PLANT SCIENCE TODAY, 2021 Vol 8(4): 889–899 https://doi.org/10.14719/pst.2021.8.4.1265 ISSN 2348-1900 (online) HORIZON e-Publishing Group RESEARCH ARTICLE Pyrenocarpous lichens in Goa with five new records to India Pallavi Randive1, Gaurav K. Mishra2, Sanjeeva Nayaka2*, D. K. Upreti2 & M. K. Janarthanam1 1 Department of Botany, Goa University, Taleigao Plateau, Goa 403 206, India Lichenology Laboratory, CSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow 226 001, India *Email: nayaka.sanjeeva@gmail.com 2 ARTICLE HISTORY ABSTRACT Received: 21 May 2021 Accepted: 27 August 2021 Available online: 29 September 2021 The pyrenocarpous lichens are the one which produces perithecial ascocarps. They are one of the prominent groups of lichens in tropical forests. Frequent incidence of pyrenocarpous taxa in lichen biota of Goa prompted us to take up the exclusive study of this group in the State. The study revealed the occurrence of 79 species belonging to 15 genera and seven families. The family Pyrenulaceae had the maximum number of 23 species, while 20 belonged to the family Porinaceae. In comparison to North Goa, South Goa is fairly well explored for lichens representing 71 species from 11 localities. The following five species are reported as new to India — Porina exserta, P. siamensis, Pyrenula dissimulans, P. pyrenastrospora and P. rinodinospora. With the addition of five new records, the Goa State now represents 165 species of lichens, out of which 48% is represented by pyrenocarpous lichens. The present study will be useful for monographic studies on pyrenocarpous lichens and for environmental monitoring studies in the area, as this can be considered as a key indicator species. KEYWORDS Arthopyreniaceae Biodiversity Porinaceae Pyrenulaceae Taxonomy Western ghats Introduction The Western Ghats traverses through the State of Goa, India and is a well-known biodiversity hotspot. Extensive lichen exploration in the State has been initiated recently, and so far, 160 species are reported (1-6). Frequent occurrence of pyrenocarpous lichens during the exploration in the State indicated their dominance which prompted us to carry out an exclusive study on this group. Pyrenocarpous are a group of lichens having perithecia as their ascocarps. Pyrenocarpous taxa commonly grow on the bark of trees, sometimes on rocks, soil or leaves, mostly in moist tropical regions of the world (7). In the past, pyrenocarpous lichens collected from Goa were included in various monographic works (814) and few new species such as Anthracothecium goaense A. Singh [current name Pyrenula gibberulosa (Vain.) Aptroot] and P. subacutalis Upreti [current name Pyrenula maravalensis Vain.] were also described. The recent floristic studies in the State reported 14 pyrenocarpous taxa from the Cotigao Wildlife Sanctuary (15) and 13 taxa from Bondla and Bhagwan Mahavir Wildlife Sanctuaries (16). A list of lichens from the Goa State reveals 118 species under 42 genera and 23 families, of which 44 were pyrenocarpous lichens (1). Further, in an exclusive survey of Cotigao Wildlife Sanctuary 36 pyrenocarpous taxa were reported under nine genera (3). In the studies on foliicolous lichens, there enumerated a total of 15 pyrenocarpous lichens (2, 4). These reports clearly indicate that Goa State has a rich diversity of pyrenocapous lichens. Therefore, carrying out an exclusive study on pyrenocarpous lichens of the State was inevitable. The ultimate aim of the study was to produce a checklist and identification key for pyrenocarpous lichens of Goa State by compilation of previous reports and by more exploration in the State. Materials and Methods The present study is based on published literature, previously collected specimens deposited in the herbarium LWG of CSIR-National Botanical Research Institute, Lucknow and freshly collected specimens from Goa. About 350 specimens from 19 localities of Goa (Fig. 1) were available for the study. Morphological and anatomical characters were examined using stereo zoom Leica S8APO and light DM2500 microscopes attached to a camera. Thin sections of perithecia were cut using a razor blade under a stereo zoom microscope. All anatomical measurements were recorded in plain water, while © Randive et al (2021). This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution and reproduction in any medium, provided the original author and source are credited (https://creativecommons.org/licenses/by/4.0/). 890 RANDIVE ET AL Fig. 1. Map of Goa showing North and South districts and localities surveyed for present study. 10% KOH was used for the detailed study of asci and ascospores. For spot tests, the usual reagents of K, C and P were used. To identify lichens substance, thin layer chromatography (TLC) was performed in solvent system C following the standard method (17). Fresh specimens were preserved in the herbarium of Goa University (GU), and a set of voucher specimens were deposited in herbarium LWG. The specimens were identified up to species level with the help of keys of earlier published literature (18-25). The classification of lichens summarized in reference 26 was followed for arranging species under their respective families. The identity of species was confirmed by matching with type specimens or well-identified specimens available at LWG. Results The study revealed the occurrence of 79 species of pyrenocarpous lichens belonging to 15 genera and seven families in Goa. Five species were recorded for the first time from the country (Table 1). The brief descriptions for the five newly recorded lichens and key for all the pyrenocarpous lichens encountered in Goa are provided. The Pyrenulaceae family was most dominant in the State, with 23 species followed by Porinaceae (20 spp.) and Monoblastiaceae (13 spp.). Among the genera, Pyrenula was dominant with 24 species, followed by Porina (19 spp.), Anisomeridium (12 spp.) and Strigula (8 spp.). All the species recorded were crustose except for Endocarpon subrosettum A. Singh & Upreti, which was squamulose. A total of 58 species were found to grow on tree trunks, branches and twigs (corticolous), followed by 16 species that grow on leaves (foliicolous), while five species grow on rocks (saxicolous). Astrothelium meristosporum (Mont. & Bosch) Aptroot & Lücking and A. scoria (Fée) Aptroot & Lücking exhibited their substrate specificity with restricted occurrence on cashew nut (Anacardium occidentale Linn.) trees. The species of Porina and Pyrenula exhibit maximum substrate diversity as they were found growing on various trees, leaves and rocks, while all Strigula species exhibit luxuriant growth only on leaves. Artocarpus integrifolius L.f., Anacardium occidentale L., Cocos nucifera L. and Terminalia elliptica Willd. were the common phorophytes for the growth of pyrenocarpous lichens. Porina interestes (Nyl.) Harm., P. tetracerae (Afz.) Müll. Arg., Pyrenula aspistea (Ach.) Ach., P. oculata A. Singh & Upreti and Trypethelium eluteriae Spreng. were the most common species of the State. PLANT SCIENCE TODAY 891 Table 1. List of pyrenocarpous lichens recorded from Goa and their distribution [Note: + = present, - = absent, CR = Crustose, SQ = Squamulose, C = Corticolous, S = Saxicolous, F = Foliicolous, BMWS = Bhagwan Mahavir Wildlife Sanctuary, CWS = Cotigao Wildlife Sanctuary, BWS = Bondla Wildlife Sanctuary, Goa Uni. = Goa University, *New record for India] Goa Uni. Melauli Panjim Ponda Radigati Valpoi Agonda BMWS Caranzol Colem CWS Mollem Pirla Quepem Quinamol Rivona Tambdisurla P. subinterstes (Nyl.) Müll. Arg. BWS 36 37 P. tetracerae (Afz.) Müll. Arg. 38 Trichothelium alboatrum Vain. Pyrenulaceae 39 Lithothelium decumbens (Müll. Arg.) Aptroot 40 Pyrenula adacta Fée 41 P. aggregata (Fée) Fée 42 P. approximans (Kremp.) Müll. Arg. 43 P. aspistea (Ach.) Ach. 44 P. breutelii (Müll. Arg.) Aptroot 45 P. brunnea Fée 46 P. castanea (Eschw.) Müll. Arg. 47 P. dissimulans (Müll. Arg.) R.C. Harris* 48 P. fetivica (Krempelh.) Müll. Arg. 49 P. gibberulosa (Vain.) Aptroot 50 P. immissa (Stirt.) Zahlbr. 51 P. leucostoma Ach. 52 P. leucotrypa (Nyl.) Upreti 53 P. mamillana (Ach.) Trevis. 54 P. maravalensis Vain. 55 P. nitidula (Bres.) R.C. Harris. Amona Arthopyreniaceae 1 Arthopyrenia alboatra (Kremp.) Müll. Arg. 2 A. finkii Zahlbr. 3 A. grisea (Schierch.) Körb. 4 A. indusiata Müll. Arg. 5 A. nidulans Müll. Arg. Monoblastiaceae 6 Anisomeridium albidoatrum (Nyl.) R.C. Harris 7 A. angulosum (Müll. Arg.) R.C. Harris 8 A. biforme (Borrer) R.C. Harris 9 A. complanatum (Makhija & Patw.) R. C. Harris 10 A. consobrinum (Nyl.) Aptroot 11 A. glaucescens (Müll. Arg.) R.C. Harris 12 A. indicum (Makhija & Patw.) R. C. Harris 13 A. palavanum (Vain.) R.C. Harris 14 A. subnexum (Nyl.) R.C. Harris 15 A. tarmugliense (Makhija & Patw.) R.C. Harris 16 A. terminatum (Nyl.) R.C. Harris 17 A. ubianum (Vain.) R.C. Harris 18 Monoblastia pellucida Aptroot Porinaceae 19 Clathroporina mastoidea (Ach.) R.C. Harris 20 Porina americana Fée 21 P. atlantica (Erich.) P. M. Jørg. 22 P. chrysophora (Stirt.) R. Sant. 23 P. conica R. Sant. 24 P. epiphylla Fée 25 P. exserta Müll. Arg.* 26 P. internigrans (Nyl.) Müll. Arg. 27 P. interestes (Nyl.) Harm. 28 P. karnatakensis Makhija, Adaw. & Patw. 29 P. nitidula Müll. Arg. 30 P. ochrostoma Makhija, Adaw. & Patw. 31 P. pallescens R. Sant. 32 P. rufula (Kremp.) Vain. 33 P. siamensis P.M. McCarthy* 34 P. subcutanea Ach. 35 P. subhibernica Upreti Habitat Sl. Family and species No. South Goa Growth form North Goa CR CR CR CR CR C C C C C - - - - - - - - - + - - - + + + + - - - - - + + - CR CR CR CR CR CR CR CR CR CR CR CR CR C C C C C C C C C C C C C - - + + + + - - - + + - + + + - + - + - - - + + + + + + - + - - - + - + - CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR C C C F F F C C C F F C F F S C C - + + - - + - - - + + - - - + + + + + - - + + + + + + + + + + + + + + - - - - + + + + - CR S - - + - - - - - - - - - + - - - - - - CR S CR F - + - - - - - - - - + - + - - + + - - - - + - - CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR - + + - + - - + - - + - + + - + - + + + + + - + + - + - + + + + + + + + - - + - - - + + - C C C C C C C C C C C C C C C C C - 892 RANDIVE ET AL 56 P. ochraceoflava (Nyl.) R. C. Harris 57 P. oculata A. Singh & Upreti 58 P. quassiaecola (Fée) Fée 59 P. pyrenastrospora Aptroot* 60 P. rinodinospora Aptroot * 61 P. subumbilicata (C. Knight) Aptroot Strigulaceae 62 Strigula antillarum (Fée) Müll. Arg. 63 S. concreta (Fée) R. Sant. 64 S. janeirensis (Müll. Arg.) Lücking 65 S. nitidula Mont. 66 S. phyllogena (Müll. Arg.) R.C. Harris 67 S. smaragdula Fr. 68 S. subelegans Vain. 69* S. subtilissima (Fée) Müll. Arg. Trypetheliaceae 70 Astrothelium luridum (Zahlbr.) Aptroot & Lücking 71 A. meristosporum (Mont. & Bosch) Aptroot & Lücking 72 A. scoria (Fée) Aptroot & Lücking Marcelaria benguelensis (Müll. Arg.) Aptroot, Nelsen 73 & Parnmen 74 Nigrovothelium bullatum Lücking, Upreti & Lumbsch 75 N. tropicum (Ach.) Lücking, M.P. Nelsen & Aptroot 76 Trypethelium eluteriae Spreng. 77 T. plicatorimosum Mahija & Patw. Verrucariaceae 78 Endocarpon subrosettum A. Singh & Upreti 79 Staurothele fissa (Taylor) Zwackh CR CR CR CR CR CR C C C C C C + - + - + - - + - - + + - + - + + + - - + - + - - - + - - - CR CR CR CR CR CR CR CR F F F F F F F F - - - - - - - - - + - - - + + + + + + + + - - - - - - CR C CR C CR C - - - - - - - - - - - - + + - - + - - - - CR C - - - - + - - - - - - - + - - - - - - CR CR CR CR C C C C + - - - - - - - + - - - - + - + + + + - - + + - - - - SQ S CR S - - + - - + - - - - - - - - - - - - - - The studied specimens belonged to nineteen localities; eleven were from North Goa, while eight localities were from South Goa. The maximum diversity of pyrenocarpous lichens is recorded in South Goa with 71 species, while North Goa recorded 29 species and 21 species are common between the two districts. It may be noted that South Goa has a more dense forest area which is also fairly well explored compared to North Goa. Among the different localities, Cotigao Wildlife Sanctuary (CWS) is the most extensively surveyed and has maximum diversity of pyrenocarpous lichens with 48 species, while Bhagwan Mahavir Wildlife Sanctuary (BMWS) has 17 and Valpoi has eight species. Ponda and Quinamol localities are poorly represented by pyrenocarpous lichens with one species each. Compared to other sites Ponda and Quinamol are anthropogenically more disturbed which may be the reason for low pyrenocarpous lichen diversity here. Distribution and ecology:—This species is found growing on the bark of Terminalia paniculata Roth in Cotigao Wildlife Sanctuary between altitude of 100– 200 m. Earlier, this species was reported from Thailand, Christmas Island, the Northern Territory, eastern Queensland, Taiwan and Tahiti (27). New records Thallus saxicolous, rimose to areolate, smooth, matt, pale greyish to green, 20–80 μm thick, ecorticate; prothallus present, grey to black. Perithecia numerous, mostly solitary, semi-immersed to ± superficial, convex to hemispherical or subconical, 0.2–0.6 mm diam., greenish-brown to black, not overgrown by the thallus; ostiole inconspicuous or in a shallow depression; centrum subglobose to ovate, 0.10–0.25 mm diam.; excipulum hyaline to pale brown or yellowish-brown, 10–15 μm thick, subhymenium 20–40 μm thick; paraphyses unbranched, 0.8–1 μm wide, periphyses absent. Asci cylindrical to elongate, 72–86 × 8–9 μm, ascospores 3-septate, cylindrical to fusiform, 14–24 × 3–4.5 μm, lacking perispore. Pycnidia present. (Fig. 2B). 1. Porina exserta Müll. Arg. in Flora 71: 548. 1888. Thallus corticolous, continuous to rimose, pale brownish to green, smooth to minutely rugulose, 35– 80 μm thick, ecorticate; prothallus lacking. Perithecial verrucae, hemispherical to subglobose, 0.35–0.96 mm diam., brownish to blackish, apex rounded; ostiole usually inconspicuous; excipulum 20–30 μm thick, pale orange-brown, centrum 0.2–0.4 mm wide; subhymenium 15–35 μm thick; paraphyses unbranched, 0.7–1.2 μm wide, periphyses absent. Asci elongate to cylindrical 110–1160 × 14–18 μm, ascospores elongate to cylindrical, (11–)15–17 (–21) septate, 44–70 × 3.5–5.5 μm. Pycnidia absent. (Fig. 2A). Chemistry:—Thallus K–, C–, KC–, PD–, UV–; no lichen substance present in TLC. Specimen examined:—INDIA, Goa: South Goa, Cotigao Wildlife Sanctuary, Endrem, N14°59ʹ51.2ʺ, E 074°11ʹ52.1ʺ, 24 February 2018, P. Randive GU-L 813 (LWG Acc. no. 36250!), on the bark. Note:—Porina exserta closely resembles P. bellendenica Müll. Arg. in having similar perithecial verrucae and ascospores, but P. bellendenica differs in having a black basal layer and 15 septate ascospores. 2. Porina siamensis P. M. McCarthy in Lichenologist 31(3): 242. 1999. Chemistry:—Thallus K–, C–, KC–, PD–, UV–; no lichen substance present in TLC. PLANT SCIENCE TODAY 893 Fig. 2. Habit (a), perithecial section (b) and ascospres (c) of newly recorded pyrenocarpous lichens. A. Porina exserta, B. P. siamensis, C. Pyrenula dissimulans. D. P. pyrenastrospora. E. P. rinodinospora (Scale bars: habit A, C–E 2 mm, B 5 mm; perithecial section A–E 200 µm; ascospores A. 50µm, B – E 30 µm). Distribution and ecology:—Porina siamensis was found growing on an open, exposed rock in Cotigao Wildlife Sanctuary at about ±200 m. Previously this species was known from Thailand (28). diam.) and ascospores are slightly larger (16–32 × 4–6 μm). In comparison, P. fortunata differs in having seven septate ascospores of size 18–33 × 5–4 μm and perithecia of 0.3–0.4 mm diam. Specimen examined:—INDIA, Goa: South Goa, Cotigao Wildlife Sanctuary, Cuncolim, Agonda, N15°13ʹ13.0ʺ, E073°97ʹ99.5ʺ, 28 December 2016, P. Randive GU-L434 (LWG Acc. no. 36251!), on rock. 3. Pyrenula dissimulans (Müll. Arg.) R.C. Harris in More Florida lichens, Incl. 10 Cent Tour Pyrenol.: 110. 1995. Note:—This species is similar to Porina chlorotica (Ach.) Müll. Arg. and P. fortunata P. M. McCarthy & Etayo in having black or dark reddish-brown perithecia. P. chlorotica also has three septate ascospores, but its perithecia are smaller (0.2–0.3 mm Thallus corticolous, corticated, smooth, continuous, brownish to yellowish, lacking pseudocyphellae. Perithecia simple, dispersed, conical, emergent, 0.25– 0.6 mm diam., black; ostioles apical black; hamathecium hyaline. Asci cylindrical to clavate, 4–8 spored, ascospores muriform, 25–55 × 14–22 μm, 894 RANDIVE ET AL lumina mostly rounded, at least in the central part of ascospores. Pycnidia not seen. (Fig. 2C). Chemistry:—Thallus K–, C–, KC–, PD–, UV–; no lichen substance present in TLC. Distribution and ecology:—This species was found growing on the tree bark of T. paniculata in the Goa University campus approximately at an altitude of 100 m. Earlier, this species was known from North and South America (29-31). Specimens examined:—INDIA, Goa: North Goa, Goa University campus, 02 October 2016, P. Randive, GUL305 (LWG Acc. no. 36252!) on bark, GU- L318 (LWG Acc. no. 36253!) on bark. Note:— Pyrenula dissimulans is close to P. oleosa R.C. Harris in having corticated thallus, apical ostioles and lacking oil inspersion in hamathecium, but the old ascospores in the latter species are filled with oil. 4. Pyrenula pyrenastrospora Aptroot in Biblthca Lichenol. 64: 165, 1997. Thallus corticolous, corticated, smooth, brownish, lacking pseudocyphellae. Perithecia mostly aggregated with fused walls, conical, 0.3–0.6mm diam.; ostioles apical or when ecentric all opining the same direction, black; hamathecium not inspersed. Asci cylindrical to clavate, 4–8 spored, ascospores brown, 3-septate, fusiform, 16–25 × 6–10 μm, lumina in a straight line, terminal lumina directly against the exospores wall. Pycnidia absent. (Fig. 2D). Distribution and ecology:—This species was found growing on the bark of Ficus benghalensis L. in Coatigao Wildlife Sanctuary and en route to SattariValpoi between altitude 100–200 m. Previously this species was known only from Papua New Guinea (32). Specimens examined:—INDIA, Goa: South Goa, Cotigao Wildlife Sanctuary, Quinomol, N15°13ʹ130ʺ, E074°11ʹ88.7ʺ, 07 December 2016, P. Randive GU-L414 (LWG Acc. no. 36255!) on bark; North Goa, SattariValpoi, N15°31ʹ66.7ʺ E074°12ʹ41.1ʺ, 07 March 2017, P. Randive GU-L 548 (LWG Acc. no. 36256!), on bark. Note:—Pyrenula rinodinospora closely resembles Pyrenula maravalensis Vain. in having similar morphology, but the latter species differs by the shorter ascospores of 20–25 µm long (18). Key to the pyrenocarpous lichens of Goa 1a. Thallus foliicolous ………………………………….…. 2 1b. Thallus otherwise …………………………..………… 17 2a. Asci functionally unitunicate, entirely thinwalled, perithecia covered by thallus, ascospores transversely 3–9 septate….…..…….. 3 2b. Asci functionally bitunicate (fissitunicate), apically thick-walled, perithecia not covered by thallus, ascospores transversely 1-septate..… 10 3a. Perithecia with 5–10 (–12) setae, soft, usually decurved, narrowly acute or bristle-like, whitish or black with white distal halves, ascospores fusiform to narrowly oblong, 7 septate, 25–35 × 4–6 µm ………………. Trichothelium alboatrum 3b. Perithecia lacking setae ….…………………………. 4 4a. Area around the ostiole or perithecial surface rough, slightly papillose or tomentose ……...… 5 4b. Area around the ostiole and perithecial surface glabrous ……………...……...………………………..…. 6 5a. Perithecial wall colourless, ascospores 5 septate, fusiform, 22–30 × 4–5 µm ... Porina pallescens 5b. Perithecial wall upper part blackish, lower part brownish, ascospores 5 septate, fusiform, 20–32 × 4–5.5 µm ……………….……...…Porina nitidula 6a. Ascospores 3 septate …..…………………………….. 7 6b. Ascospores 7–9 septate ……...…..………………….. 8 7a. Perithecial apex conical, translucent, ascospores 18–27 × 3–5 µm …………………..… Porina rufula 7b. Perithecia apex rounded, not translucent, ascospores oblong, 14–20 × 2–4 µm …………………………………….Porina chrysophora 8a. Perithecia conical, apex with prominent, short cylindrical extension, ascospores narrowly fusiform to oblong, 34–48 7 × 4–6 µm ………………..………………………..… Porina conica 8b. Perithecia apex without any extension ……….. 9 9a. Perithecia conical to wart-shaped when mature, with black dot around ostiole, ascospores narrowly fusiform, 33–63 × 4–6 µm, white prothallus sometimes present ……………….……….. ………………………...………Porina karnatakensis Chemistry:—Thallus K–, C–, KC–, PD–, UV–; no lichen substance present in TLC. Distribution and ecology:—This species was found growing on the tree bark of Tectona grandis L.f. in Cotigao Wildlife Sanctuary at altitude ±200 m. Previously this species was known from Papua New Guinea (19). Specimen examined:—INDIA, Goa: South Goa, Cotigao Wildlife Sanctuary, Bela lake, N14°57ʹ23.7ʺ, E074°09ʹ09.3ʺ, 15 January 2018, P. Randive GU-L655 (LWG Acc. no.36254!), on bark. Note:—Pyrenula pyrenastrospora is closely related P. minarum Vain., which differs in having inspersed hamathecium and partly eccentric ostioles. 5. Pyrenula rinodinospora Aptroot in Lichenologist 44(5): 611–618. 2012. Thallus corticolous, corticate, smooth, continuous, thin, brownish, without pseudocyphellae. Perithecia simple, dispersed, conical, emergent, 0.3–0.5 mm diam., black, edges without thallus covering; ostioles black, apical; hamathecium hyaline, densely inspersed with oil droplets. Asci cylindrical to clavate, 8-spored, ascospores brown, 3-septate, fusiform, without constrictions, 20–30 × 11–12.5 μm, ends mostly pointed, lumina mostly quadrangular, angles blunt, terminal lumina elongated and not separated from the end wall by an endospore layer. Pycnidia not seen. (Fig. 2E). Chemistry:—Thallus K–, C–, KC–, PD–, UV–; no lichen substance present in TLC. PLANT SCIENCE TODAY 9b. Perithecia lens shaped to hemispherical, without dark spot, prothallus absent, ascospores oblong, 26–33 × 3–4 µm ..….. Porina epiphylla 10a. Ascospores large, 35–70 × 4–8 µm, often breaking into halves (each cell often with up to 3 secondary septa), oblong, with distinct constriction at septum ....Strigula janeirensis 20a. 21a. Distal cell of the ascospore enlarged.…………. 12 11b. Distal cell not enlarged ...…………………..……… 13 12a. Thallus subcuticular, bright green, ascospores irregularly biseriate, 15–25 × 4–6 µm …....………... …………………..…………….... Strigula antillarum 12b. Thallus epiphyllous, pale greenish to bluish grey, ascospores biseriate, 15–25 × 4–6 µm ………. ……………………..……………... Strigula subelegans 13a. Ascospores breaking into halves either inside or outside the asci …………..………………………...… 14 Thallus thin (8–15 µm), bright metallic green, margin effuse, with a thin blackish line, ascospores 8–12 × 2–3 µm ….Strigula nitidula 14b. Thallus thick (15–30 µm), pale greyish green, margin crenulate to lobulate, lacking blackish line, ascospores 8–12 × 2–3 µm ……………………….. …………………………………….… Strigula concreta 15a. Thallus elobate, photobiont cells rectangular to angular-rounded, perithecia pure black, conical, delimited from surrounding thallus, ascospores 9–12 × 2–3 µm ……………….Strigula phyllogena 15b. Thallus crenulate lobate to distinctly lobate . 16 16a. Thallus with distinct lobes leaving small to large interspaces, greenish-brown, perithecia completely exposed but covered by thin thallus layer, ascospores oblong acicullar, with a slight constriction at septum, 10–18 × 2–3 µm .…………. ……..….………………………..Strigula subtilissima 16b. Thallus entire to crenulate or lobulate, sometimes whole thallus lobate-laciniate, bright green, perithecia immersed to erumpent, covered by algiferous thallus tissue up to ostiolum, ascospores 14–24 × 4–6 µm …….……….. …………………………………..Strigula smaragdula 17a. Thallus saxicolous ……………………………………. 18 22a. Thallus squmulose, squmules imbricate, up to 2.5 mm wide, perithecia 1–6 per squamule, ascospores 2 per asci, brown, muriform, 30–40 × 12–14 µm …………... Endocarpon subrosettum 18b. Thallus crustose, continuous to areolate …… 19 19a. Ascospores muriform, 2 per acus, brown, 70–90 × 15–25 µm, perithecia sunken, algal cell in the hymenium globose, thallus rimose areolate, greyish brown …………......… Staurothele fissa 19b. Ascospores transversely septate, hyaline ….. 20 Ascospores hyaline …………………………………..23 22b. Ascospores greyish brown to brown …….…… 57 23a. Ascospores simple, subglobose to ellipsoid, surface ornamented with cristae, 9–13 × 7–9 µm, thallus slightly squamulose ………..………………..... ……….………….………..….Monoblastia pellucida 23b. Ascospores septate …………………………………...24 24a. Ascospores transversely septate ….……………. 25 24b. Ascospores muriform ……………………..……….. 56 25a. Ascospores 1-septate ………….…………………….. 26 25b. Ascospores 3 or more septate..………………….. 41 26a. Perithecial wall hyphal, containing bark cells, pseudoparaphyses branched, but not anastomosing …………………………………..…….. 27 26b. Perithecial wall cellular, lacking bark cells, pseudoparaphyses slender, branched and anastomosing especially above the asci ...….. 30 27a. Ascospores 2 per asci, 1-septate, sometimes constricted at both ends and appearing as 3 septate, large (40–60 × 15–25 µm) …………….….….. ……………..…………………….. Arthopyrenia finkii 27b. Ascospores 8 per asci ………………………...……. 28 28a. Distal cell of the ascospore is larger, ascospore 30–45 × 14–18 µm, perithecia solitary………………. ……………………………….Arthopyrenia nidulans 28b. Both the cells are almost equal in size …...…. 29 29a. 17b. Thallus corticolous ………………………………….. 22 18a. Ostiole conspicuous, periostiolar region brown to black, perithecia 0.5 – 0.8 mm diam., 7–9 septate, 23–34 × 5–9 µm, fusiform with rounded ends …………………….…….. Porina subinterstes 21b. Ostiole usually inconspicuous, periostiolar area pale to dark brown or blackish, perithecia 0.30.8 mm diam., ascospores 7 septate, 24–44 × 4–7 µm, cylindrical or narrowly obclavate…………….. ………………………………...…….Porina tetracerae 13b. Ascospores not breaking ………………………….. 15 14a. Ascospores 3 septate, cylindrical to fusiform, 14– 24 × 3–5 µm, perithecia 0.2–0.6 mm diam., greenish-brown to black, not overgrown by the thallus, ostiole inconspicuous or in a shallow depression …………………..…. Porina siamensis 20b. Ascospores 7–9 septate …………………………….. 21 10b. Ascospores small, up to 25 µm long ….……….. 11 11a. 895 Ostiole mammilate, perithecia solitary to 2–3 aggregate, ascospores 25–43 × 6–11 µm……………. ……………………………...Arthopyrenia indusiata 29b. Ostiole not mammilate, perithecia solitary, ascospores 10–18 × 5–7 µm ………..……..……………. ………..…………….....….. Arthopyrenia alboatra 30a. Thallus UV+ ……………………………………...…….. 31 30b. Thallus UV– ...…………………….……………………. 35 31a. Distal cell of the ascospores is larger, ascospores 23–30 × 7–10 µm, perithecia single, ostiole eccentric .………………..……………………………………. …………………………...Anisomeridium ubianum 31b. Both the cells are of almost equal size ...…….. 32 896 32a. RANDIVE ET AL Ostiole eccentric, perithecia single, ascospores 27–35 × 8–10 µm . Anisomeridium angulosum 44a. 32b. Ostiole apical ………………………………...………... 33 33a. Ascospores smaller, 14–19 × 4–65 µm, narrowly fusiform, ascus mostly biseriate……………………... ……………………...Anisomeridium albidoatrum 33b. Ascospores >20 µm long ……………….………….. 34 34a. Ascospores 25–30 × 9–13 µm, elliptical to broadly elliptical, ascus mostly uniseriate……….. ……………………..Anisomeridium consobrinum 34b. Ascospores 30–45 × 13–16 µm, ovoid to broadly ovoid, ends somewhat pointed ……………………….. ……………………………..Anisomeridium indicum 35a. Ascospores 2–4 per ascus, 40–66 × 10–17 µm, perithecia solitary, ostiole black with white wide pore……...Anisomeridium tarmugliense 44b. Thallus smooth to uneven, olive-green to yellowish-brown, perithecia 0.2–0.3 mm diam., prominent to sessile, subglobose to barrelshaped with a flattened top, ostiolar area greyish, ascospores fusiform-ellipsoid, 20–25 × 7–10 μm …………..… Nigrovothelium tropicum 45a. Distal cell of ascospore larger in size ….……… 37 36b. Both the cells of ascospores are of almost equal in size ………………………..………..…………………. 38 37a. Ascospore 3 septate, perithecia solitary or 2–3 aggregated, ostiole apical, umbonate, 19–23 × 5– 7 µm, thallus endophloeodal…………………………... ……………………………….… Arthopyrenia grisea 45b. Ascospores more than 3 septate …………………. 46 46a. 35b. Ascospores 8 per ascus ………………………..…... 36 36a. Thallus strongly verrucose-bullate, olive-green to brownish, perithecia 0.2–0.3 mm diam., sessile, stioles visible as tiny black dots, ascospores fusiform, 20–25 × 8–10 µm …………….. …………………………...Nigrovothelium bullatum Perithecia highly crowded to pseudostromatic …………..…………………………………...…..…………. 47 46b. Perithecia solitary or aggregated, never in pseudostroma, but in verrucae ……...…....…… 48 47a. Ostiole eccentric, ascospores 19–30 × 7–12 µm ……...…….………… Anisomeridium palavanum 37b. Ostiole apical, ascospores 16–21 × 4–7 µm ………………..…………... Anisomeridium biforme Perithecia in diffuse pseudostroma, brownblack, 0.5–1.0 mm, basally covered by thallus but upper part exposed, flattened and discshaped, ascospores 12–17-septate, fusiform, 45– 55 × 6–8 μm, thallus olive-green to yellowish ……..…………… Trypethelium plicatorimosum 38b. Ostiole eccentric ……………………………………….39 47b. Perithecia in distinct pseudostroma, brownish to dark brown, covered by yellow pigment, 1–2 mm diam., ascospores 9–13-septate, fusiform, 37–52 × 8–11 μm…………………………………………….. ……….…….…………….... Trypethelium eluteriae 39a. 48a. 38a. Ostiole apical, ascospores 23–27 × 9–11 µm, fusiform, biseriate or irregularly arranged in ascus ….……………... Anisomeridum subnexum Ascospores granular ornamented, fusiform to narrowly ovoid, 19–30 × 6–8 µm……………………... ……………….……..Anisomeridium terminatum Ostiole K+ reddish, perithecia 0.45-0.6 mm diam., ascospores 7 septate, elliptic, 19–30 × 4–7 μm …………………….Clathroporina mastoidea 39b. Ascospores smooth ……...…………………………...40 48b. Ostiole K- …………...…………………………………... 49 40a. Ascospores fusiform, with pointed ends, 30–37 × 9–11 µm ………... Anisomeridium glaucescens 49a. 40b. Ascospores 30–40 × 6–8 µm …………………………….. …………………….Anisomeridium complanatum 50a. 41a. 42a. Perithecia in distinct or diffuse pseudostroma, laterally covered by thallus ……………..…….…. 43 42b. Perithecia solitary to crowded, fully exposed, black ………………………...….……………………...… 44 43a. 49b. Ascospores 6–13 septa ……………………………….51 Ascospores lumina diamond-shaped ……….... 42 41b. Ascospores lumina rectangular-shaped ………45 Ascospores >13 septa ……………………………….. 50 Perithecia brownish blackish, ostiole inconspicuous, ascospores 11–21 septa, elongate to cylindrical, 47–70 × 4–6 μm…………………………. …………………………………………….Porina exserta 50b. Perithecia greenish grey, area around ostiole black, ascospores 13-16 septate, straight to curved with pointed ends, 50–70 × 5–7 μm …………………….………….… Porina subhibernica 51a. Hamathecium inspersed with oil …………………………..……….……………….…………. 52 Perithecia in diffuse stroma, upper portion blackish brown with whitish rim, erumpent, 0.30.5 mm diam., hamathecium inspersed, ascospores 3 septate, 15–27 × 7–10 µm, fusiformellipsoid ………………...…...Astrothelium scoria 51b. Hamathecium not inspersed ………………………………...………….….….………….. 53 43b. Perithecia in distinct stroma, upper portion dark, erumpent to prominent, 0.8-1.5 mm broad, ostiolar region internally with yellow pigment, hamathecium clear, ascospores 7–11 septate, fusiform, 55–88 × 13–28 µm………………... …………………..……………Astrothelium luridum 52b. Ascospores 11–12 septate, fusiform, 89–92 × 16– 20 μm, ostiolar region black……………………………. …………..………………………....Porina americana 52a. 53a. Ascospores 7–9 septate, oblong ellipsoid, 60–85 × 12–20 μm, ostiolar region brown black……………. ……………………………………...Porina subcutanea Ostiolar region yellow to light brown, surrounded by black priostiolar rim, ascospores PLANT SCIENCE TODAY 7–9 septate, fusiform with acute ends, 43–50 × 4– 6 μm ……..………………....... Porina ochrostoma 63a. 53b. Ostiolar region variously coloured, peri ostiolar rim lacking ………………………..………….………… 54 54a. Ostiolar region brown-black, ascospores 7–8 septate, fusiform, acute ends, 32–48 × 4–7 μm….. …………..…………………………….Porina interestes Ostiolar region orange, pinkish, reddish to brownish, rarely dark brown, ascospores 7–9 (– 13) septate, fusiform with rounded ends, 44–80 × 7–13 μm ………………………...… Porina atlantica 55b. Ostiolar region orange-brown to black to black, ascospores fusform round ended, 45–60 ×11–18 μm …………………………….. Porina internigrans 56a. Thallus olive-green, yellow-orange pruinose, UV+, perithecia flattened top, solitary to irregularly grouped and confluent, covered with yellow-orange pigment, hamathecium densely inspersed, ascospores fusiform-ellipsoid, 50–80 × 17–23 µm ……………Marcelaria benguelensis Thallus pseudocyphellate, yellow-brown buff, perithecia solitary or 1–4 coalescing, ostiole indistinct, ascospores oblong ellipsoid, 32–45(– 60) × 15–27 µm ………….…… Pyrenula immissa 63b. Thallus lacking pseuodcyphellae ………………. 64 64a. 54b. Ostiolar region otherwise, ascospores much larger ……………………….……………………..……… 55 55a. 897 Ascospore 27–50 × 15–18 µm, oblong ellipsoid, perithecia solitary, completely embedded in thalline verrucae, ostiole indistinct, thallus ochre-yellowish brown …... Pyrenula oculata 64b. Ascospore <30 µm long ……………………………..65 65a. End lumina of the ascospore is elongated …..66 65b. All lumina ± rounded to angular …...…………. 67 66a. Thallus brownish to pale brown, ascomata emergent 0.3–0.5 mm diam., hemithecium densely impressed with oil droplets, ascospores 20–25 × 6–10 µm .…… Pyrenula maravalensis 66b. Thallus ochraceous yellow to brownish, ascomata semiemergent 0.5- 1.0 mm diam., hemithecium without oil droplets, ascospores 20–30 × 11–13 μm .... Pyrenula rinodinospora 67a. Ostiole mamillate papillate, dull black, perithecia solitary, ascospores oblong ellipsoid, 17‒20×7‒9 µm, thallus buff to yellow-brown ….. ………….………………………….Pyrenula mamillan 56b. Thallus light olive-green, pigmented pruina absent, UV+, flattened top, solitary to grouped, covered by thallus up to ostiole, hamathecium inspersed, ascospores oblong-fusiform, 120–220 × 25–40 µm, with thickened median septum…….. ………………….… Astrothelium meristosporum 67b. Ostiole otherwise, centrum I ± perithecia solitary or semi-solitary, ascospores ellipsoid, thallus brownish yellow to brown …...………. 68 57a. 68a. Ascospores transversely 3 septate ……..……… 58 57b. Ascospores submuriform to muriform ……… 74 58a. Perithecia in pseudostroma, 1–12 carpic, thallus yellowish-brown to brown, K+ red, ascospores ellipsoidal, 13–23 × 6–10 µm ……..………………... ………………………………..… Pyrenula leucotrypa 58b. Perithecia solitary or aggregated, not forming pseudostroma ……………..…..…..……...…………. 59 59a. Ascomata mostly aggregated, with fused walls but with separate ostioles, conical, 0.3–0.6 mm diam., ostioles apical, black, ascospores brown, 3 septate, fusiform 16–25 × 6–10 μm………………... …………………………Pyrenula pyrenastrospora 59b. Ascomata solitary ……………………………………. 60 60a. Ascospores red-brown, lumina becoming rounded when older, 15–20 × 9–10 µm, ellipsoid, hamathecium inspersed, thallus olive-brown shining…………….…….Lithothelium decumbens 68b. Psudocyphellae absent, ascocarp 1.0–2.5 (–3.0) mm in diameter, flat, convex to conicodepressed, centrum I -, ascospores 15–17 × 5–7 µm ……..………………....……. Pyrenula castanea 69a. Ostiole eccentric, perithecia solitary, ascospores 35–45 × 15–18 µm, terminal lumina separated from the exospore wall by endospore thickening ………...….………………………….. Pyrenula adacta 61b. Ostiole apical ……………………………………….…. 62 62a. Hamathecium inspersed ………..………………... 63 62b. Hamathecium not inspersed……………………….69 Thallus with pseudocyphellae, brownish to olive-green, ascospore fusiform, 22–38 × 8–15 μm, lumina rounded to somewhat angular …………..…….…….....…. Pyrenula quassiaecola 69b. Thallus lacking pseudocyphellae ….……...…… 70 70a. Terminal lumina mostly separated from exospore wall …….………………...……….………… 71 70b. Terminal lumina not separated from exospore wall ….………….....……………………………..………. 73 71a. 60b. Ascospores grey to brown, rarely red-brown and then lumina angular …...……..………..…… 61 61a. Psudocyphellae present, ascocarp 0.8–1.0 mm in diameter, conical- hemispherical, centrum I+ blue, ascospores 16–24 × 8–13 µm …………………... ………………...……………………..Pyrenula fetivica Perithecia slightly larger, 0.6–1.0 mm diam., ascospores oblong ellipsoid, 12–17 × 5–7 μm ……. ………………………………….…...Pyrenula brunnea 71b. Perithecia smaller ………….……………………….. 72 72a. Perithecia 0.2–0.6 mm diam., ascospores oblong ellipsoid, 17–22 × 9–10 μm …Pyrenula aspistea 72b. Perithecia 0.4–0.7 mm diam., ascospores oblong ellipsoid, 15–21 × 8–10 μm………………………………. …………………………………....Pyrenula aggregata 73a. End lumina elongated, ascospore 20–29 ×9–11 μm ……...………….….…. Pyrenula approximans 898 RANDIVE ET AL 73b. All lumina ± rounded to angular, ascospores 13– 16 × 7–8 μm ………………....… Pyrenula nitidula 74a. Ostioles eccentric, fused, perithecia usually in groups of 2–6, ascospores submuriform, with 5 transverse septa and few longitudinal septa, fusiform, with pointed ends, 24–32 × 10–13 μm, thallus yellowish to olive-green ……………............ ……………………………. Pyrenula subumbilicata 74b. Ostiole apical, not fused, perithecia mostly solitary ……………..…………..……………...……….. 75 75a. Thallus and perithecia yellow to orange, indistinctly to faintly papillate, ascospores ellipsoidal, 12–18 × 7–10 μm …………….…………….. ………………………….... Pyrenula ochraceoflava 75b. Thallus and perithecia otherwise ……………….76 76a. Old ascospores filled with orange oil, muriform, 25–30 × 10–15 μm, pantropical distribution …………………………………...… Pyrenula breutelii 76b. Old ascospores lacking orange oil, submuriform..…….………………….…………..…... 77 77a. Thallus yellow-brown, pseudocyphellate, UV+ yellowish, ostioles indistinct to mamillatepapillate, hamethecium not inspresed, ascospores 23–35 × 10–12 μm long ……………..…... …………………………...…… Pyrenula gibberulosa 77b. Thallus otherwise, ostioles apical, ascospores larger, up to 45–55 μm long ……………………... 78 78a. Lumina of the ascospore mostly round, ascospore 25–55 × 14–22 μm, ostioles apical, naked, black, thallus yellowishbrown……………………….Pyrenula dissimulans 78b. Lumina relatively large and angular, ascospore 24–46 × 9–20 μm, ostiolar region naked, brownish-black to black, thallus greenish straw coloured, yellow to yellow-brown …………………... …………………………………..Pyrenula leucostoma Discussion The semi-evergreen forest in the Goa region exhibits maximum diversity of pyrenocarpous species. The trees along the streams in moist, shady habitats bear luxuriant growth of pyrenolichens which mostly belong to Porina and Pyrenula species. The occurrence of pyrenocarpous lichens indicates the abundant presence of smooth-barked trees in the State. Further, the occurrence of a large number of foliicolous lichens indicates that the State has several healthy and undisturbed forests. Bhagwan Mahavir Wildlife Sanctuary as a whole and within Cotigao Wildlife Sanctuary, Avem, Bela to Zombolim,, Endrem to Tulshimol, Kuske waterfall area, Marlem, Nadkem and Ravan Donger are few such pristine habitats. With the addition of five new records, the total number of lichens species in Goa State increases to 165 species, out of which pyrenocarpous lichens represent almost half (48%). Ghats forests of Goa. It can be noted that among the protected areas within Goa, Cotigao Wildlife Sanctuary represents more number of pyrenocarpous lichens. Further, along with the forest areas within Goa, the coconut, arecanut orchards and several cultivated plants also provide suitable habitats for lichen growth. The present study will be highly useful for monographic studies on pyrenocarpous lichens of India or the world and for environmental monitoring studies in the area. Acknowledgements We are thankful to Director, CSIR-NBRI, Lucknow, for providing the laboratory facilities for the research, to Publication Ethics Committee of CSIR-NBRI for carefully scrutinizing the manuscript and approving it for publication with ms number MS/2020/08/03. Authors PR and MKJ are thankful to Goa University for financial assistance under the Ph.D. programme and to the Forest Department of Goa for permission to survey Cotigao Wildlife Sanctuary. PR is also thankful to DST for financial assistance under Women Scientist Scheme (SR/WOS-A/LS-354/2016). GKM is thankful to SERB for financial assistance under the NPDF scheme (PDF/2017/000356). Authors’ contributions PR collected fresh samples of pyrenocarpous lichens from Goa and carried out their preliminary identification. 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A world key to the species of Anthracothecium and Pyrenula. Lichenologist. 2012;44(1):5-53. https://doi.org/10.1017/ s0024282911000624 Publisher’s Note: Horizon e-Publishing Group remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. 20. Aptroot A, Ertz D, Salazar JAE, Gueidan C, Diaz JAM, Schumm F, Weerakoon G. Forty-six new species of Trypetheliaceae from the tropics. Lichenologist. 2016;48(6):609-38. https://doi.org/ 10.1017/s002428291600013x 21. Aptroot A, Cáceres MEDS. New Trypetheliaceae from the Amazon basin in Rondônia (Brazil), the centre of diversity of the genus Astrothelium. Lichenologist. 2016;48(6):693-712. https:// doi.org/10.1017/s0024282915000584 22. Aptroot A, Lücking R A revisionary synopsis of the Trypetheliaceae (Ascomycota: Trypetheliales). Lichenologist. 2016;48(6):763–982. https://doi.org/10.1017/S0024282916000487 at To cite this article: Randive P, Mishra G K, Nayaka S, Upreti D K, Janarthanam M K. Pyrenocarpous lichens in Goa with five new records to India. Plant Science Today. 2021;8(4):889–899. https://doi.org/10.14719/pst.2021.8.4.1265 Plant Science Today, published by Horizon e-Publishing Group, is covered by Scopus, Web of Science, BIOSIS Previews, Clarivate Analytics, etc. See https://horizonepublishing.com/journals/index.php/PST/indexing_abstracting