Braby Et Al 2024

Zoological Journal of the Linnean Society, 2024, XX, 1–16
https://doi.org/10.1093/zoolinnean/zlae043
Advance access publication 3 May 2024
Review
Review
How to describe a new species in zoology and avoid mistakes
Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Michael F. Braby1,*, Yu-Feng Hsu2, , Gerardo Lamas3
Division of Ecology and Evolution, Research School of Biology, Australian National University, Acton, ACT 2601 and Australian National Insect
1
Collection, National Research Collections Australia, GPO Box 1700, Canberra, ACT 2601, Australia
2
Department of Life Science, National Taiwan Normal University, Taipei, Taiwan 116, Republic of China
3
Departamento de Entomología, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Av. Arenales 1256, Apartado 14-0434, Lima-
14, Peru
*
Corresponding author. Division of Ecology and Evolution, Research School of Biology, RN Robertson Building, 46 Sullivans Creek Road, The Australian National
University, Acton, ACT 2601, Australia. E-mail: michael.braby@anu.edu.au
ABSTR ACT
Taxonomy is the science of discovering, naming, describing, diagnosing, identifying, and classifying different kinds of taxa, from species to fam-
ilies. It lays the foundation for all of the biological sciences. The rapid increase in both taxonomic descriptions and malpractice in recent decades in-
dicates a need for consistency in the procedure and quality of taxonomic research publications dealing with recognition of new taxa, name changes,
and nomenclatural acts. Indeed, there are numerous examples in the recent literature of taxonomic works that fall short of the basic procedures
and minimum standards required for naming new species according to the mandatory provisions and recommendations of the International Code
of Zoological Nomenclature, accepted standards of taxonomic best practice, and journal editorial policy. Here, we provide practical guidelines of
the procedure and key elements required to name, describe, and publish a new animal species or revise the taxonomic status of a species. We then
discuss some of the common pitfalls that should be avoided. Mistakes commonly made include failure to read the Code, to review the primary
taxonomic literature, to examine type material, to construct the taxon name correctly, to explicitly establish the new taxon, to provide adequate
typification, to clearly differentiate the new taxon, to register the publication of the new taxon name in ZooBank, or to publish the name of the new
taxon in a manner that is compliant with the Code. We provide some examples of these mistakes, mainly from butterflies (Insecta: Lepidoptera:
Papilionoidea) and, to some extent, reptiles because these taxon groups seem to have an unusually high level of poor taxonomic practice.
Keywords: code; description; new species; nomenclature; taxonomy; zoology
INTRODUCTION for scientific communication, establishing clear, universal, and

unambiguous terms of reference. It underpins all fields in the
Without taxonomy to give shape to the bricks, and systematics to biological sciences, laying the foundation for ecology, evolu-
tell us how to put them together, the house of biological science is tionary biology, conservation biology, biosecurity, primary in-
a meaningless jumble. dustry (agriculture, aquaculture, and fisheries), medicine, and
Lord Robert M. May (1990) public health, all of which depend on a robust taxonomy (e.g.
Kitching 1993, Winston 1999, 2018, Godfray 2002, Mallet and
Taxonomy is the science of discovering, naming, describing, Willmott 2003, Wheeler et al. 2004, Wilson 2004, Costello et al.
diagnosing, documenting, recognizing, identifying, and clas- 2013, Vane-Wright 2013, Braby and Williams 2016, Taxonomy
sifying different kinds of organisms (Mayr 1969, Mayr and Decadal Plan Working Group 2018, Hamilton et al. 2021, Löbl
Ashlock 1991). It is often viewed as a subset of systematics, et al. 2023).
which is the science that seeks to explore relationships between Descriptive taxonomy (alpha taxonomy) includes three main
species in an evolutionary context by providing the frame- tasks: (i) generating species-level taxonomic hypotheses and rec-
work into which species are classified—a categorical arrange- ognizing or delimiting the boundaries of these taxa through ac-
ment that gives order to the nature and patterns of variation of quisition of evidence and rigorous hypothesis-testing according
life on Earth. Taxonomy provides the international language to a given species’ concept; (ii) proposing new scientific names,
Received 24 October 2023; revised 4 February 2024; accepted 19 March 2024

© 2024 The Linnean Society of London.
This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial License (https://creativecommons.org/licenses/by-nc/4.0/),
which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited. For commercial re-use, please contact reprints@
oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our
site—for further information please contact journals.permissions@oup.com.
2 • Braby et al.
making nomenclatural acts, and proposing taxonomic opinions so that it is reinstated. If it turns out that one of the species does
(name changes) based on a set of rules and recommendations not have a binomen, then it needs to be formally named and de-
that ought to be followed; and (iii) publication based on a set of scribed as new to become available and valid. On the other hand,
procedures or protocols for best practice. Nomenclatural acts in- the taxonomic research may provide evidence to indicate that
clude type fixations or designations (typification), whereas taxo- there is a single species (i.e. accept H0) that is misclassified and
nomic opinions cause species’ names to change. The names of needs to be reclassified and transferred from the group in which
species change for various reasons (Mallet and Willmott 2003, it was originally described to a group that better reflects its true
Knapp et al. 2004, Thiele et al. 2021), but the main changes in- phylogenetic relationships.

clude synonymy, status revision, and new combinations, as il- The procedures dealing with taxonomic research that lead to
lustrated by the following scenarios. Suppose a taxonomist is nomenclatural changes, such as the recognition of new species
investigating the boundaries of a species group or complex using and name changes based on taxonomic opinions, is, in our ex-
an evidence-based hypothesis testing framework. Suppose fur- perience, where most pitfalls and problems arise in taxonomic
ther that the null hypothesis (H0) is that the complex comprises publications. For any taxonomic publication, there are basic-
only a single species, whereas the alternative hypothesis (H1) ally four different sets of rules and principles that need to be
holds that there are two species (Braby et al. 2012). Although in considered: (i) articles of the relevant code of nomenclature;
this simple case there are only two possible taxonomic conclu- (ii) recommendations of the code of nomenclature; (iii) ac-
sions, there are seven possible decisions or outcomes using such cepted standards of taxonomic best practice; and (iv) journal
a framework (Table 1). For example, it may turn out that what editorial policy and author guidelines. Schenk and McMasters
was previously considered to be two species is in fact one spe- (1948) provided international guidelines for taxonomic best
cies, in which case the younger species’ name needs to be to be practice and publication, including an outline for the descrip-
placed into the synonymy of the older species’ name as a junior tion of a new species (on p. 25), in relation to the first edition
synonym. Alternatively, what was previously considered to be a of the International Code of Zoological Nomenclature. However,
single species may in fact comprise two species (i.e. reject H0 and that pioneering work is now considerably out of date. Mayr’s
accept H1). If both species have species’ names, then it is simply a (1969) classic work on systematic zoology included a detailed
matter of revising the status of the nomen currently in synonymy chapter on how to prepare a taxonomic publication, as well as a
Table 1. Hypothesis testing and possible taxonomic decisions based on revision of a species complex involving one or two species.
Hypothesis Number of species Number of species Taxonomic decision
(before revision) (after revision)
H0 1 1 none
(1 species)
1 1 new combination (comb. nov.)
The investigated species is found to be a member of a genus that is
different from its original or current combination and is transferred
to that genus.
2 1 new synonymy (syn. nov.)
The investigated species is found to be identical to another named
species, and the binomen of that 2nd species is older. The younger
species’ name is placed into the synonymy of the older species’
name as a junior synonym.
H1 1 2 none
(2 species) The investigation reveals the presence of two species, both with avail-
able and valid names, but the 2nd species was previously unknown
from study region.
1 2 revised status (stat. rev.).
The investigation reveals the presence of two species, both with
available names, but the binomen of the 2nd species is currently in
synonymy with the other species. The 2nd species is removed from
synonymy and redescribed as a valid species.
1 2 reinstated status (reinst. stat.)
The investigation reveals the presence of two species, but the
binomen of one species was previously valid but is currently in
synonymy. The previously synonymized name is removed from
synonymy and made valid again.
1 2 new species (sp. nov.)
The investigation reveals the presence of two species, one of which
does not have a binomen. The unnamed species is named and
described as new.
Taxonomic procedures • 3
transcript of the second (1964) edition of the International Code 2005, Kaiser 2013, Winston 2018), and it deals with the regula-
of Zoological Nomenclature. More recently, Winston (1999) pub- tion of names at three different hierarchical levels: family-group
lished a comprehensive manual of best practice for describing names, genus-group names, and species-group names. The Code
new species and an overview of the international zoological and regulates these names according to a set of carefully constructed
botanical codes of nomenclature. Some of the concepts of best articles, with the overarching goal of providing stability and uni-
practice were expanded upon by Dubois (2005), Kaiser (2013), versality to nomenclature based on certain underlying principles
Kaiser et al. (2013), Wüster and Kaiser (2023), and Denzer and (Knapp et al. 2004, Polaszek and Wilson 2005, Winston 2018,
Kaiser (2023). Šlapeta (2013) provided brief guidelines to assist Ceríaco et al. 2023). The articles consist of rules, which are man-

with describing and publishing a new species. However, there is datory provisions, and recommendations, which are advisory
no modern account that sets out succinctly how species ought to statements for best practice. The rules include information on
be named, described, or revised and published (but see: Winston what taxonomic ranks are considered to be valid, how scientific
1999: 84–86). With the rapid increase in both taxonomic pub- names must be constructed and spelt, how capitalization is to
lications and species’ descriptions or delineation in recent dec- be used, how authorities of names (i.e. the author’s name of a
ades (Isaac et al. 2004, Joppa et al. 2011, Bacher 2012, Costello taxon) are to be constructed to avoid ambiguity, and what abbre-
et al. 2013, 2015a, b) there is a pressing need for consistency in viations must be used for new nomenclatural acts. For instance,
the procedure, nomenclature, and quality of taxonomic papers, if the generic assignment of a species has been changed so that
which has been clearly lacking in several works (Wüster et al. the new combination differs from the original combination, then
2021). Hence, the aim and main focus of this paper is to provide the authority citation (and date, if cited) of the species is placed
practical guidelines of the procedure and components required in parentheses. Other rules include details on typification (des-
to describe and publish a new species or to revise the taxonomic ignation of type specimens and fixation of names), synonymy,
status of a species, especially scientists new to the field of tax- principle of priority (use of different names for the same taxon is
onomy (students or individuals without access to mentors) and prohibited), principle of homonymy (use of the same name for
editors of journals that publish taxonomic works. Before dis- different taxa is prohibited), emendation (intentional change in
cussing the important elements of a taxonomic paper, we briefly the original spelling of an available name), principle of the first
comment on nomenclature because it is an important adjunct to reviser, reversal of precedence, and gender agreement among
taxonomy. In this work, we limit our discourse to zoology. other matters when dealing with existing names. Importantly,
the Code makes a clear distinction between available names and
valid names. A species may have many available names but only
ZOOLOGICAL NOMENCL ATURE one valid name, which is the correct name for the taxon in an
Each organism has a unique scientific name that is composed author’s taxonomic judgement.
of two parts, generic and specific, which is referred to as the It is crucial to emphasize that the Code only regulates the use
binominal system (ICZN 1999, Knapp et al. 2004), often in- of names and does not interfere with the quality of science or
correctly referred to as the ‘binomial’ system (Schenk and process of taxonomy, such as the data (evidence) used to delimit
McMasters 1948). The generic or genus name forms the group species and identify independent evolutionary lineages, regard-
name and is always a noun. The specific name is the descriptive less of the quality of the work (Wüster et al. 2001, Knapp et al.
part and may be a noun or an adjective. For animals, the zoo- 2004, Borrell 2007, Krell 2021, Wüster et al. 2021). In other
logical nomenclatural system started with the 10th edition of words, the Code is scientifically neutral and does not interfere
Linnaeus’ Systema Naturae published in 1758 (Winston 1999, with scientific opinion or judgment. The Code also has a Code
2018, Knapp et al. 2004). of Ethics (listed under Appendix A), which provides a set of re-
The nomenclature of most organismal Kingdoms is sup- commendations on various matters, such as avoidance of offen-
ported by a set of rules and recommendations, referred to as sive names, and how to proceed if it is discovered that another
codes (Mayr 1969, Mayr and Ashlock 1991, Knapp et al. 2004). person is working on the same undescribed taxon, but these
There are three separate international codes of nomenclature, all cannot be enforced.
of which operate independently. For animals, the relevant code
is the International Code of Zoological Nomenclature published
by the International Commission on Zoological Nomenclature THE KEY ELE MENTS OF A TA XONOMIC PAPER
(ICZN), which was established in 1895, and ratified by the For a new specific group name (i.e. species or subspecies) to be
International Union of Biological Sciences. Because the codes available and valid under the current edition of the Code, there
are independent, the same taxon name can be used as valid in are four basic elements or criteria that need to be met: (i) the
each code. For example, the generic name Pieris has been used for name of the new species (in Latin or latinized) and a clear indi-
both animals and plants—in zoology the genus Pieris Schrank, cation that it is intended to be new by the author; (ii) the des-
1801 is an insect (butterfly) in the family Pieridae, whereas in ignation or fixation of a holotype; (iii) a description and/or
botany the genus Pieris D.Don is a vascular flowering plant in the diagnosis that purports to differentiate the taxon; and (iv) sci-
family Ericaceae. entific publication in a manner that ensures multiple identical
The International Code of Zoological Nomenclature (hereafter copies of the work containing the name are freely available or by
referred to as the Code) is currently in its fourth edition (ICZN purchase. However, to satisfy both the rules and recommenda-
1999). The next (fifth) edition of the Code is expected to be tions of the ICZN and the accepted standards of scientific best
published later this decade (Rheindt et al. 2023). The Code is a practice and journal editorial policy, the following list of items, in
system or convention, not a science (Knapp et al. 2004, Dubois our view, constitutes the minimum set of information required
Table 2. Example of how to set out the classification headers (higher-level taxon names), type species, species’ name header, figures, and synonymy of a species and subspecies (modified from
4 •
Beaver et al. 2023).

LYCAENIDAE Leach, 1815
OGYRINI Waterhouse & Lyell, 1914
Braby et al.
Ogyris Angas, 1847

Type species: Ogyris amaryllis Hewitson, 1862 by subsequent monotypy.
Ogyris arcana M.R.Williams & Hay, 2001 stat. rev.
(Figs. 1, 3e–h, 6e–h)
Ogyris otanes C.Felder & R.Felder, 1865.—Waterhouse & Lyell, 1914: 121; Waterhouse, 1932: 184; Common & Waterhouse, 1972: 342; Williams et al., 1992: 55–60; Hay et al. [1994]: 31, pl. 4,
fig. 7–9. [Genus Ogyris Westwood, 1851]. [Misidentifications in literature].
Ogyris sp.—Field, 1987: 113.
Ogyris otames C.Felder & R.Felder, 1865.—Hay, 1989: 42, 43. [Misspelling of O. otanes].
Ogyris otanes C.Felder & R.Felder, 1865 ‘local forms 2 and 3’.—Dunn and Dunn, 1991: 367. [Genus Ogyris Westwood, 1851].
Ogyris otanes (C.Felder & R.Felder, 1865) ‘south-western form’.—Braby, 2000: 710–712, pl. 50, fig. 3b. [Genus Ogyris Angas, 1847].
Ogyris otanes (C.Felder & R.Felder, 1865).—Williams et al., 2012: 50–51. [Misidentification in literature]. [Genus Ogyris Angas, 1847].
Ogyris otanes arcana M.R.Williams & Hay, 2001: 59–61, fig. 5–8; Sands & New, 2002: 288; Braby, 2010: 35; Schmidt et al., 2014: 473–484; Braby, 2016: 266–267, 350; Sankowsky, 2020: 315.
Ogyris halmaturia waterhouseri (Bethune-Baker, 1905) stat. rev.
(Figs. 1, 4g–l, 7e–h.)
Ogyris idmo Hewitson, 1862.—Miskin, 1890: 24; Anderson & Spry, 1893: 101, 104; Lower, 1893: 9; Waterhouse, 1903a: 29. [Misidentifications in literature or incorrectly considered to be a
senior synonym of O. halmaturia].
Ogyris waterhouseri Bethune-Baker, 1905: 273–274; Waterhouse and Lyell, 1908: 162, 165–166; Kershaw, 1908: 163–164; Seitz, 1926: 940; Peters, 1971: 26. [Genus Ogyris Westwood, 1851].
Ogyris waterhouseri (Bethune-Baker, 1905).—Braby and Douglas, 2008: 315–329, fig. 1–3, 14–17. [Genus Ogyris Angas, 1847].
Ogyris idmo waterhouseri Bethune-Baker, 1905. — Waterhouse & Lyell, 1914: 122, pl. 18; Burns, 1931: 131–132; Common, 1964: 96; Burns & Rotherham, 1969: 98; D’Abrera, 1971: 320;
McCubbin, 1971: 84; Quick, 1972: 9–10.
Ogyris idmo waterhousei Bethune-Baker, 1905.—Common, 1964: 96. [Misspelling of O. waterhouseri].
Ogyris halmaturia waterhouseri Bethune-Baker, 1905.—Tindale, 1923: 348.
Ogyris idmo halmaturia Tepper, 1890.—Waterhouse, 1932: 179–180;Common and Waterhouse, 1972: 341–342; Crosby, 1974a: 65; Fisher, 1978: 193–194, pl. 10, fig. 11–13; Common and
Waterhouse, 1981: 479; Dunn and Dunn, 1991: 366, 457; Douglas, 1995: 6–10, fig. 1; Grund, 1999: 47–48; Grund, 2003: 71. [Genus Ogyris Westwood, 1851]. [Considered to be a senior
synonym of O. waterhouseri].
Ogyris idmo halmaturia (Tepper, 1890).—Edwards, 1996: 250; Hunt et al., 1998: 113–116; Moore, 1999: 12; Field, 1999: 252–254, 259, fig. 13–16; Braby, 2000: 396, 712–714, pl. 50, fig. 5b;
Edwards et al., 2001: 254–255; Sands and New, 2002: 282–284; Braby, 2004: 242–243, 316; New et al., 2007: 245. [Genus Ogyris Angas, 1847]. [Considered to be a senior synonym of O.
waterhouseri].
Ogyris halmaturia (Tepper, 1890).—Grund, 2010: 114–120; Braby et al., 2011: 29–36; New, 2011: 21, 99, 103–104, 148; Schmidt et al., 2014: 473–484; Braby, 2016: 268–269, 350. [Genus
Ogyris Angas, 1847].

for a taxonomic publication, whether it be of a new species, re- The term comb. nov. is used for a new combination (e.g. spe-
view of a species group or revision of a genus or higher rank (see cies’ name transferred to a genus that is different from its original
also: Winston 1999). There is generally a set order of how the combination); stat. rev. is used for revised status (e.g. subspecies’
items are arranged as headings and subheadings, although this is name treated as full species’ name); and syn. nov. is used for a new
by no means universal, and each publication has its own style of synonymy (e.g. species’ name considered to be a junior synonym
how taxonomic papers are structured. The following order pro- and, therefore, invalid). Another frequent abbreviation is reinst.
vides a logical and coherent framework for the key elements of a stat., meaning, for instance, a previously synonymized name that
taxonomic work. is made valid again, or a name previously downgraded to subspe-

cies status that is restored to valid species status. Combinations of
Classification header these abbreviations ought to be used when two or more changes
The higher-level taxon name, together with the authority citation are made, for example, syn. nov. et stat. rev. means that a spe-
and date, are listed as a heading (Table 2). Often two headings cies is being placed in synonymy but treated as a valid taxon
are given: the family-group name is listed first, followed by the (e.g. subspecies); stat. rev. et comb. nov. means that the status
genus-group name as a second heading. However, some journals of a species is being revised and transferred to a different genus.
require other higher categories of classification as headings, for It should be noted that these terms are guidelines only according
example, names of the superfamily, subfamily, tribe or subtribe to journal editorial policy (e.g. the author guidelines) and are not
to which the species belongs. considered by the Code (see Recommendation 16A: means of ex-
plicitly indicating names as intentionally new).
Type species: Every genus (and subgenus) must have a type spe- The following information should be included under the
cies, and it should be listed below the genus-group name with heading of the taxon name and authority citation.
a statement of how it was designated (Table 2). Type species
of genera are fixed in several ways, but the most common three Figures: New or revised species ought to be illustrated and re-
are: (i) by original designation, which means by the describer ferred to in the figures, with figure numbers listed in the first line
(author) in the original publication; (ii) by monotypy, which after the species’ name heading (Table 2). Although the illustra-
means by the author when only a single species was placed in a tions do not need to include the name-bearing type specimen,
genus in the original publication; or (iii) by subsequent designa- best practice is to include types and many journals now insist on
tion, which means by a reviser when two or more species were this. Illustrations of types (and their labels) of previously named
assigned to a genus for which no type species was fixed in the species and junior synonyms are not essential, but they can be
original publication. For new genera, it must be clearly indicated particularly useful if there is doubt over their existence, authen-
that a type species is being designated for the first time, for ex- ticity, or identity.
ample, by the term ‘type species’ and the name should be given
in its original binomen. ZooBank registration number: For papers published in electronic
only (online) journals, it is now mandatory for the work to be
Species’ name header registered in ZooBank prior to publication (ICZN 2012). The
If the species has not previously been named, then two elements Code does not require registration of the new taxa, although
are required in the heading for the species’ name: (i) the taxon most journals now require that newly described species (as well
name in binominal form; followed by (ii) the term ‘sp. nov.’, ‘sp. as new genera, subgenera, and subspecies) also be registered in
n.’, ‘new species’, or ‘species nova’. The abbreviations sp. nov. or ZooBank. Once registered, the ZooBank registration number
sp. n. actually mean species nova or new species, and one of must be listed in the publication itself.
these terms must follow the taxon name to indicate explicitly
that the author is proposing a name for a species new to science. Type locality: The place of origin from which the name-bearing
If one author or a subset of authors of a multi-authored paper type specimen (holotype, lectotype, or neotype) was collected
are describing the new species, then those author(s) must appear defines the type locality for that species, and it must be given.
after the new name, followed by a comma, and then the term ‘sp.
nov.’ or ‘sp. n.’, so that is clear who is the author(s). Once a new Synonymy. The synonymy section is a critically important part of
species has been described and diagnosed, subsequent reference the paper, and it must be understood clearly. The synonymy con-
to the species in the manuscript need not include ‘sp. nov.’, except sists of a summary of the full history of the nomenclature of the
in the captions of the Tables and Figures. species—all the species’ names and combinations used, plus as
If, however, the species has previously been named and is complete a list of all previous references pertaining to the species
being revised, then four elements are required in the heading for as possible (Table 2). It also provides the author’s conclusions re-
the species’ name: (i) the taxon name in binominal form; (ii) the garding taxonomic assignment of the species and to the validity
authority citation; (iii) the date of publication; and (iv) the kind of names that have been applied by previous workers in the past.
of taxonomic change being made, which appears as a term after The style or convention used for the synonymy varies according
the year (Table 2). The authority citation is given by the person’s to the journal, but it is customary to cite in chronological order:
family name and the date as the year of publication; if there are (i) the species’ name; and for each name (ii) the bibliographic
two authors, both author names are cited and joined by ‘and’ or reference, including the author, date (year) of publication, and
an ampersand ‘&’. If there are more than three authors, then the page number(s) and/or figure or plate number(s). For example,
first author is followed by ‘et al.’. For taxonomic changes, the main Hay et al. [1994], on page 31 and plate 4, figures 7–9, referred to
terms commonly used are ‘comb. nov.’, ‘stat. rev.’, and ‘syn. nov.’. the species Ogyris arcana M.R.Williams & Hay, 2001 under the
6 • Braby et al.
name O. otanes C.Felder & R.Felder, 1865 (Table 2). Generally, in entomology), it may be prudent to be selective about the
the synonymy section only applies to established species that are number of paratypes designated or to list them in an appendix or
being revised and it is not required for new species. However, for as online supplementary material/supporting information.
a new species that was previously confused with another species,
there may be a history of names in which the species was pre- Other material: In studies where the status of a species has been
viously classified—these constitute misidentifications and they reviewed, revised, synonymized, or transferred to a different
ought to be included as ‘misidentifications in the literature’. genus, all other specimens examined and used in the differenti-
ation of that species should be listed after the type material. It is
Material examined important to list this material because it allows others to repeat

and test your observations at some point in the future (i.e. check
Type material: The type specimens examined should be listed the primary material evidence on which the data was obtained).
along with their deposition for all name-bearing types (holo-
type, lectotype, or neotype). When designating a holotype, the Description
specimen should be clearly labelled in a way that will unmistak- The description section is mandatory for new species, and it
ably identify it, and it must be accompanied by a statement of should be presented in telegraphic style in which non-essential
intent that it will be (or is) deposited in a collection, and a state- grammar (verbs and adjectives) is omitted to save space. The de-
ment indicating the name and location of that collection. Ideally, scription should include the standardized vocabulary and all the
it should be lodged and registered in a public institution that is essential taxonomic characters used as the standard in the tax-
a recognized repository (i.e. museum of national/international onomy of the group concerned (Winston 1999). For described
standing). Deposition of types in public institutions is not man- species that involve name changes, or where the original descrip-
datory for holotypes or lectotypes, only for neotypes, but it is tion was found to be inadequate, a redescription may be neces-
strongly recommended. The size, sex (if applicable), and devel- sary.
opmental stage should also be given. It is important to document
the label data [i.e. locality (including geographical coordinates), Diagnosis
altitude, date, collector name, and other collection details], The diagnosis should comprise a differential diagnosis that
registration number, and the name of the institution/collection clearly provides information on how the species differs from
in which the specimen is held. Publication of all information on most similar or closely related species based on at least one
the labels accompanying the holotype ensures that the specimen unique character state of a distinguishing character that is easily
will be recognized in future. recognizable (Rheindt et al. 2023). The diagnosis section is al-
Lectotypes and neotypes are treated similarly to holotypes. ways shorter than the description section. Winston (1999) and
If a lectotype is being selected from two or more syntypes, then Borkent (2021) recommend that the diagnosis section should be
the specimen must be clearly labelled to ensure recognition. For brief, consisting of a minimal set of statements that provide clear
lectotype designations to be valid, the term ‘lectotype’ must be and precise information that distinguish a given specimen from
used and the published work must contain a statement of the other species. That information is usually based on comparative
taxonomic purpose of the designation. In other words, the work morphological character states, but other evidence (e.g. DNA,
must state that a lectotype is being designated for the first time immature stages, behaviour, and biology) that distinguishes the
so that it is clear that the name is being fixed to the species, for species can be used, although sometimes it is better to include
example, by the words ‘new lectotype designation’ or ‘lectotype this information in the remarks section (Šlapeta 2013) but not
hereby designated’. in the discussion.
If no name-bearing type of a species-group taxon is believed
to be extant (i.e. the holotype, lectotype, or syntypes are lost or Remarks
destroyed), or if the existing name-bearing type cannot be deter- The taxonomic remarks or comments section is not mandatory,
mined so that the correct application of the name is in doubt (i.e. but it is highly recommended (Šlapeta 2013). It is generally used
the name is a nomen dubium), a single specimen may be desig- to make comments about the history of nomenclature, type ma-
nated as a neotype, which then becomes the name-bearing type. terial, and typification. Other information, such as history of dis-
However, designation of neotypes should be done only under ex- covery and distinguishing characters not listed in the diagnosis
ceptional circumstances in order to solve a complex taxonomic (e.g. immature stages, and reproductive structures), can be in-
problem, not as a matter of curatorial routine; besides that, often cluded in the remarks.
at times ‘lost’ types have been rediscovered, invalidating care-
lessly proposed neotypes. Etymology
Once the holotype or lectotype has been fixed, labelled, and The etymology section provides the derivation of the name, why
listed, it is essential to label all the remaining specimens exam- it was chosen, and how it was composed (Winston 1999). It is
ined, which automatically become paratypes or paralectotypes. only included for newly described taxa. Inclusion of the ety-
The label data and repository of all paratypes or paralectotypes mology is highly recommended but is not mandatory under the
should then be listed in the paper. These types have no name- Code, although many journals require it. There are four main
bearing function, but it is important to mention them because types of scientific names that taxonomists regularly use: (i)
they provide material evidence that was used to inform the ori- descriptive names; (ii) toponyms; (iii) eponyms; and (iv) mis-
ginal description, diagnosis, and extent of variation. If, however, cellaneous names. Descriptive names are used to describe the
there are large numbers of specimens (which frequently happens appearance of a species or pertain to some distinguishing feature
or combination of characters; they may be simple (as an adjec- (Cook et al. 2010). In addition to species’ concepts, authors may
tive, or as a noun in apposition), or they may be compound in have different views or opinions regarding the delineation of the
form (e.g. adverb-adjective, adjective-adjective, adjective-noun, species under investigation. These views may be either broad or
noun-noun). Toponyms refer to geographical location names or narrow. Taxonomists with a broad species’ concept (the so called
landmarks, and they may be constructed as a noun or as an ad- ‘lumpers’) tend to recognize fewer species and more synonyms
jective. Eponyms or patronyms are bestowed to commemorate and the term sensu lato or s.l. may be given after the species’ name
or honour a particular person, such as the discoverer (field col- or for the species complex, whereas taxonomists with a narrow
lector) or someone who has made a significant contribution to concept (the so called ‘splitters’) tend to recognize more species

the field of science. Miscellaneous names are essentially other and fewer synonyms and the term sensu stricto or s.s. may be ap-
names to the exclusion of categories (i)–(iii) listed above, such plied to the residual species’ name. Note, this is a different ap-
as cartoon or fictional characters that may nonetheless convey proach to polytypic species with several subspecies (subordinate
some properties of the species or an arbitrary combination taxa) in which the term s.l. may be used to refer to a species and
of letters of the Latin alphabet provided it is pronounceable all of its constituent subspecies, whereas the term s.s. refers only
with vowels. A good source for Latin names is Brown’s (1956) to the nominotypical subspecies.
Composition of Scientific Words.
DISCUSSION
OTHER ELE MENTS OF A TA XONOMIC PAPER Taxonomy is an essential activity based on the testing of hypoth-
A taxonomic paper should follow the standard format of a sci- eses and delimiting or refining species boundaries of organisms
entific research paper (Winston 1999). Thus, it still needs to (Mayr 1969, Mayr and Ashlock 1991, Yeates et al. 2011, Thiele
include an abstract, introduction, methods, discussion, acknow- et al. 2021). As new evidence is acquired, species boundaries
ledgements, and bibliography or references sections, and some- may need to be refined, which inevitably leads to name changes,
times the results section. The introduction should clearly state either through the delimitation and description of new species
the aims of the study, the reason for describing the new species or alteration to the taxonomic status of species (Table 1).
in the first place, and include sufficient background information Descriptive taxonomy is not a difficult science, but it is one
to place the work in context. The results section is particularly in which it is very easy to make mistakes (Winston 1999), per-
useful where the taxonomic revision is based on a phylogenetic haps because it can be a laborious, tedious, and time-consuming
hypothesis or morphometric data with statistical analyses. Some scientific discipline, but also because authors, and editors, are
journals now include the results section before the formal taxo- frequently unaware of basic taxonomic procedures (Šlapeta
nomic section. Keys to species are particularly useful for larger 2013). Work that contains errors cannot be retracted once it
revisionary works and actually help inform the diagnoses of is published, and either you or someone else will have to deal
each taxon (Borkent 2021). Several other sections are often in- with the consequences (and likely embarrassment!). Good tax-
cluded in a taxonomic publication; they are not essential but add onomy requires an understanding of nomenclature (rules of
value to the overall quality of the science. For instance, sections the Code), a sound knowledge of the historical literature, and
on variation (including sample size used to establish the range the whereabouts of type material. Before revising or describing
of variation), distribution, biology, ecology (e.g. host associ- anything as new, a few simple steps ought to be followed (Table
ations), and conservation status, if known, are very useful. The 3). First, conduct taxonomic research using all available speci-
discussion section is essential and must place the results in rela- mens and other resources. Once the scientific research is com-
tion to the aim or purpose of the study, any questions raised or pleted and taxonomic conclusions have been reached in relation
hypotheses advanced in the introduction, and place the study in to proposed species hypotheses (Table 1), then determine how
broader context of other work. A succinct summary explaining the names ought to be used (steps 2–4). A good understanding
how a particular taxonomic conclusion was reached based on of nomenclature necessitates being familiar with rules, recom-
evaluation of the available evidence is also very useful in the dis- mendations, and terminology of the ICZN (1999) (step 2).
cussion. Then develop a list of all available names that have been pub-
Few taxonomic papers cite the species’ concept adopted and lished based on review of the primary taxonomic literature of
the underlying criteria, and hence evidence required, to test the species and all similar or closely related species containing
those concepts (Cook et al. 2010, Yeates et al. 2011). The spe- the original descriptions of those species (step 3). It is crucial to
cies’ concept is important because different concepts can lead ensure that your species has not previously been named and de-
to vastly different conclusions based on the same evidence. For scribed, and your newly proposed name has not previously been
example, a set of metapopulations or lineage may be regarded as used—otherwise you may end up creating a junior synonym or
a single species under the biological species’ concept but may a junior homonym that is unavailable or invalid. Then locate and
be regarded as two or more sister-species under any one of the examine, and preferably photograph, the type specimens (if they
phylogenetic species’ concepts, which can have profound impli- exist), especially the name-bearing types, pertaining to those
cations for biodiversity conservation (Agapow et al. 2004, Isaac names so that the identity of each available name is clearly es-
et al. 2004, Balakrishnan 2005, de Queiroz 2007, Garnett and tablished (step 4). This step is crucial because it ensures that the
Christidis 2017, Thiele et al. 2021). Thus, it is important to state names and identities of species have been correctly established.
in the methods the species’ concept used so that it is clear to The next tasks (steps 5–10) are more operational and in-
the reader how the conclusions regarding species’ delimitation clude constructing the new species’ name correctly, designating
were reached based on interpretation of the available evidence a holotype and illustrating it, preparing the description and
8 • Braby et al.
Table 3. Guidelines for the description of a new species based on 11 practical steps (modified from Šlapeta 2013).
Step 1 Conduct taxonomic research and determine species boundaries in relation to testable and falsifiable hypotheses.
Step 2 Become familiar with rules, recommendations and terminology of the International Code of Zoological Nomenclature.
Step 3 Review primary taxonomic literature and collate all available names (i.e. previously described names) and descrip-
tions of similar species.
Step 4 Examine (and photograph) type material of available names of similar species.
Step 5 Construct new species’ name correctly.

Step 6 Select holotype (name-bearing type specimen) that is representative of new species.
Step 7 Prepare high quality illustrations of type material, especially holotype.
Step 8 Prepare description according to accepted taxonomic standards for the taxon group.
Step 9 Prepare differential diagnosis based on comparative evidence.
Step 10 Register manuscript (and preferably new species’ name) in ZooBank for electronic-only publication.
Step 11 Publish research findings in reputable taxonomic journal.
diagnosis, and registering the title and author(s) of the manu- intentions, noting that ‘In the future, a formal taxonomic de-
script in ZooBank for electronic-only publications. Finally, once scription and nomenclature will require the discovery of whole
the manuscript is complete, publish your findings in a reput- animals and a type specimen’, Manis mysteria Gu et al., 2023 is
able peer-reviewed scientific journal that has a track record of not available because it contravenes numerous provisions of the
publishing quality systematic works or an Editorial Board that Code, suggesting the authors were not aware of basic taxonomic
is familiar with taxonomic procedures (step 11). These publica- procedures regardless of the quality of their science, which was
tions take on many forms, from short papers describing a single published in the journal PNAS. Similarly, Lamb (2023) recently
species to systematic revisions and large monographs revising proposed the name Vanessa kiwa Lamb, 2023 for a putative new
numerous species within a higher taxonomic group. Although species of nymphalid butterfly from New Zealand’s North Island
there is no requirement for taxonomic works to be published (Poverty Bay). Although the name of the new species was latin-
in peer-reviewed scientific journals, scientific best practice is ized, there was no explicit intention to indicate that the taxon
to publish authoritative taxonomic papers in refereed journals was new (Article 16.1), no designation of a holotype (Article
(Wüster and Kaiser 2023)—the peer-review process is by far the 73), no explicit statement to indicate in which collection name-
best mechanism to control the quality of publications on tax- bearing type material is located (Article 16.4), and no descrip-
onomy. tion or differentiation (Article 13), just a vague comparison with
Despite these simple guidelines (Table 3), we have observed the two subspecies of Vanessa gonerilla (Fabricius, 1775) that
that a number of taxonomic publications in recent decades fall hardly justified a ‘diagnosis’. Beside the fact that the taxonomic
short of the basic procedures and minimum standards required literature was not reviewed or type material of V. gonerilla was
for naming new taxa. We provide some examples of a number examined, the name V. kiwa Lamb, 2023 is unavailable because
of pitfalls and mistakes to avoid in relation to several manda- the mandatory provisions of the Code were not followed. We ex-
tory provisions and recommendations of the Code and accepted pand on these and other non-compliance issues below.
standards of taxonomic best practice and journal editorial policy.
Our examples are mainly from the insect order Lepidoptera, par- Failure to review the primary taxonomic literature
ticularly butterflies (Papilionoidea), because this taxon group Failure to compile a list of available names (synonyms) based on
seems to be rife with ‘malpractice’, but examples from other review of the taxonomic literature can result in enormous prob-
groups with high levels of ‘malpractice’, such as reptiles and am- lems for other workers. For example, Miller (1995) published
phibians (Wüster et al. 2001, Kaiser 2013, 2014, Kaiser et al. a checklist of the neotropical Castniidae (Lepidoptera), but
2013, Wüster et al. 2021, Denzer and Kaiser 2023, Wüster and it became apparent that the primary taxonomic literature had
Kaiser 2023), are given. not been examined, which resulted in numerous inaccuracies,
including omissions of available names, misspellings, wrong au-
Failure to read the Code thority citations, incorrect dates of publication, and incorrect
Once the species boundary or delimitation has been established type localities (Lamas 1995). A consequence of this inaction
it is crucial to have a good working knowledge of the rules and was taxonomic misinformation and inflation—the 134 species
recommendations of the Code to ensure that new names are code recognized in Miller’s list were reduced to 81 species in Lamas’
compliant. However, it is apparent that some authors are simply list, largely due to synonymy.
not aware that a code of zoological nomenclature even exists. Article 23 of the Code sets out the Principle of Priority. The
For example, Gu et al. (2023) proposed the name Manis mysteria valid name of a taxon is the oldest available name (the senior
Gu et al., 2023 (Pholidota: Manidae) for a new cryptic species synonym) applied to it. The purpose of this provision is to pro-
of pangolin discovered in China; however, they neither clearly vide nomenclatural stability in cases in which a taxon has two
indicated that the name was intended to be new (Article 16.1) or more names bestowed upon it. This duplication of names
nor designated a holotype (Article 73), and, moreover, they did happened frequently in historical times due to taxonomists
not provided a description (Article 13). Despite the authors working on the same species in different places or countries, or
through the description of different sexes as different species. Failure to examine type material
The easiest way to avoid duplicating species’ names and creating Recommendation 73B (Preference for specimens studied by
nomenclatural confusion is to become familiar with all of the author) of the Code states ‘An author should designate as holo-
available names published in the taxonomic literature. However, type a specimen actually studied by him or her, not a specimen
even in modern times there are still cases in which known spe- known to the author only from descriptions or illustrations in
cies are being named and described. For example, Fujioka the literature’. Denzer and Kaiser (2023) recommended that
(1992) described the butterfly Laeosopis praetextatus Fujioka, best taxonomic practice is for authors to include a statement
1992 (Lycaenidae), but in the following year D’Abrera (1993) in their manuscript indicating that type specimens were phys-

described the same species under the name Laeosopis hoenei ically examined in order for their publications to be acceptable
D’Abrera, 1993. Under Article 23 of the Code, Fujioka’s name is for the purposes of zoological nomenclature. The main reasons
available and valid (senior synonym), whereas D’Abrera’s name for examining type material are to ensure that the nomen of the
is available but invalid (junior synonym). In other cases, syn- taxon has been correctly applied by subsequent workers, and that
onyms arise from competition between taxonomists working the name-bearing type is identifiable. If the name-bearing type is
on the same taxa, a practice strongly discouraged by the Code indeterminate, an author may request the Commission, under its
under its Code of Ethics. For example, Yago (2004) was under- plenary power, to replace the unidentifiable name-bearing type
taking a systematic revision of the butterfly subtribe Horagina with a neotype (Article 75.5). However, there are numerous ex-
(Lycaenidae) and the finished work included the description of amples in the literature in which authors undertaking taxonomic
a new species, Horaga uedai Yago, 2004, which was published on revisions fail to examine name-bearing type material. Such ac-
20 January 2004. However, Saito and Seki [2004] described the tions are not in breach of the Code, but it is poor taxonomic
same species, under the name Horaga takanamii Seki & Saito, practice not to examine type material. For example, in a system-
[2004], a few weeks earlier by side-stepping the peer-review pro- atic revision of the butterfly genus Taractrocera Butler, [1870]
cess and publishing in a magazine [issue Number 38 of Butterflies (Hesperiidae) based on comparative morphology, de Jong
was released to its members on 1 January 2004 (U. Yoshinobu, (2004) did not examine type material of the species Taractrocera
pers. comm.)]. Thus, the valid name for the species is Horaga ilia Waterhouse, 1932. At that time the species was considered to
takanamii Seki & Saito, [2004] (senior synonym) and not be polytypic, composed of two subspecies: the nominotypical
Horaga uedai Yago, 2004 (junior synonym). subspecies from northern Australia, and T. ilia beta Evans, 1934
Article 52 of the Code sets out the Principle of Homonymy. from mainland New Guinea. De Jong (2004) only examined ma-
The purpose of this provision is to ensure that two or more dif- terial of the subspecies T. ilia beta for the diagnosis of the species
ferent taxa are not denoted by the same name. In such cases, T. ilia. More seriously, types of neither subspecies were exam-
only the oldest name (the senior homonym) can be used ined. Subsequent examination and comparison of the types of
as the valid name. For example, Lambert et al. (2010a) de- both subspecies revealed that T. beta Evans, 1934 is, in fact, a spe-
scribed a new genus of fossil sperm whale from the Miocene cies distinct from T. ilia (Braby and Zwick 2015), leading to an
of Peru in the prestigious journal Nature, not realizing that the erroneous diagnosis of T. ilia in the earlier study.
name Leviathan Lambert et al., 2010 (junior homonym) was
pre-occupied by a mammoth Leviathan Koch, 1841 (senior Failure to construct the taxon name correctly
homonym), thus rendering their new genus name unavailable. Articles 31–34 of the Code deal with the formation of species-
The authors subsequently published a corrigendum Lambert et group names, original spellings, and subsequent spellings. The
al. (2010b) and proposed Livyatan Lambert et al., 2010 as a re- scientific name (genus- and species-group) must be in Latin
placement name for Leviathan Lambert et al., 2010. In another or latinized (Article 11.9), with correct endings when formed
example, Fujioka (1993) described the butterfly subspecies from personal names (Article 31.1). However, some authors
Ravenna nivea howarthi Fujioka, 1993 (Lycaenidae) in a paper fail to follow these rules. For example, the snake Acanthophis
dated 15 April 1993, not realizing that the same name had been wellsei Hoser, 1998 was described in honour of Richard Wells.
used for the same taxon a month earlier by Saigusa (1993), as However, this name was improperly constructed because under
Ravenna nivea howarthi Saigusa, 1993 published on 5 March the Code the genitive ending for a man is ‘i’ and for a woman
1993. Thus, Ravenna nivea howarthi Fujioka, 1993 is a junior is ‘ae’ (Article 31.1.2). The name was subsequently redescribed
homonym (and a junior synonym) of the valid name Ravenna and emended to Acanthophis wellsi Hoser, 1998 by Aplin and
nivea howarthi Saigusa, 1993, which is the senior homonym Donnellan (1999), although this action is an unjustified emend-
(and senior synonym). ation, and the name A. wellsi should be regarded as an incorrect
To avoid creating junior synonyms or junior homonyms, subsequent spelling and, therefore, unavailable (Articles 33.3,
three useful resources to check for names are: 33.4), even though the original spelling was erroneous. Other
examples of erroneous endings constructed by herpetologists
• Nomenclator Zoologicus (Neave 1939–40, 1950–1975): that required emendation (Articles 33.2, 50.4) were reviewed by
http://insecta.bio.spbu.ru/z/nomenclator_zoologicus_ Wüster et al. (2001).
PDF.htm
• Zoological Record: https://clarivate.com/products/ Ethics: Some names are particularly cumbersome or poorly
scientific-and-academic-research/research-discovery- constructed and difficult to pronounce. Long-winded names
and-workflow-solutions/webofscience-platform/web- such as the amphipod Gammaracanthuskytodermogammarus
of-science-zoological-records/ loricatobaicalensis Dybowski, 1926 are not particularly helpful
• Google Scholar: https://scholar.google.com/ in terms of providing clear and unambiguous communication.
10 • Braby et al.
Although G. loricatobaicalensis was technically available under Hesperillini Grund, 1998; Trapezitini Grund, 1998; Proeidosini
the Principle of Binominal Nomenclature (Article 5), the work Atkins, 2005; Mesodinina Atkins, 2012; Hesperillina Atkins,
where it was published was suppressed for nomenclatural pur- 2012; Toxidina [sic] Atkins, 2012; Trapezitini Atkins, 2012;
poses by the ICZN under its plenary powers (Opinion 105, Anisyntina Atkins, 2017; Proeidosina Atkins, 2017; Toxidini
Direction 32), and thus the name is unavailable. The Code recom- [sic] Atkins, 2017. However, for each higher taxon no type genus
mends that ‘Authors should exercise reasonable care and consid- was given. Thus, these names are not available because the pro-
eration in forming new names to ensure that they are chosen posals contravene the rules of the Code for typification of types
with their subsequent users in mind and that, as far as possible, in the family group (Article 63. Name-bearing types) and names

they are appropriate, compact, euphonious, memorable, and do published after 1999 (Article 16.2. Family-group names: type
not cause offence’ (Recommendation 25C. Responsibility of au- genus to be cited). There were also other issues with these names,
thors forming new names). Although it is not explicitly stated in for instance, the authors failed to provide a diagnosis (nomina
the Code, it is considered unethical to name a species after your- nuda—see diagnosis below), or construct the name correctly, or
self, but it does happen. For example, Wall (1907) named the the names were junior synonyms and homonyms (Toussaint et al.
snake Bungarus walli Wall, 1907 after himself and, more recently, 2022). For instance, Hesperillini Grund, 1998 and Hesperillina
Kaiser et al. (2013) identified an author who named no less than Atkins, 2012 are junior synonyms and homonyms of Hesperillini
43 eponyms that included his surname. Voss, 1952, and Trapezitini Grund, 1998 and Trapezitini Atkins,
2012 are junior synonyms and homonyms of Trapezitinae
Eponyms: Recent proposals to change names, particularly ep- Waterhouse & Lyell, 1914.
onyms, on ethical grounds (Hammer and Thiele 2021, Guedes At the species-level, the name-bearing types (holotype,
et al. 2023), or to replace long-established scientific names with lectotype, neotype, or syntypes) are particularly important be-
new indigenous names (Gillman and Wright 2020), have been cause they fix the name to the species (Articles 72–75). When
rejected by the taxonomic community (Ceríaco et al. 2023). describing a new species, it is now mandatory to designate a
Such proposals contravene the fundamental aim of the Code, holotype. Furthermore, it is advisable to lodge the type spe-
which is to promote nomenclatural stability and universality cimen in an institutional collection and give the name of the
without constraining taxonomic judgement. institution in which it has been deposited (Recommendation
16C. Preservation and deposition of type specimens). When
Failure to explicitly establish the new taxon designating a holotype for a new species, it is essential to se-
Article 16.1 of the Code states that every new name published after lect a specimen that is representative of the species, that is, one
1999 must be explicitly indicated as intentionally new. Authors that shows most, or all of the diagnostic characters of the spe-
are advised to make their intentions explicit by using in the cies. Other type material should also be listed correctly and
heading of new names abbreviations of Latin terms, such as ‘fam. referred to as paratypes, not as ‘cotypes’ (Recommendation
nov.’, ‘gen. nov.’, ‘sp. nov.’, ‘ssp. nov.’, or an equivalent expression 73E. Avoidance of the term ‘cotype’) or ‘topotypes’ (speci-
(e.g. ‘species nova’, ‘new species’). Failure to follow Article 16.1 mens originating from the type locality), although the Code
will render the new name(s) invalid. For example, Stradomsky does allow use of the term ‘allotype’ to indicate a specimen of
(2016) proposed several new subtribes of polyommatine butter- opposite sex to the holotype (Recommendation 72A. Use of
flies (Lycaenidae) (i.e. Pithecopina Stradomsky, 2016, Azanina the term ‘allotype’). Failure to provide adequate typification
Stradomsky, 2016, Theclinesthina Stradomsky, 2016, Jamidina will render the name unavailable. Moreover, failure to comply
Stradomsky, 2016, Uranothaumatina Stradomsky, 2016, with the requirements of Article 16.4 (fixation of name-bearing
Zizulina Stradomsky, 2016, Oboroniina Stradomsky, 2016, and types for species-group names to be explicit) will make names
Fameganina Stradomsky, 2016), but he did not explicitly indi- published after 1999 unavailable—there are several cases of
cate that the names were intentionally new, thus failing Article authors who have forgotten to provide statements about the
16.1 (All names: intention of authors to establish new nominal name(s) and location(s) of the collection(s) where the holo-
taxa to be explicit). Moreover, he did not cite the type genus for types are (or will be) deposited. For example, Viloria et al.
each taxon, thus failing Article 16.2 (see typification below). (2003) proposed a new species of butterfly, Redonda bordoni
Viloria & Pyrcz, 2003 (Nymphalidae), from the high Andes of
Failure to provide adequate typification South America in the Proceedings of the Royal Society of London.
Articles 61–75 of the Code deal with the Principle of Typification, They provided a diagnosis; however, the description was rele-
which applies to nominal taxa in the family, genus, and species gated to an online appendix, which is no longer accessible.
groups. For family-group names, the name-bearing type is the More seriously, however, no type data and no repository for
‘type genus’ (Article 63); for genus-group names, the name- type material were provided. Thus, the name Redonda bordoni
bearing type is the ‘type species’ (Article 67); and for species- Viloria & Pyrcz, 2003 is unavailable because it does not satisfy
group names, the name-bearing type is the ‘holotype’ or syntypes the provisions of Article 16.4.
(Article 73), ‘lectotype’ (Article 74) or ‘neotype’ (Article 75).
For all taxa described after 1930, failure to designate a type Type material: We recommend that types be based on preserved
genus for a new family-group name or a type species for a new material deposited in collections, rather than on photographs of
genus-group name will render the taxon name unavailable. For live animals (i.e. photography-based taxonomy—Donegan 2008,
example, Grund (1998) and Atkins (2005, 2012, 2017) collect- Pape 2016, Thorpe 2017), because specimens are the primary
ively proposed 11 tribal and subtribal names for the butterfly sub- evidence on which names are based. They also provide far more
family Trapezitinae (Hesperiidae), viz: Mesodinini Grund, 1998; characters for an objective diagnosis and description, evidence
that can be verified and replicated by future workers, than sec- information to allow anyone to distinguish the new species from
ondary evidence from photographs (Dubois and Nemésio 2007, similar or closely related species (Rheindt et al. 2023). However,
Krell and Wheeler 2014, Rocha et al. 2014, Ceríaco et al. 2016, some authors fail to provide this information. For example, in re-
Krell and Marshall 2017). Perhaps the most notorious example vising the diverse Neotropical butterfly genus Catasticta Butler,
of photography-based taxonomy is the description of the in- 1870 (Pieridae), Reissinger (1972) divided the genus into sev-
famous Loch Ness Monster Nessiteras rhombopteryx Scott & eral subgenera, of which four were new (Archonoia Reissinger,
Rines, 1975, a hoax published in Nature. Irrespective of the or- 1972, Pierinoia Reissinger, 1972, Leodontoia Reissinger,
ganisms lack of biological reality (Halstead et al. 1976, Lawton 1972, and Hesperochoia Reissinger, 1972). However, Reissinger

1996), Scott and Rines (1975) provided no name-bearing type, (1972) did not provide descriptions or diagnoses for them
with the entire description based on a rather poor set of photo- and thus they are nomenclaturally unavailable (Wojtusiak
graphs of purported animal parts of a plesiosaur-like reptile, all of 1998, Lamas and Bollino 2004). Subsequently, Eitschberger
which have since proven to be fraudulent (Campbell 1986, Ohl and Racheli (1998) recognized two of Reissinger’s names,
2018). One of the cornerstones of scientific work is repeatability, Leodontoia and Hesperochoia, as distinct genera and provided
and specimens lodged in a museum allow other researchers to putative distinguishing characters, thus inadvertently making
examine them so that the observations can be repeated. For in- them available. Therefore, the available names are now credited
stance, if future research reveals that taxonomic changes need to to Eitschberger and Racheli (1998), as Leodontoia Eitschberger
be made (e.g. presence of a second, cryptic species based on new & Racheli, 1998 and Hesperochoia Eitschberger & Racheli, 1998,
evidence), the holotype specimen can be re-examined for verifi- rather than to Reissinger (1972). Similarly, Smart (1975) estab-
cation and acquisition of additional data, a task that may not be lished the subgeneric name Antipodolycaena Smart, 1975 for the
possible from photographs (Ceríaco et al. 2016). Museum speci- New Zealand coppers Lycaena Fabricius, 1807 (now placed in
mens also provide opportunities for future research in ways yet the genus Boldenaria Zhdanko, 1995). Smart (1975) published
to be realized with the emergence of new technologies (Krell and the name in an appendix of a popular book and listed Lycaena
Wheeler 2014, Rocha et al. 2014). However, we acknowledge boldenarum White, 1862 as the type species, but he did not pro-
that the preservation of complete specimens may not always be vide a diagnosis or description (he just stated ‘… certain small
possible, particularly for threatened species that have exceedingly but significant differences in pattern element, particularly on the
low population numbers, such as the recently discovered, and de- undersides...’). Hence, Antipodolycaena Smart, 1975 is not avail-
scribed, Galapagos pink land iguana Conolophus marthae Gentile able because it does not satisfy the provisions of Article 13.
& Snell, 2009 from the Galapagos Islands (Donegan 2009, Gentile Many authors of taxonomic papers prepare the diagnosis as
and Snell 2009), although in such cases the collection of a single or a shortened version of the description without comparison to
a few specimens to serve as types is not likely to cause extinction other similar species, and thus the diagnoses fail to be contrastive
(Rocha et al. 2014). Type specimens should never be permanently and/or state-specific (Rheindt et al. 2023). We discourage such
deposited in private collections due to the risk of them being lost poor taxonomic practice because the goal of a differential diag-
or destroyed when the owner of the collection dies or the collec- nosis is to clearly specify which character states, of one or more
tion is broken up (Knapp et al. 2004). Some journals demand that characters, distinguish the species from other species.
type and other specimens are lodged in public institutions.
Failure to register the publication in ZooBank
Failure to differentiate the new taxon For electronic publications, the work describing the new name
Article 13 of the Code sets out the requirements for new names must be registered in ZooBank for it to be available (Yeates 2009,
published after 1930. For new names to be available they must ‘be Rosenberg et al. 2012, Zhang 2012, Krell and Pape 2015, Braby
accompanied by a description or definition that states in words 2018). However, following the amendment of the Code by the
characters that are purported to differentiate the taxon’ (Article ICZN (2012), there have been hundreds, if not thousands, of ex-
13.1.1). Further, the Code goes on to say that ‘When describing amples of new species published in e-journals where this has not
a new nominal taxon, an author should make clear his or her pur- happened (M.S. Harvey, pers. comm.). For example, the paper
pose to differentiate the taxon by including with it a diagnosis, that describing a new species of frog (Buergeria otai) from Taiwan in
is to say, a summary of the characters that differentiate the new PLoS ONE (Wang et al. 2017) was not registered in ZooBank
nominal taxon from related or similar taxa’ (Recommendation and thus the name is not available. The authors subsequently re-
13A. Intent to differentiate). Names published after 1930 that submitted to the same journal a correction (Wang et al. 2018),
fail to conform to Article 13 are not available and are known as which included the ZooBank registration number, but they
nomina nuda (singular nomen nudum). Thus, for a new species, inadvertently omitted details concerning the description and
the description and diagnosis need to be useful and to convince typification, so that the name constituted a nomen nudum (an
the reader that the new species is biologically meaningful and can unavailable name—see diagnosis above) with no holotype (see
be justified. That justification is based on evidence, which is the typification above). The authors finally made the name of their
data used to test species boundaries and to delimit the species. new species available by sidestepping the ZooBank registra-
This evidence may include morphological, molecular (Tautz tion altogether and publishing the description (in Chinese and
et al. 2003, Cook et al. 2010), biological, ecological, or behav- English) in the print (hard copy) magazine Nature Conservation
ioural characters, and ideally an integrative approach that com- Quarterly (Wang et al. 2020). Hence, the correct nomencla-
bines traits from all of these data sources (Will et al. 2005, Wiens ture for the species is Buergeria otai Wang et al., 2020 and not
2007). In other words, the diagnosis should provide sufficient Buergeria otai Wang et al., 2017, but it took three attempts!
12 • Braby et al.
Failure to publish the new taxon name 2000 (see reviews by: Aplin 1999, Wüster et al. 2001, Williams
Article 8 of the Code sets out the criteria to be met for new names et al. 2006, Kaiser et al. 2013, Kaiser 2014, Wüster et al. 2021).
to be published. A published work ‘Must be issued for the purpose Although the names are technically available according to the
of providing a public and permanent scientific record, must be rules of the Code, the robust scientific evidence upon which they
obtainable free of charge or by purchase, and must have been pro- should have been established was fundamentally lacking—for
duced in an edition containing simultaneously obtainable copies instance, the descriptions and diagnoses were inadequate, often
by a method that assures numerous identical and durable copies’ failing to demonstrate the biological validity or the reality of the
(Article 8.1). Further, the Code recommends that new scientific species concerned, or the authors designated type specimens

names ought to be published in journals that will be captured that they had not examined (contravening Recommendation
by the Zoological Record database. Names printed in abstracts 73B), such as through nomenclatural harvesting of phylogen-
or posters at conference meetings or congresses, or manuscript etic clades (Denzer and Kaiser 2023). Such mass-naming or
names placed on a label attached to a specimen, or photographs, mass-changing of taxa based on poor or non-scientific practice
or names posted on the Internet are considered unpublished and, is known as ‘taxonomic vandalism’ (Kaiser et al. 2013, Wüster
therefore, are not available. For example, C. Lee coined the name et al. 2021). Proposals to declare names and nomenclatural
Luehdorfia longicaudata for a new and endangered butterfly from acts published in controversial taxonomic works as unavailable
the Qinling Mountains in China in an oral presentation at the through the Code’s requirements for publication have been re-
28th annual meeting of the Lepidopterological Society of Japan jected by the ICZN because the proposals are a direct violation
in 1981, but the name is not available since it was not published of the Code (Krell 2021). Instead, such works are best resolved
according to the rules of Article 8. However, the following year, by prevailing usage within the scientific community who will
an abstract of his lecture concerning the geographical distribu- either accept or reject the proposed taxonomy. In the case of rep-
tion and habitat of the new species, together with a brief descrip- tiles, the herpetological community has recommended not to
tion and comparison with two other species of Luehdorfia Krüger, adopt the names proposed by ‘taxonomic vandals’ (Kaiser 2013,
1878 from China, appeared in print in a non-peer-reviewed news 2014, Kaiser et al. 2013). Moreover, Wüster et al. (2021) have
bulletin (translated in Japanese) (Lee 1982). Thus, the valid proposed a new concept to deal with such extreme acts of taxo-
name for this species is Luehdorfia longicaudata Lee, 1982 and nomic vandalism—they coined the term ‘aspidonyms’, which
not Luehdorfia longicaudata Lee, 1981 or Luehdorfia taibai Chou, are scientific names proposed to overwrite unscientific names
1984, a proposed replacement name (nomen novum) (Inomata established by non-scientific methods as a last resort to stabilize
1995), even though the publication process was convoluted and scientific nomenclature. The aspidonyms are either senior syn-
scientifically poor. onyms or replacement names (junior synonyms), but it remains
The Code does not explicitly exclude new names from avail- to be seen what position the ICZN adopts on the matter of taxo-
ability in higher degree theses and dissertations, but most tax- nomic vandalism.
onomists have generally assumed that they do not constitute
published works within the provisions of Article 8 of the Code Fake holotypes: Fabricated or fictitious types based on fake species
unless they are issued ‘For the purpose of providing a public and are particularly serious and hinder the advancement of scientific
permanent scientific record’ (Article 8.1.1); they are ‘Obtainable, knowledge—they simply add to the burden of junior synonyms
when first issued, free of charge or by purchase’ (Article 8.1.2); that someone else has to resolve (Ohl 2018). Yet, there are several
and they are ‘Produced in an edition containing simultaneously examples of species, especially butterflies, in which new species
obtainable copies by a method that assures numerous identical have been described and published based on fakes—we know of
and durable copies’ (Article 8.1.3). If these provisions are not at least nine in the past few decades ( Johnson and Matusik 1988,
satisfied, authors should regard theses and dissertations as un- 1989, Okano 1989, Johnson 1990, 1992, Johnson et al. 1990,
published work. 1997). In these cases, the holotypes of the fake species were ac-
tually composite specimens, created by joining parts belonging
Other issues of ‘malpractice’ to different species, often belonging to different genera (and
Taxonomic vandalism: Taxonomy is perhaps the only field of sci- sometimes from different biogeographical regions). Under the
ence in which there is no requirement for its scientific products provisions of Article 1.3 of the Code, names of species based on
to be published through the critical peer-review process (Wüster hypothetical concepts may be excluded (i.e. a taxonomic concept
et al. 2021). Although there have been proposals to change the that when published contained no animal then known to exist
rules of the Code so that new species can be described only in nature, past or present). In addition, under the provisions of
through a formal accreditation process, such as in peer-reviewed Article 73.1.5 of the Code, parts of the chimeric holotypes may be
journals (Borrell 2007, Yeates 2009, Kaiser 2013, Kaiser et al. excluded from the holotype to clarify identification of the name
2013, Denzer and Kaiser 2023, Wüster and Kaiser 2023), these in order to recognize it as a junior synonym (Yagishita et al. 1993,
recommendations have yet to be endorsed and implemented by Peggie et al. 1995, Robbins and Nicolay 1999, Robbins and Lamas
the ICZN. Thus, taxonomic works may be published privately 2002, Bálint and Benyamini 2020). In other cases, the holotypes
in self-published works, which potentially opens the door to of fictitious species were manufactured by applying paint to spe-
poor-quality work or even non-scientific work. For example, in cimens of known species to give an entirely different appearance
the past four decades (since 1984) over 2000 new names of rep- (Vane-Wright and Whalley 1985) or the holotype was stolen and
tiles and amphibians have been published in non-accredited out- replaced with a fake comprising a painted specimen of a different
lets by two authors, of which c. 90% have been published since species (Braby and Eastwood 2019).
CONCLUSION phylogeny of the Hesperiidae. News Bulletin of the Entomological

Society of Queensland 2005;33:24–34.
Taxonomy lays the foundation for all of the biological sciences, Atkins AF. A list of butterflies (Papilionoidea) and skipper butterflies
including ecology, evolutionary biology, biodiversity conserva- (Hesperioidea) found within a 10km diameter area centered on the
tion, biosecurity, primary industry, medicine, and public health. village of Eudlo in the hinterland of the sunshine coast with a note
However, the prevalence of poor taxonomic practice and even on their migratory habits. Metamorphosis Australia. Magazine of the
‘malpractice’ (taxonomic vandalism, fictitious species based on Butterfly and Other Invertebrates Club 2012;66:4–11.
Atkins AF. Skipper butterflies (Hesperoidea [sic]) and bar codes: how old
fake types) indicates a need for consistency in the procedure are they, and how do the Aussie species fit in to the scheme of things?
and quality of taxonomic research publications dealing with (Part one: the sub-family Trapezitinae and their ilk). Metamorphosis

the description of new species, changes of species’ names (i.e. Australia. Magazine of the Butterfly and Other Invertebrates Club
synonymy, status revision, new combination), and nomencla- 2017;86:4–21.
tural acts (typification). Authors (and editors) need to be aware Bacher S. Still not enough taxonomists: reply to Joppa et al. Trends in
Ecology and Evolution 2012;27:65–6; author reply 66. https://doi.
of basic taxonomic procedures and be familiar with the sets of
org/10.1016/j.tree.2011.11.003
rules and principles at four different levels: (i) articles of the cur- Balakrishnan R. Species concepts, species boundaries and species identi-
rent International Code of Zoological Nomenclature; (ii) recom- fication: a view from the tropics. Systematic Biology 2005;54:689–93.
mendations of the Code; (iii) accepted standards of taxonomic https://doi.org/10.1080/10635150590950308
best practice of the taxon group; and (iv) journal editorial policy Bálint Z, Benyamini D. The identity of the nomen dubium Penaincisalia
and author guidelines. Robust taxonomy requires an under- patagonaevaga Johnson, 1990 (Lepidoptera: Lycaenidae,
Polyommatinae). Opuscula Zoologica 2020;51:213–7. https://doi.
standing of nomenclature (rules and recommendations of the org/10.18348/opzool.2020.2.213
Code), familiarity of the primary taxonomic literature, and the Beaver EP, Braby MF, Glatz RV et al. Systemtatic revision of the Ogyris
whereabouts of name-bearing type material, which ought to idmo (Hewitson, 1862) species group (Lepidoptera: Lycaenidae):
be physically examined. Moreover, descriptions of new species implications for the conservation management of Australia’s most
should construct the taxon name correctly, explicitly establish threatened butterflies. Invertebrate Systematics 2023;37:677–701.
the new taxon, provide adequate typification, clearly differen- https://doi.org/10.1071/is23032
Borkent A. Diagnosing diagnoses—can we improve our tax-
tiate the new taxon, and publish the name of the new taxon in a onomy? Zookeys 2021;1071:43–8. https://doi.org/10.3897/
manner that is compliant with the Code, preferably in an authori- zookeys.1071.72904
tative peer-reviewed scientific journal. It is recommended that Borrell B. The big name hunters. Nature 2007;446:253–5. https://doi.
the taxonomic section of a taxonomic publication include the org/10.1038/446253a
following elements: classification header (including type species Braby MF. Are taxonomic publications involving nomenclatural acts
of the genus-group name); species’ name header in binominal on early view code compliant? Austral Entomology 2018;57:371–6.
https://doi.org/10.1111/aen.12372
form, followed by the term ‘sp. nov.’ or ‘sp. n.’ for new species, Braby MF, Eastwood RG. Revised taxonomic status of Pseudalmenus
or the authority citation, date of publication, and kind of taxo- barringtonensis Waterhouse, 1928 stat. rev. (Lepidoptera: Lycaenidae):
nomic change being made for described species being revised; uncovering Australia’s greatest taxonomic fraud. Invertebrate
figure references (illustrations of name-bearing type), ZooBank Systematics 2019;33:530–43.
registration number (for papers published in electronic-only Braby MF, Williams MR. Biosystematics and conservation biology: crit-
ical scientific disciplines for the management of insect biological di-
journals), type locality, and the synonymy section; material versity. Austral Entomology 2016;55:1–17. https://doi.org/10.1111/
examined (including type material, and other material); descrip- aen.12158
tion; diagnosis; remarks (including, but not limited to, history Braby MF, Zwick A. Taxonomic revision of the Taractrocera ilia
of nomenclature, type material, typification, and history of dis- (Waterhouse) complex (Lepidoptera: Hesperiidae) from
covery); and the etymology of new taxa. north-western Australia and mainland New Guinea based on mor-
phological and molecular data. Invertebrate Systematics 2015;29:487–
509. https://doi.org/10.1071/is15028
ACKNOWLEDGE MENTS Braby MF, Eastwood RG, Murray N. The subspecies concept in butterflies:
has its application in taxonomy and conservation biology outlived its
We are most grateful to Hinrich Kaiser, Bruce Halliday, and Mark usefulness? Biological Journal of the Linnean Society 2012;106:699–
Harvey for reviewing the manuscript, providing constructive advice 716. https://doi.org/10.1111/j.1095-8312.2012.01909.x
and substantially improving the quality of this work. Brown RW. Composition of Scientific Words. A Manual of Methods and
a Lexicon of Materials for the Practice of Logotechnics, revised edn.
Washington, DC: Smithsonian Institution Press, 1956.
REFERENCES Campbell S. The Loch Ness Monster: The Evidence. Wellingborough,
Agapow P-M, Bininda-Emonds ORP, Crandall KA et al. The impact of Northamptonshire: Aquarian Press, 1986, 128.
species concept on biodiversity studies. The Quarterly Review of Ceríaco LMP, Gutiérrez EE, Dubois A. Photography-based taxonomy is
Biology 2004;79:161–79. https://doi.org/10.1086/383542 inadequate, unneccessary, and potentially harmful for biological sci-
Aplin KP. ‘Amateur’ taxonomy in Australian herpetology—help or ences. Zootaxa 2016;4196:435–45.
hindrance? Monitor, Journal of the Victorian Herpetological Society Ceríaco LMP, Aescht E, Ahyong ST et al. Renaming taxa on ethical
1999;10:104–9. grounds threatens nomenclatural stability and scientific communica-
Aplin KP, Donnellan SC. An extended description of the Pilbara death tion. Zoological Journal of the Linnean Society 2023;197:283–6.
adder, Acanthophis wellsi Hoser (Serpentes: Elapidae), with notes Cook LG, Edwards RD, Crisp MD et al. Need morphology always
on the desert death adder, A. pyrrhus Boulenger, and identification be required for new species descriptions? Invertebrate Systematics
of a possible hybrid zone. Records of the Western Australian Museum 2010;24:322–6. https://doi.org/10.1071/is10011
1999;19:277–98. Costello MJ, May RM, Stork NE. Can we name Earth’s species before
Atkins AF. Skipper butterflies: their origins? A preliminary discussion they go extinct? Science 2013;339:413–6. https://doi.org/10.1126/
on the higher classification of the higher systematics and a proposed science.1230318
14 • Braby et al.
Costello MJ, Lane M, Wilson S et al. Factors influencing when species Isaac NJB, Mallet J, Mace GM. Taxonomic inflation: its influence on
are first named and estimating global species richness. Global Ecology macroecolgy and conservation. Trends in Ecology and Evolution
and Conservation 2015a;4:243–54. https://doi.org/10.1016/j. 2004;19:464–9. https://doi.org/10.1016/j.tree.2004.06.004
gecco.2015.07.001 Johnson K. Penaincisalia, a new genus of ‘elfin’-like butterflies from the
Costello MJ, Vanhoorne B, Appeltans W. Conservation of biodiversity high Andes (Lepidoptera: Lycaenidae). Pan-Pacific Entomologist
through taxonomy, data publication, and collaborative infrastructures. 1990;66:97–125.
Conservation Biology 2015b;29:1094–9. https://doi.org/10.1111/ Johnson K. Genera and species of the Neotropical ‘elfin’-like hairstreak
cobi.12496 butterflies (Lepidoptera, Lycaenidae, Theclinae). Reports of the
D’Abrera B. Butterflies of the Holarctic Region. Part III Nymphalidae Museum of Natural History, University of Wisconsin 1992;22:1–279.
(concl.), Libytheidae, Riodinidae & Lycaenidae. Victoria: Hill House, Johnson K, Matusik D. Five new species and one new subspecies of butter-

1993, 335–524, i–vii. flies from the Sierra de Baoruco of Hispaniola. Annals of the Carnegie
Denzer W, Kaiser H. Naming and gaming: The illicit taxonomic practice Museum 1988;57:221–54. https://doi.org/10.5962/p.215185
of ‘nomenclatural harvesting’ and how to avoid it. Journal of Zoology Johnson K, Matusik D. Addendum: a new species of Tmolus (Lycaenidae)
2023;320:161–8. https://doi.org/10.1111/jzo.13061 from Hispaniola. In: Schwartz A (ed.), The butterflies of Hispaniola.
Donegan TM. New species and subspecies descriptions do not and should Gainesville: University of Florida Press, 1989, 525–9.
not always requrie a dead type specimen. Zootaxa 2008;1761:37–48. Johnson K, Eisele RC, MacPherson BN. The ‘hairstreak butterflies’
Donegan TM. Type specimens, samples of live individuals and the (Lycaenidae, Theclinae) of north-western Argentina. II. Strymon,
Galapagos Pink Land Iguana. Zootaxa 2009;2201:12–20. sensu stricto. Bulletin of the Allyn Museum 1990;130:1–77.
Dubois A. Proposed rules for the incorporation of nomina of higher- Johnson K, Austin GT, Le Crom J-F et al. The Strephonina, a new
ranked zoological taxa in the International Code of Zoological infratribe of the Eumaeini with description of fourteen new genera
Nomenclature. 1. Some general questions, concepts and terms of bio- (Lycaenidae). Revista de Theclinae Colombianos 1997;1:1–61.
logical nomenclature. Zoosystema 2005;27:365–426. de Jong R. Phylogeny and biogeography of the genus Taractrocera Butler,
Dubois A, Nemésio A. Does nomenclatural availability of nomina of new 1870 (Lepidoptera: Hesperiidae), an example of Southeast Asian-
species or subspecies require the deposition of vouchers in collec- Australian interchange. Zoologische Mededelingen 2004;78:383–415.
tions? Zootaxa 2007;1409:1–22. Joppa LN, Roberts DL, Pimm SL. The population ecology and social be-
Eitschberger U, Racheli T. Catasticta studies (Lepidoptera: Pieridae). haviour of taxonomists. Trends in Ecology and Evolution 2011;26:551–
Neue Entomologische Nachrichten 1998;41:4–94. 3. https://doi.org/10.1016/j.tree.2011.07.010
Fujioka T. Zephyrus (Theclini butterflies) in the world (2)—genus Kaiser H. The Taxon Filter, a novel mechanism designed to facilitate
Ussuriana and Laeosopis. Butterflies (Tokyo) 1992;2:3–18. the relationship between taxonomy and nomenclature, vis-à-vis the
Fujioka T. Zephyrus (Theclini butterflies) in the world (3)—genus utility of the Code’s Article 81 (the Commission’s plenary power).
Antigius and allied genera. Butterflies (Tokyo) 1993;4:3–20. Bulletin of Zoological Nomenclature 2013;70:293–302. https://doi.
Garnett ST, Christidis L. Taxonomy anarchy hampers conservation. org/10.21805/bzn.v70i4.a17
Nature 2017;546:25–7. https://doi.org/10.1038/546025a Kaiser H. Best practices in herpetological taxonomy: errata and addenda.
Gentile G, Snell H. Conolophus marthae sp. nov. (Squamata, Iguanidae), Herpetological Review 2014;45:257–68.
a new species of land iguana from the Galápagos archipegalo. Zootaxa Kaiser H, Crother BI, Kelly CMR et al. Best practices; in the 21st century,
2009;2201:1–10. taxonomic decisioins in Herpetology are acceptable only when sup-
Gillman LN, Wright SD. Restoring indigenous names in taxonomy. ported by a body of evidence via peer-review. Herpetological Review
Communications Biology 2020;3:609. https://doi.org/10.1038/ 2013;44:8–23.
s42003-020-01344-y Kitching RL. Biodiversity and taxonomy: impediment or opportunity.
Godfray HCJ. Challenges for taxonomy. Nature 2002;417:17–9. https:// In: Moritz C, Kikkawa J (eds), Conservation Biology in Australia and
doi.org/10.1038/417017a Oceania. Chipping Norton, Australia: Surrey Beatty, 1993, 253–68.
Grund R. The identification of Gahnia Forst & Forst. F. (Cyperaceae) Knapp S, Lamas G, Lughadha EN et al. Stability or stasis in the names
eating Hesperiidae (Lepidoptera) using immature stages. Victorian of organisms: the evolving codes of nomenclature. Philosophical
Entomologist 1998;28:20–32. Transactions of the Royal Society of London, Series B: Biological Sciences
Gu TT, Wu H, Yang F et al. Genomic analysis reveals a cryptic pangolin 2004;359:611–22. https://doi.org/10.1098/rstb.2003.1445
species. Proceedings of the National Academy of Sciences of the United Krell F-T. Suppressing works of contemporary authors using the Code’s
States of America 2023;120:1–12. publication requirements is neither easy nor advisable. Bulletin of
Guedes P, Alves-Martins F, Arribas JM et al. Eponyms have no place in Zoological Nomenclature 2021;78:61–7.
21st-century biological nomenclature. Nature Ecology and Evolution Krell F-T, Marshall SA. New species described from photographs:
2023;7:1157–60. https://doi.org/10.1038/s41559-023-02022-y Yes? No? Sometimes? A fierce debate and a new declaration of the
Halstead LB, Goriup PD, Middleton JA. The Loch Ness Monster. Nature ICZN. Insect Systematics and Diversity 2017;1:3–19. https://doi.
1976;259:75–6. https://doi.org/10.1038/259075c0 org/10.1093/isd/ixx004
Hamilton CA, Shockley FW, Simmons R et al. The future for a prominent Krell F-T, Pape T. Electronic publications need registration in ZooBank
taxonomy. Insect Systematics and Diversity 2021;5:1–2. to be available. Bulletin of Zoological Nomenclature 2015;72:245–51.
Hammer TA, Thiele KR. (119–122) Proposals to amend Articles 51 https://doi.org/10.21805/bzn.v72i3.a2
and 56 and Division III, to allow the rejection of culturally offen- Krell F-T, Wheeler QD. Specimen collection: plan for the future. Science
sive and inappropriate names. Taxon 2021;70:1392–4. https://doi. 2014;344:815–6.
org/10.1002/tax.12620 Lamas G. A critical review of J.Y. Miller’s checklist of the neotrop-
Hay RW, Houston TF, Williams AAE et al. Bring Back the Butterflies. ical Castniidae (Lepidoptera). Revista Peruana de Entomología
Butterfly Gardening for Western Australians. Perth: Western Australian 1995;37:73–87.
Museum, [1994], viii+72. Lamas G, Bollino M. Revisional notes on the ‘amastris’ group of Catasticta
ICZN. International Code of Zoological Nomenclature, 4th edn. London: Butler, 1870 (Lepidoptera: Pieridae), with descriptions of new spe-
The International Trust for Zoological Nomenclature, 1999. cies and subspecies. Zootaxa 2004;605:1–19.
ICZN. Amendment of Articles 8, 9, 10, 21 and 78 of the International Lamb J. Vanessa kiwa. Moths and Butterflies of New Zealand Trust
Code of Zoological Nomenclature to expand and refine methods of 2023;46:11–2.
publication. Bulletin of Zoological Nomenclature 2012;69:161–9. Lambert O, Bianucci G, Post K et al. The giant bite of a new raptorial sperm
Inomata T. On the original description of three Luehdorfia taxa in China whale from the Miocene epoch of Peru. Nature 2010a;466:105–8.
(Papilionidae). Tyô to Ga 1995;45:239–41. https://doi.org/10.1038/nature09067
Lambert O, Bianucci G, Post K et al. Corrigendum: The giant bite of a Smart P. The International Butterfly Book. London: Salamander Books,
new raptorial sperm whale from the Miocene epoch of Peru. Nature 1975, 275.
2010b;466:1134–1134. https://doi.org/10.1038/nature09381 Stradomsky BV. A molecular phylogeny of subfamily Polyommatinae
Lawton JH. Nessiteras Rhombopteryx. Oikos 1996;77:378–80. https:// (Lepidoptera: Lycaenidae). Caucasian Entomological Bulletin
doi.org/10.2307/3545927 2016;12:145–56. https://doi.org/10.23885/1814-3326-2016-12-1-
Lee C. Distribution and habitats of three Luehdorfia species in China. 145-156
Yadoriga 1982;107/108:38–40. Tautz D, Arctander P, Minelli A et al. A plea for DNA taxonomy. Trends
Löbl I, Klausnitzer B, Hartmann M et al. The silent extinction of species in Ecology and Evolution 2003;18:70–4. https://doi.org/10.1016/
and taxonomists—an appeal to science policymakers and legislators. s0169-5347(02)00041-1
Diversity 2023;15:1053. https://doi.org/10.3390/d15101053 Taxonomy Decadal Plan Working Group. Discovering Diversity: A Decadal

Mallet J, Willmott K. Taxonomy: renaissance or tower of babel? Trends Plan for Taxonomy and Biosystematics in Australia and New Zealand
in Ecology and Evolution 2003;18:57–9. https://doi.org/10.1016/ 2018–2028. Canberra and Wellington: Australian Academy of
s0169-5347(02)00061-7 Science and Royal Society Te Apārangi, 2018.
May RM. Taxonomy as destiny. Nature 1990;347:129–30. https://doi. Thiele KR, Conix S, Pyle RL et al. Towards a global list of accepted spe-
org/10.1038/347129a0 cies I. Why taxonomists sometimes disagree, and why this matters.
Mayr E. Principles of Systematic Zoology. New York: McGraw-Hill, 1969. Organisms Diversity and Evolution 2021;21:615–22. https://doi.
Mayr E, Ashlock PD. Principles of Systematic Zoology, 2nd edn. New York: org/10.1007/s13127-021-00495-y
McGraw-Hill, 1991. Thorpe SE. Is photography-based taxonomy really inadequate, un-
Miller JY. Castniidae. In: Heppner JB (ed.), Atlas of Neotropical Lepidoptera. necessary, and potentially harmful for biological sciences? A reply to
Checklist: Part 2. Hyblaeoidea-Pyraloidea-Tortricoidea. Gainesville: Ceríaco et al. (2016). Zootaxa 2017;4226:449–50.
Association for Tropical Lepidoptera/Scientific Publishers, 1995, Toussaint EFA, Braby MF, Müller CJ et al. Molecular phylogeny, system-
133–7, 176–7. atics and generic classification of the butterfly subfamily Trapezitinae
Neave SA. Nomenclator Zoologicus. A List of the Names of Genera and (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the
Subgenera in Zoology from the Tenth Edition of Linnaeus 1758 to the end Linnean Society 2022;195:1407–21.
of 1935. London: The Zoological Society of London, 1939–40. Vane-Wright RI. Taxonomy, methods of. In: Levin SA (ed.), Encyclopedia
Neave SA. Nomenclator Zoologicus. (Vol. 6 edited by M.A. Edwards & of Biodiversity, 2nd edn. Waltham, MA: Academic Press, 2013, 97–111.
A.T. Hopwood; Vol. 7 by M.A. Edwards & H.G. Vevers). London: The Vane-Wright RI, Whalley PES. Linnaeus’ fabulous butterfly. The Linnean
Zoological Society of London, 1950–75. 1985;1:19–24.
Ohl M. The Art of Naming. (Translated by Elisabeth Lauffer). Cambridge, Viloria AL, Pyrcz TW, Wojtusiak J et al. A brachypterous butterfly?
MA: The MIT Press, 2018, xv+294. Proceedings of the Royal Society of London (B) (Supplement) Biology
Okano K. Descriptons of four new butterflies of the genus Delias (Lep.: Letters 2003;270:21–4.
Pieridae), with some notes on Delias. Tokurana 1989:14:1–6. Wall F. A new krait from Oudh (Bungarus walli). Journal of the Bombay
Pape T. Species can be named from photos. Nature 2016;537:307. Natural History Society 1907;17:608–11.
https://doi.org/10.1038/537307b Wang Y-H, Hsiao Y-W, Lee K-H et al. Acoustic differentiation and be-
Peggie D, Vane-Wright RI, Yata O. An illustrated checklist of the pierid havioral response reveals cryptic species within Buergeria treefrogs
butterflies of northern and central Maluku (Indonesia). Butterflies (Anura, Rhacophoridae) from Taiwan. PLoS ONE 2017;12:1–23.
(Tokyo) 1995;11:23–47. Wang Y-H, Hsiao Y-W, Lee K-H et al. Correction: acoustic differen-
Polaszek A, Wilson EO. Sense and stability in animal names. Trends in tiation and behavioral response reveals cryptic species within
Ecology and Evolution 2005;20:421–2. https://doi.org/10.1016/j. Buergeria treefrogs (Anura, Rhacophoridae) from Taiwan. PLoS ONE
tree.2005.05.016 2018;13:e0193040. https://doi.org/10.1371/journal.pone.0193040
de Queiroz K. Species concepts and species delimitation. Systematic Biology Wang Y-H, Hsiao Y-W, Lee K-H et al. Renaming a new Buergeria spe-
2007;56:879–86. https://doi.org/10.1080/10635150701701083 cies from Taiwan—correction of a nomenclatural act not available to
Reissinger E. Zur Taxonomie und Systematik der Gattung Catasticta Butler ICZN. Nature Conservation Quaerterly 2020;111:4–13.
(Lepidoptera, Pieridae). Entomologische Zeitschrift 1972;82:97–124. Wheeler QD, Raven PH, Wilson EO. Taxonomy: impediment or expedient?
Rheindt FE, Bouchard P, Pyle RL et al. Tightening the requirements for Science 2004;303:285. https://doi.org/10.1126/science.303.5656.285
species diagnoses would help integrate DNA-based descriptions in Wiens JJ. Species delimitation: new approaches for discovering
taxonomic practice. PLoS Biology 2023;21:1–13. diversity. Systematic Biology 2007;56:875–8. https://doi.
Robbins RK, Lamas G. Nomenclatural changes in the neotrop- org/10.1080/10635150701748506
ical Eumaeini (Lepidoptera: Lycaenidae: Theclinae). Revista Will KW, Mishler BD, Wheeler QD. The perils of DNA barcoding and the
Brasileira de Zoologia 2002;19:197–214. https://doi.org/10.1590/ need for integrative taxonomy. Systematic Biology 2005;54:844–51.
s0101-81752002000500015 https://doi.org/10.1080/10635150500354878
Robbins RK, Nicolay S. Taxonomy and nomenclature of Strymon istapa Williams D, Wüster W, Fry BG. The good, the bad and the ugly: Australian
and S. columella (Lycaenidae: Theclinae: Eumaeini). Journal of the snake taxonomists and a history of the taxonomy of Australia’s ven-
Lepidopterists Society 1999;52:318–27. omous snakes. Toxicon 2006;48:919–30. https://doi.org/10.1016/j.
Rocha LA, Aleixo A, Allen G et al. Specimen collection: an essen- toxicon.2006.07.016
tial tool. Science 2014;344:814–5. https://doi.org/10.1126/ Wilson EO. Taxonomy as a fundamental discipline. Philosophical
science.344.6186.814 Transactions of the Royal Society of London, Series B: Biological Sciences
Rosenberg G, Krell F-T, Pyle R. Call to register new species in ZooBank. 2004;359:739. https://doi.org/10.1098/rstb.2003.1440
Nature 2012;491:40. https://doi.org/10.1038/491040b Winston JE. Describing Species. Practical Taxonomic Procedure for Biologists.
Saigusa T. A study on new subspecies of the tribe Theclini from eastern New York: Columbia University Press, 1999.
Asia (Lepidoptera, Lycaenidae). Zephyrus Researches 1993;1:12–22. Winston JE. Twenty-first century biological nomenclature—the enduring
Saito K, Seki Y. Four new taxa of Lycaenidae from South-East Asia power of names. Integrative and Comparative Biology 2018;58:1122–
(Lepidoptera). Butterflies (Tokyo) [2004];38:44–7. 31. https://doi.org/10.1093/icb/icy060
Schenk ET, McMasters JH. Procedure in Taxonomy (Revised Edition by A.M. Wojtusiak J. North Andean cloud forest and subparamo species of the
Keen and S.W. Muller). Stanford, CA: Standford University Press, 1948. genus Catasticta: C. chrysolopha Kollar C. spectrum Reissinger and C.
Scott P, Rines R. Naming the Loch Ness monster. Nature 1975;258:466– abdita new species. Lambillionea 1998;98:595–9.
8. https://doi.org/10.1038/258466a0 Wüster W, Kaiser H. Bungled Bungarus: lessons from a venomous snake
Šlapeta J. Ten simple rules for describing a new (parasite) spe- complex illustrate why taxonomic decisions belong in taxonomy-
cies. International Journal for Parasitology: Parasites and Wildlife competent journals. Zootaxa 2023;5297:139–43. https://doi.
2013;2:152–4. org/10.11646/zootaxa.5297.1.9
16 • Braby et al.
Wüster W, Bush B, Keogh JS et al. Taxonomic contributions in the ‘amateur’ Horagina. Transactions of the Lepidopterological Society of Japan
literature: comments on recent descriptions of new genera and species 2004;55:13–25.
by Raymond Hoser. Litteratura Serpentium 2001;21:67–79, 86–91. Yeates DK. Viva La Revolución! Designing the digital renaissance in zoo-
Wüster W, Thomson SA, O’Shea M et al. Confronting taxonomic van- logical taxonomy. Australian Journal of Entomology 2009;48:189–93.
dalism in biology: conscientious community self-organization can https://doi.org/10.1111/j.1440-6055.2009.00703.x
preserve nomenclatural stability. Biological Journal of the Linnean Yeates DK, Seago A, Nelson L et al. Integrative taxonomy, or iterative
Society 2021;133:645–70. taxonomy? Systematic Entomology 2011;36:209–17. https://doi.
Yagishita A, Nakano S, Morita S. An Illustrated List of the Genus Delias Hübner org/10.1111/j.1365-3113.2010.00558.x
of the World. Tokyo: Khepera Publishers, 1993, xiv+384, xiv+410+vi pls. Zhang Z-Q. A new era in zoological nomenclature and taxonomy: ICZN
Yago M. A new species of Horaga Moore (Lepidoptera, Lycaenidae, accepts e-publication and launches ZooBank. Zootaxa 2012;3450:8.

Lycaeninae, Theclini), with a key to the species of the subtribe https://doi.org/10.11646/zootaxa.3450.1.2

Braby Et Al 2024

Uploaded by

Copyright:

Available Formats

Braby Et Al 2024

Uploaded by

Document Information

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Braby Et Al 2024

Uploaded by

Copyright:

Available Formats

Zoological Journal of the Linnean Society, 2024, XX, 1–16

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

INTRODUCTION for scientific communication, establishing clear, universal, and

Received 24 October 2023; revised 4 February 2024; accepted 19 March 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Beaver et al. 2023).

Ogyris Angas, 1847

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

CONCLUSION phylogeny of the Hesperiidae. News Bulletin of the Entomological

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

Downloaded from https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlae043/7664331 by guest on 15 May 2024

You might also like