Opuscula Philolichenum, 18: 390-395. 2019. *pdf effectively published online 4October2019 via (http://sweetgum.nybg.org/philolichenum/)

Notes on the (, ) in South America: Two new species from Peru, and a new combination

JOHN A. ELIX1

ABSTRACT. – Tetramelas peruviensis Elix and T. weberianus Elix are described as new to science from high altitudes in Peru. The new combination, T. coquimbensis (C.W. Dodge) Elix is proposed for coquimbensis and a key to the eight species of Tetramelas present in South America is given.

KEYWORDS. – Biodiversity, fungal systematics, .

INTRODUCTION

Phylogenetic studies have confirmed that Tetramelas Norman constitutes a well-founded, monophyletic segregate of Buellia sens. lat. (Helms et al. 2003, Nordin 2004, Nordin & Tibell 2005). Generic characters include the presence of polychloroxanthones (arthothelin, 6-O-methylarthothelin, isoarthothelin or 2,5,7-trichloro-3-O-methylnorlichexanthone), commonly curved, 1–3 septate ascospores with pointed apices that show - or Callispora-type thickenings in early ontogeny (Giralt et al. 2009), bacilliform conidia and a predominantly Arctic-Antarctic or alpine-subalpine distribution (Kalb 2004, Elix 2018). Up to the present five species have been reported from South America, T. anisomerus (Vain.) Elix from Argentina and (Calvelo & Fryday 2006, Calvelo & Liberatore 2002, Galloway & Quilhot 1998), T. fuegiensis Elix, H. Mayrhofer & J.M. Rodr. from Argentina (Elix et al. 2018), T. pulverulentus (Anzi) A. Nordin & Tibell from Argentina (Nordin 2000), T. regiomontanus Marbach from Chile, Colombia and Venezuela (Kalb 2004, Marbach 2000) and T. thiopolizus (Nyl.) Giralt & Clerc from Venezuela (Giralt & Clerc 2011). In this paper I describe two new species of Tetramelas from high elevation in Peru and propose a new combination for the genus. A key to the eight species of Tetramelas present in South America is given.

MATERIALS AND METHODS

Observations and measurements of thallus and apothecium anatomy, asci, ascospores and conidia were made on hand-cut sections mounted in water and treated with 10% potassium hydroxide (K). Asci were also observed in Lugol’s Iodine (I), with and without pre-treatment in K. Medullary sections were treated with 10% sulfuric acid (H2SO4) and apothecial sections with 50% nitric acid (N). Ascospore measurements are presented below in the following format: smallest measurement–mean–largest measurement and are based on 50 observations. Chemical constituents were identified by thin-layer chromatography (TLC), high- performance liquid chromatography (HPLC, Elix 2014) and comparison with authentic samples.

TAXONOMIC SECTION

Tetramelas coquimbensis (C.W.Dodge) Elix, comb. nov. MYCOBANK #831053

1JOHN A. ELIX – Research School of Chemistry, Building 137, Australian National University, Canberra, A.C.T. 2601, Australia – e-mail: [email protected]

390 ≡ Buellia coquimbensis C.W.Dodge, Nova Hedwigia 12: 349. 1967[1966]. TYPE: CHILE. . ELQUI : La Serena, 1965, on coastal rocks, G. Follmann 14765, (FH!, holotype).

DISCUSSION. – The morphological and chemical features of this species are consistent with it belonging to Tetramelas. The ascospores exhibit characteristic Physconia- or Callispora-type wall thickenings in early ontogeny, very similar to those of T. peruviensis (Fig. 1B). Furthermore, this species has short bacilliform conidia and contains polychloroxanthones (Bungartz et al. 2016). Ideally it would have been more satisfactory to have molecular analyses to confirm the generic placement of this species (as well as the two species described below), but unfortunately only aged specimens were available. This species contains 2,5,7-trichloro-3-O-methylnorlichexanthone (major) and isoarthothelin (minor) rather than 6-O- methylarthothelin and arthothelin as reported previously (Bungartz et al. 2016).

Specimens examined. – CHILE. , ELQUI PROVINCE: La Serena, 1965, on coastal rocks, G. Follmann 16228 (KASSEl, from the type locality and erroneously labelled “type”); Pichidangui, coastal headlands just N of village, 10 m, 14.xi.1976, on boulders lying at foot of rocky tor, W.A. Weber & B. Johnston s.n. = Lichenes Exsiccati COLO No. 546 (CANB, distributed as Buellia flavoareolata (Nyl.) Müll. Arg.).

Tetramelas peruviensis Elix, sp. nov. MYCOBANK #830138 FIGURE 1.

Similar to Tetramelas nelsonii but differs in having broader ascospores, 7–12 µm wide, and in containing isoarthothelin and medullary calcium oxalate.

TYPE: PERU. DEPT. DE ICA: western foothills of the Andes 36–40 km E of Nazca, on road to Puquio, alt. 2000–2500 m, steep slope and ridgetop with large boulders of rhyolite-tufa, in Bursera- Neoraimondia association, 25.iii.1978, , on rock, W.A. Weber & J. Kohn s.n. = Lichenes Exsiccati COLO No. 582 (CANB!, holotype; CANB!, GZU!, isotypes).

DESCRIPTION. – Thallus areolate to squamulose or sublobate, sometimes forming rosettes to 20 mm wide and 1.2 mm thick; areoles rounded, 0.5–3 mm wide, separate to crowded and contiguous, sublobate at margins; upper surface mustard-yellow to orange-brown, dull, smooth, convex, epruinose; prothallus not apparent; photobiont cells 7–19 µm wide; medulla white, containing calcium oxalate (H2SO4+), I–. Apothecia 0.5–1.2 mm wide, lecideine, separate and ±round, broadly adnate to sessile; disc black, epruinose, plane to markedly convex with age. Proper exciple distinct but excluded in older, convex apothecia, in section 30–40 µm thick; outer part aeruginose-dark brown, K−, N+ purple-brown, brown within. Hypothecium 200−250 µm thick, dark brown to deep red-brown, K+ orange to pale brown solution, N+ orange-brown. Epihymenium 15–25 µm thick, dark olive-brown to aeruginose-black, K+ blue-green, N+ purple-brown. 75–85 µm thick, colourless, not inspersed but with some paraphyses 3–6 µm wide, containing oil droplets. Subhymenium 20−30 µm thick, pale brown. Paraphyses 1–2 µm wide, simple to sparsely branched, with apices 3.5–5 µm wide and aeruginose caps. Asci of the Bacidia-type, with 8 or fewer . Ascospores initially of the Callispora- or Physconia-types, then of the Buellia-type, 1-septate, brown to dark brown, ellipsoid to broadly fusiform, 12–[16.4]–22 × 7–[8.6]–12 µm, sometimes becoming constricted at the septum, rarely curved; outer -wall finely ornamented (microrugulate). Pycnidia immersed, punctiform; conidia bacilliform to elongate-ellipsoid, 5–6.5 × 1–1.5 µm

Chemistry. – 2,5,7-trichloro-3-O-methylnorlichexanthone (major), isoarthothelin (minor or trace), ±atranorin (minor), 2,5-dichloronorlichexanthone (trace) or isoarthothelin (major), ±atranorin (minor), 2,5- dichloronorlichexanthone (trace). Spot tests: thallus K+ yellow, C+ orange, KC+ deep orange, P–, UV+ orange.

ETYMOLOGY. – The species is named after its occurrence in Peru.

391

Figure 1. Tretramelas peruviensis (from the holotype). A, thallus and apothecia. B, line drawing of ascospore ontogeny. Scales = 1.0 mm in A and 10µm in B.

ECOLOGY AND DISTRIBUTION. – At present, the new species is known only from the type locality. Associated species included an unidentified Caloplaca species and Tetramelas weberianus, described below.

DISCUSSION. – In many respects the new species resembles T. nelsonii (Darb.) Elix, which is known from the South Shetland Islands and continental Antarctica (Lamb 1968; Øvstedal & Lewis Smith 2001; Elix 2018). Both are characterized by the presence of an aeruginose, N+ purple epihymenium and similar- sized ascospores and conidia and similar apothecial anatomy including similar reactions of the hypothecium. However, T. nelsonii contains 6-O-methylarthothelin and 4,5-dichloro-6-O-methylnorlichexanthone, lacks medullary calcium oxalate (H2SO4–) and has narrower ascospores, 6–8 µm wide, that sometimes develop endo-septa, then appearing 2- or 3-septate (Lamb 1968). In addition, some specimens of T. nelsonii have a weakly amyloid medulla. The new species resembles T. coquimbensis, a superficially similar species from coastal Chile (Bungartz et al. 2016, Dodge 1967). Both lichens are characterized by the presence of an

392

Figure 1. Tretramelas weberianus (from the holotype). A, thallus and apothecia. B, line drawing of ascospore ontogeny. Scales = 1.0 mm in A and 10µm in B.

aeruginose, N+ purple-brown epihymenium, 1-septate ascospores, bacilliform conidia and similar apothecial anatomy including the reactions of the hypothecium and epihymenium, as well as identical chemistry. However, T. coquimbensis has smaller ascospores, 12–[14.4]–15 × 5–[7.8]–9 µm, a compact, black prothallus and lacks oil paraphyses in the hymenium.

393 Tetramelas weberianus Elix, sp. nov. MYCOBANK #830139 FIGURE 2.

Similar to Tetramelas granulosus, but differs in having an aeruginose, N+ purple-brown epihymenium, a non-amyloid medulla that contains calcium oxalate and in containing isoarthothelin rather than 6-O-methylarthothelin.

TYPE: PERU. DEPT. DE ICA: western foothills of the Andes 36–40 km E of Nazca, on road to Puquio; alt. 2000–2500 m, steep slope and ridgetop with large boulders of rhyolite-tufa, in Bursera- Neoraimondia association, 25.iii.1978, on rock, W.A. Weber & J. Kohn s.n. = Lichenes Exsiccati COLO No. 581 (CANB!, holotype; CANB!, GZU!, isotypes).

DESCRIPTION. – Thallus areolate to subsquamulose, forming rosettes to 20 mm wide and 1.2 mm thick; areoles irregular, angular, 0.2–1 mm wide, separate to crowded and contiguous; marginal areoles flattened, sublobate, to 2 mm wide; upper surface mustard yellow to yellow-brown, dull, smooth, pruinose; prothallus not apparent; photobiont cells 7–19 µm wide; medulla white, containing calcium oxalate (H2SO4+), I–. Apothecia 0.3–1.3 mm wide, lecideine, separate and ±round, sessile; disc black, epruinose, plane to markedly convex with age. Proper exciple distinct but excluded in older, convex apothecia, in section 50–80 µm thick; outer part aeruginose-black, K−, N+ purple-brown, dark brown within. Hypothecium 200−250 µm thick, dark brown to brown-black, K−. Epihymenium 15–20 µm thick, dark olive-brown to aeruginose-black, K+ blue-green, N+ purple-brown. Hymenium 75–85 µm thick, colourless, not inspersed. Subhymenium 20−25 µm thick, pale to mid-brown; paraphyses 1–2 µm wide, simple to sparsely branched, with apices 5–6 µm wide and dark aeruginose caps. Asci of the Bacidia-type, with 8 or fewer spores. Ascospores initially of the Callispora- or Physconia-types, then of the Buellia-type, 1-septate or often becoming 3-septate, brown, ellipsoid to broadly fusiform or oblong, 15–[20.1]–26 × 8–[9.3]–12 µm, becoming constricted at the septum, rarely curved; outer spore-wall finely ornamented (microrugulate). Pycnidia immersed, punctiform; conidia bacilliform to elongate-ellipsoid, 4.5–6.5 × 1–1.5 µm.

CHEMISTRY. – Isoarthothelin (major), ±atranorin (minor), 2,5-dichloronorlichexanthone (trace). Spot tests: thallus K+ yellow, C+ orange, KC+ deep orange, P–, UV+ orange.

ETYMOLOGY. – This species is named in honour of William A. Weber, American botanist and lichenologist and co-collector of the type specimen.

ECOLOGY AND DISTRIBUTION. – At present, the new species is known only from the type locality. Associated species included an unidentified Caloplaca species and Tetramelas peruvianus described above.

DISCUSSION. – The new species resembles Tetramelas granulosus (Darb.) A. Nordin, known from the Antarctic Peninsula, South Georgia, South Orkney Islands and the South Shetland Islands (Lamb 1968; Nordin 2000; Øvstedal & Lewis Smith 2001). Both lichens have predominantly 3-septate ascospores, similar conidia and apothecial anatomy and similar (K–) dark brown pigmented hypothecia. However, T. granulosus differs in having an amyloid medulla that lacks medullary calcium oxalate (H2SO4–), a brown N– epihymenium, and in containing 6-O-methylarthothelin.

KEY TO TETRAMELAS IN SOUTH AMERICA

1 Thallus lichenicolous (on foliose ) ...... T. pulverulentus 1: Thallus not lichenicolous ...... 2

2 Thallus growing on bark, old wood, mosses or plant debris ...... 3 2: Thallus growing directly on rocks ...... 4

3 Hypothecium with yellow pigment reacting K+ violet; ascospores 14–25 × 7–9 µm T. thiopolizus 3: Hypothecium without yellow pigment, K–; ascospores 27–31 × 10–11 µm...... T. regiomontanus

394 4 Epihymenium dark brown or dark olive-brown, N– ...... T. anisomerus 4: Epihymenium aeruginose, N+ red-violet to purple-brown ...... 5

5 Medulla amyloid, I+ blue-violet; calcium oxalate absent ...... T. fuegiensis 5: Medulla non-amyloid, I–; calcium oxalate present ...... 6

6 Ascospores commonly 3-septate ...... T. weberianus 6: Ascospores 1-septate ...... 7

7 Ascospores 12–[16.4]–22 × 7–[8.6]–12 µm; oil paraphyses present; prothallus absent .. T. peruviensis 7: Ascospores 12–[13.9]–15 × 5–[7.8]–9 µm; oil paraphyses absent; black prothallus present ...... T. coquimbensis

ACKNOWLEDGEMENTS

I thank Emeritus Professor William A. Weber for generously distributing his many excellent exsiccati collections and to two anonymous reviewers for their helpful suggestions for improving my draft manuscript.

LITERATURE CITED

Bungartz, F., J.A. Elix, K. Kalb and M. Giralt. 2016. New and overlooked species from the Galapagos Islands: The generic concept of reassessed. The Lichenologist 48: 489–516. Calvelo, S. and A.M. Fryday. 2006. New reports of lichens from Argentine Tierra del Fuego and the Falkland Islands (Islas Malvinas). The Bryologist 109: 372–380. Calvelo, S. and S. Liberatore. 2002. Catálogo de los líquenes de la Argentina. Kurtziana 29: 7–170. Dodge, C.W. 1966. New lichens from Chile. Nova Hedwigia 12: 307–352. Elix, J.A. 2014. A Catalogue of Standardized Thin-Layer Chromatographic Data and Biosynthetic Relationships for Substances, 3rd edition. Published by the author, Canberra. 323 pp. Elix, J.A. 2018. New combinations of Tetramelas (Caliciaceae, Ascomycota) and a key to the species in Antarctica. Australasian Lichenology 83: 42–47. Elix, J.A., H. Mayrhofer and J.M. Rodriguez. 2018. Two new species, a new combination and four new records of saxicolous buellioid lichens (Ascomycota, Caliciaceae) from southern South America. Australasian Lichenology 83: 3–13. Galloway, D.J. and W. Quilhot. 1998. Checklist of Chilean lichen-forming and lichenicolous fungi. Gayana Botánica 55: 111–185. Giralt, M. and P. Clerc. 2011. Tetramelas thiopolizus comb. nov., with a key to all known species of Tetramelas. The Lichenogist 43: 417–425. Giralt, M., G. Paz-Bermúdez and J.A. Elix. 2009. The saxicolous xanthone-containing species of the genus Buellia s.l. (Physciaceae, Ascomycota) in the Iberian Peninsula. Nova Hedwigia 89: 321–334 Helms G., T. Friedl and G. Rambold. 2003. Phylogenetic relationships of the Physciaceae inferred from rDNA sequence data and selected phenotypic characters. Mycologia 95: 1078–1099. Kalb, K. 2004. New or otherwise interesting lichens. II. Bibliotheca Lichenologica 88: 301–329. Lamb, I.M. 1968. Antarctic lichens II. The genera Buellia and Rinodina. British Antarctic Survey Reports 61: 1–129. Marbach, B. 2000. Corticole und lignicole Arten der Flectengattung Buellia sensu lato in den Subtropen und Tropen. Bibliotheca Lichenologica 74: 1–384. Nordin, A. 2000. Taxonomy and phylogeny of Buellia species with pluriseptate spores (, Ascomycotina). Symbolae Botanicae Upsalienses 33: 1–117. Nordin, A. 2004. New species of Tetramelas. The Lichenologist 36: 355–359 Nordin, A. and L. Tibell. 2005. Additional species of Tetramelas. The Lichenologist 37: 491–498 Øvstedal, D.O. and R.I. Lewis Smith 2001. Lichens of Antarctica and South Georgia. A guide to their identification and ecology. Cambridge, Cambridge University Press. 411 pp.

395