Joum. Hallor; Bot. Lab. No. 68: 279- 284 (June 1990)

THE GENERA AND IN

D. K. UPRETI1 AND B. BUDEL2

ABSTRACT. Five species of Pettuta and two species of Heppia are reported from India. Of the seven species, Heppia tutosa, H. trichophora, Pettuta euptoca, P. obscurans, P. patellata, P. tortuosa and P. zahtbruckneri, only H. lrichophora and P. euptoca were previously known from Inida. No lichen compounds were detected in all taxa by TLC.

INTRODUCTION The lichen family has so far been represented in India by two species, Heppia trichophora Awas. et S. Singh and Peltula euploca (Ach.) Poelt ex Pisut, reported from Mt. Abu, by Awasthi and Singh (1979). Wetmore (197 1) and Swinscow and Krog (1979), in revising the American and East African species of Heppia and Peltula respectively, treated both the genera within the family Heppiaceae. Blidel (1986 in; Eriksson & Hawksworth 1987a, p. 149) included the Peltula in the new family , on the basis of ascocarp ontogeny together with the unique polysporous and unitunicate-rostrate ascus structure and function. Furthermore, Blidel (1 987b) described three new species and made three new combinations of Peltula while revising the genus from South Africa and added six more species to the ten known for Africa, previously described by Swinscow and Krog (1979) from East Africa. This paper deals with five species of Peltula and two of Heppia from India. The taxonomic account and a key for species identification are provided.

MATERIAL The study is based on material lodged at Lucknow University, Lucknow Herbarium (LWU) and the herbarium of the National Botanical Research Institute, Lucknow (LWG). Type material from the herbaria of the Botanical Museum, Helsinki (H) and Naturhistorisches Museum, Wien (w) was also investigated in this study.

K EY TO THE INDIAN SPECIES OF HEPPlA AND PELTULA la. Thallus sorediate ...... Peltula euploca 1b . Thallus non sorediate ...... 2 2a. Apices of lobules densely trichoma tic ...... Heppia trichophora 2b. Apices non-trichomatic ...... 3

I Cryptogamic Botany Section, National Botanical Research Institute, Lucknow- 226 001, India. 2 Fachbereich Biologie/Botanik, Philipps Universitat, Karl-v.-Frisch-Str., D-3550 Marburg/Lahn, F. R. Germany. 280 Journ. Hattori Bot. Lab. No. 68 I 990

3a. Asci 8-spored ...... Heppia lutosa 3b. Asci multispored ...... 4 4a. Apothecia immersed, disc punctiform, squamules multicarpous ...... 6 4b. Apothecia superfical, disc expanded, squamules I-(rarely 3- 5) carpous...... 5 5a. Thallus upper surface reticulate rugose, margins upwards and smooth, forming a rim ...... Peltula patellata 5b. Thallus upper surface not so; rosette-shaped ...... Peltula obscurans 6a. Thallus subfruticose, ascending, lobes cylindrical, upright, top flattened ...... Peltula tortuosa 6b. Thallus squamulose, lobes inflated, not ascending, horizontal .. Peltula zahlbruckneri

SPECIES I. Heppia [utosa (Ach.) NyJ., Syn. Lich. 2: 45. 1855. - Collema lutosum Ach., Syn. Lich.: 309. 1814. Type: Germany (Holotype: H-ACH, not seen). Fig. 1. Thallus squamulose, olive to greenish grey; squamules 1- 7mm; flat-concave, roundish or irregular; margins slightly lobulate; upper surface rough and cracked; attached by a tuft of hyphae. Thallus 150--300 flm, upper cortex 30--70 flm thick; algal layer 40--100 flm or scattered throughout the thallus; hyphal cells anticlinally arranged within the medullary region; no lower cortex developed. Apothecia mostly one per squamule; immersed, disc 0.3- J.Omm expanded, without thalloid rim; epithecium yellowish, sometimes reddish brown, K -; hymenium 100--130flm thick; J + vinose red; paraphyses simple, septate; asci clavate without gelationus sheath, 60--75 x 15- 20 flm; 8-spored, spores hyaline, simple, oval-globose, 10--15 x 6-10 flm . Remarks: H. lutosa is characterized by flat, concave, roundish to irregular, olive to greenish-grey squamules. The asci and spores of the Indian species are somewhat smaller than in specimens from other parts of the world. This might be due to the immature stage of the ascocarpia in the Indian specimen. The other Indian species, H. trichophora, is differentiated by the presence of hyaline trichomes at the apices of the lobules. Specimen examined: U.P.; Allahabad, on black soil, Sepl. 1926, Naturhistorisches Museum, Wien, Dudgeon 729 (LWG) .

2. Heppia trichophora Awas. et Singh, Norw. J. Bot. 26: 92, 1979. Type: India, Rajasthan, Sirohi district, Mt. Abu, Kanrter Kund area, on rock, Singh 78 .151 (Holotype: LWU). Corticolous and saxicolous, thallus grey to greyish brown; squamules 1- 2 mm diameter, dichotomously to irregularly divided; apices of lobules flattened to swollen, densely trichoma tic; upper surface smooth, under surface erhizinate, attached by edge to the substrate. Thallus 50--100 flm thick, homoiomerous, paraplectenchymatous throughout, cells rounded to angular, cortex 10 flm thick, cellular; photobiont a member of the family Scytonemataceae, cells sphaerical, 4-10 flm in diameter. Apothecia absent. Specimens examined: Rajasthan, Sirohi district, Ml. Abu, all. ca. 1350m, on bark, 10.1.1978, Singh 78 .157 (LWU); Sirohi darbar area, all. ca. 1230 m; on rock, 9.1.1978, Singh 78.28 (LWG) .

3. Peltula euploca (Ach.) Poelt ex Pisut, Acta Rer. Nat. Mus. Slov. 13: 8, 1967. - Lichen euplocus Ach., Suec. Prodr.: 141, 1798. Holotype: Sweden; Westring (H-ACH). Fig. 2. Thallus squamulose, umbilicate, squamules grey green; peltate or orbicular to irregularly D. K. UPRETl & B. BODEL: Lichen genera Heppia and Petluta in India 281

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FIGS. 1--6. Fig. l. Heppia tutosa, fertile squamule with immersed apothecia, ascus. Fig. 2. Pettula euptoca, sterile umbilicate squamule with sorediate margin, ascus. Fig. 3. Peltuta obscurans, fertile squamule with adnate apothecium, ascus. Fig. 4. Petluta patellala, fertile squamule, ascus. Fig. 5. Peltula tortuosa, fertile fruticose thallus, ascus. Fig. 6. Peltula zahlbruckneri, fertile sq uamule, ascus. 282 Journ. Hattori Bot. Lab. No. 68 I 990 lobulate, imbricate, 0.75- 15mm in diameter, margins down-rolled, entire or slightly lobulate, sorediate; soredia granular; attached to substratum by umbi-Iicus. Thallus 200-450/lm thick; upper cortex absent; algal layer 60-1 SO /lm thick; medulla with loose hyphae, 70-120 /lm thick; lower cortex paraplectenchymatous, 4-9 cells and 40-80 /lm thi.ck. Apothecia 3- 7 per squamule, totally immersed; disc punctiform or expanded, up to O.S mm, lacking thalloid rim; epithecium pale yellow, K -; hymenium 100-250/lm high, I + blue; paraphyses simple, capitate; asci 60-100 x 10-20 /lm, polysporous; spores ellipsoid, 3- S x 2- 3/lm. Remarks: P. euploca is the most frequently occurring species in India. It is charcterized by peltate or orbicular sometimes polyphyllous and sorediate squamules; when fertile, soredia are often lacking. The asci and spores are not fully developed in the Indian specimens. Specimens examined: Gujarat, Banas Kantha district, on rock, 1976, s.n. (LWG). Jammu & Kashmir, Pahalgam, all. 2134-2438 m, on exposed boulders, 2. 6. 1977, Dange 77-41 (LWU) . Madhay Pradesh, Jabalpur district, Mandan Mahal, 600 m, on rock, 24. I. 1980, Upreti & Misra 80--296 (LWU). Bedinagar Hills, on rock, 25. I. 1980, Upreti & Misra 80--316 (LWU). Tamil Nadu, Nilgiri Hills, 1800m, on stones, 1.1.1971 , Awasthi & Singh. 71 -1 (LWU). Ootacomund, Valley Hill View, on stones, I. 12. 1973, Singh 73-526 (LWU). , Mirzapur district, Shivpur Hills, alt. 500-600 m, on rock, 3. 2. 1980, Upreti & Misra 80--601, --{)15, --{)19 (LWU). Sirsi Falls, all. 600 m, on rocks, 4. 2.1980, Upreti & Misra 80--599, - 600 (LWU). Wyndom Falls, Kotwa, on rocks, 5. 2. 1980, Upreti & M isra 80-{)32/B, --{)40 (LWU). Uttar kashi district, harsil, 2400m, on rock, 22. 4. 1962, Singh 97530 (LWG).

4. Peltula obscurans (Nyl.) Gyelnik, Rep. Spec. Nov. Regn. Veg. 38: 308, 1935. - Endocarpiscum obscurans Nyl., Bull. Soc. Linn. Normand. 2(6): 309, 1872. Typus: France, Collioure, 1872, Nylander (Lectotype: H-NYL 30900, Wetmore, 1971). Fig. 3. Saxicolous, squamulose thallus, squamules greenish grey to olive, 0.7S-2.0mm in diam., rounded to angular, plain to convex, sometimes deeply lobed, rosette-shaped, attached by umbilicus. Thallus 100-250 /lm thick; epinecral layer 8- 30/lm, algal layer 40-100/lm thick; medulla of loose hyphae, 10-30/lm thick; lower cortex 3- S cell-layered, 15-40/lm thick. Apothecia 1- 2 (rarely 3) per squamule, disc brown-orange, up to 0.4 mm in diam., with thalloid rim, epithecium 1-, K-; hymenium 120-150/lm high, 1+ vinose-red; asci clavate, with gelatinous sheath, 72-80 x IS- 20 /lm, multispored, spores hyaline, simple, oval, 2- 3 x 1- 2/lm (immature ascocarp). Remarks: P. obscurans is close to P. michoacanensis (B. de Lesd.) Wetm. in size ofsquamules, but the latter has deeply concave squamules and the ascocarp lacks a thalloid rim. Specimens examined: Uttar Pradesh, Mirzapur district, Shivpur, 500--{)00 m, on rocks, 3.2.1980, Upreti & Misra 80--597 (LWU). Wyndom Falls, Kotwa, 500 m, on rocks, 5.2.1980, Upreti & Misra 80--{)33 (LWU). Rae Bareli district, Pithan village, on tiles, Singh 97600, 97617 (LWG). , Jabalpur, Karia Pathar, 550 m, on rocks, 25. I. 1980, Upreti & Misra 80--320 (LWU).

S. Peltula patellata (Bag!.) Swinscow et Krog, Norw. J. Bot. 26: 221, 1979. - Acarospora patellata Bagl., Nuov. Giorn. Bot. Ital. 7: 24S . Holotype: Ethiopia, Beccari 74 (FI, not seen). Fig. 4. Thallus squamulose, olive-brown, squamules 2.0-3.0mm in diam., concave to plane, orbicular to angulate or lobulate or ascending towards margin, upper surface reticulate-rugose, attached by a more or less central tuft of hyphae. Thallus IS0-2S0 /lm thick; epinecral layer 20-40/lm, algal layer S0-90 /lm thick; medulla paraplectenchymatous, partly composed of longitudinally disposed hyphae; lower cortex paraplectechymatous, 40-80/lm thick. Apothecia 1- (rarely S) per squamule, 0.5-1.0mm in diam., emergent; disc reddish brown, with thalloid D. K. UPRETI & B. BODEL: Lichen genera Heppia and Peltula in India 283 rim; epithecium K -; hymenium 140-200 Ilm high, I + blue, paraphyses simple to sparingly branched; asci clavate, with serriate gelatinous sheath, 50-80 x 15- 20 Ilm (immature), multispored; spores hyaline, simple, globose, 3--4 x 2- 3Ilm. Remarks: P. patellata is close to P. obscurans, but the latter has deeply lobed, imbricate squamules with a smooth upper surface. Specimens examined: Uttar Pradesh, Lucknow district, National Botanical Gardens, on wall, 4. 8. 1963, Singh 97584, 97586 (LWG). Rana Pratap Marg, on wall, 10.1978, Singh 85620 (LWG). Dilkusha Garden, on plaster of grave, 9. 8. 1988, Upreti 201831 (LWG).

6. Peltula tortuosa (Nees) Wetm., Ann. Mo. Bot. Gdn. 57: 205, 1971. -Dufourea tortuosa Nees, Horae Phys. Berol.: 43, 1820. Type: Venezuela, Orinoco River, Humboldt (Lectotype: H-NYL 30890). Fig. 5. Thallus subfruticose, olive to brown to brownish black, lobes flattened, upright, somewhat branched, partly terete, top mostly flattened, surface smooth, fixed to the substratum by a central attachment. Thallus lobes 1- 2 mm long, 0.5-1.0 mm broad, epinecral layer ca. 51lm, outer part of algal layer paraplectenchymatous, algal layer 50-100 Ilm thick; medulla of loose hyphae. Apothecia rarely present, immersed in lobes, 1 per lobe; up to 0.5 mm in diam., disc punctiform, to 0.2 mm wide, brown-black; epithecium pale yellowish, 9- l81lm thick, K -; hymenium 90-150 Ilm high, 1+ vinose-red; paraphyses simple, septate; asci clavate, 50- 70 x 18- 20 Ilm (immature), multispored; spores hyaline, simple, globose, 2--4 x 2- 3Ilm. Remarks: P. tortuosa specimens from India are smaller than specimens from the American continent and resemble South African specimens. Because the material was not fully developed, no further conclusions can be made. Specimens examined: Uttar Pradesh, Mirzapur district, Sirsi Falls, 600 m, on rocks, 4. 2. 1980, Upreti & M ista 80-609, -627 (LWU). Wyndom Falls, Kotwa, 500 m, on rocks, 5.2.1980, Upreti & Misra 80-632 (LWU).

7. Peltula zahlbruckneri (Hasse) Wetm., Ann. Mo. Bot. Gdn. 57: 205, 1971. - Heppia zahlbruckneri Hasse, Bryologist 14: 100, 1911. Type: California, Rubio Canyon, San Gabriel Gelage bei Pasadena, Kingman 1159 (Holotype: w; Isotype: FH). Fig. 6. Thallus squamulose, squamules greenish-grey, inflated, minutely lobed at margins, convex or flattened when young, 1--4mm wide, attached to substratum by umbilicus. Squamules 160-350 Ilm thick, epinecrallayer 40-60 Ilm, algal layer 70-150 Ilm thick; algal cells sometimes scattered to the lower cortex; medulla of loose hyphae, 40-80 Ilm thick, lower cortex 4--7 cell-layered, 30-60 Ilm thick. Apothecia 1-5 per squamule, immersed, disc punctiform to 0.3-0.75 mm expanded, lacking thalloid rim; epithecium yellowish, K -; hymenium 130-170 Ilm thick, 1+ vinose-red; asci clavate, 60-90 x 10-151lm, polysporous; spores hyaline, simple, globose, 2--4 x 2- 3Ilm; paraphyses simple to sparsely branched, septate. Remarks: Young scales of P. zahlbruckneri can be easily confused with young scales of P. euploca or P. cylindrica Wetm. SpecimeDs examined: Uttar Pradesh, Mirzapur district, Wyndom Falls, Kotwa, 500 m, on rocks, Upreti & Misra 80-645 (LWU). Shivpur Hills, 500--600m, on rocks, Upreti & Misra 80-596 (LWU).

ACKNOWLEDGEMENTS We wish to express our sincere thanks to Dr. D. D. Awasthi and the curators of the herbaria of the Botanical Museum, Helsinki (H) and Naturhistorisches Museum, Wien (w) for the loan 284 Journ. Hattori Bot. Lab. No. 68 I 990 of material. D. K. Upreti is grateful to Dr. Ajay Singh for his valuable suggestions, to the Director of the National Botanical Research Institute, Lucknow for providing facilities, and to Mr. Murari Ranjan for assisting in laboratory work. B. Biidel gratefully acknowledges the support of the Deutsche Forschungsgemeinschaft.

REFERENCES

Awasthi, D. D. & S. R. Singh. 1979. New or otherwise interesting from Mt. Abu, Rajasthan, India. Norw. 1. Bot. 26: 91 - 97. Blidel, B. 1987a. Toxonomy and biology of the lichen genus Pe/tu/a Nyl. In Progress and problems in lichenology in the eighties. In E. Peve1ing (ed.), Bibl. Licheno!. 25: 209- 217. ---. 1987b. Zur Biologie und Systematik der Flechtengattungen Heppia und Peltu/a im slid lichen Afrika. Bib!. Lichenol. 23: 1- 105. Eriksson, O. & D. L. Hawksworth. 1986. Notes on ascomycete systematics. Nos 1- 224. Systema ascom. 5: 114-174. Swinscow, T. D. V. & H. Krog. 1979. The lichen genera Heppia and Peltula in East Africa. Norw. l. Bot. 26: 213-224. Wetmore, C. M. 1971 . The lichen family Heppiaceae in North America. Ann. Mo. Bot. Gdn 57: 158- 209.