Angiactis, a new crustose genus in the Roccellaceae, with species from Bermuda, the Gala´pagos Islands and Australia

ANDRE´ APTROOT ABL Herbarium, Gerrit van der Veenstraat 107, NL-3762 XK Soest, The Netherlands e-mail: [email protected]

LAURENS B. SPARRIUS BIO.DIV, Vrijheidslaan 27, NL-2806 KE Gouda, The Netherlands e-mail: [email protected]

SCOTT LAGRECA Department of Botany, the Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom e-mail: [email protected]

FRANK BUNGARTZ Botany Department, Charles Darwin Research Station, Isla Santa Cruz, Gala´pagos; Postal Address: Charles Darwin Foundation, Av. 6 de Diciembre N 36-109 y Pasaje California, Post Box 17-01-3891, Quito, Ecuador e-mail: [email protected]

ABSTRACT. The new genus Angiactis Aptroot & Sparrius is described, with three species, A. bermudensis LaGreca sp. nov. from Bermuda, A. littoralis (Kantvilas) Aptroot & Sparrius comb. nov. from Australia and A. spinicola Aptroot & Sparrius sp. nov. from the Gala´pagos. The new genus differs from Lecanographa in the presence of a thalline exciple, and from Llimonaea in the non-corticate thallus and presence of a thalline exciple.

KEYWORDS. Angiactis, Arthoniales, Lecanographa, Llimonaea.

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During the last decade, a number of old crustose example, Egea and Torrente (1994) split Roccellaceae genera have been revised by various s.l. into several taxa, based on apothecium shape and workers, revealing new characters. As part of this pigments, and gave more attention to ascospore and process, several larger genera in the family have been ascus characters. As part of this work, related genera reduced in size, such as Enterographa (Sparrius 2004) such as Llimonaea (Torrente & Egea 1991) were and Plectocarpon (Ertz et al. 2006). During these described. studies, new circumscriptions of genera were made, More recently, Kantvilas (2004) described and many new species have been described. For Lecanographa littoralis, a species quite different from

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Table 1. Main differences between Angiactis and four related genera.

Angiactis Dirina Lecanographa Llimonaea Schismatomma

Thalline exciple present present absent present present Thallus cortex absent present absent present absent Ascus type Grumulosa Abietina Grumulosa Grumulosa Abietina Ascospore color hyaline hyaline hyaline dark brown hyaline Ascospore wall thick thick thin or thick thick thin Excipulum color carbonized carbonized carbonized carbonized hyaline Excipular KOH none none green or none green green or none reaction

Llimonaea and Lecanographa, but placed tentatively anastomosantibus, ascosporibus leptoseptatibus clavatis in the latter genus awaiting future examination. In 3–7-septatibus hyalinis gelatinosis. the following two years, we collected two similar TYPE: Angiactis littoralis (Kantvilas) Aptroot & species in coastal habitats. While visiting the Sparrius. Gala´pagos Islands for two weeks in 2005, the first Description. Thallus thick; cortex absent; author noticed that the spiny Scutia bushes near the medulla cretaceous; photobiont . seashore at the Charles Darwin Research Station were Ascomata irregularly rounded to elongate, with a covered with an unknown, crustose Roccellaceae. thick thalline margin; disc black, often white During fieldwork conducted by the first and last pruinose; excipulum and hypothecium dark brown authors in the following year, this remarkable lichen to black/carbonized, KOH-; hymenium not was found on other hosts at additional localities in inspersed; paraphysoids slender, anastomosing, non- the Gala´pagos, but always close to the sea. It was septate, not separating easily in water or KOH; asci of initially thought to be a new species of the small the Grumulosa-type, with KI+ blue inner wall and genus Llimonaea, but it differs in several characters. blue ring in the apical dome; ascospores hyaline, During a collecting trip to the Bermudas the same thin-walled, asymmetrical, clavate, 3–7-septate, year, the third author collected material from surrounded by a thin perispore. Conidiomata known maritime rocks which appeared to belong to the only from A. bermudensis, punctiform, immersed, same, undescribed genus. Here we formally describe black; conidia rod-shaped, non-septate, hyaline. this new genus as Angiactis in the Roccellaceae, with Chemistry. Thallus C+ and KC+ red (often in three species known, mainly differing in ascomatal patches), KOH-, PD-, UV- or UV+ violet or whitish. shape, chemistry and substrate. Gyrophoric acid and/or erythrin; sometimes traces of lecanoric acid or methylgyrophorate detected with TLC. MATERIAL AND METHODS Specimens for this study were collected in the Etymology. The name Angiactis is derived from the Greek ‘‘acceion’’ (receptacle) and ‘‘a toz’’ field or borrowed from HO. TLC was carried out [ using solvent A (Orange et al. 2001). Measurements (probably also meaning ‘‘shaped’’, here used as in the of ascospore sizes are based on at least 25 ascospores name of the related genus Lecanactis) referring to the per specimen. fruiting bodies covered by a thalline excipulum. Discussion. Genera with elongate (lirelline) Angiactis Aptroot & Sparrius, gen. nov. apothecia are scarce and most, like Lecanographa and Genus novum Roccellacearum, Lecanographae et , have ascospores with thick septa and lack Llimonaeae affine, sed ab ea differt essentialiter thallo a thalline margin around the apothecia. Angiactis is crustaceo cretaceo non corticato, apotheciis rotundis vel probably closely related to Llimonaea, which has a elongatis, marginibus thallinibus, excipulo corticate thallus, ascospores that become brown and hypotecioque fusco vel niger, KOH non reagens, ascis an excipulum that is KOH+ green. Lecanographa, in cylindricis, Grumulosa-typis, paraphysibus gracilibus which the type species of Angiactis was first

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Table 2. Main differences between the three known species of Angiactis.

A. bermudensis A. littoralis A. spinicola

Ascomatal shape rounded to stellate rounded, unbranched elongate, often forked Ascomatal position immersed sessile, mostly constricted below sessile, mostly adnate Chemistry (major substances) erythrin only gyrophoric acid only erythrin and gyrophoric acid Ascospore length 17–28 mm 11–24 mm 17–25 mm Ascospore septa 3–4(–6) 3–6(–7) (3–)4–6 Ecology saxicolous on limestone; saxicolous on limestone, and corticolous; coastal coastal also on dead twigs; coastal Distribution Bermuda Australia Gala´pagos described—albeit with considerable hesitation and a often wider underneath the disc than on the edges, lengthy discussion of the differentiating characters dark reddish brown to black in section, with crystals; (see discussion in Kantvilas 2004)—has no thalline hypothecium dark reddish brown, ca. 40 mm tall, margin around the true exciple. An overview of the with crystals, often extending downwards into the differences between Angiactis, Dirina, Lecanographa, thallus; hymenium pale reddish brown, ca. 50–60 mm Llimonaea and Schismatomma are given in Table 1. tall, sometimes eroding away in the center with age; A fifth genus, Sigridea differs in having a corticate paraphysoids ca. 1 mm thick, highly branched and thallus containing psoromic acid in all species and anastomosing, tips not swollen; epithecium ca. the Abietina-type ascus. 10 mm tall, red-brown, with brown crystals of ca. 2– We studied a Tasmanian specimen of the type 3 mm diam., KOH-; asci 40–50 3 11–15 mm, species, growing on soft limestone rock, and found it cylindrical, 8-spored, of the Grumulosa-type; to be close to the following new species from ascospores hyaline, narrowly clavate to fusiform, Bermuda and the Gala´pagos Islands. The main thin-walled, (17–)25(–28) 3 5–10 mm, 3–4(–6)- differences between all three species are listed in septate, perispore present, ca. 1 mm thick. Table 2. Conidiomata black, punctiform, immersed, ca. 100– 150 mm in diameter, with a brown-orange pigment in Angiactis bermudensis LaGreca, sp. section, KOH-; conidia rod-shaped, hyaline, ca. 4–6.5 nov. Figs, 1, 2A 3 1.5(–2.5) mm. Angiactis rupes calcares incolens, thallo cretaceo, Chemistry. Thallus C+ red (at least around apotheciis rotundis vel lirelliformis, ascosporibus vulgo apothecia or apothecium margins), KOH-, PD-, UV+ triseptatibus vel tetraseptatibus 17–28 mm longis 5– violet. Erythrin plus traces of two unidentified 10 mm latis, erythrinum continens. compounds were found with TLC. These TYPE: BERMUDA. WARWICK PARISH: Stonehole Bay unidentified compounds were visible in solvent A National Park, off South Road, on shaded, dry only. No gyrophoric or lecanoric acid was detected. underhangs of limestone cliffs along the coast, Etymology. Named after the island group where 32u159180N, 64u489480W, 17 m alt., 16 Aug 2006, the type was collected. LaGreca 1543 (BM, holotype). Distribution and ecology. Endemic to the Description. Thallus to 4 cm in diam., Bermuda Islands. Saxicolous on limestone rock on continuous, scurfy-crustose, chalk-white; cortex shaded, dry underhangs on or near the coast. absent; medulla white, cretaceous, with crystals; Additional specimens examined. BERMUDA. ST. photobiont Trentepohlia; prothallus indistinct. GEORGE’S PARISH: Town of St. George, The Unfinished Ascomata ellipsoid to elongate, sometimes forked or Church on Folly Road, on brick/mortar on inside stellate, solitary (not confluent), (0.2–)0.5(–1.0) 3 wall of church, 10 Aug 2006, LaGreca 1544 (BM); St. (0.1–)0.3(–0.4) mm, immersed in thallus; disc pitch David’s Island, Great Head Park, on steep maritime black, sometimes white pruinose; thalline margin cliff adjacent to ‘‘The Crack,’’ 19 Aug 2006, LaGreca thick, irregular, white; excipulum to 10 mm wide, 1545 & J. McCabe (ABL, BM, HO).

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Figure 1. Angiactis bermudensis, LaGreca 1544 (BM). Habit of the thallus with ascomata. Scale: each increment 5 1 mm.

Angiactis littoralis (Kantvilas) Aptroot & containing crystals; photobiont Trentepohlia; Sparrius, comb. nov. Figs. 2B, 3 prothallus indistinct. Ascomata numerous, adnate to Lecanographa littoralis Kantvilas, Symb. Bot. Upsal. basically constricted, rounded, unbranched, to ca. 34(1): 197. 2004. 0.3–1 mm diam.; disc black, coarsely white pruinose; TYPE: AUSTRALIA. WESTERN AUSTRALIA: Cape thalline margin prominent, 0.06–0.1 mm wide of the Leeuwin, on dead Melaleuca twigs in coastal same color and structure as the thallus, ecorticate; heathland, 34u229S, 115u079E, 9 Oct 1992, Kantvilas excipulum dark brown, KOH-, to 10 mm wide; & Jarman 320/92 (HO, holotype; BM, GZU, PERTH, UPS, epithecium olive-brown, 12–20 mm thick, KOH-; isotypes). hypothecium brown to black, often extending Description. Thallus white to pale yellowish downwards, upper layer (subhymenium) hyaline to cream, continuous, scurfy-crustose to byssoid; cortex orange-brown, 25–80 mm tall, KOH-; hymenium 60– absent; medulla white, cretaceous, to 1 mm thick, 75(–100) mm tall, hyaline, K/I+ pale blue;

Figure 2 A–C. Angiactis ascospores. A. A. bermudensis, LaGreca 1544 (BM). B. A. littoralis, Kantvilas 311/97 (HO). C. A. spinicola, Aptroot 63413 (CDS, holotype). Scale: bar 5 5 mm.

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Figure 3. Angiactis littoralis, Kantvilas & Jarman 320/92 (BM, isotype). Habit of the thallus with ascomata. Scale: bar 5 5 mm. paraphysoids slender, ca. 1 mm wide, anastomosing; Angiactis spinicola Aptroot & Sparrius, sp. asci cylindrical, 65–90 3 10–16 mm, approximating nov. Figs. 2C, 4 the Grumulosa-type and lacking any discernible Angiactis epiphyticus, thallo cretaceo, apotheciis amyloid sheath between the inner ascus wall and the lirelliformibus, ascosporibus vulgo tetraseptatibus 17– ascoplasm visible in younger asci; ascospores 3–6(–7)- 25 mm longis 3–6 mm latis, acidum gyrophoricum septate, hyaline, narrowly ellipsoid to fusiform, methylgyrophoratum erythrinumque continens. slightly curved, 11–24 3 4–8 mm, surrounded by a TYPE: ECUADOR. GALA´ PAGOS ISLANDS: Santa Cruz 0.5–1 mm thick perispore. Conidiomata not observed. Island, coastal cliffs E of Puerto Ayora near Charles Chemistry. Thallus C+ reddish, KOH-, PD-, Darwin Research Station, on spines of Scutia spicata, UV-. Gyrophoric acid and lecanoric acid (trace) 0u449450S, 90u179390W, 20 m alt., 29 May 2005, detected with TLC. Aptroot 63413 (CDS 30168, holotype; ABL, BM, isotypes). Distribution and ecology. Endemic to Australia Description. Thallus white to a bit ochraceous, (Tasmania and Western Australia). continuous, to 5 cm diam.; cortex absent; medulla Note. Many of the characters above are derived white, cretaceous, to 1 mm thick, with crystals; from Kantvilas (2004). The drawing of the ascomata photobiont Trentepohlia; prothallus indistinct. in Kantvilas (2004) shows structures that can be Ascomata numerous, sessile, rounded to usually interpreted as interhymenial strands, but these are in elongate, often forked, ca. 0.1–0.5 3 0.3–1.0 mm; fact folded margins of the apothecia and have no disc black, sometimes white pruinose; thalline taxonomic value. The same structures can also margin ca. 0.2 mm wide of the same color and sometimes be found in Angiactis bermudensis. structure as the thallus; excipulum dark brown, Additional specimen examined. AUSTRALIA. KOH-, to 10 mm wide; epithecium red-brown, TASMANIA: Flinders Island, Cave Beach, on soft tertiary KOH-; hypothecium dark brown to black, often limestone, 1997, Kantvilas 311/97 (HO). extending downwards, upper layer (subhymenium)

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Figure 4. Angiactis spinicola, Aptroot 63413 (CDS, holotype). Habit of the thallus with ascomata. Scale: bar 5 1 mm.

hyaline to orange-brown, 40–100 mm tall; Selected additional specimens. ECUADOR. hymenium ca. 50 mm tall, hyaline; paraphysoids ca. GALA´ PAGOS ISLANDS: Santa Cruz Island, Puerto Ayora, 1 mm wide, anastomosing; asci clavate-cylindrical, near the Charles Darwin Research Station, Grumolosa-type, 8-spored, 70–90 3 15–20 mm; 00u449320S, 90u189100W, ca. 1 m alt., on bark of ascospores (3–)4–6-septate, hyaline, clavate, Rhizophora twigs in mangrove swamp, 24 May 2005, straight, 17–25 3 (3–)5–6 mm, surrounded Aptroot 63065 (CDS 29793); along the S coast of Santa by a 1–2 mm thick perispore. Conidiomata not Cruz close to the Charles Darwin Research Station, observed. 00u449390S, 90u189020W, ca.1 m alt., among bare Chemistry. Thallus C+ red (at least around coastal lava rocks with Cryptocarpus pyriformis, apothecia or apothecium margins), KOH-, PD-, UV+ occasionally in between on bark of Cryptocarpus whitish. Erythrin, gyrophoric acid and methyl pyriformis twigs, 10 Feb 2006, Bungartz 3424 (CDS gyrophorate detected with TLC. 27130). Etymology. Named after the preference of this species for spines and twigs. ACKNOWLEDGMENTS Distribution and ecology. Endemic to the Our friend Leo Spier is thanked for doing the TLC work for Gala´pagos. This species is common and abundant on Angiactis spinicola. Gintaras Kantvilas provided us with the additional specimen of A. littoralis. Alan Tye is warmly thanked spines of Scutia spicata, twigs of mangroves and twigs for his invitation to investigate the lichen flora of the of Cryptocarpus pyriformis on the coastal cliffs close Gala´pagos. We are grateful to Norman Robson for proofread- to the Charles Darwin Research Station, Santa Cruz. ing the Latin descriptions. Pat Wolseley assisted with the initial It was not encountered during trips to other islands examinations of A. bermudensis, and Wolfgang Sterrer assisted of the Gala´pagos. with the photography of that species. Keith Babuszczak is

The Bryologist bryo-111-03-01.3d 1/2/08 13:21:45 6 Aptroot et al.: Angiactis (Roccellaceae), gen. nov. 0 thanked for his assistance with the digital image of A. Ertz, D., C. Christnach, M. Wedin & P. Diederich. 2006. A bermudensis. Grants for fieldwork to SLG from the Bermuda world monograph of the genus Plectocarpon (Roccellaceae, Biological Station for Research, Inc. (BBSR) and the Natural Arthoniales). Bibliotheca Lichenologica 91: 1–155. History Museum are gratefully acknowledged. James McCabe, Kantvilas, G. 2004. A contribution to the Roccellaceae in the staff of the Bermuda Aquarium, Museum & Zoo (BAMZ), Tasmania: new species and notes on Lecanactis and allied and the staff of the Bermuda Department of Conservation genera. Symbolae Botanicae Upsalienses 34(1): 183–203. Services—especially Anne Glasspool, Wolfgang Sterrer, and Orange, A., P. W. James & F. J. White. 2001. Microchemical Jack Ward—are thanked for their assistance with the Bermuda methods for the identification of . British Lichen fieldwork. This publication is contribution number 1057 of the Society, London. 101 pp. Charles Darwin Research Station, and contribution number Sparrius, L. B. 2004. A monograph of Enterographa. Bibliotheca 137 of the Bermuda Biodiversity Project (BBP), Bermuda Lichenologica 89: 1–141. Aquarium, Natural History Museum and Zoo. Torrente, P. & J. M. Egea. 1991. Llimonaea, a new genus of lichenized fungi in the order Opegraphales (Ascomycotina). LITERATURE CITED Nova Hedwigia 52: 239–245. Egea, J. M. & P. Torrente. 1994. El ge´nero de hongos liquenizados Lecanactis (Ascomycotina). Bibliotheca Lichenologica 54: 1–205. ms. received July 2, 2007; accepted January 15, 2008.

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