Endophytic fungal communities in woody perennials of three tropical forest types of the Western Ghats,southern India

Fungal endophytes of tropical trees are expected to be exceptionally species rich as a consequence of high tree diversity in the tropics and the purported host restriction among the endophytes. Based on this premise, endophytes have been regarded as a focal group for estimating fungal numbers because their possible hyperdiverse nature would reflect significantly global fungal diversity. We present our consolidated ten-year work on 75 dicotyledonous tree hosts belonging to 33 families and growing in three different types of tropical forests of the NBR in the Western Ghats, southern India. We conclude that endophyte diversity in these forests is limited due to loose host affiliations among endophytes. Some endophytes have a wide host range and colonize taxonomically disparate hosts suggesting adaptations in them to counter a variety of defense chemicals in their hosts. Furthermore, such polyphagous endophytes dominate the endophyte assemblages of different tree hosts. Individual leaves may be densely colonized but only by a few endophyte species. It appears that the environment (the type of forest in this case) has a larger role in determining the endophyte assemblage of a plant host than the taxonomy of the host plant. Thus, different tropical plant communities have to be studied for their endophyte diversity to test the generalization that endophytes are hyperdiverse in the tropics, estimate their true species richness, and use them as a predictor group for more accurate assessment of global fungal diversity.     

Leaf shedding and weather in tropical dry-seasonal forest shape the phenology of fungi – Lessons from two years of monthly surveys in southwestern Panama

In the present study, conducted in a secondary dry-seasonal forest in the pacific lowlands of southwestern Panama over 2 years, fungal diversity is linked to plant phenology, litter, and climatic data. Agaricales fungi showed maximum species richness at the beginning of rainy seasons, probably due to the important litter accumulation during the dry season and the increase in humidity favoring fungal growth. Species richness declined during the wet season possibly due to torrential rains, moulds, and decreasing availability of nutrients. Occurrence of foliar pathogenic microfungi correlated negatively with flushing of new leaves at the beginning of the rainy season. Their incidence increased during the wet season and remained high during the dry season. Synchronization of leaf shedding in most tree species significantly reduced the yearly incidence of foliar pathogenic fungi causing an annual turn-over of fungal pathogens that probably contributes to maintain a high diversity of plant pathogenic species.     

Host associations and beta diversity of fungal endophyte communities in New Guinea rainforest trees

Processes shaping the distribution of foliar fungal endophyte species remain poorly understood. Despite increasing evidence that these cryptic fungal symbionts of plants mediate interactions with pathogens and herbivores, there remain basic questions regarding the extent to which dispersal limitation and host specificity might shape fungal endophyte community composition in rainforests. To assess the relative importance of spatial pattern and host specificity, we isolated fungi from a sample of mapped trees in lowland Papua New Guinea. Sequences of the internal transcribed spacer (ITS) region were obtained for 2079 fungal endophytes from three sites and clustered into molecular operational taxonomic units (MOTUs) at 95% similarity. Multivariate analyses suggest that host affinity plays a significant role in structuring endophyte community composition whereas there was no evidence of endophyte spatial pattern at the scale of tens to hundreds of metres. Differences in endophyte communities between sampled trees were weakly correlated with variation in foliar traits but not with tree species relatedness. The dominance of relatively few generalist endophytes and the presence of a large number of rare MOTUs was a consistent observation at three sites separated by hundreds of kilometres and regional turnover was low. Host specificity appears to play a relatively weak but more important role than dispersal limitation in shaping the distribution of fungal endophyte communities in New Guinea forests. Our results suggest that in the absence of strong ecological gradients and host turnover, beta diversity of endophyte communities could be low in large areas of contiguous forest.     

Are tropical fungal endophytes hyperdiverse?

Fungal endophytes are ubiquitous fungi that inhabit healthy plant tissues without causing disease. Endophytes have been found in every plant species examined to date and may be important, but often overlooked, components of fungal biodiversity. In two sites in a lowland, moist tropical forest of central Panama, we quantified endophyte colonization patterns, richness, host preference, and spatial variation in healthy leaves of two co-occurring, understory tree species [ Heisteria concinna (Olacaceae) and Ouratea lucens (Ochnaceae)]. From 83 leaves, all of which were colonized by endophytes, we isolated 418 endophyte morphospecies (estimated 347 genetically distinct taxa), most of which were represented by only a single isolate (59%). Among morphospecies encountered in more than one leaf (nonsingletons), we found evidence of host preference and spatial heterogeneity using both morphospecies frequencies and presence/absence records. Based on these data, we postulate that tropical endophytes themselves may be hyperdiverse and suggest that extrapolative estimates that exclude them will markedly underestimate fungal species diversity.     

Arbuscular mycorrhizal communities in tropical forests are affected by host tree species and environment

Arbuscular mycorrhizal (AM) fungi are mutualists with plant roots that are proposed to enhance plant community diversity. Models indicate that AM fungal communities could maintain plant diversity in forests if functionally different communities are spatially separated. In this study we assess the spatial and temporal distribution of the AM fungal community in a wet tropical rainforest in Costa Rica. We test whether distinct fungal communities correlate with variation in tree life history characteristics, with host tree species, and the relative importance of soil type, seasonality and rainfall. Host tree species differ in their associated AM fungal communities, but differences in the AM community between hosts could not be generalized over life history groupings of hosts. Changes in the relative abundance of a few common AM fungal species were the cause of differences in AM fungal communities for different host tree species instead of differences in the presence and absence of AM fungal species. Thus, AM fungal communities are spatially distinguishable in the forest, even though all species are widespread. Soil fertility ranging between 5 and 9 Mg/ha phosphorus did not affect composition of AM fungal communities, although sporulation was more abundant in lower fertility soils. Sampling soils over seasons revealed that some AM fungal species sporulate profusely in the dry season compared to the rainy season. On one host tree species sampled at two sites with vastly different rainfall, relative abundance of spores from Acaulospora was lower and that of Glomus was relatively higher at the site with lower and more seasonal rainfall.     

Odocoileus virginianus diet (Artiodactyla: Cervidae) in a temperate forest of Northern Oaxaca, Mexico

The Sierra Madre de Oaxaca region, located in the Northern state of Oaxaca, Mexico, is an area of forest ecosystems subject to high exploitation rates, although in some areas its temperate forests are conserved by indigenous community initiatives that live there. We analyzed the diet of white tailed-deer (Odocoileus virginianus) in the localities of Santa Catarina Lachatao and San Miguel Amatlán from June 1998 to August 1999. Sampling was done during both the wet and dry seasons, and included the observation of browsing traces (238 observations), microhistological analysis of deer feces (28 deer pellet-groups), and two stomach content analysis. The annual diet of white-tailed deer was composed of 42 species from 23 botanical families. The most represented families in the diet of this deer were Fagaceae, Asteraceae, Ericaceae and Fabaceae. There were significant differences in the alpha diversity of the diet during the wet and dry seasons (H'=2.957 and H'=1.832, respectively). The similarity percentage between seasons was 56%. Differences in plant species frequency were significantly higher during the wet season. Herbaceous plants made up the greatest percentage of all the species consumed. The preferred species throughout the year were Senecio sp. (shrub), Sedum dendroideum (herbaceous), Arctostaphylos pungens (shrub) and Satureja macrostema (shrub). Diet species richness was found to be lower than that observed in a tropical forest (Venezuela), tropical dry forest (Mexico) and temperate deciduous and mixed forest (Mexico), but similar to the diet species richness observed in a tropical dry forest (Costa Rica) and temperate coniferous and deciduous forests (USA).     

Spatial patterns of fungal endophytes in a subtropical montane rainforest of northern Taiwan

Fungal endophytes of plants are ubiquitous and important to host plant health. Wood-inhabiting and foliar endophyte communities from multiple tree hosts were sampled at multiple spatial scales across the Fushan forest dynamics plot in northern Taiwan, using culture-free, community DNA amplicon sequencing methods. Fungal endophyte communities were distinct between leaves and wood, but the mycobiomes were highly variable across and within tree species. Despite this, host tree species was an important predictor of mycobiome community-composition. Within a single common tree species, “core” mycobiomes were characterized using co-occurrence analysis. The spatial co-occurrence patterns of these few species of fungal endophytes appear to explain the strong host effect. For wood endophytes, a consistent core mycobiome coexisted with the host across the extent of the study. For leaf endophytes, the core fungi resembled a more dynamic, “gradient” model of the core microbiome, changing across the topography and distance of the study.     

Divergent rainforest tree microbiomes between phases of the monsoon cycle,host plants and tissues

• The Australian Monsoon Tropics (AMT) contain some of the most biodiverse forests on the continent. Little is known about the dynamics of rainforest plant microbiomes in general, and there have been no community-level studies on Australian rainforest endophytes, their seasonality, tissue and host specificity.
• We tested whether community composition of tropical tree endophytes (fungi and bacteria) differs: (i) at different points during a monsoon cycle, (ii) between leaf and stem tissues, (iii) between forest microclimates (gully/ridge), and between (iv) host plant species, and (v) host plant clade, using amplicon sequencing of the bacterial 16S and fungal ITS2 gene regions.
• Results indicated that the composition of rainforest plant microbiomes differs between wet and dry seasons, which may be explained by physiological shifts in host plants due to annual climate fluctuations from mesic to xeric. Endophyte microbiomes differed between leaves and stems. Distinct fungal communities were associated with host species and clades, with some trees enriched in a number of fungal taxa compared to host plants in other clades. Diversity of bacterial endophytes in plant stems increased in the dry season.
• We conclude that the microbiomes of tropical plants are responsive to monsoonal climate variation, are highly compartmentalised between plant tissues, and may be partly shaped by the relatedness of their host plants.

     

A comparison of fungal endophytic community diversity in tree leaves of rural and urban temperate forests of Kanto district,eastern Japan

To clarify the effects of forest fragmentation and a change in tree species composition following urbanization on endophytic fungal communities, we isolated fungal endophytes from the foliage of nine tree species in suburban (Kashiwa City, Chiba) and rural (Mt. Wagakuni, Ibaraki; Mt. Takao, Tokyo) forests and compared the fungal communities between sites and host tree species. Host specificity was evaluated using the index of host specificity (Si), and the number of isolated species, total isolation frequency, and the diversity index were calculated. From just one to several host-specific species were recognized in all host tree species at all sites. The total isolation frequency of all fungal species on Quercus myrsinaefolia, Quercus serrata, and Chamaecyparis obtusa and the total isolation frequency of host-specific species on Q. myrsinaefolia, Q. serrata, and Eurya japonica were significantly lower in Kashiwa than in the rural forests. The similarity indices (nonmetric multidimensional scaling (NMS) and C_MH) of endophytic communities among different tree species were higher in Kashiwa, as many tree species shared the same fungal species in the suburban forest. Endophytic fungi with a broad host range were grouped into four clusters suggesting their preference for conifer/broadleaves and evergreen/deciduous trees. Forest fragmentation and isolation by urbanization have been shown to cause the decline of host-specific fungal species and a decrease in β diversity of endophytic communities, i.e., endophytic communities associated with tree leaves in suburban forests were found to be depauperate.     

Fungal endophyte communities in above- and belowground olive tree organs and the effect of season and geographic location on their structures

Studies comparing fungal endophytes between above- and belowground woody crop organs and the factors that may structure their communities are lacking. Due to its great impact on the Mediterranean Basin, the olive tree was chosen for the isolation of endophytic fungi from roots, leaves and twigs in two seasons in north-eastern Portugal. Nine hundred seventy-six isolates belonging to 38 fungal species were obtained. Phomopsis columnaris, Fusarium oxysporum and Trichoderma gamsii were the most frequently isolated, collectively representing 69% of the isolates. Fungal diversity in the roots was higher than in the aboveground organs and higher in spring than autumn. Endophyte community similarity between the above- and belowground organs and between seasons was low. Species composition also varied spatially, with the fungal composition of the roots varying more among locations than that of the aboveground organs. Our results suggest olive tree endophyte community structure is affected by plant organ, location and season.     

High butterfly beta diversity between Brazilian cerrado and cerrado–caatinga transition zones

Tropical dry forests are among the most diverse and threatened habitats in the world, yet they are rarely protected and remain poorly studied. In Brazil, dry forests are naturally fragmented and embedded within various biomes, thus making it important to assess biotic homogeneity among geographically separated forest fragments. We sampled 7732 individuals belonging to 48 species to quantify the diversity of fruit-feeding butterfly communities at four Brazilian dry forest sites, and found differences in community structure between northern and central sites. Species richness per plot was the same in both areas, but abundance per plot was higher in northern sites. Species composition differed between sites mostly due to species of Satyrinae. Additive partitioning showed that beta diversity corresponded to 70.1% of all diversity. Rather than species loss, beta diversity primarily represented species turnover that was potentially driven by differences in the surrounding habitats. Butterfly community composition and abundance were influenced by vegetation where abundance increased with tree density and basal area, and decreased with the average tree height. Butterfly species richness and abundance were higher in the wet season than in the dry season, and all species sampled in the dry season were present in the wet season. Differences in community structure across relatively short geographic distances in the same type of habitat highlight the importance of conserving tropical dry forest fragments to ensure the maintenance of butterfly diversity and, presumably, other insects.     

Epiphytic orchids in tropical dry forests of Yucatan,Mexico – Species occurrence,abundance and correlations with host tree characteristics and environmental conditions

Tropical dry forests have been less studied in terms of their resident epiphyte flora compared to wet forests. We studied five species of epiphytic orchids in two dry forest fragments differing in tree composition, stature and rainfall regime. We compared the vertical distribution within the host tree, epiphyte-host associations and seasonal variation in microclimatic conditions in a tropical dry deciduous (Celestún) and a semi-deciduous forest (Kaxil-Kiuic) of the Yucatan Peninsula, Mexico, during the wet, early dry and dry seasons. Light, vapor pressure deficit, air temperature, and dew were measured on two heights (1.5 and 3.5 m) of the host with the highest abundance of orchids. Surprisingly, orchid abundance was higher in the Celestún deciduous forest, the site with low precipitation. High epiphyte abundance in the middle canopy stratum of the hosts in both forests was arguably related to a favorable combination of micro-environmental factors. In both forests, about 90% of all orchids grew on a single host tree species. Although bark roughness and the area of the substrate were the most important host characteristics that influenced the abundance of orchids in the Celestún deciduous forest, this did not explain this preference. Climatic variation was greater among seasons than between microenvironments in the host trees. The most abundant species, Encyclia nematocaulon, had a great capacity to occupy different strata in both forests, even in tree tops with very large micro-environmental fluctuations during the year.     

Abundance and Leaf Damage Caused by Gall‐Inducing Insects in a Mexican Tropical Dry Forest1

We evaluated the temporal and spatial patterns of abundance and the amount of damage caused by gall‐inducing insects (GII) in deciduous and riparian habitats in a seasonal tropical dry forest in Mexico. Plants occurring in these habitats differ in their phenology and moisture availability. Deciduous habitats are seasonal and xeric, while riparian habitats are aseasonal and mesic. We found 37 GII species and each one was associated with a specific plant species. In total, 19 species (51.3%) were present in deciduous habitats, 13 species (35.2%) in riparian habitats, and only 5 species (13.5%) occurred in both. Abundance and leaf damage by GII were greater in deciduous than in riparian habitats during the wet season. For each GII species that occurred in both habitats, host plant species supported greater abundance and leaf damage by GII in deciduous habitats during the wet season. These results indicate a greater association of GII species with host plants in deciduous than in riparian habitats during the wet season. In riparian habitats, 11 plant species (61.1%) had greater density of GII in the dry than in the wet season. Similarly, leaf damage by GII was significantly greater in the dry than in the wet season in riparian habitats for 12 plant species (66.7%). Dry forest plants of riparian habitats presented two peaks of leaf‐flushing: GII colonized leaves produced in the first peak at the beginning of the wet season, and accumulated or recolonized leaves in the second peak at the beginning of the dry season. The levels of leaf damage by GII detected in this study in the rainy season were considerably higher than those obtained for folivorous insects in other neotropical forests, suggesting that this GII guild might have an important impact on their host plant species in this tropical community.     

The incidence and implications of clouds for cloud forest plant water relations

Although clouds are the most recognisable and defining feature of tropical montane cloud forests, little research has focussed on how clouds affect plant functioning. We used satellite and ground‐based observations to study cloud and leaf wetting patterns in contrasting tropical montane and pre‐montane cloud forests. We then studied the consequences of leaf wetting for the direct uptake of water accumulated on leaf surfaces into the leaves themselves. During the dry season, the montane forest experienced higher precipitation, cloud cover and leaf wetting events of longer duration than the pre‐montane forest. Leaf wetting events resulted in foliar water uptake in all species studied. The capacity for foliar water uptake differed significantly between the montane and pre‐montane forest plant communities, as well as among species within a forest. Our results indicate that foliar water uptake is common in these forest plants and improves plant water status during the dry season.     

Ecology and diversity of leaf litter fungi during early-stage decomposition in a seasonally dry tropical forest

Leaf litter samples of 12 dicotyledonous tree species (belonging to eight families) growing in a dry tropical forest and in early stages of decomposition were studied for the presence of litter fungi. Equal-sized segments of the leaves incubated in moist chambers were observed every day for 30 d for the presence of fungi. Invariably, the fungal assemblage on the litter of each tree species was dominated by a given fungal species. The diversity of fungi present in the litter varied with the tree species although many species of fungi occurred in the litter of all 12 species. A Pestalotiopsis species dominated the litter fungal assemblage of five trees and was common in the litter of all tree species. The present study and earlier studies from our lab indicate that fungi have evolved traits such as thermotolerant spores, ability to utilize toxic furaldehydes, ability to produce cell wall destructuring enzymes and an endophyte-litter fungus life style to survive and establish themselves in fire-prone forests such as the one studied here. This study shows that in the dry tropical forest, the leaf litter fungal assemblage is governed more by the environment than by the plant species.     

Disturbance and Seasonal Dynamics of Mycorrhizae in a Tropical Deciduous Forest in Mexico1

Mycorrhizal fungi were sampled in a deciduous tropical forest on the Pacific coast of Mexico during different seasons and in natural treefall gaps and pastures. All 12 plant species sampled in the forest were arbuscular mycorrhizal. The percent root infection and spore production were closely related to the phenology of the plants. Most tree species and all herbaceous species had the highest infection in the summer rainy season, but two species, Opuntia excelsa and Jacquinia pungens, had highest infection in the dry season. Unusually high rainfall during the dry season was associated with increased infection but not increased spore production. Spore density was low for all species at all sample times, except at the beginning of the July 1993 rainy season in, when we observed up to 28 spores/g soil. The percent cover of shrubs or herbs did not increase in gaps after two years, and we observed no colonizing seedlings. No plant species with cover higher than 2.7 percent occurred exclusively in gaps or forest. The percent mycorrhizal infection did not differ significantly between gaps and forest. Spore counts were as high in the gaps as in the forest in two of the three gaps but lower in the third gap. The lack of significant response of plants in these gaps after two years differed from the rapid response in tropical rainforests. It is likely related to the small size of the gaps and to light infiltration to the forest floor. Pastures were dominated by two species of exotic grasses and one species of mycorrhizal fungus, whereas forests had 15 fungal species. The slow regrowth of vegetation in gaps was not limited by mycorrhizal fungi, since they were still abundant after the treefalls, but recovery in pastures could be affected by low fungal diversity and dominance of grasses.     

Diversity and community composition of ectomycorrhizal fungi in a dry deciduous dipterocarp forest in Thailand

Large forest areas of South-East Asia, are dominated by the Dipterocarpaceae tree family, which contains many important timber species. Unlike many other tropical trees, Dipterocarpaceae rely on ectomycorrhizal (ECM) root symbiosis for their mineral nutrition. This study aims to document the richness and community composition of ECM fungi in a dry deciduous forest in Thailand. Combining morphological and molecular identification methods revealed 69 species of ECM fungi that belong to 17 phylogenetic lineages. The /russula-lactarius, /tomentella-thelephora, /sordariales, /sebacina and /cantharellus lineages were the most species-rich. The fungal richness is comparable to other tropical rain forest sites, but the phylogenetic community structure has elements of both tropical and temperate ecosystems. Unlike tropical rain forests, the Cenococcum geophilum complex was one of the most frequent fungal taxa that had a relatively high ITS genetic diversity over the small sampling area. This study provides the first snapshot insight into the fungal community of dry dipterocarp forests. However, it is necessary to broaden the spatial and temporal scales of sampling to improve our understanding of the below-ground relations of dry and humid tropical forests.     

Bird communities of dry forests and oak woodland of western Mexico

Bird species-richness and densities were studied in spring and autumn of 1991 and 1992 in dry forest and oak woodland in western Mexico. Classification of the vegetation resulted in the identification of six dry forest types: from thorn forests, dominated by Acacia , through broad-leaved tree forest (typical dry forest of western Mexico) and small-leaved tree forest to Ipomoea forest. Oak woodland and mixed woodland, which are a transition between dry forest and the more mesic oak woodland, formed two further forest types. In the study area as a whole, bird densities were higher in autumn, following the rainy season, than in spring, which is the dry season, and the beginning of the breeding season. Densities in the two autumns were very similar (mean 93 ± 7.8 birds per 2.8 ha) but differed significantly between spring 1991 (57 ± 2.2 per 2.8 ha) and spring 1992 (70 ± 3.2 per 2.8 ha), possibly due to unseasonal rain in 1992. In autumn, bird densities were highest in thorn forests and lowest in broad-leaved tree forests and oak woodland. In spring 1991, bird densities were highest in Prosopis thorn forest and mixed woodland, and lowest in oak woodland. Bird species-richness was low in oak woodland, in both seasons, and high in mixed woodland and Ipomoea forest in spring. Bird species composition between forest types was more similar in the spring than autumn. The spring similarity in species composition is explained by the presence of generalist species in times of food scarcity during the dry season, and by the homogeneity of the predominantly leafless vegetation. Comparison with the bird species composition of a dry forest in Venezuela and the Pacific coast of Mexico indicates that the sites in the present study have an intermediate number of species. Although the present study recorded a smaller number of bird species than on the Pacific coast, 16 were different, expanding the total for Mexican dry forest by 14% from 118 to 134.     

Phyllostomid Bat Community Structure and Abundance in Two Contrasting Tropical Dry Forests1

Although tropical wet forests are generally more diverse than dry forests for many faunal groups, few studies have compared bat diversity among dry forests. I compared ground level phyllostomid bat community structure between two tropical dry forests with different precipitation regimes. Parque National Palo Verde in northwestern Costa Rica represents one of the wettest tropical dry forests (rainfall 1.5 m/yr), whereas the Chamela‐Cuixmala Biosphere Reserve on the Pacific coast of central Mexico represents one of the driest (750 mm/yr). Mist net sampling was conducted at the two study sites to compare changes in ground level phyllostomid bat community structure between regions and seasons. Palo Verde was more diverse than Chamela and phyllostomid species showed low similarity between sites (Classic Jaccard = 0.263). The distinct phyllostomid communities observed at these two dry forest sites demonstrates that variants of tropical dry forest can be sufficiently different in structure and composition to affect phyllostomid communities. At both dry forest sites, abundance of the two most common foraging guilds (frugivores and nectarivores) differed between seasons, with greatest numbers of individuals captured coinciding with highest chiropterophilic resource abundance.     

Wet- vs. Dry-Season Transpiration in an Amazonian Rain Forest Palm Iriartea deltoidea

The wet and dry seasons in tropical rain forests can differ in precipitation, soil moisture and irradiance more significantly than often assumed. This could potentially affect the water relations of many tree species that may exhibit either increased transpiration in the dry season as a response to the increased irradiance or decreased transpiration as a result of decreases in soil moisture and increases in atmospheric vapor pressure deficit (VPD). Atmospheric data, soil moisture data and sap fluxes in Iriartea deltoidea palms were measured in eastern Ecuador during the wet and dry seasons. There were no differences between total daily sap fluxes in I. deltoidea palms during the wet and dry seasons; however, evaporative demand was significantly higher in the dry season and therefore, transpiration was more restricted by stomatal closure during the dry season than the wet season. This is likely the result of larger atmospheric VPD during the dry season compared with the wet season and possibly the result of reduced soil moisture availability. Additionally, based on published tree abundances in this area, measured sap fluxes in I. deltoidea were scaled up to the hectare level. Transpiration from I. deltoidea palms was estimated to be around 0.03 mm/d, which could represent about 1 percent of total transpiration in this area of the Amazon rain forest. If climate change predictions for more lengthy tropical dry periods are realized, greater stomatal control of dry-season sap flux has the potential to become even more prevalent in tropical species.