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How not to delimit taxa: A critique on a recently proposed “pragmatic classification” of jumping spiders (Arthropoda: Arachnida: Araneae: Salticidae)



Modern taxonomy and systematics profit from an invaluable tool that has been developed in the course of more than a century by intense discussions and negotiations of generations of zoologists and palaeontologists: The International Code of Zoological Nomenclature (ICZN 1999, 2012). The main goal of the Code is “to promote stability and universality in the scientific names of animals and to ensure that the name of each taxon is unique and distinct” (Melville 1995, ICZN 1999: 2). The provisions of the Code are generally accepted and thoroughly applied by the scientific community. Exceptions, such as the one described below, are very rare.
Accepted by G. Ruiz: 5 Dec. 2018; published: 18 Jan. 2019
ISSN 1175-5326 (print edition)
(online edition)
Copyright © 2019 Magnolia Press
Zootaxa 4545 (3): 444
How not to delimit taxa: a critique on a recently proposed pragmatic
classification” of jumping spiders (Arthropoda: Arachnida: Araneae: Salticidae)
Natural History Museum Bern, Switzerland
Institute of Ecology and Evolution, University of Bern, Switzerland
Hummeltal, Germany
Laboratório Especial de Coleções Zoológicas, Instituto Butantan, São Paulo, SP, Brazil
Department of Molecular Biology and Genetics, Democritus University of Thrace, Alexandroupolis, Greece
Department of Arachnology, Zentrum für Naturkunde, Universität Hamburg, Germany
Department of Zoology & Entomology, University of the Free State, Bloemfontein, South Africa
Department of Terrestrial Zoology, Western Australian Museum, Australia. Adjunct: School of Biological Sciences, University of
Western Australia, Australia
Arachnology, Senckenberg Research Institute, Frankfurt am Main, Germany
Institute for Biological Problems of the North, Magadan, Russia
Department of Zoology, National Museum of Nature and Science, Japan
Corresponding author. E-mail:
Modern taxonomy and systematics profit from an invaluable tool that has been developed in the course of more than a
century by intense discussions and negotiations of generations of zoologists and palaeontologists: The International Code
of Zoological Nomenclature (ICZN 1999, 2012). The main goal of the Code is “to promote stability and universality in
the scientific names of animals and to ensure that the name of each taxon is unique and distinct” (Melville 1995, ICZN
1999: 2). The provisions of the Code are generally accepted and thoroughly applied by the scientific community.
Exceptions, such as the one described below, are very rare.
The recent biodiversity crisis (e.g. Barnosky et al. 2011) and the insufficient number of taxonomists (e.g. Bacher
2012) lead to various approaches for improving the taxonomic procedure and for speeding up taxonomic progress
intended to describe the world’s species richness before large parts of it have become extinct, and to provide science-
based strategies against the loss of biodiversity. The usage of a variety of methods, including textual differential
diagnoses and descriptions, morphometrics, accurate illustrations, stacked photographs, SEM pictures and Micro-CT
representations of taxonomically relevant structures, as well as a wide range of molecular data, has become good practice
in taxonomy and systematics. As a consequence, modern taxon descriptions are much more comprehensive than in
earlier times. However, sometimes the amplification of various methods and techniques may cause complications rather
than enhancing good taxonomic practice. This may especially be true in cases where these tools are seen as exchangeable
rather than complementary, and newer approaches completely replace the more time-consuming textual descriptions or
pencil- and ink drawings (e.g., Coleman 2006; cf. Jäger 2016).
Recently, a new approach, termed “pragmatic classification” was suggested by Prószyński (2017a) in the taxonomy
of salticid spiders. Prószyński has an enormous reputation in this field, and Salticidae is the most species-rich family of
spiders, currently comprising more than 6100 extant and extinct species worldwide, representing ca. 13% of the global
spider diversity (World Spider Catalog 2018). They inhabit almost all types of land ecosystems and are therefore a target
group for describing, classifying and cataloguing the world’s terrestrial biodiversity. Prószyński considers word
descriptions and diagnoses of salticid spider taxa as mostly superfluous and even misleading. In a series of
publications on salticid taxonomy, he stated that the “traditional system of Salticidae (…) is insufficient to accommodate
hundreds of new taxa” (Prószyński 2016a: 4), and therefore, an “alternative classification of Salticidae” (Prószyński
2017a: 3) is needed. He based his approach on a large-scale “synthetic comparison of main diagnostic drawings”
(Prószyński 2016a: 5), and he proposed to select diagnostic characters of genera and species “by precise drawings of
palps, epigyne, spermathecae and ducts (…) and dismissal of translation of these drawings into words”. Because such a
Zootaxa 4545 (3) © 2019 Magnolia Press ·
set of drawings should not need further explanation, “dutifully made tedious descriptions of external appearances and
measurements of body parts (…) are almost useless” while, in contrast, “graphic definitions appear unequivocal”
(Prószyński 2017b: 37). He underlined his view several times as the “translation of appearance of characters into words
is too imprecise and often misleading” (Prószyński 2017a: 9), “the traditional way of documenting species by
descriptions with words and by routine measurements is particularly ineffective”, “lengthy description could be replaced
by color photographs of 3-4 key aspects of spider body appearance” (Prószyński 2018: 132−133), and so on. He further
claims that his system is provisional and intended purely for identification purposes. This system should exist in “parallel
to more theoretical system of affinities and phylogeny” (Prószyński 2017a: 4). Later (Prószyński 2018: 176), he extended
the usage of “pragmatic classification” to help “explain presumable relationships between biological entities, such as
species and genera”, while further stating that his classification “does not accept changes based on supposed gene
differences if they are not congruent with morphological premises” (Prószyński 2018: 177).
Based on his “pragmatic classification” approach, Prószyński created numerous new salticid genera in a series of
papers (e.g., Prószyński 2016a, 2018). This dynamic intervention into taxonomic practice has caused intense discussions
of the World Spider Catalog editorial team (most of them are co-authors of this contribution) with taxonomists from all
over the world, including Prószyński himself. Thus, we address some problems with “pragmatic classification” that seem
to us worthy of clarification.
(1) Disregard of ICZN rules. First of all, ICZN article 13.1.1 clearly states that, in order to be available, a new
name published after 1930 must “be accompanied by a description or definition that states in words characters that are
purported to differentiate the taxon” (emphasis by us; ICZN 1999, 2012). Therefore, a “pragmatic classification”, with its
disregard of textual descriptions and especially diagnoses, violates this article, or at least its meaning: There must be
evidence justifying a taxon, and this evidence must be explained. If it is not, this is an appeal to authority, not to
evidence. Furthermore, Prószyński’s (2017b: 37) claim that “graphic definitions appear unequivocal” is not supported by
facts — a glimpse on the different diagnostic figures for many European spider species in the database “Spiders of
Europe” (Nentwig et al. 2018) demonstrates the opposite. Good scientific drawings are always abstractions of reality,
underlining the relevant structures and neglecting the irrelevant ones. They do not only vary by their unique artistry, but
their interpretation may be subject to “the eye of the beholder”. Therefore, comparative illustrations (as important as they
are) cannot be regarded as a scientifically sound substitute for textual diagnoses.
(2) Disregard of the need to explain evidence. With its refusal to explain the evidence that should be found in
illustrations, Prószyński leaves it up to the reader to find this evidence by him- or herself. Therefore, the conclusions
expressed in a “pragmatic classification” are literally unjustified, and we suspect that in many cases they are likely false.
We do not know of any other field of science where simply presenting any evidence without explaining it, would be
accepted by the community.
(3) Descriptive taxonomy vs systematics (an old story). With “pragmatic classification”, an explicit classification
is created in parallel with the one that aims to reflect the taxa’s phylogeny. This reminds us of the long-lasting “Hennig vs
Mayr” dispute, but this discussion was already decided decades ago in favour of the former. There is only a single
objective base for a classification in systematics, and this is the unique process of phylogeny — thus, one could even
discuss a violation of the basic principle of objectivity in natural science by Prószyński’s “pragmatic classification”. The
creation of a parallel classification that disregards the information from molecular (and indeed, also morphological;
Maddison & Hedin 2003; Maddison 2015) phylogenies is even more bizarre, as “pragmatic classification” should also
shed light on the relationships of species and genera (Prószyński 2018). This procedure seems unacceptable, given the
important progress in salticid phylogeny in recent years, exhibited in a series of excellent contributions summarized by
Maddison (2015), which led to a comprehensive salticid classification, with placements for 96% of fossil and extant
genera. With the “pragmatic classification” approach, a severely retrograde scientific step has occurred, and confusion
and chaos in future salticid systematics seem to become an unavoidable outcome if such a system would be implemented
or widely accepted.
(4) Disregard of modern scientific methods. Prószyński (2017b) also rejects new taxonomic methods other than
molecular ones, in particular morphometrics (citations above). We cannot see any rational foundation for this, as modern
morphometrics are successfully applied in numerous taxonomic groups (Zelditch et al. 2012), especially in arthropods.
The enormous power of measurements and especially of ratios in connection with multivariate statistical methods has
been shown convincingly (Baur & Leuenberger 2011). Morphometrics may even provide a satisfying “loophole” in
diagnosing “cryptic” species by eye (e.g., Baur et al. 2014), that up till now could only be characterized by molecules (a
drawback that Prószyński himself laments).
(5) Suprageneric names ending with -INES. Prószyński (2017a: 9) created numerous suprageneric names ending
with “-INES” as “informal GROUPS OF GENERA” written entirely in capital letters to indicate that these are neither
subfamilial nor other names ruled by the Code. In spite of this, Prószyński (2017a) attributed to these informal names his
authorship with 2016 as year of publication, which does not correspond to the year of a valid publication concerning this
· Zootaxa 4545 (3) © 2019 Magnolia Press
matter, but rather to his webpage (Prószyński 2016b). Examples for such names are ASTIAINES Prószyński, 2016
(compare Astieae Simon, 1901), CHRYSILLINES Prószyński, 2016 (compare Chrysilleae Simon, 1901), PELLENINES
Prószyński, 2016 (compare Pelleninae Petrunkevitch, 1928), and so on. In contrast to his own publication (Prószyński
2017a), he uses the heading “subfamilies” at his webpage (Prószyński 2016b: Part I). It is obvious that the creation of
these names brings nothing but chaos in salticid systematics, and they should therefore be ignored by the community, or
at best, be considered as incorrect spelling of the respective available names. Furthermore, we do not understand how a
journal mostly dedicated to taxonomy and systematics (in this case “Ecologica Montenegrina”,
em) can accept this type of papers, claiming to have passed them through a peer-reviewing process. Many of the basic
errors highlighted above would have been pointed out by even a moderately competent reviewer familiar with basic
salticid systematics and taxonomic principles. In our view, this is nothing but scientific malpractice.
In order not to be misunderstood, we want to point out that we consider Prószyński’s lifework as truly exceptional
and feel a deep respect for it. But we must conclude that his “pragmatic classification” approach brings only limited
insights to salticid taxonomy and systematics. The editorial team members of the World Spider Catalog therefore already
did and will make accessible all taxonomic and nomenclatural information from the cited papers (this is indeed the
purpose of the Catalog; Nentwig et al. 2015; World Spider Catalog 2018), but we will reserve the right not to implement
all of the suggested changes for the structure of the Catalog to promote taxonomic stability in the Salticidae.
Bacher, S. (2012) Still not enough taxonomists. Trends in Ecology and Evolution, 27, 65–66.
Barnosky, A.D., Matzke, N., Tom iya , S., Wo gan , G.O.U., Swartz, B., Quental, T.B., Marshall, C., McGuire, J.L., Lindsey, E.L.,
Maguire, K.C., Mersey, B. & Ferrer, E.A. (2011) Has the earth’s sixth mass extinction already arrived? Nature, 471, 51–57.
Baur, H., Kranz-Baltensperger, Y., Cruaud, A., Rasplus, J.-Y., Timokhov, A.V. & Gokhman, V.E. (2014) Morphometric analysis and
taxonomic revision of Anisopteromalus Ruschka (Hymenoptera: Chalcidoidea: Pteromalidae) — an integrative approach.
Systematic Entomology, 39, 691–709.
Baur, H. & Leuenberger, C. (2011) Analysis of ratios in multivariate morphometry. Systematic Biology, 60, 813–825.
Coleman, C.O. (2006) Substituting time-consuming pencil drawings in arthropod taxonomy using stacks of digital photographs.
Zootaxa, 1360, 61–68.
ICZN [International Commission on Zoological Nomenclature] (1999) International Code of Zoological Nomenclature. Fourth
edition. International Trust for Zoological Nomenclature, London, 126 pp.
ICZN (2012) International Code of Zoological Nomenclature. 4
Edition. [Incorporating Declaration 44, amendments of Article
74.7.3, with effect from 31 December 1999 and the Amendment on e-publication, amendments to Articles 8, 9, 10, 21 and 78, with
effect from 1 January 2012]. Available from: (accessed 31 May 2018)
Jäger, P. (2016) A plea for taxonomic drawings and against pure photo-taxonomy. Indian Journal of Arachnology, 5 (1–2), 61–66.
Maddison, W.P. (2015) A phylogenetic classification of jumping spiders (Araneae: Salticidae). Journal of Arachnology, 43, 231–292.
Maddison, W.P. & Hedin, M.C. (2003) Jumping spider phylogeny (Araneae: Salticidae). Invertebrate Systematics, 17, 529–549.
Melville, R.V. (1995) Towards stability in the names of animals. A history of the International Commission on Zoological
Nomenclature 1895–1995. International Trust of Zoological Nomenclature, London, 92 pp.
Nentwig, W., Gloor, D. & Kropf, C. (2015) Spider taxonomists catch data on web. Nature, 528, 479.
Nentwig, W., Blick, T., Gloor, D., Hänggi, A. & Kropf, C. (2018) Spiders of Europe. Version 05.2018. Available from: http:// (accessed 31 May 2018)
Prószyński, J. (2016a) Delimitation and description of 19 new genera, a subgenus and a species of Salticidae (Araneae) of the world.
Ecologica Montenegrina, 7, 4–32.
Prószyński, J. (2016b) Monograph of Salticidae (Araneae) of the world 1995–2015. Part 1 & 2. Available from: http:// & (accessed 31 May 2018)
Prószyński, J. (2017a) Pragmatic classification of the world's Salticidae (Araneae). Ecologica Montenegrina, 12, 1–133.
Prószyński, J. (2017b) Revision of the genus Sitticus Simon, 1901 s. l. (Araneae: Salticidae). Ecologica Montenegrina, 10, 35–50.
Prószyński, J. (2018) Review of genera Evarcha and Nigorella, with comments on Emertonius, Padilothorax [sic], Stagetillus, and
description of five new genera and two new species (Araneae: Salticidae). Ecologica Montenegrina, 16, 130–179.
World Spider Catalog (2018) World Spider Catalog. Version 19.0. Natural History Museum Bern. Available from:
(accessed 31 May 2018)
Zelditch, M.L., Swiderski, D. & Sheets, H.D. (2012) Geometric Morphometrics for Biologists: A Primer. Second Edition. Elsevier
Academic Press, London, 488 pp.
... In 816 of the 1,021 papers (≈80 %) at least one new species was described. Conversely, the remaining 205 papers (≈20%) did not provide new species descriptions, but included, amongst others, papers publishing redescriptions of poorly known species and descriptions of unknown sexes (mainly papers in the Correspondence category), new country or regional records, national checklists of particular groups (e.g., Zonstein & Marusik 2013), catalogues, or discussed general issues in the taxonomy or biodiversity of arachnids (e.g., Platnick & Raven 2013;Kropf et al. 2019). ...
... The number of authors per araneological paper in Zootaxa ranged from one to thirteen. The article with the exceptionally high number of thirteen authors was a position paper on the delimitation of higher taxa in jumping spiders (Kropf et al. 2019). The highest number of co-authors in taxonomic papers describing new taxa (species and genera) was eight. ...
... In 816 of the 1,021 papers (≈80 %) at least one new species was described. Conversely, the remaining 205 papers (≈20%) did not provide new species descriptions, but included, amongst others, papers publishing redescriptions of poorly known species and descriptions of unknown sexes (mainly papers in the Correspondence category), new country or regional records, national checklists of particular groups (e.g., Zonstein & Marusik 2013), catalogues, or discussed general issues in the taxonomy or biodiversity of arachnids (e.g., Platnick & Raven 2013;Kropf et al. 2019). ...
... The number of authors per araneological paper in Zootaxa ranged from one to thirteen. The article with the exceptionally high number of thirteen authors was a position paper on the delimitation of higher taxa in jumping spiders (Kropf et al. 2019). The highest number of co-authors in taxonomic papers describing new taxa (species and genera) was eight. ...
Full-text available
Zootaxa published more than a thousand papers on Araneae from 2002 to the present, including descriptions of 3,833 new spider species and 177 new genera. Here we summarise the key contributions of Zootaxa to our current knowledge of global spider diversity. We provide a historical account of the researchers that have actively participated as editors, and recognize the more than 1,000 reviewers without whom none of this would have been possible. We conduct a simple analysis of the contributions by authors and geographic region, which allows us to uncover some of the underlying trends in current spider taxonomy. In addition, we examine some of the milestones in twenty years of spider systematic research in Zootaxa. Finally, we discuss future prospects of spider taxonomy and the role that Zootaxa and its younger sister journal Megataxa will play in it. We would like to dedicate this contribution to the memory of Norman I. Platnick, a crucial figure in the advancement of spider systematics.
... Supplemental data for this article can be accessed here. recent splitting of Evarcha into five genera was not supported phylogenetically, and it is not accepted (Blick and Marusik 2018;Kropf et al. 2019;World Spider Catalog 2020). ...
... While our results agree with Prószyński (2018) that Evarcha should be split, the division supported by our analysis is different from what he chose, as Colopsus does not fall with either Evacin or Evalba or with Evarcha s. str. Several arguments have been presented against splitting of Evarcha (Blick & Marusik, 2018;Kropf et al. 2019). Counterarguments have also been put forward (Breitling 2019). ...
The Sri Lankan jumping spider species first described as Colopsus cancellatus and its close relatives are an understudied yet charismatic part of the fauna of the island. Here, using molecular sequence data (cytochrome c oxidase subunit I, 18S rRNA, 28S rRNA, and histone H3), and a variety of methods we tested the validity of this genus and recover three well-supported clades in the ML analysis: Colopsus and Evarcha in two separate clades with Burmattus in-between. In the MP analysis Colopsus and Evarcha form two separate clades. The Sri Lanka Colopsus previously misplaced in Evarcha formed a distinct clade in both the ML and MP topologies. Thus, Colopsus is restored as a distinct genus. Pancorius is recovered as non-monophyletic. We additionally describe the following new species: Colopsus cinereus sp. nov., C. ferruginus sp. nov., C. magnus sp. nov. and C. tenuesi sp. nov., Evarcha latus sp. nov., Pancorius alboclypeus sp. nov., P. altus sp. nov and P. athukoralai sp. nov. Pancorius is recorded for the first time from Sri Lanka.
... 18S rDNA sequences are not characters, but suites of characters-the characters are individual nucleotide positions. In a similar fashion, a morphological diagram cannot alone be used to describe a new species, even if it encompasses all the key features (as an example, see [4]). Rule 13.1.1 ...
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Recently Greay et al. (Parasit Vectors 11:197, 2018) described several new Apicomplexa parasites from domestic companion animals in Australia. Harris (Parasit Vectors 12;172, 2019) highlighted that these descriptions did not conform to the International Code of Zoological Nomenclature (ICZN) rules. Despite Harris (2019) clearly noting “molecular characters can be used to satisfy article 13.1.1 of the code”, in a reply Greay et al. (Parasit Vectors 12:178, 2019) incorrectly state “Harris considers the eight new species…invalid on the basis that only molecular characters were provided”. This was not the case. The ICZN has strict rules regarding species descriptions for good reasons. Here I reiterate why the forms described by Greay et al. (2018) are not valid.
... Some taxa, such as spiders and butterflies, have their own world catalogs in which the most important available taxonomic information is compiled. However, these databases are meant to track the current situation; hence, it is beyond their scope to regulate the validity of descriptions or names, although the editorial team may sporadically decide to exclude taxonomic information (Kropf et al. 2019). Transferring the responsibility of policing the legality of preserved specimens to scientific societies is neither feasible nor desirable, although scientific societies certainly should participate by having an explicit code of ethics and strongly encouraging their affiliates to publish only in peer-reviewed venues. ...
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Illegal transfer of wildlife has 2 main purposes: trade and scientific research. Trade is the most common, whereas scientific research is much less common and unprofitable, yet still important. Biopiracy in science is often neglected despite that many researchers encounter it during their careers. The use of illegally acquired specimens is detected in different research fields, from scientists bioprospecting for new pharmacological substances, to taxonomists working on natural history collections, to researchers working in zoos, aquariums, and botanical gardens. The practice can be due to a lack of knowledge about the permit requirements in different countries or, probably most often, to the generally high level of bureaucracy associated with rule compliance. Significant regulatory filters to avoid biopiracy can be provided by different stakeholders. Natural history collection hosts should adopt strict codes of conduct; editors of scientific publications should require authors to declare that all studied specimens were acquired legally and to cite museum catalog numbers as guarantee of best practices. Scientific societies should actively encourage publication in peer-reviewed journals of work in which specimens collected from the wild were used. The International Commission on Zoological Nomenclature could require newly designated types based on recently collected specimens to be accompanied by statements of deposition in recognized scientific or educational institutions. We also propose the creation of an online platform that gathers information about environmental regulations and permits required for scientific activities in different countries and respective responsible governmental agencies and the simplification of the bureaucracy related to regulating scientific activities. This would make regulations more agile and easier to comply with. The global biodiversity crisis means data need to be collected ever faster, but biopiracy is not the answer and undermines the credibility of science and researchers. It is critical to find a modus vivendi that promotes compliance with regulations and scientific progress. Article impact statement: Although poorly debated, illegal collection of wildlife for science is a situation faced by scientists working from bioprospecting to taxonomy and natural history. This article is protected by copyright. All rights reserved.
... I was resuscitated by a single, righteous man, a person whom I never know before -Rainer Breitling. He has asked me whether I read Kropf et al (2019) paper -I did not. So he has sent me PDF of it, and let me read his article written out of compassion and in my defense, which later, after biased delaying tactics of Zootaxa, had to be published in other journal and has finally appeared on May 7th, 2019 in Ecologica Montenegrina 21: 62-69. ...
The paper comments discussion on how to delimit taxa of Salticidae, initiated by Kropf et al. (2019). It discusses a number of important general issues, including the relative value of precise diagnostic drawings and photos compared to verbal definitions of taxa, which is relevant to hundreds of current taxonomic publications. To illustrate key principles, the paper also analyses the validity of the recent synonymization of the genera Junxattus, Lechia and Orcevia, evaluating the sufficiency and relevance of previously published morphological and molecular data.On the basis of the above (documented on Figs 1-4 below), the following momenclatural changes are introduced: Gen. Junxattus Prószyński & Deeleman-Reinhold, 2012 - revalidated - (type species Junxattus daiqini); Gen. Lechia Zabka, 1985 - revalidated - (type species Lechia squamata); Gen. Orcevia Thorell, 1890 - revalidated - (type species Orcevia keyserlingii).The placement of female of Laufeia concava and male (only!) of Laufeia squamata are erroneous and require generic transfer. Lechia minuta (Prószyński, 1992) is misplaced too, belonging neither to Laufeia nor to Euoprys, it should be provisionally listed as Lechia (according to latest documented reference) until revisionary research of fauna of its area yields additional material to clarify its correct placement.
... was recently split into a number of separate genera (Prószyński 2016(Prószyński , 2017, following an earlier subdivision of the European members into several subgenera by Lohmander (1944). This action was part of a larger series of revisions that has recently come under severe criticism (Kropf et al. 2019). The question to ask in the context of the barcode analysis is obviously whether this extensive splitting of Sitticus is justified. ...
DNA barcode sequencing has rapidly become one of the most powerful tools for biodiversity assessments. Beyond its original uses for the identification of animal species, including the discovery of cryptic diversity in difficult taxonomic groups, the growing public sequence datasets also offer opportunities for more wide-ranging applications. This contribution shows how barcode data can provide useful complementary information to assist taxonomic decision making at the genus level. An analysis of public barcode datasets for 10 diverse spider families, covering more than 3400 species and morphospecies, reveals numerous examples where sequence similarities either strongly support or convincingly refute recent controversial genus assignments. The following nomenclatorial changes are suggested based on a combined assessment of morphological evidence and the barcode analysis: Acantholycosa = Pardosa (syn. nov.); Piratula = Pirata (syn. nov.); Pulchellodromus, Philodromimus, Tibellomimus, Artanes, and Emargidromus = subgenera of Philodromus (stat. nov.); Cryptachaea riparia = Parasteatoda riparia (comb. nov.); Ohlertidion = Heterotheridion (syn. nov.); Saaristoa = Aphileta (syn. nov.); Aphileta microtarsa = Eulaira microtarsa (comb. conf.);
... A large number of anapophysate species, as well as many species inquirendae, often known only from female or juvenile specimens, typically the type material, remain incertae sedis within Xysticus, pending additional analysis. One could question whether the new combinations proposed here are a potential threat to nomenclatural stability (analogous to the case discussed by Kropf et al. 2019) and whether any major rearrangement would better be published as part of a -final‖ comprehensive revision of the group. Obviously, the present work does build on and complement several major revisions, which have already proposed a large number of new combinations without any adverse effect on nomenclatural practice. ...
The phylogenetic relationships and taxonomy of the crab spider genus Xysticus and its closest relatives (i.e., the tribe Coriarachnini, also including, e.g., Ozyptila, Coriarachne and Bassaniana) have long been controversial, with several alternative classifications being proposed, none of which has gained universal acceptance. As Coriarachnini is largely confined to the Holarctic region, the main target area of recent DNA barcoding projects for spiders, a large amount of genetic data for the group is now publicly available. The results of a phylogenetic analysis of this sequence dataset are largely congruent with earlier morphology-based results regarding the evolutionary structure of the group. In particular, they highlight the fact that Xysticus s. lat. is a paraphyletic assembly and that several species groups need to be placed in separate genera to achieve monophyly of Xysticus s. str. Similarly, Coriarachne and Bassaniana appear as independent clades rather than a joined monophyletic Coriarachne s. lat. In contrast, further subdivision of Ozyptila is not supported by the genetic data. Importantly, the analysis also shows that anapophysate members of Xysticus s. lat. form two widely separated groups: a primarily anapophysate division, also including Coriarachne and Bassaniana, at the base of Xysticus s. lat., and a secondarily anapophysate clade deeply nested within Xysticus s. str. This might explain some of the earlier difficulties when trying to define generally accepted subgroups within Xysticus s. lat. The phylogenetic scaffold based on barcode sequences is sufficiently dense and well resolved to attempt the tentative and provisional placement of the majority of species in Xysticus s. lat. in the independent genera Xysticus s. str., Bassaniodes, Psammitis and Spiracme as a starting point for a future more formal revision of the group.
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The systematics of sitticine jumping spiders is reviewed, with a focus on the Palearctic and Nearctic regions, in order to revise their generic classification, clarify the species of one region (Canada), and study their chromosomes. A genome-wide molecular phylogeny of 23 sitticine species, using more than 700 loci from the arachnid Ultra-Conserved Element (UCE) probeset, confirms the Neotropical origins of sitticines, whose basal divergence separates the new subtribe Aillutticina (a group of five Neotropical genera) from the subtribe Sitticina (five genera of Eurasia and the Americas). The phylogeny shows that most Eurasian sitticines form a relatively recent and rapid radiation, which we unite into the genus Attulus Simon, 1868, consisting of the subgenera Sitticus Simon, 1901 (seven described species), Attulus (41 described species), and Sittilong Prószyński, 2017 (one species). Five species of Attulus occur natively in North America, presumably through dispersals back from the Eurasian radiation, but an additional three species were more recently introduced from Eurasia. Attus palustris Peckham & Peckham, 1883 is considered to be a full synonym of Euophrys floricola C. L. Koch, 1837 (not a distinct subspecies). Attus sylvestris Emerton, 1891 is removed from synonymy and recognized as a senior synonym of Sitticus magnus Chamberlin & Ivie, 1944. Thus, the five native Attulus in North America are Attulus floricola , A. sylvestris , A. cutleri , A. striatus , and A. finschi . The other sitticines of Canada and the U.S.A. are placed in separate genera, all of which arose from a Neotropical radiation including Jollas Simon, 1901 and Tomis F.O.Pickard-Cambridge, 1901: (1) Attinella Banks, 1905 ( A. dorsata , A. concolor , A. juniperi ), (2) Tomis ( T. welchi ), and (3) Sittisax Prószyński, 2017 ( S. ranieri ). All Neotropical and Caribbean “ Sitticus ” are transferred to either Jollas (12 species total) or Tomis (14 species). Attinella (three species) and Tomis are both removed from synonymy with Sitticus ; the synonymy of Sitticus cabellensis Prószyński, 1971 with Pseudattulus kratochvili Caporiacco, 1947 is restored; Pseudattulus Caporiacco, 1947 is synonymized with Tomis . Six generic names are newly synonymized with Attulus and one with Attinella . Two Neotropical species are described as new, Jollas cupreussp. nov. and Tomis manabitasp. nov. Forty-six new combinations are established and three are restored. Three species synonymies are restored, one is new, and two are rejected. Across this diversity of species is a striking diversification of chromosome complements, with X-autosome fusions occurring at least four times to produce neo-Y sex chromosome systems (X 1 X 2 Y and X 1 X 2 X 3 Y), some of which ( Sittisax ranieri and S. saxicola ) are sufficiently derived as to no longer preserve the simple traces of ancestral X material. The correlated distribution of neo-Y and a base autosome number of 28 suggests that neo-Y origins occurred preferentially in lineages with the presence of an extra pair of autosomes.
The paper gives a counterpoint to the recent critique of the “pragmatic classification” of jumping spiders (Arthropoda: Arachnida: Araneae: Salticidae).
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We use an integrative taxonomic approach to revise the genus Anisopteromalus. In particular, we apply multivariate ratio analysis (MRA), a rather new statistical method based on principal component analysis (PCA) and linear discriminant analysis (LDA), to numerous body measurements and combine the data with those from our molecular analysis of Cytb and ITS2 genetic markers (on a subset of species) and all available published data on morphology, karyology, behaviour, host associations and geographic distribution. We demonstrate that the analysis of quantitative characters using MRA plays a major role for the integration of name-bearing types and thus for the association of taxa with names. Six species are recognized, of which two are new: A. cornis Baur sp.n. and A. quinarius Gokhman & Baur sp.n. For Anisopteromalus calandrae (Howard), a well-known, cosmopolitan parasitoid of stored-product pests, we have selected a neotype to foster continuity and stability in the application of this important name. The species was sometimes confused with the related A. quinarius sp.n., another cosmopolitan species that is frequently encountered in similar environments. We also show that several species originally described or later put under Anisopteromalus actually belong to different genera: Cyrtoptyx camerunus (Risbec) comb.n.; Meraporus glaber (Szelényi) comb.n.; Dinarmus schwenkei (Roomi, Khan & Khan) comb.n. Neocatolaccus indicus Ayyar & Mani is confirmed as a junior synonym of Oxysychus sphenopterae (Ferrière) syn.n. and Anisopteromalus calandrae brasiliensis (Domenichini) stat.rev. must be considered as a valid but doubtful taxon. This published work has been registered in ZooBank,
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The most time-consuming part of a taxonomic description is making the illustrations. This contribution shows how to save time by omitting the pencil drawings of arthropod appendages and replacing them by stacks of microphotographs. These are imported into a drawing software package on a computer in order to make a publication-ready line drawing, a technique described in detail in Coleman (2003). The photographic method requires a special treatment of the appendages which is also shown.
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The analysis of ratios of body measurements is deeply ingrained in the taxonomic literature. Whether for plants or animals, certain ratios are commonly indicated in identification keys, diagnoses, and descriptions. They often provide the only means for separation of cryptic species that mostly lack distinguishing qualitative characters. Additionally, they provide an obvious way to study differences in body proportions, as ratios reflect geometric shape differences. However, when it comes to multivariate analysis of body measurements, for instance, with linear discriminant analysis (LDA) or principal component analysis (PCA), interpretation using body ratios is difficult. Both techniques are commonly applied for separating similar taxa or for exploring the structure of variation, respectively, and require standardized raw or log-transformed variables as input. Here, we develop statistical procedures for the analysis of body ratios in a consistent multivariate statistical framework. In particular, we present algorithms adapted to LDA and PCA that allow the interpretation of numerical results in terms of body proportions. We first introduce a method called the "LDA ratio extractor," which reveals the best ratios for separation of two or more groups with the help of discriminant analysis. We also provide measures for deciding how much of the total differences between individuals or groups of individuals is due to size and how much is due to shape. The second method, a graphical tool called the "PCA ratio spectrum," aims at the interpretation of principal components in terms of body ratios. Based on a similar idea, the "allometry ratio spectrum" is developed which can be used for studying the allometric behavior of ratios. Because size can be defined in different ways, we discuss several concepts of size. Central to this discussion is Jolicoeur's multivariate generalization of the allometry equation, a concept that was derived only with a heuristic argument. Here we present a statistical derivation of the allometric size vector using the method of least squares. The application of the above methods is extensively demonstrated using published data sets from parasitic wasps and rock crabs.
This paper, dedicated to search for identification methods of genera of Salticidae (Araneae), presents prototype of a “Handbook of Jumping Spiders Identification”, based on morphology of palps, spermathecae and ducts, as well as some other easily noticeable characters. It includes diagnostic drawings of representative species of each genus, additional survey of diversity of these characters in 4800 recognizable species is available instantly, by hyperlinks provided to parallel Internet "Monograph of the Salticidae (Araneae) of the World 1995-2016".Part I "Introduction to alternative classification of Salticidae" by Prószyński (2016a), accessible at: [too large to be published whole as a PDF]. The work contains methodological suggestions on how the proposed system could be improved and further developed. Partial revision of the present taxonomic system of Salticidae is included.The paper provides diagnoses and diagnostic drawings to genera of Salticidae, grouped to facilitate identification into morphologically coherent, informal groups of genera. There are following provisional groups proposed: AEURILLINES, AMYCINES, AMYCOIDA VARIA, ASTIAINES, BELIPPINES, CHRYSILLINES, COCALODINES, COLONINES [= former Thiodininae], DENDRYPHANTINES, DIOLENINES, EUODENINES, EUOPHRYINES, EUPOAINES, EVARCHINES, HABRONATTINES, HARMOCHIRINES, HELIOPHANINES, HISPONINES, HYLLINES, ICIINES, LAPSIINES, LIGONIPEINES, LYSSOMANINES, MENEMERINES, MYRMARACHNINES, NOTICIINES, PELLENINES, PSEUDICIINES, SIMAETHINES, SITTICINES, SPARTAEINES, THIRATOSCIRTINAE, YAGINUMAELLINES, YLLENINES. There is also temporary UNCLASSIFIED group and display of exemplary FOSSILS. The proposals of grouping and delimitation have working character, pending further research and tests.The following synonyms and combinations (new, corrected or reinstated) are listed in the paper together with their documentation and/or discussions. They have been accumulated during 22 years of work on database, but are printed for the first time only now (location of their documentation in the text below can be quickly found using computer searching facility).Aelurillus stanislawi (Prószyński, 1999) (male from Israel) = Rafalus stanislawi Prószyński, 1999, Aelurillus stanislawi Azarkina, (2006) (nec Prószyński, 1999) = Aelurillus minutus Azarkina, 2002, Amphidraus manni (Bryant 1943) = Nebridia manni Bryant 1943, Amphidraus mendica (Bryant 1943) = Nebridia mendica Bryant 1943, Amphidraus semicanus (Simon, 1902) = Nebridia semicana Simon, 1902, Bianor incitatus Thorell, 1890 (in part) = Stichius albomaculatus Thorell, 1890, Bryantella smaragdus (Crane, 1945) = Bryantella smaragda (Crane, 1945), Chinattus undulatus (Song & Chai, 1992) (in part, male) = Chinattus szechwanensis (Prószyński, 1992), Colyttus kerinci (Prószyński & Deeleman-Reinhold, 2012) = Donoessus kerinci Prószyński & Deeleman-Reinhold, 2012, Colyttus nigriceps (Simon, 1899) = Donoessus nigriceps (Simon, 1899), Colyttus striatus (Simon, 1902) = Donoessus striatus (Simon, 1902), Cytaea severa (Thorell, 1881) (in part) = Cytaea alburna Keyserling, 1882, Euophrys minuta Prószynski, 1992 ) = Lechia minuta (Prószynski, 1992 ), Laufeia daiqini (Prószyński & Deeleman-Reinhold, 2012) = Junxattus daiqini Prószyński & Deeleman-Reinhold, 2012, Laufeia kuloni (Prószynski & Deeleman-Reinhold, 2012) = Orcevia kuloni Prószynski & Deeleman-Reinhold 2012, Laufeia keyserlingi (Thorell, 1890) = Orcevia keyserlingi (Thorell, 1890), Laufeia eucola (Thorell, 1890) = Orcevia eucola (Thorell, 1890), Laufeia perakensis (Simon, 1901) = Orcevia perakensis (Simon, 1901), Laufeia proszynskii Song, Gu & Chen, 1988 = Orcevia proszynskii (Song, Gu & Chen, 1988), Laufeia squamata ( Żabka, 1985 ) = Lechia squamata Żabka, 1985, Maevia C. L. Koch, 1846 (in part) = Paramaevia Barnes, 1955, Maevia hobbsae Barnes, 1958 = Paramaevia hobbsae Barnes, 1958, Maevia michelsoni Barnes, 1958 = Paramaevia michelsoni (Barnes, 1958), Maevia poultoni Peckham & Peckham, 1909 = Paramaevia poultoni (Peckham & Peckham, 1901),Maratus anomaliformis (Żabka, 1987) = "Lycidas" anomaliformis Żabka, 1987, Metaphidippus felix (Peckham & Peckham, 1901) = Messua felix (Peckham & Peckham, 1901), Monomotapa principalis Wesolowska, 2000 = Iranattus principalis (Wesolowska, 2000), Myrmarachne exasperans (Peckham & Peckham, 1892) = Emertonius exasperans Peckham & Peckham, 1892, Myrmarachne melanocephala MacLeay, 1839 (in part) = Myrmarachne ramosa Badcock, 1918, Myrmarachne melanocephala MacLeay, 1839 (in part) = Myrmarachne contracta (Karsch, 1880), Myrmarachne melanocephala MacLeay, 1839 (in part) = Myrmarachne albicrurata Badcock, 1918, Myrmarachne melanocephala MacLeay, 1839 (in part) = Myrmarachne lateralis Badcock, 1918, Myrmarachne melanocephala MacLeay, 1839 (in part) = Myrmarachne providens Simon, 1901, Myrmavola globosa (Wanless, 1978) = Toxeus globosus (Wanless, 1978) (self-correction), Omoedus albertisi (Thorell, 1881) = Zenodorus albertisi (Thorell, 1881), Omoedus arcipluvii (Peckham, Peckham, 1901) = Zenodorus arcipluvii (Peckham, Peckham, 1901), Omoedus asper (Karsch, 1878) = Ascyltus asper (Karsch, 1878), Omoedus bernsteini (Thorell, 1881) = Zenodorus bernsteini (Thorell, 1881), - Omoedus brevis Zhang J., Maddison, 2012 = Zenodorus brevis (Zhang J., Maddison, 2012), Omoedus cyanothorax (Thorell, 1881) = Pystira cyanothorax (Thorell, 1881), - Omoedus durvillei (Walckenaer, 1837) = Zenodorus durvillei (Walckenaer, 1837)- Omoedus danae (Hogg, 1915) = Zenodorus danae Hogg, 1915, - Omoedus darleyorum Zhang J., Maddison, 2012 = Zenodorus darleyorum (Zhang J., Maddison, 2012),Omoedus ephippigerus (Simon, 1885) = Pystira ephippigera (Simon, 1885), Omoedus karschi (Thorell, 1881) = Pystira karschi (Thorell, 1881), Omoedus lepidus (Guerin, 1834) = Zenodorus lepidus (Guerin, 1834), Omoedus metallescens (Koch L., 1879) = Zenodorus metallescens (Koch L., 1879), Omoedus meyeri Zhang J., Maddison, 2012 = Zenodorus meyeri (Zhang J., Maddison, 2012), Omoedus microphthalmus (Koch L., 1881) = Zenodorus microphthalmus (Koch L., 1881), Omoedus nigripalpis (Thorell, 1877) = Pystira nigripalpis (Thorell, 1877)]. Omoedus obscurofemoratus (Keyserling, 1883) = Zenodorus obscurofemoratus (Keyserling, 1883), Omoedus omundseni Zhang J., Maddison, 2012 = Zenodorus omundseni (Zhang J., Maddison, 2012), Omoedus orbiculatus (Keyserling, 1881) = Zenodorus orbiculatus (Keyserling, 1881), Omoedus papuanus Zhang J., Maddison, 2012 = Zenodorus papuanus (Zhang J., Maddison, 2012), Omoedus ponapensis (Berry, Beatty, Prószyński, 1996) = Zenodorus ponapensis Berry, Beatty, Prószynski, 1996, Omoedus semirasus (Keyserling, 1882) = Zenodorus semirasus (Keyserling, 1882), Omoedus swiftorum Zhang J., Maddison, 2012 = Zenodorus swiftorum (Zhang J., Maddison, 2012), Omoedus tortuosus Zhang J., Maddison, 2012 = Zenodorus tortuosus (Zhang J., Maddison, 2012), Omoedus versicolor (Dyal, 1935) = Pystira versicolor Dyal, 1935, [Unrecognizable species of Zenodorus: Omoedus jucundus (Rainbow, 1912) = Zenodorus jucundus (Rainbow, 1912), Omoedus juliae (Thorell, 1881) = Zenodorus juliae (Thorell, 1881), Omoedus marginatus (Simon, 1902) = Zenodorus marginatus (Simon, 1902), Omoedus niger (Karsch, 1878) = Zenodorus niger (Karsch, 1878), - Omoedus pupulus (Thorell, 1881) = Zenodorus pupulus (Thorell, 1881), - Omoedus pusillus (Strand, 1913) = Zenodorus pusillus (Strand, 1913), Omoedus rhodopae (Hogg, 1915) = Zenodorus rhodopae (Hogg, 1915), Omoedus syrinx (Hogg, 1915) = Zenodorus syrinx Hogg, 1915, Omoedus variatus (Pocock, 1899) = Zenodorus variatus (Pocock, 1899), Omoedus varicans (Thorell, 1881) = Zenodorus varicans Thorell, 1881, Omoedus wangillus (Strand, 1911) = Zenodorus wangillus Strand, 1911], Pellenes ostrinus (Simon, 1884) (in part) = Pellenes diagonalis Simon, 1868, Pseudicius alter Wesolowska, 1999 = Afraflacilla altera (Wesolowska, 1999), Pseudicius arabicus (Wesolowska, van Harten, 1994) = Afraflacilla arabica Wesolowska, van Harten, 1994, Pseudicius bipunctatus Peckham, Peckham, 1903 = Afraflacilla bipunctata (Peckham, Peckham, 1903), Pseudicius braunsi Peckham, Peckham, 1903 = Afraflacilla braunsi (Peckham, Peckham, 1903), Pseudicius datuntatus Logunov, Zamanpoore, 2005= Afraflacilla datuntata (Logunov, Zamanpoore, 2005), Pseudicius elegans (Wesolowska, Cumming, 2008) = Afraflacilla elegans (Wesolowska, Cumming, 2008), Pseudicius eximius Wesolowska, Russel-Smith, 2000 = Afraflacilla eximia (Wesolowska, Russel-Smith, 2000), Pseudicius fayda Wesolowska, van Harten, 2010 = Afraflacilla fayda (Wesolowska, van Harten, 2010), Pseudicius flavipes Caporiacco, 1935 = Afraflacilla flavipes (Caporiacco, 1935), Pseudicius histrionicus Simon, 1902 = Afraflacilla histrionica (Simon, 1902), Pseudicius imitator Wesolowska, Haddad, 2013 = Afraflacilla imitator (Wesolowska, Haddad, 2013), Pseudicius javanicus Prószynski, Deeleman-Reinhold, 2012 = Afraflacilla javanica (Prószynski, Deeleman-Reinhold, 2012), Pseudicius karinae (Haddad, Wesolowska, 2011) = Afraflacilla karinae (Haddad, Wesolowska, 2011), Pseudicius kraussi Marples, 1964 = Afraflacilla kraussi (Marples, 1964), Pseudicius mikhailovi Prószynski, 1999 = Afraflacilla mikhailovi (Prószynski, 1999), Pseudicius mushrif Wesolowska, van Harten, 2010 = Afraflacilla mushrif (Wesolowska, van Harten, 2010), Pseudicius philippinensis Prószynski, 1992 = Afraflacilla philippinensis (Prószynski, 1992), Pseudicius punctatus Marples, 1957 = Afraflacilla punctata (Marples, 1957), Pseudicius refulgens Wesolowska, Cumming, 2008 = Afraflacilla refulgens (Wesolowska, Cumming, 2008), Pseudicius reiskindi Prószynski, 1992 = Afraflacilla reiskindi (Prószynski, 1992), Pseudicius roberti Wesolowska, 2011 = Afraflacilla roberti (Wesolowska, 2011), Pseudicius spiniger (Pickard-Cambridge O., 1872) = Afraflacilla spiniger (Pickard-Cambridge O., 1872), Pseudicius tamaricis Simon, 1885 = Afraflacilla tamaricis (Simon, 1885), Pseudicius tripunctatus Prószynski, 1989 = Afraflacilla tripunctata (Prószynski, 1989), Pseudicius venustulus Wesolowska, Haddad, 2009 = Afraflacilla venustula (Wesolowska, Haddad, 2009), Pseudicius wadis Prószynski, 1989 = Afraflacilla wadis (Prószynski, 1989), Pseudicius zuluensis Haddad, Wesolowska, 2013 = Afraflacilla zuluensis (Haddad, Wesolowska, 2013), Servaea incana (Karsch, 1878) (in part) = Servaea vestita ( L. Koch, 1879), Sidusa extensa (Peckham & Peckham, 1896) = Cobanus extensus (Peckham & Peckham, 1896), Sidusa Peckham & Peckham, 1895 (in part) = Cobanus F. O. Pickard-Cambridge , 1900, Sidusa Peckham & Peckham, 1895 (in part) = Wallaba Mello-Leitão, 1940, Stagetillus elegans (Reimoser, 1927) = "Padillothorax" elegans Reimoser, 1927, Stagetillus taprobanicus (Simon, 1902) = "Padillothorax" taprobanicus Simon, 1902, Telamonia besanconi (Berland & Millot, 1941) = Brancus besanconi (Berland & Millot, 1941), Telamonia fuscimana (Simon, 1903) = Brancus fuscimanus (Simon, 1903), Telamonia longiuscula (Thorell, 1899) = Hyllus longiusculus (Thorell, 1899), Telamonia thoracica (Thorell, 1899) [="Viciria"thoracica: Prószyński, 1984 = Hyllus thoracicus (Thorell, 1899), - Thiania sundevalli (Thorell, 1890) = Nicylla sundevalli Thorell, 1890, Thiania spectrum (Simon, 1903) = Thianitara spectrum Simon, 1903, Thiania thailandica (Prószyński & Deeleman-Reinhold, 2012) = Thianitara thailandica Prószyński & Deeleman-Reinhold, 2012, Viciria albocincta Thorell, 1899 = Hyllus albocinctus (Thorell, 1899), Yaginumaella striatipes (Grube, 1861) (in part) = Yaginumaella ususudi Yaginuma, 1972.
The genus Hasarius Simon, 1871, is revised following methodology recommended by "Pragmatic classification" of Prószynski (2017). Structure of the genus is insufficiently known, containing at present single cosmopolite species and a few of uncertain congeners. The paper introduces the following nomenclatorical corrections: Hasarius adansoni: Jastrzebski, 2010b: 321, f. 1, 4-5 (female only) = Hasarius tropicus Jastrzebski, 2010-correction of identification. Species: kweilinensis Prószynski, 1992, orientale Zabka, 1985, dactyloides Xie, Peng & Kim, 1993 listed variably in combinations with generic names Habrocestum, Habrocestoides, Hasarius and Chinattus seem to be misplaced in these genera and deserve transfer to own new genus. Qualification of 6 species of Hasarius as "nomina dubia" by the WSC (ver. 18.5) after Roewer's (1954[1955]: 1523-1524) is changed to "pending revision" because of existence of preserved type specimens. Synonymy of the genera Gedea Simon, 1902 and Meata Zabka, 1985 proposed recently by Maddison (2015) without published documentation are not recognized here until proof will appear printed.
19 new genera, one new subgenus and one new species of salticid spider are delimited and described. These are: Logunyllus gen. n., Marusyllus gen. n., Myrmage gen. n., Myrmagua gen. n., Myrmanu gen. n., Myrmapana gen. n., Myrmapeni gen. n., Myrmaplata gen. n., Myrmatheca gen. n., Myrmavola gen. n., Myrmele gen. n., Nandicius gen. n., Nepalicius gen. n., Okinawicius gen. n., Orienticius gen. n., Psenuc gen. n., Rudakius gen. n., Sittipub gen. n., Toxeus gen. reinstated. New subgenus Pellenes (Pellap) Prószyński, 2015 subgen. n. is also delimited. One misidentified species is renamed, Myrmavola yamasakii Prószyński, 2016 sp. n., and described as new species. I propose to consider name Helicius kimjoopili Kim, 1995 a "nomen dubium". Following synonyms are rejected and species reinstated to their current combinations: Tasa koreana Suguro & Yahata, 2014 (female) = Nepalicius koreanus (Wesolowska, 1981) comb. n., Tasa koreana Suguro & Yahata, 2014 (male) = Tasa nipponica Bohdanowicz & Prószyński, 1987, Pseudicius tokaraensis Suguro & Yahata, 2014 (male) = Nepalicius koreanus (Wesolowska, 1981) comb. n., Pseudicius tokaraensis Suguro & Yahata, 2014 (female 1) = Okinawicius okinawaensis (Prószyński, 1992) comb. n., Pseudicius tokaraensis Suguro & Yahata, 2014 (female 2) = Okinawicius tokaraensis (Bohdanowicz & Prószyński, 1987) comb. n. Also American Pseudicius siticulosus Peckham & Peckham, 1909 = Metaphidippus [?] siticulosus (Peckham, Peckham, 1909) comb. n., Myrmarachne paludosa (Simon, 1900) = Panachraesta paludosa Simon, 1900. I confirm hereby original genus placement of Emertonius exasperans Peckham & Peckham, 1892, as seconded by Prószyński & Deeleman-Reinhold, 2010: 164-167, 169-17, but disregarded by World Spider Catalog, ver. 2016.
The classification of jumping spiders (Salticidae) is revised to bring it into accord with recent phylogenetic work. Of the 610 recognized extant and fossil genera, 588 are placed at least to subfamily, most to tribe, based on both molecular and morphological information. The new subfamilies Onomastinae, Asemoneinae, and Eupoinae, and the new tribes Lapsiini, Tisanibini, Neonini, Mopsini, and Nannenini, are described. A new unranked clade, the Simonida, is recognized. Most other family-group taxa formerly ranked as subfamilies are given new status as tribes or subtribes. The large long-recognized clade recently called the Salticoida is ranked as a subfamily, the Salticinae, with the name Salticoida reassigned to its major subgroup (the sister group to the Amycoida). Heliophaninae Petrunkevitch and Pelleninae Petrunkevitch are considered junior synonyms of Chrysillini Simon and Harmochirina Simon respectively. Spartaeinae Wanless and Euophryini Simon are preserved despite older synonyms. The genus Meata Żabka is synonymized with Gedea Simon, and Diagondas Simon with Carrhotus Thorell. The proposed relationships indicate that a strongly ant-like body has evolved at least 12 times in salticids, and a strongly beetle-like body at least 8 times. Photographs of living specimens of all 7 subfamilies, 30 tribes, and 13 subtribes are presented.
A phylogenetic analysis of five sequenced genes (28S, 16S, EF1-α , CO1, ND1) from 81 genera of jumping spiders (Salticidae) and five outgroups supports the monophyly of the Dendryphantinae and Euophryinae and refines the concepts of the Plexippinae and Pelleninae. The clade that excludes lyssomanines and spartaeines and contains the bulk of salticid species is formally named as the Salticoida. The previously proposed clade delimited by an embolus articulated and separated from the tegulum by a developed distal hematodocha (as opposed to fused immovably to the tegulum) is rejected, suggesting the 'free embolus' evolved independently several times. Three major clades are discovered, the Marpissoida (including Dendryphantinae, Marpissinae and smaller groups such as synagelines), the Plexippoida (plexippines plus pellenines) and the Amycoida (including Amycinae, Sitticinae, Hyetusseae, Hurieae, Synemosyninae). The amycoids form a large neotropical radiation from which only a single known group (Sitticus and Attulus) has reached the Old World. The marpissoids also constitute a major New World group with relatively few species in the Old World. In contrast, the Plexippoida is predominantly an Old World group (except for the spectacular radiation of Habronattus in North America), as is the Heliophaninae. These results suggest that much of salticid diversification occurred after the separation of the continents of the Old World and New World.