Takayuki Shiono

Phylogenetic properties of Tertiary relict flora in the east Asian continental islands: imprint of climatic niche conservatism and in situ diversification

&NA; Understanding biodiversity patterns on islands has long been a central aim in ecology and conservation biology. Island‐specific biogeographical processes play substantial roles in the formation of endemic biota. Here, we examined how climate niche conservatism and geohistorical factors are interactively associated with in situ diversification of Tertiary relict flora in the east Asian continental islands. We generated two novel datasets for species distribution and phylogeny that included all of the known vascular plant species in Japan (5575). Then we tested phylogenetic signal of climatic tolerance, in terms of absolute minimum temperature and water balance, and explored environmental predictors of phylogenetic structure (evolutionary derivedness and clustering) of species assemblages. Although phylogenetic signal of climatic tolerance was significant across the phylogeny of most species, the strength of climatic niche conservatism differed among ferns, gymnosperms, angiosperm trees, and angiosperm herbs. For angiosperm trees, cold temperatures acted as environmental filters that generated phylogenetic derivedness/clustering of species assemblages. For fern and angiosperm herb species, however, phylogenetic properties were not associated with climatic harshness. These contrasting patterns among groups reflected climate niche evolution in vascular plants with different growth forms and traits; for example, diversification of angiosperm trees (but not fern and herb) occurred in response to historical climatic cooling. More importantly, geographical constraints contributed to evolutionary radiation that resulted from isolation by distance from the continent or by elevation. Quaternary climate change was also associated with clade‐specific radiation in refugial habitats. The degree to which geographical, geological, and palaeoclimatic variables explain the phylogenetic structure underscores the importance of isolation‐ and habitat‐stability‐related geohistorical processes in driving in situ diversification despite climatic niche conservatism. We propose that the highly endemic flora of the east Asian islands resulted from the interplay of idiosyncratic regional factors, and ecological and evolutionary processes, such as climate niche assembly and adaptive/nonadaptive radiation.

Functional response of plant communities to clearcutting: management impacts differ between forest vegetation zones

Summary Understanding of the ecological impacts of logging practices on biodiversity and associated ecosystem processes is essential for developing sustainable forest management approaches. We documented the impacts of clearcutting on the functional structure of tree and herbaceous communities in hemiboreal, cool-temperate, warm-temperate and subtropical forests in the Japanese archipelago and identified forest vegetation that is vulnerable to deterioration of important ecosystem functions. We combined species data for leaf, stem, flower and fruit traits related to productivity, nutrient cycling and habitat quality for wildlife with phytosociological vegetation data from unmanaged and previously clearcut forests, then calculated functional structure indices [community mean (CM) of trait values, functional richness (FRic) and functional divergence (RaoQ)] of plant communities. For tree species, functional structure indices of specific leaf area (SLA), leaf carbon and nitrogen concentrations, maximum height, wood density and flower size differed between unmanaged and clearcut forests, while for herb species, only maximum height differed between the two forest types. Functional structure indices showed divergent patterns across forest vegetation zones. In hemiboreal, cool-temperate and warm-temperate forests, the CMs of SLA and leaf nitrogen concentration were greater and that of leaf carbon concentration was smaller in clearcut than in unmanaged forests. Although clearcut forests had greater species richness than unmanaged forests, FRic and/or RaoQ of maximum height were smaller in clearcut forests. In hemiboreal and cool-temperate forests, FRic and RaoQ of SLA were also smaller in clearcut forests. In contrast, subtropical forests showed no differences in species richness and functional structures between unmanaged and clearcut forests. Synthesis and applications. Functional redundancy of plant communities differs among traits and among forest vegetation zones. After intensive logging, hemiboreal, cool-temperate and warm-temperate forests were more vulnerable to the loss of ecosystem functions related to leaf and stem traits of tree species than were subtropical forests, which appeared relatively resilient. Locally adaptive management to maintain multiple ecosystem functions should be developed based on the degree of functional complementarity among plant species in forest communities.

Using tree functional diversity to evaluate management impacts in a subtropical forest

The trait-based approach has received much research attention as it provides a heuristic framework for evaluating the ecological impacts of anthropogenic activities on communities and ecosystems. In this study, functional diversity (or structure) measures, such as functional richness, functional evenness, functional divergence, and functional composition, were used to examine management impacts on subtropical forests on the Ryukyu Islands of Japan. Functional indices were compared in tandem with taxonomic diversity indices between three forest types with different management histories: intact old-growth forests, secondary forests after clear-cutting, and abandoned Pinus luchuensis plantations. Species diversity indices were not significantly different among the three forest types. In contrast, functional diversity indices were significantly different among intact forests and managed forests. Functional richness and functional evenness were significantly lower in secondary forests than in intact forests and P. luchuensis plantations. Functional divergence was significantly higher in secondary forests and P. luchuensis plantations than in intact forests. These differences suggest that management activities affected niche space and the patterns of niche differentiation among component species in the functional space of managed forests. Community weighted means for each functional trait were also different among the forest types. The managed forests had greater leaf thickness, leaf dry matter content and maximum height, and lower specific leaf area and leaf nitrogen concentration than intact forests. These differences in functional composition of traits suggested potential functional impacts. This study demonstrated the utilization of species functional traits and community functional structure as a tool of natural experiment for assessing impacts of forest management practices on woodland ecosystems. It was also suggested that logging activities that include large-scale clear-cutting or establishment of P. luchuensis plantations may be incompatible with the conservation of natural ecosystem properties in subtropical forests.

Multiple filters affect tree species assembly in mid-latitude forest communities

Species assembly patterns of local communities are shaped by the balance between multiple abiotic/biotic filters and dispersal that both select individuals from species pools at the regional scale. Knowledge regarding functional assembly can provide insight into the relative importance of the deterministic and stochastic processes that shape species assembly. We evaluated the hierarchical roles of the α niche and β niches by analyzing the influence of environmental filtering relative to functional traits on geographical patterns of tree species assembly in mid-latitude forests. Using forest plot datasets, we examined the α niche traits (leaf and wood traits) and β niche properties (cold/drought tolerance) of tree species, and tested non-randomness (clustering/over-dispersion) of trait assembly based on null models that assumed two types of species pools related to biogeographical regions. For most plots, species assembly patterns fell within the range of random expectation. However, particularly for cold/drought tolerance-related β niche properties, deviation from randomness was frequently found; non-random clustering was predominant in higher latitudes with harsh climates. Our findings demonstrate that both randomness and non-randomness in trait assembly emerged as a result of the α and β niches, although we suggest the potential role of dispersal processes and/or species equalization through trait similarities in generating the prevalence of randomness. Clustering of β niche traits along latitudinal climatic gradients provides clear evidence of species sorting by filtering particular traits. Our results reveal that multiple filters through functional niches and stochastic processes jointly shape geographical patterns of species assembly across mid-latitude forests.

Macroscale estimates of species abundance reveal evolutionary drivers of biodiversity

Evolutionary processes underpin the biodiversity on the planet. Theories advocate that the form of the species abundance distribution (SAD), presented by the number of individuals for each species within an ecological community, is intimately linked to speciation modes such as point mutation and random fission. This prediction has rarely been, however, verified empirically; the fact that species abundance data can be obtained only from local communities critically limits our ability to infer the role of macroevolution in shaping ecological patterns. Here, we developed a novel statistical model to estimate macroscale SADs, the hidden macroecological property, by integrating spatially replicated multispecies detection-nondetection observations and the data on species geographic distributions. We determined abundance of 1,248 woody plant species at a 10 km grid square resolution over East Asian islands across subtropical to temperate biomes, which produced a metacommunity (i.e. species pool) SAD in four insular ecoregions along with its absolute size. The metacommunity SADs indicated lognormal-like distributions, which were well explained by the unified neutral theory of biodiversity and biogeography (UNTB) with protracted speciation, a mode of speciation intermediate between point mutation and random fission. Furthermore, the analyses yielded an estimate of speciation rate in each region that highlighted the importance of geographic characteristics in macroevolutionary processes and predicted the average species lifetime that was congruent with previous estimates. The estimation of macroscale SADs plays a remarkable role in revealing evolutionary diversification of regional species pools.

Environmental filters shaping angiosperm tree assembly along climatic and geographic gradients

Co-ordinating Editor: Leandro Duarte Abstract Question: Globalscale forest censuses provide an opportunity to understand diversification processes in woody plant communities. Based on the climatic or geographic filtering hypotheses associated with tropical niche conservatism and dispersal limitation, we analysed phylogenetic community structures across a wide range of biomes and evaluated to what extent regionspecific processes have influenced largescale diversity patterns of tree species communities across latitude or continent. Location: Global. Methods: We generated a data set of species abundances for 21,379 angiosperm woody plants in 843 plots worldwide. We calculated net relatedness index (NRI) for each plot, based on a single global species pool and regional species pools, and phylogenetic βdiversity (PBD) between plots. Then, we explored the correlations of NRI with climatic and geographic variables, and clarified phylogenetic dissimilarity along geographic and climatic differences. We also compared these patterns for South America, Africa, the IndoPacific, Australia, the Nearctic, Western Palearctic and Eastern Palearctic. Results: NRI based on a globalscale species pool was negatively associated with precipitation and positively associated with Quaternary temperature change. PBD was positively associated with geographic distance and precipitation difference between plots across tropical and extratropical biomes. Moreover, phylogenetic dissimilarity was smaller in extratropical regions than in regions including the tropics, although temperate forests of the Eastern Palearctic showed a greater dissimilarity within extratropical regions. Conclusions: Our findings support predictions of the climatic and geographic filtering hypotheses. Climatic filtering (climatic harshness and paleoclimatic change) relative to tropical niche conservatism played a role in sorting species from the global species pool and shaped the largescale diversity patterns, such as the latitudinal gradient observed across continents. Geographic filtering associated with dispersal limitation substantially contributed to regional divergence of tropical/extratropical biomes among continents. Old, longstanding geographic barriers and recent climatic events differently influenced evolutionary diversification of angiosperm tree communities in tropical and extratropical biomes.

A paradox of latitudinal leaf defense strategies in deciduous and evergreen broadleaved trees

The classical “low latitude–high defense” hypothesis is seldom supported by empirical evidence. In this context, we tested latitudinal patterns in the leaf defense traits of deciduous broadleaved (DB) and evergreen broadleaved (EGB) tree species, which are expected to affect herbivore diversity. We examined the co-occurrence of leaf defense traits (tannin and phenol content, leaf mechanical strength, leaf dry matter content, leaf mass per area, and leaf thickness) in 741 broadleaved tree species and their correlations with species geographical range in East Asian island flora. We discovered contrasting latitudinal defense strategy gradients in DB and EGB tree species. DB species employed chemical defenses (increasing tannin and phenol content) at higher latitudes and physical defenses (softer and thinner leaves) at lower latitudes, whereas EGB tree species exhibited opposite latitudinal defense patterns. The “low latitude high defense” hypothesis included a paradoxical aspect in chemical and physical defense traits across broadleaved tree species. To reconcile paradoxical defense strategies along the latitudinal gradient, we conclude that interactive correlations among leaf traits are controlled by leaf longevity, which differs between DB and EGB tree species.

Correction to: A paradox of latitudinal leaf defense strategies in deciduous and evergreen broadleaved trees

The article “A paradox of latitudinal leaf defense strategies in deciduous and evergreen broadleaved trees”, written by Saihanna Saihanna, Tomoe Tanaka, Yu Okamura, Buntarou Kusumoto, Takayuki Shiono, Toshihide Hirao, Yasuhiro Kubota, and Masashi Murakami, was originally published electronically on the publisher’s internet portal (currently SpringerLink) on 01 June 2018 without open access.

How well are biodiversity drivers reflected in protected areas? A representativeness assessment of the geohistorical gradients that shaped endemic flora in Japan

Protected areas function as a lifeboat that can preserve the origins and maintenance of biodiversity. We assessed the representativeness of biodiversity in existing protected areas in Japan using a distribution dataset and phylogenetic tree for 5565 Japanese vascular plant species. We first examined the overlap of species distribution with the existing protected areas and identified the minimum set representing all plant species. Second, we evaluated the relative importance of environmental variables in explaining the spatial arrangement of protected areas using a random forest model. Finally, we clarified how potential drivers of plant diversity were sufficiently captured within the protected areas network. Although the protected areas captured the majority of species, nearly half of the minimum set areas were selected from outside the existing protected areas. The locations of existing protected areas are mainly associated with geographical and socio-economic factors rather than key biodiversity features (including evolutionary distinctiveness). Moreover, critical biodiversity drivers, which include current climate, paleoclimatic stability, and geographical isolation, were biasedly emulated within the existing protected areas. These findings demonstrate that current conservation planning fails to represent the ecological and evolutionary processes relevant to species sorting, dispersal limitation, and allopatric speciation. In particular, under-representativeness of historically stable habitats that function as evolutionary hotspots or refugia in response to climate changes may pose a threat to the long-term persistence of Japan’s endemic biota. This study provides a fundamental basis for developing prioritization measures to retain species assembly processes and in situ diversification along current climatic and geohistorical gradients.