A scientometric investigation on the impact of the PhyloCode proposal on the scientific literature

SCHELESKY-PRADO, DANIEL C.; FALASCHI, RAFAELA L.; AFFONSO, IGOR P.

doi:10.1590/0001-3765202420231178

Abstract

In 2000, the first draft of PhyloCode was made public, an alternative naming code that does not follow Linnaean rankings and is based on the phylogenetic relationships of taxa in cladograms. In this study, the impact of the PhyloCode on scientific literature was analyzed from its first appearance in the literature to 2021. We investigated the areas that have most assimilated the proposal, the criticisms that have arisen over time, and whether there has been growing adherence to it up to the present day. The analyzed data were obtained from the Clarivate Analytics Web of Science database, where 121 articles that used or discussed the PhyloCode were found. Initially, there was an increase in publications, but in mid-2004, there was a downward trend, which was more noticeable after 2008. Results suggest that despite the criticisms, the proposal has been used in research in several areas, mainly in Zoology and Botany, as an alternative to the Linnaean ranking system. Most articles have been published in Systematics and Taxonomy and discuss the functionality of the proposed code. Despite the proposal’s potential, its acceptance can be considered relatively low and it still generates discussions, just like any scientific novelty.

Key words Classification; nomenclature; scientometrics; taxonomic ranks

INTRODUCTION

Biological systematics, the science of classifying living organisms based on morphological characteristics and molecular data, boasts a rich history dating back centuries (Manktelow 2010). Early taxonomic endeavors can be traced to ancient civilizations, such as the pharmacopoeias of the Chinese Emperor Shen Nung (around 3000 BC) and Egyptian depictions of medicinal plants around 1500 BC (Manktelow 2010). Nevertheless, the framework of biological systematics as we know it today found its foundations through the pioneering work of the Swedish naturalist Carl Linnaeus (1707–1778), who developed his classification system, heavily influenced by the fundamentals of the Greek philosopher Aristotle (384 BC–322 BC) (Amorim 2002). Notably, in his “Species Plantarum” (1753) and in the tenth edition of “Systema Naturae” (1758), Linnaeus introduced the enduring binomial nomenclature for plants and animals, respectively, both perpetuated to this day (Amorim 2002).

It is important to recognize that Linnaeus was not a solitary naturalist in the domain of biological nomenclature during this time. Early and contemporary scholars, such as the English naturalist John Ray (1627–1705) and the German physician and botanist Augustus Quirinus Rivinus (1652–1723), made substantial contributions to this field. Ray classified plants based on morphological attributes, whereas Rivinus consistently advocated for a nomenclatural rule requiring shared generic names within the same plant genus (Nicolau 2017). These contributions collectively laid the foundation for modern biological systematics.

Traditional nomenclature codes, notably the International Code of Zoological Nomenclature (ICZN) and the International Code of Nomenclature for algae, fungi, and plants (ICN), have long relied on the Linnaean ranking system to represent various taxonomic levels, from Kingdom to Class and beyond (International Code of Zoological Nomenclature 1999, Turland et al. 2018). Although this hierarchical ranking system lacks intrinsic biological (and evolutionary) significance and is perceived as somewhat arbitrary by certain scholars, it serves as a practical means of associating groups with taxonomic ranks, being useful for the organization and nomenclature of taxa (Amorim 2002).

Notwithstanding Linnaeus’ significant contributions to systematics, some of his ideas are incongruent with contemporary biological knowledge (Nicolau 2017). Linnaeus’ system lacked an evolutionary perspective. The theory of evolution, which would come to the fore a century later, was still in its formative stages during Linnaeus’ time (Manktelow 2010, Nicolau 2017). Linnaeus’ primary focus was on classification, nomenclature, and description, rather than delving into the intricate web of evolutionary relationships among organisms.

A wave of subsequent contributions to biological systematics, such as Darwin’s and Wallace’s evolutionary theory, and later genetics, molecular studies, simulations, and computational modeling, culminated in the emergence of Phylogenetic Systematics in the 20th century (Manktelow 2010, Nicolau 2017). This school of thought was notably distinguished by its concern for unraveling the evolutionary history of taxa, moving beyond mere utilitarian classification.

In 2000, an alternative proposal for biological classification, known as PhyloCode, was introduced. This innovative nomenclature code dispensed with traditional Linnaean rankings, instead focusing on the relationships between taxa in cladograms (Cantino & de Queiroz 2000, 2019). The adoption and reception of the PhyloCode in the scientific community have sparked diverse reactions, with some fervently supporting and using it (Gauthier & de Queiroz 2001, Langer 2001) whereas others vehemently criticize it (Benton 2000, Platnick 2012).

It is pertinent to emphasize that critics of the PhyloCode have often cited a failure to fully grasp its principles (de Queiroz & Donoghue 2011). A pivotal aspect of this discussion pertains to understanding the core of the PhyloCode, which centers on phylogenies — hypotheses elucidating relationships among taxa forming clades. Importantly, the PhyloCode primarily focuses on naming clades, with species names continuing to be governed by traditional rank-based codes (Cantino & de Queiroz 2019). As such, the PhyloCode can coexist with established codes and does not necessitate the use of categorical ranks, making rank-associated endings, such as “idae” for zoological families and “-aceae” for botanical families, irrelevant to the composition of taxa (de Queiroz & Donoghue 2011).

Furthermore, taxon names within the PhyloCode are based on phylogenetic definitions, falling into three types: apomorphy-based, minimum-clade definition (formerly node-based), and maximum-clade definition (formerly stem-based/branch-based) (Gauthier & de Queiroz 2001, Cantino & de Queiroz 2019). In these definitions, traditional nomenclatural types are supplanted by specifiers, such as specific apomorphies in apomorphy-based definitions or internal/external reference taxa in minimum and maximum-clade definitions. Additionally, the International Clade Names Repository, RegNum (https://phyloregnum.org/), has been proposed as a repository for clade names, each accompanied by a registration number and additional pertinent information (Gauthier & de Queiroz 2001, Cantino & de Queiroz 2019).

Within this context, this study aims to scientometrically investigate the impact of the PhyloCode proposal on scientific literature from its inception in 2000 to December 8th, 2021. Scientometrics, a valuable tool for gaging scientific progress, leverages bibliometrics to glean insights into published scientific articles, research institutions, scientific journals, and knowledge domains. To achieve this goal, quantitative and impact indicators in bibliometrics are employed, encompassing measures of scientific activity and impact (da Silva & Bianchi 2001). This investigation offers insights on reception and incorporation of this approach among scientists, discussing dialogues and assessments concerning its advantages and constraints.

MATERIALS AND METHODS

Data were collected using the database of the Clarivate Analytics Web of Science (WoS) database, searching the terms “PhyloCode*” and “PhyloCod*” (if there were articles that used this prefix, however, no article with a term other than PhyloCode was found). The temporal delimitation was between the years 2000 and 2021, a period of 22 years. The articles were collected until December 8th, 2021.

To analyze the content of the articles, some basic information was collected, such as the title, author(s), date when the article was published, number of citations, and journal in which it was published. The area of biology in which the PhyloCode was inserted was also added. Articles that simply applied the PhyloCode to a taxonomic group or discussed why it is harmful to the group’s classification were considered in the areas of zoology, botany, mycology, and phycology. Articles that essentially discussed PhyloCode theoretically and the practical consequences, positive or negative, without applying to a taxonomic group or just using some group as an example, were considered in the area of Systematics and Taxonomy. There was a categorization regarding the use of the code for each article: concordant (whether using it for the classification and nomenclature of a group or just defending its use and theoretical scope), discordant (that is, criticizing its use and demonstrating its flaws), or even indifferent (as in cases of articles that only mentioned that there was a discussion on the subject at a scientific event or not showing a preference for its use or disuse of it). Also, the country of institution of the corresponding authors and the taxonomic groups of the papers (fossil, extant or both, only for the articles in the field of zoology) were computed.

In a scientometric research, the quantitative indicators of scientific activity use the number of publications. Thus, the number of publications per year, by area of Biology, by journal, and if the articles were concordant, discordant, or indifferent to the PhyloCode were tabulated and interpreted through graphs. Impact indicators of the articles use the number of citations of the publications and the impact indicators of the journals use the number of citations of the respective journals (da Silva & Bianchi 2001). Here, the number of citations per publication were also graphically represented and interpreted.

A regression tree analysis was employed to examine potential temporal patterns in the publication count of papers on PhyloCode. This approach involves dividing the predictor variable into segments based on similar values of the response variable. The segmentation process is repeated until the number of observations within a segment becomes small (De’Ath & Fabricius 2000). The analysis was conducted using the package ‘rpart’ (Therneau & Atkinson 2010) within the R environment (R Development Core Team 2021).

RESULTS

A total of 121 publications addressed this subject. Among them, there are articles that explain the principles, use, and apply the methodology to name groups, cite or criticize the PhyloCode. Regarding the number of publications per year (Figure 1), there was a peak in 2005 and another in 2007, both with 14 articles published. Most articles were published between 2003 and 2007 (58 articles, approximately 47%), and few articles were published since 2012, with a maximum of five articles per year.

Figure 1
Number of papers indexed by the Institute for Scientific Information (ISI) published on PhyloCode between 2000 and 2021. The dashed line indicates the year (2004.5) in which the regression tree partitioned the data in two segments.

Regarding the areas of Biology that used the PhyloCode (Figure 2), Systematics and Taxonomy had the highest number of articles, 72, followed by Zoology with 29, and Botany with 12 articles (together they account for more than 90%). However, there are also articles related to Ecology, Phycology, and Mycology, a total of eight articles. Regarding the number of publications per journal, only 17 of the 58 journals have more than one publication related to the PhyloCode (Figure 3). As highlights, we have the following journals: Systematic Biology, 15, Cladistics, 12, and Taxon, nine. The other journals have seven or less publications.

Figure 2
Number of articles published on PhyloCode by area of Biology.

Figure 3
Number of articles published on PhyloCode per journal.

Regarding the number of citations per article, we have 100 articles with up to 50 citations and 13 articles with 51 to 100 citations (Figure 4). The rest of the citation classes have five or fewer articles each (101 to 150 = five, 151 to 200 = one, 201 to 250 = zero, 251 to 300 = one, 301 to 350 = one). Regarding the articles being concordant, discordant, and indifferent, we have 50 concordant, 48 discordant, and 23 indifferent (see Appendix).

Figure 4
Number of articles published on PhyloCode by the number of citations (in classes).

Regarding the countries of the institutions of the corresponding authors, there are 22 countries (Figure 5), of which the United States of America, France, and the United Kingdom stand out in relation to the number of publications (46, 19, and ten, respectively). France has more concordant than discordant articles, the United Kingdom is the opposite, and the United States of America has a similar number of concordant and discordant articles (Table I).

Figure 5
Number of articles published on PhyloCode per country.

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Table I
Number of concordant, discordant and indifferent articles per country.

In zoology articles, eight phyla of animals were represented in the works: Annelida, Arthropoda, Brachiopoda, Chordata (only Vertebrata), Echinodermata, Mollusca, Porifera, and Platyhelminthes (Figure 6). The phyla with the most articles are Chordata (12), and Porifera (six). Most publications were of extant taxa (21), some of both extant and fossil taxa (seven) and only one exclusively of fossils (see Appendix).

Figure 6
Number of articles published on PhyloCode per zoological phylum.

The area of biology that most debated the PhyloCode was Systematics, something to be expected since a large part (about 44.63%) of the articles dealt with discussions between those who supported and those who argued against the use of the code, in addition to some indifferent articles. The rest of the articles (about 27.3%) are, in short, about the application of PhyloCode to generate biological classifications on an experimental basis. The major areas of Biology that presented the most expressive numbers of publications were Zoology and Botany, but Phycology, Ecology, and Mycology presented a small fraction of publications (6.6% of the total).

Most of the articles (50) are compliant with the PhyloCode, i.e., they defend, explain and/or use the code to generate classifications. However, a very close number (48) disagreed, i.e., they criticized the code and discouraged its use. A smaller portion of the articles are indifferent (23), i.e., they only mention the PhyloCode and do not show a preference for using it or not.

DISCUSSION

The findings indicate a significant surge in annual article publications related to the PhyloCode from its inception in the early 2000s until around mid-2004. However, after 2008, there was a noticeable decline in the number of publications, marking a consistent downward trend in this area of study up to 2021. These trends suggest that the initial years following the introduction of the PhyloCode saw extensive debates, resulting in numerous articles that presented both support and criticism of the code, along with practical applications for classification. Subsequently, the decline in publications may reflect diminishing interest in the subject and a gradual waning of the tool’s application.

A possible explanation for this decline is that, in 2003, seven discordant articles were published, two of which had more than 50 citations and only one concordant article. Furthermore, in April of the following year, the article with the most citations was published, Wheeler (2004), which was a discordant article. This may have discouraged discussions about the PhyloCode.

The fact that most of the articles deal with the pros and cons of the proposal to try to predict problems in its application explains the concentration of publications on the subject in journals dedicated exclusively to the area of systematics, such as Systematic Biology, Cladistics, and Taxon.

Regarding citations, the relatively small number of articles with more than 100 citations suggests that the studies did not significantly impact the scientific literature. Apart from a few articles that reached a relatively high number of citations (an article with 192, another with 290, and another with 302 were the record holders in the number of citations), the low number of citations may reflect the non-acceptance of the PhyloCode by many taxonomists or even the lack of knowledge about it.

Researchers who oppose PhyloCode present the following arguments: traditional codes can be “fixed” instead of requiring the implementation of a new code, and phylogenetic nomenclature brings problems such as confusion and instability, in addition to reducing the amount of information.

Regarding stability, Langer (2001) argues that it can be interpreted in two ways: the definition of the name of the taxon, i.e., a name must designate only a single taxon and a taxon must not be designated by different names. This concept of stability was adopted by the defenders of PhyloCode, like de Queiroz & Gauthier (1994), in which the code is regarded as stable. Another way to interpret stability is through taxon circumscription, i.e., identifying which taxa are included in a group referred to by a name. This definition was adopted by Nixon & Carpenter (2000), who were opponents of PhyloCode. On that occasion, the code is unstable because when different phylogenetic hypotheses are proposed and used, different taxa may be included in a group referred to by a name.

Regarding the amount of information, de Queiroz & Donoghue (2011) argue that the comparison made by Platnick (2009) is inappropriate. Platnick uses “three-taxon statements”, i.e., propositions about phylogenetic relationships. He compares the information contained in the phylogeny of Acrodonta (a group that traditionally contains the families Agamidae and Chamaeleonidae) in two moments: in the first, the phylogeny in which both families form monophyletic groups, and the classification is based on Linnaean ranking. In the second moment, a new phylogenetic hypothesis is presented, in which the two groups of Agamidae from the first hypothesis no longer form a monophyletic group, one of them being phylogenetically more closely related to the Chamaeleonidae than the other group and the classification used is the minimum-clade definition. Platnick states that in the first case, there are 63 three-taxon statements, i.e., it can be stated that two certain terminal taxa are closer to each other than to a third. In the second case, there are 18 three-taxon statements, resulting in a loss of 71% of the information.

However, de Queiroz & Donoghue (2011) demonstrated that such a comparison is inappropriate, as the amount of information is directly related to the phylogenetic hypothesis used and not to the chosen classification system (Table II). Therefore, the hypothesis of the first case has 63 three-taxon statements, regardless of being classified with Linnaean ranking or phylogenetically, and the second hypothesis presents 18 three-taxon statements, regardless of the system that is applied. Thus, the phylogenetic definitions and Linnaean rankings have the same amount of information relative to the three-taxon statements.

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Table II
Comparison made by Platnick (2009).

To elucidate the question concerning changes in phylogenetic hypotheses and the consequent changes in the classification and names of taxa, we can look at the example of the tribe Bibionini (Diptera, Bibionidae, Bibioninae). The taxon has the following genera: Bibio Geoffroy, 1764; Bibiodes Coquillett, 1904; Bibionellus Edwards, 1935; and Enicoscolus Hardy, 1961 (Pinto & Amorim 2000). Phylogenetic studies indicate that Bibio is probably not monophyletic (Fitzgerald 2004, Skartveit & Willassen 1996). So, if one or more genera of Bibionini are taxa that make Bibio paraphyletic, what should be done to solve this taxonomic problem?

When using traditional nomenclature, several options are available. One possibility is to synonymize the genus Bibiodes with Bibio. Furthermore, if Bibiodes is monophyletic, it could be considered a subgenus of Bibio. For example, Bibiodes halteralis Coquillett, 1904 would then receive the name Bibio (Bibiodes) halteralis Coquillett, 1904. This option would result in a change in the ranking of Bibiodes from genus to subgenus, and the name Bibio would cover more taxa, in line with the hypothesis that this genus is monophyletic. A second option would involve dividing the genus Bibio into new genera, with each monophyletic subgroup receiving a new name, except for Bibiodes and other genera that are different from Bibio. The subgroup that includes the type species of the genus Bibio, namely Bibio hortulanus (Linnaeus), would retain the name Bibio. The second option thus involves proposing new generic names for each monophyletic subgroup, with Bibiodes remaining as a separate genus. This option would result in a decrease in the definition and limits of Bibio.

The PhyloCode provides another option: a taxon of a certain taxonomic category can be included in another taxon of the same category; thus, Bibiodes and other genera of Bibionini could be included in Bibio without changing their names, thus maintaining both the names of the genera, as well as the species belonging to them.

In general, when a scientific novelty appears, there is a natural period to absorb the knowledge, acceptance, and academic incorporation. With the PhyloCode, it is not different. Like any new proposal, some supporters see its potential, but there are also many critics. The latter form a heterogeneous group, which ranges from those who claim to find qualities in the proposal (Frost et al. 2009) to those who are completely contrary, suggesting that the code is unnecessary, harmful, and meaningless (Platnick 2012).

Something interesting to note is that the PhyloCode does not bring a revolutionary idea in the sense that its contents are already known and applied, such as phylogenies, the search for monophyletic groups, and the very understanding of the diversity of life because of biological evolution. What is new is how to apply these concepts in the nomenclature differently from other codes, and in this attempt, it brings new ways of naming supra specific taxa, using phylogenetic definitions, abandoning the obligation of Linnaean ranking and its hierarchy, allowing, for example, that taxa with the same ending are not mutually exclusive.

Another important point is that it exclusively concerns the nomenclature. The method to perform systematics remains the same with or without the PhyloCode. The search for phylogenetic hypotheses that best portray the evolutionary history of a given group predates the code. Therefore, traditional codes already deal with the inevitable and multiple changes in phylogenetic hypotheses. Phylogenies can change, the difference between the PhyloCode and the other codes is how to transpose a new hypothesis into a new classification, new names, names that become obsolete, new ranges of taxa, and definitions used. From the same phylogeny, these codes introduce two ways of generating a classification. Therefore, it is up to the researcher to analyze whether it is advantageous for their data to use the PhyloCode exclusively, use it together with traditional codes, or just use the latter.

Both traditional and PhyloCode nomenclatures offer advantages and limitations. For example, PhyloCode offers greater taxonomic stability (de Queiroz & Gauthier 1994), which can be beneficial for researchers seeking to create a more accurate and consistent system. However, traditional codes offer greater nomenclatural stability (Nixon & Carpenter 2000), which may be important for researchers seeking to maintain continuity with established naming conventions. The choice between the two is not always clear-cut because each offers distinct advantages depending on the researcher’s goals and priorities. Ultimately, the decision of which code to use is personal and may vary depending on the specific group being studied and the researcher’s goals.

It is worth noting that PhyloCode can be used in conjunction with traditional codes, allowing researchers to incorporate both nomenclatures into their work. This flexibility allows researchers to tailor their classification systems to their specific needs and preferences. Regardless of which code is used, the phylogeny of a group will be known, and the choice of nomenclature will depend on the researcher’s priorities and goals.

This research revealed a trajectory in publications regarding the PhyloCode subject that emerges in debates and publications shortly after its introduction but declines in interest over time. The fluctuating trends in articles suggest that the idea is plausible and received attention from the scientific community, characterized by initial enthusiasm, but followed by a subsequent skepticism. It is expected that discussions surrounding the PhyloCode may continue, although with varying degrees of fervor and engagement. Ultimately, the future trajectory of articles on this topic will likely rely on upcoming debates, emerging research findings and the evolving of taxonomic methodologies.

ACKNOWLEDGMENTS

We thank Diego Aguilar Fachin, Carla S. Pavanelli and anonymous reviewers for providing valuable suggestions and comments that significantly enhanced the quality of this manuscript. This article was based on the undergraduate thesis of DCSP (2021), whose advisors were IPA and RLF. DCSP received a scholarship from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), Brazil (process 131837/2022-2), IPA thanks CNPq (process 200018/2024-8) and RLF thanks the Fundação de Amparo à Pesquisa do Estado de Goiás -FAPEG (process 202410267000126).

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APPENDIX.

APPENDIX. List of articles studied chronologically descending. The article is identified by the author(s) and year of publication. The data presented are: area of Biology (Area), number of citations (Citations), if the article is concordant, discordant, or indifferent (CDI), country of institution of the corresponding authors (Country), and the phylum, including if the taxa studied are fossil, extant or both (Phylum).

Article	Area	Citations	CDI	Country	Phylum*
Sakamoto & Ortega 2021	Systematics and Taxonomy	0	Indifferent	Brazil	-
Maxwell et al. 2021	Zoology	0	Concordant	Australia	Mollusca (both)
Penagos Zuluaga et al. 2021	Botany	0	Concordant	United States of America	-
Sangster & Mayr 2021	Zoology	0	Concordant	Netherlands	Chordata (extant)
Joyce et al. 2021	Zoology	10	Concordant	Switzerland	Chordata (both)
Brower 2020	Systematics and Taxonomy	0	Discordant	United States of America	-
Guayasamin & Trueb 2020	Systematics and Taxonomy	0	Indifferent	Ecuador	-
Chen & Field 2020	Zoology	0	Concordant	United Kingdom	Chordata (extant)
Laurin 2019	Systematics and Taxonomy	0	Concordant	France	-
Fucikova et al. 2019	Phycology	8	Concordant	United States of America	-
Hibbett et al. 2018	Mycology	10	Concordant	United States of America	-
Kraichak et al. 2018	Mycology	33	Discordant	Thailand	-
Cohen 2018	Zoology	3	Discordant	United Kingdom	Brachiopoda (both)
Crowl & Cellinese 2017	Botany	0	Concordant	United States of America	-
Leal et al. 2017	Zoology	6	Concordant	Brazil	Porifera (extant)
Wu et al. 2016	Zoology	3	Concordant	China	Arthropoda (extant)
Leslie 2015	Ecology	2	Indifferent	United States of America	-
Miller et al. 2014	Botany	8	Concordant	Australia	-
Bennett & Balick 2014	Botany	44	Discordant	United States of America	-
Brummitt 2014	Botany	19	Discordant	United Kingdom	-
Wojciechowski 2013	Botany	15	Concordant	United States of America	-
Olesen & Richter 2013	Zoology	24	Concordant	Denmark	Arthropoda (extant)
De Queiroz & Donogue 2013	Systematics and Taxonomy	5	Concordant	United States of America	-
Cellinese et al. 2012	Systematics and Taxonomy	10	Concordant	United States of America	-
Gazave et al. 2012	Zoology	71	Concordant	France	Porifera (extant)
Platnick 2012	Systematics and Taxonomy	9	Discordant	United States of America	-
Cardenas et al. 2012	Zoology	136	Concordant	France	Porifera (extant)
De Queiroz & Donogue 2011	Systematics and Taxonomy	6	Concordant	United States of America	-
Wagner et al. 2011	Zoology	17	Discordant	United States of America	Chordata (extant)
Near et al. 2011	Zoology	119	Concordant	United States of America	Chordata (extant)
Aldous et al. 2011	Systematics and Taxonomy	15	Indifferent	United States of America	-
Ward 2011	Zoology	31	Discordant	United States of America	Arthropoda (extant)
Cardenas et al. 2011	Zoology	14	Concordant	France	Porifera (extant)
Assis 2011	Systematics and Taxonomy	4	Discordant	Brazil	-
Wagner 2010	Zoology	1	Discordant	Germany	Chordata (extant)
Gazave et al. 2010	Zoology	34	Concordant	France	Porifera (extant)
Bethoux 2010b	Systematics and Taxonomy	0	Concordant	Australia	-
Mazaris et al. 2010	Ecology	28	Indifferent	Greece	-
Bethoux 2010a	Systematics and Taxonomy	21	Discordant	Germany	-
Dubois 2010	Systematics and Taxonomy	10	Discordant	France	-
Laurin 2010	Systematics and Taxonomy	27	Concordant	France	-
Frost et al. 2009	Zoology	10	Discordant	United States of America	Chordata (extant)
Guayasamin et al. 2009	Zoology	115	Concordant	Ecuador	Chordata (extant)
Laurin & Bryant 2009	Systematics and Taxonomy	5	Concordant	France	-
Mishler 2009	Systematics and Taxonomy	27	Indifferent	United States of America	-
McGuire et al. 2009	Zoology	60	Concordant	United States of America	Chordata (both)
Nakada et al. 2008	Phycology	121	Concordant	Japan	-
Fitzhugh 2008	Systematics and Taxonomy	22	Indifferent	United States of America	-
Laurin 2008	Systematics and Taxonomy	24	Concordant	France	-
Martin & Benton 2008	Zoology	22	Discordant	France	Chordata (both)
Dayrat et al. 2008	Systematics and Taxonomy	30	Concordant	United States of America	-
Wolsan 2007b	Systematics and Taxonomy	5	Indifferent	Poland	-
Lee & Skinner 2007	Systematics and Taxonomy	7	Concordant	Australia	-
Benton 2007	Systematics and Taxonomy	13	Discordant	United Kingdom	-
Dubois 2007c	Systematics and Taxonomy	22	Discordant	France	-
Cantino et al. 2007	Botany	192	Concordant	United States of America	-
Shenoy et al. 2007	Mycology	124	Indifferent	China	-
Dubois 2007b	Systematics and Taxonomy	33	Discordant	France	-
Wolsan 2007a	Systematics and Taxonomy	3	Indifferent	Poland	-
Ereshefsky 2007	Systematics and Taxonomy	29	Indifferent	Canada	-
Fisher et al. 2007	Botany	12	Concordant	United States of America	-
Saucede et al. 2007	Zoology	33	Concordant	France	Echinodermata (fossil)
Dubois 2007a	Systematics and Taxonomy	38	Indifferent	France	-
Hillis 2007	Systematics and Taxonomy	48	Concordant	United States of America	-
Seberg & Petersen 2007	Systematics and Taxonomy	4	Indifferent	Sweden	-
Muona 2006	Zoology	5	Indifferent	Finland	Annelida (extant)
Dubois 2006b	Systematics and Taxonomy	16	Discordant	France	-
Dubois 2006a	Systematics and Taxonomy	22	Indifferent	France	-
Monsch 2006	Systematics and Taxonomy	5	Discordant	Poland	-
Stevens 2006	Systematics and Taxonomy	20	Discordant	United States of America	-
Rieppel 2006	Systematics and Taxonomy	38	Discordant	United States of America	-
De Queiroz 2006	Systematics and Taxonomy	39	Concordant	United States of America	-
Kuntner & Agnarsson 2006	Systematics and Taxonomy	34	Indifferent	Slovenia	-
Bertrand & Harlin 2006	Systematics and Taxonomy	9	Indifferent	Sweden	-
Fisher 2006	Botany	15	Concordant	United States of America	-
Willems et al. 2006	Zoology	51	Concordant	Belgium	Platyhelminthes (extant)
Laurin et al. 2005	Systematics and Taxonomy	20	Concordant	France	-
Pickett 2005b	Systematics and Taxonomy	9	Discordant	United States of America	-
Tang & Lu 2005	Systematics and Taxonomy	3	Discordant	China	-
Lobl & Leschen 2005	Systematics and Taxonomy	16	Indifferent	Switzerland	-
Sereno 2005	Systematics and Taxonomy	69	Indifferent	United States of America	-
Martens & Segers 2005	Systematics and Taxonomy	8	Discordant	Belgium	-
Dayrat 2005	Zoology	13	Concordant	United States of America	Mollusca (extant)
Antonov 2005	Systematics and Taxonomy	1	Indifferent	Russia	-
Korf 2005	Systematics and Taxonomy	18	Discordant	Canada	-
Agapow & Sluys 2005	Ecology	13	Indifferent	United Kingdom	-
Pfeil & Crisp 2005	Botany	35	Discordant	Australia	-
Dayrat & Gosliner 2005	Zoology	20	Concordant	United States of America	Mollusca (extant)
Pickett 2005a	Systematics and Taxonomy	21	Discordant	United States of America	-
Dubois 2005	Systematics and Taxonomy	70	Discordant	France	-
Bertrand & Hogskola 2004	Systematics and Taxonomy	0	Indifferent	Sweden	-
Pickett 2004	Systematics and Taxonomy	0	Discordant	United States of America	-
Stevenson 2004	Systematics and Taxonomy	2	Discordant	United States of America	-
Nygren 2004	Zoology	61	Concordant	Sweden	Annelida (extant)
Groves 2004	Systematics and Taxonomy	55	Discordant	Australia	-
Joyce et al. 2004	Zoology	290	Concordant	United States of America	Chordata (both)
Pavlinov 2004	Systematics and Taxonomy	6	Discordant	Russia	-
Sluys et al. 2004	Systematics and Taxonomy	14	Discordant	Netherlands	-
Donoghue & Gauthier 2004	Systematics and Taxonomy	19	Concordant	United States of America	-
Kraus 2004	Systematics and Taxonomy	8	Discordant	Germany	-
Wheeler 2004	Systematics and Taxonomy	302	Discordant	United States of America	-
Pleijel & Rouse 2003	Systematics and Taxonomy	44	Concordant	France	-
Nixon 2003b	Systematics and Taxonomy	0	Discordant	United States of America	-
Gao & Sun 2003	Systematics and Taxonomy	3	Discordant	China	-
Janovec et al. 2003	Botany	7	Discordant	United States of America	-
Kojima 2003	Systematics and Taxonomy	17	Discordant	Japan	-
Carpenter 2003	Systematics and Taxonomy	34	Discordant	United States of America	-
Keller et al. 2003	Systematics and Taxonomy	79	Discordant	United States of America	-
Nixon 2003a	Systematics and Taxonomy	53	Discordant	United States of America	-
Sennblad & Bremer 2002	Botany	56	Concordant	Sweden	-
Bryant & Cantino 2002	Systematics and Taxonomy	48	Concordant	Canada	-
Berry 2002	Systematics and Taxonomy	8	Discordant	United States of America	-
Brummitt 2002	Systematics and Taxonomy	84	Discordant	United Kingdom	-
Forey 2002	Systematics and Taxonomy	31	Discordant	United Kingdom	-
Langer 2001	Systematics and Taxonomy	8	Concordant	United Kingdom	-
Kress & DePriest 2001	Systematics and Taxonomy	1	Discordant	United States of America	-
Gauthier & De Queiroz 2001	Zoology	89	Concordant	United States of America	Chordata (both)
Benton 2000	Systematics and Taxonomy	97	Discordant	United Kingdom	-
McNeill 2000	Systematics and Taxonomy	14	Indifferent	United Kingdom	-

*Only for Zoology articles.

Publication Dates

Publication in this collection
25 Nov 2024
Date of issue
2024

History

Received
25 Oct 2023
Accepted
25 Aug 2024

This is an open-access article distributed under the terms of the Creative Commons Attribution License

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	First Moment	Second Moment
Phylogenetic hypothesis	both families are monophyletic, Agamidae and Chamaeleonidae	Agamidae is not monophyletic
Used classification	Linnean ranking	node-based (PhyloCode)
Three-taxon statements	63	18