Putting the Biological Species Concept to the Test: Using Mating Networks to Delimit Species

Although interfertility is the key criterion upon which Mayr’s biological species concept is based, it has never been applied directly to delimit species under natural conditions. Our study fills this gap. We used the interfertility criterion to delimit two closely related oak species in a forest stand by analyzing the network of natural mating events between individuals. The results reveal two groups of interfertile individuals connected by only few mating events. These two groups were largely congruent with those determined using other criteria (morphological similarity, genotypic similarity and individual relatedness). Our study, therefore, shows that the analysis of mating networks is an effective method to delimit species based on the interfertility criterion, provided that adequate network data can be assembled. Our study also shows that although species boundaries are highly congruent across methods of species delimitation, they are not exactly the same. Most of the differences stem from assignment of individuals to an intermediate category. The discrepancies between methods may reflect a biological reality. Indeed, the interfertility criterion is an environment-dependant criterion as species abundances typically affect rates of hybridization under natural conditions. Thus, the methods of species delimitation based on the interfertility criterion are expected to give results slightly different from those based on environment-independent criteria (such as the genotypic similarity criteria). However, whatever the criterion chosen, the challenge we face when delimiting species is to summarize continuous but non-uniform variations in biological diversity. The grade of membership model that we use in this study appears as an appropriate tool.     

Is the Species Flock Concept Operational? The Antarctic Shelf Case

There has been a significant body of literature on species flock definition but not so much about practical means to appraise them. We here apply the five criteria of Eastman and McCune for detecting species flocks in four taxonomic components of the benthic fauna of the Antarctic shelf: teleost fishes, crinoids (feather stars), echinoids (sea urchins) and crustacean arthropods. Practical limitations led us to prioritize the three historical criteria (endemicity, monophyly, species richness) over the two ecological ones (ecological diversity and habitat dominance). We propose a new protocol which includes an iterative fine-tuning of the monophyly and endemicity criteria in order to discover unsuspected flocks. As a result nine « full » species flocks (fulfilling the five criteria) are briefly described. Eight other flocks fit the three historical criteria but need to be further investigated from the ecological point of view (here called « core flocks »). The approach also shows that some candidate taxonomic components are no species flocks at all. The present study contradicts the paradigm that marine species flocks are rare. The hypothesis according to which the Antarctic shelf acts as a species flocks generator is supported, and the approach indicates paths for further ecological studies and may serve as a starting point to investigate the processes leading to flock-like patterning of biodiversity.     

浅析物种概念的演变历史

本文是一篇关于物种概念演变的简述。生物学家用不同的方法或标准划分物种, 就形成了不同的物种概念, 如生物学物种、形态学物种、生态学物种、进化物种、系统发生或支序物种, 或它们的组合, 等等。它们都揭示了物种属性的特定侧面, 都是不同物种客观存在的真实反映, 但都无法令所有人满意。对真核生物来说, 无论它们在形态上的差别有多大, 生殖隔离(不能产生可育的后代)应该是两个群体能否真正分化成不同物种的关键, 这种隔离机制可以是地理的、行为的或其他方式; 而生殖隔离总会伴随着一些形态或遗传上的变化, 虽然这些特征可能与生殖隔离本身并无多大关系, 但往往成为分类学家或分子进化生物学家区分种的依据,对已经灭绝的化石物种来说, 生殖隔离的物种划分方式就无能为力了。如何准确定义一个物种依然充满着矛盾, 因为基于生殖隔离的物种概念不实用, 而实用的物种概念(如形态学物种)又被认为是人为的。     

Assessment of Rarity Category for Higher Aquatic Plants

A spatiotemporal analysis of higher aquatic plant flora in the water bodies of the city of Kyiv (Ukraine) has been used to develop local protection criteria for 22 rare and protected species. These criteria differ considerably from the regional ones due to the significant transformation of the environment in an urban landscape. The rare component of macrophyte flora in the water bodies of the city have become poorer over the past 40–50 years; two species have disappeared from the urban flora, seven species are characterized as critically endangered, five species are endangered, four are vulnerable and at a high extinction risk, and four more species are characterized by a low risk of extinction.     

Generic ecological impact assessments of alien species in Norway: a semi-quantitative set of criteria

The ecological impact assessment scheme that has been developed to classify alien species in Norway is presented. The underlying set of criteria enables a generic and semi-quantitative impact assessment of alien species. The criteria produce a classification of alien species that is testable, transparent and easily adjustable to novel evidence or environmental change. This gives a high scientific and political legitimacy to the end product and enables an effective prioritization of management efforts, while at the same time paying attention to the precautionary principle. The criteria chosen are applicable to all species regardless of taxonomic position. This makes the assessment scheme comparable to the Red List criteria used to classify threatened species. The impact of alien species is expressed along two independent axes, one measuring invasion potential, the other ecological effects. Using this two-dimensional approach, the categorization captures the ecological impact of alien species, which is the product rather than the sum of spread and effect. Invasion potential is assessed using three criteria, including expected population lifetime and expansion rate. Ecological effects are evaluated using six criteria, including interactions with native species, changes in landscape types, and the potential to transmit genes or parasites. Effects on threatened species or landscape types receive greater weightings.     

Prioritizing species for conservation planning

The efforts to protect biological diversity must be prioritized because resources for nature conservation are limited. Conservation prioritization can be based on numerous criteria, from ecological integrity to species representation, but in this review I address only species-level prioritization. Criteria used for species prioritization range from aesthetical to evolutionary considerations, but I focus on the aspects that are biologically relevant. I distinguish between two main aspects of diversity that are used as objectives: Maintenance of biodiversity pattern, and maintenance of biodiversity process. I identify two additional criteria typically used in species prioritization that serve for achieving the objectives: The species’ need of protection, and cost and effectiveness of conservation actions. I discuss how these criteria could be combined with either of the objectives in a complementarity-based benefit function framework for conservation prioritization. But preserving evolutionary process versus current diversity pattern may turn out to be conflicting objectives that have to be traded-off with each other, if pursued simultaneously. Although many reasonable criteria and methods exist, species prioritization is hampered by uncertainties, most of which stem from the poor quality of data on what species exist, where they occur, and what are the costs and benefits of protecting them. Surrogate measures would be extremely useful but their performance is still largely unknown. Future challenges in species prioritization lie in finding ways to compensate for missing information.     

Descriptions and phylogenetic systematics of Myxobolus spp. from cyprinids in Algonquin Park, Ontario

Eight species of Myxobolus were collected from four species of cyprinids in Algonquin Park, Ontario. On the basis of spore morphology, five of these species are described as new and two are redescribed. The evolutionary relationships among these eight species were studied using partial small subunit ribosomal DNA (ssu-rDNA) sequence data. The resulting cladograms, which were highly resolved and with strongly supported relationships, allowed for the evaluation of spore morphology, host specificity, and tissue tropism, criteria traditionally used in species identification. These criteria, recently criticized for creating artificial rather than natural taxonomic groupings, were evaluated for their reliability in the systematics of the species examined. The data showed that distantly related species often infect the same host and tissue, and that closely related species often occur in different hosts. Morphologically similar species are more closely related to each other and the taxonomy based on spore morphology is consistent with the relationships depicted in the phylogenies. These results suggest that spore morphology is better than host specificity and tissue tropism as a species character, as well as for determining evolutionary relationships among the species of Myxobolus examined.     

The cladistic solution to the species problem

The correct explanation of why species, in evolutionary theory, are individuals and not classes is the cladistic species concept. The cladistic species concept defines species as the group of organisms between two speciation events, or between one speciation event and one extinction event, or (for living species) that are descended from a speciation event. It is a theoretical concept, and therefore has the virtue of distinguishing clearly the theoretical nature of species from the practical criteria by which species may be recognized at any one time. Ecological or biological (reproductive) criteria may help in the practical recognition of species. Ecological and biological species concepts are also needed to explain why cladistic species exist as distinct lineages, and to explain what exactly takes place during a speciation event. The ecological and biological species concepts work only as sub-theories of the cladistic species concept and if taken by themselves independently of cladism they are liable to blunder. The biological species concept neither provides a better explanation of species indivudualism than the ecological species concept, nor, taken by itself, can the biological species concept even be reconciled with species individualism. Taking the individuality of species seriously requires subordinating the biological, to the cladistic, species concept.     

Species concepts and species delimitation

The issue of species delimitation has long been confused with that of species conceptualization, leading to a half century of controversy concerning both the definition of the species category and methods for inferring the boundaries and numbers of species. Alternative species concepts agree in treating existence as a separately evolving metapopulation lineage as the primary defining property of the species category, but they disagree in adopting different properties acquired by lineages during the course of divergence (e.g., intrinsic reproductive isolation, diagnosability, monophyly) as secondary defining properties (secondary species criteria). A unified species concept can be achieved by treating existence as a separately evolving metapopulation lineage as the only necessary property of species and the former secondary species criteria as different lines of evidence (operational criteria) relevant to assessing lineage separation. This unified concept of species has several consequences for species delimitation, including the following: First, the issues of species conceptualization and species delimitation are clearly separated; the former secondary species criteria are no longer considered relevant to species conceptualization but only to species delimitation. Second, all of the properties formerly treated as secondary species criteria are relevant to species delimitation to the extent that they provide evidence of lineage separation. Third, the presence of any one of the properties (if appropriately interpreted) is evidence for the existence of a species, though more properties and thus more lines of evidence are associated with a higher degree of corroboration. Fourth, and perhaps most significantly, a unified species concept shifts emphasis away from the traditional species criteria, encouraging biologists to develop new methods of species delimitation that are not tied to those properties.     

Comparing Multiple Criteria for Species Identification in Two Recently Diverged Seabirds

Correct species identification is a crucial issue in systematics with key implications for prioritising conservation effort. However, it can be particularly challenging in recently diverged species due to their strong similarity and relatedness. In such cases, species identification requires multiple and integrative approaches. In this study we used multiple criteria, namely plumage colouration, biometric measurements, geometric morphometrics, stable isotopes analysis (SIA) and genetics (mtDNA), to identify the species of 107 bycatch birds from two closely related seabird species, the Balearic (Puffinus mauretanicus) and Yelkouan (P. yelkouan) shearwaters. Biometric measurements, stable isotopes and genetic data produced two stable clusters of bycatch birds matching the two study species, as indicated by reference birds of known origin. Geometric morphometrics was excluded as a species identification criterion since the two clusters were not stable. The combination of plumage colouration, linear biometrics, stable isotope and genetic criteria was crucial to infer the species of 103 of the bycatch specimens. In the present study, particularly SIA emerged as a powerful criterion for species identification, but temporal stability of the isotopic values is critical for this purpose. Indeed, we found some variability in stable isotope values over the years within each species, but species differences explained most of the variance in the isotopic data. Yet this result pinpoints the importance of examining sources of variability in the isotopic data in a case-by-case basis prior to the cross-application of the SIA approach to other species. Our findings illustrate how the integration of several methodological approaches can help to correctly identify individuals from recently diverged species, as each criterion measures different biological phenomena and species divergence is not expressed simultaneously in all biological traits.     

A new methodology for compiling national Red Lists applied to butterflies (Lepidoptera, Rhopalocera) in Flanders (N-Belgium) and the Netherlands

The compilation of the Red Lists of butterflies in Flanders and the Netherlands was based on two criteria: a trend criterion (degree of decline) and a rarity criterion (actual distribution area). However, due to the large difference in mapping intensity in the two compared periods, a straightforward comparison of the number of grid cells in which each species was recorded, appeared inappropriate. To correct for mapping intensity we used reference species that are homogeneously distributed over the country, that have always been fairly common and that did not fluctuate in abundance too much during this century. For all resident species a relative presence in two compared periods was calculated, using the average number of grid cells in which these reference species were recorded as a correction factor. The use of a standardized method and well-defined quantitative criteria makes national Red Lists more objective and easier to re-evaluate in the future and facilitates the comparison of Red Lists among countries and among different organisms. The technique applied to correct for mapping intensity could be useful to other organisms when there is a large difference in mapping intensity between two periods.     

Comparison of Biological,Molecular, and Morphological Methods of Species Identification in a Set of Cultured Panagrolaimus Isolates

We have developed a molecular barcode system that uses the small subunit ribosomal RNA (SSU) sequence to define molecular operational taxonomic units (MOTU) of soil nematodes. Here we attempt to differentiate five cultured isolates of a taxonomically difficult genus, Panagrolaimus, using morphological, molecular, and biological (breeding) criteria. The results indicated that the five culture populations belonged to two reproductively isolated species. The available morphological criteria, including scanning electron microscopy (SEM), were insufficient to differentiate among them, and all five could be classified as one morphospecies. Within-culture variation of the morphometrical data did not discern between the two biological species. Sequence data clearly separated the populations into two groups that supported the breeding results. Given this study represented only five populations of one genus, we suggest a congruence of MOTU analysis with the biological species concept. This multifaceted approach is promising for future identification of nematodes as it is simple, comparable, and transferable.     

物种概念及其界定

半世纪以来,物种概念的定义备受关注,不同研究方向的生物学家提出24种不同或至少有分歧的物种概念,根据其不同的物种概念,物种的界定和物种的数量会出现很大的差异。人们普遍认同:物种是进化分离的微居群谱系,但把谱系分离过程中获得的特征如生殖隔离、可鉴定性、单系统发生等视为鉴定物种次级特征却有不同的声音。该文提出统一的物种概念,把谱系进化分离作为物种界定的唯一而又必要的特征,把谱系分离过程中获得的次级特征作为界定谱系分离的证据。鉴于此,物种概念间的分歧就会化解。其一,物种概念化与物种界定明显分开,不再混淆;其二,谱系的次级特征只与物种界定有关,在某种程度上为谱系分离提供证据;第三,若能把合理解释的任何一个特征作为某物种客观存在的证据,这样更多的特征更能确定谱系分离;最后最重要的是,统一物种概念使我们解放思想,扬弃传统的物种界定标准,探求物种界定的新思路。     

A molecular approach to the phylogeny of Acanthamoeba

Isoenzyme electrophoresis of three different enzyme systems was used to compare 71 strains assigned to the 15 currently recognized species of Acanthamoeba. A phylogenetic (cladistic) analysis of the zymograms indicated an arrangement of strains in 15 distinguishable lineages, but not all corresponding to current taxonomic assignments. Five of the groups corresponded to the recognized species A. castellanii, A. culbertsoni, A. griffini, A. lenticulata and A. royreba. But none of these groups consisted of only strains which had been previously assigned to each respective species. The type-equivalent strains for two species, A. hatchetti and A. tubiashi, were not closely aligned to any other strain and thus are considered to be monotypic. Strains of A. triangularis, A. astronyxis and A. palestinensis occurred together in a single group suggesting possible synonymy; however, on morphologic criteria, the strains assigned to these species are readily distinguishable. Strains assigned to A. polyphaga and A. rhysodes were interspersed throughout the other species groups. The strains of these two species were either misidentified or the species could not be recognized. Two groups previously not recognized as unique formed distinctive clusters which could be considered as new species. The analysis also made it possible to place strains which had previously been identified only to genus into species complexes. These results therefore suggest that previous criteria which have been used to classify Acanthamoeba are not adequate for fully resolving taxa at the species level.     

Validation of grassland indicator species selected for result-oriented agri-environmental schemes

A list of plant species indicating species-rich grassland was validated for use in an agri-environmental programme in the Federal State of Brandenburg (Germany). It has become common practice that suggested lists of indicator species pass through a process of coordination involving several experts and stakeholders. Therefore, an ‘aptitude test’ for indicator species was needed that would be transparent as well as comprehensible. The starting list of potentially suitable indicator species was filtered using criteria that removed rare, toxic and ubiquitous species and retained representative species relevant for extensively managed grassland habitats of Brandenburg and species that are easy to detect and identify. Vegetation surveys conducted on 50-m transects in about 120 grassland fields in Brandenburg served as the database for the validation. The data were grouped by the mean moisture levels of the sites. We examined the influence of the presence or absence of potentially suitable indicator species on diversity quality criteria for each site group using a single species testing procedure. Besides floristic species diversity (overall species number), the number of species indicating extensive use and the number of Red List species are criteria that can be used. In addition, weighted indices were computed which pool together the three quality criteria to establish a ranking of the pre-selected species. The quality criteria can be differentially combined depending on the needs of the individual project.     

On areas of endemism,with an example from the African Restionaceae

Areas of endemism are central to cladistic biogeography. The concept has been much debated in the past, and from this has emerged the generally accepted definition as an area to which at least two species are endemic. Protocols for locating areas of endemism have been neglected, and to date no attempt has been made to develop optimality criteria against which to evaluate competing hypotheses of areas of endemism. Here various protocols for finding areas of endemism are evaluated--protocols based on both phonetic and parsimony analyses, on both unweighted data and data weighted by various criteria. The optimality criteria used to compare the performance of the methods include the number of species included in the areas of endemism, the number of areas delimited, and the degree of distributional congruency of the species restricted to each area of endemism. These methods are applied to the African Restionaceae in the Cape Floristic Region. Parsimony methods using weighted data are shown to perform best on the combination of all three optimality criteria. By varying the weighting parameters, the size of the areas of endemism can be varied. This provides a very useful tool for locating areas of endemism that satisfy prespecified scale criteria.     

Using taxonomically unbiased criteria to prioritize resource allocation for oceanic island species conservation

Oceanic islands have been the grand stage of documented extinctions. In view of limited resources, efficient prioritization is crucial to avoid the extinction of taxa. This work lists the top 100 management priority species for the European archipelagos of the Macaronesian region (Azores, Madeira and the Canary Islands), taking into account both their protection priority and their management feasibility. Bryophytes, vascular plants, molluscs, arthropods and vertebrates were scored by species experts following two sets of criteria: (i) protection priority, including ecological value, singularity, public institutions’ management responsibilities and social value; (ii) management feasibility, including threats knowledge and control feasibility, external socio-economical support for management and biological recovery potential. Environmental managers weighted the same criteria according to their management importance. Final species scores were determined by the combination of both species valuation and criteria weighting. Vascular plants dominate the Top 100 list, followed by arthropods and vertebrates. The majority of listed taxa are endemic to one archipelago or even to a single island. The management feasibility criteria did not dictate that all taxa must be eminently endangered, as for most of the species it should be relatively easy to control threats. The main advantages of this process are the independent participation of scientists and conservation managers, the inclusion of criteria on both protection priority and management feasibility and the taxonomically unbiased nature of the process. This study provides a potentially useful biodiversity conservation tool for the Macaronesian archipelagos that could be readily implemented by the respective regional governments in future legislation.     

贵州省两栖动物生存状况评价及其保护需求初探

针对我国野生动物保护的实情,为便于各省区有关部门制订野生濒危动物的保护等级,通过对国内外物种濒危等级标准的全面分析,建议省级保护物种濒危等级的数量标准为:濒危级En分布面积为1个县;易危级T分布面积小于3个县;关注级C分布面积小于5个县.结合贵州省两栖动物已有文献资料数据及作者近三十年来野外考察数据,按上述指标对贵州省已知的72种两栖动物的濒危等级进行了评估.