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.     

Perspectives: Towards a language for mapping relationships among taxonomic concepts

Abstract

Taxonomic concepts (sensu Berendsohn) embody the underlying meanings of scientific names as stated in a particular publication, thus offering a new way to resolve semantic ambiguities that result from multiple revisions of a taxonomic name. This paper presents a comprehensive and powerful language for representing the relationships among taxonomic concepts. The language features terms and symbols for concept relationships within a single taxonomic hierarchy, or between two related but independently published hierarchies. Taxonomic concepts pertaining to a single hierarchy are characterised by parent/child relationships, whereas those pertaining to two independent hierarchies may have the following basic relationships: congruence, inclusion (non‐symmetrical, relative to the side of comparison), overlap, and exclusion. The relationships are asserted by specialists who have the option to add or subtract concepts on one or both sides of a relationship equation in order to reconcile differences between non‐congruent taxonomic perspectives. The terms ‘and’, ‘or’ and ‘not’ are available, respectively, to connect multiple simultaneously or alternatively valid relationship assessments, or to explicitly negate the validity of a relationship. The language also permits the decomposition of a relationship according to the intensional (property referencing) and ostensive (member pointing) aspects of the compared taxonomic concepts. Adopting the concept relationship language will facilitate a more precise documentation of similarities and differences in multiple succeeding taxonomic perspectives, thereby preparing the stage for an ontology‐based integration of taxonomic and related biological information.     

A synthetic phylogeny of freshwater crayfish: insights for conservation

Phylogenetic systematics is heading for a renaissance where we shift from considering our phylogenetic estimates as a static image in a published paper and taxonomies as a hardcopy checklist to treating both the phylogenetic estimate and dynamic taxonomies as metadata for further analyses. The Open Tree of Life project (opentreeoflife.org) is developing synthesis tools for harnessing the power of phylogenetic inference and robust taxonomy to develop a synthetic tree of life. We capitalize on this approach to estimate a synthesis tree for the freshwater crayfish. The crayfish make an exceptional group to demonstrate the utility of the synthesis approach, as there recently have been a number of phylogenetic studies on the crayfishes along with a robust underlying taxonomic framework. Importantly, the crayfish have also been extensively assessed by an IUCN Red List team and therefore have accurate and up-to-date area and conservation status data available for analysis within a phylogenetic context. Here, we develop a synthesis phylogeny for the world''s freshwater crayfish and examine the phylogenetic distribution of threat. We also estimate a molecular phylogeny based on all available GenBank crayfish sequences and use this tree to estimate divergence times and test for divergence rate variation. Finally, we conduct EDGE and HEDGE analyses and identify a number of species of freshwater crayfish of highest priority in conservation efforts.     

Species identification in the genus Saprolegnia (Oomycetes): Defining DNA-based molecular operational taxonomic units

The lack of a robust taxonomy in the genus Saprolegnia is leading to the presence of incorrectly named isolates in culture collections and of an increasing number of misassigned sequences in DNA databases. Accurate species delimitation is critical for most biological disciplines. A recently proposed approach to solve species delimitation (taxonomic diagnosis system) of difficult organisms is the definition of molecular operational taxonomic units (MOTUs). We have used 961 sequences of nrDNA ITS from culture collections (461 sequences) and GenBank (500 sequences), to perform phylogenetic and clustering optimization analyses. As result, we have identified 29 DNA-based MOTUs in agreement with phylogenetic studies. The resulting molecular clusters support the validity of 18 species of Saprolegnia and identify 11 potential new ones. We have also listed a number of incorrectly named isolates in culture collections, misassigned species names to GenBank sequences, and reference sequences for the species. We conclude that GenBank represents the main source of errors for identifying Saprolegnia species since it possesses sequences with misassigned names and also sequencing errors. The presented taxonomic diagnosis system might help setting the basis for a suitable identification of species in this economically important genus.     

Visual exploration of alternative taxonomies through concepts

A graphical user interface is presented that allows users of taxonomic data to explore concept relationships between conflicting but related taxonomic classifications.Ecological analyses that use taxonomic metadata depend on accurate naming of specimens and taxa, and if the metadata involves several taxonomies, care has to be taken to match concepts between them. To perform this accurately requires expert-defined concept relationships, which are more complex yet more representative than the simple one-to-one mappings found through simple name matching, and can accommodate nomenclatural changes and differences in classification technique (cf ‘lumpers’ versus ‘splitters’). In the SEEK-Taxon (Scientific Environment for Ecological Knowledge) project we aim to help users of taxonomic datasets untangle and understand these relationships through a prototype visual interface which graphically displays these relationship structures, allowing users to comprehend such information and more accurately name their data.     

Cross-cultural variation in the development of folk ecological reasoning

Two studies examined children's reasoning about biological kinds in populations that vary in formal education and direct experience with the natural world, a Western (urban U.S.) and a Non-Western population (Tanna, Vanuatu). Study 1 examined children's concepts of ecological relatedness between species (N = 97, 5–13-year-olds). U.S. children provided more taxonomic explanations than Ni-Vanuatu children, who provided more ecological, physiological, and utility explanations than U.S. children. Ecological explanations were most common overall and more common among older than younger children across cultures. In Study 2, children (N = 106, 6–11-year-olds) sorted pictures of natural kinds into groups. U.S. children were more likely than Ni-Vanuatu children to categorize a human as an animal and the tendency to group a human with other animals increased with age in the U.S. Despite substantial differences in cultural, educational, and ecological input, children in both populations privileged ecological reasoning. In contrast, taxonomic reasoning was more variable between populations, which may reflect differences in experience with formal education.     

Aphyly: A Systematic Designation for a Taxonomic Problem

A taxon is aphyletic when it is deemed to be non-monophyletic or unresolved, therefore aphyletic taxa are a taxonomic problem rather than an evolutionary anomaly. A problem arises in systematics when taxonomic names assigned to aphyletic taxa are treated as if they were natural groups. In the absence of a taxonomic and systematic revision, anomalous taxa should be labelled as aphyletic without recourse to phylogenetic inference (i.e., interpretation). Doing so avoids the validation of aphyletic names and the creation of dubious results in fields that rely on systematic and taxonomic data.     

Definitions in phylogenetic taxonomy: critique and rationale

A general rationale for the formulation and placement of taxonomic definitions in phylogenetic taxonomy is proposed, and commonly used terms such as "crown taxon" or "node-based definition" are more precisely defined. In the formulation of phylogenetic definitions, nested reference taxa stabilize taxonomic content. A definitional configuration termed a node-stem triplet also stabilizes the relationship between the trio of taxa at a branchpoint, in the face of local change in phylogenetic relationships or addition/deletion of taxa. Crown-total taxonomies use survivorship as a criterion for placement of node-stem triplets within a taxonomic hierarchy. Diversity, morphology, and tradition also constitute heuristic criteria for placement of node-stem triplets.     

Outlier detection in BLAST hits

Background

An important task in a metagenomic analysis is the assignment of taxonomic labels to sequences in a sample. Most widely used methods for taxonomy assignment compare a sequence in the sample to a database of known sequences. Many approaches use the best BLAST hit(s) to assign the taxonomic label. However, it is known that the best BLAST hit may not always correspond to the best taxonomic match. An alternative approach involves phylogenetic methods, which take into account alignments and a model of evolution in order to more accurately define the taxonomic origin of sequences. Similarity-search based methods typically run faster than phylogenetic methods and work well when the organisms in the sample are well represented in the database. In contrast, phylogenetic methods have the capability to identify new organisms in a sample but are computationally quite expensive.

Results

We propose a two-step approach for metagenomic taxon identification; i.e., use a rapid method that accurately classifies sequences using a reference database (this is a filtering step) and then use a more complex phylogenetic method for the sequences that were unclassified in the previous step. In this work, we explore whether and when using top BLAST hit(s) yields a correct taxonomic label. We develop a method to detect outliers among BLAST hits in order to separate the phylogenetically most closely related matches from matches to sequences from more distantly related organisms. We used modified BILD (Bayesian Integral Log-Odds) scores, a multiple-alignment scoring function, to define the outliers within a subset of top BLAST hits and assign taxonomic labels. We compared the accuracy of our method to the RDP classifier and show that our method yields fewer misclassifications while properly classifying organisms that are not present in the database. Finally, we evaluated the use of our method as a pre-processing step before more expensive phylogenetic analyses (in our case TIPP) in the context of real 16S rRNA datasets.

Conclusion

Our experiments make a good case for using a two-step approach for accurate taxonomic assignment. We show that our method can be used as a filtering step before using phylogenetic methods and provides a way to interpret BLAST results using more information than provided by E-values and bit-scores alone.

     

A warning for ecologists and conservation biologists using species checklists: How the European marine fauna ‘lost’ all of its 16 <Emphasis Type="Italic">Discodoris</Emphasis> species (Mollusca: Gastropoda)

The European marine fauna used to be considered to include 16 species of Discodoris sea slugs until a recent worldwide revision demonstrated that there is not a single Discodoris species in European waters. This exemplary case illustrates the fact that species checklists do not accurately represent biodiversity unless they are based on sound taxonomic work in which (1) the status of every available species name has been addressed, i.e. whether it is valid, synonymous, or of doubtful application, and (2) classification reflects phylogenetic relationships. It is argued that taxonomic revisions are critically needed, because the status of species names can only be addressed properly through revisions. It is discussed that fields which depend on taxonomic data, such as conservation biology and ecology, might be affected deeply if problematic species names (synonyms and nomina dubia) have not been recognized. Consequently, it is proposed that a taxon that has not been revised be red-flagged in checklists, so that non-taxonomists will know which species names should be applied with caution or not at all.     

Delimitation despite discordance: Evaluating the species limits of a confounding species complex in the face of mitonuclear discordance

The delimitation of species is an essential pursuit of biology, and proper taxonomies are crucial for the assessment and conservation management of organismal diversity. However, delimiting species can be hindered by a number of factors including highly conserved morphologies (e.g., cryptic species), differences in criteria of species concepts, lineages being in the early stages of the speciation or divergence process, and discordance between gene topologies (e.g., mitonuclear discordance). Here we use a taxonomically confounded species complex of toads in Central America that exhibits extensive mitonuclear discordance to test delimitation hypotheses. Our investigation integrates mitochondrial sequences, nuclear SNPs, morphology, and macroecological data to determine which taxonomy best explains the divergence and evolutionary relationships among these toads. We found that a three species taxonomy following the distributions of the nuclear SNP haplotypes offers the best explanation of the species in this complex based off of the integrated data types. Due to the taxonomic instability of this group, we also discuss conservation concerns in the face of improper taxonomic delimitation. Our study provides an empirical and integrative hypothesis testing framework to assess species delimitation hypotheses in the face of cryptic morphology and mitonuclear discordance and highlights the importance that a stable taxonomy has over conservation‐related actions.     

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.     

The 'species problem' and testing macroevolutionary hypotheses

Species lists change for a variety of reasons, including new information and preferences for different species concepts. Uncertainty over species numbers is potentially damaging to tests of proposed correlates of species richness, particularly if taxonomic changes are biased toward some clades over others. We investigate the effects of this error and bias by testing the same suite of macroevolutionary hypotheses in seven different arrangements of primate taxonomy. This is the first time that the effects of the ‘species problem’ have been systematically investigated in this way. Primates are an excellent model system for examining the effects of taxonomic uncertainty: species numbers have doubled in the past two decades, with the fastest growth in the Neotropics. We found that different variables were significantly associated with species richness in each taxonomic arrangement. However, there were no significant differences among taxonomies in the regression slopes for any predictor variable. We found no tendency for significant correlations to occur in taxonomies with more species, suggesting that the results cannot be explained by a lack of power in the smaller taxonomies. The findings are discussed with reference to the wider implications for testing macroevolutionary hypotheses.     

Assessing DNA Barcoding as a Tool for Species Identification and Data Quality Control

In recent years, the number of sequences of diverse species submitted to GenBank has grown explosively and not infrequently the data contain errors. This problem is extensively recognized but not for invalid or incorrectly identified species, sample mixed-up, and contamination. DNA barcoding is a powerful tool for identifying and confirming species and one very important application involves forensics. In this study, we use DNA barcoding to detect erroneous sequences in GenBank by evaluating deep intraspecific and shallow interspecific divergences to discover possible taxonomic problems and other sources of error. We use the mitochondrial DNA gene encoding cytochrome b (Cytb) from turtles to test the utility of barcoding for pinpointing potential errors. This gene is widely used in phylogenetic studies of the speciose group. Intraspecific variation is usually less than 2.0% and in most cases it is less than 1.0%. In comparison, most species differ by more than 10.0% in our dataset. Overlapping intra- and interspecific percentages of variation mainly involve problematic identifications of species and outdated taxonomies. Further, we detect identical problems in Cytb from Insectivora and Chiroptera. Upon applying this strategy to 47,524 mammalian CoxI sequences, we resolve a suite of potentially problematic sequences. Our study reveals that erroneous sequences are not rare in GenBank and that the DNA barcoding can serve to confirm sequencing accuracy and discover problems such as misidentified species, inaccurate taxonomies, contamination, and potential errors in sequencing.     

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.