The Lichenized Genus Cora (Basidiomycota: Hygrophoraceae) in Mexico: High Species Richness, Multiple Colonization Events, and High Endemism

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Plant and Fungal Systematics 64(2): 393–411, 2019 ISSN 2544-7459 (print) DOI: 10.2478/pfs-2019-0026 ISSN 2657-5000 (online)

The lichenized genus Cora (Basidiomycota: Hygrophoraceae) in Mexico: high species richness, multiple colonization events, and high endemism

Bibiana Moncada1,3, Rosa Emilia Pérez-Pérez2 & Robert Lücking3*

Abstract. In a continued effort to catalog the numerous phylogenetically detected and Article info predicted species of Cora in the Americas, we focus here on the diversity of the genus in Received: 23 Apr. 2019 Mexico and the phylogenetic relationships of the taxa present in this area. Based on pre- Revision received: 4 Oct. 2019 vious results and new collections, 12 taxa are recognized in Mexico, including eight new Accepted: 14 Oct. 2019 Published: 2 Dec. 2019 species and one new subspecies. The 12 taxa form 11 unrelated lineages within the genus, indicating multiple independent colonization from Central and South America. While the Associate Editor new subspecies is nested within a species known from the northern Andes in South Amer- Adam Flakus ica, the other species are all putative endemics for Mexico, resulting in endemism of 92% at species level and 100% at taxon level. Considering the rather narrow area of origin of the sequenced specimens in southeastern Mexico and the previously documented range of Cora including the northwestern part of the country, plus the underlying topography, we predict that the 12 species and subspecies now known represent only about 20% of the total richness of Cora in the country, and that many more endemic lineages are to be found in the western and northwestern parts (Sierra Madre Occidental). The new taxa from Mexico formally introduced in this study are Cora benitoana sp. nov., with a strongly projecting, cyphelloid hymenophore; C. buapana sp. nov., with elongate, finger-like and partly branched appendages on the lower medullary hyphae; C. dewisanti subsp. mexicana subsp. nov., with a marginally protruding hymenophore; C. guzmaniana sp. nov., with a partly setose lobe surface; C. ixtlanensis sp. nov., a phenotypically cryptic species similar but unrelated to C. terrestris; C. lawreyana sp. nov., with globose hyphal appendages; C. marusae sp. nov., a phenotypically cryptic species similar but unrelated to C. comaltepeca; C. totonacorum sp. nov., a phenotypically cryptic species similar but phylogenetically distant to C. davidia; and C. zapotecorum sp. nov., with a very thinly pilose lobe surface. Key words: Cora casasolana, Cora comaltepeca, Cora dulcis, Frullania, Rhizonema, Cen- tral Volcanic Belt, Isthmus of Tehuantepec, Sierra Madre de Chiapas, Sierra Madre Oriental

Introduction

With a share of about 1%, Basidiomycota form only 93 of which have so far been formally described, and a small proportion of all lichenized fungi (Lücking et al. over 450 species predicted (Lücking et al. 2013, 2014, 2017a). However, the genus Cora, until recently believed 2017b). Although Cora has been demonstrated to occur to comprise only a single species (Parmasto 1978; Chaves on islands in the South Atlantic, in tropical Africa and et al. 2004), has been shown to be hyperdiverse, with in Asia (Lücking et al. 2015, 2017b), over 95% of the nearly 200 species already detected using ITS barcoding, known species are restricted to the Americas, the genus being most diverse in the northern Andes but also found as far north as Mexico and Florida, and as far south as 1 Licenciatura en Biología, Universidad Distrital Francisco José de southern Chile (Lücking et al. 2014, 2017b). Caldas, Cra. 4 No. 26D-54, Torre de Laboratorios, Herbario, Bogotá D.C., Colombia; Research Associate, Science & Education, The Field The occurrences of Cora in Mexico and Florida Museum, 1400 South Lake Shore, Chicago, IL 60605, USA mark the northernmost distribution of the genus, which 2 Escuela de Biología, Benemérita Universidad Autónoma de Puebla, is remarkable, given that the latitudinal distribution range Puebla, México of Cora reaches only 28° north in Mexico and Florida 3 Botanischer Garten und Botanisches Museum, Freie Universität Berlin, Königin-Luise-Straße 6–8, 14195 Berlin, Germany but 40° south in southern Chile, a difference of more than * Corresponding author e-mail: [email protected] 1300 km. The reasons for the absence of Cora in western

This work is licensed under the Creative Commons BY-NC-ND 4.0 License 394 Plant and Fungal Systematics 64(2): 393–411, 2019

North America north of Mexico are unknown. For Mex- To place the sequenced collections from Mexico into ico, until recently only a single species had been reported, a broader context allowing assessment of the sampling under the names C. glabrata, C. pavonia, Dictyonema effort, we assembled a database of known occurrences glabratum and D. pavonium (Imshaug 1956; Ryan et al. of Cora from Mexico from CNALH (https://lichenportal. 1996; García et al. 2000; Guzmán 2008; Lücking et al. org/cnalh) and GBIF (https://www.gbif.org), searching 2009). The first sequence data for a Mexican specimen under the names C. glabrata, C. pavonia, D. glabratum were published by Lücking et al. (2009, 2014) as D. gla- and D. pavonium (Table S2). All collections were then bratum and Dictyonema sp., respectively. After the formal mapped using Map Maker (https://www.darrinward.com/ recognition of Cora as a genus separate from Dictyonema lat-long). s.str. (Dal Forno et al. 2013; Lücking et al. 2013), a much broader sampling of Cora revealed the presence of at Results and discussion least nine lineages in Mexico, none of them conspecific with C. glabrata s.str. or C. pavonia s.str., two species Phylogeny. The best-scoring maximum likelihood tree likely endemic to the Caribbean (Lücking et al. 2013, matches the topology obtained by Lücking et al. (2017b), 2015). Thus far, three new species from Mexico have dividing Cora into two large, well-supported clades, clade been formally described: C. casasolana, C. comaltepeca I and clade II (96% and 100% bootstrap support, respec- and C. dulcis. A further lineage was found nested within tively; Fig. 1, Fig. S1). The Mexican specimens form 12 C. dewisanti, a species known from the northern Andes in lineages, all except two unrelated to each other and more Colombia, Venezuela and Ecuador (Lücking et al. 2017b). or less evenly distributed over the tree, with three lineages For the present study we included additional material in clade I and nine in the larger clade II (Fig. 1, Fig. S1). from Mexico to reassess the diversity and evolutionary Eight represent new species and one a new subspecies, origins of this genus at its northernmost western limit in all formally described below. the Americas. We present an updated phylogeny based Five lineages are represented by singletons (C. casaso- on the fungal ITS barcoding marker, revealing 12 distinct lana, C. comaltepeca, C. dulcis, C. lawreyana, C. totonaco- lineages, eight of which are formally introduced as new rum), whereas seven lineages (C. benitoana, C. buapana, species and one as new subspecies in this paper. C. dewisanti subsp. mexicana, C. guzmaniana, C. ixtlanensis, C. marusae, C. zapotecorum) contain between Material and methods two and 16 specimens each (Table 1, Fig. S1). Of the non-singleton lineages, seven are reciprocally mono- Fresh material of Cora was collected in Mexico by the phyletic, whereas one is nested within a known spe- first two authors between 2015 and 2017 in the states cies, C. dewisanti (Fig. S1). The two Mexican lineages of Oaxaca and Veracruz. Specimens were studied at the C. cacasolana and C. totonacorum (both singletons) form Botanical Garden and Botanical Museum (Berlin, Ger- a sister clade (Fig. S1), thus resulting in 11 independent many), using standard techniques of light microscopy lineages present in Mexico (Fig. 1). and thin-layer chromatography (Orange et al. 2010). We The closest relatives of these 11 Mexican lineages employed a standardized protocol for morphological, are in four cases species from Costa Rica, in six cases anatomical and chemical characters to characterize the species from Colombia and adjacent areas in the northern material (Lücking et al. 2013, 2017b; Vargas et al. 2014). Andes (Venezuela, Ecuador), and in one case unresolved Voucher specimens were deposited at EB-BUAP, with (Fig. S1). The Mexican species with Costa Rican rela- duplicates at B. tives are C. marusae (with C. minor and C. paraminor), Part of the material had already been sequenced to C. benitoana (with C. barbulata), C. casasolana and provide the framework for an earlier, large-scale taxo- C. totonacorum (with C. haledana), and C. ixtlanensis nomic paper in which three species from Mexico were (with C. terrestris; Fig. S1). also formally described (Lücking et al. 2017b). In addi- tion to the 36 already published ITS sequences in that Phenotype characters. Including the aforementioned study, here we generated a further 18 new ITS sequences, species, eight of the 12 Mexican taxa can be compared following previously published protocols (Dal Forno in their phenotype characters (see descriptions and key et al. 2013; Lücking et al. 2017c). The newly gener- below) directly to closely related and phenotypically char- ated sequences were added to the global dataset of 653 acterized species (Fig. S1; see also Ariyawansa et al. 2015; sequences (including two outgroup taxa) assembled in Lücking et al. 2017b). Thus, C. marusae (Mexico) agrees the previous study (Lücking et al. 2017b), for a total of with C. minor and C. paraminor (both Costa Rica) in the 671 sequences (Table 1, Table S1), and were manually epiphytic growth, small size, and formation of soredia, but aligned (File S1) after checking their position using the differs in the presence of papilliform hyphal appendages local blast function in BioEdit 7 (Hall 1999). The data- on the lower medullary hyphae and in the forest-green set was subjected to a maximum likelihood search using rather than olive-green fresh color. Cora benitoana (Mex- RAxML 8.2.8 on the CIPRES Science Gateway server ico) and C. barbulata (Costa Rica) share the epiphytic (Miller et al. 2010; Stamatakis 2015), with non-parametric growth habit and the cyphelloid hymenophore, as well bootstrapping using 402 replicates (automatically deter- as the absence of papilliform hyphal appendages on the mined based on a saturation criterion) under the universal lower medullary hyphae, but differ in habitat ecology GTR-GAMMA model. (forest vs. paramo), size (medium-sized vs. large), surface B. Moncada et al.: The lichenized genus Cora in Mexico 395

Cora_guzmaniana_MON6660_Mexico-Perez-1335 Cora_guzmaniana_MON6666_Mexico-Perez-1298 Cora_guzmaniana_MON6658_Mexico-Perez-1333

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Cora_spec-93_KX772707_Colombia

Cora_spec-95_KJ780629_Ecuador

Cora_spec-94_KJ780557_Brazil

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Cora_yukiboa_KX772744_Puerto-Rico

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Cora_spec-84_KJ780636_Ecuador Cora_pseudocorani_KJ802417_BoliviaCora_setosa_KJ780535_Colombia

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Cora_paraciferrii_KJ780428_Colombia Cora_davibogotana_KX772429_Colombia Cora_paraciferrii_KX772649_Colombia Cora_davibogotana_KJ780505_Colombia Cora_paraciferrii_KJ780493_Colombia Cora_putumayensis_KX772608_Colombia Cora_paraciferrii_KX772517_Colombia Cora_paraciferrii_KJ780458_Colombia Cora_putumayensis_KX772611_Colombia Cora_paraciferrii_KJ780459_Colombia Cora_paraciferrii_KJ780431_Colombia Cora_dalehana_KJ780434_Colombia Cora_paraciferrii_KJ780423_Colombia Cora_spec-60_KX772697_Colombia Cora_paraciferrii_KX772458_Colombia Cora_spec-60_KX772662_Colombia Cora_paraciferrii_KX772612_Colombia Cora_spec-60_KX772679_Colombia Cora_paraciferrii_KX772598_Colombia Cora_paraciferrii_KX772501_Colombia Cora_spec-60_KX772678_Colombia Cora_paraciferrii_KX772647_Colombia Cora_spec-60_KX772718_Colombia Cora_paraciferrii_KX772518_Colombia Cora_spec-58_KJ780637_Ecuador Cora_ciferrii_KJ780387_Ecuador Cora_spec-59_KJ780638_Ecuador Cora_ciferrii_KX772448_Ecuador Cora_spec-57_KX772431_Colombia Cora_ciferrii_KX772451_Ecuador Cora_byssoidea_KF443234_Colombia Cora_ciferrii_KX772449_Ecuador Cora_ciferrii_KX772450_Ecuador Cora_byssoidea_KJ802425_Colombia Cora_ciferrii_KX772659_Colombia Cora_viliewoa_EU825956_Costa-Rica Cora_ciferrii_KJ780386_Ecuador Cora_viliewoa_KJ780634_Ecuador Cora_ciferrii_KJ780393_Ecuador Cora_viliewoa_KX772504_Colombia Cora_ciferrii_KJ780383_Ecuador Cora_viliewoa_KX772503_Colombia Cora_ciferrii_KX772729_Colombia Cora_ciferrii_KX772688_Colombia Cora_spec-52_KX772529_Colombia Cora_ciferrii_KX772703_Colombia Cora_spec-52_KX772685_Colombia Cora_ciferrii_KX772669_Colombia Cora_verjonensis_KX772457_Colombia Cora_ciferrii_KX772702_Colombia Cora_verjonensis_KJ780467_Colombia Cora_ciferrii_KJ780389_Ecuador Cora_palustris_KJ780597_Costa-Rica Cora_ciferrii_KJ780390_Ecuador Cora_ciferrii_KJ780380_Ecuador Cora_palustris_KJ780613_Costa-Rica Cora_ciferrii_KJ780382_Ecuador Cora_palustris_DQ917656_Costa-Rica Cora_ciferrii_KJ780620_Ecuador Cora_spec-56_KX772438_Colombia Cora_ciferrii_KJ780554_Ecuador Cora_udebeceana_KJ780520_Colombia Cora_ciferrii_KJ780384_Ecuador Cora_pikynasa_KX772511_Colombia Cora_ciferrii_KF443238_Ecuador Cora_spec-54_KX772521_Colombia Cora_ciferrii_KJ780385_Ecuador Corella_tomentosa_KJ780617_Costa-Rica Cora_spec-53_KX772520_Colombia Corella_zahlbruckneri_KJ780592_Brazil Cora_spec-53_KX772488_Colombia Cora_spec-1_KJ780426_Colombia Cora_comaltepeca_KX772556_Mexico Cora_spec-1_KJ780539_Colombia Cora_spec-55_KJ780668_Colombia Cora_spec-1_KJ780417_Colombia Cora_spec-55_KJ780672_Colombia Cora_spec-1_KJ780540_Venezuela Cora_spec-1_KJ780470_Colombia Cora_spec-55_KJ780675_Colombia Cora_pseudobovei_KJ780372_Bolivia Cora_spec-48_KJ780674_Colombia Cora_parabovei_KJ780571_Bolivia Cora_spec-48_KJ780669_Colombia Cora_bovei_KJ780579_Chile Cora_spec-47_KJ780581_Ecuador Cora_bovei_KJ780580_Chile Cora_spec-50_KX772466_Brazil Cora_palaeotropica_KX772743_Sri-Lanka Cora_spec-51_KJ780576_Brazil Cora_palaeotropica_KX772742_Sri-Lanka Cora_caraana_KX772512_Brazil Cora_celestinoa_KX772595_Colombia Cora_celestinoa_KX772596_Colombia Cora_spec-49_KJ780652_Brazil Cora_spec-2_KX772666_Colombia Cora_inversa_KJ780374_Colombia Cora_spec-2_KX772713_Colombia Cora_inversa_KF443237_Colombia Cora_spec-2_KX772730_Colombia Cora_inversa_KF443236_Colombia Cora_quillacinga_KX772701_Colombia Cora_fuscodavidiana_KJ780411_Colombia Cora_inversa_KJ780476_Colombia Cora_urceolata_KJ780452_Colombia Cora_inversa_KJ780446_Colombia Cora_urceolata_KJ780677_Colombia Cora_inversa_KJ780471_Colombia Cora_corelleslia_KX772433_Colombia Cora_inversa_KF922372_Colombia Cora_corelleslia_KJ780498_Colombia Cora_inversa_KF922371_Colombia Cora_spec-3_KX772580_Colombia Cora_spec-4_KJ780588_Costa-Rica Cora_inversa_KF922370_Colombia Cora_spec-5_KX772686_Colombia Cora_inversa_KJ780475_Colombia Cora_spec-5_KX772684_Colombia Cora_inversa_KX772502_Colombia Cora_spec-5_KX772690_Colombia Cora_inversa_KX772496_Colombia Cora_barbifera_KX772682_Colombia Cora_inversa_KX772525_Colombia Cora_schizophylloides_KX772719_Colombia Cora_hirsuta_KJ802426_Colombia Cora_inversa_KX772471_Colombia Cora_hirsuta_KJ780447_Colombia Cora_spec-46_KX772628_Ecuador Cora_hirsuta_KF443235_Colombia Cora_spec-6_KX772478_Colombia Cora_arborescens_KJ780599_Costa-Rica Cora_spec-7_KJ780404_Venezuela Cora_casanarensis_KJ780530_Colombia Cora_spec-8_KX772663_Colombia Cora_casanarensis_KJ780532_Colombia Cora_spec-9_KX772712_Colombia Cora_spec-10_KX772476_Colombia Cora_casanarensis_KX772705_Colombia Cora_spec-11_KX772657_Colombia Cora_casanarensis_KX772740_Colombia Cora_spec-12_KX772484_Colombia Cora_casanarensis_KX772579_Colombia Cora_rothesiorum_KX772526_Colombia Cora_casanarensis_KX772739_Colombia Cora_rothesiorum_KJ780490_Colombia Cora_casanarensis_KX772738_ColombiaCora_itabaiana_KX772630_Brazil Cora_rothesiorum_KJ780445_Colombia Cora_itabaiana_KX772629_Brazil Cora_rothesiorum_KJ780444_Colombia Cora_spec-45_KJ780647_Brazil Cora_marusae_KX772553_Mexico Cora_spec-45_KJ780641_Brazil Cora_marusae_KX772570_Mexico Cora_marusae_KX772542_Mexico Cora_spec-44_KJ780642_Brazil Cora_marusae_KX772541_Mexico Cora_spec-43_KJ780558_Brazil Cora_marusae_KX772564_Mexico Cora_strigosa_KF443241_Peru Cora_minor_KJ780363_Costa-Rica Cora_strigosa_KJ802413_Peru Cora_minor_EU825968_Costa-Rica Cora_strigosa_KJ802410_Peru Cora_paraminor_KJ780610_Costa-Rica Cora_paraminor_KJ780611_Costa-Rica Cora_spec-42_KX772635_Brazil Cora_paraminor_KJ780608_Costa-Rica Cora_paraminor_KJ780609_Costa-Rica Cora_spec-41_KJ780556_Ecuador Cora_spec-14_KX772725_Colombia Cora_spec-17_KX772715_Colombia Cora_zapotecorum_KX772546_Mexico Cora_spec-15_KX772522_Colombia Cora_zapotecorum_KX772568_Mexico Cora_spec-15_KX772527_Colombia Cora_spec-39_KJ780577_Brazil Cora_spec-16_KJ802427_Colombia Cora_zapotecorum_KX772566_MexicoCora_spec-39_KX772465_Brazil Cora_spec-16_KJ780375_Colombia Cora_spec-39_KJ780665_Brazil Cora_pastorum_KX772689_Colombia Cora_spec-39_KJ780662_Brazil Cora_pastorum_KX772660_Colombia Cora_davicrinita_KX772599_Colombia Cora_spec-39_KX772622_Brazil Cora_davicrinita_KX772510_Colombia Cora_davicrinita_KX772653_Colombia Cora_spec-38_KX772528_Colombia Cora_davicrinita_KX772652_Colombia Cora_davicrinita_KX772661_Colombia Cora_spec-38_KX772505_Colombia Cora_davicrinita_KJ780474_Colombia Cora_spec-38_KX772459_Colombia Cora_davicrinita_KX772733_Colombia Cora_spec-38_KX772575_Colombia Cora_davicrinita_KJ780555_Ecuador Cora_spec-38_KX772576_Colombia Cora_davicrinita_KJ780481_Colombia Cora_davicrinita_KJ780416_Colombia Cora_soredavidia_KJ780649_Brazil Cora_davicrinita_KJ780440_Colombia Cora_soredavidia_KJ780650_Brazil Cora_davicrinita_KJ780442_Colombia Cora_maxima_KJ802420_Bolivia Cora_campestris_KX772463_Brazil Cora_soredavidia_KJ780601_Costa-RicaCora_dewisanti_KX772591_Colombia Cora_campestris_KJ780578_Brazil Cora_gigantea_KX772439_Colombia Cora_dewisanti_KX772474_Colombia Cora_gigantea_KJ780515_Colombia Cora_dewisanti_KX772446_Ecuador Cora_aff-accipiter_KF443232_Venezuela Cora_dewisanti_KX772523_Colombia Cora_accipiter_KJ780478_Colombia Cora_accipiter_KX772499_Colombia Cora_dewisanti_KX772519_Colombia Cora_accipiter_KX772603_Colombia Cora_dewisanti_KJ780469_Colombia Cora_accipiter_KX772602_Colombia Cora_dewisanti_KJ780412_Colombia Cora_accipiter_KJ780457_Colombia Cora_accipiter_KJ780464_Colombia Cora_dewisanti_KJ780676_Colombia Cora_accipiter_KJ780430_Colombia Cora_dewisanti_KX772687_Colombia Cora_accipiter_KX772594_Colombia Cora_accipiter_KJ780453_Colombia Cora_dewisanti_KJ780403_Venezuela Cora_accipiter_KJ780494_Colombia Cora_dewisanti_KX772692_Colombia Cora_accipiter_KJ780501_Colombia Cora_dewisanti_KJ780407_Colombia Cora_accipiter_KJ780433_Colombia Cora_accipiter_KX772498_Colombia Cora_dewisanti_KJ780449_Colombia Cora_accipiter_KX772593_Colombia Cora_dewisanti_KX772447_Ecuador Cora_accipiter_KX772601_Colombia Cora_accipiter_KX772592_Colombia Cora_dewisanti_KJ780414_Colombia Cora_accipiter_KX772597_Colombia Cora_dewisanti_KJ780482_Colombia Cora_accipiter_KX772681_Colombia Cora_accipiter_KJ780495_Colombia Cora_dewisanti_KJ780409_Colombia Cora_accipiter_KX772691_Colombia Cora_dewisanti_KJ780484_Colombia Cora_arachnodavidea_KJ780492_Colombia Cora_arachnodavidea_KJ780477_Colombia Cora_dewisanti_KJ780486_Colombia Cora_spec-36_KJ780670_Colombia Cora_dewisanti_KJ780405_Colombia Cora_cyphellifera_KF443242_Ecuador Cora_cyphellifera_KJ780631_Ecuador Cora_cyphellifera_KX772677_Colombia Cora_cyphellifera_KX772675_Colombia Cora_barbulata_KJ780602_Costa-Rica Cora_barbulata_EU825961_Costa-Rica Cora_barbulata_KJ780605_Costa-Rica Cora_barbulata_KJ780604_Costa-Rica Cora_dewisanti-mexicana_KX772559_Mexico Cora_barbulata_KJ780606_Costa-Rica Cora_benitoana_KX772555_Mexico Cora_dewisanti-mexicana_KX772544_Mexico Cora_benitoana_KX772557_Mexico Cora_benitoana_KX772567_Mexico Cora_dewisanti-mexicana_KX772552_Mexico Cora_benitoana_KX772606_Mexico Cora_benitoana_KX772550_Mexico Cora_dewisanti-mexicana_KX772549_Mexico Cora_benitoana_KX772561_Mexico Cora_arachnoidea_KF443233_Venezuela Cora_dewisanti-mexicana_KX772543_Mexico Cora_arachnoidea_KX772711_Colombia Cora_arachnoidea_KX772485_Colombia Cora_dewisanti-mexicana_KX772548_Mexico Cora_arachnoidea_KX772533_Colombia Cora_arachnoidea_KX772664_Colombia Cora_dewisanti-mexicana_KX772560_Mexico Cora_arachnoidea_KX772656_Colombia Cora_arachnoidea_KX772670_Colombia Cora_dewisanti-mexicana_KX772558_Mexico Cora_arachnoidea_KX772671_Colombia Cora_arachnoidea_KX772655_Colombia Cora_dewisanti-mexicana_KX772538_Mexico Cora_arachnoidea_KX772735_Colombia Cora_aff-arachnoidea_KX772680_Colombia Cora_aff-arachnoidea_KX772651_Colombia Cora_dewisanti-mexicana_EU825957_Mexico Cora_arachnoidea_KX772708_Colombia Cora_dewisanti_KJ780667_Colombia Cora_arachnoidea_KJ780489_Colombia Cora_dewisanti-mexicana_KX772562_Mexico Cora_arachnoidea_KJ780438_Colombia Cora_dewisanti_KX772732_ColombiaCora_spec-35_KX772717_Colombia Cora_arachnoidea_KJ780465_Colombia Cora_arachnoidea_KJ780436_Colombia Cora_dewisanti-mexicana_KX772563_Mexico Cora_arachnoidea_KJ780439_Colombia Cora_spec-35_KX772731_Colombia Cora_arachnoidea_KX772614_Colombia Cora_arachnoidea_KJ780496_Colombia Cora_dewisanti-mexicana_KX772545_MexicoCora_spec-35_KX772721_Colombia Cora_arachnoidea_KJ780487_Colombia Cora_arachnoidea_KX772524_Colombia Cora_arachnoidea_KX772530_Colombia Cora_dewisanti-mexicana_KX772569_MexicoCora_spec-34_KX772736_Colombia Cora_arachnoidea_KJ780432_Colombia Cora_arachnoidea_KJ780437_Colombia Cora_arachnoidea_KJ780485_Colombia Cora_spec-33_KX772709_Colombia Cora_arachnoidea_KJ780408_Colombia Cora_arachnoidea_KJ780420_Colombia Cora_arachnoidea_KJ780462_Colombia Cora_arachnoidea_KJ780463_Colombia Cora_spec-33_KX772734_Colombia Cora_arachnoidea_KX772668_Colombia Cora_arachnoidea_KX772683_Colombia Cora_arachnoidea_KX772445_Ecuador Cora_arachnoidea_KJ780468_Colombia Cora_spec-31_KX772483_Colombia Cora_arachnoidea_KJ780480_Colombia Cora_arachnoidea_KX772590_Colombia Cora_arachnoidea_KX772722_Colombia Cora_arachnoidea_KX772456_Colombia Cora_arachnoidea_KX772650_Colombia Cora_arachnoidea_KX772710_Colombia Cora_arachnoidea_KX772714_Colombia Cora_arachnoidea_KX772672_Colombia Cora_arachnoidea_KX772479_Colombia Cora_spec-31_KX772481_Colombia Cora_arachnoidea_KX772486_Colombia Cora_arachnoidea_KX772489_Colombia Cora_arachnoidea_KX772728_Colombia Cora_arachnoidea_KX772477_Colombia Cora_spec-32_KX772532_Colombia Cora_arachnoidea_KX772726_Colombia Cora_spec-32_KX772470_Colombia Cora_spec-30_KX772723_Colombia Cora_spec-29_KJ780502_ColombiaCora_spec-28_KX772573_Brazil Cora_spec-29_KJ780429_ColombiaCora_spec-28_KJ780661_Brazil Cora_spec-27_KJ780654_Brazil Cora_spec-29_KX772646_Colombia Cora_spec-27_KX772637_Brazil

Cora_spec-26_KJ780518_Colombia Cora_spec-25_KX772706_Colombia

Cora_leslactuca_KJ780524_Colombia Cora_leslactuca_KJ780507_Colombia Cora_spec-23_KJ780527_Colombia Cora_leslactuca_KX772497_ColombiaCora_spec-23_KJ780516_Colombia Cora_leslactuca_KX772494_ColombiaCora_spec-24_KX772493_Colombia Cora_spec-24_KJ780435_Colombia Cora_spec-24_KX772720_Colombia Cora_elephas_KJ780406_Colombia Cora_elephas_KJ780422_Colombia Cora_elephas_KJ780418_Colombia Cora_elephas_KJ780424_Colombia Cora_elephas_KJ780448_Colombia Cora_elephas_KJ780454_Colombia Cora_elephas_KJ780413_Colombia Cora_elephas_KJ780450_Colombia Cora_elephas_KJ780456_Colombia Cora_elephas_KJ780410_Colombia Cora_elephas_KJ780455_Colombia Cora_elephas_KX772600_Colombia Cora_elephas_KX772507_Colombia Cora_spec-20_KX772464_Brazil Cora_elephas_KX772613_Colombia Cora_spec-22_KJ780523_Colombia Cora_spec-22_KJ780521_Colombia

Cora_spec-21_KX772605_Colombia Cora_spec-21_KX772604_Colombia Cora_elephas_KJ780396_Ecuador Cora_spec-19_KX772694_Colombia Cora_spec-19_KX772695_Colombia

Cora_elephas_KJ780499_Colombia Cora_elephas_KJ780497_Colombia

Cora_elephas_KX772452_Ecuador Cora_elephas_KJ780451_Colombia Cora_elephas_KJ780425_Colombia Cora_elephas_KJ780419_Colombia Cora_elephas_KJ780415_Colombia

Cora_aturucoa_KX772607_Colombia Cora_spec-18_KJ780479_Colombia Cora_spec-18_KJ780500_Colombia Cora_aturucoa_KX772615_Colombia

Cora_rubrosanguinea_KJ780400_Ecuador Cora_rubrosanguinea_KJ780395_Ecuador Cora_gomeziana_KJ780598_Costa-Rica

Cora_caucensis_KX772716_Colombia

Cora_arachnoidea_KX772673_Colombia

Cora_hochesuordensis_KJ802415_Bolivia Cora_hochesuordensis_KJ780367_Bolivia

Figure 1. Circular cladogram of the genus Cora (outgroup Corella) based on 669 ITS ingroup OTUs, showing the phylogenetic position of specimens from Mexico (orange). For the full tree with terminal labels and branch support, see Fig. S1.

(glabrous vs. setose) and the formation of projecting, but they are clearly distinct phylogenetically, differing shortly stipitate basidiocarps in C. benitoana. The three consistently in five positions in the ITS (File S1). In the species C. casasolana, C. totonacorum (both Mexico) and global ITS phylogeny (Fig. S1) they appear rather close, C. haledana (Costa Rica) agree in having medium-sized with C. terrestris receiving low support (62%) on a short thalli and lobes with more or less greenish fresh color branch, but the resolution and support within this terminal (olive-green to dark green) and the papilliform hyphal clade increase when analyzed separately (not shown). All appendages on the lower medullary hyphae. However, species in this terminal clade, also including C. caliginosa, they all differ in their substrate ecology, C. casasolana C. auriculeslia, C. ciferrii, C. paraciferrii and C. squami- being terrestrial and the other two species epiphytic; Cora formis, are terrestrial and very closely related, but differ totonacorum can further be distinguished in the formation partly in phenotype and substrate ecology, some growing of soredia. directly on soil and others between grasses or bryophytes, Cora ixtlanensis (Mexico) and C. terrestris (Costa and the species are largely allopatric, so this entire clade Rica) both grow terrestrially, have medium-sized thalli appears to represent a very recent radiation. and lobes with olive-grey color when fresh, share the corti- The sister-group relationship between C. lawreyana cioid hymenophore with concentric arrangement, and lack (Mexico) and C. minutula (Ecuador) is not supported papilliform appendages on the lower medullary hyphae; and both form long branches, which may explain that both can be considered phenotypically cryptic species the two are quite distinct phenotypically: medium-sized 396 Plant and Fungal Systematics 64(2): 393–411, 2019

Table 1. Voucher information and GenBank accession numbers for the Mexican specimens used in this study. Newly generated accessions are bolded. For GenBank accession numbers of the remaining, non-Mexican specimens used, see Table S1.

Genus Species ITS Accession Country Collectors Number Cora benitoana KX772550 Mexico B. Moncada & R.-E. Pérez-Pérez 10193 Cora benitoana KX772555 Mexico B. Moncada & R.-E. Pérez-Pérez 10180 Cora benitoana KX772557 Mexico B. Moncada & R.-E. Pérez-Pérez 10306 Cora benitoana KX772561 Mexico B. Moncada & R.-E. Pérez-Pérez 10199 Cora benitoana KX772567 Mexico B. Moncada & R.-E. Pérez-Pérez 10207 Cora benitoana KX772606 Mexico B. Moncada & R.-E. Pérez-Pérez 10183 Cora buapana KX772554 Mexico B. Moncada & R.-E. Pérez-Pérez 10305 Cora buapana KX772571 Mexico B. Moncada & R.-E. Pérez-Pérez 10197 Cora casasolana KX772540 Mexico B. Moncada & R.-E. Pérez-Pérez 10257 Cora comaltepeca KX772556 Mexico B. Moncada & R.-E. Pérez-Pérez 10233 Cora dewisanti subsp. mexicana EU825957 Mexico R. S. Egan 17538 Cora dewisanti subsp. mexicana KX772538 Mexico B. Moncada & R.-E. Pérez-Pérez 10300 Cora dewisanti subsp. mexicana KX772543 Mexico B. Moncada & R.-E. Pérez-Pérez 10297 Cora dewisanti subsp. mexicana KX772544 Mexico B. Moncada & R.-E. Pérez-Pérez 10228 Cora dewisanti subsp. mexicana KX772545 Mexico B. Moncada & R.-E. Pérez-Pérez 10278 Cora dewisanti subsp. mexicana KX772548 Mexico B. Moncada & R.-E. Pérez-Pérez 10195 Cora dewisanti subsp. mexicana KX772549 Mexico B. Moncada & R.-E. Pérez-Pérez 10196 Cora dewisanti subsp. mexicana KX772552 Mexico B. Moncada & R.-E. Pérez-Pérez 10301 Cora dewisanti subsp. mexicana KX772558 Mexico B. Moncada & R.-E. Pérez-Pérez 10185 Cora dewisanti subsp. mexicana KX772559 Mexico B. Moncada & R.-E. Pérez-Pérez 10302 Cora dewisanti subsp. mexicana KX772560 Mexico B. Moncada & R.-E. Pérez-Pérez 10176 Cora dewisanti subsp. mexicana KX772562 Mexico B. Moncada & R.-E. Pérez-Pérez 10154 Cora dewisanti subsp. mexicana KX772563 Mexico B. Moncada & R.-E. Pérez-Pérez 10322 Cora dewisanti subsp. mexicana KX772569 Mexico B. Moncada & R.-E. Pérez-Pérez 10198 Cora dulcis KX772551 Mexico B. Moncada & R.-E. Pérez-Pérez 10288 Cora guzmaniana MN475936 Mexico R.-E. Pérez-Pérez & D. León-González 1325 Cora guzmaniana MN475948 Mexico R.-E. Pérez-Pérez & D. León-González 1323 Cora guzmaniana MN475950 Mexico R.-E. Pérez-Pérez & D. León-González 1328 Cora guzmaniana MN475940 Mexico R.-E. Pérez-Pérez & D. León-González 1329 Cora guzmaniana MN475941 Mexico R.-E. Pérez-Pérez & D. León-González 1330 Cora guzmaniana MN475944 Mexico R.-E. Pérez-Pérez & D. León-González 1331 Cora guzmaniana MN475938 Mexico R.-E. Pérez-Pérez & D. León-González 1332 Cora guzmaniana MN475949 Mexico R.-E. Pérez-Pérez & D. León-González 1333 Cora guzmaniana MN475945 Mexico R.-E. Pérez-Pérez & D. León-González 1334 Cora guzmaniana MN475947 Mexico R.-E. Pérez-Pérez & D. León-González 1335 Cora guzmaniana MN475942 Mexico R.-E. Pérez-Pérez & D. León-González 1336 Cora guzmaniana MN475951 Mexico R.-E. Pérez-Pérez & D. León-González 1337 Cora guzmaniana MN475943 Mexico R.-E. Pérez-Pérez & D. León-González 1338 Cora guzmaniana MN475939 Mexico R.-E. Pérez-Pérez & D. León-González 1327 Cora guzmaniana MN475937 Mexico R.-E. Pérez-Pérez & D. León-González 1326 Cora guzmaniana MN475946 Mexico R.-E. Pérez-Pérez & D. León-González 1298 Cora ixtlanensis KX772539 Mexico B. Moncada & R.-E. Pérez-Pérez 10264 Cora ixtlanensis KX772547 Mexico B. Moncada & R.-E. Pérez-Pérez 10205 Cora ixtlanensis KX772565 Mexico B. Moncada & R.-E. Pérez-Pérez 10293 Cora lawreyana MN475935 Mexico R.-E. Pérez-Pérez & G. Águila-Rodríguez 1168 Cora marusae KX772541 Mexico B. Moncada & R.-E. Pérez-Pérez 10212 Cora marusae KX772542 Mexico B. Moncada & R.-E. Pérez-Pérez 10319 Cora marusae KX772553 Mexico B. Moncada & R.-E. Pérez-Pérez 10178 Cora marusae KX772564 Mexico B. Moncada & R.-E. Pérez-Pérez 10316 Cora marusae KX772570 Mexico B. Moncada & R.-E. Pérez-Pérez 10222 Cora totonacorum MN475934 Mexico R.-E. Pérez-Pérez & G. Águila-Rodríguez 1189b Cora zapotecorum KX772546 Mexico B. Moncada & R.-E. Pérez-Pérez 10318 Cora zapotecorum KX772566 Mexico B. Moncada & R.-E. Pérez-Pérez 10307 Cora zapotecorum KX772568 Mexico B. Moncada & R.-E. Pérez-Pérez 10206 B. Moncada et al.: The lichenized genus Cora in Mexico 397 vs. small, with vs. without papilliform appendages on the lower medullary hyphae, and with a corticioid hymenophore vs. soredia. Cora guzmaniana (Mexico) and C. galapagoensis (Galapagos) agree in the epiphytic growth and medium-sized thallus and lobes, as well as in the concentrically arranged, corticioid hymenophore and the presence of papilliform appendages on the lower medullary hyphae, but the first species differs in the pigmented hyphal layers enclosing the photobiont layer and in the coralloid-spinulose appendages on the lower medullary hyphae. Finally, Cora dewisanti subsp. mexicana (Mexico) differs consistently from the nominal subspecies (known from Colombia, Venezuela and Ecuador) in the much larger thallus and lobes and in the often marginally protruding hymenophore. Thus, while the new species shared certain phenotyp- ical characters between closely related lineages, most of the novel taxa are diagnosable by at least one character, supporting their taxonomic recognition not only on molecular grounds. The only exceptions are the morphologically cryptic species C. ixtlanensis and C. terrestris.

Associations of lichen thalli with bryophytes. Nota- bly, of the 54 sequenced specimens studied here, 43 were associated with liverwort mats of the genus Frullania (Fig. 2), either exclusively (14) or with further bryophyte taxa in between, and then usually with the Frullania species being dominant (see also descriptions below). It was recently shown that Frullania may act as a reservoir for the cyanobacterial photobiont Rhizonema (Cornejo & Scheidegger 2016), which is the photobiont present in Cora lichens and their relatives, as well as in several cyanobacterial ascolichens (Lawrey et al. 2009; Cornejo & Scheidegger 2016). Since Cora disperses predomi- nantly by basidiospores which require de-novo symbi- osis to form a new thallus, the notable association of Cora thalli with Frullania mats may be explained by the presence of non-lichenized Rhizonema colonies in these liverworts. A similar association was found in the related Acantholichen galapagoensis, which predom- inantly grows on thick epiphytic Frullania mats (Dal Forno et al. 2017). An alternative explanation would be for these associations to be opportunistic, given the ubiq- uity of Frullania mats in the habitats where most Cora Figure 2. Examples of thalli of various Cora specimens associated with Frullania mats. A – C. lawreyana (holotype, field image); B – C. bua- species abound. This is contradicted by the observation pana (isotype, rewetted); C – C. dewisanti subsp. mexicana (Moncada that the frequently sorediate species, C. marusae, which & Pérez-Pérez 10300, rewetted). does not require a new photobiont to propagate, was in only one out of five cases associated withFrullania mats More sophisticated methods including next-gen sequenc- and in the other cases with other bryophytes. These other ing are required to test this hypothesis. bryophytes, usually mosses, do not develop water sacs or analogous structures with cyanobacteria. The difference Species richness and putative endemism. The 54 in the association with Frullania mats in non-sorediate sequenced specimens mostly originate from southeastern vs. sorediate taxa is statistically significant (Chi-Square Mexico, from the states of Veracruz and Oaxaca, with one test with Yates correction: χ = 16.6, p < 0.001). We there- specimen also from southwestern Mexico in the state of fore hypothesize that also in the case of Cora, Frullania Jalisco (Fig. 3). Overall, Cora has been recorded mostly mats may act as a reservoir for Rhizonema to form thalli from this area, but also from northwestern, western, cen- through relichenization. Examination of associated Frul- tral and southeasternmost Mexico, from the states of Chi- lania mats revealed the presence of a variety of cyanobac- huahua, Sinaloa, Durango, Jalisco, México D.F., Puebla, teria both on the stem and leaf surfaces and in the water Tlaxcala and Chiapas (Fig. 3). Thus, while the sequenced sacs, but none unequivocally identifiable as Rhizonema. material chiefly represents the eastern coniferous and 398 Plant and Fungal Systematics 64(2): 393–411, 2019

Figure 3. Map showing recorded distribution of Cora in Mexico. Red circles: occurrence records from CNALH and GBIF; blue circles: occurrence records of sequenced specimens included in this study. The grids indicate those used in the previous global richness prediction analysis (see Lücking et al. 2014).

cloud forests ranging from the Sierra Madre Oriental (2–)11–17 species per grid for the central volcanic belt through the central volcanic belt to the Sierra Madre and the Sierra Madre del Sur (grids 20–23, 30–31), and del Sur, Cora also extends far into the (north-)western 36 per grid for the Sierra Madre de Chiapas extending coniferous forests of the Sierra Madre Occidental. The far into Guatemala (grid 32). The 12 taxa based on the sequenced material from Veracruz and Oaxaca represents sequenced material correspond to grids 20, 22 and 23, for invariably endemic lineages that are here interpreted as which two, 17 and five species were predicted, respec- 11 species and one subspecies (see taxonomic treatment tively. Our findings documented one, two and ten taxa below). Given the pronounced depression east of the for the three grids, respectively, with one (C. dewisanti Sierra Madre del Sur along the Isthmus of Tehuantepec, subsp. mexicana) overlapping between grids 20 and 23. between Oaxaca and Chiapas, it is likely that these spe- Notably, there was no overlap between the two taxa found cies will remain Mexican endemics, even with further in grid 22 (Veracruz) and the ten taxa in grid 23 (Oaxaca). sampling eastwards into Chiapas and Guatemala. We The predicted grid total for the Mexican grids, excluding also expect that the material previously recorded from grid 36 with overlaps with Guatemala, was 80. Given the central and (north-)western Mexico represents additional, estimated mean grid predicted range of 2.66 for Cora unrecognized species, mostly different from those now species across the Americas (Lücking et al. 2014), this recognized for southeastern Mexico, given the underlying would result in 80/2.66 = 30 predicted species for Mex- topography, with separation between the Sierra Madre ico excluding Chiapas. This number appears reasonable, Occidental, Oriental, and del Sur. Unfortunately, sim- considering that the thus-far sequenced material chiefly ple revision of the previously reported collections is not covers southeastern Mexico west of Chiapas, whereas, sufficient to clarify this, as fresh material is required to with the exception of a single specimen from Jalisco, assess field characters and substrate ecology and to gen- the majority of the documented range for the genus in erate sequence data for accurate placement of these taxa. Mexico, specifically central Mexico and the entire Sierra In the study by Lücking et al. (2014), who predicted Madre Occidental, has not yet been sampled for molecular global species richness in the genus Cora using a quan- analysis. We hypothesize that additional species unveiled titative, linear extrapolation technique based on a 370 in this vast area will largely be Mexican endemics. In × 300–370 km2 grid map (Fig. 3), Mexico was repre- contrast, the entirely undocumented but high predicted sented by 22 grids, with a single sequenced specimen richness for Chiapas, likely adding another 30 species, from Jalisco (grid 20). Grid-based species richness was will at least partly be shared with Guatemala. All together, estimated at 2–3 species per grid in the Sierra Madre we can thus expect about 60 species of Cora for Mexico, Occidental (grids 02, 07, 08, 12), up to eight species per only 12 (20%) of which are now known. For compari- grid in the Sierra Madre Oriental (grids 04, 09, 10, 14), son, thus far 108 species are recognized for the territory B. Moncada et al.: The lichenized genus Cora in Mexico 399 of Colombia, which is little more than half the size of aeruginous-green; medulla 30–50 µm thick, hydrophobic; Mexico, 44 of which (40%) have been formally described clamp connections absent, papilliform hyphal appendages (Lücking et al. 2013, 2017b). absent. Hymenophore cyphelloid, developed partly as The 12 sequenced taxa known so far from Mexico irregular, resupinate patches mostly towards lobe margins, originated from 11 independent migration events, with partly as shortly stipitate, strongly protruding basidiocarps the closest relatives identified from either Costa Rica or beyond lobe margins, resupinate patches 10–30 mm long the northern Andes. This pattern is rather remarkable, and 5–10 mm broad, with white (fresh) to cream-colored given that most of these lineages were found growing in (in the herbarium), smooth surface and setose, involute close geographic proximity, and suggests that multiple margins, protruding basidiocarps distinctly cyphelloid, lineages of Cora migrated northwards from South Amer- up to 10 mm long and broad, with setose upper surface; ica more or less simultaneously, at least on geological hymenophore in section 100–150 µm thick, resting on timescales. This is supported by the terminal position of medullary layer or entirely protruding; hymenium com- all Mexican lineages and the observation that in a third posed of numerous palisade-like basidioles and basidia; of these lineages a Costa Rican species is the closest basidioles 20–30 × 5–6 µm; basidia 30–40 × 5–7 µm, relative (Fig. S1). 4-sterigmate; basidiospores not observed. Chemistry: No substances detected by TLC.

Taxonomy Etymology. This new species is dedicated to Benito Pablo Cora benitoana Moncada, R.-E. Pérez & Lücking, Juárez García, Mexican president from 1857 to 1872, of sp. nov. (Fig. 4A–D) indigenous origin (Zapotec) from the state of Oaxaca. A progressive reformer, Benito Juárez is considered the MycoBank MB 832769 ‘Mexican Abraham Lincoln’ and is the only individual Diagnosis: A (medium-sized to) large Cora growing epiphyt- whose birthday is a national holiday in Mexico. ically on tree trunks over bryophytes, fresh with dark grey- green, distinctly narrowly undulate lobes lacking color zonation, Distribution and ecology. Cora benitoana is so far with a glabrous surface and with distinct, whitish, glabrous to known from a small area in Oaxaca, growing epiphytically thinly pilose margins; hymenophore cyphelloid, in part forming on tree trunks in cloud forest (bosque mesófilo) between strongly protruding dorsiventral basidiocarps with setose upper 2000 and 3000 m. surface; differing from the closely relatedC. barbulata (known from Costa Rica) in the glabrous lobe surface, strongly projecting basidiocarps, and forest habitat. Notes. This new species is well characterized by its unusually large, cyphelloid hymenophore developed Type: Mexico: Oaxaca. Santiago Comaltepec, Cerro Pelón; mostly towards the lobe margins and partly as entirely 17°34′40.2″N; 96°30′21.6″W; 2990 m; cloud forest (bosque mesófilo), 17 May 2015, B. Moncada & R.-E. Pérez-Pérez 10193 protruding, shortly stipitate basidiocarps closely resem- (EB-BUAP – holotype!; B – isotype!). bling those of the related, appressed-filamentous genus Cyphellostereum (Dal Forno et al. 2013). There are several ITS barcoding sequence(s): KX772550 (isotype), other mid-sized to large epiphytic species with (dark) KX772555, KX772557, KX772561, KX772567, green color when fresh, but most feature a typically KX772606 (paratypes). adnate, concentric-reticulate hymenophore over all of Description. Thallus epiphytic on tree trunks, usually the lobe underside and most are smaller. Cora barbulata, over mosses and liverworts (often Frullania), foliose, described from Costa Rica (Ariyawansa et al. 2015), is up to 20 cm across, composed of 10–30 semicircular, most closely related and shares with C. benitoana the imbricate lobes; individual lobes 3–6 cm wide and 2–4 cm epiphytic growth habit and the cyphelloid hymenophore, long, moderately to frequently branched and with short, as well as the absence of papilliform hyphal appendages radially branching sutures, surface dark grey-green when on the lower medullary hyphae, but differs in the termi- fresh, without concentric color zonation, with distinct, nally more-branched lobes, the partially setose, more or involute, light greyish margins, light bluish grey when less even lobe surface, the paramo habitat, and the ubica- dry and becoming light grey in the herbarium, without tion of the hymenophore on the lobe underside, without distinct concentric color zonation. Upper surface distinctly forming protruding basidiocarps. A similar cyphelloid narrowly undulate and somewhat pitted when fresh, hymenophore which also agrees in size and epiphytic shallowly undulate to almost even when dry, glabrous; habit is otherwise known from C. cyphellifera, known involute margins glabrous to thinly pilose; lower sur- from Ecuador and Colombia (Lücking et al. 2013), but face ecorticate, felty-arachnoid (representing the exposed in that species the thallus is light aeruginous when fresh. medulla), whitish when fresh, becoming yellowish white The two species species are related but clearly set apart in the herbarium. Thallus in section 150–200 µm thick, phylogenetically (see Fig. S1). with upper cortex, photobiont layer and medulla; upper Additional specimens examined. MEXICO: Oaxaca. San- cortex diffusely viaduct-shaped, formed by a 20–30 µm tiago Comaltepec, Cerro Pelón; 17°34′40.2″N; 96°30′21.6″W; thick layer of loosely packed, irregularly arranged to peri- 2990 m; cloud forest (bosque mesófilo), 17 May 2015, B. Mon- clinal, 4–6 µm thick hyphae supported by a 30–50 µm cada & R.-E. Pérez-Pérez 10180, 10183, 10193, 10199, 10207 high ‘medullary’ layer of spaced groups of anticlinal, (EB-BUAP, B – paratypes!). Santiago Comaltepec, Cerro Las 4–6 µm thick hyphae; photobiont layer 50–70 µm thick, Antenas; 17°35′16.5″N; 96°27′2.7″W; 2190 m; cloud forest 400 Plant and Fungal Systematics 64(2): 393–411, 2019

Figure 4. Habit of new Cora taxa. A–D – C. benitoana (holotype and isotype), showing rewetted thallus with basidiocarps (A), close-up of basidiocarps on dry thallus in the field (B) and rewetted thallus (C), and lobe underside with marginal hymenophore (D). E, F – C. buapana (isotype), showing rewetted thallus (E), lobe underside with corticioid-concentrical hymenophore (F) and partly branched-coralloid papillae on lower medullary hyphae (insets).

(bosque mesófilo), 17 May 2015, B. Moncada & R.-E. Pérez- lower medullary hyphae, and from other similar-sized epiphytic Pérez 10306 (EB-BUAP, B – paratypes!). species with hyphal appendages in the grey rather than green fresh color. Cora buapana Moncada, R.-E. Pérez & Lücking, Type: Mexico: Oaxaca. Santiago Comaltepec, Cerro Las sp. nov. (Fig. 4E–F) Antenas; 17°35′16.5″N; 96°27′2.7″W; 2190 m; cloud forest (bosque mesófilo), 17 May 2015, B. Moncada & R.-E. Pérez- MycoBank MB 832770 Pérez 10305 (EB-BUAP – holotype!; B – isotype!). Diagnosis: A medium-sized Cora growing epiphytically on branches of trees or shrubs over and between bryophytes, ITS barcoding sequence(s): KX772554 (isotype), fresh with dark grey to olive-grey lobes lacking distinct color KX772571 (paratype). zonation, with a glabrous surface; hymenophore corticioid and distinctly concentrically arranged; differing from C. lawreyana Description. Thallus epiphytic on branches of trees and in the elongate, finger-like, partly branched appendages on the shrubs, usually over and between mosses and liverworts B. Moncada et al.: The lichenized genus Cora in Mexico 401

(often Frullania), foliose, up to 15 cm across, composed lobes. The sympatric C. benitoana (see above) can be of 5–25 semicircular, slightly imbricate lobes; individual readily distinguished by the dark grey-green, distinctly lobes 1.5–3 cm wide and 1–2(–3) cm long, rather fre- undulate lobes and the cyphelloid hymenophore forming quently branched at base, with thin radially branching strongly protruding basidiocarps, and anatomically by the sutures that often expose photobiont clusters to appear absence of papilliform appendages on the lower medul- granulose-sorediate, surface dark grey to olive-grey lary hyphae. The otherwise very similar but allopatric when fresh, without concentric color zonation, with dis- and only distantly related C. lawreyana differs chiefly tinct, involute, light greyish margins, white-grey when in the short, mostly globose hyphal appendages in the dry and in the herbarium, without distinct concentric lower medulla. color zonation. Upper surface uneven when fresh and dry, glabrous; involute margins glabrous; lower surface Additional specimens examined. MEXICO: Oaxaca. San- ′ ″ ′ ″ ecorticate, felty-arachnoid (representing the exposed tiago Comaltepec, Cerro Pelón; 17°34 40.2 N; 96°30 21.6 W; 2990 m; cloud forest (bosque mesófilo), 17 May 2015, medulla), whitish when fresh, becoming yellowish white B. Moncada & R.-E. Pérez-Pérez 10197 (EB-BUAP, B – para- in the herbarium. Thallus in section 250–300 µm thick, types!). with upper cortex, photobiont layer and medulla; upper cortex viaduct-shaped, formed by 20–30 µm thick layer Cora dewisanti subsp. mexicana Moncada, R.-E. Pérez of rather densely packed, ± periclinal, 4–6 µm thick & Lücking, subsp. nov. (Fig. 5A–C) hyphae supported by 50–80 µm high ‘medullary’ layer MycoBank MB 832771 of spaced groups of anticlinal, 4–6 µm thick hyphae; photobiont layer 70–120 µm thick, aeruginous-green; Diagnosis: A large to very large Cora growing terrestrially medulla 30–70 µm thick, hydrophobic; clamp connec- between bryophytes and other lichens, fresh with olive-grey lobes lacking distinct color zonation except when dry, with tions absent but lower medullary hyphae with scattered, a glabrous surface and with distinct, whitish, glabrous to thinly papilliform to elongate-finger-like or flask-shaped, partly pilose margins; hymenophore corticioid-cyphelloid, often pro- branched hyphal appendages 3–7 µm long and 2–3 µm truding beyond the lobe margins; differing from subsp.dewisanti thick. Hymenophore corticioid, forming resupinate, linear in the much larger thallus and lobes and in the hymenophore and partly branched and anastomosing patches across lobe often protruding beyond the lobe margins, as well as the forest underside in distinctly concentric arrangement, patches habitat ecology. 3–15 mm long and 0.5–1 mm broad, with white (fresh) to Type: Mexico: Oaxaca. Santiago Comaltepec, Cerro Pelón; cream- or flesh-colored (in the herbarium), smooth surface 17°34′40.2″N; 96°30′21.6″W; 2990 m; cloud forest (bosque and glabrous to very minutely pilose, slightly involute mesófilo), 17 May 2015, B. Moncada & R.-E. Pérez-Pérez 10196 margins; hymenophore in section 100–130 µm thick, rest- (EB-BUAP – holotype!; B – isotype!). ing on thick medullary layer; hymenium composed of ITS barcoding sequence(s): KX772549 (isotype), numerous palisade-like basidioles and scattered basidia; EU825957, KX772538, KX772543, KX772544, basidioles 20–30 × 5–6 µm; basidia 30–35 × 5–7 µm, KX772545, KX772548, KX772552, KX772558, 4-sterigmate; basidiospores not observed. Chemistry: No KX772559, KX772560, KX772562, KX772563, substances detected by TLC. KX772569 (paratypes). Etymology. The epithet honors the Benemérita Univer- Description. Thallus terrestrial between mosses and sidad Autónoma de Puebla (BUAP), the oldest and larg- liverworts (often Frullania) and other lichens (Hypo- est university in the state of Puebla, founded in 1578 as trachyna), foliose, up to 30 cm across, composed of a Jesuit college. 10–30 semicircular, imbricate lobes; individual lobes Distribution and ecology. Cora buapana has an ecol- 3–8 cm wide and 2–6 cm long, moderately to frequently ogy similar to that of C. benitoana, so far known from branched and with short, radially branching sutures, a small area in Oaxaca, growing epiphytically on tree surface olive-grey when fresh, without concentric color trunks in cloud forest (bosque mesófilo) between 2000 zonation, with distinct, involute, whitish to greyish mar- and 3000 m. It is apparently rarer than the preceding gins, light bluish grey when dry and becoming grey with species. yellowish margins in the herbarium, with distinct concentric color zonation of paler and darker areas. Upper Notes. Cora buapana forms part of a rather large surface even to shallowly undulate when fresh, undu- group of medium-sized, epiphytic, non-sorediate spe- late-rugose when dry, glabrous; involute margins gla- cies described from Central America (Costa Rica), the brous to shortly pilose, characteristically enrolled when northern and central Andes (Colombia, Ecuador, Bolivia) dry, forming triangular patterns across lobes; lower sur- and the Galapagos (Dal Forno et al. 2017; Lücking et al. face ecorticate, felty-arachnoid (representing the exposed 2017b). Of these, C. boleslia (Bolivia), C. haledana medulla), whitish when fresh, becoming yellowish white (Costa Rica), C. pastorum (Colombia), C. santacruzen- in the herbarium. Thallus in section 300–400 µm thick, sis (Galapagos) and C. viliewoa (Costa Rica) agree in with upper cortex, photobiont layer and medulla; upper the formation of papilliform appendages on the lower cortex diffusely viaduct-shaped, formed by 150–200 µm medullary hyphae and in the corticioid, concentrically thick layer of loosely and irregularly arranged, 4–6 µm arranged hymenophore (where known), but differ in the thick hyphae; photobiont layer 70–100 µm thick, orange- aeruginous to emerald- or olive-green fresh color of the brown above, olive-green below; medulla 30–70 µm 402 Plant and Fungal Systematics 64(2): 393–411, 2019

thick, hydrophobic; clamp connections absent, papil- Cora guzmaniana Moncada, R.-E. Pérez & Lücking, liform hyphal appendages absent. Hymenophore cor- sp. nov. (Fig. 5D–F) ticidoid-cyphelloid, developed as rounded to elongate, MycoBank MB 832772 resupinate patches, mostly towards lobe margins and Diagnosis: A medium-sized Cora growing epiphytically on tree often protruding beyond them, sometimes also extend- trunks over bryophytes, fresh with dark grey lobes lacking dis- ing over nearby bryophytes, patches 5–30 mm long and tinct color zonation, with a mostly glabrous surface but with 2–5 mm broad, often irregularly aggregate, with white indistinct concentric lines of scattered setae, especially towards to cream-colored, smooth surface and felty, involute the margin, and with distinct, whitish, glabrous to thinly pilose margins; hymenophore in section 100–150 µm thick, margins; lower medullary hyphae with coralloid, often spinulose resting on thick medullary layer; hymenium composed of hyphal appendages; hymenophore corticioid, forming elongate, numerous palisade-like basidioles and basidia; basidioles concentrically arranged patches; differing from C. buapana 20–35 × 5–7 µm; basidia 30–35 × 5–7 µm, 4-sterigmate; chiefly in the partly setose lobe surface and the larger, coral- basidiospores not observed. Chemistry: No substances loid and spinulose hyphal appendages on the lower medullary hyphae. detected by TLC. Type: Mexico: Oaxaca. San Vicente, Nuñú, Teposcolola; Etymology. The subspecific epithet refers to the distinc- 17°24′N; 97°26′W; 2616 m; conifer forest (bosque de Juni- tive distribution of this lineage as a putative Mexican perus), 14 November 2017, R.-E. Pérez-Pérez & D. León- endemic. González 1327 (EB-BUAP – holotype!; B – isotype!).

Distribution and ecology. This is so far the most ITS barcoding sequence(s): MN475939 (isotype), widely distributed taxon in Mexico, known from the MN475936, MN475937, MN475938, MN475940, states of Jalisco and Oaxaca, suggesting a distribution MN475941, MN475942, MN475943, MN475944, along the central volcanic belt. The nominal subspecies MN475945, MN475946, MN475947, MN475948, is also rather widely distributed in the northern Andes MN475949, MN475950, MN475951 (paratypes). (Colombia, Venezuela, Ecuador; Lücking et al. 2017b), indicating increased dispersal capacity in this lineage as Description. Thallus epiphytic, on Juniperus flaccida compared to most other species of Cora (Lücking et al. between mosses and other lichens (Flavoparmelia, Leu- 2014). Whereas subsp. dewisanti grows in paramos above codermia and Parmotrema), foliose, up to 10 cm across, 3000 m in the northern Andes, subsp. mexicana has so composed of 5–10 semicircular, partly imbricate lobes; far been found only in cloud forest (bosque mesófilo) individual lobes 1–3(–4) cm wide and 1–2 cm long, mod- between 2000 and 3000 m. erately to frequently branched and with short, radially branching sutures, surface dark grey when fresh, without Notes. Cora dewisanti subsp. mexicana is phyloge- concentric color zonation but more olive-grey towards netically nested within C. dewisanti, where it forms margin, with distinct, involute, olive-grey margins, a monophyletic clade differing from the nominal sub- becoming white to light grey in the herbarium, without species consistently in four insertions in the ITS. While distinct concentric color zonation. Upper surface (un-) the nominal subspecies appears to be widespread in even when fresh, shallowly undulate to almost even when Colombia, Venezuela and Ecuador, subsp. mexicana dry, mostly glabrous but with thin concentric lines of has so far been found only in Mexico. Together with short, erect, thin, white setae, especially towards margin; the nested position, this merits recognition at infra- involute margins glabrous to thinly pilose; lower sur- species level rather than as a distinct species. The tax- face ecorticate, felty-arachnoid (representing the exposed onomic distinction is supported by the morphometric medulla), whitish when fresh, becoming yellowish white and ecological differences, with subsp.mexicana devel- in the herbarium. Thallus in section 300–400 µm thick, oping much larger thalli and lobes and a more robust with upper cortex, photobiont layer, and medulla; upper hymenophore not protruding beyond the lobe margins, cortex diffusely viaduct-shaped, formed by 100–200 µm and growing in forest habitats, but otherwise agreeing thick layer of loosely packed, irregularly arranged, 4–6 µm in overall morphology. thick hyphae; photobiont layer 100–150 µm thick, olive- green above, aeruginous-green below, sometimes above Additional specimens examined. MEXICO: Jalisco. Parque Nacional Volcán Nevado de Colima, beyond entrance station in and below the photobiont layer with a 20–30 µm thick, La Joya area near the campground; 19°35′N, 103°36′W, 3415 m; orange-brown-pigmented layer of irregularly arranged 26 December 2006, R. Egan 17538 (OMA). Oaxaca. Ixtlán hyphae; medulla 30–50 µm thick, hydrophobic; clamp de Juárez, Desviación a San Pedro de Yaneri; 17°24′29.9″N; connections absent but lower medullary hyphae with 96°29′59″W; 2978 m; cloud forest (bosque mesófilo), 17 May numerous shortly coralloid hyphal appendages 5–15 µm 2015, B. Moncada & R.-E. Pérez-Pérez 10154, 10176, 10322 long and 5–10 µm thick, appearing somewhat spinulose (EB-BUAP, B – paratypes!). Santiago Comaltepec, Cerro Las due to short, thin hyphal outgrowths 2–3 µm long and ′ ″ ′ ″ Antenas; 17°35 16.5 N; 96°27 2.7 W; 2190 m; cloud forest 1–2 µm broad. Hymenophore corticioid, forming con- (bosque mesófilo), 17 May 2015, B. Moncada & R.-E. Pérez- centrically arranged, resupinate, rounded to elongate and Pérez 10278, 10297, 10300, 10301, 10302 (EB-BUAP, B – paratypes!). Santiago Comaltepec, Cerro Pelón; 17°34′40.2″N; sometimes branched patches across lower lobe surface, 96°30′21.6″W; 2990 m; cloud forest (bosque mesófilo), 17 May patches 1–5 mm long and 0.5–1 mm broad, with white 2015, B. Moncada & R.-E. Pérez-Pérez 10185, 10195, 10198, (fresh) to cream-colored (in the herbarium), smooth sur- 10228 (EB-BUAP, B – paratypes!). face and glabrous to minutely pilose, involute margins; B. Moncada et al.: The lichenized genus Cora in Mexico 403

Figure 5. Habit of new Cora taxa. A–C – C. dewisanti subsp. mexicana (Moncada & Pérez-Pérez 10300), showing dry thallus in the field (A) and rewetted lobes (B) with partially protruding hymenophore, and lobe underside with marginal hymenophore (C). D–F – C. guzmaniana (D Pérez-Pérez & León-González 1334, E–F isotype), showing dry thallus with concentric rows of setae towards the lobe margins (D), rewetted thallus (E) and lobe underside with corticioid-concentric hymenophore (F). hymenophore in section 70–100 µm thick, resting on (Guzmán-Dávalos 2016; Mata & Salmones 2016; Guz- a multilayered medullary layer including older hymenial mán-Dávalos & Cifuentes 2018). layers; hymenium composed of numerous palisade-like basidioles and basidia; basidioles 20–30 × 5–7 µm; Distribution and ecology. This new species is thus far basidia 30–35 × 5–7 µm, 4-sterigmate; basidiospores known from only a single locality in Oaxaca, growing not observed. Chemistry: No substances detected by TLC. rather abundantly in patches of Juniperus trees immersed in upper montane conifer forest. Etymology. The epithet of this new species honors Gastón Guzmán Huerta [1932–2016], the most prom- Notes. Cora guzmaniana is another medium-sized epi- inent Mexican mycologist, and colleague and mentor phytic species rather similar to C. buapana, including in of the second author, for his paramount contributions the fresh color, but differs from the latter in the partly to mycology in Mexico and Latin America as a whole setose lobe surface, especially towards the margins, and 404 Plant and Fungal Systematics 64(2): 393–411, 2019 in the peculiar, coralloid and often appearing spinulose Pérez-Pérez & D. León-González 1298, 1323, 1325, 1326, 1328, hyphal appendages on the lower medullary hyphae. The 1329, 1330, 1331, 1332, 1333, 1334, 1335, 1336, 1337, 1338 only other medium-sized epiphytic species with a partly (EB-BUAP, B – paratypes!). setose lobe surface and lower medullary hyphal papillae is C. garagoa from Colombia (Lücking et al. 2017b), Cora ixtlanensis Moncada, R.-E. Pérez & Lücking, which differs in the emerald-green, broadly undulate sp. nov. (Fig. 6A–C) lobes, the adnate-confluent hymenophore, and the small, MycoBank MB 832773 largely unbranched papilliform appendages on the lower medullary hyphae. Diagnosis: A medium-sized Cora growing terrestrially between bryophytes and other lichens, fresh with dark grey to olive-grey Additional specimens examined. MEXICO: Oaxaca. San lobes lacking distinct color zonation except for a darker, mar- Vicente, Nuñú, Teposcolola; 17°24′N; 97°26′W; 2616 m; coni- ginal zone exposing the photobiont (best visible when drying), fer forest (bosque de Juniperus), 14 November 2017, R.-E. with a glabrous surface and lacking papilliform appendages on

Figure 6. Habit of new Cora taxa. A–C – C. ixtlanensis (holo- and isotype), showing dry thallus in the field (A), rewetted lobes (B), and lobe underside with corticioid-concentrical hymenophore (C). D–F – C. lawreyana (holo- and isotype), showing dry thallus in the field (D), rewetted lobes (E), lobe underside with corticioid-concentrical hymenophore (F) and globose papillae on lower medullary hyphae (insets). B. Moncada et al.: The lichenized genus Cora in Mexico 405 the lower medullary hyphae; hymenophore corticioid, forming Notes. Cora ixtlanensis is closely related and very sim- resupinate, elongate, concentrically arranged patches; similar ilar to C. terrestris described from Costa Rica (Lücking to C. terrestris and differing from the latter only in molecular et al. 2017b). Both are terrestrial and form medium-sized characters. thalli and lobes with olive-grey color when fresh. They Type: Mexico: Oaxaca. Santiago Comaltepec, Cerro Pelón; also share the corticioid hymenophore with resupinate, 17°34′40.2″N; 96°30′21.6″W; 2990 m; cloud forest (bosque elongate patches in a concentric arrangement, and both mesófilo), 17 May 2015, B. Moncada & R.-E. Pérez-Pérez 10205 lack papilliform appendages on the lower medullary (EB-BUAP – holotype!; B – isotype!). hyphae. Since the two lineages are phylogenetically ITS barcoding sequence(s): KX772547 (isotype), distinct, differing consistently in five positions in the KX772539, KX772565 (paratypes). ITS, we consider this to be one of the few examples of cryptic speciation known so far within the genus. Other Description. Thallus terrestrial between bryophytes medium-sized terrestrial species with more or less grey (Breutelia, Thuidium) and other lichens (Cladonia), fresh color are C. celestinoa described from Colombian foliose, up to 15 cm across, composed of 10–20 semi- paramos and C. palustris known from montane forest in circular, adjacent to slightly imbricate lobes; individual Costa Rica, which agree in the type of hymenophore and lobes 1.5–3 cm wide and 1–2.5 cm long, rather frequently the absence of papilliform appendages on the lower med- branched at base, with short radially branching sutures, ullary hyphae and hence can also be considered cryptic, surface dark grey to olive-grey when fresh, without although their upper cortex appears more dense and not concentric color zonation, with distinct, involute, light distinctly viaduct-shaped (Lücking et al. 2017b). Notably, greyish margins, light grey when dry and in the herbar- the two are not related to each other or to C. ixtlanensis ium, without distinct concentric color zonation except for and C. terrestris. Cora ixtlanensis grows sympatrically a darker marginal zone exposing the photobiont layer. with C. casasolana and C. dewisanti subsp. mexicana; the Upper surface uneven to shallowly undulate when fresh first differs in the olive-green fresh color and the papilli- and dry, glabrous; involute margins glabrous; lower sur- form appendages on the lower medullary hyphae, whereas face ecorticate, felty-arachnoid (representing the exposed the second forms much larger thalli and lobes, with the medulla), whitish when fresh, becoming yellowish white hymenophore becoming partly cyphelloid and developed in the herbarium. Thallus in section 250–350 µm thick, mainly towards the lobe margins. with upper cortex, photobiont layer and medulla; upper cortex viaduct-shaped, formed by 20–30 µm thick layer Additional specimens examined. MEXICO: Oaxaca. San- of rather densely packed, periclinal, 4–6 µm thick hyphae tiago Comaltepec, Cerro Pelón; 17°34′40.2″N; 96°30′21.6″W; supported by 50–80 µm high ‘medullary’ layer of spaced 2990 m; cloud forest (bosque mesófilo), 17 May 2015, B. Mon- cada & R.-E. Pérez-Pérez 10264, 10293 (EB-BUAP, B – para- groups of anticlinal to oblique, 4–6 µm thick hyphae; types!). photobiont layer 100–200 µm thick, olive-green above, aeruginous-green below; medulla 30–70 µm thick, hydrophobic; clamp connections absent, papilliform hyphal Cora lawreyana Moncada, R.-E. Pérez & Lücking, appendages absent. Hymenophore corticioid, forming sp. nov. (Fig. 6D–F) resupinate, shortly elongate linear patches across lobe MycoBank MB 832774 underside in distinctly concentric arrangement, patches Diagnosis: A medium-sized (to large) Cora growing epiphyti- 3–10(–20) mm long and 0.5–1 mm broad, with whit- cally on tree trunks over bryophytes, fresh with aeruginous-grey ish (fresh) to cream- or flesh-colored (in the herbarium), to marginally olive-green lobes lacking distinct color zonation, smooth surface and glabrous to minutely pilose, slightly with a glabrous surface and with distinct, whitish, glabrous involute margins; hymenophore in section 100–150 µm to thinly pilose margins; hymenophore corticioid, forming thick, resting on thick medullary layer; hymenium com- resupinate, elongate, concentrically arranged patches; lower medullary hyphae with short, mostly subglobose appendages; posed of numerous palisade-like basidioles and scattered differing from C. buapana chiefly in the thicker, subglobose basidia; basidioles 20–30 × 5–7 µm; basidia 25–35 × hyphal appendages. 5–7 µm, 4-sterigmate; basidiospores not observed. Chem- Type: Mexico: Veracruz. La Cortadura Ecological Reserve; istry: No substances detected by TLC. 19°29′N; 97°2′W; 2088 m; cloud forest (bosque mesófilo), Etymology. We dedicate this new species to the Ixtlán 21 March 2017, R.-E. Pérez-Pérez & G. Águila-Rodríguez 1168 (EB-BUAP – holotype!; B – isotype!). community of the town of Ixtlán de Juárez in the state of Oaxaca, renowned for their sustainable ecosystem man- ITS barcoding sequence: MN475935 (isotype). agement and ecotourism (Bray 2016; Vázquez-Maguirre 2018; Pazos-Almada & Bray 2018). Description. Thallus epiphytic on branches of Baccha- ris, between mosses and liverworts (Frullania) and other Distribution and ecology. This new species is thus far lichens (Leucodermia, Parmotrema and Usnea), foliose, known from only a single location, growing terrestri- up to 10 cm across, composed of 5–10 semicircular, partly ally in an upper montane cloud forest (bosque mesófilo) imbricate lobes; individual lobes 1–3(–4) cm wide and which is thus far the location richest in documented Cora 1–3 cm long, moderately to frequently branched and lineages, with six species and one subspecies, includ- with distinct, radially branching sutures, surface dark ing the also terrestrial C. casasolana and C. dewisanti aeruginous-grey to olive-green towards margins when subsp. mexicana. fresh, without concentric color zonation, with distinct, 406 Plant and Fungal Systematics 64(2): 393–411, 2019

involute, light grey margins, becoming white to light (bosque mesófilo), 17 May 2015, B. Moncada & R.-E. Pérez- grey in the herbarium, without distinct concentric color Pérez 10316 (EB-BUAP – holotype!; B – isotype!). zonation. Upper surface uneven when fresh, ± even to ITS barcoding sequence(s): KX772564 (isotype), rugose when dry, glabrous; involute margins glabrous KX772541, KX772542, KX772553, KX772570 (para- to thinly pilose; lower surface ecorticate, felty-arach- types). noid (representing the exposed medulla), whitish when fresh, becoming yellowish white to bluish grey in the Description. Thallus epiphytic on tree trunks, often herbarium. Thallus in section 200–250 µm thick, with overgrowing mosses and liverworts (including Frulla- upper cortex, photobiont layer, and medulla; upper cor- nia, Metzgeria, Plagiochila), foliose, up to 3 cm across, tex viaduct-shaped, formed by 10–20 µm thick layer of composed of 5–10 semicircular, adjacent to subimbri- loosely packed, periclinal, 4–6 µm thick hyphae sup- cate lobes; individual lobes 0.5–1(–1.5) cm wide and ported by 30–50 µm high ‘medullary’ layer of spaced 0.5–0.7(–1) cm long, sparsely branched, without radially groups of irregularly arranged to anticlinal, 4–6 µm thick branching sutures, surface dark forest-green when fresh, hyphae; photobiont layer 50–80 µm thick, aeruginous- to without concentric color zonation, with distinct, involute emerald-green; medulla 50–100 µm thick, hydrophobic; margins covered by lighter olive-brown to olive-grey sore- clamp connections absent but lower medullary hyphae dia, surface becoming white-grey and soredia becoming with numerous papilliform, unbranched, mostly glo- dark olive-grey in the herbarium. Upper surface uneven bose hyphal appendages 2–3(–5) µm long and 2–3 µm to shallowly narrowly undulate when fresh, undulate thick. Hymenophore corticioid, distinctly concentrically to rugose when dry, glabrous, with diffuse concentric arranged, forming resupinate, elongate to linear, partly bands of scattered to dense soredia; involute margins branched to anastomosing patches across lower lobe sur- glabrous, densely sorediate; lower surface ecorticate, face, patches 1–10 mm long and 0.5–1 mm broad, with appressed felty-arachnoid (representing the exposed white (fresh) to cream-colored (in the herbarium), smooth medulla) and with photobiont layer partially protruding, surface and glabrous to minutely pilose, involute margins; whitish when fresh, becoming light yellowish grey in hymenophore in section 100–120 µm thick, resting on the herbarium. Thallus in section 250–300 µm thick, medullary layer or entirely protruding; hymenium com- with upper cortex, photobiont layer and medulla; upper posed of numerous palisade-like basidioles and basidia; cortex viaduct-shaped, formed by 20–30 µm thick layer basidioles 20–35 × 5–7 µm; basidia 30–40 × 5–7 µm, of loosely packed, periclinal, 4–6 µm thick hyphae sup- 4-sterigmate; basidiospores not observed. Chemistry: No ported by 50–100 µm high ‘medullary’ layer of spaced substances detected by TLC. groups of irregularly arranged to oblique, 4–6 µm thick hyphae; photobiont layer 80–150 µm thick, olive-green Etymology. We are delighted to dedicate this new species above, aeruginous-green below, soredia aeruginous-green; to our colleague and friend James D. (‘Jim’) Lawrey on medulla 30–50 µm thick; clamp connections absent but the occasion of his 70th birthday. Jim has made numer- lower medullary hyphae with numerous papilliform, ous contributions to lichenology in such diverse fields as mostly unbranched hyphal appendages 3–5 µm long and ecology, lichenicolous fungi, and the evolution of basid- 1.5–3 µm thick. Hymenophore not observed. Chemistry: iolichens and their photobionts. No substances detected by TLC.

Distribution and ecology. Known from only a single Etymology. We dedicate this new species to our col- locality in cloud forest (bosque mesófilo) in the state of league, mentor and friend María de los Ángeles Herrera- Veracruz. Campos (‘Marusa’) for her important contributions to the development of modern lichenology in Mexico. Notes. Cora lawreyana is similar to the apparently allopatric C. buapana and differs from the latter and other similar Distribution and ecology. Found sympatrically with species (discussed under C. buapana) chiefly in the mostly Cora benitoana and C. buapana (see above) in Oax- globose appendages on the lower medullary hyphae. aca, growing epiphytically on tree trunks in cloud forest (bosque mesófilo) between 2000 and 3000 m. Cora marusae Moncada, R.-E. Pérez & Lücking, sp. nov. (Fig. 7A–B) Notes. Cora marusae is morphologically and anatomically similar to three other small (to medium-sized) epiphytic MycoBank MB 832775 species producing soredia but with a glabrous, undulating Diagnosis: A small, epiphytic Cora with abundant, marginal lobe surface and featuring papilliform appendages on the and partly laminal soredia and marked color contrast between lower medullary hyphae. All three are olive-green rather fresh and dry lobes: fresh lobes with dark forest-green color and than forest-green, although this difference may be difficult lighter olive-brown to olive-grey soredia, and dry lobes with white-grey color and dark olive-grey soredia; lower medullary to discern. The hymenophore in the somewhat related hyphae with papilliform appendages; differing from the morpho- C. rothesiorum known from Colombia is also unknown, logically cryptic but phylogenetically unrelated C. comaltepeca but that species differs from C. marusae in habitat ecol- in the dark forest-green rather than olive-green lobes when fresh, ogy, being found in paramo vegetation (Lücking et al. and in the much more abundant soredia. 2017b). In both C. comaltepeca (Mexico) and C. soreda- Type: Mexico: Oaxaca. Santiago Comaltepec, Cerro Las vidia (Costa Rica and Brazil) the hymenophore is known Antenas; 17°35′16.5″N; 96°27′2.7″W; 2190 m; cloud forest and is distinct in the two species: corticioid-concentric B. Moncada et al.: The lichenized genus Cora in Mexico 407 in the first, and cyphelloid and disc-shaped in the second of soredia also on the lobe surface. The taxon most likely (Lücking et al. 2017b). Both are unrelated to C. maru- to be confused is the sympatric C. comaltepeca (Lücking sae and C. rothesiorum. Thus, hymenophore type may et al. 2017b), which is, however, phylogenetically very provide a good diagnostic character in these otherwise distinctive; it usually produces only a few soredia but highly similar species. Cora minor and C. paraminor the available material so far is too sparse to assess this are closely related to C. marusae and also feature small variation quantitatively. In both species the color contrast sorediate lobes, but the soredia are marginal and the two between fresh and dry thalli is also remarkable, with the species lack hyphal appendages on the lower medullary soredia being lighter than the lobe surface in the hydrated hyphae. Within Mexico, C. marusae is a very character- stage and becoming much darker in the herbarium. Cora istic and readily recognized species, due to the heavily totonacorum (see below) is also similar but produces larger sorediate margins and the formation of concentric bands lobes; it is not closely related to C. marusae.

Figure 7. Habit of new Cora taxa. A, B – C. marusae (A isotype, B Moncada & Pérez-Pérez 10178), showing rewetted (A) and dry (B) lobes with largely marginal soredia. C, D – C. totonacorum (isotype), showing rewetted (C) and dry (D) lobes with largely marginal soredia; lobe in D with attack by unidentified fungus. E, F – C. zapotecorum (isotype), showing rewetted lobes with thin setae (E) and lobe underside with corticioid-concentrical, partly confluent hymenophore (F). 408 Plant and Fungal Systematics 64(2): 393–411, 2019

Additional specimens examined. MEXICO: Oaxaca. San- Distribution and ecology. Known from only a single tiago Comaltepec, Cerro Pelón; 17°34′40.2″N; 96°30′21.6″W; locality in cloud forest (bosque mesófilo) in the state of 2990 m; cloud forest (bosque mesófilo), 17 May 2015, B. Veracruz, growing sympatrically with C. lawreyana. Moncada & R.-E. Pérez-Pérez 10178, 10212, 10222, 10319 (EB-BUAP, B – paratypes!). Notes. Thus far there are three other medium-sized epiphytic species known to produce soredia and to feature Cora totonacorum Moncada, R.-E. Pérez & Lücking, papilliform appendages on the lower medullary hyphae: sp. nov. (Fig. 7C–D) Cora davidia (Colombia to Ecuador), C. inversa (Colom- MycoBank MB 832776 bia) and the widespread C. hawksworthiana (Costa Rica Diagnosis: A medium-sized Cora growing epiphytically on tree to Chile). All three occur in paramo to puna vegetation trunks, fresh with olive-grey to olive-green lobes with some or temperate grasslands, but agree with C. totonacorum color zonation, with a glabrous surface and with marginal in the more or less olive-green color. Cora inversa differs soredia; lower medullary hyphae with papilliform appendages; from the other three species in the rather large papilliform similar and related to the allopatric C. davidia but differing appendages on the lower medullary hyphae, whereas the phylogenetically in a large number of substitutions and indels. remaining three taxa are more or less cryptic morpholog- Type: Mexico: Veracruz. La Cortadura Ecological Reserve; ically and anatomically. Whereas C. hawksworthiana and 19°29′N; 97°2′W; 2088 m; cloud forest (bosque mesófilo), C. inversa are only distantly related, C. davidia is close to 21 March 2017, R.-E. Pérez-Pérez & G. Águila-Rodríguez C. totonacorum, although the two differ phylogenetically 1189b (EB-BUAP – holotype!; B – isotype!). in 13 substitutions and two indels in the ITS. ITS barcoding sequence: MN475934 (isotype). Cora zapotecorum Moncada, R.-E. Pérez & Lücking, Description. Thallus epiphytic on branches of Bac- sp. nov. (Fig. 7E–F) charis, between mosses and liverworts (Frullania) and others lichens (Leucodermia, Parmotrema and Usnea), MycoBank MB 832777 foliose, up to 7 cm across, composed of 3–5 semicir- Diagnosis: A small (to medium-sized) Cora growing epi- cular, adjacent to subimbricate lobes; individual lobes phytically on branches of trees or shrubs over and between 1–3 cm wide and 1–2 cm long, ascending, sparsely to bryophytes, fresh with olive-grey lobes lacking distinct color zo- moderately branched, without distinct, radially branch- nation, with a mostly glabrous surface and with concentric areas ing sutures, surface dark olive-grey to olive-green when of appressed setae (erect when wet), and with distinct, whitish, fresh (towards margins), with some concentric color glabrous to thinly pilose margins; hymenophore ± cyphelloid, zonation alternating with lighter grey zones, with thin, forming rounded to elongate patches; differing fromC. hochesuordensis and C. schizophylloides in the very thinly pilose lobe flat to involute margins covered by olive-green soredia, surface and the corticioid-concentrical hymenophore. surface becoming white-grey and soredia becoming dark olive-grey in the herbarium. Upper surface even to very Type: Mexico: Oaxaca. Santiago Comaltepec, Cerro Pelón; 17°34′40.2″N; 96°30′21.6″W; 2990 m; cloud forest (bosque shallowly undulate when fresh, shallowly undulate to mesófilo), 17 May 2015, B. Moncada & R.-E. Pérez-Pérez 10206 rugose when dry, glabrous or with scattered, thin, white (EB-BUAP – holotype!; B – isotype!). setae towards margins; involute margins glabrous to minutely pilose; lower surface ecorticate, felty-arach- ITS barcoding sequence(s): KX772568 (isotype), noid (representing the exposed medulla), whitish when KX772546, KX772566 (paratypes). fresh, becoming light grey in the herbarium. Thallus in section 250–300 µm thick, with upper cortex, photobiont Description. Thallus epiphytic on branches of trees and layer and medulla; upper cortex distinctly viaduct-shaped, shrubs, usually over and between mosses and liverworts formed by 30–50 µm thick layer of rather densely packed, (often Frullania), foliose, up to 8 cm across, composed periclinal, 4–6 µm thick hyphae supported by 50–70 µm of 5–15 semicircular, distant to slightly imbricate lobes; high ‘medullary’ layer of loosely arranged, ± anticlinal, individual lobes 0.5–1.5 cm wide and 0.5–1 cm long, fre- 4–6 µm thick hyphae; photobiont layer 50–100 µm thick, quently branched at margins to appear marginally crenate olive-green above, aeruginous-green below; medulla to lobate, without radially branching sutures, surface dark 30–60 µm thick; clamp connections absent but lower olive-grey when fresh, without concentric color zonation, medullary hyphae with numerous papilliform, mostly with distinct, involute, light greyish margins, white-grey unbranched hyphal appendages 3–5 µm long and 2–3 µm when dry and in the herbarium, without distinct concentric thick. Hymenophore not observed. Chemistry: No sub- color zonation. Upper surface uneven when fresh and dry, stances detected by TLC. mostly glabrous but with concentric areas of strongly appressed setae arranged radially, becoming erect when Etymology. The epithet honors the indigenous Totonac wet; involute margins glabrous to thinly pilose; lower sur- people, who inhabit the states of Veracruz, Puebla and face ecorticate, felty-arachnoid (representing the exposed Hidalgo. Their language belongs to a small, isolated lan- medulla), whitish when fresh, becoming yellowish white guage family without any known relatives (Sandstrom in the herbarium. Thallus in section 300–350 µm thick, & García-Valencia 2005; Mackay & Trechsel 2018). with upper cortex, photobiont layer and medulla; upper During the 19th century the Totonac were the most cortex viaduct-shaped, formed by 20–30 µm thick layer important producers of vanilla (Rain & Lubinsky 2011; of rather densely packed, irregularly arranged to peri- Cameron 2012). clinal, 4–6 µm thick hyphae supported by 50–100 µm high B. Moncada et al.: The lichenized genus Cora in Mexico 409

‘medullary’ layer of spaced groups of anticlinal, 4–6 µm 2(1) Thallus and lobes medium-sized; mature lobes up to thick hyphae; photobiont layer 150–200 µm thick, olive- 3 cm broad ...... C. totonacorum green above, aeruginous-green below; medulla 30–70 µm Thallus and lobes small, mature lobes usually up to 1 cm thick, hydrophobic; clamp connections absent but lower broad, rarely up to 1.5 cm ...... 3 medullary hyphae with scattered, papilliform, (sparsely) 3(2) Soredia abundant and dense; hymenophore unknown branched hyphal appendages 3–5 µm long and 2–4 µm ...... C. marusae thick. Hymenophore corticioid, forming resupinate, elon- Soredia sparse; hymenophore usually present, corticioid, gate to irregularly rounded patches across lobe underside, forming resupinate, elongate, concentrically arranged in more or less concentric arrangement and often closely patches ...... C. comaltepeca adjacent to confluent, patches 2–5 mm long and 1–2 mm broad, with white (fresh) to cream-colored or flesh-colored 4(1) Thallus epiphytic, often overgrowing bryophytes . .5 (in the herbarium), smooth surface and glabrous, slightly Thallus terrestrial, growing between bryophytes and other involute margins; hymenophore in section 100–130 µm lichens ...... 9 thick, resting on medullary layer; hymenium composed of 5(4) Thallus and lobes small, mature lobes usually up to numerous palisade-like basidioles and scattered basidia; 1 cm broad, rarely up to 1.5 cm; lobe surface with basidioles 20–30 × 5–7 µm; basidia 25–35 × 5–7 µm, thin, concentric bands of scattered setae; lower med- 4-sterigmate; basidiospores not observed. Chemistry: No ullary hyphae with papilliform hyphal appendages . substances detected by TLC...... C. zapotecorum Thallus and lobes medium-sized to large, mature lobes Etymology. The epithet of this new species honors the usually over 2 cm broad; lobe surface in most species Zapotec people, one of the most important indigenous glabrous; if concentric bands of setae present, then pho- civilizations in Mesoamerica, concentrated in the area tobiont layer enclosed by upper and lower layer of pig- today covered by the state of Oaxaca. The cultural history mented hyphae and lower medullary hyphal appendages of the Zapotec goes back to 500 BCE and perhaps as far coralloid-spinulose ...... 6 th back as 1150 BCE, and lasted until the early 16 century 6(5) Hymenophore cyphelloid, developed along lobe margins, (Marcus & Flannery 1996). in part strongly protruding to form flabelliform basidiocarps with setose upper surface; thallus and lobes large, Distribution and ecology. Found sympatrically with mature lobes 3–6 cm broad, with distinctly narrowly Cora benitoana, C. buapana and C. marusae (see above) undulate surface; lower medullary hyphae without papill- in Oaxaca, growing epiphytically on branches in cloud iform hyphal appendages ...... C. benitoana forest (bosque mesófilo) between 2000 and 3000 m. Hymenophore corticioid, developed across lobe underside and forming elongate, concentrically arranged, resu- Notes. Thus far, only two other small (to medium-sized) pinate patches; thallus and lobes medium-sized, mature epiphytic species are known to produce setae on the lobe lobes usually up to 3 cm, rarely up to 4 cm broad, with surface and feature papilliform appendages on the lower uneven to at most very shallowly undulate surface; medullary hyphae: C. hochesuordensis from the Boliv- lower medullary hyphae with distinct, papilliform hyphal ian puna and C. schizophylloides from the Colombian appendages ...... 7 paramo (Lücking et al. 2017b). Both differ in the more 7(6) Lobe surface with concentric bands of scattered, erect densely pilose-setose lobe surface and the distinctly setae; lower medullary hyphal appendages coral- cyphelloid hymenophore. Within Mexico, C. zapote- loid-spinulose ...... C. guzmaniana corum is most similar to the sympatric C. marusae but Lobe surface glabrous; lower medullary hyphal append- lacks soredia and differs in the partly pilose-setose lobe ages mostly unbranched and short ...... 8 surface. 8(7) Lower medullary hyphal appendages short, mostly glo- Additional specimens examined. MEXICO: Oaxaca. bose ...... C. lawreyana Santiago Comaltepec, Cerro Las Antenas; 17°35′16.5″N; Lower medullary hyphal appendages elongate and fin- 96°27′2.7″W; 2190 m; cloud forest (bosque mesófilo), 17 May ger-like or flask-shaped, partly branched . C. buapana 2015, B. Moncada & R.-E. Pérez-Pérez 10307 (EB-BUAP, B – paratypes!). Santiago Comaltepec, Cerro Pelón; 17°34′40.2″N; 9(4) Hymenophore corticioid-cyphelloid, developed as indi- 96°30′21.6″W; 2990 m; cloud forest (bosque mesófilo), 17 May vidual patches towards margins; thallus and lobes very 2015, B. Moncada & R.-E. Pérez-Pérez 10318 (EB-BUAP, large, mature lobes 3–8 cm broad; lower medullary B – paratypes!). hyphae without hyphal appendages ...... C. dewisanti subsp. mexicana Key to the species of Cora thus far known Hymenophore when present corticioid, developed across from southeastern Mexico entire lobe underside and forming elongate, concentrically arranged, resupinate patches; thallus and lobes 1 Lobes with marginal (to laminal) soredia (sometimes medium-sized, mature lobes usually up to 3 cm broad, sparse); thallus always epiphytic; lower medullary hyphae rarely up to 4 cm; lower medullary hyphae with or with- always with papilliform hyphal appendages . . . . 2 out hyphal appendages ...... 10 Lobes lacking soredia; thallus epiphytic or terrestrial; 10(9) Lobes dark grey to olive-grey when fresh; lower med- lower medullary hyphae with or without papilliform ullary hyphae without hyphal appendages . . . . . hyphal appendages ...... 4 ...... C. ixtlanensis 410 Plant and Fungal Systematics 64(2): 393–411, 2019

Lobes olive-green to aeruginous-green when fresh; Moncada, B., Nuankaew, S., Parnmen, S., de Azevedo Santiago, lower medullary hyphae with papilliform hyphal A. L. C. M., Sommai, S., Song, Y., de Souza, C. A. F., de Sou- appendages ...... 11 za-Motta, C. M., Su, H. Y., Suetrong, S., Wang, Y., Wei, S. -F., Wen, T. C., Yuan, H. S., Zhou, L. W., Réblová, M., Fournier, J., 11(10) Lobes aeruginous-green when fresh; growing in montane Camporesi, E., Luangsa-ard, J. J., Tasanathai, K., Khonsanit, A., forest; upper cortex collapsed-compacted; hymenophore Thanakitpipattana, D., Somrithipol, S., Diederich, P., Millanes, corticioid, forming elongate, concentrically arranged, A. M., Common, R. S., Stadler, M., Yan, J. Y., Li, X., Lee, H. W., resupinate patches ...... C. dulcis Nguyen, T. T. T., Lee, H. B., Battistin, E., Marsico, O., Vizzini, A., Vila, J., Ercole, E., Eberhardt, U., Simonini, G., Wen, H. -A., Lobes olive-green (to olive-grey) when fresh; growing Chen, X. -H. 2015. Fungal diversity notes 111–252 – taxonomic in subalpine vegetation; upper cortex viaduct-shaped; and phylogenetic contributions to fungal taxa. Fungal Diversity hymenophore unknown ...... C. casasolana 75: 27–274. Bray, D. B. 2016. Muir and Pinchot in the Sierra Norte of Oaxaca: Acknowledgements Governance of forest management and forest recovery in Pueblos Mancomunados. World Development Perspectives 4: 8–10. This study was supported in part by three grants from the Na- Cameron, K. 2012. Vanilla Orchids: Natural History and Cultivation. tional Science Foundation: TICOLICHEN – The Costa Rican Timber Press, Portland, OR. Lichen Biodiversity Inventory (DEB 0206125 to The Field Mu- Chaves, J. L., Lücking, R., Sipman, H. J. M., Umaña, L. & Navarro, E. seum; PI Robert Lücking), Neotropical Epiphytic Microlichens 2004. A first assessment of the Ticolichen biodiversity inventory – An Innovative Inventory of a Highly Diverse yet Little Known in Costa Rica: the genus Dictyonema (Polyporales: Atheliaceae). Group of Symbiotic Organisms (DEB 0715660 to The Field The Bryologist 107: 242–249. Museum; PI R. Lücking), and Phylogenetic Diversity of Myco- Cornejo, C. & Scheidegger, C. 2016. Cyanobacterial gardens: the liver- bionts and Photobionts in the Cyanolichen Genus Dictyonema, wort Frullania asagrayana acts as a reservoir of lichen photobionts. with Emphasis on the Neotropics and the Galapagos Islands Environmental Microbiology Reports 8: 352–357. (DEB 0841405 to George Mason University; PI J. Lawrey; Dal Forno, M., Lawrey, J. D., Sikaroodi, M., Bhattarai, S., Gillevet, Co-PIs: R. Lücking, P. Gillevet). We thank Dr. Gonzalo Cas- P. M., Sulzbacher, M. & Lücking, R. 2013. Starting from scratch: tillo-Campos from INECOL, A.C. (Instituto de Ecología, A.C.) evolution of the lichen thallus in the basidiolichen Dictyonema for financial support to study lichens in La Cortadura Ecolog- (Agaricales: Hygrophoraceae). Fungal Biology 117: 584–598. ical Reserve. Comparative material from Costa Rica was also Dal Forno, M., Bungartz, F., Yánez-Ayabaca, A., Lücking, R. & Lawrey, collected during two lichen courses of the Organization for J. D. 2017. High levels of endemism among Galapagos basidioli- Tropical Studies (OTS). chens. Fungal Diversity 85: 45–73. García, A. Z., Nash, T. H. III and Herrera-Campos, M. A. 2000. Lichen Supplementary electronic material decline in desierto de los leones (Mexico City). The Bryologist 103: 428–442. Figure S1. Complete, best-scoring phylogenetic ML tree of the ITS Guzmán, G. 2008. Diversity and use of traditional Mexican medicinal sequences of Cora used in this study. Supported branches are thickened fungi. A review. International Journal of Medicinal Mushrooms and bootstrap support values are indicated above the branches. Samples 10: 209–217. from Mexico are highlighted in orange. Download file Guzmán-Dávalos, L. 2016. Gastón Guzmán (1932–2016). Acta Botánica Table S1. Genbank accession numbers and geographic origin for the Mexicana 115: 5–8. ITS sequences of Cora used in this study. Download file Guzmán-Dávalos, L. & Cifuentes, J. 2018. Gastón Guzmán, 26 August Table S2. Georeference data for specimens of Cora reported from 1932–12 January 2016. Mycologia 110: 791–794. Mexico used to generate the map in Fig. 3, including specimens available through GBIF and CNALH and the specimens analysed (sequenced) in Hall, T. A. 1999. BioEdit: a user-friendly biological sequence alignment the present work. Download file editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98. File S1. Complete alignment of the ITS sequences of Cora used in this Imshaug, H. A. 1956. 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