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Invertebrate Systematics

Invertebrate Systematics

Invertebrate Systematics publishes significant contributions and reviews on the systematics, phylogeny and biogeography of all invertebrate taxa. Read more about the journalMore

Editor-in-Chief: Gonzalo Giribet

Publishing Model: Hybrid. Open Access options available.

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Latest

These articles are the latest published in the journal. Invertebrate Systematics is published under a continuous publication model. More information is available on our Continuous Publication page.

Published online 15 September 2025

IS24087The phylogenetic placement of the enigmatic pseudoscorpion family Menthidae (Pseudoscorpiones): a revised superfamily assignment based on new molecular data

Mark S. Harvey 0000-0003-1482-0109, Ligia R. Benavides 0000-0003-4387-1220, Terrence L. Miller, Julia G. Cosgrove 0000-0003-2776-699X, Gonzalo Giribet 0000-0002-5467-8429 and Michael G. Rix 0000-0001-5086-3638
 

A picture of Specimen of Thenmus aigialites (left) and phylogenetic analysis (right) that reveals that the newly collected menthid pseudoscorpion belongs to the superfamily Neobisioidea.

Pseudoscorpions are an ancient arachnid group with a fossil record that extends to the Devonian, with all modern families having likely evolved during the Mesozoic. One of the rarest pseudoscorpion families, Menthidae, is sporadically distributed around the world, and ever since its description has been included in the superfamily Garypoidea. Based on new Sanger sequencing and phylotranscriptomic data, Menthidae are inferred to be a member of the superfamily Neobisioidea, and the sister-group to a clade that includes Gymnobisiidae, Neobisiidae and some Syarinidae. (Image credit: Mark Harvey.)

IS24087 Abstract  |  IS24087 Full Text  | IS24087PDF (2.9 MB)  | IS24087Supplementary Material (357 KB)   Open Access Article

Published online 04 September 2025

IS25016Molecular phylogenetic analysis of the family Chlorogomphidae (Odonata, Anisoptera)

Thomas Schneider, Andy Vierstraete 0000-0001-5005-8882, Oleg E. Kosterin 0000-0001-5955-4057, Dietmar Ikemeyer 0000-0003-0751-3266, Fang-Shuo Hu 0000-0003-1269-8904, Tom Kompier 0000-0001-9488-7746 and Henri J. Dumont
 

Two photographs of Chlorogomphus dragonflies, C. aritai on the left and and C. suzukii on the right.

Phylogenetic analysis of Chlorogomphidae, using both nuclear and mitochondrial markers, revealed that the family is best treated as containing a single genus, Chlorogomphus. Chlorogomphus montanus is synonymised to Chlorogomphus nasutus, and Chlorogomphus urolobatus is synonymised to Chlorogomphus infuscatus. The previous synonymy of Chlorogomphus suzukii and Chlorogomphus tunti is confirmed. Other potential synonyms are discussed, including C. nasutus, which actually refers to two (not three) species – herein considered as C. nasutus and C. satoi – leaving the genus with 56 species. (Image credit: left, Tom Kompier; right, Fang-Shuo Hu.)

IS25016 Abstract  | PDF (3 MB) - $60.00   | IS25016Supplementary Material (393 KB)
Published online 12 August 2025

IS25033Phylogenomic placement of a new genus and species of jumping spiders (Araneae: Salticidae: Spartaeini) based on UCE data

Yi Ni 0009-0003-3239-3697, Kun Yu 0000-0002-5517-4484 and Junxia Zhang 0000-0003-2179-3954
 

Pictures of the new genus and species of salticid showing the female body, male palp and female epigyne (left to right).

Jumping spiders (Salticidae) are a diverse group, and understanding their evolutionary relationships is crucial for biodiversity studies. This research addresses the classification of an enigmatic lineage within the subtribe Spartaeina, previously unresolved due to limited specimens. With the updated phylogeny using ultraconserved element data, we provided a solid phylogenetic framework for Spartaeina and established a new genus, Protaeus. These findings enhance our knowledge of phylogeny and diversity of Spartaeina, and highlight the need for further exploration in this fascinating group. (Image credit: Yi Ni.)

IS25033 Abstract  | PDF (7.5 MB) - $60.00   | IS25033Supplementary Material (9.4 MB)
Published online 12 August 2025

IS25023Molecular phylogeny provides new insights into the classification of the order Spirostreptida (Arthropoda: Diplopoda), with description of Pericambala foveiformis, sp. nov. (Cambalidea: Pericambalidae) using integrated evidence

Hong-Ru Xu, Kuang Zhang, Hui Wang, Wei-Xin Liu 0000-0001-9122-8868 and Liang Li 0000-0003-2338-1099
 

Gene maps of two spirostreptid millipedes. The new species (right) has the largest diplopod mitogenome currently known.

A new millipede species Pericambala foveiformis, sp. nov. is described using integrated methods. The mitogenome of P. foveiformis with 17,467 bp in length, is the largest millipede mitogenome currently known. Molecular phylogenies provided new insights into the phylogenetic relationships of higher taxa within the order Spirostreptida. (Image credit: Liang Li.)

IS25023 Abstract  | PDF (4.4 MB) - $60.00   | IS25023Supplementary Material (1.3 MB)
Published online 06 August 2025

IS24049The tables have turned: taxonomy, systematics and biogeography of the Acropora hyacinthus (Scleractinia: Acroporidae) complex

Sage H. Rassmussen 0000-0001-5077-6905, Peter F. Cowman, Andrew H. Baird, Augustine J. Crosbie, Andrea M. Quattrini, Victor Bonito, Frederic Sinniger, Saki Harii, Patrick C. Cabaitan, Nur Fadli, Chun-Hong Tan, Julia Yun-Hsuan Hung 0009-0000-6809-1859, Teina Rongo, Danwei Huang, Tuikolongahau Halafihi and Tom C. L. Bridge
 

Photograph of Acropora tersa with tabular colony morphology, which dominate many shallow-water coral reef communities throughout the Indo-Pacific

Genomic approaches suggest that traditional taxonomy does not accurately reflect the true diversity or systematic relationships of corals. We combine molecular analysis and morphology of type material to revise the Acropora hyacinthus group. Our results show significantly higher species richness than currently accepted, with five new species and nine reinstated species, demonstrating narrower species geographic ranges than previously assumed. Given the key ecological role that Acropora play on Indo-Pacific reefs, our findings are relevant for reef conservation and management. (Photograph by Tom Bridge.)

IS24049 Abstract  |  IS24049 Full Text  | IS24049PDF (9.1 MB)  | IS24049Supplementary Material (13.5 MB)   Open Access Article

Published online 23 July 2025

IS25029Phylogeny and evolution of male genitalia in Withiidae (Arachnida: Pseudoscorpiones)

Catalina Romero-Ortiz 0000-0001-8939-7814, Mark S. Harvey 0000-0003-1482-0109, Ligia R. Benavides 0000-0003-4387-1220, Sebastian Cuadrado-Rios 0000-0002-1060-820X and Carlos E. Sarmiento 0000-0003-4012-8108
 

Examples of long and short type ejaculatory canals of male Withiidae and their evolutionary rate of change from long to short.

Pseudoscorpions show great promise for studying the evolution of genitalia because of their many unique sexually related features. We estimated a dated phylogeny based on morphology and genes for Withiidae to understand how quickly certain body parts changed. Sensory organs were found to change faster than genitalia, suggesting that environmental adaptation might exert stronger selective pressures than sexual selection in shaping their morphology. Our study also highlights the importance of considering both morphological features and genes when examining evolution. (Image credit: Catalina Romero-Ortiz.)

IS25029 Abstract  | PDF (1.5 MB) - $60.00   | IS25029Supplementary Material (587 KB)
Published online 18 July 2025

IS25009Taxonomy and systematics of the Australasian gum nut orb-weaving spider genus Carepalxis (Araneae, Araneidae)

Pedro de S. Castanheira 0000-0002-0623-1622, Dimitar Dimitrov 0000-0001-5830-5702, Renner L. C. Baptista, Nikolaj Scharff and Volker W. Framenau
 

Frontal view of a female Carepalxis montifera showing the paired cephalic humps, which are characteristic of the genus.

The molecular phylogeny of the gum nut orb-weaving spider genus Carepalxis was investigated, revealing a polyphyletic Carepalxis: its Australasian representatives were part of the informal clade ‘backobourkiines’, and the Neotropical species were related to the genus Ocrepeira. A revised Carepalxis, recognised by the presence of two cephalic humps in females and two enlarged tibial spines in males, is composed of seven endemic Australian species, including four new species described here, and C. beelzebub, which also occurs in New Guinea. (Photography by Pedro Castanheira.)

IS25009 Abstract  |  IS25009 Full Text  | IS25009PDF (20.2 MB)  | IS25009Supplementary Material (2.6 MB)   Open Access Article

Published online 11 July 2025

IS25025Cryptic species, mitochondrial phylogenomics and historical biogeography in the endemic genus Schistodesmus (Bivalvia, Unionidae) from China

Kaiyu Hou, Xianan Wang, Junli Jia, Xiongjun Liu, Xiaoping Wu, Dandong Jin, Jianmei An and Ruiwen Wu 0000-0002-8936-6054
 

Picture of shells of five Schistodesmus species.

An integrative taxonomic approach, including morphological and molecular data, revealed three new cryptic species: Schistodesmus luqiaoensis, Schistodesmus tongpenensis, and Schistodesmus xinyuensis. Maternal mitochondrial genomes of the first two species and Schistodesmus lampreyanus were sequenced, and interspecific relationships elucidated based on mitochondrial phylogenomics. Molecular clock analysis and biogeographical reconstruction based on fossil-calibrated dating indicated that the ancestor of Schistodesmus originated in the paleo-Yangtze River Basin during the Eocene (c. 50.74 Ma) and began diversifying c. 18.68 Ma in the Neogene. (Photography by Kaiyu Hou.)

IS25025 Abstract  | PDF (1.9 MB) - $60.00   | IS25025Supplementary Material (1.4 MB)
Published online 03 July 2025

IS24101The phylogeny and systematics of the Costellariidae (Caenogastropoda: Turbinelloidea) revisited

Alexander Fedosov 0000-0002-8035-1403, Philippe Bouchet 0000-0001-5864-8676, Aart Dekkers, Sandro Gori, Shih-I Huang, Yuri Kantor 0000-0002-3209-4940, Thomas Lemarcis, Maxwell Marrow, Claudia Ratti, Gary Rosenberg, Richard Salisbury, Sofia Zvonareva and Nicolas Puillandre
 

Phylogenomic tree of Costellaridae showing the relationships of the nine new genera and their shell forms.

The family Costellariidae is a charismatic, species-rich, yet poorly studied group of marine gastropod molluscs. Recent analyses of DNA sequences and shell features have failed to embrace the complexity of the group. Here, we analyse the morphology, DNA barcodes and genomic data from over 200 species, with emphasis on the South Pacific and Caribbean faunae, describing 9 new genera and 23 new species. Our results update the phylogeny and systematics of Costellariidae, corroborating patterns of species diversity in tropical seas. (Image credit: Alexander Fedosov.)

IS24101 Abstract  | PDF (14.7 MB) - $60.00   | IS24101Supplementary Material (11.9 MB)
Published online 01 July 2025

IS24084Does morphological diversity in amphidromous Macrobrachium (Decapoda: Caridea: Palaemonidae) reflect species homology or habitat partitioning?

Valentin de Mazancourt 0000-0003-3739-603X, Pierre Feutry, Coline Bernard 0009-0002-2902-2871, Gérard Marquet, Philippe Keith 0000-0002-1600-4909 and Magalie Castelin 0000-0002-9984-2122
 

A gradient map showing how clades of Macrobrachium have developed morphological adaptations depending on the current at various elevations.

We apply rigorous taxonomic and phylogenetic frameworks to shed light on both the taxonomy and evolution of amphidromous freshwater shrimps. Automatic species delineation methods based on molecular data revealed the existence of several species complexes within Macrobrachium. Results suggest habitat-driven speciation modes, where species differentiate in similar habitats distributed across the Indo-Pacific. The morphology of amphidromous Macrobrachium reflects numerous conserved adaptations to flow velocity. (Image credit: Valentin de Mazancourt.)

IS24084 Abstract  | PDF (2.1 MB) - $60.00   | IS24084Supplementary Material (1.1 MB)
Published online 01 July 2025

IS24103Paedomorphic adaptations in a new Heterostigma species: a novel strategy for ascidians to live in soft-bottom habitats

Riccardo Virgili 0000-0003-0536-7939, Valentina Tanduo, Salvatore D’Aniello, Angelo Fontana, Xavier Turon and Fabio Crocetta
 

Three images that show sessile to motile phenotype shift of Heterostigma monniotae, its complete mitochondrial genome, and the phylogenetic placement within Pyuridae ascidians.

Heterostigma is a poorly known genus of pyurid ascidian, deemed to inhabit the interstitial space between coarse sand grains. A new species, Heterostigma monniotae, is described, updating the taxonomy of the genus and clarifying its phylogenetic position through analyses of single loci and complete mitochondrial genome. In addition, observations on live specimens revealed an ontogenetic shift in mature specimens from a sessile to a motile phenotype. These new findings shed light on the evolutionary implications of this adaptive strategy. (Image credit: R. Virgili.)

IS24103 Abstract  | PDF (4.4 MB) - $60.00   | IS24103Supplementary Material (16.6 MB)
Published online 25 June 2025

IS25007Molecular and morphological data uncover a striking new genus of dragon millipedes in Thailand, with alternately long and short legs (Diplopoda, Polydesmida, Paradoxosomatidae)

Ruttapon Srisonchai 0000-0002-7142-0999, Henrik Enghoff, Natdanai Likhitrakarn, Somsak Panha and Chirasak Sutcharit
 

Picture showing a new dragon millipede genus from Thailand with alternately long and short legs (heteropody).

Dragon millipedes recently collected in southern Thailand exhibited a unique morphological trait: alternating long and short legs on each diplosegment (heteropody), a feature previously unseen in the family Paradoxosomatidae. Phylogenetic analyses using morphology and genetic data strongly support the monophyly of all dragon millipede genera. The new specimens form a distinctly separate clade for which we establish a new monotypic genus, Alternaxytes, characterised by unique gonopodal features and heteropodous legs, which are an example of convergent evolution within millipedes. (Photographs and image credit: Ruttapon Srisonchai.)

IS25007 Abstract  | PDF (7.3 MB) - $60.00   | IS25007Supplementary Material (123 KB)
Published online 27 May 2025

IS25003Addressing taxonomic shortfalls in Neotropical gastrotrichs: a new genus and species of freshwater Gastrotricha (Chaetonotida: Paucitubulatina) from Brazil

Axell K. Minowa 0000-0002-2962-6053, Alexander Kieneke, Maria Balsamo, Loretta Guidi and André R. S. Garraffoni
 

A distribution map of a new Brazilian gastrotrich, an image showing molecular phylogenetic relationships, and a SEM image showing upper body details.

The knowledge about Gastrotricha in Brazilian territory is still far from complete. In our work we describe a new species and new genus distributed across a wide latitudinal range. The genus is characterised by a novel cephalic cuticular ornamentation, called leafed scale, and supported by molecular phylogeny. In our work we reinforce the importance of using methods of integrative taxonomy for uncovering gastrotrich diversity. (Image credit: Axell Minowa.)

IS25003 Abstract  | PDF (12.5 MB) - $60.00   | IS25003Supplementary Material (4.1 MB)
Published online 13 May 2025

IS24079A new large violin spider from Argentina, with a discussion on the homology and evolution of female genital structures in Loxosceles (Araneae: Sicariidae)

Ivan L. F. Magalhaes 0000-0003-3728-3270
 

Photograph of the female of a new species of violin spider (Loxosceles) from Parque Nacional Talampaya, north-west Argentina.

A new species of violin spider, Loxosceles galianoa, sp. nov. is described from north-western Argentina as a member of the laeta group. A survey of the diversity of the female genitalia in Loxosceles, combined with a phylogeny of its species groups, suggests that the female genitalia is more diverse than that of males in this genus. It is hypothesised that the medically significant Loxosceles laeta likely originated in Peru, not Chile. (Photograph by I. L. F. Magalhaes.)

IS24079 Abstract  | PDF (16 MB) - $60.00   | IS24079Supplementary Material (673 KB)

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Collections

Collections are a curation of articles relevant to a topical research area

Invertebrate Systematics of the Americas and Beyond: Spotlight for ICIM-6

To mark the sixth International Congress on Invertebrate Morphology (ICIM-6) in Chile, Invertebrate Systematics is offering a free-to-read Virtual Issue. This special collection highlights recent research from the the Pacific region and beyond, showcasing the remarkable work of local researchers. Dive into the latest discoveries from deep-sea squat lobsters to Antarctic echinoderms, free to read until November 2025.

Last Updated: 01 Aug 2025

Myriapods and Onychophorans—their Systematics, Biogeography and Species Delimitation

Myriapoda and Onychophora constitute two clades of mostly soil-dwelling multi-legged panarthropods, once thought to form the clade Uniramia, but now genomic evidence strongly suggest that Onychophora is the sister group of Arthropoda, where myriapods are nested. The articles compiled in this Collection include several examples of the integrative work we publish in our journal, and we welcome further submissions of your best work on myriapods and onychophorans.

Last Updated: 19 Jun 2017

30 Years of Invertebrate Systematics

Invertebrate Systematics has focused on the publication of primary research and review articles on all aspects related to systematics of invertebrates, including taxonomic papers with a focus on evolution and ecology. To celebrate 30 years of publication of Invertebrate Systematics, this Collection contains some of our most highly cited papers.

Last Updated: 14 Dec 2016

Malacology

In this Collection, we present articles covering a diversity of taxa (Bivalvia, Cephalopoda, Gastropoda, Polyplacophora) and approaches (molecular, morphological – including morphometrics – and total evidence) to better understand molluscan diversification at different time-scales. These articles encompass research and species found globally, highlighting both the interdisciplinary and collaborative nature of the science published in Invertebrate Systematics.

Last Updated: 30 Jun 2016

Decapoda

Invertebrate Systematics has published many significant studies of the Decapoda, of which 12 are showcased in this Collection. Launched to coincide with the 8th International Crustacean Congress in Frankfurt, Germany (18-23 August 2014), this Collection spans molecular and morphological phylogenetic and revisionary taxonomic studies of crabs, crayfish, freshwater and marine shrimps.

Last Updated: 08 Aug 2014

Hymenoptera

Invertebrate Systematics has published many important studies on hymenopterans, 14 of which are showcased in this Collection. Launched to coincide with the 8th International Congress of Hymenopterists in Cusco, Peru (20-25 July 2014), this significant collection comprises molecular and morphological phylogenetic and taxonomic studies on a wide range of families.

Last Updated: 01 Jul 2014

Arachnology

This Collection, launched to coincide with the 19th International Congress of Arachnology in Taiwan in June 2013, comprises 12 important studies on arachnids from Invertebrate Systematics. Together, the publications in this Collection reveal key aspects of the systematics, biology and biogeography of these invertebrate animals.

Last Updated: 01 Jun 2013

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