Massive aerial roots affect growth and form of Dracaena draco

Key message

Large aerial roots grow out from the branches of injured Dracaena draco trees. They integrate with the trunk or cause the branches to break off the tree and deform it.

Abstract

Dracaena draco, the dragon tree, is an iconic monocot of the Canary Islands with a tree-like growth habit and some distinctive features that are unique in the plant kingdom. We report about the massive aerial roots in this tree. They appear on trees that are injured or under environmental stress and affect growth form and the whole life of the plant. We analysed the growth of these roots and tested our findings in experiments on plants. Clusters of these roots emerge from the bases of the lowest branches and growing down they may reach the soil. Descending along the trunk, they cling tightly to the trunk, integrate with it and contribute to its radial growth. This may explain (1) why the trunk of a mature D. draco tree looks fibrous, as if made of many individual strands, and (2) how some trees reach enormous radial dimensions. Alternately, a large, 2–5 m high, multi-segmented branch with aerial roots at its base, may break off the tree and grow on its own, as a mammoth off-cut, perhaps the largest known in the plant kingdom. This detachment would cause a significant reduction in the size of the crown and deform its original, highly organized pattern of branching. In the extreme condition this may result in the destruction of the mother plant.     

Epigenetic regulation of adaptive responses of forest tree species to the environment

Epigenetic variation is likely to contribute to the phenotypic plasticity and adaptative capacity of plant species, and may be especially important for long‐lived organisms with complex life cycles, including forest trees. Diverse environmental stresses and hybridization/polyploidization events can create reversible heritable epigenetic marks that can be transmitted to subsequent generations as a form of molecular “memory”. Epigenetic changes might also contribute to the ability of plants to colonize or persist in variable environments. In this review, we provide an overview of recent data on epigenetic mechanisms involved in developmental processes and responses to environmental cues in plant, with a focus on forest tree species. We consider the possible role of forest tree epigenetics as a new source of adaptive traits in plant breeding, biotechnology, and ecosystem conservation under rapid climate change.     

Cluster analysis of spatial patterns in Malaysian tree species

Tree species in tropical rain forests exhibit a rich panoply of spatial patterns that beg ecological explanation. The analysis of tropical census data typically relies on spatial statistics, which quantify the average aggregation tendency of a species. In this article we develop a cluster-based approach that complements traditional spatial statistics in the exploration and analysis of ecological hypotheses for spatial pattern. We apply this technique to six study species within a fully mapped 50-ha forest census in peninsular Malaysia. For each species we identify the scale(s) of spatial aggregation and the corresponding tree clusters. We study the correlation between cluster locations and abiotic variables such as topography. We find that the distribution of cluster sizes exhibits equilibrium and nonequilibrium behavior depending on species life history. The distribution of tree diameters within clusters also varies according to species life history. At different spatial scales, we find evidence for both niche-based and dispersal-limited processes producing spatial pattern. Our methodology for identifying scales of aggregation and clusters is general; we discuss the method's applicability to spatial problems outside of tropical plant ecology.     

Genome-scale phylogenetics: inferring the plant tree of life from 18,896 gene trees

Phylogenetic analyses using genome-scale data sets must confront incongruence among gene trees, which in plants is exacerbated by frequent gene duplications and losses. Gene tree parsimony (GTP) is a phylogenetic optimization criterion in which a species tree that minimizes the number of gene duplications induced among a set of gene trees is selected. The run time performance of previous implementations has limited its use on large-scale data sets. We used new software that incorporates recent algorithmic advances to examine the performance of GTP on a plant data set consisting of 18,896 gene trees containing 510,922 protein sequences from 136 plant taxa (giving a combined alignment length of >2.9 million characters). The relationships inferred from the GTP analysis were largely consistent with previous large-scale studies of backbone plant phylogeny and resolved some controversial nodes. The placement of taxa that were present in few gene trees generally varied the most among GTP bootstrap replicates. Excluding these taxa either before or after the GTP analysis revealed high levels of phylogenetic support across plants. The analyses supported magnoliids sister to a eudicot + monocot clade and did not support the eurosid I and II clades. This study presents a nuclear genomic perspective on the broad-scale phylogenic relationships among plants, and it demonstrates that nuclear genes with a history of duplication and loss can be phylogenetically informative for resolving the plant tree of life.     

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.     

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.     

How to tell a shrub from a tree: A life‐history perspective from a South African savanna

The ecological differences between ‘shrubs’ and ‘trees’ are surprisingly poorly understood and clear ecological definitions of these two constructs do not exist. It is not clear whether a shrub is simply a small tree or whether shrubs represent a distinct life‐history strategy. This question is of special interest in African savannas, where shrubs and trees often co‐dominate, but are often treated uniformly as ‘woody plants’ even though the tree to shrub ratio is an important determinant of ecosystem functioning. In this study we use data from a long‐term fire experiment, together with a trait‐based approach to test (i) if woody species usually classified as shrubs or trees in African savanna differ in key traits related to disturbance and resource use; and (ii) if these differences justify the interpretation of the two growth forms as distinct life‐history strategies. We measured for 22 of the most common woody plant species of a South African savanna 27 plant traits related to plant architecture, life‐history, leaf characteristics, photosynthesis and resprouting capacity. Furthermore we evaluated their performance during a long‐term fire experiment. We found that woody plants authors call (i) shrubs; (ii) shrubs sometimes small trees; and (3) trees responded differently to long‐term fire treatments. We additionally found significant differences in architecture, diameter‐height‐allometry, foliage density, resprouting vigour after fire, minimum fruiting height and foliar δ¹³C between these three woody plant types. We interpret these findings as evidence for at least two different life‐history‐strategies: an avoidance/adaptation strategy for shrubs (early reproduction + adaptation to minor disturbance) and an escape strategy for trees (promoted investment in height growth + delayed reproduction).     

Plant community composition and structural characteristics of an invaded forest in the Galápagos

Non-native species have invaded habitats worldwide, greatly impacting the structure and function of native communities and ecosystems. To better understand mechanisms of invasion impacts and how to restore highly impacted and transformed ecosystems, studies are needed that evaluate invader effects on both biotic communities and structural characteristics. On Santa Cruz Island in Galápagos we compared biotic (plant species richness, diversity, and community composition) and structural (canopy openness, forest height, and leaf litter) characteristics of a relic forest dominated by an endemic and highly threatened tree and a forest dominated by an invasive tree. The forests are located within the historical distribution of the endemic tree, which now occupies only 1% of its original extent. We found that the invaded forest had 42% lower native plant species richness and 17% less plant diversity than the endemic tree dominated forest. Additionally, with the invader there was 36% greater non-native plant species richness, 37% higher non-native plant diversity, and highly dissimilar plant composition when compared to the endemic-dominated forest. Additionally, the invaded forest had a more open and taller tree canopy and greater leaf litter cover than native forest. The presence of the invasive tree and the associated forest structural changes were the primary factors in models that best explained higher non-native diversity in the invaded forest. Our correlational results suggest that an invasive tree has significantly altered plant assemblage and forest structural characteristics in this unique ecosystem. Experiments that remove the invader and evaluate native plant community responses are needed to identify thresholds for practical restoration of this threatened and biologically unique native forest.     

Quantifying the degree of self-nestedness of trees: application to the structural analysis of plants

In this paper, we are interested in the problem of approximating trees by trees with a particular self-nested structure. Self-nested trees are such that all their subtrees of a given height are isomorphic. We show that these trees present remarkable compression properties, with high compression rates. In order to measure how far a tree is from being a self-nested tree, we then study how to quantify the degree of self-nestedness of any tree. For this, we define a measure of the self-nestedness of a tree by constructing a self-nested tree that minimizes the distance of the original tree to the set of self-nested trees that embed the initial tree. We show that this measure can be computed in polynomial time and depict the corresponding algorithm. The distance to this nearest embedding self-nested tree (NEST) is then used to define compression coefficients that reflect the compressibility of a tree. To illustrate this approach, we then apply these notions to the analysis of plant branching structures. Based on a database of simulated theoretical plants in which different levels of noise have been introduced, we evaluate the method and show that the NESTs of such branching structures restore partly or completely the original, noiseless, branching structures. The whole approach is then applied to the analysis of a real plant (a rice panicle) whose topological structure was completely measured. We show that the NEST of this plant may be interpreted in biological terms and may be used to reveal important aspects of the plant growth.     

生长型分类方案不同导致森林生态系统植物功能性状的统计偏差

将植物划分为不同的生长型来统计植物功能性状特征,是当前植物性状研究中常用的方法;但生长型分类方案的不同很可能造成植物功能性状统计分析的偏差,对此偏差的评估却尚未见报道。根据植物志描述及野外调查实际情况,将生长型划分为3种不同的分类方案：分类1：根据植物志信息划分为传统意义的乔木和灌木;分类2：根据树高和胸径划分乔木、小乔木和灌木;分类3：仅根据树高划分乔木层与灌木层的乔木和灌木。以东部亚热带常绿阔叶林区域的浙江金华北山35种优势阔叶木本植物的枝叶性状为研究对象,比较不同生长型分类对植物枝叶性状统计数据的影响。结果表明：（1）与传统的分类1相比,分类2对乔木植物枝叶性状影响的显著程度要高于分类3,而对灌木植物枝叶性状的影响程度低于分类3;但不同生长型分类方案中乔木和灌木的枝叶性状总体差异不显著。而与分类2小乔木相比,分类1以及分类2内部的乔木和灌木生长型的性状与分类2小乔木差异非常明显;（2）将不同生长型植物再划分为不同生活型后,不同生长型分类方案对性状统计的影响增大。无论是常绿还是落叶生活型的小乔木,其与不同生活型乔木和灌木的性状差异仍然显著。可见,不同的生长型分类方案可造成植物功能性状统计的差异;把小乔木植物这一功能类群划分出来,能更好地反映森林生态系统性状特征的差异性。     

Altitudinal Variations in Reproductive Allocation of Bergenia purpurascens (Saxifragaceae)

The strategy of resource allocation between vegetative and reproductive functions, quantitative relationship between size and reproductive output are central aspects of plant life history. To test the tactics of resource allocation and its altitudinal trend, we examined the reproductive allocation (RA) of Bergenia purpurascens (Saxifragaceae), in six populations along a shady slope in Sejila Mountain of southeast Tibet, at an altitude gradient from 4 200 m to 4 640 m. Our results showed that (1) with increasing altitude, vegetative biomass, reproductive biomass, total aboveground biomass, flower number per plant and length of flower stalk decreased significantly, but the number of leaves did not change greatly. However, the change of RA did not show a monotonic trend when altitude increased, shifting from significantly decreasing below the tree line to slightly increasing above it; (2) vegetative biomass was positively correlated with reproductive biomass, but negatively correlated with RA in all populations, but the level of significance was different among the populations; (3) RA decreased with individual size in all populations, whereas the relationship between absolute resource allocated to reproduction and individual size was allometric; (4) reproductive allometry and a size threshold for reproduction did exist in this alpine perennial, but the obvious altitudinal trend was only found along the populations below the tree line, not above it. We then concluded the altitude could not fully explain the change of resource allocation strategy of this alpine perennial, and different effects of size and habitat on RA may result from various environmental constraints along the altitudinal gradient or genetic background. Therefore, each individual within a population will follow its own developmental trajectory shaped by its genotype and the habitats. The most innovative finding was plant adaptation and resource trade off might be sharply altered at the tree line, which is a sensitive area in alpine mountains. Further investigations are needed to better understand the relationship between the reproductive allocation and changing environmental conditions.     

Interacting effects of warming and drought on regeneration and early growth of Acer pseudoplatanus and A. platanoides

Climate change is acting on several aspects of plant life cycles, including the sexual reproductive stage, which is considered amongst the most sensitive life‐cycle phases. In temperate forests, it is expected that climate change will lead to a compositional change in community structure due to changes in the dominance of currently more abundant forest tree species. Increasing our understanding of the effects of climate change on currently secondary tree species recruitment is therefore important to better understand and forecast population and community dynamics in forests. Here, we analyse the interactive effects of rising temperatures and soil moisture reduction on germination, seedling survival and early growth of two important secondary European tree species, Acer pseudoplatanus and A. platanoides. Additionally, we analyse the effect of the temperature experienced by the mother tree during seed production by collecting seeds of both species along a 2200‐km long latitudinal gradient. For most of the responses, A. platanoides showed higher sensitivity to the treatments applied, and especially to its joint manipulation, which for some variables resulted in additive effects while for others only partial compensation. In both species, germination and survival decreased with rising temperatures and/or soil moisture reduction while early growth decreased with declining soil moisture content. We conclude that although A. platanoides germination and survival were more affected after the applied treatments, its initial higher germination and larger seedlings might allow this species to be relatively more successful than A. pseudoplatanus in the face of climate change.     

Food selection in <Emphasis Type="Italic">Microtus arvalis</Emphasis>: the role of plant functional traits

We offered captive common voles (Microtus arvalis) a choice of 11 plant species (representing four ecological groups) growing in vivaria. Selection was evaluated by measuring (1) the biomass of each plant species consumed and (2) functional and life-history plant traits. The legume Trifolium pratense, known for its high nutrient level, and well accessible rosette forbs creating the highest biomass at the soil ground level, were mostly preferred. Voles avoided mainly grasses and the creeping forb Thymus pulegioides. The experiment showed that foraging was strongly plant species-specific. We assessed whether plant functional traits explain selective foraging in common voles. To explore this, we reanalyzed Holišová’s (1959) data about common vole stomach contents and plant trait databases. Regression tree analysis indicated that plant guild and life span were the best predictors of dietary selection, with a probability exceeding 0.5 that voles would eat more grasses and/or legumes than forbs. These results do not correspond with the feeding trial. We suggest that the voles usually consume grasses in the field because grasses are abundant and readily available, but prefer protein-rich forbs when possible.     

The Role of Soil Nutrients in Boundaries between Mangrove and Herbaceous Assemblages in a Tropical Estuary

We aimed to determine the role of soil nutrients in the herbaceous‐tree boundary in a tropical mangrove forest. We assembled plots in this boundary such that each was initially covered by ~50% of each life form. We added NPK to three of the plots monthly and monitored them, along with the control plots, for 18 mo. In plots with added NPK, mangrove branches grew over and shaded the herbaceous plants, resulting in the presence of mostly bare soil. The mangrove trees eventually covered ~80% of the experimental plots, but the proportion of cover in the control plots remained unaltered. We conclude that soil nutrient level modulates the mangrove‐herbaceous plant interaction in steps, and from a given level there is a shift in the dominant life form. This mechanism leads to a sharp transition over a smooth nutritional gradient.     

Effects of gene flow on phenotype matching between two varieties of Joshua tree (Yucca brevifolia; Agavaceae) and their pollinators

In animal‐pollinated plants, local adaptation to pollinator behaviour or morphology can restrict gene flow among plant populations; but gene flow may also prevent divergent adaptation. Here, we examine possible effects of gene flow on plant–pollinator trait matching in two varieties of Joshua tree (Agavaceae: Yucca brevifolia). The two varieties differ in strikingly in floral morphology, which matches differences in the morphology of their pollinators. However, this codivergence is not present at a smaller scale: within the two varieties of Joshua tree, variation in floral morphology between demes is not correlated with differences in moth morphology. We use population genetic data for Joshua tree and its pollinators to test the hypotheses that gene flow between Joshua tree populations is structured by pollinator specificity, and that gene flow within the divergent plant–pollinator associations ‘swamps’ fine‐scale coadaptation. Our data show that Joshua tree populations are structured by pollinator association, but the two tree varieties are only weakly isolated – meaning that their phenotypic differences are maintained in the face of significant gene flow. Coalescent analysis of gene flow between the two Joshua tree types suggests that it may be shaped by asymmetric pollinator specificity, which has been observed in a narrow zone of sympatry. Finally, we find evidence suggesting that gene flow among Joshua tree sites may shape floral morphology within one plant–pollinator association, but not the other.     

Resource tracking and its conservation implications for an obligate frugivore (Procnias tricarunculatus,the three‐wattled bellbird)

In Monteverde, Costa Rica, the vulnerable Three‐wattled Bellbird (Procnias tricarunculatus) feeds primarily upon the fruit of Lauraceae species during its reproductive and post‐reproductive seasons. To understand and advance appropriate conservation measures, this study identified the bellbird's foraging challenges in its search for a temporally and spatially fluctuating resource. Although there are at least 96 species of Lauraceae found in the five life zones of Monteverde, the distinct distributions of tree species both among and within life zones require the bellbirds to track seasonal fruiting across the various zones. In this 6‐year study, we monitored the fruiting of tree species and bellbird abundance in 24 study plots within its post‐reproductive life zone, the Premontane Wet forest, to identify preferred bellbird food resources and how the fruiting of these species drives the spatial distribution of the bellbird. Our research revealed phenological patterns of annual, biennial, and triennial fruiting with high levels of fruiting synchrony within several identified key fruit species. Of critical conservation importance is that no single species of Lauraceae produced a consistent food supply for bellbirds each year. Therefore, even within life zones, the bellbird's survival depends on its mobility to search for and obtain fruit, as well as the availability of fruits of multiple tree species. The conservation implications include focused attention on multiple core areas within given life zones, protection of existing forest and remnant trees, and forest restoration with plantings of multiple tree species. We suspect that other tropical frugivorous species face similar conservation challenges.     

Niches of temperate tree species converge towards nutrient‐richer conditions over ontogeny

Niche changes during a species’ lifespan are known as ontogenetic niche shifts. These shifts reflect changes in resource availability, requirements, organisms’ foraging ability and/or size‐dependent biotic interactions. In the plant kingdom, however, this issue remains poorly covered. We investigated nutritional niche shifts over the ontogeny of 23 temperate tree species (among nine phylogenetic families) by a synchronic approach. We used 1963 temporary phytoecological surveys conducted throughout metropolitan French forests. The realised niches of three life‐history stages (seedling: <0.5 m; sapling: >0.5 m and < 8 m; tree: >8 m) in each tree species were modelled on the basis of presence/absence data and the main factors of species distribution (energy, water and nutritional resources). We computed the nutritional optima and amplitudes by bootstrapping partial response curves for the C:N ratio and base saturation rate. We assessed changes in these niche parameters over ontogeny and also evaluated the relative importance of ontogenetic shifts in the differentiation of nutritional niche among the selected tree species. The tree stage was found to occur mainly at higher nutrient availability than the seedling (+16.3% on the nutritional gradient) or sapling (+11.1%) stages. In addition, nutritional niches of tree species exhibited, successively, a niche enlargement in eutrophic conditions and a niche restriction in oligotrophic conditions during growth. These global nutritional niche shifts observed over the species’ lifespan contributed moderately but significantly to the niche separation in temperate tree communities (up to 4.5%). We interpreted niche shifts as a response to an increase in nutritional requirements over ontogeny, leading to an intra‐specific selection where individuals established in eutrophic soils have the maximal fitness. Biotic interactions and temporal changes in the environment may secondarily enhance or counteract the process. The importance of ontogenetic niche shifts requires consideration in the study of species autecology and plant community organisation.     

Integrating biogeography,threat and evolutionary data to explore extinction crisis in the taxonomic group of cycads

Will the ongoing extinction crisis cause a severe loss of evolutionary information accumulated over millions of years on the tree of life? This question has been largely explored, particularly for vertebrates and angiosperms. However, no equivalent effort has been devoted to gymnosperms. Here, we address this question focusing on cycads, the gymnosperm group exhibiting the highest proportion of threatened species in the plant kingdom. We assembled the first complete phylogeny of cycads and assessed how species loss under three scenarios would impact the cycad tree of life. These scenarios are as follows: (1) All top 50% of evolutionarily distinct (ED ) species are lost; (2) all threatened species are lost; and (3) only all threatened species in each IUCN category are lost. Finally, we analyzed the biogeographical pattern of cycad diversity hotspots and tested for gaps in the current global conservation network. First, we showed that threatened species are not significantly clustered on the cycad tree of life. Second, we showed that the loss of all vulnerable or endangered species does not depart significantly from random loss. In contrast, the loss of all top 50% ED , all threatened or all critically endangered species, would result in a greater loss of PD (Phylogenetic Diversity) than expected. To inform conservation decisions, we defined five hotpots of diversity, and depending on the diversity metric used, these hotspots are located in Southern Africa, Australia, Indo‐Pacific, and Mexico and all are found within protected areas. We conclude that the phylogenetic diversity accumulated over millions of years in the cycad tree of life would not survive the current extinction crisis. As such, prioritizing efforts based on ED and concentrating efforts on critically endangered species particularly in southern Africa, Australia, Indo‐Pacific, and Mexico are required to safeguarding the evolutionary diversity in the cycad tree of life.     

A trait‐based plant economic framework can help increase the value of reforestation for conservation

While reforestation is gaining momentum to moderate climate change via carbon sequestration, there is also an opportunity to use tree planting to confront declining global biodiversity. Where tree species vary in support of diversity, selecting appropriate species for planting could increase conservation effectiveness. We used a common garden experiment in Borneo using 24 native tree species to examine how variation among tree species in their support of beetle diversity is predicted by plant traits associated with “acquisitive” and “conservative” resource acquisition strategies. We evaluate three hypotheses: (1) beetle communities show fidelity to host identity as indicated by variation in abundance and diversity among tree species, (2) the leaf economic spectrum partially explains this variation as shown by beetle preferences for plant species that are predicted by plant traits, and (3) a small number of selected tree species can capture higher beetle species richness than a random tree species community. We found high variation among tree species in supporting three highly intercorrelated metrics of beetle communities: abundance, richness, and Shannon diversity. Variation in support of beetle communities was predicted by plant traits and varied by plant functional groups; within the dipterocarp family, high beetle diversity was predicted by conservative traits such as high wood density and slow growth, and in non‐dipterocarps by the acquisitive traits of high foliar K and rapid growth. Using species accumulation curves and extrapolation to twice the original sample size, we show that 48 tree species were not enough to reach asymptote levels of beetle richness. Nevertheless, species accumulation curves of the six tree species with the highest richness had steeper slopes and supported 33% higher richness than a random community of tree species. Reforestation projects concerned about conservation can benefit by identifying tree species with a disproportional capacity to support biodiversity based on plant traits.