Buntarou Kusumoto

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.

Community structure and topographic distribution of lianas in a watershed on Okinawa, south-western Japan

We investigated spatial distribution of lianas, and examined relationships between distribution and topography in a secondary forest on the northern part of the main island of Okinawa, south-west Japan. All lianas with a stem diameter exceeding 2 cm at a point 1.3 m from the root were inventoried, and stem diameter and root position were measured within a small watershed (16.0625 ha). Geographical information systems (GIS) were used to divide the watershed into 257 cells in total, each cell being 25 m × 25 m. Three topographic parameters were measured in each cell: slope angle, slope aspect and relief. In total, 930 stems from 20 species of lianas were recorded. Distribution of all liana stems and 14 of 15 species analysed were biased toward concave sites. Abundance of total lianas was higher in concave sites than in convex sites. Our results suggest that distribution and abundance of lianas are strongly affected by topography, and that lianas tend to be distributed in sites with more water and nutrients, which are formed by relief in the Okinawa forest. Factors affecting stem density differed among species.

Stand structure and plant species occurrence in forest edge habitat along different aged roads on Okinawa Island, southwestern Japan

We compared stand structure and plant species occurrence on the shoulders of 3-, 10- and 20-year-old roads in a subtropical evergreen broad-leaved forest to document temporal changes in edge effects of forest roads. We recorded 154 vascular species in the study plots, including 3 non-native species. We used generalized linear mixed models to analyze changes in forest structure and plant species composition in relationship to the distance from forest roads. The spatial patterns in stand structure at different distances from roads differed with road age. The large canopy openness on the edge of 3-year-old roads decreased with time after the construction. A progressive decrease in tree height on roadsides was observed after the road construction, suggesting tall trees could not withstand the dry and windy roadside environment. The edge effect on the canopy tended to be larger at higher elevation sites. The spatio-temporal pattern of species occurrence based on distance from roads differed by species. Typical pioneer species such as Schima wallichii and Eurya japonica increased along the edge, while less aggressive pioneer species and understory species decreased. As time passed after the road construction, some climax and understory species decreased at the forest edge, while other climax and understory species increased. The modeling methods proposed in this study could be applied to different roadside and edge study sites.

A theory for ecological survey methods to map individual distributions

Spatially explicit approaches are widely recommended for ecosystem management. The quality of the data, such as presence/absence or habitat maps, affects the management actions recommended and is, therefore, key to management success. However, available data are often biased and incomplete. Previous studies have advanced ways to resolve data bias and missing data, but questions remain about how we design ecological surveys to develop a dataset through field surveys. Ecological surveys may have multiple spatial scales, including the spatial extent of the target ecosystem (observation window), the resolution for mapping individual distributions (mapping unit), and the survey area within each mapping unit (sampling unit). We developed an ecological survey method for mapping individual distributions by applying spatially explicit stochastic models. We used spatial point processes to describe individual spatial placements using either random or clustering processes. We then designed ecological surveys with different spatial scales and individual detectability. We found that the choice of mapping unit affected the presence mapped fraction, and the fraction of the total individuals covered by the presence mapped patches. Tradeoffs were found between these quantities and the map resolution, associated with equivalent asymptotic behaviors for both metrics at sufficiently small and large mapping unit scales. Our approach enabled consideration of the effect of multiple spatial scales in surveys, and estimation of the survey outcomes such as the presence mapped fraction and the number of individuals situated in the presence detected units. The developed theory may facilitate management decision-making and inform the design of monitoring and data gathering.

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.

Spatially Explicit Approach To Population Abundance Estimation In Field Surveys

The abundance of species is a fundamental consideration in ecology and conservation biology. Although broad models have been proposed to estimate the population abundance using existing data, available data is often limited. With no information available, a population estimation will rely on time consuming field surveys. Typically, time is a critical constraint in conservation and often management decisions must be made quickly under the data limited situation. Depending on time and budgetary constraints, the required accuracy of field survey changes significantly. Hence, it is desirable to set up an effective survey design to minimize time and effort of sampling given required accuracy. We examine a spatially-explicit approach to population estimation using spatial point processes, enabling us to explicitly and consistently discuss various sampling designs. We find that the accuracy of abundance estimation varies with both ecological factors and survey design. Although the spatial scale of sampling does not affect estimation accuracy when the underlying individual distribution is random, it decreases with the sampled unit size if individuals tend to form clusters. These results are derived analytically and checked numerically. Obtained insights provide a benchmark to predict the quality of population estimation, and improve survey designs for ecological studies and conservation.

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.

Correction to: A theory for ecological survey methods to map individual distributions

The original version of this article unfortunately contained a mistake. The x-axis in Figs. 4-6 in the original version of this article should be replaced with the x-axis shown in the improved figures below. This does not change the calculations and conclusions.