Properties of phylogenetic trees generated by Yule-type speciation models

We investigate some discrete structural properties of evolutionary trees generated under simple null models of speciation, such as the Yule model. These models have been used as priors in Bayesian approaches to phylogenetic analysis, and also to test hypotheses concerning the speciation process. In this paper we describe new results for three properties of trees generated under such models. Firstly, for a rooted tree generated by the Yule model we describe the probability distribution on the depth (number of edges from the root) of the most recent common ancestor of a random subset of k species. Next we show that, for trees generated under the Yule model, the approximate position of the root can be estimated from the associated unrooted tree, even for trees with a large number of leaves. Finally, we analyse a biologically motivated extension of the Yule model and describe its distribution on tree shapes when speciation occurs in rapid bursts.     

The prior probabilities of phylogenetic trees

Bayesian methods have become among the most popular methods in phylogenetics, but theoretical opposition to this methodology remains. After providing an introduction to Bayesian theory in this context, I attempt to tackle the problem mentioned most often in the literature: the “problem of the priors”—how to assign prior probabilities to tree hypotheses. I first argue that a recent objection—that an appropriate assignment of priors is impossible—is based on a misunderstanding of what ignorance and bias are. I then consider different methods of assigning prior probabilities to trees. I argue that priors need to be derived from an understanding of how distinct taxa have evolved and that the appropriate evolutionary model is captured by the Yule birth–death process. This process leads to a well-known statistical distribution over trees. Though further modifications may be necessary to model more complex aspects of the branching process, they must be modifications to parameters in an underlying Yule model. Ignoring these Yule priors commits a fallacy leading to mistaken inferences both about the trees themselves and about macroevolutionary processes more generally.     

A new balance index for phylogenetic trees

Several indices that measure the degree of balance of a rooted phylogenetic tree have been proposed so far in the literature. In this work we define and study a new index of this kind, which we call the total cophenetic index: the sum, over all pairs of different leaves, of the depth of their lowest common ancestor. This index makes sense for arbitrary trees, can be computed in linear time and it has a larger range of values and a greater resolution power than other indices like Colless’ or Sackin’s. We compute its maximum and minimum values for arbitrary and binary trees, as well as exact formulas for its expected value for binary trees under the Yule and the uniform models of evolution. As a byproduct of this study, we obtain an exact formula for the expected value of the Sackin index under the uniform model, a result that seems to be new in the literature.     

Linking Trees on Farms with Biodiversity Conservation in Subsistence Farming Systems in Nepal

The Kingdom of Nepal extends 800 km east to west along the southern slopes of the Himalayas. Because of its biogeographical position, Nepal contains biological elements of both the Indo-Malayan and Palaearctic realms. Beside the trees in forest ecosystems, a large number of tree species are maintained on farms as part of subsistence farming systems. The role of these trees in ensuring the sustainability of agricultural production and the importance of traditional farming systems for the conservation of crop diversity have been well documented. However, the status of farm trees and their role in biodiversity conservation are poorly documented. This paper presents a case study of two villages in the western middle hills of Nepal. It highlights the role of traditional agroforestry practices for the conservation of tree diversity and argues that farms can be considered biodiversity reservoirs. Farm trees help to reduce pressure on community and government forests and create a favorable environment for many plant and animal species. Farm trees also provide social functions in that households with many farm trees no longer exercise their communal rights to extract grass products from community forests, which in turn benefits poor and disadvantaged households. The paper discusses possibilities to improve the role of farm trees in biodiversity conservation. It argues for the development of mechanisms such as tax exemptions and conservation credits that provide benefits to rural communities as compensation for their local and global environmental services including biodiversity conservation.     

National biodiversity risk assessment: a composite multivariate and index approach

Due to the shortage of financial resources for international conservation assistance, the setting of priorities for this assistance is an important issue. A national biodiversity risk assessment index (NABRAI) is constructed to quantify national conservation performances and identify nation states of critical conservation concern. The index, which contains measures of biodiversity stock, flow and response measures, attempts to overcome several weaknesses present in other models used to prioritize nations for conservation assistance. Multivariate analyses of the index as well as economic and biodiversity resources reveal significant positive correlations between the NABRAI values and population density as well as land area exposed to high disturbance intensity. The combination of the multivariate analyses and the interpretation of NABRAI values allows for prioritization of biodiversity risk among the global community and can thus serve as an indicator of current priorities for policy makers. The present study also suggests two methods to incorporate a better understanding of biodiversity risk in models of conservation priorities; by including a wider range of variables and by developing a theoretical foundation for the relationship between the categories of variables used in the model.     

The equivalence of two phylogenetic biodiversity measures: the Shapley value and Fair Proportion index

Most biodiversity conservation programs are forced to prioritise species in order to allocate their funding. This paper contains a mathematical proof that provides biological support for one common approach based on phylogenetic indices. Phylogenetic trees describe the evolutionary relationships between a group of taxa. Two indices for computing the distinctiveness of each taxon in a phylogenetic tree are considered here—the Shapley value and the Fair Proportion index. These indices provide a measure of the importance of each taxon for overall biodiversity and have been used to prioritise taxa for conservation. The Shapley value is the biodiversity contribution a taxon is expected to make if all taxa are equally likely to become extinct. This interpretation makes it appealing to use the Shapley value in biodiversity conservation applications. The Fair Proportion index lacks a convenient interpretation, however it is significantly easier to calculate and understand. It has been empirically observed that there is a high correlation between the two indices. This paper shows the mathematical basis for this correlation and proves that as the number of taxa increases, the indices become equivalent. Consequently in biodiversity prioritisation the simpler Fair Proportion index can be used whilst retaining the appealing interpretation of the Shapley value.     

基于空间分析的保护生物学研究

 保护生物学家和生态学家早就认识到只有准确地辨识保护对象的空间位置、 范围、 及其相邻的关系(例如边缘)和连接度, 以及依存的地形和气候等生境条件, 才能发现生物种群和生境在空间的扩散与收缩、 增长与灭绝的动态, 揭示分布的格局, 从而系统、 全面地了解保护对象和生境的存在状态、 破碎程度和变化趋势, 进行有效的自然保护。 得益于新兴的空间分析技术, 保护生物学自20世纪90年代以来取得了很大的进步。基于空间分析的保护生物学研究是最近10年左右大力发展的新保护生物学的重要基础。 该文结合作者的研究工作,综述了基于空间分析的保护生物学项目, 探讨了保护生物学发展历史、 主要研究方法与应用、 以及今后的可能发展趋势。 在生物多样性的丰度和分布的空间解绎部分,通过综述世界保护监测中心的图解全球生物多样性的工作, 如国家尺度的生物多样性水平、 植物多样性的分布中心和维管束植物科的多样性等的空间分布 ,介绍了 Dobson等图示美国主要濒危植物、 鸟类、 鱼类和软体动物等4个主要类群在县(County) 为基本空间单位上分布的空间格局, 讨论了生物多样性空间解绎的意义。在第二部分用世界资源研究所的全球森林监测(Global forest watch)项目, 美国的国家保护缺失区分析(GAP analysis)项目, 美国林务局的无路自然区域(Roadless area)保护项目和加拿大自然审计(Nature audit)项目, 以及北美和东亚生物多样性空间分布的比较分析和生物入侵的空间分析等具体实例来说明生物多样性空间分布变化比较分析方法的应用。 过去20年来, 面向空间格局的生态学和保护生物学研究得到了快速的发展, 特别是空间格局的描述、 由地统计演变而成的空间统计、 地理信息系统、 基于个体(或栅格)的空间解绎模拟模型、 基于斑块(Patch)的种群理论及其发展(如复合种群理论, 源 汇模型等)等。在第三部分, 以美国森林破碎度空间格局分析和美国太平洋西北演替后期森林的空间格局分析为例, 介绍了空间格局分析在保护生物学中的应用。 同时介绍了澳大利亚保护生态学家Lindenmayer 和美国著名景观生态学家Franklin 2002年提出的模板(Matrix)保护理论,把保护的眼光不局限在面积不多而且分散的保护区中,应注意景观模板和保护区相邻的原生区域的综合保护, 这样将大大扩展保护的范围, 并且平衡保护与发展的关系。最后, 介绍了在保护生物学中已有一定应用的空间模型和模拟, 包括了空间解绎模型(Spatial explicit model)、 基于过程(Process-based)的空间模拟模型、 面向代理(Agent-based)的空间适应模拟模型(SWAM)以及与此有关的动态全球植被模型(DGVM)。 通过上面的讨论和综述, 预测一个新的保护生物学的分支: 空间保护生物学, 已经逐渐成熟问世, 这门基于现代信息技术和空间技术的新学科已经而且还将为全球生物多样性的研究和保育作出重大的贡献。     

Forest biodiversity in a changing climate: which logic for conservation strategies?

Climate change has direct and indirect impacts on forest ecosystems worldwide. In this context, changing site conditions and altered disturbance regimes as well as forest management responses are challenging the conservation of biodiversity in forests. Climate-induced dynamics and uncertainties related to future forest ecosystem development are calling into question current conservation strategies and concepts. Given the longevity of trees, slow development rates of forest ecosystems and slow migration rates of many forest species, the planning of adaptation measures in response to climate change are especially difficult though highly important for forest biodiversity conservation. This paper introduces a special issue with eight contributions which deal with a variety of aspects of forest biodiversity conservation in the face of climate change. More specifically, the papers address direct impacts of climate change on forest biodiversity, adaptation measures for forest and conservation management, as well as resulting challenges for conservation strategies and concepts. In conclusion, adaptation measures that enhance diversity and provide different options for future action, thereby maintaining ecosystems’ resilience, as well as conservation management operating on a landscape level, are promoted as being beneficial for coping with uncertainties related to climate change. Adaptive management, which constantly reviews conservation goals and measures, and which takes into account both science-based and local ecological knowledge on climate change can be a valuable tool to inform decisions for forest biodiversity conservation.     

Constraining prior probabilities of phylogenetic trees

Although Bayesian methods are widely used in phylogenetic systematics today, the foundations of this methodology are still debated among both biologists and philosophers. The Bayesian approach to phylogenetic inference requires the assignment of prior probabilities to phylogenetic trees. As in other applications of Bayesian epistemology, the question of whether there is an objective way to assign these prior probabilities is a contested issue. This paper discusses the strategy of constraining the prior probabilities of phylogenetic trees by means of the Principal Principle. In particular, I discuss a proposal due to Velasco (Biol Philos 23:455–473, 2008) of assigning prior probabilities to tree topologies based on the Yule process. By invoking the Principal Principle I argue that prior probabilities of tree topologies should rather be assigned a weighted mixture of probability distributions based on Pinelis’ (P Roy Soc Lond B Bio 270:1425–1431, 2003) multi-rate branching process including both the Yule distribution and the uniform distribution. However, I argue that this solves the problem of the priors of phylogenetic trees only in a weak form.     

Spatial heterogeneity in the Mediterranean Biodiversity Hotspot affects barcoding accuracy of its freshwater fishes

Incomplete knowledge of biodiversity remains a stumbling block for conservation planning and even occurs within globally important Biodiversity Hotspots (BH). Although technical advances have boosted the power of molecular biodiversity assessments, the link between DNA sequences and species and the analytics to discriminate entities remain crucial. Here, we present an analysis of the first DNA barcode library for the freshwater fish fauna of the Mediterranean BH (526 spp.), with virtually complete species coverage (498 spp., 98% extant species). In order to build an identification system supporting conservation, we compared species determination by taxonomists to multiple clustering analyses of DNA barcodes for 3165 specimens. The congruence of barcode clusters with morphological determination was strongly dependent on the method of cluster delineation, but was highest with the general mixed Yule‐coalescent (GMYC) model‐based approach (83% of all species recovered as GMYC entity). Overall, genetic morphological discontinuities suggest the existence of up to 64 previously unrecognized candidate species. We found reduced identification accuracy when using the entire DNA‐barcode database, compared with analyses on databases for individual river catchments. This scale effect has important implications for barcoding assessments and suggests that fairly simple identification pipelines provide sufficient resolution in local applications. We calculated Evolutionarily Distinct and Globally Endangered scores in order to identify candidate species for conservation priority and argue that the evolutionary content of barcode data can be used to detect priority species for future IUCN assessments. We show that large‐scale barcoding inventories of complex biotas are feasible and contribute directly to the evaluation of conservation priorities.     

Landscape patterns of species-level association between ground-beetles and overstory trees in boreal forests of western Canada (Coleoptera, Carabidae)

Spatial associations between species of trees and ground-beetles (Coleoptera: Carabidae) involve many indirect ecological processes, likely reflecting the function of numerous forest ecosystem components. Describing and quantifying these associations at the landscape scale is basic to the development of a surrogate-based framework for biodiversity monitoring and conservation. In this study, we used a systematic sampling grid covering 84 km(2) of boreal mixedwood forest to characterize the ground-beetle assemblage associated with each tree species occurring on this landscape. Projecting the distribution of relative basal area of each tree species on the beetle ordination diagram suggests that the carabid community is structured by the same environmental factors that affects the distribution of trees, or perhaps even by trees per se. Interestingly beetle species are associated with tree species of the same rank order of abundance on this landscape, suggesting that conservation of less abundant trees will concomitantly foster conservation of less abundant beetle species. Landscape patterns of association described here are based on characteristics that can be directly linked to provincial forest inventories, providing a basis that is already available for use of tree species as biodiversity surrogates in boreal forest land management.     

Root location in random trees: A polarity property of all sampling consistent phylogenetic models except one

Neutral macroevolutionary models, such as the Yule model, give rise to a probability distribution on the set of discrete rooted binary trees over a given leaf set. Such models can provide a signal as to the approximate location of the root when only the unrooted phylogenetic tree is known, and this signal becomes relatively more significant as the number of leaves grows. In this short note, we show that among models that treat all taxa equally, and are sampling consistent (i.e. the distribution on trees is not affected by taxa yet to be included), all such models, except one (the so-called PDA model), convey some information as to the location of the ancestral root in an unrooted tree.     

Towards a theoretical basis for ecosystem conservation

Humans have altered the environment so severely that extinction events are now occurring at rates unprecedented in modern history. In order to slow this trend, conservation actions must be taken to protect biodiversity, beyond just saving flagship species. Some governmental and conservation organizations have responded by committing to ecosystem conservation but, as yet, there is no coherent strategy for how this can be carried out. This report introduces many of the theoretical aspects that will need to be considered for the development of a coherent ecosystem conservation policy. The approach includes analyzing a hierarchy of interaction-based local coexistence mechanisms within a regional and historical context. This approach points toward the need for prioritizing sensitive habitats using local interaction models; considering the effects of historical exploitation mechanisms, which are now often missing; and evaluating regional diversity influences further to identify circumstances where system-wide habitat improvements deserve more emphasis.     

Yule-generated trees constrained by node imbalance

The Yule   process generates a class of binary trees which is fundamental to population genetic models and other applications in evolutionary biology. In this paper, we introduce a family of sub-classes of ranked trees, called Ω-trees, which are characterized by imbalance of internal nodes. The degree of imbalance is defined by an integer 0≤ω$0\le \omega$. For caterpillars  , the extreme case of unbalanced trees, ω=0$\omega =0$. Under models of neutral evolution, for instance the Yule model, trees with small ω are unlikely to occur by chance. Indeed, imbalance can be a signature of permanent selection pressure, such as observable in the genealogies of certain pathogens. From a mathematical point of view it is interesting to observe that the space of Ω-trees maintains several statistical invariants although it is drastically reduced in size compared to the space of unconstrained Yule trees. Using generating functions, we study here some basic combinatorial properties of Ω-trees. We focus on the distribution of the number of subtrees with two leaves. We show that expectation and variance of this distribution match those for unconstrained trees already for very small values of ω.     

Using a goal programming approach to design and evaluate protected areas for the conservation of multiple dimensions of biodiversity

The decline and loss of biodiversity provoked by human activities have caused ecologists and conservationists to center their attention on the design of conservation priority areas (PAs), focusing mainly on species conservation in terms of richness, rarity and/or vulnerability. However, biodiversity has multiple dimensions, evolutionary processes have recently been labeled the ‘missing component’ of conservation strategies, and increasingly more authors are suggesting that the ecological, evolutionary and historical aspects of biodiversity are key components of conservation planning. In this study we develop a prioritization system to design conservation PAs using the wild terrestrial mammals of the Iberian Peninsula as an example. We aim to contribute to the design of more suitable PAs by integrating ecological components of biodiversity (species richness, vulnerability and rarity), evolutionary aspects (accumulated genetic diversification) and historical information relevant to the study area. After selecting a set of biodiversity indicators, we applied a multi-objective technique (extended goal programming) to construct a combined index, where values in the top 90th percentile were then used to select the PAs. According to our most efficient and satisfactory results, some areas highlighted for their conservation are currently categorized as PAs, however, we found that it would be necessary to reconsider their extent, especially in northern Spain, where the historical aspects of biodiversity (the missing component) are more widely present. The need to determine PAs is unquestionable. However, it should also be a priority to move towards a model of sustainable and fair development.     

Measuring and comparing the accuracy of species distribution models with presence–absence data

Species distribution models have been widely used to predict species distributions for various purposes, including conservation planning, and climate change impact assessment. The success of these applications relies heavily on the accuracy of the models. Various measures have been proposed to assess the accuracy of the models. Rigorous statistical analysis should be incorporated in model accuracy assessment. However, since relevant information about the statistical properties of accuracy measures is scattered across various disciplines, ecologists find it difficult to select the most appropriate ones for their research. In this paper, we review accuracy measures that are currently used in species distribution modelling (SDM), and introduce additional metrics that have potential applications in SDM. For the commonly used measures (which are also intensively studied by statisticians), including overall accuracy, sensitivity, specificity, kappa, and area and partial area under the ROC curves, promising methods to construct confidence intervals and statistically compare the accuracy between two models are given. For other accuracy measures, methods to estimate standard errors are given, which can be used to construct approximate confidence intervals. We also suggest that as general tools, computer‐intensive methods, especially bootstrap and randomization methods can be used in constructing confidence intervals and statistical tests if suitable analytic methods cannot be found. Usually, these computer‐intensive methods provide robust results.     

Stand-level management of plantations to improve biodiversity values

As conservation reserves expand, the likelihood that they will capture areas degraded by previous land use increases. Ecological restoration of such areas will therefore play an increasing role in biodiversity conservation. On the New South Wales North Coast, recent expansion in the conservation estate has captured over 300 softwood and hardwood plantations, many with understoreys dominated by exotic weeds. Here we present an overview of the practices we have adopted in managing flooded gum (Eucalyptus grandis) plantations infested with lantana (Lantana camara) to enhance their biodiversity value. Experiments designed to overcome barriers limiting regeneration of native forest in conjunction with measurement of soil and plant responses yielded insights into the management of former timber plantations for biodiversity. Canonical Correspondence Analysis indicated that the level of canopy retention (or logging intensity) within sites consistently explained the greatest amount of variation in plant community composition (32–38% post-treatment). Thinning and burning stimulated regeneration of native species. Retained canopy cover was proportional to the richness or abundance of native woody shrubs, understorey trees and native perennial herbs, indicating that management intensity can be varied to promote a range of conservation values. A state-and-transition model summarising purported management actions and likely outcomes for these plantations is presented. This is the first time plantations have been managed solely for biodiversity. Logging income means that plantation restoration can be cost-neutral, and the positive influence of a cover crop of trees means that plantation management may generally be manipulated to promote biodiversity conservation.     

Statistical Properties of Pairwise Distances between Leaves on a Random Yule Tree

A Yule tree is the result of a branching process with constant birth and death rates. Such a process serves as an instructive null model of many empirical systems, for instance, the evolution of species leading to a phylogenetic tree. However, often in phylogeny the only available information is the pairwise distances between a small fraction of extant species representing the leaves of the tree. In this article we study statistical properties of the pairwise distances in a Yule tree. Using a method based on a recursion, we derive an exact, analytic and compact formula for the expected number of pairs separated by a certain time distance. This number turns out to follow a increasing exponential function. This property of a Yule tree can serve as a simple test for empirical data to be well described by a Yule process. We further use this recursive method to calculate the expected number of the n-most closely related pairs of leaves and the number of cherries separated by a certain time distance. To make our results more useful for realistic scenarios, we explicitly take into account that the leaves of a tree may be incompletely sampled and derive a criterion for poorly sampled phylogenies. We show that our result can account for empirical data, using two families of birds species.