f u n g a l e c o l o g y 1 9 ( 2 0 1 6 ) 1 5 5 e1 6 8 available at www.sciencedirect.com ScienceDirect journal homepage: www.elsevier.com/locate/funeco Relationships between terrestrial and freshwater lignicolous fungi R. KODSUEBa,b,*, S. LUMYONGb, E.H.C. MCKENZIEc, A.H. BAHKALId, K.D. HYDEe a Faculty of Science and Technology, Pibulsongkram Rajabhat University, Phitsanulok 65000, Thailand Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand c Landcare Research, Private Bag 92170, Auckland, New Zealand d Botany and Microbiology Department, King Saud University, College of Science, P.O. Box: 2455, Riyadh 1145, Saudi Arabia e Center of Excellence in Fungal Research, Mae Fah Luang University, Tasud, Chiang Rai 57100, Thailand b article info abstract Article history: This study investigated the communities of fungi on Magnolia liliifera wood in terrestrial Received 16 May 2015 and freshwater habitats in northern Thailand. Freshly fallen samples of M. liliifera wood Revision received 26 August 2015 were collected from the forest floor, oven dried, and then placed in the stream or adjacent Accepted 27 August 2015 to the stream on the forest floor as baits. The samples were collected and examined after 3 Available online 21 October 2015 and 6 months of submersion and periodically over 29 months from the forest floor. Thirty- Corresponding editor: € rlocher Felix Ba seven species of lignicolous fungi were discovered from 50 bait samples of wood sub- Keywords: baits). The fungal communities on wood in freshwater were distinct from those in the Aquatic fungi terrestrial habitat. Seventeen species of fungi overlapped between freshwater and terres- Fungal succession trial habitats, but only five out of 234 species overlapped between all habitats (freshwater, Lignin natural terrestrial and terrestrial bait). Corynespora cassiicola (60% frequency of occurrence) merged in the stream. Fungi obtained from submerged baits were compared with those found on wood of M. liliifera in terrestrial habitats (both naturally terrestrial and terrestrial Degradation was the most common taxon found on natural terrestrial samples, while Lasiodiplodia Fungal communities theobromae (43% frequency of occurrence) was the dominant species from terrestrial baits. Candelabrum brocchiatum (26%) was the most common species from submerged baits. The common genera of fungi obtained from submerged baits were similar to those reported in other submerged wood studies. ª 2015 Elsevier Ltd and The British Mycological Society. All rights reserved. Introduction Freshwater fungi are any species relying on freshwater for some part of their life cycle, or any species colonizing substrata that are predominantly aquatic or semi-aquatic in nature (Thomas, 1996; Wong et al. 1998). In other words, they are fungi whose habitats may be clearly of an aquatic nature, or that colonize submerged plant parts in freshwater environments (Wong et al. 1998). Freshwater fungi thus are a ubiquitous and diverse group of organisms that colonize substrata found in aquatic or semi-aquatic environments (Luo et al., 2004; Fryar et al., 2005; Pascoal et al., 2005; Sakayaroj * Corresponding author. Faculty of Science and Technology, Pibulsongkram Rajabhat University, Phitsanulok 65000, Thailand. E-mail address: kodsueb@yahoo.com (R. Kodsueb). http://dx.doi.org/10.1016/j.funeco.2015.09.005 1754-5048/ª 2015 Elsevier Ltd and The British Mycological Society. All rights reserved. 156 R. Kodsueb et al. et al., 2005; Vijaykrishna and Hyde, 2006; Shearer et al., 2007; Hyde et al. 2016). They are taxonomically diverse, and flourish € nczo € l and in various ecological niches (Cai et al., 2003; Go vay, 2003; 2004; Fryar et al., 2004a; b; Vijaykrishna and Hyde, Re 2006; Shearer et al., 2007). According to the above definition, some terrestrial fungi, which release spores that are dispersed in water, are regarded as freshwater fungi (Luo et al., 2004). Shearer (1993), however, mentioned that the presence of spores in water alone may not be an appropriate definition. Moreover, there are several species of fungi (e.g. Jahnula species) that only occur in freshwater habitats and only those can be confidently classified as freshwater fungi (Wong et al., 1998). Some ascomycetes categorized as freshwater fungi have also been reported from other habitats e.g. terrestrial and marine habitats. Freshwater ascomycetes can be divided into four groups based on their occurrence: (1) genera that are known only from freshwater habitats; (2) genera found in both freshwater and terrestrial habitats; (3) genera from freshwater and marine habitats; and (4) genera that are found in freshwater, marine and terrestrial habitats (Table 1, Vijaykrishna and Hyde, 2006). Vijaykrishna and Hyde (2006) investigated the origin of freshwater ascomycetes using molecular sequence data, and showed that freshwater ascomycetes (and marine fungi) originated from terrestrial fungi. The freshwater fungi that grow on submerged wood are known as lignicolous freshwater fungi (Hyde et al., 2016), while those that grow on leaves are known as aquatic leafinhabiting hyphomycetes or Ingoldian fungi (Graça et al., 2016). These fungi are generally studied by different methods (Chan et al., 2000; Tsui et al., 2000), with woody lignicolous samples being placed in moist chambers and leaves being placed in aerated bubbles chambers (Graça et al., 2016). The communities on wood and leaves therefore differ, although the few studies where woody litter has been placed in bubble € nczo € l and Re vay, chambers have revealed Ingoldian fungi (Go 1993; Sridhar et al., 2010). Here we use moist chamber incubation so our study is confined to lignicolous freshwater fungi, although we briefly discuss leaf-inhabiting fungi. Conidial fungi with staurospores, the Ingoldian fungi, are commonly reported from freshwater habitats (Descals and € nczo € l and Re  vay, 2006). Ando (1992) introMoralejo, 2001, Go duced the term “terrestrial aquatic hyphomycetes” for the fungi which possess staurospores found in terrestrial habitats (e.g. litter, roots and soils). Some species in this group are listed in Table 2. Relationships between freshwater and terrestrial lignicolous fungi There is little information concerning the relationship between freshwater and terrestrial lignicolous fungi. Cai et al. (2006b) studied the fungal communities on bamboo in a freshwater stream and in the riparian region, and found that different fungal communities occur in each habitat, with low overlap of species, and differing dominant species. In Thailand, the relationship between freshwater fungi and terrestrial fungi on palms has been investigated by comparing the occurrence of fungi on the same substratum in different habitats (e.g. Pinnoi et al., 2006; Pinruan et al., 2007; Boonyuen et al., 2014). Pinnoi et al. (2006) studied fungal diversity on different parts of the palm, Eleidoxa conferta, including those from terrestrial (dry and damp materials) and submerged habitats. Boonyuen et al. (2014) showed partial overlap in species between the two habitats with submerged samples yielding the most fungal records. Similar work has been reported by Pinruan et al. (2007), who studied the fungal diversity on another palm, Licuala longicalycata. However, they found that the dry material supported most fungal diversity. Boonyuen et al. (2014) showed that wood-inhabiting fungi are diverse, however, they differ between studies depending on tree species, geography and the exposure period. They mentioned that freshwater fungi are slow to cause wood decay since they are soft rotters, as compared to white rotters and brown rotters that predominate in wood decay in terrestrial environments (Yuen et al., 2000). Woody litter may fall into streams and rivers. It is not clear whether the fungi decaying terrestrial forest wood continue to thrive when submerged in water. In this study, we investigated fungi on Magnolia liliifera wood from freshwater and terrestrial habitats to see if they are the same or different. This study examined whether fungi on decaying wood on the forest floor are also present on wood submerged in water. Materials and methods Study sites This study was undertaken in an evergreen forest nearby the Medicinal Plant Garden in Doi Suthep-Pui National Park, Table 1 e Common freshwater ascomycete genera with terrestrial, marine/terrestrial or no counterparts (from Vijaykrishna and Hyde, 2006). Freshwater only Freshwater/Terrestrial Freshwater/Marine Freshwater/Marine/Terrestrial Aquaticola Cataractispora Jahnula Mamillisphaeria Pseudoproboscispora Rivulicola Torrentispora Annulatascus Ascotaiwania Byssosphaeria Cercophora Kirschsteiniothelia Ophioceras Pseudohalonectria Aniptodera Halosarpheia Nais Quintaria Savoryella Anthostomella Didymella Lophiostoma Massarina Phomatospora Saccardoella Vibrissea Terrestrial and freshwater lignicolous fungi 157 Table 2 e Staurosporous aquatic hyphomycetes reported from fallen leaves, fallen twigs and litter (modified from Ando, 1992). Species Alatosessilispora bibrachiata Alatospora acuminata Alatospora pulchella Anguillospora crassa Anguillospora longissima Anguillospora rosea Arborispora dolichovirga Arborispora palma Articulospora foliicola Articulospora tetracladia Calcarispora hiemalis Ceratosporium cornutum Ceratosporium verrucosum Clavariopsis aquatica Clavatospora longibranchiaga Colispora elongata Crucella subtilis Culicidospora aquatica Culicidospora gravida Curicispora ombrogena Dacryoscyphus chrysochilus Dendrospora erecta Dendrospora fastuosa Dendrospora fusca Dendrospora nana Dendrospora tenella Dicranidion fissile Digitodochium rhodoleucum Diplocladiella alta Dwayaangam colodena Dwayaangam cornuta Dwayaangam yakuensis Flagellospora curvula Flagellospora fusarioides Fontanospora fusiramosa Gemmulina botryosa Geniculosora inflata Gyoerffyella biappendiculata Gyoerffyella gemellipara Gyoerffyella rotula Heliscella stellata Heliscina campanulata Heliscus lugdunensis Ingoldiella hamata Isthmotricladia britannica Isthmotricladia rotundata Lateriramulosa ainflata Lateriramulosa minitriangularia Lemoniera aquatica Lemoniera filiformis Lemoniera terrestris Lunulospora curvula Margaritispora aquatica Microstella pluvioriens Mycocentrospora acerina Ordus tribrachiatus Pleuropedium multiseptatum Pleuropedium tricladioides Porocladium aquaticum Retiarius bovicornutus Speiropsis pedatospora References Ando (1992) € nzo € l and Re vay (1983), Sokolski et al. (2006) Go Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Ando (1992) Matsushima (1975) € nczo € l and Re vay (1983), Sokolski et al. (2006) Go Sokolski et al. (2006) Ando (1992) Kirschner and Chen (2004) € nczo € l and Re vay (1983), Sokolski et al. (2006) Go Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Ando (1992) Kirschner and Yang (2005) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Ando (1992) Tubaki and Kubono (1989) Kirschner and Chen (2004) Sokolski et al. (2006) Sokolski et al. (2006) Ando (1992) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Kodsueb (2007) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Kodsueb (2007) Sokolski et al. (2006) Matsushima (1983) Matsushima (1975) Sokolski et al. (2006) € nczo € l and Re vay (1983), Sokolski et al. (2006) Tubaki (1958), Go Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Ando (1992) Sokolski et al. (2006) Ando (1992) Sokolski et al. (2006) Sokolski et al. (2006) Sokolski et al. (2006) Ando (1992) Kodsueb (2007) (continued on next page) 158 R. Kodsueb et al. Table 2 e (continued ) Species Speiropsis rogergoosensis Taeniospora descalsii Taeniospora gracilis Tetrachaetum elegans Tetracladium furcatum Tetracladium setigerum Tetraploa aristata Titaea clakeae Titaeella capnophila Tricellula taiwanensis Tricladiella pluvialis Tricladium angulatum Tricladium splendens Tridentaria sp. Trifurcospora irregularis Trinacrium subtile Tripospermum acerinum Tripospermum infalcata Tripospermum myrti Triscelophorus monosporus Trisulcosporium acerinum Varicosporium elodeae Varicosporium giganteum Volucrispora ornithomorpha Chiang Mai Province, northern Thailand. The sampling site for the submerged succession experiment was located in the same area as for the terrestrial experiment. The stream was 30 cm to 1 m wide, 15 cm to 45 cm deep and passed through secondary forest. Unpolluted water flowed throughout the year, with flooding during the rainy season. The stream comprised a stony, sandy and gravel bottom. Sample collection and examination One hundred and ten dead branches of Magnolia liliifera (ca 30 cm long, 1e2.5 cm diameter) that had fallen to the ground under M. liliifera trees were collected and oven-dried at 70  C for 1 week and used for the submersion and terrestrial succession study. The submerged wood bait samples were tied separately with nylon thread and submerged in the study stream at Doi Suthep-Pui on 18 November 2002. Twenty-five wood baits were randomly collected from the stream at each sampling time along the stream of the study site. It was planned to collect samples at 3, 6, 9 and 12 months. However, in the rainy season, most of the baits were washed away by very strong flooding, while other baits were buried in silt and sand. Therefore, baits were collected only twice e 17 February 2003 (month 3; 25 samples) and 20 May 2003 (month 6; 25 samples). Sixty samples (ca 30 cm long, 1e2.5 cm diameter) of naturally decaying M. liliifera, that had fallen to the ground under the trees, were also collected from areas alongside the stream for general fungal diversity assessment. For terrestrial succession studies, two-hundred living samples (ca 30 cm long, 1e2.5 cm diameter, with bark) were cut and collected from living trees. Samples were oven-dried at 70  C for 1 week and tied References Keshava Prasad and Bhat (2002) Sokolski et al. (2006) Sokolski et al. (2006) € nczo € l and Re vay (1983), Sokolski Go Sokolski et al. (2006) € nczo € l and Re vay (1983), Sokolski Go Ando (1992) Ando (1992) Ando (1992) Sokolski et al. (2006) Ando (1992) € nczo € l and Re vay (1983), Sokolski Go Sokolski et al. (2006) Ando (1992) Ando (1992) Ando (1992) Ando (1992) Ando (1992) Sokolski et al. (2006) Sokolski et al. (2006) Ando (1992) € nczo € l and Re vay (1983), Sokolski Go Sokolski et al. (2006) € nczo € l and Re vay (1983) Go et al. (2006) et al. (2006) et al. (2006) et al. (2006) with labelled plastic tags. Dried samples (terrestrial baits) were then placed on the ground under M. liliifera trees on 9 July 2002. Ten baits were randomly collected from the field at each sampling time (15d, 1, 3, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 29 months, Kodsueb et al. 2008a, b). Fungi on these samples for each experiment were examined and identified following 1 week of incubation and periodically examined for up to 1 month. The fungi were identified and recorded. Herbarium material is maintained at CMU (Chiang Mai University). Fungi were identified by morphological characters using relevant texts and references (e.g. Ellis, 1971, 1976; Carmichael et al., 1980; Sutton, 1980; Sivanesan, 1984; € hlich and Hyde, 2000; Hyde et al., 2000; Lu and Hyde, Fro 2000; Grgurinovic, 2003; Taylor and Hyde, 2003; Tsui and Hyde, 2003; Wang et al., 2004; Wu and Zhuang, 2005; Cai et al., 2006a). Statistical analyses A 3-dimensional correspondence analysis (Anonymous, 1995) was performed to examine the differences in fungal communities on M. liliifera in freshwater and terrestrial habitats. The results are presented in terms of percentage occurrence of fungi (Table 3), where taxa with a percentage occurrence higher than ten are regarded as dominant species. Shannon indices (H’) were used to express species diversity of a community (Shannon and Weaver, 1949), and species area curves were used to determine the adequacy of the sample size (10 samples as the least number). Similarity of fungal communities (Magurran, 2004; Diserud and Ødegaard, 2007) was calculated from PC-ORD version 4.0 (McCune and Mefford, 1999). Terrestrial and freshwater lignicolous fungi 159 Table 3 e Percentage occurrence of fungi found on submerged Magnolia liliifera wood baits. Taxa Candelabrum brocchiatum Berkleasmium minutissimum Ellisembia opaca Aquaticola hyalomura Phaeoisaria clematidis Sporoschisma uniseptatum Acanthophyses-like structure Annulatascus velatisporus Helicosporium lumbricoides Nectria haematococca Ophioceras arcuatisporum Aquaphila albicans Dactylaria lignicola Hyphomycete 2 Nais inornata Nodulisporium sp. Savoryella lignicola Trichoderma sp. Verticillium sp. 4 Acremonium sp. Acrogenospora sphaerocephala Unitunicate ascomycete sp. 5 Coelomycete sp. 4 Coelomycete sp. 5 Cylindrocladium sp. Dactylaria hyalina Dactylella sp. 3 Didymosphaeria sp. Lasiodiplodia theobromae Gonytrichum chlamydosporium var. simile Hyphomycete sp. 3 Hypoxylon kretzschmarioides Lasiosphaeria immersa Nectria sp. Ophiochaeta lignicola Pleurothecium recurvatum Pseudoproboscispora aquataca Percentage occurrence Overall percentage occurrence 3 months submerged 6 months submerged 4 12 28 8 12 4 48 28 4 12 4 12 12 12 12 12 12 8 8 4 4 8 8 4 8 4 8 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 Fig 1 e Species Accumulation curve. 26 20 16 10 8 8 6 6 6 6 6 4 4 4 4 4 4 4 4 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 160 R. Kodsueb et al. Table 4 e A comparison of overlapping taxa recovered on Magnolia liliifera wood from different habitats. Taxa Habitats Naturally occurring Acanthophyses-like structure Acanthostigma minutum Acremonium sp. Acrodictys deightonii Acrodictys globulosa Acrodictys sp. Acrogenospora sphaerocephala Amphisphaeria sp. Annulatascus velatisporus Anthostomella cf. limitata Anthostomella ludoviciana Anthostomella sp. Aquaphila albicans Aquaticola ellipsoidae Aquaticola hyalomura Arthrobotrys sp. Ascotaiwania wulai Bactrodesmium longispora Bactrodesmium ramosium Basidiomycete sp. 1 Basidiomycete sp. 2 Berkleasmium concinnum Berkleasmium minutissimum Bionectria sp. Bisporella sp. Bisporomyces lignicola Bitunicate ascomycete sp. 1 Bitunicate ascomycete sp. 2 Bitunicate ascomycete sp. 3 Boerlagiomyces grandisporus Brachydesmiella caudata Canalisporium caribense Canalisporium cf. caribense Canalisporium exiguum Canalisporium pallidum Canalisporium sp. Canalisporium sp. nov.? Candelabrum brocchiatum Candelabrum sp. Catenularia malabarica Cercophora arenicola Cercophora sp. Cercospora sp. Cercosporella sp. Chaetomium globosum Chaetopsina fulva Chaetosphaeria sp. Chalara hyalina Cheiromyces sp. Chloridium botryoideum Chloridium lignicola Chloridium sp. Chloridium viride Clonostachys cylindrospora Codinaea sp. Coelomycete sp. 1 Coelomycete sp. 2 Coelomycete sp. 3 Coelomycete sp. 4 Coelomycete sp. 5 Coprinus sp. Cordana sp. Corynespora cassiicola þ þ þ þ þ Submerged baits Terrestrial baits þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ Terrestrial and freshwater lignicolous fungi 161 Table 4 e (continued ) Taxa Habitats Naturally occurring Cryptophiale udagawae Cryptophialoidea unilateralis Cylindrocarpon candidum Cylindrocladium sp. Dactylaria biseptatum Dactylaria cf. lakebarrinensis Dactylaria hyalina Dactylaria irregularis Dactylaria lignicola Dactylaria longidentata Dactylaria sp. 1 Dactylaria sp. 2 Dactylaria sp. 3 Dactylella sp. 1 Dactylella sp. 2 Dactylella sp. 3 Daldinia concentrica Diaporthe sp. 1 Diaporthe sp. 2 Diaporthe sp. 3 Diatrypella quercina Dictyochaeta australiensis Dictyochaeta simplex Dictyochaeta sp. Dictyochaeta uliginicola Dictyosporium digitatum Dictyosporium subramanianii Didymosphaeria sp. Dischloridium regenerans Dischloridium sp. Discomycete sp. Discomycetoidea aequatorialis Dokmaia monthadangii Dothidotthia sp. Ellisembia adscendens Ellisembia brachyphus Ellisembia opaca Ellisembia sp. 1 Ellisembia sp. 2 Eutypa sp. Exosporium ampullaceum Fusarium sp. 1 Fusarium sp. 2 Gliocladium sp. Gonytrichum chlamydosporium var. simile Gonytrichum macrocladum Gonytrichum sp. Guignardia sp. Halotthia posidoniae Harpographium sp. Helicoma viridis Helicomyces bellus Helicomyces macrofilamentosus Helicomyces roseus Helicomyces sp. Helicosporium pallidum Helicosporium aureum Helicosporium lumbricoides Helicosporium vegetum Helicosporium velutinum Henicospora longissima Henicospora sp. Submerged baits þ þ þ þ Terrestrial baits þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ (continued on next page) 162 R. Kodsueb et al. Table 4 e (continued ) Taxa Habitats Naturally occurring Hilberina caudata Hyalosynnema magnoliae sp. nov. Hyphomycete sp. 1 Hyphomycete sp. 2 Hyphomycete sp. 3 Hypoxylon fragiforme Hypoxylon kretzschmarioides Idriella mycoyonoidea Keissleriella fusispora Kiessleria sp. Kirschsteiniothelia thujina Lachnum sp. Lachnum virgineum Lasiodiplodia theobromae Lasiosphaeria immersa Massarina cf. walkerii Massarina sp. Melanocephala australiensis Melanocephala cupulifera Melanocephala sp. Melanochaeta hemipsila Mirandina dactylelloides Monacrosporium sp. Monodictys peruviana Monodictys sp. Mycosphaerella sp. Nais inornata Nakataea serpens Nakataea sp. Nectria aureo-fulva Nectria coccinea Nectria galligena Nectria haematococca Nectria sp. 1 Nectria sp. 2 Nectriella cf. microspora Nectriopsis sp. Neta compacta Neta sp. Nodulisporium sp. Ophioceras arcuatisporum Ophioceras sp. Ophiochaeta lignicola Orbilia xanthostigma Paecilomyces sp. Penicillium sp. Periconiella sp. 1 Periconiella sp. 2 Phaeoisaria clematidis Phaeosphaeria cf. canadensis Phaeosphaeria sp. 1 Phaeosphaeria sp. 2 Phaeostalagmus altissimus Phoma sp. Phomopsis sp. Pleurocatena acicularis Pleurophragmium sp. Pleurothecium recurvatum Pseudobotrytis terrestris Pseudoproboscispora aquataca Ramichloridium fasciculatum Ramichloridium lignicola Ramichloridium sp. Rhinocladiella cf. intermedia Submerged baits Terrestrial baits þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ Terrestrial and freshwater lignicolous fungi 163 Table 4 e (continued ) Taxa Habitats Naturally occurring Rhinocladiella mansonii Saccardoella sp. Savoryella lignicola Scolecobasidium sp. Spadicoides magnoliae sp. nov. Sporidesmium sp. 1 Sporidesmium sp. 2 Sporidesmium sp. 3 Sporoschisma saccardoi Sporoschisma uniseptatum Sporothrix sp. Stachybotrys chlorohalonata Stibella sp. Stilbella aciculosa Stilbohypoxylon quisquiliarum Tapesia fusca Torula herbarum Trichoderma cf. viride Trichoderma lignorum Trichoderma sp. Tubeufia cylindrothecia Tubeufia paludosa Unitunicate ascomycete sp. 1 Unitunicate ascomycete sp. 2 Unitunicate ascomycete sp. 3 Unitunicate ascomycete sp. 4 Unitunicate ascomycete sp. 5 Veronaea botryosa Verticillium sp. 1 Verticillium sp. 2 Verticillium sp. 3 Verticillium sp. 4 Verticillium tenerum Volutella ramkumarii Xylaria carpophila Xylaria filiformis Xylaria hypoxylon Xylaria longipes Xylaria magnoliae Xylaria polymorpha Xylaria sp. 1 Xylaria sp. 2 Xylaria sp. 3 Xylaria sp. 4 Xylomyces aquaticus Submerged baits Terrestrial baits þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ þ *Note: Bold indicates overlapping species from all habitats. Results Determination of adequacy of sampling size Species area curves for each collection of M. liliifera almost reached an asymptote because the slopes of the curves declined with the increase of sample size. At about 10 samples, the slopes were close to zero (Fig 1). Although the curve did not completely level off, the number of samples was large enough to obtain a highly representative result. Fungal diversity on submerged wood baits, and naturally occurring terrestrial and terrestrial bait samples of Magnolia liliifera Two hundred and thirty four taxa were identified from 250 wood samples from terrestrial wood, terrestrial baits and freshwater baits. Naturally occurring samples (terrestrial) yielded the greatest diversity (species richness and number of fungi per sample) with 83 taxa from 60 wood samples. This was followed by terrestrial baits (yielding 163 taxa from 140 wood samples) and submerged baits (37 taxa from 50 wood samples). 164 R. Kodsueb et al. Table 5 e Number of overlapping taxa on woody litter of Magnolia liliifera from three habitats (number in brackets is the similarity index). Terrestrial naturally occurring Submerged Baits Submerged baits Terrestrial baits 8 (0.13) 32 (0.26) - 14 (0.14) *Overlapping between all habitats ¼ five species (similarity index ¼ 0.17) Thirty-seven taxa were identified from 50 submerged M. liliifera wood baits (Table 3). The most common taxa were Candelabrum brocchiatum (on 26% of wood samples), Berkleasmium minutissimum (20%), Sporidesmium sp. (16%), Aquaticola hyalomura (10%) and Sporoschisma uniseptatum (8%). Only 14 fungal taxa were obtained from samples submerged for 3 months, while 31 taxa were obtained from samples submerged for 6 months (Table 3). Eighteen species (49% of total species) were represented by only one record and can be regarded as rare. Five taxa were unidentified including one ascomycete, two coelomycetes and two hyphomycetes since there was not enough material and some were immature. The list of overlapping fungi obtained from the three habitats is shown in Table 4, with only five taxa overlapping between all habitats. Of the 214 taxa recorded from terrestrial samples (baits and natural occurrence), only six (2.8% of all terrestrial taxa) were overlapping at 3 months and 17 (7.9% of all terrestrial taxa) overlapping at 6 months (Table 5). Fungal communities on Magnolia liliifera wood from different habitats 3D-correspondence analysis of fungi on wood of M. liliifera from different habitats (terrestrial naturally occurring, terrestrial baits and submerged baits) showed three fungal communities; naturally occurring terrestrial; MLN, submerged baits; MLS and terrestrial baits; MLT (Fig 2). The percentages of abundance for each fungal taxon obtained from submerged baits are listed in Table 3, and those from naturally occurring and terrestrial baits in Kodsueb et al. (2008a; b). Dominant species from the three habitats, species richness, species evenness, number of fungi per sample, Shannon diversity index (H) and Simpson diversity index (D) of each habitat are shown in Table 6. distribution arrangement of fungal community based on the frequencies of occurrence of fungal taxa in each sample types, corroborated the existence of two distinct habitats supporting distinct fungal communities (Fig 2). Eight taxa overlapped on naturally occurring terrestrial and submerged baited samples (similarity index ¼ 0.13). There were 14 overlapping taxa on submerged baits and terrestrial baits and 32 overlapping taxa on naturally occurring terrestrial samples and terrestrial baits (similarity index of 0.14 and 0.26, respectively). Discussion Species composition and species abundance This is the first study investigating the differences in fungal communities on Magnolia wood in a freshwater stream as compared to the same wood in the adjacent riparian terrestrial habitat in a tropical rainforest. The dominant fungi on naturally occurring terrestrial samples were similar to those from terrestrial baits, but significantly different from those on submerged baits (Table 6). The dominant fungi in the present study also differed from those usually found to be common on terrestrial wood (Huhndorf and Lodge, 1997; Crites and Dale, 1998; Allen et al., 2000). Asexual fungi were the dominant group on wood in all habitats. Basidiomycetes were present only on terrestrial samples, with very low species numbers; they were absent in the freshwater habitat. This is consistent with previous studies where basidiomycetes were rare or absent on submerged wood (Wong et al., 1998; Shearer et al., 2007; Hyde et al., 2016). The most common species on freshwater baits was Candelabrum brocchiatum. Corynespora cassiicola was dominant on natural terrestrial wood and Lasiodiplodia theobromae was dominant on terrestrial baits. The most common taxa on Magnolia liliifera from freshwater baits overlapped with those Comparison between fungal communities on terrestrial woody litter and submerged woody litter 3D-correspondence analysis (Fig 2) showed that the fungal communities on naturally occurring terrestrial M. liliifera wood (MLN) and terrestrial baited wood (MLT) are more similar to each other than to those on submerged wood baits (MLS) (see similarity index in Table 5). The fungal communities inhabiting terrestrial samples revealed higher similarities considering species richness and fungal taxa composition than those in the submerged habitat. The 3D correspondence analysis, which provided an integrated overview of Fig 2 e 3D-correspondence analysis of fungal succession on woody litter of Magnolia liliifera (MLN [ naturally occurring terrestrial samples, MLS [ submerged baits, MLT [ terrestrial baits). Terrestrial and freshwater lignicolous fungi 165 Table 6 e Diversity indices of saprotrophic fungi recovered from Magnolia liliifera wood. Naturally terrestrial Number of samples examined Average number of fungi per wood sample Species richness Species evenness Shannon indices Simpson indices Dominant species (percentage occurrence) Submerged baits Terrestrial baits (MLT) 60 1.4 50 0.7 140 1.2 83 0.890 3.923 0.9640 Corynespora cassiicola (60) Phaeoisaria clematidis (20) Anthostomella ludoviciana (16.7) Canalisporium caribense (16.7) Diaporthe sp. 2 (16.7) Brachydesmiella cauata (13.3) Massarina sp. (13.3) Sporidesmium sp. 1 (13.3) Ellisembia brachyphus (11.7) Phomopsis sp. (11.7) 37 0.899 3.245 0.9439 Candelabrum brocchiatum (26) Berkleasmium minutissimum (20) Ellisembia opaca (16) Aquaticola hyalomura (10) Phaeoisaria clematidis (8) Sporoschisma uniseptatum (8) Acanthophyses-like structure(6) Annulatascus velatisporus(6) Helicosporium lumbricoides(6) Nectria haematococca(6) Ophioceras arcuatisporum(6) 163 0.834 4.248 0.9731 Lasiodiplodia theobromae (42.9) Nectria coccinea (37.1) Xylaria carpophila (23.6) Canalisporium pallidum (17.1) Dactylaria hyaline (17.1) Chloridium viride (15.7) Phaeostalagmus rossicus (15) Gonytrichum macrocladum (14.3) Dactylaria biseptata (13.6) Boerlagiomyces grandisporus (11.4) Coprinus sp. (11.4) on other unidentified naturally occurring wood submerged in the same stream (Kodsueb, pers. obs.), where Acanthophyseslike structure, Annulatascus velatispora, Helicosporium lumbricoides, Ophioceras arcuatisporum and Sporoschisma uniseptatum were the most common species. brocchiatum, Savoryella lignicola and Sporoschisma uniseptatum (Hyde and Goh, 1998; Tsui et al., 2000; Ho et al., 2002; Fryar et al., 2004a, b; Tsui and Hyde, 2004). Nais inornata and Nectria haematococca were found both here and in temperate freshwater (Shearer and von Bodman, 1983; Shearer and ZareMaivan, 1988). Are the fungi on terrestrial Magnolia liliifera wood different to those on submerged wood samples? Habitat recurrence of fungi on Magnolia liliifera Fungal successions and general diversity studies of fungal communities on decomposing wood have been performed worldwide in freshwater ecosystems (Goh and Hyde, 1996; Ho et al., 2001, 2002; Tsui et al., 2000, 2001a, b; 2002; 2003; Sivichai et al., 2000, 2002; Maria and Sridhar, 2004). There were some overlapping species in M. liliifera wood baits compared to fungi living on other wood in freshwater ecosystems in previous studies, including Annulatascus velatispora, Aquaphila albicans, Aquaticola hyalomura, Candelabrum brocchiatum, Helicosporium lumbricoides, Ophioceras arcuatisporum, Nais inornata, Pseudoproboscispora aquataca, Savoryella lignicola and Sporoschisma uniseptatum (Ho et al., 2001; 2002; Tsui et al., 2000; 2001a, b; 2002; 2003; Sivichai et al., 2000; 2002). Berkleasmium minutissimum, the dominant fungus on submerged baits of M. liliifera has not been recorded as dominant in other studies. Conversely, some common genera obtained from submerged wood worldwide e.g. Halosarpheia, Jahnula and Lophiostoma, were not found in the present study. A possible explanation for this may be the short time that the bait samples were submersed compared with the naturally occurring samples or the wood type. Several studies have shown that different fungal communities occur in temperate and tropical freshwater habitats (Goh and Hyde, 1996; Wong et al., 1998; Ho et al., 2001; Kane et al., 2002; Tsui and Hyde, 2003; Tsui et al., 2003; Shearer et al., 2007). The taxa obtained in this study are similar to those of other tropical areas and less similar to those of temperate regions. Species which overlap with those of other tropical studies include Annulatascus velatispora, Candelabrum Fungal species found on Magnolia liliifera varied between habitats. From a total of 1,074 collections (234 taxa), 20 taxa were unique to the freshwater samples and 197 taxa were unique to the terrestrial samples, with only 17 overlapping species. Some fungi collected from submerged wood are typical freshwater fungi (e.g. Annulatascus velatispora, Aquaticola hyalomura, Ophioceras arcuatisporum, Nais inornata, Pseudoproboscispora aquataca, Savoryella lignicola and Sporoschisma uniseptatum), which are believed to have adaptation mechanisms for living in freshwater habitats (Hyde and Goh, 2003; Cai et al., 2006b). On the other hand, typical terrestrial fungi such as Hypoxylon sp., Guignardia sp., Massarina cf. eburnea and Didymosphaeria sp. were recorded from terrestrial wood. However, Lasiodiplodia theobromae, which has previously been assumed to be unique to the terrestrial environment, was recorded twice from submerged wood. Similarly, species of Nectria, which are common terrestrial fungi on wood, bark and other fungi (Samuels, 1976) and believed not to grow well in aquatic habitats (Fryar et al., 2004b), were found on submerged wood in the present study and in some previous studies (Willoughby and Archer, 1973; Shearer and von Bodman, 1983; Ho et al., 2002). In previous studies, some species of terrestrial fungi that were present on wood before submergence, survived and were found again after submergence (Shearer and Webster, 1991; Fryar et al., 2004b). It is not clear whether these fungi are habitat exclusive and continue to thrive in a freshwater environment. It is unlikely that the fungi on submerged wood in the present study originated from the wood prior to submersion, since it had been oven-dried before submergence, 166 though some resistant spores may have survived. Fryar et al. (2004b) believed that differences in fungal assemblages at the beginning and after 3 months submergence might be attributed to a difference in substratum rather than the assemblage of fungi already present in the sample. The similarity of fungi in the present study in terrestrial (naturally occurring and baited samples) and freshwater habitats (submerged baits) was low (17 overlapping species), suggesting that most fungi growing on terrestrial wood do not thrive in freshwater habitats, concurring with previous studies (Fryar et al., 2004b; Cai et al., 2006b; Pinnoi et al., 2006; Pinruan et al., 2007). Why do submerged samples support lower fungal diversity? The present finding of less diversity on submerged wood concurs with that of Pinruan et al. (2007). Generally, longer freshwater studies with prolonged sampling, yield greater fungal diversity (Shearer and Crane, 1986; Ho et al., 2001; Tsui et al., 2001b, 2003b). Compared to other freshwater studies, fewer samples were examined in the present study and the submergence period was shorter (Ho et al., 2002; Sivichai et al., 2002), which may partly explain why more species were obtained from terrestrial samples of M. liliifera than from the freshwater samples. However, only between 25 and 80 taxa were found on submerged wood in streams in 21 studies in the tropics and subtropics (Ho et al., 2001; Cai et al., 2003; Fryar et al., 2004b), indicating terrestrial fungi are much more diverse. Another reason for the low numbers of fungi on submerged samples is that few taxa are adapted for a submerged existence in freshwater ecosystems. These adaptations have been discussed in Hyde et al. (2016). Acknowledgements Kevin D. Hyde thanks the Chinese Academy of Sciences, project number 2013T2S0030, for the award of Visiting Professorship for Senior International Scientists at Kunming Institute of Botany. 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