Lichenologist 33(6): 477–482 (2001) doi:10.1006/lich.2001.0360, available online at http://www.idealibrary.com on

Paralethariicola aspiciliae (, ), a new and species of lichenicolous fungi

V. CALATAYUD, J. ETAYO and P. DIEDERICH

Abstract: The generic name Paralethariicola gen. nov. is introduced for a lichenicolous very similar to Lethariicola, but differing by the presence of excipular hairs, asci with a hemiamyloid apical ring and a more flattened apex, curved or slightly helicoid ascospores withpointed ends, and thicker paraphyses. The single species, Paralethariicola aspiciliae sp. nov., is lichenicolous on Aspicilia, and is known from continental Spain and Sardinia.  2001 The British Society

Introduction who further proposed the relocation of the genus Skyttea from the Ostropales to the Recent lichenological studies on Leotiales. The cleistohymenial, i.e. initially Buntsandstein (Triassic rocks) in eastern closed, apothecia, and the presence of peri- Spain have shown that this substratum hosts physes support the inclusion of our fungus in a rich and interesting flora of and the Ostropales, family Odontotremataceae lichenicolous fungi (Calatayud 1998). (Sherwood-Pike 1987). It differs from the Additional fieldwork, including the first sur- lichenicolous genus Lethariicola Grummann, vey of several minor outcrops, has increased and also from the other genera of the family, the number of taxa known in the region (e.g. by a number of important features, and the Calatayud & Etayo 1999). The most inter- description of a new genus seems to be esting discoveries include a lichenicolous appropriate. fungus parasitic on Aspicilia contorta, which cannot be identified with the current litera- ture. It has cleistohymenial apothecia, an exciple with periphyses and marginal hairs, Material and Methods asci with a thickened hemiamyloid apical wall and a distinct hemiamyloid ring, The material was examined with standard micro- scopical techniques. Photographs were taken with an and hyaline, 3-septate, curved, falcate or Olympus SZH10 stereomicroscope and an Olympus slightly helicoid ascospores. Lichenicolous, BX50 microscope with DIC. The amyloid reactions ostropalean and leotialean genera sharing were tested with Lugol’s reagent, either without (I) or some of these features have recently been with pre-treatment with KOH (K/I). Microscopical measurements were made in water. The ascospore keyed out by Diederich & Etayo (2000), range (in dead state) was calculated after rejecting manually 10% of the highest and 10% of the lowest values measured; the mean values are given in italics, V. Calatayud: Fundación CEAM, C/Charles R. Darwin and the extremes within parentheses. Excipular pig- 14, Parc Tecnològic, ES-46980 Paterna, València, ments have been studied in water, potassium hydroxide Spain. (K) and nitric acid (N). J. Etayo: Navarro Villoslada 16, 3a dcha., ES-31003 The comparison with species of Lethariicola is based Pamplona, Navarra, Spain. on unpublished results by Diederich, Etayo & P. Diederich: Musée national d’histoire naturelle, Zhurbenko, referring to the complete available material 25 rue Munster, L-2160 Luxembourg, Luxembourg. of specimens belonging to this genus.

0024–2829/01/060477+06 $35.00/0  2001 The British Lichen Society 478 THE LICHENOLOGIST Vol. 33

Taxonomy 3-septatis, (21–)22–27(–31)3–3·5 m. In thallo lichenum generis Aspicilia crescens. Paralethariicola Calatayud, Etayo & Typus: Spain, Region of Aragón, Prov. Teruel, be- Diederich gen. nov. tween Puebla de Arenoso and Olba, close to Los Lucas, beside the main road, c. 2kmEofOlba,c. 408N, A genere Lethariicola praecipue margine excipuli 037W, alt. c. 700 m, on Aspicilia contorta subsp. hoff- piloso, ascis annulo hemiamyloideo, ascosporis curva- manniana, on slightly carbonated sandstone, 14 iii tis, falcatis vel parum helicoideis et paraphysibus 1999, V. Calatayud s. n. (MA-Fungi 49914— latioribus differt. holotypus; hb. Calatayud 67, hb. Diederich and hb. Typus: Paralethariicola aspiciliae Calatayud, Etayo & Etayo—isotypi). Diederich Ascomata initially perithecioid and closed, (Figs 1 & 2) with the disc entirely covered by the exciple, opening by a pore, becoming deeply urceolate Ascomata initially immersed, later when mature. Exciple paraplectenchymatous, erumpent, mostly closed and with a small dark reddish brown in its outer part and central pore when dry, but opening to hyaline towards the hymenium; excipular pig- become urceolate and exposing the disc ment extracellular, not granulose, reddish when moist, arising singly or in groups, brown, K+ more greyish brown, N+ orange sometimes confluent, dark reddish brown, brown; excipular margin with brown hairs the oldest with a margin markedly striate, (but hyaline or pale brownish in the lower 200–500 m diam.; pore irregular or part, close to the hymenium) that are I, rounded. Exciple laterally dark reddish K/I; periphysoids well developed on the brown in its uppermost and outer part, inner excipular surface, developed up to the hyaline towards the hymenium, 50–65 m base of the asci and paraphyses, perpen- thick, basally dark reddish brown, 20– dicular to the paraphyses, hyaline, septate, 35 m; excipular cellspolygonal or some- branched, I,K/I (but plasma I+ red- what elongate, 4–7 m diam.; excipular dish). Hymenium hyaline, hymenial gel I+ hairs c. 10–253–4 m; inner part of the directly red (not bluish, then red), K/I+ blue; lateral exciple provided with abundant, short subhymenium and epihymenium hyaline. and hyaline periphysoids, 6–122–2·5 m. Paraphyses simple, septate, apically not or Hymenium 60–75 m high; subhymenium hardly thickened. Asci subcylindrical when 10–15 m high. Paraphyses 2–3(–4) m young, clavate or slightly saccate when ma- thick. Asci 8-spored, 40–559–11 m. ture, basally with a short stipe, apically Ascospores twisted within the asci, 3-septate, slightly flattened, wall apically thickened, with a relatively small number of minuscule with a distinct ocular chamber, I+ red, K/I+ guttules (observed in K), I,KI, (21–) blue, except the apical dome which is K/I 22–24·8–27(–31)3–3·5 m, length/width or pale blue (similar to the Lecanora-type), ratio (6·3–)7·0–8·0–9·0(–10·3) [n=36]. with a distinct hemiamyloid ring (in some asci difficult to observe). Ascospores curved, falcate Hosts. On the thallus of Aspicilia contorta or slightly helicoid, with more or less pointed subsp. hoffmanniana and of an unidentified ends, transversely septate, smooth-walled, Aspicilia species. The infected parts are lacking a visible perispore, hyaline. Anamorph bleached and partly destroyed by the fungus unknown. Vegetative hyphae indistinct. (appearing as if they had been grazed by insects), which makes it easy to find in the Etymology. From para- (Greek, similar to) field. and Lethariicola, its most closely related genus. Etymology. The epithet ‘aspiciliae’ refers to the host genus. Paralethariicola aspiciliae Calatayud, Etayo & Diederich sp. nov. Ecology and distribution. Paralethariicola Paralethariicola insignis ascomatibus 200–500 m aspiciliae is known from two Mediterranean diam., ascis 8-sporis, 40–559–11 m, et ascosporis localities. The type was gathered in a locality 2001 Paralethariicola aspiciliae—Calatayud et al. 479

F.1. Paralethariicola aspiciliae (holotype). A, habitus; B, closer view of an ascoma, showing the striate margin; C, section through an ascoma; D, exciple; E, periphysoids (lower part, hyaline) and hairs (upper part, brown), observed in K; F, paraphyses in K/I; G, asci in K/I; H, detail of ascus apex in K/I; I & J, ascospores in I. Scales: A=1 mm; B=0·5 mm; C=100 m; D=25 m; E–J=10 m. F.2. Paralethariicola aspiciliae (holotype). A, section through the exciple of a mature ascoma, with periphysoids (inner part) and hairs (apical part, brown); B, ascospores; C, transverse section through a young ascoma, still without excipular hairs; A,B&Cobserved in water; D, hymenium with asci in several developing stages, observed in K/I; E, several ascomata, the youngest with radial fissures and the most mature ones striate and more flattened. Scales: A,B&D=10m; C=50 m; E=0·25 mm. 2001 Paralethariicola aspiciliae—Calatayud et al. 481 in central-eastern Spain (region of Aragón, both taxa. This clearly indicates a close Prov. Teruel), on a relatively small out- phylogenetic relationship. On the other growth of slightly carbonated red sandstone, hand, genera within the Odontotremataceae completely surrounded by calcareous rocks. are often separated by minor differences. If The second specimen is from Sardinia; this P. aspiciliae was included in an enlarged sample was published as Pleospilis sp. and as concept of Lethariicola, then several other Skyttea sp. by Nimis & Poelt (1987: 184 & genera within the family would logically also 222). have to be united. In our opinion, all the ff Additional material examined. Italy: Sardinia: Prov. di erences enumerated above strongly sup- Sàssari, Valle dei Nuraghi, 1·5 km SE der Bahnstation port the distinction of two distinct genera, at von Torralba, c. 350 m, beweidete, frühjahrsfeuchte least as long as narrow generic concepts are Rasen, Gräben, Wegränder, Mauern, vereinzelte used within the whole family. niedere Silikatblöcke, 1986, H. Mayrhofer 6091 (GZU). The genus Phragmiticola Sherwood was the only hitherto recognized member of Discussion the Odontotremataceae in which asci have an amyloid apical ring. Because of this The new fungus strongly resembles species character Sherwood-Pike (1987) hesitated of the lichenicolous genus Lethariicola to include the genus in the Ostropales, but, (Lumbsch & Hawksworth 1990; Diederich, following Baral (pers. comm.), the mono- Etayo & Zhurbenko, unpublished data). typic genus is a typical member of this order. Both share similar cleistohymenial asco- Furthermore, the exciple of Phragmiticola mata, a similar excipular structure, the same presents verrucose marginal hairs (Baral, excipular pigments, hyaline, transversely pers. comm.). We believe that Paralethari- septate (in Lethariicola rarely submuriform), icola aspiciliae should nevertheless not be multiguttulate ascospores, and similar peri- included in Phragmiticola, as in the latter: the physoids, paraphyses and asci. Species of hymenium and the ascus wall (except that of Lethariicola are distinguished by the absence apical ring) are non-amyloid; the apical ring of excipular hairs and the absence of a is euamyloid (fide Baral, pers. comm., versus hemiamyloid apical ring in the ascus wall hemiamyloid in Paralethariicola); the excipu- (the ascus wall is either K/I or pale bluish, lar structure, especially near the margin, is but never with any special apical structures not exactly the same; the excipular hairs are visible by light microscopy). Furthermore, verrucose; the ascospores have a distinct, ascospores in Lethariicola are generally thick perispore; the asci have a narrower, ellipsoid, rarely narrowly ellipsoid, but never conical apex, with a narrower apical ring; bent or helicoid, and usually with a rounded and the ecology is different, the only known apex; the asci in Lethariicola are apically species of Phragmiticola being saprophytic more rounded, whilst those in Paralethari- on Phragmites (Sherwood-Pike 1987; Baral, icola aspiciliae have a distinctly flattened pers. comm.). apex; finally the paraphyses in species of The genera Skyttea Sherwood, D. Lethariicola are rarely over 1·5 m in diam- Hawksw. & Coppins and Spirographa eter, exceptionally up to 2 m, whilst Zahlbr. (syn. Pleospilis Clem.), under which those of P. aspiciliae are distinctly thicker, the Sardinian specimen has initially been 2–3(–4) m in diameter. published (Nimis & Poelt 1987) are very For a long time we hesitated between the distinct (see Diederich & Etayo 2000). inclusion of P. aspiciliae within Lethariicola Paralethariicola aspiciliae also shows some and the description of a distinct genus. common features with the lichen genus Ontogenetically, the ascomata of Para- Ramonia Stizenb. As described by Sherwood lethariicola are similar to those of Lethariicola, (1977), this genus is characterized by with the excipular hairs in Paralethariicola having pinkish or light-coloured, immersed developing only at a later stage (see Fig. 2C); ascomata and a two-layered margin, with an also the excipular pigment is the same in exciple of ‘textura angularis’ with, on the 482 THE LICHENOLOGIST Vol. 33 inner surface a layer of periphysoids. A dark Generalitat Valenciana and Bancaixa for support to the coloured species, R. nigra Coppins, was CEAM. eventually described by Coppins (1987). This author pointed out that at least two R taxa, R. nigra and R. chrysophaea (Pers.) Calatayud, V. (1998) Líquenes y hongos liquenícolas de Veˇzda, have an amyloid ring-like structure in rocas no carbonatadas en el Sistema Ibérico oriental e the ascus apex. Paralethariicola differs from Islas Columbretes. Unpublished Thesis, Universitat Ramonia by its hairy exciple and hemiamy- de València. Calatayud, V. & Etayo, J. (1999) Codonmyces and loid asci (they are non-amyloid in species of Lichenostella, two new genera of lichenicolous Ramonia), and by its curved, transversely conidial fungi. Lichenologist 31: 593–601. septate ascospores. Although the shape, sep- Coppins, B. J. (1987) The genus Ramonia in the British tation and ascospore number are very vari- Isles. Lichenologist 19: 409–417. Diederich, P. & Etayo, J. (2000) A synopsis of the able within the genus Ramonia, species with genera Skyttea, Llimoniella and Rhymbocarpus curved or helicoid ascospores have not been (lichenicolous , Leotiales). Lichenolo- reported so far. The ecology is also different, gist 32: 423–485. doi: 10.1006/lich.2000.0290. since the genus Ramonia includes only Lumbsch, H. T. & Hawksworth, D. L. (1990) The lichenized species, most of them epiphytic. species of Lethariicola Grumm. (Odontotremata- ff ceae). Bibliotheca Lichenologica 38: 325–333. In spite of these di erences, there are Nimis, P. L. & Poelt, J. (1987) The lichens and remarkable similarities between both genera, lichenicolous fungi of Sardinia (Italy): an anno- and the systematic position of Ramonia tated list. Studia Geobotanica 7, Supplement 1: should be reconsidered to establish if it defi- 1–269. Purvis, O. W., Coppins, B. J., Hawksworth, D. L., nitely belongs to the Ostropales, rather than James, P. W. & Moore, D. M. (Eds) (1992) The to the (e.g. Purvis et al. 1992; see Lichen Flora of Great Britain and Ireland. London: also the discussion by Coppins 1987). Natural History Museum Publications. Sherwood, M. A. (1977) The ostropalean fungi. I. We are most grateful to the curator of GZU for the loan Mycotaxon 5: 1–277. of the Sardinian specimen of the new fungus, and to Dr Sherwood-Pike, M. A. (1987) The ostropalean fungi. EmmanuëlSérusiaux and Hans-Otto Baral for precious III. The Odontotremataceae. Mycotaxon 28: comments on the manuscript. V.C. wishes to thank the 137–177.

Accepted for publication 9 October 2001