Introduction
The genus Pyrenula is a group of crustose lichens typically growing on smooth, shaded bark. It comprises c. 745 named taxa worldwide (Aptroot Reference Aptroot1991), though this includes some taxa now excluded from the genus as well as many synonyms. It is most speciose in the tropics, with 42 species in Australia (Aptroot Reference Aptroot2009) and 55 species knownfrom the small country of Costa Rica alone (Aptroot et al. Reference Aptroot, Lücking, Sipman, Umaña and Chaves2008), but with only 10 species in the whole of Europe (Smith et al. Reference Smith, Aptroot, Coppins, Fletcher, Gilbert, James and Wolseley2009), with an extra one on the Azores (Aptroot et al. Reference Aptroot, Rodrígues, Schumm, Camara, Gabriel, Borges, Costa, Cunha, Gabriel, Gonçalves, Martins, Melo, Parente, Raposeiro and Rodrigues2010). Here, 169 species are recognized in the genus. The genus Anthracothecium is a probably related group in which 155 taxa have been described, of which only five species are currently accepted (Aptroot Reference Aptroot2009).
The genera have never been monographed, but several recent regional revisions are in existence, notably for North America (Harris Reference Harris1989, Reference Harris1995), Costa Rica (Aptroot et al. Reference Aptroot, Lücking, Sipman, Umaña and Chaves2008), India (Upreti Reference Upreti1998), Papua New Guinea (Aptroot et al. Reference Aptroot, Diederich, Sérusiaux and Sipman1997), New Zealand (Galloway Reference Galloway2007), and Australia (Aptroot Reference Aptroot2009). More or less complete species lists exist from the above-mentioned countries, various European countries, as well as from the Azores (Aptroot et al. Reference Aptroot, Rodrígues, Schumm, Camara, Gabriel, Borges, Costa, Cunha, Gabriel, Gonçalves, Martins, Melo, Parente, Raposeiro and Rodrigues2010), Japan (Harada et al. Reference Harada, Okamoto and Yoshimura2004; Kashiwadani & Aptroot Reference Kashiwadani, Aptroot and Moon2009), South Korea (Moon & Aptroot Reference Moon and Aptroot2009), Hong Kong (Aptroot & Sipman Reference Aptroot and Sipman2001), Taiwan (Aptroot Reference Aptroot2003), the Seychelles (Schumm & Aptroot Reference Schumm and Aptroot2010) and Panama (Etayo & Aptroot Reference Etayo, Aptroot, Castroviejo and Ibanez2006).
While working on the Pyrenula species from such widely separated countries as Costa Rica, the Seychelles, Papua New Guinea and Australia, it was striking that a significant proportion of the flora is shared between them. There are a handful of very common and widespread species, and many more which are much rarer but also widespread. It is surprising that recently described species from one continent often turn up on other continents when the Pyrenulas of that region are studied in detail. Apparently, relatively few species are much more local and only rarely are they locally common.
In the process of making the local revisions, many types were investigated as it is usually not enough to rely only upon the original description. In addition, published reports on type studies can be used. At least some species are pantropical and could thus have been described first from a distant corner of the earth (as seen from the viewpoint of the local revision). A further reason was that most descriptions of Pyrenula and Anthracothecium species miss one or more, or even all, of the following characters, which are generally necesary for a certain identification: UV-reaction, hamathecium inspersion, shape of the ascospore lumina (especially whether there is an endospore between the end lumina and the wall) and disto- or euseptation (essential to distinguish Pyrenula and Anthracothecium s. str.). None of the publications from the 18th and 19th centuries cite any of these characters, except sometimes a description or illustration of the ascospore lumina. In the first half of the 20th century, the inspersion was often noted, but UV-reactions were only noted in the second half of the 20th century. In some cases, the type needed could not be borrowed but curators kindly checked the UV-reaction.
Still, not all types have been recently studied. In the recent papers mentioned above, or in the relatively few other modern papers dealing with these genera, less than half of the described taxa are treated in some way.
Over the years, several thousands of specimens belonging to these genera have been studied, and information about distribution and morphology, including specimens sent for identification and isotypes of described species, kept accumulating.
It remained unclear whether the recently published local treatments dealt with all accepted species, or whether a vast number of additional species exist, for which in general only the original description (often in latin, and with one or more of the key characters missing) and the type specimen (rarely more) are in existence. This paper endeavours to answer this question, by keying out all species that are known and summarizing the recognized synonymy, while adding to the synonymy all taxa that can be reliably interpreted on the basis of original observations or of published type studies. In order to make the treatment comprehensive, all taxa that are excluded from Pyrenula and Anthracothecium and its generic synonyms have also been listed. This still leaves some taxa with unknown or uncertain disposition. They are partly listed at the end but partly listed as a synonym of the most likely species, preceded by a question mark. In two cases, this hinges on only one character that is unknown (one hamathecium inspersion, the other the UV-reaction). Some of these names may take priority when clarified.
Synonyms are listed roughly by date of the basionym; all combinations and other homotypic names are grouped together; if their disposition is uncertain, they together are preceded by only one question mark. No further details are given as to place of publication, type specimens and so forth, as this would make the key much longer and less focused. New synonyms are not noted explicitly, but a few explanatory notes are added, such as where a lectotype is designated. All valid names in Pyrenula and Anthracothecium are treated, also those below species level. The treatment of invalid and illegitimate names is less comprehensive; such names are marked as ‘nom. inval.’ or ‘nom. illeg.’ respectively. A complication is that sometimes in the past there was often no intention to propose a new name. New names were sometimes the result of mistakes, and in some cases were meant to replace previous names, but a direct reference was missing as it was considered superfluous. The authors could not know that direct citation of the replaced name would be retroactively imposed upon them by future designers of the Code.
During the research, many type and other specimens have been photographed. A selection of these illustrations will be made available to all on www.tropicallichens.net.
In order to make the key more accessible, it also contains some species and genera that are often confused with Pyrenula and Anthracothecium, and for these taxa the synonymy is given as well. In fact, the key is also complete for Blastodesmia, Sulcopyrenula and Eopyrenula, but not for others such as Aptrootia, Architrypethelium, and Lithothelium, of which only the corticolous brown-spored taxa are treated.
Only six studied species were undescribed, and these are included as ‘ined.’ with a provisional name. They will be described elsewhere in the near future. A few new combinations were necessary. New combinations and new names are formalized in a separate section, nomenclatural novelties, following the key. The taxonomy of the species is mostly that of the recent published treatments, which apply a rather robust species concept. Notable exceptions to the currently accepted taxonomy are noted in the key. At first glance, the systematics of this group might seem rather schematical, with a few characters that in combination define schematic groups. However, there are often additional characters or rather trends associated with these key characters, and the accepted species seem to make sense also in terms of ecology and distribution. The known distribution is noted in each instance.
The generic concept applied here is that in current use since Harris (Reference Harris1989) and Aptroot (Reference Aptroot1991). Table 1 gives some characteristics of all genera currently accepted in the family Pyrenulaceae, as well as Granulopyrenis, a species of which turned out to group inside the family in a recent phylogenetic study (Gueidan et al. Reference Gueidan, Villaseñor, Hoog, de Gorbushina, Untereiner and Lutzoni2008). Most genera are well characterized morphologically, and all but Pyrenula contain few species. The family has so far received little attention in phylogenetic studies, and in most genera so far not a single species has been sequenced. The available phylogenetic trees (e.g. Gueidan et al. Reference Gueidan, Villaseñor, Hoog, de Gorbushina, Untereiner and Lutzoni2008) do, however, suggest that at least some of the well-characterized small genera (notably Anthracothecium, Granulopyrenis and Pyrgillus) are ingroups of Pyrenula in the current sense, making Pyrenula currently paraphyletic. A wider species sampling is needed to evaluate this. It is not inconceivable that some of the genera currently recognized in the Pyrenulaceae are ingroups of Pyrenula, but it should be noted here that the species of, for example, Pyrgillus and Mazaediothecium are not treated in the keys below because they are not pyrenocarpous. In any case, the focus of the present paper is on species and their identification, not on genera and their delimitation.
Materials and Methods
The morphology of all specimens was studied with an Olympus SZX7 stereomicroscope and an Olympus BX50 compound microscope with differential interference contrast optics. Chemistry was investigated using short-wave UV light. Names were mostly checked in the original literature.
Morphology of Pyrenula and Anthracothecium
In general, pyrenocarpous lichens are poor in key characters, as they have a very simple thallus and a uniform, carbonized ascoma wall structure. Species of Pyrenula and Anthracothecium are constant in certain characters, while they are quite plastic in others. This can be deduced for instance from the variation within large specimens that grow all around a tree, especially when this tree is at the margin of a forest or at the seashore, where situations differ on different sides.
The thallus is in close contact with the substratum, and therefore is much influenced by its differences. Significant thallus characters are the following: 1) thallus ecorticate or corticate; 2) thallus with or without pseudocyphellae (also called maculae or pockets of hyaline crystals); 3) thickness (0·3 mm is often a relevant border); 4) UV-reaction (yellow or negative; whitish reflection is scored as negative); 5) coloration by anthraquinones (KOH+ purple, is occasionally orange, red or yellow, and superficial or in the medulla). The KOH+ brown reaction often noticed in older descriptions is an artefact of an interaction with the bark. A few species have special thallus characters, such as a bullate thallus or a silvery or creamy colour. In general, thallus colour is variable within a species (and indeed within a thallus). No significant characters have been noted in the algae (all trentepohlioid) or the hyphal structure. A black prothallus is often present, but is not a reliable character. Soredia and isidia are always absent.
The ascomata are closed pseudothecia with a uniformly carbonized wall. The shape varies from conical to globose, and these extremes are fixed species characters, but more subtle differences in shape (e.g. elongation) are generally induced by the substratum. The amount of wall material underneath the ascoma is also largely correlated with the primary shape and, furthermore, substratum-induced, so it does not represent an independent key character. The presence of anthraquinones (KOH+ purple) is a useful character, just as with the thallus. A red, orange or yellow coloration can be present outside the ascomata, and red crystals are sometimes present inside. The size class of the ascomata is usually a reliable character; at least four classes can be distinguished: up to 0·4 mm diam., 0·4–0·7 mm diam., 0·7–2·0 mm diam. and over 2·0 mm diam. These measurements relate to the outward extension of the carbonization.
The organization of the ascomata on the other hand is an important, and usually constant, character. Ascomata can be simple (only aggregated as if by chance when crowded), or mostly sideways aggregatd with common walls but separate ostioles (formerly often classified in Melanotheca) or joined with partly common ostioles (species formerly mainly in Parmentaria or Pleurotheciopsis when ascospores muriform, or in Pyrenastrum when ascospores transversely septate). In the latter case, the ostioles are not apical but lateral or eccentric. Ostioles can also be eccentric to lateral while the ascomata are not joined (species formerly mainly in Parathelium). This is a reliable character for most species, but one or two species are variable in this respect. When the ostioles are skewed but all point in roughly the same direction, it is an environmentally-induced form of a species with apical ostioles, usually a collection from a slanting tree. The shape and colour of the ostioles, which has been given much attention in descriptions, is on the other hand quite variable, even within one specimen, and differences are mostly age-related.
The hamathecium can be inspersed with oil droplets or not (or only in the upper half), and there are no distinguishing characters in the filaments. Iodine reactions with IKI can be orange, blue or absent, somewhat correlated with inspersion but also depending on development. A fleeting KOH+ green reaction has been noted in one species only.
The number of ascospores per ascus is more or less constant. Asci can have the ascospores strictly uniseriate or more or less biseriate. No variation has been observed in the ascus structure and size differences are fully dependent on size and number of the ascospores.
The ascospores provide the majority of the key characters. In order to evaluate the ascospore characters, fully developed but not overmature ascospores should be investigated. Measurements should be taken from some of the larger, most coloured ascospores with intact lumina. A section often also contains many smaller, immature ascospores, as well as very large, overmature ascospores in which the lumina are starting to disintegrate. There is considerable variation in ascospore size within and between collections, more so in the species with large, muriform ascospores, where a chance reduction in the number of ascospores per ascus (one suppressed mitotic division) can cause a near doubling of ascospore size.
Other notable ascospore characters are: cilia (rare); colour (grey or brown; orange oil in postmature ascospores or not); transverse, submuriform or muriform septation; number of septa; overall shape (broadly ellipsoid, ellipsoid or fusiform to elongated); curvature (straight or bent); shape of the ends (rounded, pointed or acuminate at one or two ends); shape of the lumina (rounded, angular, diamond-shaped, some or all elongated); the position of the end lumina (directly against the exospore wall orseparated from the wall by a layer of endo spore); presence or absence of eusepta or bands of dark granules at the tips or between the lumina (not always a reliable character); or the presence of an additional layer of endospore around the lumina.
World key to the species of Pyrenula, Anthracothecium and similar genera
Ascospores submuriform to muriform ... Key A
Ascospores only transversely septate ... Key B
Key A. Corticolous pyrenocarpous lichens with brown, (sub)muriform ascospores
Synopsis for quick access (only first alternative given except at the end)
1 Hamathecium filaments distinctly branched to anastomosing ... 2
4 Hamathecium filaments unclear or absent ... Agonimia allobata
5 Young ascospores with euseptata only; mature ascospores still mainly euseptate ... ... Anthracothecium s. str., 6
10 Ascospores strongly flattened, with two rows of lumina (lozenge-shaped) ... Sulcopyrenula, 11
14 Ascospores submuriform ... 15
22 Ascomata and/or thallus or medulla with yellow, orange or red anthraquinones ... ... 23
28 Ostioles lateral ... 29
36 Ascospores <25 µm long ... 37
40 Old ascospores filled with orange oil ... Pyrenula breutelii
41 Ascospores >80 µm long, mostly 2 per ascus ... 42
Ascospores <80 µm long, mostly 4–8 per ascus ... 45
Main Key
1 Hamathecium filaments distinctly branched to anastomosing over their whole length ... 2
Hamathecium filaments mostly unbranched or unclear ... 4
2(1) Ascospores 200–330 µm long; New Zealand ... ... Aptrootia elatior (Stirt.) Aptroot
[Ascidium elatius Stirt., Leptotrema elatius (Stirt.) Müll. Arg., Thelotrema elatius (Stirt.) Hellb., Laurera elatior (Stirt.) D. J. Galloway, Anthracothecium monosporum Müll. Arg., Polyblastiopsis monospora (Müll. Arg.) Upreti & Ajay Singh, Julella monospora (Müll. Arg.) D.D. Awasthi]
Ascospores 20–40 µm long ... 3
3(2) Ascospores smooth, with pointed ends; California; ... ... Thelenella hassei (Zahlbr.) H. Magn.
(Microglaena hassei Zahlbr., Microglaena sychnogonoides Zahlbr.)
Ascospores warted, with rounded ends; pantropical ... ... Mycomicrothelia decipiens (Müll. Arg.) R.C. Harris
(Anthracothecium decipiens Müll. Arg, Bogoriella subpersicina Zahlbr., Anthracothecium corcovadense Malme, Ornatopyrenis muriformis Aptroot)
Note: The new synonym Bogoriella subpersicina Zahlbr. is the type and only species of Bogoriella Zahlbr., which antedates Mycomicrothelia Rehm by a decade, necessitating either conservation of the little-used generic name Mycomicrothelia, or a transfer of the currently accepted species to Bogoriella.
4(1) Hamathecium filaments unclear or absent ... ... Agonimia allobata (Stizenb.) P. James
[Verrucaria allobata Stizenb., Polyblastia allobata (Stizenb.) Zsch.]
Hamathecium filaments mostly unbranched ... 5
5(4) Young ascospores with euseptata only; mature ascospores still mainly euseptate ... ... Anthracothecium s. str., 6
Septation only or mostly distoseptate, with notably thickened endospore ... 10
6(5) Ascomata mostly single ... 7
Ascomata mostly aggregated with a shared ostiole ... 9
7(6) Ostiole apical ... 8
Ostiole lateral; pantropical ... ... Anthracothecium australiense (Müll. Arg.) Aptroot
[Pleurothelium australiense Müll. Arg., Pleurotheliopsis australiense (Müll. Arg.) Zahlbr., Bottaria collospora Vain., Anthracothecium collosporum (Vain.) Zahlbr., Pleurotheliopsis nanam Zahlbr., Anthracothecium nanum (Zahlbr.) R.C. Harris, Parmentaria nana (Zahlbr.) R.C. Harris, Parmentaria subastroidea var. subsimplex Müll. Arg., Astrothelium congregans Eckf. (nom. inval.), ? Verrucaria guineensis Nyl., Anthracothecium guineense (Nyl.) Müll. Arg., Pleurotheliopsis erigens Kashiw., Anthracothecium erigens (Kashiw.) H. Harada]
8(7) Ascospores 6–8 per ascus; pantropical ... ... Anthracothecium prasinum (Eschw.) R.C. Harris
[Verrucaria prasina Eschw., Acrorixis prasina (Eschw.) Trevis., Anthracothecium eschweileri Müll. Arg., Verrucaria praelustris Krempelh., Anthracothecium praelustre (Krempelh.) Müll. Arg., Trypethelium pallidum C. Knight, Parmentaria pallida (C. Knight) Shirley, Verrucaria thwaitesii Leight., Verrucaria borbonica Nyl., Sporodictyon borbonicum (Nyl.) Trevis., Anthracothecium borbonicum (Nyl.) Müll. Arg., Anthracothecium angulatum Zahlbr., Anthracothecium angulatum var. majus Zahlbr., Anthracothecium majus (Zahlbr.) Kashiw. & Kurok., Julella luzonensis P. Henn., Titanella luzonensis (P. Henn.) Syd. & P. Syd., Pleamphisphaera luzonensis (P. Henn.) Höhnel, ? Anthracothecium speciosum f. iturupiense O. B. Blum, Anthracothecium pseudoborbonicum Upreti & Ajay Singh]
Ascospores 1–4 per ascus; pantropical ... ... Anthracothecium macrosporum (Hepp) Müll. Arg.
[Verrucaria macrospora Hepp, Anthracothecium doleschallii A. Massal., Verrucaria andamanica Nyl., Anthracothecium andamanicum (Nyl.) Müll. Arg., Bottaria columellata Vain., Anthracothecium columellatum (Vain.) Zahlbr., Anthracothecium manipurense Müll. Arg., ? Verrucaria luteonitens Nyl., Anthracothecium luteonitens (Nyl.) Zahlbr., Julella dactylospora Rehm, Anthracothecium japonicum Kashiw. & Kurok., Pyrenula neojaponica H. Harada, Anthracothecium megaspermum Patw. & Makhija, Anthracothecium indicum Ajay Singh]
9(6) Ascospores 6–8 per ascus; Australasian, possibly also Africa ... ... Anthracothecium gregale (C. Knight) Aptroot
[Trypethelium gregale C. Knight, Parmentaria gregalis (C. Knight) Müll. Arg., Trypethelium subplanum C. Knight, Parmentaria subplana (C. Knight) Müll. Arg.]
Ascospores 1–2 per ascus; pantropical ... ... Anthracothecium interlatens (Nyl.) Aptroot comb. nov.
10(5) Ascospores strongly flattened, with two rows of lumina (lozenge-shaped) ... ... Sulcopyrenula, 11
Ascomata not flattened, circular in transverse section ... 14
11(10) Thallus UV− ... 12
Thallus UV+ yellow ... 13
12(11) Hamathecium inspersed; neotropical ... ... Sulcopyrenula canellae-albae (Fée) H. Harada
[Pyrenula canellae-albae Fée, Verrucaria canellae-albae (Fée) Nyl., Sporodictyon canellae-albae (Fée) Trevis., also as Sporodictyon canellae (Fée) Trevis., Anthracothecium canellae-albae (Fée) Müll. Arg, Bottaria canellae-albae (Fée) Vain., Verrucaria punctuliformis Stizenb., Anthracothecium punctuliforme (Stizenb.) Müll. Arg.]
Hamathecium not inspersed; pantropical ... ... Sulcopyrenula staurospora (Tuck.) H. Harada
[Pyrenula staurospora Tuck., Anthracothecium staurosporum (Tuck.) Zahlbr., Anthracothecium tetraspermum Riddle]
13(11) Ascospores ellipsoid, c. 2 times as long as wide; neotropical ... ... Sulcopyrenula cruciata Aptroot
Ascospores nearly globose; neotropical ... ... Sulcopyrenula subglobosa (Riddle) Aptroot comb. nov.
14(10) Ascospores submuriform, only one or a few of the median cells with longitudinal septa ... 15
Ascospores muriform (when ascospores small, logically only few septa can be present; when in doubt, start with the first alternative) ... 22
15(14) Ostiole lateral ... 16
Ostiole apical ... 18
16(15) Ascospores <35 µm long ... 17
Ascospores 45–65 µm long; neotropical ... Pyrenula erumpens R.C. Harris
17(16) Ascospores 15–22 µm long; Australasian ... ... Lithothelium hieroglyphicum Aptroot
Ascospores 24–32 µm long; Australasian ... ... Pyrenula subumbilicata (C. Knight) Aptroot
[Trypethelium subumbilicatum C. Knight, Parmentaria subumbilicata (C. Knight) Müll. Arg., Pyrenastrum knightii Müll. Arg. Pyrenula immersa Müll. Arg.]
18(15) Ascospores <35 µm long ... 19
Ascospores 34–72 µm long ... 21
19(18) Ascospores 14–23 µm long, hamathecium not inspersed; eastern palaeotropical (Australia to Japan) ... Lithothelium nanosporum (C. Knight) Aptroot
[Trypethelium nanosporum C. Knight, ? Verrucaria pusilla Ach., Anthracothecium pusillum (Ach.) Müll. Arg., Parmentaria microspora Müll. Arg., Anthracothecium laevigatum Müll. Arg., Pyrenula neolaevigata H. Harada, Lithothelium submuriforme R.C. Harris & Aptroot]
Ascospores 23–35 µm long ... 20
20(19) Ascospores 23–35 µm long, hamathecium not inspersed; eastern palaeotropical ... ... Pyrenula gibberulosa (Vain.) Aptroot comb. nov.
Ascospores 23–33 µm long, hamathecium inspersed; eastern palaeotropical (India) ... Pyrenula darjeelingensis Jagadeesh Ram & G.P. Sinha
21(18) Ascospores with pointed ends, hamathecium not inspersed; eastern palaeotropical (Australia to Japan) ... Pyrenula subvariolosa (C. Knight) Aptroot
[Anthracothecium subvariolosum C. Knight, Verrucaria subvariolosa C. Knight, Trypethelium planum C. Knight, Parmentaria plana (C. Knight) Shirley, Trypethelium umbilicatum C. Knight, Parmentaria umbilicata (C. Knight) Shirley, Bottaria umbilicata (C. Knight) Müll. Arg., Anthracothecium asahinae Kashiw. & Kurok., Pyrenula asahinae (Kashiw. & Kurok.) H. Harada]
Ascospores with rounded ends, hamathecium inspersed; neotropical ... ... Pyrenula novemseptata Vain.
[Anthracothecium novemseptatum (Vain.) R.C. Harris, Anthracothecium varians R.C. Harris]
Note: All specimens examined, including some studied by Harris, have distoseptate spores.
22(14) Ascomata and/or thallus or medulla with yellow, orange or red anthraquinones ... ... 23
Anthraquinones absent ... 28
23(22) Thallus warts internally with soft orange medulla; Philippines ... ... Pyrenula endocrocea Aptroot ined.
Anthraquinone outside ... 24
24(23) Ascomata and/or thallus red ... 25
Ascomata and/or thallus yellow to orange ... 27
25(24) Ostiole apical, hamathecium inspersed; neotropical ... ... Pyrenula cruentata (Müll. Arg.) R.C. Harris
[Bottaria cruentata Müll. Arg., Anthracothecium cruentatum (Müll. Arg.) Müll. Arg., Trypethelium cruentatum Nyl. (nom. nud.), Bottaria cruentata var. chlorotica Müll. Arg.]
Ostiole lateral, hymenium not inspersed ... 26
26(25) Ascomata partly fused with joint ostioles, ascospores with up to 6 lumina per tier; Caribbean ... Pyrenula kermesina R.C. Harris
Ascomata solitary, ascospores with up to 2 lumina per tier; Pacific ... ... Pyrenula palmarum (Krempelh.) R.C. Harris
[Verrucaria palmarum Krempelh., Anthracothecium palmarum (Krempelh.) Vain., Bottaria palmarum (Krempelh.) Vain.]
27(24) Ascospores 10–23 µm long, with 3 primary septa; pantropical ... ... Pyrenula ochraceoflava (Nyl.) R.C. Harris
[Verrucaria ochraceoflava Nyl., Anthracothecium ochraceoflavum (Nyl.) Müll. Arg., Sporodictyon ochraceoflavum (Nyl.) Trevis., Verrucaria ochraceoflava f. nudior Nyl., Anthracothecium ochraceoflavum f. nudius (Nyl.) Zahlbr., Bottaria nudior (Nyl.) Vain., Verrucaria denudata f. ochrotropa Nyl., Bottaria ochrotropa (Nyl.) Vain., Anthracothecium denudatum var. ochrotropum (Nyl.) Zahlbr., Anthracothecium ochrotropum (Nyl.) Zahlbr., Bottaria ochraceoflava var. ochrotropa (Nyl.) Vain., Verrucaria albescens ssp. subochracea Nyl., Bottaria subochracea (Nyl.) Vain., Bottaria albescens ssp. subochracea (Nyl.) Vain., Anthracothecium albescens var. subochraceum (Nyl.) Zahlbr., Anthracothecium subochraceum (Nyl.) Zahlbr., Anthracothecium coccineum Müll. Arg., Bottaria coccinea (Müll. Arg.) Vain., Verrucaria vitellina Stizenb., Anthracothecium vitellinum (Stizenb.) Müll. Arg., Anthracothecium ochroxanthum Müll. Arg., Bottaria ochrotropa var. cruenta Räsänen, Bottaria ochrotropa var. cinnamomea Räsänen]
Note: Specimens with small ascospores with two layers of lumina are ... ... Pyrenula ochraceoflava var. pacifica P.M. McCarthy
[Bottaria rosea Vain., Anthracothecium roseum (Vain.) Zahlbr.]
Ascospores 23–35 µm long, with 5–7 primary septa; neotropical ... ... Pyrenula ochraceoflavens (Nyl.) R.C. Harris
[Verrucaria ochraceoflavens Nyl., Anthracothecium ochraceoflavens (Nyl.) Zahlbr., Bottaria ochraceoflava ssp. ochraceoflavens (Nyl.) Vain.]
28(22) Ostioles lateral ... 29
Ostioles apical (here species of the non-pyrenocarpous genus Leptotrema would key out also, characterized by stiff paraphyses in a dense layer) ... 36
29(28) Ascospores <70 µm long ... 30
Ascospores >70 µm long ... 32
30(29) Ascomata at least partly fused with a more or less joint ostiole ... 31
Ascomata all single, ascospores 15–33 µm long; eastern palaeotropical (Andaman Islands) ... Pyrenula microspora (Nagarkar & Patw.) Upreti
(Pleurothelium microsporum Nagarkar & Patw., Pleurotheliopsis andamanensis Ajay Singh & Upreti, Pyrenula subandamanica Upreti)
31(30) Ascospores 25–45 µm long; pantropical ... ... Pyrenula astroidea (Fée) R.C. Harris
[Parmentaria astroidea Fée, Verrucaria aspistea var. astroidea (Fée) Nyl., Pyrenastrum astroideum (Fée) Eschw., Verrucaria astroidea (Fée) Nyl., Parmentaria cinchonarum Fée, Verrucaria libricola var. cinchonarum (Fée) Nyl., ? Pyrenastrum album Eschw., Pyrenastrum album ssp. coronatum (Eschw.) Eschw., Pyrenastrum americanum Spreng., Pyrenastrum astroideum var. duplicatum Nyl., Heufleridium pentagastricum Müll. Arg., Parmentaria baileyana Müll. Arg., also as Parmentaria baileyi Müll. Arg., Parmentaria zenkeri Müll. Arg., Pleurotheliopsis asahinae Zahlbr., Parmentaria pluricarpa Ajay Singh, Pyrenula pluricarpa (Ajay Singh) Upreti, Parmentaria mamillata Ajay Singh, Pyrenula mamillata (Ajay Singh) Upreti, Anthracothecium immersum Patw. & Makhija, Parmentaria immersa (Patw. & Makhija) Ajay Singh, Pyrenula karnatakensis Upreti, Parmentaria andamanica Upreti & Ajay Singh, Pyrenula bicarpa Upreti]
Ascospores 45–70 µm long; pantropical ... ... Pyrenula ravenelii (Tuck.) R.C. Harris
[Pyrenastrum ravenelii Tuck., Parmentaria ravenelii (Tuck.) Müll. Arg., Pyrenastrum gemmeum Tuck., Verrucaria pyrenastroides C. Knight, Astrothelium pyrenastroides (C. Knight) C. Knight, Parmentaria pyrenastroides (C. Knight) Müll. Arg., Pyrenula pyrenastroides (C. Knight) D.J. Galloway, Astrothelium prostratum Stirt., Parmentaria prostrata (Stirt.)Müll.Arg., Heufleridium prostratum (Stirt.) Müll.Arg., Parmentaria consanguinea Müll. Arg.,? Pleurothelium inclinatum Müll. Arg., Anthracothecium inclinatum (Müll. Arg.) Kashiw., Parmentaria prostrata (Stirt.) Müll.Arg., Pyrenula prostrata (Stirt.) D. J. Galloway, Astrothelium ochrocleistum Nyl., Bathelium megaspermum var. tasmanicum Jatta, ? Parmentaria chevallieri B. de Lesd., Laurera megasperma var. tasmanica (Jatta) Zahlbr., Parmentaria ceylenensis Upreti & Ajay Singh, Pyrenula nova-granadensis Upreti & Ajay Singh]
32(29) Ascospores 2 per ascus, 135–200 µm long; pantropical ... ... Pyrenula lyoni (Zahlbr.) Aptroot comb. nov.
Ascospores 4–8 per ascus ... 33
33(32) Ascospores 100–135 µm long ... 34
Ascospores 50–110 µm long ... 35
34(33) Ascomata mostly fused; pantropical ... ... Pyrenula schiffneri (Zahlbr.) Aptroot comb. nov.
Ascomata rarely fused; neotropical ... Pyrenula chilensis (Fée) R.C. Harris
(Parmentaria chilensis Fée)
35(33) Ascospores 70–110 µm long; N. Atlantic (W. Europe & Macaronesia) ... ... Pyrenula hibernica (Nyl.) Aptroot
[Verrucaria hibernica Nyl., Verrucaria pyrenuloides var. hibernica (Nyl.) Carroll, Anthracothecium hibernicum (Nyl.) A.L. Sm., Polyblastia hibernica (Nyl.) Arnold]
Ascospores 50–90 µm long; eastern palaeotropical (India) ... ... Pyrenula ceylonensis (Ajay Singh & Upreti) Aptroot comb. nov.
36(28) Ascospores <25 µm long ... 37
Ascospores >25 µm long ... 40
37(36) Hamathecium inspersed, ascospores 20–25 µm long; Borneo ... ... Pyrenula borneensis Aptroot ined.
Hamathecium not inspersed ... 38
38(37) Thallus UV+yellow; pantropical ... Pyrenula confinis (Nyl.) R.C. Harris
[Verrucaria confinis Nyl., Sporodictyon confine (Nyl.) Trevis., Anthracothecium confine (Nyl.) Müll. Arg., Bottaria confinis (Nyl.) Vain., Anthracothecium corticatum Müll. Arg., Pyrenula corticata (Müll. Arg.) R. C. Harris, Verrucaria albescens Nyl., Sporodictyon albescens (Nyl.) Trevis., Anthracothecium albescens (Nyl.) Müll. Arg., Bottaria connectens Vain.]
Thallus UV− ... 39
39(38) Ascospores 15–22 µm long; pantropical ... ... Pyrenula parvinuclea (Meyen & Flot.) Aptroot
[Verrucaria parvinuclea Meyen & Flot., Anthracothecium parvinucleum (Meyen & Flot.) Zahlbr., Bottaria parvinuclea (Meyen & Flot.) Vain., Verrucaria denudata Nyl., Sporodictyon denudatum (Nyl.) Trevis., Anthracothecium denudatum (Nyl.) Zahlbr, Bottaria denudata (Nyl.) Vain., Bottaria parameroides Vain., Anthracothecium parameroides (Vain.) Müll. Arg., Bottaria subconnectens Vain., Anthracothecium awasthii Ajay Singh, Anthracothecium angolense Upreti & Ajay Singh]
Ascospores 7–10 µm long; India ... Pyrenula nanospora (Ajay Singh) Upreti
(Anthracothecium nanosporum Ajay Singh)
40(36) Old ascospores filled with orange oil ... ... Pyrenula breutelii (Müll. Arg.) Aptroot comb. nov.
Old ascospores without orange oil ... 41
41(40) Ascospores >80 µm long, mostly 2/ascus ... 42
Ascospores <80 µm long, mostly 4–8/ascus ... 45
42(41) Hamathecium inspersed, ascospores 90–200 µm long; pantropical ... ... Pyrenula globifera (Eschw.) Aptroot
[Verrucaria globifera Eschw., Sporodictyon globiferum (Eschw.) Trevis., Anthracothecium globiferum (Eschw.) Müll. Arg., ? Pyrenula variolosa Pers., Anthracothecium variolosum (Pers.) Müll. Arg., Verrucaria variolosa (Pers.) Mont., Polyblastia variolosa (Pers.) Trevis., Sporodictyon variolosum (Pers.) Trevis., Bottaria variolosa (Pers.) Vain., Verrucaria epapillata Nyl., Sporodictyon epapillatum (Nyl.) Trevis., Anthracothecium epapillatum (Nyl.) Müll. Arg., Bottaria epapillata (Nyl.) Vain., Verrucaria operta Nyl., Anthracothecium opertum (Nyl.) Müll. Arg., Anthracothecium paraguayense Malme, ? Anthracothecium globiferum var. microsporum Ajay Singh, Anthracothecium austroindicum Ajay Singh]
Note: neotropical specimens are rarely UV+ yellow. Such specimens are possibly worth recognition at species level. For the time being, the UV− reaction of specimens of this species should be noted.
Hamathecium not inspersed ... 43
43(42) Thallus without pseudocyphellae, ascospores 80–140(–155) µm long; pantropical ... Pyrenula platystoma (Müll. Arg.) Aptroot comb. nov.
Thallus with pseudocyphellae ... 44
44(43) Ascospores 80–110 µm long; neotropical ... ... Pyrenula neosandwicensis Aptroot nom. nov.
Ascospores 115–180 µm long; pantropical ... ... Pyrenula duplicans (Nyl.) Aptroot
[Verrucaria duplicans Nyl., Anthracothecium duplicans (Nyl.) Müll. Arg., Anthracothecium exsertum (Krempelh.) Müll. Arg., Verrucaria exserta Krempelh., Verrucaria interponens Nyl., Polyblastia interponens (Nyl.) Müll. Arg., Anthracothecium interponens (Nyl.) Müll. Arg., Anthracothecium maculatum Nagarkar & Patw., Verrucaria assamiensis Stirt., Parmentaria assamiensis (Stirt.) Zahlbr., Anthracothecium assamiense (Stirt.) Ajay Singh, Anthracothecium pseudocyphellatum Ajay Singh, Anthracothecium pustuliferum Ajay Singh]
45(41) Hamathecium inspersed, ascospores 30–45 µm long; India ... ... Pyrenula sublaevigata (Patw. & Makhija) Upreti
(Anthracothecium sublaevigatum Patw. & Makhija)
Hamathecium not inspersed ... 46
46(45) Lumina relatively large and angular, with up to 6 between 2 primary septa;
pantropical ... Pyrenula leucostoma Ach.
[Anthracothecium leucostomum (Ach.) Malme, Sporodictyon feei var. leucostomum (Ach) Trevis.,? Pyrenula subcutanea Fée (nom. illeg.), Anthracothecium subcutaneum Müll. Arg., Pyrenula libricola Fée, Verrucaria libricola (Fée) Nyl., Anthracothecium libricolum (Fée) Müll. Arg., Bottaria libricola (Fée) Vain., Sporodictyon feei var. libricolum (Fée) Trevis., Sporodictyon feei Trevis., ? Anthracothecium javanicum (Hepp) Zahlbr., Verrucaria javanica Hepp, Verrucaria paramera Nyl., Bottaria paramera (Nyl.) Vain., Anthracothecium paramerum (Nyl.) Müll. Arg., Verrucaria analepta var. americana Ach., Anthracothecium americanum (Ach.) Müll. Arg., Verrucaria emergens Leight., Anthracothecium emergens (Leight.) Zahlbr., Bottaria paramera f. pallidoalba Vain., Anthracothecium paramerum f. pallidoalbum (Vain.) Zahlbr., Bottaria endococcinea Vain., Anthracothecium endococcineum (Vain.) Zahlbr., Bottaria submucosa Vain., Anthracothecium submucosum (Vain.) Zahlbr., Bottaria erythrinae Vain., Anthracothecium erythrinae (Vain.) Zahlbr., Pyrenula pachycheila Tuck., Anthracothecium pachycheilum (Tuck.) Zahlbr., Parmentaria rappii Zahlbr., Anthracothecium fraternale Zahlbr., Anthracothecium obscuratum Upreti & Ajay Singh]
Lumina mostly round, at least in the central part of the ascospore with more than 6 between 2 primary septa ... 47
47(46) Ascospores >50 µm long ... 48
Ascospores <50 µm long ... 49
48(47) Ascospores with rounded ends; pantropical ... ... Pyrenula pyrenuloides (Mont.) R.C. Harris
[Trypethelium pyrenuloides Mont., Bathelium pyrenuloides (Mont.) Trevis., Verrucaria pyrenuloides (Mont.) Nyl., Pyrenastrum pyrenuloides (Mont.) Nyl., Anthracothecium pyrenuloides (Mont.) Müll. Arg., Bottaria pyrenuloides (Mont.) Vain., ? Pyrenula cinerosa Ach., Anthracothecium cinerosum (Ach.) Müll. Arg., Verrucaria pyrenoica Ach., also as Verrucaria pyrinoica Ach., Sporodictyom pyrenoicum (Ach.) Trevis., also as Sporodictyom pyrinoicum (Ach.) Trevis., Parmentaria pyrenoica (Ach.) Müll. Arg, also as Parmentaria pyrinoica (Ach.) Müll. Arg, Verrucaria variolosa f. pyrenoica (Meyen & Flot.) Nyl., Anthracothecium goniostomum Müll. Arg., Anthracothecium amphitropum Müll. Arg., Verrucaria globifera ssp. depressa Eschw., Verrucaria depressa (Eschw.) Meyen & Flot., Spermatodium depressum (Eschw.) Trevis., Anthracothecium depressum (Eschw.) Müll. Arg., Anthracothecium sandwicense var. convexum Zahlbr., Anthracothecium cristatellum Nagarkar & Patw., Anthracothecium leightonii Patw. & Makhija]
Ascospores with pointed ends; pantropical ... ... Pyrenula papillifera (Nyl.) Aptroot comb. nov.
49(47) Ascospores 25–35 µm long; palaeotropical ... ... Pyrenula welwitschii (Upreti & Ajay Singh) Aptroot comb. nov.
Ascospores >35 µm long ... 50
50(49) Ascospores 11–15 µm wide; pantropical ... Pyrenula thelomorpha Tuck.
[also as Pyrenula thelemorpha Tuck., Bottaria thelomorpha (Tuck.) Vain., also as Bottaria thelemorpha (Tuck.) Vain., Anthracothecium thelomorphum (Tuck.) Zahlbr., also as Anthracothecium thelemorphum (Tuck.) Zahlbr., Verrucaria cellulosa C. Knight, Anthracothecium cellulosum (C. Knight) Müll. Arg.]
Ascospores 14–22 µm wide ... 51
51(50) Old ascospores filled with colourless oil; neotropical ... ... Pyrenula oleosa R.C. Harris
Old ascospores without oil, shriveling when old; neotropical ... ... Pyrenula dissimulans (Müll. Arg.) R.C. Harris
(Pleurothelium dissimulans Müll. Arg., Anthracothecium cascarillae Müll. Arg.)
Key B. Corticolous pyrenocarpous lichens with brown, transversely septate ascospores
Synopsis for quick access (only first alternative given except at the end)
1 Ascospores 2-septate ... 2
3 Hamathecium anastomosing, ascospores >90 µm long ... Architrypethelium, 4
5 Ascospores nearly only euseptate, lumina rectangular ... 6
15 Ascospores red-brown and lumina at least becoming rounded when older ... ... Lithothelium, 16
20 Ostioles pointing in various directions, mostly eccentric to lateral ... 21
39 Ascospores with many hyaline cilia at the ends ... 40
41 Ascospores at least partly or seemingly with more than 3 septa ... 42
53 Ascospores mostly >50 µm long ... 54
59 Lumina not in a straight line, somewhat to strongly zig-zag ... 60
61 Ascomata and/or thallus with yellow, orange or red anthraquines on the outside ... 62
70 Ascomata mostly aggregated, with fused walls but with separate ostioles ... 71
80 Thallus ecorticate, whitish ... 81
85 Old ascospores with orange oil ... 86
87 Terminal lumina all directly against the exospore wall ... 88
107 Hamathecium inspersed ... 108
119 Thallus UV+yellow ... 120
123 Ascospores mostly >25 µm long ... 124
135 Ascospores mostly 21–25 µm long ... 136
– Ascospores mostly <21 µm long ... 142
Main Key
1 Ascospores 2-septate (no Pyrenula species with 1-septate ascospores are known, corticolous pyrenocarpous lichens with 1-septate brown ascospores include species of Clypeopyrenis, Distopyrenis, Distothelia, Parapyrenis, Granulopyrenis, and Mycomicrothelia) ... 2
Ascospores at least 3-septate ... 3
2(1) Ascospore septation symmetrical; pantropical ... ... Pyrenula lineatostroma Aptroot
(Melanotheca indica Nyl., Pyrenula subindica Upreti, Melanotheca indica var. vaga Nyl.)
Ascospore septation strongly asymmetrical; Madagascar ... ... Lacrymospora parasitica Aptroot
3(1) Hamathecium anastomosing, ascospores >90 µm long ... ... Architrypethelium, 4
Hamathecium mostly unbranched, ascospores <90 µm long ... 5
4(3) Ostioles eccentric; neotropical ... Architrypethelium nitens (Fée) Aptroot
[Verrucaria nitens Fée, Pyrenula nitens (Fée) Fée, Pyrenula nitida var. americana Fée (nom. inval.), Pyrenastrum seminudum Mont., Architrypethelium pyrenuloides (Mont.) Aptroot, Parathelium ernstianum Müll. Arg., Pleurothelium ernstianum (Müll. Arg.) Müll. Arg., Parathelium superans Müll. Arg., Splanchnonema superans (Müll. Arg.) O. Erikss.]
Ostioles apical; neotropical ... Architrypethelium uberinum (Fée) Aptroot
[Porina uberina Fée, Porophora uberina (Fée) Spreng., Pyrenula uberina (Fée) Fée, Pertusaria uberina (Fée) A. Massal., Trypethelium uberinum (Fée) Nyl., Verrucaria uberina (Fée) Trevis., Stromatothelium uberinum (Fée) Trevis., Verrucaria megalospora Krempelh., Parathelium megalosporum (Krempelh.) Müll. Arg., Trypethelium uberinoides Nyl.]
5(3) Ascospores nearly only euseptate, lumina rectangular, at most a bit rounded in the corners, end cells often paler than middle cells (here would also key out the fungus Hysterium pulicare, which has a slit-like ostiole) ... 6
Ascospores clearly distoseptate, lumina of different shape than the outer wall ... ... 15
6(5) Ascospores only 3-septate, macroconidia 3-septate or unknown ... 7
Ascospores 3–7-septate, macroconidia either 1–7-septate or unknown ... 12
7(6) Ascoma wall dense ... 8
Ascoma wall cellular ... Eopyrenula, 10
8(7) Ascospores smooth, 14–17 µm long; neotropical ... ... Pyrenula tenuisepta R.C. Harris
Ascospores verrucose, 30–47 µm long ... Mycomicrothelia, 9
9(8) Ascomata at least partly fused with a more or less joint ostiole; Japan ... ... Mycomicrothelia collospora (Vain.) Aptroot
[Pyrenula collospora Vain., Melanotheca collospora (Vain.) Zahlbr.]
Ascomata all single; Australasian ... ... Mycomicrothelia queenslandica (Müll. Arg.) Sipmn & Aptroot
[Microthelia queenslandica Müll. Arg., Ornatopyrenis queenslandica (Müll. Arg.) Aptroot]
10(7) Ascospores 4–5 µm wide; New World temperate ... ... Eopyrenula parvispora R.C. Harris & Aptroot
Ascospores 5·5–7·5 µm wide ... 11
11(10) Ascospores mostly <15 µm long; Atlantic (W. Europe) ... ... Eopyrenula avellanae Coppins
Ascospores mostly >15 µm long; Atlantic (W. Europe) ... ... Eopyrenula grandicula Coppins
12(6) Ascospores 3(–5)-septate, macroconidia 1-septate; Old World temperate ... ... Eopyrenula leucoplaca (Wallr.) R.C. Harris
[Verrucaria leucoplaca Wallr., Pyrenula leucoplaca (Wallr.) Körb., also as Pyrenula leucophaea (Wallr.) Körb., Pyrenula alba var. leucoplaca (Wallr.) Schaer., Spermatodium leucoplacum (Wallr.) Trevis., Leptosphaeria leucoplaca (Wallr.) Vain., ? Pyrenula alni A. Massal., Pyrenula alba A. Massal., Pyrenula quercus A. Massal., Verrucaria quercus (A. Massal.) Garov., Pyrenula schaereri A. Massal., Pyrenula leucoplaca var. umbrosa Körb., Pyrenula farrea var. umbrosa (Körb.) Zahlbr., Pyrenula leucoplaca f. umbrosa (Körb.) Migula, Pyrenula leucoplaca var. chrysoleuca Körb., Pyrenula leucoplaca f. chrysoleuca (Körb.) Migula, Sagedia chiomela Norm., Porina chiomela (Norm.) Zahlbr., Pyrenula glabrata f. cinerea Haszl., Pyrenula laevigata f. cinerea (Haszl.) Zahlbr., ? Blastodesmia albonigra Zahlbr.]
Ascospores (3–)5–7-septate, macroconidia 3–7-septate or unknown ... 13
13(12) Ascospores mostly 37–47 µm long, constricted at the septa; Europe ... ... Blastodesmia nitida A. Massal.
[Polyblastia nitida (A. Massal.) Tevis., Verrucaria massalongoi Garov., Verrucaria circumfusa Nyl., Pyrenula circumfusa (Nyl.) Trevis., Sagedia circumfusa (Nyl.) Haszl.]
Ascospores mostly <35 µm long ... Eopyrenula, 14
14(13) Ascospores >6 µm wide, macroconidia (5–)7-septate; Atlantic (W. Europe) ... ... Eopyrenula septemseptata Coppins
Ascospores <6 µm wide, macroconidia 3(–4)-septate; New world temperate ... ... Eopyrenula intermedia Coppins
[Pyrenula leucoplaca var. pluriloculata Fink, Pyrenula farrea var. pluriloculata (Fink) Zahlbr.]
15(5) Ascospores red-brown and lumina at least becoming rounded when older (species on rock not keyed out here) ... Lithothelium, 16
Ascospores grey to brown, rarely red-brown and then lumina angular ... ... Pyrenula, 20
16(15) Thallus UV+yellow; neotropical (Costa Rica) ... ... Lithothelium fluorescens Aptroot & Sipman
Thallus UV− ... 17
17(16) Ascospores 3-septate ... 18
Ascospores 7-septate ... 19
18(17) Ascospores 15–20 µm long; eastern palaeotropical ... ... Lithothelium decumbens (Müll. Arg.) Aptroot
[Parathelium decumbens Müll.Arg., Pyrenula decumbens (Müll.Arg.) Upreti]
Ascospores 25–40 µm long; northern temperate ... ... Lithothelium phaeosporum (R.C. Harris) Aptroot
(Plagiocarpa phaeospora R.C. Harris)
19(17) Ascospores 30–40 µm long; northern temperate ... ... Lithothelium septemseptatum (R.C. Harris) Aptroot
(Plagiocarpa septemseptata R.C. Harris)
Ascospores 55–80 µm long; New World temperate ... ... Lithothelium macrosporum (R.C. Harris) Aptroot
(Plagiocarpa macrospora R.C. Harris)
20(15) Ostioles pointing in various directions, mostly eccentric to lateral ... 21
Ostioles apical or, when eccentric, all pointing in the same direction ... 39
21(20) Ascospores 5–15-septate ... 22
Ascospores 3-septate ... 24
22(21) Ascospores 5-septate, 42–55 µm long; neotropical (Colombia) ... ... Pyrenula pleiomera (Nyl.) Zahlbr.
(Verrucaria pleiomera Nyl.)
Ascospores 9–15-septate ... 23
23(22) Ascospores 9–13-septate, 65–90 × 17–22 µm ascomata single ... ... Pyrenula fusispora (Malme) Aptroot comb. nov.
Ascospores 11–15-septate, 50–70 × 4·5–6 µm, ascomata with 5–15 ostioles fused. ... Pyrenula tokyensis (Müll. Arg.) H. Harada
(Pyrenastrum tokyense Müll. Arg., also as Pyrenastrum tokyoense Müll. Arg.)
24(21) Thallus UV+yellow ... 25
Thallus UV− ... 26
25(24) Ascospores 21–23 µm long; Hawaii ... Pyrenula hawaiiensis Aptroot ined.
Ascospores 32–45 µm long; neotropical (Brazil) ... ... Pyrenula crassiuscula (Malme) Aptroot
(? Pyrenula copalchina Fée, Parathelium crassiusculum Malme, Parathelium crassiusculum f. chlorophorae Malme)
26(24) Terminal lumina directly against the exospore wall ... 27
Terminal lumina separated from the exospore wall by endospore thickening ... ... 31
27(26) Ascomata at least partly fused with joint ostioles ... ... Pyrenula subgregantula Müll. Arg.
[Pyrenastrum personatum Malme, Pyrenula personata (Malme) R.C. Harris, Pyrenastrum fulvum Malme, Pyrenula fulvella R.C. Harris]
Ascomata single ... 28
28(27) Ascospores 13–25 µm long ... 29
Ascospores 25–65 µm long ... 30
29(28) Ascospores 16–25 µm long; pantropical ... Pyrenula circumfiniens Vain.
[Parathelium subferrugineum Malme, Parathelium subferrugineum f. expallescens Malme, Pyrenula subferruginea (Malme) R.C. Harris]
Note: the completely lateral position of the ostiole in the type of Pyrenula circumfiniens was not noticed before.
Ascospores 13–16 µm long; neotropical ... Pyrenula elliptica Müll. Arg.
(Lectotype (designated here): Cuba, Wright, Verr. Cub. 218a p.p. (G); the specimen in G of 218b p.p. under the name Pyrenula elliptica is Pyrenula mamillana)
30(28) Ascospores 45–65 µm long; neotropical ... Pyrenula erumpens R.C. Harris
(Parathelium emergens Nyl. ex Müll. Arg.)
Ascospores 25–35 µm long; pantropical ... ... Pyrenula cuyabensis (Malme) R.C. Harris
(Parathelium cuyabense Malme)
31(26) Ostiole outside with a red ring; New world temperate ... ... Pyrenula wetmorei R.C. Harris
Ostiole without red colour ... 32
32(31) Ascomata mostly aggregated with a shared ostiole ... 33
Ascomata nearly all single ... 35
33(32) Ascospores 12–24 µm long; pantropical ... ... Pyrenula septicollaris (Eschw.) R.C. Harris
[Pyrenastrum septicollare Eschw., Parmentaria septicollare (Eschw.) Trevis., Astrothelium septicollare (Eschw.) Leight., Pyrenula irregularis Fée, Pyrenastrum irregulare (Fée) Müll. Arg., Pyrenastrum fuscum Mont., Porodothion acharii Mont., Astrothelium pyrenastraeum Nyl., Pyrenastrum pyrenastraeum (Nyl.) Zahlbr., Pyrenastrum depressum Müll. Arg., Pyrenastrum bicolor Vain., Pyrenula polillensis Vain., Melanotheca polillensis (Vain.) Zahlbr., Pyrenastrum microsporum Malme, Pyrenastrum pruinosum C.W. Dodge, Pyrenastrum erumpens C.W. Dodge, Pyrenastrum parathelioides C.W. Dodge, Pyrenula laureriformis Aptroot]
Notes. The new synonym Pyrenula irregularis (holotype: G-FEE seen) was described in the same year as Verrucaria septicollare but seems to be the younger name. The completely lateral position of the ostiole and the joint ostioles in the type of Pyrenula polillensis were not noticed before. Study of additional specimens showed that Pyrenula laureriformis falls within the variation of Pyrenula septicollare.
Ascospores >25 µm long ... 34
34(33) Ascospores 30–45 µm long; neotropical ... ... Pyrenula cryptothelia (Müll. Arg.) Aptroot & Etayo
[Pyrenastrum cryptothelium Müll.Arg., Astrothelium cryptothelium (Müll. Arg.) Nyl., Pyrenastrum depauperatum Malme]
Ascospores 25–35 µm long; neotropical ... ... Pyrenula cubana (Müll. Arg.) R.C. Harris
(Pyrenastrum cubanum Müll.Arg., Pyrenastrum cubanum var.obtectum Malme, Pyrenastrum cubanum var. intermedium Malme)
35(32) Ascospores 13–17 µm long, hamathecium inspersed; neotropical (Florida). ... ... Pyrenula wheeleri R.C. Harris
Ascospores >17 µm long, hamathecium not inspersed ... 36
36(35) Ascospores 28–45 µm long (compare also Pyrenula minarum, with partly eccentric ostioles) ... 37
Ascospores 17–28 µm long ... 38
37(36) Ascospores 35–45 µm long; pantropical (America and Africa) ... ... Pyrenula adacta Fée
[Verrucaria punctella var. adacta (Fée) Nyl., Pyrenula punctella var. adacta (Fée) Müll. Arg., Parathelium indutum Nyl., Pleurothelium indutum (Nyl.) Müll. Arg., Parathelium martinicanum Vain., Pyrenula martinicana (Vain.) R.C. Harris (nom. illeg.), Pyrenula caraibica Aptroot, ? Pyrenula pulchella Müll. Arg., Pyrenula marginatula Müll. Arg., Pyrenula acaciae Vain., Parathelium dilutum Malme]
Ascospores 28–32 µm long; probably pantropical (only known from Brazil and Papua New Guinea) ... Pyrenula gahavisukana Aptroot
(Parathelium dilutum var. catervaria Malme)
38(36) Ascospores with at least one pointed end; temperate Northern Hemisphere, extending to the tropics ... Pyrenula acutispora Kalb & Hafellner
[Pyrenula kakouettae Sérus. & Diederich, Pyrenula alnicola R.C. Harris (nom. inval.)]
Ascospores with rounded ends; New World ... ... Pyrenula microtheca R.C. Harris
(Parathelium microcarpum Riddle)
39(20) Ascospores with many hyaline cilia at the ends (not to be confused with germ tubes which are one per lumen) ... 40
Ascospores without cilia ... 41
40(39) Ascospores 3-septate; Australasian (Papua New Guinea) ... ... Pyrenula ciliata Aptroot
Ascospores 5–7-septate; neotropical (Panama) ... Pyrenula hirsuta Etayo
41(39) Ascospores at least partly with more than 3 septa, or with 3 septa and long tails at one or both ends, thus seemingly more than 3-septate ... 42
Ascospores all 3-septate ... 52
42(41) Ascospores at least seemingly 4–7-septate ... 43
Ascospores 7–17-septate, more than 4 times as long as wide ... 48
43(42) Ascospores with tails at both ends; Australia ... ... Pyrenula bicuspidata Müll. Arg.
(Melanotheca oxyspora Müll. Arg.)
Ascospore without tails, at most pointed ... 44
44(43) Ascospores more than 4 times as long as wide; neotropical ... ... Pyrenula melanophthalma (Mont.) Trevis.
[Verrucaria melanophthalma Mont., Melanotheca melanophthalma (Mont.) Müll. Arg., Trypethelium melanophthalmum (Mont.) Nyl., Stromatothelium melanophthalma (Mont.) Trevis., Verrucaria infida Nyl., Pyrenula infida (Nyl.) Müll. Arg.]
Ascospores less than 4 times as long as wide ... 45
45(44) Ascospores <30 µm long ... 46
Ascospores >29 µm long ... 47
46(45) Old ascospores with orange oil, thallus often with pseudocyphellae; pantropical ... ... Pyrenula sexlocularis (Nyl.) Müll. Arg.
[Verrucaria sexlocularis Nyl., ? Verrucaria cyrtospora Stirt., Pyrenula cyrtospora (Stirt.) Nyl., Verrucaria concatervans Nyl., Melanotheca concatervans (Nyl.) Zahlbr., Pyrenula concatervans (Nyl.) R.C. Harris, Anthracothecium seminudum Müll. Arg., Anthracothecium hexamerum Müll. Arg., Pyrenula atroalbella Vain., Pyrenula flavofulvescens Vain., Melanotheca cinerata Zahlbr., Pyrenula sexlocularis var. xanthoplaca Zahlbr.]
Note: the specimens until recently called Pyrenula concatervans with 3-septate ascospores are now kept separate as Pyrenula bahiana; therefore a name-change of the 5-septate specimens from Pyrenula concatervans to Pyrenula sexlocularis is welcome.
Ascospores without orange oil, thallus without pseudocyphellae; South Africa ... ... Pyrenula wilmsiana Mül. Arg.
47(45) Ascospores 29–35 µm long, hamathecium not inspersed; Australasian (Papua New Guinea) ... Pyrenula sexluminata Aptroot nom. nov.
Ascospores 30–55 µm long, hamathecium inspersed; pantropical ... ... Pyrenula caracasana Müll. Arg.
(Pyrenula mangiferae Vain., Pyrenula megapotamica Malme)
48(42) Ascospores with a long tail at one end; Pacific (Mariana Islands) (here the temperate fungal genus Rebentischia keys out) ... ... Pyrenula flagellata H. Harada
Ascospores without tails (here the common fungal genus Navicella keys out, characterized by elongate ostioles) ... 49
49(48) Ascospores 7–17-septate ... 50
Ascospores (5–)7-septate ... 51
50(49) Ascospores 12–17-septate; eastern palaeotropical (Ryukyu Islands, Japan) ... ... Pyrenula cylindrica Kashiw.
Ascospores 7–11-septate; eastern palaeotropical (India) ... ... Pyrenula subcylindrica Jagadeesh Ram & Upreti
Note: the enigmatic Melanotheca pusilla (Jatta) C.W. Dodge from Ethiopia would key out here. It is reported to have 9-septate ascospores.
51(49) Ascospores 30–38 µm long; pantropical ... Pyrenula montagnei Müll. Arg.
Ascospores 53–70 µm long; Australasian (New Zealand), also neotropical? ... ... Pyrenula moniliformis (C. Knight) Müll. Arg.
(Verrucaria moniliformis C. Knight)
52(41) Ascospores more than 4 times as long as wide; neotropical (Brazil) ... ... Pyrenula fusoluminata Aptroot
Ascospores less than 4 times as long as wide ... 53
53(52) Ascospores mostly >50 µm long ... 54
Ascospores mostly <50 µm long ... 59
54(53) Lumina not in a straight line, somewhat zig-zag; neotropical (Brazil) ... ... Pyrenula hoehneliana Zahlbr.
Lumina in a straight line or ascospores bent, not zig-zag ... 55
55(54) Terminal lumina directly against the exospore wall ... 56
Terminal lumina separated from the exospore wall by endospore thickening ... ... 58
56(55) Thallus UV+ yellow; neotropical ... Pyrenula praelucida (Mont.) Trevis.
(Verrucaria praelucida Mont.)
Thallus UV− ... 57
57(56) Hamathecium not inspersed; pantropical ... Pyrenula rockii Zahlbr.
Hamathecium inspersed; neotropical ... Pyrenula subpraelucida Müll. Arg.
[?(inspersed?) Pyrenula quadruplans Vain.]
58(55) Ascospores 75–90 µm long, thallus with crystalline papillae; neotropical (Costa Rica) ... Pyrenula montocensis Lücking
Ascospores (45–)50–60 µm long., thallus without papillae but with pseudocyphellae; pantropical ... Pyrenula immissa (Stirt.) Zahlbr.
(Verrucaria immissa Stirt., Pyrenula cuprescens A. Zahlbr., Pyrenula laii Aptroot)
59(53) Lumina not in a straight line, somewhat to strongly zig-zag ... 60
Lumina in a straight line, not zig-zag ... 61
60(59) Thallus UV+ yellow; pantropical ... Pyrenula aggregans Vain.
Thallus UV−; pantropical ... Pyrenula papilligera (Leight.) Müll. Arg.
(Verrucaria papilligera Leight., Anthracothecium fulvum Müll. Arg., Pyrenula neofulva Ajay Singh)
61(59) Ascomata and/or thallus with yellow, orange or red anthraquinones on the outside ... 62
Ascomata and thallus without external pigments ... 70
62(61) Terminal lumina directly against the exospore wall ... 63
Terminal lumina separated from the exospore wall by endospore thickening ... ... 66
63(62) Ascomata at least partly yellow, hamathecium not inspersed; neotropical (Panama, Costa Rica, Puerto Rico, Hawaii) ... ... Pyrenula luteopruinosa Etayo & Aptroot
Ascomata or thallus orange or red, hamathecium inspersed ... 64
64(63) Ascomata red, thallus UV−; Australasian (Lord Howe Island) ... ... Pyrenula howeana Aptroot
Thallus or ascomata orange, thallus UV+yellow (when thallus patchily orange brown and UV−, see Pyrenula occidentalis) ... 65
65(64) Ascomata orange, ascospores 17–20 µm long; eastern palaeotropical (Korea) ... ... Pyrenula sipmanii Aptroot & K. H. Moon
Thallus orange, ascospores 19–22 µm long; eastern palaeotropical (Thailand) ... ... Pyrenula aurantiopileata Aptroot
66(62) Thallus orange; neotropical ... Pyrenula cerina Eschw.
[Verrucaria cerina Eschw., Spermatodium cerinum (Eschw.) Trevis., Pyrenula aurantiaca Fée, Verrucaria aurantiaca (Fée) Nyl., Pyrenula cerina var. expallens Zahlbr.]
Thallus and/or ascomata at least partly red ... 67
67(66) Ascospores 15–19 µm long, hamathecium not inspersed ... 68
Ascospores >19 µm long, hamathecium inspersed ... 69
68(67) Ascospores 15–17 µm long, ascomata mostly aggregated, with fused walls but with separate ostioles; neotropical (Costa Rica) ... ... Pyrenula rubroanomala Aptroot & Lücking
Ascospores 17–19 µm long, ascomata simple, red coloration only around the ostiole; neotropical ... Pyrenula rubrostoma R.C. Harris
69(67) Ascospores 19–22 µm long; eastern palaeotropical (Java) ... ... Pyrenula rubrojavanica Aptroot ined.
Ascospores 27–35 µm long; pantropical ... Pyrenula cruenta (Mont.) Vain.
[Trypethelium cruentum Mont., Stromatothelium cruentum (Mont.) Trevis., Melanotheca cruenta (Mont.) Müll. Arg., ? Verrucaria sinapisperma Fée, Sphaeromphale sinapisperma (Fée) Trevis., Anthracothecium sinapispermum (Fée) Müll. Arg., Trypethelium rubrum C. Knight, Melanotheca rubra (C. Knight) C. Knight, Trypethelium cinnabrinum C. Knight, Melanotheca cinnabarina (C. Knight) C. Knight, Melanotheca rubescens C. Knight, Verrucaria circumrubens Nyl., Pyrenulia circumrubens (Nyl.) B de Lesd., Pyrenulia circumrubens var. erythrinosa B de Lesd., Verrucaria circumrubens var. rubrotecta Stirt., Pyrenulia circumrubens var. rubrotecta (Stirt.) Shirley, Trypethelium cruentum var. subdecolor Nyl., Melanotheca cruenta f. subdecolor (Nyl.) Zahlbr., Pyrenula subdecolor (Nyl.) R.C. Harris, Trypethelium subincruentum Nyl., Melanotheca subincruenta (Nyl.) Zahlbr., ? Trypethelium coccinatum Stizenb., Trypethelium purpurascens Stizenb., Melanotheca purpurascens (Stizenb.) Müll. Arg., Trypethelium connivens Stirt. (nom. illeg.), Melanotheca connivens Zahlbr., Trypethelium cruentulum Nyl., Melanotheca cruentula (Nyl.) Zahlbr., Melanotheca ornata Müll. Arg., Trypethelium ornatum (Müll. Arg.) Hellb. (nom. illeg.), Trypethelium oblitescens Stirt., Pyrenastrum oblitescens (Stirt.) Makhija & Patw., Pyrenula gravenreuthii Stein, Pyrenula rubromaculata Vain., Melanotheca rubromaculata (Vain.) Zahlbr.]
70(61) Ascomata mostly aggregated, with fused walls but with separate ostioles ... 71
Ascomata mostly simple, only aggregated as by chance when crowded (if difficult to decide, start with the first alternative) ... 80
71(70) Old ascospores with red oil; neotropical (Puerto Rico) ... ... Pyrenula concastroma R.C. Harris
Old ascospores without red oil ... 72
72(71) Terminal lumina directly against the exospore wall; Australasian (Papua New Guinea) ... Pyrenula pyrenastrospora Aptroot
Terminal lumina separated from the exospore wall by endospore thickening ... ... 73
73(72) Hamathecium inspersed, ostioles sometimes partly eccentric, ascospores 25–40 µm long; pantropical ... Pyrenula minarum Vain.
(Pyrenula minarum var. colorans Malme)
Hamathecium not inspersed ... 74
74(73) Ascospores mostly >21 µm long ... 75
Ascospores mostly <20 µm long ... 78
75(74) Ascospores mostly >25 µm long ... 76
Ascospores mostly <25 µm long ... 77
76(75) Thallus with pseudocyphellae; neotropical (Cuba) ... ... Pyrenula wrightii (Müll. Arg.) R.C. Harris
(Melanotheca wrightii Müll. Arg.)
Thallus without pseudocyphellae; eastern palaeotropical (Sri Lanka) ... ... Pyrenula zeylanica Upreti & Ajay Singh
77(75) End lumina elongated; neotropical (Costa Rica) ... ... Pyrenula subsoluta (Müll. Arg.) Aptroot
(Melanotheca subsoluta Müll. Arg.)
All lumina more or less rounded; eastern palaeotropical ... ... Pyrenula leucotrypa (Nyl.) Upreti
[Trypethelium leucotrypum Nyl., Melanotheca leucotrypa (Nyl.) Müll. Arg., ? Trypethelium erumpens Stirt. (nom. illeg.), Melanotheca stirtoniana Müll. Arg., Trypethelium stirtonianum (Müll. Arg.) Hellb., Melanotheca negrosensis Herre]
78(74) Ascospores with dark bands between the lumina; neotropical (Florida) ... ... Pyrenula atrolaminata R.C. Harris
Ascospores without dark bands ... 79
79(78) Ascospores mostly >15 µm long; pantropical ... ... Pyrenula anomala (Ach.) Vain.
[Trypethelium anomalum Ach., Melanotheca anomala (Ach.) A. Massal., Melanotheca achariana Fée, Pyrenula achariana (Fée) Vain., Mycoporum acharii G. Mey., Porodithion acharii (G. Mey.) Mont., Trypethelium nudum Fée, Coenoicia nuda (Fée) Trevis, Celothelium nudum (Fée) Trevis., Trypethelium favulosum Ach., Trypethelium scoria Fée (nom. illeg.), Pseudopyrenula scoria (Fée) Vain. (nom. illeg.),? Trypethelium inconspicuum C.F.W. Meissn., Trypethelium fuscum Krempelh., Pyrenula fusca (Krempelh.) Vain., Melanotheca fusca (Krempelh.) Müll. Arg., Melanotheca aggregata Müll. Arg., Melanotheca arthonioides var. grisea Müll. Arg., Melanotheca eschweileri var. grisea (Müll. Arg.) Zahlbr., Melanotheca foveolata Müll. Arg., Trypethelium nigritulum Nyl., Melanotheca achariana var. angolensis Vain., Melanotheca angolensis (Vain.) C. W. Dodge, Melanotheca subdissidens (Vain.) Zahlbr., Pyrenula obscurascens Vain., Trypethelium anomalum var. obscurascens (Vain.) Zahlbr., Melanotheca achariana var. obscurascens (Vain.) Zahlbr., Melanotheca obscurascens (Vain.) C. W. Dodge, Pyenula anomaloides Vain., Melanotheca irregularis Zahlbr., Melanotheca arthonioides var. machaerii Malme, Melanotheca arthonioides var. lueheae Malme, Melanotheca cameroonensis C. W. Dodge, Melanotheca porosa C. W. Dodge, Trypethelium microsporum Makhija & Patw.]
Ascospores mostly <15 µm long; pantropical ... ... Pyrenula arthoniotheca Upreti
[Porothelium arthonioides Eschw., Verrucaria arthonioides (Eschw.) Eschw., Porodothion arthonioides (Eschw.) Trevis., Melanotheca arthonioides (Eschw.) Müll. Arg., Pyrenula arthonioides (Eschw.) Vain. (nom. illeg.), Syngenosorus eschweileri Trevis., Melanotheca eschweileri (Trevis.) Zahlbr., ? Pyrenula subdissidens Vain., Melanotheca nigeriensis C.W. Dodge]
Note: it remains uncertain whether this species is not mostly immature material of the previous species.
80(70) Thallus ecorticate, whitish ... 81
Thallus corticate (may be difficult to observe in poor specimens; when in doubt try first alternative) ... 85
81(80) Thallus UV+yellow; pantropical ... Pyrenula cocoes Müll. Arg.
(Pyrenula rugulosa Müll. Arg., Pyrenula insularum H. Magn.)
Thallus UV− ... 82
82(81) Hamathecium inspersed; eastern paleotropical (Philippines) ... ... Pyrenula albothallina Vain.
Hamathecium not inspersed ... 83
83(82) Thallus white to grey, dull, ascomata <0·7 mm diam; pantropical ... ... Pyrenula microcarpa Müll. Arg.
(Pyrenula melaleuca Müll. Arg., Pyrenula albella Müll. Arg., Pyrenula microcarpoides Müll. Arg., Pyrenula conspurcata Müll. Arg., Pyrenula alboostiolata Vain., ? Pyrenula laevigata var. meizospora Vain., Pyrenula cinerea Zahlbr., Pyrenula texana Tuck. ex R.C. Harris)
Thallus cream to silvery or unapparent, smooth; temperate ... 84
84(83) Ascospores <18 µm long, hamathecium KOH−, pycnidia present; known from temperate regions in the Northern Hemisphere and reported from Chile, so probably cosmopolitan in temperate regions ... Pyrenula coryli A. Massal.
[Verrucaria coryli (A. Massal.) Nyl., Arthopyrenia coryli (A. Massal) Müll. Arg., Arthopyrenia glabrata var.coryli (A. Massal.) H. Olivier, Microthelia glabrata var. coryli (A. Massal.) Boistel, Mycopyrenula coryli (A. Massal.) Vain.]
Ascopores >18 µm long, hamathecium sometimes KOH+blue (fleeting), pycnidia absent; temperate Northern Hemisphere ... ... Pyrenula laevigata (Pers.) Arnold
[Verrucaria laevigata Pers., Pyrenula alba var. laevigata (Pers.) Trevis., Verrucaria glabrata Ach., Microthelia glabrata (Ach.) Boistel, Arthopyrenia glabrata (Ach.) H. Olivier, Pyrenula glabrata (Ach.) A. Massal., Porina glabrata (Ach.) Sandst., Verrucaria nitida f. albolutea Grognot, Verrucaria gemmata var. minor Nyl., Pyrenula glabrata f. major Krempelh., Pyrenula laevigata f. major (Krempelh.) Zahlbr., Pyrenula glabrata f. microcarpa Hepp, Pyrenula laevigata f. microcarpa (Hepp) Arnold, Pyrenula alba var. microcarpa (Hepp) Trevis.]
85(80) Old ascospores with orange oil ... 86
Old ascospores without orange oil ... 87
86(85) Ascospores 25–35 µm long; pantropical ... Pyrenula bahiana Malme
(Pyrenula crystalligera H. Magn.)
Note: this species has until recently been called Pyrenula concatervans, including material now called Pyrenula sexlocularis. Although a few specimens of Pyrenula sexlocularis are known that contain both 3- and 5-septate ascospores, the majority of the specimens in this group have either only 3- (in Pyrenula bahiana) or 5-septate ascospores (in Pyrenula sexlocularis).
Ascospores (30–)35–51 µm long; eastern palaeotropical (India, Thailand) ... ... Pyrenula thailandica Aptroot ined.
87(86) Terminal lumina all directly against the exospore wall (if anthraquinones present on thallus and/or ascomata, compare 55) ... 88
Terminal lumina mostly (at least in mature ascospores) separated from the exospore wall by endospore thickening ... 107
88(87) Thallus UV+ yellow ... 89
Thallus UV− ... 90
89(88) Hamathecium inspersed; northern temperate ... ... Pyrenula pseudobufonia (Rehm) R.C. Harris
(Clypeosphaeria pseudobufonia Rehm, Pyrenula neglecta R.C. Harris, Pyrenula shirabeicola Kurok. & S. Nakan.)
Hamathecium not inspersed; pantropical ... Pyrenula cocoes Müll. Arg.
(Pyrenula rugulosa Müll. Arg., Pyrenula insularum H. Magn.)
90(88) Hamathecium inspersed (in some cases only in the upper part) ... 91
Hamathecium not inspersed ... 101
91(90) End lumina elongated ... 92
All lumina more or less rounded to angular ... 94
92(91) Ascospores >25 µm long ... 93
Ascospores 20–25 µm long; pantropical (compare also Pyrenula acutalis which is inspersed only in the upper part) ... Pyrenula maravalensis Vain.
(Pyrenula subacutalis Upreti)
93(92) Ascospores 26–30 µm long; Australasian (Papua New Guinea) ... ... Pyrenula rinodinospora Aptroot ined.
Ascospores 30–50 µm long; eastern palaeotropical (India) ... ... Pyrenula kurzii Ajay Singh & Upreti
94(91) Ascospores all <16 µm long; pantropical ... Pyrenula cayennensis Müll. Arg.
(Pyrenula hunana Zahlbr.)
Ascospores partly >16 µm long ... 95
95(94) Ascospores >40 µm long ... 96
Ascospores <40 µm long ... 97
96(95) Ascospores 42–50 µm long, mostly straight; eastern palaeotropical (India) ... ... Pyrenula subcamptospora Upreti
Note: the description describes the ascospores as curved, but they were mostly straight in the isotype.
Ascospores 45–52 µm long, mostly curved; neotropical ... ... Pyrenula cryptostoma (Nyl.) Müll. Arg.
(Verrucaria cryptostoma Nyl., Pyrenula camptospora Malme)
97(95) Ascospores partly >30 µm long ... 98
Ascospores all <30 µm long ... 99
98(97) Ascospores with black granular material around the ends; Australasian (Papua New Guinea) ... Pyrenula grossa Aptroot
Ascospores without black granular material; palaeotropical (Seychelles) ... ... Pyrenula fulva (Krempelh.) Müll. Arg
(Pyrenula marginata var. fulva Krempelh, Pyrenula supracongruens Aptroot & Schumm)
99(97) Thallus with patches of orange-brown coloration which are KOH+ purple. Known from temperate regions on the Northern Hemisphere and South Africa, so probably cosmopolitan ... ... Pyrenula occidentalis (R.C. Harris) R.C. Harris
[Pyrenula neglecta ssp. occidentalis R.C. Harris, Pyrenula glabrata f. incusa Flot., Pyrenula laevigata f incusa (Flot.) Zahlbr., Pyrenula harrisii Hafellner & Kalb]
Thallus without coloured, KOH+ purple patches ... 100
100(99) Hamathecium inspersed only in the upper part; neotropical ... ... Pyrenula acutalis R.C. Harris
Hamathecium totally inspersed; pantropical ... ... Pyrenula fetivica (Krempelh.) Müll. Arg.
[Verrucaria fetivica Krempelh., Pyrenula subcongruens Müll. Arg., Verrucaria fibrata Stirt., Pyrenula fibrata (Stirt.) Zahlbr., Pyrenula albida Müll. Arg., Verrucaria obtusior Nyl., Pyrenula obtusior (Nyl.) Zahlbr., Verrucaria glabriuscula Nyl., Pyrenula glabriuscula (Nyl.) Vain., ? Pyrenula truncata Müll. Arg., Pyrenula lamprocarpa Müll. Arg., Pyrenula rhombospora Müll. Arg., Pyrenula approximata Vain., Pyrenula feracissima Vain., Pyrenula mastospora Vain., ? Pyrenula samarana Vain., Pyrenula transparens Zahlbr., Pyrenula sandwicensis Zahlbr., Pyrenula sublateritia Zahlbr., Pyrenula quercuum Zahlbr., Pyrenula dussii Malme, Pyrenula subcremea Malme, Pyrenula commixta Malme, Pyrenula japonica Kurok., Pyrenula citriformis R.C. Harris]
Note: specimen H-NYL 1403 is here selected as the lectotype of Verrucaria obscurior, although specimen 1402 is identical.
101(90) End lumina elongated (all three species rare and possibly not clearly separated) ... 102
All lumina more or less rounded to angular ... 104
102(101) Ascospores 20–29 µm long, ascomata <0·7 mm diam ... 103
Ascospores 27–40 µm long, ascomata >0·7 mm diam; pantropical ... ... Pyrenula vernicosa (Krempelh.) Müll. Arg.
(Verrucaria vernicosa Krempelh., ? Pyrenula tricolor Müll. Arg.)
103(102) Ascospores 20–29 µm long, ascomata 0·5–0·7 mm diam; eastern palaeotropical ... ... Pyrenula finitima Müll. Arg.
(Pyrenula oxyspora Müll. Arg., Pyrenula oxysporiza Zahlbr.)
Ascospores 20–29 µm long, ascomata <0·5 mm diam; eastern palaeotropical ... ... Pyrenula approximans (Krempelh.) Müll. Arg.
[Verrucaria approximans Krempelh., Verrucaria nodulata Stirt., Pyrenula nodulata (Stirt.) Zahlbr.]
104(101) Ascomata > 2 mm diam; eastern palaeotropical (Philippines) ... ... Pyrenula irosina Vain.
Ascomata < 2 mm diam. ... 105
105(104) Ascospores 11–13 µm long; palaeotropical (Seychelles) ... ... Pyrenula infracongruens Aptroot & Schumm
Ascospores >13 µm long ... 106
106(105) Ascospores at least partly >16 µm long; pantropical ... ... Pyrenula nitidula (Bres.) R.C. Harris
[Melanomma nitidulum Bres., ? Verrucaria dealbata C. Knight, Pyrenula dealbata (C. Knight) Müll. Arg., Verrucaria glabrata var. cinereoalba C. Knight, Pyrenula marginata f. diminuens Nyl., Verrucaria marginata f. diminuens (Nyl.) Nyl., Pyrenula mamillana f. diminuens (Nyl.) Zahlbr., Pyrenula paraensis Müll. Arg., Verrucaria inflata Stirt., Pyrenula cinereoglauca Zahlbr., Bottaria dimorpha Vain., Anthracothecium dimorphum (Vain.) Zahlbr., Pyrenula laevigata var. microspora Vain., Pyrenula subsimplex Vain., Pyrenula griseola Vain., Melanotheca griseola (Vain.) Zahlbr., Pyrenula platysporella Zahlbr., Pyrenula plittii R.C. Harris, Pyrenula andamanica Ajay Singh & Upreti]
Note: the bands of dark granular material mentioned as a distinguishing character of Pyrenula plittii are in this ascospore type only a development state (postmature ascospores), visible also in some other species, notably often in Pyrenula finitima.
Ascospores 14–16 µm long; eastern palaeotropical ... ... Pyrenula mastophoriza (Nyl.) Zahlbr.
(Verrucaria mastophoriza Nyl., Anthracothecium peltophorum Müll. Arg., Pyrenula neopeltophora Ajay Singh, Pyrenula glabrescens Vain.)
Note: the ascospore size shows much variation in this and the previous species, also within one section. They might be one species.
107(87) Hamathecium inspersed ... 108
Hamathecium not inspersed ... 119
108(107) Central lumina strongly elongated; northern temperate ... ... Pyrenula subelliptica (Tuck.) R.C. Harris
[Clypeosphaeria imperfecta Ellis & Everh., Pyrenula imperfecta (Ellis & Everh.) R.C. Harris, Anthracothecium pauciloculare Herre]
Central lumina not strongly elongated ... 109
109(108) Ascospores constricted at the central septum; neotropical (Costa Rica) ... ... Pyrenula minae Aptroot & Lücking
Ascospores not constricted, at most becoming constricted when post mature ... ... 110
110(109) Ascomata mostly <0·7 mm diam ... 111
Ascomata mostly >0·7 mm diam ... 114
111(110) Ascospores 27–50 µm long; eastern palaeotropical (India) ... ... Pyrenula oculata Ajay Signh & Upreti
Ascospores <25 µm long ... 112
112(111) Ascospores 13–18 µm long; neotropical. ... Pyrenula laetior Müll. Arg.
(Pyrenula gregantula Müll. Arg.)
Ascospores 18–25 µm long ... 113
113(112) Ascospores 18–25 µm long; thallus and/or acomata with patches of rusty anthraquinone; known from temperate regions on the Northern Hemisphere and South Africa, so probably cosmopolitan ... ... Pyrenula occidentalis (R.C. Harris) R.C. Harris
[Pyrenula neglecta ssp. occidentalis R.C. Harris, Pyrenula glabrata f. incusa Flot., Pyrenula laevigata f. incusa (Flot.) Zahlbr., Pyrenula harrisii Hafellner & Kalb]
Ascospores 18–20 µm long; thallus and ascomata without any anthraquinone; eastern palaeotropical (Singapore) ... ... Pyrenula subglabrata (Nyl.) Müll. Arg.
(Verrucaria subglabrata Nyl.)
114(110) Ascospores 30–45 µm long ... 115
Ascospores <30 µm long ... 116
115(114) Ascomata conical, emergent, sides spreading; eastern palaeotropical ... ... Pyrenula interducta (Nyl.) Zahlbr.
(Verrucaria interducta Nyl.)
Ascomata globose, immersed; neotropical ... ... Pyrenula mastophoroides (Nyl.) Zahlbr.
[Verrucaria mastophoroides Nyl., Verrucaria mastophoroides var. flavicans Nyl., Pyrenula mastophoroides var. flavicans (Nyl.) Zahlbr.]
116(114) Ascospores mostly <21 µm long ... 117
Ascospores mostly >20 µm long ... 118
117(116) Ascospores mostly <17 µm long, biseriate in the ascus; pantropical ... ... Pyrenula castanea (Eschw.) Müll. Arg.
(Verrucaria hymnothora ssp. castanea Eschw., Spermatodium amazonicum Trevis., Pyrenula limae Vain., Pyrenula mamillana var. bataana Vain., Pyrenula limayensis Vain., Pyrenula submarginata Vain.)
Notes: Pyrenula submarginata is lectotypified here with specimen no. 23 (TUR-Vainio 31312); the other two original specimens do not have an inspersed hymenium. The ascospores of Pyrenula limayensis are shorter than in the description, up to 15 µm long.
Ascospores mostly 17–21 µm long, uniseriate in the ascus; pantropical ... ... Pyrenula mamillana (Ach.) Trevis.
[Verrucaria mamillana Ach., Verrucaria kunthii Fée., Pyrenula kunthii (Fée) Fée, Verrucaria cinchonae Fée, Pyrenula cinchonae (Fée) Fée (nom. illeg.), Verrucaria phaea Eschw. (nom. illeg.), Trypethelium ocellatum Zenker, Verrucaria marginata Hook., Pyrenula marginata (Hook.) Trevis., Verrucaria xyloides Eschw., Pyrenula xyloides (Eschw.) Müll. Arg., Melanomma subconicum Ellis & Everh., Pyrenula velata Müll. Arg., Verrucaria punctiformis Eschw. (nom. illeg.), ? Verrucaria papilligera Krempelh. (nom.illeg.), Verrucaria imitans Nyl., Pyrenula imitans (Nyl.) Zahlbr., Pyrenula lagoensis Müll. Arg., Verrucaria warmingii Krempelh., Melanotheca subsimplex Müll.Arg., Pyrenula warmingii (Krempelh.) Müll. Arg., ? Pyrenula mamillana var. subconfluens Vain, Pyrenula subconfluens (Vain.) Vain., Pyrenula subglabriuscula Vain., ? Pyrenula subglabriuscula var. natalensis Vain., Pyrenula philippina Vain., Pyrenula comirana Vain., Pyrenula marginata var. australasiatica Vain., Pyrenula fuscolurida Vain., Pyrenula atrofuscescens Vain., Pyrenula cinereovelata Vain., Pyrenula affinis Malme, Pyrenula oligocarpa Malme, ? Pyrenula philippina var. oceanica Räsänen ex Sbarbaro, Pyrenula columellata Upreti & Ajay Singh, Pyrenula elegans Ajay Singh & Upreti]
118(116) Ascospores rounded, uniseriate in the ascus; pantropical ... ... Pyrenula massariospora (Starb.) R.C. Harris
[Clypeosphaeria massariospora Starb., Starbaeckiella massariospora (Starb.) H. Syd. & Syd., Pyrenula pachyspora Vain., Pseudopyrenula majuscula H. Magn.]
Ascospores at least at one end pointed, biseriate in the ascus; neotropical ... ... Pyrenula acutalis R.C. Harris
119(107) Thallus UV+yellow (when material well preserved, sometimes only part of the thallus reacting) ... 120
Thallus UV− or greenish/whitish reflecting ... 123
120(119) Ascospores mostly >24 µm long ... 121
Ascospores mostly <24 µm long ... 122
121(120) Ascospores 25–40 µm long, without black granules; neotropical (Costa Rica) ... ... Pyrenula andina Aptroot
Ascospores 36–45 µm long, with black granules at the tips; temperate America ... ... Pyrenula caryae R.C. Harris
122(120) Ascospores 9–11 µm long, ellipsoid; Australian ... ... Pyrenula xanthominuta Aptroot
Ascospores 14–21 µm long, fusiform, pointed; cosmopolitan ... ... Pyrenula dermatodes (Borrer) Schaer.
[Verrucaria dermatodes Borrer, Verrucaria nitida var. dermatodes (Borrer) Leight., Pyrenula nitida var. dermatodes (Borrer) Trevis., ? Pyrenula viridescens Fée, Spermatodium viridescens (Fée) Trevis., Pyrenula mollis Fée, ? Pyrenula porinoides Fée (nom. illeg.), Melanotheca feeana Müll. Arg., Pseudopyrenula galactina Shirley, Pyrenula galactina (Shirley) Kantvilas, Pyrenula chloroplaca Shirley, Chrooicia inconspicua (C.F.W. Meissn.) Trevis., Melanotheca inconspicua (C.F.W. Meissn.) Müll. Arg., Pyrenula hypophyta (Nyl.) Müll. Arg., Verrucaria hypophyta Nyl., Verrucaria micromma Nyl. (nom. illeg.), Pyrenula micromma Shirley (nom. illeg.), Verrucaria micromma var. leucomma Nyl., Pyrenula micromma var. leucomma (Nyl.) Trevis., Pyrenula occulta var. leucomma (Nyl.) Zahlbr., Verrucaria punctella var. exstans Nyl., Pyrenula punctella var. exstans (Nyl.) Müll. Arg., Pyrenula pinguis var. exstans (Nyl.) Zahlbr., Verrucaria arverna Nyl., Melanotheca arverna (Nyl.) Boistel, Verrucaria arthoniza C. Knight, Pyrenula arthoniza (C. Knight) Müll. Arg., Verrucaria nitida var. pseudonitidella C. Knight, Pyrenula pseudonitidella (C. Knight) D.J. Galloway, Pyrenula annulata Müll. Arg., Pyrenula subimmersa Müll. Arg, Pyrenula diffracta Müll. Arg., Verrucaria occulta C. Knight, Pyrenula occulta (C. Knight) Müll. Arg., Pyrenula hypophytoides Harm., Verrucaria obvoluta Nyl., Pseudopyrenula obvoluta (Nyl.) Zahlbr., Pyrenula obvoluta (Nyl.) R.C. Harris & Aptroot, Verrucaria achroopora Nyl., Pyrenula achroopora (Nyl.) Arnold, Verrucaria glabratula Nyl., Pyrenula glabratula (Nyl.) Arnold, Verrucaria subtrahens Nyl., Pyrenula subtrahens (Nyl.) Müll. Arg., Pyrenula porinella Vain., ?(UV test needed) Pyrenula nitidella var. exstantior Vain., Pyrenula stramineoatra Vain., ? Pyrenula chondrina Zahlbr., Pyrenula tunicata Zahlbr., Pyrenula straminescens Zahlbr., Pyrenula schutschensis Zahlbr., Pyrenula lucifera R.C. Harris]
Note: somewhat variable, especially in UV-reaction, but always with pointed ascospores and thallus at least partly covering the ascomata. The oldest name found for specimens that are not reacting with UV is Pyrenula subtrahens (Nyl.) Müll. Arg.
123(119) Ascospores mostly >25 µm long ... 124
Ascospores mostly <25 µm long ... 135
124(123) Ascospores 36–45 µm long, with or without black granules at the tips ... 125
Ascospores <40 µm long, without black granules at the tips ... 126
125(124) Ascospores 36–45 µm long, with black granules at the tips; temperate America ... ... Pyrenula caryae R.C. Harris
Ascospores 36–45 µm long, without black granules at the tips; neotropical ... ... Pyrenula subducta (Nyl.) Müll. Arg.
[Verrucaria subducta Nyl., ? Verrucaria subducta var. retracta Nyl., Pyrenula subducta var. retracta (Nyl.) Zahlbr., Verrucaria marginata var. convexa Nyl., Verrucaria convexa (Nyl.) Nyl., Pyrenula convexa (Nyl.) Müll. Arg.]
126(124) End lumina elongated, ascospores 24–29 µm long; neotropical (Virgin Islands) ... ... Pyrenula spectata R.C. Harris
(also as Pyrenula expectata R.C. Harris)
End lumina not elongated ... 127
127(126) Growing on rock, ascospores with dark bands between the lumina, 25–30 µm long; neotropical (Brazil) ... Pyrenula quarzitica Aptroot
Note: the only Pyrenula species that is known from rock.
Growing on bark, ascospores without dark bands ... 128
128(127) Ascomata mostly >0·7 mm diam ... 129
Ascomata mostly <0·7 mm diam ... 131
129(128) Thallus with pseudocyphellae; temperate regions of the old world ... ... Pyrenula macrospora (Degel.) Coppins & P. James
[Verrucaria nitida f. elaeodes Leight., Pyrenula nitida f. elaeodes (Leight.) A.L. Sm., Pyrenula nitida var. macrospora Degel., Pyrenula chlorospila var. macrospora (Degel.) Maas Geest., Pyrenula nitida var. grandispora Barchalov]
Thallus without pseudocyphellae ... 130
130(129) Ascospores 24–29 µm long, thallus thick; eastern palaeotropical (Papua New Guinea) ... Pyrenula media Aptroot
Ascospores 29–40 µm long, thallus thin; pantropical ... ... Pyrenula complanata (Mont.) Trevis.
[Verrucaria complanata Mont., Spermatodium complanata (Mont.) Trevis., Pyrenula macrocarpa A. Massal., Verrucaria ectypa Krempelh., Pyrenula ectypa (Krempelh.) Zahlbr., Verrucaria introducta Stirt., Pyrenula introducta (Stirt.) Zahlbr., Pyrenula cordatula Zahlbr.]
131(128) Ascospores with diamond-shaped lumina; temperate regions on both hemispheres ... Pyrenula chlorospila (Nyl.) Arnold
[Verrucaria chlorospila Nyl., Pyrenula nitida var. nitidella f. chlorospila (Nyl.) Keissl., Pyrenula nitidella var. chlorospila (Nyl.) Szatala, Pyrenula pinguis Chevall. (nom. illeg.), Verrucaria nitida var. pinguis (Westr.) Nyl., Verrucaria pinguis Westr., Pyrenula nitens f. pinguis (Westr.) Zahlbr., Pyrenula nitida f. pinguis (Westr.) Zahlbr., Arthopyrenia nitida f. pinguis (Westr.) H. Olivier, Arthopyrenia nitida var. nitidella f. pinguis (Westr.) Boistel, Verrucaria chlorospiloides Nyl., Verrucaria nitida var. phaeospila Nyl., Pyrenula nitida var. phaeospila (Nyl.) Zahlbr., ? Verrucaria olivaceofusca C. Knight, Pyrenula knightiana Müll. Arg., Pyrenula mamillana var. erubescens Zahlbr., ? Verrucaria glabrata var. deprimens C. Knight, Pyrenula deprimens (C. Knight) D.J. Galloway, Pyrenula nitidella var. maculata R.C. Harris, Pyrenula maculata (R.C. Harris) R.C. Harris]
Note: specimen H-NYL 1165 (Portugal, Welwitsch 1840) is selected as the lectotype of Verrucaria nitida var. phaeospila Nyl. It also includes (and even more) Pyrenula macrospora, but the smaller syntype H-NYL 1166 shows what element was meant.
Ascospores with somewhat rounded or quadrangular lumina ... 132
132(131) Ascospores with dark granules between the lumina; temperate American ... ... Pyrenula macounii R.C. Harris
Ascospores without dark bands ... 133
133(132) Ascospores 32–42 µm long; temperate America, extending to Japan ... ... Pyrenula punctella (Nyl.) Trevis.
(Verrucaria punctella Nyl., Pyrenula impressa Müll. Arg.)
Ascospores mostly 25–37 µm long; tropical ... 134
134(133) Central lumina much wider than long, ascomata conical, emergent, thallus without pseudocyphellae; pantropical ... ... Pyrenula mastophora (Nyl.) Müll. Arg.
(Verrucaria mastophora Nyl., Pyrenula mastophora var. australis Malme, ? Pyrenula sessilis H. Magn.)
Central lumina more or less rounded, ascomata somewhat rounded, often partly immersed in the thallus, thallus often with (generally sparse) pseudocyphellae; pantropical ... Pyrenula quassiaecola (Fée) Fée
[also as Pyrenula quassiicola (Fée) Fée, Verrucaria quassiaecola Fée, also as Verrucaria quassiicola Fée, Verrucaria nitida var. quassiaecola (Fée) Nyl., also as Verrucaria nitida var. quassiicola (Fée) Nyl., Pyrenula pinguis Fée, Verrucaria pinguis (Fée) Spreng., ? Pyrenula vitrea (Eschw.) Müll. Arg., Verrucaria vitrea Eschw., Leiophloea vitrea (Eschw.) Trevis., Pyrenula flaventior Müll. Arg., Verrucaria flaventior (Müll. Arg.) Stirt., ? Verrucaria crassescens Stirt., Pyrenula crassescens (Stirt.) Müll. Arg., Stigmatidium confluens C. Knight, Stigmatidium prominulum C. Knight, ? Pyrenula mammillaris (Hepp) Zahlbr., Verrucaria mammillaris Hepp, Verrucaria subpunctella Nyl., Pyrenula subpunctella (Nyl.) Müll. Arg., Verrucaria mastophora Nyl., ? Verrucaria subnitida Nyl. (nom. inval.), Pyrenula subnitida Müll. Arg., ? Verrucaria glabrata var. homalisma C. Knight, Pyrenula homalisma (C. Knight) D.J. Galloway, Verrucaria baileyi C. Knight, Pyrenula baileyi (C. Knight) Shirley, Trypethelium papillatum C. Knight, Parmentaria papillata (C. Knight) Shirley, Pyrenula glaziovii Müll. Arg., Pyrenula ferax Müll. Arg., Pyrenula olivaceofusca Müll. Arg., Pyrenula nitidans Müll. Arg., Pyrenula pulchella var. cinerascens Müll. Arg., Pyrenula adacta var. cinerascens (Müll. Arg.) Zahlbr., Pyrenula punctella var. emergens Müll. Arg., Pyrenula pinguis var. emergens (Müll. Arg.) Müll. Arg., Verrucaria pinguis var. emergens (Müll. Arg.) Stizenb., Pyrenula emergens (Müll. Arg.) Vain., Pyrenula subcuprea Müll. Arg., Pyrenula virescens Müll. Arg., Pyrenula endostega Müll. Arg., Pyrenula parvula Müll. Arg., Pyrenula mastophorizans Müll. Arg., Pyrenula marmorata Müll. Arg., Pyrenula defossa Müll. Arg., Pyrenula coerulescens Müll. Arg., also as Pyrenula caerulescens Müll. Arg., Pyrenula obscurata Müll. Arg., Pyrenula subvelata Müll. Arg. (nom. illeg.), Pyrenula pseudovelata M. Choisy, Pyrenula gibberosa Vain., Pyrenula eucalypta Vain., ? Pyrenula mozambica Vain., Pyrenula orofensis Vain., Pyrenula euphorbiae Vain., Pyrenula martinicana Vain., Pyrenula rizalensis Vain., Melanotheca rizalensis (Vain.) Zahlbr., Pyrenula punctifera Vain., Pyrenula pallidofulvescens Vain., Anthracothecium olivaceocinereum Vain., Pyrenula vanoverberghii Vain., Pyrenula trombetana Vain., Pyrenula pallidofulvescens var. fulvostraminea Vain., Pyrenula oculifera Vain., Pyrenula obscurior Vain., Pyrenula pudica Zahlbr., Pyrenula manhaviensis Zahlbr., Pyrenula pertusarina Zahlbr., Pyrenula kelungana Zahlbr., Pyrenula chungii Zahlbr., Pyrenula nebulosa Zahlbr., Pyrenula albidopunctata Zahlbr., Pyrenula emersa Malme, Pyrenula bonariensis Malme, Pyrenula fulvescens Malme, Pyrenula emersa var. rissoensis Malme, Pyrenula plumbea Malme, Pyrenula fissa H. Magn., Pyrenula obscura Räsänen]
135(123) Ascospores mostly 21–25 µm long ... 136
Ascospores mostly <21 µm long ... 142
136(135) Ascomata c. 3–4 mm diam; Japan ... Pyrenula gigas Zahlbr.
Ascomata <3 mm diam ... 137
137(136) Ascomata with red, KOH+ purple crystals inside ... 138
Ascomata without red crystals ... 139
138(137) Ascomata <0·5 mm diam; temperate regions of Europe and Asia ... ... Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
[Verrucaria nitida var. nitidella Flörke ex Schaer., Verrucaria nitidella (Flörke ex Schaer.) Nyl., Bunodea nitida var. nitidella (Flörke ex Schaer.) Beltr., Pyrenula nitida var. nitidella (Flörke ex Schaer.) Schaer., Arthopyrenia nitida var. nitidella (Flörke ex Schaer.) H. Olivier, Pyrenula nitidella var. cintrana Welw., Pyrenula nitida var. minor Hepp, also as Pyrenula nitida var. minima Hepp Pyrenula nitida var. minor f. pinicola Hepp, also as Pyrenula nitida var. minima f. pinicola Hepp, Pyrenula nitidella f. pinicola (Hepp) Zahlbr., Pyrenula nitidella var. nigrescens B. de Lesd., Pyrenula nitida var. nitidella f. nigrescens (B. de Lesd.) Keissl., Pyrenula nitida var. aequata Zahlbr., Pyrenula nitida var. nitidella f. fuscata Suza]
Ascomata >0·5 mm diam; temperate regions of Europe and Asia ... ... Pyrenula nitida (Weigel) Ach.
[Lichen alveolatus Scop., Sphaeria nitida Weigel, Lichen nitidus (Weigel) Ach., Verrucaria nitida (Weigel) Schrad., Bunodea nitida (Weigel) Beltr., Arthopyrenia nitida (Weigel) H. Olivier., Verrucaria maxima DC., Verrucaria nitida var. maxima (DC.) March., Lichen populneus Ach., Verrucaria populnea (Ach.) DC, Pyrenula glabrata f. pachyderma Haszl., Pyrenula laevigata f. pachyderma (Haszl.) Zahlbr., Pyrenula nitida f. pachyderma (Haszl.) Szatala, Pyrenula nitida f. chevalieri M. Choisy, Verrucaria nitida var. populi Saint-Amans, Verrucaria nitida f. flavescens Malbr., Arthopyrenia nitida f. flavescens (Malbr.) H. Olivier, Arthopyrenia nitida var. nitidella f. flavescens (Malbr.) Boistel, Pyrenula nitida f. flavescens (Malbr.) Zahlbr., Verrucaria nitida f. squamata Malbr., Arthopyrenia nitida f. squamata (Malbr.) H. Olivier, Pyrenula nitida f. squamata (Malbr.) Zahlbr., Verrucaria nitida var. major Schaer., Pyrenula nitida var. major (Schaer.) Schaer., Pyrenula nitida f. virens Servít & Nádvorník, Pyrenula nitida f. diffracta Erichs., Pyrenula nitida f. crassa Barchalov]
139(137) Ascospores with angular diamond-shaped lumina; temperate America ... ... Pyrenula micheneri R.C. Harris
Ascospores with rounded or quadrangular lumina ... 140
140(139) Ascospores with bands of dark granules between the lumina; pantropical ... ... Pyrenula pyrgillospora Aptroot
Ascospores without bands of dark granules ... 141
141(140) Ascospores with at least one pointed end; temperate Northern Hemisphere, extending to the tropics ... Pyrenula acutispora Kalb & Hafellner
[Pyrenula kakouettae Sérus. & Diederich, Pyrenula alnicola R.C. Harris (nom. inval.)]
Ascospores with rounded ends; eastern palaeotropical (India) ... ... Pyrenula submastophora Ajay Singh & Upreti
142(135) Ascospores 7–10 µm long; Iran ... ... Pyrenula minutissima Aptroot, Valadbeigi & Sipman ined.
Ascospores >10 µm long ... 143
143(142) Ascospores mostly <15 µm long ... 144
Ascospores mostly >15 µm long ... 146
144(143) Ascomata >0·7 mm diam; pantropical ... ... Pyrenula atropurpurea (Eschw.) Müll. Arg.
(Verrucaria atropurpurea Eschw.)
Ascomata <0·7 mm diam ... 145
145(144) Ascospores 6–8 µm wide; pantropical ... Pyrenula brunnea Fée
[Microthelia shirleyana Müll. Arg., Pyrenula shirleyana (Müll. Arg.) Aptroot, ? Pyrenula subgriseola Vain., Melanotheca subgriseola (Vain.) Zahlbr.]
Ascospores 4–6 µm wide; pantropical ... Pyrenula aspistea (Ach.) Ach.
[Verrucaria aspistea Ach., Polyblastia aspistea (Ach.) Trevis., Pyrenula nitida var. aspistea (Ach.) Trevis, ? Pyrenula porinoides Ach., Ocellularia porinoides (Ach.) Spreng., Verrucaria porinoides (Ach.) Mont., Segestria porinoides (Ach.) Trevis., Pyrenula heteroclita Ach., Verrucaria heteroclita (Ach.) Spreng., Verrucaria aggregata f. heteroclita (Ach.) Nyl., Pyrenula heteroclita ssp. minuscula Ach., Pyrenula heteroclita ssp. denigrata Ach., Pyrenula glauca (Fée) Müll. Arg., Verrucaria glauca Fée, Spermatodium glaucum (Fée) Trevis., ? Verrucaria viridescens Fée, Spermatodium viridescens (Fée) Trevis., Verrucaria decolorata Fée, Verrucaria bonplandiae Fée, Pyrenula bonplandiae (Fée) Fée, Pyrenula subtrahens var. microspora (Krempelh.) Zahlbr., Verrucaria subtrahens var. microspora Krempelh., Verrucaria seriata Hepp, Pyrenula seriata (Hepp) Müll. Arg., Verrucaria aggregata Nyl., Verrucaria aggregata f. segregata Nyl., Pyrenula segregata (Nyl.) Müll. Arg, Pyrenula subaggregata Müll. Arg., Pyrenula velatior Müll. Arg., Pyrenula nigrocincta Müll. Arg., Pyrenula umbilicatula Müll. Arg., Pyrenula atrofusca Müll. Arg., Pyrenula minutula Müll. Arg., Pyrenula exigua Müll. Arg., Pyrenula dispora Müll. Arg., Pyrenula peltophora Müll. Arg., Pyrenula indusiata Müll. Arg., Massaria bataanensis Rehm, Pyrenula bataanensis (Rehm) Shoemaker & P.M. LeClair, Pyrenula apayaoensis Vain., Pyrenula guimarana Vain., Pyrenula bilirana Vain., Pyrenula ochracea Szatala, Pyrenula rubidopunctata Szatala, Pyrenula aquila R.C. Harris, Pyrenula subrizalensis Ajay Singh & Upreti]
Note: only the lectotype (designated here) of Pyrenula umbilicatula:(Cuba, Wright, Verr. Cub. 218a p.p.; G, as Pertusaria umbilicatula) is Pyrenula aspistea; the specimen in G of 218b p.p. under the name Pertusaria umbilicatula is Pyrenula mamillana.
146(143) Thallus at least partly 0·3 mm thick ... 147
Thallus thinner ... 148
147(146) Ascospores at least partly constricted at the septa; Australasian (Papua New Guinea) ... Pyrenula montana Aptroot
Ascospores not constricted; neotropical (Puerto Rico) ... ... Pyrenula psoriformis Zahlbr.
148(146) Ascomata >0·7 mm diam ... 149
Ascomata <0·7 mm diam ... 151
149(148) Thallus with pseudocyphellae; New Zealand ... ... Pyrenula deliquescens (C. Knight) Müll. Arg.
(Verrucaria deliquescens C. Knight, Verrucaria astata C. Knight, Pyrenula consociata Zahlbr.)
Thallus without pseudocyphellae ... 150
150(149) Lumina rounded; eastern palaeotropical (India) ... ... Pyrenula scutata (Stirt.) Zahlbr.
(Verrucaria scutata Stirt., Pyrenula pileata Vain.)
Note: examination of the type showed that the name Pyrenula pileata has recently been incorrectly used for Pyrenula fetivica.
Lumina angular; pantropical ... Pyrenula balia (Krempelh.) R.C. Harris
[Verrucaria balia Krempelh., Pseudopyrenula balia (Krempelh.) Müll. Arg., Verrucaria marginata var. santensis Nyl., Verrucaria santensis (Nyl.) Nyl., Pyrenula mamillana var. santensis (Nyl.) Trevis., Pyrenula marginata var. santensis (Nyl.) Tuck., Pyrenula santensis (Nyl.) Müll. Arg., Pyrenula nitida var. commutata Trevis., Verrucaria hymnothora Krempelh. (nom. illeg.), Pyrenula deplanata Müll. Arg., Pyrenula longislandica Ajay Singh & Upreti]
151(148) Ascospores with extra endospore layer around lumina; neotropical (Florida, Louisiana) ... Pyrenula rubrostoma R.C. Harris
Ascospores without extra endospore layers ... 152
152(151) Ascospores with dark bands between the lumina; North America extending to Japan ... Pyrenula confoederata R.C. Harris
Ascospores without dark bands ... 153
153(152) Ascomata <0·4 mm diam ... 154
Ascomata >0·4 mm diam; pantropical (UV− material of Pyrenula dermatodes keys out here as well) ... Pyrenula aggregata (Fée) Fée
[Verrucaria aggregata Fée, Spermatodium aggregatum (Fée) Trevis., Melanotheca aggregata (Fée) Müll. Arg. (nom. illeg.), Verrucaria subnitidella Nyl., Pyrenula subnitidella (Nyl.) Müll. Arg., Pyrenula costaricensis Müll. Arg., Pyrenula gracilior Müll. Arg., Pyrenula virens Müll. Arg., Pyrenula tenella Müll. Arg., Pyrenula fuscoolivacea Vain., Pyrenula aspisteoides Vain., Pyrenula parva Vain., Pyrenula irrubescens Vain., Pseudopyrenula awajiensis Vain., Pyrenula awajiensis (Vain.) Kashiw., Pyrenula oblonga Zahlbr., Pyrenula athallina H. Magn., Pyrenula nuda Ajay Singh & Upreti]
Note: the type of Pyrenula aggregata shows only crowded ascomata, no joint walls.
154(153) Ascospores 18–20 µm long; neotropical ... Pyrenula tristissima Vain.
Ascospores <18 µm long; neotropical ... Pyrenula minor Fée
[Spermatodium minus (Fée) Trevis.]
Nomenclatural novelties
Anthracothecium interlatens (Nyl.) Aptroot comb. nov
MycoBank No.: MB563101
Basionym: Astrothelium interlatens Nyl., Bull. Soc. Linn. Normand., ser. 2, 2: 134 (1868); type: New Caledonia, Lifu, 1864, Thiébaut (H-NYL 82 —lectotype, selected here).
Parmentaria interlatens (Nyl.) Müll. Arg., Parmentaria toowoombensis Müll. Arg., Anthracothecium toowoombense (Müll. Arg.) Aptroot, Astrothelium interlatens var. nudatum Nyl., Parmentaria interlatens var. nudatum (Nyl.) Zahlbr., Parmentaria grossa Müll. Arg., Parmentaria subastroidea Müll. Arg., Parmentaria denudata Zahlbr., Plagiothelium australiense Stirt., Parmentaria australiense (Stirt.) Müll. Arg., Parmentaria chungii Zahlbr., Parmentaria obtecta Zahlbr., Parmentaria capensis Zahlbr., Anthracothecium capense (Zahlbr.) K.P. Singh & G.P. Singha, Parmentaria nilamburensis Makhija & Patw., Parmentaria indica Upreti & Ajay Singh.
Note: specimen H-NYL 80 (New Caledonia, Lifu, Thiébout 1864) is selected as lectotype. The specimens H-NYL 77, 78 and 82 are identical.
Pyrenula breutelii (Müll. Arg.) Aptroot comb. nov
MycoBank No.: MB563102
Basionym: Anthracothecium breutelii Müll. Arg., Flora 68: 339 (1885); holotype: St Thomas, Breutel, ex hb. Hampe 1877 (G).
Anthracothecium maculare Zahlbr.; Pyrenula macularis (Zahlbr.) R.C. Harris, Anthracothecium speciosum Zahlbr.
Notes. Anthracothecium breutelii is the oldest epithet for this distinctive species. It was not surprising that an older name should be found for this common and widespread pantropical species. The holotype of Anthracothecium speciosum is mixed and contains also Pyrenula leucostoma, but the material distributed in Zahlbruckner's Lichenes Rariores Exsiccati 252 is all Pyrenula macularis.
Pyrenula ceylonensis (Ajay Singh & Upreti) Aptroot comb. nov
MycoBank No.: MB563103
Basionym: Pleurotheliopsis ceylonensis Ajay Singh & Upreti, Geophytology 16: 262 (1986).
Parmentaria anamalaiensis Upreti & Ajay Singh, Pyrenula anamalaiensis (Upreti & Ajay Singh) Upreti, Parmentaria oligocarpa Ajay Singh, Pyrenula suboligocarpa Upreti.
Pyrenula fusispora (Malme) Aptroot comb. nov
MycoBank No.: MB563104
Basionym: Parathelium fusisporum Malme, Ark. Bot. 19(1): 17 (1924); holotype: Brazil, Matto Grosso, Curumbá, Malme, 19 July 1894 (S).
Pyrenula gibberulosa (Vain.) Aptroot comb. nov
MycoBank No.: MB563105
Basionym: Bottaria gibberulosa Vain., Annal. Acad. Sci. Fenn., ser. A, 15(6): 329 (1921).
Anthracothecium gibberulosum (Vain.) Zahlbr., Pyrenula subochraceoflavens Upreti, Anthracothecium goaense Ajay Singh.
Note: an isotype studied of Pyrenula subochraceoflavens revealed the submuriform ascospores and the lack of orange pigmentation (it is not more pigmented than Pyrenula occidentalis).
Pyrenula lyoni (Zahlbr.) Aptroot comb. nov
MycoBank No.: MB563106
Basionym: Parmentaria lyoni Zahlbr., Ann. Mycol. 10: 363 (1912).
Parmentaria lyoni f. straminescens Zahlbr., Polyblastiopsis negrosensis Herre, Anthracothecium keralense Patw. & Makhija, Parmentaria keralensis (Patw. & Makhija) Ajay Singh.
Pyrenula neosandwicensis Aptroot nom. nov
MycoBank No.: MB563107
Nom. nov. pro: Anthracothecium sandwicense Zahlbr., Ann. Mycol. 10: 361 (1912).
Anthracothecium sandwicense var. globosum Zahlbr.
Pyrenula papillifera (Nyl.) Aptroot comb. nov
MycoBank No.: MB563108
Basionym: Verrucaria papillifera Nyl., Espos. Synopt. Pyrenocarp: 42 (1858).
Anthracothecium papilliferum (Nyl.) Müll. Arg., Sporodictyon papilliferum (Nyl.) Trevis., Verrucaria thwaitesii Leight., Anthracothecium thwaitesii (Leight.) Müll. Arg., Bottaria thwaitesii (Leight.) Vain., Bottaria mucosa Vain., Anthracothecium mucosum (Vain.) Zahlbr., Pyrenula mucosa (Vain.) R.C. Harris, Bottaria albidopallens Vain., Anthracothecium albidopallens (Vain.) Zahlbr., Anthracothecium chrysophorum Zahlbr., Anthracothecium badioatrum Ajay Singh.
Note: Anthracothecium chrysophorum Zahlbr. was described as having a yellow medulla, but there is no pigment present in the type and only specimen.
Pyrenula platystoma (Müll. Arg.) Aptroot comb. nov
MycoBank No.: MB563109
Basionym: Anthracothecium platystomum Müll. Arg., Revue Mycol. 10: 124 (1888); lectotype: Paraguay, Guarapi, 1878 Balansa 4172, (G, also 2 identical isolectotypes).
Anthracothecium oculatum Müll. Arg., Pyrenula neoculata Aptroot, Bottaria impressa Vain., Anthracothecium impressum (Vain.) Zahlbr., Anthracothecium biferum Zahlbr., Bottaria himalayensis Räsänen, Anthracothecium himalayense (Räsänen) D.D. Awasthi, Anthracothecium vermiculare Kashiw. & Kurok., also as Anthracothecium vermicularis Kashiw. & Kurok., Pyrenula vermicularis (Kashiw. & Kurok.) H. Harada, Anthracothecium pseudohimalayense Ajay Singh, Anthracothecium himalayense var. pseudohimalayense (Ajay Singh) Ajay Singh, Anthracothecium mulleri Patw. & Makhija, Polyblastiopsis muelleri Upreti & Ajay Singh, Anthracothecium lifuense Upreti & Ajay Singh, Anthracothecium bengalense Ajay Singh.
Pyrenula schiffneri (Zahlbr.) Aptroot comb. nov
MycoBank No.: MB563110
Basionym: Parmentaria schiffneri Zahlbr., Denkschr. math.-naturw. Classe, Akad. Wissensch. Wien 83: 95 (1909).
Anthracothecium falsarium Zahlbr., Pyrenula falsaria (Zahlbr.) R.C. Harris, Pyrenula verruculosa Upreti & Ajay Singh.
Note: the ascospores of Parmentaria schiffneri are much smaller than mentioned in the original description.
Pyrenula sexluminata Aptroot nom. nov
MycoBank No.: MB563111
Nom. nov. pro: Pyrenula quinqueseptata Aptroot, Bibliotheca Lichenologica 64: 167 (1997).
Pyrenula welwitschii (Upreti & Ajay Singh) Aptroot comb. nov
MycoBank No.: MB563112
Basionym: Anthracothecium welwitschii Upreti & Ajay Singh, Feddes Repert. Spec. Nov. Regni Veg. 99: 151 (1988).
Pyrenula himalayana Upreti.
Note: an isotype studied of Pyrenula himalayana showed that it contains no anthraquinones on the thallus or ascomata.
Sulcopyrenula subglobosa (Riddle) Aptroot comb. nov
MycoBank No.: MB563113
Basionym: Anthracothecium subglobosum Riddle, in Riddle & Millspaugh, Bahama Fl.: 527 (1920).
The curators of the lichen herbaria of B, BM, BRI, CANB, G, GZU, H, HO, L, LI, M, S, TNS, TUR, UPS and W are warmly thanked for their often prompt cooperation. Robert Lücking is thanked for putting his pictures of lichen types (TROPILIT project) at my disposal.
Index with all names in Anthracothecium and Pyrenula, with their disposition (A and B refer to the two keys, numbers to the key entries)
Anthracothecium albescens (Nyl.) Müll. Arg. A38 = Pyrenula confinis (Nyl.) R.C. Harris
Anthracothecium albescens var. subochraceum (Nyl.) Zahlbr. A27 = Pyrenula ochraceoflava (Nyl.) R.C. Harris
Anthracothecium albidopallens (Vain.) Zahlbr. A48 = Pyrenula papillifera (Nyl.) Aptroot
Anthracothecium americanum (Ach.) Müll. Arg. A46 = Pyrenula leucostoma Ach.
Anthracothecium amphitropum Müll. Arg. A48 = Pyrenula pyrenuloides (Mont.) R.C. Harris
Anthracothecium andamanicum (Nyl.) Müll. Arg. A8 = Anthracothecium macrosporum (Hepp) Müll. Arg.
Anthracothecium angolense Upreti & Ajay Singh A39 = Pyrenula parvinuclea (Meyen & Flot.) Aptroot
Anthracothecium angulatum Zahlbr. A8 = Anthracothecium prasinum (Eschw.) R.C. Harris
Anthracothecium angulatum var. majus Zahlbr. A8 = Anthracothecium prasinum (Eschw.) R.C. Harris
Anthracothecium anoistum (Stirt.) Zahlbr. = Polymeridium proponens (Nyl.) R.C. Harris
Anthracothecium asahinae Kashiw. & Kurok. A21 = Pyrenula subvariolosa (C. Knight) Aptroot
Anthracothecium assamiense (Stirt.) Ajay Singh A44 = Pyrenula duplicans (Nyl.) Aptroot
Anthracothecium aurantiacum (Eschw.) Müll. Arg. (nom. illeg.) = Astrothelium conicum aggr.
Anthracothecium aurantium (Eschw.) Müll. Arg. = Astrothelium conicum aggr.
Anthracothecium australiense (Müll. Arg.) Aptroot A7
Anthracothecium austroindicum Ajay Singh A42 = Pyrenula globifera (Eschw.) Aptroot
Anthracothecium awasthii Ajay Singh A39 = Pyrenula parvinuclea (Meyen & Flot.) Aptroot
Anthracothecium badioatrum Ajay Singh A48 = Pyrenula papillifera (Nyl.) Aptroot
Anthracothecium bengalense Ajay Singh A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium biferum Zahlbr. A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium borbonicum (Nyl.) Müll. Arg. A8 = Anthracothecium prasinum (Eschw.) R.C. Harris
Anthracothecium brasilianum M. Choisy = Astrothelium conicum aggr.
Anthracothecium breutelii Müll. Arg. A40 = Pyrenula breutelii (Müll. Arg.) Aptroot
Anthracothecium capense (Zahlbr.) K. P. Singh & G. P. Sinha A9 = Anthracothecium interlatens (Nyl. ) Aptroot
Anthracothecium canellae-albae (Fée) Müll. Arg. A12 = Sulcopyrenula canellae-albae (Fée) H. Harada
Anthracothecium cascarillae Müll. Arg. A51 = Pyrenula dissimulans (Müll. Arg.) R.C. Harris
Anthracothecium cellulosum (C. Knight) Müll. Arg. A50 = Pyrenula thelomorpha Tuck.
Anthracothecium chrysophorum Zahlbr. A48 = Pyrenula papillifera (Nyl.) Aptroot
Anthracothecium cinerosum (Ach.) Müll. Arg. A48 = ? Pyrenula pyrenuloides (Mont.) R.C. Harris
Anthracothecium coccineum Müll. Arg. A27 = Pyrenula ochraceoflava (Nyl.) R.C. Harris
Anthracothecium collosporum (Vain.) Zahlbr. A7 = Anthracothecium australiense (Müll. Arg.) Aptroot
Anthracothecium columellatum (Vain.) Zahlbr. A8 = Anthracothecium macrosporum (Hepp) Müll. Arg.
Anthracothecium confine (Nyl.) Müll. Arg. A38 = Pyrenula confinis (Nyl.) R.C. Harris
Anthracothecium corcovadense Malme A3 = Mycomicrothelia decipiens (Müll. Arg.) R.C. Harris
Anthracothecium corticatum Müll. Arg. A38 = Pyrenula confinis (Nyl.) R.C. Harris
Anthracothecium cristatellum Nagarkar & Patw. A48 = Pyrenula pyrenuloides (Mont.) R.C. Harris
Anthracothecium cruentatum (Müll. Arg.) Müll. Arg. A25 = Pyrenula cruentata (Müll. Arg.) R.C. Harris
Anthracothecium decipiens Müll. Arg. A3 = Mycomicrothelia decipiens (Müll. Arg.) R.C. Harris
Anthracothecium denudatum (Nyl.) Zahlbr. A39 = Pyrenula parvinuclea (Meyen & Flot.) Aptroot
Anthracothecium denudatum var. ochrotropum (Nyl.) Müll. Arg. A27 = Pyrenula ochraceoflava (Nyl.) R.C. Harris
Anthracothecium depressum (Eschw.) Müll. Arg. A48 = Pyrenula pyrenuloides (Mont.) R.C. Harris
Anthracothecium desquamans Müll. Arg. = a thelotremoid Graphidaceae, most recently as Leptotrema desquamans (Müll. Arg.) Patw. & Makhija
Anthracothecium dimorphum (Vain.) Zahlbr. B106 = Pyrenula nitidula (Bres.) R.C. Harris
Anthracothecium doleschallii A. Massal. A8 = Anthracothecium macrosporum (Hepp) Müll. Arg.
Anthracothecium duplicans (Nyl.) Müll. Arg. A44 = Pyrenula duplicans (Nyl.) Aptroot
Anthracothecium emergens (Leight.) Zahlbr. A46 = Pyrenula leucostoma Ach.
Anthracothecium eminentior (Nyl.) Müll. Arg. = Porina eminentior Nyl.
Anthracothecium endococcineum (Vain.) Zahlbr. A46 = Pyrenula leucostoma Ach.
Anthracothecium epapillatum (Nyl.) Müll. Arg. A42 = Pyrenula globifera (Eschw.) Aptroot
Anthracothecium erigens (Kashiw.) H. Harada A7 = Anthracothecium australiense (Müll. Arg.) Aptroot
Anthracothecium erythrinae (Vain.) Zahlbr. A46 = Pyrenula leucostoma Ach.
Anthracothecium eschweileri Müll. Arg. A8 = Anthracothecium prasinum (Eschw.) R.C. Harris
Anthracothecium euthelium (Nyl.) Zahlbr., Verrucaria euthelia Nyl. = Astrothelium sp.
Anthracothecium exsertum (Krempelh.) Müll. Arg. A44 = Pyrenula duplicans (Nyl.) Aptroot
Anthracothecium falsarium Zahlbr. A34 = Pyrenula schiffneri (Zahlbr.) Aptroot
Anthracothecium fraternale Zahlbr. A46 = Pyrenula leucostoma Ach.
Anthracothecium fulvum Müll. Arg. B60 = Pyrenula papilligera (Leight.) Müll. Arg.
Anthracothecium gibberulosum (Vain.) Zahlbr. A20 = Pyrenula gibberulosa (Vain.) Aptroot
Anthracothecium globiferum (Eschw.) Müll. Arg. A42 = Pyrenula globifera (Eschw.) Aptroot
Anthracothecium globiferum var. microsporum Ajay Singh A42 = ? Pyrenula globifera (Eschw.) Aptroot
Anthracothecium goaense Ajay Singh A20 = Pyrenula gibberulosa (Vain.) Aptroot
Anthracothecium goniostomum Müll. Arg. A48 = Pyrenula pyrenuloides (Mont.) R.C. Harris
Anthracothecium gregale (C. Knight) Aptroot A9
Anthracothecium guineense (Nyl.) Müll. Arg. A7 = Anthracothecium australiense (Müll. Arg.) Aptroot
Anthracothecium hexamerum Müll. Arg. B46 = Pyrenula sexlocularis (Nyl.) Müll. Arg.
Anthracothecium hians Müll. Arg. = a thelotremoid Graphidaceae, recently known as Thelotrema neohians Patw. & Makhija
Anthracothecium hibernicum (Nyl.) A.L. Sm. A35 = Pyrenula hibernica (Nyl.) Aptroot
Anthracothecium himalayense (Räsänen) D.D. Awasthi A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium himalayense var. pseudohimalayense (Ajay Singh) Ajay Singh A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium immersum Patw. & Makhija A31 = Pyrenula astroidea (Fée) R.C. Harris
Anthracothecium impressum (Vain.) Zahlbr. A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium inclinatum (Müll. Arg.) Kashiw. A31 = ? Pyrenula ravenelii (Tuck.) R.C. Harris
Anthracothecium indicum Ajay Singh A8 = Anthracothecium macrosporum (Hepp) Müll. Arg.
Anthracothecium interlatens (Nyl.) Aptroot A9
Anthracothecium interponens (Nyl.) Müll. Arg. A44 = Pyrenula duplicans (Nyl.) Aptroot
Anthracothecium japonicum Kashiw. & Kurok. A8 = Anthracothecium macrosporum (Hepp) Müll. Arg.
Anthracothecium javanicum (Hepp) Zahlbr. A6 = ? Pyrenula leucostoma Ach.
Anthracothecium keralense Patw. & Makhija A32 = Pyrenula lyoni (Zahlbr.) Aptroot
Anthracothecium laevigatum Müll. Arg. A19 = Lithothelium nanosporum (C. Knight) Aptroot
Anthracothecium leightonii Patw. & Makhija A48 = Pyrenula pyrenuloides (Mont.) R.C. Harris
Anthracothecium leucostomum Ach. A46 = Pyrenula leucostoma Ach.
Anthracothecium libricolum (Fée) Müll. Arg. A46 = Pyrenula leucostoma Ach.
Anthracothecium lifuense Upreti & Ajay Singh A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium lugescens (Nyl.) Zahlbr. = Campylothelium lugescens (Nyl.) Upreti & Ajay Singh
Anthracothecium luteonitens (Nyl.) Zahlbr. A8 = ? Anthracothecium macrosporum (Hepp) Müll. Arg.
Anthracothecium macrosporum (Hepp) Müll. Arg. A8
Anthracothecium maculare Zahlbr. A40 = Pyrenula breutelii (Müll. Arg.) Aptroot
Anthracothecium maculatum Nagarkar & Patw. A44 = Pyrenula duplicans (Nyl.) Aptroot
Anthracothecium majus (Zahlbr.) Kashiw. & Kurok. A8 = Anthracothecium prasinum (Eschw.) R.C. Harris
Anthracothecium manipurense Müll. Arg. A8 = Anthracothecium macrosporum (Hepp) Müll. Arg.
Anthracothecium megaspermum Patw. & Makhija A8 = Anthracothecium macrosporum (Hepp) Müll. Arg.
Anthracothecium melasporum (Taylor) Müll. Arg. = Henrica melaspora (Taylor) Savić & Tibell
Anthracothecium monosporum Müll. Arg. A2 = Aptrootia elatior (Stirt.) Aptroot
Anthracothecium mucosum (Vain.) Zahlbr. A48 = Pyrenula papillifera (Nyl.) Aptroot
Anthracothecium mulleri Patw. & Makhija A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium nanosporum Ajay Singh A39 = Pyrenula nanospora (Ajay Singh) Upreti
Anthracothecium nanum (Zahlbr.) R.C. Harris A7 = Anthracothecium australiense (Müll. Arg.) Aptroot
Anthracothecium novemseptatum (Vain.) R.C. Harris A21 = Pyrenula novemseptata Vain.
Anthracothecium obscuratum Upreti & Ajay Singh A46 = Pyrenula leucostoma Ach.
Anthracothecium ochraceoflavens (Nyl.) Zahlbr. A27 = Pyrenula ochraceoflavens (Nyl.) R.C. Harris
Anthracothecium ochraceoflavum (Nyl.) Müll. Arg. A27 = Pyrenula ochraceoflava (Nyl.) R.C. Harris
Anthracothecium ochraceoflavum f. nudius (Nyl.) Zahlbr. A27 = Pyrenula ochraceoflava (Nyl.) R.C. Harris
Anthracothecium ochrotropum (Nyl.) Zahlbr. A27 = Pyrenula ochraceoflava (Nyl.) R.C. Harris
Anthracothecium ochroxanthum Müll. Arg. A27 = Pyrenula ochraceoflava (Nyl.) R.C. Harris
Anthracothecium oculatum Müll. Arg. A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium oligosporum Müll. Arg. = a thelotremoid Graphidaceae, recently known as Leptotrema oligosporum (Müll. Arg.) Patw. & Makhija
Anthracothecium olivaceocinereum Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Anthracothecium opertum (Nyl.) Müll. Arg. A42 = Pyrenula globifera (Eschw.) Aptroot
Anthracothecium pachycheilum (Tuck.) Zahlbr. A46 = Pyrenula leucostoma Ach.
Anthracothecium palmarum (Krempelh.) Vain. A26 = Pyrenula palmarum (Krempelh.) R.C. Harris
Anthracothecium papilliferum (Nyl.) Müll. Arg. A48 = Pyrenula papillifera (Nyl.) Aptroot
Anthracothecium paraguayense Malme A42 = Pyrenula globifera (Eschw.) Aptroot
Anthracothecium parameroides (Vain.) Müll. Arg. A39 = Pyrenula parvinuclea (Meyen & Flot.) Aptroot
Anthracothecium paramerum (Nyl.) Müll. Arg. A46 = Pyrenula leucostoma Ach.
Anthracothecium paramerum f. pallidoalbum (Vain.) Zahlbr. A46 = Pyrenula leucostoma Ach.
Anthracothecium parvinucleum (Meyen & Flot.) Zahlbr. A39 = Pyrenula parvinuclea (Meyen & Flot.) Aptroot
Anthracothecium pauciloculare Herre B108 = Pyrenula subelliptica (Tuck.) R.C. Harris
Anthracothecium peltophorum Müll. Arg. B106 = Pyrenula mastophoriza (Nyl.) Zahlbr.
Anthracothecium planiusculum (Nyl.) Müll. Arg. = Anisomeridium planiusculum (Nyl.) R.C. Harris
Anthracothecium platystomum Müll. Arg. A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium praelustre (Krempelh.) Müll. Arg. A8 = Anthracothecium prasinum (Eschw.) R.C. Harris
Anthracothecium prasinum (Eschw.) R.C. Harris A8
Anthracothecium pseudoborbonicum Upreti & Ajay Singh A8 = Anthracothecium prasinum (Eschw.) R.C. Harris
Anthracothecium pseudocyphellatum Ajay Singh A44 = Pyrenula duplicans (Nyl.) Aptroot
Anthracothecium pseudohimalayense Ajay Singh A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium punctuliforme (Stizenb.) Müll. Arg. A12 = Sulcopyrenula canellae-albae (Fée) H. Harada
Anthracothecium pusillum (Ach.) Müll. Arg. A19 = Lithothelium nanosporum (C. Knight) Aptroot
Anthracothecium pustuliferum Ajay Singh A44 = Pyrenula duplicans (Nyl.) Aptroot
Anthracothecium pyrenuloides (Mont.) Müll. Arg. A48 = Pyrenula pyrenuloides (Mont.) R.C. Harris
Anthracothecium rhodesianum C.W. Dodge = unknown
Anthracothecium roseum (Vain.) Zahlbr. A27 = Pyrenula ochraceoflava var. pacifica P. M. McCarthy
Anthracothecium sandwicense Zahlbr. A44 = Pyrenula neosandwicensis Aptroot
Anthracothecium sandwicense var. convexum Zahlbr. A48 = Pyrenula pyrenuloides (Mont.) R.C. Harris
Anthracothecium sandwicense var. globosum Zahlbr. A44 = Pyrenula neosandwicensis Aptroot
Anthracothecium seminudum Müll. Arg. B46 = Pyrenula sexlocularis (Nyl.) Müll. Arg.
Anthracothecium sinapispermum (Fée) Müll. Arg. B69 = ? Pyrenula cruenta (Mont.) Vain.
Anthracothecium speciosum Zahlbr. A40 = Pyrenula breutelii (Müll. Arg.) Aptroot
Anthracothecium speciosum f. iturupiense O.B. Blum A8 = ? Anthracothecium prasinum (Eschw.) R.C. Harris
Anthracothecium staurosporum (Tuck.) Zahlbr. A12 = Sulcopyrenula staurospora (Tuck.) H. Harada
Anthracothecium subcutaneum Müll. Arg. A46 = Pyrenula leucostoma Ach.
Anthracothecium subglobosum Riddle A13 = Sulcopyrenula subglobosa (Riddle) Aptroot
Anthracothecium sublaevigatum Patw. & Makhija A45 = Pyrenula sublaevigata (Patw. & Makhija) Upreti
Anthracothecium submucosum (Vain.) Zahlbr. A46 = Pyrenula leucostoma Ach.
Anthracothecium subochraceum (Nyl.) Zahlbr. A27 = Pyrenula ochraceoflava (Nyl.) R.C. Harris
Anthracothecium subvariolosum C. Knight A21 = Pyrenula subvariolosa (C. Knight) Aptroot
Anthracothecium tetraspermum Riddle A12 = Sulcopyrenula staurospora (Tuck.) H. Harada
Anthracothecium thelemorphum (Tuck.) Zahlbr. A50 = Pyrenula thelomorpha Tuck.
Anthracothecium thelomorphum (Tuck.) Zahlbr. A50 = Pyrenula thelomorpha Tuck.
Anthracothecium thwaitesii (Leight.) Müll. Arg. A48 = Pyrenula papillifera (Nyl.) Aptroot
Anthracothecium toowoombense (Müll. Arg.) Aptroot A9 = Anthracothecium interlatens (Nyl.) Aptroot
Anthracothecium varians R.C. Harris A21 = Pyrenula novemseptata Vain.
Anthracothecium variolosum (Pers.) Müll. Arg. A42 = ? Pyrenula globifera (Eschw.) Aptroot
Anthracothecium vermiculare Kashiw. & Kurok. A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium vermicularis Kashiw. & Kurok. A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Anthracothecium vitellinum (Stizenb.) Müll. Arg. A27 = Pyrenula ochraceoflava (Nyl.) R.C. Harris
Anthracothecium welwitschii Upreti & Ajay Singh A49 = Pyrenula welwitschii (Upreti & Ajay Singh) Aptroot
Pyrenula acaciae Vain. B37 = Pyrenula adacta Fée
Pyrenula achariana (Fée) Vain. B79 = Pyrenula anomala (Ach.) Vain.
Pyrenula achroopora (Nyl.) Arnold B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula acutalis R.C. Harris B100, B118
Pyrenula acutispora Kalb & Hafellner B38, B141
Pyrenula adacta Fée B37
Pyrenula adacta var. cinerascens (Müll. Arg.) Zahlbr. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula addubitans (Stirt.) Zahlbr. = fungus, cf. Pleospora
Pyrenula aenea (Wallr.) Rabenh. = Porina aenea (Wallr.) Zahlbr.
Pyrenula aethiobola (Wahlenb.) Ach. = Verrucaria aethiobola Wahlenb.
Pyrenula affinis Malme B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula aggregans Vain. B60
Pyrenula aggregata (Fée) Fée B153
Pyrenula alba A. Massal. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula alba var. laevigata (Pers.) Trevis. B84 = Pyrenula laevigata (Pers.) Arnold
Pyrenula alba var. leucoplaca (Wallr.) Schaer. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula alba var. microcarpa (Hepp) Trevis. B84 = Pyrenula laevigata (Pers.) Arnold
Pyrenula albella Müll. Arg. B83 = Pyrenula microcarpa Müll. Arg.
Pyrenula albida Müll. Arg. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula albidopunctata Zahlbr. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula albissima (Fée) Trevis. = Polypyrenula albissima (A. Massal.) Aptroot
Pyrenula alboostiolata Vain. B83 = Pyrenula microcarpa Müll. Arg.
Pyrenula albothallina Vain. B82
Pyrenula alni A. Massal. B12 = ? Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula alnicola R.C. Harris B38, B141 = Pyrenula acutispora Kalb & Hafellner
Pyrenula alutacea (Wallr.) Schaer. = Verrucaria fuscella (Turner) Winch & Thornhill
Pyrenula americana (A. Massal.) Trevis. = Anisomeridium americanum (A. Massal.) R.C. Harris
Pyrenula analepta (Ach.) Trevis. (nom. illeg.) = Arthopyrenia analepta (Ach.) A. Massal.
Pyrenula analepta Fée = ? Anisomeridium americanum (A. Massal.) R.C. Harris
Pyrenula analepta var. americana Fée = ? Anisomeridium americanum (A. Massal.) R.C. Harris
Pyrenula anamalaiensis (Upreti & Ajay Singh) Upreti A35 = Pyrenula ceylonensis (Ajay Singh & Upreti) Aptroot
Pyrenula andamanica Ajay Singh & Upreti B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula andina Aptroot B121
Pyrenula annularis (Spreng.) Fée = Trypethelium annulare (Spreng.) Mont.
Pyrenula annulata Müll. Arg. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula anomala (Ach.) Vain. B79
Pyrenula anomaloides Vain. B79 = Pyrenula anomala (Ach.) Vain.
Pyrenula antoniae (Krempelh.) van Overeem-de Haas = Trypethelium cinereorosellum Krempelh.
Pyrenula apayaoensis Vain. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula approximans (Krempelh.) Müll. Arg. B103
Pyrenula approximata Vain. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula aquila R.C. Harris B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula arctecincta Fée = unknown
Pyrenula areolata Ach. = Staurothele areolata (Ach.) Lettau
Pyrenula arthonioides (A. Massal.) Trevis. = Tomasellia arthonioides (A. Massal.) A. Massal.
Pyrenula arthonioides (Eschw.) Vain. B79 = Pyrenula arthoniotheca Upreti
Pyrenula arthoniotheca Upreti B79
Pyrenula arthoniza (C. Knight) Müll. Arg. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula asahinae (Kashiw. & Kurok.) H. Harada A21 = Pyrenula subvariolosa (C. Knight) Aptroot
Pyrenula aspistea (Ach.) Ach. B145
Pyrenula aspisteoides Vain. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula astroidea (Fée) R.C. Harris A31
Pyrenula athallina H. Magn. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula atroalbella Vain. B46 = Pyrenula sexlocularis (Nyl.) Müll. Arg.
Pyrenula atrofusca Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula atrofuscescens Vain. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula atrolaminata R.C. Harris B78
Pyrenula atropurpurea (Eschw.) Müll. Arg. B144
Pyrenula aurantiaca Fée B66 = Pyrenula cerina Eschw.
Pyrenula aurantiopileata Aptroot B65
Pyrenula awajiensis (Vain.) Kashiw. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula bahiana Malme B86
Pyrenula baileyi (C. Knight) Shirley B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula balia (Krempelh.) R.C. Harris B150
Pyrenula bataanensis (Rehm) Shoemaker & P.M. LeClair B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula bayrhofferi (Zwackh) Hepp = Thelopsis rubella Nyl.
Pyrenula betulae (A. Massal.) Trevis. = Arthopyrenia grisea (Schleich. ex Schaer.) Körb.
Pyrenula bicarpa Upreti A31 = Pyrenula astroidea (Fée) R.C. Harris
Pyrenula bicuspidata Müll. Arg. B43
Pyrenula biformis (Borrer) Hepp = Anisomeridium biforme (Borrer) R.C. Harris
Pyrenula bilirana Vain. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula boberskiana Körb. = fungus, cf. Leptosphaeria
Pyrenula bonariensis Malme B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula bonplandiae (Fée) B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula borneensis Aptroot ined. A37
Pyrenula brachysperma Müll. Arg. = mixture of fungi, earlier classified in Dipyrenis
Pyrenula breutelii (Müll. Arg.) Aptroot A40
Pyrenula brunnea Fée B145
Pyrenula caerulescens Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula camptospora Malme B96 = Pyrenula cryptostoma (Nyl.) Müll. Arg.
Pyrenula canellae-albae Fée A12 = Sulcopyrenula canellae-albae (Fée) H. Harada
Pyrenula caraibica Aptroot B37 = Pyrenula adacta Fée
Pyrenula carcasana Müll. Arg. B47
Pyrenula carpinea (Wallr.) Trevis. = Porina aenea (Wallr.) Zahlbr.
Pyrenula cartilaginea Fée = Trypethelium cartilagineum (Fée) Aptroot
Pyrenula cartilaginea A. Massal. (nom. illeg.) = nomen nudum
Pyrenula caryae R.C. Harris B121, B125
Pyrenula castanea (Eschw.) Müll. Arg. B117
Pyrenula catalepta Schaer. = Verrucaria fuscella (Turner) Winch & Thornhill
Pyrenula catervaria (Fée) A. Massal. = Astrothelium variolosum (Ach.) Müll.Arg.
Pyrenula cayennensis Müll. Arg. B94
Pyrenula cerasi (Schrad.) Trevis. = Arthopyrenia cerasi (Schrad.) A. Massal.
Pyrenula cerasi var. vera Naeg. = Arthopyrenia cerasi (Schrad.) A. Massal.
Pyrenula ceratina Fée = Trypethelium ceratinum (Fée) R.C. Harris
Pyrenula cerina Eschw. B66
Pyrenula cerina f. expallens Zahlbr. B66 = Pyrenula cerina Eschw.
Pyrenula ceylonensis (Ajay Singh & Upreti) Aptroot A35
Pyrenula chilensis (Fée) R.C. Harris A34
Pyrenula chionea (Mont.) Trevis. = Polymeridium chioneum (Mont.) R.C. Harris
Pyrenula chloroplaca Shirley B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula chlorospila (Nyl.) Arnold B131
Pyrenula chlorospila var. macrospora (Degel.) Maas Geest. B129 = Pyrenula macrospora (Degel.) Coppins & P. James
Pyrenula chlorotica (Ach.) Trevis. = Porina chlorotica (Ach.) Müll. Arg.
Pyrenula chondrina Zahlbr. B122 = ? Pyrenula chondrina Zahlbr.
Pyrenula chungii Zahlbr. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula ciliata Aptroot B40
Pyrenula cinchonae Fée B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula cinchonae (Ach.) Tuck. = Arthopyrenia cinchonae (Ach.) Müll. Arg.
Pyrenula cinerea Zahlbr. B83 = Pyrenula microcarpa Müll. Arg.
Pyrenula cinerella (Flot. ex Zwackh) Branth & Rostrup = Mycomicrothelia melanospora (Hepp) D. Hawksw.
Pyrenula cinerella var. quadriloculata Fink = a coelomycete
Pyrenula cinerella ssp. quadriloculata (Fink) Fink = a coelomycete
Pyrenula cinereoglauca Zahlbr. B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula cinereovelata Vain. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula cinerosa Ach. A48 = ? Pyrenula pyrenuloides (Mont.) R.C. Harris
Pyrenula cinnamomea (Mont.) Trevis. = Astrothelium sp.
Pyrenula circumfiniens Vain. B29
Pyrenula circumfusa (Nyl.) Trevis. B13 = Blastodermia nitida A. Massal.
Pyrenula circumrubens (Nyl.) B. de Lesd. B69 = Pyrenula cruenta (Mont.) Vain.
Pyrenula circumrubens var. erythrinosa B. de Lesd. B69 = Pyrenula cruenta (Mont.) Vain.
Pyrenula circumrubens var. rubrotecta (Stirt.) Shirley B69 = Pyrenula cruenta (Mont.) Vain.
Pyrenula citriformis R.C. Harris B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula clandestina Ach. = Ocellularia clandestina (Ach.) Müll. Arg.
Pyrenula clopima (Wahlenb.) Ach. = Staurothele clopima (Wahlenb.) Th. Fr.
Pyrenula coactella (Stirt.) Upreti = Melanothecopsis coactella (Stirt.) C.W. Dodge
Pyrenula cocoes Müll. Arg. B81, B89
Pyrenula coerulescens Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula collospora Vain. B9 = Mycomicrothelia collospora (Vain.) Aptroot
Pyrenula columellata Upreti & Ajay Singh B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula comirana Vain. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula commixta Malme B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula complanata (Mont.) Trevis. B130
Pyrenula composita Ach. = a non-lichenized fungus
Pyrenula concastroma R.C. Harris B71
Pyrenula concatervans (Nyl.) R.C. Harris B46 = Pyrenula sexlocularis (Nyl.) Müll. Arg.
Pyrenula confinis (Nyl.) R.C. Harris A38
Pyrenula confoederata R.C. Harris B152
Pyrenula congregans Nyl.= nomen nudum
Pyrenula conica Müll. Arg. = Saccardoella sp. (a non-lichenized fungus)
Pyrenula consociata Zahlbr. B149 = Pyrenula deliquescens (C. Knight) Müll. Arg.
Pyrenula conspurcata Müll. Arg. B83 = Pyrenula microcarpa Müll. Arg.
Pyrenula convexa (Nyl.) Müll. Arg. B125 = Pyrenula subducta (Nyl.) Müll. Arg.
Pyrenula copalchina Fée B25 = ? Pyrenula crassiuscula (Malme) Aptroot
Pyrenula cordatula Zahlbr. B130 = Pyrenula complanata (Mont.) Trevis.
Pyrenula corticata (Müll. Arg.) R.C. Harris A38 = Pyrenula confinis (Nyl.) R.C. Harris
Pyrenula corticola R.K. Verma & Kamal = unknown
Pyrenula coryli A. Massal. B84
Pyrenula costaricensis Müll. Arg. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula crassescens (Stirt.) Müll. Arg. B134 = ? Pyrenula quassiaecola (Fée) Fée
Pyrenula crassiuscula (Malme) Aptroot B25
Pyrenula cruenta (Mont.) Vain. B69
Pyrenula cruentata (Müll. Arg.) R.C. Harris A25
Pyrenula cryptostoma (Nyl.) Müll. Arg. B96
Pyrenula cryptothelia (Müll. Arg.) Aptroot & Etayo B34
Pyrenula crystalligera H. Magn. B86 = Pyrenula bahiana Malme
Pyrenula cubana (Müll. Arg.) R.C. Harris B34
Pyrenula cuprescens Zahlbr. B58 = Pyrenula immissa (Stirt.) Zahlbr.
Pyrenula cuyabensis (Malme) R.C. Harris B30
Pyrenula cylindrica Kashiw. B50
Pyrenula cyrtospora (Stirt.) Nyl. B46 = Pyrenula sexlocularis (Nyl.) Müll. Arg.
Pyrenula darjeelingensis Jagadeesh Ram & G.P. Sinha A20
Pyrenula dealbata (C. Knight) Müll. Arg. B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula decumbens (Müll. Arg.) Upreti B18 = Lithothelium decumbens (Müll. Arg.) Aptroot
Pyrenula defossa Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula deliquescens (C. Knight) Müll. Arg. B149
Pyrenula deplanata Müll. Arg. B150 = Pyrenula balia (Krempelh.) R.C. Harris
Pyrenula deprimens (C. Knight) D.J. Galloway B131 = ? Pyrenula chlorospila (Nyl.) Arnold
Pyrenula dermatodes (Borrer) Schaer. B122
Pyrenula diffracta Müll. Arg. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula diluta (Fée) Tuck. = Pseudopyrenula diluta (Fée) Müll. Arg
Pyrenula discissa (Nyl.) Zahlbr. = Melanotrema meiospermum (Nyl.) A. Frisch
Pyrenula discolor Ach. = Ampliotrema discolor (Ach.) Kalb
Pyrenula dispersa (Wallr.) Schaer. = Verrucaria sp.
Pyrenula dispora Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula dissimulans (Müll. Arg.) R.C. Harris A51
Pyrenula duplicans (Nyl.) Aptroot A44
Pyrenula dussii Malme B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula ectypa (Krempelh.) Zahlbr. B130 = Pyrenula complanata (Mont.) Trevis.
Pyrenula elaeina (Borrer) Schaer. = Verrucaria elaeina Borrer
Pyrenula elegans Ajay Singh & Upreti B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula elliptica Müll. Arg. B29
Pyrenula emergens (Müll. Arg.) Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula emersa Malme B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula emersa var. rissoensis Malme B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula endococcinea (Nyl.) Willey = Stigmidium tabacinae (Arnold) Triebel
Pyrenula endocrocea Aptroot ined. A23
Pyrenula endoleuca Fée = unknown
Pyrenula endostega Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula endoxantha Vain. = Pseudopyrenula endoxantha (Vain.) Zahlbr.
Pyrenula enteroleuca Spreng. = Stictis urceolatum (Ach.) Gilenstam
Pyrenula epapillata Fée = Astrothelium variolosum (Ach.) Müll.Arg.
Pyrenula epidermidis (Ach.) Trevis. = Leptorhaphis epidermidis (Ach.) Th. Fr.
Pyrenula erumpens R.C. Harris A16, B30
Pyrenula eschweileri (Mont.) Trevis. = Phaeographis lobata (Eschw.) Müll. Arg.
Pyrenula eucalypta Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula euphorbiae Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula exigua Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula expectata R.C. Harris B126 = Pyrenula spectata R.C. Harris
Pyrenula falklandica (Nyl.) Zahlbr. = Lithothelium falklandicum (Nyl.) Aptroot
Pyrenula fallaciosa (Stizenb. ex Arnold) Willey = Julella fallaciosa (Stizenb. ex Arnold) R.C. Harris
Pyrenula falsaria (Zahlbr.) R.C. Harris A34 = Pyrenula schiffneri (Zahlbr.) Aptroot
Pyrenula farrea (Ach.) Branth & E. Rostrup = Verrucaria sp.
Pyrenula farrea var. pluriloculata (Fink) Zahlbr. B14 = Eopyrenula intermedia Coppins
Pyrenula farrea var. umbrosa (Körb.) Zahlbr. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula feeana Trevis. = ? Anisomeridium americanum (A. Massal.) R.C. Harris
Pyrenula feracissima Vain. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula ferax Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula fetivica (Krempelh.) Müll. Arg. B100
Pyrenula fibrata (Stirt.) Zahlbr. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula fimbriata Fée = a non-lichenized fungus
Pyrenula finitima Müll. Arg. B103
Pyrenula fissa H. Magn. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula flagellata H. Harada B48
Pyrenula flaventior Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula flavofulvescens Vain. B46 = Pyrenula sexlocularis (Nyl.) Müll. Arg.
Pyrenula fraxini (A. Massal.) Trevis. = Arthopyrenia fraxini A. Massal.
Pyrenula friesii Trevis. = nomen nudum
Pyrenula fuliginea Ach. = Thelignya fuliginea (Ach.) A. Massal.
Pyrenula fulva (Krempelh.) Müll. Arg. B98
Pyrenula fulvella R.C. Harris B27 = Pyrenula subgregantula Müll. Arg.
Pyrenula fulvescens Malme B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula funckii Spreng. = Verrucaria funckii (Spreng.) Zahlbr.
Pyrenula fusca (Krempelh.) Vain. B79 = Pyrenula anomala (Ach.) Vain.
Pyrenula fuscata Pers. = unknown, type lost
Pyrenula fuscolurida Vain. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula fuscoolivacea Vain. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula fusiformis Hepp = Porina aenea (Wallr.) Zahlbr.
Pyrenula fusispora (Malme) Aptroot B23
Pyrenula fusoluminata Aptroot B52
Pyrenula gahavisukana Aptroot B37
Pyrenula galactina (Shirley) Kantvilas B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula gaudichaldii (Fée) Pers. = Trypethelium tropicum (Ach.) Müll. Arg.
Pyrenula gaudichaudii (Fée) Pers. = Trypethelium tropicum (Ach.) Müll. Arg.
Pyrenula gelatinosa (Ach.) Schaer. = Agonimia gelatinosa (Ach.) M.A. Brand & Diederich
Pyrenula gemmata (Ach.) Naeg. = Acrocordia gemmata (Ach.) A. Massal.
Pyrenula gemmata var. macrocarpa (Körb.) Willey = Anisomeridium macrocarpum (Körb.) V. Wirth.
Pyrenula gemmata var. sphaeroides (Wallr.) Hepp = Acrocordia gemmata (Ach.) A. Massal.
Pyrenula gemmata var. tersa (Krempelh.) Kickx = Acrocordia gemmata (Ach.) A. Massal.
Pyrenula gemmifera (Tayl.) Willey = Endococcus propinquus (Körb.) D. Hawksw
Pyrenula gibberosa Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula gibberulosa (Vain.) Aptroot A20
Pyrenula gibbosa Ach. = Rimularia gibbosa (Ach.) Coppins, Hertel & Rambold.
Pyrenula gigas Zahlbr. B136
Pyrenula glabrata (Ach.) A. Massal. B84 = Pyrenula laevigata (Pers.) Arnold
Pyrenula glabrata f. cinerea Haszl. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula glabrata f. grisea Nyl. = unknown
Pyrenula glabrata f. incusa Flot. B99, B113 = Pyrenula occidentalis (R.C. Harris) R.C. Harris
Pyrenula glabrata f. major Krempelh. B84 = Pyrenula laevigata (Pers.) Arnold
Pyrenula glabrata f. microcarpa Hepp B84 = Pyrenula laevigata (Pers.) Arnold
Pyrenula glabrata f. pachyderma Haszl. B138 = Pyrenula nitida (Weigel) Ach.
Pyrenula glabratula (Nyl.) Arnold B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula glabrescens Vain. B106 = Pyrenula mastophoriza (Nyl.) Zahlbr.
Pyrenula glabriuscula (Nyl.) Vain. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula glauca (Fée) Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula glaziovii Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula globifera (Eschw.) Aptroot A42
Pyrenula globosa Pers. = unknown, type lost
Pyrenula gracilior Müll. Arg. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula gravenreuthii Stein B69 = Pyrenula cruenta (Mont.) Vain.
Pyrenula gregantula Müll. Arg. B112 = Pyrenula laetior Müll. Arg.
Pyrenula griseola Vain. B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula grossa Aptroot B98
Pyrenula guayaci (Fée) Trevis. = Parapyrenis guayaci (Fée) Aptroot
Pyrenula guimarana Vain. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula hagmannii Redinger = unknown
Pyrenula harrimannii (Sm.) Trevis. = Verrucaria hochstetteri Fr.
Pyrenula harrisii Hafellner & Kalb B99, B113 = Pyrenula occidentalis (R.C. Harris) R.C. Harris
Pyrenula hawaiiensis Aptroot ined. B25
Pyrenula henatomma (Ach.) Ach. = Ocellularia henatomma (Ach.) Müll. Arg.
Pyrenula heppii (Naeg.) Müll. Arg. = a non-lichenized fungus
Pyrenula heppii ssp. carpinea Hepp = a non-lichenized fungus
Pyrenula heppii ssp. fraxini Hepp = a non-lichenized fungus
Pyrenula herrei Fink = Arthopyrenia plumbaria (Stizenb.) R.C. Harris
Pyrenula heterochroa (Mont.) Trevis. = Trypethelium aeneum (Eschw.) Zahlbr.
Pyrenula heteroclita Ach. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula heteroclita ssp. denigrata Ach. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula heteroclita ssp. minuscula Ach. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula hiascens Ach. = Verrucaria hochstetteri Fr
Pyrenula hibernica (Nyl.) Aptroot A35
Pyrenula himalayana Upreti A49 = Pyrenula welwitschii (Upreti & Ajay Singh) Aptroot
Pyrenula hirsuta Etayo B40
Pyrenula hoehneliana Zahlbr. B54
Pyrenula homalisma (C. Knight) D.J. Galloway B134 = ? Pyrenula quassiaecola (Fée) Fée
Pyrenula howeana Aptroot B64
Pyrenula hunana Zahlbr. B94 = Pyrenula cayennensis Müll. Arg.
Pyrenula hyalospora (Nyl.) Tuck. = Lithothelium hyalosporum (Nyl.) Aptroot
Pyrenula hydrela (Ach.) Schaer. = Verrucaria denudata Zschacke
Pyrenula hypophyta (Nyl.) Müll. Arg. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula hypophytoides Harm. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula imitans (Nyl.) Zahlbr. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula immersa Müll. Arg. A17 = Pyrenula subumbilicata (C. Knight) Aptroot
Pyrenula immissa (Stirt.) Zahlbr. B58
Pyrenula imperfecta (Ellis & Everh.) R.C. Harris B108 = Pyrenula subelliptica (Tuck.) R.C. Harris
Pyrenula impressa Müll. Arg. B133 = Pyrenula punctella (Nyl.) Trevis.
Pyrenula incrustans Körb. = Phaeospora parasitica Arnold
Pyrenula indica A. Massal. = unknown
Pyrenula indusiata Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula infernalis (Mont.) Trevis. = Anisomeridium infernale (Mont.) R.C. Harris
Pyrenula infida (Nyl.) Müll. Arg. B44 = Pyrenula melanophthalma (Mont.) Trevis.
Pyrenula infracongruens Aptroot & Schumm B105
Pyrenula insularum H. Magn. B81, B89 = Pyrenula cocoes Müll. Arg.
Pyrenula interducta (Nyl.) Zahlbr. B115
Pyrenula introducta (Stirt.) Zahlbr. B130 = Pyrenula complanata (Mont.) Trevis.
Pyrenula irosina Vain. B104
Pyrenula irregularis Fée B33 = Pyrenula septicollaris (Eschw.) R.C. Harris
Pyrenula irrubescens Vain. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula isabellina Vain. (as isabellinum) = unknown
Pyrenula japonica Kurok. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula kakouettae Sérus. & Diederich B38, B141 = Pyrenula acutispora Kalb & Hafellner
Pyrenula kamatii Muthukumar & Tarar = nom. inval.
Pyrenula karnatakensis Upreti A31 = Pyrenula astroidea (Fée) R.C. Harris
Pyrenula kelungana Zahlbr. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula kermesina R.C. Harris A26
Pyrenula knightiana Müll. Arg. B131 = ? Pyrenula chlorospila (Nyl.) Arnold
Pyrenula kunthii (Fée) Fée B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula kurzii Ajay Singh & Upreti B93
Pyrenula lactea (A. Massal.) Tuck. = Julella lactea (A. Massal.) M.E. Barr
Pyrenula laetior Müll. Arg. B112
Pyrenula laevigata (Pers.) Arnold B84
Pyrenula laevigata f. cinerea (Haszl.) Zahlbr. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula laevigata f. grisea (Nyl.) Zahlbr. = unknown
Pyrenula laevigata f. incusa (Flot.) Zahlbr. B99, B113 = Pyrenula occidentalis (R.C. Harris) R.C. Harris
Pyrenula laevigata f. major (Krempelh.) Zahlbr. B84 = Pyrenula laevigata (Pers.) Arnold
Pyrenula laevigata f. microcarpa (Hepp) Arnold B84 = Pyrenula laevigata (Pers.) Arnold
Pyrenula laevigata f. pachyderma (Hazsl.) Zahlbr. B138 = Pyrenula nitida (Weigel) Ach.
Pyrenula laevigata var. meizospora Vain. B83 = ? Pyrenula microcarpa Müll. Arg.
Pyrenula laevigata var. microspora Vain. B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula lagoensis Müll. Arg. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula laii Aptroot B58 = Pyrenula immissa (Stirt.) Zahlbr.
Pyrenula lamprocarpa Müll. Arg. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula laureriformis Aptroot B33 = Pyrenula septicollaris (Eschw.) R.C. Harris
Pyrenula leightontii Muthukumar & Tarar = nom. inval.
Pyrenula leprieurii Trevis. = a Celothelium sp.
Pyrenula leucocephala (Ach.) Ach. = Schismatomma abietinum (Erhr.) A. Massal.
Pyrenula leucocephala var. amphibola Ach. = Schismatomma abietinum (Erhr.) A. Massal.
Pyrenula leucochlora (Müll. Arg.) Willey = Anisomeridium leucochlorum (Müll. Arg.) R.C. Harris
Pyrenula leucophaea (Wallr.) Körb. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula leucoplaca (Wallr.) Körb. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula leucoplaca f. chrysoleuca (Körb.) Migula B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula leucoplaca f. umbrosa (Körb.) Migula B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula leucoplaca var. chrysoleuca Körb. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula leucoplaca var. pluriloculata Fink B14 = Eopyrenula intermedia Coppins
Pyrenula leucoplaca var. umbrosa Körb. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula leucostoma Ach. A46
Pyrenula leucostoma Fée = Arthopyrenia planorbis (Ach.) Müll.Arg.
Pyrenula leucotrypa (Nyl.) Upreti B77
Pyrenula libricola Fée A46 = Pyrenula leucostoma Ach.
Pyrenula limae Vain. B117 = Pyrenula castanea (Eschw.) Müll. Arg.
Pyrenula limayensis Vain. B117 = Pyrenula castanea (Eschw.) Müll. Arg.
Pyrenula lineatostroma Aptroot B2
Pyrenula lithina (Ach.) Ach. = Staurothele sp.
Pyrenula longislandica Ajay Singh & Upreti B150 = Pyrenula balia (Krempelh.) R.C. Harris
Pyrenula lucifera R.C. Harris B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula luteopruinosa Etayo & Aptroot B63
Pyrenula lyoni (Zahlbr.) Aptroot A32
Pyrenula macounii R.C. Harris B132
Pyrenula macrocarpa A. Massal. B130 = Pyrenula complanata (Mont.) Trevis.
Pyrenula macrocarpa (Fée) A. Massal. = Astrothelium variolosum (Ach.) Müll.Arg
Pyrenula macrospora (Degel.) Coppins & P. James B129
Pyrenula macularis (Zahlbr.) R.C. Harris A40 = Pyrenula breutelii (Müll. Arg.) Aptroot
Pyrenula maculata (R.C. Harris) R.C. Harris B131 = Pyrenula chlorospila (Nyl.) Arnold
Pyrenula mamillana (Ach.) Trevis. B117
Pyrenula mamillana f. diminuens (Nyl.) Zahlbr. B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula mamillana var. bataana Vain. B117 = Pyrenula castanea (Eschw.) Müll. Arg.
Pyrenula mamillana var. erubescens Zahlbr. B131 = Pyrenula chlorospila (Nyl.) Arnold
Pyrenula mamillana var. santensis (Nyl.) Trevis. B150 = Pyrenula balia (Krempelh.) R.C. Harris
Pyrenula mamillana var. subconfluens Vain. B117 = ? Pyrenula mamillana (Ach.) Trevis.
Pyrenula mamillata (Ajay Singh) Upreti A31 = Pyrenula astroidea (Fée) R.C. Harris
Pyrenula mammillaris (Hepp) Zahlbr. B134 = ? Pyrenula quassiaecola (Fée) Fée
Pyrenula mangiferae Vain. B47 = Pyrenula carcasana Müll. Arg.
Pyrenula manhaviensis Zahlbr. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula maravalensis Vain. B92
Pyrenula marcida Fée = Trypethelium marcidum (Fée) Müll.Arg.
Pyrenula margacea (Wahlenb.) Ach. = Verrucaria margacea (Wahlenb.) Wahlenb.
Pyrenula marginata Hook. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula marginata f. diminuens Nyl. B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula marginata var. australasiatica Vain. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula marginata var. fulva Krempelh. B98 = Pyrenula fulva (Krempelh.) Müll. Arg.
Pyrenula marginata var. santensis (Nyl.) Tuck. B150 = Pyrenula balia (Krempelh.) R.C. Harris
Pyrenula marginatula Müll. Arg. B37 = Pyrenula adacta Fée
Pyrenula marmorata Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula martinicana Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula martinicana (Vain.) R.C. Harris B37 = Pyrenula adacta Fée
Pyrenula massalongiana Trevis. = Strigula stigmatella (Ach.) R.C. Harris
Pyrenula massariospora (Starb.) R.C. Harris B118
Pyrenula mastoidea Ach. = Porina mastoidea (Ach.) Müll. Arg.
Pyrenula mastophora (Nyl.) Müll. Arg. B134
Pyrenula mastophora var. australis Malme B134 = Pyrenula mastophora (Nyl.) Müll. Arg.
Pyrenula mastophoriza (Nyl.) Zahlbr. B106
Pyrenula mastophorizans Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula mastophoroides (Nyl.) Zahlbr. B115
Pyrenula mastophoroides var. flavicans (Nyl.) Zahlbr. B115 = Pyrenula mastophoroides (Nyl.) Zahlbr.
Pyrenula mastospora Vain. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula maura (Wahlenb.) Schaer. = Verrucaria maura Wahlenb.
Pyrenula media Aptroot B130
Pyrenula megalospora Fink = Acrocordia megalospora (Fink) R.C. Harris
Pyrenula megapotamica Malme B47 = Pyrenula carcasana Müll. Arg.
Pyrenula melaleuca Müll. Arg. B83 = Pyrenula microcarpa Müll. Arg.
Pyrenula melanophthalma (Mont.) Trevis. B44
Pyrenula melanospora Hepp = Mycomicrothelia melanospora (Hepp) D. Hawksw.
Pyrenula micheneri R.C. Harris B139
Pyrenula microcarpa Müll. Arg. B83
Pyrenula microcarpoides Müll. Arg. B83 = Pyrenula microcarpa Müll. Arg.
Pyrenula microciba Ach. = Catapyrenium cinereum (Pers.) Körb.
Pyrenula micromma Shirley B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula micromma var. leucomma Nyl. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula microscopica Müll. Arg. = Mycoporopsis microscopica (Müll. Arg.) Riedl
Pyrenula microspora (Nagarkar & Patw.) Upreti D30
Pyrenula microtheca R.C. Harris B38
Pyrenula minae Aptroot & Lücking B109
Pyrenula minarum Vain. B73
Pyrenula minarum var. colorans Malme B73 = Pyrenula minarum Vain.
Pyrenula minor Fée B154
Pyrenula minuta Naeg. = Strigula affinis (A. Massal.) R.C. Harris
Pyrenula minutissima Aptroot, Valadbeigi & Sipman ined. B142
Pyrenula minutula Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula mollis Fée B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula moniliformis (C. Knight) Müll. Arg. B51
Pyrenula montagnei Müll. Arg. B51
Pyrenula montana Aptroot B147
Pyrenula montocensis Lücking B58
Pyrenula mozambica Vain. B134 = ? Pyrenula quassiaecola (Fée) Fée
Pyrenula mucosa (Vain.) R.C. Harris A48 = Pyrenula papillifera (Nyl.) Aptroot
Pyrenula muscorum (A. Massal.) Hepp = Strigula stigmatella (Ach.) R.C. Harris
Pyrenula muscorum var. faginea (Schaer.) Hepp = Strigula stigmatella (Ach.) R.C. Harris
Pyrenula myriocarpa Fée = Mycomicrothelia wallrothii (Hepp) D. Hawksw.
Pyrenula naegelii Hepp = Strigula glabra (A. Massal.) V. Wirth
Pyrenula nanospora (Ajay Singh) Upreti A39
Pyrenula nebulosa Zahlbr. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula neglecta R.C. Harris B89 = Pyrenula pseudobufonia (Rehm) R.C. Harris
Pyrenula neglecta ssp. occidentalis R.C. Harris B99, B113 = Pyrenula occidentalis (R.C. Harris) R.C. Harris
Pyrenula neoculata Aptroot A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Pyrenula neofulva Ajay Singh B60 = Pyrenula papilligera (Leight.) Müll. Arg.
Pyrenula neojaponica H. Harada A8 = Anthracothecium macrosporum (Hepp) Müll. Arg.
Pyrenula neolaevigata H. Harada A19 = Lithothelium nanosporum (C. Knight) Aptroot
Pyrenula neopeltophora Ajay Singh B106 = Pyrenula mastophoriza (Nyl.) Zahlbr.
Pyrenula neosandwicensis Aptroot A44
Pyrenula netrospora Naeg. = Strigula affinis (A. Massal.) R.C. Harris
Pyrenula nigrescens (Pers.) Ach. = Verrucaria nigrescens Pers.
Pyrenula nigrescens var. areolata Schaer. = Verrucaria nigrescens Pers.
Pyrenula nigrocincta Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula nitens (Fée) Fée = unknown
Pyrenula nitens f. pinguis (Chevall.) Zahlbr. B131 = Pyrenula chlorospila (Nyl.) Arnold
Pyrenula nitida (Weigel) Ach. B138
Pyrenula nitida f. chevallieri M. Choisy B138 = Pyrenula nitida (Weigel) Ach.
Pyrenula nitida var. commutata Trevis. B150 = Pyrenula balia (Krempelh.) R.C. Harris
Pyrenula nitida f. crassa Barchalov B138 = Pyrenula nitida (Weigel) Ach.
Pyrenula nitida f. diffracta Erichsen B138 = Pyrenula nitida (Weigel) Ach.
Pyrenula nitida f. elaeodes (Leight.) A.L. Sm. B129 = Pyrenula macrospora (Degel.) Coppins & P. James
Pyrenula nitida f. flavescens (Malbr.) Zahlbr. B138 = Pyrenula nitida (Weigel) Ach.
Pyrenula nitida f. pachyderma (Hazsl.) Szatala B138 = Pyrenula nitida (Weigel) Ach.
Pyrenula nitida f. pinguis (Westr.) Zahlbr. B131 = Pyrenula chlorospila (Nyl.) Arnold
Pyrenula nitida f. squamata (Malbr.) Zahlbr. B138 = Pyrenula nitida (Weigel) Ach.
Pyrenula nitida f. virens Servít & Nádvorník B138 = Pyrenula nitida (Weigel) Ach.
Pyrenula nitida var. aequata Zahlbr. B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitida var. americana Fée B4 = Architrypethelium nitens (Fée) Aptroot
Pyrenula nitida var. aspistea (Ach.) Trevis. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula nitida var. dermatodes (Borrer) Trevis. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula nitida var. grandispora Barchalov B129 = Pyrenula macrospora (Degel.) Coppins & P. James
Pyrenula nitida var. macrospora Degel. B129 = Pyrenula macrospora (Degel.) Coppins & P. James
Pyrenula nitida var. major (Schaer.) Schaer. B138 = Pyrenula nitida (Weigel) Ach.
Pyrenula nitida var. minima Hepp B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitida var. minima f. pinicola Hepp B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitida var. minor Hepp B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitida var. minor f. pinicola Hepp B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitida var. nitidella (Flörke ex Schaer.) Schaer. B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitida var. nitidella f. chlorospila (Nyl.) Keissl. B131 = Pyrenula chlorospila (Nyl.) Arnold
Pyrenula nitida var. nitidella f. fuscata Suza B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitida var. nitidella f. nigrescens (B. de Lesd.) Keissl. B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitida var. phaeospila (Nyl.) Zahlbr. B131 = Pyrenula chlorospila (Nyl.) Arnold
Pyrenula nitidans Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg. B138
Pyrenula nitidella f. pinicola (Hepp) Zahlbr. B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitidella var. chlorospila (Nyl.) Szatala B131 = Pyrenula chlorospila (Nyl.) Arnold
Pyrenula nitidella var. cintrana Welw. B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitidella var. maculata R.C. Harris B131 = Pyrenula chlorospila (Nyl.) Arnold
Pyrenula nitidella var. exstantior Vain. B122 = ? Pyrenula dermatodes (Borrer) Schaer.
Pyrenula nitidella var. nigrescens B. de Lesd. B138 = Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.
Pyrenula nitidula (Bres.) R.C. Harris B106
Pyrenula nodulata (Stirt.) Zahlbr. B103 = Pyrenula approximans (Krempelh.) Müll. Arg.
Pyrenula nova-granadensis Upreti & Ajay Singh A31 = Pyrenula ravenelii (Tuck.) R.C. Harris
Pyrenula novemseptata Vain. A21
Pyrenula nuda Ajay Singh & Upreti B153 = Pyrenula aggregata (Fée) Fée
Pyrenula oblonga Zahlbr. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula oblongata (Müll. Arg.) Willey = Mycomicrothelia wallrothii (Hepp) D. Hawksw.
Pyrenula obovata (Stirt.) Shirley = Mycomicrothelia obovata (Stirt.) D. Hawksw.
Pyrenula obscura Räsänen B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula obscurascens Vain. B79 = Pyrenula anomala (Ach.) Vain.
Pyrenula obscurata Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula obscurior Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula obtecta G. Merr. = nomen nudum
Pyrenula obtusior (Nyl.) Zahlbr. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula obvoluta (Nyl.) R.C. Harris & Aptroot B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula occidentalis (R.C. Harris) R.C. Harris B99, B113
Pyrenula occulta (C. Knight) Müll. Arg. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula occulta var. leucomma (Nyl.) Zahlbr. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula ocellata (Ach.) Ach., Verrucaria ocellata Ach. = type lost
Pyrenula ocellata (Leight.) Zahlbr. (nom. illeg.), Verrucaria ocellata Leight. (nom. illeg.) = all illeg.
Pyrenula ochracea Szatala B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula ochraceoflava (Nyl.) R.C. Harris A27
Pyrenula ochraceoflava var. pacifica P.M. McCarthy A27
Pyrenula ochraceoflavens (Nyl.) R.C. Harris A27
Pyrenula oculata Ajay Singh & Upreti B111
Pyrenula oculifera Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula oleagina Fée = unknown
Pyrenula oleosa R.C. Harris A51
Pyrenula oligocarpa Malme B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula olivacea (Pers.) Hepp = Porina borreri (Trevis.) D. Hawksw. & P. James
Pyrenula olivacea (Fr.) Schaer. = Thelidium olivaceum (Fr.) Körb.
Pyrenula olivaceofusca Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula olivaceofusca C. Knight B131 = ? Pyrenula chlorospila (Nyl.) Arnold
Pyrenula orofensis Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula oxyspora (Nyl.) Körb. = Leptorhaphis epidermidis (Ach.) Th. Fr.
Pyrenula oxyspora Müll. Arg. B103 = Pyrenula finitima Müll. Arg.
Pyrenula oxysporiza Zahlbr. B103 = Pyrenula finitima Müll. Arg.
Pyrenula pachycheila Tuck. A46 = Pyrenula leucostoma Ach.
Pyrenula pachyspora Vain. B118 = Pyrenula massariospora (Starb.) R.C. Harris
Pyrenula pallidofulvescens Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula pallidofulvescens var. fulvostraminea Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula palmarum (Krempelh.) R.C. Harris A26
Pyrenula papillifera (Nyl.) Aptroot A48
Pyrenula papilligera (Leight.) Müll. Arg. B60
Pyrenula papularis (Fr.) Schaer. = Thelidium papulare (Fr.) Arnold
Pyrenula papuliformis Eckf. = Arthopyrenia papuliformis (Eckf.) Zahlbr.
Pyrenula paraensis Müll. Arg. B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula parva Vain. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula parvinuclea (Meyen & Flot.) Aptroot A39
Pyrenula parvula Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula peltophora Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula perpusilla (Nyl.) Willey = Endococcus perpusillus Nyl.
Pyrenula personata (Malme) R.C. Harris B27 = Pyrenula subgregantula Müll. Arg.
Pyrenula pertusarina Zahlbr. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula pertusarioidea Krempelh. = ? Polymeridium proponens (Nyl.) R.C. Harris
Pyrenula philippina Vain. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula philippina var. oceanica Räsänen ex Sbarbaro B117 = ? Pyrenula mamillana (Ach.) Trevis.
Pyrenula pileata Vain. B150 = Pyrenula scutata (Stirt.) Zahlbr.
Pyrenula pinguis Fée B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula pinguis Chevall. B131 = Pyrenula chlorospila (Nyl.) Arnold
Pyrenula pinguis var. emergens (Müll. Arg.) Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula pinguis var. exstans (Nyl.) Zahlbr. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula planiuscula (Nyl.) Tuck. = Anisomeridium planiusculum (Nyl.) R.C. Harris
Pyrenula planorbis (Ach.) Trevis. = Arthopyrenia planorbis (Ach.) Müll.Arg.
Pyrenula platysporella Zahlbr. B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula platystoma (Müll. Arg.) Aptroot A43
Pyrenula pleiomera (Nyl.) Zahlbr. B22
Pyrenula pleiomeriza (Nyl.) Zahlbr. = Polymeridium sp.
Pyrenula plittii R.C. Harris B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula plumbea Malme B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula pluricarpa (Ajay Singh) Upreti A31 = Pyrenula astroidea (Fée) R.C. Harris
Pyrenula polillensis Vain. B33 = Pyrenula septicollaris (Eschw.) R.C. Harris
Pyrenula polycarpa (Körb.) Hepp = Anisomeridium biforme (Borrer) R.C. Harris
Pyrenula porinella Vain. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula porinoides Ach. B145 = ? Pyrenula aspistea (Ach.) Ach.
Pyrenula porinoides Fée B122 = ? Pyrenula dermatodes (Borrer) Schaer.
Pyrenula porphyria G. Mey. = Staurothele sp.
Pyrenula porphyria Schaer. = Staurothele sp.
Pyrenula porrecta (Krempelh.) Müll. Arg. = unknown
Pyrenula praelucida (Mont.) Trevis. B56
Pyrenula prorecta (Krempelh.) Müll. Arg. = unknown
Pyrenula prostans (Mont.) Trevis. = Arthopyrenia cinchonae (Ach.) Müll. Arg.
Pyrenula prostrata (Stirt.) D.J. Galloway A31 = Pyrenula ravenelii (Tuck.) R.C. Harris
Pyrenula pseudobufonia (Rehm) R.C. Harris B89
Pyrenula pseudonitidella (C. Knight) D.J. Galloway B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula pseudovelata M. Choisy B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula psoriformis Zahlbr. B147
Pyrenula pudica Zahlbr. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula pulchella Müll. Arg. B37 = ? Pyrenula adacta Fée
Pyrenula pulchella var. cinerascens Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula punctella (Nyl.) Trevis. B133
Pyrenula punctella var. adacta (Fée) Müll. Arg. B37 = Pyrenula adacta Fée
Pyrenula punctella var. emergens Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula punctella var. exstans (Nyl.) Müll. Arg. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula punctifera Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula punctiformis (A. Massal.) Trevis. = Arthopyrenia punctiformis A. Massal.
Pyrenula punctiformis f. lactea (Anzi) Rabenh. = Strigula stigmatella (Ach.) R.C. Harris
Pyrenula punctiformis var. analepta (Ach.) Naeg. = Arthopyrenia analepta (Ach.) A. Massal.
Pyrenula punctiformis var. analepta f. coryli Hepp = Arthopyrenia analepta (Ach.) A. Massal.
Pyrenula punctiformis var. atomaria (Ach.) Hepp = Leptorhaphis atomaria (Ach.) Szatala
Pyrenula punctiformis var. cinereopruinosa (Schaer.) Hepp = Arthopyrenia cinereopruinosa (Schaer.) A. Massal.
Pyrenula punctiformis var. cinereopruinosa f. buxicola Hepp (nom. inval.) = Arthopyrenia cinereopruinosa (Schaer.) A. Massal.
Pyrenula punctiformis var. cinereopruinosa f. galactites Hepp = Arthopyrenia cinereopruinosa (Schaer.) A. Massal.
Pyrenula punctiformis var. cinereopruinosa f. hederae Hepp = Arthopyrenia cinereopruinosa (Schaer.) A. Massal.
Pyrenula punctiformis var. cinereopruinosa f. lactea Hepp = Arthopyrenia cinereopruinosa (Schaer.) A. Massal.
Pyrenula punctiformis var. cinereopruinosa f. pinicola Hepp = Arthopyrenia cinereopruinosa (Schaer.) A. Massal.
Pyrenula punctiformis var. fallax (Nyl.) Willey = Arthopyrenia analepta (Ach.) A. Massal.
Pyrenula punctiformis var. fallax f. betulae Hepp = Arthopyrenia analepta (Ach.) A. Massal.
Pyrenula punctiformis var. vera f. acerina Hepp. = Arthopyrenia analepta (Ach.) A. Massal.
Pyrenula punctiformis var. vera f. fraxini Hepp = Arthopyrenia fraxini A. Massal.
Pyrenula punctiformis ssp. lactea (Anzi) Hepp = Strigula stigmatella (Ach.) R.C. Harris
Pyrenula pupula Ach. = Trypethelium pupula (Ach.) R.C. Harris
Pyrenula pygmaea (Körb.) Tuck. = Muellerella pygmaea (Körb.) D. Hawksw.
Pyrenula pyrenastroides (C. Knight) D.J. Galloway A31 = Pyrenula ravenelii (Tuck.) R.C. Harris
Pyrenula pyrenastrospora Aptroot B72
Pyrenula pyrenuloides (Mont.) R.C. Harris A48
Pyrenula pyrgillospora Aptroot B140
Pyrenula quadruplans Vain. B57 = ? Pyrenula subpraelucida Müll. Arg.
Pyrenula quarzitica Aptroot B127
Pyrenula quassiaecola (Fée) Fée B134
Pyrenula quassiicola (Fée) Fée B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula quercus (A. Massal.) Trevis. = Cyrtidula quercus (A. Massal.) Minks
Pyrenula quercus A. Massal. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula quercuum Zahlbr. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula quinqueseptata (Hepp) Tuck. = Thelidium quinqueseptatum (Hepp) Arnold
Pyrenula quinqueseptata Aptroot B47 = Pyrenula sexluminata Aptroot
Pyrenula ravenelii (Tuck.) R.C. Harris A31
Pyrenula rhombospora Müll. Arg. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula rhyponta (Ach.) Trevis. = Arthopyrenia rhyponta (Ach.) A. Massal.
Pyrenula rimicola Müll. Arg. = Phaeospora rimosicola (Leight. ex Mudd) Hepp ex Stein
Pyrenula rinodinospora Aptroot ined. B93
Pyrenula rizalensis Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula rockii Zahlbr. B57
Pyrenula rubidopunctata Szatala B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula rubroanomala Aptroot & Lücking B68
Pyrenula rubrojavanica Aptroot ined. B69
Pyrenula rubromaculata Vain. B69 = Pyrenula cruenta (Mont.) Vain.
Pyrenula rubrostoma R.C. Harris B68, B151
Pyrenula rudis Fée = unknown, type lost
Pyrenula rugulosa Müll. Arg. B81, B89 = Pyrenula cocoes Müll. Arg.
Pyrenula salicis (A. Massal.) Trevis. = Arthopyrenia salicis A. Massal.
Pyrenula samarana Vain. B100 = ? Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula sandwicensis Zahlbr. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula santensis (Nyl.) Müll. Arg. B150 = Pyrenula balia (Krempelh.) R.C. Harris
Pyrenula schaereri A. Massal. B12 = Eopyrenula leucoplaca (Wallr.) R.C. Harris
Pyrenula schiffneri (Zahlbr.) Aptroot A34
Pyrenula schutschensis Zahlbr. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula scutata (Stirt.) Zahlbr. B150
Pyrenula segregata (Nyl.) Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula septicollaris (Eschw.) R.C. Harris B33
Pyrenula seriata (Hepp) Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula sessilis H. Magn. B134 = ? Pyrenula mastophora (Nyl.) Müll. Arg
Pyrenula sexlocularis (Nyl.) Müll. Arg. B46
Pyrenula sexlocularis var. xanthoplaca Zahlbr. B46 = Pyrenula sexlocularis (Nyl.) Müll. Arg.
Pyrenula sexluminata Aptroot B47
Pyrenula shirabeicola Kurok. & S. Nakan. B89 = Pyrenula pseudobufonia (Rehm) R.C. Harris
Pyrenula shirleyana (Müll. Arg.) Aptroot B145 = Pyrenula brunnea Fée
Pyrenula sipmanii Aptroot & K.H. Moon B65
Pyrenula spectata R.C. Harris B126
Pyrenula spadicea (Wallr.) Schaer. = Staurothele sp.
Pyrenula sphaerica Pers. = unknown, type lost
Pyrenula sphaeroides (Wallr.) Hepp = Acrocordia gemmata (Ach.) A. Massal.
Pyrenula staurospora Tuck. A12 = Sulcopyrenula staurospora (Tuck.) H. Harada
Pyrenula stictica Link = Arthonia cinereopruinosa (Körb.) Schaer.
Pyrenula stramineoatra Vain. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula straminescens Zahlbr. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula subacutalis Upreti B92 = Pyrenula maravalensis Vain.
Pyrenula subaggregata Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula subandamanica Upreti B30 = Pyrenula microspora (Nagarkar & Patw.) Upreti
Pyrenula subaperta Ach. = arthonialean pycnidia
Pyrenula subcamptospora Upreti B96
Pyrenula subcinerea (Nyl.) Tuck. = Polymeridium subcinereum (Nyl.) R.C. Harris
Pyrenula subconfluens (Vain.) Vain. B117 = ? Pyrenula mamillana (Ach.) Trevis.
Pyrenula subcongruens Müll. Arg. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula subcremea Malme B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula subcuprea Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula subcutanea Fée A46 = Pyrenula leucostoma Ach.
Pyrenula subcylindrica Jagadeesh Ram & Upreti B50
Pyrenula subdecolor (Nyl.) R.C. Harris B69 = Pyrenula cruenta (Mont.) Vain.
Pyrenula subdissidens Vain. B79 = ? Pyrenula arthoniotheca Upreti
Pyrenula subducta (Nyl.) Müll. Arg. B125
Pyrenula subducta var. retracta (Nyl.) Zahlbr. B125 = Pyrenula subducta (Nyl.) Müll. Arg.
Pyrenula subelliptica (Tuck.) R.C. Harris B108
Pyrenula subfarinosa Fée = a non-lichenized fungus
Pyrenula subferruginea (Malme) R.C. Harris B29 = Pyrenula circumfiniens Vain.
Pyrenula subglabrata (Nyl.) Müll. Arg. B113
Pyrenula subglabriuscula Vain. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula subglabriuscula var. natalensis Vain. B117 = ? Pyrenula mamillana (Ach.) Trevis.
Pyrenula subgregantula Müll. Arg. B27
Pyrenula subgriseola Vain. B145 = ? Pyrenula brunnea Fée
Pyrenula subimmersa Müll. Arg. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula subindica Upreti B2 = Pyrenula lineatostroma Aptroot
Pyrenula sublaevigata (Patw. & Makhija) Upreti A45
Pyrenula sublateritia Zahlbr. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula submarginata Vain. B117 = Pyrenula castanea (Eschw.) Müll. Arg.
Pyrenula submastophora Ajay Singh & Upreti B141
Pyrenula submersa (Borrer) Schaer. = Verrucaria denudata Zschacke
Pyrenula subnitida Müll. Arg. B134 = ? Pyrenula quassiaecola (Fée) Fée
Pyrenula subnitidella (Nyl.) Müll. Arg. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula subochraceoflavens Upreti A20 = Pyrenula gibberulosa (Vain.) Aptroot
Pyrenula suboligocarpa Upreti A35 = Pyrenula ceylonensis (Ajay Singh & Upreti) Aptroot
Pyrenula subpraelucida Müll. Arg. B57
Pyrenula subprostans (Nyl.) Tuck. = Anisomeridium subprostans (Nyl.) R.C. Harris
Pyrenula subpunctella (Nyl.) Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula subpunctiformis (Nyl.) Eckf. = Arthopyrenia atomarioides Müll. Arg.
Pyrenula subrizalensis Ajay Singh & Upreti B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula subsimplex Vain. B106 = Pyrenula nitidula (Bres.) R.C. Harris
Pyrenula subsoluta (Müll. Arg.) Aptroot B77
Pyrenula subtrahens (Nyl.) Müll. Arg. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula subtrahens var. microspora (Krempelh.) Zahlbr. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula subumbilicata (C. Knight) Aptroot A17
Pyrenula subvariolosa (C. Knight) Aptroot A21
Pyrenula subvelata Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula sulphurea (Mont.) Zahlbr. = arthonialean crust cf. Herpothallon
Pyrenula supracongruens Aptroot & Schumm B98 = Pyrenula fulva (Krempelh.) Müll. Arg.
Pyrenula tenella Müll. Arg. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula tenuisepta R.C. Harris B8
Pyrenula tessellata (Turner) Ach. = Verrucaria viridula (Schrad.) Ach.
Pyrenula tetracerae Ach. = Porina tetracerae (Ach.) Müll. Arg.
Pyrenula texana Tuck. ex R.C. Harris B83 = Pyrenula microcarpa Müll. Arg.
Pyrenula thailandica Aptroot ined. B86
Pyrenula thelemorpha Tuck. A50 = Pyrenula thelomorpha Tuck.
Pyrenula thelena (Ach.) Trevis. = Mycomicrothelia thelena (Ach.) D. Hawksw.
Pyrenula thelomorpha Tuck. A50
Pyrenula tokyensis (Müll. Arg.) H. Harada B23
Pyrenula transparens Zahlbr. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula tremulae (Körb.) Hepp = Leptorhaphis tremulae Körb.
Pyrenula tricolor Müll. Arg. B102 = ? Pyrenula vernicosa (Krempelh.) Müll. Arg.
Pyrenula triphracta (Nyl.) Willey = Phaeospora parasitica (Lönnr.) Arnold
Pyrenula tristissima Vain. B154
Pyrenula trombetana Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula tropica (Ach.) Trevis. = Trypethelium tropicum (Ach.) Müll. Arg.
Pyrenula truncata Müll. Arg. B100 = Pyrenula fetivica (Krempelh.) Müll. Arg.
Pyrenula trypanea Ach. = Ocellularia terebrata (Ach.) Müll. Arg.
Pyrenula tunicata Zahlbr. B122 = Pyrenula dermatodes (Borrer) Schaer.
Pyrenula uberina (Fée) Fée B4 = Architrypethelium uberinum (Fée) Aptroot
Pyrenula umbilicatula Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula umbonata Ach. = Pyrenocarpon thelostomum (Ach. ex J. Harriman) Coppins & Aptroot
Pyrenula umbrata Ach. = sterile crust
Pyrenula vanoverberghii Vain. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula variolosa Pers. A42 = ? Pyrenula globifera (Eschw.) Aptroot
Pyrenula velata Müll. Arg. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula velatior Müll. Arg. B145 = Pyrenula aspistea (Ach.) Ach.
Pyrenula ventosicola (Mudd) Willey = Muellerella ventosicola (Mudd) D. Hawksw.
Pyrenula vermicellifera (Kunze) Link = Opegrapha vermicellifera (Kunze) J.R. Laundon
Pyrenula vermicularis (Kashiw. & Kurok.) H. Harada A43 = Pyrenula platystoma (Müll. Arg.) Aptroot
Pyrenula vernicosa (Krempelh.) Müll. Arg. B102
Pyrenula verrucarioides Fée (nom. illeg.) = an Anisomeridium
Pyrenula verrucosa Ach. = Polyblastia verrucosa (Ach.) Lönnr.
Pyrenula verruculosa Upreti & Ajay Singh A34 = Pyrenula schiffneri (Zahlbr.) Aptroot
Pyrenula virens Müll. Arg. B153 = Pyrenula aggregata (Fée) Fée
Pyrenula virescens Müll. Arg. B134 = Pyrenula quassiaecola (Fée) Fée
Pyrenula viridescens Fée B122 = ? Pyrenula dermatodes (Borrer) Schaer.
Pyrenula vitrea (Eschw.) Müll. Arg. B134 = ? Pyrenula quassiaecola (Fée) Fée
Pyrenula volvarioides Fée = Stictis sp.
Pyrenula wallrothii Hepp = Mycomicrothelia wallrothii (Hepp) D. Hawksw.
Pyrenula warmingii (Krempelh.) Müll. Arg. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula welwitschii (Upreti & Ajay Singh) Aptroot A49
Pyrenula wetmorei R.C. Harris B31
Pyrenula wheeleri R.C. Harris B35
Pyrenula willeyana Müll. Arg. = Mycomicrothelia willeyana (Müll. Arg.) D. Hawksw
Pyrenula wilmsiana Müll. Arg. B46
Pyrenula wrightii (Müll. Arg.) R.C. Harris B76
Pyrenula xanthominuta Aptroot B122
Pyrenula xyloides (Eschw.) Müll. Arg. B117 = Pyrenula mamillana (Ach.) Trevis.
Pyrenula zeylanica Upreti & Ajay Singh B76
Pyrenula zwackhii Hepp = Mycoporum antecellans (Nyl.) R.C. Harris
Disposition of the excluded taxa in genera which are synonymous to Pyrenula
Melanotheca acervulans Nyl. = Zwackhiomyces dispersus (J. Lahm ex Körb.) Triebel & Grube
Melanotheca aciculifera Nyl. = Celothelium aciculiferum (Nyl.) Vain.
Melanotheca agminella Nyl. = Celothelium sp.
Melanotheca apogyra Nyl. = Anthostomaria apogyra (Nyl.) Theiss. & Syd.
Melanotheca arthoniella Nyl. = Dermatina arthoniella (Nyl.) Riedl
Melanotheca arthonioides (A. Massal.) Nyl. = Tomasellia arthonioides (A. Massal.) A. Massal.
Melanotheca brasiliensis Nyl. = Tomasellia brasiliensium (Nyl.) Zahlbr.
Melanotheca coactella Stirt. = Melanothecopsis coactella (Stirt.) C.W. Dodge
Melanotheca coarctella Stirt. = Melanothecopsis coactella (Stirt.) C.W. Dodge
Melanotheca collospora (Vain.) Zahlbr. = Mycomicrothelia collospora (Vain.) Aptroot (keyed out though)
Melanotheca cumingiana (Stirt.) Müll. Arg. = Laurera cumingii (Mont.) Zahlbr.
Melanotheca diffusa Leight. = Tomasellia diffusa (Leight.) J. Lahm
Melanotheca esenbeckiana Fée = Arthopyrenia esenbeckiana (Fée) R.C. Harris
Melanotheca gelatinosa (Chev.) Nyl. = Tomasellia gelatinosa (Chevall.) Zahlbr.
Melanotheca glomerulosa Arnold = alpine calcicolous fungus
Melanotheca homostegia (Nyl.) H. Olivier = Homostegia piggotii (Berk. & Broome) P. Karst.
Melanotheca insularis Hult. = Melanopsamma lettauiana (Keissl.) Vouaux
Melanotheca ischnobela Nyl. = Celothelium ischnobelum (Nyl.) M.B. Aguirre
Melanotheca leightonii (Hepp) Garov. = Tomasellia gelatinosa (Chevall.) Zahlbr.
Melanotheca macularis (Hampe ex A. Massal.) Th. Fr. = Mycomicrothelia macularis (Hampe ex A. Massal) Keissl.
Melanotheca rhaphidiza Stirt. = Celothelium aciculiferum (Nyl.) Vain.
Melanotheca simplicella Nyl. = Celothelium sp.
Melanotheca sinensis Krempelh. = Melanothecopsis sinensis (Krempelh.) C.W. Dodge
Melanotheca subpuncta Nyl. = Melanopsamma lettauiana (Keissl.) Vouaux
Melanotheca superveniens Nyl. = Perigrapha superveniens (Nyl.) Hafellner
Parathelium oblongulum Stizenb. = Anisomeridium sp.
Parathelium polysemum Nyl. = Lithothelium polysemum (Nyl.) Aptroot
Parathelium trichosporum Stizenb. = Celothelium sp.
Pyrenastrum album ssp. verrucarioides Eschw. = Polymeridium catapastum (Nyl.) R.C. Harris
Pyrenastrum cinnamomeum Eschw. = Astrothelium cinnamomeum (Eschw.) Müll. Arg.
Pyrenastrum clandestinum (Fée) Müll. Arg. = Astrothelium crassum (Fée) Aptroot
Pyrenastrum eustomum Mont. = Astrothelium eustomum (Mont.) Müll. Arg.
Pyrenastrum gallicum Spreng. = a non-lichenized fungus
Pyrenastrum lageniferum (Ach.) Müll. Arg = description refers to a mixture of 2 taxa
Pyrenastrum plicatum Eschw. (orth. variant) = Astrothelium pyrenastrosulphureum nom. nov. ined.
Pyrenastrum sulphureum Eschw. = Astrothelium pyrenastrosulphureum nom. nov. ined.
Pyrenastrum sulphureum ssp. plicatum Eschw. = Astrothelium pyrenastrosulphureum nom. nov. ined.
Not recently studied, of unknown identity:
Melanotheca coccorum (A. Massal.) Zahlbr., Micromma coccorum A. Massal.
Melanotheca crotonoidea Hue
Melanotheca oculea Stizenb.
Parathelium brodiei Stirt.
Parathelium nidificans Stirt.
Pleurotheliopsis salvata (Müll. Arg.) Zahlbr., Pleurotheliums salvatum Müll. Arg.
Pyrenastrum chilense Mont., Verrucaria chilensis (Mont.) Nyl.
Pyrenastrum compositum Hampe, Bottaria composita (Hampe) A. Massal.
Pyrenastrum echinatum Eschw.
Pyrenastrum macrospermum Mont.
Pyrenastrum trypethelioides Eschw.