1363 Lich 44.4 03 Aptroot 449..456

The Lichenologist 44(4): 449–456 (2012) 6 British Lichen Society, 2012 doi:10.1017/S0024282912000084

A new terricolous Trapelia and a new Trapeliopsis (Trapeliaceae, Baeomycetales) from Macaronesia

Andre´ APTROOT and Felix SCHUMM

Abstract: Trapeliopsis gymnidiata, a terricolous species from the Canary Islands, is described as new to science. It has previously been confused with T. wallrothii, but differs by the soft, partly decorticate isidia, or rather gymnidia, that leave inconspicuous scars after falling off. Trapeliopsis wallrothii is known only from high mountains in Macaronesia, while T. gymnidiata is a lowland species. Trapelia rubra from Madeira is also described as new to science. It grows in similar places and resembles some morphs of Trapelia coarctata, but differs by the nearly squamulose areoles, the crenulate, seemingly sorediate, apothecium margin that does not become excluded, and especially by the patchy red colour (skyrin) in the medulla. Key words: Canary Islands, Gomera, Gran Canaria, La Palma, lichen, Madeira, new species, Portugal, Spain, Tenerife Accepted for publication 31 January 2012

Introduction have been combined in Trapelia at one time (Wirth 1987). However, recent DNA inves- Trapelia and Trapeliopsis were among the first tigations (Lumbsch et al. 2007) have sup- genera to be separated from, or resurrected ported the view that both are distinct genera to accommodate, species of Lecidea, which (Lumbsch et al. 2007). until the 1970s encompassed species that The taxonomy of these species complexes are now classified in over 30 different genera remains essentially unchanged in Europe in about 20 families in three different orders. since 1984, as can be seen from the key in In a classic paper, Coppins & James (1984) Smith et al. (2009) which is virtually identi- transferred some further species, notably cal, and is still complete for most of Europe. crustose taxa, to Trapeliopsis that until that However, while preparing a second volume time contained only squamulose forms, and of Schumm (2008), it became apparent that described some species that were widely two different, probably not even closely re- known under informal names such as ‘‘Tra- lated, species are currently called Trapeliopsis pelia spec.’’ or ‘‘Lecidea granulosa with orange wallrothii (Flo¨rke in Spreng.) Hertel & Gotth. pigment’’, and presented a key to the British Schneid. in Macaronesia. They differ, for species of the two genera. They also clarified example, in the type of isidia. Trapeliopsis the distinction between Trapelia coarctata wallrothii s. str. has firm, often solitary, corti- (Turner ex Sm.) M. Choisy and T. glebulosa cate isidia, as illustrated, for example, by (Sm.) J. R. Laundon, a matter that has been Coppins & James (1984) and Wirth (1995). much debated before and since as the species These isidia can be hand-sectioned with a seems to exhibit discrete variations, but their razor blade and after falling off leave a con- classification still stands firm. The separation spicuous scar. The other species has softer, of Trapeliopsis from Trapelia was repeatedly more clustered, partly decorticated isidia, questioned, and nearly all Trapeliopsis species which are often called gymnidia and are rather common in the Peltigerales and Phys- A. Aptroot: ABL Herbarium, G.v.d.Veenstraat 107, ciaceae; these cannot be hand-sectioned with- NL-3762 XK Soest, The Netherlands. out crumbling. Both occur in Macaronesia, Email: [email protected]. F. Schumm: Mozartstr. 9, D-73117 Wangen, Germany. while one of them is more widely distributed

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occurring also in Europe. Trapeliopsis wallro- Typus: Portugal, Madeira, Camacha, Valle Paraiso, thii has previously erroneously been reported 36170 N, 3250 E, on soil between basalt of footpath, 750 m alt., 3 June 2011, A. Aptroot 69725 (BR—holoty- from America, where currently three similar, pus; ABL—isotypus). but different species are thought to occur (McCune et al. 2002). No species of Trapelia (Fig. 1) or Trapeliopsis have previously been described from Macaronesia, but the occurrence of Thallus consisting of tightly appressed nodu- several widespread species has been noted. lose areoles, whitish, corticate, slightly glossy, There are three Lecidea species described mostly heavily pruinose, grey where not prui- from Macaronesia that are not accounted nose. Areoles roundish to incised, convex, c. for in recent publications (Tavares 1952), 1 mm diam., but partly incised and mostly viz. Lecidea azorica Nyl. from the Azores, aggregated to form thallus patches of 05– which is according to the description possi- 20 cm diam. Medulla partly bright red. Algae bly a Porpidia, Lecidea homosemoides Harm. green, c. 10 mm diam. from the Canary Isles which is a maritime Apothecia usually present, sessile, cupular, saxicolous species, and Lecidea conglobata up to 1 mm diam., circular, thalline margin (Hepp) Zahlbr. which is an illegitimate hom- white, crenulate with irregular rough sur- onym. None of these are identical with the face; disc concave, pink, often white pruinose. species treated here. The additional species Apothecium initials show as roughened whit- of Trapeliopsis from Macaronesia is therefore ish areas on the thallus, somewhat resembling described below as new, as well as an unde- soredia or the scars left by fallen-off isidia scribed Trapelia species with red medulla in Trapeliopsis wallrothii. Hymenium pink; that occurs in the same region. hypothecium brown, partly bright red. Asci Trapelia-type. Ascospores 8 per ascus, hyaline, ellipsoid, 18–23 9–12 mm. Pycnidia not observed. Material and Methods Secondary chemistry.ThallusC-- ,K-- ,KC-- , Identiﬁcation and descriptive work was carried out in Soest using an Olympus SZX7 stereomicroscope and P-- . Anthraquinone (skyrin, TLC) in red an Olympus BX50 compound microscope with inter- parts of the medulla and hypothecium, which ference contrast, connected to a Nikon Coolpix digital react K+ purple. camera, and in Wangen using a Wild M3 stereomicros- kope, an Olympus BX51 compound microscope with interference contrast, a Canon EOS 40D camera with Ecology and distribution. On compacted MP-E 65 mm and a Mic HM 560 cryotome. Sections soil, for example between rocks on footpath, were mounted in tap water or iodine (I or IKI), or en- at lowland to mid elevations. Known from hanced with lactophenol Cotton Blue (LCB). When Madeira and the Canary Islands. nothing is indicated, tap water was used. All measure- ments were made on material mounted in water. The specimens are deposited in ABL, B, BR, L, and hb. Discussion. This species resembles some Schumm. TLC was carried out following the protocol morphs of Trapelia coarctata, but differs by of Orange et al. (2001), with all three standard solvents the nearly squamulose areoles, the crenulate, and observation in UV before and after charring with seemingly sorediate, apothecium margin that sulphuric acid. does not become excluded, and especially by the patchy red colour (skyrin) in the medulla. Moreover, it occurs only on compacted soil, The Species while Trapelia coarctata also, and even mainly, occurs on other substrata such as rock and Trapelia rubra Aptroot & Schumm sp. timber. nov. Additional specimens examined. Spain: Canary Islands: MycoBank No: MB 564322 Gran Canaria, 8 km W. of Teror, near Pinos de Galdar, A crustose, nearly squamulose, terricolous Trapelia with 28030 N, 15370 W, 1295 m alt., 1999, F. Schumm 5314 crenate and seemingly sorediate apothecia and skyrin. (hb. Schumm), 5304 (ABL).

Fig.1. Trapelia rubra (holotype). A, habitus; B, section through apothecium with margin; C, ascus (dark areas I+ reddish); D, ascospores. Scales: A ¼ 1 mm; B ¼ 100 mm; C & D ¼ 10 mm.

Trapeliopsis gymnidiata Aptroot & T. wallrothii can be hand-sectioned with a Schumm sp. nov. razor blade and leave a conspicuous, well delimited scar when falling off. Both species MycoBank No: MB 564323 occur in Macaronesia, where true T. wallrothii A crustose terricolous Trapeliopsis with mostly non- is only known from high mountains, while T. corticate isidia, large apothecia and gyrophoric acid. Typus: Spain, Canary Islands, Gran Canaria, Em- gymnidiata occurs at lower elevations. balse del Mulato, 275490 N154160 W, 760 m alt., 31 Additional specimens examined. Spain: Canary Islands: December 1998, F. Schumm 5390 (B—holotypus), 5401 Gran Canaria, Pinar de Tamadaba west of Artenara (ABL—isotypus), 5369, 5398, 5370 (hb. F. Schumm— 28020 N, 15410 W, 1094 m alt., 2000, F. Schumm isotypi). 8129 (hb. Schumm); Gran Canaria, 3 km north-west of Artenara, Embalse de Lugarejos, 28020 N, 154050 W, (Figs 2 & 3A–D) 340 m alt., 1999, F. Schumm 5113 (hb. Schumm); La Palma, Los Cancajos south of Santa Cruz, 283870 N, Thallus consisting of tightly appressed 17460 W, 30 m alt., 2007, F. Schumm 12987 (hb. areoles that form a thick crust and can be Schumm); La Gomera, Los Barranquillos north-east of 0 0 slightly radially arranged along the margins, Arure, 28 091 N, 17 183 W, 1020 m alt., 2003, F. Schumm 10430 (hb. Schumm); Tenerife, Ladera de pinkish to lead grey, corticate, dull. Areoles Guı¨mar, 400 m alt., 1983, A. Aptroot 11568 (ABL). angular, a little incised, flat to convex along the margins, c. 1 mm diam., aggregated to Trapeliopsis wallrothii (Flo¨rke) form thallus patches of 05–5 cm diam. Hertel & Gotth. Schneid. Isidioid propagules or gymnidia (mostly non- corticate isidia) are formed in defined sora- in Schneider, Biblioth. Lichenol. 13: 153 (1980).—Lecidea wallrothii Flo¨rke, in Sprengel, Neue Entdeckungen im gan- lium-like places, c. 1 mm diam, soft, un- zen Umfang der Pflanzenkunde 2: 96 (1820); type: Ger- branched or gnarled, whitish grey, usually many, Halle, Kro¨llwitz, Wallroth (L0833383—isotype). flattened and at most corticate at one side, c. Lecidea salweii Borrer, in Hooker, Supplement to 02 03–05 mm, easily detached, leaving English Botany 3: 2861 (1841), also as ‘‘salweei’’ or ‘‘salweyi’’; type: Great Britain, n. v. (see Laundon & inconspicuous, not clearly delimited, whitish Waterfield 2007: 386) scars. Medulla whitish. Algae green, c. 10 mm diam. (Figs 3E & F, 4) Apothecia usually present, sessile, up to See Smith et al. (2009) for description. 2 mm diam., margin often flexuose and un- Ecology and distribution. On compacted dulating, without thalline margin; disc convex, soil, in Macaronesia and S Europe at higher brownish grey, margin lead grey, persistent, elevations, in the rest of Europe at lowland margin closely appressed to the thallus. Hy- to mid elevations. Known from Macaronesia menium hyaline; epihymenium golden brown; and Europe. subhymenium hyaline; hypothecium with pinkish crystals. Asci Trapelia-type. Ascospores 8per Notes on author citation and typification.As ascus, hyaline, ellipsoid, 80–110 45–55 can be seen from the original paper, the orig- mm. inal description is doubtlessly provided by Pycnidia not observed. Flo¨rke; Sprengel was only the editor of the journal, in which various papers appeared Secondary chemistry. Thallus C+ fleetingly (hence the title: ‘‘ganzen Umfang der Pflan- pink, K-- ,KC-- ,P-- . Gyrophoric acid (TLC). zenkunde’’). The author should therefore be Ecology and distribution. On compacted soil, Flo¨rke in (not ex) Spreng., and ‘‘in Spreng.’’ at lowland to mid elevations. Known only can be left out at will. Schneider (1980) cites from the Canary Islands. a collection from H. N. Bolander, originat- ing from North America, as type. Bolander Discussion. This species strongly resembles was born long after the original description T. wallrothii, but has softer, more clustered, of the species. As no indication is given by partly decorticated isidia or rather gymnidia, Schneider (1980) whether or not original which cannot be hand-sectioned without material still exists, it can be seen as a mis- crumbling. The distinctly corticate isidia of take rather than a neotypification. Coppins

Fig.2.Trapeliopsis gymnidiata. A, habitus with gymnidia; B & C, habitus also with apothecia; D, gymnidia; E, section through gymnidium (dark areas LCB+ blue). (B, D & E, holotype; A & C, Schumm 10430). Scales: A ¼ 5 mm; B ¼ 2 mm; C & D ¼ 1 mm; E ¼ 20 mm.

Fig.3.A–D,Trapeliopsis gymnidiata (holotype). A, section through gymnidium (dark areas LCB+ blue); B, section through apothecium with margin (dark areas LCB+ blue); C, ascus (dark areas IKI+ bluish); D, ascospores. E & F, Trapeliopsis wallrothii (isotype in L). E, label of the packet; F, habitus with scars where isidia have fallen off. Scales: A ¼ 200 mm; B ¼ 100 mm; C & D ¼ 5 mm; F ¼ 1 mm.

Fig.4. Trapeliopsis wallrothii (Schumm 8484). A, habitus with globose isidia; B, habitus also with apothecium; C, section through thallus with attached isidium; D, section through apothecium with margin (dark areas LCB+ blue); E, ascus (dark areas I+ reddish); F, ascospores. Scales: A ¼ 2 mm; B ¼ 1 mm; C & D ¼ 100 mm; E & F ¼ 5 mm.

& James (1984) suggest that in case the References original material proves untraceable, a speci- Coppins, B. J. & James, P. W. (1984) New or interesting men from the contemporary G. F. W. Meyer British lichens V. Lichenologist 16: 241–264. could be selected as neotype, but they re- Laundon, J. R. & Waterfield, A. (2007) William Borrer’s lichens in the Supplement to the English Botany 1829– frained from neotypification. In the Koerber 1866. Botanical Journal of the Linnean Society 154: collection (Stammherbar) in Leyden, a spe- 381–392. cimen (L0833383, Fig. 3E & F) is present Lumbsch, H. T., Schmitt, I., Mangold, A. & Wedin, M. which was collected by Wallroth and labelled (2007) Ascus types are phylogenetically misleading in Trapeliaceae and Agyriaceae (Ostropomycetidae, with the unpublished name ‘‘Lecanora coarc- Ascomycota). Mycological Research 111: 1133–1141. tata v[ar]. macrocymatia Wallr.’’ It was appar- McCune, B., Camacho, F. & Ponzetti, J. (2002) Three ently sent to Flo¨rke, who described it as new species of Trapeliopsis on soil in Western North ‘‘Lecidea wallrothii Flk.’’ The species has thus America. Bryologist 105: 78–85. been named after the collector, Wallroth. Orange, A., James, P. W. & White, F. J. (2001) Micro- chemical Methods for the Identification of Lichens. Subsequently, the type collection was divided London: British Lichen Society. and distributed over several herbaria, which Schneider, G. (1980) [‘1979’] Die Flechtengattung Psora was common practice at this time. The speci- sensu Zahbruckner. Bibliotheca Lichenologica 13: 1– men in Leyden is certainly part of this type 291. Schumm, F. (2008) Flechten Madeiras, der Kanaren und specimen, even though it does not bear a Azoren. Wangen: published by the author. collecting locality or date. In the original Smith, C. W., Aptroot, A., Coppins, B. J., Fletcher, A., publication, only the occurrence of this spe- Gilbert, O. L., James, P. W. & Wolseley, P. A. (eds) cies in Kro¨llwitz (near Halle in Germany) is (2009) The Lichens of Great Britain and Ireland. Lon- mentioned, which should thus be the type don: British Lichen Society. Tavares, C. N. (1952) Contribution to the lichen flora of locality. Macaronesia I. Lichens from Madeira. Portugaliae Details of illustrated specimen. Portugal: Madeira: Pau´l Acta Biologia, ser. B, 3(3): 308–391. da Serra, near Urze, 324530 N, 170650 W, 1400 m Wirth, V. (1987) Die Flechten Baden-Wu¨rttembergs. Stutt- alt., 2001, F. Schumm 8484 (hb. Schumm). gart: Ulmer. Wirth, V. (1995) Die Flechten Baden-Wu¨rttembergs. Ed. 2. Pat Wolseley is warmly thanked for sending photocopies Stuttgart: Ulmer. of the description of Lecidea salweii. The collection manager in Leyden, Gerard Thijsse, is warmly thanked for his prompt sending of specimens from that herbarium.