem, Genera of Fungi 41. 1909.

 

Under the present nomenclatural rules concerning superfluous names

(Stafleu et al., 1972), Pyrenillium is a synonym of Porina (see discus-

 

sion, p. 37).

Syngenosorus Trev., Conspect. Verruc. 15. 1860.

 

Zahlbruckner lists this genus as a synonym of Tomasellia. There

 

were two species originally included in the genus, but one was with a
question mark. The unquestioned species (and logical lectotype),

S, arthoniellus (Nyl.) Trev. [Melanotheca arthoniella Nyl.] has

 

 

194

muriform spores (Riedl, 1962) and thus cannot be referred to Tomasellia.

 

Trichotrema Clem., Genera of Fungi 41. 1909.

 

This genus is placed in synonymy with Pleurotrema in Zahlbruckner

 

(1921-22). It is unknown to me but would seem to be excluded from my

concept of Pleurotrema by the long acicular spores and is perhaps better

 

placed in or near Leptorhaphis.

 

B. Excluded Species Names

Arthopyrenia rappii Zahlbr., Ann. Mycol. 33: 34. 1935.

 

The type specimen was not available.

Leiophloea clypeata Riedl, Sydowia 16: 267. 1963.

The type specimen was not available.

flycoporellum epistigmellum Nyl. in Hasse, Lich. Calif. ed. 2,
Addenda. 1898.
This species is a lichen parasite and synonymous with Pharcidia

dispersa (Lahm) Wint. according to Keissler (1930).

VIII.

DISPOSITION OF THE PYRENOLICHEN NAMES INCLUDED IN HALE AND

CULBERSON (1970) ACCORDING TO THE TREATMENT IN THIS WORK

ARTHOPYRENIA

atractospora Zahlbr.

 

alba (Schrad.) Zahlbr.
ambigua Zahlbr.
analepta (Ach.) Mass.

analeptella (Nyl.) Arn.

 

bifera Zahlbr.
cerasi (Schrad.) Mass.

cinchonae (Ach.) Mfill. Arg.

cinereopruinosa (Schaer.) Massal.

 

conformis“(Nyl.) Mull. Arg.
conoidea (Fr.) Zahlbr.
dimidiata Fink

distans (Wi11.) Zahlbr.

epidermidis (DC.) Mass.

 

fallax (Nyl.) Arn.
finkii Zahlbr.
gemmata (Ach.) Mass.

halodytes (Nyl.) Arn.

hyalospora (Nyl.) Fink

 

leucochlora Mfill. Arg.

 

macrocarpa (K6rb.) Zahlbr.

 

195

excluded as misidentifications

= Anisomeridium

 

mostly = Arthopyrenia padi

 

excluded as misidentifications

excluded as misidentifications

= Anisomeridium biforme ?

 

= Acrocordia

 

= Anisomeridium carinthiacum
= Anisomeridium
excluded, see Harris (1973)

= Arthopyrenia lapponina

 

= Acrocordia megalospora
excluded as misidentifications

= Pyrenocollema

 

= Plagiocarpa

 

= Anisomeridium

 

excluded as misidentifications

196

parvula Zahlbr. Anisomeridium biforme

 

ll

Arthopyrenia cinereopruinosa

pinicola (Hepp) Mass.

prospersella (Nyl.) Zahlbr. Pyrenocollema

 

punctiformis auct. Arthopyrenia padi

 

 

quinqueseptata (Nyl.) Fink Polymeridium

 

 

'0

rappi Zahlbr. , type not available
rhyponta (Ach.) Mass. excluded as misidentifications

sanfordensis Zahlbr. Anisomeridium

 

 

sphaeroides auct. Acrocordia cavata

 

 

sublitoralis (Leight.) Arn. Pyrenocollema halodytes

 

 

subpunctiformis Nyl. Arthopyrenia atomarioides

 

 

MYCOPORELLUM

californicum Zahlbr. Tomasellia

 

 

difforme (Minks) Fink

Ox)

. type not found

hassei Zahlbr.

Tomasellia lactea

 

sparsellum (Nyl.) Mfill. Arg. Tomasellia

 

PORINA
carpinea (Pers.) Zahlbr.

cestrensis (Michen.) Mfill. Arg.

 

chlorotica (Ach.) Mfill. Arg.

 

cinerea (Pers.) Zahlbr. excluded as misidentifications
hibernica P. James & Swinsc.

lectissima (Fr.) Zahlbr.

 

mamillosa (Th. Fr.) Vain.

mastoidea (Ach.) Mull. Arg. excluded as misidentifications

197

nucula Ach.
olivacea (Pers.) A. L. Sm.
plumbaria (Stizenb.) Hasse

rhaphidosperma M811. Arg.

 

salicina Mfill. Arg.

subcinerea (Nyl.) Zahlbr.

 

thaxteri Sant.

viridiseda (Nyl.) Zahlbr.

 

PSEUDOPYRENULA

pupula (Ach.) Mull. Arg.

PYRENULA

herrei Fink

THELOPSIS
isiaca Stizenb.

subporinella Nyl.

 

TRYPETHELIUM
aeneum (Eschw.) Zahlbr.

catervarium (Fee) Tuck.

 

eluteriae Spreng.

exocanthum Thck.

 

mastoideum Ach.

 

ochroleucum (Eschw.) Nyl.

 

scorites Tuck.

excluded as misidentifications

= Arthopyrenia

 

?, type not seen

= Polymeridium

 

= Strigula

N. Amer. records = Trypethelium
floridanum

 

 

Arthopyreniagplumbaria

 

.T. isiaca, fid. VEZda (1969)

N. Amer. records = Trypethelium sp.

 

_T_. virens

?, type not seen

198

tropicum (Ach.) Mfill. Arg.

virens Tuck. ex Michen.

IX. ALPHABETICAL LIST OF EXSICCATI EXAMINED

Anzi It. 3

Anzi Lang. 347
471

Anzi Ven.

Arn. 1262
1573

Arn. Mon.

Britz. 51
185
218
385
386
388

Claud. 50

Cum. I: 18
24

II: 1

1

Desm. ser.

Erb. II: 1
9

Fellm. 221
222

Fr. 242
243
244
274

Harm. Loth. 450

85

121
129
130
132

420
449

0
9
10
89

II.

20
30

Arthopyrenia fraxini

Arthopyrenia lapponina
Arthopyrenia rhyponta

Arthopyrenia rhyponta
Arthopyrenia cinereopruinosa
Arthopyrenia cerasi
Anisomeridium biforme

Arthopyrenia cinereopruinosa
Tomasellia arthonioides

Arthopyrenia cinereopruinosa
N I!

Arthopyrenia cinereopruinosa
Arthopyrenia padi
n ’9
Arthopyrenia cinereopruinosa
" " , A. padi
Arthopyrenia padi

Arthopyrenia lapponina
Arthopyrenia lyrata
Mycoglaena quercicola
Arthopyrenia lyrata
Mycoglaena quercicola

Anisomeridium biforme

Arthopyrenia salicis
Arthopyrenia lapponina

Arthopyrenia lapponina
Arthopyrenia fraxini

Arthopyrenia fraxini

Arthopyrenia padi

Arthopyrenia lapponina, A. megalospora
Acrocordia conoidea

Arthopyrenia cerasi

199

Hepp 105-107
450-452
453
455—456
457
458
464
953
954

Kern. 2774
3136
3534

Krypt. Vind. 68, 68b

180

268a
268b

269

375

468

468b
468c
469, 469b
861

1021
1356
1523
1646
1763a, b
2152
3141
3477

Leight. 31
100
197
288
344

Lich. Colo. 67

Lojk. Univ. 199
279

Magn. 127
242
377

Malbr. 97
199
250
397

200

Arthopyrenia cinereopruinosa
Arthopyrenia lapponina
Arthopyrenia fraxini, A. lapponina
Arthopyrenia cinereopruinosa
Arthopyrenia cerasi

Strigula affinis

Strigula stigmatella

Anisomeridium biforme

Arthopyrenia antecellens

Arthopyrenia padi
Arthopyrenia rhyponta
Arthopyrenia salicis

Tomasellia arthonioides
Strigula stigmatella
Arthopyrenia lapponina
Arthopyrenia fraxini
Arthopyrenia cinereopruinosa
Anisomeridium biforme
Arthopyrenia salicis
Arthopyrenia padi
Arthopyrenia salicis
Pyrenocollema halodytes
Mycoglaena myricae
Arthopyrenia rhyponta
Pyrenocollema tichothecioides
Arthopyrenia padi
Mycoglaena meridionalis
Arthopyrenia lapponina
Acrocordia cavata

Strigula phaea

Mycoglaena myricae

Acrocordia conoidea
Anisomeridium biforme
Arthopyrenia cinereopruinosa

Arthopyrenia salicis
H H

Arthopyrenia padi

Strigula stigmatella
Pyrenocollema epigloea

Mycoglaena myricae
Pyrenocollema halodytes
Tomasellia gelatinosa

Arthopyrenia lapponina
Arthopyrenia cinereopruinosa
Tomasellia gelatinosa
Acrocordia conoidea

Malbr. 398
400

Mass. 43
106
127
184
185-186
197-200
201
202-203
219
298A, B
299
350

Merr. I: 147
156
164
281
289

II: 5

20
38
93

Mig. 24
74
100

Moug. 364
557

Norrl. 149
393

Nyl. Par. 91
148

Nyl. Pyr. 50

Pisut 77
151

Rab. 89
145
146
203
390
476
483
561

201

Acrocordia cavata
Arthopyrenia cerasi

Mycoglaena meridionalis

Arthopyrenia cerasi

Arthopyrenia salicis

Arthopyrenia fraxini

Arthopyrenia lapponina

Arthopyrenia cinereopruinosa
" " , A. salicis
If If

Arthopyrenia cerasi

Arthopyrenia fraxini

H I?

Strigula affinis

Anisomeridium biforme
Tomasellia californica, T. eschweileri
Anisomeridium biforme
Arthopyrenia fraxini
Strigula phaea
I! H
Arthopyrenia fraxini
Arthopyrenia lyrata
Mycoglaena meridionalis

Pyrenocollema halodytes
H I!

Mycoglaena myricae

Arthopyrenia cinereopruinosa, A. lapponina
Arthopyrenia rhyponta, Arthopyrenia padi

Arthopyrenia salicis
Arthopyrenia cerasi

Anisomeridium biforme
Arthopyrenia lapponina

Arthopyrenia cerasi

Arthopyrenia padi
Acrocordia conoidea

Acrocordia cavata
Arthopyrenia cerasi
Arthopyrenia fraxini
Arthopyrenia salicis
Arthopyrenia padi

I! It
Anisomeridium biforme
Strigula affinis

Rab. 574
579
598
623
630
658
659
704
780
943

Rel. Hasse 79
113

Rel. Suza 6
9

Rel. Tuck. 130
Samp. 22

Schaer. 287

Stenh. 89
180

Suza 122

Trev. 29
41-43

54

VEZda Bohem. 62

vezda Sel. 130

577
1229
Zahl. 190
241
Zw. 815
1197

202

Strigula sychnogonioides
Strigula stigmatella
Acrocordia conoidea

Strigula stigmatella
Arthopyrenia cinereopruinosa
Arthopyrenia fraxini
Arthopyrenia cinereopruinosa
Tomasellia arthonioides
Tomasellia gelatinosa
Arthopyrenia salicis

Anisomeridium biforme
Tomasellia lactea

Arthopyrenia lapponina
Tomasellia arthonioides

Arthopyrenia cinchonae
Arthopyrenia cerasi
Arthopyrenia cinereopruinosa, A. lapponina

Arthopyrenia lapponina
Arthopyrenia padi

Tomasellia arthonioides
Arthopyrenia cinereopruinosa
n . n
Tomasellia gelatinosa
Acrocordia conoidea
Pyrenocollema halodytes
Strigula sychnogonioides

Acrocordia cavata

Anisomeridium biforme
Trypethelium floridanum

Tomasellia arthonioides
Mycoglaena myricae

LITERATURE CITED

X. LITERATURE CITED

Ahmadjian, V. 1958. A guide for the identification of algae occurring
as lichen symbionts. Bot. Not. 111: 632-644.

. 1967. A guide to the algae occurring as lichen symbionts.
Phycologia 6: 127-160.

 

Ainsworth, G. C. 1971. Ainsworth & Bisby's Dictionary of Fungi.
Commonwealth Mycological Institute, Kew. 663 pp.

Anderson, R. A. 1967. Additions to the lichen flora of North America
- II. Bryologist 70: 339-343.

Brodo, I. M. 1968. The lichens of Long Island, New York: A vegeta-
tional and floristic analysis. New York State Mus. & Sci.
Serv. Bull. 410, 330 pp.

Chadefaud, M. 1969. Une interpretation de la paroi des ascospores
septées, notamment celles des Aglaospora et des Pleospora.
Bull. Soc. Mycol. France 85: 145-157.

 

. 1973. Les asques et la systematique des Ascomycetes.
Bull. Soc. Mycol. France 89: 127-170.

 

Choisy, M. 1954. La nomenclature générique des Endocarpaceae, Lichens
Ascomycetes. Bull. Soc. Mycol. France 70: 162-181.

Clements, F. E. 1909. The Genera of Fungi. H. W. Wilson Co.,
Minneapolis. 227 pp.

Cooke, W. B. & D. L. Hawksworth. 1970. A preliminary list of the
families proposed for fungi. Mycol. Papers 121, 86 pp.

Fink, B. 1910. The lichens of Minnesota. Contr. U. S. Natl. Mus.
14(1): 1-269.

Fries, T. 1861. Genera Heterolichenum europaea recognita. Edquist,
Upsala. 116 pp.

Hale, M. E. & W. L. Culberson. 1970. A fourth checklist of the lichens
of the continental United States and Canada. Bryologist
73: 499-543.

Harris, R. C. 1973. The corticolous pyrenolichens of the Great Lakes
region. Michigan Bot. 12: 3-68.

203

204

Harris, R. C. 1975. Belonia, Scoliocarpon and Robergea. Contr. Univ.
Michigan Herb. 11(2). [in press]

 

Herre, A. 1942. Some noteworthy lichens from Florida. Bryologist
45: 180.

Janex-Favre, M. C. 1971. Recherches sur l'ontogenie, l'organisation et
les asques de quelques Pyrenolichens. Rev. Bryol. Lichénol.
37: 428-649.

Johnson, G. T. 1959. The Trypetheliaceae of Mississippi. Mycologia
51: 741-750.

Keissler, K. 1930. Die Flechtenparasiten. In L. Rabenhorst, Kryptogamen-
Flora 8: 1-712.

. 1936-1938. Pyrenulaceae bis Mycoporaceae, Coniocarpineae.
In L. Rabenhorst, Kryptogamen-Flora. 9(1 Abt., 2 Teil):
1'8460

 

Korf, R. P. & J. K. Rogers. 1967. A new term, the schizotype, and the
concept of implicit typification. Taxon 16: 19-23.

Lanjouw, J. & F. A. Stafleu. 1964. Index Herbariorum. Regnum
Vagatabile 31, 251 pp.

Lawrence, G. et a1. 1968. Botanico-Periodicum-Huntianum. Hunt
Botanical Library, Pittsburgh. 1063 pp.

Lynge, B. 1920-1922. Index specierum et varietatum lichenum quae
collectionibus "Lichenes exsiccati" distibutae sunt.
Pars II. Nyt Mag. Naturvidensk. 57-60: 1-316.

. 1939. Index collectionum "Lichenes exsiccati" Suppl. 1.
Nyt Mag. Naturvidensk. 79: 233-323.

 

Magne, F. 1946. Anatomie et morphologie comparées des asques de
quelques Lichens. Rev. Bryol. Lichénol. 15: 204-209.

Malme, G. O. 1929. Porinae et Phylloporinae in Itinere Regnelliano
primo collectae. Ark. Bot. 23A(1): 1-37.

Massalongo, A. B. 1856. Genera Lichenum aliquot nova. Flora 39: 281-
286.

. 1860. Esame comparativo di alcune generi di licheni.
Atti R. Ist. Venet., Sci. Lett. ed Arti, ser. 3, 5: 247-276,
313-337.

 

Morgan-Jones, G. 1972. Studies on lichen asci. II. Further examples
of the bitunicate type. Lichenologist 5: 275-282.

205

Morgan-Jones, G. 1973. Endoascosporic cells in three pyrenocarpous
lichen genera. Canad. J. Bot. 51: 493-495.

, & T. D. V. Swinscow. 1965. On the genus Micrgglaena
K5rb. Lichenologist 3: 42-54.

 

 

Mfiller, E. & J. A. von Arx. 1962. Die Gattungen der didymosporen
Pyrenomyceten. Beitr. Kryptogamenfl.Schweiz 11(2): 1-922.

Mfiller, J. 1888. Pyrenocarpeae Feeanae. Mem. Soc. Phys. Geneve

Poelt, J. 1969. Bestimmungschlfissel Européischer Flechten. Cramer,
Lehre. 757 pp.

. 1974. Classification. Appendix A, p. 599-632. In
Ahmadjian, V. & M. E. Hale, [eds.]. The Lichens.
Academic Press, New York & London.

 

Printz, H. 1940. Trentepohliaceae. Nytt Mag. Naturvidensk. 80:
137-210.

Pyatt, F. B. 1974. Lichen Propagules, p. 117-145. In Ahmadjian, V. &
M. E. Hale, [eds.]. The Lichens. Academic Press, New York
& London.

Richardson, D. H. S. & G. Morgan-Jones. 1964. Studies on lichen asci.
I. The bitunicate type. Lichenologist 3: 72-83.

Riedl, H. 1962. Die Arten der Gattung Mycoporellum Mfill. Arg. sensu
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. 1963. Die Arten der Gattung Mycgporellum .... II.
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. 1971. Zur Kenntnis von Polyblastiopsis Zahlbr. und
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Rudolph, E. & R. M. Giesy. 1966. Electron microscope studies of lichen
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Santesson, J. 1970. Some occurrences of the anthraquinone parietin
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Santesson, R. 1939. Amphibious Pyrenolichens I. Ark. Bot.
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Stafleu, F.

Swinscow, T.

206

A. et a1. 1972. International Botanical Code of Nomenclature.
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D. V. 1962. Pyrenocarpous lichens: 3 The genus Porina
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Lichenologist 2: 169-171.

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Lichenologist 3: 72-85.

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Trevisan, V.

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Licheni. Spighe e Paglie, p. 1-20.

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Vainio, E.

1921. Lichenographia Fennica I. Pyrenolichenes. Acta Soc.
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Willey, H.

Tschechoslowakei. I. Folia Geobot. Phytotax. 5: 307-337.

1892. Enumeration of the Lichens found in New Bedford,
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Zahlbruckner, A. 1903-1907. Lichenes (Flechten). B. Spezieller Teil.

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207

Zahlbruckner, A. 1921-1922. Catalogus Lichenum Universalis. Vol. 1.
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INDEX TO TAXA

XI.

Acrocordia
Acrocordiaceae
Acrocordiomyces
adnexum

aeneum
aeruginella
affinis

albidum
albisedum

amaura

ambiguum
americana (Strigula)
americana (Tomasellia)
anacardii
analepta
Anisomeridium
annulata
antecellens
arthonioides
Arthopyrenia
Arthopyreniaceae
Arthopyreniella

Arthopyreniomyces

INDEX TO TAXA

34

74

37

208

Aspidothelium
Astrotheliaceae
Astrothelium
Athrismidium
atomaria
atomarioides
atractospora
Beckhausia
Belonia
bermudana
Bertossia
betulae
bifera
biforme
brachyspora
buxicola
caesia
californica
carinthiacum
catapastum
cavata
cedrina

Celothelium

26

148

El
78, 79

61

31

146

74

56

69, 7_1_
104
104

50

_12_4
80, g
M
_1_5_3_

94

46

192

cerasi
cestrensis
Chlorodothis
chlorotica

aff. chlorotica
Ciferriolichen
cinchonae
cinerea
cinereopruinosa
cinerescens
cineriseda
cinnamomeum
clypeata
comparatula
complanata
confluens (Arthopyrenia)
confluens (Pseudopyrenula)
connivens
conoidea
consequella
conspurcata
contendens
crategina
Cryptotheliaceae
cubanum

degelii

£1, 72

143

74
142
1g
194
158
13.2

2
161

138

KO
U!

126

deserticola

detergens

dimidiata

distans

Ditremis

elabens

elegans

eluteriae

epigloea

epistigmellum
erubescens

eschweileri
exasperatum
exocanthum

faginea

falklandica

fallacior

fallaciosa
fallacissima

fallax

feeanum

finkii (Anisomeridium)
finkii (Arthopyrenia)
floridana (Arthopyrenia)
floridanum (Trypethelium)

fraxini

86
47
107
ME
118

50

1g
194
187
80, 84
126.3
188
143
164

63

76

44
186

54

Geisleria
Geisleriomyces
gelatinosa
gemmata
Giacominia
Gleophragmiaceae
Gloenoblastiaceae
Gloenodictyaceae
halodytes

hassei

hederae

herrei (Arthopyrenia)
herrei (Pyrenula)
heterospora
Heufleria
horridulum
Hyalophragmiaceae
hyalospora
hypothallina
imshaugii
Inoderma
inordinata
inspersum
isabellinum
isiaca

jamesii

210

26, 131
132

80, 85
93

36

163

166

163

 

Jatteolichen 36
Jatteomyces 36
lactea 80, 86
lapponina '56
Laureraceae 26, 149
lectissima 113
aff. lectissima lZ§_
Leiophloea 37, 192
Lembidium 97
leptalea .114
Leptorhaphis 31
leucochlorum .119
leucoplaca 86
leucostoma 52
ligustri 51
Lithothelium .164
lyrata .12
macrocarpa 164
macrospora (Acrocordia) 93
macrospora (Arthopyrenia) 96
macrosporum (Anisomeridium) .112
macularis .88
Magmopsis 192
mamillosa 114
mastoideum 186
megalospora (Acrocordia) 2§_

megalospora (Arthopyrenia)
melathelia
meridionalis
Mesopyrenia
Microglaena
Microglaenaceae
microspora

minor

minus
Monoblastia
Monoblastiaceae
Mycarthopyrenia
Mycoarthopyrenia
Mycociferria
Mycoglaena
Mycoporaceae
Mycoporellum
Mycoporopsis
myricae

nigratum
nitidula (Porina)
nitidula Strigula)
Normandina
Nothostroma
nucula

aff. nucula

90, 91

90
35
36

35

34
78, 79

78, 79

182
E3.
_14_1

21, 28

78

oblongens
ochroleucum
ochrothelizum
octomera

padi
pallescens
pallidum
palmicola
Paratheliaceae
parvula
peranomala
phaea
Phyllobathelium
pinicola
Plagiocarpa
Plagiotrema
planior
planorbiculata
planorbis
platyspora
pleiomerellum
Pleosporaceae
Pleurotrema
Pleurotremataceae
plumbaria

polycarpum

187
_1_8_2_

_1_7_6

61

187

182

90, 91
163

104

75

102, 1g
26

50

1_65_

160

63

Polymeridium

polysemum 118,
Porina

Porinaceae
porospora
portoricensis
prospersella
prostans
Prototylium
Pseudarthopyrenia
Pseudopyrenia
Pseudopyrenula
Pseudosagedia
punctiformis
Pyrenillium 37,
Pyrenocollema

Pyrenulaceae 24, 27,
quercicola

quinqueseptatum
Raciborskiella

rappii

rhaphidosperma

rhyponta

rubella

Sagediomyces

salicis 61,

212

£55;
119
1E
166

69
187
_1_23

47
193
121
193
lfl
166

61
193
_1_2_1_

163

salweyi 93
sanfordense 46, 112_
Santessoniolichen 36
Santessoniomyces 36
saxicola .139
Sciodothis 78
scoria 187
sparsella §9_
Sporoschizon 33
stigmatella jig;
Strigula 1§l_
Strigulaceae 25, 29'
Strigulomyces 132
subcinereum _159
subglobosa 93
subgregaria 161
subimitans 159
submuriformis 144
subprospersella 147
subprostans .113
subpunctiformis 44
subpungens .116
sychnogonioides 26, 145_
Syngenosorus 193
tamarindi .114
taxodii 68

taylori

tenuis

cf. tetracerae
thaxteri
Telopsis
Thrombium
tichothecioides
Tomasellia
Tomaselliomyces
tremelloides
Trichotheliaceae
Trichothelium
Trichotrema
Trypetheliaceae 24,

Trypethelium

tuckeri

27,

23,

148,

125
166
319
194

180

tumida «

vainii

Verrucariaceae 23,
verrucarioides
versicolor
violascens
virella

virens

viridiseda
wetmorei
willeyanum
wilsonii
Xanthopyrenia
Xanthopyreniaceae
xylogena

zwackhii

154
147
26, 29
154
E
164
142
1_88
£5
76
_1_1_6_
1:11
121
90
97

43

FIGURES

214

Figures 1-6. Arthgpyrenia antecellens (Nyl.) Arn.

 

England, Larbalestier ZZ (H-NYL 767) [holotype]. 1. Spores.

 

2. Microconidia. 3. Ascus.‘

British Columbia, Brodo 11603 (CAN). 4. Spores. 5. Cross section

 

of ascocarp. 6. Ascus.

Figures 7-11. Arthopyrenia atomarioides Mfill. Arg.

 

Cuba, Wright Verr. Cub. II: 629 (G) [holotype]. 7. Spores.
8. Ascus.
Florida, 1887 Calkins (MICH) [isotype of Verrucaria subpunctiformis

Nyl.]. 9. Spores. 10. Ascus. 11. Cross section of ascocarp.

Figures 12-15. Arthopyrenia atractospora Zahlbr.

 

Florida, Rapp 137 (W) [holotype]. l2. Spores. l3. Microconidia.

l4. Ascus. 15. Cross section of ascocarp.

Figures l6-19. Arthopyrenia cedrina (Zahlbr.) R. C. Harris

 

Florida, Rapp 135 (W) [holotype]. 16. Spores. l7. Ascus.

18. Microconidia. 19. Cross section of ascocarp.

‘ Scale bar represents 10 p for spores, 25 p for the asci and 250 u

for the ascocarp cross section.

 

215

 

ANT ECELLENS

 

 

ATDMAHIDIDES

 

ATnACTosponA

CEDHINA

216

Figures 20-23. Arthopyrenia cerasi (Schrad.) Massal.

 

France, collector unknown (MSC). 20. Spores. 21. Microconidia.

22. Ascus. 23. Cross section of ascocarp.

Figures 24-30. Arthopyrenia cinchonae (Ach.) Mull. Arg.

 

French Guiana, Leprieur 215 (P-MONT) [holotype of Verrucaria

 

 

prostans Mont.]. 24. Spores. 25. Microconidia. 26. Cross section of
ascocarp. 27. Ascus.

Florida, Harris 2054 (MSC). 28. Spores.

 

Puerto Rico, Britton et 21. 4183 (MICH). 29. Spores.

Florida, Calkins (US). 30. Ascus.

Figures 31-35. Arthopyrenia cinereopruinosa (Schaer.) Massal.

 

England, Leight. 197 (FH). 31. Spores. 32. Microconidia.
33. Ascus. 34. Cross section of ascocarp.

Newfoundland, Waghorne 367 (US ex MO). 35. Spores.

 

217

WOOD

 

CERASI

 

 

 

 

CINEREDPHUINDBA

218

Figures 36-39. Arthopyrenia confluens R. C. Harris

"In Cort. Cascarillae ex hb. Hampe, 1877" (G) [holotype of

Tomasellia leucostoma Mfill. Arg.]. 36. Spores. 37. Microconidia. ,

38. Cross section of a group of ascocarps. 39. Ascus.

Figures 40-43. Arthopyrenia degelii R. C. Harris

 

Tennessee, ll.IX.l939 Degelius (US ex MO) [isotype].

40. Spores.

(US) [holotype]. 41. Asci. 42. Spores. 43. Cross section of ascocarp.

Figures 44-48. Arthopyrenia fraxini Massal.
Italy, ex hb. Massalongo (FH). 44. Spores.
Maine, 20.IV.1910 Merrill (FH). 45. Microconidia.

47. Ascus. 48. Cross section of ascocarp.

Figures 49-52. Arthopyrenia herrei R. C. Harris

 

California, Herre 889 (FH) [holotype]. 49. Spores.

51. Microconidia. 52. Cross section of ascocarp.

46. Spores.

50. Ascus.

219

 

 

 

FRAXINI

U

 

 

QED) - .
50 Hanna

)‘4

220

Figures 53-59. Arthopyrenia lapponina Anzi

 

Italy, Anzi Lang. 347 (FH-TUCK 4084) [isotype]. 53. Spores.
54. Microconidia. 55. Cross section of ascocarp. 56. Ascus.
New Hampshire, 1886 [Farlow] (FH). 57. Spores.

Finland, 1852 Nylander (H-NYL 955) [holotype of Arthopyrenia

 

epidermidis var. fallax Nyl.]. 58. Spores. 59. Ascus.

 

Figures 60-64. Arthopyrenia megalospora Lannr.

 

Maine, 20.X.1912 Merrill (FH). 60. Spores. 61. Ascus. 62. Cross
section of ascocarp.

Sweden, Fr. 244 (FH-TUCK 4067) [isotype]. 63. Spores. 64. Ascus.

Figures 65-68. Arthopyrenia minor R. C. Harris

 

Florida, Harris 1545 (MSC) [holotype]. 65. Spores. 66. Ascus.

 

67. Cross section of ascocarp.

Louisiana, 6.VI.1894 Langlois (US). 68. Microconidia.

221

 

A
m
N
D
p
p
A
L

 

 

MEGALDSPDRA

222

Figures 69-72. Arthopyrenia oblongens Zahlbr. ex R. C. Harris

 

Florida, IV.1930 Rapp (MICH) [holotype]. 69. Spores. 70. Ascus.

71. Cross section of ascocarp. 72. Microconidia.

Figures 73-79. Arthopyrenia padi Rabenh.

British Columbia, Brodo 12915 (CAN). 73. Ascus. 74. Spores.

 

Switzerland, Rab. 390 (MICH) [isotype]. 75. Spores. 76. Asci.
77. Cross section of ascocarp. 78. Microconidia.

Maine, Plitt 108 (MICH). 79. Spores.

Figures 80-87. Arthopyrenia planorbis (Ach.) Mull. Arg.

 

"In America ad Crotonis Cascarillae" (H-ACH) [holotype].

80. Spores.

Florida, Hale 36942 (US). 81. Spores. 82. Ascus. 83. Micro-

 

conidia.
Cuba, Wright Verr. Cub. 405 (US) [isotype of Arthopyrenia

planorbiculata Mull. Arg.]. 84. Spores. 85. Microconidia. 86. Ascus.

 

87. Cross section of ascocarp.

223

82

 

 

 

224

Figures 88-92. Arthopyrenia plumbaria (Stizenb. in Hasse) R. C. Harris
California, Herre 493 (FH). 88. Spores. IX.1912 Hasse (FH).
89. Microconidia. 90. Spores. 91. Cross section of ascocarp.

92. Ascus.

Figures 93-96. Arthopyrenia rhyponta (Ach.) Massal.

 

Sweden, collector unknown (BM-ACH 289) [isotype?]. 93. Spores.

94. Microconidia. 95. Cross section of ascocarp. 96. Ascus.

Figures 97-100. Arthopyrenia salicis Massal.
Italy, Mass. 127 (FH-TUCK 4085). 97. Spores. 98. Cross section

of ascocarp. 99. Microconidia. 100. Asci.

Figures 101-104. Arthopyrenia taxodii R. C. Harris
Texas, Ravenel §9_(FH-TUCK 4022) [holotype]. 101. Ascus.

102. Spores. 103. Cross section of ascocarp. 104. Microconidia.

1___AJ\___

225

   

 

 

 

 

 

\
y a
Q 94 96
HHYPDNTA
/®\
““73
100
103
102 5::5/

TAXDDH ‘“

226

Figures 105-108. Arthopyrenia annulata R. C. Harris

 

Florida, Britton & Britton 846 (FH-RIDD) [holotype]. 105. Spores.

 

106. Ascus. 107. Cross section of ascocarp.

Figures 109-113. Arthopyrenia bifera Zahlbr.

 

Florida, Rappiég (W) [holotype]. 109. Spores. 110. Cross section

of ascocarp. Harris 1637 (MSC). lll. Spores. 112. Microconidia.

 

113. Ascus.

227

 

 

 

228

Figures 114-118. Arthopyrenia lyrata R. C. Harris

 

Florida, Merr. II: 38 (MSC) [holotype]. 114. Microconidia.

115. Ascus. 116. Spores. 117. Cross section of ascocarp.

 

Louisiana, Tucker 7026 (LSU). 118. Spores. J

 

Figures 119-125. Mycoglaena meridionalis (Zahlbr.) Szat. }
Italy, Mass. 43 (FH-TUCK 4084). 119. Spores. 120. Microconidia.
121. Ascus.

Albania, Latzel 5890 (W) [holotype of Arthopyrenia peranomala

 

 

Zahlbr.]. 122. Spores.
Yugoslavia, Krypt. Vind. 1646 (US) [isotype]. 123. Ascus.

124. Spores. 125. Cross section of ascocarp.

Figures 126-130. Mycoglaena myricae (Nyl.) R. C. Harris

 

Michigan, Harris 7652 (MSC). 126. Spores.

 

Finland, 1861 Fellman (H) [holotype of Verrucaria fallacissima

 

Nyl.]. 127. Spores. 128. Cross section of ascocarp. 129. Microconidia.

130. Asci.

229

 

 

H7

28

 

 

 

LYRATA

 

MYHICAE

MERIDIDNALIS'

230

Figures 131-134. Mycoglaena quercicola R. C. Harris

 

Iowa, 1895 Fink (US) [holotype]. 131. Spores. 132. Microconidia.

133. Cross section of ascocarp. 134. Ascus.

Figures 135-137. Mycoglaena wetmorei R. C. Harris

 

Texas, Wetmore l9648-B (MIN) [holotype]. 135. Spores. 136. Asci.

 

137. Cross section of ascocarp.

 

Figures 138-140. Tomasellia americana (Minks ex Willey) R. C. Harris

Massachusetts, Willey 742 (FH-TUCK 3735). 138. Spores.

 

139. Asci. 140. Cross section of ascocarp.

Figures 141-144. Tomasellia arthonioides (Massal.) Massal.

 

Bulgaria, 29.V.l929 Szatala (US). 141. Spores. 142. Microconidia.

143. Cross section of ascocarp. 144. Ascus.

Figures 145-149. Tomasellia californica (Zahlbr.) R. C. Harris

 

California, Hasse 1003 (W) [holotype]. 145. Spores.

 

(FH) [isotype]. 146. Ascus. 147. Cross section of ascocarp.
148. Microconidia.

Hawaiian Islands, Heller 2051 (MSC). 149. Spores.

 

231

 

GUEHCICDLA

/’ - ‘ FM.

. f ’ 1)
5 )1. 1-1:,»
m-Ad

 

 

232

Figures 150-155. Tomasellia eschweileri (Mfill. Arg.) R. C. Harris

 

Brazil, Collector unknown (MSC) [fragment of holotype].
150. Spores. 151. Ascus.
Iowa, VIII.1895 Fink (MICH). 152. Spores. 1

Florida, Harris 2502-B (MSC). 153. Microconidia. 154. Ascus.

 

155. Cross section of ascocarp. »

Figures 156-159. Tomasellia gelatinosa (Chev.) Zahlbr.

 

France, Collector unknown (MSC). 156. Spores.
Maine, 15.IV.1910 Merrill (FH). 157. Cross section of ascocarp.

158. Ascus. 159. Spores.

Figures 160-166. Tomasellia lactea (Ach.) R. C. Harris

 

West Indies, Swartz (UPS) [isotype]. 160. Spores. 161. Ascus.
162. Cross section of ascocarp.

California, Hasse 1321 (W) [holotype of Mycoporellum hassei

 

 

Zahlbr.]. 163. Spores.

Florida, Harris 29l6-A (MSC). 164. Microconidia.

 

Brazil, Glaziou (US) [isotype of Mycoporopsis leucoplaca Mfill. Arg.].

 

165. Spores. 166. Ascus.

   

9456135181

155

 

 

162

' . 166
165

LACTEA

 

234

Figures 167-169. Tomasellia macularis (Minks ex Willey) R. C. Harris
Massachusetts, Willey 8500 (FH-TUCK 3731). 167. Spores.

168. Cross section of ascocarp. 169. Ascus.

Figures 170-173. Tomasellia sparsella (Nyl.) R. C. Harris
Columbia, Lindig 1863 (H-NYL 4292) [holotype]. 170. Spores.
171. Ascus. 172. Cross section of ascocarp.

Florida, Harris 2799 (MSC). 173. Microconidia.

Figures 174-178. Acrocordia cavata (Ach.) R. C. Harris 12 V5zda
Switzerland, [Schleicher?].4423 (H-ACH) [holotype].
174. Spores.
North Carolina, Harris 3193-B (MSC). 175. Cross section of
ascocarp. 176. Microconidia. 177. Spores.

Maine, 23.VIII.1911 Merrill (FH). 178. Ascus.

 

235

  
  

SPAHSELLA

 

 

175

((60

$176

CAVATA [
177

 

236

Figures 179-182. Acrocordia conoidea (Fr.) K6rb.

 

Michigan, Harris 9474 (MICH). 179. Spores. 180. Ascus.

 

181. Cross section of ascocarp. 182. Microconidia.

Figures 183-186. Acrocordia megalospora (Fink) R. C. Harris

 

Minnesota, Fink 209 (MIN) [isolectotype]. 183. Spores. 184. Ascus
(half normal magnification). 185. Microconidia. 186. Cross section of

ascocarp.

 

237

 

 

 

186

 

 

 

 

 

 

238

Figures 187-190. Anisomeridium adnexum (Mfill. Arg.) R. C. Harris

 

Brazil, Puiggari 240 (G) [lectotype]. 187. Spores. 188. Micro-

conidia. 189. Cross section of ascocarp. 190. Ascus.

Figures 191-194. Anisomeridium albisedum (Nyl.) R. C. Harris

 

Florida, Calkins (H-NYL 717) [lectotype]. 191. Spores.

192. Microconidia. 193. Ascus. 194. Cross section of ascocarp.

195-199. Anisomeridium ambiguum (Zahlbr.) R. C. Harris
Florida, Rapp 142 (W) [lectotype]. 195. Ascus. 196. Spores.
Harris 2200 (MSC). 197. Macroconidia. 198. Microconidia. 199. Cross

section of ascocarp.

239

 

' / 9L , \-

I89

ADNEXUM

 

 

 

199

AMBIGUUM

 

I90

 

240

Figures 200-207. Anisomeridium biforme (Borr.) R. C. Harris

California, Ha§§e_259 (FH) [isotype of Arthopyrenia parvula
Zahlbr.]. 200. Spores. 201. Microconidia. 202. Ascus.

Massachusetts, 1921 Robbins (FH). 203. Spores. 204. Cross section
of ascocarp. 205. Ascus.

British Columbia, Macoun §4_(FH). 206. Macroconidia.

England, Mudd (US). 207. Spores.

Figures 208-212. Anisomeridium carinthiacum (J. Stein.) R. C. Harris
Austria, Steiner (US ex MO) [isosyntype]. 208. Spores.

209. Ascus. 210. Cross section of ascocarp.
Connecticut, 1895 Gregg (MICH) [holotype of Arthopyrenia dimidiata

Fink ex Hedr.]. 211. Spores. 212. Microconidia.

Figures 213-218. Anisomeridium distans (Willey) R. C. Harris
Massachusetts, Willey (MICH) [isotype]. 213. Spores. 214. Ascus.

215. Microconidia.

Ohio, Moldenke 14399 (US ex MO). 216. Spores. 217. Cross section

of ascocarp. 218. Macroconidia.

241

/

204

 

207
_/
/ ( ,( >-
217“) \Z:““’ [:2
271—0*

    

. _2
67 212

CARINTHIACUM

 

\)

    
   

DISTANS Q

242

Figures 219-224. Anisomeridium feeanum (Mfill. Arg.) R. C. Harris

 

[West Indies] (G) [holotype]. 219. Spores. 220. Ascus.
221. Cross section of ascocarp. 222. Microconidia.

Florida, Thaxter 22_(FH). 223. Spores. 224. Ascus.

Figures 225-229. Anisomeridium finkii R. C. Harris

 

Puerto Rico, Fink 2052 (US) [holotype]. 225. Spores. 226. Cross
section of ascocarp. 227. Ascus.

Florida, XII.1897 Thaxter (MICH). 228. Spores. 229. Microconidia.

Figures 230-235. Anisomeridium leucochlorum (Mfill. Arg.) R. C. Harris

 

Massachusetts, Willey (US) [isotype].- 230.spores. 231. Micro-
conidia. 232. Ascus. 233. Cross section of ascocarp.

Illinois, Wolf (US). 234. Spores. 235. Macroconidia.

243

00.19 2022.220 221

FEEANUM

(0)7

-_.- -‘H-‘

226
.'22% Q2297 FINKII

"Him

233

 

 

 

   

 

‘0 O O 232
0 ©
0 23‘ @@
_EC @ 235

234 LEUCOCHLDHUM

244

Figures 236-241. Anisomeridium macrosporum R. C. Harris

Florida, Harris 279l-A (MSC) [holotype]. 236. Spores.

 

237. Longitudinal section of ascocarp. 238. Microconidia. 239. Ascus.

Harris 2785 (MSC). 240. Cross section of ascocarp. Harris 2829-C

 

(MSC). 241. Macroconidia.

Figures 242-245. Anisomeridium sanfordense (Zahlbr.) R. C. Harris
Florida, Rapp 134 (W) [holotype]. 242. Spores. 243. Ascus.

244. Microconidia. 245. Cross section of ascocarp.

245

 

 

 

240
MAcnosponum

\‘
r U" /V,. 000.
\ 243 0244 . 245
b ’2”

SANFORDENSE

 

246

Figures 246-250. Anisomeridium subprostans (Nyl.) R. C. Harris

 

South Carolina, Ravenel (H-NYL 7378) [holotype]. 246. Spores.
247. Ascus. 248. Cross section of ascocarp. 249. Microconidia.

Louisiana, 26.X.1896 Langlois (US). 250. Macroconidia.

Figures 251-256. Anisomeridium tamarindi(E%mfl R. C. Harris

 

Guadeloupe (G) [lectotype]. 251. Spores. 252. Ascus.
(G) [isolectotype]. 253. Cross section of ascocarp. 254. Ascus.

Florida, Thaxter 313 (MICH). 255. Microconidia. 256. Macroconidia.

 

Figures 257-261. Anisomeridium tuckeri R. C. Harris

 

Louisiana, Tucker 10003 (LSU) [holotype]. 257. Spores.

 

258. Microconidia. 259. Macroconidia. 260. Ascus. 261. Cross section

of ascocarp.

Figures 262-265. Anisomeridium willeyanum (R. C. Harris) R. C. Harris

 

Michigan, Harris 818 (MSC) [holotype]. 262. Spores. 263. Cross

 

section of ascocarp. 264. Microconidia. 265. Ascus.

247

 

 

:\\

256

TAMAHINDI

O m ‘0 08%258
(302490 G Q30

TSUBPRDSTANS

99:“ L11)
. /
992% @

261' ‘lr
WILLEYANUM . TUCKER!-

 

 

 

 

248

Figures 266-269. Pleurotrema anacardii (Vain.) R. C. Harris
Guadeloupe, Duss 498 (TUR) [holotype]. 266. Ascus.
267. Longitudinal section of ascocarp. 268. Spores.

Florida, Small 7598 (US). 269. Spores.

Figures 270-272. Pleurotrema inspersum Mull. Arg.

Cuba, Wright Verr. Cub. 123 (US) [isotype]. 270. Spores.

271. Ascus. 272. Longitudinal section of ascocarp.

Figures 273-274. Eyrenocollema caesia (Nyl.) R. C. Harris

France, I.1923 Crozals (US ex MO). 273. Spores. 274. Ascus.

Figures 275-278. Pyrenocollema epigloea (Nyl.) R. C. Harris
Yugoslavia, Lojk. Univ. 249 (MICH) [isotype]. 275. Spores.

276. Ascus. 277. Microconidia. 278. Cross section of ascocarp.

249

 

267

 

 

 

 

275

/m\

CAESIA EPIGLDEA

 

migf “~M.

250

Figures 279-283. Ryrenocollema halodytes (Nyl.) R. C. Harris

 

Massachusetts, 1890 Willez (H-NYL 984) [holotype of Verrucaria
conseguella Nyl.]. 279. Spores. 280. Asci. 281. Microconidia.

Rhode Island, 3l.VIII.l963 Wetmore (CAN). 282. Cross section of
ascocarp.

California, Bonar (FH) [isotype of Didymella conchae Bonar?].

 

283. Spores.

Figures 284—288. Pyrenocollema imshaugii R. C. Harris

 

California, Imshaug 17678 (MSC). 284. Spores. 285. Ascus.

 

286. Microconidia. 287. Cross section of ascocarp. 288. Symbiotic

algae.

Figures 289-292. Pyrenocollema prospersella (Nyl.) R. C. Harris

 

Illinois, Calkins (H-NYL 991) [holotype]. 289. Spores.
290. Ascus. 29l. Cross section of ascocarp. (MICH). 292. Micro—

conidia.

Figures 293-295. gyrenocollema saxicola (Massal.) R. C. Harris

Switzerland, collector unknown (FH). 293. Spores. 294. AscuuS-

295. Microconidia.

 

 

251

119 1119

 

Q § a
p 286
@3’9
QC
HALODYTES 288

287

.: IMSHAUGII

294

O
291

PHDBPEHSELLA BAXICDLA

 

 

 

252

Figures 296—300. Eyrenocollema tichothecioides (Arn.) R. C. Harris
Yugoslavia, Krypt. Vind. 1356 (US). 296. Spores. 297. Micro-
conidia. 298. Ascus.

Michigan, Harris 73l-C (MSC). 299. Spores. 300. Cross section.<3f

 

ascocarp.

Figures 301-308. Strigula affinis (Massal.) R. C. Harris

 

England, XI.l964 James (BM) [holotype of Geisleria jamesii Swirnsc:-:]o
301. Spores. 302. Macroconidia.

Louisiana, Tucker 9056-A (MSC). 303. Spores. 304. Microconidiaio

 

Haiti, Wetmore 2903 (MSC). 305. Spores. 306. Macroconidia.

 

Austria, Rab. 561 (MICH). 307. Cross section of ascocarp.

308. Ascus.

Figures 309-314. Strigula americana R. C. Harris

Iowa, 1896 EEEE.(MSC) [holotype of Arthopyrenia tenuis R. C. Hfilfxrisj'
309. Spores. 310. Microconidia. 311. Macroconidia. (MIN) [isotyIDEi] '
312. Cross section of ascocarp.

Louisiana, Tucker 9035 (LSU). 313. Ascus.

Florida, Harris 2405 (MSC). 314. Spores.

 

 

253

   
  

299

 

 

 

 

312

QCB O
03” 1

313 AMERICANA

 

254

Figure 315. Strigula complanata (Fee) Mont.

 

Florida, Harris 1598 (MSC). 315. Spores.

Figures 316-320. Strigula connivens R. C. Harris

 

Florida, Thaxter 12_(MICH) [holotype]. 316. Spores. 317. Asci.
318. Microconidia. 319. Macroconidia. 320. Cross section of joined

ascocarps.

Figures 321-322. Strigula elegans (Fee) Mull. Arg.

 

Florida, Thaxter 427 (MICH). 321. Spores.

Cuba, Ekman L-13l (MSC). 322. Macroconidia.

Figures 323-325. Strigula hypothallina R. C. Harris

 

Florida, Harris 320l-A (MSC) [holotype]. 323. Spores.

 

324. Cross section of ascocarp. 325. Ascus.

Figures 326-327. Strigula nitidula Mont.

 

Florida, XI.1922 RaEE (MICH). 326. Spores. 327. Ascus.

 

Figures 328-336. Strigula phaea (Ach.) R. C. Harris

West Indies, coll. unknown (S) [isotype]. 328. Spores.
329. Microconidia. 330. Asci. 331. Cross section of ascocarp.

Cuba, Wright Verr. Cub. 10 (MSC) [fragment of‘holotype of 3913133
cineriseda Mull. Arg.]. 332. Spores. 333. Macroconidia.

Florida, Calkins (H-NYL 718) [syntype of Verrucaria viridi§§§2;
f. albiseda Nyl.]. 334. Spores. 335. Ascus. 336. Cross section Of

ascocarp.

A

 

255

Q
a
Q

 

 

317

   

 

320
CONNIVENS

. '3
324

HYPDTHALLINA

NITIDULA ((11 (>13 9

'11:,
@Q @@ @333 8.84!@

@332 . PHAEA

     

 

 

 

 

 

 

 

 

 

Q

 

256

Figures 337-341. Strigula stigmatella (Ach.) R. C. Harris

 

Germany, Persoon (H-ACH) [lectotype]. 337. Spores. 338. Macro-
conidia.
Michigan, Harris 7534 (MSC). 339. Ascus. Harris 6052 (MSC).

340. Spores. Harris 7447 (MSC). 341. Cross section of ascocarp.

Figures 342-345. Strigula submuriformis (R. C. Harris) R. C. Harris

 

Pennsylvania, Cum. I. 299. (MSC) [holotype]. 342. Spores.
343. Cross section of ascocarp. 344. Ascus.

Minnesota, Fink 11 (MIN). 345. Macroconidia.

Figures 346-348. Strigula sychnogonioides (Nitschke) R. C. Harris

 

Germany, Rab. 574 (MICH) [isotype]. 346. Spores.
Switzerland, HeEE 938 (EM). 347. Cross section of ascocarp.

348. Ascus.

Figures 349-353. Strigula taylori (Nyl.) R. C. Harris

 

Ireland, Carroll (EM). 349. Spores. 350. Asci. 351. Micro—

conidia. 352. Macroconidia. 353. Cross section of ascocarp.

 

 

257

 

 

 

STIGMATELLA

 

 

 

 

SUBMURIFDFIMIS

 

 

 

TAYLDHI

SYCHNDGDNIDIDES

wH—‘JL

258

Figures 354-359. Strigula viridiseda (Nyl.) R. C. Harris

 

French Guiana, Melinon (H-NYL 719) [holotype]. 354. Spores.
355/ Ascus. 356. Cross section of ascocarp. 357. Microconidia.

Bermuda, Farlow (H-NYL 985) [holotype of Verrucaria bermudana

 

Nyl.]. 358. Spores. 359. Macroconidia.

Figures 360—366. Strigula wilsonii (Ridd.) R. C. Harris

 

Cuba, Britton et 21. 15288 (FH-RIDD) [holotype]. 360. Spores.
361. Microconidia.

Puerto Rico, Fink 1431 (MICH) [holotype of Porina vainii Fink

 

in Hedr.J. 362. Spores. 363. Cross section of ascocarp. 364. Macro-

conidia. 365. Ascus. (FH) [isotype]. 366. Ascus.

Figures 367-37l. Polymeridium albidum (Mull. Arg.) R. C. Harris

 

Brazil, Martius (m) [holotype]. 367. Spores. 368. Cross
section of ascocarp. 369. Ascus.

Florida, Harris 2889-A (MSC). 370. Spores. 371. Microconidia.

 

 

259

 

  

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260

Figures 372-378. Polymeridium catapastum (Nyl.) R. C. Harris

 

Colombia, Lindig 2869 (FH—TUCK 4086) [holotype]. 372. Spores.
373. Ascus.

Florida, Harris 2429-8 (MSC). 374. Cross section of ascocarp. j

 

375. Old spore. 376. Ascus. Harris 1472 (MSC). 377. Spores.

378. Microconidia.

 

Figures 379-382. Polymeridium contendens (Nyl.) R. C. Harris

 

Florida, Harris 2642 (MSC). 379. Spores.
Colombia, Lindig 2877 (H-NYL 7331) [holotype]. 380. Ascus.

381. Spores. 382. Cross section of ascocarp.

Figures 383-387. Polymeridium exasperatum R. C. Harris

 

Florida, Harris 2829-A (MSC) [holotype]. 383. Spores.

 

384. Ascus. 385. Microconidia. 386. Longitudinal section of ascocarp.

Harris 2814-8 (MSC). 387. Old spore.

 

 

261

 

 

 

382

CDNTENDENS

 

 

 

EXASPEHA‘I‘UM

 

262

Figures 388-391. Polymeridium pleiomerellum (Mfill. Arg.) R. C. Harris

 

Florida, Harris 2900 (MSC). 388. Spores. 389. Cross section of

ascocarp. 390. Ascus. 391. Microconidia.

Figures 392-395. Polymeridium quinqueseptatum (Nyl.) R. C. Harris

 

South Carolina, Ravenel (H-NYL 591) [holotype]. 392. Spores.

393. Ascus. 394. Microconidia. 395. Cross section of ascocarp.

Figures 396-399. Polymeridium subcinereum (Nyl.) R. C. Harris

 

Texas, Wright §§_(H-NYL 1529) [holotype]. 396. Spores.
397. Microconidia. 398. Cross section of ascus. (FH-TUCK 4022)

[isotype]. 399. Ascus.

Figures 400-405. Pseudopyrenula subgregaria Mull. Arg.

 

Cuba, Wright Verr. Cub. 80 (US) [isotype]. 400. Cross section
of ascocarp. 401. Spores. 402. Ascus.
Florida, Harris 1521 (MSC). 403. Ascus. 404. Spores.

Louisiana, Tucker 10005 (LSU). 405. Microconidia.

 

263

 

 

 

 

 

 

 

 

 

 

SUBCINEREUM SUBGHEGAHIA

267

 

 

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