The phylogenetic placement of the enigmatic pseudoscorpion family Menthidae (Pseudoscorpiones): a revised superfamily assignment based on new molecular data
Mark S. Harvey
A B * , Ligia R. Benavides C , Terrence L. Miller D , Julia G. Cosgrove C , Gonzalo Giribet C and Michael G. Rix D
A
B
C
D
Abstract
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.
Keywords: Arachnida, Iocheirata, morphology, Neobisioidea, phylogeny, phylotranscriptomic analysis, Sanger sequencing.
References
Aberer AJ, Kobert K, Stamatakis A (2014) ExaBayes: massively parallel Bayesian tree inference for the whole-genome era. Molecular Biology and Evolution 31, 2553-2556.
| Crossref | Google Scholar | PubMed |
Altenhoff AM, Levy J, Zarowiecki M, Tomiczek B, Warwick Vesztrocy A, Dalquen DA, Müller S, Telford MJ, Glover NM, Dylus D, Dessimoz C (2019) OMA standalone: orthology inference among public and custom genomes and transcriptomes. Genome Research 29, 1152-1163.
| Crossref | Google Scholar | PubMed |
Arabi J, Judson ML, Deharveng L, Lourenço WR, Cruaud C, Hassanin A (2012) Nucleotide composition of CO1 sequences in Chelicerata (Arthropoda): detecting new mitogenomic rearrangements. Journal of Molecular Evolution 74, 81-95.
| Crossref | Google Scholar | PubMed |
Ballesteros JA, Santibáñez-López CE, Baker CM, Benavides LR, Cunha TJ, Gainett G, Ontano AZ, Setton EVW, Arango CP, Gavish-Regev E, Harvey MS, Wheeler WC, Hormiga G, Giribet G, Sharma PP (2022) Comprehensive species sampling and sophisticated algorithmic approaches refute the monophyly of Arachnida. Molecular Biology and Evolution 39, msac021.
| Crossref | Google Scholar | PubMed |
Beier M (1932) Pseudoscorpionidea I. Subord. Chthoniinea et Neobisiinea. Tierreich 57, 1-258 [In German].
| Google Scholar |
Beier M (1948) Phoresie und Phagophilie bei Pseudoscorpionen. Österreichische Zoologische Zeitschrift 1, 441-497 Available at https://www.zobodat.at/pdf/OEZ_01_0441-0497.pdf [In German].
| Google Scholar |
Beier M (1952) On some Pseudoscorpionidea from Malaya and Borneo. Bulletin of the Raffles Museum 24, 96-108 Available at https://lkcnhm.nus.edu.sg/wp-content/uploads/sites/11/app/uploads/2017/06/24brm096-108.pdf.
| Google Scholar |
Beier M (1955) Pseudoscorpione im baltischen Bernstein aus dem Geologischen Staatsinstitut in Hamburg. Mitteilungen aus dem Mineralogisch-Geologischen Staatsinstitut in Hamburg 24, 48-54.
| Google Scholar |
Benavides LR (2025) Datasets and Supplementary material of ‘The phylogenetic placement of the enigmatic pseudoscorpion family Menthidae (Pseudoscorpiones): a revised superfamily assignment based on new molecular data’. Harvard Dataverse 2025, V2 [Dataset, published 17 October 2024].
| Crossref | Google Scholar |
Benavides LR, Cosgrove JG, Harvey MS, Giribet G (2019) Phylogenomic interrogation resolves the backbone of the Pseudoscorpiones Tree of Life. Molecular Phylogenetics and Evolution 139, 106509.
| Crossref | Google Scholar | PubMed |
Chamberlin JC (1930) A synoptic classification of the false scorpions or chela-spinners, with a report on a cosmopolitan collection of the same. Part II. The Diplosphyronida (Arachnida-Chelonethida). Annals and Magazine of Natural History 5(10), 1-48.
| Crossref | Google Scholar |
Chamberlin JC (1931) The arachnid order Chelonethida. Stanford University Publications, Biological Sciences 7(1), 1-284.
| Google Scholar |
Derkarabetian S, Baker CM, Hedin M, Prieto CE, Giribet G (2021) Phylogenomic re-evaluation of Triaenonychoidea (Opiliones: Laniatores), and systematics of Triaenonychidae, including new families, genera and species. Invertebrate Systematics 35, 133-157.
| Crossref | Google Scholar |
dos Reis M, Yang Z (2011) Approximate likelihood calculation on a phylogeny for Bayesian estimation of divergence times. Molecular Biology and Evolution 28(7), 2161-2172.
| Crossref | Google Scholar | PubMed |
Engel R (2012) Novel discovery of lamellar papillae on the grooming organ in Synsphyronus (Garypidae: Pseudoscorpiones). Arthropod Structure & Development 41, 265-269.
| Crossref | Google Scholar | PubMed |
Gao Z, Zhang F, Harvey MS (2025) New Asian pseudoscorpions improve the phylogenetic resolution of Garypinoidea (Pseudoscorpiones). Invertebrate Systematics 39, IS24098.
| Crossref | Google Scholar | PubMed |
Gardini G (2015) The species of the pseudoscorpion genus Pseudoblothrus (Pseudoscorpiones: Syarinidae) in Italy (on Italian pseudoscorpions XLVIII). Arachnologische Mitteilungen 49, 21-33.
| Crossref | Google Scholar |
Geißler C, Kotthoff U, Hammel J, Harvey MS, Harms D (2022) The first fossil of the pseudoscorpion family Ideoroncidae (Arachnida: Pseudoscorpiones): a new taxon from the mid-Cretaceous of northern Myanmar. Cretaceous Research 130, 105030.
| Crossref | Google Scholar |
Harms D, Dunlop JA (2017) The fossil history of pseudoscorpions (Arachnida: Pseudoscorpiones). Fossil Record 20, 215-238.
| Crossref | Google Scholar |
Harms D, Harvey MS, Roberts JD, Loria SF (2024) Tectonically driven climate change and the spread of temperate biomes: Insights from dragon pseudoscorpions (Pseudotyrannochthoniidae), a globally distributed arachnid lineage. Journal of Biogeography 51, 1032-1048.
| Crossref | Google Scholar |
Harvey MS (1992) The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Taxonomy 6, 1373-1435.
| Crossref | Google Scholar |
Harvey MS (1993) The systematics of the Hyidae (Pseudoscorpionida: Neobisioidea). Invertebrate Taxonomy 7, 1-32.
| Crossref | Google Scholar |
Harvey MS (1998) Pseudoscorpion groups with bipolar distributions: a new genus from Tasmania related to the Holarctic Syarinus (Arachnida, Pseudoscorpiones, Syarinidae). Journal of Arachnology 26, 429-441.
| Google Scholar |
Harvey MS (2006) New species and records of the pseudoscorpion family Menthidae (Pseudoscorpiones). Records of the Western Australian Museum 23, 167-174.
| Crossref | Google Scholar |
Harvey MS (2016) The systematics of the pseudoscorpion family Ideoroncidae (Pseudoscorpiones, Neobisioidea) in the Asian region. Journal of Arachnology 44, 272-329.
| Crossref | Google Scholar |
Harvey MS (2023) A preliminary phylogeny for the pseudoscorpion family Garypinidae (Pseudoscorpiones: Garypinoidea), with new taxa and remarks on the Australasian fauna. Invertebrate Systematics 37, 623-676.
| Crossref | Google Scholar |
Harvey MS, Du Preez G (2014) A new troglobitic ideoroncid pseudoscorpion (Pseudoscorpiones: Ideoroncidae) from southern Africa. Journal of Arachnology 42, 106-110.
| Crossref | Google Scholar |
Harvey MS, Muchmore WB (1990) The systematics of the family Menthidae (Pseudoscorpionida). Invertebrate Taxonomy 3, 941-964.
| Crossref | Google Scholar |
Harvey MS, Muchmore WB (2013) The systematics of the pseudoscorpion family Ideoroncidae (Pseudoscorpiones: Neobisioidea) in the New World. Journal of Arachnology 41, 229-290.
| Crossref | Google Scholar |
Harvey MS, Šťáhlavský F (2010) A review of the pseudoscorpion genus Oreolpium (Pseudoscorpiones: Garypinidae), with remarks on the composition of the Garypinidae and on pseudoscorpions with bipolar distributions. Journal of Arachnology 38, 294-308.
| Crossref | Google Scholar |
Harvey MS, Lopes PC, Goldsmith GR, Halajian A, Hillyer MJ, Huey JA (2015) A novel symbiotic relationship between sociable weaver birds (Philetairus socius) and a new cheliferid pseudoscorpion (Pseudoscorpiones: Cheliferidae) in southern Africa. Invertebrate Systematics 29, 444-456.
| Crossref | Google Scholar |
Harvey MS, Abrams KM, Beavis AS, Hillyer MJ, Huey JA (2016a) Pseudoscorpions of the family Feaellidae (Pseudoscorpiones: Feaelloidea) from the Pilbara region of Western Australia show extreme short-range endemism. Invertebrate Systematics 30, 491-508.
| Crossref | Google Scholar |
Harvey MS, Huey JA, Hillyer MJ, McIntyre E, Giribet G (2016b) The first troglobitic species of Gymnobisiidae (Pseudoscorpiones, Neobisioidea), from Table Mountain (Western Cape Province, South Africa) and its phylogenetic position. Invertebrate Systematics 30, 75-85.
| Crossref | Google Scholar |
Harvey MS, Cosgrove JG, Harms D, Selden PA, Shih C, Wang C-C (2018) The oldest chthonioid pseudoscorpion (Arachnida: Pseudoscorpiones: Chthonioidea: Chthoniidae): a new genus and species from mid-Cretaceous Burmese amber. Zoologischer Anzeiger 273, 102-111.
| Crossref | Google Scholar |
Harvey MS, Hillyer MJ, Carvajal JI, Huey JA (2020) Supralittoral pseudoscorpions of the genus Garypus (Pseudoscorpiones: Garypidae) from the Indo-West Pacific region, with a review of the subfamily classification of Garypidae. Invertebrate Systematics 34, 34-87.
| Crossref | Google Scholar |
Harvey MS, Burger MAA, Abrams KM, Finston TL, Huey JA, Perina G (2023) The systematics of the pseudoscorpion genus Indohya (Pseudoscorpiones: Hyidae) in Australia. Zootaxa 5342, 1-119.
| Crossref | Google Scholar | PubMed |
Henderickx H (2005) A new Geogarypus from Baltic Amber (Pseudoscorpiones: Geogarypidae). Phegea 33, 87-92.
| Google Scholar |
Hoang DT, Chernomor O, von Haeseler A, Minh BQ, Vinh LS (2018) UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35(2), 518-522.
| Crossref | Google Scholar | PubMed |
Judson MLI (1997) Catalogue of the pseudoscorpion types (Arachnida: Chelonethi) in the Natural History Museum, London. Occasional Papers on Systematic Entomology 11, 1-54.
| Google Scholar |
Judson MLI (2000) Electrobisium acutum Cockerell, a cheiridiid pseudoscorpion from Burmese amber, with remarks on the validity of the Cheiridioidea (Arachnida, Chelonethi). Bulletin of the Natural History Museum, Geology 56, 79-83.
| Google Scholar |
Judson MLI (2003) Baltic amber pseudoscorpions (Arachnida, Chelonethi): a new species of Neobisium (Neobisiidae) and the status of Obisium rathkii Koch & Berendt. Geodiversitas 25, 445-450.
| Google Scholar |
Judson MLI (2009) Cheliferoid pseudoscorpions (Arachnida, Chelonethi) from the Lower Cretaceous of France. Geodiversitas 31, 61-71.
| Crossref | Google Scholar |
Judson MLI (2012) Reinterpretation of Dracochela deprehendor (Arachnida: Pseudoscorpiones) as a stem-group pseudoscorpion. Palaeontology 55, 261-283.
| Crossref | Google Scholar |
Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30, 772-780.
| Crossref | Google Scholar | PubMed |
Katoh K, Misawa K, Kuma K-i, Miyata T (2002) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research 30, 3059-3066.
| Crossref | Google Scholar | PubMed |
Kearse M, Moir R, Wilson A, Stones-Havas S, Cheung M, Sturrock S, Buxton S, Cooper A, Markowitz S, Duran C, Thierer T, Ashton B, Meintjes P, Drummond A (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28(12), 1647-1649.
| Crossref | Google Scholar |
Kolesnikov VB, Turbanov IS, Eskov KY, Propistsova EA, Bashkuev AS (2022) First non-amber Mesozoic pseudoscorpion from Upper Triassic deposits of eastern Europe, with a description of two new fossil subfamilies (Arachnida, Pseudoscorpiones, Feaellidae). Papers in Palaeontology 8(5), e1466.
| Crossref | Google Scholar |
Lartillot N, Philippe H (2004) A Bayesian mixture model for across-site heterogeneities in the amino-acid replacement process. Molecular Biology and Evolution 21(6), 1095-1109.
| Crossref | Google Scholar | PubMed |
Le SQ, Gascuel O (2008) An improved general amino acid replacement matrix. Molecular Biology and Evolution 25, 1307-1320.
| Crossref | Google Scholar | PubMed |
Mahnert V (1984) Beitrag zu einer besseren Kenntnis der Ideoroncidae (Arachnida: Pseudoscorpiones), mit Beschreibung von sechs neuen Arten. Revue Suisse de Zoologie 91, 651-686 [In German].
| Crossref | Google Scholar |
Mahnert V (2007) Pseudoscorpions (Arachnida: Pseudoscorpiones) of the Socotra Archipelago, Yemen. Fauna of Arabia 23, 271-307.
| Google Scholar |
Manni M, Berkeley MR, Seppey M, Zdobnov EM (2021a) BUSCO: assessing genomic data quality and beyond. Current Protocols 1, e323.
| Crossref | Google Scholar |
Manni M, Berkeley MR, Seppey M, Simão FA, Zdobnov EM (2021b) BUSCO update: novel and streamlined workflows along with broader and deeper phylogenetic coverage for scoring of eukaryotic, prokaryotic, and viral genomes. Molecular Biology and Evolution 38(10), 4647-4654.
| Crossref | Google Scholar |
Minh BQ, Schmidt HA, Chernomor O, Schrempf D, Woodhams MD, von Haeseler A, Lanfear R (2020) IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution 37, 1530-1534.
| Crossref | Google Scholar | PubMed |
Muchmore WB (1971) Phoresy by North and Central American pseudoscorpions. Proceedings of the Rochester Academy of Science 12, 79-97.
| Google Scholar |
Muchmore WB (1973) New and little known pseudoscorpions, mainly from caves in Mexico (Arachnida, Pseudoscorpionida). Bulletin of the Association for Mexican Cave Studies 5, 47-62.
| Google Scholar |
Muchmore WB (1984) Troglobochica, a new genus from caves in Jamaica, and redescription of the genus Bochica Chamberlin (Pseudoscorpionida, Bochicidae). Journal of Arachnology 12, 61-68.
| Google Scholar |
Muchmore WB (1996) A remarkable new genus and species of Pseudoscorpionida (Syarinidae) from a cave in Arizona. Southwestern Naturalist 41, 145-148.
| Google Scholar |
Muchmore WB (1998) Review of the family Bochicidae, with new species and records (Arachnida: Pseudoscorpionida). Insecta Mundi 12, 117-132.
| Google Scholar |
Murienne J, Harvey MS, Giribet G (2008) First molecular phylogeny of the major clades of Pseudoscorpiones (Arthropoda: Chelicerata). Molecular Phylogenetics and Evolution 49, 170-184.
| Crossref | Google Scholar | PubMed |
Nassirkhani M, Shoushtari RV (2015) The first record of the family Menthidae Chamberlin (Arachnida: Pseudoscorpiones) from Iran. The Journal of Zoology Studies 1, 27-31.
| Google Scholar |
Ontano AZ, Gainett G, Aharon S, Ballesteros JA, Benavides LR, Corbett KF, Gavish-Regev E, Harvey MS, Monsma S, Santibáñez-López CE, Setton EVW, Zehms JT, Zeh JA, Zeh DW, Sharma PP (2021) Taxonomic sampling and rare genomic changes overcome long-branch attraction in the phylogenetic placement of pseudoscorpions. Molecular Biology and Evolution 38, 2446-2467.
| Crossref | Google Scholar | PubMed |
Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67, 901-904.
| Crossref | Google Scholar | PubMed |
Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3), 539-542.
| Crossref | Google Scholar | PubMed |
Röschmann LM, Harvey MS, Hou Y, Harms D, Kotthoff U, Hammel JU, Ren D, Loria SF (2024) First fossil species of family Hyidae (Arachnida: Pseudoscorpiones) confirms 99 million years of ecological stasis in a Gondwanan lineage. PeerJ 12, 17515.
| Crossref | Google Scholar | PubMed |
Schawaller W, Shear WA, Bonamo PM (1991) The first Paleozoic pseudoscorpions (Arachnida, Pseudoscorpionida). American Museum Novitates 3009, 1-24.
| Google Scholar |
Trifinopoulos J, Nguyen L-T, von Haeseler A, Minh BQ (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research 44(W1), W232-W235.
| Crossref | Google Scholar | PubMed |
World Pseudoscorpiones Catalog (2025) World Pseudoscorpiones Catalog. (Natural History Museum) Available at https://wac.nmbe.ch/order/pseudoscorpiones/3 [Verified 22 January 2025]
Yang Z (2007) PAML 4: phylogenetic analysis by maximum likelihood. Molecular Biology and Evolution 24, 1586-1591.
| Crossref | Google Scholar | PubMed |
Yang Z, Rannala B (2006) Bayesian estimation of species divergence times under a molecular clock using multiple fossil calibrations with soft bounds. Molecular Biology and Evolution 23, 212-226.
| Crossref | Google Scholar | PubMed |
Zaragoza JA (2010) Arcanobisium, a remarkable new genus, representing a new subfamily with a relictual distribution from eastern Spain (Arachnida: Pseudoscorpiones: Syarinidae). Zootaxa 2491, 41-60.
| Crossref | Google Scholar |
