Differential effects of tropical arbuscular mycorrhizal fungal inocula on root colonization and tree seedling growth: implications for tropical forest diversity

The potential for mycorrhizae to influence the diversity and structuring of plant communities depends on whether their affinities and effects differ across a suite of potential host species. In order to assess this potential for a tropical forest community in Panama, we conducted three reciprocal inoculation experiments using seedlings from six native tree species. Seeds were germinated in sterile soil and then exposed to arbuscular mycorrhizal fungi in current association with naturally infected roots from adults of either the same or different species growing in intact forest. The tree species represent a range of life histories, including early successional pioneers, a persistent understory species, and emergent species, typical of mature forest. Collectively, these experiments show: (i) the seedlings of small-seeded pioneer species were more dependent on mycorrhizal inocula for initial survival and growth; (ii) although mycorrhizal fungi from all inocula were able to colonize the roots of all host species, the inoculum potential (the infectivity of an inoculum of a given concentration) and root colonization varied depending on the identity of the host seedling and the source of the inoculum; and (iii) different mycorrhizal fungal inocula also produced differences in growth depending on the host species. These differences indicate that host–mycorrhizal fungal interactions in tropical forests are characterized by greater complexity than has previously been demonstrated, and suggest that tropical mycorrhizal fungal communities have the potential to differentially influence seedling recruitment among host species and thereby affect community composition.     

林火对植物根围丛枝菌根真菌多样性的影响

林火是森林生态系统的一种主要干扰因子,以青岛市三标山林火迹地为研究对象,采集荆条(Vitex negundo)、胡枝子(Lespedeza bicolor)、花木蓝(Indigofera kirilowii)、青花椒(Zanthoxylum schinifolium)和野青茅(Deyeuxia arundinacea)5种优势植物根围土壤,研究不同林火强度对丛枝菌根(AM)真菌多样性的影响。结果表明,AM真菌侵染率和孢子密度随火灾强度的加强而降低;非过火区植物根围土壤中,分离鉴定出AM真菌3属11种,轻度过火区分离鉴定出AM真菌3属10种,中度过火区分离鉴定出AM真菌3属9种,重度过火区分离鉴定出AM真菌3属8种。过火区AM真菌种丰度低于非过火区。过火区和非过火区AM真菌的重要值和优势种不同,非过火区植物根围的优势种是地球囊霉(Glomus geosporum)、台湾球囊霉(G.taiwanensis)、分支巨孢囊霉(Gigaspora ramisporophora)、极大巨孢囊霉(Gi.gigantean)、福摩萨球囊霉(G.formosanum)、悬钩子球囊霉(G.rubiforme)、柯氏无梗囊霉(Acaulospora koskei)和松蜜无梗囊霉(A.thomii);轻度过火区植物根围的优势种是地球囊霉和台湾球囊霉;中度过火区的是台湾球囊霉和地球囊霉(野青茅除外);重度过火区植物根围的优势种是地球囊霉。不同强度的过火区对AM真菌群落组成有不同程度的影响。认为林火降低植物根围土壤中AM真菌多样性。     

Terrestrial vertebrates promote arbuscular mycorrhizal fungal diversity and inoculum potential in a rain forest soil

We examined whether terrestrial vertebrates affected the arbuscular mycorrhizal fungal spore communities and mycorrhizal inoculum potential (MIP) of a tropical rain forest soil by comparing plots where terrestrial vertebrates had been excluded for 3 years to adjacent control plots. We extracted spores from soil using sucrose density gradient centrifugation and assayed MIP by growing seedlings of maize ( Zea mays ) and a rain forest tree ( Flindersia brayleana ) in intact soil cores from exclosure and control plots. Control plots had significantly higher spore abundance, species richness and diversity than exclosures. Spore community composition also differed significantly between exclosure and control plots. Seedlings of both plant species grown in control cores had significantly higher arbuscular-mycorrhizal colonization than those grown in exclosure cores. This study suggests that loss of vertebrates could alter rates of mycorrhizal colonization with consequences for community and ecosystem properties.     

Environmental filtering structures fungal endophyte communities in tree bark

The factors that control the assembly and composition of endophyte communities across plant hosts remains poorly understood. This is especially true for endophyte communities inhabiting inner tree bark, one of the least studied components of the plant microbiome. Here, we test the hypothesis that bark of different tree species acts as an environmental filter structuring endophyte communities, as well as the alternative hypothesis, that bark acts as a passive reservoir that accumulates a diverse assemblage of spores and latent fungal life stages. We develop a means of extracting high‐quality DNA from surface sterilized tree bark to compile the first culture‐independent study of inner bark fungal communities. We sampled a total of 120 trees, spanning five dominant overstorey species across multiple sites in a mixed temperate hardwood forest. We find that each of the five tree species harbour unique assemblages of inner bark fungi and that angiosperm and gymnosperm hosts harbour significantly different fungal communities. Chemical components of tree bark (pH, total phenolic content) structure some of the differences detected among fungal communities residing in particular tree species. Inner bark fungal communities were highly diverse (mean of 117–171 operational taxonomic units per tree) and dominated by a range of Ascomycete fungi living asymptomatically as putative endophytes. Together, our evidence supports the hypothesis that tree bark acts as an environmental filter structuring inner bark fungal communities. The role of these potentially ubiquitous and plant‐specific fungal communities remains uncertain and merits further study.     

西双版纳热带次生林中的丛枝菌根调查

对西双版纳热带次生林中13个科的26种植物根系的丛枝菌根真菌（Arbuscular mycorrhizal fongii,AMF）侵染情况进行了研究,并从这些植物的根围土壤中分离鉴定了隶属于球囊霉属（Clomus）、巨孢囊霉属（Gigaspora）、盾巨孢囊霉属（Scutellospora）和无梗囊霉属（Acaulospora）的11种丛枝菌根真菌。该地次生林中AMF的孢子密度为13—29个／100g土壤,平均为19个;种的丰富度在4．9之间（平均为6）;平均频度为53．8％;相对多度为3．2％-26．5％;物种多样性指数和均匀度指数分别为0．94和0．93。丛枝菌根的侵染率达到44．8％．57．2％（平均为50．9％）;球囊霉属（Glomus）和无梗囊霉属（Acattlospora）是热带次生林根围土壤中菌根真菌的优势类群。     

Molecular diversity of arbuscular mycorrhizal fungi and patterns of host association over time and space in a tropical forest

We have used molecular techniques to investigate the diversity and distribution of the arbuscular mycorrhizal (AM) fungi colonizing tree seedling roots in the tropical forest on Barro Colorado Island (BCI), Republic of Panama. In the first year, we sampled newly emergent seedlings of the understory treelet Faramea occidentalis and the canopy emergent Tetragastris panamensis, from mixed seedling carpets at each of two sites. The following year we sampled surviving seedlings from these cohorts. The roots of 48 plants were analysed using AM fungal-specific primers to amplify and clone partial small subunit (SSU) ribosomal RNA gene sequences. Over 1300 clones were screened for random fragment length polymorphism (RFLP) variation and 7% of these were sequenced. Compared with AM fungal communities sampled from temperate habitats using the same method, the overall diversity was high, with a total of 30 AM fungal types identified. Seventeen of these types have not been recorded previously, with the remainder being similar to types reported from temperate habitats. The tropical mycorrhizal population showed significant spatial heterogeneity and nonrandom associations with the different hosts. Moreover there was a strong shift in the mycorrhizal communities over time. AM fungal types that were dominant in the newly germinated seedlings were almost entirely replaced by previously rare types in the surviving seedlings the following year. The high diversity and huge variation detected across time points, sites and hosts, implies that the AM fungal types are ecologically distinct and thus may have the potential to influence recruitment and host composition in tropical forests.     

Intra‐annual plasticity of growth mediates drought resilience over multiple years in tropical seedling communities

Precipitation patterns are changing across the globe causing more severe and frequent drought for many forest ecosystems. Although research has focused on the resistance of tree populations and communities to these novel precipitation regimes, resilience of forests is also contingent on recovery following drought, which remains poorly understood, especially in aseasonal tropical forests. We used rainfall exclusion shelters to manipulate the interannual frequency of drought for diverse seedling communities in a tropical forest and assessed resistance, recovery and resilience of seedling growth and mortality relative to everwet conditions. We found seedlings exposed to recurrent periods of drought altered their growth rates throughout the year relative to seedlings in everwet conditions. During drought periods, seedlings grew slower than seedlings in everwet conditions (i.e., resistance phase) while compensating with faster growth after drought (i.e., recovery phase). However, the response to frequent drought was species dependent as some species grew significantly slower with frequent drought relative to everwet conditions while others grew faster with frequent drought due to overcompensating growth during the recovery phase. In contrast, mortality was unrelated to rainfall conditions and instead correlated with differences in light. Intra‐annual plasticity of growth and increased annual growth of some species led to an overall maintenance of growth rates of tropical seedling communities in response to more frequent drought. These results suggest these communities can potentially adapt to predicted climate change scenarios and that plasticity in the growth of species, and not solely changes in mortality rates among species, may contribute to shifts in community composition under drought.     

The ecology of predispersal insect herbivory on tree reproductive structures in natural forest ecosystems

Plant-insect interactions are key model systems to assess how some species affect the distribution, the abundance, and the evolution of others. Tree reproductive structures represent a critical resource for many insect species, which can be likely drivers of demography, spatial distribution, and trait diversification of plants. In this review, we present the ecological implications of predispersal herbivory on tree reproductive structures by insects (PIHR) in forest ecosystems. Both insect's and tree's perspectives are addressed with an emphasis on how spatiotemporal variation and unpredictability in seed availability can shape such particular plant-animal interactions. Reproductive structure insects show strong trophic specialization and guild diversification. Insects evolved host selection and spatiotemporal dispersal strategies in response to variable and unpredictable abundance of reproductive structures in both space and time. If PIHR patterns have been well documented in numerous systems, evidences of the subsequent demographic and evolutionary impacts on tree populations are still constrained by time-scale challenges of experimenting on such long-lived organisms, and modeling approaches of tree dynamics rarely consider PIHR when including biotic interactions in their processes. We suggest that spatially explicit and mechanistic approaches of the interactions between individual tree fecundity and in sect dynamics will clarify predictions of the demogenetic implications of PIHR in tree populations. In a global change context, further experimental and theoretical contributions to the likelihood of life-cycle disruptions between plants and their specialized herbivores, and to how these changes may gen erate novel dynamic patterns in each partner of the interaction are increasingly critical.     

Biochar soil amendments in prairie restorations do not interfere with benefits from inoculation with native arbuscular mycorrhizal fungi

Little of the historical extent of tallgrass prairie ecosystems remains in North America, and therefore there is strong interest in restoring prairies. However, slow‐growing prairie plants are initially weak competitors with the fast‐growing yet short‐lived weedy plant species that are typically abundant in recently established prairie restorations. One way to aid establishment of slow‐growing plant species is through adding soil amendments to prairie restorations before planting. Arbuscular mycorrhizal (AM) fungi form mutualisms with the roots of most terrestrial plants and are particularly important for the growth of slow‐growing prairie plant species. As prairie ecosystems are adapted to fires that leave biochar (charred organic material) in the soil, adding biochar as well as AM fungal strains from undisturbed remnant prairies into the soil of prairie restorations may improve restoration outcomes. Here, we test this prediction during the first four growing seasons of a prairie restoration. When prairie plant seedlings were inoculated prior to planting into the field with AM fungi derived from remnant prairies, that one‐time inoculation significantly increased growth of five of the nine tested plant species through at least two growing seasons. This long‐term benefit of AM fungal inoculation was unaffected by biochar addition to the soil. Biochar application rates of at least 10 tons/ha significantly decreased Coreopsis tripteris growth but acted synergistically with AM fungal inoculation to significantly improve survival of Schizachyrium scoparium. Overall, inoculation with native AM fungi can help promote prairie plant establishment, but concomitant use of biochar soil amendments had relatively little effect.     

Facilitation between woody and herbaceous plants that associate with arbuscular mycorrhizal fungi in temperate European forests

In late‐successional environments, low in available nutrient such as the forest understory, herbaceous plant individuals depend strongly on their mycorrhizal associates for survival. We tested whether in temperate European forests arbuscular mycorrhizal (AM) woody plants might facilitate the establishment of AM herbaceous plants in agreement with the mycorrhizal mediation hypothesis. We used a dataset spanning over 400 vegetation plots in the Weser‐Elbe region (northwest Germany). Mycorrhizal status information was obtained from published resources, and Ellenberg indicator values were used to infer environmental data. We carried out tests for both relative richness and relative abundance of herbaceous plants. We found that the subset of herbaceous individuals that associated with AM profited when there was a high cover of AM woody plants. These relationships were retained when we accounted for environmental filtering effects using path analysis. Our findings build on the existing literature highlighting the prominent role of mycorrhiza as a coexistence mechanism in plant communities. From a nature conservation point of view, it may be possible to promote functional diversity in the forest understory through introducing AM woody trees in stands when absent.     

Increased Productivity and Reduced Seed Predation Favor a Large‐seeded Palm in Small Atlantic Forest Fragments

Large‐seeded plants are especially vulnerable to the loss of seed dispersers in small forest fragments. The palm Attalea humilis goes against this trend by reaching high abundances in small remnants. Productivity, seed dispersal and seed predation of A. humilis were investigated in two large (2400 and 3500 ha) and three small (19, 26 and 57 ha) Atlantic Forest fragments in southeastern Brazil. Palms in the small fragments produced more female inflorescences, resulting in a higher fruit production in these places. Seed dispersal rates were higher in the large fragments, where scatter hoarding was more frequent. Scolytine beetles were the main seed predators and damaged a larger number of seeds in small fragments, but predation by rodents and bruchine beetles was low irrespective of fragment size. As scolytines do not necessarily kill the seeds, low predation by bruchines and rodents, together with its own high productivity, allow A. humilis to be more abundant in small fragments despite the scarcity of its main dispersers. This increased abundance, by its turn, can increase competitive interactions between A. humilis and other plants in small fragments. Thus, abundance patterns of A. humilis are a good example of fragmentation affecting the balance of ecological interactions in a complex way, emphasizing the role of preserving ecological processes for conserving biodiversity in fragmented tropical landscapes. Abstract in Portuguese is available at http://www.blackwell‐synergy.com/loi/btp .     

海南霸王岭热带雨林常见植物丛枝菌根真菌调查

对海南霸王岭热带雨林的12科16种常见植物的丛枝菌根状况进行了调查,用碱解离-酸性品红染色法进行了真菌鉴定。结果表明,13种植物形成典型的丛枝菌根,占所调查植物的81%;3种植物没有形成丛枝菌根,占所调查植物的19%。用湿筛沉淀法从这些植物根际土壤中共分离鉴定出了3属11种丛枝菌根真菌(AMF),即无梗囊霉属(Acaulos-pora)3种,球囊霉属(Glomus)7种,巨孢囊霉属(Gigaspora)1种;其中,球囊霉属是样地的优势属。在AMF中,孔窝无梗囊霉(A.foveata)分离频率最高,在14种植物的根际土中都有发现;此外,大果球囊霉(G.macrocarpum)的相对多度最大,为59%,具有最强的产孢能力。同时,在11种植物的根中发现了深色有隔内生真菌(DSE),占调查植物的69%;其中,11种植物同时被DSE和AMF感染。     

Drivers of aboveground wood production in a lowland tropical forest of West Africa: teasing apart the roles of tree density,tree diversity,soil phosphorus,and historical logging

Tropical forests currently play a key role in regulating the terrestrial carbon cycle and abating climate change by storing carbon in wood. However, there remains considerable uncertainty as to whether tropical forests will continue to act as carbon sinks in the face of increased pressure from expanding human activities. Consequently, understanding what drives productivity in tropical forests is critical. We used permanent forest plot data from the Gola Rainforest National Park (Sierra Leone) – one of the largest tracts of intact tropical moist forest in West Africa – to explore how (1) stand basal area and tree diversity, (2) past disturbance associated with past logging, and (3) underlying soil nutrient gradients interact to determine rates of aboveground wood production (AWP). We started by statistically modeling the diameter growth of individual trees and used these models to estimate AWP for 142 permanent forest plots. We then used structural equation modeling to explore the direct and indirect pathways which shape rates of AWP. Across the plot network, stand basal area emerged as the strongest determinant of AWP, with densely packed stands exhibiting the fastest rates of AWP. In addition to stand packing density, both tree diversity and soil phosphorus content were also positively related to productivity. By contrast, historical logging activities negatively impacted AWP through the removal of large trees, which contributed disproportionately to productivity. Understanding what determines variation in wood production across tropical forest landscapes requires accounting for multiple interacting drivers – with stand structure, tree diversity, and soil nutrients all playing a key role. Importantly, our results also indicate that logging activities can have a long‐lasting impact on a forest's ability to sequester and store carbon, emphasizing the importance of safeguarding old‐growth tropical forests.     

Woody plant diversity in relation to environmental factors in a seasonally dry tropical forest landscape

Questions

Water availability is known to be a first‐order driver of plant diversity; yet water also affects fire regimes and soil fertility, which, in turn, affect plant diversity. We examined how precipitation, fire and soil properties jointly determine woody plant diversity. Specifically, we asked how woody plant diversity varies along a sharp precipitation gradient (about 600–1,800 mm mean annual precipitation [MAP ]within a ~45‐km distance) exhibiting considerable variation in long‐term fire burn frequency and soil fertility, in a southern Indian seasonally dry tropical forest (SDTF ) landscape.

Location

Mudumalai, Western Ghats, India.

Methods

Woody plants ≥1‐cm DBH were enumerated in 19 1‐ha permanent plots spanning a range of tropical vegetation types from dry thorn forest, through dry and moist deciduous forest to semi‐evergreen forest. Burn frequencies were derived from annual fire maps. Six measures of surface soil properties – total exchangeable bases (Ca + Mg + K), organic carbon (OC ), total N, pH , plant available P and micronutrients (Fe + Cu + Zn + Mn) were used in the analyses. Five measures of diversity – species richness, Shannon diversity, the rarefied/extrapolated versions of these two measures, and Fisher's α – were modelled as functions of MAP , annual fire burn frequency and the principal components of soil properties.

Results

Most soil nutrients and OC increased with MAP , except in the wettest sites. Woody productivity increased with MAP , while fire frequency was highest at intermediate values of MAP . Woody plant diversity increased with MAP but decreased with increasing fire frequency, resulting in two local diversity maxima along the MAP gradient – in the semi‐evergreen and dry thorn forest – separated by a low‐diversity central region in dry deciduous forest where fire frequency was highest. Soil variables were, on the whole, less strongly correlated with diversity than MAP .

Conclusions

Although woody plant diversity in this landscape, representative of regional SDTF s, is primarily limited by water availability, our study emphasizes the role of fire as a potentially important second‐order driver that acts to reduce diversity in this landscape.

     

Common spatial patterns of trees in various tropical forests: Small trees are associated with increased diversity at small spatial scales

Tropical forests are notable for their high species diversity, even on small spatial scales, and right‐skewed species and size abundance distributions. The role of individual species as drivers of the spatial organization of diversity in these forests has been explained by several hypotheses and processes, for example, stochastic dilution, negative density dependence, or gap dynamics. These processes leave a signature in spatial distribution of small trees, particularly in the vicinity of large trees, likely having stronger effects on their neighbors. We are exploring species diversity patterns within the framework of various diversity‐generating hypotheses using individual species–area relationships. We used the data from three tropical forest plots (Wanang—Papua New Guinea, Barro Colorado Island—Panama, and Sinharaja—Sri Lanka) and included also the saplings (DBH ≥ 1 cm). Resulting cross‐size patterns of species richness and evenness reflect the dynamics of saplings affected by the distribution of large trees. When all individuals with DBH ≥1 cm are included, ~50% of all tree species from the 25‐ or 50‐ha plot can be found within 35 m radius of an individual tree. For all trees, 72%–78% of species were identified as species richness accumulators, having more species present in their surroundings than expected by null models. This pattern was driven by small trees as the analysis of DBH >10 cm trees showed much lower proportion of accumulators, 14%–65% of species identified as richness repellers and had low richness of surrounding small trees. Only 11%–26% of species had lower species evenness than was expected by null models. High proportions of species richness accumulators were probably due to gap dynamics and support Janzen–Connell hypothesis driven by competition or top‐down control by pathogens and herbivores. Observed species diversity patterns show the importance of including small tree size classes in analyses of the spatial organization of diversity.     

Soil fungal communities of grasslands are environmentally structured at a regional scale in the Alps

Studying patterns of species distributions along elevation gradients is frequently used to identify the primary factors that determine the distribution, diversity and assembly of species. However, despite their crucial role in ecosystem functioning, our understanding of the distribution of below‐ground fungi is still limited, calling for more comprehensive studies of fungal biogeography along environmental gradients at various scales (from regional to global). Here, we investigated the richness of taxa of soil fungi and their phylogenetic diversity across a wide range of grassland types along a 2800 m elevation gradient at a large number of sites (213), stratified across a region of the Western Swiss Alps (700 km²). We used 454 pyrosequencing to obtain fungal sequences that were clustered into operational taxonomic units (OTUs). The OTU diversity–area relationship revealed uneven distribution of fungal taxa across the study area (i.e. not all taxa are everywhere) and fine‐scale spatial clustering. Fungal richness and phylogenetic diversity were found to be higher in lower temperatures and higher moisture conditions. Climatic and soil characteristics as well as plant community composition were related to OTU alpha, beta and phylogenetic diversity, with distinct fungal lineages suggesting distinct ecological tolerances. Soil fungi, thus, show lineage‐specific biogeographic patterns, even at a regional scale, and follow environmental determinism, mediated by interactions with plants.     

Sapling growth rates reveal conspecific negative density dependence in a temperate forest

Local tree species diversity is maintained in part by conspecific negative density dependence (CNDD). This pervasive mechanism occurs in a variety of forms and ecosystems, but research to date has been heavily skewed toward tree seedling survival in tropical forests. To evaluate CNDD more broadly, we investigated how sapling growth rates were affected by conspecific adult neighbors in a fully mapped 25.6 ha temperate deciduous forest. We examined growth rates as a function of the local adult tree neighborhood (via spatial autoregressive modeling) and compared the spatial positioning of faster‐growing and slower‐growing saplings with respect to adult conspecific and heterospecific trees (via bivariate point pattern analysis). In addition, to determine whether CNDD‐driven variation in growth rates leaves a corresponding spatial signal, we extended our point pattern analysis to a static, growth‐independent comparison of saplings and the next larger size class. We found that negative conspecific effects on sapling growth were most prevalent. Five of the nine species that were sufficiently abundant for analysis exhibited CNDD, while only one species showed evidence of a positive conspecific effect, and one or two species, depending on the analysis, displayed heterospecific effects. There was general agreement between the autoregressive models and the point pattern analyses based on sapling growth rates, but point pattern analyses based on single‐point‐in‐time size classes yielded results that differed markedly from the other two approaches. Our work adds to the growing body of evidence that CNDD is an important force in temperate forests, and demonstrates that this process extends to sapling growth rates. Further, our findings indicate that point pattern analyses based solely on size classes may fail to detect the process of interest (e.g., neighborhood‐driven variation in growth rates), in part due to the confounding of tree size and age.