Ceci n'est pas une pipe: names, clades and phylogenetic nomenclature

An introduction is provided to the literature and to issues relating to phylogenetic nomenclature and the PhyloCode, together with a critique of the current Linnaean system of nomenclature. The Linnaean nomenclature fixes taxon names with types, and associates the names with ranks (genus, family, etc.). In phylogenetic nomenclature, names are instead defined with reference to cladistic relationships, and the names are not associated with ranks. We argue that taxon names under the Linnaean system are unclear in meaning and provide unstable group–name associations, notwithstanding whether or not there are agreements on relationships. Furthermore, the Linnaean rank assignments lack justification and invite unwarranted comparisons across taxa. On the contrary, the intention of taxon names in phylogenetic nomenclature is clear and stable, and the application of the names will be unambiguous under any given cladistic hypothesis. The extension of the names reflects current knowledge of relationships, and will shift as new hypotheses are forwarded. The extension of phylogenetic names is, therefore, clear but is associated to (and thus dependent upon) cladistic hypotheses. Stability in content can be maximized with carefully formulated name definitions. A phylogenetic nomenclature will shift the focus from discussions of taxon names towards the understanding of relationships. Also, we contend that species should not be recognized as taxonomic units. The term ‘species’ is ambiguous, it mixes several distinct classes of entities, and there is a large gap between most of the actual concepts and the evidence available to identify the entities. Instead, we argue that only clades should be recognized. Among these, it is useful to tag the smallest named clades, which all represent non-overlapping groups. Such taxa  – LITUs (Least Inclusive Taxonomic Units) – are distinguished from more inclusive clades by being spelled with lower-case initial letter. In contrast to species, LITUs are conceptually straightforward and are, like other clades, identified by apomorphies.     

Stems,nodes, crown clades,and rank‐free lists: is Linnaeus dead?

Recent radical proposals to overhaul the methods of biological classification are reviewed. The proposals of phylogenetic nomenclature are to translate cladistic phylogenies directly into classifications, and to define taxon names in terms of clades. The method has a number of radical consequences for biologists: taxon names must depend rigidly on the particular cladogram favoured at the moment, familiar names may be reassigned to unfamiliar groupings, Linnaean category terms (e.g. phylum, order, family) are abandoned, and the Linnaean binomen (e.g. Homo sapiens) is abandoned. The tenets of phylogenetic nomenclature have gained strong support among some vocal theoreticians, and rigid principles for legislative control of clade names and definitions have been outlined in the PhyloCode. The consequences of this semantic maelstrom have not been worked out. In pratice, phylogenetic nomenclature will bc disastrous, promoting confusion and instability, and it should be abandoned. It is based on a fundamental misunderstanding of the difference between a phylogeny (which is real) and a classification (which is utilitarian). Under the new view, classifications are identical to phlylogenies, and so the proponents of phylogenetic nomenclature will end up abandoning classifications altogether.     

Classification of apocynaceae s.l. according to a new approach combining Linnaean and phylogenetic taxonomy

A new approach to a nomenclatural system, including elements from both Linnaean and phylogenetic nomenclature, is proposed. It is compatible with the existing Linnaean system, including "standard names" corresponding to principal and secondary ranks, and uses a variant of the definitions from the Phylocode system. A new infrafamilial classification, using this nomenclatural approach, of the Apocynaceae s.l. (i.e., including the Asclepiadaceae) based mainly on analyses of rbcL and ndhF data is discussed. Twenty-one tribes and four rankless taxa are defined.     

Should taxon names be explicitly defined?

Overviews are provided for traditional and phylogenetic nomenclature. In traditional nomenclature, a name is provided with a type and a rank. In the rankless phylogenetic nomenclature, a taxon name is provided with an explicit phylogenetic definition, which attaches the name to a clade. Linnaeus’s approach to nomenclature is also reviewed, and it is shown that, although the current system of nomenclature does use some Linnaean conventions (e.g., certain rank-denoting terms, binary nomenclature), it is actually quite different from Linnaean nomenclature. The primary differences between traditional and phylogenetic nomenclature are reviewed. In phylogenetic nomenclature, names are provided with explicit phylogenetic definitions, whereas in traditional nomenclature names are not explicitly defined. In phylogenetic nomenclature, a name remains attached to a clade regardless of how future changes in phylogeny alter the clade’s content; in traditional nomenclature a name is not “married” to any particular clade. In traditional nomenclature, names must be assigned ranks (an admittedly arbitrary process), whereas in phylogenetic nomenclature there are no formal ranks. Therefore, in phylogenetic nomenclature, the name itself conveys no hierarchical information, and the name conveys nothing regarding set exclusivity. It is concluded that the current system is better able to handle new and unexpected changes in ideas about taxonomic relationships. This greater flexibility, coupled with the greater information content that the names themselves (i.e., when used outside the context of a given taxonomy or phytogeny) provide, makes the current system better designed for use by all users of taxon names.     

Forum – Taxonomic Stability is Ignorance

Absolute nomenclatural stability is undesirable in phylogenetic classifications because they reflect changing hypotheses of cladistic relationships. De Queiroz and Gauthier's (1990: Syst. Zool. 39, 307–322; 1992: A. Rev. Ecol. Syst. 23, 449–480; 1994: Trends Ecol. Evol. 9, 27–31) alternative to Linnaean nomenclature is concluded to provide stable names for unstable concepts. In terms of communicating either characters shared by species of a named taxon or elements (species) included in a taxon, de Queiroz and Gauthier's system is less stable than the Linnaean system. Linnaean ranks communicate limited information about inclusivity of taxa, but abandonment of ranks results in the loss of such information. As cladistic hypotheses advance, taxa named under de Queiroz and Gauthier's system can change their level of generality radically, from being part of a group to including it, without any indicative change in its spelling. The Linnaean system has been retained by taxonomists because its hierarchic ranks are logically compatible with nested sets of species, monophyletic groups, and characters. Other authors have offered conventions to increase the cladistic information content of Linnaean names or to replace them with names that convey cladistic knowledge in greater detail; de Queiroz and Gauthier sacrifice the meaning of taxon names and categorical ranks in favor of spelling stability.     

Exploring the tempo of species diversification in legumes

Whatever criteria are used to measure evolutionary success – species numbers, geographic range, ecological abundance, ecological and life history diversity, background diversification rates, or the presence of rapidly evolving clades – the legume family is one of the most successful lineages of flowering plants. Despite this, we still know rather little about the dynamics of lineage and species diversification across the family through the Cenozoic, or about the underlying drivers of diversification. There have been few attempts to estimate net species diversification rates or underlying speciation and extinction rates for legume clades, to test whether among-lineage variation in diversification rates deviates from null expectations, or to locate species diversification rate shifts on specific branches of the legume phylogenetic tree. In this study, time-calibrated phylogenetic trees for a set of species-rich legume clades – Calliandra, Indigofereae, Lupinus, Mimosa and Robinieae – and for the legume family as a whole, are used to explore how we might approach these questions. These clades are analysed using recently developed maximum likelihood and Bayesian methods to detect species diversification rate shifts and test for among-lineage variation in speciation, extinction and net diversification rates. Possible explanations for rate shifts in terms of extrinsic factors and/or intrinsic trait evolution are discussed. In addition, several methodological issues and limitations associated with these analyses are highlighted emphasizing the potential to improve our understanding of the evolutionary dynamics of legume diversification by using much more densely sampled phylogenetic trees that integrate information across broad taxonomic, geographical and temporal levels.     

A higher-level taxonomy for hummingbirds

In the context of a recently published phylogenetic estimate for 151 hummingbird species, we provide an expanded informal taxonomy, as well as a formal phylogenetic taxonomy for Trochilidae that follows the precepts of the PhyloCode, but remains consistent with the hierarchical nomenclature of the Linnaean system. We compare the recently published phylogenetic hypothesis with those of prior higher-level and more taxonomically circumscribed phylogenetic studies. We recommend the recognition of nine new clade names under the PhyloCode, eight of which are consistent with tribes and one with a subfamily under the Linnaean system.     

A framework infrageneric classification of Carex (Cyperaceae) and its organizing principles

Phylogenetic studies of Carex L. (Cyperaceae) have consistently demonstrated that most subgenera and sections are para- or polyphyletic. Yet, taxonomists continue to use subgenera and sections in Carex classification. Why? The Global Carex Group (GCG) here takes the position that the historical and continued use of subgenera and sections serves to (i) organize our understanding of lineages in Carex, (ii) create an identification mechanism to break the ~2000 species of Carex into manageable groups and stimulate its study, and (iii) provide a framework to recognize morphologically diagnosable lineages within Carex. Unfortunately, the current understanding of phylogenetic relationships in Carex is not yet sufficient for a global reclassification of the genus within a Linnean infrageneric (sectional) framework. Rather than leaving Carex classification in its current state, which is misleading and confusing, we here take the intermediate steps of implementing the recently revised subgeneric classification and using a combination of informally named clades and formally named sections to reflect the current state of our knowledge. This hybrid classification framework is presented in an order corresponding to a linear arrangement of the clades on a ladderized phylogeny, largely based on the recent phylogenies published by the GCG. It organizes Carex into six subgenera, which are, in turn, subdivided into 62 formally named Linnean sections plus 49 informal groups. This framework will serve as a roadmap for research on Carex phylogeny, enabling further development of a complete reclassification by presenting relevant morphological and geographical information on clades where possible and standardizing the use of formal sectional names.     

生物谱系命名法规简评

基于林奈命名法和林奈分类系统的生物分类系统已经存在250多年并仍然为广大生物学工作者使用,由此产生的国际动物、植物、细菌的命名法规亦执行了100年(1905年,国际植物命名法规第1版产生),并在不断修订.随着分类方法的不断进步,林奈分类系统的一些缺陷逐渐显露,一种被称为生物谱系命名法规(PhyloCode)的新的命名法出现在人们眼前.这种基于系统发育系统学的命名法规一经问世就引起诸多争论,但是,作为一种新的命名法规,无论与传统的命名法规融合还是独立发展,对于已有的分类系统都是一个新的机遇和挑战.     

Towards a new classification system for legumes: Progress report from the 6th International Legume Conference

Legume systematists have been making great progress in understanding evolutionary relationships within the Leguminosae (Fabaceae), the third largest family of flowering plants. As the phylogenetic picture has become clearer, so too has the need for a revised classification of the family. The organization of the family into three subfamilies and 42 tribes is outdated and evolutionarily misleading. The three traditionally recognized subfamilies, Caesalpinioideae, Mimosoideae, and Papilionoideae, do not adequately represent relationships within the family. The occasion of the Sixth International Legume Conference in Johannesburg, South Africa in January 2013, with its theme “Towards a new classification system for legumes,” provided the impetus to move forward with developing a new classification. A draft classification, based on current phylogenetic results and a set of principles and guidelines, was prepared in advance of the conference as the basis for discussion. The principles, guidelines, and draft classification were presented and debated at the conference. The objectives of the discussion were to develop consensus on the principles that should guide the development of the classification, to discuss the draft classification's strengths and weaknesses and make proposals for its revision, and identify and prioritize phylogenetic deficiencies that must be resolved before the classification could be published. This paper describes the collaborative process by a large group of legume systematists, publishing under the name Legume Phylogeny Working Group, to develop a new phylogenetic classification system for the Leguminosae. The goals of this paper are to inform the broader legume community, and others, of the need for a revised classification, and spell out clearly what the alternatives and challenges are for a new classification system for the family.     

Second Meeting of the International Society for Phylogenetic Nomenclature: a Report

The Second Meeting of the International Society for Phylogenetic Nomenclature (ISPN) convened at Yale University in New Haven from June 28 to July 2, 2006. In addition to contributed talks, the conference included symposia on phylogenetic nomenclature of species, phyloinformatics, and implementing phylogenetic nomenclature. Other discussion focused on recent controversial additions to the draft PhyloCode concerning the choice of names for total clades, and the Committee on Phylogenetic Nomenclature (CPN) was encouraged to revisit this issue. A proposal to permit emendation of phylogenetic definitions without CPN approval under certain circumstances was well received, and there was wide support for a proposed mechanism to use Linnaean binomina in the context of phylogenetic nomenclature without extending the PhyloCode to govern species names. The ISPN Council voted to expand the CPN from 9 to 12 members.     

Naming diversity in an evolutionary context: Phylogenetic definitions of the Roucela clade (Campanulaceae/Campanuloideae) and the cryptic taxa within

In recent times, evolution has become a central tenet of taxonomy, but nomenclature has consistently been decoupled from the tree‐thinking process, often leading to significant issues in reconciling traditional (Linnaean) names with clades in the Tree of Life. Recent evolutionary studies on the Roucela clade, a group of endemic plants found in the Mediterranean Basin, motivated the establishment of phylogenetic concepts to formally anchor clade names on the Campanuloideae (Campanulaceae) tree. These concepts facilitate communication of clades that approximate traditionally defined groups, in addition to naming newly discovered cryptic diversity in a phylogenetic framework.     

Species names in phylogenetic nomenclature

Linnaean binomial nomenclature is logically incompatible with the phylogenetic nomenclature of de Queiroz and Gauthier (1992, Annu. Rev. Ecol. Syst. 23:449-480): The former is based on the concept of genus, thus making this rank mandatory, while the latter is based on phylogenetic definitions and requires the abandonment of mandatory ranks. Thus, if species are to receive names under phylogenetic nomenclature, a different method must be devised to name them. Here, 13 methods for naming species in the context of phylogenetic nomenclature are contrasted with each other and with Linnaean binomials. A fundamental dichotomy among the proposed methods distinguishes those that retain the entire binomial of a preexisting species name from those that retain only the specific epithet. Other relevant issues include the stability, uniqueness, and ease of pronunciation of species names; their capacity to convey phylogenetic information; and the distinguishability of species names that are governed by a code of phylogenetic nomenclature both from clade names and from species names governed by the current codes. No method is ideal. Each has advantages and drawbacks, and preference for one option over another will be influenced by one's evaluation of the relative importance of the pros and cons for each. Moreover, sometimes the same feature is viewed as an advantage by some and a drawback by others. Nevertheless, all of the proposed methods for naming species in the context of phylogenetic nomenclature provide names that are more stable than Linnaean binomials.     

Names,numbers and indentations: a guide to post-Linnaean taxonomy

The vast majority of biological taxonomists use the Linnaean system when constructing classifications. Taxa are assigned Linnaean ranks and taxon names are devised according to the Linnaean rules of nomenclature. Unfortunately, the Linnaean system has become theoretically outdated. Moreover, its continued use causes a number of practical problems. This paper begins by sketching the ontological and practical problems facing the Linnaean system. Those problems are sufficiently pressing that alternative systems of classification should be investigated. A number of proposals for an alternative system are introduced and evaluated. The best aspects of those proposals are brought together to form a post-Linnaean system, and a comparison of the Linnaean and post-Linnaean systems is conducted. The final section of this paper considers not only the theoretical reasons for replacing the Linnaean system, but also the practical feasibility of adopting an alternative system.     

The Evolution of the Linnaean Hierarchy

The Linnaean system of classification is a threefold system of theoretical assumptions, sorting rules, and rules of nomenclature. Over time, that system has lost its theoretical assumptions as well as its sorting rules. Cladistic revisions have left it less and less Linnaean. And what remains of the system is flawed on pragmatic grounds. Taking all of this into account, it is time to consider alternative systems of classification.     

On the lack of good scientific reasons for the growing phylogeny/classification gap

An increasing number of phylogenetic analyses is no longer translated into classifications. The resulting phylogeny/classification gap is undesirable because the precise transmission of phylogenetic insights depends on the frequent revision of Linnaean names. The move away from classifying has numerous correlates. These include: an expanded pool of researchers who are able to produce phylogenetic estimates, a mismatch between the properties of molecular phylogenies and the requirements for verbal Linnaean definitions, the emphasis of statistical representations over the creation and evaluation of scientific terms, and a partial disconnect between the processes of nomenclature and taxonomy. The “taxonomic concept” approach allows systematists to express their varying perspectives with a high precision and can therefore help reduce the aforementioned gap. The phylogeny/classification link must persist in order to ensure community‐wide access to, and continued testing of, the products of systematic research. © The Willi Hennig Society 2005.     

Is the Neotropical flora ready for the PhyloCode?

Nomenclatural systems are structured around classification, and together they enable increasingly informed communication about biological diversity. Challengers of Linnaean classification and nomenclature have proposed the PhyloCode, a new set of rules that would govern the way systematists classify and name the diversity of life. Monographs and floras are two fundamental vehicles for communicating information about plant diversity. These works provide a comprehensive foundation of botanical research upon which other scientific studies are based. Information conveyed by monographs and floras is utilized directly or indirectly both within and outside the scientific arena by a wide range of consumers, such as educators, agronomists, ecologists, conservationists, amateur naturalists, and even lawmakers, to name a few. Both classification and nomenclature are essential to the process of synthesis that leads to monographic and floristic treatments and the communication that they facilitate. Conversion to a new system would have far-reaching consequences for the flow of information from systematics to other scientific disciplines, and to society. The purposes of this article are to address the proposed conversion from the perspective of monographic and floristic research focused on Neotropical plant diversity and to point out some difficulties in applying the PhyloCode to the Neotropical flora. Although we welcome improvements in the current nomenclatural system, we conclude that the PhyloCode is not prepared to replace the Linnaean system as a new way to communicate information about Neotropical plant diversity.     

Least-inclusive taxonomic unit: a new taxonomic concept for biology

Phylogenetic taxonomy has been introduced as a replacement for the Linnaean system. It differs from traditional nomenclature in defining taxon names with reference to phylogenetic trees and in not employing ranks for supraspecific taxa. However, 'species' are currently kept distinct. Within a system of phylogenetic taxonomy we believe that taxon names should refer to monophyletic groups only and that species should not be recognized as taxa. To distinguish the smallest identified taxa, we here introduce the least-inclusive taxonomic unit (LITU), which are differentiated from more inclusive taxa by initial lower-case letters. LITUs imply nothing absolute about inclusiveness, only that subdivisions are not presently recognized.     

浅评当今植物系统学中争论的三个问题——并系类群、谱系法规和系统发育种概念

一般来讲,进化学派承认分支学派对系统学的研究作出了有意义的贡献,如应用分支分析方法重建系统发育,应用共有衍征确定分类群之间的分支关系以及应用外类群方法来判断性状的极性等,都对系统学的方法有所改进。但分支学派的致命缺点是拒绝接受并系类群。我们属于进化学派,认为并系类群是可以接受的。例如,根据分子资料分析,Zabelia属可以包括于Abelia属内。Zabelia属不但在花粉上和Abelia属不同,可能由于它占有了新的生态位,获得了新的特征,如叶柄基部膨大两两联合,并宿存以保护腋芽。有理由认为它们应独立成属,并不由于Zabelia属从Abelia属分出而使后者成为一个并系类群而把它们合并。分支学派的一些学者认为生物名称作为交流的工具和生物信息储存系统应有明晰的、唯一的和稳定的特性。但具等级的林奈命名系统并不具有这些特性来命名分支和种。最后,PhyloCode被提出。PhyloCode对分支的命名方法有3种,即分支结点定义、分支基干定义和衍征定义。我们认为林奈命名系统作为传媒系统在生物学界的应用已近250年,若要废弃它而采用PhyloCode,必然会在命名方面引起一片混乱。但我们并不是说PhyloCode的拥护者所提出的建议一无是处,我们建议他们宜继续进行研究。由于应用生物学种概念于植物界产生了许多问题,因此多为植物系统学家所抛弃。分支学派的兴起,推动了系统发育种概念的提出。该概念基于3个特征,即自征、区别特征和基本排它,因此分别命名为自征种概念、特征种概念和谱系种概念。事实上,目前大多数植物系统学家仍然应用着形态–地理学种概念,但我们在划分种时,必须有尽可能多的资料,特别是要将传粉、繁育系统、分子系统学资料和形态学资料结合起来。     

Paraphyletic group,PhyloCode and phylogenetic species—the current debate and a preliminary commentary

In this essay, three currently hotly debated issues in biological systematics, i.e., the paraphyletic group, the PhyloCode, and the phylogenetic species concept, have been briefly reviewed. (1) It is widely acknowledged that cladistics has made some positive contributions to the study of systematics. In particular, the employment of outgroup analysis for assessing character polarities, the application of synapomorphies to the inference of relationships between taxa, and the use of cladistic methods for reconstructing phylogeny, have all greatly facilitated the improvement of systematic approaches. A fatal flaw in cladistics is its refusal to accept paraphyletic groups. Frankly, we are adherents and practitioners of phyletics, and hence consider paraphyletic groups to be acceptable. For example, an AFLP analysis has shown that Zabelia (Caprifoliaceae) can be included in Abelia, but the members in Zabelia differ from those in Abelia not only in pollen morphology, but also in having persistent petioles dilated and connate at base, thus enclosing axillary buds, characters of adaptive significance obtained possibly when Zabelia members entered a new ecological niche, so we consider that they are better treated as two independent genera, though indeed such a treatment makes Abelia paraphyletic. (2) Some cladists pointed out that as the tool for communication and the system for information storage and retrieval, biological nomenclature is required to be unambiguous, unique and stable. They criticise the Linnaean rank-based system of nomenclature for failing to satisfy such requirements for the naming of clades and species. To address this problem, the PhyloCode is proposed in recent years, in which three definitions for clade naming are given, i.e., the node-based, the stem-based, and the apomorphy-based. We are of the opinion that since the Linnaean binominal system of botanical nomenclature has existed for nearly 250 years, the rejection of this system and the adoption of the PhyloCode would create a state of chaos in botanical nomenclature. This does not mean that there exist no merits in the proposals made by the PhyloCode supporters. We suggest that further studies should be conducted for its practical application. (3) It has been well known that there are many problems with the application of the biological species concept in plants, and thus at the present time the majority of plant systematists actually seldom use this concept in their practical work. The rapid development of cladistic approach has motivated the proposal of the phylogenetic species concept. This species concept is established based on three criteria, i.e., the autamorphy, the diagnosability and the basal exclusivity, hence the autamorphy species concept, the diagnosability species concept, and the genealogical concept are created respectively. Nevertheless, the morpho-geographical species concept is still predominantly adopted in plant systematics. When using this species concept, however, we should also take into account the data from other sources, particularly those from pollination biology, breeding system and molecular systematics.