nova hedwigia 98 1 2 0001 0029 aptroot wm

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© 2013 J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, www.borntraeger-cramer.deGermany. DOI: 10.1127/0029-5035/2013/0143 0029-5035/2013/0143 $ 7.25

Nova Hedwigia Vol. 98 (2014) Issue 1–2, 1–29 published online October 15, 2013; published in print February 2014

ArticleC

A refined species concept in the tropical lichen genus Polymeridium (Trypetheliaceae) doubles the number of known species, with a worldwide key to the species

André Aptroot1 and Marcela Eugenia da Silva Cáceres2

1 ABL Herbarium, G.v.d.Veenstraat 107, NL-3762 XK Soest, The Netherlands2 Departamento de Biociências, Universidade Federal de Sergipe, CEP: 49500-000, Itabaiana, Sergipe, Brazil

With 5 figures

Abstract: A refined species concept is applied in the tropical corticolous lichen genus Polymeridium (Trypetheliaceae). While up to now many species were considered to be variable in phenotypical characters, such as hamathecium inspersion and/or presence of lichexanthone, these characters are here accepted as species-specific characters. This classification is in line with the taxonomy in other groups of lichenized fungi. The monotypic genus Exiliseptum is included here in Polymeridium, as it differs in only one character from several species of Polymeridium and has no unique characters. The number of accepted species is thus raised from 27 to 53, including 13 new species, viz.: Polymeridium albidoreagens, P. albidovarians, P. alboflavescens, P. catapastoides, P. costaricense, P. inspersum, P. multiforme, P. multiseptatum, P. neuwirthii, P. pyrenastroides, P. submuriforme, P. tribulationis, and P. xanthoreagens. The following new combinations are proposed: Polymeridium albopruinosum, P. amylosporum, P. bengoanum, P. brachysporum, P. jordanii, P. microsporum, P. ocellatum, P. refertum, P. siamense, P. stramineoatrum, P. subvirescens, and P. suffusum. An identification key is provided to all accepted species.

Key words: Australia, Brazil, corticolous lichens, Costa Rica, Guyana, Indonesia, new species, Philippines, Trypetheliales, Venezuela.

Introduction

The genus Polymeridium is a conspicuous member of corticolous microlichens in Trypetheliaceae, characterized by black perithecioid ascomata, anastomosing interthecial filaments and association with trentepohlioid algae. Within the family, it is characterized by only slightly rounded ascospore lumina and a thin, whitish thallus. It is often confused with species of Pseudopyrenula, which share the family-specific characters and have similar whitish thalli, but differ in having ascospore lumina that become diamond-shaped during maturation. Harris (1993) recognized 19 species in his revision of the genus Polymeridium. He applied a broad species concept, accepting

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variation in chemistry and hamathecial inspersion within species. Since that time, this species concept has been widely adopted, and subsequently only seven species have been added to the genus (Aptroot et al. 1995, Aptroot & Ferraro 2001, Aptroot et al. 2013b).

Species of Polymeridium are generally rare or at least rarely reported, and not locally abundant, but it was already noted by Harris (1993) that at least some regions of Brazil are rich in species. Our observations confirm this and we can add that Polymeridium is even abundant on trees in at least some areas with Caatinga vegetation in Brazil.

Harris (1993) applied a rather broad species concept, and the characters absence/presence of lichexanthone and absence/presence of hamathecial inspersion were not used as distinguishing characters for species. However, we came to realize that within a certain geographic area generally only one combination of these characters is present within each morphological species, showing that these characters are not random variation. Therefore, with the additional information of numerous (over 200) collections now available, the taxonomy of the species in the genus is revised here.

Material and methods

Identification and descriptive work was carried out in Itabaiana, Universidade Federal de Sergipe, using a Leica EZ4 stereomicroscope and a Leica DM500 compound microscope, and also in Soest using an Olympus SZX7 stereomicroscope and an Olympus BX50 compound microscope with interference contrast, connected to a Nikon Coolpix digital camera. Sections have been mounted in tap water, in which also all measurements were taken, but the reaction of ascospores to IKI (Lugol’s solution) has also been observed. The specimens from this study are preserved in ISE. The chemistry of the type specimens has been investigated by spot reactions (KOH 10%), under long-wave UV light and by thin-layer chromatography (TLC) using solvent A (Orange et al. 2001).

Results and discussion

The presence or absence of lichexanthone in a lichen thallus is usually a constant character, and it is considered characteristic for species or even species groups in foliose and fruticose lichens (Kalb 1987, Sipman et al. 2009). Interestingly, the substance occurs only in predominantly tropical lichen groups and is largely absent in lichens in temperate to arctic regions. Lichexanthone is notably common in Hypotrachyna (Sipman et al. 2009) and Pyxine (Kalb 1987). It is also considered a valid species character in numerous families of crustose lichens, and also in other genera of Trypetheliaceae. Recent revisions of speciose microlichen groups, such as Graphis (Lücking 2009, Lücking et al. 2009), thelotremoid Graphidaceae (Rivas Plata et al. 2010), and Pyrenula (Aptroot 2012) invariably accept the presence or absence of lichexanthone as a species-specific character. This is supported by phylogenetic studies in these groups (Rivas Plata et al. 2012, 2013). Also in other groups, such as Arthoniales, lichexanthone is regarded to be a valuable species-specific character. In a recent study on Ochrolechia (Brodo & Lendemer 2012), the presence or absence of lichexanthone was even regarded to be of higher taxonomic value than the mode of reproduction. Although occasionally specimens are found that contain lichexanthone

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only in low concentrations or only in part of the thallus, this can usually be explained by poor preservation or partial decay (and hence decomposition of the substance) already occurring in the field. The acceptance of lichexanthone-containing and lichexanthone-devoid specimens within one species in Polymeridium was aberrant and has not been confirmed by molecular data (Nelsen et al. submitted).

The same applies to the character of hamathecial inspersion. Hamathecial inspersion occurs in species of various lichen groups. The morphology differs between families, probably depending both on the hamathecial gel and on the oil causing the inspersion. To date, it is generally accepted as a valid species character, even in groups where the inspersion can be fine or where fresh specimens may have a similar (but less regular) inspersion that soon vanishes leading to false positives, such as Graphidaceae (Lücking et al. 2009) or Pyrenulaceae (Aptroot 2012, Aptroot et al. 2013a). In Trypetheliaceae, the hamathecial inspersion is a clear-cut character: relatively large globules in gel which often become distorted (oval to elongate). Some variation in size occurs (cf. Fig 4D & 4G) but the character is usually easily interpreted. Some species groups show only one of the character states; e.g. none of the species in the large Astrothelium conicum aggregate are inspersed, and inspersion with yellow oil only occurs in two species groups within the family.

Application of a narrower species concept in Polymeridium leads to the recognition of 53 species, 38 of which are known from Brazil, which remains the center of diversity of this genus. It is remarkable that the vast majority of the Polymeridium specimens present in NE Brazil contain lichexanthone in the thallus and lack hamathecial inspersion. Molecular studies are needed to test whether this is due to adaptation or a shared ancestor.

The genus is very homogenous in many characters, such as thallus organization (non-corticate and white with only few exceptions), ascomatal wall and shape (although locally the separate species can often be recognized by differences in ascoma dimensions), hamathecial filaments, ascus-shape. The monotypic genus Exiliseptum is included here in Polymeridium, as it differs in only two characters (submuriform ascospores and corticate thallus) from Polymeridium dithecium. These characters, although unusual, are not unique within Polymeridium, as P. corticatum has a corticate thallus and P. cinereonigricans and P. submuriforme have submuriform ascospores.

Three groups can be distinguished within the genus at present, each of which may turn out to be phylogenetically distinct groups (Nelsen et al. submitted).

The P. sulphurescens-group is characterized by relatively broadly ellipsoid ascospores, the tendency to have a white ring around the ostiole and the often exserted to superficial, somewhat barrel-shaped ascomata, which are similar to Trypethelium tropicum (Ach.) Müll. Arg. Trypethelium tropicum differs strongly by the angular ascospore lumina, but is phylogenetically close and may fall into this group.

The P. proponens-group is characterized by muriform ascospores and usually eccentric and often fused ostioles.

The core group of the genus includes the remaining species with transversely septate or submuriform ascospores.

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The key not only gives the characters needed for identification, but gives key characters for all species, at least ostiole orientation (and fusion where applicable), hamathecial inspersion, absence/presence of lichexanthone (given as UV-reaction), ascospores septation and dimensions.

Worldwide key to the species of Polymeridium

1 Ascospores muriform or submuriform .......................................................................................21* Ascospores only transversely septate .......................................................................................11

2 Ascospores muriform, with over 10 cells, hamathecium not inspersed .....................................32* Ascospores submuriform, with ≤ 10 cells ..................................................................................9

3 Ostiole apical, thallus UV–, ascospores 25–29 × 11–13 μm ........................................................ ............................................................................ P. julelloides E.L.Lima, M.Cáceres & Aptroot3* Ostiole eccentric, sometimes fused and/or ascomata fused ........................................................4

4 Thallus UV+ yellow ...................................................................................................................54* Thallus UV– ...............................................................................................................................7

5 Ascomata deeply immersed in the bark, ascospores 2/ascus, 45–70 × 17–25 μm ....................... ......................................................................P. immersum Aptroot, A.A.Menezes & M.Cáceres5* Ascomata erumpent, ascospores 8/ascus ....................................................................................6

6 Ascospores 35–54 × 12–19 μm .................................................. P. proponens (Nyl.) R.C.Harris6* Ascospores 55–75 × 19–21 μm ...................P. paraproponens Aptroot, M.Cáceres & E.L.Lima

7 Ascospores 55–80 × 17–25 μm .................................... P. campylothelioides Aptroot & Sipman7* Ascospores under 55 μm long ....................................................................................................8

8 Ostiole with red, K+ green pigment inside, ascospores 33–55 × 11–15 μm ................................ .............................................................P. isohypocrellinum Xavier-Leite, M.Cáceres & Aptroot8* Ostiole without red pigment, ascospores 33–55 × 12–21 μm ....P. amylosporum (Vain.) Aptroot

9 Ostiole eccentric, fused, thallus corticate, UV+ yellow, hamathecium not inspersed, ascospores 15–16 × 6–8 μm ............................................................P. ocellatum (Müll. Arg.) Aptroot9* Ostiole apical, thallus not corticate, UV– .................................................................................10

10 Hamathecium inspersed, ascospores 18–20 × 6–7.5 μm ...................... P. submuriforme Aptroot10* Hamathecium not inspersed, ascospores 30–45 × 9–11 μm ......................................................... .........................................................................................P. cinereonigricans (Vain.) R.C.Harris

11 Ostiole eccentric, pointing in various directions ......................................................................1211* Ostiole at least in majority apical (or rarely all pointing sideways in the same direction) .......20

12 Ostioles and/or ascomata at least partly fused, hamathecium not inspersed ............................1312* Ostioles not fused .....................................................................................................................15

13 Ascospores 7–9-septate, 28–35 × 6–7.5 μm, thallus UV+ yellow..........P. dithecium R.C.Harris13* Ascospores 3-septate ................................................................................................................14

14 Thallus UV+ yellow, ascospores 28–32 × 9–11 μm ..... P. pyrenastroides R.C.Harris ex Aptroot14* Thallus UV–, ascospores 26–30 × 8–10 μm .......................................... P. biloculare R.C.Harris

15 Thallus UV+ yellow, hamathecium not inspersed, ascospores 3-septate, 24–33 × 6–10 μm ....... ............................................................................................... P. stramineoatrum (Vain.) Aptroot15* Thallus UV– .............................................................................................................................16

16 Hamathecium inspersed ............................................................................................................1716* Hamathecium not inspersed ......................................................................................................18

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17 Hamathecial inspersion hyaline, ascospores 3–6-septate, 22–26 × 7–9 μm ................................. ............................................................................................................P. refertum (Stirt.) Aptroot17* Hamathecial inspersion yellow, KOH+ purple, ascospores 8–12-septate, 40–55 × 9–11 μm, often breaking into part spores .......................................................... P. flavothecium R.C.Harris

18 Ascospores 3-septate, 30–35 × 12–13 μm ............................................... P. simulans R.C.Harris18* Ascospores 7–13-septate ..........................................................................................................19

19 Ascospores 7–9-septate, 40–50 × 8–9 μm ................................................P. neblinae R.C.Harris19* Ascospores 9–13-septate, (35–)50–75 × 12–16 μm ........................P. pleurothecium R.C.Harris

20 Hamathecium not inspersed ......................................................................................................2120* Hamathecium inspersed ............................................................................................................38

21 Thallus UV+ yellow .................................................................................................................2221* Thallus UV– .............................................................................................................................30

22 Ascospores 3-septate ................................................................................................................2322* Ascospores 4–13-septate ..........................................................................................................28

23 Ascospores 13–17 × 4–6 μm ................................P. albopruinosum (Makhija & Patw.) Aptroot23* Ascospores mostly over 16 μm long .........................................................................................24

24 Ascospore with septa IKI+ violet, 16–23 × 5–7 μm ........................... P. amyloideum R.C.Harris24* Ascospore septa IKI – ...............................................................................................................25

25 Ascospores 16–19 × 4.5–6 μm ..................................................... P. bengoanum (Vain.) Aptroot25* Ascospores mostly over 19 μm .................................................................................................26

26 Ascomata mostly exserted, ostiole flat and with white ring, ascospores relatively broad, ellipsoid, 17–25 × 7–11 μm ........................................................................ P. xanthoreagens Aptroot26* Ascomata erumpent, ostiole not flat and never with white ring, ascospores elongate .............27

27 Ascospores 20–23 × 6–8 μm ..................P. albidoreagens Aptroot, A.A.Menezes & M.Cáceres27* Ascospores 25–33 × 7–10 μm ...................................................P. catapastum (Nyl.) R.C.Harris

28 Ascospores (4–)5(–7)-septate, 19–26 × 5–7 μm ........................................ P. multiforme Aptroot28* Ascospores at least 7-septate ....................................................................................................29

29 Thallus not corticate, ascospores 7–11-septate, 28–39 × 6–9 μm ....... P. alboflavescens Aptroot29* Thallus corticate, ascospores 9–13-septate, 32–36 × 6–7 μm ...................................................... ..................................................................... P. corticatum A.A.Menezes, M.Cáceres & Aptroot


31 Ascospores 11–16 × 3–6 μm ...................................P. microsporum (Makhija & Patw.) Aptroot31* Ascospores mostly over 17 μm long .........................................................................................32

32 Ascomata mostly exserted, ostiole flat and with white ring, ascospores relatively broad, ellipsoid, 19–27 × 6.5–11 μm ..............................................P. sulphurescens (Müll.Arg.) R.C.Harris32* Ascomata erumpent, ostiole not flat and never with white ring, ascospores elongate .............33

33 Ascospores (15–)16–20 × 5–7.5 μm ........................................P. subcinereum (Nyl.) R.C.Harris33* Ascospores mostly over 20 μm .................................................................................................34

34 Ascospores (18–)20–23 × 6–8(–9) μm ................................. P. albidum (Müll. Arg.) R.C.Harris34* Ascospores over 23 μm .............................................................................................................35

35 Medulla white, ascospores 24–32 × 6–11 μm .......................................P. catapastoides Aptroot35* Medulla partly yellow, KOH+ purple, aascospores 30–34 × 9–10 μm ......................................... .........................................................................................................P. endocrocinum R.C.Harris

36 Ascospores (4–)5(-7)- septate, 21–30 × 5–9 μm ......................P. jordanii (C.W.Dodge) Aptroot36* Ascospores at least 7-septate ....................................................................................................37

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37 Ascospores 7–11-septate, (24–)28–39 × 6–9 μm ........ P. albocinereum (Krempelh.) R.C.Harris37* Ascospores 13–17-septate, 85–150 × 20–30 μm ..............P. bambusicola Aptroot & L.I.Ferraro

38 Thallus UV+ yellow .................................................................................................................3938* Thallus UV– .............................................................................................................................44


40 Ascospores 13–17 × 4–6 μm .................................................P. brachysporum (Malme) Aptroot40* Ascospores mostly over 17 μm .................................................................................................41

41 Ascospores 17–20 × 5–7 μm ...............................................P. pyrenuloides (Müll.Arg.) Aptroot41* Ascospores mostly over 20 μm .................................................................................................42

42 Ascospores 20–23 × 6–8 μm .................................................................P. albidovarians Aptroot42* Ascospores 24–33 × 6–10 μm ................................................. P. subvirescens (Leight.) Aptroot

43 Ascospores 4–7-septate, 18–28 × 5–7 μm ..P. multiseptatum Aptroot, A.A. Menezes & Cáceres43* Ascospores 8–12-septate, 40–50 × 9–11 μm .............................P. chioneum (Mont.) R.C.Harris


45 Ascospores 12–16 × 4–6 μm .....................................................P. contendens (Nyl.) R.C.Harris45* Ascospores mostly over 16 μm .................................................................................................46

46 Ascomata mostly exserted, ostiole flat and with white ring, ascospores relatively broad, ellipsoid, 20–24 × 7–10 μm ............................................................................ P. tribulationis Aptroot46* Ascomata erumpent, ostiole not flat and never with white ring, ascospores elongate .............47

47 Ascospores 16–20 × 3–5 μm ...................................................................... P. inspersum Aptroot47* Ascospores mostly over 20 μm .................................................................................................48

48 Ascospores 20–23 × 5–7 μm ............................................................ P. siamense (Vain.) Aptroot48* Ascospores over 23 μm .............................................................................................................49

49 Ascospores smooth, 24–30 × 6–9 μm .............................................P. suffusum (Knight) Aptroot49* Ascospores with rough wall, 26–34 × 6–9 μm ...................... P. glaucoatrum (Vain.) R.C.Harris

50 Ascospores with rough wall, 7-septate, 30–33 × 9–10 μm ......................... P. neuwirthii Aptroot50* Ascospores smooth ...................................................................................................................51

51 Ascospores (4–)5(-7)- septate, 18–28 × 4–7 μm ..............P. quinqueseptatum (Nyl.) R.C.Harris51* Ascospores at least 7-septate ....................................................................................................52

52 Ascospores 7–11-septate, 25–36 × 5–9 μm ................. P. pleiomerellum (Müll.Arg.) R.C.Harris52* Ascospores 8–12-septate, 40–50 × 9–11 μm ...........................................P. costaricense Aptroot

Taxonomy

Polymeridium (Müll.Arg.) R.C.Harris, in Tucker & Harris, Bryologist 83: 12 (1980)

Arthopyrenia sect. Polymeridium Müll.Arg., Flora 46: 317 (1883).

Typus: Arthopyrenia contendens (Nyl.) Müll.Arg. [= Polymeridium contendens (Nyl.) R.C.Harris]

Exiliseptum R.C.Harris, Acta Amazon. (Suppl.) 14: 65 ('1984') [1986]. Typus: Exiliseptum ocellatum (Müll.Arg.) R.C.Harris

DescripTion: Thallus white to yellowish or grey, usually ecorticate. Ascomata simple or fused with partly shared walls or only the ostioles fused, black, without pseudostromatic

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tissues, globose to pyriform, erumpent from the substratum. Ostiole apical to lateral, brown to black or with red anthraquinone. Hamathecium colourless, sometimes inspersed with hyaline or red (KOH+ green) oil droplets. Ascospores 4–8 per ascus, IKI– or rarely IKI+ violet, irregularly biseriate, colourless, ellipsoid to fusiform with rounded to subacute ends, symmetrically 3–13-septate to muriform, not constricted at the septa, sometimes surrounded by a thin to 2–5 μm thick gelatinous sheath; septa not thickened; lumina rectangular, slightly rounded but not diamond-shaped. Conidiomata rather rare. Chemistry: Thallus occasionally with lichexanthone, anthraquinones rarely present in ostiole or hamathecium.

Polymeridium albidoreagens Aptroot, A.A.Menezes & M.Cáceres, sp. nov. Fig. 5

MycoBank: MB802982.

Diagnosis: Like Polymeridium albidum but thallus UV+ yellow.

Typus: Brazil. ronDônia: Porto Velho, Parque Natural Municipal; alt. 100 m; on tree bark; 16 Nov. 2012, M.E.S.Cáceres & A.Aptroot 15194 (ISE – holotype; ABL – isotype).

DescripTion: Thallus ecorticate, whitish grey, UV+ yellow. Ascomata 0.2–0.4 mm diam., centrum globose, erumpent, solitary. Ostiole apical. Hamathecium not inspersed. Ascospores 8/ascus, IKI–, 3-septate, 20–23 × 6–8 μm, not ornamented. Chemistry: lichexanthone.

noTe: Additional material referable to this species was cited by Harris (1993) from Brazil.

aDDiTional speciMens exaMineD: Brazil. ceará: Chapada do Araraipe; alt. 800 m; May 2012, A.A.Menezes 276 (ISE 15901); 754 (ISE 15902).

Polymeridium albidovarians Aptroot, sp. nov. Figs 1A, 3A

MycoBank: MB802983.

Diagnosis: Like Polymeridium albidum but thallus UV+ yellow and hamathecium inspersed.

Typus: inDonesia. Java: Kawi-complex, Tjemarakandang; alt. 2700 m; on twig; 1937, P.Groenhart 5755 (L – holotype; ABL – isotype).

DescripTion: Thallus ecorticate, yellowish white, UV+ yellow. Ascomata 0.2–0.4 mm diam., centrum globose, erumpent, solitary. Ostiole apical. Hamathecium inspersed with oil droplets. Ascospores 8/ascus, IKI–, 3-septate, 20–23 × 6–8 μm, not ornamented. Chemistry: lichexanthone.


Polymeridium albidum (Müll.Arg.) R.C.Harris, Acta Amazon. (Suppl.) 14: 69 ('1984') [1986]

Arthopyrenia albida Müll.Arg., Flora 67: 664 (1884). Typus: Brazil. Bahia: Caetité, G.Martius (G – holotype, not seen).

illusTraTion: Aptroot, Fl. Australia 57: 539, fig. 171C (2009).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole apical. Hamathecium not inspersed. Ascospores IKI–, 3-septate, (18–)20–23 × 6–8(–9) μm. Chemistry: No substances detected.

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DisTriBuTion: Pantropical; USA (Harris 1993), Colombia (Sipman et al. 2008), Brazil, Mozambique (Harris 1993), Sarawak (Harris 1993), Papua New Guinea (Aptroot et al. 1997), Australia (Aptroot 2009). Incorrectly reported from Thailand.

new counTry recorDs: venezuela. MeriDa: Sucre, Funca Los Topos, H.J.M.Sipman & M.Morales 38060, 1996 (B). Bolivia. la paz: Sud Yunga, Alto Beni, Sapecho, Acebey & Krömer, 995b, 1999 (B). Malaysia. Johor: Gunung Pulai Forest Reserve, H.J.M.Sipman, B.Tan & Farida 46439, 2000 (B)

Polymeridium albocinereum (Krempelh.) R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 625 ('1991') [1993]

Verrucaria albocinerea Krempelh., Flora 59: 524 (1876); Arthopyrenia albocinerea (Krempelh.) Müll.Arg., Flora 66: 318 (1883). Typus: Brazil. rio De Janeiro: A.Glaziou 5025 (M – lectotype, not seen).

Pseudopyrenula pentameria Vain., Ann. Soc.Zool. Bot. Fenn. Vanamo 1(3): 55 (1921). Typus: Thai-lanD. Doi Suthep, C.C. Hosseus (G – holotype, not seen).

illusTraTion: McCarthy, Australas. Lichenol. Newslett. 26: 17, fig. 1A (1995).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole apical. Hamathecium not inspersed. Ascospores IKI–, 7–11-septate, (24–)28–39 × 6–9 μm. Chemistry: No substances detected.

DisTriBuTion: Probably pantropical; Brazil, Tanzania (Aptroot & Alstrup), India (Upreti & Nayaka 2006), Thailand.

new counTry recorD: coloMBia. caqueTa: Ararauara, road to airstrip, H.J.M.Sipman & J.Duiven-voorden 27826, 1988 (B)

Polymeridium alboflavescens Aptroot sp. nov. Figs 1B, 3B, 4A

MycoBank: MB802984.

Diagnosis: Like Polymeridium albocinereum but thallus UV+ yellow.

Typus: venezuela. aMazonas: Alto Orinoco, 15 km W of Esmeralda; alt. 110 m; on tree bark; 14 Feb 1997, J.Hafellner & H.Komposch 991-6-4 (GZU – holotype).

DescripTion: Thallus ecorticate, whitish grey, UV+ yellow. Ascomata 0.3–0.5 mm diam., centrum globose, erumpent, solitary. Ostiole apical. Hamathecium not inspersed. Ascospores 8/ascus, IKI–, 7–11-septate, 28–39 × 6–9 μm, not ornamented. Chemistry: lichexanthone.


aDDiTional speciMens exaMineD: Brazil. ceará: Chapada do Araraipe; alt. 800 m; May 2012, A.A.Menezes 166 (ISE 15903), 289 (ISE 15904), 671 (ISE 15905), 521 (ISE 15906), 763 (ISE 15907), 765 (ISE 15908), 770 (ISE 15909), 285 (ISE 15910), 265 (ISE 15911), 289 (ISE 15912), 494 (ISE 15913), 124 (ISE 15914).

Polymeridium albopruinosum (Makhija & Patw.) Aptroot comb. nov.

MycoBank: MB802985

Trypethelium albopruinosum Makhija & Patw., J. Hattori Bot. Lab. 73: 193 (1993). Typus: inDia. anDaMan islanDs: Middle Andaman, Dhanimali; on tree bark; 26 Dec 1985, P.G.Patwardhan & M.B.Nagarkar AMH 85.2342 (ABL, BM – isotypes).

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DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia solitary. Ostiole apical. Hamathecium not inspersed with oil droplets. Ascospores IKI–, 3-septate, 13–17 × 4–6 μm. Chemistry: lichexanthone.

DisTriBuTion: Pantropical: Brazil (Harris 1993, as P. contendens, "Thallus UV+; hymenium not inspersed"), India.

new counTry recorD: Tanzania. Bagmoyo Distr., Mapinga, Changwahela, V.Alstrup TZ4072b, 1999 (C, ABL).

Fig. 1. Habitus of Polymeridium species, all from holotype. A. P. albidovarians, B. P. alboflavescens, C. P. catapastoides, D. P. inspersum, E. P. multiforme, F. P. neuwirthii. Bar = 0.2 mm.

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Polymeridium amyloideum R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 626 ('1991') [1993].

Typus: Brazil. pará: Serra do Cachimbo, L.Brako 6107 (NY – isotype).

illusTraTion: Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 628, fig. 1–2 ('1991') [1993].

DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia solitary joint ostiole. Ostiole lateral. Hamathecium not inspersed. Ascospore septa IKI+ violet, 3-septate, 16–23 × 5–7 μm. Chemistry: lichexanthone.

DisTriBuTion: Neotropical (only known from Brazil).

Polymeridium amylosporum (Vain.) Aptroot comb. nov.

MycoBank: MB802986.

Thelenella amylospora Vain., Acta Soc. Fauna Flora Fenn. 7(2): 218 (1890); Polyblastiopsis amylospora (Vain.) Zahlbr., Catal. Lich. Univ. 1: 347 (1922); Campylothelium amylosporum (Vain.) R.C.Harris, in Tucker & Harris, Bryologist 83: 7 (1980). Typus: Brazil. rio De Janeiro: Sepetiba; on tree bark; E.A.Vainio, Lich. Bras. 419 (UPS – isotype).

Polyblastiopsis dealbens Fink, in C.Hedrick, Mycologia 25: 307 (1933). Typus: USA. souTh carolina: Green (MICH-Fink 11252, lectotype, selected by R.C.Harris, Contr. Univ. Michigan Herb. 11: 96 (1975), not seen).

illusTraTions: Tucker & Harris, Bryologist 83: 8, fig. 14–17 (1980); Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 628, fig. 12 ('1991') [1993]; Aptroot, Lucking, Sipman, Umaña & Chaves, Biblioth. Lichenol. 97: 86, fig. 14C (2008).

DescripTion: Thallus ecorticate, white, UV-. Perithecia solitary. Ostiole lateral. Hamathecium not inspersed, filaments profusely anastomosing. Ascospores 8/ascus, IKI+ violet, muriform, 33–55 × 12–21 μm, with partly oblique septa, not ornamented, wall not thickened. Chemistry: no substances.

DisTriBuTion: Pantropical: Hawai’i, USA (Harris 1993), Mexico, Costa Rica (Aptroot et al. 2008), Puerto Rico (Harris 1993), Brazil (Harris 1993), South Africa (Harris 1993), Seychelles (Seaward & Aptroot 1999), Hong Kong (Aptroot & Seaward 1999), Fiji (Harris 1993), New Caledonia, Papua New Guinea (Aptroot et al. 1997), Australia (Aptroot 2009). The species has recently often been reported as P. proponens; older reports of P. amylosporum may refer to either this species or P. proponens).

new counTry recorD: inDia. West Begal: Sundarbans Bisosphere Reserve, Chamta, T.A.M.Jagadeesh Ram 13571, 2003 (ASSAM, ABL).

Polymeridium bambusicola Aptroot & Ferraro, Bonplandia 10: 140 (2000).

Typus: argenTina. corrienTes: Dep. Mercedes, Macrosistema Iberá, Estancia Rincón del Diablo, M.M.Arbo, A.Schinini, V.Marunak & F.Zanier 8269a (ABL – isotype).

illusTraTion: Aptroot & Ferraro, Bonplandia 10: 140, fig. 1 (2000).

DescripTion: Thallus absent, UV–, not lichenized. Perithecia solitary. Ostiole apical. Hamathecium not inspersed. Ascospores IKI–, 13–17-septate, 85–150 × 20–30 μm. Chemistry: No substances detected.

DisTriBuTion: Neotropical (only known from Argentina).

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Polymeridium bengoanum (Vain.) Aptroot comb. nov.

MycoBank: MB802987.

Pseudopyrenula bengoana Vain., in Hiern, Cat. Afr. Pl. 2(2): 457 (1901). Typus: angola. Quifandango; on tree bark; F.M.J.Welwitsch 437 (BM – isotype).

DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia solitary. Ostiole apical. Hamathecium not inspersed with oil droplets, filaments profusely anastomosing. Ascospores 8/ascus, IKI–, 3-septate, 16–19 × 4.5–6 μm, not ornamented, wall not thickened. Chemistry: lichexanthone.

DisTriBuTion: Pantropical: Venezuela (Harris 1993), Angola, Australia (Aptroot 2009, sub P. subcinereum).

new counTry recorDs: usa. california: Fern Canyon, Redwood National Park, C.M.vanHerk, 1995 (ABL). souTh africa. naTal: Oribi Gorge Nature Reserve, alt. 450 m, D.Triebel & G.Rambold 7303, 1990 (ABL, M).

Polymeridium biloculare R.C.Harris, Acta Amazon. (Suppl.) 14: 70 ('1984') [1986]. Typus: Brazil. roraiMa: Manaus-Boa Vista-road, W.R.Buck (NY– isotype).

illusTraTion: Harris, Acta Amazon. (Suppl.) 14: 78, fig. 8 ('1984') [1986].

DescripTion: Thallus ecorticate, white, UV–. Perithecia 2 aggregated with joint ostiole. Ostiole lateral. Hamathecium not inspersed. Ascospores IKI–, 3-septate, 26–30 × 8–10 μm, surrounded by a gelatinous sheath. Chemistry: No substances detected.


Polymeridium brachysporum (Malme) Aptroot comb. nov.

MycoBank: MB802988.

Trypethelium brachysporum Malme, Ark. f. Bot. 19(1): 30 (1924). Typus: Brazil. MaTo grosso: Cuiabá; on tree bark; G.O.A.Malme 2055 (BM – isotype).

DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia solitary. Ostiole apical. Hamathecium inspersed with oil droplets. Ascospores IKI–, 3-septate, 13–17 × 4–6 μm. Chemistry: lichexanthone.


Polymeridium campylothelioides Aptroot & Sipman, in Aptroot, Diederich, Sérusiaux & Sipman, Biblioth. Lichenol. 57: 37 (1995).

Typus: papua new guinea. MaDang prov.: Laing Island in Hansa Bay near Bogia, H.J.M.Sipman 34767 (B – holotype).

illusTraTions: Aptroot, Diederich, Sérusiaux & Sipman, Biblioth. Lichenol. 57: 46, fig. 2e; 48, fig. 7 (1995).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary or 2 aggregated with common ostiole. Ostiole lateral. Hamathecium not inspersed, filaments profusely anastomosing. Ascospores 8/ascus, IKI-, muriform, 55–80 × 17–25 μm, with partly oblique septa, not ornamented, wall not thickened. Chemistry: no substances.

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DisTriBuTion: Asian: Taiwan (Aptroot 2003), Papua New Guinea. Incorrectly reported from Venezuela (Komposch & Hafellner 2006); this is probably P. paraproponens.

Polymeridium catapastoides Aptroot sp. nov. Figs 1C, 3C, 4B

MycoBank: MB802989.

Diagnosis: Like Polymeridium catapastum but thallus UV-.

Typus: ausTralia. queenslanD: Cape Tribulation, N of Daintree; on tree bark; 4 Jul 1983, M.E.Hale 64184 (US – holotype; ABL – isotype).

DescripTion: Thallus ecorticate, yellowish white, UV–. Ascomata 0.4–0.6 mm diam., centrum globose, erumpent, solitary. Ostiole apical or partly lateral. Hamathecium not inspersed with oil droplets. Ascospores 4–8/ascus, IKI–, 3-septate, 24–32 × 6–11 μm, not ornamented. Chemistry: no substances.

noTe: Also reported from Malaysia (Harris 1993) and Thailand (Aptroot et al. 2007, as P. catapastum).

aDDiTional speciMens exaMineD: ausTralia. queenslanD: Atherton Tablelands, Crater Lake; 1999, A.Aptroot 46182 (ABL). inDonesia. Java: Mt. Gede, Cibodas; 1939, P.Groenhart 3219 (ABL, L). papua new guinea. chiMBu: Mount Wilhelm area, near Lake Piunde; Mar 1987, A.Aptroot 18279 (ABL). Brazil. ceará: Chapada do Araraipe; May 2012, A.A.Menezes 776 (ISE 15915), 777 (ISE 15916), 777a (ISE 15917), 778 (ISE 15918), 522 (ISE 15919), 760 (ISE 15920), 525 (ISE 15921), 543 (ISE 15922), 046 (ISE 15923), 525 (ISE 15924), 454 (ISE 15925), 152 (ISE 15926), 508 (ISE 15927).

Polymeridium catapastum (Nyl.) R.C.Harris, Acta Amazon. (Suppl.) 14: 70 ('1984') [1986]

Verrucaria catapasta Nyl., Acta Soc. Sci. Fenn. 7: 488 (1863); Arthopyrenia catapasta (Nyl.) Müll. Arg., Flora 66: 318 (1883). Typus: coloMBia. Bogotá, A.Lindig 2869 (BM – isotype).

Pyrenastrum album var. verrucarioides Eschw., in Mart., Flora Bras. 1: 147 (1833); Arthopyrenia tumida Müll.Arg., Flora 67: 669 (1884). Typus: Brazil. Bahia: G.Martius (G – lectotype, selected by R.C.Harris, Acta Amazon. (Suppl.) 14: 70 ('1984') [1986], not seen).

DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia solitary. Ostiole apical. Hamathecium not inspersed. Ascospores IKI–, 3-septate, 25–33 × 7–10 μm. Chemistry: lichexanthone.

DisTriBuTion: Neotropical: USA (Harris 1993), Bahamas (Harris 1993), Cayman Islands (Harris 1993), Cuba (Harris 1993), El Salvador (Sipman 2001), Guyana, Colombia (Sipman et al. 2008), Venezuela (Harris 1993), Brazil (Harris 1993).

Polymeridium chioneum (Mont.) R.C.Harris, Lichenogr. Thomsoniana 141 (1998)

Verrucaria chionea Mont., Ann. Sci. Nat. Bot. sér. 2, 19: 58 (1843); Pyrenula chionea (Mont.) Trevis., Spighe e Paglie: 17 (1853); Pseudopyrenula chionea (Mont.) Zahlbr., Catal. Lich. Univ. 1: 355 (1922). Typus: french guiana. F.R.Leprieur 613 (PC-Mont – lectotype, selected by R.C.Harris, Lichenogr. Thomsoniana 141, 1998, not seen).

Polymeridium pleiomeroides (Müll.Arg.) R.C.Harris, Acta Amazon. (Suppl.) 14: 70 ('1984') [1986]; Arthopyrenia pleiomeroides Müll.Arg., Bot. Jahrb. Syst. 6: 407 (1885). Typus: cuBa. C.Wright, Verr. Cub. 76 (G – holotype, not seen).

illusTraTion: Aptroot, Lucking, Sipman, Umaña & Chaves, Biblioth. Lichenol. 97: 86, fig. 14A (2008).

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DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia solitary. Ostiole apical. Hamathecium inspersed, filaments profusely anastomosing. Ascospores 8/ascus, hyaline, IKI–, 8–12-septate, 40–50 × 9–11 μm, not ornamented, wall not thickened. Chemistry: lichexanthone.

DisTriBuTion: Neotropical: Cuba, Venezuela (Komposch & Hafellner 2006), French Guiana, Guyana (Sipman & Aptroot 1992), Brazil (Harris 1993).

Polymeridium cinereonigricans (Vain.) R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 631 ('1991') [1993]

Thelenella cinereonigricans Vain., Acta Soc. Fauna Fl. fenn. 7(2): 216 (1890); Clathroporina cinereonigricans (Vain.) Zahlbr., Catal. Lich. Univ. 1: 416 (1922). Type: Brazil. Minas gerais: Carassa, E.A.Vainio, Lich. Bras. 657 (BM – isotype).

illusTraTion: Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 628, fig. 3 ('1991') [1993].

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole apical. Hamathecium not inspersed. Ascospores IKI–, submuriform, 8–11 × 1–2-septate, 30–45 × 9–11 μm. Chemistry: No substances detected.

DisTriBuTion: Neotropical: Hawai’i, Brazil.

Fig. 2. Habitus of Polymeridium species, all from holotype. A. P. pyrenastroides, B. P. submuriforme, C. P. tribulationis, D. P. xanthoreagens. Bar = 0.2 mm.

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Polymeridium contendens (Nyl.) R.C.Harris, in Tucker & Harris, Bryologist 83: 12 (1980)

Verrucaria contendens Nyl., Acta Soc. Sci Fenn. 7: 492 (1863); Arthopyrenia contendens (Nyl.) Müll.Arg., Flora 66: 317 (1883). Typus: coloMBia. San Antonio, Lindig 2877 (BM – isotype).

Arthopyrenia atroalba Vain., Acta Soc. Fauna Fl. fenn. 7(2): 234 (1890). Typus: Brazil. Minas gerais: Lafayette, E.A.Vainio, Lich. Bras. 334 (BM – isotype).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole apical. Hamathecium inspersed with oil droplets. Ascospores IKI–, 3-septate, 12–16 × 4–6 μm. Chemistry: No substances detected.

DisTriBuTion: Neotropical: USA (Harris 1993), Costa Rica (Aptroot et al. 2008), Colombia, Brazil. Incorrectly reported from Australia, Papua New Guinea and Tanzania.

new counTry recorDs: guyana. rupununi DisTr.: Monkey Pod Landing, SW of Mt. Makarapan, P.J.M.Maas et al. 7373a, 1988 (ABL). venezuela. Bolivar: Cerro Guaiquinima, alt. 1000 m, H.J.M.Sipman 26547, 1990 (B).

Polymeridium corticatum A.A.Menezes, M.Cáceres & Aptroot, Lichenologist 45: 546.

Typus: Brazil. ceará: Chapada do Araripe, on bark of tree, 15 May 2012, A.A. Menezes P1-A3-T-CA (ISE – holotype).

DescripTion: See Aptroot et al. (2013b).

DisTriBuTion: Neotropical (Brazil). Polymeridium costaricense Aptroot sp. nov.

MycoBank: MB802990.

Diagnosis: Like Polymeridium chioneum but thallus UV–.

Typus: cosTa rica. guanacasTe: Palo Verde National Park, trail to Rio Tempisque, 0–50 m; on tree bark; 28 Mar 2003, S.Will-Wolf 12512d (INB – holotype; WIS – isotype).

illusTraTion: Aptroot, Lucking, Sipman, Umaña & Chaves, 2008: 86, Fig. 14a, as P. chioneum.

DescripTion: Thallus ecorticate, whitish grey, UV–. Ascomata 0.4–0.6 mm diam., centrum globose, erumpent, solitary. Ostiole apical. Hamathecium inspersed, filaments profusely anastomosing. Ascospores 8/ascus, hyaline, IKI–, 8–12-septate, 40–50 × 9–11 μm, not ornamented, wall not thickened. Chemistry: No substances.

noTe: The type specimen was reported and illustrated as P. chioneum by Aptroot et al. (2008).

aDDiTional speciMens exaMineD: Brazil. ceará: Chapada do Araraipe; May 2012, A.A.Menezes 176 (ISE 15928), 775 (ISE 15929), 486 (ISE 15930), 123 (ISE 15931), 138 (ISE 15932).

Polymeridium dithecium R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 632 ('1991') [1993].

Typus: venezuela. aMazonas: Río Casiquiare, K.Kubitzki (NY– holotype).


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DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia 2 aggregated with joint ostiole. Ostiole lateral. Hamathecium not inspersed. Ascospores IKI–, 7–9-septate, 28–35 × 6–7.5 μm. Chemistry: Thallus with lichexanthone.

DisTriBuTion: Neotropical (only known from Venezuela).

Polymeridium endocrocinum R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 633 ('1991') [1993].

Typus: Brazil. aMazonas: Serra Aracá plateau, G.Samuels 521 (NY– holotype).


Fig. 3. Ascospores of Polymeridium species, all from holotype. A. P. albidovarians, B. P. albo-flavescens, C. P. catapastoides, D. P. inspersum, E. P. multiforme, F–G. P. neuwirthii (showing ornamentation in two different focal view), H–J P. pyrenastroides, (H with immature ascospore and J overmature ascospore), K. P. submuriforme, L. P. tribulationis, M. P. xanthoreagens. Bar = 10 μm.

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DescripTion: Thallus ecorticate, white, UV-. Perithecia solitary, with orange, KOH+ purple pigment inside. Ostiole apical. Hamathecium not inspersed. Ascospores IKI–, 3-septate, 30–34 × 9–10 μm. Chemistry: Pigment KOH+ purple.


Polymeridium flavothecium R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 633 ('1991') [1993].

Typus: DoMinican repuBlic. DisT. nacional: Sierra Prieta, 18 km NW of Villa Melia, R.C.Harris 20160 (NY– holotype).


DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole lateral. Hamathecium with yellow, KOH+ red granules below, filaments profusely anastomosing. Ascospores 8/ascus, hyaline, IKI–, 8–12-septate, 40–55 × 9–11 μm, not ornamented, breaking into part spores. Chemistry: Pigment KOH+ purple.

DisTriBuTion: Neotropical (Dominican Republic, Costa Rica).

Polymeridium glaucoatrum (Vain.) R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 634 ('1991') [1993]

Arthopyrenia glaucoatra Vain., Ann. Acad. Sci Fenn. Ser. A, 19(15): 12 (1923). Typus: philippines. luzon: Benguet prov., Pauai, R.C.McGregor (TUR – holotype, not seen).

DescripTion: Thallus ecorticate, white, UV-. Perithecia solitary. Ostiole apical. Hamathecium inspersed. Ascospores IKI–, 3-septate, 26–34 × 6–9 μm, granular ornamented. Chemistry: no substances.

DisTriBuTion: Asian: Sri Lanka (Harris 1993), Philippines.

Polymeridium immersum Aptroot, A.A.Menezes & M.Cáceres, Lichenologist 45: 546.

Typus: Brazil. ronDônia: Porto Velho, UNIR Federal University campus S of city, on bark of tree, c. 100 m alt., 8 Mar. 2012, M.E.S.Cáceres & A.Aptroot 11138 (ISE – holotype; ABL – isotype).


Neotropical (Brazil).

Polymeridium inspersum Aptroot sp. nov. Figs 1D, 3D, 4D

MycoBank: MB802991.

Diagnosis: Like Polymeridium subcinereum but hamathecium inspersed.

Typus: ausTralia. queenslanD: Kuranda, Jumrum Creek track; on twig; Mar 1988, A. & M.Aptroot 22755 (CANB – holotype; ABL – isotype).

DescripTion: Thallus ecorticate, pinkish grey, UV–. Ascomata 0.4–0.6 mm diam., centrum globose, erumpent, solitary. Ostiole apical. Hamathecium inspersed with oil droplets, filaments profusely anastomosing. Ascospores 8/ascus, IKI–, 3-septate, 16–20 × 3–5 μm, not ornamented, wall not thickened. Chemistry: no substances.

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Fig. 4. Hamathecium of Polymeridium species, all from holotype. A. P. alboflavescens, B. P. cata-pastoides, C. P. tribulationis, D. P. inspersum, E. P. multiforme, F. P. submuriforme, G. P. neuwirthii, H. P. xanthoreagens. Bar = 20 μm.

noTe: Pantropical; material referable to this species was reported by Harris (1993) from Brazil and the Philippines. The type and the other specimen from Australia were reported as P. subcinereum by Aptroot (2009).aDDiTional speciMens exaMineD: ausTralia. queenslanD: Bellenden Ker National Park, centre peak, 1988, A. & M.Aptroot 22508 (ABL). Brazil. ceará: Chapada do Araraipe; May 2012, A.A.Menezes 779 (ISE 15933), 780 (ISE 15934), 781 (ISE 15935).

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Polymeridium isohypocrellinum Xavier-Leite, M.Cáceres & Aptroot, Lichenologist 45: 548.Typus: Brazil. paraíBa: Reserve Muralha, on bark of tree, 16 Jul. 2012, A.B.X.Leite T1/A3 (ISE – holotype).



Polymeridium jordanii (C.W.Dodge) Aptroot comb. nov.MycoBank: MB802992.

Pseudopyrenula jordanii C.W.Dodge, Nova Hedwigia, Beiheft 12: 11 (1964). Typus: sierra leone. Colony, Sugar Loaf Mountain; on tree bark; F.C.Deighton M44440 (FH-Dodge – holotype).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole apical. Hamathecium not inspersed, filaments profusely anastomosing. Ascospores 8/ascus, IKI–, 4–7-septate, 21–30 × 5–9 μm, not ornamented, wall not thickened. Chemistry: no substances detected.

DisTriBuTion: Pantropical: USA (Harris 1993), Puerto Rico (Harris 1993), Mozambique (Harris 1993), Sierra Leone, Hong Kong (Harris 1993).new counTry recorDs: seychelles. alDaBra: Cinque Cases, R.J.Hnatiuk 730181, 1973 (BM, ABL). yeMen. socoTra: Hadibu, M.Schultz 14047c, 1997 (ABL, HBG). philippines. palawan: Iwahig, P.Diederich 13323, 1994 (hb. Diederich, ABL). inDonesia. Bali: Nusa Dua, T.Gandjar, 1995 (ABL).

Polymeridium julelloides E.L.Lima, M.Cáceres & Aptroot, Lichenologist 45: 548.Typus: Brazil. pernaMBuco: Buíque, Vale do Catimbau National Park, on bark of tree, c. 900 m alt., 3 Oct. 2011, E.L.Lima 145 (ISE – holotype).



Polymeridium microsporum (Makhija & Patw.) Aptroot comb. nov.MycoBank: MB802993.

Trypethelium microsporum Makhija & Patw. J. Hattori Bot. Lab. 73: 201 (1993). Typus: inDia. anDaMan islanDs: Middle Andaman, Yeratta; on tree bark; 24 Dec 1985, P.K.Sethy & M.B.Nagarkar AMH 85.2210 (ABL, BM – isotypes).


DisTriBuTion: Asian (India).new counTry recorDs: papua new guinea. easTern highlanDs: Mount Gahavisuka, A.Aptroot 31245, 1992 (ABL). hawai’i. hawai’i islanD: Volcano National Park, alt. 1200 m, A. & M.Aptroot 26520, 1989 (ABL). fiJi. Koroninia Research Station, Fairman, 1969 (ABL, L).

Polymeridium multiforme Aptroot, sp. nov. Figs 1E, 3E, 4E

MycoBank: MB802994.

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Diagnosis: Like Polymeridium quinqueseptatum but thallus UV+ yellow and hamathecium not inspersed.

Typus: guyana. rupununi: trail at base of Mt. Makarapan; on Myrsinaceae tree bark; 20 Sep 1988, P.J.M.Maas 7586c (B – holotype; ABL – isotype).

DescripTion: Thallus ecorticate, whitish grey, UV+ yellow. Ascomata 0.2–0.3 mm diam., centrum globose, erumpent, solitary. Ostiole apical. Hamathecium not inspersed, filaments profusely anastomosing. Ascospores 8/ascus, IKI–, 4–7-septate, 19–26 × 5–7 μm, not ornamented, wall not thickened. Chemistry: lichexanthone.

noTe: Neotropical; material referable to this species was reported by Harris (1993) from Brazil and Venezuela.

aDDiTional speciMens exaMineD: Brazil. ceará: Chapada do Araraipe; May 2012, A.A.Menezes 535 (ISE 15937), 042 (ISE 15938), 543 (ISE 15939), 543 (ISE 15940), 774 (ISE 15941), 759 (ISE 15942).

Polymeridium multiseptatum Aptroot, A.A.Menezes & Cáceres sp. nov.

MycoBank: MB802995.

Diagnosis: Like Polymeridium quinqueseptatum but thallus UV+ yellow.

Fig. 5. Polymeridium albidoreagens – isotype. A. Thallus with ascomata, B. Thallus with ascomata under UV light, C. hamathecium, D. Ascospores. Bar = 2 mm in A & B, 20 μm in C and 10 μm in D.

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Typus: Brazil. ceará: Chapada do Araraipe; alt. 800 m; on tree bark; May 2012, A.A.Menezes 782 (ISE 15943 – holotype); 783 (ISE 15944 – paratype), 784 (ISE 15945 – paratype), 785 (ISE 15946 – paratype), 128 (ISE 15947 – paratype).

DescripTion: Thallus ecorticate, whitish grey, UV+ yellow. Ascomata 0.2–0.4 mm diam., centrum globose, erumpent, solitary. Ostiole apical. Hamathecium inspersed with oil droplets, filaments profusely anastomosing. Ascospores 8/ascus, IKI–, 4–7-septate, 18–28 × 5–7 μm, not ornamented, wall not thickened. Chemistry: lichexanthone.

noTe: Additional material referable to this species was cited by Harris (1993) from Angola.

Polymeridium neblinae R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 635 ('1991') [1993].

Typus: venezuela. aMazonas: Cerro de la Neblina, W.R.Buck 10788 (NY – holotype).


DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole lateral. Hamathecium not inspersed, filaments profusely anastomosing. Ascospores 8/ascus, hyaline, IKI–, 7–9-septate, 40–50 × 8–9 μm, not ornamented. Chemistry: No substances detected.

DisTriBuTion: Neotropical (only known from Venezuela).

Polymeridium neuwirthii Aptroot sp. nov. Figs 1F, 3F–G, 4G

MycoBank: MB802996.

Diagnosis: Like Polymeridium quinqueseptatum but hamathecium inspersed with very large oil droplets and ascospores rough and consistently 7-septate.

Typus: venezuela. Puerto Ajacucho, at Orinoco; on twig of shrub; 10 Nov 2005, G.Neuwirth 8409 (B – holotype; ABL, hb. Neuwirth – isotypes).

DescripTion: Thallus ecorticate, pinkish grey, UV–. Ascomata 0.4–0.6 mm diam., centrum globose, erumpent, solitary. Ostiole apical. Hamathecium inspersed with very large oil droplets. Ascospores 8/ascus, IKI–, 7-septate, 30–33 × 9–10 μm, wall roughly ornamented. Chemistry: No substances detected.

noTe: The name is in honour of the collector of the type and only specimen, Gerhard Neuwirth.

Polymeridium ocellatum (Müll.Arg.) Aptroot comb. nov.

MycoBank: MB802997.

Polyblastia ocellata Müll.Arg., Flora 65: 402 (1882); Verrucaria ocellata Leight., Trans. Linn. Soc. London 35: 458 (1866) non Hoffm. (1790); Polyblastiopsis ocellata (Müll.Arg.) Zahlbr., Catal. Lich. Univ. 1: 351 (1922); Exiliseptum ocellatum (Müll.Arg.) R.C.Harris, Acta Amazon. (Suppl.) 14: 66 ('1984') [1986]. Typus: Brazil. aMazonas: São Gabriel; on tree bark; R. Spruce 244 (BM – isotype).

illusTraTions: Leighton, Trans. Linn. Soc. London 35: pl. 56, fig. 23 (1866); Harris, Acta Amazon. (Suppl.) 14: 78, fig. 6 ('1984') [1986].

DescripTion: Thallus greyish green (lichexanthone-colour), smooth, UV+ yellow. Perithecia pyriform, immersed in the bark, 2–4 fused with joint ostioles. Ostioles

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eccentric. Hamathecium not inspersed, IKI–, filaments profusely anastomosing. Ascospores 8/ascus, hyaline, IKI–, submuriform, 15–16 × 6–8 μm, fusiform, with constrictions.

DisTriBuTion: Neotropical (Brazil).

Polymeridium paraproponens Aptroot, M.Cáceres & E.L.Lima, Lichenologist 45: 546.

Typus: Brazil. ronDônia: Porto Velho, UNIR Federal University campus S of city, on bark of tree, c. 100 m alt., 8 Mar. 2012, M.E.S.Cáceres & A.Aptroot 11162 (ISE – holotype; ABL – isotype).


Neotropical (Brazil). A record from P. aff. proponens from Venezuela (Komposch & Hafellner 2006) probably belongs to this species.

Polymeridium pleiomerellum (Müll.Arg.) R.C.Harris, in Egan, Bryologist 90: 164 (1987)

Arthopyrenia pleiomerella Müll.Arg., Bot. Jahrb. Syst. 6: 406 (1885). Typus: cuBa. c.Wright, Verr. Cub. 107 (G – holotype, not seen).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole apical. Hamathecium inspersed with oil droplets. Ascospores IKI–, 7–11-septate, 25–36 × 5–9 μm. Chemistry: No substances detected.

DisTriBuTion: Probably pantropical: USA (Harris 1993), Mexico (Harris 1993), Jamaica (Harris 1993), Puerto Rico (Harris 1993), Cuba, Brazil (Harris 1993), Papua New Guinea (Aptroot et al. 1997).

Polymeridium pleurothecium R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 636 ('1991') [1993].

Typus: guyana. Mt Ayanganna, Potaro, G.Samuels 5085 (NY– holotype).

illusTraTions: Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 628, fig. 10–11 ('1991') [1993]; Aptroot, Lucking, Sipman, Umaña & Chaves, Biblioth. Lichenol. 97: 86, fig. 14B (2008).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole lateral. Hamathecium not inspersed, filaments profusely anastomosing. Ascospores 8/ascus, hyaline, IKI–, 9–13-septate, (35–)50–75 × 12–16 μm, not ornamented, wall not thickened. Chemistry: No substances detected.

DisTriBuTion: Pantropical: Costa Rica (Aptroot et al. 1997), Venezuela (Komposch & Hafellner 2006), Guyana, Australia (Aptroot 2009).

new counTry recorDs: papua new guinea. MaDang prov.: Foothills of Finisterre Range, A.Aptroot 33169, 1992 (ABL).

Polymeridium proponens (Nyl.) R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 637 ('1991') [1993]

Verrucaria proponens Nyl., Bull. Soc. Linn. Normandie, sér. 2, 2: 130 (1868); Campylothelium proponens (Nyl.) Müll.Arg. (1892); Polyblastiopsis proponens (Nyl.) Zahlbr., Catal. Lich. Univ. 1: 351 (1922). Typus: new caleDonia. loyalTy islanDs: Lifu, C.Thiébaut (H-Nyl – holotype, not seen).

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Verrucaria anoista Stirt., Proc. Phil. Soc. Glasgow 13: 191 (1881); Anthracothecium anoistum (Stirt.) Zahlbr., Catal. Lich. Univ. 1: 458 (1922); Campylothelium anoistum (Stirt.) Patw. & Makhija, Biovigyanam 7: 44 (1981). Typus: inDia. Assam, G.Watt (BM – lectotype, selected by Patw. & Makhija, Biovigyanam 7: 44, 1981).

Campylothelium album Müll.Arg., Bull. Soc. Roy. Bot. Belg. 30(1): 89 (1891). Typus. cosTa rica. punTarenas: Baia de Salinas, 1890, G.Pittier 5173 (G– holotype).

Campylothelium nitidum Müll.Arg., Nuovo Giorn. Bot. Ital. 23: 401 (1891). Typus: ausTralia. queenslanD: Brisbane, F.M.Bailey 460 (G – holotype).

illusTraTions: Patwardhan & Makhija, Biovigyanam 7: 45, fig. 1–2 (1981); Aptroot, Fl. Australia 57: 544, fig. 172A (2009).

DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia solitary. Ostiole lateral. Hamathecium not inspersed, filaments profusely anastomosing. Ascospores 8/ascus, IKI+ violet, muriform, 35–54 × 12–19 μm, with partly oblique septa, not ornamented, wall not thickened. Chemistry: lichexanthone.

DisTriBuTion: Pantropical: Hawai’i, USA (Harris 1993), Mexico, Puerto Rico (Harris 1993), Costa Rica (Aptroot et al. 2008), Guyana (Harris 1993), Venezuela (Harris 1993), Brazil (Harris 1993), Canary Isles (Tenerife, leg. Etayo 12417, det. Aptroot), India (Jagadeesh Ram et al. 2005), Papua New Guinea (Aptroot et al. 1997), Australia (Aptroot 2009). The species has recently often been reported in a wide sense, including P. amylosporum; older reports of P. amylosporum may refer to either this species or P. proponens).

Polymeridium pyrenastroides R.C.Harris ex Aptroot sp. nov. Figs 2A, 3H–J

MycoBank: MB802998.

Diagnosis: Like Polymeridium biloculare but thallus UV+ yellow.

Typus: venezuela. aMazonas: Alto Orinoco, 15 km W of Esmeralda, W bank of Surumoni; alt. 110 m; on Qualea sp. tree bark; 15 Feb 1997, J.Hafellner & H.Komposch 209-6-41 (GZU – holotype). Venezuela: Same locality as the holotype, J.Hafellner & H.Komposch 3380, 1998 (GZU – paratypes).

DescripTion: Thallus ecorticate, whitish grey, UV+ yellow. Ascomata 0.4–0.6 mm diam., triangular in surface view, erumpent, pyriform, mostly 2–5 fused with joint ostiole. Ostioles lateral, up to 0.6 mm long and often surfacing separated from the ascomata. Hamathecium not inspersed. Ascospores 8/ascus, IKI–, 3-septate, 28–32 × 9–11 μm, not ornamented, surrounded by a firm 2–3 μm thick layer of gel. Chemistry: lichexanthone.

noTe: The name refers to the similarity with Pyrenastrum because of the fused ostioles; the name was found on the herbarium specimen and in an accompanying manuscript made available by Josef Hafellner & Harald Komposch.

Polymeridium pyrenuloides (Müll.Arg.) Aptroot comb. nov.

MycoBank: MB803007.

Arthopyrenia pyrenuloides Müll.Arg., Mém. Soc. Phys. Genève 30(3): 27 (1888); Verrucaria pyrenuloides Fée, Essai Crypt. Exot., Suppl. Révis. (Paris): 86 (1837), non Ach. (1814); Arthopyrenia verrucarioides Zahlbr., Catal. Lich. Univ. 1: 312 (1921) nom. superfl. Typus: souTh aMerica. Tree bark of Cinchona, A.L.A.Fée s.n. (G – lectotype, selected by R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 639 ('1991') [1993], not seen).

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Arthopyrenia paulensis Zahlbr., Akad. Wiss. Wien, Math.-Naturwiss. Kl. Denkschr. 83: 89 (1909). Typus: Brazil. são paulo: S.Cruz, Bella Vista; on tree bark; R. von Wettstein & G.Schiffner s.n. (W – holotype, not seen).

DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia solitary. Ostiole apical. Hamathecium inspersed with oil droplets, filaments profusely anastomosing. Ascospores 8/ascus, IKI–, 3-septate, 17–20 × 5–7 μm, not ornamented, wall not thickened. Chemistry: lichexanthone.

DisTriBuTion: Probably pantropical: Brazil, Australia (Aptroot 2009, part of the specimens cited as P. subcinereum, e.g. J.A.Elix 37743).

Polymeridium quinqueseptatum (Nyl.) R.C.Harris, in Tucker & Harris, Bryologist 83: 12 (1980)

Verrucaria quinqueseptata Nyl., Expos. Syn. Pyrenocarp. 58 (1858); Arthopyrenia quinqueseptata (Nyl.) Müll. Arg., Flora 68: 326 (1885). Type: usa. souTh carolina: Ravenel (H-Nyl – holotype, not seen).

Verrucaria pustulosa Stirt., Proc. Roy. Philos. Soc. Glasgow 13: 191 (1881); Polyblastiopsis pustulosa (Stirt.) Zahlbr., Cat. Lich. Univ. 1: 351 (1922). Type: inDia. assaM: Tezpore, G.Watt (BM – isotype).

Arthopyrenia comparatula Müll.Arg., Bot. Jhrb. Syst. 6: 406 (1885). Type: cuBa. C.Wright, Verr. Cub. 152 (G – holotype, not seen).

Pseudopyrenula polyphragmia Vain., Bol. Soc. Broteriana sér. 2, 6: 178 (1930). Type: MozaMBique. Palma, A.Pires de Lima 303 (TUR-Vainio 34713 – lectotype, selected by R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 639 ('1991') [1993], not seen).

illusTraTions: McCarthy, Australas. Lichenol. Newslett. 26: 17, fig. 1B (1995); Aptroot, Lucking, Sipman, Umaña & Chaves, Biblioth. Lichenol. 97: 86, fig. 14D (2008).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole apical. Hamathecium inspersed, filaments profusely anastomosing. Ascospores 8/ascus, IKI–, 4–7-septate, 18–28 × 4–7 μm, not ornamented, wall not thickened. Chemistry: no substances detected.

DisTriBuTion: Pantropical: USA (Harris 1993), Jamaica (Harris 1993), Cuba, Costa Rica (Aptroot et al. 2008), Venezuela (Harris 1993), Brazil (Harris 1993), Mozambique, Seychelles (Seaward & Aptroot 2009), India, Thailand, Philippines (Harris 1993), Australia (Aptroot 2009).new counTry recorDs: neTherlanDs anTilles. sT. eusTaTius: Courtar Mountain, Quill, H.J.M.Sipman 56815b, 2008 (B).

Polymeridium refertum (Stirt.) Aptroot comb. nov.MycoBank: MB802999.

Trypethelium refertum Stirt., Proc. Roy. Phil. Soc. Glasgow 11: 321 ('1878') [1879]; Plagiotrema refertum (Stirt.) Makhija & Patw., J. Hattori Bot. Lab. 73: 209 (1993). Typus: inDia. Nilgiris; on tree bark; 1814, G.Watt s.n. (BM – isolectotype, selected by U.Makhija & P.G.Patwardhan, J. Hattori Bot. Lab. 73: 209, 1993).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole lateral. Hamathecium inspersed. Ascospores 3–6-septate, predominantly 4-septate, 22–26 × 7–9 μm. Chemistry: No substances detected.

DisTriBuTion: Asia (India).

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noTe: This species resembles P. quinqueseptatum but for the lateral ostiole.

Polymeridium siamense (Vain.) Aptroot comb. nov.MycoBank: MB803000.

Arthopyrenia siamensis Vain., Ann. Soc.Zool. Bot. Fenn. Vanamo 1(3): 55 (1921). Typus: ThailanD. chiang Mai: Chiang Mai, Doi Suthep; on tree bark; C.C.Hosseus s.n. (G – holotype).

Arthopyrenia obvelata Vain., Ann. Acad. Sci Fenn. Ser. A, 19(15): 11 (1923). Typus: philippines. luzon: Benguet prov., Pauai; on tree bark; P.McGregor s.n. (TUR – holotype).


DisTriBuTion: Australasian (Australia, Philippines, Thailand).new counTry recorD: guyana. rupununi DisTr.: Kuyuwini, M.Jansen-Jacobs et al. 2574, 1991 (ABL).

Polymeridium simulans R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 641 ('1991') [1993]. Typus: Brazil. aMazonas: 5 km E of Borba, Nelson 1310 (NY– holotype).


DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole lateral. Hamathecium not inspersed, filaments profusely anastomosing. Ascospores 8/ascus, IKI-, 3-septate, 30–35 × 12–13 μm, not ornamented, wall not thickened. Chemistry: Thallus without lichexanthone.

DisTriBuTion: Neotropical (Brazil).new counTry recorDs: guyana. poTaro-siparuni: Kaieteur Falls, P.DePriest, E.Chistenson & C.Kelloff 9343, 1994 (B). venezuela. Bolivár: Canaima National Park, alt. 600 m, F.Berger 18504, 2003 (ABL, hb. Berger).

Polymeridium stramineoatrum (Vain.) Aptroot comb. nov.MycoBank: MB803001.

Arthopyrenia stramineoatra Vain., Acta Soc. Fauna Flora Fenn. 7(2): 234 (1890). Typus: Brazil. Minas gerais: Carassa; on tree bark; E.A.Vainio, Lich. Bras. 1566 (TUR – holotype, not seen).

DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia solitary. Ostioles mostly eccentric. Hamathecium not inspersed. Ascospores IKI–, 3-septate, 24–30 × 7–10 μm. Chemistry: lichexanthone.

DisTriBuTion: Neotropical (Brazil).new counTry recorD: venezuela. aMazonas: Alto Orinoco, 15 km W of Esmeralda, alt. 110 m, J.Hafellner & H.Komposch 209-6-4, 1997 (GZU).

Polymeridium subcinereum (Nyl.) R.C.Harris, in Tucker & Harris, Bryologist 83: 12 (1980)

Verrucaria subcinerea Nyl., Expos. Syn. Pyrenocarp. 37 (1858); Pyrenula subcinerea (Nyl.) Tuck., Gener. Lich.: 273 (1872); Porina subcinerea (Nyl) Zahlbr., Catal. Lich. Univ. 1: 405 (1922). Typus: USA. Texas: C.Wright (H-Nyl – holotype, not seen).

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Arthopyrenia subimitans Müll.Arg., Bull. Soc. Roy. Bot. Belg. 32(1): 169 (1893). Typus: cosTa rica. punTarenas: Boruca, Tonduz, G.Pittier, Pl.Costaric. Exs. 6287 (BM – isotype).

Pseudopyrenula follmannii C.W.Dodge, Nova Hedwigia 12: 308 (1966). Typus: chile. aconcagua: Los Molles, G.Follmann 14852-D (FH-Dodge – holotype).

Massarina operculicola M.Morelet, Bull. Soc. Sci. Nat. Archéol. Toulon & Var 36: 14 (1980). Typus: france. Entrecasteaux, M.Morelet 1276 (herb. M.Morelet, Champenoux – holotype ['PFN', 'CNRF 898', 'INRA, Nancy']).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole apical. Hamathecium not inspersed with oil droplets, filaments profusely anastomosing. Ascospores 8/ascus, IKI–, 3-septate, (15–)16–20 × 5–7 μm, not ornamented, wall not thickened. Chemistry: no substances detected.

DisTriBuTion: Subcosmopolitan: USA (Harris 1993), Dominican Republic (Harris 1993), Costa Rica (Aptroot et al. 2008), Venezuela (Komposch & Hafellner 2006), Colombia (Sipman et al. 2008), Brazil (Harris 1993), Chile, France, Mozambique (Harris 1993), Philippines, Australia (Aptroot 2009, partly referring to other species; M.E. Hale 69141 from Lamington forest (US, ABL) is correct).

new counTry recorDs: guyana. rupununi DisTr.: Monkey Pod Landing, SW of Mt. Makarapan, P.J.M.Maas et al. 7373c, 1988 (ABL). papua new guinea. MaDang prov.: Jais Aben Resort, A.Aptroot 38248, 1995 (ABL).

Polymeridium submuriforme Aptroot sp. nov. Figs 2B, 3K, 4F

MycoBank: MB803002.

Diagnosis: Like Polymeridium cinereonigricans but ascospores much smaller and hamathecium inspersed.

Typus: philippines. leyTe: Leyte, between Camp Pandong Bato and Mt. Agipo; on tree bark; 17 Aug 2000, F.Schumm & S.Schwarz 774 (B – holotype).

DescripTion: Thallus ecorticate, pinkish grey, UV–. Ascomata 0.3–0.6 mm diam., centrum globose, erumpent, solitary, hemisphaerical-conical with round top, wall expanding up to 0.2 mm sideways in older ascomata. Ostiole apical. Hamathecium inspersed. Ascospores 6–8/ascus, IKI–, submuriform, 4–5 × 0–1-septate with partly oblique septa, 18–20 × 6–7.5 μm, not ornamented, surrounded by a 1–2 μm wide gelatinous sheath. Chemistry: No substances.

Polymeridium subvirescens (Leight.) Aptroot comb. nov.

MycoBank: MB803003.

Verrucaria subvirescens Leight., Trans. Linn. Soc. London 25: 488 (1866); Porina subvirescens (Leight.) Zahlbr., Cat. Lich. Univ. 1: 406 (1922). Typus: Brazil. aMazonas: Santarém; on tree bark; R.Spruce, Lich. Amaz. And. 358 (BM – lectotype, selected by R.C.Harris, Acta Amazon. (Suppl.) 14: 70 ('1984') [1986]).

DescripTion: Thallus ecorticate, white, UV+ yellow. Perithecia solitary. Ostiole apical or partly lateral. Hamathecium inspersed with oil droplets. Ascospores IKI–, 3-septate, 24–33 × 6–10 μm. Chemistry: lichexanthone.

DisTriBuTion: Probably pantropical: Brazil, Australia (Aptroot 2009, as P. catapastum).

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Polymeridium suffusum (Knight) Aptroot comb. nov.

MycoBank: MB803004.

Verrucaria suffusa Knight, Trans. Proc. New Zealand Inst. 15: 356 (1883); Arthopyrenia suffusa (Knight) Müll.Arg., Bull. Soc. Roy. Bot. Belgique 31(2): 40 (1894). Typus: new zealanD. norTh islanD: Wellington; on tree bark; 20 Jan 1882, C.Knight s.n. (G – isotype).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole apical or partly lateral. Hamathecium inspersed with oil droplets. Ascospores IKI–, 3-septate, 24–30 × 6–9 μm. Chemistry: No substances detected.

DisTriBuTion: Pantropical: Cuba (Harris 1993), Sarawak (Harris 1993), New Zealand.

new counTry recorDs: puerTo rico. DisTr. ponce: Caribbean National Forest, alt. 1150 m, A. & M.Aptroot 25625, 1989 (ABL). cosTa rica. prov. hereDia: Montaña Azul, alt. 1530 m, F.Berger 14078, 2000 (ABL, hb. Berger). reunion. Bourg-Murat, Col de Bellevue, alt. 1617 m, F.Schumm 15158, 2009 (Hb.Schumm).

Polymeridium sulphurescens (Müll.Arg.) R.C.Harris, Lichenogr. Thomsoniana 147 (1998).

Arthopyrenia sulphurescens Müll.Arg, Flora 65: 518 (1882); Pseudopyrenula sulphurescens (Müll.Arg.) Müll.Arg., Flora 66: 249 (1883). Typus: ausTralia. queenslanD: Toowoomba, C.H.Hartmann (G – holotype).

Arthopyrenia oculata Müll.Arg., Rep. Australas. Assoc. Advancem. Sci. 1895: 451 (1895); Polymeri-dium oculatum (Müll.Arg.) R.C.Harris, Bol. Mus. Paraense Emílio Goeldi, Ser. Bot., 7: 635 ('1991') [1993]. Type: ausTralia. queenslanD: C.Knight 136 (G – holotype).

Arthopyrenia suboculata Müll.Arg., Bull. Herb. Boissier 3: 325 (1895). Type: ausTralia. queenslanD: C.Knight 199 (G – holotype).

illusTraTion: Aptroot, Fl. Australia 57: 544, fig. 172B (2009).

DescripTion: Thallus ecorticate, white, UV–. Perithecia solitary. Ostiole apical to partly skewed, usually surrounded by a ring. Hamathecium not inspersed. Ascospores IKI–, 3-septate, 19–27 × 6.5–11 μm, generally surrounded by a 2–4 μm thick gelatinous sheath. Chemistry: No substances.

DisTriBuTion: Australasian: Papua New Guinea (Aptroot et al. 1997), Australia; rarely also on rock.

Polymeridium tribulationis Aptroot sp. nov. Figs 2C, 3L, 4C

MycoBank: MB803005.

Diagnosis: Like Polymeridium sulphurescens but hamathecium inspersed.

Typus: ausTralia. queenslanD: Cape Tribulation, 2 km of main road between Oil Palms and Coopers Creek; on tree bark; 4 Jul 1983, M.E.Hale 641717 (CANB – holotype; US, ABL – isotype).

DescripTion: Thallus ecorticate, yellowish white, UV–. Ascomata 0.4–0.6 mm diam., centrum globose, erumpent, solitary, superficial, mostly covered by thallus. Ostiole apical, usually surrounded by a ring. Hamathecium inspersed with oil droplets. Ascospores 8/ascus, IKI–, 3-septate, 20–24 × 7–10 μm, not ornamented. Chemistry: no substances.

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Polymeridium xanthoreagens Aptroot sp. nov. Figs 2D, 3M, 4H

MycoBank: MB803006.

Diagnosis: Like Polymeridium sulphurescens but thallus UV+ yellow.

Typus: ausTralia. queenslanD: E of Mareeba, Davies Creek Road 17 km S of Kennedy Highway; on tree bark; 7 Jul 1983, M.E.Hale 64168 (CANB – holotype; US,ABL – isotype).

DescripTion: Thallus ecorticate, yellowish white, UV+ yellow. Ascomata 0.4–0.6 mm diam., centrum globose, erumpent, solitary, superficial, mostly covered by thallus. Ostiole apical, partly eccentric, usually surrounded by a ring. Hamathecium not inspersed. Ascospores 8/ascus, IKI–, 3-septate, 17–25 × 7–11 μm, not ornamented, generally surrounded by a 2–5 μm thick gelatinous sheath. Chemistry: lichexanthone.

noTe: Specimen M.E.Hale 69187 was reported as P. sulphurescens by Aptroot (2009).

aDDiTional speciMens exaMineD: ausTralia. queenslanD: S of Atherton, Herberton Range, Plath Road logging head, 9 km W on Plath Road, off Kennedy Highway; on tree bark; 1983, M.E.Hale 64178 (ABL, US); Mt. Windsor logging area, NW of Mossman; on tree bark; 1983, M.E.Hale 69187 (ABL, US).

Acknowledgements

The curators of the herbaria mentioned are thanked for their cooperation, and many colleagues for sending identified and unidentified material over the years. Dr. U.Makhija is especially thanked for placing some isotypes at my disposal. The CNPq – Conselho Nacional de Desenvolvimento Científico e Tecnológico is thanked for a research grant to MESC (Processo 501633/2009-0) and for financial support to the collecting trips by MESC (CNPq-PPBio Semiárido Processo 558317/2009-0). The Stichting Hugo de Vries-fonds is thanked for travel support to AA. Leo Spier is thanked for performing thin-layer chromatography.

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Manuscript submitted January 11, 2013; accepted March 20, 2013.

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