The spatial distribution of fungi on decomposing alder leaves in a freshwater stream

Summary Transects were cut from alder leaves incubated in a freshwater stream and plated as quadrats so that fungal isolates could be mapped by reconstruction of each transect. Initially there was fewer than one aquatic hyphomycete colonist per quadrat, but the mode increased to 6–7 then progressively decreased to <1. Numbers of species of aquatic hyphomycetes per quadrat rose and fell similarly with a maximum mode of 3–4, as did species per transect with a maximum of 11 and a diversity of 16, comprising 6 dominant species and about 10 occasional species. The latter showed no pattern of appearance but the dominant group was established early and persisted in a dynamic equilibrium. Aquatic hyphomycetes were initially randomly distributed but developed progressively into clumped consortia which persisted after peak colonization, declining as leaf degradation became total. Colonies of the most persistent aquatic hyphomycete species were initially discrete,developing into a complex network of overlapping colonies and species, no two of which showed positive association. These complexes broke down to large colonies of a few species and finally to 1–2 small colonies. The pattern of isolates of the 18 genera of other fungi was the reverse of that for aquatic hyphomycetes. Only Cladosporium, Epicoccum and Fusarium were important colonizers. The first two appear to be inhibited by aquatic hyphomycetes, but were found to degrade substrates representative of cell-wall polymers vigorously whereas aquatic hyphomycetes showed varied degradative ability. Leaf transects were examined by S.E.M. and epifluorescent microscopy so that hyphal colonization could be followed at progressive stages of leaf degradation. Bacteria on transects were patchily distributed, the temporal pattern indicating inhibition by aquatic hyphomycetes and colonization of senescent hyphae.     

Studies on the aquatic hyphomycetes of a sulfur spring in the Western Ghats, India

Studies on the occurrence of aquatic hyphomycetes were carried out in Panekal sulfur spring in the Western Ghats, India by incubation of leaf litter and analysis of natural foam and of induced foam. Sampling was done once every three months over a period of two years from September 1989 to June 1991. The temperature, pH, dissolved oxygen, and sulfide content of water were also measured. No fungi were observed within the spring, whereas 16 species belonging to 13 genera were isolated from two outflow sites of the stream. The percent frequency of Triscelophorus monosporus was high (24.0%). The temperature of water in the spring ranged between 30.0 and 38.5°C and the sulfide content between 3.2 and 4.3 mg 1^–1. Studies showed that sulfide water (4.0 mg 1^–1). from the spring inhibited the growth of the colonies of Dactylella aquatica, Phalangispora constricta, Tetracladium setigerum, Vermispora cauveriana, and Wiesneriomyces laurinus. When the leaves colonized by aquatic hyphomycetes were incubated at different temperatures in sulfur-spring water and stream water separately, sporulation was not observed in any of the fungi at and above 35°C except Phalangispora constricta, which could sporulate at 35°C. At lower temperatures (15–30°C) relatively fewer species were found to sporulate in sulfur-spring water than in stream water. Correspondence to: K.M. Kaveriappa     

Colonization,Species Composition,and Conidial Production of Aquatic Hyphomycetes on Chirpine Needle Litter in a Freshwater Kumaun Himalayan Stream

Aquatic hyphomycetes colonizing the submerged chirpine (pinus roxburghii SARG .) needle litter in a high altitude, Kumaun Himalayan stream were studied. 15 species belonging to different genera of aquatic Hyphomycetes have been recognized as the colonizers of chirpine needle litter. Clavariopsis aquatica, Heliscus lugdunensis, Lunulospora cymbiformis, Triscelophorus acuminatus and T. monosporus were found with a high frequency of occurrence. The conidial production was highest in Flagellospora penicillioides, however, Campylospora chaetocladia, L. cymbiformis and T. acuminatus had less number of conidia per unit area of pine needles. The chirpine needle litter decomposition in the freshwater habitat is also discussed.     

The molecular phylogeny of aquatic hyphomycetes with affinity to the Leotiomycetes

Aquatic hyphomycetes play a key role in decomposition of submerged organic matter and stream ecosystem functioning. We examined the phylogenetic relationships among various genera of aquatic hyphomycetes belonging to the Leotiomycetes (Ascomycota) using sequences of internal transcribed spacer (ITS) and large subunit (LSU) regions of rDNA generated from 42 pure cultures including 19 ex-types. These new sequence data were analyzed together with additional sequences from 36 aquatic hyphomycetes and 60 related fungi obtained from GenBank. Aquatic hyphomycetes, characterized by their tetraradiate or sigmoid conidia, were scattered in nine supported clades within the Helotiales (Leotiomycetes). Tricladium, Lemonniera, Articulospora, Anguillospora, Varicosporium, Filosporella, and Flagellospora are not monophyletic, with species from the same genus distributed among several major clades. The Gyoerffyella clade and the Hymenoscyphus clade accommodated species from eight and six different genera, respectively. Thirteen aquatic hyphomycete taxa were grouped in the Leotia-Bulgaria clade while twelve species clustered within the Hymenoscyphus clade along with several amphibious ascomycetes. Species of Filosporella and some species from four other aquatic genera were placed in the Ascocoryne-Hydrocina clade. It is evident that many aquatic hyphomycetes have relatives of terrestrial origin. Adaptation to colonize the aquatic environment has evolved independently in multiple phylogenetic lineages within the Leotiomycetes.     

The ecology of aero-aquatic hyphomycetes

We observed 35 species of aero-aquatic hyphomycetes belonging to 21 genera and 4 unidentified taxa of the Clavariopsis, Dactilella, Helicoon, and Tetracladium genera. Substrate preferences were detected in some species. Species such as Anguillospora longissima, Tetracladium marchalianum, and Fusarium aqueductum were found on different types of substrata. Only 11 species are developed on pine needles, while 20 species were observed on leaves. Eight species of aquatic hyphomycetes (Alatospora acuminata, Angullospora aquatica, Lemonniera aquatica, Tetracladium setigerum, Tricladium angulatum, Tripospermum campelopardus, Septonema secedens, and Spermospora sp.) were revealed on leaves taken from water habitats, whereas 15 species were found in litter. The jointly grown species did not demonstrate any inhibiting effects on each other except for Tripospermum campelopardus, which inhibited the growth of other species (Table 3). The microcyclic development of a Dactilella sp. was detected for the first time while being incubated on leaves in the laboratory. The frequency of appearance changes, and the diversity of the species of aquatic hyphomycetes appears to be higher on intact leaves than on skeletonized ones. The seasonal dynamics of aquatic hyphomycetes with two peaks of mass conidia development (vernal and sharper autumnal) were described. Some species were observed throughout the entire vegetation season. The conidial development in the leaf samples incubated in the laboratory lasted for 30–40 days. Therefore, the method of incubation for 7–10 days recommended in the literature did not allow for the complete investigation of species diversity in aquatic hyphomycetes.     

Do Invertebrate Activity and Current Velocity Affect Fungal Assemblage Structure in Leaves?

In this study we assessed the effect of current velocity and shredder presence, manipulated in artificial channels, on the structure of the fungal assemblage colonizing alder (Alnus glutinosa (L.) Gaertner ) leaves incubated in coarse and fine mesh bags. Fungal sporulation rates, cumulative conidial production and number of species of aquatic hyphomycetes were higher in leaves exposed to high rather than to low current velocity. The opposite was observed regarding Simpson's index (D) on the fungal assemblage. Some species of aquatic hyphomycetes were consistently stimulated in high current channels. No effect of shredders or of mesh type was observed. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Biodiversity of freshwater fungi

There are more than 600 species of freshwater fungi with more known from temperate, as compared to tropical regions. These includeca 340 ascomycetes, 300 deuteromycetes, and a number of lower fungi which are not discussed here.Aniptodera, Annulatascus, Massarina, Ophioceras andPseudohalonectria are common freshwater ascomycetes, which appear to be well adapted for this lifestyle either in their ascospore types or their competitive-degradative characters. The most common genera of wood-inhabiting deuteromycetes includeCancellidium, Dactylaria, Dictyosporium andHelicomyces. They are categorized into four groups depending on their form and life style: the ingoldian hyphomycetes; the aero-aquatic hyphomycetes; the terrestrial-aquatic hyphomycetes; and the submerged-aquatic hyphomycetes. The adaptations of aquatic fungi for their dispersal and subsequent attachment to new substrates are discussed.     

Leaf Decomposition in a Mountain Stream in the Sultanate of Oman

Decomposition of Juglans regia leaves was studied in fine and coarse mesh bags in a permanent mountain stream in Oman. A rapid initial mass loss, attributed to leaching, was followed by a more gradual decline. Daily exponential decay rates (k) calculated over 32 days were 0.011 (fine mesh litter bags) and 0.014 (coarse mesh litter bags). The difference between bag types was not significant, suggesting limited impact of leaf‐shredding invertebrates. Ergosterol levels on leaves from fine mesh bags peaked at 0.3 mg g^–1 AFDM after 16 days of stream exposure. During the experimental period, which followed the annual leaf fall, the concentration of aquatic hyphomycete conidia in the stream varied between 82 and 1362 l^–1. Based on the morphology of conidia found in the water column or released from leaves, we identified 14 species of aquatic hyphomycetes. Tetracladium apiense was the most common taxon (62.2% of conidia in water column during the field experiment). Three other Tetracladium species contributed another 8%. Plating out leaf particles yielded common epiphytic taxa such as Alternaria sp., Aureobasidium pullulans and Phoma sp. The measured metrics of leaf decay in this desert stream fall within the range of values observed in temperate and tropical streams, with clear evidence for an early leaching phase, and no evidence of a strong impact of leaf shredders. The community of aquatic hyphomycetes appears impoverished. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Temperature and Sporulation of Aquatic Hyphomycetes

Temperature appears to be an important factor affecting the occurrence and distribution of aquatic hyphomycetes, the dominant leaf litter-decomposing fungi in streams. We compared conidium production by eight species of aquatic hyphomycetes grown on yellow poplar leaves in stream-simulating microcosms at three temperatures (15, 20, and 25°C). The greatest conidium production occurred at 15°C for one species, 20°C for two species, and 25°C for two species. Two species produced similar numbers of conidia at 20 and 25°C, and one species produced similar numbers of conidia at all three temperatures. Linear growth rates were determined on malt extract agar. Six species had the same pattern of temperature responses for growth on malt extract agar as for sporulation on leaves, as shown by the positive correlations between the two parameters at the three temperatures. The species examined also exhibited differences in number of conidia produced from a similar amount of leaf material at a given temperature. These differences appeared to be due primarily to differences in individual conidium mass (determined by weighing conidia produced from cultures), as shown by the relationship of the type Y = k/X (r² = 0.96), where Y is the number of conidia produced, X is the individual conidium mass in milligrams, and k is a constant empirically determined to be 2.11. This finding supports the hypothesis that aquatic hyphomycetes allocate similar amounts of their resources to reproduction but vary with respect how these resources are partitioned into reproductive units (conidia).     

Colonization pattern of aquatic hyphomycetes on leaf packs in subtropical stream

Steralized leaf pieces of eight plants (Eucalyptus rostrata, Phoenix dactylifera, Phragmites australis, Musa nana, Salix subserrata, Cyperus alopecuroides, Ricinus communis and Eichhornia crassipes) were submerged in the Nile stream. Thirty-nine species of aquatic hyphomycetes were colonized on the plant leaves. Eucalyptus was the best substratum (30 species) for colonization by aquatic hyphomycetes. Phoenix (14 species), Phragmites (11 species), Salix (9 species), Musa (8 species), Ricinus and Cyperus (6 species) were also good substrata for aquatic hyphomycetes colonization. Echhornia (aquatic plant) was not suitable for aquatic hyphomycetes colonization. Alatospora acuminata, Triscelophorus monosporus and Tetracladium marchalianum were found to be the major colonizers on all leaf materials. Temperature and dissolved oxygen were the highest physico-chemical parameters affecting the aquatic hyphomycetes colonization. These results indicate that aquatic hyphomycetes have been shown to be successful colonizers on plants leaf in river Nile in Egypt as subtropical region. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of drying and freezing autumn leaves on leaching and colonization by aquatic hyphomycetes

1. Drying or freezing autumn-shed leaves of Betula papyrifera, Ulmus americana and Acer saccharum increased the leaching of soluble substances. The difference between fresh and treated leaves was most pronounced in birch. 2. Dried and frozen leaves supported more species of aquatic hyphomycetes during early decay, and conidium production from these fungi was higher. 3. Anguillospora filiformis was the most prolific spore producer on most samples. It declined in later stages of decay, when Articulospora tetradadia became more common. Crucella subtilis, an unspecialized parasite of aquatic hyphomycetes, was quite common in many samples. 4. In the autumn of 1990, 50% of elm and maple leaves, and 30% of birch leaves were shed before the air temperature dropped below 0°C; 10% (elm, maple) to 30% (birch) experienced more than one freezing period before being shed. 5. Water-soluble extracts from leaves inhibited radial growth of four fungal species. Maple extracts generally had the greatest inhibitory effect, followed by birch and elm.     

Endophytic aquatic hyphomycetes in roots of riparian tree species of two Western Ghat streams

Bubble chamber incubation of surface sterilized segments of root bark and xylem of 10 riparian tree species of the Sampaje (475–500 m asl) and V?Badaga (765–800 m asl) stream reaches of the Western Ghats yielded 20 species of endophytic aquatic hyphomycetes. Anguillospora crassa, A. longissima and Cylindrocarpon sp. were among the top five species in streams. A two-way ANOVA showed significantly higher species richness and counts of conidium in the tree species of Sampaje compared to V?Badaga (p < 0.001), while two variables were not significantly different between bark and xylem. The total number of species recovered was slightly higher in bark than in xylem (14–19 vs. 13–17 spp.) and the average species richness between tissues did not differ significantly except for one tree species (Madhuca neriifolia: p < 0.05). The release of conidia from bark of only three tree species was significantly higher than from xylem (M. neriifolia and Canarium strictum: p < 0.05; Vateria indica: p < 0.01). Sørensen’s similarity index for bark as well as xylem between tree species was higher in Sampaje stream than in V?Badaga stream (0.45–0.78 vs. 0.25–0.61). The diversity of aquatic hyphomycetes in bark and xylem was higher in the trees of Sampaje than V?Badaga (3.1–3.3 vs. 2.7). A cluster analysis of aquatic hyphomycetes in bark and xylem resulted in two groups coinciding with the two streams. The results of this study revealed that assemblage and diversity of endophytic aquatic hyphomycetes in riparian tree roots are high in the mid-altitude Sampaje stream as previously documented for saprotrophic aquatic hyphomycetes.     

Diversity and activity of aquatic fungi under low oxygen conditions

1. The objective was to test whether a decrease in oxygen concentration in streams affects the diversity and activity of aquatic hyphomycetes and consequently leaf litter decomposition. 2. Senescent leaves of Alnus glutinosa were immersed for 7 days in a reference stream, for fungal colonization, and then incubated for 18 days in microcosms at five oxygen concentrations (4%, 26%, 54%, 76% and 94% saturation). Leaf decomposition (as loss of leaf toughness), fungal diversity, reproduction (as spore production) and biomass (ergosterol content) were determined. 3. Leaf toughness decreased by 70% in leaves exposed to the highest O₂ concentration, whereas the decrease was substantially less (from 25% to 45%) in treatments with lower O₂. Fungal biomass decreased from 99 to 12 mg fungi g⁻¹ ash‐free dry mass on exposure to 94% and 4% O₂ respectively. Sporulation was strongly inhibited by reduction of dissolved O₂ in water (3.1 × 10⁴ versus 1.3 × 10³ spores per microcosms) for 94% and 4% saturation respectively. 4. A total of 20 species of aquatic hyphomycetes were identified on leaves exposed to 94% O₂, whereas only 12 species were found in the treatment with 4% O₂ saturation. Multidimensional scaling revealed that fungal assemblages exposed to 4% O₂ were separated from all the others. Articulospora tetracladia, Cylindrocarpon sp. and Flagellospora curta were the dominant species in microcosms with 4% O₂, while Flagellospora curvula and Anguillospora filiformis were dominant at higher O₂ concentrations. 5. Overall results suggest that the functional role of aquatic hyphomycetes as decomposers of leaf litter is limited when the concentration of dissolved oxygen in streams is low.     

The cooler the better: Increased aquatic hyphomycete diversity in subtropical streams along a neotropical latitudinal gradient

Aquatic hyphomycetes are microbial decomposers in freshwater environments that, together with detritivores, play an essential role in the functioning of low-order streams. Here, we evaluated aquatic hyphomycetes communities associated with decomposing leaves of Nectandra megapotamica, a common Neotropical riparian tree, along a subtropical-tropical latitudinal gradient. Two forest streams located in subtropical regions and 3 in tropical regions were selected. We identified 29 species of aquatic hyphomycetes, 22 (75.8%) in subtropical streams and 15 (51.7%) in tropical streams. We also found a higher fungal biomass in subtropical streams. However, the amounts of leaf mass loss did not differ between regions, but the values were higher in summer than in winter. High temperature, pH and electrical conductivity values, as well as low dissolved oxygen levels, negatively affected spore production. These results suggest that the subtropical-tropical gradient is an important predictor of aquatic hyphomycete diversity; however, the observed species had different sensitivities to local environmental factors.     

Effects of Zinc on Leaf Decomposition by Fungi in Streams: Studies in Microcosms

The effect of zinc on leaf decomposition by aquatic fungi was studied in microcosms. Alder leaf disks were precolonized for 15 days at the source of the Este River and exposed to different zinc concentrations during 25 days. Leaf mass loss, fungal biomass (based on ergosterol concentration), fungal production (rates of [1-¹⁴C]acetate incorporation into ergosterol), sporulation rates, and species richness of aquatic hyphomycetes were determined. At the source of the Este River decomposition of alder leaves was fast and 50% of the initial mass was lost in 25 days. A total of 18 aquatic hyphomycete species were recorded during 42 days of leaf immersion. Articulospora tetracladia was the dominant species, followed by Lunulospora curvula and two unidentified species with sigmoid conidia. Cluster analysis suggested that zinc concentration and exposure time affected the structure of aquatic hyphomycete assemblages, even though richness had not been severely affected. Both zinc concentration and exposure time significantly affected leaf mass loss, fungal production and sporulation, but not fungal biomass. Zinc exposure reduced leaf mass loss, inhibited fungal production and affected fungal reproduction by either stimulating or inhibiting sporulation rates. The results of this work suggested zinc pollution might depress leaf decomposition in streams due to changes in the structure and activity of aquatic fungi.     

水生丝孢菌研究进展

本文回顾了自1942年水生丝孢菌发现以来,该类群的研究进展。世界范围内已有约80属、200种的水生丝孢菌被报道。中国发现20属、28种。12属中的15种水生丝孢菌的有性阶段已发现,它们大多数是子囊菌,少数为担子菌。生理学研究证实,该类真菌普遍地可以分泌胞外多糖酶,从而能较好地利用多种碳源。其它生理特性,如氮源和矿物的需求等,本文也作了综述。一些植物寄生性水生丝孢菌也有报道。生态学研究表明,秋季和冬季是水生丝孢菌大发生季节。它们在水环境中的重要作用,文中也作了较详细的讨论。最后涉及的是水生丝孢菌的传布和陆生情况的发现。本文还对该类群真菌的研究前景作了展望。     

Lignocellulolytic Enzyme Production by Aquatic Hyphomycetes Species Isolated from the Nile's Delta Region

Twenty-six species of aquatic hyphomycetes were isolated from woody sources (unidentified wood segments, leaf skeletons and neck of leaves and bark) in the North River Nile (Delta region). Alatospora acuminata, Anguillospora crassa, Flagellaspora penicillioides, Lunulospra curvula, Tetracladium marchalianum and Triscelophorus monosporus were the most common species. Temperature was the highest physico-chemical parameter affecting the aquatic hyphomycetes occurrence. Twelve species of hyphomycetes, isolated from woody substrates, were screened for their ability to produce extracellular lignocellulolytic enzymes on solid media. The enzymes tested included: endoglucanase, endoxylanase, beta-glucosidase, laccase, peroxidase, polyphenoloxidase, tyrosinase and beta-xylosidase. Three species, A. acuminata, F. penicillioides, T. monosporus, were positive for all tested enzymes. Also, A. longissima was positive for all enzymes except lignin-peroxidase. The ability to produce cellulase was 100% for all species while only, four species were positive for lignin-peroxidase. The ability of the species to produce other lignocellulotic enzyme ranged from 50% to 83%. Freshwater hyphomycetes have been shown to produce a rich array of enzymes able to degrade the polysaccharides of plant debris.     

Studies on synthetic hybrids of british species of Senecio: I Senecio viscosus L. × S. vulgaris L.

Summary

Four aquatic hyphomycetes and one terrestrial fungus were examined for their responses to the phenoxy herbicides (±)-MCPP and 2,4-D as both single and binary preparations with respect to hyphal extension, sporulation and respiration. Hyphal extension of all species was unaffected at concentrations less than 100 mg l^?1. At higher concentrations there was a variable inhibitory response to the herbicides but no clear pattern was observed between the five fungi. The binary herbicide mixture had a weak synergistic effect on inhibition of growth rate. For the four aquatic hyphomycetes sporulation was reduced at several herbicide concentrations, but there was no consistent reduction over the experimental period. Flagellospora curvula and Clavariopsis aquatica showed increased sporulation at 100 and 1000 mg l^?1 for only some herbicide combinations. The respiration rates of the fungi varied with species and herbicide concentration and ranged between stimulation at 100 mg l^?1 to inhibition at 4000 mg l^?1. The results indicate that the five fungi are not likely to be severely effected by the phenoxy herbicides at concentrations normally occurring in the field. The possible effects of these herbicides on nutrient cycling are briefly discussed.     

Fungal Propagules and DNA in Feces of Two Detritus-Feeding Amphipods

Aquatic shredders (leaf-eating invertebrates) preferentially ingest and digest leaves colonized by aquatic hyphomycetes (fungi). This activity destroys leaf-associated fungal biomass and detritial resources in streams. Fungal counter-adaptations may include the ability to survive passage through the invertebrate's digestive tract. When fecal pellets of Gammarus tigrinus and Hyalella azteca were incubated with sterile leaves, spores of nine (G. tigrinus) and seven (H. azteca) aquatic hyphomycete species were subsequently released from the leaves, indicating the presence of viable fungal structures in the feces. Extraction, amplification, and sequencing of DNA from feces revealed numerous fungal phylotypes, two of which could be assigned unequivocally to an aquatic hyphomycete. The estimated contributions of major fungal groups varied depending on whether 18S or ITS sequences were amplified and cloned. We conclude that a variable proportion of fungal DNA in the feces of detritivores may originate from aquatic hyphomycetes. Amplified DNA may be associated with metabolically active, dormant, or dead fungal cells.     

Natural UVR Does Not Affect Decomposition by Aquatic Hyphomycetes

We performed field and laboratory experiments to evaluate the effect of solar radiation (UVR and PAR) on leaf litter decomposition, fungal biomass and sporulation rates, in the Andean Patagonia, where high UVR levels are common. Leaves of Alnus glutinosa exposed to three treatments, normal radiation (PAR + UVR), protected from UVR and protected from total radiation (SHADE) by plastic films lost 31–37% of their mass. Leaves of Nothofagus pumilio lost 61–64% of their mass under the same conditions. For both leaf species, differences in mass losses among treatments were not statistically significant. Sporulation rates were significantly lower in the SHADE treatment. Fungal biomass accounted for 6.2 to 7.1% of leaf mass, without significant differences among treatments. In the laboratory, leaf discs of A. glutinosa colonized by single species of aquatic hyphomycetes (Articulospora tetracladia, Flagellospora curta or Lunulospora curvula) and exposed to or protected from UVR did not differ in mass loss and sporulation rates. Pure cultures of two fungal species grew at the same rates when exposed to light (PAR and PAR + UVR) or to the SHADE. In summary, we found no evidences that current high levels of UV radiation affect litter decomposition mediated by aquatic hyphomycetes. (© 2010 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)