Short communicationQuantification of water resource utilization efficiency as the main driver of plant diversity in the water-limited ecosystems
Graphical abstract
Introduction
The community ecology has been developing rapidly with many theories attempting to explain the patterns of species diversity and abundance. The neutral theory, metapopulation and metacommunity theories, generalized fraction, Poisson distribution, and entropy theory and so on have been developed in the past 20 years (Hubbell, 2005; Volkov et al., 2005; McGill, 2010; Violle et al., 2012; Harte and Newman, 2014). These theories start with radically different assumptions, and the theories seem extremely different from each other. To form a model with reasonable hypothesis, verifiable and convincible predictions, further research is still needed.
In essence, the distribution of plant diversity is mainly related to the niches of different species. Obviously, the establishment of a biodiversity niche model from the perspective of resource competition is of great significance to the in-depth analysis of the mechanisms which is the foundation of the formation and maintenance of biodiversity as well as the improvement of prediction accuracy.
The most important and basic resource for plant communities is water (Tilman, 1982; Jobbágy et al., 2011; Silvertown et al., 2015), especially in arid areas where water resources are scarce. Water determines the global distribution pattern vegetation and the vegetation types within a region (Jirka et al., 2009; Jobbágy et al., 2011). At the local scale, water also affects carbon dioxide capture, soil nutrient availability, plant growth and development, microbial activity, and interspecific interactions. Although the hydrological niche of plant communities has been verified by experiments in many studies (Silvertown et al., 2015; García-Baquero et al., 2016; Letten et al., 2015), the current theoretical research literature is insufficient. At present, niche theory based on resource competition fails to explain the diversity of tropical rain forest (Hubbell, 2005).
In water-limited terrestrial ecosystem, with environmental filtration and biological filtration (intra- and inter-specific interactions), plant community species can maximize utilization efficiency of community water resource. At the individual level of the plant, there is a trade-off between relative growth rate (RGR) and water use efficiency (WUE) in different species of a community (Huxman and Venable, 2013). At the population level, each species has its own niche, and there is niche differentiation even for the same resource (Li; et al., 2000;nMcGill, 2010). Niche differentiation is conducive to avoid competition and achieve species coexistence. From the perspective of water resource utilization, this niche differentiation ensures that more available water resources are fully utilized. At the same time, different species also have niche overlapped because of the overlap of their functional traits (Violle et al., 2012). When the niche overlaps, the competition for available water resources within the community is the most intense, which stands for the most efficient use of available water resources. At the community level, available water resources have spatiotemporal changes, and their distribution is fractal distribution (Bird et al., 2000). In order to make full use of available water resources, based on species interactions, the community is surely to obtain characteristics related to fractal characteristics (Harte and Newman, 2014). Although a community cannot evolve directly through genetic change as species, communities evolve by improving the capacity of the entire community to exist (Verboef and Morin, 2010). Therefore, the community eventually realizes the utilization efficiency of available water resources to the maximum extent to attain the status of an ecologically stable community and to match the living environment (Trautz et al., 2017). In this paper, in water-limited terrestrial ecosystem, plant community species can maximize water resource utilization efficiency to the community, including water resource utilization characteristics at the individual and population level, all of which are used as the basis for our model construction.
On the other hand, in the development of plant diversity theory, the central unanswered question is what determines the total number of individuals () and the number of species () (McGill, 2010). and are always inputs in existing diversity theory. What drives these? Despite McGill calling the unified theory, a theory of biodiversity, in every case the species richness, , and number of individuals, are inputs to the model rather than predictions (Mutshinda et al., 2009; McGill, 2010; Violle et al., 2012). To date the greatest success in the study of these factors has been empirical (i.e. looking for correlations with environmental variables), where factors like productivity, climate, and altitude seem important. This paper attempts to answer this question based on the theory of the water niche (The main context is shown in Fig. 1).
Based on interspecific and intraspecific interactions in water niche and the maximum utilization efficiency of available water resources in an ecologically stable community, a water niche theoretical model of plant diversity is established and also validated using published survey data. This model we intend to answer in what determine S (the number of species) and N (the total number of individuals) as well as what is the driver of the species-area relationship.
Section snippets
Establishment of the plant diversity model based on water niche
The formation of plant community diversity involves environmental filtration and biological filtration. In water-limited plant communities, biological filtration of interspecific and intraspecific interactions is based on the trade-offs between the utilization efficiency of water resources and the relative growth rate (Huxman and Venable, 2013). Stable communities are established when water resource utilization is maximized (Zuppinger-Dingley et al., 2014; Trautz et al., 2017). This
Species richness and the number of individuals in plant communities based on a water niche model
The above model has a unique solution, and the optimal structure of the community can be determined. From the above results, we know that the number of individuals () and the relationship between the species richness () and the maximum power of the plant community () are as follows (for additional details please see supporting material S1):where ,where is the maximum power of the plant community; is the total quantity of available water
Experimental verification
Due to similarities in habitat, the maximum power of the plant community is proportional to the area of the community, and the average community fitness of the unit area is constant: .
From equations and , the species-area curve (Volkov, 2005; McGill, 2010; He and Hubbell, 2011) is expressed aswhere, , and .
This is a new species area curve, but the ecological significance of its parameters can be clearly seen. In the formula for the species-area
Discussion
In this paper, the theoretical model based on water niche is deducted and verified by theoretical and data verification. This model can explain the main characteristics of plant diversity and is an enrichment and improvement of existing plant diversity theories. Our model deduction is based on maximizing the utilization efficiency of available water resources of an ecologically stable community in water-limited terrestrial ecosystem. Maximizing the efficiency of available water resources in
Data Accessibility
All data from published articles
Authors’ contribution
ZQH, TL, and BLL designed the study; ZQH and BLL established the model, and ZQH drafted the manuscript; TL edited the manuscript, performed analyses, and assisted with preparation of figures; TW, HFL, XRH, OYYN and BZ generated figures and assisted with analyses; all authors contributed to subsequent revisions.
Funding
The research presented in this paper is supported by the National Science Foundation of China (Grant Nos. 31770461 and 31260099), Project of international scientific and technological cooperation of the corps science and Technology Bureau (2016AH001), and the Key Technology R & D Program (Grant No. 2014BAC14B02).
Declaration of Competing Interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
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