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foliicolous species may be a general phenomenon. It is seen in the P. leptalea group (Segestria/Polycornum) and in the unrelated Thelenellaceae. Further, both the setose and non-setose species of Trichothelium share the tendency towards long, multiseptate ascospores, only very rarely becoming somewhat muriform and growing if not on leaves, in wet, even semiaquatic habitats. The presence of setae has no predictive value and even if it were to be demonstrated that the setose taxa form a monophyletic clade, I would be reluctant to recognize more than a single genus, Trichothelium. The problem of how to deal with a single species or group with a single strikingly different character has always been a problem, e.g., placement of the compound-leaved Acer negundo. It is my understanding the theory (but perhaps not the practice) of modern phylogenetic approaches is that the preponderance of evidence should outweigh any single character, e.g., the abandonment of distinctive single character lichen genera by the merger of Coelocaulon into Cetraria or the realignment of Ionaspis/Hymenelia (abandoning the photobiont character). With the exception of the foliicolous species, the distribution of Trichothelium is amphitropical. I also include Zamenhofia in Trichothelium. It was tentatively maintained by Hafellner and Kalb on the basis of lacking a chitinoid ascus ring and lacking pigments. I am very hesitant to give possible "loss mutations" generic status. Also the assay used for the pigments was the crude application of KOH, etc., and also there seems to be some disagreement , "... some species with a layer reacting K+ red" (Purvis et al., 1992). There is general agreement among recent studies that the original characters for Zamenhofia, isidia and periphyses, are meaningless at the generic level. (I do part company with McCarthy (1993) who dismissed the use of isidia at any level. I agree in a couple of the cases he cited that the "isidia" are due to abnormal growth of the photobiont but isidiate species are not uncommon, especially in Clathroporina where sterile isidiate thalli are common and isidia may be the main means of reproduction in tropical wet forests.) When it comes to Hafellner and Kalb's residual Porina, I would move in the other direction, splitting rather than lumping. Part of the impetus for this approach came from McCarthy's (1995) discovery that if one merges Clathroporina with Porina (as he would and I whole-heartedly agree), the older name is Clathroporina. The groups involved are those centered around P. lectissima, P. nucula and Clathroporina eminentior. The first, for which the name Segestria Fr. is available, can be separated by lack of oxalate crystals in thallus and ascomata, small, ± naked ascomata, small ascospores and corticolous/saxicolous species basically amphitropical. Hafellner and Kalb pointed out two species intermediate between this group and the P. nucula group. I suggest that they are actually reduced members (in adaptation to living on leaves) of the P. nucula group. The P. nucula group is separated from Clathroporina not on ascospore sepatation (transverse vs. muriform) but on thallus characters which lead to very different aspects. Clathroporina has a smooth, often shiny, highly organized thallus with a cortex, photobiont ± organized in vertical rows, a layer of oxalate crystals and a dark hypothallus. The thallus is not intimately involved with the substrate but separated by the hypothallus and often peels away becoming almost foliose. The ascomata are ± immersed, sessile or protruding from the lower side of the thallus (well illustrated by McCarthy, 1995). To Clathroporina as defined by its type, C. eminentior, I would add the P. mastoidea group, differing in having transversely septate ascospores. Hafellner and Kalb recognized Clathroporina on ascus characters: lack of chitinoid ring and presence of a constriction near the tip in young asci. I consider usage of such characters as grasping at straws but they could be right, in which case a new home would have to be found for the P. mastoidea group. Porina nucula, the type of Porina, has a less organized, often verrucose thallus lacking a dark hypothallus with sessile ascomata, emergent and often constricted at the base. Again ascospore septation is not a character as several species with muriform ascospores are referable to Porina in this restricted sense. These genera are also supported by ecology and distribution. Clathroporina is mainly a genus of wet, dark, mostly tropical forests. It is one of the few groups, like Phyllopsora and Thelotremataceae, that are adapted to the lower levels of tropical rainforest and Caribbean wet broadleaf forest. (Is the development of a hypothallus an adaptation to this habitat?) Porina may be basically subtropical, adapted to more open forests with higher light and probably less moisture. The generic definitions suggested here are rather crude but by using them as a hypothesis, additional characters can be examined in detail to evaluate their utility. It is possible that conidial characters may support the new generic arrangement. At this time there is too little data about microconidia. Macroconidia are reported (Purvis et al. 1992) for two species referred here to Trichothelium. They may also occur in Segestria leptalea but I need several more occurrences to hypothesize an anamorphic connection. Ascospore tendencies also give some support, large with a strong tendency to become muriform in Clathroporina, medium to large with little tendency to become muriform in Porina, small, few septate in Segestria, long, 160
multiseptate in Trichothelium. Ascomatal setae are confined to Segestria and Trichothelium. Clathroporina also differs in having more organized isidia, the ostiolar area is often black and there are no foliicolous species. The recognition of these four genera, Clathroporina, Porina, Segestria and Trichothelium, is somewhat analogous to generic segregation in Parmelia, distinctive in aspect (characters of the association) but ± uniform in characters of the asci and ascospores. I feel that the genera thus constituted have considerable predictive value regarding probable ascospore type, ecology and distribution. There seem to be four basic thallus/ascomata paradigms with rather continuous ascospore variation within each. This can be seen in the specific keys below where ascomata play only a small role. I simply do not understand the ascospore variation in the Trichotheliaceae. As pointed out by Aptroot and Sipman, it seems to encompass a different order of magnitude from that which mycologists are accustomed. In fact I consider this one of the distinctive features defining Trichotheliaceae. However, no one, myself included, has put the time and effort in to analyzing the situation. This leaves a situation where "species" are intuitively defined and one person's speculation is a good as another's. I consider my treatment below speculative at best and subject to considerable correction. Most of the species treated by McCarthy (1995) remain in Clathroporina under the new circumscription. Porina tahitensis (Räsänen) McCarthy and Clathroporina nuculastrum Müll. Arg. belong in Porina as defined here. Caribbean Porina firmula Müll. Arg. and P. subfirmula Riddle have very odd, placodioid thalli and may be considered for generic recognition. Two other species that may be referable to Segestria but are anomalous because of the pale pinkish ascomata and long, multiseptate ascospores and occur in eastern North America are an isidiate taxon reported from Long Island, New York reported by Brodo as Porina nucula (Porina sp. 4 in thesis), and a non-isidiate one from New Hampshire (Porina sp. 2 in thesis). I have not had time to reexamine the material, especially with regard to generic disposition, and as they are both very rare, I have decided to omit them here. KEY TO GENERA 1. Ascomata usually small, essentially naked, with at most a thin covering of thallus (occasionally immersed in substrate); ascomata and thallus lacking oxalate crystals; ascomatal wall either some shade of red (Porina-yellow, Sagedia-red) or brown to blackish, often with blue or violet tones (Pseudosagedia-violet); some species with setose ascomata; muriform ascospores very rare; considerable diversity on leaves; distribution, except for foliicolous species, amphitropical .......................................... 2 2. Ascomata blackish; wall often with blue or violet tints, without orange or red tints, KOH+ purplish or KOH-(Pseudosagedia-violet); with a very strong tendency for muitiseptate, elongate (even filiform) ascospores; setose species not uncommon........................................................................................ Trichothelium 2. Ascomata reddish, sometimes darkening to black, lacking blue or violet tints, KOH+ more intensely orange or red (Porina-yellow); ascospores short, mostly 3-7-septate; setose species rare .......................................................................Segestria 1. Ascomata usually rather large, surrounded by well developed thalline wart or immersed in thallus; wart and thallus containing oxalate crystals; ascomatal wall some shade of yellow or red (Porina-yellow); ascomata not setose; muriform ascospores not uncommon; few species on leaves; distribution essentially tropical and subtropical ............................................................................................................................. 3 3. Black hypothallus lacking; thallus firmly attached to substrate, poorly organized, often verrucose; ascomata superficial, often somewhat constricted at base; few muriform-spored species; some follicolous species known; mainly species of open, drier? forest or scrub habitats, perhaps mainly subtropical ...............................Porina 161
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PART 1 INTRODUCTION Sadly this intr
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...................................
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1. Ascomata immersed or emergent; n
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ascospores 18-24 × 3 µm; Louisian
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2. Ascospores 1(-2)/ascus..........
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3. Ascomata immersed or sessile, bl
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GYROSTOMUM Fr. Only the single spec
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Sarcographia labyrinthica (Ach.) M
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to expose black rim around pore; co
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4. Ascospores light to dark brown..
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2. Apothecia minute, ca. 0.2 mm tal
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three collections of C. stemonea te
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...................................
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COLLEMA Wigg. [The non-lichenized l
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PILOCARPACEAE Zahlbr. There are mor
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doubt the occurrence of this specie
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22. Podetia forming abruptly flared
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11. Isidia cylindrical ............
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12. Medulla yellow; thallus not sor
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3. Soredia farinose, not derived fr
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Lake County U. baileyi (Stirton) Za
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10. Ascospores 22-41 × 11-21 µm..
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6. Underside white or purplish gray
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PD+ dirty orangish; ascospores 4-ce
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9. Medulla PD+ orange .............
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hope, leads to more intensive study
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Chaetothyriales and a number of par
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2. Ascomata scattered; clypeus larg
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Duval County.......................
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Arthopyrenia atomarioides Müll. Ar
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ascomata, muriform ascospores) and
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1. Ascoma simple, with only a singl
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Mycoporum compositum (Massal.) R. C
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XANTHOPYRENIACEAE Zahlbr. It has be
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Pyrenocollema tichothecioides (Arn.
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The family is defined here more nar
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a basal fringe; ascospores 20-30 ×
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above). The selection of piece B is
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Arthopyrenia exasperata R. C. Harri
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immersis clypeo lato et microconidi
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9. Ascospores small, 16-22 × 8-9
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?Polyblastiopsis alboatra Zahlbr. i
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Additional specimens. FLORIDA. Coll
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5. Endospore absent or poorly devel
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1. Ascospores small, under 20 µm l
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Distopyrenis pachyospora Aptroot Ap
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Tomasellia nigrescens Müll. Arg.,
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illustration and distribution, P. c
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12. Ascospores 30-42(-53) × 11-15
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33. Ascospores 44-50(-56) × 17-24
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47. Endospore very reduced; ascospo
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celled, becoming filled with reddis
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5.VIII.1990, Kalb 23189, not seen,
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Aptroot, for which I have seen the
Page 109 and 110: The type collection has some ascosp
Page 111 and 112: Pseudopyrenula endochrysea Vainio,
Page 113 and 114: TRYPETHELIUM Sprengel 1. Thallus or
Page 115 and 116: 2). Trypethelium variolosum Ach., A
Page 117 and 118: 1. Ascomata present; pycnidia prese
Page 119 and 120: 14 above...........................
Page 121 and 122: This species was treated in 1990 as
Page 123 and 124: subclavate, 180-220 × 45-55 µm, w
Page 125 and 126: Monoblastia borinquensis R. C. Harr
Page 127 and 128: 9. Ascospores biseriate or irregula
Page 129 and 130: conidia in a single row; macroconid
Page 131 and 132: 33. Ascospores broadly elliptical,
Page 133 and 134: 50. Medulla UV-; cortex UV+ yellow;
Page 135 and 136: 69. Growing in aquatic or semiaquat
Page 137 and 138: Anisomeridium angulosum (Müll. Arg
Page 139 and 140: Arthopyrenia glaucescens Müll. Arg
Page 141 and 142: 16.5-23 µm. As I have unpacked onl
Page 143 and 144: Species distincta macropycnidiis ro
Page 145 and 146: the species or strain of the photob
Page 147 and 148: univ. 1: 188. 1921. Phyllobathelium
Page 149 and 150: and S. tagananae are transferred fr
Page 151 and 152: 1.0 mm diam.; wall black, thinner b
Page 153 and 154: this to mean Massalongo intended (i
Page 155 and 156: 5. Ascospores basically 4/ascus, 11
Page 157 and 158: Thelenella humilis R. C. Harris, sp
Page 159: Afro-montagnard Centre-oriental: en
Page 163 and 164: Pyrenula mastoidea Ach., Ges. Natur
Page 165 and 166: junk heap. These variants (sp. 6, s
Page 167 and 168: Hedwigia 67: 274. 1927. Type not se
Page 169 and 170: dark green-blue in section; on bamb
Page 171 and 172: Trichothelium isidiatum could be re
Page 173 and 174: Arthopyrenia planiuscula...........
Page 175 and 176: Distopyrenis pachyospora ..........
Page 177 and 178: Mycocalicium subtile...............
Page 179 and 180: Polyblastiopsis sphaerica..........
Page 181 and 182: Thelenella humilis.................
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