Research Progress on the Theory of Delineation and Zoning of Nature Reserves in China and Discussion of Strategies

Na ZHANG Jingbo ZHOU

Abstract The construction of nature reserves plays an essential role in protecting natural resources and the ecological environment, especially the conservation of rare and endangered species, which also has a vital function in maintaining the ecological security of the national territory. China has built up a comprehensive network of nature reserves after waves of development, but there are still many problems with the management of nature reserves, such as the definition of boundaries, zoning theory, zoning methods, and legal protection. The definition of boundaries is a tool for effective management of nature reserves, but achieving scientific and effective boundary definition is a difficult problem for the construction and management of reserves. It is also a problem that must be solved to achieve sustainable development. This paper compared three fundamental theories of the conservation of species in nature reserves, namely island biogeography, metapopulation, and landscape ecology, from connotations, applications and advantages, and disadvantages, and discussed the progress of theoretical research to provide valuable references for the definition and planning of effective boundaries of nature reserves.

Key words Nature reserves; Island biogeography; Metapopulation; Landscape ecology

Received: August 26, 2021  Accepted: August 28, 2021

Supported by the Central Universities Basic Research Operations Project (JS2019HGXJ0027).

Na ZHANG (1983-) , female, P. R. China, PhD, lecturer, devoted to research about ecological management of protected areas.

*Corresponding author. Jingbo ZHOU (1967-), female, P. R. China, professor, senior engineer. E-mail: 515911792@qq.com.

China built its first nature reserve, the Zhaoqing Dinghushan Nature Reserve in Guangdong Province in 1956, but the construction of nature reserves in China experienced a long period of stagnation from the 1960s to 1980s and has developed rapidly since the reform and opening up［1-2］. The number of nature reserves in China had reached 2 750 by the end of 2020, with a total area of 147.33 million hectares, accounting for about 14.88% of the country’s land area［3］. The construction of nature reserves plays a vital role in protecting natural resources and the ecological environment, especially the conservation of rare and endangered species, which also has an essential function in maintaining the ecological security of the national territory［4-5］. However, after waves of development, there are still many problems in achieving the scientific management of nature reserves, such as defining boundaries, zoning theory, zoning methods, and legal protection［6-7］. At present, the fundamental theories of nature reserves include: island biogeography theory, metapopulation theory, and landscape ecology theory. However, these theories and methods are only based on the study of the conservation of target species while ignoring the inner ecological properties of nature reserves, and therefore, there are still certain shortcomings［8-9］. It is necessary to study them in-depth further to strengthen the establishment and effective management of nature reserves.

Island Biogeography Theory

Basic theories

MacAr-thur and Wilson proposed the island biogeographic equilibrium theory in 1967, which describes the relationship between equilibrium points in island biomes and rates of opening and extinction by predicting the possible results of a given set of conditions using a quantitative theoretical model, the first time the relationship between species richness and area and degree of isolation was described in dynamic terms［10-11］. This theory suggests that the species richness in an island depends on the migration and extinction rates of species and that the increase or decrease of these two processes leads to changes in species richness, with the rate of migration and extinction depending on the size of the island and the degree of isolation,  i.e. , there are two effects: the area effect and the distance affect—the extinction rate decreasing with increasing island size and the migration rate decreasing with increasing degree of isolation［12-13］. When the rate of migration equals the rate of extinction, the number of species in the island reaches a dynamic equilibrium,  i.e. , the number of species is relatively stable, but the composition of species is constantly changing and renewing［10］. A large number of islands with their separate biomes or ecosystems have given island biogeography a special place in overall biogeography, and its theory has been applied to the study of island habitats widely. The number of biological species on an island depends on the island’s size, age, the diversity of habitats, the possibility for pioneers to enter the island from the source and the richness of the sources, and the balance between the migration rate of new species and extinction rate of extant species［14-15］.

The application of island biogeography

In the 1970s and 1980s, the application of island biogeography theory to guide the design of wildlife reserves became a hot topic of research, especially after May presented a succinct summary of the bird studies that were the basis for a general utility perspective on the conservation of certain species［16-17］. However, at that time, he pointed out that the models of wildlife reserves did not match up with potentially important biological facts［18］. However, the species-area relationship has an essential place in island biogeography theory, which is one of the fundamental theories in the early development of conservation biology and is used in the design of nature reserves and the conservation of endangered species widely［19］.

In the extensive literature on the application of island biogeography to guide the design of wildlife reserves, there has been debate on whether to establish one large nature reserve or several smaller ones (the so-called SLOSS principle)［20］. Hypothesis one: (i) The larger the protected area, the better; (ii) one large reserve is better than several small reserves with the same total area; (iii) for some special habitats and taxa, it is better to design several reserves with the closer distance between them; (iv) it is better to connect nature reserves with access roads to increase the migration rate of species; and (v) to avoid the "peninsula effect," it is better to make the reserves circular［21］. Hypothesis two: For species with high dispersal, translocation, and colonization abilities, small reserves may hold more species when the sum of several small reserves equals a large reserve, while for species with very low dispersal, translocation, and colonization abilities, large reserves may hold more species［22］. In 1933, Wright  et al. ［23］ proposed a design idea that is still important today: the concept of minimum population and minimum area and the ratio of the reserve area to its circumference ten years later. In 1971, Smiberloff attempted to test the scientific validity of the population-area hypothesis by an experiment, which showed that groups of reserves were richer in species than a larger reserve. However, the experiment was not representative due to its small number and shortcomings, such as increasing extinction rates with few populations［24］. Whereas, in fact, island biogeography theory is neutral to the SLOSS principle-species richness depended on the colonization capacity of species on a site-Simberloff  et al. ［25］ considered on its own, the theory could predict that several small reserves would maintain a more significant number of species than the equivalent one with a large area. Tj［26］ applied a species diversity model to explain the factors influencing the optimum number of reserves concerning the species and area curves of two or more reserves, pointing out that it is more appropriate to establish a small number of reserves with larger areas if the overlap of species between reserves is high, and that the number of species increases with the area. In contrast, the minimum area for population survival, spatial aggregation, and uneven species richness also increase. He also noted that if the establishment of a reserve emphasizes the conservation of a rare species rather than a majority of species, then an enormous reserve is the best option［27］.

Contradictions and deficiencies of island biogeography

Island biogeography played a vital role in the emergence of the theory of nature reserves. However, many studies have shown that the subject was not mature enough, and its application was limited, especially in nature reserves［28］. Higgs pointed out that the application of island biogeography theory to the design of nature reserves is controversial. The species-area relationship suggests that whether to support the establishment of several small island groups or one island of equal size depends on the proportion of species shared by each island group. Other factors ( e.g. , irreversibility and the effects of natural disasters and species extinction after establishing the reserve) may also support any of the above strategies for establishing two or more island groups or a large protected area of equal size. Consequently, island biogeography and related ecological theories have generated conflicting views on conservation strategies［29］. Daniel  et al. ［30］ argued that island biogeographic equilibrium theory is partially applicable to ecosystems. Although it is subject to turnover (extinction and migration) at a certain point in time, the problem lies in the vague definition of this specific point in time, with a representative notion that the turnover of specific populations and sites is so slow that the equilibrium point cannot be reached at all. The application of island biogeography theory to conservation practice is too early. From both theoretical and practical perspectives, the main conclusion of its application—the establishment of a stand-alone reserve with the maximum area is incorrect for multiple biologically viable site conditions. The costs and irreversibility of large-scale conservation programs require a cautious attitude towards applying a theory without full validation.

In fact, the island biogeography theory itself has some limitations, such as attributing all biological characteristics of islands to one variable, the number of species, and ignoring critical ecological factors such as competition, predation, reciprocal symbiosis, and evolution; and the island biogeography theory emphasizes species richness at equilibrium, while the equilibrium assumption has been questioned more often in practice.

Metapopulation Theory

Connotation and development of metapopulation theory

After the research wave of the 1970s, the accuracy and scope of application of island biogeography theory became controversial, especially since the late 1980s, when more and more ecological articles began to question the veracity of island biogeography theory, and people began to turn their attention to metapopulation theory slowly［31］. No matter in natural environments or artificial landscapes, due to habitat fragmentation, species inhabit these fragmented spatial ‘islands,’ which exist as a metapopulationand species often undergo extinction through the metapopulation stage［32］. Therefore, metapopulation has become a hot topic of research in conservation biology in recent years. Additionally, metapopulation theory has received much attention in population biology and conservation biology because the metapopulations of many local populations that are easy to extinction exist in these fragmented landscapes, as a balance of random extinction and recolonization of populations［33］.

Levins proposed the concept of a metapopulation in 1969, defining it as a "population of populations,"  i.e. , a collection of local populations within a relatively independent area, which becomes a whole through a degree of individual migration. Metapopulation studies are centered on a spatial perspective as a network of habitat patches and explore the dynamics of extinction and recolonization among local populations within these patch networks［34-35］. In the 1990s, Hanski  et al. ［36］ proposed a new definition of the metapopulation, giving a new dimension to metapopulation theory. He argued that spatially realistic metapopulation theory is a model of species dynamics in highly fragmented landscape environments, focusing on stochastic patch occupancy models and the presence and extinction of focal species in habitat patches［37］. Chinese scholar Wu［38］  believed that metapopulation theory is about the movement and decreasing of populations in a landscape patch assemblage and the interaction between spatial patterns and population ecological processes. Current research about metapopulation theory has focused on population dynamics, particularly concerning changes in migration due to the extinction and recolonization of each small population within a population, and these studies are essential for the conservation of organisms as they provide an empirical basis for assessing the effects of connectivity between geometric habitat patches and local and regional populations, and a framework for predicting future habitat loss and intra-population segregation［39-40］.

Model and application of metapopulation theory

In modern spatially realistic metapopulation models, population survival is assessed using a stochastic occupancy model［41］. The Levins model is a deterministic approximation of the dynamic variables of the homogeneous stochastic occupancy patch model, in which habitat patches are divided into settled and unsettled, and the true size of local populations is ignored［42］. It assumes that all populations have a constant risk of extinction and that the probability of population establishment is proportional to the proportion of patches with populations settled (P, migration source) and the proportion of patches currently unsettled (1- P , migration target). Based on the above assumptions, the changing rate of  P  is  dPdt=cP(1-P)-eP  (where  c  and  e  are parameters for colonization and extinction respectively), and the balance value of  P  is:  P=1-e/c , as can be seen from the Levins mode, if  e/c <1, metapopulation would be able to survive ( P >0) sustainably［37］. Assuming that the patches are identical and that the dynamics are independent, the variation rate of anyone occupied patch is also identical. The rate of variation of the likelihood of patch i being occupied in a randomly occupied patch model in a heterogeneous environment a system of n equations gives deterministic continuous-time approximation for a network of n patches,  dpi/dt=Ci(p)(1-pi)-Ei(p)pi , where  Ci(p)  is the colonization rate when patch i is empty,  Ei(p)  is the extinction rate when patch i is occupied, p is the possession probability of  n , and if  Ei(p)/Ci(p)≥1 , then the population would become extinct［32］. As the risk of population extinction decreases with increasing island (or patch) size and the probability of colonization decreases with increasing island (or patch) isolation［42］, species will first disappear from the smallest habitat patches as their size decreases and their isolation increases. Species can persist a little longer in larger habitat patches, and if islands are ‘too’ small or ‘too’ far away from each other, the metapopulation will become extinct faster.

As seen from the above, the core of metapopulation research is to consider these geographic spaces as networks of habitat patches and explore the population dynamics of extinction and recolonization among local populations within these patch networks. Metapopulation theory has been studied extensively, and many scholars have conducted studies on protozoa, amphibians, and vertebrate brown bears, such as Hanski on the population dynamics of the  Melitaea cinxia  L in the Finnish, Meyer  et al.  on forest frogs in Europe, and Graighead  et al. &nbsp;on brown bears［43-44］. These studies suggest that the long-term persistence of endangered species requires at least a number of well-connected habitat patches and that the habitat patches should be ideally spaced to meet migration requirements. In fact, the metapopulation theory and the island biogeography theory have the same roots, as both theories explore the conservation of endangered species and biodiversity. Both adopt an area-segregation paradigm, but the island biogeography focuses on the effects of the scale of habitat fragmentation and structural segregation of species composition and the richness of species on islands［45］. In contrast, the concept of metapopulation dynamics has focused on connectivity and changes within populations and regional persistence conditions for species with unstable local populations, avoiding local or even the eventual extinction of species［46］.

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Landscape Ecology

Connotation and research progress of landscape ecology theory

Human activities have caused enormous impacts on biodiversity at all levels and hierarchies, with landscape fragmentation and habitat destruction being the main reasons for the accelerated rate of species extinction globally. Meanwhile, habitat loss and isolation are linked to land conversion caused by human activities and also pose the most serious threat to the planet’s biodiversity. Hence, landscape ecology and landscape design and planning principles provide an essential platform for researching habitat fragmentation［47］. In 1939, the German geobotanist C. Troll proposed the concept of landscape ecology in his study of land use in East Africa by aerial film interpretation, which was used to represent the analysis of the interrelationships of the natural-biological complexes that govern a regional unit, and was derived mainly from the close integration of the two scientific perspectives of landscape and ecology in biology［48］. Although the concept of landscape ecology was introduced in the 1930s, it was only in the 1980s that landscape ecology was developed well, and meanwhile, it brought new ideas, theories, and methods to the nature reserves theory.

Landscape ecology research focuses on the following aspects: the formation and dynamics of spatial heterogeneity or patterns, the interaction between spatial heterogeneity and ecological processes, the classification structure characteristics of landscapes, the interrelationship between pattern-process-scale, and the feedback relationship between human activities and landscape structure and function. The multi-scale and multi-dimensional coupled study of landscape patterns and ecological processes is based on a specific region and a particular target-oriented division of land units or ecosystem types. It combines small-scale observational studies and large-scale spatial dynamic simulations through land evaluation and model integration to build a coupled model system with good adaptability based on achieving regional targets［49］. In the development of landscape ecology, island biogeography has made the most outstanding contribution to landscape ecology by cleverly linking the spatial characteristics of ‘island’ landscape elements to species populations in a theoretical formula, which has made a significant contribution to the development of landscape ecology［50］. The significance of landscape planning and design in biodiversity conservation has attracted a great deal of attention from biologists, as Wilson said: "As a developing profession, landscape design will play a key role in the conservation of biodiversity, through the clever arrangement of woodlands, green belts, water systems, reservoirs, and artificial pools and lakes, it is possible to keep biodiversity at a high level despite the increasing artificiality of the environment." Landscape design is the redistribution of resources on a specific scale and proposes solutions for the rational use of landscape resources by studying the influence of landscape patterns on ecological processes, and thus landscape ecology construction and landscape structure design have an important role in biodiversity conservation［51］.

Application of landscape ecology theory

In the process of spatial analysis of landscapes, establishing the interconnection of patterns and processes and the infiltration of other theories such as island biogeography and infiltration theory towards landscape ecology, many landscape indices have been developed to describe landscape patterns and their changes, such as the number and size of patches, the number of patch sub-dimensions, the number of edges and diversity between landscape elements, dominance and spread［52］. Fu  et al. ［53］ proposed that species diversity in patches is related to the following patch characteristics in the following order: S=f ［+habitat diversity+/-disturbance+area+age+landscape heterogeneity-degree of isolation-boundary discontinuity］, from the equation above it can be seen that species diversity is significantly related to patch area and therefore in the design of nature reserves for the conservation of rare and endangered species and the maintenance of stable ecosystems, the area of the reserve is the main factor, while other factors such as the degree of isolation, age, and shape of the patch are secondary. The ability to quantify landscape structure is a prerequisite for studying landscape function and change, and thus, the development of methods to quantify landscape structure has been given much attention. Nonetheless, there are currently many formal indicators in the numerous landscape indices, meaning that they cannot solve the problems that exist in the current reality, so efforts have mainly been made to adapt the data to meet the needs of specific research objectives and the user-generated computer programs have been used for analysis［54］. The landscape pattern index is the primary method to quantify the landscape pattern and ecological processes, but many landscape pattern indices are in trouble because it is challenging to integrate the pattern and processes together. Chen  et al. ［55］ introduced the concepts of "source" and "gather" in atmospheric pollution into landscape ecology in order to provide a basis for quantitative analysis of landscape patterns and ecological processes and to evaluate the suitability of landscape spatial patterns and provide a way to design biodiversity through landscape assessment models to analyze the role of different landscape types concerning target species. Wu  et al. ［56］ proposed to use dissipative structure theory and hierarchical system theory to analyze and discuss problems such as ecological balance, conservation of biodiversity, protection of natural landscapes, and restoration and reconstruction of habitat ecosystems. Zhou  et al. ［57］ and Wu  et al. ［58］ discussed the relationship between landscape structure and biodiversity conservation, such as the relationship between the size, shape, and type of patches in the landscape and biodiversity, the role of corridors in biodiversity conservation, and the edge effect and biodiversity and the effect of human disturbance on biodiversity.

The landscape design pathways for biodiversity conservation can be divided into two main types: a species-based landscape design pathway and a landscape element-based design pathway. The former identifies species firstly and then designs the landscape pattern according to the ecological characteristics of the species, while the latter takes landscape elements of various scales as conservation objects and designs the landscape pattern according to their spatial location and relationships ［59-60］. Since the 1990s, there has been growing attention to the theory and application of spatial and temporal scales and heterogeneity in landscape ecology, which can influence the flow and spread of resources, species, or disturbances across the landscape, thus with a significant impact on the functions and processes of nature reserve landscapes. The spatial heterogeneity of nature reserves includes three aspects: spatial composition (type, number, and area ratio of ecosystems), spatial configuration (spatial distribution of ecosystems), and spatial correlation (the extent and scale of spatial association of ecosystems and parameters,  etc. ). The landscape pattern of nature reserves determines the distribution of species, resources, and the environment. The study of the human disturbance impact on protected areas and its distinction from natural disturbance will provide theoretical guidance for managing nature reserve landscapes.

Conclusions

The fundamental theories of nature reserves include island biogeography theory, metapopulation theory, and landscape ecology theory. The island biogeography theory was the first to be applied to studying the size of protected areas. The metapopulation theory was proposed during Levins’ research on island biogeography theory, focusing on the size of the smallest population, while the landscape ecology theory is the most commonly applied theory of nature reserves. Each of the 3 theories has specific strengths and weaknesses.

In summary, all three theories consider the boundary definition of reserves from the perspective of the species under conservation, exploring the extent of habitats that meet the sustainable existence of protected species, ignoring the ecological issues of their attributes in the region, and failing to research the population-area relationship of protected species from the overall regional ecological security pattern, which is something that needs to be further improved in the research of nature reserve boundary definition and zoning management.

TheoryCore content AdvantagesLimitations

Island biogeography (1967) The dynamics of species migration and extinction rates determine the species richness on islands. It allows the study of species conservation to move from describing species through qualitative comparisons to verifying the mechanisms of species formation through field simulations; it enriches theories of biogeography and ecology and promotes the knowledge and understanding of the geographical distribution and dynamic patterns of species diversity, and it may have more theoretical significance for the conservation of species diversity by studying the patterns of species change at the community level. It focuses only on the number of species on an island concerning its size, rather than the size and number of individuals within the same species, making it challenging to reflect the heterogeneity of the environment; it does not take into account other critical ecological factors that determine the structure of island communities, such as competition, predation, mutualistic symbiosis, and evolution.

Metapopulation theory (1969) The dynamics of individual migration between local populations and the conditions for species persistence. It considers the exchange between individuals of the same species as well as the genetic exchange between individuals within a species, thus providing a more effective methodological guide for the conservation of endangered species; it combines ecological processes and GIS analysis tools through a spatially explicit model, and the results are spatially visualized, thus enhancing the predictive power of process models and the spatial analysis capabilities of GIS. It assumes that local populations are homogeneous, but that real patch networks vary greatly in patch size, quality, and connectivity, and that absolutely homogeneous habitat do not exist; it assumes that the system is in equilibrium and can only be applied to habitat networks with a large number of patches, and that the survival of the current metapopulation may be deceptive.

Landscape ecology theory (1933) The study of the influence of landscape patterns on ecological processes and the proposal of rational use of landscape resources based on landscape analysis and comprehensive evaluation plays a vital role in biodiversity conservation. Integrating the function of spatially interacting horizontal ecological zones with longitudinal studies, the research pays excellent attention to the development of methods for quantifying landscape structure and has used user-generated computer programs for analysis. There are many quantitative indicators and a lack of science in analyzing ecological functions and processes in practice; its theory needs further improvement.

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