A First Phylogenetic Assessment of Dictyonema S.Lat. in Southeastern North America Reveals Three New Basidiolichens, Described in Honor of James D

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Plant and Fungal Systematics 64(2): 383–392, 2019 ISSN 2544-7459 (print) DOI: 10.2478/pfs-2019-0025 ISSN 2657-5000 (online)

A first phylogenetic assessment of Dictyonema s.lat. in southeastern North America reveals three new basidiolichens, described in honor of James D. Lawrey

Manuela Dal Forno1,2*, Laurel Kaminsky 3, Roger Rosentreter 4, R. Troy McMullin5, André Aptroot 6 & Robert Lücking7,8

Abstract. Three species of lichenized basidiomycetes in the Dictyonema clade from south- Article info eastern North America are described as new to science: Cyphellostereum georgianum, Received: 21 May 2019 C. jamesianum and Dictyonema lawreyi, all with a crustose-filamentous growth form. Revision received: 23 Aug. 2019 Based on ITS sequences, the species form well-supported monophyletic clades in a phy- Accepted: 25 Aug. 2019 logeny and are represented by at least two specimens each. They are also distinguishable Published: 2 Dec. 2019 by morphological and anatomical characters. These new findings emphasize the importance Associate Editor of lichenological studies in North America, especially in historically understudied taxo- Adam Flakus nomic groups, such as basidiolichens. This study is dedicated to James D. Lawrey on the occasion of his 70th birthday. Key words: Lichen systematics, Hygrophoraceae, United States, biodiversity

Introduction

The Dictyonema clade is the largest group of basidiolichens 2013; Lücking et al. 2014, 2017b). The clade currently (Lücking et al. 2017a). It belongs to tribe Arrheniae, sub- has 137 accepted species in five genera: Acantholichen, family Lichenomphaloideae and family Hygrophoraceae Cora, Corella, Cyphellostereum and Dictyonema (s.str.); in the order Agaricales (Lawrey et al. 2009; Lodge et al. however, a few historical genus names that have an uncer- 2014). Hygrophoraceae comprises over 600 species, most tain taxonomic status persist (Lücking et al. 2013). All five of which are non-lichenized, except for the Dictyonema of the accepted genera occur in the New World, whereas clade and the genus Lichenomphalia (Lodge et al. 2014). other regions almost exclusively feature Cyphellostereum Species in the Dictyonema clade occur worldwide but the and Dictyonema. highest diversity is in the Neotropics (Dal Forno et al. Aside from Mexico, the Dictyonema clade is sparsely represented in North America. In the ‘Cumulative Check- list for the Lichen-forming, Lichenicolous and Allied 1 Department of Botany, Smithsonian Institution, National Museum of Fungi of the Continental United States and Canada’ th Natural History, 10 St & Constitution Ave NW, Washington, DC, (Esslinger 2018), four taxa from this clade are listed: 20560, USA 2 Research Associate, Botanical Research Institute of Texas, Fort Worth, Cora glabrata [syn.: C. pavonia, Dictyonema glabratum, TX, 76107, USA D. pavonium], D. moorei, D. phyllogenum and D. seri- 3 Digitization Coordinator, McGuire Center for Lepidoptera and Bio- ceum. Two additional names on the list are considered diversity, Florida Museum of Natural History, University of Florida, synonyms or misidentifications: Dictyonema guadalu- Gainesville, FL 32611, USA 4 Dept of Biology, Boise State University, 1910 University Ave, Boise pense [= D. sericeum] and D. irpicinum (Esslinger 2018). ID, 83725 The genera Acantholichen, Corella and Cyphellostereum 5 Canadian Museum of Nature, Research and Collections, PO Box 3443, were not reported in Esslinger’s list but D. phyllogenum Station D, Ottawa, ON, K1P 6P4, Canada is now classified in Cyphellostereum (Lücking et al. 6 Laboratório de Botânica / Liquenologia, Instituto de Biociências, Universidade Federal de Mato Grosso do Sul, Avenida Costa e Silva 2013), so this genus is recognized from North America. s/n, Bairro Universitário, CEP 79070-900, Campo Grande, MS, Brazil. The taxonomic status of D. guadalupense is unclear but 7 Botanical Garden and Botanical Museum Berlin, Königin-Luise-Straße certainly it is not a synonym of D. sericeum (Lücking 6–8, 14195 Berlin, Germany et al. 2013). Given that the Dictyonema clade has been 8 Research Associate, Science & Education, The Field Museum, 1400 South Lake Shore, Chicago, IL, 60605, USA shown to contain many more species than proposed by * Corresponding author e-mail: [email protected] Parmasto (1978), including several hundred alone in the

This work is licensed under the Creative Commons BY-NC-ND 4.0 License 384 Plant and Fungal Systematics 64(2): 383–392, 2019

genera Cora and Dictyonema (Lücking et al. 2014, 2017b; and obtained molecular data to aid in their classification. Dal Forno 2015), it is safe to say that, with the possible We detected three distinct lineages, all new to science exception of D. moorei, none of the names listed for North – two in the genus Cyphellostereum and one in Dic- America actually represent an accurate identification of tyonema, which are described here. the taxa occurring in this region. th As of February 20 , 2019, the ‘Consortium of North Material and methods American Lichen Herbaria’ (CNALH; http://lichenportal. org/) contained 111 records of continental United States Taxon sampling and phylogenetic analysis. This specimens identified as Cora or Dictyonema, with the study is part of a broad systematics study of the Dic- exception of one misidentification ofAcantholichen , which tyonema clade. Most specimens from North America we revised. All collections are from the southeastern United included here were already sequenced and included in States and the majority of them are from Florida (FL, 90 prior phylogenetic studies (Dal Forno 2015; Dal Forno total, 31 in Cora and 59 in Dictyonema), with a few speci- et al. 2017). However, to account for newly available ITS mens from North Carolina (NC, 2), South Carolina (SC, 7), sequences not included in previously published datasets, Georgia (GA, 4), and Louisiana (LA, 5), all in Dictyonema. we generated new MAFFT v1.3.7 alignments (Katoh A single specimen (Dictyonema sp.) is mentioned from & Toh 2005) on Geneious v11.1.5 (Kearse et al. 2012; Texas, and two samples lack information regarding their File S1 and File S2) and ran new phylogenetic analyses state provenance. Different names have been applied to on RAxML-HPC2 on XSEDE (v8.2.10) in the CIPRES identify these specimens: Cora glabrata and C. pavonia portal (Miller et al. 2010), following the same laboratorial (FL), ‘Dictyonema cincinnatum’ (as ‘Dichonema cincin- and computational procedures as described in Dal Forno natum’; FL, SC), D. glabratum (FL, ≡ Cora glabrata), (2015) and Dal Forno et al. (2013, 2017). Generated trees D. guadalupense (FL), D. irpicinum (FL, GA), D. moorei were visualized in FigTree v1.4.3 and edited in Photoshop (FL), D. pavonium (FL, ≡ Cora pavonia), D. reticulatum Illustrator. GenBank numbers for the new species are (SC), D. sericeum (FL, GA, LA, NC, SC) and D. spon- listed in the taxonomy section and in Table S1. giosum (FL). Dictyonema sericeum is also cited for Ala- bama by Parmasto (1978). These names agree with those Morphological analysis. Morphological characters were included in Esslinger’s list (2018), with the exception analyzed under a Leica M165 C stereoscope following of D. spongiosum and the apparently unpublished name standard procedures for the group (Lücking et al. 2013). ‘Dichonema cincinnatum’, interpreted in CNALH as ‘Dic- Anatomical thallus characters were investigated by light tyonema cincinnatum’. The latter is based on two collec- microscopy using a Leica DM4 B compound microscope, tions by the same collector (B. M. Davis) from FL and with thallus pieces mounted in tap water. Microscopic SC and likely a herbarium-working name. It should not be measurements were made at 400× and 1000× in water. confused with the cyanobacterium Scytonema cincinnatum Images were taken with the aforementioned microscopes described from France, neither is it derived from the city of with a Leica DFC450 digital camera and processed with Cinncinati in Ohio; rather, the epithet cincinnatum (Latin) Leica Application Suite (LAS) X software. In addition, refers to ‘curly hair’, i.e. the typical morphology of many we used an Olympus DSX 100 compound microscope filamentous Dictyonema species. with Microscope Control v2.1.5 software to generate Given the much-revised phylogenetic classification stacked images. of genera and species in the Dictyonema clade (Lücking et al. 2013, 2017b; Dal Forno et al. 2013, 2017; Dal Results and discussion Forno 2015) and that identifications of North American material rely almost entirely on Parmasto’s (1978) out- Maximum likelihood phylogenetic analysis based on dated monograph, virtually nothing is known about the the ITS fungal barcoding marker grouped the sequenced diversity and range of this clade in the United States. material into three distinct, strongly supported lineages The only reported species that might actually be present (Figs. 1, 2). Two lineages belong to the genus Cyphelloste- in North America north of Mexico is the demonstrably reum, and the two form a strongly supported sister group widespread Dictyonema moorei, which is well charac- relationship (Fig. 1), indicating a strong geographic signal terized by 2–3 rows of cyanobacterial cells inside the for this clade. The closest relative is C. imperfectum from fungal sheath. Dictyonema irpicinum is restricted to the Guatemala but this relationship is not well supported. The Paleotropics and D. guadelupense likely to the Caribbean, third lineage is a new species in Dictyonema s.str., related whereas D. sericeum and D. spongiosum form shelf-like to two undescribed species from Brazil and Costa Rica thalli and also appear to be restricted to the Caribbean (Fig. 2). The relationship between this new species of (Lücking et al. 2013). The genus Cora has so far only Dictyonema from the United States and the species from been found in Florida. and all specimens studied by us Brazil is not well supported, although the clade with these belong to a single undescribed species that will be treated species and the taxon from Costa Rica is. in a separate paper. Compared to the 111 collections of Dictyonema s.lat. In order to achieve a more accurate picture of the listed in CNALH (see above), the eight specimens stud- diversity and taxonomy of filamentous species of Dic- ied here with molecular data are a small subset covering tyonema s.lat. present in North America, we collected the easternmost part of the reported distribution range fresh material throughout the southeastern United States of the Dictyonema clade in southeastern North America. M. Dal Forno et al.: A fi rst phylogenetic assessment of Dictyonema s.lat. in southeastern North America 385

OUTGROUP_Eonema_pyriformis_Hjm_18581_SWED_EU118605 MDF712_Cyphellostereum_spec_Luecking_36060_HAWA MDF426_Cyphellostereum_spec_DalForno_1798_ECUA MDF350_Cyphellostereum_spec_DalForno_1740_CORI MDF433_Cyphellostereum_spec_DalForno_1825_ECUA MDF461_Cyphellostereum_spec_DalForno_1926_ECUA MDF457_Cyphellostereum_spec_DalForno_1915_ECUA MDF462_Cyphellostereum_spec_DalForno_1927_ECUA DIC333_Cyphellostereum_spec_Luecking_17013_CORI R04_Cyphellostereum_spec_Luecking_sn_CORI_EU825976 MDF329_Cyphellostereum_spec_DalForno_1729_CORI NSF324_Cyphellostereum_pusiolum_DalForno_3051_JAM DNA7668_Cyphellostereum_spec_Moncada_6684_COLO DNA7678_Cyphellostereum_spec_Moncada_6708_COLO MON1746_Cyphellostereum_spec_Coca_sn_COLO MON1747_Cyphellostereum_spec_Coca_sn_COLO MDF526_Cyphellostereum_spec_DalForno_2026_BRAZ DIC511_Cyphellostereum_spec_Luecking_33338_COLO DIC441_Cyphellostereum_spec_Luecking_34061_COLO MDF606_Cyphellostereum_spec_DalForno_2056_BRAZ MDF664_Cyphellostereum_spec_Spielmann_sn_BRAZ MDF662_Cyphellostereum_spec_Spielmann_sn_BRAZ MDF663_Cyphellostereum_spec_Spielmann_sn_BRAZ MDF610_Cyphellostereum_spec_DalForno_2064_BRAZ MDF375_Cyphellostereum_spec_DalForno_2430_BRAZ DNA7757b_Cyphellostereum_spec_Sulzbacher_1480_BRAZ DNA7758t_Cyphellostereum_spec_Sulzbacher_1480_BRAZ DIC115_Cyphellostereum_imperfectum_Luecking_Type_GUAT MDF680_Cyphellostereum_georgianum_McMullin_12290_USA MDF295_Cyphellostereum_georgianum_Rosentreter_17755_USA MDF561_Cyphellostereum_jamesianum_Harris_59781b_USA MDF563_Cyphellostereum_jamesianum_Aptroot_72323_PUERI LK370_Cyphellostereum_jamesianum_Kaminsky_370_USA DIC127_Cyphellostereum_spec_RivasPlata_2138a_PHIL MDF431_Cyphellostereum_spec_DalForno_1813_ECUA MDF567_Cyphellostereum_spec_DalForno_2203_PUERI_JCF67IH01CC9YM MDF417_Cyphellostereum_spec_DalForno_1797_ECUA MDF415_Cyphellostereum_spec_DalForno_1792_ECUA MDF358_Cyphellostereum_spec_DalForno_1758_CORI MDF323_Cyphellostereum_spec_DalForno_1723_CORI MDF320_Cyphellostereum_spec_DalForno_1720_CORI MDF460_Cyphellostereum_spec_DalForno_1923_ECUA DIC158_Cyphellostereum_phyllogenum_Lumbsch_sn_FIJI MDF176_Cyphellostereum_unoquinoum_Bungartz_9475_GALA MDF356_Cyphellostereum_spec_DalForno_1756_CORI MDF354_Cyphellostereum_spec_DalForno_1754_CORI MDF120_Cyphellostereum_galapagoense_Bungartz_8517_GALA MDF126_Cyphellostereum_galapagoense_Yanez_1545_GALA MDF721_Cyphellostereum_spec_Luecking_36242_HAWA MDF703_Cyphellostereum_spec_Luecking_35632_HAWA MDF704_Cyphellostereum_spec_Luecking_35633_HAWA MDF720_Cyphellostereum_spec_Luecking_36241_HAWA MDF719_Cyphellostereum_spec_Luecking_36240_HAWA MDF722_Cyphellostereum_spec_Luecking_36243_HAWA MDF723_Cyphellostereum_spec_Luecking_36244_HAWA MDF717_Cyphellostereum_spec_Luecking_36238_HAWA MDF710_Cyphellostereum_spec_Luecking_35898a_HAWA

0.2 Figure 1. Phylogenetic tree of the genus Cyphellostereum inferred by maximum likelihood in RAxML using ITS fungal barcoding (n = 57, alignment 864 bp). Bolded branches have bootstrap values (BS) over 85% and are considered well-supported.

The sequences generated from the North American mate- The eight samples included in our study represent rial did not cluster with the names currently in the ‘North three previously unrecognized lineages but they only American Lichen Checklist’ (Esslinger 2018). This sup- represent the fi rst step towards a comprehensive study ports the notion that the taxa of this clade present in of Dictyonema s.lat. in North America. We hope that North America do not correspond to the names applied by these results will alert workers to the unrecognized diver- Parmasto (1978) but likely represent regional endemics. sity in this clade. Given our prior global sampling, we As shown in the species discussion, remarks and fi gures anticipate that additional new species await discovery in herein, the three new taxa show distinctive morphologies. North America, particularly due to the high diversity of These diff erences may help reassign previously collected unique habitats in this area. This result would be con- material to the current taxonomy. However, there is a nota- sistent with our detailed studies of Cora in the Americas ble level of cryptic speciation in Dictyonema s.lat. which (Lücking et al. 2014, 2017b) and the Dictyonema clade requires a broader molecular dataset to properly assess in the Galapagos (Dal Forno et al. 2017), which demon- the phylogenetic diversity, distribution and morphology strated that most species in this group are endemic to of these basidiolichens in North America. small areas. 386 Plant and Fungal Systematics 64(2): 383–392, 2019

R04_Cyphellostereum_spec_Luecking_sn_CORI_EU825976 MDF388_Dictyonema_spec_Spielmann_11217_BRAZ MDF381_Dictyonema_spec_DalForno_2448_BRAZ MDF249_Dictyonema_spec_Boom_40318_MASC NSF097_Dictyonema_spec_Gostel_247_MADA MDF244_Dictyonema_spec_Boom_40730_MASC MDF245_Dictyonema_spec_Boom_40588_MASC DIC156_Dictyonema_phyllophilum_Lumbsch_19812_FIJI DIC156_Dictyonema_phyllophilum_Lumbsch_19812_FIJI_GXTHBOB01BTOR6 DIC125_Dictyonema_irpicinum_Lumbsch_19837e_FIJI_GXTHBOB01A9XWR DIC129_Dictyonema_spec_RivasPlata_2143_PHIL DIC129_Dictyonema_spec_RivasPlata_2143_PHIL_GXTHBOB01B0JV9 MDF283_Dictyonema_spec_Gumboski_4390_BRAZ MDF260_Dictyonema_spec_Beck_E415_ECUA_JCF67IH01B1HGN MDF459_Dictyonema_spec_DalForno_1920_ECUA DIC123_Dictyonema_metallicum_Luecking_26255_ECUA DIC123_Dictyonema_metallicum_Luecking_26255_ECUA_GXTHBOB01BN86H MDF370_Dictyonema_spec_DalForno_1770_CORI MDF488_Dictyonema_spec_DalForno_1983_ECUA MDF227_Dictyonema_spec_Luecking_34694_CORI_JCF67IH01DRH9R DIC501_Dictyonema_spec_Luecking_33529_COLO MDF476_Dictyonema_spec_DalForno_1974_ECUA MDF464_Dictyonema_spec_DalForno_1933a_ECUA MDF256_Dictyonema_spec_MercadoDiaz_sn_PUERI MDF255_Dictyonema_spec_MercadoDiaz_sn_PUERI_IXR852001DRKP3 MDF490_Dictyonema_spec_DalForno_1986_ECUA MDF485_Dictyonema_spec_DalForno_1981c_ECUA MDF071_Dictyonema_pectinatum_DalForno_1221_GALA MDF025_Dictyonema_pectinatum_DalForno_1170_GALA_JCF67IH01C25K3 MDF025_Dictyonema_pectinatum_DalForno_1170_GALA (collapsed) Dictyonema moorei DIC101_Dictyonema_spec_Vela_sn_PERU_GXTHBOB01B0I7Y DIC057_Dictyonema_spec_Luecking_16561_CORI DIC121_Dictyonema_spec_Luecking_25561_GUAT_GXTHBOB01A6E17 DIC116_Dictyonema_spec_Luecking_25551b_GUAT_GXTHBOB01BMUMT DIC116_Dictyonema_spec_Luecking_25551b_GUAT MDF434_Dictyonema_spec_DalForno_1826_ECUA (collapsed) Dictyonema barbatum NSF310_Dictyonema_sericeum_DalForno_3036_JAM DIC630_Dictyonema_spec_Moncada_6336_COLO MDF259_Dictyonema_spec_Beck_E400_ECUA DIC596_Dictyonema_spec_Spielmann_10246_BRAZ MDF263_Dictyonema_spec_Beck_E425c_ECUA MDF262_Dictyonema_spec_Beck_E425b_ECUA DIC100_Dictyonema_hapteriferum_Vela_sn_PERU_GXTHBOB01BWLNQ DIC138_Dictyonema_hapteriferum_Wilk_8868_BOLI MDF650_Dictyonema_spec_Chkovsky_sn_BRAZ MDF600_Dictyonema_spec_DalForno_600_BRAZ MDF284_Dictyonema_spec_Gerlach_791_BRAZ_IXR852001BQM9L MDF284_Dictyonema_spec_Gerlach_791_BRAZ_IXR852001BGI3T MDF261_Dictyonema_spec_Beck_E425a_ECUA (collapsed) Dictyonema discocarpum DIC618_Dictyonema_spec_Luecking_35425_COLO (collapsed) Dictyonema giganteum MDF491_Dictyonema_spec_DalForno_1987_ECUA MDF327_Dictyonema_spec_DalForno_1727_CORI MDF325_Dictyonema_spec_DalForno_1725_CORI MDF384_Dictyonema_spec_Gumboski_5016_BRAZ MDF383_Dictyonema_spec_Gumboski_5015_BRAZ MDF348_Dictyonema_spec_DalForno_1748_CORI DIC336_Dictyonema_spec_Amtoft_3095_CORI MDF081_Dictyonema_spec_Jonitz_592_PERU MDF363_Dictyonema_spec_DalForno_1763_CORI DIC404_Dictyonema_spec_Luecking_33854_PUERI MDF586_Dictyonema_spec_Aptroot_75495_TENE MDF246_Dictyonema_spec_Boom_45707_CANA_JCF67IH01C0IGW DIC566_Dictyonema_spec_Luecking_35284_COLO DIC523_Dictyonema_spec_Luecking_33365_COLO DIC565_Dictyonema_spec_Luecking_35282_COLO MDF333_Dictyonema_spec_DalForno_1733_CORI DIC055_Dictyonema_spec_Luecking_15340_CORI MDF230_Dictyonema_spec_Yanez_2500_ECUA DIC114_Dictyonema_spec_Luecking_30060_BRAZ_GXTHBOB01BGIWK DIC113_Dictyonema_spec_Luecking_30062_BRAZ MDF660_Dictyonema_spec_Spielmann_10131_BRAZ MDF103_Dictyonema_spec_DalForno_1070a_BRAZ MDF521_Dictyonema_spec_DalForno_2021_BRAZ MDF520_Dictyonema_spec_DalForno_2020_BRAZ DIC113_Dictyonema_spec_Luecking_30062_BRAZ_GXTHBOB01BLYJS MDF341_Dictyonema_spec_DalForno_1741_CORI MDF340_Dictyonema_spec_DalForno_1740_CORI DIC339_Dictyonema_spec_Luecking_18053_CORI MDF617_Dictyonema_spec_DalForno_2092b_BRAZ MDF619_Dictyonema_spec_DalForno_2096_BRAZ MDF673_Dictyonema_spec_Spielmann_8837_BRAZ MDF658_Dictyonema_spec_Spielmann_8843_BRAZ MDF241_Dictyonema_spec_CaceresAptroot_13598_BRAZ DIC053_Dictyonema_aeruginosulum_Nelsen_3754_CORI DIC331_Dictyonema_aeruginosulum_Trest_1569_CORI MDF351_Dictyonema_spec_DalForno_1741_CORI MDF616_Dictyonema_spec_DalForno_2085_BRAZ MDF463_Dictyonema_spec_DalForno_1928_ECUA MDF635_Dictyonema_spec_DalForno_2132_BRAZ MDF489_Dictyonema_spec_DalForno_1984_ECUA MDF484_Dictyonema_spec_DalForno_1981b_ECUA MDF626_Dictyonema_spec_DalForno_2115_BRAZ MDF468_Dictyonema_spec_DalForno_1936_ECUA MDF469_Dictyonema_spec_DalForno_1961_ECUA (collapsed) Dictyonema ramificans MDF258_Dictyonema_spec_Beck_E398_ECUA MDF233_Dictyonema_spec_Gumboski_3549_BRAZ MDF651_Dictyonema_spec_Koch_sn_BRAZ DIC338_Dictyonema_spec_Luecking_15353_CORI DIC362_Dictyonema_spec_Moncada_sn_COLO Figure 2. Phylogenetic tree of the genus Dictyonema inferred by maximumMDF324_Dictyonema likelihood_spec_DalForno_1724_CORI in RAxML using ITS fungal barcoding (n = 231, alignment 938 bp). Bolded branches have bootstrap values (BS) over 70% andMDF321_ are consideredDictyonema_spec_DalForno_1721_CORI well-supported. For original tree, see Figure S1. MDF228_Dictyonema_spec_Blanchon_5141_NEWZE MDF247_Dictyonema_spec_Boom_39933_MASC MDF248_Dictyonema_spec_Boom_40086_MASC_JCF67IH01AQQRU DIC134_Dictyonema_applanatum_Wilk_8982_BOLI_GXTHBOB01AKHJX MDF079_Dictyonema_spec_Jonitz_655_ECUA_JCF67IH01BOG67 MDF544_Dictyonema_spec_DalForno_2044_BRAZ MDF236_Dictyonema_spec_Gumboski_2343a_BRAZ MDF237_Dictyonema_spec_Gumboski_2343b_BRAZ DIC062_Dictyonema_hernandezii_Luecking_15243a_CORI R10_Dictyonema_hernandezii_Luecking_sn_CORI DIC122_Dictyonema_spec_Luecking_26258_ECUA DIC122_Dictyonema_spec_Luecking_26258_ECUA_GXTHBOB01BIQM4 DIC263_Dictyonema_spec_Luecking_31306_BRAZ NSF259_Dictyonema_spec_DalForno_2878_BRAZ MDF741_Dictyonema_spec_Coca_6958_COLO_LSU MDF577_Dictyonema_spec_Boom_44764_PANA MDF601_Dictyonema_spec_DalForno_2050_BRAZ MDF208_Dictyonema_lawreyi_Lawrey_1600_USA LK409a_Dictyonema_lawreyi_Kaminsky_409_USA LK409b_Dictyonema_lawreyi_Kaminsky_409_USA DIC059_Dictyonema_spec_Luecking_17200_CORI MDF337_Dictyonema_spec_DalForno_1737_CORI R02_Dictyonema_spec_Luecking_sn_CORI DIC337_Dictyonema_spec_Luecking_18008_CORI R01_Dictyonema_spec_Luecking_sn_CORI MDF239_Dictyonema_spec_Gumboski_2445_BRAZ MDF238_Dictyonema_spec_Gumboski_2399_BRAZ DIC410_Dictyonema_spec_Luecking_33915_PUERI MDF275_Dictyonema_huaorani_DavisYost_1051_ECUA DIC411_Dictyonema_spec_Luecking_33917_PUERI DIC407_Dictyonema_spec_Luecking_33907_PUERI DIC408_Dictyonema_spec_Luecking_33910_PUERI DIC412_Dictyonema_spec_Luecking_33920_PUERI DIC406_Dictyonema_spec_Luecking_33901_PUERI DIC413_Dictyonema_spec_Luecking_33936_PUERI MDF423_Dictyonema_spec_DalForno_1803_ECUA (collapsed) Dictyonema subobscuratum MDF242_Dictyonema_spec_CaceresAptroot_13632_BRAZ (collapsed) Dictyonema obscuratum MDF296_Dictyonema_spec_Aptroot_18078_BRAZ_IXR852001DMKPQ MDF571_Dictyonema_spec_DalForno_2221_PUERI_IXR852001DFWPA MDF296_Dictyonema_spec_Aptroot_18078_BRAZ DIC334_Dictyonema_spec_Luecking_15327_CORI DIC330_Dictyonema_spec_Luecking_17252i_CORI (collapsed) Dictyonema darwinianum

0.2 R04_Cyphellostereum_spec_Luecking_sn_CORI_EU825976 MDF388_Dictyonema_spec_Spielmann_11217_BRAZ MDF381_Dictyonema_spec_DalForno_2448_BRAZ MDF249_Dictyonema_spec_Boom_40318_MASC NSF097_Dictyonema_spec_Gostel_247_MADA MDF244_Dictyonema_spec_Boom_40730_MASC MDF245_Dictyonema_spec_Boom_40588_MASC DIC156_Dictyonema_phyllophilum_Lumbsch_19812_FIJI DIC156_Dictyonema_phyllophilum_Lumbsch_19812_FIJI_GXTHBOB01BTOR6 DIC125_Dictyonema_irpicinum_Lumbsch_19837e_FIJI_GXTHBOB01A9XWR DIC129_Dictyonema_spec_RivasPlata_2143_PHIL DIC129_Dictyonema_spec_RivasPlata_2143_PHIL_GXTHBOB01B0JV9 MDF283_Dictyonema_spec_Gumboski_4390_BRAZ MDF260_Dictyonema_spec_Beck_E415_ECUA_JCF67IH01B1HGN MDF459_Dictyonema_spec_DalForno_1920_ECUA DIC123_Dictyonema_metallicum_Luecking_26255_ECUA DIC123_Dictyonema_metallicum_Luecking_26255_ECUA_GXTHBOB01BN86H MDF370_Dictyonema_spec_DalForno_1770_CORI MDF488_Dictyonema_spec_DalForno_1983_ECUA MDF227_Dictyonema_spec_Luecking_34694_CORI_JCF67IH01DRH9R DIC501_Dictyonema_spec_Luecking_33529_COLO MDF476_Dictyonema_spec_DalForno_1974_ECUA MDF464_Dictyonema_spec_DalForno_1933a_ECUA MDF256_Dictyonema_spec_MercadoDiaz_sn_PUERI MDF255_Dictyonema_spec_MercadoDiaz_sn_PUERI_IXR852001DRKP3 MDF490_Dictyonema_spec_DalForno_1986_ECUA MDF485_Dictyonema_spec_DalForno_1981c_ECUA MDF071_Dictyonema_pectinatum_DalForno_1221_GALA MDF025_Dictyonema_pectinatum_DalForno_1170_GALA_JCF67IH01C25K3 MDF025_Dictyonema_pectinatum_DalForno_1170_GALA (collapsed) Dictyonema moorei DIC101_Dictyonema_spec_Vela_sn_PERU_GXTHBOB01B0I7Y DIC057_Dictyonema_spec_Luecking_16561_CORI DIC121_Dictyonema_spec_Luecking_25561_GUAT_GXTHBOB01A6E17 DIC116_Dictyonema_spec_Luecking_25551b_GUAT_GXTHBOB01BMUMT DIC116_Dictyonema_spec_Luecking_25551b_GUAT MDF434_Dictyonema_spec_DalForno_1826_ECUA (collapsed) Dictyonema barbatum NSF310_Dictyonema_sericeum_DalForno_3036_JAM DIC630_Dictyonema_spec_Moncada_6336_COLO MDF259_Dictyonema_spec_Beck_E400_ECUA DIC596_Dictyonema_spec_Spielmann_10246_BRAZ MDF263_Dictyonema_spec_Beck_E425c_ECUA MDF262_Dictyonema_spec_Beck_E425b_ECUA DIC100_Dictyonema_hapteriferum_Vela_sn_PERU_GXTHBOB01BWLNQ DIC138_Dictyonema_hapteriferum_Wilk_8868_BOLI MDF650_Dictyonema_spec_Chkovsky_sn_BRAZ MDF600_Dictyonema_spec_DalForno_600_BRAZ MDF284_Dictyonema_spec_Gerlach_791_BRAZ_IXR852001BQM9L MDF284_Dictyonema_spec_Gerlach_791_BRAZ_IXR852001BGI3T MDF261_Dictyonema_spec_Beck_E425a_ECUA (collapsed) Dictyonema discocarpum DIC618_Dictyonema_spec_Luecking_35425_COLO (collapsed) Dictyonema giganteum MDF491_Dictyonema_spec_DalForno_1987_ECUA MDF327_Dictyonema_spec_DalForno_1727_CORI MDF325_Dictyonema_spec_DalForno_1725_CORI MDF384_Dictyonema_spec_Gumboski_5016_BRAZ MDF383_Dictyonema_spec_Gumboski_5015_BRAZ MDF348_Dictyonema_spec_DalForno_1748_CORI DIC336_Dictyonema_spec_Amtoft_3095_CORI MDF081_Dictyonema_spec_Jonitz_592_PERU MDF363_Dictyonema_spec_DalForno_1763_CORI DIC404_Dictyonema_spec_Luecking_33854_PUERI MDF586_Dictyonema_spec_Aptroot_75495_TENE MDF246_Dictyonema_spec_Boom_45707_CANA_JCF67IH01C0IGW DIC566_Dictyonema_spec_Luecking_35284_COLO DIC523_Dictyonema_spec_Luecking_33365_COLO DIC565_Dictyonema_spec_Luecking_35282_COLO MDF333_Dictyonema_spec_DalForno_1733_CORI DIC055_Dictyonema_spec_Luecking_15340_CORI MDF230_Dictyonema_spec_Yanez_2500_ECUA DIC114_Dictyonema_spec_Luecking_30060_BRAZ_GXTHBOB01BGIWK DIC113_Dictyonema_spec_Luecking_30062_BRAZ MDF660_Dictyonema_spec_Spielmann_10131_BRAZ MDF103_Dictyonema_spec_DalForno_1070a_BRAZ MDF521_Dictyonema_spec_DalForno_2021_BRAZ MDF520_Dictyonema_spec_DalForno_2020_BRAZ DIC113_Dictyonema_spec_Luecking_30062_BRAZ_GXTHBOB01BLYJS MDF341_Dictyonema_spec_DalForno_1741_CORI MDF340_Dictyonema_spec_DalForno_1740_CORI DIC339_Dictyonema_spec_Luecking_18053_CORI MDF617_Dictyonema_spec_DalForno_2092b_BRAZ MDF619_Dictyonema_spec_DalForno_2096_BRAZ MDF673_Dictyonema_spec_Spielmann_8837_BRAZ MDF658_Dictyonema_spec_Spielmann_8843_BRAZ MDF241_Dictyonema_spec_CaceresAptroot_13598_BRAZ DIC053_Dictyonema_aeruginosulum_Nelsen_3754_CORI DIC331_Dictyonema_aeruginosulum_Trest_1569_CORI MDF351_Dictyonema_spec_DalForno_1741_CORI MDF616_Dictyonema_spec_DalForno_2085_BRAZ MDF463_Dictyonema_spec_DalForno_1928_ECUA MDF635_Dictyonema_spec_DalForno_2132_BRAZ MDF489_Dictyonema_spec_DalForno_1984_ECUA M. Dal Forno et al.: A fi rst phylogenetic assessment of DictyonemaMDF484_ s.lat.Dictyonema in southeastern_spec_DalForno_1981b_ECUA North America 387 MDF626_Dictyonema_spec_DalForno_2115_BRAZ MDF468_Dictyonema_spec_DalForno_1936_ECUA MDF469_Dictyonema_spec_DalForno_1961_ECUA (collapsed) Dictyonema ramificans MDF258_Dictyonema_spec_Beck_E398_ECUA MDF233_Dictyonema_spec_Gumboski_3549_BRAZ MDF651_Dictyonema_spec_Koch_sn_BRAZ DIC338_Dictyonema_spec_Luecking_15353_CORI DIC362_Dictyonema_spec_Moncada_sn_COLO MDF324_Dictyonema_spec_DalForno_1724_CORI MDF321_Dictyonema_spec_DalForno_1721_CORI MDF228_Dictyonema_spec_Blanchon_5141_NEWZE MDF247_Dictyonema_spec_Boom_39933_MASC MDF248_Dictyonema_spec_Boom_40086_MASC_JCF67IH01AQQRU DIC134_Dictyonema_applanatum_Wilk_8982_BOLI_GXTHBOB01AKHJX MDF079_Dictyonema_spec_Jonitz_655_ECUA_JCF67IH01BOG67 MDF544_Dictyonema_spec_DalForno_2044_BRAZ MDF236_Dictyonema_spec_Gumboski_2343a_BRAZ MDF237_Dictyonema_spec_Gumboski_2343b_BRAZ DIC062_Dictyonema_hernandezii_Luecking_15243a_CORI R10_Dictyonema_hernandezii_Luecking_sn_CORI DIC122_Dictyonema_spec_Luecking_26258_ECUA DIC122_Dictyonema_spec_Luecking_26258_ECUA_GXTHBOB01BIQM4 DIC263_Dictyonema_spec_Luecking_31306_BRAZ NSF259_Dictyonema_spec_DalForno_2878_BRAZ MDF741_Dictyonema_spec_Coca_6958_COLO_LSU MDF577_Dictyonema_spec_Boom_44764_PANA MDF601_Dictyonema_spec_DalForno_2050_BRAZ MDF208_Dictyonema_lawreyi_Lawrey_1600_USA LK409a_Dictyonema_lawreyi_Kaminsky_409_USA LK409b_Dictyonema_lawreyi_Kaminsky_409_USA DIC059_Dictyonema_spec_Luecking_17200_CORI MDF337_Dictyonema_spec_DalForno_1737_CORI R02_Dictyonema_spec_Luecking_sn_CORI DIC337_Dictyonema_spec_Luecking_18008_CORI R01_Dictyonema_spec_Luecking_sn_CORI MDF239_Dictyonema_spec_Gumboski_2445_BRAZ MDF238_Dictyonema_spec_Gumboski_2399_BRAZ DIC410_Dictyonema_spec_Luecking_33915_PUERI MDF275_Dictyonema_huaorani_DavisYost_1051_ECUA DIC411_Dictyonema_spec_Luecking_33917_PUERI DIC407_Dictyonema_spec_Luecking_33907_PUERI DIC408_Dictyonema_spec_Luecking_33910_PUERI DIC412_Dictyonema_spec_Luecking_33920_PUERI DIC406_Dictyonema_spec_Luecking_33901_PUERI DIC413_Dictyonema_spec_Luecking_33936_PUERI MDF423_Dictyonema_spec_DalForno_1803_ECUA (collapsed) Dictyonema subobscuratum MDF242_Dictyonema_spec_CaceresAptroot_13632_BRAZ (collapsed) Dictyonema obscuratum MDF296_Dictyonema_spec_Aptroot_18078_BRAZ_IXR852001DMKPQ MDF571_Dictyonema_spec_DalForno_2221_PUERI_IXR852001DFWPA MDF296_Dictyonema_spec_Aptroot_18078_BRAZ DIC334_Dictyonema_spec_Luecking_15327_CORI DIC330_Dictyonema_spec_Luecking_17252i_CORI (collapsed) Dictyonema darwinianum

0.2 Figure 2. Continued.

Taxonomy to bright yellow, 5–7 μm wide × 5–7 μm high, mostly circular; clamp connections observed in loose hyphae Cyphellostereum georgianum Dal Forno, McMullin forming the thallus and in hymenophore. Hymenophore & Lücking, sp. nov. (Fig. 3) forming irregular patches, resupinate to corticioid; patches MycoBank MB 832378 up to 14 mm long and 2 mm wide, white; hymenophore in section up to 100 μm thick, composed of 2–4 μm thick Diagnosis: Characterized by the opaque, crustose, dark green hyphae. Basidioles mostly rounded to clavate, 3–4 μm thallus formed by dense fi brils embedded in an inconspicuous matrix, with abundant corticioid white basidiocarps, and a fungal wide × 12–18 μm high. Basidia and basidiospores not sheath consisting of irregular, thin, branched hyphae surrounding observed. the cells, partly furnished with clamps. Etymology. The epithet refers to the state where the type Type: United States, Georgia, McIntosh County, Harper Lake specimens were collected. Park, 31°28′18.9″N (31.47192), 81°36′22.8″W (−81.60632), alt. 25 m (78 ft), on trunk of deciduous tree; 29 March 2013, R. T. Distribution and ecology. Currently only known from McMullin 12290 (CANL – holotype!; US – isotype!). the type locality, where it grows on trunks of trees such as GenBank ITS barcoding sequences: KY861606 (holotype), Quercus. The locality where it was collected has canopy KY861529 (paratype) dominated by Pinus taeda, with a midstory of Quercus hemisphaerica, Q. geminata, Ilex opaca, Magnolia gran- Description. Thallus epiphytic on tree trunks, crus- diﬂ ora and Carya sp., and shrub vegetation with Serrenoa tose-fi lamentous growth form, developing a continuous repens, Lyonia ferruginea, L. lucida (near wetlands), Vac- mat at ‘base’, with small irregular elevated thick ‘patches’ cinium arboreum, V. tenellum, V. stamineum, Hamamelis formed by denser fi brils. Upper surface with densely fun- virginiana and Ditrysinia fruticosa. gal-cyanobacterial fi brils, dark opaque green, with yellowish hue in elevated ‘patches’ but with bluish ‘base’. Remarks. Although possibly originating from a com- Thallus composed entirely by cyanobacterial fi laments mon ancestor, this Georgian species diff ers from its sister wrapped in fungal sheath; individual cells 7–12.5 μm species in South Carolina and Puerto Rico by its dense wide × 5–10 μm high, green to green-bluish, covered by crustose habit, forming a continuous dark green mat yet thin elongated fungal cells; heterocytes frequent, pale with clearly elevated thick patches of dense fi brils, while 388 Plant and Fungal Systematics 64(2): 383–392, 2019

Figure 3. Cyphellostereum georgianum. Crustose thallus growing directly on bark and overgrowing bryophytes, with white steroid hymenophores. A–C – general view of thallus observed under dissecting scope. Note reproductive structures, i.e., hymenophores, in A as resupinate white patches; D–F – general view of thallus observed under light microscope. In E, note lichenized fibrils (cyanobacterial cells wrapped in fungal hyphae) and free-living green algae surrounding them. Note clamp connections in F (arrows). Images from specimen collected by T. McMullin 12290 (A–C, E) and from specimen collected by R. Rosentreter 17755 (D, F). Scales: A, B = 2 mm; C = 4 mm; D, F = 30 μm; E = 40 μm.

the sister species has a thinner and more bluish thallus 81°36′22.8″W (−81.60632), alt. 25 m (78 ft), on bark of Quer- with more defined (separated) fibrils. Close-up obser- cus; 29 March 2013, R. Rosentreter 17755 (SRP – paratype!). vations under a dissecting scope show short yellowish fibrils. The thallus lacks organized layers but parts of it Cyphellostereum jamesianum Dal Forno & Kaminsky, are dominated by a gelatinous matrix formed by pho- sp. nov. (Fig. 4) tosynthesizing organisms (possibly cyanobacteria and MycoBank MB 832354 green algae). Diagnosis: Characterized by the crustose, dark green thallus Additional specimens examined. United States, Georgia, formed by loose individual fibrils, and a fungal sheath consisting McIntosh County, Harper Lake Park, 31°28′18.9″N (31.47192), of irregular, thin, branched hyphae surrounding the cells. M. Dal Forno et al.: A first phylogenetic assessment of Dictyonema s.lat. in southeastern North America 389

Type: UNITED STATES, South Carolina, Berkeley County, fungal-cyanobacterial fibrils, dark opaque green to Francis Marion National Forest along Forest Service Road 204F, green-bluish. Thallus composed entirely of cyanobac- 0.25 miles S of McConnel’s Landing, 1.3 miles N of jct w/ terial filaments wrapped in fungal sheath; filament cells Forest Service Road 204, 33°14′29″N, 79°31′16″W, alt. ~ 12 m, 11–13(–14) μm wide × 4–6(–8) μm high, green-bluish, on trunk of 4″ diam. oak; 3 December 2013, R. C. Harris 59781 covered by thin elongated fungal cells; heterocytes fre- (NYBG – holotype!). quent, pale to bright yellow; clamp connections observed GenBank ITS barcoding sequences: KY861586 (holotype), (in fresh material from Puerto Rico). Hymenophore (only MN046975 (FL paratype), KY861587 (PR paratype). one present) in development forming a round patch, resu- pinated, 0.5 mm long × 0.5 mm wide, white, smooth, flat. Description. Thallus epiphytic on tree trunks, associ- ated with bryophytes, crustose-filamentous growth form, Etymology. This species honors the work of our esteemed developing a mat of ± individual to slightly interwoven colleague, mentor, and friend, Dr. James D. Lawrey, who

Figure 4. Cyphellostereum jamesianum. All images from A. Aptroot 72323. A–C – general view of thallus observed under dissecting scope; E–F – general view of thallus observed under light microscope. Scales: A = 200 μm; B = 400 μm; C–E = 100 μm; F = 50 μm. 390 Plant and Fungal Systematics 64(2): 383–392, 2019

contributed to the discovery that the genus Cyphelloste- Remarks. This species is sister to C. georgianum from reum belongs in the Dictyonema clade (Lawrey et al. Georgia, and both share the crustose-filamentous growth 2009). form. However, under a dissecting scope the species are distinct: C. georgianum has a thallus organized in dense Distribution and ecology. This species is known from mats with elevated thick patches of tangled fibrils (Fig. 3), three localities: two in the southeastern United States and while C. jamesianum forms loose, individual to slightly one in Puerto Rico. The type (from South Carolina) was interwoven fungal-cyanobacterial fibrils (Fig. 4). collected in a mature open pine-oak woodland; the Flor- ida specimen is known from old-growth oak-dominated Additional specimens examined. UNITED STATES, deciduous forest that floods seasonally, and the Puerto Florida, Sarasota County, Myakka River State Park, off the hiking Rican specimen from a disturbed forest. trail north of the canopy walkway trail; 27°14′39″N, –082°18′W,

Figure 5. Dictyonema lawreyi. All images from J.D. Lawrey 1600 (holotype). Crustose thallus growing directly on bark. A–C – general view of thallus observed under dissecting scope. Note white distinct prothallus and hypothallus; D–F – general view of thallus observed under light microscope. Note jigsaw-puzzle-shaped fungal sheath hyphae. Scales: A, C = 200 μm; B – 500 μm; D, F = 50 μm; E = 25 μm. M. Dal Forno et al.: A first phylogenetic assessment of Dictyonema s.lat. in southeastern North America 391

14 m; old-growth oak-dominated seasonally flooded hardwood the fibrils are very distinct: the Brazilian species forms forest, on oak; 16 January 2018, L. Kaminsky 370 (FLAS, dense mats of packed interwoven fibrils, while the Costa US – paratypes!). PUERTO RICO, Utuado, Río Abajo State Rican species form small tufts of interwoven erect fibrils. Forest, ~ 65 km WSW of San Juán and 15 km S of Arecibo, Under a microscope these morphologies differ greatly from access road in forest beyond camp ground; 18°19′N, 66°43′W, D. lawreyi, which has sparse, loosely interwoven, bright 380 m; lowland evergreen and seasonal evergreen wet forest on karst, disturbed primary and old-growth secondary forest, turquoise fibrils resting on a white hypothallus. on tree bark; 20 February 2014, A. Aptroot 72323 (US, ABL, Additional specimens examined. UNITED STATES, Florida, F, UPR – paratypes!). Putnam County, Ordway-Swisher Biological Station, Southeast of Lake Suggs. Habitat: deciduous hardwood forest. At edge of Dictyonema lawreyi Dal Forno, Kaminsky & Lücking, deciduous hardwood forest and longleaf pine habitat boundary. sp. nov. (Fig. 5) 29.684450°, –82.013689°, 29 m; on base of Quercus, 15 Decem- MycoBank MB 832355 ber 2016, L. Kaminsky 409 (FLAS, US – paratypes!). Note: One of the sequences for this specimen is short (159bp) and Diagnosis: A turquoise appressed-filamentousDictyonema with therefore not submitted to GenBank. However, this sequence a well-developed, arachnoid, white prothallus and loosely ar- is available in the alignment of File S2. ranged and prostrate turquoise filaments resting on a white hypothallus. Acknowledgments Type: United States, Florida, Marion County, Ocala National Forest, 2.2 miles N of Florida Highway 40 on Florida Highway We would like to thank the National Science Foundation for 19, Mormon Branch on E side of road, 29°12′N, 81°39′W, on support through a Postdoctoral Research Fellowship in Biology Magnolia, 2 January 1996; J. D. Lawrey 1600 (US – holotype!; (PRFB 1609022; PI M. Dal Forno) and the grant ‘Phylogenetic GMUF – isotype!). diversity of mycobionts and photobionts in the cyanolichen GenBank ITS barcoding sequences: KY861507 (holotype), genus Dictyonema, with emphasis on the Neotropics and the MN046973 (paratype). Galapagos Islands’ (DEB 0841405; PI J. Lawrey; Co-PIs R. Lücking, P. Gillevet). The South Carolina material was col- Description. Thallus epiphytic on tree trunks, with crus- lected with support from the National Science Foundation (DEB tose-filamentous growth form, white hypothallus extend- 1145511; PI J. Lendemer; Co-PI R. Harris). We also thank Dr. James Lendemer for organizing the Tuckerman Workshop in ing to form distinct prothallus with continuous, fine mat of Georgia in 2013, where samples of Cyphellostereum georgianum turquoise fibrils on top. Upper surface with very loosely were collected, and Dr. Richard C. Harris for collecting the interwoven and irregular fungal-cyanobacterial fibrils. type material of C. jamesianum. The Ordway-Swisher Biolog- Patches up to 5 cm long, ± continuous. Thallus composed ical Station is thanked for funding (Matthew Smith JumpStart mainly of cyanobacterial filaments wrapped in fungal award FLA-PLP-005561), and the Florida Department of En- sheath; filament cells 12–17.5 μm wide × 3–8 μm high, vironmental Protection for granting permission to collect study green-bluish, covered by jigsaw-puzzle-shaped fungal material (Permit # 11181610). cells; heterocytes rare, mostly hyaline to sparsely pale yellow, 8–10 μm wide × 4–5 μm high; clamp connections Supplementary electronic material not observed. Immature hymenophore possibly present, Figure S1. Phylogenetic tree of the genus Dictyonema inferred by forming resupinate, irregular, small white patches, but maximum likelihood in RAxML using ITS fungal barcoding (n = 231, hymenium was observed. alignment 938 bp). Tree visualized in FigTree v1.4.3 and edited in Photoshop Illustrator. Download file Etymology. This species is also dedicated to Dr. James D. Table S1. List of specimens included in this study. Download file Lawrey, whose pioneering work helped to redefine the cir- File S1. MAFFT alignment generated on Geneious used to generate cumscription of the genus Dictyonema s.str. (Lawrey et al. the phylogenetic tree of the genus Cyphellostereum (n = 57, alignment 2009) and who is the collector of the holotype specimen. 864 bp). Download file File S2. MAFFT alignment generated on Geneious used to generate Distribution and ecology. This species is known from the phylogenetic tree of the genus Dictyonema (n = 231, alignment three specimens, all epiphytic in hardwood forests in 938 bp). Download file Florida. The specimen from Ordway-Swisher was collected where an oak-dominated deciduous forest begins References to intergrade with pine. Dal Forno, M. 2015. Evolution and Diversity of the basidiolichen Remarks. This species is a typical-looking appressed-fil- clade Dictyonema (Agaricales: Hygrophoraceae). PhD Disserta- amentous Dictyonema, but the color more resembles spe- tion. 308 pp. cies belonging to the Cyphellostereum imperfectum group. Dal Forno, M., Lawrey, J. D., Sikaroodi, M., Bhattarai, S., Gillevet, However, anatomically it is clearly a Dictyonema due to P. M., Sulzbacher, M. & Lücking, R. 2013. Starting from scratch: the presence of jigsaw-puzzle-shaped fungal cells wrap- evolution of the lichen thallus in the basidiolichen Dictyonema (Agaricales: Hygrophoraceae). Fungal Biology 117: 584–598. ping around the cyanobacterial cells. The species from Bra- zil, which appears in Figure 2 as the sister species (without Dal Forno, M., Bungartz, F., Yánez-Ayabaca, A., Lücking, R. & Lawrey, J. D. 2017. High levels of endemism among Galapagos basidioli- support), and the species from Costa Rica that together chens. Fungal Diversity 85: 45–73. form a well-supported clade with Dictyonema lawreyi, are Esslinger, T. L. 2018. A Cumulative Checklist for the Lichen-Forming, also crustose-filamentous with a clear white prothallus. Lichenicolous and Allied Fungi of the Continental United States and Nonetheless, both (unnamed) species are dark green and Canada, Version 22. Opuscula Philolichenum 17: 6–268. 392 Plant and Fungal Systematics 64(2): 383–392, 2019

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